Turk J Bot (2016) 40: 402-411 © TÜBİTAK doi:10.3906/bot-1508-2 Turkish Journal of Botany http://journals.tubitak.gov.tr/botany/ Research Article Alyssum amasianum (Brassicaceae), a new species from North Anatolia, Turkey 1, 2 3 Osman KARABACAK *, Ahmet DURAN , Mustafa ÇELİK Department of Biology, Polatlı Faculty of Science and Literature, Gazi University, Ankara, Turkey 2 Department of Biology, Faculty of Science, Selçuk University, Konya, Turkey 3 Department of Biotechnology, Faculty of Science, Selçuk University, Konya, Turkey 1 Received: 03.08.2015 Accepted/Published Online: 05.02.2016 Final Version: 07.06.2016 Abstract: Herein a new species, Alyssum amasianum, is described and illustrated from North Anatolia. The new species is very similar to A. hirsutum subsp. caespitosum but differs in the shape and indumentum of leaves, trichome type of the fruits, and margin of the seeds. The ecology, biogeography, and conservation status of the new species are discussed. The leaf, fruit, and seed surface micromorphology of A. amasianum, A. hirsutum subsp. caespitosum, A. hirsutum subsp. hirsutum, A. strigosum subsp. strigosum, and A. xanthocarpum were also examined by scanning electron microscope. Key words: Alyssum, Alysseae, Cruciferae, SEM, Turkey 1. Introduction The family Brassicaceae is distributed worldwide across all continents except for Antarctica (Koch and Kiefer, 2006). It consists of 49 tribes, about 321 genera, and 3660 species (Al-Shehbaz, 2012). The Brassicaceae, or mustard family, is easily distinguished from other flowering plant families with floral and fruit morphology by the cruciform corolla, tetradynamous stamens, and a siliqua often with a septum (Koch et al., 2012). The tribe Alysseae is composed of 24 genera and about 277 species, 114 of which are placed in Alyssum (Warwick et al., 2006; Al-Shehbaz, 2012; Španiel et al., 2015). The genus Alyssum is distributed primarily in Europe, Southwest Asia, and northern Africa (Al-Shehbaz, 1987; Al-Shehbaz et al., 2006). However, a few species are dispersed in Central Asia, Siberia, and North America (Dudley, 1964a, 1964b). In a recent taxonomic and phylogenetic study of the genus (Španiel et al., 2015), two sections of Alyssum were raised to genus level, i.e. Alyssum sect. Odontarrhena and A. sect. Meniocus. In addition, A. homalocarpum and A. antiatlanticum were described in the new genus Cuprella Salmerón-Sánchez, Mota & Fuertes. Based on the latest taxonomic situation (Španiel et al., 2015) and having this new species addition, the number of Alyssum species has reached 54 in Turkey, 25 of which are endemic. This further stresses that Turkey is the main center of diversity for the genus (Dudley, 1965; Davis et al., 1988; Yıldırımlı, 2000; Mutlu, 2012; Özhatay et al., 2013). After separation * Correspondence: okarabacak35@gmail.com 402 of the genus Alyssum, there are 38 species in Odontarrhena and 7 species in Meniocus in Turkey. In the latest taxonomic position, Alyssum s.str. includes two clades; one of them is taxa of A. sect. Alyssum that are mostly annual and perennial and the other one is A. sect. Gamosepalum, a few perennials of A. sect. Alyssum, and annual A. dasycarpum (Resetnik et al., 2013; Španiel et al., 2015). The members of the section Alyssum are characterized by monomorphic sepals free; unilaterally or bilaterally winged, dentate, or appendaged filaments; dehiscent monomorphic or dimorphic indumentum or, rarely glabrous and valves equally or unequally inflated fruits; and biovulate loculi (Dudley, 1964a, 1964b). The section Alyssum contains 43 taxa in Turkey, 19 of which are endemic (Dudley, 1965; Orcan and Mısırdalı, 2000; Orcan and Binzet, 2006, 2009). Interesting specimens of Alyssum belongs to the sect. Alyssum were collected during field trips in Borabay village (Amasya, North Anatolia) in 2013 and 2014. They were found distinct from all known species and most similar to A. hirsutum subsp. caespitosum. The present study aimed to describe a new Alyssum species from Turkey based on morphology and micromorphology of leaf, fruit, and seed features. 2. Materials and methods The specimens were cross-checked with the keys from Flora Iranica (Hedge, 1968), Flora of Iraq (Townsend, KARABACAK et al. / Turk J Bot 1980), Flore du Liban and De La Syrie (Bouloumoy, 1930), Flora of USSR (Bush, 1970), Flora of Cyprus (Meikle, 1977), and Flora Europaea (Ball and Dudley, 1964) and additional literature sources were investigated (Dudley, 1965; Davis et al., 1988; Orcan and Mısırdalı, 2000; Yıldırımlı, 2000; Orcan and Binzet, 2006, 2009; Mutlu, 2012; Özhatay et al., 2013). The specimens were cross-checked with the material housed at various herbaria (E, B, W, K, GAZI, EGE, HUB, KNYA, and ANK). For the scanning electron microscope (SEM) investigations, leaves, fruit, seed, and pollen were directly mounted on prepared stubs and coated with gold. Photographs were taken using a JEOL-JSM-6490 LV SEM after being coated with a Hummle VII Sputter Coater. The averages and standard deviations of the measurements were calculated. The terminology given by Stearn (1996), Prentince (1979), Barthlott (1981), and El Naggar (2005) was used for the description of the SEM aspects. The terminology used for pollen description has been followed as suggested by Erdtman (1952). Alyssum amasianum Karabacak & A.Duran sp. nov. (Figures 1–2). Type: Turkey. A5 Amasya: Taşova, Borabay village to Başyurt Yayla, 9 km, 1382 m, serpentine slopes, roadsides, 16.08.2013, 40°48′011″N 36°07′005″E, A.Duran 9733 & O.Karabacak (holotype: KNYA; isotypes: GAZI, ANK). Diagnosis: Alyssum amasianum is similar to A. hirsutum subsp. caespitosum. It mainly differs from A. hirsutum subsp. caespitosum because it has petals 5 × 1.5–2 mm, retuse or slightly emarginate (not 4.5 × 1 mm, deeply emarginate); long filament appendage connate for about half of their length (not dilated at the base); short filament appendage apex acute (not tridentate); fruiting pedicel 3–4 mm (not 5–6 mm); fruit 8–9 mm and stellate trichomes with mostly 8–10 subequal adpressed rays and bigger tuberculate trichomes with 3–6 unequal rays (not 5–6 mm, sparsely covered with smaller stellate trichomes with 8–10 subequal adpressed rays and much longer spreading simple tuberculate trichomes); style 3–4 mm (not 2–2.5 mm); seed dark brown, 3–3.5 × 2–2.5 mm (not pale brown, 2 × 1.5 mm). Description: Annual or rarely biennial herb, up to 7 cm long, mostly branched below. Stem erect and leaves densely covered with unequally branched stellate trichomes. Leaves oblanceolate, entire; lower surface of leaves densely stellate, 8–12 subequal rays with tuberculate; lower cauline leaves 15–20 × 6–10 mm; upper cauline leaves 6–15 × 2–8 mm. Inflorescence raceme, 5–10-flowered, condensed at anthesis, fruiting raceme slightly elongated. Sepals narrowly ovate, deciduous, 4 × 1.5–2 mm, with a thin scarious margin, outer surface sparsely stellate and branched tuberculate trichomes. Petals yellow, 5 × 1.5–2 Figure 1. Alyssum amasianum (photo by A Duran). mm, retuse or shortly emarginate, outer surface stellate trichome; limbs narrow, about twice as broad as claw. Long filaments 4 mm, appendages connate for about half of their length, acute; short filaments 3.5 mm, appendage 1 mm joined at the base then c. 1.5 mm free, acute. Anthers 1–1.25 mm, yellow. Style 3–4 mm long and dilated at the base, stellate trichome, stigma truncate. Fruiting pedicels 3–4 mm, erect to patent, densely stellate pubescent. Fruits 8–9 × 6–8 mm, suborbicular in outline, indumentum consisting of stellate trichomes with mostly 8–10 subequal subpatent rays and bigger tuberculate trichomes with 3–6 unequal rays, c. 2 mm long, loculi 2 ovulate. Seeds dark brown, elliptic, 3 × 2 mm, wing c. 0.2–0.3 mm wide, slightly undulate. Fl. & Fr. August and September. Distribution and ecology: Alyssum amasianum is an endemic species that is confined to the Başyurt plateau in Borabay village (Amasya Province). It is a Euro-Siberian element. Alyssum amasianum grows in open forest of Pinus nigra Arn. together with Rubus canescens DC. var. canescens, Scutellaria salviifolia Bentham, Sedum pallidum M.Bieb. var. pallidum, Hypericum perforatum L., Astragalus plumosus Willd. var. nitens (Freyn & Bornm.) Chamb. & Matthews, Anthemis kotschyana Boiss. var. kotschyana, Salvia tomentosa Miller, Dryopteris filix-mas (L.) Schott, Linaria sp., and Campanula sp. International Union for Conservation of Nature (IUCN) red list category: Alyssum amasianum is known from the type locality and this area is smaller than 2.5 km2 (Criteria B1). Animals are overgrazing in this area. Because of this, the species is under threat. Groundwater and surface streams are seasonally overflowing because of heavy rainfall. Hence, their destruction is leading to the reduction in the number of plants (Criteria A). The mature individual members of the population constitute 250 specimens of flowers (Criteria C2). Therefore, it should be considered critically endangered (CR) according to the IUCN Red List Criteria (IUCN, 2010). 403 KARABACAK et al. / Turk J Bot Figure 2. Alyssum amasianum (from the holotype). a- habit, b- long filament, c- short filament, d- petal, e- fruit, f- seed. Alyssum hirsutum subsp. caespitosum: g- long filament, h- short filament, i- petal, j- fruit, k- seed. 3. Discussion Alyssum amasianum is very similar to A. hirsutum subsp. caespitosum. However, there are some significant morphological and ecological differences between them (Table). In the tribe Alysseae, trichome morphology has been usually used as a diagnostic character (Dudley, 1964b; AlShehbaz, 1987; Ančev, 1991; Ančev and Goranova, 2006). Alysseae is characterized by having most often adpressed or subpatent stellate indumentum (trichomes with 3 or more rays originating from one point), sometimes an admixture of simple trichomes can be observed, and rarely trichomes are malpighiaceous (Warwick et al., 2008). On fruit surface, Alyssum amasianum has tuberculate and stellate dimorphic trichomes. Tuberculate trichomes stalked with 3–6 unequal rays with tuberculate surface and 404 smooth base. Stellate trichomes with subequal 8–10 rays, mostly straight and tapering on the tip. On the other hand, both A. hirsutum subsp. caespitosum and A. hirsutum subsp. hirsutum have simple, unequally bifurcate tuberculate (rare), and stellate trichomes that have 8–10 subequal rays. Both Alyssum strigosum subp. strigosum and A. xanthocarpum have 2 equally bifurcate tuberculate trichomes. However, the first one has stellate trichomes with 6 subequal rays; the latter one has 8–10 subequal rays (Figure 3). In leaf indumentum, A. amasianum has stellate trichomes with 8–12 subequal rays, but A. hirsutum subsp. hirsutum has stellate trichomes with 6–8 subequal rays. Alyssum hirsutum subsp. caespitosum, A. strigosum subsp. strigosum, and A. xanthocarpum have stellate trichomes with 6 subequal rays and tuberculate trichomes with 2 (–4) arms and smooth surface (Figure 4). KARABACAK et al. / Turk J Bot Table. Diagnostic characters of Alyssum amasianum and A. hirsutum subsp. caespitosum. Characters Alyssum amasianum Alyssum hirsutum subsp. caespitosum Stems to 7 cm tall, branched 3–4 cm tall, unbranched Leaves lower obovate and upper oblanceolate, sparsely oblanceolate, densely stellate trichome, lower 15–20 × stellate and 2 (–4) arms tuberculate trichome, 8–10 × 6–10 mm and upper 6–15 × 2–8 mm 2–3 mm Sepals 4 × 1.5–2 mm, deciduous, outer surface sparsely stellate and branched tuberculate trichomes 3 × 1 mm, deciduous, outer surface sparsely stellate trichomes Petals 5 × 1.5–2 mm, retuse or shortly emarginated 4.5 × 1 mm, deeply emarginated Long filaments 4 mm and appendages connate for about half of their 3.5 mm dilated at the base length Short filaments 3.5 mm and appendage free, acute c. 3 mm and appendage free, tridentate Anthers 1–1.25 mm 0.5–0.6 mm Fruit pedicels 3–4 mm 5 mm Fruits 8–9 mm, stellate and 3–6 unequal rays with tuberculate trichomes 5–6 mm, sparsely stellate and simple or rarely unequally bifurcate tuberculate trichomes Styles 3–4 mm 2–2.5 mm Seeds dark brown, 3–3.5 × 2–2.5 mm, margin irregular pale brown, 2 × 1.5 mm, margin smooth Alyssum amasianum, A. hirsutum subsp. caespitosum, and A. hirsutum subsp. hirsutum have tricolpate apertures with reticulate tecta and perprolate pollen grains (Figure 5). Based on the exine sculpturing pattern, size, and shape of the pollen grains, there are no significant differences among the taxa. Alyssum amasianum has dark brown seeds with irregular margins. On the other hand, A. hirsutum subsp. caespitosum, A. hirsutum subsp. hirsutum, A. strigosum subsp. strigosum, and A. xanthocarpum have pale brown seeds with smooth margins. Based on the epidermal cell patterns of testae obtained from SEM, three different surface sculpturing patterns are seen: reticulate, reticulate– papillate, and papillate. While A. amasianum, A. hirsutum subsp. hirsutum, and A. xanthocarpum have reticulate sculpturing patterns, A. hirsutum subsp. caespitosum has a reticulate–papillate and A. strigosum subsp. strigosum has only a papillate sculpturing pattern. While Alyssum amasianum and A. xanthocarpum have similar anticlinal cell boundaries (i.e. flat and raised), their periclinal cell walls differ from one another. For example, A. amasianum has flat, but A. xanthocarpum has convex periclinal cell walls. Both A. hirsutum subsp. caespitosum and A. hirsutum subsp. hirsutum have raised anticlinal cell boundaries; however, they differ from one another by their periclinal cell wall development as A. hirsutum subsp. caespitosum has a papillate pattern and A. hirsutum subsp. hirsutum has concave periclinal cell walls with some intercellular area. In addition, A. strigosum subsp. strigosum has channeled anticlinal cell boundaries and raised-papilate periclinal cell walls (Figure 6). Alyssum amasianum is located in a transitional zone between the Euro-Siberian and Irano-Turanian phytogeographical regions. However, A. hirsutum subsp. caespitosum and A. hirsutum subsp. hirsutum are distributed in the Irano-Turanian phytogeographical region. In addition, there is a phenologic difference between A. amasianum, A. hirsutum subsp. caespitosum, and A. hirsutum subsp. hirsutum. While Alyssum amasianum flowers in August, the latter two taxa flower from May to June. 3.1. Additional specimens examined (paratypes): Turkey: A5 Amasya: Taşova, Borabay village to Başyurt Yayla, 9 km, 1363 m, 26.08.2014, 256966 N, 4520721 E (UTM), O.Karabacak & M.Çelik 9294 (GAZI, KNYA). 3.2. Additional specimens examined (similar taxa): A. hirsutum subsp. caespitosum: Turkey: C2 Muğla: Köyceğiz, Sultaniye, Günlük bucağı çevresi, 15 m, aluviyal düzlük, 15.04.1991, A. Güner 8610 & M. Vural, H.Duman, A.Dönmez, B.Mutlu (HUB); C2 Muğla: Köyceğiz Sultaniye arası, Günlük bucağı çevresi, 10 m, metamorfik çakıllı arazi, aluviyal düzlük, 17.03.1991, A.Güner 8209 & H.Duman, H.Şağban (GAZI); C2 Muğla: Köyceğiz, Sultaniye, Kersele köyü, 20 m, serpentin taşlık arazi, 14.04.1992, A.Güner 10426 & H.Duman, A.Dönmez, H.Şağban (HUB); C2 Muğla: Köyceğiz, Ekincik köyü, İskele-Kurşuncuk feneriKaraçay arası, 0–30 m, kayalık yamaçlar, kızıl çam ormanı, 05.04.1991, A.Güner 8293 & H.Duman, H.Şağban (HUB); 405 KARABACAK et al. / Turk J Bot Figure 3. Scanning electron micrographs of the fruits. Alyssum amasianum: a- indumentum of fruit, b- stellate trichomes, ctuberculate trichomes; A. hirsutum subsp. caespitosum: d- indumentum of fruit, e- stellate trichomes, f- tuberculate trichomes; A. hirsutum subsp. hirsutum: g- indumentum of fruit, h- stellate trichomes, i- tuberculate trichomes; A. strigosum subsp. strigosum: jindumentum of fruit, k- stellate trichomes, l- tuberculate trichomes; A. xanthocarpum: m- indumentum of fruit, n- stellate trichomes, o- tuberculate trichomes. 406 KARABACAK et al. / Turk J Bot Figure 4. Scanning electron micrographs of the leaf trichomes. Alyssum amasianum (a–c); A. hirsutum subsp. caespitosum (d–f); A. hirsutum subsp. hirsutum (g–i); A. strigosum subsp. strigosum (j–l); A. xanthocarpum (m–o). 407 KARABACAK et al. / Turk J Bot Figure 5. Scanning electron micrographs of the pollens. Alyssum amasianum: a- general shape of pollen, b- pollen ornamentation; A. hirsutum subsp. caespitosum: c- general shape of pollen, d- pollen ornamentation; A. hirsutum subsp. hirsutum: e- general shape of pollen, f- pollen ornamentation. C4 Konya: Beyşehir, 1 km E of Nazım Bey tepesi, 1150 m, stony pastures, among Astragalus, 29.04.1961, H.Demiriz 4459 (E). A. hirsutum subsp. hirsutum: Turkey: B5 Niğde: Aksaray, Ihlara vadisi, 17.06.1986, S.Erik 3844 & İ.Verten (HUB); B3 Ankara: Polatlı, Polatlı’nın 18 km batısı, Acıkır mevkii (topçu atış okulu), 850 m, korunmuş step, 16.05.1991, T.Ekim 3793 & Z.Aytaç, H.Duman (GAZI); B5/6 Yozgat: Pinus sylvestris orman açıklığı, c. 1400–1500 m, kalkerli alan, 04.06.1980, T.Ekim 4953 (ANK); B3 408 Akşehir: Sultan dağları, Cankurtaran köyü karşısı, 1600– 1700 m, Y.Akman 13830 (ANK); B4 Ankara: ElmadağKırıkkale, approx. 7–10 km W of Kırıkkale, on S side of river, serpentine rock slope above highway, 21.05.1996, R.D.Reeves 1678 & U.Krämer (E). A. strigosum subsp. strigosum: Turkey: B4 Ankara: Polatlı, Mehmet Akif mahallesi çevresi, step, 17.04.2015, 39°35′922″N 32°08′331″E, 923 m, G134 (KNYA). A. xanthocarpum: Turkey: B9 Van: Erciş, Şehirpazarı köyü çevresi, step, 26.06.2005, 39°13′935″N 43°25′725″E, KARABACAK et al. / Turk J Bot Figure 6. Scanning electron micrographs of the seeds. Alyssum amasianum: a- seed shape, b- seed ornamentation; A. hirsutum subsp. caespitosum: c- seed shape, d- seed ornamentation; A. hirsutum subsp. hirsutum: e- seed shape, f- seed ornamentation; A. strigosum subsp. strigosum g- seed shape, h- seed ornamentation; A. xanthocarpum i- seed shape, j- seed ornamentation. 409 KARABACAK et al. / Turk J Bot 2250 m, OK 3722 (KNYA); Taşkapı köyü ile Ganissipi Y. arası, step, 29.06.2006, 39°18′158″N 43°26′648″E, 2200 m, OK 4854 (KNYA). 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