Turkish Journal of Turk J Bot (2019) 43: 386-394 http://journals.tubitak.gov.tr/botany/ © TÜBİTAK Research Article doi:10.3906/bot-1806-50

Hedysarum alamutense (-), a new from Iran, and its phylogenetic position based on molecular data

1 1, 2 2 Haniyeh NAFISI , Shahrokh KAZEMPOUR-OSALOO *, Valiollah MOZAFFARIAN , Mohammad AMINI-RAD  1 Department of Biology, Faculty of Biological Sciences, Tarbiat Modares University, Tehran, Iran 2 Research Institute of Forests and Rangelands, Agricultural Research, Education, and Extension Organization (AREEO), Tehran, Iran

Received: 25.06.2018 Accepted/Published Online: 14.01.2019 Final Version: 06.05.2019

Abstract: alamutense, a new species in the tribe Hedysareae DC. (Fabaceae), is described and illustrated. It belongs to the traditionally recognized Hedysarum L. section Multicaulia (Boiss.) B.Fedtsch., which extends over the West Alborz Mountains in northern Iran. This species is characterized by greenish stems and corolla persisting in fruiting stage and has mostly 1–2-jointed, unarmed, biconvex pods with short hairs becoming bald early, and flattened margins. Phylogenetic analyses of molecular data clearly unite this species with H. formosum Fisch. & C.A.Mey. ex Basin. The new species is characterized by 4 singleton nucleotide substitutions, suggesting it as a distinct taxon. Phylogenetic inferences are consistent with the interpretations of morphological features. Moreover, the lack of nucleotide variation among individual samples of H. formosum indicated that the species has not been diversified at the population level and reinforced our findings.

Keywords: Alborz Mountains, Hedysarum, phylogeny, section Multicaulia,

1. Introduction (Boissier, 1872; Rechinger, 1952; Chrtková-Žertová, 1968; The Hedysarum L., encompassing ca. 180 species, is Hedge, 1970; Fedtschenko, 1972; Townsend, 1974, 1984; the largest genus of the tribe Hedysareae, Fabaceae (Lock, Ranjbar et al., 2007; Xu and Choi, 2010) and compared 2005; Xu and Choi, 2010; Amirahmadi et al., 2014). In with the herbarium specimens deposited in TARI, W, and the last 3 decades new findings, mostly new species from LE. It was concluded that the specimens belonging to H. Iran, have increased the species number to 38 (Rechinger, subsect. Multicaulia ought to be considered an undescribed 1952, 1984; Ranjbar et al., 2006, 2007, 2008; Ranjbar, 2010; species with affinity to H. formosum. Here, it is described Akrami et al., 2011; Amirabadi-Zadeh, 2011; Dehshiri as H. alamutense sp. nov. The specimens in vegetative and et al., 2012; Dehshiri, 2013; Amirahmadi et al., 2014; fruiting stages were deposited at the herbaria of TARI and Bidarlord et al., 2015; Dehshiri and Goodarzi, 2016). Tarbiat Modares University (TMUH). A molecular study Most of the species of the genus in Iran belong to H. sect. based on concatenated nrDNA ITS + plastid (trnL-trnF Multicaulia (Boiss.) B.Fedtsch., and recent phylogenetic and matK) sequences of 16 representatives (23 accessions) studies (Duan et al., 2015; Liu et al., 2017; Nafisi et al., of Hedysarum was undertaken to infer the phylogenetic 2019) indicated that this section is composed of 2 clades of position and identity of H. alamutense. The recent findings traditionally recognized subsects: Multicaulia, Subacaulia from Iran together with the new species described herein (Boiss.) B.H.Choi & H.Ohashi, and Crinifera (Boiss) bring the species number to 39 in Iran. Choi & H.Ohashi. One clade is composed of 12 of species from the traditionally recognized subsections Multicaulia 2. Materials and methods and Crinifera. The second clade is a large assemblage of representatives of all 3 subsections. 2.1. Taxon sampling During a botanical trip to the West Alborz Mountains, Five individuals of the new species were collected by the Qazvin Province (northern Iran), in June 2013, we authors during field collection in June 2013. The collected encountered an unknown population belong to Hedysarum. and herbarium specimens were cross-checked with the The newly collected specimens were cross-checked with the various Hedysarum accounts given in the relevant references specific descriptions of the genus in the relevant references (Chrtková-Žertová et al., 1968; Hedge, 1970; Fedtschenko, * Correspondence: [email protected] 386

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1972; Townsend, 1974; Rechinger, 1984; Ranjbar et al., A list of species included in the phylogenetic analysis is 2007). Quantitative traits from the lowest to the highest given in Table 1. values were presented in the species description. The new 2.2. Molecular studies species belonging to H. sect. Multicaulia is described as Total genomic DNA was isolated from dried and fresh H. alamutense. The specimens in vegetative and fruiting materials using the CTAB method of Doyle and Doyle stages were deposited at the herbaria of TARI and TMUH. (1987). Polymerase chain reaction (PCR) was carried out

Table 1. Taxa included in the nrDNA ITS, trnL-F, and matK analyses.

GenBank accession no. Species (Voucher, source) nrDNAITS/cpDNA trnL-F/ matK Iran, Qazvin, Alborz Mountain, Alamut-e H. alamutense Nafisi & Kaz. Osaloo Gharbi District Kazempour-Osaloo & Bahadori LC466942*/LC466950*/ LC466946 105171(TARI) Iran, Qazvin, Alborz Mountain, Alamut-e Gharbi LC466943*/LC466951*/ H. alamutense Nafisi & Kaz. Osaloo District Kazempour-Osaloo 105173 (TARI) LC466947* H. candidissimum Freyn Turkey, Erzincan, Ganer 15114 (GAZI) LC404209/LC404292/LC404362 Iran, Chaharmahal and Bakhtiari, Mozaffarian, H. criniferum Boiss. LC404214/-/LC404365 57360 (TARI) Iran, Kohkiluyeh and Boyer-Ahmad, Yasuj, H. elymaiticum Boiss. & Hausskn. LC404219/LC404298/ LC404367 Ghahraman et al., 22526 (TUH) Azerbaijan, Nakhcivan, Grossgeim & Elinskaya H. formosum Fisch. & C.A.Mey. ex Basin. KP338165/ KP338286/ KP338542 s.n. (US 2394108), Iran, Azerbaijan, Khoy, Qotur, Kazempour- H. formosum Fisch. & C. A. Mey. ex Basin. LC404225/LC404303/LC404372 Osaloo & Nafisi 1394-3 (TMUH) Iran, Azerbaijan, Khoy, Seyed Tajaddin, H. formosum Fisch. & C.A.Mey. ex Basin. LC466944*/LC466952*/LC466948* Kazempour-Osaloo 1391-3 (TMUH) H. formosum Fisch. & C.A.Mey. ex Basin. Iran, Azerbaijan, Maku, Alizade 6741 (TMUH) LC466945*/LC466953*/ LC466949* H. huetii Boiss. Turkey, Artvin, Aytaç 86574 (GAZI) LC404233/LC404310/ LC404378 H. ibericum M.Bieb. Armenia, Vitek et al. 0005008 (W) LC404236/LC404312/ - H. kopetdaghi Boriss. Iran, Khorasan, K.H. Rechinger 5143-b (US) KP338172/KP338548/ KP338292 H. kotschyi Boiss. Turkey, Kahramanmaraş, Aytaç 3957 (GAZI) LC404241/LC404316/ LC404383 H. pestalozzae Boiss. Turkey, Adana, Vural 7517 (GAZI) LC404255/LC404329/ LC404396 Turkey, Malatya, Stainton & Henderson, 5468 H. pogonocarpum Boiss. LC404258/LC404331/ - (MSB) H. syriacum Boiss. Turkey, Ankara, Turgut 6304 (GAZI) LC404269/LC404342/ LC404408 Iran, Khorasan, Rabat-Sefid, Rafei, 30763 H. renzii Rech.f. LC404262/LC404335/ LC404401 (FUMH) H. singarense Boiss. & Hausskn. Iraq, Mosul, Rechinger, 0007043(W) LC404266/LC404339/ LC404405 H. varium Willd. Turkey, Kirikkale, Birden 1133 (GAZI) LC404272/LC404345/ LC404410 H. varium Willd. Iraq, Atrush, C. Chapman 9351 (US) KP338191/KP338309/ KP338567 H. varium Willd. Turkey, İçel, Sorger, 0004415 (W) LC404273/LC404346/ - H. varium Willd. Turkey, Erzincan, Altusoes 5627 (GAZI) LC404274/LC404347/ LC404411 Iran, Qazvin, Kazempour-Osaloo et al. 2013 H. wrightianum Aitch. & Baker LC404279/LC404352/ LC404415 (TMUH) viciifolia Scop. Spain, Huesca, Podlech 6892 (MSB) AB854512/AB854555/AB854604 Iran, Bandar-Abbas, Ahangarian & Kazempour cuneifolia Arn. AB329706/AB854558/AB854611 Osaloo 2006-4 (TMUH)

387 NAFISI et al. / Turk J Bot with 20 µL of the final volume of mixture containing 1.0 Osaloo & Bahadori 105171 (holotype: TARI, isotype: µL of template DNA (5 ng/µL), 0.5 µL of each primer (10 TMUH). pmol/µL), 10 µL of 2X Taq DNA polymerase Master Mix Diagnosis: Hedysarum alamutense is more closely Red (Amplicon, Cat. No. 180301, Germany), and 8.0 µL of related to H. formosum than to H. varium Willd., having sterile water. The nrDNA ITS region was amplified using the peduncles shorter than (rarely equal to) supporting leaves primers ITS5m (Sang et al., 1995) and ITS4 (White et al., (not longer); calyx indumentum not dense, calyx teeth 1990). PCR cycles consisted of predenaturation at 94 °C for 1.5–3 times as long as tube (not equal to 1.5 times); corolla 3 min followed by 30 cycles of 50 s at 94 °C for template with narrow wings, pale violet spot on the tip of keel or denaturation, 40 s at 53 °C for primer annealing, and 55 s at very rarely on wings, never on the standard, and persistent 72 °C for primer extension. A final elongation step of 7 min at in fruiting stage. The new species is distinct in possessing 72 °C was performed. The trnL-F region was amplified using greenish stems and long-lasting corolla and the following the universal “c” and “f” primers (Taberlet et al., 1991). The pod characteristics: biconvex, compressed with flattened PCR profile for this fragment consisted of predenaturation at margins and 1–2-segmented, covered with short hairs, 94 °C for 2:30 min followed by 30–48 cycles of denaturation becoming bald early, and without any bristles (Figure 1). at 94 °C for 50 s, annealing at 53 °C or 58 °C for 40 s, and H. formosum is characterized by grayish stems, deciduous elongation at 72 °C for 50 s. A final elongation step of 7 min corolla in fruiting stage, and dorsoventrally flattened and at 72 °C was performed. The partial matK gene was amplified 2–5-segmented pods, densely covered with long-lasting using the trnK685F (Steele and Wojciechowski, 2003) and short hairs and long, stiff bristles (Figure 2). The diagnostic matK1324R (Amirahmadi et al., 2014) primers. The PCR traits of the new species as well as H. formosum and H. profile for this fragment consisted of predenaturation at 80 varium are represented in Table 2. °C for 5 min followed by 33–41 cycles of denaturation at 95 Description: Perennial, (35–)50 cm long; stems °C for 50 s, annealing at 51–55 °C for 50 s, and elongation numerous, branched, erect, striate, zigzag, up to 7 mm in at 72 °C for 50–140 s with a final elongation step of 7 min diameter, greenish and foliate, subglabrous toward tip. at 72 °C. PCR products were separated by electrophoresis in 1% agarose gel stained with ethidium bromide and Stipules 3.5–7 mm long, herbaceous, with adpressed hairs, were photographed with a UVI gel documentation system in dried state brownish, lower ones connate, narrowly (UVItec, UK). PCR products along with the primers used triangular-acuminate from a wide base, upper ones free, for amplification were sent to Macrogen Inc. (Seoul, South subulate. Leaves imparipinnate 10–15.5 cm long; leaflets Korea) for Sanger sequencing. Sequences for marker datasets (3.4–)5–7 pairs, 12–21 × 4–10 mm, ovate to ovate-elliptic, of each taxon were edited using FinchTV 1.4 and aligned above glabrous and beneath more or less densely hairy. with the Web-based version of MUSCLE (Edgar, 2004) 20–50-flowered, axis of the up to under default parameters followed by manual adjustments. 19 cm and curved at mature stage; 3.5–10 cm Since both clades presented here are congruent in both long, shorter than supporting leaves; 2–4 mm long, nuclear and plastid datasets, we combined all of the datasets. brownish, hairy; bracteole 1 mm long, shorter than calyx Sequences generated in this study have been deposited in tube. Flowers existing in fruiting stage, erect in youth, GenBank (Table 1). hanged in fruiting stage. Calyx 5–7 mm, teeth 1.5–3 times The best models of sequence evolution were selected as long as tube. Corolla 12–17 mm long, over 3 times longer using the program MrModeltest version 2.31 based on the than calyx; standard 12–17 × 6–9 mm, obovate, deeply Akaike information criterion (AIC) (Posada and Buckley, emarginated at the apex, gradually attenuate at the base, 2004). Phylogenetic analyses of the sequence data were cream with large yellow spot at the base of standard; wings performed by Bayesian inference (BI) using MrBayes 3.2 10–13 × 1.5–3 mm, slightly shorter than keel, blade oblong (Ronquist et al., 2012) and maximum parsimony (MP) lanceolate, apex acute, claw 1.5 mm long, auricle 1 mm long, method as implemented in PAUP* version 4.0b10 (Swofford, very rarely with pale violet spot on the tip; keel 10–14 × 4–7 2002). Branch support values were calculated with 1000 mm, triangular, apex subacute; claw 2–3 mm long; auricle bootstrap replicates (Felsenstein, 1985). 0.5 mm long with pale violet spot on the tip, disappearing in mature flowers. Pods 1–2 (rarely 3) jointed, 5–7 × 4–5 mm, 3. Results elliptic, biconvex with flattened margin, ribbed reticulum, 3.1. Taxonomic treatment covered with white, short hairs becoming bald early. Hedysarum alamutense H. Nafisi & Kaz. Osaloo sp. nov. Phenology: Flowering and fruiting time is during June– (Figure 1) July. Type: Iran. Qazvin, Alborz Mountains, Alamut-e Gharbi Distribution and habitat: Hedysarum alamutense is District, Rajai Dasht, 1408 m, 10.06.2013, Kazempour- distributed over the West Alborz Mountains and grows in 1 Nylander JAA (2004). MrModeltest v2. Program distributed by the author. Uppsala, Sweden: Evolutionary Biology Centre, Uppsala University [online]. https://www.abc.se/~nylander/.

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B

C D

Figure 1. Hedysarum alamutense (from the holotype: TARI). A–B) Habit of plant. Scale in Figure B = 4.1 cm. C) Pod (5×); D) wings, standard, keel, androecium, calyx, gynoecium. red marl soils at 1300 m. The species is accompanied by Representative specimens: H. alamutense: Iran. annual grasses (Aegilops cylindrica Host, Bromus tectorum Qazvin, Alborz Mountains, Alamut-e Gharbi District, L., and Avena fatua L.). Rajai Dasht, 1300 m, 2013, 10 June, Kazempour-Osaloo Etymology: The specific epithet corresponds to the and Bahadori 105172 (Paratype: TARI, TMUH), Qazvin, type locality, Alamut area, West Alborz Mountains in Alborz Mountains, Alamut-e Gharbi District, 5 km after Qazvin Province. Rajai Dasht, 1300 m, 2013, 13 June, Kazempour-Osaloo

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Figure 2. A) Habit of Hedysarum formosum from the type specimen locality, plant height 25 cm. B–C) Pod. Bristles as long as width of pod.

105173 (TARI, TMUH), Zandjan to Mianeh, Mozaffarian not Paratype:), İçel, Sorger, 0004415 (W), Uşak, Balansa, 87122 (TARI). Plantes, Balansa 00010233 (W), Denizli, Boissier 00010252 H. formosum: Iran, West Azerbaijan, Urmia, Tarighi (W), Ankara, Bornmüller 00010251 (W), Denizli, Speta and Amini 1343 (TARI), West Azerbaijan, Khoy, Qotur 0006457 (W); Ankara, Sorger 0013372 (W), Çankırı, Speta valley, Rechinger 0005883 (W), West Azerbaijan, Khoy, 0006449 (W); İçel Sorger 0004415 (W); Ankara, Speta Qotur, Rechinger 0020785 (W), Iran, West Azerbaijan, 0006463 (W), Kastamonu, Edmondson 0008354 (W). Maku, Alizade 6741(TMUH), West Azerbaijan, Khoy, Armenia, Yerevan, Grossheim 0021933 (W). Seyed Tajaddin, Kazempour-Osaloo et al. 1391-3 (TMUH), Iran, Azerbaijan, Khoy, Qotur valley Kazempour-Osaloo 3.2. Key to some species of H. section Multicaulia and Nafisi 1394-3 (TMUH). Iraq, Kirkuk, Rechinger 1a. Plant with silvery, dense adpressed indumentum; 0013914 (W). leaflets obovate; wings 1/2 of keel, with a triangular acute H. varium: Iraq, Atrush, C. Chapman 9351 (US: auricle ...... H. singarense photo), Atrush, C. Chapman 9351 (US: photo). Turkey, 1b. without silvery indumentum; leaflets not Kırıkkale, Birden 1133 (GAZI: photo); Erzincan, Kemaliye, obovate (except H. candidissimum), wings ±shorter than Kayıkbaşı, yol kenarı 861 m, 04. 06. 2007 H. Altıözlü 5627 keel, with oblong lanceolate blade and acute apex and (GAZI: photo, this plant is named H. pogonocarpum, curved acute auricles ...... 2

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Table 2. Diagnostic morphological traits of Hedysarum alamutense, H. formosum, and H. varium.

H. alamutense H. formosum H. varium Stem color Greenish Grayish Grayish Leaflet pairs (3,4–)5–7 (3,5–)6–8(–9,10) (4–6–)7–9(–10) Mostly 1.5–3× shorter than Mostly 2–3× shorter than 1.5–2× longer than supporting leaf Peduncle supporting leaf, rarely equal supporting leaf, rarely equal or equal to Calyx teeth 1.5–3 times as long as tube 1.5–3 times as long as tube 1.5 times as long as tube Calyx indumentum Sparsely appressed Sparsely appressed Densely appressed Corolla Persistent in fruiting stage Deciduous in fruiting stage Persistent in fruiting stage Creamy with yellow spot at the Creamy with yellow spot at the 1- Creamy with yellow spot at the base of standard and temporal base of standard and temporal base of standard with temporal or Corolla color pale violet spot on the tip of keel pale violet spot on the tip of keel lasting violet spot on the tip of keel, and very rarely on wings and very rarely on wings wings, and sometimes standard 2- Purplish yellow Wing width 1.5–3 mm 1.5–3 mm 3–4 mm Pod segments 1–2, biconvex 2–5, flattened 2–3, flattened Long-lasting dense short hairs Pod surface Short hairs becoming bald early Hairy with short seta along with long stiff bristles

2a. Peduncle shorter than leaves ...... 3 11b. Flower pale violet, leaves 4–5 pairs, loment without 2b. Peduncle longer than leaves ...... 4 setae ...... H. ibericum 3a. Pod 2–5-segmented, flattened, hairy, with long stiff 3.3. Phylogenetic relationships bristles ...... H. formosum The concatenated sequence dataset (nrDNAITS,trn L-F, 3b. Pod 1–2-segmented, biconvex, hairy, without any and matK) for 18 analyzed taxa includes 2477 nucleotide bristle ...... H. alamutense sites, of which 288 (11.62%) are variable and 131 (5.2%) 4a. Setae longer than width of pods, pod width 8–16 are parsimoniously informative. The MP analysis resulted mm ...... 5 in a single tree with a consistency index of 0.861 and a 4b. Pods with short setae or without setae ...... 6 retention index of 0.948, along with bootstrap values 5a. Pod segments 12–16 mm long; leaflets ca. 12 × 6 (not shown). The MP tree is almost the same, in terms mm ...... H. kotschyi of bootstrap support and topology, as the BI tree. In the 5b. Pod segments 8 mm long; leaflets ca. 10 × 4 mm ...... BI tree the new species formed a well-supported sister ...... H. pogonocarpum group relationship with H. formosum in a clade (clade “B”) 6a. Plant with dense felted indumentum ...... comprising representatives of H. sect. Multicaulia...... H. candidissimum 6b. Plants without sericeous or felted indumentum .... 7 4. Discussion 7a. Leaves with 1–5 pairs of leaflets ±broad ovate or Phylogenetic analyses of the concatenated dataset fleshy ...... 8 generated 2 well-supported clades of ingroup taxa. 7b. Leaves with 4–9 pairs of leaflets, not broad ovate or Clade A was composed of some representatives of fleshy ...... 9 the traditionally recognized subsections Multicaulia, 8a. Pod 1–2-segmented ...... H. elymaiticum Crinifera, and Subacaulia (Choi and Ohashi, 2003) 8b. Pod 2–4-segmented ...... H. huetii including H. wrightianum Aitch. & Baker, H. renzi Rech.f., 9a. Stems 15–30 cm; leaflets 2.5–4 mm broad; loment H. criniferum Boiss., and H. kopetdaghi. Boriss. Clade B ±densely tomentose ...... H. pestalozzae consisted of 12 representatives from the traditionally 9b. Stems 15–50 cm; leaflets 2–12 mm broad; loment recognized subsections Multicaulia and Crinifera. not densely tomentose ...... 10 Hedysarum singarense Boiss. & Hausskn. was identified as 10a. Pod segments without setae ...... H. syriacum sister to the remaining species of the clade. The remaining 10b. Pod segments with or without setae ...... 11 species, in turn, constructed 2 well-supported subclades: 11a. Flower cream or purplish-cream, leaves with (4– one with 7 species encompassing H. pestalozzae Boiss., H. 6–)7–9(–10) pairs, loment with setae ...... H. varium varium Willd., H. syriacum Boiss., H. elymaiticum Boiss.

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Figure 3. Bayesian tree based on combined nuclear and plastid dataset of some representatives of sect. Multicaulia. Branch lengths are proportional to the number of nucleotide changes, as indicated above branches. Posterior probability and bootstrap values resulting from Bayesian inference and maximum parsimony analyses are given under branches, respectively.

& Hausskn., H. huetii Boiss., H. candidissimum Freyn in bristles, and corolla persisting in . On the other hand, Bull., and H. ibericum M.Bieb. and a second including H. H. formosum is characterized by dorsoventrally flattened pogonocarpum Boiss., H. kotschyi Boiss., H. formosum, and and 2–5-segmented pods, densely covered with long- H. alamutense. Hedysarum alamutense with 2 accessions is lasting short hairs and long stiff bristles, and diminishing well united with 4 accessions of H. formosum (Figure 3). corolla in fruit. In addition, the separation of H. varium These 2 species do share several traits including peduncles from the 2 species is supported by phylogenetic analyses shorter than supporting leaves, shape and size of wing and confirmed through morphological characters: grayish , calyx indumentums not dense, calyx teeth 1.5–3 appearance, peduncles longer than supporting leaves, times as long as tube and narrow wings, and pale violet sometimes purplish yellow petal color, violet spot on the spot on the tip of keel or very rarely on wings. Thus, the standard, and short setae on the pod surface (Table 2; position of H. alamutense in the reconstructed molecular Figure 3). phylogeny is congruent with our interpretation of its Hedysarum alamutense is characterized by 4 singleton morphological characters. The persuasive morphological nucleotide substitutions, 1 for nrDNA ITS and 3 for matK characters distinguishing H. alamutense from H. formosum and trnL-F, suggesting this species as a distinct taxon, are height and stature of plant, biconvex and compressed as are many of the closest species in Hedysarum. On the 1–2-segmented pods with flattened margins, covered other hand, no nucleotide variation was detected among 4 with short hairs becoming bald early and without any sampled plants of H. formosum, indicating that the species

392 NAFISI et al. / Turk J Bot has not been diversified at the population level. The new membranous, and transparent appendix, sometimes very species is a narrow endemic known only from the West slender and hardly objective, which begins after the pod Alborz Mountains, Iran, whereas H. formosum has a broad outline. The pod margin, a sticky and hairy part of the pod biogeographical distribution ranging from northwestern derived from the patching of parts without seed, is present Iran and northeastern Turkey to southern Transcaucasia. in other sections of Hedysarum. Considering the increase in Hedysarum species in Iran to 38 in the last 3 decades, the genus is susceptible to a high Acknowledgments speciation rate. In classifying new taxa it is necessary to The help of Ernst Vitek of herbarium W during a visit of define some new morphological characters. In this study the first author is highly appreciated. We would like to we described some new identifying characters between the thank the curators of the herbarium TARI for providing 2 closest species. herbarium facilities and herbaria GAZI, US, and W for There is merit in redefining pod “margin” and “wing” providing photos of the examined species. This work was to draw attention to a common misunderstanding in the supported by the Tarbiat Modares University Research identification of species of the genus. A specific character Council. of H. sect. Hedysarum is a wing in pods. The wing is a fine,

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