Population dynamics and energy budget of Marionina southerni (Cernosvitov) (Enchytraeidae,Oligochaeta) in the shallow littoral of Lake Esrom,Denmark

Marionina southerni (Cernosvitov) was numerically the dominant oligochaete in a shallow, strongly exposed surf zone locality (0–1 m depth) in the mesotrophic Lake Esrom. It comprised 21,500 ind m^–2 or nearly 50% of the total oligochaete community, which otherwise was dominated by Nais spp. and the lumbriculid Stylodrilus heringianus Claparède. M. southerni appears to have a 1-y life cycle. Individual biomass ranged from 3 to 48 µg ash free dry weight. Annual net production at the site was 5.1 kcal m^–2 (size-frequency method) with a P/B ratio of 2.5. This is about 0.5% of the estimated mean zoobenthic production in the littoral zone from 0 to 2 m depth in Lake Esrom. At 5° C M. southerni showed a constant oxygen uptake down to 25% oxygen saturation, but practically no regulatory respiration was found at 20 °C. In the field M. southerni was never found in sediment with less than 60% oxygen saturation. The annual community respiration of M. southerni was 14.2 kcal m^–2, and annual assimilation thus made up 19.3 kcal m^–2 with a net production efficiency of 26.5%.     

Abundance,population dynamics and production of Chironomidae (Diptera) in an ul traoligotrophic lake in South Greenland

Macrozoobenthos of the ultraoligotrophic Lake 95 (61°N, 46°W, 8 ha, z_max=18 m, ) is composed of about 14 taxa dominated by 12 Chironomidae species. Abundance, life cycle, biomass and production were estimated for the six dominant taxa. Abundance declined fromca. 4150 at 2.5 m depth toca. 1400 ind m^–2 at 16 m depth and averagedca. 3200 ind m^–2 on a lakewide basis. By numbers,Heterotrissocladius changi andH. oliveri dominated the average fauna.H. changi was common at the 2.5 m and 5 m depth stations, whereasH. oliveri dominated from 5 m depth downwards. Chironomids showed mainly a 1-yr life cycle, but apparently bothHeterotrissocladius species had two contemporary cohorts with emergence in midsummer and late autumn/early spring, respectively. Average annual ratio was 4.2 and 4.6 forH. oliveri andH. changi, respectively. Annual production varied from 0.3 g ash-free dry weight (AFDW) m^–2 y^–1 at 16 m depth to 1.6 g AFDW m^–2 y^–1 at 2.5 m depthH. changi contributed 45%, fiveMicropsectra spp. 17% andH. oliveri 15% to total average production, which on a lakewide basis wasca. 1.1 g AFDW or 25 kJ m^–2 y^–1. Lake 95 thus belongs at the very low end of measured lake zoobenthic productions, which range from 10 kJ m^–2 y^–1 in Arctic lakes toca. 1600 kJ m^–2 y^–1 in highly eutrophic shallow lakes.     

The Life History and Occurrence of Sialis lutaria L. (Megaloptera) in the Littoral Zone of Lake Esrom,Denmark

The life history and distribution of Sialis lutaria L. in the exposed littoral zone of Lake Esrom, Denmark were examined during 1979/80. Sialis showed an annual life cycle with mean annual abundance of 20 and 40 ind m⁻² at 0.4 and 2.0 m depth, respectively. Growth peaked during July with rates of about 10 % d⁻¹. Net annual production averaged 0.2 g DW at 0.4 m depth and 0.8 g DW at 2.0 m depth. The annual P/B-ratios were estimated in the range of 4.6-7.1.     

Comparative bioenergetics of permanent and temporary pond populations of the freshwater clam,Musculium partumeium (Say)

The population energetics of a temporary and a permanent pond population of Musculium partumeium in Southwest Ohio were studied. In the permanent pond (surface area = 396 m², maximum depth = 0.7 m) the population was bivoltine and iteroparous whereas in the temporary pond (surface area = 1042 m², maximum depth = 0.9 m) the population was usually univoltine and semelparous.Growth and biomass were assessed as total organic carbon and total nitrogen to provide estimates of productivity and seasonal changes in C:N for each generation. Productivity (non-respired assimilation = growth + reproduction; N-R.A. = G + R) was 6939 mgC·m^–2·a^–1 (3858 and 3353 mgC·m^–2·a^–1 for each generation) and 1661 mgC·m^–2·a^–1 for the permanent and temporary pond populations respectively. The average standing crop biomass (B) was 606.8 mgC·m^–2 (357.5 and 249.3 mgC·m^–2 for each generation) and 231.9 mgC·m^–2 with overall productivity: biomass ratios of 11.4 and 7.2 for the permanent pond and temporary pond populations respectively.Respiration rates were converted to carbon equivalents (respired assimilation = R.A.) and used to evaluate the components of total assimilation (T.A. = R.A. + N-R.A.) and the efficiency of partitioning this energy to N-R.A. for G and R. When expressed as a percentage, the production efficiencies (100 × N-R.A.:T.A.) were 50.4 and 62%, and the reproductive efficiencies (100 × R:N-R.A.) were 26.4 and 18% for the permanent and temporary pond populations respectively. The reproductive efficiencies for populations of these viviparous clams are greater than those for most oviparous molluscs.The comparative information on the energetics of these populations does not completely fit any theoretical consideration of reproductive effort or life-history strategy. These data are discussed in relation to selection for population success in temporary ponds.Funded in part by grants to Albert J. Burky from the Ohio Biological Survey and the University of Dayton Research CouncilFunded in part by grants to Albert J. Burky from the Ohio Biological Survey and the University of Dayton Research Council     

Laboratory investigation on community composition, emergence patterns and biomass of wood-inhabiting Chironomidae (Diptera) from a sandy lowland stream in Central Europe (Germany)

Wood-inhabiting chironomid communities were investigated from June 1998 to July 1999 by laboratory rearing of randomly collected submerged branches from a headwater and mid-reach site of a sandy lowland stream, separated by an impoundment used for fishery purposes. Total annual emergence (males and females) from headwater samples was higher (2551 ind m^–2 y^–1) compared to the mid-reach (1576 ind m^–2 y^–1), which could be due to disturbances caused by frequent impoundment openings resulting in high discharge events. Chironomid community from branches comprised three subfamilies, with Orthocladiinae (18 species, 2189 ind m^–2 y^–1) clearly predominating at the headwater (total of 36 species). Mid-reach samples (total of 48 species) showed similar emergent numbers of Orthocladiinae (19 species, 786 ind m^–2 y^–1) and Chironominae (26 species, 764 ind m^–2 y^–1). Tanypodinae were caught very rarely in the laboratory emergence (12 ind m^–2 y^–1 at both sites). Shannon-Wiener diversity index for the mid-reach chironomid community was higher (2.52) than for the headwater community (1.68). Chironomid species composition on woody debris was similar between stream sites, with a Sørensen index of 0.75, but showed different dominance structures indicated by a Wainstein index of 0.26. Total adult biomass (in the manner of dry mass) during the study period was higher for wood-dwelling chironomids from the headwater (158.2 mg m^–2 y^–1) compared to the mid-reach (123.8 mg m^–2 y^–1), but individual biomass was higher for mid-reach chironomids (0.079 mg ind^–1 vs. 0.062 mg ind^–1 at the headwater), indicating the predominance of larger species. Total biomass of wood-inhabiting chironomids in the investigated lowland stream was low compared to other field emergence studies, which could be attributed to the laboratory approach for investigating the emergence from a single substrate type (submerged wood) instead of the integrative field surveys where chironomids from all habitats were caught. Main reason for the lack of chironomid species closely associated to wood in this sandy lowland stream could be infrequent but episodic disurbances caused by the anthropogenic induced highly fluctuating discharge regime of the downstream study site.     

Crustacean assemblage and environmental characteristics of a man-made solar saltwork in southern France,with emphasis on anostracan (Branchiopoda) population dynamics

Physical and chemical variables, anostracan populations (Artemia parthenogenetica and Branchinella spinosa) and other biota were studied during 1996–1997 in a Camargue saltern (max. depth 1 m). The taxonomic composition and density of macroinvertebrates were investigated twice monthly, based on benthic substrate and water column samples. Fauna was composed of three groups in terms of numerical importance. The benthic macroinvertebrates were represented only by nematodes (< 50 ind. m^–2 to > 500 ind. m^–2 in November–December and May respectively). The zooplankton was dominated by crustaceans, one cladoceran, Moina salina (ranging from 670 to 2350 ind. m^–2 in spring), two anostracans, Artemia parthenogenetica (< 50 ind. m^–2 in autumn), and Branchinella spinosa (max. 190 ind. m^–2 in December to absent in April), and two copepods, Cletocamptus retrogressus (max. density 2000 ind. m^–2 in November), and Eurytemora velox (max. density 650 ind. m^–2 in February–March). Insects (Chironomidae, Culicidae) were rare, with mean densities < 1 ind. m^–2. The phenology of each crustacean population is discussed in relation to physical and chemical water variables. Salinity appeared to be of greatest importance regulating the population abundance.     

Energy metabolism of Chironomus anthracinus (Diptera: Chironomidae) from the profundal zone of Lake Esrom,Denmark, as a function of body size,temperature and oxygen concentration

The profundal zone of Lake Esrom, Denmark has a dense population of Chironomus anthracinus, which survives 2–4 months of oxygen depletion each summer during stratification. The metabolism of 3rd and 4th instar larvae was examined in regard to variation in biomass and temperature. Respiration at air saturation was described by a curvilinear multiple regression relating oxygen consumption to individual AFDW and temperature. At 10 °C and varying oxygen regimes the O₂ consumption and CO₂ production of 4th instar larvae were almost unaltered from saturation to about 3 mg O₂ l^–1, but decreased steeply below this level. The respiratory quotient increased from 0.82 at saturation to about 3.4 at oxygen concentrations near 0.5 mg O₂ l^–1. This implied a shift from aerobic to partially anaerobic metabolism. At 0.5 mg O₂ l^–1 the total energy production equalled 20% of the rate at saturation of which more than one third was accounted for by anaerobic degradation of glycogen. This corresponded to a daily loss of 12 µg mg AFDW^–1 or approximately 5% of the body reserves. At unchanged metabolic rate the glycogen store would last three weeks, but long term oxygen deficiency causes a further suppression of the energy metabolism in C. anthracinus.     

Life cycle and growth ofTrichostegia minor (Curtis) in temporary woodland pools (Trichoptera: Phryganeidae)

Larvae of the caddisTrichostegia minor (Curtis) were collected from four woodland pools in The Netherlands, three of which are temporary, from August 1986 till June 1987. Eggs and larvae of this species proved to be very well adapted to drought, freezing, strongly fluctuating pH and alkalinity levels and prolonged oxygen deficit. The life cycle ofT.minor in a small woodland marsh overgrown byCalla palustris took one year. Adult flight period started at the end of May. Oviposition took place independent of water. Hatching of the eggs started in September and was probably induced by immersion. During the larval stage from September until May, 5 instars could be distinguished by the size of the head capsule. Growth of instars I, II and III during autumn was moderate. Most larvae overwintered as instar III or IV. Possibly there was a larval diapause during winter. In spring rapid growth to instar V took place prior to pupation. Growth rate, expressed as the increase of mean individual dry weight was highest from March to April (2.05±0.75% DW.m^–2.d^–1). In extremely shallow water growth in spring was initially more rapid compared to growth in deeper water. During winter the growth rate decreased to 0.038±0.071% DW.m^–2.d^–1. Net annual production based on the changes of momentary biomass was 183.2±31.7 mg DW.m^–2.y^–1 or 177.2±31.3 mg AFDW.m^–2.y^–1. Production loss during the winter season was 75.1±10.8 mg DW.m^–2.y^–1 or 72.3±10.6 mg AFDW.m^–2.y^–1.     

Crustacean plankton in Høylandet

Crustacean plankton was studied in 12 lakes in theHøylandet area in 1986–87. Basic lake characteristicsare elevation 134–415 m, surface area 9–530 ha, pH 5.9–7.1,conductivity (25 °C) 12–40 µS cm^-1 andSecchi depth 4–9 m. Number ofspecies present varied between 3 and 11. Populationnumbers between 4000 and 400 000 per m² andbiomasses were within the range 30 to1800 mg m^-2 dry weight. Cladocera dominated overCopepoda in lakes with allopatric brown trout (Salmo trutta L.), on the contrary to lakes also populatedby Arctic charr (Salvelinus alpinus (L.)). Thesevariations are caused by differences in elevation,lake morphometry, water quality, fish predation andthe general distribution of the species. The largestlakes at lowest elevation were richest in species. Theacid sensitive genus Daphnia was represented by 3species. The lakes Storgrønningen (530 ha) andRøyrtjønna (27 ha) were sampled monthly in theice-free seasons of 1986–89, and Storgrønningen moreintensively from June to November in 1987 and 1988. The same6 species of Cladocera and 5 of Copepoda were presentin both lakes. Their life cycles were traditional orknown from several other Scandinavian lakes. Meanseasonal biomasses were of the range600–750 mg m^-2. At the species level, there wereconsiderable variations between years inStorgrønningen and particularly in Røyrtjønna. Noeffects of human impacts on the crustacean planktonwere found. The Høylandet lakes are representative forScandinavian oligotrophic to almost ultra-oligotrophiclakes. Storgrønningen is well qualified as a referencesystem. The between year variations in Røyrtjønna areso extreme, that any human impact could only be traced at alevel causing the extinction of species.     

Transport of coarse particulate organic matter in an Idaho river,USA

I investigated organic matter transport in the Henry's Fork of the Snake River, Idaho, USA, from August 1987 to November 1988. Mean discharge during the study was 15 m³ s^–1. Screens were used to sample very coarse (> 6 mm) transported aquatic macrophyte material (VCTMM). Drift nets were used to sample coarse (1–6 mm) and fine (0.25–1 mm) transported particulate organic matter (CTOM and FTOM). Mean monthly concentration of VCTMM was 0.064 mg AFDWl^–1 and was significantly higher than CTOM (0.024 mg AFDW l^–1) and FTOM (0.036 mg AFDW l^–1). VCTMM concentration was highest in December (0.163 mg AFDW l^–1) and lowest in May (0.018 mg AFDW l^–1). The sample position along a transect across the channel had a significant effect on the amount of transported organic matter collected in many months. The concentration of debris from individual species tracked the standing stock of that species during the growing season. In Fall, a dramatic increase in VCTMM corresponded to a decrease in macrophyte standing stock. FTOM and CTOM concentrations were highest in January (CTOM: 0.048; FTOM: 0.111 mg AFDW l^–1), lowest in November 1988 (<0.006 mg AFDW l^–1), were not correlated with discharge, and were inversely correlated with the standing stocks of macrophytes upstream, probably because macrophyte beds influenced the retentiveness of the channel. Standing stock of aquatic macrophytes was highest in September–October (5.2 kg wet weight m ^–2) and lowest in February (1.7 kg wet weight m^–2). Annual movement of particulate organic matter past the sampling point was about 45 000 kg AFDW, of which 21 000 kg was VCTMM, 8 000 kg was CPOM, and 16 000 kg was FPOM.     

On the distribution patterns and population dynamics of sublittoral and profundal oligochaeta fauna from Lake Banyoles (Catalonia,NE Spain)

Oligochaeta assemblages from Lake Banyoles were composed of one species of Naididae and eight species of Tubificidae, Potamothrix heuscheri (8–1350 ind. m^–2) being the most abundant and widely distributed species, followed by Limnodrilus hoffmeisteri (25–858 ind. m^–2) and Psammoryctides barbatus (12–792 ind. m^–2). The remaining species were Potamothrix hammoniensis, Potamothrix bavaricus, Limnodrilus claparedeanus, Branchiura sowerbyi, Aulodrilus pigueti and Dero digitata, all of which occurred at lower densities and with restricted distributions.Regardless of sampling depth, maximum densities of oligochaetes were found in winter (up to 5,142 ind. m^–2 in December) and secondarily in June. Immature Tubificidae with hair setae and mature P. heuscheri were present and dominant throughout the year. Cestode parasites were recorded infesting L. hoffmeisteri and P. heuscheri populations.Oligochaeta densities increased with depth, but low oxygen concentrations (less than 1 mg l^–1) during a long period (4 months) in some basins of the lake acted as a key factor in reducing the density of worms and the species richness to one species, Potamothrix heuscheri.     

Seasonal growth of the submerged macrophyte Ceratophyllum demersum L. in mesotrophic Lake Vechten in relation to insolation,temperature and reserve carbohydrates

Seasonal growth of a Ceratophyllum demersum stand in Lake Vechten, The Netherlands, varied considerably during 1978, '79 and '81. The biomass was highest in 1981 (max. 115 g ash-free dry weight m^–2). The 1978 growth trend was similar to that of 1981 with slightly lower net annual production (max. 92 g AFDW m^–2). The vegetation almost disappeared in 1979 (max. 24 g AFDW m^–2). These large differences in biomass could not be explained by differences in insolation but were mainly attributed to the severe winter of 1978–'79 which slowed plant development and full maturity was attained only after the diurnal insolation had decreased.Reserve carbohydrates were primarily starch and water-soluble sugars. Starch concentrations varied between 71.5 and 24.7 mg g^–1 AFDW in the winter and summer, and total sugar concentrations between 24.9 and 20.2 mg g^–1 AFDW, respectively. The non-reducing sugars tended to predominate in summer and the reducing sugars in winter. The value of these reserves for maintenance, growth and survival was assessed by calculating plant carbon budgets for several seasons.It was concluded that C. demersum relies heavily on its developmental timing and energy-conserving physiological mechanisms to survive extended periods of stress and in these respects is analogous to a typical terrestrial shade plant.     

Living conditions, abundance and biomass of under-ice fauna in the Storfjord area (western Barents Sea, Arctic) in late winter (March 2003)

The under-ice habitat and fauna were studied during a typical winter situation at three stations in the western Barents Sea. Dense pack ice (7–10/10) prevailed and ice thickness ranged over <0.1–1.6 m covered by <0.1–0.6 m of snow. Air temperatures ranged between –1.8 and –27.5°C. The ice undersides were level, white and smooth. Temperature and salinity profiles in the under-ice water (0–5 m depth) were not stratified (T=–1.9 to –2.0°C and S=34.2–34.7). Concentrations of inorganic nutrients were high and concentrations of algal pigments were very low (0.02 g chlorophyll a l^–1), indicating the state of biological winter. Contents of particulate organic carbon and nitrogen ranged over 84.2–241.3 and 5.3–16.4 g l^–1, respectively, the C/N ratio over 11.2–15.5 pointing to the dominance of detritus in the under-ice water. Abundances of amphipods at the ice underside were lower than in other seasons: 0–1.8 ind. m^–2 for Apherusa glacialis, 0–0.7 ind. m^–2 for Onisimus spp., and 0–0.8 ind. m^–2 for Gammarus wilkitzkii. A total of 22 metazoan taxa were found in the under-ice water, with copepods as the most diverse and numerous group. Total abundances ranged over 181–2,487 ind. m^–3 (biomass: 70–2,439 g C m^–3), showing lower values than in spring, summer and autumn. The dominant species was the calanoid copepod Pseudocalanus minutus (34–1,485 ind. m^–3), contributing 19–65% to total abundances, followed by copepod nauplii (85–548 ind. m^–3) and the cyclopoid copepod Oithona similis (44–262 ind. m^–3). Sympagic (ice-associated) organisms occurred only rarely in the under-ice water layer.     

Life history, population dynamics and production of eastern mosquitofish, Gambusia holbrooki (Pisces, Poeciliidae), in rice fields of the lower Mondego River Valley, western Portugal

The introduced population of Gambusia holbrooki from the rice fields of the lower Mondego River Valley, Portugal, was studied for 15 months, relating their life cycle and population dynamics with its production, in order to assess the role of the species in the energy flow and secondary production in this type of agro-ecosystem. Two main annual cohorts (1995 and 1996 cohorts) were identified. The females outnumbered males and the average female/male-ratio was 4. The inspection of ovary developmental stages of this viviparous fish, revealed that the most important reproductive period was between April and August. The first recruits were recorded in June and were present thereafter until October. Males from the parental cohort died before August, whereas parental females could survive until October. Mean adjusted fecundity (number of embryos divided by female standard length) peaked in July 1996 (0.95) and in June 1997 (1.05). Females reached greater sizes, had a higher growth rate and lived longer than males. Annual production was estimated at 3.101 g.m^–2.year^–1 (ash-free dry weight, AFDW), the average biomass at 2.896 g.m^–2 (AFDW), and the P/B ratio was 1.071. A conjugation of life history, population dynamics, production and ecological traits (e.g. fast growth, reduced longevity, viviparity, high productivity, an intermediate position in food chain, and no special habitat requirements for reproduction) clearly show that the populations of G. holbrooki, introduced into rice fields all over the world, may play an important role in the structure and functioning of the biological communities of these important agro-ecosystems.     

Phosphorus release from resuspended sediment in the shallow and wind-exposed Lake Arresø, Denmark

Wind-induced sediment resuspension occurs frequently in the shallow and eutrophic Lake Arresø, Denmark. The impact of resuspension on internal phosphorus loading was investigated by laboratory experiments studying P-release from the undisturbed sediment surface and by experiments simulating resuspension events.Phosphorus release from undisturbed sediment sampled in May and August was 12 mg and 4 mg m^–2 d^–1, respectively. During experimental simulation of resuspension, soluble reactive phosphate (SRP) increased by 20–80 µg l^–1, which indicates that a typical resuspension event in the lake would be accompanied by the release of 150 mg SRP m^–2. The internal P loading induced by resuspension is estimated to be 60–70 mg m^–2 d^–1, or 20–30 times greater than the release from undisturbed sediment.SRP release during simulation of resuspension was mainly dependent on the equilibrium conditions in the water column and was basically independent of the increase in suspended solids and the duration of resuspension. A second simulation of resuspension conducted 26 hours later, did not result in any further release of SRP from sediment sampled in May. In contrast, there was an additional SRP release from sediment sampled in August, indicating that an exchangable P pool, capable of altering equilibrium conditions, is built up between resuspension events.It is concluded that resuspension, by increasing the P flux between sediment and water, plays a major role in the maintenance of the high nutrient level in Lake Arresø. A relatively high release rate is maintained during resuspension because of the low Fe:P ratio and the high concentration of NH₄Cl-extractable P in the sediment.     

Impact of long-term alternations of discharge and spate on the chironomid community in the lowland Widawka River (Central Poland)

Spatial and temporal distribution, abundance and production of macroinvertebrate communities were estimated over two years in a fifth-order section of the Widawka River. Discharge of this river has been increased artificially by coal mine water inputs. Additionally, during the second year, one of the highest discharges of the current 20-year period was recorded. Chironomidae were co-dominant in macrobenthos, both in a straight reach (WIA) and in a meandering site (WIB). More mosaic habitats resulted in higher densities of midges, reaching 6215 ind.m^–2 in year 1 and 1141 ind.m^–2 in year 2 at WIA, while at WIB 896 densities were ind.m^–2 and 257 ind.m^–2, respectively. Flooding affected the distribution and abundance of the chironomid assemblages. Recolonization by psammophilous Polypedilum began after the various microhabitats were buried with sand. Chironomid production was estimated on a species-specific basis for the dominant taxa. In year 1 (mean annual water temperature 10.0° C) chironomid production was 12.4 g dry wt m^–2 yr^–1 1 at WIA and 1.9 g dry wt m^–2 yr^–1 at WIB. These values sharply decreased in year 2 (mean annual water temperature 9.8° C) reaching 1.9 g dry wt m^–2 yr^–1 at WIA and 0.4 g dry wt m^–2 yr^–1 at WIB, as effects of the high spate.     

Modelling the recovery of hypertrophic L. Glumsø (Denmark)

Diversion of sewage from L. Glumsø reduced phosphorus loading from 6.0 g P.m^–2.yr^–1 to 1.6 g P.m^–2.yr^–1. Chlorophyll levels during summer were reduced from 6–800 mg Chl.m^–3 to about 200 mg Chl.m^–3 mainly by extended periods with phosphorus limitation. Internal phosphorus loading was significant in the first 2 years after nutrient reduction. Predictions of the recovery were made by both simple, empirical models and with complex, dynamic model versions. The actual responses of L. Glumsø were compared with both previously published predictions and predictions made with improved model versions. Objective functions of 0.18 and global correlation coefficients of 0.89 could be achieved.     

The drift of zooplankton and microzoobenthos in the river Strandaelva,western Norway

The drift of zooplankton (rotifers, cladocerans, cyclopoid copepods) and microscopical zoobenthos (mainly bdelloid rotifers and small chironomid larvae) was investigated by filtering samples of river water. The number of drifting benthic rotifers varied between 1 000 and 6 000 ind. m^–3 in the lake inlet, and between 30 and 500 ind. m^–3 in the lake outlet, without any seasonal trend. The number of drifting insect larvae was approx. equal in the lake inlet and outlet, with a maximum in summer (250–300 ind. m^–3) and minimum in winter (ca. 10 ind. m^–3). Increasing water flow resulted in an increasing number of drifting zoobenthos. Downstream from the lake, the number of drifting benthic rotifers was increasing from approx. 300 ind. m^–3 in the outlet to 6 500 ind. m^–3 3.4 km downstream, while the number of insect larvae was ca. 100 ind. m^–3 in the outlet and leveled off at approx. 300 ind. m^–3 after 200 m. The number of drifting zooplankton in the lake outlet varied between 20 and 2 000 ind. m^–3 for crustaceans, and between 300 and 20 000 ind. m^–3 for rotifers, both with a maximum in late summer/autumn and a minimum in winter. The number of drifting zooplankton decreased by some 45% in the first 200 m from the lake outlet, but some zooplankton was still found in the drift 3.4 km downstream. The largest species was removed first from the drift. The diurnal variation in the number of drifting zooplankton in lake outlets appear to be related to the vertical migration in the lake, i.e. the largest number drifting when most animals are in the upper water layers.Contribution from the Voss Project, University of OsloContribution from the Voss Project, University of Oslo     

Emergence patterns of univoltine and bivoltine Ips typographus (L.) populations and associated natural enemies

Control measures aiming at reducing bark beetle populations and preserving their natural enemies require a sound knowledge on their overwintering and emergence behaviour. These behavioural traits were investigated in univoltine and bivoltine populations of the European spruce bark beetle (Ips typographus [L.], Coleoptera: Scolytinae) and its predators and parasitoids over several consecutive years. In univoltine populations, roughly 50% of the bark beetles left their brood trees in fall together with most parasitoids and some significant predatory flies and beetles. In bivoltine populations, <10% of the second bark beetle generation emerged in fall and the remainder overwintered under the bark of their brood trees. Likewise, most predatory beetles and flies spent wintertime with their prey under the bark, while most parasitic wasps emerged in fall. The spring emergence of bivoltine predatory beetles was found to occur up to 3 weeks earlier than that of I. typographus, while that of the predatory flies and the parasitoids was delayed by up to 1 month. In univoltine populations, the bark beetles emerged several weeks prior to most antagonistic taxa. In the heat year 2003, three I. typographus generations were produced at the lower location, 36% of the third generation emerged in fall, while the proportions of overwintering predators remained largely the same as in previous years. Similar to their host, more parasitoids left their brood trees in fall after warm years. The results show that sanitation felling during winter probably kills most bark beetles in bivoltine populations, but also eliminates many natural enemies. In univoltine populations, sanitation felling might be less detrimental to both I. typographus and natural enemies because a fair fraction of their populations will already have left the trees before cutting. Warmer climates may affect the interactions of bark beetles and natural enemies and thus the impact of control measures.     

Eight years of internal phosphorus loading and changes in the sediment phosphorus profile of Lake Søbygaard,Denmark

During each of the first 8 years following an 80–90% reduction in external phosphorus loading of shallow, hypertrophic Lake Søbygaard, Denmark in 1982, phosphorus retention was found to be negative. Phosphorus release mainly occurred from April to October, net retention being close to zero during winter. Net internal phosphorus loading was 8 g P m^–2 y^–1 in 1983 and slowly decreased to 2 g P m^–2 y^–1 in 1990, mainly because of decreasing sediment phosphorus release during late summer and autumn. The high net release of phosphorus from Lake Søbygaard sediment is attributable to a very high phosphorus concentration and to a high transport rate in the sediment caused by bioturbation and gas ebullition. Sediment phosphorus concentration mainly decreased at a depth of 5 to 20 cm, involving sediment layers down to 23 cm. Maximum sediment phosphorus concentration, which was 11.3 mg P g^–1 dw at a depth of 14–16 cm in 1985, decreased to 8.6 mg P g^–1 dw at a depth of 16–18 cm in 1991. Phosphorus fractionation revealed that phosphorus release was accompanied by a decrease in NH₄Cl-P + NaOH-P and organic phosphorus fractions. HCl-P increased at all sediment depths. The Fe:P ratio in the superficial layer stabilized at approximately 10. Net phosphorus release can be expected to continue for another decade at the present release rate, before an Fe:P ratio of 10 will be reached in the sediment layers from which phosphorus is now being released.