Ecological wood anatomy of NepaleseRhododendron (Ericaceae). 2. Intraspecific variation

The intraspecific relationship of selected wood anatomical characters with stem diameter, plant height, and altitude was investigated in four NepaleseRhododendron species:R. anthopogon, R. lepidotum, R. campanulatum, andR. arboreum, i.e., two shrubs, one subtree, and one tree. We studied 23 to 27 specimens for each species.R. anthopogon grew from 3,380 to 4,950 m,R. lepidotum from 2,060 to 4,720 m,R. campanulatum from 2,790 to 4,140 m, andR. arboreum from 1,430 to 3,460 m. Multiple regression analysis and actual distribution of character values show that the wood anatomical characters having a significant correlation with non-anatomical factors differ between species. The number of significant characters are two inR. anthopogon andR. campanulatum, five inR. lepidotum, and nine inR. arboreum and tends to be small in species having smaller altitudinal ranges. Average pore area, most strongly correlated with non-anatomical factors in interspecific variation, is significantly correlated only inR. lepidotum andR. arboreum. The general trends in intraspecific variation among four NepaleseRhododendron species differ from the trends found in interspecific variation within the genus and are characteristic of each species.     

Ontogenetic wood anatomy of tree and subtree species of Nepalese Rhododendron (Ericaceae) and characterization of shrub species

Ontogenetic trends in the wood structure of Nepalese Rhododendron were studied in 15 specimens of two tree and four subtree species. Average growth ring width was constant from pith to bark in spite of occurrences of extremely narrow, false, or discontinuous rings. Vessel density, vessel area, vessel element length, and multiseriate ray height generally had an initial increase or decrease to 1.5 cm radius and near plateau or slight decrease or increase outward. Multiseriate ray density and area percentage were variable between specimens without a clear pattern. Ontogenetic trends from pith to fully mature wood in trees plus subtrees were inferred by treating the measurements in the present study with those of mature individuals in a previous study. Comparison of trends in trees plus subtrees and those in shrubs lead to ecological or systematic groupings. Vessel features showed that alpine shrub species have distinctly small, numerous vessels composed of short vessel elements. Multiseriate ray features indicated a systematic difference between the trees plus subtrees of subgenus Hymenanthes and the shrubs of subgenus Rhododendron. Vessel features of alpine shrubs may be an adaptation against frequent freeze-thaw cycles or the result of growth stress imposed by the severe alpine environment.     

Wood anatomy of Gentianaceae, tribe Helieae, in relation to ecology, habit, systematics, and sample diameter

Twenty collections representing one species each ofSymbolanthus andTachia, and 17 species ofMacrocarpaea were studied by means of light microscopy and scanning electron microscopy (SEM). Wood details show that the three genera form a coherent group;Tachia differs from the others in only a few minor characters. Because the species studied form a natural group, wood variations within Helieae offer the basis for correlations and interpretations with respect to habit and ecology. Diameter of stems studied proves to be an important variable that must be taken into account. Correlations with stem diameter include wider vessels in outer wood of wider samples. This would correspond to deeper penetration of reliable water tables by roots of helioid trees or large shrubs. Ray height decreases with increase in stem diameter, an indication of paedomorphosis. Rays of all species are paedomorphic in histology by virtue of relative paucity or even absence of procumbent cells in multiseriate rays. Pseusoscalariform lateral wall pitting of vessels is also a feature characteristic of paedomorphosis. The assemblage of paedomorphic features correlates well with the conclusion, reached by authors who used cladistic methods, that Gentianaceae other than Gentianeae are derived from suffrutescent prennials. The Mesomorphy Ratio, which incorporates three vessel features, correlates with leaf length and with stem diameter. All Helieae are mesophytic, but to various degrees. Septate fiber-tracheids, where present, are typically near vessels and form a substitute for or an addendum to vasicentric axial parenchyma as a mechanism for photosynthate storage. Vestured pits occur on lateral wall pits of vessels of all Helieae, but not on the fibertracheids. Vestured pits show diversity withinMacrocarpaea, a feature of possible systematic significance.     

Ecological wood anatomy ofAlnus nepalensis (Betulaceae) in East Nepal

Wood anatomical characters ofAlnus nepalensis growing in East Nepal are evaluated against three non-anatomical factors: tree height, diameter at breast height (DBH), and altitude. Samples were taken from the outermost part of the trunk of five canopy trees at 11 localities between 790 and 2,740 m above sea level. Tree height ranged from 10 to 28 m, and DBH ranged from 15 to 80 cm. Altitude and tree height are correlated with all the vessel characters studied. Among wood anatomical characters, vessel characters measured from cross sections are strongly correlated with one another, and are also correlated with vessel element and fiber-tracheid length. Multiple regression analysis using non-anatomical factors as independent variables resulted in significant correlation at 1% level in all pore characters, vessel element length, perforation plate bar number, and fibertracheid length. Regression coefficients of significant regressions are usually largest for altitude. For wood structure ofAlnus nepalensis in East Nepal, 23 to 42% of the variation is affected by non-anatomical factors. The large contribution of altitude is considered to be an indirect measure of the effect of temperature.     

A search for phylogenetically informative wood characters within Lecythidaceae s.l

The wood structure of 71 species representing 24 genera of the pantropical Lecythidaceae s.l., including the edible Brazil nuts (Bertholletia excelsa) and the spectacular cannon-ball tree (Couroupita guianensis), was investigated using light and scanning electron microscopy. This study focused on finding phylogenetically informative characters to help elucidate any obscure evolutionary patterns within the family. The earliest diverging subfamily Napoleonaeoideae has mixed simple/scalariform vessel perforations, scalariform vessel-ray pitting, and high multiseriate rays, all features that are also present in Scytopetaloideae. The wood structure of Napoleonaea is distinct, but its supposed close relative Crateranthus strongly resembles Scytopetaloideae. The isolated position of Foetidia (Foetidioideae) can be supported by a unique type of vessel-ray pitting that is similar in shape and size to intervessel pitting (distinctly bordered, <5 μm). The more derived Planchonioideae and Lecythidoideae share exclusively simple perforations and two types of vessel-ray pitting, but they can easily be distinguished from each other by the size of intervessel pitting, shape of body ray cells in multiseriate rays, and the type of crystalliferous axial parenchyma cells. The anatomical diversity observed is clearly correlated with differences in plant size (shrubs vs. tall trees): the percentage of scalariform perforations, as well as vessel density, and the length of vessel elements, fibers, and multiseriate rays are negatively correlated with increasing plant size, while the reverse is true for vessel diameter.     

The Ecological Secondary Xylem Anatomy of 7 Desert Species of Leguminosae

The wood anatomy of 7 species (Ammopiptanthus mongolicus, Amorpha fruticosa, Halimodendron halodendron, Hedysarum mongolicum. Hedysarum scoparium, Lespedeza bicolor and Robinia pseudoacacia)of Leguminosae, which grow in desert regions of Northern China, is described in details. A comparative study on the quantitative wood anatomical characters among the species is made. Except some anatomical characters in A. fruticosa were larger in vessel diameter, thin walled in vessels and libriform fibres, all the rest six species showed a general similarities:vessel frequency/sq, mm very numerous and percentage of multiple vessels high; vessel elements very short, perforations simple and in almost horizontal end walls, intervessel bordered pits alternate and vestured; libriform fibres very short, and usually with thickened walls, and with simple pits; average ray height very low, and with multiseriate as well as uniseriate. However, there are differences in other characters, e. g. vessel distribution, percentage of solitary vessels; spiral thickenings present or absent; amount of axial parenchyma and distribution; ray frequency and type; crystals present or absent, and crystal distribution, if present. According to these anatomical diversities, a key to the identification of the 7 species is given. In this article, the relation between the structure of wood and the environmental influences has been discussed.     

Anatomical wood variation of Buddleja cordata (Buddlejaceae) along its natural range in Mexico

Buddleja cordata is an evergreen species of wide distribution in Mexico that is represented by shrubs and trees. Wood variability of B. cordata was evaluated in relation to plant size as well as latitude, altitude, soils, and climatic data. Canonical correlation analysis (CCA) showed that two canonical correlations are significant (Wilks' λ, p<0.0001) and explained 76% of total variance. Redundancy analysis revealed that the first pair of canonical variates are significant, thus the canonical variate, named distribution, represents a gradient of maximum temperature of the warmest period, annual temperature range, and latitude in its area of distribution; whereas the canonical variate named wood represents vessel density, fiber length, and plant size, best associated to the environmental gradient. Vessel density expressed by its distribution in latewood and porosity type showed that ring-porosity is common in individuals from high latitudes. Temperatures below zero or lack of rainfall during several months might induce porosity variability in B. cordata as suggested by CCA, but was not related to phenology since the species is evergreen along its latitudinal range. Plant size was also influenced by extreme temperature and rainfall. Shorter plants are distributed in the northern population or driest sites located in north-central Mexico, and in addition, fiber length followed an allometric relation with individuals' height. Wood characters in B. cordata as for simple perforation plate, helical thickenings, type of intervascular and vessel-ray pits, scanty paratracheal parenchyma, and heterogeneous type IIB rays were not correlated with plant size, climate, and soil parameters or species distribution. These features are common with other species of Buddleja.An erratum to this article can be found at     

Comparative wood anatomy ofCoriaria

Wood anatomy ofCoriaria was surveyed to clarify generic features on the basis of 14 species collected from various regions of the World to cover the whole range of geographic distribution and habitual variation. Wood anatomy ofcoriaria is considerably uniform, and the species share a combination of the following features: 1) pores are thin-walled, polygonal in outline and mostly in multiples; 2) vessel elements and libriform fibers are very short; 3) perforation plates are exclusively simple; 4) intervessel pits are alternate; 5) vascular tracheids are present; 6) wood parenchyma is vasicentric and sometimes confluent; 7) rays are heterogeneous and large. Its species differ in several characters, such as distinctness of growth rings, pore size, pore patterns, type and abundance of wood parenchyma, and distinctness of storied structure. Comparisons among species indicate that the species of the Northern Hemisphere show a tendency toward having semi-ring porosity, while those of the Southern and Western Hemisphere have diffuse porosity. The other infrageneric variations appear to be related to different habits of the species rather than to geographic distribution. Small trees mostly have confluent and vasicentric parenchyma composed of fusiform cells and distinctly storied tissues, while shrubs and herbs have less abundant parenchyma which is vasicentric and comprises strands of two to four cells and indistinctly storied tissues.     

Geographic variation in seed traits within and among forty‐two species of Rhododendron (Ericaceae) on the Tibetan plateau: relationships with altitude,habitat, plant height,and phylogeny

Seed mass and morphology are plant life history traits that influence seed dispersal ability, seeding establishment success, and population distribution pattern. Southeastern Tibet is a diversity center for Rhododendron species, which are distributed from a few hundred meters to 5500 m above sea level. We examined intra‐ and interspecific variation in seed mass and morphology in relation to altitude, habitat, plant height, and phylogeny. Seed mass decreased significantly with the increasing altitude and increased significantly with increasing plant height among populations of the same species. Seed mass differed significantly among species and subsections, but not among sections and subgenera. Seed length, width, surface area, and wing length were significantly negative correlated with altitude and significantly positive correlated with plant height. Further, these traits differed significantly among habitats and varied among species and subsection, but not among sections and subgenera. Species at low elevation had larger seeds with larger wings, and seeds became smaller and the wings of seeds tended to be smaller with the increasing altitude. Morphology of the seed varied from flat round to long cylindrical with increasing altitude. We suggest that seed mass and morphology have evolved as a result of both long‐term adaptation and constraints of the taxonomic group over their long evolutionary history.     

Dendrometric characteristics as indicators of pressure of Afzelia africana Sm. dynamic changes in trees found in different climatic zones of Benin

Afzelia africana is a forest species used by local inhabitants for various purposes, especially as forage to feed cattle, as medicinal plant and its wood is used to make furniture or for cooking. Its utilisation in its current form constitutes a threat to this species. However, the lack of data on this species is a hindrance towards drawing up an efficient program for its sustainable management. In order to fill in some gaps in the knowledge of A. africana tree populations, dendrometric characteristics of this species were studied within different climatic zones where it occurs in Benin. Data collected on each of them included height and diameter, and with regard to the levels of pressure, five categories were defined namely: null, weak, moderate, severe and very severe. As far as diameter is concerned every size was taken into consideration in all the climatic zones. However, average diameter and height of the A. africana individuals varied significantly according to climatic zones. Anthropogenic pressure increased while moving from the humid zone towards the drier zone. Moreover, there was a noticeable significant change in the level and quality of pressure between trees found in the different climatic zones in the sense that the lower the height of the trees, the more severe the level of pressure exerted. Such a relationship was not significant when one considers tree diameter in accordance with the climatic zones in the country.     

红锥家系遗传变异与优良家系选择

为选择红锥(Castanopsis hystrix)优良家系,对2–11年生红锥家系的生长性状和遗传参数进行了研究分析。结果表明,家系间在树高、胸径和单株材积上的差异达显著或极显著水平,家系的树高、胸径和单株材积受中等或中等以上遗传控制,具有很强的家系和家系内单株选择潜力;家系的单株材积变异幅度较大,胸径居中,树高的较小,显示出红锥家系间存在着丰富的变异;家系和单株遗传力随林龄的增大逐渐变小,到7–9年生时逐渐稳定。年度生长性状的相关分析表明,树高、胸径和单株材积的遗传相关系数随林龄逐渐增大,到7年生时最大,分别达到0.9602、0.9340和0.9849,之后趋于稳定,因此,7年生可作为早期选择的适合年龄。结合早期选择及形质指标,11年生时最终选择出优良家系10个,其平均树高、胸径和单株材积分别为13.95 m、14.34 cm和0.1229 m3,平均遗传增益分别为12.24%、18.69%和51.59%,其通直度、圆满度、分枝角度、最大分枝、冠幅、枝下高等形质指标也提高7.71%~12.94%。     

Gen-ecology of capsella bursa-pastoris from an altitudinal transsect in the Alps

Summary Seeds were collected in wild populations of Capsella bursa-pastoris growing along a cline from low elevations to the high mountain region in Switzerland. Progeny were grown in open field random block experiments and a number of relevant characters was measured. Germination behaviour showed no relationsship to the place of origin and exhibits considerable phenotypic plasticity. Flowering, plant height, rosette diameter and leaf forms displayed genotypic variations that were correlated with altitude. Along the considered altitude gradient phenotypic plasticity was overruled by genetic variation. Later flowering genotypes replaced the earlier flowering genotypes along the topocline which indicates retarded maturity and prolongation of the life cycle at high altitude.     

Testing the generality of above‐ground biomass allometry across plant functional types at the continent scale

Accurate ground‐based estimation of the carbon stored in terrestrial ecosystems is critical to quantifying the global carbon budget. Allometric models provide cost‐effective methods for biomass prediction. But do such models vary with ecoregion or plant functional type? We compiled 15 054 measurements of individual tree or shrub biomass from across Australia to examine the generality of allometric models for above‐ground biomass prediction. This provided a robust case study because Australia includes ecoregions ranging from arid shrublands to tropical rainforests, and has a rich history of biomass research, particularly in planted forests. Regardless of ecoregion, for five broad categories of plant functional type (shrubs; multistemmed trees; trees of the genus Eucalyptus and closely related genera; other trees of high wood density; and other trees of low wood density), relationships between biomass and stem diameter were generic. Simple power‐law models explained 84–95% of the variation in biomass, with little improvement in model performance when other plant variables (height, bole wood density), or site characteristics (climate, age, management) were included. Predictions of stand‐based biomass from allometric models of varying levels of generalization (species‐specific, plant functional type) were validated using whole‐plot harvest data from 17 contrasting stands (range: 9–356 Mg ha^?1). Losses in efficiency of prediction were <1% if generalized models were used in place of species‐specific models. Furthermore, application of generalized multispecies models did not introduce significant bias in biomass prediction in 92% of the 53 species tested. Further, overall efficiency of stand‐level biomass prediction was 99%, with a mean absolute prediction error of only 13%. Hence, for cost‐effective prediction of biomass across a wide range of stands, we recommend use of generic allometric models based on plant functional types. Development of new species‐specific models is only warranted when gains in accuracy of stand‐based predictions are relatively high (e.g. high‐value monocultures).     

Aboveground biomass allocation and water content relationships in Mediterranean trees and shrubs in two climatological regions in Israel

Thisstudy investigated the variation along basipetal gradients of the relationshipsbetween the foliage/wood allocation ratios of biomass and of water content, inMediterranean trees and shrubs, at two different locations along a climaticgradient. Understanding of the biomass allocation and water relations inMediterranean trees and shrubs provides useful information on growth patternsofthese species, and on resource dynamics of these plant communities. Twoexperimental sites were selected along a climatological transect that runs fromthe foothills of the Judean Hills to the northern Negev desert in Israel. Ateach site, 16 quadrats of 10 × 10 m (eight on south-facingslopes and eight on north-facing slopes) were marked. The aboveground biomassofdominant tree and shrub species were estimated. Main branches of trees andshrubs were cut, their foliage and wood biomass were separately weighed, andtheir respective water contents were determined. The species studied includedthe evergreen sclerophylls, Quercus calliprinos, Phillyrealatifolia and Pistacia lentiscus, and thesemi-deciduous species, Cistus creticus, Coridothymuscapitatus and Sarcopoterium spinosum. Theresults indicated that the foliage/wood ratio decreased from the periphery ofthe crown to the interior of the trees and shrubs: foliage biomass and waterwere mainly limited to the top 30 cm of the crown in all studiedspecies. Leaves had higher relative water contents than woody tissues in theupper part of the crown. However; when the whole tree or shrub was considered,the relative water content was found to be mostly allocated to the woodystructures. The results are discussed in terms of biomass allocation in variouslife forms of the eastern-Mediterranean plant communities and how they areaffected by slope aspect and climatic conditions.     

What causes geographical variation in the species–area relationships? A test from forests in China

The increase of species richness with sampling area and the decrease with latitude and altitude are two of the most frequently studied patterns in biogeography. However, few studies have simultaneously examined these two patterns to investigate how species–area relationships (SAR) vary with latitude and altitude. In this study, we explore the spatial patterns of SAR in forests in China by investigating numbers of species by life form group (trees, shrubs and herbs) in 32 nested plots from 12 mountains ranging from 18.7°N to 51.9°N in latitude and from 300 to 3150 m in altitude. The slopes of the power law SAR (z‐values) decreased with increasing latitude for all life forms except herbaceous plants, and also decreased with increasing altitude for all life forms but not for shrubs. Latitude and altitude, as well as their interactions, together explained 65.4, 61.8, 48.9 and 45.3% of the variation in z‐values for overall species, trees, shrubs and herbaceous plants, respectively. In addition, actual evapotranspiration affected SAR significantly, but this effect varied significantly among life forms. We concluded that there are significant geographical patterns of SAR for China's forests, which is primarily controlled by energy availability.     

桥山栎林群落结构特征与物种多样性相关关系分析

为了解森林群落结构特征与物种多样性之间的相关关系,以黄土高原桥山林区典型麻栎纯林、麻栎阔叶混交林和麻栎油松混交林为研究对象,调查分析了群落结构特征,计算物种重要值及物种多样性,并进行了冗余分析,结果表明:(1)麻栎油松混交林主要以乔木层胸径、树高、枝下高等最高;麻栎阔叶混交林以灌木层盖度、地径、冠幅、高度等最高;麻栎纯林以草本地径、草本盖度及草本冠幅最高。(2) 3种类型林分乔木层重要值最高的均为麻栎(Quercus acutissima),灌木层为狼牙刺(Sophora viciifolia)、南蛇藤(Celastrus orbiculatus)、胡枝子(Lespedeza bicolor),草本层为苔草(Carex tristachya)。(3)麻栎油松混交林乔、灌层物种多样性较高,麻栎阔叶混交林以草本层物种多样性最高。(4)不同类型麻栎林群落结构特征与物种多样性关系有差异。松栎混交林中,对物种多样性影响最大的为乔木胸径、新稍长及灌木高度;麻栎阔叶混交林对物种多样性影响最大的为灌木层高度及冠幅;麻栎纯林中,对物种多样性影响最大的为乔木胸径。(5)麻栎油松混交林、麻栎阔叶混交林的胸径、树高、物种多样较麻栎纯林高,具有较高木材生产能力和生态防护功能,是未来森林经营培育的方向。     

岷江源区两种优势针叶树当年生小枝性状与生物量分配随海拔的分异规律

当年生小枝具有较少的次生组织,同时是植物分支系统中最具活力的部分,木本植物当年生小枝性状与生物量分配的海拔变化是理解物种对不同生境适应策略的重要内容。通过分析青藏高原东缘岷江源区两种优势亚高山针叶乔木(紫果云杉和岷江冷杉)当年生小枝性状(茎长、茎粗、比茎长)与不同部位器官(茎、叶)的生物量随天然生境的海拔(3500-3550 m、3650-3700 m和3800-3850 m)变化,尝试揭示两物种当年生小枝在不同海拔下的生物量权衡及其生长策略。结果表明:(1)不同海拔下茎生物量种间差异大于种内差异,3650-3700 m处的茎生物量变异最大(128.4%)。(2)云杉的茎生物量、总叶生物量与海拔间存在显著的负相关关系(P<0.05),比茎长与海拔呈显著正相关(P<0.05);冷杉的总叶生物量与海拔间为显著负相关关系(P<0.05)。(3)随着海拔的升高,云杉茎生物量分配比逐渐从33.0%降低到27.0%;而冷杉的茎生物量分配比则从23.0%渐增至28.0%。(4)3500-3550 m和3650-3700 m两处的云杉茎生物量与总叶生物量、茎长与茎粗呈异速生长关系;3500-3550 m、3650-3700 m和3800-3850 m三处的冷杉茎生物量与总叶生物量、茎长与茎粗一直呈现异速生长关系。两种针叶树的茎生物量分配比及相关性状随海拔的变化差异表明冷杉更适应高海拔的胁迫环境。     

How to tell a shrub from a tree: A life‐history perspective from a South African savanna

The ecological differences between ‘shrubs’ and ‘trees’ are surprisingly poorly understood and clear ecological definitions of these two constructs do not exist. It is not clear whether a shrub is simply a small tree or whether shrubs represent a distinct life‐history strategy. This question is of special interest in African savannas, where shrubs and trees often co‐dominate, but are often treated uniformly as ‘woody plants’ even though the tree to shrub ratio is an important determinant of ecosystem functioning. In this study we use data from a long‐term fire experiment, together with a trait‐based approach to test (i) if woody species usually classified as shrubs or trees in African savanna differ in key traits related to disturbance and resource use; and (ii) if these differences justify the interpretation of the two growth forms as distinct life‐history strategies. We measured for 22 of the most common woody plant species of a South African savanna 27 plant traits related to plant architecture, life‐history, leaf characteristics, photosynthesis and resprouting capacity. Furthermore we evaluated their performance during a long‐term fire experiment. We found that woody plants authors call (i) shrubs; (ii) shrubs sometimes small trees; and (3) trees responded differently to long‐term fire treatments. We additionally found significant differences in architecture, diameter‐height‐allometry, foliage density, resprouting vigour after fire, minimum fruiting height and foliar δ¹³C between these three woody plant types. We interpret these findings as evidence for at least two different life‐history‐strategies: an avoidance/adaptation strategy for shrubs (early reproduction + adaptation to minor disturbance) and an escape strategy for trees (promoted investment in height growth + delayed reproduction).     

Modelling the distribution of diameter growth along the stem in Scots pine

This paper presents an empirical model for the distribution of diameter growth along the stem in Scots pine (Pinus sylvestris L.) and for the consequent stem form over time. First, the distribution of annual mass growth in the stem is determined as a function of the total annual growth in stem mass, current stem mass and the distribution of the latter along the stem. Second, the distribution of diameter growth is obtained by converting the fraction of annual growth in the stem mass at a given height in the stem into the thickness of the annual ring at the same height. Application of the model to Scots pine data sets including both young and mature trees not used in parameter estimation showed that the model was capable of reconstructing the distribution of diameter growth from the stem butt to the apex and from the pith to the stem surface at any height in the stem in both young and mature trees. The resulting empirical model was also linked to a physiological, process-based model in order to study its performance in a simulated stand. Simulations representing trees grown in unthinned and thinned Scots pine stands with trees of different status (from dominant to suppressed) showed that the response in tree growth to thinning in terms of the distribution of diameter growth along the stem was quite realistic relative to measured data.