Intraspecific variation in the water relations of Salix arctica,an arctic-alpine dwarf willow

Summary The seasonal and diurnal water relations were investigated within arctic and alpine populations of the dwarf willow Salix arctica. Marked differences that were habitat dependent (e.g. xeric vs. mesic) occured both within and between the populations. The environmental variables that most affected plant water balance and the bulk tissue water relations were soil water potential (_soil) and the leafto-air water vapor pressure gradient (w), however, low soil temperature (<4.0° C) also had a marked effect in the wet to mesic habitats. The effects of declining _soil and increased w were most pronounced in the plants growing in xiric habitats in both populations. Stomatal response to increased w was two-fold greater in alpine versus arctic plants and is hypothesized to have arisen in response to more frequent exposure to the higher evaporative conditions of alpine existance. Seasonal fluctuations in the osmotic potential closely followed changes in _soil, suggesting that these were active rather that passive changes. Additionally, plants from xeric habiats had a lower bulk tissue elastic modulus (more elastic tissues) in both arctic and alpine populations. The osmotic and elastic properties enhanced turgor maintenance over a broad range of leaf water potentials and during periods when w was high. Turgor maintenance also correlated to continued transpiration despite fluctuations in soil and atmospherically induced water deficits. Arctic habitats have a shorter growing season, lower soil temperatures due to the presence of permafrost, but higher soil water potentials and lower leaf-to-air vapor pressure gradients than alpine habitats. The observed variation in patterns of stomatal conductance and in values of tissue water relations characteristics between arctic and alpine populations of S. arctica is hypothesized to have arisen in response to these different environmental regimes which represent different selective regimes that occur along the arctic-alpine environmental continuum inhabited by this wide ranging species.     

Divergence in germination traits among arctic and alpine populations of <Emphasis Type="Italic">Koenigia islandica</Emphasis>: light requirements

Light is known to regulate conservative germination strategies and the formation of seed banks. Although these strategies are crucial to survival in tundra environments—especially for annuals—light requirements for germination in arctic–alpine species are seldom investigated. Furthermore, environmental differences between arctic and alpine regions are expected to lead to evolutionary divergence among conspecific populations in seed germination strategies. In this study, we report important differences in germination light requirements among six arctic and alpine populations of the annual Koenigia islandica. Light had little effect on germination of the seeds from Iqaluit (Nunavut, Canada), Yukon (Canada), and Jasper (Alberta, Canada), whereas the seeds from the most severe climates, Svalbard (Norway) and Colorado (USA), had strong light requirements. Stratification of the seeds had little influence on their germination light requirements, with the exception of the population from Dovre (Norway), in which it induced a strong light requirement. Possible adaptive explanations and some implications of these observed germination patterns are discussed.     

The impact of temperature regimes on development,dormancy breaking and germination of dwarf shrub seeds from arctic,alpine and boreal sites

It has been suggested that the infrequent sexual reproduction of arctic dwarf shrubs might be related to the harsh environmental conditions in which they live. If this is the case, then increases in temperature resulting from global climate change might drastically affect regeneration of arctic species. We examined whether recruitment of Empetrum nigrum ssp. hermaphroditum and Vaccinium uliginosum (hereafter E. nigrum and V. uliginosum) was affected by temperature during three reproductive stages: seed development, dormancy breakage and germination. Seeds were collected from an arctic, an alpine (only E. nigrum) and a boreal site with different climates; stored at different winter temperatures and incubated for germination at different temperatures. Seeds of V. uliginosum developed in the boreal region had a higher percentage germination than did seeds developed in the Arctic. In contrast, seeds of E. nigrum from the arctic site had a higher or similar percentage germination than did seeds from the alpine and boreal sites. Increased winter temperatures had no significant effect on resulting germination percentage of E. nigrum. However, V. uliginosum seeds from the arctic site suffered increased fungal attack (and thus decreased germination) when they were stratified under high winter temperatures. Seeds of both species increased germination with increased incubation temperatures. Our results suggest that both species would increase their germination in response to warmer summers. Longer summers might also favour the slow-germinating E. nigrum. However, increased winter temperatures might increase mortality due to fungal attack in V. uliginosum ecotypes that are not adapted to mild winters.     

Habits and relative population densities of some hydrophilids in California rice fields

Populations of the hydrophilids Berosus styliferus Horn, Hydrophilus triangularis Say, and Tropisternus lateralis (Fabricius) were observed in northern California rice paddies. Relative adult and larval density for each species was determined using aquatic light traps adjacent to the water margin, and at 5 m and 30 m distances from the levee. Egg cases were monitored by recording numbers per unit area.The seasonal patterns of abundance indicate each species has a univoltine life cycle. All stages of B. styliferus and H. triangularis are more abundant nearer the levees. T. lateralis adults are commonly found at the levees during the first 10 weeks post-flood, whereas T. lateralis larvae and egg cases are most numerous at the 30 m distance.Colonication of the paddies occurs through water connections, flight, and adult overwintering.     

Overcrowding as one of the causes of dispersal of young Tree Sparrows

Capsule Low and variable encounter rates of birds in fragmented arctic‐alpine habitats add difficulty to monitoring their breeding populations.

Aims To quantify seasonal variation in the encounter rates (apparent abundance) of breeding birds in arctic‐alpine habitats in Scotland.

Methods Birds were sampled from 15 repeated linear transects between April and August in 2005 and 2006. glmms (and for scarcer species glms) were used to investigate how the apparent abundance of different species varied between months and years.

Results Three arctic‐alpine specialists (Rock Ptarmigan, Eurasian Dotterel and Snow Bunting) were recorded. The 24 other species recorded included more widely distributed upland species, generalists that also used arctic‐alpine habitats and also some transient species from lower altitude. Overall encounter rates were low (only exceeding 1 bird km^?1 in any month for one species; Meadow Pipits) with marked variation between months. The pattern of seasonal variation in encounter rates varied markedly between species.

Conclusions Low encounter rates and marked variation in apparent abundance will render more difficult efforts to monitor birds in marginal and fragmented areas of arctic‐alpine habitats. Particularly relevant is the potential for changes in the timing of breeding and seasonal movements to influence encounter rates and be falsely interpreted as changes in actual abundance. Monitoring in arctic‐alpine habitats should include both specialist and non‐specialist birds of that habitat, as the latter may be more numerous and, therefore, provide supplementary evidence of temporal or seasonal change.     

Ecological and evolutionary responses of an arctic plant to variation in microclimate and soil

The arctic and alpine regions are predicted to experience some of the highest rates of climate change, and the arctic vegetation is expected to be especially sensitive to such changes. Understanding the ecological and evolutionary responses of arctic plant species to changes in climate is therefore a key objective. Geothermal areas, where natural temperature gradients occur over small spatial scales, and without many of the confounding environmental factors present in latitudinal and other gradient studies, provide a natural experimental setting in which to examine the response of arctic–alpine plants to increasing temperatures. To test the ecological and evolutionary response of the circumpolar alpine bistort Persicaria vivipara to temperature, we collected plant material and soil from areas with low, intermediate and high soil temperatures and grew them at three different temperatures in a three-factorial growth chamber experiment. At higher experimental soil temperatures, sprouting was earlier and plants had more leaves. Sprouting was earlier in soil originating from intermediate temperature and plants had more leaves when grown in soil originating from low temperatures. We did not find evidence of local adaptation or genetic variation in reaction norms among plants originating from areas with low, intermediate and high soil temperature. Our findings suggest that the alpine bistort has a strong plastic response to warming, but that differences in soil temperature have not resulted in genetic differentiation. The lack of an observed evolutionary response may, for example, be due to the absence of temperature-mediated selection on P. vivipara, the low rate of sexual recombination, or high levels of gene flow balancing differences in selection. When placed within the context of other studies, we conclude that arctic–alpine plant species often show strong plastic responses to spring warming, while evidence of evolutionary responses varies among species.     

高山林线交错带高山杜鹃的凋落物分解

凋落物分解是维持生态系统生产力、养分循环、土壤有机质形成的关键生态过程。高山林线交错带是陆地生态系统中对气候变化响应的敏感区域。季节变化和海拔梯度上的植被类型差异可能会影响该区域凋落物的分解,进而对高山生态系统的碳氮循环产生重要影响。采用凋落物分解袋的方法,研究了川西高山林线交错带优势种高山杜鹃(Rhododendron lapponicum)凋落叶在雪被期和生长季的分解特征。结果显示:(1)季节变化和植被类型对高山杜鹃凋落物的分解均具有显著影响(P0.05),凋落叶的质量损失主要发生在生长季且在高山林线最大,暗针叶林中雪被期的质量损失略高于生长季,但差异不显著;(2)林线交错带上高山杜鹃凋落叶分解缓慢,一年干物质失重率为9.62%,拟合分解系数k为0.145;(3)高山杜鹃凋落叶的质量变化主要体现在纤维素降解显著且集中在雪被期,木质素无明显降解,在高山林线上C/N、C/P、木质素/N变化幅度较小且C、N、P的释放表现得稳定而持续。结果表明,季节性雪被对林线交错带内高山杜鹃分解的影响不仅局限在雪被期内,雪被融化期间频繁的冻融作用和雪融水淋洗作用可能会促进高山杜鹃凋落物在生长季初期的分解。总的来看,在气候变暖的情景下,雪被的缩减、生长季的延长和高山杜鹃群落的扩张可能加速高山林线交错带高山杜鹃凋落物的分解。     

Instar sizes,life cycles,and food habits of five Rhyacophila (Trichoptera: Rhyacophilidae) species from the Appalachian Mountains of South Carolina,U.S.A.

The larval head widths at each instar, life cycles, and food habits of late instars were determined for five species of Rhyacophila from two Appalachian mountain streams in South Carolina, U.S.A. Rhyacophila acutiloba Morse & Ross was univoltine with two cohorts, one emerging in the spring and another presumably emerging in early autumn. Rhyacophila fuscula (Walker), R. nigrita Banks, and R. carolina Banks were apparently multicohort, univoltine species with extended flight periods. Rhyacophila minor Banks was univoltine with a spring emergence. All species were predaceous and consumed mainly Plecoptera nymphs and Trichoptera larvae.     

Demographic consequences of age-structure in extreme environments: population models for arctic and alpine ptarmigan

Organisms living in arctic and alpine environments are increasingly impacted by human activities. To evaluate the potential impacts of global change, a better understanding of the demography of organisms in extreme environments is needed. In this study, we compare the age-specific demography of willow ptarmigan (Lagopus lagopus) breeding at arctic and subalpine sites, and white-tailed ptarmigan (L. leucurus) breeding at an alpine site. Rates of egg production improved with age at the alpine and subalpine sites, but the stochastic effects of nest and brood predation led to similar rates of annual fecundity among 1-, 2-, and 3+-year-old females. All populations had short generation times (T<2.7 years) and low net reproductive rates (R ₀<1.2). Stable age distributions were weighted towards 1-year-old females in willow ptarmigan (>59%), and to 3+-year-old females in white-tailed ptarmigan (>47%). High damping ratios (ρ>3.2) indicated that asymptotic estimates were likely to match natural age distributions. Sensitivity and elasticity values indicated that changes in juvenile survival would have the greatest impact on the finite rate of population change (λ) in willow ptarmigan, whereas changes to the survival of 3+-year-old females would have a greater effect in white-tailed ptarmigan. High survivorship buffers white-tailed ptarmigan in alpine environments against the potential effects of climate change on annual fecundity, but may make the species more sensitive to the effects of pollutants or harvesting on adult survival. Conversely, processes that reduce annual fecundity would have a greater impact on the population viability of willow ptarmigan in arctic and subalpine environments. If these same demographic patterns prove to be widespread among organisms in extreme environments, it may be possible to develop general recommendations for conservation of the biological resources of arctic and alpine ecosystems.     

Caddisfly life histories along permanence gradients in high-altitude wetlands in Colorado (U.S.A.)

SUMMARY 1. Larvae of cased caddisflies (Limnephilidae and Phryganeidae) are among the most abundant and conspicuous invertebrates in northern wetlands. Although species replacements are often observed along permanence gradients, the underlying causal mechanisms are poorly understood. In this paper, we report on the distributional patterns of caddisflies in permanent and temporary high‐altitude ponds, and how those patterns reflect differences in life history characteristics that affect desiccation tolerance (fundamental niches) versus constraints related to biotic interactions (realised niches). 2. Species (Hesperophylax occidentalis and Agrypnia deflata) that were encountered only in permanent ponds are restricted in distribution by life history (no ovarian diapause, aquatic oviposition, and/or inability to tolerate desiccation). Although the egg masses of H. occidentalis tolerate desiccation, the larvae leave the protective gelatinous matrix of the egg mass because adults oviposit in water. 3. Three species (Asynarchus nigriculus, Limnephilus externus and L. picturatus) have life history characteristics (rapid larval growth, ovarian diapause and terrestrial oviposition of desiccation‐tolerant eggs) that should facilitate the use of both permanent and temporary habitats. However, A. nigriculus is rare or absent in most permanent ponds, and L. externus and L. picturatus are rare or absent in most temporary ponds. Experimental data from a previous study on the combined effects of salamander predation and interspecific interactions among caddisflies (e.g. intraguild predation) suggest that biotic interactions limit each species to a subset of potentially exploitable habitats. 4. Many wetland invertebrates exhibit species replacements along permanence gradients, but few studies have separated the relative importance of the effects of drying per se from the effects of biotic interactions. Our results emphasise the complementary roles of comparative data on life histories and experimental data on competition and predation for understanding invertebrate distributions along permanence gradients.     

Effects of season,food deprivation and re-feeding on leptin,ghrelin and growth hormone in arctic foxes (<Emphasis Type="Italic">Alopex lagopus</Emphasis>) on Svalbard,Norway

The arctic fox (Alopex lagopus) is a medium-sized predator of the high Arctic experiencing extreme seasonal fluctuations in food availability, photoperiod and temperature. In this study, the plasma leptin, ghrelin and growth hormone (GH) concentrations of male arctic foxes were determined during a food deprivation period of 13 days and the subsequent recovery in November and May. Leptin, ghrelin and GH were present in arctic fox plasma in amounts comparable to other carnivores. The plasma leptin concentrations did not react to food deprivation unlike in humans and rodents. However, the leptin levels increased during re-feeding as an indicator of increasing energy reserves. The relatively high ghrelin–leptin ratio, decrease in the plasma ghrelin concentration, an increase in the circulating GH concentrations and the observed negative correlation between plasma ghrelin and free fatty acid levels during fasting suggest that these hormones take part in the weight-regulation and energy metabolism of this species by increasing fat utilisation during food deprivation. The results strengthen the hypothesis that the actions of these weight-regulatory hormones are species–specific and depend on seasonality and the life history of the animals.Abbreviations FFA free fatty acid - GH growth hormone - RMR resting metabolic rate Communicated by G. Heldmaier     

Phylogeographic patterns in Leccinum sect. Scabra and the status of the arctic-alpine species L. rotundifoliae

We investigated inter- and intraspecific phylogenetic relationships in the ectomycorrhizal fungal genus Leccinum section Scabra. Species of this section are exclusively associated with Betula and occur throughout the Northern Hemisphere. We compared the phylogenetic relationships of arctic, alpine, boreal and temperate accessions of section Scabra based on DNA sequences of the single-copy nuclear gene Gapdh and the multiple-copy nuclear region 5.8S-ITS2. Exclusively arctic lineages were not detected in species that occur both in arctic-alpine or boreal regions, except in L. rotundifoliae that was restricted to cold climates. L. scabrum and L. holopus showed an intercontinental phylogeographic pattern, and L. variicolor showed a pattern unrelated to geographical distribution. Molecular clock estimates indicated that L. rotundifoliae is as old as other species in section Scabra. Individual gene trees suggest that interspecific hybridisation occurred several times in the evolution of section Scabra.     

The salinity tolerance of freshwater macroinvertebrate eggs and hatchlings in comparison to their older life-stages: a diversity of responses

The tolerance to high salinity of older life stage macroinvertebrates could underestimate the effects of increasing salinity on freshwater macroinvertebrates. The salinity tolerance of the eggs and/or hatchlings of Glyptophysa gibbosa (Planorbidae), Glyptophysa aliciae, Glacidorbis sp. (Glacidorbidae), a Glossiponiidae, Piona cumberlandis (Pionidae), and Chironomus sp. (Chironomidae) were determined. The salinity tolerances of younger life-stages of species studied here and elsewhere were then compared to salinity tolerances of their mature aquatic or dominant life-stage. A diversity of responses have been observed with some species having similar salinity tolerances in all life-stages examined, whilst the eggs or hatchlings of other species had salinity tolerances ranging from 4% to 88% of their older life stages. On present knowledge, this diversity of responses presents some difficulties for simple rules of thumb to approximate sensitivity of young life-stages of freshwater macroinvertebrates inferred from their dominant stage’s tolerance.     

Vulnerability of alpine stream biodiversity to shrinking glaciers and snowpacks

Climate change poses a considerable threat to the biodiversity of high latitude and altitude ecosystems, with alpine regions across the world already showing responses to warming. However, despite probable hydrological change as alpine glaciers and snowpacks shrink, links between alpine stream biota and reduced meltwater input are virtually unknown. Using data from the French Pyrénées, we demonstrate that taxonomic richness and total abundance of stream macroinvertebrates increase significantly as meltwater (snow melt and glacier melt) contributions to river flow decrease. Macroinvertebrate species showed a gradation of optimum meltwater conditions at which they persist. For example: Habroleptoides berthelemyi (Ephemeroptera), Perla grandis (Plecoptera) and Rhithrogena spp. (Ephemeroptera) increased in abundance when meltwater contributions to streamflow decrease, whereas in contrast, Rhyacophila angelieri (Trichoptera) and Diamesa latitarsis spp. (Diptera) decreased in abundance. Changes in alpine stream macroinvertebrate community composition as meltwater contributions decline were associated with lower suspended sediment concentration, and higher water temperature, electrical conductivity and pH. Our results suggest α diversity (at a site) of streams presently fed by meltwaters will increase with future meltwater reductions. However, β diversity (between‐sites) will be reduced as snow melt and glacier melt decrease because the habitat heterogeneity associated with spatiotemporal variability of water source contributions will become lower as meltwater contributions decline. Extinction of some endemic alpine aquatic species (such as the Pyrenean caddis fly R. angelieri) is predicted with reduced meltwater inputs, leading to decreases in γ diversity (region). Our identification of significant links between meltwater production and stream macroinvertebrate biodiversity has wider implications for the conservation of alpine river ecosystems under scenarios of climate change induced glacier and snowpack loss.     

Thermotolerance of Photosystem 2 of Three Alpine Plant Species Under Field Conditions

The species specific response of photosystem 2 (PS2) efficiency and its thermotolerance to diurnal and seasonal alterations in leaf temperature, irradiance, and water relations were investigated under alpine field conditions (1 950 m) and in response to an in situ long-term heat treatment (+3 K). Three plant species were compared using the naturally occurring microstratification of alpine environments, i.e. under contrasting leaf temperatures but under similar macroclimatic conditions. Thermotolerance of PS2 showed a high variability in all three species of up to 9.6 K. Diumal changes (increases or even decreases) in PS2 thermotolerance occurred frequently with a maximum increase of +4.8 K in Loiseleuria procumbens. Increasing leaf temperatures and photosynthetic photon flux density influenced thermotolerance adjustments. Under long-term heating (+3 K) of L. procumbens canopies with infra-red lamps, the maxima of the critical (T_c) and the lethal (T_p) temperature of PS2 increased by at least 1 K. Thermotolerance of the leaf tissue (LT₅₀) increased significantly by +0.6 K. The effects of slight water stress on thermotolerance of PS2 were species specific. High temperature thresholds for photoinhibition were significantly different between species and increased by 9 K from the species in the coldest microhabitat to the species in the warmest. Experimental heating of L. procumbens canopies by +3 K caused a significant (p>0.01) upward shift of the high temperature threshold for photoinhibition by +3 K. Each species appeared to be very well adapted to the thermal conditions of its microhabitat as under the most frequently experienced daytime leaf temperatures no photoinhibition occurred. The observed fine scale thermal adjustment of PS2 in response to increased leaf temperatures shows the potential to optimise photosynthesis under varying environmental conditions as long as the upper thermal limits are not exceeded.     

SYSTEMATIC INFERENCES FROM VARIATION IN ISOZYME PROFILES OF ARCTIC AND ALPINE CESPITOSE FESTUCA (POACEAE)

Variation in isozyme patterns was used to assess species boundaries in North American arctic and alpine representatives of the Festuca ovina L. complex. Isozyme profiles, in combination with chromosome number, delimit four discrete entities within the complex: F. brevissima Jurtzev (diploid); F. aggr. auriculata Drobov (diploid); F. brachyphylla Schultes (hexaploid); and tetraploid populations corresponding in morphology to F. baffinensis Polunin (arctic Canada) and F. minutiflora Rydberg (alpine United States). Although no fixed difference was detected between isozyme profiles of the latter two taxa, they are morphologically distinct. Thus variation in isozymes, morphology, and chromosome number delimits five taxa within the F. ovina complex in North America. Some alleles observed in the polyploid taxa were not detected among the diploids, and some observed in F. brachyphylla, the hexaploid taxon, were not detected in either the diploid or the tetraploid species. One possible explanation for these occurrences is that the North American polyploids originated in Eurasia, where many other potential diploid and tetraploid progenitors occur.     

Factors influencing species diversity in saline waters of Death Valley,USA

Salinity is a major factor influencing the distributions and abundances of aquatic macroinvertebrates of saline waters in Death Valley, California, USA. A general pattern of declining numbers of species with increasing salinity is seen in Death Valley waters. Some species are restricted to low salinities, others are found only in highly saline pools, and still others are widely distributed over a broad range of salinities.Salinity alone cannot explain distributions seen in the field. Distributions and abundances of species such as the caddisfly Limnephilus assimilis Banks are broader than would be predicted on the basis of laboratory studies of salinity and temperature. I present evidence that for such species, biotic factors such as reduced predation at high salinities may compensate for increased physiological stress.     

Ecological life histories of the three aquatic nuisance plants,Myriophyllum spicatum,Potamogeton crispus and Elodea canadensis

The life histories of Myriophyllum spicatum L., Elodea canadensis Michx., and Potamogeton crispus L., serious aquatic nuisances in many regions of the world, are reviewed to provide insights into the life style of successful aquatic nuisance plants. Specifically, their distribution and spread in North America; their life cycle, productive and reproductive potential; and their ecosystem relationships are reviewed. Hopefully this review will improve a manager's ability to deal with aquatic nuisance problems. It also provides suggestions for basic research needed to develop more effective management practices.It was found that all three species possess a number of adaptations, including an ability to rapidly propagate vegetatively, an opportunistic nature for obtaining nutrients, a life cycle that favors cool weather, and a number of mechanisms which enhance photosynthetic efficiency, which allow them to proliferate.These three species do provide benefits to the ecosystem through their roles in materials cycling and energy flow. Therefore, management of these species should take an integrated approach which recognizes these benefits.The life history information available about the three species varies tremendously; however, a better understanding of resource gain and allocation is needed to manage all three species. Specifically, more research is needed to provide a better understanding of: 1) the role bicarbonate plays in photosynthesis, 2) the role roots play in supplying CO₂ to the plabts, 3) resource accumulation and allocation under different temperature and light regimes, 4) resource allocation on a seasonal basis, and 5) nutrient cycling under different management regimes.     

Environmental drivers of invertebrate population dynamics in Neotropical tank bromeliads

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Aquatic macroinvertebrate community structure of aNymphoides peltata-dominated and macrophyte-free site in an oxbow lake

The aquatic macroinvertebrates in two freshwater biotopes,viz. aNymphoides peltata-dominated site and a macrophyte-free site, were studied quantitatively in a shallow alkaline oxbow lake of the river Waal, the main branch of the river Rhine in The Netherlands. The research comprised the analysis of water, sediment and macrophyte samples.In the macrophyte-free site Oligochaeta and Nematocera, particularly of the collector gatherer functional feeding group, dominated the prevailing benthic community. The total macroinvertebrate biomass ranged here from 0.3 to 0.9 g ash-free dry weight per m² of biotope.Species richness, densities, and biomass of macroinvertebrates were considerably higher in the biotope dominated byNymphoides peltata. Many taxa were found associated with the aboveground macrophyte. The sediment compartment, however, contributed most to the total density and biomass of macroinvertebrates. Nematocera and Oligochaeta were the most abundant fauna groups, whereas the largest share in total biomass was provided by clams (Mollusca). The biomass of the total macroinvertebrate community in theNymphoides-dominated site ranged from 6.2 to 7.5 g ash-free dry weight per m² of biotope. The biomass of the aboveground phytophilous fauna ranged from 0.1 to 0.6 g ash-free dry weight per m² of biotope. In September, when theNymphoides peltata vegetation was in its senescent phase, the largest numbers and the highest biomass of phytophilous macroinvertebrates were observed. The contribution of the shredder functional feeding group was high in this period. This, and the overall high abundance of fauna with a detritivorous mode of life, indicates the importance of macrophyte detritus as input to food chains.