Rhaponticoides schmidii (Wagenitz) Negaresh, 2015

Negaresh, Kazem, Khoshroo, Sayed Mohammad Reza, Karamian, Roya & Joharchi, Mohammad Reza, 2015, A revision of Rhaponticoides (Asteraceae, Cardueae-Centaureinae) from Iran, Phytotaxa 213 (2), pp. 87-101 : 92-99

2. Rhaponticoides schmidii (Wagenitz) Negaresh , comb. nov. ( Figs. 4–6 )

Basionym: Centaurea schmidii Wagenitz (1980: 330) . Type:— IRAN. Prov. Khorasan: Bar village, in stepposis montanis, 1780–2530 m, 28 June 1956, Schmid 6291 (holotype W!, isotypes E!, G!).

Plants perennial, herbaceous, arising from very thick rhizomes up to 3 cm in diam., green throughout, 70–100 cm tall, densely covered with white lanate pubescence at base, other parts glabrous; remains of stems of the previous year present. Stems several, 5 to 30(–40), sometimes single, erect or ascending, usually branched in upper part, sometimes producing fairly small sterile capitula, relatively sturdy, sulcate, densely leafy in lower part, other parts of stem aphyllous, 4–5(–6) mm in diam. at base. Leaves firm, pinnate and distinctly serrate. Basal and lower cauline leaves petiolate, lanceolate in outline, 17–30 cm long, pinnatisect, segments in 6–8(–9) pairs; terminal segments wider, ovate, 2–4 × (1.5–) 3–5 cm; lateral segments oblong or lanceolate, (1–)3–5 × (0.3–) 0.8–1.5 cm, decurrent along the rachis. Median cauline leaves subsessile, 13–20 cm long, pinnatisect, segments in ca. 8 pairs; segments equal in size, oblong, sometimes lanceolate, decurrent along the rachis. Upper cauline leaves increasingly reduced, sessile, lyrate, with 1–3 pairs of segments; terminal segment longer, narrowly oblong, denticulate, with a distinct mucro up to 2 mm long; lateral segment small, acute at apex, entire. Capitula few (2–3), solitary on short to long peduncles (3–15 cm long). Involucres oblong or subglobose, truncate at base, 25–28 × 25–30(–35) mm. Phyllaries multiseriate, coriaceous, yellowish-brown, becoming dark-brown apically, conspicuously longitudinally striate with brown-blackish lines, rounded at apex. Outer phyllaries broadly ovate, 3–7 × 3–6 mm; middle phyllaries ovate-oblong to oblong, 12–15 × 6–8 mm; inner phyllaries oblong, 13–20 × 2–3 mm. Appendages large, totally concealing the phyllaries, conspicuously longitudinally striate with brown-blackish nerves, membranous-scarious, orbicular, the inner ones broadly oblong, 8–10 mm long, 14–16 mm wide, strongly decurrent, denticulate, irregular lacerate or strongly undulate. Receptacle densely covered with long white bristles. Flowers yellow in vivo, sulphury in sicco; central flowers hermaphrodite, 35–40 mm long, distinctly 5-veined, anther tube pale yellow, exserted; peripheral flowers sterile, slightly radiant, with staminodes, 4–5-lobed, the lobes filiform. Achenes asymmetrically subcylindrical-truncate or oblong, sometimes oblong-undulate, compressed at margins, 9–11 mm long, 3–4 mm wide, brown, dark brown to blackish, rarely yellowish and shiny, transversally wrinkled in the upper part, smooth towards base, glabrous; insertion areole large, ca. 2 mm long, yellowish. Pappus multiseriate, persistent, scabrous, brownish to blackish, 5–7 mm long; inner pappus with unequal bristles: adaxial bristles 7–10 long, abaxial bristles 4–6 mm long.

Taxonomic and distributional remarks: — Rhaponticoides schmidii is a rare endemic of the vicinities of Bar village in Khorasan Province, NE Iran ( Fig. 3 ). It is the only species of the genus Rhaponticoides with that special pappus type (inner pappus with unequal bristles: adaxial bristles 7–10 mm long, abaxial bristles 4–6 mm long). By this morphological character, R. schmidii is similar to members of the genera Mantisalca Cassini (1818: 142) and Oligochaeta ( Candolle 1838: 671) Koch (1843: 42) , but differs from them by many characters. Rhaponticoides schmidii is also the only species of R. sect. Iranicae with the pappus shorter than achenes (vs. pappus as long as achenes). It grows in steppes, rocky or stony slopes, dry rubble, on lake margins, scattered between Rosmarinus and Amygdalus shrubs, in montane zones, at elevations of 1750–2600 m ( Fig. 5 ). It closely resembles R. lachnopus in its habit.

This species is also related to R. jerhardii ( Agababian 1997: 897) Ranjbar & Negaresh (2014a: 14) , especially because of the indumentum, leaf dissection and color of flowers. However, it differs from R. jerhardii in its stem being densely leafy in the lower part, upper parts without leaves (vs. the stem being foliate loosely throughout), terminal segments of leaves wider than lateral ones (vs. terminal segments as large as lateral ones), and also inner pappus shorter than outer, inner pappus with unequal bristles: adaxial bristles 7–10 mm long, abaxial bristles 4–6 mm long (vs. inner pappus as long as the outer). Rhaponticoides schmidii grows at higher elevations than those of R. jerhardii .

Chromosome numbers: —Unknown.

Selected specimens examined:— IRAN. Prov. Khorasan: NW Neyshabur, Bar village , 1800 m, 25 June 2002, Joharchi & Nasseh 34279 ( FUMH!, duplicate BASU!) ; NW Neyshabur, Bar village , 2000 m, 29 June 2006, Zangui 34877 ( FUMH!, duplicate BASU!) ; NW Neyshabur, W mountains of the Bar village toward waterfall, 36° 31′ 2.01″ N, 58° 46′ 3.43″ E, 1920 m, 9 June 2013, Joharchi 45068 ( FUMH!, duplicate herb. Negaresh!: two sheets) .

II. Rhaponticoides sect. Ruthenicae (Dobrocz.) Negaresh , comb. & stat. nov.

Basionym: Centaurea subgen. Centaurea ser. Ruthenicae Dobrocz. in Dobroczajeva & Kotov (1962: 43). Centaurea subgen. Centaurea sect. Centaurea subsect. Ruthenicae Agababian (1997: 894) . Type:— Centaurea ruthenica Lam.

Plants densely covered with white lanate pubescence at base, other parts glabrous. Leaves entirely divided, pinnatipartite, margin regular denticulate or cartilaginous-denticulate, sometimes entire. Involucres medium-sized, 15–20(–25) × 10–20 mm, ovoid or ovoid-oblong, rarely cupuliform. Phyllaries broadly ovoid to suborbicular, without distinct appendages, or with very narrow scarious margins up to 1(–2.5) mm wide, margin entire to slightly undulate. Flowers yellow, peripheral radiant, staminodes indistinct. Achenes 6–8 mm long; pappus usually shorter than achenes, (4–)6–13(–16) mm long, inner bristles usually strongly shorter than outer ones.

Taxonomic and distributional remarks: — Rhaponticoides sect. Ruthenicae is the largest section of Rhaponticoides and comprises 15 species. It is also the most widespread section of the genus, ranging from SE Spain to S Europe, E Turkey to Magnolia and NW China. Rhaponticoides sect. Ruthenicae is the only section whose members are present in the Eurasian-Circumboreal region. Altogether, members of Rhaponticoides sect. Ruthenicae belong to Eurasian-Circumboreal, Mediterranean and Irano-Turanian elements.

3. Rhaponticoides bachtiarica (Boiss. & Hausskn.) Martins (2007: 43) . Serratula bachtiarica Boissier & Haussknecht in Boissier (1875: 587). Centaurea bachtiarica (Boiss. & Hausskn.) Hayek & Bornmüller in Hayek (1926: 272). Type:— IRAN. Prov. Chahar Mahal and Bakhtiari: Tang-e Mahmuod, in declivibus dumetosis et in planitie Ardell, Bachtiari, July 1868, Haussknecht s.n. (holotype G!, isotypes JE!: two sheets, W!). Fig. 7 .

Plants perennial, herbaceous, glabrous, green-brownish throughout, 80–100 cm tall. Stems erect, simple or branched above (with a few short branches), sulcate, densely leafy in lower to median parts. Leaves coriaceous, pinnate. Basal and lower cauline leaves long-petiolate, pinnatipartite, oblong in outline; segments in 7–8 pairs, 4.5–7 × 0.5–2.2 cm, lanceolate, oblong or lanceolate-oblong, basally asymmetric, decurrent along the rachis, margin usually entire, sometimes remotely inconspicuously dentate, acute at apex.Median cauline leaves subsessile, 15–17 cm long, lanceolate in outline; segments in 6–7 pairs, 1.5–6(–7) × 0.2–1 cm, sometimes terminal segment longer, oblong, sometimes ± retrorse, decurrent along the rachis, margin entire, acuminate at apex. Upper cauline leaves sessile, simple, undivided, 6–8 × 0.7–0.9 cm, lanceolate, narrowed toward base, entire. Capitula few, 2-cephalous, solitary on long peduncles (up to 15 cm long). Involucres cupuliform, truncate at base, ca. 22 × 15–16 mm. Phyllaries multiseriate, imbricate, appressed, glabrous, yellowish-green, longitudinally striate, obtuse, with narrow scarious pellucid entire margin (1–2.5 mm wide). Outer phyllaries suborbicular, 2–5 × 2–6 mm; middle phyllaries broadly oblong or oblong-orbicular, 10–12 × 6–8 mm; inner phyllaries oblong or narrowly oblong, 14–16 × 3–5 mm; scarious margin lacerate. Receptacle densely covered with long white bristles. Flowers pale yellow. Achenes immature; pappus double, persistent, whitish, outer ca. 6 mm long, inner shorter.

Taxonomic and distributional remarks: — Rhaponticoides bachtiarica is a very rare endemic to Chahar Mahal and Bakhtiari Province, S Iran ( Fig. 3 ). It is only known from the type locality (Tang-e Mahmoud) and probably grows in steppes of montane zones. Rhaponticoides bachtiarica is similar to R. ruthenica in its indumentum, dissection of leaves and color of flowers. However, it differs from R. ruthenica by its basal and lower cauline leaves with 7–8 pairs of segments, margin usually entire, sometimes remotely inconspicuously dentate (vs. segments in 12–15 pairs, margin regularly denticulate or cartilaginous-denticulate), capitula few, 2-cephalous (vs. several, 2–7-cephalous), involucres cupuliform (vs. ovoid), appendages of phyllaries 1–2.5 mm (vs. 0.5 mm) wide, and also pappus ca. 6 mm (vs. 4–6 mm) long.

Chromosome numbers: —Unknown.

4. Rhaponticoides ruthenica (Lam.) M.V. Agab. & Greuter in Greuter (2003: 61). Centaurea ruthenica Lamarck (1785: 663) . Centaurium ruthenicum (Lam.) Koch (1851: 417) . Type:— AFGHANISTAN. Ghorat: Darrah-e Ghuk prope Puni, 2200 m, 6 June 1971, Podlech 21933 (KU! neotype designated here by Negaresh). Fig. 8 .

Centaurea schwarzenbergiana Schur (1855: 4) . Bielzia schwarzenbergiana (Schur) Schur (1886: 409) . Type:— ROMANIA. Cluj, Schur s.n. (JE! lectotype designated here by Negaresh, isolectotypes JE!, M!).

Plants perennial, herbaceous, green throughout, 40–150 cm tall, densely covered with white lanate pubescence at base, other parts glabrous. Stems solitary or few, erect, usually branched in upper part, rarely simple, sulcate, loosely leafy throughout, 5–6 mm in diam. at base. Leaves entirely divided, pinnatipartite, margin regular denticulate or cartilaginous-denticulate. Lower and median cauline leaves petiolate, segments in 12–15 pairs, linear or linearlanceolate, 3–7 × (0.1–) 0.3–0.9 cm, terminal segment equal with lateral ones and usually decurrent along the rachis, acuminate at apex. Upper cauline leaves increasingly reduced, sessile, segments in 5–6 pairs, linear to narrowly linear, terminal segment longer, 1–3 mm long, shallowly denticulate or entire. Capitula several, 2–7-cephalous, solitary on short to long peduncles (1–15 cm long). Involucres ovoid, truncate at base, 20–25 × 14–20 mm. Phyllaries multiseriate, coriaceous, yellowish-brown, becoming dark-brown apically, conspicuously longitudinally striate with green or brown lines, rounded at apex. Outer phyllaries broadly ovate, 2–6 × 3–6 mm, middle phyllaries ovate-oblong to oblong, 7–12 × 6–7.5 mm, both without a distinct appendage, or with a very narrow scarious margin ca. 0.5 mm wide; margin entire to slightly undulate. Inner phyllaries oblong, 13–20 × 2–3 mm, with brown scarious margins, oblong, lacerate or undulate. Receptacle densely covered with long white bristles. Flowers yellow in vivo, sulphury in sicco; central flowers hermaphrodite, 22–24 mm long, anther tube pale yellow, exserted; peripheral flowers sterile, radiant, with staminodes, 4–5-lobed, the lobes filiform. Achenes transversally wrinkled, 6–8 mm long. Pappus double, persistent, whitish or brownish; outer pappus 4–6 mm long, inner pappus strongly shorter, 1–2 mm long.

Taxonomic and distributional remarks: — Rhaponticoides ruthenica , even in narrow circumscription, is widely distributed in Afghanistan, Armenia, Belarus, Caucasus, China, Kazakhstan, Kyrgyzstan, Romania, NW Iran, NW Pakistan, SE Turkey, Tajikistan, Ukraine, Uzbekistan, W Siberia, Middle Asia and Central Europe ( Tzvelev 1963, Wagenitz 1980, Bădărău et al. 2001, Duran et al. 2014, Ranjbar & Negaresh 2014a). Rhaponticoides ruthenica is the only species of the genus that is distributed in both Mediterranean and Irano-Turanian regions. This species has the widest distribution range among the species of the genus. Rhaponticoides ruthenica grows in steppes, forest steppes and mesoxerophytic bushy submediterranean habitats from west Palaearctis ( Bădărău et al. 2001). In addition, it can be expected to grow on stony slopes, meadows, follow fields, outcrops of chalk and limestone, in mid-montane zone, at elevations of 1600–3300 m. In Iran, R. ruthenica closely resembles R. bachtiarica in its habit.

Chromosome numbers: — Tonian (1980), Agababian & Goukasian (1994) and Gaffari & Shahraki (2001) reported the same chromosome number 2 n = 2 x =30 for Centaurea ruthenica .

Selected specimens examined:— IRAN. Prov. Azerbaijan: Chalil Kuh , in faucibus NW Selvana, in rupium fissuris, 1750–2000 m, Rechinger 48895 ( W!) ; Ghogeh Dagh, W Bazaragn , 2100–2250 m, Lamond 4994 ( E!) ; W. Rechinger 43984 ( E!, W!) . AFGHANISTAN. Ghorat: Darrah-e Ghuk prope Puni , 2200 m, 6 June 1971, Podlech 21933 ( KU!) ; Wakhan: in valle Daryao Baroghil , 3200–3300 m, Anders 7850 ( W!) ; Bamian: Band-e Amir ad lacum Band-e Panir , 2800 m, Rechinger 18407-b ( W!) . KYRGYZSTAN. 17 August 1998, Robertson et al. 5851 ( E!) ; 25 August 1998, Robertson et al. 6003 ( E!) . TURKEY. B9 Ağrı: Patnos, Karakuyu village road, 1681 m, 8 August 2009, 39°20’040’’N, 42°44’521’’E, Duran & Doğan 8730 ( KNYA!) .

Figures

FIGURE 3. Distribution of Rhaponticoides bachtiarica (1), R. ruthenica (2), R. lachnopus (3), and R. schmidii (4) in Iran.

FIGURE 4. Rhaponticoides schmidii (Wagenitz) Negaresh (Schmid 6291, E). A. close up of involucre. B. habit. C. close up of achenes with pappus. Scale A = 1 cm, B = 5 mm.

FIGURE 5. Rhaponticoides schmidii (Wagenitz) Negaresh (Joharchi & Nasseh 34279, FUMH). A. inflorescence. B. habit and habitat. Scale A = 10 cm, B = 20 cm.

FIGURE 6. Rhaponticoides schmidii (Wagenitz) Negaresh (Joharchi & Nasseh 34279, FUMH). A. capitule (focused on several dark brownish nerves). B. capitule with two pollinators. C. focused on median and lower cauline leaves with serrate margins. Scale A = 1 cm, C = 2 cm.

FIGURE 7. Rhaponticoides bachtiarica (Boiss. & Hausskn.) Martins (Haussknecht s.n., W).

FIGURE 8. Rhaponticoides ruthenica (Lam.) M. V.Agab. & Greuter (Podlech 21933, KU). Scale A = 2 cm.