Longitudinal distribution of macroinvertebrate assemblages in a glacially influenced stream system in the Italian Alps

1. The longitudinal distribution of macroinvertebrates was investigated in June, August and September 1996 and 1997 in the Conca glacial stream and its tributary (Italian Alps; 46°N, 10°E). The principal aim was to test the 22 model that predicts the succession of faunal groups downstream of the glacial snout in relation to water temperature and channel stability. The effect of a non‐glacial tributary on the taxonomic richness and density patterns occurring in the glacial stream was also considered. 2. Channel stability showed an atypical longitudinal trend in the Conca glacial stream, being high in the upper part with Pfankuch Index values between 30 and 33. Water temperature exceeded 6 °C at all stations, with average values below 2 °C occurring only within 700 m from the glacial snout. 3. Taxonomic richness and diversity increased downstream. Taxonomic richness in the glacial stream (at about 1.5 km from the glacier) was comparable with the tributary and the reach after the confluence. Abundance also increased downstream in the glacial stream, but not as greatly as the number of taxa. 4. At higher taxonomic levels, the community structure in the tributary stations appeared to be similar to the two stations in the glacial stream just upstream of the confluence. The effect of the tributary was evident mainly at the genus or species level of the Chironomidae community. Some taxa found in the non‐glacial stream (e.g. Cricotopus fuscus, Eukiefferiella coerulescens, Metriocnemus sp., Paratrichocladius rufiventris, P. skirwitensis, Rheocricotopus effusus and Smittia sp.) were found also in the Conca stream but only after the confluence. 5. The upper glacial reach (within 700 m from the glacier snout) was dominated by the chironomid Diamesa spp. Less than 400 m from the glacier other Diamesinae (Pseudokiefferiella parva) and a few Orthocladiinae, especially Orthocladius (Euorthocladius) rivicola gr., colonized the stream. Some Diamesinae maintained relatively dense populations at mean water temperature around 5 °C, while some Orthocladiinae colonized reaches with mean water temperature <3 °C. 6. Contrary to the 22 model, Dipteran families such as Empididae and Limoniidae were more abundant in the upper stations than Simuliidae; non‐insects such as Nematoda and Oligochaeta were also numerous at some sites. Leuctridae, Taeniopterygidae and Nemouridae were the first Plecoptera to appear upstream, while Chloroperlidae were restricted to the lower reaches. Among Ephemeroptera, Heptageniidae were more abundant than Baetidae in the glacial sites. 7. In this glacial system channel stability and maximum temperature did not show the expected longitudinal trend and thus a typical kryal community was confined within 700 m from the glacier snout where summer mean water temperature was below 4 °C.     

Longitudinal zonation of macroinvertebrates in an Ecuadorian glacier‐fed stream: do tropical glacial systems fit the temperate model?

1. The ecology of glacier‐fed streams at temperate latitudes has been intensely studied in recent years, leading to the development of a well‐validated conceptual model on the longitudinal distribution of macroinvertebrate communities downstream of the glacier margin (Freshwater Biology, 2001a; 46 , 1833). However, to our knowledge, the ecology of tropical glacier‐fed streams has not yet been studied. 2. We sampled benthic macroinvertebrates and measured environmental variables at nine sites between 4730 and 4225 m altitude along a 4.3 km stretch of a glacier‐fed stream 40 km south of the equator in the Ecuadorian Andes. Our goal was to study the longitudinal distribution of the fauna in relation to environmental factors and to compare this with the conceptual model based on temperate–arctic glacier‐fed streams. 3. Total density of invertebrates differed considerably at the two highest altitude sites; 4600 m^?2 at a pro‐glacial lake outlet and only 4 m^?2 at a site originating directly from the glacier snout. Otherwise, there was a downstream decrease in density to about 825 m^?2 at the three lowest sites. Taxon richness increased with distance from the glacier, very similar to the pattern predicted. A total of 28 taxa were collected; two at the glacier snout, seven at the nearby pro‐glacial lake outlet, 13 at site 2 (<400 m from the glacier) and 20 at the lowest sites. 4. The numerical percentage of Chironomidae (Diptera) decreased downstream from 100 to 44%. The subfamily Podonominae was numerous at the highest sites but became much less important further downstream. The Orthocladiinae were important both in numbers and species at all sites, while Diamesinae were numerous only in the middle of the reach studied and were completely absent from the upper three sites. The limited importance of the Diamesinae, and its replacement by Podonominae, is different from the pattern typically observed in north‐temperate glacier‐fed streams. This could be because of the fact that the genus Diamesa is missing from the Neotropics. 5. Stream temperature and channel stability explained most of the variability in faunal composition and richness, supporting the model. Stability increased systematically downstream while temperature did not. Surprisingly, no classical kryal zone (T_max < 4 °C) was found, as even the site closest to the glacier snout (50 m) had a T_max of 15 °C and no site had T_max < 8 °C. We propose that this might be a general feature of equatorial glacial streams.     

When the going gets tough,the tough get going: The enigma of survival strategies in harsh glacial stream environments

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Why on the snow? Winter emergence strategies of snow‐active Chironomidae (Diptera) in Poland

A long‐term study of adult non‐biting midges (Chironomidae) active in winter on the snow in mountain areas and lowlands in Poland yielded 35 species. The lowland and mountain communities differed significantly in their specific composition. The mountain assemblage was found to be more diverse and abundant, with a substantial contribution from the subfamily Diamesinae, whereas Orthocladiinae predominated in the lowlands. Orthocladius wetterensis Brundin was the most characteristic and superdominant species in the winter‐active chironomid communities in both areas. Only a few specimens and species of snow‐active chironomids were recorded in late autumn and early winter. The abundance of chironomids peaked in late February in the mountain and lowland areas with an additional peak in the mountain areas in early April. However, this second peak of activity consisted mainly of Orthocladiinae, as Diamesinae emerged earliest in the season. Most snow‐active species emerged in mid‐ and late winter, but their seasonal patterns differed between the 2 regions as a result of the different species composition and the duration of snow cover in these regions. Spearman's rank correlation coefficient tests yielded positive results between each season and the number of chironomid individuals recorded in the mountain area. A positive correlation between air temperature, rising to +3.5 °C, and the number of specimens recorded on the snow in the mountain community was statistically significant. The winter emergence and mate‐searching strategies of chironomids are discussed in the light of global warming, and a brief compilation of most important published data on the phenomena studied is provided.     

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.     

Egg development and hatching success in alpine chironomids

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A note on chironomids (diptera) of temporary pools in the National Park of Circeo,Central Italy

Preliminary observations on chironomid assemblages in 9 temporary pools of the National Park of Circeo (Central Italy) are reported. A total of 15 genera or species groups (6 Orthocladiinae, 3 Tanypodinae, 1 Tanytarsini and 5 Chironomini) were recorded during March and April, 1986.Psectrotanypus varius, Polypedilum nubeculosum gr.,Chironomus thummi gr. andC. plumosus gr. were the most abundant and frequent taxa in the nine pools. Almost all chironomids collected are eurytopic and widely distributed in Europe, including Italian waters. Only the finding ofGymnometriocnemus is reported in this paper as a new record for Central Italy. Similarity among pools and among taxa (coefficient of Jaccard) shows a major occurrence of aquatic Orthocladiinae in smaller pools and of Chironomini in larger pools. This relationship between chironomid assemblages and pool sizes can be partly related to the duration of the wet phase which affects chironomid species according to their survival strategies.     

The response of Chironomidae (Diptera) to a long-term exclusion of terrestrial organic matter

We examined the effects of a seven-year detrital exclusion on chironomid assemblages in an Appalachian headwater stream. We hypothesized that litter exclusion would lead to a reduction in all chironomids at both the subfamily and generic levels because organic matter serves as both food and habitat in these headwater streams. Tanytarsini total abundance and biomass significantly declined after litter exclusion. Before litter exclusion, Tanytarsini average abundance was 4271 ± 1135 S.E. m⁻² and 625 ± 98 after litter exclusion. Biomass was 3.57 ± 0.96 mg AFDM m⁻² before litter exclusion and 1.03 ± 0.9 after exclusion. In contrast, Orthocladiinae abundance and biomass did not change because a psammanophilic chironomid, Lopescladius sp., and other Orthocladiinae genera did not decline significantly. Overall chironomid taxa richness and diversity did not change as a result of litter exclusion. However, Canonical Correspondence Analysis (CCA) of genus-level biomass did show a clear separation between the litter exclusion stream and a reference stream. Separation of taxa between the two streams was due to differences in fine (r ² = 0.39) and coarse (r ² = 0.36) organic matter standing stocks and the proportion of small inorganic substrates (r ² = 0.39) present within a sample. As organic matter declined in the litter exclusion stream, overall chironomid biomass declined and the chironomid community assemblage changed. Tanytarsini were replaced by Orthocladiinae in the litter exclusion stream because they were better able to live and feed on biofilm associated with inorganic substrates. Handling editor: K. Martens     

Weak altitudinal pattern of overall chironomid richness is a result of contrasting trends of subfamilies in high-altitude ponds

The decline of species richness with altitude is one of the most obvious patterns in ecology and results from the combination of ecological and evolutionary mechanisms. In harsh high-altitude environments, the effect of altitude usually overrules other environmental variables related to biodiversity. Studies using species richness along altitude gradients in high altitude are relatively numerous for lakes, but not for ponds. However, due to their special features, such as small size, high isolation and regional variability, ponds have been proved to be different systems compared to lakes. In high-altitude waterbodies, species of the family Chironomidae often dominate in benthic invertebrate communities and thus serve as an ideal model to study aquatic community changes along an altitude gradient. However, due to the time-consuming processing and expertise needed to identify the species, chironomids are often excluded from regular surveys. In the present study, we sampled 66 Tatra ponds over a 1100-m altitude gradient for benthic invertebrates, with special attention to chironomids. Out of the total 122 taxa collected, Chironomidae constituted the richest group with 58 taxa, being present in all the study ponds. The most diverse pond supported 13 chironomid taxa, and mean diversity was 6 taxa/pond. While total invertebrate richness decreased with altitude, chironomid richness showed only a weak negative response to altitude. The proportion of total chironomid diversity made up of Tanypodinae and Chironominae subfamilies decreased with altitude, while the opposite trend was recorded for the proportion of Diamesinae and Orthocladiinae.     

The influence of glacial runoff on stream macroinvertebrate communities in the Taillon catchment, French Pyrénées

1. Benthic communities were sampled from five sites within a glacial catchment in the Cirque du Gavaranie, French Pyrénées, over two consecutive years (i) to investigate whether longitudinal patterns in zoobenthic communities exist downstream of a glacial margin and (ii) to identify the principal environmental variables influencing such patterns. 2. There was a distinct zonation of communities with increasing distance from the glacial margin. Ordination of the zoobenthic distribution indicated sites were separated by the relative contributions of taxa rather than their presence or absence. A shift in community composition and diversity separated a kryal type community dominated by Diamesa spp., Prosimulium spp., Eriopterini and Empididae at ≥2200 m a.s.l., from a more rhithral community of Orthocladiinae, Ephemeroptera, Plecoptera and Trichoptera at 1900 m a.s.l. 3. Chironomidae showed a defined gradient in distribution from Pseudokiefferiella parva and Diamesa latitarsis groups close to the glacier, through D. zernyi and D. cinerella groups, Orthocladius, Parametriocnemus and Micropsectra further downstream with Rheocricotopus, Corynoneura and Nilotanypus furthest from the glacial margin. Diamesa cinerella/zernyi group was the most euryzonal taxon. 4. Gradients in channel and hydraulic stability, groundwater input and mean water temperature were identified as the principal environmental variables associated with the downstream distribution gradient of zoobenthos. Diamesa, Empididae, Eriopterini and Nematoda were most tolerant of channel and hydraulically unstable and cold water habitats. Simuliidae (Prosimulium), Crenobia alpina, Rhyacophila, Chaetopterygini, Drusus rectus, Capnioneura, Orthocladius and Parametriocnemus were associated with intermediate conditions. Corynoneura, Tanypodinae, Perlidae, Chloroperlidae, Agapetus fuscipes and Coleoptera were least tolerant of channel and hydraulic instability and low water temperature.     

Selective predation by four darter (Percidae) species on larval chironomids (Diptera) from a Mississippi stream

We determined seasonal foraging modes of four sympatric darter species, Etheostoma lynceum, E. stigmaeum, E. swaini and Percina nigrofasciata, from Beaverdam Creek, Mississippi (USA) at two scales of taxonomic resolution: (1) chironomid prey identified to family and (2) chironomid prey identified to genus/species. When chironomids were identified to family, high proportional similarity (PS) and low niche breadth (NB) values suggested the darters fed opportunistically on a relatively small number of available prey taxa. In contrast, when chironomids were identified to genus/species, concordant low PS and NB values suggested the darters fed like classic specialists, selecting a small number of prey taxa relative to prey availability in the resource base. The darters selected just one to four chironomid taxa from 52 available taxa across seasons. Our study shows that the scale of taxonomic resolution used to identify darter prey may influence the characterization of darter foraging modes.     

Ecology of alpine streams

1. This review examines ecological conditions and zoobenthic communities of kryal, krenal and rhithral streams of the alpine zone. Altitudinal and biogeographical faunal patterns are also analysed. 2. Kryal segments, fed by glacial meltwater, are characterized by low temperatures (T_max4°C) and large diel flow fluctuations in summer. The water may be clear or turbid from suspended rock flour. Fishes and higher plants are absent. The macroalga Hydrurus foetidus may be abundant in kryal and other alpine stream types of the Holarctic. The highly restricted cosmopolitan fauna of glacial brooks consists of diamesine chironomids, sometimes accompanied by simuliids. Sparse food resources include algae and allochthonous (aeolian) organic matter. 3. Rhithral segments in alpine catchments are characterized by soft water, a hydrograph dominated by an extended period of snowmelt runoff, and a broader temperature range than kryal or krenal biotopes. Bryophytes, macroalgae (chrysophytes, chlorophytes, cyanophytes, rhodophytes) and epiphytic and epilithic diatoms constitute the flora. A relatively diverse zoobenthos includes four orders of insects (Flecoptera, Ephemeroptera, Trichoptera, Diptera), turbellarians, acarines, oligochaetes and nematodes. 4. Krenal segments, fed by groundwater, are typically calcareous with summer-cool and winter-warm thermal conditions, high water clarity, and constant flow regimes. Bryophytes and macroafgae are accompanied by a rich diatom flora. The zoobenthos consists of a composite of kryal and rhithral elements with few crenobionts. Zoobenthos species richness values are intermediate between those of kryal and rhithral segments, whereas densities in perennial, well-oxygenated springbrooks far exceed those in other alpine stream types. 5. Downstream faunal changes are most predictable in kryal segments where chironomids of the genus Diamesa are the predominant, if not sole, members of the zoobenthos in the upper zone of glacial brooks, the metakryal. Where T_max exceeds about 2°C the transition to the hypokryal occurs and Diamesa is co-dominant with simuliids. These largely stenozonal headwater forms decline downstream where T_max exceeds about 4°C, concomitant with a marked increase in the euryzonal mountain fauna. 6. Species occurring in alpine rhithral biotopes tend to be euryzonal forms at their upper alrirudinal limits, whereas the lower elevation mountain stream fauna consists of species with narrower distribution limits. There is, however, a precipitous drop in mean altitudinal range from the alpine rhithral to the kryat because of the stenozonal nature of the glacial brook fauna. 7. The view that effects of temperature on generation time and mutation rate determine the speed at which selection proceeds is consistent with altitudinal species richness patterns exhibited by zoobenthos along the alrirudinal gradient and may provide an evolutionary explanation for the low faunal diversity in alpine headwaters. 8. With increasing altitude, mountain ‘islands’ become progressively insular as area decreases and isolation increases. For a cold-adapted stream fauna the insular nature of mountain tops is greatest in the tropics. Nonetheless, alpine stream faunas generally     

The macroinvertebrate communities of two contrasting Norwegian glacial rivers in relation to environmental variables

1. Macroinvertebrate communities in two Norwegian glacial rivers, one in the western fjords (Dalelva) and one in the eastern mountains (Leirungsåi), were investigated during three time periods in 1996 and 1997.
2. Channel stability variables (substratum heterogeneity/Pfankuch index/hydraulic stress) and water temperature accounted for 54% of the total inertia in the principal components analysis (PCA) ordination of environmental variables. The importance of these variables was confirmed by cluster analysis.
3. The two rivers were well separated in the ordinations, with Leirungsåi showing much greater heterogeneity. This is explained by differences in altitudinal range, terrestrial vegetation and the importance and nature of tributary inputs.
4. Channel stability and temperature were also important in determining faunal communities in the two glacial rivers, supporting the main determining variables in the conceptual model of glacial streams ( Milner & Petts, 1994 ). However, clear temporal differences were apparent in the data, the two rivers being more similar during the summer period of high discharge dominated by glacial meltwater. During spring and especially during autumn environmental conditions and the macroinvertebrate fauna differed both within and between rivers.
5. Diamesinae dominated in the upper reaches of both rivers, with Orthocladiinae becoming more common downstream. The dominance of Diamesinae persisted further down Dalelva because of the continued influence of glacial tributaries, whereas in Leirungsåi the influence of non-glacial tributaries led to a change towards a greater proportion of Orthocladiinae. Lakes modified macroinvertebrate communities in both river systems.     

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

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Does biogeographical history matter? Diversity and distribution of lotic midges (Diptera: Chironomidae) in the Australian Wet Tropics

Abstract We examined broad scale patterns of diversity and distribution of lotic Chironomidae (Diptera) within the Wet Tropics bioregion of northern Queensland, Australia. Field surveys across broad latitudinal and altitudinal gradients within the Wet Tropics revealed a fauna of 87 species‐level taxa in 49 genera comprising three main elements: a small genuinely tropical fraction, and larger cosmopolitan and Gondwanan components. The latter group originated when Australia, as part of the ancient Gondwana supercontinent, was situated over Antarctic latitudes with a cooler, wetter climate than today. In the Wet Tropics, cool Gondwanan taxa occurred predominantly in upland and shaded lowland sites, but no species appeared narrowly temperature restricted, and there was no faunal zonation with altitude. Most chironomid species occurred at all latitudes within the Wet Tropics, with no evidence for an enduring effect of the historical rainforest contractions on current‐day distribution patterns. These findings contrast with those for aquatic faunas elsewhere in the world and for the terrestrial Wet Tropics fauna. We relate this to the generally broad environmental tolerances of Australian chironomids, and comment on why the latitudinal diversity gradient does not apply to the Australian chironomid fauna.     

Longitudinal distribution of macroinvertebrates in two glacier-fed New Zealand rivers

1. Environmental variables, benthic algal biomass and macroinvertebrate fauna were examined from September 1999 to January 2000 (austral summer) along two glacier-fed rivers in South Island, New Zealand.
2. The rivers were characterized by high flow variability, high turbidity and physically disturbed beds. Water temperature ranged from <1 °C near the glacier margin to 10 °C further downstream.
3. Epilithic algal biomass was very low (<0.1 mg m^–2) in months characterized by heavy rainfall, but ranged from 1.1 to 14.4 mg m^–2 following an extended period with negligible precipitation.
4. Abundance and diversity of invertebrates in both rivers was low. Dominant taxa were Chironomidae (Orthocladiinae, Podonominae, Diamesinae), although mayfly species ( Deleatidium : Leptophlebiidae) also occurred at most sites. A species of Eukiefferiella (Orthocladiinae) was collected at all sites and was the most abundant invertebrate close to the glacier margins. No meiofauna were found in either river.
5. Faunal diversity increased at the lowermost stations where species of Plecoptera, Trichoptera, Coleoptera and non-chironomid Diptera also occurred.
6. The faunas of the two New Zealand rivers conformed to the conceptual model of Milner & Petts (1994) in that taxon richness increased downstream with water temperature. However, invertebrate abundance increased downstream in only one of the two rivers. Also in contrast to the model predictions, Leptophlebiidae and Orthocladiinae, rather than Diamesinae, dominated the fauna at the coldest sites.     

Temporal and spatial distribution of chironomid larvae and oligochaetes in two Dutch storage reservoirs

Long-term changes in distribution and taxonomic composition of chironomid larvae and oligochaetes in two water storage reservoirs in The Netherlands were studied. A succession among the chironomid species was observed. In the last 10–15 years chironomid densities varied. Compared with other lakes chironomid densities were high. Densities of chironomids were higher in the littoral zone than in the profundal zone. The opposite was found for oligochaetes. Densities and composition of the chironomid fauna in the two reservoirs were similar.Procladius, Tanytarsus andHarnischia dominated at all depths. However, pupal exuviae samples showed some differences in chironomid taxonomic composition between the two reservoirs. Orthocladiinae, rarely found in bottom samples, abounded in pupal exuviae samples.     

Macroinvertebrate stream communities along regional and physico-chemical gradients in Western Greenland

1. Macroinvertebrates were collected and physico‐chemical variables measured at 16 stream sites in Western Greenland during July 1999. Eight sites were located on Disko Island in an arctic oceanic climate and eight sites in the Kangerlussuaq area close to the icecap where the climate is arctic continental. The streams had different water sources (glacial, groundwater, snowmelt and lake water). 2. The streams showed pronounced differences in water temperature (2.2–17.3 °C), concentrations of suspended solids (0–2400 mg L^?1), and conductivity (10–109 μS cm^?1). Principal component analysis (PCA) analysis of the physico‐chemical variables separated the Disko Island sites into a distinct group, whereas the sites in the Kangerlussuaq area were more dispersed. 3. A total of 56 macroinvertebrate species were found, including 31 species of Chironomidae, the most abundant of which was Orthocladius thienemanni. Diamesa sp. was only the sixth most abundant chironomid taxon. Species composition varied between sites, and abundance varied from about 20 individuals m^?2 in a glacier fed stream to more than 16 000 m^?2 in a lake outlet. 4. The macroinvertebrate communities of the 16 streams were separated into five TWINSPAN groups reflecting water source, irrespective of region. Lake outlets and ground‐water‐fed streams had the highest species richness and abundance, temperature and bed stability, while glacier‐fed streams were characterized by low species richness, abundance, temperature, bed stability and high concentrations of suspended solids. Macroinvertebrate species richness was positively correlated with water temperature and negatively with bed stability. Conductivity was positively correlated with invertebrate abundance. 5. The results of this study suggest that the source of stream water can be used to predict invertebrate community composition in Greenlandic streams and thus the effects of changes in water balance and flow regime, and to identify sites of special conservation interest.     

Spatial variability in macroinvertebrate assemblages along and among neighbouring equatorial glacier‐fed streams

1. During the past two decades, understanding of the structure and function of glacier‐fed stream ecosystems at temperate latitudes has increased substantially. In contrast, information on their tropical counterparts is very limited. We studied three neighbouring glacier‐fed streams in the tropical Andes of Ecuador. Our main goals were (i) to determine overall longitudinal patterns in density, taxon richness and the composition of macroinvertebrate assemblages and driving factors in equatorial glacial streams and (ii) to examine variability among replicate streams in faunal metrics and assemblages, and stream‐specific effects of supposed environmental key factors. 2. We measured four geographical and 17 environmental factors and collected five Surber samples (500 cm²) of macroinvertebrates at each of nine sites, three sites along three streams. The streams were located 1–5 km apart. In each stream, the three sites were placed at comparable distances from the glacier and were grouped as ‘upper’ (50–200 m), ‘middle’ (1.5 km) and ‘lower’ sites (3.5–5.6 km). 3. In total, 2200 individuals (64% chironomids) were collected and 47 taxa (30 dipterans, 18 of these Chironomidae) identified. Density ranged from 176 to 372 ind. m^?2, and the number of taxa ranged from 2 to 6 at the upper sites and 868–3044 ind. m^?2 and 21–27 taxa at the lower sites. Density, number of taxa, rarefied richness and axis‐1 coordinates from a MDS ordination increased logarithmically with distance from the glacier. These faunal metrics were equally related to altitude and glacier per cent of catchment and correlated with maximum conductivity, mean temperature, mean daily maximum temperature and a channel stability index. As expected, the mean difference in distance decay in similarity was higher at the upper (47% km^?1) than at the lower reaches (20% km^?1) of the streams. 4. The number of taxa varied among sites within the upper and middle groups, but not among the lower sites. In contrast, but in accordance with our expectation, assemblage composition did not differ among upper sites but did so at middle and lower sites, following a supposed decrease in environmental harshness along the streams. Relationships between faunal metrics and the four environmental variables mean temperature, the stability index, chlorophyll a and coarse particulate organic matter also varied among the three streams. Generalised linear model analyses revealed that temperature interacted with stream on macroinvertebrate density, while chlorophyll a had a significant effect on the number of taxa in interaction with stream and stability. 5. The basic predictions of the Milner et al. (2001a) , model regarding longitudinal faunal patterns and temperature and stability as main driving factors were met by our three replicate equatorial glacial streams. Qualitative departures from the model were mainly because of zoogeographical differences. We demonstrated that variability in assemblages between comparable sites in closely situated streams was considerable, and the effect of key environmental factors varied among streams and interacted with other factors. Quantifying spatial variation in benthic assemblages may help us foresee possible consequences for biodiversity as a result of glacial retreat.     

A preliminary survey of the Chironomidae (Diptera) of Costa Rica,with emphasis on the lotic fauna

Extensive collections of Chironomidae were made in Costa Rica, Central America, during 1986 and 1987. Fifty-five genera and at least 148 species belonging to the subfamilies Podonominae, Tanypodinae, Orthocladiinae and Chironomidae were found. Chironominae and Orthocladiinae predominated. Only one species of Podonominae was collected. Tanypodinae was represented by many genera, but species richness was low.Cricotopus was the most widespread and diverse genus of Orthocladiinae. Among the Chironominae the generaPolypedilum, Pseudochironomus, Tanytarsus andRheotanytarsus showed high species richness. Several species were collected that could not be assigned to genus. A number of range extensions were recorded for taxa found in the Neotropical region for the first time and for Neotropical taxa recorded outside of South America for the first time. The Costa Rican chironomid fauna consists of cosmopolitan, holarctic and neotropical components. There is probably an endemic Central American chironomid fauna at the species level.