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

• 1 The objective of this study, which is based on forty-two species of hydrophytes and helophytes, is to investigate: (i) relationships among species traits; (ii) habitat utilization by species; (iii) the relationship between species traits and habitat utilization; (iv) trends in species traits in the framework of spatial–temporal habitat variability, and if trends match predictions from the river habitat templet; and (v) trends in species richness in the framework of spatial–temporal habitat variability, and if trends match predictions of the patch dynamics concept.
• 2 Two data sets were used for this analysis: species traits (mainly reproductive and morphological characteristics) were documented from the literature; and species distribution across eight habitat types was from field surveys conducted in the floodplain of the Upper Rhone River, France. This information was structured by a fuzzy coding technique and analysed by ordination methods.
• 3 Several species traits, which are related to disturbances and reflect resistance (e.g. attachment to soil or substrate) or resilience (e.g. potential for regeneration of an individual), are closely related for aquatic macrophytes.
• 4 Habitat utilization by aquatic macrophytes separates the habitat types along a gradient of connectivity with the main channel, which corresponds to a gradient in flood disturbance frequency and the permanence of the different water-bodies.
• 5 The relationship between species traits and habitat utilization is highly significant, indicating that a particular set of habitat types is used by taxa with a particular set of species trait modalities.
• 6 Observations in one habitat templet (in which scaling of the templet is primarily based on water level fluctuations for the temporal variability axis and on substrate characteristics for the spatial variability axis) generally do not support predictions on trends in species traits but do support predictions on trends in species richness.
• 7 Observations in an alternative habitat templet (in which scaling of the templet is based on frequency of flood scouring for the temporal variability axis and on heterogeneity of the substrate for the spatial variability axis) support theoretical predictions on trends for about half of the species traits for which predictions were available. However, trends in species richness in this alternative habitat templet are only partly in agreement with predictions.

     

Theoretical habitat templets, species traits, and species richness: a synthesis of long-term ecological research on the Upper Rhône River in the context of concurrently developed ecological theory

• 1 Based on information obtained from analysis of thirteen taxonomic groups of plants and animals occurring in the alluvial floodplain habitats of the Upper Rhône River, France, we synthesize results obtained on: (i) relationships among species traits; (ii) habitat utilization by species; (iii) the relationship between species traits and habitat utilization; (iv) trends in species traits in a framework of spatial and temporal variability; and (v) tests of trends predicted for species traits and species richness in the framework of spatial and temporal habitat variability in terms of the river habitat templet and patch dynamics concept.
• 2 Species traits describing reproductive characteristics, food, and size had the closest relationships with each other in the various correspondence analyses performed. Faunal and floral separation by species traits produced groupings similar to those based on traditional taxonomy.
• 3 Two major gradients appear in the utilization of the floodplain habitats: a vertical gradient from interstitial to superficial habitats; and a transverse gradient from the main channel to oxbow lakes, temporary waters, and terrestrialized habitats.
• 4 For the majority of the groups examined, a statistically significant relationship was evident between the structure of the species trait and habitat utilization arrays. For these groups, the characteristics of the habitat act as a templet for species traits. Moreover, species trait modalities (i.e. categories defining traits) were significantly arranged along the axis of spatial and temporal variability for most groups, which indicates that such variability acts as a templet for species traits.
• 5 Species traits did not conform to predictions of the river habitat templet because the observed modality sequences did not follow the trends predicted in a framework of spatial and temporal variability. Moreover, there was no clear pattern in the distribution of species traits along an axis of temporal variability for groups of organisms having different sizes, which is a correlate of longevity, nor did modalities of species traits that occur under conditions of low temporal variability also tend to occur under conditions of high spatial variability (or vice versa). Clearly, species traits occur as alternative suites of characteristics in various groups of organisms.
• 6 The patch dynamics concept, which predicts that highest species richness occurs at intermediate levels of temporal variability and highest levels of spatial variability, was supported by observations in only two of the thirteen groups exaniined, and only partially (for spatial variability) when all 548 taxa were examined together.
• 7 The predictions of the river habitat templet and patch dynamics concepts were not supported, perhaps because templet theories do not yet accommodate alternative suites of characteristics and trade-offs between combinations of traits, or perhaps because the single scale of variability considered in the analyses, the inhomogeneity of the available biological information, and the aggregation of spedes traits that were used created methodological problems.
• 8 Ecologically sound river management polides eventually may be based on two key points that emerged from this synthesis: that the habitat acts as a templet for spedes traits, and that composite taxonomic groupings represent relatively homogeneous assemblages of spedes trait modalities.
• 9 The use of statistical approaches developed in this project to analyse other long-term data sets may clarify questions about the applicability of habitat templet theories to river ecology, and hasten development of ecologically sound river management policies.

     

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

• 1 To test predictions of the river habitat templet and the patch dynamics concept, trends in species traits and species richness of aquatic beetles were related to the spatial-temporal variability of eighteen habitat types in the alluvial floodplain of the French Upper Rhône River. One hundred and twenty species of beetles were used in this analysis.
• 2 The basic information was obtained either from the literature (for most of the species traits) or from observations made at approximately 500 sampling sites in the Brégnier-Cordon and Jons sections over the past 19 years (for habitat utilization). This information was structured by a fuzzy coding technique and examined by ordination analyses.
• 3 Analyses of the relationships among nineteen species traits revealed a clear distinction according to traits such as body form (for adults), functional feeding type and food (adults and larvae), attachment to the substrate and dissemination potential (adults and larvae), and patterns of aquatic and/or terrestrial life of adults and larvae. Species traits such as number of descendants per reproductive cycle, and number of reproductive cycles per year or per individual showed less contrast, because these traits are rather homogeneous in aquatic beetles.
• 4 Analyses of the habitat utilization by the aquatic beetles revealed a vertical gradient that separates interstitial from superficial habitats, and a transverse gradient for the superficial habitats, which extends from the main channel towards permanent oxbow lakes and temporary waters.
• 5 The significant relationship betweeen species traits and habitat utilization demonstrates that most beetle species use a particular set of habitat types with a particular set of species trait modalities.
• 6 Species traits of aquatic beetles are homogeneous but evidently very successful and are adapted to many potential conditions of spatial–temporal variability. Because of this homogeneity, observations on aquatic beetles do not support trends of traits in the framework of spatial–temporal variability predicted from the river habitat templet.
• 7 The observed species richness of aquatic beetles is low in habitat types with a low spatial–temporal variability, increases as spatial variability increases, and tends to be highest at intermediate temporal variability. This pattern matches predictions of the patch dynamics concept.

     

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

• 1 For Trichoptera occurring in two sites of the Upper Rhône River (France) we examine: (i) relationships among species traits; (ii) habitat utilization of Trichoptera species; (iii) the relationship between species traits and habitat utilization; (iv) trends of species traits in the framework of spatial–temporal habitat variability to test predictions of the habitat templet concept; and (v) trends of species richness in the framework of spatial–temporal habitat variability to test predictions of the patch dynamics concept.
• 2 Of the sixteen species traits selected, twelve have high correlation ratios for the seventy-five species used in this analysis; these traits are related to behavioural, morphological, or physiological aspects. Traits related to reproduction or life cycle have lower correlation ratios.
• 3 An ordination by species traits separates the five main families into three groups: (i) Hydropsychidae and Polycentropodidae; (ii) Hydroptilidae; and (iii) Leptoceridae and Limnephilidae. An ordination of the habitat utilization of the species in ten habitats indicates that the Hydropsychidae occur preferentially in the main channel, Hydroptilidae, Polycentropodidae, and Limnephilidae occur in backwaters or oxbow lakes, and the Leptoceridae are ubiquitous.
• 4 The Hydropsychidae exhibit a relationship between species traits and habitat utilization, i.e. they use similar habitat types with similar species traits. The species traits of the other four families are similar but their habitat utilization is quite different.
• 5 The Hydropsychidae occur in lowest spatial–temporal variability habitats and Limnephilidae in the highest. Therefore, net spinners and filterers are characteristic of habitats with a low spatial–temporal variability, whereas shredders and case makers using plant material are characteristic of habitats with high spatial–temporal variability. The trends in species traits show little agreement with trends predicted from the river habitat templet.
• 6 Trends of species richness in the framework of spatial and temporal variability do not follow the predictions of the patch dynamics concept because richness is similar in all superficial habitats. This implies that each habitat, in spite of large differences in their spatial and temporal variability, offers Trichoptera a similar but limited number of ecological niches.

     

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

• 1 This paper summarizes twenty years of ecological research on aquatic oligochaetes of the Upper Rhône River and its alluvial floodplain (France). Species traits of fifty species of the ninety taxa recorded from two areas Gons and Brégnier-Cordon) were used to examine the relationships among species traits, habitat utilization of these species, whether a relationship exists between species traits and habitat utilization, and the applicability of predictions from the river habitat templet and the patch dynamics concept in the framework of spatial and temporal habitat variability. We used fourteen habitat types and sixteen species traits in this analysis.
• 2 When examined by correspondence analysis, species traits separate the Naididae (with a higher potential for reproduction, small size, high mobility, and opportunistic diet) from all other families.
• 3 Habitat utilization by oligochaetes demonstrates two gradients: a vertical gradient that arranges species by their affinity for interstitial habitats (stygophily) and a transversal gradient that arranges them by their affinity for main channel habitats (rheophily).
• 4 No significant relationship was found between species traits and habitat utilization in a co-inertia analysis.
• 5 Trends observed for species traits within the framework of spatial-temporal habitat variability show only minor agreement with predictions of the river habitat templet.
• 6 Species richness is generally higher in superficial and interstitial habitats that are permanently connected with the main channel, and peaks in the superficial parapotamons (backwaters that are permanently connected with the main channel) characterized by intermediate levels of spatial as well as temporal variability; this pattern only partially fits with predictions of the patch dynamics concept.

     

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: amphibians in the Upper Rhône River and its floodplain

• 1 The purpose of this study was to test predictions of the habitat templet and the patch dynamics concept by analysing the relationship between either the species traits or species richness of amphibians and the spatial–temporal variability of eight habitat types of the Upper Rhône River and its floodplain, France.
• 2 The information on species traits of the twelve amphibian species was accumulated primarily from the literature; that on habitat utilization was based on field surveys. The information was ‘fuzzy coded’, analysed by ordination techniques, and finally linked to the spatial–temporal variability of habitat types elaborated elsewhere in this issue.
• 3 After elimination of the variance caused by the differences between urodeles and anurans, correspondence analysis of the matrix of species traits demonstrated that the flexibility of the temporal pattern of reproduction was the major source of variation among amphibian species. A less significant source of variation within anurans was related to traits usually linked to the concept of r–K selection (potential number of descendants per reproductive cycle and ratio of egg size to female size).
• 4 A correspondence analysis of habitat utilization by amphibian species separated four groups of habitat types, corresponding to different degrees of influence that the active channel had on adjacent waters in the floodplain (i.e. from habitats that are most frequently disturbed by floods, to habitats of braided sections, old oxbow lakes, and the more terrestrialized habitats).
• 5 A co-inertia analysis of the relationship between species traits and habitat utilization separated two habitat types (terrestrialized former meanders and oxbow lakes) from the others. Species of terrestrialized habitats had a reproductive display, a low number of descendants per reproductive cycle, and a high ratio of egg size to female size. Species of oxbow lakes were early or synchronous breeders, and laid a single clutch.
• 6 Neither the predictions of the habitat templet concept nor those of the patch dynamics concept were validated. Species traits were not significantly correlated to the axes of the spatial–temporal variability templet. The highest species richness was observed in habitat types with the highest temporal variability; in addition, richness did not peak at highest spatial variability.
• 7 Deviation from the predictions of the river habitat templet could be explained by: (i) the over-simplification of this model with regard to the evolution of the complex life cycles of the amphibians; and (ii) an underestimation of the importance of phylogenetic constraints and the evolution of community interactions.

     

Theoretical habitat templets, species traits, and species richness: ostracods (Crustacea) in the Upper Rhône River and its floodplain

• 1 Ostracods occurring at two sections of the Upper Rhône River, France, were examined to determine relationships among species traits, habitat utilization, the relationship between species traits and habitat utilization, and trends in species traits and species richness in the context of spatial and temporal variability of habitats. Twenty regularly sampled species were used in this study and fifteen species traits were considered.
• 2 Throe groups can be distinguished according to their species traits: group 1 has species of mixed sizes with high reproductive rates, short life span, spherical shape, long swimming bristles, low thigmotactism, and high resistance to desiccation; group 2 has medium-sized species with low reproductive rates, long life span, low or no tolerance to desiccation, geometric (trapezoidal, triangular) or streamlined carapace shape, no swimming bristles, and a strong thigmotactism; group 3 has the largest species with parthenogenetic reproduction, medium-sized swimming bristles, and flattened or cylindric carapace shape.
• 3 Ostracod habitat utilization segregates the superficial and interstitial habitats along a gradient from the main channel to the abandoned arms and to the temporary waters.
• 4 The co-structure (= relationship) between species traits and habitat utilization indicates that the species use particular habitats with a particular set of species trait modalities. Species with long life spans, late maturity, low fecundity, and low migratory ability are restricted to the interstitial habitats; the epigean species with long life spans, large size, and parental care are more abundant in permanent flowing and standing surface waters; the epigean species with short life spans, high migratory ability, and high tolerance to desiccation are more abundant in temporary ponds.
• 5 The analyses of the distribution of the species traits in a river habitat templet of spatial and temporal variability emphasized that the main disturbance structuring the Rhône River ostracod assemblage is desiccation.
• 6 Of the trends predicted for species traits in the framework of the river habitat templet, five (size, body form, attachment, reproductive technique, and mobility) are clearly opposite for ostracods (because the predictions were mainly established for flood-related disturbances) but four (life span, number of reproductive cycles per year, age at first reproduction, and desiccation tolerance) are in agreement.
• 7 No trends in ostracod species richness in the framework of spatial–temporal habitat variability were evident.

     

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.

     

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

• 1 For five orders of Insecta (Plecoptera, Ephemeroptera, Odonata, Trichoptera, and Coleoptera) in two sites of the Upper Rhône River (France), the following are examined: (i) relationships among nineteen species traits; (ii) habitat utilization of species; (iii) the relationship between species traits and habitat utilization; and (iv) trends of species traits and species richness in a templet of spatial-temporal habitat variability.
• 2 The species traits having the highest correlations correspond to reproduction, life cycle, nutritional, and morphological features. Species trait characteristics of Coleoptera are distinctly contrasted with those of Plecoptera and Ephemeroptera; Odonata and Trichoptera are intermediate to these orders.
• 3 The distribution of species in fourteen habitat types of the Upper Rhône River floodplain demonstrates a transverse gradient from the main channel to the oxbow lakes and the temporary water habitats, and a vertical gradient from interstitial to superficial habitats.
• 4 Despite a significant relationship between species traits and habitat utilization, superposition between species traits and habitat utilization is limited. At the order level, species form usually one (Ephemeroptera, Trichoptera, Odonata) or several (Coleoptera) groups of relatively homogeneous species traits; however, the species of each of these groups utilize rather different habitat types.
• 5 Only for some life history traits, e.g. the minimum age of reproduction or the number of reproductive cycles per year, do the trends observed in the framework of spatial—temporal variability of habitat types agree with the predictions from the river habitat templet. This mismatch mainly results from the unique phylogenetic history of the Coleoptera compared with that of the other four orders.
• 6 Species richness peaks at an intermediate level of temporal variability; however, it does not gradually increase with increasing spatial variability, nor increase from low to intermediate temporal variability.

     

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.

     

Temporal and spatial environmental variability in the Upper Rhône River and its floodplain

• 1 This paper develops a framework of spatial and temporal variability for a habitat typology of the Upper Rhône River (France) and its alluvial floodplain that is based on about 17 years of data collection and analysis. The aim was to provide a scale of spatial-temporal variability for river habitat templet predictions on trends in species traits and species richness.
• 2 In developing this framework, eight physical-chemical variables were available and could be considered for twenty-two habitat types: seventeen superficial (surface) and five interstitial (0.5 m below the substrate surface). These habitat types were selected in two areas (Jons and Brégnier-Cordon) after geomorphological considerations and because of differences in their biological characteristics.
• 3 The data sets used were processed by a ‘fuzzy coding’ method using, for each variable, the frequency distribution (by modalities = categories) of all measurements and monthly means over an annual scale. Two tables were produced; the first corresponded to an expression of the total variability, and the second represented an evaluation of the temporal variability.
• 4 Each of these tables was analysed by correspondence analysis, which provided factorial scores that were used to calculate, by habitat type and by variable, a total variability and a temporal variability in terms of cumulated variability of factorial scores for the eight physical–chemical variables. The rationale in describing variability from these two tables is that total variability equals temporal variability plus spatial variability. The spatial variability was then determined by the difference between total and temporal variability. From this procedure, a positioning of the twenty-two habitat types on the spatial and temporal variability axes was obtained.
• 5 The estimate of spatial variability did not consider any error term that may have occurred in the above model; it was then tested by an independent assessment of the spatial variability using thirteen variables in nine major habitat types. A high correlation between the two ways of assessing spatial variability (r = 0.85, P < 0.004) underscored the reliability of the spatial variability that was calculated previously.
• 6 The river habitat templet obtained for the Upper Rhône and its alluvial floodplain appears to be appropriate to test the predictions on patterns of species traits and species richness in the framework of spatial and temporal variability.

     

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

• 1 To test predictions of the river habitat templet and the patch dynamics concept, twenty species traits and the species richness of eighty-one bird species were examined in the context of the spatial – temporal variability of ninety-one sites at Jons, a section of the Upper Rhône River, France.
• 2 The basic information was obtained either from the literature (for species traits) or by observation, then structured by a fuzzy coding technique, and examined using multivariate analyses.
• 3 The relationships between species traits separated three groups of birds: (i) large species that are long lived, have long breeding cycles, and use open sites to feed, breed, and sing (e.g. Podicipidae, Laridae); (ii) medium-sized species with intermediate longevities and breeding cycles that use less open sites (e.g. Columbidae, Corvidae); and (iii) small species that are short lived, have short breeding cycles, and use closed sites (e.g. Sylviidae).
• 4 The distribution of birds in the floodplain habitats of the river was closely related to the spatial–temporal variability of sites.
• 5 No trends in species traits were observed along a gradient of spatial–temporal variability of the sites.
• 6 Species richness of birds was relatively constant over the gradient of spatial–temporal variability of the ninety-one sites, but decreased at low spatial and high temporal variability because of human activities and vegetation changes that occurred at these sites.
• 7 These observations on species traits and richness were related to the high mobility of birds (which is a scale phenomenon) and the high intercorrelation between parameters used to determine the spatial and temporal variability of the sites (which makes separation of the spatial from the temporal variability impossible).

     

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.

     

Stream community structure in relation to spatial and temporal variation: a habitat templet study of two contrasting New Zealand streams

• 1 The physical characteristics of two contrasting streams, and habitat types within these streams, are described in terms of a two-dimensional physical habitat templet in which disturbance frequency and the availability of spatial refugia are the temporal and spatial axes.
• 2 It is predicted that habitats experiencing a high disturbance frequency and low refuge availability will be characterized by a low invertebrate species diversity, a low biomass of epilithic algae and particulate organic matter and a community made up of mobile, weedy species. Habitats having a low disturbance frequency and high refuge availability will be characterized by a diverse community containing sedentary and specialist species, with high algal and particulate organic matter levels.
• 3 A lower median substrate particle size and higher shear stress regime in Timber Creek were indicative of a higher disturbance frequency than in the Kyeburn. Substrate diversity was lower in Timber Creek than in the Kyeburn and indicated that the availability of refugia was lower in Timber Creek. In both streams, pools were found to have a higher disturbance frequency and lower availability of refugia than riffles.
• 4 Invertebrate species diversity, the biomass of epilithic algae and particulate organic matter and the representation of sedentary species, filter feeders and shredders were higher in the more temporally stable and spatially heterogeneous Kyeburn. The community of Timber Creek, frequently disturbed and having low refuge availability, had a high proportion of mobile and weedy species, with the highly mobile, generalist-feeding Deleatidium spp. (Ephemeroptera; Leptophlebiidae) being the most dominant organisms.
• 5 The predictions made about stream community structure and species characteristics in relation to disturbance frequency and the availability of spatial refugia are generally supported. Now a larger scale investigation is required to test the generality of the predictions. We conclude that the habitat templet approach offers a sound framework within which to pose questions in stream ecology.

     

Metacommunity structure of aquatic gastropods in a river floodplain: the role of niche breadth and drift propensity

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Species traits in relation to temporal and spatial heterogeneity in streams: a test of habitat templet theory

1. The habitat templet approach depends on defining templet axes appropriate to the organism(s) of interest, predicting the traits of species associated with different parts of the templet, and testing these predictions in a range of habitats whose positions in the templet have been determined. 2. In this study of thirty-five benthic insect taxa at fifty-four tributary sites of the Taieri River on the South Island of New Zealand, we chose as the temporal axis the intensity/frequency of disturbance, defined in terms of bed movement during high discharge events. As the spatial axis, we postulated that three features would provide refugia and therefore ameliorate disturbance—percentage of the bed with low shear stress, percentage of the bed made up of large substratum particles and availability of interstitial space in the bed—from which we derived a combined multivariate refugium axis. 3. More disturbed communities contained a significantly higher percentage of individuals possessing the following traits: small size, high adult mobility, habitat generalist (each predicted to confer resilience in response to disturbance), clinger, streamlined/flattened and with two or more life stages outside the stream (each predicted to confer resistance in the face of disturbance). When analyses were performed on the percentage of taxa having particular traits, the predicted positive relationships with average bed movement were found for high adult mobility and habitat generalist traits. 4. The percentage of variance in trait scores explained by intensity of disturbance was generally higher in sites with less refugia available and lower in sites further from the headwaters. The percentage of variance explained was higher in sites recently subject to a major high discharge disturbance, suggesting that disturbances tend to strengthen the pattern of preponderance of resilience/resistance traits. 5. We mapped insect taxa onto the two-dimensional templet, following Grime et al.’s triangular terrestrial plant classification. The full variety of resistance and resilience traits were represented in insect species throughout the templet, but taxa associated with more disturbed conditions generally displayed a larger number of resilience and resistance traits, combined, than taxa associated with more stable stream beds.     

Life-history strategies in freshwater macroinvertebrates

• 1 Explaining spatial and temporal differences in species assemblages is a central aim of ecology. It requires a sound understanding of the causal mechanisms underlying the relationship of species with their environment. A species trait is widely acknowledged to be the key that links pattern and process, although the enormous variety of traits hampers generalization about which combination of traits are adaptive in a particular environment.
• 2 In three steps, we used species traits to match species and environment, and chose lentic freshwater ecosystems to illustrate our approach. We first identified key environmental factors and selected the species traits that enable the organism to deal with them. Secondly, we investigated how investments in these traits are related (e.g. through trade‐offs). Thirdly, we outlined 13 life‐history strategies, based on biological species traits, their interrelations known from life‐history theory and their functional implications.
• 3 Species traits and environmental conditions are connected through life‐history strategies, with different strategies representing different solutions to particular ecological problems. In addition, strategies may present an integrated response to the environment as they are based on many different traits and their interrelationships. The presence and abundance of (species exhibiting) different life‐history strategies in a location may therefore give direct information about how a particular environment is experienced by the species present.
• 4 Life‐history strategies can be used to (i) explain differences in species assemblages either between locations or in different periods; (ii) compare waterbodies separated by large geographical distances, which may comprise different regional species pools or span species distribution areas and (iii) reduce often very complex, biodiverse assemblages into a few meaningful, easily interpretable relationships.

     

Environmental heterogeneity and dispersal limitation explain different aspects of β‐diversity in Neotropical fish assemblages

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