Les forêts sacrées de Guinée : entre écologie et conservation

Guinea has very little ecological data available regarding its sacred forests. This study shows the important conservation role of these forests in a local context of strong human impacts. We present four representative case studies from the Sudano-Guinean zone in Upper Guinea. Our phytoecological inventories recorded a total of 431 species, 312 genera, and 87 families including 242 species, 187 genera, and 64 families in the forests. Twelve plant groups have been identified, including five in forests and seven in the surrounding vegetation. Geomorphology, texture, soil hydromorphism, microclimate, and human impacts are significant in explaining the spatial distribution of plant groups. The study reveals that in these Sudano-Guinean savannas, vegetation is dominated by micro-phanerophytes. The Guinean-Congolese flora is better represented in gallery forests.     

Influences of ecology and biogeography on shaping the distributions of cryptic species: three bat tales in Iberia

To determine what shapes the distributions of cryptic species, we aimed to unravel ecological niches and geographical distributions of three cryptic bat species complexes in Iberia, Plecotus auritus/begognae, Myotis mystacinus/alcathoe and Eptesicus serotinus/isabellinus (with 44, 69, 66, 27, 121 and 216 records, respectively), considering ecological interactions and biogeographical patterns. Species distribution models (SDMs) were built using a presence‐only technique (Maxent), incorporating genetically identified species records with environmental variables (climate, habitat, topography). The most relevant variables for each species’ distribution and respective response curves were then determined. SDMs for each species were overlapped to assess the contact zones within each complex. Niche analyses were performed using niche metrics and spatial principal component analyses to study niche overlap and breadth. The Plecotus complex showed a parapatric distribution, although having similar biogeographical affinities (Eurosiberian), possibly explained by competitive exclusion. The Myotis complex also showed Eurosiberian affinities, with high overlap between niches and distribution, suggesting resource partitioning between species. Finally, E. serotinus was associated with Eurosiberian areas, while E. isabellinus occurred in Mediterranean areas, suggesting possible competition in their restricted contact zone. This study highlights the relevance of considering potential ecological interactions between similarly ecological species when assessing species distributions. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 ,150–162.     

Vole mound effects and disturbance rate in a mediterranean plant community under different grazing and irrigation regimes

A factorial field experiment was used to assess the influence of soil-disturber mammals in the structure of a 9-year-old Mediterranean annual plant community subjected to different sheep grazing and irrigation regimes. We estimated the disturbance rate (mound building activity) by Mediterranean voles, their effects on vegetation and the mechanisms of these effects during a period of vole outbreak. The effects on vegetation were analysed at the levels of species, functional groups and plant community. Disturbance rate was high and voles can disturb the entire soil surface once every four or five years. The availability of certain trophic resources (perennial plants) appeared to drive vole expansion in the experimental plots and it was independent of the irrigation and grazing treatments. Mound building activities largely affected vegetation but conserved plot differences. Total vegetation cover, absolute cover of all functional groups, mean vegetation height and species richness were less on mounds than on undisturbed ground. These effects did not change the relative abundance of annuals, perennials, grasses and forbs. Only the relative abundance of small-seeded species decreased on mounds. As the proportion of these seeds was similar in both types of patches, we suggest that small-seeded species had more difficulties for germinating or emerging when they are buried during mound formation. Irrigation and sheep grazing promoted large changes in the vegetation parameters but these effects were, in general, similar on mounds and undisturbed ground. Our results show that the availability of germinable seeds may be the major limitation for mound revegetation, probably due to the scarcity of seeds existing at the depths from which soils are excavated. Our results also suggested a resource limitation on mounds. The results provide additional evidence that soil disturbances by small herbivore mammals exert relevant ecological effects on abandoned Mediterranean croplands. We discuss the ecological implications of vole mound-building activities for plant succession, plant species conservation and forage resource availability for livestock. This revised version was published online in June 2006 with corrections to the Cover Date.     

Influences of the individualistic concept of vegetation on syntaxonomy

The individualistic concept of vegetation is based on three principles: (1) vegetation continuum, (2) ecological and chorological individuality of species and (3) multidimensional variability of vegetation. All principles reflect phenomena existing in nature. The conclusions arising from these principles are these: (1) plant communities (phytocoenoses) are not natural objects (wholes) as such but merely mixtures of plant individuals coexisting on the same site as the result of migration and environmental selection, (2) the classification of vegetation is quite arbitrary if not pseudoscientific, and (3) vegetation phenomena can be reduced to phenomena of plant individuals.The integrated concept of vegetation considers the phytocoenoses as real functional systems (wholes) where plant populations are integrated both by the environment and by interactions among and within plant populations. The principles of the individualistic concept, but not the above conclusions arising from them, are compatible with the integrated concept. Phytocoenoses limit the individualistic performance of species through restriction of ecological amplitudes and shifting of ecological optima (as compared with physiological amplitudes and optima). The individualistic behaviour of species does not exclude the formation of coenological groups of species which form a basis for the classification of vegetation according to floristic-coenological criteria. Under natural conditions, plant communities are usually interconnected by transition zones to form a vegetation continuum. Phenomena of the vegetation continuum do not cause greater difficulties in syntaxonomy than those of polymorphic taxonomic groups in plant taxonomy.     

Are dominant plant species more susceptible to leaf‐mining insects? A case study at Saihanwula Nature Reserve,China

Dominant species significantly affect interspecific relationships, community structure, and ecosystem function. In the field, dominant species are often identified by their high importance values. Selective foraging on dominant species is a common phenomenon in ecology. Our hypothesis is that dominant plant groups with high importance values are more susceptible to leaf‐mining insects at the regional level. Here, we used the Saihanwula National Nature Reserve as a case study to examine the presence–absence patterns of leaf‐mining insects on different plants in a forest‐grassland ecotone in Northeast China. We identified the following patterns: (1) After phylogenetic correction, plants with high importance values are more likely to host leafminers at the species, genus, or family level. (2) Other factors including phylogenetic isolation, life form, water ecotype, and phytogeographical type of plants have different influences on the relationship between plant dominance and leafminer presence. In summary, the importance value is a valid predictor of the presence of consumers, even when we consider the effects of plant phylogeny and other plant attributes. Dominant plant groups are large and susceptible targets of leaf‐mining insects. The consistent leaf‐mining distribution pattern across different countries, vegetation types, and plant taxa can be explained by the “species‐area relationship” or the “plant apparency hypothesis.”     

Vegetation–environment relationships and ecological species groups of an Arizona Pinus ponderosa landscape, USA

Pinus ponderosa forests occupy numerous topographic and soil complexes across vast areas of the southwestern United States, yet few data exist on species distributions and vegetation–environment relationships for these environmentally diverse landscapes. We measured topography, soils, and vegetation on 66, 0.05-ha plots within a 110,000-ha P. ponderosa landscape in northern Arizona, USA, to discern vegetation–environment relationships on this landscape. We analyzed associations of environmental variables with plant communities and with single-species distributions, and we classified ecological species groups (co-occurring plant species exhibiting similar environmental affinities). Gradients in community composition paralleled gradients in soil texture, available water, organic C, total N, and geographic precipitation patterns. Soil parent material, affected by the presence or absence of volcanic activity, is a primary factor constraining vegetation patterns on this landscape. Using discriminant analysis, we built a model that correctly classified the most important of four grasses (Bouteloua gracilis, Muhlenbergia montana, Sporobolus interruptus, or Festuca arizonica) on 70–80% of plots based on five environmental variables related to soil moisture and resource levels. We also classified 52 of the 271 detected plant species into 18 ecological species groups. Species groups ranged from Phacelia and Bahia groups occupying xeric, volcanic cinder soils low in organic C and total N, to Festuca and Lathyrus groups characterizing moist, loam and silt loam soils. We applied the species groups by estimating P. ponderosa diameter increment in a regression tree using abundances of species groups. The most rapid P. ponderosa diameter growth of 5 mm/year occurred on plots with high importance of the Festuca and Lathyrus groups. Our results on this semi-arid landscape support several general ecological species group principles chiefly developed in temperate regions, and suggest that vegetation–environment research has great potential for enhancing our understanding of P. ponderosa forests occupying vast areas of the southwestern United States.     

Optical trait indicators for remote sensing of plant species composition: Predictive power and seasonal variability

Most plant species feature similar biochemical compositions and thus similar spectral signals. Still, empirical evidence suggests that the spectral discrimination of species and plant assemblages is possible. Success depends on the presence or absence of faint but detectable differences in biochemical (e.g., pigments, leaf water and dry matter content) and structural properties (e.g., leaf area, angle, and leaf structure), i.e., optical traits. A systematic analysis of the contributions and spatio-temporal variability of optical traits for the remote sensing of organismic vegetation patterns has not yet been conducted. We thus use time series of optical trait values retrieved from the reflectance signal using physical models (optical trait indicators, OTIs) to answer the following questions: How are optical traits related among patterns of floristic composition and reflectance? How variable are these relations in space and time? Are OTIs suitable predictors of plant species composition?We conducted a case study of three temperate open study sites with semi-natural vegetation. The canopy reflectance of permanent vegetation plots was measured on multiple dates over the vegetation period using a field spectrometer. We recorded the cover fractions of all plant species found in the vegetation plots and extracted gradients of species composition from these data. The physical PROSAIL leaf and canopy optical properties model was inverted with random forest regression models to retrieve time series of OTIs for each plot from the reflectance spectra. We analyzed these data sets using correlation analyses. This approach allowed us to assess the distribution of optical traits across gradients of species composition. The predictive performance of OTIs was tested in relation to canopy reflectance using random forest models.OTIs showed pronounced relationships with floristic patterns in all three study sites. These relationships were subject to considerable temporal variability. Such variability was driven by short-term vegetation dynamics introduced by local resource stress. In 72% of all cases OTIs out-performed the original canopy reflectance spectra as indicators of plant species composition. OTIs are also easier to interpret in an ecological sense than spectral bands or features. We thus conclude that optical traits retrieved from reflectance data have a high indicative value for ecological research and applications.     

Monitoring shifts in plant diversity in response to climate change: a method for landscapes

Improved sampling designs are needed to detect, monitor, and predict plant migrations and plant diversity changes caused by climate change and other human activities. We propose a methodology based on multi-scale vegetation plots established across forest ecotones which provide baseline data on patterns of plant diversity, invasions of exotic plant species, and plant migrations at landscape scales in Rocky Mountain National Park, Colorado, USA. We established forty two 1000-m² plots in relatively homogeneous forest types and the ecotones between them on 14 vegetation transects. We found that 64% of the variance in understory species distributions at landscape scales were described generally by gradients of elevation and under-canopy solar radiation. Superimposed on broad-scale climatic gradients are small-scale gradients characterized by patches of light, pockets of fertile soil, and zones of high soil moisture. Eighteen of the 42 plots contained at least one exotic species; monitoring exotic plant invasions provides a means to assess changes in native plant diversity and plant migrations. Plant species showed weak affinities to overstory vegetation types, with 43% of the plant species found in three or more vegetation types. Replicate transects along several environmental gradients may provide the means to monitor plant diversity and species migrations at landscape scales because: (1) ecotones may play crucial roles in expanding the geophysiological ranges of many plant species; (2) low affinities of understory species to overstory forest types may predispose vegetation types to be resilient to rapid environmental change; and (3) ecotones may help buffer plant species from extirpation and extinction.     

Patterns of plant diversity at high altitudes on the Qinghai-Tibetan Plateau

Aims To describe the biodiversity patterns of plants along an altitudinal gradient on the Qinghai-Tibetan Plateau and to clarify the bias in plant specimen records at high altitude.Methods We conducted a large-scale investigation of vegetation at a wide range of altitudes, focusing on a high-altitudinal range (3?200–5?200 m) at different locations on the Qinghai-Tibetan Plateau. We then compared the altitudinal distribution of plant species obtained from our field investigation with that in plant specimen records from published sources and an online database.Important findings Our data provide evidence that altitude plays a large role in regulating species composition on the Qinghai-Tibetan Plateau. We could not, however, detect a clear relationship between altitude and species richness, although a weak monotonically increasing trend of richness was detected with increasing altitude. According to specimen records, most species have been sampled at a wide range of altitudes, and the average range of 145 species is>2?000 m. Despite this wide range, more than half of the species we observed were at higher altitudes than the specimen records indicate. High-altitude areas have probably been so poorly sampled that only a small fraction of the resident species has been recorded. This study clearly shows the regional bias of specimen records in the Qinghai-Tibetan Plateau.     

浙江瑞安公益林群落结构及其与环境的相关性

运用CCA排序、TWINSPAN等级分类、物种多样性以及种间联结等分析方法探讨了浙江省瑞安市公益林群落中地形因子对植物空间分布格局的影响以及主要优势物种的种间关系,以期解释公益林群落特征及其与环境因子的关系。结果表明,瑞安市公益林物种分布受海拔、坡度和坡向(总体上可解释69.1%的环境因子)的影响。运用TWINSPAN等级分类,在种-环境CCA排序结果的基础上将群落大致划分为13类群丛,其中以马尾松(Pinus massoniana)、杉木(Cunninghamia lanceolata)和柳杉(Cryptomeria fortunei)为优势种的群丛个数最多;群落的多样性水平总体较高,以马尾松、杉木和柳杉为优势种的群丛因其具有较好的地形条件而多样性最高;群落垂直层次的物种多样性由大到小依次为:灌木层乔木层草本层,同时3个垂直层次中主要优势种的正负关联比低(均小于2),其中乔木层优势种的正负关联比最高,说明研究区群落结构尚不稳定,但乔木层相对比较稳定。研究结果为研究区公益林管理提供理论支撑。     

Bryophyte diversity and distribution along an altitudinal gradient on a lava flow in La Réunion

Non‐vascular plant distribution patterns were examined in three microhabitats along an altitudinal gradient on a recent lava flow of the Piton de la Fournaise volcano (La Réunion, Mascarene archipelago). The uniform nature of the lava flow provides an excellent system to study the relationship between altitude and species diversity and distribution, and at the same time avoiding confusing multiple effects of substrate and vegetation heterogeneity. Non‐vascular plants were surveyed with quadrats within an altitudinal range from 250 m to 850 m a.s.l. Fine‐scale variations in bryophyte communities between three ecological microhabitats (the ground and on the rachises of two fern species) were investigated. Three specific questions were addressed: (1) What is the species diversity of bryophyte communities on a 19‐year‐old lava flow? (2) How does altitude influence the diversity and distribution of bryophytes on a lava flow? (3) Does microhabitat variation control bryophyte diversity? In our study, bryophyte diversity increased with altitude. Unexpectedly, species richness was very high; 70 species of bryophytes were recorded including nine new records for the island. Diversity was also controlled by ecological microhabitats. Bryophyte species were structured into six categories according to altitude and microhabitat preferences. Results suggested that the high diversity of these cryptic organisms on this lava flow is fostered in part by their host substrate and their adaptative strategies on new substrates. On a broader scale, it was concluded that lava flows as primary mineral environments are important to conserve, as they support a high diversity of pioneer organisms that constitute the early stages of the development of La Réunion's remnant lowland rainforest.     

水坝工程对澜沧江中游陆生植物的生态风险评估

以澜沧江中游小湾水库库周为研究区,在1997年(小湾水库建设前)澜沧江中游小湾水库下游3个样地[输电区(1号样地)、变电站和进站公路区(2号样地)和水库移民区(3号样地)]植被调查基础上,2010年(小湾水库建设后)分别对3个样地进行植被组成、盖度和优势种观测,评估了水坝工程对澜沧江中游陆地植物的生态风险.结果表明:3个样地上优势种的总和优势度在水坝建设前后均发生了明显变化.本研究根据物种类型(优势种和非优势种)及物种总和优势度变率将生态风险划分为无风险/极低风险(0)、低风险(Ⅰ)、中风险(Ⅱ)、高风险(Ⅲ)、极高风险(Ⅳ)5个等级.研究区处于Ⅲ级生态风险(高风险)的物种数量最多,部分物种处于Ⅳ级生态风险(极高风险);3号样地处于Ⅲ级(高风险)和Ⅳ级(极高风险)生态风险的物种比例明显高于l、2号样地.水坝建设导致的物种水平生态风险主要表现为原生物种减少、外来物种大量入侵等.应采取有效的保护措施降低陆生植物物种水平水坝生态风险.     

黄土丘陵区延河流域潜在植被分布预测与制图

潜在植被的分布预测与制图对植被恢复规划具有重要的指导价值.利用广义相加模型（generalized additive model,GAM）,结合GIS空间分析技术和环境梯度分层采样技术,为延河流域24个地带性物种建立了分布模型,并在考虑群落内部物种种间关系及其分布概率的基础上,对物种分布进行运算,模拟预测了延河流域37种植物群落的分布状况和延河流域的潜在植被分布.结果表明: 研究区植被分布预测值与实际调查值间的差异不显著,预测的植被空间分布较好地反映了延河流域潜在的植被分布状况,表明该模型具有较好的预测能力,对于区域植被恢复的目标设定和恢复规划具有重要意义.     

黄土高原林草植被建设的地带性特征

 针对黄土高原现有人工林草植被存在的主要问题,依据天然植被地带性分布规律和实测资料,分析了以水分为主的生态条件及林草植被对水分利用的地带性特征,提出了黄土高原林草植被建设的群落学原理和关键技术。即：依据植被地带分布规律指导人工林草植被建设;选择地带性植被优势种作为主要造林种草的植物种;模拟天然植被结构实行乔灌草复层混交是快速建造稳定植被的科学途径。根据一些“适地适树”树种并不能正常成林的现象,提出造林应强调“适地适林”的原则,同时给出适宜黄土高原不同植被地带林草植被建设的主要树草种和伴生种或四旁绿化树种。     

A palaeoecological model for the interpretation of wild plant species

The interpretation of subfossil records of wild plant species with respect to both environmental conditions and past vegetation is complicated by the following: (1) production and dispersal of plant remains including diaspores, (2) the formation of the soil flora, (3) taphonomic processes and differential preservation that act on subfossil assemblages and (4) methods applied to produce subfossil records. Whereas the similarity between recent plant communities and seed banks is often weak, the relationship between past vegetation and subfossil assemblages is still more complicated. It is therefore unlikely that macrofossil assemblages derived from soil samples can be considered as pure samples representing particular palaeobiocoenoses. The assumption that plant communities, in the past, may have been in some way aberrant with respect to composition and that the ecological ranges of species varied during the Quaternary has to be rejected, if not based on well considered assumptions or evidence from pure samples. Only if a sufficient number of suitable studies is available, which enable evaluation between all kinds of plant communities and their respective seed floras, can progress be made with regard to the reconstruction of past vegetation and environmental conditions. As long as these data are not available, the ecological interpretation of particular subfossil assemblages isolated from soil samples has to be carefully evaluated within their particular context.     

Distribution of alien vs. native plant species in roadside communities along an altitudinal gradient in Tenerife and Gran Canaria (Canary Islands)

Roadside plant communities were studied along two roads following an altitudinal gradient in Gran Canaria and Tenerife (Canary Islands). Our aim was to investigate variation in plant species richness, particularly of the alien flora, along a gradient from coastal shrubland to summit vegetation (1950 m a.s.l. in Gran Canaria, 2300 m in Tenerife) in relation to variation in habitat factors (altitude, habitat structure, roadside disturbance, distance to urban nuclei). We compared different species groups that were classified in terms of their biogeographical status, origin and life form. Altitude was the most important factor determining species richness and composition along both roadside transects. Alien plants showed a unimodal distribution pattern along the altitudinal gradient, with less species and lower abundance at low and high altitudes, and highest abundance at intermediate altitude. Alien plant species were also relatively more frequent near urban centres. The number of native and alien species was significantly positively correlated along the altitudinal gradient. Both alien and native, non-endemic species showed differences in their distribution along the altitudinal gradient according to their biogeographical affinities and climatic tolerances. Despite considerable differences in species pools these patterns were consistent among the two islands. Environmental (abiotic) stress is proposed as a primary, altitude-related factor acting as a filter against most alien plants at coastal and high-mountain altitudes. A higher frequency or intensity of disturbance at intermediate altitudes may be a further causal factor promoting alien plants in this zone. Future management efforts to control alien plants along roads should, therefore, concentrate on intermediate altitudinal zones of the higher Canary Islands.     

The tick Ixodes ricinus: distribution and climate preferences in the western Palaearctic

In this study, multivariate spatial clustering on monthly normalized difference vegetation index (NDVI) maps is used to classify ecological regions over the western Palaearctic. This classification is then used to delineate the distribution and climate preferences of populations (clades) of the tick Ixodes ricinus L. (Acari: Ixodidae) from a geographically extensive dataset of tick records and a gridded 2.5-km resolution climate dataset. Using monthly layers of the NDVI, regions of similar ecological attributes were defined and nine populations with significant differences in critical climate parameters (P< 0.005) were detected. Grouping of tick records according to other categories, such as political divisions, a 4 degrees x 4 degrees grid overlying the study area, or the CORINE) and USGS) vegetation classification schemes did not provided significantly separated populations (P = 0.094-0.304). Factor analysis and hierarchical tree clustering provided an ecological overview of these tick clades: two Mediterranean and one Scandinavian (western) clades are clearly separated from a node that includes clades of different parts of central Europe and the British Isles, with contrasting affinities between the different clades. The capture records of these ecologically separated clades produce a clear bias when bioclimate envelope modelling is applied to the mapping of habitat suitability for the tick in the western Palaearctic. The best-performing methods (Cohen's kappa = 0.834-0.912) use partial models developed with data from each ecoregion, which are then overlapped over the region of study. It is concluded that the use of ecologically derived ecoregions is an objective step in assessing the presence of ecologically different clades, and provides a guide in the development of data partitioning for habitat suitability modelling.     

Novelty predictors for shrub (and climbers) ecological niche modeling,based on their successional stage

The identification of species' environmental predictors constitute a key challenge for decision making, especially when using ecological niche modeling based on these drivers and when presence points are limited. More specifically, shrub species are affected by ecosystem dynamics, and appear in degraded formations, in dense mid-stage vegetation formations, or under late climax-forest canopy. In this study, we tested novelty predictors to understand the drivers that affect the selected species distribution in the Mediterranean biome, targeting different vegetation successional stages, and further improve ecological models' performance, when presence points are limited. Land surface temperature (LST) in association with temperature related predictors, allowed differentiating between species thriving in the understory of the forest canopy, from those that are co-dominant with dense vegetation cover and a third group/species, thriving in degraded vegetation. In addition, the Normalized Difference Vegetation Level Index (NDVI) played a key role in the models for species growing in highly degraded ecological niches such as Spartium junceum, Calicotome villosa, but also forest-fringe vegetation like the climber Hedera helix. Our study highlights the importance of integrating remote sensed predictors, combined with appropriate climate drivers, when using ecological niche modeling.     

中国西南喀斯特植物群落演替特征及驱动机制

植被恢复是西南喀斯特石漠化治理和生态重建的关键步骤。通过总结已有的结果和分析中国科学院环江喀斯特试验站部分试验数据,研究了西南喀斯特植被演替过程中群落的动态特征,结果发现:西南喀斯特沿北、中、南亚热带至北热带的海拔递减、气温和降雨量升高的梯度格局下,群落的高度、盖度增加,密度、科属种数量和物种多样性呈递减趋势;不同区域沿草丛、灌丛、灌乔丛、乔丛的顺向演替方向上,群落的高度、盖度增加、密度减小,科属种数量及物种多样性除北亚热带持续增加外,其他3个区域的最大值均出现在灌乔丛,乔丛反而有所下降;西南喀斯特植被演替是生物驱动因素与环境阻力相互作用的结果,不同区域不同演替阶段植被演替的建群种和关键种不同,其植物群落的稳定性受多样性和结构性的共同控制,因地质和生态环境恶劣,植被退化容易恢复难,充分理解和认识植被演替规律对西南喀斯特植被恢复和生态重建意义重大。     

Ecological amplitude in Silene nutans in relation to allozyme variation at the western margin of its distribution

Abstract. At the western border of its geographical range, Silene nutans (Caryophyllaceae) has evolved two groups of parapatric populations showing distinct allozyme patterns and apparently occurring on different bedrock types. This study tests the hypothesis that these groups represent edaphic ecotypes. With this in view, the ecological amplitude of 36 populations of Silene nutans from Belgium was investigated and their synecology specified using vegetation composition and soil parameters; Ellenberg indicator values were used for ecological interpretation of the vegetation analyses. The results provide evidence that allozyme and habitat variations are correlated, the two groups occurring on contrasting soil with distinct vegetation types. One group is restricted to alkaline soils and typically occurs in open calcareous grasslands, fringes and scrub. The other group is characterized by wider autecological and synecological ranges with a bimodal pH- distribution, occurring on two kinds of siliceous soils: (1) neutral soils, supporting vegetation with a high species richness comprising many mesophilous species, and (2) CaCO₃ -free bedrocks characterized by acidophilous, species-poor, open dry grasslands and woodlands. It also appears that factors not related to soil chemistry, such as microclimate, are involved in the ecological specialization of the two ecotypes. It is argued that these ecotypes exemplify adaptive radiation and parapatric speciation at the margin of a species distribution area.