Spatial and temporal variation of microbial respiration rates in a blackwater stream

1. The extent of spatial and temporal variation of microbial respiration was determined in a first-order, sand-bottomed, blackwater stream on the coastal plain of south-eastern Virginia, U.S.A.
2. Annual mean respiration rates (as g O₂ m^–3 h^–1) differed significantly among substrata: leaf litter, 12.9; woody debris, 2.4; surface sediment, 0.8; hyporheic sediment, 0.4; water column, 0.003. Rates associated with wood were higher than those with leaves when expressed per unit surface area.
3. Highest respiration rates on leaves, wood and in the water column occurred during the summer, whereas rates in the sediments were greatest during the late autumn and winter. Water temperature, as well as particulate organic matter and nitrogen content of the substrata, was correlated positively with respiration rates.
4. A stepwise multiple regression showed that temperature and nitrogen content together explained 88% of the variation in respiration rates of leaves and wood. In contrast, particulate organic matter content and nitrogen content explained 89–90% of the variation in respiration in the sediments. Although water temperature was a significant factor in the sediment multiple regressions, its addition as an independent variable improved the regression models only slightly.
5. Annual mean respiration in the stream channel, based on the proportional amount of respiration occurring associated with each type of substratum during each month, was 1.1 kg O₂ m^–2 yr^–1. Seventy per cent of respiration in the stream occurred in the hyporheic zone, 8–13% occurred in the surface sediment, leaf litter or woody debris, and < 1% occurred in the water column. Approximately 16% of total detritus, or 40% of non-woody detritus, stored in the stream during the year was lost to microbial respiration.     

Spatial and temporal variation of microbial respiration rates in a blackwater stream

1. The extent of spatial and temporal variation of microbial respiration was determined in a first-order, sand-bottomed, blackwater stream on the coastal plain of south-eastern Virginia, U.S.A.
2. Annual mean respiration rates (as g O₂ m^–3 h^–1) differed significantly among substrata: leaf litter, 12.9; woody debris, 2.4; surface sediment, 0.8; hyporheic sediment, 0.4; water column, 0.003. Rates associated with wood were higher than those with leaves when expressed per unit surface area.
3. Highest respiration rates on leaves, wood and in the water column occurred during the summer, whereas rates in the sediments were greatest during the late autumn and winter. Water temperature, as well as particulate organic matter and nitrogen content of the substrata, was correlated positively with respiration rates.
4. A stepwise multiple regression showed that temperature and nitrogen content together explained 88% of the variation in respiration rates of leaves and wood. In contrast, particulate organic matter content and nitrogen content explained 89–90% of the variation in respiration in the sediments. Although water temperature was a significant factor in the sediment multiple regressions, its addition as an independent variable improved the regression models only slightly.
5. Annual mean respiration in the stream channel, based on the proportional amount of respiration occurring associated with each type of substratum during each month, was 1.1 kg O₂ m^–2 yr^–1. Seventy per cent of respiration in the stream occurred in the hyporheic zone, 8–13% occurred in the surface sediment, leaf litter or woody debris, and < 1% occurred in the water column. Approximately 16% of total detritus, or 40% of non-woody detritus, stored in the stream during the year was lost to microbial respiration.     

The responses of a lotic mayfly Nousia sp. (Ephemeroptera: Leptophlebiidae) to moving water and light of different wavelengths

1. Although laboratory studies of the behaviour of aquatic macroinvertebrates are common, there has been little critical evaluation of the importance of test conditions to them. We used a common Australian leptophlebiid mayfly, Nousia sp., to investigate responses to light, wavelength of light, presence or absence of cover and still or flowing water.
2. Nousia sp. showed substantial qualitative differences in behaviour, as measured by movement, when there was no refuge (in the form of a crevice beneath a tile) present in the experimental arena.
3. We found no evidence of diel periodicity in activity in Nousia sp.
4. Nousia sp. did not respond to infra-red, red or green light at a flux density of 18–19 μmol m^–2 s^–1.
5. Nymphs were three times more likely to remain stationary in flowing water (mean velocity 0.10 m s^–1) than in still water.
6. We conclude that generalized assumptions about test conditions for experiments designed to quantify laboratory behaviour in benthic macroinvertebrates are unjustified and that evaluation of the individual requirements of test species should be conducted routinely.     

Micro-environmental characteristics of filamentous algal communities in flowing freshwaters

SUMMARY. 1. Filamentous algae in flowing freshwaters can represent a spatially and temporally distinct sub-habitat for epiphytic diatom communities. This sub-habitat is described in a low discharge, spring-fed stream with extensive filamentous green algal mats, and in a tuft of Cladophora glomerata (L.) Kützing from a large river.
2. Oxygen micro-electrodes, a thermistor current velocity probe, a standard pH probe and water chemistry were employed to assess spatial heterogeneity. Temporal patterns of epiphyte colonization were evaluated on filamentous artificial substrates.
3. There were steep spatial gradients in the low discharge stream. At mid-day, O₂ ranged from 0–1.5 times air-saturated O₂ concentrations, pH varied from 7.25–8.0, and current velocity spanned 0–0.5 m s⁻¹. Areas near the surface of algal mats had high O₂, pH and current velocity. These patterns were correlated with epiphyte community structure.
4. In the interior of C. glomerata tufts O₂ concentration was raised and current velocity depressed compared to the surrounding water, even when external current velocity was as high as 0.4 m s⁻¹.
5. After thirty-five days of colonization of artificial substrate in the low discharge environment, epiphyte communities were similar to those on filamentous atgae. Epiphyte diversity on artificial substrates subsequently decreased compared to natural substrates as did the similarity between the types on substrates, suggesting that microscale renewal of epiphyte habitat (growth of filamentous algae) maintains high epiphytic diversity.     

Sediment microbial respiration in a synoptic survey of mid‐Atlantic region streams

1. The rate of microbial respiration on fine‐grained stream sediments was measured at 196 first‐ to third‐order sites in the mid‐Atlantic region of the United States.
2. Sample collection took place between April and July in 1993, 1994 and 1995.
3. Study streams were randomly selected from the United States Environmental Protection Agency's (USEPA) River Reach File (RF3) using the sample design developed by USEPAs Environmental Monitoring and Assessment Programme (EMAP).
4. Respiration rate ranged from 0 to 0.622 g O ₂ g^–1 ash‐free dry mass (AFDM) h^–1, and was significantly correlated with stream temperature, water chemistry and physical habitat.
5. Respiration rate was significantly higher in streams from the Chesapeake drainage area compared to those from the Ohio drainage area, and in streams from the Northern Appalachian Plateau and Uplands and Central Appalachian Valleys ecoregions compared with streams from the Blue Ridge and Central Appalachian Plateau ecoregions.
6. Respiration was not significantly different among stream orders or between years.     

Denitrification in river sediments: relationship between process rate and properties of water and sediment

1. A survey was made of denitrification and nitrous oxide (N₂O) production in river sediments at fifty sites in north‐east England during one season in order to investigate the relationship between rates and environmental factors likely to influence these processes. The sites were chosen to represent a wide range of physical and chemical conditions. Denitrification rate and N₂O production were measured within 5 h of sampling using the slurry acetylene blockage technique.
2. Denitrification rate ranged from less than 0.005–260 nmol N g^–1 DW h^–1, tending to increase in a downstream direction. N₂O production ranged from negative values (net consumption) to 13 nmol N₂O‐N g^–1 DW h^–1 and accounted for 0–115% of the N gases produced.
3. Denitrification rate and N₂O concentration in the sediment were correlated positively with nitrate concentration in the water column, water content of the sediment and percentage of fine (< 100 μm) particles in the sediment.
4. The variation in denitrification rate was satisfactorily explained (64% total variance) by a model employing measurements of water nitrate and water content of sediments. No simple or multiple relationship was found for N₂O production.     

Positive anemotaxis by Varroa mites: responses to bee odour plumes and single clean‐air puffs

Abstract.The stimuli and mechanisms mediating host location and host choice by the bee mite, Varroa jacobsoni (Oudemans), are currently unknown. It is shown that Varroa can use single clean‐air puffs and bee‐odour plumes in a wind tunnel as directional cues. Varroa turned nearly straight upwind in response to single 0.1‐s puffs of clean air directed at 90° to the their anterior‐posterior axis. They turned significantly further to their left side (104°) than to their right (76°), but showed no difference in latency to initiation of the turns (means of 63.3 ms vs. 62.6 ms, respectively). They also followed bee‐odour plumes in a wind tunnel. When released in odour and control plumes mid‐way between the plume's origin and the downwind end of the tunnel, mites responding to bee‐odour walked upwind in, or along the edge of, the odour plume with 38% making contact with the odour delivery tube; mites in clean air did not walk upwind along the air stream, and none made contact with the air delivery tube. Walking speeds were not different between the bee‐odour and control groups (0.28 vs. 0.29 cm s^–1); there were also no differences in the turning rates (96.85 vs. 97.16 deg s^–1 and 388.08 vs. 379.18 deg cm^–1, respectively). Under all conditions, mites walked in a zigzag fashion.     

Vertical distribution and abundance of invertebrates within the sandy substrate of a low-gradient headwater stream

SUMMARY. 1 The vertical distribution of invertebrates (>0.053 mm) was studied in a sandy-bottomed, first-order stream on the Coastal Plain of Virginia, U.S.A. Invertebrate species composition, abundance and bio-mass were determined monthly over one year at sediment depth intervals of 0–1, 1–5, 5–15,15–30 and 30–40 cm.
2. The subsurface community was numerically dominated by species of Chironomidae, Nematoda and Crustacea, while much of the biomass was due to early instars of several species of Trichoptera.
3. Invertebrate density and biomass decreased significantly with depth in the substrate (ANOVA; P <0.05). Annual mean density decreased from 1,346,844 individuals m⁻³ at the surface to 13,578 individuals m⁻³ at 15–30 cm. Annual mean biomass decreased from 66.30 g m⁻³ at the surface to 0.44 g m⁻³ at 15–30 cm.
4. Dissolved oxygen decreased markedly from the surface to the 5 cm depth in the substrate, anaerobic conditions often occurring below 10 cm. Density and biomass both showed a significant positive relationship with dissolved oxygen concentration (Linear regression; P <0.05).
5. Physical forces were important in structuring the subsurface invertebrate community. Besides low dissolved oxygen concentration, sediment scouring resulting from storm discharge dramatically reduced density and biomass     

Measurements of phosphorus uptake by macrophytes and epiphytes from the LaPlatte River (VT) using 32P in stream microcosms

1. Phosphorus (P) uptake by macrophytes and epiphytes from the LaPlatte River (VT) was examined in the laboratory by adding ³²PO₄‐P to recirculating stream microcosms.
2. Water, plugs of sediment and plants were removed from the river and placed into the microcosms. ³²PO₄‐P was then added either to the water or the sediment, and its incorporation into plants and epiphytes was monitored over 3 days. Uptake was examined at both ambient (5 μg L^–1) and increased (50 μg L^–1) soluble reactive phosphorus (SRP) concentrations. A computer program was developed to fit curves to the radiotracer data and calculate rate constants for the simultaneous transfer of ³²P among compartments.
3. Both macrophytes and epiphytes removed P from the water, but epiphyte uptake of P was more rapid. Phosphate enrichment stimulated P uptake by both macrophytes and epiphytes. Macrophytes also obtained P from the sediment. The relative contribution of P to macrophytes from the water vs. that from the sediment appeared to vary with SRP in the overlying water. Accurate estimates of rates of P uptake from sediments by macrophytes were difficult to obtain however, due to very low and highly variable unit rate constants for P uptake and uncertainty about the magnitude of the phosphate pool available for uptake.
4. SRP concentrations were greater in the overlying water than in the sediment pore water of stream microcosms in the present study. Numerous reports in the literature have suggested that this condition favours uptake by macrophyte stems and leaves rather than by roots.
5. Phosphate uptake from the water by macrophytes in shallow streams may be more common than for macrophytes in lakes.     

In situ responses to elevated CO2 in tropical forest understorey plants

1. Plants growing in deep shade and high temperature, such as in the understorey of humid tropical forests, have been predicted to be particularly sensitive to rising atmospheric CO₂. We tested this hypothesis in five species whose microhabitat quantum flux density (QFD) was documented as a covariable. After 7 (tree seedlings of Tachigalia versicolor and Beilschmiedia pendula ) and 18 months (shrubs Piper cordulatum and Psychotria limonensis, and grass Pharus latifolius ) of elevated CO₂ treatment ( c. 700 μl litre^–1) under mean QFD of less than 11 μmol m^–2 s^–1, all species produced more biomass (25–76%) under elevated CO₂.
2. Total plant biomass tended to increase with microhabitat QFD (daytime means varying from 5 to 11μmol m^–2 s^–1) but the relative stimulation by elevated CO₂ was higher at low QFD except in Pharus .
3. Non-structural carbohydrate concentrations in leaves increased significantly in Pharus (+ 27%) and Tachigalia (+ 40%).
4. The data support the hypothesis that tropical plants growing near the photosynthetic light compensation point are responsive to elevated CO₂. An improved plant carbon balance in deep shade is likely to influence understorey plant recruitment and competition as atmospheric CO₂ continues to rise.     

Water temperature in a stream gravel bed and implications for salmonid incubation

SUMMARY. 1. Water temperatures were recorded at hourly intervals in the gravel of a trout spawning area in a small stony stream at depths of 0–20 cm in 1985–86 and 0– 40 cm in 1987.
2. As depth within gravel increased, the size of the daily fluctuations reduced and their time of occurrence was delayed by about 12 min cm⁻¹' in 1985–86 and about 6 min cm⁻¹ in 1987.
3. From October to February mean temperatures at 20 cm depth were, on average, 0.5°C higher than those at the gravel surface. This reflected elevated daily minima more than it reflected elevated daily maxima.
4. From March to July daily minima were lower and daily maxima were higher in the stream than in the gravel. Consequences were: (a) an appreciable increase in mean daily range at all depths in the gravel during the summer, (b) higher daily means (by an average of about 0.4°C) in the water than in the gravel in May to August.
5. Some implications for the early development of salmonid fish are considered.     

A continuous-flow system for measuring in vitro oxygen and nitrogen metabolism in separated stream communities

SUMMARY. 1. This paper describes a continuous-flow system that we have used extensively for measurement of oxygen exchange and nitrogen transformations in different communities of lowland streams, i.e. the water, the sediment, and the macrophyte—biofilm community. The system, which is set up on the stream bank, consists of several macrophyte and sediment chambers equipped with a double-flow system that ensures an internal water velocity close to that in the stream and which, by continuously renewing the water, mimics diel fluctuation in stream temperature and water chemistry. Water temperature and dissolved oxygen are measured continuously and the data stored on magnetic tape. Water samples are collected in a refrigerator and analysed using standard chemical procedures.
2. The application of the system is illustrated using results obtained during a 2-day summer experiment in a shallow macrophyte—rich stream. The biological processes in the stream were mainly associated with the macrophyte—biofilm community and the sediment, those in the water being negligible. Oxygen release was confined to the macrophyte—biofilm community, the sediment consuming oxygen both by day and by night. Whole-system gross production and dark respiration occurred at similar rates (6–7g O₂ m⁻² day⁻¹), net balance being about zero. Inorganic nitrogen was consumed both by the sediment and to a greater extent by the macrophytes, the diel average consumption being 1g N m⁻² day⁻¹.
3. The sum of the activity in the macrophyte and sediment chambers corresponded to the overall activity of the stream section as determined by upstream/downstream mass balance. This indicates that the results obtained with the continuous-flow chambers realistically describe the oxygen and the nitrogen metabolism of the stream.     

Endurance swimming of European eel

A long‐term swim trial was performed with five female silver eels Anguilla anguilla of 0·8–1·0 kg ( c . 80 cm total length, L _T) swimming at 0·5 body lengths (BL) s⁻¹, corresponding to the mean swimming speed during spawning migration. The design of the Blazka‐type swim tunnel was significantly improved, and for the first time the flow pattern of a swim tunnel for fish was evaluated with the Laser‐Doppler method. The velocity profile over three different cross‐sections was determined. It was observed that 80% of the water velocity drop‐off occurred over a boundary layer of 20 mm. Therefore, swim velocity errors were negligible as the eels always swam outside this layer. The fish were able to swim continuously day and night during a period of 3 months in the swim tunnel through which fresh water at 19° C was passed. The oxygen consumption rates remained stable at 36·9 ± 2·9 mg O₂ kg⁻¹ h⁻¹ over the 3 months swimming period for all tested eels. The mean cost of transportation was 28·2 mg O₂ kg⁻¹ km⁻¹. From the total energy consumption the calculated decline in fat content was 30%. When extrapolating to 6000 km this would have been 60%, leaving only 40% of the total energy reserves for reproduction after arriving at the spawning site. Therefore low cost of transport combined with high fat content are crucial for the capacity of the eel to cross the Atlantic Ocean and reproduce.     

The distances travelled by downstream-moving trout fry, Salmo trutta, in a Lake District stream

SUMMARY. 1. Field experiments were performed in the day and night at six modal water velocities (range 10–52cm s⁻¹), using: (i) newly- emerged fry without neutral buoyancy; (ii) older fry in poor condition (weight well below that expected for resident fry); (iii) older fry in good condition (weight similar to that of resident fry); (iv) dead fry.
2. An exponential model described the return rate of fry to the stream bottom; the mean distance travelled downstream varied considerably between the four fry categories, but always increased linearly with increasing water velocity.
3. Results were similar for dead fry and newly-emerged fry released at night; 50% of the fry returned to the bottom in 10–11 s and nearly all returned in c . 70s, the maximum distance travelled ranging from c . 7 m at 10 cm s⁻¹ to c . 37m at 52cm s⁻¹, Newly-emerged fry released in the day returned slightly faster (54s for 99% return to bottom).
4. Older fry in poor condition returned to the bottom slightly faster in the day than at night, but took about 2 min and travelled about twice the distance covered by dead fry. Older fry in good condition returned to the bottom at the fastest rate (3–6s for 50% and c . 30s for the rest), and travelled only about half (at night) or a third (in day) of the distance covered by dead fry.
5. The implications of this investigation are discussed and it is concluded that, apart from water velocity, the age and condition of the fry were the two most important factors affecting their downstream movement.     

Concentrations and annual loading of dissolved organic matter in a small moorland stream

SUMMARY. 1. Dissolved organic matter (DOM) was measured at 8h intervals over a 1 year period in a stream draining 51 ha of moorland with peaty soils.
2. DOM concentrations increased with increasing stream discharge from low flow values of 0–3 mg 1⁻¹ to maximum values of 30 mg 1⁻¹. There were also seasonal differences of up to 13mgl⁻¹ between August (maximum) and February, and differences of about 2.5 mg l⁻¹ between rising and falling stage samples.
3. Seasonal variation was closely related to mean temperature.
4. Total loss over the year was 168 kg ha⁻¹ DOM (84 kg ha⁻¹ C), larger than previous estimates for upland sites.     

Immobilization of dissolved organic matter from groundwater discharging through the stream bed

SUMMARY. 1. In laboratory experiments, 9.7–25.7% of dissolved organic carbon (DOC) in groundwater (at concentrations of 18.7–24.8 mg 1-⁻¹) was immobilized after perfusion through 8-cm-deep (22-cm-diameter) cores of stony stream-bed substratum.
2. This represented immobilization rates of 7.1–23.5 mg m⁻² h⁻¹ or, extrapolated across the year, potential immobilization rates within the stream bed of 62.2–205.9g m⁻² yr⁻¹. Actual rates in the entire stream bed were probably higher because perfusion through the experimental cores did not reduce groundwater DOC concentrations to levels measured in the adjacent stream.
3. Natural concentrations of dissolved free amino acids (DFAAs) in groundwater were generally unchanged following perfusion through the cores, suggesting the maintenance of a dynamic equilibrium in their concentrations.
4. Selective enrichments of amino acids in groundwater (up to 20-fold) were entirely immobilized following perfusion, indicating their rapid retention and flux in this environment. Thus, immobilization of the bulk DOC in stream-bed cores probably did not reflect net reductions in dissolved free, low-molecular-weight material, with higher molecular weight, more 'refractory' material being immobilized instead.
5. We conclude that groundwater can contribute substantial amounts of DOC, both high and low molecular weight, to a stream ecosystem. The stream bed is the site at which much of this material could be initially immobilized and made available to the stream trophic structure.     

Egg-to-fry survival of the sea trout in some streams of Gotland

The quality of the surface water of Gotland (Sweden) streams was good, and did not limit sea trout egg-to-fry (ETF) survival. The oxygen concentration of the interstitial water was positively correlated to the permeability of the stream bed, and to the geometric mean diameter of the substratum. When the oxygen at 15 cm inside the stream bed was undiminished from surface values, the permeability of the stream bed was at least 6000 cm h^–1 and the geometric mean diameter of the substratum at least 15.0 mm. When the interstitial oxygen concentration increased, the interstitial ammonium concentration decreased, the NO₂ concentration remained stable and the NO₃ concentration increased. The ETF survival up to hatching was highly dependent upon the oxygen concentration of the interstitial water. For survival from hatching to emergence, the geometric mean diameter of the substratum was the most important factor. ETF survival >50% was observed where the interstitial oxygen concentration averaged at least 10mg I-¹, the substratum permeability at least 2000cm h-¹ and the substratum geometric mean diameter at least 15 mm. Other factors, such as overdigging and drying out of the spawning grounds could increase the ETF mortality.     

Biomass and oxygen dynamics of the epiphyte community in a Danish lowland stream

SUMMARY. 1. We examined the abundance and oxygen metabolism of epiphytic organisms on the dominant macrophyte, Potamogeton pectinatus , in headwaters of the eutrophic River Suså. Microbenthic algae were abundant in the stream during spring and macrophytes during summer.
2. The low macrophyte biomass in spring supported a dense epiphyte cover whereas the high macrophyte biomass during summer had a thin epiphyte cover of 10–100-fold lower abundance per unit area of macrophyte surface. The epiphyte community was dominated by microalgae in spring and by heterotrophs, probably bacteria, during summer. This seasonal shift was shown by pronounced reductions of the chlorophyll a content (from 2–3% to 0.1–0.7% of organic DW), the gross photosynthetic rate (from 20–85 to 3–15 mg O₂, g-¹ organic DW h⁻¹) and the ratio of gross photosynthesis to dark respiration in the epiphyte community (from 5–18 to 1). The reduced contributions of epiphytic microalgae correlated with reduced light availability during summer.
3. Both the density and the photosynthetic activity of epiphytic algae were low on a stream area basis relative to those of microbenthic algae and macrophytes. Rapid variations in water velocity and extensive light attenuation in water and macrophyte stands probably constrained the development of epiphytic algae. The epiphyte community was more important in overall stream respiration, contributing c. 10% to total summer respiration and c. 20% to summer respiration within the predominantly heterotrophic microbial communities on sediments and macrophyte surfaces.     

Lough Ennell: laboratory studies on sediment phosphorus release under varying mixing, aerobic and anaerobic conditions

SUMMARY. 1. The exchange of phosphorus between the epilimnetic (shallow zone) sediment and water column in Lough Ennell was investigated in laboratory experiments using five intact cores.
2. Variations in water mixing, sediment suspension and aerobic–anaerobic oxygen status in the water column and its effects on sediment phosphorus release rates were determined.
3. Experimental results indicated that phosphorus release is possible under both aerobic and anaerobic conditions. Aerobic release (0.025 mg P l⁻¹ over 5 days) was possible up to the point when mass resuspension of sediment occurred. Anaerobic release for the same period and mixing conditions was 0.183 mg Pl⁻¹.
4. The release rate under aerobic conditions at 10°C equates to an internal areal loading of 0.134 g P m⁻² yr^−1, which is approximately 17% and 30% of the average total phosphorus and orthophosphate loadings respectively for the period 1974–79.
5. The results clearly implicate aerobic inorganic phosphorus release from the epilimnetic sediments as a significant source of this nutrient to the overlying water column and is likely a major factor in the continuing eutrophic status in the lake.     

Age determination and growth of turbot and brill in the Adriatic Sea: reversal of the seasonal pattern of otolith zone formation

The growth of two commercially important flatfish, turbot ( Psetta maxima ) (L.) and brill ( Scophthalmus rhombus ) (L.), was investigated in the Adriatic using whole otoliths (sagittae) and stained otolith sections. At variance with the pattern usually observed in temperate seas, the opaque zone was found to be laid down in autumn and winter, and the translucent zone in spring and summer. Growth rates differed according to sex, with the females attaining greater body lengths. The von Bertalanffy growth parameters were: L_∞=66.2 cm, K=0.31 years^–1, and t₀=–0.14 years for turbot males, L_∞=81.5 cm, K=0.21 years^–1, and t₀=–0.48 years for turbot females; L_∞=40.2 cm, K=0.49 years^–1, and t₀=–1.03 years for brill males; L_∞=50.1 cm, K=0.27 years^–1, and t₀=–1.75 years for brill females. Growth rates and maximum age recorded for turbot were comparable to those reported in the North Sea.