Odontarrhena bertolonii subsp. scutarinum,

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 : 8-10

2. O. chalcidica ( Janka 1872: 175) Španiel, Al-Shehbaz, D. A.German & Marhold in Španiel et al. (2015: 2484).

(≡) Alyssum chalcidicum Janka. ≡ A. murale subsp. chalcidicum (Janka) Contandriopoulos (1969: 323) .

Lectotype (designated here):— GREECE. “in montibus inter Hierisso et Gomati Chalcidices; in arvis atque locis silvaticis ad agrorum margines”, 04/08/1871, V. Janka , WU1938-0001420 , WU!

Isolectotypes:—“in montibus inter Hierisso et Gomati Chalcidices; in arvis atque ad agrorum margines”, Herb. Haussknecht, JE00003077 , JE !; “in montibus inter Hierisso et Gomati Chalcidices”, B 10 0058222, B !; GH00018476 , GH !; Herb. Churchill, K000484603 , K !; Herb. Reichenbach, W1889-0038856 , W !; “In campi ad silvarum margines inter pag. Hierisso et Gomati chalcidices”, FI010117 , FI !; ex Herb. Groves, FI018015 , FI !; JE00003078 , JE !; Herb. Halácsy, WU0033143 , WU !

(=) A. markgrafii O.E.Schulz in Markgraf (1926: 422), syn. nov. ≡ O. markgrafii (O.E.Schulz) Španiel, Al-Shehbaz, D. A.German & Marhold in Španiel et al. (2015: 2485).

Neotype (designated here):— ALBANIA. “ prefettura di Elbasan, distretto di Librazhd, pietraia calcarea presso la vetta del Gur i Pishkashit, 1080 m, 41° 5’44.53”N, 20°31’10.12”E ”, 14/07/2016, I. Bettarini, L. Cecchi, A. Coppi & F. Selvi, FI050424 , FI!

Isoneotypes:—B!, Herb. Cecchi no. 3335!

The original collections (syntypes) of A. markgrafii (“ Çermenika: Gur i Pishkashit bei Qukës, 1100 m, Kalk, den Gipfel bedeckend”, 19/06/1924, F. Markgraf no. 773; “ Shpat: Shelcan, offene Serpentinschuttflur, massenhaft, 800 m ”, 01/06/1924, F. Markgraf no. 501) were kept in B but were destroyed and no duplicates could be traced in the other European herbaria hosting Markgraf’s materials ( E, TIR, Z). We collected material from both type localities above ; the neotype selected here comes from the first one and fits the original description of the species ( O. E. Schulz in Markgraf 1926) .

(=) A. janchenii Nyár. in Novák (1927: 109), syn. nov.

Lectotype (designated here):— ALBANIA. “Nord-Albanien: Umgebung von Shkodra. Abhänge des Kleinen Bardanjolt, Serpentin”, 08/06/1916, E. Janchen , WU0073381 , WU!

Isolectotype:—WU0073382, WU!

Other original material:— ALBANIA. “in aridis serpentinis infra Scutari et Renci”, 11/06/1897, A. Baldacci, Iter Albanicum Quintum no. 241, BM000750162 [flowering specimen only], BM !; K000484615 [flowering specimen only], K !; WU00679672 , WU! “ Im Schotter des Nerfuša-Baches bei dessen Einmündung in den Drin (Östl. Von Skodra)”, 18/06/1916, E. Janchen, WU0033158 , WU !

Although the name was first published in a paper on the flora of Serbia ( Novák 1927), it was explicitly based on Nyárády’s diagnosis of Albanian collections; in a later publication Nyárády himself (1928: 93–96), mentioned the provenance of the original material by Janchen, which accounts for the specific epiteth.

(=) A. bertolonii subsp. scutarinum Nyárády (1928: 101) , syn. nov. ≡ O. bertolonii subsp. scutarina (Nyár.) Španiel, Al-Shehbaz, D. A.German & Marhold in Španiel et al. (2015: 2485).

Lectotype (designated here):— ALBANIA. “in aridis serpentinis / infra Scutari et Renci”, 09/08/1897, A. Baldacci, Iter Albanicum Quintum no. 241, WU0033163 , WU!

Isolectotypes:— BM000750162 [fruiting specimen only], BM !; K000484615 [fruiting specimen only], K !

(=) A. chlorocarpum var. subellipticum Nyárády (1928: 121) , syn. nov.

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

(=) A. punctatum f. divergens Nyárády (1928: 86) , syn. nov.

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

(=) A. elatius Meyer (2011: 64) , syn. nov. ≡ O. elatior (F.K.Mey.) Španiel, Al-Shehbaz, D. A.German & Marhold in Španiel et al. (2015: 2484; ‘elatius’).

Holotype:— ALBANIA. “ Korça, Mali i Moravës, bei Drenova , ca. 1100–1200 m, Serpentin. 12/09/1961, F. K. Meyer, Flora Albanica no. 6148, JE00016684 , JE!

Flowering stems erect 10–60(100) cm, thick and somewhat rigid in fruit. Basal sterile shoots rarely numerous when present, with leaves up to 20 × 6 mm, spathulate, obtuse, usually gutter-like folded, greenish above with a single layer of hairs, greyish below for two layers of hairs with 6–20 rays, 0.3–0.6 mm across. Leaves on flowering stems linear to narrowly spathulate, longer than those on sterile shoots, often incurved and with smaller leaves at the axil. Inflorescence a broad, compound corymb, 3- to 4-branched. Partial racemes usually with 10–20 fruits. Fruiting pedicels delicate, erecto-patent, the lowermost sometimes slightly flexuous. Sepals 1.5–2 × 0.7–1.0 mm. Petals 2–3 mm. Style 0.8–1.5 mm. Siliculae 2.5–4.3 mm long, elliptic to suborbicular, retuse, rounded or subacute at apex, obtuse to rounded at base, symmetrical; valves flat to slightly inflated, not undulate, indistinctly veined; hairs on valves absent or sparse, never overlapping, 0.15–0.3 mm across, with 8–16 appressed rays. Seeds 1.5–2.0 mm long, including a wing 0.3–0.5 mm wide. Figs. 3B , 6B, C , 7B, C , 8 .

Phenology. Flowering from June to July, fruit ripening from July to August ( Fig. 4 ).

Chromosome number. 2n = 32 + 0–4B ( Fig. 5D, E ). Counts were performed on plants from five populations: near Pogradec (FI050416), Mt. Shpat near Elbasan (one of the two type localities of O. markgrafii, FI 050423), Mt. Shebenik near Skënderbej (FI050428), Renc near Skhodër (type locality of O. bertolonii subsp. scutarina and A. janchenii, FI 050420) and Mt. Morave near Drenova, type locality of O. elatior (FI052165; Fig. 5E ). Supernumerary B-chromosomes were observed in the population from Mt. Shpat ( Fig. 4D ) and probably from Pogradec. Plants of O. muralis (Waldst. & Kit.) Endl. from the type locality in Romania (Deva), investigated here for the first time, were instead diploid with 2n = 16 ( Fig. 5A ). Previous counts in the A. murale complex from other Balkan countries reported both 2n = 16 and 2n = 32 ( Jalas et al. 1996).

Distribution and ecology. Widely distributed throughout Albania, especially in the northern, central and eastern parts, less frequent in the south ( Fig. 1 ). It grows on ultramafic rocks and other peridotites, less frequently on schists and flysch, from 50 to 1300 m a.s.l. It is a synanthropic plant clearly preferring disturbed habitats such as ruderal sites, mining areas, road margins, cultivated and abandoned lands, dry pasturelands and fields.

Nickel accumulation. Shoot Ni-levels in this species were variable, though always well above 1000 μg g-1 dw when in plants from serpentine soil. The maximum concentration, detected in plants from Pogradec, was over five times higher than the minimum value detected in plants from mixed serpentine-schist soil south of Erseke ( Table 1).

Comments. Odontarrhena chalcidica belongs to the species complex of O. muralis and was included until recent times within the latter, as a subspecies (Contandriopoulos 1967) or as “variant” ( Ball & Dudley 1993, Jalas et al. 1996, Cecchi et al. 2010). At present, however, it is mostly recognized as a separate species and indicated from most of the southern Balkan countries: Serbia, Kosovo, Macedonia, Greece and possibly Bulgaria ( Hartvig 2002, Marhold 2011, Spaniel et al. 2015). Examination of O. muralis in the type locality in Romania (citadel of Deva, FI052171) confirmed that O. chalcidica differs by the less densely pubescent stems, the obtuse leaves, the smaller flowers, the more slender fruiting pedicels, and especially the silicles with at most few, scattered and minute trichomes (vs. dense in O. muralis , Fig. 6G ). It also differs ecologically in its clear preference for soils with high levels of Mg, especially serpentine, while O. muralis is usually found on limestone soils or other substrates (including volcanic rocks as in the type locality), but not on serpentine (see also Hartvig 2002).

As in the Greek populations, the Albanian plants of O. chalcidica show very broad variations in size, branching degree of flowering stems, shape and hairs on the silicle valves, even at the individual level. At a close examination of specimens from their respective type localities, both O. markgrafii and O. bertolonii subsp. scutarina resulted clearly within the range of phenotypic variation of this species and are here included in its circumscription, in contrast with most of the literature (Ball & Dudley 1963, Jalas et al. 1996, Spaniel et al. 2015). When tested on numerous individuals complete of all parts, the characters used to separate these taxa in treatments based on herbarium material appeared artificial. The former species ( O. markgrafii ) is reported to differ from O. muralis by substantially the same characters of O. chalcidica (more slender habit, smaller petals, silicles glabrous or with sparse hairs). Schulz (in Markgraf 1926: 422) indicated that O. markgrafii differs from typical O. chalcidica by the smaller flowers and the trichomes on the stem, which are oblong and with less numerous (6–10), longer rays (vs. circular “disciform” hairs with 10–20, short rays, fig. 7B, C). However, we observed that these two morphologies are connected by a continuous series of intermediate types and most of these are found in plants from localities in N and C Albania, including the typical ones of O. markgrafii . This shows the scarce taxonomic value of this character and supports inclusion of O. markgrafii in O. chalcidica , as proposed by Hayek (1927). Odontarrhena bertolonii subsp. scutarina was mainly distinguished from O. muralis in view of the narrower silicles with elliptic shape, subacute at the apex. For this character the Albanian plants were referred to the Italian endemic O. bertolonii (Desv.) Jord. & Fourr. , with similar silicles, though Nyárády himself (1930, 1949) correctly excluded this affinity in his last papers. Indeed, plant habit, leaf shape and branching pattern of the inflorescence as displayed in the field are clearly different in these two taxa, which supports the phylogenetic distance between them revealed by molecular data ( Cecchi et al. 2010). Based on these, the Albanian plants belong to the O. muralis clade, while O. bertolonii is included in a different clade together with other W Mediterranean endemics ( Cecchi et al. 2013). Odontarrhena elatior (‘elatius’) was described based on a single fruiting collection from the region of Korça ( Meyer 2011). Close examination of material from the type locality on the Morava massif (Appendix 1) showed that the diagnostic characters mentioned in the protologue are still in the range of variation of O. chalcidica . In some plants, the elliptic, subacute silicles and the numerous sterile shoots with small leaves, uncommon in typical O. chalcidica , suggest possible hybridization with O. moravensis , the other taxon in this area.

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 8. O. chalcidica: A, F) habit (flowering and fruiting specimens of different stature); B) cauline leaves of flowering (left) and sterile (right) shoots, showing their upper and lower surface on the left and on the right, respectively; C) lateral fruiting raceme; D) silicle in lateral, inner and outer view. Original drawing by L. Cecchi (based on FI050844 and FI050882).