Taxonomy of Quercus crassifolia (Fagaceae) and morphologically similar species in Mexico

In this work, ten species morphologically similar to Q. crassifolia are circumscribed based on a phylogenetic species concept. Examination of these species suggests that Q. dysophylla and Q. radiata are of hybrid origin; the putative parents of Q. dysophylla are Q. crassifolia and Q. crassipes, while Q. radiata is likely the result of hybridization between Q. coccolobifolia and Q. conzattii or Q. urbanii. A lectotype of Q. crassifolia is designated here. Dichotomous keys, as well as distribution ranges and photographs, are presented.     

Natural hybridization and hybrid zones between Quercus crassifolia and Quercus crassipes (Fagaceae) in Mexico: morphological and molecular evidence

Hybrid zones provide interesting systems to study genetic and ecological interaction between different species. The correct identification of hybrids is necessary to understand the evolutionary process involved in hybridization. An oak species complex occurring in Mexico formed by two parental species, Quercus crassifolia H. & B. and Q. crassipes H. & B., and their putative hybrid species, Q. dysophylla, was analyzed with molecular markers (random amplified polymorphic DNA [RAPDs]) and morphological tools in seven hybrid zones (10 trees per taxa in each hybrid zone) and two pure sites for each parental species (20 trees per site). We tested whether geographic proximity of hybrid plants to the allopatric site of a parental species increases its morphological and genetic similarity with its parent. Seventeen morphological traits were measured in 8700 leaves from 290 trees. Total DNA of 250 individuals was analyzed with six diagnostic RAPD primers. Quercus crassifolia differed significantly from Q. crassipes in all the examined characters. Molecular markers and morphological characters were highly coincident and support the hypothesis of hybridization in this complex, although both species remain distinct in mixed stands. Clusters and a hybrid index (for molecular and morphological data) showed that individuals from the same parental species were more similar among themselves than to putative hybrids, indicating occasional hybridization with segregation in hybrid types or backcrossing to parents. Evidence does not indicate a unidirectional pattern of gene flow.     

Landscape-level spatial genetic structure in Quercus acutissima (Fagaceae)

Quercus acutissima (Fagaceae), a deciduous broad-leaved tree, is an important forest element in hillsides of South Korea. We used allozyme loci, Wright's F statistics, and multilocus spatial autocorrelation statistics to examine the distribution of genetic diversity within and among three local populations and the spatial genetic structure at a landscape scale (15 ha, 250 × 600 m) on Oenaro Island, South Korea. Levels of genetic diversity in Q. acutissima populations were comparable to mean values for other oak species. A moderate but significant deficit of heterozygotes (mean F(IS) = 0.069) was detected within local populations and low but significant differentiation was observed among populations (F(ST) = 0.010). Spatial autocorrelation analyses revealed little evidence of significant genetic structure at spatial scales of 100-120 m. The failure to detect genetic structure within populations may be due to intraspecific competition or random mortality among saplings, resulting in extensive thinning within maternal half-sib groups. Alternatively, low genetic differentiation at the landscape scale indicates substantial gene flow among local populations. Although wind-borne pollen may be the primary source of gene flow in Q. acutissima, these results suggest that acorn movement by animals may be more extensive than previously anticipated. Comparison of these genetic data for Oenaro Island with a disturbed isolated inland population suggests that population-to-population differences in internal genetic structure may be influenced by local variation in regeneration environment (e.g., disturbance).     

Quercus tuitensis (Fagaceae, Quercus sect. Lobatae), a new deciduous oak from western Jalisco, Mexico

Quercus tuitensis, a deciduous red oak known only from the Sierra El Tuito on the Pacific slopes of western Jalisco, Mexico, is described, illustrated, and compared toQ. praineana Trel. In addition,Q. aequivenulosa andQ. coffeaecolor are reaffirmed as synonyms ofQ. praineana.     

Tuberculate ectomycorrhizae of angiosperms: The interaction between Boletus rubropunctus (Boletaceae) and Quercus species (Fagaceae) in the United States and Mexico

Tuberculate ectomycorrhizae (TECM) are unique structures in which aggregates of ectomycorrhizal roots are encased in a covering of fungal hyphae. The function of TECM is unknown, but they probably enhance the nitrogen nutrition and disease resistance of host plants. Trees in the Pinaceae form TECM with species of Rhizopogon and Suillus (Suillineae, Boletales). Similar tubercules are found with diverse angiosperms, but their mycobionts have not been phylogenetically characterized. We collected TECM in Mexico and the USA that were similar to TECM in previous reports. We describe these TECM and identify both the plant and fungal symbionts. Plant DNA confirms that TECM hosts are Quercus species. ITS sequences from tubercules and sclerotia (hyphal aggregations that serve as survival structures) matched sporocarps of Boletus rubropunctus. Phylogenetic analyses confirm that this fungus belongs to the suborder Boletineae (Boletales). This is the first published report of TECM formation in the Boletineae and of sclerotia formation by a Boletus species. Our data suggest that the TECM morphology is an adaptive feature that has evolved separately in two suborders of Boletales (Suillineae and Boletineae) and that TECM formation is controlled by the mycobiont because TECM are found on distantly related angiosperm and gymnosperm host plants.     

Effect of a red oak species gradient on genetic structure and diversity of Quercus castanea (Fagaceae) in Mexico

Incipient reproductive barriers are a common characteristic of oak species. Disruption of these barriers promotes changes in diversity and genetic structure of the species involved. Quercus castanea is a red oak with wide geographic distribution in Mexico, which presents atypically high morphological variability when it occurs in sympatry with other red oak species, suggesting that hybridization may explain the observed variation. We tested if the genetic structure and diversity levels of Q. castanea are related to the number of red oak species growing in sympatry. In total, 14 microsatellite (SSRs) primers (six nSSRs and eight cpSSRs) were used in 120 Q. castanea individuals (20/site) belonging to six populations, where the number of red oak species associated varied from zero to five. Results showed a positive and significant relationship between the genetic diversity of Q. castanea and the number of red oak species growing in sympatry, regardless of the marker type or the parameter of genetic diversity analyzed. Also, we found a higher genetic differentiation of Q. castanea populations using cpSSRs in comparison with nSSRs. Our results suggest that temperate forests with high red oaks species richness co-dominated by Q. castanea promote the increase in this species genetic diversity. From a conservation perspective, high genetic diversity levels of foundation species such as Q. castanea may have positive cascade effects extending to other species in the community.     

Environmental suitability for Agrilus auroguttatus (Coleoptera: Buprestidae) in Mexico using MaxEnt and database records of four Quercus (Fagaceae) species

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Lectotypification of Quercus arbutifolia (Fagaceae) and the taxonomic treatment of Quercus subsect. Chrysotrichae

The name Quercus arbutifolia Hickel & A. Camus is lectotypifed. In addition, leaf epidermal features of Q. obovatifolia C. C. Huang, Q. arbutifolia and four other closely related species, Q. daimingshanensis (S. K. Lee) C. C. Huang, Q. merrillii Seemen, Q. litseoides Dunn and Q. tiaoloshanica Chun & W. C. Ko, were studied and compared using herbarium collections. Based on a comparison of leaf epidermal and other key taxonomic features, we conclude that Q. obovatifolia and Q. arbutifolia are identical. We thus reduce Q. obovatifolia to a new synonym of Q. arbutifolia. A key to the studied species is included.     

Microsatellite markers for northern red oak (Fagaceae: Quercus rubra)

We provide primer sequences for 14 (GA) _n microsatellite loci developed from northern red oak, an important timber species. We screened loci using two sets of samples. A parent–offspring set included DNA from seven acorns collected from one mother tree along with maternal DNA, to determine that all progeny carried a maternal allele at each locus. The other set was comprised of 10 adult trees sampled from Indiana old‐growth forest, providing a measure of diversity revealed by each locus.     

Competitive suppression of Quercus douglasii (Fagaceae) seedling emergence and growth

Reduced recruitment of blue oak (Quercus douglasii) seedlings in California grasslands and woodlands may result from shifts in seasonal soil water availability coincident with replacement of the native perennial herbaceous community by Mediterranean annuals. We used a combination of container and field experiments to examine the interrelationships between soil water potential, herbaceous neighborhood composition, and blue oak seedling shoot emergence and growth. Neighborhoods of exotic annuals depleted soil moisture more rapidly than neighborhoods of a perennial grass or "no-neighbor" controls. Although effects of neighborhood composition on oak seedling root elongation were not statistically significant, seedling shoot emergence was significantly inhibited in the annual neighborhoods where soil water was rapidly depleted. Seedling water status directly reflected soil water potential, which also determined the extent and duration of oak seedling growth during the first year. End-of-season seedling height significantly influenced survival and growth in subsequent years. While growth and survival of blue oak seedlings may be initially constrained by competition with herbaceous species, subsequent competition with adult blue oak trees may further contribute to reduced sapling recruitment.     

Two new species of oak (Fagaceae, Quercus sect. Lobatae) from the Sierra Madre del Sur, Mexico

Two new species of oak (Fagaceae) with narrowly ellipticlanceolate leaves from the Sierra Madre del Sur of western Jalisco, Mexico, are described and illustrated.Quercus cualensis is a local endemic andQ. iltisii a more widespread species; they are closely related toQ. salicifolia Née.     

Spatial and genetic structure of two sandhills oaks: Quercus laevis and Quercus margaretta (Fagaceae)

Quercus laevis Walt, (turkey oak) and Q. margaretta Ashe (scrubby post oak) are important scrub oaks in the sandhills forest communities of the Coastal Plain of the southeastern United States. We used allozyme loci and Ripley's L-statistics to examine clonal structure and spatial dispersion in these species. Q. laevis greater than 1.5 m in height were randomly dispersed on a scale of 0–40 m; smaller individuals (< 1.5 m) were slightly clustered on a scale of 0–12 m. Larger individuals separated by ≤1 m had 15% probability of being ramets of the same clone. Q. margaretta showed strong clustering on a scale of 0–20 m. Stems separated by ≤ 1 m had a 71% probability of being ramets of the same clone. Clonal offspring were strongly clustered about the presumed clonal parent: 50% fell within 0.50 m of this individual. Simulation modeling and direct comparison of adult and juvenile genotypes indicated that acorns are dispersed on a scale of tens of meters for both species, suggesting animal vectors such as squirrels or blue jays.     

Limited hybridization between Quercus lobata and Quercus douglasii (Fagaceae) in a mixed stand in central coastal California

Many oak species are interfertile, and morphological and genetic evidence for hybridization is widespread. Here we use DNA microsatellite markers to characterize hybridization between two closely related oak species in a mixed stand in central coastal California, Quercus lobata (valley oak) and Q. douglasii (blue oak) (Fagaceae). Genotypes from four microsatellite loci indicate that many alleles are shared between the two species. However, each species harbors unique alleles, and allele frequencies differ significantly. A Bayesian analysis of genetic structure in the stand identified two highly differentiated genetic clusters, essentially corresponding to species assignment based on morphology. Data from the four loci were sufficient to assign all 135 trees to one of the two species. In addition, five putative hybrid individuals having intermediate morphologies could be assigned genetically to one or the other species, and all but one had low probability of hybrid ancestry. Overally, only six (4.6%) trees showed >0.05 probability of hybrid ancestry, in all cases their probabilities for nonhybrid ancestry were substantially higher. We conclude that adult hybrids of Q. douglasii × Q. lobata are rare at this site and plasticity in morphological characters may lead to overestimates of hybridization among Quercus species.     

Clonal and genetic structure of two Mexican oaks: Quercus eduardii and Quercus potosina (Fagaceae)

Quercus eduardii and Q. potosina are dominant oak species in Sierra Fría, Aguascalientes, Mexico. These species have been exploited for multiple purposes since the 16th century. Both species produce clonal offspring through root suckering and acorns through sexual reproduction. To understand clonality for the implementation of the most adequate actions for the conservation of these species, we addressed the following questions: (a) what is the spatial clonal structure of both species? (b) How much clonal and genetic diversity is maintained in these species? Random Amplified Polymorphic DNAs (RAPDs) were used as molecular markers for these analyses. Genets of both species have few ramets and these grow close the parent tree. Autocorrelation analyses at the ramet level showed an aggregated distribution at short distances and a random spatial distribution at larger distances. Also, at the genet level the autocorrelation analyses showed a random distribution. Clonal diversity was high in both species (Q. eduardii: D=0.963, G/N=0.60; Q. potosina: D=0.985, G/N=0.65). Genetic diversity was high within populations (Q. eduardii: H _e =0.33±0.11; Q. potosina: H _e =0.35±0.11). Low levels of genetic differentiation among populations were observed (Q. eduardii ? _st =0.19, P < 0.002; Q. potosina ? _st =0.13, P < 0.002). Both species maintain high levels of clonal and genetic diversity, probably due to successful sexual reproduction, which allows gene flow among populations. Conservation and/or reforestation programs must include seed collections and germplasm banks. Due to the small genet size and the high clonal diversity of these species, seeds can be collected in any place in Sierra Fría, Aguascalientes.     

Epigenetic marks in the mature pollen of Quercus suber L. (Fagaceae)

We have analysed the distribution of epigenetic marks for histone modifications at lysine residues H3 and H4, and DNA methylation, in the nuclei of mature pollen cells of the Angiosperm tree Quercus suber; a monoecious wind pollinated species with a protandrous system, and a long post-pollination period. The ultrasonic treatment developed for the isolation of pollen nuclei proved to be a fast and reliable method, preventing the interference of cell wall autofluorescence in the in situ immunolabelling assays. In contrast with previous studies on herbaceous species with short progamic phases, our results are consistent with a high level of silent (5-mC and H3K9me2) epigenetic marks on chromatin of the generative nucleus, and the prevalence of active marks (H3K9me3 and H4Kac) in the vegetative nucleus. The findings are discussed in terms of the pollination/fertilization timing strategy adopted by this plant species.     

Polyphenol deposition in leaf hairs of Olea europaea (Oleaceae) and Quercus ilex (Fagaceae)

The subcellular localization (cytoplasm, vacuoles, cell walls) of polyphenol compounds during the development of the multicellular nonglandular leaf hairs of Olea europaea (scales) and Quercus ilex (stellates), was investigated. Hairs of all developmental stages were treated with specific inducers of polyphenol fluorescence, and the bright yellow-green fluorescence of individual hairs was monitored with epifluorescence microscopy. During the early ontogenetic stages, bright fluorescence was emitted from the cytoplasm of the cells composing the multicellular shield of the scales of O. europaea. Transmission electron micrographs of the same stages showed that these cells possessed poor vacuolation and thin cell walls. The nucleus of these cells may be protected against ultraviolet-B radiation damage. The progressive vacuolation that occurred during maturation was followed by a shifting of the bright green-yellow fluorescence from the perinuclear region and the cytoplasm to the cell walls. The same trends were observed during the development of the nonglandular stellate hairs of Quercus ilex, in which maturation was also accompanied by a considerable secondary thickening of the cell walls. Despite the differences in morphology, high concentrations of polyphenol compounds are initially located mainly in the cytoplasm of the developing nonglandular hairs, and their deposition on the cell walls takes place during the secondary cell wall thickening. These structural changes during the development of the leaf hairs make them a very effective barrier against abiotic (uv-B radiation) and probably biotic (pathogenic) stresses.     

Post-fire recolonization of dominant epiphytic lichen species on Quercus hypoleucoides (Fagaceae)

Following a forest fire (27 500 ha) in 1994, post-fire recolonization of Quercus hypoeleucoides by epiphytic lichens was documented as changes in lichen cover, number of small thalli, specific factors that affected reestablishment of lichens, and modes of dispersal. Three sites in the Chiricahua Mountains (Arizona, USA) were chosen according to the severity of fire damage—unburned, moderately burned, and severely burned. From 1994 through 1997, the amount of dead lichen cover significantly increased at the moderately burned site. For the same time period, the amount of live lichen cover significantly increased at the severely burned site. Numbers of new thalli increased significantly at the severely burned site each year but only in the last year (1996–1997) for the moderately burned site. Bark texture and proximity to trees with lichens were among the most important physical factors for recolonization. The most important means of dispersal for Flavopunctelia praesignis was fragmentation. For Punctelia hypoleucites, the primary means of dispersal was spores. Increases in live lichen cover and numbers of new thalli occur faster in severely burned areas probably due to the loss of lichens on tree trunks, which provides space and a lack of competition.