ACTA BOT. CROAT. 77 (2), 2018 119

Acta Bot. Croat. 77 (2), 119–125, 2018  CODEN: ABCRA 25
DOI: 10.2478/botcro-2018-0006 ISSN 0365-0588
 eISSN 1847-8476
 

Application and limitation of molecular data and 
essential oil content in identification of Leutea 
elbursensis Mozaff in northern Iran
Samane-Sadat Emami-Tabatabaei1, Kambiz Larijani2, Iraj Mehregan1*

1 Department of Biology, Science and Research branch, Islamic Azad University, Tehran, Iran
2 Department of Chemistry, Science and Research branch, Islamic Azad University, Tehran, Iran

Abstract – In this paper, the internal transcribed spacer (ITS) sequences, genetic structure and the chemical composi-
tion of essential oils of four populations belonging to Leutea elbursensis and Leutea petiolaris, two species endemic to 
northern Iran, are analyzed. Phylogenetic analysis based on the ITS data showed that all accessions of L. elbursensis 
formed a monophyletic clade, and L. elbursensis was a sister to the rest of Leutea species. Results of amplified fragment 
length polymorphism (AFLP) analysis performed on the total genome showed that all individuals presented in the 
study belonged to two different genetic clusters. The individuals belong to L. petiolaris had a different genetic structure 
and yielded no traceable amount of essential oils. The essential oil obtained from the ripe fruits of L. elbursensis yielded 
0.5–0.6% of volatile essential oils. In total, 15–29 volatile natural components were identified on the basis of their mass 
spectra characteristics and retention indices, in which α-pinene (33.18–43.22%), β-pinene (32.4–40.9%) were the ma-
jor constituents. Our results indicate that L. elbursensis is a distinct species, segregated from the other species based on 
morphology, ITS data and AFLP profile. In addition, despite the relatively uniform genetic structure of L. elbursensis, 
the chemical composition of essential oil could be highly affected by different factors.

Keywords: chemotaxonomy, Iran, phylogeny, α-pinene, β-pinene

* Corresponding author, e-mail: iraj@daad-alumni.de; imehregan@srbiau.ac.ir

Introduction
Plants are an important component of traditional food, 

but are also central to healthy diets of the modern urban 
population (Benjak et al. 2005, Ercisli 2009, Ercisli et al. 2010, 
Rop et al. 2014, Canan et al. 2016, Zorenc et al. 2016). Es-
sential oils obtained from plants have applications in food, 
chemistry, pharmacy, medicine and perfumery (Hay and 
Waterman 1993, Mehregan and Ghannadi 2013). Members 
of the family Apiaceae (Umbelliferae) are well known for 
the production of essential oils with high pharmaceutical 
and economic value (Olivier and van Wyk 2013). Based on 
the most recent researches, the plant family Apiaceae with 
about 430 genera and 3800 species has a worldwide distribu-
tion, especially in the northern hemisphere (Stevens 2012). 
Apiaceae has about 110 genera and 400 species distributed 
in Iran (Mozaffarian 2007). The genus Leutea Pimenov be-
longs to the “Ferula group” including Ferula L., Dorema L. 
and Leutea (Kurzyna-Młynik et al. 2008). The genus Ferula 
includes more than 170 species distributed in central and 
southwestern Asia, and the Mediterranean region includ-
ing northern Africa (Pimenov and Leonov 1993, Kurzyna-

Młynik et al. 2008). It has about 30 aromatic species in Iran 
(Mozaffarian 2007). The genus Dorema has seven species 
in Iran (Mozaffarian 2007). The taxonomic status of Leu-
tea is more complex. The genus Leutea with its few species 
is limited to SW Asia (Pimenov 1987). The taxonomy of the 
group has changed considerably, especially since publication 
of Flora Iranica, volume 162 (Pimenov 1987). The type of 
the genus, i.e. Leutea petiolaris was first effectively published 
as part of the genus Ferula as F. petiolaris DC. (de Candolle 
1830). Boissier (1872) integrated L. petiolaris with other spe-
cies of the genus Peucedanum L. sect. Juncea. He described 
two other new species, which are today known to be part of 
Leutea. Pimenov (1987) transferred all six known species of 
the group to the new genus named Leutea Pimenov. Spalik 
and Downie integrated the whole Leutea species into the ge-
nus Ferula (see Kurzyana-Mlynik et al. 2008). Recent works 
of Panahi et al. (2015) suggested that species of Leutea should 
be re-established from Ferula again. 

Leutea elbursensis Mozaff. (syn.: Ferula elbursensis (Mo-
zaff.) Spalik et. S. R. Downie), an endemic to northern Iran, 



EMAMI-TABATABAEI S.-S., LARIJANI K., MEHREGAN I.

120 ACTA BOT. CROAT. 77 (2), 2018

is a glabrous perennial tough herb, 1.5 to 3 m high, with 
compound pinnate leaves with tubular lobes. It has com-
pound umbels consisting of yellow 5-merous flowers. The 
fruits are schizocarp, compressed, elliptic and up to 10 × 4 mm 
in size (Mozaffarian 2007, Kanani et al. 2013). L. elbursen-
sis can be distinguished from the other species by its nar-
row pedicels, ovate-elliptic fruits, greenish-yellow petals, and 
large and strong habits (Mozaffarian 2007). Despite the re-
cent publication date of its name, many populations of L. el-
bursensis have been known for a long time. Material of L. el-
bursensis was first considered part of L. cupularis (Pimenov 
1987), and was later segregated from it by Mozaffarian in 
2003 (Kurzyna-Młynik et al. 2008). Mozafarian (2007) iden-
tifield two different species in the north of Iran (Tehran prov-
ince) i.e. L. petiolaris and L. elbursensis. These taxonomic 
complexities have resulted in ambiguity attaching to those 
works already performed on three species L. elbursensis, L. 
cupularis and L. petiolaris (Masoudi et al. 2004, Kurzyna-
Młynik et al. 2008, Alipour et al. 2015).

Members of the family Apiaceae in Iran are widely stud-
ied for their essential oil contents (Olivier et al. 2013, Sa-
faeian et al. 2015). Most of those studies are single records 
based on the material collected from a single locality. Chemi-
cal composition of the essential oils depends on many fac-
tors such as collection time, geographical locality and ge-
netic structure (Munoz-Bertomeu et al. 2007). Therefore, a 
single report of essential oil composition for a given species 
could not be generalized for all of its populations. Looking 
for their chemotaxonomic value, Kanani et al. (2011) stud-
ied chemical composition of the essential oils for 18 Ferula 
species from Iran. 

Since its introduction in 1995, amplified fragment length 
polymorphism (AFLP) has become a popular research tool 
for detecting genetic structure of populations as well as dif-
ferences at intra-species level (Vos et al. 1995). 

In this paper we try to find out if different populations 
growing wild in the north of Iran (Tehran province) belong 
to the same species. Material from the western part of the 
province was previously identified as L. elbursensis by Mozaf-
farian (2007). We used analysis of ITS (internal transcribed 
spacer) of the nuclear genome to find the phylogenetic place-
ment of those plants. In addition, AFLP fingerprinting tech-
niques are used for distinguishing the genetic structures of 
different populations. We also aim to study the possible cor-
relation between the genetic structure and the essential oil 
profile of L. elbursensis populations collected from different 
localities in northern Iran (Tehran province), using GC/MS 
and AFLP fingerprinting techniques. Finally, we will try to 
clarify the taxonomic position of L. elbursensis.

Materials and methods
Between August and September 2013, plants from three 

populations of L. elbursensis and one population of L. pet-
iolaris were collected from northern Iran, the province of 
Tehran (Tab. 1). Plant materials were identified by authors 
after Mozaffarian (2007) and vouchers were deposited in the 

herbarium of Islamic Azad University, Science and Research 
branch, Tehran, Iran (IAUH).

Fresh leaf fragments from at least six individuals from 
each population (24 individuals in total) were taken and 
gradually dried in silica-gel pearls. Total DNA was extract-
ed for each individual sampled using NucleoSpin® Plant II 
kit after the manufacturer’s manual (Machery-Nagel, Du-
eren, Germany). The complete internal transcribed spacer 
(ITS) region of the DNA was amplified using the primer 
pair AB101 (5ʹ- ACG AAT TCA TGG TCC GGT GAA GTG 
TTC G – 3ʹ) and AB102 (5ʹ-TAG AAT TCC CCG GTT CGC 
TCG CCG TTA C – 3ʹ) (Douzery et al. 1999), in a PCR re-
action under the following conditions: a pretreatment of 5 
minutes at 95 °C, 35 cycles of 30 seconds at 95 °C, 30 sec-
onds at 50 °C, and 1 minute 30 seconds at 72 °C, and a final 
extension of 7 minutes at 72 °C. The complete ITS region 
was sequenced on an ABI 3730 sequencer machine (Applied 
Biosystems, Waltham, Massachusetts, USA). Sequences were 
visually checked and edited with Sequencher 4 (Gene Codes 
Corporation, Ann Arbor, MI USA), and then aligned using 
MacClade 4.08 (Maddison and Maddison 2000), alongside 
additional sequences taken from the GeneBank. Ferula vio-
lacea Korovin and F. olivacea (Diels) H. Wolff. were chosen 
as outgroup taxa after Panahi et al. (2015). Maximum parsi-
mony (MP) analysis of the ITS dataset was performed with 
PAUP* (Swofford 2002). Bayesian analysis (BA) of the ITS 
dataset was performed using MrBayes v3.1.2 (Huelsenbeck 
and Ronquist 2001).

The AFLP™ procedure for this work followed Vos et al. 
(1995), and Scalone et Albach (2012) with the following 
modifications: three primer combinations used for selective 
PCR were E38-HEX labelled (5ʹ – GAC TGC GTA CCA ATT 
CAC T – 3ʹ) combined with M57 (5ʹ- GAT GAG TCC TGA 
GTA ACG G – 3ʹ), E45-FAM labelled (5ʹ-GAC TGC GTA CCA 
ATT CAT G – 3ʹ) combined with M54 (5ʹ – GAT GAG TCC 
TGA GTA ACC T – 3ʹ), and E40-NED labelled (5ʹ- GAC TGC 
GTA CCA ATT CAG C – 3ʹ) with M55 (5ʹ-GAT GAG TCC 
TGA GTA ACG A – 3ʹ). PCR products of each sample were 
combined equally, and 2 μl of this multiplex product was run 
with 7.75 μl HiDi formamide (Applied Biosystems) and 0.25 
μl internal size standard GeneScan ROX (Applied Biosys-
tems) on an ABI 3730 automated capillary sequencer. Frag-
ments were analyzed and scored using GeneMarker 2.4.1 
(SoftGenetics). Structure 2.3.4 (Pritchard et al. 2000) was 
used for analyzing the genetic structure of populations. An 
analysis of molecular variance (AMOVA) test was performed 
using GenAlEx 6.5 (Peakall and Smouse 2012).

100 g of the ripened and dried fruits of the plant for each 
population were chopped in distilled water and a hydro-dis-
tilled fraction of it was isolated by hydrodistillation for 3 
hours. We used a Hewlett Packard 5972A mass selective de-
tector coupled with a Hewlett Packard 6890 gas chromato-
graph, equipped with a cross-linked 5% PH ME siloxane HP-
5MS capillary column (50 m × 0.25 mm, film thickness 0.32 
μm) for gas chromatography-mass spectrometry (GC-MS) 
analysis. The following GC operating conditions were used: 
carrier gas, helium with a flow rate of 1 mL min–1; column 



MOLECULAR IDENTIFICATION OF LEUTEA ELBURSENSIS

ACTA BOT. CROAT. 77 (2), 2018 121

temperature, 60 °C with 7 °C temperature increase per min-
ute, up to 230 °C; injector and detector temperatures, 280 °C; 
volume injected, 0.1 μL of the oil; split ratio, 1:25. In addition, 
the following MS operating parameters were used: ionization 
potential, 70 ev; resolution, 1000; ion source temperature, 
200 °C. Components in the oil were identified based on GC 
retention indices relative to n-alkanes and computer match-
ing with the Wiley 275.L library, as well as by comparison of 
the fragmentation patterns of the mass spectra with those 
reported in the literature (Adams 1995, Masoudi et al. 2004, 
Alipour et al. 2015).

SPSS v. 20 (IBM corporation) was used to perform clus-
ter analyses based on the chemical components of essential 
oils with Ward’s method (Ward 1963).

Results 
Figure 1 shows the phylogeny of different species of Leu-

tea based on the Bayesian analysis (BA) of the ITS region. 
Two species of Ferula i.e. F. violacea and F. olivacea, were in-
cluded in the tree as outgroups. Posterior probabilities (PP) 
are indicated by numbers above each clade. Bootstrap sup-
ports (BS) for those clades also retrieved in the maximum 
parsimony (MP) analysis are indicated by the numbers be-
low each clade (Fig. 1). As seen on the phylogenetic tree, spe-
cies of the genus Leutea are mainly grouped into two clades: 

clade A) a firmly supported clade including all samples of L. 
elbursensis with posterior probability (PP) = 1.00 and boot-
strap support (BS) = 100%, and clade B) another well sup-
ported clade including a polythomy of eight species with PP 
= 1.00 and BS = 85%. Leutea elbursensis is sister to rest of spe-
cies (Fig. 1). All our three accessions of L. elbursensis togeth-
er with another two samples taken from the Genbank (both 
from NW of Tehran) formed a monophyletic clade. All four 
samples are taken from a relatively small locality in N Iran 
(central Elburz) with distances of less than 100 km among 
them. The monophyletic clade B presents a polythomy of dif-
ferent species. These include three samples of L. petiolaris, 
two samples of L. cupularis, and one sample of each species 
L. galucopruinosa, L. rechingeri, L. polyscias, L. avicennae, L. 
gracillima and L. nematoloba (Fig. 1). Two samples of L. cu-
pularis collected from the Dena mountain ranges (SW Iran) 
formed a monophyletic clade with strong support (PP = 1, 
BS = 85%). Different samples of L. petiolaris did not form 
a monophyletic clade. The results showed that our samples 
from the area clearly belong to two different species. Phylo-
genetic relationships within this clade are not essentially re-
solved. This clade consists of samples collected from a larger 
region involving distances of hundreds of kilometres.  

In the selective PCR of AFLP analysis, the E38-M57 
primer combination yielded 62 bands, the E45-M54 primer 
combination 36 bands, and the E40-M55 primer combina-

Tab. 1. Populations of Leutea elbursensis from northern Iran included in the molecular and phytochemical analysis with voucher informa-
tion and GenBank accession numbers.

No. Species Locality Herbarium number
Genbank
number

1 Leutea elbursensis Mozaff. Iran: Tehran, Emamzadeh-Davoud,35.898, 51.288, 2400 m Emami 14391 (IAUH) KP793684

2 Leutea elbursensis Mozaff. Iran: Tehran, Morad-Abad, 35.796, 51.322, 1800 m
Emami 14393 
(IAUH)

KP793683

3 Leutea elbursensis Mozaff. Iran: Tehran, Hesarak35.796, 51.305, 1700 m Emami 14394 (IAUH) KP793686

4 Leutea petiolaris (DC.) Pimenov Iran: Tehran, Dizin,36.043, 51.441, 3400 m
Emami 14392
(IAUH)

KP793685

5 Leutea elbursensis Mozaff. Iran: Tehran, Karaj, 35.933, 51.067, 1600 m
Valiejo-Roman et al.,
526 (MW)

AY941276
AY941304

6 Leutea elbursensis Mozaff. Iran: NW of Tehran, Souleghan Mozaffarian & Jamzad 33570 (TARI) KJ660832

7 Leutea glaucopruinosa (Rech. fil.) Akhani & Salimian Iran: Mazandaran
Termeh & Zargari 040483-E (W 
0023608)

KJ660833

8 Leutea petiolaris (DC.) Pimenov Iran: Azarbayejan Mozaffarian & Massoumi 78152 (TARI) KJ660836

9 Leutea petiolaris (DC.) Pimenov Iran: Tehran,35.767, 51.950, 2400 m Valiejo-Roman et al. 63 (MW)
AY941278
AY941306

10 Leutea rechingeri (Leute) Pimenov Iraq: Suleymanieh, Mt. Algurd Rechinger 11416 (W 05825) KJ660838

11 Leutea cupularis (Boiss.) Pimenov Iran: SW, Dena Mts., 2900-3100 m Valiejo-Roman et al.235 (MW)
AY941277
AY941305

12 Leutea cupularis (Boiss.) Pimenov Iran: SW, Dena Mts., 3500-3900 m Assadi & Mozaffarian. 31236 (TARI) KJ660831
13 Leutea polyscias Pimenov Iran: Manjil, 1650 m Mozaffarian 64227 (TARI) KJ660837
14 Leutea gracillima Pimenov Iran: NE, Golestan Park. Akhani 12060 (W 1999-03655) KJ660834
15 Leutea nematoloba (Rech.f.) Pimenov Iran: N, Chalus Valley. Rechinger & Rechinger 6668 (W 02835) KJ660835
16 Ferula violaceae Korovin SW Asia Valiejo-Roman et al., 119899 (MW) AF077891
17 Ferula olivacea (Diels) H. Wolff. China Chamberlain, Ming, Yuan & al. 229 (E) EF560691



EMAMI-TABATABAEI S.-S., LARIJANI K., MEHREGAN I.

122 ACTA BOT. CROAT. 77 (2), 2018

tion 36 bands. The AMOVA test results showed that of 100% 
total variation, 34% was among regions, 6% was among pop-
ulations and 60% was within populations (Tab. 2). Analysis 
of the populations using Structure 2.3.4 software showed that 
all individuals belonged to two different clusters showed here 
with different colours (Fig. 2). As seen in Figure 2, individu-
als of L. petiolaris from “Dizin” population are homogenous 
in their genetic structure (cluster “a”). All six individuals 
collected from “Dizin” (L. petiolaris) had genetic structures 
90–100 % consisting of cluster “a” alleles. Genetic profiles of 
three populations of L. elbursensis mainly consist of a differ-
ent cluster (cluster b). Genetic structures of all six individuals 
from the “Emamzadeh-Davoud” population consisted 100 % 
of alleles from cluster b. Except for one individual from the 
“Hessarak” population and another one individual from the 
“Morad-Abad” population, all other individuals from those 
two populations were 97-100% made by alleles from cluster 
“b”. AFLP profiles of two species are clearly different. This 
clearly shows that two different species are distributed in the 
area N and NW of Tehran. 

This is the first report of chemical composition of essen-
tial oils extracted from the ripened fruits of L. elbursensis. 
The essential oils obtained from dried fruits were clear, pale 
yellow liquids. The essential oil content of L. elbursensis was 
between 0.5% (w/w; in the Emamzadeh-Davoud population) 
– 0.6% (w/w; in Hessarak and Moradabad populations). Be-
tween 15 to 29 natural compounds were identified, account-
ing for 98.7 – 100% of the oils (Tab. 3). Ripened fruits of 
L. petiolaris collected for two successive years did not yield 

Fig. 1. Phylogenetic tree obtained from the Bayesian analysis of 
the internal transcribed spacer region of different species of Leutea 
alongside two Ferula species as outgroup (total characters = 442; 
constants = 299; parsimony informatives = 60. Model of evolution 
= SYM +G; A-C constitution rate = 1.7930; A-G constitution rate 
= 2.2561; A-T constitution rate = 2.0384; C-G constitution rate = 
0.4189; C-T constitution rate = 5.9481; G-T constitution rate = 
1.0000. Gamma distribution rate = 0.6465). Numbers above clades 
are posterior probabilities. Numbers below clades are the bootstrap 
supports (100 replicates) for those clades retrieved in the maxi-
mum parsimony analyses (bootstrap supports less than 50% are 
not shown).

Tab. 2. AMOVA test results showing the variations of Leutea elbur-
sensis from northern Iran: within populations, among populations 
(pops) and among regions. Df – degrees of freedom; SS- sum of 
the squares; MS – mean squares; Est. Var. – estimation of variance.

Source Df SS MS Est. Var. %
Among regions 1 124.972 124.972 10.602 34
Among pops 2 59.111 29.556 1.798 6
Within pops 20 375.333 18.767 18.767 60
Total 23 559.417 173.295 31.167 100

Fig. 2. Amplified fragment length polymorphism finger printing results showing the genetic profile of 24 individuals collected from 
northern Iran (Leutea elbursensis from Hessarak, Morad-Abad and Emamzadeh (E-) Davoud; L. petiolaris from Dizin). 



MOLECULAR IDENTIFICATION OF LEUTEA ELBURSENSIS

ACTA BOT. CROAT. 77 (2), 2018 123

any traceable essential oil. The highest number of compo-
nents was identified in the Moradabad population (29 com-
ponents). We also identified 20 components for the “Hes-
sarak” population. The smallest number of components was 
15 and was identified in the “Emamzade-Davoud” popula-
tion. The main components identified in our three popula-
tions were α –pinene (33.18 – 43.22%) and β-pinene (32.4 
– 40.9%). There were few other important but less abundant 
components (ca. 5% or more) i.e. myrtenol (5.6%), trans-
verbenol (5.6%) and isobornyl acetat (4.95%) which were 
identified only in the Hesarak population. 

Discussion
Different populations of L. elbursensis had identical ITS 

sequences and relatively similar genetic structures. Its seg-
regation from L. petiolaris was supported by ITS and AFLP 
data. This segregation is also supported by morphological 
and geographical evidence. Leutea elbursensis has unique tall 

Tab. 3. Composition of the essential oils of different populations of Leutea elbursensis from northern Iran. KI – Kovats index.

Number Component name KI
Percentage of components in each population 

Morad-Abad Hessarak Emamzadeh-Davoud
1 α-Thujene 930 1.38 0.54 1.2
2 α-Pinene 939 39.77 33.18 43.22
3 Camphene 954 1.32 0.57 1.24
4 Thuja-2,4 (10)-diene 960 0.14 0 0
5 Sabinene 975 2.05 0.71 1.9
6 β-Pinene 979 36.1 32.4 40.9
7 Myrcene 990 0 1 0
8 Р-Cymene 1024 0.35 0 0
9 Limonene 1029 3.14 1.9 3.5

10 β-Phelandrene 1029 0.91 0.64 0
11 Cineol 1031 0.78 0 0.8
12 z-β-Ocimene 1037 0.18 0 0
13 γ-Terpinene 1059 0.19 0 0
14 α-Campholenal 1126 0.41 0.53 0
15 Trans-Pinocarveol 1139 3.5 0 1.3
16 Trans- Verbenol 1144 0.16 5.62 0
17 Pinocarvone 1164 0.9 1.52 0.8
18 α-Terpineol 1188 0.33 0.78 0.46
19 Myrtenol 1195 1.34 5.6 1.2
20 Verbenone 1205 0.19 0 0
21 endo-Fenchyl acetate 1220 1.32 3.02 0.8
22 Isobornyl acetate 1285 1.36 4.95 1.73
23 cis-Pinocarvyl acetate 1312 0.25 0.75 0
24 Myrtenyl acetate 1326 0.57 2.12 0.45
25 Neryl acetate 1361 0.26 2.01 0.5
26 Daucene 1381 0.52 0.81 0
27 α-Trans Bergamotene 1434 0.1 0 0
28 z-β-Farnesene 1442 0.15 0 0
29 Pentadecane 1500 0.2 0 0
30 Trans-β-Guaiene 1502 0.2 0 0
31 α-Copaen-11-01 1539 0 0.8 0

Total 98.07 99.45 100

flowering stems 1.5–3 m high, while other species are clearly 
shorter (up to 1.5 m high). Leutea species have an allopat-
ric distribution. Leutea elbursensis is endemic to N and NW 
Tehran, especially the rocky slopes of Souleghan valley (Mo-
zaffarian 2007). Despite the similarities in morphology, ITS 
sequence and genetic structure, the essential oil contents of 
our three populations were highly variable. 

The results showed that Leutea plants distributed in the 
region N and NW of Tehran do not belong to a single species. 
Pimenov (1987) mistakenly identified all herbarium material 
collected from the area under L. cupularis. We showed here 
that they belong to two different species. Our results are in 
agreement with Mozaffarian (2007), who identified the ma-
terial from “central Elburz” under two different species i.e. 
L. elbursensis and L. petiolaris. These results indicate that F. 
petiolaris and F. elbursensis are two different entities. Regard-
less of the disputable taxonomic position of the genus Leu-
tea, our results clearly showed that L. elbursensis is a distinct 
species sister to the rest of Leutea species (Fig. 1). 



EMAMI-TABATABAEI S.-S., LARIJANI K., MEHREGAN I.

124 ACTA BOT. CROAT. 77 (2), 2018

The genus Ferula s. l. (including Leutea) in Iran is widely 
studied chemically. We compared our results to one of the 
most comprehensive studies, that performed by Kanani et 
al. (2011) on 18 different populations belonging to different 
species of Ferula (Fig. 3). Our analysis showed that the essen-
tial oil profile of our three samples was most similar to that of 
a sample from F. stenocarpa Boiss. & Hausskn. All these four 
samples formed a distinct group “rich” and “balanced” of α –
Pinene (33.18 – 48.8%) and β-Pinene (30.1 – 40.9%), and this 
group is also related to another group consisted of F. gum-
mosa Boiss. and F. galbaniflua Boiss. & Buhse with a higher 
amount of β-Pinene (26.8 – 69.2%) and a lower amount of α–
Pinene (1.4–33.9%) (Ghannadi and Amree 2002, Ghasemi et 
al. 2005, Jahansouz et al. 2008, Talebi Kouyakhi et al. 2008, 
Kanani et al. 2011). The chemical composition of essen-
tial oils of Leutea glaucopruinosa Rech.f. Yassa et al. (2003) 
showed no close similarity to our samples (“LG” in Figure 
3). The chemical composition of Dorema glabrum Fisch. & 
C. A. Mey. Delnavazi et al. (2015) is similar to that of some 
Ferula species. Comparing the dendrogram shown in Fig-
ure 3 with the Leutea phylogeny (Fig. 1) and the phylogeny 
obtained by Kurzyna-Mlynik et al. (2008), it becomes clear 
that similarities in chemical composition of the essential oils 
of the “Ferula group” does not reflect the phylogenies based 
on the molecular data.

Regarding their uniform ITS sequences and similar AFLP 
profiles, the essential oil profiles of different populations of 
L. elbursensis were very variable, most probably as the result 
of different ecological conditions. The chemical composition 
of essential oils in the family Apiaceae can be very variable, 
especially according to the plant parts the oils are extracted 
from (Kanani et al. 2011). Alipour et al. (2015) reported dif-
ferent chemical compositions for the essential oils extract-
ed from different parts of L. cupularis. They found following 
major components in different parts of the plant: ð-2-Carene 
(15.81%) and DL-Limonene (25.04%) in flowers; β-Pinene 
(13.87%), β-Ocimene (9.05%), Bornyl angelate (6.55%) and 
allo-Ocimene (6.08%) in leaves; and ð-3-Carene (8.38%), 
α-Terpinyl isobutyrate (8.69%) and Bornyl angelate (7.45%) 
in stems. In addition, the chemical composition of essential 
oil can be highly affected by ecological conditions and ge-
netic structure (Munoz-Bertomeu et al. 2007). Therefore, 
we here suggest that a standard method should be used for 
analyzing and interpreting the data obtained from essential 
oils. As mentioned above, different parts of the plants might 
yield essential oils with different contents. Ripened fruits of 
the members of the family Apiaceae could be an appropriate 

References

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Fig. 3. Dendrogram showing classification of different populations 
of Ferula spp. based on the similarities in essential oil composi-
tion. Amount of components for Leutea populations are shown 
in Table 3 and for other populations are presented in Kanani et al. 
(2011). Numbers on the x-axis represent values of dissimilarity. LE1 
– Leutea elbursensis, Moradabad; LE2 – L. elbursensis, Hessarak; 
LE3 – L. elbursensis, Emamzadeh-Davoud); LE4 – L. elbursensis, 
Karaj; OV1-4 – Ferula ovina; SZ – F. szowitsiana; MI – F. micro-
colea; PP – F. persica var. persica; PL – F. persica var. latisecta; ST 
– F. stenocarpa; GA1-2 – F. galbaniflua; GU – F. gummosa; BE – F. 
behboudiana; OO – F. oopoda; HI – F. hirtella; MA – F. macrocolea; 
FL – F. flabelliloba; AS – F. assa-foetida; OR – F. orientalis; HE – F. 
hezarlalehzarica; FO – F. foetida; DI – F. diversivittata; LA – F. lati-
secta); LG – L. glaucopruinosa; DG – Dorema glabrum.

source for obtaining essential oils. Regarding those issues, we 
here suggest that the chemical composition of essential oils 
cannot be used as a trusted taxonomic tool, at least in case of 
the “Ferula-group”. 

Acknowledgments
The authors would like to thank Mrs R. Amini for her 

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