Ecosystem response to earlier ice break‐up date: Climate‐driven changes to water temperature,lake‐habitat‐specific production,and trout habitat and resource use

Climate warming has yielded earlier ice break‐up dates in recent decades for lakes leading to water temperature increases, altered habitat, and both increases and decreases to ecosystem productivity. Within lakes, the effect of climate warming on secondary production in littoral and pelagic habitats remains unclear. The intersection of changing habitat productivity and warming water temperatures on salmonids is important for understanding how climate warming will impact mountain ecosystems. We develop and test a conceptual model that expresses how earlier ice break‐up dates influence within lake habitat production, water temperatures and the habitat utilized by, resources obtained and behavior of salmonids in a mountain lake. We measured zoobenthic and zooplankton production from the littoral and pelagic habitats, thermal conditions, and the habitat use, resource use, and fitness of Brook Trout (Salvelinus fontinalis). We show that earlier ice break‐up conditions created a "resource‐rich" littoral–benthic habitat with increases in zoobenthic production compared to the pelagic habitat which decreased in zooplankton production. Despite the increases in littoral–benthic food resources, trout did not utilize littoral habitat or zoobenthic resources due to longer durations of warm water temperatures in the littoral zone. In addition, 87% of their resources were supported by the pelagic habitat during periods with earlier ice break‐up when pelagic resources were least abundant. The decreased reliance on littoral–benthic resources during earlier ice break‐up caused reduced fitness (mean reduction of 12 g) to trout. Our data show that changes to ice break‐up drive multi‐directional results for resource production within lake habitats and increase the duration of warmer water temperatures in food‐rich littoral habitats. The increased duration of warmer littoral water temperatures reduces the use of energetically efficient habitats culminating in decreased trout fitness.     

Seasonal patterns of sediment loading and benthic invertebrate community dynamics in Lake Tanganyika, Africa

1. This study investigated the spatiotemporal dynamics of benthic invertebrate communities located close to the mouths of the Kalambo and Lunzua Rivers, at the southern end of Lake Tanganyika, and subject to varying degrees of sediment load. 2. Metrics of exposure to riverine sediments were associated inversely with abundance and diversity of benthic organisms at sampling locations adjacent to both river mouths, indicating a major effect of riverine sediments on the structure of near‐shore benthic communities. 3. A lower abundance of benthos was found at the mouth of the Lunzua River, which exports significantly higher sediment loads than the Kalambo. 4. Seasonal cycles of abundance and diversity in the benthos varied with distance from river mouths. This has important implications for monitoring programmes investigating benthic communities close to the mouths of rivers. 5. Our findings suggest that recent increases in the sediment loading of Lake Tanganyika, owing to anthropogenic catchment disturbance, impact significantly on biological diversity throughout the lake littoral, which provides habitat for the majority of species in the lake.     

Niche segregation of coexisting Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) constrains food web coupling in subarctic lakes

1. Generalist fish species are recognised as important couplers of benthic and pelagic food‐web compartments in lakes. However, interspecific niche segregation and individual specialisation may limit the potential for generalistic feeding behaviour. 2. We studied summer habitat use, stomach contents and stable isotopic compositions of the generalist feeder Arctic charr coexisting with its common resource competitor brown trout in five subarctic lakes in northern Norway to reveal population‐level and individual‐level niche plasticity. 3. Charr and trout showed partial niche segregation in all five lakes. Charr used all habitat types and a wide variety of invertebrate prey including zooplankton, whereas trout fed mainly on insects in the littoral zone. Hence, charr showed a higher potential to promote habitat and food‐web coupling compared to littoral‐dwelling trout. 4. The level of niche segregation between charr and trout and between pelagic‐caught and littoral‐caught charr depended on the prevailing patterns of interspecific and intraspecific resource competition. The two fish species had partially overlapping trophic niches in one lake where charr numerically dominated the fish community, whereas the most segregated niches occurred in lakes where trout were more abundant. 5. In general, pelagic‐caught charr had substantially narrower dietary and isotopic niches and relied less on littoral carbon sources compared to littoral‐caught conspecifics that included generalist as well as specialised benthivorous and planktivorous individuals. Despite the partially specialised planktivorous niche and thus reduced potential of pelagic‐dwelling charr to promote benthic–pelagic coupling, the isotopic compositions of both charr subpopulations suggested a significant reliance on both littoral and pelagic carbon sources in all five study lakes. 6. Our study demonstrates that both interspecific niche segregation between and individual trophic specialisation within generalist fish species can constrain food‐web coupling and alter energy mobilisation to top consumers in subarctic lakes. Nevertheless, pelagic and littoral habitats and food‐web compartments may still be highly integrated due to the potentially plastic foraging behaviour of top consumers.     

Depth-Related Effects on a Meiofaunal Community Dwelling in the Periphyton of a Mesotrophic Lake

Periphyton is a complex assemblage of micro- and meiofauna embedded in the organic matrix that coats most submerged substrate in the littoral of lakes. The aim of this study was to better understand the consequences of depth-level fluctuation on a periphytic community. The effects of light and wave disturbance on the development of littoral periphyton were evaluated in Lake Erken (Sweden) using an experimental design that combined in situ shading with periphyton depth transfers. Free-living nematodes were a major contributor to the meiofaunal community. Their species composition was therefore used as a proxy to distinguish the contributions of light- and wave-related effects. The periphyton layer was much thicker at a depth of 30 cm than at 200 cm, as indicated by differences in the amounts of organic and phototrophic biomass and meiofaunal and nematode densities. A reduction of the depth-level of periphyton via a transfer from a deep to a shallow location induced rapid positive responses by its algal, meiofaunal, and nematode communities. The slower and weaker negative responses to the reverse transfer were attributed to the potentially higher resilience of periphytic communities to increases in the water level. In the shallow littoral of the lake, shading magnified the effects of phototrophic biomass erosion by waves, as the increased exposure to wave shear stress was not compensated for by an increase in photosynthesis. This finding suggests that benthic primary production will be strongly impeded in the shallow littoral zones of lakes artificially shaded by construction or embankments. However, regardless of the light constraints, an increased exposure to wave action had a generally positive short-term effect on meiofaunal density, by favoring the predominance of species able to anchor themselves to the substrate, especially the Chromadorid nematode Punctodora ratzeburgensis.     

Morphological divergence between three Arctic charr morphs – the significance of the deep‐water environment

Morphological divergence was evident among three sympatric morphs of Arctic charr (Salvelinus alpinus (L.)) that are ecologically diverged along the shallow‐, deep‐water resource axis in a subarctic postglacial lake (Norway). The two deep‐water (profundal) spawning morphs, a benthivore (PB‐morph) and a piscivore (PP‐morph), have evolved under identical abiotic conditions with constant low light and temperature levels in their deep‐water habitat, and were morphologically most similar. However, they differed in important head traits (e.g., eye and mouth size) related to their different diet specializations. The small‐sized PB‐morph had a paedomorphic appearance with a blunt head shape, large eyes, and a deep body shape adapted to their profundal lifestyle feeding on submerged benthos from soft, deep‐water sediments. The PP‐morph had a robust head, large mouth with numerous teeth, and an elongated body shape strongly related to their piscivorous behavior. The littoral spawning omnivore morph (LO‐morph) predominantly utilizes the shallow benthic–pelagic habitat and food resources. Compared to the deep‐water morphs, the LO‐morph had smaller head relative to body size. The LO‐morph exhibited traits typical for both shallow‐water benthic feeding (e.g., large body depths and small eyes) and planktivorous feeding in the pelagic habitat (e.g., streamlined body shape and small mouth). The development of morphological differences within the same deep‐water habitat for the PB‐ and PP‐morphs highlights the potential of biotic factors and ecological interactions to promote further divergence in the evolution of polymorphism in a tentative incipient speciation process. The diversity of deep‐water charr in this study represents a novelty in the Arctic charr polymorphism as a truly deep‐water piscivore morph has to our knowledge not been described elsewhere.     

Lake morphometry predicts the degree of habitat coupling by a mobile predator

Habitat coupling is an ecosystem process whereby semi-discontinuous habitats are connected through the movement of energy and nutrients by chemical, physical or biological processes. One oft-cited example is that of littoral–pelagic coupling in lakes. Theory has argued that such habitat coupling may be critical to food web dynamics, yet there have been few empirical studies that have quantified ecological factors that affect the degree of habitat coupling in ecosystems. Specifically, the degree to which habitat coupling occurs across important physical gradients has largely been ignored. To address this, we investigate the degree of littoral habitat coupling (i.e. the degree to which a top predator lake trout, Salvelinus namaycush, derives energy from the littoral zone) along a gradient of lake shape, where lake shape modifies the relative quantity of coupled epilimnetic benthic and pelagic habitats within each lake. Herein we demonstrate that littoral habitat coupling is intensified in simple circular lakes compared to their reticulate counterparts in seven Canadian Shield lakes. Although the more reticulate lakes had larger areas of epilimnetic benthic habitat, littoral food sources comprised 11% compared to 24% of lake trout diet in reticulate and circular lakes, respectively. This heightened interaction in circular lakes also appears to translate into increased omnivory in more circular lakes compared to reticulate lakes such that lake trout of circular lakes have a significantly lower trophic position than lake trout of reticulate lakes (F_1,5=6.71 p=0.05). These results suggest that it is the accessibility of littoral production via thermal refugia, and not the amount of littoral production, that determines the degree to which lake trout couple littoral and pelagic habitats in lakes.     

Combined influence of river discharge and wind on littoral nematode communities of a river mouth area of Lake Constance

The littoral nematode community adjacent to a river mouth (River Schussen) on Lake Constance (Germany) was studied from February 1999 to January 2000 in order to determine the influence of stress resulting from fluctuations in river discharge on local nematode assemblages. Additionally, the influence of wind as a second important stress factor was considered. Six sample sites were chosen, reflecting a gradient of river influence within the broader river mouth area. Nematode communities, varying in a mean range from 121 to 165 ind/10 cm², were found to differ significantly in terms of abundance, feeding type composition and species diversity. Deposit feeders were most abundant at all sites followed by chewers. Deposit feeders were affected mainly by wind events, while species diversity and the occurrence of chewers were influenced mainly by river discharge. The impact of both these stress factors was modified by a third variable, water level. Moderate and high levels of combined habitat stress led to significant changes in community structure. Under conditions of calm weather and low discharge, reduced species diversity and an increased predominance of deposit feeders were observed. In most cases, species diversity was found to be higher under moderate stress conditions, an observation that offers support for Connel’s Intermediate Disturbance Hypothesis.     

Responses of Deep‐Sea Meiobenthic Communities to Sediment Disturbance Simulating Effects of Polymetallic Nodule Mining

During a benthic impact experiment (BIE) carried out during 1995–1997 by the Interoceanmetal Joint Organization (IOM) at an abyssal site in the North‐East Pacific, sediment disturbance mimicking that resulting from polymetallic nodule extraction was created with a specialised device (the Benthic Disturber). The effects of the disturbance on meiobenthic communities were assessed immediately after disturbance and 22 months later. A reduction in meiobenthos abundance, observed immediately following impact, was not significant; neither were changes in composition of the meiobenthos which was dominated by nematodes and harpacticoids. The lack of any significant numerical response is probably accounted for by the moderate degree of disturbance in this study, compared with other BIE‐type experiments. On the other hand, statistically significant changes in both meiofauna abundance and vertical distribution profiles in the changed sediment within the Disturber tracks were recorded. After 22 months, a significant increase in overall meiobenthos abundance was detected in that part of the test site affected by increased resuspended sediment settlement and receiving natural phytodetrital inputs. Certain taxon‐specific responses on the part of nematodes and harpacticoids were noted both immediately after the disturbance and 22 months later. They were explained by the effects of sediment physical reworking and responses to phytodetrital enrichment. The results presented should aid in developing experimental designs, on both temporal and spatial scales, of future deep‐sea tests aimed at assessing the scale and consequences of man‐made impacts.     

Interim Responses of Benthic and Snag‐Dwelling Macroinvertebrates to Reestablished Flow and Habitat Structure in the Kissimmee River,Florida, U.S.A.

A critical component in the effort to restore the Kissimmee River ecosystem is the reestablishment of an aquatic invertebrate community typical of free‐flowing rivers of the southeastern United States. This article evaluates early responses of benthic and snag‐dwelling macroinvertebrates to restoration of flow and habitat structure following Phase I construction (interim period) of the Kissimmee River Restoration Project. Replicate benthic and snag samples were collected from remnant river channels in Pool A (Control site), and Pool C, the site of the first phase of restoration (Impact site). Samples were collected quarterly for 2 years prior to construction (baseline) and monthly or quarterly for 3 years following Phase I construction and restoration of flow. Baseline benthic data indicate a community dominated by taxa tolerant of organic pollution and low levels of dissolved oxygen, including the dipterans Chaoborus americanus (Chaoboridae) and the Chironomus/Goeldichironomus group (Chironomidae). Baseline snag data indicate a community dominated by gathering‐collectors, shredders, and scrapers. Passive filtering‐collector invertebrates were rare. Following restoration of flow, benthic invertebrate communities are numerically dominated by lotic taxa, including bivalves and sand‐dwelling chironomids (e.g. Polypedilum spp., Cryptochironomus spp., and Tanytarsini). Snags within the Phase I area support an invertebrate community dominated by passive filtering‐collectors including Rheotanytarsus spp. (Chironomidae) and Cheumatopsyche spp. (Hydropsychidae). Results indicate that restoration of flow has resulted in ecologically significant changes to the river habitat template not observed in Pool A. Observed shifts in benthic and snag macroinvertebrate community structure support previously developed hypotheses for macroinvertebrate responses to hydrologic restoration.     

The respiration of common benthic invertebrate species from the shallow littoral zone of Lake Esrom,Denmark

To enable estimation of the total assimilation of benthic populations, we measured the oxygen consumption of macroinvertebrates from the littoral zone of a eutrophic lake. The animals were collected all the year round, and their respiration was measured at field temperatures using a closed-bottle method. Multiple regressions relating the rate of oxygen consumption to temperature and body size were established for 22 taxa, including data derived from the literature for meiofaunal groups.     

Gene expression remodelling and immune response during adaptive divergence in an African cichlid fish

Variation in gene expression contributes to ecological speciation by facilitating population persistence in novel environments. Likewise, immune responses can be of relevance in speciation driven by adaptation to different environments. Previous studies examining gene expression differences between recently diverged ecotypes have often relied on only one pair of populations, targeted the expression of only a subset of genes or used wild‐caught individuals. Here, we investigated the contribution of habitat‐specific parasites and symbionts and the underlying immunological abilities of ecotype hosts to adaptive divergence in lake–river population pairs of the cichlid fish Astatotilapia burtoni. To shed light on the role of phenotypic plasticity in adaptive divergence, we compared parasite and microbiota communities, immune response, and gene expression patterns of fish from natural habitats and a lake‐like pond set‐up. In all investigated population pairs, lake fish were more heavily parasitized than river fish, in terms of both parasite taxon composition and infection abundance. The innate immune response in the wild was higher in lake than in river populations and was elevated in a river population exposed to lake parasites in the pond set‐up. Environmental differences between lake and river habitat and their distinct parasite communities have shaped differential gene expression, involving genes functioning in osmoregulation and immune response. Most changes in gene expression between lake and river samples in the wild and in the pond set‐up were based on a plastic response. Finally, gene expression and bacterial communities of wild‐caught individuals and individuals acclimatized to lake‐like pond conditions showed shifts underlying adaptive phenotypic plasticity.     

Salinity and fish effects on Salton Sea microecosystems: benthos

The Salton Sea, the largest lake in California, has a surface elevation 69 m below sea level which is maintained predominantly by the balance of agricultural runoff and evaporation. The lack of outflowing streams is resulting in a gradual buildup of salts in the lake, increasing the salinity. A 15 month microcosm experiment was conducted to determine the effects of salinity and tilapia ( Oreochromis mossambicus) on an assemblage of benthic and planktonic Salton Sea algae and invertebrates. This article reports the responses of the benthic invertebrates. Microcosms (312 l fiberglass tanks) were set up without tilapia at 30, 39, 48, 57, and 65 g · l-1. Additional microcosms were set up with tilapia at 39 and 57 g · l-1. In the absence of fish Gammarus mucronatus dominated the benthos at the lower salinities, and Trichocorixa reticulata and the larvae of Ephydra riparia were most abundant above 48 g · l-1. The most abundant meiofaunal species included the harpacticoid copepod. Cletocamptus deitersi, three nematodes, the rotifer Brachionus plicatilis, ciliates, including Condylosoma sp. and Fabrea salina, two foraminiferans including Quinqueloculina sp., and a large flagellate. Most meiofaunal species responding to salinity were most abundant at 65 g · l-1, especialy after 6 months when Gammarus dominated the lower salinities. The tilapia reduced the abundance of macrofaunal species, especially at 39 g · l-1, and generally increased the abundance of meiofaunal species and ciliates. The microcosm benthic macro- and meiofaunal communities were most likely structured by Gammarus, salinity and tilapia. Gammarus reduced the other species by predation and changing the detritus from an algal base to a fecal pellet base. Gammarus was itself reduced by tilapia and by reduced reproductive success above 39 g · l-1. More species were therefore able to compete at higher salinities and in the presence of tilapia. Tilapia also affected the benthos by depositing loosely packaged fecal material which may support more meiofaunal species than either the robust Gammarus fecal pellets that were abundant at 39 g · l-1 or the algae-fecal pellet mix at 57 g · l-1. This revised version was published online in July 2006 with corrections to the Cover Date.     

Importance of scalar and riparian habitat effects for assessment of ecological status using littoral diatoms

The classification of waterbodies under the Water Framework Directive is dependent on the ability of monitoring programmes to reflect habitat quality using biotic elements including benthic diatom communities. This study investigated the influence of specific riparian habitats, of mixed woodland, grassland and lake artificial structures such as jetties and slipways, on benthic diatom assemblages in nine lakes across gradients of total phosphorus, alkalinity and in the presence or absence of Dreissena polymorpha. The heterogeneity of the benthic diatom assemblages at riparian and lake scale was assessed by taking three replicates per site category per lake, following standard European Union protocols. Canonical correspondence analysis (CCA) and mixed effect modelling was used to investigate the main environmental controls on assemblage structure. Non-metric Multidimensional Scaling (NMDS) was used to examine patterns in assemblage structure. No single environmental gradient was found to control benthic diatom composition, with differences among assemblages influenced both by riparian habitat type within lakes and interaction of multiple environment gradients, including presence of D. polymorpha. Greater control was exerted on community structure at the lake than local riparian scale. The influence of scalar factors on diatom assemblages increased with increasing scale. We recommend that for effective monitoring and assessment of ecological status, standard sampling protocols should include localised littoral habitats with individual samples pooled across riparian habitat types, thereby accounting for both multiple environmental and spatial controls on community structure.     

The effects of predation by juvenile fish on the meiobenthic community structure in a natural pond

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An Ecosystem Model for Exploring Lake Restoration Effects on Fish Communities and Fisheries in Florida

Developing quantitative ecosystem–scale expectations of habitat restoration projects and examining trade‐offs associated with alternative approaches has been a challenge for restoration ecology. Many of the largest freshwater lake restoration projects have occurred in Florida to remediate degradation to vegetated littoral habitats resulting from stabilized water levels, but effects across lake food‐webs have not been assessed. We developed an ecosystem model using Ecopath with Ecosim and Ecospace for a generalized large, eutrophic Florida Lake to explore how simulated restoration activities could influence fish communities with emphasis on sport fish abundance. We modeled three habitat restoration scenarios: (1) “no control,” (2) a “10‐year control” that restored littoral habitat every 10 years, and (3) a “combined control” scenario that restored littoral habitat every 10 years with maintenance controls between 10‐year periods. Our “combined control” scenario provided the largest long‐term habitat restoration benefits for sport fish abundance and the fisheries they support. In Ecospace, we simulated a littoral habitat restoration project that reduced lake‐wide tussock coverage from 30 to 15%. Ecospace predicted positive benefits to sport fish and fisheries following the restoration simulation and highlighted the importance of habitat edge effects, spatial design of habitat restoration projects, and sampling designs for evaluating restoration projects.     

Seasonal and diel patterns of invertebrate drift in different alpine stream types

1. We examined the seasonal and diel patterns of invertebrate drift in relation to seston and various habitat characteristics in two each of four different kinds of alpine streams [rhithral (snow‐fed) lake outlets, rhithral streams, kryal (glacial‐fed) lake outlets and kryal streams]. Samples were collected at four times of the day (dawn, midday, dusk and midnight) during three seasons (spring, summer and autumn). 2. Habitat characteristics differed mainly between rhithral and kryal sites, with the latter having higher discharge and turbidity, lower water temperature, and higher concentrations of ammonium, and particulate and soluble reactive phosphorus. Seasonality in habitat characteristics was most pronounced for kryal streams with autumn samples being more similar to rhithral sites. 3. The concentration of seston was lowest in the glacial‐influenced lake outlets and slightly higher in the stream sites; no seasonal or diel patterns were evident. 4. The density of drifting invertebrates averaged less than 100 m^?3 and was lowest (<10 m^?3) at three of the four kryal sites. Taxon richness and diversity were lowest at rhithral lake outlets. Chironomidae dominated the drift as well as benthic communities and <30% of benthic taxa identified were found in the drift. 5. Drifting invertebrates showed no consistent seasonal pattern. However, density tended to be highest in spring at rhithral sites and in autumn at kryal sites. No diel periodicity in drift density was found at any site and the lack of diel pattern may be a general feature of high altitude streams. 6. Glacially influenced habitat parameters were a major factor affecting drift in these alpine streams, whereas no clear differences were observed between streams and lake outlets. Our findings indicate that invertebrate drift in alpine streams is primarily influenced by abiotic factors, and therefore, substantially differs from patterns observed at lower altitude.     

Permian macroburrows as microhabitats for meiofauna organisms: an ancient behaviour common in extant organisms

Meiofaunal organisms are indirectly influenced by the activity of benthic macroinvertebrates within the sediment, which plays a role in modifying physical and chemical characteristics of the habitat. The association of meiofaunal organisms and macroburrows is well known in modern environments, but the record of this relationship in the geological record is still incipient. This study documents diminutive burrows (Helminthoidichnites tenuis) associated with the surface of macroburrows (Palaeophycus tubularis) in Early Permian deposits. The cylindrical shape and meandering to loop trajectory of the diminutive burrows indicate that they were produced by small free‐living meiofaunal nematodes. Apparently, P. tubularis (open burrow) constituted a favourable microhabitat for nematodes, providing the following: (1) protection against erosive processes and meiofauna predators; (2) oxygen access to more in‐depth layers within the sediment; (3) temperature stabilization; and (4) food supply due to mucus impregnation in the macroburrow walls by the Palaeophycus tracemaker. The association between H. tenuis and P. tubularis constitutes the first fossil record of a symbiotic relationship between meiofaunal nematodes and macrobenthic organisms (polychaetes). It also suggests that ecological strategies such as mutualism or commensalism, which are common between extant nematodes and macrobenthic invertebrates, were available in the behavioural programme of these organisms since the Early Permian.     

Some like it deep: Intraspecific niche segregation in ruffe (Gymnocephalus cernua)

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The lentic and lotic characteristics of habitats determine the distribution of benthic macroinvertebrates in Mediterranean rivers

1. The importance of flow‐related factors to benthic organisms, as well as the role of habitat conditions in shaping aquatic communities during low‐flow periods, have been recognised. Despite this, the preferences of macroinvertebrates to the ratio of lentic to lotic habitats at the reach scale have not been accurately quantified in most instances.
2. Aquatic invertebrates and habitat features in a range of temporary rivers in Sardinia were investigated. The investigation focused on the flow‐related characteristics that contribute to defining the lentic–lotic condition of the river reaches. The relation of habitat features to benthic taxa distributions was assessed using multidimensional scaling. The main aim of the paper was to quantify the responses of taxa to the different lentic and lotic habitat conditions by applying hierarchical logistic regressions. Finally, taxon optima were aligned along the lentic–lotic gradient and the responses of different taxonomic groups compared.
3. Unbroken waves and imperceptible flow were correlated with benthic taxa variability, suggesting local hydraulics and turbulence have a major role in regulating community composition. The overall lentic–lotic character of the river reaches was also clearly related to the benthic taxa distribution. More than 80% of taxa were significantly related to the lentic–lotic gradient, and an asymmetrical response curve was the predominant model.
4. Benthic groups showed taxon optima clustered in different ranges of the lentic–lotic gradient. Odonata, Coleoptera, Hemiptera, and Mollusca preferred clearly lentic conditions. Diptera mainly ranged on the lotic side of the gradient, while Trichoptera were relatively uniformly distributed across the gradient. Ephemeroptera taxa clustered in intermediate lentic–lotic conditions, with two species preferring extremely lentic habitats. In general, optima converged at intermediate and extremely lentic conditions, presumably due, respectively, to the coexistence of different lentic and lotic features and to the highly diverse environmental characteristics under extremely lentic situations.
5. These results support the conclusion that dissimilar ecological factors act on benthic taxa along the lentic–lotic range and species favouring different lentic–lotic conditions are subjected to pressures of different nature. This should not be ignored when defining species preferences and studying community structure or relationships between species in Mediterranean rivers, which cyclically vary their habitat composition. In addition, the uneven distribution of optima of different groups along the lentic–lotic gradient might affect macroinvertebrate metrics when assessing ecological status or establishing reference conditions under variable climatic conditions.

     

Species–area relationship within benthic habitat patches of a tropical floodplain river: An experimental test

The curvilinear relationship between species richness and habitat area (species–area relationship (SAR)) is a fundamental ecological pattern. The relationship is often viewed from a long‐term perspective across relatively large spatial scales, reflecting a balance between immigration and extinction dynamics. We explored whether predictions of SAR also manifest over short time periods (days) in benthic habitat patches of a dynamic floodplain river where littoral faunal assemblages are continuously assembled and disassembled with changing water levels. We examined the relationship of patch size with faunal abundance (i.e. fish and aquatic invertebrates), taxonomic richness, trophic group richness and overall assemblage composition. Strong taxa–area relationships emerged despite the relatively short experimental time period (21 days); larger patches had more taxa and trophic groups. For the smallest patches, taxonomic richness was especially sensitive to abundance of individuals; abundance of individuals was a less important predictor of taxonomic and trophic group richness for the largest patches. Despite the relatively short time frame for study within this temporally dynamic ecosystem, our findings indicate a strong SAR for fishes and macroinvertebrates inhabiting patchy habitats in the littoral zone of this tropical river.