Odontarrhena decipiens 2018

Cecchi, Lorenzo, Bettarini, Isabella, Colzi, Ilaria, Coppi, Andrea, Echevarria, Guillaume, Pazzagli, Luigia, Bani, Aida, Gonnelli, Cristina & Selvi, Federico, 2018, The genus Odontarrhena (Brassicaceae) in Albania: Taxonomy and Nickel accumulation in a critical group of metallophytes from a major serpentine hot-spot, Phytotaxa 351 (1), pp. 1-28 : 11-12

3. O. decipiens ( Nyárády 1928: 113) L.Cecchi & Selvi , comb. nov.

(≡) Alyssum decipiens Nyár.

Lectotype (designated here):— GREECE. “in summis saxosis montis Smolika distr. Konitza”, 18/07/1896, A. Baldacci , Iter Albanicum (Epiroticum) Quartum no. 206, WU0043221 , WU!

Isolectotype:—“in saxosis regionis Abietinarum m. Smolika supra Kerasovo distr. Konitza”, BM000750128, BM!

We were unable to trace a third specimen whose image appeared in a later publication by Nyárády (1929b: 44).

(=) A. kosaninum var. obovatifolium Nyárády (1928: 97) , syn. nov.

Holotype:— ALBANIA. “Lurija”, 06/07/1913, N. Košanin , BEO07360 [right specimen], BEO!

(=) A. balkanicum Nyárády (1930: 393) , syn. nov. [Latin diagnosis published later by Markgraf (1931: 337)]. − A. balkanicum f. elatum Nyárády (1930: 394) , nom. inval. − A. balkanicum var. elatum Nyárády (1949: 157) , nom. inval.

Neotype (designated here):— ALBANIA. “ Prefettura di Durazzo (Durrës), Distretto di Croia (Krujë), versante occidentale del Maja e Liqenit, pietraia serpentinosa sul primo tornante a gomito subito oltre il passo di Shtamës, 1010 m, 41°31’11.42”N, 19°52’51.85”E ”, 18/07/2016, I. Bettarini, L. Cecchi, A. Coppi & F. Selvi, FI050442 , FI!

Isoneotypes:— B !; FI050893 , FI !; K!; JE!; TIR!; Herb. Cecchi no. 3372!

Alyssum balkanicum was originally described in two forms, later treated as varieties; “f. elatum ” is here selected as the one corresponding to the type, and is thus invalid with respect to the autonym ( ICN Art. 26.2; McNeill et al. 2012). Designation of a neotype is required since the following original collections (syntypes, all listed as “f. elatum ”) were lost during the war and no duplicates were found in E, TIR or Z: “Nord-Albanien: Landschaft Matja [ Mat river ] (östlich Kruja) […] Zalli Germanit, Serpentinschutt, 500 m ü.d. M.”, 12/07/1928, F. Markgraf & W. Pieper, no. 1614 & 1615 ; Proj Rrepës zw. Ben [Bejne] und Cëruj [Ceruje], Serpentingeröll, 400 m ü.d. M.”, 27/05/1928, F. Markgraf & W. Pieper, no. 1118 ; “Nord-Albanien: Östliche Matja, Mal i Alamanit, Stan i Alamanit, Serpentinschutt, 1400 m ü.d. M.”, 15/07/1928, F. Markgraf & W. Pieper, no. 1688 ; “Nord-Albanien: Stammesgebiet Merdita, Östteil des Mal i Shenjit (Bajrak Oroshi), Qaf e Mshkallës am FuSS der Zepa, 1200 m ü.d. M.”, 24/07/1928, F. Markgraf & W. Pieper, no. 1848. The selected neotype was collected in the locality to the east of Krujë and fits the original description of the species .

(=) A. balkanicum f. depressum Nyárády (1930: 394) , syn. nov. ≡ A. balkanicum var. depressum (Nyár.) Nyárády (1949: 157) .

Neotype (designated here):— ALBANIA. “ Prefettura di Dibër, Distretto di Mat , lungo la sterrata che sale da ovest al passo di Murrë (Qafë Murrë), tra i villaggi di Lis e Vig, a margine del bosco, 870 m, 41°38’6.28”N, 20° 7’26.80”E ”, 12/06/2017, L. Cecchi & F. Selvi, FI050830 , FI!

The neotype is from the Lurë mountains, as mentioned in the original description ( Nyárády 1930: 395): “Nord-Albanien: Stammesgebiet Lurja, Kunora e Lurës, Qaf e Thëres (Südöstlich Kumbla [Kumbull]), Wiese, Serpentin, 1500 m ü. D. M.”, 21/07/1928, F. Markgraf & W. Pieper, no. 1810. The other original collection was: “Mittel-Albanien: Gur i Topit (Quellgebiet des Shkumbins), SW.-Seite, Serpentinschutt westl. Qaf e Shalësit, 1500 m ü. D. M.”, 27/06/1928, F. Markgraf & W. Pieper, no. 1540.

(=) A. balkanicum var. calvescens Nyárády (1949: 158) , syn. nov.

Neotype (designated here):— ALBANIA. “ prefettura di Kukës, distretto di Tropojë, Fierzë, pietraia di serpentino e arbusteto basso lungo la strada presso il ponte sul lago omonimo, 220 m, 42°16’ 5.92”N, 20° 1’34.54”E ”, 12/07/2016, I. Bettarini, L. Cecchi, A. Coppi, C. Gonnelli, P. Meerts & F. Selvi, FI050444 , FI!

The original collection of this taxon was from the same area of the neotype designated here (lower Valbona valley close to Fierzë): ALBANIA. “Unt. Valbone-Tal”, sine die, sine coll., “Herb. Schütt Bremen” (not found in BREM).

(=) A. balkanicum f. microcarpum Nyárády (1949: 158) , syn. nov.

Neotype (designated here):— ALBANIA. “ prefettura di Durazzo (Durrës), distretto di Croia (Krujë), versante occidentale del Maja e Liqenit, pietraia serpentinosa sul primo tornante a gomito subito oltre il passo di Shtamës, 1010 m, 41°31’11.42”N, 19°52’51.85”E. Serpentino”, 06/10/2017, I. Bettarini & F. Selvi, FI052160 , FI!

The original collections used to describe this taxon (syntypes) were: “ Albania media: Kruja, Östl. der qaf e Shtamёs, serp. 900 m ”, sine die, F. Markgraf ; “ Matja , Qafa e Müreş ”, sine die, F. Markgraf .

(=) A. lurense Meyer (2011: 64) , syn. nov. ≡ O. lurensis (F.K.Mey.) Španiel, Al-Shehbaz, D. A.German & Marhold in Španiel et al. (2015: 2485).

Holotype:— ALBANIA. “Lura, Kunora e Lures, ca. 1300 m ”, 02/08/1959, F. K. Meyer, Flora Albanica no. 4627, JE00016685 , JE!

Flowering stems 20–90 cm, erect, robust. Sterile shoots at the base always present and more or less abundant, often forming densely leafy cushions. Basal leaves up to 25 × 9 mm, broadly spathulate and obtuse, greenish with scattered hairs above, white-silvery below for dense, overlapping hairs with 10–15 rays 0.3–0.6 mm across. Leaves on flowering stems numerous, narrowly oblong-spathulate, flat. Inflorescence broadly corymbose, dense, with branching of third and, more commonly, fourth level. Partial racemes usually short and dense, with up to 15 fruits. Fruiting pedicels erecto-patent, never flexuous. Sepals 2.2 × 0.5–0.8 mm. Petals 1.8–2.8 mm. Style 0.8–1.0 mm. Siliculae (2.5)3.5(4) × (1.6)2.0(2.7) mm, narrowly to broadly elliptic, acute to subacute, usually symmetrical; valves not undulate when ripe, indistinctly veined; valves glabrous to glabrescent, ca. 0.2 mm across, 5–10 rayed. Seeds 2.0– 2.2 mm long, including a wing of 0.4–05 mm. Figs. 3C , 6D , 7D , 9 .

Phenology. Flowering from late June to mid August, fruits ripening from the end of July to the first half of October ( Fig. 4 ).

Chromosome number. 2n = 32+0–2B ( Fig. 5F ); plants from N (south of Gjegian, region of Püke, FI050443) and C Albania (east of Krujë, FI050442).

Distribution and ecology. Mainly on the outcrops of N and C Albania ( Fig. 1 ; Appendix 1). It grows strictly on ultramafic rocks, from 220 to 1900 m a.s.l., both in anthropogenic sites, such as fields and pasturelands on shallow soil, and natural habitats such as rocky slopes, screes, stony and gravelly soils and mountain grasslands. Also present in Kosovo, Macedonia and N Greece.

Nickel accumulation. The lowest concentration was detected in plants from Fierzë, the highest in those from the Krujë region, with a difference of over 9000 μg g-1 dw ( Table 1).

Comments. This species was described based on a Baldacci’s collection from Mt. Smolikas in N Greece, and interpreted as a hybrid between O. smolikana and O. muralis (Nyárády 1927) . It was then included in O. muralis s.l. ( Ball & Dudley 1993, Jalas et al. 1996) or O. chalcidica , though it was stated that its identity remains uncertain ( Hartvig 2002). Our Albanian collections, with clearly intermediate characters between O. smolikana subsp. glabra and O. chalcidica , match the description and type collection of O. decipiens and can be readily referred to this taxon. Its distinctness is supported by the stability of the diagnostic characters in large populations, which are widely distributed in C and N Albania. These are mostly found within the ranges of O. chalcidica and O. smolikana , at an intermediate altitude, and have likely resulted from extensive homoploid hybridization at the contact zone between populations of the latter two taxa, both 2n = 32. The same process has likely occurred in the area of Mt. Smolikas, where O. smolikana subsp. smolikana and O. chalcidica occur at different altitudes ( Hartvig 2002). The separate status of O. decipiens is supported by the various later names that were published for it based on material from the mountains of C Albania, such as A. kosaninum var. obovatifolium , A. balkanicum s.l. and A. lurense . To date, the conspecificity of these taxa was not recognized due to the difficulty in the study of type collections, scarcity of herbarium material and field observations. This applies especially to A. balkanicum (mentioned as a synonym of A. bertolonii subsp. scutarinum in Ball & Dudley 1993, and Jalas et al. 1996), whose type material was lost during 2 nd World War. Major distinguishing characters of A. decipiens are the tall and robust habit, the numerous stems arising from base, each with numerous sterile shoots bearing large, spathulate leaves with white-silvery abaxial surface; the cauline leaves are also larger and denser ( Fig. 9 ). In addition, flowering and fruiting in this species are considerably delayed compared with both O. chalcidica and O. smolikana subsp. glabra , regardless of altitude ( Fig. 4 ). Plants of O. decipiens from higher altitudes with lower stems and habit similar to that of the latter species (described as A. balkanicum f. depressum ) differ by the later flowering and the silicle morphology (see key to species).

Figures

FIGURE 1. Distribution of Odontarrhena in Albania; the range of O. chalcidica is given as light grey area, the other taxa as symbols (in legend); major serpentine outcrops are represented as dark grey spots and numbered from the north to the south. 1: Tropoja, 2: Krrabi, 3: Gomsiqe, 4: Puke, 5: Kukesi, 6: Lure, 7: Skenderbeu, 8: Bulqize, 9: Shebenik, 10: Shpati, 11: Vallamara, 12: Devolli, 13: Voskopoja, 14: Morava (after Bani et. al. 2017, modified).

FIGURE 3. The Albanian taxa of Odontarrhena in their natural environment:A) O. albiflora (Mt. Tathë); B) O. chalcidica (Pishkash); C) O. decipiens (Bulqizë); D) O. moravensis (Voskopojë); E) O. rigida (Mt. Shpat); F) O. smolikana subsp. glabra (Mt. Shebenik).

FIGURE 4. Phenological differences between taxa of Odontarrhena in Albania. Bars in light grey indicate the time interval between the decade in which at least 50% of the population is in flower and the decade when full flowering (100%) of the population occurs; bars in dark grey indicate the same interval for the fruiting process (50%-100% of the population with ripe fruits).

FIGURE 5. Micrographs of chromosome metaphase plates of: A) O. muralis (Romania, Deva, type locality, FI052171), 2n = 16; B) O. rigida, 2n =16; C) O. moravensis, 2n = 16; D) O. chalcidica (“A. markgrafii”, FI050423), 2n = 32 + 4B; E) O. chalcidica (type locality of A. elatius, FI052165), 2n = 32; F) O. decipiens (FI050443), 2n = 32+2B; G) O. smolikana subsp. glabra, 2n = 32; H) O. albiflora, 2n = 32. Scale bar = 5 μm. The B-chromosomes are indicated by arrow heads.

FIGURE 6. SEM micrographs of silicles of: A) O. albiflora (FI050840); B) O. chalcidica (neotype of A. markgrafii, FI050424); C) O. chalcidica (Greece, near Thessaloniki, isolectotype, FI010117); D) O. decipiens (FI050445); E) O. smolikana subsp. glabra (FI050433); F) O. moravensis (FI050441); G) O. muralis (Romania, Deva, type locality, FI050439); H) O. rigida (neotype, FI050434). Scale bar = 2mm.

FIGURE 7. SEM micrographs of trichomes on the median part of flowering stems of:A)O. albiflora (FI050840); B) O.chalcidica (neotype of A. markgrafii, FI050424); C) O. chalcidica (FI050422); D) O. decipiens (FI050445); E) O. smolikana subsp. glabra (FI050433); F) O. moravensis (FI050441); G) O. muralis (FI050439 type locality, Romania); H) O. rigida (neotype, FI050434). Scale bar = 500 μm.

FIGURE 9. O. decipiens. A) habit (flowering specimen and fruiting shoot); B) leaf of sterile shoot, showing its upper (on the left) and lower surface (right); C) lateral fruiting racemes; D) closed and open silicles of different size and shape. Original drawing by L. Cecchi (based on FI050442 and FI052160).