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Molecular phylogeny of Acer monspessulanum L. subspecies from Iran inferred using the ITS region of nuclear ribosomal DNA

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Khademi H, Mehregan I, Assadi M, Nejadsatari T, Zarre S. 2015. Molecular phylogeny of Acer monspessulanum L. subspecies from Iran inferred using the ITS region of nuclear ribosomal DNA. Biodiversitas 17: 16-23. This study was carried out on the Acer monspessulanum complex growing wild in Iran. Internal transcribed spacer (ITS) sequences for 75 samples representing five different subspecies of Acer monspessulanum were analyzed. Beside this, 86 previously published ITS sequences from GenBank were used to test the monophyly of the complex worldwide. Phylogenetic analyses were conducted using Bayesian inference and maximum parsimony. The results indicate that most samples of A. monspessulanum species from Iran were part of a monophyletic clade with 8 samples of A. ibericum from Georgia, A. hyrcanum from Iran and one of A. sempervirens from Greece (PP= 1; BS= 79%). Our results indicate that use of morphological characteristics coupled with molecular data will be most effective.
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Molecular phylogeny of Acer monspessulanum L. subspecies from Iran inferred using the ITS region of nuclear ribosomal D
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B I O D I V E R S IT A S
ISSN: 1412-033X
Volume 17, Number 1, April 2016 E-ISSN: 2085-4722
Pages: 16-23 DOI: 10.13057/biodiv/d170103
Molecular phylogeny of Acer monspessulanum L. subspecies from Iran
inferred using the ITS region of nuclear ribosomal DNA
HANIF KHADEMI1, IRAJ MEHREGAN1,, MOSTAFA ASSADI2, TAHER NEJADSATARI1,
SHAHIN ZARRE3
1Department of Biology, Science and Research Branch, Islamic Azad University, Hesarak-1477893855, Tehran. Iran, email: iraj@daad-alumni.de
2Research Institute of Forest and Rangelands, National Botanical Garden of Iran, P.O. Box 13185-116, Tehran, Iran
3Department of Plant Sciences, School of Biology, College of Science, University of Tehran, P.O. Box 14155-6455, Tehran, Iran
Manuscript received: 29 September 2015. Revision accepted: 16 November 2015.
Abstract.Khademi H, Mehregan I, Assadi M, Nejadsatari T, Zarre S. 2015. Molecular phylogeny of Acer monspessulanum L.
subspecies from Iran inferred using the ITS region of nuclear ribosomal DNA. Biodiversitas 17: 16-23. This study was carried out on
the Acer monspessulanum complex growing wild in Iran. Internal transcribed spacer (ITS) sequences for 75 samples representing five
different subspecies of Acer monspessulanum were analyzed. Beside this, 86 previously published ITS sequences from GenBank were
used to test the monophyly of the complex worldwide. Phylogenetic analyses were conducted using Bayesian inference and maximum
parsimony. The results indicate that most samples of A. monspessulanum species from Iran were part of a monophyletic clade with 8
samples of A. ibericum from Georgia, A. hyrcanum from Iran and one of A. sempervirens from Greece (PP= 1; BS= 79%). Our results
indicate that use of morphological characteristics coupled with molecular data will be most effective.
Keywords: Biogeography, maple, phylogeny, Sapindaceae
INTRODUCTION
The genus Acer L. is a member of Sapindaceae that
mainly grows in tropical and subtropical regions. This
genus is one of the most diverse tree genera in the Northern
Hemisphere with approximately 129 species and is the largest
tree genus of the northern hemisphere besides Quercus
(Fang 1966; Grimm et al. 2006). A number of characteristics
are shared among all members of Acer. The arrangement of
the two winged pericarps (samaras) in the fruits ranges from
subparallel to diverging at about a right angle.
A number of characters that are beneficial for
identification of species in the field may have evolved
independently (lobe shape, margin of lobes). These traits
may be highly variable within species like pubescence of
lower leaf surface (Grimm et al. 2007). Several different
florescence types, including racemes, panicles, corymbs
and spikes, occur in this genus. These variations make
infrageneric divisions very difficult. Species delimitation
and phylogenic relationships within the genus Acer are also
very controversial (Kholie 1967; Judd et al. 2002). Fang
(1966) proposed a different system in which the genus was
divided into two subgenera, mainly on the basis of simple
versus compound leaves (Koidzumi 1911). In Ogata’s
system (Ogata 1967), the genus was classified into 26
sections (Momotani 1962). In 1970, Murray published his
monograph of the Aceraceae with 7 subgenera, 24 sections
and 35 series within Acer (Murray 1970). Ogata’s system
was essentially followed by Xu (1966), with some
additions and amendments. More recently de Jong (1994)
recognized only 19 series in 16 sections, providing a quite
different arrangement from those of other authors (Pax
1902; Xu 1966, 1998; Ogata 1967; Xu et al. 2008). Some
researchers discussed the infrageneric phylogenetic
relationships in the genus by analyzing gross morphology,
seed proteins, fossils and geographic distributions, but the
conclusions were not in consensus (Momotani 1962;
Rechinger 1969; Pax 1985, 1986; Wolfe and Yanai 1987;
Thorne 1992).
Acer monspessulanum is a medium-sized deciduous
tree or densely branched shrub that grows to a height of 10-
15 m (rarely to 20 m) (Fontaine 2011).The trunk is up to 75
cm diameter, with smooth, dark grey bark on young trees,
becoming finely fissured on old trees. Among similar
maples is most easily distinguished by its small three-lobed
leaves, 3-6 cm long and 3-7 cm wide, glossy dark green,
sometimes a bit leathery, and with a smooth margin, with a
2-5 cm petiole. The leaves fall very late in autumn,
typically in November. The flowers are produced in spring,
in pendulous, yellow to white corymbs 2-3 cm long. The
samaras are 2-3 cm long with rounded nutlets (Rushforth
1999; van Gelderen and van Gelderen 1999).
Acer monspessulanum from Acer section (van Gelderen
et al. 1994) has distinct small, 3-lobed leaves, while its
close relatives A.hyrcanum and A.opalus normally have 5-
lobed leaves. Acer ibericum displays dimorphic leaves that
are 5-lobed in juvenile plants and sucker shoots, and 3-
lobed in older plants (Grimm et al. 2007). Acer
monspessulanum fossils are fairly common in Late
Miocene and Pliocene floras from southern Europe and
south-western Asia (Kvacek et al. 2002; Sachse 2004).
The ITS is highly variable nuclear region suitable for
phylogenetic reconstruction of closely related taxa. The
utility of this marker has already been investigated in other
KHADEMI et al. –Molecular phylogeny of Acer monspessulanum
17
plant groups including trees, e.g. Acer (Tian et al. 2002;
Grimm et al. 2006; Grimm et al. 2007) and Crataegus (Zarrei
et al. 2014, 2015) and bulbs (Zarrei et al. 2009).
The aim of this study is to clarify taxonomy and to
delimit Acer monspessulanum subspecies that grow in Iran
using ITS marker and comparing results with
morphological traits.
MATERIALS AND METHODS
Plant samples
The ITS sequencing was performed on 75 individuals
from 15 populations of Acer monspessulanum distributed
in Iran. Population name, localities, altitude, and herbarium
number for each population are shown in Table 1. The
plant specimens were identified in the Department of
Biology, Science and Research Branch of Islamic Azad
University in Tehran, by the aid of local and regional
Floras, and voucher specimens of the plants with numbers
14821-14835 were deposited in the IAUH (Table 1). The
specimens were collected during July and December 2014.
Wherever possible, five trees from at least 50 m distant
from each other were sampled randomly from each
population. Fresh leaves were collected and kept in 50 CC
falcon tubes, filled with Silica Gel, for the purpose of
drying them (Chase and Hill 1991). The leaves were then
used as a DNA extraction source.
DNA extractions and ITS amplification
Total DNA was extracted following a modified CTAB
protocol of Doyle and Doyle (1990) using the DNeasy
Plant Mini kit (Qiagen, Germany). We amplified the
Internal Transcribed Spacer region (ITS1-5.8S-ITS2) of the
nuclear ribosomal DNA using primer combinations 18S
(forward primer 5'-CCT TMT CAT YTA GAG GAA GGA
G-3') and 28S (reverse primer 5'-CCG CTT ATT KAT
ATG CTT AAA-3'). The PCR protocol for ITS region
included: 34 cycles of 18 seconds denaturation (94°C), 30
seconds annealing (53°C), and 60 seconds elongation (72°
C), with two additional minutes elongation (Gaskin and
Schaal 2003). The quality of PCR products was checked by
electrophoresis on a 1.0% agarose gel and then visualized
under UV light.
Table 1. List of Acer monspessulanum subspecies investigated in our analysis and their morphological characters and locality in Iran
(small= up to 2×2 cm, large=2-3 × 3.5-4 cm)
Taxon
Locality with herbarium numbers and
GenBank accession numbers
Major features of
morphological traits(Rechinger
1969)
A. monspessulanum ssp.
turcomanicum (Pojark.) Rech.
f.
Iran: Khorasan Shomali, 45 km N of Shirvan, Golul-Sarani,
2302 m, Basiri 14823 (IAUH)
Leaves: large
Loculus inside: hairy
Loculus outside: sparsely hairy
A. monspessulanum ssp.
ibericum (M.B.) Yaltirik
Iran: Azarbayejan Sharghi, Kaleybar, Arasbaran forest, Venigh,
1070 m, Masoud, 14821 (IAUH), KT587662
A. monspessulanum ssp.
ibericum (M.B.) Yaltirik
Iran: Azarbayejansharghi, Kaleybar, Arasbaran forest, Tuali, 850
m, Masoud, 14822 (IAUH), KT587663
A. monspessulanum ssp.
Ibericum (M.B.) Yaltirik
Iran: Golestan, Gorgan, Golestan National Park, 677 m,
Khademi, 14833 (IAUH), KT587665
A. monspessulanum ssp.
ibericum (M.B.) Yaltirik
Iran: Mazandaran, Amol, Haraz road, Chelav, 737 m, Khademi,
14834 (IAUH), KT587661
Leaves: large
Loculus inside: hairy
Loculus outside: glabrous
Lower surface midrib: glabrous
A. monspessulanum ssp.
assyriacum (Pojark.) Rech.
Iran: Kordestan, Mariwan, Mohhamadeh village toward
Benavechele, 1550 m, Khademi, 14828 (IAUH), KT587655
A. monspessulanum ssp.
assyriacum (Pojark.) Rech.
Iran: Kordestan, Mariwan, Mohhamadeh village toward
Benavechele, 1510 m, Khademi, 14829 (IAUH), KT587653
A. monspessulanum ssp.
assyriacum (Pojark.) Rech.
Iran: Kermanshah, Jawanroud toward Salas, 1585 m, Khademi,
14832 (IAUH), KT587654
Leaves: large
Loculus inside: hairy
Loculus outside: glabrous
Lower surface midrib: sparsely
hairy
A. monspessulanum ssp.
cinerascens (Boiss.) Yaltirik
Iran: Fars, Marwdasht, Jahanabad village, 1756 m, Khademi,
14824 (IAUH), KT587656
A. monspessulanum ssp.
cinerascens (Boiss.) Yaltirik
Iran: Fars, Marwdasht, Bizjan village, Dorodzan Dam, 1715 m,
Khademi, 14825 (IAUH), KT587657
A. monspessulanum ssp.
cinerascens (Boiss.) Yaltirik
Iran: Fars, Marwdasht, Chav road, 1823 m, Khademi, 14826
(IAUH). KT587658
A. monspessulanum ssp.
cinerascens (Boiss.) Yaltirik
Iran: Fars, Bayza. Tang Tir forest, 1632 m, Khademi, 14827
(IAUH), KT587659
A. monspessulanum ssp.
cinerascens (Boiss.) Yaltirik
Iran: Kohgiloye-va-Boir Ahmad, Gachsaran, Gachsaran, 15 km
to Choram, After Abrigoon, Deel neck, 1600 m, Mehrgan, 14835
(IAUH)
Leaves: small
Loculus inside: densely hairy
Loculus outside: glabrous
A. monspessulanum ssp.
persicum (Pojark.) Rech.
Iran: Kerman, 25 km from Dalfard toward Jiroft, 980 m,
Meyjani, 14830 (IAUH)
A. monspessulanum ssp.
persicum (Pojark.) Rech.
Iran: Kerman, Meyjan, 1218 m, Meyjani, 14831 (IAUH),
KT587664
Leaves: small
Loculus inside: glabrous
Loculus outside: sparsely hairy
B I O D I V E R S IT A S
17 (1): 16-23, April 2016
18
Table 2. List of taxa used in our analysis with their GenBank
accession numbers.
Region
Taxon
Iran
A. hyrcanum ssp. hyrcanum
Iran
A. hyrcanum ssp. hyrcanum
Iran
A. hyrcanum ssp. hyrcanum
Iran
A. hyrcanum ssp. hyrcanum
Georgia
A. ibericum
Georgia
A. ibericum
Georgia
A. ibericum
Georgia
A. ibericum
Georgia
A. ibericum
Georgia
A. ibericum
Georgia
A. ibericum
Georgia
A. ibericum
Georgia
A. ibericum
Georgia
A. ibericum
Georgia
A. ibericum
France
A. monspessulanum
France
A. monspessulanum
France
A. monspessulanum
France
A. monspessulanum
France
A. monspessulanum
France
A. monspessulanum
France
A. monspessulanum
France
A. monspessulanum
France
A. monspessulanum
France
A. monspessulanum
Bulgaria
A. monspessulanum
Bulgaria
A. monspessulanum
Bulgaria
A. monspessulanum
Bulgaria
A. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
Spain
A. monspessulanum ssp. monspessulanum
Spain
A. monspessulanum ssp. monspessulanum
Spain
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
France
A. monspessulanum ssp. monspessulanum
AM238384
France
A. monspessulanum ssp. monspessulanum
AM238385
France
A. monspessulanum ssp. monspessulanum
AM238386
France
A. monspessulanum ssp. monspessulanum
AM238387
France
A. monspessulanum ssp. monspessulanum
AM238388
France
A. monspessulanum ssp. monspessulanum
AM238391
France
A. monspessulanum ssp. monspessulanum
AM238393
France
A. monspessulanum ssp. monspessulanum
AM238394
France
A. monspessulanum ssp. monspessulanum
AM238395
France
A. monspessulanum ssp. monspessulanum
AM238396
France
A. monspessulanum ssp. monspessulanum
AM238397
France
A. monspessulanum ssp. monspessulanum
AM238398
France
A. monspessulanum ssp. monspessulanum
AM238399
France
A. monspessulanum ssp. monspessulanum
AM238401
Germany
A. monspessulanum ssp. monspessulanum
AM238402
Germany
A. monspessulanum ssp. monspessulanum
AY605315
Spain
A. monspessulanum ssp. monspessulanum
AY605316
Spain
A. monspessulanum ssp. monspessulanum
AY605317
Spain
A. monspessulanum ssp. monspessulanum
AY605318
France
A. monspessulanum ssp. monspessulanum
AY605319
France
A. monspessulanum ssp. monspessulanum
AY605320
France
A. monspessulanum ssp. monspessulanum
AY605321
France
A. monspessulanum ssp. monspessulanum
DQ366124
France
A. monspessulanum ssp. monspessulanum
DQ366125
France
A. monspessulanum ssp. monspessulanum
DQ366126
France
A. monspessulanum ssp. monspessulanum
DQ366127
France
A. monspessulanum ssp. monspessulanum
DQ366128
France
A. monspessulanum ssp. monspessulanum
AM238327
Cyprus
A. obtusifolium
AM238328
Cyprus
A. obtusifolium
AM238329
Cyprus
A. obtusifolium
AM238330
Cyprus
A. obtusifolium
AM238331
Syria
A. obtusifolium
AM238332
Syria
A. obtusifolium
AM238417
France
A. opalus
AM238418
France
A. opalus
AM238419
France
A. opalus
AM238420
France
A. opalus
AM238421
France
A. opalus
AM238422
France
A. opalus
AM238334
Greece
A. sempervirens
AM238335
Greece
A. sempervirens
AM238336
Greece
A. sempervirens
AM238337
Greece
A. sempervirens
AM238340
Greece
A. sempervirens
AM238341
Greece
A. sempervirens
AM238342
Greece
A. sempervirens
AM238343
Greece
A. sempervirens
AM238344
Greece
A. sempervirens
AM238348
Greece
A. sempervirens
AM238349
Greece
A. sempervirens
AM238350
Greece
A. sempervirens
AM238351
Greece
A. sempervirens
AY605349
Greece
A. sempervirens
AY605350
Greece
A. sempervirens
AY605351
Greece
A. sempervirens
AY605352
Greece
A. sempervirens
AY605353
Greece
A. sempervirens
DQ366122
Greece
A. sempervirens
DQ366123
Greece
A. sempervirens
AY605361
Iran
A. velutinum
Note: ssp. = subsp. = sub species
KHADEMI et al. –Molecular phylogeny of Acer monspessulanum
19
Phylogenetic analyses
Phylogenetic reconstructions were performed with 15
samples from each of 75 accessions (15 populations)
belonging to five subspecies of Acer monspessulanum from
Iran (Table 1). In addition, we used the ITS sequence of 86
accessions of Acer from GenBank. List of non-Iranian taxa
used in our analysis with GenBank accession numbers are
shown in Table 2. We used ITS sequences of Acer
velutinum Boiss. from GenBank as the outgroup based on
the earlier studies including Grimm et al. (2006) (Table 2).
The 3' region of the 18S rDNA, the 5' region of the 26S
rDNA, and the whole ITS1-5.8S rDNA-ITS2 region were
sequenced for all the taxa, and these were compared to
sequences produced for other maples. Forward and reverse
sequences were visually compared and edited, and then
aligned using Sequencher 4 software (Gene Codes
Corporation, Ann Arbor, MI, USA). In addition to our
sequences, 86 ITS sequences from other taxa were taken
from GenBank (Table 2). All ITS sequences were
assembled and aligned using MacClade 4 (Maddison and
Maddison 2005). The parsimony analyses were performed
using PAUP*4.0b10 (Swofford 2002), with the following
options: heuristic search with 1,000 random-addition-
sequence replicates; tree bisection-reconnection (TBR)
branch swapping; saving all most parsimonious trees.
Character state changes were treated as equally weighted.
Relative clade support was estimated using 1,000 bootstrap
replicates in PAUP* via full heuristic searches and simple
taxon addition. Clades with a bootstrap value of 50% or
more were considered as robustly supported nodes. The
consistency index (CI) and retention index (RI) were
calculated to assess the amount of homoplasy present in the
data. The best-fitting substitution model (TrN+I) was
determined under the Akaike Information Criterion (AIC;
Akaike 1974) using Modeltest 3.7 (Posada and Crandall
1998). The Bayesian analysis (BA) of the ITS datasets
were performed using MrBayes v3.1.2 (Huelsenbeck and
Ronquist 2001). TrN+I is a transitional model with six
rates. For the ITS dataset, the TrN+I model was chosen.
The amount of proportion of invariable sites(I) was 0.6732.
RESULTS AND DISCUSSION
The data set of the ITS region included 675 characters,
43 of them parsimony informative. Strict consensus tree
(length of 138 steps, consistency index (CI) = 0.703,
retention index (RI) = 0.904) is shown in Figure 1. Figure 2
shows tree from Bayesian analysis using MrBayes. All
sampled species of Iranian Acer were part of a
monophyletic clade with Posterior Probability (PP) = 1 and
Bootstrap Support (BS) = 79% (Clade N; Figure 1). Since
Iranian A. monspessulanum origin are from Mediterranean
(Rechinger 1969), we compare other studies that have
involved North Africa samples, and found that European,
North Africa and Asia Minor samples are in one clade as Acer
core clade with BS 75% and PP 0.96 (Grimm et al. 2007).
Our maximum parsimony results (Figure 1) indicate
that Iranian A. monspessulanum subspecies (13 populations)
in clade N are closely related to eight A. ibericum samples
from Georgia, one sample of A. hyrcanum from Iran and
two samples of A. sempervirens from Greece (pp= 1, BS=
79). This agrees with results reported by Grimm et al.
(2007) where different taxa of one of one group fall into
three lineages. In their results, Acer monspessulanum and
A.ibericum +A. hyrcanum group together in the graph and
long proximal edges indicate that they are most closely
related. Clones of A. monspessulanum are distinct and
placed near the center of the graph (Grimm et al. 2007).
Clade O that includes six specimens of A. ibericum
from Georgia with one specimen of A. monspessulanum
subsp. turcomanicum from Iran and one specimen of A.
sempervirens from Greece (PP= 0.79; BS= 62%) has
proved Rechinger results about Iranian Acer origin. Clade
M comprises 4 species of A. monspessulanum from Bulgaria
with PP= 0.8 and BS= 77%. Clades M and N together are
in clade C (PP= 0.95). This close relation between Iranian
and Bulgarian Acer monspessulanum species samples support
this notion that they have an origin in Mediterranean region
(Rechinger 1969). Clade L has two species from France,
one A. monspessulanum and A. monspessulanum ssp.
monspessulanum. Clade K include clade L with another A.
monspessulanum ssp. monspessulanum from France (PP=
0.84). Clade J includes clades K and L with four taxa of A.
monspessulanum ssp. monspessulanum from France and
Spain. Clade I have three A. monspessulanum ssp.
monspessulanum from France with PP= 0.91. Clade H
include 17 A. monspessulanum ssp. monspessulanum with
A. opalus, all from France (PP= 0.97). Each clades of G
and F has two A. monspessulanum ssp. monspessulanum
from France with PP= 1 and BS= 80%. Clade E comprise
clades F, G, H, I, J, K and L from France and Spain. Clade
D has only one species A. monspessulanum from France
that with clade E are in clade B with BS= 50%. Clade P
that has one A. velutinum from Iran, consider as out-group
in our analysis. Data analysis indicates that the
classification of species according presence or absence of
hairs in inner or outer surface of loculus is a true
morphological characteristic for delimitation of subspecies
in Acer monspessulanum.
The observed polytomies in clades E and N indicated
that these taxa are taxonomically closely related and there
were not enough time passed since divergence from their
ancestral taxa (Zarrei et al. 2009). More divergent markers,
i.e. low-copy nuclear genes, could potentially resolve these
branches.
Based on our results using Bayesian analysis, some well
resolved clades were present (Figure 2). Clade A comprise
two subclades, E and F. Clade E includes four A.
monspessulanum specimens from Bulgaria and clade F
includes 27 taxa (PP= 1), that 13 of them are A.
monspessulanum subspecies from Iran and others are A.
ibericum (from Georgia), A. sempervirens (from Greece)
and A. hyrcanum subsp. hyrcanum (from Iran), this clade
proved Mediterranean origin of Iranian Acer (clade F;
Figure 2). One of our taxa (A. monspessulanum subsp.
ibericum) placed in clade L with five A. ibericum from
Georgia and an Acer sempervirens from Greece with high
support (PP= 0.79). This placement indicates that A.
monspessulanum and A. sempervirens are closely related
together (Grimm et al. 2007).
B I O D I V E R S IT A S
17 (1): 16-23, April 2016
20
Figure 1.Phylogenetic relationships of 13 samples of Acer from Iran, on the basis of the analysis of internal transcribed spacer (ITS)
sequences. Numbers above branches are Bayesian posterior probabilities. Numbers below branches are bootstrap values
KHADEMI et al. –Molecular phylogeny of Acer monspessulanum
21
Figure 2. Bayesian tree. Numbers on nodes represent posterior probability values
B I O D I V E R S IT A S
17 (1): 16-23, April 2016
22
Clade B (Figure 2) have two subclades M and N (PP=
0.99) that include European Acer monspessulanum (from
France, Spain and Germany). Clades G and N show
polytomies since comprise species their morphology are
closely related together.
Clade C (Figure 2) have four taxa, three of them are A.
sempervirens from Greece and one A. obtusifolium from
Cyprus (PP= 1) that in Grimm et al. (2007) study In
Europe, their distribution ranges from northern, cool-
temperate latitudes (southern Sweden, A. pseudoplatanus
L., naturalized) throughout central, western, and south-
eastern Europe (A. hyrcanum Fischer & Meyer, A. opalus
P. Miller, A. pseudoplatanus) to the subtropical
Mediterranean (A. monspessulanum L., A. opalus,A.
heldreichii Orphanides ex Boissier, A. sempervirens L., and
A. obtusifolium Sibthorp & Smith), and with an eastward
expansion to Asia Minor, the Caucasus, and Iran (A.
trautvetteri Medvedev, A. hyrcanum,A.ibericum
Bieberstein ex Willdenow, A. monspessulanum,A.
pseudoplatanus, and A. velutinum Boissier) (Grimm et al.
2007).
Subspecies geographical distribution show that
assyriacum subspecies from Kordestan and Kermanshah
provinces are near together and support with Bayesian
analysis (PP= 0.82). A. monspessulanum subsp. ibericum
collected from Arasbaran forest (Azarbayejansharghi
province) placed in clade L (Figure 2) with A. ibericum
from Georgia (PP = 0.79) that from geographical
distribution approach is justifiable. All Iranian Acer
monspessulanum in the present survey made a clade with
two A. sempervirens from Greece and eleven A. ibericum
from Georgia and one A. hyrcanum subsp. hyrcanum from
Iran with high Bayesian support (PP= 1).
Because ITS results couldn’t delimitate on subspecies
level we used from morphological traits. Some
morphological features are important for identification of
subspecies A. monspessulanum, such as size of the leaves,
lower surface midrib hair and loculus inside and outside
base on presence or absence of hair. One of the most
important characteristic traits for distinguish between some
subspecies of A. monspessulanum, is presence or absence
of hair inside and outside their loculus. Based on the size of
the leaves we have two groups, (i) small (up to 2×2 cm)
that has two subspecies (ssp. persicum and ssp.
cinarescens) and (ii) large (2-3×3.5-4 cm) with three
subspecies (ssp. turcomanicum, ssp. assyriacum and ssp.
ibericum). Lower surface midrib hair separate only two
subspecies of A. monspessulanum, one of them is glabrous
(ssp. ibericum) and the other is sparsely hairy (ssp.
assyriacum), so this trait is not a proper discriminative
factor for other three subspecies. Loculus outside hair has
two state, (i) glabrous (ssp. turcomanicum, ssp. persicum)
and (ii) sparsely hairy (ssp. ibericum, ssp. assyriacum, ssp.
cinarescens). We have three group base on loculus inside
hair, (i) hairy (ssp. turcomanicum, ssp. ibericum, ssp.
assyriacum), (ii) densely hairy (ssp. cinarescens) and (iii)
glabrous (ssp. persicum) (Wolfe and Yanai 1987), so the
most important and discriminative character to detect
subsp. persicum from subsp. cinarescens is loculus inside
hair.
The internal transcribed spacer of the nuclear region
(ITS) is a widely used molecular marker for reconstruction
of evolutionary patterns in plant kingdom. It has been used
both in the higher taxonomic level (i.e. family) as well as
lower even below the species rank (Zarrei et al. 2014). Our
results indicate that this marker could be potentially
valuable in delineating subspecies boundaries in maple
species. The limiting factor is that this marker is not well
diverged in some groups. A more divergent molecular
marker such as low copy nuclear genes and intergenic
nuclear spacers could potentially be helpful. Based on our
experiences working on Acer and results of others studies
on tree taxa (Zarrei et al. 2014); we suggest combining our
ITS DNA sequences with additionally markers to increase
the power of our phylogenetic analysis and improve
resolution of unresolved clades. Such strategies have been
applied before (see Zarrei et al. 2015). The implication of
next generation sequencing data has been already proven in
revolving systematic problem with closely related species
(Liston et al. 2015).
ACKNOWLEDGEMENTS
This article is extracted from first author’s Ph.D. thesis.
We would like to thank from Islamic Azad University,
Science and Research Branch, Tehran, Iran for providing
the facilities necessary to carry out the work and Masoud,
Basiri and Meyjani for their help in plant material
collection. We thank the invaluable suggestions of an
anonymous reviewer on early draft of this manuscript.
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One table includes a statistical summary of flowering-plant taxa: c235 000 species of 12 615 genera, 440 families, and 711 subfamilies and undivided families in 28 superorders, 70 orders, and 75 suborders of Angiospermae. Three other tables summarize the indigenous distribution of the families and subfamilies of Angiospermae about the world. -from Author
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