Longitudinal and seasonal pattern of insect emergence in alpine streams

Prevailing water sources and/or regional climate are known to have an important influence on hydromorphology and chemistry of high alpine streams, affecting biology and phenology of aquatic insects considerably. Seven reaches in two different stream types (glacial and non-glacial) in the European Central Alps were investigated along a longitudinal gradient above the tree line to elucidate community structure and emergence patterns of aquatic insects. Aquatic insect emergence was dominated by chironomid taxa in both streams (95.0% in the glacial vs. 90.5% in the spring-fed stream). Emergence rate was much higher in the non-glacial stream, with Chironomidae 638.9 ind. m⁻² d⁻¹ and EPT (Ephemeroptera, Plecoptera, Trichoptera) 20.3 ind. m⁻² d⁻¹ (annual mean), compared to the glacial stream (Chironomidae 132 ind. m⁻² d⁻¹ and EPT 7.0 ind. m⁻² d⁻¹). Whereas, in the glacial stream a richer and more diverse species composition was found at lower elevations, emergence rate and emerging taxa numbers were higher at higher altitude in the non-glacial stream. Seasonal comparisons also showed a significant difference between the two streams. In the glacial stream maximum emergence was in April/May, whereas, in the non-glacial stream in July. A comparison with similar studies carried out in alpine streams showed that abundance and biomass of emerging insects were relatively low in the glacial stream. The continuous emergence throughout the summer is another example of insect life-cycle adaptation to the harsh environmental conditions in glacial streams: most likely, emergence during the warmer summer months, where the probability of experiencing favourable climate conditions on land is higher than for the rest of the year, was an evolutionary advantage for many glacial stream taxa.     

Runoff and the longitudinal distribution of macroinvertebrates in a glacier‐fed stream: implications for the effects of global warming

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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.     

Controls on periphyton biomass in heterotrophic streams

1. Headwater streams of the Hubbard Brook Experimental Forest (HBEF) are typically characterised by a periphyton assemblage of low biomass and diversity. However, periphyton blooms have been observed following catchment deforestation experiments and occasionally during the annual spring thaw before canopy leaf‐out. 2. There is pronounced seasonal variation in both nutrient and light availability in HBEF streams. Stream water nitrogen (N) concentrations and light levels are higher before canopy leaf‐out and after leaf senescence and are lower during the growing season. Periphyton accrual rates also change seasonally; they are highest in spring prior to leaf‐out and significantly lower during summer and in autumn. 3. Periphyton biomass rarely responded positively to in‐situ experimental enrichment with nitrogen or phosphorus. In the summer, nutrient enrichment overall had no effect on periphyton biomass, while outside the growing season N enrichment had inhibitory effects on periphyton. 4. Despite these experimental results, surveys of ambient chlorophyll a concentrations in streams across the HBEF demonstrated no relationship between streamwater dissolved inorganic N or P concentrations and benthic chlorophyll a. 5. Our results suggest that HBEF periphyton communities are not closely regulated by nutrient availability, even during periods of high light availability. The inhibitory effects of nutrient enrichment outside the growing season are interesting, but further research is necessary to elucidate the mechanisms driving these responses.     

Does rapid glacial recession affect feeding habits of alpine stream insects?

1. Glacial retreat, accompanied by shifts in riparian vegetation and glacier meltwater inputs, alters the energy supply and trophic structure of alpine stream food webs. Our goal in this study was to enhance understanding of dietary niches of macroinvertebrates inhabiting different alpine streams with contrasting glacial and non‐glacial (groundwater, precipitation, snowmelt) water inputs in conjunction with seasonal and habitat‐specific variation in basal resource availability.
2. We measured a range of stream physico‐chemical attributes as well as carbon and nitrogen isotopes (δ¹³C, δ¹⁵N) of macroinvertebrates and primary food sources at seven sites across seasons within a Swiss glaciated catchment (Val Roseg) undergoing rapid glacial retreat (1–2 km between 1997 and 2014). Sampling sites corresponded to streams used in a previous (1997/1998) study within the same alpine catchment.
3. Physico‐chemical attributes showed wide variation in environmental conditions across streams and seasons. Significant correlation among physico‐chemical proxies of glacier meltwater (phosphate‐P, total inorganic carbon, conductivity, turbidity) and macroinvertebrate δ¹³C, δ¹⁵N, and size‐corrected standard ellipse area (a proxy for feeding niche width) values showed that the extent of glacial water input shapes the energy base among alpine streams. Feeding niche differences among common alpine stream insect taxa (Chironomidae, Baetidae, Heptageniidae) were not significant, indicating that these organisms probably are plastic in feeding behaviour, opportunistically relying on food resources available in a particular stream and season.
4. Seasonal trends in macroinvertebrate δ¹³C largely followed patterns in periphyton δ¹³C values, indicating that autochthonous resources were the main consumer energy source within the stream network, as shown previously. The overall range in macroinvertebrate δ¹³C (?33.5 to ?18.4‰) and δ¹⁵N (?6.9 to 6.7‰) values also corresponded to values measured in the previous study, suggesting that macroinvertebrates altered diets in line with changes in environmental conditions and food resources during a period of rapid glacial retreat. Our results suggest that environmental changes brought on by rapid glacial retreat have not yet caused a profound change in the trophic structure within these fluvial networks.

     

Physical and biological changes to a lengthening stream gradient following a decade of rapid glacial recession

In mountains, environmental gradients are steep in both terrestrial and aquatic systems, and climate change is causing upward shifts of physical and biological features of these gradients. Glacial streams are an interesting system to evaluate such shifts both because streams have a linear nature (for simplicity of analysis), and because the stream habitat will at least temporarily lengthen as it follows receding glaciers upward. The Tschierva Glacier, Swiss Alps, receded 482 m upstream from 1997 to 2008. We tested the null hypothesis that the physical and biological stream gradient below this glacier maintained the same structure between these time periods, but simply shifted upward following the glacial source. We compared longitudinal patterns of water temperature and zoobenthic community structure in 1997 and 2007–2008 during three seasons (spring, summer, fall) along the uppermost ca. 5 stream km. Upward shifts were evident, including colonization of the newly exposed stream reaches by cold‐adapted taxa, and the appearance in 2007/2008 of four lower‐altitude species that were previously absent. Overall, however, results rejected the null hypothesis, instead revealing significant changes in gradient structures. These included a more steeply increasing temperature profile downstream of the glacier and increased amplitude of seasonal community turnover in 2007/2008 vs. 1997. Long‐term (1955–2007) flow records revealed increasing short‐term and seasonal hydrologic variability, which might have influenced the increased intra‐annual community variability. The steepening of the temperature gradient was likely caused by a warming lake‐outlet tributary upon which glacial influence was diminished between 1997 and 2007/2008. These results suggest that upward‐shifting gradients in glacial streams can involve complex interactions with other landscape elements and that local‐scale climate response can progress even more rapidly than the rate of glacial recession.     

Effect of periphyton biomass on hydraulic characteristics and nutrient cycling in streams

The effect of periphyton biomass on hydraulic characteristics and nutrient cycling was studied in laboratory streams with and without snail herbivores. Hydraulic characteristics, such as average water velocity, dispersion coefficients, and relative volume of transient storage zones (zones of stationary water), were quantified by performing short-term injections of a conservative tracer and fitting an advection-dispersion model to the conservative tracer concentration profile downstream from the injection site. Nutrient cycling was quantified by measuring two indices: (1) uptake rate of phosphorus from stream water normalized to gross primary production (GPP), a surrogate measure of total P demand, and (2) turnover rate of phosphorus in the periphyton matrix. These measures indicate the importance of internal cycling (within the periphyton matrix) in meeting the P demands of periphyton. Dense growths of filamentous diatoms and blue-green algae accumulated in the streams with no snails (high-biomass streams), whereas the periphyton communities in streams with snails consisted almost entirely of a thin layer of basal cells of Stigeoclonium sp. (low-biomass streams). Dispersion coefficients were significantly greater and transient storage zones were significantly larger in the high-biomass streams compared to the low-biomass streams. Rates of GPP-normalized P uptake from water and rates of P turnover in periphyton were significantly lower in high biomass than in low biomass periphyton communities, suggesting that a greater fraction of the P demand was met by recycling in the high biomass communities. Increases in streamwater P concentration significantly increased GPP-normalized P uptake in high biomass communities, suggesting diffusion limitation of nutrient transfer from stream water to algal cells in these communities. Our results demonstrate that accumulations of periphyton biomass can alter the hydraulic characteristics of streams, particularly by increasing transient storage zones, and can increase internal nutrient cycling. They suggest a close coupling of hydraulic characteristics and nutrient cycling processes in stream ecosystems.     

Patterns of macroinvertebrate traits along three glacial stream continuums

1. Glacier‐fed streams are characterised by low spatial but high temporal heterogeneity, manifested in seasonal and diurnal discharge and suspended sediment peaks induced by glacial runoff. These streams shelter macroinvertebrate communities adapted to such harsh environmental conditions. Studies relating macroinvertebrate traits to environmental conditions in glacial streams could provide important insights into the structure and function of glacial stream communities. 2. From data collected in three glacial streams from the central Swiss and southern French Alps, we analysed the relationships among six biological traits to define five groups of macroinvertebrate taxa with similar suites of traits. 3. The longitudinal distribution of the five groups and of individual traits was analysed, as well as their variation according to a glaciality index combining water temperature, conductivity, suspended solids and substrate stability. 4. The trait diversity along the three streams showed a strong upstream‐downstream gradient. The upper reaches were dominated by a single group of taxa characterised by small, crawling, deposit feeders. The other trait‐based groups appeared progressively downstream. 5. Changes in the relative frequency of trait‐based groups along the glaciality gradient highlighted the dominance of all‐rounder resistant/resilient traits in the three streams and confirmed that environmental conditions in the glacial streams are too harsh or uniform to allow macroinvertebrate communities to develop alternative suites of traits. The findings are discussed in relation to the question of trait coding in the available literature.     

Autochthonous primary production in southern Amazon headwater streams: Novel indicators of altered environmental integrity

The riparian forest reduces the amount of light entering streams, which limits autochthonous primary production. The aim of this study was to evaluate temporal variation of autochthonous primary production in pristine and altered streams, with the goal of identifying indicators of change in environmental integrity in the southern Brazilian Amazon. We evaluated free algal biomass in the water column, the presence of periphyton, and the richness and cover of aquatic herbaceous plants in 20 streams (10 pristine and 10 altered, i.e., with riparian deforestation) during the dry period, at the beginning of the rainy period, and at the end of the rainy period. In altered streams, we recorded the presence of macroscopic periphyton and the amount of algal biomass varied between the dry and flood seasons. Variations in hydrological periods did not contribute to changes in algal biomass in pristine streams; we did not observe the presence of macroscopic periphyton these streams. In altered streams, 23 aquatic herbaceous species were identified, versus only four in the pristine streams. Results showed that riparian deforestation contributes to increased autochthonous primary production, which is also influenced by different hydrological periods, with algae and aquatic herbaceous plants responding differently to dry and rainy periods. The responses of these primary producers confirm their role as important bioindicators of change in the environmental integrity of southern Amazonian streams.     

Effects of land cover on sediment regime and fish assemblage structure in four southern Appalachian streams

SUMMARY 1. We examined the relationship between catchment land cover, sediment regime and fish assemblage structure in four small streams in the upper Little Tennessee River basin of North Carolina. Study streams drained similar sized catchments (17–31 km²) with different fractions of non-forested land cover. Non-forested land cover was <3% in two 'reference' streams, whereas it was 13 and 22% in two 'disturbed' streams. Land cover data were compared with sediment transport data (suspended and bedload), benthic habitat data (embeddedness, substratum composition and coverage of fines) and fishes collected in autumn 1997.
2. Suspended sediment concentration was significantly higher in disturbed streams during both baseflow and stormflow. During baseflow disturbed streams nearly always exceeded 10 nephelometric turbidity units (NTU), whereas reference streams never exceeded this threshold. The difference in suspended sediment concentration between reference and disturbed streams was more consistent at baseflow than at stormflow. Therefore, baseflow turbidity may be a useful indicator of potential stream degradation.
3. Disturbed sites had five- to nine-fold more bedload transport than reference sites. Both embeddedness and streambed instability increased with increasing non-forested land cover.
4. Relative abundance of fishes requiring clean cobble/gravel substratum for spawning was lower in disturbed streams, whereas relative abundance of mound-building cyprinids, their nest associates and fishes that excavate nests in soft sediments (centrarchids) was higher. Relative abundance of fishes spawning in benthic crevices and gravel (BC + G) declined as the proportion of non-forested land cover increased. This study supports growing evidence that human-induced sedimentation alters stream fish assemblages.     

Benthic Algae in High Altitude Streams of the Alps – a Neglected Component of the Aquatic Biota

This is a review on benthic algae from streams situated above the tree line in the Alps. It integrates published and unpublished data from alpine streams in Austria and in the Trento Province (Northern Italy). The main focus is on the structural and taxonomic composition of benthic algae including macro- and micro-algae and their contribution to the epilithic biofilm and the stream food-web. The environmental conditions relevant to algae within the two major stream types, the glacial (glacier-fed) and non-glacial krenal (spring-fed) stream are discussed. The paper considers both the maximum possible structural complexity of transverse algae zonation in cascading alpine/subalpine stream segments, and the effects of glacial water on species richness in the Central Alps in Austria. Autecological data are given for 46 common diatoms from 42 sites in the Central Eastern and Southern Central Alps and for 30 algae in addition to diatoms for 22 streams in the Central Eastern Alps. Since data on responses of benthic algae to the harsh conditions in high altitude Alpine streams are very scarce, results from literature and the author’s experiences from these and other mountain stream types are synthesized to formulate major objectives for future research in benthic high altitude Alpine stream ecology.     

Longitudinal and seasonal distribution patterns of the benthic fauna of an alpine glacial stream (Val Roseg, Swiss Alps)

1. Seasonal changes in longitudinal patterns of environmental conditions and macroinvertebrate community distributions were examined in an alpine glacial stream (Roseg River, Switzerland). 2. Physico‐chemical parameters reflected seasonal changes in glacial influence via shifts in water sources and flowpaths (glacial meltwater versus ground water), and were best described by turbidity, particulate phosphorus and specific conductance. High nitrogen concentrations indicated snowmelt was the main water source in June. 3. Macroinvertebrate densities and taxon richness were highest during spring (4526 m^–2 and 16 taxa, all sites combined) and late autumn/early winter (8676–13 398 m^–2 with 16–18 taxa), indicating these periods may be more favourable for these animals than summer when glacial melting is maximal. Diamesa spp. (Chironomidae) dominated the fauna at the upper three sites (>95% of zoobenthos) and were abundant at all locations. Other common taxa at lower sites (1.2–10.6 km downstream of the glacier terminus) included other chironomids (Orthocladiinae, Tanytarsini), the mayflies Baetis alpinus and Rhithrogena spp., the stoneflies Leuctra spp. and Protonemura spp., blackflies (Simulium spp., Prosimulium spp.), and Oligochaeta. 4. Co‐inertia analysis revealed a strong relationship between environmental conditions and benthic macroinvertebrate assemblages. Furthermore, it elucidated temporal variability in longitudinal response patterns, as well as a similarity in temporal patterns among individual sites. 5. Our results suggest that zoobenthic gradients are not solely related to temperature and channel stability. Seasonal shifts in sources and pathways of water (i.e. extent of glacial influence), and periods of favourable environmental conditions (in spring and late autumn/early winter) also strongly influenced zoobenthic distributions.     

Can increased glacial melting resulting from global change provide attached algae with transient protection against high irradiance?

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凉水和帽儿山地区低级溪流生境和水质状况

为研究不同植被景观土地利用下低级溪流的生境状况和水质差异,对凉水国家级自然保护区红松原始林内和帽儿山国家森林公园境内天然次生林植被和农田背景下各3条溪流生境和水质状况进行调查。对溪流生物特性（悬浮藻、附着藻）和理化性质（温度、混浊度、溶解氧（DO）、pH、NH4^＋-N、NO3^-N、PO4^3-P、总氮（TN）和总磷（11P））以及细小颗粒有机物质（FPOM）和粗大有机物质（CPOM））进行测定。研究结果表明,凉水地区原始林溪流的生境状况好于帽儿山地区的次生林,帽儿山地区农田溪流生境最差。原始林溪流具有稳定的溪底生物生活基质,稳水区和急流区均匀分布,稳水区尺度变化大,受淀积物沉降干扰小,河道较弯曲,河岸稳定,河岸植被覆盖度高;次生林溪流以急流区为主,稳定基质相对较差,并受到一定程度的淀积物沉降干扰,河岸尚稳定,有一定程度的人为干扰;农田溪流基质不稳定,受到强烈淀积物沉降影响,渠道化严重,河岸带植被严重破坏。3种景观背景下溪流总磷（11P）、溶解氧（DO）、混浊度、温度、氮磷比值（N／P）（P〈0．05）存在显著差异。原始林溪流NH4^＋、DO、TP、TN、悬浮藻浓度和pH较高、附着藻数量较多,温度较低、FPOM和CPOM的数量较少;次生林溪流的NO3^--N、N／P和TDIN较高。 浊度较低;农田溪流浊度、温度、PO4^3--P较高,DO和pH较低,附着藻数量较少。景观尺度上的土地利用对溪流生境具有深刻的影响,同时决定溪流的水质状况。     

Temporal Patterns and Environmental Correlates of Macroinvertebrate Communities in Temporary Streams

Temporary streams are characterised by short periods of seasonal or annual stream flow after which streams contract into waterholes or pools of varying hydrological connectivity and permanence. Although these streams are widespread globally, temporal variability of their ecology is understudied, and understanding the processes that structure community composition in these systems is vital for predicting and managing the consequences of anthropogenic impacts. We used multivariate and univariate approaches to investigate temporal variability in macroinvertebrate compositional data from 13 years of sampling across multiple sites from autumn and spring, in South Australia, the driest state in the driest inhabited continent in the world. We examined the potential of land-use, geographic and environmental variables to predict the temporal variability in macroinvertebrate assemblages, and also identified indicator taxa, that is, those highly correlated with the most significantly associated physical variables. Temporal trajectories of macroinvertebrate communities varied within site in both seasons and across years. A combination of land-use, geographic and environmental variables accounted for 24% of the variation in community structure in autumn and 27% in spring. In autumn, community composition among sites were more closely clustered together relative to spring suggesting that communities were more similar in autumn than in spring. In both seasons, community structure was most strongly correlated with conductivity and latitude, and community structure was more associated with cover by agriculture than urban land-use. Maintaining temporary streams will require improved catchment management aimed at sustaining seasonal flows and critical refuge habitats, while also limiting the damaging effects from increased agriculture and urban developments.     

Effects of forestry on the thermal habitat of Dolly Varden (<Emphasis Type="Italic">Salvelinus malma</Emphasis>)

Forestry activities in riparian areas are known to affect stream communities considerably. Not only do riparian deforestation resulting from agriculture or urbanization developments affect stream communities but extensive commercial plantation and forestry practices can alter stream environments adjacent to remaining, intact or secondary forests. Because forestry often includes the construction of logging roads through the riparian zone, this can directly degrade stream environments. Twelve streams in the Shiretoko Peninsula, Hokkaido were investigated so as to determine the effects of forestry practices on stream temperature, periphyton biomass, grazer (benthic invertebrates) biomass and Dolly Varden (Salvelinus malma Walbaum) biomass. The greater the proportion of planted area in the catchment, the higher the stream temperature. Stream temperature directly affects periphyton biomass and Dolly Varden biomass negatively. Neither stream temperature nor periphyton biomass predicted grazer biomass, whereas a positive correlation was found between grazer biomass and Dolly Varden biomass that forage on invertebrates. The overall results indicated that Dolly Varden in the Shiretoko streams were negatively affected by forestry practices and the resultant stream temperature increases. Without effective future riparian forest management, the complex effects of both riparian disturbance and ongoing global warming could further reduce Dolly Varden populations in the region.     

Quantifying the direct and indirect effects of flow‐related disturbance on stream fish assemblages

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Autochthonous production in shallow littoral zones of five floodplain rivers: effects of flow,turbidity and nutrients

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The response of stream periphyton to Pacific salmon: using a model to understand the role of environmental context

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Seasonal flow variation allows 'time-sharing' by disparate aquatic insect communities in montane desert streams

1. Flow variation can drive major abiotic changes in stream environments between seasons. Theoretically, disparate biotic communities could be maintained during different seasons at a single site if suitable refuges and colonist sources were available. Using isolated montane desert streams in south‐east Arizona as a model system, we hypothesised that two disparate aquatic insect faunas (montane temperate and neotropical) could be maintained at the same sites through strong seasonal variation in abiotic conditions. 2. We collected aquatic insects representing 59 families from seven streams during high‐flow (March–April) and low‐flow (June) sampling periods across two years. We assessed changes in aquatic insect community and functional feeding group composition by habitat (riffle, pool) and season (high flow, low flow). 3. Within sites, wetted stream area decreased by an average of 97% between high‐flow (predominately riffles) and low‐flow (predominately pools) seasons. Community composition likewise showed strong seasonal patterns; the montane temperate fauna was strongly associated with the high‐flow season while neotropical hemipterans and coleopterans were associated with the low‐flow season. Increased water temperature was significantly associated with this shift from temperate to neotropical assemblages. 4. Functional feeding group composition shifted dramatically by season. The proportion of predators increased from 24.5% (high flow) to 75.2% (low flow) while collector–filterers and shredders declined from 38.4% (high flow) to 1.7% (low flow). 5. We suggest that habitat ‘time‐sharing’ by disparate communities is facilitated via strong seasonal variation in temperature and flow and the presence of high elevation refuges or diapause stages for temperate montane taxa to survive the dry season.