Red Oak Species Research Articles

The effects of environmental heterogeneity on the morphological similarity of toothed-leafed red oak species (Acutifoliae; <i>Quercus</i>) in Mexico and Central America

Background: In plants, environmental gradients can promote similar morphophysiological traits in species that inhabit areas with similar climatic conditions. Here, to evaluate possible causes of morphological similarity, we analyzed leaf morphological and functional variation in toothed-leaf oaks and its relationships with environmental and geographic variables. Hypotheses: We expected that species with a greater degree of environmental overlap would exhibit similar leaf morphology, and those with a broader range of environmental tolerance would exhibit greater leaf morphological variation. Studied species: Acutifoliae group (Fagaceae, Quercus) Study sites and dates: Leaf samples were collected from adult trees distributed in diverse forest types between 495 and 2,566 meters above sea level (m asl) in Mexico and Guatemala. Methods: We compared patterns of leaf variation in a total of 2,934 leaves from 640 individuals corresponding to 68 populations of ten oak species using univariate and multivariate analyses. We also characterized the environmental conditions where each of the species occurred using the available geographical records and bioclimatic variables. Results: We found morphological overlap in leaf traits in oak species occurring in similar climatic environments. Additionally, oak species with a greater range of environmental distributions showed greater leaf shape morphological variation. Conclusions: The environment influenced the generation of similar leaf morphologies among oak species despite geographical distance and phylogenetic relationships.

Growth and productivity of red oak (Q.Rubra L.) forest cultures in the North-West Caucasus

The paper presents the results of the study of red oak (Q. rubra L.) cultures located on the territory of «Kurjip district forestry» of the Republic of Adygea (western part of the North Caucasus). The characteristic of red oak is given as a tree species, less demanding to soil conditions in comparison with other forest-forming species and possessing high decorative properties, due to which it is used for solitary plantings, as hedges along highways, arrays in parks, squares and other objects of landscape architecture. It combines well both in pure plantings and in combination with petiole oak, hornbeam and aspen. Red oak can grow on sandy loam and loamy podzolic soils without stagnant moisture, mainly on the banks of rivers, it grows worse on calcareous soil. The tree species due to well-developed root system is resistant to wind, also characterized by good resistance to damage by entomo- and phyto-pests. The growth and productivity of red oak and related species were analyzed in the work, and the graphs of growth progress by height and diameter of crops were plotted. It was found that in more favorable growing conditions red oak is not inferior to petiole oak in growth and productivity. Keywords: North-West Caucasus, Forest Cultures, Red Oak, Common Oak, Non-Native Species, Growth, Productivity.

A haplotype-resolved chromosome-scale genome for Quercus rubra L. provides insights into the genetics of adaptive traits for red oak species.

Northern red oak (Quercus rubra L.) is an ecologically and economically important forest tree native to North America. We present a chromosome-scale genome of Q. rubra generated by the combination of PacBio sequences and chromatin conformation capture (Hi-C) scaffolding. This is the first reference genome from the red oak clade (section Lobatae). The Q. rubra assembly spans 739 Mb with 95.27% of the genome in 12 chromosomes and 33,333 protein-coding genes. Comparisons to the genomes of Q. lobata and Q. mongolica revealed high collinearity, with intrachromosomal structural variants present. Orthologous gene family analysis with other tree species revealed that gene families associated with defense response were expanding and contracting simultaneously across the Q. rubra genome. Quercus rubra had the most CC-NBS-LRR and TIR-NBS-LRR resistance genes out of the nine species analyzed. Terpene synthase gene family comparisons further reveal tandem gene duplications in TPS-b subfamily, similar to Q. robur. Phylogenetic analysis also identified four subfamilies of the IGT/LAZY gene family in Q. rubra important for plant structure. Single major QTL regions were identified for vegetative bud break and marcescence which contain candidate genes for further research, including a putative ortholog of the circadian clock constituent cryptochrome (CRY2) and eight tandemly duplicated genes for serine protease inhibitors, respectively. Genome-environment associations across natural populations identified candidate abiotic stress tolerance genes and predicted performance in a common garden. This high-quality red oak genome represents an essential resource to the oak genomics community which will expedite comparative genomics and biological studies in Quercus species.

Forest composition and red oak (Quercus sp.) response to elevation gradients across greentree reservoirs

Elevation gradients within forested wetlands have long been recognized for their role in defining species composition through factors such as hydrology and soil characteristics. Greentree reservoirs (GTRs) are levee-impounded tracts of bottomland hardwood forest flooded throughout the winter months to provide habitat for overwintering waterfowl. Artificial flooding of GTRs alters the forest composition due to flood frequency, depth, and duration in combination with slight changes in topography. To evaluate the effect of elevation gradients, soil properties, and management techniques in the overstory species composition and red oak (Quercus spp.) species abundance, we inventoried 662 plots across 12 independent GTRs in eastern Arkansas. In the lower elevations ranging from 50.98 to 54.99 ​m above sea level, the importance value index (IVI) was highest for nuttall oak (Quercus texana) and overcup oak (Quercus lyrata), whereas IVI shifted to cherrybark oak (Quercus pagoda) in the higher elevations ranging from 54.99 to 58.00 ​m. Alpha diversity did not differ by elevation gradient, soil property, or management technique within GTRs. Beta diversity, using non-metric multi-dimensional scaling (NMDS) analysis, indicated site-specific variability significantly correlated with the environmental predictors, including elevation (R2 = ​0.57), easting (R2 = ​0.47), soil texture (R2 = ​0.21), and pH (R2 = ​0.12). Red oak species-specific mixed-effects modeling of abundance response using Poisson distribution suggested an inverse correlation of nuttall oak and a direct correlation of cherrybark oak abundance with elevation. However, willow oak (Quercus phellos) abundance was not significantly affected by elevation but was by silt loam soil texture and restoration management techniques. These findings will aid management efforts to reduce the dominance of less desirable species that are prominent under specific environmental conditions and promote the dominance of more desirable species. Ultimately GTR sustainability is increasingly important amid the unpredictable impacts of climate change on the preferred red oak species that are economically, ecologically, and environmentally valuable to the sustaining economy of the local community and managing habitats for wildlife.

Growth Response of Red Oaks to Climatic Conditions in the Lower Mississippi Alluvial Valley: Implications for Bottomland Hardwood Restoration with a Changing Climate

Bottomland hardwood forests (BHFs) offer a wide range of ecosystem services that are of high environmental and socioeconomic value. Yet, nearly 70% of BHFs in the southern United States have been lost during the past 100 years primarily due to land use change including agricultural expansion, calling for restoration efforts. We estimated the statistical relationship of the annual radial growth rate of three red oak species with climatic conditions and tree age using the tree ring data collected from a BHF plantation in the Arkansas Delta region. These species were Cherry bark oak (Quercus pagoda), Shumard oak (Quercus shumardii), and Nuttall oak (Quercus texana). The destructive sampling method was employed to obtain tree growth data and the cross-dating method was used for tree age determination. A log-linear regression model was estimated to uncover the statistical relationship between annual tree ring growth rate and climatic conditions. We identified the most critical time windows of climate variables that affect the growth of these trees. We found that the average temperature in October of the previous year and the minimum temperature between December of the previous year and January of the current year were positively associated with the radial growth rate in the current year although the maximum temperature from January to August and total precipitation from April to July of the current year were negatively correlated with the growth rate. Compared to Cherry bark and Shumard oaks, Nuttall oak was less sensitive to a rise in the minimum temperature between December and January. The projected climate change is likely to create slightly more favorable overall climatic conditions for these oak species in the region. Our findings suggest that these three red oak species are well suited for the study region for restoring BHFs, especially with a changing climate.

Wildfire Variable Toxicity: Identifying Biomass Smoke Exposure Groupings through Transcriptomic Similarity Scoring.

The prevalence of wildfires continues to grow globally with exposures resulting in increased disease risk. Characterizing these health risks remains difficult due to the wide landscape of exposures that can result from different burn conditions and fuel types. This study tested the hypothesis that biomass smoke exposures from variable fuels and combustion conditions group together based on similar transcriptional response profiles, informing which wildfire-relevant exposures may be considered as a group for health risk evaluations. Mice (female CD-1) were exposed via oropharyngeal aspiration to equal mass biomass smoke condensates produced from flaming or smoldering burns of eucalyptus, peat, pine, pine needles, or red oak species. Lung transcriptomic signatures were used to calculate transcriptomic similarity scores across exposures, which informed exposure groupings. Exposures from flaming peat, flaming eucalyptus, and smoldering eucalyptus induced the greatest responses, with flaming peat grouping with the pro-inflammatory agent lipopolysaccharide. Smoldering red oak and smoldering peat induced the least transcriptomic response. Groupings paralleled pulmonary toxicity markers, though they were better substantiated by higher data dimensionality and resolution provided through -omic-based evaluation. Interestingly, groupings based on smoke chemistry signatures differed from transcriptomic/toxicity-based groupings. Wildfire-relevant exposure groupings yield insights into risk assessment strategies to ultimately protect public health.

Stem profile of red oaks in a bottomland hardwood restoration plantation forest in the Arkansas Delta (USA)

Bottomland hardwoods are among the most diverse and productive forest ecosystems in the southeastern United States and are critically important for the provision of timber and non-timber ecosystem services. Red oaks, the dominant species in this group of forests, are of high ecological and economic value. Stem profile models are essential for accurately estimating the merchantable volume of oak trees, which is also closely indicative of total tree biomass and other ecosystem services given their allometric relationships. This study aims to develop and compare stem profiles among three red oak species in an 18-year old plantation forest using destructive sampling. Sixty trees randomly selected from an oak restoration plantation in the Arkansas Delta were felled for measuring the diameter-outside-bark (DOB) and diameter-inside-bark (DIB) at different stem heights. These sample composed of twenty trees from each of three species: cherry bark oak (CBO - Quercus pagoda Raf), Nuttall oak (NUT - Quercus texana Buckley), and Shumard oak (SHU - Quercus shumardii Buckl). Multiple models, including the segmented-profile model, form-class profile model, and second-and third-order polynomial models were fitted and compared. Results demonstrate that the form-class profile model was the best fitted for CBO and NUT, whereas the third-order polynomial model was the best for SHU. CBO tends to grow taller and has a higher wood density than NUT and SHU. These findings will inform restoration and management decisions of bottomland hardwood forests, especially red oaks in the region.

ПРОДУКТИВНОСТЬ КУЛЬТУР ДУБА КРАСНОГО КУРДЖИПСКОГО УЧАСТКОВОГО ЛЕСНИЧЕСТВА РЕСПУБЛИКИ АДЫГЕЯ

The results of a study of red oak crops located on the territory of the Kurdzhip precinct forestry of the Republic of Adygea (western part of the North Caucasus) are presented. The characteristic of red oak, as a tree species, is less demanding on soil conditions compared to other forest-forming species and has high decorative properties, due to which it is used alone for planting in open areas, as a hedge along roads, individual tracts in parks, squares in the form of oak groves. It goes well both in pure stands and in combination with pine, beech, hornbeam, elm, ash, linden, mountain ash, maple. It is able to grow on sandy loam and loamy podzolic soils without stagnant moisture, mainly along river banks. It grows poorly on calcareous and moist soils. Due to its well-developed root system, wood is resistant to wind. Red oak is characterized by good resistance to damage by entomo - and phyto pests [1, 5]. The paper analyzes the taxation indicators of red oak and related species crops, and plots of growth progress along the height and diameter of the crops are constructed. The growth and productivity advantage of the oak over red oak has been established.

Nondestructive Evaluation of Red Oak and White Oak Species

Abstract The objective of this article is to evaluate the relationship between the dynamic modulus of elasticity (MOEd), which was obtained with acoustic-based nondestructive testing (NDT) methods, and static bending properties of two domestic hardwood oak species. The mechanical properties were conducted using static modulus of elasticity (MOE) and modulus of rupture (MOR) in radial and tangential directions. Mechanical tests were performed according to ASTM D143 on small clear, defect-free specimens from the two tree species: red oak (Quercus rubra) and white oak (Quercus alba). The MOEd was determined by two NDT methods and three longitudinal vibration methods based on the fast Fourier transform. The destructive strength values obtained in this study were within the expected range for these species. The MOE was best predicted by NDT methods for both species but also had a strong capability to predict MOR.

Soil nitrogen assimilation of 1-year oak seedlings: Implication for forest fertilization and management

Reserves in cotyledons and nutrients in soil are essential for the early development of seedlings of oaks. However, much less is known to what extent the reserves in seeds and/or soil nutrients contribute to seedling development and growth at early stages, especially at multi-species level. We carried out an indoor experiment in which the soil was enriched in 15N to quantify N uptake and its distribution in the seedlings of five red and white oak species. Dry masses of seedlings did not vary significantly with N fertilization rates, reflecting the low morphological and physiological plasticity of oak seedlings to soil fertility. However, dry masses of roots of white oak species were much higher than those of red oak species. Soil-derived N incorporated into the seedlings of the five oak species increased with seedling growth and enhanced 15N fertilization rates. Soil-derived N in the seedlings of white oak species was much higher than those of red oak species. Acorn-derived N in seedlings decreased with N fertilization rates, implying that acorn N mobilization can be accelerated under low soil nutrient level. We found higher soil N/acorn N ratio in the seedlings of white oak species than those of red oak species, indicating the rapid uptake of soil N in the seedlings of white oak species. Acorn N rather than soil N played a crucial role in supporting oak seedling development and growth at the early stage. Our results provide insight into the role of acorn N and soil fertility to 1-year oak seedlings and have great implication for forest fertilization and management.

Біолого-екологічні особливості поширення омели білої (Viscum album) в умовах міста Вінниця

У межах населених пунктів, зокрема й у Вінниці, омела біла набула інтенсивного поширення. Ураження дерев омелою призводить як до погіршення стану дерев, їх подальшого всихання, так і до зниження естетичної оцінки насаджень. Незважаючи на інтенсивне пошкодження деревних насаджень омелою у Вінниці, достатньо стійкими виявилися дерева більшості хвойних видів, зокрема: сосна, ялина, дугласія. Із листяних видів достатньо високою стійкістю відрізняється дуб, граб, бук. Незначною мірою пошкоджується омелою каштан, дуб червоний, горіх та ін. Поряд із цим, більшість алейних насаджень, зокрема тополі та липи, істотно пошкоджені цим напівпаразитом. Це зумовлено не лише особливістю деревних порід, але й і їхнім віком. Зокрема, більшість дерев старшого віку із наявними пошкодженнями стовбурової частини та гілок інтенсивніше уражаються омелою. За результатами проведених досліджень виявлено, що вік дерев, які найбільш уражені омелою білою, становить 50-60 років і з віком ступінь ураження зростає, а частка ураження дерев віком 70-80 років становить 25-29 %. Ступінь ураження дерев омелою білою визначено за 5-бальною шкалою, згідно з якою найвищий бал ураження виявлено у тополі чорної та дельтовидної (5 балів), найнижчий – у ялини звичайної, липи серцелистої (1-2 бали). Розглядаючи види Populus за інтенсивністю заселення їх Viscum album, встановлено, що низький і високий ступені ураження мають 26 % дерев, середній – 28 %, а дуже високий – 30 %. Найбільшою часткою дерев з високим і дуже високим ступенем ураження відзначаються P. nigra та P. deltoids – 22 і 32 %.

Molecular evolution of the internal transcribed spacers in red oaks (Quercus sect. Lobatae)

Previous studies of the Internal Transcribed Spacers of the nuclear ribosomal DNA (ITS) in sections Quercus (white oaks), Protobalanus (intermediate or golden cup oaks), Cerris (Cerris oaks), and Ilex (Ilex oaks) suggest that ITS regions undergo full concerted evolution in oaks; however, ITS evolution patterns in red oaks (section Lobatae) are unknown due to scant representation in published work. To determine whether full concerted evolution occurs in red oaks, the purpose of this study was to examine ITS sequences from 40 red oak species. The results show incomplete concerted evolution and the presence of three ITS ribotypes of lengths 505, 609, 601 bp, hereafter referred to as ITS-S (small), I ITS-M (medium), and ITS-L (large), respectively. Thirty species had only one ribotype (ITS-M), nine species had two ribotypes (different combinations of ITS-L, ITS-M, and ITS-S), and only one species had all three ribotypes. Furthermore, examination of these three ribotypes showed that only ITS-M is putatively functional and ITS-L and ITS-S are pseudogenes. Bayesian analysis strongly supported (100%) two pseudogenes clades but provided weak support for the monophyly of a putative functional clade (ITS-M); moreover, within the “functional” clade, species relationships were uncertain and, in most cases, sequences from the same species failed to group together. The results of the current study suggest that ITS may not be appropriate for phylogeny reconstruction of red oaks due to low levels of interspecific variation and incomplete concerted evolution.

The Influence of Dormancy Break Requirements on Germination and Viability Responses to Winter Submergence in Acorns of Three Bottomland Red Oak (Sect. Lobatae) Species

Abstract The objective of this study was to examine the relation of dormancy break requirements to winter submergence effects on germination and viability in cherrybark, willow, and Nuttall oak acorns. Acorns were submerged in a greentree reservoir and received 0, 21, 42, 63, or 84 days of winter submergence, followed by 8 weeks of incubation in four temperature regimes (15/6°, 20/10°, 25/15°, and 30/20° C). Winter submergence substituted for cold stratification requirements of cherrybark oak acorns, and dormancy break was achieved with 63 days of submergence. Although winter submergence exerted a positive effect on germination in Nuttall and willow oak acorns, dormancy break was not achieved in acorns of either species. Germination percentages were highest in the 30/20° C incubation temperature—cherrybark oak acorns (75–83 percent), followed by Nuttall oak (33–55 percent) and willow oak (14–52 percent). Among treatment combinations, all willow oak nongerminants were viable. Viability loss in cherrybark and Nuttall oak acorns was greatest in the 63- and 84-day treatments. The dynamics of acorn germination in response to winter hydrologic regime is influenced by submergence duration that may, or may not, satisfy dormancy break requirements in acorns of these red oak species.

Responses of seedling growth and survival to post‐germination cotyledon removal: An investigation among seven oak species

Abstract Rodents regularly rely on emerged epicotyls to locate and remove cotyledons still containing valuable nutrients. However, the extent to which acorn characteristics influence tolerance to post‐germination predation has received little attention. Here, we investigated the impact of cotyledon removal following epicotyl emergence on seedling performance and survival of seven oak (Quercus) species. We imitated cotyledon predation at different stages of seedling establishment and development in order to detect effects on seedling height, leaf number and tissue/component mass. Seedling growth and survival were negatively affected by cotyledon loss regardless of oak species. However, these negative effects decreased as the epicotyl length at which cotyledons were removed increased. We also found that there was a threshold epicotyl length above which seedling survival and performance were relatively unaffected in white oak species compared to red oak species. Following cotyledon removal, early germinating white oak (section Quercus) seedlings survived and/or grew better than the late germinating red oak (section Lobatae) seedlings. This was likely caused by a difference in dependence on cotyledon reserves, which ultimately affected the ability of seedlings to tolerate cotyledon removal. Synthesis. From an evolutionary perspective, this is likely to follow from the early germination in white oaks and the ability of seed consumers to locate young seedlings from the emerging epicotyls. Our study has implications for forest regeneration by suggesting additional opportunities for white oak species to establish following epicotyl emergence. Future studies should consider quantifying the rates of post‐germination cotyledon loss.

Experimental evidence for selection against hybrids between two interfertile red oak species

Abstract Reproductive isolation between related oak species within one taxonomic section is incomplete. Even though pre- and post­zygotic isolation mechanisms have been described for interfer­tile oak species, natural hybridization is common in contact zones between related oaks. The apparent restriction of inter­specific hybrids between ecologically divergent species to intermediate environments in contact zones suggests postzy­gotic isolation via environmental selection against hybrids in parental environments. Overrepresentation of hybrids in seeds as compared to adult trees provides additional indirect evi­dence for selection against hybrids. Here, we used genetic assignment analyses in progeny obtained from a sympatric stand of Quercus rubra and Quercus ellipsoidalis, two interfertile species with different adaptations to drought, to characterize the number of hybrids and “pure” species in the non-germinated acorns and in seedlings. The occurrence of 43.6 % F1 hybrids and introgressive forms among the non-germinated acorns and their very low frequency in the seedlings (9.3 %) is to our knowledge the first direct evidence for early selection against hybrids in oaks possibly as result of genetic incompatibilities. Additionally, we found a signature of positive selection on EST-SSR PIE200 in Q. rubra which needs further confirmation. These results contribute to our under­standing of reproductive isolation and divergence between interfertile oak species with different ecological adaptations.

Leaf morphological and genetic variation between Quercus rubra and Quercus ellipsoidalis: comparison of sympatric and parapatric populations

Species boundaries in oaks are often not clear-cut, which is potentially a result of interspecific hybridization with trait introgression and phenotypic plasticity. Quercus rubra L. and Quercus ellipsoidalis E.J. Hill are two interfertile partially sympatric red oak species (section Lobatae) with different adaptations to drought. Quercus ellipsoidalis is the most drought tolerant of the North American red oak species and is characterized by deep tap roots, a shrubby growth and by deeply dissected leaves. Genetic differentiation between species is low for most molecular markers. However, one genic microsatellite in a CONSTANS-like (COL) gene, FIR013, was previously identified as outlier locus under strong divergent selection between species. In this study, we analyzed leaf morphometric traits in neighboring (parapatric) Q. rubra/Q. ellipsoidalis populations and in one sympatric population from the same region along an environmental gradient. Using multivariate statistics of leaf traits both species showed distinct bimodal frequency distributions for the first canonical discriminant function with some overlap in the phenotypic extremes, especially in the sympatric population. Leaf dissection traits showed strong and consistent differentiation between species in sympatric and parapatric populations, while differentiation for leaf size was lower in the sympatric population under more similar environmental conditions. Leaf phenotypes in F1 hybrids and introgressive forms suggested maternal effects and introgression of leaf traits between species. The association of outlier gene copy number at FIR013 with species-discriminating leaf traits in Quercus rubra can be a reflection of population differences since outlier gene copy number and population membership show significant collinearity. Similar environmental selection pressures on outlier alleles and leaf shape could also have resulted in this association. In future studies, segregating full-sib families could be used to test whether outlier alleles and associated genomic regions are indeed associated with leaf traits or other species-discriminating characters.

Growth, survival and sunfleck response of underplanted red oaks (Quercus spp., section Erythrobalanus) along a topographic gradient in southern New England

An understanding of the regeneration niche is critical to our ability to refine silvicultural approaches aimed at regenerating and maintaining diverse, mixed species stands. However, few studies have directly tested differences in habitat associations among closely related trees of the same genus, particularly to assess habitat partitioning at the regeneration stage. To evaluate differences in regeneration niche, we established a common garden experiment with five red oak species (Quercus spp., section Erythrobalanus) under-planted across a topographic gradient (valley, midslope and ridge positions) in mixed hardwood forests of Southern New England. Represented were three red oak species common to intermediate site conditions of southern New England (Q. rubra L, Q. velutina Lam., and Q. coccinea Münchh.), and two species commonly associated with more extreme site conditions - Q. ilicifolia Wangenh. (xeric, skeletal ridge tops) and Q. palustris Münchh. (river floodplains). Seedlings were caged to prevent deer browse, and seedling height and survival were measured annually for nine growing seasons. Response to sunflecks, a primary source of light for advance regeneration-dependent species that establish in the forest understory, was measured using a LiCor 6400 photosynthesis system. Overall, survival was highest for Q. rubra, followed by Q. velutina, Q. coccinea, Q. ilicifolia and finally Q. palustris. The pattern among species held for all topographic positions, with higher survival rates on the ridgetops, and lower survival in valley positions. Height development was highest in the valley, intermediate in the ridge, and lowest in the midslope positions. Survival patterns were positively correlated with light availability, and negatively correlated with site fertility. Based on net photosynthetic rates, Q. rubra, Q. velutina, and Q. coccinea were more responsive to sunflecks than Q. ilicifolia and Q. palustris, and maintained higher photosynthetic rates over the measurement period. Greater survival of all species in upper topographic position reflects in part adaptations of the red oak group in this region to higher light and lower soil fertility conditions. But sunflecks likely play a critical role as well, judging from the positive correlation of survival with understory light (43–80% of which occurs as sunflecks), the strong height development in the light-limited valleys (where sunflecks should be most critical), and the clear alignment of species sunfleck response with species survival patterns. Differences among the oaks in photosynthetic response to sunflecks may thus help explain their habitat associations and contribute to niche partitioning among these related species. These findings have important silvicultural implications, particularly for enrichment planting of different red oaks and the timing of overstory removal treatments to facilitate establishment of regeneration for growth into the canopy.

The evolution and diversification of the red oaks of the California Floristic Province (Quercus section Lobatae, series Agrifoliae).

The California Floristic Province (CA-FP) is a unique and diverse region of floral endemism, yet the timing and nature of divergence and diversification of many lineages remain underexplored. We seek to elucidate the evolutionary history of the red oaks of the CA-FP, the Agrifoliae. We collected PstI-associated RAD-seq data as well as morphometrics from individuals of the four species across their ranges, including varieties and hybrids. Phylogeny and divergence times were estimated. We analyzed morphological differentiation in over 70 plants using PCA and assessed species delimitation and admixture using genotype clustering analysis in over 40 plants. We find that the Agrifoliae are monophyletic and sister to all other red oak species. Within the Agrifoliae, all species are supported, with Quercus kelloggii sister to a clade of subevergreen taxa: (Quercus agrifolia - (Q. parvula + Q. wislizeni)). Molecular and morphometric analyses are equivocal for named varieties. Notably, Q. parvula var. tamalpaisensis appears to be part of a hybrid swarm between Q. parvula and Q. wislizeni. Dating estimates were concordant with previous hypotheses and geological evidence, with diversification occurring between 10 and 20 million years ago. The Agrifoliae represent a geographically discrete, early-diverging red oak lineage that diversified during the period of drying and warming associated with Sierran uplift during the middle Miocene. Molecular differentiation within the clade supports the current taxonomy, including an east-west species level pattern (Q. parvula and Q. wislizeni) and north-south intraspecific patterns to some degree, although the latter require additional study.

Evidence for environment-dependent introgression of adaptive genes between two red oak species with different drought adaptations.

Introgression of adaptive alleles between hybridizing species is likely an important mechanism to generate new genetic variation for adaptation to rapidly changing environmental conditions. Oaks provide a model for the study of adaptive gene introgression because environmental selection maintains high interspecific differentiation at a few outlier loci and species-specific adaptations despite recurrent interspecific gene flow. Previously, we identified a CONSTANS-like gene under strong divergent selection between drought-tolerant Quercus ellipsoidalis and drought-averse Quercus rubra. Neighboring (parapatric) populations of both species were fixed for alternate alleles, Q. ellipsoidalis for allele 138 and Q. rubra for allele 141. In this study, we analyzed introgression of these outlier alleles in sympatric and in parapatric Q. rubra/Q. ellipsoidalis populations. We found evidence that environmental selection affected the level and direction of outlier allele introgression. Thus, in the face of symmetric interspecific gene flow, outlier allele introgression was asymmetric and introgression of allele 138 into Q. rubra was consistently higher than introgression of allele 141 into Q. ellipsoidalis in sympatric populations, but the opposite pattern was found in parapatric populations. Furthermore, the rate of introgression of outlier alleles between species in sympatric stands was related to soil quality (i.e., soil water holding capacity, nutrient availability). This pattern suggests that introgression of adaptive genes between these two red oak species is strongly affected by environmental selection and is an important mechanism for species' adaptation to changing environmental conditions in a changing climate.

Differentiation in the water-use strategies among oak species from central Mexico.

Oak species (Fagaceae: Quercus) differ in their distribution at the landscape scale, specializing to a certain portion of environmental gradients. This suggests that functional differentiation favors habitat partitioning among closely related species. To elucidate the mechanisms of species coexistence in oak forests, we explored patterns of interspecific variation in functional traits involved in water-use strategies. We tested the hypothesis that oak species segregate along key trade-offs between xylem hydraulic efficiency and safety, and between hydraulic safety and drought avoidance capacity, leading to species niche partitioning across a gradient of aridity. To do so, we quantified biophysical and physiological traits in four red and five white oak species (sections Lobatae and Quercus, respectively) across an aridity gradient in central Mexico. We also explored the trade-offs guiding species differentiation, particularly between the drought tolerance versus water acquisition capacity, and determined whether the water-use strategy was associated with the portion of the environmental gradient that the species occupy. In a trait-by-trait analysis, we detected differences between white and red oak species. However, a larger part of the variation was explained at the species rather than at the section level. We detected two primary axes of trait covariation. The first exhibited differences between species with dense tissues and species with soft tissues (the tissue construction cost axis); however, the oak sections did not constitute separate groups, while the second suggested a trade-off between xylem resistance to cavitation and tree deciduousness. As expected, the water-use strategies of the species were related to the environment; oak species from arid areas had more deciduousness and a higher instantaneous water-use efficiency. In contrast, their humid counterparts had less deciduousness and had a xylem that was more resistant to embolisms. Altogether, these results suggest that aridity filters closely related species, resulting in habitat partitioning and niche divergence.