1Department of Biology, University of Central Missouri, Warrensburg, MO 64093,
and Botanical Research Institute of Texas, Fort Worth, TX 76102. 2Department of
Botany, The Field Museum, Chicago, IL 60605. 3University of Maine at Fort Kent,
Fort Kent, ME 04743. *Corresponding author – haroldkeller@hotmail.com.
Great Smoky Mountains National Park’s First Lichen
Bio-Quest
Harold W. Keller1,*, Joseph S. Ely1, H. Thorsten Lumbsch2,
and Steven B. Selva3
Abstract - The first Lichen Bio-Quest was held at the Great Smoky Mountains
Institute at Tremont near Townsend, TN, on 19–20 June, 2004. More than 30 participants
included high school teachers and students, Park volunteers and staff, area
residents, and professional lichenologists. The primary goal was first to provide an
educational component, including lichen morphology, growth forms, terminology,
and identification using lecture and video-microscopy presentations, followed by
a field component collecting lichens in different habitats. H. Thorsten Lumbsch,
an expert on crustose lichens, and Steven B. Selva, an expert on calicioid (stubble)
lichens, served as instructors, foray captains, and helped identify specimens. Lowerelevation
collection sites were located in the Tremont area (Lumber Ridge Trail and
Spruce Flats Falls Trail) and ranged from 405–550 m. High-elevation sites (Indian
Gap, Spruce-Fir Nature Trail, and the Balsam Mountain Road area) ranged from
1094 to 1706 m. Eighty-eight lichen and lichenicolous fungi species were identified,
including 10 new published Park and Tennessee records. The new lichen records
were: Aspicilia caesiocinerea, Calicium glaucellum, Chaenotheca brunneola, Placynthiella
icmalea, Trapelia glebulosa, T. placodioides, and Trapeliopsis fl exuosa.
The new lichenicolous fungi records were: Mycocalicium subtile, Phaeocalicium
polyporaeum, and Sphinctrina turbinata.
Introduction
One of the world's most biologically diverse temperate regions, Great
Smoky Mountains National Park (GSMNP), comprises approximately
210,566 ha, with roughly 40,000 ha of old-growth forest (Snell et al. 2003).
This area is known for high tree-species diversity, including tertiary relics
that survived in eastern North America and eastern Asia, but became extinct
in Europe. This type of distribution has been known to occur in higher plants,
but has only recently been discovered for lichens (Yoshimura 1968). Since
Yoshimura’s pioneering work, this distribution type has been discussed
repeatedly in the macrolichen literature (e.g., Culberson 1972, Kurokawa
1972, Yoshimura 1987). Miyawaki (1994) also listed numerous crustose
lichens with the same type of distribution. The importance of GSMNP as a
glacial refugium for widely disjunct lichen species is demonstrated by those
89
The Great Smoky Mountains National Park All Taxa Biodiversity Inventory:
A Search for Species in Our Own Backyard
2007 Southeastern Naturalist Special Issue 1:89–98
90 Southeastern Naturalist Special Issue 1
species that occur in eastern North America as well as in the Pontis region
(Turkey), and in eastern Asia.
There are few studies devoted solely to the lichen biota of GSMNP. Degelius
(1941), in his treatment of the lichen fl ora of North America, provided
data collected from GSMNP. He listed 206 species for the Park, including 15
species new to science. Dey (1978) identified 178 species of macrolichens in
the high-mountain areas of the southern Appalachians, which included areas
of the GSMNP. He recorded 4% of the species as endemic and also noted
continental disjuncts known only from Japan. Skorepa (1972) listed lichens
from the Park in his checklist, and DePriest (1984) included records in a bibliography
of southern Appalachian lichens. Ciegler et al. (2003) collected
lichens from the tree canopy in GSMNP and noted geographical disjuncts
generally found in more-northern regions.
Esslinger (2006) has created a collective checklist of lichenicolous,
lichen-forming, and allied fungi for North America. The most recent version
(10 April 2006) is available at the North Dakota State University
web site (http://www.ndsu.nodak.edu/instruct/esslinge/chcklst/chcklst7.
htm) listing 721 genera and 7734 species. The number of lichen species
published from GSMNP, and/or reported as part of the GSMNP checklist
updated periodically by the National Park Service, is approximately 563,
with an estimated total number between 600–700 species (Becky Nichols,
GSMNP, Gatlinburg, TN, pers. comm.). Many of the recent additions to the
lichen checklist and species new to science are the result of specialists in
the crustose lichens exploring the GSMNP and completing regional monographs
or papers (Tønsberg 2002, 2004; Printzen and Tønsberg 2003, 2004).
“Bio-Quest” and/or “Bio-Blitz” are terms used for special field surveys
and biotic inventories that occur within limited time duration, usually 24 to
48 hours. These events are designed to increase the public’s awareness of
biodiversity in a given area, and also to survey for new taxonomic records
and new species. If the area is small, containing several hundred hectares,
every taxonomic group is collected and identified. Larger geographic areas,
such as GSMNP, require a number of expert taxonomists to survey and
inventory as many habitats as possible, concentrating on the collection and
identification of a single target group of organisms such as lichens. The All
Taxa Biodiversity Inventory (ATBI), with staff and financial support from
Discover Life in America, Inc., is an initiative to identify all of the life
forms in GSMNP (Sharkey 2001). One approach for obtaining this data is
to support taxonomists to survey, inventory, and identify different groups
of organisms in diverse fields such as bryology, entomology, lichenology,
mycology, myxomycology, phycology, pteridology, and tardigradology. Another
approach is to sponsor Bio-Quests that may produce new records and
new species.
The purpose of this paper is to report the results of the first Lichen
Bio-Quest for GSMNP held at the Great Smoky Mountains Institute at
2007 H.W. Keller, J.S. Ely, H.T. Lumbsch, and S.B. Selva 91
Tremont (GSMIT) on 19–20 June 2004. Our primary goal was to bring together
school teachers, students, Park volunteers and staff, area residents,
parataxonomists, and professional lichenologists to teach and learn how
to collect and identify lichens in GSMNP. The objectives of the Lichen
Bio-Quest included an educational component that answered the following
questions: What is a lichen? Where do lichens grow? How do you collect
and preserve lichen specimens? How do you recognize growth forms?
How do you use lichen terminology? How do you identify lichens using
picture keys? A secondary objective was to inventory and contribute to an
annotated checklist of lichens associated with specific high-elevation and
low-elevation sites in GSMNP as part of the ATBI. The final objective was
to curate, identify, and characterize new lichen records for GSMNP, and
enter geo-referenced collection data into the ATBI database.
Methods
The Lichen Bio-Quest was held at the Great Smoky Mountains Institute
at Tremont (GSMIT), which is within GSMNP, from 8:00 am to 6:00 pm on
19–20 June 2004. More than 30 individuals registered and participated. Two
lichenologists, H. Thorsten Lumbsch, an expert on crustose lichens, and Steven
B. Selva, an expert on calicioid (stubble) lichens, served as instructors
and foray captains, and also helped identify specimens. Saturday morning
and early afternoon were devoted to lectures and field training of the participants.
Keith Langdon, Inventory and Monitoring Coordinator, GSMNP,
gave a brief introduction to collecting in the Park, with special remarks
about habitat and distribution of the federally endangered lichen species
Cetradonia linearis (Evans) J.-C. Wei & Ahti. H. Thorsten Lumbsch, Department
of Botany, The Field Museum, presented a lecture that covered lichen
symbiosis, morphology (growth forms and terminology), reproduction,
physiology, ecology, importance, systematics, and taxonomic characters.
Steven B. Selva, University of Maine at Fort Kent, presented a lecture on the
use of calicioid (stubble) lichens as environmental indicators of old-growth
forests and described morphological characters that distinguish this group
of lichens. Participants viewed lichen specimens using microscope video
cameras to illustrate morphology, terminology, and taxonomic characters.
Examples were shown representing the crustose, foliose, and fruticose lichen
growth forms. Time constraints limited late-afternoon foray activities
to the GSMIT grounds and nearby areas.
The next day was spent as an all-day foray; one team went to highelevation
sites in the vicinity of the Spruce-Fir Nature Trail, and another
team collected along the Heintooga Ridge Road and Balsam Mountain Road.
There were only 2 to 3 hours to picture-key and identify specimens after
our return to GSMIT. Many reference books on lichens were available for
perusal and were used for practice keying, but the most user-friendly book
for the beginner, with pictures and keys, was Brodo et al. (2001).
92 Southeastern Naturalist Special Issue 1
Field collection sites
Low-elevation sites at about 405 to 550 m were surveyed by the Bio-
Quest teams near GSMIT, Blount County, TN along the Lumber Ridge Trail
and along the Spruce Flats Falls Trail. The Lumber Ridge Trail is dominated
by Tsuga canadensis (L.) Carrière (eastern hemlock) and is located on a
moist northwest-facing slope. The Spruce Flats Falls Trail is in a mixed
community of T. canadensis, Liriodendron tulipifera L. (yellow poplar),
Carya ovata (Mill.) K. Koch (shagbark hickory), and Aesculus fl ava Ait.
(yellow buckeye), and is very similar to a cove hardwood forest with a
southwest-facing drier slope. The Spruce Flats Falls trailhead and the watertower
glade area at GSMIT (site of three new lichen records for the Park)
were dominated by Pinus virginiana Mill. (Virginia pine), Rhus copallina
L. (winged sumac), Danthonia spp. (oatgrasses), and Schizachyrium scoparium
(Michx.) Nash (little bluestem).
A foray team of three people concentrated on collecting calicioid
lichens from high-elevation sites at Indian Gap (1524 m) and along the
Spruce-Fir Nature Trail (1620 m), which are areas dominated by Abies
fraseri (Pursh) Poir. (Fraser fir) and Picea rubens Sarg. (red spruce).
The other teams followed Heintooga Ridge Road, which leads to the
Balsam Mountain Campground area (1706 m), and then followed Balsam
Mountain Road down to Lower Beech Gap Trail (1094 m). This portion
of the Park is dominated by Abies fraseri and Picea rubens at the highest
elevations and mixed deciduous trees such as Acer rubrum L. (red
maple), Acer saccharum Marshall (sugar maple), Aesculus flava, Betula
alleghaniensis Britton (yellow birch), and Prunus serotina Ehrh. (black
cherry) downslope. Boulders and rock outcrops along the road also provided
habitats for a variety of lichens.
Lichen taxa listed in the Results section are arranged alphabetically by
genus and species, as recognized by Esslinger (2006). An asterisk (*) indicates
a new lichen species for the Park’s checklist and/or a new published
record for the state of Tennessee. Collection details are given for this noteworthy
species group. Representative species are grouped under the field
sites where they were collected, without repeating duplicate or noteworthy
species. Some species were sight-identified by expert lichenologists, and
voucher specimens do not exist. The Universal Transverse Mercator (UTM)
coordinate system was used to record the global positioning system (GPS)
units. Voucher specimens for the calicioid lichens collected by Steven B.
Selva were deposited in the herbarium at the University of Maine at Fort
Kent (UMFK) and GSMNP; specimens collected by H. Thorsten Lumbsch
were deposited at the Field Museum (F).
Results
Noteworthy species collected during the Lichen Bio-Quest were identified,
and voucher specimens and new Park records were processed. There
2007 H.W. Keller, J.S. Ely, H.T. Lumbsch, and S.B. Selva 93
were 136 lichen and lichenicolous fungi taxa collected, representing 88
species and 10 new published Park records. There were 57 genera, 29 families,
5 suborders, and 7 orders inventoried for GSMNP. The majority of
species recorded belong to the Lecanorales (n = 76, 59.5%) with 46 genera
and 19 families. Parmeliaceae (n = 26), Cladoniacaeae (n = 9), Lecanoraceae
(n = 8), Physciaceae (n = 6), and Lobariaceae (n = 6) contained
the majority of other taxa recorded. These taxonomic groups represented
nearly 63.6% of the species encountered. The majority of these species are
foliose and fruticose lichens, which are larger, more conspicuous, and more
likely collected by first-time collectors.
Two stubble lichens that were new to the Park belong to the Caliciales;
Calicium glaucellum (Caliciaceae) and Chaenotheca brunneola (Coniocybaceae).
Phaeocalicium polyporaeum and Mycocalicium subtile were also
new records and represent two non-lichenized fungi that morphologically
resemble stubble lichens, but belong to an unrelated group of saprophytic
fungi (Mycocaliciaceae, Mycocaliciales). All of these lichens are often
overlooked because of their small size (less than 1–2 mm) and stalked habit
more similar to myxomycetes and other fungi. This group is understudied,
even by lichenologists, as evidenced by a recent (2005) two-week summer
collecting trip that resulted in 23 new records for calicioid lichens and fungi
for the Park (S.B. Selva, unpubl. data).
Noteworthy species
*Aspicilia caesiocinerea (Nyl. ex Malbr.) Arnold. Siliceous, on exposed
schistose rocks. All saxicolous lichen specimens were growing on siliceous
rocks in an open and sunny ravine with sparse vegetation of pines and shrubs
(UTM 17S 0257111E 3946498N) at the end of Spruce Flats Falls Trail near
GSMIT, 19 June 2004, Lumbsch 19277e. This is a common species on siliceous
rocks in eastern North America. It probably has been overlooked since
natural areas with exposed rocks are frequent in GSMNP.
*Calicium glaucellum Ach. Lignicolous on standing conifer snags (UTM
17S 0278493E 3943338N) near Indian Gap parking lot, 20 June 2004, Selva
9334A; lignicolous on standing Abies fraseri snags (UTM 17S 0277192E
3941683N), Spruce-Fir Nature Trail, Selva 9336. This is a fairly common
species in northeastern North America, which suggests that some calicioid
lichens have been overlooked in GSMNP.
*Chaenotheca brunneola (Ach.) Müll. Arg. Lignicolous on standing
Abies fraseri snags (UTM 17S 0278294E 3943313N) near Indian Gap
parking lot, 20 June 2004, Selva 9333, 9334B. This is perhaps the second
most common calicioid species known (S.B. Selva, pers. observ.). Brodo et
al. (2001) describes a northwestern, northeastern, and Florida distribution
for this species.
*Mycocalicium subtile (Pers.) Szatala. Lignicolous on a fallen, decaying
Quercus alba L. (white oak) log covering an extensive area of several
94 Southeastern Naturalist Special Issue 1
m2 (UTM 17S 0256460E 3947502N), Lumber Ridge Trail near GSMIT, 19
June 2004, HWK 4784, Selva 9331B. This lichenicolous fungus is probably
the most common calicioid species known. The substrate is most often
wood rather than bark, which was the case in GSMNP. Species typically
found on lignum, such as Mycocalicium subtile, also are more frequently
found on bark in aging forests (S.B. Selva, pers. observ.). The black, stalked
habit with a rounded capitulum of the apothecium appears similar in gross
morphology to stalked myxomycete species in the genus Licea (H.W. Keller,
pers. observ.).
*Phaeocalicium polyporaeum (Nyl.) Tibell. On a polypore bracket
fungus, Trichaptum pergamenum (Fr.) G. Cunn., growing on standing dead
Liquidambar styracifl ua L. (sweet gum), Selva 9331A, HWK 4783 and
Betula lenta L. (black birch), Selva 9332A (UTM 17S 025640E 3947500N),
Lumber Ridge Trail behind dorm at GSMIT, 19 June 2004.
*Placynthiella icmalea (Ach.) Coppins & P. James. On wood of an old
picnic table bench on grounds of GSMIT next to Spruce Flats Falls Trailhead
(UTM 17S 0256507E 3947014N), 19 June 2004, Lumbsch 19274.
This inconspicuous crustose lichen is among the most common lichens in
the temperate northern hemisphere and is often a pioneer on wood and soil.
This wooden picnic table was totally encrusted with lichens. Two new Park
lichen records were found here while we ate lunch. This is a common and
widespread species and should be found more frequently in the Park.
*Sphinctrina turbinata (Pers.: Fr.) De Not. Growing over the lichen
Pertusaria macounii (Lamb) Dibben on Fagus grandifolia Ehrh. (American
beech) (UTM 17S 0278630E 3943389N) along Indian Gap Trail near parking
lot, 20 June 2004, Selva 9335A.
*Trapelia glebulosa (Sm.) J.R. Laundon. On siliceous rocks near ground
behind water tower in glade near GSMIT (UTM 17S 0256589E 3947066N),
19 June 2004, Lumbsch 19278e. This is another pioneer lichen species expected
in the Park since it is a common and cosmopolitan species.
*Trapelia placodioides Coppins & P. James. On exposed siliceous rocks,
behind water tower in glade near GSMIT (UTM 17S 0256589E 3947066N),
19 June 2004, Lumbsch 19270a. This sorediate species was described in
1984 and was formerly included in T. glebulosa; however, it differs in having
a continuous, sorediate thallus. The distribution of this species in North
America is not well known, but it is very common in Europe.
*Trapeliopsis fl exuosa (Fr.) Coppins & P. James. On wood of an old picnic
table bench on grounds of GSMIT next to Spruce Flats Falls Trailhead
(UTM 17S 0256507E 3947014N), Lumbsch 19274a. This is another example
of a common species overlooked in the Park, often a pioneer colonizer
on wood.
Representative species from field sites
Lumber Ridge Trail near GSMIT: Graphis scripta (L.) Ach., Hypotrachyna
imbricatula (Zahlbr.) Hale, Lepraria lobificans Nyl., Porpidia
2007 H.W. Keller, J.S. Ely, H.T. Lumbsch, and S.B. Selva 95
albocaerulescens (Wulfen) Hertel & Knoph, and Psilolechia lucida
(Ach.) M. Choisy.
Spruce Flats Falls Trail and glade near GSMIT: Anaptychia palmulata
(Michaux) Vainio, Bacidia schweinitzii (Fr. ex E. Michener) A. Schneider,
Baeomyces rufus (Hudson) Rebent., Buellia stillingiana J. Steiner, Cetrelia
olivetorum (Nyl.) Culb. & C. Culb., Chrysothrix chlorina (Ach.) J.R. Laundon,
Cladonia chlorophaea (Flörke ex Sommerf.) Sprengel, C. coniocraea
(Flörke) Sprengel, C. fimbriata (L.) Fr., C. macilenta Hoffm., C. squamosa
Hoffm., Dibaeis baeomyces (L.f.) Rambold & Hertel, Flavoparmelia baltimorensis
(Gyelnik & Fóriss) Hale, F. caperata (L.) Hale, Heterodermia
casarettiana (A. Massal.) Trevisan, Hypocenomyce scalaris (Ach.) M.
Choisy, Hypogymnia physodes (L.) Nyl., Hypotrachyna livida (Taylor)
Hale, Imshaugia aleurites (Ach.) S.F. Meyer, Ionaspis lacustris (With.) Lutzoni,
Lecanora allophana Nyl., L. hybocarpa (Tuck.) Brodo, L. imshaugii
Brodo, L. polytropa (Hoffm.) Rabenh., L. thysanophora Harris, Lepraria
lobificans Nyl., L. neglecta (Nyl.) Erichsen, L. membranacea (Dickson)
Vainio, Parmelia squarrosa Hale, Parmotrema perforatum (Jacq.) A. Massal.,
P. reticulatum (Taylor) M. Choisy, Pertusaria amara (Ach.) Nyl., P.
velata (Turner) Nyl., Platismatia tuckermanii (Oakes) Culb. & C. Culb.,
Pseudevernia consocians (Vainio) Hale & Culb., Punctelia rudecta (Ach.)
Krog, Rhizocarpon obscuratum (Ach.) A. Massal., Sticta beauvoisii Delise,
Tuckermannopsis chlorophylla (Willd.) Hale, T. ciliaris (Ach.) Gyelnik,
Usnea pensylvanica Mot., U. strigosa (Ach.) Eaton, and Xanthoparmelia
conspersa (Ehrh. ex Ach.) Hale.
Balsam Mountain Road: Allocetraria oakesiana (Tuck.) Randlane &
Thell, Bryoria furcellata (Fr.) Brodo & D. Hawksw., Cetrelia cetrarioides
(Duby) Culb. & C. Culb., Cladonia furcata (Hudson) Schrader, C. peziziformis
(With.) J.R. Laundon, C. pleurota (Flörke) Schaerer, Coccocarpia
palmicola (Sprengel) Arv. & D.J. Galloway, Coenogonium pineti (Ach.)
Lücking & Lumbsch, Fuscopannaria leucosticta (Tuck.) P.M. Jørg., Heterodermia
leucomela (L.) Poelt., Hypotrachyna revoluta (Flörke) Hale,
Lasallia papulosa (Ach.) Llano, Lecanora caesiorubella Ach., L. subrugosa
Nyl., Leptogium cyanescens (Rabenh.) Körber, Lobaria pulmonaria
(L.) Hoffm., L. quercizans Michaux, Menegazzia terebrata (Hoffm.) A.
Massal., Nephroma parile (Ach.) Ach., Ochrolechia androgyna (Hoffm.)
Arnold, Peltigera polydactylon (Necker) Hoffm., Physcia aipolia (Ehrh.
ex Humb.) Fürnr., Porpidia crustulata (Ach.) Hertel & Knoph, P. macrocarpa
(DC.) Hertel & A.J. Schwab, Pseudocyphellaria crocata (L.) Vainio,
Pyxine sorediata (Ach.) Mont., Ramalina americana Hale, Stereocaulon
dactylophyllum Flörke, Umbilicaria mammulata (Ach.) Tuck., and Usnea
subgracilis Vainio.
Discussion
This type of field excursion has advantages and disadvantages. There
should be a limit placed at 25 persons or less and at least five foray captains
96 Southeastern Naturalist Special Issue 1
to serve as experts for different lichen taxonomic groups, or about a 5:1 instructor-
participant ratio. With inexperienced middle school, high school,
and university students, teachers, and general public groups collecting
lichens, fungi, or myxomycetes, one-on-one personal attention often is
required to help with identifications and to answer questions (H.W. Keller,
pers. observ.). We placed no restrictions on who could register so that the
general public could benefit from learning more about lichens. However,
because of the complexity of lichen terminology, chemicals used to test
color reactions, the need for both dissecting and compound microscopes,
and the number of species involved (approximately 600–700 in GSMNP),
only a cursory introduction was possible. In addition, a Bio-Quest of this
type is useful as an educational tool, but is not practical for novices to conduct
survey work. Participants learned what was and was not a lichen, and
how to recognize lichen growth forms. Participants also gained a better
appreciation of the presence of lichens and the important role they play in
different ecosystems in GSMNP. Obviously, two days is not enough time
to master lichen identification using dichotomous keys or even picture
keys. Students who want to learn more in-depth details about lichen morphology
and identification should consider attending summer field courses
on lichens at field stations such as the Humboldt Field Research Institute
in Maine (www.eaglehill.us).
A list of 88 species and 10 new published Park lichen records, including
three lichenicolous fungi, represents a successful Bio-Quest. The
new records (all common species) indicate that certain groups, such as
the stubble lichens, are understudied and are often overlooked even by
lichenologists. This is due in part to their small size and the requirement
that at least a 10x hand lens be used to scan wood or bark surfaces. Additionally,
many lichen habitats are often overlooked, such as rock surfaces
and outcrops restricted to smaller areas (saxicolous lichens), inside rocks
(endolithic), on the bark of living trees (corticolous), and especially in
the tree canopy (Ciegler et al. 2003, Keller 2004). Targeting specific
habitats that are difficult to access will increase the chances of discovering
new records and species.
An important objective of many Bio-Quests is to increase the general
public's awareness of life forms in city and state parks, nearby natural areas,
and even forested areas in their own backyard. However, all of the new
Park records were collected by professional lichenologists, suggesting that
a certain level of expertise is needed to discover new records and species.
Acknowledgments
Special thanks go to Jeanie Hilten from Discover Life in America and Michelle
Prysby from GSMIT, who, along with staff, provided the necessary pre-meeting
registration announcements and logistical support of equipment, supplies, maps, and
food arrangements for the participants. Erin Fanning, an undergraduate student at the
University of Central Missouri, assisted with the preparation of the lichen species
2007 H.W. Keller, J.S. Ely, H.T. Lumbsch, and S.B. Selva 97
list and database management. We greatly appreciate participants who volunteered to
car-pool people and gear to collection sites. The editorial commentary and detailed
suggestions of Becky Nichols, Glen H. Mittelhauser, and anonymous reviewers
greatly improved the content of this paper. This Lichen Bio-Quest was financially
supported in part by Discover Life in America Award #2004-6, National Geographic
Society Committee for Research and Exploration Award #7272-02, and National Science
Foundation DEB Award #0343447 to H.W. Keller.
Literature Cited
Brodo, I.M., S.D. Sharnoff, and S. Sharnoff. 2001. Lichens of North America. Yale
University Press, New Haven, CT. 795 pp.
Ciegler, A., U.H. Eliasson, and H.W. Keller. 2003. Tree canopy lichens of the Great
Smoky Mountains National Park. Evansia 20:114–131.
Culberson, W.L. 1972. Disjunctive distributions in the lichen-forming fungi. Annals
of the Missouri Botanical Garden 59:165–173.
Degelius, G. 1941. Contributions to the lichen fl ora of North America. II. The lichen
fl ora of the Great Smoky Mountains. Arkiv för Botanik 30A:1–80.
DePriest, P. 1984. Southern Appalachian lichens: An indexed bibliography. US Department
of the Interior, National Park Service, Atlanta, GA. Research/Resources
Management Report SER-70. 38 pp.
Dey, J.P. 1978. Fruticose and foliose lichens of the high-mountain areas of the Southern
Appalachians. Bryologist 81:1–93.
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, Fargo, ND. Available at http://www.ndsu.nodak.edu/instruct/
esslinge/chcklst/chcklst7.htm. First posted 1 December 1997; most recent update
10 April 2006.
Keller, H.W. 2004. Tree canopy biodiversity: Student research experiences in
Great Smoky Mountains National Park. Systematics and Geography of Plants
74:47–65.
Kurokawa, S. 1972. Probable mode of differentiation of lichens in Japan and eastern
North America. Pp. 139–146, In A. Graham (Ed.). Floristics and Paleofl oristics
of Asia and Eastern North America. Elsevier, Amsterdam, The Netherlands.
Miyawaki, H. 1994. Lecanora imshaugii, a lichen of eastern North America and
eastern Asia. Bryologist 97:409–411.
Printzen, C., and T. Tønsberg. 2003. Four new species and three new apothecial pigments
from the lichen genus Biatora. Bibliotheca Lichenologica 16:133–145.
Printzen, C., and T. Tønsberg. 2004. New and interesting Biatora-species, mainly
from North America. Acta Universitatis Upsaliensis, Symbolae Botanicae Upsalienses
34(1):343–357.
Sharkey, M.J. 2001. The All Taxa Biological Inventory of the Great Smoky Mountains
National Park. Florida Entomologist 84:556–564.
Skorepa, A.C. 1972. A catalog of lichens reported from Tennessee. Bryologist
75:481–500.
Snell, K.L., H.W. Keller, and U.H. Eliasson. 2003. Tree canopy myxomycetes and new
records from ground sites in the Great Smoky Mountains National Park. Castanea
68:97–108.
98 Southeastern Naturalist Special Issue 1
Tønsberg, T. 2002. Additions to the lichen fl ora of North America XI. Bryologist
105:122–125.
Tønsberg, T. 2004. Additions to the lichen fl ora of Great Smoky Mountains National
Park. ATBI Quarterly 3(1):6.
Yoshimura, I. 1968. The phytogeographical relationships between the Japanese and
North American species of Cladonia. Journal of the Hattori Botanical Laboratory
31:227–246.
Yoshimura, I. 1987. Taxonomy and speciation of Anzia and Pannoparmelia. Bibliotheca
Lichenologica 25:185–195.