Variation in morphological traits in a recent hybrid zone between closely related Quercus liaotungensis and Q. mongolica (Fagaceae)

Aims Hybridization usually leads to gene introgression between related species in hybrid zones, associated with complex patterns of morphological variation. Nevertheless, previous studies have tended to ignore the effects of geographic variation in hybridization rates on species taxonomy. This study aims to investigate the variation of morphological traits between two sympatric and taxonomically confused oak species, Quercus liaotungensis and Q. mongolica, and reveal the effects of hybridization rates on morphological traits and the taxonomic boundary.Methods We used seven microsatellite loci to evaluate species status and measured 15 morphological traits in 26 trees in the recent hybrid zone between Q. liaotungensis and Q. mongolica, and we characterized the differences between the two oak species and their hybrids for the investigated traits.Important findings Molecular analyses indicated that 74% of 78 sampled maternal trees were hybrids between Q. liaotungensis and Q. mongolica although the observed morphological variation suggested that they had remained distinct species. Across all of the differentiated leaf and reproductive traits, the hybrids expressed patterns similar to Q. liaotungensis, which may suggest dominant expression of parental characters. These results are consistent with our expectation that hybrids will be difficult to distinguish from parental species in a recent hybrid zone.     

雷公青冈和滇南青冈学名订正

分布于华南的雷公青冈Quercus hui Chun和分布于云南及广西的滇南青冈Q.austroglauca(Y.T.Chang)Y.T.Chang长期被认为是我国特有植物,实则分别为越南北部分布的Q.auricoma A.Camus和云南东南部分布的龙迈青冈Q.lungmaiensis(Hu)C.C.Huang & ...     

A phenetic study of the taxonomic delimitation of Quercus acutifolia and Q. conspersa (Fagaceae)

A phenetic study ofQuercus acutifolia andQ. conspersa (Fagaceae) is carried out. The study was based on morphological characters, and included cluster analysis and principal component analysis. The results revealed that the characters employed so far to separateQuercus acutifolia andQ. conspersa seem to be quite unreliable. Nevertheless, both species can still be recognized as taxonomically different entities, although mostly based on characters of leaf margins. We propose that the individuals that have leaves with dentate-aristate margins correspond toQ. acutifolia, whereas those that have leaves with entire and not dentate-aristate margins belong toQ. conspersa.     

Leaf epidermis of Quercus subgen. Cyclobalanopsis (Oerst.) Schneid. (Fagaceae)

The characters of leaf epidermis (mainly epidermal cells, stomata, trichomes) have been examined in 48 species of Quercus subgen. Cyclobalanopsis of the Fagaceae. The adaxial leaf epidermis was investigated under light microscope. The epidermal cells of this surface are of two types in terms of the outline of their anticlinal wall. One type is characterized by having straight wall, and this type has been observed in most of the species studied, while the other is characterized by having sinuate or sinuolate wall, and this type has been found only in eight species, such as Q. blakei and Q. hui. There exist two types of trichome-bases on this surface: in one type the bases are singlecelled, and this type has been found in most of the species studied, such as Q. oxyodon and Q. augustinii; while in the other the bases consist of a group of cells, and this type has been found only in nine species, such as Q. thorelii and Q. chungii. The abaxial leaf epidermis was investigated under light microscope and scanning electron microscope. The epidermal cells and the trichome-bases on this surface under LM are in morphology essentially similar to those on the adaxial surface. The stomata under LM include three types: cyclocytic, subcyclocytic and anomocytic. The stomatal density is of some diagnostic value to distinguish between closely related species, e.g. the stomatal density in Q. glauca is higher than that in Q. schottkyana. Under SEM the stomata are elliptical or roundish. The most noteworthy feature of the adaxial leaf epidermis is the trichomes. Eight types of trichomes have been observed under SEM: simple-uniseriate, solitary, papillae, stellate, fused-stellate, stipitate-fasciculate, multiradiate and jellyfish-like. The first type, i.e. the simple-uniseriate trichome, is glandular, other six types, i.e. the solitary, papillae, stellate, fused-stellate, stipitate-fasciculate and multiradiate trichomes, are non-glandular, and the last one, i.e. the jellyfish-like trichome, may represent an intermediate type in structure between the glandular and the non-glandular trichomes. Papillae are very common in subgen. Cyclobalanopsis although in the Fagaceae they were previously reported only in a few species of Lithocarpus. The jellyfish-like trichome, observed only in Q. sichouensis, is reported in the Fagaceae for the first time. The evolutionary trends of trichomes are discussed. The trichomes seem to evolve toward the direction of increasing complexity in structure. Papillae are considered as the pedomorphism of trichomes. From papillae the solitary trichomes and then the stellate ones have evolved. The complicated trichome types with more arms may be produced from relatively simple ones with fewer arms by means of varions fusion: several stellate trichomes are fused to form the fused-stellate ones by basal fusion, and by the same way the fused-stellate ones to form the stipitate-fasciculate ones; the multiradiate trichomes are formed by the stellate ones, in which the arms emerge in a variety of seemingly random directions from a typically rounded common base. Based on the above results, it is considered that the trichome features are of important significance for a better understanding of the infrageneric division of Quercus and the phylogenetic relationships between this genus and the other genera in the Fagaceae. The very common presence of papillae in Quercus subgen. Cyclobalanopsis and in a few species of Lithocarpus may indicate that subgen. Cyclobalanopsis is a quite natural group on the one hand, and may have close relationship with Lithocarpus on the other hand. The thin-walled, jellyfish-like trichomes found in Q. sichouensis are in structure similar to the “glandular” peltate trichomes in Trigonobalanus, a genus generally regarded as the most primitive in subfam. Quercoideae, indicating that the affinity between subgen. Cyclobalanopsis and Trigonobalanus may becloser than that between subgen. Quercus and Trigonobalanus.     

栎属青冈亚属（壳斗科）的叶表皮研究

利用光镜和扫描电镜观察了栎属青冈亚属Quercus subgen. Cyclobalanopsis 48种植物的叶表皮,尤以对叶下表皮的毛被特征观察较为仔细。共观察到8种不同类型的叶表皮毛：单列毛、单毛、乳突、星状毛、溶和星状毛、具柄束毛、多出毛和水母状毛。其中乳突在青冈亚属中较常见,而在壳斗科其他属中仅在石栎属 Lithocarpus 少数种类中有报道;水母状毛首次在壳斗科中发现。毛被可能遵循以下的演化规律：乳突→单毛→星状毛;星状毛依照从简单→复杂的演化途径,分化出各种形态各异和结构复杂的毛系。初步讨论了毛被以及叶表皮其他特征(如毛基细胞和表皮细胞的形态、气孔的类型和密度等)的分类和系统学意义。     

Typification of the Linnaean name Ononis crispa and the related O. zschackei (Fabaceae)

The typification of the Linnaean name Ononis crispa and the related O. zschackei (Fabaceae) is discussed. Original material for O. crispa was traced at LINN, but found to be heterogeneous. In addition, the illustration by Magnol cited in the protologue represents O. aragonensis. The name is lectotypified using a specimen at LINN to preserve the current usage of the name and to avoid any ambiguity in the interpretation of the lectotype, an epitype is selected. Since no original material is extant for the name O. zschackei, a neotype is designated from a specimen preserved at P and collected in Mallorca (Balearic Islands, Spain).     

The effects of seed size, cotyledon reserves, and herbivory on seedling survival and growth in Quercus rugosa and Q. laurina (Fagaceae)

In a greenhouse experiment, seedling survival of two oak species (Quercus rugosa and Q. laurina) was greatly affected by the excision of cotyledons 1 mo after germination, with a greater impact on Q. laurina. The effect of seed size was also significant for both species, with a positive correlation between seed mass and survival and growth. The effect of cotyledon excision on seedling growth persisted throughout the first growing season in Q. rugosa and was not analyzed for Q. laurina due to the low number of seedlings that survived cotyledon excision. Seed size significantly affected seedling height, diameter, leaf area, and biomass at 6 mo. Seed size and cotyledon retention affected the ability of Q. rugosa to recover from herbivory, as both factors had a significant effect on relative growth rates after aerial biomass removal. The results show that seedlings originating from large seeds can better endure loss of cotyledons and aerial biomass and thus are better equipped to confront stress early in their lives.     

中国栎属植物的数量分类研究

在全面衡量中国栎属49种4变种性状特征的基础上,观测了25个定性性状和18个数量性状。通过聚类分析,可以将中国栎属分为5个特征明显的表征群,即5组:麻栎组(Section Aegilops)、槲栎组(Section Quercus)、高山栎组(Section Brachylepides)、巴东栎组(Section Engleriana)和橿子栎组(Section Echinolepides)。在主成分分析中,虽然性状的累积贡献率增长不明显,前3个主成分累积贡献率仅有50.4%,但结果仍然显示出和聚类分析同样的结果。说明栎属的分类性状仍存在主要方面,只不过在演化过程中由于性状变异的多样性,产生了多个不同的表征类群。分析结果对我们解决个别有疑问种的分类地位也有启示作用。因此,数量分类方法对解决栎属组间及种间的分类问题有重要的意义。     

Clonal population structure and genetic variation in sand-shinnery oak,Quercus havardii (Fagaceae)

We investigated clonal population structure and genetic variation in Quercus havardii (sand-shinnery oak), a deciduous rhizomatous shrub that dominates vegetation by forming uninterrupted expanses of ground cover over sandy deposits on the plains of western Texas, western Oklahoma, and eastern New Mexico. Isozyme electrophoresis (15 loci coding 11 enzymes) was used to recognize and map clones arrayed in a 2000-m transect (50-m sample intervals) and a 200 × 190 m grid (10-m sample intervals). Ninety-four clones were discovered, 38 in the transect and 56 in the grid, resulting in an estimated density of ～15 clones per hectare. Clones varied greatly in size (～100-7000 m), shape, and degree of fragmentation. The larger clones possessed massive interiors free of intergrowth by other clones, while the smaller clones varied in degree of intergrowth. The population maintained substantial levels of genetic variation (P = 60%, A = 2.5, H(exp) = 0.289) comparable to values obtained for other Quercus spp. and for other long-lived perennials. The population was outcrossing as evidenced by conformance of most loci to Hardy-Weinberg expected genotype proportions, although exceptions indicated a limited degree of population substructuring. These data indicate that despite apparent reproduction primarily through vegetative means, Q. havardii possesses conventional attributes of a sexual population.     

Genetic Variation in Fragmented Forest Stands of the Andean Oak Quercus humboldtii Bonpl. (Fagaceae)1

Quercus humboldtii is a montane forest dominant species in Colombia, which has experienced significant habitat loss. Using three microsatellite loci, we compared the genetic diversity of adults and seedlings in fragments of small and large size. Results show high genetic diversity, comparable to other temperate oak species (H_o= 0.813, H_e= 0.780, and f=?0.044). However, allelic richness reduction in seedlings of the most fragmented part of the landscape, suggested restricted gene flow and risk of future genetic bottlenecks, if larger tracts of forest disappear.     

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.     

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.     

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.     

Population structure, genetic diversity, and clone formation in Quercus chrysolepis (Fagaceae)

Stands of canyon live oak (Quercus chrysolepis, Fagaceae) are maintained for fuelwood, fire management, recreation, and as habitat for wildlife. Information about the link between the oak's reproductive ecology and its extent of genetic diversity is important in developing land management policies that will maintain the long-term viability of populations. Basal sprouting is the primary means of reproduction following fire or cutting, and stands frequently include groups of visibly connected trees in a clustered distribution that suggests cloning. We determined the extent to which clusters of trees were clonal and defined the spatial pattern and diversity of genotypes for six populations across nearly the entire east-west extent of the San Bernardino Mountains in southern California. We mapped over 100 trees at each of five sites and genotyped each tree for allozymes at seven polymorphic loci. We identified clones using these multilocus genotypes and detected an average of 34.4 ± 7.3 (SD) clones per site, most of which had unique genotypes. In general, clustered trees belong to single clones and most clones consist of few trees (mean = 3.4 ± 0.6 trees per clone). However, clone size increased significantly with increased individual heterozygosity, suggesting that selection may favor highly heterozygous clones. Clonal diversity and evenness were high relative to reports for most other clonal species; an average of 97% of clones had distinct genotypes, and Simpson's index of diversity averaged 0.95 ± 0.02. Population genetic analyses of 319 clones from six sites revealed high genetic diversity within sites (mean HS = 0.443). Only a small proportion of the total genetic diversity was explained by variation among sites (mean GST = 0.018), which is consistent with high gene flow among sites (Nm = 9.5). We found no significant substructure among plots within sites, and fixation indices within sites were generally small, suggesting that either little inbreeding occurs, and/or few inbred progeny survive. However, spatial autocorrelation analysis of clones indicated fine-scale genetic structure at distances under 4 m, possibly due to limited seed dispersal. Our data suggest that guidelines for seed collection of canyon live oak for use in restoration can be specified in a manner similar to that recommended for conifer species within the region studied.     

倒卵叶青冈——广西壳斗科一新记录种

报道了广西壳斗科植物一新记录种——倒卵叶青冈,并对倒卵叶青冈的形态进行了补充描述,同时对该种的生存现状进行评价,为这一物种的保护提供基础资料。     

Anatomical and developmental study of petrified Quercus (Fagaceae) fruits from the Middle Miocene, Yakima Canyon, Washington, USA

The first reported petrified acorns to show internal anatomical structure are here described from Middle Miocene (～15.6 million years old) chert of the Columbia River Basalt Group in Yakima Canyon, Washington. Quercus hiholensis Borgardt et Pigg sp. nov. is described from anatomical and morphological fruit features, as well as a little recognized anatomical feature, the umbilical complex. Acorns, each comprising a nut and its cupule, are up to 15.3 mm long and 18.8 mm wide with helically arranged, imbricate, tuberculate cupule scales. They show basal aborted ovules, short styles, broad stigmas, and lack grooves in their cotyledons. These characters and the developmental pattern seen in these fossil acorns demonstrate that Q. hiholensis conforms to genus Quercus (Fagaceae), subgenus Quercus, section Quercus (the white oaks). The correspondence of Q. hiholensis to the modern section Quercus reveals that the derived floral and fruit characters that distinguish section Quercus within the genus had evolved by the Middle Miocene.     

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.     

Electrophoretic Analysis of Genetic Variation within and between Populations of Quercus cerris,Q. pubescens,Q. petraea and Q. robur (Fagaceae) from Eastern Austria

Allozyme variation at 14 loci was studied electrophoretically in 19 population samples of Quercus cerris, Q. pubescens, Q. petraea, and Q. robur. Genetic variation, measured as mean heterozygosity, was found to be high within populations but low among populations. A significant deficit of heterozygotes was observed in most of the populations of Q. pubescens, Q. petraea, and Q. robur. At the species level, Q. cerris (sect. Cerris) appears to be clearly separated from the rest (sect. Quercus). Q. pubescens, Q. petraea, and Q. robur form a cluster of closely-related species. The degree of genetic differentiation among these species is low, i.e. in the range normally found among con-specific populations. Nevertheless, allozyme data allow the elaboration of a dendrogram which is in reasonable accordance with the taxonomic classification, but also supports the importance of hybridization and introgression. This is also underlined by the fact that the allelic differentiation at the Got-2 locus corresponds with the phenotypic classification (petraea-like, hybrids, and robur-like) in samples from mixed populations.     

Development and characterization of polymorphic microsatellite markers in Cyclobalanopsis glauca (Fagaceae)

? Premise of the study: We isolated and characterized polymorphic microsatellite loci in Cyclobalanopsis glauca (Fagaceae), an evergreen broadleaved monoecious tree, to provide tools for analyzing parentage and mating system. ? Methods and Results: Thirteen polymorphic microsatellite markers were developed and tested in three C. glauca populations. The number of alleles per locus varied from two to 22. The observed and expected heterozygosities within populations were 0.000-0.967 and 0.033-0.949, respectively. ? Conclusions: These polymorphic primers showed high levels of polymorphism within tested populations, and can be used in parentage analysis and mating system estimation of C. glauca.