Reichardia macrophylla (Vis. & Pančić) Pančić, Fl. Serbiae, 1874

Conti, Fabio, Niketić, Marjan, Vukojičić, Snežana, Siljak-Yakovlev, Sonja, Barina, Zoltán & Lakušić, Dmitar, 2015, A new species of Reichardia (Asteraceae, Cichorieae) from Albania and reevaluation of R. macrophylla, Phytotaxa 236 (2), pp. 121-134 : 126-132

Reichardia macrophylla (Vis. & Pančić) Pančić, Fl. Serbiae : 460 (1874). Figs. 5 , 7 .

≡ Picridium macrophyllum Visiani & Pančić (1870: 3 , tab. 16).

— Picridium macrogaranum Pančić ex Nyman (1879: 461) , nom. inval. (pro syn. of P. macrophyllum Vis. & Pančić ).

Type:—“ Picridium mokragoranum / Vis. & P / [ SERBIA.] Mokra gora in rupib[us], [1]866”, Pančić s.n. (lectotype designated here: BEOU 11147).

Note: —The species was first named as Picridium macrophyllum . Reichardia picroides (as P. crassifolium Willk. ) and the West Asian R. dichotoma (as P. dichotomum DC. ) were listed as the most similar species ( Visiani & Pančić 1870). On the second occasion Pančić (1874) gave the current name, R. macrophylla , but without indication of the basionym and direct citation of the original reference. Instead of that, the citation of original authors (“Vis. & Pančić”) can be considered as an indirect reference to the basionym and the name R. macrophylla can be treated as a valid new combination ( McNeill et al. 2012, Art. 41.3 and 41.4), homotypic with P. macrophyllum . In Pančić’ s time this plant was only known from the vicinity of Mokra Gora in western Serbia. The corresponding geographical name P. macrogaranum (copied from Pančić’ s label in error for “ P. mokragoranum ”) was subsequently mentioned in synonymy ( Nyman 1879).

It appears that after the discovery of the species Pančić firstly used the name P. mokragoranum as provisional. We found four specimens collected by Pančić before 1870 which could belong to original material used in the protologue ( BEOU 11147, BEOU 11148, BP 437534, PAD 0045565). Among them we have selected and designated as a lectotype the Pančić’ s specimen labelled as “ Picridium mokragoranum ” ( BEOU 11147). It should be noted that no specimens and additional names were cited in the protologue of P. macrophyllum ( Visiani & Pančić 1870) . However, several facts indicate the connection of this specimen with the original material: Pančić’ s handwriting, material collected in 1866 and additional diagnostic label with comparison of P. macrophyllum with P. crassifolium . Moreover, on another sheet of the same collection there is a label reading “ Picridium / lingulato & crassifolio / affine / Mokra gora“ ( P. lingulatum Boissier 1841: 390 , nom. inval., also refers to R. picroides ). The specific epithet ( “ macrophyllum Vis. & P.”) and the new generic placement (“ Reichardia ”) were additionally written later by Pančić on the same label. This well preserved and recognizable material clearly points to the original description and illustration.

Taxonomic relationship:—Although the clear differentiation between R. macrophylla , R. picroides and R. dichotoma was presented in the protologue ( Visiani & Pančić 1870), R. macrophylla was neglected for a long time, and it was considered a synonym of R. picroides ( Sell 1976) or R. dichotoma ( Greuter 2006 +; Hand et al. 2009+).According to the latter authors R. dichotoma is known from two disjunct areas, one in the former Yugoslavia and the second in Asia Minor, Syria, Lebanon, Transcaucasia and Iran. However, many relevant Balkan botanists ( Siljak-Yakovlev 1981, 1986, 1996; Šilić 1984, 2006; Stevanović et al. 1995; Niketić 2014; Tomović et al. 2014) have supported the original identity of R. macrophylla as an endemic Balkan species, vicarious to Western Asiatic R. dichotoma .

Reichardia macrophylla differs from closely related R. dichotoma by a simple stem or synflorescence (not strongly divaricately branched), larger flowering capitula (3–5 cm vs. 2–3.5 cm in diameter), outer phyllaries never curved at the apex, relatively wider basal leaves in the upper part (oblong-ovate vs. linear-spathulate), etc. It also differs from R. picroides and R. albanica by larger capitula and leaves and longer achenes (4–5 mm vs. 3–4.5 mm). It never has papillose leaves, its ligules are not striped dorsally like those in R. picroides , and their colour is bright yellow or sulphur-coloured (vs. light yellow in R. albanica ), and also its outer phyllaries are ovate, etc.

The aforementioned morphology of leaves refers to the typical populations from Serbia and eastern Bosnia. In the southwestern part of the species range ( Herzegovina, Montenegro) leaves are more xeromorphic as in R. dichotoma and R. albanica (linear-spathulate, coriaceous, with sinuate and spinulose margin). These two groups of populations may be referred to two distinct infraspecific taxa.

Karyology:— A comparative analysis of karyotypes of the species R. picroides and R. macrophylla has shown significant karyotypic differences ( Siljak-Yakovlev 1981). From the basic chromosome number x = 7 (2 n =14) for R. picroides and x = 9 (2 n =18) for R. macrophylla , it can be seen that they are two completely different species. At the same time, although R. macrophylla and R. dichotoma have the same chromosome number (2 n =18), the karyotypes of the two species are well differentiated. The chromosome formula are 2 n =18=12 metacentric chromosomes (2 with intercalary secondary constrictions— SC) + 6 submetacentrics (4 satellite chromosomes—sat) for R. dichotoma and 2 n = 18 = 8 metacentrics (2 sat) + 8 submetacentrics + 2 subtelocentrics (2 sat) for R. macrophylla (for more of detail see Siljak-Yakovlev 1996). In addition, Giemsa C-banding data revealed a different distributional pattern of constitutive heterochromatin on chromosomes of each species (6 bands for R. dichotoma and 18 for R. macrophylla ) and therefore confirmed the strong differentiation between two species also at the chromosomal level ( Siljak-Yakovlev 1986, 1996). The karyotype of R. albanica is closer to that of R. dichotoma because the two species possess the same chromosome formula (6 metacentric and 3 submetacentric chromosome pairs).

The genome size (2 C DNA value) assessed by flow cytometry also showed the certain difference from R. dichotoma and the other two taxa: 5.25 pg for R. macrophylla from Bosnia and Herzegovina ( Siljak-Yakovlev et al. 2010) and 5.54 pg for R. dichotoma from Lebanon (BouDagher-Kharrat et al. 2013) and 5.22 pg for R. albanica (present work). According to the classification of endemics based on karyological characters ( Favarger & Siljak-Yakovlev 1986; Siljak-Yakovlev & Peruzzi 2012) these three species, with same diploid chromosome number, belong to the category of paleo-schizoendemics.

Distribution and habitat:— Reichardia . macrophylla is endemic to the central part of Balkan Peninsula (east Dinarides), distributed in Bosnia and Herzegovina, Montenegro, western Serbia and northern Albania ( Fig. 6 ). Although its occurrence is reported from Croatia ( Hayek 1931, Greuter 2006 + as R. dichotoma, Hand et al. 2009 + as R. dichotoma ), we did not find any relevant data to confirm this. The possible place where this plant could be found in Croatia is foothills of Mt. Orjen, located near the tripoint of Montenegro, Bosnia and Herzegovina and Croatia. Greuter (2006 +) and Hand et al. (2009+) believe that the presence of R. dichotoma in Albania is doubtuful. Hayek (1924) reported R. macrophylla from Albania: “Auf felsigem Boden unter Kiefern und in Geröllhalden östlich ober Abata, zirka 1700 bis 1750 m ”. Jávorka (1926) also reported R. macrophylla from Albania (Hekurave); Schütt (in mscr.) reported it from Qafa e Pejes (also in the Albanian Alps) and recently, Meyer (2011) reported it from the same place and two additional places around Thethi. The record of Jávorka is vouchered (BP 437532, BP 437533), an additional specimen is collected by Jakucs (BP 589912), and together with our recent record (see below) we can confirm the occurrence of R. macrophylla in Albania.

Here it is important to note that Gallego et al. (1980) reported R. dichotoma for Albania on the basis of a poor specimen collected by G.E. Post in 1855 and kept in E (E 00480970). Its label states no country but only the locality “Resheny”. It is know from the database of Royal Botanic Garden of Edinburg (http://elmer.rbge.org.uk/bgbase/ vherb/bgbasevherb.php?cfg=bgbase/vherb/collectorfull.cfg&coll__id=GEPT&queryRow=53) that Post collected in Lebanon, Libya, Syria and Turkey. For this reason we consider the record of R. dichotoma from Albania by Gallego et al. (1980) as not reliable.

Populations of R. macrophylla are part of the vegetation of xerophilous rocky grounds (Festuco-Brometea), screes (Thlaspietea rotundifolii), subalpine grasslands (Elyno-Seslerietea) and open black pine ( Pinus nigra ) and Bosnian pine ( Pinus heldreichii ) woodland, on limestone and dolomites, at altitudes between 500–1750 m a.s.l.

New chorological data ( Fig. 6 ):

ALBANIA. Prokletije : Sesi [Thethi], foothill of Pejasi in the gorge [ MGRS 34 T CM99 ], limestone rocks, 22 August 1938, Černjavski s.n. ( BEO 27057 ) ; Prokletije : Sesi [Thethi], gorge in limestone blocks in sparse beech forest [ MGRS 34 T CM99 ], August 1938, Černjavski s.n. ( BEO 27058 ) ; District of Tropojë (Rrethi i Tropojës) ; above the valley of brooklet përroi i Lugu i Vranes, above village Peraj; in rocky beech forest, on limestone [ MGRS 34 T DM08 ], 42.33291 N 19.85645 E, 1394 m.s.m, 18 July 2014, Barina, Papp, Sárospataki 24089 ( BP) .

BOSNIA AND HERZEGOVINA. Sutjeska canyon [ MGRS 34 T CP01 ], 11 August 2005, Stevanović 20189 ( BEOU) ;

MONTENEGRO. Cetinje , Orlov krš [ MGRS 34 T CM29 ], 750 m. s.m, Jun 1908, Bierbach, s.n. ( P 03863464 ) ; Mt Durmitor : Boljske Grede [ MGRS 34 T CN37 , CN46 , CN47 ], 1750 m.s.m, rocky crevices, limestone, 25 August 1999, Niketić, s.n. ( BEO) ; Mt Durmitor : Sušica canyon, between Razmlin and Tara river [ MGRS 34 T CN38 ], limestone, 17 July 1994, Jovanović, Lakušić, Bulić, Vukojičić, Benić, Pavić 480/94 ( BEOU) ; Durmitor : Bavan—Ravni Bolj [ MGRS 34 T CN38 ], 1400 m.s.m, limestone, 27 July 1989, Lakušić, Stanić, Đorđević 549/89 ( BEOU) ; Mt Durmitor : Sušica canyon, between Sušica Lake and Tara river [ MGRS 34 T CN38 , CN39 ], Ostryo-Carpinion, limestone, 17 July 1994, Jovanović, Lakušić, Bulić, Vukojičić, Benić, Pavić 827/94 ( BEOU) ; Nikšić : Vilusi, Ilino brdo [ MGRS 34 T CN03 ], near the road, limestone, 25 July 2003, Niketić, Tomović 17653 ( BEOU) ;

SERBIA. Jadovnik: Žuta Stijena [MGRS 34T DN09], 43.25102 N 19.7271 E, 784.9 m.s.m, rocky places, Festuco-Brometea, limestone, 26 July 2012, Lakušić, Kuzmanović, Janković, Đurović 35024 (BEOU).

Figures

FIGURE 5. A–B. Reichardia albanica from locus classicus (A—habit, B—capitulum; 26 June 2011, photo D. Lakušić); C–D. R. macrophylla from locus classicus (C—habit, D—capitulum; 06 June 2014, photo M. Niketić).

FIGURE 6. Distribution of Reichardia albanica and R. macrophylla (UTM Grid zone 33 and 34T; dots correspond to basic MGRS squares 10 × 10 km).

FIGURE 7. Reichardia macrophylla (Vis. & Pančić) Pančić, lectotype.