1. Hydrobiological changes were assessed along an altitudinal transect of eighteen to twenty-three tributaries from 6(X) to 3750 m in two adjacent river systems in east-central Nepal. The transect incorporated catchments under terraced agriculture at the lowest altitudes in the Likhu Khola, through streams in forest, alpine scrub and tundra at higher altitudes in Langtang. 2. Diatoms, bryophytes, macroinvertebrates and fish all showed pronounced altitudinal changes in assemblage composition as shown by TWINSPAN and DECORANA. A few taxa were restricted to streams at high altitude, but many more occurred only at lower altitudes where taxon richness increased substantially despite catchment disturbance by terraced agriculture. 3. Diatoms characteristic of lower altitude streams were mostly motile, epipelic or episammic Navicula and Nitzschia spp., which occur typically at greater electrolyte and nutrient concentrations. Those characteristic of higher and steeper sites included attached Fragilaria spp. and prostrate Achnanthes spp., tolerant of turbulent flow. 4. Cover by bryophytes varied within catchment type; high altitude springs supported dense mats, unlike streams fed by ice and glaciers. Taxa confined to low altitudes included those characteristic of humid subtropical conditions. 5. Invertebrate families occurring only at lower altitudes included a range of burrowers and pool dwellers. Numerically, filter feeding Hydropsychidae and Simuliidae dominated streams in terraced and forested catchments, whereas grazing baetid mayflies dominated higher altitude streams in scrub and tundra. 6. The combined density and biomass of at least six fish species in the Likhu Khola were 23-250 (per 100 m"^), and 86-1282 g wet mass (per 100 m"^), respectively. No fish were found in Langtang streams, probably because torrential headwaters prevented colonization. 7. Our data confirm that altitudinal transitions in stream biota are pronounced in the Himalaya of Nepal, but are likely to reflect a wide array of potential influences.• -in the distributions of freshwater oreanisms are undescribed. Polar, high altitude, tropical and subAlthough freshwater ecology is maturing into an tropical regions are among those known inadequately, experimental and process-orientated science, there and together they cover a large proportion of the are still large areas of the world where basic patterns earth's land surface. As a result, much ecological 309
SUMMARY 1. Surprisingly few data compare the apparent responses of diatoms and macroinvertebrates to metals in streams. We examined variation in metals, diatoms and macroinvertebrates between 51 streams in metal‐mining areas of Wales and Cornwall, U.K., using a survey design with multiple reference and polluted sites. 2. To quantify variations in metals between sites, we calculated cumulative criterion unit (CCU) scores, a recently defined measure of total stream metal concentration and toxicity, to account for additive effects of each metal relative to putative toxic thresholds. We compared assemblage responses among epilithic diatoms and macroinvertebrates to CCU scores or individual metal concentrations using correlation and detrended correspondence analysis (DCA). 3. Macroinvertebrate diversity, richness and total abundance declined and evenness increased with increasing copper concentrations. Trends with CCU scores were significant but less pronounced. Some individual macroinvertebrate taxa varied significantly in abundance with CCU scores, copper or zinc, but overall assemblage composition correlated only with manganese, pH and nitrate. 4. Among diatoms, pH and conductivity explained the major variations in assemblage composition, and neither diversity, richness nor evenness varied with metal concentration. Nevertheless, the single strongest predictor of diatom assemblages on ordination axis 2 was the CCU score. The abundances of some macroinvertebrate taxa, particularly grazers, also explained significant variations in diatom assemblages that were linked to both metals and acid–base status. 5. Diatom species apparently tolerant of high metal concentrations included Psammothidium helveticum, Eunotia subarcuatoides, Pinnularia subcapitata and Sellaphora seminulum. Of these, P. helveticum, E. subarcuatoides and P. subcapitata were abundant at lower pH than S.seminulum and might indicate metal enrichment over different pH ranges. Sensitive species included Fragilaria capucina var. rumpens, Achnanthes oblongella and Tabellaria flocculosa. 6. We conclude that macroinvertebrates at these sites reflected metal pollution most strongly through variations in diversity while effects on diatoms were best reflected by changes in assemblage composition. We suggest that, with further refinement, CCU scores might be useful in evaluating the possible effects of metal pollution on benthic organisms in European rivers.
1. Using a replicated survey design at the catchment scale, we compared the composition and diversity of diatom communities in three stream groups in the Middle Hills of Nepal: semi‐natural reference (Arun Valley), enriched by agricultural runoff (Likhu Khola) and grossly polluted by sewage (Kathmandu Valley). We sampled riffles at all sites; in the Kathmandu Valley and Likhu Khola we also sampled diatoms in pools and on vegetation. 2. Species richness and diversity H′ were significantly higher in agricultural streams than in either organically polluted streams or references. Community composition, as shown by principal components analysis, varied significantly between all three stream types due to differences in the abundances of species characteristic of organic enrichment (Kathmandu Valley), moderate enrichment (Likhu Khola) or undisturbed hillstreams (Arun Valley). 3. Few species varied in relative abundance between pools, riffles and vegetation so that variations in community composition and diversity were stronger between stream groups than between habitats. Samples from any one habitat produced only 75.7% (± 8.6 SD) of the species from three habitats, so that surveys aimed at recording biodiversity may need more comprehensive habitat coverage than surveys for biomonitoring. 4. We conclude that diatom communities can indicate different sources of pollution in Nepalese streams, and advocate further studies to develop this indicator potential over a wider area of the Himalaya. Comparisons between replicate groups of streams selected a priori helped to clarify effects which were sometimes obscured by other survey designs.
Summary 1. Diatoms are recognised as indicators in temperate streams, but only recently have assessments begun of their value in indicating stream quality in the tropics and sub‐tropics. Here, we extend previous studies by assessing stream diatom assemblages in relation to water quality and habitat character in the Kathmandu Valley, and in the Middle Hills of Nepal and northern India. We also assessed whether the U.K. Trophic Diatom Index (TDI) was sufficiently portable to reveal pollution in Himalayan rivers. In the more urbanised and highly agricultural Kathmandu Valley, we compared diatom response to water quality classes indicated by a local invertebrate index, the Nepalese Biotic Score (NEPBIOS). 2. Thirty and 53 streams in the Kathmandu Valley (2000) and Middle Hills (1994–96), respectively, were sampled in October and November during stable flows following the monsoon. Diatoms were collected in riffles, water samples taken for chemical analysis, and habitat character of the stream channel, bank and catchment assessed using river habitat surveys. In the Kathmandu Valley, macroinvertebrates were collected by kick‐sampling. 3. In total, 113 diatom taxa were found in the Kathmandu Valley streams and 106 in the Middle Hills. Of 168 taxa recorded, 62 occurred only in the Kathmandu Valley, 56 only in the Middle Hills and 50 were common to both areas. Most taxa found only in the Kathmandu Valley belonged to the genus Navicula while most taxa confined to the Middle Hills were Achnanthes, Fragilaria and Gomphonema. 4. In the Kathmandu Valley, richness and diversity increased significantly with K, Cl, SO4 and NO3, but declined significantly with Al, Fe, surfactants and phenols. Richness here also varied with habitat structure, being lowest in fast flowing, shaded streams with coarse substrata in forested catchments. In all streams combined, richness increased significantly with Si, Na and PO4, but declined significantly with increasing pH, Ca and Mg. 5. Diatom assemblage composition in the Kathmandu Valley strongly reflected water chemistry as revealed by cations (K, Na, Mg, Ca), anions (Cl, SO4), nutrients (NO3, PO4, Si), and also substratum composition, flow character and catchment land use. The commonest taxa in base‐poor forested catchments were Achnanthes siamlinearis, A. subhudsonis, A. undata and an unidentified Gomphonema species; Cocconeis placentula and Navicula minima in agricultural catchments; and Mayamaea atomus var. alcimonica, M. atomus var. permitis, and Nitzschia palea at polluted sites near settlements. Diatom assemblages in none‐agricultural catchments of the Kathmandu Valley and Middle Hills were similar, but they contrasted strongly between urban or agricultural catchments of the Kathmandu Valley and the less intensively farmed catchments of the Middle Hills. 6. In keeping with variations in assemblage composition, most streams in the Kathmandu Valley had higher TDI values (33–87, median = 64) and more pollution tolerant taxa (0–78%, median = 16) than streams in the Middle Hills (25–82, median 4...
1. Acid-base status has major effects on diatoms, but there is little information on their shortterm response to changing acidity. This is despite the use of diatoms as bioindicators in streams where acid episodes are important during rainstorms (hours to days) or snowmelt (days to weeks). In the Llyn Brianne experimental catchments (Wales, UK), we attempted to mimic the effects of short-term acidification by (i) reciprocally transplanting diatoms between two streams of contrasting acidity and (ii) using acid-diffusing substrates. 2. Diatom diversity decreased rapidly on substrata transplanted from the circumneutral into the acidic stream, and increased in the reciprocal transplantation. Changes in dominant taxa occurred within three days in the acidic stream because of the rapid growth of Eunotia exigua, and by nine days in the circumneutral stream because of the proliferation of Achnanthidium minutissimum. Transplants were near indistinguishable from ambient assemblages by day 12. 3. There were no effects of enclosures on assemblage composition, but diatoms responded more rapidly to altered chemistry in enclosures with coarse mesh (26 · 50 mm) than finer mesh (320 lm). 4. Chemical diffusing substrates comprised terracotta tiles attached to dosing reservoirs that created locally acid (using H 2 SO 4 ) or metal-rich conditions (using MnSO 4 ) in the circumneutral stream over 26 days. Diatom responses were compared with reference substrates dosed with deionised or circumneutral stream water, and we also assessed whether effects were moderated by macroinvertebrate grazers. 5. Surface pH was lower by 1-2 pH units on acid-dosed substrates than on reference tiles or in surrounding streamwater. Grazed assemblages on acid-dosed substrates differed significantly from ungrazed reference assemblages, acquiring significantly greater relative abundance of Eunotia spp. However, the magnitude of response was less than in the between-stream transplantations either because (i) metal exposure and base cation concentrations differed between the transplants and dosing substrates or (ii) diatom response to reduced pH on the diffusing substrates was restricted by the scarcity of acidobiontic diatoms in the circumneutral stream. Similar filter, founder or dominance effects might also affect diatom responses to real acid episodes.6. These data show that diatom assemblages can respond rapidly and directly to changes in acid-base status, but short-term acidification might affect diatoms more rapidly than subsequent recovery. Because the experimental methods used were imperfect representations of episodic effects, diatom response to real acid events requires further field evaluation.
High altitude ecosystems have important natural ecological functions but are under increasing impacts from human activities and climate change. A detailed analysis of the water chemistry of Lake Rara, a high mountain lake in western Nepal, was carried out in October 2015 and April 2016. A total of 31 water samples were collected.Major ions (Ca 2+ , Mg 2+ , Na + , K + , SO 4 2−
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