Theoretical and Applied Genetics 44, 49-- 57 (1974) | b y Springer-Verlag 1974

Survival and Growth in Scots Pine (Pinus silvestris L.) Provenance Experiments

in N o r t h e r n S w e d e n

VILHELMS EICHE and ENAR ANDERSSON D e p a r t m e n t of F o r e s t Genetics, R o y a l College of F o r e s t r y , S t o c k h o l m (Sweden)

Summary. Genotype -- environment interaction has been investigated in Scots pine provenance experimental series, laid out during 1952--1955 in Sweden. Results obtained from six experimental plantations in the county of Norrbotten during a period of 20 years (including 1970) are the subject of discussion. The genecological variation of climatic hardiness, survival and growth -- rate are shown graphically and in statistical analyses. Response of seed sources to the transfer of populations from their native habitats to the experimental sites has been analysed. The changed length of the growing season in connection with the transfer of populations is taken as the criterion of climatic differences on natural habitats and experimental sites.

I. Introduction D u r i n g the two last decades two p r o v e n a n c e exp e r i m e n t a l series w i t h Scots p i n e h a v e a t t r a c t e d considerable i n t e r e s t in Sweden. One of t h e m was laid o u t b y S t e f a n s s o n in t950 a n d t951, a n d t h e o t h e r b y E i c h e i n c o o p e r a t i o n w i t h the staff of the R o y a l College of F o r e s t r y d u r i n g 1952--1955. A n a c c o u n t of S t e f a n s s o n ' s series was r e n d e r e d b y Stefansson, t 9 6 5 ; S t e f a n s s o n a n d Sinko, t 9 6 7 ; b y H a g n e r , 1970 a n d b y R e m r 6 d , t973. A r e p o r t on E i c h e ' s series was m a d e b y Eiche, 1962, 1966; L a n g l e t , t 9 6 8 ; Eiche a n d G u s t a f s s o n , 1970 a n d E r i k s s o n , 1972. D a t a o b t a i n e d from e x p e r i m e n t a l p l a n t a t i o n s l o c a t e d in the c o u n t y of N o r r b o t t e n (Fig. 1) are a c c o u n t e d for in the present report, w h i c h deals w i t h E i c h e ' s series.

2. Material and M e t h o d s The provenance material in this series consists of 100 different provenances, originating from habitats systematically chosen in Sweden and Norway from native stands and from Germany and Holland (Fig. 1). The most northern population originates from F i n n m a r k in Norway, 6 9 ~ Of these 100 provenances 90 are represented by the progency of 25--30 single trees in each population. The total n u m b e r of single trees whose progenies (families) were tested in these experiments is 2200. Seeds were obtained after open pollination. Four standard populations were used throughout the whole experimental series. They are: no. 10, Korpilombolo, Norrbotten, 66~ N; no. 23, Mal~tr/~sk, V/~sterbotten, 65~ no. 42, Sveg, J/imtland, 6 2 ~ and no. 74, Eckersholm, J6nk6ping, 570 36' N. The experimental series was laid out during 1952--1955 on 30 localities, distributed over the main part of the country, with a total area of 36 hectares. The most northern experimental plantation (EP) is situated near Kiruna. On the EPs unilorm tending of plants was carried out. During t954 to 1962 the plants were sprayed with CaS-solution to protect them against snow-blight fungus (Phacidium infestans Karst.). Most of the EPs were fenced. Experimental series described in the present report were laid out with the view of studying the climatic hardiness

of the plants, their survival, growth-rate and quality, when transferring populations in meridional direction, from north to south and vice versa, or on a vertical scale, i. e. from a higher to a lower altitude and vice versa, respectively, compared with the locality of the EP. On each experimental site 7, 14 or 20 populations were tested in four replications according to modified Youden square design, One of these populations is a local strain and two are standard ones. The remaining are two plus two provenances, whose natural habitats lie to the north or to the south of the experimental locality, respectively. One of these provenances originates from a higher altitudinal range and the other from a lower one, compared with the altitude of the experimental site. I n experiments with 14 (or more) populations, the additional 7 (or more) populations were transferred from more remote habitats. In all EPs each population was represented by 20 families and 13 offspring plants per family, i. e. 260 individuals per population. Mortality and all changes in the composition of the plants were registered on all EPs north of 60 ~ N every year, and on other EPs with an interval of a few years. A general survey of the growth-rate of the provenance material and of its quality was carried out in Norrland in 1967 and 1970, and in other parts of the country during 1968--1970. Six of the thirty EPs are laid out in Norbotten with 4-year old plants (2+2). Out of these six plantations the following five 1 plantations will be accounted for, namely: No.

Locality

~o

Hs

Y

27 29 24 25 22

Kiruna, Kauppinen G~llivare, Linalombolo K~bdalis, Rahaberget Korpilombolo, Ohtanaj~rvi Nlvsbyn, Aspl6vberg

67 ~ 50" 67 ~ t4" 66 ~ 16' 66 o 56" 65 ~ 38"

360 475 440 200 60

100 98 105 113 127

E P 29 lies near the tree limit in vertical direction, E P 27 is the most northern E P in this experimental series and the position of E P 24 is in upland near the Arctic Circle. Harsh climate is a c o m m o n characteristic for these three EPs. Climatic conditions of E P 25 are more 1 The sixth plantation (EP 26) is identical with E P 29, b u t is located in an extreme frost locality.

50

V. E i c h e a n d E. A n d e r s s o n : S u r v i v a l a n d G r o w t h in Scots P i n e ( + ) a n d d o w n w a r d ( - - ) ; Y -- s h o r t e n i n g (--) or l e n g t h e n i n g ( + ) of t h e g r o w i n g season. CM ( c u m u l a t i v e m o r t a l i t y ) : p e r c e n t a g e of s u r v i v e d t r e e s f r o m 260, i. e., f r o m t h e n u m b e r of p l a n t s i n t h e y e a r t h e e x p e r i m e n t was laid out. Tree g r o w t h : g -- m e a n n u m b e r of t r e e s p e r r e p l i c a t i o n , a p p l i e d in a n a l y s e s of v a r i a n c e ( m e a n n u m b e r of p l a n t s p e r r e p l i c a t i o n i n 1954 was 65), t - - u n w e i g h t m e a n of h e i g h t i n c r e m e n t in c m in 1970, h -- u n w e i g h t m e a n of h e i g h t in c m for t w e n t y - y e a r old trees. 3" S u r v i v a l

of Provenance

Material

D a t a o n t h e m o r t a l i t y r a t e of s i n g l e p r o v e n a n c e s in each experimental plantation (EP) are shown on t w o t y p e s of m o r t a l i t y d i a g r a m s : t . C u m u l a t i v e m o r t a l i t y (CM) a n d 2. S i n g l e y e a r m o r t a l i t y ( S Y M ) . Results obtained comprise u n t i l t h e a u t u m n of t 9 7 0 .

analyses

carried

out

//

,

/

I

,,'/ /

/

"21- ~. . . . . . . . . . . . . . . . . . . . . . .

"/

.i

/ // / ~ /

I i

.,"? Y I I

,~.~-..~

n

511

i

I /

Fig. 2. Cumulative mortality (CM) in E P 27 (see Table t) T a b l e t. Experimental plantation 27, Norrbotlen, Kiruna,

Kauppinen. qJ: 67~ o', 2 : 2 o ~ Fig. 1. Provenance trials with Scots pine (Pinus silvestris L.) laid out in 1952--1955. Experimental plantations are denoted by triangles and populations by black dots. Interrupted lines drawn from the Arctic Circle in NW-SE direction denote t h e southern border line of the county of Norrbotten, which occupies the northern p a r t of Sweden m o d e r a t e t h a n t h o s e of E P s 24, 27 a n d 29. T h e c l i m a t e of E P 22 is t h e m i l d e s t of all t h e five E P s .

Symbols in the Table above and in Tables 1 --5 P R : no. -- n u m b e r of p o p u l a t i o n ; 9 -- n o r t h l a t i t u d e ; -- e a s t . l o n g i t u d e f r o m G r e e n w i c h ; Hs - - a l t i t u d e a b o v e t h e sea level ill m ; Y -- l e n g t h of g r o w i n g season, i. e., t h e n u m b e r of d a y s w i t h m e a n t ~ 6 ~ per year (O. L a n g l e t , t936). T r a n s f e r of P R : 9 -- d i s t a n c e in m e r i d i o n a l d i r e c t i o n f r o m t h e n a t u r a l h a b i t a t of t h e P R t o t h e site of E P in kin, in n o r t h e r l y d i r e c t i o n ( + ) a n d i n s o u t h e r l y d i r e c t i o n ( - - ) ; H~ - - d i s t a n c e i n v e r t i c a l d i r e c t i o n i n m, u p w a r d

PR

No

~ Locality

", Hs: 360, Y: loo

Transfer of P R

~

Hs

Y

~

Hs

km

m

Y

2 69055 " 23015 ' 2O 104 - - 2 3 2 + 3 4 0 - 4 Norway, Alta, Aronnes 7 67050 ' 20~ 355 100 -+ 5Norrbotten, Kiruna, Kauppinen 8 67050 " 20~ " 480 94 ---120 + 6 Norrbotten, Kiruna, Nokutosvara t 0 66050 , 23009 , 185 t 1 4 t i l l +t75--14 Norrbotten, Korpilombolo 19 65~ ' 23018 " 3O 128 + 2 2 3 + 3 3 0 - - 2 8 N o r r b o t t e n , Kalix, M6rtr~isk 23 65008 " 18053 , 305 t 1 8 + 3 0 0 + 5 5 - - 1 8 V / i s t e r b o t t e n , Malg, S t r f m f o r s 10t 66003 " 17~ " 460 105 + 1 9 8 - - 1 0 0 5 Norrbotten, Arjeplog

CM t970

%

98.1 77.7 98.9 96.9 95.0 93.9 93.5

Theoret. Appl. Genetics, Vol. 44, No. 2

V. Eiche and E. Andersson: Survival and Growth in Scots Pine

51

Table 2. Experimental plantation 29, Norrbotten, G~illivare, Linalombolo. 9:67~ ", "t,: 2o~ ", Hs: 475, Y: 98 PR No

qp

Locality 4 9 10 11 2O 23 32

Transfer of PR ~.

Hs

Y

qo

Hs

km

m

Y

69007 , 18~ 10o ~05 --210 + 3 7 5 - 7 Norway, M~lselv, Moen, Olsberg 67018 ' 14025" 30 119 -- 7 +445 --21 Norway, Bod6, Mulstrand 66~ " 23009 " 185 114 + 45 +29O--16 Norrbotten, Korpilombolo 67~ 20~ ' 470 98 -+ 5Norrbotten, Giillivare, Linalombolo 65038 , 21007, 75 127 +178 + 4 0 0 - - 2 9 Norrbotten, _~lvsbyn, Aspl6vberg 65008 , 18053" 305 118 +234 + 1 7 0 - - 2 0 V~tsterbotten, Mal&, Str6mfors 64034 , 18015" 510 111 + 2 9 7 - - 3 5 - - 1 3 V/isterbotten, Lycksele, Storberget

CM t970 %

~ i e,

A\

99.6

/ / ,".~,

"

, 'ilJt\ ~\

~

-1/\, },

94.2

, ....

, 't

~=:~

,

t~

\I

"..9:ql

\\,o

83.5 78.5 98.5 96.9

Fig. 3. Single year mortality (SYM) in EP 27 (see Table t) ~0e%

99.2

The lowest m o r t a l i t y rate (CM) in E P 27, Kiruna, is to be found within the local population 7 (Table t, Fig. 2) which surrounds the site. The rate of survival in t970 was 22~o. The other populations had only a few plants left intact, for instance, population t 9 - - 5 % ; population t 0 - - 3 % ; population 23--60/0 . Population 8 from the same neighbourhood, but with a h a b i t a t which lies t 20 m higher, appeared to be the least h a r d y (1%). This population is a typical representative for marginal populations with a low viability. The local population t l in E P 29 (Table 2) lies close to the experimental site. As seen from the CM diagram (Fig. 4), the m o r t a l i t y curve of this population occupies the lowest position. The climatic hardiness of the local population 1t (EP 29) and the local population 7 (EP 27) is similar. Oa, b o t h EPs local populations distinguish themselves b y a higher degree of hardiness t h a n other populations. Among other populations in E P 29, population t0 shows a rate of survival up to 16%. The remaining populations have only a few plants intact. Single year m o r t a l i t y finds its expression in a wavelike line, t h a t is, SYM varies from year to year. Plant death was caused primarily b y cold damage during some years with extreme climatic conditions. Main types of plant damage were: basal stem girdle and frost canker caused b y cold during late winter in some years, and die-back of the top o f plants caused b y late spring frosts (Eiche, t962, 1966; Sakai, t968; Eiche and Gustafsson, t970). The effect of climatic extremes on non-hardy populations is obvious. As a rule, the SYM curves show different kinds of peaks. Often a peak is caused b y the delayed effect of injuries, which had arisen one or two years earlier. For instance, m o r t a l i t y during 1965--t968 was caused b y injuries which appeared during late winter in t965. Sometimes trouble comes m u c h later t h a n expected, as for example, in E P 27 (Fig. 3) and Theoret. Appl. Genetics, Vol. 44, No. 2

5o

/d/~

'("

-

..... --?~' /

cM Ep~9

"/ / 7

.-

i

"

to

/

"

///

Fig. 4. Cumulative mortality (CM) in EP 29 (see Table 2) so "4

~. /

ii

ii'/Ai,

,'

~i ,

ill i

" / ~ J}'!

SYM EP29

,/,,'

.. \,,_. . . ..'

~ Y'-V'--'~

.

,

,.

.-....-%

"%~~'~/

."

l~,.

-

.

It

Y

,,'/y

,~ ,,,/

," ";/

~ i'

!i

Fig. 5. Single year mortality (SYM) in EP 29 (see Table 2) E P 29 (Fig. 5) when the plants were already t4 years old. The SYM diagrams of E P 27 and those of E P 29 are similar. Results obtained from E P 24, Kt~bdalis, are shown in Table 3 and Figs. 6 and 7. Populations 6 and 7 possess a great degree of climatic hardiness and their

52

V. Eiche and E. Andersson: Survival and Growth in Scots Pine Table 3. Experimental plantation 24, Norrbotten, Kdbdalis, Rahaberget. q~: 66~ ,~: 19~ ", Hs: 44 o, Y : so5 PR No

Transfer of PR

q)

~

Hs

Y

Locality 2

qn

Hs

km

m

3 6

7 t0 13 14 23 28 34 38 42 101 107

Estimated trees 1970 CM 1970

Y

69055 " 23015 , 20 1o4 -406 +420 Norway, Alta, Aronnes 68047 " 19~ 25O 100 --280 + 1 9 0 Norway, Mglselv, Dividalen 67045 ' 23022 " 225 107 - - t 6 5 +215 Norrbotten, Kitki6joki 67~ " 20~ 355 100 - - t 7 6 + 85 Norrbotten, Kiruna, Kauppinen 66053 " 23003 " 175 114 -- 69 +265 Norrbotten, Korpilombolo, Smedberg 66018 , 14010 , 75 123 -- 4 +365 Norway, Mo i Rana, Bgsmo 66005 , 20045 " 130 121 + 20 + 3 1 0 Norrbotten, Spikseleg 65008 " 18053" 3O5 118 + 1 2 6 +135 V/~sterbotten, Mal~, Str6mfors 64014 " 19043 " 200 128 +226 +240 Vgsterbotten, Vindeln, Svartberget 63~ 18047 , 8 144 +336 +432 V/isternorrland, 0rnsk61dsvik, Alfredshem 63010 , 13004 , 540 117 +345 --100 Jgmtland, Vallboggrden 62o03 , 14~ 385 132 + 4 6 9 + 55 Jitmtland, Sveg, Malmb/icken 66003 , 17~ 460 105 + 24 -- 20 Norrbotten, Arjeplog 63008 , 16039 , 455 122 + 3 4 9 -- 15 J/imtland, Bispfors, Torresj61andet

%

i

cr~ E.Pz4

,r

/I

"1

.: I

71.5

17.oo

18.49

204.72

+

5

91.9

5.00

17.35

191.36

-- 2

43.9

35.00

23.46

248.73

+

5

44.6

32.75

21.08

222.87

--9

59.2

26.00

22.45

235.92

--18

83.0

10.50

17.32

197.81

--16

88.9

7.25

23.14

235.78

--13

79.1

12.75

23.24

243.64

--23

95.8

2.50

16.27

182.73

--39

100.0

0.25

4.25

47.50

--12

98.9

0.75

t3.75

152.50

--27

t00.0

0.25

3.75

52.00

--

83.5

10.75

21.50

214.95

--17

98.1

1.25

8.00

84.87

2.18

2.70

29.78

n a t e l y , t h e local p o p u l a t i o n is n o t r e p r e s e n t e d in this E P . P o p u l a t i o n 101 f r o m A r j e p l o g is t h e one which comes closest to t h e local p o p u l a t i o n . P o p u l a t i o n 42, Sveg, an d p o p u l a t i o n 34, A l f r e d s h e m , p e r i s h e d alr e a d y in 1964 an d t967, r e s p e c t i v e l y . Single y e a r m o r t a l i t y c u r v e s (Fig. 7) show, on t he whole, t h e s a m e course of d e v e l o p m e n t , as was t h e / :

sYM Ep 2~

/.@2):/

/ io

~,l.r" l

I

.... "7,."~"