Seeing the forest through the trees: comprehensive inference on individual mating patterns in a mixed stand of Quercus robur and Q. petraea

Background and Aims

Sexual reproduction is one of the most important moments in a life cycle, determining the genetic composition of individual offspring. Controlled pollination experiments often show high variation in the mating system at the individual level, suggesting a persistence of individual variation in natural populations. Individual variation in mating patterns may have significant adaptive implications for a population and for the entire species. Nevertheless, field data rarely address individual differences in mating patterns, focusing rather on averages. This study aimed to quantify individual variation in the different components of mating patterns.

Methods

Microsatellite data were used from 421 adult trees and 1911 seeds, structured in 72 half-sib families collected in a single mixed stand of Quercus robur and Q. petraea in northern Poland. Using a Bayesian approach, mating patterns were investigated, taking into account pollen dispersal, male fecundity, possible hybridization and heterogeneity in immigrant pollen pools.

Key Results

Pollen dispersal followed a heavy-tailed distribution (283 m on average). In spite of high pollen mobility, immigrant pollen pools showed strong genetic structuring among mothers. At the individual level, immigrant pollen pools showed highly variable divergence rates, revealing that sources of immigrant pollen can vary greatly among particular trees. Within the stand, the distribution of male fecundity appeared highly skewed, with a small number of dominant males, resulting in a ratio of census to effective density of pollen donors of 5·3. Male fecundity was not correlated with tree diameter but showed strong cline-like spatial variation. This pattern can be attributed to environmental variation. Quercus petraea revealed a greater preference (74 %) towards intraspecific mating than Q. robur (36 %), although mating preferences varied among trees.

Conclusions

Mating patterns can reveal great variation among individuals, even within a single even-age stand. The results show that trees can mate assortatively, with little respect for spatial proximity. Such selective mating may be a result of variable combining compatibility among trees due to genetic and/or environmental factors.     

Dissecting apple tree architecture into genetic, ontogenetic and environmental effects: QTL mapping

The present study aimed to dissect tree architectural plasticity into genetic, ontogenetic and environmental effects over the first 4 years of growth of an apple F1 progeny by means of quantitative traits loci (QTL) mapping. Both growth and branching processes were phenotyped on the consecutive annual shoots of different axes within a tree. For each studied trait, predicted values (best linear unbiased predictors, BLUPs) of the genotypic (G) effect or its interaction with tree age (G×A) and climatic year (G×Y) were extracted from mixed linear models of repeated data. These BLUPs, which are independent from autocorrelations between repeated measurements, were used for QTL mapping. QTL detection power was improved by this two-step approach. For each architectural process, numerous QTLs were detected and some particularly interesting co-localised in common genomic regions, for internode lengthening, top diameter, and number and percentage of axillary shoots. When several QTLs were detected for a given trait, global models were estimated, which explained a maximum of 40% of the total variance for both internode length and top diameter and 28% for branching. QTLs detected for BLUPs of G×Y effects were interpreted as resulting from the interaction between genetic maximal potential of growth and climatic factors, while those for G×A effects were interpreted in relation to tree ontogeny. Most of the latter ones were found to be concomitant with key development stages during which the trait average started to decrease, but with different magnitudes depending on genotype. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Pure species in a continuum of genetic and morphological variation: sympatric oaks at the edge of their range

Background and Aims Studies on oaks (Quercus spp.) have often been hampered by taxonomic confusion, a situation further compounded by the occurrence of extensive interspecific hybridization. In the present study, a combination of genetic and morphological analyses was used to examine sympatric populations of Q. petraea and Q. robur at the north-western edge of their ranges in Northern Ireland, since it had previously been suggested that hybridization could facilitate the apparent rapid, long-distance dispersal of oaks following the glaciations.Methods Samples were collected from 24 sites across Northern Ireland that had been previously designated as ancient or semi-natural woodland. Genotypes were obtained from a total of 950 trees using 12 nuclear microsatellite loci, and admixture coefficients were calculated based on a Bayesian clustering approach. Individuals were also classified as Q. petraea, Q. robur or hybrids based on two objective morphometric characters shown previously to delineate pure individuals effectively. Genetically ‘pure’ individuals of both species, as defined by the Bayesian clustering, were also genotyped for five chloroplast microsatellites.Key Results Genetic and morphological analyses both indicated the presence of pure individuals of both species, as well as a continuum of intermediates. There was a good agreement between the molecular and morphological classification, with a generally clear separation between pure individuals.Conclusions Despite millennia of hybridization and introgression, genetically and morphologically pure individuals of both Q. petraea and Q. robur can be found at the edge of their range, where both species occur sympatrically. The high proportion of individuals exhibiting introgression compared with previous studies may reflect the historical role of hybridization in facilitating dispersal following the glaciations. This is further supported by the significantly higher chloroplast diversity in Q. robur compared with Q. petraea.     

A stochastic model of tree architecture and biomass partitioning: application to Mongolian Scots pines

Background and Aims

Mongolian Scots pine (Pinus sylvestris var. mongolica) is one of the principal species used for windbreak and sand stabilization in arid and semi-arid areas in northern China. A model-assisted analysis of its canopy architectural development and functions is valuable for better understanding its behaviour and roles in fragile ecosystems. However, due to the intrinsic complexity and variability of trees, the parametric identification of such models is currently a major obstacle to their evaluation and their validation with respect to real data. The aim of this paper was to present the mathematical framework of a stochastic functional–structural model (GL2) and its parameterization for Mongolian Scots pines, taking into account inter-plant variability in terms of topological development and biomass partitioning.

Methods

In GL2, plant organogenesis is determined by the realization of random variables representing the behaviour of axillary or apical buds. The associated probabilities are calibrated for Mongolian Scots pines using experimental data including means and variances of the numbers of organs per plant in each order-based class. The functional part of the model relies on the principles of source–sink regulation and is parameterized by direct observations of living trees and the inversion method using measured data for organ mass and dimensions.

Key Results

The final calibration accuracy satisfies both organogenetic and morphogenetic processes. Our hypothesis for the number of organs following a binomial distribution is found to be consistent with the real data. Based on the calibrated parameters, stochastic simulations of the growth of Mongolian Scots pines in plantations are generated by the Monte Carlo method, allowing analysis of the inter-individual variability of the number of organs and biomass partitioning. Three-dimensional (3D) architectures of young Mongolian Scots pines were simulated for 4-, 6- and 8-year-old trees.

Conclusions

This work provides a new method for characterizing tree structures and biomass allocation that can be used to build a 3D virtual Mongolian Scots pine forest. The work paves the way for bridging the gap between a single-plant model and a stand model.     

Pathogenicity of Bursaphelenchus xylophilus on Pines in Minnesota and Wisconsin

The pinewood nematode, Bursaphelenchus xylophilus, was inoculated into established native jack and red pines (Pinus banksiana and P. resinosa) and exotic Austrian pine (P. nigra) in Minnesota and Wisconsin forests during summer 1981. The nematode isolates did not kill established nonstressed pine trees growing in the forest. However, the same nematode isolates killed pine seedlings under greenhouse conditions. Girdling the main stem of some trees to induce stress resulted in the death of the majority of inoculated and noninoculated branches of Austrian and jack pines, but no branch death was observed on red pine. Greater numbers of nematodes were extracted from branches of inoculated, girdled trees than from nongirdled trees. The mean number of nematodes extracted from branches of inoculated, nongirdled trees was 0.3 - 14 nematodes per gram of wood.     

思茅松天然林树冠结构模型

以云南省普洱市思茅区思茅松天然林为研究对象,采用枝解析调查了34株思茅松样木的树冠数据,分析了一级枝枝长、枝径、着枝角度、弦长和树冠半径5个树冠形状变量的变化规律,分别构建其预估模型;分析了树冠结构变化,分别构建了一级枝轮枝高度预估模型、一级枝枝条数量预估模型和一级枝枝条数量累积预估模型,并采用独立样本进行模型统计精度检验。结果表明:8个预估模型的预测效果良好,精度达到91%以上,尤其是一级枝着枝角度模型和一级枝轮枝高度模型预测精度达到97%以上。研究结果合理准确描述思茅松树冠结构的变化,为思茅松天然林的经营管理提供科学依据。     

Leaf-traits and growth allometry explain competition and differences in response to climatic change in a temperate forest landscape: a simulation study

Background and Aims

The ability to simulate plant competition accurately is essential for plant functional type (PFT)-based models used in climate-change studies, yet gaps and uncertainties remain in our understanding of the details of the competition mechanisms and in ecosystem responses at a landscape level. This study examines secondary succession in a temperate deciduous forest in eastern China with the aim of determining if competition between tree types can be explained by differences in leaf ecophysiological traits and growth allometry, and whether ecophysiological traits and habitat spatial configurations among PFTs differentiate their responses to climate change.

Methods

A temperate deciduous broadleaved forest in eastern China was studied, containing two major vegetation types dominated by Quercus liaotungensis (OAK) and by birch/poplar (Betula platyphylla and Populus davidiana; BIP), respectively. The Terrestrial Ecosystem Simulator (TESim) suite of models was used to examine carbon and water dynamics using parameters measured at the site, and the model was evaluated against long-term data collected at the site.

Key Results

Simulations indicated that a higher assimilation rate for the BIP vegetation than OAK led to the former''s dominance during early successional stages with relatively low competition. In middle/late succession with intensive competition for below-ground resources, BIP, with its lower drought tolerance/resistance and smaller allocation to leaves/roots, gave way to OAK. At landscape scale, predictions with increased temperature extrapolated from existing weather records resulted in increased average net primary productivity (NPP; +19 %), heterotrophic respiration (+23 %) and net ecosystem carbon balance (+17 %). The BIP vegetation in higher and cooler habitats showed 14 % greater sensitivity to increased temperature than the OAK at lower and warmer locations.

Conclusions

Drought tolerance/resistance and morphology-related allocation strategy (i.e. more allocation to leaves/roots) played key roles in the competition between the vegetation types. The overall site-average impacts of increased temperature on NPP and carbon stored in plants were found to be positive, despite negative effects of increased respiration and soil water stress, with such impacts being more significant for BIP located in higher and cooler habitats.     

Effect of Copper Sulfate and Lead Acetate on Infection of Pines with Bursaphelenchus xylophilus

Treatment of 3-year-old Scots, white, and Austrian pine seedlings with copper sulfate or lead acetate significantly affected energy homeostasis and oleoresin production in the seedlings but did not induce wilting of the seedlings. Inoculation of copper sulfate-treated or lead acetate-treated white, Scots, and Austrian pine seedlings with the white pine specific pathotype of Bursaphelenchus xylophilus, VPSt-1, caused a significant increase in oleoresin production, stressed energy homeostasis, and induced rapid wilting of the seedlings. Scots pine lost tolerance and Austrian pine lost resistance to VPSt-1 after the seedlings were treated with either copper sulfate or lead acetate. These results suggest that environmental pollution may significantly affect susceptibility of pines to B. xylophilus and may have a role in establishment of this nematode in uninfested areas.     

Genomic Differences among Pathotypes of Bursaphelenchus xylophilus

Total genomic DNA from Bursaphelenchus xylophilus pathotypes MPSy-1 and VPSt-1 and from B. mucronatus was digested with restriction endonucleases. DNA fragments were electrophoretically separated, Southern blotted to nitrocellulose, and hybridized to genomic DNA from one of the isolates. The resulting hybridization patterns indicate genomic differences in repetitive DNA sequences among these populations. Greatest differences were seen between B. xylophilus and B. mucronatus, but genomic differences were also apparent between B. xylophilus pathotypes MPSy-1 and VPSt-1 and between a population from P. nigra in New Jersey and a population of a mucronate form from Abies balsamea in Quebec, Canada.     

Seasonal, daily and diurnal variations in the stable carbon isotope composition of carbon dioxide respired by tree trunks in a deciduous oak forest

The stable C isotope composition (δ¹³C) of CO₂ respired by trunks was examined in a mature temperate deciduous oak forest (Quercus petraea). Month-to-month, day-to-day and diurnal, measurements were made to determine the range of variations at different temporal scales. Trunk growth and respiration rates were assessed. Phloem tissue was sampled and was analysed for total organic matter and soluble sugar ¹³C composition. The CO₂ respired by trunk was always enriched in ¹³C relative to the total organic matter, sometimes by as much as 5‰. The δ¹³C of respired CO₂ exhibited a large seasonal variation (3.3‰), with a relative maximum at the beginning of the growth period. The lowest values occurred in summer when the respiration rates were maximal. After the cessation of radial trunk growth, the respired CO₂ δ¹³C values showed a progressive increase, which was linked to a parallel increase in soluble sugar content in the phloem tissue (R = 0.95; P < 0.01). At the same time, the respiration rates declined. This limited use of the substrate pool might allow the discrimination during respiration to be more strongly expressed. The late-season increase in CO₂ δ¹³C might also be linked to a shift from recently assimilated C to reserves. At the seasonal scale, CO₂ δ¹³C was negatively correlated with air temperature (R = −0.80; P < 0.01). The diurnal variation sometimes reached 3‰, but the range and the pattern depended on the period within the growing season. Contrary to expectations, diurnal variations were maximal in winter and spring when the leaves were missing or not totally functional. By contrast to the seasonal scale, these diurnal variations were not related to air temperature or sugar content. Our study shows that seasonal and diurnal variations of respired ¹³C exhibited a similar large range but were probably explained by different mechanisms.     

Characterization of a Nonparasitic Isolate of Bursaphelenchus xylophilus

Bursaphelenchus xylophilus isolate MPSy-1av was subcultured from pathotype MPSy-1. MPSy-1av is nonparasitic and does not establish in Pinus sylvestris, P. strobus, P. nigra, or P. taeda. This isolate produces ethanol as an end product of carbohydrate metabolism, whereas its parent pathotype, MPSy-1, does not. Alcohol dehydrogenase activity was easily detectable in homogenates of MPSy-1av but barely detectable in some homogenates of MPSy-1. Genomic differences were seen between MPSy-1 and M PSy-1av by restriction endonuclease analysis of total nematode DNA, and hybridization of DNA fragments to the alcohol dehydrogenase gene from Drosophila.     

Isometric scaling of above- and below-ground biomass at the individual and community levels in the understorey of a sub-tropical forest

Background and Aims Empirical studies and allometric partitioning (AP) theory indicate that plant above-ground biomass (M_A) scales, on average, one-to-one (isometrically) with below-ground biomass (M_R) at the level of individual trees and at the level of entire forest communities. However, the ability of the AP theory to predict the biomass allocation patterns of understorey plants has not been established because most previous empirical tests have focused on canopy tree species or very large shrubs.Methods In order to test the AP theory further, 1586 understorey sub-tropical forest plants from 30 sites in south-east China were harvested and examined. The numerical values of the scaling exponents and normalization constants (i.e. slopes and y-intercepts, respectively) of log–log linear M_A vs. M_R relationships were determined for all individual plants, for each site, across the entire data set, and for data sorted into a total of 19 sub-sets of forest types and successional stages. Similar comparisons of M_A/M_R were also made.Key Results The data revealed that the mean M_A/M_R of understorey plants was 2·44 and 1·57 across all 1586 plants and for all communities, respectively, and M_A scaled nearly isometrically with respect to M_R, with scaling exponents of 1·01 for all individual plants and 0·99 for all communities. The scaling exponents did not differ significantly among different forest types or successional stages, but the normalization constants did, and were positively correlated with M_A/M_R and negatively correlated with scaling exponents across all 1586 plants.Conclusions The results support the AP theory’s prediction that M_A scales nearly one-to-one with M_R (i.e. M_A ∝ M_R^≈1·0) and that plant biomass partitioning for individual plants and at the community level share a strikingly similar pattern, at least for the understorey plants examined in this study. Furthermore, variation in environmental conditions appears to affect the numerical values of normalization constants, but not the scaling exponents of the M_A vs. M_R relationship. This feature of the results suggests that plant size is the primary driver of the M_A vs. M_R biomass allocation pattern for understorey plants in sub-tropical forests.     

Carbohydrate Catabolism in Populations of Bursaphelenchus xylophilus and in B. mucronatus

Genotypically different host specific pathotypes of Bursaphelenchus xylophilus have been identified. These pathotypes elicit different responses in pines depending on susceptibility, tolerance, or resistance. Continued passage of some of these pathotypes on fungal cultures leads to conversion to nonparasitic populations. These populations metabolize carbon substrates to ethanol by an anaerobic pathway, while operating some level of a phosphoenolpyruvate (PEP)-succinate pathway to excrete succinate-lactate and malate. On the other hand, parasitic populations metabolize glucose to lactate-succinate, mainly by a PEP-succinate pathway, and maintain redox balance through glycerol production. Ethanol and malate are not excreted by parasitic populations.     

A comparative analysis of leaf shape of wheat, barley and maize using an empirical shape model

Background and Aims

The phenotypes of grasses show differences depending on growth conditions and ontogenetic stage. Understanding these responses and finding suitable mathematical formalizations are an essential part of the development of plant and crop models. Usually, a marked change in architecture between juvenile and adult plants is observed, where dimension and shape of leaves are likely to change. In this paper, the plasticity of leaf shape is analysed according to growth conditions and ontogeny.

Methods

Leaf shape of Triticum aestivum, Hordeum vulgare and Zea mays cultivars grown under varying conditions was measured using digital image processing. An empirical leaf shape model was fitted to measured shape data of single leaves. Obtained values of model parameters were used to analyse the patterns in leaf shape.

Key Results

The model was able to delineate leaf shape of all studied species. The model error was small. Differences in leaf shape between juvenile and adult leaves in T. aestivum and H. vulgare were observed. Varying growth conditions impacted leaf dimensions but did not impact leaf shape of the respective species.

Conclusions

Leaf shape of the studied T. aestivum and H. vulgare cultivars was remarkably stable for a comparable ontogenetic stage (leaf rank), but differed between stages. Along with other aspects of grass architecture, leaf shape changed during the transition from juvenile to adult growth phase. Model-based analysis of leaf shape is a method to investigate these differences. Presented results can be integrated into architectural models of plant development to delineate leaf shape for different species, cultivars and environmental conditions.     

Phylogeny of the cycads based on multiple single-copy nuclear genes: congruence of concatenated parsimony,likelihood and species tree inference methods

Background and aims

Despite a recent new classification, a stable phylogeny for the cycads has been elusive, particularly regarding resolution of Bowenia, Stangeria and Dioon. In this study, five single-copy nuclear genes (SCNGs) are applied to the phylogeny of the order Cycadales. The specific aim is to evaluate several gene tree–species tree reconciliation approaches for developing an accurate phylogeny of the order, to contrast them with concatenated parsimony analysis and to resolve the erstwhile problematic phylogenetic position of these three genera.

Methods

DNA sequences of five SCNGs were obtained for 20 cycad species representing all ten genera of Cycadales. These were analysed with parsimony, maximum likelihood (ML) and three Bayesian methods of gene tree–species tree reconciliation, using Cycas as the outgroup. A calibrated date estimation was developed with Bayesian methods, and biogeographic analysis was also conducted.

Key Results

Concatenated parsimony, ML and three species tree inference methods resolve exactly the same tree topology with high support at most nodes. Dioon and Bowenia are the first and second branches of Cycadales after Cycas, respectively, followed by an encephalartoid clade (Macrozamia–Lepidozamia–Encephalartos), which is sister to a zamioid clade, of which Ceratozamia is the first branch, and in which Stangeria is sister to Microcycas and Zamia.

Conclusions

A single, well-supported phylogenetic hypothesis of the generic relationships of the Cycadales is presented. However, massive extinction events inferred from the fossil record that eliminated broader ancestral distributions within Zamiaceae compromise accurate optimization of ancestral biogeographical areas for that hypothesis. While major lineages of Cycadales are ancient, crown ages of all modern genera are no older than 12 million years, supporting a recent hypothesis of mostly Miocene radiations. This phylogeny can contribute to an accurate infrafamilial classification of Zamiaceae.     

Water stress-induced xylem hydraulic failure is a causal factor of tree mortality in beech and poplar

Background and Aims

Extreme water stress episodes induce tree mortality, but the physiological mechanisms causing tree death are still poorly understood. This study tests the hypothesis that a potted tree''s ability to survive extreme monotonic water stress is determined by the cavitation resistance of its xylem tissue.

Methods

Two species were selected with contrasting cavitation resistance (beech and poplar), and potted juvenile trees were exposed to a range of water stresses, causing up to 100 % plant death.

Key Results

The lethal dose of water stress, defined as the xylem pressure inducing 50 % mortality, differed sharply across species (1·75 and 4·5 MPa in poplar and beech, respectively). However, the relationships between tree mortality and the degree of cavitation in the stems were similar, with mortality occurring suddenly when >90 % cavitation had occurred.

Conclusions

Overall, the results suggest that cavitation resistance is a causal factor of tree mortality under extreme drought conditions.     

Scanning Electron Microscopy of Xiphinema,Longidorus, and Californidorus Stylet Morphology

Stylet ultrastructure of five Xiphinema, four Longidorus, and three Californidorus species was compared by scanning electron microscopy. Morphological differences were seen in the odontophores and odontostyle bases between the genera and some of the species. All Xiphinema studied had well-developed odontophore flanges; the Longidorus species lacked flanges, except for weakly developed ones in L. diadecturus; and none of the Californidorus had flanges. Three sinuses were present in the odontophores of all species. The sinuses varied in length depending upon species. In Xiphinema and Californidorus the odontostyle bases had distinct overlapping collars, but in Longidorus the collars were absent except for L. diadecturus. The odontostyle-odontophore junction from a lateral view appeared as a slanted transverse line in all the species, but in a dorsal view of Xiphinema and Californidorus it was V-shaped. Dorsal longitudinal seams of the odontostyle and odontophore were observed in all the species. The dorsally located odontostyle aperture was ca. 1 μm from the anterior end in all species, except in one Longidorus sp. it was ca. 4 μm from the end.     

Modelling and predicting the spatial distribution of tree root density in heterogeneous forest ecosystems

Background and Aims In mountain ecosystems, predicting root density in three dimensions (3-D) is highly challenging due to the spatial heterogeneity of forest communities. This study presents a simple and semi-mechanistic model, named ChaMRoots, that predicts root interception density (RID, number of roots m^–2). ChaMRoots hypothesizes that RID at a given point is affected by the presence of roots from surrounding trees forming a polygon shape.Methods The model comprises three sub-models for predicting: (1) the spatial heterogeneity – RID of the finest roots in the top soil layer as a function of tree basal area at breast height, and the distance between the tree and a given point; (2) the diameter spectrum – the distribution of RID as a function of root diameter up to 50 mm thick; and (3) the vertical profile – the distribution of RID as a function of soil depth. The RID data used for fitting in the model were measured in two uneven-aged mountain forest ecosystems in the French Alps. These sites differ in tree density and species composition.Key Results In general, the validation of each sub-model indicated that all sub-models of ChaMRoots had good fits. The model achieved a highly satisfactory compromise between the number of aerial input parameters and the fit to the observed data.Conclusions The semi-mechanistic ChaMRoots model focuses on the spatial distribution of root density at the tree cluster scale, in contrast to the majority of published root models, which function at the level of the individual. Based on easy-to-measure characteristics, simple forest inventory protocols and three sub-models, it achieves a good compromise between the complexity of the case study area and that of the global model structure. ChaMRoots can be easily coupled with spatially explicit individual-based forest dynamics models and thus provides a highly transferable approach for modelling 3-D root spatial distribution in complex forest ecosystems.     

Highly local environmental variability promotes intrapopulation divergence of quantitative traits: an example from tropical rain forest trees

Background and Aims

In habitat mosaics, plant populations face environmental heterogeneity over short geographical distances. Such steep environmental gradients can induce ecological divergence. Lowland rainforests of the Guiana Shield are characterized by sharp, short-distance environmental variations related to topography and soil characteristics (from waterlogged bottomlands on hydromorphic soils to well-drained terra firme on ferralitic soils). Continuous plant populations distributed along such gradients are an interesting system to study intrapopulation divergence at highly local scales. This study tested (1) whether conspecific populations growing in different habitats diverge at functional traits, and (2) whether they diverge in the same way as congeneric species having different habitat preferences.

Methods

Phenotypic differentiation was studied within continuous populations occupying different habitats for two congeneric, sympatric, and ecologically divergent tree species (Eperua falcata and E. grandiflora, Fabaceae). Over 3000 seeds collected from three habitats were germinated and grown in a common garden experiment, and 23 morphological, biomass, resource allocation and physiological traits were measured.

Key Results

In both species, seedling populations native of different habitats displayed phenotypic divergence for several traits (including seedling growth, biomass allocation, leaf chemistry, photosynthesis and carbon isotope composition). This may occur through heritable genetic variation or other maternally inherited effects. For a sub-set of traits, the intraspecific divergence associated with environmental variation coincided with interspecific divergence.

Conclusions

The results indicate that mother trees from different habitats transmit divergent trait values to their progeny, and suggest that local environmental variation selects for different trait optima even at a very local spatial scale. Traits for which differentiation within species follows the same pattern as differentiation between species indicate that the same ecological processes underlie intra- and interspecific variation.