Allium undulatipetalum İ.Genç & N.Özhatay, 2015

Deniz, İsmail Gökhan, Genç, İlker & Sari, Duygu, 2015, Morphological and molecular data reveal a new species of Allium (Amaryllidaceae) from SW Anatolia, Turkey, Phytotaxa 212 (4), pp. 283-292 : 284-289

Allium undulatipetalum İ.Genç & N.Özhatay sp. nov. ( Figs. 1 , 2 , 3 , 4A–F )

Allium undulatitepalum is related to A. orientale Boiss. and it differs from the latter species by bulb globose (not ovoid); leaves 2–5 cm wide (not 0.8–2 cm wide); inflorescence semispherical (not fastigiate-semispherical); perigone segments after anthesis only reflexed (not reflexed and twisted outwards), 6–7 × 3.5–6 mm (not 5–6 × 2–3 mm), edges often undulate (not straight), white with green midvein (not pinkish white-pink with green or pinkish midvein); ovary purplish black in every stage (not green to turn purple); capsule globose (not elliptic to pear shaped); seed testa cells angularly round (not elliptic-oblong), well convex and with many prominent verrucae (not one or two-rowed with less prominent verrrucae).

Type:— TURKEY. C3 Antalya: Akseki, Çaltılıçukur Village, Salamut Plateau , calcareous stony and grassy slopes close to Cedrus libani forests, 36°53N, 31°55E, 1600 m, 9 May 1982, T. Ekim, M. Koyuncu s.n. (holotype ISTE 54419!; isotypes AEF!) .

Bulb globose, 2–3 cm in diameter, outer tunics blackish, disintegrating, inner tunics white. Scape 20–50 cm long above ground, slightly flexuous, cylindrical, 2–3 mm in diameter, basal part green or flushed carmine. Leaves 3–5, linearlanceolate, 2–5 cm wide and 7–25 cm long, green, with narrow white margin, smooth. Spathe most often completely split in 2–3 triangular parts, initially adpressed to the pedicels and later deflexed, faintly purplish with somewhat darker veins. Inflorescence semispherical at flowering time, subspherical in fruit, dense, 4–8.5 cm in diameter. Pedicels cylindrical, up to 2 cm long, almost equal, greenish or slightly carmine-flushed. Perigone nearly campanulate. Tepals obovate-orbiculate, obtuse at apex, often undulate, 3.5–6 mm wide and 6–7 mm long, white with green midvein. Filaments 4/5 as long as tepals, fleshy, basally united, triangular, white. Anthers yellow. Ovary depressed-globose with three furrows, dark purplish-black. Style cylindrical. Capsule globose, with three longitudinal furrows, 5–6 mm wide. Seeds ovate to broadly ovate, rugose, blackish, 3–4.2 mm long. The seed testa cells angularly round, convex, with many prominent verrucae and omega-like (Ω) undulations with moderate wavelength. Flowering in May–June, fruiting in June–August. 2n = 16.

Distribution, habitat and ecology:— Allium undulatitepalum is a local endemic restricted to the Salamut, Güzle and Çimi Plateaus in Antalya, southwestern Anatolia ( Fig. 1E ). It is a territory belonging to the Mediterranean floristic region. The new species colonizes only calcareous stony and grassy slopes close to Cedrus libani forests, between 1400–1700 m of elevation ( Fig. 1D ).

Etymology: — The species epithet is derived from its undulate tepals representing the one of the main characters which distinguish it from other similar species ( Fig. 2 ).

Molecular analysis: — The ITS sequences of A. undulatitepalum had ≥ 99% nucleotide identity with A. orientale and also ≥ 96% with A. nigrum according to the BLAST analysis. The A. orientale , A. multibulbosum and A. nigrum alliances are related to each other morphologically, but occur in different clades ( Fig. 3 ). Allium undulatitepalum is closely related to, but separate from A. orientale , occurring in the same clade. These molecular results supported the data obtained from the morphological studies with reliable bootstrap values. For ITS studies, alongside of A. undulatitepalum accessions some sequences of related taxa in the subgenus were retrieved from NCBI nucleotide database and compared. According to records of the NCBI and Fritsch et al. (2010), all A. orientale specimens were collected from Antalya (FM177375–177377) and some A. multibulbosum (FM177370) specimens from İzmir Province in Turkey, respectively. The accessions FM177368–177370 were cited as A. nigrum in Gurushidze et al. (2008), then these samples were considered as A. multibulbosum ( Fritsch et al. 2010, Fragman-Sapir & Fritsch 2011) ( Fig. 3 ). A. nigrum was included in molecular studies with specimens collected in Demre County, Antalya. Phylogenetic relationships among the studied Allium species, based on ITS sequences were in agreement with previous studies ( Gurushidze et al. 2008, Fritsch et al. 2010).

Karyotype: — In the present study, the somatic chromosome number of A. undulatitepalum was determined as 2n = 16 ( Fig. 4F ). The karyotype formula of the new species consists of twelve chromosome pairs with centromeres in median position and four pairs with centromeres in submedian position (12m + 4 smsat). The shortest chromosome pairs is 11.42 μm long, the longest is 15.13 μm and the haploid chromosome length is 107.1 μm. The basic chromosome number of the species in sect. Melanocrommyum is x = 8. The majority of Turkish species have 2n = 16 chromosomes ( Genç 2010). In the present study, the somatic chromosome number of A. undulatitepalum was determined to be 2 n = 16 ( Fig. 4F ). Allium orientale is similar to the new species in its diploid character 2n = 16 ( Bartolo et al. 1984, Pogosian & Seisums 1992, Genç et al. 2013) but different cytotypes of A. orientale were reported as being triploid or tetraploid 2n = 24, 32 from Cyprus ( Tzanoudakis 1999). In the present study, the karyotype formula of A. undulatitepalum was determined as 12m + 4 smsat. Conversely, according to Genç et al. (2013), A. orientale has 10m + 6 smsat chromosomes. Allium undulatitepalum shows also chromosomes larger than A. orientale .

Pollen characters: — A. undulatitepalum and A. orientale have monosulcate pollen grains and the exine ornamentation is perforate, striate and rugulate in both species ( Fig. 4E, 4G ). Also, the pollen shape (based on LA/SA ratio) is prolate in distal view, and circular in polar view ( Fig. 4D ).According to LM measurements, A. undulatitepalum : long axis (28.3–)32.66(–36.75) ± 1.96 μm, short axis (14.7–)18.54(–24.15) ± 2.32 μm; A. orientale : long axis (28.35–)32.57(–36.75) ± 1.74 μm, short axis (16.8–)20.05(–23.1) ± 1.64 μm. Results of the SEM and LM studies indicate that sulcus extends from distal face to polar points but does not reach the proximal face and that sulcus ends are rounded in both species. Palynological characters and measurement data from A. undulatitepalum and A. orientale are very similar and are consistent with the results of previous studies conducted on different species of the sect. Melanocrommyum ( Özhatay & Koçyiğit 2009, Deniz et al. 2013).

Seed characters: — The seeds are ovate to broadly ovate, blackish and rugose in A. undulatitepalum ( Fig. 4A ). When compared from an identical midpoint position on the dorsal surface of the seeds, the dominant shape of A. undulatitepalum testa cells is angularly rounded while those of the related A. orientale are elliptic-oblong ( Fig. 4C, 4I ). The new species presented in the study has depressed, omega-like (Ω) undulated anticlinal walls and strongly convex, prominently well developed verrucate periclinal walls ( Fig. 4B, 4C ). On the other hand, A. orientale has one or two rowed and less prominent verrucae on the periclinal walls ( Fig. 4H, 4I ). It is reported that the seeds of subg. Melanocrommyum are black, generally ovate and somewhat larger than those of most Allium species belonging to other groups ( Neshati & Fritsch, 2009). Many species of subg. Melanocrommyum share convex periclinal walls with verrucate sculptures and S-, U- and omega-like (Ω) undulated anticlinal walls. Transitions also occurred between these types ( Fritsch et al. 2006, Neshati & Fritsch 2009, Celep et al. 2012).

Taxonomic relationships: — Allium nigrum and A. orientale are widely distributed and variable members of the sect. Melanocrommyum . They were long considered as a taxonomically problematic species. The neotypification of A. nigrum was defined by Seisums (1998), some variants occurring in Israel were described as new species separate from A. nigrum and A. orientale by Fragman-Sapir & Fritsch (2011), and useful morphological qualitative-quantitative characters separating A. nigrum and A. cyrilli were presented by Peruzzi et al. (2012). As a result of these studies, A. multibulbosum was accepted as a valid species different from A. nigrum again.Also, it was understood that A. orientale has a more narrow distribution in Turkey. Due to this confusion, some herbarium specimens of A. undulatitepalum were identified as A. nigrum or A. orientale in the past. As a result of our studies and investigations, A. undulatitepalum is morphologically related to A. orientale . The detailed morphologic differences between A. undulatitapalum and A. orientale are presented in Table 1. At the first glance, A. undulatitepalum is clearly distinguishable from A. orientale by broad and undulate tepals, the dark purplish-blackish ovarium in every stage, the semispherical and dense inflorescence, and by broad leaves.

Additional specimens examined of Allium undulatitepalum (paratyes):— TURKEY. C3 Antalya: Manavgat, Güzle Upland, meadows, 1400 m, 7 May 1982, T. Ekim, M. Koyuncu 5267b (ISTE 54418!); Akseki, Çaltılıçukur Village, Salamut Upland, 1600 m, 9 May 1982, T. Ekim, M. Koyuncu 5285a (ISTE 54420!);Akseki, Çimi Village, Çimi Upland, Aldürbe area, 1500–1700 m, 22 May 1983, M. Koyuncu 6035, S. Erik (AEF 19392!); Akseki, Çaltılıçukur Village, Salamut Plateau, calcareous slopes 1640 m, 7 May 2008, İ. Genç et al. 1182 (ISTE 91484!); ibidem, 13 June 2012, H. Sümbül, C. Aykurt, İ.G. Deniz 4578 (AKDU!; NCBI Accession No: KP881225); ibidem, 14 May 2014, İ.G. Deniz 5545 (AKDU!; NCBI Accession No: KP881223); ibidem, 5546 (AKDU!; NCBI Accession No: KP881224), ibidem, 5547 (AKDU!); ibidem, 5 August 2014, İ.G. Deniz 5827 (AKDU!).

Figures

FIGURE 1. Allium undulatitepalum: A. Top view of inflorescence, B. Habit, C. Side view of inflorescence, D. Habitat, E. Distribution map of A. undulatitepalum (Ì) and A. orientale (z) (Photos: İ.G.Deniz & İ.Genç).

FIGURE 2. Shape of tepals, filaments, and capsule valves of Allium undulatitepalum and A. orientale (all scale bars 1 mm).

FIGURE 3. Maximum likelihood tree of ITS sequences with bootstrap values from 6 Allium species (GeneBank accession numbers are given behind the Allium names, abbreviations starting with “KP” are listed in the ‘specimens examined’ section).

FIGURE 4. Allium undulatitepalum (A–F) and A. orientale (G–I): A. Seed, B. Periclinal walls, C & I. Testa cells and undulation of anticlinal walls, D. Pollen grains, E & G. Exine ornamentation, F. Mitotic metaphase chromosomes 2n = 16, H. Periclinal and anticlinal walls (Specimens examined:A–C Deniz 5827, D–E Deniz 5547, F Genç 1182, G Genç 1360, F Genç 1363).