As a result of lichenological exploration of the Erzi Nature Reserve in the Republic of Ingushetia, North Caucasus, Russia, we report 266 species of lichenized fungi, 13 of lichenicolous fungi and seven of non-lichenized saprophytic fungi, representing 115 genera. Twenty-three species are recorded for the first time for North Caucasus, of which 16 are new for the Caucasus and ten are new for Russia (Arthonia protoparmeliopseos, Aspicilia viridescens, Buellia abstracta, Lecidella viridans, Micarea soralifera, Rinodina straussii, R. tunicata, R. zwackhiana, Schizoxylon alboatrum and Verrucaria fuscoatroides). Of the 16 species new for the Caucasus, seven are recorded for the first time for Asia (Buellia abstracta, Micarea nigella, M. soralifera, Ramonia chrysophaea, Rinodina tunicata, Schizoxylon alboatrum and Verrucaria elevata). In addition, two taxa listed in the Red Data Book of Russia are recorded: Leptogium burnetiae and Usnea florida.
Version of record first published online on 9 November 2017 ahead of inclusion in December 2017 issue.
Introduction
The Caucasus is one of the world's biodiversity hotspots (Krever & al. 2001) and one of the most diverse regions of Russia (Geltman & al. 1998). In contrast to the relatively well-studied lichen flora of the W Caucasus, its C and E parts (including the Republic of Ingushetia) are still poorly explored. In North Caucasus, seven Nature Reserves include the most important natural ecosystems with great biological diversity and natural features of exceptional scientific interest. The lichen flora of six of these has been studied (Urbanavichus & Urbanavichene 2004, 2015, 2017; Urbanavichus & al. 2010); only the Erzi Nature Reserve remains to be studied. The present study aims at improving our knowledge of the Reserve's lichen flora and, together with other recent contributions on Caucasian lichens, to serve as an official documentation of newly discovered/reported species for the region and for Russia.
Research area
The Erzi Nature Reserve (Kotiev & al. 2015) covers an area of 35,292 ha and is located on the N slope of the Greater Caucasus in the Sunhza and Dzheirakh districts of the Republic of Ingushetia, c. 35 km S of Magas city (Fig. 1). In Russia the Reserve is bordered by the Republic of Northern Ossetia (Severnaya Osetiya) and the Republic of Chechnya along the main Caucasian ridge, with the Republic of Georgia to the south. The altitude ranges from 850 to 3031 m (the highest peak Mt Khakhalgi). The landscape features mountain range spurs over deep river valleys, steep S slopes and gently sloping N slopes. Rivers in the Reserve include the Assa and Armkhi, which feed into the Terek river. The outcropping rocks are predominantly of shales and sandstones of Miocene age; Jurassic calcareous rock formations are rare. About onethird of the Reserve is forested, mostly on the lower N slopes. The forests are mostly oak (Quercus petraea (Matt.) Liebl.) and birchpine (Betula pendula Roth., Pinus kochiana Klotzsch ex K. Koch) communities. Underbrush at the mid-level includes sea buckthorn (Hippophae rhamnoides L.), willow (Salix caprea L.), grey alder (Alnus incana (L.) Moench) and in some places extensive hazel (Corylus avellana L.). Approaching the sub-alpine zone the brush features Caucasian rhododendron (Rhododendron caucasicum Pall.). Above 2000 m, but below the alpine zone at 2500 m, is a belt of mountain forest-steppe: wheatgrass-wormwood-cereal xerophytic steppes on shallow meadow soils. Although the Reserve is rather small, it supports about 18 % (c. 1100 species) of the Caucasian flora. The climate of Erzi is characterised by long, cold winters, and short, cool summers. The average number of days of precipitation per year is 113. The average temperature is -1.4°C in January and 26.4°C in July. The average annual precipitation ranges from 350–400 mm below 1200 m altitude to 500–600 mm above 2000 m. Over 160 historical and cultural sites (martial towers, temples, necropolis tracts, sacred groves, and structures from ancient, medieval and later periods) are also protected by the Reserve.
Material and methods
The lichen specimens were collected in the following seven localities:
1 Right-hand bank of Tetritskali river, 42°49′53″N, 44°55′19″E, 1280–1290 m, Pinus kochiana forest, 29 June 2016.
2 Targim tower complex on right-hand bank of Assa river, 42°50′00″N, 44°56′29″E, 1080 m, floodplain forest with Alnus incana, Hippophae rhamnoides and Salix caprea, 30 June 2016.
3 Between Leymi and Egikhal tower complexes, 42°50′02-07″N, 44°54′28-44″E, 1340–1450 m, Pinus kochiana forest and stony SE slope, 30 June 2016.
4 Right-hand bank of a nameless stream (right tributary of Tetritskali river), 42°49′43″N, 44°54′14″E, 1350–1360 m, Alnus incana and Salix caprea forest, 1 July 2016.
5 Around Ozdig tower complex, 42°49′30″N, 44°54′52″E, 1500–1600 m, Pinus kochiana and Betula pendula forest with Populus tremula, Pyrus sp. and Tilia cordata Mill. on N slope, 1 & 5 July 2016.
6 Left-hand bank of Assa river, 42°49′55″N,44°56′18″E, 1090–1100 m, steppe meadow and grassland with calcareous and sandstones boulders, 2 & 6 July 2016.
7 Vovnushki tower complex on right-hand bank of Guloykhi river, 42°48′01″N, 44°59′41″E, shale cliffs and single trees of Alnus incana with bushes of Juniperus communis L., 1220–1250 m, 4 July 2016.
Our results are based on investigations from the field trips made between 29 June and 6 July 2016. Air-dried samples were examined using a stereo microscope, a light microscope and the usual spot tests with standard identification methods for lichenized and lichenicolous fungi; thin layer chromatography (TLC) was undertaken where necessary (Orange & al. 2001). A bullet “•” marks the analysed specimens. Remarkable species and species difficult to identify in the field were collected. Common macrolichens and some crustose lichens were not collected in the field, only registered in each locality. The specimens are kept in the herbarium of the Komarov Botanical Institute, St. Petersburg (LE), and in the private herbarium of G. Urbanavichus (“herb. Urbanav.”). All taxa listed here were collected by the authors. For nomenclatural aspects, mainly Hafellner & Türk (2016) and Lawrey & Diederich (2016) were followed, except for some of the Teloschistaceae taxa.
Results
Checklist of taxa
Symbols used before a species name are as follows: # = lichenicolous fungi; + = non-lichenized fungi; * = new for Russia; † = new for the Caucasus; ‡ = new for North Caucasus (i.e. not new for the whole Caucasus); ! = new for Asia; and after species name “•” = analysed specimens (see Material and methods).
Species names are cited with their authorship. They are followed by locality numbers (loc. 1–7, see Material and methods) and, in parentheses under each locality, collection numbers (e.g. IU ER-16/1-009) and the herbarium (LE or herb. Urbanav.). Collection numbers of the first author are preceded by GU and those of the second author by IU.
Substrates are indicated, with abbreviations as follows: Ai = Alnus incana, Bp = Betula pendula, Cb = Carpinus betulus L., Ca = Corylus avellana, Jc = Juniperus communis, Pk = Pinus kochiana, Pt = Populus tremula, Pr = Prunus sp., Py = Pyrus sp., Qp = Quercus petraea, Sa = Sorbus aucuparia L., Sc = Salix caprea, Sn = Sambucus nigra L., Tc = Tilia cordata.
For certain species, additional comments are provided.
#Abrothallus microspermus Tul. — loc. 1 (IU ER-16/1-009, LE); on thallus of Flavoparmelia caperata on Pk.
Absconditella lignicola Vězda & Pišut — loc. 1 (IU ER-16/1-010, LE), 3 (GU ER-16/3-127, herb. Urbanav.), 5 (GU ER-16/5-257, LE); on wood of pine log, stump.
Acarospora badiofusca (Nyl.) Th. Fr. — loc. 7 (GU ER-16/7-445, herb. Urbanav.); on shale.
Acarospora cervina A. Massal. — loc. 3, 6 (in sample GU ER-16/6-379, herb. Urbanav.); on calcareous rocks.
Acarospora veronensis A. Massal. — loc. 7 (GU ER-16/7-446, herb. Urbanav.); on shale.
Acarospora versicolor Bagl. & Carestia — loc. 6 (GU ER-16/6-366, herb. Urbanav.); on sandstones.
Acrocordia gemmata (Ach.) A. Massal. — loc. 2 (GU ER-16/2-048, herb. Urbanav.); on Ai.
Agonimia opuntiella (Buschardt & Poelt) Vězda — loc. 6, 7 (GU ER-16/7-447, herb. Urbanav.); on plant debris.
Agonimia tristicula (Nyl.) Zahlbr. — loc. 3, 6 (GU ER-16/6-368, herb. Urbanav.); on plant debris.
Alyxoria varia (Pers.) Ertz & Tehler — loc. 2 (GU ER-16/2-091, herb. Urbanav.); on Ai.
Amandinea punctata (Hoffm.) Coppins & Scheid. — loc. 1 (IU ER-16/1-011, LE), 5, 7 (in sample GU ER-16/7-511, herb. Urbanav.); on Jc, pine log, and rocks.
Anaptychia ciliaris (L.) Körb. — loc. 2, 4, 5; on Ai, Pt, Py.
Anema tumidulum Henssen ex P. M. Jørg. & al. — loc. 3 (GU ER-16/3-130, herb. Urbanav.); on calcareous rocks.
Arthonia atra (Pers.) A. Schneid. — loc. 4 (IU ER-16/4-236, LE); on Sc.
#Arthonia epiphyscia Nyl. — loc. 2 (GU ER-16/2-050, herb. Urbanav.); on thallus of Physcia aipolia on Ai.
Arthonia fusca (A. Massal.) Hepp — loc. 3 (GU ER-16/3-131, herb. Urbanav.), 6 (GU ER-16/6-369, herb. Urbanav.); on calcareous rocks.
#Arthonia phaeophysciae Grube & Matzer — loc. 2 (GU ER-16/2-051, herb. Urbanav.); on thalli of Phaeophyscia hirsuta and Phaeophyscia orbicularis on Ai.
*†#Arthonia protoparmeliopseos Etayo & Diederich — loc. 6 (GU ER-16/6-370, LE); on apothecia of Protoparmeliopsis muralis on calcareous rocks. Described from Luxembourg and Spain (Etayo & Diederich 2009) and recently reported from Turkey (Yazici & Etayo 2014) and North America (Kocourková & Knudsen 2015).
Arthonia punctiformis Ach. — loc. 5 (GU ER-16/5-260, herb. Urbanav.); on Tc.
Arthonia radiata (Pers.) Ach. — loc. 2, 4 (IU ER-16/4-212, LE); on Ai.
Arthopyrenia analepta (Ach.) A. Massal. — loc. 5 (GU ER-16/5-261, herb. Urbanav.); on Tc.
Arthrosporum populorum A. Massal. — loc. 2 (GU ER-16/2-053, herb. Urbanav.); on Ai.
Aspicilia cf. desertorum (Kremp.) Mereschk. nom. illeg. — loc. 6 (GU ER-16/6-372, herb. Urbanav.), 7; on calcareous sandstones.
*‡Aspicilia viridescens (A. Massal.) Hue — loc. 7 (GU ER-16/7-450, herb. Urbanav.); on shale. Also recorded from Armenia in Transcaucasia (Harutyunyan & al. 2011).
Athallia pyracea (Ach.) Arup & al. — loc. 2; on Ai.
Bacidia pycnidiata Czarnota & Coppins — loc. 1, 4 (IU ER-16/4-213, LE); on mosses Sc and pine log.
Bacidia rubella (Hoffm.) A. Massal. — loc. 4 (IU ER-16/4-214, LE), 5; on Ai, Pt, Sc.
‡Bacidina arnoldiana (Körb.) V. Wirth & Vězda — loc. 3 (GU ER-16/3-132, herb. Urbanav.); on calcareous rocks. Recently reported from Armenia in Transcaucasia (Gasparyan & al. 2015).
Bagliettoa calciseda (DC.) Gueidan & Cl. Roux — loc. 6 (GU ER-16/6-373, herb. Urbanav.); on calcareous rocks.
Blastema hungarica (H. Magn.) Arup & al. — loc. 1 (IU ER-16/1-016, LE), 3; on Jc, Pk.
Blennothallia crispa (Huds.) Otálora & al. — loc. 7; on soil.
Bryoria capillaris (Ach.) Brodo & D. Hawksw. — loc. 5 (IU ER-16/5-263, LE); on Pk.
Bryoria fuscescens (Gyeln.) Brodo & D. Hawksw. — loc. 5 (IU ER-16/5-264, LE); on Pk, Tc.
*†!Buellia abstracta (Nyl.) H. Olivier — loc. 6 (GU ER-16/6-374, LE); on sandstones. Also known from Eu-rope and North and South America (Giralt & al. 2011; Knudsen & Kocourková 2012). Our specimens lackvisible thallus; the spores are 4.5-5 × 10-12 µm.
Buellia disciformis (Fr.) Mudd — loc. 4 (IU ER-16/4-215, LE); on Ai.
Buellia erubescens Arnold — loc. 2 (GU ER-16/2-054, herb. Urbanav.); on Ai.
Buellia griseovirens (Turner & Borrer ex Sm.) Almb. — log. 1, 3; on Jc, Pk.
Calicium glaucellum Ach. — loc. 5 (IU ER-16/5-265, LE); on Pk.
Calicium lucidum (Th. Fr.) M. Prieto & Wedin — loc. 1, 3 (IU ER-16/3-154, LE); on Pk.
Calicium trabinellum (Ach.) Ach. — log. 3, 5; on Pk.
Callome multipartita (Sm.) Otálora & al. — loc. 3, 6 (GU ER-16/6-396, herb. Urbanav.); on calcareous rocks.
Calogaya biatorina (A. Massal.) Arup & al. — log. 3, 6, 7; on shale, calcareous rocks.
Calogaya pusilla (A. Massal.) Arup & al. — loc. 7 (GU ER-16/7-454, herb. Urbanav.); on shale.
Calogaya saxicola (Hoffm.) Vondrák — loc. 2, 7 (GU ER-16/7-455, herb. Urbanav.); on calcareous sandstones, shale.
Caloplaca asserigena (J. Lahm) Della Torre & Sarnth. — loc. 1, 5 (IU ER-16/5-267, LE); on Pk.
Caloplaca cerina (Hedw.) Th. Fr. — loc. 2, 5, 6, 7; on Ai, Jc, Qp.
Caloplaca conversa (Kremp.) Jatta — loc. 6; on calcareous rocks.
Caloplaca inconnexa (Nyl.) Zahlbr. — loc. 6 (GU ER-16/6-379, herb. Urbanav.); on thallus of Acarospora cervina on calcareous rocks.
Caloplaca obscurella (J. Lahm ex Körb.) Th. Fr. — loc. 4 (IU ER-16/4-216, LE); on Ai.
Caloplaca raesaenenii Bredkina — loc. 6 (GU ER-16/6-386, herb. Urbanav.); on plant debris.
Caloplaca stillicidiorum (Vahl) Lynge — loc. 6; on plant debris.
Candelaria concolor (Dicks.) Stein — loc. 2, 5, 7; on Ai, Py, shale.
Candelariella aurella (Hoffm.) Zahlbr. — loc. 3, 6, 7; on Jc, calcareous rocks, shale.
Candelariella efflorescens R. C. Harris & W. R. Buck — loc. 1 (IU ER-16/1-017, LE, dubl. in ALTB), 2, 4, 5, 6; on Ai, Jc, Pk, Tc.
Candelariella lutella (Vain.) Räsänen — loc. 4, 5; on Ai.
Candelariella oleaginescens Rondon — loc. 6; on calcareous rocks.
Candelariella rhodax Poelt & Vězda — loc. 3, 6 (GU ER-16/6-392, herb. Urbanav.), 7; on calcareous rocks, shale.
Candelariella vitellina (Hoffm.) Müll. Arg. — loc. 7; on shale.
Catillaria nigroclavata (Nyl.) Schuler — loc. 2 (GU ER-16/2-062, herb. Urbanav.); on Ai.
†#Cercidospora epicarphinea (Nyl.) Grube & Hafellner — loc. 6 (GU ER-16/6-393, herb. Urbanav.); on apothecia of Pyrenodesmia variabilis on calcareous rocks. Already reported from NW Russia (Alstrup & al. 2005) and S Ural mountains (Merkulova & Urbanavichus 2006).
Cetrelia cetrarioides (Delise & Duby) W. L. Culb. & C. F. Culb. • — loc. 4 (IU ER-16/4-219, LE), 5; on Ai, Tc. (TLC — perlatolic acid).
Cetrelia olivetorum (Nyl.) W. L. Culb. & C. F. Culb. • — loc. 2 (GU ER-16/2-063, herb. Urbanav.), 5; on Ai, Tc. (TLC — olivetoric acid).
Chaenotheca chrysocephala (Turner ex Ach.) Th. Fr. — loc. 1, 5; on Pk.
Chaenotheca furfuracea (L.) Tibell — loc. 5; on the roots of Bp.
Chaenotheca trichialis (Ach.) Th. Fr. — loc. 5; on pine wood.
Chaenotheca xyloxena Nádv. — loc. 5 (IU ER-16/5-278, LE); on pine wood.
+Chaenothecopsis pusiola (Ach.) Vain. — loc. 5 (IU ER-16/5-279, LE); on pine wood.
+Chaenothecopsis savonica (Räsänen) Tibell — loc. 5 (IU ER-16/5-280, LE); on pine wood.
+Chaenothecopsis viridireagens (Nádv.) A. F. W. Schmidt — loc. 5 (IU ER-16/5-281, LE); on pine wood.
Chrysothrix candelaris (L.) J. R. Laundon — loc. 4, 5; on Sc, Tc.
Circinaria calcarea (L.) A. Nordin & al. — loc. 6; on calcareous rocks.
Cladonia coniocraea (Flörke) Spreng. — loc. 1, 5; on Pk.
Cladonia foliacea (Huds.) Willd. — loc. 2, 3, 6, 7; on soil.
Cladonia fimbriata (L.) Fr. — loc. 1, 3, 5; on Pk.
Cladonia macilenta Hoffm. — loc. 3; on pine log.
Cladonia symphycarpa (Flörke) Fr. — loc. 3; on soil.
Coenogonium pineti (Ach.) Lücking & Lumbsch — loc. 5; on Tc.
Collema flaccidum (Ach.) Ach. — loc. 5; on Pt, Tc.
Collema furfuraceum (Arnold) Du Rietz — loc. 7; on plant debris on shale.
Collema subflaccidum Degel. — loc. 7; on plant debris on shale.
Cryptodiscus foveolaris (Rehm) Rehm — loc. 5 (IU ER-16/5-287, LE); on pine wood.
Dermatocarpon miniatum (L.) W. Mann — loc. 1, 2, 3, 5, 6, 7; on shale, calcareous rocks.
Dimelaena oreina (Ach.) Norman — loc. 2, 6, 7 (GU ER-16/7-470, herb. Urbanav.); on shale, small granite stones.
Diploschistes muscorum (Scop.) R. Sant. — loc. 7; on soil.
Diplotomma hedinii (H. Magn.) P. Clerc & Cl. Roux — loc. 6 (GU ER-16/6-401, herb. Urbanav.); on calcareous rocks.
‡Diplotomma lutosum A. Massal. — loc. 7 (GU ER-16/7-472, herb. Urbanav.); on shale. Also recorded from Georgia in Transcaucasia (Barkhalov 1983).
Enchylium polycarpon (Hoffm.) Otálora & al. — loc. 6, 7; on calcareous rocks.
Enchylium tenax (Sw.) Gray — loc. 3, 7; on calcareous soil.
Endocarpon pallidum Ach. — loc. 3 (GU ER-16/3-156, herb. Urbanav.); on calcareous rocks.
Endocarpon pusillum Hedw. — loc. 7 (GU ER-16/7-473, herb. Urbanav.); on soil.
#Endococcus pseudocarpus Nyl. — loc. 6 (GU ER-16/6-402, herb. Urbanav.); on thallus of Callome multipartita on calcareous rocks.
Evernia divaricata (L.) Ach. — loc. 5; on Pr.
Evernia prunastri (L.) Ach. — loc. 1, 3, 5; on Bp, Pk, Pr, Py, Tc, snag.
Flavoparmelia caperata (L.) Hale — loc. 1 (in sample IU ER-16/1-009, LE), 2, 3, 4, 5, 6, 7; on Ai, Cb, Jc, Pk, Pr, Py, Sc, Tc.
Flavoplaca coronata (Kremp. ex Körb.) Arup & al. — loc. 1, 2, 3, 5, 6, 7; on calcareous rocks.
Flavoplaca flavocitrina (Nyl.) Arup & al. — loc. 3, 6 (GU ER-16/6-382, herb. Urbanav.); on calcareous sandstones, calcareous rocks.
Flavoplaca oasis (A. Massal.) Arup & al. — loc. 6 (in sample GU ER-16/6-373, herb. Urbanav.); on thallus of Bagliettoa calciseda on calcareous rocks.
Flavopunctelia flaventior (Stirt.) Hale — loc. 1, 2, 3, 4, 5, 6, 7; on Ai, Jc, Pk, shale.
Flavopunctelia soredica (Nyl.) Hale — loc. 1, 2, 3, 4, 5, 6, 7; on Ai, Jc, Pk, Tc, shale.
Graphis scripta (L.) Ach. — loc. 2, 4, 5; on Ai, Bp, Tc.
†Gyalideopsis helvetica van den Boom & Vězda — loc. 5 (GU ER-16/5-295, LE); on wood of pine log. Also recently reported from S Siberia (Urbanavichene 2015).
Gyalolechia flavorubescens (Huds.) Søchting & al. — loc. 2; on Ai.
Gyalolechia flavovirescens (Wulfen) Søchting & al. — loc. 3, 6 (in sample GU ER-16/6-370, LE); on calcareous sandstones, sandstones.
Heterodermia speciosa (Wulfen) Trevis. — loc. 4; on Ai.
Heteroplacidium compactum (A. Massal.) Gueidan & Cl. Roux — loc. 6 (GU ER-16/6-406, herb. Urbanav.); on calcareous rocks.
Hyperphyscia adglutinata (Flörke) H. Mayrhofer & Poelt — loc. 2, 7; on Ai, Jc, shale.
†Hyperphyscia granulata (Poelt) Moberg — loc. 7 (GU ER-16/7-478, herb. Urbanav.); on shale. Also reported from S Siberia (Urbanavichus & Urbanavichene 2007).
Hypocenomyce scalaris (Ach.) M. Choisy — loc. 3; on Pk.
Hypogymnia physodes (L.) Nyl. — loc. 1, 3, 4, 5; on Pk, Sc.
Hypogymnia tubulosa (Schaer.) Hav. — loc. 1, 3, 5; on Pk.
Lathagrium auriforme (With.) Otálora & al. — loc. 7; on mossy calcareous rocks.
Lathagrium cristatum (L.) Otálora & al. — loc. 2, 3, 6, 7; on calcareous rocks, calcareous sandstones.
Lathagrium fuscovirens (With.) Otálora & al. — loc. 3, 7; on calcareous rocks, shale.
Lathagrium undulatum (Flot.) Otálora & al. — loc. 6; on calcareous rocks.
Lecania croatica (Zahlbr.) Kotlov — loc. 4 (IU ER-16/4-227, LE), 5; on Ai, Sc.
Lecania naegelii (Hepp) Diederich & van den Boom — loc. 2 (in sample GU ER-16/2-051, herb. Urbanav.), 4; on Ai.
Lecanora albella (Pers.) Ach. — loc. 1, 5 (IU ER-16/1-029, LE, GU ER-16/5-299, herb. Urbanav.); on Pk, Tc.
Lecanora albellula (Nyl.) Th. Fr. — loc. 3; on pine wood.
Lecanora allophana Nyl. — loc. 2, 5; on Ai, Tc.
Lecanora argopholis (Ach.) Ach. — loc. 7 (GU ER-16/7-479, herb. Urbanav.); on shale.
Lecanora chlarotera Nyl. — loc. 2; on Ai.
Lecanora intumescens (Rebent.) Rabenh. — loc. 5 (IU ER-16/5-301, LE); on Pr, Pt, Tc.
Lecanora meridionalis H. Magn. — loc. 2 (GU ER-16/2-077, herb. Urbanav.); on Ai.
Lecanora populicola (DC.) Duby — loc. 5; on Pt.
‡Lecanora reagens Norman — loc. 7 (GU ER-16/7-482, herb. Urbanav.); on shale. Previously recorded from Georgia in Transcaucasia (Barkhalov 1983).
Lecanora subcarpinea Szatala — loc. 2 (GU ER-16/2-081, herb. Urbanav.); on Ai.
Lecanora symmicta (Ach.) Ach. — loc. 2, 3; on Ai, Pk.
Lecanora thysanophora R. C. Harris • — loc. 4 (IU ER-16/4-229, LE), 5; on Ai, Tc.
‡Lecidea nylanderi (Anzi) Th. Fr. — loc. 3 (GU ER-16/3-170, herb. Urbanav.); on Pk. Previously recorded from Azerbaijan in Transcaucasia (Barkhalov 1983).
Lecidella achristotera (Nyl.) Hertel & Leuckert — loc. 2 (GU ER-16/2-083, herb. Urbanav.); on Ai.
Lecidella flavosorediata (Vězda) Hertel & Leuckert — loc. 2 (GU ER-16/2-084, herb. Urbanav.), 3 (GU ER-16/3-171, herb. Urbanav.), 5; on Ai, Pk.
Lecidella patavina (A. Massal.) Knoph & Leuckert — loc. 6 (GU ER-16/6-410, herb. Urbanav.), 7; on calcareous rocks and shale.
Lecidella stigmatea (Ach.) Hertel & Leuckert — loc. 3, 6 (GU ER-16/6-411, herb. Urbanav.); on calcareous rocks.
*†Lecidella viridans (Flot.) Körb. — loc. 7 (GU ER-16/7-485, herb. Urbanav.); on shale. Also known from many regions of Europe, and North and South America (Knoph & Leuckert 2004).
Lepraria finkii (B. de Lesd.) R. C. Harris • — loc. 5 (IU ER-16/5-307, LE); on Tc.
Leptogium burnetiae C. W. Dodge — loc. 4 (IU ER-16/4-230, LE), 5, 7 (GU ER-16/7-486, herb. Urbanav.); on Sc, Tc, mossy shale.
Leptogium cyanescens (Rabenh.) Körb. — loc. 5 (IU ER-16/5-309, LE); on Tc.
Leptogium saturninum (Dicks.) Nyl. — loc. 4, 5, 7; on Ai, Sn, Tc, mossy shale.
#Lichenochora obscuroides (Linds.) Triebel & Rambold — loc. 2 (in sample GU ER-16/2-051, herb. Urbanav.); on thalli of Phaeophyscia orbicularis and Phaeophyscia pusilloides on Ai.
#Lichenostigma cosmopolites Hafellner & Calat. — loc. 7 (GU ER-16/7-488, herb. Urbanav.); on thallus of Xanthoparmelia stenophylla on shale.
#Lichenothelia renobalesiana D. Hawksw. & V. Atienza — loc. 6 (in sample GU ER-16/6-373, herb. Urbanav.); on thallus of Bagliettoa calciseda on calcareous rocks.
†Lichinella stipatula Nyl. — loc. 3, 6 (GU ER-16/6-413, herb. Urbanav.); on calcareous rocks. Also known from S Ural mountains and S Siberia (Urbanavichus 2010).
Lobothallia alphoplaca (Wahlenb.) Hafellner — loc. 2; on sandstones.
Lobothallia radiosa (Hoffm.) Hafellner — loc. 2, 3, 6; on calcareous rocks and sandstones.
Melanelixia albertana (Ahti) O. Blanco & al. — loc. 4 (IU ER-16/4-232, LE), 5; on Ai, Pr.
Melanelixia fuliginosa (Fr. ex Duby) O. Blanco & al. — loc. 1, 2 (GU ER-16/2-088, herb. Urbanav.); on Ai, Jc.
Melanelixia glabra (Schaer.) O. Blanco & al. sensu lato — loc. 2, 4, 5; on Ai, Pt, Tc. The recently resurrected taxon M. epilosa (J. Steiner) A. Crespo & al. (Leavitt & al. 2016) and the species M. glabra are not morphologically distinguishable. Probably our Ingushetian specimens refer to M. epilosa, but it is not certain without molecular sequence data. We treat our specimens in a broad sense as M. glabra sensu lato.
Melanelixia glabratula (Lamy) Sandler & Arup — loc. 5; on Bp, Tc.
Melanelixia subargentifera (Nyl.) O. Blanco & al. — loc. 4, 7; on Ai, shale.
Melanelixia subaurifera (Nyl.) O. Blanco & al. — loc. 1, 3, 4, 5; on Ai, Sc, Tc, pine wood.
Melanohalea exasperata (De Not.) O. Blanco & al. — loc. 5; on Ai, Bp, Qp.
Melanohalea exasperatula (Nyl.) O. Blanco & al. — loc. 2, 5; on Ai, Bp, Pk, Pr, Qp, Tc.
†!Micarea nigella Coppins • — loc. 5 (IU ER-16/5-317, LE); on pine wood. Also known from NW Russia (Urbanavichus 2010).
*†!Micarea soralifera B. Guzow-Krzemińska & al. • — loc. 5 (IU ER-16/5-318, LE); on pine wood. A recently described species from the Czech Republic and Poland (Guzow-Krzemińska & al. 2016).
Montanelia tominii (Oxner) Divakar & al. — loc. 7; on shale.
#Muellerella lichenicola (Sommerf. ex Fr.) D. Hawksw. — loc. 3 (GU ER-16/3-176, herb. Urbanav.), 6 (in sample GU ER-16/6-370, LE); on thallus of Lecidella stigmatea, and apothecia of Gyalolechia flavovirescens and Xanthocarpia crenulatella on calcareous rocks.
+Mycocalicium subtile (Pers.) Szatala — loc. 5; on wood.
Myriolecis crenulata (Hook.) Śliwa & al. — loc. 2, 6, 7; on calcareous sandstones, shale.
Myriolecis dispersa (Pers.) Śliwa & al. — loc. 3; on calcareous rocks.
Myriolecis persimilis (Th. Fr.) Śliwa & al. — loc. 2, 5; on Ai, Tc.
Myriolecis pruinosa (Chaub.) Śliwa & al. — loc. 3; on calcareous rocks.
Myriolecis semipallida (H. Magn.) Śliwa & al. — loc. 2, 3, 6, 7; on calcareous sandstones, shale.
Neocatapyrenium rhizinosum (Müll. Arg.) Breuss — loc. 3 (GU ER-16/3-177, herb. Urbanav.); on calcareous soil.
Neofuscelia pulla (Ach.) Essl. — loc. 7; on shale.
Nephroma parile (Ach.) Ach. — loc. 5; on Sc, Tc.
#Nesolechia oxyspora (Tul.) A. Massal. — loc. 7 (GU ER-16/7-497, herb. Urbanav.); on thallus of Xanthoparmelia stenophylla on shale.
Ochrolechia arborea (Kreyer) Almb. — loc. 1, 3, 5; on Jc, Pk.
Opegrapha vulgata (Ach.) Ach. — loc. 5 (IU ER-16/5-322, LE); on Tc.
Parmelia barrenoae Divakar & al. — loc. 5 (GU ER-16/5-323, herb. Urbanav.); on Tc.
Parmelia sulcata Taylor — loc. 1, 2, 3, 4, 5, 6, 7; on Ai, Bp, Pk, Pr, Py, Sa, Sc, Tc.
Parmelina pastillifera (Harm.) Hale — loc. 5 (IU ER-16/5-325, LE); on Bp, Tc.
Parmelina tiliacea (Hoffm.) Hale — loc. 1, 5 (IU ER-16/5-326, LE); on Bp, Pk, Pr, Tc.
Parmotrema perlatum (Huds.) M. Choisy • — loc. 2, 4 (IU ER-16/4-239, LE); on Ai. (TLC — stictic acid).
Parmotrema stuppeum (Taylor) Hale • — loc. 5 (IU ER-16/5-327, LE); on Tc. (TLC — salazinic acid).
Peltigera canina (L.) Willd. — loc. 5; on soil.
Peltigera didactyla (With.) J. R. Laundon — loc. 7; on mossy rocks.
Peltigera elisabethae Gyeln. — loc. 5; on mossy log.
Peltigera lepidophora (Nyl. ex Vain.) Bitter — loc. 7; on soil above shale.
Peltigera praetextata (Flörke ex Sommerf.) Zopf — loc. 5; on Pt, Tc, log.
Peltigera rufescens (Weiss) Humb. — loc. 3; on soil.
Peltula bolanderi (Tuck.) Wetmore — loc. 3, 7 (GU ER-16/7-495, herb. Urbanav.); on calcareous sandstones and shale.
Peltula euploca (Ach.) Poelt — loc. 6, 7 (GU ER-16/7-496, herb. Urbanav.); on shale.
+Peridiothelia fuliguncta (Norman) D. Hawksw. — loc. 5 (GU ER-16/5-331, herb. Urbanav.); on Tc.
Phaeophyscia cernohorskyi (Nádv.) Essl. — loc. 6, 7 (GU ER-16/7-498, herb. Urbanav.); on plant debris above calcareous sandstones and shale.
Phaeophyscia ciliata (Hoffm.) Moberg — loc. 2, 5; on Ai, Sn.
Phaeophyscia endophoenicea (Harm.) Moberg — loc. 2, 5; on Ai, Sn.
Phaeophyscia hirsuta (Mereschk.) Essl. — loc. 2, 5; on Ai, Jc, Sc.
Phaeophyscia insignis (Mereschk.) Moberg — loc. 7 (GU ER-16/7-499herb. Urbanav.); on shale.
Phaeophyscia kairamoi (Vain.) Moberg — loc. 2, 4, 5; on Ai, Pt, Sc, Sn.
Phaeophyscia nigricans (Flörke) Moberg — loc. 2, 7; on Ai, branches of shrubs.
Phaeophyscia orbicularis (Neck.) Moberg — loc. 2, 5; on Ai.
Phaeophyscia pusilloides (Zahlbr.) Essl. — loc. 2, 4, 5; on Ai, Sc, Tc.
Phaeophyscia sciastra (Ach.) Moberg — loc. 1, 2, 3, 5, 6, 7; on calcareous rocks, sandstones, shale.
Phlyctis argena (Ach.) Flot. — loc. 1; on Jc.
Physcia adscendens (Fr.) H. Olivier — loc. 1, 2, 3, 4, 5, 6; on all plant and rock substrates.
Physcia aipolia (Ehrh. ex Humb.) Fürnr. — loc. 2 (in sample GU ER-16/2-050, herb. Urbanav.), 4, 5; on Ai, Pr, Pt.
Physcia caesia (Hoffm.) Fürnr. — loc. 2, 3, 5, 6, 7; on calcareous rocks, sandstones.
Physcia dimidiata (Arnold) Nyl. — loc. 6, 7; on shale, sandstones.
Physcia dubia (Hoffm.) Lettau — loc. 7; on shale.
Physcia stellaris (L.) Nyl. — loc. 2, 5, 6; on Ai, Bp, Jc, Qp.
Physcia subalbinea Nyl. — loc. 7; on shale.
Physcia tenella (Scop.) DC. — loc. 4, 5; on Ai, Pk.
Physcia tribacia (Ach.) Nyl. — loc. 2 (GU ER-16/2-106, herb. Urbanav.); on Ai.
Physcia vitii Nádv. — loc. 1, 2, 4, 5; on Ai, Jc, Pr, Sn.
Physciella chloantha (Ach.) Essl. — loc. 2, 4, 5, 6, 7; on Ai, Bp, Sn, and all rock substrates.
Physconia distorta (With.) J. R. Laundon — loc. 2, 4, 5, 6; on Ai, Pr.
Physconia grisea (Lam.) Poelt — loc. 7; on plant debris, shale.
Physconia muscigena (Ach.) Poelt — loc. 7; on soil, plant debris above shale.
Physconia petraea (Poelt) Vězda & Poelt — loc. 7; on rocks.
Physconia rossica G. Urban. — loc. 3, 7 (GU ER-16/7-510, herb. Urbanav.); on plant debris above shale.
Placidium rufescens (Ach.) A. Massal. — loc. 6 (GU ER-16/6-426, herb. Urbanav.); on calcareous rocks.
Placocarpus schaereri (Fr.) Breuss — loc. 3, 6; on calcareous rocks.
Placynthiella dasaea (Stirt.) Tønsberg — loc. 1, 3; on wood of pine log.
Placynthium nigrum (Huds.) Gray — loc. 3, 6; on calcareous rocks.
Placynthium posterulum (Nyl.) Hens sen — loc. 6 (GU ER-16/6-429, herb. Urbanav.); on calcareous rocks.
†#Polycoccum clauzadei Nav.-Ros. & Cl. Roux — loc. 7 (GU ER-16/7-511, herb. Urbanav.); on thallus of Rusavskia sp. on rocks. Previously known only from the Republic of Buryatia, S Siberia (Urbanavichene & Urbanavichus 2009).
Protoparmeliopsis muralis (Schreb.) M. Choisy — loc. 1, 2, 3, 5, 6, 7; on all rock substrates.
Pseudevernia furfuracea (L.) Zopf — loc. 1, 3, 5; on Jc, Pk.
Pseudoschismatomma rufescens (Pers.) Ertz & Tehler — loc. 4 (IU ER-16/4-237, LE); on Ai.
Psora decipiens (Hedw.) Hoffm. — loc. 3; on calcareous soil.
Punctelia borreri (Sm.) Krog — loc. 4, 7; on Ai, shale.
Punctelia jeckeri (Roum.) Kalb — loc. 1, 2, 3, 4, 5; on Ai, Jc, Pk, Pr, shale.
Punctelia subrudecta (Nyl.) Krog — loc. 5, 7; on Tc, shale.
Pycnora praestabilis (Nyl.) Hafellner — loc. 3; on Pk.
Pyrenodesmia chalybaea (Fr.) A. Massal. — loc. 6; on calcareous rocks.
Pyrenodesmia variabilis (Pers.) A. Massal. — loc. 3, 6; on calcareous rocks.
Ramalina asahinana Zahlbr. — loc. 2, 4, 5; on Ai, Pt, Pr.
Ramalina farinacea (L.) Ach. — loc. 2, 4; on Ai, Sc.
Ramalina pollinaria (Westr.) Ach. — loc. 2, 4, 5; on Ai, Sc, Tc.
Ramalina sinensis Jatta — loc. 2, 5; on Ai, Pr.
Ramalina subgeniculata Nyl. — loc. 4; on Ai.
†!Ramonia chrysophaea (Pers.) Vězda — loc. 5 (IU ER-16/5-353, LE); on Sn. Previously reported from the S Ural mountains (Urbanavichene & al. 2013). A very rare or overlooked species, not reported outside of Europe.
Rhizocarpon disporum (Nägeli ex Hepp) Müll. Arg. — loc. 7 (GU ER-16/7-514, herb. Urbanav.); on shale.
Rhizoplaca chrysoleuca (Sm.) Zopf — loc. 6, 7; on shale, small granite stones.
Rinodina bischoffii (Hepp) A. Massal. — loc. 6 (GU ER-16/6-431, herb. Urbanav.); on calcareous rocks.
Rinodina exigua (Ach.) Gray — loc. 2 (GU ER-16/2-115, herb. Urbanav.); on Ai.
Rinodina oleae Bagl. — loc. 2 (GU ER-16/2-116, herb. Urbanav.); on Ai.
Rinodina septentrionalis Malme — loc. 1 (IU ER-16/1-043, LE); on Pk.
Rinodina sophodes (Ach.) A. Massal. — loc. 2 (GU ER-16/2-117, herb. Urbanav.); on Ai.
*†Rinodina straussii J. Steiner — loc. 7 (GU ER-16/7-516, LE); on shale. Also known from S Asia, the European Alps and North America (Sheard 2010).
*†!Rinodina tunicata H. Mayrhofer & Poelt — loc. 6 (GU ER-16/6-517, herb. Urbanav.); on calcareous rocks. A mainly Mediterranean species occurring on calcareous rocks at middle altitudes in the mountains of S Europe and N Africa (Giralt & Llimona 1997).
*†Rinodina zwackhiana (Kremp.) Körb. — loc. 7 (GU ER-16/7-518, LE); on shale. Also known from S Europe, Asia (China) and North America (Sheard 2010).
Rusavskia papillifera (Vain.) S. Y. Kondr. & Kärnefelt — loc. 2, 3, 6, 7; on calcareous rocks.
Sarcogyne privigna (Ach.) A. Massal. — loc. 7 (GU ER-16/7-519, herb. Urbanav.); on shale.
Sarcogyne regularis Körb. — loc. 3 (GU ER-16/3-193, herb. Urbanav.); on calcareous rocks.
*†!+Schizoxylon alboatrum Rehm — loc. 5 (GU ER-16/5-354, LE); on Tc. A relatively rare species in only a few European countries and North America (Sherwood 1977).
Spilonema revertens Nyl. — loc. 7 (GU ER-16/7-520, herb. Urbanav.); on shale.
Staurothele frustulenta Vain. — loc. 7 (GU ER-16/7-521, herb. Urbanav.); on shale.
+Stenocybe pullatula (Ach.) Stein — loc. 2, 4, 5; on Ai.
Synalissa ramulosa (Hoffm. ex Bernh.) Fr. — loc. 3; on calcareous rocks.
Thallinocarpon nigritellum (Lettau) P. M. Jørg. — loc. 3, 6; on calcareous rocks and sandstones.
‡Thermutis velutina (Ach.) Flot. — loc. 6 (GU ER-16/6-433, herb. Urbanav.); on thallus of Toninia cinereovirens on calcareous rocks. Recently reported from Armenia in Transcaucasia (Gasparyan & al. 2015).
Thyrea confusa Henssen — loc. 6; on calcareous rocks.
Toninia candida (Weber) Th. Fr. — loc. 3; on calcareous rocks.
Toninia cinereovirens (Schaer.) A. Massal. — loc. 3, 5, 6; on calcareous rocks.
Toninia toniniana (A. Massal.) Zahlbr. — loc. 3; on calcareous rocks.
Trapeliopsis flexuosa (Fr.) Coppins & P. James — loc. 1, 3; on Pk, log.
Umbilicaria cylindrica (L.) Delise ex Duby — loc. 7; on shale.
Umbilicaria hirsuta (Sw. ex Westr.) Hoffm. — loc. 7; on shale.
Usnea cavernosa Tuck. — loc. 5; on Pk.
Usnea dasopoga (Ach.) Nyl. — loc. 5; on Pk.
Usnea florida (L.) F. H. Wigg. — loc. 1 (IU ER-16/1-045, LE), 2 (GU ER-16/2-119, herb. Urbanav.), 3, 4, 5, 6, 7; on Jc, Pk, Pr, Py, Qp, Tc.
Usnea hirta (L.) F. H. Wigg. — loc. 1, 3, 5; on Jc, Pk, snag.
Usnea lapponica Vain. — loc. 5 (GU ER-16/5-361, herb. Urbanav.); on Tc.
Usnea wasmuthii Räsänen — loc. 5 (IU ER-16/5-362, LE); on Tc.
Variospora aurantia (Pers.) Arup & al. — loc. 3, 6; on calcareous rocks.
Variospora velana (A. Massal.) Arup & al. — loc. 3; on calcareous rocks.
†!Verrucaria elevata (Nyl.) Zschacke — loc. 6 (GU ER-16/6-437, herb. Urbanav.); on calcareous rocks. Recently reported from the Leningrad region (Pykälä & al. 2012).
Verrucaria furfuracea (B. de Lesd.) Breuss — loc. 6; on calcareous rocks.
*‡Verrucaria fuscoatroides Servít — loc. 3 (GU ER-16/3-202, herb. Urbanav.); on calcareous rocks. Recently reported from Abkhazia in Transcaucasia (Urbanavichus & Urbanavichene 2012). An apparently rather widespread but still poorly known species described from Germany; also reported from a few other European countries and North America (Breuss 2007; Breuss & Berger 2010). In her excellent revision of Verrucaria in Poland, Krzewicka (2012) suggested that V. fuscoatroides may be a synonym of V. nigroumbrina (A. Massal.) Servít.
Verrucaria macrostoma Dufour ex DC. — loc. 2 (GU ER-16/2-120, herb. Urbanav.); on calcareous sandstones.
Verrucaria nigrescens Pers. — loc. 7 (GU ER-16/7-525, herb. Urbanav.); on shale.
Verrucaria sphaerospora Anzi — loc. 3, 6 (GU ER-16/6-439, herb. Urbanav.); on calcareous rocks.
Verrucaria tristis (A. Massal.) Kremp. — loc. 3 (GU ER-16/3-204, herb. Urbanav.); on calcareous rocks.
Xanthocarpia crenulatella (Nyl.) Frödén & al. — loc. 3, 6; on calcareous rocks.
Xanthoparmelia camtschadalis (Ach.) Hale — loc. 2, 3, 6; on soil.
Xanthoparmelia conspersa (Ehrh. ex Ach.) Hale — loc. 7; on shale.
Xanthoparmelia stenophylla (Ach.) Ahti & D. Hawksw. — loc. 2, 7; on shale.
Xanthomendoza fallax Søchting & al. — loc. 3, 7; on calcareous rocks, shale.
Xanthomendoza fulva (Hoffm.) Søchting & al. — loc. 6; on calcareous rocks.
Xanthomendoza ulophyllodes (Räsänen) Søchting & al. — loc. 2; on Ai.
Xanthoria aureola (Ach.) Erichsen — loc. 3, 7; on calcareous rocks, shale, mossy rocks.
Xanthoria calcicola Oxner — loc. 2, 6, 7; on sandstones, shale.
Xanthoria parietina (L.) Th. Fr. — loc. 1, 2, 3, 4, 5, 6, 7; on Ai, Cb, Jc, Pk, Pr, Py, Qp, Sc, Tc.
Xylographa parallela (Ach.) Fr. — loc. 3, 5; on pine wood. #Zwackhiomyces coepulonus (Norman) Grube & R. Sant. — loc. 7 (GU ER-16/7-532, herb. Urbanav.); on thallus of Calogaya biatorina on shale.
Conclusions
The inventory of lichenized and lichenicolous fungi taxa in Erzi Nature Reserve is essential for protecting the Caucasian lichen flora, and for future floristic, taxonomic, systematic and ecological studies, as well as for the monitoring of future changes in the composition of the flora. We currently recognize 286 species as occurring in the Erzi Nature Reserve (266 species of lichenized fungi, 13 of lichenicolous fungi and seven of non-lichenized saprophytic fungi), representing 115 genera. Twentythree species are recorded for the first time for North Caucasus, of which 16 are new for the Caucasus and ten are new for Russia (Arthonia protoparmeliopseos, Aspicilia viridescens, Buellia abstracta, Lecidella viridans, Micarea soralifera, Rinodina straussii, R. tunicata, R. zwackhiana, Schizoxylon alboatrum and Verrucaria fuscoatroides). Of the 16 species new for the Caucasus, seven are recorded for the first time for Asia (Buellia abstracta, Micarea nigella, M. soralifera, Ramonia chrysophaea, Rinodina tunicata, Schizoxylon alboatrum and Verrucaria elevata). In addition, two of the 42 taxa listed in the Red Data Book of Russia (Trutnev & al. 2008) are recorded, namely Leptogium burnetiae (Near Threatened category) and Usnea florida (Vulnerable category).
Acknowledgements
This work was supported by a grant of the Russian Foundation for Basic Research, No. 15-29-02396 “North Caucasian Lichen Flora: taxonomic structure, diversity, specificity, systematics of individual taxa and contribution to the diversity of the lichen flora of Russia”. We are grateful to Liza Khaykharoeva and staff of the Erzi Nature Reserve for supporting our research and to Prof. Mark Seaward who kindly corrected the English. We also thank Dr. Harrie J. M. Sipman (Botanic Garden and Botanical Museum Berlin) and an anonymous reviewer for their comments on an earlier version of this paper. Special thanks are also due to Igor Lavrinenko (Komarov Botanical Institute, St. Petersburg) for technical assistance with the map (Fig. 1).
