Pseudocamelina glaucophylla (DC.) N.Busch

Esmailbegi, Shokouh, Lysak, Martin A., Rahiminejad, Mohammad Reza, Mirtadzadini, Mansour, Mummenhoff, Klaus & Al-Shehbaz, Ihsan A., 2017, A taxonomic revision of the genus Pseudocamelina (Brasssicaceae, tribe Thlaspideae), Phytotaxa 313 (1), pp. 117-129 : 123-125

Pseudocamelina glaucophylla (DC.) N.Busch in Grossheim (1927: 214). ≡ Nasturtium glaucophyllum de Candolle

(1821: 195). ≡ Cochlearia glaucophylla (DC.) Boissier (1842: 170) . Type:— IRAN (as Persia). Tehran: Tehran, Olivier

(holotype, P- 02272504!).

Cochlearia camelinae Boissier (1842: 172) , nom. illeg., non Cochlearia camelinae ( Fischer & Meyer 1835: 34) Ledebour (1841: 160) , in nota. ≡ Pseudocamelina camelinae N. Busch (1928: 113) . Described from: “[P. M. R. Aucher-Eloy] N. 331 et 4164, Persia.” Type:— IRAN. Without locality data, P. M. R. Aucher-Eloy 4164 (lectotype first-step designation by Rechinger (1968: 221) and second step by Miller (1978: 28): G-BOIS-00332266!; isolectotypes, BM-000582866, E-00381433!, G-00389747!, G-00389748!, K-000484395!, P- 02272509!, P- 02272510!, P- 02272511!). Syntype: [Iran], Persia, 1835, Aucher-Eloy 331 (P-05413238!).

Cochlearia violacea Boissier (1842: 171) . ≡ Pseudocamelina violacea (Boiss.) N. Busch (1928: 115) . Described from: [P. M. R. Aucher-Eloy] N. 91, Persia.” Type:—[ IRAN]. “Persia. Aucher-Eloy-Herbier d’Orient 91.” (lectotype first-step designation by Rechinger (1968: 222) and second-step by Miller (1978: 28): G-BOIS-00332267!; isolectotypes, G-00371872! [de Candolle herbarium], G-00371889!, P- 02272519!, P- 02272520!).

Description:—Herbs biennial, glabrous throughout or rarely pilose, glaucous. Stems simple or rarely few branched at base, (25 −) 30 − 70 cm tall, somewhat dichotomously branched above base, ascending, flexuous. Basal leaves rosulate, petiole 0.5 − 2.5(− 3) cm long; blade fleshy, orbicular to obovate or oblong to spatulate, (2 −)4 − 8.4 × 0.4 − 2.1 cm, entire or dentate; uppermost leaves linear to oblanceolate, entire. Racemes many flowered, lax, elongated considerably in fruit, flexuous; fruiting pedicels divaricate, filiform, straight, 4 − 19 mm long. Sepals oblong, 2.5 − 4(− 4.5) mm long, erect, lateral pair not saccate at base; petals spatulate, white, sometimes becoming purplish when dry, 3.5 − 4.5 × 1 − 1.8 mm, obscurely differentiated into blade and claw; filaments erect, 2 − 3.2 mm long, tetradynamous; anthers oblong, ca. 1 mm long; ovules 10 − 20 per ovary Fruit linear or oblong, (3 −)5 − 18(− 25) × 1 − 1.5 mm; style 0.4 − 1 mm long. Seeds uniseriate to biseriate, pale brown, smooth, oblong, 1 − 1.5 × 0.6 − 0.8 mm. 2 n =14.

Distribution:—endemic to Iran at 500 − 3900 m.

Representative specimens:— IRAN. Without exact locality P. M. R. Aucher-Eloy 90 (BM, P), 4137 (BM, P).

Azarbaijan: 13 km below Mianeh on road to Zanjan , 1050 m, 22 June 1971, J. Lamond 4231 ( E) ; Qezel Owzan (Kizil Uzun), 1200 m, 22 June 1971, K. H. Rechinger 42234 ( G, W) ; Einal Zeinal , between Tabriz and Talkheh Rud (Atschi Tschai), 1700 m, 20 July 1971, K. H. Rechinger 43548 ( G, K, W) ; N of Tabriz , 1700 m, 20 July 1971, J. Lamond 5177 ( E) ; Mianeh , Ghaflankuh , Zanjan road, 1150 m, 21 July 1974, H. Foroughi & M. Assadi 13714 ( W) ; Kivi , 90 km ENE Mianeh versus khalkhal, 1400 m, 15 July 1971, K. H. Rechinger 43325 ( W) ; SE of Ardabil, east of Hir , Darband region, 1921 m, 17 June 2015, M. Mirtadzadini et al. 1806 ( MIR) ; Takab , 2.5 km W of Takht-e Solaiman temple, 2156 m, 19 June 2015, M. Mirtadzadini et al. 1812 ( MIR) ; Hashtjin to Khalkhal , W of Aq-Dagh Mt. , after Sousahab village, near Mir-e Jin village, 2027 m, 7 August 2012, M. Mirtadzadini 1813 ( MIR). Chaharmahal Bakhtiari: Bakhtiari, 40 km S. Gandoman, H. Pabot 2272 ( G) ; Bakhtiari ( Tang-e-Sayad protected region), Marghak, 2550 m, 3 June 1974, K. H. Rechinger 47239 ( G, K, W) ; 21 km from Gandoman to Lordegan, 2210 m, 22 June 2015, M. Mirtadzadini 1756 ( MIR) ; Shahr-e Kord to Farsan , 2 July 2004, M. Mirtadzadini 1800 ( MIR). Fars : Abadeh , Shahrmian Kushk-e Zar , Dareh-e Gol , 2550 − 3000 m, 4 June 1975, P. Wendelbo & H. Foroughi 17859 ( E). Hamadan: Hamadan, Kuh-e Khorzane, Sabeti 136 ( W). Isfahan : S of Isfahan, Semirom waterfall, 1 Jul 2006, M. Mirtadzadini s.n. ( MIR) ; between Isfahan and Tehran , May 1859, A. G. Bunge s.n. ( P) ; between Yezd and Isfahan , May 1859, A. G. Bunge 13 ( P) ; Belehsuh-Kuh , between Dameneh and Khunsar, 2770 m, 10 August 1966, J. C. Archibald 3083 ( E, W) ; Qashqai , between Shahreza and Semirom, 2600 m, 5 June 1974, K. H. Rechinger 47361 ( E, W) ; Qashqai , Darreh Abshar, Semirom, 6 June 1974, K. H. Rechinger 47475 ( G, W) ; Ostan 10, between Isfahan and Abadeh, 1500 − 2500 m, 27 April 1956, F. Schmid 5420, 5382 ( G) ; Isfahan , Balehsun-Kuh, between Damaneh and Khunsar, J. C. Archibald 3083 ( W) ; Qashqai , Kuh-e Surmandeh (Kuh-e Alijuq) N of Semirom, 2700 − 3900 m, 7 June 1974, K. H. Rechinger 47500 ( W) ; W mountain of Baq-Bahadoran, 7 June 2001, M. Mirtadzadini 1798 ( MIR) ; between Tiran and Damane, S of Kord-e Olia, 2450 − 2600 m, 26 May 2010, M. Mirtadzadini 1799 ( MIR) ; Varian , near Ardestan, 2000 m, 25 June 1975, H. Foroughi & M. Assadi 18052 ( E) ; Semirom , Vanak , Dalan Kuh, 1900 m, 5 June 2001, M. R. Parishani 14163 (Isfahan) ; Isfahan , 1837, Aucher-Eloy 90 ( G). Kashan : Niasar to Delijan , Barzok village to Mt. Karkas , 23 May 2010, M. Mirtadzadini 1805 ( MIR). Kermanshah : 19 km N of Paveh , H. Pabot 1255 ( G). Kurdistan : Bijar to Zanjan, 5 km up to Bianlo village, 1650 m, 7 October 2002, H. Maroofi & S. Naseri 5574 (Sanandaj). Luristan: Durud, 2133 m, 28 May 1941, W. N. Koelz 17865 ( E, MICH, W) ; Oshtoran Kuh , 16 − 17 June 1974, J. Renz 48204 ( W) ; 33 km from Shul- Abad to Aligoodarz, 2760 m, 30 July 1999, M. Mirtadzadini 1803 ( MIR) ; 12 km from Aligodarz to Khomein Mt , 2600 − 2850 m, 3 July 1977, Runemark & Lazari 26557 ( E). Markazi: 4 km W Arak, 1950 m, 7 June 1959, H. Pabot 1152 ( G) ; E of Arak , 1645 m, 3 May 1966, J. C. Archibald 1735 ( E, K) ; 19 km N of Saveh , 1370 m, 22 May 1961, H. Pabot 7255 ( G) ; Sefid Khani Mt. , 10 July 1966, Anonymous 12441 ( TUH). Mazandaran: Lar Valley , 2450 m, 3 July 1974, P. Wendelbo & M. Assadi 13415 ( E, W) ; Haraz valley , above Panjab in dry slope, 1000 m, 11 May 1959, P. Wendelbo 638 ( E, W) ; Alborz , near Elika, 2700 m, Gauba 152 ( W) ; Alborz , Gochisar , NE Karaj, 2133 m, 18 June 1962, P. Furse 2636 ( E, W) ; Kudjur , near Elika, Gauba 135 ( W) ; Callus Valley , 2400 m, 9 June 1937, K. H. Rechinger 960 ( BM, W) ; Nasan to Yoush, 22 July 2015, M. Mirtadzadini 1791 ( MIR) ; between Yush and Nasan, 21 Hune 2011, M. Mirtadzadini 1792a ( MIR) ; Heraz road, near Ab-Askar (near Sad Lake), 12 August 2008, M. Mirtadzadini 1793 ( MIR) ; Karaj-Chalus road, Pol-e Zanguleh Kalavanga, Dunassi, 2900 m, 8 July 1999, H. Nazarian 33243 ( TUH) ; Gochisar , NE of Karaj, Furse 3636 ( E, W) ; Kendovan , Gachsar, 2980 m, 17 July 1972, S. C. Klein 3734, 3712 ( E, G) ; Kendovan , Gachsar, a proximite de l’ancienne route du col, 2980 m, 17 July 1972, S. C. Klein 3726 ( G) ; N of Kandovan Pass , 3110 m, 25 June 1966, J. C. Archibald 2481 ( E, W) ; Kandovan , 22 June 1966, Anonymous 12556 ( TUH) ; N of Karaj , S of Kandovan tunnel, 2691 m, 2 August 2012, M. Mirtadzadini 1790b ( MIR) ; Chalus road, W of Kandovan tunnel, 2887 m, 25 June 2015, M. Mirtadzadini 1807 ( MIR) ; Alborz mountain chain, Chalus road, after Karaj dam, Mowrud village, 2340 m, 21 July 2010, M. Mirtadzadini 1808 ( MIR) ; Karaj to Chalus, N of Nesa village, 2229 m, 13 June 2015, M. Mirtadzadini 1825 ( MIR) ; Karaj to Chalus, S of Gatshsar, W of Talaqan crossroad, 2251 m, 13 June 2015, M. Mirtadzadini 1826 ( MIR) ; Nesa in Chalus road, 7 May 1984, A. Ghahreman & Sheikhaleslami 12444 ( TUH) ; Tochal Mt. , 2200 m, 7 June 1972, R. Alava 10385 ( E). Qazvin : Vessieh, 1500 m, 1 September 1948, K. H. Rechinger 6851 ( E, G, MO, W) ; between Qazvin and Rasht, 500 − 1200 m, 5 − 6 September 1956, F. Schmid 6586 ( E, G, W). Semnan: Kavir, Siah Kuh, 5 − 35 km a Karavan − Sari Shah Abbas, 24 May 1974, K. H. Rechinger 46293 ( E, G, MO, W). Tehran : Mt. Alborz near Derbend, 8 June 1843, T. Kotschy 259 ( BM, E, G, MO, P, W) ; Hawuz-e Sultan , 1868, C. Haussknecht s.n. ( BM, P, W) ; 1 km N of Abeali , 2040 m, 11 July 1960, H. Pabot 1290 ( G) ; Tochal , Ferasad, 1800 m, 28 May 1902, J. F. N. & A. Bornmüller 6182 ( E, P) ; Tehran , P. M. R. Aucher-Eloy 4163 ( BM, P) ; Tochal , Scheheristanek, 2200 m, 6 June 1902, J. F. N. Bornmüller 6274 ( P) ; between Tudischk and Ku-pahi , 11 May 1889, A. G. Bunge s.n. ( P) ; Dih-korghon , 1856, Seidlitz s.n., ( P) ; Mt. Tochal , Hafthauz, 1300 − 1500 m, 4 July 1948, P. Aellen 1079 ( G, MO, W) ; 20 km SE of Tehran, 1050 m, 13 June 1962, H. Pabot 7371 ( G, W) ; Au N-O du village de Kelvan , S of Naz (Mishenou), 2860 m, 21 June 1971, S. C. Klein 1204, 1215, 1043, 1066, 1075 ( G) ; Tochal , 2990 m, 16 June 1973, S. C. Klein 4613 ( G) ; Imam Zadeh Hashem , Polour, 2500 m, 30 June 1974, S. C. Klein 7637 ( G) ; 20 km SE of Tehran , H. Pabot 7371 ( G) ; Tochal , near Scheheristanek, 2200 m, 7 June 1902, J. F. N. Bornmüller 6183 ( BM, G, P, W) ; between Shemshak and Dizin, 2900 m, 25 June 1977, K. H. Rechinger 57215 ( E, K, G, W) ; Vashan , Kuh-e Lazar, F. Termé & Matine 31792e ( E) ; Bijin , about 44 km from Tehran on road to Qom, 1050 m, 20 May 1974, A. Shirdelpur & H. Amini 11613 ( W) ; 157 km from Hamadan on road to Ghazvin , 1890 m, 19 July 1974, M. Assadi & H. Amini 13630 ( E, W) ; Road of Qom , 900 m, 2 May 1977, H. Amin & Rejamand 32867 ( W) ; 30 km N of Karaj ( K. 32 km W of Tehran), 2 July 1969, J. S. Anderson & I. C. Petersen 151 ( E, K, W) ; Karaj , Kuh-e Dasht, 7 May 1937, K. H. Rechinger 372, 766, 977 ( W) ; Alborz , Tochal, 2600 m, 30 May 1902, J. F. N. Bornmüller 6184 ( PRC) ; Tehran , Darake, 16 June 2000, M. Mirtadzadini 1794 ( MIR) ; Tochal , 2000 m, 19 June 2015, S. Esmailbegi 1795, 1796, 1979 ( MIR) ; Darband , 1971 m, 18 June 2015, S. Esmailbegi 1802, 1804 ( MIR) ; N of Karaj dam, 1803 m, 26 June 2015, M. Mirtadzadini 1814, 1815 ( MIR) ; 30 km E of Demavand, on the pass SE of Taar lake , 3046 m, 12 July 2015, M. Mirtadzadini 1816 ( MIR) ; Emamzadeh Davoud Mt. , 2750 m, 25 June 2015, M. Mirtadzadini 1817, 1818 ( MIR) ; N of Dizin Pass , 3349 m, 12 June 2015, M. Mirtadzadini 1820 ( MIR) ; 17 km NE of Karaj, W of Arangeh village , 1915 m, 24 June 2015, M. Mirtadzadini 1821 ( MIR) ; 15 km NE of Karaj near Arangeh , 1696 m, 24 June 2015, M. Mirtadzadini 1822 ( MIR) ; Dizin , W of Gajereh, 3592 m, 3 August 2012, M. Mirtadzadini 1823 ( MIR) ; Dizin Pass , near ski run, 3130 m, 3 August 2012, M. Mirtadzadini 1824 ( MIR) ; road of Qom 50 km from Tehran, Abdol-Abad, 19 June 1967, S. Mobayen 12442 ( TUH) ; N of Tehran, Tochal Mt., 2000 m, 19 June 2015, S. Esmailbegi 1810, 1811 ( MIR) ; Damavand , Anonymous 12443 ( TUH) ; Taleghan , Mehran, 2050 m, 6 June 1991, Sanei & A. A. R. Maasoumi 100392 ( TUH) ; above Darband and Mt. Kuhha-ye Tochal, 2000 − 3800 m, 29 June 1973, J. Sojak 7278, 7265 ( PR). Yazd: Yazd, Deh-Bala, Shirkuh mountain, 3700 − 4000 m, 21 June 1975, H. Foroughi & M. Assadi 17969 ( E, G). Zanjan: 4 km from Mahneshan to Aqkand, 1792 m, 19 June 2015, M. Mirtadzadini et al. 1809 ( MIR) .

Notes:— Pseudocamelina glaucophylla is by far the most widespread and variable species of the genus, a conclusion reached earlier by Miller (1978). The variation is evident in every morphological feature, including plant height, size, shape, and margin of basal leaves, branching of the inflorescence, length of fruiting pedicles, flower size, ovule number, fruit size and shape, and seed arrangement. The longest fruit in the species (2 − 3 cm × ca. 1 mm) were observed in plants of Alborz Mountain, especially in Province Mazandaran. By contrast, the shortest and widest fruit (3 − 8 × 1.4 − 1.8 mm) are scattered throughout the species range and typically biseriate seeds were found in Rechinger 46293 (K, W) that was collected from central Iran, Kavir protected region, Siah-Kuh.

Forms of Pseudocamelina glaucophylla with straight rachis and pedicels shorter than fruit were recognized by Boissier (1867) as Cochlearia violacea (petals purple) and C. camelinae (petals white). However, the examination of a wealth of material of this “three-species” complex, both in the field and in many herbaria, revealed that fruit shape, pedicel length, presence or absence of indumentum, flower color when dry, length ratio of fruiting pedicel to fruit are definitely unreliable, and Miller’s (1978) treatment of the three taxa as a single polymorphic species is fully justified. Indeed, these characters are not inherited in a single, consistent pattern to justify the recognition of additional taxa.

Rechinger (1968) did not specify which of the three duplicates in the Geneva herbaria is the lectotype of Pseudocamelina violacea . Boissier annotated P- 02272519 and another sheet that he did not cite (P-55348479), but the collection number and year of collection of the latter sheet are 90 and 1835, respectively.