Opuscula Philolichenum, 13: 80-83. 2014. *pdf effectively published online 20June2014 via (http://sweetgum.nybg.org/philolichenum/)

A new species of Lecidea (Lecideaceae, Lichenized Ascomycetes) from the mountains of California

1,2 3 KERRY KNUDSEN & JANA KOCOURKOVÁ

ABSTRACT. – Lecidea oreophila, a species similar to L. tessellata, is described from the mountains of California (Sierra Nevada, San Bernardino, San Jacinto) from sites above 1958 meters in elevation. The new species produces 2′-O-methylmicrophyllinic acid with or without confluentic acid.

KEYWORDS. – Alpine lichens, montane lichens, taxonomy.

INTRODUCTION

The Lecidea species of the Sonoran Desert Region in western North America were recently monographed by Hertel and Printzen (2004). That monograph was an important step forward in the study of the North American lichen flora because the genus Lecidea, like other crustose genera such as Acarospora and Aspicilia, was still poorly understood at the beginning of the 21st century. Nonetheless, the problems in Lecidea s. str. in western North America are far from resolved. The delineation of species and infraspecific taxa in the L. fuscoatra and L. atrobrunnea groups needs re-examination (for instance see Hutten et al. 2013). A number of endolithic species including L. cruciaria Tuck. and L. kingmanii (Hasse) Hertel & S. Ekman are in need of revision. Several new taxa have been discovered (Knudsen et al. 2013). In this paper we describe one of these new taxa from the mountains of California.

MATERIALS AND METHODS

Specimens were examined using standard microscopical techniques with an Olympus SZX 7 stereomicroscope or an Olympus BX 51 fitted with Nomarski differential interference contrast. Specimens were compared with material in the Herbarium Mycologium of J. Kocourková and K. Knudsen from the mountains of the Czech Republic and specimens in the herbaria of GZU, NY and UCR. Secondary metabolites were studied using standardized thin-layer chromatography in Solvent C (Culberson & Ammann 1979, Culberson & Johnson 1982, Orange et al. 2001, 2010). Hand cut sections were studied in water and 10% KOH (K). Amyloid reactions were tested in Lugol’s iodine [I] with and without pretreatment with K. Nitric acid 70% was used for chromatic reactions of inspersions. Ascospores and conidia were measured in water with an accuracy of 0.5 μm and given in the form “(minimum–) mean minus standard deviation–average–mean plus standard deviation (–maximum)” rounded to nearest 0.5 microns and followed by the number of measurements (n); the length/breadth ratio is indicated as l/b and given in the same way.

1 KERRY KNUDSEN – Department of Ecology, Faculty of Environmental Sciences, Czech University of Life Sciences, Prague, Kamýcká 129, Praha 6 - Suchdol, CZ–165 21, Czech Republic 2Herbarium, Department of Botany and Plant Sciences, University of California, Riverside CA 92521– 0124, U.S.A. – e-mail: [email protected] 3 JANA KOCOURKOVÁ – Department of Ecology, Faculty of Environmental Sciences, Czech University of Life Sciences, Prague, Kamýcká 129, Praha 6 - Suchdol, CZ–165 21, Czech Republic – e-mail: [email protected]

80 THE NEW SPECIES

Lecidea oreophila K. Knudsen & Kocourk., sp. nov. Mycobank #809259. FIGURE 1.

Similar to Lecidea tessellata, but has a dark hypothecium and produces 2′-O-methylmicrophyllinic acid as the primary substance, with or without accessory confluentic acid.

TYPE: U.S.A. CALIFORNIA. RIVERSIDE CO.: San Bernardino National Forest, San Jacinto Mountains, Black Mountain, Black Mountain Fire Outlook area, 33°49′28″N, 116°45′31.4″W, 2349 m, 10.viii.2012, on large granite boulders, K. Knudsen 15013 (UCR!, holotype).

DESCRIPTION. – Thallus to 10.0 cm or more in diameter, but often smaller, irregularly areolate with deep fissures between areoles, not flat or thin in appearance, areoles to 2.0 mm wide, to 0.5 mm thick, variable in shape, vegetatively dividing, broadly attached. Upper surface rugulose, light to dark gray, sometimes slightly bluish. Apothecia to 1.0 mm wide, margin thin and persistent, disc flat to convex, epruinose, arising from the areoles, usually higher than thallus surface, vegetatively dividing, first appearing on top of the areoles, eventually expanding to give the appearance of occuring between areoles. Cortex 10–30 μm thick, of round to angular cells, mostly 4–5 μm wide, becoming gelatinized below a thin epinecral layer. Algal layer 80–120 μm, uninterrupted, algal cells 8–15 μm wide. Medulla I+ violet, 0.1–0.4 mm thick, often obscured by crystals, prosoplectenchymatous, hyphae 4–6 μm in width, thin-walled, sometimes rugulose, branching, continuous with attaching hyphae. Exciple 60–90 μm wide, of radiating hyphae, hyphae 1.5–2.0 μm in width, apices expanded to 5 μm, thin outer layer green-black, internally hyaline. Epihymenium green-black, ca. 10 μm tall, N+ reddish- purple. Hymenium 60–80 μm tall. Asci clavate, 8–spored, mostly 40–50 × 20–40 μm. Ascospores hyaline, thin-walled, (12–)12–13–14 × (4–)4.5–5.5–6 μm (n=20), l/b = 2.0–2.5–3.0(–3.5). Paraphyses 1.5–2.0 μm wide, rarely branching, apices expanded up to 5 μm. Subhymenium 30–40 μm tall, pigmented light or dark brown in thin section, darker in thick section, N+ reddish-purple, not easily distinguished from hypothecium. Hypothecium 80–100 μm, pigmented light to dark brown, N+ reddish-purple. Pycnidia, ostiolate, not graphidoid, appearing as dark dots. Conidia hyaline, (9.0–)10–13–15.5(–20) × 1–2 μm (n=40).

CHEMISTRY. – 2′-O-methylmicrophyllinic acid (major), with or without confluentic acid (major), with or without a trace of 2′-O-methylperlatolic acid.

ETYMOLOGY. – The epithet “oreophila” means “mountain lover” and refers to the occurrence of the species in the mountains of California above 1958 meters.

ECOLOGY AND DISTRIBUTION. – Lecidea oreophila is locally frequent. It occurs above 1958 meters on granite in saxicolous montane communities usually associated with members of the Lecidea atrobrunnea complex. It is known from scattered localities in the mountain of central and southern California (Sierra Nevada Mountains; San Bernardino Mountains; San Jacinto Mountains).

DISCUSSION. – Lecidea oreophila belongs to Lecidea sensu stricto based on ascus stain (Hertel & Printzen 2004) and is similar to L. tessellata Flörke. The irregularly areolate thallus with apothecia usually higher than the areoles separates L. oreophila from the regularly areolate thallus of L. tessellata which has been described as having “half-sunken” apothecia (Hertel & Printzen 2004; for image of a typical thallus see Brodo et al. 2001). Lecidea tessellata has graphidoid pycnidia, a usually hyaline hypothecium, and the consistent production of confluentic acid (Hertel & Printzen 2004). Lecidea oreophila differs in having non-graphidoid pycnidia, a dark subhymenium and hypothecium in thin section, and the consistent production of 2′-O-methylmicrophyllinic acid with or withour accessory confluentic acid. In California, L. tessellata is the only species of the genus in montane communities with which L. oreophila could be confused. In southern California, based on 25 collections from UCR (UCR Herbarium 2014), L. tessellata is common but was not collected above 2056 m and often occurs at lower elevations down to 635 m in the Santa Ana Mountains. Lecidea tessellata also occurs in the Little San Bernardino Mountains of Joshua Tree

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Figure 1, holotype of Lecidea oreophila from the San Jacinto Mountains, K. Knudsen 15013 (UCR), scale = 1.0 mm.

National Park in the southwestern Mojave Desert (Knudsen et al. 2013; image in Sharnoff 2014). During our fieldwork throughout the region where L. oreophila is known to occur, L. tessellata was not found in any communities with L. oreophila. Lecidea oreophila could also be confused with L. lapicida (Ach.) Ach. which is not known from western North America (Hertel 2006, Hertel & Printzen 2004). Lecidea lapicida has ascospores of the same general length as L. oreophila but the mature ascospores are generally more broadly ellipsoid (6–8 μm wide vs. 4–6 μm in L. oreophila; fide Aptroot et al. 2009). In his excellent monograph on Lecidea s. str., though Schwab gave different and smaller spore sizes for L. lapicida, he nonetheless considered the ascospores of L. lapicidia to be broadly ellipsoid (Schwab 1986). The ascospores of L. oreophila are usually ellipsoid, though immature spores in the ascus can be globose in early ontogeny. Lecidea lapicida differs chemically from L. oreophila in lacking 2′-O-methylmicrophyllinic acid and confluentic acid, and in producing norstictic acid (in L. lapicida var. pantherina (Hofm.) Ach. [=L. pantherina (Hoffm.) Ach.] ) or the stictic acid complex (in L. lapicidia var. lapicida) (Aptroot et al. 2009). Lecidea confluens is rare in North America (Hertel 2006) with only one report from California (Knudsen 2012). These sites could be Pleistocene relics. The ascospores of L. confluens are 9–13 × 4.4–5.6 μm in size (Schwab 1986) and overlap the size range of ascospores in L. oreophila, with L. oreophila having generally longer ascospores (12–14 μm). Lecidea confluens has a flatter, smoother, regularly areolate thallus (see illustration in Wirth et al. 2013) rather than the irregularly areolate and rugulose thallus of L. oreophila (figure 1 herein). Lecidea confluens consistently produces confluentic acid and 2′-O- methylperlatolic acid and sometimes stictic acid (Aptroot et al. 2009). Lecidea oreophila differs in producing 2′-O-methylmicrophyllinic acid consistently, sometimes with accessory confluentic acid and 2′- O-methylperlatolic acid. It is quite possible that L. oreophila is related to L. confluens as well as L. lapicidia and thus should be included in future molecular phylogenetic studies of Lecidea s. str.

82 Based on its few recorded occurrences, we currently consider Lecidea oreophila to be rare. In the San Jacinto Mountains, the species could become extirpated by wildfires. The 2013 Mountain Fire in the San Jacinto Mountains, which devastated over 30-square-miles of suitable habitat where L. oreophila could have occured, prevented more fieldwork aimed at locating additional populations. Fortunately the fire did not extirpate any of the populations of L. oreophila reported in this paper.

Additional specimens examined. – U.S.A. CALIFORNIA. MONO CO.: Inyo National Forest, Tioga Pass, 37°55′40.7″N 119°15′21.3″W, 2930 m, 13.vii.2012, on granite outcrops, K. Knudsen et al. 14732 (NY), K. Knudsen et al. 14740.1 (UCR), Knudsen et al. 14740.2 (FH). RIVERSIDE CO.: Mount San Jacinto State Park, San Jacinto Mountains, San Jacinto State Wilderness Area, near trail to Round Valley, 33°48′27″N 116°48′39.6″W, 2676 m, 17.xi.2006, on granite boulders, K. Knudsen 7892 & L. Kuzina (UCR); same locality, 33°48′39.6″N 116°39′02″W, 2699 m, 3.vi.2013, on granite boulders, K. Knudsen 15815 (UCR). SAN BERNARDINO CO.: San Bernardino National Forest, San Bernardino Mountains, 2N02, crossing at Arrastre Creek, pinyon-juniper woodland, 34°15′19.6″N 116°44′39.2″W, 1959 m, 20.iii.2014, on granite outcrop, Knudsen et al. 16510 (NY, hb. Kocourková and Knudsen)

ACKNOWLEDGEMENTS

We thank our reviewers for their comments and suggestions. We thank the curators of GZU and NY. We thank James C. Lendemer for the thin-layer chromatography of specimens. We thank USDA Forest Service botanists Scott Eliason, Diane Iketa, Kate Kramer, and Sue Weiss for their enthusiasm and support of our montane research as well as Ken Kietzer of Mount San Jacinto State Park. The holotype was collected preparing a lichen workshop for the FS and the specimens on Tioga Pass were collected while preparing for a FS lichen walk for the public. The work of both authors Kerry Knudsen and Jana Kocourková was financially supported by the grant “Environmental aspects of sustainable development of society” 42900/1312/3166 from the Faculty of Environmental Sciences, Czech University of Life Sciences Prague.

LITERATURE CITED

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