Further contributions to the genus Rinodina (Physciaceae, Lecanoromycetidae): two species new to science and a new record for the Canadian High Arctic

Herzogia 25 (2), 2012: 125 143 125 Further contributions to the genus Rinodina (Physciaceae, Lecanoromycetidae): two species new to science and a new record for the Canadian High Arctic John W. Sheard *, James C. Lendemer, Toby Spribille, Göran Thor & Tor Tønsberg Abstract: Sheard, J. W., Lendemer, J. C., Spribille, T., Thor, G. & Tønsberg, T. 2012. Further contributions to the genus Rinodina (Physciaceae, Lecanoromycetidae): two species new to science and a new record for the Canadian High Arctic. Herzogia 25: 125 143. Two species of Rinodina are described as new to science (R. buckii and R. bullata) and a third is reported for the first time from arctic Canada (R. ochracea). Rinodina buckii, a sorediate species that resembles R. willeyi is described from disjunct localities in East Asia and the Appalachian Mountains of eastern North America. Rinodina bullata, a species with a peculiar minutely areolate thallus that is frequently sterile, is described from the central Appalachian Mountains of eastern North America. Rinodina ochracea, originally described from Greenland, is reported from the Northwest Territories in Canada. The newly described species are illustrated with both colour light micrographs (CLMs) and scanning electron micrographs (SEMs), and maps of their geographic distributions are presented. CLMs and SEMs of R. excrescens and R. willeyi are also provided for comparison. Zusammenfassung: Sheard, J. W., Lendemer, J. C., Spribille, T., Thor, G. & Tønsberg, T. 2012. Weitere Beiträge zur Kenntnis der Flechtengattung Rinodina (Physciaceae, Lecanoromycetidae): zwei neue Arten und ein Erstnachweis für die kanadische Arktis. Herzogia 25: 125 143. Zwei neue Arten der Flechtengattung Rinodina, R. buckii und R. bullata, werden beschrieben und eine dritte, R. ochracea, wird erstmals für das nordamerikanische Festland nachgewiesen. Rinodina buckii ist eine sorediöse Art mit Ähnlichkeiten zu R. willeyi, welche nach dem jetzigen Kenntnisstand disjunkt in Ostasien und im Appalachengebirge des östlichen Nordamerikas vorkommt. Rinodina bullata, eine häufig sterile Art mit winzig areoliertem Lager, wird ebenfalls aus den Appalachen beschrieben. Die ursprünglich aus Grönland beschriebene Rinodina ochracea wird aus den kanadischen Northwest Territories nachgewiesen. Die neu beschriebenen Arten sowie vergleichend R. excrescens und R. willeyi werden anhand von mikroskopischen Farbfotos und rasterelektronischen Aufnahmen abgebildet und bekannte Vorkommen mittels Verbreitungskarten dargestellt. Key words: Lichenized Ascomycetes, phytogeography, sterile crust, soredia, blastidia. Introduction New species and new records of the genus Rinodina (Ach.) Gray continue to be added to the North American flora (Sheard 2011, Sheard et al. 2011, Lendemer et al. 2012) despite this crustose genus being relatively well known on the continent (Sheard 2010). Here we describe two new species from eastern North America, R. buckii Sheard and R. bullata Sheard & Lendemer, and report a new record from the high arctic, R. ochracea Lynge. Rinodina buckii * corresponding author

126 Herzogia 25 (2), 2012 also occurs in Japan, Korea and the Russian Far East, exhibiting a similar distribution to R. chrysidiata Sheard (Lendemer et al. 2012) and R. excrescens Vain. (Galanina et al. 2011). The total number of Rinodina species now known from North America is 105. Materials and Methods The study is based primarily on specimens collected by the authors (JL, TS, GT and TT) and loans from the herbaria cited in the acknowledgements. Surface observations of specimens were made using a Wild M5 stereomicroscope. Thallus measurements were taken at 25 magn ification and rounded to the nearest 0.05 mm. Internal ascomatal measurements were made on vertical sections (c. 25 µm thick), cut with a Leitz freezing microtome, at 50 magnification to an accuracy of 5 µm using a Wild M20 compound microscope. Internal ascospore measurements were taken at 500 magnification using a Wild vernier micrometer (scale of 0.1 µm) to an accuracy of 0.5 µm. They are quoted as the range between the 25 th and 75 th percentiles with the 5 th and 95 th percentiles indicated in brackets. Observations of ascospore wall structure were made with an oil immersion lens at a magnification of 1250. Ascospore structure of fresh collections is conveniently revealed by heating water mounted slide sections over a methyl alcohol burner. For species with a conglutinate hymenium this also has the effect of aiding the release of ascospores from the ascus on applying gentle pressure to the cover slip. Light micrographs from a dissecting microscope were prepared following the methods outlined by Lendemer (2011) and scanning electron micrographs were prepared following Lendemer & Elix (2010). The species Rinodina buckii Sheard sp. nov. [Mycobank 801362] Figs 1, 2A B, 3A B, 4 and 6B. Type: U.S.A. North Carolina. Swain Co.: Great Smoky Mountains National Park, Hyatt Ridge Trail between Beech Gap Trail and Enloe Creek Trail, upland ridge forest with Picea rubens, Quercus rubra, and Acer saccharum grading into Quercus alba, Q. montana, Castanea dentata with Kalmia latifolia, Rhododendron maximum, and Vaccinium understory, on fallen branch, 6 Aug. 2009, J. C. Lendemer 19269 & E. A. Tripp (NY, holotype). Diagnosis. Thallus thin, areolate; areoles becoming verrucose, verrucae developing raised soralia; apothecia erumpent, ascospores Teichophila-type, (19.5 )22.0 25.0( 28.0) (10.5 ) 11.5 13.0( 15.0) µm; pannarin present, zeorin usually present. Etymology. Named after William Bill R. Buck (1950 present), noted bryologist and the first person to collect this new species. Description. Thallus thin, dark grey, sometimes brownish, initially of plane, isolated areoles c. 0.20 mm wide (Figs 1B & D, 4A & B), coalescing to form larger areoles to 0.40 1.00 mm wide, sometimes becoming continuous; quickly becoming verrucose, verrucae c. 0.20 mm diam., breaking into soredia (Figs 1C & E, 4D F); vegetative propagules always present except in youngest, marginally isolated areoles; soralia limited to verrucae at first (Figs 4B & C), 0.15 0.60 mm wide; soredia 40 65 µm diam., raised, finally often dissolving areoles (Figs 1F, 4F and 6B); thallus margin determinate or indeterminate, prothallus dark, fimbriate, or not evident. Apothecia erumpent, then broadly attached, finally becoming more narrowly att ached at base (Fig. 1A), 0.35 0.70 mm diam.; disc black, sometimes grey pruinose, pers istently plane or rarely developing a central umbo; thalline margin prominent, c. 0.10 mm wide, thalline layer sometimes

Sheard et al.: Further contributions to the genus Rinodina 127 Fig. 1: Morphology of Rinodina buckii (A C from Lendemer 10408, D F from Lendemer 19241). A sectioned, narrowly attached apothecium. B F development of the thallus in which the convex areoles (B) develop soralia on raised verrucae (C D) that eventually dissolve the verrucae and areoles (E F). Scales = 0.5 mm: F, 0.25 mm: A E. thin, poorly formed, translucent when wet, or thick and sored iate; excipular ring often present, confluent with thalline margin. Apothecial anatomy: thalline exciple 50 100 µm wide laterally; cortex poorly differentiated, 5 10 µm wide when present, epinecral layer sometimes present, 5 10 µm wide; crystals present in cortex and med ulla (pannarin); cortical cells 3.0 5.0 µm wide, not pigmented; algal cells 9.0 14.5 µm in diameter; proper exciple 10 30 µm wide laterally, some-

128 Herzogia 25 (2), 2012 Fig. 2: Comparison of the Teichophila-type ascospores of Rinodina buckii (Tønsberg 22507) and Pachysporariatype I ascospores of R. willeyi (Lendemer 32335). A B Rinodina buckii. A Immature spores with Physcia-like, relatively angular lumina, mostly with narrow canals and thickened walls at the apices. B Mature spores with more rounded lumina although mostly retaining some apical wall thickening. Tori are visible in various foci. C D Rinodina willeyi. C Immature spores with irregularly rounded lumina, broad canals and relatively thin apical walls. D Mature spores with well rounded lumina and well developed tori. Scales = 10 µm. times lightly pigmented, expanding to 40 70 µm wide above; hypothecium mostly light brown, 40 90 µm tall; hymenium hyaline, 110 130 µm high, not inspersed, paraphyses 2.0 3.0 µm wide, conglutinate, apices expanded to 4.0 4.5 µm wide, lightly pigmented, immersed in dispersed pigment forming a red-brown epihymenium. Asci c. 60 90 20 30 µm. Ascospores 4 or 8/ascus, development type A, often asynchronous (4 mature, 4 immature), Teichophila-type (Sheard 2010), (19.5 )22.0 25.0( 28.0) (10.5 )11.5 13.0( 15.0) µm, l/w ratio (1.6 )1.8 2.0( 2.3) (n = 186), lumina canals broad, poorly formed, apical walls thickened at first, Physcia-like (Fig. 2A), typically becoming more rounded at maturity (Pachysporaria-like), but apical thickening mostly retained (Fig. 2B), rarely Mischoblastia-like when overmature; prominent torus present at maturity; walls becoming darkly pigmented; not ornamented. Pycnidia not seen. Chemistry. Spot tests, K, C, KC, P+ cinnabar; secondary substances, pannarin (in cortex and medulla), ± zeorin (the latter sometimes in low concentration and not detected in Tønsberg 22507, 22936a). Pannarin is absent from the epihymenium. Ecology and distribution. Rinodina buckii appears to be another example of a lichen species with a disjunct distribution between the Appalachian Mountains of eastern North America and Eastern Asia (Yoshimura 1968, Culberson 1972, Kurokawa 2006, Sheard et al. 2008, Galanina et al. 2011, Lendemer et al. 2012). In eastern North America the species has been found throughout the Appalachian Mountains, although it appears to be most common in the southern portion of the mountain range (Fig. 3A). It has been collected primarily on the bark of maples (Acer) and

Sheard et al.: Further contributions to the genus Rinodina 129 Fig. 3: Geographic distributions of Rinodina buckii, North America (A), East Asia (B) and Rinodina bullata (C). less commonly on the bark of other hardwoods (Betula, Carya, Fraxinus and Quercus) at low to middle elevations (225 1535 m). On two occasions it has been found growing associated with R. adirondackii H.Magn., an esorediate species with which it otherwise could be mistaken. The collections of this species seen from Eastern Asia were made in Hokkaido, Japan, South Korea and far eastern Russia (Fig. 3B). In Japan Rinodina buckii has been collected primarily on Abies and also on Alnus, Malus, Phellodendron, Quercus, Sorbus and Taxus at elevations of 15 750 m. The single Russian collection examined occurred on Picea at 340 m. Southward in South Korea the species has been collected on Abies and Pinus at higher elevations of 1300 1375 m.

130 Herzogia 25 (2), 2012 Fig. 4: Scanning electron micrographs of Rinodina buckii (all from Lendemer 19241). A F development of the thallus in which A, thalline verrucae (t) develop soralia (s), initiating in B C, and more developed in D, eventually dissolving the verrucae leaving only masses of aggregated soredia weakly held together by gelatinized hyphae (E F). Scale bars as indicated. Discussion. Fertile specimens of Rinodina buckii might be mistaken for R. adirondackii H.Magn. because of the erumpent nature of their young apothecia which later become more narrowly attached. Closer inspection, however, will reveal that the thallus of R. buckii becomes verrucose, at least in part, rather than remaining plane, and that the verrucae develop irregu larly shaped, raised soralia, eventually dissolving the areoles into aggregations of coarse

Sheard et al.: Further contributions to the genus Rinodina 131 Fig. 5: Morphology of Rinodina willeyi (A C from Lendemer 18989, D F from Lendemer 23081). A widely attached apothecium, the margin typically irregular in outline. B D areoles with marginal soralia. E F soralia spreading over areole surfaces. Scales = 0.25 mm. soredia (see Figs 1B F and 4B F). These aggregations of soredia can eventually overlap and appear to form a continuous leprose crust (Figs 1F, 4F and 6B), although this process happens more sparingly on thalli in the fertile state. Another critical difference between the two species is that the spores of R. adirondackii are larger and possess a typical Pachysporaria-type I structure including nearly polygonal lumina in some immature spores (Sheard 2010).

132 Herzogia 25 (2), 2012 Fig. 6: Scanning electron micrographs comparing verrucate soralia of Rinodina buckii (B Lendemer 19241) to marginal soralia of R. willeyi (A Lendemer 23081), t = thallus, s = soralium, e = young soralium emerging from small thalline verruca. Scale bars as indicated. Due to the combination of frequent sterility, production of lichenized diaspores, and areolate thallus, Rinodina buckii is likely to be confused with several morphologically similar corticolous species that are largely sympatric in eastern North America. Three of these species have sorediate, areolate thalli, namely R. degeliana Coppins, R. efflorescens Malme, and R.

Sheard et al.: Further contributions to the genus Rinodina 133 willeyi ** Sheard & Giralt. Of these, the one most likely to be confused with R. buckii is R. willeyi because both species produce pannarin and zeorin. Indeed, all but one of the specimens of R. buckii in NY were originally identified as R. willeyi. However, R. willeyi usually has a lighter grey thallus (Fig. 5A E), does not develop verrucae (Figs 5B E and 6A), and has soralia originating on the areole margins (Figs 5B F, 6A and 7B E) that do not typically fully diss olve the areoles leaving only aggregations of soredia (see Fig. 6, and compare Figs 1F and 4F of R. buckii to Figs 5B F and 7B F of R. willeyi). The soredia of R. willeyi are also smaller (< 40 µm diam.) than those of R. buckii. While R. willeyi has ascospores of a similar size to those of R. buckii they differ in belonging to the Pachysporaria-type I group rather than the Teichophila-type (Fig. 2C D). The ascospores of R. buckii have a more variable lumina shape during development. In the later stages of development they may have Pachysporaria-type I tendencies but we prefer to place them in the Teichophila-type for the present, reflecting the rather variable internal structure earlier in their development (Sheard & Mayrhofer 2002, Sheard 2010) despite the fact that this is perhaps the least well understood spore type in the genus. The large range of ascospore size and shape quoted above is at least partly due to asci with only four spores having larger spores. Rinodina efflorescens is chemically similar to R. buckii in the production of pannarin, however that species only produces zeorin as a trace accessory to pannarin (Tønsberg 1992, Lendemer unpublished data) and always produces the xanthone secalonic acid (Tønsberg 1992, Sheard 2010). The presence of the latter substance typically gives the soralia of R. efflorescens a distinctive yellowish colouration that is often easily seen even in the field with a hand lens. The thallus of R. efflorescens also differs from that of R. buckii in that the soralia typically remain discrete, are not raised and rarely dissolve the entire thallus. When fertile, the two species are easily distinguished by their ascospores since those of R. efflorescens are smaller and belong to the Physcia-type (Sheard 2010). Rinodina degeliana is least likely to be confused with R. buckii because it produces atranorin rather than pannarin in addition to zeorin (Tønsberg 1992, Sheard 2010). Rinodina degeliana also has marginal soralia that typically do not expand to fully dissolve the areoles. The areoles of R. degeliana are typic ally lighter grey than those of R. buckii and are further distinguished by their almost white, marginal soredia. The spores of R. degeliana are smaller and belong to the Physcia-type when immature and later the Physconia-type, making the two species readily separable when fertile. The only sorediate Rinodina species previously reported from eastern Asia is R. xanthophaea (Nyl.) Zahlbr. (Kurokawa & Kashiwadani 2006). This species has a pigmented thallus cont aining xanthones, has Physcia-type spores when fertile and therefore presents no possibility for confusion with R. buckii (Lendemer et al. 2012). An emended version of the key from Sheard (2010) is presented below to facilitate the identification of Rinodina buckii. 45 Soredia in discrete soralia; similar to colour of thallus; apothecia not common; never with pann arin in epihymenium; rarely southwestern... 45a 45a(45) Thallus minutely verrucose verrucae quickly becoming sorediate, initially forming small, isol ated, raised soralia, later spreading over thallus surface... R. buckii 45a Thallus not verrucose soralia not raised... 46 ** Following Art. 60.11 and Rec. 60C.1.A of the ICBN the epithet of Rinodina willeyi should be spelled with a single i rather than with a double ii.

134 Herzogia 25 (2), 2012 Fig. 7: Scanning electron micrographs of Rinodina willeyi (all from Lendemer 23081). A apothecium and thallus. B D development of marginal soredia. E F soralia spreading to the surface of the areoles. Scale bars as indicated. Additional specimens examined: Japan. Hokkaido. Ishikari Prov.: Kamikawa-gun, Kamikawa-cho, Obako Tourist Centre, 10 June 1995, on Alnus, G. Thor 14612, on Abies, 14624 (UPS). Kitami Prov.: Rishiro-to Island, Rishiri-gun, Rishirifuji-cho, 2.5 km S Sakae, 01 June 1995, on Abies, T. Tønsberg 22385 (BG); Namura village, 02 June 1995, on Abies, G. Thor 13943 (UPS); Oniwaki area, 02 June 1995, on Abies, T. Tønsberg 22506 (BG), T. Tønsberg 22507 (BG); Shari-gun, Shari-cho, Shiretoko National Park, 06 June 1995, on Abies, T. Tønsberg 22818 (BG), T. Tønsberg 22820c (BG), T. Tønsberg 22822 (BG); on Taxus, G. Thor 14249 (UPS); Soya-gun, Sarufutsu-mura, 14 km NW Kamatombetsu town, 04 June 1995, on Malus, T. Tønsberg 22579 (BG). Kushiro Prov.: Akkeshi-gun, Hamanaka-cho, 08 June 1995, on Sorbus, T. Tønsberg 22935a (BG), T. Tønsberg 22936a pro parte (BG); 3 km E Hichiripputo Lake,

Sheard et al.: Further contributions to the genus Rinodina 135 Fig. 8: Rinodina bullata (A, C E from Lendemer 12193, B from Lendemer 16730, and F from Lendemer 25485). A C narrowly attached apothecia with entire margins and plane discs. D F morphological variation in the bullate thalline areoles: a well developed form with overlapping areoles (D), a dispersed form with slightly overlapping areoles (E), and a young thallus with immature apothecia developing on several areoles (F). Scales = 0.5 mm: A B and D; 0.25 mm: C, E and F. on Alnus, G. Thor 14450 (UPS); Kushiro-gun, Kushiro-cho, 13 km E Kushiro, 08 June 1995, on Sorbus, T. Tønsberg 23020 (BG), on Taxus, T. Tønsberg 23025, 23026 (BG). Nemuro Prov.: Shiretoko Peninsula, Menashi-gun, Rausocho, 07 June 1995, on Quercus, T. Tønsberg 22905 (BG). Teshio Prov.: Rumoi-gun, Obira-cho, 21 km E Obira, 28 May 1995, on Abies, T. Tønsberg 21996 (BG); Teshio-gun, Toyotomi-cho, Rishiri-Rebun National Park, 30 May

136 Herzogia 25 (2), 2012 1995, on Abies, T. Tønsberg 22185 (BG). Tokachi Prov., Achiro-gun, 3 km WSW Kitaichijo village, 09 May 1995, on Quercus, G. Thor 14564 (UPS). Russia. Khabarovskiy Krai. Sonakh River, Amgun River region, c. 7.7 km NW main Berezoviy-Badzhal route, logging road, 25 July 2009, T. Spribille 31397 & L. Yakovchenko (GZU). South Korea. Cheju-Do. Cheju Island, Eorimok National Park Office, NW slope Mt. Halla, 21 May 2001, on Pinus, G. Thor 17213 (UPS). Gangwon-Do, Yanggyang-gun, Osaeki-ri, Sorak-San National Park, Mt. Dachong, 20 Oct. 2006, on Abies, G. Thor 20595 (UPS), G. Thor 20620 (UPS). U.S.A. Maine. Hancock Co.: T28 MS, SE slopes of Lead Mountain, terminus of gravel road off ME9, 11 June 2010, on Fraxinus, J. C. Lendemer 23161 (NY). New York. Greene Co.: Catskill Mountains, N end of Mink Hollow and W slope of Sugarloaf Mountain, 08 Oct. 2007, on Acer, J. C. Lendemer 9803 & A. Moroz (NY). Ulster Co.: Catskill Park, trail to Belleayre Mt. from Lost Clove Trail Head Parking Area, 10 May 1996, on Quercus, R. C. Harris 38481 (NY). North Carolina. Clay Co.: Nantahala National Forest, 1 1.5 mi N of US 64 on Buck Creek Rd., vicinity of Doe Knob, 10 Nov. 2007, on Betula, J. C. Lendemer et al. 10408 (NY). Haywood Co.: Great Smoky Mountains National Park, E slopes above (E of) Caldwell Fork, 4 Aug. 2009, on Acer, J. C. Lendemer 18874 (NY). Macon Co.: Nantahala National Forest, N slope of Wayah Bald, 12 Oct. 1998, on bark, W. R. Buck 35134 (NY, w/ R. adirondackii). Swain Co.: Great Smoky Mountains National Park, Beech Gap Trail between Straight Fork Rd./Balsam Mountain Rd. along Hyatt Ridge Trail, 06 Aug. 2009, on Acer, J. C. Lendemer 19241 & E. Tripp (NY); Great Smoky Mountains National Park, Heintooga Ridge Rd., Polls Gap, 29 June 2002, on Fagus, T. Tønsberg 30922 (BG), along Heintooga Ridge Rd./Blue Ridge Parkway Extension, between Parkway and National Park border, 20 June 2002, on Acer, T. Tønsberg 30904 (BG), Balsam Mountain Rd., 20 June 2002, on Acer, T. Tønsberg 30932 (BG). Pennsylvania. Cameron Co.: Moshannon State Forest, Grant Trail just E of Cameron/Elk County line, slopes of Mile Run, 31 Aug. 2010, on Carya, J. C. Lendemer 24018 (NY). Monroe Co.: Delaware Water Gap National Recreation Area, Community Drive Wetlands, 17 Sept. 2005, on Acer, J. C. Lendemer 51610-A (NY). Tennessee. Blount Co.: Great Smoky Mountains National Park, Gregory Bald Trail, 12 Oct. 2010, on Acer, J. C. Lendemer et al. 26798 (NY); Great Smoky Mountains National Park, Cades Cove Mountain, summit pass Rich Mountain Rd., 14 June 2002, on Acer, T. Tønsberg 30744a (BG). Rinodina bullata Sheard & Lendemer sp. nov. [Mycobank 801364] Figs 8 10. Type: U.S.A. Ohio. Scioto Co.: Shawnee State Forest, Copperhead Lookout Tower, 0.75 mi S of Bear Creek Lake, 21 May 2006, on Quercus, J. C. Lendemer et al. 7239 (NY [Barcode #0169444], holotype; NY [Barcode #01220500]; SASK, isotypes). Diagnosis. Thallus light grey, areoles minute, loosely attached, bullate; apothecia narrowly attached; spores Pachysporaria-type II, (13.0 )14.5 16.5( 17.5) (7.5 )8.0 9.0( 10.0) µm, lightly pigmented with spherical lumina at maturity; atranorin in cortex. Etymology. The epithet bullata is derived from the Latin bullatus and refers to the distinct ive morphology of the areoles that comprise the thallus. Description. Thallus light grey, entirely of minute, loosely attached, bullate areoles, 0.10 0.15( 0.20) mm diam., discrete at first, becoming confluent; surface matt; thallus margin indeterminate, prothallus not seen; soredia and isidia absent (Figs 8 9). Apothecia narrowly attached when present, usually not contiguous, 0.30 0.60 mm diam.; disc brown, persistently plane; thalline margin concolourous with thallus, 0.05 0.10 mm wide, entire or beaded, persistent; excipular ring absent (Figs 8A C). Apothecial anatomy: thalline exciple 50 110 µm wide: cortex poorly differentiated, 5 10 µm wide; crystals present in cortex (atranorin), absent in medulla; cortical cells 4.0 5.0 µm wide, not pigmented; algal cells 11.0 15.0 µm diam.; proper exciple hyaline, 5 10 µm wide laterally, expanding to c. 20 µm above; hypothecium 25 60 µm tall, hyaline; hymenium not inspersed, 80 90 µm high; paraphyses 1.5 2.0 µm wide, moderately conglutinate, apices 3.5 4.5 µm wide, lightly pigmented and immersed in dispersed pigment forming a light red- to orange-brown epihymenium; asci 50 60 15 16 µm. Ascospores 8/ascus, Type A development, Pachysporaria-type II (Sheard 2010), (13.0 )14.5 16.5( 17.5) (7.5 )8.0 9.0( 10.0) µm, l/w ratio (1.6 )1.7 1.9( 2.1) (n = 68), lumina ± triangular with rounded angles at first, becoming ± spherical; narrow torus present at maturity; walls light

Sheard et al.: Further contributions to the genus Rinodina 137 Fig. 9: Scanning electron micrographs of Rinodina bullata (all from Lendemer 16730). A gross morphology of thallus showing bullate nature of thalline areoles (selected representatives marked with t ) that do not form blastidia and an immature apothecium (a). B detail of apothecium (a). C detail of bullate areoles (t). Scale bars as indicated. pigmented, not ornamented (Fig. 10). Pycnidia apically positioned on bullate areoles, c. 0.05 mm diam.; conidia bacilliform, 4.5 6.0 1.0 1.5 µm. Chemistry. Spot tests, K+ yellow, C, KC, P+ faint yellow; secondary metabolite, atranorin in cortex. Ecology and distribution. Rinodina bullata is so far known only from a small region of eastern North America (throughout Pennsylvania and in southeastern Ohio) where it occurs

138 Herzogia 25 (2), 2012 Fig. 10: Mature Pachysporaria-type II ascospores of Rinodina bullata (Lendemer 17688). Arrows point to septal discs in different optical sections, tori not present. Note small subspherical lumina and very thick walls. Scale = 10 µm. in open, middle-elevation (300 900 m), xeric, secondary forests, often on ridge tops (Fig. 3C). The majority of populations of R. bullata encountered have been found on the bark of hardwoods, especially oak (Quercus) and maple (Acer), where it is rarely accompanied by other species of crustose lichens. Occasionally it has been found growing on the dead branches of hemlock (Tsuga), again only in the company of a handful of other crustose lichens. The species is most often found growing with populations of Chrismofulvea dialyta (Nyl.) Marbach, R. efflorescens, and R. excrescens and appears to belong to a community of crustose lichens that tolerate, and even thrive, in xeric regenerating hardwood forests where competition from other lichens is relatively low. Considering the relatively inconspicuous colour and size of the thallus, it is likely that the species is more widespread in the central Appalachians and Ohio Valley, and has simply been overlooked previously. Indeed Rinodina bullata is probably an example of the negative impact of collector bias on our knowledge of sterile, asexually reproducing crustose lichens (Harris & Lendemer 2010). Discussion. This species is characterized by its small Pachysporaria-type II spores, a thallus that produces atranorin in the cortex and loosely attached, bullate areoles. Morphologically Rinodina bullata is most similar to R. excrescens, a sympatric species that produces pannarin rather than atranorin, typically has a darker coloured thallus, and larger areoles (> 0.15 mm wide). While the two species are similar in their bullate thalline morphology, the thallus of R. excrescens often produces tiny blastidia along the edges of the areoles (Figs 11B D and 12B F) whereas no such blastidia are produced in R. bullata (Figs 8D F and 9A & C). Instead, the minute bullate areoles of R. bullata are loosely attached to the substrate (Figs 8D and 9C) and potentially act as lichenized diaspores, performing the same function as the blastidia in R. excrescens. When fertile, R. excrescens can easily be separated from R. bullata by its larger, Physcia-type ascospores (Sheard 2010). Mature ascospores are, however, often difficult to find in R. bullata, even in material with abundant apothecia. This may suggest that they are seasonally differentiated, and that the species primarily reproduces asexually by means of the loosely attached areoles. It may not often rely on sexual diaspores for reproduction. The new species is also similar to an undescribed taxon with the herbarium name Rinodina roborina in ASU that has similar small spores, atranorin in the cortex, and occurs in Mexico. It also possesses small, convex areoles but they ultimately merge to form a continuous, verrucate thallus and the verrucae themselves are not loosely attached. Although related by spore type this taxon is almost certainly distinct from R. bullata. Rinodina bullata also has a superficial res emblance to R. roboris var. armeriicola Matzer & Sattler, a rare taxon which grows on Armeria maritima and Calluna on small islands and headlands in the extreme southwest of England and Wales (Mayrhofer et al. 1993, Giavarini et al. 2009). This taxon has a densely granular thallus and also contains atranorin. However, it has spores belonging to Pachysporaria-type I rather than the -type II of R. bullata. The spores of R. roboris var.

Sheard et al.: Further contributions to the genus Rinodina 139 Fig. 11: Morphology of Rinodina excrescens (A D from Lendemer 17542A, E and F from Lendemer 16465). A Sectioned, narrowly attached, apothecium. B F morphological variation of the thallus illustrating typical form with bullate areoles (B D) as well as the form with marginally divided, lobate areoles (E F). Scales = 0.5 mm: A and B; 0.25 mm: C F. armeriicola are slightly shorter than those of R. roboris var. roboris (Giralt & Mayrhofer 1994, Sheard et al. 2010) which is even less likely to be confused with R. bullata because of its verrucose thallus and large apothecia up to 1.6 mm in diameter. An emended version of the key from Sheard (2010) is presented below to facilitate the identification of R. bullata.

140 Herzogia 25 (2), 2012 Fig. 12: Scanning electron micrographs of Rinodina excrescens (all from Lendemer 17549) illustrating the areoles (a) and the development of blastidia (b). A an areole (a) without blastidia. B an areole (a) with two blastidia (b) developing on the surface. C D areoles (a) developing marginal blastidia (b). E F areoles (a) developing blastidia (b) on the surface. Scale bars as indicated. 71 Thallus areolate or rimose-areolate; young spores not swollen at septum in KOH... 72 72(71) Average spore size < 18.0 µm long; lower cortex not expanded or to 30 µm deep... 72a 72 Average spore size > 18.0 µm long; lower cortex expanded to > 35 µm deep... 73 72a(72) Thalline margins of some apothecia not completely formed; spores Physcia-type, darkly pigm ented with a prominent torus, western... R. boulderensis 72a Thalline margins entire or beaded, spores Pachysporaria-type II, lightly pigmented, narrow torus only at maturity, eastern... R. bullata Additional specimens examined: U.S.A. Pennsylvania. Centre Co.: Bald Eagle State Forest, Stover Gap, 14 Sept. 2010, on Acer, J. C. Lendemer 25507 (NY); Bald Eagle State Forest, above Winkleblech Vista, 14 Sept. 2010, on Acer, J. C. Lendemer 25485 (NY). Clearfield Co.: Moshannon State Forest, Caledonia Pike at jct w/ PR 12112, 30 Aug. 2010, on Acer, J. C. Lendemer 23710 (NY). Luzerne Co.: Ricketts Glen State Park, Grand View Trail between

Sheard et al.: Further contributions to the genus Rinodina 141 Red Rock Quarry (PA 487) and Firetower at Luzerne/Columbia/Sullivan County line, 19 Sept. 2010, on Quercus, J. C. Lendemer 25716 & J. Kunsman (NY). Pike Co.: Delaware Water Gap National Recreation Area, Pocono Environmental Education Center, 18 Sept. 2005, on Pinus, J. C. Lendemer et al. 5045 (NY), J. C. Lendemer et al. 5074 (NY). Schuylkill Co.: Weiser State Forest, E end of Manhantango Mountain, N of T409/Taylorville Rd., 20 May 2009, on Quercus, J. C. Lendemer 17688 & R. C. Harris (NY). Tioga Co.: Tioga State Forest, W rim of Pine Creek Gorge, vicinity of Barbour Rock, 13 May 2009, on Quercus, J. C. Lendemer 16692-A (NY), on Betula, J. C. Lendemer 16730 (NY). Wayne Co.: Lacawac Sanctuary, ridge E of lodge, above Wallenpaupack Ledges, 29 June 2008, on Tsuga, J. C. Lendemer 12193 (NY). Rinodina ochracea Lynge, Medd. Grønland 118 (8): 188 (1937). Type: Greenland. West Greenland, Disko Island, Ritenbenk coal mine, 13 July 1871, on calcareous rock, T. M. Fries s.n. (O!, holotype; S!, isotype). Discussion. This species has been described and discussed in detail by Mayrhofer & Sheard (1988). It is characterized by its small Bicincta-type ascospores (Sheard 2010), (12.5 )13.5 15.0( 16.0) (8.5 )8.5 9.5( 10.0) µm, l/w ratio (1.4 )1.5 1.6( 1.7) (n = 100), and particularly by the pigmented bands that are located just below the apex of each cell rather than in the more usual medial position of most other species with this spore type. Rinodina luridata (Körb.) H.Mayrhofer, Scheid. & Sheard also has small, Bicincta-type spores but it is a western temperate species that does not occur in the Arctic (Sheard 2010). Both species are morphologically variable but R. ochracea typically has small thalli with convex areoles whereas R. luridata often has relatively large thalli with plane or rugose areoles. Previously, Rinodina ochracea was only known from West Greenland on calcareous substrates where it has been collected with both R. calcigena (Th.Fr.) Lynge and R. endophragmia I.M.Lamb (Mayrhofer & Sheard 1988). One of the present collections (Gould 920A), was also accompanied by R. calcigena. Due to thallus variability in the extreme Arctic envir onment the three species are not easily distinguished by their vegetative morphology (see discussion in Mayrhofer & Sheard 1988). They are, however, easily distinguished by ascospore size and structure. Rinodina calcigena has much larger Bischoffii-type ascospores (Sheard 2010) and R. endophragmia has somewhat larger Bicincta-type spores with the pigmented bands around the middle of the cells. An emended version of the key from Sheard (2010) is presented below to facilitate the identification of R. ochracea. 25(21) Spores averaging <15.0 µm long, Bicincta-type, average spore l/w ratio <1.6... 25a 25 Spores averaging >15.0 µm long, belonging to other spore types, average spore l/w ratio >1.6... 26 25a(25) Western temperate distribution in U.S.A.... R. luridata 25a High Arctic distribution... R. ochracea Specimens examined: Canada. Northwest Territories: Banks Island, Thomsen River, 20 July 1990, on calcareous pebble, W. A. Gould 920A (MIN), W. A. Gould 920B (MIN, with R. calcigena). Greenland: West Greenland, Disko Island, Asuk, 7 Aug. 1952. P. Gelting 18853b (C); Nugssuaq Peninsula, 13 July 1871, T. M. Fries s.n. (O, S); Søndre Strømfjord, 25.VI.1979, on carbonatite, V. Alstrup s.n. (CANL). Acknowledgments The authors are grateful to the curators of ASU, C, CANL, MIN, NY, O, S and UPS for the loan of specimens. J. Lendemer s work on this project was supported by NSF Award DEB-1145511. Fieldwork during which Lendemer collected material used for this study was funded in part by the City University of New York, NSF Award DEB- 110433, Southern Appalachian Botanical Society, Tallassee Foundation, and Western Pennsylvania Conservancy. Special thanks to the staff of Great Smoky Mountains National Park for facilitating the studies of J. C. Lendemer, T.

142 Herzogia 25 (2), 2012 Tønsberg, and their colleagues in that preserve. Lendemer s access to the SEM was facilitated by Lehman College of the City University of New York with logistical support from M. Baxter. T. Spribille s work in the Russian Far East was funded by the Austrian Science Foundation (FWF grant P21052-B16) and made possible by the Botanical Garden Institute of the Russian Academy of Sciences, Vladivostok, with special thanks to Lidia Yakovchenko. The stay in Japan of G. Thor 1994-1996 was granted by a post-doctoral fellowship from the Japan Society for the Promotion of Science (JSPS) and a fellowship for Priority area research in Japan from the JSPS. The field trip in Korea by G. Thor 2006 was financially supported by a grant to K. H. Moon (no. 052-052-040) from the Core Environmental Technology Development Project for Next Generation funded by the Ministry of Environment of the Korean Government. References Culberson, W. L. 1972. Disjunctive distributions in the lichen forming fungi. Annals of the Missouri Botanical Garden 59: 165 173. Galanina, I. A., Yakovchenko, L. S., Tsarenko, N. A. & Spribille, T. 2011. Notes on Rinodina excrescens in the Russian Far East (Physciaceae, lichenized Ascomycota). Herzogia 24: 59 64. Giavarini, V., James, P. W. & Purvis. O. W. 2009. Rinodina (Ach.) Gray (1821). In: Smith, C. W. et al. (eds.). The lichens of Great Britain and Ireland. Pp. 812 826. London: British Lichen Society and The Natural History Museum. Giralt, M. & Mayrhofer, H. 1994. Four corticolous species of the genus Rinodina (lichenized Ascomycetes, Physciaceae) containing atranorin in southern Europe and adjacent regions. Nova Hedwigia 59: 129 142. Harris, R. C. & Lendemer, J. C. 2010. A review of Lecania croatica (syn. Catillaria croatica) in North America. Opuscula Philolichenum 8: 41 49. Kurokawa, S. 2006. Phytogeographical elements in the lichen flora of Japan. Journal of the Hattori Botanical Laboratory 100: 721 738. Kurokawa, S. & Kashiwadani, H. 2006. Checklist of Japanese lichens and allied fungi. Tokyo: National Science Museums Monographs No. 33. Lendemer, J. C. 2011. A standardized morphological terminology and descriptive scheme for Lepraria (Stereocaulaceae). Lichenologist 43: 379 399. Lendemer, J. C. & Elix, J. A. 2010. Two new species of Chrysothrix from eastern North America. Opuscula Philolichenum 8: 51 58. Lendemer, J. C., Sheard, J. W., Thor, G. & Tønsberg, T. 2012. Rinodina chrysidiata, a new species from far eastern Asia and the Appalachian Mountains of North America. Lichenologist 44: 179 187. Mayrhofer, H. & Sheard, J. W. 1988. Four notable saxicolous species of the lichenized Ascomycete genus Rinodina from the Arctic. Bryologist 91: 106 112. Mayrhofer, H., Matzer, M. & Sattler, J. 1993. A revision of the Atlantic-Mediterranean Rinodina beccariana and related taxa (lichenized Ascomycetes, Physciaceae). Nova Hedwigia 57: 281 304. Sheard, J. W. 2010. The lichen genus Rinodina (Ach.) Gray (Lecanoromycetidae, Physciaceae) in North America, north of Mexico. Ottawa, Ontario: NRC Research Press. Sheard, J. W. 2011. Rinodina pityrea discovered for the first time from North America. Bibliotheca Lichenologica 106: 291 295. Sheard, J. W. & Mayrhofer, H. 2002. New species of Rinodina (Physciaceae, lichenized Ascomycetes) from Western North America. Bryologist 105: 645 672. Sheard, J. W., Knudsen, K., Mayrhofer, H. & Morse, C. A. 2011. Three new species of Rinodina (Physciaceae) and a new record from North America. Bryologist 114: 453 465. Sheard, J. W., Lendemer, J. C. & Tripp, E. A. 2008. Buellia japonica (Physciaceae), a new lichen record for North America. Bryologist 111: 124 127. Sheard, J., Tønsberg, T. & Johnsen, J. I. 2010. Rinodina orculata and R. roboris new to Fennoscandia. Graphis Scripta 22: 43 46. Tønsberg, T. 1992. The sorediate and isidiate, corticolous, crustose lichens of Norway. Sommerfeltia 14: 1 331. Yoshimura, I. 1968. The phytogeographical relationships between the Japanese and North American species of Cladonia. Journal of the Hattori Botanical Laboratory 31: 227 246. Manuscript accepted: 16 October 2012.

Sheard et al.: Further contributions to the genus Rinodina 143 Addresses of the authors John W. Sheard, Dept. of Biology, University of Saskatchewan, 112 Science Place, Saskatoon, SK S7N 5E2, Canada. E-mail: john.sheard@usask.ca James C. Lendemer, Institute of Systematic Botany, The New York Botanical Garden, Bronx, New York, 10458-5126, U.S.A. E-mail: jlendemer@nybg.org Toby Spribille, Institute of Plant Sciences, University of Graz, Holteigasse 6, 8010 Graz, Austria; current address: Division of Biological Sciences, University of Montana, 32 Campus Drive, Missoula, MT 59812, U.S.A. E-mail: toby.spribille@mso.umt.edu Göran Thor, Department of Ecology, Swedish University of Agricultural Sciences, P.O. Box 7044, 750 07, Uppsala, Sweden. E-mail: goran.thor@slu.se Tor Tønsberg, Museum of Natural History, University of Bergen, Allégaten 41, Postboks 7800, 5020 Bergen, Norway. E-mail: tor.tonsberg@um.uib.no