Does biofilm origin matter? Biofilm responses to non-flow period in permanent and temporary streams

1. In some regions, climate change is increasing the variability of rainfall and the frequency of extreme events such as drought. Consequently, non-flow periods have grown in length and frequency, both in temporary and in formerly permanent streams. Water abstraction for human use may further prolong these dry periods.
2. We analysed the resistance and resilience of biofilms from permanent and temporary streams to non-flow conditions. This was achieved by exposing cobbles (collected from permanent and temporary streams) with intact biofilm to 31 days of non-flow, followed by 20 days of stream flow in artificial stream channels. Biofilm resistance and resilience were assessed at a structural (algal biomass, pigment composition, and algae and cyanobacteria composition) and functional level (photosynthetic efficiency and community metabolism).
3. Algal taxa in biofilms from permanent and temporary streams differed throughout the experiment. Biofilms from permanent streams were less resistant to non-flow than those from temporary streams at structural level. Permanent stream biofilms also presented lower resilience at a structural level, but responded similarly to temporary stream biofilms at a functional level.
4. Our investigation shows how the non-flow period disturbed permanent stream biofilms, and suggests that temporary stream biofilms will have greater adaptive capacity as hydroperiod becomes shorter due to climate change.

     

Putting the meio- into stream ecology: current findings and future directions for lotic meiofaunal research

• 1 There is a paucity of research on epigean freshwater lotic meiofauna. This may result from a previous emphasis on interstitial (groundwater and hyporheic) meiofauna and/or a reliance on sampling methodologies in lotic systems which are inappropriate for meiofauna.
• 2 Meiofauna contribute much to the diversity of lotic ecosystems. Species lists for seven streams reveal that meiofauna contribute 58–82% of total species numbers, with rotifers and chironomids dominating most systems. The absence of taxonomic keys for most meiofaunal taxa in large areas of the world precludes a wider analysis of their contribution to lotic diversity and an assessment of biogeographical patterns and processes.
• 3 The trophic and functional role of meiofauna in lotic ecosystems is unclear. There are few estimates of meiofaunal production in freshwaters and biomass spectra have produced conflicting results for lotic meiofauna. Present static estimates suggest that the contribution of meiofauna to lotic productivity and biomass is small to moderate, but further studies incorporating a temporal component may provide a more realistic picture of the total contribution of meiofauna to biomass size spectra.
• 4 Meiofauna differ from macroinvertebrates in several respects apart from size and conceptual models for lotic ecosystems should include all metazoans if they are to be truly representative.
• 5 Information on the basic ecology of certain lotic meiofauna (i.e. nematodes, tardigrades, microturbellarians) is urgently required. For those groups whose distributional patterns are better understood (e.g. microcrustaceans), the mechanisms underpinning these patterns should be explored. It is essential that the importance of meiofauna is recognised by lotic ecologists; the only realistic way forward is for greater collaboration among meiofaunal ecologists and taxonomists and other lotic scientists.

     

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

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Global and regional patterns in lotic meiofauna

• 1 Parsimony analysis of endemicity (PAE) was used to assess patterns in the distribution of harpacticoid copepods (all freshwater species and stream species only) at global and regional scales. These analyses provided a focus for reviewing large scale patterns and processes in freshwater meiofauna.
• 2 On a global scale, PAE suggested that large‐scale biogeographical events have been most important in shaping present‐day distributions in the Canthocamptidae. A small proportion (4%) of canthocamptid species were widespread (i.e. occurred in more than one biogegraphical region), suggesting that dispersal may also play a role in determining distribution at the species level. Global distribution patterns for other meiofauna suggest varying roles for dispersal and vicariant events. No consistent latitudinal trends in species diversity were evident, although a lack of distributional data for many regions, and uncertainty over the status of many cosmopolitan species, precludes more robust analyses. Molecular techniques should prove useful in identifying truly cosmopolitan taxa.
• 3 On a regional scale, a PAE within Western Europe demonstrated a clear link between the distribution of canthocamptid species and the extent of the Last (Wiechselian) glaciation. Northern and southern areas of Europe contain distinctive harpacticoid faunas and the recolonisation of northern Europe appears to have been from the Balkans rather than other Mediterranean peninsulae. The high harpacticoid diversity in southern Europe, may reflect a lack of glacial disruption of groundwater habitats.
• 4 A PAE of lotic data for harpacticoid copepods within the Holarctic reflected the global PAE for freshwater harpacticoids as a whole, but not the regional PAE. A high proportion of stream‐dwelling harpacticoids are widespread species, but only one (Bryocamptus zschokkei) was found in streams across the Holarctic. Other cosmopolites were restricted to streams in Europe or North America, suggesting that species‘ niche requirements might differ among regions. There appeared to be some convergence in the composition of lotic copepod communities in terms of the number of species within genera.
• 5 We conclude that large‐scale processes inevitably have a major influence on the local composition of lotic meiofaunal communities, but that the relative importance of small scale vs. large scale processes is unclear at present, largely due to a paucity of suitable data.

     

What drives small‐scale spatial patterns in lotic meiofauna communities?

• 1 Lotic meiofaunal communities demonstrate extremely variable dynamics, especially when viewed at small spatial scales (≤ metres). Given the limited amount of research on lotic meiofauna, we chose to organise our discussion of their small‐scale spatial patterns around the dominant factors we believe drive their spatial distributions in streams. We separate scale‐dependent effects that structure lotic meiofauna into biotic factors (e.g. predation, food quantity/quality, dispersal) and abiotic factors (e.g. local flow dynamics and substratum characteristics).
• 2 The impact of predation on the distribution of meiofauna varies with the scale over which predators forage (e.g. fish predation influences meiofauna in different ways and at broader spatial scales than do invertebrate predators), the type of streambed substrata in which the predator‐prey interactions occur, and the dispersal ability of different meiofauna. The latter is greatly influenced by predator and prey (meiofauna) interactions with the flow environment.
• 3 Organic matter influences the small‐scale distribution of meiofauna in streams. Both its quality as food (as indicated by C:N content, ATP content, or microbial biomass) and its spatial distribution on the streambed, influence meiofauna patchiness, community structure and life history characteristics. As a habitat, the structure that organic matter provides (e.g. wood or leaves) can influence predator‐prey interactions, offer materials for case‐building and offer refugia during disturbance events ‐ all of which influence the small‐scale spatial distribution of meiofauna.
• 4 Stream flow influences the distribution of meiofauna at broad scales (10s–100s of metres), primarily because of the high susceptibility of meiofauna to passive drift; small‐scale interactions between flow and substrata are also important, however, particularly at more localised (≤ metre) scales. At both scales, substratum particle size is important to interstitial‐dwelling fauna, influencing the probability of passive drift by meiofauna as well as local microhabitat conditions (e.g. dissolved oxygen; upwelling/downwelling in the hyporheic zone) and, thus, the small‐scale distribution among microhabitats.
• 5 In general, the processes governing the distribution of meiofauna at small scales cannot be separated entirely from those processes working at larger scales. A conceptual diagram is presented illustrating the relative importance of various factors in influencing the spatial patterns of meiofauna and over what scales these factors act.

     

Wide variation in a tiny space: The microdistribution of meiobenthos in an artificial pond

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An unexpected source of invertebrate community recovery in intermittent streams from a humid continental climate

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Distribution and feeding of the predatory nematode Anatonchus dolichurus (Mononchoidea) in the Dokka delta (Norway) and its impact on the benthic meiofauna

• 1 A dense population of a large (over 6.5mm long), semiaquatic, predatory nematode Anatonchus dolichurus was found in the delta of the River Dokka, Norway. This is the first time it has been reported in surface fresh waters.
• 2 Nematode distribution was related to water depth, with maximum abundance (over 24000 ind.m^?2) occurring in shallow areas (0.5–2.0m). Sediments at all stations with a high density of A. dolichurus were dry and exposed to air and ice during winter and early spring, and overgrown with macrophytes during summer.
• 3 This predator contributed a greater fraction of total numbers and biomass of the nematode fauna than predatory forms in other known freshwater nematode assemblages. It constituted up to 15.6% of numbers and up to 90% of total nematode biomass.
• 4 Anatonchus dolichurus preyed on Nematoda, small Oligochaeta (mainly Enchyrraeidae) and Chironomidae. It was able to reduce the density of these meiofaunal taxa when present in large numbers in experimental cores.

     

The importance of meiofauna to lotic ecosystem functioning

• 1 Although meiofauna occur in large numbers in many streams, almost nothing is known about their functional role.
• 2 In other systems, meiofauna influence microbial and organic matter dynamics through consumption and bioturbation. Given that these are important processes in streams, meiofauna have the potential to influence lotic function by changing the quality and availability of organic matter as well as the number and biotic activity of benthic microbes. Selective feeding by meiofauna has the potential to alter the availability of nutrients and organic carbon.
• 3 Meiofauna generally contribute only a small amount to metazoan production and biomass in streams, although exceptions occur. Within a stream, the relative importance of meiofauna may reflect whether the temporary or permanent meiofauna dominate the meiobenthos as well as the season when samples are collected.
• 4 We suggest stream conditions (small sediment grain size, restricted interstitial flow) under which meiofauna have the greatest likelihood of influencing stream ecosystem function.
• 5 Important areas for future research include addressing whether meiofauna feed selectively, whether meiofauna are links or sinks for carbon in streams, and whether bioturbation by meiofauna influences stream ecosystem processes in a predictable manner.

     

Parallels and contrasts between intermittently freezing and drying streams: From individual adaptations to biodiversity variation

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Invertebrate assemblages of pools in arid‐land streams have high functional redundancy and are resistant to severe drying

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Taxonomic and functional homogenisation of macroinvertebrate communities in recently intermittent Alpine watercourses

1. Mountain streams in southwestern European Alps are currently shifting from perennial to intermittent flow due to the combined effects of climate change and local anthropogenic pressures. Given that flow intermittency is a recently documented phenomenon in the Alps, only scattered studies have investigated functional and taxonomical diversity of benthic invertebrate communities in recently intermittent Alpine streams.
2. We used a hierarchical sampling design to investigate patterns in taxonomic and functional diversity of benthic invertebrate communities in 13 recently intermittent Alpine streams in north-west Italy. in April 2017, we sampled benthic communities in two reaches of each stream with different hydrological conditions: a control reach, with permanent flow; and an intermittent reach, which recently experienced non-flow periods in summer.
3. We tested for the response of taxonomic richness at multiple spatial scales by partitioning total diversity into the average richness of local communities and the richness due to variation among local communities both within and among reaches. By partitioning total diversity (γ) into its local (α) and turnover (β) components we showed a decrease in local and regional species richness both within and among reaches, whereas variation among communities was significantly lower in intermittent reaches at the reach scale only.
4. The analysis of multidimensional trait space of macroinvertebrate communities in reaches with different hydrological conditions revealed a significant reduction of functional diversity, dispersion, and evenness in intermittent reaches. There was trait overdispersion in intermittent reaches, as these hosted both typical Alpine taxa and organisms adapted to flow intermittency. In particular, we observed the replacement of taxa with aquatic respiration and those preferring medium- to fast-flowing oligotrophic waters by taxa adapted to lentic habitats, air breathing and with larval dormancy phases.
5. These results indicate that recent flow intermittency has caused drastic changes in benthic invertebrate communities in Alpine streams. Our work highlights the importance of integrating taxonomic and functional diversity to thoroughly assess the impacts of flow intermittency.

     

Trophic relationships: integrating meiofauna into a realistic benthic food web

• 1 This paper summarises the most important contributions on trophic relationships of lotic meiofauna. In contrast to marine research, the few quantitative studies of the freshwater meiobenthos have shown that these invertebrates not only take up particulate/fine organic matter, but also dissolved organic substances attached to organic particles. In lotic ecosystems, further estimates of grazing rate and bacterial/algal ingestion rate are needed, particularly in situ measurements.
• 2 The effects of macroinvertebrate predators upon meiofauna are still under debate. Depending on the type of experiments (laboratory vs. field) it seems that macrofauna may or may not affect meiofauna. Field samples and analyses of gut contents of larval tanypod chironomids have shown that the impact upon meiofauna was low and larvae were nonselective predators. Predation amounted to 2.2% of the combined prey density and prey consumption averaged 1.3 individuals per predator individual per year.
• 3 Adding taxonomic resolution by including the meiofaunal component within lotic food webs distinctly increases the number of total species and, as a consequence, changes food web statistics. Webs that included meiofauna revealed that these metazoans contributed substantially to the percentage of intermediate species (species with predators and prey). The resolution of dietary analyses of major consumers of macro‐ and meiobenthos showed that many stream invertebrates feed on meiofauna.

     

Intertidal meiofauna of a high-latitude glacial Arctic fjord (Kongsfjorden, Svalbard) with emphasis on the structure of free-living nematode communities

Meiofauna communities of four intertidal sites, two sheltered and two more exposed, in Kongsfjorden (Svalbard) were investigated in summer 2001 at two different tidal levels (i.e. the low-water line and close below the driftline, referred to as mid-water (MW) level). A total of seven meiofaunal higher taxa were recorded with nematodes, oligochaetes and turbellarians being numerically dominant. Mean total meiofaunal densities ranged between 50 ind. 10 cm⁻² and 903 ind. 10 cm⁻². Our data showed a clear decrease in total meiofaunal densities with increasing coarseness of the sediment. Total meiofaunal biomass varied from 0.2 g dwt m⁻² to 2 g dwt m⁻² and, in general, was high even at low meiofaunal densities, i.e. larger interstitial spaces in coarser sediments supported larger meiofauna, especially turbellarians. The results on the vertical distribution of meiofauna contrasted sharply with typical meiobenthic depth profiles on other beaches, probably in response to ice-scouring and concomitant salinity fluctuations. Oligochaetes were the most abundant taxon, with a peak density of 641 ind. 10 cm⁻² at Breoyane Island. They were mainly comprised of juvenile Enchytraeidae, which prohibited identification to species/genus level. Nematode densities ranged between 4 ind. 10 cm⁻² and 327 ind. 10 cm⁻². Nematodes were identified up to genus level and assigned to trophic guilds. In total, 28 nematode genera were identified. Oncholaimus and Theristus were the most abundant genera. The composition of the nematode community and a dominance of predators and deposit feeders were in agreement with results from other arctic and temperate beaches. Nematode genus diversity was higher at the more sheltered beaches than at the more exposed ones. Low-water level stations also tended to harbour a more diverse nematode communities than stations at the MW level. Differences in nematode community structure between low- and MW stations of single beaches were more pronounced than community differences between different beaches and were mainly related to resources quality and availability.     

Role of meiofauna in estuarine soft-bottom habitats*

Meiofauna are ubiquitous in estuaries worldwide averaging 10⁶ m^?2. Abundance and species composition are controlled primarily by three physical factors: sediment particle size, temperature and salinity. While meiofauna are integral parts of estuarine food webs, the evidence that they are biologically controlled is equivocal. Top down (predation) control clearly does not regulate meiofaunal assemblages. Meiofauna reproduce so rapidly and are so abundant that predators cannot significantly reduce population size. Food quantity (bottom up control) also does not appear to limit meiofaunal abundance; there is little data on the effect of food quality. In estuarine sediments meiofauna: (i) facilitate biomineralization of organic matter and enhance nutrient regeneration; (ii) serve as food for a variety of higher trophic levels; and (iii) exhibit high sensitivity to anthropogenic inputs, making them excellent sentinels of estuarine pollution. Generally mineralization of organic matter is enhanced and bacterial production stimulated in the presence of meiofauna. Tannins from mangrove detritus in northern Queensland appear to inhibit meiofaunal abundance and therefore the role of meiofauna in breakdown of the leaves. Meiofauna, particularly copepods, are known foods for a variety of predators especially juvenile fish and the meiofaunal copepods are high in the essential fatty acids required by fish. The copepod’s fatty acid composition is like that of the microphytobenthos they eat; bacterial eaters (nematodes?) do not have the essential fatty acids necessary for fish. Most contaminants in estuaries reside in sediments, and meiofauna are intimately associated with sediments over their entire life-cycle, thus they are increasingly being used as pollution sentinels. Australian estuarine meiofauna research has been concentrated in Queensland, the Hunter River estuarine system in New South Wales, and Victoria’s coastal lagoons. Studies in northern Queensland have primarily concentrated on the role of nematodes in mineralization of organic matter, whereas those from southern Queensland have concentrated on the role of meiofauna as food for fish and as bacterial grazers. The New South Wales studies have concentrated on the Hunter River estuary and on the structure and function of marine nematode communities. In Victoria, several fish have been shown to eat meiofauna. The Australian world of meiofaunal research has hardly been touched; there are innumerable opportunities for meiofaunal studies. In contaminated estuarine sediments reduced trophic coupling between meiofauna and juvenile fish is a basic ecological question of habitat suitability, but also a question with relevance to management of estuarine resources. Because meiofauna have short lifecycles, the effects of a contaminant on the entire life-history can be assessed within a relatively short time. The use of modern molecular biology techniques to assess genetic diversity of meiofauna in contaminated vs uncontaminated sediments is a promising avenue for future research. Much of the important meiofaunal functions take place in very muddy substrata; thus, it is imperative to retain mudflats in estuaries.     

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.     

Meiofaunal prominence and benthic seasonality in a coastal marine ecosystem

Summary The muds of a shallow (7 m) site in Narragansett Bay, Rhode Island contained higher abundances of meiofauna (averaging 17×10⁶ individuals per m² and ash free dry weight of 2.9 g/m² during a 3 year period) than have been found in any other sediment. The majority of sublittoral muds, worldwide, have been reported to contain about 10⁶ individuals per m². This difference is attributed primarily to differences in sampling techniques and laboratory processing.Extremely high meiofaunal abundances may have also occurred because Narragansett Bay sediments were a foodrich environment. While the quantity of organic deposition in the bay is not unusually high for coastal waters, this input, primarily composed of diatom detritus, may contain an unusually high proportion of labile organics. Furthermore, meiofauna could have thrived because of spatial segregation of meiofauna and macrofauna. While meiofauna were concentrated at the sediment-water interface, most macrofauna were subsurface deposit feeders. Macrofaunal competition with, and ingestion of meiofauna may thus have been minimized.The seasonal cycles of meiofauna and macrofauna were similar. Highest abundances and biomass were observed in May and June and lowest values in the late summer and fall. Springtime increases of meiofaunal abundance were observed in all depth horizons, to 10 cm. We hypothesize that phytoplankton detritus accumulated in the sediment during the winter and early spring, and that the benthos responded to this store of food when temperatures rose rapidly in the late spring. By late summer, the stored detritus was exhausted and the benthos declined.     

Meiobenthos in mangrove areas in eastern Africa with emphasis on assemblage structure of free-living marine nematodes

A survey was conducted to examine spatial variations in the population density of major meiofaunal taxa and the assemblage structure of free-living marine nematodes within 5 mangrove areas on the west and east coast of Zanzibar. Meiofauna densities in surface sediments (0–5 cm) ranged from 205 to 5263 ind. 10 cm², being on average 1493 ind. 10 cm². Of the 17 major taxa recorded, nematodes dominated (64–99%) in all samples while harpacticoid copepods were usually second most abundant. Within all areas the numbers of meiofauna were very variable and significant differences among areas were only detected for oligochaetes and turbellarians. Densities of nematodes, harpacticoids, polychaetes and turbellarians were, however, significantly (P<0.001) higher at low water stations compared with mid and high water stations. Harpacticoids were negatively correlated with the numbers of fiddler crab (Uca spp.) burrows. Other correlations between environmental factors (grain size, temperature, salinity, oxygen tension, prop root density, fiddler crab burrows) and major meiofaunal taxa were non-significant. A total of 94 nematode genera were recorded from four mangrove areas. The most abundant and frequent genera were Microlaimus and Spirinia, followed by Desmodora and Metachromadora. Representatives of the genera most common in current study are found all over the globe. There was a high variation in nematode assemblage structure within and between sampling areas indicating the absence of a well defined nematode assemblage confined to mangrove areas. In a hypersaline area diversity was much reduced and where salinity was over 100%. the fauna was restricted to 3 nematode genera, Microlaimus, Theristus and Bathylaimus. Multidimensional scaling ordination (MDS) of the nematode genera separated samples taken from low water stations from other stations, the assemblage structure being significantly different at the low water stations. Numbers of selective deposit feeders were negatively correlated with average grain size and positively correlated with silt content.     

Ecological effects of perturbation by drought in flowing waters

• 1 Knowledge of the ecology of droughts in flowing waters is scattered and fragmentary, with much of the available information being gathered opportunistically. Studies on intermittent and arid‐zone streams have provided most of the information.
• 2 Drought in streams may be viewed as a disturbance in which water inflow, river flow and water availability fall to extremely low levels for extended periods of time. As an ecological perturbation, there is the disturbance of drought and the responses of the biota to the drought.
• 3 Droughts can either be periodic, seasonal or supra‐seasonal events. The types of disturbance for seasonal droughts are presses and for supra‐seasonal droughts, ramps.
• 4 In droughts, hydrological connectivity is disrupted. Such disruption range from flow reduction to complete loss of surface water and connectivity. The longitudinal patterns along streams as to where flow ceases and drying up occurs differs between streams. Three patterns are outlined: ‘downstream drying’, ‘headwater drying’ and ‘mid‐reach drying’.
• 5 There are both direct and indirect effects of drought on stream ecosystems. Marked direct effects include loss of water, loss of habitat for aquatic organisms and loss of stream connectivity. Indirect effects include the deterioration of water quality, alteration of food resources, and changes in the strength and structure of interspecific interactions.
• 6 Droughts have marked effects on the densities and size‐ or age‐structure of populations, on community composition and diversity, and on ecosystem processes.
• 7 Organisms can resist the effects of drought by the use of refugia. Survival in refugia may strongly influence the capacity of the biota to recover from droughts once they break.
• 8 Recovery by biota varies markedly between seasonal and supra‐seasonal droughts. Faunal recovery from seasonal droughts follows predictable sequences, whilst recovery from supra‐seasonal droughts varies from one case to another and may be marked by dense populations of transient species and the depletion of biota that normally occur in the streams.
• 9 The restoration of streams must include the provision of drought refugia and the inclusion of drought in the long‐term flow regime.

     

Disentangling the multiple effects of stream drying and riparian canopy cover on the trophic ecology of a highly threatened fish

1. Understanding risks to aquatic systems posed by changing drought regimes is particularly important for the conservation of already threatened taxa. However, little is known about how local environmental conditions, especially those in heavily human‐influenced situations, interact with regional shifts such as droughts to alter realised impacts on aquatic communities, including threatened top predators.
2. Here, we investigated the combined effects of stream drying intensity and riparian canopy cover on the trophic interactions of critically endangered kōwaro, or Canterbury mudfish (Neochanna burrowsius) in an agricultural area of New Zealand. Fish populations and their potential prey, both terrestrial and aquatic, as well as environmental variables, including riparian canopy cover and drying measured with stage loggers, were sampled over eight visits to 24 sites spanning orthogonal drying and canopy gradients. Stable isotope ratios, ¹³C/¹²C and ¹⁵N/¹⁴N, were used to investigate trophic links between mudfish and their terrestrial and aquatic prey across these gradients.
3. When non‐native willows (predominantly Salix fragilis) dominated the riparian canopy, increased tree cover led to elevated drying intensity, probably driven by their relatively high water demands compared to other trees. However, in the absence of willows, canopy cover had no effect on drying intensity. Although this was the only direct link between these two environmental factors, they had opposing effects on kōwaro populations, which will be important for management under drought.
4. Increased drying intensity contributed to elevated abundance of microcrustacea and aquatic Diptera larvae, and an increase in the relative abundance of kōwaro juveniles. However, drying‐affected kōwaro populations also had fewer large reproductive adults and elevated δ¹⁵N values, probably driven by physiological limitations and an increase in kōwaro cannibalism, respectively.
5. By comparison, increased canopy cover enhanced input of terrestrial invertebrates, a food resource for larger kōwaro, leading to elevated kōwaro δ¹³C values, no effects on δ¹⁵N values, and higher relative abundance of large kōwaro in shaded streams compared to unshaded streams. Thus, the riparian canopy cover was able to offset some of the effects of drying.
6. Overall, we found no interactions between drying intensity and canopy cover affecting kōwaro. However, their opposing effect highlights the important role local conditions such as riparian canopies play on aquatic communities and their potential role as a restoration tool to mitigate the effects of large‐scale shifts such as drought.