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Silva Fennica 42(1)
research notes
www.metla.fi/silvafennica · ISSN 0037-5330
The Finnish Society of Forest Science · The Finnish Forest Research Institute
Differences in Leaf Morphology between
Quercus petraea
and
Q. robur
Adult and
Young Individuals
Adam Boratynski, Katarzyna Marcysiak, Amelia Lewandowska, Anna Jasinska,
Grzegorz Iszkulo and Jaroslaw Burczyk
Boratynski, A., Marcysiak, K., Lewandowska, A., Jasinska, A., Iszkulo, G. & Burczyk, J.
2008. Differ-
ences in leaf morphology between
Quercus petraea
and
Q. robur
adult and young individuals.
Silva Fennica 42(1): 115–124.
The characters of
Quercus robur
and
Q. petraea
leaves are of main taxonomic value and the
adult trees of both species can be distinguished on them. However, young individuals, mostly
seedlings but also saplings, are told to be undistinguishable or only partly distinguishable on
the leaf morphology. The aim of the study was to verify this hypothesis on the basis of bio-
metrical analyses of leaf characteristics of adults trees and saplings in two mixed oak woods,
one located close to the north-eastern limit, the other about 400 km inside of the
Q. petraea
range in Poland. The analysis of discriminations and minimum spanning tree on the squares
of Mahalanobis distances were analysed to find differences between
Q. robur, Q. petraea
and intermediate adults and saplings. The differences between saplings of
Q. robur
and
Q.
petraea
were found lower than between adult trees. Nevertheless, the biometrical analysis
confirmed determination of saplings in the field.
Keywords
morphological diversity, plant variation, biometry,
Quercus,
leaf polymorphism
Addresses
Boratynski, Jasinska
and
Iszkulo,
Polish Academy of Sciences, Institute of Dendrol-
ogy, 5 Parkowa str., 62-035 Kórnik, Poland;
Marcysiak, Lewandowska
and
Burczyk,
Kazimierz
Wielki University, Institute of Biology and Environment Protection, 12 Ossolinskich str.,
85-064 Bydgoszcz, Poland
E-mail
borata@man.poznan.pl
Received
13 February 2007
Revised
29 October 2007
Accepted
20 November 2007
Available at
http://www.metla.fi/silvafennica/full/sf42/sf421115.pdf
115
Silva Fennica 42(1), 2008
research notes
1 Introduction
Pedunculate oak
– Quercus robur
L. and sessile
oak –
Q. petraea
(Matt.) Liebl. are well dis-
tinguishable on the morphological characters of
their leaves, acorns and cupulas (Kotschy 1862,
Schwarz 1937, Krahl-Urban 1959, Schwarz 1964,
Rushton 1983, Amaral Franco 1990, Bacilieri et
al. 1995). The leaves, as the most easily accessible
parts of the trees and easily measured with utiliza-
tion of electronic accessories and software have
been recently used with high success in exten-
sive biometrical study through Western Europe
(Dupouey 1983, Kleinschmit et al. 1995, Kremer
et al. 2002). However, the seedlings and even
saplings are frequently told to be very similar
and quite undistinguishable on the leaf characters
(Aas 1993, Rushton 1993, Kremer et al. 2002,
Boratynska et al. 2006).
The aim of the present study was the biometri-
cal verification of correctness of in the field deter-
mination of adult trees and about 12–18 years old
saplings, which appeared spontaneously under
canopy of two different, adult mixed stands of
Q.
robur
and
Q. petraea
in Poland. The white oaks
hybridize frequently (Bacilieri et al. 1996, Dodd
and Afzal-Rafii 2004, Boratynska et al. 2006,
Giertych 2006). The reviewed biometrical works
have been performed predominantly in the West-
ern Europe, partly within range of
Q. pubescens
(Meusel et al. 1965, Jalas and Suominen 1976,
Dupont 1990, Boratynski et al. 2006), which
could also influence the variation of local popula-
tions of
Q. robur
and
Q. petraea.
Central Europe
is outside the ranges of oak species other than
Q.
robur
and
Q. petraea,
which makes the compara-
tive morphological study much simpler.
2 Material and Methods
2.1 Plant Material and Field Works
The two seed collection stands distinguished for
forest gene conservation and seed production
(Matras 1996) were selected in: 1) Jamy and
2) Legnica Forest Offices (Table 1). The stands
have been thinned as a result of oak decay during
1980–1990 and the cutting down of adult trees
116
and saplings of
Fagus sylvatica
and
Carpinus
betulus
(unpublished data of Jamy and Legnica
Forest Offices). Natural regeneration appeared
in both stands, in Jamy as a result of elimination
of competition of other tree species and herbs by
soil preparation and in Legnica spontaneously.
The saplings in the Jamy stand were formed over
85% of the stand area in 1986. The dead oak trees
were removed in subsequent years and the first
selective clearings were made in 2000, removing
mainly saplings of
Fagus sylvatica, Acer pseudo-
platanus
and other tree species (personal com-
munications of foresters). The stand in Legnica
had not been prepared for self-sowing but, in
spite of that, natural regeneration was abundant.
The site conditions of both stands are similar
(unpublished data of Forest Offices). Both stands
are mixed with similar participation of
Q. robur
and
Q. petraea
and only inconspicuous addition
of hybrids. The form of mixing is random, stem
by stem or rarely 3–4-tree clusters.
The taxonomic status of adult trees was deter-
mined over the entire area of both stands. Leaf,
acorn and cupula morphology were applied
during determination in the field by visual assess-
ment. Every tree was subsequently assigned into
one of the three categories: 1)
Q. robur,
2)
Q.
petraea
and 3) hybrid (intermediate morphol-
ogy). The trees of latter category have been dis-
tinguished on the basis of intermediate length
of leaf petiole, the shape of leaf blade, the pres-
ence/absence of nerves between leaf lobes (Aas
1993, Kremer et al. 2002), and length of cupula
peduncle (Schwartz 1937, 1964, Boratynska et
al. 2006). The determination was conducted in
September, when acorns with cupulas were well
visible using binoculars. The trees with not stable
characters (see Kremer et al. 2002: 783 and 784,
as unclassified) were only scarcely represented
in both stands (Table 1). Then plots 120 m long
and 40 m wide were established in central parts
of each stand. The 59 trees in the Jamy and 95
in the Legnica stands (Table 2) were sampled on
the plots for the biometrical verifications. Ten
leaves were collected from central parts of long
shoots, from south-facing, insolated parts of the
crown of every tree, at an altitude of 8–9 m. The
leaves from the central part of the first spring
longitudinal increment of shoots were used, as
the most typically developed (Staszkiewicz 1970,
Boratynski et al.
Differences in Leaf Morphology between
Quercus petraea
and
Q. robur
Adult and Young Individuals
Table 1.
Sampled stands composed of
Quercus robur
and
Q. petraea.
Locality
Geographic
coordinates
Abbreviation
Age
Area
[ha]
Participation of taxa
[%]
Q. robur
Q. petraea
Hybrids
Jamy Forest Division,
Forestry Jamy, no. 96c
Legnica Forest Division,
Forestry Karczewiska, no. 315d
E18°53´
N53°35´
E16°10´
N51°19´
Jamy
Legnica
125
154
5.00
10.83
57.0
53.5
42.2
44.4
0.8
2.1
Table 2.
Adult trees and saplings of
Quercus robur,
Q. petraea
and intermediate individuals on the
analysed plots and transects in the stands of Jamy
and Legnica.
Taxon
Stand
Number of individuals
Trees
biometrically
analysed on
plots
Saplings
biometrically
analysed on
transects
Table 3.
Characters of leaves analyzed biometrically.
Number of
character
Character
Q. robur
Q. petraea
Intermediate
Q. robur
Q. petraea
Intermediate
Jamy
Legnica
23
33
3
35
54
6
27
46
27
10
79
9
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
Aas 1993, Rushton 1993, Kremer et al. 2002,
Boratynska et al. 2006, Chałupka 2006). A total
of 1550 leaves were measured and analysed.
The saplings, individuals higher than 0.3 m
(Harmer 2001), growing at a distance of about
1 m from each other were determined. Then,
ten leaves from every sapling were collected for
biometrical verification in every stand. The 100
saplings were sampled in Jamy and 98 in Legnica
stand. A total of 1980 leaves from saplings were
biometrically compared (Table 2).
2.2 Leaf Biometry
Leaves were analysed biometrically using 16
measured, 1 evaluated and 7 synthetic characters
(Table 3), according to the methods used earlier
by Kremer et al. (2002) and Borazan and Babac
(2003).
Area of leaf blade (A)
Circumference of leaf blade (P)
Length of leaf blade (LL)
Maximum width of leaf blade (LW)
Width of leaf blade in mid-length
Width of leaf blade in 90% of length
Petiole length (PL)
Area of petiole
Length of apical lobe
Width of apical lobe
Length of the longest side lobe (a)
Length of lobe below the longest side
lobe (b)
Depth of sinus between lobes a and b
Length of the second side nerve
Number of lobes (NL)
Number of nerves between the lobes (NV)
Type of leaf base (BS)
PR = 100 × PL/(LL + PL)
PV = 100 × NV/NL
LWR = 100 × LW/LL
100 × (Width of leaf blade at 90% of
length/maximum width)
100 × (Width of leaf blade at mid-length/
maximum width)
100 × (Width of leaf blade at 90% of
length/width of leaf blade at mid-length)
AP = A/P
The distributions of character values were
verified using Shapiro-Wilks’ test and frequency
histograms. The average values of particular char-
acters were calculated separately for subpopula-
tions of adult and young individuals of each taxon
117
Silva Fennica 42(1), 2008
research notes
distinguished in each stand to find their taxonomic
importance. The similarity/dissimilarity among
adult and young subpopulations of each taxon
in each stand was determined using discrimina-
tion analysis (Zar 1999, Sokal and Rohlf 2003).
The latter was performed on the evaluated (17)
and synthetic (18–24) characters only, to avoid a
possible influence of environmental differences
(Kremer et al. 2002). The square of the shortest
Mahalanobis’ distances between distinguished
subpopulations of adult trees and saplings of
Q.
robur, Q. petraea
and hybrids has been analysed
for each stand separately to find the relationships
between them (Sokal and Rohlf 2003).
The leaves were measured with a scanner and
WinSeedle software (Regent Inc.). Statistical
analyses were performed using STATISTICA 6
software (StatSoft).
3 Results
Most of the leaf characters (Table 3) presented
the unimodal frequency distribution. Only length
and area of the leaf petiole (characters 7 and 8),
the length of the longest side lobe, number of
veins between lobes, ratio of petiole length to
leaf length and ratio of width of leaf blade at
mid-length to maximal width (characters 11, 16,
18 and 22, respectively) have shown a slightly
biased frequency distribution.
The character values were found to be pre-
dominantly higher for adult trees than for sap-
lings. Additionally, the measured characters had
generally higher values in the Legnica than in
the Jamy stands (Table 4). In some cases aver-
age values of characters for adult trees from
Jamy were similar to the sapling characteristics
in the Legnica population, as for example in the
area of leaf blade, maximal width of leaf blade,
width of leaf blade in mid-length, width of leaf
blade in 90% of length, length of the longest lobe
and lobe below the longest side lobe (characters
1, 4, 5, 6, 11 and 12, respectively) (Table 4). It
blurred the differences between species and made
difficult the analyses and hybrid identification
in the field. Nevertheless, the hybrid adult trees
had values of particular characters intermediate
between
Q. robur
and
Q. petraea
adult trees, and
118
hybrid saplings intermediate between of
Q. robur
and
Q. petraea
saplings. It concerns mostly such
characters as length and area of petiole, length of
lobe below the longest side lobe, depth of sinus
between lobes a and b, length of second side vein,
number of lobes, number of veins between lobes,
and type of leaf base (characters 7, 8, 12, 13, 14,
15, 16 and 17, respectively, see Table 4).
The petiole length, number of veins between
lobes, type of leaf blade base, ratio of length
of petiole to leaf length and number of veins to
number of lobes (character 7, 16, 17, 18 and 19,
respectively) differed between taxa. The length
of petiole (character 7) had significantly higher
values for leaves of adult trees of
Q. petraea
and
hybrids. The leaves of
Q. robur
had a higher
number of veins between lobes (character 16)
than of
Q. petraea,
independently of age. The
leaves of adult
Q. robur
had a typically auriculate
type of blade base and the highest value of this
character (17). The ratio of petiole to leaf length
(character 18) had the highest values in the group
of adult trees of
Q. petraea
and the intermedi-
ate individuals. The ratio of number of veins to
number of lobes (character 19) had the highest
values for adults of
Q. robur
and the lowest for
Q. petraea.
The average values of some characters dif-
fered between populations and between adult
trees and saplings within populations (Table 4).
The number of characters differing
Q. robur
and
Q. petraea
at a statistically significant level is
higher for subpopulations of adults.
Q. robur
differed statistically significantly from
Q. petraea
in both, population and age categories
in length of petiole, width of apical lobe, depth of
sinus between longest and lying below side lobes,
number of veins between lobes, proportion of pet-
iole length to leaf length, ratio of number of veins
between side lobes to number of side lobes and
proportion of width of leaf blade at mid-length to
maximal width (characters 7, 10, 13, 16, 18, 19
and 22). More characters of adults differentiated
significantly between species in the Legnica than
in the Jamy stand. Of particular interest was the
fact that the type of leaf blade base, used as one
of the most important traits, differed statistically
significantly only between adult trees.
The adult trees of
Q. robur
and
Q. petraea
were
the most morphologically distant in the discrimi-
Boratynski et al.
Table 4.
Average values of characters of
Q. robur, Q. petraea
and hybrid subpopulations of mother trees and saplings in Jamy and Legnica stands; 1–24 character
numbers as in Table 3; P-JD –
Q. petraea
mother trees in Jamy stand; R-JD –
Q. robur
mother trees in Jamy stand; X-JD – hybrid mother trees in Jamy stand,
P-JM –
Q. petraea
saplings in Jamy; R-JM –
Q. robur
saplings in Jamy; X-JM – hybrid saplings in Jamy; P-LD –
Q. petraea
mother trees in Legnica; R-LD
–
Q. robur
mother trees in Legnica; X-LD – hybrid mother trees in Legnica; P-LM –
Q. petraea
saplings in Legnica; R-LM –
Q. robur
in Legnica; X-LM
– hybrid saplings in Legnica.
Character number
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
Sub-
population
1
2
3
4
Differences in Leaf Morphology between
Quercus petraea
and
Q. robur
Adult and Young Individuals
P-JD
R-JD
X-JD
P-JM
R-JM
X-JM
P-LD
R-LD
X-LD
P-LM
R-LM
X-LM
36.9 37.2
38.3 40.1
36.8 44.5
26.7 31.9
25.7 33.4
29.3 33.4
57.2 41.9
54.3 43.8
62.9 53.2
40.8 35.4
44.3 37.8
37.3 36.4
9.8
10.6
9.3
9.3
9.7
10.0
11.6
12.0
12.7
10.2
10.8
10.0
5.65 4.89 2.48 1.65 0.20 0.64 1.13 3.99 3.34 2.31 1.33 5.93 0.49 3.87 14.28 8.8 58.0 44.0 86.6 52.0
5.86 4.75 2.74 0.86 0.11 0.82 1.38 4.00 2.96 1.73 0.99 5.09 2.49 7.48 7.43 50.8 56.2 46.9 81.3 59.5
6.15 5.33 3.04 1.44 0.18 0.59 1.06 4.45 3.37 2.05 1.07 5.17 1.04 4.35 13.08 21.1 66.1 49.1 86.0 58.4
4.59 3.89 2.18 0.86 0.10 0.45 0.83 3.24 2.62 1.85 1.03 6.15 0.82 2.67 8.43 14.3 49.9 47.3 84.7 57.8
4.51 3.60 2.17 0.66 0.08 0.58 0.93 3.14 2.29 1.39 0.92 5.97 2.27 3.42 6.34 39.3 46.8 47.8 80.1 62.0
4.82 3.95 2.32 0.73 0.09 0.57 1.01 3.36 2.61 1.68 1.03 6.15 1.66 2.68 6.82 28.6 48.5 47.8 81.7 60.2
7.41 6.67 3.33 2.01 0.39 0.52 1.15 4.85 4.09 2.93 1.28 6.84 0.39 3.69 14.87 5.8 63.9 44.9 90.0 50.4
7.34 6.25 3.67 1.04 0.14 0.59 1.29 4.80 3.83 2.52 1.10 5.92 2.68 7.00 8.02 45.8 61.1 49.9 85.0 59.6
8.11 7.16 3.35 2.01 0.27 0.48 1.25 4.48 3.81 2.71 1.14 6.69 1.71 3.34 13.79 27.1 63.7 41.0 88.4 47.5
6.16 5.42 2.66 1.23 0.14 0.50 0.94 4.18 3.45 2.54 1.20 6.58 0.38 2.57 10.73 6.1 60.1 42.9 87.8 49.4
6.36 5.42 2.94 0.94 0.11 0.57 1.08 4.31 3.32 2.31 1.09 6.18 1.33 3.30 7.85 23.7 58.2 46.1 84.9 55.3
6.10 5.40 2.52 0.99 0.11 0.56 1.02 4.15 3.15 2.17 1.08 6.29 0.94 2.55 8.93 15.3 60.9 41.0 88.6 47.1
0.97
0.93
0.81
0.82
0.75
0.85
1.34
1.21
1.15
1.11
1.10
1.00
119

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