Table 1. Information on the crosses conducted in the Ahvenlampi Scots pine seed orchard in 2013. Grafts of three maternal genotypes were cross-pollinated with pollen from seven paternal genotypes from two other Scots pine orchards and pollen from surrounding forests. The cross-combinations with the original genotype codes, the genotype provenance effective temperature sums (d.d.) plus latitudes and the genotype codes applied in the tables and figures are listed. The letters of the genotype codes denote the following: E, K and P = southern, central and northern Finland, respectively; M = maternal genotype; S = southern paternal genotype; N = northern paternal genotype; OP = southern pollen mixture from forests surrounding the orchard of the controlled crosses.
Cross-combination maternal * paternal genotype with the original codes Maternal genotype provenance d.d. and (latitude) Paternal genotype provenance d.d. and (latitude) Applied maternal genotype code Applied paternal genotype code
K205 Open pollinated 1095 (62°34´) 1075 (62°45´) M1 OP
K205 * K543 1095 (62°34´) 1049 (62°50´) M1 S1
K205 * K652 1095 (62°34´) 1112 (62°22´) M1 S2
K205 * K890 1095 (62°34´) 1090 (62°50´) M1 S3
K205 * P573 1095 (62°34´) 692 (68°33´) M1 N1
K205 * P575 1095 (62°34´) 692 (68°33´) M1 N2
K205 * P577 1095 (62°34´) 692 (68°33´) M1 N3
K205 * P625 1095 (62°34´) 692 (68°33´) M1 N4
K214B Open pollinated 1117 (62°37´) 1075 (62°45´) M2 OP
K214B * K543 1117 (62°37´) 1049 (62°50´) M2 S1
K214B * K652 1117 (62°37´) 1112 (62°22´) M2 S2
K214B * K890 1117 (62°37´) 1090 (62°50´) M2 S3
K214B * P573 1117 (62°37´) 692 (68°33´) M2 N1
K214B * P575 1117 (62°37´) 692 (68°33´) M2 N2
K214B * P577 1117 (62°37´) 692 (68°33´) M2 N3
K214B * P625 1117 (62°37´) 692 (68°33´) M2 N4
E2856 Open pollinated 1142 (62°02´) 1075 (62°45´) M3 OP
E2856 * K543 1142 (62°02´) 1049 (62°50´) M3 S1
E2856 * K652 1142 (62°02´) 1112 (62°22´) M3 S2
E2856 * K890 1142 (62°02´) 1090 (62°50´) M3 S3
E2856 * P573 1142 (62°02´) 692 (68°33´) M3 N1
E2856 * P575 1142 (62°02´) 692 (68°33´) M3 N2
E2856 * P577 1142 (62°02´) 692 (68°33´) M3 N3
E2856 * P625 1142 (62°02´) 692 (68°33´) M3 N4
Table 2. Information concerning the natural Scots pine forests included in this study as a source of seed material. The forest locality, latitude, effective temperature sum (d.d.) and ripening year of the seed lot are listed.
Forest locality Latitude d.d. Ripening year
Kittilä 67°22´ 760 1988
Pello 66°48´ 860 1988
Puolanka 65°05´ 880 1997
Pudasjärvi 65°35´ 900 1997
Suomussalmi 64°57´ 980 1991
Pyhäjärvi 63°34´ 1000 1997
Lestijärvi 63°39´ 1020 1991
Rautavaara 63°28´ 1050 2002
Keuruu 62°16´ 1100 2003
Korpilahti 61°55´ 1150 1997
Rauma 60°58´ 1275 2002
Table 3. Results from the ANOVA tests (Model 1). The tests investigated the effect of the Scots pine paternal genotype on the timing of budset (Week) and autumn frost hardening (Br %) of first-year seedlings. Model 1: Y = Intercept + M + P + M*P, where Y = Week or Br %; M = maternal genotype; P = paternal genotype.
Dependent variable: Week Dependent variable: Br %
df F Sig. df F Sig.
Corrected model 20 11.058 <0.001 20 7.264 <0.001
Intercept 1 274849.954 <0.001 1 2951.919 <0.001
Maternal genotype 2 19.581 <0.001 2 7.889 0.001
Paternal genotype 6 26.370 <0.001 6 19.676 <0.001
Maternal genotype * Paternal genotype 12 1.981 0.041 12 0.894 0.558
Error 63 60
df = the number of degrees of freedom; F = F ratio; Sig. = p-value; Week = the calendar week when the terminal buds appeared; Br % = the percentage browning of primary needles.
Table 4. Tukey post hoc test results for the ANOVA tests (Model 1). Pairwise comparisons of the paternal genotype effect (mean difference) on the timing of budset (Week) and autumn frost hardening (Br %) of first-year seedlings of Scots pine. OP, N1–N4 and S1–S3 are codes for the paternal genotypes. The letters of the codes denote the following: S = southern paternal genotype; N = northern paternal genotype; OP = southern pollen mixture from forests surrounding the orchard of the controlled crosses.
Dependent variable: Week Dependent variable: Br %
(I) Paternal
genotype
(J) Paternal
genotype
Mean
difference (I-J)
p-value Signi-
ficance
Mean
difference (I-J)
p-value Signi-
ficance
OP N1 1.313 <0.001 *** 23.548 <0.001 ***
N2 –0.077 1.000 39.812 <0.001 ***
N3 2.322 <0.001 *** 24.480 <0.001 ***
N4 1.199 <0.001 *** 29.038 <0.001 ***
S1 –0.203 0.991 0.942 1.000
S2 0.735 0.072 –0.425 1.000
S3 0.491 0.495 13.663 0.060
N1 N2 –1.389 <0.001 *** 16.264 0.009 **
N3 1.009 0.003 ** 0.933 1.000
N4 –0.113 1.000 5.490 0.912
S1 –1.516 <0.001 *** –22.606 <0.001 ***
S2 –0.578 0.285 –23.973 <0.001 ***
S3 –0.822 0.028 * –9.885 0.358
N2 N3 2.398 <0.001 *** –15.332 0.017 *
N4 1.276 <0.001 *** –10.774 0.228
S1 –0.127 1.000 –38.870 <0.001 ***
S2 0.812 0.031 * –40.237 <0.001 ***
S3 0.568 0.307 –26.149 <0.001 ***
N3 N4 –1.123 0.001 *** 4.558 0.966
S1 –2.525 <0.001 *** –23.538 <0.001 ***
S2 –1.587 <0.001 *** –24.905 <0.001 ***
S3 –1.831 <0.001 *** –10.817 0.248
N4 S1 –1.403 <0.001 *** –28.096 <0.001 ***
S2 –0.464 0.567 –29.463 <0.001 ***
S3 –0.708 0.093 –15.375 0.021 *
S1 S2 0.938 0.007 ** –1.367 1.000
S3 0.694 0.107 12.721 0.101
S2 S3 –0.244 0.974 14.088 0.067
***p-value < 0.001, **p-value < 0.01, *p-value < 0.05; Week = the calendar week when the terminal buds appeared; Br % = the percentage browning of primary needles.
1

Fig. 1. Average a. timing of budset (Week) and b. frost hardiness (Br %) of first-year Scots pine seedlings from twenty-four cross-combinations presented as bar charts. Cross-combinations (columns) are grouped by the three maternal genotypes M1–M3 on the x-axis. Cross-combinations are labelled with the paternal genotype code. The letters of the paternal genotype codes denote the following: S = southern paternal genotype; N = northern paternal genotype; OP = southern pollen mixture from forests surrounding the orchard of the controlled crosses. Week denotes the calendar week when the terminal buds appeared and Br % denotes the percentage browning of primary needles. Error bars represent the 95% confidence intervals.

Table 5. Results from the linear regression analyses (Model 2). The analyses quantified the effect of the parental forest locality (Ldd) on the timing of budset (Week) and frost hardening (Br %) of first-year Scots pine seedlings. Model 2: Y = Intercept + Ldd, where Y = Week or Br %; Ldd = average effective temperature sum at the natural forest location.
Dependent variable: Week Dependent variable: Br %
ANOVA ANOVA
df F p-value df F p-value
Regression 1 136.104 <0.001 Regression 1 140.091 <0.001
Residual 42 Residual 42
Coefficients Coefficients
Model Unstandardized coefficients Model Unstandardized coefficients
B SE p-value B SE p-value
(Constant) 28.982 0.488 <0.001 (Constant) –103.522 13.597 <0.001
Ldd 0.0057 0.0005 <0.001 Ldd 0.160 0.013 <0.001
df = the number of degrees of freedom; F = F ratio; B = estimate; SE = estimate standard error; Week = the calendar week when the terminal buds appeared; Br % = the percentage browning of primary needles.
Table 6. Results from the ANCOVA tests (Model 3). Model 3 was applied to quantify the effect of the paternal genotype provenance on the timing of budset (Week) and autumn frost hardening (Br %) of controlled-cross Scots pine seedlings. Model 3: Y = Intercept + M + Pdd, where Y = Week or Br %; M = maternal genotype; Pdd = average effective temperature sum at the original location of the paternal genotype, which corresponds to the provenance of the paternal genotype.
Dependent variable: Week Dependent variable: Br %
Tests of between-subject effects Tests of between-subject effects
Source df F p-value Partial Eta Squared Source df F p-value Partial Eta Squared
Corrected Model 3 9.868 <0.001 0.270 Corrected Model 3 28.445 <0.001 0.526
Intercept 1 4644.29 <0.001 0.983 Intercept 1 5.529 0.021 0.067
Pdd 1 14.498 <0.001 0.153 Pdd 1 72.706 <0.001 0.486
Maternal genotype 2 7.553 0.001 0.159 Maternal genotype 2 6.598 0.002 0.146
Error 80 Error 77
Parameter estimates Parameter estimates
Parameter B SE p-value Parameter B SE p-value
Intercept 31.515 0.493 <0.001 Intercept 9.606 6.721 0.157
Pdd 0.002 0.0005 <0.001 Pdd 0.062 0.007 <0.001
df = the number of degrees of freedom; F = F ratio; Partial Eta Squared = SSEffect*(SSEffect + SSError)–1, SS = sum of squares; B = coefficient estimate; SE = estimate standard error; Week = the calendar week when the terminal buds appeared; Br % = the percentage browning of primary needles.
2

Fig. 2. Comparison of 95% confidence intervals of regression coefficients 0.5*βLdd and βPdd. Detection of the difference in the provenance effect between the Scots pine paternal genotypes of the natural forests and the paternal genotypes of the southern seed orchards by comparing the 95% confidence intervals of 0.5*βLdd and βPdd in the case of a. the timing of budset (Week) and b. frost hardening (Br %) of first-year seedlings. 0.5*βLdd denotes half of the regression coefficient of the independent variable Ldd from Model 2, where Ldd is the average effective temperature sum at the natural forest locality; βPdd denotes the regression coefficient of the covariate Pdd from Model 3, where Pdd is the average effective temperature sum at the original location of the seed orchard paternal genotype.