Ecology of muskoxen in Jameson Land, northeast Greenland

Muskoxen Ovibos moschatus in Jameson Land exist at a density of somewhat more than 1 km⁻² of useable habitat and select moist meadows and snow bed vegetation for summer grazing and wind-exposed, dry dwarf shrub heath vegetation in winter. Graminoids dominate the winter diet and willows are the main component of the summer diet. Quality of the winter diet, as measured by the protein to fiber ratio is about one fourth that of the summer diet. During summer muskoxen supplement dietary sodium by using mineral licks. Muskoxen, especially females, retain considerable unused fat reserves through the winter and these are drawn upon during the post-calving period of lactation. Alternate year breeding is a common occurrence. Calves are frequently not weaned before the end of their first winter. Mean calf mortality is relatively low in the absence of significant predation and annual removal by hunting Inuits approaches the annual increment.     

Foraging strategies and seasonal diet optimization of muskoxen in West Greenland

Various aspects of optimal foraging and seasonal diet composition of bulls (bachelor and dominant), cows, subadults, and yearlings of muskoxen Ovibos moschatus were investigated in West Greenland during the following seasons: calving, post-calving, summer, rut and mid-winter. The following hypotheses were tested: (1) muskoxen maximize daily energy intake during spring and summer, (2) dominant bulls monopolizing cows during the rutting season shift from an energy maximizing to a time minimizing foraging strategy in order to maximize the time available for reproductive activities, and (3) muskoxen employ a time minimizing foraging strategy during winter to conserve energy. As forage quality changed throughout the short Arctic growing season, muskoxen responded by changing the proportions of daily time spent feeding on graminoids (Cyperaceae, Poaceae) and dicots (Salix, Betula), respectively. This seasonal variation in the relative proportion of daily feeding time spent ingesting graminoids followed approximately the energy maximization prediction over the periods calving to rut. Neither time minimizing nor random foraging could explain the observed diets in this period, thus confirming hypothesis 1. Dominant bulls did not shift to the time minimizing strategy as predicted by hypothesis 2. However, during the pre-rutting and rutting seasons bulls deviated from the other sex/age classes by failing to obtain the daily maximum energy predicted by the model, as a result of a higher proportion of time allocated to agonistic and sexual behaviour. During winter, none of the sex/age classes employed a time minimizing strategy, so rejecting hypothesis 3. Instead, muskoxen were found to maximize Na intake, indicating that Na is of major importance for winter survival. The results emerging from a linear programming model with constraint settings varying over seasons confirm that the constraint parameters applied are indeed important limiting factors for muskoxen in natural populations.     

Diet selection by vertebrate herbivores in the high arctic of Greenland

Microhistological analysis of fecal droppings indicates that graminoids (sedges, grasses, and rushes) and the only willow present, Salix arctica , are primary food items of the muskox Ovibos moschatus , arctic hare Lepus arcticus , and collared lemming Dicrostonyx groenlandicus in northern Greenland (79°-83°N). Plant biomass available during the peak growing season varied with vegetation types from < 2 g m⁻² in polar barrens to 34 g m⁻² in sedge fens. The clustered distribution of vegetation minimizes competition among vertebrate herbivores.     

Winter foraging by muskoxen: a hierarchical approach to patch residence time and cratering behaviour

We examined the temporal and spatial patterns of feeding behaviours of muskoxen during winter in the High Arctic. Pawing motions (to uncover forages beneath snow cover) were strongly aggregated into temporal bouts. Similarly, feeding stations (areas exploitable without motion of the forelegs) were aggregated into spatial patches. Muskoxen responded to greater snow accumulation at feeding sites by increasing the rates of pawing, rates of pawing bouts, number of pawing strokes per bout, and station residence times. Patch residence times showed little relationship to snow or forage abundance because, as muskoxen increased station residence times, they decreased the number of stations per patch. Muskoxen displaced one another from feeding stations more frequently as snow thickness and group size increased. Time spent at feeding stations was positively correlated to travel costs, in accordance with the marginal value model of patch residence. The model was not supported, however, at the scale of the feeding patch. The results indicate that behavioural responses of muskoxen to foraging conditions differ across scales.     

Planktic cyanophytes and their ecology in the shallow Lake Mikri Prespa, Greece

Cyanophytes dominated the phytoplankton of shallow Lake Mikri Prespa during the period May 1990 to September 1992 (76.5 and 52.0% of the total phytoplankton biomass in 1990–1991 and 1991–1992, respectively). Biomass peaks were observed in autumn (from 5.2 to 34.5 g m^-3) when low dissolved inorganic nitrogen, high phosphate phosphorus and low Z_cu to Z_mix ratio prevailed. The dominant species were Microcystis aeruginosa and M. wesenbergii. These represented 64 and 86% of the cyanophyte biomass in 1990–1991 and 1991–1992, respectively and revealed similar patterns of seasonality forming biomass peaks in late summer - autumn. Small chroococcalean cyanophytes (< 2 μm) showed also similar temporal distributions. Of the filamentous cyanophytes, the most important species was Anabaena lemmermannii var. minor which formed peaks in late summer and autumn (2.6 and 1.1 g m^-3 in 1990 and 1992, respectively). Rainfall and the N:P ratio were probably the main factors influencing the seasonality of all of the filamentous cyanophytes in the lake. Cyanophytes, tended to increase at temperatures higher than 16 °C and at inorganic nitrogen concentrations lower than 100 μg 1^--¹. The frequent mixing of the water column did not seem to prohibit the substantial increase of the group.     

A carbon budget for nematodes, rotifers and tardigrades in a Swedish coniferous forest soil

Nematodes, rotifers and tardigrades from a Swedish pine forest soil were investigated in a monthly sampling programme lasting for one year. The monthly mean values of number, biomass and oxygen consumption were estimated. All groups fluctuated rather much over the year with a summer minimum (1.1 ˙ 10⁶ animals m^-2) and a winter maximum (6.3 ˙ 10⁶ animals m^-2) for the nematodes. The reasons for these fluctuations are discussed in relation to fluctuations of water content and temperature of the soil. On an annual basis a carbon budget was calculated which gave the following values for this fauna; consumption 4.3, production 0.6, respiration 0.9 and defecation 2.8 g carbon m^-2.     

Temporal and spatial distribution of planktic cyanobacteria in Lake Kastoria, Greece, a shallow, urban lake

The temporal and spatial distribution of planktic cyanobacteria and some environmental parameters were studied in the shallow, urban Lake Kastoria, Greece from June 1996 to June 1997. Water temperature varied from 6–27 °C, pH from 7.5–8.9 and dissolved O₂ concentration from 0.7–12 mg m^-3 10^-3. The mean annual Chl a concentration was 83 mg Chl a m^-3 indicative of the eutrophic-hypertrophic state of the lake. Cyanobacterial biomass ranged from 11–238 g FW m^-3, constituting about 90% of the total phytoplankton biomass throughout the year. Cyanobacterial biomass was non-uniformly distributed both vertically and horizontally from August to November 1996 and resulted mainly from the distribution of Microcystis. Seven cyanobacterial taxa were reported for the first time in Lake Kastoria. Six taxa were dominant: Microcystis aeruginosa, M. flos-aquae, M. novacekii. Limnothrix redekei, Anabaena sp. and Cylindrospermopsis raciborskii. The dominant cyanobacterial taxa can be grouped on the basis of their distribution patterns (1) Microcystis species: maximum biomass occurring at pH > 8, temperature 12–17 °C, depth < 0.2 m; (2) Anabaena sp. and Cylindrospermopsis raciborskii : maximum biomass at temperatures 23–26 °C; (3) Limnothrix redekei : maximum biomass at temperatures 6–27 °C. Usually, non-uniform, vertical distributions of cyanobacterial biomass were associated with the formation of temperature, pH and O₂ gradients. L. redekei was considered to be a key lake organism since it contributed up to 59% of the cyanobacterial biomass. Interestingly, three of the dominant cyanobacteria Microcystis aeruginosa, Anabaena sp. and Cylindrospermopsis raciborskii belong to genera that include toxin-producing species.     

Seasonal hyperphagia does not reduce digestive efficiency in an Arctic grazer

Muskoxen (Ovibos moschatus) consume fibrous plants that grow rapidly over the short Arctic summer. We studied responses of eight castrated male muskoxen to a diet of grass hay and mineral supplements during spring, autumn, and winter. Animals gained body mass in spring (239+/-39 kg) as body fat content increased from 26% to 38% of ingesta-free mass in winter without changes in lean mass and protein. Intakes of dry matter (DM) increased by 74% between spring and autumn as digestible energy increased from 554 to 923 kJ kg(-0.75) d(-1) during mass gain. Digestibility of cellulose (72%-76%) was not affected by increasing food intake between spring and autumn but was reduced to 65% in winter. Digestibility of nitrogen compounds was 61%-66%, even though intake increased by 134% between spring and autumn. Excess dietary nitrogen from hay and supplements increased urea concentrations in plasma and urine. High loads of solutes such as potassium did not affect plasma or urinary osmolality but were associated with increased rates of glomerular filtration and urinary excretion. Low intakes of sodium from grasses may limit intake and digestion during summer, but high food intakes can support deposition of nitrogen, calcium, magnesium, copper, and zinc in body tissue even when dietary concentrations are low. Seasonal increases in digestive and metabolic functions allow muskoxen to rapidly accumulate energy and nutrients in body tissue during the short season of plant growth.     

Life cycle, production, microdistribution and diet of the damselfly Euphaea decorata (Odonata: Euphaeidae) in a Hong Kong forest stream

Euphaea decorata in Tai Po Kau Forest Stream (Hong Kong) was univoltine. Most recruitment took place in summer, and larval growth proceeded throughout the year. Life-cycles recorded in 1977–78 and 1978–79 were similar. Annual production estimates, using the removal-summation, instantaneous growth and size-frequency methods, were more similar for the 1978–79 generation (ranging from 158.7–174.7; mean 1671 mg dry wt m^-2) than for the 1977–78 generation (93.9-173-6; mean 131.7 mg dry wt m^-2). Mean biomass was similar for both generations (ranging from 33.5–33.9 mg dry wt m^-2), and mean P/B ratios were 3.9 1977–78) and 5.0 (1978–79). These are the first estimates of annual production by an Oriental stream insect.
Larvae were most abundant at microsites in the middle of the stream. Multiple regression analysis indicated that substratum characteristics were a major determinant of microdistribution. Euphaea decorata apparently favoured poorly-sorted sediments with highly peaked grain size-frequency distributions, containing few fine particles.
The carnivorous larvae showed ontogenetic changes in diet. Small individuals consumed mainly chironomid (Diptera) larvae; the diet expanded to include (successively) larvae of Ephemeroptera and Trichoptera as E. decorata grew. Seasonal changes in diet were also apparent, although larval diets during spring and summer were similar. There was also considerable overlap between autumn and winter diets. Ontogenetic influences upon prey consumed were not sufficient to account for the observed seasonal differences.     

Habitat selection as a hierarchy: the spatial scales of winter foraging by muskoxen

We studied the winter resource selection of muskoxen Ovibos moschatus in the High Arctic using a nested hierarchy of spatial scales 1) population range, 2) travel routes, 3) feeding sites (l e clusters of feeding craters), 4) feeding craters, and 5) diet (I e plant species) We found that, generally, patterns of selection remained consistent across all levels At successively smaller scales, muskoxen selected for higher graminoid abundance and particularly for thinner, softer snow cover, although we did not reject the hypothesis of random travel route selection Muskoxen uncovered forages from beneath the snow cover, by cratering, near the flonstic and nival extremes of availability Selection was consistently biased toward use of water sedge, Carex aquatilis As scale changed, however, muskoxen showed reversals of preference for some other forage species Diet was dominated by C aquatilis and cotton sedge, Eriophorum angustifolium , species characteristic of lowland meadows During spring melt, muskoxen moved to snow-free uplands to feed Dietary quality, as revealed by fecal nitrogen, increased at this time The consistency of the results across scales implied that these local levels of habitat selection occurred within one scaling domain     

Sampling strategy related to fish distribution, with particular reference to the Norfolk Broads

Large-scale movements of fish within a complex freshwater system such as the Norfolk Broads can give rise to erroneous sampling data if the extent of these movements is not fully appreciated. The Norfolk Broads habitat is briefly described, and data are presented showing that mean estimates of standing crop in the broads vary from <1gm^-2 in the winter to 9.4 gm^-2 in the summer. The widespread nature of the phenomenon is demonstrated by winter and summer data from five broads, and a more detailed picture of the seasonal changes is shown by an intensive 2-year study on one of these broads. Winter aggregations of fish have been found in sites adjacent to rivers connected with the broads, with densities of up to 36.7 fish m^-2 and a biomass of 1787 gm^-2. Accurate overall estimates of fish standing crop can only be obtained when fish are maximally dispersed during the summer months.     

Life cycle and population dynamics of Trochosa terricola Thorell (Araneae: Lycosidae) in a Norfolk grass heath

Abstract. 1. The population of the lycosid Trochosa terricola Thorell was sampled from April 1973 to August 1975 at Weeting Heath NNR, a Breckland grass heath.
2. Four sampling methods were compared for efficiency. Hand searching gave density estimates between 38.3 and 70.1% of heat extraction.
3. The temperature range in the sward at +1 cm was –5°C to 39°C with January and July means of 3.2°C and 17.4°C.
4. Eight male and nine female instars were determined and the life cycle extended over 2 or 3 years.
5. Adults were nocturnal but the juveniles diurnal. An annual diplochrone activity pattern was observed for adult males.
6. The horizontal distribution within the sward was aggregated, the structure and microhabitat being important determining factors. The population density was greater in moist, young Festuca spp. tussocks. An equation relating population density to habitat characteristics was derived.
7. The overall population density ranged from 14.0 m^-2 to 76.0 m^-2 and was maximal in autumn after breeding. The population biomass was greatest during autumn (291.2 mg d.wt m^-2).
8. Mean number of juveniles emerging from an egg sac was 77.3 (first sac) and 38.0 (second sac). The natality in 1973 was 66.8 individuals m^-2.
9. The survivorship curve until maturity varied between types I and III in different years.
10. The population dynamics were compared and are discussed in the light of other data. The variable population characteristics suggested that Den Boer's "spreading of risk" theory applied to the T.terricola population.     

Energy partitioning in the Antarctic collembolan Cryptopygus antarcticus

Abstract. 1. Consumption, production and assimilation rates were determined for two age groups of Crypropygus antarcticus to give an estimate of energy utilization, and to investigate low temperature adaptation in its energy partitioning.
2. Feeding selectivity shown in laboratory preference tests was supported by gut analysis of field animals from contrasting sites. Although moulting rate was not significantly affected by food type, rates of growth were slowest and mortality highest when fed on a non-preferred substrate.
3. Both a radio labelling and a more direct method for measuring dry weight consumed gave similar results for Cvpropygus feeding on algae. The consuniption rate for animals when feeding on algae was lower than that on moss peat. The assimilation efficiency for immature animals feeding on algae was 46% and for mature animals was 19%; the values when feeding on moss peat were 7% and lo%, respectively, The net production efficiency ranged from 35%(inimatures) to 13% (matures) and was similar on both substrates.
4. Food consumption exceeded assimilation over the range 2.5–10°C, but the two converged from 2.5 to 0°C. Immature Cryptopygus maintained a net positive energy balance over 0–10°C, whilst below 1S°C respiration exceeded assimilation for mature individuals.
5. An estimate of the annual dry matter consumption (7 g m^-1 y^-1) by Ctypropygus in a moss turf at Signy Island agrees with one based on respiration data alone (Davis, 1981). The consumption at an alga-dominated site was c . 26 g m^-2 y^-l, and Crypropygus may have a locally limiting effect on net priniary production at such sites.     

The effect of light intensity and spectrum on the incidence of first feeding by larval haddock

Under full–spectrum white light, feeding success of haddock Melanogrammus aeglefinus first feeding larvae, as measured both by proportion of larvae feeding and mean prey consumed, peaked at 1·7-18 μmol s^-1 m^-2. Feeding was significantly reduced at lower and higher intensities. A similar result was observed for larvae feeding under blue (470 nm) light, with significantly greater feeding success at intermediate light intensity (1·8 μmol s^-1 m^-2). When different light qualities were compared, larvae had significantly greater feeding success when exposed to blue (470 nm) light than either full-spectrum white or green (530 nm) light. Haddock larvae were capable of prey capture under all light treatments tested, indicating a necessary degree of adaptive flexibility in feeding response. The results are consistent with predisposition of haddock larvae to optimal feeding in a visual environment comparable with open ocean nursery grounds. Information on the impact of light on haddock first feeding can be incorporated into models of larval growth, survival, year-class strength and recruitment, and assist in developing husbandry protocols to maximize larval survival in aquaculture.     

Surface enlargement in the rumen of free-ranging muskoxen (<Emphasis Type="Italic">Ovibos moschatus</Emphasis>)

The intraruminal papillation pattern indicates the degree of rumen contents stratification and is related to the feeding niche of a ruminant. Muskoxen (Ovibos moschatus) display a variety of morphophysiological adaptations typical for grazers. We investigated the intraruminal papillation of 22 free-ranging muskoxen from five different months by comparing the surface enlargement factor both between seasons and between individual rumen regions. The seasonal pattern of rumen papillation indicated a distinct seasonality in food quality. The intraruminal papillation indicated a moderate degree of rumen contents stratification typical for intermediate feeders. The nutritional ecology of muskoxen is characterised by specific morphophysiological adaptations to a grass-dominated diet that nevertheless allow extensive seasonal use of browse forage.     

Kinetic changes with temperature of phosphoenolpyruvate carboxylase from a CAM plant

Abstract. Purified and crude phosphoenolpyruvate carboxylase from the CAM plant Kalanchoë daigremontiana Hamet et Perrier ( Bryophyllum diagremontianum ) was assayed at temperatures between 10 and 45° C. The optimum temperature of the enzyme activity changed with substrate availability and effector concentration in the assay. l -malate inhibited the enzyme activity and lowered the optimum temperature. Glucose-6-phosphate raised the optimum temperature to 43°C. K _m values for phosphoenolpyruvate increased with assay temperature from 0.12 mol m^-3 at 15° C to 0.36 mol^m−3 at 35° C. Inhibition by malate increased with temperature and acidity of the assay. In the crude enzyme 50% of control activity was inhibited by 1.65 mol m^-3 malate at 15° C and by 0.5 mol m^-3 at 35° C (at pH 7.0). With purification malate sensitivity was lost ( K _i values for malate at least 10 times higher). The shift in optimum temperatures for PEP-carboxylase activity thus results from changes in the kinetic parameters with temperature and allosteric effectors. The often low optimum temperatures for CO₂ fixation observed in nature may thus be the result of substrate and effector concentrations in the cytoplasm and the antagonistic effect of temperature on substrate affinity and effector efficiency on phosphoenolpyruvate carboxylase.     

Quantum yield and energy dependence for photooxidation of chlorophyllide in greening wheat

Dark grown leaves of wheat were irradiated with red light of different intensities, at a temperature close to 0°C. The rate of photoreduction of the protochlorophyllide 650-form into chlorophyllide 684-form was measured. On continued irradiation the chlorophyllide 684-form was photodecomposed. By comparing the rates of the two processes the quantum yield for photooxidation of the chlorophyllide 684-form was calculated. The quantum yield was 2°10^-5 at an intensity of 2200 W m^-2, and increased with decreasing light intensity to 3.2°10^-5 at an intensity of 170 W m^-2.     

Seasonal changes in biom ass of macrophytes on shaded and unshaded sections of the River Lambourn, England

SUMMARY. Monthly changes in the biomass of Ranunculus, Berula and Callitriche were recorded on two 50-m sites on the River Lambourn at Bagnor between March 1971 and October 1973. On the unshaded site. Ranunculus was the dominant macrophyte and characteristically its biomass changed from 20–40 g dry wt m^-2 in March to around 400 g dry wt m^-2 at the end of the growing season. On the shaded site, where Berula was the dominant macrophyte. the biomass of Ranunculus was lower than on the unshaded site. There was no significant difference between the mean biomasses of Berula on these two sites but Callitriche had a signifieantly higher biomass on the shaded site. Estimates of the total biomass of macrophyte on each site were calculated and the biomass of macrophyte removed during cutting operations each summer was also assessed.     

Dietary salt intake and its relevance to ionic regulation in freshwater salmonids

Dietary sodium intake for freshwater salmonids feeding in the wild (invertebrate diet) or in captivity (pellet diet) was calculated and compared with published branchial sodium influx values. Dietary sodium intake (mmol kg⁻¹ per month) increases from winter minimum values of 5 and 30-40 to reach maximum values in summer of 175 and 240 for invertebrate and pellet diet, respectively. In summer, dietary sodium intake for fish feeding in the wild was of the same magnitude as branchial sodium influx. The implications of dietary sodium intake for sodium balance in freshwater fish are discussed.     

The distribution and abundance of young (I +–II +) perch, Perca fluviatilis L., in a deep eutrophic lake, England

Seasonal changes in the vertical and horizontal distribution of young (I +–II +) perch in Rostherne Mere, a deep (max. depth 31.0 m) eutrophic lake are described using sonar. During the summer (June–August), perch were present within the thermocline (9–14° C) between 7 and 10 m. There was no significant difference between samples during this period, nor was there any evidence of shoaling. From September onwards perch moved into deeper waters. By late winter fish had become patchy in their distribution. Total numbers, during the summer, were estimated to be 988 ha ¹ with 95% confidence limits of ±115 (biomass 98.8 kg ha ⁻¹). Densities (no. m⁻³) fell by over 90% after the autumn overturn. The implications for predators are discussed. Acoustic sampling provides a useful, non-destructive method of sampling eutrophic lakes on a routine basis. Information obtained is of ecological, behavioural and physiological interest.