Opuscula Philolichenum, 11: 313-321. 2012. *pdf effectively published online 26November2012 via (http://sweetgum.nybg.org/philolichenum/)

Studes in and lichenicolous fungi – no. 16

1 2 JAMES C. LENDEMER AND RICHARD C. HARRIS

ABSTRACT. – The type material of Bacidia leucampyx was examined and the name found to be conspecific with B. schweinitzii. It is formally placed in synonymy with B. schweinitzii. Examination of the type and original material of Pertusaria shenandoensis revealed the name is based on immature specimens of Pertusaria plittiana. The chemical variability of P. plittiana is discussed and the name P. shenandoensis is placed in synonymy with P. plittiana. The following taxa are reported for the first time from North America: Arthonia colombiana (on from Maine, U.S.A. and Quebec, Canada), A. coronata (on Cladonia digitata, Flavoparmelia caperata, and Parmeliopsis hyperopta in Maine, U.S.A. and Quebec, Canada), and Rosellinula haplospora (on cinerea from the Yukon Territory, Canada).

INTRODUCTION

During the course of routine organizational work in the herbarium of The New York Botanical Garden (NY) in 2012 we uncovered new information relating to two poorly understood and overlooked names of lichens described from eastern North America: Bacidia leucampyx (Tuck.) Dodge and Pertusaria shenandoensis Dibben. Review of the literature and loans of material from other herbaria allowed us to finally resolve the identity of these names. We present the results of our findings in the present contribution, which continues the series which ended last with the publication of Knudsen et al. (2011). In addition to the resolution of two little used names we report three lichenicolous fungi for the first time from North America, based on material collected during recent fieldwork.

MATERIAL AND METHODS

This study is based primarily on specimens collected by the authors and deposited in the herbarium of the New York Botanical Garden (NY). In addition to this material, specimens were borrowed on loan from DUKE, FH, HBG and US. Chemical and morphological investigations were carried out using the methods outlined in detail by Lendemer et al. (2008).

TAXONOMIC SECTION

1. Arthonia colombiana Etayo, Bibliotheca Lichenologica 84: 23. 2002. TYPE: COLOMBIA: Nariño, Pasto, Serranía de Morasurco, 3000-3300 m, en Cladonia sp, 25.vii.1998, J. Etayo 15934 (COL[n.v.], holotype; hb. Etayo[n.v.], isotype).

DISCUSSION. – The lichenicolous species Arthonia colombiana (fig. 1) is here reported for the first time from North America. It is the only lichenicolous Arthonia with orange ascomatal pigment (KOH+ red purple) reported to occur on Cladonia squamules. The ascospores are 1-septate, soon becoming dark and warted, and 13-16 x 5-6 µm in size (12-15 x 4.5-5.5 µm fide Etayo (2002)). In North American material the ascomata are orangish to dark brown and the host squamules appear bleached around the ascomata apparently due to the death of the Cladonia photobiont and replacement by scattered filaments of

1 JAMES C. LENDEMER – Institute of Systematic Botany, The New York Botanical Garden, Bronx NY 10458-5126, U.S.A. – e-mail: [email protected] 2 RICHARD C. HARRIS – Institute of Systematic Botany, The New York Botanical Garden, Bronx NY 10458- 5126, U.S.A. – e-mail: [email protected]

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Figure 1, Arthonia colombiana (Buck 53854: A and B; Harris 57525: A[inset], C and D). A-B, infection on podetia (A) and squamule (B) of a host thallus of Cladonia squamosa, note the yellowish-orange discoloration around the apothecia where A. colombiana appears to form a thallus. A[inset], ascospores. C- D, cross section of apothecium mounted in water (C) and then flooded with KOH (D). Scales = 0.25 mm: A and B; 100 μm: C and D; 20 μm A[inset].

Trentepohlia. Scottish material of this taxon was also found to be associated with Trentepohlia and as such A. colombiana was proposed to be a “lichenicolous ” by Coppins and Aptroot (2009). Arthonia colombiana was described from unidentified Cladonia species occurring at high elevations in Colombia (Etayo 2002). It has also been reported from Scotland where it was found on C. pyxidata (L.) Hoffm. and C. squamosa Hoffm. (Coppins & Aptroot 2009). The North American populations we have encountered are most commonly on C. squamosa, although the species has also been found on C. ochrochlora Flörke. Based on the available material, and the previous literature reports, it appears that this species has an oceanic distribution in the Northern Hemisphere (e.g., Clerc & May 2007, Degelius 1935, Jørgensen 1996).

Specimens examined. – CANADA. QUEBEC. M.R.C. DE LA HAUTE-GASPÉSIE: Parc national de la Gaspésie, along Sentier de la Grande Traversée Ouest, from parking for Chute Sainte-Anne to ca. 1.5 km along Rivière Sainte-Anne, Thuja-Abies balsamea forest along river, 4.vii.2012, on Cladonia squamosa, R.C. Harris 57651 (NY). U.S.A. MAINE. HANCOCK CO.: Rt. 182, west of Rt. 1, trail to Tunk Mountain, near Salmon Pond and Mud Pond, Acer rubrum, Picea rubens, Tsuga canadensis, 97 m, 1.ix.2001, on C. squamosa, M.S. Cole 9254 & D. Hawksworth (NY). WASHINGTON CO.: Town of Beals, Great Wass Island Preserve, Picea rubens-Abies balsamea coastal forest, 29.vi.2012, on C. squamosa, R.C. Harris 57522 (NY), on C. ochrochlora, R.C. Harris 57525 (NY); Town of Cutler, Cutler Public Reserve Land, Coastal Trail, SE of ME 191, ca. 3 mi NE of village of Cutler, mixed conifer-hardwood forest, 4.vii.2008, on C. squamosa, W.R. Buck 53854 (NY).

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Figure 2, Arthonia coronata on Parmeliopsis hyperopta (all from Harris 57546). A-B, infection developed on soralia of the host thallus. C, squash mount of apothecium and associated thallus tissue from the host. D, asci, ascospores, fragmented spines mounted in IKI. Scales = 0.25 mm: A and B; 100 μm: C; 20 μm: D.

2. Arthonia coronata Etayo, Bull. Soc. Linn. Provence 47: 95. 1996. TYPE: FRANCE: Fôret Communale de St.-Pée-sur-Nivelle, St.-Pée, 6.iii.1994, sobre F. caperata en viejos robles de fonde de valle, J. Etayo 12603 (MA-Lich[n.v.], holotype; hb. Etayo[n.v.], isotype).

DISCUSSION. – Arthonia coronata (fig. 2), a distinctly odd lichenicolous , is here newly reported for North American. The species is immediately recognizable by the black ascomata bristling with short brown setae and by growing among the soredia of various host lichens. The ascospores are 1-septate, colorless to pale brown, and 11-14 x 3.5-5 µm in size (Etayo 1996). Etayo (1996) discussed the possibility that A. coronata may not be correctly placed in Arthonia which seems probable since the setose ascomata are unique. It is tempting to speculate that the setae serve to anchor the ascomata to the highly unstable soredia. Arthonia coronata was described from the soralia of Flavoparmelia caperata (L.) Hale and the two collections from Maine are also on this host. Etayo (1996) reported it from France, Spain and the Canary Islands (Gomera). It has since been found in Scotland on Cladonia ochrochlora Flörke and C. subulata (L.) F. H. Wigg. (Coppins & Aptroot 2009). The specimens from Quebec that we have examined were found on C. digitata (L.) Hoffm. and the soralia of Parmeliopsis hyperopta (Ach.) Arnold. The only unifying theme as regards host taxa seems to be the production of soredia. As is the case for A. colombiana reported above, based on the available data A. coronata appears to have an oceanic distribution in the Northern Hemisphere.

Specimens examined. – CANADA. QUEBEC. M.R.C. DE LA HAUTE-GASPÉSIE: Parc national de la Gaspésie, Sentier du Mont-Olivine, from parking area to slopes of Mont Olivine, coniferous forest, 2.vii.2012, on Cladonia digitata, R.C. Harris 57534 (NY), on soralia of Parmeliopsis hyperopta,

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Figure 3, lectotype material of Biatora leucampyx (Bumstead s.n., FH-TUCK!). A-B, morphology of the thallus. C-E, detail of the morphology of the apothecia, including one apothecium (E) with a pruinose margin. F, section of an apothecium mounted in water. Scales: 2.0 mm: A; 1.0 mm: B; 0.25 mm: C-E; 200 μm: F.

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R.C. Harris 57546 (NY). U.S.A. MAINE. HANCOCK CO.: Township of T9 SD, Donnell Pond Maine Public Reserved Lands, Black Mountain Trail from Schoodic Beach parking area, wet conifer forest, 25.vi.2012, on soralia of Flavoparmelia caperata, R.C. Harris 57444 (NY). WALDO CO.: Town of Lincolnville, Ducktrap River Preserve, ca. 4 mi NE of Lincolnville Center on ME 52, hardwood floodplain forest, 28.v.2009, on soralia of F. caperata, R.C. Harris 55505 (NY).

3. Bacidia schweinitzii (Fr. ex. Tuck.) A. Schneid., Guide Study Lich., p. 110. 1898. Biatora schweinitzii Fr. ex Tuck. in Darlington, Fl. Cestrica, ed. 3, p. 447. 1853. TYPE: U.S.A. PENNSYLVANIA. CHESTER CO.: sine loc., on trunks, sine date, E. Michener s.n. [Michener’s Lichens, book 5, pg. 7, no. 126] (BPI[n.v.], lectotype [selected by Ekman (1996)]). Biatora luteola var. schweinitzii (Fr. ex Tuck.) Tuck., Gen Lich., p. 166. 1872.

Syn. nov. Biatora leucampyx Tuck., Syn. N. Amer. Lich., 2: 47. 1888. TYPE: U.S.A. MASSACHUSETTS. [WITHOUT COUNTY]: “Berkshire”, 1845, W. Bumstead s.n. (FH-TUCK 2872 [HUH-BARCODE 00197667]!, lectotype selected here). Bacidia leucampyx (Tuck.) Dodge, Rhodora, 28: 160. 1926.

DISCUSSION. – While organizing our reprint collection we came across a publication by C.W. Dodge documenting the lichens of the Gaspe Peninsula in Quebec, Canada (Dodge 1926). In reading this publication we discovered that Dodge had proposed the new combination Bacidia leucampyx for the name Biatora leucampyx. Since we were unfamiliar with both of these names we searched for them in Index Fungorum and on the checklist of North American lichens (Esslinger 2011), both without success. Using Dodge’s citation however, we were able to locate the protologue and determine that based on the description the taxon was likely conspecific with the common eastern North American species Bacidia schweinitzii (Ekman 1996). In an effort to resolve the identity of this long overlooked name we borrowed the type material from the Tuckerman herbarium at FH. Examination of the only specimen filed under this name in FH-TUCK (fig. 3) confirmed our suspicion that Bacidia leucampyx should be treated as a synonym of B. schweinitzii as is formally proposed here. It should be noted that although only a single specimen labeled Biatora leucampyx was located in the Tuckerman herbarium at FH we have chosen to select the specimen as the lectotype rather than simply assume its status as the holotype. This is due to the paucity of data given in the protologue, where Tuckerman stated only that he had seen material collected on beech (Fagus) in western Massachusetts. The precise locality, collector, and date of collection were not included in the protologue despite the presence of these data on the label associated with the lectotype. Thus we cannot conclude with certainty that Tuckerman did not have additional material available to him at the time of the description of B. leucampyx that was subsequently distributed to other herbaria or lost.

4. Pertusaria plittiana Erichsen, Repert Spec. Nov. Reg. Veg., 41: 77. 1936. TYPE: MARYLAND. BALTIMORE CO.: Gunpowder River, 7.vi.1910, on rocks along right bank, C.C. Plitt 255 (HBG! lectotype [selected by Dibben (1980)]; NY!, US?[n.v.]3, isolectotypes).

Syn. nov. Pertusaria shenandoensis Hale & Dibben in Dibben, Publications in Biology and Geology, Milwaukee Public Museum, 5: 108. 1980. TYPE U.S.A. VIRGINIA. PAGE CO.: Shenandoah National Park, along trail to Little Stony Man (Mt.), 1.vi.1971, on rocks in sheltered woods, M.E. Hale s.n. (US!, holotype).

3There are two specimens of Plitt 255 at US. One of these, US Barcode 00517446[!] was annotated by Dibben in 1971 as an isoype of Pertusaria plittiana. Dibben’s TLC annotation slip reports only norstictic acid and a trace of connorstictic acid. Our examination of this specimen revealed that it represents Aspicilia cinerea (L.) Korb. and not P. plittiana (fig 4E and F). Based on the original notes associated with this specimen, which illustrate an and ascospore of typical P. plittiana, it is possible that the specimen that was originally present in the packet became disassociated after Dibben annotated the specimen. There is no photograph of the second specimen, US Barcode 00348614, available online however we suspect that it may represent a true isolectotype of P. plittiana.

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Figure 4, comparison of the lectotype of Pertusaria plittiana (A, thallus; B, fertile verruca), holotype of P. shenandoensis (C, thallus; D, sterile verrucae), and non-type specimen of Aspicilia cinerea at US annotated as an isotype of P. plittiana by Dibben (E, thallus; F, apothecia). Scale bars = 2.0 mm: C; 1.0 mm: A and E; 0.5 mm: B, D, and E.

318 DISCUSSION. – Pertusaria shenandoensis is an enigmatic taxon that has puzzled us, and evidently other authors, since the time of its description. The species was described by Dibben (1980) on the basis of several sterile specimens from the Shenandoah Mountains of Virginia in eastern North America. The protologue includes a detailed description that alludes to the presence of sterile verrucae (i.e. warts typical of Pertusaria s. str.). Considering our extensive collection efforts and studies of sterile crustose lichens in eastern North America, we have always wondered why we have not encountered P. shenandoensis. Indeed, a sterile crustose species that produces norstictic acid and gyrophoric acid would be very distinctive. Further, the existence of a lichen species endemic to a single locality in the central Appalachians seemed highly improbable. However if such a species did exist, it would certainly warrant significant conservation attention. Recently we became aware of the online key to Pertusaria (Archer & Elix undated [accessed 2011]) and noticed that P. shenandoensis was included among the species of “uncertain status”. Review of the protologue led us to wonder if, in fact, Dibben had described the species we have been referring to as “Phlyctis petraea”. That species is common throughout the Appalachians, and one of us (JCL) had recently seen abundant material from the vicinity of the type locality of P. shenandoensis while visiting CANL. Hoping that we had finally found a published name for Phlyctis “petraea”, we borrowed the paratype of Pertusaria shenandoensis that had been deposited in DUKE. Study of the specimen revealed it to be a sterile thallus of the common eastern North American species P. plittiana. Thin Layer Chromatography (TLC) of the specimen revealed only the presence of the norstictic acid aggregate. We suspect that Dibben’s (1980) report of gyrophoric acid from the species might have been the result of contamination from adjacent thalli of Trapelia placodioides Coppins & P. James, which could potentially resemble P. plittiana when young. Examination of the holotype of P. shenandoensis at US (figs. 4, C and D) confirmed that it has the same chemistry as the paratype at DUKE and also represents a sterile specimen of P. plittiana growing adjacent to young thalli of T. placodioides. When Dibben (1980) described Pertusaria shenandoensis he compared it to P. plittiana but considered the two species to differ significantly in chemistry. Dibben (1980) considered P. plittiana to produce norstictic acid, connorstictic acid, perlatolic acid, and stenosporic acid, with the first three compounds occurring in variable concentrations. Conversely, he considered P. shenandoensis to produce only norstictic acid with a trace of gyrophoric acid. The chemistry of Pertusaria plittiana reported by Dibben (1980) is contradicted however, by our examination of the lectotype at HBG (figs. 4A and B) and an isolectotype at NY. TLC of both specimens detected the presence of only norstictic and connorstictic acids. Thus it would seem that the type collections of P. plittiana and P. shenandoensis do not actually differ in chemistry. As such we place P. shenandoensis in synonymy with P. plittiana here. During our fieldwork in North America we have encountered populations of Pertusaria plittiana whose chemistry agreed with that published by Dibben (1980) as well as populations that produced only norstictic and connorstictic acids. It should be noted that populations of P. plittiana with the latter chemistry have also been reported from China (Zhao et al. 2004). Here, we assert that the populations referred to P. shenandoensis, as well as those represented by the type collection of P. plittiana, likely represent a chemical variant of P. plittiana in which stenosporic and perlatolic acids are absent or not detectable with TLC. It would be logical to conclude that if two of the substances produced in P. plittiana can be present in highly variable concentrations as Dibben (1980) proposed then the remaining ones can be as well.

Additional specimen of Pertusaria shenandoensis examined. – U.S.A. VIRGINIA. PAGE CO.: Shenandoah National Park, Stoney Man Mtn., ½ mi up trail from Little Stoney Man parking lot, 10.iv.1972, on flat rock surface by trail, M.J. Dibben 22437 (DUKE!).

5. Rosellinula haplospora (Nyl.) R. Sant., Syst. Ascom. 5(2): 311. 1986. Endococcus haplosporus Th. Fr. & Almq., Bot. Notiser [unnumbered volume for 1867]: 109. 1867. TYPE: SWEDEN: Herjedalen, 1866, Funnesdalens kalkbrott, on Aspicilia cinerea, S. Almquist s.n. (UPS[n.v.], holotype) Muellerella haplospora (Th. Fr. & Almq.) Arnold, Flora, Jena 57: 155. 1874. Roselliniella haplospora (Th. Fr. & Almq.) Hafellner, Herzogia 7: 147. 1985.

319 DISCUSSION. – In conjunction with collection efforts aimed at understanding the genus Lepraria s.l. in North America JCL conducted fieldwork in the Yukon Territory of Canada. Among the interesting discoveries made during this trip was a collection of the lichenicolous fungus Rosellinula haplospora that had infected a thallus of Aspicilia cinerea (L.) Körber. This is the first report of R. haplospora from North America. It can easily be recognized by its peritheciate fruiting bodies with interascal filaments, polysporous asci and simple, brown ascospores (7-11 x 4.5-6.5 μm in size fide Hafellner (1985) and Ihlen and Wedin (2008)). Although occurring on unrelated hosts, Muellerella polyspora Müll. Arg. (on Arthonia radiata and Opegrapha atra; corticolous members of the Arthoniomycetes) and M. hospitans Stizenb. (on Bacidia spp. and Lecania fuscella; corticolous members of the Ramalinaceae) could be confused with R. haplospora. Both of these taxa lack interascal filaments and have differ in characters of the ascospores (fide Hawksworth et al. 2008); smaller (2.5-3.5(-4) x (1.5-)2-2.5 µm) and nearly subglobose in M. hospitans, smaller in M. polyspora (5-7(-9.5) x 2.5-3(-3.5) µm).

Specimen examined. – CANADA. YUKON TERRITORY: NW slope of Needle Mt., E of Annie Lake Rd. above N end of Annie Lake, 16.3 km W of jct of Annie Lake Rd. and Klondike Hwy. (Yukon 2), 4.vi.2011, on Aspicilia cinerea on rock, J.C. Lendemer 28782 (NY).

ACKNOWLEDGEMENTS

Alan Archer and Kendra Driscoll are thanked for their reviews of the manuscript. We thank the curators and collections managers of DUKE, FH, HBG, and US for the loan of specimens used in this study. Irwin Brodo is thanked for facilitating the first author’s visit to CANL, and both Irwin and Fenja Brodo are thanked for their hospitality during the same visit. This study was produced as part of NSF Award DEB-145511. The lichenicolous fungi reported from Maine were collected during educational outreach conducted by the authors at the Humboldt Field Research Institute in Steuben, and those reported from Quebec were collected as part of outreach conducted by the authors at the 2012 Tuckerman Workshop held on the Gaspe Peninsula. The first author’s fieldwork in the Yukon Territory was supported by NSF Award DEB-1110433.

LITERATURE CITED

Archer, A.W. and J.A. Elix. [undated]. A Preliminary World-wide Key to the Lichen Genus Pertusaria. http://www.rbgsyd.nsw.gov.au/science/Plant_Diversity_Research/Key_to_Pertusaria. [website, accessed 13 November 2011] Clerc, P. and P.F. May. 2007. Usnea flammea () new for North America. The Bryologist, 110(1): 126-128. Coppins, B.J. and A. Aptroot. Arthonia Ach. (1806) In: Smith, C.W., A. Aptroot, B. J. Coppins, A. Fletcher, O. L. Gilbert, P. W. James and P. A. Wolseley (eds.) 2009. The Lichens of Great Britain and Ireland. 2nd ed. British Lichen Society, London. pp. 153-171. Degelius, G. 1935. Das ozeanische Element der Strauch- und Laubflechtenflora von Skandinavien. Acta Phytogeographica Suecica, 7: 411. Dibben, M.J. 1980. The Chemosystematics of the Lichen Genus Pertusaria in North America North of Mexico. Publications in Biology and Geology No. 5, Milwaukee Public Museum Press, Milwaukee. 162 pp. Dodge, C.W. 1926. Lichens of the Gaspé peninsula, Quebec. Rhodora, 28: 157-161. Ekman, S. 1996. The corticolous and lignicolous species of Bacidia and Bacidina in North America. Opera Botanica, 127: 1-148. Esslinger, T.L. 2011. 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 (#17) 16 May 2011), Fargo, North Dakota. Etayo, J. 1996. Aportación a la flora liquénica de las Islas Canarias. I. Hongos liquenícolas de Gomera. Bulletin de la Société Linnéenne de Provence, 47: 93-110. Etayo, J. 2002. Aportación al Conocimiento de los Hongos Liquenícolas de Colombia. Bibliotheca Lichenologica 84, J. Cramer, Berlin, Stuttgart. 154 pp. Hafellner, J. 1985. Studien uber lichenicole Pilze und Flechten III. Die Gattung Roselliniella Vainio emend. Haf. (Ascomycotina, Dothideales). Herzogia, 7: 145-162. Hawksworth, D.L., V. Atienza and B.J. Coppins. 2008. DRAFT Artificial Keys to the Lichenicolous Fungi of Great Britain, Ireland, the Channel Islands, Iberian Peninsula, and Canary Islands, Third Draft Edition for Testing. Distributed by the authors. 36pp. Ihlen, P.G. and M. Wedin. 2008. An annotated key to the lichenicolous (including mitosporic morphs) of Sweden. Nova Hedwigia, 86(3-4): 275-365.

320 Jørgensen, P.M. 1996. The oceanic element in the Scandinavian lichen flora revisited. Symbolae Botanicae Upsaliensis, 31(3): 297-317. Knudsen, K., J.C. Lendemer and R.C. Harris. 2011. studies in lichens and lichenicolous fungi - no. 15: miscellaneous notes on species from eastern North America. Opuscula Philolichenum, 9: 45-75. Lendemer J.C., K. Knudsen and J.A. Elix. 2008. Lepraria friabilis, a new species from Eastern North America. Opuscula Philolichenum 5: 61-66. Zhao, Z-T., Q. Ren and A. Aptroot. 2004. An annotated key to the lichen genus Pertusaria in China. The Bryologist, 107(4): 531-541.

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