Новости Сист. Низш. Раст. 51 — Novosti Sist. Nizsh. Rast. 51
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Новости сист. низш. раст. — Novosti Sist. Nizsh. Rast. 51: 204–219. 2017 The genus Parmelina (Parmeliaceae) in the Russian Caucasus I. N. Urbanavichene Komarov Botanical Institute RAS, Professor Popov Str., 2, St. Petersburg, 197376, Russia; [email protected] Abstract. The present study provides a brief survey of the genus Parmelina in the Russian Caucasus (Western Transcaucasia, Western, Central and Eastern Caucasus) with a special emphasis on species distribution. The study area is located in the larg- est nature reserves of the Russian Caucasus — Kavkazskiy, Utrish, Severo-Osetinskiy, Erzi. This paper contributes to a better understanding of the genus in the Caucasus, focusing mainly on the author’s collections in different regions of the Northern Cau- casus in 2008–2016. Four species have been identifi ed from the Russian Caucasus — Parmelina carporrhizans, P. pastillifera, P. quercina and P. tiliacea. Notes on their morphology, anatomy, ecology and distribution are based on data obtained from a study of ca. 20 specimens of each species. The presence of whitish macules on the up- per surface of P. carporrhizans (after Clerc, Truong, 2008) as an important diagnostic feature was shown. The Caucasian material shows the differences in size and propor- tions of ascospores, size and shape of pycnoconidia between critical species P. car- porrhizans and P. quercina. An identifi cation key for Caucasian Parmelina species is composed for the fi rst time and can be used for central regions of Russia. Keywords: lichens, genus Parmelina, taxonomic review, key to species, Cauca- sus, nature reserves, Russia. Род Parmelina (Parmeliaceae) на Российском Кавказе И. Н. Урбанавичене Ботанический институт им. В. Л. Комарова РАН, лаборатория лихенологии и бриологии, ул. Профессора Попова, д. 2, Санкт-Петербург, 197376, Россия; [email protected] Резюме. Представлен краткий обзор рода Parmelina для основных россий- ских районов Кавказа — Западного, Центрального, Восточного и Северо-Запад- ного Закавказья. Район исследований включает территории четырех крупней- ших на Кавказе российских заповедников — Кавказского, Северо-Осетинского, Утриш, Эрзи. Особое внимание уделено морфологическим, анатомическим при- знакам и распространению выявленных видов. Анализ особенностей строения и распространения представителей рода Parmelina в изученных районах основан на сборах 2008–2016 гг. Четыре вида рода Parmelina были выявлены для россий- ской части Кавказа — P. carporrhizans, P. pastillifera, P. quercina and P. tiliacea. Данные получены при изучении в среднем более чем 20 образцов каждого вида. В качестве важного диагностического признака, согласно работе Clerc и Truong (2008), использовано наличие беловатых макул на верхней поверхности у P. carporrhizans. На кавказском материале показаны различия между видами P. carporrhizans и P. quercina, заключающиеся в размерах и пропорциях аско- спор, а также в размерах и форме пикноконидий. Впервые составлен ключ для 204 Urbanavichene. Parmelina in the Russian Caucasus кавказских видов рода Parmelina, который может быть использован и при изуче- нии лихенофлоры центральных регионов России. Ключевые слова: лишайники, род Parmelina, таксономический обзор, ключ для определения видов, Кавказ, природные заповедники, Россия. The Parmelina Hale species have been studied by numerous research- ers. There are many papers on the morphology and taxonomy of the ge- nus Parmelina, as well as a lot of molecular data and population gene- tics studies in the Mediterranean, Southern and Central European regions (Alors et al., 2014; 2017; Argüello et al., 2007a, b; Clerc, Truong, 2008; Núñez-Zapata, 2013; Núñez-Zapata et al., 2011; 2015). The present pa- per continues the series of those on the lichen fl ora the Russian Caucasus (Urbanavichene, Urbanavichus, 2014, 2016; Urbanavichus, Urbanaviche- ne, 2014, 2015). The aim of this study is to refi ne our knowledge of the morphological and anatomical features in the delimitation of Parmelina species. Research by Núñez-Zapata (2013), combining molecular, chemi- cal, geographical and ecological characters within the genus Parmelina, especially P. quercina s. l., served as the basis for our work. Since its establishing by Hale (1976), the circumscription of the genus Parmelina has undergone signifi cant changes: of the 47 species, only 9 species remained (Elix, Hale, 1987; Divakar et al., 2010; Nelson et al., 2012), namely P. quercina (Willd.) Vain., P. carporrhizans (Taylor) Poelt et Vĕzda, P. pastillifera (Harm) Hale, P. tiliacea (Hoffm.) Hale, P. atri- cha (Nyl.) P. Clerc, P. coleae Argüello et A. Crespo, P. cryptotiliacea A. Crespo et Núñez-Zapata, P. gyrophorica Elix et al. and P. yalungana (Zahlbr.) P. R. Nelson et Kepler. Only three species of Parmelia now treated as Parmelina were mentioned by Rassadina (1971) from Russia and the Caucasus in general: Parmelia quercina (Willd.) Vain., P. carpor- rhizans Tayl. and P. scortea Ach. (P. scortea f. scortea and P. scortea f. coerulescens Harm). Currently four species of Parmelina are identifi ed from the Russian Caucasus, namely P. carporrhizans, P. pastillifera, P. quercina and P. tili- acea. All Parmelina species are common in the lowlands up to ca. 1850 m a. s. l. in the virgin or slightly anthropogenically disturbed woods. The habitats of Parmelina species in the Caucasus are confi ned to broadleaved and mixed coniferous-broadleaved forests. In the studied territory P. quer- cina, P. pastillifera and P. carporrhizans are corticolous, and only P. tili- acea may be saxicolous. 205 Новости сист. низш. раст. — Novosti Sist. Nizsh. Rast. 51: 204–219. 2017 Material and methods Specimens of Parmelina were collected in the Russian Caucasus in 2008–2016. The study is based on the author’s collections and mostly fi eld identifi cations (with light microscope). In such cases the author’s name is not specifi ed in the lists of examined specimens below. Some specimens were collected by the author together with G. Urbanavichus and J. Vondrák. Only representative specimens are deposited in the Herbarium of the Komarov Botanical Institute and marked by the Herbarium abbreviation (LE). The presented results are based on ca. 80 fi eld identifi cations and representative specimens of Parmelina from the Caucasian nature re- serves — Utrish (coll. by I. Urbanavichene et G. Urbanavichus), Kavkaz- skiy (coll. by I. Urbanavichene, G. Urbanavichus et J. Vondrák), Severo- Osetinskiy (coll. by I. Urbanavichene et G. Urbanavichus), Erzi (coll. by I. Urbanavichene et G. Urbanavichus). Morphology and anatomical structure of the species were studied by Zeiss microscopes (Axio Scope A1, Stemi 2000CS). Selected specimens were analyzed for secondary metabolites by high performance thin-layer chromatography (HPTLC) (Arup et al., 1993). When searching for new and cryptic species and identifi cation of problematic species P. quercina and P. carporrhizans, the ITS 1 and 2, including 5,8S rDNA sequences were obtained. The key with morphological and anatomical characters, based on an analysis of our herbarium material from the Russian Caucasus, uses in- dicative characteristics, including the ascospores length to width ratio (abbreviated as l/w) measured in 20–30 well developed ascospores and pycnoconidia of fertile specimens. The subdivision of the Caucasus in the lists of studied specimens below follows Menitskiy (1991): Western Caucasus, Central Caucasus, Eastern Caucasus, North-Western-Transcaucasia, and Western Transcaucasia. Results The genus Parmelina Hale is characterized by a foliose greyish white thallus with linear to sublinear lobes and cilia in the axils or margins, lecanorine apothecia, white medulla with black lower cortex; simple, black rhizines reaching the edge of the lobes; ellipsoid ascospores, thin- walled excipular cells darkly pigmented at the edge of the apothecium; presence of lecanoric or gyrophoric acid in the medulla (C+ rose-red), atranorin in the cortex (K+ yellow) (Hale, 1976; Clerc, Truong, 2008; Crespo et al., 2010; Divakar et al., 2010; Nelson et al., 2012; Núñez-Za- pata, 2013). The morphologically closest genus Myelochroa (Asahina) 206 Urbanavichene. Parmelina in the Russian Caucasus Fig. Location of the Caucasian nature reserves/ 1 — Utrish, 2 — Kavkazskiy, 3 — Severo-Osetinskiy, 4 — Erzi. 3 — Severo-Osetinskiy, 1 — Utrish, 2 Kavkazskiy, 207 Новости сист. низш. раст. — Novosti Sist. Nizsh. Rast. 51: 204–219. 2017 Elix et Hale is distinguished from Parmelina by chemistry (atranorin in medulla) and K+ yellow and C– reactions of the medulla. In the review, the most important characters for determination of Par- melina species are presented. The main synonyms for every species are given. Parmelina carporrhizans (Taylor) Hale, 1974, Phytologia, 28(5): 482. ≡ Parmelia carporrhizans Taylor, 1847, in Hook., London J. Bot. 6: 163. ≡ Parmelina quercina (Willd.) Halle var. carporrhizans (Taylor) V. Wirth, 1994, Stuttgarter Beitr. Naturk., A, 517: 62. Upper surface of thallus light grey, smooth, maculate, apothecia com- mon, rhizinate on underside, usually numerous at the centre of thallus. Well-developed ascospores ellipsoid, mostly c. 10 μm or more long, (7.0)9.0–11.7(12.8) × (5)5.1–5.8(7) μm, l/w = (1.56)1.65–2.0(2.1). Co- nidiomata black, immersed pycnidia, frequently present. Pycnoconidia always acicular (5.8)6.7–10.0(11.0) × 1 μm. Contains atranorin in the cortex (K+ yellow); lecanoric acid in the me- dulla (C+ and KC+ rose-red). In the Russian Caucasus, P. carporrhizans was found mostly in broad- leaved or in coniferous-broadleaved forests.