Dynamics of coexisting Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus populations in a high-Arctic fjord

We studied the population dynamics of Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus in Billefjorden, Svalbard (78°40N). All three species reproduced in the fjord with different timing. The maximum abundance of Calanus spp. copepodite stages peaked on the 11th of July (29,000 ind m^–2). C. glacialis was the dominant species accounting for 60–80% of the total Calanus abundance. C. finmarchicus appear to thrive in the fjord despite the low temperatures (–1.86°C to 5°C) and accounted for 20–30% of the total population. C. hyperboreus contributed less to the total abundance (5–20%). A 1-year life cycle is suggested for C. finmarchicus and C. hyperboreus in the fjord, C. glacialis has a 1- to 2-year life cycle. Highest mortality rates were observed for copepodite stage CV in C. finmarchicus and C. glacialis (0.09 and 0.075 d^–1, respectively) and for females in C. hyperboreus (0.149 d^–1). Mortality of copepodite stages was substantially lower in C. glacialis than in the other species. This is particularly obvious in the early and numerous copepodite stages (CI + CII) during the period of recruitment to these stages. This suggests that differences in secondary production in Arctic pelagic ecosystems are controlled partly by population loss rates.     

Lipid composition of copepodite stages and adult females of Calanus glacialis in Arctic waters of the Barents Sea

Summary The composition of lipid classes and their component fatty acid are described for copepodite stages III, IV, V, and adult females of Calanus glacialis sampled from Arctic waters of the Barents Sea during summer. Was esters were the principal component of the lipid in all copepodite stages examined, averaging 73% over all the stages. The proportion of triacylglycerols varied from 1.5% to 10.5% of total lipid among copepodite stages. The lipids of adult females contained lower levels of wax esters and higher levels of triacylglycerols than copepodite stages III, IV and V. Fatty alcohols of wax esters from copepods sampled in June and July were dominated by 20:1 (n-9) and 22:1 (n-11) alcohols with the proportion of 20:1 (n-9) increasing from stages III to adult female. 14:0 and 16:0 alcohols were the principal fatty alcohols of wax esters of a sample comprising mainly of copepodites stage III taken in August. 16:1 (n-7), 20:1 (n-9) and 20:5 (n-3) were the major fatty acids in the was esters of animals captured in June and July whereas 18:1 (n-9) predominated in the August sample. The polar lipids of the copepodite stage III from August also contained lower levels of polyunsaturated fatty acids than from all stages of copepods from June and July. The data are discussed in relation to life cycle strategies and trophic aspects of Calanus glacialis in the Arctic pelagic community of the Barents Sea.     

Differences in food delivered to chicks by males and females of little auks ( Alle alle ) on South Spitsbergen

Sex differences in the quality and quantity of food loads brought to little auk (Alle alle) chicks were investigated in a large colony in Hornsund (South Spitsbergen). Adults returning to the colony were caught in mist-nets and food loads were taken from their gular pouch. The sex of each bird was determined by means of molecular methods. Females brought significantly more food per single load than males in terms of wet weight (30% more on average), number of prey items (46%) and energy contents (39%). However, there was no difference between the sexes in the size of prey taken. Energy-rich Calanus glacialis, originating from cold Arctic waters, was the most frequent prey item and made up the majority of food loads brought by both males and females (75 and 72%, respectively). This indicates that both sexes forage mainly in Arctic waters. However, differences in the proportion of cold water Calanus species (C. glacialis and C. hyperboreus), warm water Calanus species (C. finmarchicus), as well as other taxa, between males and females may suggest different ways of exploiting the feeding area.     

Body sizes,development rates,and genome sizes among Calanus species

Data on embryonic and larval development times (D) of Calanus species are analysed using Blehrádek's temperature (T) function, D = a (T – )^b, with b = – 2.05 as in previous studies. Among these species, for embryonic duration varies directly with temperatures in their geographical ranges and a is related to egg diameter. Using and b from embryonic durations, the fitted values of a for older stages are related to body sizes. Roughly estimated nucleus numbers in single adult females of C. finmarchicus, glacialis and hyperboreus were similar at 72 000, 85 000, and 96 000 respectively. Genome sizes (2C) of adult females are ca. 13 pg DNA in C. finmarchicus and pacificus, ca. 17 pg in C. sinicus, ca. 21 pg in C. helgolandicus and marshallae, and ca. 25 pg in C. glacialis and hyperboreus. These correspond roughly to body sizes and temperature-corrected development rates, quite precisely so in the sibling pair C. finmarchicus and C. glacialis, suggesting that, given similar nucleus numbers, there is nucleotypic control of whole-organism characteristics.     

Distribution of dominant calanoid copepod species in the Greenland sea during late fall

Summary Between 6 November and 12 December 1988, vertical distributions of Calanus finmarchicus, C. hyperboreus, C. glacialis and Metridia longa were studied at three stations in the Arctic water of the Greenland Sea Gyre (GSG) and compared with two stations in the Atlantic water (AW) of the Westspitsbergen Current. Nine depth strata down to 3,000 m were sampled. C.finmarchicus was most abundant in AW, C. hyperboreus in GSG, M. longa showed no preference and C. glacialis was rare everywhere. Stage composition differed with species and water mass. Vertical distribution varied also with water mass in all species but C. hyperboreus, which was always centered between 1,000 and 1,500 m. The other species were concentrated in the upper 300 m in AW and between 1,000 and 1,500 m in GSG, although not all stages followed this trend. Ontogenetic vertical migration leads to significant dislocations of organic matter from the euphotic zone to great depth in the Greenland Sea. In egg production experiments, C. hyperboreus spawned up to 149 eggs female^–1 day^–1. Implications of stage composition, sex ratio, and gonad maturation on copepod life cycles are discussed.     

Assimilation and biosynthesis of lipids in Arctic Calanus species based on feeding experiments with a C labelled diatom

The dominant Arctic Ocean and North Atlantic copepods Calanus hyperboreus, Calanus glacialis, and Calanus finmarchicus were collected in the Greenland Sea and fed ¹³C labelled diatom Thalassiosira weissflogii to follow the transfer and assimilation of carbon, lipid, and individual fatty acids and alcohols. The diatom was grown with ¹³C for 3 to 5 days and fed then to the copepods. During the feeding period of 14 days, total carbon increased in the copepodite stages V of C. hyperboreus and C. finmarchicus, whereas carbon remained almost constant in C. glacialis females. However, total lipid increased in all species and stages. Highest lipid accumulation occurred in C. hyperboreus in which nearly all lipids were exchanged already after 11 days of feeding. In the other species lipid accumulation made up between 22% (C. finmarchicus) and 45% of total lipid (C. glacialis). The proportion of wax esters was high ranging from 76% of total lipid in C. glacialis to 92% in C. finmarchicus. The fatty acid composition of the alga was dominated by 16:1(n-7), 16:0, 20:5(n-3), and 22:6(n-3). The composition of the copepods was similar because of feeding already on diatoms in the field. In addition, the monounsaturated fatty acids and alcohols, 20:1(n-9) and 22:1(n-11), were major components of the copepod lipids. During the feeding period the highest ¹³C labelling was always found in the C₁₆ polyunsaturated fatty acids and in the 16:1(n-7) alcohol. Because these components occurred only in trace amounts in the copepods they totally originated from the diet explaining the high labelling. It is noteworthy that the 16:1(n-7) alcohol originated only from the corresponding dietary and not from the abundant internal fatty acid. The long-chain monounsaturated fatty acids and alcohols, 20:1(n-9) and 22:1(n-11), are not existent in phytoplankton and have to be produced de novo. They were less labelled in the smaller species but highly ¹³C enriched in C. hyperboreus. Although dietary fatty acids were generally retained by the copepods it seems that fatty acids or even lipids were selectively accumulated and turned over due to bodily requirements, and thus, essential polyunsaturated fatty acids were preferentially retained. During feeding mixing, accumulation, and exchange of internal and dietary fatty acids and alcohols occurred as well as utilisation of lipids from both sources for metabolic requirements. The differences in lipid assimilation fit to the different life strategies of the copepods.     

Spatial distribution of digestive enzyme activities of Calanus finmarchicus and C.hyperboreus in Fram Strait/Greenland Sea

Digestive enzymes of copepodite V (CV) Calanus finmarchicusand C.hyperboreus from two different depths were compared duringMIZEX 1984 (June/July) at stations in the ice and open water.CV of both species from 500–200 m showed reduced enzymeactivity, indicating that they were in a resting stage. In moultingexperiments at the end of June using CV from 100–0 m moultingwas delayed and began only after 3 weeks in C.finmarchicus andafter 3 months in C.hyperboreus. These results suggest thatthe deep CV populations are the seed of the new overwinteringstock. In surface CV C.finmarchicus and C. hyperboreus enzymeactivities were generally much higher than in deep CV. In neitherspecies were enzyme activities correlated with chlorophyll concentrations.Activities in C.finmarchicus reflected overall phytoplanktondistribution and were highest in the marginal ice zone, whereasthey decreased under the ice except for polynya stations onthe East Greenland Shelf. In surface C.hyperboreus digestiveenzymes were not correlated with those of C.finmarchicus, pointingto different diets or regulatory mechanisms. Enzyme activitywas lowest in the marginal ice zone and increased under theice. High activities were found at polynya stations and otherclose pack ice. The utilization of ice algae by C.hyperboreuscould explain these discrepancies in digestive enzyme activitiesof both species.     

The lipid biochemistry of calanoid copepods

Calanus species, particularly those in high latitudes, can accumulate large oil reserves consisting predominantly of wax esters. These wax esters consist predominantly of 16:0, 20:1 (n–9) and 22:1 (n–11) fatty alcohols, mainly formed de novo by the animals from non-lipid dietary precursors, esterified with various fatty acids that are often polyunsaturated fatty acids and largely of dietary, phytoplanktonic origin. Wax ester formation is maximal in copepodite stages IV and V. The lipids are elaborated not primarily for buoyancy regulation but as a source of metabolic energy during overwintering, particularly for reproduction. Large quantities of wax esters are utilised for gonadal development when stage V copepodites mature to females. Development of stage V copepodites to males is not accompanied by wax ester utilisation but males consume large amounts of these lipids in physical activity during reproduction. The role of wax esters in the life history of calanoids is illustrated with particular reference to a comparison of Calanus finmarchicus and Metridia longa in Balsfjord, northern Norway.     

Protein polymorphisms in six species of the genus Calanus

Isoelectric focusing was used to study total protein patterns and allozyme variations of six species of the genus Calanus. Each species could be characterized by total protein patterns. The results of the allozyme study indicated, in agreement with previous morphological studies, that the six Calanus species belong to three different groups: the C. finmarchicus group C. finmarchicus, C. glacialis and C. marshallae the C. helgolandicus group (C. helgolandicus, C. pacificus), and C. hyperboreus, which stands apart. There is no indication that there are more loci coding for the proteins studied in species with larger genome sizes. Nor is the degree of enzyme polymorphism related to genome size in these species.     

Integrating conventional microscopy and molecular analysis to analyse the abundance and distribution of four Calanus congeners in the North Atlantic

Analysis of the demographic structure of Calanus species inthe North Atlantic presents particular difficulties due to theoverlapping spatial distributions of four main congeneric species(Calanus finmarchicus, Calanus helgolandicus, Calanus glacialisand Calanus hyperboreus). These species have similar morphologies,making microscopic discrimination only possible between someof the species at late copepodite or adult stages. However,molecular techniques now offer the possibility of screeningsignificant numbers of specimens and unambiguously identifyingthem to species, regardless of developmental stage. Unfortunately,the processing rate of specimens by molecular methods is stilltoo low to offer a realistic alternative to microscopy for analysisof samples from large field surveys. Here, we outline and testan approach involving the use of molecular methodology in conjunctionwith conventional microscopy to assess the species assignmentof developmental stage abundances of Calanus congeners. Ourstudy has highlighted many important methodological issues.First, it cannot be assumed that the species composition ishomogeneous across the development stages; applying proportionalspecies composition of adults to morphologically undistinguishableearlier development stages can result in error. The second importantconclusion is that prosome length may be a highly unreliablediscriminator of C. finmarchicus and C. glacialis.     

Lipid composition of Calanus finmarchicus from the north sea and the arctic. A comparative study

• 1.1. Females, copepodid stages V and IV of Calanus finmarchicus were collected in Fram Strait area of the Arctic and in the northern North Sea to compare their lipid composition.
• 2.2. For the comparison only copepods were considered which contained more than 8% of 18:4 fatty acid and high amounts of wax esters to exclude seasonal and spatial variabilities and different reproductive status of females.
• 3.3. Animals are heavier in the Fram Strait area than in the North Sea with similar lipid proportion of dry weight and wax ester proportion of total lipid.
• 4.4. Only some statistical significant differences exist between the fatty acid and alcohol compositions. The levels of 16:0 acid and alcohol and of 22:1 alcohol are higher and of 20:1 acid and alcohol are lower in the North Sea than in the Arctic.

     

Reproduction of Calanus glacialis in the Laptev Sea, Arctic Ocean

Abundance and reproductive biology (gonad maturation and egg production) of the Arctic copepod Calanus glacialis were studied in the Laptev Sea and adjacent Arctic Ocean in September 1993 and from July to September 1995. Both abundance and reproductive activity were subject to strong spatial and seasonal variability, which was related to the ice cover, feeding conditions and circulation pattern. Maximum abundance of the C. glacialis population was generally confined to the outer shelf and slope with depths between 50 and 1000 m. During both cruises, highest egg production rates and largest number of young copepodite stages were observed in the eastern Laptev Sea, where the development of the C. glacialis population seems to follow the opening of the “Siberian Polynya”. In the western part, which is usually covered by pack ice, females were all immature, and no young stages were found. However, females responded quickly to a temporary opening of the ice there in 1995 and spawned. Starvation experiments showed that food-independent reproduction fuelled by internal energy resources was at least partly responsible for relatively high egg production rates at low ambient food concentrations. Egg production rates in starved females were considerably higher than those previously reported. Accepted: 14 June 2000     

The effects of temperature on metabolic rates of different life stages of Calanus glacialis in the Barents Sea

Summary The effects of temperature on rates of respiration, excretion and gut evacuation were examined for copepodite stages and adult female Calanus glacialis collected in areas close to the ice-edge during the arctic summer in the Barents Sea. The various life history stages responded differently to acute temperature changes above the in situ temperature (ca.-1.7°C). Respiration rates of early copepodite stages (C I to C IV) were very variable whereas excretion rates declined with increasing temperature in the range from-1.7°C to +5°C. Rate of oxygen consumption of adult females were independent of temperature between-1.7° and +5°C, but increased as temperature increased from +5° to 10°C. Rates of excretion of copepodite stage V and adult females were independent of temperature in the range from-1.7° to +2°C, whereas excretion of copepodite stages III and IV was negatively related to temperature in the range from-1.7° to +5°C. In C IV, C V and adult females the instantaneous rate of gut evacuation increased with increasing temperature. The different response patterns of metabolic rates of small copepodite stages, copepodite stage V and adult females C. glacialis to acute temperature changes suggest that the capacity for adjustment of ammonia excretion is better developed in C V's and adult females than in the younger life stages.     

Chick diet of dovekies Alle alle in Northwest Greenland

This paper reports the results of an investigation of dovekies (Alle alle) breeding at Hakluyt Island in the Thule District, Northwest Greenland. The paper presents a comprehensive examination of the chick diet of dovekies in the species' most important breeding area. Food delivered to nestlings consisted primarily of copepods Calanus hyperboreus and C. glacialis in the copepodite stages IV and V. Themisto libellula, other amphipods, decapods and fish larvae in small numbers were found in the chick diet. The zooplankton in the chick diet is presented both in terms of numbers and dry weight. The characteristic copepod of high-arctic waters, C. hyperboreus, was found to be the single most important species in the chick diet, contributing approximately 54% of the dry food weight. The food samples revealed low diversity, and the three species C. hyperboreus, C. glacialis and T. libellula made up 91.5% of the total zooplankton dry weight. The results from this paper are compared with previous dovekie-chick diet studies. Accepted: 14 June 2000     

Warming of Subarctic waters accelerates development of a key marine zooplankton Calanus finmarchicus

Recent observations confirm the rising temperatures of Atlantic waters transported into the Arctic Ocean via the West Spitsbergen Current (WSC). We studied the overall abundance and population structure of the North Atlantic keystone zooplankton copepod Calanus finmarchicus, which is the main prey for pelagic fish and some seabirds, in relation to selected environmental variables in this area between 2001 and 2011, when warming in the Arctic and Subarctic was particularly pronounced. Sampling within a 3‐week time window each summer demonstrated that trends in the overall abundance of C. finmarchicus varied between years, with the highest values in “extreme” years, due to high numbers of nauplii and early copepodite stages in colder years (2001, 2004, 2010), and contrary to that, the fifth copepodite stage (C5) peaking in warm years (2006, 2007, 2009). The most influential environmental variable driving C. finmarchicus life cycle was temperature, which promoted an increased C5 abundance when the temperature was above 6°C, indicating earlier spawning and/or accelerated development, and possibly leading to their development to adults later in the summer and spawning for the second time, given adequate food supply. Based on the presented high interannual and spatial variability, we hypothesize that under a warmer climate, C. finmarchicus may annually produce two generations in the southern part of the WSC, what in turn could lead to food web reorganization of important top predators, such as little auks, and induce northward migrations of fish, especially the Norwegian herring.     

Lipids in copepodite stages of <Emphasis Type="Italic">Calanus glacialis</Emphasis>

Calanus glacialis is a key herbivore in Arctic shelf seas. It feeds on primary producers and accumulates large energy reserves, primarily as wax esters. Lipid classes, fatty acids (FAs) and fatty alcohols (FAlcs) from copepodite stage II (CII) to adult females (AF) from Kongsfjorden, Svalbard, were studied in May 2004. Wax esters were the dominating lipid class in all stages, ranging from 34% of total lipids in CII to 60% in CIII–CV. Triacylglycerols increased from 8% of total lipids in CII to 23% in AF. In the earlier stages, 16:1n7 and 16:0 FAs and FAlcs were the major components of the neutral lipids, whereas the later stages were mainly characterized by the long-chained FAs and FAlcs 20:1n9 and 22:1n11. C. glacialis utilizes the short spring bloom to build up lipid reserves, mainly as wax esters, and it also incorporates effectively essential polyunsaturated FAs such as 20:5n3 and 22:6n3 in its polar lipids.     

Lipids and trophic interactions of ice fauna and pelagic zooplankton in the marginal ice zone of the Barents Sea

Gammarus wilkitzkii, Apherusa glacialis, Onismus nanseni, Onismus glacialis, Boreogadus saida, Parathemisto libellula and Calanus hyperboreus, collected in late June in the Barents Sea marginal ice zone, contained substantial levels (28–51% of the dry mass) of total lipid, the highest levels (51% and 41% respectively) being in  A. glacialis and  C. hyperboreus. Neutral lipids were present in greater amounts than polar lipids in all species. Triacylglycerols were major neutral lipids in A. glacialis, G. wilkitzkii and O. nanseni; triacylglycerols and wax esters were present in similar amounts in O. glacialis; higher levels of wax esters than triacylglycerols occurred in P. libellula; wax esters greatly exceeded triacylglycerols in C. hyperboreus, the opposite being true for B. saida. Diatom fatty acid markers were prominent in the triacylglycerols of G. wilkitzkii, O. nanseni, O. glacialis and, particularly, of  A. glacialis; 20:1(n-9) and 22:1(n-11) moieties were abundant in wax esters of G. wilkitzkii, O. nanseni, O. glacialis, P. libellula and  C. hyperboreus, and in triacylglycerols of B. saida. We deduce that  A. glacialis feeds mainly on ice algae and phytodetritus, G. wilkitzkii and the Onismus spp. feed on calanoid copepods as well as ice algae, whereas P. libellula and especially B. saida feed extensively on calanoid copepods. Accepted: 17 May 1998     

Seasonal changes in zooplankton composition in the Barents Sea, with special attention to Calanus spp. (Copepoda)

The Barents Sea is an important area with respect to fisheriesresources (i.e. capelin and cod). In May, June and August 1981zooplankton biomass was measured along a transect at 30°E,from the ice border southwards. A maximum was recorded in Atlanticwater by the end of June (>100 g wet weight m^–2 InAugust the biomass values were relatively low south of the Polarfront and increased northwards into Arctic water (–50g m^–2 The species composition was influenced by the distributionof cold Arctic water and warmer Atlantic water. The zooplanktonwas dominated by the copepods Calanus finmarchicus and C. glacialis;the former is regarded as an Atlantic species and C. glacialisas an Arctic species.     

Contrasting physiological responses to future ocean acidification among Arctic copepod populations

Widespread ocean acidification (OA) is modifying the chemistry of the global ocean, and the Arctic is recognized as the region where the changes will progress at the fastest rate. Moreover, Arctic species show lower capacity for cellular homeostasis and acid‐base regulation rendering them particularly vulnerable to OA. In the present study, we found physiological differences in OA response across geographically separated populations of the keystone Arctic copepod Calanus glacialis. In copepodites stage CIV, measured reaction norms of ingestion rate and metabolic rate showed severe reductions in ingestion and increased metabolic expenses in two populations from Svalbard (Kongsfjord and Billefjord) whereas no effects were observed in a population from the Disko Bay, West Greenland. At pH_T 7.87, which has been predicted for the Svalbard west coast by year 2100, these changes resulted in reductions in scope for growth of 19% in the Kongsfjord and a staggering 50% in the Billefjord. Interestingly, these effects were not observed in stage CV copepodites from any of the three locations. It seems that CVs may be more tolerant to OA perhaps due to a general physiological reorganization to meet low intracellular pH during hibernation. Needless to say, the observed changes in the CIV stage will have serious implications for the C. glacialis population health status and growth around Svalbard. However, OA tolerant populations such as the one in the Disko Bay could help to alleviate severe effects in C. glacialis as a species.