Low resource availability limits weed invasion of tropical savannas

The savanna biome is one of the least invaded among global biomes, although the mechanisms underpinning its resistance to alien species relative to other biomes is not well understood. Invaders generally are at the resource acquisitive end of functional global plant trait variation and in low-resource savanna environments we might expect that successful invaders will only outperform native species under resource rich or highly disturbed conditions. However, invaders may also directly exploit resource stressed environments using resource conservative traits in some situations. It’s also possible that successful invaders and native species largely overlap in their trait profiles indicating site specific environmental factors are responsible for invader success in particular contexts rather than a general trait and functional divergence between invaders and native species. To address these various hypotheses, we compared a suite of morphological and physiological traits in graminoid and herbaceous native and co-occurring invasive plant species across a range of habitats in savannas of the Kimberley region of northern Australia. Invader grass species had traits associated with resource acquisition and fast growth rates, such as high SLA and leaf nutrient contents. In contrast, dominant native perennial grasses had traits characteristic of resource conservation and slow growth in resource stressed conditions. Trait profiles among invasive forbs and legumes exhibited stress tolerant traits relative to their native counterparts. Invaders also displayed strong divergence in reproductive traits, suggesting diverse responses to disturbance not indicated by leaf economic traits alone. These results suggest that savannas may be resistant to invaders with resource acquisitive traits due to their strong resource limitation.     

Zoogeography of a South African Province: A framework for management

State‐level conservation in South Africa is structured around distinct political entities (i.e. municipalities). This is problematic because an ecological approach that considers species distribution is required to delineate meaningful management units. To do so, vegetation types can be used as management units—however, it is uncertain whether vertebrate communities are associated with vegetation types as defined by the national vegetation map. Here, we investigate mammal diversity patterns within and among biomes (savannah and grassland) and bioregions and test whether different mammal communities were associated with different vegetation types. We used an extensive database of species occurrences in the North West Province. We found that species richness was higher in the savannah than grassland biome. Beta diversity was higher within the savannah than grassland biome, due to greater environmental heterogeneity, though one grassland bioregion was similar to the savannah bioregions. Mammal communities were significantly different among bioregions, but not biomes, suggesting mammal communities are congruent with vegetation type at finer scales (i.e. bioregional), but not at coarser scales (biomes). It thus makes sense to use a bioregional framework to design mammal management strategies. The invasion of grasslands by savannah species should be monitored, specifically given the predicted changes in climate.     

Does seasonality drive spatial patterns in demography? Variation in survival in African reed warblers Acrocephalus baeticatus across southern Africa does not reflect global patterns

Among birds, northern temperate species generally have larger clutches, shorter development periods and lower adult survival than similarly‐sized southern and tropical species. Even though this global pattern is well accepted, the driving mechanism is still not fully understood. The main theories are founded on the differing environmental seasonality of these zones (higher seasonality in the North). These patterns arise in cross‐species comparisons, but we hypothesized that the same patterns should arise among populations within a species if different types of seasonality select for different life histories. Few studies have examined this. We estimated survival of an azonal habitat specialist, the African reed warbler, across the environmentally diverse African subcontinent, and related survival to latitude and to the seasonality of the different environments of their breeding habitats. Data (1998–2010) collected through a public ringing scheme were analyzed with hierarchical capture‐mark‐recapture models to determine resident adult survival and its spatial variance across sixteen vegetation units spread across four biomes. The models were defined as state‐space multi‐state models to account for transience and implemented in a Bayesian framework. We did not find a latitudinal trend in survival or a clear link between seasonality and survival. Spatial variation in survival was substantial across the sixteen sites (spatial standard deviation of the logit mean survival: 0.70, 95% credible interval (CRI): 0.33–1.27). Mean site survival ranged from 0.49 (95% CRI: 0.18–0.80) to 0.83 (95% CRI: 0.62–0.97) with an overall mean of 0.67 (95% CRI: 0.47–0.85). A hierarchical modeling approach enabled us to estimate spatial variation in survival of the African reed warbler across the African subcontinent from sparse data. Although we could not confirm the global pattern of higher survival in less seasonal environments, our findings from a poorly studied region contribute to the study of life‐history strategies.     

GIRAE: a generalised approach for linking the total impact of invasion to species' range,abundance and per-unit effects

The total impact of an alien species was conceptualised as the product of its range size, local abundance and per-unit effect in a seminal paper by Parker et al. (Biol Invasions 1:3–19, 1999). However, a practical approach for estimating the three components has been lacking. Here, we generalise the impact formula and, through use of regression models, estimate the relationship between the three components of impact, an approach we term GIRAE (Generalised Impact?=?Range size × Abundance × per-unit Effect). We discuss how GIRAE can be applied to multiple types of impact, including environmental impacts, damage and management costs. We propose two methods for applying GIRAE. The species-specific method computes the relationship between impact, range size, abundance and per-unit effect for a given species across multiple invaded sites or regions of different sizes. The multi-species method combines data from multiple species across multiple sites or regions to calculate a per-unit effect for each species and is computed using a single regression model. The species-specific method is more accurate, but it requires a large amount of data for each species and assumes a constant per-unit effect for a species across the invaded area. The multi-species method is more easily applicable and data-parsimonious, but assumes the same relationship between impact, range size and abundance for all considered species. We illustrate these methods using data about money spent managing plant invasions in different biomes of South Africa. We found clear differences between species in terms of money spent per unit area invaded, with per-unit expenditure varying substantially between biomes for some species—insights that are useful for monitoring and evaluating management. GIRAE offers a versatile and practical method that can be applied to many different types of data to better understand and manage the impacts of biological invasions.

     

Using auto covariance method for functional discrimination of membrane proteins based on evolution information

Membrane transporters are critical in living cells. Therefore, the discrimination of the types of membrane proteins based on their functions is of great importance both for helping genome annotation and providing a supplementary role to experimental researchers to gain insight into membrane proteins’ function. There are a lot of computational methods to facilitate the identification of the functional types of membrane proteins. However, in these methods, the local sequence environment was not integrated into the constructed model. In this study, we described a new strategy to predict the functional types of membrane proteins using a model based on auto covariance and position-specific scoring matrix. The novelty of the presented approach is considering the distribution of different positions of functional conservation sites in protein sequences. Thereby, this model adequately takes into account the long-range correlation between such sites during sequential evolution. Fivefold cross-validation test shows that this method greatly improves the prediction accuracy and achieves an acceptable prediction accuracy of 87.51%. The result indicates that the current approach might be an effective tool for predicting the functional types of membrane proteins only using the primary sequences. The code and dataset used in this article are freely available at .     

高海拔和高纬度生境的相似性:综述以及青藏高原鸟类肾上腺皮质激素对应激反应的初步研究

在介绍性教科书、保护计划甚至基础著作中 ,常常将北极区和高山生物群系等同看待。高山和北极区生物群系无树的广阔区域 ,草本和非禾本科草本植物的初级生产力总量低 ,食物网单一。此外 ,除热带高山生境可能有例外 ,该两种环境都具有明显的季节性。但是 ,对于这些表面上相似的环境以前很少进行比较 ,而两者之间应具有明显的不同。我们假定 :如果环境相似 ,栖息在这些环境里的鸟类会对意外的应激刺激表现出相似的反应。相反 ,如果这些环境很不同 ,那么鸟类对剧烈应激的反应可进行调整以适应当地的不同环境 ,而不应一成不变。本文初步报道了青藏高原某些繁殖鸟类的肾上腺皮质反应 ,并与已发表的栖息于北极区苔原生态系统鸟类的数据进行了比较。白腰雪雀 (Onychostruthustaczanowskii)和棕颈雪雀 (Pyrgilaudaruficollis)被认为是高山代表物种 ,随着标准化捕捉 -处理 -抑制的应激刺激 ,它们表现出典型的血浆肾上腺皮质酮含量升高。但是 ,不同于许多北极区的代表物种 ,它们对应激的肾上腺皮质反应并无任何季节性变化。研究结果表示 :生活在高山和北极区环境下的鸟类物种可能具有激素反应以适应其生存环境所施加的生态挑战 [动物学报 49(1) :1～ 19,2 0 0 3]。     

Microscale methods to assemble mammalian cells into tissue-like structures

Different cell types make up tissues and organs hierarchically and communicate within a complex, three-dimensional (3D) environment. The in vitro recapitulation of tissue-like structures is meaningful, not only for fundamental cell biology research, but also for tissue engineering (TE). Currently, TE research adopts either the top-down or bottom-up approach. The top-down approach involves defining the macroscopic tissue features using biomaterial scaffolds and seeding cells into these scaffolds. Conversely, the bottom-up approach aims at crafting small tissue building blocks with precision-engineered structural and functional microscale features, using physical and/or chemical approaches. The bottom-up strategy takes advantage of the repeating structural and functional units that facilitate cell-cell interactions and cultures multiple cells together as a functional unit of tissue. In this review, we focus on currently available microscale methods that can control mammalian cells to assemble into 3D tissue-like structures.     

Forest vegetation across the tropical Pacific: A biogeographically complex region with many analogous environments

The tropical realm of the Pacific contains many islands with closely similar (analogous) environmental settings. Due to the `filter effect' of the ocean, these are occupied by historically different species assemblages. This results in a unique biogeographic complexity not found in any of the continental tropical regions. The paper presents a hierarchical approach to the study of vegetation in analogous environments with data illustrated by a series of diagrams. It begins with differentiating island types and climatic zones. Then it focuses on Pacific-wide biomes, thereafter on island landscape profiles, and finally, on stand-level profiles as a method to compare forest structure and composition among islands in analogous environments at the scale of relevés. The conclusions emphasize the urgent need for intensifying conservation-oriented research and capacity building in all island countries of the Pacific (not only Hawaii) to protect their indigenous biodiversities for sustainable uses and the health of their ecosystems. Attention is also drawn to the scientific advantage of using the Pacific-Asia Biodiversity Transect (PABITRA) Network, since it connects the island areas across biogeographic boundaries in form of an experimental design. This design refers to analogous island environments filled with different sets of biodiversities, which follow a trend of impoverishment with indigenous founder species from west to east.     

Global to community scale differences in the prevalence of convergent over divergent leaf trait distributions in plant assemblages

Aim The drivers of species assembly, by limiting the possible range of functional trait values, can lead to either convergent or divergent distributions of traits in realized assemblages. Here, to evaluate the strengths of these species assembly drivers, we partition trait variance across global, regional and community scales. We then test the hypothesis that, from global to community scales, the outcome of co‐occurring trait convergence and divergence is highly variable across biomes and communities. Location Global: nine biomes ranging from subarctic highland to tropical rain forest. Methods We analysed functional trait diversity at progressively finer spatial scales using a global, balanced, hierarchically structured dataset from 9 biomes, 58 communities and 652 species. Analyses were based on two key leaf traits (foliar nitrogen content and specific leaf area) that are known to drive biogeochemical cycling. Results While 35% of the global variance in these traits was between biomes, only 15% was between communities within biomes and as much as 50% occurred within communities. Despite this relatively high within‐community variance in trait values, we found that trait convergence dominated over divergence at both global and regional scales through comparisons of functional trait diversity in regional and community assemblages against random (null) models of species assembly. Main conclusions We demonstrate that the convergence of traits occurring from global to regional assemblages can be twice as strong as that from regional to community assemblages, and argue that large differences in the nature and strength of abiotic and biotic drivers of dominant species assembly can, at least partly, explain the variable outcome of simultaneous trait convergence and divergence across sites. Ultimately, these findings stress the urgent need to extend species assembly research to address those scales where trait variance is the highest, i.e. between biomes and within communities.     

荒漠植物不同功能群性状特征及其与土壤环境的关系

研究不同功能群植物性状差异及其与土壤环境关系对于充分掌握植物的环境适应策略至关重要。以艾比湖流域为研究区,利用荒漠植物的植物高度、叶片碳、氮、磷、硫、钾、钙、钠、镁含量等9个性状,将高、低土壤水盐环境下的植物划分为5个功能群,分析不同功能群的植物组成、性状差异及其与土壤环境的关系。结果表明：(1)不同土壤水盐环境下,其植物功能群组成不同;其中白刺、胡杨和罗布麻在两个土壤水盐环境下的功能群中均存在。(2)植物的功能性状在不同土壤水盐环境下也发生了适应性的变化。高土壤水盐环境下3个功能群的植物高度、叶片碳、氮、磷和钙含量显著高于低土壤水盐环境功能群(P<0.05);低土壤水盐环境下2个功能群的植物叶片硫、钠和镁含量高于高土壤水盐环境功能群。(3)土壤含水量(SVWC)、电导率(EC)、pH以及土壤磷含量对荒漠植物功能性状影响较大。在高土壤水盐环境下,EC、pH与植物高度,叶片钾、钙含量正相关,与叶片硫含量负相关;在低土壤水盐环境下,SVWC、EC与植物高度呈显著正相关(P<0.05)。研究有助于理解荒漠植物对极端环境的适应对策,为保护荒漠地区生物多样性提供理论依据。     

Taxonomic and functional β-diversity of ants along tree plantation chronosequences differ between contrasting biomes

The replacement of natural by anthropogenic habitat changes biological communities in any biome. Variations in environmental conditions along the chronosequence of tree plantations may act as a gradient of environmental filtering where the gain or loss of species occurs. It is expected that environmental filtering increases with the decrease in environmental similarity between the plantations and the natural habitat. Young tree plantations are structurally more similar to grasslands than to mature plantations, which in turn are structurally more similar to subtropical forest than young plantations. This study compares patterns of beta diversity across exotic pine plantation chronosequences in contrasting biomes. We predict that taxonomic and functional beta diversity between plantation and the natural habitat assemblages increase with plantation age in grasslands and decrease in the subtropical forest. We sampled epigean ants and measured environmental variables at 54 plantations of different ages and natural habitats in grassland and forest biomes in Argentina. Taxonomic and functional beta diversity between natural habitat and pine plantations were estimated through dissimilarity indexes of turnover and nestedness. To assess the response of beta diversity estimators to plantation age we performed general linear and non-linear models. Results revealed opposite beta diversity patterns between biomes along the plantation cycle. Turnover increased and nestedness decreased with declining environmental similarity between pine plantations and the natural habitats; changes in the identity of the species were coupled to changes in their functional characteristics. Thus, a given environmental gradient may produce different diversity patterns depending on the regional species pool. Forestry practices that generate environmental conditions similar to natural environments could help to conserve species from the natural habitat.     

Can ants be used as ecological indicators of restoration progress in dynamic environments? A case study in a revegetated riparian zone

Ant assemblages are focal ecological indicators of progress in mine-site restoration, often showing increasing species richness with restoration age. Certain functional groups also behave in predictable ways in response to disturbance and changes in the environment. Whether these ant responses can be applied to other types of restoration and ecosystems is unknown, especially in dynamic environments and where gradients may not be as severe as in mine-site restoration. Ant assemblages would be expected to perform poorly as ecological indicators in dynamic environments because such environs are subject to periodic disturbance of important habitat features. Indeed, periodic disturbance may limit the predictive power of any ecological indicator. In this study, we trapped ants on two separate occasions to compare ant assemblages among four riparian habitat types (Unplanted grassland, Young revegetation, Older revegetation and Mature woodland). These habitat types were assumed to represent progressive stages of restoration. In contrast to the findings of others, species richness was variable among replicate locations of the same habitat type, and did not differ among the four habitat types. Also in contrast to what others have found for functional groups, dolichoderines were equally abundant in all habitat types and did not decrease in abundance with vegetation maturity. While generalized myrmicines and opportunists became more common with maturation of the vegetation, they did not replace dolichoderines as the most common ants. Surprisingly, the relative abundance of Subordinate Camponotini, a functional group considered to be of limited use in discriminating structural types, increased across the restoration gradient. There were also fairly distinct species assemblages associated with unplanted grassland and mature woodland. Communities in revegetated habitats were intermediate of these extremes, suggesting there is a level of predictiveness to their response to revegetation in this system. While species richness and a functional group approach would be of little use in this environment, species composition would provide a useful gauge of restoration progress. Ant species richness and functional group metrics have repeatedly been advocated as ecological indicators. Given our results, we caution against the blind application of metrics that have not been validated in the context in which they are to be applied.     

Functional Roles of Centridini Oil Bees and Malpighiaceae Oil Flowers in Biome‐wide Pollination Networks

Interactions between oil‐collecting bees and oil‐producing flowers are a very specialized mutualism, whose natural history is well known at the organism and population levels. In this study, we assessed these interactions at the biome level with a network approach, and hypothesized that widespread bee and plant species would occupy different ecological functional roles (Eltonian niches) in different biomes. Furthermore, we expected the most important functional roles in each network to be occupied more frequently by Byrsonima oil flowers and Centris oil bees, which share the longest coevolutionary history in the Neotropics. By compiling data from 40 articles on oil flower interactions within the Malpighiaceae family, we built six networks for different Brazilian biomes. We assessed the ecological functional role of each species in pollination networks of oil flowers through the metric known as ‘network functional role’. Although 90 percent of the species occupied peripheral roles in each network, some were found to occupy highly central roles. Oil flowers of the genera Byrsonima and Banisteriopsis and oil bees of the genera Centris and Epicharis were the most important species in all networks, as they made a disproportionally high number of interactions (hubs), or helped bind together different modules (connectors). Our findings suggest that functional roles vary geographically and seem to be affected by local conditions in different biomes. Furthermore, coevolutionary history seems to play an important role in determining functional roles in oil flower networks, although other factors are probably also important, especially the degree of specialization in this kind of interaction.     

Integrated and independent evolution of heteromorphic sperm types

Sperm are a simple cell type with few components, yet they exhibit tremendous between-species morphological variation in those components thought to reflect selection in different fertilization environments. However, within a species, sperm components are expected to be selected to be functionally integrated for optimal fertilization of eggs. Here, we take advantage of within-species variation in sperm form and function to test whether sperm components are functionally and genetically integrated both within and between sperm morphologies using a quantitative genetics approach. Drosophila pseudoobscura males produce two sperm types with different functions but which positively interact together in the same fertilization environment; the long eusperm fertilizes eggs and the short parasperm appear to protect eusperm from a hostile female reproductive tract. Our analysis found that all sperm traits were heritable, but short sperm components exhibited evolvabilities 10 times that of long sperm components. Genetic correlations indicated functional integration within, but not between, sperm morphs. These results suggest that sperm, despite sharing a common developmental process, can become developmentally and functionally non-integrated, evolving into separate modules with the potential for rapid and independent responses to selection.     

Variation in plant responses to neighbors at local and regional scales

Differences in the responses of plant species to neighbors may determine their distribution among contrasting environments, but no studies have compared variation in competitive or facilitative abilities both within and among environments. We determined whether the responses of plants to interspecific interactions varied at large scales (between environments) and small scales (among sites within an environment) across a tree line. We separated the effects of above- and belowground interactions on seedlings of grasses and trees grown in prairie or forest using vegetation removals at several sites. Species interactions generally had no significant effect on transplant survival. Competition reduced seedling growth by about 33%-89% in both prairie and forest environments. Despite the strong suppression of growth by neighbors, environment and species effects contributed more to variation in transplant performance than did neighbor removals. Responses to neighbors varied among transplant species but generally did not vary significantly between environments or among sites. With vegetation removed, grasses grew significantly faster in prairie and trees grew faster in forest. Thus, in the absence of neighbors, species showed distinct preferences for the environment in which they are most abundant. In summary, the responses of grasses and woody species to neighbors did not vary significantly at either large (between environments) or small (among sites) scales. These results suggest that species responses to interspecific interactions do not vary strongly with environment or smaller-scale site effects.     

Guilds or functional groups: does it matter?

Although most researchers use the terms "guild" and "functional group" more or less synonymously, these two concepts bear different meanings. The guild concept refers primarily to the mechanisms of resource sharing by species in a competitive context whereas the functional groups concept is concerned with how a resource or any other ecological component is processed by different species to provide a specific ecosystem service or function. In many cases but not necessarily all, the two concepts are the two "faces" or "sides" of the same coin: the sharing by species of a similar resource is the guild facet (structural), while the ecosystem processes these species eventually perform through resource exploitation is the functional group facet. The two concepts differ in that competitive relationships within groups of species are not the focus of the functional group approach, exactly as processes or functions are not the focus of the guild approach. A group of species can be considered either as a guild or a functional group depending on the question addressed. Guild and functional group membership is independent of phylogenetic relationships but because species tend to share similar life history traits and adaptations through common evolutionary history, guild and functional group associates are often closely related. The concept of guild has had broader application in animal studies than in plant studies, whereas the reverse is true for the concept of functional group. Recent methodological advances to objectively partition species into guilds and functional groups, taking into consideration the most relevant characters or traits for delineating them, provide the means to construct an operational framework for making in situ and ex situ experiments that are urgently needed for a better understanding of the role of species in ecosystem functioning, especially in relation to global change concerns.     

The phylogeny and biogeography of Hakea (Proteaceae) reveals the role of biome shifts in a continental plant radiation

The frequency of evolutionary biome shifts during diversification has important implications for our ability to explain geographic patterns of plant diversity. Recent studies present several examples of biome shifts, but whether frequencies of biome shifts closely reflect geographic proximity or environmental similarity of biomes remains poorly known. We explore this question by using phylogenomic methods to estimate the phylogeny of Hakea, a diverse Australian genus occupying a wide range of biomes. Model‐based estimation of ancestral regions indicates that Hakea began diversifying in the Mediterranean biome of southern Australia in the Middle Eocene–Early Oligocene, and dispersed repeatedly into other biomes across the continent. We infer around 47 shifts between biomes. Frequencies of shifts between pairs of biomes are usually similar to those expected from their geographic connectedness or climatic similarity, but in some cases are substantially higher or lower than expected, perhaps reflecting how readily key physiological traits can be modified to adapt lineages to new environments. The history of frequent biome‐shifting is reflected in the structure of present‐day assemblages, which tend to be more phylogenetically diverse than null‐model expectations. The case of Hakea demonstrates that the radiation of large plant clades across wide geographic areas need not be constrained by dispersal limitation or conserved adaptations to particular environments.     

The cultural transmission of acquired variation: Effects on genetic fitness

In a cultural species individuals acquire some aspects of their phenotypes by imitating conspecifics. Presumably, cultural transmission has some adaptive function in such species. Some authors have suggested that one such function may be the ability to transmit phenotypes that have been modified by learning or some other form of phenotypic plasticity. To examine this hypothesis we have compared the genetic fitness of cultural and noncultural individuals in three different models. In each model the environment is assumed to be variable, either in space or time, and learning modifies each individual's phenotype so as to increase the individual's fitness in the local environment. Cultural individuals transmit the modified phenotype to their offspring, while noncultural individuals do not. The models are different in the following ways: two models assume a dichotomous phenotype, one in a temporally varying environment and one in a spatially varying environment; the third model assumes a quantitative phenotype in a temporally variable environment. The two models which assume a temporally varying environment yield similar results. Genetic transmission is favored in highly autocorrelated environments while cultural transmission in environments with intermediate autocorrelation. In the spatially varying model cultural transmission is always favored.     

Does size matter? Examining the drivers of mammalian vocalizations

Previous studies of the vocalization frequencies of mammals have suggested that it is either body mass or environment that drives these frequencies. Using 193 species across the globe from the terrestrial and aquatic environments and a model selection approach, we identified that the best‐supported model for minimum and maximum frequencies for vocalization included both body mass and environment. The minimum frequencies of vocalizations of species from all environments retained the influence of body mass. For maximum frequency however, aquatic species are released from such a trend with body mass having little constraint on frequencies. Surprisingly, phylogeny did not have a strong impact on the evolution of the maximum frequency of mammal vocalizations, largely due to the pinniped species divergence of frequency from their carnivoran relatives. We demonstrate that the divergence of signal frequencies in mammals has arisen from the need to adapt to their environment.     

Clonal vs leaf-height-seed (LHS) traits: which are filtered more strongly across habitats?

Plant functional traits are now frequently used instead of species identity to identify how plant species co-exist in assemblages. One notion is that species inhabiting the same environment have more characteristics in common than species from different habitats, leading to different prevailing dominant traits along environmental gradients, and also to a lesser diversity of traits in habitats that impose a stronger filter on these traits. Though such patterns have been demonstrated for different environmental drivers and different traits, studies using easily available traits connected to above ground processes (i.e. traits of the leaf-height-seed, or LHS, strategy scheme) are largely overrepresented in these analyses. Here we combined data on clonal and bud bank traits, representing the ability to reproduce and spread vegetatively, with LHS trait data and examined how these traits varied in relation to the vegetational composition of 29 Central-European habitat types. Our analysis focused on determining whether clonal/bud bank or LHS traits play an important role for environmental filtering along gradients approximated by Ellenberg indicator values (EIV) across these habitats. Our results show that clonal and bud bank traits are at least as – if not more – important for the differentiation of the 29 habitat types. Overall, diversity and dominance of clonal and bud bank traits was more strongly correlated with gradients of light availability, temperature, moisture, soil reaction, and nutrient availability across these habitats than it was the case for traits of the leaf-height-seed scheme. Our results call for a stronger integration of belowground traits into the functional traits approach in plant ecology and for an extension of efforts to collect such data.