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Megalaria beechingii (lichenized Ascomycota), a new species from eastern North America

Authors:
Opuscula Philolichenum, 4: 39-44. 2007.
Megalaria beechingii (lichenized ascomycota), a New
Species from Eastern North America
JAMES C. LENDEMER1
ABSTRACT. Megalaria beechingii, a new species from the southern Appalachian Mountains, USA is
described.
INTRODUCTION
Recently, I received a number of interesting collections made by Sean Beeching from the southern
Appalachian Mountains of North America. Among the specimens was a puzzling saxicolous crustose
lichen that, upon further study, was recognized as an undescribed species of Megalaria. A trip to the
locality allowed me to gather additional ecological data as well as ample material to allow detailed study
and to serve as a type specimen. Subsequent examination of undetermined material in the herbarium of the
New York Botanical Garden (NY) resulted in the discovery of several additional collections of the species.
The material is here described as Megalaria beechingii.
MATERIALS & METHODS
Specimens were initially studied using a standard light dissecting microscope. Hand cut sections
were prepared using a razor blade and by wetting the thallus/apothecia with water. Apothecial anatomy,
pigments, and the characteristics of the asci and ascospores were examined following the methods of
Ekman (1996). Photographs were prepared using a CoolPix 950 Digital Camera with the images assembled
and edited (for grayscale and improved contrast) in Adobe Photoshop. TLC was performed using solvent C
following the methods of Culberson and Kristinsson (1970). Selected specimens used for comparison are
cited in Appendix I.
TAXONOMIC SECTION
Megalaria beechingii Lendemer sp. nov.
Ab Megalario pulvereo substrato rupicola, hypothecio brunneo (K+ violaceus), et acido
fumarprotocetrarico nullo differt.
TYPE: USA. GEORGIA. RABUN CO.: Lake Burton Wildlife Management Area, vicinity of Popcorn
Overlook, just south of US Route 76, north of Lake Burton, ~7.5 miles east of Clayton, Lake Burton Quad.,
elev. ca. 750 ft., humid overgrown dunite barren, on ridge-top, above tributary to Lake Burton, south-facing
slope, pine (Pinus) – oak (Quercus) forest above, mixed hardwoods (Acer, Nyssa, Quercus) below, on large
boulders, 17.IX.2006, J.C. Lendemer et al. 7700 (NY, holotype; CANL, PH-HBL, UCR, isotypes)
DESCRPITION. Thallus crustose, saxicolous, blue-gray to greenish-gray, areolate with areoles
resting on a well developed white prothallus; prothallus white, fibrous in the available material; areoles
1 JAMES C. LENDMER – Dept. of Botany, The Academy of Natural Sciences of Philadelphia, 1900
Benjamin Franklin Pky., Philadelphia, PA, 19103, USA. – e-mail: lendemer@acnatsci.org
39
arising from thalline warts near the growing edge of the thallus, initially + globose and convex, becoming
flattened with age and aggregating as the thallus grows until forming a + continuous areolate crust;
granules (granulose soredia) forming from the apical breakdown of areoles (plate 1, figure 3). Apothecia
biatorine, epruinose, purple-black to jet black, initially plane and eventually becoming convex; margin
concolorous with disc, persistent, becoming + excluded in convex apothecia; proper exciple laterally 80–
100µm thick, occasionally with few large crystals, the interior dark brown-black (K–, HNO3–) contrasting
distinctly with the hyaline rim and the hypothecium, the outermost cells of the rim with variable amount of
bluish-green pigment (blue-green, K+ intense blue-green, HNO3+ red, often faint in mature apothecia),
composed of loose radiating hyphae, continuous under the hypothecium; hypothecium brown-red to dark
brownish black (K+ purple-red, HNO3+ intense brown), 50µm high; hymenium 100–110µm high, hyaline
though often with streaks of pigment from hypothecium in the lower portions; epihymenium with variable
blue-black pigmentation (K+ purplish, HNO3+ red), often with bluish excipular pigment extending from
rim of proper exciple to adjacent portions; paraphyses unbranched to little branched above, + anastomosed
below; asci Bacidia-type, cylindrical to + clavate, octosporous; ascospores broadly ellipsoid, often bent
toward one side (kidney bean-shaped), hyaline, 1-septate, thick walled, not halonate, (12.7) –13.8– (14.8) x
(4.6) –5.1– (5.4)µm (n=15; xbar + 0.25s); pycnidia not seen.
CHEMISTRY. – Atranorin and zeorin by TLC. Spot tests, K+ yellow, KC+ yellowish, C-, PD-, UV-.
ECOLOGY & DISTRIBUTION. – Initially Megalaria beechingii was known only from the type locality,
a small ridge-top dunite barren in the southern Appalachian Mountains of eastern North America. The
barren is bisected by a major highway and is somewhat overgrown. It is located on a south-facing slope
above a small stream, and the topography and proximity to water have created a micro-habitat with high
humidity; as was evidenced by the luxuriant growth of both corticolous and saxicolous macrolichens. It
should be noted that, whereas many normally corticolous species were found on rocks at the locality, M.
beechingii was exclusively saxicolous. The species is also remarkably abundant at the type locality,
seemingly the dominant crustose lichen on many exposed rocks.
The dominance of M. beechingii at the type locality, coupled with the lack of previous collections
(excepting a sterile collection made by R.C. Harris in the vicinity of the type locality) at first led me to
assume the species was a narrow endemic. However, a search of undetermined material in the herbarium of
the New York Botanical Garden revealed two additional collections from Clay Co., North Carolina.
Associated saxicolous species found with M. beechingii include Acarospora dispersa H. Magn.,
Diploschistes actinostomus (Pers.) Zahlbr., D. scruposus (Schreb.) Norman, Fellhanera sp., Gomphillus
americanus Essl., Hypotrachyna spp., Lasallia papulosa (Ach.) Llano, Parmotrema reticulatum (Taylor)
M. Choisy, Pertusaria plittiana Erichsen, and Phlyctis petraea ined.
DISCUSSION. – The genus Megalaria Hafellner was established by Hafellner (1984) to
accommodate a single species, M. grossa (Pers. ex Nyl.) Hafellner, that had previously been placed in
Catillaria A. Massal. or Catinaria Vain. The type species, M. grossa, differs significantly from the other
sixteen species that have subsequently been placed in the genus by others (Ekman and Tønsberg 1996,
Fryday 2004a, 2004b, Galloway 2004, Schreiner and Hafellner 1992), perhaps most notably in having a
different ascus type and photobiont.
When Ekman and Tønsberg (1996) described the corticolous species M. brodoana from the
Pacific Northwest of North America, they took the opportunity to discuss the problems associated with the
circumscription of Megalaria, particularly the seemingly isolated position of the type species. Ekman and
Tønsberg (1996) concluded that whereas differences exist among the taxa referred to Megalaria there are
also substantial similarities. The authors thus proposed two alternative circumscriptions of Megalaria, one
restricting the genus to the type species M. grossa, and the other adopting a “broad” concept of the genus
reflecting the similarities in apothecial anatomy and tendencies in ascus type. Ekman and Tønsberg (1996)
adopted the latter circumscription of the genus, which was subsequently confirmed with molecular data
(Ekman 2001).
The new species differs most obviously from M. pulverea by its saxicolous habit and dark
pigmented hypothecium, the two species are otherwise quite similar and, M. pulverea (Borrer) Hafellner &
E. Schreiner has been reported from North America (Esslinger 2006). Interestingly, while the species are
also chemically similar in the production of atranorin and zeorin, M. beechingii seems to constantly lack
fumarprotocetraric acid, which is sometimes present in M. pulverea (fumarprotocetraric acid is present in a
collection of M. pulverea from Rabun Co., Georgia (NY!)).
40
Figures 1-3. Megalaria beechingii (holotype). Figure 1. Thallus with distinct prothallus and formation of
areoles along growing edge. Figure 2. Apothecium. Figure 3. Formation of granulose soredia from the
apical breakdown of areoles.
Figure 4. Known distribution of Megalaria beechingi, in eastern North America.
41
Within the larger, broad context of Megalaria the new species is distinguished by the unique
combination of thalline and apothecial characters, saxicolous habit, and chemistry. Megalaria beechingii is
one of only three currently recognized, species of Megalaria that are not corticolous. The three other
saxicolous and terricolous species of Megalaria being only recently recognized (Fryday 2004a, 2004b).
None of these taxa have been reported from eastern North America, and all three differ in apothecial or
thalline characters. Coppins (1992) reported several saxicolous specimens of M. grossa from Europe but
that species has ascospores and apothecia much larger than those of M. beechingii and is not known to
produce lichen substances (Ekman and Tønsberg 1996).
ADDITIONAL SPECIMENS EXAMINED . USA. GEORGIA. RABUN CO.: 12.2. mi. W of Clayton, 11.vi.1991,
R.C. Harris 13796 (NY, sterile); vicinity of Popcorn Overlook, 27.vii.2006, S.Q. Beeching s.n. (PH-HBL)
NORTH CAROLINA. CLAY CO.: Nantahala National Forest, along Buck Creek Road, 1.2 mi. NNW of
US 64, 9.x.1998, R.C. Harris 42695 (NY), R.C. Harris 42729 (NY).
ACKNOWLEDGEMENTS
First and foremost I thank Sean Beeching for sending this and other interesting species from the
southern Appalachians as well as for accompanying me for several days of field work in northern Georgia and adjacent
North Carolina. Irwin Brodo, Alan Fryday and Richard Harris are thanked for helpful discussion and criticism of the
manuscript. Also, Andy Moroz for his companionship during field work and the curators of CANL, MSC, and NY for
locating and loaning comparative material of other Megalaria species used in this study.
LITERATURE CITED
Coppins, B.J. 1992. Megalaria. In: The Lichen Flora of Great Britain and Ireland (O.W. Purvis, B.J. Coppins, D.L.
Hawksworth, P.W. James & D.M. Moore, eds.): pp. 365-366. Natural History Museum/British Lichen Society,
London.
Culberson, C.F. and H. Kristinsson. 1970. A standardized method for the identification of lichen products. Journal of
Chromatography, 46: 85-93.
Ekman, S. 2001. Molecular phylogeny of the Bacidiaceae (Lecanorales, lichenized Ascomycota). Mycological
Research, 105(7): 783-797.
Ekman, S. 1996. The corticolous and lignicolous species of Bacidia and Bacidina in North America. Opera Botanica,
127: 1-148.
Ekman, S. & T. Tønsberg. 1996. A new species of Megalaria from the North American west coast, and notes on the
generic circumscription. The Bryologist, 99(1): 34–40.
Esslinger, T.L. 2006. 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.nodak.edu/instruct/esslinge/chcklst/chck lst7.htm (First Posted 1 December 1997, Most Recent
Update 10 April 2006), Fargo, North Dakota.
Fryday, A.M. 2004a. A new species of Fuscopannaria with a green photobiont, and other taxonomic innovations and
new records of lichenized-fungi from Alaska. The Bryologist, 107(2): 173–179.
Fryday, A.M. 2004b. New species and records of lichenized fungi from Campbell Island and the Auckland Islands,
New Zealand. In: Contributions to Lichenology. Festschrift in Honour of Hannes Hertel. (P. Döbbeler, & G.
Rambold, eds.): pp. 127-146. J. Cramer in der Gebrüder Borntraeger, Berlin, Stuttgart.
Galloway, D.J. 2004. New lichen taxa and names in the New Zealand mycobiota. New Zealand Journal of Botany. 42:
105–120.
Schreiner, E. & J. Hafellner. 1992. Sorediöse, corticole Krustenflechten im Ostalpenraum. I. Die Flechtenstoffe und die
gesicherte Verbreitung der besser bekannten Arten. Bibliotheca Lichenologica, 45, J. Cramer, Berlin, Stuttgart.
291 pp.
APPENDIX I
SELECTED SPECIMENS EXAMINED OF ADDITIONAL MEGALARIA SPECIES . M. BRODOANA. CANADA. BRITISH
COLUMBIA.: Queen Charlotte Islands, Moresby Island, Tasu, 52º46’N, 132º02’W, in Picea-Thuja forest on
Gowing Island, on Almnus [sic] rubra above beach, 29.vii.1967, I.M. Brodo et al. 12844 (CANL, PARATYPE);
Queen Charlotte Islands, Moresby Island, Jedway, 52º18’N, 131º13’W, along road to foot of Harriet Harbour
and near stream inlet, Picea-Tsuga-Alnus stand at shore, on Tsuga at edge of forest, 26.vii.1967, I.M. Brodo
et al. 12552 (CANL, PARATYPE ), I.M. Brodo et al. 12569 (CANL, PARATYPE). M. COLUMBIANA. CANADA. BRITISH
COLUMBIA.: Didney, on alder trunks, 12.xii.1912, J. Macoun 59 (CANL, ISOLECTOTYPE). M. GROSSA. SPAIN.
CANARY ISLANDS.: Tenerife, Fayal-brezal with scattered pines, between Agua Garcia and Monte de La
42
Esperanza, 1100 m., 26.ii.1964, central mountain range, H.A. Imshaug et al. 35856 (MSC); Fayal-brezal zone,
ridge from Piedra Chinobre to Pico de Limante, 800 m., 21.ii.1964, Sierra de Anaga, H.A. Imshaug et al.
35673A (MSC). M. IMSHAUGII. NEW ZEALAND.: Campbell Island, Rock outcrops and tussock grass-land on
summit ridge of Beeman Hill, 600-650 ft., 24.xii.1969, R.C. Harris 4535 (MSC, PARATYPE), R.C. Harris 4537
(MSC, PARATYPE). M. JEMTLANDICA. USA. ALASKA. BARROW.: Dry bank north of North Meadow Lake,
70º18’40”N, 156º39’15”W, 0 m., 18.vii.2001, A. Fryday 8370 (MSC). M. LAURERI. CANADA. ONTARIO.:
Nipissing District, Lookout Trail, Algonquin Provincial Park, ix.1960, on Acer, H.A. Imshaug 26876 (MSC).
USA. MICHIGAN. CHEBOYGAN CO.: Moist hardwoods in gorge of Carp Creek, University of Michigan Biological
Station, 10.vii.1967, on Fagus, R.C. Harris 3655 (MSC). M. MELANOTROPA. NEW ZEALAND .: Campbell Island,
Dracophyllum scrub on north side of Preserverance Harbour, about half-mile north of Beeman Station,
22.i.1970, R.C. Harris 5694 (MSC). M. PULVEREA. BRITISH ISLES.: England, V.C. 3 South Devon, Dartmoor,
Okehampton, Throwleigh, Blackaton Brook, Grid Ref. 20/65-91, on bole of Quercus in sheltered, rather
moist woodland, 14.xii.1975, P.W. James s.n. (CANL, FERTILE).
43
44
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This is a taxonomic revision of the corticolous and lignicolous taxa of the genera Bacldia and Bacidina (Lecanoraceae, Lecanorales, lichenized Ascomycotina) in the continental United States and Canada. Twenty-seven species of Bacidia (one of which is divided into two subspecies) and twelve species of Bacidina are recognized. The morphology, secondary chemistry, ecology, and phytogeography of these spceies are discussed, and a key is provided. Apothecium pigmentation, thallus composition, presence of crystals in the proper exciple, hymenium height, thickness of paraphyses, spore shape and size, in Bacidina sometimes also conidial type and tholus structure, have proved to be the most important characters to distinguish betwcen species. Circumscriptions of Bacidia and Bacidina are attempted. Tholus structure, cell structure of the proper exciple, presence of goniocysts, presence of crystals in the thallus cortex, and size of conidiogenous cells are the most valuable characters in the delimitation of these genera. Nine new taxa are described: Bacidia diffracta, B. helicospora, B. salmonea, Bacidina aenea, Bn. californica, Bn. crystallifera, Bn. ramea, Bn, squamellosa, and Bn. varia. Twelve new combinations are introduced: Bacidia campalea, Bacidia laurocerasi subsp. idahoensis, Bacidina assulata, Bn. egenuloidea, Byssoloma meadii, Fellhanera floridana, Herteliana alaskensis, Lecania stigmatella, L, subfuscula, Pachyphiale gyalizella, Ropalospora phaeoplaca, and Ophioparma rubricosa. In addition to the newIy described species, 12 species of Bacidia and four species of Bacidina are correctly reported from the study area for the first time. The genera Psorella and Toniniopsis are reduced into synonymy with Bacidia.
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