Introduction
The grass family (Poaceae), with about 12,000 species and 780 genera, ranks as the second largest monocot family and the fifth largest plant family in the world (Clayton and Renvoize 1986,Kellogg 2015,Christenhusz and Byng 2016,Soreng et al. 2017). In the checklist of the family Poaceae in Turkey, Cabi and Doğan (2012) reported that Turkey hosts 146 grass genera including 547 species (658 taxa). After the publication of the checklist of the Poaceae, still more new grass taxa were recorded or described for the Turkish flora (e.g.Cabi et al. 2013,2015a,b,Doğan et al. 2015,Cabi and Soreng 2016,Cabi et al. 2018,Terzioğlu and Özkan 2020,Behçet and Yapar 2021).
Festuca L. is a genus of the family Poaceae with about 636 species, distributed in Europe, Africa, temperate Asia, tropical Asia, Australasia, the Pacific, North America, South America and Antarctica (Clayton and Renvoize 1986,Watson and Dallwitz 1992,Clayton et al. 2006-onwards,Martínez-Sagarra and Devesa 2019,POWO 2021). It is characterized by its perennial habit, paniculate inflorescence, usually with 4–5 flowered spikelets, rounded lemmas on the back and linear hilum running the length of the caryopsis. Festuca is not a well-researched genus for the Turkish flora, probably because of the high phenotypic plasticity and variability at the taxon level, which makes identification demanding. Taxonomic opinion since the first major monograph of Hackel (1882) on the genus, has varied, and it has been divided into several subgenera and sections or even segregate genera (Saint-Yves 1922,Tzvelev 1971,Alexeev 1977,1978,1981,1986,Clayton and Renvoize 1986,Aiken et al. 1997). In recent years, molecular phylogenetic studies conducted on the genus Festuca and its putative relatives have demonstrated the paraphyly of Festuca s.l. (Darbyshire and Warwick 1992,Charmet et al. 1997,Gaut et al. 2000,Torrecilla and Catalán 2002,Catalán et al. 2004,2007, Inda et al. 2008,Foggi et al. 2012). Clayton and Renvoize (1986) and Aiken et al. (1997) recognized the following subgenera: Drymanthele Krecz. and Bobrov, Helleria Alexeev, Leucopoa (Griseb.) Hack., Schedonorus (P. Beauv.) Peterm., Subulatae (Tzvelev) Alexeev, Xanthochloa (Kriv.) Tzvelev and Festuca. Then, the subgen. Leucopoa, subgen. Schedonorus, subgen. Drymochloa Holub and subgen. Parafestuca E.B. Alexeev were recognizedas segregated genera (Foggi et al. 2012). Additionally, species from the F. paniculata complex were recently separated into a separate genus Patzkea (L.) G.H.Loos (Loos 2010).
The subgenus Festuca is the largest and taxonomically most complex group (Hackel 1882,Saint-Yves 1922,Markgraf-Dannenberg 1980) in this genus. Drawing on molecular data, Catalán et al. (2007) proposed a new circumscription of the subgenus, recognizing only the sections Festuca, Aulaxyper Dumort. andEskia Willk., and later, Inda et al. (2008) added the sections Dimorphae Joch. Müll. and Catalán, Amphigenes (Janka) Tzvel., Scariosae Hack., Pseudoscariosa Krivot, Lojaconoa Catalán and Joch. Müll. and Subbulbosae Nyman ex Hack.
This genus is especially diverse in the northern hemisphere, and it is likely that the Eurasian region was its center of origin (Catalán et al. 2004,Inda et al. 2008). The genus Festuca is represented by 66 taxa of which 27 are endemic to Turkey according to Cabi and Doğan (2012) and were arranged in three subgenera and four sections (Markgraf-Dannenberg 1985).
Antalya is the richest province of Turkey in terms of plant diversity (Deniz and Aykurt 2014). A large part of the acreage of Antalya province is the mountainous Western Taurus range, and this area is one of the important endemism centers of Turkey. During the project, in which we investigated the Poaceae diversity of the high mountainous area of Antalya province, a large number of Poaceae samples were collected and evaluated in detail in terms of morphology. Festuca is one of the genera with the largest number of species in this area. The taxa of this genus, which is taxonomically highly complex, occurring in the Western Taurus Mountains, are included in Sect. Festuca Subsect. Festuca, except F. gigantea (L.) Vill. and F. anatolica Markgr.-Dann. subsp. anatolica. The species of this subsection were evaluated under five groups, which are Frigida, Laevis, Indigesta, Valesiaca and Ovina, in Turkey (Markgraf-Dannenberg 1985). The most useful character to segregate species groups within the different taxonomic sections is the arrangement of the sclerenchyma, and a remarkable variability of this character is found in the species of Festuca Sect. Festuca (Martínez-Sagarra et al. 2017).
For Festuca taxonomy, leaf anatomical characters may be the most important diagnostic character (e.g.Markgraf-Dannenberg 1985,Foggi and Rossi 1996,Martínez-Sagarra et al. 2017). The anatomical characteristics of the species distributed in the subalpine and alpine zones of the Western Taurus Mountains are given in this study. The leaf anatomical features of the new species have been compared with morphologically similar species within the same group and the species included in the other species groups of the Sect. Festuca Subsect. Festuca occurring in Turkey.
Material and methods
A large number of Festuca specimens were collected during field studies for a floristic survey for the project named “A research study on the grass family (Poaceae) distributed in the high mountain zone in Antalya province” between the years 2018 and 2020. The morphological characteristics of all these specimens were recorded both in the field and in the laboratory. Specimens collected were evaluated by the use of related literature (Tzvelev 1971,Alexeev 1977,Markgraf-Dannenberg 1980,1985,Clayton and Renvoize 1986,Watson and Dallwitz 1992,Kellogg 2015) and the specimens were deposited in AKDU and Nezahat Gökyiğit Botanical Garden (NGBG). The overall morphology of the specimens was examined under a stereo-binocular microscope. The new species described here is known from only its type locality. We visited this locality three times between the years 2019 and 2020 and collected a few individuals without damaging the population. Ten individuals were evaluated for morphological determination of F. albomontana Aykurt, Çıngay, Sümbül & Cabi. We used standard measurements and terminology in accordance with Hackel (1882), Markgraf-Dannenberg (1985), Ellis (1987), Fuente and Ortúñez (1998), Foggi et al. (1999), Martínez-Sagarra and Devesa (2019).
For the anatomical studies, the tiller leaves taken from herbarium samples were used. Leaves were kept in distilled water for c. five minutes. Transverse sections ± 0.05 mm thick were cut from pieces of tiller leaf fixed in styrofoam, by hand using a razor blade. This was done under a stereomicroscope using reflected light.
Results
The occurrence of nine Festuca species (10 taxa) was determined in the high mountainous area (Western Taurus) of Antalya province with the project mentioned above. Apart from F.anatolica subsp. anatolica (Sect. Variae) and F. gigantea (Sect. Bromoides), all the remaining Festuca species in the Western Taurus, F. callieri (Hack.) Markgr. subsp. callieri, F. callieri subsp. zederbaueri Markgr.-Dann., F. elwendiana Markgr.-Dann., F. valesiaca Schleich. ex Gaudin, F. jeanpertii (St.-Yves) F. Markgr. subsp. jeanpertii, F. pinifolia (Hack. ex Boiss.) Bornm. var. pinifolia, F. sipylea (Hack.) Markgr.-Dann., and F. albomontana described here as a new species, are included in the Sect. Festuca Subsect. Festuca. This subsection is divided into five groups according to Flora of Turkey. Considering the new species described here, the species representing each group are distributed in the Western Taurus Mountains: Festuca albomontana in the Frigida group, F. jeanpertii subsp. jeanpertii in the Laevis group, F. pinifolia var. pinifolia in the Indigesta group, F. valesiaca, F. callieri (subsp. callieri and subsp. zederbaueri) and F. elwendiana in the Valesiaca group, and F. sipylea in the Ovina group. The transverse sections taken from the tiller leaf blades of these taxa are shown inFig. 1. F. pinifolia var. pinifolia and F. sipylea have sclerenchyma forming a continuous ring. All the remaining taxa except for F. pinifolia var. pinifolia have sclerenchyma forming abaxial strands. In F. pinifolia var. pinifolia, sclerenchyma strands are located both abaxially and adaxially. Festuca alpina group species are distinguished by relatively small strands.
The transverse section of tiller leaves of the new species is more or less rounded, polygonal, not closed and not V-shaped. There are 5 vascular bundles and 5–8 small sclerenchyma strands, also with 2 very small submarginal accessory sclerenchyma strands, located under the abaxial side of the epidermis in F. albomontana. The median strand is generally slightly larger than the others. The 3 adaxial ribs are prominent.
Festuca albomontana is a chasmophytic species growing on rock crevices of the subalpine zone of Akdağ (Alanya, Antalya) and known only from its type location. The bedrock of this location is called Payallar formation and consists of Paleozoic schist, phyllite, quartzist, quartz etc. This formation is represented by lowly metamorphosed schists with quartzite, nodular limestone, marble dolomite interlayers (Şenel et al. 2016).
Densely tufted new fescue is morphologically characterized by 1. its fine, soft and glabrous leaves; 2. simple panicles with few spikelets; 3. lemmas with a long awn. A detailed comparison of both the morphological and anatomical characters of the new species and its morphologically close relatives are shown inTab. 1.
| Characters | F. albomontana | F. alfrediana (Foggi et al. 2012,POWO 2021) | F. alpina (Foggi et al. 2012,POWO 2021) | F. cratericola (Markgraf-Dannenberg 1985,POWO 2021) | F. sommieri (Markgraf-Dannenberg 1985,POWO 2021) | |
| Habit | More or less densely tufted grass with ascending vegetative shoots | More or less densely tufted grass, sometimes with prostrate-ascending basal vegetative shoots | More or less densely tufted grass, sometimes with prostrate-ascending basal vegetative shoots | Densely caespitose grass | Densely caespitose, glaucous grass | |
| Culms (length and indumentum) | 6–14 cm long, smooth, glabrous and distinctly fused for their entire length |
5–30 cm long, smooth and glabrous for their entire length | 3–21 cm long, smooth and glabrous along entire length | 10–15 cm long, thin, glabrous | 5–15 cm long, culm-internodes distally glabrous | |
| Sheats of tiller leaves | Papyraceous, scabrid on back, sometimes minutely so, completely fused, sometimes decaying into fibres | Papyraceous, glabrous, completely fused, sometimes decaying into fibres | Papyraceous, glabrous, fused for entire length, sometimes decaying into fibres | Glabrous, decaying into irregular fibres | Glabrous, irregularly decaying into transverse, undulate fibres | |
| Ligula | Ciliate at apex | With an eciliate membrane | With an eciliate membrane | Shortly ciliate, scabrid | With an eciliate membrane | |
| Panicle length (cm) | 2.4–4.5 | 1.5–4.2 | 1.2–3.6 | 3–4 cm | 1.5–3.5 cm | |
| Panicle branches | Glabrous, smooth, sometimes few scabridulous towards apex | Generally scabrid | Generally scabrid | Scabrid | Generally glabrous | |
| Spikelet | 5–7 mm long, with 2–4 flowers |
6.6–8.2 mm long, with 4–5(–6) flowers |
4.1–6.4 mm long, with (2–)3–4(–5) flowers. | 7–7.3 mm long, with 3–4 flowers | 3–flowered one 7–7.5 mm, with 2–3(–5) flowers | |
| Lower glume | Subulate, 2.5–2.8 mm long |
Subulate, 2.5– 3.5 mm long |
Subulate, 1.8–3.3 mm long | Lanceolate | Lanceolate, c. 3.5 mm long | |
| Upper glume | Subulate, 3.2–3.5 mm long | Subulate, 3.6–4.8 mm long |
Subulate, 2.7– 4.1 mm long | Oblong-lanceolate, 4–4.6 mm long | Oblong-lanceolate, 6–6.4 mm long | |
| Lemma | Subulate, 3.7–4.2 mm long, c. 1 mm wide, glabrescent to scabrid in the distal part |
Subulate, 4.3–5.7 mm long, 0.4–0.7 mm wide, slightly scabrid in the distal part |
Subulate, 2.8–4.3 mm long, 0.3–0.6 mm wide | Lanceolate, 4.5–5.2 mm long, 1.1–1.2 mm wide, scabrid in upper part. | Oblong-lanceolate, 5.7–6.9 mm long, 1.4–1.6 mm wide, scabrid in upper part | |
| Lemma awn length | 3.5–4.5 mm | 2.6–5.3 mm | 1.2–2.6 mm | 1,5–2 mm (c. 1/3 × lemma) | To 9 mm, often 2 × lemma | |
| Palea | 3.5–3.8 mm long, more or less finely hairy along its entire length |
4.1–5.2 mm long, more or less finely hairy along the carena. |
2.5–4.1 mm long, glabrous or sparsely finely hairy on the carena | 4.5–5.2 mm long | 5.7–6.9 mm long | |
| Anther length (mm) | 1.5–2 | 0.9–2.2 | 0.6–1.4 | 2.3–2.5 | 1.8–2 | |
| Tiller leaf anatomical features | ||||||
| Transverse section of tiller leaf blades | More or less rounded, polygonal (not closed), not V-shaped | Elliptic, V-shaped or slightly closed polygonal | V-shaped or slightly closed polygonal | Irregularly V-shaped, slightly closed, polygonal | Nearly rounded, not closed | |
| Number of vascular bundles | 5 | (3–)5 | 3 to 5 | 5 | 5(–7) | |
| Number of sclerenchyma stands | 5–8 small sclerenchyma strands, also with 2 very small submarginal accessory sclerenchyma strands |
3 narrow sclerenchyma strands, sometimes with two others submarginal
| 3 small sclerenchyma strands, sometimes with two others submarginal | 7, very thin, subequal, median strand of only 5–8 cells | 3, very small | |
| Number of ribs | 3 distinct adaxial ribs, without or with 1–2 minute hairs | 3 adaxial ribs of which two lateral ribs are generally only slightly developed | 3 adaxial ribs, with two lateral ribs generally less developed, with few hairs | 1 | Indistinct | |
Discussion
In Turkey, the species included in the Frigida group, which are F. cratericola Markgr.-Dann., F. sommieri Litard. and F. bushiana (St.-Yves) Tzvelev, are low plants of high alpine habitats, with short, simple panicles. These species are characterized by their (1) leaf sheaths closed nearly at the mouth and weakly decaying into fibres; (2) leaf blades which are usually fine, with 5 veins, l(–3) ribs, and 3–7 sclerenchyma strands (Markgraf-Dannenberg 1985) and they resemble species in the F. alpina group in terms of both morphological and anatomical features. Foggi et al. (2012) indicated that the F. alpina group, which comprises several small species typically found in exposed (mostly calcareous) rocky habitats in the high mountains of Europe, is characterized by tiller leaf sheaths completely closed or open in their upper half. The leaf anatomical features of this group and their systematic significance were evaluated by Martínez-Sagarra et al. (2017) and a leaf model was presented for the “F. alpina group” species: The leaves are conduplicate and quite narrow (0.34–0.62 mm) in transverse section. The number of vascular bundles is 3–5 and the sclerenchyma strands are quite thin. The species of this group are also characterized by their small size, flexible leaves, and short panicles with few spikelets (López et al. 2016). Festuca alpina Suter, F. alfrediana Foggi & Signorini, F. glacialis (Miégev ex Hack.) K. Richt. and F. frigida (Hack.) K. Richt. were included in this group (Foggi et al. 2012,2017). In recent years, it has been seen that the species with the morphological and anatomical features mentioned above are grouped under the “F. alpina group” rather than the “Frigida group”. For this reason, the new species described here is evaluated in the F. alpina group and it has a geographically isolated position in the Mediterranean region of Turkey. The other representatives of this group also have a limited distribution area and they are also known from a single location according to the Flora of Turkey. Festuca bushiana and F. sommieri occur in alpine habitats in the Black Sea region, Rize and Trabzon provinces, respectively. The other species, F. cratericola, of this group is endemic to Erciyes Mountain in Central Anatolia.
Here, we compared the new species with morphologically and anatomically similar taxa, i.e., F. sommieri, F. cratericola, F. alpina and F.alfrediana, by using literature information (e.g.Markgraf-Dannenberg 1985,Foggi et al. 2012,POWO 2021). Festuca alpina and F. alfrediana are also the two most common species in this group, growing in rocky habitats across the European mountain chains. According to Foggi et al. (2012), F. olympica Vetter from Greece and F. pirinica I.Horvat ex Markgr.-Dann. from Bulgaria both have open leaf-sheaths and therefore they do not belong to the Alpina group.
The lemmas with long awns are one of the most remarkable morphological features of F. albomontana,and the new species most resembles F. sommieri and F. alfrediana with respect to this character. However, the leaf anatomical features of the new species are quite different from both of them. The number of sclerenchyma strands and having 3 distinct ribs on the adaxial side of the leaf cross-section are the most useful diagnostic characters for separation of the new species from its close relatives.
Taxonomic treatment
Festuca albomontana Aykurt, Çıngay, Sümbül & Cabi sp. nov. (Figs. 2,3).
Type: Turkey: Antalya, Alanya, north of Akdağ (White Mountain), rock crevices near little lake, 2100 m elevation, 36°41´22.98´´ N / 32°16´47.76´´ E, 17.07.2019, C. Aykurt 5525 & H. Sümbül (holotype: AKDU; isotype: AKDU).
Diagnosis:Festuca albomontana differs from the related species F. sommieri by its subulate and 2.5–2.8 mm long lower glumes (vs. lanceolate and c. 3.5 mm long), subulate and 3.2–3.5 mm long upper glumes (vs. lanceolate and 6–6.4 mm long), 5–7 sclerenchyma strands (vs. 3), and 3 distinct adaxial ribs (vs. indistinct).
Description: More or less densely tufted grass with ascending vegetative shoots. Basal innovations intravaginal. Culms 6–14 cm long, smooth, glabrous along entire length. Leaves filiform, smooth, soft, minutely scabrid on margins, acute to obtuse, few scabrid at apex; ligule c. 0.4 mm long, auriculate, ciliate at apex. Tiller leaf sheaths papyraceous, scabrid on back, sometimes minutely so, completely fused, sometimes decaying into fibres; blades 2–8 × 0.03–0.05 cm; transverse section of tiller leaf blades polygonal, not closed, more or less orbicular, slightly angled, with 5 vascular bundles, 5–8 small sclerenchyma strands, also with 2 very small submarginal accessory sclerenchyma strands (the median strand is generally slightly larger than the others); with 3 distinct adaxial ribs with a few minute hairs. Culm leaf sheaths glabrous, closed to mouth; blades 1–2.5 × 0.03–0.04 cm. Panicle 2.4–4.5 cm long, with spreading short branches; branches glabrous, smooth. Spikelets 5–7 mm long with 2–4 florets; rachilla scabrid. Glumes unequal, subulate, upper glume distinctly longer than the lower glume, glabrous; lower glumes 2.5–2.8 × 0.5–0.9 mm, 1-veined; upper glumes 3.2–3.5 × 0.9–1.2 mm, 3-veined. Lemmas 3.7–4.2 × c. 1 mm, subulate, glabrescent to scabrid in the distal part; awn 3.5–4.5 mm long, minutely scabrid. Palea 3.5–3.8 × c. 1 mm, more or less finely hairy along its entire length. Anthers c. 1.5–2 mm long. Ovary glabrous. Caryopsis thin and glabrous.
Phenology: Flowering period starts in July, continues towards end of August. Fruiting period is late August.
Etymology: The specific epithet ‘albomontana’ refers to the Ak Mountain where the new species was discovered. “Ak Mountain” in translates as “Alba Montana”.
Habitat, ecology and conservation status: Festuca albomontana is a local species endemic to the high mountain zone of the Ak Mountain (Alanya, Antalya), inhabiting schistose rock crevices at an elevation of 2100 m. Within this area, the new species is associated with plants such as Eremopoa capillaris R.R. Mill, Poa davisii Bor, Melica persica Kunth subsp. jacquemontii (Decne. at Jacquem.) P.H. Davis, Paracaryum lithospermifolium (Lam.) Grande subsp. cariense (Boiss.) R.R.Mill, var. cariense, Lamium eriocephalum Benth., Stachys citrina Boiss. & Heldr. ex Benth. subsp. citrina, and Galium sorgerae Ehrend. & Schönb.-Tem.
The area of occupancy of the species is limited to less than 10 km2, and the total number of mature individuals is approximately 50. Overgrazing by goat herds was noticed near the type locality. Therefore, this new species should be considered as “critically endangered” (CR B2 ab(v); D) according to the IUCN categories (IUCN, 2019).
The identification key for species of the F. alpina group in Turkey
1a. Leaf blades fine, 0.25–0.5(-0.6) mm broad; sclerenchyma strands very slender………………………………………2
1b. Leaf blades stouter, (0.3–)0.4–0.65(–0.8) mm broad; sclerenchyma strands 3 strong………F. bushiana
2a. Lemma awn c. 1/3 × lemma length…………………………………………………………………………F. cratericola
2b. Lemma awn 1–2 × lemma length………………………………………………………………………………………………3
3a. Tiller leaves with 5–8 sclerenchyma strands, also with 2 very small submarginal accessory sclerenchyma strands; adaxial ribs 3, distinct…………………………………………………………………………………………………………………F. albomontana
3b. Tiller leaves with 3 sclerenchyma strands; adaxial ribs indistinct…………………………………………F. sommieri
Additional specimens examined:
F. callieri subsp. callieri: TURKEY. Antalya, Kaş, Akdağ, rocky areas near İkizgöller, 2355 m, 03.07.2020, C. Aykurt 5641 (AKDU). F. callieri subsp. zederbaueri: TURKEY. Antalya, Alanya, Gökbel Plateau, 1700 m, 18.06.2019, C. Aykurt 5248 (AKDU). F. elwendiana: Antalya: İbradı, around Derme Plateau, 1900 m, 24.06.2020, C. Aykurt 5634 (AKDU).F. valesiaca: TURKEY. Antalya: Alanya, Sarıveliler road, 1500 m, 17.06.2019, C. Aykurt 5131b (AKDU). F. pinifolia var. pinifolia: TURKEY. Antalya, Kaş, Akdağ, above İkizgöller, 2450 m, 03.07.2020, C. Aykurt 5649 (AKDU), F. sipylea: TURKEY. Antalya, Gazipaşa, Taşeli Plateau, above Mihrap Plateau, on rocks, 2015 m, 22.06.2020, C. Aykurt 5602 (ADDU). F. jeanpertii subsp. jeanpertii: TURKEY.Antalya: Elmalı, between Yuva and Seki, stony slopes, 1450 m, C. Aykurt 4813 (AKDU). F. alpina: AUSTRIA. Kärnten: Bezirk Spittal an der Drau, Innerfragant, Wanderweg von der Fraganter Hütte zum Schobertörl, hinter der Eggerhütte, 1900 m, 13 June 2010, C. Gilli, A. Berger (WU0067019). Österreich: Bezirk Liezen, Gemeinde Bad Mitterndorf, Katastralgemeinde Klachau; ca. 7 km ESE Bad Mitterndorf, um den Gipfel des Grimming, 2340 m, K. Zernig, (GJO0035680). F. bushiana: RUSSIA. Caucasus:Stavropol Krai, Karachay-Cherkessia & Kabardino-Balkaria18 June 1924, A. Shchukina (MW0649040). Krasnodar Krai & Adygea, North Ossetia, 2350 m, 30 June 1971, Khudyakova (MW0649040). Ingushetia and Chechnya, 27 June 1979, V.G. Onipchenko (MW0658393). North Ossetia, Ingushetia & Chechnya, 13 Aug 1981, V. V. Shcherbakov (MW0649045). F. cratericola: TURKEY. Kayseri: Erciyes Mountain, E-Setie, 2900 m, 7 June 1979, Carle, R. & Kurschner, H. (E00407254). Erdschias-dagh, auf feuchten Stellen beim nördlichen Krater, 2900 m., 24 June1924, Markgraf-Dannenberg, I., (W1916-0007506). F. sommieri: GEORGIA. Tschernomorsky (Province de la Mer Noire): M. Adzitouko (sources de la Mdzymta [Mzymta]), 2850 m, Markgraf-Dannenberg, I (1973-04), (W1916-0007510). RUSSIA. Caucasus:Stavropol Krai, Karachay-Cherkessia & Kabardino-Balkaria, 17 June 1994, Onipchenko V. G., (MW0649225). Caucasus boreali-occidentalis, in valle fl. Urup, mons Bolschaja Azgara, in rupibus subalpinae, 12 June 1971, E. B. Alexeev, (NMNH14809073). TURKEY. Rize:pass from Ikizdere to Ispir, 2600-2900 m, Sorger 82-119-60 (W).