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Opuscula Philolichenum, 4: 57-68 Opuscula Philolichenum, 6: 157–174. 2009. The Fellhanera silicis group in eastern North America 1 2 RICHARD C. HARRIS & JAMES C. LENDEMER ABSTRACT. – A revision of Fellhanera in eastern North America resulted in the discovery of an undescribed complex of closely related taxa differing only in their methods of reproduction and/or the size of their dispersal units. The following species of Fellhanera are described as new to science: F. eriniae, F. fallax, F. granulosa, F. hybrida, F. minnisinkorum, F. montesfumosi, and F. silicis. Additional records for F. floridana are documented. The lichenicolous fungi found on members of the F. silicis group are discussed, including the new species Phaeosporobolus fellhanerae. INTRODUCTION The lichen genus Fellhanera Vězda has long been known from North America from F. bouteillei (Desmaz.) Vězda. Although most species of Fellhanera (including F. bouteillei) are foliicolous, several corticolous and saxicolous species have recently been recognized (Aptroot et al. 1998, Ekman 1996, Sérusiaux et al. 2001, Sparrius & Aptroot 2000, van den Boom 2004). The genus is poorly known in North America, with most of the species having been recently reported (Goward et al. 1996) or described (van den Boom 2004). When the first author began studying the lichens of the Ozark Ecoregion he recognized a saxicolous Fellhanera that had been previously confused with superficially similar saxicolous species including Bacidia granosa (Tuck.) Zahlbr. (= B. coprodes (Körb.) Lettau) and Micarea erratica (Körb.) Hertel et al. A description of the taxon was included in an online treatment of undescribed Ozark lichens under the name Fellhanera silicis. Several years later, while collecting in the Delaware Water Gap National Recreation Area, we collected material of two additional Fellhanera species that were briefly discussed in print (Harris & Lendemer 2005). At that time we had intended to formally describe all three taxa in an upcoming publication, however further field work resulted in the accumulation a large amount of additional material some of which represented additional undescribed taxa. Finally, nearly a decade after the discovery of F. silicis, we have developed a preliminary treatment of Fellhanera in eastern North America that is presented here. In our region the genus is represented primarily by a complex of taxa that we hypothesize has only recently undergone speciation. With the exception of F. floridana (Tuck.) S. Ekman, which belongs to a group of orange fruited tropical species, all of the species so far discovered in eastern North America are nearly identical in characteristics of the apothecia and overall aspect of the thallus. Interestingly, the differences among these taxa center around methods of dispersal. A core group of species have identical ascospores and differ only in conidial size and the presence or absence of vegetative propagules. Two additional species differ further in having larger ascospores with more transverse septa (5–7 septa vs. 3 septa in most species). In this treatment we first provide a key to the genus Fellhanera in North America with a supplementary key to normally sterile members of the F. silicis group (section I). Secondly, we provide formal descriptions and discussions of the members of the F. silicis group and include additional records of the little–known tropical species F. floridana (section II). A brief discussion of the origin and subsequent evolution of the F. silicis group follows the descriptive section of the treatment (section III). While revising specimens of Fellhanera we encountered two lichenicolous fungi on thalli of F. granulosa; these are also documented (section IV). 1 RICHARD C. HARRIS – Institute of Systematic Botany, The New York Botanical Garden, Bronx, NY, 10458-5126, U.S.A. – e–mail: [email protected] 2 JAMES C. LENDEMER – Institute of Systematic Botany, The New York Botanical Garden, Bronx, NY, 10458-5126, U.S.A. – e–mail: [email protected] 1571 Plate 1. Conidia of Fellhanera (all scales =20 µm). Figure 1, F. silicis (Lendemer 2140, NY). Figure 2, F. hybrida (holotype). Figure 3, F. montesfumosi (holotype). Figure 4, F. fallax (holotype). Figure 5, F. minnisinkorum (holotype). Figure 6, F. erinae (holotype). MATERIALS AND METHODS This study is based primarily on material collected by the authors throughout eastern North America over the last decade. All of this material is deposited in the herbarium of The New York Botanical Garden (NY). Specimens were studied with a Zeiss Stemi 200–C dissection microscope and images of the thallus and apothecia were captured using an Olympus DP20 digital camera using Microsuite Special Edition. Microscopic characters were measured in water with an Olympus BX51 microscope and images were captured in the same manner as above. Measurements of ascospores and conidia are based on the average (= xbar) of the number of measurements made (= n) plus or minus one standard deviation (= 1SD) from the average. Measurements are given in the following format: (xbar–1SD)–xbar–(xbar+1SD); sample size (n) is also provided. The chemistry of representative specimens of all species was studied with thin layer chromatography (TLC) using solvent A and following the standardized methods of Culberson and Kristinsson (1970). Common spot test reagents were also routinely used (e.g. following Brodo et al. 2001) and throughout the manuscript the following abbreviations are used for these: K or KOH for 10% solution of potassium hydroxide, N for concentrated HNO3, and C for Chlorox bleach. 1581 1591 TAXONOMIC TREATMENT Although the bulk of the genus Fellhanera is foliicolous, the taxa described here are all either corticolous, lignicolous, or saxicolous. It is this anomalous ecology that first drew our attention to the group. This ecology coupled with similarities in the anatomy of the thallus, sexual reproductive structures and chemistry (all taxa TLC–) led us to postulate a close relationship among the species. Despite the anomalous habit of these species, all share the basic characters of the genus Fellhanera: a cellular exciple, asci of Pilocarpaceae–type, colorless septate ascospores, and bowling pin or rod shaped conidia. Fellhanera silicis is the most commonly collected and geographically widespread taxon, and as such we believe it occupies a core position in the group. This hypothesis is reflected in the detailed description of this taxon that is provided at the beginning of the taxonomic treatment. As all the other species found in the study area, excluding F. floridana, deviate from F. silicis only in several key characters, the descriptions of these taxa are abbreviated and intended primarily to illustrate differences from F. silicis. I. – KEY TO FELLHANERA IN NORTH AMERICA While species of the Fellhanera silicis group are common and widespread in eastern North America, the identification of a given specimen can be difficult. While one can often collect apotheciate specimens at a given locality, it is most common to also find (and collect) populations and individual thalli that produce only pycnidia. Pycnidia are almost always present, conspicuous, and abundant (only in F. granulosa are they inconspicuous). They are also extremely variable in size and form; initially they are subglobose, producing conidia when they are at ~50 µm in diameter, and become gaping and irregularly cupuliform when mature (~200–300 µm in diameter). We have utilized differences in these structures to circumscribe the taxa recognized in the present treatment. This means that one is forced to locate pycnidia and measure conidia to identify a given specimen. A key for sterile specimens is provided following the main key below. 1. Thallus foliicolous; apothecia pale orange–yellow; ascospores 1–septate………………..F. bouteillei 1. Thallus not foliicolous; apothecia dark brown–black; ascospores > 3–septate………………….....…2 2. Ascus Bacidia–type; conidia filiform; on calcareous substrates ……………….Bacidia coprodes 2. Ascus Pilocarpaceae–type; conidia bowling pin or short bacillar; when saxicolous, on acidic rock……………………………………………………………………………………………...3 3. Thallus with minute blastidia or goniocysts, rarely composed entirely of isidioid blastidia..4 4. Ascospores 5–9–septate; corticolous; Great Smoky Mountains….......….….. F. eriniae 4. Ascospores 3–septate; common………………………………..…………………….....5 5. Ascospores ovoid; saxicolous; Sonoran………………….....……………..F. nashii 5. Ascospores fusiform–clavate; substrate various; eastern North America……...….6 6. Conidia 4–5 µm ................................................................................................7 7. Thallus thin, composed of small areoles; mostly corticolous/lignicolous; pigment in epihymenium green or absent; northern U.S. and Ozarks ………….…………........................................................ F. minnisinkorum 7. Thallus thicker, rimose; mostly saxicolous; pigment in epihymenium brown or absent; mostly Appalachian…................................ F. granulosa 6. Conidia 5–7 µm ................................................................................ F. hybrida 3. Thallus without lichenized diaspores……………………………….…………...…………..8 8. Ascospores 5–6–septate; Great Smoky Mountains.….…….................. F. montesfumosi 8. Ascospores 3–septate …………………..........…………...………….…….…..…...…. 9 1601 9. Apothecia orange to orange–brown; corticolous; Florida ……...……. F. floridana 9. Apothecia dark, brown to blackish; mostly saxicolous; not from Florida............. 10 10. Exciple less distinct, variously pigmented;
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