NUTLET AND FLOWER MORPHOLOGICAL AND MICROMORPHOLOGICAL STUDIES ON ONOSMA L. (BORAGINACEAE) IN IRAN ‫ﺷﻨﺎﺳﻲ‬ ‫رﻳﺨﺖ‬ ‫ﻣﻄﺎﻟﻌﺎت‬ ‫ﺷﻨﺎﺳﻲ‬ ‫رﻳﺨﺖ‬ ‫رﻳﺰ‬ ‫و‬ ‫ﺑﺮﺧﻲ‬ ‫ﻓﻨﺪﻗﻪ‬ ‫و‬ ‫ﮔﻞ‬ ‫ﻫﺎي‬ ‫ﮔﻮﻧﻪ‬ ‫از‬

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NUTLET AND FLOWER MORPHOLOGICAL AND MICROMORPHOLOGICAL
STUDIES ON ONOSMA L. (BORAGINACEAE) IN IRAN
M. Arabameri, A.R. Mehrabian & M. Sheidai
Received 2014. 03. 04; accepted for publication 2014. 09. 03
Arabameri, M., Mehrabian, A.R. & Sheidai, M. 2014. 12. 31: Nutlet and flower morphological and
micromorpholoical studies on Onosma L. (Boraginaceae) in Iran.- Iran. J. Bot.20 (2): 211-227.Tehran.
The morphology and micromorphology of flower and nutlet in 29 Iranian Onosma species were examinated
comparatively using SEM method in order to evaluate and clarify their taxonomic values and specific emphasis was
on their role in classification. Our study highlights the importance of the floral and nutlet characters in relation to
identification of most of the studied species based on the PCoA and cluster analysis. Besides, Palynological,
morphological and molecular studies confirmed our results. Although there are some taxa which cannot be
determined solely using the mentioned characters, but these characters accompanied with other important characters
are useful tools in distinguishing many taxa and solving some complexities in Onosma. Moreover if molecular
evidence (chloroplast and nuclear sequences) confirm our results, the divergence of Onosma orientalis and Onosma
rostellata from the genus Onosma is expexcted, then these taxa may be transferred to different genera. On the other
hand, combinations of these characters along with morphological and molecular evidence provide a perfect view for
taxonomy and comphrensive taxonomical revision of Onosma.
Mahnaz Arabameri, Ahmadreza Mehrabian (correspondence <A_mehrabian@sbu.ac.ir>) & Massoud Sheidai,
Department of plant Sciences, Faculty of Biological Sciences, Shahid Beheshti University, P. O. Box, 1983969411,
Tehran, Iran.
KEY WORD: Boraginaceae; Onosma; morphology; micromorphology; taxonomy; SEM; Iran
ﻲﺳﺎﻨﺷ ﺖﺨﻳر تﺎﻌﻟﺎﻄﻣ ﻲﺳﺎﻨﺷ ﺖﺨﻳر ﺰﻳر و ﻲﺧﺮﺑ ﻪﻗﺪﻨﻓ و ﻞﮔ يﺎﻫ ﻪﻧﻮﮔ زاOnosma L. (Boraginaceae) ناﺮﻳا رد
،يﺮﻣﺎﻋ بﺮﻋ زﺎﻨﻬﻣﻚﻴﺗﺎﻤﺘﺴﻴﺳ ﺪﺷرا ﻲﺳﺎﻨﺷرﺎﻛ يﻮﺠﺸﻧاد -ناﺮﻳا ،ناﺮﻬﺗ ،ﻲﺘﺸﻬﺑ ﺪﻴﻬﺷ هﺎﮕﺸﻧاد ،ﻲﺘﺴﻳز مﻮﻠﻋ هﺪﻜﺸﻧاد ،ﻲﻫﺎﻴﮔ يژﻮﻟﻮﻛا
،نﺎﻴﺑاﺮﺤﻣ ﺎﺿرﺪﻤﺣاﻫﺎﻴﮔ مﻮﻠﻋ هوﺮﮔ رﺎﻳدﺎﺘﺳاناﺮﻳا ،ناﺮﻬﺗ ،ﻲﺘﺸﻬﺑ ﺪﻴﻬﺷ هﺎﮕﺸﻧاد ،ﻲﺘﺴﻳز مﻮﻠﻋ هﺪﻜﺸﻧاد ،ﻲ
،ﻲﻳاﺪﻴﺷ دﻮﻌﺴﻣناﺮﻳا ،ناﺮﻬﺗ ،ﻲﺘﺸﻬﺑ ﺪﻴﻬﺷ هﺎﮕﺸﻧاد ،ﻲﺘﺴﻳز مﻮﻠﻋ هﺪﻜﺸﻧاد ،ﻲﻫﺎﻴﮔ مﻮﻠﻋ هوﺮﮔ دﺎﺘﺳا
رد رﺬﺑ و ﻞﮔ ﻲﺳﺎﻨﺷ ﺖﺨﻳر ﺰﻳر و ﻲﺳﺎﻨﺷ ﺖﺨﻳر29 ﻲﻧاﺮﻳا ﻪﻧﻮﮔOnosma ﻪﺑ ﻪﺴﻳﺎﻘﻣ ترﻮﺻ ﻧوﺮﺘﻜﻟا ﭗﻜﺳوﺮﻜﻴﻣ زا هدﺎﻔﺘﺳا ﺎﺑ يا ياﺮﺑ هرﺎﮕﻧ ﻲ
ﺖﺳا ﻪﺘﻓﺮﮔ ترﻮﺻ يﺪﻨﺑ هدر رد ﺎﻫ نآ ﺶﻘﻧ ﺮﺑ هﮋﻳو ﺪﻴﻛﺎﺗ ﺎﺑ ﺎﻫ نآ ﻲﻜﻴﻣﻮﻧﻮﺴﻛﺎﺗ يﺎﻫ شزرا ﻦﻴﻴﺒﺗ و ﻲﺑﺎﻳزرا . ندﻮﻤﻧ ﻪﺘﺴﺟﺮﺑ ﺐﺒﺳ ﺎﻣ ﻪﻌﻟﺎﻄﻣ
ﺧ ﺰﻴﻟﺎﻧآ و ﻲﻠﺻا يﺎﻫ ﻪﻔﻟﻮﻣ ﻪﺑ ﻪﻳﺰﺠﺗ ﺰﻴﻟﺎﻧآ سﺎﺳا ﺮﺑ هﺪﺷ ﻪﻌﻟﺎﻄﻣ يﺎﻫ ﻪﻧﻮﮔ ﺐﻟﺎﻏ ﻲﻳﺎﺳﺎﻨﺷ ﺎﺑ طﺎﺒﺗرا رد ﻪﻗﺪﻨﻓ و ﻞﮔ تﺎﻔﺻ ﺖﻴﻤﻫا هﺪﺷ يا ﻪﺷﻮ
ﺖﺳا .ﺪﻧا هدﻮﻤﻧ ﺪﻴﻳﺎﺗ ار ﺎﻣ ﺞﻳﺎﺘﻧ ﻲﻟﻮﻜﻟﻮﻣ و ﻲﺳﺎﻨﺷ ﺖﺨﻳر ،ﻲﺳﺎﻨﺷ هدﺮﮔ تﺎﻌﻟﺎﻄﻣ ،هوﻼﻋ ﻪﺑ . ﻲﻳﺎﻬﻨﺗ ﻪﺑ ﻪﻛ ﺪﻧراد دﻮﺟو ﻲﻳﺎﻫ نﻮﺴﻛﺎﺗ ﻪﭼﺮﮔا
ﻮﻤﻧ فﺮﻃﺮﺑ و ﺺﻴﺨﺸﺗ رد ﻲﻤﻬﻣ راﺰﺑا ،ﺮﮕﻳد ﺖﻴﻤﻫا ﺎﺑ تﺎﻔﺻ ﺎﺑ ﻲﻫاﺮﻤﻫ ﻪﺑ تﺎﻔﺻ ﻦﻳا ﺎﻣا ،ﺪﻨﺘﺴﻴﻧ ﻲﻳﺎﺳﺎﻨﺷ ﻞﺑﺎﻗ هﺪﺷ ﺮﻛذ تﺎﻔﺻ سﺎﺳاﺮﺑ ند
ﺲﻨﺟ ﻲﻜﻴﻣﻮﻧﻮﺴﻛﺎﺗ يﺎﻫ ﻲﮔﺪﻴﭽﻴﭘ
Onosma ﺪﻨﺷﺎﺑ ﻲﻣ . ﻲﻟﻮﻜﻟﻮﻣ ﺪﻫاﻮﺷ ﺮﮔا)ﻲﺘﺳﻼﭘوﺮﻠﻛ و يا ﻪﺘﺴﻫ يﺎﻫ ﻲﻟاﻮﺗ ( ﻲﻳاﺮﮔاو هرﺎﺑرد ار ﺎﻣ ﺞﻳﺎﺘﻧ
Onosma orientalis وOnosma rostellata ﺖﺒﺴﻧﺲﻨﺟ ﻪﺑ Onosma ﻞﻘﺘﻨﻣ يﺮﮕﻳد يﺎﻫ ﺲﻨﺟ ﻪﺑ ﺪﻨﻧاﻮﺗ ﻲﻣ ﺎﻫ نﻮﺴﻛﺎﺗ ﻦﻳا ،ﺪﻨﻳﺎﻤﻧ ﺪﻴﻳﺎﺗ
ﺪﻧﻮﺷ .ﺳ زاﻲﻜﻴﻣﻮﻧﻮﺴﻛﺎﺗ ﻲﻨﻴﺑزﺎﺑ نﻮﻣاﺮﻴﭘ ﻲﻌﻣﺎﺟ شﺮﮕﻧ ﻲﻟﻮﻜﻟﻮﻣ و ﻲﺘﺧﺎﻨﺷ ﺖﺨﻳر ﺪﻫاﻮﺷ ﺎﺑ هاﺮﻤﻫ ﻞﮔ و رﺬﺑ تﺎﻔﺻ ﺐﻴﻛﺮﺗ ،ﺮﮕﻳد يﻮ
Onosma
ﺪﻧروآ ﻲﻣ ﻢﻫاﺮﻓ.
INTRODUCTION
Onosma L. is a genus of the tribe Lithospermeae
Dumort. belonging to the large family Boraginaceae,
comprising ca. 150 species (Mehrabian et al. 2012;
Kolarčik 2010; Cecchi & Selvi 2009) distributed
mainly in western, central Asia and in the

IRAN. J. BOT. 20 (2), 2014 M. Arabameri & al. 212
Mediterranean area (Jávorka 1906; Meusel et al. 1978).
These taxa are considered as a taxonomically difficult
group.Taxonomic treatments within the genus are
highly controversial and many closely related taxa were
described based on minor morphological differences.
Due to the similarities among the Onosma taxa, there
are many problems in their identification (Binzet &
Akcin 200 9). Moreover, the species are distinguished
on the basis of indumentum leading to several mistakes
in taxonomy of the genus in the past (Ball 1972; Maggi
2008). Due to mentioned complexities, Riedl (1978)
emphasized to providing the useful evidence in a new
classification. Khatoon et al (1994) did the first study
of the ultra-morphology of Boraginaceae including
Onosma. Moreover many authors have used trichome
micro-morphology (Khatamsaz 2002) and Palynology
(Binzet 2011; Qureshi & Qaiser 1987; Binzet & Orcan
2003a & b; Mehrabian et al. 2012) for character
evaluation in Onosma. Moreover corolla epidermis
ornamentations appear to be of great value in some
flowering plants i.e. Fabales (Christensen & Hansen
1998; Hammett et al. 1994), Rubus
(Sharifnia & Shakib
2012) and Onosma (Riedl 1978).
Nutlet morphology has provided useful evidence on
evolutionary classification and systematics of flowering
plants (Corner 1976). Besides, they have been used as
the most important character in Boraginaceae (Baillon
1888; Gurke 1893; Hilger 1985; Al-Shehbaz 1991;
Riedl 1997). Mentioned characters are valuable
evidence that can be used for delimitation of taxa in
Boraginaceae (Gray 1884; Langstrom & Chase 2002)
and specially in Onosma (Akcin 2009; Binzet & Akcin
2009).
A comprehensive study on morphological and
micro- morphological characters in Onosma is lacking.
Furthermore, the taxonomic values of these characters
is rather ambiguous. On the other hand, it is important
to understand the diversity that exists and the
prominent characteristics of the nutlet and flower when
identifying Onosma
species. The objective of the
present study is to clarify the taxonomic values of
mentioned characters and an specific emphasis was on
their role in classification and determination of Onosma
at infrageneric and interspecific levels. In addition,
attempts were made to identify unique characters for
each taxon and solving some taxonomic complexities.
MATERIALS AND METHODS
A-Micromorphological assessment
In our research, nutlet and petal morphology and
micro-morphology of 29 species of the genus Onosma
from Iran was studied on the basis of SEM and LM
microscopy techniques. Voucher data are presented in
table 1. Materials used for this study were taken from
wild populations as well as samples in HSBU, W and
TARI (herbarium abbreviations according to Thiers
2008).
A total of 1-3 populations for each taxon were studied
on petal and nutlet morphology and micro-morphology.
From each population, 1-3 plant specimens were used
and from each plant specimen at least 3-5 petals and 5-
10 nutlets were investigated and prepared for scanning
electron microscopy (SEM) and light microscopy
(LM).
For SEM studies, nutlets and petals were mounted
on stubs using double-sided adhesive tape. Samples
were coated with 12.5-15 nm of gold. Afterward,
coated samples were examined and photographed with
Cam Scan- MV 2300 Electron Microscope (fig.1).
The measurements taken from the petal (table 2) and
nutlet (table 3) were made by Image Tolls version 3.
The general terminology follows Riedl (1967); Davis
(1978) and Harris & Harris (2001). The measurements
taken from the flowers were 4 quantitative and 11
qualitative characters (table 2) as follows: calyx length,
calyx lobe width, calyx accrescent, calyx sections,
calyx trichome color, corolla lobe length, corolla
length, corolla shape, corolla color, corolla lobe shape,
corolla trichome, nectar trichome, exsertion of anther,
anthers adherence, corolla epidermis ornamentation and
nutlet measurements are including 5 quantitative and 4
qualitative characters including nutlet orientation,
nutlet shape, nutlet ventral keel, nutlet length, nutlet
width, nutlet epidermis ornamentation, nutlet beak
length/ beak length, nutlet length and nutlet
length/nutlet width.
B-Statistical analysis
For grouping of the studied species on the basis of
petal and nutlet morphology and micro-morphology,
mentioned data were standardized (mean = 0, variance
= 1). For morphological analysis, quantitative
characters were coded as multistate characters and used
for further analysis. Mentioned data were used for
cluster analysis, including UPGMA (Unweighted
Paired Group Using Average Method) and principal
coordinate analysis (PCoA), (Rholf 1972) and SPSS
ver. 9 software were used for statistical analysis.
(PCoA polts can be found in figs 3, 6 and 8 and cluster
diagrams are in figs 2, 5, 7).
RESULTS
A-Flower
Floral morphology and micro-morphology showed
high variability among the mentioned taxa. The flower
characters are discussed below:

213 Morphological Studies on Onosma IRAN. J. BOT. 20 (2), 2014
Table 1. List of species used in this study.
S
p
ecies Localit
y
Collector & Voucher
Onosma cornuta Riedl Kordestan, bi
j
ar, 1931
m
Mehrabian, HSBU-2010274
Onosma cornuta Riedl Kermanshah, Between Eslam Abad and Ivan,
1420m Mozaffarian, TARI-74735
Onosma straussii Riedl Markazi, Arak, Gavar, 1990
m
Mehrabian, HSBU-2010212
Onosma rasche
y
ana Boiss. Zan
j
an, Mahneshan, Bel
g
he Mt, 1298
m
Mehrabian, HSBU-281
Onosma rasche
y
ana Boiss. Azarba
yj
an, Oromieh, 1750
m
Zehzad, HSBU-2010310
Onosma alborosea Fisch. Khoram Abad, 1650
m
Riedl & Laraze, W-103
Onosma microcar
p
a DC. Azarba
yj
an,Khalkhal, 1793
m
Mehrabian, HSBU-2010252
Onosma microcar
p
a DC. Azarbai
j
an, Kho
y
to Ghotor, 1700
m
Mehrabian, HSBU-2010242
Onosma microcar
p
a DC. Tehran, Dizin, 2316
m
Mehrabian, HSBU-2010244
Onosma kurdica Te
pp
ne
r
Kordestan , Hamze Arab Mt, 2510
m
Lamond&Termeh, WU-06112
Onosma kilouyensisBoiss. Kermanshah,Gahvare to kuzaran, 1469m Mehrabian&Mohamadi, HSBU-
260
Onosma sericea willd. Kermanshah,Gardaneh-e-Ghala
j
eh, 1923
m
Mehrabian, HSBU-265
Onosma sericea willd. Kermanshah, Bi
j
ar, 1700
m
Mehrabian, HSBU-2011276
Onosma nervosa Riedl Kordestan,15 Km Marivan to sananda
j
, 1878
m
Mozaffarian, TARI-74656
Onosma dasytricha Boiss. 30 Km Noor Abad, near Robat, 1600m Assadi &Abuhamze, TARI-
38361
Onosma rostellata Lehm. Kermanshah ,Pave, ba
y
an
g
an, 1421
m
Mehrabian, HSBU-2010225
Onosma stenosiphon Boiss. Tehran, 13km Firuzkuh toward Semnan, 2000m Assadi& Mozaffrian, TARI-
35237
Onosma stenosi
p
hon Boiss. Kerman , Bi
j
ar Mt, 1857
m
Kanani, HSBU-2011-121
Onosma elwendica wettst. Kermanshah, Ghardaneh-e-Ghala
j
eh, 1923m Mehrabian, HSBU-268
Onosma dichroanta Boiss. Golestan, Golestan National Park, 1500
m
Heidari, et al. HSBU-2007300
Onosma sabalanica Poner
t
Azarba
yj
an, Meshkin Shahr, Sabalan Mt, 4762
m
Mehrabian, HSBU-2010-201
Onosma sabalanica Poner
t
Azarbai
j
an, Arasbaran Protected Area,2000m Zehzad, HSBU-83898
Onosma olivieri Boiss. Kermanshah, 2103
m
Zar
g
ani, IRAN-2901
Onosma olivieri Boiss. Kermashah, Nosoud to nodesheh, 1800
m
Mehrabian, HSBU-2010900
Onosma longiloba Bge. Semnan, 20km Mohamad abad from
Firuzkuh,22.5.2012, Pahlevani, 2100
m
Iranshahr, HSBU-2010100
Onosma
p
ach
yp
oda Boiss. Azarabi
j
an, Tabriz, Mishoda
g
h Mt. 1800
m
Mehrabian, HSBU-2010602
Onosma bulbotrica DC. Zan
j
an,nik Pe
y
to Mahneshan, 1608
m
Mehrabian, HSBU-2010290
Onosma bulbotrica DC. Kermanshah, Gilane-Gharb, Ghala
j
eh, 1630m Mehrabian, HSBU-2010263
Onosma bulbotrica DC. Ghazvin, Takestan to Abhar, 1400m Mehrabian, HSBU-2010254
Onosma kotschyi Boiss. Esfahan, Semirom Padena, 2397m Mehrabian &Mostafavi, HSBU-
123
Onosma orientalis L. Fars between Shiraz and Kazerun, 2162m Ghorbani&Habibi, HSBU-
2010225
Onosma orientalis L. Bushehr, Boraz
j
an, Dalaki, Tan
g
-e Eram, Mozaffarian, TARI-74152
Onosma subsericea Fre
y
n Azarba
yj
an, between kiwi to Khal
k
hal, 1920
m
Rechin
g
er, W-07869
Onosma subsericea Freyn Azarbaijan, Khalkhal to Givi, 1800m Wendelbo& Assadi, SBU-2010-
232
Onosma hebebulba DC. Kermanshah, Eslam Abad to Ilam, 1437
m
Sharif, TARI-5329
Onosma as
p
errima Bornm. Fars , Abade, 2007
m
Behbudi, W-7148
Onosma as
p
errima Bornm.
N
orabad, Doshman ziari, 2500
m
Mehrabian, TARI-45722
Onosma bodeana Bornm. Tehran , Sohanak, 1832
m
Mehrabian, HSBU-272
Onosma
g
aubae Bornm. Semnan,Shahmirzad,nizva Mt, 2115m Zar
g
ani, TARI-2794
Onosma
g
aubae Bornm. Alborz,
N
ear the Amirkabir Dam, 1900
m
Mehrabian, HSBU-
Onosma armena DC. Zan
j
an, Mahneshan, 1296
m
Mehrabian, HSBU-2010654
Onosma
p
lat
yp
h
y
lla Riedl Lorestan, Ali
g
udarz, Ghalikoh, 3121
m
Rechin
g
er, W-05714
Onosma platyphylla Riedl Lorestan, 45km Dorood, 1600m Assadi& Mozaffarian, TARI-
37007
Onosma macrophylla
Bornm. Kurdestan, Baneh, 2200m Rechinger, W-05718

IRAN. J. BOT. 20 (2), 2014 M. Arabameri & al. 214
Calyx
Calyx length is varied between 6mm (O. rostellata)
to 22.5mm (O. kurdica). Calyx lobe width: 0.74mm (O.
rostellata) to 6.42 mm (O. straussii). Calyx with non-
accrescent strucure is most common, but some showed
accrescent structure as follow : O. pachypoda, O.
cornuta, O. dichroantha. Calyx usually is sectioned at
the base, but O. cornuta and O. dichroantha lack this
trait and are divided near to base.
Corolla morphology
Corolla length showed variation between 10 mm
(O. kotschyi) to 33 mm (O. dichroantha). But
dominantly it is between 15-24 mm. Corolla lobe
length is ranged between 0.8 mm (O. dasytricha, O.
rostellata) to 6 mm (O. longiloba). Moreover a wide
range of taxa showed wide corolla lobe, but linear
(long) lobes were only found in O. rostellata and O.
platyphylla. Corolla dominantly show yellow color, but
there are other colors that includ: white (O.
bulbotricha, O. Gaubae), red-pink (O. rascheyana, O.
straussii), blue-violet (O. dichroantha, O. dasytricha).
Corolla shape shows high variability among studied
species: tubular (O. rostellata, O. orientalis, O.
sabalanica), campanulate (O. olivieri , O. bulbotricha),
tubular-campanulate (O. kilouyensis. O. platyphylla),
tubular-clavate (O. Gaubae, O. alborosea), funnel-
campanulate (only O. starussii), clavate (O. sericea, O.
rascheyana ), Moreover two types of nectary were
observed: most species showed glaber nectar, but O.
bulbotricha and O. stenosiphon
are with tomentose
trichomes and O. cornuta, and O. orientalis by villous
trichomes. Furthermore corolla epidermis represent
glaber (O. sabalanica, O. Gaubae) to tomentose (O.
dichroantha, O. bodeana) structure.
Anthers are in 3 groups: they are predominantly
connected to the base. Other patterns are connected
over the stamen (only in O. rostellata) and free (O.
asperrima
and O. kotschyi).
Data on type of nectary, corolla trichomes and
continuity of anthers have not been recognized
previously in the treatments of the Iranian Onosma
species.
Corolla micromorphology
Five different patterns of petal epidermis
ornamentations are observed in studied Onosma species
as below:
Type 1: This type is characterized by rugose
ornamentation that is covered with coarse reticulate
lines. Mentioned type is seen in 3 subsections as
follows: sect. Asterotrich a(O. hebebulba), sect.
Haplotricha (O. bodeana) and sect. Heterotricha ( O.
nervosa ) (figs.1.1-3).
Type 2: This type is characterized by farinaceous
ornamentations that is covered with small granules and
is finely mealy. This type was observed in O. longiloba
(fig.1.10) and O. kurdica
(fig. 1.11).
Type 3: This type is characterized by Papilllate
(verrucate/tuberculate) ornamentations, having minute,
rounded protuberances. This type is observed in O.
rascheyana (fig1.12) and O. sabalanica (figs. 1.13,14).
Type 4: This type is characterized by aculeate (prickly)
ornamentations, with sharp prickles, no spine, and no
thorny appendages. Onosma platyphylla
(figs.1.15,16)
is the only species that shows mentioned characters.
Type 5: This type is characterized by scurfy
ornamentations, having small granules and is covered
by tomentose trichomes. A wide range of taxa are
classified in this type O. sericea
(fig.1.17), O. Gaubae
(fig.1.18), O. rostellata
(fig.1.19), O. orientalis
(fig.1.20) O. asperrima (fig.1.21), O. cornuta
(fig.1.22), O. kilouyensise (fig1.23), O. pachypoda
(fig1.24), O. macrophyla (fig1.25), O. straussii
(fig.1.26), O. albo-rosea (fig.1.27) and O. armena
(fig.1.28). Petal epidermis ornamentation is reported
here for the first time on Iranian Onosma.
The UPGMA trees of flower characters (fig. 2) showed
a relatively high cophenetic correlation (r >0.80). In
general, two main clusters are observed. PCA and
Cluster analysis didn’t show clearly sectional divisions.
B- Nutlet morphology
Nutlts Size varies greatly in studied taxa. Nutlet
length varies from 1.71 mm (O. rascheyana) to 8.18
mm (O. alborosea). But dominant range is 4-5.5 mm.
Nutlet width showed variation between 2 mm (O.
orientalis ) to 5.5 mm (O. olivieri). Nutlet beak length
varies from 0.8 mm (O. orientalis) to 3.08 mm (O.
rascheyana). Moreover beak length/nutlet length varies
between 0.12 mm (O. olivieri) to 0.43 mm (O.
rostellata) and nutlet length/nutlet width varies
between 0.59 mm (O. rascheyana) to 2.08 mm (O.
sabalanica).
Nutlet shapes are broad obvoate in O. nervosa (fig.
4.3), O. cornuta (fig.4.21)
O. rostellata (fig.4.31), O.
kilouyense (fig.4.15), O. kotschyi (fig.4.39), O.
microcarpa (fig.4.11), O. albo-rosea (fig.4.9), O.
elwendica (fig.4.41), O. olivieri (fig.4.17) ovate in O.
dichroantha (fig. 4. 5), O. orientalis (fig.4.33), O.
sabalanica (fig.4.23), O. sericea (fig.4.29), O.
stenosiphon (fig.4.25), O. rascheyana (fig.4.1), O.
dasytricha (fig.4.19), O. longiloba (fig4.37), pyramid
in O. bulbotricha (fig.4.27) and O. straussii (fig. 4. 7),
and rhombic in O. kurdica (Fig. 4.13). Broad ovate is
the most common shape in the studied species.
Moreover nutlet orientation is incurved (only in O.
orientalis),(fig. 4.33) and straight (in other species).

215 Morphological Studies on Onosma IRAN. J. BOT. 20 (2), 2014
Fig. 1: Corolla epidermis ornamentation (1-28) and nectar trichomes (29-32). 1, O. hebebulba; 2, O. bodeana; 3-O.
nervosa; 4, O. bulbotrica; 5, O. dichroanta; 6, O. microcarpa; 7, O. stenosiphon; 8, O. subsericea; 9,O. elwendica;
10, O. longiloba; 11, O. kurdica; 12, O. rascheyana;13-14, O. sabalanica; 15-16, O. platyphylla; 17, O. sericea;
18,O. gaubae; 19, O. rostellata; 20, O. orientalis.
1-Onosma hebebulba 2-Onosma bodeana 3-Onosma nervosa 4-Onosma bulbotrica
200
µ
200
µ
200
µ
200
µ
5-Onosma dichroanta 6-Onosma microcarpa 7-Onosma stenosiphon 8-Onosma subsericea
200
µ
200
µ
200
µ
200
µ
9-Onosma elwendica 10-Onosma longiloba 11-Onosma kurdica 12-Onosma rascheyana
200
µ
200
µ
200
µ
200
µ
13-Onosma sabalanica 14-Onosma sabalanica 15-Onosma platyphylla 16-Onosma platyphylla
50
µ
200
µ
50
µ
200
µ

IRAN. J. BOT. 20 (2), 2014 M. Arabameri & al. 216
Fig. 1: Continued Corolla epidermis ornamentation (21-28) and nectar trichomes (29-32). 21, O. asperrima; 22,O.
cornuta; 23, O. kilouyensis; 24,O. pachypoda; 25, O. macrophyla; 26, O. straussii; 27, O. albo-rosea; 28, O.
armena; 29, O. straussii; 30,O. cornuta; 31, O. stenosiphon; 32,O. bulbotricha.
Furthermore nutlet ventral keel can be found in O.
rascheyana (fig. 4.1), O. kurdica (fig.4.13), O.
elwendica (fig.4.41), O. nervosa (fig4.31).
Nutlet micromorphology
Six types of nutlet epidermis ornamentations are
defined for the studied taxa as follows:
Type I: Epidermal layer is characterized by rugose or
bullate shape covered with coarse reticulate lines. This
type can be divided to two subtypes: first subtype is
seen in . rascheyana (fig.4. 2), O. nervosa (fig. 4.4), O.
dichroanta (fig.4.6), O. straussii (fig.4.8), O. albo-
rosea (fig.4.10), O. microcarpa (fig.4.12), O. kurdica
(fig.4.14), O. kilouyense (fig.4.16), O. olivieri
(fig.4.18), and second one only was observed in O.
dasytricha (fig.4.20).
Type II: Epidermal layer is wrinkled. It is observed in
O. cornut (fig.4.22) and O. sabalanica (fig.4.24).
Type III: Epidermal layer is rugose-striate including
parallel coarse reticulate lines. Mentioned type is
observed O. stenosiphon (fig.4.26), O. bulbotricha
(fig.4.28) and O. sericea (fig.4.30).
Type IV: Epidermal layer is farinaceous (granular) that
is finely mealy and covered with small granules.
Onosma rostellata (fig.4.32) shows mentioned
structure.
Type V: Epidermal layer is papillate (tuberculate or
verrucate) having minute, rounded protuberances.
Onosma orientalis (fig. 4.34) is classified in this type.
Type VI: Epidermal layer is relatively to completely
smooth, O. elwendica (fig.4.42), O. longiloba
(fig.4.38), O. pachypoda (fig.4.36) and O. kotschyi
(fig.4.40) are in this type.
The UPGMA trees of nutlet evidence (fig. 5) showed a
relatively high cophenetic correlation (r >0.80). In
general, three main clusters can be shown. PCoA an
Cluster analysis showed a relatively acceptable
sepration at sectional level.
DISCUSSION
Measuring the size of plant organs and parts is
important in description and identification. Within each
taxon, the size of the flower parts is very variable.
Calyx length and width were used in some references

Table 2. Measurements taken from the flower characters. Calyx accrescent (ab: absence, ex: exsistence), sectioned calyx, (1: in base, 2: close to base);
corolla color (1: yellow, 2: white, 3: red-pink, 4: blue-violet), corolla shape (1: tubular, 2: campanualte, 3: tubular campanulate, 4: tubular-clavate, 5:
funnel-campanulate, 6: clavate), corolla lobe (1: broad, 2: linear), corolla epidermis ornamentation (I:reticulate, II:farinaceous, III:papillate, IV:aculeate,
V:scurfy); anther exsertion (1: inside, 2: ouside), anther adhrence(1: free, 2: at base, 3: along side); nectar trichome (ab: absence, ex: exsistence ).
Characters
Species
Calyx
length
(mm)
(A)
Calyx lobe
width
(mm)
(B)
Calyx
accresce
nt
(C)
Calyx
section
(D)
Calyx
trichome
color
(O)
Corolla
lobe
length
(mm)
(E)
Corolla
length
(mm)
(F)
Corolla
shape
(G)
Corolla
color
(H)
Corolla
lobe
shape
(M)
Anther
exsertion
(I)
corolla
epidermis
ornamentati
on
(J)
Anthers
Adherenc
e
(k)
Corolla
trichome
(L)
Nectar
tricho
me
(N)
O. rostellata 6
5-8.5
0.74
0.6-0.9 ab 1 0 0.8±0.2 10
7-11 1 3 1 2 V 3 ex ab
O. orientalis 10
6-12
0.8
0.7-1 ab 1 0 2.5±0.5 12
10-13 1 4 1 2 V 2 ab ex
O. asperrima 8.5
6-12
1.14
1-1.3 ab 1 0 1±0.2 18
16-22 3 1 1 1 V 1 ex ab
O. bodeana 20.5
18-22
2.16
2-2.5 ab 1 0 1.8±0.2 25
19-27.5 3 1 1 1 I 2 ab ab
O. bulbotrica
17.5
12.2-
18.9
1.5
1.2-2 ab 1 0 2±0.35 20
17-23 1 2 1 1 I 2 ab ex
O. cornuta 11
9-14
4.71
4-7 ex 2 0 1.7±0.3
14.5
14-16 3 1 1 1 V 2 ex ex
O. dichroanta 20
18-25
2.86
2.5-3 ex 2 0 2±0.35 33
21-38 3 4 1 1 I 2 ab ab
O. Gaubae 13.5
12-15
1.1
0.9-1.3 ab 1 0 2±0.35 17
15-18 4 1 1 1 V 2 ex ab
O. kilouyensis 15
14-17
5.85
5-8 ab 1 0 1.8±0.3 20
18-21 3 3 1 1 V 1 ex ab
O. kotschyi 8.5
7-11
1.2
1.1-1.3 ab 1 0 2±0.35 9
7-10 1 1 1 1 _ 1 ex ab
O. longiloba 11
10-13.5
3.85
3-5 ab 1 0 6±0.6 9
8-12.5 1 4 2 1 II 1 ab ab
O. microcarpa 10
8-14
1.07
1-1.3 ab 1 1 1.1±0.2
18.5
15-25 3 1 1 1 I 2 ex ab
O. pachypoda 17
14-18
0.81
0.7-1 ex 1 0 1.5±0.2 26
24.5-29 6 4 1 1 V 1 ex ab
O. platyphylla 10
8-13
0.97
0.8-1.2 ab 1 0 3±0.4 13
12-15 3 1 2 2 IV 1 ex ab

Table 2. Continued.
O. sabalanica 16-15 4.85
2-7 ab 1 1 2.8±0.4 17
15-20 2 1 1 1 III 2 ex ab
O. sericea 14
12-16
7.6
7-9 ab 1 0 1.9±0.3
17.5
15-20 6 1 1 1 V 2 ex ab
O. stenosiphon 11.2
7.3-14.2
1.44
1-2 ab 1 0 1±0.2 12
7.3-14 1 4 1 1 I 2 ab ex
O. subsericea 20.8
18.3-25
2.61
2.3-3 ab 1 0 1±0.2 26.5
22-28 2 1 1 1 I 3 ex ab
O. elwendica 16
14-17
5.85
5-8 ex 2 0 2±0.35 19
18-20.5 3 4 1 1 I 2 ab ab
O.
macrophyllla
11
10-13.5
4.71
4-6 ab 1 0 1.5±0.2 18
17-19 2 1 1 1 V 2 ex ab
O. nervos 11
10-13
1.7
0.8-2 ex 1 1 1.3±0.2 13
12-15 2 3 1 1 I 2 ab ab
O. olivieri 15.5
13-16.6
2.5
2-2.9 ab 1 0 1.2±0.2 21
20-23 2 1 1 1 _ 2 ex ab
O. straussii 9.5
7-10
6.42
5-7 ab 1 0 2±0.3 17.5
16-20 5 3 1 2 V 2 ex ab
O. albo-rosea 15
14-17
8.8
5-11 ab 1 0 2±0.4 24
22-27 4 3 1 1 V 2 ex ab
O. armena 9.5
9-10
0.85
0.8-0.9 ab 1 0 1.8±0.2 18
16-20 1 1 1 1 V 2 ab ab
O. dasytricha 18
16-20
3.94
2-4.7 ab 1 0 0.8±0.2 24
20-28 4 4 1 1 _ 2 ex ab
O. hebebulba 16
15-18
1.6
1.2-1.9 ab 1 0 1.8±0.2 15
14-17 3 1 1 1 I 2 ab ab
O. kurdica 22.5
20-23.5
0.9
0.8-1 ab 1 0 0.8±0.2 25
23-27 6 3 1 2 II 2 ab ab
O.rascheyana 13.5
12-16.5
1.5
1-2.1 ab 1 0 1±0.2 22.5
20-25.2 6 3 1 1 III 1 ab ab

219 Morphological Studies on Onosma IRAN. J. BOT. 20 (2), 2014
Fig 2. UPGMA tree of studied taxa based on flower character.
Fig 3. PCoA plot of studied taxa based on flower evidence.

IRAN. J. BOT. 20 (2), 2014 M. Arabameri & al. 220
Fig. 4: nutlet micromorphology of studied species. 1-2, Onosma rascheyana; 3-4 O. nervosa; 5-6, O. dichroanta; 7-
8, O. straussii; 9-10, O. albo-rosea;11-12, O. microcarpa; 13-14, O. kurdica; 15-16 O. kilouyensis; 17-18, O.
olivieri; 19-20, O. dasytricha; 21-22, O. cornuta; 23-24, O. sabalanica.
1-Onosma rasche
y
ana 2-Onosma rasche
y
ana 3-Onosma nervosa 4-Onosma nervosa
5-Onosma dichroanta 6-Onosma dichroanta 7-Onosma straussi
i
9-Onosma albo-rosea 10-Onosma albo-rosea 11-Onosma microcarpa 12-Onosma microcarpa
13-Onosma kurdica 14-Onosma kurdica 15-Onosma kilouyens 16-Onosma kilouyensis
18-Onosma olivieri 19-Onosma dasytricha 20-Onosma dasytricha
2m
50µ
200µ
50µ
2m
500µ
17-Onosma olivieri
2m
200µ
200µ
8-Onosma straussii
200µ
2m
200µ
2m
200µ
2m
200µ
21-Onosma cornuta 22-Onosma cornuta 23-Onosma sabalanica 24-Onosma sabalanica
1m
200µ
1m
50µ
3m
5m
3m
5m

221 Morphological Studies on Onosma IRAN. J. BOT. 20 (2), 2014
Fig. 4: Continued. nutlet micromorphology of studied species. 25-26, O. stenosiphon; 27-28, O. bulbotrica; 29-30,
O. sericea; 31-32, O. rostellata; 33-34, O. orientalis; 35-36, O. pachypoda; 37-38, O. longiloba; 39-40, O. kotschyi;
41-42, O. elwendica.
25-Onosma stenosiphon 26-Onosma stenosiphon 27-Onosma bulbotrica 28-Onosma bulbotrica
29-Onosma sericea 30-Onosma sericea 31-Onosma rostellata 32-Onosma rostellata
33-Onosma orientalise 34-Onosma orientalise 35-Onosma pachypoda 36-Onosma pachypoda
37-Onosma longiloba 38-Onosma longiloba 39-Onosma kotschyi 40-Onosma kotschyi
2m 200
µ
2m 200
µ
200
µ
1m
200
µ
2m
1m 100
µ
2m 50
µ
200
µ
1m
200
µ
2m
41-Onosma elwendica 42-Onosma elwendica
200
µ
2.5m

IRAN. J. BOT. 20 (2), 2014 M. Arabameri & al. 222
Table 3. Measurements taken from the nutlet characters. Nutlet orientation (1:straight, 2: incurved), nutlet shape (ov: ovate, wo:
wide ovate, py: Pyramid:, rh: rhombic), nutlet ventral keel (ex: exsistence, A. absence, M: middle), nutlet epidermis
ornamentation (typeI: rugose, typeII: wrinkled, type3: rugose-striate, type4: farinaceous, type5: papillae, type 6: smooth).
Character
p
ecies
Nutlet
orienta
tion
(A)
Nutlet
shape
(B)
Nutlet
ventral
keel
(C)
Nutlet
epidermis
ornament
ation
(D)
Nutlet
width
(mm)
(E)
Nutlet
length
(mm)
(F)
Nutlet
beak
length
(mm)
(G)
Beak
length/nutl
et length
(H)
Nutlet
length/nutlet
width
(I)
O. rostellata
Lehm. 1 wo ex IV 3.37
3-3.5 3.38
3-3.6 1.43±0.2 0.43
0.39-0.47 1.00
0.85-1.2
O. orientalis 2 ov ab V 2
1.5-1.8 2.25
2.3-2.5 0.8±0.2 0.44
0.32-0.34 1.55
1.27-1.66
O. bulbotrica 1 py m III 4.47
4-4.9 4.67
4-5 0.86±0.2 0.18
0.17-0.21 0.92
0.81-1.25
O. cornuta 1 Wo ex II 2.8
2.6-3 4.15
4-4.3 1±0.2 0.26
0.23-0.27 1.27
1.15-1.72
O. dichroanta 1 ov ab I 2.97
2.9-3 4.33
4-5 1.23±0.2 0.28
0.24-0.30 1.46
1.33-1.72
O. kilouyensis 1 wo ex I 3.83
3.5-4 5.28
5-6 1±0.2 0.20
0.16-0.21 1.29
1.25-1.71
O. longiloba 1 ov ab VI 2.23
2-2.5 3.46
3.2-3.6 0.87±0.2 0.27
0.24-0.27 1.46
1.28-1.80
O. kotschyi 1 wo ab VI 2.11
1.5-2.5 3.77
3.5-4 1.82±0.2 0.49
0.44-0.52 1.82
1.4-2.6
O. microcarpa 1 wo ex I 2
1.7-2.5 2.57
2.5-3 0.9±0.2 0.34
0.3-0.36 1.19
1-1.76
O. pachypoda 1 wo ab VI 3.35
3.2-3.5 5.5
5.3-5.7 1.6±0.2 0.29
0.28-0.30 1.64
1.51-1.78
O. sabalanica 1 ov m II 2
1.8-2.1 4.06
3.7-4.3 1±0.2 0.27
0.23-0.27 2.08
1.76-2.38
O. sericea 1 ov ex III 3.64
3-4 4.8
4.4-5 1.62±0.2 0.34
0.32-0.36 1.33
1.1-1.66
O. s
t
enosiphon 1 ov ab III 2.06
2-2.2 4.08
4-4.2 0.83±0.2 0.20
0.19-0.20 1.95
1.81-2.10
O. straussii 1 py m I 4.3
4.1-4.5 5.47
5.3-5.7 1.2±0.2 0.22
0.21-0.22 1.36
1.20-1.33
O. albo-rosea 1 wo ab I 5.78
5.5-6 8.18
7.8-8.3 0.93±0.2 0.12
0.11-0.12 1.37
1.30-1.50
O. dasytricha 1 ov m I 5.21
5-5.4 6.63
6-7 2.72±0.3 0.40
0.38-0.45 1.44
1.11-1.5
O. rascheyana 1 ov ab I 2.88
2.8-3 1.71
1.5-2 3.07±0.4 1.67
1.53-2.04 0.59
0.50-0.71
O. kurdicua 1 rh ex I 4
3.8-4.2 8
7.5-8.3 3.2±0.4 0.4
0.38-0.42 0.8
1.78-2.18
O. elwendica 1 wo ex VI 3.86
3.5-4 5.28
5-6 1.36±0.2 0.18
0.22-0.27 1.36
1.25-1.71
O. nervosa 1 wo ex I 3.8
3.7-4 4.04
3.9-4.3 1.31±0.2 0.32
0.30-0.33 1.06
0.97-1.16
O. olivieri 1 wo m I 5.5
5.3-5.7 7.5
7.3-7.7 2±0.3 0.26
0.25-0.27 1.36
1.28-1.45
(Shishkin 1953) as a weak-effect character, but
along with other diagnostic characters, they can be used
in determination of Onosma taxa. Our study revealed
taxonomic significance of the corolla length in some
Onosma taxa, for example O. kotschyi (corolla length
ca. 9mm) and O. asperrima (corolla length ca. 25mm)
as closely related species, can be distinguished based
on this character. Moreover Riedl (1967) referred to it
as a valuable character. Furthermore Peruzzi &
Passalacqua. (2008) and Mehrabian et al. (2012) used
corolla length for interpretation of population
variability in Onosma.
Six kinds of corolla shapes can be distinguished in
studied taxa. In our study it appeared as an important
diagnostic character at species level in some taxa and
as a clear character is able to distinguish some groups
that faced taxonomical complexities. On the basis of
this character O. sericea, O. cornuta and O. elwendica
as a complex taxa are determinable. Moreover Riedl
(1967) and Davis (1978) confirmed the importance of
corolla shape in taxonomy of the genus.
Wodehouse (1935) and Lee (1979) have reported a
direct correlation between corolla and pollen size. But
Binzet (2011) in Onosma proved that there was no
correlation between corolla size and pollen size.
Mehrabian et al. (2012) proved Some species with
large corolla i.e. O. alborosea and O. bulbotrichum
have longer pollens (18-21µm and 15-18 µm

223 Morphological Studies on Onosma IRAN. J. BOT. 20 (2), 2014
Fig 5. UPGMA tree based on nutlet character.
Fig. 6. PCoA plot of studied taxa based on nutlet character.

IRAN. J. BOT. 20 (2), 2014 M. Arabameri & al. 224
respectively) but some taxa , i.e. O. microcarpa and
O. rostellata with shorter corolla showed larger pollen
(16-19µm and 16-22 µm respectively). Our results
confirm those provided by Binzet (2011).
Corolla color varies considerably. Besides it showed
some variations in diverse ecological (especially soil
and climate) conditions (Mehrabian 2011) in different
populations. For instance O. pachypoda and O.
bodeana showed variability in diverse ecological
habitats. Furthermore, several species showed
similarity in corolla color. This character is hardly
reliable but as a subsidiary diagnostic character along
with other important characters can be used to
distinguish Onosma taxa.
In the present study, the nectary trichomes of the
studied taxa appeared to be diagnostic in restricted taxa
i.e. in O. straussii (fig.1.29), O. cornuta
(fig.1.30), O.
stenosiphon (fig.1.31) of the sect. Onosma and O.
orientalis (fig.1.32) of the sect. Protonosma. Therefore
as a weak delimitation character can be used along with
other characters for definition of limited taxa.
Corolla lobe is hardly reliable, nevertheless, O.
longiloba (ca. 6mm) and O. paltyphylla (ca. 3.5mm)
are distinguished from other taxa by short corolla lobe
(less than 2.8 mm). Calyx accrescent was observed in
two subsections for example O. cornutua, O.
dichroantha, O. pachypoda belonging to subsect.
Haplotricha, O. elwendica and O. nervosa from
subsect. Heterotricha. Hence it can be used as a
diagnostic character for delimitation of some closely
and hardly distinguishable taxa. For instance based on
calyx accrescent O. elwendica can easily be
distinguished from its close species O. olivieri. Calyx
division did not show taxonomic importance, most
species of Onosma have shown division at the base but
only O. cornuta, O. dichroantha belonging to subsect.
Haplotricha and O. elwendica from subsect.
Heterotrica represent calyx division near the base. Also
Calyx trichome color showed weak valuae in taxonomy
of Onosma, because only O. sabalanica (subsect.
Haplotricha) , O. elwendica and O. nervosa (subsect.
Asterotricha) showed this properties. Riedl (1968);
Davis (1978); Shishkin (1953); Akcin (2009) used
mentioned features as a diagnostic characters in
Onosma .
Anther features have proved their abilities for
delimitation of Onosma species. Anthers are connected
along side in sect. Podonosma, but they are free to
connected at the base in other sections. Nevertheless,
they are diagnostic characters as follow : O. asperrima
with free anthers is distinguishable from very closely
relatad species O. kotschyi with connected anthers at
base and O. pachypoda with free anthers is diffreable
from O. bodeana with connected anthers at base.
Onosoma rostellata of sect. Protonosma with
connected anthers in along sides, ellipsodal pollen is
different from the other Onosma taxa which have
prolate or subprolate pollen shape (Mehrabian et al.
2012) differences in molecular characters (Mehrabian
et al. 2011) and the granulate ornamentation in nutlet
surface are the diagnostic characteristics that brings the
idea of transfering this onosma species to a different
genus. Moreover Onosma orientalis (section.
Podonosma Boiss.) with divergence in morphology,
differences in molecular (Mehrabian et al. 2011),
trichome characters and tuberculate ornamentation of
nutlet surface shows differences from the other
Onosma taxa. Complementary studeis are needed to
prove its taxonomic status within Boraginaceae, i.e.
phylogenetic stdies besed on chloroplast and nuclear
genome sequences.
Corolla epidermis showed five ornamental patterns.
These detailed patterns showed clearly boundaries of
species. Some types are unique characteristcs for
special species. Type IV was only observed in O.
platyphylla (subsect. Haplotricha) and type I was
observed only in sect. Onosma. Moreover type III and
type II can be seen in sect. Onosma. On this baisis,
these features accompained by other morphological
characters are useful for species determination and
solving some taxonomical complexities in Onosma.
Akcin (2009) have used petal morphology as a valuable
character.
Some nutlet studies in Boraginaceae (Selvi et al.
2006) proved nutlet morphology to be important for
differentiation of these taxa at the infrageneric
categories.
Nutlet orientation follows two patterns: straight, and
incurved. Onosma orientalis (sect.Podonosma) showed
high divergence compared to other onosma sections,
therefore as a unique chracter it is valuable for
infrageneric delimitation in Onosma.
Nutlet size and witdh in some cases appeared to be
important among studeid species. Onosma kurdica, O.
bulbotricha and O. starussii showed specific shapes
that help to distinguish them from the other species.
Moreover some species showed very different size for
example O. alborosea that is differable from its closely
related taxon O. dasytricha. Nutlet ventral keel showed
diversity in studied species, due to its appearance in all
sections and subsections, this character is not useful
for delimitation of species but as a valuable character
showed diagnostic ability to define closely related
species in complicated species groups. For example O.
kurdiua (by ventral Keel) and O. rascheyana (no
ventral keel) are distinguished.
Nutlet ornamentation represent five patterns in
studied taxa. Each section have specific pattern as

225 Morphological Studies on Onosma IRAN. J. BOT. 20 (2), 2014
Fig. 7: Combined UPGMA tree based on nutlet and flower characters.
Fig. 8. Combined PCoA plot based on nutlet and flower characters.

IRAN. J. BOT. 20 (2), 2014 M. Arabameri & al. 226
follow: O. orientalis (Sect. Podonosma) by Papillate
pattern and O. rostellata (Sect. Protonosma) with
farinaceous pattern. Furthermore there are rugose
(O.rascheyana, O. nervosa), rugose-striate (O.
pachypoda, O. longiloba and smooth pattern (O.
bulbotricha, O. stenosiphon) in sect.Onosma.
Accordingly, this character is useful in recognition of
Onosma at sectional level, and as a valuable character
in species delimitation. Moreover some complex
species groups (groups with many overlapping
morphological features) such as O. bulbotricha, O.
straussii, O. cornuta and O. sericea are distinguisable
based on mentioned patterns. Besides, some closely
related taxa like O. rascheyana and O. kurdica can be
sepretaed by this character. PCoA and cluster analysis
showed their efficiency for sectional delimitation, but
not useful to seprate the subsections. Binzet and Akcin
(2009) and Akcin & Binzet (2011) had the same
results.
Cluster analysis based on palynological (Mehrabian
et al. 2012), morphological and molecular (Mehrabian
et al. 2012) evidence confirmed our logic on diagnosis
value of studied charcters at sectional and species level.
The results of the present study highlight the
importance of the morphology and micro-morphology
of floral and nutlet characters in relation to
identification and delimitation of most of the studied
species and improvement of taxonomic knowledege
about Onosma. Although there are some taxa which
cannot be determined solely using the mentioned
characters, but these characters are useful tools in
distinguishing many taxa and solving some
complexities in Onosma. On the other hand the
combination of these characters along with
morphological, palynological and molecular evidence
provide a perfect view for taxonomy and careful
taxonomical revision of Onosma.
ACKNOWLEDGEMENT
We would like to thank Dr. Vitek head of Botany
Department of Natural History Museum of Wien and
Prof. Dr. Till head of Wien University Herbarium for
their support and help during our stay in Wien.
Moreover we are very grateful to Dr. Amini Rad and
Mr. Pahlevani from Iran Herbarium, Dr. Assadi and Dr.
Massoumi of Botany Division of the Research Institute
of Forests and Rangelands.
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