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.     

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.     

Influence of water abstraction on the macroinvertebrate community gradient within a glacial stream system: La Borgne d'Arolla, Valais, Switzerland

1. Water abstraction from glacial rivers is an important characteristic of hydroelectric power schemes in Alpine regions. Streams in the Valais region of Switzerland have been particularly affected. 2. Invertebrate distributions are described in La Borgne d'Arolla, a glacial stream with icemelt-, snowmelt- and groundwater-dominated tributaries. The icemelt-dominated streams have been affected by abstractions for more than 30 years. 3. The glacial streams contain only Chironomidae (Diamesa), and are devoid of fauna for between 200 and 500 m below the glacier snouts. 4. Immediately below the water intakes the streams are intermittent, flowing only during system purges and high floods, and are devoid of fauna for short distances (<1.5km). 5. Further downstream, abstraction of glacial meltwater increases the importance of snowmelt and groundwater, increasing water temperatures, improving water clarity and increasing the length of krenal/rhithral streams at the expense of kryal streams. 6. A community including Chironomidae, Simuliidae, Baetidae, Nemouridae, Limnephilidae and Chloroperlidae occurs as soon as a permanent flow is maintained by tributary runoff, and the channel becomes stable. 7. A wide range of taxa inhabit snowmelt- and groundwater-dominated tributary streams with stable channels, often at much higher altitudes than the main river. The tributaries provide sources for rapid colonization of the main channel following ice retreat or physical disturbance. 8. Purges and high floods are important disturbances within the main channel. Recovery may be rapid because of drift from tributaries, but sites influenced by frequent disturbances have reduced faunas in comparison to stable channel sites. 9. This study supports the model proposed by Milner & Petts (1994) and shows that deterministic responses of macroinvertebrate communities may be observed to changes of temperature, turbidity, flow regime and channel stability.     

A comparison of benthic macroinvertebrate assemblages among different types of alpine streams

1. Benthic macroinvertebrate assemblages were compared among a diverse array of first‐order alpine tundra streams of the Swiss Alps. 2. A principal components analysis separated sites into three main groups: rhithral streams, rhithral lake outlets, and kryal sites including outlets and streams. Rhithral streams contained the most diverse and taxon rich assemblages, being colonised by both non‐insect taxa and Ephemeroptera, Plecoptera, Trichoptera and Diptera. 3. Rhithral lake outlets supported high densities of non‐insect taxa such as Oligochaeta, Nemathelminthes and crustaceans. Despite low taxon richness, kryal sites had high Ephemeroptera and Plecoptera abundances. Chironomidae were most common at all sites. 4. Collector‐gatherers were dominant at all sites, whereas filter‐feeders were rare. Scrapers and shredders were more common in streams than lake outlets. 5. Water temperature and algal standing crops were higher at rhithral lake outlets than rhithral streams, perhaps providing more favourable habitat for non‐insect taxa. Glacial runoff was the dominant factor influencing macroinvertebrate assemblages of kryal streams and kryal lake outlets. Alpine lakes influenced the environmental conditions of their outlets and, consequently, their macroinvertebrate assemblages unless being constrained by a glacial influence.     

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.     

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 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.     

Macroscale ecotone effects for the stream macrobenthic fauna and communities of the eastern Novosibirsk region

It is suggested that the eastern Novosibirsk region appeared as a margin between the Western and the Eastern Palaearctic zones in regard to the faunas of stream communities of rheophilic invertebrates. The western (European) elements of the fauna prevail in the lowland areas; but both faunas, eastern and western, combine at Salairsky Ridge. In addition, some differences between these two faunas are observed, when the species of the western origin dominate in the soft-bottom and phytophylic communities and refer to the European fauna. On the other hand, the eastern Siberian species are the key elements in the rhithral (hard-bottom) communities and somehow in krenal communities. Seventeen regional types of rheophilic communities are described.     

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.     

Bacterial structures and ecosystem functions in glaciated floodplains: contemporary states and potential future shifts

Glaciated alpine floodplains are responding quickly to climate change through shrinking ice masses. Given the expected future changes in their physicochemical environment, we anticipated variable shifts in structure and ecosystem functioning of hyporheic microbial communities in proglacial alpine streams, depending on present community characteristics and landscape structures. We examined microbial structure and functioning during different hydrologic periods in glacial (kryal) streams and, as contrasting systems, groundwater-fed (krenal) streams. Three catchments were chosen to cover an array of landscape features, including interconnected lakes, differences in local geology and degree of deglaciation. Community structure was assessed by automated ribosomal intergenic spacer analysis and microbial function by potential enzyme activities. We found each catchment to contain a distinct bacterial community structure and different degrees of separation in structure and functioning that were linked to the physicochemical properties of the waters within each catchment. Bacterial communities showed high functional plasticity, although achieved by different strategies in each system. Typical kryal communities showed a strong linkage of structure and function that indicated a major prevalence of specialists, whereas krenal sediments were dominated by generalists. With the rapid retreat of glaciers and therefore altered ecohydrological characteristics, lotic microbial structure and functioning are likely to change substantially in proglacial floodplains in the future. The trajectory of these changes will vary depending on contemporary bacterial community characteristics and landscape structures that ultimately determine the sustainability of ecosystem functioning.     

Disruption of a longitudinal pattern in environmental factors and benthic fauna by a glacial tributary

SUMMARY 1. In the upper Rhône catchment (Swiss Alps), modifications in the longitudinal pattern of environmental conditions and the benthic macroinvertebrate fauna were investigated in a glacier-fed stream (Rhône) at its confluence with a smaller glacier-fed tributary (Mutt) in June, August and September 1998. The distance to the source glacier was greater for the Mutt than for the Rhône.
2. Environmental conditions were harsher for the biota in the main stream upstream of the confluence than in the tributary. The tributary upstream of the confluence was characterised by higher taxonomic richness and abundance of the zoobenthos than the Rhône upstream.
3. Although environmental conditions in the main stream were little modified by the tributary, the fauna was richer and more diverse below the confluence. During the period of ice melt, colonisation from the Mutt led to the occurrence of faunal elements atypical of glacial streams in the main glacial stream upstream of the confluence, where water temperature remains below 4 °C.
4. Although contributing an average of only 10% to the Rhône discharge, the Mutt tributary is suggested to be the faunal driver of the system.     

Changes in the benthic fauna and flora after application of temephos to a stream on Mt. Tsukuba

A dose of 2 ppm of temephos in emulsifiable formulation was applied to a mountain stream to observe its effects on the zoobenthos and the subsequent changes in benthic algae. Most zoobenthos in the treated region drifted but there were slight differences among taxa. Ephemeroptera and Plecoptera disappeared but some Trichoptera and Diptera survived. Chironomids recolonized earlier than other invertebrates and reached a higher density level than before the treatment. A bloom of benthic algae occurred following the destruction of fauna. The increase in the standing crop of algae was accompanied by an increase in the number of species. Among these, Achnanthes lanceolata, Meridion circulare and Tetraspora gelatinosa were predominant. The end of the bloom began with the recovery of the zoobenthos population.     

Microdistribution of chironomids (Diptera: Chironomidae) in Alpine streams: an autoecological perspective

Chironomid communities from three glacial and three non-glacial high mountain streams in three Alpine river basins were analyzed (Conca, Niscli, Cornisello, NE Italy, 46°N, 10°E). Eighteen sampling reaches belonging to five stream types (kryal, subkryal, glacio-rhithral, kreno-rhithral, outlet) were investigated. At each reach, geomorphological, physical, chemical and biological data were collected. Field surveys were carried out during three periods per year from 1996 to 1998: immediately after spring snowmelt, in mid-summer and in early autumn. In all, 439 zoobenthos samples were collected from 5 to 10 microhabitats of 0.1m² in each reach and date using a standard pond net (mesh size of 250m). About 50% of individuals collected were chironomids (26673 specimens, 53 taxa), with densities ranging from 4 to 2652indm^–2. With few exceptions, they dominated as number of taxa and individuals in all reaches. Chironomid subfamilies Diamesinae and Orthocladiinae were most abundant, especially in glacial reaches, where Diamesa spp. constituted up to 100% of the total fauna. Chironomid distribution was analyzed in relation to 37 abiotic variables, referring to stream origin, hydrology, geomorphology, physics and chemistry. Diamesa steinboecki, D. latitarsis gr. A and Pseudokiefferiella parva were the taxa best associated with glacial conditions (i.e. high channel instability or presence of bedrock, high suspended solids and total phosphorous content, low conductivity and silica content, highly variable diel discharge and low mean temperature), while Pseudodiamesa branickii, Corynoneura spp., Eukiefferiella spp., Parorthocladius nudipennis, Tvetenia calvescens/bavarica, Thienemaniella spp. and Micropsectra atrofasciata were mostly associated with non-glacial conditions. Substratum particle size, water depth, current velocity, the presence of riffles/pools and of mosses/algae (Hydrurus foetidus) were the major factors affecting microdistribution of chironomids in the investigated streams.     

Macroinvertebrate community structure in relation to environmental variables in a Swiss glacial stream

1. Benthic macroinvertebrate distribution was examined in relation to channel characteristics (including stability), substratum, hydraulic variables, primary production (chlorophyll a ) and coarse particular organic matter (CPOM) in an alpine glacial stream, the Mutt (Upper Rhône valley, Switzerland). Co-inertia analysis and canonical correspondence analysis were used to identify the major environmental gradients influencing community variations.
2. The Mutt (length: 3.6 km, altitudinal range: 1800–3099 m a.s.l.) exhibited typical characteristics of a kryal stream. Average summer temperature remained below 2 °C immediately downstream from the snout but was on average 5 °C higher 1700 m downstream. Seasonal variations in water sources were evidenced by the high late-summer (September) contribution of groundwater with increased conductivity.
3. Sixty-six taxa were recorded from the five reaches sampled at three periods (snowmelt, ice melt and low water in late summer) in 1996 and 1997, of which 29 were Chironomidae. Three taxa of Diamesinae were the first colonizers of the stream below the glacier, but 16 taxa, including Ephemeroptera, Plecoptera and Trichoptera, were already recorded 200 m downstream. Water depth, channel slope and Pfankuch's Index of channel stability were strongly correlated with the longitudinal faunal gradient. Maximum temperature, current velocity and water conductivity were also correlated, but to a lesser extent.
4. The rapid incorporation of non-chironomid taxa into the stream community represented a departure from Milner & Petts's (1994) conceptual model of invertebrate succession downstream of glacial margins. The results confirmed that glacial stream communities are primarily driven by physical determinants.     

Chironomid (Diptera: Chironomidae) communities in six European glacier-fed streams

1. A study on glacial stream ecosystems was carried out in six regions across Europe, from Svalbard to the French Pyrenees. The main aim was to test the validity of the conceptual model of 38 with regard to the zonation of chironomids of glacier‐fed rivers along altitudinal and latitudinal gradient. 2. Channel stability varied considerably, both on the latitudinal and altitudinal scale, being lowest in the northern regions (Svalbard, Iceland and Norway) and the Swiss Alps. Water temperature at the upstream sites was always <2 °C. 3. There was a prominent difference in taxonomic richness between the Alpine and the northern European regions, with a higher number of taxa in the south. In all regions, the chironomid community was characterized by the genus Diamesa and the subfamily Orthocladiinae. Of a total of 63 taxa recorded, two (Diamesa bertrami and Orthocladius frigidus) were common in all the regions except Svalbard. 4. On the basis of cluster analysis, seven distinct groups of sites were evident amongst glacial‐fed systems of the five regions (Pyrenees excluded). This classification separated the glacier‐fed streams on geographical, latitudinal and downstream gradients. 5. Canonical Correspondence Analysis (CCA) of environmental variables was carried out using 41 taxa at 105 sites. Slope, water depth, distance from source, water temperature and the Pfankuch channel stability index were found to be the major explanatory environmental variables. The analysis separated Diamesinae and typical upstream orthoclads from the other chironomids by low temperature and high channel instability. 6. In all six regions, Diamesa was present closest to the glacier. Within 200 m of the glacier snout, other genera of Diamesinae were found together with Orthocladiinae. Pioneer taxa like Diamesa species coexisted with later colonizers like Eukiefferiella minor/fittkaui in relatively unstable channels. 7. The longitudinal succession of chironomid assemblages across altitudinal and latitudinal gradients in glacial streams followed the same pattern, with similar genera and groups of species. The general aspects of the conceptual model of 38 were supported. However, Diamesa species have wider temperature limits than predicted and other Diamesinae as well as Orthocladiinae colonize metakryal habitats.     

Ammonium uptake in alpine streams in the High Tatra Mountains (Slovakia)

Uptake of NH _inf4 ^sup+ -N by streambed biota of mountain brooks was studied in the alpine zone of the High Tatra Mountains. Experiments were performed involving in situ dosing of ammonium directly to the acidified stream and incubations of ammonium and streambed bryophytes in enclosures within a range of pH from 4.45 to 8.10.NH _inf4 ^sup+ -N uptake length decreased with decreasing stream discharge, while comparable values of discharge-normalized uptake lengths were found during two in situ experiments.Maximum uptake rates of NH _inf4 ^sup+ -N obtained during the incubation of bryophytes (6 to 11 mg m^–2 h^–1) were comparable with results of two in situ experiments (8 and 12 mg m^–2 h^–1). The average NH _inf4 ^sup+ -N uptake rates observed during incubations lasting 3 to 5 hours (4.3 mg m^–2 h^–1) were not related to the pH of stream water. Nitrification of about 50% of the NH _inf4 ^sup+ -N added was observed in non-acidified streams, but was negligible in acidified streams. Significant photoinhibition of nitrification was observed in non-acidified streams during enclosure experiments.     

Pleistocene origin and population history of a neoendemic alpine butterfly

Alpine environments underwent dramatic transformation during glacial–interglacial cycles, with the consequence that geographical, ecological and demographic changes of alpine populations provided the opportunity for formation of neoendemic species. Several biogeographical models have been proposed to account for the unique history of alpine populations, with different expectations of genetic divergence and speciation. The expanding alpine archipelago model proposes that alpine populations expand spatially and demographically during glacial events, dispersing between mountain ranges. Under this model, alpine populations are unlikely to diverge in isolation due to substantial interpopulation gene flow. In contrast, the alpine archipelago refuge model proposes that gene flow during glacial phases is limited and populations expand demographically during interglacial phases, increasing genetic isolation and the likelihood of speciation. We assess these models by reconstructing the evolutionary history of Colias behrii, a morphologically and ecologically distinct alpine butterfly restricted to the California Sierra Nevada. C. behrii exhibits very low genetic diversity at mitochondrial and nuclear loci, limited population structure and evidence of population expansion. C. behrii and Rocky Mountain C. meadii share identical mitochondrial haplotypes, while in contrast, nuclear data indicate common ancestry between C. behrii and Cascades Range Colias pelidne. The conflict in gene genealogies may be a result of recent expansion in North American Colias, but an isolation with migration analysis indicates that genetic patterns in C. behrii might result from differential introgression following hybridization. Based on the timing of population expansion and gene flow between mountain ranges, the expanding alpine archipelago model is supported in C. behrii.     

THE BENTHIC MACROINVERTEBRATE FAUNA OF A SOUTH AFRICAN MOUNTAIN STREAM AND ITS RESPONSE TO FIRE

Summary

The effects of a late-summer prescribed burn on the temperature and benthic macroinvertebrate fauna of a south-western Cape mountain stream were investigated over a period of 12 months. Temperature and discharge regimes appear well-defined and relatively predictable from year to year. As in other mediterranean-type ecosystems, seasonal changes in the structure of the invertebrate community and the relative abundance of different feeding groups appear to be associated primarily with changes in the physical environment. Distinctive summer and winter communities were identified, with chironomids dominating the fauna in summer and simuliids dominant in winter. Although the riparian vegetation was only slightly damaged by the fire, a heavy, aseasonal leaf-fall occurred shortly afterwards. The canopy remained sparse for approximately four months. Stream temperature in the post-burn year was not demonstrably affected by increased exposure to solar radiation, however, probably because the canopy remained open during the winter months. The fire appeared to have little effect on the invertebrate fauna. Apart from five rare elements of the biota, all species recorded in the pre-burn year were present in the post-burn year and in similar densities. It is concluded that the riparian vegetation is of major importance in maintaining the integrity of the stream environment.     

On the abundance and activity pattern of zoobenthos inhabiting a tropical reef area,Cebu, Philippines

A benthic faunal study was carried out in the tidal area of Mactan Island (Cebu, Philippines). The area was subdivided along a transect from the beach to the reef according to benthic assemblages. The sediments are largely composed of calcareous skeletal remains of the indigenous biota and surrounding calcareous rocks. The content of protein and carbohydrates of the sediment was estimated, providing an approximation of organic matter in terms of feeding efficiency. Total number of zoobenthos, both as regards the sediment samples and as to the epifaunal communities associated with seaweeds, is rather uniformly distributed justifying the 95% confidence level (P>0.05). Distinct differences are apparent in abundance values of individual taxa. Although the study area showed the expected distribution pattern, with dominance of Nematoda (39%) living in sediment and Harpacticoida (36–66%) dwelling on Thalassia and algae, Polychaeta reveal a dominant attraction to both these habitats. The reasons for this phenomenon are discussed in relation to the absolute lack of macrofaunal predators The zoobenthos adjust their distribution and activity to fluctuating conditions of the environment. Light is mainly suggested as stimulating diel migration activities of the benthic fauna, moving upwards from the sediment to the algae and Thalassia during daytime. In a field experiment the zoobenthos was investigated for digestion activity over a diurnal cycle. The results reveal that feeding activity of zoobenthos follows a diel cycle showing maximum activity during the morning and evening obviously influenced by changes of light.     

Stream habitat fragmentation — a threat to biodiversity

Biodiversity is undisturbed rhithral streams in central Europe is high, with about 1000 resident metazoan species; over 600 insect species occur in the Fulda river (Germany). Longitudinal downstream shift of dominance from rheobiontic to rheophilous and finally to ubiquituos rheoxenic taxa in the potamal is described. Present downstream importance of ubiquituous species probably results from replacement of original potamal communities, present faunas being surrogates. Species losses through human impact are well documented for fish. The case of Plecoptera (10 potamal species either altogether extinct, extinct in Central Europe or extremely endangered) suggests that potamal invertebrates suffered as severe losses as did fish.Human impact on major rivers was so severe also because they occur at distances beyond average dispersal capacity of the fauna, i.e. are widely separate ecological islands, with known risk of species losses. In contrast, faunal exchange between adjacent headwater streams in mountains with intact stream nets is easy, certainly for amphibious insects. However, damage to rhithral streams is becoming increasingly frequent. This fragments stream nets, turning also upper parts of drainage systems into ecological islands, with danger of extinctions. Rhithral biodiversity is thought to be much more endangered by human impact than is presently recognized.