Leaf damage and functional traits along a successional gradient in Brazilian tropical dry forests

Herbivory has significant impacts on individual plants and plant communities, both at ecological and evolutionary time scales. In this context, this study aims to evaluate herbivore damage and its relationship with leaf chemical and structural traits, nutritional status, and forest structural complexity along a successional gradient. We predicted that trees in early successional stages support conservative traits related to drought tolerance (high specific leaf mass and phenolics), whereas trees in light-limited, late successional stages tend to enhance light acquisition strategies (high nitrogen content). We sampled 261 trees from 26 species in 15 plots (50 × 20 m; five per successional stage). From each tree, twenty leaves were collected for leaf trait measures. Phenolic content increased whereas specific leaf mass and nitrogen content decreased from early to late stages. However, leaf damage did not differ among successional stages. Our results partially corroborate the hypothesis that early successional plants in tropical dry forests exhibit leaf traits involved in the conservative use of water. The unexpected decrease in nitrogen content along the chronosequence is likely related to the fact that thinner leaves with low specific leaf mass could have less nitrogen-containing mesophyll per unit area. Mechanisms affecting herbivory intensity varied across scales: at the species level, leaf damage was negatively correlated with tannin concentration and specific leaf mass; at the plot level, leaf damage was positively affected by forest structural complexity. Herbivory patterns in tropical forests are difficult to detect because abiotic factors and multiple top-down and bottom-up forces directly and indirectly affect herbivores.     

Plant hydraulics and photosynthesis of 34 woody species from different successional stages of subtropical forests

It is important to understand the ecophysiological characters of plants when exploring mechanisms underlying species substitution in the process of plant succession. In the present study, we selected 34 woody species from different stages of secondary succession in subtropical forests of southern China, and measured their hydraulic conductivity, gas exchange rates, leaf nutrients and drought‐tolerance traits such as xylem resistance to cavitation, turgor loss point and carbon isotope ratio. Principal component analysis revealed that early‐, mid‐ and late‐successional species were significantly separated along axis 1, which was strongly associated with hydraulic‐photosynthetic coordination. In contrast to species distributed in late‐successional forest, early‐successional species had the highest hydraulic conductivity, net photosynthetic rates, photosynthetic nitrogen and phosphorus use efficiencies, but had the lowest photosynthetic water‐use efficiency. However, changes of the measured drought‐tolerance traits of the 34 species along the succession did not demonstrate a clear trend – no significant correlations between these traits and plant successional stages were found. Moreover, the trade‐off between hydraulic efficiency and safety was not identified. Taken together, our results suggested that hydraulic efficiency and photosynthetic function, rather than drought tolerance, play an important role in species distributions along plant succession in subtropical forests.     

Plant functional traits under cattle grazing and fallow age scenarios in a tropical dry forest of Northwestern Mexico

Evaluating plant functional traits helps to understand how plants respond to changing environmental conditions and resource availability associated with disturbance events. Livestock production is one of the primary drivers of tropical forest loss and degradation. Livestock alter environmental conditions within the forest by grazing, trampling and nutrient inputs, which in turn can influence species composition and functional traits of species. Understanding how livestock influence functional traits along a successional gradient is poorly understood. Here, we studied the effect of cattle grazing and fallow age on plant functional traits and soil nutrients in secondary and old-growth tropical dry forests. We analyzed plant functional traits of the most important species in successional and old-growth forest communities in both cattle present and cattle excluded plots. Our results showed the effects of cattle grazing and fallow age on plant functional traits, with fallow age explaining more variation than cattle grazing. In early succession, functional traits were associated with water conservation (thicker leaves, lower specific leaf area), and in later successional they were linked with sunlight conservation (larger height, higher specific leaf area). The presence of large fruits and seeds in advanced successional sites suggests high resource availability, which may help plants to successfully reproduce. Moreover, under cattle grazing some functional traits are associated with herbivory defense (high foliar dry weight and thick leaves). Even though N and C increased as succession advanced, the sites with cattle grazing had higher NH4 and NO3 concentrations as a result of fecal deposition. Plant functional traits responded to fallow age than to cattle grazing. Our study showed that cattle exclusion, as a management and biodiversity conservation strategy, contributes positively to soil nutrition. Thus, fallow age and cattle exclusion facilitate soil recovery and allows establishing species with suitable functional attributes for overcoming environmental filters in abandoned cattle fields.     

Impacts of Land Abandonment on Vegetation: Successional Pathways in European Habitats

Changes in traditional agricultural systems in Europe in recent decades have led to widespread abandonment and colonization of various habitats by shrubs and trees. We combined several vegetation databases to test whether patterns of changes in plant diversity after land abandonment in different habitats followed similar pathways. The impacts of land abandonment and subsequent woody colonization on vegetation composition and plant traits were studied in five semi-natural open habitats and two arable habitats in six regions of Europe. For each habitat, vegetation surveys were carried out in different stages of succession using either permanent or non-permanent plots. Consecutive stages of succession were defined on a physiognomic basis from initial open stages to late woody stages. Changes in vegetation composition, species richness, numbers of species on Red Lists, plant strategy types, Ellenberg indicator values of the vegetation, Grime CSR strategy types and seven ecological traits were assessed for each stage of the successional pathway. Abandonment of agro-pastoral land-use and subsequent woody colonization were associated with changes in floristic composition. Plant richness varied according to the different habitats and stages of succession, but semi-natural habitats differed from arable fields in several ecological traits and vegetation responses. Nevertheless, succession occurred along broadly predictable pathways. Vegetation in abandoned arable fields was characterized by a decreasing importance of R-strategists, annuals, seed plants with overwintering green leaves, insect-pollinated plants with hemi-rosette morphology and plants thriving in nutrient-rich conditions, but an increase in species considered as endangered according to the Red Lists. Conversely, changes in plant traits with succession within the initially-open semi-natural habitats showed an increase in plants thriving in nutrient-rich conditions, stress-tolerant plants and plants with sexual and vegetative reproduction, but a sharp decrease in protected species. In conclusion, our study showed a set of similarities in responses of the vegetation in plant traits after land abandonment, but we also highlighted differences between arable fields and semi-natural habitats, emphasizing the importance of land-use legacy.     

Combined use of leaf size and economics traits allows direct comparison of hydrophyte and terrestrial herbaceous adaptive strategies

Background and Aims

Hydrophytes generally exhibit highly acquisitive leaf economics. However, a range of growth forms is evident, from small, free-floating and rapidly growing Lemniden to large, broad-leaved Nymphaeiden, denoting variability in adaptive strategies. Traits used to classify adaptive strategies in terrestrial species, such as canopy height, are not applicable to hydrophytes. We hypothesize that hydrophyte leaf size traits and economics exhibit sufficient overlap with terrestrial species to allow a common classification of plant functional types, sensu Grime''s CSR theory.

Methods

Leaf morpho-functional traits were measured for 61 species from 47 water bodies in lowland continental, sub-alpine and alpine bioclimatic zones in southern Europe and compared against the full leaf economics spectrum and leaf size range of terrestrial herbs, and between hydrophyte growth forms.

Key Results

Hydrophytes differed in the ranges and mean values of traits compared with herbs, but principal components analysis (PCA) demonstrated that both groups shared axes of trait variability: PCA1 encompassed size variation (area and mass), and PCA2 ranged from relatively dense, carbon-rich leaves to nitrogen-rich leaves of high specific leaf area (SLA). Most growth forms exhibited trait syndromes directly equivalent to herbs classified as R adapted, although Nymphaeiden ranged between C and SR adaptation.

Conclusions

Our findings support the hypothesis that hydrophyte adaptive strategy variation reflects fundamental trade-offs in economics and size that govern all plants, and that hydrophyte adaptive strategies can be directly compared with terrestrial species by combining leaf economics and size traits.     

Differences in xylem and leaf hydraulic traits explain differences in drought tolerance among mature Amazon rainforest trees

Considerable uncertainty surrounds the impacts of anthropogenic climate change on the composition and structure of Amazon forests. Building upon results from two large‐scale ecosystem drought experiments in the eastern Brazilian Amazon that observed increases in mortality rates among some tree species but not others, in this study we investigate the physiological traits underpinning these differential demographic responses. Xylem pressure at 50% conductivity (xylem‐P₅₀), leaf turgor loss point (TLP), cellular osmotic potential (π_o), and cellular bulk modulus of elasticity (ε), all traits mechanistically linked to drought tolerance, were measured on upper canopy branches and leaves of mature trees from selected species growing at the two drought experiment sites. Each species was placed a priori into one of four plant functional type (PFT) categories: drought‐tolerant versus drought‐intolerant based on observed mortality rates, and subdivided into early‐ versus late‐successional based on wood density. We tested the hypotheses that the measured traits would be significantly different between the four PFTs and that they would be spatially conserved across the two experimental sites. Xylem‐P₅₀, TLP, and π_o, but not ε, occurred at significantly higher water potentials for the drought‐intolerant PFT compared to the drought‐tolerant PFT; however, there were no significant differences between the early‐ and late‐successional PFTs. These results suggest that these three traits are important for determining drought tolerance, and are largely independent of wood density—a trait commonly associated with successional status. Differences in these physiological traits that occurred between the drought‐tolerant and drought‐intolerant PFTs were conserved between the two research sites, even though they had different soil types and dry‐season lengths. This more detailed understanding of how xylem and leaf hydraulic traits vary between co‐occuring drought‐tolerant and drought‐intolerant tropical tree species promises to facilitate a much‐needed improvement in the representation of plant hydraulics within terrestrial ecosystem and biosphere models, which will enhance our ability to make robust predictions of how future changes in climate will affect tropical forests.     

浙江天童常绿阔叶林不同演替阶段木本植物的水力结构特征

水力结构是植物应对环境形成的与水分运输相关的形态策略.探索不同演替阶段和群落不同高度层植物的水力结构特征, 有助于理解植物的水分运输和利用策略.该研究以浙江天童常绿阔叶林演替前中后期群落的上层木(占据林冠层的树种)和下层木(灌木层物种)为对象, 测定了演替共有种(至少存在于两个演替阶段的物种)和更替种(仅存在于某一演替阶段的物种)的枝边材比导率,叶比导率和胡伯尔值, 以及边材疏导面积,末端枝总叶面积和枝条水势, 分析植物水力结构在群落上层木和下层木间以及在演替阶段间的差异, 及其与枝叶性状的相关关系.结果显示: (1)上层木植物边材比导率和叶比导率显著高于下层木植物(p < 0.05); (2)上层木和下层木的边材比导率与叶比导率在演替阶段间均无显著差异(p > 0.05); 上层木的胡伯尔值在演替阶段间无显著差异, 下层木的胡伯尔值随演替显著下降(p < 0.05); (3)上层木共有种仅边材比导率随演替进行显著降低(p < 0.05), 更替种的3个水力结构参数在演替阶段间无显著差异; 下层木共有种水力结构参数在演替阶段间无明显差异, 更替种仅胡伯尔值随演替减小(p < 0.05); (4)植物边材比导率与枝疏导面积和末端枝所支撑的总叶面积显著正相关(p < 0.01), 胡伯尔值与枝条水势及末端枝总叶面积显著负相关(p < 0.01).以上结果表明: 天童常绿阔叶林演替各阶段上层木比下层木具有更大的输水能力和效率; 随着演替进行, 上层木与下层木的共有种和更替种边材比导率的相反变化表明上层木水力结构的变化可能由微生境变化引起, 而下层木水力结构特征的变化可能由物种更替造成.     

Hydraulic architecture of evergreen broad-leaved woody plants at different successional stages in Tiantong National Forest Park,Zhejiang Province,China

Aims Hydraulic architecture is a morphological strategy in plants to transport water in coping with environmental conditions. Change of hydraulic architecture for plants occupying different canopy layers within community and for the same plant at different successional stages reflect existence and adaptation in plant's water transportation strategies. The objective of this study was to examine how hydraulic architecture varies with canopy layers within a community and with forest succession.Methods The study site is located in Tiantong National Forest Park, Zhejiang Province, China. Hydraulic architectural traits studied include sapwood-specific hydraulic conductivity, leaf-specific hydraulic conductivity, Huber value, sapwood channel area of twigs, total leaf area per terminal twig, and water potential of twigs. We measured those traits for species that occur in multiple successional stages (we called it "overlapping species") and for species that occur only in one successional stage (we called it "turnover species") along a successional series of evergreen broadleaved forests. For a given species, we sampled both overstory and understory trees. Hydraulic architectural traits between overstory and understory trees in the same community and at successional stages were compared. Pearson correlation was used to exam the relationship between hydraulic architectural traits and the twig/leaf traits.Important findings Sapwood-specific hydraulic conductivities and leaf-specific hydraulic conductivities were significantly higher in overstory trees than those in understory trees, but did not significantly differ from successional stages. Huber value decreased significantly for understory trees, but did not change for overstory trees through forest successional stages. For overstory trees, a trend of decreasing sapwood-specific hydraulic conductivity was observed for overlapping species but not for turnover species with successional stages. In contrast, for understory trees, a trend of decreasing Huber values was observed for turner species but not for overlapping species with successional stages. Across tree species, sapwood-specific hydraulic conductivity was positively correlated with sapwood channel area and total leaf area per terminal twig size. Huber value was negatively correlated to water potential of twigs and total leaf area per terminal twig size. These results suggest that water transportation capacity and efficiency are higher in overstory trees than in understory trees across successional stages in evergreen broadleaved forests in Tiantong region. The contrasting trends of sapwood-specific hydraulic conductivity between overlapping species and turnover species indicate that shift of microenvironment conditions might lead to changes of hydraulic architecture in overstory trees, whereas species replacement might result in changes of hydraulic architecture in understory trees.     

Phenology and Stem Diameter Increment Seasonality in a Costa Rican Wet Tropical Forest

The relationship between phenology and tree stem diameter increment is largely unexplored in tropical species, especially in wet tropical forests. To explore links between these phenomena, we measured stem diameter increment and phenology of ten canopy tree species from a range of functional types in the Atlantic lowlands of Costa Rica to test for seasonal and interannual patterns. We measured stem diameter increment using band dendrometers and visually assessed leaf and reproductive phenology monthly from 1997 to 2000. We categorized the species into groups based on patterns of leaf exchange and reproduction. Species were either deciduous with synchronous or asynchronous leaf drop, or evergreen with continuous or seasonal leaf flushing. Flowering occurred supra-annually, annually, or continuously. Of the ten species studied, four species, Cecropia insignis, Dipteryx panamensis, Lecythis ampla, and Simarouba amara , had consistent seasonal stem diameter increment patterns in both years. Dipteryx panamensis and L. ampla were deciduous with synchronized leaf drop . Cecropia insignis was evergreen and produced new leaves continuously. Simarouba amara , also evergreen, exchanged leaves over a brief period once a year. We tested whether stem diameter increment was correlated to phenology using logistic regression. Leaflessness significantly explained patterns in stem diameter increment but reproductive phenology did not. Deciduous trees were 2.6–9.3 times more likely to grow less than average the month following leaffall than in months when trees had full crowns.     

Amazonia trees have limited capacity to acclimate plant hydraulic properties in response to long‐term drought

The fate of tropical forests under future climate change is dependent on the capacity of their trees to adjust to drier conditions. The capacity of trees to withstand drought is likely to be determined by traits associated with their hydraulic systems. However, data on whether tropical trees can adjust hydraulic traits when experiencing drought remain rare. We measured plant hydraulic traits (e.g. hydraulic conductivity and embolism resistance) and plant hydraulic system status (e.g. leaf water potential, native embolism and safety margin) on >150 trees from 12 genera (36 species) and spanning a stem size range from 14 to 68 cm diameter at breast height at the world's only long‐running tropical forest drought experiment. Hydraulic traits showed no adjustment following 15 years of experimentally imposed moisture deficit. This failure to adjust resulted in these drought‐stressed trees experiencing significantly lower leaf water potentials, and higher, but variable, levels of native embolism in the branches. This result suggests that hydraulic damage caused by elevated levels of embolism is likely to be one of the key drivers of drought‐induced mortality following long‐term soil moisture deficit. We demonstrate that some hydraulic traits changed with tree size, however, the direction and magnitude of the change was controlled by taxonomic identity. Our results suggest that Amazonian trees, both small and large, have limited capacity to acclimate their hydraulic systems to future droughts, potentially making them more at risk of drought‐induced mortality.     

Variations and Trade‐offs in Functional Traits of Tree Seedlings during Secondary Succession in a Tropical Lowland Rain Forest

The seedling stage is generally the most important bottleneck for the successful regeneration of trees in forests. The traits of seedlings, particularly biomass allocation and root traits, are more easily quantified than the traits of adults. In this study, we tested the hypothesis that seedling traits vary and trade‐off tracking the changing environment during secondary succession. We measured the major morphological traits of 27 dominant species and the major environmental factors in a chronosequence (30‐yr‐old fallow, 60‐yr‐old fallow, and old growth forest) after shifting cultivation in a tropical lowland rain forest on Hainan Island, China. The 30‐yr‐old fallow had higher light and nutrient availability, and the older forests had higher soil water content. Redundancy analysis based on species abundance and environmental factors revealed groups of seedlings that dominate in different stages of succession. Seedlings in different stages of succession had different strategies of biomass allocation for harvesting resources that varied in availability. Species characteristic of younger forest had higher allocation to roots and higher specific leaf area, while species characteristic of older forest had higher allocation to leaves. Our study suggests that the variations and trade‐offs in the major functional traits of tree seedlings among successional classes may reflect changes in environmental conditions during succession.     

Plant phenology,leaf traits and leaf litterfall of contrasting life forms in the arid Patagonian Monte,Argentina

Question: Do coexisting plant life forms differ in overall phenology, leaf traits and patterns of leaf litterfall? Location: Patagonian Monte, Chubut Province, Argentina. Methods: We assessed phenology, traits of green and senesced leaves and the pattern of leaf litterfall in 12 species of coexisting life forms (perennial grasses, deciduous shrubs, evergreen shrubs). Results: We did not identify differences in phenology, leaf traits and patterns of leaf litterfall among life forms but these attributes contrasted among species. Independent of the life form, the maintenance of green leaves or vegetative growth during the dry season was mostly associated with leaves with high leaf mass per area (LMA) and high concentration of secondary compounds. Low LMA species produced low litterfall mass with low concentration of secondary compounds, and high N concentration. High LMA species produced the largest mass of leaf litterfall. Accordingly, species were distributed along two main dimensions of ecological variation, the dimension secondary compounds in leaves ‐ length and timing of the vegetative growth period (SC ‐ VGP) and the dimension leaf mass per area ‐ leaf litterfall mass (LMA ‐ LLM). Conclusions: Phenology, leaf traits and leaf litterfall varied among species and overlapped among life forms. The two dimensions of ecological variation among species (SC ‐ VGP, LMA ‐ LLM) represent distinct combinations of plant traits or strategies related to resource acquisition and drought tolerance which are reflected in the patterns of leaf litterfall.     

Drought resistance in early and late secondary successional species from a tropical dry forest: the interplay between xylem resistance to embolism,sapwood water storage and leaf shedding

The mechanisms of drought resistance that allow plants to successfully establish at different stages of secondary succession in tropical dry forests are not well understood. We characterized mechanisms of drought resistance in early and late‐successional species and tested whether risk of drought differs across sites at different successional stages, and whether early and late‐successional species differ in resistance to experimentally imposed soil drought. The microenvironment in early successional sites was warmer and drier than in mature forest. Nevertheless, successional groups did not differ in resistance to soil drought. Late‐successional species resisted drought through two independent mechanisms: high resistance of xylem to embolism, or reliance on high stem water storage capacity. High sapwood water reserves delayed the effects of soil drying by transiently decoupling plant and soil water status. Resistance to soil drought resulted from the interplay between variations in xylem vulnerability to embolism, reliance on sapwood water reserves and leaf area reduction, leading to a tradeoff of avoidance against tolerance of soil drought, along which successional groups were not differentiated. Overall, our data suggest that ranking species' performance under soil drought based solely on xylem resistance to embolism may be misleading, especially for species with high sapwood water storage capacity.     

Fit for succession – community structure and life strategies of leafhoppers in urban brownfields

Abstract.  1. Urban brownfields offer an excellent opportunity to study successional processes. Changes in the frequencies of biological traits during succession are of particular interest. They shed light on the general reasons why species emerge and vanish during the course of succession.
2. Leafhopper (Hemiptera: Auchenorrhyncha) occurrence data (3763 species observations) of 194 species were studied. Data was collected on 246 brownfield plots, aged 0–40 years, in two cities in Northern Germany.
3. Four categorical traits were studied: host-plant type, phagy, dormancy, and voltinism. In these traits, two aspects were analysed: (1) changes in trait category frequencies during succession, and (2) distribution of trait categories (i) within the brownfield species pool versus the German species pool and (ii) within species observations.
4. Trait categories showed clear successional trends. Young successional stages were related to feeding on herbs, polyphagy, egg overwintering, and two generations per year.
5. By analysing combinations of two traits, species could be assigned to four functional groups: species associated with young, intermediate and old sites, and one group indifferent to site age. The pioneer group comprised the least number of species, but the highest number of observations.
6. Categories associated with young site age were over-represented in the brownfield species pool. Moreover, within this already biased species pool, species with pioneer trait categories occurred with higher frequency. For the slow colonisers among leafhoppers, brownfields seem to be habitats that are hard to exploit.     

海岛植物不同演替阶段植物功能性状与环境因子的变化规律

植物功能性状与环境之间的关系是功能性状研究的重点,环境因子驱使植物功能性状发生变化,进而推动群落发生演替。以平潭岛4个不同演替阶段的森林植被(灌草丛、针叶林、针阔混交林、常绿阔叶林)为研究对象,结合不同群落演替阶段的物种特征和群落结构,分析海岛不同演替阶段茎、叶功能性状的变化规律,以及功能性状与环境因子的关系。结果表明:(1)随着演替的进行,土壤养分和水分逐渐增加,土壤pH逐渐下降。比叶面积(SLA)、叶片氮含量(LNC)、叶片磷含量(LPC)、茎氮含量(SNC)、茎磷含量(SPC)下降后上升,叶厚度(LT)、叶片碳含量(LCC)、茎碳含量(SCC)与之相反,叶干物质含量(LDMC)、茎组织密度(STD)逐渐上升。(2)冗余分析表明,演替早期植物主要分布在土壤pH、容重高的贫瘠环境,拥有较高SLA、SNC、SPC、LPC的性状组合;演替后期植物主要分布在土壤养分和水分高的肥沃环境,拥有较高的STD、LDMC、LCC、LNC的性状组合。其中,土壤有机质和全氮含量是影响海岛植物演替过程中功能性状变化的关键环境因子。研究海岛植物功能性状与环境之间的关系随演替的变化规律,探讨各演替阶段功能性状和环境特征,以及功能性状如何响应环境变化。旨在为今后选择合适的树种进行海岛植被修复和重建提供依据。     

Shift in community structure in an early‐successional Mediterranean shrubland driven by long‐term experimental warming and drought and natural extreme droughts

Global warming and recurring drought are expected to accelerate water limitation for plant communities in semiarid Mediterranean ecosystems and produce directional shifts in structure and composition that are not easily detected, and supporting evidence is scarce. We conducted a long‐term (17 years) nocturnal‐warming (+0.6°C) and drought (?40% rainfall) experiments in an early‐successional Mediterranean shrubland to study the changes in community structure and composition, contrasting functional groups and dominant species, and the superimposed effects of natural extreme drought. Species richness decreased in both the warming and drought treatments. Responses to the moderate warming were associated with decreases in herb abundance, and responses to the drought were associated with decreases in both herb and shrub abundances. The drought also significantly decreased community diversity and evenness. Changes in abundance differed between herbs (decreases) and shrubs (increases or no changes). Both warming and drought, especially drought, increased the relative species richness and abundance of shrubs, favoring the establishment of shrubs. Both warming and drought produced significant shifts in plant community composition. Experimental warming shifted the community composition from Erica multiflora toward Rosmarinus officinalis, and drought consistently shifted the composition toward Globularia alypum. The responses in biodiversity (e.g., community biodiversity, changes of functional groups and compositional shifts) were also strongly correlated with atmospheric drought (SPEI) in winter–spring and/or summer, indicating sensitivity to water limitation in this early‐successional Mediterranean ecosystem, especially to natural extreme droughts. Our results suggest that the shifts in species assembles and community diversity and composition are accelerated by the long‐term nocturnal‐warming and drought, combined with natural severe droughts, and that the magnitude of the impacts of climate change is also correlated with the successional status of ecosystem. The results thus highlight the necessity for assessing the impacts on ecosystemic functioning and services and developing effective measures for conserving biodiversity.     

Allocation strategies and seed traits are hardly affected by nitrogen supply in 18 species differing in successional status

Species performance depends on ecological strategies, revealed by suites of traits, conferring different relative ecological advantages in different environments. Although current knowledge on plant strategies along successional gradients is derived from studies conducted in situ, actually quantifying these strategies requires disentangling the effects of environmental factors from intrinsic differences between species.Here we tested whether allocation strategies and seed traits differ among successional stages and nitrogen levels. To this aim, we assessed biomass and nitrogen allocations and seed traits variations for 18 species, differing in life history and belonging to three stages of a Mediterranean old-field succession. These species were grown as monocultures in an experimental garden under limiting and non-limiting nitrogen supply.Early successional species allocated allometrically more nitrogen and proportionally more biomass to reproduction, and set more seeds than later successional species. Seed mass increased with successional status and was negatively related to seed number. Early successional species thus produced more but less-provisioned seeds, suggesting better colonization abilities. These patterns were not the sole consequence of the replacement of annuals by perennials along the successional gradient, since comparable trends were also observed within each life history. Allocation patterns were generally not altered by nitrogen supply and the higher nitrogen content in vegetative organs of plants grown under high nitrogen supply was not retranslocated from leaves to seeds during seed development.We therefore conclude that differences in plant ecological strategies in species characteristics from contrasting successional stages appear to be intrinsic properties of the studied species, and independent from environmental conditions.     

Reddish spring colouring of deciduous leaves: a sign of ecotype?

Several deciduous woody plant species produce anthocyanins during leaf development in spring and again during leaf senescence in autumn. The leaves of Betula pendula Roth (silver birch) commonly exhibit transient reddening in juvenile leaves under northern growing conditions, with the intensity of the red colour varying among individual trees. The objective of our study was to test the hypothesis that the accumulation of foliar anthocyanins during spring in leaves of B. pendula is an ecotypic response. Chlorophyll fluorescence ratio (Fv/Fm), leaf reflectance and anthocyanin concentrations were measured, in relation to phenology in spring, summer and autumn from birches used for landscaping with either red or green-emergent leaves. The results suggest that (1) the trees with green- or red-emergent juvenile leaves represent different populations, and (2) that the red-emergent leaves senesced earlier, indicating that (3) trees with red-emerging leaves belong to a more northern ecotype than the trees with green-emerging leaves. The role of anthocyanin synthesis in a northern radiation environment is discussed.     

Differential drought responses between saplings and adult trees in four co-occurring species of New England

Tree-ring characteristics in four species were examined to address whether co-occurring mature trees of different successional status respond differently to drought, and whether saplings of these species have a greater response to drought than mature trees. We examined saplings and mature trees of paper birch, yellow birch, red maple and sugar maple, which varied in successional status (shade-tolerance) and co-occurred at Harvard Forest, Petersham, Mass., USA. Three drought events in 1964–1966, 1981 and 1995 were identified using climate data. For mature trees, there was no significant interspecific difference in relative changes in ring-width index (RWI) during the 1964–1966 and 1995 drought events. However, the interspecific difference was significant in the 1981 drought event. Response function analysis for mature trees showed that the radial growth of sugar maple was mainly controlled by spring and summer precipitation, red maple by spring and summer precipitation and temperature, yellow birch by winter and summer precipitation, and spring and summer temperature, and paper birch by spring and summer precipitation and spring temperature. Saplings of sugar maple and yellow birch, but not red maple and paper birch, showed significant positive correlations between RWI and annual total precipitation. In the 1995 drought event, saplings and mature trees of red maple and paper birch differed significantly in drought responses, but this was not true in sugar maple and yellow birch. Our results do not support a generally greater response in saplings than in mature trees, nor an early- versus late successional difference in drought responses.     

Phenology of a common roadside fig in Sarawak

The phenology of a dioecious fig (Ficus fulva, Reinw. ex Bl.; 25 female, 26 male trees) was studied at Lambir Hills National Park, Sarawak. Dioecious fig phenology provides an excellent opportunity to investigate the influence of climate and sexual specialization on the obligate fig–fig pollinator/ovule parasite interaction. Leaf phenology was strongly correlated between sexes. Trees dropped leaves during drought and initiated new leaf growth after the renewal of rain. Before the production of large crops of syconia, trees shed their leaves and then new leaves and syconia were initiated together. Syconia were produced in synchronous crops with asynchrony between trees maintaining a relatively even production of syconia within the tree group. Syconia abortion on male but not female trees, was negatively correlated with the proportion of trees with male phase syconia. A severe drought in early 1998 significantly disrupted the phenology thereafter. The duration of crop development was approximately twice as long on female trees as on males, and total syconia production was much higher on male trees. Plots of syconia diameter versus dry weight suggest sexual specialization in the investment profile during crop development. Male trees also sometimes produced a small crop of syconia immediately before a large crop, probably to supply wasps for the main crop. Sexes had different growth strategies with male trees growing more as small individuals and slightly delaying reproduction. Diameter at breast height was significantly correlated with total syconia production in male trees but not in females. Syconia production was best predicted by canopy width.