Hedysarum hyrcanum Bornm. & Gauba (1935: 39)

Nafisi, Haniyeh, Kazempour-Osaloo, Shahrokh, Kaveh, Akram & Mahmoodi, Mohammad, 2021, A Taxonomic Revision of the Genus Hedysarum L. (Fabaceae-Hedysareae) in Iran, Phytotaxa 511 (2), pp. 79-110 : 92-100

Hedysarum hyrcanum Bornm. & Gauba (1935: 39)

Typical characters:— Plant multicaule; branches with acute angles, rootstock thick. Leave with 4‒7 pairs of leaflets, sometimes highly remote; petiole decreasing from 4.5 cm in basal to topmost as to 0.5 cm long. Stipules ± 5 mm long, pale yellow-white, scarious, connate, acute, superiors lanceolate. Bract 5 mm long. Peduncles 1.5‒2 times as long as subtending leave, as long as to longer than stems. Calyx 6‒7.5 mm long. Corolla violet, 12‒15 mm long, standard as long as or longer than the keel, emarginate at the apex; wings 4‒7.5 mm. Pods glabrescent, not flattened, not rugose, nerves not elevated, segment 9 × 6 mm (Appendix 1).

Distribution:— Tehran and Azarbaijan, Iran.

─ ─ var. hyrcanum Type:— IRAN. Tehran: Karaj, Elburs, Kuh Dashteh , 2200 m, 10 July 1935, Gauba (B- 100277870!) .

Hedysarum hyrcanum var. viridissimum Bornm. & Gauba , l.c.: 112. Type:— Tehran: Karaj, Elburs, Kuh Daschteh , 2200 m, 10 July 1935,

E. Gauba (B-100277870).

Hedysarum elbursense Bornm. & Gauba (1940: 49) . Type:—Karaj, Kuh Dashteh, 2200 m, 24 May 1935, D. E. Gauba 1473 (B-

100591017!).

Typical characters:— Leaflets green, upper side loosely hairy. Pods orbicular-elliptic, without setae.

Representative specimens:— IRAN. Azarbaijan: Tabriz, Misho dagh, 15 June 2009, H . Razifard 40220 ( TUH) .

Tehran: Karaj, Elburs, Kuh Daschteh , 2200 m, 10 July 1935, E. Gauba (B-100277870); Damavand, Havir, 5 June 2015, V. Mozzafarian s.n.1,2 ( TUH); Kuh Dashteh near Karaj, 2200 m, 24 May 1935, D. E. Gauba 1473 (B-100591017!) .

Note:— H. hyrcanum var. incanescens Born. & Gauba , mentioned in Flora Iranica (1984) is not an accepted name in IPNI. Any specimen with pods with setae isn’t collected from Iran except introduced specimens by Gauba from the same locality of var. hyrcanum , 22 July 1935 (B-100277870!). Hedysarum elbursense as the sole specimen of this species collected from the same locality of H. hyrcanum var. hyrcanum type specimen, retrieved in H. kopetdaghi group of the plastid tree topology and share the shortness of calyx and ratio of the keel to standard with that, as well.

Hedysarum neyshaboricum Ranjbar (2010: 20) . Type: — IRAN. Khorasan: North of Neyshabor, around Abshar , Bar village , 1850–1900 m, 20 May 1996, A. Rafeie & H . Zangooei 27013 (holotype, FUMH-27013!; isotype, BASU) .

Typical characters:— Plant many stemmed, up to 50 cm, rootstock tough, appressed pubescent. Stem 6–19 cm long. Leave 11–14 cm long, with 2‒4 pairs of ovate-lanceolate, 20‒55 mm long, obtuse to round or rarely slightly retuse, mostly shortly mucronulate leaflets. Stipules 7‒9 × 4‒5 mm, light brown-scabrous, semi-connate; the lower ones triangular, upper ones narrower and acuminate from a wide base; petioles decreasing from lower leave 2.9–4.5 to upper ones 0.8–2.5 cm. Peduncles 20‒27 cm, longer than stems and 1.5‒2x as long as supporting leave in the flowering stage. Bract 5‒6 mm long. Bracteoles shorter than the calyx tube. Calyx 6–7 mm; calyx teeth 3–5 mm. Corolla dark purple, 2.4‒2.5 times as long as calyx; standard 15‒17 mm long, longer than the keel, glabrous, obcordate at the apex; wings 7‒8 mm long; keel 13‒14 mm. pods not seen.

Distribution:— It is a narrowly endemic species known only from the type locality, distributed in the dry zone of the mountainous region near Abshar, Bar village north of Neyshabor, Khorasan province.

Representative specimens:— IRAN. Khorasan: North of Neyshabor, around Abshar, Bar village , 1850–1900 m, 20 May 1996, A. Rafeie & H. Zangooei 27013 (holotype, Holotype, FUMH-27013) .

Note:— This species characteristics show a close relationship with those in H. paucifoliolatum . The length diversity of plant, peduncle, and corolla between them may refer to phenologic diversities, which need molecular surveys.

Hedysarum glabrifoliolatum Ranjbar (2008: 25) . Type:— IRAN. Khorasan: Kuchan, Farog to Oghaz-Kohneh , Ghaleh Safa mountain , 1750 m a.s.l., 23 June 1997, Faghinia & Zangooie 29350 (holotype: FUMH!)

Typical characters:— Plant glabrous. Stems 5.5‒10.5 cm, internodes 1–4 cm long. Stipule 4‒11 mm long, pale brownwhite, sparsely hairy, membranous, with brownish strips, connate at ¾ long, triangular to narrowly triangular-acuminate. Leave 5.5‒7 cm, with 1‒2 pairs of 19‒48 mm long, ovate-elliptic leaflets to lanceolate, round or obtuse, mostly tiny mucronate, glabrous to sparsely hairy. Inflorescence 6‒9 cm to up to 18 cm in fruit stage. Bract up to 5 mm long. Peduncles 7 cm and up to 16 cm in fruit stage, 1.5‒4.4 times as long as stems, 1‒2 times as long as subtending leave in fruit stage. Calyx 8 mm; calyx teeth 5 mm. Corolla 1.2‒1.86x as long as calyx; standard 15‒20 mm, slightly longer than the keel, obcordate at the apex; wings 6 mm; keel 12‒19 mm long. Pod 1‒4-jointed, compressed, transversely netted veined, glabrous, and without setae (Appendix 1).

Distribution:— It is endemic to northeastern Iran from the steppe and clay zones surrounding the Shah-Jahan mountain, Iran.

Representative specimens:— IRAN. Khorasan: Farooj, Curran-e kord, from the river toward resource, eastern slopes after canyon, 07 June 2009, F . Memariani & Zangooei 42903a,b ( FUMH); North of Farooj, 1‒2 km north of Safa castle, toward new Oqaz, 16 June 2009, F . Memariani & Zangooei 43110 ( FUMH); North of Farooj, 1 km north of Safa castle, toward Shokranlou, gypsi and serpantin hills, 01 June 2009, F . Memariani & Zangooei 42828 ( FUMH) .

Hedysarum paucifoliolatum Ranjbar & Olanj (2008: 25) . Type:— IRAN. Northern Khorasan: Shirvan to Bajgiran , Alkhas , 1800 m a.s.l., 9 June 1998, Faghihnia & Zangooie 31470 (holotype: FUMH!, isotype: BASU).

Typical characters:— Perennial, up to 35 cm tall, many stemmed, covered with short appressed pubescens. Stems internodes up to 3.5 cm long. Stipules 4‒6 × 2‒3 mm, pale brown-white, scarious, semi-connate, triangular, acuteacuminate at the apex, adnate to the petiole for 1‒1.5 mm, more or less with appressed hairs. Leave 3.5‒6.5 cm long, 1‒2(3‒4-) paired; leaflets 20–38 mm long, ovate or ovate-lanceolate, obtuse to rounded or slightly emarginate, mostly tiny mucronate. Inflorescence 7‒9 cm, 10‒16.5 cm in fruit stage. Peduncles as long as to shorter than subtending leave, 7–13 mm long, growing up to 16.5 at fruit stage. Bract 4‒5 mm long, membranous, with one vein. Calyx 5‒6.5 mm long; calyx teeth 3‒5 mm long. Corolla violet; standard 10‒14 mm long, slightly obcordate at apex; wings 4‒5 mm; keel 11‒15.2 mm long. Pods 1‒5-jointed, compressed, joints 6–8 mm long, without setae, transversely netted veined, unripe pods hairy at margins.

Distribution:— Endemic to N Khorasan, Iran.

Representative specimens:— IRAN. North of Khorasan: Khor, Kalat mount., 1350 m, 06 June 1988, M . R . Johartchi & Zangooei 16808 ( FUMH); Esfarayen, Ordaghan village in Milanlu rural district, 1670 m, 26 June 1995, M . Ranjbar 26013 ( FUMH); Shirvan to Bajgiran, Alkhas , 1800 m, 9 June 1988, Faghihnia & Zangooei 31470 ( FUMH); Bajgiran-Shirvan area , Oqaz Tazeh, Faruj, 5 km to South Castle from Shokranloo, 58° 13' 089" E, 37° 27' 956" N, 1626 m, 1 June 2009, S . Zarre, Y . Salmaki & A . Ebrahimi 38079 ( TUH) .

Hedysarum marandense Mozaffarian, Akrami & Maassoumi (2011: 17) . Type:— IRAN. Azarbaijan : Marand, between Zunuzagh and Kuhkamar, 2083‒2328 m, 25 June 2009, V. Mozaffarian & S . Akrami 93757 ( TARI!)

Typical characters:— Perennials, 25‒40 cm tall, many stemmed, appressed sericeous-hairy, caudex relatively thin. Stems 7.5‒15 cm, erect or ascending, stipitate. Stipules, 8‒9 mm long, bicuspidate, with white sericeous hairs, lower ones thick, upper ones ± scarious. Leave 7.5‒12 cm long, upper leaflets 5‒7 paired, lower ones 2‒4 paired; leaflets greenish-gray above, dense sericeous beneath, mucronulate, the topmost leaflets without a petiole. Peduncle longer or shorter than the stem, slightly longer than subtending leave. Bract 4 mm long, white, many striate, appressed hairy. Bracteoles 2, almost shorter than the tube, 3‒4 mm long. Calyx 8‒9 mm long; calyx teeth 5‒6 mm long. Corolla fuchsia, in sicco violet, 2.4‒2.75 times as long as calyx; standard slightly longer than the keel, 22‒24 × 15‒16 mm, deeply cleft, claw 6 mm long; wings 11‒13 mm long, 3/5x as long as the keel; keel 20 mm long. Pod with (-1)(2‒3)(5‒ 6-) joints, 8‒10 mm long, biconvex in the center, ribbed, hairy, with purple setae, turning to yellow, up to half of the pods, 1.5‒4.5 mm long (Fig 3; Appendix 1).

Distribution:— Endemic to Marand in Azarbaijan province of Iran.

Representative specimens:— IRAN. Azarbaijan: Marand, between Zunuzagh and Kuhkamar , 2083‒2328 m, 27 June 2009, V . Mozaffarian & S . Akrami 93757a,b ( TARI). Marand, between Zunuzagh and Kuhkamar , 27 June 2015, Sh. Kazempour-Osaloo & H . Nafisi 01-03-1394, 02-03-1394, 03-03-1394, 04-03-1394, 05-03-1394, 06-03-1394 ( TMUH); 19 km after Ahar to Tabriz , 38 21 56 N, 46 51 38 E, 1826 m, 18 June 2015, F . Attar , M . Mirtadzadini , A . Rastegar & Asgari 46235 ( TUH) .

Hedysarum garinense Dehshiri & Maassoumi (2012: 30) . Type:— IRAN. Hamedan province: Nahavand city, Gamasiab fountain, Garin Mountain , Yal Kabud pike, 34° 00' N, 48° 23' E, 2900–3000 m a.s.l., 23 July 2009, Dehshiri & Zarrini 97460 (holotype: TARI!, isotype: TARI)

Typical characters:— Plant caespitose, up to 17 cm high, with thick rootstock, multicaule, appressed sericeous white hairy vegetatively. Stems numerous, many-branched, suberect or ascending, thin, cylindrical, grooved, up to 7 cm long, 1–2 mm thick, arising from a woody rootstock, densely appressed sericeous white hairy. Stipules yellow-white, membranous, 2–4 mm long. the lower ones longer than the upper, highly connated, triangular, hairy. Leave 3‒10 cm, with 4–6 pairs of more or less remote leaflets. Peduncles 3–10 cm, hardly longer than supporting leave, longer than or as long as stem. Bract 2 mm and in fruit stage up to 4 mm long, villous. Bracteoles nearly as long as the tube. Calyx 5–6 mm long; calyx teeth 3.5–4.5 mm long, 2.5–3.0 times as long as the tube. Corolla 10–12 mm long, 2 times as long as calyx, emarginate at apex, pale yellow at first, later purple-magenta, at drying mauve; standard 7–8 mm long, 2/3x as long as the keel; wings 5–6 mm long, half as long as the keel; keel 10–12 mm long. Pods 1–2-jointed, joints elliptic, biconvex in the center, 10 × 7 mm, with 5 mm setae, slightly shorter than the width of pods (Appendix 1).

Distribution:— Endemic to the northern slope of the Garin Mountain, south of Nahavand in Hamedan province, Iran.

Representative specimens:— IRAN. Hamedan province: Nahavand city, Gamasiab fountain, Garin Mountain , Yal Kabud pike, 34° 00' N, 48° 23' E, 2900–3000 m a.s.l., 23 July 2009, Dehshiri & Zarrini 97460 (holotype: TARI) .

Hedysarum damghanicum Rech. f. (1979: 34). Type:— IRAN. Semnan: Damghan, Ahuvan neck, 35° 47' N, 53° 45' E, 1800 m, 21 May 1977, K. H. Rechinger 55373 (G-00365780 [Photo!]; Isotype, W-0007088!; TARI!); Semnan- Damghan , in declivibus argilloso-conglomeraticis jugi Ahvan (Ahuvan) 35°47' N, 53°45' E inter Semnan et Damghan, 1800 m, 07 June 1977, K. H. Rechinger 56498 (Isotype, W-0052998!)

Typical characters:— Plant suffrutex, multicaule, covered with appressed, greyish-green hairs, rootstock thick. Stem 4‒9 cm long, mostly with 2 leave. Leave with 5‒6 pairs of obovate, emarginated or truncate, mucronate, and sometimes remote leaflets, both sides tomentose; both basal and stem leave with long petioles. Stipules 5‒8 mm, lower ones tomentose, upper ones white, scarious ± hairy, long bicuspidate, with two veins. Inflorescence up to 16 cm long at fruit stage. Peduncle shorter than to as long as stems and supporting leave. Bract 3–4 mm long, Bracteoles 2, 0.5‒1 mm long, 1/4‒-1/2 calyx tube. Calyx 5.5‒7 mm long, calyx teeth 1.8‒2.25 time as long as tube. Corolla 2.4‒2.5 times as long as calyx, fresh petals pink with dark striate, in drying state mauve; standard 12‒14 × 9 mm, 3/4x as long as the keel, obcordate at apex, with a white spot at the base, in drying stages dark violet, with a small white spot at base; wings 7‒12 × 3‒4 mm, 0.42x as long as the keel; keel 13‒17 × 7 mm, in dry flowers very pale pink. Pods with 1‒2(3-) small, rugose joints, densely white tomentose, with scattered, tiny yellow setae at the center ( Fig 4 ; Appendix 1).

Distribution:— Endemic to Damghan, Iran.

Representative specimens:— IRAN. Semnan: Damghan, in declivibus argilloso-conglomeraticis jugi Ahuvan, 35°47' N, 53°45' E inter Semnan et Damghan, 1800 m., 21 May 1977, K. H . Rechinger 55373 ( W, WU); In declivibus argilloso-conglomeraticis jugi Ahvan ( Ahuvan ) 35°47' N, 53°45' E inter Semnan et Damghan, 1800 m., 07 June 1977, K. H . Rechinger 56498 ( W); 75 km of Damghan, 3 km of roadside assistance of Ahuvan , 12 May 2009, Sh. Kazempour- Osaloo & A . Amirahmadi 22-02-1388 a,b,c ( TMUH); Tehran-Mashhad road, Low hills along the road, 23 miles east of Semnan , elev. 1800 m., 07 July 1972, R . Alava & Iranshahr 10834 ( W) .

Hedysarum criniferum Boiss., Diagn. (1843: 2) . Type:― Isphahan: 32°44'22" N, 54°11'52" E, s.d., P. M. R. Aucher- Eloy 1081 (Syntype, G-00330388; Isosyntype G-00382780[Photo!]), 32°44'22" N, 54°11'52" E, s.d., P. M. R. Aucher- Eloy 4446 (Syntype G-00330389[Photo!]); 32°44'22" N, 54°11'52" E, s.d., P. M. R. Aucher-Eloy 4439 (Syntype G- 00330390[Photo!]; Isosyntype G-00382779[Photo!])

Hedysarum ecbatanum Beck, Denkschr. (1886: 51) . Type:― Hamadân: In agro Ecbatanensi (Media), wegränder bei Abatabad nächst Hamedan, 34.77° N, 48.55° E, June 1882, J. E. Polak s.n. (WU-0073037).

Hedysarum orumiehense Ranjbar (2010: 20) . Type : Iran. West Azarbaijan , Orumieh , ShahrChai dam, near Band, 1530 m, 11 June 2005 (fl., fr.), M. Ranjbar 7092 (holotype, BASU; isotypes, FUMH, MO, TARI, W) .

Typical characters:— Stem 10‒21 cm long. Leave (5‒6)7‒9 paired; leaflets 9‒16 mm long, elliptic-linear, more or less remote, truncate with tiny mucron, or acuminate, the topmost leaflets without petioles. Peduncle as long as to shorter than subtending leave and as long as to longer than stems. Bract 2.5‒5 mm long. Bracteoles 2, 1/2 of the calyx tube, 1‒ 1.5 mm. Calyx 5‒7.5(11‒13-) mm long, calyx teeth 1‒3x as long as tube. Corolla 13‒18 mm long, mostly 2.5 times as long as calyx; standard 11-15mm long, 5/7 to 6/7x as long as the keel, obcordate at the apex, in fresh state magenta, with white, obcordate spot up to the base; wings white, 6–9 mm, 0.33‒0.66x as long as the keel; keel 12‒18 mm long, blade white, tinged with pink at the tip. Pod with (-1‒2)3‒4 joints, joints 6‒8 × 5‒7 mm, sub-reticulated, sub-glabrous, sub-rugose, with long, reddish than yellow setae, slightly shorter than the width of pods (Appendix 1).

Distribution:— Isfahan, Tehran, Arak, Hamedan, Kermanshah, Tabriz, Kurdestan, Kohkilu-e, and Boyer Ahmad, Iran.

Representative specimens:— IRAN. Isphahan: 32°44'22" N, 54°11'52" E, s.d., P. M. R. Aucher-Eloy 1081 (Syntype, G-00330388[photo!]; Isosyntype G-00382780[photo!]), 32°44'22" N, 54°11'52" E, s.d., P. M. R. Aucher- Eloy 4446 (Syntype G-00330389[photo!]); 32°44'22" N, 54°11'52" E, s.d., P. M. R. Aucher-Eloy 4439 (Syntype G- 00330390[photo!]; Isosyntype G-00382779[photo!]); Musiabad ca. 90 km West of Isfahan, sedimentary hills, 6000- 6500 feet, H. S. Gentry 14893 (US [Photo!]) ; Ghameshloo protected area, Kahvare to Ghameshloo , 2200 m, May 1996, Yosofi 874 (Isfahan); Ghameshloo protected area, Tappeh Khan Ali , 2000‒2100 m, May 1996, Yosofi 981 ( TARI) .

Hamadân: In agro Ecbatanensi (Media), wegränder bei Abatabad nächst Hamedan, 34.77° N, 48.55° E, June 1882, J . E . Polak s.n. (WU-0073037). Azarbaijan: 50‒ 30 km E Tabriz, 38°05' N, 46°18' E, versus Mianeh , 37°23' N, 47°45' E, 31 July 1965, K . H . Rechinger 32627 ( W) . Kermanshah: Gahvareh, Mela Ferdin , 1800 m, June 1992, Nemati & Mirabdali 3530 (Kermanshah); Gahvareh, 1 km to Tootshami, 1500 m, 30 May 2004, Nemati & Mirabdali 6804 (Kermanshah); Gahvareh, Hovaru, 1500 m, 29 June 1998, Nemati 6066 (Kermanshah) . Tehran: Arak to Mahallat, Latedar , 2100‒2500 m, June 1984, Mozaffarian & Maassoumi 47927 ( TARI), Saveh , Ghatghabad , Ghazmabad , June 1991, A . Ghahreman & V . Mozaffarian 9926 ( TARI) . Qazvin: Avaj , ca. 3 km from Avaj to Hamedan (close to Sultanbolagh), N . 35° 34', E. 49° 13', 1980 m, 24 June 2005, V . Mozaffarian 87332 ( TARI). Kohgiluyeh and Boyer-Ahmad : Sisakht , kuhgol and Kokhedan , 2200 m, May 1995, V . Mozaffarian 74436 ( TARI) . Kurdistan: Kauleh , between Sanandaj and Saqes, steep N . and N. W . facing clay slepes, 2070 m, 21 May 1966, J . C . Archibald 2137 ( W) .

Notes:— As mentioned earlier, the ratio of standard to keel in Hedysarum is supposed to be for mature flowers. As a matter of fact, in this species, the standard is as long as the keel in immature flowers and longer than keel in mature ones.

Hedysarum melanotrichum (Boiss.) H. Nafisi & Kaz.Osaloo Comb. Nov. ≡ Hedysarum criniferum var. melanotrichum Boiss. Type:— Between Aghkent and Bagh, on the road between Tehran and Tabriz (Bunge!)

Description:— Perennials, 21‒33 cm long, rootstock thick, many stemmed, external stems decumbent and internals erect, sometimes with the remnant of stems of previous years in the rootstock. Stem sulcate, up to 17 cm long, branched only from upper sides, internodes developed 6‒11 cm long. Leave imparipinnate, with (-4‒7)8‒11 pairs of mostly narrow elliptic, more or less remote leaflets, upper leave with very short to none petioles; leaflets 6‒12 × 2‒4 mm, length/width of leaflets vary between 3 to 4 times, upper surface much less hairy than beneath, the topmost leaflets without petioles. Stipules 3.5‒6 mm long, white, membranous, striped, and hairy to glabrescent, basal stipules semiconnate, shortly bicuspidate from a wide base, upper ones, with free lanceolate cuspids. Peduncles as long as to shorter, rarely longer than subtending leave. Bract 2‒5 mm long. Bracteoles 2, delicate, 1‒1.2 mm long, one half of the calyx tube. Calyx size growing, 4‒8 mm, the teeth in fruit stage, up to 1‒3.3 times as long as tube. Corolla 11‒17 mm long, white and or bicolor of white-magenta; standard 9‒14 × 7‒9 mm long, 5/7‒ 1x as long as the keel, obcordate at the apex, gradually attenuate to the base, magenta with a white spot up to the base; wings white, 5‒9 mm long, 0.41‒0.53 times as long as the keel; keel white with pink tints at the apex, 11‒17 mm long, blade white tinged with light violet at the tip. Pods asymmetric, elliptic to semi-ovate, 10-12(14‒17-) × (-7) 8‒10 mm, mostly 1‒2 and rarely 3‒4-segmented, with dark purple or navy blue turning to yellow setae in drying, as long as or up to 2.5 times longer than the width of pods ( Fig 4 ; Appendix 1).

Distribution:— Azarbaijan, Hamedan, Zanjan, and Isfahan provinces of Iran.

Representative specimens:— IRAN. Azarbaijan: 52 km of Urmia toward Oshnavieh, Gardane , 1800 m, 16 June 2012, Sh. Kazempour-Osaloo & A . Amirahmadi 27-03-1391 ( TMUH); Urmia, Ghasemloo valley , Natural resource preserved region, 1500 m, 7 June 2000, Heidari, Shanaki & Alizadeh 1710 (Zanjan). Tekab , Qholan valley, 1900 m, 17 August 2018, B . Alizadeh & S . Ghasempoor 5430 (Urumia). Mianeh , Bozgoush Mount region , Kuh-e Bozgoush from Varzegan valleys, 37° 41' N, 37° 41' E, 1960 m, May 2007, V . Mozaffarian 93360 ( TARI). Hamedan: Avaj , 1800‒2000m, 1 June 1988, V . Mozaffarian 64437 ( TARI); Avaj , 1800–2000m, 1 June 1988, V . Mozaffarian 64437 ( TARI); Avaj , 1800–2000m, 1 June 1988, V . Mozaffarian 64437 ( TARI); Zanjan: Gjareh-Pushtlu, north of Sohrin village , 2000 m, 11 June 1998, A . Mousavi 2385 (Zanjan); 40km toward Tabriz road, Abdol village , 1900 m, 18 June 1997, A . Mousavi 1692 (Zanjan); Qeydar , between Chokhoryort and Avouch Bolagh village, 2000m, 19 June 1997, A . Mousavi 1695 (Zanjan); Mahneshan, Dar-Bande Habash , 2200 m, 6 July 1994, A . Mousavi 249 (Zanjan); between Zanjan and Soltaniyeh, Hidaj , Pirsega village 15 June 2014, Sh. Kazempour-Osaloo et al. 25-03-1393 ( TMUH); Hidaj , Sarve Jahan village , 2000 m, 13 June 1998, A . Mousavi 2262 (Zanjan); 20 km of Zanjan-Tabriz road, 26 km Esfejin to Mirjan road, Agh-Bolagh village , 36° 40' 45" N, 48° 02' 49" E, 2212 m, 01 June 2013, M . Mahmoodi 99658 ( TARI) 20 km of Zanjan-Tabriz road, 26 km Esfejin to Mirjan road, Agh-Bolagh village , 36° 40' 45" N, 48° 02' 40" E, 2212 m, 01 June 2013, M . Mahmoodi 99657 ( TARI). Isfahan: Chelgard, Daran , 32°59′19″ N, 50°24′46″ E, Kemintak , 2000 m, 19 June 1973, Iranshahr & Moussavi 15521- E ( W); Semiron, Vanak to Dalan-Kuh, Shams-Abad and Ganat, 2200 m, June 1984, M . Nowroozi & J . Bozorgi 3624 ( TARI); Semiron, Vanak to Dalan-Kuh , Shams-Abad and Ganat , 2200 m, June 1984, M . Nowroozi & J . Bozorgi 3622 ( TARI) . Kohkiluye and Boyer Ahmad: Between Yasuj and Kakan, against to Cheshme Chenar village , 2200 m, 26 May 1995, V . Mozaffarian 74448 ( TARI) .

Discussion:— The specimen collected from Aghkent et Bagh on the road between Tehran and Tabriz (Bunge!), classified by Boissier (1872) as a variety of Hedysarum criniferum , distinguishing with large pods and dark colorful setae. In the tree topology resulted from molecular phylogenetic analyses using nrDNAITS dataset, the group with large pods possesses four singleton nucleotide site differences in comparison with the members of H. criniferum group and placed out of this group in a polytomy. The diagnostic key characters in identifying H. melanotrichum are the size and shape of pods. As a matter of fact, however, mostly corolla and calyx size in H. criniferum group are longer and pods mostly 3–5 jointed, but the number, size, and shape of leaflets, size of flowers, the ratio of standard to the keel, the ratio of the wing to the keel, size of calyx and bract, the ratio of calyx teeth to the tube, number of pods and color of setae in both groups involved in overlaps.

The specimens collected from Zanjan have wider leaflets even in upper leave and calyx size reaches up to 6.5 mm long, whereas the specimen from Urmia 27-03-1391 (TMUH) comprises slender leaflets and calyx size reaches up to 8 mm (in fruit stage) and bigger pod joints (14–17 mm). Moreover, we detected populations with monochrome white flowers and populations with magenta tint at the apex of the standard petal growing in Zanjan.

H. varium with standard petal longer than the keel, wings oblong-lanceolate, slightly shorter than the keel, having a dark yellow spot at the middle of standard and peduncles longer than or as long as supporting leave, is distinct from H. melanotrichum with keel longer than standard, wings oblong-obovate, 1/3x keel, monochrome, white flowers, and peduncles as long as or shorter than subtending leave, which is highly supported with phylogenetic analyses, too.

It is merit to note that the careful examination of specimens cited in Ranjbar et al. (2006) as H. singarense Boiss. & Hausskn. ex Boiss. , confirmed to be erroneously identified. As our morphological and phylogenetic studies supported the affinity of the examined samples to H. melanotrichum , placed in the H. subsect. Crinifera clade. Whereas, the actual specimens belonging to H. singarense ( Iraq, 11635 (W)), characterized with appressed silky hairy stems, 3‒5 pairs of oblong or ovate leaflets, 10‒25 × 7‒16 mm, silvery with closely appressed hairs on both side and with triangular acute auricles in petal wing, nested in the H. subsect. Multicaulia clade (Nafisi et al. 2019a; Figs. 1 , 2 ; Appendix 1, 2). Bract 2‒4 mm long. Bracteoles 2, 1‒ 1.5 mm long, ½ calyx tube.

Hedysarum atropatanum Bunge ex Boiss. (1872: 2) . Type :— Azarbaijan : In herbidis promontorii ad pag. Seidihadzi, Khoy, N 54 °11'52ʺ, EN 32°44'22ʺ, 17 May 1828, J. N. Szovits 236 (Syntype, G-00330385 [photo!])

Typical characters:— Plant suffrutex, multicaule, branches with obtuse angles, rootstock thick. Leave with (-4)(5‒ 8)(9‒11-) pairs of sometimes remote leaflets. Peduncle shorter to as long as the stem, equal to slightly longer than subtending leave. Bract 2‒4 mm long. Bracteoles 2, 1‒ 1.5 mm long, ½ calyx tube. Calyx teeth 1.5‒2 times as long as tube. Corolla magenta, 2.5 times as long as calyx; standard 11‒15 mm long, ¾ keel, in fresh state magenta, then fuchsia and mauve, with a white spot at the base, obcordate at the apex; wings 7‒8 mm long; keel 14‒16 mm long, blade not white or pale. Pods with and without setae ( Fig 4 ; Appendix 1).

Distribution:— W Iran, and S Transcaucasia.

─ ─ var. atropatanum

Typical characters:— Pods with long setae, up to slightly shorter than the width of pods

─ ─ var. glabrum H. Nafisi , var. nov. Type:— Azarbaijan : Khoy , Seyyed Tajeddin, 25 June 2015, Sh. Kazempour- Osaloo & H . Nafisi 2015 -02, 2015-04, 2015-05 (TMUH).

Typical characters:— Pods without setae or with black spots.

Representative specimens:— IRAN. Azarbaijan: Khoy, Seyyed Tajeddin , 25 June 2015, Sh. Kazempour-Osaloo & H . Nafisi 2015 -01, 2015-02, 2015-06, 2015-07, 16-03-1394 ( TMUH); In herbidis promontorii ad pag. Seidihadzi, Khoy , 54°11'52" N, 32°44'22" E, 17 May 1828, J . N . Szovits 236 (Syntype, G-00330385[photo!]); inter Marand and Sufian , 1600‒1750 m, 6 July 1971, K . H . Rechinger 41801 ( W); 3 km S Khoy versus Shahpur , 1500 m, 12 June 1971, K . H . Rechinger 41190 ( W); Inter Tehran and Tabriz , 0 m, 32°44'22" N, 54°11'52" E, June 1859, A . A . von Bunge s.n. (Syntype, G-00330386 [photo!]); Tabriz, Mishodagh , 27 June 2015, Sh. Kazempour-Osaloo & H . Nafisi 2015 -03 (TMUH)

Notes:— Pods are unribbed at the early stage of growth. There are a couple of specimens collected from the type locality, Seyyed Tajeddin, having pods with and without setae. The specimen ( var. glabrum ) collected from Mishodagh, somewhat far away from the type locality, has pods without setae.

Hedysarum fallacinu m Rech.f. & Aellen (1951: 75). Type:— IRAN. Khorasan: 36.88° N, 59.28° E, Montes Hazar Masdjid, inter Gosh et Talgor, ca 1800 m, 07 June 1948, K. H. Rechinger et al. 5143a (Holotype, W-0005666!) Hedysarum longipedunculatum Ranjbar & Karamian (2008: 26) . Type:— IRAN. Northern Khorasan : West Bojnord , between Darkash and

Havar , 1350 m a.s.l., 8 Jul 2001, Johartchi 33858 (holotype: FUMH, isotype: Bu-Ali-Sina Univ. herbarium) .

Description:— Plant rhizome suffrutescent, few-or multicaulis, few branched mostly from the base, at first stages with short white hairs, then glabrescent. Stems 2‒56 cm, zigzag. Lower leave with 3‒6 pairs and upper leave with (7‒9)(10-) pairs of leaflets; leaflets elongated elliptic, 10‒30 × 4‒7 mm, or short obovate distinctly emarginated, 15‒29 × 7‒11 mm, upper side glabrescent, lower side loosely or densely appressed. Stipules membranous, glabrous, pale brown. Inflorescence short, and many-flowered before anthesis, post-anthesis 4‒13 cm long. Peduncles 11‒31 cm, as long as, shorter, or longer than stems, exceeding the leave. Calyx (-6)(8‒10) mm. Corolla 1.7‒2 times as long as calyx, violet-mauve; standard 12‒18 × 7‒8 mm long, mostly as long as, or slightly longer than the keel, emarginate; wings 5‒6 mm long, rounded at the apex, blades 3‒5 × 2‒3.5 mm, auricles ca. 1 mm long, claw ca. 2 mm long; keel 11‒18, blades subtriangular, subacute at the apex, 8‒13 × 4‒6 mm, claw 2.5‒3 mm long. Pods 1‒2(3-)-jointed, densely short appressed, and 1.5‒3 mm long, purple setae.

Distribution:— Khorasan province and Golestan national park in Gorgan, Iran.

Representative specimens:— Iran. Khorasan: West of Bojnord, between Darkash and Havar, 1000‒1200 m, 10 May 2000, M. R. Johartchi & Zangooei 32768 ( FUMH); Northwest Neishabour, Abshar Bar , 2000 m a.s.l., 29 June 2003, Zangooei 38890 ( FUMH); Hazar Masjid , inter Gash et Tolgor , ca 1800 m, 07 June 1948, K. H. Rechinger et al. 5143a, c ( W); West of Bojnurd , between Darkash and Havar, 1350 m, 08 July 2001, M. R. Johartchi 33858 (holotype, FUMH); MRSWP, vic Almeh, 20 June 1969, J. Hassinger & D. Ferguson 25336 ( W); West of Bojnurd , 13 km of Bojnurd to Hesar Hoseini, 1681 m, 18 May 2016, M. R. Johartchi & F. Memariani 45791 ( FUMH); West of Bojnurd , 3 km West of Juzak, natural resource Gorogh, 1‒3 km by road, 1320‒1370 m, 23 May 2010, F. Memariani & M. Arjmandi 43690 ( FUMH); West of Bojnurd , Gorkhod sanctuary zone, Bash Kalage road to Chale Khuki, Artemisia-Juniperus co mmunity, 1800 m, 19 May 2010, F. Memariani & M. Arjmandi 43684 ( FUMH); Bojnurd, Gorkhod sanctuary zone, between Razavi village and Gharabil Robat, 1600 m, 31 May 1998, M. Ranjbar 31184 ( FUMH); Bojnurd, Robat Garabil, 3 km of old road of Jajarm, 1600 m, 29 May 1998, Hojjat & Zangooei 31084 ( FUMH); Bojnurd to Espharayen, 49 km of Mahphan, 1500 m, 23 May 1983, M. R. Johartchi & Mahvan 10286 ( FUMH); Bojnurd, 85 km, Shahpasand road, 1270 m, 29 May 1972, F. Foroughi 6359 ( TARI); Gorgan: 1 km after Kosh-Yeylagh to Shahrud, 36° 51' 32" N, 55° 20' 60" E, 1165 m, 02 June 2009, S. Zarre, Y. Salmaki & A. Ebrahimi 38090 ( TUH); Mohammad Reza Shah wildlife Park, Almeh, 1750 m, 20 June 1974, Wendelbo & Foroughi 12744 a,b ( W); Park-e Golestan, Almeh , 1300‒1700 m, 5 June 1987, A. Ghahreman & V. Mozaffarian 5923a, b, c ( TUH); Park-e Golestan, Almeh , 1350‒1700 m, 05 June 1987, V. Mozaffarian & Abouhamzeh 59062 ( TARI), (Appendix 1) .

Discussion:— According to nrDNA ITS phylogenetic tree, H. longipedunculatum and H. fallacinum (both from type specimens) formed a subclade, with the highest support value. Also, the subclade indicates a close relationship with H. sauzakense , H. bucharicum and H. malurense , as H. sauzakense group, which despite the significant differences of shape and number of leaflets, all share pods with setae detached from H. kopetdaghi group with pods without setae. H. fallacinum also shares short wing petals and elliptic and numerous leaflets with H. bucharicum . Morphologic experiments are handled with molecular phylogenetic analyses of nrDNA ITS and the combined trn L-F and mat K dataset which confirmed the separation of two groups (Nafisi et al. 2019a). Also, both species possess seven singleton nucleotide differences in their nuclear and plastid sequences, which is of significant value among Hedysarum species.

In the plastid tree topology, H. longipedunculatum and H. fallacinum formed a clade, but other members were placed in a polytomic relationship with this clade (Nafisi et al. 2019a; Fig 1 , 2 , Appendix 2).

The specimens 21927a,b,c (MSB-0243879!; W-020192!), distributed in Afghanistan, precisely determined by Podlech as H. cf. sauzakense , possess 4–5 paired leaflets, 6‒11 cm long peduncles, 12‒14 mm long corollas, 6‒7 mm long calyxes in share with H. sauzakense , but distinctly different in leaflet shapes. Considering the overlapping distribution area of H. sauzakense and H. malurense , the hypothesis is that, H. cf. sauzakense suppose to be a hybrid between H. sauzakense and H. malurense or probably H. bucharicum and through subsequent hybridizations and immigration to the northeast of Iran, H. fallacinum is developed ( Fig 5 ). H edysarum cf. sauzakense despite high resemblances to 21927a-b-c, differs in the number of leaflets and ratio of wing petal to keel.

It is noteworthy that some of the descriptive and diagnostic characters of H. longipedunculatum ( Ranjbar et al., 2008) are not confirmed here. As a matter of fact, the calyx/ corolla ± 1/2 (not ± as long as corolla) and wing petal/ keel= 0.3‒0.5x (not ± as long as the keel). Calyx size in H. fallacinum group was not more than 10 mm long, but 6–8 mm (e.g. 5143a (W-0005666), 12744a (W-00017847); TARI-25336) and or 8‒10 mm (e.g. 6359, 59062 (TARI), 5143b (W-00017848).

Hedysarum macranthum (Freyn & Sint.) B.Fedtsch. (1948:13) . ≡ Hedysarum micropterum (Bunge in Boiss) B.Fedtsch. subsp. macranthum Freyn & Sint. (1906: 6(2)). Type:— TURKMENISTAN: Ahal, Ashgabat, Suluklü (Saratowka), the limits of Iran, in mountains above Nephton, 04‒26 May 1900, P. E. E. Sintenis 237 (syntypes, W-0006790!, WU- 0096350[Photo!]); 392 (syntypes, W-00010229!, JE-00010230, W-455134!, W-0006789!, WU-0096351[Photo!]) Typical characters:— Plant 40‒60 cm, multicaule. Stipules membranous, 6–8 mm long, connate from base to the middle, bicuspidate. Leave with (-4)5(6-) paired. Leaflets of first years shorter and elliptic. Peduncles a little bit shorter or as long as subtending leave. Bract 8‒9.5 mm. Bracteoles 2, as long as calyx tube, 3 mm long. Calyx 9‒11 mm long, calyx teeth 3‒4x as long as tube. Corolla 1.48‒1.6x as long as calyx; standard 11‒15 mm, 5/6x as long as the keel, white with pale violet shades, and a large cream spot at the base; wings 6‒7 mm long, 2/5x as long as the keel; keel 13‒18 mm long, white with green nerves and pale violet and sometimes green shades. Pods 2‒3(4-)-joined, rugose, 6 mm long, with setae as long as width od pods (Appendix 1).

Distribution:— Turkmenistan and Khorasan province of Iran.

Representative specimens:— IRAN. Khorasan: Kalat-e-Naderi, Khesht mountains , 1500 m, 02 June 1997, Faghihnia & Zangooei 29030 ( FUMH) ; North east of Kalat-e-Naderi, Kalat toward Khesht Naderi , 1200 m, 24 May 1994, Faghihnia & Zangooei 24030a,b ( FUMH) ; 10 km of Kalat Naderi to Archangan , 1100 m, 01 June 1997, Faghihnia & Zangooei 28959 ( FUMH) ; Mashhad road, Kalat, 15 km of Kalat , 1000 m, 30 June 1992, Faghihnia & Zangooei 22105 ( FUMH) ; inter Ghuchan and Dar Gaz, begining of Ali Bolagh road, 05 June 2003, Ajani & Zangooei 34705 ( FUMH) .

Notes:— Examining several specimens of this species indicate that in contrast to the description of species, the keel is many times longer than standard, not shorter. Some specimens from Turkmenistan (W-00010229, W-00010230, W-0096351, W-0006789, Iso W-455134, WU-0096350, WU-0006790,) were also observed.

Hedysarum gypsophilum Dehshiri (2016: 53) . Type:— IRAN. Lorestan: Khorramabad-Andimeshk highway, 83 km from Khorramabad towards Andimeshk and 97 km from Andimeshk towards Khorramabad , Khargushan mountain , 33° 04' N, 48° 14' E, 1200–1300 m a.s.l., 15 May 2013, M. M. Dehshiri 51553 (holotype IRAN; isotype TARI)

Typical characters:— Plant 15‒40 cm tall, multicaule, many-branched, distributed in low mountain slopes, gypsumrich soils. Stipules yellow, membranous, 4–6 mm long, bicuspids 2 mm long. Leave (-2)3‒5(6-) paired; leaflets 15‒ 32 mm long, oblong, remote, obtuse-marginate, sparingly pubescent to glabrescent above. Peduncles shorter than subtending leave. Bract 5‒8 mm long. Bracteoles 2, 2 mm long. Calyx 8‒10 mm; calyx teeth 6‒7 mm, 2.5‒3x as long as tube. Corolla 1.3x as long as calyx, pink, 10.5‒12 mm, at dry violet; standard 10‒10.5 mm long, slightly obcordate; wings 3.5‒4.5 mm long; keel 10.5‒12 mm long, as long as to slightly longer than standard. Pods 1–2(3-) jointed; joints flattened, widely elliptic, 8–10 mm long, transversely ribbed, pubescent, densely covered with very short appressed sericeous white hairs and beset along nerves with numerous, slightly shorter than the width of pods, purplish setae (Appendix 1).

Distribution:— It is only known from the type locality and vicinities of eastern slops of the Khargushan mountain (close to Pole Zal), southern of Khorramabad (Lorestan Province).

Representative specimens:— IRAN. Lorestan: Khorramabad, 83 km from Khorramabad to Andimeshk , Khargushan mountain , 33° 04'N, 48° 14'E, 1200‒1300 m, 15 May 2013, M. M . Dehshiri 51553 (Holotype, Isotype; TARI!) .

Figures

FIGURE 1. Fifty percent majority rule consensus tree resulting from Bayesian inference of the nrDNA ITS dataset. Numbers above branches are posterior probabilities (PP) and numbers below the branches are likelihood bootstrap values; values <50% are not shown. Numbers after species names are herbarium numbers, given only if more than one accession per species is included along with distribution areas.

FIGURE 2. Fifty percent majority rule consensus tree resulting from Bayesian inference of the combined plastid dataset (trnL-F, matK). Numbers above branches are posterior probabilities (PP) and numbers below the branches are likelihood bootstrap values; values <50% are not shown. Numbers after species names are herbarium numbers, given only if more than one accession per species is included along with distribution areas.

FIGURE 4. E. H. damghanicum. Photo from E. Amirahmadi. 1E: Plant. 2E: Pod.; F. H. alamutense. Photos from Sh. Kazempour-Osaloo and Sh. Bahadori. 1F: Plant. 2F: Pod.; G. H. melanothricum. Photos from M. Mahmoodi. 1G: Plant. 2G,3G,4G: Pods.; H. H. atropatanum. 1H: Plant. 2H: Standard petal. 3H: Keel petal. 4H: Pods of (var. atropatanum) 5H: Pods of (var. glabrum).

FIGURE 5. Distribution areas of species of H. sauzakense group and H. kopetdaghi.