PHYLOGENY OF THE GENUS EREMURUS (ASPHODELACEAE) BASED ON
MORPHOLOGICAL CHARACTERS IN THE FLORA IRANICA AREA
K. Naderi Safar, S. Kazempour Osaloo & M. Zarrei
Received 22.11.2008. Accepted for the publication 15.04.2009
Naderi Safar, K., Kazempour Osaloo, S. & Zarrei, M. 2009: 06 30 Phylogeny of the genus Eremurus
(Asphodelaceae) based on morphological characters in the Flora Iranica area. -Iran. J. Bot. 15(1): 27-35. Tehran.
A phylogenetic analysis of Eremurus (Asphodelaceae) based on morphological data is presented. A total of twentyfive characters including 20 gross morphological, 4 palynological and 1 chromosomal characters were analyzed to
reconstruct phylogenetic relationships for 24 taxa of Eremurus and Trachyandra malosana plus Asphodelus
tenuifolius and two Asphodeline species as outgroups. Maximum parsimony approach as implemented in PAUP*
using heuristic search and branch swapping option of tree bisection-reconnection were used for phylogenetic
analyses. The analysis of characters with successive weighting using rescaled consistency index generated more
resolved and supported trees than the analysis with equally weighting characters. The analyses showed that T.
malosana is well allied with a clade of Eremurus species. Among Eremurus species analyzed, E. persicus and a
subclade of E. kopetdaghensis and E. luteus comprise the basal most branches as sisters to the remainder. Eremurus
subgenus Henningia with 15 taxa was paraphyletic; whereas, nine members of Eremurus subgenus Eremurus
formed a well supported monophyletic group for which E. furseorum of the former subgenus was sister taxon.
Sections Henningia (of subgenus Henningia) and Eremurus (of subgenus Eremurus) are also appeared to be nonmonophyletic while the section Ammolirion (of subgenus Eremurus) is monophyletic. Medium size pollen grain,
(25-50 µm) and chromosome number of 2n=14 are shared by Eremurus and Trachyandra. Three characters
including non-branched inflorescence, basifixed stamens and ellipsoidal pollen grains are synapomorphies for the
genus Eremurus solely. Tubular/campanulate flowers, incurved tepals and tepals with 3-5 nerves are
synapomorphies for the subgenus Eremurus. Similarly exerted stamen is a synapomorphy for the subgenus
Eremurus and E. forseorum (from subgenus Henningia). The tubular flowers (rather than campanulate) and the
thinnest pollen exine are putative synapomorphies for the section Ammolirion. The remaining characters were
evolved homoplasiously or ambiguously among these taxa.
Kosar Naderi Safar & Shahrokh Kazempour Osaloo (correspondence skosaloo@modares.ac.ir;
skosaloo@yahoo.com), Department of Plant Biology, Faculty of Basic Sciences, Tarbiat Modares University, P.O.
Box 14115-175, Tehran, Iran. −Mehdi Zarrei, Department of Biology, Faculty of Sciences, Shahed University of
Tehran, Po Box 18151-159, Tehran, Iran.
Key words. Asphodelaceae, Phylogeny, Morphology, Eremurus, Iran.
( ﺑﺮ اﺳﺎس ﺻﻔﺎت رﻳﺨﺖﺷﻨﺎﺳﻲ در ﻣﻨﻄﻘﻪ ﻓﻠﻮرا اﻳﺮاﻧﻴﻜﺎAsphodelaceae) ﻓﻴﻠﻮژﻧﻲ ﺟﻨﺲ ﺳﺮﻳﺶ
. داﻧﺸﺠﻮي ﻛﺎرﺷﻨﺎﺳﻲ ارﺷﺪ ﮔﺮوه زﻳﺴﺖﺷﻨﺎﺳﻲ ﮔﻴﺎﻫﻲ داﻧﺸﮕﺎه ﺗﺮﺑﻴﺖ ﻣﺪرس،ﻛﻮﺛﺮ ﻧﺎدري ﺻﻔﺎر
. اﺳﺘﺎدﻳﺎر ﮔﺮوه زﻳﺴﺖﺷﻨﺎﺳﻲ ﮔﻴﺎﻫﻲ داﻧﺸﮕﺎه ﺗﺮﺑﻴﺖ ﻣﺪرس،ﺷﺎﻫﺮخ ﻛﺎﻇﻢ ﭘﻮر اوﺻﺎﻟﻮ
. اﺳﺘﺎدﻳﺎر ﮔﺮوه زﻳﺴﺖﺷﻨﺎﺳﻲ داﻧﺸﮕﺎه ﺷﺎﻫﺪ ﺗﻬﺮان،ﻣﻬﺪي زارﻋﻲ
در ﻣﺠﻤﻮع.( ﺑﺮ اﺳﺎس داده ﻫﺎي ﺣﺎﺻﻞ از ﺻﻔﺎت رﻳﺨﺖﺷﻨﺎﺳﻲ ﻣﻨﻄﻘﻪ ﻓﻠﻮرا اﻳﺮاﻧﻴﻜﺎ اراﺋﻪ ﻣﻲﺷﻮدEremurus) آﻧﺎﻟﻴﺰ ﻓﻴﻠﻮژﻧﻲ ﺟﻨﺲ ﺳﺮﻳﺶ
( در ﻧﺎﺣﻴﻪ ﻓﻠﻮرا اﻳﺮاﻧﻴﻜﺎ و ﮔﻮﻧﻪEremurus) ﺗﺎﻛﺴﻮن از ﺟﻨﺲ ﺳﺮﻳﺶ24 ﺻﻔﺖ رﻳﺨﺖﺷﻨﺎﺳﻲ ﺑﺮاي ﺑﺎزﺳﺎزي رواﺑﻂ ﻓﻴﻠﻮژﻧﻲ25
ﻣﻮرد آﻧﺎﻟﻴﺰ ﻗﺮار، ﺑﻪ ﻋﻨﻮان ﺑﺮون ﮔﺮوهAsphodeline و دو ﮔﻮﻧﻪ از ﺟﻨﺲAsphodelus tenuifolius ﺑﻪ اﺿﺎﻓﻪTrachyandra malosana
( وheuristic) ﺑﺎ اﺳﺘﻔﺎده از ﺟﺴﺘﺠﻮي اﺑﺘﻜﺎريPAUP* ﺗﻌﺒﻴﻪ ﺷﺪه در ﻧﺮم اﻓﺰارMaximum parsimony آﻧﺎﻟﻴﺰ ﻓﻴﻠﻮژﻧﻲ ﺑﻪ روش.ﮔﺮﻓﺖ
IRAN. JOURN. BOT. 15 (1), 2009
Naderi Safar & al. 28
از آﻧﺎﻟﻴﺰ ﺻﻔﺎت ﺑﺎ وزندﻫﻲ ﻣﺘﻮاﻟﻲ ﺑﺎ اﺳﺘﻔﺎده از ﺷﺎﺧﺺ ﺳﺎزﮔﺎري.ﮔﺰﻳﻨﻪ ﺗﺒﺎدل ﺷﺎﺧﻪ دوﻧﻴﻤﻪ ﺷﺪن و اﺗﺼﺎل ﻣﺠﺪد درﺧﺖ ﺻﻮرت ﮔﺮﻓﺖ
آﻧﺎﻟﻴﺰ ﻧﺸﺎن داد ﻛﻪ.ﺗﺼﺤﻴﺢ ﺷﺪه درﺧﺘﺎﻧﻲ ﺑﺎ رواﺑﻂ ﻓﻴﻠﻮژﻧﻲ ﺑﻬﺘﺮ و ﺣﺪود اﻃﻤﻴﻨﺎن ﻛﻼدﻫﺎي ﺑﻴﺸﺘﺮ ﻧﺴﺒﺖ ﺑﻪ وزندﻫﻲ ﻳﻜﺴﺎن ﺻﻔﺎت ﺑﺪﺳﺖ آﻣﺪ
E. persicus ،Eremurus در ﻣﻴﺎن ﮔﻮﻧﻪﻫﺎي آﻧﺎﻟﻴﺰ ﺷﺪه از. ﻣﺘﺤﺪ ﺷﺪه اﺳﺖEremurus ﺑﻪ ﺧﻮﺑﻲ ﺑﺎ ﻛﻼد ﻣﺘﺸﻜﻞ از ﮔﻮﻧﻪﻫﺎيT. malosana
در. ﻫﺴﺘﻨﺪEremurus اوﻟﻴﻦ ﺷﺎﺧﻪﻫﺎ و ﮔﺮوه ﺧﻮاﻫﺮي ﺑﺮاي ﺳﺎﻳﺮ ﮔﻮﻧﻪﻫﺎيE. luteus وE. kopetdaghensis ﺑﻪ اﺿﺎﻓﻪ زﻳﺮ ﻛﻼدي ﺷﺎﻣﻞ
ﺑﻪ. ﺗﺎﻛﺴﻮن ﺗﻚ ﺗﺒﺎر ﻣﻲﺑﺎﺷﺪ9 ﺑﺎEremurus در ﺣﺎﻟﻲ ﻛﻪ زﻳﺮﺟﻨﺲ، ﺗﺎﻛﺴﻮن ﭘﻴﺮاﺗﺒﺎر اﺳﺖ15 ﺑﺎHenningia زﻳﺮﺟﻨﺲEremurus ﺟﻨﺲ
در ﺣﺎﻟﻲ ﻛﻪ.( ﻧﻴﺰ ﺗﻚﺗﺒﺎر ﻧﻴﺴﺘﻨﺪEremurus )از زﻳﺮﺟﻨﺲEremurus ( وHenningia )از زﻳﺮﺟﻨﺲHenningia ﻧﻈﺮ ﻣﻲرﺳﺪ ﺑﺨﺸﻪﻫﺎي
(25- 50 µm) ﺷﻴﺎرﻫﺎي ﮔﺮده ﺑﺎ اﻧﺪازه ﻣﺘﻮﺳﻂ، ﺻﻔﺖ آﻧﺎﻟﻴﺰ ﺷﺪه25 از ﻣﻴﺎن.( ﺗﻚ ﺗﺒﺎر اﺳﺖEremurus )از زﻳﺮﺟﻨﺲAmmolirion ﺑﺨﺸﻪ
ﭘﺮﭼﻢ ﻫﺎي، ﺳﻪ ﺻﻔﺖ ﺷﺎﻣﻞ ﮔﻞ آذﻳﻦ ﻏﻴﺮ ﻣﻨﺸﻌﺐ. ﻣﺸﺘﺮكاﻧﺪTrachyandra وEremurus ﺑﺮاي ﺟﻨﺲﻫﺎي2n= 14 و ﻋﺪد ﻛﺮوﻣﻮزوﻣﻲ
ﮔﻠﭙﻮشﻫﺎي ﺧﻤﻴﺪه ﺑﻪ ﺳﻤﺖ،اﺳﺘﻜﺎﻧﻲ/ ﮔﻞ ﻫﺎي ﻟﻮﻟﻪاي.ﭘﺎﻳﻪ ﭼﺴﺐ و ﺷﻴﺎر ﻫﺎي ﮔﺮده اي ﺑﻴﻀﻮي ﺳﻴﻨﺎﭘﻮﻣﻮرﻓﻲﻫﺎي وﻳﮋه ﺟﻨﺲ ﺳﺮﻳﺶ ﻫﺴﺘﻨﺪ
ﻫﻤﭽﻨﻴﻦ ﭘﺮﭼﻢﻫﺎي ﺑﻴﺮون زده ﻣﺨﺘﺺ زﻳﺮ ﺟﻨﺲ. ﻫﺴﺘﻨﺪEremurus رﮔﻪ ﺑﻮدن ﮔﻠﭙﻮشﻫﺎ ﺳﻴﻨﺎﭘﻮﻣﻮرﻓﻲﻫﺎي زﻳﺮ ﺟﻨﺲ5 ﺗﺎ3 داﺧﻞ و
ﮔﻞﻫﺎي ﻟﻮﻟﻪاي )ﻧﻪ اﺳﺘﻜﺎﻧﻲ( و ﻧﺎزﻛﺘﺮﻳﻦ اﮔﺰﻳﻦ ﮔﺮدهاي ﺻﻔﺎت ﻣﺸﺘﺮك.( اﺳﺖHenningia )از زﻳﺮﺟﻨﺲE. forseorum وEremurus
. ﺑﻘﻴﻪ ﺻﻔﺎت ﺑﻪ ﺻﻮرت ﻫﻢﻧﻤﺎ ﻳﺎ ﻣﺒﻬﻢ در ﻣﻴﺎن اﻳﻦ ﺗﺎﻛﺴﻮنﻫﺎ ﺗﻜﺎﻣﻞ ﻳﺎﻓﺘﻪاﻧﺪ. ﻣﻲ ﺑﺎﺷﻨﺪAmmolirion ﺑﺮاي اﻋﻀﺎي ﺑﺨﺸﻪ
INTRODUCTION
Eremurus M. Bieb. (Asphodelaceae) with 45 species
worldwide (Mabberley 1990) is distributed over large
area in Central Asia, Caucasia, Afghanistan, Iran,
Pakistan, Iraq, Turkey, Lebanon, India and China
(Wendelbo & Furse 1969). Its centre of diversity is in
Central Asia (Hedge & Wendelbo 1963). In the Flora
Iranica area, Asphodeloideae as a subfamily of
Liliaceae has three genera including Eremurus,
Asphodelus L. and Asphodeline Reichenb (Wendelbo
1982). Seven species of Eremurus have been recorded
in Iran, and 24 species described for the Flora Iranica
area. Among them only Eremurus kopetdaghensis M.
Pop. ex B. Fedtsch. is sub-endemic to Iran. It is also
distributed in Turkmenistan (Wendelbo 1982).
Angiosperm Phylogeny Group (APG 1998) determined
Asphodelaceae as a family of Asparagles.
Xanthorrhoeaceae sensu lato as a large unit
circumscribed by APG II (APG 2003) includes
Asphodelaceae,
Hemerocallidaceae
and
Xanthorrhoeaceae s. str.
Eremurus is well known for its large colourful
racemes. It is distinguished from Asphodelus with not
branched inflorescence and from Asphodeline by
lacking membranous leaf sheaths at the base of
inflorescence. Eremurus has been divided into two
subgenera and three sections (Wendelbo 1982). The
subgenus Eremurus is characterized by light browngreen or cream tubular/campanulate flowers, incurved
tepals and tepals with 3−5 nerves abaxially and exerted
filaments. Whereas, the subgenus Henningia has white,
pink or yellow rotate flowers, mostly non-exerted
filaments and tepals one nerved abaxially.
Apart from several studies conducted by Wendelbo
(1962, 1964, 1968, 1982, 1985) and Wendelbo & Furse
(1969) which are mainly concentrated on Central Asia,
there is no intensive contribution to this genus in Iran.
In addition to this fact that most species are of great
horticultural potential, this genus has several other
commercial usages such as, extracting natural glue
from rhizomes, edible young leaves as vegetable and
very beautiful ornamental flowers (Kamenetsky &
Rabinowitch 1999).
The goals of the present study are: (1) to evaluate
phylogenetic status of the Eremurus and its
infrageneric subdivisions including subgenera and
sections and (2) to examine evolutionary trend of
morphological characters in the context of the
hypothesized phylogeny.
MATERIALS AND METHODS
Twenty-four taxa representing three currently
recognized sections belonging to the two subgenera of
Eremurus and one species of Trachyandra (T.
malosana (Baker) Oberm.), as the closest genus to
Eremurus (Chase & al. 2000), were included in the
analysis. Asphodelus tenuifolius and the two
Asphodeline species were selected as outgroups.
Characters used in the phylogenetic analysis were
obtained through examination of fresh materials in the
field and herbarium specimens deposited at four major
herbaria of Iran (TARI, IRAN, TUH & FUMH, see
Holmgren & Holmgren 1998 for herbarium acronyms)
29 Cladistic analysis of Eremurus in Iran
IRAN. JOURN. BOT. 15 (1), 2009
Table 1. List of morphological characters of Eremurus species used in phylogenetic analyses.
1. Height of plant: ≤ 70 cm (0) > 70cm (1)
Scape:
2. Indumentum: Pubescent (0) Glabrous (1)
Bract:
3. Shape: Subulate (0) Lanceolate (1) Ovoid (2)
4. Margin: Glabrous (0) Ciliate (1)
5. Surface: Glabrous (0) Hairy (1)
Leaves indumentum:
6. Margin: Glabrous (0) Ciliate (1)
7. Surface: Glabrous (0) Hairy (1)
Inflorescence:
8. Branched (0) Non branched (1)
9. Length: ≤ 30 cm (0) >30 cm (1)
10. No. of flowers: ≤ 70 (0) >70 (1)
Flower:
11. Flower shape: Subrotate (0) Tubular (1) Campanulate (2)
12. Tepal color: White-cream (0) Pink (1) Yellow (2) Brown (3)
13. Tepal length: ≤ 15 mm (0) >15 mm (1)
14. Tepal width: Equal (0) Unequal (1)
15. Tepal tip: Erect or recurved (0) Incurved (1)
16. Tepal nerve: 1 (0) 3-5 (1)
Stamen:
17. Anther type: Dorsifixed (0) Basifixed (1)
18. Stamen position: Non exerted (0) Exerted (1)
Fruit:
19. Shape: Globose (0) Ellipsoidal (1) Pyriform (2)
20. Surface sculpturing: Present (0) Absent (1)
Pollen grain:
21. Shape: Oblate-spheroidal (0) Ellipsoidal (1)
22. Size: Large (50-100 µm) (0) Medium (25-50µm) (1)
Exine:
23. Thickness: Thick, > 1.5 µm (0) Thin, ≤ 1.5 µm (1)
24. Surface sculpturing: Rugulate-perforate (0) Microreticulate (1) Distinct reticulate (2)
Chromosome number:
25. 2n=28 (0) 2n=14(1)
or adopted from appropriate references (Wendelbo
1982; Matthews 1986; Fedchenco 1968; Kosenko &
Sventorzhetskaya 1999; Kativu 2001).
Twenty-five informative characters with relevant
character states used in the present analysis were given
in Table 1. The polarity of characters was determined
using the outgroup method (Maddison & al. 1984).
Phylogenetic analyses were performed on the data
matrix (Table 2) using maximum parsimony method
(MP) as implemented in PAUP* version 4.0b10
(Swofford 2002) installed in a Macintosh computer. All
characters were considered as equally weighted. The
heuristic search option was selected using 100
replications of random addition sequence with
ACCTRAN optimization and TBR (tree bisection
reconnection) branch-swapping with MulTrees on and
steepest descent off. Analyses were then conducted
using a successive re-weighting strategy (Farris 1969)
in order to improving the trees indices and decreasing
the effect of characters showing high homoplasy on
tree topologies. Weights were assigned to characters
using the re-weight characters option based on the
rescaled consistency index (RC) (Farris 1989) with a
base weight of one. When the tree length and
consistency index (CI), retention index (RI) and RC
remained unchanged in successive rounds, these trees
were accepted as the successive re-weighting trees. In
both analyses, supports for clades were evaluated by
bootstrapping (Felsenstein 1985) using 20000
replications with the heuristic search option, simple
addition sequence, TBR branch swapping and
MulTrees off.
Character evolution was surveyed on the strict
consensus tree resulting from successive weighting of
IRAN. JOURN. BOT. 15 (1), 2009
Naderi Safar & al. 30
Table 2. Data matrix used in the phylogenetic analyses of Eremurus and its related genera. Missing data are coded
as “?” a= {01}.
________________________________________________________________
Taxon
characters
_______________________________________________________________
Asphodelus tenuifolious
0010010000100000000000010
Asphodeline szovitsii
0000010000001100000?00000
A. dendroides
0100010000000?00000100000
Eremurus inderiensis
0001111101100011110111111
E.cappadocicus
00010001??130011110111111
E.comosus
10110111?0110011110111111
E.dolichomischus
10110111?1130011110111111
E.spectabilis subsp.
spectabilis
1101000111200111110011021
E.spectabilis subsp.
subalbiflorus
1101011111200011111011021
E.korshinskii
1101000100231011110111021
E.soogdianus
100101a1?0200111110011021
E.fuscus
11010001?1220011110111021
E.kopetdaghensis
0111010100011100101111021
E.persicus
0021111100011100100111021
E.stenophyllus subsp.
stenophyllus
1100010111020100100111021
E.stenophyllus subsp.
aurantiacus
1001011111020100100111021
E.olgae
1100000110011100100111021
E.luteus
0101010100021100101011021
E.kaufmanii
10210111?1001000100111021
E.himaliacus
11010101?1001000100111021
E.aitchisonii
11010101?0011000100111021
E.bucharicus
1001010110000100100011021
E.bacterianus
10011111??001100100111021
E.afghanicus
10011111?1000000100111021
E.suworowiia
0010101?0020000100111021
E.roseolus
10010111?1011100102111021
E.furseorum
10001101?1010000110111021
Trachyandra malosana
0020011001000000000001011
characters with ACCTRAN optimization and
interpreting polytomies as multiple speciation events
(hard polytomies) using MacClade version 4.3
(Maddison & Maddison 2004).
RESULTS
The first phylogenetic analysis using equally weighted
characters generated 423 most parsimonious trees of 86
steps with CI= 0.360 and RI= 0.656. The strict
consensus tree of the analysis was not shown. The
relationships among Eremurus species remain
unresolved in this tree. The second analysis based on
re-weighted characters by rescaled consistency index
yielded 60 well resolved and supported trees with
length= 20.2 steps, CI= 0.694, RI= 0.901. The strict
consensus tree of the analysis is shown in Fig. 1. In
both analyses, Trachyandra malosana is closest taxa to
Eremurus. Among Eremurus species analyzed,
Eremurus persicus (Jaub. & Spach) Boiss. and a
subclade of E. kopetdaghensis and E. luteus Baker, as
unresolved branches, comprised the basal most taxa.
The remaining species formed a large polytomy of
several individual branches and two weakly subclades
(Fig. 1.).
DISCUSSION
Monophyly and infrageneric relationships
of Eremurus
The present analysis based on morphological characters
indicates that Eremurus is a monophyletic taxon
31 Cladistic analysis of Eremurus in Iran
IRAN. JOURN. BOT. 15 (1), 2009
Fig. 1. Strict consensus of the 60 most parsimonious trees obtained from morphological characters after successive
weighting by rescaled consistency index. Numbers above branches are bootstrap values. Numbers <50% were not
indicated.
(bootstrap=99%). Our molecular analyses based upon
both nrDNA ITS and chloroplast trnL-F sequences is
also consistent with this finding (Naderi & Kazempour
Osaloo 2008, and Naderi & al. unpubl. data).
According to latest treatment of Wendelbo (1982),
Eremurus has been divided into two subgenera,
Eremurus and Henningia. The former subgenus is
composed of sections Eremurus and Ammolirion, and
the latter one includes only section Henningia.
Phylogenetic analyses of the morphological data
suggested that the subgenus Eremurus is well
supported monophyletic group (bootstrap=91%). The
subgenus is characterized by three synapomorphies
including tubular/campanulate flowers, incurved tepals
and tepals with 3−5 abaxial nerves. The section
Eremurus is not monophyletic while the section
Ammolirion is monophyletic (bootstrap=81%). This
section is distinguished from the section Eremurus by
having tubular flowers rather than campanulate flowers
and by the thinnest pollen exine (Wendelbo 1982;
Kosenko & Sventorzhetskaya 1999). The subgenus
Henningia has white, pink or yellow subrotate flowers,
non-exerted filaments and erect or recurved tepals with
one abaxial nerve. The present phylogeny shows that
these characters are primitive (symplesiomorphies),
since they are shared by Trachyandra, the closest
relative of Eremurus (Chase & al. 2000; Devey & al.
2006) and outgroup taxa (see Figs. 2B & 3A).
Therefore, the subgenus Henningia (and section
Henningia) is paraphyletic. Eremurus furseorum
Wendelbo is the only species of Henningia that allied
with Eremurus subgenus Eremurus. They do share
exerted stamens.
IRAN. JOURN. BOT. 15 (1), 2009
Naderi Safar & al. 32
Evolutionary trend of characters
REFERENCES
MacClade reconstruction of character evolution
indicated that all 25 characters but characters no. 14
and 20 were evolved unambiguously. Characters 22
(medium-sized pollen grain, 25-50 µm) and 25
(Chromosome number of 2n=14) are shared by
Eremurus and Trachyandra. The three characters
including non-branched inflorescence (character 8),
basifixed anthers (character 17) and ellipsoidal pollen
grains (character 21) are synapomorphies for the genus
Eremurus solely (Fig. 2A). As noted at the previous
part, tubular/campanulate flowers (character 11)
incurved tepals (character 15) and tepals with 3−5
abaxial nerves (character 16) are synapomorphy for the
subgenus Eremurus (Figs. 2B & 3A). Similarly,
character 18 (exerted stamens) is a synapomorphy for
the subgenus Eremurus and E. forseorum (from
subgenus Henningia). The tubular flowers (character
11) and the thinnest pollen exine (character 23) are
putative synapomorphies for the section Ammolirion
(Fig. 3B).
Characters 1, 4, 9, 10, 13 and 24 were undergone
reversal evolution among Eremurus species. For
example, the plesiomorphic state for characters 10 and
24 are number of flowers < 70 and pollen exine with
mircoreticulate sculpturing, respectively. The number
of flowers > 70 and pollen exine with distinct reticulate
sculpturing are derived (Figs. 4A & 4B). Likewise,
characters 2, 3, 5, 6, 7, 12 and 19 were evolved in
parallel evolution among Eremurus species and
Trachyandra malosana. For instance, evolutionary
trend for characters 3, having three character sates, was
occurred from subulate bract as a plesiomorphic
condition to both lanceolate (in Eremurus comosus O.
Fedtsch., E. dolichomischus Vved. & Wendelbo and E.
kopetdaghensis) and ovate (in E. kaufmanii Regel, E.
persicus and T. malosana). Similar results were found
for the evolution of character no. 19 (fruit shape). The
analysis indicated that globose fruit is plesiomorphic
and fruits with ellipsoidal and pyriform shape are
derived. According to this study, ellipsoidal fruits that
are found in both E. kopetdaghensis and E. luteus as
well as in the far related E. spectabilis subsp.
subalbiflorus evolved in parallel, and pyriform fruit is
unique to E. roseolus Vved. (Figs. 5A & 5B).
Chase, M. W., Bruijn, A.Y., Cox, A. V., Reeves, G. P.,
Rudall, J., Johnson, M. A. T. & Eguiarte, L. E.
2000:
Phylogenetics
of
Asphodelaceae
(Asparagales): An Analysis of Plastid rbcL and
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374-385.
Farris, J. S. 1989: The Retention index and the rescaled
consistency index.− Cladistics 5: 417-419.
Fedchenco, B. A. 1968: Eremurus. In. Flora of the
U.S.S.R, vol. IV: 27- 40.-Jerusalem. (English
translation).
Felsenstein, J. 1985: Confidence limits on phylogenies:
An approach using the bootstrap. -Evolution.
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Hedge, I. & Wendelbo, P. 1963: Notes on the giant
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Holmgren, P. K. & Holmgren, N. H. 1998:
[continuously updated]. Index Herbariorum: A
global directory of public herbaria and associated
staff. −New York Botanical Garden's Virtual
Herbarium. http://sweetgum.nybg.org/ih/
Kamenetsky, K. & Rabinowitch, E. 1999: Flowering
response of Eremurus to post- harvest temperature.
- Scientia Horticulturae 79: 75-86
Kativu, S. 2001: Trachyandra. In. Flora Zambesiaca
vol. 12, part3. -Kew (http://epic.kew.org).
Kosenko, V & Sventorzhetskaya, O. 1999. Pollen
Morphology in the family Asphodelaceae
(Asphodeleae, Kniphofieae). – Grana 38: 218-227.
Mabberley, D. J. 1990: The Plant Book. -Cambridge
University Press, Cambridge.
Maddison, W. P., Donoghue, M. J. & Maddison, D. R.
1984: Outgroup analysis and parsimony. -Syst.
Zool. 33:83-103.
Maddison, D. R. & Maddison, W. P. 2004: MacClade4:
Analysis of phylogeny and character evolution.
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Massachusetts.
Matthews, W. A. 1986: Eremurus. In. Flora of Turkey
and the East Aegean Island vol. 8: 86-87. Edinburgh.
Naderi, K. & Kazempour Osaloo, S. 2008: Molecular
phylogeny of the genus Eremurus (Asphodelaceae)
based on cpDNA trnL-F sequences. -Proceedings of
15th National and 3rd International Conference of
Biology. p. 37.-Tehran.
ACKNOWLEDGEMENTS
This research was supported by a research fund of
Tarbiat Modares University. This work represents
partial fulfillment of the requirement for obtaining M.
Sc. degree by K. Naderi from Tarbiat Modares
University.
33 Cladistic analysis of Eremurus in Iran
IRAN. JOURN. BOT. 15 (1), 2009
A
Character 8: Inflorescence status
0 branched
1 non-branched
Character 1 7: Stamens
position
0 Dorsifixed
1 Basifixed
B
Asphodelus tenuifolious
Asphodeline szovitsii
Ashodeline dendroides
Eremurus inderiensis
E. cappadocicus
E. comosus
E. dolichomischus
E.korshinskii
E. fuscus
E. spectabilis subsp. spectabilis
E. soogdianus
E. spectabilis subsp. subalbiflorus
E. furseorum
E. stenophyllus subsp. stenophyllus
E. stenophyllus subsp. aurantiacus
E. suworowii
E. olgae
E. kaufmanii
E. himaliacus
E. aitchisonii
E. bucharicus
E. bacterianus
Character 11:Flower shape
E. afghanicus
E. roseolus
0 Subrotate
E. kopetdaghensis
E. luteus
E. persicus
T. malosana
Character 21: Pollen grain
Asphodelus tenuifolious
Asphodeline szovitsii
Ashodeline dendroides
Eremurus inderiensis
E. cappadocicus
E. comosus
E. dolichomischus
E.korshinskii
E. fuscus
E. spectabilis subsp. spectabilis
E. soogdianus
E. spectabilis subsp. subalbiflorus
E. furseorum
E. stenophyllus subsp. stenophyllus
E. stenophyllus subsp. aurantiacus
E. suworowii
E. olgae
E. kaufmanii
E. himaliacus
E. aitchisonii
E. bucharicus
E. bacterianus
E. afghanicus
E. roseolus
E. kopetdaghensis
E. luteus
E. persicus
T. malosana
1 Tubular
2 Campanulate
0 Oblate-spheroidal
1 Ellipsoidal
Fig. 2. MacClade reconstruction of character evolution of: A (characters 8, 17, 21); B (character 11)
on the tree.
A
Character 15: Number of
tepal nerve
0 (1)
1 (3-5)
Character 16: Tepal Tip
0 Erect or recarved
1 Incurved
B
Asphodelus tenuifolious
Asphodeline szovitsii
Ashodeline dendroides
Eremurus inderiensis
E. cappadocicus
E. comosus
E. dolichomischus
E.korshinskii
E. fuscus
E. spectabilis subsp. spectabilis
E. soogdianus
E. spectabilis subsp. subalbiflorus
E. furseorum
E. stenophyllus subsp. stenophyllus
E. stenophyllus subsp. aurantiacus
E. suworowii
E. olgae
E. kaufmanii
E. himaliacus
E. aitchisonii
E. bucharicus
E. bacterianus
E. afghanicus
Character 23: Pollen exin
E. roseolus
thickness
E. kopetdaghensis
E. luteus
E. persicus
T. malosana
0 > 1.5 µm
1 ≤ 1.5 (1)
Asphodelus tenuifolious
Asphodeline szovitsii
Ashodeline dendroides
Eremurus inderiensis
E. cappadocicus
E. comosus
E. dolichomischus
E.korshinskii
E. fuscus
E. spectabilis subsp. spectabilis
E. soogdianus
E. spectabilis subsp. subalbiflorus
E. furseorum
E. stenophyllus subsp. stenophyllus
E. stenophyllus subsp. aurantiacus
E. suworowii
E. olgae
E. kaufmanii
E. himaliacus
E. aitchisonii
E. bucharicus
E. bacterianus
E. afghanicus
E. roseolus
E. kopetdaghensis
E. luteus
E. persicus
T. malosana
Fig. 3. MacClade reconstruction of character evolution of: A (characters 15, 16); B (character 23) on the tree.
IRAN. JOURN. BOT. 15 (1), 2009
A
Character 10: Number
of flowers
0 ≤ 70
Asphodelus tenuifolious
Asphodeline szovitsii
Ashodeline dendroides
Eremurus inderiensis
E. cappadocicus
E. comosus
E. dolichomischus
E.korshinskii
E. fuscus
E. spectabilis subsp. spectabilis
E. soogdianus
E. spectabilis subsp. subalbiflorus
E. furseorum
E. stenophyllus subsp. stenophyllus
E. stenophyllus subsp. aurantiacus
E. suworowii
E. olgae
E. kaufmanii
E. himaliacus
E. aitchisonii
E. bucharicus
Character 24: Exine Surface
E. bacterianus
ornamentation
E. afghanicus
E. roseolus
0 Rugulate-Perforate
E. kopetdaghensis
E. luteus
E. persicus
T. malosana
1 >70
Naderi Safar & al. 34
1 Microreticulate
2 Reticulate
B
Asphodelus tenuifolious
Asphodeline szovitsii
Ashodeline dendroides
Eremurus inderiensis
E. cappadocicus
E. comosus
E. dolichomischus
E.korshinskii
E. fuscus
E. spectabilis subsp. spectabilis
E. soogdianus
E. spectabilis subsp. subalbiflorus
E. furseorum
E. stenophyllus subsp. stenophyllus
E. stenophyllus subsp. aurantiacus
E. suworowii
E. olgae
E. kaufmanii
E. himaliacus
E. aitchisonii
E. bucharicus
E. bacterianus
E. afghanicus
E. roseolus
E. kopetdaghensis
E. luteus
E. persicus
T. malosana
Fig. 4. MacClade reconstruction of character evolution of: A (character 10); B (character 24) on the tree.
A
Character 3: Bracts Shape
0 Subulate
1 Lanceolate
2 Ovoid
Asphodelus tenuifolious
Asphodeline szovitsii
Ashodeline dendroides
Eremurus inderiensis
E. cappadocicus
E. comosus
E. dolichomischus
E.korshinskii
E. fuscus
E. spectabilis subsp. spectabilis
E. soogdianus
E. spectabilis subsp. subalbiflorus
E. furseorum
E. stenophyllus subsp. stenophyllus
E. stenophyllus subsp. aurantiacus
E. suworowii
E. olgae
E. kaufmanii
E. himaliacus
E. aitchisonii
E. bucharicus
Character 19: Fruit Shape
E. bacterianus
E. afghanicus
0 Globose
E. roseolus
E. kopetdaghensis
E. luteus
E. persicus
T. malosana
1 Ellipsoidal
2 Pyriform
B
Asphodelus tenuifolious
Asphodeline szovitsii
Ashodeline dendroides
Eremurus inderiensis
E. cappadocicus
E. comosus
E. dolichomischus
E.korshinskii
E. fuscus
E. spectabilis subsp. spectabilis
E. soogdianus
E. spectabilis subsp. subalbiflorus
E. furseorum
E. stenophyllus subsp. stenophyllus
E. stenophyllus subsp. aurantiacus
E. suworowii
E. olgae
E. kaufmanii
E. himaliacus
E. aitchisonii
E. bucharicus
E. bacterianus
E. afghanicus
E. roseolus
E. kopetdaghensis
E. luteus
E. persicus
T. malosana
Fig. 5 MacClade reconstruction of character evolution of: A (character 3); B (character 19) on the tree.
35 Cladistic analysis of Eremurus in Iran
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