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In Eastern North America

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In Eastern North America Opuscula Philolichenum, 6: 1-36. 2009. A synopsis of the lichen genus Heterodermia (Physciaceae, lichenized Ascomycota) in eastern North America 1 JAMES C. LENDEMER ABSTRACT. – A preliminary treatment of the members of the genus Heterodermia occurring in eastern North America is provided. A key to species is included with color images of many species that have not been adequately illustrated previously. A new species, Heterodermia erecta, is described from high elevations of the southern Appalachians. Heterodermia microphylla is provisionally excluded from the study area as all specimens examined were misidentifications of other taxa. A norstictic acid deficient strain of H. neglecta is recognized pending further study of the material which may warrant recognition as a distinct species consisting of disjunct populations in the Canadian Maritimes, southern Appalachian Mountains, and central Florida. INTRODUCTION The roots of this treatment are grounded in nearly five years of field work in the Appalachian Mountains of eastern North America, combined with collecting expeditions throughout the southeastern coastal plain and the Ozarks. Over this time I have collected hundreds specimens of Heterodermia Trevis. and visited dozens of localities, with some localities hosting more than half a dozen species. While most specimens were easily identified, problems quickly began to accumulate and consultation of the literature often only resulted in further confusion. Conversations with other lichenologists working in the region made it clear that they had encountered similar problems. This led me to compile a key to the genus Heterodermia in eastern North America to aid in the routine identification of specimens. The revision of the material held by The New York Botanical Garden (NY) led to the discovery of significant range extensions (and reductions) as well as the recognition that the taxonomic limits and concepts of some species remained obscure despite the large amount of recent literature on the subject (Dey 1978; Harris 1995; Kurokawa 1962, 1973; Moberg & Nash 2002). Thus, I decided to attempt a preliminary treatment of Heterodermia species occurring in eastern North America the results of which are provided here. The present work is not intended to be the final word on the subject, but rather a guide that will both facilitate identification of specimens and highlight areas where taxonomic problems remain. Despite being a genus of charismatic and conspicuous macrolichens, the status of several North American Heterodermia species remains unclear. Hopefully this treatment represents a first step toward filling in the gaps in our knowledge. MATERIALS AND METHODS This study is limited primarily to herbarium materials at The New York Botanical Garden (NY). Normally one would prefer to review as much material as possible for such a study, however due to the time and resources required to properly identify specimens it was simply impractical to revise the material held in other large herbaria. Heterodermia is an attractive and conspicuous genus, and as such it is frequently collected even by non–lichenologists. An attempt has been made to treat all taxa currently reported from eastern North America. Synonymies are not provided and type specimens are only cited when they are particularly relevant to a problem. Otherwise synonymies and type specimens can be found 1 JAMES C. LENDEMER – Cryptogamic Herbarium, Institute of Systematic Botany, The New York Botanical Garden, Bronx, NY, 10458-5126, U.S.A. – e–mail: [email protected] 1 11 in any number of publications on Heterodermia (particularly Kurokawa 1962, and 1998). Similarly, more extensive descriptions can be found in Kurokawa (1962) and Moberg and Nash (2002). All specimens were studied dry using a Bausch & Lomb StereoZoom 7 dissecting microscope and images were captured using an Olympus DP20 digital camera with Microsuite Special Edition. Illustrations were prepared using Adobe Photoshop. Following Kurokawa (1973) I have made an attempt to analyze the terpenoid profiles of each taxon in addition to the other secondary metabolites such as norstictic acid and salazinic acid which are routinely used for identification. A summary of these is provided below (Plate 1). While authors subsequent to Kurokawa (1973) do not appear to have emphasized the presence of terpenoids other than zeorin in most species, these do appear to be species specific in some cases and, as such, could prove particularly useful in future studies. Chromatography was performed using solvents A and C following the standardized methods of Culberson and Kristinsson (1970). Plate 1. Representative chromatograms of Heterodermia species recognized in this treatment run in Solvent C, based on North American material. TAXONOMIC SECTION 2 22 I. – KEY TO HETERODERMIA IN EASTERN NORTH AMERICA 1. Thallus lobes adnate, attached to the substrate, obviously foliose……………………………………...…2 2. Soredia/isidia/phyllidia absent; apothecia usually present; underside ecorticate, pale….H. hypoleuca 2. Soredia/isidia/phyllidia present; apothecia usually absent; underside various………………………..3 3. Thallus sorediate; soralia marginal or terminal but not laminal……………………………….…4 4. Underside ecorticate…………………………………………………………………………5 5. Underside evenly orange pigmented (pigment K+ purple)….………….....H. obscurata 5. Underside not pigmented, or pigmented in a manner different than above (e.g. yellow pigmented (pigment K–), purple pigmented, or pigmented only in small patches at the lobe tips)….……………………………………………………………………………6 6. Underside with small patches of orange pigment (pigment K+ purple) only at the lobe tips………………………………………………………………………………...7 7. Medulla K+ red, norstictic acid present, terpenoid 1 absent…...….H. neglecta 7. Medulla K–, norstictic acid absent; terpenoid 1 present…….…H. neglecta s.l. 6. Underside not pigmented, or pigmented yellow/orange and the pigment K–…..…8 8. Underside not pigmented yellow…………………………………………...…9 9. Lobes haphazardly erect, especially in central portions of the thallus; soralia subterminal, not reflexed, not wider than the width of the lobes; montane Appalachians……………………………………….…H. erecta 9. Lobes lying flat and plane; soralia reflexed or not; not montane Appalachians…………………………………………………………...10 10. Rhizines thyrsiform2 and pale; soralia labriform and strongly reflexed, wider than the width of the lobes; terpenoid 1 absent….... ……...………………………………………….…H. galactophylla 10. Rhizines squarrose and dark; soralia marginal, not as above, not wider than the width of the lobes; terpenoid 1 present…………… …………………………………………………...….…H. japonica 8. Underside pigmented yellow (pigment K–)…………………………………11 11. Lobes long and narrow, strap–like (see plate 3); medulla K– (norstictic acid absent); rare…………………………………...…H. appalachensis 11. Lobes not as above (see plate 4); medulla K+ red (norstictic acid present) or K–…………………………………………………………12 12. Pigment confined to the lobe tips; coastal plain.…H. japonica s. lat. 12. Pigment central and spreading towards thallus margins, but not confined to the lobe tips; Appalachian/Piedmont.…H. casarettiana 4. Underside corticate………………………………………………………………………...13 13. Medulla K– (salazinic acid absent)……………………………………...…H. speciosa 13. Medulla K+ yellow turning red…………………………………………………...…14 14. Medulla with salazinic acid; thallus large; common and usually corticolous; coastal plain/piedmont…………………………………………………H. albicans 2 thyrsiform – Throughout this treatment the term is used following Kurokawa (1962) to denote a rhizine in which the branching resembles the type of inflorescence referred to as a thryse in vascular plants. See plate 7, figure 2, herein. 3 33 14. Medulla with norstictic acid; thallus small; rare and usually saxicolous; Appalachians………………………………………………………….H. pseudospeciosa 3. Thallus with laminal isidia/isidioid–soredia or with minute marginal phyllidia/lobules……….15 15. Thallus with minute marginal phyllidia/lobules………………………….…H. squamulosa 15. Thallus with laminal isidia/isidioid–soredia……………………………………………...16 16. Underside pigmented orange (pigment K+ purple); medulla K– (salazinic acid absent); SE……………………………………………………………………H. crocea 16. Underside not pigmented orange; medulla K+ red (salazinic acid present); mostly northern/central………………………………………………………….H. granulifera 1. Thallus lobes not adnate, often ascending; appearing fruticose……………………………………….…17 17. Soredia absent; apothecia usually present………………………………………………..H. echinata 17. Soredia present; lobes strap–like………………………………………………………………...…18 18. Underside irregularly yellow pigmented; medulla without salazinic acid; rare………………… ……………………………………………………………………………...…H. appalachensis 18. Underside not pigmented yellow; medulla with or without salazinic acid; common…………19 19. Marginal cilia red pigmented (pigment an anthraquinone, K+ purple); medulla K– (salazinic acid lacking)……………………………………………...…..…H. cf. leucomela 19. Marginal cilia not red pigmented; medulla K+ red (salazinic acid or norstictic acid)……20 20. Medulla with salazinic acid; Appalachian…………………………H. leucomela s. str. 20. Medulla with norstictic acid; Florida…………………………………H. leucomela s.l. II. – THE SPECIES 1. Heterodermia albicans (Pers.) Swinscow & Krog PLATE 2 (PAGE 7). DESCRIPTION. – Thallus foliose, sorediate, upper surface gray to blue–gray; lobes long, plane, epruinose, marginally spreading and flat, centrally overlapping; medulla white; underside corticate, pale to light brown at centrally; rhizines pale, simple, primarily marginal; soralia marginal and becoming laminal in central
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