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Micarea Byssacea New to North America and Micarea Hedlundii New to Maine, Michigan and Quebec

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Micarea Byssacea New to North America and Micarea Hedlundii New to Maine, Michigan and Quebec Opuscula Philolichenum, 13: 84-90. 2014. *pdf effectively published online 25June2014 via (http://sweetgum.nybg.org/philolichenum/) Micarea byssacea new to North America and Micarea hedlundii new to Maine, Michigan and Quebec 1 2 ANNINA LAUNIS & LEENA MYLLYS ABSTRACT. – Micarea byssacea is reported new to North America from the coastal region of Maine. Micarea hedlundii is reported new to the states of Maine and Michigan (U.S.A.) and the province of Quebec (Canada). Micarea hedlundii was previously reported for North America only from New Brunswick, Canada and California, U.S.A. Both species have likely been overlooked in North America owing to their inconspicuous thalli. Further studies are needed to fully understand the ecology and distribution of M. hedlundii, M. byssacea and allied species in North America. KEYWORDS. – New records, biogeography, ecology, taxonomy, crustose lichens. INTRODUCTION Micarea Fr. is a crustose lichen genus (Lecanoromycetes, Ascomycota) containing almost 100 species worldwide (Coppins 2009, Czarnota 2007). However, phylogenetic analyses based on molecular data clearly show that the genus is paraphyletic (Andersen & Ekman 2005, Serusiaux et al. 2010), even after the introduction of a new genus Brianaria Ekman & Svensson for the M. sylvicola group (Ekman & Svensson 2014). The M. prasina group, which includes the type species of the genus, M. prasina Fr., currently consists of 17 species (Czarnota & Guzow-Krzemińska 2010), which all have a “micareoid” photobiont (a coccoid green alga with cells 4–7 µm in diameter), immarginate apothecia, branched paraphyses and an ascus of the Micarea-type (Hafellner 1984). All the species in this group occur on bark, especially of old trees or on soft lignum. The majority of the species produce the Sedifolia-grey pigment (K+ violet, C+ violet), which is typically present in the apothecia and pycnidia (Coppins 1983, Czarnota & Guzow-Krzemińska 2010). The taxonomy of Micarea is incompletely known and many species are still undercollected, probably owing to their small size and inconspicuous thalli. Micarea prasina has been divided into several species during the last two decades, including the M. micrococca complex that is composed of M. byssacea (Th. Fr.) Czarnota, Guzow- Krzemińska & Coppins, M. micrococca (Körb.) Gams ex D. Hawskw. s. str. and a yet unnamed taxon (Czarnota & Guzow- Krzemińska 2010). According to the 19th edition of the North American lichen checklist (Esslinger 2014), 40 Micarea species are known from North America, of which eight belong to the M. prasina group. Here we present the first records of M. byssacea, for North America and of M. hedlundii Coppins for the states of Maine and Michigan (U.S.A.) and the province of Quebec (Canada). The recent molecular study by Czarnota and Guzow-Krzemińska (2010) confirms that both species belong to the M. prasina group. MATERIALS AND METHODS Micarea specimens were collected by the first author during the Crustose Lichens and Sterile Crustose Lichens workshops at the Eagle Hill Institute, Maine, in June 2012. Specimens were identified 1 ANNINA LAUNIS – Botany Unit, Finnish Museum of Natural History, P.O. Box 7, FI-00014 University of Helsinki, Finland. – e-mail: [email protected] 2 LEENA MYLLYS – Botany Unit, Finnish Museum of Natural History, P.O. Box 7, FI-00014 University of Helsinki, Finland. – e-mail: [email protected] 84 with a dissecting or compound microscope. Ascospore dimensions and other anatomical measurements were made in water and are presented based on the cited collections and the measurements of Czarnota and Guzow-Krzemińska (2010). Chemical spot tests were performed under a compound microscope using sodium hypochlorite (C) and 10% potassium hydroxide (K) (Orange et al. 2001). Pigments were defined following Coppins (1983), Meyer and Printzen (2000) and Czarnota (2007). Specimens were further studied using thin-layer chromatography following Culberson and Kristinsson (1970) and Orange et al. (2001). Specimens are deposited in the Finnish Museum of Natural History (H) and the New York Botanical Garden (NY). THE SPECIES Micarea byssacea (Th. Fr.) Czarnota, Guzow- Krzemińska & Coppins FIGURE 1B. CHARACTERIZATION OF THE SPECIMENS EXAMINED. – Thallus effuse, green to olive-green and minutely granular, composed of small goniocysts. Photobiont micareoid (cells ± globose, 4–7 µm). Apothecia variable in size and color even in the same collection, usually numerous, immarginate and mostly adnate, sometimes convex to hemispherical, 0.2–0.6 mm in diameter, whitish, cream, whitish-grey, olive-grey, or grey to blackish grey, often with paler outer part. Hymenium hyaline, but in darker apothecia the epihymenium is greyish, K± violet, C± reddish violet due to the presence of Sedifolia-grey which is confined to the gel matrix. Hypothecium hyaline or slightly yellowish. Ascus clavate or cylindrical-clavate. Ascospores oblong, oblong-ovoid, ellipsoid, 0(–1) septate, (6–)8–12(–13) x 2.7–3.5(–4.2) µm. Pycnidia, not always present, usually immersed to sessile and white to greyish-white (K± violet, C± reddish violet). In North American material, however, pycnidia are sometimes emergent and dark grey. Mesoconidia (3.8– )4.5–5.5 x 1.2–1.5 µm and microconidia 5–7.5(–8) x 0.8–1 µm. CHEMISTRY. – Methoxymicareic acid detected by TLC and the Sedifolia-grey pigment detected by chemical spot tests (K and C). DISTRIBUTION AND ECOLOGY. – Micarea byssacea was previously known only from Central and Northern Europe. It is likely widespread but overlooked in North America, often reported as M. prasina or M. micrococca (sensu Coppins) (see below for notes). In eastern Maine, M. byssacea was collected from two localities in Washington County: from Great Wass Island Reserve in the Town of Beals and from Eagle Hill in the Town of Steuben. Collections are both from humid natural and semi-natural coastal forests. In Europe, Micarea byssacea is a forest epiphyte occurring on the bark of several tree species such as Pinus spp., Picea spp. and Betula spp. It usually occurs on trees with a low bark pH, although one collection from North America is from Acer, which sometimes has less acid bark. On rather rare occasions, the species has been found on more or less hard lignum of decaying stumps of coniferous trees. Almost all known collections of M. byssacea are from natural or semi-natural forests. The species obviously thrives in shaded habitats with humid microclimates. However, the distribution and ecological preferences of M. byssacea are still incompletely known both in North America and Europe. NOTES. – Coppins (1983) considered Micarea prasina to be a variable species needing taxonomic revision and it has subsequently been divided into several species based on differences in morphology, chemistry and/or DNA characters (e.g. Coppins & Tønsberg 2001, van den Boom & Coppins 2001). Micarea micrococca was separated from M. prasina, which contains micareic acid, mainly by the presence of methoxymicareic acid (Coppins 2002). However, based on a molecular analysis of the M. prasina group, Czarnota and Guzow- Krzemińska (2010) found that M. micrococca sensu Coppins is not monophyletic and includes, in fact, three separate lineages: M. micrococca s. str. (hereafter referred to as M. micrococca), M. byssacea, and a yet unnamed taxon. In spite of their similarity in secondary chemistry with all producing methoxymicareic acid, the species have some differences in morphology and ecology. Below we discuss the differences between M. prasina (fig. 1A), M. bysscaea (fig. 1B) and M. micrococca (fig. 1C), based on both European and North American material. Because of the taxonomic uncertainties, and the lack of collections from North America, the unnamed taxon is not discussed thoroughly here. Czarnota and Guzow- Krzemińska (2010) considered the unnamed taxon to be cryptic and an intermediate between M. 85 Figure 1, morphology of Micarea species discussed in this paper (A–D and F) and geographic distribution of M. hedlundii in North America (E). A, apothecia and thallus of M. prasina (Launis 229106, H). B, apothecia and thallus of M. byssacea (Launis 66128, H). C, apothecia and thallus of M. micrococca (Launis 1010131, H). D, olive-green thallus composed of goniocysts and stalked lilac pycnidia coated with white tomentum in M. hedlundii (Launis 67119, H). E, known geographic distribution of M. hedlundii in North America. F, stalked pycnidium releasing mesoconidia in M. hedlundii (Launis 66125, H) Scales = 1.0 mm in A-D, 100 µm in E. 86 micrococca and M. byssacea. However, our unpublished results and personal comments from P. Czarnota show that some morphological characters are found. These characters include small (0.1–0.3 mm) and adnate to hemispherical apothecia with the Sedifolia-grey pigment, or without the pigment but with smaller ascospores than in M. micrococca. If specimens like this were to be collected from North America they should be sequenced to determine whether they belong to the unnamed taxon. Morphologically, Micarea byssacea resembles M. micrococca and M. prasina, but M. prasina differs from the first two species by the production of micareic acid and by mainly growing on soft lignum. Although all three species have immarginate apothecia, a thallus composed of goniocysts, and almost no differences in spore sizes, M. prasina can quite easily be identified with TLC and also by its preference for decaying lignum instead of bark. Similarly to Micarea micrococca, M. byssacea produces methoxymicareic acid and occurs mainly on acid bark. However, M. byssacea differs in the presence of the Sedifolia-grey pigment (K+ violet, C+ violet), as well as in its larger, adnate and usually darker colored apothecia (Czarnota & Guzow- Krzemińska 2010). In M. micrococca the apothecia are colorless throughout in section and always pale externally, without any darker pigmentation. In M. byssacea and M. prasina the pigmentation of the apothecia is variable, probably in response to the amount of light. Therefore, in shaded habitats the apothecia can be very pale or whitish with no Sedifolia-grey pigment, and these forms are not always easy to separate from those of M.
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