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Variation in spring and autumn frost tolerance amongprovenances of Russian larches (Larix Mill.)Thröstur Eysteinsson a , Lars Karlman b c , Anders Fries d , Owe Martinsson c & BrynjarSkúlason ea Iceland Forest Service , Egilsstadir, Icelandb Department of Ecology and Management , Swedish University of Agricultural Sciences ,Umeå, Swedenc Jämtlands Institute for Rural Development , Bispgården, Swedend Department of Forest Genetics and Plant Physiology , Swedish University of AgriculturalSciences , Umeå, Swedene North Iceland Regional Afforestation Project , Akureyri, IcelandPublished online: 13 May 2009.

To cite this article: Thröstur Eysteinsson , Lars Karlman , Anders Fries , Owe Martinsson & Brynjar Skúlason (2009)Variation in spring and autumn frost tolerance among provenances of Russian larches (Larix Mill.), Scandinavian Journal ofForest Research, 24:2, 100-110, DOI: 10.1080/02827580902773470

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ORIGINAL ARTICLE

Variation in spring and autumn frost tolerance among provenancesof Russian larches (Larix Mill.)

THROSTUR EYSTEINSSON1, LARS KARLMAN2,3, ANDERS FRIES4,

OWE MARTINSSON3 & BRYNJAR SKULASON5

1Iceland Forest Service, Egilsstadir, Iceland, 2Department of Ecology and Management, Swedish University of Agricultural

Sciences, Umea, Sweden, 3Jamtlands Institute for Rural Development, Bispgarden, Sweden, 4Department of Forest Genetics

and Plant Physiology, Swedish University of Agricultural Sciences, Umea, Sweden, and 5North Iceland Regional Afforestation

Project, Akureyri, Iceland

AbstractSpring and autumn frost tolerance was measured using material from a range-wide (50�678 N, 38�1588 E) provenance trialof four Russian larch species (Larix sukaczewii Dyl., L. sibirica Ledeb., L. gmelinii Rupr. and L. cajanderi Mayr.) growing innorthern Sweden. Shoots were collected in early May and late September and frozen at �8, �12, �16 and �208C.Cambial damage was assessed visually after development under ideal conditions for 2 weeks. Differences in frost damageamong provenances were highly significant in both spring and autumn. Autumn frost damage was significantly correlatedwith provenance latitude and longitude and spring frost damage was significantly correlated with provenance longitude butnot latitude. Frost damage was not correlated with provenance elevation. North-western provenances showed the leastdamage and far-eastern provenances the greatest damage in both spring and autumn. A possible explanation for less springfrost damage to north-western provenances is adaptation to maritime conditions in proximity to the Barents Sea, which isoften ice free in late winter. This would counteract early loss of frost tolerance and bud flushing if warm spells occurred inlate winter. North-eastern Siberian provenances did not show similar adaptation and may exhibit increased spring frostdamage if global warming eventually results in the Arctic Ocean north of Siberia becoming ice free in late winter.

Keywords: Adaptation, frost tolerance, larch, Larix, provenance trials.

Introduction

Larches (Larix Mill.) have their greatest area of

distribution in Russia, covering an area of roughly

280 million ha, about 37% of the forested area of

Russia (Martinsson & Lisinski, 2007) or almost 7% of

the world’s total forested areas (FAO). The Russian

larches have been divided into various species. There

are, according to taxonomists, at least two species,

Larix sibirica Ledeb. and L. gmelinii Rupr., and several

subspecies and hybrids (Milyutin & Vishnevetskaya,

1995). Bobrov (1972) separates L. cajanderi Mayr.

from L. gmelinii and Dylis (1981) suggests a separate

western species, L. sukaczewii Dyl., to be distin-

guished from L. sibirica. Phylogenetic studies (Ba-

shalkhanov et al., 2003; Khatab et al., 2008) support

Dylis and in this paper L. cajanderi and L. sukaczewii

are distinguished as separate species (Figure 1).

The species easily hybridize, however, and the

distribution is more or less continuous across Russia.

Thus, when studying clinal trends in adaptation, the

populations of Russian larches included here can be

treated together at the genus level. They provide an

opportunity to study adaptation over a very wide

geographical range; over 188 latitude and more than

1208 longitude.

In boreal forest trees, adaptation to seasonal

climatic conditions is essential, with frost hardiness

in spring and autumn among the most important

adaptational traits to consider when selecting prove-

nances for planting in forestry. When and how

quickly frost hardiness is lost in spring and built up

in autumn reflect adaptation to maritime versus

continental climate (Howe et al., 2003; Persson

et al., 2006), with temperature sum and day length

Correspondence: T. Eysteinsson, Iceland Forest Service, Midvangi 2-4, Egilsstadir, IS-700 Iceland. E-mail: throstur@skogur.is

Scandinavian Journal of Forest Research, 2009; 24: 100�110

(Received 15 September 2008; accepted 22 January 2009)

ISSN 0282-7581 print/ISSN 1651-1891 online # 2009 Taylor & Francis

DOI: 10.1080/02827580902773470

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as proximal drivers (Dormling et al., 1968; Simak,

1970). There are therefore likely to be both latitu-

dinal and longitudinal components to the variation

in these traits within a species or species complex.

Larch is often damaged by frost if the provenance

is poorly adapted to the climate where it is planted.

Examples of this can be found both in Sweden and

Iceland, where L. sibirica originating from south

central Siberia is largely considered to have failed

(Eysteinsson et al., 1994; Abaimov et al., 1998). The

seed sources were in the Krasnoyarsk area and

Khakasia where the climate is probably too con-

tinental compared to Sweden and especially Iceland.

These trees generally survived frost injuries but often

sustained secondary damage due to pathogens. Frost

injuries also resulted in poor stem form by killing the

terminal shoot. Eysteinsson and Skulasson (1995)

showed a positive correlation between frost resis-

tance and growth and stem form in young plants.

Therefore, good frost resistance is extremely impor-

tant when the goal is to produce high-quality timber.

The sensitivity to frost among European larch

(L. decidua Mill.) and L. sukaczewii when planted in

Sweden was studied by Simak (1969, 1970, 1979).

Major findings were that autumn frost resistance was

strongly connected with the photoperiodic response

of the introduced larches. Southern provenances

experience longer days and colder temperatures

during autumn in northern Scandinavia than they

are adapted to, leading to poor hardening and

sensitivity to early frosts.

Artificial freezing tests have been recognized as a

good complement to provenance testing in the field

for determining frost tolerance in conifers

(Andersson, 1992; Aitken & Adams, 1997; Persson

et al., 2006). High correlation between artificial

freezing tests and field performance has been shown

for Scots pine at the provenance level (Nilsson &

Andersson, 1987) and for coastal Douglas fir (Aitken

& Adams, 1997).

Beginning in 1996, the Russian�Scandinavian

larch project started to collect and test larch prove-

nances representing the entire range of larches in

Russia. Seed was collected over a 5-year period and

field trials were established in 10 countries (Mar-

tinsson & Takata, 2005). This undertaking is now

providing access to material in provenance and

family trials, allowing study of range-wide adaptation

in Russian larches that has until now not been

feasible. This study aims to determine variation

and clinal trends in spring and autumn frost toler-

ance, which are indicators of adaptation of growth

rhythm to climate. From a practical standpoint, it

Figure 1. Location of the 28 provenances, two seed stands and two seed orchards as numbered in Table I. The provenance trial site where

the shoots were collected is marked T. The six different patterns indicate larch species and their hybridization zones 1�Larix sukaczewii

Dyl., 2�L. sibirica Ledeb., 3�L.�czekanowskii, 4�L. gmelinii Rupr., 5�L. gmelinii�L. cajanderii, 6�L. cajanderi Mayr. Map based on

Milyutin and Vishnevetskaya (1995) and modified according to Schmidt (1995) and Putenikhin and Martinsson (1995).

Frost tolerance in Russian larches 101

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provides insight into provenance selection for var-

ious conditions.

Materials and methods

Twigs for the freezing tests were collected in a

provenance field trial established in 2003 in Jarvtrask,

Sweden (65811? N, 19831? E, 410 m a.s.l.). From

each of the 28 provenances (Table I), 15 trees were

randomly selected to be included in the tests, five trees

of each provenance in three different blocks. For

comparison, material originating from two Russian

seed stands (Ivanovo and Irkutsk) and two Nordic

seed orchards [Ostteg (Sweden) and Lassinmaa (Fin-

land)] growing in the same trial was included. The

Russian seed stands contain local material from their

respective regions, the Ostteg seed orchard is com-

posed of L. sukaczewii mostly of Archangelsk origin

selected in plantations in northern Sweden and the

Lassinmaa seed orchard contains L. sukaczewii of

Raivola origin selected in Finnish plantations. In total,

shoots from 480 trees were tested.

Five 10 cm long twigs were cut from the top half of

the stem of each sampled tree. Only trees taller than

50 cm were included in the sampling frame. The

twigs for the spring test were collected on 7�8 May

2006, when buds on about half of the provenances

had started to swell and the earliest provenances had

started to produce needles. The twigs were labelled,

put in a plastic bag and then into an insulated box

(cooler). As the temperature was high during collec-

tion it was important to place the twigs in cool

storage as soon as possible. In each bag, moist moss

was placed with the twigs to avoid drying (B.

Andersson, personal communication, May 2006).

The twigs were then sent by air transport to Iceland

and were received within 24 h.

Collection of twigs for the autumn freezing test

took place on 26�27 September 2006 and followed

the same procedure as collection during spring. All

the trees had set bud but the provenances were in

various states of needle senescence, some having

dropped their needles while others were still fully

green.

Table I. Geographical origin of the 28 larch provenances, two seed orchards and two seed stands used in the freezing tests.

No. Region Provenance Latitude (N) Longitude (E) Elevation (m a.s.l.) Species

1 Arkhangelsk Onega 64801? 38815? 110 L. suk.

2 Arkhangelsk Emtsa 63800? 40821�25? 100�120 L. suk.

3 Arkhangelsk Shalakusha 62809? 40819? 120 L. suk.

4 Niz. Novgorod Vetluga 57830? 45810? 145 L. suk.

5 Komi Usinsk 66800? 57848? 75 L. suk.

6 Yamalia Kharp 66856? 65845? 130 L. suk.

7 Yamalia Labytnangi 66828? 66839? 40 L. suk.

8 Khanti-Mansi Beloyarsk 63841? 66844? 60 L. sib.

9 Perm Osa 57819? 55827? 160 L. suk.

10 Sverdlovsk Visim 57830? 59848? 350 L. suk.

11 Ufa Maginsk 55845? 56858? 370 L. suk.

12 Ufa Zilair 52813? 57825? 550 L. suk.

13 Chelyabinsk Nyazepetrovsk 56809? 59832? 460 L. suk.

14 Chelyabinsk Kyshtym 55843? 60827? 480 L. suk.

15 Chelyabinsk Zlatoust 55807? 59830? 600 L. suk.

16 Chelyabinsk Miass 54858? 60807? 380 L. suk.

17 Kemerovo Antoninovka 54812? 88842? 700a L. sib.

18 Kemerovo Mezhdurechensk 53848? 88800? 400a L. sib.

19 Kemerovo Kondoma 52848? 87824? 600a L. sib.

20 Altai Aktash 50812�16? 87803�54? 1600 L. sib.

21 Krasnoyarsk Boguchany 58839? 97830? 96�158 L. sib.

22 Sakha Zhigansk 66845�51? 123821�22? 70�90 L. caj.

23 Magadan Motykleyka 59830? 148830? 80 L. caj.

24 Magadan Sokol 59850? 150840? 60 L. caj.

25 Magadan Nyurchan 59820? 152830? 100 L. caj.

26 Khabarovsk Vanino 49808�12? 139800? 90�125 L. gme.

27 Sachalin Nogliki 51848? 143809? 50 L. gme.

28 Kamchatka Esso 55848? 158840? 700a L. gme.

29 Seed stand (Ru) Ivanovo 578 418 130a L. suk.

30 Seed stand (Ru) Irkutsk 528 1048 500a L. sib.

31 Seed orchard (Fi) Lassinmaa 62804? 25809? 107 L. suk.

32 Seed orchard (S) Ostteg 63848? 20816? 10 L. suk.

Note: aelevations estimated based on map coordinates.

L. suk.�Larix sukaczewii; L. sib�L. sibirica; L. caj.�L. cajanderi; L. gme.�L. gmelinii; Ru�Russia; Fi�Finland; S�Sweden.

102 T. Eysteinsson et al.

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Frost tolerance testing took place at the Icelandic

Agricultural University’s freeze-testing laboratory at

Modruvellir in northern Iceland directly after the

twigs arrived. The twigs were tested at five different

minimum temperatures, �4 (control), �8, �12,

�16 and �208C, one twig from each tree, i.e. a total

of 15 twigs per provenance at each temperature. They

were cooled quickly from room temperature to 48Cand then at a rate of 28C h�1 to the minimum for

each group of twigs, kept at the minimum tempera-

ture for 2 h and then warmed up again at a rate of

28C h�1. The cooling and warming rate was chosen

to roughly mimic the way a severe frost event might

occur in nature. The total duration that the twigs were

kept below freezing was longer as the minimum

temperatures were lower, a total of 9 h for the

�88C group to 21 h for the �208C group. After

freezing, the twigs were stuck in sand-filled trays and

kept in a warm (208C) greenhouse under a misting

system for 2 weeks to allow damage to develop. Frost

damage to dissected cambium and terminal buds was

then scored visually, using a 12 point scale for the

cambium, where 1�no damaged tissue, 12�no

undamaged tissue and the numbers in between

corresponded to percentile ranges (i.e. how large a

proportion of the entire shoot length had damaged

cambium, 2�1�10% of tissue damaged, 3�10�20%

damaged and so on). This method has been used

successfully to assess freezing damage to both needles

and shoots (Nilsson & Andersson, 1987; Lindgren &

Nilsson, 1992; Eysteinsson & Skulasson, 1995). Buds

were rated as either damaged or undamaged.

Statistical analysis was carried out using Sigmastat

(Systat Software, 2007). The data did not follow a

normal distribution, requiring the use of non-para-

metric tests. Friedman repeated-measures ANOVA

was used on untransformed data to analyse tem-

perature effects and provenance differences, with

post hoc Tukey tests to separate means. In the

Friedman test, the damage scores are ranked within

each provenance and temperature and a Q statistic is

calculated by dividing the sum of squared differences

between observed and expected (average) rank by

the expected (average) sum of squared differences

(Q�SSt/SSe). The Q statistic is then tested against a

chi-squared distribution.

Multiple linear regression analyses were per-

formed using overall mean damage scores (spring

and autumn separately) for each provenance as the

dependent variables, and provenance latitude, long-

itude and elevation as independent variables, yield-

ing equations in the form of:

D�k�a�Latitude�b�Longitude�c�Elevation

where D is the spring or autumn damage score, k is a

constant, and a, b and c are regression coefficients.

By using mean damage scores over all freezing

temperatures, the data conformed to assumptions

of statistical normality and equal variance, making

them suitable for parametric testing. This analysis

was carried out over all of the provenances from

Russia (excluding the two Nordic seed orchards).

The L. sukaczewii provenances were then analysed

separately, because suspicion arose that they showed

a different pattern in spring damage with respect to

latitude from the other species.

Results

Cambial damage scores and bud damage scores were

highly correlated (Spearman r�0.625 for spring and

0.927 for autumn, pB0.0001). To avoid repetition,

only the cambial damage results are presented here.

Freezing temperatures

Overall, the freezing temperatures used were suffi-

cient to produce a range of frost damage among the

provenances tested (Figure 2). In the spring test the

unfrozen (�48C) control shoots of the five north-

easternmost provenances [Zhigansk (no. 22), Motyk-

leyka (23), Sokol (24), Nyurchan (25) and Esso (28)]

along with the Irkutsk seed stand material (30) were

damaged, indicating loss of frost tolerance and frost

damage before testing. The two south-easternmost

provenances [Vanino (26) and Nogliki (27)] were

severely damaged at �88C but others were mostly

tolerant to either �12 or �168C. No provenances

were absolutely free of damage at �128C or below,

but three provenances, Kharp (6), Maginsk (11) and

Emtsa (2), were only slightly damaged at �208C(damage score B3).

123456789

101112

4 -8 -12 -16 -20 4 -8 -12 -16 -20Temperature °C

Cam

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b

c c

aab

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Spring test Autumn test

Figure 2. Overall cambial damage by freezing temperature in the

spring (early May) and autumn (early October) tests. Letters

above the bars indicate significant differences (pB0.05).

Frost tolerance in Russian larches 103

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In the autumn test, damage increased as test

temperatures became progressively colder, with sig-

nificantly greatest damage at �208C (pB0.0001).

Only the two south-easternmost provenances [Va-

nino (26) and Nogliki (27)] were severely damaged

(damage score �7) at �128C. Three northern

provenances, Usinsk (5), Labytnangi (7) and

Zhigansk (22), were undamaged and two others,

Kharp (6) and Beloyarsk (8), along with the Ostteg

(32) seed orchard material, were only slightly

damaged at �208C (damage score B3).

Autumn frost tolerance

The variation among provenances in frost damage

was smaller in autumn than in spring (Figure 3).

North-western L. sukaczewii provenances exhibited

the lowest amount of autumn frost damage, along

with the northern-most L. cajanderi provenance

Zhigansk (22). With the exception of Zhigansk,

the far-eastern L. cajanderi and L. gmelinii prove-

nances showed most damage, especially the two

southern ones, Nogliki (27) and Vanino (26). The

L. sibirica provenances, along with some southern

Ural L. sukaczewii provenances, showed intermedi-

ate damage.

Multiple linear regression analysis of autumn

damage by provenance latitude, longitude and

elevation yielded the following equation:

Autumn damage�19:77�(0:29�Latitude)

�(0:02�Langitude); SE�0:72

Provenance latitude and longitude both contributed

significantly (R2�0.531 and 0.356 respectively,

pB0.001) in explaining variation in autumn frost

damage (farther north�less damage, farther east�more damage) (Table II, Figure 4).

No significant elevational effect was observed in

either spring or autumn. In fact, there was a weak,

counterintuitive trend showing increasing spring and

autumn damage with increasing elevation (Figure 5).

Spring frost tolerance

Over all test temperatures combined, provenances

from north-western Russia and the Urals (L. sukac-

zewii) were least damaged in the spring test

(Figure 6). Of the north-western provenances, only

the Ivanovo seed stand (29) showed significantly

greater damage than the least damaged provenances,

while the Ural provenances (nos 9�16) showed

greater variation in damage than other geographical

provenance groupings (Figure 6). All the L. cajan-

derii and L. gmelinii provenances from the Siberian

far east were severely injured with no significant

differences among them. Southern Siberian L.

sibirica provenances showed intermediate damage,

with only the Irkutsk seed stand (30) showing

significantly more damage than the least damaged

provenances within that species.

Multiple linear regression analysis of spring da-

mage by provenance latitude, longitude and eleva-

tion yielded the following equation:

Spring damage�0:55�(0:07�Longitude);

SE�1:53

Provenance longitude was the only significant

(R2�0.761, pB0.001) predictor of frost damage

score in the spring test (farther east�more damage)

(Table II, Figure 7).

Taken separately, L. sukaczewii showed a different

pattern in spring frost tolerance with respect to

latitude than all species combined. In L. sukaczewii,

multiple linear regression analysis of spring damage

1

2

3

4

5

6

7

8

9

10

11

12

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7 La

by22

Zhi

g6

Kha

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o32

Öst

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Akt

a31

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s1

One

g2

Em

ts10

Vis

i15

Zla

t21

Bog

u4

Vet

l12

Zila

11 M

agi

13 N

yaz

29 Iv

an16

Mia

s25

Nyu

r9

Osa

30 Ir

ku14

Kys

h17

Ant

o18

Mez

23 M

oty

19 K

ond

28 E

sso

24 S

oko

26 V

ani

27 N

ogl

Provenances

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L. sukaczewii

L. gmelinii

L. sibirica

L. cajanderi

Figure 3. Autumn cambial damage by provenance. Mean values for all five test temperatures combined (1�undamaged, 12�100%

damage). Provenances under each horizontal line did not differ significantly in damage (pB0.05).

104 T. Eysteinsson et al.

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by latitude, longitude and altitude yielded the

following equation:

Spring damage�16:16�(0:20�Latitude);

SE�1:53

Provenance latitude was the only significant

(p�0.024) predictor of spring frost damage for L.

sukaczewii (Figure 8).

Discussion

Freezing temperatures

Several provenances were undamaged at �208C in

one or both tests. Although the range of freezing

temperatures used in the tests was sufficient to

differentiate among most of the provenances, lower

temperatures, possibly much lower, would have been

required to test the full range of frost tolerance found

in Russian larches in spring and autumn.

Autumn frost tolerance

The autumn test yielded results that were in agree-

ment with previous research, with northern prove-

nances showing less frost damage than southern ones

at similar longitudes. There was also a significant

longitudinal component to autumn frost damage,

with eastern provenances showing more damage

than western ones at similar latitudes. An explana-

tion might be an autumn maritime effect, i.e.

adaptation in the far-eastern material to later arrival

Table II. Regression coefficients and their standard errors, coefficients of determination (R2) and p values from multiple linear regression of

spring and autumn frost damage with provenance latitude, longitude and elevation.

Regression coefficient SE of coefficient R2 p

Spring

Latitude �0.107 0.0722 0.104 0.149 (ns)

Longitude 0.0665 0.00696 0.761 B0.001

Elevation �0.000387 0.00108 0.00587 0.722 (ns)

Total 0.781 B0.001

Autumn

Latitude �0.296 0.0332 0.531 B0.001

Longitude 0.0178 0.00320 0.365 B0.001

Elevation �0.00224 0.000494 0.00297 0.775 (ns)

Total 0.839 B0.001

Note: ns�not significant.

0

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48505254565860626466

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tum

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East

Lon

gtiu

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North Latitude

Figure 4. Autumn frost damage for all provenances by latitude

and longitude. The plane is a plot of the multiple linear regression

equation: Autumn damage�19.773 � (0.296�Lat.)�(0.0178�Long.). Both latitude and longitude were significant (pB0.001)

predictors of autumn frost damage.

Elevation (m a.s.l.)0 200 400 600 800 1000 1200 1400 1600 1800

Mea

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amag

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ore

- A

utu

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0

1

2

3

4

5

6

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8

Figure 5. Scatterplot of autumn frost damage by elevation along

with the regression line that best fits the data points and 95%

confidence intervals. Autumn frost damage was not significantly

(p�0.775) correlated with elevation and the same was true of

spring frost damage.

Frost tolerance in Russian larches 105

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of autumn frosts compared to inland areas because

of proximity to the Pacific Ocean, still warm from

the summer.

Longitudinal differentiation in autumn frost hardi-

ness has also been observed in Scots pine (Pinus

sylvestris L.). Comparing populations from Scandi-

navia and Russia of corresponding latitudes,

Andersson and Fedorkov (2004) found that Russian

populations (from a more continental climate) were

clearly more frost resistant than Scandinavian ones,

concluding that not only latitude but also to

some extent longitudinal origin (degree of continen-

tality) determines autumn frost hardiness in Scots

pine.

The effect of provenance elevation on autumn

frost tolerance did not yield a discernible pattern,

although the highest elevation provenance,

Aktash (20), was more frost tolerant than other

0

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10

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48505254565860626466

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rin

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Lon

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North Latitude

Figure 7. Spring frost damage for all provenances by latitude and

longitude. The plane is a plot of the multiple linear regression

equation: Spring damage�6.039 � (0.0914�Lat.)�(0.0642�Long.). Only longitude was a significant (pB0.001) predictor of

spring frost damage over all provenances.

Figure 8. Spring frost damage, for Larix sukaczewii provenances

only, by latitude and longitude. The plane is a plot of the multiple

linear regression equation: Spring damage�16.157 � (0.204�Lat.) � (0.000115�Long.). Latitude was the only significant

(p�0.024) predictor of spring frost damage when only L.

sukaczewii was included.

1

2

3

4

5

6

7

8

9

10

11

12

6 K

har

11 M

agi

2 E

mts

5 U

sin

31 L

ass

32 Ö

stt

3 S

hal

7 La

by

9 O

sa

13 N

yaz

8 B

elo

15 Z

lat

14 K

ysh

1 O

neg

4 V

etl

18 M

ezh

29 Iv

an

17 A

nto

12 Z

ila

10 V

isi

20 A

kta

21 B

ogu

19 K

ond

16 M

ias

30 Ir

ku

25 N

yur

27 N

ogl

26 V

ani

28 E

sso

24 S

oko

23 M

oty

22 Z

hig

Provenance

Cam

bia

l dam

age

spri

ng

L. sukaczewii

L. sibirica

L. gmelinii

L. cajanderi

Figure 6. Spring cambial damage by provenance. Mean values for all five test temperatures combined (1�undamaged, 12�100%

damage). Provenances under each horizontal line did not differ significantly in damage (pB0.05).

106 T. Eysteinsson et al.

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provenances from similar latitudes. Observation of

an elevational trend was compromised by southern,

higher elevation provenances being compared to

more northern lowland provenances. This resulted

in provenance differences in autumn frost tolerance

being small, both generally between north and

south and especially between low and higher

elevations. Furthermore, the differences in elevation

between most of the provenances were not great.

Therefore, these results should not be interpreted

to mean that provenance elevation is unimportant

in the development of spring and autumn frost

tolerance.

Spring frost tolerance

The provenances least damaged in spring were the

north-western L. sukaczewii provenances, including

the ones from farther north than the test site

[Usinsk (5), Kharp (6) and Labytnangi (7)], along

with some more southern Ural provenances and the

comparison seed orchard material from Lassinmaa

and Ostteg (Figure 6). At similar longitudes, the

northern Arkhangelsk provenances (1�3) showed

less spring frost damage than the more southern

Nizhniy Novgorod (4) and Ivanovo (29), and the

northern Komi and Ob-Ural provenances (5�8) had

significantly less damage than the southern Ural

provenances (9�16) (Figure 8). In other words,

northern L. sukaczewii provenances stayed frost

tolerant longer in spring than more southern ones

growing in the same common-garden trial in

Sweden.

The present results for L. sukaczewii are in

contrast with what has been found in studies of

Norway spruce and larch at high latitudes. In

general, experience from provenance testing is that

northern populations are more susceptible to spring

frost damage than more southern populations when

grown on the same site owing to a lower heat sum

requirement for budburst (Eriksson et al., 2006).

Thus, Carswell and Morgenstern (1995), studying

the phenology and growth of nine larch species

tested in New Brunswick, Canada, found that

northern and high-elevation provenances were

more damaged by spring frost than southern ones.

Furthermore, Simak (1979) compared the climate at

similar latitudes in northern Sweden and western

Russia and found that spring starts earlier but

progress at a slower rate in Sweden. This would,

according to Simak, lead to earlier budburst for

same-latitude material planted in Sweden, thus

increasing the risk of early spring frost damage. To

reach a similar temperature regime in Sweden as in

Russia, L. sukaczewii should, according to Simak, be

transferred 3�5 degrees latitude north. The results

here do not support that conclusion for L. sukacze-

wii. However, the same pattern as in Simak (1979),

but considerably weaker, occurred in this study for

L. sibirica, L. gmelinii and L. cajanderii. Thus, the

pattern seen in L. sukaczewii is surprising and calls

for an explanation.

Overall, provenance longitude explained the varia-

bility in spring frost damage to a much greater extent

than latitude. Longitude is a proxy for the continen-

tality of climate, i.e. how likely it is that warm

temperatures occur in winter. In northern continen-

tal areas, the likelihood of winter temperatures above

freezing is practically nil until the arrival of spring.

Trees from such areas have only minimal adaptation

against losing frost hardiness when temperatures rise

above freezing in late winter. Winter temperatures

above freezing occur more often in proximity to an

ocean in the direction from which air masses often

arrive, but at northern latitudes frosts can also occur

until late spring. Northern provenances adapt to

boreal maritime climates in the same way that

more southern provenances adapt to warm spells

in winter, by developing a higher chilling require-

ment and/or heat sum requirement for loss of

frost hardiness and initiation of growth (Hannerz,

1994; Leinonen, 1996; Leinonen & Hanninen,

2002).

With the exception of shore-bound ice, the

Barents Sea is often ice free in late winter as far

east as Novaya Zemlya (Polar Research Group,

2008). This means that the north-western L. sukac-

zewii provenances are only a few hundred kilometres

from open ocean and thus under maritime influence

at least occasionally. They have adapted to tolerate

warm spells in late winter, in contrast to more

continental provenances, which in the case of L.

sukaczewii are found farther south.

Northern Sweden is also under considerable mar-

itime influence, and the north-eastern provenances

Zhigansk (22), Sokol (24), Motykleyka (23) and Esso

(28) were already damaged before the freeze testing

commenced (i.e. the control shoots were damaged).

Since they all had good to intermediate frost tolerance

in the autumn test, damage probably occurred in

winter or early spring, indicating poor adaptation to a

maritime climate. The south-eastern provenances (26

and 27) are not far behind in that respect. Thus, the

geographical pattern in adaptation to a continental

climate that results in quick loss of frost hardiness in

spring covers the entire range of latitude in eastern

Siberia but narrows to an inland tongue towards

the west.

The clearest indication of adaptation to maritime

climate was the lack of spring frost damage in the

provenances Usinsk, Kharp and Labytnangi (5�7)

compared to Zhigansk (22), which was most severely

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damaged. They are all from roughly 668 N latitude,

with the three western provenances in proximity to

the ice-free Barents Sea, but Zhigansk in the middle

of a very large late-winter ‘‘continent’’ that stretches

across the frozen Arctic Ocean and North America

as well as Eurasia. The three western provenances

have adapted to avoid early loss of frost hardiness as

a response to occasional late winter warm spells,

while the eastern provenance did not have such

adaptation and was damaged in the maritime climate

of the trial in northern Sweden. This could have

serious implications with regard to global warming.

If the Arctic Ocean north of Siberia becomes ice

free in late winter, resulting in a more maritime

climate for northern Siberia, the incidence of winter/

spring frost damage to larch growing there could

increase.

It should be noted that these four northernmost

provenances show poor growth and Zhigansk

(no. 22) has the lowest survival in the provenance

trial where the freeze-test material was collected

(Karlman & Martinsson, 2007), a common out-

come when trees are planted farther south than

their origin. In the case of the three western

provenances, however, poor growth performance

cannot be attributed to poor spring or autumn frost

tolerance.

Comparison with seed stand and seed

orchard material

The comparison seed stand material [Ivanovo (29)

and Irkutsk (30)] differs from other provenances

only in that the seed used in establishing the

Jarvtrask field trial was collected in stands that

have been managed for seed production, i.e. by

thinning that presumably involved selection for

growth and form. This might have had an effect on

adaptation in the resulting progeny, but there is no

indication of that here. Both spring and autumn frost

tolerance in the Ivanovo material was consistent with

its geographical location within the range of L.

sukaczewii. The Irkutsk material was the least frost

tolerant of the L. sibirica provenances in spring,

which is consistent with the fact that it is also the

easternmost provenance of that species in the trial.

Thus, both seed stands fit well with the general

trends in frost tolerance observed among the prove-

nances in this study.

Most L. sukaczewii planted in the Nordic coun-

tries in recent years has been of Raivola (Lintula)

origin (Redko & Malkonen, 2005).

The two comparison seed orchards [Lassinmaa

(31) and Ostteg (32)] are comprised of L. sukaczewii

originating mostly in north-west Russia, but with the

parental clones selected in Finnish and Swedish

progeny and provenance trials (Martinsson &

Lesinski, 2007). They behaved as the north-west

Russian provenances, with little damage in both

spring and autumn, and thus also fit well with the

trends in this study. Apparently, the selection process

used when establishing the seed orchards did not

result in changes in spring and autumn frost

tolerance, as compared to provenances of similar

origin.

Conclusions

Even though the provenances have sometimes been

treated as groups to facilitate discussion, the frost

tolerance test results presented here did not yield

groupings of provenances with clear differences

between species. Instead, they showed a roughly

continuous variation in frost tolerance (Figures 3

and 6), mostly trending east�west for spring frost

tolerance and north�south trending in autumn

(Figures 4 and 7). Larix sukaczewii did show a

latitudinal trend in spring frost tolerance, which

may be explained by adaptation to maritime condi-

tions in north-west Russia.

When considering adaptation, latitude and long-

itude are proxies for climatic factors, with longitude

mostly indicating maritime versus continental condi-

tions because of the orientation of the Eurasian

landmass and prevailing westerly winds, while lati-

tude represents the arrival of warm temperatures in

spring and differences in day length in late summer

and autumn. The strong longitudinal effect in spring

for larch in northern Russia indicates adaptation to

the different sea ice cover east and west of Novaya

Zemlya.

Frost tolerance testing cannot by itself be the only

criterion for selecting provenances for use in forestry

in a specific area. It can, however, provide clues to

narrow the search. When selecting Russian larch

provenances for use in forestry in the Nordic

countries, provenances ‘‘downwind’’ and in fairly

close proximity to open ocean in spring seem to be

the best adapted from the standpoint of frost

hardiness in spring and autumn. These are the

north-western provenances, including the Lassinmaa

and Ostteg seed orchard material. This study did not

reveal any Russian larch provenances better adapted

to climatic conditions in the Nordic countries than

the ones already in use. However, some western

Russian, Ural and southern Siberian provenances

are also likely to be sufficiently adapted for use in

forestry in the Nordic countries. Far-eastern prove-

nances, regardless of latitude, are most likely to

suffer from spring frost damage in more maritime

regions.

108 T. Eysteinsson et al.

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Acknowledgements

This work was supported by grants from the Kempe

Foundation and the Northern Periphery project

SIBLARCH. We are grateful to Lena Helin, Tom

Gards, Lidia Kovler, Kurt Olsson, Nicole Suty,

Freyr Ævarsson, Aleksey Fedorkov, Alexandra Ber-

kutenko, Antti Lukkarinen and Seppo Ruotsalainen

for their assistance in fieldwork and help with

gathering information. Dag Lindgren and Bjorn

Hanell made valuable comments on the manuscript.

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