Ecology of alpine streams

SUMMARY 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 max 4°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