The Lichenologist 46(3): 413–439 (2014) 6 British Lichen Society, 2014 doi:10.1017/S0024282913000911
Unveiling a surprising diversity in the lichen genus Micarea (Pilocarpaceae)inRe´union (Mascarenes archipelago, Indian Ocean)
A. Maarten BRAND, Pieter P. G. VAN DEN BOOM and Emmanue¨lSE´ RUSIAUX
Abstract: Detailed anatomical and chemical studies conducted on recent collections made in almost all suitable habitats on Re´union, a small remote tropical island in the Indian Ocean, yielded a surpris- ing diversity in the widespread lichen genus Micarea (Pilocarpaceae, Lecanorales). Twenty-one species are recognized, including 13 described here as new to science. They are: Micarea alectorialica, M. bebourensis, M. borbonica, M. boryana, M. cilaoensis, M. hyalinoxanthonica, M. isidiosa, M. melano- prasina, M. pseudocoppinsii, M. pseudolignaria, M. sublithinella, M. takamakae and M. tenuispora. Notes on local ecology and important biogeographical features are also given and a key to the species is pro- vided. Isidiiform areolae are reported for the first time in the genus (M. isidiosa and M. tenuispora), as well as the production of protolichesterinic and confluentic acids (M. sublithinella and M. takamakae, respectively). Two groups within the genus are species-rich on the island: the M. peliocarpa group with possibly 5 species, including 3 new to science, and the M. prasina group with 4 species, includ- ing 2 new to science. Micarea levicula is reported here for the first time since its description, and the status of the material that can be referred to M. micrococca s. lat. needs further study. Key words: lichen diversity, lichenized Ascomycota, new species, Southern Hemisphere, taxonomy, tropical habitats Accepted for publication 16 September 2013
Introduction widespread and speciose genus (Micarea Fr., Pilocarpaceae, Lecanorales) to describe each The species is the fundamental unit of any species and formulate a sound taxonomic study on the biodiversity and ecosystems of treatment of all species found in this limited, our planet, and yet, many groups of organ- but very diverse, territory in the tropics. By isms have only a small fraction of their species showing that the genus Micarea is species- formally described (Gaston 2000). Fungi as rich in Re´union, with many undescribed a major phylum present in almost all biomes species, we provide further data for the represent a remarkable example: the number better assessment of the total number of un- of species is expected to be much higher than described lichen taxa (Lu¨cking et al. 2009; a million, while only 100 000 are currently Bass & Richards 2011). described (Blackwell 2011; Hawksworth Although there are many species of Micarea 2012). We performed detailed anatomical in the Southern Hemisphere, the few mono- and chemical studies of lichen material re- graphic treatments of the genus, however mas- cently collected in Re´union (a small and terful, have addressed only European species remote island in the Indian Ocean) in a (Coppins 1983, 2009; Czarnota 2007). Mo- lecular inferences resolved the genus into A. M. Brand: Klipperwerf 5, 2317 DX, Leiden, The different clades and thus it is polyphyletic Netherlands. (Andersen & Ekman 2005; Se´rusiaux et al. P. P. G. van den Boom: Arafura 16, NL-5691 JA, Son, 2010; Schmull et al. 2011): M. crassipes is The Netherlands. the type species of the distantly related genus E. Se´rusiaux (corresponding author): Evolution and Helocarpon, M. sylvicola and its relatives Conservation Biology Unit, University of Lie`ge, Sart Tilman B22, B-4000 Lie`ge, Belgium. belong to the Psoraceae, and all other species Email: [email protected] form a supported but poorly resolved clade 414 THE LICHENOLOGIST Vol. 46 with all accessions of the Ectolechiaceae and at high elevations. A working list of lichen the Pilocarpaceae s. str. nested into it. Further species found on the island was published phylogenetic studies with several loci are recently (van den Boom et al. 2011), based therefore needed before a well-supported de- on collections gathered over the last 15 years. limitation of the genus can be adopted. A total of 463 taxa were recorded, including Phylogenetic inferences from mtSSU se- several rare species, large expansion of the quences (Czarnota & Guzow-Krzemin´ska distribution range for others, and the detec- 2010) were used to investigate the species tion of an austral element (species whose dis- delimitation within the well-known, albeit tribution range is limited to the Southern notoriously difficult, Micarea prasina group. Hemisphere). Finally, analysis of the status Three lineages were resolved within M. of species endemic to the island pointed to micrococca, two being formally recognized at the urgent need for taxonomical studies in species level [M. byssacea (Th. Fr.) Czarnota speciose and poorly known genera. Micarea et al.andM. micrococca (Ko¨rb.) Gams ex is one of these and the taxonomy of species Coppins s. str.]. This study assumed that a found on the island is the aim of this paper. phylogenetic tree produced with data from a single locus can disentangle a difficult group of species, whereas gene trees represent ‘local Materials and Methods optima’ that require additional tests and This study is mainly based on extensive collections of appropriate statistical methods to yield a ro- Micarea, made by the authors on the island in 2008, bust species tree (Maddison 1997; Edwards and hosted in LG and the private herbaria of the first 2008; Knowles & Kubatko 2010; Camargo two authors (h refers to the private herbarium of the et al. 2012a, b; Carstens et al. 2013). It never- collector). Further specimens gathered in 2009 by one theless demonstrates that the variation within of us (ES), with Nicolas Magain, were also included. Anatomical data were gathered and measured in dilute the M. prasina group is quite substantial and KOH for conidia and paraphyses width, and on material requires detailed investigation. mounted in water for all other characters. In each collec- Even when excluding the species that be- tion, c. 5 well-developed ascospores representing the size long elsewhere in the Lecanorales, the genus and shape variation detected, conidia and paraphyses were measured up to 0 1 mm accuracy using camera is very diverse and occurs on all continents, lucida drawings. including the Antarctic, and its diversity is Chemical compounds were studied using several estimated at c. 90 species (Coppins 2009). methods: medulla under UV light; melting point and The area best sampled and studied is microcrystallization; thin-layer chromatography (TLC) certainly Europe (Czarnota 2007; Coppins using solvent system A (toluene/1,4-dioxane/acetic acid 180:45:5) or C (toluene/acetic acid 170:30) when gyro- 2009). Asia (from the western Himalaya to phoric acid was expected (Huneck & Yoshimura 1996; Japan) and tropical mountains in the three Orange et al. 2010) and visualization of spots with sul- continents represent unexplored and highly phuric acid sprayed over the plates, followed by heating promising biomes for studying the diversity at 110 Cforc. 5 min. Insoluble pigments are characterized of the genus. following the classification of Meyer & Printzen (2000). Re´union is a small island of 2512 km2 in the Indian Ocean, part of the Mascarene ar- chipelago along with two other islands, Mau- Results ritius and Rodrigues. It has a volcanic origin, Chemical compounds and one of the most active volcanoes in the world (Piton de la Fournaise) is located on Here we present data on the chemical com- its eastern side, with an elevation of 2631 m. pounds detected in our material. Details are Otherwise it is a serrated mountain range provided only for unusual compounds or (highest point at 3069 m at the Piton des those that can be difficult to distinguish. The Neiges), dissected in huge ‘cirques’ and numeral in TLC rubric refers to the height, very deep gorges. Its position in the tropical measured in mm, after a 45 mm run; H refers zone makes its climate warm and seasonal, to the spot colour after charring in H2SO4; although the temperature can be quite low U2 refers to UV examination at 254 nm, 2014 Micarea on Re´union Island—Brand et al. 415 while U3 refers to 360 nm; and finally extract thiophanic acid, two compounds are worth description refers to acetone extraction on a mentioning: alectorialic acid, diagnostic for microscope slide. M. alectorialica sp. nov., and a fatty acid (pro- tolichesterinic acid) reported here for the first Methoxymicareic acid (Elix et al. 1984) ¼ time in the genus and diagnostic for M. subli- prasina A (sensu Coppins 1983): TLC 26; thinella sp. nov. Another fatty acid, close to U2+, U3e white (‘‘blue-white’’ in Coppins rangiformic acid, was also detected in M. 1983); H yellow-brown; U3 after charring alectorialica sp. nov. dark brown; crystals not melting on heating Insoluble pigments have been characterized (or only after prolonged boiling). by Meyer & Printzen (2000); the following Micareic acid (Elix et al. 1984) ¼ prasina have been detected in our material (H being B(sensu Coppins 1983): TLC 26; U2+, the reaction to HCl, and N to HNO3): U3e white (‘‘blue’’ but less than prasina A in Coppins 1983); H (orange-) yellow; Cinereorufa-green: bright aeruginose, K+ U3 after charring vivid yellow; crystals green, H-- , N+ red, found in Micarea alec- melting on gentle heating. torialica sp. nov., M. boryana sp. nov., M. Prasinic acid (Elix et al. 1984) ¼ prasina incrassata, M. isidiosa sp. nov., M. lignaria, C(sensu Coppins 1983): TLC 24; U2+, M. melanoprasina sp. nov., M. pseudocop- U3e white (‘‘grey or pale mauve’’ or ‘‘col- pinsii sp. nov., M. pseudolignaria sp. nov., ourless’’ in Coppins 1983, depending on M. takamakae sp. nov. and M. tenuispora solvent system used); H grey; U3 after sp. nov. In M. borbonica sp. nov., M. cil- charring dark blue-grey; extract soon milky aoensis sp. nov., M. cinerea, M. peliocarpa by forming rod-shaped crystals. and M. pseudocoppinsii sp. nov., the same Methylhiascic acid: TLC 12; U2+ grey, U3-- ; pigment is detected but has a slightly dif- H grey; can only be separated from gyro- ferent colour (greyish blue and not bright phoric acid by TLC in solvent C (Orange aeruginose as in other species) and is a et al. 2010), not in A. It was found in surface incrustation of hyphae and not dif- Micarea borbonica sp. nov. and M. peliocarpa. fuse in the hymenial gelatine (as in other Unknown 1 ¼ unknown in M. melanoprasina species). We consider it a variant of the sp. nov.: TLC 31; U2+, U3-- ; H pale Cinereorufa-green that requires further greenish grey; U3 after charring light investigation. greenish grey; crystals melting on gentle Laurocerasi-brown: pale reddish brown, K+ heating, soon recrystallizing; extract clear, purple-brown, H-- ,N-- , found in Micarea gummy, without crystals; a substance prob- bebourensis sp. nov. ably related to micareic acid. Sedifolia-grey: greyish, K+ violet, found in Unknown 2 ¼ most probably confluentic acid, Micarea hedlundii and M. prasina. or a related compound, detected in M. Superba-brown: pale brown, K+ dull brown, takamakae sp. nov.: TLC 24; U2+, U3-- ; found in Micarea incrassata. H pale yellowish with greyish rim; extract Unknown 4: violet-red, K+ green, H-- , form- gummy, with radial wrinkles, later turning ing dark blue granules in N which get milky. more greenish when further applying K, Unknown 3 ¼ unknown in M. pseudolignaria found in the hymenium of Micarea isidiosa sp. nov.: TLC 28; U2+, U3-- ; H yellow- sp. nov. The granules are quite similar to brown (as in methoxymicareic acid); U3 those found in Lecidea hypnorum but, in after charring dark brown (as in methoxy- this species, they do not require application micareic acid); extract clear, gummy, white of N to be detected. within the ring, amorphous, not polarizing; Unknown 5: this substance can be detected small crystals in thallus not melting. at the goniocysts centre of Micarea hedlundii as tiny oily drops, which are not soluble in Besides the well-known compound argopsin acetone, and therefore cannot be analyzed and a xanthone that we tentatively refer to as with TLC. 416 THE LICHENOLOGIST Vol. 46
Additional brown pigment(s) that lack char- of Micarea so far restricted to the Southern acteristic reactions and have thus not been Hemisphere (Coppins & Kantvilas 1990; classified by Meyer & Printzen (2000) were Kantvilas & Elix 1994; Coppins 1999; Apt- also observed. root 2002; Fryday 2004; Ca´ceres et al. 2013) In several species (Micarea bebourensis sp. are: Micarea almbornii Coppins, M. corallo- nov., M. boryana sp. nov., M. levicula, M. thallina M. Ca´ceres et al., M. endoviolascens melanoprasina sp. nov. and M. prasina), and Coppins, M. intersociella (Sirt.) Coppins, M. in some collections of others (M. isidiosa sp. magellanica (Mu¨ll. Arg.) Fryday (¼ M. aus- nov., M. lignaria and M. pseudolignaria sp. troternaria Coppins & Kantvilas), M. pannar- nov.), the thallus hyphae are I+ light blue ica Fryday, M. subgranulans (Vain.) Aptroot after soaking in KOH (hemiamyloid); the re- and M. subternaria (Vain.) Aptroot. Micarea action is often weak and not always constant. poliocheila (Vain.) Aptroot, restricted to Such a hemiamyloid reaction has also been Brazil (Minas Gerais and Rio de Janeiro), detected in many European species of Mi- has been transferred to Bilimbia by Kalb carea but, to our knowledge, has never been (2007). found in other genera of the Lecanorales. Interestingly, habitats at the highest eleva- tions (>2000 m), and more precisely out- Taxonomy (Table 1) crops in dry and exposed conditions above We could detect and delimit 21 species in cloud level, yielded three species that are our material, out of which only eight can be either widespread in the temperate and assigned to an already described species; the boreal zones of the Northern Hemisphere other 13 are thus described as new to science (M. erratica and M. lignaria) or bipolar, being in this paper. A key for their identification is also found in subantarctic islands (M. incras- presented below. sata). Such observations demonstrate the im- Two subgeneric entities yielded several pressive dispersion capacity of lichen species new species and can be described as speciose (Honegger 1993; Romeike et al. 2002). groups on the island. They are the M. pelio- Micarea is absent in all habitats at low ele- carpa group and the M. prasina group (sensu vations on the island, including the rocky Coppins 1983). The first one comprises M. shores, open and anthropogenic vegetation, borbonica sp. nov., M. boryana sp. nov., M. recent or old lava flows, and remnants of cinerea, M. peliocarpa and possibly M. pseudo- low elevation rainforest dominated by the coppinsii sp. nov., while the second includes Sapotaceae (e.g. at the nature reserve of M. levicula, M. melanoprasina sp. nov., M. ‘Mare Longue’ near St-Philippe). Following prasina and possibly M. hyalinoxanthonica the vegetation typology provided by Cadet sp. nov. The latter group further includes (1977), Strasberg et al. (2005) and Lacoste material that contains methoxymicareic acid et al. (2011), we recognized six vegetation and belongs to M. micrococca s. lat. (Coppins types that accommodate species of Micarea 2009); its taxonomy needs further study and (Fig. 1): is not dealt with in this paper. Montane forest with large boles of Acacia Ecology and biogeography (Table 2) heterophylla, either in natural stands or in plantations, at >1500 m elevation. Interestingly, no species known so far only Montane forest known as ‘Bois de couleurs from the Southern Hemisphere was detected des Hauts’, easily characterized by the in our material, although one (M. boryana sp. abundance of tree species in the genera nov.) seems to be close to Micarea mutabilis, Dombeya (Malvaceae), Monimia and Tam- described and as yet known only from Tas- bourissa (Monimiaceae), and others in the mania (Australia). The austral element de- Euphorbiaceae, Rubiaceae, Rutaceae, Nuxia tected in the working list of lichens present verticillata, and locally the native bamboo on Re´union (van den Boom et al. 2011) can- Nastus borbonicus and tree ferns (Cyathaea), not thus be consolidated. The other species mainly at elevations >1300 m. 2014 Micarea on Re´union Island—Brand et al. 417
Table 1. Summary of the taxonomy adopted for material of Micarea from Re´union and substantiation of the decision taken
Species name Substantiation of the decision taken in this study
M. alectorialica sp. nov. Alectorialic acid is otherwise only present in M. magellanica and M. submilliaria which do not develop soralia M. bebourensis sp. nov. Easily recognized by its tuberculate thallus, globose and almost stipitate, light to dark brown apothecia M. borbonica sp. nov. Close to M. peliocarpa and differing by its 7-septate, longer and larger ascospores M. boryana sp. nov. Easily recognized by its narrowly clavate to needle-like ascospores, mainly 3-septate; a member of the M. peliocarpa group M. cilaoensis sp. nov. Easily recognized by its Lecania-like apothecia, close to M. denigrata which differs by its smaller macroconidia M. cinerea (Schaer.) Hedl. Identical with the material from Europe available to us, except for the absence of methylhiascic acid M. erratica (Ko¨rb.) Hertel Identical with the material from Europe available to us et al. M. hedlundii Coppins Identical with the material from Europe and East Africa available to us M. hyalinoxanthonica sp. nov. Distinguished from the other two xanthone-producing species by thallus, apothecia and ascospore characters; may belong to the M. prasina group M. incrassata Hedl. Almost identical with the material from Europe available to us
M. isidiosa sp. nov. Unique combination of isidiiform areolae and bacillar, 3–7-septate ascospores M. levicula (Nyl.) Coppins A representative of the M. prasina group, distinguished by production of gyrophoric acid; M. viridileprosa differs by thallus characters and narrower ascospores M. lignaria (Ach.) Held. Identical with the material from Europe available to us, although TLC analysis could not be performed M. melanoprasina sp. nov. A representative of the M. prasina group, distinguished by its conspicuous blackish prothallus and production of an unknown substance M. peliocarpa (Anzi) Identical with the material from Europe available to us Coppins & R. Sant. M. prasina Fr. Identical with the material from Europe, except for the K+ violet pigment which is present in the hypothecium and not in the epihymenium M. pseudocoppinsii sp. nov. Differs from M. coppinsii by shorter ascospores and absence of methylhiascic acid; probably a member of the M. peliocarpa group M. pseudolignaria sp. nov. Differs from M. lignaria by its 1-septate ascospores and different chemistry M. sublithinella sp. nov. Differs from M. lithinella by its 0–1-septate and larger ascospores and production of protolichesterinic acid M. takamakae sp. nov. Easily distinguished by small ascospores (8 8–10 2 3 1–3 8 mm) and production of confluentic acid M. tenuispora sp. nov. Unique combination of isidiiform areolae with narrowly clavate to needle-like, 3–5-septate ascospores
Wet montane thickets with tall ericoid species Open and partially grazed forests, dominated including Erica reunionensis,withexuberant by Sophora denudata, on the SW side of the development of ground bryophyte cover Piton de la Fournaise, at c. 1800–2000 m. (including species of Sphagnum), at c. Dry and exposed outcrops, at high elevations 1700–1900 m. (>2000 m) with xerophytic bryophytes. 418 THE LICHENOLOGIST Vol. 46
Table 2. Distribution of the representatives of the genus Micarea in the different habitats where they grow on Re´union
Montane Montane Montane forest forest Wet forest Disturbed Species name dominated (‘Bois de montane dominated Outcrops habitat (all in the genus by Acacia couleur des ericoid by Sophora at high below 700 m Micarea) heterophylla Hauts’) thickets denudata elevation elevation alectorialica ——X——— bebourensis —X———— borbonica XXXX—— boryana —X———— cilaoensis —X———— cinerea ——XX—— erratica ————X— hedlundii —X———— hyalinoxanthonica —X———— incrassata ————X— isidiosa ——X——— levicula X————— lignaria ————X— melanoprasina X————— peliocarpa XXX——— prasina X————— pseudocoppinsii X————— pseudolignaria ——X——— sublithinella XX———— takamakae —————X tenuispora —X————
The numbers of species of Micarea found in Like many isolated islands and archipelagos each habitat are presented in Table 2. As ex- all over the world, Re´union’s original biota pected, the most diverse habitat is the mon- has been devastated by human impact and tane forest ‘Bois de couleurs des Hauts’ with the local fauna and flora are much altered by nine species, including five new species which exotic species; therefore conservation evalua- have only been found here (M. bebourensis sp. tion and planning require detailed studies of nov., M. boryana sp. nov., M. cilaoensis sp. biodiversity processes (Lagabrielle et al. 2009). nov., M. hyalinoxanthonica sp. nov. and M. In Re´union, the native vascular flora is esti- tenuispora sp. nov.). It is immediately fol- mated at c. 500 species and more than 3500 lowed by the Acacia heterophylla forests, with have been introduced, 62 being highly inva- seven species. The soft, rather acidic, quick- sive (Baret et al. 2006). The question may drying and often peeling bark of that en- therefore arise regarding the status of lichen demic tree seems to be a very suitable habitat species observed on the island: are some spe- for many species of Micarea. The wet ericoid cies exotic? Although we have no clue at all thickets shelter six species, with two species to support this hypothesis, the question may new to science (M. alectorialica sp. nov. and be relevant for species growing on introduced M. isidiosa sp. nov.) being restricted to it. tree species, such as Cryptomeria japonica,a Furthermore, the Sophora open forest has species widely planted on Re´union. The case only two species, including M. borbonica sp. of three newly described species, M. boryana, nov. which is the only species present in all M. cilaoensis and M. tenuispora,isillustrative types of montane forest. The most distinctive as they have been detected only on trunks of M. takamakae sp. nov. is only known from a Cryptomeria. However, the acidic and peeling non-montane forest habitat. bark of this tree makes it a suitable habitat for 2014 Micarea on Re´union Island—Brand et al. 419
Fig. 1. Photographs of the main habitats for Micarea on Re´union. A, typical peeling trunk of Acacia heterophylla, near Savane Cimetie`re (NNW of Piton de la Fournaise): seven species of Micarea were found in this habitat; B, typical view of understorey of the montane ‘Bois de couleurs des Hauts’ (ForeˆtdeBe´bour): the richest habitat for Micarea with nine species; C, wet montane ericoid thickets with Erica reun- ionensis (along the track from Be´louve to Caverne Dufour): six species occur in this spectacular habitat, including one new species with soralia and producing alectorialic acid, and one new species with isidia; D, open forest dominated by Sophora denudata, along road to Bourg Murat (WNW of Piton de la Fournaise): only two species were found in this habitat, including the new M. borbonica, the most ubiquitous species on the island; E, exposed outcrops at high elevations near the Piton de la Fournaise: three species occur in such habitats: M. erratica, M. lignaria, both wide- spread in the Northern Hemisphere, and M. incrassata, a bipolar species; F, lava flow of April 2007 within the ‘Enclos’, photographed in May 2008: no species of Micarea were found on old or recent lava flows at low elevation. In colour online. 420 THE LICHENOLOGIST Vol. 46 pioneer and fast-growing taxa such as Mi- the Azores, an archipelago in the North At- carea, and is also easy to sample. We there- lantic Ocean with a high rainfall level and a fore postulate that these three species are much degraded vegetation, where Cryptomeria native to Re´union. That Micarea grows easily is also widely planted (Purvis & James 1993). on planted Cryptomeria is also observed in
Key to the species of Micarea in Re´union 1 Thallus with gyrophoric acid or alectorialic acid, C+ red ...... 2 Thallus without gyrophoric and alectorialic acids, C-- or C+ orange ...... 12 2(1) Thallus with genuine convex soralia ...... 3 Thallus not sorediate ...... 4 3(2) Thallus with alectorialic acid (K+ yellow, P+ yellow)...... M. alectorialica Thallus with gyrophoric acid (K-- ,P-- )...... M. pseudocoppinsii 4(2) Thallus with, or entirely made of, isidiiform areolae, ascospores long and narrow (>20 mm long and <3 mm wide) ...... 5 Thallus without isidiiform areolae ...... 6 5(4) Ascospores bacillar, straight to slightly curved; aeruginose hymenium contrasting with reddish purple hypothecium ...... M. isidiosa Ascospores acicular; hymenium colourless and hypothecium pale brown to dark bluish ...... M. tenuispora 6(4) Ascospores 0–1(–2)-septate ...... 7 Ascospores 3-septate or more ...... 9 7(6) Thallus formed of vivid green goniocysts; apothecia white ...... M. levicula Thallus not formed of goniocysts; apothecia white or grey to brown ...... 8 8(7) Thallus thin, uneven; macroconidia 38–48 mm...... M. cilaoensis Thallus tuberculate; macroconidia unknown ...... M. bebourensis 9(6) Ascospores fusiform, mainly or only 7-septate...... 10 Ascospores mainly or only 3-septate...... 11 10(9) Ascospores 3 4–4 4 mm wide; macroconidia 57–78 1 2–1 5 mm....M. cinerea Ascospores 3 0–4 1 mm wide; macroconidia 30–40 1 0–1 3 mm...... M. borbonica 11(9) Ascospores fusiform, 3 6–4 7 mm wide, less than 17 mm long . . . . . M. peliocarpa Ascospores narrowly clavate to needle-like, 2 0–2 5 mm wide, more than 17 mmlong ...... M. boryana 12(1) Thallus with xanthones, C+ orange-yellow ...... M. hyalinoxanthonica Thallus without xanthones, C-- ...... 13 13(12) Thallus composed of goniocysts ...... 14 Thallus not composed of goniocysts...... 18 14(13) Pycnidia stipitate, with tiny hairs, thallus with K+ violet pigment . . . M. hedlundii Pycnidia not stipitate, without hairs, or pycnidia absent ...... 15 15(14) Thallus with conspicuous black prothallus; presence of Cinereorufa-green pigment in thallus and apothecia and unknown pigment in apothecia . . . M. melanoprasina Thallus without dark prothallus ...... 16 2014 Micarea on Re´union Island—Brand et al. 421
16(15) Apothecia dark; micareic or confluentic acid present...... 17 Apothecia whitish to brownish; methoxymicareic acid present...... M. micrococca s. lat. (not treated) 17(16) Apothecia with K+ violet pigment; micareic acid present...... M. prasina Apothecia without K+ violet pigment but epihymenium blue-green, K-- ; confluentic acid present...... M. takamakae 18(13) Ascospores (1–)3–7-septate ...... M. lignaria Ascospores 0–1-septate ...... 19 19(18) Saxicolous; no lichen substances...... 20 Corticolous or terricolous; lichen substances present ...... 21 20(19) Apothecia marginate, up to 0 4 mm diam.; epihymenium aeruginose ...... M. erratica Apothecia immarginate, up to 0 6 mm diam., epihymenium brown, locally with blue- green pigment ...... M. incrassata 21(19) Ascospores 13–15 5–6 mm; protolichesterinic acid present . . . . M. sublithinella Ascospores 8–9 3–4 mm; chemistry different, not a fatty acid ...... M. pseudolignaria
The Species formed by strongly conglutinated radiating hyphae. Hypothecium pale or light bluish. Micarea alectorialica Brand, van den Hymenium c. 55 mm high; epihymenium dark Boom & Se´rus. sp. nov. blue. Paraphyses c. 1 5 mm thick, apically branched and anastomosing. Asci clavate, c. MycoBank No.: MB807670 46 12 mm, tholus of the Micarea-type. As- Thallus areolate, sorediate, soralia to 0 6 mm diam.; cospores 8 per ascus, fusiform 17 0–20 0 apothecia convex, up to 0 7 mm, shining black; asco- 4 5–5 6 mm, (1–)3-septate. spores fusiform, 17–20 4 5–5 6 mm, (1–)3-septate; production of alectorialic acid. Micropycnidia numerous, superficial be- Type: Re´union, ForeˆtdeBe´bour, path from Be´louve tween areolae, black, c. 70 mm; microconidia to cabane Dufour, 21 5 100 S, 55 31 360 E, 1890 m, narrow, fusiform with tapering ends, 6 6– wet montane ericoid thickets, on Erica, 2 June 2008, 8 3 0 9–1 0 mm. M. Brand 58863 (LG—holotype; hb. v.d. Boom, hb. Brand—isotype). Chemistry. Alectorialic acid (K+ yellow, P+ (Fig. 2A) yellow, C+ red) in thallus and soralia; fatty acid also present (extract as rangiformic acid, Thallus c. 5 cm across, formed by dispersed but larger agglomerations). Pigment in thallus areolae; areolae first globose, 0 1–0 2mm and apothecia belonging to Cinereorufa- across, pale-coloured, then bursting into a green. soralium; soralia to 0 6(–0 8) mm diam., con- vex, sometimes aggregated to almost stipitate, Etymology. The new species is named after white to light greenish; soredia 25–35 mm. the chemical compound that makes this spe- Photobiont micareoid, cells c. 5 0–5 5 mm. cies easy to recognize amongst the species of Cortex absent, but in young areolae upper Micarea present on Re´union. layer with more crystals; soredia strongly in- Habitat and distribution.OnErica stems spersed with crystals (alectorialic acid). at 1900–2000 m, in wet montane ericoid Apothecia up to 0 7 mm diam., shining thickets, abundant. black, convex without margin, often complex, and then made of fused small apothecia. Notes. Alectorialic acid is rare amongst Excipulum bluish in outer parts, inside pale, species of Micarea as it is so far detected in 422 THE LICHENOLOGIST Vol. 46
Fig. 2. Photographs of the new species described from Re´union. A, Micarea alectorialica sp. nov. (holotype); B, M. bebourensis sp. nov. (holotype); C, M. borbonica sp. nov. (holotype); D, M. boryana sp. nov. (holotype); E, M. cilaoensis sp. nov. (holotype); F, M. hyalinoxanthonica sp. nov. (holotype); G, M. isidiosa sp. nov. (holotype); H, M. pseudocop- pinsii sp. nov. (holotype). Scale ¼ 1 mm. In colour online. 2014 Micarea on Re´union Island—Brand et al. 423 only two species: M. magellanica (Mu¨ll. Arg.) dark, partly pale), almost globose, with con- Fryday (¼ M. austroternaria Coppins & Kant- stricted base. Excipulum greyish brown in vilas) restricted to the Southern Hemisphere outer parts, pale inside. Hypothecium pale (Coppins & Kantvilas 1990; Fryday 2004), (reddish) brown, conglutinate. Hymenium and M. submilliaria (Nyl.) Coppins, a mostly c. 50–60 mm high; epihymenium patchily Northern Hemisphere species (Coppins brownish. Paraphyses 0 7–1 1 mm thick, 2009), but both do not produce soralia, branched and strongly anastomosing; some although the surface of the latter is usually paraphyses with a thickened apex (1 8 mm). eroding to form sorediate patches. Only Asci cylindrical, c. 40–53 10 mm, tholus of three other species of Micarea produce sora- the Psora-type. Ascospores 8 per ascus, ovate to lia: M. coppinsii Tønsberg, M. pseudocoppinsii shortly bacillar, 11–14 3 3–4 3 mm, 1(–3)- (here described as new) and M. viridileprosa septate. Coppins & van den Boom (soralia usually Micropycnidia inconspicuous, immersed, not clearly delimited and forming a leproid with brownish top, c. 35 mm. Microconidia surface); they can all be distinguished by the bacillar with rounded ends, some slightly production of gyrophoric acid (van den Boom curved, 5 9–7 1 0 9–1 1 mm. & Coppins 2001; Coppins 2009). Chemistry. Gyrophoric acid (K-- ,P-- ,C+ Additional specimen examined. Re´union: same locality red), mainly in the thallus, epihymenium, as the type, 21 050 3100S, 55 300 4100E, 1910–2000 m, wet ericoid thickets with E. reunionensis and Phylica nitida, on excipulum and pycnidia wall. Brownish pig- Erica, 9 xi 2009, N. Magain & E. Se´rusiaux s. n. (LG). ment belonging to Laurocerasi-brown, in epihymenium, slightly brownish, K+ purple- brown; subhymenium and hymenium in K Micarea bebourensis Brand, van den slightly rose-coloured. Boom & Se´rus. sp. nov. Etymology. This new species is named after MycoBank No.: MB807671 the Be´bour forest, one of the largest and best Thallus typically tuberculate when young, tubercles preserved montane forests on Re´union, where 0 10–0 15 mm across; apothecia up to 0.6 mm, greyish many interesting lichen species grow. brown; ascospores ovate to shortly bacillar, 11–14 3 3–4 3 mm, 1(–3)-septate; production of gyrophoric acid. Habitat and distribution. On introduced Type: Re´union, ForeˆtdeBe´bour, trail to Casse´ de Ta- tree species, including Cryptomeria japonica kamaka, 21 07 50 S, 55 34 50 E, 1340 m, wet montane and decaying wood, in lower montane forest rainforest, on rotting standing trunk, 1 June 2008, P. at 1200–1500 m. van den Boom 40344 (LG—holotype; hb. v.d. Boom, hb. Brand—isotype). Notes. The almost globose apothecia, (Fig. 2B) sometimes almost stipitate, and the thallus tubercles (best observed in young thalli) rep- Thallus up to 2 cm diam., pale to brownish resent the most obvious characters to distin- greyish or greenish, uneven, typically tuber- guish this species. Its generic assignment to culate when young, with tubercles 0 10– Micarea is questionable as the asci are cylin- 0 15 mm broad and high, eventually formed drical (not clavate as in other species of the of coarse and agglomerated granules, first core group of Micarea) and the tholus is pale, eventually breaking up at the top of much akin to the Psora-type (sensu Ekman tubercules or at the margins of granules, and et al. 2008). The anatomy of apothecia (glo- forming a rough blue-grey surface (resem- bose apothecia with a constricted base and bling tiny soralia, but no genuine soredia are development of excipular tissue) makes it formed). Cortex absent, often with thin, clear close to Micarea lignaria and M. pseudolignaria epinecral layer. Photobiont micareoid. Thallus sp. nov.; M. bebourensis is easily differentiated filled with crystals (gyrophoric acid). from both by the absence of an aeruginose Apothecia up to 0 6 mm diam, light to dark pigment and the production of gyrophoric greyish brown, often unevenly coloured (partly acid. 424 THE LICHENOLOGIST Vol. 46
Additional specimens examined. Re´union: same locality slip), 30–42 1 0–1 3 mm (measured under as the type, P. van den Boom 40370 (h); Be´louve, pressed cover slip), faintly 3-septate. Micro- 21 3 740 S, 55 32 290 E, 1500 m, montane forest, on planted Cryptomeria, 2008, M. Brand 58215 (h); Plaine- pycnidia inconspicuous, semi-superficial, c. des-Palmistes, Ravine Se`che, 21 8 630 S, 55 35 310 E, 35 mm, pale. Microconidia narrow fusiform 1200 m, planted exotic trees in sheltered valley, on Cryp- with tapering ends, 5–7(–10) 0 8–1 0 mm. tomeria, 2008, M. Brand 58141 (h); ForeˆtdeBe´bour, path W of Col de Be´bour, 21 07 20 S, 55 33 60 E, 1490 Chemistry. Gyrophoric and methylhiascic m, Cryptomeria plantation, on Cryptomeria, 2008, P. van acids (K-- ,P-- , C+ red), mainly in thallus, den Boom 40322 (h). hymenium, excipulum and pycnidia. Pigment in apothecia belonging to Cinereorufa-green. Micarea borbonica Brand, van den Boom & Se´rus. sp. nov. Etymology. The name of this new species refers to the old name of the island, ‘Ile Bour- MycoBank No.: MB807672 bon’ in French. It is indeed characteristic of Thallus small, to 1 5 cm, dull white to pale greenish; all habitats, natural or anthropogenic, in the apothecia up to 0 5 mm diam., whitish to pale brownish or pale to dark bluish grey; ascospores fusiform with montane forest zone of the island. rounded ends, 15–27 3 0–4 1 mm, 3–7-septate; mac- roconidia strongly and repeatedly curved sigmoid, 30– Habitat and distribution. On native and 42 1 0–1 3 mm, indistinctly 3-septate; production of exotic trees (Acacia heterophylla, Cryptomeria, gyrophoric acid. Sophora denudata, Platanus and on branches Type: Re´union, WNW of Piton de la Fournaise, along of Erica), and decorticated wood, between road to Bourg Murat, 21 11 90 S, 55 36 90 E, 1970 m, small forest with Sophora denudata trees, on Sophora, 4 800–2000 m. June 2008, P. van den Boom 40644 (LG—holotype; hb. v.d. Boom, hb. Brand—isotype). Notes. Ascospores in this new species are typically 7-septate, although 3–6-septate (Fig. 2C) ones can be found usually in immature or poorly developed apothecia. Micarea borbon- Thallus small, to 1 5 cm across, dull white ica is closely related to M. peliocarpa, a spe- to pale greenish, uneven, c. 50 mm high or cies also found in Re´union; it differs in hav- consisting of small granules, some of them ing convex to subglobose areoles, longer and resembling small isidia. Upper part c. 15 mm narrower ascospores (13 6–17 0 3 6–4 7 without algae, of loosely interwoven hyphae. mm in collections of M. peliocarpa from Re´- Photobiont micareoid, cells c.5mm, photo- union), and is usually 7-septate (mostly 3- biont layer inspersed with crystals (gyrophoric septate in M. peliocarpa). A further related acid). species is M. cinerea, also found on Re´union, Apothecia up to 0 2–0 5 mm diam., whit- that can be distinguished by larger spores (24– ish to pale brownish or pale to dark bluish 30 3 4–5 0 mm) and much longer macroco- grey, often coloured only partially, with pale nidia (57–78 1 2–1 5 mm). See further margin, flat or slightly convex to almost notes under M. cinerea and M. peliocarpa. globose, adnate with constricted base. Excip- Additional specimens examined. Re´union: Piton de la ulum pale or bluish grey in outer parts, 0 pale inside. Hypothecium colourless. Hyme- Fournaise, 0 5kmNWofGıˆte du Volcan, 21 12 07 S, 55 41 530 E, 2040 m, mixed wood on N-slope, on Aca- nium c. 55 mm high; epihymenium colourless cia heterophylla, 2008, M. Brand 58361, 58376 (h); or partly blue-grey. Paraphyses 1 3–1 6 mm Plaine-des-Palmistes, 21 6 980 S, 55 37 130 E, 1100 m, thick, branched and anastomosing. Asci Platanus branches in garden, 2008, M. Brand 58696 clavate, 33–40 12–15 mm, tholus of the (h); ibid., on decorticated wood of telegraph pole, 2008, P. van den Boom 40621 (h); Cilaos, Foreˆtdu Micarea-type. Ascospores 8 per ascus, fusi- Grand Matarum, path to Cabane Dufour, 21 7 080 S, form with rounded ends, 15–27 3 0–4 1 55 29 190 E, 1770 m, wet montane ericoid thickets on mm, (3–)7-septate, often curved. steep SW slope, on thin twigs, 2008, M. Brand 59713 Macropycnidia not rare, globose, 100– (h); WNW of Piton de la Fournaise, along road to Bourg Murat, 21 11 60 S, 55 37 50 E, 2090 m, disturbed wood 140 mm, top often blue-grey; macroconidia with Sophora denudata,onSophora, P. van den Boom strongly and repeatedly curved sigmoid 40665 (h); ESE of Le Tampon, NNE of Petite-ıˆle, (when observed under unpressed cover 21 18 90 S, 55 35 40 E, 800 m, open place with mature 2014 Micarea on Re´union Island—Brand et al. 425
Cryptomeria trees, on Cryptomeria, 2008, P. van den crystals of gyrophoric acid, mainly near the Boom 40806 (h). apex. Mesoconidia ovate, 4 0–4 5 (1 3–) 1 5–1 6 mm. Micropycnidia rarely present, in- Micarea boryana Brand, van den Boom conspicuous, with a dark brownish wall. &Se´rus. sp. nov. Microconidia fusiform, 4 0–4 5(–5 0) 1 0– MycoBank No.: MB807673 1 1 mm. Thallus areolate, greenish to greyish brown, eroding Chemistry. Gyrophoric acid (K-- ,P-- ,C+ when old; apothecia to 0 4(–0 6) mm, grey to bluish red) detected in thallus, but also in pale parts black; ascospores narrowly clavate to needle-like, 16– 27 2 0–2 5 mm, (1–)3(–4)-septate; production of of apothecia and in mesopycnidia. Pigment gyrophoric acid. in thallus and apothecia belonging to Ciner- Type: Re´union, Cirque de Cilaos, N of Cilaos, Foreˆt eorufa-green. Brown pigment in hypothecium du Grand Matarum, trail to Caverne Dufour, 21 07 30 S, K+ reddish to purple-brown. 55 29 20 E, montane forest, 1420 m, on Cryptomeria,31 May 2008, P. van den Boom 40264 (LG—holotype; hb. Etymology. This new species is named after v.d. Boom, hb. Brand—isotype). the French officer and first lichen collector (Fig. 2D) on the island, J. B. G. M. Bory de Saint- Vincent (1778–1846) who described several Thallus up to 5 cm across, greenish to grey- spectacular and now well-known species, ish brown, continuous, made of small flat to such as Cladonia candelabrum, C. giganteum, convex or even subglobose areolae, with a Lobaria retigera, Stereocaulon vulcani and Sticta thin endo- to epiphloeodal dark bluish black ambavillaria. prothallus made of narrow hyphae, colour- Habitat and distribution. Found at several less or partly dark brownish or bluish hyphae. localities on the island, between 1200–1500 Areolae either flat (0 1–0 6 mm broad, 0 05 m, always epiphytic on the exotic Cryptomeria mm thick) or subglobose (granules c. 0 1 japonica. mm across), eroding at the margins in old thalli, slightly shiny. Cortex absent, but a Notes. We first assigned these collections thin, clear epinecral layer can be observed. to Micarea mutabilis Coppins & Kantvilas, Photobiont micareoid, cells c. 5–8 mm. Thal- described from Tasmania (Coppins & Kant- lus filled with crystals of gyrophoric acid. vilas 1990). According to the original descrip- Apothecia up to 0 4(–0 6) mm diam., grey tion, the latter, however, clearly differs by to bluish black, sometimes with a pale mar- several important characters: absence of pro- gin, first flat, without a distinct margin, then thallus, larger asci (40–50 9–10 mm), asco- semiglobose. Excipulum with radiating hyphae, spores longer and more septate [21–41 varying in colour (often in same apothecium) 2 2–3 0 mm; (1–)5–7(–8)-septate], absence from pale to superficially bluish and centrally of conspicuous pycnidia producing mesoco- light brown. Hypothecium pale or brownish. nidia (the mesoconidia described for that Hymenium c. 30–55 mm high; epihymenium species are actually microconidia). We believe patchily greyish-bluish. Paraphyses 1 0–1 2 such discrepancies are worth species recogni- mm thick, branched and anastomosing. Asci tion. Other species with needle-shaped or 25–37 10–15 mm, tholus of the Micarea- acicular ascospores are (Coppins 2009): type. Ascospores 8 per ascus, in one bundle, Micarea globulosella (Nyl.) Coppins, distin- narrowly clavate to needle-like, 16 0–27 0 guished by its black and subglobose apothecia 2 0–2 5 mm, (1–)3(–4)-septate, sometimes and smaller ascospores [(10–)12–19(–24) apically slightly enlarged and tapering to- 2 0–2 5(–3 0) mm]; M. synotheoides (Nyl.) wards their basal ends. Coppins distinguished by the absence of Mesopycnidia often present and conspicu- chemical compounds and variable asco- ous, sessile, c. 160–230 mm diam. and 200– spores [needle-like, fusiform to rod-shaped, 270 mm in height, dark brownish, with white (0–)1–3-septate, 14–35(–43) 1 8–2 5(–3 0) blob of conidia on top; pigment brown to mm]; M. pycnidiophora Coppins & P. James, greenish brown or aeruginose, K-- ; wall with together with M. stipitata Coppins & P. 426 THE LICHENOLOGIST Vol. 46
James, both being easily recognized by their found: microconidia bacilliform, 5 2–7 1 numerous, stipitate pycnidia; and finally M. 0 9–1 1 mm. tenuispora sp. nov., described here as new -- -- from Re´union, which is easily recognized by Chemistry. Gyrophoric acid (K ,P ,C+ its isidiiform thallus areolae. red) detected in thallus, hymenium, excipu- lum, pycnidia. Pigment in apothecia belong- Additional specimens examined. Re´union: same locality ing to Cinereorufa-green. as the type, 2008, P. van den Boom 41041 (h); Plaine- des-Palmistes, Ravine Se`che, 21 8 630 S, 55 35 310 E, Etymology. The name chosen for this new 1200 m, planted exotic trees in wood in sheltered valley, species refers to the most spectacular cirque on Cryptomeria,2008,M. Brand 58141, 58146 (h), P. van den Boom 39776 (h); Be´louve, 21 3 740 S, 55 32 290 E, on the island, the Cirque de Cilaos, where 1500 m, disturbed montane forest, on Cryptomeria, this rare species has been discovered. 2008, M. Brand 58206 (h). Habitat and distribution. Known from a single locality, on the exotic Cryptomeria Micarea cilaoensis Brand, van den japonica at the margin of disturbed natural Boom & Se´rus. sp. nov. forest, at 1420 m elevation. MycoBank No.: MB807675 Notes. Micarea cilaoensis could be confused Thallus thin, uneven, consisting of slightly convex gran- with a species of Lecania because of its apoth- ules, green; apothecia to 0 4 mm, adnate, with con- ecia with a brownish disc and whitish margin, stricted base; ascospores ovoid to ellipsoid, 11–12 and its well-developed excipulum. However, 4 0–4 4 mm, 0–1-septate; production of gyrophoric acid. Type: Re´union, Cirque de Cilaos, Foreˆt du Grand examination of the hamathecium and ascus- Matarum, trail to Caverne Dufour, 21 07 30 S, type immediately point to the genus Micarea. 55 29 20 E, 1420 m, montane forest, on Cryptomeria,31 Also remarkable are the large ampulliform May 2008, P. van den Boom 40230 (LG—holotype; hb. pycnidia producing macro- and microconi- v.d. Boom, hb. Brand—isotype). dia together. Micarea cilaoensis is close to M. (Fig. 2E) denigrata (Fr.) Hedl., a widespread species in the Northern Hemisphere which also produces Thallus rather extensive (3 cm across), large ampulliform pycnidia; the latter differs green, thin (65–90 mm), uneven, made of by its narrower ascospores [(7–)9–16(–18) slightly convex granules. Cortex absent. Photo- 2 0–3 3(–3 5) mm] and much smaller biont micareoid, cells c.5–9mm. Photobiont macroconidia (12–24 c. 1 mm; Coppins layer inspersed with crystals of gyrophoric 2009). acid. Apothecia up to 0 4 mm diam., whitish to Additional specimen examined. Re´union: same locality as the type, P. van den Boom 40229 (h, hb Brand 62255). greyish with pale margin, flat or slightly con- vex, adnate with constricted base. Excipulum Micarea cinerea (Schaer.) Hedl. well developed, made of thin reticulate hy- phae, filled with crystals, colourless. Hypothe- Thallus up to c. 1 cm across, made of small cium colourless. Hymenium c.35–42mmhigh; granules (c. 0 1–0 2 mm diam.), some of epihymenium colourless or faintly greenish them resembling small isidia, white to pale grey, in K unchanged or locally brownish. grey. Paraphyses 1 3 mm thick, branched and anas- Apothecia to 0 3 mm diam., whitish, beige, tomosing. Asci clavate, c.28 12 mm, tholus pale yellowish brown or dark bluish grey, of the Micarea-type. Ascospores 8 per ascus, often unevenly coloured, with a slightly pale ovoid to ellipsoid, 11–12 4 0–4 4 mm, 0– margin, flat or convex to almost globose, 1-septate. adnate with constricted base. Hymenium c. Pycnidia not rare, superficial or semi- 50 mm; epithecium yellowish or more rarely immersed, white, ampulliform, c. 150 mm bluish, K-- . Hypothecium hyaline. Asci 37– diam., top with crystals; macroconidia straight 45 15–20 mm, tholus of the Micarea- or more or less curved, 38–48 0 9–1 1 type. Ascospores 8 per ascus, fusiform, 24– mm, non-septate, cylindrical with rounded 30 3 4–5 0 mm, (3–)7-septate, straight to ends; in same pycnidia, microconidia can be often curved. 2014 Micarea on Re´union Island—Brand et al. 427
Pycnidia large, c. 200 mm diam., partly Chemistry. No lichen substances detected. immersed in areoles. Macroconidia long and Habitat and distribution. On volcanic rocks slightly flexuose, 7–10-septate, 57–78 1 2– in meadow, at c. 2000 m elevation. 1 5 mm. Notes. This single collection falls within Chemistry. Gyrophoric acid (K-- ,P-- ,C+ the variation of European specimens of that red), mainly in thallus, hymenium, excipu- species, including the size and height of lum and pycnidia wall; methylhiascic acid apothecia, and we have no doubt it belongs absent. Pigment in apothecia belonging to to the widespread M. erratica. The species Cinereorufa-green. has been recognized as the sole representative of the new genus Leimonis (Harris 2009), an Habitat and distribution. On Erica in wet option not retained so far (Coppins 2009). montane ericoid thickets at 2000 m, and on trunk of Sophora at similar elevation. Specimen examined. Re´union: track Bourg Murat to Piton de la Fournaise, 21 12 630 S, 55 36 570 E, 1970 Notes. In European collections of this m, volcanic rock outcrops in meadow on NW slope, on stones, 2008, M. Brand 59027 (h). species, methylhiascic acid is detected as well as gyrophoric acid; this acid has not Micarea hedlundii Coppins been detected in the material from Re´union. See further data under M. borbonica. Micarea Thallus less than 1 cm across, made of cinerea is a widespread species as it occurs abundant and aggregated goniocysts, forming in Europe, North and Central America, Asia coralloid masses, dark greenish to greyish. and Australia (Tasmania), according to Goniocysts 17–32 mm diam., locally with Coppins (2009). coarse crystals. Hyphae partly with brown- grey pigment (K+ violet). Photobiont mi- Specimens examined. Re´union: ForeˆtdeBe´bour, trail careoid, c. 5 mm. from Gıˆte de Be´louve, 3 5 km to the south-west, to Caverne Mussard, 21 05 30 S, 55 31 30 E, 1980 m, wet Apothecia absent. montane ericoid thickets, 1980 m, on Erica, 2008, P. Pycnidia stipitate; stipe 0 24–0 40 mm van den Boom 40526, 40555 (h, hb Brand 61471); high, 100–160 mm thick, surface with loose, WNW of Piton de la Fournaise, along road to Bourg curly, grey tomentum, made of c. 1 7 mm Murat, 21 11 90 S, 55 36 90 E, small open forest with Sophora denudata,onSophora, 1970 m, 2008, P. van thick hyphae; pycnidia c. 140 mm, 1–3(–5) den Boom 40644A (h). on a stipe. Mesoconidia elliptic, c. 4 2–5 2 1 6–2 1 mm. Micarea erratica (Ko¨rb.) Hertel et al. Chemistry. Unknown 5 at the centre of Thallus of scattered whitish areoles; areo- goniocysts, and Sedifolia-grey. lae up to 0 5 mm diam., with a c. 15 mm thick Habitat and distribution. On soft rotting epinecral layer. Photobiont not micareoid, cells wood of standing trunks in montane forest, to 15 mm. at 1400–1900 m elevation. Apothecia up to 0 4 mm broad, black, flat, with a distinct margin. Excipulum of conglu- Notes. Apothecia are absent in our material tinated radiate hyphae, outer part aerugi- from Re´union, but otherwise we could not nose, inner part hyaline. Hypothecium dark detect any differences with available collec- brown. Hymenium c. 25 mm; epihymenium tions from Western Europe, from the Alber- aeruginose. Paraphyses simple, c. 1 8 mm tine Rift in Africa and most recent descrip- thick. Asci c. 25 10 mm, tholus of the tions (Czarnota 2007; Coppins 2009). The Micarea-type. Ascospores 8 per ascus, nar- recently described M. tomentosa Czarnota & rowly ellipsoid, c.7 3 mm. Coppins (Czarnota 2007), known from cen- Mesopycnidia numerous, c. 70–100 mm, tral and northern Europe (Suija et al. 2008), immersed, upper side dark bluish, lower also has stalked pycnidia but differs by the side unpigmented. Mesoconidia c. 3 2–4 5 complete absence of pigment H in the gonio- 1 1–1 9 mm. cysts. 428 THE LICHENOLOGIST Vol. 46
Specimens examined. Re´union: Cilaos, ForeˆtduGrand Chemistry. Xanthone (thiophanic acid?) Matarum, along path to Cabane Dufour, 21 6 290 S, in thallus and apothecia (K+ yellow; P-- ,C+ 55 29 580 E, 1870 m, low forest on steep SW slope, on Dombeya tree, 2008, M. Brand 59739 (h); Foreˆtde and KC+ orange). Crystals in thallus and Be´bour, trail to Takamaka, 21 05 90 S, 55 34 30 E, apothecia, in streaks in hymenium and ex- 1370 m, wet rainforest, on rotting standing trunk, 2008, cipulum. P. van den Boom 40883 (h).—Rwanda: Virunga volca- noes, southern flank of the Karisimbi, c. 3400 m, open Etymology. The name chosen combines the forest with Hagenia abyssinica and Hypericum revolutum, chemical compound produced by this new on big bole of Hagenia, 1974, J. Lambinon 74/1524 (LG; confirmed by B. J. Coppins, 1992); ibid., species and a diagnostic character of its 01 290 12 5600S, 029 280 37 3800E, c.3200m,onHagenia, apothecia, being translucent when wet. x 2010, E. Se´rusiaux s. n. (LG). Habitat and distribution. Only known from the type locality. It grows on decaying liver- Micarea hyalinoxanthonica Brand, worts on bole of a Dombeya in rainforest van den Boom & Se´rus. sp. nov. (c. 1400 m). MycoBank No.: MB807676 Notes. Young globose apothecia of no Thallus thin, pale greenish, made of goniocysts; apothe- more than 100 mm diam. contain already cia up to 0 6 mm, adnate pale beige, translucent when ripe asci. TLC gives the same result as with wet; ascospores bacillar, 13 2–17 0 3 2–3 6 mm, 3- septate; production of xanthone (thallus and apothecia Micarea xanthonica Coppins & Tønsberg, C+ orange). which contains thiophanic acid (Coppins & Type: Re´union, ForeˆtdeBe´bour, Sentier de Taka- Tønsberg 2001). This species can be distin- maka, 21 06 480 S, 55 34 020 E, 1380 m, low trees in mon- guished by its thallus entirely made of dense tane forest, on Dombeya, 7 June 2008, M. Brand 59293 goniocysts, very similar to that of M. prasina, (LG—holotype; hb. v.d. Boom, hb. Brand—isotype). apothecia rare or even absent, and ascospores (Fig. 2F) (0–)1(–3)-septate. A further species contain- ing xanthone is Micarea isabellina Coppins & Thallus c. 3 cm diam., pale greenish, thin, Kantvilas, so far only known from Tasmania, overgrowing decaying bryophytes, soft, felty, growing on rocks or terricolous (Coppins & made of small (20–30 mm diam.) goniocysts. Kantvilas 1990). It can be distinguished by Photobiont micareoid, 5–8 mm, individually its areolate to irregularly warty thallus, black surrounded with a mantle of furcate hyphae, apothecia and larger ascospores (19–26 partly aggregated in goniocysts; thallus also 3 5–4 0 mm). Furthermore, M. hyalinoxan- with strait, thick-walled hyphae (c. 2 mm) and thonica can be distinguished by its ascospores sometimes including crystals. that are distinctly constricted at septae, with Apothecia abundant, 0 1–0 5(–0 6) mm more or less globose cells. diam., adnate with constricted base, pale beige, translucent when wet, rarely greyish, Micarea incrassata Hedl. first flat, with hardly distinct margin, then convex. Excipulum thin, chondroid, with Thallus c. 2 cm across, composed of dis- thin, branching hyphae, outer parts some- persed to adjacent areolae on rock, over- times with protruding hyphae, giving a hairy growing mats of cyanophyta (Stigonema); appearance. Hypothecium pale or patchily light areolae beige greenish, globose to convex, brownish. Hymenium c. 50 mm high, strongly 0 1–0 2 mm. Photobiont micareoid, cells c. 5 conglutinated, colourless. Paraphyses c. 1 3 mm. Cortex absent, thallus without crystals. mm thick, branched and anastomosing. Asci Genuine cephalodia absent, but thallus clearly clavate, c. 45–50 12 mm; tholus of Mi- associated with Stigonema, the cells of which carea-type. Ascospores 8 per ascus, bacillar, are clearly intermingled with hyphae within 13 2–17 0 3 2–3 6 mm, 3-septate, dis- the areolae. tinctly constricted at septa, with more or less Apothecia up to 0 6 mm diam., black, con- globose cells. vex without margin. Excipulum hardly distinct Pycnidia absent. from hymenium. Hypothecium dark brown. 2014 Micarea on Re´union Island—Brand et al. 429
Hymenium c. 40 mmhigh;epihymenium brown, 29 8 2 8–2 9 mm, 3–7-septate; production of gyro- locally with blue-green pigment. Paraphyses phoric acid. Type: Re´union, ForeˆtdeBe´bour, path from Be´louve c. 1 8 mm thick, mostly simple, occasionally to cabane Dufour, 21 05 100 S, 55 31 360 E, 1890 m, branched. Asci clavate, c. 27–30 8–10 mm, wet montane ericoid thickets, on Erica, 2 June 2008, M. tholus of the Micarea-type. Ascospores 8 per Brand 58862 (LG—holotype; hb. v.d. Boom, hb. ascus, ellipsoid, 8 6–10 0 4 2–5 0 mm, Brand—isotype). non-septate, wall c. 0 3 mm thick. (Fig. 2G) Micropycnidia scarce, between areolae, wall dark brown, c.60mm; microconidia bacil- Thallus up to 3 cm diam., greenish to lar, 5 7–6 8 1 0–1 1 mm. brownish, discontinuous, made of small glo- bose or isidia-like areolae on thin endo- to Chemistry. No lichen substance. Pigment epiphloeodal prothallus with narrow colour- in thallus and apothecia belonging to Ciner- less or partly brownish or bluish hyphae. Are- eorufa-green, pigment in hypothecium be- olae first granular (0 1 mm across) eventually longing to Superba-brown. becoming isidiiform (0 30 0 05 mm), un- Habitat and distribution. Only known from branched or forked. Cortex absent. Photobiont a single collection, on recently exposed vol- micareoid. Thallus filled with crystals (gyro- canic rock at high elevation. phoric acid). Apothecia up to 0 4 mm diam., black, first Notes. Our single collection is very close to flat, without distinct margin, then semiglo- the European material of Micarea incrassata bose. Excipulum with radiating hyphae, pur- that we could examine. It differs, however, plish, with few crystals. Hypothecium reddish by its constantly 0-septate ascospores (vs. 0– or bluish. Hymenium c. 40 mmhigh;epihyme- 2-septate in European material), smaller asci nium and hymenium intense blue-green. Par- and thus lower hymenium, and absence of aphyses c. 1 4 mm thick, branched. Asci c. 30– a chemical compound (European material 34 10 mm, tholus of the Micarea-type. has thallus crystals, not dissolving in K or Ascospores 8 per ascus, bacillar, straight to HCl, but slowly in acetone). Pending further slightly curved, 22 5–29 8 2 8–2 9 mm, studies, we don’t consider these variations 3–7-septate. to be worth taxonomic recognition. Micropycnidia inconspicuous, immersed, Micarea incrassata is a further example of a with dark bluish wall. Microconidia fusiform, species widespread in the Northern Hemi- 3 7–4 1 0 9–1 0 mm. sphere and also detected in Australia (New South Wales) and subantarctic islands, and Chemistry. Gyrophoric acid (K-- ,P-- ,C+ that occurs patchily on summits in tropical red), mainly in thallus, but also detected areas. A similar example is Fuscopannaria in apothecia. Blue pigment in thallus and praetermissa, present on high mountains in apothecia K+ intense green, N+ red and H+ East Africa and in Re´union (van den Boom darker, most probably belonging to Ciner- et al. 2011). eorufa-green; red pigment mainly present in hymenium (unknown 4) K+ green, N pro- Specimen examined. Re´union: Piton de la Fournaise, ducing a precipitate of dark blue granules, S slope of Pas de Bellecombe, 21 12 60 S, 55 41 80 E, 2250 m, recent volcanic rock on steep S-slope, 2008, H-- . M. Brand 58333 (h). Etymology. The name refers to the isidioid Micarea isidiosa Brand, van den Boom areolae forming the thallus. &Se´rus. sp. nov. Habitat and distribution. On Erica stems in MycoBank No.: MB807677 wet ericoid thickets at elevations between Thallus formed of isidioid granules growing on partly 1900–2000 m. dark prothallus, greenish to brownish; apothecia up to 0 4 mm in diam, black; hypothecium purplish or reddish; Notes. The thallus of this species seems ascospores bacillar, straight to slightly curved, 22 5– to be isidiate; however, the ‘isidia’ are not 430 THE LICHENOLOGIST Vol. 46 outgrowths on a thallus or areolae as they Habitat and distribution. On natural stands grow directly on the prothallus; they thus of Acacia heterophylla; so far only found near represent isidiiform areolae. The thallus and the Piton de la Fournaise, where it grows apothecia of this species are much akin to with the related M. prasina. those of M. tenuispora, also described as new in this paper; the bacillar ascospores, more Notes. Micarea levicula has been recog- vivid colours in apothecium section, with an nized as a member of the M. prasina group aeruginose tinge in the epihymenium con- by van den Boom & Coppins (2001), on the trasting with a reddish purple hypothecium, basis of its type and only available collection and the unique precipitate of dark blue gran- from Cuba (H-Nyl. 20762). The short de- ules in N in the hymenium are the diagnostic scription provided matches the material col- characters. lected in Re´union. Micarea viridileprosa Cop- Additional specimens examined. Re´union: Foreˆtde pins & van den Boom, also a member of the Be´bour, path from Be´louve to cabane Dufour, 3 km prasina group with gyrophoric acid, differs by from Gıˆte Be´louve, 21 05 100 S, 55 31 360 E, 1890 m, the loose, non-coralloid and smaller, more tall ericoid thickets in montane forest, on Erica, 2008, soredia-like goniocysts, the localization of the 0 0 M. Brand 58865 (h); ibid., 21 05 3 S, 55 31 3 E, 1980 gyrophoric acid crystals on the outer parts of m, on Erica, 2008, P. van den Boom 40529 (h). the hyphae surrounding the goniocysts, and finally by the narrower spores (c. 11–14 Micarea levicula (Nyl.) Coppins 3 0–3 4 mm). Micarea viridileprosa is so far Thallus diffuse, vivid green, made of deli- known from western Europe, where it can cately and finely coralloid goniocysts; gonio- be quite common, and Tasmania (van den cysts 25–53 mm diam. or cylindrical (c. 50 Boom & Coppins 2001; Coppins 2009). The 18 mm), round to ovate, often more or less recently described M. corallothallina (Ca´ceres angular, fused to form branched coralloid et al. 2013) from NE Brazil has a thallus structures up to 150 mm high, made of con- made of ‘‘irregularly densely branched isidi- glutinated hyphae and micareoid photobiont oid granules of c. 50 mm thick’’ but can be (cells c. 6 mm), inspersed with crystals (gyro- distinguished by the absence of any chemical phoric acid), locally some hyphae protrude coumpounds. to form low papillae on the surface. Specimens examined. Re´union: NNW of Piton de Apothecia up to 0 5 mm diam., white, la Fournaise, trail along Ravine Savane Cimetie`re, adnate, slightly constricted at base, without 21 11 60 S, 55 37 50 E, 2050 m, mixed forest with ma- ture Acacia heterophylla,onAcacia, 2008, E. Se´rusiaux any visible margin, convex. Excipulum nar- s. n. (LG), P. van den Boom 39979, 40001 (h); Piton row (<10 mm thick), formed by radiating, de la Fournaise, 0 5kmNWofGıˆte du Volcan, conglutinate hyphae; surface of excipulum 21 12 820 S, 55 41 180 E, 2040 m, Acacia heterophylla sometimes rough because of protruding wood on N-slope, 2008, M. Brand 58353 (h). hyphae. Hypothecium colourless. Hymenium 50–55 mm high. Paraphyses 1 3–1 6 mm Micarea lignaria (Ach.) Hedl. thick, branched and anastomosing. Asci c. 39–55 8–12 mm, tholus of the Micarea- Thallus consisting of small (100 mm across), type. Ascospores 8 per ascus, ellipsoid, 10 3– convex, light grey granules over an algal layer 10 8 3 7–4 1 mm, 0–1-septate. growing on saxicolous mosses. Photobiont mi- Micropycnidia occasionally present, small, careoid, cells round, c.6 5–8 0 mm. white; microconidia 9 8 0 8 mm, narrow Apothecia c. 0 25 mm broad, convex, fusiform. Mesoconidia occasionally present, black, immarginate. Hymenium c. 60 mm, 5 5 1 3 mm. epihymenium blue. Ascospores 8 per ascus, fusiform, c. 20 0 5 2 mm, 5–7-septate. Chemistry. Gyrophoric acid (K-- ,P-- ,C+ red), crystals present only at the centre of Chemistry. Pigment in apothecia belonging the goniocysts. to Cinereorufa-green. 2014 Micarea on Re´union Island—Brand et al. 431
Habitat and distribution. On steep rock face epihymenium colourless to aeruginose. Para- at 2450 m. physes 1 1–1 5 mm thick, branched and anas- tomosing. Asci c. 32–35 12–13 mm; tholus Notes. This collection is very small, but all of the Micarea-type. Ascospores 8 per ascus, characters match those of European collec- oval-oblong to slightly clavate, 8 5–10 5 tions. The thallus is P+ red, but is too small 2 8–4 0 mm, 0–1-septate, septum not in for chemical analysis; the shape of crystals in middle of the spore but somewhat to the the thallus is identical to that of argopsin, the wider end (old spores occasionally 3-septate). lichen substance diagnostic of M. lignaria. Micropycnidia occasionally present, small The ascospores are rather small but fall with- (15–30 mm), immersed, colourless; microco- in the range of European material. The spe- nidia shortly bacilliform, 4 0–4 5 1 0 mm. cies is widespread as, according to Coppins (2009), it occurs in Europe, North and Chemistry. Unknown 1 in thallus (K-- ,P-- , South America, and Asia (Siberia). C-- ). Pigment in thallus and apothecia be- longing to Cinereorufa-green. Specimen examined. Re´union: Cilaos, path Le Bloc 0 to cabane Dufour, Coˆteau de Kerve´guen, 21 6 72 S, Etymology. The name of this new species 55 29 700 E, 2450 m, basalt of W-exposed rock face, 2008, M. Brand 59904 (h). refers to the bluish black prothallus that makes it very characteristic amongst species of the M. prasina group, to which it belongs. Micarea melanoprasina Brand, van den Boom & Se´rus. sp. nov. Habitat and distribution. On trunks of trees, including Acacia heterophylla, and on branches MycoBank No.: MB807678 of shrubs (Gaertnera vaginata, Chassalia cor- Thallus to 5 cm, pale green on black prothallus; alloides, Dombeya sp.) in montane rainforest apothecia 0 2–0 5 mm, dark grey to black; ascospores (1400–1900 m). oval-oblong to slightly clavate, 8 5–10 5 2 8–4 0 mm, 0–1-septate; production of unkown compound. Notes. A distinctive species, easily recog- Type: Re´union, ‘Re´serve naturelle de la Roche Ecrite’, track to the summit, 20 580 600S, 55 260 2600E, nized because of its dark prothallus and finely c. 1500 m, montane forest, on Acacia heterophylla,4 coralloid-goniocystose thallus, very much November 2009, N. Magain & E. Se´rusiaux s. n. (LG— akin to that of M. prasina. Micarea melano- holotype; hb. v.d. Boom, hb. Brand—isotype). prasina is indeed related to the M. prasina (Fig. 3A & B) group and can be distinguished by the pre- sence of Cinereorufa-green in the thallus Thallus to 5 cm across, pale green, made of and apothecia, as well as the unnamed brown coralloid goniocysts growing on bluish black pigment in the apothecia. Micarea subvirides- prothallus; thallus surrounded by bluish black cens (Nyl.) Hedl. has a dark bluish green prothallus, consisting of c. 2 mm thick blue- thallus, made of dense aggregated gonio- black, irregularly intricated hyphae; gonio- cysts, dark apothecia, larger ascospores (10– cysts 14–30 mm, round to ovate, fused to 18 4–6 mm) and contains prasinic acid form branched coralloid structures to 200 (Coppins 2009). mm high, composed of conglutinated hyphae Additional specimens examined. Re´union: Be´louve in a gelatinized matrix and micareoid photo- forest, 21 03 930 S, 55 32 930 E, 1600 m, pristine mon- biont (cells c. 4–5 mm), inspersed with small tane forest, on tree, 27 v 2008, E. Se´rusiaux s. n. (LG); crystals (melting in boiling water forming Bois de sans Souci, 21 1 230 S, 55 22 100 E, 1350 m, oily drops, recrystallizing after cooling). montane forest, on Gaertnera vaginata, 2008, M. Brand 59994 (h); 21 01 220 S, 55 22 230 E, 1380 m, open Apothecia 0 2–0 5mmdiam.,greytoblack, shrub woodland, degraded montane forest, on Chassalia adnate, without visible margin, convex. Excip- corallioides, 2008, M. Brand 60103 (h); E of Le Tampon, ulum colourless or dark blue, formed by radi- ForeˆtdeNotre-DamedelaPaix,21 15 10 S, 55 36 50 E, ating hyphae. Hypothecium pale or more or 1720 m, montane forest, on Dombeya, 2008, P. van den less blue-aeruginose, and brownish (K+ pur- Boom 40775 (h); Cirque de Cilaos, Foreˆt du Grand Matarum, along path to Cabane Dufour, 21 07 040 S, plish brown) in lower part. Hymenium 33–37 55 29 230 E, 1870 m, montane forest, on Dombeya, mm high, more or less aeruginose near base; 2008, M. Brand 59740 (h). 432 THE LICHENOLOGIST Vol. 46
Fig. 3. Photographs of the new species of Micarea described from Re´union. A & B, M. melanoprasina sp. nov. (hol- otype); C, M. pseudolignaria sp. nov. (holotype); D, M. sublithinella sp. nov. (holotype); E, M. takamakae sp. nov. (holotype); F, M. tenuispora sp. nov. (holotype). Scale ¼ 1 mm. In colour online.
Micarea peliocarpa (Anzi) Coppins & part c. 15 mm without algae, of loosely inter- R. Sant. woven hyphae. Photobiont micareoid, photo- biont layer inspersed with crystals (gyrophoric Thallus rather small, up to 1 5 cm across, acid). pale greenish, uneven, c. 50 mm high. Upper 2014 Micarea on Re´union Island—Brand et al. 433
Apothecia up to 0 6 mm diam., whitish or Specimens examined. Re´union: St-Paul, road to Piton dark bluish grey, often patchily coloured, Maı¨do, 21 03 240 S, 55 21 390 E, 1500 m, edge of open woodland, on large Acacia heterophylla, 2008, M. Brand with pale margin, flat or low convex, almost 59488 (h); Cilaos, Plateau du Petit Matarum, path from globose, adnate with constricted base. Outer Le Bloc to Cabane Dufour, 21 06 960 S, 55 29 220 E, parts of excipulum pale or bluish grey, pale 1900 m, montane forest, on Acacia heterophylla, 2008, inside. Hypothecium colourless. Hymenium c. M. Brand 59756 (h); ibid., 21 07 30 S, 55 29 20 E, 1420 40–60 mm high; epihymenium colourless or m, montane forest, on tree, 2008, P. van den Boom 40282 (h); Piton de la Fournaise, 0 5kmNWofGıˆte du patchily blue-grey, if pigmented then pig- Volcan, 21 12 80 S, 55 41 180 E, 2040 m, on Acacia het- ment also descending in hymenium. Para- erophylla in wood on N-slope, 2008, M. Brand 58356a, physes 1 1–1 4 mm thick, branched and anas- 58356b (h); ForeˆtdeBe´bour, path Be´louve to cabane 0 tomosing. Asci clavate, c. 35–55 12–15 mm, Dufour, 3 km from Gıˆte Be´louve, 21 05 10 S, 55 31 360 E, 2040 m, wet high ericoid thickets, on Erica, tholus of the Micarea-type. Ascospores 8 per 2008, M. Brand 58864 (h). ascus, fusiform with rounded ends, 3-septate, 13 6–17 0 3 6–4 7 mm. Micarea prasina Fr. Macropycnidia not rare, globose, c. 120– 150 mm, top blue-grey or not; macroconidia Thallus diffuse, c. 1 cm across, green, thin curved (but not spirally), 45–58 1 1–1 3 (50 mm), granular, made of goniocysts 30– mm, indistinctly 3-septate. Micropycnidia rare, 40 mm diam., round, dark greyish green; inconspicuous, semi-superficial, c. 45 mm, goniocysts composed of thin hyphae inspersed pale. Microconidia narrowly fusiform with with small crystals (melting in boiling water). tapering ends, 6 3–8 1 0 7–0 9 mm. Photobiont micareoid, cells c. 4–6 mm. Apothecia up to 0.3 mm diam., dark grey Chemistry. Gyrophoric and methylhiascic to dull black, convex, adnate with narrow acids (K-- ,P-- , C+ red), mainly in thallus, base, without any visible margin. Excipulum hymenium, excipulum and pycnidia. Pig- in lower part of apothecia c. 40 mmwide, ment in thallus and apothecia belonging to formed of radiating hyphae. Hypothecium dark Cinereorufa-green. grey-brown (K+ violet). Hymenium to 50 mm high. Paraphyses c. 1 2–1 3 mm thick, Habitat and distribution. Mostly on soft bark branched and anastomosing. Asci c. 40– of shaded Acacia heterophylla trees, at 1500– 42 9–12 mm, tholus of the Micarea-type. 2000 m; also found on Erica, and sometimes Ascospores 8 per ascus, ovoid-ellipsoid, 9 3– overgrowing mosses. 12 8 3 8–4 3 mm, (0–)1-septate. Notes. Within the speciose Micarea pelio- Micropycnidia sparse, white or grey; micro- carpa group in Re´union, we recognize M. conidia 6 3–7 1 0 9–1 1 mm, narrow fusi- borbonica sp. nov. with 7-septate fusiform as- form, straight or slightly curved. cospores, and thus close to M. cinerea (see Chemistry. Micareic acid. With Sedifolia- further comments under M. borbonica), M. grey pigment in inner excipulum, hypothe- boryana sp. nov. with 3-septate narrowly clav- cium and in diffuse streaks in hymenium. ate ascospores (see under that species) and also M. peliocarpa s. str. which encompasses Habitat and distribution. On bark of Acacia collections with consistently 3-septate, fusi- heterophylla, intermixed with the related M. form ascospores. The size of ascospores falls levicula, and so far only found in natural in the range of European populations as pro- stands of Acacia heterophylla near the Piton vided by Coppins [2009: (11–)15–23(–24) de la Fournaise. 3–5(–6) mm]. However, in the Re´union populations the ascospores are on average Notes. Our material differs from Micarea wider than in Europe: 13 6–17 0 4 1–4 7 prasina, as it is now circumscribed in Europe mminRe´union vs. 12 2–22 5 3 5–4 2 mm (Coppins 2009), by the location of the K+ in Europe (unpublished results, based on violet pigment in the hypothecium instead studies of 35 collections). of the epihymenium. Otherwise the thallus 434 THE LICHENOLOGIST Vol. 46 made of finely dissected and coralloid gonio- grey, pale inside, partly incrusted with gyro- cysts, and the production of micareic acid, phoric acid crystals. Hypothecium colourless. are diagnostic for M. prasina. We refrain Hymenium c. 50–55 mm high; epihymenium from describing a new species on that sole colourless or blue-grey in patches. Paraphyses basis. A K+ violet hypothecium is otherwise 1 7–1 9 mm thick, simple or branched, only known from M. endoviolascens Coppins strongly conglutinate. Asci clavate, for from South Africa (Coppins 1999), easily example 32–47 11–16 mm, tholus of the distinguished by its corticated squamules, Micarea-type. Ascospores 8 per ascus, bacillar- and M. hypoviolascens Czarnota & Coppins ellipsoid, 11 5–14 7 4 2–5 7 mm, (0–)3- from a single locality in W Scotland (Czar- septate, straight to curved. nota & Coppins 2005), distinguished by its Pycnidia not found. areolate thallus and production of an un- known substance. Chemistry. Gyrophoric acid (K-- ,P-- ,C+ red), mainly in thallus and soredia, some- Specimens examined. Re´union: NNW of Piton de la Fournaise, trail along Ravine Savane Cimetie`re, times in excipulum. Pigment in apothecia 21 11 60 S, 55 37 50 E, 2050 m, mixed forest with belonging to Cinereorufa-green. mature Acacia heterophylla,onAcacia, 2008, P. van den Boom 39976 (h); Piton de la Fournaise, c.0 5kmNW Etymology. The name refers to another of Gıˆte du Volcan, 21 12 820 S, 55 41 180 E, 2040 m, on Acacia heterophylla in wood on N-slope, 2008, M. species of Micarea, M. coppinsii, which was Brand 58355 (h). dedicated to our talented colleague and friend, Dr Brian J. Coppins.
Micarea pseudocoppinsii Brand, van Habitat and distribution. On soft bark of den Boom & Se´rus. sp. nov. Acacia heterophylla or overgrowing mosses, MycoBank No.: MB807679 between 1500–2050 m elevation. Thallus diffuse, small, pale greenish, with convex soralia; apothecia to 0 4 mm, white, pale orange or partly bluish Notes. Genuine soralia are rare in Micarea. grey, convex; ascospores bacillar-ellipsoid, 11 5–13 0 Beside M. pseudocoppinsii, they are only known 4 2–5 7 mm, (0–)3-septate; production of gyrophoric in two other species: M. alectorialica, here acid. described as new and easily identified by its Type: Re´union, NNW of Piton de la Fournaise, trail along Ravine Savane Cimetie`re, 21 12 060 S, black apothecia and production of alector- 55 41 460 E, 2050 m, mixed forest with mature Acacia ialic acid, and M. coppinsii Tønsberg, also heterophylla,onAcacia, 28 May 2008, P. van den Boom producing gyrophoric acid. The latter species 39991 (LG—holotype; hb. v.d. Boom, hb. Brand differs in producing methylhiascic acid to- 61466—isotype). gether with gyrophoric acid, and has fusi- (Fig. 2H) form, sometimes slightly clavate, longer as- cospores (20–28 4–5 mm)(Coppins 2009). Thallus diffuse, small (1 5 cm across) pale Micarea viridileprosa Coppins & van den Boom greenish, made of small granules or gonio- has a bright, green, leproid thallus without cysts, locally with soralia; goniocysts 18–48 clearly delimited soralia and has smaller asco- mm diam., round to ovoid, partly coalescing spores [8–12(–14) 2 5–4 0 mm]. Micarea but not coralloid. Photobiont micareoid, c. pseudocoppinsii is most likely related to the 5–8 mm. Soralia sometimes numerous, round, group of M. peliocarpa as it has the same convex, 0 3–0 6 mm broad; soredia light type of apothecia, and to M. peliocarpa which green, very fine, 11–32 mm diam. differs by its less-developed thallus and lack Apothecia up to 0 4 mm diam., white, pale of soralia. orange or partly bluish grey, convex, adnate with constricted base, without any visible Additional specimen examined. Re´union: Foreˆtde Be´bour, near Gıˆte de Be´louve, 21 03 740 S, 55 32 310 E, margin. Excipulum formed by radiating, con- 1520 m, old plantations of Acacia heterophylla trees, on glutinate hyphae, outer parts pale or bluish Acacia, 2008, P. van den Boom 39861(h). 2014 Micarea on Re´union Island—Brand et al. 435
Micarea pseudolignaria Brand, van den Notes. This new species looks very much Boom & Se´rus. sp. nov. like M. lignaria (Ach.) Hedl., also found in a single locality in Re´union, but has smaller 1- MycoBank No.: MB807680 septate ascospores and a different chemistry. Thallus muscicolous, areolate, greyish green; apothecia In M. lignaria, ascospores are 3–7-septate, to 0 6 mm, shining black, convex; ascospores ellipsoid, 8 0–9 2 3 0–3 3 mm, (0–)1-septate; production of 16–28(–38) 4 3–5 7(–7 0) mm, and the unknown substance. thallus produces argopsin (var. lignaria)or Type: Re´union, ForeˆtdeBe´bour, path Be´louve to unidentified xanthones (var. endoleuca). Fur- cabane Dufour, 21 04 770 S, 55 31 640 E, 1740 m, soil thermore, M. lignaria has a greenish hue in in ericoid thickets, 2 June 2008, M. Brand 58846 (LG—holotype; hb. v.d. Boom, hb. Brand—isotype). the lower part of the hymenium, which is never inspersed. The rare M. ternaria (Nyl.) (Fig. 3C) Veˇzda, known only from the Northern Hemi- sphere, has a similar thallus and apothecia Thallus c. 2 cm across, overgrowing decay- but differs by the lack of lichen substances ing bryophytes, uneven, made of coalescent, and (1–)3-septate, 13–22 3 5–5 0 mmasco- 0 2–0 3 mm diam., slightly convex, greyish spores (Coppins 2009). greenish areolae. Photobiont micareoid, cells c. 5 mm. Cortex absent but areolae partly cov- ered by a hyaline epinecral layer. Thallus Micarea sublithinella Brand, van den without crystals, except for lower part of pho- Boom & Se´rus. sp. nov. tobiont layer more or less opaque due to oily drops and tiny crystals. MycoBank No.: MB807681 Apothecia up to 0 6 mm diam., shining Thallus thin, with flattened rounded areoles, green; black, convex without any distinct margin, apothecia up to 0 4 mm diam.; ascospores obovoid, 12 5–15 0 5 0–5 8 mm, 0–1-septate; production of often coalescing to form compound apothe- protolichesterinic acid. cia. Excipulum with radiating conglutinated Type: Re´union, NNW of Piton de la Fournaise, hyphae, outer parts brownish to bluish, col- trail along Ravine Savane Cimetie`re, 21 12 060 S, 0 ourless inside. Hypothecium colourless. Hy- 55 41 46 E, 2050 m, mixed forest with mature Acacia heterophylla,onAcacia, 28 May 2008, P. van den Boom menium c. 40 mm high, lower part more or 39980 (LG—holotype; hb. v.d. Boom, hb. Brand— less inspersed; epihymenium blue. Paraphyses isotype). 1 5 mm thick, strongly branched and anasto- mosing. Asci clavate, c. 35 9 mm, tholus of (Fig. 3D) the Micarea-type. Ascospores 8 per ascus, ellip- Thallus c. 2 cm across, thin (25–50 mm), soid, 8 0–9 2 3 0–3 3 mm, (0–)1-septate, green, continuous or consisting of flat wall c. 0 3 mm thick. rounded areoles growing over a thin, slightly Micropycnidia superficial (i.e. on thin thal- shiny film over the substratum. Photobiont lus overgrowing leafy liverworts), c. 75 mm micareoid, cells c. 5–6 mm, thallus with a diam., wall dark from bluish pigment. Micro- few crystals. conidia bacillar, 4 6–5 8 0 9–1 0 mm. Apothecia numerous and small, up to 0 4 Chemistry. Unknown 3 in thallus (K-- ,P-- , mm diam. (deformed apothecia up to 0 5 C-- ), most probably present as tiny oil drop- mm), dull, light brownish, broadly adnate, lets or crystals. Pigment in apothecia belong- immarginate, convex to semiglobose. Excip- ing to Cinereorufa-green. ulum thin (less than 10 mm). Hypothecium pale. Hymenium c. 55–75 mm high; epihyme- Etymology. The name refers to the wide- nium colourless or faintly brownish, with spread M. lignaria, which could be easily con- crystals in vertical streaks. Paraphyses 1 0 fused with this new species. mm thick, branched. Asci c. 50–70 15–16 mm, tholus of the Micarea-type. Ascospores Habitat and distribution. Only known from 8 per ascus, obovoid (apical part slightly the type locality. On soil, among rich stands broader), 12 5–15 0 5 0–5 8 mm, 0–1- of Cladonia, in montane ericoid shrubs. septate, wall thin (c. 0 2 mm). 436 THE LICHENOLOGIST Vol. 46
Pycnidia not found. the bark or, when superficial, goniocysts covered by clear necrotic tissue. Photobiont -- -- Chemistry. Protolichesterinic acid (K ,P , micareoid, cells c. 5–7 mm. Crystals detected -- C ), with crystals mainly detected in epihy- in goniocysts. menium, but also present in thallus. Apothecia very numerous and small (up Etymology. The name refers to M. lithi- to 0 2 mm diam.), black, adnate with con- nella, a widespread species that could be stricted base to nearly stipitate, first flat easily confused with this new species. without any distinct margin, then convex, often forming clusters. Excipulum well devel- Habitat and distribution. On wood of rotting oped in under part, made of chondroid tis- trunks, or on trees, including Acacia hetero- sue; outer part hyaline, inner part locally phylla, in montane forests. brown. Hypothecium pale, opaque because of Notes. Most characters of Micarea sublithi- numerous small crystals. Subhymenium red- nella fit the the traditional concept of the dish brown. Hymenium c. 35 mm high; epihy- genus Micarea (sensu Coppins 1983), but the menium dull blue-green and brownish, with larger asci and production of fatty acid set it small dark granules in K. Paraphyses 1 4 mm apart. In the field it can easily be confused thick, branched. Asci c. 27–30 11–13 mm, with M. lithinella (Nyl.) Hedl., a widespread tholus of the Micarea-type. Ascospores 8 per species in the Northern Hemisphere, but this ascus, ellipsoid, 8 8–10 2 3 1–3 8 mm, 0– species is a pioneer saxicolous species in dis- 1-septate, wall c. 0 3 mm thick. Pycnidia not found. turbed habitats, has smaller asci (35–50 8– 13 mm), smaller and simple ascospores (6 5– Chemistry. Unknown 2, assumed to be 9 5 2 5–4 0 mm) and no compounds de- confluentic acid or a related compound (K-- , tected (Coppins 2009). Species related to P-- ,C-- ). Pigment in epihymenium belonging M. lithinella, such as M. farinosa Coppins & to Cinereorufa-green. Aptroot, also lack compounds. Additional specimens examined. Re´union: same locality Etymology. The name refers to a spot on as the type, E. Se´rusiaux s. n. (LG); ForeˆtdeBe´bour, the northern flank of the island, ‘Le Casse´ trail to Takamaka, 21 05 90 S, 55 34 30 E, 1370 m, wet de Takamaka’, a landscape of dramatic montane forest, on trees, 2008, P. van den Boom 40872 beauty. (h); E of Le Tampon, Foreˆt de Notre-Dame de la Paix, 21 15 10 S, 55 36 50 E, 1720 m, montane forest, on tree, 2008, P. van den Boom 40745 (h). Habitat and distribution. Only known from the type locality, at rather low elevation (620 Micarea takamakae Brand, van den m) in a high rainfall area, on decaying wood Boom & Se´rus. sp. nov. of standing trunk. MycoBank No.: MB807682 Notes. Micarea takamakae has an excipu- Thallus semi-endoxylic, made of goniocysts, whitish; lum and hymenium very much akin to those apothecia up to 0 2 mm diam., adnate; ascospores ellip- of M. erratica, but all other characters are quite soid, 8 8–10 2 3 1–3 8 mm, 0–1-septate; production different and set it apart from that species. of unknown substance. Confluentic acid has never been reported in Type: La Re´union, WSW of St-Benoıˆt, along the road to Takamaka, c. 3 km SW of Abondance les Hauts, the genus but all chemical features point to 21 03 50 S, 55 38 50 E, 620 m, disturbed area with that substance. Small ascospores are a fur- planted trees, on decaying tree, 11 June 2008, P. van ther interesting feature of this new species, den Boom 41009 (LG—holotype; hb. v.d. Boom, hb. amongst the species found on Re´union and Brand—isotype). not belonging to the M. micrococca-prasina (Fig. 3E) aggregate.
Thallus inconspicuous, diffuse, partly en- Micarea tenuispora Brand, van den doxylic on soft, decaying wood, whitish, Boom & Se´rus. sp. nov. made of dispersed, round goniocysts, c. 30– 75 mm diam., either in empty dead cells of MycoBank No.: MB807683 2014 Micarea on Re´union Island—Brand et al. 437
Thallus very thin to endophloeodal, isidiate or made of Etymology. The name was chosen to em- isidiiform areolae, isidia small cylindrical to coralloid, phasize the thin ascospores of this species. up to 0 3 mm high; apothecia to 0 9 mm, dark grey to black, ascospores acicular, narrowly clavate, 31–35 2 2–2 5 mm, 3–5-septate; production of gyrophoric acid. Habitat and distribution. On bark of the Type: Re´union, Cirque de Cilaos, Foreˆt du Grand exotic Cryptomeria japonica, between 1200– Matarum, trail to Caverne Dufour, 21 07 30 S, 1500 m elevation, at edges of disturbed natu- 55 29 20 E, 1420 m, montane forest, on Cryptomeria,31 ral forests. May 2008, P. van den Boom 40231 (LG—holotype; hb. v.d. Boom, hb. Brand—isotype). Notes. Micarea tenuispora is easily distin- (Fig. 3F) guished by its ‘isidiate’ thallus and 3–5- septate ascospores. Isidia or isidiiform thal- Thallus to 5 cm across, pale greenish lus areolae are reported in only two species to brownish, discontinuous, thin (c. 50 mm of Micarea, both described here as new. For thick), consisting of scattered areolae, soon differences to M. isidiosa sp. nov., see under forming isidia or becoming isidiiform, on that species. thin endo- to epiphloeodal dark bluish hypo- thallus with narrow, colourless or locally dark Additional specimen examined. Re´union: Cilaos, Foreˆt du Grand Matarum, 21 07 170 S, 55 29 100 E, 1450 m, brownish or bluish hyphae; mature thallus disturbed margin of montane forest, on Cryptomeria, always with isidiiform areolae (c.0 2mm 2008, M. Brand 58685 (h). high, 0 05–0 07 mm thick). Cortex absent, We are greatly honoured to publish our paper in this often with thin epinecral layer. Photobiont special issue of The Lichenologist for the 65th birthday of micareoid. Thallus filled with crystals of our most distinguished colleague and friend, and the gyrophoric acid. world expert on Micarea taxonomy, Dr Brian J. Coppins. Apothecia up to 0 8 mm diam., sessile to Field studies in Re´union were made possible with the substipitate, black, usually with a pale margin help and advice from the Parc National de La Re´union, when young, first flat, without distinct rim, especially through the courtesy of Mr B. Lequette and then usually but not always semiglobose. Mr J. M. Pause´. Dr Cl. Ah-Peng and Prof. D. Strasberg of the University of La Re´union in Saint-Denis were also Excipulum with radiating hyphae, varying in very helpful. We thank them all most sincerely. We colour (often in the same apothecium) from further thank the Senior Editor of The Lichenologist, pale (then with crystals) to superficially bluish Prof. P. D. Crittenden, and the three Editors of this and centrally light brown. Hypothecium pale special volume (Dr Alan Fryday, Dr Gintaras Kantvilas brown to dark bluish. Hymenium c. 55 mm and Dr Chris Ellis). Both referees contributed to the final version of this manuscript as their notes and suggestions high; epihymenium greyish bluish in patches. were very helpful: we thank them warmly. Paraphyses c. 1 3 mm thick, branched and anastomosing. Asci c. 44 11 mm, tholus of the Micarea-type. Ascospores 8 per ascus, in References one bundle, sometimes twisted, acicular, Andersen, H. L. & Ekman, S. (2005) Disintegration apically inflated and tapering towards the of the Micareaceae (lichenized Ascomycota): a molecular phylogeny based on mitochondrial rDNA base, 31–35 2 2–2 5 mm, 3–5-septate. sequences. Mycological Research 109: 21–30. Micropycnidia scarce, inconspicuous, ses- Aptroot, A. (2002) New and interesting lichens and sile, c. 110 mm, with blue-green top, brown- lichenicolous fungi in Brazil. Fungal Diversity 9: 15– ish wall. Microconidia narrow fusiform, aver- 45. Baret, S., Rouget, M., Richardson, D. M., Lavergne, C., age 3 8–4 1 1 1–1 2 mm. Egoh, D., Dupont, J. & Strasberg, D. (2006) Cur- rent distribution and potential extent of the most in- vasive alien species on La Re´union (Indian Ocean, Mascarene Islands). Austral Ecology 31: 747–758. Chemistry. Gyrophoric acid (K-- ,P-- ,C+ Bass, D. & Richards, T. A. (2011) Three reasons to re- red), mainly in thallus, but also in pale parts evaluate fungal diversity ‘on Earth and in the ocean’. of apothecia and in pycnidia. Pigment in thal- Fungal Biology Reviews 25: 129–164. Blackwell, M. (2011) The fungi: 1, 2, 3 . . . 5.1 million lus and apothecia belonging to Cinereorufa- species? American Journal of Botany 98: 426–438. green. Brown pigment in hypothecium K+ Ca´ceres, M. E. S., Mota, D. A., de Jesus, L. S. & Aptroot, reddish to purple. A. (2013) The new lichen species Micarea corallo- 438 THE LICHENOLOGIST Vol. 46
thallina from Serra da Jibo´ia, an Atlantic rainforest Islands, New Zealand. Bibliotheca Lichenologica 88: enclave in Bahia, NE Brazil. Lichenologist 45: 371– 127–146. 373. Gaston, K. J. (2000) Global patterns in biodiversity. Cadet, T. (1977) La ve´ge´tation de l0 Ile de La Re´union: e´tude Nature 405: 220–227. phytoe´cologique et phytosociologique. Ph.D. thesis, Uni- Harris, R. C. (2009) Four novel lichen taxa in the lichen versite´ d’Aix-Marseille. biota of eastern North America. Opuscula Philoliche- Camargo, A., Avila, L. J., Morando, M. & Sites, J. W. num 6: 149–156. (2012a) Accuracy and precision of species trees: Hawksworth, D. L. (2012) Global species numbers of effects of locus, individual, and base pair sampling fungi: are tropical studies and molecular approaches on inference of species trees in lizards of the Liolaemus contributing to a more robust estimate? Biodiversity darwinii group (Squamata, Liolaemidae). Systematic and Conservation 21: 2425–2433. Biology 61: 272–288. Honegger, R. (1993) Developmental biology of lichens. Camargo, A., Morando, M., Avila, L. J. & Sites, J. W. Tansley Review No. 60. New Phytologist 125: 659– (2012b) Species delimitation with ABC and other 677. coalescent-based methods: a test of accuracy with Huneck, S. & Yoshimura, I. (1996) Identification of simulations and an empirical example with lizards Lichen Substances. Berlin, Heidelberg: Springer- of the Liolaemus darwinii complex (Squamata: Lio- Verlag. laemidae). Evolution 66: 2834–2849. Kalb, K. (2007) New or interesting lichens. III. Biblio- Carstens, B. C., Pelletier, T. A., Reid, N. M. & Satler, theca Lichenologica 95: 297–316. J. D. (2013) How to fail at species delimitation. Kantvilas, G. & Elix, J. A. (1994) Ramboldia, a new Molecular Ecology 22: 4369–4383. genus in the lichen family Lecanoraceae. Bryologist Coppins, B. J. (1983) A taxonomic study of the lichen 97: 296–304. genus Micarea in Europe. Bulletin of the British Knowles, L. L. & Kubatko, L. S. (2010). Estimating Museum´ (Natural History), Botany Series 11(2): 17– species trees: an introduction to concepts and 214. models. In Estimating Species Trees. Practical and Coppins, B. J. (1999) Two new species of Micarea from Theoretical Aspects (L. L. Knowles & L. S. Kutbako, South Africa. Lichenologist 31: 559–565. eds): 1–14. Hoboken: Wiley-Blackwell. Coppins, B. J. (2009) Micarea.InThe Lichens of Great Lacoste, M., Delbosc, P. & Picot, F. (2011) Typologie Britain and Ireland. (C. W. Smith, A. Aptroot, B. J. Descriptive des Habitats Naturels et Semi-naturels de La Coppins, A. Fletcher, O. L. Gilbert, P. W. James & Re´union, Version De´cembre 2011. Rapport technique P. A. Wolseley, eds): 583–606. London: British no 8. Saint-Leu, Re´union: Conservatoire Botanique Lichen Society. de Mascarin. Coppins, B. J. & Kantvilas, G. (1990) Studies on Micarea Lagabrielle, E., Rouget, M., Payet, K., Wistebaar, N., in Australasia I. Four new species from Tasmania. Durieux, L., Baret, S., Lombard, A. & Strasberg, Lichenologist 22: 277–288. G. (2009) Identifying and mapping biodiversity pro- Coppins, B. J. & Tønsberg, T. (2001) A new xanthone- cesses for conservation planning in islands: a case containing Micarea from Northwest Europe and the study in Re´union Island (Western Indian Ocean). Pacific Northwest of North America. Lichenologist Biological Conservation 142: 1523–1535. 33: 93–96. Lu¨cking, R., Rivas Plata, E., Chaves, J. L., Uman˜a, L. & Czarnota, P. (2007) The Lichen Genus Micarea (Lecanor- Sipman, H. J. M. (2009) How many tropical lichens ales, Ascomycota) in Poland. Polish Botanical Studies are there really? Bibliotheca Lichenologica 100: 399– No. 23.Krako´w: W. Szafer Institute of Botany, Polish 418. Academy of Sciences. Maddison, W. P. (1997) Gene trees in species trees. Czarnota, P. & Coppins, B. J. (2005) A second Micarea Systematic Biology 46: 523–536. with a hypothecial K+ violet pigment. Lichenologist Meyer, B. & Printzen, C. (2000) Proposal for a standar- 37: 477–479. dized nomenclature and characterization of insolu- Czarnota, P. & Guzow-Krzemin´ska, B. (2010) A phylo- ble lichen pigments. Lichenologist 32: 571–583. genetic study of the Micarea prasina group shows Orange, A., James, P. W. & White, F. J. (2010) Micro- that Micarea micrococca includes three distinct line- chemical Methods for the Identification of Lichens. ages. Lichenologist 42: 7–21. London: British Lichen Society. Edwards, S. V. (2008) Is a new and general theory of Purvis, O. W. & James, P. W. (1993) Studies on the molecular systematics emerging? Evolution 63: 1–19. lichens of the Azores. Part 1. Caldeira Faial. Arqui- Ekman, S., Andersen, H. L. & Wedin, M. (2008) The pe´lago, Life and Marine Sciences 11A: 1–15. limitations of ancestral state reconstruction and the Romeike, J., Friedl, T., Helms, G. & Ott, S. (2002) evolution of the ascus in the Lecanorales (Lichenized Genetic diversity of algal and fungal partners in Ascomycota). Systematic Biology 57: 141–156. four species of Umbilicaria (lichenized Ascomy- Elix, J. A., Jones, A. J., Lajide, L., Coppins, B. J. & James, cetes) along a transect of the Antarctic Peninsula. P. W. (1984) Two new diphenyl ethers and a new Molecular Biology and Evolution 19: 1209–1217. depside from the lichen Micarea prasina Fr. Austra- Schmull, M., Mia˛dlikowska, J., Pelzer, M., Stocker- lian Journal of Chemistry 37: 2349–2364. Wo¨rgo¨tter, E., Hofstetter, V., Fraker, E., Hodkin- Fryday, A. M. (2004) New species and records of liche- son, B. P., Reeb, V., Kukwa, M., Lumbsch, H. T., nized fungi from Campbell Island and the Auckland et al. (2011) Phylogenetic affiliations of members of 2014 Micarea on Re´union Island—Brand et al. 439
the heterogeneous lichen-forming fungi of the genus Suija, A., Lo˜hmus, P. & Motieju#naite˙, J. (2008) New Lecidea sensu Zahlbruckner (Lecanoromycetes, As- Estonian records. Lichens and lichenicolous fungi. comycota). Mycologia 103: 983–1003. Folia Cryptogamica Estonica 44: 156–159. Se´rusiaux, E., Brand, A. M., Motieju#naite˙, J., Orange, van den Boom, P. P. G. & Coppins, B. J. (2001) Micarea A. & Coppins, B. J. (2010) Lecidea doliiformis be- viridileprosa sp. nov., an overlooked lichen species longs to Micarea, Catillaria alba to Biatora, and Bia- from Western Europe. Lichenologist 33: 87–91. tora ligni-mollis occurs in western Europe. Bryologist van den Boom, P. P. G., Brand, M., Ertz, D., Kalb, K., 113: 333–344. Magain, N., Masson, D., Schiefelbein, U., Sipman, Strasberg, D., Rouget, M., Richardson, D. M., Baret, H. J. M. & Se´rusiaux, E. (2011) Discovering the S., Dupont, J. & Cowling, R. M. (2005) An assess- lichen diversity of a remote tropical island: working ment of habitat diversity and transformation on La list of species collected on Re´union (Mascarene Re´union Island (Mascarene Islands, Indian Ocean) archipelago, Indian Ocean). Herzogia 24: 325–349. as a basis for identifying broad-scale conservation priorities. Biodiversity and Conservation 14: 3015– 3032.