Lipids and life strategy of Arctic Calanus

Figures & data

Figure 1.  Time-related plankton blooms in the Arctic Oceans (modified after Zenkevitch 1963).

Figure 2.  The current system of the Arctic Seas.

Figure 3.  Generalized seasonal migration and stage development of Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus. Upper and lower line delineates the general depth of the population.

Figure 4.  Life span of Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus in Arctic waters. The numeration generalizes a gradient of geographic regions from the southern Barents Sea to the North Pole as indicated in Figure 1.

Table I. Life span of Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus in Arctic waters.

Species	Life span (years)	Area	References
Calanus finmarchicus	1	East Greenland Barents Sea Norwegian Sea Foxe Basin West Greenland Current Davis Strait Barents Sea	With 1915 Jaschnov 1939; Manteiphel 1941; Kamshilov, 1955 Ostvedt 1955; Lie 1968 Grainger 1965 MacLellan 1967; Madsen et al. 2008 Sekerak et al. 1976 Falk-Petersen et al. 1999
	2	Canadian archipelago	Longhurst et al. 1984
Calanus glacialis	1	West Greenland Current Davis Strait Nova Scotia Fram Strait	MacLellan 1967; Sekerak et al. 1976 Huntley et al. 1983 Runge et al. 1986 Smith 1990
	1–2	Barents Sea	Falk-Petersen et al. 1999
	2	Arctic Ocean Resolute area Barents Sea White Sea	Grainger 1965 Conover & Huntley 1991 Jaschnov 1961, 1970; Tande et al. 1985; Slagstad & Tande 1990; Pedersen et al. 1995 Prygunkova 1974
	2–3	Barrow Strait	Conover et al. 1991
	3	White Sea	Kosobokova 1999
Calanus hyperboreus	1	Davis Strait	Sekerak et al. 1976
	1–2	East Greenland Norwegian fjords Norwegian Sea Canadian archipelago Fram Strait	Ussing 1938 Wiborg 1954; Matthews et al. 1978; Østvedt 1955 Grainger 1959 Smith 1990
	2 2	North of Iceland Canadian archipelago	Gislason & Astthorsson 1998 Longhurst et al. 1984
	2–3	Arctic Ocean West Spitsbergen Current	Harding 1966; Dawson 1978 Hirche 1997
	3	Arctic Ocean Resolute area	Harding 1966; Dawson 1978; Pavshtiks 1983 Conover & Huntley 1991
	3–4	Canadian Greenland Sea Gyre	Conover 1988; Conover et al. 1991; Conover & Siferd 1993 Hirche 1997
	3–5	Barents Sea	Falk-Petersen et al. 1999

Table II. Over-wintering stages and spawning periods of Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus in Arctic waters.

Species	Area	Stages	Spawning	References
C. finmarchicus	Canadian archipelago Barents Sea Greenland Sea (Northeast Water Polynya) Greenland Sea North of Iceland Greenland Sea	CIII CIV–CV CIV–CV CV CV Female	April–May April–May August–October	Longhurst et al. 1984 Pedersen et al. 1995 Hirche et al. 1994 Hirche 1991 Hirche & Kwasniewski 1997 Gislason & Astthorsson 1998 Hirche 1989b
C. glacialis	Baffin Bay Canadian archipelago Barents Sea Basin White Sea White Sea Greenland Sea (Northeast Greenland (Young Sound) Water Polynya) West Greenland Davis Strait Fram Strait North of Iceland David Strait Barents Sea (central part) Barents Sea (around 78^o N) Barents Sea (central part) Polar basin Resolute area Laptev Sea (east part)	CIII CIII CIII–CIV CIV CIV CIV and CV CIV CV–CVI CV–CVI CV–CVI	May–June May–June February–March June March March–April Late winter February March–April April May–June May–October May–July June–July	Grainger 1963 Longhurst et al. 1984; Tande et al. 1985; Pavshtiks 1977; Kosobokova 1986 Prygunkova 1968 Kosobokova & Pertzova 1990 Kosobokova 1999 Hirche 1991; Hirche et al. 1994; Nielsen et al. 2007 MacLellan 1967 Huntley et al. 1983 Smith 1990 Gislason & Astthorsson 1998 Sekerak et al. 1976; Tande 1991 Pedersen et al. 1995 Falk-Petersen et al. 1999; Tande et al. 1985; Slagstad & Tande 1990; Johnson 1963 Conover & Huntley 1991 Michel et al. 2006 Kosobokova & Hirche 2001
C. hyperboreus	Canadian archipelago North Water (Canada) Greenland Sea Greenland Sea Off Norway North of Iceland Arctic Ocean Foxe Basin Resolute Passage Greenland Sea	CIII CIV, CV CIII CIII and CIV CIII–CIV CIV CV	Winter October–March January–February Late winter January–May May–June January–April	Longhurst et al. 1984 Ringuette et al. 2002 Hirche et al. 1994; Hirche & Niehoff 1996; Hirche 1997 Wiborg 1954 Gislason & Astthorsson 1998 Johnson 1963 Grainger 1965 Conover 1988; Conover et al. 1991 Conover & Huntley 1991 Hirche & Kwasniewski, 1997

Table III. Prosome length (mm), wet mass (WW), dry mass (DW) and lipid mass (LW) as mg ind^–1, and % total lipid of DW in different developmental stages of Calanus finmarchicus, C. glacialis and C. hyperboreus. Length from Hirche et al. (1994), WW, DW and % total lipid of DW from Scott et al. (2000). DM of stages I–II are calculated after Mauchlene (1989).

Species	Stage	Length (mm)	Wet mass (mg ind^−1)	Dry mass (mg ind^−1)	Lipid mass (mg ind^−1)	% lipid
Calanus finmarchicus	I	0.68		0.0016
	II	0.93		0.004
	III	1.31		0.013
	IV	1.6–2–2	0.28	0.04	0.02	53
	V	2.0–3.0	0.56	0.13	0.04	34
	F	2.4–3.2	1.08	0.26	0.08	31
Calanus glacialis	II	1.31		0.019
	III	1.81		0.04
	IV	2.2–2–8	0.39	0.09	0.05	56
	V	3.0–4.0	2.38	0.62	0.38	61
	F	3.2–4.6	2.60	0.64	0.45	70
Calanus hyperboreus	I	1.04		0.05
	II	1.63		0.032
	III	2.47		0.142
	IV	3.4–4.0	1.91	0.51	0.28	54
	V	4.5–6.0	6.44	1.58	1.03	65
	F	6.2–7.0	12.09	2.92	1.81	62

Figure 5.  Seasonal variation in the specific gravity (buoyancy) of Calanus finmarchicus and Calanus hyperboreus (after Koegeler et al. 1987).

Table IV. Fatty acid composition of polar lipids of Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus from the Arctic.

	Calanus finmarchicus				Calanus glacialis					Calanus hyperboreus
Location	NBS^1		FSW^3	HB^11	NBS^1	KGF^8			FSW^3	NBS^1	FSW^3	AO^9
Year	1986		89 + 90	1975	1986	2004			1989 + 90	1986	1989 + 90
Season	spring		sum.	winter	spring	spring			sum.	spring	sum.	sum.	aut.	winter
Stage	CV	AF	AF	CV	AF	CIV	CV	AF	AF	AF	AF	CIV–V	CIV–V	CIV–V
Replicate	n=1	n=1	n=2		n=1	n=6	n=6	n=3	n=4	n=1	n=3	n=1	n=1	n=2
14:0	4.4	3.9	3.3	4.6	6.1	6.3	6.5	4.9	5.2	7.3	4.0	3.0	2.0	3.0
15:0	0.6	0.4	nd	0.8	0.4	0.4	0.4	0.4	1.8	nd	0.8	nd	nd	nd
16:0	33.2	27.0	25.8	21.5	31.8	10.9	9.5	9.3	25.8	29.1	25.5	12.0	14.0	15.0
16:1(n–9)	nd	nd	nd	0.8	1.6	0.5	0.3	0.3	nd	nd	nd	nd	nd	nd
16:1(n–7)	5.4	2.7	1.1	2.8	4.0	7.2	9.6	6.9	4.6	6.3	2.5	3.0	3.0	5.0
16:1(n–5)	1.6	0.6	0.2	1.2	0.8	0.3	0.3	0.1	0.1	1.0	0.9	nd	nd	nd
18:0	3.1	1.8	3.6	2.2	1.2	1.3	1.1	1.1	2.3	2.3	2.3	1.0	nd	4.0
18:1(n–9)	6.4	4.1	2.5	5.1	3.7	2.6	2.5	3.2	6.4	4.1	5.1	11.0	11.0	13.0
18:1(n–7)	2.4	2.2	1.0	2.2	4.4	1.1	1.0	1.3	3.7	3.5	1.4	nd	nd	nd
18PUFAs	4.4	5.6	4.6	0.0	2.6	6.7	5.4	5.2	2.2	1.5	3.1	2.0	3.0	2.0
20:1(n–9)	1.4	0.9	0.2	4.0	1.9	6.0	6.9	5.9	0.9	0.8	0.6	nd	nd	nd
20:1(n–7)	nd	nd	nd	0.3	0.2	0.1	0.1	0.3	nd	0.1	nd	nd	nd	nd
20:4(n–6)	0.2	0.3	0.2	0.1	0.2	0.2	0.3	0.2	nd	0.2	0.2	nd	nd	nd
20:4(n–3)	0.5	0.7	0.2	0.2	0.3	0.0	0.0	0.0	nd	0.5	nd	nd	nd	nd
22:1(n–11)	0.7	0.1	0.2	6.7	nd	3.3	3.9	4.2	nd	0.1	1.2	nd	nd	nd
22:1(n–9)	0.5	0.2	nd	0.9	0.2	0.4	0.5	0.6	nd	nd	nd	nd	nd	nd
20:5(n–3)	12.6	22.5	19.2	13.2	21.7	30.0	26.9	33.0	16.5	21.4	18.6	19.0	18.0	18.0
24:1(n–9)	2.7	1.4	nd	1.57	1.2	0.4	0.4	0.1	nd	2.1	nd	nd	nd	nd
22PUFAs	10.4	20.6	37.6	24.2	13.0	14.0	13.7	17.0	30.5	11.7	33.8	43.0	39.0	36.5
Saturated	41.3	33.1	32.7	29.1	39.5	18.9	17.4	15.6	35.1	38.7	32.6	16.0	16.0	22.0
Monounsat	21.1	12.2	5.2	25.5	18.0	21.8	25.6	22.9	15.7	18.0	11.7	14.0	14.0	18.0
PUFAs	28.1	49.7	61.8	37.7	37.8	50.9	46.2	55.3	49.2	35.3	55.7	64.0	60.0	56.5
20:5(n–3) + 16:1(n–7)	18.0	25.2	20.3	16.0	25.7	37.2	36.5	39.9	21.1	27.7	21.1	22.0	21.0	23.0
16:1(n–7)/16:0	0.2	0.1	0.0	0.1	0.1	0.7	1.0	0.7	0.2	0.2	0.1	0.3	0.2	0.3
Northern Barents Sea (NBS), Kongsfjorden (KGF), Fram Strait East (FSE), Fram Strait West (FSW), Arctic Ocean (AO), Hudson Bay (HB) and North Water Polynia (NWP). The numbers from 1–11 refer to the original work: 1 = Falk-Petersen 1986 (unpublished), 2 = Kattner et al. 1989, 3 = Alberts et al. 1996, 4= Scott et al. 2002, 5 = Falk-Petersen/Norwegian Polar Institute database (unpublished), 6 = Søreide et al. 2007, 7 = Leu (unpublished), 8 = Leu et al. 2006, 9 = Lee 1974, 10 = Stevens et al. 2004, 11 = Mayzaud (unpublished). * = neutral lipid, ** = total lipid.

Table V. Fatty alcohol composition of wax esters of Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus from the Arctic.

	Calanus finmarchicus
Location	NBS^1		KGF^4		KGF^5		FSE^2			FSE^6			FSW^2		FSW^3	AO^6	HB^11
Year	1986		1997		1998		1984			2003			1984		1989 + 90	2003	1975
Season	spring		autumn		autumn		summer			spring	autumn	winter	summer		summer	autumn	winter
Stage	CV	AF	CV	AF	CV	AF	CIV	CV	AF	CV	CV	CV	AF	CV	AF	CV	CV
Replicates	n=1	n=1	n=13	n=12	n=6	n=6	n=4	n=3	n=3	n=3	n=3	n=6	n=1	n=1	n=2	n=3
14:0	0.2	0.1	1.9	2.1	3.1	1.7	3.0	1.9	1.2	2.9	2.1	2.1	0.4	1.1	3.9	1.5	2.0
14:1	0.9	0.6	nd	nd	0.1	0.1	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	0.1
15:0	nd	nd	0.3	0.3	0.2	0.2	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	0.4
16:0	7.5	6.0	8.4	8.0	9.6	7.3	12.4	8.8	7.5	8.4	8.1	11.3	3.0	6.6	14.6	4.4	11.8
16:1(n–7)	3.3	2.7	6.2	6.8	6.2	6.8	2.5	1.6	1.4	5.1	2.4	2.1	0.7	1.8	3.4	1.9	1.9
18 total	8.4	7.2	5.9	5.6	4.8	5.4	5.2	4.1	3.4	1.5	2.7	3.4	1.9	4.0	0.0	1.1	7.5
20:1(n–9)	42.2	48.5	35.3	35.6	39.9	41.5	28.2	42.1	40.6	38.9	41.0	43.7	39.8	41.0	39.3	36.8	33.4
20:1(n–7)	nd	nd	0.7	2.3	0.5	1.4	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	0.6
22:1(n–11)	24.6	25.0	39.0	36.5	33.1	31.6	38.4	41.4	45.7	39.6	38.9	33.0	54.3	45.2	38.8	50.0	32.4
22:1(n–9)	nd	nd	0.3	0.8	0.6	2.8	nd	nd	nd	3.6	4.8	4.4	nd	nd	nd	4.3	1.7
20:1 + 22:1	66.8	73.5	75.4	75.2	74.1	77.4	66.5	83.5	86.3	82.1	84.6	81.0	94.1	86.2	78.1	91.0	68.1
22:1(n–11)/ 20:1(n–9)	0.6	0.5	1.1	1.0	0.8	0.8	1.4	1.0	1.1	1.0	0.9	0.8	1.4	1.1	1.0	1.4	1.0
Northern Barents Sea (NBS), Kongsfjorden (KGF), Fram Strait East (FSE), Fram Strait West (FSW), Arctic Ocean (AO), Hudson Bay (HB) and North Water Polynia (NWP). The numbers from 1–11 refer to the original work: 1 = Falk-Petersen 1986 (unpublished), 2 = Kattner et al. 1989, 3 = Alberts et al. 1996, 4= Scott et al. 2002, 5 = Falk-Petersen/Norwegian Polar Institute database (unpublished), 6 = Søreide et al. 2007, 7 = Leu (unpublished), 8 = Leu et al. 2006, 9 = Lee 1974, 10 = Stevens et al. 2004, 11 = Mayzaud (unpublished). * = neutral lipid, ** = total lipid.

Table

	Calanus glacialis
Location	NBS^1	NBS^6*		KGF^4		KGF^5		KGF^8		FSE^6*		FSW^3	AO^6*
Year	1986	2004	2003	1997		1998		2004		2003		1989 + 90	2003
Season	spring	spring	autumn	autumn		autumn		spring		autumn	winter	summer	autumn
Stage	AF	CV	CV	CV	AF	CV	AF	CV	AF	CV	CV	AF	CV
Replicates	n=1	n=3	n=9	n=9	n=8	n=5	n=6	n=6	n=3	n=7	n=6	n=4	n=3
14:0	0.7	5.7	3.2	2.1	1.5	3.4	2.6	4.3	2.8	3.1	3.5	2.1	3.3
14:1	1.3	nd	nd	0.2	0.2	nd	nd	nd	nd	nd	nd	nd	nd
15:0	nd	nd	nd	0.3	0.3	0.3	0.2	nd	nd	nd	nd	nd	nd
16:0	5.1	13.1	10.3	9.4	7.7	9.6	8.2	12.7	12.1	8.8	13.2	9.3	9.2
16:1(n–7)	3.0	5.4	4.4	4.2	3.8	5.8	4.3	8.6	8.6	4.1	4.7	5.3	2.7
18 total	4.6	2.1	1.8	5.9	5.2	4.0	3.8	1.7	2.3	1.6	2.6	0.0	1.8
20:1(n–9)	63.4	47.4	46.4	41.3	45.2	38.7	36.5	45.5	45.1	47.3	48.5	58.4	45.6
20:1(n–7)	1.0	nd	nd	0.7	1.9	2.3	1.3	nd	nd	nd	nd	nd	nd
22:1(n–11)	16.1	21.8	24.4	31.1	28.4	32.7	41.4	24.7	23.2	29.8	22.5	25.0	31.6
22:1 n–9	1.3	4.4	5.2	2.5	3.3	1.9	1.4	4.0	6.0	5.2	4.9	nd	5.8
20:1 + 22:1	81.8	73.7	76.0	75.6	78.8	75.6	80.7	74.1	74.2	82.3	75.9	83.4	83.0
22:1(n–11)/20:1(n–9)	0.3	0.5	0.5	0.8	0.6	0.8	1.1	0.5	0.5	0.6	0.5	0.4	0.7

Table

	Calanus hyperboreus
Location	NBS^1	NBS^6*		KGF^4		KGF^5		FSE^2**			FSE^6*		FSW^2**			FSW^3	AO^6*		AO^10
Year	1986	2004	2003	1997		1998		1984			2003		1984			1989 + 90	2003
Season	spr.	spr.	aut.	autumn		autumn		summer			spr.	wint.	summer			summer	autumn		sum.	aut.	wint.
Stage	AF	CV	CV	CV	AF	CV	AF	CIV	CV	AF	CV	AF	CIV	CV	AF	AF	CV	AF	CIV–V	CIV–V	CIV–V
Replicates	n=1	n=3	n=9	n=12	n=14	n=6	n=3	n=5	n=3	n=3	n=6	n=3	n=2	n=2	n=3	n=3	n=6	n=6	n=1	n=1	n=2
14:0	0.5	8.1	5.2	3.3	2.6	4.6	5.2	6.4	4.4	2.0	4.9	2.1	3.4	3.1	2.1	4.4	4.1	2.0	1.0	2.0	3.0
14:1	4.0	nd	nd	nd	nd	0.0	0.0	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd
15:0	10.7	nd	nd	0.3	0.3	0.2	1.2	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd
16:0	1.6	13.0	10.2	8.8	6.5	9.4	4.2	7.8	7.5	4.1	7.5	4.7	11.1	7.9	5.7	11.2	8.0	3.6	5.0	8.0	33.5
16:1(n–7)	nd	1.4	2.6	3.5	2.0	3.3	2.2	0.5	0.3	0.1	1.7	0.6	2.2	1.1	1.2	1.6	1.3	0.6	2.0	4.0	5.0
18 total	1.8	0.5	0.5	3.3	2.3	2.5	1.4	1.3	1.2	0.4	0.7	0.7	0.7	0.8	0.7	0.0	0.6	0.4	5.0	8.0	2.0
20:1(n–9)	33.6	33.1	29.5	28.1	27.7	26.4	28.7	28.8	32.0	31.3	29.4	24.5	27.4	27.4	34.3	27.8	29.0	27.6	35.0	41.0	23.0
20:1(n–7)	3.5	nd	nd	2.4	3.1	0.0	4.9	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd
22:1(n–11)	29.3	34.1	39.8	47.3	52.8	53.5	47.1	55.0	54.7	61.6	44.5	50.7	55.0	59.8	55.9	55.0	44.0	49.2	48.0	34.0	30.5
22:1 n–	8.4	6.0	8.4	1.5	1.3	0.1	5.1	nd	nd	nd	8.3	10.6	nd	nd	nd	nd	9.3	13.2	nd	nd	nd
20:1 + 22:1	74.8	73.2	77.7	79.3	84.9	80.0	85.9	83.7	86.7	92.9	82.2	85.8	82.4	87.2	90.2	82.8	82.3	90.0	83.0	75.0	53.5
22:1(n–11)/20:1(n–9)	0.9	1.0	1.4	1.7	1.9	2.0	1.6	1.9	1.7	2.0	1.5	2.1	2.0	2.2	1.6	2.0	1.5	1.8	1.4	0.8	1.3

Table VI. Fatty acid composition of wax esters of Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus from the Arctic.

	Calanus finmarchicus
Location	NBS^1		NBS^6*	KGF^5				FSE^2**			FSE^5		FSE^6*			FSW^3	FSW^2**		FSW^5		AO^5		AO^6*	HB^11
Year	1986		2004	1998		1997		1984			1998		2003	2003	2003	89 + 90	1984		1998		1998		2003	1975
Season	spring		spring	autumn		autumn		summer			autumn		spr.	aut.	wint.	summer	summer		autumn		autumn		autumn	winter
Stage	CV	AF	CV	CV	AF	CV	AF	CIV	CV	AF	CV	AF	CV	CV	CV	AF	CV	AF	CV	AF	CV	AF	CV	CV
Replicates	n=1	n=1	n=3	n=6	n=6	n=11	n=11	n=3	n=3	n=4	n=2	n=3	n=3	n=3	n=6	n=2	n=1	n=1	n=3	n=3	n=6	n=6	n=3
14:0	25.2	36.0	16.8	15.6	13.6	9.4	8.8	12.1	14.7	12.7	9.5	9.4	12.0	18.5	23.4	26.3	18.0	21.2	23.3	14.4	17.9	11.4	18.6	10.0
15:0	1.5	1.3	0.6	1.3	2.0	0.7	0.6	nd	nd	nd	2.2	1.3	0.4	0.9	1.2	0.7	nd	nd	0.9	0.7	1.1	0.8	0.9	0.8
16:0	11.2	8.6	7.8	9.8	9.4	7.8	6.3	8.6	8.2	10.1	5.3	7.1	6.2	10.0	11.9	9.8	9.3	8.8	10.7	8.2	9.3	6.7	8.6	11.9
16:1(n–9)	nd	nd	0.6	nd	nd	nd	nd	nd	nd	nd	nd	nd	0.4	0.7	0.5	nd	nd	nd	nd	nd	nd	nd	0.4	1.1
16:1(n–7)	9.9	8.9	10.1	18.7	21.1	20.9	25.2	4.3	4.4	5.2	19.2	12.1	7.5	6.5	8.2	6.7	6.5	7.3	12.3	18.4	18.9	28.2	8.8	9.7
16:1(n–5)	1.7	1.5	0.1	nd	nd	0.1	0.2	nd	nd	nd	nd	nd	0.5	0.7	0.1	0.9	nd	nd	nd	nd	nd	nd	0.8	2.1
16PUFA	1.3	0.7	13.1	2.7	2.9	4.9	4.3	1.8	2.3	1.7	5.1	4.1	9.0	2.5	2.1	2.0	2.3	1.2	2.6	5.6	3.5	3.2	2.3	1.1
18:0	0.6	0.6	0.8	0.6	0.6	0.3	0.5	nd	nd	nd	0.2	0.7	0.5	0.7	0.7	0.9	nd	nd	0.6	0.3	0.5	0.3	0.6	0.8
18:1(n–9)	6.4	4.3	2.8	4.3	5.3	2.1	3.2	4.3	4.4	5.3	2.7	3.7	2.5	4.5	6.4	5.3	5.3	5.7	4.1	3.7	5.0	3.5	5.3	7.6
18:1(n–7)	0.7	0.5	0.5	1.1	2.4	1.5	1.6	nd	nd	nd	1.3	0.7	0.6	0.4	0.5	0.3	nd	nd	0.5	1.2	0.8	1.3	0.5	1.5
18:4(n–3)	0.8	0.0	7.0	8.2	5.1	1.8	3.7	16.7	16.2	9.0	5.5	4.2	9.2	12.6	11.1	13.7	9.3	1.6	1.6	4.6	1.4	1.8	5.6	3.2
18PUFA (other)	3.3	1.0	1.0	2.8	1.8	2.7	1.9	0.0	0.0	0.0	1.9	0.4	0.9	3.1	3.7	2.7	0.0	0.0	2.3	3.1	2.3	1.0	2.2	3.9
20:0	0.1	0.2	0.2	nd	nd	0.0	0.1	nd	nd	nd	nd	nd	0.6	0.2	0.2	nd	nd	nd	nd	nd	nd	nd	0.4	0.3
20:1(n–11)	nd	nd	0.5	nd	nd	0.3	0.3	nd	nd	nd	nd	nd	1.1	0.4	0.7	nd	nd	nd	nd	nd	nd	nd	1.2	0.3
20:1(n–9)	10.1	13.3	9.3	10.4	11.9	15.3	13.8	5.2	7.6	8.3	17.8	17.7	10.0	9.3	8.5	7.8	9.6	13.4	14.1	13.9	13.5	19.4	9.5	13.6
20:1(n–7)	0.3	0.3	0.1	0.0	0.8	2.6	1.2	nd	nd	nd	0.6	0.2	1.3	0.1	0.1	0.9	nd	nd	0.0	0.0	0.0	1.1	0.1	0.7
20:4(n–6)	0.2	0.1	0.1	nd	nd	0.9	0.4	nd	nd	nd	nd	nd	1.2	0.4	0.3	0.4	nd	nd	nd	nd	nd	nd	1.6	0.2
20:4(n–3)	nd	0.1	0.0	0.9	0.6	0.6	0.9	nd	nd	nd	0.8	0.1	3.2	0.0	0.0	0.5	nd	nd	0.7	0.6	0.5	0.7	4.0	0.5
22:1(n–11)	12.1	15.3	6.5	10.4	9.5	8.6	10.9	6.7	9.2	8.2	11.6	11.0	13.5	10.4	7.4	7.0	12.0	15.7	15.4	10.9	14.9	11.2	15.4	15.7
22:1(n–9)	1.1	1.7	0.7	0.4	1.4	1.9	0.9	nd	nd	nd	1.1	0.0	1.2	0.9	0.6	0.2	nd	nd	0.0	0.0	0.2	1.2	1.4	1.3
22:1(n–7)	nd	nd	0.1	nd	nd	nd	nd	nd	nd	nd	nd	nd	0.3	0.1	0.1	nd	nd	nd	nd	nd	nd	nd	0.4	0.2
20:5(n–3)	0.9	nd	16.0	8.7	8.0	11.3	11.5	16.2	12.9	14.6	10.8	15.5	12.3	9.9	6.7	11.4	9.8	7.4	8.0	11.8	6.8	5.5	5.9	6.5
24:1(n–9)	0.2	1.0	0.3	nd	nd	nd	nd	nd	nd	nd	nd	nd	0.4	0.4	0.3	nd	nd	nd	nd	nd	nd	nd	0.6	0.3
22:6(n–3)	0.0	0.0	1.4	1.3	1.2	1.2	1.7	13.0	9.4	13.8	2.3	1.6	1.9	3.2	1.5	2.2	7.7	9.8	0.8	0.5	0.8	0.8	1.1	2.6
22PUFAs (other)	0.0	0.0	0.3	0.7	0.6	0.6	0.7	0.0	0.0	0.0	0.7	9.7	0.1	0.6	0.3	0.2	0.0	0.0	0.4	0.6	0.4	0.8	0.0	0.1
Saturated	38.6	46.7	26.2	27.2	25.5	18.3	16.2	20.7	22.9	22.8	17.2	18.5	19.6	30.2	37.4	37.7	27.3	30.0	35.5	23.7	28.8	19.2	29.2	23.9
Monounsat	42.5	46.8	31.5	45.2	52.3	53.1	57.2	20.5	25.7	27.0	54.3	45.5	39.3	34.5	33.4	29.1	33.4	42.1	46.3	48.1	53.4	65.9	44.5	54.0
PUFAs	6.5	1.9	39.0	25.3	20.2	24.0	25.0	47.7	40.8	39.1	27.1	35.6	37.9	32.1	25.8	33.1	29.1	20.0	16.4	26.9	15.8	13.8	22.8	18.1
20:5(n–3) + 16PUFA + 16:1(n–7)	12.1	9.6	39.2	30.0	32.0	37.1	40.9	22.3	19.6	21.5	35.1	31.7	28.9	18.9	17.0	20.1	18.6	15.9	23.0	35.8	29.2	36.9	17.1	17.3
16:1(n–7)/16:0	0.9	1.0	1.3	1.9	2.3	2.7	4.0	0.5	0.5	0.5	3.6	1.7	1.2	0.7	0.7	0.7	0.7	0.8	1.2	2.2	2.0	4.2	1.0	0.8

Table

	Calanus glacialis
Location	NBS^1	NBS^6*	KGF^7**		KGF^8*		KGF^4		KGF^5		FSE^5		FSE^6*		FSW^3	FSW^5		AO^5		AO^6*
Year	1986	2003	2003		2004		1997		1998		1998		2003		89 + 90	1998	1998	1998	1998	2003
Season	spring	autumn	spring		spring		autumn		autumn		autumn		aut.	win.	summer	autumn		autumn		autumn
Stage	AF	CV	AF	CIV	AF	CV	AF	CV	AF	CV	AF	CV	CV	CV	AF	AF	CV	AF	CV	CV
Replicates	n=1	n=9	n=3	n=6	n=3	n=6	n=7	n=8	n=6	n=6	n=1	n=3	n=7	n=6	n=4	n=3	n=1	n=6	n=6	n=3
14:0	14.0	9.9	4.8	5.8	8.1	8.9	5.6	7.9	10.5	8.4	9.3	14.4	8.7	16.4	13.1	11.9	11.1	8.6	8.1	8.0
15:0	0.3	0.6	0.7	0.5	0.3	0.5	0.3	0.6	1.6	1.2	0.0	0.5	0.4	0.9	0.3	0.7	0.5	0.4	0.4	0.4
16:0	2.2	6.5	17.3	8.8	5.7	8.4	4.2	6.6	5.4	5.3	4.3	7.4	5.5	9.1	6.1	5.4	5.6	4.0	4.7	5.1
16:1(n–9)	nd	0.6	0.2	0.3	0.8	0.4	nd	nd	nd	nd	nd	nd	0.5	0.6	nd	nd	nd	nd	nd	0.6
16:1(n–7)	13.5	14.5	1.5	10.9	19.6	16.3	23.2	17.8	25.0	25.0	18.6	20.0	12.1	16.5	32.9	23.8	21.4	34.7	26.0	8.5
16:1(n–5)	2.0	0.7	0.8	1.0	0.9	0.5	nd	nd	nd	nd	nd	nd	0.6	0.1	0.3	nd	nd	nd	nd	0.7
16PUFA	1.2	5.9	1.1	10.0	2.6	4.0	3.6	2.4	5.0	4.7	6.8	5.1	8.0	3.0	1.2	3.7	5.2	3.5	4.4	4.2
18:0	0.4	0.5	1.5	1.7	1.1	1.0	0.3	0.5	0.3	0.2	0.0	0.0	0.5	0.7	nd	0.1	0.2	0.4	0.2	0.3
18:1(n–9)	2.4	4.0	2.2	3.1	4.6	3.4	3.6	5.6	3.6	3.9	3.3	3.5	3.5	6.1	5.5	4.8	4.1	3.0	3.6	3.8
18:1(n–7)	0.7	0.8	0.2	1.2	1.4	1.1	1.2	0.9	1.1	0.9	1.3	1.3	0.7	0.9	1.1	0.8	0.7	1.5	1.0	0.8
18:4(n–3)	0.0	7.1	1.7	4.6	7.0	5.8	3.5	6.5	4.3	5.8	7.6	5.8	8.2	8.3	0.5	3.4	4.3	2.0	4.2	7.8
18PUFAs (other)	0.6	2.5	2.2	2.4	2.2	1.5	1.3	2.6	2.9	1.6	2.0	1.9	1.3	3.1	1.3	2.2	1.6	1.5	1.8	6.9
20:0	0.1	0.1	0.2	0.5	0.3	0.2	nd	nd	nd	nd	nd	nd	0.3	0.2	0.1	nd	nd	nd	nd	0.6
20:1(n–11)	nd	0.1	0.3	0.9	0.4	0.2	nd	nd	nd	nd	nd	nd	0.6	0.4	nd	nd	nd	nd	nd	0.9
20:1(n–9)	34.5	15.8	8.0	14.4	18.7	13.2	20.5	16.6	15.6	16.9	18.7	13.0	14.8	13.2	23.0	18.0	17.9	20.3	19.5	13.8
20:1(n–7)	1.0	0.2	0.4	1.1	1.0	0.3	1.0	0.3	0.2	0.0	0.0	0.0	0.2	0.2	1.0	0.0	0.0	1.0	0.0	1.5
20:4(n–6)	0.3	0.2	0.4	0.2	0.2	0.2	0.2	0.3	nd	nd	nd	nd	1.2	0.2	nd	nd	nd	nd	nd	1.7
20:4(n–3)	nd	0.0	0.7	1.7	nd	nd	0.7	0.9	0.6	0.6	0.7	0.7	1.5	0.0	nd	0.6	0.6	0.6	0.6	2.7
22:1(n–11)	16.8	8.0	3.6	5.7	6.5	6.4	10.3	10.7	8.9	10.4	12.5	12.1	9.7	6.7	8.3	10.0	11.0	8.0	11.0	9.8
22:1(n–9)	4.9	1.2	1.1	1.1	1.8	0.8	1.8	0.9	1.0	0.2	0.0	0.0	1.3	0.9	2.0	0.0	0.0	1.8	1.3	1.4
22:1(n–7)	nd	0.1	0.2	0.5	0.4	0.1	0.4	0.1	nd	nd	nd	nd	0.3	0.1	nd	nd	nd	nd	nd	0.5
20:5(n–3)	0.6	13.9	21.7	12.2	5.8	16.3	12.9	11.7	10.4	11.6	11.4	11.2	14.0	6.9	2.7	12.3	13.2	6.4	9.4	12.6
24:1(n–9)	1.0	0.3	nd	nd	0.7	0.4	nd	nd	nd	nd	nd	nd	0.3	0.3	nd	nd	nd	nd	nd	0.4
22:6(n–3)	0.0	1.4	19.3	5.1	2.4	6.7	1.5	2.4	1.1	1.5	1.7	1.7	2.2	1.1	0.8	0.9	0.9	0.6	1.7	2.7
22PUFAs (other)	0.0	2.1	0.9	0.6	0.7	0.9	0.8	1.0	0.9	0.9	0.7	0.6	0.3	0.3	0.0	0.5	0.5	0.5	0.9	1.2
Saturated	17.0	17.7	24.6	17.3	15.5	18.9	10.4	15.5	17.8	15.0	13.6	22.3	15.3	27.3	19.6	18.1	17.3	13.4	13.4	14.4
Monounsat	76.8	46.4	18.5	40.2	56.8	43.0	62.2	52.8	55.4	57.3	54.5	49.9	44.7	46.1	74.1	57.2	55.2	70.3	62.3	42.6
PUFAs	2.7	33.1	48.0	36.9	21.0	35.4	24.5	27.8	25.2	26.6	31.0	27.0	36.6	23.0	6.5	23.5	26.2	15.0	23.0	39.8
20:5(n–3) + 16PUFA+ 16:1(n–7)	15.3	34.3	24.2	33.1	28.1	36.6	39.7	31.9	40.4	41.3	36.9	36.3	34.1	26.5	36.8	39.7	39.8	44.6	39.8	25.3
16:1(n–7)/16:0	6.1	2.2	0.1	1.2	3.4	1.9	5.5	2.7	4.6	4.7	4.3	2.7	2.2	1.8	5.4	4.4	3.8	8.7	5.5	1.7

Table

	Calanus hyperboreus
Location	NBS^1	NBS^6*	NBS^6*	KGF^5		KGF^4		FSE^2**			FSE^5		FSE^6*	FSW^3	FSW^2**			FSW^5		AO^9			AO^5		AO^6*			NWP^10
Year	1986	2004	2003	1998		1997		1988			1998		2003	89 + 90	1984			1998					1998		2003	2003		1999
Season	spr.	spr.	aut.	autumn		autumn		summer			autumn		winter	summer	summer			autumn		sum.	aut.	win.	autumn		spring	autumn		autumn
Stage	AF	CV	CV	CV	AF	CV	AF	CIV	CV	AF	CV	AF	AF	AF	CIV	CV	AF	CV	AF	CIV-V	CIV-V	CIV-V	CV	AF	CV	CV	AF	CV
Replicates	n=1	n=3	n=9	n=6	n=4	n=14	n=13	n=3	n=3	n=3	n=3	n=4	n=3	n=3	n=2	n=2	n=3	n=2	n=4	n=1	n=1	n=2	n=1	n=1	n=6	n = 7	n = 7	n = 27
14:0	4.6	4.9	3.6	4.6	4.2	3.3	3.3	3.4	2.6	2.9	2.9	3.6	4.5	6.4	6.6	6.7	3.6	0.9	3.1	2.0	5.0	6.0	6.1	3.9	2.2	2,9	3,0	3,1
15:0	0.2	0.1	0.2	0.4	0.7	0.2	0.2	nd	nd	nd	2.0	0.1	0.3	nd	nd	nd	nd	0.3	0.2	nd	nd	nd	0.6	0.0	0.0	0,1	0,2	0,1
16:0	2.5	2.7	2.8	3.4	2.8	2.3	2.8	3.3	2.6	3.1	2.0	2.7	3.4	5.8	8.0	5.4	3.9	3.2	2.3	3.0	4.0	5.0	2.2	8.3	1.5	2,0	2,2	3,1
16:1(n–9)	nd	0.2	0.6	nd	nd	nd	nd	nd	nd	nd	nd	nd	0.8	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	0.4	0,6	0,1	nd
16:1(n–7)	20.9	7.2	12.6	nd	nd	21.7	19.3	2.6	2.8	3.7	nd	nd	12.1	11.7	12.8	14.2	16.9	nd	nd	24.0	22.0	30.0	nd	nd	6.1	6,7	7,1	17,9
16:1(n–5)	1.2	0.0	0.4	nd	nd	nd	nd	nd	nd	nd	nd	nd	0.3	0.7	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	0.3	0,4	0,1	0,3
16PUFA	3.0	5.2	6.4	4.2	4.9	4.8	3.1	1.7	1.6	2.1	3.4	3.0	1.9	3.5	3.1	1.5	2.6	7.5	6.0	6.0	5.0	0.0	3.5	0.0	5.4	3,4	2,6	7,4
18:0	0.6	0.4	0.5	0.4	0.1	0.3	2.1	nd	nd	nd	0.1	0.5	0.6	0.6	nd	nd	nd	0.0	0.1	2.0	nd	nd	0.0	0.0	0.6	0,2	0,3	0,2
18:1(n–9)	4.1	3.4	2.7	3.6	2.6	2.6	0.3	4.0	4.1	3.8	3.9	3.7	3.7	5.8	6.7	6.8	5.0	1.7	2.2	3.0	8.0	7.5	6.7	45.5	2.5	3,1	2,2	2,0
18:1(n–7)	1.0	1.0	1.2	1.4	1.4	1.5	2.2	nd	nd	nd	1.3	1.4	1.3	1.6	nd	nd	nd	1.1	1.2	nd	nd	nd	1.4	1.4	0.8	0,9	0,9	1,3
18:4(n–3)	1.3	31.6	12.8	4.7	10.8	6.9	0.7	19.5	23.3	4.4	11.1	16.1	4.3	6.2	3.1	1.8	2.9	3.2	3.8	0.0	6.0	0.0	1.1	7.1	22.5	9,4	10,6	3,2
18PUFAs (other)	0.3	30.6	11.6	3.3	9.4	2.0	0.9	0.0	0.0	15.1	9.8	0.0	2.7	5.2	0.0	0.0	0.2	2.0	0.3	14.0	0.0	2.0	13.4	0.0	2.3	5,1	1,8	1,7
20:0	nd	0.1	0.1	nd	nd	nd	nd	nd	nd	nd	nd	nd	0.7	0.8	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	0.5	0,4	0,5	nd
20:1(n–11)	nd	1.0	0.4	nd	nd	nd	4.9	nd	nd	nd	nd	nd	1.8	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	2.1	1,4	1,7	nd
20:1(n–9)	27.4	11.3	13.7	14.1	18.3	15.3	nd	25.9	15.7	15.2	12.9	16.8	14.9	19.0	4.1	18.0	19.5	7.0	12.2	2.0	22.0	19.0	15.7	10.0	11.5	13,9	13,4	7,1
20:1(n–7)	1.6	0.6	1.1	1.0	2.7	1.8	17.4	nd	nd	nd	0.2	1.2	1.4	1.5	nd	nd	nd	1.1	1.9	nd	nd	nd	0.0	0.0	0.7	1,0	1,5	1,2
20:4(n–6)	0.5	0.1	0.2	nd	nd	nd	nd	nd	nd	nd	nd	nd	1.0	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	1.3	1,5	1,4	nd
20:4(n–3)	0.2	0.0	0.0	1.1	0.8	0.9	nd	nd	nd	nd	1.1	1.4	5.9	0.7	nd	nd	nd	0.6	0.7	5.0	nd	13.0	1.1	0.0	3.9	3,7	5,3	nd
22:1(n–11)	11.1	7.8	10.9	11.9	11.6	15.6	12.0	8.3	16.7	18.2	10.0	11.7	18.1	17.3	12.8	18.4	17.7	5.4	7.1	nd	15.0	7.5	23.5	11.3	12.7	15,2	15,0	6,8
22:1(n–9)	7.6	1.4	2.8	1.9	3.1	0.7	14.1	nd	nd	nd	0.7	2.7	5.0	3.2	nd	nd	nd	0.0	1.5	nd	nd	nd	0.0	2.6	2.6	3,9	6,2	1,3
22:1(n–7)	nd	0.1	0.2	nd	nd	nd	3.7	nd	nd	nd	nd	nd	1.3	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	0.7	0,8	1,3	0,2
20:5(n–3)	4.1	8.0	14.5	9.6	10.8	13.1	0.8	12.6	9.2	10.4	15.9	14.6	5.6	7.0	10.4	8.2	12.6	24.5	19.4	35.0	12.0	23.0	5.6	4.0	9.5	13,0	11,5	27,1
24:1(n–9)	0.6	0.1	0.3	nd	nd	nd	nd	nd	nd	nd	nd	nd	0.7	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	nd	0.1	0,3	0,4	nd
22:6(n–3)	0.0	3.7	5.2	2.1	4.8	3.2	0.1	11.0	10.9	6.4	7.2	9.6	3.0	2.4	7.5	10.7	3.4	1.9	2.6	3.0	1.0	4.0	2.5	3.1	5.8	5,6	6,1	5,0
22PUFAs (other)	0.0	0.4	1.3	3.8	0.0	1.2	0.9	0.0	0.0	4.6	5.1	0.0	0.0	0.5	0.0	0.0	4.1	2.1	0.7	0.0	3.0	0.0	1.6	0.0	0.3	0,7	0,0	1,8
Saturated	7.9	8.2	7.2	8.7	7.8	6.1	8.4	6.7	5.3	6.1	6.9	6.9	9.4	13.6	14.5	12.1	7.5	4.4	5.7	7.0	9.0	11.0	8.9	12.2	4.8	5,6	6,1	6,6
Monounsat	75.5	34.1	47.0	33.9	39.7	59.2	73.8	40.7	39.3	40.9	29.2	37.5	61.6	60.8	36.3	57.4	59.2	16.4	26.2	29.0	67.0	64.0	47.3	70.8	40.5	48,1	50,0	38,0
PUFAs	10.4	52.5	42.4	35.9	26.1	32.1	6.5	25.1	45.0	43.0	53.7	43.2	24.4	25.5	19.7	22.2	25.8	41.9	33.5	61.0	23.0	41.0	28.9	7.6	51.0	42,4	39,2	46,3
20:5(n–3) + 16PUFA + 16:1(n–7)	28.0	20.4	33.5	13.8	15.6	39.6	23.2	16.9	13.6	16.2	19.4	17.6	19.6	22.2	26.2	23.9	32.1	32.1	25.4	65.0	39.0	53.0	9.1	4.0	21.0	23,1	21,1	52,4
16:1(n–7)/16:0	8.4	2.7	4.6	0.0	0.0	9.4	7.0	0.8	1.1	1.2	0.0	0.0	3.6	2.0	1.6	2.7	4.4	0.0	0.0	8.0	5.5	6.0	0.0	0.0	4.0	3,4	3,3	5,8

Figure 6.  Hierarchical cluster analysis of fatty alcohol composition for C. hyperboreus, C. glacialis and Calanus finmarchicus collected in different sectors of the European Arctic Ocean. Symbols: Ch = C. hyperboreus, Cg = C. glacialis, Cf = C. finmarchicus

Figure 7.  PLS-DA for the three main species of Calanus sampled in the Northern Barents Sea and Kongsfjorden based on the fatty acid composition of wax esters. Projection on the plane defines by the first two discriminant functions (DS1 = 45.9% and DS2 = 33.2% total inertia). Factor loading for the fatty acid descriptors. Italic: centroid of the factorial group. Ind = indices for each Calanus species (see text).

Figure 8.  PLS-DA for the three main species of Calanus sampled in the eastern Fram Strait and Arctic Ocean based on the fatty acid composition of wax esters. Projection on the plane defines by the first two discriminant functions (DS1 = 37.7% and DS2 = 32.3% total inertia). Factor loading for the fatty acid descriptors. Italic: centroid of the factorial group. Ind = indices for each Calanus species (see text).

Figure 9.  PLS-DA for the three main species of Calanus sampled in the western Fram Strait based on the fatty acid composition of wax esters. Projection on the plane defines by the first two discriminant functions (DS1 = 36% and DS2 = 32% total inertia). Factor loading for the fatty acid descriptors. Italic: centroid of the factorial group. Ind = indices for each Calanus species (see text).

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