Opuscula Philolichenum, 18: 11-16. 2019. *pdf effectively published online 8February2019 via (http://sweetgum.nybg.org/philolichenum/)

The first confirmed report of bullata from North America

SAMUEL R. BRINKER1 AND KERRY KNUDSEN2

ABSTRACT. – The first confirmed North American record of is reported from a granite cliff in southeastern , Canada. The ecology and conservation status of the population is discussed. A taxonomic account of the species, including a description based on the newly collected material, is provided.

KEYWORDS. – Acarosporaceae, biodiversity, , cliffs.

INTRODUCTION

Acarospora bullata Anzi was described from the mountains of Lombardy in Italy and reported from California and Iran (Magnusson 1929). Based on Magnusson’s identification of a H.E. Hasse specimen from California as A. bullata, the species was reported from the Sonoran Desert region (Knudsen 2007). It was also reported from South America (Knudsen et al. 2008). A later molecular phylogenetic analysis showed that specimens identified as A. bullata from western North America were conspecific with A. rosulata (Th. Fr.) H. Magn., which was previously known only from Norway (Knudsen et al. 2010). The specimens from South America published as A. bullata were then revised to A. subcastanea (Nyl.) Hue (Knudsen 2012). As a result, A. bullata was no longer recognized as occurring in the Americas (Esslinger 2018). Here we present the first confirmed record of A. bullata for North America.

MATERIALS AND METHODS

The specimens examined were studied with standard microscopy and chemical spot tests for lichens following Brodo et al. (2001). The newly collected specimen was compared with an isotype deposited in the University of Helsinki herbarium (H). The map was generated with ESRI ArcMap 10.3.1. Field photographs by the first author were taken with a Canon EOS 7D body with a Canon EF 16-35 F4L and EF 100/2.8 macro lens.

RESULTS AND DISCUSSION

Cliffs support distinct flora and serve as refugia for some lichens, providing unique microhabitats, particularly for rare species. Due to their vertical inclination and gravitational stresses, the accretion of water and soil are limited on cliffs, thereby restricting vegetation establishment and reducing competition from vascular plants (Clebsch & Walker 1988, Farris 1998, Haig et al. 1999). Plant growth can be further limited in exposed situations by wind, frost wedging, and rapid fluctuations in temperature, or low light levels in sheltered, shaded situations (Cohen et al. 2015, MDNR 2003).

1SAMUEL R. BRINKER – Ontario Ministry of Natural Resources and Forestry, Natural Heritage Information Centre, 300 Water St. Peterborough, Ontario K9J 3C7, Canada. – e-mail: [email protected] 2KERRY KNUDSEN – Department of Ecology, Faculty of Environmental Sciences, Czech University of Life Sciences, Prague, Kamýcká 129, Praha 6 - Suchdol, CZ–165 00, Czech Republic. – e-mail: [email protected]

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Figure 1. General location of collecting site at Mazinaw cliff in southeastern Ontario, Canada.

Cliffs often provide stable environments, by escaping the deleterious effects of natural disturbances such as fire, and anthropogenic pressures such as logging due to inaccessibility (Larson & Kelly 1991, Krajick 1999, Larson et al. 2000). The presence of intact, remnant old-growth Thuja occidentalis-dominated (with individual stems up to 1,032 years old) limestone cliff and talus ecosystems in southern Ontario, such as along the , has been well documented (Larson & Kelly 1991, Kelly et al. 1992). In glaciated regions, cliffs were some of the first exposed substrates to become colonized post-glaciation, and likely provided early, non-competitive pathways for the movement of species in otherwise unsuitable environments from glacial refugia (Braun 1928, Butters & Abbe 1953, Walker 1978). Indeed, cliffs have been shown to support disjunct arctic-alpine or Western Cordilleran species in Ontario that survive as relicts of a presumably wider or more continuous distribution (e.g., Brayshaw 1964, Brinker 2017, Reznicek 1972). The floras of calcareous cliffs have been relatively well studied and shown to often have a preponderance of rare vascular plants and lichens (e.g., Boggess et al. 2017, Gilbert et al. 1982, Matthes et al. 2000, Müller et al. 2004, Vogler & Reisch 2011). But there is a paucity of literature on diversity or rare lichens associated with acidic cliffs. This is perhaps not surprising given their comparatively lower weatherability, impermeability and productivity (Shure 1999). Lendemer (2011) highlighted the importance of acidic rock overhang communities in supporting rare lichens in Pennsylvania, but this habitat harbours different lichen communities than exposed acidic cliffs. Acarospora bullata was collected during targeted lichen inventories of a shoreline cliff by the first author in Bon Echo Provincial Park in central Frontenac County, Ontario (Fig. 1). The park supports a provincially significant 100-meter-high west-facing granite rock face adjacent to the 145-meter-deep waters of Mazinaw Lake (Fig. 2A). Known as the Mazinaw Cliff, it is the largest in the region and forms part of a

12 protected natural and cultural heritage reserve long revered for its unusual cliff flora, including stunted old- growth Thuja occidentalis L. more than 900 years old (Dougan & Associates 1995). The park is dominated by metaplutonic rocks including gabbro and granite with minor occurrences of metasedimentary and metavolcanic rocks (Easton 1994). The cliff feature is part of a two kilometer long reverse thrust fault zone extending north and south of Mazinaw Lake containing exposures of fine-grained Abinger granite and mafic diabase dikes resulting in a complex assemblage of topographic and geologic exposures making the rock face quite heterogeneous (Easton 1994). The 100-meter-tall granitic Mazinaw Cliff is the central tourist attraction at Bon Echo Provincial Park, which received 208,212 visitors in 2017, and is the fifth most used provincial park in terms of camping in the province. The cliff protects one of the largest collections of prehistoric Indigenous pictographs in North America and has over 150 recognized climbing routes on its face with a rich climbing history dating back to 1956 (OMNR 1991; C. Richards & L. Roach, pers. comm.). A past study of authorized and unauthorized climbing showed increased vegetation damage and declines in lichen abundance along climbing routes on Mazinaw Cliff (Dougan & Associates 1995), leading to restrictions on certain routes to promote resource protection. Despite these actions, continuing concern over impacts from climbing and more recently from recreational boating along the cliff/shore interface has spurred interest in more thorough inventories of the park’s biodiversity. The xerothermic granite cliff on Mazinaw Lake is unique in the region and supports other rare saxicolous species such as Acarospora sinopica (Wahlenb.) Körb., Lecanora epanora (Ach.) Ach, and Lichinella nigritella (Lettau) P.P. Moreno & Egea also documented by the first author with vouchers housed at the Canadian Museum of Nature (CANL). The negative impacts of climbing on cryptogam cliff communities is well established (e.g., Adams & Zniewski 2012, Farris 1998, Nuzzo 1996, Thiel & Spribille 2007) although large gaps in knowledge still exist (Holzschuh 2016). It is imperative that active monitoring and enforcement occurs to safeguard the rare cliff flora that is the focus of this paper. We suggest that any future planned or proposed climbing routes should be properly surveyed for rare lichens before they are approved. Additional study is required to better understand the status of A. bullata in North America. Its presence at the site in Ontario suggests it may occur elsewhere that xerothermic granite cliffs exist. This study illustrates the importance of granitic cliffs to support rare lichens and that knowledge of their biodiversity is still incomplete.

TAXONOMIC SECTION

Acarospora bullata Anzi, Atti della Società Italiana di Scienze Naturali 11(4): 165. 1868. TYPE: ITALY. LOMBARDY: Valfura, village of Uzza and St. Antonio, 1400 m, on mica schist, 1868, M. Anzi s.n. = Lich. Langob. Exs. 532 (MRSN[n.v.]?, holotype; H!, M[digital image!], isotypes).

DESCRIPTION. – Thallus determinate, up to 5 cm in diameter, of clustered bullate areoles, 0.5–2.0 mm wide, up to 0.8 mm high, in center with outer lobes discontinuous or surrounding whole thallus, lobes convex, ca. 2 mm long, usually 1 mm wide, sometimes imbricate, tips usually fan-shaped, fibrillate. Upper surface light brown to olive, epruinose, smooth to rugulose. Epicotex thin, less than 10 µm thick. Upper cortex 45–60 µm thick, upper layer brown, lower layer hyaline, cells distinct 2.0–4.0 µm in diameter, mostly round. Algal layer continuous, uninterrupted by hyphal bundles, 70–120 µm thick, algal cells 10–16 µm in diam. Medulla up to 600 µm thick of hyphae 3–4 µm wide. Apothecia usually one per areole, sometimes 2 or 3, immersed when young, eventually expanding up to 1 mm wide, with thalline margin, older disc brown and distinctly rugulose or gyrose. Parathecium ca. 20 µm wide, not distinctly expanded around the disc. Epihymenium reddish-brown ca. 15 µm high. Hymenium 90–140 µm high, paraphyses 1.5–2.0 µm wide at midlevel, apices expanded 3–4 µm, hymenial gel IKI+ dark blue or red (euamyloid or hemiamyloid). Asci 70–80 × 20–25 µm, ascospores several hundred per ascus, 3–5 × 1.5–2.0 µm. Subhymeniun IKI+ blue thin 10–15 µm tall. Hypothecium thin, ca.10 µm tall.

CHEMISTRY. – Gyrophoric acid. Spot tests (cortex): C+ pinkish-red in thin section, KC+ pinkish- red in thin section, K-, P-, UV-. Medulla C-, KC-, K-, P-, UV-.

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Figure 2. A, portion of Mazinaw cliff on Upper Mazinaw Lake in Bon Echo Provincial Park where A. bullata was discovered. B, detail of A. bullata showing aerolate thallus, elongated outer lobes and rugulose apothecia. C, cluster of A. bullata thalli on vertical rock face. Note that both B and C are field photos and were taken without scales.

ECOLOGY AND DISTRIBUTION. – The species occurs on steeply inclined surfaces on mica schist and various types of siliceous rock. It is widespread but not common at higher elevations in the Alps in Europe (Austria, France, Italy, Switzerland; Magnusson 1929, Nimis et al. 2018). The occurrence is here confirmed for the first time from North America (Canada). Reports from Iran or Asia probably represent a different taxon and need further study (Knudsen et. al. 2010).

14 TAXONOMIC DISCUSSION. – Acarospora bullata is recognized by its determinate thallus with fan- shaped lobes (Fig. 2B & 2C), KC+ positive reaction for gyrophoric acid in the cortex, rugulose apothecial discs, and occurrence on non-calcareous rock. In North America, it is most similar to A. rosulata. Acarospora rosulata differs from A. bullata in lacking large determinate thalli with fan-shaped lobes and distinctly rugulose apothecial discs and can be easily identified from the description and picture in Knudsen et al. (2010). Specimens identified as A. bullata in North American herbaria can be easily verified from the description and image in this paper.

Additional specimen examined. – CANADA. ONTARIO. FRONTENAC CO.: Bon Echo Provincial Park, east shore of Upper Mazinaw Lake, ~270 m., exposed W-facing cliff along lakeshore, 30.v.2018, on granite above epilittoral zone, S.R. Brinker 6509 (hb. Kocourková & Knudsen).

ACKNOWLEDGEMENTS

The manuscript benefited from the helpful comments of James Lendemer, and an anonymous reviewer. We thank the curators of the University of Helinski herbarium (H) for providing a loan of an isotype of A. bullata. We gratefully acknowledge Mike Burrell, Travis Cameron, Clark Richards and Lisa Roach for assisting with fieldwork and providing logistical support. A permit to collect in provincial parks was provided by the Ministry of Natural Resources and Forestry. We thank P.L. Nimis (Italy) for information on Anzi’s herbaium. The work of Kerry Knudsen 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.

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