Documente Academic
Documente Profesional
Documente Cultură
http://journals.tubitak.gov.tr/agriculture/
Research Article
Accepted: 30.01.2013
Printed: 25.09.2013
Abstract: The existence of Quercus virgiliana Ten. is a subject of debate among botanists and silviculturists. It was considered a
distinct species, sometimes as an intraspecific taxonomic unit of Q. pubescens L., or was not even recognized as a taxon or infrataxon.
This disagreement with regard to the taxonomic classification is explained by the morphological similarities between Q. virgiliana
and Q. pubescens, with a small overlap between particular leaf and fruit traits. The main objective of this study was to evaluate the
macromorphological fruit and leaf descriptors in Romanian populations of pubescent oaks. We wanted to find which traits discriminate
between the 2 taxa. By using 7 microsatellite markers, we checked whether the 2 taxa have different genetic structures. A total of 918
individuals were sampled for morphological analyses from 20 stands across Romania. The length of cupula peduncle showed the highest
discriminating power, but also leaf characters such as abaxial laminar pubescence, lamina length, sinus width, and length of lamina
at largest width helped to discriminate between the 2 taxa. The analysis performed at 7 microsatellite loci revealed very little genetic
differentiation between Q. pubescens and Q. virgiliana. Different genetic assignment tests provided no support for 2 genetic entities in
our sample. This could be explained by the restricted number of loci used. However, the morphological differences found in this study
suggest that Q. virgiliana is an intraspecific taxonomic unit of Q. pubescens rather than a distinct species.
Key words: Fruit morphology, leaf morphology, microsatellites, Quercus pubescens, Quercus virgiliana
1. Introduction
Quercus virgiliana has been taxonomically classified
in many ways. According to different taxonomic
classifications, it is sometimes treated as a distinct species
(Schwarz 1993; ofletea and Curtu 2007; Doni 2008),
sometimes as an intraspecific taxonomic unit of Q.
pubescens, or sometimes it is not recognized or nominated
as a taxon or infrataxon (Camus 19361954; Nixon
1993; Bussotti and Grossini 1997; Menitsky, 2005). This
disagreement regarding the taxonomic classification of
the 2 taxa is derived from their morphological similarities,
with small overlaps between particular leaf traits (lamina
length, petiole length, number of lobes) and fruit
descriptors (peduncle length, cup scales shape), according
to various descriptions (Schwarz 1993; Doni 2008).
Among these morphological descriptors, the length of the
cupula peduncle, which is usually longer in Q. virgiliana, is
considered essential for differentiating the 2 oak taxa from
the series Lanuginosae (Bartha 2001; Trinajsti 2007). Most
analyses of leaf morphological characters in oaks from
the series Lanuginosae carried out in Europe could not
* Correspondence: nic.sofletea@unitbv.ro
632
633
assigned to
INT
VIR
Total
PUB
INT
VIR
Total
PUB
VIR
Total
PUB
Stand
Aiud
45
50
45
50
13
14
Bdeana
15
18
17
50
15
18
17
50
11
Berca
48
50
48
50
Breana Rocani
49
50
17
18
Ciucurova
15
16
15
16
Clisura Dunrii
14
14
Comanca
47
50
47
50
Cri
48
50
42
44
Deveselu
42
50
42
50
Dumbrveni
45
50
33
38
10
10
Grboavele
38
38
22
22
Hoia
37
50
37
50
Jugureni
50
50
Mcin
48
50
48
50
Mirslu
42
50
42
50
Peti
49
50
49
50
Rducneni
40
50
40
50
Sclaia
40
50
16
10
uncuiu
30
11
50
30
11
50
10
Vldila
45
50
45
50
Total
787
89
42
918
623
89
42
754
70
26
96
634
Beckman dye
Primer concentration
(uM)
Allele size
(bp)
ssrQpZAG112
di
D4
0.20
82112
ssrQpZAG96
di
D3
0.80
140180
ssrQpZAG11
di
D3
0.60
242289
ssrQpZAG110
di
D4
0.90
205243
ssrQpZAG87
di
D3
0.55
103183
ssrQpZAG7
di
D4
0.65
116157
ssrQpZAG20
di
D3
0.80
159213
Locus
635
Table 3. Mean and standard deviations of the 15 fruit and leaf descriptors for all sampled trees and separated for each taxon.
Q. pubescens (623 oaks)
4.2 0.9
3.7 1.1
4.2 0.86
4.0 0.7
3.9 0.7
4.0 0.71
Number of lobes
9.7 1.4
9.3 1.6
9.5 1.47
2.2 1.3
2.2 1.3
2.2 1.3
82.1 13.7
96.8 21.2
83.5 15.1
14.3 3.4
14.9 4.1
14.4 3.58
29.2 5.1
30.8 5.7
29.2 5.31
12.2 3.9
15.1 5.4
12.5 4.18
43.5 8.4
51.6 11.3
44.3 9.15
Lamina shape
52.9 5
53.5 4
53.0 5.06
Petiole ratio
14.8 2.9
13.5 3.1
14.8 3.02
57.5 12.2
50.8 13.4
56.6 12.27
Percentage venation
23.5 14.1
23.9 13.5
23.4 13.95
35.7 4.4
32.3 4.3
35.3 4.4
Descriptor
636
4.0 2.1
22.9 8
5.8 5.44
100
60
40
20
0
Breana Roscani
Cris
Macin
Berca
Petis
Deveselu
Vladila
Comanca
Ciucurova
Dumbraveni
Miraslau
Sacalaia
Raducaneni
Badeana
Garboavele
Aiud
Hoia
Suncuius
Linkage distance
80
Factor 2: 17.52%
4
2
0
2
4
6
4
Q . pubescens
Q. virgiliana
2
4
6
8
Factor 1: 24.77%
individuals with intermediate morphology
8
6
Factor 2: 17.94%
4
2
0
2
8
Q. pubescens
Q. virgiliana
2
Factor 1: 51.77%
637
ssrQpZAG96
ssrQpZAG110
ssrQpZAG11
ssrQpZAG87
ssrQpZAG7
ssrQpZAG20
Taxon
Na
Ne
Ho
He
Q. pubescens
14
5.8
11.18
0.857
0.828
Q. virgiliana
12
6.6
11.67
0.808
0.849
Q. pubescens
22
14.8
17.44
0.899
0.932
Q. virgiliana
15
11.2
14.87
0.84
0.91
Q. pubescens
22
14.88
0.825
0.833
Q. virgiliana
12
3.9
12
0.583
0.741
Q. pubescens
35
12.6
21.52
0.809
0.921
Q. virgiliana
21
15.2
20.71
0.64
0.934
Q. pubescens
13
3.9
9.02
0.729
0.744
Q. virgiliana
2.7
0.708
0.631
Q. pubescens
21
12.7
16.49
0.871
0.921
Q. virgiliana
16
11
15.79
0.88
0.909
Q. pubescens
18
7.1
13.32
0.87
0.859
Q. virgiliana
15
6.5
15
0.917
0.846
Mean Q. pubescens
21
14.79
0.837
0.862
Mean Q. virgiliana
14
13.86
0.768
0.832
17,3
8.5
14.32
0.802
0.847
Overall mean
Footnote: Na = number of alleles, Ne = effective number of alleles, A = allelic richness, Ho = observed heterozygosity, He
= expected heterozygosity.
638
4. Discussion
In general, the leaf morphological survey indicated that
there are small differences between the sampled stands
from Romania and others from the rest of Europe. With
respect to lamina length, it can be observed that the total
mean value for all 918 trees (83.5 mm) is higher than the
values reported in recent studies done on Q. pubescens
in Italy (Bruschi et al. 2000), Croatia (kvorc et al. 2005;
Franji et al. 2006), and Slovenia (Jere and Bati 2007).
In this context, the slightly larger size of leaves in some
stands from Romania could be due to the presence of the
42 Italian oaks and 89 intermediate individuals between
the 2 taxa in the sampled stands, or by the specific habitat
conditions at the northern natural distribution range of
pubescent oak. Kleinschmit (1993) reported the existence
of adaptive reactions in European oaks, which are driven
by environmental conditions. It was also reported that
the transition from glabrous to pubescent leaves is due to
the environmental conditions related mainly to soil water
content (Drake and Muller-Bombois 1993). Thus, more
densely pubescent oak individuals are generally located
in drier sites. This was also observed during our field
exploration. Smaller but hairier leaves were sampled in
drier sites.
Compared to the values from other studies (Dupouey
and Badeau 1993; Bruschi et al. 2000), the mean number
of lobes is smaller in our study. In contrast, the average
value that we obtained for petiole length is higher than
those reported in Croatia (kvork et al. 2005; Franji et
al. 2006) and Slovenia (Jere and Bati 2007). According
to Dupouey and Badeau (1993), the pilosity and number
of intercalary veins are very important discriminant
morphological descriptors between Q. robur, Q. petraea,
and Q. pubescens. In our study, the mean value for all stands
for number of intercalary veins is 2 times higher than that
in France (Dupouey and Badeau 1993), indicating a high
level of leaf irrigation at the northern natural distribution
range of Q. pubescens. Instead, the values found for the
length of cupula peduncle are smaller than those reported
for Q. pubescens from similar studies in France (Dupouey
and Badeau 1993) and Slovenia (Jere and Bati 2007).
In a similar study on leaf morphology of the Q. roburQ.
petraea complex (Ponton et al. 2004), it was shown that
leaf characters are not independent from each other as
shown by a positive correlation between them. In addition,
our findings suggest that there is also a positive correlation
between leaf dimensional characters and length of cupula
peduncle. However, taxonomic determinations, which use
a combination of characters (lamina length, lobe width,
petiole length, length of cupula peduncle) proved to be
informative, but other relevant characteristics should be
taken into account.
With regard to PCA, the lack of separation of Q.
virgiliana from Q. pubescens was also reported in Turkey
639
Acknowledgments
This study was supported by the Sectoral Operational
Program Human Resources Development (SOP HRD),
financed from the European Social Fund, and by the
Romanian Government under the contract number
POSDRU/88/1.5/S/59321 and by CNCS-TE-73-2010
project. We are grateful to Andras Tothpal and numerous
friends and colleagues from the Forest Districts for
supporting us during the fieldwork. We also thank Viorel
Roca, park director, and Bogdan Bjenariu, biologist at the
National Park Munii Mcinului, for allowing us to collect
samples of pubescent oak. The authors would like also to
thank the 2 anonymous reviewers for their constructive
comments, which helped to improve the manuscript.
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