On the Origin of Brucellosis in Bison of Yellowstone National Park: A Review

Brucellosis caused by Brucella abortus occurs in the free-ranging bison ( Bison bison ) of Yellowstone and Wood Buffalo National Parks and in elk ( Cervus elaphus ) of the Greater Yellowstone Area. As a result of nationwide bovine brucellosis eradication programs, states and provinces proximate to the national parks are considered free of bovine brucellosis. Thus, increased attention has been focused on the wildlife within these areas as potential reservoirs for transmission to cattle. Because the national parks are mandated as natural areas, the question has been raised as to whether Brucella abortus is endogenous or exogenous to bison, particularly for Yellowstone National Park. We synthesized diverse lines of inquiry, including the evolutionary history of both bison and Brucella , wild animals as Brucella hosts, biochemical and genetic information, behavioral characteristics of host and organism, and area history to develop an evaluation of the question for the National Park Service. All lines of inquiry indicated that the organism was introduced to North America with cattle, and that the introduction into the Yellowstone bison probably was directly from cattle shortly before 1917. Fistulous withers of horses was a less likely possibility. Elk on winter feedgrounds south of Yellowstone National Park apparently acquired the disease directly from cattle. Bison presently using Grand Teton National Park probably acquired brucellosis from feedground elk.     

Prey risk allocation in a grazing ecosystem.

Understanding the behaviorally mediated indirect effects of predators in ecosystems requires knowledge of predator-prey behavioral interactions. In predator-ungulate-plant systems, empirical research quantifying how predators affect ungulate group sizes and distribution, in the context of other influential variables, is particularly needed. The risk allocation hypothesis proposes that prey behavioral responses to predation risk depend on background frequencies of exposure to risk, and it can be used to make predictions about predator-ungulate-plant interactions. We determined non-predation variables that affect elk (Cervus elaphus) group sizes and distribution on a winter range in the Greater Yellowstone Ecosystem (GYE) using logistic and log-linear regression on surveys of 513 1-km2 areas conducted over two years. Employing model selection techniques, we evaluated risk allocation and other a priori hypotheses of elk group size and distributional responses to wolf (Canis lupus) predation risk while accounting for influential non-wolf-predation variables. We found little evidence that wolves affect elk group sizes, which were strongly influenced by habitat type and hunting by humans. Following predictions from the risk allocation hypothesis, wolves likely created a more dynamic elk distribution in areas that they frequently hunted, as elk tended to move following wolf encounters in those areas. This response should dilute elk foraging pressure on plant communities in areas where they are frequently hunted by wolves. We predict that this should decrease the spatial heterogeneity of elk impacts on grasslands in areas that wolves frequently hunt. We also predict that this should decrease browsing pressure on heavily browsed woody plant stands in certain areas, which is supported by recent research in the GYE.     

Seasonal variation and diversity of rotifers in Ehoma lake, Nigeria

The zooplankton and physicochemistry of Ehoma lake were studied from March 2005 to August 2006 in order to evaluate seasonal trend in species composition and abundance in relation to water quality. Forty two (42) species of rotifers dominated by Keratella tropica tropica were encountered. Species richness declined by nearly 50% from dry season values during the rainy season. Species richness was higher in the littoral zone during the dry season, the reverse was the case during the rainy season. Species abundance followed similar trend as species richness. Abundance was closely associated with dissolved oxygen, biological oxygen demand and width of the lake and these in turn varied significantly between seasons. Shannon-Weaver diversity varied from 2.2 to 3.8 bit ind(-1) and significantly between seasons (p<0.05). Anuraeopsis navicula navicula and Euchlanis dilatata dilatata Ehrenbreg were the only species gained during the flood periods.     

Sex-based differences in density-dependent sociality: an experiment with a gregarious ungulate

For animals living in natural or semi-natural settings, empirical data on how sociality changes in response to increasing population density are few, especially with respect to true conspecific density and not group size. However, insight into this line of research may be far-reaching--from understanding density dependence in sexual selection to improving models of disease transmission. Using elk (Cervus elaphus Linnaeus) held in enclosures, we conducted sex-stratified experiments to test how the frequency of dyadic pairings (interaction rate) and their quality (duration) responded to manipulations in exposure to density. Using proximity-logging radio collars we recorded when and for how long individuals shared a space within 1.4 m of each other. As predicted, males increased their interaction rate as density increased. Female interaction rates, however, increased initially as density increased but soon declined to become indistinguishable from rates at low density. Females interacted for longer periods at medium densities, whereas male interaction length clearly decreased as density increased. We highlight a sexually dichotomous, density-dependent response in sociality that has yet to be reported. In addition to furthering our understanding of sociobiology (e.g., implications of time constraints presented by density on dyadic interactions), our results have implications for managing communicable disease in gregarious species of livestock and wildlife.     

Daily and seasonal feeding rhythmicity of <Emphasis Type="Italic">Palaemonetes varians</Emphasis> (Leach 1814) from southwestern Europe

The feeding activity rhythm of Palaemonetes varians (Leach 1814) was studied by collecting stomachs over a 24-h cycle in four different seasons of 1996 and spring–summer of 1997 to assess a regulation of this behaviur dependent upon seasonal variations in photoperiod length. A total of 6,692 stomachs were collected from a population reared in an extensive earthen pond located in the Guadalquivir Estuary salt marshes (Spain). Stomach fullness (f) classification was performed with a stereomicroscope by means of a four-point visual qualitative scale. Results showed that a high proportion of stomachs were empty in all seasons considered and a great variability in the data occurred irrespective of the time of the day. The number of full stomachs at each catch were represented over 24-h for each sampling season. Time series showed the occurrence of a feeding activity rhythm phased at day–night transitions. The sunset peak occurred in all months of sampling, while the sunrise one occurred in summer. Results are discussed in relation to available data on behavior, moulting, and reproduction of this and other species of the same genus.     

Analysis of water quality factors influencing the speciation of inorganic nitrogen using GRA

Based on the water quality data from 2006 to 2008, grey relational analysis (GRA) is used to analyze factors that may have influence on the speciation of inorganic nitrogen in the Chengdu section of middle Min river. The results show that water temperature, changing from 20.2 +/- 2.7, 13.4 +/- 5.7 and 16.8 +/- 5.6 degrees C, is the first restrictive factor for the speciation of inorganic nitrogen; it is negatively correlated with the ratio of total ammonia nitrogen to total inorganic nitrogen contents [m(AN)/m(TIN)] in three different periods of wet season, dry season and year-round. The average pH values for years, in wet and dry periods are 7.6 +/- 0.4, 7.3 +/- 0.3 and 7.8 +/- 0.2, respectively, and have different influences in different seasons. It is the second restrictive factor and positive correlation between pH and m(AN)/m(TIN) in wet season and through the year yet it is the fourth factor in dry seasons. The values of dissolved oxygen (DO), which are 4.6 +/- 1.4, 4.6 +/- 2.4, 4.6 +/- 2.0 respectively, is the third factor and negatively correlates with m(AN)/m(TIN) in third different periods. The chemical oxygen demand (COD) indirectly inhibits the nitrifying bacteria because the DO is depleted in the decomposition of organic matter by heterotrophic bacteria, showing the positive correlation.As the alkalinity can meet the requirement of nitrification in wet season and through the year, it is not restrictive factor. However, it is the second restrictive factor in dry season because of low content of alkalinity inhibiting the growth of nitrifying bacteria.     

Population density and the intensity of sexual selection on body length in spatially or temporally restricted natural populations of a seed bug

Summary Population density affects the dynamics of mate acquisition and the opportunity for sexual selection in natural populations of the seed bug, Neacoryphus bicrucis Say (Hemiptera : Lygaeidae). The opportunity for sexual selection and the intensity of directional sexual selection on body length increased as the population density declined within a season for a population in a small, disjunct patch of host plant, Senecio anonymus. In a larger, dispersed population, both measures of selection were greater in host plant patches of low rather than high adult density when the population was sampled at peak density. Under conditions of higher density, males were more likely to share plants, larger males were less likely to monopolize patches of host plant to which females were attracted for mating, and smaller males were more likely to mate in the presence of large males. Thus, resource defense polygyny collapsed under high density, obviating the advantage of size in territory control, and resulted in scramble competition among males for mates. The population exhibited significant additive genetic variation for body length. This suggests that natural selection acting on other components of fitness favors smaller size or that the direction of sexual selection on size fluctuates between generations in response to the between-year variation in population density. Thus, strong sexual selection appears to impose a significant genetic load.     

Bill color, not badge size, indicates testosterone-related information in house sparrows

The honesty of ornamental signals of quality is often argued to be enforced via costs associated with testosterone. It is still poorly understood, however, how seasonal variation of testosterone within individuals is related to the timing and extent of ornament development. Here, we studied inter- and intra-individual variability of plasma testosterone levels in a population of 150 captive male house sparrows (Passer domesticus) through the course of a full year. We further analyzed the relationship between plasma testosterone levels and two sexually dimorphic ornaments: badge size and bill coloration. Also, because of a known negative relation between molt and circulating testosterone levels, we analyzed the relationship between ornamentation and molt status during the fall. We found that testosterone levels increased towards the breeding season and decreased before the onset of annual molt. However, within individuals, relative testosterone titers demonstrated low repeatability between seasons. Plasma testosterone levels were not correlated with badge size in any season but were correlated strongly with bill coloration during all periods, except the breeding season when variation in bill color was low. Finally, we found that bill coloration strongly correlated with molt status during fall. Our results indicate that bill coloration, not badge size, is the best ornamental indicator of a “running average” of male testosterone in house sparrows and therefore the best potential indicator of qualities and/or behavioral strategies associated with testosterone.     

Seasonal changes in the structure of rhesus macaque social networks

Social structure emerges from the patterning of interactions between individuals and plays a critical role in shaping some of the main characteristics of animal populations. The topological features of social structure, such as the extent to which individuals interact in clusters, can influence many biologically important factors, including the persistence of cooperation, and the rate of spread of disease. Yet, the extent to which social structure topology fluctuates over relatively short periods of time in relation to social, demographic, or environmental events remains unclear. Here, we use social network analysis to examine seasonal changes in the topology of social structures that emerge from socio-positive associations in adult female rhesus macaques (Macaca mulatta). Behavioral data for two different association types (grooming and spatial proximity) were collected for females in two free-ranging groups during two seasons: the mating and birth seasons. Stronger dyadic bonds resulted in social structures that were more tightly connected (i.e., of greater density) in the mating season compared to the birth season. Social structures were also more centralized around a subset of individuals and more clustered in the mating season than those in the birth season, although the latter differences were mostly driven by differences in density alone. Our results suggest a degree of temporal variation in the topological features of social structure in this population. Such variation may feed back on interactions, hence affecting the behaviors of individuals, and may therefore be important to take into account in studies of animal behavior.     

Breeding success of tree swallows along a gradient of agricultural intensification

The intensification of agricultural practices has been identified as the main cause of population decline in farmland birds since the 1960s in both Europe and North America. Although the links between species richness or abundance and various components of agricultural intensification are well established, the mechanisms underlying these trends have rarely been addressed along a gradient of intensification or have been quantified at only one spatial scale. Here we quantified the influence of landscape structure on the nest box occupancy and breeding success of Tree Swallows (Tachycineta bicolor) at seven spatial scales (1, 2, 3, 4, 5, 10, and 20 km radii) over a 10 200-km2 gradient of agricultural intensification in southern Québec, Canada. A network of 400 nest boxes distributed among 40 farms was visited every two days over three breeding seasons, 2004-2006. Nest box occupancy decreased with the proportion of intensive cultures (maize, cereals, and soybeans) in the landscape, especially when manure heaps and tanks were abundant, and was also determined by local variables (i.e., nest box clearance, interspecific competition) and by previous-year fledging success. Clutch size decreased as the breeding season progressed and with the proportion of intensive cultures in the landscape, with no consistent variation across spatial scales. Hatching success was not related to any landscape variables but increased with clutch size. Both the number of fledglings and fledging probability increased with the proportion of extensive cultures (hayfields, pastures, and fallows). These effects increased with spatial scale and reached a plateau at the 5 km radius: the maximum distance from the nest reached by foraging Tree Swallows. Our results can likely be attributed to lower food availability in intensive cultures compared to extensive ones. This study suggests that several components of breeding that impact on population structure and dynamics of insectivorous birds will be negatively affected by agricultural intensification.     

Mineral licks: motivational factors for visitation and accompanying disease risk at communal use sites of elk and deer

Free-ranging cervids acquire most of their essential minerals through forage consumption, though occasionally seek other sources to account for seasonal mineral deficiencies. Mineral sources occur as natural geological deposits (i.e., licks) or as anthropogenic mineral supplements. In both scenarios, these sources commonly serve as focal sites for visitation. We monitored 11 licks in Rocky Mountain National Park, north-central Colorado, using trail cameras to quantify daily visitation indices (DVI) and soil consumption indices (SCI) for Rocky Mountain elk (Cervus elaphus) and mule deer (Odocoileus hemionus) during summer 2006 and documented elk, mule deer, and moose (Alces alces) visiting licks. Additionally, soil samples were collected, and mineral concentrations were compared to discern levels that explain rates of visitation. Relationships between response variables; DVI and SCI, and explanatory variables; elevation class, moisture class, period of study, and concentrations of minerals were examined. We found that DVI and SCI were greatest at two wet, low-elevation licks exhibiting relatively high concentrations of manganese and sodium. Because cervids are known to seek Na from soils, we suggest our observed association of Mn with DVI and SCI was a likely consequence of deer and elk seeking supplemental dietary Na. Additionally, highly utilized licks such as these provide an area of concentrated cervid occupation and interaction, thus increasing risk for environmental transmission of infectious pathogens such as chronic wasting disease, which has been shown to be shed in the saliva, urine, and feces of infected cervids.     

Clustering of the abundance of West Nile virus vector mosquitoes in Peel Region,Ontario, Canada

Understanding the spatial–temporal distribution of vector mosquitoes is essential in designing an efficient mosquito control strategy to reduce the risk of the mosquito-borne disease. In this paper, we apply a non-parametric clustering method, CLUES, to the surveillance data of West Nile virus vector mosquitoes collected by light traps in Peel Region, Ontario, during the mosquito seasons in 2004–2010. In order to obtain robust and reliable results, a statistical smoothing procedure LOWESS is applied to the original time series data. It was found that the mosquito trap sites can be clustered into three groups. The weather impact on the mosquito abundance of each clustered group are similar, while the interannual variability and the highest abundance and peak time in each mosquito season are different. The impact of weather factors on this clustering is investigated.     

Sardine (Sardina pilchardus) spawning seasonality in European waters of the northeast Atlantic

Egg data from ichthyoplankton monitoring sites in the western English Channel (1988–2003) and northern Spain (1990–2000) and macroscopic maturity data from biological samples of purse seine landings in western and southern Iberia (1980–2004) are used to describe the spawning seasonality of sardine (Sardina pilchardus) in European waters of the northeast Atlantic using generalised additive models. The fitted models reveal a double peak in spawning activity during early summer and autumn in the western Channel, a wider spring peak off northern Spain and a broad winter season in the western and southern Iberian Peninsula. At all sites, a high probability of spawning activity was observed over at least 3 months of the year, with the duration of the season increasing with both decreasing latitude and increasing fish size. Off western and southern Iberia there are indications that the spawning season has been of longer duration in recent years for all size classes (reaching in some cases 8 months of the year for large fish). These patterns are in general agreement with existing literature and theoretical expectations of sardine spawning being driven locally by the seasonal cycle of water temperature, assuming preferences for spawning at 14 –15°C and avoidance for temperatures below 12°C and above 16°C. Regional quotient plots indicated that spawning tolerance to higher temperatures increases progressively with decreasing latitude. Despite the weak evidence for geographical differences in temperature tolerance that may have some genetic origin, the degree of spatio-temporal overlap in sardine-spawning activity within Atlantic European waters is unlikely to promote any reproductive isolation in that area.     

Food abundance affects energy intake and reproduction in frugivorous female Assamese macaques

In most mammals, female fertility and reproduction are strongly influenced by nutritional status and, therefore, by foraging conditions. Here, we investigate the relationship between food resources, feeding competition, energy intake and reproduction in a group of wild female Assamese macaques (Macaca assamensis) in northeastern Thailand. Over 2,100 h of data on feeding behaviour, energy intake and activity budgets were combined with data on resource characteristics, female reproduction and physical condition. We found that an increase in food availability had a positive effect on female energy intake and conception rates. In addition, it appeared that females incurred energetic costs during lactation and that females with a better physical condition during the mating season were more likely to conceive. The annual birth season occurred a few months before the annual peak in food availability, causing peak lactation to coincide with a period of high food availability. This suggests that females use the increased food abundance to compensate for the energetic costs of lactation. Neither energy intake rates nor activity budgets were influenced by female dominance rank, even during periods when the levels of contest competition were predicted to be high. In line with this, we found no evidence for rank-related differences in reproduction. The apparently limited influence of feeding competition in female Assamese macaques adds to the debate on the extent to which patterns in feeding competition and fitness can reliably be predicted based on ecological conditions. We suggest that this may partially be resolved by including potential competition-reducing mechanisms into the predictive framework.     

Complex social structure of southern flying squirrels is related to spatial proximity but not kinship

Social individuals have organized relationships that affect fitness and so a species' tendency to be social has important implications for its population ecology, gene flow, and its distribution in space and time. We quantitatively examined the social structure of southern flying squirrels (Glaucomys volans) and tested for a role of kinship and prior familiarity in predicting social structure. To quantify social structure, we monitored nest group composition of southern flying squirrels. All squirrels at the study site were marked with passive integrated transponder (PIT) tags and nest cavity entrances were monitored with automated PIT tag recorders for a period of 28 months. Squirrels were genotyped at eight microsatellite loci. Permutation tests of associations suggested that individuals nested with other specific individuals more often than expected by chance. The lagged association rate indicated that relationships were stable and persisted across seasons and years. Multiple summer nest associates came together in winter to form larger nest groups which were likely important for social thermoregulation. A measure of prior familiarity, but not kinship, was related to the proportion of time individuals nested together during winter. We suggest that the evolution of sociality in southern flying squirrels is driven largely by mutually beneficial behaviors related to social thermoregulation although other, as of yet, unidentified mechanisms are needed to explain sociality in the warm season. We hypothesize that minimum group size requirements associated with social thermoregulation could explain the absence of this species in patchy landscapes and aspects of range boundary dynamics near their northern range boundary.     

Habitat use pattern of tidal mud and sand flats by shorebirds (charadriiformes) wintering in southern India

Habitat use pattern and status of shorebirds in southern India were investigated in six tidal flats comprising three mudflats and three sand flats on the east coast of southern India. Totally, 7757 shorebirds belonging to 21 species were recorded during September 2000–March 2002. The birds preferred mudflats over sand flats. Density, diversity and richness of shorebirds were relatively higher in tidal flats, especially in mudflats. The shorebirds preferred mudflats over sand flats as stopover sites and sites for refueling the required energy during migration. Rhythmic changes in tidal pattern also influenced the allochthonous nutrients and prey species, which also attracted the migratory shorebirds. However, the abundance of shorebirds with reference to tidal flats *season*tide and habitats*seasons * tide had strong variations (P < 0.001). During migratory season, bird density, diversity and species richness were observed higher than in other seasons.     

Irruptive population dynamics in Yellowstone pronghorn.

Irruptive population dynamics appear to be widespread in large herbivore populations, but there are few empirical examples from long time series with small measurement error and minimal harvests. We analyzed an 89-year time series of counts and known removals for pronghorn (Antilocapra americana) in Yellowstone National Park of the western United States during 1918-2006 using a suite of density-dependent, density-independent, and irruptive models to determine if the population exhibited irruptive dynamics. Information-theoretic model comparison techniques strongly supported irruptive population dynamics (Leopold model) and density dependence during 1918-1946, with the growth rate slowing after counts exceeded 600 animals. Concerns about sagebrush (Artemisia spp.) degradation led to removals of >1100 pronghorn during 1947-1966, and counts decreased from approximately 700 to 150. The best models for this period (Gompertz, Ricker) suggested that culls replaced intrinsic density-dependent mechanisms. Contrary to expectations, the population did not exhibit enhanced demographic vigor soon after the termination of the harvest program, with counts remaining between 100 and 190 animals during 1967 1981. However, the population irrupted (Caughley model with a one-year lag) to a peak abundance of approximately 600 pronghorn during 1982-1991, with a slowing in growth rate as counts exceeded 500. Numbers crashed to 235 pronghorn during 1992-1995, perhaps because important food resources (e.g., sagebrush) on the winter range were severely diminished by high densities of browsing elk, mule deer, and pronghorn. Pronghorn numbers remained relatively constant during 1996-2006, at a level (196-235) lower than peak abundance, but higher than numbers following the release from culling. The dynamics of this population supported the paradigm that irruption is a fundamental pattern of growth in many populations of large herbivores with high fecundity and delayed density-dependent effects on recruitment when forage and weather conditions become favorable after range expansion or release from harvesting. Incorporating known removals into population models that can describe a wide range of dynamics can greatly improve our interpretation of observed dynamics in intensively managed populations.     

Estimating the influence of overhead transmission power lines and landscape context on the density of little bustard Tetrax tetrax breeding populations

Collision with conductors and earth cables is a known impact generated by transmission power lines, however there is virtually no information on how these infrastructures might affect bird distribution in a landscape context. With this work we specifically hypothesise that transmission power lines may affect the occurrence of a threatened bird, the little bustard (Tetrax tetrax). To test this hypothesis we used a Stochastic Dynamic Methodology (StDM), analysing the effects of power lines in a landscape perspective and simulating population trends as a response to power line installation and habitat changes induced by agricultural shifts in southern Portugal. The data used in the dynamic model construction included relevant gradients of environmental conditions and was sampled during the breeding seasons of 2003-2006. Transmission power lines were significantly avoided by the little bustard and the developed StDM model showed that the distance to these utility structures is the most important factor determining breeding densities in sites with suitable habitat for the species, which possibly leads to displacement of populations and habitat fragmentation. The model simulations also provided the base to analyse the cumulative effects caused by the habitat degradation that can ultimately lead to the extinction of local populations. Within priority conservation sites, the dismantling of existing transmission lines should be considered whenever possible, in order to ensure adequate breeding habitat. The model is considered useful as an auxiliary tool to be used in environmental impact assessments, management and conservation studies.     

Length-structured analysis of the reproductive season of anchovy and common sardine off central southern Chile

The anchovy (Engraulis ringens) and common sardine (Strangomera bentincki) are two small pelagic fish which have a similar reproductive strategy off central southern Chile. The seasonal reproductive dynamics of both the species was investigated by taking into account the gonadosomatic index (GSI) and visual maturity data as a function of female size for the period 1993–1999, and also the seasonal pattern in condition factor per size classes. Larger females of common sardine (repeat spawners) have a reproductive peak earlier in the season than younger females at first maturity, while larger anchovy have a reproductive peak delayed as compared with first-time spawners. The condition factor of females exhibited an inverse cycle with the seasonal pattern of GSI, and delayed by approximately 6 months. Although larger females tend to present better and wider condition than smaller females, the condition of females seems to be delayed in larger females probably mediated by the seasonal pattern in food availability. This “energy storage strategy” in spring and summer time seems to be size-dependent and past energetic reserves could also affect the egg production and timing of reproduction. We concluded that the reproductive season of anchovy and common sardine is different and mediated by the length structure in the seasonal upwelling system off central southern Chile.     

Life cycle, seasonal vertical distribution and feeding of Calanus finmarchicus in Skagerrak coastal water

 Day/night zooplankton sampling in Kosterfjorden, Sweden, gave information on population succession, vertical distribution and feeding of Calanus finmarchicus over 17 months. Copepodid Stage 1 and 2 (C-I, C-II) were present from December to August, indicating reproduction during most of the year. Mating and breeding for the overwintering generation mainly took place in February/March, resulting in peak abundance of C-I in March/April. Secondary breeding periods were in April and July/August, but the resulting recruitment from these were low. The relative recruitment success in the first spring was around 13 times higher than during the second spring. Low temperature in the surface water early in the year and depression of the spring phytoplankton bloom may have caused the failure in the second year. Population biomass peaked at >15 g dry weight m⁻² during the numeric peak of the youngest stages in March/April. A more sustainable level of high biomass of 8 to 10 g dw m⁻² was gradually built up during summer, mainly due to a continuous accumulation of C-Vs. Adults and C-Vs comprised the overwintering population, with 7 to 14% and 85 to 93%, respectively, for the 2 years, but only C-Vs staying in the deep water were in a resting state. Adult males showed a strong diurnal vertical migration (DVM) of the usual type from spring to early autumn and a reversed DVM during the cold season. They were in a feeding state throughout the year. Adult females showed the usual type of DVM during summer to autumn, but commonly a reversed DVM during winter to spring. They were usually in a feeding state, with no pronounced differences between surface and deep water or between day and night. C-Vs aggregated in the deep water from October to March and performed DVM in April to June. They were commonly feeding in the deep water between March and June but showed no or very low feeding activity there from July to February. C-Vs in the surface water were commonly feeding and showed the highest proportion of feeding in autumn, when the population in the deep water was inactive. C-V constituted up to nearly 100% of the population biomass, and therefore must be of profound ecological importance. Defined by this dominant role, the population of C. finmarchicus can be characterised as having an active period of feeding, reproduction and development from February to July with a following 6 to 7 months of resting in the deep water, when development is arrested and no feeding occurs. Received: 1 October 1999 / Accepted: 27 April 2000