Viola pseudaetolica Tomović, Melovski & Niketić, 2016

Tomović, Gordana, Niketić, Marjan, Lazarević, Maja & Melovski, Ljupčo, 2016, Taxonomic reassessment of Viola aetolica and Viola elegantula (V. sect. Melanium, Violaceae), with descriptions of two new species from the Balkan Peninsula, Phytotaxa 253 (4), pp. 237-265 : 242-255

Viola pseudaetolica Tomović, Melovski & Niketić , sp. nov. ( Figs. 5 & 6 a-c)

= V. saxatilis [ subsp. aetolica ] var. pseudolutea Hayek, Akad. Wiss. Wien, Math.-Naturwiss. Kl., Denkschr. 99: 134 (1924). Holotype (designated here):— ALBANIA. “Nord-Albanien / 1916 und 1918. / No. 13 / Viola alpestris ssp. aetolica (B.H.) / n. v. pseudosudetica Dfl. et Hay. / Distrikt Auf steinigen Wiesen des / Maranaj, nördl. von Skutari; ca. 1230 m. / 1.VI / [Legit] I. Dörfler” (WU! 2776).

– “ V. lutea ” sensu Rohlena, Sitzungsber. Königl. Böhm. Ges. Wiss. Prag : 28 (1904), non Hudson, Fl. Angl.: 331 (1762).

– “ V. alpestris subsp. aetolica ” sensu W. Becker, Beih. Bot. Centralbl. 26(2): 345 (1910), p.p., non (Boissier & Heldreich) W. Becker, loc. cit.

– “ V. saxatilis subsp. aetolica ” sensu Hayek, Denkschr. Kaiserl. Akad. Wiss., Wien. Math.-Naturwiss. Kl. 99: 133 (1924), non (Boissier & Heldreich) Hayek, loc. cit.; Hayek, Repert. Spec. Nov. Regni Veg. Beih. 30(1): 516 (1925) [Prodr. Fl. Penins. Balc. 1]; Rohlena, Preslia 20–21: 104 (1942).

– “ V. aetolica ” sensu Diklić , Flora SR Srbije, 3: 161 (1972), non Boissier & Heldreich in Boissier, Diagn. Pl. Orient. ser. 2, 6: 24 (1859); Erben, Mitt. Bot. Staatssamml. München 21(2): 565 (1985); Micevski, Flora na Republika Makedonija, 1(3): 531 (1995).

Type :— REPUBLIC OF MACEDONIA, Mt Jablanica , Kokalo, planinsko pasište na varovnička podloga, 1850 m, 15–Jul–2006, Melovski, Lj., Tomović, G., Niketić, M. (holotype: MKMEL! 07396; isotypes: BEOU! 17138; BEO! A26 ; MKNH; W!)

Typification: —An inspection of the Herbarium of Vienna University (WU) disclosed original material of V. saxatilis [ subsp. aetolica ] var. pseudolutea Hayek , collected by J. Dörfler 1916. from Mt. Maja e Maranajt in northern Albania (WU–Hal! 2776). August Hayek, as the author of this variety, initially wrote a different name on the label, “ V. alpestris ssp. aetolica (B.H.) / n. v. pseudosudetica Dfl. et Hay. ”. Although the final epithet from the label (“ pseudosudetica ”) does not mach the protologue ( “ pseudolutea ”) ( Hayek 1924), the author cites the same locality, date of gathering and the collecting number in the protologue, and also notes that the plant habitually resembles V. lutea subsp. sudetica (Willd.) W. Becker, Beih. Bot. Centralbl. 18(2): 388 (1905), which clearly shows that this is a holotype material of V. saxatilis var. pseudolutea . Although the epithet “ pseudolutea ” belongs to the oldest validly published name, in the case of raising to specific rank it will become an elder homonym of V. pseudolutea Schur, Enum. Pl. Transsilv. : 87 (1866), therefore it would not be acceptable as a valid for this species ( McNeill et al. 2012, Art. 38.1). Consequently, in this paper another name, V. pseudaetolica , is given, based on detailed description of the type from a more complete recent material (MKMEL, BEOU, BEO).

Description: —Usually short-lived perennial (rarely annual or biennial), (10)15–30(40) cm height, little- to muchbranched from base. Shoots 3–15(25) cm long, erect to ascending, sparsely puberulent to densely pubescent, green. Lower leaves with longer petioles than those from middle and upper; blade ovate to orbicular, obtuse or rounded, coarsely crenate, glabrescent or shortly pubescent along the veins and margin and sometimes on surface. Stipules of lower leaves small, oblong to oblong-spathulate, entire or pinnately divided, with 1–2 lateral segments on the both sides. Middle and upper leaves with shorter petioles; blade oblong-ovate to lanceolate or oblanceolate, about 1.5 times longer than stipules, gradually attenuated at the base into the petiole, obtuse or acute, crenate, glabrescent or shortly pubescent along the veins and margins and sometimes on surface. Stipules of middle and upper leaves pinnately divided; glabrescent or shortly pubescent, the undivided terminal part relative large, with 1–3 lateral segments on the inner and 2–7 on the outer side, tapering gradually downwards. Peduncles (4) 7–13.5 cm, glabrous or puberulent; bracts in upper third of peduncle. Flowers medium size 1.8–2.5 × 1–2.2 cm, bright yellow; lower surface of petals often becoming blue-violet at the end of flowering period. Calyx: upper sepals (6.4)8.6–9.9(12) × (2.3)3.1–4.2(5.2) mm; lateral (7.4)9.5–11(13) × (2.1)2.8–3.5(4.7) mm; lower (8.8)10.8–12.5(14.5) × (2.7)3.4–4.6(5.4) mm. Sepals ovate-lanceolate to ovate, acute, rarely obtuse, attenuate at base, puberulent to pubescent on back, sometimes becoming glabrous in fruit, ciliate at margin; margin with 1–3 conspicuous teeth on both sides, rarely entire. Appendages c. 1/4 as long as sepals, narrow to wide rectangular, trapezoid or semiorbicular, attenuate at base, coarsely dentate. Corolla: upper petals (10.2)12.3–14.3(16.1) × (8.3)10.3–12.4(15.7) mm, obovate to orbiculate; lateral (10.3)12.3–13.9(15.7) × (6.1)7.9–9.7(11) mm, ovate to irregularly ovate, papillose-hairy at the upper margins; lower without spur (10)11.4– 12.7(13.9) × (9.4)11.5–13(14.4) mm slightly triangular to cordate. Spur glabrous, straight to slightly curved upwards, greenish-yellow to greenish-violet, (6.7)7.5–8.5(9.8) × (1.7)1.9–2.1(2.4) mm; ratio of length/width: 3.8–4.3. Seeds ellipsoid, pale to dark brown. Flowering period: May–July.

Etymology: —The specific epithet is derived from the great similarity with V. aetolica .

Chromosome number: —2n = 16 (counted on plants from Mt Jablanica; Fig. 7a ).

Distribution: — V. pseudaetolica is distributed in the western parts of the Republic of Macedonia (Mt Bistra, Mt Dešat and Mt Jablanica), southern Serbia (Mt Koritnik, Mt Paštrik, Mt Mokra Gora and Mts Prokletije), northern Albania (Mt Mali e Đalices, Mt Mali e Shebenikut, Mt Maja e Nermajn, Mt Maja e Cukalit, Mt Maja e Maranajt and Gryka e Liqenit), Montenegro (Mt Rumija, Mt Lovćen, Mt Orjen and Mts Prokletije) and the southeastern parts of Bosnia and Herzegovina (surrounding of Trebinje). In Serbia, possible literature records of the presence of V. pseudaetolica (as V. aetolica ) for Mt Golija ( Nikolić et al. 1986, Gajić 1989) and Mt Zlatar ( Obratov 1992) have not been confirmed. The following literature records have to be treated as erroneous: Nikolić & Diklić (1958, as V. saxatilis subsp. aetolica for Mt Jablanik in nortwestern Serbia), Diklić (1958, as V. saxatilis subsp. aetolica for Mt Miroč in northeastern Serbia), Lakušić & Niketić (1988) and Krivošej (1997), as V. aetolica for Mts Šar Planina in southern Serbia). These records were confirmed by an inspection of plant material from BEO and BEOU herbarium collections. On the basis of the literature sources for Albania ( Markgraf 1931, Erben 1985, Meyer 2011) and Montenegro ( Rohlena 1942), as well as on the general distribution range, it is possible that V. pseudaetolica also grows in northern, eastern and central parts of Albania (Puka, Lura, Mt Korab, Mt Maja e Skenderbegut, Mt Maja e Dajtit, Mt Maja e Grope and Mt Guri i Topit) and in northwestern Montenegro (Mts Pivske Planine). In addition, possible presence of V. pseudaetolica is in Mt Velež in Bosnia and Herzegovina ( Beck 1918, as V. aetolica ) has not been confirmed due to lack of herbarium material from this locality ( Fig. 8 ).

Habitat and ecology: — V. pseudaetolica grows at the elevation from (870–)1000 to 2000(–2400) m. on carbonate or silicate geological substrates. It inhabits mountainous and high-mountainous pastures and rocky grounds of the vegetation classes Daphno–Festucetea, Festuco –Seslerietea and Juncetea trifidi. It can also be found in scree vegetation of the classis Drypetea spinosae and rarely at the edges of pine forests ( Pinus heldreichii or P. peuce ) or scrub vegetation of Pinus mugo or Juniperus sibirica ( Fig. 6d ).

Discussion: —Already Rohlena (1904, as V. lutea Huds. ) indirectly noticed that plants from one locality in Montenegro (Mt Balj in vicinity of Andrijevica) differ from V. aetolica in having denticulate sepals. This observation was supported by Hayek (1924) who, besides this character, pointed out that specimens of this taxon resembles to large individuals of V. lutea subsp. sudetica . However, according to the same author, the latter subspecies had not been found in the Balkan Peninsula. On the basis of these characters, Hayek described a new variety, V. saxatilis [ subsp. aetolica ] var. pseudolutea Hayek (1924: 134) , from the locality Mt Maja e Maranajt north from Skodra lake.

V. pseudaetolica differs from the closely related V. aetolica by the larger and bright yellow flowers, the relatively narrower and often denticulate sepals and a longer spur ( Table 2). On the basis of these characters V. pseudaetolica is also similar to V. elegantula and V. kopaonikensis ( Table 3).

V. aetolica is distributed in Greece (Epirus: Mts North and Sout Pindhos), East Central Greece, Sterea Ellas and Peloponnisos) ( Raus 1986), southern part of the Republic of Macedonia (Mt Galičica) ( Erben 1985, Micevski 1995, pro parte) and southeastern Albania (Mt Gramos), while literature records for Mt Mali i Nemereckes ( Meyer 2011) and Mt Ostrovices ( Erben 1985, Meyer 2011) remain questionable ( Fig. 8 ). Micevski (1995) considered that V. aetolica also grows in Mt Dautica, Mt Ljuben and Mt Stogovo. However, during inspection of the herbarium material (MKNH) from these mountains, some other species from the V. sect. Melanium were recognized. Although Delipavlov (1979) and Anchev (1992) considered that V. aetolica is distributed in Bulgaria (Mts. Central Stara Planina, Central and Western Rhodopes), an examination of the plant material from the herbarium collections SOA and SOM did not confirm these literature records.

Spatial differentiation among populations of V.pseudaetolica and V.aetolica corresponds to their phytogeographical regions. Although the main part of the V. pseudaetolica range is situated on the southeastern Dinarides its southern boundary extending to Mt Jablanica in the northern part of the Scardo–Pindic phytogeographic provinces. This mountain is also referred to as the southern boundary for some other endemics with similar distribution, such as Tanacetum larvatum (Griseb. ex Pant.) Hayek , Euphorbia montenegrina (Bald.) K. Malý ex Rohlena ( Niketić et al. 2014) and Pimpinella serbica (Vis.) Benth. & Hook.f. ex Drude (Niketić, M., Tomović, G. 2006, pers. obs.). These orophytes often grow in the zone of Pinus heldreichii or P. peuce , while further south the population of V. aetolica generally follows the Abies borisii-regis and A. cephalonica zones.

Additional specimens examined:— REPUBLIC OF MACEDONIA. Bistra, blizu s. Galičnik, pod Bardeš , [ MGRS 34 T DM70 ], pasište, 1500 m, 28.04.2001, Melovski, Lj. ( MKMEL! 07337, 07338, 07339, 07340, 07341) ; Bistra, Carevec , [ MGRS 34 T DM71 ], pasište, 1750 m, 28.04.2001, Melovski, Lj. ( MKMEL! 07354) ; Bistra, Carevec– Ivanova Krasta , [ MGRS 34 T DM71 ], pasture on the stony site, limestone, 1680 m, 17.05.2008, Melovski, Lj. ( MKMEL! 07353) ; Bistra, Frčec , [ MGRS 34 T DM70 ], pasište, 1600 m, 06.06.2003, Melovski, Lj. ( MKMEL! 07355) ; Bistra, pomeđu Frčec i Govedarnik, [ MGRS 34 T DM70 ], trevesto kamenito, pokraj pat, 1800 m, 30.05.2003, Melovski, Lj., Arsovska, S. ( MKMEL! 07352) ; Bistra, s. Galičnik, [ MGRS 34 T DM70 ], grassy site on stony ground, limestone, 1450 m, Anonymus ( MKMEL! 07342) ; Bistra, Korita, below Rusin , [ MGRS 34 T DM70 ], montane pasture on deep soil, limestone, 1580 m, 16.05.2013, Melovski, Lj. ( MKMEL! 07321) ; Bistra, s. Galičnik–Karšija, Kuculova Livada , [ MGRS 34 T DM70 ], livada, 1350 m, 05.05.2002, Melovski, Lj. ( MKMEL! 07343) ; Bistra, sproti s. Galičnik, Kuculova Livada , [ MGRS 34 T DM70 ], 1400 m, 01.05.1983, Melovski, Lj. ( MKMEL! 07344, 07345, 07346) ; Bistra, Rusin , [ MGRS 34 T DM70 ], pasište na kamenjar, varovnik, 1650 m, 01.05.2000, Melovski, Lj. ( MKMEL! 07325) ; Bistra, podnožjeto na Rusin brdo, [ MGRS 34 T DM70 ], livadi i pasišta, varovnik, 1550 m, 03.05.1983, Melovski, Lj. ( MKMEL! 07322, 07323, 07324) ; Bistra, Rusin , at the sheep-fold “Rusin”, [ MGRS 34 T DM70 ], subalpine pasture on deep soil, calcareous ground, 1700 m, 16.05.2013, Melovski, Lj. ( MKMEL! 07316, 07317, 07318, 07319, 07320) ; Bistra, Toni Voda , [ MGRS 34 T DM71 ], pasište, 1700 m, 16.05.2003, Melovski, Lj. ( MKMEL! 07350, 07351) ; Bistra, Toni Voda , [ MGRS 34 T DM71 ], pasište, 1720 m, 28.04.2001, Melovski, Lj. ( MKMEL! 07348) ; Bistra , Toni Voda, [ MGRS 34 T DM71 ], pasište na dlaboka počva, 1650 m, 30.05.2003, Melovski, Lj., Arsovska, S. ( MKMEL! 07349) ; Bistra, Veli Vrv, [ MGRS 34 T DM70 ], S slope, subalpine pasture on stony ground, limestone, 1700–1850 m, 27.06.2015, Melovski, Lj. ( MKMEL! 07326, 07327) ; Bistra, Venec, nad s. Rosoki , [ MGRS 34 T DM70 ], otvoreno trevesto vo bukova šuma, 1500 m, 16.05.2003, Melovski, Lj. ( MKMEL! 07356) ; Bistra, Veli Vrv, [ MGRS 34 T DM70 ], S slope, subalpine pasture on stony ground in juniper shrub, limestone, 1800 m, 27.06.2015, Melovski, Lj. ( MKMEL! 07328, 07329) ; Bistra, Veli Vrv –summit area, S slope, [ MGRS 34 T DM70 ], subalpine pasture on stony ground, limestone, 1820 m, 27.06.2015, Melovski, Lj. ( MKMEL! 07330, 07331) ; Bistra, Veli Vrv –summit area, S slope, [ MGRS 34 T DM70 ], subalpine pasture on stony ground, limestone, 27.06.2015, 1850–1900 m, Melovski, Lj. ( MKMEL! 07332, 07333, 07334, 07335, 07336) ; Debarsko, Krčin , nad s. Gorno Kosovrasti, [ MGRS 34 T DM60 ], po planinskite livadi, 1530 m, 19.07.1960, Micevski, K. s.n. ( MKNH!) ; Debarsko, Krčin , nad s. Gorno Kosovrasti, [ MGRS 34 T DM60 , po varovitite mesta, 1400 m, 19.07.1960, Micevski, K. s.n. ( MKNH!) ; Debarsko , Krčin, vo bukova šuma, [ MGRS 34 T DM60 ], 1550 m, 19.07.1960, Micevski, K. s.n. ( MKNH!) ; Debar, Krčin , [ MGRS 34 T DM60 ], po planinskite livadi vo pojasot na bukovata šuma, 23.06.1968, Micevski, K. s.n. ( MKNH!) ; m. Jablanica, 07.1921, Soška, Th. ( BEO! 08504) ; mt. Jablanica, od Crnog Kamena prema Šećernoj glavi, [ MGRS 34 T DL56 , DL66 ], livadska biljna grupacija stenovitog nagiba, 16.07.1939, Černjavski, P. ( BEO! 08355) ; mt. Jablanica, na putu Crni Kamen – Šećerna glava, probna površina, [ MGRS 34 T DL56 , DL66 ], zatvorena biljna grupacija na jako tamno crvenom tlu, 15.07.1939, Černjavski, P. ( BEO! 08354) ; mt. Jablanica , na putu za Šećernu Glavu, [ MGRS 34 T DL56 ], c. 2000 m, 15.07.1939, Černjavski, P. ( BEO! 08356) ; m. Jablanica , in pascuis alpinis, solo silic., 18.07.1921, Košanin, N. (No. 25) ( BEOU! 41114) ; m. Jablanica, silikat, 07.1921, Košanin, N. (No. 16) ( BEOU! 41115) ; m. Jablanica, in jugo, solo silic., 07.1921, Soška, Th. (No. 43) ( BEOU! 41368) ; Jablanica, Crni kamen, [ MGRS 34 T DL66 ], visokoplaninski pašnjaci, kamenjari i stene, krečnjak+silikat, 1600–1800 m, 18.07.2006, Niketić, M., Tomović, G., Melovski, Lj. 21572 ( BEOU!) s.n. ( BEO!) ; Jablanica, Strižek, [ MGRS 34 T DL67 ], visokoplaninski pašnjaci, kamenjari, stene i sipari, krečnjak, 1800–2200 m, 19.07.2006, Niketić, M., Tomović, G., Melovski, Lj. 21635 ( BEOU!) ; s.n. (BEO!); Jablanica, Gornja Belica , pokraj patot za „Krstec“, pod Čuma, [ MGRS 34 T DL66 ], 1400–1550 m, 06.08.1986, Micevski, K. s.n. ( MKNH!) ; Jablanica , nad s. Vevčani, [ MGRS 34 T DL66 ], po silikatnite karpi, c. 1800 m, 25.07.1963, Micevski, K. s.n. ( MKNH!) ; Jablanica, nad Vevčanska Lokva (Ezero), [ MGRS 34 T DL66 ], silikaten kamenjar, 2000 m, 08.06.2003, Melovski, Lj., Arsovska, S,. Šušlevska, M. ( MKMEL! 07365, 07387) ; Jablanica, među Podgorečko i Vevčansko Ezero, [ MGRS 34 T DL66 ], pasište, 1950 m, 08.06.2003, Melovski, Lj., Arsovska, S., Šušlevska, M., ( MKMEL! 07388, 07389) ; Jablanica, Čumin Vrv, [ MGRS 34 T DL56 ], pasište na varovnički kamenjar, 2050–2080 m, 12.07.2006, IDSB ’06 ( MKMEL! 07394) ; Jablanica , Čuma, [ MGRS 34 T DL66 ], varovnički kamenjar, 1750–1900 m, 15.06.2006, Melovski, Lj., Veličkovska, M. ( MKMEL! 07366) ; Jablanica , Tri Šiljka, [ MGRS 34 T DL65 ], at the edge of the beech forest, 1750–1800 m, 16.06.2013, Hristovski, S. ( MKMEL! 07393) ; Jablanica, above Labuniška Lokva (upper lake)– Ljapi , [ MGRS 34 T DL56 ], subalpine pasture, 1950 m, 21.06.2008, Melovski, Lj. ( MKMEL! 07390) ; Jablanica, s. Gorna Belica, [ MGRS 34 T DL66 ], varovnički kamenjar, 1500 m, Melovski, Lj., Arsovska, S., Šušlevska, M. ( MKMEL! 07386) ; Jablanica, pod Krstec (nad Dupka), [ MGRS 34 T DL66 ], erodirana silikatna podloga, 1700 m, 15.06.2006, Melovski, Lj., Veličkovska, M. ( MKMEL! 07363, 07364) ; Jablanica, pod Vevčanska Lokva (Ezero), [ MGRS 34 T DL66 ], pasište na silikatna podloga, 1900–1980 m, 08.06.2003, Melovski, Lj., Arsovska, S., Šušlevska, M. ( MKMEL! 07384) ; Jablanica , Krstec, [ MGRS 34 T DL56 ], red stony ground, 1850 m, 17.06.2009, Hristovski, S. ( MKMEL! 07385) ; SERBIA. Bertisci, na Babinom polju, [ MGRS 34 T DN21 ], nekošene livade, 19.07.1933, Černjavski, P., Rudski, I., Lindtner, V. ( BEO! 08450) ; Bertisci, m. Bogićevica, Babino polje, [ MGRS 34 T DN21 ], u smrčevoj šumi, 19.07.1933, Černjavski, P., Rudski, I., Lindtner, V. ( BEO! 08472) ; Bertisci, m. Bogićevica, blizu žandarmerijske stanice na Dobroškoj planini, [ MGRS 34 T DN21 ], 18.07.1933, Černjavski, P., Rudski, I., Lindtner, V. ( BEO! 08474) ; Bertisci, m. Derviš Kom, [ MGRS 34 T DN21 ], na morenama od granitnih blokova, 17.07.1933, Černjavski, P., Rudski, I., Lindtner, V. ( BEO! 08501) ; Bertisci, mt. Nedžinat , [ MGRS 34 T DN22 ], na krečnjačkim stenama u cirku i pod vrhom u subalpskom regionu, 28.07.1933, Černjavski, P., Rudski, I., Lindtner, V. ( BEO! 08475) ; Bertisci, mt. Nedžinat , [ MGRS 34 T DN22 ], na krečnjačkim stenama u borovoj šumi blizu subalpskih livada, 28.07.1933, Černjavski, P., Rudski, I., Lindtner, V. ( BEO! 08466) ; Bertisci, m. Pločica–Bogićevica, blizu Tropoje , [ MGRS 34 T DN21 ], pored puta u kamenjarima, 18.07.1933, Černjavski, P., Rudski, I., Lindtner, V. ( BEO! 08473) ; Bertisci, m. Koprivnik, [ MGRS 34 T DN31 , DN32 ], in pascuis subalpinis, 06.1923, Soška, Th. ( BEOU! 41121) ; Bertisci, m. Koprivnik ( Peć ), [ MGRS 34 T DN31 , DN32 ], in pascuis subalpinis 15.06.1923, Soška, Th. ( BEOU! 41122) ; Bertisci, m. Koprivnik ( Peć ), [ MGRS 34 T DN31 , DN32 ], in mughetis, 15.06.1923 Soška, Th. ( BEOU! 41123) ; Bertiscus (Alpes boreales albanicae), in monte Lumbardska planina prope oppidum Peć ( Ipek ), [ MGRS 34 T DN22 , DN32 ], c. 2100 m, substr. calc. 08.– 09.07.1933, Rechinger fil., Scheffer 534 ( W! 6669) ; Pločica supra man. Dečani, [ MGRS 34 T DN21 , DN31 ], 1933, Lindtner, V. s.n. ( BEO!) ; sa Mokre Gore, Petrović, D. ( BEOU! 41298) ; Prokletije, m. Kurvala, [ MGRS 34 T DN31 ], pod vrhom, na livadama, 16.07.1933, Černjavski, P., Rudski, I., Lindtner, V. ( BEO! 08539) ; Prokletije, m. Kurvala (Pločica), [ MGRS 34 T DN21 , DN31 ], škriljci, 2100–2200 m, 16.07.1933, Černjavski, P., Rudski, I., Lindtner, V. ( BEO! 08465) ; Prokletije, pod Kurvalom, Ločanska planina, [ MGRS 34 T DN31 ], na suvatima iznad šumskog pojasa, 15.07.1933, Černjavski, P., Rudski, I., Lindtner, V. ( BEO! 08470) ; Prokletije, m. Ločanska Bistrica – Pločica, blizu žandarmerijske stanice, [ MGRS 34 T DN21 , DN31 ], u šiblju na livadi, 15.07.1933, Černjavski, P., Rudski, I., Lindtner, V. ( BEO! 08505) ; Prokletije , Koprivnik, [ MGRS 34 T DN31 , DN32 ], vlažni mugus, 11.07.1963, Janković, M. ( BEOU! 41293) ; Prokletije , Lumbardske planine, kod doma, [ MGRS 34 T DN22 , DN32 ], munikova šuma, južna padina, 22.05.1959, Janković, M. ( BEOU! 41297) ; Prokletije, Prilepske planine, potez Raški do– Savine vode– Amšor–Žljeb–Kovraga , [ MGRS 34 T DN21 , DN22 ], 28.05– 07.06.1960, Janković, M. ( BEOU!41295) ; Prokletije, Rugovska klisura, Slane poljane, [ MGRS 34 T DN32 ], munikova šuma, 06.07.1960, Janković, M. ( BEOU! 41294) ; Paštrik, [ MGRS 34 T DM67 ], 19.07.1980, Džukić, G., Milojević, B. ( BEOU! 1034, 1071, 1093, 1117) ; Paštrik bei Prizren, auf dem Berge, [ MGRS 34 T DM67 ], 1800 m, 20.– 21.07.1936, Behr, O., Behr, E. ( W! 00487) ; distrikt Hasi, Paštrik, [ MGRS 34 T DM67 ], an felsigen Hängen in der Gipfelregion , c. 1800 m, 16.05.1916, Dörfler, I. 479 ( WU! 2776) ; Ms Koritnik, [ MGRS 34 T DM65 , DM66 ], 05.1937, Oehm, H. ( BEO! 08506) ; ALBANIA. Mali e Shebenikut, Darkat e Librazhd–Katund , [ MGRS 34 T DL56 ], pasište, 1700 m, 02.06.2006, Melovski, Lj. ( MKMEL! 07347) ; m. Jalica, in saxosis pag. Fagi , [ MGRS 34 T DM55 ], 10.05.1913, Košanin, N. ( BEOU! 41117) ; m. Jalica, in glareosis calcaris pag. Fagi , [ MGRS 34 T DM55 ], 10.05.1913, Košanin, N. ( BEOU! 41118) ; m. Jalica , [ MGRS 34 T DM55 ], zaravan, 14.07.1919, Košanin, N. ( BEOU! 41332) ; distr. Scutari, m. Cukali [ MGRS 34 T CM96 ], in saxosis elatis, 15.06.1897, Baldaccii, A. 154 ( W! 1594) ; distr. Scutari, m. Cukali , [ MGRS 34 T CM96 ], in saxosis elatis, 15.06.1897, Baldaccii, A. 154 ( WU! 5274) ; Distrikt Šala, Felsige Hänge an der Fuša Nermajns bei Abata, [ MGRS 34 T DM08 ], ca. 1800 m, 22.06.1916, Dörfler, I. 163 ( WU! 2776) ; nördl. von Skutari, Auf steinigen Wiesen des Maranaj , [ MGRS 34 T CM87 ], ca. 1230 m, 1.06.1916, Dörfler, I. No. 13 ( WU! 2776) ; MONTENEGRO. Prokletije, m. Bjelič, [ MGRS 34 T DN 00], krečnjaci alpiskog pojasa, 22.07.1933, Černjavski, P., Rudski, I., Lindtner, V. ( BEO! 08538) ; Prokletije, dolina Bjeluhe, [ MGRS 34 T DN22 ], livade 15.07.1973, Nikolić, V., Diklić, N., Bogdanović, M. s.n. ( BEO!) ; Prokletije, Djel, livade na padini prema Bjeluhi, [ MGRS 34 T DN22 ], 27.07.1933, Černjavski, P., Rudski, I., Lindtner, V. ( BEO! 08502) ; Prokletije, na putu Plav – Djel, [ MGRS 34 T DN21 , DN22 ], livade u bukovoj šumi, 1450 m, 27.07.1933, Černjavski, P., Rudski, I., Lindtner, V. ( BEO! 08451) ; Bertiscus, Murino, in valle fl. Lim prope Andrijevica , [ MGRS 34 T DN02 ], 29.06.1938, Lemperg, Fr. s.n. ( BEO!) ; 29.06.1938, Lemperg, F. 707 ( W! 11400) ; Prokletije, Grebaja, Volušnica , [ MGRS 34 T CN90 ], sipari Drypetea spinosae, krečnjak, 1314 m, 16.07.2010, Lakušić, D. 31166 ( BEOU!) ; Prokletije, Maja Šćapica, [ MGRS 34 T DN10 ], silikat, 08.07.2005, Niketić, M., Tomović, G. 19878 ( BEOU!) ; Kučke Prokletije, Širokar – Bigeze, [ MGRS 34 T CN81 ], 42° 34’ 44.2’’ N, 19° 35’ 16.4’’ E, rudine ( Festuco – Seslerietea ), krečnjak, 1884 m, 19.06.2008, Lakušić, D. 27263 ( BEOU!) ; Prokletije, Ćaf Bor – Sapica , [ MGRS 34 T DN00, DN10], Seslerietalia comosae, silikat, 1950 m, 07.07.1995, Stevanović, V., Lakušić, D., Niketić, M., Bulić, Z. Hadžiablahović, S. 859/95 ( BEOU!) ; Prokletije, Ćaf Bor – Sapica , [ MGRS 34 T DN00, DN10], Juniperion sibiricae, silikat, 1950 m, 07.07.1995, Stevanović, V., Lakušić, D., Niketić, M., Bulić, Z. Hadžiablahović, S. 890/95 ( BEOU!) ; Prokletije, Ćaf Bor – Sapica, [ MGRS 34 T DN00, DN10], lokve, Pinetum peucis, silikat, 1950 m, 08.07.1994, Lakušić, D., Niketić, M. 369/94 ( BEOU!) ; Prokletije, Čakor, Devojački krš, [ MGRS 34 T DN12 ], 03.08.2007. Niketić, M. 24857 ( BEOU!) ; Bertiscus, Alpes boreales albanicae, in declivibus orientalibus sub jugo Čakor prope pagum Plav, [ MGRS 34 T DN12 ], 1300– 1700 m, Rechinger fil. et Scheffer 925 ( W! 6668) ; östl. Murino an der Passhöhe , [ MGRS 34 T DN02 ], ostwärts in Kalkshutt, 1720 m, mont. Orien, [ MGRS 34 T CN01 ], in saxosis calcareis alpinis, 07.1916, Adamović, L. ( WU! 5274) ; Crkvice, Krivošije, [ MGRS 34 T CN01 ], 18.05.06 Schneider, J. ( W! 8671) ; Jelovi do, Krivošije, [ MGRS 34 T CN01 ], 16.04.06 Schneider, J. ( W! 8670) ; Lovčen am Weg nach Cettinje, Nathere Orte an bushigen Abhangen, [ MGRS 34 T CM29 ], 8000– 1000 m, 17.– 18.05.1886, Bornmüller, J. 1003 ( BEOU! 41120) ; Orlov Krch, [ MGRS 34 T CM36 ], 04.1907, Bierbarch ( BEOU! 41119) ; Rumija, Sozina, prevoj Petrovac – Skadarsko jezero, [ MGRS 34 T CM37 ], 28.04.2005, Niketić, M., Tomović, G., Lazarević, P., Tomašević, M. 19694 ( BEOU!) ; Megjurečka planina prope fines Turcorum dist. Primorije, [ MGRS 34 T CM55 ], in saxosis, 15.06.1898, Baldaccii, A. 53 ( W! 332, WU! 5274) ; Sutorman Loftan, [ MGRS 34 T CM46 ], 06.1841, Ebel s.n. ( W!) ; Sutorman, [ MGRS 34 T CM46 ], 05.2009, Anačkov, G. s.n. ( BUNS!) ; BOSNIA AND HERZEGOVINA. circa Trebinje, [ MGRS 34 T BN83 ], in pascuis, 06.1888, Adamović, L. ( WU! 84) .

Viola kopaonikensis Pančić ex Tomović & Niketić , sp. nov. ( Figs. 9 & 10 a-c); Niketić, Botanica Serbica 38(2): 226 (2014), nom. inval. (nom. propos.).

– V. kopaonikensis Pančić 1888 , Srpska Kralj. Bot. Bašta Beograd 1887: 10, nom. inval. (nom. nud.) – “ V. lutea ” sensu Pančić 1874 , Fl. Prin. Serb.: 158, non Smith, Fl. Brit. 1: 248 (1800), p.p. – “ V. alpestris subsp. aetolica ” sensu W. Becker, Beih. Bot. Centr. 26(2): 345 (1910), non (Boissier & Heldreich) W. Becker, loc. cit. – “ V. saxatilis subsp. aetolica ” sensu Hayek, Denkschr. Kaiserl. Akad. Wiss., Wien. Math.-Naturwiss. Kl. 99: 133 (1924), non (Boissier &

Heldreich) Hayek, loc. cit. ; Hayek, Repert. Spec. Nov. Regni Veg. Beih. 30(1): 516 (1925) [Prodr. Fl. Penins. Balc. 1]. – “ V. aetolica ” sensu Diklić , Flora SR Srbije, 3: 161 (1972) non Boissier & Heldreich in Boissier , Diagn. Pl. Orient. ser. 2, 6: 24 (1859). Type:— SERBIA. Central Serbia, Mt Kopaonik, Šiljak [“Treska”] peak, [ MGRS 34 T DN88 ], 43.25985 N ; 20.78522 E, vegetacija stena i rudina, ultramafit, 1604 m, 26–June–2004, Niketić, M., Tomović, G., Novčić, R. 18933 (holotype: BEOU! 17139; isotypes: BEO!

A 27; MKMEL! 07500; WU!)

Typification: —During a comprehensive examination of the Pančić’ s collection, Herbarium Pancicianum (BEOU), original material of V. kopaonikensis was not found. In the Herbarium of the Royal Botanic Garden Edinburgh there is a specimen of this plant which was cultivated in the botanical garden in Belgrade in 1887 (06.1887, leg. et det. J. Bornmüller 00327619 E!). Given that a year later, the booklet on the plants cultivated in the Belgrade Botanical Garden in 1887 was already published ( Pančić 1888), it can be assumed that Pančić named his species on the basis of the mentioned specimen. However, according to ICN ( McNeill et al. 2012, Art. 38.1) this name is not validly published (nomen nudum) and, as such, it could not be typified or validated. Although in this paper the name of a new species was retained and ascribed to Pančić, type specimens are still selected from a more complete recent material (BEOU 40661; BEO A 27).

Description: —Usually short-lived perennial (rarely annual or biennial), (8)15–20(25) cm height, branched from base. Shoots pubescent in lower part, green to canescent. Lower leaves with longer petioles than those from middle and upper; blade ovate to orbicular, obtuse or rounded, coarsely crenate, pubescent. Stipules of lower leaves small, oblong to oblong-spathulate, entire or two to three-partite. Middle and upper leaves with shorter petioles; blade oblong-ovate to lanceolate or oblanceolate, 1.5–2 times longer than stipules, gradually attenuated at the base into the petiole, obtuse or acute, crenate, pubescent. Stipules of middle and upper leaves pinnately divided, pubescent; the undivided terminal part relative large, with 1–3 lateral segments on the inner and 2–6 on the outer side, tapering gradually downwards. Peduncles 4–10 cm, puberulent to pubescent in lower part, glabrescent in middle part and densely papillose at top; bracts in upper third of peduncle. Flowers small to medium size, 1.6–2.3 × 1.3–1.9 cm, bright yellow; lower surface of petals often becoming blue-violet at the end of flowering period. Calyx: upper sepals (5.2)7.4–9.5(11.8) × (2.2)2.9–3.6(5.4) mm; lateral (6.3)8.3–10.6(13.4) × (2)2.6–3.4(4.8) mm; lower (7.7)9.4–11.9(15.2) × (2.5)3.2–4.2(5.9) mm. Sepals broadly lanceolate to ovate-lanceolate or elliptic-lanceolate, mostly acute, attenuate at base, mostly densely pubescent on back, sometimes becoming glabrous in fruit, ciliate at margin; margin with 1–3 ± conspicuous teeth on both sides, or often entire. Appendages 1/4–1/3 as long as sepals, narrow to wide rectangular, trapezoid or semiorbicular, attenuate at base, coarsely dentate. Corolla: upper petals (8.5)10–12.3(16) × 6(7.4)–9.4(11.8) mm, obovate to orbiculate; lateral (9)10.4–12.7(15.7) × (5)6.1–7.2(9.9) mm, ovate to irregularly ovate, papillose-hairy at the upper margins; lower without spur (8.2)10.1–11.8(13.8) × (8)9.7–11.3(13.8) mm slightly triangular to cordate, usually longitudinally folded along the midrib. Spur glabrous, straight to slightly curved upwards, yellow to yellow-violet, (6.3)7.1–8.1(9.6) mm × (1.5)1.7–2(2.2) mm; ratio of length/width: 3.9–4.6. Seeds ellipsoid, pale to dark brown. Flowering period: May–July.

Etymology: —The specific epithet is retained from the original source ( Pančić 1888) and derived from the type locality Mt Kopaonik in Central Serbia. Although it is recommended by the Melbourne Code (Rec. 23Aj, McNeill et al. 2012) that the names of the species with a wider range should not refer to restricted geographical attribute, in the honour of Dr. Josif Pančić, who first suggested the name “ Viola kopaonikensis ” we decided to keep this epithet.

Chromosome number: —2n = 20 (counted on plants from the type locality in Mt Kopaonik and Mt Rogozna; Fig. 7b ).

Distribution: — V. kopaonikensis is an endemic species of the Republic of Serbia, restricted to several mountains in Central Serbia (Mt Kopaonik and Mt Željin) and southwestern Serbia (Mt Rogozna and Pešter plateau). Literature data for “ V. saxatilis subsp. aetolica ” ( Pavlović 1953, 1955) and “ V. aetolica ” ( Diklić 1972) from the ultramafic mountain pastures of Mt Ozren are erroneous. ( Fig. 11 ).

Habitat and ecology: — V. kopaonikensis inhabits xerophile montane, subalpine and alpine rocky grounds, on ultramafic geological substrate, at the elevation from 1100 to 1900 m. The habitats of this endemic species in Mt Kopaonik are subalpine and alpine rocky grounds of vegetation class Festuco –Seslerietea, as well as montane rocky grounds of the class Festuco –Brometea at the elevations from 1500 to 1900 m. In Mt Rogozna, as well as on the Pešter plateau, this plant was found at the elevations between 1100 and 1350 m. At these localities, V. kopaonikensis also inhabits montane rocky grounds of vegetation class Festuco –Brometea ( Fig. 10d ).

Discussion: —The name “ Viola kopaonikensis Panč. (nov. spec.)” was introduced in the botanical literature by Pančić in his last published paper ( Pančić 1888).After that, only Adamović (1909) accepted “ V.kopaonikensis ” as a good narrow endemic species. However, as Pančić gave no formal botanical description, this name could not be accepted as a valid in the botanical literature. According to Becker (1910) “ V. kopaonikensis Panč. ” is the synonym of V. alpestris subsp. aetolica (Boissier & Heldreich) Becker (1910: 345) . Hayek (1925) includes the name “ V. Kopaonicensis Panč. in sched.” in “ V. saxatilis subsp. aetolica ”. This opinion was followed by Diklić (1972) who included the violet from Mt Kopaonik in the synonymy of V. aetolica . The name V. kopaonikensis was no longer mentioned in the recent floristic literature ( Valentine et al. 1968, Erben 1985).

Although the most authors confused Pančić’ s violet with V. aetolica , our comparative analysis of morphological characters of flowers, as well as the number of chromosomes in populations from Mt Kopaonik, Mt Rogozna and the Pešter plateau (2n = 20), suggest that individuals from these localities do not belong to V. aetolica (2 n = 16), but to a new species, V. kopaonikensis . Populations of V. kopaonikensis are the most similar to V. elegantula by the following characters: narrow spur (1.7–1.9 mm), gradually narrowed toward the apex and linear-lanceolate to ovate-lanceolate sepals. V. elegantula also has the same chromosome number (2n = 20, Erben 1985) as V. kopaonikensis . On the other hand, V. kopaonikensis differs from V. elegantula by a denser indumentum, relatively longer middle leaves, smaller and strictly yellow flowers and a relatively shorter spur ( Table 3). In these morphological characters V. kopaonikensis is similar to V. aetolica , which could explain their previous taxonomic treatment.

The closely related species V. elegantula was described by Schott (1857: 167, “Dalmatia (Prologh)”), on the basis of the plant material collected by Roberto de Visiani (as “ V. gracilis Sibth. et Sm. / Verlica Mt Ghniat”, PAD). Type locality is actually in Mt Dinara in Bosnia and Herzegovina and lectotypification was done by Erben (1985). V. elegantula is distributed in Croatia ( Erben 1985, Nikolić 2015), Bosnia and Herzegovina ( Beck 1918, Erben 1985, Lubarda 2013), Montenegro ( Rohlena 1942, Erben 1985), Albania ( Erben 1985, Qosja et al. 1992, Meyer 2011), Republic of Macedonia ( Erben 1985, Micevski 1995) and Serbia ( Diklić 1972, Tomović et al. 2014) ( Fig. 11 ). It inhabits different habitat types: mesophyllous montane, subalpine and alpine rocks, rocky grounds and pastures and forest (spruce, beech, pine) edges within elevation range from 700 to 2400 m. V. elegantula can be found on carbonate and silicate geological substrates and the only ultramafic localities of this species are in Mt Ozren and Mt Giljeva Planina in southwestern Serbia. Nevertheless, on these two ultramafic mountains, V. kopaonikensis was not found.

We propose that V. kopaonikensis may have been derived from V. elegantula by peripatric edaphic speciation, and some differences between these two species (e.g. lower growth and denser indumentum) could reflect adaptations of V. kopaonikensis to the harsh ecological conditions of the ultramafic soils. Such microevolutionary processes were described for other species on heavy metal soils ( Wierzbicka & Rostański 2002, Hildebrandt et al. 2006). However, morphological characters such as relatively longer middle leaves, smaller flowers and relatively wider spur goes in favour that V. kopaonikensis represents a different and well recognized species. Besides, edaphic stenoendemics are not so unusual in the Balkan Peninsula, only several species from V. sect. Melanium grow on very restricted and specific geological substrates, such as V. vourinensis Erben (1985: 500) , V. beckiana Fiala ex Beck (1896: 197) and V. allchariensis Beck (1894: 6) .

Additional specimens examined:— SERBIA. Kopaonik, in saxosis, 07.1864, Pančić, J. (BEOU! 3221); Kopaonik, 1885, Pančić, J. (culta in Horto Belgr. 06.1887, Bornmueller, J.) (gold gella) (E! 00327619); Kopaonik, in alpinis, 05.1903, Bierbach (WU! 125); Kopaonik, Anonymus s.n. (BEO!); Kopaonik, Anonymus (BEO! 08590); Kopaonik, Anonymus (BEOU! 41242); Kopaonik, Jadovnik, istočna padina, [MGRS 34TDN79], pašnjak, 14.07.1951, Pavlović, Z. s.n. (BEO!); Kopaonik, Jadovnik, [MGRS 34T DN79], vrh, kamenjar, 28.05.1951, Pavlović, Z. s.n. (BEO!); Kopaonik, Kopaonička kula, [MGRS 34T DN88], stene, 13.07.1938, Rudski, I. s.n. (BEO!); Kopaonik, Kukavica, [MGRS 34T DN79], ass. Poa molinieri-Plantago carinata, 14.07.1951, Pavlović, Z. s.n. (BEO!); Kopaonik, Kukavica, [MGRS 34T DN79], 10.08.1986, Lakušić, D., Niketić, M. (BEOU! 41103); Kopaonik, Kukavica, [MGRS 34T DN79], u zajednici Carex humilis – Bromus erectus , krečnjak, 1700–1800 m, 07.08.1986, Lakušić, D., Niketić, M. (BEOU! 41104); Kopaonik, Kukavica, [MGRS 34T DN79], serpentinit, 1960 m, 27.06.2004, Niketić, M., Tomović, G. 18901 (BEOU!), s.n. (BEO!); Kopaonik, Kukavica–Kozje stene, [MGRS 34T DN79], pašnjak, 08.06.2001, Grahek, I. s.n. (BUNS!); Kopaonik, Nebeske stolice, [MGRS 34T DN88], pašnjak, 11.07.1951, Pavlović, Z. s.n. (BEO!); Kopaonik, Nebeske stolice, [MGRS 34T DN88], 16.07.1988, Anonymus (BEOU! 41102); Kopaonik, Nebeske stolice, [MGRS 34T DN88], stene i kamenjari Silenetum serbicae, serpentinit, 1900 m, 07.07.1989, Lakušić, D. 4/89 (BEOU! 41101); Kopaonik, Nebeske stolice, [MGRS 34T DN88], 1900 m, 07.05.1991, Lakušić, D. 677/91 (BEOU! 41106); Kopaonik, Nebeske stolice, [MGRS 34T DN88], serpentinit, 1900 m, 22.05.2002, Lakušić, D. 16008 (BEOU!); Kopaonik, Nebeske stolice, [MGRS 34T DN88], serpentinit, 1700–1800 m, 26.06.2004, Niketić, M., Tomović, G. 18916 (BEOU!), s.n. (BEO!); Kopaonik, Nebeske stolice, [MGRS 34T DN88], serpentinit, 09.07.2005, Niketić, M., Tomović, G. 19915 (BEOU!), s.n. (BEO!); Kopaonik, ad Bajgora ka Ostrem Koplju, uz Šalju, [MGRS 34T EN06], 12.04.1913, Košanin, N. (BEOU! 41107); Kopaonik, Oštro Koplje, [MGRS 34T EN06], 12.04.1913, Košanin, N. (BEOU! 41108); Kopaonik, Ostro Koplje–Bare, [MGRS 34T EN06], 30.06.1914, Soška, Th. (BEOU! 41109); Kopaonik, Ostro Koplje– Kačandol, [MGRS 34T EN06], 29.06.1914, Soška, Th. (BEOU! 41110); Kopaonik, Kačandol–Bela stena, [MGRS 34T EN06], 30.06.1914, Soška, Th. (BEOU! 41111); Kopaonik, Suvo rudište, [MGRS 34T DN88], jugoistočne padine, pašnjak, 26.05.1951, Pavlović, Z. s.n. (BEO!); Kopaonik, Suvo rudište, [MGRS 34T DN88], jugoistočni greben, pašnjak, 26.05.1951, Pavlović, Z. s.n. (BEO!); Kopaonik, Šiljak [“Treska”], vrh, [MGRS 34T DN88], 1620m, 17.07.1951, Pavlović, Z. s.n. (BEO!); Kopaonik, Šiljak [“Treska”], južne padine, [MGRS 34T DN88], kamenjari, serpentinit, 1500–1600 m, S exp. 21.07.1987, Lakušić, D. (BEOU! 41105); Kopaonik, Šiljak [“Treska”], [MGRS 34T DN88], serpentinit, 09.07.2005, Niketić, M., Tomović, G., Novčić, R. 19923 (BEOU!), s.n. (BEO!); Kopaonik, Šiljak [“Treska”], [MGRS 34T DN88], serpentinit, 08.06.2012, Vukojičić, S., Tomović, G., Zlatković, B., Jakovljević, K., Kabaš, E. 35320 (BEOU!); Mt Kopanik, Šiljak [“Treska”] peak, [MGRS 34T DN88], visokoplaninski pašnjak, serpentinit, 1604 m, 28.05.2014, Tomović, G., Jakovljević, K., Đurović, S., Buzurović, U. 40661 (BEOU!), s.n. (BEO!); Željin in Kruševaces Kr., [MGRS 34T DP81], 08.1852, Pančić, J. (BEOU! 3246); Pešter, Trojan brdo, [MGRS 34T DN37], 43°06’33.2’’ N, 20°09’36.5’’ E, pašnjaci sa dominacijom Carex humilis i Bromus erectus , serpentinit, 1350 m, 08.07.2010, Niketić, M., Tomović, G. 30749 (BEOU!); Pešter, Trojan brdo, [MGRS 34T DN37], kamenjari, serpentinit, 03.07.2008, Lazarević, P. (BEOU! 16314); Pešter, Tuzinje, kota 1213, [MGRS 34T DN27], kamenjari, serpentinit, 1222 m, 11.05.2014, Niketić, M., Tomović, G., Buzurović, U. 40640 (BEOU!), s.n. (BEO!); Pešter, Tuzinje, kota 1213, [MGRS 34T DN27], kamenjari, serpentinit, 1222 m, 28.05.2014, Tomović, G., Jakovljević, K., Đurović, S., Buzurović, U. 40665 (BEOU!); Rogozna, inter Grižani Han et Karaula, [MGRS 34T DN76], 06.1914, Soška, Th. (BEOU! 41100); Rogozna, Velika Skakavica, [MGRS 34T DN76], 43°02’26.8’’ N, 20°43’04.9’’ E, pašnjaci sa dominacijom Carex humilis i Bromus erectus , serpentinit, 1166 m, 08.07.2010, Niketić, M., Tomović, G. 30742 (BEOU!); Rogozna, Pasji potok–Vinorog, [MGRS 34T DN77], 43°07’45.4’’ N, 20°38’46.5’’ E, pašnjaci pored puta, serpentinit, 1100 m, 15.05.2011, Niketić, M., Tomović, G. 32156 (BEOU!); Rogozna, Vinorog, vrh, [MGRS 34T DN77], 43°07’56.4’’ N, 20°38’06.5’’ E, pašnjaci, serpentinit, 1200–1220 m, 15.05.2011, Niketić, M., Tomović, G. 32177 (BEOU!).

Figures

FIGURE 5. Viola pseudaetolica. A. lower leaf with stipule; B. middle leaf with stipule; C. upper leaf with stipule; D. flower, front view. E. flower, lateral view with spur; F. lower sepal; G. middle sepal; H. upper sepal; Scale bar = 10 mm.Artists: Ivana Janković & Rastko Ajtić.

FIGURE 6. Viola pseudaetolica Tomović, Melovski & Niketić, sp. nov. (from the locus classicus). a) habitus. b) flower, lateral view with spur. c) flower, front view. d) high-mountain pastures in Mt Jablanica. Photographs: Marjan Niketić.

FIGURE 7. Metaphase chromosome plates: a) Viola pseudaetolica 2n = 16 (population 4—Mt. Jablanica). b) Viola kopaonikensis 2n = 20 (population 10—Mt. Rogozna). Photographs: Uroš Buzurović & Maja Lazarević.

FIGURE 8. Distribution of Viola aetolica (black marks) and Viola pseudaetolica (red marks) in the Balkan Peninsula. All literature data are related to Viola aetolica. All doubtful and erroneous literature data refer to V.aetolica, but in the north and northwest they are attributed to V. pseudaetolica (red marks). Population numbers correspond to Pop. № in Tab. 1.

FIGURE 9. Viola kopaonikensis. A. lower leaf with stipule; B. middle leaf with stipule; C. upper leaf with stipule; D. flower, front view. E. flower, lateral view with spur; F. lower sepal; G. middle sepal; H. upper sepal; Scale bar = 10 mm.Artist: Ivana Janković.

FIGURE 10. Viola kopaonikensis Tomović & Niketić, sp. nov. (from the locus classicus). a) habitus. b) flower, lateral view with spur. c) flower, front view. d) high-mountain pastures on ultramafic substrate in Mt Rogozna. Photographs: Marjan Niketić.

FIGURE 11. Distribution of Viola elegantula (black marks) and Viola kopaonikensis (red marks) in the Balkan Peninsula. Doubtful records for V. aetolica are attributed to V. kopaonikensis (red marks). Population numbers correspond to Pop. № in Tab. 1.