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.     

Species traits help explain butterfly habitat use in a tropical forest

1. When species can access all parts of the landscape, species-sorting metacommunity theory predicts that community composition depends on habitat choice and interactions with other species and the environment. These filtering processes can also depend on species' traits.
2. The authors investigated how traits mediate a species-sorting process in determining butterfly community composition in a naturally patchy landscape in the tropical Western Ghats, India. The authors asked, do traits mediate access to certain habitats and does seasonality affect these patterns? The authors surveyed 56 habitat patches in three habitat types: laterite plateau grasslands, ridge grassland, and moist-deciduous forest, in a 65-km² landscape.
3. Non-palatable butterflies showed similar occurrences across seasons and habitats, but palatable butterflies were less commonly encountered in open habitats in the dry season. Polyphagous butterflies occurred infrequently in the dry season in laterite habitats, potentially indicating emigration or diapause patterns are linked to diet breadth.
4. All species were present in all habitats, implying dispersal does not limit access to different habitat patches, consistent with the species-sorting metacommunity concept. Nevertheless, butterfly occurrence was strongly influenced by the interaction of mobility and habitat type with sedentary species occurring less often in low-resource open laterite patches than mobile species.
5. Species sorting is typically regarded as occurring directly through environmental filters, but here the authors suggest that the environmental filter acts through movement limitations. Studies integrating landscape heterogeneity and species characteristics will help us better understand metacommunities and species distributions in nature.

     

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.     

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.

     

Environmental restraints and life strategies: a habitat templet matrix

Summary Four basic environmental restraints on life are deduced from the requirements of life's inherent order laws. Possible life strategies to contend with these restraints are listed. The various combinations of the restraints are subsequently investigated, and appropriate combinations of life strategies are fitted. This model is finally tested against insect case histories in various environments, and is demonstrated to explain some combinations of characteristics of insects in ecosystems not covered by the r-K or r-K-A continua. The role of heterochrony in achieving appropriate life strategies is briefly discussed.     

Species traits for future biomonitoring across ecoregions: patterns along a human-impacted river

1. Current budgets for environmental management are high, tend to increase, and are used to support policy and legislation which is standardized for large geographic units. Therefore, the search for tools to monitor the effects of this investment is a major issue in applied ecology. Ideally, such a biomonitoring tool should: (1) be as general as possible with respect to its geographic application; (2) be as specific as possible by separating different types of human impact on a given ecosystem; (3) reliably indicate changes in human impact of a particular type; and (4) be derived from a sound theoretical concept in ecology. 2. We developed an approach to biomonitoring which matches these ‘ideal’ characteristics by focusing on numerous, general biological species traits (e.g. size, number of descendants per reproductive cycle, parental care, mobility) and on the habitat templet concept, which relates trends in these general species traits to disturbance patterns. Using the French Rhône River and benthic macroinvertebrates as an example, we have used the data to demonstrate a general framework and the potential of our approach rather than to produce a ready-made tool. Our data covered a large river and its major tributaries, which has a catchment that crosses ecoregions, and known gradients and discontinuities in human impact. 3. We applied multivariate analyses to evaluate how the distribution of species traits in invertebrate communities could discriminate environmental differences along the Rhône in comparison to traditionally used approaches (e.g. community structure, based on species abundances, or ecological species traits, such as velocity preferences and pollution tolerance). Invertebrate community structure expressed in terms either of the abundance or the traits of species reliably indicated differences in overall human impact. The community structure based on biological traits was less confounded by natural spatial gradients and reliably indicated human impact, while community structure based on ecological traits was the most confounded by natural spatial gradients and was the poorest indicator of human impact. Community structure based on species abundances was an intermediate indicator of human impact. 4. These results indicate that a revision of biomonitoring approaches which have been based on a single aspect of the biological responses may be warranted. The biological traits of species could separate the different types of human impact. Therefore, the use of these traits in biomonitoring could improve existing multi-metric approaches. Future research has to show if the general applicability of species traits allows the development of a unique biomonitoring tool for running waters of the European Union, for running waters in temperate climates on several continents, for freshwater, marine and terrestrial systems, and/or for global biodiversity assessment.     

The stream as a habitat templet for bryophytes: species' distributions along gradients in disturbance and substratum heterogeneity

1. The structure of bryophyte communities in streams in relation to habitat characteristics, especially disturbance, is described. Disturbance in rivers is quantified as movement of the stream bed, whereas in small streams water level fluctuation is used as an indicator of disturbance frequency. 2. Canonical correspondence analysis differentiated frequently disturbed sites from more stable ones. The existence of a disturbance gradient was confirmed in a subset of the study sites using long-term records of discharge variation. A parallel change was detected in the species composition of bryophyte communities with low-statured, potentially fast colonizers dominating the disturbed end and large perennial species the stable end of the gradient. 3. A consistent pattern of zonation of bryophyte species was found along the gradient from continually submersed to persistently dry conditions in small streams and lake outlets. An abrupt increase in species richness occurred at or just above the water line, where facultatively aquatic species tolerant of both conditions formed the bulk of the community. 4. The relationship between species richness and standing crop in stream bryophyte communities was consistent with the hump-backed model of Grime (1979), especially at the within-habitat scale. Quadrats of low and very high standing crop were characterized by low species richness, while peak richness was observed at intermediate standing crops. 5. A few perennial species (e.g. Fontinalis spp. and Rhynchostegium riparioides) capable of monopolizing space dominated the most stable habitats. Species composition in low biomass sites was more variable, yet only one basic growth-form (small-statured species with high allocation to spore production) seemed possible in these highly disturbed streams. In habitats of intermediate biomass, small-scale disturbances apparently allow the formation of a more varied bryophyte community. 6. A habitat templet for stream bryophyte life strategies and community structure is presented. Disturbance is proposed to be the factor filtering out traits unsuitable for a given environment. The potential of stream bryophytes for testing and developing general ecological theory is emphasized.     

Species traits and the response of open‐habitat species to forest edge in landscape mosaics

Human driven changes in land‐use have increased the need to understand how landscape structure affects species distribution. We studied how forest edges affected the distribution of birds in grasslands recently encroached by forest patches. We investigated how species’ biological traits influenced their response to vegetation change near forest edges. We censured birds along 300‐m line transects run into the open habitat perpendicularly to forest edges. We recorded habitat variables and landscape context along each transect and characterized edges and forest patches. We recorded 33 bird species in 153 transects for a total of 654 individuals. We analyzed species response to edges with generalized linear mixed models. Habitat preference was prevalent to explain species response to forest edges. The abundance of open‐habitat birds such as skylark Alauda arvensis decreased significantly in the vicinity of edges. This negative response extended within 150 m from the edge. The effect was disproportionately higher in open‐habitat species with high conservation concern. The abundance of species feeding or/and breeding in both forest and open habitat, such as woodlarks Lullula arborea, sharply increased near edges (positive edge response). Abundance of shrub and non‐shrub dependent species increased with distance to edge. The two species groups did no differ in abundance/distance to edge relationship. Intensity of species response to forest edges varied among transects in relation to transect vegetation characteristics. Edge length or aspect, diet and nest height had no direct effect. We discuss the possible role of variation in resources and nest predation risk to explain observed patterns.     

Fish habitat modelling as a tool for river management

This paper presents an integrated modelling approach to simulate and assess ecological effects of physical habitat changes in rivers. An ecohydraulic simulation tool was created by combining a 1D hydraulic model based on HEC-RAS software and the fish habitat module of CASiMiR, a fuzzy logic-based ecohydraulic modelling system. This tool was applied on a river stretch commonly occurring in Belgium and elsewhere in Europe. In particular the effect of weir removal on habitat suitability for bullhead (Cottus gobio L.) was simulated. Physical conditions of the studied stretch after weir removal were simulated with a hydraulic model. CASiMiR linked these conditions to ecological expert knowledge to calculate habitat suitability for three life stages of bullhead at four different flow rates based on fuzzy logic. Results indicated that after weir removal, habitat suitability increased significantly for all life stages and all flow rates. The presented approach is promising regarding fish community assessment and ecological river engineering.     

Coexistence and host use by a large community of Pierid butterflies: habitat is the templet

Summary We report studies on the butterfly-hostplant communities in the species-rich area of west central Morocco. Pieridae feeding on Capparales form two distinct ecological guilds: inflorescence feeders and folivores. Several members of each guild may synchronously occur in sympatry. Substantial levels of cannibalism and inter-specific predation occur amongst the inflorescence feeders. No evidence was obtained for segregation of butterfy species on different hosts. Host plants included in the diet of specialists were also used by generalists. Despite substantial differences observed in laboratory trials of larval survivorship on different hostplants, results were congruent for all Pieridae, suggesting that little one-to-one insect-host coevolution has occurred. Host specialization was instead related to the year-to-year stability of host numbers in an area. High levels of pierid infestation occurred on host species with numerically stable populations. Host numerical stability was correlated with habitat type. There is little evidence for segregation of competing inflorescence feeders by hostplant species, but some evidence for segregation by habitat type (particularly by shading levels). We interpret our results as indicating that the hostplant affiliations of Moroccan Capparales-feeding Pieridae are subject to (at best) diffuse coevolutionary effects from hosts and competitors, and are strongly influenced by habitat characteristics.     

The evolution of coloniality in birds in relation to food, habitat, predation, and life-history traits: a comparative analysis

Coloniality in birds has been intensively studied under the cost and benefit approach, but no general conclusion can be given concerning its evolutionary function. Here, we report on a comparative analysis carried out on 320 species of birds using the general method of comparative analysis for discrete variables and the contrast method to analyze the evolution of coloniality. Showing a mean of 23 convergences and 10 reversals, coloniality appears to be a rather labile trait. Colonial breeding appears strongly correlated with the absence of feeding territory, the aquatic habitat, and nest exposure to predators but was not correlated with changes in life-history traits (body mass and clutch size). The correlation of coloniality with the aquatic habitat is in fact explained by a strong correlation with the marine habitat. Unexpectedly, we found that the evolution toward a marine habitat in birds was contingent on coloniality and that coloniality evolved before the passage to a marine life. These results-along with the lack of transitions from the nonmarine to marine habitat in solitary species and the precedence of the loss of feeding territoriality on the passage to a marine life-contradict most of the hypotheses classically accepted to explain coloniality and suggest that we use a different framework to study this evolutionary enigma.     

Woody species colonisation in relation to habitat productivity

A study was conducted to analyse the effect of habitat productivity on woody species colonisation. Three soil types were distinguished: a relatively poor sandy soil type (1), a somewhat richer sandy type (2) and a relatively rich sandy loamy type (3). Chronosequences were established on these three soil types of 38 (type 1), 20 (type 2) and 54 years (type 3) after abandonment. In total 117 vegetation relevées were used to analyse life form change and species responses during old field succession via regression models. On the rich soil type the colonisation rate of woody species was slower than on the poor soil type. This can be explained by higher abundances of perennial species during the first 20 years after abandonment on the rich soil type in contrast to the poor soil type. Perennial species may delay the woody species colonisation. First they close the bare ground which inhibits germination and next they compete with woody seedlings for light, water and nutrients. The effect of habitat productivity on woody species colonisation can only be determined appropriately by taking life history traits into account. Early successional 'pioneer' woody species dispersed by wind have less difficulties colonising old fields than late successional 'forest' species; they colonise old fields prior to the development of a dense perennial sward. Forest species depend on animals to be dispersed which are attracted by vegetation structure. In ± 30 years on the poor soil type and in ± 45 years on the rich soil type woody species become dominant relative to other life forms. Forest species like Quercus robur L. invaded relatively early (<5 years) in contrast to other studies which probably coincides with the distance to seed sources (forest edges).     

Community structure and the habitat templet: ants in the tropical forest canopy and litter

The tropical forest canopy and litter differ in physical structure, resource availability, and abiotic conditions. We used standardized bait experiments in the canopy and litter of four neotropical tree species to explore how these differences shape the behavior, morphology, and diversity of ant assemblages. Ant activity (biomass at a bait after 32 min) was higher in the canopy, and higher on protein baits than carbohydrate baits. Aggressive bait defense occurred more frequently in the canopy (60%) than in the litter (32%), but was not associated with tree species or bait type in either habitat. The median size of workers of species in the canopy and litter was nearly identical, but body size distribution was unimodal in the canopy and bimodal in the litter. The colony size of the most aggressive species was an order of magnitude larger in the canopy. Species richness at a bait was relatively uniform across tree species and habitats. Litter and canopy shared no species, but overlap among tree species was three times higher in the litter assemblages. Litter assemblages showed less activity, less interference, less differentiation across the landscape, and different size distributions than canopy assemblages. The canopy and litter templets subsume a number of environmental gradients that combine to shape ant community structure.     

河流生物栖息地调查及评估方法

河流生物栖息地的研究是河流生态修复和流域环境管理的重要基础.本文概述了河流生物栖息地的分类、调查方法（包括调查范围、采样密度、采样季节和数据记录方式）以及调查数据处理等;并从物种与栖息地适宜性关系的角度出发,对目前国际上主要的栖息地评估方法（评估模型）的应用特点和应用范围进行了评述;最后,在探讨发达国家关于栖息地研究现状的基础上,提出了我国河流治理应从生物栖息地角度出发进行生态恢复建设.     

Performance of bottom ramps to mitigate gravel habitat bottlenecks in a channelized lowland river

The long‐term performance of measures to restore in‐stream habitat in gravel bed rivers is uncertain in the presence of impoundments, land use pressures, and fine sediment inputs. The goal of this study was to evaluate the longer‐term performance of five bottom ramps, designed to facilitate fish passage, and constructed similarly to artificial riffles to provide compensatory gravel riverbed habitat for benthic invertebrates and lithophilic, coarse‐substrate‐preferring fish in a channelized lowland river. Bottom ramp age did not significantly influence habitat conditions indicated by a lack of correlations with the percentage of fine sediment less than 2 mm, the organic content of the substrate, and the years since construction. A significant decrease in the relative abundances of coarse‐substrate‐preferring benthic invertebrates corresponding to project construction age was found, but there were no significant differences in the density of rheophilic, fast‐flow‐preferring or lithophilic fish species among sites. This study presents substantial evidence that similarly constructed bottom ramps in comparable environmental settings provided sufficient habitat for sensitive benthic invertebrates and fish to be present for over 13 years. However, a sudden decrease in habitat suitability cannot be excluded in the long‐term if there is a fine sediment deposition threshold, which results in ramps becoming full. Nevertheless, bottom ramps are recommended as effective measures to enhance longitudinal connectivity, fish passage, and gravel habitat provision in channelized lowland rivers.