Headwater Stream Communities Research Articles

Phytoplankton in headwater streams: spatiotemporal patterns and underlying mechanisms

Phytoplankton are key members of river ecosystems wherein they influence and regulate the health of the local environment. Headwater streams are subject to minimal human activity and serve as the sources of rivers, generally exhibiting minimal pollution and strong hydrodynamic forces. To date, the characteristics of phytoplankton communities in headwater streams have remained poorly understood. This study aims to address this knowledge gap by comparing phytoplankton communities in headwater streams with those in plain rivers. The results demonstrated that within similar watershed sizes, lower levels of spatiotemporal variability were observed with respect to phytoplankton community as compared to plain rivers. Lower nutrient levels and strong hydrodynamics contribute to phytoplankton growth limitation in these streams, thereby reducing the levels of spatiotemporal variation. However, these conditions additionally contribute to greater phytoplankton diversity and consequent succession towards Cyanophyta. Overall, these results provide new insights into the dynamics of headwater stream ecosystems and support efforts for their ecological conservation.

Extreme climatic events alter the aquatic insect community in a pristine German stream

As a result of ongoing climate change, extreme climatic events (ECEs) are expected to become more frequent and severe. The high biodiversity of riverine ecosystems is susceptible to ECEs, especially to water temperature (extreme heat and extreme cold) and discharge-related (flood and drought) events. Long time series are needed to unravel the effects of ECEs on ecological communities. Here, we used 20 years (1986–2005) of unusually high-resolution data from a pristine first-order stream in Germany. Daily recordings of species-level identified aquatic insect (Ephemeroptera, Plecoptera, Trichoptera: EPT) emergence, water temperature and discharge data were used to examine the effects of four types of ECEs (extreme heat, extreme cold, flood, and drought events) on insect abundance, common taxonomic diversity metrics, and selected traits after five different time lags (2 weeks, 1, 3, 6, and 12 months). Extreme heat events increased from 1.8 ± 1.9 SE events per year before 2000 to 5.3 ± 1.9 SE events per year after 2000. Water temperature-related ECEs restructured the EPT community in abundance, species richness, and traits (community temperature index: CTI, and dispersal capacity metric: DCM). The strongest effects on the EPT community were found when it was exposed to multiple ECEs and 1 and 3 months after an ECE. The changing frequencies and durations of ECEs, especially the increasing frequency of extreme heat events and the negative cumulative effects of ECEs, paint a worrisome picture for the future of EPT communities in headwater streams. High-resolution, long-term data across sites is needed to further disentangle the effects of different ECE stressors.

Brown trout density effects on the macroinvertebrate and biofilm communities of headwater streams: a cage experiment

Salmo trutta is a key species of headwater streams that is able to influence the structure of these ecosystems through predation. Populations of this species are being threatened by climate change and hydrological alteration. In order to understand how their disappearance would affect the ecosystem structure of headwater streams, we used cage experiments to analyse different biofilm and macroinvertebrate structural parameters and the biodiversity of their communities under different trout densities. Due to a heavy rain event, our experiment was destroyed on the 10th day, but the role of trout as stream ecosystemic structure drivers was observed, as multiple parameters were modulated by their presence or absence. In the high-density treatments, a significant reduction in the biomass of macroinvertebrate families that fed on biofilms was observed due to direct predation on them, which caused the biofilms to be more autotrophic. A decrease in brown trout population densities could cause cascading trophic effects on the ecosystem structure of headwater streams

Cryptic declines of small, cold-water specialists highlight potential vulnerabilities of headwater streams as climate refugia

Increasing temperatures and climate-driven disturbances like wildfire are a growing threat to many species, including cold-water specialists. Montane areas and cold streams are often considered climate refugia that buffer communities against change. However, climate refugia are often species-specific, and despite growing awareness that life histories and habitat requirements shape responses to change, small or non-game species are often under-represented in monitoring and planning programs. A recent study in Montana, USA, revealed much larger warming-related declines in occupancy for small, non-game slimy sculpin (Cottus cognatus) between 1993 and 1995 and 2011–2013 than for two socially valued salmonid fishes that shape regional conservation efforts. To broaden insight into climate change vulnerabilities of headwater stream communities, we analyzed data for Rocky Mountain tailed frogs (Ascaphus montanus) that were collected during those same electrofishing surveys for fishes from 241 stream reaches. Tailed frogs occupy small, cold streams and have several life-history traits that make them sensitive to environmental change. We used a Bayesian framework to estimate occupancy, colonization, and extinction dynamics relative to forest canopy, estimated stream temperature, and wildfire effects. Tailed frog occupancy decreased by 19 % from 1993 to 1995 to 2011–2013. Changes in occupancy were linked with increased extinction and reduced colonization where there were fire-driven reductions in canopy cover, and reduced colonization of stream reaches that warmed on average 0.8 °C during the study. Our results highlight extensive extirpations for oft-overlooked species and emphasize the importance of including species with diverse habitat requirements and life histories in conservation planning.

Top-Down Effects of Salamanders on Macroinvertebrates in Fishless Headwater Streams

Salamanders are the top predators in many fishless headwater streams, and intraguild interactions among stream salamanders are well documented. However, little is known about the top-down effects of salamanders on stream food webs or how intraguild interactions mediate these effects. To investigate the effects of salamanders on macroinvertebrate communities of headwater streams, we conducted an experiment in stream mesocosms to test for effects of two stream salamander species, namely, Eurycea bislineata and Gyrinophilus porphyriticus, alone or in combination, on benthic and emerging macroinvertebrate density, biomass, and community composition. We also assessed intraguild interactions between these salamander species by comparing Eurycea bislineata survival and G. porphyriticus growth in single-species versus two-species treatments. Gyrinophilus porphyriticus reduced benthic macroinvertebrate densities when alone but not when co-occurring with E. bislineata. There were no effects of salamanders on benthic macroinvertebrate biomass or community composition and no effects on emerging macroinvertebrate density, biomass, or community composition. Eurycea bislineata survival decreased and G. porphyriticus weight increased in two-species treatments, suggesting that intraguild predation was occurring. Overall, although some of our findings are equivocal, these results suggest that salamanders can exert top-down control on macroinvertebrate communities in fishless headwater streams, decreasing benthic macroinvertebrate density. But this effect is dependent on the salamander species present, and can be removed by intraguild interactions between salamander species.

Multi-region assessment of chemical mixture exposures and predicted cumulative effects in USA wadeable urban/agriculture-gradient streams

Chemical-contaminant mixtures are widely reported in large stream reaches in urban/agriculture-developed watersheds, but mixture compositions and aggregate biological effects are less well understood in corresponding smaller headwaters, which comprise most of stream length, riparian connectivity, and spatial biodiversity. During 2014–2017, the U.S. Geological Survey (USGS) measured 389 unique organic analytes (pharmaceutical, pesticide, organic wastewater indicators) in 305 headwater streams within four contiguous United States (US) regions. Potential aquatic biological effects were evaluated for estimated maximum and median exposure conditions using multiple lines of evidence, including occurrence/concentrations of designed-bioactive pesticides and pharmaceuticals and cumulative risk screening based on vertebrate-centric ToxCast™ exposure-response data and on invertebrate and nonvascular plant aquatic life benchmarks. Mixed-contaminant exposures were ubiquitous and varied, with 78% (304) of analytes detected at least once and cumulative maximum concentrations up to more than 156,000 ng/L. Designed bioactives represented 83% of detected analytes. Contaminant summary metrics correlated strong-positive (rho (ρ): 0.569–0.719) to multiple watershed-development metrics, only weak-positive to point-source discharges (ρ: 0.225–353), and moderate- to strong-negative with multiple instream invertebrate metrics (ρ: −0.373 to −0.652). Risk screening indicated common exposures with high probability of vertebrate-centric molecular effects and of acute toxicity to invertebrates, respectively. The results confirm exposures to broad and diverse contaminant mixtures and provide convincing multiple lines of evidence that chemical contaminants contribute substantially to adverse multi-stressor effects in headwater-stream communities.

Aquatic insect communities in headwater streams of Ciliwung River watershed, West Java, Indonesia

Abstract. Wakhid, Rauf A, Krisanti M, Sumertajaya IM, Maryana N. 2021. Aquatic insect communities in headwater streams of Ciliwung River watershed, West Java, Indonesia. Biodiversitas 22: 30-41. Ciliwung is one of the major rivers flowing from Cisarua (Bogor) to Jakarta. This study was conducted to assess and compare aquatic insect biodiversity and community structure in various headwater streams with respect to different land-use types. Ten stream reaches of the first and second orders representing three different land-use types in the surrounding area (forest, tea plantation, village) were selected. At each stream reach, the aquatic insects were collected using a D-net (0.4 mm mesh). Physico-chemical properties of water were directly measured in the field. Samples were collected every two months from April until December 2018. A total of 26531 aquatic insect specimens belonging to 75 species/morphospecies, 46 families, and 10 orders were recorded. The order Trichoptera was the most abundant comprising 51.28% of the total collected insects, followed by Diptera (30.5%), Ephemeroptera (7.52%), and Plecoptera (6.25%). The highest proportion of functional feeding group was filtering-collectors (61.34%), followed by shredders (16.29%), gathering-collectors (15.32%), and predators (6.94%). Abundance of Diptera and Plecoptera was significantly higher in forest streams, while Ephemeroptera in plantation streams, and Trichoptera in village streams. The peak population of Diptera, Ephemeroptera, and Trichoptera occurred in June (dry period), while Plecoptera in December (wet period). The forest streams exhibited the highest species richness and diversity. Analysis of similarity (ANOSIM) showed a low dissimilarity among streams of the three land uses (Global R=0.217; P<0.001). The non-metric multidimensional scaling (NMDS) analysis showed that forest streams somewhat separated from plantation and village streams. The study provided important information on aquatic insect biodiversity, and indicated the need for forest area preservation in the headwater streams of Ciliwung River watershed.

Functional and trophic diversity of tropical headwater stream communities inferred from carbon, nitrogen and hydrogen stable isotope ratios

Tropical freshwaters support an immense diversity of fishes and invertebrates but are understudied in comparison to temperate systems. This is especially true of headwater streams, as only a small number of studies has assessed the trophic dynamics underpinning food web structure in these streams. We used stable isotopes of carbon, nitrogen and hydrogen to determine the resource use of dominant invertebrate guild and fishes in seven headwater streams in Eastern and Western Thailand, and assessed the functional and trophic diversity of each community using isotope food web metrics. Benthic invertebrates (95% credibility interval: 37–85%) and fishes (39–79%) obtained most of their resources from autochthonous sources in each stream but allochthonous and autochthonous specialists were evident in each community. We observed an increase in isotopic diversity of fishes associated with increasing stream size, but this was primarily driven by an increase in the range of isotope ratios of allochthonous and autochthonous food web endmembers rather than an increase in functional diversity. Maximum trophic position did increase with stream size. The snakehead, Channa gachua, was enriched in 2H relative to all other fishes, possibly reflecting facultative air breathing by this species. Fish communities in the headwater streams analysed filled a variety of trophic niches, predominantly fuelled by autochthonous primary production.

Seasonal variation in the strength of interference competition among headwater stream predators

Abstract Vertebrate communities in headwater streams are assumed to be regulated through competitive and predatory interactions. Although documented predation is rare, studies regularly report competitive dominance by fish that, as larger competitors reliant on aquatic habitat, exclude semi‐aquatic salamanders to marginal stream habitat. However, it is unclear whether fish interact with stream‐breeding salamanders through indirect effects such as competition for resources (e.g. food or cover) or fear (i.e. threat of predation) nor is it known whether these interactions are consistent through time. This study used a novel caging approach to determine if competitive outcomes between a headwater fish and salamanders were regulated primarily through resource depletion (exploitative competition) or behavioural avoidance (interference competition). We paired banded sculpin (Cottus carolinae) and larval red salamanders (Pseudotriton ruber) of similar body size in independent flow through mesocosms with intra‐ and inter‐specific pairs allowed to interact physically or non‐physically. The experiment was repeated in the autumn and in the spring when stream salamander larvae begin to transform into terrestrial juveniles. Banded sculpin negatively influenced growth of red salamanders regardless of whether they were allowed to physically interact, suggesting interference competition and behavioural avoidance. This asymmetrical effect was strongest in the spring when salamanders underwent metamorphosis at higher rates in the presence of fish. However, in the autumn, the effects were more balanced between the two species with salamanders impacting fish through exploitative competition. By studying the temporal relationships between two competitors and using a caging method novel to competition studies, we established that the outcomes of competition are dependent on season and may vary in type relative to the timing of life‐history events. For this community, these results suggest that outcomes of competition are highly dependent on season and could indicate a biotic mechanism maintaining headwater salamander distributions through source–sink dynamics. Our results also suggest that, in this species interaction, it may be unwarranted to assume that the outcomes of competition at one time represent the complex relationships regulating community interactions.

Alternatives for the biomonitoring of fish and phytoplankton in tropical streams

Biomonitoring programs need to balance accurate responses in assessments of changes in biological communities with sampling that is fast and low cost. In this study, we evaluated the concordance among fish and phytoplankton communities of streams. We tested the cross-taxa surrogacy, taxonomic, numerical resolution and ecological substitute group (habitat use and trophic guilds) resolution with Procrustes analyses aim of simplifying the biomonitoring process. We collect a total fish abundance of 8,461 individuals, represented by the ecological classes of habitat, including benthic, nektonic, nektobenthic, marginal and trophic guilds by detritivore, terrestrial invertivore, aquatic invertivore, piscivore, algivore and herbivore. We sampled a phytoplankton total density of 1,466.68 individuals/ml, represented by four Morphology-Based Functional Groups and nine Reynolds Functional Groups. Our results don’t support the use of substitute groups among fish and phytoplankton. For fish, habitat use and trophic guild are good surrogates for species-level data. Additionally, our results don’t support the use of functional groups as surrogates for phytoplankton. We suggest the use of higher taxonomic levels (genus and family) and record only the occurrence of species and/or genus for fish and phytoplankton. Our findings contribute to decreasing the costs and time of biomonitoring programs assessments and/or conservation plans on fish and phytoplankton communities of headwater streams.

Temporal variability in stream fish assemblage metrics and implications for long-term monitoring

High natural variability in the condition of fish communities in headwater streams complicates detection of long-term responses to changes in water quality. As a result, little is known about the impacts and recovery of fishes from acid deposition in streams of New York. Twenty-one fish metrics from annual electrofishing surveys at 13 streams sites in the Catskill and Adirondack mountains were assessed to quantify temporal variability and identify effective monitoring strategies for detecting change in headwater stream fish assemblages. Metrics included the density and biomass of Brook Trout populations and entire fish communities using length-, area-, and effort-based standardization techniques. Linear mixed models were used to estimate changes in coefficients of variation (CV) for different classes of metrics and standardization techniques, and a simulation-based power analysis was conducted to assess differences in statistical power for each metric with various sampling designs. Metric CV varied significantly as a result of standardization technique and whether metrics were calculated for the entire community or Brook Trout only. The sampling effort necessary to detect a 30% change with power of 0.80 was strongly correlated with CV (R2 = 0.77). The number of sampling events at the 13 sites needed to detect this change ranged from 1 to >60, suggesting metric selection can strongly affect statistical power and the resources necessary to detect change. Thus, metric selection is a complex decision that must consider study objectives and biological relevance, in addition to natural variability and statistical power. However, adequate statistical power was achieved at relatively small sample sizes using certain metrics, indicating that fish communities in headwater streams can be a valuable component of long-term assessments of acidification impacts and recovery.

Temporal and spatial dynamics in microbial community composition within a temperate stream network.

The water column of streams hosts a unique microbial community that is distinct from the microbial communities of the stream benthos and surrounding soil. This community is shaped by complex interacting forces, including microbial dispersal from surrounding environments and in-stream selection. However, how the processes structuring stream communities change over space and time remains poorly understood. In this study, we characterize spatial and temporal trends in microbial community composition throughout a stream network spanning first through fifth order streams. We found that the microbial communities of headwater streams are compositionally diverse, with low representation of freshwater microbial taxa and high representation of soil and sediment-associated taxa. In three out of five seasonal samplings, a successional pattern was identified in which phylotype richness and compositional heterogeneity decreased while the proportion of known freshwater taxa increased with increasing cumulative upstream dendritic distance. However, in two samplings, streams instead exhibited uniformly high microbial diversity across the watershed, and the fraction of freshwater taxa showed no relationship with dendritic distance. Overall, our data suggest that the successional processes that drive microbial diversity in streams are highly dynamic and can be disrupted at landscape scales, potentially in response to variation in temperature and precipitation.

Difference in benthic invertebrate communities of headwater streams can be detected using a short elevation gradient

Communities of benthic invertebrates from a series of intermittent headwater streams spanning over a short elevation gradient (i.e., ~ 180–500 m) were investigated in April–July 2010 and May–July 2011. The main purpose of this study was to better understand whether the potential future effect of environmental change on biological communities of the Precambrian Shield’s freshwaters can be detected using elevation as a substitute for time. Since obtaining long-term environmental data is a time-consuming process, substituting space for time could instead generate similar information in a shorter time. In this study, environmental differences associated with short elevation gradient were correlated with differences in benthic invertebrate communities. Therefore, elevation gradient provides a spatial proxy for anticipated future environmental change impacts over time. It was determined that water temperature accounts for the greatest variation in communities along the elevation. Many community metrics such as abundance, functional feeding groups, diversity, and evenness were significantly different based on the difference in elevation. Result indicates that even a short elevation gradient can potentially be used as a surrogate to look at the effect of environmental change.

Spatio-Temporal Variability in Benthic Macroinvertebrate Communities in Headwater Streams in South Korea

Comprehensive research on the structural and functional variability of benthic macroinvertebrate communities within headwater streams is limited, despite the fact that the majority of streams within a watershed are headwater streams that form the primary link between terrestrial and aquatic ecosystems. Therefore, we investigated the structure and function of benthic macroinvertebrate communities in four headwater streams at two different spatial scales (i.e., sampling sites (i.e., reaches) >samples (i.e., riffles)) over three seasons (i.e., spring, summer and autumn) of the year. Community indices, functional feeding guilds and habit trait guilds varied significantly depending on the seasons rather than on sites in two-way ANOVA based on spatial (i.e., sampling sites) and seasonal effects in each headwater stream. Non-metric multidimensional scaling analyses showed the differences between communities according to the considered spatial and temporal scales. At the individual stream scale, the differences between samples followed seasonal variation more than spatial differences. Site differences became more important when performing an ordination within a single season (i.e., spring, summer, and autumn). Continued research and monitoring employing both multidisciplinary and multidimensional approaches are required to maintain macroinvertebrate diversity within headwater streams.

Benthic Collector and Grazer Communities Are Threatened by Hemlock Woolly Adelgid-Induced Eastern Hemlock Loss

Eastern hemlock (Tsuga canadensis (L.) is a foundation species in eastern North America where it is under threat from the highly invasive, exotic hemlock woolly adelgid (Adelges tsugae). Eastern hemlock is especially important in riparian areas of Central and Southern Appalachia, so we compared the spatial and temporal composition of benthic collector-gatherers, collector-filterers, and grazers in headwater streams with hemlock-dominated riparian vegetation to those with deciduous tree-dominated riparian vegetation to evaluate the extent to which adelgid-induced hemlock loss could influence composition and abundance of these two functional feeding groups. We found differences in benthic invertebrate abundance and family-level diversity based on riparian vegetation and sampling approach, and, often, riparian vegetation significantly interacted with location or season. Collector-gatherers and grazers were more abundant in eastern hemlock streams in the summer, when hemlock litter is readily available and deciduous litter is relatively sparse. Riparian eastern hemlock appears to exert considerable influence on benthic invertebrate functional feeding group composition in headwater stream communities, as expected with a foundation species. With the loss of eastern hemlock due to adelgid-induced mortality, we should expect to see alterations in spatial and temporal patterns of benthic invertebrate abundance and diversity, with potential consequences to both benthic and terrestrial ecosystem function.

Edge-mediated effects of forest fragments on the trophic structure of stream fish

The aquatic communities of headwater streams are highly influenced by the surrounding matrix. The edge effects of forest fragments are known to influence terrestrial communities; however, their influence on aquatic communities remains poorly understood. The aim of this study was to investigate whether and how edge effects can influence stream environmental variables and fish trophic structure. We sampled stream reaches located within forest fragments (192–2480 ha) and stream reaches located on the forest edge during wet and dry periods to describe local habitat variables and fish trophic structure. We found high dissimilarity in the trophic structure (>60%) between stream positions and between seasons. Aquatic invertivores contributed to the majority of the observed dissimilarity. Omnivores and algivores were more abundant on edge reaches and were notably associated with high water temperature. Herbivores and terrestrial invertivores were more abundant in the interior reaches mostly during the wet season and were associated with high dissolved oxygen. The scarcity of riparian forest cover on the edge reaches not only caused physical changes but also influenced the fish trophic structure, therefore providing evidence of edge-mediated effects.

Effects of conservation status of the riparian vegetation on fish assemblage structure in neotropical headwater streams

The aims of this study were to investigate the relationships between the riparian vegetation and fish assemblages (structure, composition and diet) in forested (Riparian Cover ≥80%; Canopy cover ≥90%) and deforested (RC < 80%; CC < 90%) headwater streams and to test how those assemblage characteristics respond temporally in streams with different conservation conditions. We sampled in five Neotropical headwater streams during dry and wet season in areas of natural and degraded riparian vegetation on central Brazil. The fish assemblage structure was related to the conservation status of the stream, and no differences were observed in the assemblages between seasons. Allochthonous food items represented 47% of the fishes’ diet, followed by items of unknown origin and autochthonous items. Only Astyanax sp. and Astyanax rivularis showed evidence of temporal diet variation. The consumption of a wide spectrum of food items was an outstanding feature, indicating the fishes’ capacity to adapt to ecosystem changes. Differences between natural and deforested streams demonstrated the importance of the riparian zone as an indirect selective force on diets and composition of fish communities in headwater streams, suggesting that changes in the structure and composition of riparian vegetation may significantly affect the integrity of fish communities.

Salinity as a Limiting Factor for Biological Condition in Mining‐Influenced Central Appalachian Headwater Streams

AbstractRecent studies have found that Appalachian coal mining causes increased surface water salinity, and that benthic macroinvertebrate communities in salinized mining‐influenced streams differ from communities in streams draining unmined areas. Understanding the role of salinity in shaping these communities is challenging because such streams are often influenced by a variety of stressors in addition to salinity. We characterized associations of salinity with biotic condition while isolating salinity from other stressors through rigorous site selection. We used a multimetric index of biotic condition to characterize benthic macroinvertebrate communities in headwater streams in the Central Appalachian Ecoregion of Virginia across a gradient of sulfate‐dominated salinity. We found strong negative seasonal correlations between biotic condition and three salinity measures (specific conductance, total dissolved solids, and SO42− concentration). We found no evidence to suggest stressors other than salinity as significant influences on biotic condition in these streams. Our results confirm negative associations of salinity with benthic macroinvertebrate community condition, as observed in other studies. Thus, our findings demonstrate that elevated salinity is an important limiting factor for biological condition in Central Appalachian headwater streams.

Non-native rainbow trout change the structure of benthic communities in headwater streams of the Cape Floristic Region, South Africa

Introduced rainbow trout Oncorhynchus mykiss have invaded many headwater streams in the Cape Floristic Region (CFR) and depleted, or elimi- nated, native fish populations. However, the question of whether trout invasions also have consequences for lower trophic levels in these systems has not been addressed. We used a broad-scale comparative study in the upper Breede River catchment (CFR) to evaluate differences in benthic community structure between sites on headwater streams with and without trout, and thereby infer community-level impacts of trout. There were differences in invertebrate abundance and assem- blage composition, and algal biomass, between sites with and without trout. Specifically, the abundance of certain herbivorous invertebrate taxa was higher, and the biomass of benthic algae lower, at sites invaded by trout. This pattern implies that trout have induced a trophic cascade by releasing herbivorous invertebrates from predation, leading to an increase in grazing pressure and a consequent indirect decrease in the biomass of benthic algae; a pattern that contrasts with the majority of studies investigating community-level impacts of introduced trout elsewhere. These findings, together with comparisons of environmental condi- tions between invaded and uninvaded sites, indicate that trout invasions have changed the structure and function of benthic communities in these streams.

The invasibility of small headwater streams by an emerging invader, Clarias gariepinus

African sharptooth catfish Clarias gariepinus has been spread globally and is proven to be invasive with demonstrable impacts on a variety of biota in recipient ecosystems. There are concerns that this species will invade headwater stream communities that are already considered fragile and heavily impacted by non-native invasive fishes. There is however no information on the ability of C. gariepinus to penetrate or establish in these environments. This paper uses data collected over a seven year period from three episodic headwater streams in the Cape Floristic Region of South Africa to assess the ability of C. gariepinus to invade headwater habitats from downstream source populations. Data demonstrate that C. gariepinus invaded all three headwater tributaries and was able to penetrate previously non-invaded threatened fish refugia. Clarias gariepinus however failed to establish in headwaters and their presence was casual with individuals migrating from mainstem sources. Headwater incursions were dynamic and C. gariepinus reinvaded streams following their extirpation through drought. These results indicate headwater streams to be invasible by C. gariepinus and an assessment of potential impact on headwater communities requires urgent attention.