Leaf spectral signatures differ in plant species colonizing habitats along a hydrological gradient

Aims We aimed at determining differences in the leaf spectral signatures of plant species groups growing in habitats along the hydrological gradient of an intermittent wetland and to define leaf traits that explain their variability. We want to contribute to the understanding of the causes for plant spectrum variability at leaf and community levels.Methods We measured leaf reflectance spectra (300–887nm) of representative plant species from different habitats and analyzed spectral differences among species groups. To explain leaf spectra variability within a group, we performed detailed analyses of leaf morphological and biochemical traits in selected species.Important findings The reflectance spectra of the different species groups differed most in the green, yellow and red spectral ranges. The reflectance spectra of submerged leaves of hydrophytes with simple structures were explained by their biochemical traits (carotenoids), while for more complex aerial leaves, morphological traits were more important. In submerged and natant leaves of amphiphytes, total mesophyll and spongy tissue thickness were the most important traits, and these explained 44% and 47%, respectively, of the spectrum variability of each plant group. In general, the redundancy analysis biplots show that samples of different plant species colonizing the same habitat form separate clusters and are related to the explanatory variables in different ways. The redundancy analysis biplots of helophytes and wet meadow species show clustering of graminoids and dicots into two distinct groups. Leaf encrustation (prickle hair properties and epidermis thickness) is important for graminoids, while leaf thickness and specific leaf area have more important roles in dicots. Our results show that knowledge of the species composition and leaf traits is necessary to interpret the reflectance spectra of such plant communities.     

Vertical gradients in leaf trait diversity in a New Zealand forest

Leaves come in a remarkable diversity of sizes and shapes. However, spatial patterns in leaf trait diversity are rarely investigated and poorly resolved. We used a hierarchical approach to evaluate vertical variability in leaf morphology (i.e., leaf trait diversity) in 16 common tree and shrub species inhabiting a New Zealand forest. Height-related heterogeneity in leaf area, specific leaf area, circularity and length to width ratio was analyzed at three scales: (1) among leaves within plants, (2) among plants within species and (3) among species within functional groups (i.e., trees vs. shrubs). Results were scale dependent. Among-leaf morphological diversity was unrelated to plant height. Among-individual morphological diversity increased with the average height of each species, indicating that taller plant species express a greater range of leaf traits than shorter species. Among-species morphological diversity was higher in shrubs than in trees. We hypothesize that scale-dependent patterns in leaf trait diversity result from scale-dependent adaptations to forest environmental conditions. As trees grow from the forest floor into the canopy, they are exposed to a range of environmental conditions, which may select for a range of leaf traits through ontogeny. Conversely, shrubs never reach the forest canopy and may instead be differentially adapted to suites of environmental conditions associated with different stages of forest recovery from tree-fall disturbances. Overall results indicate that vertical patterns in leaf trait diversity exist. However, their strength and directionality are strongly scale-dependent, suggesting that different processes govern leaf shape diversity at different levels of ecological organization.     

不同水氮条件下水稻冠层反射光谱与植株含水率的定量关系

研究了不同土壤水氮条件下水稻 (Oryzasativa) 冠层光谱反射特征和植株水分状况的量化关系。结果表明, 水稻冠层近红外光谱反射率随土壤含水量的降低而降低, 短波红外光谱反射率随土壤含水量的降低而升高。相同土壤水分条件下, 高氮水稻的冠层含水率高于低氮水稻的冠层含水率 ;同一水分条件下, 高氮处理的可见光区和短波红外波段光谱反射率低于低氮处理, 近红外波段光谱反射率高于低氮处理。发现拔节后比值植被指数 (R810 /R460 ) 与水稻叶片含水率和植株含水率呈极显著的线性相关, 模型的检验误差 (RootmeansquareError, RMSE) 分别为 0.93和 1.5 0。表明比值植被指数R810 /R460 可以较好地监测不同生育期水稻叶片和植株含水率。     

鼎湖山南亚热带常绿阔叶林叶功能性状沿群落垂直层次的种内变异

探究功能性状沿着环境梯度如何变化一直以来是基于性状的群落生态学的核心问题之一。尽管功能性状存在种内和种间变异, 但种内变异沿环境梯度如何变化仍有待探究。本文以鼎湖山南亚热带常绿阔叶林1.44 ha塔吊样地内16个树种的2,820个个体为研究对象, 探究4种叶功能性状(比叶面积、叶干物质含量、叶厚度和叶面积)沿群落垂直层次的种内变异。首先, 利用随机效应线性模型量化塔吊样地内的种内变异和种间变异; 其次, 利用Kmeans函数将森林的垂直层次划分为灌木层、亚冠层和林冠层, 并通过构建回归模型探究叶功能性状在群落垂直层次中的种内变异格局。最后, 应用混合线性模型和单因素方差分析的方法探究叶功能性状沿垂直层次的种内变异是否具有物种依赖性。结果表明: 在局域群落中, 并非所有叶功能性状的种内变异都低于种间变异; 叶功能性状在不同垂直层次的种内变异格局存在显著差异, 且种内变异与垂直范围呈正相关; 叶功能性状的种内变异具有较强的物种依赖性, 因此树种差异相对于小环境解释了更多的性状变异; 此外, 不同叶功能性状的种内变异沿垂直层次的变化趋势并不一致。本研究发现种内变异对于物种共存具有重要作用。     

Disparate relatives: Life histories vary more in genera occupying intermediate environments

Species within clades are commonly assumed to share similar life history traits, but within a given region some clades show much greater variability in traits than others. Are variable clades older, allowing more time for trait diversification? Or do they occupy particular environments, providing a wider range of abiotic or biotic opportunities for the establishment and maintenance of diverse trait attributes? Does environmental opportunity increase trait variability across all species, or is it specific to species belonging to the same clade, increasing only within-clade trait variability? We studied the variability of six life-history traits (initiation of flowering, duration of flowering, plant life span, seed mass, stress tolerance, type of reproduction) within 383 angiosperm genera from Central Europe distributed along six abiotic gradients. We compared patterns of within-genus variability to those present in the entire dataset, independent of genus membership. We found that trait variability differed strongly between genera, but did not depend on their age. Trait variability was higher within genera occupying intermediate positions along regional abiotic environmental gradients, compared with patterns across the entire dataset (and unbiased by geographical sampling, family membership or species richness). Increasing trait variability within genera reflected increasing independence of traits from the abiotic environment. We conclude that intermediate abiotic environments play an important role in maintaining and possibly generating the striking diversity of life history traits present within certain clades. They may do so by relaxing the abiotic constraints on the evolution and maintenance of species traits within clades.     

Spatial distribution of functional traits of bryophytes along environmental gradients in an Atlantic Forest remnant in north-eastern Brazil

ABSTRACT

Background: Functional trait-based approaches have been used to identify patterns of plant species diversity and composition related to environmental variability/changes. Bryophytes are rapidly affected by perturbations and thus their traits and distribution are expected to show well-defined relationships with environmental variability.

Aim: To quantify the impacts of fragmentation on the diversity and structure of epiphytic bryophytes to vertical and horizontal environmental gradients in an Atlantic Forest remnant.

Methods: Six functional traits related to water relations and light tolerance were recorded and one of them, the leaf lobule area in liverworts, was selected for morphometric measurements along the gradients analysed. Functional diversity and composition metrics of bryophytes along gradients were compared.

Results: Functional diversity changed little along the vertical and horizontal gradients. Conversely, the functional composition of traits changed markedly. Traits related to water storage, such as the presence and area of lobules, and to protection against excess light incidence, such as dark pigments, were more numerous in the canopy and at the fragment edge.

Conclusions: Functional composition is more correlated with the vertical and horizontal gradients than functional diversity. The lobule of liverworts stood out as the most relevant trait to explain the adaptive strategies of bryophytes.     

Greater variation in boreal plant community composition and community-level traits at local- than regional-scale

Questions

Rapid climate change in northern latitudes is expected to influence plant functional traits of the whole community (community-level traits) through species compositional changes and/or trait plasticity, limiting our ability to anticipate climate warming impacts on northern plant communities. We explored differences in plant community composition and community-level traits within and among four boreal peatland sites and determined whether intra- or interspecific variation drives community-level traits.

Location

Boreal biome of western North America.

Methods

We collected plant community composition and functional trait data along dominant topoedaphic and/or hydrologic gradients at four peatland sites spanning the latitudinal extent of the boreal biome of western North America. We characterized variability in community composition and community-level traits of understorey vascular and moss species both within (local-scale) and among sites (regional-scale).

Results

Against expectations, community-level traits of vascular plant and moss species were generally consistent among sites. Furthermore, interspecific variation was more important in explaining community-level trait variation than intraspecific variation. Within-site variation in both community-level traits and community composition was greater than among-site variation, suggesting that local environmental gradients (canopy density, organic layer thickness, etc.) may be more influential in determining plant community processes than regional-scale gradients.

Conclusions

Given the importance of interspecific variation to within-site shifts in community-level traits and greater variation of community composition within than among sites, we conclude that climate-induced shifts in understorey community composition may not have a strong influence on community-level traits in boreal peatlands unless local-scale environmental gradients are substantially altered.     

Review of optical-based remote sensing for plant trait mapping

Plant trait data have been used in various studies related to ecosystem functioning, community ecology, and assessment of ecosystem services. Evidences are that plant scientists agree on a set of key plant traits, which are relatively easy to measure and have a stable and strong predictive response to ecosystem functions. However, the field measurements of plant trait data are still limited to small area, to a certain moment in time and to certain number of species only. Therefore, remote sensing (RS) offers potential to complement or even replace field measurements of some plant traits. It offers instantaneous spatially contiguous information, covers larger areas and in case of satellite observations profits from their revisit capacity.In this review, we first introduce RS concepts of light–vegetation interactions, RS instruments for vegetation studies, RS methods, and scaling between field and RS observations. Further we discuss in detail current achievements and challenges of optical RS for mapping of key plant traits. We concentrate our discussion on three categorical plant traits (plant growth and life forms, flammability properties and photosynthetic pathways and activity) and on five continuous plant traits (plant height, leaf phenology, leaf mass per area, nitrogen and phosphorous concentration or content). We review existing literature to determine the retrieval accuracy of the continuous plant traits. The relative estimation error using RS ranged between 10% and 45% of measured mean value, i.e. around 10% for plant height of tall canopies, 20% for plant height of short canopies, 15% for plant nitrogen, 25% for plant phosphorus content/concentration, and 45% for leaf mass per area estimates.The potential of RS to map plant traits is particularly high when traits are related to leaf biochemistry, photosynthetic processes and canopy structure. There are also other plant traits, i.e. leaf chlorophyll content, water content and leaf area index, which can be retrieved from optical RS well and can be of importance for plant scientists.We underline the need that future assessments of ecosystem functioning using RS should require comprehensive and integrated measurements of various plant traits together with leaf and canopy spectral properties. By doing so, the interplay between plant structural, physiological, biochemical, phenological and spectral properties can be better understood.     

Relationships between endophyte diversity and leaf optical properties

A single tropical plant species can harbour hundreds of endophyte species within its tissues. Beyond this, little is known about the relationship between endophyte colonization, leaf traits and spectral properties of leaves. We explore these relationships in Coccoloba cereifera, a plant well known for its symbiotic properties. Endophyte richness in C. cereifera was statistically correlated with leaf traits such as water content, the ratio of fresh weight/dry weight and polyphenol/leaf specific weight. Endophyte diversity was also related to spectral vegetation indices of chlorophyll content. The associations among endophyte diversity, leaf traits and spectral reflectance pose new questions and present new opportunities to better understand plant–fungal symbioses and related leaf optical properties.     

Canopy phylogenetic, chemical and spectral assembly in a lowland Amazonian forest

? Canopy chemistry and spectroscopy offer insight into community assembly and ecosystem processes in high-diversity tropical forests, but phylogenetic and environmental factors controlling chemical traits underpinning spectral signatures remain poorly understood. ? We measured 21 leaf chemical traits and spectroscopic signatures of 594 canopy individuals on high-fertility Inceptisols and low-fertility Ultisols in a lowland Amazonian forest. The spectranomics approach, which explicitly connects phylogenetic, chemical and spectral patterns in tropical canopies, provided the basis for analysis. ? Intracrown and intraspecific variation in chemical traits varied from 1.4 to 36.7% (median 9.3%), depending upon the chemical constituent. Principal components analysis showed that 14 orthogonal combinations were required to explain 95% of the variation among 21 traits, indicating the high dimensionality of canopy chemical signatures among taxa. Inceptisols and lianas were associated with high leaf nutrient concentrations and low concentrations of defense compounds. Independent of soils or plant habit, an average 70% (maximum 89%) of chemical trait variation was explained by taxonomy. At least 10 traits were quantitatively linked to remotely sensed signatures, which provided highly accurate species classification. ? The results suggest that taxa found on fertile soils carry chemical portfolios with a deep evolutionary history, whereas taxa found on low-fertility soils have undergone trait evolution at the species level. Spectranomics provides a new connection between remote sensing and community assembly theory in high-diversity tropical canopies.     

Filtering processes in the assembly of plant communities: Are species presence and abundance driven by the same traits?

Question: Is the response of plant traits to environment at the community level similar when considering species abundance and when considering species presence only? Location: Mountain grasslands, central Argentina. Methods: We used data from 57 floristic samples, ordinated through DCCA along moisture and grazing gradients combined with trait values from 85 species (plant height, leaf area, leaf thickness leaf toughness and SLA). For each sample, we calculated the weighted average (considering species abundance) and the simple average (considering only species presence). Through multiple regressions we analysed how each average (dependent variable) responded to moisture and grazing (DCCA scores along Axes 1 and 2, respectively, as independent variables). Results: Weighted averages of all traits were significantly associated to both gradients, while simple averages did not always respond. In some cases the responses followed similar but weaker trends than the responses of weighted averages, but in other cases these responses were qualitatively different. Traits more associated with size (plant height, leaf area, leaf thickness) responded more consistently (similar trends for both averages) to grazing than to moisture, while traits more associated with plant resource acquisition (SLA, leaf toughness) responded more consistently to moisture than to grazing. Conclusion: The trait values and combinations which determine the probability of species presence are not necessary the same as those which determine their probability of becoming abundant. To understand community assembly rules, both species presence and species abundance should be taken into account as the result of different, although closely linked, filtering processes.     

Plant trait responses to the environment and effects on ecosystem properties

A combined analysis of plant trait responses to the environment, and their effects on ecosystem properties has recently been proposed. In this study, we related the trait composition of plant communities to soil nutrients and disturbance as environmental drivers and to productivity, decomposition and soil carbon as ecosystem properties. We surveyed two sites, one comprising intensively grazed and fertilized grasslands, the other consisting of semi-natural grassland and open heathland. Species abundance and trait values of 49 species were recorded in 69 plots, as well as parameters describing soil resources, land-use disturbances, and ecosystem properties. Our main goal was to test whether the average or the diversity of the trait values of the vegetation had stronger effects on ecosystem properties (mass ratio vs. diversity hypothesis). Structural equation modeling was used to perform a simultaneous analysis of trait responses and effects. Specific leaf area and leaf nutrient contents were always negatively correlated with stem dry matter content and canopy height, indicating greater investments in supportive and nutrient-conserving tissue as plants increased in size. In the agricultural site, disturbance was the single most important factor decreasing plant height, while leaf traits such as specific leaf area and leaf nutrient contents increased with soil resources in heathlands. Productivity was directly or indirectly driven by leaf traits, and investments in structural tissue increased standing biomass and soil carbon. Different environmental drivers in the two sites produced opposing leaf trait effects on litter decomposition. Ecosystem properties were explained by the community mean trait value as predicted by the mass ratio hypothesis. Evidence for effects of functional diversity on productivity and other ecosystem properties was not detected, suggesting that diversity–productivity relationships depend on the length of the investigated environmental gradients. We conclude that changes in community composition and dominance hierarchies deserve the most attention when ecosystem properties must be maintained.     

Plant ecophysiological processes in spectral profiles: perspective from a deciduous broadleaf forest

The need for progress in satellite remote sensing of terrestrial ecosystems is intensifying under climate change. Further progress in Earth observations of photosynthetic activity and primary production from local to global scales is fundamental to the analysis of the current status and changes in the photosynthetic productivity of terrestrial ecosystems. In this paper, we review plant ecophysiological processes affecting optical properties of the forest canopy which can be measured with optical remote sensing by Earth-observation satellites. Spectral reflectance measured by optical remote sensing is utilized to estimate the temporal and spatial variations in the canopy structure and primary productivity. Optical information reflects the physical characteristics of the targeted vegetation; to use this information efficiently, mechanistic understanding of the basic consequences of plant ecophysiological and optical properties is essential over broad scales, from single leaf to canopy and landscape. In theory, canopy spectral reflectance is regulated by leaf optical properties (reflectance and transmittance spectra) and canopy structure (geometrical distributions of leaf area and angle). In a deciduous broadleaf forest, our measurements and modeling analysis of leaf-level characteristics showed that seasonal changes in chlorophyll content and mesophyll structure of deciduous tree species lead to a seasonal change in leaf optical properties. The canopy reflectance spectrum of the deciduous forest also changes with season. In particular, canopy reflectance in the green region showed a unique pattern in the early growing season: green reflectance increased rapidly after leaf emergence and decreased rapidly after canopy closure. Our model simulation showed that the seasonal change in the leaf optical properties and leaf area index caused this pattern. Based on this understanding we discuss how we can gain ecophysiological information from satellite images at the landscape level. Finally, we discuss the challenges and opportunities of ecophysiological remote sensing by satellites.

     

Importance of grazing and soil acidity for plant community composition and trait characterisation in coastal dune grasslands

Question: Does grazing by large herbivores affect species composition or community‐wide variation in plant functional traits? Location: Dune grasslands at the Belgian coast. Methods: Plant cover and soil data were collected in 146 plots that were randomly selected at 26 grazed and ungrazed grassland sites. Plant community composition was assessed by Detrended Correspondence Analysis and mean values of plant trait categories were calculated across the plots. Results: Differentiation of plant composition and community‐wide plant trait characteristics was largely determined by grazing, soil acidity and their interaction. In ungrazed situations, a clear floristic distinction appears between acidic (non‐calcareous) and alkaline (calcareous) grasslands. In grazed situations, these floristic differences largely disappeared, indicating that grazing results in a decrease of natural variation in species composition. At higher soil pH, a larger difference in plant community composition and community‐wide plant traits was observed between grazed and ungrazed plots. In ungrazed situations, shifts in plant functional traits along the acidity gradient were observed. Conclusions: Grazing is responsible for shifts in plant community composition, and hence a decrease in plant diversity among grasslands at opposing acidity conditions in coastal dune grasslands. Therefore, care should be taken when introducing grazing as a system approach for nature conservation in dune grasslands as it may eliminate part of the natural variation in plant diversity along existing abiotic gradients.     

Reflectance and transmittance spectra of leaves and shoots of 22 vascular plant species and reflectance spectra of trunks and branches of 12 tree species in Japan

This data paper reports spectral reflectance and transmittance data of leaves from 21 terrestrial vascular plant species (seven herbaceous, and 14 broadleaf and long-needle coniferous tree species) and of shoots from one short-needle coniferous tree species. The reflectance spectra of branches of one tree species, of the trunks of 12 tree species and ground surface of one deciduous broad-leaf forest are also reported. Optical measurements and leaf samplings were made at five sites on Honshu Island, Japan, which are typical vegetation types in East Asia, i.e., grassland, paddy field, and deciduous broad-leaf or coniferous forests. The collection and measurements were conducted for main species in each site. To include other common vegetation types in East Asia, such as evergreen broad-leaf or coniferous forests, the sample collection and the measurements were conducted at gardens and an experimental forest. Leaves of ten deciduous species were measured at different phenological stages from leaf expansion to senescence since those species shows significant seasonal changes in spectral reflectance and transmittance of leaves. Leaves at different position in a canopy (e.g., sunlit versus shaded leaves) were also measured for eight of 21 species. The spectral reflectance and transmittance from both adaxial and abaxial sides of the all leaves or needles, expect Picea abies needles. The measurements of the leaves were conducted with a spectroradiometer attached via an optical fiber to an integrating sphere. Two types of integrating spheres were used: a model LI-1800-12 (Li-Cor) and an RTS-3ZC integrating sphere (Analytical Spectral Devices). A leaf clip accessory was also used instead of an integrating sphere for measuring the leaves of two species. All data were measured within the 350–2,500-nm spectral range with 1-nm steps between measurements but the data obtained by LI-1800 is unavailable in 1,650–1,740, 1,890–1,950, and 2,050–2,500 nm because of a large amount of noise. These data are used as input parameters in a radiative transfer model designed to estimate the leaf area index from radiation reflected from a canopy surface.     

Quantifying relationships between traits and explicitly measured gradients of stress and disturbance in early successional plant communities

Questions: How can one explicitly quantify, and separately measure, stress and disturbance gradients? How do these gradients affect functional composition in early successional plant communities and to what extent? Can we accurately predict trait composition from knowledge of these gradients? Location: Southern Quebec, Canada. Methods: Using eight environmental variables measured in 48 early successional plant communities, we estimated stress and disturbance gradients through structural equation modelling. We then measured 10 functional traits on the most abundant species of these 48 communities and calculated their community‐level mean and variance weighted by the relative abundance of each species. Finally, we related these community‐weighted means and variances to the estimated stress and disturbance gradients using general linear models or generalized additive models. Results: We obtained a well‐fitting measurement model of the stress and disturbance gradients existing in our sites. Of the 10 studied traits, only average plant reproductive height was strongly correlated with the stress (r²=0.464) and disturbance (r²=0.543) gradients. Leaf traits were not significantly related to either the stress or disturbance gradients. Conclusions: The well‐fitting measurement model of the stress and disturbance gradients, combined with the generally weak trait–environment linkages, suggests that community assembly in these early successional plant communities is driven primarily by stochastic processes linked to the history of arrival of propagules and not to trait‐based environmental filtering.     

Predicting leaf traits of herbaceous species from their spectral characteristics

Trait predictions from leaf spectral properties are mainly applied to tree species, while herbaceous systems received little attention in this topic. Whether similar trait–spectrum relations can be derived for herbaceous plants that differ strongly in growing strategy and environmental constraints is therefore unknown. We used partial least squares regression to relate key traits to leaf spectra (reflectance, transmittance, and absorbance) for 35 herbaceous species, sampled from a wide range of environmental conditions. Specific Leaf Area and nutrient‐related traits (N and P content) were poorly predicted from any spectrum, although N prediction improved when expressed on a per area basis (mg/m² leaf surface) instead of mass basis (mg/g dry matter). Leaf dry matter content was moderately to good correlated with spectra. We explain our results by the range of environmental constraints encountered by herbaceous species; both N and P limitations as well as a range of light and water availabilities occurred. This weakened the relation between the measured response traits and the leaf constituents that are truly responsible for leaf spectral behavior. Indeed, N predictions improve considering solely upper or under canopy species. Therefore, trait predictions in herbaceous systems should focus on traits relating to dry matter content and the true, underlying drivers of spectral properties.     

Clonal Growth Strategies Along Flooding and Grazing Gradients in Atlantic Coastal Meadows

Specific composition and species clonal traits were characterized along combined flooding and grazing gradients to answer two questions. i) To what extent does the interaction of flooding and grazing influence the clonal characteristics of the vegetation? ii) Are the effects of both environmental factors independent or interactive? This study was carried out in a wet meadow along the Atlantic coast (France). Three plant communities (hygrophilous, meso-hygrophilous and mesophilous) were distinguished along a flooding gradient and five levels of grazing pressure were controlled through an experimental design (from no grazing to heavy grazing). We monitored species composition and retrieved, for each species, the type of clonal growth organs (CGOs) and clonal traits from the CLO-PLA3 database. We identified two syndromes of clonal traits: ??above-ground splitters?? and ??below-ground integrators??. Clonal traits played a key role in plant assembly in the studied meadows. The interaction of both environmental factors selected for particular syndromes of clonal traits; however, flooding had a stronger filtering effect than grazing. The hygrophilous community was dominated by above-ground splitters, whereas the meso-hygrophilous vegetation was dominated by below-ground integrators. In the mesophilous community, clonal composition was the most diverse and shared clonal traits with the vegetation of both the hygrophilous and meso-hygrophilous communities. Grazing impact on CGOs and clonal traits differed between plant communities, i.e., the effect of grazing was modulated by the flooding regime. This study confirmed that vegetation responses to grazing might depend on the pool of traits, primarily filtered by environmental factors such as flooding.