Effects of emergence phenology, taxa tolerances and taxonomic resolution on the use of the Chironomid Pupal Exuvial Technique in river biomonitoring

1. We studied chironomid communities of four rivers in south‐eastern Finland, differing in their water quality, during summer 2004 using the Chironomid Pupal Exuvial Technique, CPET. The aims of the study were to (i) test the adequacy of the generic‐level identification in the CPET method, (ii) define the emergence patterns of chironomid taxa classified as intolerant to organic pollution, (iii) assess the tolerance levels of intolerant chironomids and (iv) identify taxa most indicative of good water quality. 2. Procrustean rotation analysis indicated very strong concordance between the ordinations using either species or genus‐level data, suggesting that generic‐level identification of chironomids is adequate for biomonitoring based on CPET. However, when only a few taxa occur in great numbers, it may be advisable to identify these to the species level, especially if these taxa are important indicators of the impact in question. 3. The detection of a particular species may require accurate timing of sampling, whereas a species‐rich genus might be detected throughout a season. Given that the emergence of chironomid species may vary from year‐to‐year and between sampling sites, community differences detected at the species level may be related to between‐site variation in species’ emergence patterns rather than true differences in species composition. 4. Indicator species analysis (IndVal) showed that the distribution and abundance of intolerant chironomid taxa differed strongly among the studied rivers. Some of the intolerant taxa were restricted to unimpacted conditions, whereas others occurred mainly in impacted rivers. Thus, the indicator status of some genera (e.g. Eukiefferiella, Parametriocnemus, Stempellinella and Tvetenia) needs reassessment.     

Grazing by oligochaetes and snails on epiphytes

SUMMARY. 1. The isotope ³³P was used to assess the effect of grazitig by oligochaetes (mainly Stylaria lacttstris L.) and the snail, Lymnaea peregra (Muller), on epiphytes within an Equisetum fluviatile L. stand.
2. Two 1 m² polystyrene enclosures were set up within the emergent macrophyte zone of the lake. At the start of the experiment ³³P-solution was mixed with the water in both tanks. Algal and animal samples for ³³P- analysis were collected during the peak occurrence of epiphytes in June.
3. Phosphorus release rates from the animals through defaecation and excretion were measured in the laboratory. The grazing rate of oligochaetes was 2.2–4.1 mg P m⁻² (of bottom) d⁻¹ of which about two - thirds was released and recycled. The oligochaete density ranged from 13,400 to 20,900 m⁻². The snails (25 ind. m⁻²) grazed 1.2-1.5 mg P m⁻²d⁻¹of whieh about a quarter was released through defaecation and excretion.
4. Daily consumption by the oligochaetes and snails corresponded to 22–45% of the average phosphorus uptake by the epiphytes.     

Reduction of ecosystem productivity and respiration during the European summer 2003 climate anomaly: a joint flux tower, remote sensing and modelling analysis

The European CARBOEUROPE/FLUXNET monitoring sites, spatial remote sensing observations via the EOS‐MODIS sensor and ecosystem modelling provide independent and complementary views on the effect of the 2003 heatwave on the European biosphere's productivity and carbon balance. In our analysis, these data streams consistently demonstrate a strong negative anomaly of the primary productivity during the summer of 2003. FLUXNET eddy‐covariance data indicate that the drop in productivity was not primarily caused by high temperatures (‘heat stress’) but rather by limitation of water (drought stress) and that, contrary to the classical expectation about a heat wave, not only gross primary productivity but also ecosystem respiration declined by up to more than to 80 gC m⁻² month⁻¹. Anomalies of carbon and water fluxes were strongly correlated. While there are large between‐site differences in water‐use efficiency (WUE, 1–6 kg C kg⁻¹ H₂O) here defined as gross carbon uptake divided by evapotranspiration (WUE=GPP/ET), the year‐to‐year changes in WUE were small (<1 g kg⁻¹) and quite similar for most sites (i.e. WUE decreased during the year of the heatwave). Remote sensing data from MODIS and AVHRR both indicate a strong negative anomaly of the fraction of absorbed photosynthetically active radiation in summer 2003, at more than five standard deviations of the previous years. The spatial differentiation of this anomaly follows climatic and land‐use patterns: Largest anomalies occur in the centre of the meteorological anomaly (central Western Europe) and in areas dominated by crops or grassland. A preliminary model intercomparison along a gradient from data‐oriented models to process‐oriented models indicates that all approaches are similarly describing the spatial pattern of ecosystem sensitivity to the climatic 2003 event with major exceptions in the Alps and parts of Eastern Europe, but differed with respect to their interannual variability.     

Influence of geographical extent on typology- and model-based assessments of taxonomic completeness of river macroinvertebrates

1. River InVertebrate Prediction and Classification System (RIVPACS)‐type predictive models are increasingly used to assess the biological condition of freshwaters, but management schemes may also be based on a priori groupings of similar water bodies (typologies) to control for natural variation in biota. The two approaches may lead to disagreements of the biological status of a site, depending on, for example, the spatial scale at which assessments are conducted. 2. We used data from 96 reference and 134 potentially impacted sites from Western and Central Finland to compare RIVPACS‐type models and a simple size‐based typology of rivers for the assessment of taxonomic completeness (the quotient of the Observed‐to‐Expected number of predicted taxa, O / E) of riffle macroinvertebrates. We specifically examined how geographical extent influences bioassessment performance (accuracy, precision and sensitivity to detect impact) of the two approaches. To fully examine the behaviour of the O / E‐index with the two approaches at differing spatial scales, we performed all assessments with a full range of thresholds for predicted taxa occurrence probabilities (p_t from 0+ to 0.9). 3. Both approaches performed consistently better than the corresponding null models. At the larger extent (i.e. assessment encompassing the whole study area), the RIVPACS‐approach performed in all aspects better than the typology‐approach. However, at the smaller extent (i.e. regional assessments) the RIVPACS‐type models and the typologies showed similar accuracy to predict the actual fauna (mean E), similar precision (SD) of cross‐validated O / E and similar sensitivity to detect sites with human impairment. 4. SD(O / E) decreased (i.e. precision increased) consistently with increasing p_t. However, both approaches were most sensitive at intermediate p_t:s (c. 0.2–0.6) when taxa with low predicted occurrence probabilities were excluded. 5. Our results show that RIVPACS‐type predictive models are less susceptible to variations in spatial scale, whereas the performance of a priori typologies increases with decreasing spatial extent. Thus, RIVPACS‐type models are more efficient for large‐scale bioassessments, but at restricted spatial scales, or with an otherwise biologically meaningful stratification, simple a priori classifications can be equally useful for the assessment of taxonomic completeness of river macroinvertebrates.     

Cladoceran community responses to biomanipulation and re‐oligotrophication in Lake Vesijärvi,Finland, as inferred from remains in annually laminated sediment

1. We studied the role of zooplankton in biomanipulation and the subsequent recovery phase in the Enonselkä basin of Lake Vesijärvi, using subfossil cladocerans in annually laminated sediment. Measures to restore the Enonselkä basin included reduction in external nutrient loading and mass removal of plankti‐ and benthivorous fish. Water clarity increased and the lake changed from a eutrophic to a mesotrophic state. However, some signs of increased turbidity were observed after 5–10 years of successful recovery. 2. Annual laminae in a freeze core sample were identified and sliced, based on the seasonal succession of diatoms. Cladoceran remains and rotifer eggs were counted, and Daphnia ephippia and Eubosmina and Bosmina ephippia and carapaces were measured. Annual changes in pelagic species composition were studied with principal component analysis. Individual species abundance, size measurements and various cladoceran‐based indices or ratios (commonly used to reconstruct changes in trophic state and fish predation) were tested for change between four distinct periods: I (1985–1988) dense fish stocks, poor water quality; II (1989–1992) fish removal; III (1993–1997) low fish density, improved water quality; IV (1998–2002) slightly increased fish density and poorer water quality. 3. After the removal of fish, the mean size of Daphnia ephippia and Eubosmina crassicornis ephippia and carapaces increased significantly. In contrast, the percentage of Daphnia did not increase. When based on ephippia, the ratio Daphnia/(Daphnia + E. crassicornis) increased, but the interpretation was obscured by the tolerance of fish predation by small Daphnia and by the fact that bosminids were the preferred food of roach. Moreover, ephippial production by E. crassicornis decreased in recent years. 4. The abundance of Diaphanosoma brachyurum and Limnosida frontosa increased significantly after the fish population was reduced, while that of Ceriodaphnia and rotifers decreased. 5. The expanding littoral vegetation along with improved water clarity was clearly reflected in the concentration of littoral species in the deep sediment core. The species diversity index for the entire subfossil community also increased. 6. The period of faltering recovery was characterised by greater interannual variability and an increased percentage of rotifers. Nevertheless, the mean sizes of Daphnia ephippia and E. crassicornis ephippia and carapaces indicated a low density of fish. The deteriorating water quality was apparently related to multiple stressors in the catchment after rehabilitation, such as intensified lakeshore building, as well as to exceptional weather conditions, challenging the management methods in use.     

Foraging activity and dietary spectrum of wood ants (Formica rufa group) and their role in nutrient fluxes in boreal forests

Abstract 1. We monitored three different‐sized wood ant (Formica aquilonia Yarrow) mounds over a 3‐year period in Finnish boreal forests dominated by Norway spruce (Picea abies Karst.), to assess the seasonal temperature dependency of ant activity. Additionally, we also monitored Norway spruce trees around the mounds for descending honeydew foragers. 2. The amount of collected honeydew and prey and its composition, as well as the carbon (C), nitrogen (N), and phosphorus (P) in honeydew and invertebrate prey was also investigated. 3. The number of warm days (average temperature above 20 °C) and the amount of precipitation differed among the years. Ant activity at the mounds (but not on the trees) was highly correlated with air temperature throughout the ant‐active season (May–September), but ant activity in spring and autumn was lower than in summer at similar temperatures. During all 3 years, honeydew played a major role in wood ant nutrition (78–92% of dry mass). Invertebrate prey was mainly Diptera (on average 26.2%), Coleoptera (12.5%), Aphidina (9.3%), and Arachnoida (8.5%). 4. The total amounts of C, N, and P input brought into the ant mounds in the form of food (both honeydew and prey) on the stand level were 12.6–39.0, 1.6–4.6 and 0.1–0.4 kg ha^?1 year^?1, respectively, which is equivalent to 2–6%, 12–33% and 27–58% of the fluxes in annual needle litterfall in typical boreal Norway spruce forests. Thus, wood ants can play a significant role in short term and local N and P cycling of boreal forest ecosystems.     

Sugar transport together with environmental conditions controls time lags between xylem and stem diameter changes

In the present study the seasonal patterns of time lags between diurnal xylem and whole stem diameter variations at the top and at the base of two Scots pine trees (Pinus sylvestris L.) were compared. The diameter variations were measured during the summers of 2001 and 2002. Time lags were determined using the cross‐correlation method. The lags were found to vary in time according to the different stages of growth. At the top the xylem lagged behind the whole stem between the beginning of stem growth and the end of shoot growth in both years. In 2001 the time lags at the base showed a similar behaviour during stem growth. That kind of seasonal pattern of the time lags would result from the changes in the sink strength due to changing growth rate at different parts of the tree and the differences in the annual rhythm of growth and water availability in the soil (based on precipitation measurements) between the years 2001 and 2002 were reflected in the patterns. The time lags of shrinking and swelling periods during high and low photosynthetic activity (measured using a shoot chamber) were also compared. It was found, for example, that in 2001 in the middle of the growing season at the top of the tree the whole stem lagged on average 15 min more behind the xylem on the days of high photosynthetic activity than on the days of low or moderate. These results show for the first time that the transportation of carbohydrates and variable sink activity could be detected during the growing season in field conditions using stem and xylem diameter variation measurements. Furthermore, these results provide evidence of the pressure gradient‐driven flow also in the phloem of gymnosperms.     

Differential vulnerability determines the diet of a slow-moving predatory stream insect

1. We investigated the diet and prey electivity of Rhyacophila obliterata, a slow‐moving invertebrate predator capable of hunting in high‐flow microhabitats, and quantified the components of the predation sequence of fifth‐instar larvae foraging on mobile (Baetis mayflies, Amphinemura stoneflies) versus semi‐sessile (larval blackflies) prey. 2. In the field, fifth‐instar Rhyacophila consistently took more larval blackflies than more mobile prey. In behavioural trials, the number of attacks by Rhyacophila differed significantly between prey types, mobile prey being attacked more often than blackflies. Capture success, by contrast, was highest for blackflies, whereas Amphinemura and Baetis were rarely captured. In mixed‐prey feeding trials, Rhyacophila showed strong preference for blackflies and equally strong avoidance of Amphinemura and Baetis. 3. For mobile prey, the risk of being captured by this sluggish predator is very low, so they can afford to be in close contact with it. Rhyacophila was almost unable to capture any other prey but blackflies, resulting in strong passive selection for blackflies. 4. Therefore, the diet of fifth‐instar Rhyacophila can be predicted from laboratory observations and prey behaviour is the major determinant of the diet of this invertebrate predator.     

CO2 balance of boreal, temperate, and tropical forests derived from a global database

Terrestrial ecosystems sequester 2.1 Pg of atmospheric carbon annually. A large amount of the terrestrial sink is realized by forests. However, considerable uncertainties remain regarding the fate of this carbon over both short and long timescales. Relevant data to address these uncertainties are being collected at many sites around the world, but syntheses of these data are still sparse. To facilitate future synthesis activities, we have assembled a comprehensive global database for forest ecosystems, which includes carbon budget variables (fluxes and stocks), ecosystem traits (e.g. leaf area index, age), as well as ancillary site information such as management regime, climate, and soil characteristics. This publicly available database can be used to quantify global, regional or biome‐specific carbon budgets; to re‐examine established relationships; to test emerging hypotheses about ecosystem functioning [e.g. a constant net ecosystem production (NEP) to gross primary production (GPP) ratio]; and as benchmarks for model evaluations. In this paper, we present the first analysis of this database. We discuss the climatic influences on GPP, net primary production (NPP) and NEP and present the CO₂ balances for boreal, temperate, and tropical forest biomes based on micrometeorological, ecophysiological, and biometric flux and inventory estimates. Globally, GPP of forests benefited from higher temperatures and precipitation whereas NPP saturated above either a threshold of 1500 mm precipitation or a mean annual temperature of 10 °C. The global pattern in NEP was insensitive to climate and is hypothesized to be mainly determined by nonclimatic conditions such as successional stage, management, site history, and site disturbance. In all biomes, closing the CO₂ balance required the introduction of substantial biome‐specific closure terms. Nonclosure was taken as an indication that respiratory processes, advection, and non‐CO₂ carbon fluxes are not presently being adequately accounted for.     

Impacts of a submerged plant (Elodea canadensis) on interactions between roach (Rutilus rutilus) and its invertebrate prey communities in a lake littoral zone

1. Using 5‐m² field enclosures, we examined the effects of Elodea canadensis on zooplankton communities and on the trophic cascade caused by 4–5 year old (approximately 16 cm) roach. We also tested the hypothesis that roach in Elodea beds use variable food resources as their diet, mainly benthic and epiphytic macroinvertebrates, and feed less efficiently on zooplankton. Switching of the prey preference stabilises the zooplankton community and, in turn, also the fluctuation of algal biomass. The factorial design of the experiment included three levels of Elodea (no‐, sparse‐ and dense‐Elodea) and two levels of fish (present and absent). 2. During the 4‐week experiment, the total biomass of euplanktonic zooplankton, especially that of the dominant cladoceran Daphnia longispina, decreased with increase in Elodea density. The Daphnia biomass was also reduced by roach in all the Elodea treatments. Thus, Elodea provided neither a favourable habitat nor a good refuge for Daphnia against predation by roach. 3. The electivity of roach for cladocerans was high in all the Elodea treatments. Roach were able to prey on cladocerans in Elodea beds, even when the abundance and size of these prey animals were low. In addition to cladocerans, the diet of roach consisted of macroinvertebrates and detrital/plant material. Although the biomass of macroinvertebrates increased during the experiment in all Elodea treatments, they were relatively unimportant in roach diets regardless of the density of Elodea beds. 4. Euplanktonic zooplankton species other than Daphnia were not affected by Elodea or fish and the treatments had no effects on the total clearance rate of euplanktonic zooplankton. However, the chlorophyll a concentration increased with fish in all the Elodea treatments, suggesting that fish enhanced algal growth through regeneration of nutrients. Thus, our results did not unequivocally show that Elodea hampered the trophic cascade of fish via lowered predation on grazing zooplankton. 5. In treatments with dense Elodea beds (750 g FW m^?2), chlorophyll a concentration was always low suggesting that phytoplankton production was controlled by Elodea. Apparently, the top‐down control of phytoplankton biomass by zooplankton was facilitated by the macrophytes and operated simultaneously with control of phytoplankton production by Elodea.