New records of lichens from the Russian Far East. III. Lichens of coastal habitats

1 Altai State University, Lenina Pr. 61, Barnaul, 656049, Russian Federation 2 Federal Scientific Center of the East Asia Terrestrial Biodiversity FEB RAS, Stoletiya Vladivostoka Pr. 159, Vladivostok, 690022, Russian Federation 3 Institute of Natural Sciences and Mathematics, Department of Biology, Ural Federal University, Lenina Pr. 51, Ekaterinburg, 620000, Russian Federation 4 Russian Academy of Sciences, Ural Branch: Institute Botanic Garden, 8 Marta St. 202a, Ekaterinburg, 620144, Russian Federation 5 Botany Department, Finnish Museum of Natural History, University of Helsinki, P.O. Box 7, 00014, Finland 6 E-mail: eadavydov@yandex.ru; ORCID iD: https://orcid.org/0000-0002-2316-8506 7 E-mail: lidiyakovchenko@mail.ru; ORCID iD: https://orcid.org/0000-0002-4342-7771 8 E-mail: gairka@yandex.ru; ORCID iD: https://orcid.org/0000-0001-9029-2470 9 E-mail: alexander_paukov@mail.ru; ORCID iD: https://orcid.org/0000-0001-6689-7189 10 E-mail: ivfrolov@gmail.com; ORCID iD: https://orcid.org/0000-0003-4454-3229 11 E-mail: teuvo.ahti@helsinki.fi; ORCID iD: https://orcid.org/0000-0001-7549-0962


Introduction
Russian Far East is a vast territory, adjacent to the Pacific Ocean, which stretches from the Chukotka Peninsula in the North to the Primorye Territory in the South. The area of the region is more than 3 million square kilometers, which is approximately 18 % of the total territory of Russia. It is geographically divided into three parts: arctic, northern and southern regions. The latter includes five administrative provinces: Amur Region, Jewish Autonomous Region, Primorye Territory, Sakhalin Region and the southern part of Khabarovsk Territory. The region has a mainly mountainous ground profile. Inland mountains and highlands have smooth slopes without pronounced ridges or canyons. In the territories situated closer to the Pacific Ocean high and pointed rocks and steep cliffs are common. In some regions of the Russian Far East the border of the subalpine zone descends closely to the seashore (Urusov, 2018).
The descriptions of the region, its climate, and the history of lichen studies as well as some new and interesting records of lichens for the Far East of Russia are published in our previous papers (Yakovchenko et al., 2019(Yakovchenko et al., , 2020Davydov et al., 2021). Here we report on the findings of several species new to the Russian Far East and further detailed information on rarely reported species along with their identification characters, images, and distribution.

Materials and methods
Morphological observations were made using a dissecting microscope. Cross-sections of apothecia and thalli were made by hand with a razor blade and observed after mounting in water using a microscope. Measurements of spores and conidia are presented as follows: (smallest value recorded) (X -SE) -X -(X + SE) (largest value recorded), where X is the (arithmetic) sample mean, and SE is the sample error of mean. The measurements were made with the precision of 0.5 μm. Polarized light (pol) was used for locating crystals in the sections.
Lichen substances of some species were studied by spot-tests using potassium hydroxide solution (K), sodium hypochlorite solution (C), 1,4-p-phenylendiamine (Pd), and iodine (I), and by a thin-layer chromatography (TLC) in solvent systems A, B and (Orange et al., 2001).
Buellia subdisciformis is a cosmopolitan coastal species in the xeric supralittoral (Coppins et al., 2009). In Russia it was only known from the Caucasus (Urbanavichus, 2010). It is characterized by the areolate, whitish thallus (K+ red, norstictic acid, I-) with the black hypothallus and sessile, black apothecia with a greyish proper margin, brown epihymenium and a dark red-brown hypothecium. Buellia aethalea differs by its immersed apothecia and aeruginose epihymenium; B. maritima is distinguished by the densely pruinose thallus with a chalky consistency and a paler (whitish to grey) hypothallus; Buellia taishanensis differs by its poorly-developed, darker (grey or grey-brown) thallus; B. yoshimurae -by its thallus, areolate in the center and rimose at the periphery, immersed apothecia and a medulla lacking calcium oxalate crystals.
This is a widespread saxicolous lichen in Europe and Asia. It was reported from different regions of Russia: Caucasus, South Siberia, Transbaikal Region, and the Urals (Vondrák et al., 2016). Calogaya arnoldii is a placodioid lichen with orange thallus and apothecia, without vegetative propagules. Morphologically it is very close to C. saxicola (Hoffm.) Vondrák. The differences between species are subtle and mostly concern shape and size of ascospores (Vondrák et al., 2016). Our finding is confirmed by the ITS sequence (not shown).   This is a common lichen in the Sakhalin Island, where it grows on a variety of rocks and on concrete in many maritime and inland localities. The species has a worldwide distribution and is known from Asia, Europe, Hawaii, North America, and Australia (Arup, 2006;Vondrák et al., 2016). In the Asian part of Russia the species was recorded in the Republic of Altai and in the Trans-Baikal Territory (Vondrák et al., 2016(Vondrák et al., , 2019. Flavoplaca flavocitrina is the species with a wide ecological range, which occurs on rocks and concrete, but it is also common on bark and wood. In its typical form, the species is characterized by the yellow apothecia and the yellow, areolate thallus with marginal soralia contrasting with the rest of the thallus. Lecanora swartzii is a cosmopolitan species growing on siliceous rocks in uplands (Edwards et al., 2009). In Russia it was reported from Caucasus, South Siberia as well as the northern parts of the Urals and European Russia (Urbanavichus, 2010). In the Russian Far East it grows not directly on the seashore but on gravel barrens near the coast. The species has a warty-areolate to bullate or stalked areolate, yellowish-white thallus (C+ yellow orange, K+ yellow, UV+ pale orange) and a white hypothallus. Apothecia lecanorine, plane to convex, markedly constricted at the base to stalked, with densely pruinose, yellowish-grey disc (C+ yellow orange). From the similar species, Lecanora rupi-30 Davydov E. A. et al. New records of lichens from the Russian Far East cola (L.) Zahlbr., it is easily distinguished by its C+ yellow-orange thallus and the absence of phenocortex on the thalline margin. Lecanora swartzii shares the eucorticate amphithecium with genetically and ecologically similar species, Lecanora bicinctoi-dea Blaha et Grube, but differs from the latter by its С-negative thallus, a white hypothallus and soon constricted at the base to distinctly stalked apothecia (Blaha, Grube, 2007). Chemistry: atranorin, thuringione and trace of arthothelin (TLC AP 291-11).
The species is distributed in Europe, Asia, Macaronesia, and North America (Fletcher et al., 2009;Esslinger, 2018) and occurs on siliceous or slightly basic rocks, often on walls and memorials, rarely on wood, worked timber or bark. In Russia it is known from South Siberia as well as the arctic and northern part of European Russia (Urbanavichus, 2010). It is characterized by a grayish with a yellowish tint, rimose-areolate to granular thallus; K+ yellow, C+ orange with discrete to confluent soralia, concolorous with the thallus composed of fine (ca. 20 µm) soredia. Apothecia black, lecideine, epihymenium green, hypothecium from dullyellow to brown. A similar species with vegetative propagules, Lecidella meiococca (Nyl.) Leuckert et Hertel, differs by its blastidiate thallus.

New records for the mainland of Far East
Lecidella asema (Nyl.) Knoph et Hertel (Fig. 6 (Himelbrant et al., 2019). Here we are reporting the second locality of the species in Primorye Territory, which is the first record of the species from the continental part of the Far East. Worldwide, the species is known in Europe, Asia, Macaronesia, Africa, and North America (Fletcher et al., 2009;Moon, 2013;Esslinger, 2018;Ohmura, Kashiwadani, 2018), where it usually grows on supralittoral rocks and rocks in coastal mountains, while it is infrequent in the inland territories. The species is characterized by its coarsely rimose, pale fawn-ochre thallus, C+ yellow-orange, K-, P-, and lecideine apothecia with a persistent proper margin, which is often paler than the disc. Exciple gray-green to gray-blue at the outer part and red-brown inside. The inner part of the exciple contains crystals (pol+), hypothecium deep red-brown, epihymenium blue-green. From a similar species, Lecidella carpathica Körb., it is easily distinguished by its thallus reacting yelloworange with C. Rinodina gennarii is characterized by its small (< 17.5 μm long), Dirinaria-type spores, saxicolous habitats, and a coastal distribution. It has a worldwide distribution in temperate regions of the Northern and Southern Hemispheres growing on maritime rocks (Trinkaus et al., 1999;Sheard et al., 2017). It is hard to mention the true distribution of the species in Russia because Kotlov (2008) regarded specimens growing on bark and on rock as the same species apparently describing the distribution of a mixture of two taxa, R. gennarii and R. oleae Bagl. Then, in the Checklist of lichens of Russia R. gennarii was Davydov E. A. et al. New records of lichens from the Russian Far East referred as a synonym of R. oleae (Urbanavichus, 2010), although R. gennarii is a primarily saxicolous species on maritime rocks occasionally growing on wood pilings at the seashores, whereas R. oleae is a corticolous species and is widely distributed in Eurasia and North America (Sheard, 2010;Sheard et al., 2017). In the East Asian region the confirmed records of R. gennari have been made in Japan (Sheard et al., 2017), Kunashir Island (Bredkina et al., 1992Tchabanenko, 2002), and Sakhalin Island (Konoreva et al., 2018). Here we report it for the first time to the mainland part of the Russian Far East from Primorye Territory.

The records of noteworthy species
Caloplaca atroflava (Turner) Mong. s. l. (Fig. 8 This is a common lichen in the Sakhalin Island. It grows on siliceous outcrops along the seashore and on siliceous boulders in rivers. It is known from Europe, Asia, and North America (Fletcher, Laundon, 2009). In Russia, the species is known from the north of the European part and in South Siberia and Primorye Territory (Urbanavichus, 2010;Skirina, 2016;Vondrák et al., 2019). The species is a one of the most widespread taxa of the so-called C. xerica group (sensu Vondrák et al., 2012). Caloplaca atroflava s. l. has a thin, areolate, grey thallus and zeorine apothecia with the thalline exciple of the same colour as the thallus and an orange-red disk. The proper exciple and disk in Caloplaca atroflava s. l. have a negative reaction with C (contains non-chlorinated anthraquinones). Besides a normal form, lacking vegetative propagules, Caloplaca atroflava s. l. has a form with poorly or well-developed blastidia (Vondrák, unpublished). In the Sakhalin Island we found both morphotypes, with and without blastidia.

A B
33 This lichen containing atranorin and fumarprotocetraric acid was earlier described as Cladonia kurokawae Ahti et Stenroos (Ahti et al., 1996). However, in a multilocus molecular phylogenetic study Pino-Bodas et al. (2013) synonymized this species with C. subconistea containing atranorin and psoromic acid. Further these taxa appear to be morphologically very close. Both chemotypes are widespread in China (also in Taiwan), Japan and Korea (ibid.). Cladonia subconistea is an East Asian endemic reported from the Russian Far East by Tchabanenko (2002) on the basis of the material (with psoromic acid) from the herbaria H and TU identified by T. Ahti. Cladonia subconistea is not the only species in the genus Cladonia, which has taxonomically insignificant strains with psoromic acid. These strains are also known in C. stellaris (Opiz) Pouzar et Vězda, C. symphycarpa (Ach.) Fr., C. furcata (Huds.) Schrad., and C. pocillum (Ach.) Grognot (Ahti, Stenroos, 2013). Cladonia kurokawae and Cladonia subconistea may still be separate species and require further examination.

Leptotrema lithophila Oxner
Chemistry: no substances (TLC ED1501-17, TLC ED1501-19). This rare species was described by Oxner (1960) as an endemic to Sikhote-Alin Range based on a single specimen collected in Primorye Territory. Later it was found in Kamchatka Peninsula (Mikulin, 1990). The species was listed in the checklists of lichens of the southern Far East (Tchabanenko, 2002) and Russia (Urbanavichus, 2010) as well as reported in a locality ca. 50 km to the south-west from the locus classicus (Cherdantseva et al., 2013). All reported collections of the species were made in upland localities at least 25 km from the seashore. Here we are reporting Leptotrema lithophila from the coastal habitats for the first time. Leptotrema litophila is characterized by having a crustose, grey, continuous and irregular thallus that would soon partly flake away from the rocky substrate, with numerous inclusions of calcium oxalate crystals and sunken, pruinose myriotremoid-type to lepadinoidtype apothecia. The lower surface has numerous root-like heavy-branched outgrowths. Morphologically this species resembles some representatives of Diploschistes that is not a close relative to Leptotrema.
Umbilicaria vellea (L.) Hoffm. This is a locally rare but a widespread species in the Russian Far East. In Kamchatka it grows on shaded and humid vertical slopes of old volcanic rocks as well as on the seashore cliffs (Davydov et al., 2011). Umbilicaria vellea has been reported to the Kuril Islands as a whole in "The Lichen Flora of Russia" (Davydov, 2017) on the basis of the following single collection, here documented for the first time in detail.