Opuscula Philolichenum, 14: 139-143. 2015. *pdf effectively published online 20November2015 via (http://sweetgum.nybg.org/philolichenum/)

Protoparmelia ryaniana is a synonym of Lecanora verrucariicola, which belongs in the genus Miriquidica ()

1 2 3 KERRY KNUDSEN , JANA KOCOURKOVÁ & TIM WHEELER

ABSTRACT. – Lecanora verrucariicola was described in 2004 by Bruce Ryan based on a holotype collected on Catalina Island in southern California. In 2008, the taxon was described again as Protoparmelia ryaniana from a collection from Baja California in Mexico, with the holotype of L. verrucariicola cited as a paratype. We show that Protoparmelia ryaniana is a superfluous and illegitimate synonym of L. verrucariicola. The nomenclature error is corrected, Protoparmelia ryaniana is made a synonym of Lecanora verrucariicola, and Lecanora verrucariicola is transferred to Miriquidica. KEYWORDS. – Baja California, Channel Islands, Miriquidica mexicana, Pacific Plate Bioregion, nomenclature.

INTRODUCTION

The authors are currently researching a book on the lichenized fungi of the eight islands of southern California that comprise the Channel Islands. An important part of this study is a review of rare fungal species that warrant monitoring in the future by the various relevant government agencies and NGOs responsible for the management of the islands. One rare species is Lecanora verrucariicola B.D. Ryan which was described from a holotype collected on Catalina Island and from several paratypes from Catalina Island and Santa Rosa Island, as well as from Baja California in Mexico (Ryan et al. 2004). The first author never found any type material during visits to ASU where they were published as being deposited. Later, because we found no specimens which had been identified as L. verrucariicola since the original publication of the name, and we had not collected any specimens that we could positively identify as L. verrucariicola, we assumed it was rare and potentially even had been extirpated in this short timeframe. When we began the review of the rare species of Lecanora from the islands, we found that the holotype and paratypes of L. verrucariicola were still not digitized (CNALH 2015). Further study led us to recognize that the same material was later described a second time under the name Protoparmelia ryaniana van den Boom, Sipman & Elix. Here we recognize P. ryaniana as a superfluous renaming of L. verrucariicola, discuss the species, and transfer the name to Miriquidica based on recently published molecular data.

MATERIALS AND METHODS

This study is based on a critical analysis of the literature describing the taxon, the herbarium records available online at CNALH and UCR, and an examination of specimens from B and UCR using hand sections mounted and measured in water. Photos were taken with a Pentax K3 DSLR, mounted on a Stackshot rail, and combined in Helicon Focus.

1KERRY KNUDSEN – 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] 2 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] 3 TIM WHEELER – Organismal Biology, Evolution, and Ecology Program, Division of Biological Sciences, University of Montana, Missoula, Montana 59801 – e-mail: [email protected]

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Figure 1, known geographic distribution (left) and morphology (scale = 2.0 mm, field image captured by T. Wheeler) of Miriquidica verrucariicola.

RESULTS

During the first author’s visits to ASU between 2003 and 2008 to study Acarosporaceae as well as newly described species from the Sonoran region, he could not locate the holotype or paratypes of Lecanora verrucariicola as well as several other taxa such as Adelolecia sonorae Hertel, Lecanora crassithallina van den Boom and Toninia nashii Timdal that were stated to have been deposited at ASU. At that time it was assumed that they were missing and might have still been on loan to authors of the various treatments prepared for the Lichen Flora of the Greater Sonoran Desert Region. When we began our study of Lecanora species on the Channel Islands, no specimens of L. verrucariicola had yet been databased or digitized at ASU or any other herbaria (CNALH 2015). We were perplexed since the species was described by Bruce Ryan and the type collection was made by Thomas Nash, the material should have been in ASU where they both had worked. Eventually, in rereading the last paragraph of the discussion of Lecanora verrucariicola (Ryan et al. 2004: 281) we noticed an apparent typographic error: the species was referred to as “Lecanora lichenicola” (Ryan et al. 2004). We then connected that statement with one in the discussion of Protoparmelia ryaniana where it was stated that some herbarium specimens may have be annotated as “Lecania lichenicola” or “Lecanora lichenicola” on page 393 (van den Boom et al. 2008: 393). Using the type collection numbers cited for L. verrucariicola, a search of CNALH quickly showed that while some paratypes were missing, the holotype of L. verrucariicola and one paratype at ASU had been annotated by P.G.G. van den Boom as P. ryaniana. In fact the holotype of L. verrucariicola had been listed as a paratype of P. ryaniana in the protologue of the latter name (van den Boom et al. 2008). The mystery of the missing holotype of L. verrucariicola was solved. It was at ASU filed under the name P. ryaniana. The holotype was originally determined as an unknown Lecania species by Thomas Nash, there is no annotation by Bruce Ryan, and it was later annotated by van den Boom as P. ryaniana at a later unspecified date (Walter Fertig, pers. comm.). Based on our study of the southern California islands and the adjacent coast, Protoparmelia ryaniana is a good species. We have made additional collections of it, extending its range to the central California coast as well as to West Anacapa Island, Santa Cruz Island, San Nicolas Island, and San Clemente Island (Knudsen & Kocourková 2009, 2010, 2012; UCR 2015). A comparative literary analysis of the original description of Lecanora verrucariicola and the description of P. ryaniana, and an

140 examination of the specimens of P. ryaniana made clear that Bruce Ryan had originally described P. ryaniana under the name L. verrucariicola. The only major difference between the two descriptions is that L. verrucariicola was described as containing an unknown (“CE-71; Pale yellow-gray phenolic 3-A, 5-B, 5-C”) and being spot test negative while P. ryaniana was described as containing lobaric acid and having a KC+ violet medulla. The RFI of the unknown described from L. verrucariicola matches that of lobaric acid (J.A. Elix, pers. comm.). The first specimen we identified as P. ryaniana was spot test negative and was verified as having lobaric acid (Lendemer pers. comm.; TLC in Solvent C). Once one collection was verified, we could easily identify P. ryaniana through standard microscopy and through observation of its parasitic habit in the field without further ascertaining its secondary metabolites. We had been collecting L. verrucariicola all along and it was not rare on the Channel Islands. Through a chain of mistakes, probably initiated by the death of Bruce Ryan, van den Boom and the principal editor of the Sonoran Lichen Flora, Thomas Nash must not have realized that L. verrucariicola was redescribed as P. ryaniana. Since the description of P. ryaniana included the type of an earlier name, P. ryaniana is an illegitimate and superfluous renaming of L. verrucariicola. The question remains as to whether Lecanora verrucariicola is actually a Protoparmelia. In the case of this taxon, a recent phylogenetic study found that P. ryaniana does not belong to Protoparmelia s. str. (Singh et al. 2015). The specimens used for the molecular analyses in that study were actually collections made by the authors of the present contribution. Singh et al. (2015) recovered the taxon as sister to species of the genus Miriquidica. Although further study of the relationship between the taxon and Miriquidica is needed, it is clear based on the available data the species does not belong to Protoparmelia. To correct the nomenclatural error and the confusion between L. verrucariicola and P. ryaniana, both being listed as valid species in the North America checklist (Esslinger 2015) and in the California lichen checklist (Tucker 2013), we here transfer L. verrucariicola to Miriquidica and treat P. ryaniana as an illegitimate synonym (McNeil et al. 2013). Ironically, some specimens of this taxon from Baja California were originally also treated as Miriquidica mexicana Rambold, Sipman & Hertel in the treatment of that taxon published in the Lichen Flora of the Greater Sonoran Desert Region (Nash et al. 2004). Although the holotype of M. mexicana was previously shown to be a synonym of M. scotopholis (Tuck.) B.D. Ryan & Timdal (Lendemer & Knudsen 2008), those specimens with lobaric acid and lacking miriquidic acid cited by Nash et al. (2004) as M. mexicana were later treated as P. ryaniana (van den Boom et al. 2008).

THE SPECIES

Miriquidica verrucariicola (B.D. Ryan) K. Knudsen & Kocourk., comb. nov. Mycobank #815090.

≡ Lecanora verrucariicola B.D. Ryan, Lichen Flora of the Greater Sonoran Desert Region 2: 280. 2004. TYPE: U.S.A. CALIFORNIA: LOS ANGELES CO.: Santa Catalina Island, Parsons Landing adjacent to an old quarry, 33°28′30″N 118°32′45″W, 20 m, on rock, 29.xii.1993, T.H. Nash 32184 (ASU[n.v.], holotype; hb. van de Boom[n.v.], isotype). Fig. 1.

= Protoparmelia ryaniana van den Boom, Sipman & Elix nom. illeg., syn. nov. TYPE: MEXICO. BAJA CALIFORNIA: Punta Santa Tomas, c. 30 km SW of Ensenada, 31°33′N 116°42′W, 80 m, on acidic rock, 1989, H. Sipman 24745 (B!, holotype; hb. v.d. Boom[n.v.], isotype)

DESCRIPTION. — For excellent descriptions of this taxon the reader should refer to Ryan et al. (2004) under the name Lecanora verrucariicola and van den Boom et. al (2008) under the name Protoparmelia ryaniana.

SECONDARY METABOLITES. — Lobaric acid (major or minor), oxolobaric acid (major), norlobaric acid (minor) produced in medulla (Ryan et. al. 2004, van den Boom et al. 2008). Spot tests of the medulla are either entirely negative (Ryan et al. 2004) or KC+ violet and distinctly UV+ blue-white.

ECOLOGY AND SUBSTRATE. — In coastal habitats at low elevations on non-calcareous rock, parasitic on Dimelaena radiata (Tuck.) Hale & Culb., Tephromela species, and Verrucaria subdivisa Breuss, rarely other crustose species, or independent, in mixed saxicolous communities.

141 DISTRIBUTION. — Endemic to the Pacific Plate Lichen Bioregion: along the Pacific coast from Baja California in Mexico to San Simeon in central California and on six of the Channel Islands (Anacapa, Catalina, San Clemente, Santa Cruz, San Nicolas, and Santa Rosa). (Ryan et al, 2004; van den Boom et al. 2008; Knudsen & Kocourková 2009, 2010, 2012; CNALH 2015; UCR 2015).

Selected specimens examined. — U.S.A. CALIFORNIA. LOS ANGELES CO.: San Clemente Island, west end, 32°57′53.5″N 118°33′51″W, elevation unknown, on Dimelaena radiata and basalt, 18.iv.2014, K. Knudsen 16528 & E. Howe (UCR). SAN DIEGO CO.: Point Loma Ecological Preserve, navy property S of McClelland and the waste management facility, E of cemetery, 32°42′46″N 117°15′37″W, 72 m., on granite and other crustose , 15.iv.2007, K. Knudsen 8424 (UCR). SAN LUIS OBISPO CO.: San Simeon, San Simeon State Park, solitary boulder on rim of canyon, 35°35′49″N 121°6′42″W, 141 m, on Dimelaena radiata and shale, 11.iv.2010, K. Knudsen 12146 (UCR). SANTA BARBARA CO.: Santa Cruz Island, cliff above Smuggler’s Road, 34°2′50.5″N 119°33′33.3″W, 9 m, on basalt, 30.vii.2012, K. Knudsen 14999 & J. Kocourková (UCR); Santa Rosa Island, South Point, 33°54′27″N 120°7′34″W, 198 m, on sandstone, 14.vi.2009, K. Knudsen 11439 & S. Chaney (NY, UCR). VENTURA CO.: San Nicolas Island, slope above L Canyon, 33°14′23.7″N 119°27′04.6″W, 105 m, lichenicolous on Dimelaena radiata on quartzitic cobble of Pleistocene beach terrace, 24.vii.2015, T. Wheeler 7115 & K. Knudsen (hb Wheeler).

ACKNOWLEDGEMENTS

The work of Jana Kocourková and 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. We thank Shirley Tucker (SBBG) for supporting our research of the lichen biota of the Channel Islands and the trip of Tim Wheeler and Kerry Knudsen to San Nicolas Island. For special help we thank William Fertig (ASU), Harrie Sipman (B), Imke Schmitt (Germany) and Jack Elix (CANB).

LITERATURE CITED

CNALH. 2015. Consortium of North American Lichen Herbaria. http://lichenportal.org/portal/ [Accessed June, 2015.] Esslinger, T.E. 2015. A cumulative checklist for the lichen-forming, lichenicolous and allied fungi of the continental United States and Canada. North Dakota State University: http://www.ndsu.edu/pubweb/~esslinge/chcklst/chcklst7.htm (First Posted 1 December 1997, Most Recent Version (#20) 19 April 2015), Fargo, North Dakota. Knudsen, K. and J. Kocourková. 2009. The lichens and lichenicolous fungi of the Point Loma Conservation Area, San Diego, USA. Evansia 26: 24–29. Knudsen, K. and J. Kocourková. 2010. Lichens and Lichenicolous Fungi of West Anacapa Island, Channel Islands National Park. Crossosoma 36(2): 32–49. Knudsen, K. and J. Kocourková. 2012. The Annotated Checklist of Lichens, Lichenicolous and Allied Fungi of Channel Islands National Park. Opuscula Philolichenum 11: 145–302. Lendemer, J.C. and K. Knudsen. 2008. Studies in lichens and lichenicolous fungi: further notes on North American taxa. Mycotaxon 103: 75–86. McNeil, J., F.R. Barrie, W.R. Buck, V. Demoulin, W. Greuter, D.L. Hawksworth, P.S. Herendeen, S. Knapp, K. Marhold, J. Prado, W.F. Prudhomme, G.F. Smith and N.J. Turland. 2012. International Code of Nomenclature for algae, fungi, and plants (Melbourne Code). Regnum Vegetabile 154. Germany: Koeltz Scientific Books. Nash III, T.H., C. Kainz, L. Zedd, B.D. Ryan and G. Rambold. 2004. Miriquidica. In: T.H. Nash III, B.D. Ryan, P. Diederich, C. Gries, and F. Bungartz (eds.): Lichen Flora of the Greater Sonoran Desert Region, Vol. 2. Arizona: Lichens Unlimited, Arizona State University, pp. 361–363. Ryan, B.D., H.T. Lumbsch, M.I. Messuti, C. Printzen, L. Śliwa and T.H. Nash III. 2004. Lecanora. In: T.H. Nash III, B.D. Ryan, P. Diederich, C. Gries, F. Bungartz (eds.): Lichen Flora of the Greater Sonoran Desert Region, Vol. 2. Arizona: Lichens Unlimited, Arizona State University, pp. 176–286. Singh, G., F. Dal Grande, P.K. Divakar, J. Otto, S.D. Leavitt, K. Szczeparska, A. Crespo, V.J. Rico, A. Aptroot, M.E. Cáceres, H.T. Lumbsch and I. Schmitt. 2015. Coalescent-Based Species Delimitation Approach Uncovers High Cryptic Diversity in the Cosmopolitan Lichen-Forming Genus Protoparmelia (, ). PLoS ONE 10(5): e0124625. Tucker, S.C. 2013. Catalog of Lichens, Lichenicoles and Allied Fungi (second revision). Constancea 85. http://ucjeps.berkeley.edu/constancea/85/tucker.html UCR Herbarium. 2015. http://ucr.ucr.edu/lichensflat_index.php [Accessed June, 2015].

142 van den Boom, P.P.G., H. Sipman and J.A. Elix. 2008[2007]. Protoparmelia. In: T.H. Nash III, C. Gries and F. Bungartz (eds.). Lichen Flora of the Greater Sonoran Desert Region. Volume 3.Arizona: Lichens Unlimited, Arizona State University, pp. 392–393

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