Xanthoria (sensu Arup et al. 2013 )

Vondrák, Jan, Frolov, Ivan, Davydov, Evgeny A., Yakovchenko, Lidia, Malíček, Jiří, Svoboda, Stanislav & Kubásek, Jiří, 2019, The lichen family Teloschistaceae in the Altai-Sayan region (Central Asia), Phytotaxa 396 (1), pp. 448-450 : 448-450

Xanthoria (sensu Arup et al. 2013)

One species recorded in the region. All species have large foliose thalli; vegetative diaspores are absent; apothecia are present. Literature: Arup et al. (2013), Lindblom (1997; including other present-day Xanthorioid genera).

Xanthoria parietina : 2 localities at altitudes 200 and 210 m. Both localities humid non-alpine and both at city of Barnaul. Substrate: bark of Populus . It is marginally distributed in the region and possibly restricted to lowlands north of Altai .

‘ Caloplaca ’ anularis group ( Fig. 15 )

Four species recorded in the region. Thallus is crustose, with marginal lobes; vegetative diaspores are absent; apothecia are sometimes present. Thallus typically very thick, with thick medulla (often several hundred μm) and thick cortex (tens of μm); algal and fungal stacks (sensu Vondrák & Kubásek 2013) sometimes present. Thallus and apothecia are with non-chlorinated anthraquinones. The group forms a genus-level clade in Xanthorioideae , perhaps with sister relationship to Amundsenia ( Fig. 2 ). It is broadly distributed and diverse in arid and alpine regions of Asia and probably absent from Arctic regions. Literature: Clauzade & Poelt (1972), Lee et al. (2018), Poelt & Hafellner (1980), Vondrák & Mayrhofer (2013).

‘Caloplaca’ anularis : 6 localities at altitudes 1500–2900 m, mostly in arid alpine habitats. Substrate: limestone rocks.

‘Caloplaca’ cf. bohlinii , Fig. 7A : 3 localities at altitudes 900–1750 m, in arid alpine and non-alpine habitats. Substrate: limestone rocks. It is distinguished from C. anularis by smaller, compact, orange thalli with shorter and broader marginal lobes. This species never forms ring-shaped fragments of thalli (sensu Vondrák & Mayrhofer 2013). It is also similar to C. scrobiculata , but thallus is less robust with fewer white pruinose spots, algal and fungal stacks are less distinct and marginal lobes less reduced. The little known C. bohlinii was described by Magnusson (1940) from Central Asia. Its type specimen (L 2622 in S!) has intermediate phenotype between C. anularis and C. cf. bohlinii and the identity of our material is thus uncertain.

‘Caloplaca’ scrobiculata : single specimen recorded at altitude 3000 m in arid alpine locality in Altai (Vondrák 9933). Substrate : limestone boulder .

‘Caloplaca’ zeorina (= Caloplaca anularis f. ignea ): single record at altitude 2900 m in arid alpine locality in Altai (specimen Vondrák 9928). This taxon is phenotypically distinct from C. anularis and both taxa are also separated in the ITS tree ( Fig. 15 ). Its status at species level therefore seems to be correct .

‘ Caloplaca ’ ahtii

Our single ITS sequence (MG954161) is placed in subfamily Xanthorioideae , but stands outside any known clade ( Fig. 2 ); it may belong in a genus of its own. The single species forms thin grey crusts with tiny crater-like soralia containing the pigment Sedifolia-grey. This pigment is also frequently present in outer apothecial margin, giving a grey tinge. Apothecia are yellow to orange, containing anthraquinones of chemosyndrome A. It is a typical boreal species common in taiga forests, usually epiphytic. Literature: Søchting (1994).

Caloplaca ahtii : 13 localities at altitudes 350–1490 m, mostly in humid, non-alpine habitats (forests). Substrate: bark of Populus and Salix , rarely on other trees, also on wood and exceptionally on dust-impregnated rock (Vondrák 10404).

‘ Caloplaca ’ exsecuta group ( Figs 16A, B )

Three species recorded in the region. All species are crustose, with thin or indistinct thallus. Vegetative diaspores are present in Caloplaca sorocarpa which is usually without apothecia. Apothecia are usually numerous in other species, containing anthraquinones in disc or in both disc and margin. Non-chlorinated anthraquinones predominate: parietin (major), emodin (minor) and traces of emodinal, emodic acid, parietinic acid and fallacinal (analysed specimens of C. exsecuta : Vondrák 1105, Tønsberg 46194). 7-Cl-emodin was not seen in UV, but was identified by mass spectrometry in Tønsberg 46194. Thallus and usually apothecial margin has Cinereorufa-green (sometimes only detected in prothallus); anthraquinones are absent from thallus. The group includes boreal-montane to arctic-alpine species. The three species listed below form a monophyletic group in ITS tree together with Caloplaca tornoenesis and C. nivalis (newly sequenced specimens in Table 1). The group is defined here and no relevant literature is available.

‘Caloplaca’ borealis : 2 localities at altitudes 1120–1530 m in humid non-alpine and subalpine habitats. Substrate: bark of Abies sibirica .

‘Caloplaca’exsecuta: 2 localities at altitudes 1260–2180 m in humid alpine habitats. Substrate: base-rich siliceous rocks. Colour of disc variable: yellow-orange in specimen Vondrák 11110 ( Fig. 16A ), but ferrugineous red in specimen Vondrák 11105 ( Fig. 16B ).

‘Caloplaca’sorocarpa: only recorded from a single locality at altitude 1580 m in humid subalpine habitat (specimen Vondrák 12695; Western Sayan). Substrate: twig of Grossularia shrub. Probably more frequent in the region.

‘ Caloplaca ’ conversa / conglomerata group

One species recorded in this group which is placed the subfamily Caloplacoideae and close to a group of species called “section Coccinodiscus ” sensu Poelt & Kalb (1985), see Fig. 2 . The group, as currently known to us, is morphologicaly characterised by presence of large amounts of Sedifolia-grey in all parts of the thallus and in apothecial margin. Anthraquinones are often confined to apothecial disc, but in some specimens, anthraquinones are replaced by Sedifolia-grey in discs. All specimens are epilithic. High diversity was observed in the Mediterranean basin (Vondrák, unpublished). Specimens of Rufoplaca with grey apothecial margin are sometimes very similar to some C. conversa morphotypes, but ascospores are broad and with thick septa in Caloplaca conversa / conglomerata group, but narrow with thin septa in Rufoplaca . The group involves at least two morphospecies, C. conversa (thin areolate crusts) and C. conglomerata (= C. peludella ) with thicker squamulose thallus, but more phylogenetic lineages are involved. Literature: Nimis (2016), Redchenko et al. (2012; ITS sequence of the type of C. conversa ).

‘Caloplaca’conversa sensu lato: 7 localities at altitudes 400–1230 m, mostly in arid non-alpine habitats. Substrate: base-rich siliceous or calcareous rocks (limestone, schist, gneiss), usually on xerothermic outcrops in forest-steppe zone.

‘ Caloplaca ’ haematites / aractina group

One species recorded in the region. Thallus is crustose, white-grey, without anthraquinones, epilithic, thinly areolate. Vegetative diaspores are absent. Apothecia are orange-red, with non-chlorinated anthraquinones. It is related to Pyrenodesmia , but forms a genus-level group with European and Mediterranean C. aractina and C. haematites (Frolov, in preparation). Literature: Magnusson (1940).

‘Caloplaca’ bicolor (= Caloplaca kansuensis ): 3 localities at altitudes 2050–2900 m in arid alpine habitats. Substrate: hard limestone and soft calcareous rock (schist). In other regions also known from siliceous rocks. No differences were found between types of Caloplaca bicolor (L 2616 in S) and C. kansuensis (L 2603 in S). The two names were given to two different taxa by Magnusson (1940), but we consider them synonymous.

‘ Caloplaca ’ raesaenenii

The species resembles some species of Athallia . It has indistinct thallus; vegetative diaspores are absent; apothecia are with non-chlorinated anthraquinones, small, initially immersed in substrate. It does not group with any known genus in ITS tree. Literature: Vondrák et al. (2012a).

‘Caloplaca’ raesaenenii : 12 localities at altitudes 250–2230 m, mostly in arid non-alpine habitats. Substrate: epiphytic, on bark and twigs of trees and shrubs, on plant debris and bryophytes (in steppes on calcareous substrate).

‘ Caloplaca ’ xerica group

Five species recorded in the region, but all are rare in the region. Thallus is crustose, white to dark grey, with Sedifolia-grey pigment; vegetative diaspores are present in C. soralifera (soredia, blastidia), C. teicholyta (blastidia) and C. xerica (blastidia, isidia); apothecia are present in all species, but rare in C. teicholyta ; chlorinated and non-chlorinated anthraquinones are present. The group is confined to inorganic substrates, usually on mineral-rich siliceous rocks. It is related to Pyrenodesmia but merits its own genus (Frolov, unpublished). Literature: Arup et al. (2013), Vondrák et al. (2012b).

‘Caloplaca’ atroflava : Only one specimen recorded at altitude 430 m in humid non-alpine habitat (Frolov 1519). Substrate: gneiss rock.

‘Caloplaca’percrocata: Only one specimen recorded at altitude 2300 m in humid alpine habitat (Davydov 12295). Substrate: rock.

‘Caloplaca’ soralifera : 4 localities at altitudes 410–480 m in humid and arid non-alpine habitats. Substrate: base-rich siliceous or calcareous rock.

‘Caloplaca’ teicholyta : 2 localities at altitudes 410–1900 m in arid alpine and non-alpine habitats. Substrate: limestone.

‘Caloplaca’xerica: Two specimens recorded in one locality at altitude 1480 m in humid non-alpine habitat (Frolov 175, Vondrák 10292). Substrate : gneiss rock, below overhang. This species is known to be xerophilous in Europe, but was not recorded from arid habitats in the Altai-Sayan region .

Figures

FIGURE 2. Classification within Teloschistaceae displayed on the single locus ITS tree (2372 sequences included, maximum likelihood method, branches with the bootstrap support ≥ 70% are thick). Genera and groups putatively on generic level are collapsed into single terminals. Groups occurring in southern Siberia are linked with red names on the right. Pale grey link: genera with 1–2 species; medium grey, 3–5 species; black, 6 and more species.

FIGURE 7. ‘Caloplaca’ anularis group (A) and Pachypeltis (B–E). A, ‘Caloplaca’ cf. bohlinii (Vondrák 18251) distinguished from ‘C.’ anularis by orange thallus with more frequent apothecia; B, Pachypeltis insularis (Vondrák 10340); C, Pachypeltis intrudens (Vondrák 18059); D, Pachypeltis cf. pachythallina (large squamules) on Calogaya (Vondrák 12649); E, Pachypeltis phoenicopta (Vondrák 18695) having deep orange pycnidia contrasting with pale yellow colour of thallus; F, detail of areoles of P. phoenicopta (Vondrák 18695) showing patches of white pruina that gives the typical pale yellow colour to the thallus.All bars, 1 mm.

FIGURE 15. Bayesian phylogenetic reconstruction of the ‘Caloplaca’ anularis group using the ITS DNA locus. Sequences from the Altai-Sayan region are in bold.

FIGURE 16. ‘Caloplaca’ exsecuta (A, B), Xanthocarpia (C) and an unknown ‘Caloplaca’ (D).A, ‘Caloplaca’ exsecuta (Vondrák 11110) with yellow-orange discs; B, ‘C.’ exsecuta (Vondrák 11105) with ferrugineous red discs; C, Xanthocarpia sp. (Vondrák 18686) resembling the Mediterranean X. marmorata; D, unknown ‘Caloplaca’ with unresolved ITS position within Xanthorioidae (Vondrák 18687).All bars, 1 mm.