Pollen grain morphology supports the taxonomical discrimination of Mediterranean oaks (Quercus,Fagaceae)

The exine architecture and some quantitative parameters (polar and equatorial axes and their ratio) of pollen grains allow a clear discrimination between groups of Mediterranean oak species. For the differentiation of species within groups only the quoted parameters prove useful.     

Seed-mass effects in four Mediterranean Quercus species (Fagaceae) growing in contrasting light environments

Three hypotheses have been proposed to explain the functional relationship between seed mass and seedling performance: the reserve effect (larger seeds retain a larger proportion of reserves after germinating), the metabolic effect (seedlings from larger seeds have slower relative growth rates), and the seedling-size effect (larger seeds produce larger seedlings). We tested these hypotheses by growing four Mediterranean Quercus species under different light conditions (3, 27, and 100% of available radiation). We found evidence for two of the three hypotheses, but none of the four species complied with all three hypotheses at the same time. The reserve effect was not found in any species, the metabolic effect was found in three species (Q. ilex, Q. pyrenaica, and Q. suber), and the seedling-size effect in all species. Light availability significantly affected the relationships between seed size and seedling traits. For Q. ilex and Q. canariensis, a seedling-size effect was found under all three light conditions, but only under the lowest light (3%) for Q. suber and Q. pyrenaica. In all species, the correlation between seed mass and seedling mass increased with a decrease in light, suggesting that seedlings growing in low light depend more upon their seed reserves. A causal model integrates the three hypotheses, suggesting that larger seeds generally produced larger seedlings.     

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).     

Stem xylem features in three Quercus (Fagaceae) species along a climatic gradient in NE Spain

 Stem xylem features in two evergreen Quercus species (Q. coccifera and Q. ilex) and a deciduous one (Q. faginea) were analysed along an Atlantic-Mediterranean climatic gradient in which rainfall and winter cold experience strong variation. Mean maximum vessel diameter, vessel density, vessel element length, xylem transverse sectional area, Huber value (xylem transverse sectional area per leaf area unit), theoretical leaf specific conductivity (estimated hydraulic conductance per leaf area unit) and total leaf area were determined in 3-year-old branches. Q. faginea presented the widest vessels and the highest theoretical leaf specific conductivity while Q. coccifera showed the lowest total leaf area and the highest Huber value. Studied features did not exhibit significant correlations with mean minimum January temperature in any species but did show significant relationships with rainfall. In Q. coccifera, mean maximum vessel diameter, vessel element length and theoretical leaf specific conductivity increased with higher rainfall while vessel density decreased. Mean maximum vessel diameter and total leaf area in Q. ilex increased with precipitation whereas variables of Q. faginea did not show any significant trend. Results suggest that aridity, rather than minimum winter temperature, controls stem xylem responses in the studied evergreen species. Q. faginea traits did not show any response to precipitation, probably because this species develops deep roots, which in turn makes edaphic and topographic factors more important in the control of soil water availability. In response to aridity Q. coccifera only exhibits adjustment at a xylem level by reducing its water transport capacity through a reduction of vessel diameter without changing the amount of xylem tissue or foliage, whereas Q. ilex adjusts its water transport capacity in parallel to the foliage area. Received: 13 January 1997 / Accepted: 8 April 1997     

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.     

Comparative leaf surface morphology and molecular data of five oaks of the subgenus Quercus Oerst (Fagaceae)

Abstract

Comparisons between different data sets are important in taxonomic studies, particularly for complex taxa such as the genus Quercus. However, there is scarce literature on comparative morphological and molecular analyses in oaks. Here we describe an experimental approach based on the correlation between micromorphological features and molecular data in order to characterize and discriminate among five closely related species: Q. robur, Q. petraea, Q. frainetto, Q. pubescens and Q. virgiliana.     

European and Mediterranean oaks (Quercus L.; Fagaceae): SEM characterization of the micromorphology of the abaxial leaf surface

This paper examines micromorphological characteristics (trichomes, waxes, stomata) of the abaxial leaf surface of European and Mediterranean oaks. Observations were performed In-scanning electron microscopy, and the purpose of the investigation was to ascertain whether it is possible to distinguish the different species taxonomically using these features. Several species, as well as some subspecific entities and taxa considered doubtful, were taken into consideration. Subgenera appear to be well-defined: e.g. the subgenus Quercus is characterized by waxes arranged in vertical scales, the subgenera Cenis and Sclerophyllodris by smooth waxes. They differ because the former has markedly elliptical stomata, while the latter two have roundish stomata. The different species are also fairly easy to distinguish, and this fact confirms the taxonomical validity of the parameters we have taken into consideration. In some cases subspecific entities and doublful species do not diflcr at all from the related species, but in others they present such marked variations that it is difficult even to classify them with certainty. The most critical taxonomic groups (i.e. those which include a number of specific and subspecific entities often too difficult to distinguish from each other) are the series Quercus robur – Quercus petraea,– Quercus pubescens and the Quercus faginea,– Quercus Insitanica Quercus canariensis group.     

A comparative flower and fruit anatomical study of Quercus acutissima, a biennial-fruiting oak from the Cerris group (Fagaceae)

A developmental series of flowers and fruits of Quercus acutissima (subgenus Quercus section Cerris) was collected over a growing season and examined for an intersectional, comparative anatomical study. Pistillate flowers of the current growing season, each consisting of a pistil with three long, slightly recurved styles, six tepals, and an inconspicuous ovary subtended by a few cycles of cupule scales, emerged in early May, were pollinated by mid-May, and then were quiescent for the remainder of the growing season. Flowers from the previous growing season resumed growth in mid-May, each forming three locules delimited by septa in the ovary, with two bitegmic, epitropous ovules developing in each locule. Mature embryo sacs were present by mid-July of the second growing season, although embryos were not observed until early August. Fruit maturation was complete by late September. Features that have not been described previously for the section Cerris include early-lignifying endocarp trichomes, persistent septa, and leaf primordia buttresses on the embryo. A comparison of flower and fruit developmental features with sections Quercus sensu stricto and Lobatae revealed a mosaic of shared features among the three sections.     

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.     

A hierarchical analysis of minisatellite DNA diversity in Gambel oak (Quercus gambelii Nutt.; Fagaceae)

Gambel oak is a short shrub to medium-sized tree of southwest North America for which the potential to grow in large clonal stands has been proposed. Here we use three different multilocus VNTR DNA probes (synthesized via PCR) to demonstrate that clones growing together can be identified, but individual clones exceeding 50 m in diameter do not commonly occur in the study population of Quercus gambelii . Further, using a hierarchical sampling scheme (three circular transects with diameters of 9, 1100, and 28 000 m), we estimate that for nonclonal individuals: (i) mean number of bands analysed per individual = 22.92; (ii) mean similarity (band sharing) between individuals = 0.322; (iii) mean probability that two randomly chosen individuals share all bands = 4.93 × 10^–11; and (iv) mean estimated heterozygosity = 0.796. F _ST calculated for the two nonclonal levels of the sampling hierarchy was 0.023, indicating that little genetic differentiation exists between them. These results support previous findings that, due to life-history traits of the genus and this species (out-crossing, wind-pollinated, animal-dispersed, long-lived woody perennials), gene flow is high and genetic subdivision of populations is low in oaks. At this study site, the clonal nature demonstrated for Gambel oak appears to have little detectable effect on these population genetic characteristics.     

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.     

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.     

Leaf morphology and leaf chemical composition in three Quercus (Fagaceae) species along a rainfall gradient in NE Spain

 Leaf features were examined in three Quercus species (Q. coccifera, Q. ilex and Q. faginea) along a steep rainfall gradient in NE Spain. The analyzed leaf traits were area, thickness, density, specific mass, leaf concentration of nitrogen, phosphorous, lignin, cellulose and hemicellulose, both on a dry weight basis (N_w, P_w, L_w, C_w, H_w) and on an area basis (N_a, P_a, L_a, C_a, H_a). These traits were regressed against annual precipitation and correlated with each other, revealing different response patterns in the three species. Q. faginea, a deciduous tree, did not show any significant correlation with rainfall. In Q. coccifera, an evergreen shrub, N_w, N_a, L_w, L_a and C_a increased with higher annual rainfall, while H_w decreased. In Q. ilex, an evergreen tree, leaf area, P_w and L_w increased with precipitation, whereas specific leaf mass, thickness and H_a showed the reverse response. Correlations between the leaf features revealed that specific mass variation in Q. faginea and Q. coccifera could be explained by changes in leaf density, while in Q. ilex specific leaf mass was correlated with thickness. Specific leaf mass in the three species appeared positively correlated with all the chemical components on a leaf area basis except with lignin in Q. ilex and with P in Q. ilex and Q. faginea. In these two tree species P_w showed a negative correlation with specific leaf mass. It is suggested that each species has a different mechanism to cope with water shortage which is to a great extent related to its structure as a whole, and to its habit. Received: 18 December 1995 / Accepted: 8 March 1996     

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.     

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.     

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.     

Woodland canopy structure and the light response of juvenile Quercus lobata (Fagaceae)

I investigated competition for light between canopy plants and juvenile valley oaks (Quercus lobata Nee) in a mixed-broadleaf woodland of California's northern Coast Ranges. Canopy effects on understory light supply were separated among the overlying adult valley oak, the adult's woody understory, and neighboring trees and shrubs through a series of light sampling surveys and measurements of the number, size, and spatial distribution of neighboring plants. Light supply in the understory was primarily influenced by neighboring plants, with no detectable effect of the overlying adult valley oak. Light supply in the understory averaged 25% full sun due to a high frequency of canopy gaps and a typically open understory. Seedling response to understory light supply was investigated in an experimental sunfleck gradient (10%, 19%, and 100% full sun). Between 10% and 100% full sun, seedling growth increased by 90% and the shoot:∗∗∗root ratio changed from 1.561 to 0.607. Shade seedlings were also taller and produced fewer, larger, and thinner leaves than seedlings grown in full sun. A field survey of the spatial distribution and crown morphology of saplings and young adults found 1) the distance between young valley oaks and neighboring overstory trees to increase with neighbor size, and 2) crowns of the young oaks to be skewed away from neighbors. Although shading by the canopy was only moderate, canopy effects on understory light supply may restrict juvenile recruitment of valley oak in this woodland.