Nutrient conservation increases with latitude of origin in European<Emphasis Type="Italic"> Pinus sylvestris</Emphasis> populations

J. Oleksyn; P. B. Reich; R. Zytkowiak; P. Karolewski; M. G. Tjoelker

doi:10.1007/s00442-003-1265-9

Oecologia

July 2003, Volume 136, Issue 2, pp 220–235 | Cite as

Nutrient conservation increases with latitude of origin in European Pinus sylvestris populations

Authors
Authors and affiliations

J. OleksynEmail author
P. B. Reich
R. Zytkowiak
P. Karolewski
M. G. Tjoelker

Population Ecology

First Online: 17 May 2003

Received: 03 October 2002

Accepted: 16 March 2003

516 Downloads
95 Citations

Abstract

Nutrient availability varies across climatic gradients, yet intraspecific adaptation across such gradients in plant traits related to internal cycling and nutrient resorption remains poorly understood. We examined nutrient resorption among six Scots pine (Pinus sylvestris L.) populations of wide-ranging origin grown under common-garden conditions in Poland. These results were compared with mass-based needle N and P for 195 Scots pine stands throughout the species' European range. At the common site, green needle N (r²=0.81, P=0.01) and P (r²=0.58, P=0.08) concentration increased with increasing latitude of population origin. Resorption efficiency (the proportion of the leaf nutrient pool resorbed during senescence) of N and P of Scots pine populations increased with the latitude of seed origin (r²≥0.67, P≤0.05). The greater resorption efficiency of more northerly populations led to lower concentrations of N and P in senescent leaves (higher resorption proficiency) than populations originating from low latitudes. The direction of change in these traits indicates potential adaptation of populations from northern, colder habitats to more efficient internal nutrient cycling. For native Scots pine stands, results showed greater nutrient conservation in situ in cold-adapted northern populations, via extended needle longevity (from 2 to 3 years at 50°N to 7 years at 70°N), and greater resorption efficiency and proficiency, with their greater resorption efficiency and proficiency having genotypic roots demonstrated in the common-garden experiment. However, for native Scots pine stands, green needle N decreased with increasing latitude (r²=0.83, P=0.0002), and P was stable other than decreasing above 62°N. Hence, the genotypic tendency towards maintenance of higher nutrient concentrations in green foliage and effective nutrient resorption, demonstrated by northern populations in the common garden, did not entirely compensate for presumed nutrient availability limitations along the in situ latitudinal temperature gradient.

Keywords

Scots pine Nutrient resorption Climate gradient Adaptation Needle longevity

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Notes

Acknowledgements

This study was made possible by support from the State Committee for Scientific Research (Poland) grant 6 PO4F 001 17 and the U.S. National Science Foundation (IBN-9630241).

Appendix 1

Average N and P concentration (mg g^-1) in mature green foliage (3–24 months old) of different Scots pine (Pinus sylvestris L.) stands within species' continuous range in Europe

See text for additional information. (FI, Finland; SE, Sweden; RU, Russian Federation; EE, Estonia; LT, Lithuania; DE, Germany; PL, Poland; BY, Belarus; UA, Ukraine; CZ, Czech Republic; HU, Hungary. 1, Average of different crown height; 2, Average for different crown or branch position; 3, seed orchard; 4, average of multiple stands; 5, average of stands sampled in different years; 6, Stand on silicate substrate; 7, Stand on carbonate substrate)

Country and remarks	Lat. (N)	Long. (E)	Mean annual temperature (°C)	N (mg g^-1)	P (mg g^-1)	Stand or tree age (years)	Source
FI	65.87	26.12	1.1	13.6	0.9	50	Pietiläinen (1984)
FI	64.97	26.40	1.1	10.3	1.3	70	Vuorinen et al. (2000)
FI	64.79	26.79	1.0	11.8	1.5	20	Silfverberg et al. (1999)
SE	63.67	17.97	3.8	10.8	1.4	23	Örlander et al. (1990)
FI	63.51	26.59	3.1	14.0	1.6	22	Reinikainen et al. (1983)
FI	63.35	19.32	2.3	11.5	1.6	45	Silfverberg et al. (1999)
FI, 1	62.85	30.88	2.2	14.9	1.6	50	Finer (1994)
FI, 1	62.85	30.88	2.2	14.2	1.1	50	Finer (1994)
FI	62.78	30.97	2.0	11.5	1.3	100	Helmisaari (1990)
FI	62.78	30.97	2.0	10.9	1.1	15	Helmisaari (1990)
FI	62.78	30.97	2.0	11.1	1.1	35	Helmisaari (1990)
FI, 2	62.78	30.97	2.0	11.0	1.1	35	Helmisaari (1992a)
FI, 2	62.78	30.97	2.0	10.0	1.0	100	Helmisaari (1992a)
FI	62.78	30.97	2.0	11.2	1.1	15	Helmisaari (1992a)
FI	62.23	20.83	3.7	11.3	1.3	85	Finer (1992)
FI, 1	62.23	30.83	2.4	12.2	1.2	85	Finer (1994)
FI	62.20	25.23	2.7	15.4	1.1	35	Silfverberg et al. (1999)
FI	62.12	22.18	3.7	13.6	1.5	13	Reinikainen et al. (1983)
FI	62.06	25.29	2.7	12.3	1.3	65	Silfverberg et al. (1999)
FI	61.92	23.73	3.8	12.5	1.6	30	Jokela et al. (1997)
FI	61.78	20.73	3.6	13.2	1.5	20	Raitio (1987)
RU	61.70	50.90	0.2	10.5	2.1		Rusanova et al. (1977)
FI, 3	61.57	26.30	2.6	16.0	1.6		Lukina et al. (1994)
FI	61.50	24.00	3.6	11.1	1.7	20	Nuorteva et al. (1993)
FI	61.18	21.97	4.0	14.4	1.4	40	Silfverberg et al. (1999)
SE, 2	60.82	16.50	4.9	9.7		20	Troeng et al. (1982)
FI	60.44	24.29	4.3	12.1	1.1	13	Silfverberg et al. (1999)
SE, 2	59.87	18.92	5.8	14.3	1.5		Tamm (1955)
RU, 4	58.26	84.45	4.3	12.3	1.6	8	Koshelkov et al. (1973)
RU	58.26	84.45	4.3	16.1	2.2		Koschelkov (1967)
RU	58.26	84.45	4.3	13.1	1.8		Koschelkov (1967)
RU, 4	58.26	84.45	4.3	13.2	1.8		Koschelkov (1967)
RU, 4	58.26	84.45	4.3	11.8	1.6		Koschelkov (1967)
RU, 4	58.26	84.45	4.3	11.5	1.5		Koschelkov (1967)
RU, 4	58.26	84.45	4.3	10.6	1.4		Koschelkov (1967)
RU, 4	58.26	84.45	4.3	9.7	1.4		Koschelkov (1967)
EE, 4	58.00	27.00	5.5	16.4	1.7		Porgasaar (1977)
UK, 2	57.58	-3.87	8.0	13.3	1.5	64	Wright et al. (1958)
UK	57.58	-3.87	8.0	10.6	1.2	18	Wright et al. (1958)
UK	57.58	-3.87	8.0	14.6	1.9	28	Wright et al. (1958)
UK	57.58	-3.87	8.0	14.2	1.5	64	Wright et al. (1958)
SE, 5	57.17	14.83	5.8	12.7	1.5	30	Sikström (1997)
LT	54.88		6.5	13.1	1.8		Stefan et al. (1997)
LT, 4	54.88		6.5	11.9	1.2		Skoudene (1997)
DE, 5	53.68	13.85	8.5	16.2	1.7	5–15	Heinsdorf (1973)
RU	53.33	34.20	5.3	11.0	1.7		Krasnikov (1976)
PL	53.20	23.37	6.7	15.2	1.4	15	Oleksyn et al. (unpublished data)
DE, 5	53.12	13.50	8.7	14.6	1.6	70	Trillmich et al. (1982)
DE	52.92	13.87	8.7	10.9	1.3	7	Heinsdorf (1976c)
DE	52.92	13.87	8.7	11.0	1.3	14	Heinsdorf (1976c)
DE	52.92	13.87	8.7	13.0	1.6	33	Heinsdorf (1976c)
DE	52.92	13.87	8.7	14.4	1.7	47	Heinsdorf (1976c)
DE	52.92	13.87	8.7	14.5	1.6	85	Heinsdorf (1976c)
DE	52.87	13.40	8.5	16.5	1.7	5	Heinsdorf (1966)
DE	52.87	13.40	8.5	16.6	1.6	6	Heinsdorf (1966)
DE, 5	52.87	13.75	8.7	16.2	1.6	42–46	Heinsdorf et al. (1976)
DE	52.83	13.88	8.9	14.9	1.4	35	Ritter et al. (1978)
DE	52.83	13.88	8.3	14.3	1.4	55	Riedel (1986)
DE	52.83	13.88	8.3	13.1	1.4	40	Hippeli (1976)
DE	52.66	13.73	8.5	15.8	1.6	5	Heinsdorf (1966)
DE	52.66	13.73	8.5	14.9	1.6	6	Heinsdorf (1966)
DE, 4	52.41	13.50	8.5	15.1		57	Tölle et al. (1970)
DE	52.36	14.07	8.5	13.1	1.5	5	Heinsdorf (1966)
DE	52.36	14.07	8.5	15.2	1.7	6	Heinsdorf (1966)
DE	52.36	14.07	8.5	13.1	1.4	6	Heinsdorf (1967)
DE	52.36	14.07	8.5	11.2	1.2	6	Heinsdorf (1967)
PL	52.35	20.30	7.5	15.9	1.7		Kurczynska et al. (1997)
PL, 5	52.30	23.85	6.2	13.8	1.4		Unpublished data of the Forest Research Institute in Warsaw
PL	52.25	17.07	7.7	11.5	1.3	11–15	Oleksyn et al. (unpublished data)
PL	52.25	17.07	7.7	12.6	1.8	29	Oleksyn et al. (unpublished data)
BY, 4	52.25	31.00	7.1	15.1	1.5	8–10	Morozov et al. (1981)
DE, 5	52.23	12.17	8.5	12.3	1.3	5–15	Heinsdorf (1973)
DE, 5	52.23	12.17	8.5	12.0		15–19	Heinsdorf (1978)
DE	52.23	12.17	8.5	12.7	1.3	5	Heinsdorf (1976b)
DE, 5	52.22	12.85	8.5	13.4	1.4	5–11	Heinsdorf (1973)
DE	52.20	12.77	8.5	12.6	1.3	5–6	Heinsdorf (1966)
DE	52.20	12.77	8.5	10.6	1.1	6	Heinsdorf (1967)
PL	52.17	20.90	7.5	16.7	1.7		Kurczynska et al. (1997)
DE, 5	52.05	13.60	8.5	11.0	1.2	6–16	Heinsdorf (1973)
DE	52.00	13.00	8.5	13.5	1.5	5–6	Heinsdorf (1973)
DE, 5	51.95	13.23	8.5	11.8	1.3	6–15	Heinsdorf (1973)
DE	51.95	13.23	8.5	12.3	1.5	5–6	Heinsdorf (1966)
DE	51.95	13.23	8.5	10.1	1.2	6	Heinsdorf (1967)
DE	51.90	13.22	8.5	12.3	1.3	5–14	Heinsdorf (1973)
DE	51.90	13.22	8.5	13.4		15–18	Heinsdorf (1978)
DE	51.87	13.43	8.5	17.1	1.8	5–6	Heinsdorf (1966)
DE	51.87	13.43	8.5	13.3	1.5	5–6	Heinsdorf (1967)
PL	51.80	14.97	8.0	12.0	1.4	60	Rzeznik et al. (1968)
DE	51.62	14.02	8.4	13.2	1.3	6–7	Heinsdorf (1976a)
DE	51.55	14.10	8.4	18.0	1.3	5–6	Heinsdorf (1976a)
DE	51.50	13.80	8.4	11.6	1.1	6–7	Heinsdorf (1976a)
DE	51.41	14.25	7.9	13.7	1.4	7	Heinsdorf (1987)
DE	51.40	12.88	8.5	17.1	2.0	37	Czerney et al. (1969)
DE	51.40	12.88	8.5	18.3	1.7	39	Czerney et al. (1969)
DE	51.40	12.88	8.5	17.8	1.8	52	Czerney et al. (1969)
DE	51.40	12.88	8.5	18.1	1.8	56	Czerney et al. (1969)
DE	51.40	12.88	8.5	17.8	1.8	63	Czerney et al. (1969)
DE	51.40	12.88	8.5	17.9	1.8	77	Czerney et al. (1969)
DE	51.40	12.88	8.5	16.1	1.7	88	Czerney et al. (1969)
DE	51.40	12.88	8.5	16.0	1.6	97	Czerney et al. (1969)
DE	51.40	12.88	8.5	17.6	1.4	108	Czerney et al. (1969)
DE	51.40	12.88	8.5	16.7	1.7	118	Czerney et al. (1969)
DE	51.40	13.90	9.1	13.5	1.5	50	Höhne et al. (1970)
DE, 5	51.40	14.00	8.5	10.7	1.1	7–11	Heinsdorf (1982)
DE	51.37	14.52	8.4	14.0	1.4	5–6	Heinsdorf (1976a)
DE	51.32	14.33	9.1	13.8	1.5	9	Fiedler et al. (1967)
DE	51.32	14.33	9.1	14.8	1.3	15	Fiedler et al. (1967)
DE	51.32	14.33	9.1	14.8	1.3	29	Fiedler et al. (1967)
DE	51.32	14.33	9.1	15.5	1.2	41	Fiedler et al. (1967)
DE	51.32	14.33	9.1	16.3	1.2	52	Fiedler et al. (1967)
DE	51.32	14.33	9.1	15.3	1.5	78	Fiedler et al. (1967)
DE	51.32	14.33	9.1	14.8	1.5	89	Fiedler et al. (1967)
DE	51.32	14.33	9.1	12.7	1.3	9	Fiedler et al. (1967)
DE	51.32	14.33	9.1	12.8	1.2	15	Fiedler et al. (1967)
DE	51.32	14.33	9.1	14.8	1.2	29	Fiedler et al. (1967)
DE	51.32	14.33	9.1	15.3	1.0	41	Fiedler et al. (1967)
DE	51.32	14.33	9.1	17.4	1.1	52	Fiedler et al. (1967)
DE	51.32	14.33	9.1	14.4	1.3	78	Fiedler et al. (1967)
DE	51.32	14.33	9.1	14.5	1.4	98	Fiedler et al. (1967)
DE	51.30	14.33	8.6	14.7	1.6	105	Leube et al. (1975)
DE, 5	51.30	14.45	8.5	13.3	1.5	6–16	Fiedler et al. (1970)
DE	51.28	12.98	7.3	16.1	1.6	24	Czerney et al. (1969)
DE	51.28	12.98	7.3	17.5	1.8	33	Czerney et al. (1969)
DE	51.28	12.98	7.3	19.1	1.6	37	Czerney et al. (1969)
DE	51.28	12.98	7.3	17.7	1.7	41	Czerney et al. (1969)
DE	51.28	12.98	7.3	17.6	1.6	47	Czerney et al. (1969)
DE	51.28	12.98	7.3	19.8	1.4	44	Czerney et al. (1969)
DE	51.28	12.98	7.3	21.3	1.4	47	Czerney et al. (1969)
DE	51.28	12.98	7.3	17.6	1.6	50	Czerney et al. (1969)
DE	51.28	12.98	7.3	17.6	1.4	60	Czerney et al. (1969)
DE	51.28	12.98	7.3	19.3	1.7	61	Czerney et al. (1969)
DE	51.28	12.98	7.3	17.0	1.7	65	Czerney et al. (1969)
DE	51.28	12.98	7.3	19.8	1.5	65	Czerney et al. (1969)
DE	51.28	12.98	7.3	19.9	1.5	69	Czerney et al. (1969)
DE	51.28	12.98	7.3	17.6	1.4	77	Czerney et al. (1969)
DE	51.28	12.98	7.3	17.0	1.4	80	Czerney et al. (1969)
DE	51.28	12.98	7.3	19.2	1.3	82	Czerney et al. (1969)
DE	51.28	12.98	7.3	17.6	1.4	100	Czerney et al. (1969)
DE	51.28	12.98	7.3	15.6	1.1	87	Czerney et al. (1969)
DE	51.28	12.98	7.3	18.0	1.4	97	Czerney et al. (1969)
DE	51.28	12.98	7.3	17.8	1.5	118	Czerney et al. (1969)
DE	51.28	12.98	7.3	16.5	1.1	111	Czerney et al. (1969)
DE	51.28	12.98	7.3	16.9	1.6	121	Czerney et al. (1969)
DE	51.28	12.98	7.3	17.6	1.2	122	Czerney et al. (1969)
DE	51.28	12.98	7.3	17.6	1.6	10	Czerney et al. (1969)
DE	51.28	14.42	9.1	13.3	1.5	97	Fiedler et al. (1967)
DE	51.27	14.33	9.1	15.0	1.4	107	Fiedler et al. (1967)
DE	51.27	14.33	9.1	12.5	1.4	10	Fiedler et al. (1967)
DE	51.27	14.33	9.1	14.4	1.4	17	Fiedler et al. (1967)
DE	51.27	14.33	9.1	12.4	1.5	27	Fiedler et al. (1967)
DE	51.27	14.33	9.1	12.9	1.5	41	Fiedler et al. (1967)
DE	51.27	14.33	9.1	12.8	1.5	50	Fiedler et al. (1967)
DE	51.27	14.33	9.1	13.2	1.5	61	Fiedler et al. (1967)
DE	51.27	14.33	9.1	12.7	1.5	81	Fiedler et al. (1967)
DE	51.27	14.33	9.1	13.7	1.4	110	Fiedler et al. (1967)
DE	51.27	14.33	9.1	15.2	1.3	107	Fiedler et al. (1967)
DE	51.27	14.33	9.1	11.1	1.2	10	Fiedler et al. (1967)
DE	51.27	14.33	9.1	12.4	1.4	17	Fiedler et al. (1967)
DE	51.27	14.33	9.1	12.4	1.6	27	Fiedler et al. (1967)
DE	51.27	14.33	9.1	12.1	1.5	41	Fiedler et al. (1967)
DE	51.27	14.33	9.1	11.4	1.4	50	Fiedler et al. (1967)
DE	51.27	14.33	9.1	12.4	1.5	61	Fiedler et al. (1967)
DE	51.27	14.33	9.1	13.8	1.6	81	Fiedler et al. (1967)
DE	51.27	14.33	9.1	14.4	1.5	110	Fiedler et al. (1967)
PL	51.23	18.17	7.5	13.2	0.9	29	Oleksyn et al. (unpublished data)
DE	50.90	11.60	8.7	15.0	1.6	12–13	Fiedler et al. (1987)
DE	50.77	11.63	8.0	13.5	0.8	77	Fiedler et al. (1969)
DE	50.77	11.63	8.0	14.5	1.0	58	Fiedler et al. (1969)
DE	50.77	11.63	8.0	15.8	1.0	72	Fiedler et al. (1969)
UA	50.55	48.35	7.1	16.3	1.4	40	Patlai (1973)
UA	50.55	48.35	7.1	15.6	1.7	5	Patlai (1973)
PL	50.03	20.37	7.7	12.4	1.5	15	Oleksyn et al. (unpublished data)
DE	50.00	8.69	9.8	17.9	1.2	100	Busch et al. (1997)
DE	50.00	12.30	7.1	14.3	1.6	111	Sauter (1991)
DE	50.00	12.30	7.1	14.2	1.6	80	Sauter (1991)
DE	50.00	12.30	7.1	15.0	1.6	81	Sauter (1991)
RU	49.93	36.28	6.9	16.3	1.4	40	Patlai (1973)
DE	49.77	11.93	8.0	13.3	1.4	86	Sauter (1991)
DE	49.75	11.55	8.0	14.0	1.5	70	Sauter (1991)
DE	49.65	14.08	7.9	16.0	1.5	15	Oleksyn et al. (unpublished data)
DE, 4	49.40	11.00	8.6	15.2	1.4		Gulder et al. (1993)
DE	49.26	12.32	8.0	13.5	1.6	62	Sauter (1991)
DE	49.26	12.32	8.0	13.4	1.5	70	Sauter (1991)
DE	49.26	12.32	8.0	13.9	1.5	100	Sauter (1991)
DE	49.20	12.03	8.0	14.3	1.4	105	Sauter (1991)
CZ, 6	49.10	17.57	8.9	14.7	1.4	85	Heinze (1996)
CZ, 7	49.10	17.57	8.9	17.4	1.6	85	Heinze (1996)
DE	49.03	10.98	7.6	13.2	1.5	89	Sauter (1991)
DE, 4, 5	49.00	11.00	7.6	14.1	1.4	40	Wehrmann (1961)
HU, 6	47.68	16.16	10.3	14.0	1.7	85	Heinze (1996)
HU, 7	47.68	16.16	10.3	14.5	1.4	85	Heinze (1996)
HU, 6	47.55	17.75	10.4	20.7	1.2	85	Heinze (1996)
HU, 6	47.45	17.03	9.7	17.8	1.4	85	Heinze (1996)
HU, 7	46.77	17.25	10.0	15.1	1.5	85	Heinze (1996)
HU, 6	46.68	19.68	10.8	16.3	1.6	10	Heinze (1996)

Appendix 2

Needle longevity (years) in different Scots pine (Pinus sylvestris L.) stands within species' continuous range in Europe

See text for additional information. [FI, Finland; SE, Sweden; RU, Russian Federation; EE, Estonia; PL, Poland; UA, Ukraine; GB, Great Britain. Lat, latitude (in decimal values); Long, longitude; Alt, altitude; M.a.t., mean annual temperature. Growing season length was calculated as the number of days with mean temperature ≥5°C]

Location (country)	Lat (N)	Long (E)	Alt (m)	M.a.t. (°C)	Growing season (days)	Needle longevity (years)	Reference
Murmansk (RU)	68.97	33.04	46	0.2	126	7.0	Pravdin (1969)
Jukkasjärvi (FI)	68.00	20.61	320	−1.5	111	7.4	Sylven (1916/1917)
Lokka (FI)	67.77	27.75	255	−0.7	126	5.7	Jalkanen et al. (1995)
Ainijärvi (FI)	67.75	29.48	265	−0.9	125	5.8	Jalkanen et al. (1995)
Särkijärvi (FI)	67.72	23.83	280	−1.3	128	6.2	Jalkanen et al. (1995)
Nuttio (FI)	67.67	25.97	220	−0.7	126	6.0	Jalkanen et al. (1995)
Aska (FI)	67.27	26.72	175	−0.7	126	6.0	Jalkanen et al. (1995)
Sätsi (FI)	67.25	29.22	225	−0.8	127	6.4	Jalkanen et al. (1995)
Pajala (SE)	67.22	23.40	168	0.5	137	5.8	Sylven (1916/1917)
Kalix (SE)	66.85	23.13	11	0.2	141	5.6	Sylven (1916/1917)
Ängesa (SE)	66.73	22.30	182	0.8	142	6.5	Sylven (1916/1917)
Raneträsk (SE)	66.70	20.00	263	−0.7	126	7.2	Sylven (1916/1917)
Pärlälven (SE)	66.70	18.40		−1.2	127	7.8	Sylven (1916/1917)
Kulveikonvaara (FI)	66.70	27.82	255	−0.7	133	5.5	Jalkanen et al. (1995)
Jockmock (SE)	66.59	19.89	255	−1.2	127	8.0	Sylven (1916/1917)
Kaarnijärvi (FI)	66.45	26.80	200	0.4	138	6.0	Jalkanen et al. (1995)
Storbacken (SE)	66.38	20.71	43	−1.2	126	7.4	Sylven (1916/1917)
Tennilä (FI)	66.38	26.48	135	0.4	137	5.2	Jalkanen et al. (1995)
Arjeplog (SE)	66.03	17.80	430	0.6	134	6.2	Sylven (1916/1917)
Malmesjaur (SE)	66.03	19.17		−1.2	127	6.4	Sylven (1916/1917)
Ranea (SE)	66.00	22.30	11	1.3	142	5.4	Sylven (1916/1917)
Vargisa (SE)	65.88	20.32	104	−1.2	127	7.2	Sylven (1916/1917)
Tornaea (SE)	65.83	24.13	3	1.0	138	5.2	Sylven (1916/1917)
Sivakkavaara (FI)	65.80	24.67	35	1.3	138	4.5	Jalkanen et al. (1995)
Arvidsjaur (SE)	65.58	19.17	378	−1.2	127	6.8	Sylven (1916/1917)
Övre Byske (SE)	65.40	19.72		−1.2	127	6.6	Sylven (1916/1917)
Älvsby (SE)	65.30	18.55	17	0.3	138	5.6	Sylven (1916/1917)
Stensele (SE)	65.09	17.20	327	0.3	118	6.2	Sylven (1916/1917)
Jörn (SE)	65.03	20.15	296	−1.3	118	6.4	Sylven (1916/1917)
Norsjö (SE)	64.92	19.48	300	3.1	159	5.6	Sylven (1916/1917)
Frostviken (SE)	64.65	13.77	337	2.0	148	5.4	Sylven (1916/1917)
Vilhelmina (SE)	64.62	16.65	341	1.1	144	5.6	Sylven (1916/1917)
S. Lycksele (SE)	64.60	18.67	223	1.2	144	5.6	Sylven (1916/1917)
Burträsk (SE)	64.52	20.65	81	2.4	160	6.0	Sylven (1916/1917)
Tasjö (SE)	64.22	15.90	248	0.9	144	5.7	Sylven (1916/1917)
Degerfors (SE)	64.20	19.72	217	2.5	159	5.0	Sylven (1916/1917)
Asele (SE)	64.17	17.33	312	1.0	144	5.4	Sylven (1916/1917)
Fredrika (SE)	64.08	18.40	293	1.1	144	5.8	Sylven (1916/1917)
Anundsjö (SE)	63.43	18.15	123	3.5	162	6.4	Sylven (1916/1917)
Are (SE)	63.40	13.07	371	1.7	156	6.0	Sylven (1916/1917)
Bräcke (SE)	63.38	14.00	295	2.5	156	5.8	Sylven (1916/1917)
Hallen (SE)	63.37	13.45	386	1.7	139	5.4	Sylven (1916/1917)
Östersund (SE)	63.18	14.50	367	2.5	154	6.0	Sylven (1916/1917)
Junsele (SE)	62.83	16.90	195	3.4	168	5.9	Sylven (1916/1917)
Härnösand (SE)	62.60	18.00	8	3.5	161	5.5	Sylven (1916/1917)
Medelpad (SE)	62.53	17.39	4	3.4	163	5.4	Sylven (1916/1917)
Rätan (SE)	62.47	14.55	351	2.4	159	5.6	Sylven (1916/1917)
Hede (SE)	62.41	13.50	439	0.5	140	8.0	Sylven (1916/1917)
Västra (SE)	62.40	14.00		2.4	159	5.0	Sylven (1916/1917)
Kaltila (FI)	62.00	24.10	110	2.6	160	4.3	Jalkanen et al. (1995)
Norra (SE)	61.83	14.00		2.4	158	5.0	Sylven (1916/1917)
Karelya (RU)	61.82	34.27	110	2.5	159	6.0	Bobkova et al. (1999)
Ranta-Halola (FI)	61.80	29.33	90	2.8	163	4.4	Jalkanen et al. (1995)
Särna (SE)	61.70	13.10	437	1.5	150	6.0	Sylven (1916/1917)
Vesijako (FI)	61.33	28.05	130	3.6	172	4.0	Jalkanen et al. (1995)
Österdalarna (SE)	61.28	14.55		0.9	175	5.2	Sylven (1916/1917)
Piikajärvi (FI)	61.27	22.18	45	3.6	170	4.1	Jalkanen et al. (1995)
Gästrikland (SE)	61.20	16.80		4.5	179	4.1	Sylven (1916/1917)
Transtrand (SE)	61.08	13.32	368	1.5	150	6.8	Sylven (1916/1917)
Ojajoki (FI)	60.67	24.35	130	4.5	173	3.9	Jalkanen et al. (1995)
Grönbo (SE)	60.60	15.47	215	4.9	183	3.0	Sylven (1916/1917)
Halkivaha (FI)	60.60	24.40	120	4.5	173	3.8	Jalkanen et al. (1995)
Kaksoiskivi (FI)	60.60	24.52	120	4.5	173	4.0	Jalkanen et al. (1995)
Torpparinmäki (FI)	60.53	26.45	30	4.5	175	4.2	Jalkanen et al. (1995)
Ruotsinkylä (FI)	60.35	24.95	60	4.3	171	4.1	Jalkanen et al. (1995)
Kopparberg (SE)	60.25	14.98	229	4.9	183	4.0	Sylven (1916/1917)
N Roslag (SE)	60.25	17.40		5.8	189	3.0	Sylven (1916/1917)
Örbyhus (SE)	60.23	17.70	21	5.9	189	3.5	Sylven (1916/1917)
Västerdalarna (SE)	60.17	14.50		5.8	183	4.8	Sylven (1916/1917)
Solböle (FI)	60.03	23.03	40	3.8	171	4.0	Jalkanen et al. (1995)
Kloten (SE)	59.90	15.28	263	5.4	180	3.2	Sylven (1916/1917)
Arvika (SE)	59.67	12.63	74	5.4	188	3.8	Sylven (1916/1917)
Västeras (SE)	59.58	16.60	6	5.9	194	3.0	Sylven (1916/1917)
Enköping (SE)	59.50	17.07	10	6.0	194	3.2	Sylven (1916/1917)
Älvdal (SE)	59.40	13.50		5.6	188	4.4	Sylven (1916/1917)
Karlstad (SE)	59.40	13.50	55	5.6	188	3.8	Sylven (1916/1917)
Örebro (SE)	59.30	15.20	33	5.4	185	3.0	Sylven (1916/1917)
Askersund (SE)	59.00	14.80	175	6.1	183	3.0	Sylven (1916/1917)
Tartu (EE)	58.30	26.70		5.5	191	2.5	Tullus (1991)
Northern (GB)	57.86	−4.79	203	6.7	196	3.4	Steven et al. (1959)
Wester Ross (GB)	57.55	−5.42	210	6.8	196	3.3	Steven et al. (1959)
Strath Glass (GB)	57.31	−4.91	301	6.1	191	3.2	Steven et al. (1959)
Speyside (GB)	57.16	−3.72	369	6.1	190	3.9	Steven et al. (1959)
Deeside (GB)	57.00	−3.10	325	6.4	190	3.8	Steven et al. (1959)
Great Glen (GB)	56.99	−5.11	305	7.0	238	3.5	Steven et al. (1959)
Moskovsk. obl. (RU)	56.80	37.00	100	3.8	178	6.0	Bobkova et al. (1999)
Rannoch (GB)	56.63	-4.33	294	6.7	238	3.9	Steven et al. (1959)
Southern (GB)	56.48	-4.79	252	7.2	235	3.0	Steven et al. (1959)
Moskovsk. obl. (RU)	55.80	37.60	156	4.2	178	5.0	Bobkova et al. (1999)
Bryansk. obl. (RU)	53.33	34.22	162	4.9	185	2.0	Bobkova et al. (1999)
Kornik (PL)	52.25	17.07	70	7.7	220	2.6	Reich et al. (1996a)
Lvov (UA)	49.80	24.00	325	7.3	210	3.0	Pravdin (1969)
Lvov (UA)	49.80	24.00	325	7.3	210	3.0	Pravdin (1969)
Lvov (UA)	49.80	24.00	325	7.3	210	3.0	Pravdin (1969)
Lvov (UA)	49.80	24.00	325	7.3	210	3.0	Pravdin (1969)
Lvov (UA)	49.80	24.00	325	7.3	210	3.0	Pravdin (1969)

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Authors and Affiliations

J. Oleksyn
- 1
- 2
Email author
P. B. Reich
- 1
R. Zytkowiak
- 2
P. Karolewski
- 2
M. G. Tjoelker
- 1
- 3

1.Department of Forest ResourcesUniversity of MinnesotaSt. PaulUSA
2.Institute of DendrologyPolish Academy of Sciences KórnikPoland
3.Department of Forest ScienceTexas A & M UniversityCollege StationUSA