Plant response strategies to stress and disturbance: the case of aquatic plants

The environmental factors controlling the establishment and development of plants in different ecosystems are of two types, stress and disturbance. The effects of stress or disturbance on aquatic systems are discussed in relation to the following questions:Can we predict the state and rate of recolonization after a disturbance? What are the strategies of recolonization developed by plants? How high is the resilience of a disturbed system? Two theories, the intermediate disturbance hypothesis, and the patch dynamics concept proposed to predict the composition, structure and dynamics of plants due to physical-chemical factors, were tested on two scales, that of communities and that of species, within two alluvial floodplains (the Rhine and the Rhône systems in France).With regard to the change of community on a larger scale (i.e. the whole network of the cut-off channels in the floodplain), large gradients of connection and disturbance induce high diversities within communities. Moreover, the highest flood disturbance induces a higher species richness and the occurrence of a particular species. The change in species is analysed using biological traits (morphological, reproductive or physiological). In the floodplain of the river Rhône, the response of plants corresponds well to theory, i.e. that habitats with an intermediate disturbance are richer than more or less disturbed habitats. So we can predict, through the biological traits, the functioning of a habitat. The last remaining question is that of the resilience of the system, which can be discussed in terms of species competition and the risk of biological invasion after an opening of habitat.     

Traits of benthic macroinvertebrates in semi-natural French streams: an initial application to biomonitoring in Europe

• 1 The methods used to indicate the biological state of streams are often based on taxonomic composition, and the abundance of species or other taxa. This ‘taxonomic structure’ varies among ecoregions and cannot be applied to wider geographical areas. Therefore, we assessed the species traits of benthic macroinvertebrates from semi‐natural reference sites as a potential benchmark for large‐scale biomonitoring. Our purpose was to assess the stability of community structure, based on the representation of taxa and of traits, across large gradients of geology (sedimentary to granitic), altitude (65–1982 m), geographical coordinates (0° 48′ W to 7° 20′ E and 42° 52′ to 48° 44′ N), stream order (1–5) and slope (0.5–60‰).
• 2 We used invertebrate abundance data from the 62 most natural French stream sites available. These abundance data served to weight the occurrence of ‘biological’ traits, such as reproductive characteristics, mobility, resistance forms, food, feeding habits, respiration, and ‘ecological’ traits, such as preferences for temperature, trophic level, saprobity, biogeographic distribution, longitudinal zonation, substratum and current velocity.
• 3 Multivariate analyses of taxonomic composition demonstrated a clear site gradient from lowlands to uplands and from calcareous to granitic geology. In contrast, community structure based on both biological and ecological traits was stable across environmental gradients.
• 4 The frequency distribution of biological traits indicated that the stream benthos of the ‘reference sites’ had a mixture of categories which confirmed theoretical predictions for temporally stable and spatially variable habitats. A mixture of ecological trait categories also occurred at our reference sites. Thus, semi‐natural benthic macroinvertebrate communities are functionally diverse. Moreover, we included an initial application of these traits to a case of slightly to moderately polluted sites to show that the impact of humans significantly changes this natural functional diversity.
• 5 Future studies should focus on the potential for various biological and ecological traits to discriminate different human impacts on the benthic macroinvertebrates of running waters, and on the integration of this functional application into a general ‘reference‐condition’ approach.

     

Genetic analysis of potential postglacial watershed crossings in Central Europe by the bullhead (Cottus gobio L.)

Natural colonizations across watersheds have been frequently proposed to explain the present distributions of many freshwater fish species. However, detailed studies of such potential watershed crossings are still missing. Here, we investigated potential postglacial watershed crossings of the widely distributed European bullhead (Cottus gobio L.) in two different areas along the Rhine–Rhône watershed using detailed genetic analysis. The main advantage of studying bullheads vs. other freshwater fish species is that their distribution has been lightly influenced by human activities and as such, interpretations of colonization history are not confounded by artificial transplantations. The genetic analyses of eight microsatellite loci revealed strong genetic similarities between populations of both sides of the Rhine–Rhône watershed in the Lake Geneva area, giving strong evidence for a natural watershed crossing of bullheads from the upper Rhine drainage into the Rhône drainage in the Lake Geneva area likely facilitated by the retreat of the glaciers after the last glacial maximum some 20 000 years ago. Populations from the Lake Geneva basin were genetically more similar to populations from across the watershed in the upper Rhine drainage than to populations further downstream in the lower Rhône. In contrast, populations from Belfort, an area, which was not covered by ice during the last glacial maximum, showed strong genetic differentiation between populations of the upper Rhine and Rhône drainages. Based on our results on the bullhead, we propose that glacial retreat may have eased the dispersal of numerous European freshwater fish species across several geological boundaries.     

Predicting community characteristics from habitat conditions: fluvial fish and hydraulics

1. One current approach to the prediction of community characteristics is to use models of key local-scale processes (e.g. niche dimensions) affecting individuals and to estimate the effects of these attributes over larger scales. We tested this approach, focusing on how the hydraulic habitat structures fluvial fish communities. 2. We used a recent statistical habitat model to predict fish community characteristics in eleven reaches in the Rhône river basin in France. Predictions were made ‘blindly’ since most reaches were not used to calibrate the model. The model reflects species preferences for local hydraulics. We made predictions of the fish community from the local hydraulic conditions found in the reaches under low flow conditions. The overall abundance and the relative abundance (both as indices) of fish species, specific size classes and species traits (i.e. reproductive, trophic, morphological and others) were predicted. We summarized our predictions of the relative abundance of species as two ‘community structure indices’ using Principal Component Analysis. 3. Our predictions from low-flow hydraulics were compared with long-term observations of fish communities. The relative abundance of species actually observed depended largely on zoogeographic factors within the Rhône basin which could not be predicted by the model. The model predicted 13% of the variance in the indices of relative abundance at the species level and 23% of this variance at the trait level for all zoogeographic regions combined. However, when focused on reaches within a geographic region, the model explained up to 47% of the same variance. Therefore, geographic regions act as ‘filters’ on the relative abundance of species, but hydraulics do affect fish communities within a given geographical context. 4. For the synthetic ‘community structure indices’, we obtained good predictions from hydraulics independently of the geographical context (variance explained up to 95%). These indices were linked to simple key hydraulic characteristics of river reaches (Froude and/or Reynolds number). The indices enabled interpretations of the links between hydraulics, geomorphology, discharge and community patterns. These links were consistent with existing knowledge of species and their traits. 5. In addition to the above validations, the habitat model partly explained the observed effects of impoundment on fish communities. 6. The present results show that stream hydraulics strongly impact fish community structure. Consequently, our findings confirm that community characteristics can be predicted using models of the local-scale habitat requirements of the species forming the community.     

Theoretical habitat templets, species traits, and species richness: floodplain vegetation in the Upper Rhône River

• 1 The floodplain vegetation at approximately 100 sites located in nine different habitat types of the Upper Rhône River, France, was surveyed three times over the past 27 years. Information on species traits of the higher plants comprising the Rhône floodplain vegetation was based on studies conducted between Geneva, Switzerland, and Lyon, France.
• 2 These data were structured using a ‘fuzzy coding’ technique and then examined using ordination analyses to investigate: (i) relationships among species traits; (ii) habitat utilization; (iii) the relationship between species traits and habitat utilization; and (iv) trends of species traits and species richness in the framework of spatial–temporal habitat variability to test predictions of the river habitat templet and the patch dynamics concept.
• 3 Size, number of descendants per reproductive cycle, number of reproductive cycles per individual, and the regeneration potential of an individual were positively related with each other, whereas the degree of attachment to the soil decreased, and the reproductive period shifted from autumn/late summer towards early summer/spring, as size increased.
• 4 The habitat utilization by the higher plants of the floodplain revealed a double lateral gradient: the first was from the banks of the temporary waters to terrestrial flats; the second from aggrading pebble to aggrading silt habitats. These gradients were related to gradients in water saturation, oxygen conditions, nutrient loading, and nutrient retention of the soils.
• 5 A significant relationship between species traits and habitat utilization was observed for the floodplain vegetation, i.e. plant communities used particular habitat types with a particular set of species trait modalities (= categories).
• 6 Patterns of species trait modalities were significantly related to temporal and spatial habitat variability but only modalities of the trait ‘parental care’ conformed to trends predicted from theory.
• 7 No trends were observed when species richness of different habitat types was considered in the framework of spatial–temporal habitat variability.
• 8 Although the habitats of the Upper Rhône clearly act as a templet for the species traits of the floodplain vegetation, the lack of agreement between observations and predictions on trends in species traits and richness in terms of habitat variability suggest that important elements of theory should be rejected. However, human-induced changes in these habitats are too recent when compared with the longer time periods required for floodplain vegetation to respond to such changes.

     

Fish community changes after minimum flow increase: testing quantitative predictions in the Rhône River at Pierre-Bénite, France

1. Many aspects of the flow regime influence the structure of stream communities, among which the minimum discharge left in rivers has received particular attention. However, instream habitat models predicting the ecological impacts of discharge management often lack biological validation and spatial generality, particularly for large rivers with many fish species. 2. The minimum flow at Pierre‐Bénite, a reach of the Rhône river bypassed by artificial channels, was increased from 10 to 100 m³ s^?1 in August 2000 (natural mean discharge 1030 m³ s^?1), resulting in a fivefold increase in average velocity at minimum flow. Fish were electrofished in several habitat units on 12 surveys between 1995 and 2004. 3. Principal components analysis revealed a significant change in the relative abundance of fish species. The relative abundance of species preferring fast‐flowing and/or deep microhabitats increased from two‐ to fourfold after minimum flow increase. A change in community structure confirmed independent quantitative predictions of an instream habitat model. This change was significantly linked to minimum flow increase, but not to any other environmental variables describing high flows or temperature at key periods of fish life cycle. The rapidity of the fish response compared with the lifespan of individual species can be explained by a differential response of specific size classes. 4. The fish community at Pierre‐Bénite is in a transitional stage and only continued monitoring will indicate if the observed shift in community structure is perennial. We expect that our case study will be compared with other predictive tests of the impacts of flow restoration in large rivers, in the Rhône catchment and elsewhere.     

Theoretical habitat templets, species traits, and species richness: fish in the Upper Rhône River and its floodplain

• 1 Recent developments in ecological theory concerned with habitat templets, species assemblages, and life history traits were examined for the riverine fish communities of the Upper Rhône River, France, in the context of spatial–temporal habitat variability. Relationships among species traits, habitat utilization of species, the relationship between species traits and habitat utilization, and trends of species traits and species richness in the spatial–temporal variability of the habitat types were analysed.
• 2 Relationships among twelve species traits, and utilization of eight habitats were examined for twenty-five fish species using correspondence analysis; the relationship between species traits and habitat utilization was investigated by co-inertia analysis.
• 3 Positive relationships among species traits were observed for size, fecundity, and the number of reproductive cycles per individual. However, species were not well differentiated according to the habitat utilization, except for habitats rarely connected with the main channel (i.e. two types of oxbow lakes).
• 4 No significant relationship was found between species traits and habitat utilization, nor for either species traits or species richness when examined in the framework of spatial–temporal habitat variability. Only two species traits corresponded (with slight trends) to predictions in a river habitat templet: (i) the number of descendants per reproductive cycle increased along with temporal variability; and (ii) the number of reproductive cycles per individual was either low or high at low temporal variability and intermediate at elevated temporal variability.
• 5 The discrepancy between the predictions of the river habitat templet as well as of the patch dynamics concept and the results observed for the fish in the Upper Rhône was explained in terms of scale problems, the evolutionary ecology of the European fish fauna, and the history of the Rhône River.

     

Theoretical habitat templets, species traits, and species richness: 548 plant and animal species in the Upper Rhône River and its floodplain

• 1 For practical reasons, conceptual developments in community ecology are usually based on studies of a restricted systematic group. The cooperation of thirty or so specialists in the synthesis of long-term ecological research on the Upper Rhône River, France, provided a unique occasion to investigate relationships among species traits, the habitat utilization by species, the relationship between species traits and habitat utilization, and trends of species traits and species richness in the framework of spatial-temporal habitat variability for 548 species of plants (Hyphomycetes, aquatic macrophytes, floodplain vegetation) and animals (Tricladida, Oligochaeta, several groups of Crustacea, Insecta and Vertebrata).
• 2 Using correspondence analysis, 100 modalities of eighteen species traits were examined; the resulting typology demonstrates that systematic groups are the most important elements for separating species traits such as size, fecundity of individuals, parental care, mobility, body form, and food type. Small species have an intermediate number of descendants per reproductive cycle and few reproductive cycles both per year and per individual; in contrast, large species have a high number of descendants per reproductive cycle and few reproductive cycles per year but many potential reproductive cycles per individual.
• 3 The analysis of habitat utilization in the Upper Rhône River and its floodplain by the 548 species demonstrated a vertical gradient separating interstitial from superficial habitats; a transverse gradient for superficial habitats from the main channel towards more terrestrial ones is also evident.
• 4 Because of a significant (P < 0.01) relationship between species traits and habitat utilization, traits such as size, fecundity of individuals, parental care, tolerance to variation in humidity, and respiration are arranged along the vertical and transverse habitat gradient. Size, the number of reproductive cycles per individual, and the tolerance to variation of humidity increases from permanent waters to temporary waters, aggrading habitats, and terrestrial habitats.
• 5 Species traits showed significant (P < 0.01) trends in the framework of spatial-temporal habitat variability and were compared with predictions based on the river habitat templet. Although each habitat showed a mixture of species traits at low temporal and spatial variability, and at high variability sites, trends corresponded to predictions for three traits (number of descendants per reproductive cycle, number of reproductive cycles per individual, attachment to soil or substrate) along a gradient of increasing temporal habitat variability.
• 6 The species richness of each habitat within the Upper Rhône River and its floodplain significantly (P = 0.03) increased as the spatial variability of habitats increased but there is no statistical correlation between spedes richness and temporal variability. An altemative hypothesis predicting that fewer spedes per resource occur in temporally stable habitats is also not supported.

     

Can biological traits of stream invertebrates help disentangle the effects of multiple stressors in an agricultural catchment?

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The application of habitat templets and traits to hyphomycete fungi in a mid-European river system

1. The general knowledge about hyphomycetes is summarized and traits of some of these organisms in the Rhône floodplain are described to assess compliance with the river habitat templet (RHT) and patch dynamics concept (PDC) hypotheses. This investigation corresponds with those for other taxonomic groups reported in a recent Freshwater Biology Special Issue on the Ecology of the Upper Rhône River (Statzner, Resh & Dolédec, 1994). 2. Three groups of fungi were considered: aquatic, terrestrial, and aero-aquatic hyphomycetes. 3. The main factors controlling distribution of these micro-organisms are food (mostly organic matter), dissolved oxygen and biotic interactions. 4. With contrasting strategies, hyphomycetes exploit both fallen leaves, a discrete ephemeral resource, and wood fragments, which are more durable. 5. Fungal propagules (spores, hyphae) are distributed in the water, and even the air, largely throughout the year. This explains the resilience of these organisms to disturbance. 6. A major problem is the spatial (boundary layer, dead zones) and temporal (leaf input) scale at which these micro-organisms live and their interactions at the scale adopted for the other organisms referred to in the Special Issue on the Ecology of the Upper Rhône. 7. Our scant knowledge of the microbial ecology in the Rhône floodplain, together with the scale problem and high microbial resilience to environmental stress, render the RHT and PDC hypotheses difficult to test. Only three traits could be tested on hyphomycetes, but all three confirmed the RHT hypotheses. 8. Hyphomycete communities appear to be biologically controlled in more favourable and temporally stable environments, and physico-chemically controlled in harsher and more perturbed environments.     

Theoretical habitat templets, species traits, and species richness: Plecoptera and Ephemeroptera in the Upper Rhône River and its floodplain

• 1 Ephemeroptera and Plecoptera in two sites of the Upper Rhône River (France) were examined using multivariate analyses to determine: (i) relationships among seventeen species traits; (ii) habitat utilization of the fifty-five species present; (iii) the relationship between species traits and habitat utilization; (iv) trends of species traits and species richness in a framework of spatial and temporal habitat variability.
• 2 The species traits having the highest correlation ratios correspond to reproduction or life cycle, behavioural, and morphological characteristics. According to their traits, species of Baetidae, Caenidae, and Leptophlebiidae (Ephemeroptera) are opposite species of Perlidae and Perlodidae (Plecoptera).
• 3 The distribution of species in thirteen habitat types of the Upper Rhône River floodplain demonstrates a transverse gradient from the main channel to the oxbow lakes. Plecoptera are restricted to the different main channel habitats; in contrast, Ephemeroptera families have a broader distribution with Baetidae and Leptophlebiidae occurring in most floodplain habitats.
• 4 Plecoptera exhibit a significant relationship between species traits and habitat utilization but no relationship is evident for Ephemeroptera. Baetidae use many habitat types and have diverse species traits; in contrast, Leptophlebiidae, Heptageniidae, and Caenidae use many habitat types but each family has a rather uniform set of traits.
• 5 Trends in species traits were significantly related to both the spatial and temporal variability of habitats. Considering only temporal variability, the distribution of species trait modalities (= categories) corresponded well to predictions on trends in the river habitat templet for ‘minimum age at reproduction’ and ‘potential longevity’, and in general for ‘descendants per reproductive cycle’, ‘reproductive cycles per year’, ‘potential size’, and ‘body flexibility’ trends in six other traits did not match predictions.
• 6 No trends in species richness were evident in spatial–temporal framework of habitat variability.

     

The response of life‐history traits to a new species in the community: a story of Drosophila parasitoids from the Rhône and Saône valleys

In the present study, we investigated the evolution of life‐history traits in the main species of a community, after the arrival of a new competitor. Two parasitoid species, Leptopilina heterotoma and Asobara tabida, are present throughout the Rhône and Saône valleys, whereas a third species, Leptopilina boulardi, is slowly extending its distribution northwards. In the presence of L. boulardi, competing parasitoids experience a higher mortality and lower host availability. Resources should thus be re‐allocated between traits according to these new factors. We compared life‐history traits of populations of L. heterotoma and A. tabida in areas with and without L. boulardi. As predicted by both Price's balanced mortality hypothesis and the theory of life‐history traits, we found that investment in reproduction is higher in southern populations for both native species, coupled with higher travelling abilities. However, only A. tabida paid their higher fecundity by a lower longevity. The absence of a clear trade‐off between these traits in L. heterotoma may be explained by a lower metabolic rate in southern populations. These results highlight the importance of the community change over climate in the evolution of life‐history traits in this parasitoid community. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Linking traits to species diversity and community structure in phytoplankton

In addition to answering Hutchinson’s question “Why are there so many species?”, we need to understand why certain species are found only under certain environmental conditions and not others. Trait-based approaches are being increasingly used in ecology to do just that: explain and predict species distributions along environmental gradients. These approaches can be successful in understanding the diversity and community structure of phytoplankton. Among major traits shaping phytoplankton distributions are resource utilization traits, morphological traits (with size being probably the most influential), grazer resistance traits, and temperature responses. We review these trait-based approaches and give examples of how trait data can explain species distributions in both freshwater and marine systems. We also outline new directions in trait-based approaches applied to phytoplankton such as looking simultaneously at trait and phylogenetic structure of phytoplankton communities and using adaptive dynamics models to predict trait evolution.     

Potential impact of invasive amphipods on leaf litter recycling in aquatic ecosystems

The impact of biological invasions on local biodiversity is well established, but their impact on ecosystem functioning has only been sketchily documented. However, biological invasions may impede services provided by aquatic ecosystems, such as, for example, the decomposition of organic matter, a key process in most small streams. To address this question, we experimentally quantified the leaf litter breakdown activity of native and invasive amphipod species, which are keystone species in aquatic ecosystems. The breakdown rate of each species was used to estimate the potential leaf litter recycling in the Rhône and Meurthe Rivers in sites occupied solely by native species and sites dominated by invasive species. We found that invaders were not able to compensate for the activity of native species and that the replacement of native species led to a decrease of at least 66% in the rate of leaf litter recycling. Our approach provides empirical evidence of the functional impact of non-indigenous species on leaf litter recycling, using standard protocols and literature data.     

Phylogeographical structure, postglacial recolonization and barriers to gene flow in the distinctive Valais chromosome race of the common shrew (Sorex araneus)

Using one male‐inherited and eight biparentally inherited microsatellite markers, we investigate the population genetic structure of the Valais chromosome race of the common shrew (Sorex araneus) in the Central Alps of Europe. Unexpectedly, the Y‐chromosome microsatellite suggests nearly complete absence of male gene flow among populations from the St‐Bernard and Simplon regions (Switzerland). Autosomal markers also show significant genetic structuring among these two geographical areas. Isolation by distance is significant and possible barriers to gene flow exist in the study area. Two different approaches are used to better understand the geographical patterns and the causes of this structuring. Using a principal component analysis for which testing procedure exists, and partial Mantel tests, we show that the St‐Bernard pass does not represent a significant barrier to gene flow although it culminates at 2469 m, close to the highest altitudinal record for this species. Similar results are found for the Simplon pass, indicating that both passes represented potential postglacial recolonization routes into Switzerland from Italian refugia after the last Pleistocene glaciations. In contrast with the weak effect of these mountain passes, the Rhône valley lowlands significantly reduce gene flow in this species. Natural obstacles (the large Rhône river) and unsuitable habitats (dry slopes) are both present in the valley. Moreover, anthropogenic changes to landscape structures are likely to have strongly reduced available habitats for this shrew in the lowlands, thereby promoting genetic differentiation of populations found on opposite sides of the Rhône valley.     

The role of functional traits and trade-offs in structuring phytoplankton communities: scaling from cellular to ecosystem level

Trait-based approaches to community structure are increasingly used in terrestrial ecology. We show that such an approach, augmented by a mechanistic analysis of trade-offs among functional traits, can be successfully used to explain community composition of marine phytoplankton along environmental gradients. Our analysis of literature on major functional traits in phytoplankton, such as parameters of nutrient-dependent growth and uptake, reveals physiological trade-offs in species abilities to acquire and utilize resources. These trade-offs, arising from fundamental relations such as cellular scaling laws and enzyme kinetics, define contrasting ecological strategies of nutrient acquisition. Major groups of marine eukaryotic phytoplankton have adopted distinct strategies with associated traits. These diverse strategies of nutrient utilization can explain the distribution patterns of major functional groups and size classes along nutrient availability gradients.     

长白山森林／沼泽生态交错带群落和环境梯度分析

揭示了森林－沼泽过渡带群落的结构、生产力、植物多样性等群落梯度和交错环境梯度的相关规律,并结合交错区环境梯度分析这些群落特征形成机制,为维持、保护与经营管理这一交错带生物资源提供了理论依据。将长白山地区森林和高、中、低位沼泽所形成的三大类型过渡带研究对象,采用样带网格的调查方法,并应用系统软件分析方法建立了经验回归模型,研究了森林／沼泽生态交错带群落的种类组成、群落建群种径级结构与年龄结构、植物多样性、群落生产力及其随生态交错带环境梯度变化趋势。结果表明,森林／沼泽生态交错带群落结构特征、植物多样性、群落生产力均随着交错带环境梯度的变化而呈现有规律的分布格局,沿着沼泽至森林方向的交错区环境梯度,群落建种种发生更替现象;群落种类数目呈现指数递增趋势;群落的径级结构呈现双曲线分布规律性;年龄结构一般呈三次式分布规律;揿样性呈二次式梯增分布趋势;群落生物量均呈现三次函数曲线递增趋势,表现出群落梯度和环境梯度的高度相关性。     

Traits uncover quasi-neutral community assembly in a coastal heath vegetation

Aims Trait-based approaches are increasingly being used in ecology due to their potential to explain and predict community structure and dynamic regardless of species identities. Here, we asked whether coastal heath vegetation established in stressful habitats could be assembled by quasi-neutral mechanisms based on functional trait distribution. Communities are organized through quasi-neutral dynamics if abiotic gradients and/or species interactions impose sorting on functional traits and species performances, but not on species phylogenetic relatedness and occurrence.     

Quantifying the role of deterministic assembly and stochastic drift in a natural community of Arctic mosses

The interpretation of natural plant communities frequently invokes species‐sorting controlled by niche differences along spatial environmental gradients. This process of niche structuring can be explained by reference to functional traits, which provide a mechanistic explanation for community structure. In contrast, models explaining species coexistence obviate the limiting effect of niche difference, by invoking processes which cause species‐level drift, e.g. demographic stochasticity. This paper investigates a simple habitat with strong gradients (moss communities in a patterned arctic wetland) to identify signature‐patterns under‐pinning the relative importance of deterministic assembly and stochastic drift in a natural community. First, ordination analysis was used to confirm community composition structured by a range of nine carefully selected functional traits. Second, to determine whether traits explaining community composition might also explain species richness, local species richness (sR) was compared to (1) observed trait diversity and (2) expected trait diversity based on permutation tests, which are used to simulate null community assembly for different values of sR. Traits explaining species composition, consistent with deterministic niche structuring, do not appear to maintain sR. This surprising result was explained by decomposing the community into individual pair‐wise comparisons, i.e. species niche‐differences and association (χ²). Results support deterministic processes via the sorting of species with similar and contrasting niches, at opposite ends of a composite environmental gradient. Nevertheless, stochastic drift is apparent in the random structure of a majority of pair‐wise associations; in addition, a species’ abundance was in general not related to environmental distance from response‐optima. We suggest therefore that spatial pattern in the moss community is a balance between deterministic forces with respect to species traits and controlling environmental gradients, and stochastic drift, which weakens this deterministic structure.