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MOLECULAR PHYLOGENY of the TRIBE HEDYSAREAE with SPECIAL REFERENCE to ONOBRYCHIS (FABACEAE) AS INFERRED from Nrdna ITS SEQUENCES S

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MOLECULAR PHYLOGENY of the TRIBE HEDYSAREAE with SPECIAL REFERENCE to ONOBRYCHIS (FABACEAE) AS INFERRED from Nrdna ITS SEQUENCES S MOLECULAR PHYLOGENY OF THE TRIBE HEDYSAREAE WITH SPECIAL REFERENCE TO ONOBRYCHIS (FABACEAE) AS INFERRED FROM nrDNA ITS SEQUENCES S. Ahangarian, Sh. Kazempour Osaloo & A. A. Maassoumi Ahangarian, S., Kazempour Osaloo, Sh. & Maassoumi, A. A. 2007 12 31:. Molecular phylogeny of the tribe Hedysareae with special reference to Onobrychis (Fabaceae) as inferred from nrDNA ITS sequences. -Iran. J. Bot. 13 (2): 64-74. Tehran. A total of 53 species (54 sequence accessions) representing 28 species of the tribe Hedysareae, 5 species of Vicioid clade and 19 species of Galegeae including the two Glycyrrhiza species as outgroups was included in a phylogenetic analysis using nrDNA ITS sequences with maximum parsimony method. The analysis, based on successive reweighting by rescaled consistency index, revealed that Vicioid clade is the first branch followed by Chesneya-Caragana clade and Astragalean clade as successive sisters to Hedysaroid clade. The genus Alhagi is well allied, as the basal most taxon, with the remainder of Hedysaroid clade. Ebenus and Taverniera are each monophyletic and weakly allied taxa. Hedysarum is not monophyletic, its species are dispersed across Hedysaroid clade. Sulla a segregate of Hedysarum, forms a well supported monophyletic group with the inclusion of H. humile. The enigmatic genus Eversmannia is allied with Onobrychis clade. Onobrychis itself is not monophyletic due to inclusion of the two Hedysarum species, and Onobrychis subgenus Onobrychis is not monophyletic as well, whereas Onobrychis subgenus Sisyrosema forms a strongly supported clade. In contrast to Onobrychis section Heliobrychis, sections Dendrobrychis and Onobrychis appear not to be each monophyletic. Hedysareae is circumscribed herein to include only Alhagi, Ebenus, Eversmannia, Hedysarum, Onobrychis, Sulla, Taverniera, Coretrodendron and Sartoria. Somayeh Ahangarian & Shahrokh Kazempour Osaloo (correspondence), Department of Plant Biology, Faculty of Basic Sciences, Tarbiat Modares University, Tehran 14115-175, Iran. –Ali Asghar Maassoumi, Research Institute of Forests and Rangelands, P. O. Box 13185-116, Tehran, Iran. Submitted: 2007. 10. 15 Accepted for publication on: 2007. 12. 19 Keywords. Fabaceae, Hedysareae, Molecular phylogeny, nrDNA ITS, Onobrychis, Iran. ﻓﻴﻠ nrDNA ITS Onobrychis Hedysareae ﻋﻠﻲ ﺗﻌﺪاد 53 ﮔﻮﻧﻪ (54 ﺗﻮاﻟﯽ) ﺷﺎﻣﻞ 28 ﮔﻮﻧﻪ از ﻗﺒﯿﻠﻪ Hedysareae، 5 ﮔﻮﻧﻪ از ﮐﻼد Vicioid و 19 ﮔﻮﻧﻪ از ﻗﺒﯿﻠﻪ Galegeae از ﺟﻤﻠﻪ دو ﮔﻮﻧﻪ Glycyrrhiza ﺑﻪ ﻋﻨﻮان ﺑﺮونﮔﺮوه ﺑﺎ اﺳﺘﻔﺎده از ﺗﻮاﻟﯽﻫﺎي nrDNA ITS در آﻧﺎﻟﯿﺰ ﻓﯿﻠﻮژﻧﯽ ﺑﺎ روش ﺑﯿﺸﯿﻨﻪ ﺻﺮﻓﻪﺟﻮﯾﯽ ﻗﺮار ﮔﺮﻓﺖ. آﻧﺎﻟﯿﺰ ﺑﺮاﺳﺎس وزندﻫﯽ ﻣﺠﺪد ﭘﯽدرﭘﯽ ﺑﺎ ﺷﺎﺧﺺ ﺳﺎزﮔﺎري ﺗﺼﺤﯿﺢ ﺷﺪه ﻓﺎش ﻧﻤﻮد ﮐﻪ ﮐﻼد Vicioid اوﻟﯿﻦ ﺷﺎﺧﻪاي اﺳﺖ ﮐﻪ ﺑﺪﻧﺒﺎل آن ﮐﻼد Chesneya-Caragana و ﮐﻼد Astragalean ﺑﻪ ﻋﻨﻮان ﮔﺮوهﻫﺎي ﺧﻮاﻫﺮي ﮐﻼد Hedysaroid ﻗﺮار دارﻧﺪ. ﺟﻨﺲ Alhagi ﺑﻪﻋﻨﻮان ﭘﺎﯾﻪﺗﺮﯾﻦ ﺗﺎﮐﺴﻮن ﺑﺎ ﺑﻘﯿﻪ ﮐﻼد Hedysaroid ﺑﻪ ﺧﻮﺑﯽ ﻣﺘﺤﺪ ﻣﯽﺷﻮد. ﻫﺮﮐﺪام از ﺟﻨﺲﻫﺎي Ebenus و Taverniera ﺗﮏﺗﺒﺎرﻧﺪ و ﺑﻪ ﻃﻮر ﺿﻌﯿﻒ ﺑﺎ ﻫﻢ ﻣﺘﺤﺪ ﻫﺴﺘﻨﺪ. ﺟﻨﺲ Hedysarum ﺗﮏﺗﺒﺎر ﻧﯿﺴﺖ و ﮔﻮﻧﻪﻫﺎﯾﺶ در ﺳﺮاﺳﺮ ﮐﻼد Hedysaroid ﭘﺨﺶ ﻫﺴﺘﻨﺪ. Sulla ﯾﮏ ﺟﻨﺲ ﺟﺪاﺷﺪه از Hedysarum، ﺑﺎ در ﺑﺮﮔﺮﻓﺘﻦ ﮔﻮﻧﻪ Hedysarum humile ﮔﺮوه ﺗﮏﺗﺒﺎر ﺑﻪ ﺧﻮﺑﯽ ﺣﻤﺎﯾﺖ ﺷﺪهاي را ﺗﺸﮑﯿﻞ ﻣﯽدﻫﺪ. ﺟﻨﺲ Eversmannia ﺑﺎ ﮐﻼد Onobrychis ﺧﻮﯾﺸﺎوﻧﺪ اﺳﺖ. ﺧﻮد ﺟﻨﺲ Onobrychis ﺑﻪ ﻋﻠﺖ ﻗﺮار ﮔﺮﻓﺘﻦ دو ﮔﻮﻧﻪ Hedysarum در ﻣﯿﺎن ﮔﻮﻧﻪﻫﺎﯾﺶ ﺗﮏﺗﺒﺎر ﻧﯿﺴﺖ و زﯾﺮﺟﻨﺲ Onobrychis از ﺟﻨﺲ Onobrychis ﻧﯿﺰ ﺗﮏﺗﺒﺎر ﻧﯿﺴﺖ، در ﺣﺎﻟﯿﮑﻪ زﯾﺮﺟﻨﺲ Sisyrosema ﯾﮏ ﮐﻼد ﻗﻮﯾﺎً ﺣﻤﺎﯾﺖ ﺷﺪهاي را ﻣﯽﺳﺎزد. در ﻣﯿﺎن ﺑﺨﺸﻪﻫﺎي ﭼﻨﺪ ﮔﻮﻧﻪاي آﻧﺎﻟﯿﺰ ﺷﺪه از ﺟﻨﺲ Onobrychis، ﺑﺮﺧﻼف ﺑﺨﺸﻪ Heliobrychis، ﺑﺨﺸﻪﻫﺎي Dendrobrychis و Onobrychis ﻇﺎﻫﺮاً ﺗﮏﺗﺒﺎر ﻧﯿﺴﺘﻨﺪ. در اﯾﻨﺠﺎ ﻗﺒﯿﻠﻪ Hedysareac ﻓﻘﻂ ﻣﺸﺘﻤﻞ ﺑﺮ ﺟﻨﺲﻫﺎي ,Alhagi, Ebenus, Eversmannia Hedysarum, Onobrychis, Sulla, Taverniera, Corethrodendron وSartoria ﺗﻌﯿﯿﻦ ﺣﺪود ﻣﯽﺷﻮد. 65 Phylogeny of Hedysareae IRAN. JOURN. BOT. 13 (2), 2007 INTRODUCTION 1981a), 5 species of the Vicioid clade (Sanderson and Since De Candolle (1825) onward, the taxonomic Wojciechowski 1996) and 19 species of the tribe delimitation of the tribe Hedysareae has been Galegeae including two Glycyrrhiza as outgroups undergone several treatments by various authors based on previous studies (Sanderson and (Bentham 1865; Hutchinson 1964; Polhill 1981a, 1994; Wojciechowski 1996; Hu et al. 2000; Wojciechowski et Choi and Ohashi 2003; Lock 2005). Just recently, Lock al. 2000, 2004; Wojciechowski 2003, 2005). The (2005) expanded the tribe to twelve genera including complete nrDNA ITS (ITS1, 5.8S and ITS2) region for Alhagi Adans., Calophaca Fisch ex DC., Caragana 19 taxa including Alhagi pseudalhagi (M. B.) Desv., Fabr. and Halimodendron Fisch ex DC., formerly Onobrychis (10 species of eight out of nine sections), treated in Galegeae (Polhill 1981b, 1994) in addition to Hedysarum (2 species), Taverniera (2 species), Ebenus Corethrodendron Basin., Ebenus L., Eversmannia (3 species), Eversmannia subspinosa Fisch., were Bunge, Hedysarum L., Onobrychis Mill., Sartoria determined first time by us. In the case of Boiss. & Heldr., Sulla Medik. and Taverniera DC. Halimodendron halodendron (Pall.) Voss, for which Lock (2005) briefly overviewed the taxonomic history ITS1+ITS2 were previously sequenced by Sanderson of some of controversial genera (e. g. Hedysarum, and Wojciechowski (1996), nrDNA ITS sequences Corethrodendron, Sulla and Taverniera ), and thus we do not repeat these again here. The tribe is a member of were also determined here again. For this and the remainder taxa, the sequences were obtained from gene Polhill’s temperate herbaceous group and has been traditionally placed close to the tribe Galegeae. bank (see Table 1). Available molecular data (nrDNA ITS and matK DNA Extraction. Leaf materials were sampled from sequences) showed that the Hedysaroid clade herbarium specimens deposited in the Herbarium of (composed of Hedysarum boreale Nutt., Onobrychis Research Institute of Forests and Rangelands (TARI), montana DC., Alhagi camelorum Fisch. and A. the Central Herbarium of University of Tehran (TUH), maurorum Medik.), is one of the three well-supported herbarium of Hacettepe University (HUB), and subclades, within the Inverted Repeat Lacking Clade Herbarium of National Plant Gene Bank. Voucher (IRLC) (Sanderson and Wojciechowski 1996; specimens of those taxa collected from the natural Wojciechowski et al. 2000, 2004; Wojciechowski habitats by us were deposited at the Tarbiat Modares 2003, 2005, 2006). Hedysareae as a whole occurs in University Plant Collection (see Table 1). Genomic dry open localities with a continental temperate or DNAs were isolated using the modified CTAB mediterranean climate, and is restricted to Eurasia, methods of Doyle and Doyle (1987). North America and the Horn of Africa with Socotra. Polymerase Chain Reaction (PCR). The complete After Hedysarum, with ca. 160 species, Onobrychis is nrDNA ITS region was amplified using primers ITS4 the second largest genus with ca. 130 species within the and ITS5 of White et al. (1990). Amplification was tribe (Mabberley 1990; Lock 2005). The genus is done in a DNA thermal cycler (Primus 96, MWG, restricted only to Eurasia and well represented in the Germany). The quality of PCR products were checked continental temperate and warm-temperate zones of the Irano-Turanian region. The Flora Iranica area is one of by electrophoresis in a 0.8% agarose gel (using 1X the main centers of diversity of the genus comprising TAE as the gel buffer) stained with ethidium bromide over 75 species in two subgenera divided into nine and then visualized under UV light. Nucleotides sections (Rechinger 1984). Hedysareae in general and sequences of PCR products were determined using Hedysarum (cf. Chennaoui et al. 2007) and Onobrychis cycle sequencing and an automated DNA sequencer in particular remain under-sampled and hitherto no through Gen Fanavaran Co. The same nrDNA ITS comprehensive molecular phylogenetic analysis primers ITS5 and ITS4 were used for cycle sequencing including almost all the genera has been undertaken. reactions. The present work using nrDNA ITS sequences is the Phylogenetic analyses. Sequences were aligned using first report on the phylogeny of the tribe with relatively clustal W 1.8 (Thompson et al. 1994) and adjusted its broader taxonomic sampling. The goals of our paper visually. Individual Sequences from taxa included in are, therefore, to: the present study have been deposited in DDBJ (see (1) Reconstruct the phylogenetic relationships of and Table 1. for accession numbers). Phylogenetic analyses delimit Hedysareae as a whole, (2) determine the were performed on the aligned data matrix using generic relationships within it and (3) clarify maximum parsimony method (MP) as implemented in infrageneric relationships within Onobrychis. the version 4.0b10 of PAUP* (Swofford 2002) installed in a Macintosh computer. The heuristic search option MATERIALS AND METHODS was selected using 100 replications of random addition Taxon Sampling. The sampling in this study includes sequence and TBR branch-swapping with MulTrees on 28 species of the tribe Hedysareae (sensu Polhill and steepest descent off. Analyses were conducted IRAN. JOURN. BOT. 13 (2), 2007 Ahangarian, Kazempour Osaloo & Maassoumi 66 using a successive weighting (SW) strategy (Farris were not resolved properly. Sulla, represented herein 1969). Weights were assigned to characters using the with 6 taxa, along with H. humile L., as nested within “reweight characters” option based on the rescaled it, forms a well supported monophyletic group (BP= consistency (RC) index (Farris 1989) with a base 100%) and for which, H. membranaceum Coss. & Bal. weight of 1. When the tree length and CI, RI and RC is a sister taxon. This
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