Document Type : Original Article
Authors
1 Department of Biotechnology, Iranian Research Organization for Science and Technology (IROST), Tehran, Iran
2 Botanical Garden and Botanical Museum Berlin, Freie Universität Berlin, Germany
Abstract
Keywords
INTRODUCTION
Arasbaran Biosphere Reserve is a mountainous area in the northern part of the province East Azerbaijan and belongs to the Iranian part of the Caucasian highlands situated between the Caspian and the Black Sea. The historical boundary of Arasbaran is ambiguous, but it is roughly defined here as the areas with intact forest and steppe-forest vegetation types. The area covers 160,000 hectares with a circumference of 250 km (Ghorbani, 2013; Sagheb-Talebi et al. 2014). The importance of biological diversity in Arasbaran forests led to the declaration that c. 80.000 hectares would be protected under the Man and the Biosphere (MAB) programme of the United Nations Educational, Scientific and Cultural Organization (UNESCO) in 1972 and as an IUCN Arasbaran Biosphere Reserve in 1998, see IUCN Jalili & Jamzad (1999). Locally, the area is known as “Qaradaq Mohali” (black mountain area) due to the dark appearance of the mountains at a distance. Today the area is composed of the northern part of Ahar and the eastern parts of Jolfa, Kaleybar and Varzegan counties, and includes the largest forested parts of the East Azerbaijan province.
The biological and anthropological significance of the Arasbaran Biosphere Reserve is due to the diversity of flora and fauna, as well as local cultural practices. The Arasbaran area encompasses a variety of natural features, such as high mountains, deep valleys, dense forests and vast rangelands with a very rich environmental diversity. Arasbaran Biosphere Reserve is affected by three different types of the climate resulting in a large diversity of ecosystems. Regional surveys have recorded c. 1000 vascular plant species, including 140 trees, the most important being Quercus atropatana ?atropatena?, Juniperus communis, Pistacia utlanta ?atlantica?, Carpinus betulus, Fraxinus ratuntifuliea ?rotundifolia? and Acer campestris, see Sohrabi & Alstrup (2007).
The first Iranian lichen list was based on collections made by Buhse (1860) in his biological investigation of the Transcaucasus and Hyrcanian area, and included over 50 lichen taxa from northern Iran, Armenia and some from Nakhchivan Autonomous Republic, an exclave of the Republic of Azerbaijan, but did not provide any lichenological records from along the Aras River, especially the valley between Jolfa and Kaleybar cited as "Gharadagh" (formerly named as the Arasbaran region). Based on the small collection made by the author, the first lichens, Cladonia and Lepraria spp., reported from the western part of the Arasbaran area appear in Ahti & Sohrabi (2006) and Sohrabi & Orange (2006) respectively. Additional data on parmelioid lichens from Arasbaran were reported by Sohrabi & Alstrup, (2007) and Sohrabi et al. (2007), and further records from there referred to in Seaward et al. (2008); Seaward et al. (2004); Myllys et al. (2011); Myllys et al. (2014); Velmala et al. (2014); Boluda et al. (2015); Valadbeigi, Nordin, et al. (2011); Valadbeigi, Sipman, et al. (2011).
The present study has several primary objectives to (1) demonstrate the importance of baseline information provided by regional studies of lichenological biodiversity, (2) contribute to our knowledge of lichenized and lichenicolous fungi of Iran and the Caucasus region in general, (3) provide new information on the occurrence and distribution of lichens and lichenicolous fungi of SW Asia, and (4) increase our knowledge of lichenology at a regional level for conservation purposes.
Outline of the study area
The study area is located in NW Iran (38º33´‒ 39º09´N & 46º09´‒ 47º09´E), c. 90 km from Tabriz, the capital city of the province of East Azerbaijan close to the countries Azerbaijan and Armenia (Fig. 1). It comprises the Gharadaq Mountain range (250‒2850 m) south of the Aras River, which includes high alpine meadows, semi-arid steppes, rangelands and occasional forested deep river valleys. Most of the visited localities in this study have been selected from forest areas due to their high diversity of lichens, particularly the trees and shrubs which are rich in epiphytic species. The lichens recorded grow on soil, siliceous or calcareous rocks, and mossy siliceous rocks. According to Talebi et al. (2014) and Sagheb-Talebi et al. (2001) forest types were distinguishable in accordance with the ecological adaptability of the tree species at different altitudes. Quercus spp. are widely distributed, and Juniperus stands to occur at 400‒1800 m, both in mixed or pure stands. Cornus spp., show a very narrow distribution between 900 and 1000 m.
Fig. 1. Location of the collection sites (indicated by Latin numbers) in the Arasbaran area in NW of Iran.
Five forest types were described by Sagheb-Talebi et al. (2001):
According to FAO (2004), the Arasbaran forests are the most degraded forests of NW Iran. Studies such as Abbasloo (1996); Alijanpour & Mahmoudzadeh (2007); Sarhangzadeh & Makhdoum (2003); Shanjani (2001); Sagheb-Talebi et al. (2014); Sagheb-Talebi et al. (2001) based on aerial photographs from early 1945, estimate that the forests originally covered 500,000 ha before being reduced to a mere 60,000 ha today. A more conservative estimate by FAO (2004) gives the 1958 area as 300,000 ha, and the present one as 200,000 ha.
In the Arasbaran forest area c. 60,000 ha of open forests and 37,500 ha of closed forests are protected; they are dominated by oak (Quercus macranthera, Q. petraea), hornbeam (Carpinus betulus), maple (Acer campestre, A. monspessulanum), elm (Ulmus glabra), wild cherry (Prunus avium), ash (Fraxinus excelsior), junipers (Juniperus communis, J. foetidissima, J. excelsa) and yew (Taxus baccata). The main forest associations are pure oak (Quercetum), pure hornbeam (Carpinetum), mixed oak-hornbeam (Querco-Carpinetum), mixed broadleaved, and conifer forests (Sagheb-Talebi et al. 2001). Conversion of forest topasture and logging has resulted in severe degradation and a decrease in forest cover.
A large part of the Arasbaran area is rangelands which cover the low elevations in the warmer regions, as well as higher elevations in Saygaram-daq. The vegetation of these areas can be roughly defined as either scrublandor grassland. Scrublands are found at elevations of 250 to 2200 m in different parts of the area. The most typical species are Punica granatum, Paliurus spina-christi, Rhamnus paliurus, Cotoneaster spp., and Ephedra spp., mixedwith a large diversity of other plants. Grasslands can be found at elevations of 250 to 2800 m and are dominated by different species of Poaceae (e.g. Agropyron & Stipa spp.), and herbs such as Astragalus, Stachys and Euphorbia spp., together with many annual plants. According to Assadi (1988), plant diversity of the rangelands is exceptionally high, especially on steppe sites and in open areas in the mountain forests. It is an important source of food for local indigenous breeds of livestock kept by local nomads and subsequently settled villagers over thousands of years. Today the rangelands are deteriorating due to the excessive numbers of livestock grazing these areas.
MATERIALS AND METHODS
During several field trips in 2001, 2004, 2005, 2006 and 2007, ca. 1000 specimens were collected in eight localities in the Arasbaran UNESCO-MAB Biosphere Reserve, and incorporated into the author’s personal herbarium in Daran village, Jolfa County (East Azerbaijan Province) and duplicated for ICH herbarium at the Iranian Research Organization for Science and Technology (IROST) in Tehran. So far ca. 200 specimens have been published in, e.g., Sohrabi & Sipman (2007); Seaward et al. (2008); Sohrabi & Alstrup (2007); Sohrabi & Orange (2006). The present report is based on the examination of ca. 533 further specimens. For the remaining ca. 250 specimens the taxonomic status is still unsettled, and awaiting revisional studies in groupssuch as Acarospora, Circinaria, Lecanora, Lecidea, Pertusaria, Rinodina, Rhizocarpon, Staurothele, Verrucaria.
The morphology and anatomy of specimens were studied with the aid of dissecting and compound microscopes, and supported where necessary by chemical reactions (Ryan et al. 2002). Microcrystallography and thin-layer chromatography (TLC) were done for some selected group of lichens following Orange et al. (2001). The macrolichen v. microlichen concept followed that defined by Kirk et al. (2001). The principal identification keys used were Nash et al. (2007); Gaya (2009); Smith et al. (2009); Sliwa (2007); Temina et al. (2005); Nash et al. (2002); Krzemińska (2012); Awasthi (1991); Wirth (1995); Dobson (2011); Malcolm & Galloway (1997); Ryan et al. (2002) were used. The species names were checked against Index Fungorum (http://www.speciesfungor um.org/names/names.asp) and Mycobank (Crous et al. 2004) to ensure their orthography. The examined specimens were stored in the herbarium of the Iranian research organization for Science and Technology with its acronym ICH, some in IRAN. Some reference material from the herbaria H, GB, C, TUR, and S was consulted for comparative purposes. The generic and family names are based on Wijayawardene et al. (2018).
Information on ecology and distribution, as well as nomenclature, has been mainly derived from Clauzade & Roux (1985); Clauzade et al. (1989); Smith et al. (2009); Brodo et al. (2001); John & Türk (2017); Santesson et al. (2004); Esslinger (2016); Nimis (2016); Gasparyan & Nimis (2016). Total genomic DNA was extracted from specimens 4375 (Lecidea auriculata) following Sohrabi et al. (2013).
Collecting Localities
I. 18.7.2005. Varzegan county, c. 26 km W of Varzegan and c. 6 km E of Joshin village (Kharvana), Joshin Castle, 1700-2000 m, 38.6517 N, 46.3500 E. A fairly arid site dominated by vascular plants, including Astragalus, Rosa, and numerous Poaceae spp., and many annual plants.
II. 15.7.2001. Jolfa county, Jolfa to Khoda-afarin, 5 km S of Missan village, 1000-1500 m, 38.8690 N, 46.4523 E. Slope covered by Lonicera and Juniperus, and also by Quercus and Carpinus at higher elevations.
III. 20.8.2005. Kaleybar, 21 km S of Khoda-afarin to Jolfa road, Aynaloo, 1700-1900 m, 38.8342 N, 46.7914 E. Slopes covered by Quercus and Carpinus. [with Masoomeh Ghobad-Nejhad]
IV. 19.8.2005. Kaleybar, c. 10 km S of Aras River and Khoda-afarin to Jolfa road, Dar-Aghzi village, 450 m, 39.0834 N, 46.8832 E. Slopes covered byRosa, Amygdalus and mostly by Paliurus spina- christi.
V. 19.8.2005. Kaleybar county, c. 4 km SW of Kaleybar, Galadarasi, towards Babak Castle (Bez Galasi), 1750-2500 m, 38.8686 N, 46.9683 E. Slopes covered by Quercus, Carpinus, and Crataegus.
VI. 20.8.2005. Kaleybar, c. 10 km W of Kaleybar, Hejranduost village, 1750-1850 m, 38.8686 N, 46.9683 E. Dominated by Juniperus communis and many annual species.
VII. 20.7. 2004. Jolfa county, Khoda-afarin to Jolfa road, crossing at Uoshtipin village, Hrass, 500 m, 38.8956 N, 46.4462 E. Roadside siliceous boulders.
VIII. 5.11. 2007. Kaleybar county, 11 km S of Asheqlu, along the road to Aynalu. 1250 m. 38.9387 N, 46.7628 E. Low, regenerating deciduous forest dominated by Carpinus orientalis. [with H. J. M. Sipman, U. Søchting & M. R. Asef]
RESULTS
As a result of this fieldwork in the Arasbaran UNESCO-MAB Biosphere Reserve, 227 lichenized and 36 lichenicolous fungi are reported, including 110 genera from 50 families in the Ascomycota and a single taxon in the family Corticiaceae, Basidiomycota. The majority of species listed are macrolichens with 122 species (44 epiphytic, 38 terricolous & 40 saxicolous) belonging to the following genera: Anaptychia, Bryoria,Candelaria, Catapyrenium, Cetraria, Cetrelia, Chaenotheca, Cladonia, Collema s.l., Dermatocarpon, Evernia, Flavoparmelia, Fuscopannaria, Hypogymnia, Leptogium, Lepra, Lepraria, Leprocaulon, Lobaria. Melanelixia, Melanohalea, Nephroma, Parmelia, Peltigera, Phaeophyscia, Physcia, Physconia, Protoparmeliopsis, Ramalina, Rhizoplaca, Squamarina,Toninia, Tornabea, Umbilicaria, Usnea, Xanthomendoza, Xanthoparmelia and Xanthoria, and 110 species of microlichens (31 epiphytic, 15 terricolous & 63 saxicolous) belonging to the following genera: Acarospora, Arthonia, Aspicilia, Buellia, Caloplaca, Candelariella, Catillaria, Dimelaena, Diploschistes, Immersaria, Ingvariella, Lambiella, Lecania, Lecanora, Lecidea, Lecidella, Lepra, Lobothallia, Melaspilea, Mycobilimbia, Ochrolechia, Pachyphiale, Pertusaria, Phlyctis, Placocarpus, Pleopsidium, Psilolechia, Rhizocarpon, Rinodina, Scoliciosporum, Staurothele, Thelenella, and Verrucaria. The most diverse lichenized families are Parmeliaceae (33), Physciaceae (28), Lecanoraceae (23), Teloschistaceae (15), Verrucariaceae (14), Cladoniaceae (13), Megasporaceae (12) and Ramalinaceae (11), and the most diverse genera are Cladonia (13 taxa), Lecanora (11), Peltigera (9), Xanthoparmelia (9), Umbilicaria (6), Ramalina (6), Physcia (6), Circinaria (6), Bryoria (5), Phaeophyscia (5), Physconia (5)and Lecidella (5). The most diverse lichenicolous fungi families are Mycosphaerellaceae (7 taxa), “incertae sedis” (8), Verrucariaceae (3), Polycoccaceae (2) Phaeococcomycetaceae (2) and Lichenoconiaceae (2), and the most diverse genera are Stigmidium (5 taxa), Rosellinula (2), Phaeosporobolus (2), Lichenoconium (2), Sphaerellothecium (2), Muellerella (2) and Lichenostigma (2).
List of taxa
The following data are provided: Ref. = literature reviews and reports; Syn. = important synonyms; functional group (L = lichenized fungi, LF = lichenicolous fungi, and LL = lichenicolous lichens); substrate; I to VIII = collection localities listed above, followed by the collection numbers of the samples taken at that locality; TLC = results of thin-layer chromatography; and New DNA = Genbank accession number in bold. Unless otherwise indicated, the collection numbers are those of M. Sohrabi and the specimens are deposited in the ICH herbarium at the Iranian Research Organization for Science and Technology (IROST) with duplicates in the personal herbarium of M. Sohrabi.
Abrothallus caerulescens Kotte
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Acarospora fuscata(Nyl.) Arnold
L; on siliceous rock; II: 1459. TLC: gyrophoric acid.
Acarospora laqueata Stizenb.
L; on siliceous rock; I: 3597. TLC: gyrophoric acid.
Alyxoria culmigena (Lib.) Ertz
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008) as Opegrapha culmigenaLib.
Alyxoria varia (Pers.) Ertz & Tehler
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008) as Opegrapha variaPers. L; on Carpinus base; VIII: Sipman 55427 [IRAN, B 60 0175537].
Anaptychia crinalis (Schrad.) Vězda
L;on the bark of deciduous trees; V: 4670, 4509.
Anaptychia desertorum (Rupr.) Poelt
L; on siliceous rock; I: 3517.
Anaptychia elbursiana (Szatala) Poelt
L; on siliceous or calcareous rocks; I: 3528; IV: 3682.
Anaptychia setifera Mereschk. ex Räsänen
Ref.: Jamshidi et al. (2014). L; on bark or branches of deciduous tree; II: 123, 118, 1402, 137, 1493, 1544; III: 3688; V: 4599, 4655, 4528, 4636, 4561; VIII: Sipman 55457 [IRAN, B 60 0175569].
Arthonia apotheciorum (A. Massal.) Almq.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008) as Arthonia galactinaria Leight.
Arthonia intexta Almq.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Arthonia radiata (Pers.) Ach.
L; on Carpinus base; VIII: Sipman 55447 [IRAN, B 60 0175559].
Arthonia varians (Davies) Nyl.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Arthrorhaphis aeruginosa R. Sant. & Tønsberg
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Aspicilia candida (Anzi) Hue
L; on calcareous rock; VI: 4302b.
Aspicilia cinerea (L.) Körb.
L; on siliceous rock; III: 4065, 3807, 3729. TLC: norstictic acid.
Aspicilia pavimentans (Nyl.) Hue
New to Iran; L; on calcareous rock; V: 4577. TLC: norstictic acid.
Biatora carneoalbida (Müll. Arg.) Coppins
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008) as Mycobilimbia carneoalbida(Müll.Arg.) Vitik. et al.).
Bryobilimbia hypnorum (Lib.) Fryday, Printzen & S. Ekman
New to Iran; L: on mosses; V: 4493.
Bryoria capillaris (Ach.) Brodo & D. Hawksw.
L; on mosses growing on rock; V: 4743, 4691, 4432. TLC: atranorin, barbatolic, fumarprotocetraric, and psoromic acids.
Bryoria implexa (Hoffm.) Brodo & D. Hawksw.
Ref.: Myllys et al. (2011); Boluda et al. (2015); Velmala et al. (2014). L; on mosses on rock; V: 4656a. TLC: connorstictic and norstictic acid.
Bryoria kuemmerleana (Gyeln.) Brodo & D. Hawksw.
Ref.: Velmala et al. (2014). L; on saxicolous mosses; V: 4656b. TLC: atranorin, norstictic, connorstictic, and psoromic acids.
Bryoria nadvornikiana (Gyeln.) Brodo & D. Hawksw.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008); Boluda et al. (2015). TLC: barbatolic, alectorialic, and fumarprotocetraric acids.
Buellia zoharyi Galun
L; on calcareous soil; VII: 4263. TLC: atranorin, norstictic, and stictic acids.
Calogaya biatorina (A. Massal.) Arup, Frödén & Søchting
Syn.:Caloplaca biatorina (A. Massal.) J. Steiner; L; on siliceous rock; V: 4596.
Calogaya decipiens (Arnold) Arup, Frödén & Søchting
Syn.: Caloplaca decipiens (Arnold) Blomb. & Forssell; L; on calcareous rock; VI: 4395.
Calogaya saxicola (Hoffm.) Vondrák
Syn.: Caloplaca saxicola (Hoffm.) Nordin; L; on siliceous rock; III: 4006.
Caloplaca cerina (Ehrh. ex Hedw.) Th. Fr. s.lat.
L; on bark or banches of deciduous tree; II: 105; III: 3813, 3685, 3690, 3772, 4020, 4013; V: 4645, 4467; VIII: Sipman 55442 [IRAN, B 60 0175554].
Caloplaca phaeothamnos Kalb & Poelt
New to Iran; L; on soil surfaced rock bases on fine rich soils; III: 4232.
Caloplaca stillicidiorum (Vahl) Lynge
L; on mosses; I: 3538b.
Candelaria concolor (Dicks.) Stein
L; on bark or branches of deciduous trees; V: 4475.
Candelariella kansuensis H.Magn.
Ref.: Westberg & Sohrabi (2012). L; on siliceous rocks; I: 3535.
Candelariella lutella (Vain.) Räsänen
Ref.: Westberg & Sohrabi (2012). L; on bark of deciduous trees; III: 3694.
Candelariella rosulans (Müll.Arg.) Zahlbr.
Ref.: Westberg & Sohrabi (2012). L; on siliceous rock; III: 3644.
Candelariella vitellina (Hoffm.) Müll.Arg.
Ref.: Westberg & Sohrabi (2012).L; on mossy siliceous rock; III: 3726b.
Catapyrenium cinereum (Pers.) Körb.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Catillaria chalybeia (Borrer) A. Massal.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Cetraria aculeata (Schreb.) Fr.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008) as Cetraria steppae (Savicz) Kärnefelt. Based on molecular studies, C. steppae was synonymizedunder C. aculeata in Nadyeina et al. (2013). TLC: norstictic acid. DNAfrom Iranian specimeninNadyeina et al. (2013): KU200416, KU200417, KU200418.
Cetraria islandica (L.) Ach.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008). TLC: lichesterinic, protocetraric, and fumarprotocetr-
aric acids.
Cetrelia monachorum (Zahlbr.) W.L. Culb. & C.F. Culb.
New to Iran; L; on mossy siliceous rock; V: 4468. TLC: atranorin and imbricaric acid.
Cetrelia olivetorum (Nyl.) W.L. Culb. & C.F. Culb.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008). TLC: atranorin, chloroatranorin, and olivetoric acid.
Chaenotheca furfuracea (L.) Tribell
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Circinaria calcarea (L.) Körb.
L; on calcareous rock; I: 3580. TLC: aspicilin.
Circinaria elmorei (E.D. Rudolph) Owe-Larss., A. Nordin & Sohrabi
New to Iran; L; on calcareous rock; IV: 3679.
Circinaria fruticulosa (Eversm.) Sohrabi (Fig. 2)
Ref.: Sohrabi et al. (2013). L: on soil, vagrant. TLC: none detected.
Circinaria maculata (H. Magn.) Q. Ren
L; on calcareous rocks; I: 3537, 3570, 3602; IV: 3635, 3666. TLC: none detected.
Cladonia borealis S. Stenroos
New to Iran; L; on soil; V: 4539. TLC: barbatic and usnic acids.
Cladonia cariosa (Ach.) Spreng.
Ref.: Ahti & Sohrabi (2006); Seaward et al. (2008). L; on soil; II: 152. TLC: atranorin and fumarprotocetraric acid
Cladonia chlorophaea (Flörke ex Sommerf.) Spreng.
Ref.: Ahti & Sohrabi (2006); Seaward et al. (2008). L; on soil; II: 1474, 1388. TLC: fumarprotocetraric acid complex.
Cladonia coniocraea (Flörke) Spreng.
L; on dead wood or dead part of fallen trees; III: 3780. TLC: fumarprotocetraric acid complex.
Cladonia fimbriata (L.) Fr.
Ref.: Ahti & Sohrabi (2006); Seaward et al. (2008). L; on soil and mosses; I: 3527; II: 137 III: 4225; V: 4650. TLC: fumarprotocetraric acid complex.
Cladonia foliacea (Huds.) Willd.
Ref.: Ahti & Sohrabi (2006); Seaward et al. (2008). L; on soil and mosses; II: 179, 1370, 1505; IV: 3645; V: 4537; VIII: Sipman 55455 [IRAN, B 60 0175567. TLC: usnic, fumarprotocetraric acids.
Cladonia macilenta Hoffm.
L; on soil and mosses; III: 4093. TLC: thamnolic, and barbatic acids.
Cladonia magyarica Vain.
Ref.: Seaward et al. (2008). L; on soil and mosses; V: 4553; VI: 4271. TLC: atranorin and fumarprotocetraric acid.
Cladonia pocillum (Ach.) Grognot
Ref.: Ahti & Sohrabi (2006); Seaward et al. (2008). L; on soil I: 3531; II: 130 V: 4614. TLC: fumarprotocetraric acid complex.
Cladonia pyxidata (L.) Hoffm.
Ref.: Ahti & Sohrabi (2006); Seaward et al. (2008). L; on soil; II: 1448, 141, 116; V: 4608, 4514; VI: 4328. TLC: fumarprotocetraric acid complex.
Cladonia rangiformis Hoffm.
Ref.: Ahti & Sohrabi (2006); Seaward et al. (2008). L; on soil, mostly associated with mosses; II: 120, 134; III: 4004; IV: 3656; VIII: Sipman 55463 [IRAN, B 60 0175575]. TLC: strain 1, atranorin, rangiformic acid, and fumarprotocetraric acid complex; strain 2, atranorin, rangiformic, and tr. norrangiformic acids (55463).
Cladonia rei Schaer.
L; on soil; III: 4186, 4023. TLC: homosekikaic, sekikaic acids, and fumarprotocetraric acid complex.
Cladonia subrangiformis Sandst.(Fig. 2)
Ref.: Ahti & Sohrabi (2006); Seaward et al. (2008). L; on soil, mostly associated with mosses; II: 154, 1502 III: 3795, 4056, 4084; V: 4755, 4747, 4466; VI: 4322. TLC: atranorin and fumarprotocetraric acid.
Cladonia symphycarpa (Ach.) Fr.
L; on soil; V: 4753. TLC: atranorin and norstictic acid.
Clypeococcum cladonema (Wedd.) D. Hawksw.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Collema flaccidum (Ach.) Ach.
L; on siliceous rock or bark, often over mosses; III: 4010, 4136, 3705; V: 4522, 4695, 4502.
Dermatocarpon intestiniforme (Körb.) Hasse
L; on siliceous rock; V: 4609, 4521.
Dermatocarpon miniatum (L.) W. Mann
L; on limestone, rarely on slightly calcareous or siliceous rocks; I: 3501; II: 126 III: 4101; V: 4731, 4560, 4513.
Dermatocarpon moulinsii (Mont.) Zahlbr.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Dermatocarpon vellereum Zschacke
L; on siliceous rocks; II: 1410; III: 3740; V: 4478, 4688.
Dimelaena oreina (Ach.) Norman
L; on slightly calcareous, hard, siliceous (often feldspar rich) siliceous rock; I: 3586; II: 1464, 1467; III: 4001, 3764, 3722. TLC: usnic, fumarprotocetraric, and gyrophoric acids.
Diploschistes muscorum (Scop.) R. Sant.
L; on calcareous soil and mosses, occasionally with Cladonia spp.; I: 3547; II: [1367, (F)]; V: 4628, 4511. TLC: diploschistesic and lecanoric acids.
Diploschistes scruposus (Schreb.) Norman
L; on weakly calcareous or siliceous rock; I: 3503, 3640, 3650; II: [1375, 1409, 1468, (F)], 1499, 1443; III: 4039, 4083; V: 4570; VII: 4241. TLC: diploschistesic and lecanoric acids.
Enchylium tenax (Sw.) Gray
Syn.: Collema tenax (Sw.) Ach.; L; on bare, or sparsely overgrown with mosses and higher plants, soil; II: 1500.
Endocarpon pallidum Ach.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Endococcus perpusillus Nyl.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Evernia prunastri (L.) Ach.
L; on acid bark of deciduous trees; III: 3773, 4080, 4175; V: 4588, 4452, 4554.
Flavoparmelia caperata (L.) Hale
Ref.: Sohrabi et al. (2007); Seaward et al. (2008).L; on mossy rock; VIII: Sipman 55466 [IRAN, B 60 0175577].
Gyalecta fagicola (Arnold) Kremp.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008) as Pachyphiale fagicola (Arnold) Zwackh.
Gyalolechia flavorubescens (Huds.) Søchting, Frödén & Arup
Syn.: Caloplaca flavorubescens (Huds.) J. R. Laundon; L; on bark or branches of deciduous trees; III: 4127; VIII: Sipman 55433 [IRAN, B 60 0175543].
Gyalolechia flavovirescens (Wulfen) Søchting, Frödén & Arup
Syn.: Caloplaca flavovirescens(Wulfen) Dalla Torre & Sarnth.; L; on calcareous rock; VI: 4382.
Hypogymnia austerodes (Nyl.) Räsänen
Ref.: Seaward et al. (2008); Sohrabi et al. (2007).
Hypogymnia physodes (L.) Nyl.
Ref.: Sohrabi et al. (2007); Seaward et al. (2008).
Hypogymnia vittata (Ach.) Parrique
Ref.: Sohrabi et al. (2007); Seaward et al. (2008).
Immersaria cupreoatra(Nyl.) Calatayud &Rambold
Ref.: Valadbeigi, Sipman, et al. (2011). L; on the siliceous rocks; TLC: gyrophoric acid.
Immersaria iranicaValadbeigi, Sipman & Rambold
Ref.: Valadbeigi, Sipman, et al. (2011). L; on siliceous rock. TLC: 2′-O-methylsuperphyllinic acid.
Immersaria usbekica (Hertel) M. Barbero, Nav.-Ros. & Cl. Roux
L; on siliceous rock; V: 4430, 4701, 4546, 4707. TLC: confluentic and/or gyrophoric acid.
Ingvariella bispora (Bagl.) Guderley & Lumbsch
L; on siliceous rock; VI: 4379.
Lathagrium cristatum (L.) Otálora, P.M. Jørg. & Wedin
L; over mossy calcareous rock; II: 4194, 4027; IV: 3605a, 3670.
Lambiella insularis(Nyl.) T. Sprib.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008) asRimularia insularis(Nyl.) Rambold & Hertel.
Lecania cyrtella (Ach.) Th. Fr.
L; on Carpinus base; VIII: Sipman 55448 [IRAN, B 60 0175560]. Confirmed by P. van den Boom, 2017.
Lecania dubitans (Nyl.)A. L. Sm
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Lecania koerberiana J. Lahm
L; on bark and branches of deciduous trees; VI: 4343b.
Lecanora albellula (Nyl.) Th. Fr.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008). TLC: isousnic and usnic acids.
Lecanora allophana Nly.
L; on bark free or open standing deciduous trees; II: 93, 140; III: 3692, 3700, 3703, 3706, 3742, 4082, 4106, 4164; V: 4433, 4725, 4440, 4516, 4500, 4651; VIII: Sipman 55437 [IRAN, B 60 0175548]. TLC: atranorin and chloroatranorin.
Lecanora argentata (Ach.) Malme
L; on deciduous trees, usually with smooth and cracked bark; III: 3774; VI: 4343.
Lecanora argopholis (Ach.) Ach.
L; on siliceous rock; II: 1471, 1420, 1404; III: 4033, 4055; IV: 3665, 3647; V: 4637, 4504. TLC: usnic acid,atranorin, epanorin, zeorin, and fatty acids.
Lecanora campestris (Schaer.) Hue.
L; on siliceous rock; VI: 4423.
Lecanora carpinea (L.) Vain.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008). L; epiphytic; VIII: Sipman 55437 [IRAN, B 60 0175548].
Lecanora populicola (DC.) Duby
New to Iran; L; on bark or branches of deciduous trees; V: 4660.
Lecanora reagens Norman
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Lecanora rupicola (L.) Zahlbr.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008). TLC: atranorin, eugenitol, chloroatranorin, and roccellic acid.
Lecanora symmicta (Ach.) Ach.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
*Lecidea auriculata Th. Fr.
New to Iran; L; on siliceous rock; VI: 4375. TLC: confluentic acid, 2'-O-methylanziaic acid. DNA: MK341059. In order to find the correct affiliation of this specimen, internal transcribed spacers ITS rDNA (ITS1 and ITS2) and the 5.8S ribosomal DNA gene were obtained in this study and its preliminary analysis was used for a confirmation of the species identity.
Lecidea swartzioidea Nyl.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Lecidea tessellata Flörke
L; on siliceous rocks; II: 1407, 1506; III: 3755; IV: 3683; V: 4562; VI: 3225.
Lecidella carpatica Körb
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Lecidella elaeochroma (Ach.) M. Choisy
L; on trunks and branches of deciduous trees; II: 105b; III: 4063; VIII: Sipman 55437 [IRAN, B 60 0175549].
Lecidella euphorea (Flörke) Hertel
L; on trunks and branches of deciduous trees; II: 122, 1504, 103, 1447; III: 3711, 3794, 3812, 4070b, 4247, 4028, 4235; V: 4482, 4530, 4451, 4745, 4557, 4756, 4547.
Lecidella patavina (A. Massal.) Knoph & Leuckert
L; on calcareous rocks; I: 3579; IV: 3607; VI: 4290, 4316.
Lecidella stigmatea (Ach.) Hertel & Leuckert
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Lepra albescens (Huds.) Hafellner
Syn.: Pertusaria albescens (Hud.) M. Choisy & Werner; L; on the bark of deciduous trees; III: 4061.
Lepra amara (Ach.) Hafellner
Syn.: Pertusaria amara(Ach.) Nyl.; L; on the bark of deciduous trees; III: 4079.
Lepra excludens (Nyl.) Hafellner
Syn.: Pertusaria excludens Nyl.; L; on siliceous rocks; I: 3541, 3502, 3504, 3581; V: 4439. TLC: lichexanthone, norstictic acid.
Lepraria crassissima (Hue) Lettau
L; on dead mosse; III: 4076, 4171; V: 4726. TLC: divaricatic and nordivaricatic acids, zeorin.
Lepraria finkii (B. de Lesd.) R.C. Harris
L; on the bark of a deciduous tree; III: 3814; V: 4448, 4662. TLC: atranorin, zeorin, and stictic acid with satellites.
Lepraria leuckertiana (Zedda) L. Saag
Syn.:Lecanora leuckertiana Zedda; L; on the bark of deciduous trees; III: 4204, 3760, 3695. TLC: usnic, isousnic acids, and zeorin.
Lepraria membranacea (Dicks.) Vain.
L; commonly on lime-free siliceous rock; V: 4742. TLC: Pannaric, roccellic, angardianic acids, and occasionally atranorin.
Lepraria vouauxii (Hue) R. C. Harris
Ref.: Sohrabi & Orange (2006). L; commonly grows in the shady habitat on lime-free siliceous rock; I: 3560, 3590, 3549; II: 1539. TLC: pannaric acid-6-methyl ester, roccellic, and angardianic acids.
Leprocaulon microscopicum (Vill) Gams ex D. Hawksw.
L; on thin soil layers or mosses; II: 1491; III: 4086.
Leproplaca chrysodeta (Vain.) Ahti
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008) as Caloplaca chrysodeta (Vain. ex Räsänen) Dombr.
Leptogium burnetiae C. W. Dodge
L; on mosses on rocks among trees or in sheltered habitats; II: 375b; III: 4055, 4143, 4207; V: 4489. The species is known from N America and C Europe.
Leptogium saturninum (Dicks.) Nyl.
L; on the bark of a deciduous tree; II: 92; III: 3732.
Lichenoconium pyxidatae (Oudem.) Petr. & Sydow
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Lichenoconium usneae (Anzi) D. Hawksw.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Lichenostigma cosmopolites Hafellner & Calatayud
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Lichenostigma rugosum G. Thor
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Lichenostigma verrucosum Brackel & Valadbeigi
Ref.: Valadbeigi & Brackel (2011). LF; on Aspicilia sp.
Lobarina scrobiculata (Scop.) Nyl. ex Cromb.
Ref.: Sohrabi & Alstrup, (2007); Seaward et al., (2008) as Lobaria scrobiculata (Scope.) DC.
Lobothallia alphoplaca (Wahlenb.) Hafellner
L; on siliceous rock; V: 4697.
Lobothallia radiosa (Hoffm.) Hafellner
L; on siliceous rocks; I: 3593, 3540; II: 1425; III: 4184; VI: 4362.
Marchandiomyces aurantiacus (Lasch) Diederich & Etayo
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Megaspora verrucosa (Ach.) Hafellner & V. Wirth
L; on the bark of a deciduous tree; VIII: Sipman 55434 [IRAN, B 60 0175544].
Megaspora rimisorediata Valadbeigi & A. Nordin
Ref.: Valadbeigi, Nordin, et al. (2011). L; on bark of deciduous tree.
Melanelixia glabra (Schaer.) O. Blanco et al.
Ref.: Sohrabi et al. (2007); Seaward et al. (2008). L; epiphytic; VIII: Sipman 55440 [IRAN, B 60 0175552]. TLC: lecanoric acid.
Melanelixia subargentifera (Nyl.) O. Blanco et al.
Ref.: Seaward et al. (2008); Sohrabi et al. (2007). L; epiphytic; VIII: Sipman 55456 [IRAN, B 60 0175568].
Melanelixia subaurifera (Nyl.) O. Blanco et al.
Ref.: Seaward et al. (2008); Sohrabi et al. (2007).
Melanohalea exasperatula (Nyl.) O. Blanco et al.
Ref.: Seaward et al. (2008); Sohrabi et al. (2007). L; epiphytic: VIII: Sipman 55460 [IRAN, B 60 0175572].
Melanohalea infumata (Nyl.) O. Blanco et al.
Ref.: Seaward et al. (2008); Sohrabi et al. (2007).
Melaspilea arthonioides (Fée) Nyl.
New to Iran;Syn.: Melaspilea enteroleuca (Ach.) Ertz & Diederich sensu Ertz et al. (2015); L; on Carpinus base; VIII: Sipman 55428 [IRAN, B 60 0175538].
Milospium graphideorum (Nyl.) D. Hawksw.
LF; on Opegrapha sp.;V: 4503.
Monerolechia badia (Fr.) Kalb
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008) as Buellia badia(Fr.) A. Massal.
Muellerella pygmaea (Körb.) D. Hawksw.
LF; on Dimelaena oreina; I: 3539; VI: 4302.
Mullerella ventosicola (Mudd) D. Hawksw.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Mycobilimbia tetramera (De Not.) Vitik. et al.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Myriolecis crenulata (Hook.) Śliwa, Zhao Xin & Lumbsch
Syn.:Lecanora crenulataHook.L; L; on calcareous rock; VI: 4348.
Myriolecis hagenii (Ach.) Śliwa, Zhao Xin & Lumbsch
Syn.:Lecanora hagenii(Ach.) Ach.; L; on trunk and branches of deciduous trees; II: 1439; V:4584, 4693.
Myriolecis semipallida (H. Magn.) Śliwa, Zhao Xin & Lumbsch
Syn.:Lecanora semipallida H. Magn.;L; on siliceous rock; III: 3759, 4137.
Nephroma parile (Ach.) Ach.
L; on deciduous, rarely conifer trees and on siliceous rock; V: 4686, 4487, 4621.
Niesslia peltigericola (D. Hawksw.) Etayo
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008) as Raciborskiomyces peltigericola (D. Hawksw.) M.E. Barr).
Parmelia saxatilis (L.) Ach.
Ref.: Sohrabi et al. (2007); Seaward et al. (2008).
Parmelia sulcata Taylor
Ref.: Sohrabi et al. (2007); Seaward et al. (2008). L; on mossy rock; VIII: Sipman 55469 [IRAN, B 60 0175580].
Parmeliella triptophylla (Ach.) Müll. Arg.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Parmelina tiliacea (Hoffm.) Hale
Ref.: Sohrabi et al. (2007); Seaward et al. (2008). L; on mossy rock; VIII: Sipman 55462 [IRAN, B 60 0175574].
Peltigera canina (L.) Willd.
L; usually over moss or humus layer on soil, siliceous and calcareous rocks; III: 4066, 4099; V: 4542.
Peltigera collina (Ach.) Schrader
L; on mossy siliceous rock; V: 4464.
Peltigera elisabethae Gyeln.
L; on mossy soil; V: 4557, 4709, 4435.
Peltigera horizontalis (L.) Willd.
L; on humus, often on base-rich soils, mossy rock, and mossy or decayed stumps; II: 98, 1434; III: 4075, 4036; V: 4434, 4719.
Peltigera neckeri Hepp ex Müll. Arg.
L; on mossy siliceous rock; III: 3799, 3775, 4073, 4100; V: 4583, 4499, 4497.
Peltigera ponojensis Gyeln.
L; on stony lime- and dolomite soils as well as base-rich siliceous soils; II: 142; III: 4107.
Peltigera praetextata (Flörke ex Sommerf.) Zopf (Fig. 2)
L; on mossy trunks of trees, on mossy boulders, rarely on soil; II: 1369, 1377; III: 3723, 3704, 3758, 4248, 4104, 4109; V: 4601, 4449, 4666, 4505, 4717, 4438, 4649.
Peltigera rufescens (Weiss) Humb.
L; on dolomite soils, as well as base-rich siliceous soils; I: 3505; II: 111; III: 4089, 4051; V: 4728; VIII: Sipman 55465 [IRAN, B 60 0168955].
Peltigera virescens (J. Steiner) Gyeln.
L; on base-rich siliceous soils; II: 106.
Pertusaria flavicans Lamy
New to Iran; L; on siliceous rocks; II: 1541, 1616; III: 3811, 4129.
Pertusaria pluripuncta Nyl.
New to Iran; L; on siliceous rocks; I: 3557; V: 4593, 4445, 4508.
Pertusaria pseudocorallina (Lilj.) Arnold
New to Iran;L; on siliceous rocks; I: 3508, 3577, 3585, 3500.
Pertusaria xanthoplaca Müll. Arg.
New to Iran; L; on calcareous rock; IV: 3612.
Phaeophyscia ciliata (Hoffm.) Moberg
L; on free-standing deciduous trees; V: 4715; VI: 4319; VIII: Sipman 55451 [IRAN, B 60 0175563].
Phaeophyscia nigricans (Flörke) Moberg
L; on calcareous rock; V: 4350b.
Phaeophyscia orbicularis (Neck.) Moberg
L; on calcareous rock; V: 4615; VI: 4350, 4355.
Phaeophyscia poeltii (Frey) Clauzade & Cl. Roux
New to Iran; L; on the bark of deciduous trees; II: [1418 (UPS)].
Phaeophyscia sciastra (Ach.) Moberg
L; on calcareous rock; V: 4496.
Phaeosporobolus alpinus R. Sant., Alstrup & D. Hawksw.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Phaeosporobolus usneae D. Hawksw. & Hafellner
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Phlyctis argena (Spreng.) Flot.
L; on the stems of deciduous trees; V: 4661, 4685.
Physcia adscendens H. Olivier
L; on free-standing deciduous trees, rather rare on calcareous rock; II: 94, 1532; III: 4173, 4091, 3761, 4112, 4134; V: 4447, 4630.
Physcia aipolia (Ehrh. ex Humb.) Fürnr.
L; on the bark of deciduous trees in open sites; II: [115(UPS)]; III: 4191, 4055, 3785, 4081, 3786; VIII: Sipman 55435 [IRAN, B 60 0175545].
Physcia biziana (A. Massal.) Zahlbr.
L; on the bark of deciduous trees; II: [1449,1551(UPS)].
Physcia caesia (Hoffm.) Fürnr.
L; on calcareous stone; II: [1424 (UPS)]; VI: 4361, 4363.
Physcia dubia (Hoffm.) Lettau
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Physcia stellaris (L.) Nyl.
L; frequently on branches of deciduous trees; II: 1405, 1551, 1538; V: 4454, 4460, 4668, 4517, 4702; VI: 4402, 4350c; VIII: Sipman 55441 [IRAN, B 60 0175553].
Physconia detersa (Nyl.) Poelt
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Physconia distorta (With.) J. R. Laundon
L; on bark of free-standing deciduous trees; II: 86, 1432; III: 3693, 4215, 4160; V: 4481, 4667, 4696, 4576, 4710.
Physconia enteroxantha (Nyl.) Poelt
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Physconia muscigena (Ach.) Poelt
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Physconia perisidiosa (Erichsen) Moberg
L; over mosses; I: 3574b; VIII: Sipman 55436 [IRAN, B 60 0175546].
Placidium lachneum (Ach.) de Lesd.
L; on soil surface of raw soils, soil- or rock-mosses; II: 1476, 1473; III: 3796c; VII: 4262.
Placocarpus schaereri (Fr.) Breuss
L; on limestone, rarely on dolomitic rocks; II: 1385; III: 4116c, 4224, 3752, 4078; V: 4473.
Pleopsidium flavum (Bellardi) Körb.
L; on smooth siliceous rock; II: 1431, 1349; IV: 3660, 3664.
Pleopsidium gobiense (H. Magn.) Hafellner
Ref.: Jamshidi & Shahidi (2015); Shahidi et al. (2013); Jamshidi et al. (2014).
Pleurosticta acetabulum (Neck.) Elix & Lumbsch (Fig. 2)
Ref.: Sohrabi et al. (2007); Seaward et al. (2008). L; epiphytic; VIII: Sipman 55459 [IRAN, B 60 0175571].
Polycoccum pulvinatum (Eithner) R. Sant.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Pronectria robergei (Mont. & Desm.) Lowen
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Protopannaria pezizoides (Weber) P. M. Jørg & S. Ekman
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Protoparmeliopsis muralis (Schreb.) M. Choisy s.lat.
L; on calcareous rock; I: 3582, 3587; III: 3730, 4216, 4031, 4048; V: 4660; VI: 4421.
Protoparmeliopsis peltata (Ramond) Arup, Zhao Xin & Lumbsch
Syn.: Rhizoplaca peltata (Ramond) Leuckert & Poelt; L; on siliceous rocks; II: 1378.
Protothelenella sphinctrinoides (Nyl.) Mayrhofer & Poelt
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Psilolechia lucida (Ach.) M. Choisy
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Psorula rufonigra (Tuck.) G. Schneider
L; on siliceous rocks; II: 1426; III: 3712, 3809.
Pyrenodesmia variabilis (Pers.) A. Massal.,
Syn.: Caloplaca variabilis(Pers.) Müll. Arg.; L; on siliceous rock; I: 3542.
Ramalina capitata (Ach.) Nyl.
L; on wind-exposed siliceous rock; II: 85, 1487; VI: 4280, 4288, 4297.
Ramalina farinacea(L.) Ach.
L; on twigs of deciduous trees; V: 4558.
Ramalina fraxinea (L.) Ach.
L; on free-standing deciduous trees; II: 133, 172; III: 4733, 4158.
Ramalina pollinaria (Westr.) Ach.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Ramalina polymorpha (Lilj.) Ach.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Ramalina sinensis Jatta (Fig. 2)
L; On free-standing deciduous trees with subneutral bark; II:109, 145, 1408, 1412 [1376,1455,157 (TSN)]; III: 3697, 4152, 3702, 4113, 4135, 3777; V: 4676, 4569. TLC: usnic acid.
Rhizocarpon alpicola (Anzi) Rabenh.
L; on siliceous rock; V: 4431.
Rhizocarpon geminatum Körb.
L; on siliceous rock; V: 4513.
Rhizocarpon geographicum (L.) DC.
L; on siliceous rocks; I: 3589, 3594, 3504; II: 1456, 1552, 1470; III: 3802, 4105; V: 4706, 4643, 4582, 4706, 4640; VI: 4366, 4346, 4346.
Rhizocarpon viridiatrum (Wulfen) Körb.
L; on siliceous rocks; III: 4226, 4203.
Rhizoplaca chrysoleuca (Sm.) Zopf
L; on siliceous rocks; V: 4495; VIII: Sipman 55468 [IRAN, B 60 0175579].
Rhizoplaca melanophthalma (Ramond) Leuckert & Poelt
L; on siliceous rock; I: 3534.
Rhizoplaca subdiscrepans (Nyl.) R. Sant.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Rinodina dubyana (Hepp) J.Steiner
L; on calcareous rock; VI: 4377, 4290, 4390.
Rinodina milvina (Wahlenb.) Th.Fr.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Rinodina pyrina (Ach.) Arnold
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Rinodina trachytica (Massal.) Bagl. & Carestia
New to Iran; L; on siliceous rock; III: 4000.
Romjularia lurida (Ach.) Timdal
L; on limestone along cracks covered by soil; II: 1495; VI: 4275, 4409.
Rosellinula frustulosea (Vouaux) R. Sant.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Rosellinula haplospora (Th. Fr. & Almq. ex Th. Fr.) R. Sant.
New to Iran; LF; on Aspicilia sp.; III: 4236.
Rusavskia elegans (Link) S.Y. Kondr. & Kärnefelt
Syn.: Xanthoria elegans (Link) Th.Fr.; L; on siliceous rocks; I: 3537, 3552; II: 1429, 1475; III: 4154; V: 4544, 4680.
Sclerococcum saxatile (Schaer.) Ertz & Diederich
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008) as Dactylospora saxatilis (Schaer.) Hafellner.
Scoliciosporum umbrinum (Ach.) Arnold
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Scytinium aragonii (Otálora) Otálora, P.M. Jørg. & Wedin
New to Iran; L; on soil and mosses; V: 4658.
Scytinium gelatinosum (With.) Otálora, P. M. Jørg. & Wedin
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008) as Leptogium gelatinosum (With.) J. R. Laundon).
Scytinium lichenoides (L.) Otálora, P. M. Jørg. & Wedin
Syn.:Leptogium lichenoides (L.) Zahlbr.l L; on soil and mosses; I: 1481; V: 4469, 4674, 4540; VI: 4330.
Sphaerellothecium cladoniae Alstrup and Zhurb.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Sphaerellothecium reticulatum (Zopf) Etayo
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008) as Echinothecium reticulatum Zopf).
Squamarina cartilaginea (With.) P. James
L; on a calcareous rock, rarely on calcareous stony soils; IV: 3606. 3655.
Staurothele fissa (Taylor) Zwackh
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Stigmidium congestum (Körb.) Triebel
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Stigmidium fuscatae (Arnold) R. Sant.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Stigmidium pumilum (Lettau) Matzer & Hafellner
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Stigmidium tabacinae (Arnold) Triebel
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Stigmidium xanthoparmeliarum Hafellner
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Telogalla olivieri (Vouaux) Nik. Hoffm. & Hafellner
New to Iran; LF; on Xanthoria parietina; II: 1529.
Thalloidima candidum (Weber) A.Massal.
Syn.:Toninia candida (Weber) Th. Fr.; L; on lime-rich to weakly calcareous rocks, especially in fissures; VI: 4349.
Thalloidima sedifolium (Scop.) Kistenich, Timdal, Bendiksby
Syn.:Toninia sedifolia(Scop.) Timdal; L; on calcareous soils; II: 1444, 1513; III: 4111, 4176; V: 4456; VI: 4420; VII: 4260, 4264.
Thelenella muscorum (Fr.) Vain.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Toninia tristis (Th. Fr.) Th. Fr.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Tornabea scutellifera (With.) J. R. Laundon
L; on bark or branches of a deciduous trees; III: 4133, 4221; V: 4723; VIII: Sipman 55458 [IRAN, B 60 0175570].
Tremella ramalinae Diederich
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Umbilicaria cinerascens (Arnold) Frey
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008). TLC: gyrophoric acid.
Umbilicaria cinereorufescens (Schaer.) Frey
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008). TLC: gyrophoric acid.
Umbilicaria freyi Codogno, Poelt & Puntillo
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008). TLC: gyrophoric acid.
Umbilicaria hirsuta (Sw.) Hoffm.
L; on siliceous rock; III: 3726. TLC: gyrophoric and lecanoric acids
Umbilicaria polyphylla (L.) Baumg.
L; on siliceous rock; V: 4535. TLC: umbilicaric and gyrophoric acids.
Umbilicaria vellea (L.) Hoffm.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008). TLC: gyrophoric and lecanoric acids.
Usnea glabrata (Ach.) Vain.
New to Iran;L; on bark or branches of a deciduous tree; III: 4125, 4182, 4151, 4057. TLC: protocetraric, fumarprotocetraric, salazinic, and norstictic acids.
Vahliella leucophaea (Vahl) P.M. Jørg.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008) as Fuscopannaria leucophaea (Vahl) P. M. Jørg.
Varicellaria lactea (L.) I. Schmitt & Lumbsch
New to Iran; L: on siliceous rocks; V: 4491, 4552, 4484.
Verrucaria fuscella (Turner) Winch
L; on limestone; VI: 4370.
Verrucaria tristis (A. Massal.) Trevis.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Xanthocarpia tominii (Savicz) Frödén, Arup & Søchting
New to Iran; L; on soil, VIII: Søchting US11067, US11072 [IRAN].
Xanthomendoza fallax Søchting, Kärnefelt & S.Y. Kondr
Syn.: Oxneria fallax (Arnold) S.Y. Kondr. & Kärnefelt L; on steep surfaces of weakly calcareous or siliceous rock; III: 3745, 4117, 4088; V: 4690, 4721; VI: 4396.
Xanthomendoza fulva (Hoffm.) Søchting, Kärnefelt & S. Kondratyuk
Syn.: Gallowayella fulva (Hoffm.) S.Y. Kondr. et al.; L; on free-standing deciduous trees; V: 4567.
Xanthoparmelia camtschadalis (Ach.) Hale (Fig 2)
Ref.: Sohrabi et al. (2007); Seaward et al. (2008).
Xanthoparmelia conspersa (Ach.) Hale
Ref.: Sohrabi et al. (2007); Seaward et al. (2008).
Xanthoparmelia loxodes (Nyl.) O. Blanco et al.
Ref.: Sohrabi et al. (2007); Seaward et al. (2008).
Xanthoparmelia pokornyi (Körb.) O. Blanco et al.
Ref.: Sohrabi et al. (2007); Seaward et al. (2008).
Xanthoparmelia pulla (Ach.) O. Blanco et al.
Ref.: Sohrabi et al. (2007); Seaward et al. (2008).
Xanthoparmelia ryssolea (Ach.) O. Blanco et al.
L; vagrant on soil surfaces, on calcareous soil; VI: 4391a.
Xanthoparmelia stenophylla (Ach.) Ahti & D. Hawksw.
Ref.: Sohrabi et al. (2007); Seaward et al. (2008).
Xanthoparmelia tinctina (Maheu & Gillet) Hale
Ref.: Sohrabi et al. (2007); Seaward et al. (2008).
Xanthoparmelia verruculifera (Nyl.) O. Blanco et al.
Ref.: Sohrabi et al. (2007); Seaward et al. (2008).
Xanthoria parietina (L.) Th.Fr.
L; on the bark of deciduous trees and shrubs; II: 173, 1536, 1546; III: 3716; IV: 3616, 3661; V: 4578.
Xanthoriicola physciae (Kalchbr.) D. Hawksw.
Ref.: Sohrabi & Alstrup (2007); Seaward et al. (2008).
Fig. 2. Selected lichens from Arasbaran: A. Xanthoparmelia camtschadalis (Location VII); B. Pleurosticta acetabulum (Location III); C. Cladonia subrangiformis (Location II); D. Peltigera praetextata (Location III); E. Circinaria fruticulosa (Location VII); F. Ramalina sinensis (Location III).
DISCUSSION
Of the 260 taxa recorded here, 20 taxa are additional to Seaward et al. (2008). The most updated Iranian lichen checklist contains c. 935 taxa (ca. 855 are lichenized and ca. 80 lichenicolous or allied fungi) (Sohrabi et al. unpublished data). According to Urbanavichus (2015), 1840 taxa of lichenized fungi are known for the whole Caucasus region (including c. 200 lichenicolous and 30 saprotrophic non-lichenized fungi) and the author predicts c. 3500 taxa of lichens for the Caucasus hotspot region. Moreover, currently, more than 730 and 824 lichen taxa are reported from Georgia and Azerbaijan respectively (Barkhalov 1975; Barkhalov 1983); Nakhutsrishvili 1986; Inashvili & Batsatsashvili 2010; Novruzov & Alverdieva 2013). The lichen checklist of Armenia includes 422 taxa (Harutyunyan et al. 2011; Gasparyan & Sipman 2016).
In comparison with the NW neighboring countries of Iran (Armenia and Azerbaijan), the lichen diversity of Arasbaran is still poorly explored. In 2015, as a result of the international lichenological excursion to Armenia, 176 species of lichenized and lichenicolous fungi were reported from Khosrov Forest State Reserve, Armenia (Gasparyan et al. 2015) at a distance of c. 200 km from Arasbaran.
The majority of the species located within the holarctic floristic kingdom are mainly distributed in the circumboreal and Irano-Turanian regions, while some have a widespread distribution within broad altitudinal and latitudinal ranges in different continents, traditionally considered to be cosmopolitan or subcosmopolitan (Nimis 2016). Those with a bipolar distribution are Aspicilia cinerea, Candelaria concolor, Candelariella vitellina, Cetrelia olivetorum, Chaenotheca furfuracea, Protoparmeliopsis muralis, Lecanora rupicola, Diploschistes muscorum, Ingvariella bispora, Physcia adscendens, Umbilicaria polyphylla, Lepraria lobificans, Lepraria vouauxii, Lobothallia radiosa, Parmelia sulcata, Peltigera canina, Rusavskia elegans, Rhizocarpon geographicum, and Bryobilimbia hypnorum. It is possible to differentiate many taxa into phytoclimatic groups, such as:
Arctic-alpine: Lecidella carpathica, Pleopsidium flavum, Umbilicaria cinerascens, U. cinereorufescens, and U. polyphylla.
Boreal-montane: Acarospora fuscata, Bryoria nadvornikiana, Cetraria aculeata, C.islandica, Circinaria calcarea, Cladonia borealis, Evernia prunastri, Hypogymnia physodes, Lepraria membranacea, Toninia tristis and Vahliella leucophaea
Submediterranean to mediterranean: Buellia zoharyi, Caloplaca phaeothamnos Cetraria aculeata, Cladonia magyarica, C. subrangiformis, Dermatocarpon moulinsii, D. vellereum, Lepraria leuckertiana, L. crassissima, Pertusaria pseudocorallina and Umbilicaria freyi.
An understanding of the taxonomy, ecology, and biogeography of the Caucasus and Iranian lichens (including the Arasbaran region) will be a long-term research project. Still, the mycobiota (lichenized and lichenicolous fungi) of Iran and neighboring countries are poorly explored and in general, this area hides numerous unreported and undescribed species. This paper is summarising the current knowledge of the lichenized and lichenicolous fungi in the Arasbaran Biosphere Reserve. As such it presents a small contribution to our understanding of the biota of Iran and the Caucasus region and provides additional distributional data for some poorly know lichenized and lichenicolous species.
ACKNOWLEDGEMENTS
The first author wishes to present his sincere thanks to Prof. Teuvo Ahti (Helsinki, Finland), who taught him lichenological methods, for his versatile technical assistance and various valuable comments during the identification of specimens, as well as for reviewing this manuscript. Gratefully acknowledged are also Masoomeh Ghobad-Nejhad (Tehran, Iran) for her help and cooperation and readiness to assist, and Prof. M.R.D. Seaward (Bradford, UK) for thorough correction of the manuscript. The thin-layer chromatography (TLC) work was financially supported by the Iranian Research Organization for Science and Technology (IROST) through grant no. 280121-96/1/15. The curators of H, GB, C, TUR, and S are gratefully acknowledged for making valuable material available to us. The senior author would like to thank Ali Shahed Anakhatoun, Davoud and Jafar Soleymani and Massoomeh Ghobad-Nejhad, for their help during fieldwork. The cost of the fieldwork for the first author between 2004-2006 was covered by Rostam Sohrabi (Jolfa, Iran).
Conflict of interest statement
The authors declare no conflict of interests.