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.     

Fish assemblages across a complex,tropical freshwater/marine ecotone

Synopsis Riverine fish assemblages in the temperate zone generally show strong longitudinal patterns of faunal turnover and increases in species richness with increasing stream order. We examined the composition and structure of tropical fish assemblages across a complex freshwater/marine ecotone in Tortuguero National Park on the Caribbean coast of Central America. Species turnover was high between four characteristic habitats that largely corresponded with a longitudinal gradient of stream order over distances of less than 30 km. Suites of common fish species characterized each habitat: creeks, rivers, lagoons, and the sea. In addition to the habitat endemics, several species spanned two habitat types, but only three species were collected in more than two habitats. Multivariate gradient analysis of fish assemblages reflected a gradient of habitats that to some extent corresponded to fluvial distances. Due to the unusual configuration of coastal lagoons lying parallel to the coast, the ordination gradient showed little correlation with linear distance to the coast. Environmental variables related to habitat size and salinity showed greatest correspondence with the fish assemblage ordination gradient. Invertebrate-feeding fishes were the predominant trophic group in 15 of 16 fish assemblages, and inland creek sites contained a greater proportion of herbivores and omnivores than other sites. The relative fraction of herbivorous and detritivorous fishes showed a monotonic decline along the longitudinal habitat gradient from inland to coast. Patterns of species composition and richness at Tortuguero Park appeared to agree well with earlier models of factors influencing temperate zone stream fishes. Headwaters have low aquatic primary productivity and contain small colonizing fish species subject to large fluctuations in local densities and intermittent competition. Lagoons contain both large and small species, the latter being restricted largely to shallow edge habitats by predation. Lagoons exhibit more lentic environmental conditions, experience relatively fewer periodic disturbances than headwaters, and their assemblages are inferred to be under relatively greater influence of biotic factors. Fish assemblages of rivers and caños (swampy side channels and braids) appear to be under less abiotic control than headwaters and influenced less by biotic factors than lagoons.     

Leaf functional traits vary with the adult height of plant species in forest communities

Aims Within-community variation accounts for a remarkable proportion of the variation in leaf functional traits. Plant height may be used to explain within-community variances of leaf traits because different microenvironments, especially light intensity, may occur at different heights. This study determines whether or not leaf nitrogen (N) and phosphorus (P) contents as well as leaf mass per area (LMA) are interspecifically correlated with the adult height of forest woody species. We also discuss these relationships with respect to community structure and functions of the ecosystem.Methods A total of 136 dicotyledonous woody species from 6 natural forests (3 evergreen and 3 deciduous ones) in East China (18°44′–45°25′N, 108°50′–128°05′E) were investigated. For each of the 157 species–site combinations, 6 traits were measured: plant adult height relative to the forest canopy (H R), leaf N and P contents per unit area (N area and P area), N and P contents per unit dry mass (N mass and P mass) and LMA. The total variances of each leaf trait across sites were partitioned in a hierarchical manner. The relationships between leaf traits and H R within forest communities were then analyzed using both standardized major axis regression and Felsenstein's phylogenetic independent contrasts. Relationships between evergreen and deciduous forests were compared by linear mixed models.Important findings H R is a robust predictor of leaf N area, P area and LMA, explaining 36.7, 39.4 and 12.0% of their total variations across forests, respectively. Leaf N area, P area and LMA increased with H R in all of the studied forests, with slopes that were steeper in evergreen forests than in deciduous ones. Leaf N mass and P mass showed no significant relationship with H R generally. The increase in leaf N area, P area and LMA with H R across species is assumed to maximize community photosynthesis and may favor species with larger H R .     

Variations in dung beetle assemblages across a gradient of hunting in a tropical forest

Populations of large mammals are severely depleted by hunting in tropical forests, with direct effects on plant regeneration. But indirect consequences on commensal taxa depending on them for food resources, like coprophagous beetles, are less documented. Cascading effects of species loss across Scarabaeinae are expected, with likely significant negative implications for ecosystem functions. We examined dung beetle assemblages using pitfall traps at three rain forest sites in French Guiana ranging from intact mammalian fauna (Nouragues) to moderate (Kaw) and heavy (Matoury) defaunation. The site with the most depauperate mammalian fauna showed significantly lower dung beetle species richness than the two other two sites, which were not different from each other. Mean abundance and biomass per trap were not different across sites whereas community composition strongly differed among sites. A positive correlation was observed between body size and the individual contribution to dissimilarity between Nouragues and Kaw. The species contributing the most to dissimilarity were large. By contrast, one medium-sized species, dominant in Matoury, contributed the most to dissimilarity between Matoury and other sites. Diurnal genera of large tunnellers showed a higher diversity and abundance in Nouragues compared to other sites, whereas a nocturnal genus showed no differences. Large rollers were more abundant in Kaw compared to other sites. None of the groups of small beetles but one were affected by defaunation. Our results suggest that loss of large mammal populations affects dung beetle assemblage structure and causes decreasing abundance or disappearance of large tunnellers species that have a major impact on several dung beetle-mediated ecological processes.     

Mechanisms driving plant functional trait variation in a tropical forest

Plant functional trait variation in tropical forests results from taxonomic differences in phylogeny and associated genetic differences, as well as, phenotypic plastic responses to the environment. Accounting for the underlying mechanisms driving plant functional trait variation is important for understanding the potential rate of change of ecosystems since trait acclimation via phenotypic plasticity is very fast compared to shifts in community composition and genetic adaptation. We here applied a statistical technique to decompose the relative roles of phenotypic plasticity, genetic adaptation, and phylogenetic constraints. We examined typically obtained plant functional traits, such as wood density, plant height, specific leaf area, leaf area, leaf thickness, leaf dry mass content, leaf nitrogen content, and leaf phosphorus content. We assumed that genetic differences in plant functional traits between species and genotypes increase with environmental heterogeneity and geographic distance, whereas trait variation due to plastic acclimation to the local environment is independent of spatial distance between sampling sites. Results suggest that most of the observed trait variation could not be explained by the measured environmental variables, thus indicating a limited potential to predict individual plant traits from commonly assessed parameters. However, we found a difference in the response of plant functional traits, such that leaf traits varied in response to canopy‐light regime and nutrient availability, whereas wood traits were related to topoedaphic factors and water availability. Our analysis furthermore revealed differences in the functional response of coexisting neotropical tree species, which suggests that endemic species with conservative ecological strategies might be especially prone to competitive exclusion under projected climate change.     

基于功能性状评价5种植物对热带珊瑚岛环境的适应性

植被新建是保护和改善热带珊瑚岛生态系统的关键环节,热带珊瑚岛极端干旱生境是影响植物存活和定居的主要限制因子之一,因此选取适生植物对热带珊瑚岛植被新建至关重要。通过测定在海南省文昌市苗圃和热带珊瑚岛上的草海桐(Scaevola taccada)、厚藤(Ipomoea pescaprae)、木麻黄(Casuarina equisetifolia)、花生(Arachis hypogaea)、椰子(Cocos nucifera)5种植物的光合/水力相关功能性状,探讨植物对热带珊瑚岛生境的适应性。研究发现:与对照(文昌苗圃)相比,热带珊瑚岛上的草海桐、厚藤、木麻黄最大光合速率(A_(max))均显著升高;除椰子外,其余4种植物的比叶面积(SLA)降低,长期水分利用效率升高(其中木麻黄和花生升高显著);5种植物的叶片碳含量(LC)均有不同程度降低。此外,厚藤、木麻黄、椰子的叶片导水率(K_(leaf))显著升高;厚藤和花生的叶片膨压丧失点(Ψ_(tlp))显著降低;厚藤和木麻黄的气孔导度(g_s)显著升高。研究结果表明草海桐、椰子、花生主要通过非气孔调节方式(提高K_(leaf),降低Ψ_(tlp)、SLA等)适应热带珊瑚岛干旱环境;而厚藤和木麻黄同时通过气孔调节(降低g_s)及非气孔调节(提高K_(leaf),降低Ψ_(tlp)、SLA等)两种方式促进植物碳同化和水分利用。综上所述,草海桐、厚藤、木麻黄具有更高的光合能力和水分利用效率,能有效协调碳同化和水分利用,表现出良好的适应能力,适合用于热带珊瑚岛的植被新建。     

Nutrient enrichment increased species richness of leaf litter fungal assemblages in a tropical forest

Microbial communities play a major role in terrestrial ecosystem functioning, but the determinates of their diversity and functional interactions are not well known. In this study, we explored leaf litter fungal diversity in a diverse Panama lowland tropical forest in which a replicated factorial N, P, K and micronutrient fertilization experiment of 40 × 40 m plots had been ongoing for nine years. We extracted DNA from leaf litter samples and used fungal‐specific amplification and a 454 pyrosequencing approach to sequence two loci, the nuclear ribosomal internal transcribed spacer (ITS) region and the nuclear ribosomal large subunit (LSU) D1 region. Using a 95% sequence similarity threshold for ITS1 spacer recovered a total of 2523 OTUs, and the number of unique ITS1 OTUs per 0.5–1.0 g leaf litter sample ranged from 55 to 177. Ascomycota were the dominant phylum among the leaf litter fungi (71% of the OTUs), followed by Basidiomycota (26% of the OTUs). In contrast to our expectations based on temperate ecosystems, long‐term addition of nutrients increased, rather than decreased, species richness relative to controls. Effect of individual nutrients was more subtle and seen primarily as changes in community compositions especially at lower taxonomic levels, rather than as significant changes in species richness. For example, plots receiving P tended to show a greater similarity in community composition compared to the other nutrient treatments, the +PK, +NK and +NPK plots appeared to be more dominated by the Nectriaceae than other treatments, and indicator species for particular nutrient combinations were identified.     

Fragmentation and pre-existing species turnover determine land-snail assemblages of tropical rain forest

Aim The main aims of the study were: (1) to investigate the effect of fragment age in relation to other patch‐ and landscape‐scale measures of forest fragmentation, and (2) to assess the relative importance of fragmentation, habitat degradation (i.e. degradation caused by selective logging and past shifting cultivation) and putative pre‐existing species turnover in structuring current land‐snail assemblages. Location South‐western Sri Lanka. Methods The land‐snail fauna was sampled using standardized belt transects. Fifty‐seven transects were sampled in 21 lowland rain forest fragments (c. 1–33,000 ha). The spatial arrangement of fragments in the study area was explicitly considered in an effort to take into account the non‐random nature of fragmentation and degradation and the possibility that current species composition may reflect patterns of species turnover that existed prior to fragmentation. The data set of 57 land‐snail species and 28 environmental and spatial variables was analysed using canonical correspondence analysis and partial canonical correspondence analysis. Results Fragment age, mean shape complexity (i.e. a landscape‐scale measure of shape complexity), altitude, and the spatial variables x (longitude), y (latitude) and y² explained significant variation in land‐snail species composition. None of the three nominal variables quantifying habitat degradation was significantly correlated with variation in species composition. The independent effects of fragment age and mean shape complexity were similar. The combined effect of the spatial variables alone was larger than the independent effects of fragment age, mean shape complexity or altitude, but was of the same order of magnitude. The total variation explained by the spatial variables was comparable to the total non‐spatial variation accounted for by fragment age, mean shape complexity and altitude. Main conclusions Fragment age was found to be one of only two key determinants (the other was shape complexity at the landscape scale) driving fragmentation‐related changes in community composition. The influence of pre‐fragmentation patterns of species turnover on assemblage structure can be stronger than the effects of fragmentation measures, such as age, and may override the effects of forest degradation. Thus, strong patterns of pre‐existing turnover may potentially confound interpretation of the effects of forest fragmentation and degradation.     

Similarity of plant functional traits and aggregation pattern in a subtropical forest

The distribution of species and communities in relation to environmental heterogeneity is a central focus in ecology. Co‐occurrence of species with similar functional traits is an indication that communities are determined in part by environmental filters. However, few studies have been designed to test how functional traits are selectively filtered by environmental conditions at local scales. Exploring the relationship between soil characteristics and plant traits is a step toward understanding the filtering hypothesis in determining plant distribution at local scale. Toward this end, we mapped all individual trees (diameter >1 cm) in a one‐ha subtropical forest of China in 2007 and 2015. We measured topographic and detailed soil properties within the field site, as well as plant leaf functional traits and demographic rates of the seven most common tree species. A second one‐ha study plot was established in 2015, to test and validate the general patterns that were drawn from first plot. We found that variation in species distribution at local scale can be explained by soil heterogeneity and plant functional traits. (From first plot). (1) Species dominant in habitats with high soil ammonium nitrogen and total phosphorus tended to have high specific leaf area (SLA) and relative growth rate (RGR). (2) Species dominant in low‐fertility habitats tended to have high leaf dry matter content (LDMC), ratio of chlorophyll a and b (ratioab), and leaf thickness (LT). The hypothesis that functional traits are selected in part by environmental filters and determine plant distribution at local scale was confirmed by the data of the first plot and a second regional site showed similar species distribution patterns.     

重庆市6种常见园林植物功能性状对城乡生境梯度的响应

功能性状作为联系植物、环境与生态系统的桥梁,对研究植物适应不同生境的策略意义重大。本研究以重庆主城区中华蚊母树(Distylium chinense)、鸡爪槭(Acer palmatum)、檵木(Loropetalum chinense)、构树(Broussonetia papyrifera)、银杏(Ginkgo biloba)、玉兰(Magnolia denudata)为对象,遴选植物功能性状及生物量分配指标,定量分析了其在不同城市环境下的差异性,旨在考查植物对于城市生态环境的适应策略。结果表明:除比叶面积和生物量分配比例外,其他功能性状包括营养器官和生殖器官随乡-郊-城环境梯度发生显著变化(P<0.05),表现为其数值随环境梯度显著下降;除鸡爪槭外,其余5种植物分配给繁殖器官的资源最多,分配给茎的资源最少;根据植物功能性状主成分分析,将6种植物分为3类:营养器官和生殖器官基本未受城市环境影响类,营养器官未受影响而生殖器官受损类,以及营养和生殖器官均受损类;植物功能性状综合评价显示,在森林公园中植物性状表现最好,郊区公园次之,城市公园中最差。     

Nurse species and indirect facilitation through grazing drive plant community functional traits in tropical alpine peatlands

Facilitation among plants mediated by grazers occurs when an unpalatable plant extends its protection against grazing to another plant. This type of indirect facilitation impacts species coexistence and ecosystem functioning in a large array of ecosystems worldwide. It has nonetheless generally been understudied so far in comparison with the role played by direct facilitation among plants. We aimed at providing original data on indirect facilitation at the community scale to determine the extent to which indirect facilitation mediated by grazers can shape plant communities. Such experimental data are expected to contribute to refining the conceptual framework on plant–plant–herbivore interactions in stressful environments. We set up a 2‐year grazing exclusion experiment in tropical alpine peatlands in Bolivia. Those ecosystems depend entirely on a few, structuring cushion‐forming plants (hereafter referred to as “nurse” species), in which associated plant communities develop. Fences have been set over two nurse species with different strategies to cope with grazing (direct vs. indirect defenses), which are expected to lead to different intensities of indirect facilitation for the associated communities. We collected functional traits which are known to vary according to grazing pressure (LDMC, leaf thickness, and maximum height), on both the nurse and their associated plant communities in grazed (and therefore indirect facilitation as well) and ungrazed conditions. We found that the effect of indirectly facilitated on the associated plant communities depended on the functional trait considered. Indirect facilitation decreased the effects of grazing on species relative abundance, mean LDMC, and the convergence of the maximum height distribution of the associated communities, but did not affect mean height or cover. The identity of the nurse species and grazing jointly affected the structure of the associated plant community through indirect facilitation. Our results together with the existing literature suggest that the “grazer–nurse–beneficiary” interaction module can be more complex than expected when evaluated in the field.     

Earthworms promote greater richness and abundance in the emergence of plant species across a grassland-forest ecotone

Aims Chalk grasslands are subject to vegetation dynamics that range from species-rich open grasslands to tall and encroached grasslands, and woods and forests. In grasslands, earthworms impact plant communities and ecosystem functioning through the modification of soil physical, chemical and microbiological properties, but also through their selective ingestion and vertical transportation of seeds from the soil seed bank. Laboratory experiments showed that seed–earthworm interactions are species specific, but little is known on the impact of seed–earthworm interactions in the field. The overall aim of this study was to better understand seed–earthworm interactions and their impact on the plant community. First we analyzed the composition of seedlings emerging from casts after earthworm ingestion. Then we compared seedling composition in casts to the plant composition of emerging seedlings from the soil and of the aboveground vegetation along four stages of the secondary succession of chalk grasslands.Methods Four stages of the secondary succession of a chalk grassland—from open sward to woods—were sampled in Upper Normandy, France, in February 2010. Within each successional stage (×3 replicates), we sampled the standing vegetation, soil seed bank at three soil depths (0–2, 2–5 and 5–10cm) and earthworm surface casts along transects. Soil and cast samples were water sieved before samples were spread onto trays and placed into a greenhouse. Emerging seedlings were counted and identified. Effect of successional stage and origin of samples on mean and variability of abundance and species richness of seedlings emerging from casts and soil seed banks were analyzed. Plant compositions were compared between all sample types. We used generalized mixed-effect models and a distance-based redundancy multivariate analysis.Important findings Seedling abundance was always higher in earthworm casts than in the soil seed bank and increased up to 5-fold, 4-fold and 3.5-fold, respectively, in the tall grassland, woods and encroached grassland compared to the soil surface layer. Species richness was also higher in earthworm casts than in the soil seed bank in all successional stages, with a 4-fold increase in the encroached grassland. The plant composition of the standing vegetation was more similar to that of seedlings from casts than to that of seedlings from the soil seed bank. Seedlings diversity emerging from casts in the tall and encroached grasslands tended toward the diversity found in woods. Our results indicate that earthworms may promote the emergence of seedlings. We also suggest that the loss of some plant species in the seed bank and the tall grass vegetation in intermediary successional stages modify the local conditions and prevent the further establishment of early-successional plant species.     

Modelling the effect of temperature changes on plant life-form distribution across a treeline ecotone in the tropical Andes

ABSTRACT

Background: Classifying species from the high tropical Andes into plant life forms (PLF) can provide a more functional understanding of tropical treeline dynamics. However, little is known about the potential response of PLFs to climate warming.

Aims: The objective of this work was to evaluate the response of PLFs to environmental conditions above the upper montane forest in the Venezuelan Andes and forecast their potential distribution under warming scenarios using habitat suitability models.

Methods: We classified species into nine PLFs, analysed their current distribution using CCA and non-linear multiple regressions and forecasted their potential distribution under warming scenarios. We used anatomical leaf traits for shrubs and caulescent rosettes to refine their potential responses to climate.

Results: Tree cover sharply decreased with increasing elevation, while shrub and caulescent rosette distribution differed depending on their leaf traits. Projections suggested an upslope shift of all PLFs with warming and an increase in tree cover.

Conclusions: The analysis of the distribution of PLFs, by linking plant adaptive strategies to topography/climate, allowed projecting changes in vegetation physiognomy in response to warming. Projections of a moderate increase in tree cover in the grassland páramo did not indicate the formation of closed forests above the forest line.     

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。     

A combination of functionally different plant traits provides a means to quantitatively predict a broad range of species assemblages in NW Europe

Assembly theory predicts that filtering processes will select species by their attributes to build a community. Some filters increase functional similarity among species, while others lead to dissimilarity. Assuming converging processes to be dominant within habitats, we tested in this study whether species assemblages across a wide range of habitats can be distinguished quantitatively by their mean trait compositions. In addition, we investigated how many and which traits are needed to describe the differences between species assemblages best. The approach has been applied on a dataset that included 12 plant traits and 7644 vegetation relevés covering a wide range of habitats in the Netherlands. We demonstrate that due to the dominant role of converging processes 1) the functional composition can explain up to 80% of the floristic differences between species assemblages using seven plant traits, showing that plant trait combinations provide a powerful tool for predicting the occurrence of species assemblages across different habitats; 2) to achieve a high performance, traits should be taken from different strategy components, i.e. traits that are functionally orthogonal, which does not necessarily coincide with low trait‐trait correlations; 3) the different strategy components identified in this study correspond to the strategy components of some conventional plant ecological strategy schemes (PESS) – schemes to describe the variation between individual species. However, some PESS merge traits into one strategy component that are shown to be functionally different when predicting species assemblages. If such PESS is used to predict assemblages, this leads to a loss in predictive capacity. Potentially, our new approach is globally applicable to quantify community assembly patterns. However this needs to be tested.