Comparisons of photosynthesis-related traits among understory lichens,mosses and vascular plant leaves in a high-elevation subalpine forest

对比高海拔亚高山森林林下地衣、藓类及维管植物叶片的光合相关性状 在高海拔亚高山森林中,许多地衣和藓类植物欣欣向荣,甚至成为林地表层的优势物种。尽管它们在森林生态系统中承担着不可替代的功能,但我们对其光合作用相关的功能性状水平知之甚少,对相同生境下藓类植物和维管植物的比较研究也较为匮乏。因此,本研究选择中国青藏高原东缘亚高山林下常见的3种地衣、3种藓类和4种维管植物,测定并对比了它们的碳、氮、磷含量及化学计量比、单位面积光合组织干重、叶绿素含量及光合光响应曲线的异同。研究结果表明,林下地衣的氮含量高于藓类植物,二者的氮、磷、叶绿素、光合速率及光合氮利用效率均低于维管植物叶片。不同种类地衣的光合相关性状与其生长环境相符,而林地表层的3种藓类植物和3种草本植物的功能性状在各自功能群中的种间差异均不显著。总之,这些结果表明高海拔亚高山林下地衣和藓类植物的优势与其光合相关性状关系不大,对严酷环境较强的适应能力可能是它们得以繁荣生长的关键。     

Changes in beta diversity and species functional traits differ between saplings and mature trees in an old‐growth forest

Invasion by generalist tree species can cause biotic homogenization, and such community impoverishment is likely more important in rare forest types. We quantified changes in tree diversity within Carolinian (range in Central Hardwood Forest), central (range in Central Hardwood Forest and Northern Hardwood‐Conifer Forest), and northern species [range reached Northern‐Conifer‐Hardwood/closed Boreal (spruce‐Fir) Forest] in an old forest tract in southern Canada at points surveyed 24 years apart. We asked: How did mature tree and sapling composition and abundance change for the three species’ groups? Did those changes lead to biotic homogenization? Can species’ changes be explained by community traits? We tested for differences in temporal and spatial tree β‐diversity, as well as forest composition and structure, using univariate/multivariate analyses and a community trait‐based approach to identify drivers of change. Major increases occurred in abundance for mature Acer rubrum (northern), while other species decreased (Fraxinus americana, Populus grandidentata); declines were found in A. saccharinum (central) and Cornus florida (Carolinian). Species composition of saplings, but not mature trees, changed due to replacement; no evidence for biotic homogenization existed in either cohort. As a group, northern mature tree species increased significantly, while central species decreased; saplings of pooled Carolinian species also declined. Shade tolerance in mature trees increased, reflecting successional changes, while drought tolerance decreased, perhaps due to changing temperatures, altered precipitation or ground water levels. Saplings showed declines in all traits, probably because of compositional change. Our results demonstrated that saplings can more closely reflect change in forest dynamics than mature trees, especially over short time periods. Based on sapling trends, this remnant could ultimately transition to a mesophytic hardwood stand dominated by A. rubrum and other shade‐tolerant species, creating a more homogeneous forest. While encouraging regeneration for Carolinian and central tree species could ensure high levels of diversity are conserved in the future, it is important to balance this with the primary management goal of maintaining the forest''s old‐growth characteristics.     

Photosynthesis,growth, and decay traits in Sphagnum – a multispecies comparison

Peat mosses (Sphagnum) largely govern carbon sequestration in Northern Hemisphere peatlands. We investigated functional traits related to growth and decomposition in Sphagnum species. We tested the importance of environment and phylogeny in driving species traits and investigated trade‐offs among them. We selected 15 globally important Sphagnum species, representing four sections (subgenera) and a range of peatland habitats. We measured rates of photosynthesis and decomposition in standard laboratory conditions as measures of innate growth and decay potential, and related this to realized growth, production, and decomposition in their natural habitats. In general, we found support for a trade‐off between measures of growth and decomposition. However, the relationships are not strong, with r ranging between 0.24 and 0.45 for different measures of growth versus decomposition. Using photosynthetic rate to predict decomposition in standard conditions yielded R² = 0.20. Habitat and section (phylogeny) affected the traits and the trade‐offs. In a wet year, species from sections Cuspidata and Sphagnum had the highest production, but in a dry year, differences among species, sections, and habitats evened out. Cuspidata species in general produced easily decomposable litter, but their decay in the field was hampered, probably due to near‐surface anoxia in their wet habitats. In a principal components analysis, PCA, photosynthetic capacity, production, and laboratory decomposition acted in the same direction. The species were imperfectly clustered according to vegetation type and phylogeny, so that some species clustered with others in the same section, whereas others clustered more clearly with others from similar vegetation types. Our study includes a wider range of species and habitats than previous trait analyses in Sphagnum and shows that while the previously described growth–decay trade‐off exists, it is far from perfect. We therefore suggest that our species‐specific trait measures offer opportunities for improvements of peatland ecosystem models. Innate qualities measured in laboratory conditions translate differently to field responses. Most dramatically, fast‐growing species could only realize their potential in a wet year. The same species decompose fast in laboratory, but their decomposition was more retarded in the field than that of other species. These relationships are crucial for understanding the long‐term dynamics of peatland communities.     

鼎湖山季风常绿阔叶林两种优势树种的生境适应研究:叶片功能性状和水力结构特征

植物在长期的进化和发展过程中,通过与环境相互作用形成了适应环境的形态结构及生理特征,反映了植物适应环境的生态策略;在森林群落中,地形和土壤等的变异常导致生境的异质性,从而直接或者间接影响物种的分布格局。因此,在生境异质性较强的森林群落中,植物物种分布格局与其生态适应策略有何关系,是值得关注的问题。该文以鼎湖山南亚热带季风常绿阔叶林20 hm2监测样地为平台,针对两种常绿优势树种罗伞树(Ardisia quinquegona)和光叶山黄皮(Aidia canthioides),对比研究了两种植物的叶片功能性状和水力结构特征在山脊、山坡、山谷三种不同生境中的生态适应策略,以阐明物种分布格局与其生态适应策略的关系。结果表明:罗伞树主要是通过调整叶面积(LA)、木材密度(WD)及渗透调节来适应不同生境;光叶山黄皮主要通过调整比叶面积(SLA)、WD及渗透调节,采取养分有效保存(低SLA,高干物质含量)及慢生长高存活的策略以适应不同生境,适应环境能力更强,尤其是在山脊和山坡生境;而且影响两个树种叶片功能性状和水力结构的主导土壤因子有所不同。研究结果说明罗伞树和光叶山黄皮对山脊和山坡生境比山谷更为适应,但在叶片功能性状和水力结构特征方面的生境适应策略不同。     

Tree growth traits and social status affect the wood density of pioneer species in secondary subtropical forest

Wood density (WD) is not only an important parameter to estimate aboveground biomass but also an indicator of timber quality and plant adaptation strategies to stressful conditions (i.e., windthrow, pests, and pathogens). This study had three objectives: (1) to compare WD among seven subtropical tree species; (2) to determine how tree growth traits may influence possible differences in WD between the pioneer and shade‐tolerant species; and (3) to examine whether or not WD differs by tree social status (dominant vs. suppressed trees) within species. To do this, 70 trees were destructively harvested. From each tree, disks at different stem heights were obtained and subjected to a method of stem analysis to measure whole tree level WD. The results showed that WD differed significantly among the seven species (p < .001). Their average WD was 0.537 g/cm³, ranging from 0.409 g/cm³ for Choerospondias axillaris to 0.691 g/cm³ for Cyclobalanopsis glauca. The average WD of the four pioneer species (0.497 ± 0.13 g/cm³) was significantly lower (p < .01) than that of the three shade‐tolerant species (0.589 ± 0.12 g/cm³). The WD of the pioneers had a significant positive correlation with their stem diameter at breast height (DBH), tree height (H), and tree age, but WD had a significant negative correlation with relative growth rate (RGR). In contrast, the WD of the shade‐tolerant tree species had no significant relationships with DBH, H, tree age, or RGR. The dominant trees of the pioneer species had a higher WD than the suppressed trees, whereas the shade‐tolerant species had a lower WD for dominant trees than the suppressed trees. However, the differences in WD between dominant and suppressed trees were not significant. Taken together, the results suggest that classifying species into pioneer and shade‐tolerant groups to examine the effects of tree growth traits and social status could improve our understanding of intra‐ and interspecific variation in WD among subtropical tree species.     

生长型分类方案不同导致森林生态系统植物功能性状的统计偏差

将植物划分为不同的生长型来统计植物功能性状特征,是当前植物性状研究中常用的方法;但生长型分类方案的不同很可能造成植物功能性状统计分析的偏差,对此偏差的评估却尚未见报道。根据植物志描述及野外调查实际情况,将生长型划分为3种不同的分类方案：分类1：根据植物志信息划分为传统意义的乔木和灌木;分类2：根据树高和胸径划分乔木、小乔木和灌木;分类3：仅根据树高划分乔木层与灌木层的乔木和灌木。以东部亚热带常绿阔叶林区域的浙江金华北山35种优势阔叶木本植物的枝叶性状为研究对象,比较不同生长型分类对植物枝叶性状统计数据的影响。结果表明：（1）与传统的分类1相比,分类2对乔木植物枝叶性状影响的显著程度要高于分类3,而对灌木植物枝叶性状的影响程度低于分类3;但不同生长型分类方案中乔木和灌木的枝叶性状总体差异不显著。而与分类2小乔木相比,分类1以及分类2内部的乔木和灌木生长型的性状与分类2小乔木差异非常明显;（2）将不同生长型植物再划分为不同生活型后,不同生长型分类方案对性状统计的影响增大。无论是常绿还是落叶生活型的小乔木,其与不同生活型乔木和灌木的性状差异仍然显著。可见,不同的生长型分类方案可造成植物功能性状统计的差异;把小乔木植物这一功能类群划分出来,能更好地反映森林生态系统性状特征的差异性。     

Functional traits determine heterospecific use of risk‐related social information in forest birds of tropical South‐East Asia

In birds and mammals, mobbing calls constitute an important form of social information that can attract numerous sympatric species to localized mobbing aggregations. While such a response is thought to reduce the future predation risk for responding species, there is surprisingly little empirical evidence to support this hypothesis. One way to test the link between predation risk reduction and mobbing attraction involves testing the relationship between species’ attraction to mobbing calls and the functional traits that define their vulnerability to predation risk. Two important traits known to influence prey vulnerability include relative prey‐to‐predator body size ratio and the overlap in space use between predator and prey; in combination, these measures strongly influence prey accessibility, and therefore their vulnerability, to predators. Here, we combine community surveys with behavioral experiments of a diverse bird assemblage in the lowland rainforest of Sumatra to test whether the functional traits of body mass (representing body size) and foraging height (representing space use) can predict species’ attraction to heterospecific mobbing calls. At four forest sites along a gradient of forest degradation, we characterized the resident bird communities using point count and mist‐netting surveys, and determined the species groups attracted to standardized playbacks of mobbing calls produced by five resident bird species of roughly similar body size and foraging height. We found that (1) a large, diverse subcommunity of bird species was attracted to the mobbing calls and (2) responding species (especially the most vigorous respondents) tended to be (a) small (b) mid‐storey foragers (c) with similar trait values as the species producing the mobbing calls. Our findings from the relatively lesser known bird assemblages of tropical Asia add to the growing evidence for the ubiquity of heterospecific information networks in animal communities, and provide empirical support for the long‐standing hypothesis that predation risk reduction is a major benefit of mobbing information networks.     

Plant species traits across a riparian‐zone/forest ecotone

Abstract. We examined the changes in prevalence of nine plant traits – including the presence of woody stem tissue, leaf longevity, nitrogen fixation, seed longevity, dispersal vector, pollination vector, and clonal growth form – across a riparian/forest‐understory ecotone. This ecotone, found along headwater streams in boreal mixed‐wood forests, supports four distinct vegetation zones: streambank, riparian, transition, and upland forest understory. The objective of this study was to identify specific trait patterns that may indicate functional responses to the changes in environmental factors such as nutrient availability and wind exposure that occur across the ecotone. The suites of plant species traits found in each zone were distinct, with a strong change in the prevalence of several traits. Wind and insect pollination, wind and vertebrate diaspore dispersal, and deciduous and evergreen leaves showed the greatest change in prevalence between the vegetation types. Some traits, including insect pollination and vertebrate diaspore dispersal, were strongly correlated within species. The consistent cooccurrence of pairs of traits in the same species suggests common responses by very different traits to the same environmental factor. This study demonstrates that an ecotone can be characterized not only as a discontinuity in species distributions or environmental factors, but also as a discontinuity in the trait spectrum. Examining ecotones from a trait perspective has strong potential for identifying the environmental factors and associated species functional responses that encourage the development of distinct vegetation boundaries.     

Functional traits of lianas in an Australian lowland rainforest align with post‐disturbance rather than dry season advantage

Lianas are an important component of tropical forests; they alter tree mortality and recruitment and impact biogeochemical cycling. Recent evidence suggests that the abundance of lianas in tropical forests is increasing. To understand and predict the effect of lianas on ecosystem processes in tropical forests, it is important to understand the mechanisms through which they compete with trees. In this study, we investigated the functional traits of lianas and trees in a lowland tropical forest in northeast Queensland, Australia. The site is located at 16.1° south latitude and experiences significant seasonality in rainfall, with pronounced wet and dry seasons. It is also subject to relatively frequent disturbance by cyclones. We asked the question of whether the canopy liana community at this site would display functional traits consistent with a competitive advantage over trees in response to disturbance, or in response to dry season water stress. We found that traits that we considered indicative of a dry season advantage (xylem water δ¹⁸O as an indicator of rooting depth; leaf and stem tissue δ¹³C and instantaneous gas exchange as measures of water‐use efficiency) did not differ between canopy lianas and canopy trees. On the other hand, lianas differed from trees in traits that should confer an advantage in response to disturbance (low wood density; low leaf dry matter content; high leaf N concentration; high mass‐based photosynthetic rates). We conclude that the liana community at the study site expressed functional traits geared towards rapid resource acquisition and growth in response to disturbance, rather than outcompeting trees during periods of water stress. These results contribute to a body of literature which will be useful for parameterising a liana functional type in ecosystem models.     

Tree rings provide no evidence of a CO2 fertilization effect in old‐growth subalpine forests of western Canada

Atmospheric CO₂ concentrations are now 1.7 times higher than the preindustrial values. Although photosynthetic rates are hypothesized to increase in response to rising atmospheric CO₂ concentrations, results from in situ experiments are inconsistent in supporting a CO₂ fertilization effect of tree growth. Tree‐ring data provide a historical record of tree‐level productivity that can be used to evaluate long‐term responses of tree growth. We use tree‐ring data from old‐growth, subalpine forests of western Canada that have not had a stand‐replacing disturbance for hundreds of years to determine if growth has increased over 19th and 20th centuries. Our sample consisted of 5,858 trees belonging to five species distributed over two sites in the coastal zone and two in the continental climate of the interior. We calculated annual increments in tree basal area, adjusted these increments for tree size and age, and tested whether there was a detectable temporal trend in tree growth over the 19th and 20th centuries. We found a similar pattern in 20th century growth trends among all species at all sites. Growth during the 19th century was mostly stable or increasing, with the exception of one of the coastal sites, where tree growth was slightly decreasing; whereas growth during the 20th century consistently decreased. The unexpected decrease in growth during the 20th century indicates that there was no CO₂ fertilization effect on photosynthesis. We compared the growth trends from our four sites to the trends simulated by seven Earth System Models, and saw that most of the models did not predict these growth declines. Overall, our results indicate that these old‐growth forests are unlikely to increase their carbon storage capacity in response to rising atmospheric CO₂, and thus are unlikely to contribute substantially to offsetting future carbon emissions.     

Functional traits determine tree growth and ecosystem productivity of a tropical montane forest: Insights from a long‐term nutrient manipulation experiment

Trait‐response effects are critical to forecast community structure and biomass production in highly diverse tropical forests. Ecological theory and few observation studies indicate that trees with acquisitive functional traits would respond more strongly to higher resource availability than those with conservative traits. We assessed how long‐term tree growth in experimental nutrient addition plots (N, P, and N + P) varied as a function of morphological traits, tree size, and species identity. We also evaluated how trait‐based responses affected stand scale biomass production considering the community structure. We found that tree growth depended on interactions between functional traits and the type or combination of nutrients added. Common species with acquisitive functional traits responded more strongly to nutrient addition, mainly to N + P. Phosphorous enhanced the growth rates of species with acquisitive and conservative traits, had mostly positive effects on common species and neutral or negative effects in rare species. Moreover, trees receiving N + P grew faster irrespective of their initial size relative to trees in control or to trees in other treatment plots. Finally, species responses were highly idiosyncratic suggesting that community processes including competition and niche dimensionality may be altered under increased resource availability. We found no statistically significant effects of nutrient additions on aboveground biomass productivity because acquisitive species had a limited potential to increase their biomass, possibly due to their generally lower wood density. In contrast, P addition increased the growth rates of species characterized by more conservative resource strategies (with higher wood density) that were poorly represented in the plant community. We provide the first long‐term experimental evidence that trait‐based responses, community structure, and community processes modulate the effects of increased nutrient availability on biomass productivity in a tropical forest.     

Using functional traits to predict species growth trajectories,and cross‐validation to evaluate these models for ecological prediction

Modeling plant growth using functional traits is important for understanding the mechanisms that underpin growth and for predicting new situations. We use three data sets on plant height over time and two validation methods—in‐sample model fit and leave‐one‐species‐out cross‐validation—to evaluate non‐linear growth model predictive performance based on functional traits. In‐sample measures of model fit differed substantially from out‐of‐sample model predictive performance; the best fitting models were rarely the best predictive models. Careful selection of predictor variables reduced the bias in parameter estimates, and there was no single best model across our three data sets. Testing and comparing multiple model forms is important. We developed an R package with a formula interface for straightforward fitting and validation of hierarchical, non‐linear growth models. Our intent is to encourage thorough testing of multiple growth model forms and an increased emphasis on assessing model fit relative to a model's purpose.     

Differential survival among life stages contributes to co‐dominance of Abies mariesii and Abies veitchii in a sub‐alpine old‐growth forest

Questions: Are there interspecific differences in mortality and recruitment rates across life stages between two shade‐tolerant dominant trees in a sub‐alpine old‐growth forest? Do such differences in demography contribute to the coexistence and co‐dominance of the two species? Location: Sub‐alpine, old‐growth forest on Mt. Ontake, central Honshu, Japan. Methods: From 1980 to 2005, we recorded DBH and status (alive or dead) of all Abies mariesii and A. veitchii individuals (DBH ≥ 5 cm) in a 0.44‐ha plot. Based on this 25 year census, we quantified mortality and recruitment rates of the two species in three life stages (small tree, 5 cm ≤ DBH < 10 cm; subcanopy tree, 10 cm ≤ DBH < 20 cm; canopy tree, DBH ≥ 20 cm). Results: Significant interspecific differences in mortality and recruitment rates were observed in both the small tree and sub‐canopy tree stages. In this forest, saplings (< 5 cm DBH) are mostly buried by snow‐pack during winter. As a consequence, saplings of A. mariesii, which is snow and shade tolerant, show higher rates of recruitment into the small tree stage than do those of A. veitchii. Above the snow‐pack, trees must tolerate dry, cold temperatures. A. veitchii, which can more readily endure such climate conditions, showed lower mortality rate at the subcanopy stage and a higher recruitment rate into the canopy tree stage. This differential mortality and recruitment among life‐stages determines relative dominance of the two species in the canopy. Conclusion: Differential growth conditions along a vertical gradient in this old forest determine survival of the two species prior to reaching the canopy, and consequently allow co‐dominance at the canopy stage.     

Conservation of genetic diversity in old‐growth forest communities of the southeastern United States

Question: How do studies of the distribution of genetic diversity of species with different life forms contribute to the development of conservation strategies? Location: Old‐growth forests of the southeastern United States. Methods: Reviews of the plant allozyme literature are used to identify differences in genetic diversity and structure among species with different life forms, distributions and breeding systems. The general results are illustrated by case studies of four plant species characteristic of two widespread old‐growth forest communities of the southeastern United States: the Pinus palustris – Aristida stricta (Longleaf pine – wiregrass) savanna of the Coastal Plain and the Quercus – Carya – Pinus (Oak‐hickory‐pine) forest of the Piedmont. Genetic variation patterns of single‐gene and quantitative traits are also reviewed. Results: Dominant forest trees, represented by Pinus palustris(longleaf pine) and Quercus rubra (Northern red oak), maintain most of their genetic diversity within their populations whereas a higher proportion of the genetic diversity of herbaceous understorey species such as Sarracenia leucophylla and Trillium reliquum is distributed among their populations. The herbaceous species also tend to have more population‐to‐population variation in genetic diversity. Higher genetic differentiation among populations is seen for quantitative traits than for allozyme traits, indicating that interpopulation variation in quantitative traits is influenced by natural selection. Conclusion: Developing effective conservation strategies for one or a few species may not prove adequate for species with other combinations of traits. Given suitable empirical studies, it should be possible to design efficient conservation programs that maintain natural levels of genetic diversity within species of conservation interest.