Journal of the Oklahoma Native Plant Society, Volume 6, Number 1, December 2006


Oklahoma Native Plant Record
Volume 6, Number 1, December 2006

4

THE LICHENS OF NORTH CENTRAL OKLAHOMA
By DARVIN WENDELL KECK

Submitted to the Faculty of the Graduate School of
Oklahoma State University in partial fullfillment of

the requirements for the degree of
DOCTOR OF PHILOSOPHY

August, 1961

Over 1,000 specimens of lichens were collected at 78 
collecting stations in 11 counties of North Central Oklahoma 
during 1959 and 1960. The objectives were to identify lichens 
collected in the area; to establish a record of lichen 
distribution for each county in the area; and to analyze the 
ecological relationships.

One hundred eleven species and varieties were identified, 
representing 34 genera and 20 familiies. Species occurring in 
each county were Candelaria fibrosa, Parmelia bolliana, 
Physcia syncolla, Teloschistes chrysophthalomus, and Xanthoria 
candelaria. Only ten other species occurred in eight or more 
counties, while 64 species were found in three counties or 
less and 30 of these were found in only one county each. The 
number of species per county varied from a low of 19 in 
Kingfisher County to a high of 77 in Osage County. The 
variation was very closely correlated to the two physiographic 
regions of the area. The five counties largely contained in 
the Sandstone Hills varied between 41 and 77 species with an 
average of 50. The six counties in the Redbeds Plains had 
between 19 and 33 species with an average of 26.

This variation between regions is correlated to rainfall 
which averages near 40 inches per year at the eastern boundary 
of the Sandstone Hills, but only 30 inches at the western 
boundary of the Redbeds Plains. Other factors, conditioned by 
rainfall, and also having an influence on lichen growth are 
the presence of trees, the kinds of rock, and the acidity of 
the soil.

Keck, D.W.
https://doi.org/10.22488/okstate.17.100044



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INTRODUCTION
Historical Background

The term “lichen” was used
by Theophrastus (371-287
B.C.), in his History of Plants, to
signify a superficial growth
on the bark of trees. It
referred to hepatics of the
Marchantia type rather than
to the lichens as they are
currently understood. He
did, however, describe a
species of Roccella, and
another of Usnea or
Alectoria, which is perhaps
the earliest known reference
to lichens.

Lichens were alluded to by
only a few writers during
the next 2000 years. This
reflected not only the small
amount of study in natural
history, but also the
relative lack of economic
worth of lichens (28).

Schneider (27) in 1897
wrote a history of
lichenology, recognizing the
following periods:

I. From the earliest times to the end of the
seventeenth century.

II. From 1694, when Tournefort, the first
to separate lichens taxonomically from the
bryophytes, arranged plants into classes called
genera, to 1729.

III. From 1729, when Micheli divided
lichens into different orders, to 1779.

IV. From 1779, when Weber established
definite and reasoned lichen genera based on the
structure of thallus and fruits, to 1825.

V. From 1825, when Wallroth and
Meyer each published works dealing with
detailed morphological, ecological, and
biochemical observations, to 1868.

VI. From 1868, when Schwendener
discovered the dual nature of lichens, to 1894.

Other notable milestones
could include the arrangement
of all known lichens under
their respective genera by
Acharius in 1803, and the use
of spore characters in
classification by De Notaris
in 1846.

The discovery of the dual
nature of lichens by
Schwendener, in 1868, is
often considered to be the
beginning of modern
lichenology. This discovery
led to a wide variety of
ideas about classification.
This was demonstrated in a
report by Fink (8) who
conducted a survey of leading
botanists and lichenologists
throughout the world in an
attempt to analyze ideas
regarding the relationship of
lichen components. Ideas
expressed included the
following: lichens are a
distinct group of organisms
and should be separated from
fungi; lichens should be
classified in the fungus
genera that they closely
resemble; and, lichens have a
definite relationship to
fungi, but for convenience
should be kept separated from
the fungi.

Fink (9) gave his own idea
in the following statement:
“The lichen is a fungus which
lives all or a part of its
life in parasitic relation with an
algal host and also sustains
a relation with an organic or
an inorganic substratum.”
This idea has not been widely
accepted, and, despite Fink’s



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insisting that a lichen was
a fungus, he never attempted
to place lichens in existing
fungus genera. Perhaps a
more widely accepted idea
was given by Imshaug (22)
who defined a lichen as “an
entity capable of
reproducing itself, and
consisting of two organisms,
an alga and a fungus, living
together in a state of
symbiosis, as is manifested by
some change in the anatomy,
morphology, or physiology of
at least one of its
components.”

Ideas in regard to the
relationship of lichen
components, which in turn
have influenced taxonomic
ideas, have been further
complicated by the
observance of some lichens
that are apparently
parasitic on other plants,
and even on other lichens.

While the relationship
between lichens and fungi
has long been recognized, no
serious attempt has been
made to place lichens in
recognized fungus genera.

Other problems are related
to growth form and to
chemistry. Some workers make
no distinction on growth
form, but a more common
approach is to separate
groups purely on that basis
(i.e.,“foliose”,
“fruticose”, “squamulose”).

Lichens are also known to
contain a great many unique
chemicals, and in recent
times considerable emphasis
has been placed on the
chemistry of lichen

substances as an aid to
lichen taxonomy. This is well
illustrated by the work of
Asahina (1), and by the use
of chemical tests in almost
all keys identifying lichens.

It is not, however, been
accepted without opposition,
as indicated by the following
statement by Nearing (25):

Chemical dyes determine nothing, and in
most cases differentiate species only from
certain artificial “Species” invented for the
purpose of being thus differentiated. In certain
cases chemicals may be suggestive and helpful,
but the naming of hundreds of “species” on
the basis of chemical reactions alone, is in
itself sufficient proof that the reactions do not
coincide with evident relationships, that all
determinations should rest on botanical
characters, or else all on chemical, which last,
of course, would be absurd, but no more
absurd than the present mixing of the two
methods.

This likely represents an
extreme view, but does
illustrate the diversity of
ideas relating to the
classification of lichens.

Because of the uniqueness
of lichens, it is
understandable that serious
problems still exist in
lichen taxonomy; much work
remains to be done before
they can be satisfactorily
solved.

In the United States,
lichen studies have been
confined mostly to the east,
north, and Pacific Coast
areas with the southwest
largely being bypassed.
Oklahoma has barely 100
species reported from about
one fourth of its 77



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Keck, D.W.

counties, and no one has
attempted to do a complete
floristic work for even one
county. Kansas had less than
40 species reported until
Fearing (7) collected 163
taxa including about 140
species. He suggested that
this was perhaps only one
third of the total lichens
in the state. Arkansas and
Texas each has had only a
few species reported. This
compares with 245 species
reported from Ohio by Wolfe
(34), and 335 species
reported from Washington by
Howard (21), although
neither suggested the work
was complete in those
states.

It has always been
customary to report the
substrate from which a
particular lichen was
collected. While the finer
details of environment were
not included, this has at
least served as a beginning
for ecological studies. In
recent times specific
observations have been made
about light, temperature,
humidity, and other factors
in regard to various lichen
species, although little
work has been done on the
microenvironment.

Because of the limited
literature on lichens in the
southwest, and more
especially in Oklahoma, the
need for this present work
was seen.

REVIEW OF LITERATURE
Previous studies involving

the lichens of Oklahoma have
been very limited. Only two
earlier studies have been
made wherein appreciable
numbers of lichens from
Oklahoma were collected and
identified.

The first of these was by
Hedrick (20), who identified
specimens that Prof. Robert
Stratton from Oklahoma State
University collected from
Cimarron, Delaware, Harmon,
Johnston, Kay, Mayes,
McCurtain, Murray, Osage,
Payne, and Roger Mills
counties. These counties are
widespread throughout the
state and represent a great
diversity of habitats;
however, no attempt was made
to collect all the species
occurring in any area or
county. This study included
59 species. A majority of
these were crustose forms
occurring on rocks, with
Lecidea, Caloplaca, Buellia,
and Lecanora being the genera
most frequently represented.

The second work was by Hale
(16) who collected during
1955-1956 in six contiguous
counties in eastern Oklahoma
as part of a study of lichens
in the Ozarks. These counties
were Adair, Cherokee,
LeFlore, McCurtain,
Pushmataha, and Sequoyah.
This study involved “Macro-
corticolous” lichens and
included 47 species,



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primarily from the genera:
Parmelia, Physica, Usnea,
Anaptychia, Leptogium, and
Pannaria.

Since the latter study
dealt exclusively with
foliose and fruticose
lichens growing on trees,
and the previous study dealt
largely with crustose forms,
there was little duplication
of species collected. Only
seven species were common to
both lists, and collectively
these two efforts totaled 99
species.

In addition to these two
reports, a few additional
specimens representing eight
other species and involving
the additional counties of
Comanche, Noble, and Carter
have been reported in
monographs by Berry (2),
Imshaug (22), and Thomason
(31), and in other articles
by Hale (11, 14, 15, 18).
This gives a total of 107
species reported from
Oklahoma and involves 19 of
the 77 counties where at
least some collecting is
known to have been done.

Specimens reported by
Hedrick are in the herbarium
at the University of
Michigan, and those
collected by Hale are at the
Smithsonian Institution.
Other Oklahoma specimens are
preserved in herbaria of the
Missouri Botanical Garden,
New York Botanical Garden,
University of Wisconsin, and
the private herbarium of C.
W. Dodge at St. Louis.

THE STUDY AREA
Location and Size

The area involved in this
study includes the following
11 counties: Creek, Garfield,
Grant, Kay, Kingfisher,
Lincoln, Logan, Noble, Osage,
Pawnee, and Payne. It is
situated in North Central
Oklahoma (Fig. 1). It is
near-rectangular in shape,
and is about 115 miles long
(east to west), and averages
95 miles wide with the
greatest width being about
105 miles. It covers almost
11,000 square miles which is
between one sixth and one
seventh of the total area of
Oklahoma.

The area of each county, in
square miles, as computed
from General Highway County
Maps prepared by the Oklahoma
Department of Highways is as
follows:

County Total Area
(Sq. Mi.)

Creek 988
Garfield 1068
Grant 1008
Kay 938
Kingfisher 910
Lincoln 967
Logan 771
Noble 750
Osage 2393
Pawnee 597
Payne 696

Physiographic Regions

Bruner (3) divided the area
into two physiographic
regions: the Sandstone Hill
region which occupies



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Keck, D.W.

approximately the eastern 40
per cent of the area (Fig.
2), and the Redbeds Plains.
These regions are not
sharply delimited;
therefore, a wide zone
characterized by features of
both regions is noticeable.

The Sandstone Hills region
is composed largely of
weathered Pennsylvanian
shales, with rough low hills
of the more resistant
sandstone remaining. Maximum
height of these hills is 300
to 400 feet, although the
average is much less. Large
sandstone blocks cover the
tops and slopes of hills.
The broader flat-topped
ridges have soil of
sufficient depth to support
good plant growth.

The Redbeds Plains are
composed of soft red Permian
clays and shales, with some
thin sandstones which are
usually so soft that they do
not prduce escarpments. This
region is primarily one of
the rolling plains where
hills seldom exceed 100 feet
in height. The slightly
rolling plain to the west is
in contrast to the greater
relief near and in the
Sandstone Hills region.

Climate and Vegetation

North Central Oklahoma has
a continental type of
weather which is
characterized by a
pronounced seasonal range in
temperature. It has a mean
January temperature of 34o-

39oF and a mean July
temperature of 80o – 82o F.
Extremes in temperature vary
from several degrees below 0oF
to somewhat over 100oF. There
is an average of 200-220
frost-free days throughout
the area (32).

Rainfall averages vary from
about 30 inches per year in
the west to 40 in the east,
with considerable year-to-
year variation.
There is also an accompanying
small increase in relative
humidity toward the east.

The Sandstone Hills region
is covered with a
transitional oak forest
interspersed with grass
areas, while the Redbeds
Plains region is primarily
grassland, with trees usually
confined to the flood plains
of streams (3). A vegetation
map of the area taken from an
Oklahoma Game and Fish
Department map (Fig. 3).

Higher rainfall and
humidity favor the
development of most lichens,
at least within the limits of
conditions found in the study
area. Most of the effect is
apparently direct since there
is a definite change in each
category of the lichen flora
(terricolous, saxicolous,
corticolous) with a change in
rainfall. Some of the effect
is perhaps related only to
tree growth since a few
corticolous species are
considered to be specific in
regard to substrate.

There is also a correlation
between rainfall and pH, with



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the eastern part of the
study area where the
rainfall is greater being
primarily acidic and the
western part being near
neutral to slightly basic.
Considerable influence is
also exerted on the lichen
flora by the pH (17),
although this is more
related to the micro-
environment than to the pH
condition of the general
area.

All of these conditions
are interrelated and
therefore difficult to
evalulate individually, but
there is a pronounced change
in the lichen flora between
the eastern and western
limits of the study area.

PROCEDURES
Collecting stations were

selected in each county,
with desirability as a
collecting site being given
primary consideration.
General County Highway Maps
were used to determine the
general collecting areas;
then a survey of each of
these was made to determine
a collecting station. This
normally consisted of the
area immediately around a
stream or draw so that
collecting could be done
from trees, rocks, and
partially shaded soil since
nearly all lichens grow best
on one of these sugbstrates.
Less frequently, areas were
collected from where rock
was the predominant

substrate. Observations were
also made repeatedly on
exposed soil and fence posts,
ordinarily near the general
collecting areas, but only a
very limited number of
lichens were found and
collected under these
conditions.

Consideration was also
given to the size of counties
so that collecting stations
varied in number from four in
Pawnee County to 15 in Osage
County. This is double the
minimum of two collecting
stations for any county by
Hale (16). An attempt was
made to properly distribute
these stations within
counties and between adjacent
counties.

Over 1,000 specimens were
collected and are being
deposited with the lichen
collection in the Department
of Botany and Plant Pathology
of Oklahoma State University,
with duplicates of most of
these being kept in the
author’s private collection.

The seasonal aspect is
important primarily because
some lichens have mature
spores only during certain
seasons of the year (28).
Collecting was done at three
different times: (1) from
late May to early Auugust,
1959; (2) November, 1959; and
(3) April 1960.

Seventy-four collecting
stations were visited
including at least four
stations for each county
during the summer. After the
degree of repetition was
determined to be unusually



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Keck, D.W.

high among neighboring
stations used in the summer,
it was decided that a
considerably smaller number
of stations would sufficie
during subsequent collection
periods. Twenty-one stations
were visited during the
fall. This included one new
station in Kingfisher
County, and two stations
previously visited in each
of the other counties.
Eighteen stations including
three new areas were visited
in the spring. This included
at least one in each county.

The 78 collecting areas
are shown in Fig. 4. Their
descriptions, locations, and
dates visited are as
follows:

1. A wooded hillside near
highway 99, two miles
northeast of Cleveland, in
Osage County, May 25, 1959;
November 21, 1959.

2. Along a small stream,
three miles south of Wild
Horse store in Osage
County, May 25, 1959.

3. A wooded hillside with
limestone outcrops near
highway 20, five miles west
of Skiatook, in Osage
County, May 25, 1959; April
12, 1960.

4. A blackjack thicket near
highway 23, two miles north
of Barnsdall in Osage
County, May, 25, 1959.

5. Postoak woods, one mile
west of Hula Dam near
highway 10 in Osage County,
May 26, 1959.

6. A wooded ravine near a
county road west of highway
99, 12 miles north of
Pawhuska in Osage County,
May 26, 1959; November 21,
1959’ April 13, 1960

7. Along a creek in Osage
Hills State Park, two miles
south of highway 60 in Osage
County, May 26, 1959; April
14, 1960.

8. Sandstone bluffs near
highway 99, four miles
southeast of Pawhuska in
Osage County, May 27, 1959.

9. Sandstone outcrops near
highway 20, 12 miles west of
Pawhuska in Osage County,
May 27, 1959.

10. Along Bird Creek, four
miles east of Foraker in
Osage County, May 25, 1959.

11. Along a small stream,
three miles north and two
miles west of Webb City, In
Kay County, May 27, 1959.

12. Around Phillips Lake, two
miles south of Shidler in
Osage County, May 27, 1959.

13. Along the Salt Fork
River, two miles south of
Lamont in Grant County, May
28, 1959; November 26, 1959.

14. Along Deer Creek, 13
miles north of Lamont in
Grant County, May 28, 1959.

15. Along Crooked Creek,
three miles west of Wakita
in Grant County, May 28,
1959.

16. Along a wooded draw near
highway 81, one mile south



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of Medford in Grant County,
May 28, 1959.

17. Along the Salt Fork
River, two miles north of
Pond Creek in Grant County,
May 28, 1959; November 26,
1959.

18. Along the Salt Fork
River, three miles north of
Nash in Grant County, May
29, 1959.

19. A wooded ravine, three
miles southeast of

Hillsdale in Garfield
County, May 19, 1959.

20. Along Turkey Creek,
three miles north of
Drummond in Garfield
County, May 29, 1959.

21. Along Turkey Creek,
eight miles southeast of
Drummond in Garfield
County, May 29, 1959;
November 26, 1959.

22. Along Wolf Creek, three
miles southeast of Douglas
in Garfield County, May 29,
1959.

23. Open woods near
Breckenridge, eight miles
northeast of Enid in
Garfield County, May 29,
1959.

24. Along Red Rock Creek,
eight miles north of Garber
in Garfield County, May 29,
1959.

25. A rocky ravine near
highway 64, three miles
southeast of Covington in
Garfield County, May 29,
1959; April 15, 1960.

26. Postoak woods near
highway 33, one mile north
of Coyle in Payne County,
June 1, 1959.

27. Along Cottonwood Creek,
three miles south and one-
half mile west of Guthrie in
Logan County, June 1, 1959.

28. Along Bear Creek, one-
fourth mile west of Meridian
in Logan County, June 1,
1959; November 28, 1959.

29. A wooded draw, one-half
mile south of highway 105 -
40 junction, three miles
west of Tryon in Lincoln
County, June 2, 1959;
November 28, 1959.

30. Sandstone outcrops near
highway 40, one mile north
of Warwick in Lincoln
County, June 2, 1959.

31. Along Quapaw Creek north
of highway 62, six miles
west of Meeker in Lincoln
County, June 2, 1959.

32. Postoak woods near
highway 18, three miles
south of Chandler in Lincoln
County, June 2, 1959;
November 12, 1959.

33. A persimmon grove, three
miles south and one-half
mile west of Avery in
Lincoln County, June 5,
1959.

34. Postoak woods near
highway 27, three and one-
half miles east of Shamrock
in Creek County, June 5,
1959; November 24, 1959.

35. A rocky ravine near the
Cimarron River, eight miles



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Keck, D.W.

northeast of Cushing in
Payne County, June 5, 1959.

36. A wooded ravine, one
mile west of highway 99
near Arlington in Lincoln
County, June 6, 1959; April
11, 1960.

37. Postoak woods, three
miles south and nine miles
east of Stroud, in Creek
County, June 6, 1959.

38. Postoak woods, five
miles south and three miles
east of Bristow in Creek
County, June 6, 1959.

39. A wooded ravine near
highway 66, eight miles
northeast of Bristow in
Creek County, June 6, 1959.

40. Postoak woods, two miles
west of Sapulpa and one-
half mile south of highway
66 in Creek County, June 6,
1959.

41. Along Bridge Creek near
highway 51, 21 miles east
of Hennessey, in Logan
County, June 8, 1959;
November 28, 1959.

42. Along Skeleton Creek,
seven miles east and two
miles south of Hennessey in
Kingfisher County, June 8,
1959; November 26, 1959.

43. Along Turkey Creek, two
miles west of Kingfisher in
Kingfisher County, June 8,
1959; November 26, 1959.

44. Postoak woods, two miles
southeast of Dover in
Kingfisher County, June 8,
1959.

45. Along Cooper Creek, one-
half mile south of Loyal in
Kingfisher County, June 9,
1959.

46. A wooded draw, two and
one-half miles south of
highway 33, ten miles west
of Kingfisher in Kingfisher
County, June 9, 1959.

47. A wooded ravine between
highway 33 and the Cimarron
River, 15 miles east of
Kingfisher in Kingfisher
County, June 9, 1959; April
16, 1960.

48. A rocky draw, one mile
south of highway 74 – 33
junction, ten miles west of
Guthrie in Logan County,
June 9, 1959.

49. Postoak woods, two miles
north of Crescent in Logan
County, June 9, 1959; April
16, 1960.

50. Sandstone bluffs near
highway 77, eight miles
north of Guthrie in Logan
Countyk, June 9, 1959.

51. Along Black Bear Creek,
one mile north of Morrison
in Noble County, June 14,
1959.

52. Along the Arkansas River,
ten miles south and three
miles east of Ponca City in
Noble County, June 13, 1959.

53. A wooded draw, six miles
east and three miles south
of Billings in Noble County,
June 13, 1959; November 26,
1959.

54. A rocky ravine, near
highway 64, five miles west



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of Perry in Noble County,
June 13, 1959; April 15,
1960.

55. A wooded draw, near
highway 64, five miles east
of Perry in Noble County,
June 13, 1959; April 15,
1960.

56. A wooded ravine, four
miles northeast of Ripley
in Payne County, June 26,
1959; November 24, 1959;
April 11, 1960.

57. A rocky draw near
highway 108, four miles
south of Glencoe in Payne
County, July 7, 1959.

58. A wooded draw, two miles
north of Maramec in Pawnee
County, July 7, 1959;
November 21, 1959.

59. Wooded draw and hillside
near highway 64, six miles
southeast of Cleveland in
Pawnee County, July 7,
1959; April 12, 1960.

60. A wooded ravine near the
Arkansas River, one mile
north of Blackburn in
Pawnee County, July 7,
1959.

61. Sandstone bluffs, along
highway 15, eight miles
north of Morrison, in
Pawnee County, July 7,
1959; November 21, 1959.

62. Along the Arkansas
River, five miles southeast
of Oilton, in Creek County,
August 6, 1959; November
24, 1959.

63. Sandstone bluffs near
highway 51, two miles west

of Mannford in Creek County,
August 6, 1959, April 12,
1960.

64. A wooded hillside, one
mile west of highway 99 and
four miles north of Hominy
in Osage County, August 6,
1959.

65. Along Gray Horse Creek,
seven miles east of Fairfax
in Osage County, August 6,
1959.

66. Rocky bluffs near the
Arkansas River, five miles
west of Fairfax in Osage
County, August, 6, 1959.

67. Along the Arkansas River
near highway 60, nine miles
east of Ponca City in Osage
County, August 6, 1959.

68. Along the Salt Fork River
near highway 77, five miles
northeast of Marland in Kay
County, August 8, 1959.

69. Along the Salt Fork
River, two miles southwest
of Tonkawa in Kay County,
August 8, 1959.

70. A wooded draw near
highway 11, three miles
west of Blackwell in Kay
County, August 8, 1959.

71. Along the Chikaskia
River, two miles west of
Braman in Kay County, August
8;, 1959.

72. Along Deer Creek, four
miles northeast of Newkirk
in Kay County, August 8,
1959.

73. Along the Arkansas River,
six miles south and six
miles east of Newkirk in Kay



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County, August 8, 1959;
November 21, 1959; April
13, 1960.

74. A wooded ravine, four
miles west and one mile
south of Stillwater in
Payne County, August 8,
1959; October 9, 1959.

75. A wooded draw near
highway 81, seven miles
north of Enid in Garfield
County, November 26, 1959.

76. Along a rocky wooded
draw, one-fourth mile
southwest of Tuskegee in
Creek County, April 11,
1960.

77. Along the side and top
of a tall rocky hill, one
mile west of Mounds in
Creek County, April 11,
1960.

78. Along a rocky draw, six
miles northwest of Renfrow
in Grant County, April 15,
1960.

ECOLOGY
The area studied has many

variations in habitat. These
include timberland and
grassland, a considerable
range in average rainfall,
different types of rocks
such as limestone,
sandstone, and loosely
compacted silty clay, with
shaded and exposed soil, and
each having at least some
species restricted to that
particular environment.

As a consequence of this
diversity, few lichen
species are widespread in

the study area. Only
Candelaria fibrosa, Parmelia
bolliana, Physcia syncolla,
Teloschistes chrysophthalmus,
and Xanthoria candelaria were
collected in each county.
Others which are widely
distributed, occurring in at
least eight counties, are
Acarospora citrina, Caloplaca
aurantiaca, Candelaria
concolor (two varieties),
Lecanora muralis, Parmelia
reticulate, Physcia aipolia,
P. ciliata, P. orbicularis,
P. Tribacoides, and P.
stellaris.

A total of 64 species were
found in three counties or
less, and 30 of these were
collected in only one county
each. Table I lists all
species found and their
occurrence by counties.

Corticolous Species
Growth Forms

Foliose

The foliose forms occurring
primarily on trees, but
occasionally on rocks, are
Anaptychia granulifera, A.
heterochroa, A. hypoleuca, A.
speciosa, Candelaria concolor
var. concolor, C. concolor
var. effuse, C. fibrosa,
Collema conglomeratum, c.
subfurvum, Dermatocarpon
tuckermani, Leptogium
chloromelum, L. cyanescens,
Parmelia bolliana, P.
caperata, P. haitiensis, P.
reticulate, P. rudecta,
Physcia aipolia, P. ciliate,
P. elaeina, P. grisea, P.
millegrana, P. orgicularis,



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Keck, D.W.

P. tribacoides, P.
stellaris, P. syncolla,
Pyxine caesiopruinosa,
Teloschistes chrysophthalmus
(fruticose), and Xanthoria
candelaria.

Candelaria concolor, C.
fibrosa, Parmelia bolliana,
P. reticulate, Physcia
aipolia, P. orbicularis, P.
tribacoides, P. stellaris,
P. syncolla, Teloschistes
chrysophthalmus, and
Xanthoria candelaria occur
generally throughout the
area with Parmelia bolliana
and P. reticulate being more
abundant in the east (the
Sandstone Hills region) than
elsewhere, and Physcia
aipolia, P. syncolla, and
Xanthoria candelaria more
abundant in the west (the
Redbeds Plains area).
Extreme east refers to one
or more of Creek, Pawnee,
and Osage counties.
Westernmost refers to one or
more of Grant, Garfield, and
Kingfisher counties.

Those occurring only in
the extreme east include
Anapptychia granulifera, A.
hypoleuca, Dermatocarpon
tuckermani, Physcia
millegrana, and Pyxine
caesiopruinosa.

Other occurring over a
wider area, but not found in
the west are Anaptychia
heterochroa, Collema
conglomeratum, C. subfurvum,
Leptogium chloromelum, L.
cyanescens, Parmelia
caperata, P. haitiensis, P.
rudecta, and Physcia grisea.

The 29 “macro-corticolous”
lichens indicate the gradual

change from a more luxuriant
lichen flora in the Ozarks
where Hale (16) collected 63
species, which included all
but three of the above group.
He found 47 of these in six
eastern Oklahoma counties.
Only 11 of the 29 occur in
Grant, Garfield, and
Kingfisher counties.

Crustose

Crustose lichens found on
trees include Allarthonia
caesia, Buellia punctata, B.
schaereri, Caloplaca
aurantiaca, C. cerina, C.
chrysophthalma, C.
microphylina, C. ulmorum,
Candelariella xanthostigma,
Crocynia membranacea,
Lecanora hageni, L.
piniperda, L. subfusca, L.
varia, Lepraria chlorine,
Graphis scripta, Pertusaria
leioplaca, P. multipuncta,
and P. pustulata.

Only Caloplaca aurantiaca
was found in some abundance
throughout the area, while C.
chrysophthalma, C.
microphylina, Lecanora
hageni, L. varia, Pertusaria
leioplaca, P. multipuncta,
and P. pustulata also
occurred scattered in several
counties. Caloplaca
microphylina occurred only in
the west while the others
occurred primarily in the
east.

Other species were
collected infrequently but
indicated the following

distributional pattern:
Allarthonia caesia, Crocynia
membranacea, Lecanora



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Keck, D.W.

piniperda, L. subfusca,
Lepraria chlorine, Graphis
scripta, Pertusaria
leioplaca, P. multipuncta,
and P. pustulata occurred
only in the east, and
Candelariella xanthostigma
only in the west.

Microhabitat

Most of the foliose forms
were found on rough bark of
various woody species
although Teloschistes
chrysophthalmus and
Xanthoria candelaria were
often found on dead twigs or
bark of dead trees.

Crustose forms were
divided into three
categories: Caloplaca
microphylina, Buellia
punctata, and B. schaereri
which were found n decaying
fence posts or old stumps;
Allarthonia caesia, Lecanora
hageni, L. piniperda, L.
subfusca, Graphis scripta,
Pertusaria leioplaca, and P.
pustulata which were found
on smooth bark such as the
younger parts of hickory
(Carya sp.), hackberry
(Celtis laevigata), redbud
(Cercis canadensis), and red
oak (Quercus sp.); and
Caloplaca aurantiaca, c.
cerina, c. ulmorum, and
Pertusaria multipuncta which
were found on rough bark
such as post oak (Quercus
stellata), blackjack (Q.
marilandica), cottonwood
(Populus deltoids), and elm
(Ulmus Americana).

Saxicolous Species
Growth Forms

Foliose

Lichen species growing on
rocks about equal the number
growing on trees, but there
is a greater proportion of
crustose species. There are
only a few that can properly
be termed foliose; however,
several others which approach
this form are called
“squamulose” or “near-
foliose”. The first term
applies to small bits of
thallus that have the same
morphology as a foliose
lichen except that it is much
smaller in diameter. The
latter term applies to those
forms that have a poorly
developed thallus in the
center but are thicker and
lobed at the margins.

Lichens growing on rocks in
the above categories are
Dermatocarpon miniatum,
Endocarpon pusillum, Heppia
hassei, Lecanora muralis, L.
rubina, Lecidea rufonigra, L.
russellii, Parmelia
conspersa, P. isidiata, P.
obsessa, P. stenophylla,
Physcia halei, P. subtilis,
P. teretiuscula, and Rinodina
oreina.

Lecanora muralis is the
only species that occurs
throughout the area. There
are only a few species in the
west but this likely reflects
the extreme scarcity of rocks
in the three westernmost
counties. None of the species
occurs primarily in the west,
but Lecidea rufonigra, L.
russellii, Endocarpon



18
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Keck, D.W.

pusillum, and Heppia hassei
are rather intermediate
while the other species are
found exclusively in the
east.

According to Weaver and
Clements (33), foliose forms
normally follow crustose
forms in a xerosere, but
Oosting and Anderson (26)
indicate that crustose forms
sometimes decay a rock in
such a way that this
succession is not followed.
Both conditions were found
in the study area.

Crustose

The crustose rock forms
include the following
species: Acarospora citrine,
A. fuscata, A. smaragdula,
Bacidia granosa, B. umbrina,
Buellia alboatra, B.
novomexicana, B.
retrovertens, B. spuria, B.
stigmaea, B. vilis,
Caloplaca arizonica, C.
decipiens, C.
flavovirencens, C. lobulata,
C. murorum, Candelariella
vitellina, Diploschistes
actinostomus, d. scruposus,
var. scruposus, Lecania
californica, L. perproxima,
Lecanora atra, L. calcarea,
L. dispersa, L. melaena,
Lecidea tesselina,
Pertusaria pertusa, P.
velata, Placynthium nigrum,
Sarcogyne clavus, S.
pruinosa, S. simplex,
Verrucaria calsiceda, and V.
nigrescens.

Acarospora citrine,
Candelariella vitellina,
Diploschistes actinostomus,

and Sarcogyne clavus are
relatively widespread, but
somewhat more abundant in the
east. Lecanora calcarea is
also widespread, but more
frequent in the west. Others
occurring less frequently but
still not confined to a small
area are Acarospora fuscata,
Caloplaca flavovirescens, and
Lecanora dispersa.

The other species were
collected only a few times
each, or in a small area,
with the following
distribution pattern:
Acarospora smaragdula,
Bacidia granosa, Buellia
novomexicana, B. spuria, B.
stigmaea, Lecania
californica, Lecanora atra,
L. melanea, Pertusaria
pertusa, and Verrucaria
calciseda are found only in
the east, while Buellia
alboatra and B. vilis are
found only in the west. Many
in this group occupy an
intermediate to somewhat
easterly position.

Types of Rock

Another important
consideration is the type of
rock. Those species found on
limestone are Bacidia
granosa, Buellia alboatra, B.
vilis, Dermatocarpon
miniatum, Heppia hassei,
Lecania californica, L.
perproxima, Lecanora
calcarea, and Verrucaria
calsiceda.

Those found on both
sandstone and limestone are
Acaropsora fuscata, Caloplaca
murorum, Endocarpon pusillum,



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Keck, D.W.

Lecanora muralis, Lecidea
russellii, Placynthium
nigrum, and Sarcogyne
pruinosa, while the
remaining 35 species in this
group are confined to
sandstone.

There was a slight
north to south variation
since most of the limestone
was found in the north.

Terricolous Species

The remaining species of
lichens grow on soil or
loosely compacted silty
clay, sometimes being
closely associated with
rocks but not attached
directly to them.

Included are Cladonia
apodocarpa, C. capitata, C.
chlorophaea, C. fimbriata,
C. subcariosa, C. subtenuis,
C. uncialis, Coccocarpia
cronia, Dermatocarpon
hepaticum, Diploschistes
scruposus, var. bryophila,
Lecidea decipiens, Peltigera
canina, Staurothele
diffreactella, and S.
umbrina.

The Cladonia species occur
primarily on thin, moist,
shaded soil overlying
sandstone. Cladonia capitata
and C. chlorophaea are
readily found in the east
with C. subcariosa being
restricted to a smaller area
and also of less frequent
occurrence. The other
species are very infrequent
and occur only in the
extreme west. No Cladonia
species were found in

westernmost counties,
although sterile,
unidentifiable specimens were
found only a few miles away
in adjacent counties.

Coccocarpia cronia, which
also occurs on tree bases,
and Diploschistes scruposus
var. bryophila occur only in
the east and grow among
mosses over sandstone.

Staurothele diffractella,
s. umbrina, Dermatocarpon
hepaticum, and Lecidea
decipiens were found on
exposed soil, and occur only
in the west.

SUMMARY
Over 1000 specimens of

lichens were collected in an
11-county area of North
Central Oklahoma during 1959-
60. this included 111 species
and varieties representing 34
genera and 20 families. Their
identificfation involved the
use of 24 monographic studies
and other taxonomic
literature. Keys to various
taxa and a list of all
species are included. The
order of listing families
follows Fink (10) with
nomenclature following Hale
and Culberson (13). Families
having the greatest number of
species and varieties are
Physciaceae 17, Lecanoraceae
14, Parmeliaceae 12,
Caloplacaceae 10, and
Buelliaceae 9. These five
families contain 56 per cent
of all species found. Table I
shows the distribution by
counties for each taxon. A



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Keck, D.W.

tablular view of families is
shown in Table II.

The specimens are being
deposited with the lichen
collection of the Department of
Botany and Plant Pathology at
Oklahoma State University, and
duplicates of most of these are
in the author’s private
collection. Only five species
were collected in each county,
while ten othrs were collected
in at least eight counties.
Sixty-four species were
collected in three counties or
less and 30 of these were
collected in only one county
each. There was a much more
luxuriant lichen flora in the
east than in the west, with
little north to south
variation. Ecological
relationships are discussed and
remarks on location and habitat
are given for each species in
the “List of Species and
Habitats”.

Fifty-nine additions were
made to the lichen flora of
Oklahoma. These are given
special reference in the keys.

LIST OF SPECIES AND
HABITATS

[Ed. note: Nomenclature
updated in 2006 by Douglas M.
Ladd is in brackets]

The order of listing
families follows Fink (10)
while nomenclature follows
Hale and Culberson (13).
Each number is a collecting
number for a specimen. These
are referred to only for
species which represent
additions to the lichen
flora of Oklahoma. When more
than one specimen is cited

for a species, each substrate
is cited only once, and
specimens without the
substrate immediately
following the collecting
number have the same
substrate as the one
previously listed. Taxa
preceded by an asterisk have
not been previously reported
as components of the Oklahoma
lichen flora. Those species
among this group not followed
by special literature
citations were not included
in other literature pertinent
to Oklahoma lichens. (11, 14,
15, 16, 18, 20).

Pyrenulales
Verrucariaceae

*Verrucaria calsiceda DC.[= V.
calciseda DC.] is represented by
Keck 68 on limestone, five
miles west of Skiatook, in
Osage County.

*Verrucaria nigrescens Pers.
is represented by Keck 36 on
sandstone, two miles
northeast of Cleveland, in
Osage County, and by Keck
1075 on limestone, six miles
northwest of Renfrow in
Grant County.

*Staurothele diffractella
(Nyl.) Tuck. is represented
by Keck 519 on poorly
cemented silty clay, 15
miles east of Kingfisher in
Kingfisher County.

Staurothele umbrina (Ach.)
Tuck. [= S. fissa (Taylor) Zwackh]; on
poorly cemented silty clay;
Kingfisher County.



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Keck, D.W.

Dermatocarpaceae
*Dermatocarpon hepaticum
(Ach.) Th. Fr. [= Catapyrenium
cinereum (Pers.) Körber, but these
reports probably refer, at least in part, to
Placidium squamulosum (Ach.) Breuss]
is represented by Keck 995
on exposed soil, two miles
west of Kingfisher in
Kingfisher County.

*Dermatocarpon miniatum (L.)
Mann. [reports on calcareous
substrates probably refer to D.
muhlenbergii (Ach.) Műll. Arg.] is
represented by Keck 76 on
limestone, five miles west
of Skiatook, in Osage
County, and Keck 1056, six
miles south and six miles
east of Newkirk, in Kay
County.

Dermatocarpon tuckermani
(Rav.) Zahl.[= Placidium arboretum
(Michener) Lendemer]; on post
oak, in Osage County.

Endocarpon pusillum Hedw.
[local reports probably refer to E.
pallidulum (Nyl.) Nyl.] is
represented by Keck 374 on
sandstone, one mile north
of Warwick in Lincoln
County, and by Keck 762 on
limestone, four miles
northeast of Newkirk in Kay
County.

Hysteriales
Arthoniaceae

*Allarthonia caesia Fw. [=
Arthonia caesia (Flotow) Körber] is
represented by Keck 915 on
persimmon (Diospyros
virginiana), five miles
southeast of Oilton, in
Creek County; by Keck 57 on
willow (Salix nigra), three
miles south of Wild Horse

store, and Keck 849 on
redbud, 12 miles north of
Pawhuska, both in Osage
County.

Graphidaceae
*Graphis scripta (L.)Ach. is
represented by Keck 678 on
hackberry, one mile
northwest of Blackburn in
Pawnee County, and by Keck
939 on red oak, four miles
northeast of Ripley in Payne
County.

Lecanorales
Diploschistaceae

*Diploschistes actinostomus
(Pers.)Zahl.[(Ach.) Zahlbr.] is
represented by Keck 16 on
sandstone, two miles east of
Cleveland, in Osage County,
and by Keck 440, 361, 536,
584, and 672 in Creek,
Lincoln, Logan, Noble, and
Pawnee counties,
respectively.

Diploschistes scruposus
(Schreb.) Norm., var.
scruposus; on sandstone;
Lincoln, Logan, Osage, and
Payne counties.

*Diploschistes scruposus
(Schreb.) Norm., var.
bryophila (Ehrh.) Ach. [= D.
muscorum (Scop.) R. Sant.] is
represented by Keck 792 over
mosses, five miles southwest
of Stillwater in Payne
County, and by Keck 345 over
Cladonia sp. three and one-
half miles southwest of
Tryon in Lincoln County.

Collemaceae (5)
Collema conglomeratum Hoffm.:
on post oak; Lincoln,



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Keck, D.W.

Pawnee, Lincoln, Pawnee,
and Payne counties.

Collema subfurvum (Mull.
Arg.) Degel. [= C. subflaccidum
Degl.]; on post oak and red
cedar (Juniperus
virginiana); Creek,
Lincoln, Osage, and Payne
counties.

*Leptogium chloromelum (Sw.)
Nyl. [occurs rarely in midcontinental
North America; L. millegranum Sierk is
more common] is represented
by Keck 614 on cedar, four
miles northeast of Ripley
in Payne County, and by
Keck 796 on post oak, three
miles south of Chandler in
Lincoln County.

Leptogium cyanescens (Ach.)
Khr. [(Rabenh.) Körber]; on post
oak; Lincoln County.

Heppiaceae
*Heppia hassei Zahl. [= Peltula

obscurans Nyl. var. hassei (Zahlbr.)
Wetmore] is represented by
Keck 1067 on limestone, six
miles northwest of Renfrow
in Grant County, and by
Keck 78 and 268 from Osage
and Garfield counties,
respectively.

Pannariaceae
*Placynthium nigrum (Huds.)
S. Gray is represented by
Keck 1075B on limestone,
six miles northwest of
Renfrow in Grant County,
and by Keck 1081 on loosely
compacted silty clay, 15
miles east of Kingfisher in
Kingfisher County.

*Coccocarpia cronia Tuck. [= C.
palmicola (Sprengel) Arv. & D.J.

Galloway]; on moss-covered
sandstone; Creek County.

Peltigeraceae (31)
Peltigera canina (L.) Willd.;

[probably P. praetextata (Flörke ex
Sommerf.) Zopf]; on shaded mossy
banks, usually overlying
sandstone; Creek, Pawnee,
and Payne counties.

Lecideaceae
Lecidea decipens (Ehrh.) Ach.

[= Psora decipiens (Hedw.) Hoffm.]; on
open soil over Dermatocarpon
hepaticum; Grant,
Kingfisher, and Noble
counties.

Lecidea rufonigra (Tuck.)Nyl.
[= Psorula rufonigra (Tuck.) Gotth.
Schneid.] is represented by Keck
1010 on sandstone, one-half
mile southwest of Tuskegee
in Creek County.

Lecidea russellii Tuck. [= Psora
russellii (Tuck.) A. Schneid., a terricolous
species – these reports of saxicolous
populations refer to Psora pseudorussellii
Timdal]; on sandstone and
limestone; Lincoln, Logan,
and Osage counties.

*Lecidea tesselina Tuck. [=
Lecanora oreinoides (Körber) Hertel &
Rambold] is represented by
Keck 10 on sandstone, two
miles east of Cleveland, in
Osage County, and by Keck
113, 143, 871, and 880 at
other locations in Osage
County.

*Bacidia granosa (Tuck.)
Zahl. [= Bacidia coprodes (Körber)
Lettau] is represented by Keck
75 on limestone, five miles
west of Skiatook, in Osage
County.



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Keck, D.W.

*Bacidia umbrina (Ach.)
Bausch [= Scoliciosporum umbrinum)
Ach.) Arnold] is represented by
Keck 586 on sandstone,
seven miles west of Perry
in Noble County.

Cladoniaceae (16, 24)
*Cladonia apodocarpa Robbins
is represented by Keck 1053
on shaded soil, 12 miles
north of Pawhuska in Osage
County, and by Keck 1046,
one mile west of Mounds in
Creek County.

*Cladonia capitata (Michx.)
Spreng. [= C. peziziformis (With.) J.R.
Laundon] is represented by
Keck 784 on shaded soil,
five miles southwest of
Stillwater in Payne County
and by Keck 100, 663, 998,
and 1037 in Osage, Pawnee,
Logan, and Lincoln
counties, respectively.

Cladonia chlorophaea (Fik.)
Spreng. [possibly including or
consisting of this species and/or C.
cryptochlorophaea Asahina and C. grayi
G. Merr.]; on thin soil over
sandstone; Creek, Lincoln,
Osage, Pawnee, and Payne
counties.

*Cladonia fimbriata (L.) Fr.
[possibly, but not known from Oklahoma;
these reports more likely refer to the C.
chlorophaea complex (see previous
entry] is represented by Keck
388 on thin soil over
sandstone, three miles
south of Chandler in
Lincoln County.

*Cladonia subcariosa Nyl. [=
C. polycarpoides Nyl.] is
represented by Keck 26 on
shaded soil, two miles east
of Cleveland, in Osage

County, and by Keck 392C and
663B in Lincoln and Pawnee
counties, respectively.

*Cladonia subtenuis (des
Abbayes) Evans [(Abbayes) Mattick]
is represented by Keck 847
on slightly shaded soil, 12
miles north of Pawhuska in
Osage County.

*Cladonia uncialis (L.) Web.
[(L.) F.H.Wigg.] is represented by
Keck 1054 on slightly shaded
soil, 12 miles north of
Pawhuska in Osage County.

Acarosporaceae
Sarcogyne clavus (Ram.)
Krmph. is represented by
Keck 29 on sandstone, two
miles east of Cleveland, in
Osage County, and by Keck
465, 289, 359, 540, 595, and
674 in Creek, Garfield,
Lincoln, Logan, Noble, and
Pawnee counties,
respectively.

*Sarcogyne pruinosa (Sm.)
Kbr. [=S. regularis Körber.] is
represented by Keck 725 on
sandstone, seven miles east
of Fairfax in Osage county,
and by Keck 271 and 1073 on
limestone, in Garfield and
Grant counties,
respectively.

Sarcogyne simplex (Dav.) Nyl.
[= Polysporina simplex Davies) Vězda];
on sandstone; Logan County.

Acarospora citrina (Tayl.)
Zahl.; on sandstone; Creek,
Garfield, Lincoln, Logan,
Noble, Osage, Pawnee, and
Payne counties.



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Keck, D.W.

*Acarospora fuscata (Nyl.)
Arn. [(Schrader) Arnold] is
represented by Keck 82 on
limestone, five miles west
of Skiatook, and Keck 2 on
sandstone, two miles east
of Cleveland, both in Osage
County, and Keck 1033, 659,
and 642 in Lincoln, Pawnee,
and Payne counties
respectively.

*Acarospora smaragdula (Wh.)
Th. Fr. is represented by
Keck 40 on sandstone, two
miles east of Cleveland, in
Osage County, and by Keck
368 and 653 in Lincoln and
Pawnee counties,
respectively.

Pertusariaceae
*Pertusaria leioplaca
(Ach.)DC. [DC.; this report may
include or consist of P. paratuberculifera
Dibben, which is locally common on
hardwoods in the eastern portion of the
study area] is represented by
Keck 13 on blackjack oak,
two miles east of
Cleveland, in Osage County,
and by Keck 464, 382, 573,
and 611 on various woody
species in Creek, Lincoln,
Noble, and Payne counties,
respectively.

Pertusaria multipuncta
(Turn.) Nyl.; [= P.
multipunctoides Diben] on
blackjack oak, post oak,
and hickory; Creek,
Lincoln, Osage, and Payne
counties.

Pertusaria pertusa (L.)
Tuck.[North American reports of this
taxon are based on misidentifications;
several pre-1980 Missouri specimens
determined as this species are actually P.

plittiana Erichsen, and the sandstone
substrate cited here indicates a similar
possibility]; on sandstone; Creek
and Payne counties.

*Pertusaria pustulata (Ach.)
Duby is represented by Keck
415 on post oak, three and
one-half miles east of
Shamrock in Creek County,
and by Keck 368, 128, 669,
and 269 on various woody
species in Lincoln, Osage,
Pawnee, and Payne counties
respectively.

*Pertusaria velata (Turn.)
Nyl. is represented by Keck
1028 on sandstone, one mile
northwest of Ripley in Payne
County.

Lecanoraceae
Lecanora atra (Huds.) Ach. [=

Tephromela atra (Huds.) Hafellner]; on
sandstone; Creek County.

*Lecanora calcarea (L.)Smrft.
[=Aspicilia calcarea (L.) Mudd; local
reports may include or consist of A.
contorta (Hoffm.) Kremp.) is
represented by Keck 66 on
limestone, five miles west
of Skiatook, in Osage
County, and by Keck 270,
1070, 531, 558, and 905 in
Garfield, Grant, Logan,
Noble, and Pawnee counties,
respectively.

*Lecanora dispersa (Pers.)
Rohl. [= (Pers.) Sommerf.] is
represented by Keck 240 on
cement, three miles
southwest of Hillsdale in
Garfield County, and by Keck
538, 582, and 728 on
sandstone, in Logan, Noble,
and Osage counties
respectively.



Oklahoma Native Plant Record 25
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Keck, D.W.

Lecanora hageni Ach.; [= L.
hagenii (Ach.) Ach.]on hickory,
hackberry, persimmon, pecan
(Carya illinoensis), red
cedar, and various oaks;
Creek, Kay, Lincoln, Noble,
and Payne counties.

*Lecanora melaena (Hedl.)
Fink is represented by
Keck 38 on sandstone, two
miles east of Cleveland, in
Osage county.

Lecanora muralis (Schreb.)
Rabh.; on sandstone and
limestone; Creek, Garfield,
Kay, Logan, Noble, Osage,
Pawnee, and Payne counties.

*Lecanora piniperda Kbr.
[Brodo et al. (2001) restrict the range of
this taxon to boreal North America and
the Rocky mountains] is
represented by Keck 414 on
post oak, three and one-
half miles southwest of
Avery in Lincoln County,
and by Keck 853 on redbud,
12 miles north of Pawhuska
in Osage County.

*Lecanora rubina (Vill.)
Ach. [= Rhizoplaca chrysoleuca (Sm.)
Zopf] is represented by Keck
117 on sandstone, 12 miles
north of Pawhuska in Osage
County, and by Keck 948 and
686 in Noble and Pawnee
counties, respectively.

Lecanora subfusca (L.) Ach.
[= L. allophana Nyl., but probably refers
to L. hybocarpa (Tuck.) Brodo]; on
hackberry, persimmon, red
oak, hickory, redbud, ash
(Fraxinus americana), and
red cedar; Creek, Kay,
Lincoln, Osage, Pawnee, and
Payne counties.

Lecanora varia (Ehrh.)
Ach.[(Hoffm.) Ach.; almost certainly
refers to L. strobilina (Sprengel.) Keiffer];
on hickory, persimmon, post
oak, plum (Prunus sp.),
birch (Betula nigra),
redbud, ash, and cedar;
Creek, Kay, Lincoln, Osage,
Pawnee, and Payne counties.

*Lecania californica (Zahl.)
Fink is represented by Keck
71 on limestone, five miles
west of Skiatook, in Osage
County.

*Lecania perproxima
(Nyl.)Zahl. is represented
by Keck 1015 on sandstone,
one-half mile west of
Meridian in Logan County.

*Candelariella vitellina
(Ehrh.) Mull Arg. [(Hoffm.) Műll.
Arg.]; on sandstone; Creek,
Garfield, Lincoln, Logan,
Noble, Osage, Pawnee, and
Payne counties.

*Candelariella xanthostigma
(Pers.) Lett. [(Ach.) Lettau] is
represented by Keck 188 over
Physcia sp. on cottonwood,
two miles south of Lamont in
Grant County.

Parmeliaceae
Candelaria concolor (Dicks.)

[(Dicks.) Stein.] Arn. Var.
concolor; on elm, soapberry,
(Sapindus drummondii), red
cedar, hackberry, willow,
birch, mulberry (Morus
rubra), post oak, and
blackjack oak; Creek,
Garfield, Grant, Kingfisher,
Logan, Noble, and Osage
counties.



26
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Keck, D.W.

*Candelaria concolor
(Dicks.) Arn. var. effuse
(Tuck.) Merr. & Burnh. is
represented by Keck 982 on
Willow, three miles
southeast of Hillsdale in
Garfield County, and by
Keck 323 on elm and 899 on
bois-d’Arc in Logan and
Pawnee counties,
respectively.

Candelaria fibrosa (Fr.)
Mull. Arg.; on most woody
species and on sandstone;
Creek, Garfield, Grant,
Kay, Kingfisher, Lincoln,
Logan, Noble, Osage,
Pawnee, and Payne counties.

Parmelia bolliana Mull.
Arg.; [= Punctelia bolliana (Műll. Arg.)
Krog and/or Punctelia graminicola (de
Lesd.) Egan]; on most woody
species and on sandstone;
Creek, Garfield, Grant,
Kay, Kingfisher, Lincoln,
Logan, Noble, Osage,
Pawnee, and Payne counties.

Parmelia caperata (L.) Ach.;
[= Flavoparmelia caperata (L.) Hale
and/or F. baltimorensis (Gyeln. & Fơriss.)
Hale; blackjack oak,
sandstone, and thin soil
overlying sandstone; Creek,
Lincoln, Osage, and Payne
counties.

Parmelia conspersa
(Ehrh.)Ach.[= Xanthoparmelia
conspersa (Ach.) Hale; may refer to other
isidiate taxa of Xanthoparmelia, with a
dark lower cortex, such as X. mexicana
(Gyeln.) Hale]; on sandstone;
Creek and Osage counties.

*Parmelia haitiensis (Hale)[=
Parmotrema hatiense (Hale) Hale] is
represented by Keck 944 on
sandstone, four miles

northeast of Ripley in Payne
County, and by Keck 418,
348, and 841 on post oak or
black jack in Creek,
Lincoln, and Osage counties,
respectively.

Parmelia isidiata (Anzi)
Gyel.; on sandstone; Logan,
Noble, and Pawnee counties.

Parmelia obsessa (Ach.); [=
Myelochroa obsessa (Ach.) Elix & Hale];
on sandstone; Creek County.

Parmelia reticulate (Tayl.)[=
Parmotrema reticulatum (Taylor) M.
Choisy]; on blackjack oak,
post oak, redbud, elm, red
cedar, and sandstone; Creek,
Kay, Kingfisher, Lincoln,
Logan, Noble, Osage, Pawnee,
and Payne counties.

Parmelia rudecta Ach. [Punctelia
rudecta (Ach.) Krog; may also include
Punctelia missouriensis G. Wilh. & Ladd];
on blackjack oak and
sandstone; Lincoln, Osage,
and Payne counties.

Parmelia stenophylla (Achl.)
Heug. [= Xanthoparmelia
viriduloumbrina (Gyeln.) Lendemer]; on
sandstone; Osage County.

Caloplacaceae
*Caloplaca arizonica Rudolph
non Magn. [possibly C. subsoluta (Nyl.)
Zahlbr.] is represented by Keck
94 on sandstone, two miles
north of Barnsdall in Osage
county, and by Keck 1007 and
813 in Logan and Pawnee
counties, respectively.

Caloplaca aurantiaca
(Lightf.) Th. Fr.[= C.
flavorubescens (Huds.) J.R. Laundon];
on most woody species;
Creek, Garfield, Grant, Kay,



Oklahoma Native Plant Record 27
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Keck, D.W.

Kingfisher, Lincoln, Logan,
Noble, Osage, and Payne
counties.

Caloplaca cerina (Ehrh.) Th.
Fr.; on elm and cottonwood;
Garfield and Grant
counties.

*Caloplaca chrysophthalma
Degel. [while this taxon is frequent in
the region, it is a corticolous species]
is represented by Keck 462,
748,835,689, and 308 on
various woody species in
Creek, Kay, Osage, Pawnee,
and Payne counties,
respectively.

*Caloplaca dicipiens
(Arn.)Jatta. [(Arnold) Blomb. &
Forss.] is represented by
Keck 583 and 597 on
sandstone, seven miles west
of Perry in Noble County.

*Caloplaca flavovirescens
(Wulf.) D.T. & S. is
represented by Keck 4 on
sandstone, two miles east
of Cleveland, in Osage
County, and by Keck 432,
1012, and 568 from Creek,
Lincoln, and Noble
counties, respectively.

*Caloplaca lobulata
(Floerke) Hellb. [(Flörke) de
Lesd.]is represented by Keck
92 on sandstone, two miles
north of Barnsdall in Osage
County, and by Keck 376,
527, and 814 from Lincoln,
Logan, and Pawnee counties,
respectively.

*Caloplaca microphylina
(Tuck.) Hasse is
represented by Keck 278 on
mulberry, eight miles north
of Garber in Garfield

County, and by Keck 220,
757, 510, 410, and 576,
primarily on fence posts and
dead wood, in Grant, Kay,
Kingfisher, Lincoln, and
Noble counties,
respectively.

Caloplaca murorum (Hoffm.)
Th. Fr.; [= C. saxicola (Hoffm.) Nordin
(may represent misidentification)] on
sandstone and limestone;
Garfield, Osage, and Payne
counties.

*Caloplaca ulmorum Fink [(Fink)
Fink] is represented by Keck
233 on cottonwood, three
miles southeast of Hillsdale
in Garfield County, and by
Keck 178, 746, 521, 417, and
155 on elm, cottonwood, and
dead wood in Grant, Kay
Kingfisher, Logan, and Osage
counties, respectively.

Teloschistaceae (30)
Teloschistes chrysophthalmus
(L.) Th. Fr.; on various
woody species especially
dead twigs; Creek, Garfield,
Grant, Kay, Kingfisher,
Lincoln, Logan, Noble,
Osage, Pawnee, and Payne
counties.

Xanthoria candelaria (L.)
Arn. [possibly Xanthomendoza fulva
(Hoffm.) Søchting or Xanthomendoza fallax
(Hepp) Søchting]; on various
woody species, especially
bark of dead trees; Creek,
Garfield, Grant, Kay,
Kingfisher, Lincoln, Logan,
Noble, Osage, Pawnee, and
Payne counties.



28
Oklahoma Native Plant Record

Volume 6, Number 1, December 2006

Keck, D.W.

Buelliaceae (22)
*Buellia alboatra (Hoffm.)
Th. Fr.[= Diplotomma alboatrum
(Hoffm.) Flotow, possibly including or
consisting of D. venustum (Körber)
Körber] is represented by Keck
1079 on limestone, six
miles northwest of Renfrow
in Grant County, and by
Keck 1065 in Garfield
County. Imshaug (22) lists
this species as occurring
in several northeastern and
midwestern states extending
no farther south than
Colorado.

*Buellia novomexicana B. de
Lesd. [= B. tyrolensis Körber] is
represented by Keck 2 on
sandstone, two miles east
of Cleveland, in Osage
County, and by Keck 593 on
sandstone and 86 on
limestone in Noble and
Osage counties,
respectively.

Buellia punctata (Hoffm.)
Mass. [= Amandinea punctata (Hoffm.)
Coppins & Scheid.]; on sandstone
and dead wood; Garfield,
Grant, Osage, Pawnee, and
Payne counties.

*Buellia retrovertens Tuck.
is represented by Keck 581
on sandstone, seven miles
west of Perry in Noble
County, and by Keck 675
from Pawnee County. Imshaug
(22) lists this species as
occurring from Texas, New
Mexico, and Colorado
westward to the coast.

*Buellia schaereri Dnot. [may
represent misidentifications; this is a
boreal taxon that is restricted to

coniferous substrates (possibly
Amandinea sp.?)] is represented
by Keck 509 and 513 on dead
wood, one-half mile south of
Loyal, in Kingfisher County.
Imshaug (22) lists this
species as boreal, and
reported only from states
bordering Canada.

Buellia spuria (Schaer.)
Anzi; on sandstone; Osage
County.

*Buellia stigmaea Tuck. [= B.
maculate Bungartz?] Is represented
by Keck 882 on sandstone,
two miles east of Cleveland,
in Osage County, and by Keck
439 and 711 in Creek County.
Imshaug (22) lists this
species from several
southern states but not
farther west than Missouri.

*Buellia vilis Th.Fr. is
represented by Keck 67 on
limestone, five miles west
of Skiatook, in Osage
County. Imshaug (22) lists
this species only from North
Dakota and Colorado.

Rinodina oreina (Ach.) Mass.
[=Dimelaena oreina (Ach.) Norman]; on
sandstone; Osage and Pawnee
counties.

Physciaceae (23)
*Pyxine caesiopruinosa
(Tuck.) Imshaug [possibly P.
subcinerea Stirton] is represented
by Keck 702 on red oak, five
miles southeast of Oilton,
in Creek County. Imshaug
(23) lists this species as
occurring only from several
Gulf Coast states.



Oklahoma Native Plant Record 29
Volume 6, Number 1, December 2006

Keck, D.W.

Physcia aipolia (Ehrh.)
Hampe [(Ehrh. Ex Humb.) Fűrnr.]; on
various woody species;
Garfield, Grant, Kay,
Kingfisher, Lincoln, Logan,
Noble, Osage, Pawnee, and
Payne counties.

Physcia ciliata (Hoffm.)
DR.[= Phaeophyscia ciliata (Hoffm.)
Moberg]; on soapberry, elm,
cottonwood, post oak, and
ash; Garfield, Grant, Kay,
Lincoln, Noble, Osage,
Pawnee, and Payne counties.

Physcia elaeina (Sm.) A. L.
Sm. [= Hyperphyscia adglutinata
(Flörke) H. Mayrhofer & Poelt]; on
cottonwood and hickory;
Garfield, Grant, and Osage
counties

Physcia grisea (Lam.) Zahl.
[probably Physconia leucoleiptes (Tuck.)
Essl.]; on blackjack oak,
post oak, hickory, and elm;
Creek, Lincoln, Noble, and
Osage counties.

*Physcia halei Thomson is
represented by Keck 816 on
sandstone, eight miles
north of Morrison, in
Pawnee County, and by Keck
115, 134, and 875 from
Osage County.

Physcia millegrana Degel.;
on birch; Osage County.

Physcia orbicularis (Neck.)
Poetsch. [= Phaeophyscia orbicularis
(Necker) Moberg, but probably other
sorediate species including P. adiastola
(Essl.) Essl, P. insignis ) Mereschk.)
Moberg, P. pusilloides (Zahlbr.) Essl. or
P. rubropulchra (Degel.) Essl.]; on
various woody species and
limestone; Creek, Garfield,
Kay, Kingfisher, Logan,

Noble, Osage, and Payne
counties.

Physcia stellaris (L.) Nyl.;
on various woody species and
sandstone; Creek, Garfield,
Kay, Kingfisher, Lincoln,
Logan, Noble, Osage, Pawnee,
and Payne counties.

*Physcia syncolla Tuck. [=
Hyperphyscia syncolla (Tuck ex Nyl.) Kalb]
is represented by Keck 1041
on hackberry, one mile west
of Mounds in Creek County,
and by Keck 197, 737, 480,
383, 319, 556, 156, 645, and
635 on various woody species
from Garfield, Grant, Kay,
Kingfisher, Lincoln, Logan,
Noble, Osage, Pawnee, and
Payne counties,
respectively.

Physcia teretiuscula (Ach.)
Lynge [= P. dubia (Hoffm.) Lettau;
possibly Speerschneidera euploca (Tuck.)
Trevisan or Physcia subtilis Degel.]; on
sandstone; Creek County.

Physcia tribacoides Nyl.;
[possibly P. americana G. Merr.] on
post oak, blackjack oak,
elm, hackberry, persimmon,
red cedar, and sandstone;
Creek, Grant, Kay, Lincoln,
Logan, Osage, and Payne
counties.

Anaptychia granulifera (Ach.)
Mass.; [= Heterodermia granulifera
(Ach.) W.L. Culb.] on post oak;
Osage County.

Anaptychia heterochroa Vain.[=
Heterodermia obscurata (Nyl.) Trevisan];
on various oaks, hickory,
and sandstone; Creek,
Lincoln, Osage, Pawnee, and
Payne counties.



30
Oklahoma Native Plant Record

Volume 6, Number 1, December 2006

Keck, D.W.

Anaptychia hypoleuca (Muhl.)
Vain. [= Heterodermia hypoleuca
(Muhl.) trevisan]; on post oak
and red oak; Creek and
Osage counties.

Anaptychia speciosa (Wulf.)
Mass. [= Heterodermia speciosa
(Wulfen) Trevisan]; on sandstone;
Payne County.

Lichenes Imperfecti
Leprariaceae

*Lepraria chlorina Ach.
[= Chrysothrix chlorina (Ach.) J.R.
Laundon, but probably Chrysothrix
candelaris (L.) J.R. Laundon] is
represented by Keck 1052 on
birch, in Osage Hills State
Park in Osage County.

*Crocynia membranacea
(Dicks.) Zahl. [probably
Lepraria lobificans Nyl.] is
represented by Keck 307 on
post oak, one mile north of
Coyle in Payne County, and
by Keck 422, 353, 315, 562,
and 140 on various oaks or
sandstones from Creek,
Lincoln, Logan, Noble, and
Osage counties,
respectively.

SELECTED BIBLIOGRAPHY
*General works not cited in text

1. Asahina, Y. 1954. Chemistry of
Lichen Substances. Tokyo.

2. Berry, E. C. 1941. A
Monograph of the Genus Parmelia in
North America, North of Mexico
Ann. Mo. Bot. Gard.
28:31-146.

3. Bruner, W. E. 1931. The
Vegetation of Oklahoma.
Ecological Monographs.
1:100-188.

*4. Culberson, William L.
1955. A Guide to the Literature on the
Lichen Flora and Vegetation of the United
States. U.S.D.A. Agri. Res.
Service. Plant Disease
Epidemics and
Identification Section.
Beltsville, MD. Special
Publication 7:1-54.

5. Degelius, Gunnar. 1954. The
Lichen Genus Collema in Europe.
Symb. Bot. Upsal. 12
(2):1-499.

6. Evans, A. W. 1930. The
Cladoniae of Connecticut. Trans.
Conn. Acad. Arts and
Sciences. 30:357-510.

7. Fearing, O. S. 1952.
Preliminary Study of the Taxonomy and
Ecology of Kansas Lichens. Univ.
Kans. Sci. Bull. 35:543-
575.

8. Fink, Bruce. 1911. The Nature
and Classification of Lichens—I. Views and
Arguments of Botanists Concerning
Classification. Mycologia.
3:231-269.

9. Fink, Bruce. 1913. The Nature
and Classification of Lichens—II. The
Lichen and Its Algal Host.
Mycologia. 5:97-166.

10. Fink, Bruce. 1935. The Lichen
flora of the United States. Univ. of
Mich. Press, Ann Arbor.

11. Hale, Mason E., Jr. 1952.
Studies on the Lichen Rinodina Oreina in
North America. Bull. Tor.
Bot. Cl. 79:251-259.

12. Hale, Mason E., Jr. 1955.
Xanthoparmelia in North America I. The
Parmelia Conspersa-Stenophylla group.
Bull. Tor. Bot. Cl.
82:9-21.

13. Hale, Mason E., Jr. and
William L. Culberson.
1956. A Checklist of the Lichens of the
United States, Canada, and Alaska.
Castenea. 21:73-105.



Oklahoma Native Plant Record 31
Volume 6, Number 1, December 2006

Keck, D.W.

14. Hale, Mason E, Jr. 1956.
Chemical Strains of the Parmelia
Conspersa-Stenophylla Group in South
Central United States. Bull.
Tor. Bot. Cl. 83:218-
220.

15. Hale, Mason E., Jr.
1956. A Note on Lichenes Americani
Exsiccati, fascicle I. The
Bryologist. 59:41-43.

16. Hale, Mason E., Jr.
1957. Corticolous Lichen Flora of the
Ozark Mountains. Trans. Kans.
Acad. Sci. 60:155-160.

17. Hale, Mason E., Jr.
1957. Lichen Handbook.
Smithsonian Institution
Washington, D. C.

18. Hale, Mason E., Jr.
1958. The Status of Usnea
Diplotypus in North America. The
Bryologist. 61:247-248.

19. Hale, Mason E., Jr.
1959. New or Interesting Species of
Parmelia in North America. The
Bryologist. 62:20-21.

20. Hedrick, Joyce. 1930.
Lichens from the State of Oklahoma.
Papers Mich. Acad.
Sci., Arts, and
Letters. 12:101-110.

21. Howard, Grace E. 1950.
Lichens of the State of Washington.
Univ. of Wash. Press,
Seattle.

22. Imshaug, H. A. 1951. The
Lichen-forming Species of the Genus
Buellia in the United States and Canada.
Univ. Microfilms Publ.
2607, Ann Arbor,
Michigan.

23. Imshaug, H. A. 1957. The
Lichen Genus Pyxine in North and
Middle America. Trans. Amer.
Micros. Soc. 76:246-
269.

24. Luttrell, E. S. 1954. The
Cladoniaceae of Virginia. Lloydia.
17:275-306.

25. Nearing, G. G. 1947. The
Lichen Book. Published by the
author, Ridgewood, New
Jersey.

26. Oosting, H.J. and L. E.
Anderson. 1937. The Vegetation
of a Barefaced Cliff in Western North
Carolina. Ecology. 18:280-
292.

27. Schneider, Albert. 1897.
A Text-Book of General Lichenology.
Wm. N. Clute and
Company, Binghamton.

28. Smith, Annie L. 1921.
Lichens. Cambridge Univ.
Press, Cambridge.

29. Snyder, L.C. 1917. Geography
of Oklahoma. Okla. Geol.
Survey. Bull. 27:1-325.

30. Thomson, John W., Jr.
1949.The Teloschistaceae of Wisconsin-
Papers on Wisconsin Lichens III. Amer.
Midl. Nat. 41:1-68.

31. Thomson, John W., Jr.
1950. The Species of Peltigera of North
America, North of Mexico. Amer.
Midl. Nat. 44:1-68.

32. United States Department
of Agriculture. 1941.
Yearbook of Agriculture.
Climate and Man. (Washington,
D. C.) Gov’t. Printing
Office: 1165-1174.

33. Weaver, John E. and
Frederick E. Clements.
1938. Plant Ecology. McGraw-
Hill Book Co., New York:
66-68.

34. Wolfe, John N. 1940. A
Catalog of the Lichens of Ohio. Ohio
Biol. Sur. Bull. 36:1-50.

*35. Zahlbruckner, A. 1922-
40. Catalogus Lichenus Universalis.
Leipzig. 10 vol.



32
Oklahoma Native Plant Record

Volume 6, Number 1, December 2006

Keck, D.W.

TABLE I SPECIES OCCURRENCY BY COUNTIES

C
r
e
e
k

G
a
r
f
i
e
l
d

G
r
a
n
t

K
a
y

K
i
n
g
f
i
s
h
e
r

L
i
n
c
o
l
n

L
o
g
a
n

N
o
b
l
e

O
s
a
g
e

P
a
w
n
e
e

P
a
y
n
e

Acarospora
citrine

X X X X X X X X

A. fuscata X X X X X

A.smaragdula X X X

Allarthonia
caesia

X X

Anaptychia
granulifera

X

A. heterochroa X X X X X

A. hypoleuca X X

A. speciosa X

Bacidia granosa X

B. umbrina X

Buellia
alboatra

X X

B. novomexicana X X X

B. punctata X X X X X

B. retrovertens X X

B. schaereri X

B. spuria X

B. stigmaea X X

C G G K K L L N O P P



Oklahoma Native Plant Record 33
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Keck, D.W.

B. vilis X

Caloplaca
arizonica

X X X

C. aurantiaca X X X X X X X X X X

C. cerina X X

C.
chrysophthalma

X X X X X X

C. decipiens X

C.
flavovirescens

X X X X

C. lobulata X X X X

C. microphylina X X X X X X

C. murorum X X X

C. ulmorum X X X X X X

Candelaria
concolor var.
concolor

X X X X X X X

C. concolor
var. effusa

X X X

C. fibrosa X X X X X X X X X X X

Candelariella
vitellina

X X X X X X X

C. xanthostigma X

Cladonia
apodocarpa

X X

C G G K K L L N O P P



34
Oklahoma Native Plant Record

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Keck, D.W.

C. capitata X X X X X

C. chlorophaea X X X X X

C. fimbriata X

C. subcariosa X X X

C. subtenuis X

C. uncialis X

Coccocarpia
cronia

X

Collema
conglomeratum

X X X X

C. subfurvum X X X X

Crocynia
membranacea

X X X X X X

Dermatocarpon
hepaticum

X X X

D. miniatum X X

D. tuckermani X

Diploschistes
actinostomus

X X X X X X X

D. scruposus
var. scruposus

X X X X

D. scruposus
var. bryophila

X X

Endocarpon
pusillum

X X

Graphis scripta X X

C G G K K L L N O P P



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Heppia hassei X X X

Lecania
californica X
L. perproxima X

Lecanora atra X

L. calcarea X X X X X X X

L. dispersa X X X X X

L. hageni X X X X X

L. melaena X

L. muralis X X X X X X X X

L. piniperda X X

L. rubina X X X

L. subfusca X X X X

L. varia X X X X X X

Lecidea
decipiens

X X X

L. rufonigra X X

L. russellii X X X

L. tesselina X



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C
r
e
e
k

G
a
r
f
i
e
l
d

G
r
a
n
t

K
a
y

K
i
n
g
f
i
s
h
e
r

L
i
n
c
o
l
n

L
o
g
a
n

N
o
b
l
e

O
s
a
g
e

P
a
w
n
e
e

P
a
y
n
e

Lepraria
chlorine

X

Leptogium
chloromelum

X X

L. cyanescens X

Parmelia
bolliana

X X X X X X X X X X X

P. caperata X X X X

P. conspersa X X

P. haitiensis X X X X

P. isidiata X X X

P. obsessa X

P. reticulata X X X X X X X X X

P. rudecta X X X

P. stenophylla X

Peltigera
canine

X X X

Pertusaria
leioplaca

X X X X X X

P. multipuncta X X X X

P. pertusa X X

P. pustulata X X X X X

P. velata X

C G G K K L L N O P P



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Physcia aipolia X X X X X X X X X X

P. ciliata X X X X X X X X

P. elaeina X X X

P. grisea X X X X

P. halei X X

P. millegrana X

P. orbicularis X X X X X X X X

P. stellaris X X X X X X X X X X

P. subtilis X X

P. syncolla X X X X X X X X X X X

P. teretiuscula X

P. tribacoides X X X X X X X

Placynthium
nigrum

X X X

Pyxine
caesiopruinosa

X

Rinodina oreina X X

Sarcogyne
clavus

X X X X X X X

S. pruinosa X X X

C
r
e

G
a
r

G
r
a

K
a
y

K
i
n

L
i
n

L
o
g

N
o
b

O
s
a

P
a
w

P
a
y



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S. simplex X

Staurothele
diffractella

X

S. umbrina X

Teloschistes
chrysophthalmus

X X X X X X X X X X X

Verrucaria
calsiceda

X

V. nigrescens X X

Xanthoria
candelaria

X X X X X X X X X X X

Total species 44 29 24 21 19 45 32 33 77 41 42



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TABLE II TABULAR VIEW OF THE FAMILIES

Families Genera
Species and
subordinate
taxa

Verrucariaceae 2 4
Dermatocarpaceae 2 4
Arthoniaceae 1 1
Graphidaceae 1 1
Diploschistaceae 1 3
Collemaceae 2 4
Heppiaceae 1 1
Pannariaceae 2 2
Peltigeraceae 1 1
Lecideaceae 2 6
Cladoniaceae 1 7
Acarosporaceae 2 6
Pertusariaceae 1 5
Lecanoraceae 3 14
Parmeliaceae 2 12
Caloplacaceae 1 10
Teloschistaceae 2 2
Buelliaceae 2 9
Physciaceae 3 17
Leprariaceae 2 2
TOTALS 20 34 111



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TAXONOMIC TREATMENT
Keys to Various Taxa

Key to Classes, Orders, and Families
1. Thallus of more or less definite form, usually with ascocarps.

Ascolichenes
1. Thallus of entangled hyphae, without ascocarps . . . . . .

Lichenes Imperfecti
Ascolichenes

1. Hymenium produced in a perithecium . . . . . . .Pyrenulales
1. Hymenium produced in an apothecium

2. Thallus rudimentary; apothecia irregular, linear or
oblong. . . . . . . . . . . . . . . . . . . . Hysteriales

2. Thallus commonly well developed; apothecia more or less
round or cuplike . . . . . . . . . . . . . . Lecanorales

Pyrenulales
1. Thallus crustose . . . . . . . . . . . . . . Verrucariaceae
1. thallus squamulose or foliose. . . . . . . Dermatocarpaceae

Hysteriales
1. Apothecia without an exciple . . . . . . . . .Arthonicaceae
1. Apothecia with an exciple. . . . . . . . . . . Graphidaceae

Lecanorales
1. Phycobiont a species of Myxophyceae

2. Thallus squamulose to foliose, taking its form from that
of the phycobiont . . . . . . . . . . . . . .Collemaceae

2. Thallus foliose to granulose; not taking its form from
the phycobiont
3. Thallus large, plainly foliose . . . . . Peltigeraceae
3. Thallus smaller, somewhat foliose to granulose

4. Spores many per ascus . . . . . . . . . Heppiaceae
4. spores 8 per ascus . . . . . . . . . . Pannariaceae

1. Phycobiont a species of Chlorophyceae
5. Apothecia with both thalloid and proper exciples. . . .

Diploschistaceae
5. Apothecia with either thalloid or proper exciple, but

not with both
6. Thallus two-fold, having both squamules and podetia. .

Cladoniaceae
6. Thallus otherwise

7. Spores brown
8. Thallus crustose to scaly . . . . . . Buelliaceae
8. Thallus foliose . . . . . . . . . . . Physciaceae

7. Spores hyaline
9. Spores very large, commonly up to 200 microns



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long . . . . . . . . . . . . . . . Pertusariaceae
9. spores smaller

10. Thallus plainly foliose to fruticose
11. Thallus generally small, yellowish in color

Teloschistaceae
11. Thallus larger, not yellowish. Parmeliaceae

10. Thallus crustose to squamulose, or near
foliose
12. Spores minute, many per ascus . . . . . . .

Acarosporaceae
12. Spores larger, usually eight per ascus

13. Apothecia with proper exciple . . . . . .
Lecideaceae

13. Apothecia with thalloid exciple
14. Apothecia usually yellowish, spores

usually one-septate with cells polar
Caloplacaceae

14. Apothecia rarely yellowish, spores
non-septate, septate, or muriform, but
not polar . . . . . . . . Lecanoraceae

Lichenes Imperfecti
The only family representing this group is Leprariaceae.



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KEY TO GENERA AND SPECIES

Pyrenulales
Verrucariaceae

1. Spores non-septate . . . . . . . . . . . . . 1. Verrucaria
1. Spores muriform. . . . . . . . . . . . . . . 2. Staurothele

1. Verrucaria
1. Thallus grayish-white . . . . . . . . . . . . V. calciseda
1. Thallus brownish or greenish to black . . . . V. nigrescens

2. Staurothele
1. Spores two per ascus . . . . . . . . . . . . . . S. umbrina
1. Spores eight per ascus . . . . . . . . . . .S. diffractella

Dermatocarpaceae
1. Spores non-septate . . . . . . . . . . . . 1. Dermatocarpon
1. Spores muriform. . . . . . . . . . . . . . . 2. Endocarpon

1. Dermatocarpon
1. Thallus umbilicate, foliose . . . . . . . . . . D. miniatum
1. Thallus adnate, squamulose

2. Squamules round with elevated margins, not contiguous
D. hepaticum

2. Squamules lobed and overlapping. . . . . . D. tuckermani
2. Endocarpon

The only species representing this genus is E. pusillum.

Hysteriales
Arthoniaceae

The only genus representing this family is Allarthonia and the
only species is A. caesia.

Graphidaceae
The only genus representing this family is Graphis and the
only species is G. scripta.

Lecanorales
Collemaceae

1. Cortex of interwoven hyphae. . . . . . . . . . . 1. Collema
2. Cortex plectenchymatous. . . . . . . . . . . . 2. Leptogium

1. Collema
1. Thallus isidiate . . . . . . . . . . . . . . . C. subfurvum
2. Thallus without isidia . . . . . . . . . . C. conglomeratum

2. Leptogium
1. Thallus brown to black, lobes crenate. . . . L. chloromelum
2. Thallus bluish-slate colored, lobes entire . . L. cyanescens



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Heppiaceae
The only genus representing this family is Heppia and the only
species is H. hassei.

Pannariaceae
1. Thallus with lower cortex wanting. . . . . . 1. Placynthium
1. Thallus with lower cortex. . . . . . . . . . 2. Coccocarpia

1. Placynthium
The only species representing this genus is P. nigrum.

2. Coccocarpia
The only species representing this genus is C. cronia.

Peltigeraceae
The only genus representing this family is Peltigera and the
only species is P. canina.

Diploschistaceae
The only genus representing this family is Diploschistes.
1. Proper exciple thick, radiately striate. . .D. actinostomus
1. Proper exciple thin, minutely toothed

2. Apothecia often 1 mm across, on soil and rocks. . . . .
D. scruposus var. scuposus

2. Apothecia usually 0.3 mm or less, on moss and Cladonia
sp. . . . . . . . . . . . . . D. scruposus var. bryophila

Lecideaceae
1. Spores non-septate . . . . . . . . . . . . . . . 1. Lecidea
1. Spores septate . . . . . . . . . . . . . . . . . 2. Bacidia

1. Lecidea
1. Thallus crustose . . . . . . . . . . . . . . . L. tesselina
1. Thallus squamulose

2. Squamules brownish to green on the edge . . L. rufonigra
2. Squamules whitish on the edge

3. Apothecia blackish. . . . . . . . . . . . L. decipiens
3. Apothecia reddish-brown . . . . . . . . . L. russellii

2. Bacidia
1. Spores acicular. . . . . . . . . . . . . . . . . B. umbrina
1. Spores fusiform . . . . . . . . . . . . . . . . B. granosa

Cladoniaceae
The only genus representing this family is Cladonia.
1. Podetia extremely short or absent. . . . . . C. apodocarpa
1. Podetia longer

2. Podetia repeatedly branched, squamules evanescent
3. Podetia dichotomously branched. . . . . . C. subtenuis
3. Podetia variously branched, but not dichotomous. . . .

C. uncialis



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2. Podetia little branched, squamules persistent.
4. Podetia cup-shaped or trumpet-shaped

5. Podetia granular. . . . . . . . . . C. Chlorophaea
5. Podetia with fine, dusty soredia . . . C. fimbriata

4. Podetia not forming cups
6. Fruits at least twice as broad as the podetia. . .

C. capitata
6. Fruits barely broader than the podetia . . . . .

C. subcariosa

Acarosporaceae
1. Apothecia with thalloid exciple . . . . . . . 1. Acarospora
1. Apothecia with proper exciple . . . . . . . . 2. Sarcogyne

1. Acarospora
1. Thallus whitish-yellow to green. . . . . . . . A. citrina
1. Thallus brown

2. Aeroles lobed, margins elevated . . . . . . A. fuscata
2. Aeroles entire, margins depressed . . . . A. smaragdula

2. Sarcogyne
1. Apothecia pruinose . . . . . . . . . . . . . . S. pruinosa
1. Apothecia not pruinose

2. Apothecia 0.2-1.0 mm. across, red when wet. . S. simplex
2. Apothecia 0.7-2.0 mm. across, not red when wet S. clavus

Pertusariaceae
The only genus representing this family is Pertusaria.
1. Spores four per ascus. . . . . . . . . . . . . P. leioplaca
1. Spores one or two per ascus

2. Fruiting knobs sorediate . . . . . . . . .P. multipuncta
2. Fruiting knobs esorediate

3. Fruiting knobs postulate, some finally opening into a
disk. . . . . . . . . . . . . . . . . . . P. pustulata

3. Fruiting knobs not postulate, or if postulate, then
not opening into a disk
4. Spores one per ascus. . . . . . . . . . P. velata
4. Spores two per ascus. . . . . . . . . . P. pertusa

Lecanoraceae
1. Thallus yellow . . . . . . . . . . . . . . 1. Candelariella
1. Thallus not yellow

2. Spores aseptate. . . . . . . . . . . . . . . 2. Lecanora
2. Spores septate . . . . . . . . . . . . . . . 3. Lecania

1. Candelariella
1. Thallus persistent; on rocks. . . . . . . . . C. vitellina
1. Thallus evanescent; on trees. . . . . . . . C. xanthostigma



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2. Lecanora
1. Epithecia intensely black . . . . . . . . . . . . . L. atra
1. Epithecia variously colored, but not black

2. Thallus aerolate to squamulose. . . . . . . . .L. rubina
2. Thallus crustose or near-foliose

3. Thallus near-foliose. . . . . . . . . . . . L. muralis
3. Thallus crustose

4. Apothecia 0.1-0.25 mm. across. . . . . L. piniperda
4. Apothecia somewhat larger to much larger

5. Apothecia pruinose
6. Spores subglobose, 9-15 microns wide . . . . .

L. calcarea
6. Spores ellipsoid, 406 microns wide . . . . . .

L. hageni
5. Apothecia not pruinose

7. Apothecia yellowish to greenish . . . . . . . . L. varia
7. Apothecia light brown or darker

8. Thallus evanescent . . . . . . . . . . . . L. dispersa
8. Thallus persistent

9. Apothecia light brown, exciples persistent. . . . .
L. subfusca

9. Apothecia dark brown to blackish,
exciples often disappearing. . . . . . . L. melaena

3. Lecania
1. Apothecia pruinose . . . . . . . . . . . . . L. californica
1. Apothecia not pruinose . . . . . . . . . . . L. perproxima

Parmeliaceae
1. Thallus and apothecia yellow; spores many per ascus. . . .

1. Candelaria
1. Thallus and apothecia rarely yellow; spores eight per ascus

2. Parmelia
1. Candelaria

1. Exciple fibrillose below. . . . . . . . . . . . .C. fibrosa
1. Exciple not fibrillose

2. Thallus reduced to granulose squamules, or passing into
a powdery crust . . . . . . . . . . . . . . . C. concolor

var. effusa
2. Thallus granulose on margins only . . . . . . . C. concolor

var. concolor
2. Parmelia

1. Soredia present
2. Medulla yellowish . . . . . . . . . . . . . . P. obsessa
2. Medulla white

3. Soredia on upper surface . . . . . . . . . P. caperata
3. Soredia marginal . . . . . . . . . . . . P. reticulata



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1. Soredia absent
4. Isidia present

5. Thallus yellowish-green. . . . . . . . . . P. isidiata
5. Thallus bluish or gray

6. Upper surface with white dots . . . . . P. rudecta
6. Upper surface without white dots. . . P. haitiensis

4. Isidia absent
7. Thallus yellowish-green

8. Thallus tightly adnate . . . . . . . . P. conspersa
8. Thallus loosely attached . . . . . . P. stenophylla

7. Thallus bluish to gray. . . . . . . . . . . P. borreri

Caloplacaceae
The only genus representing this family is Caloplaca.
1. Thallus sorediate

2. Soredia dull orange to blackish. . . . . C. microphylina
2. Soredia yellow

3. Thallus smooth and continuous . . . C. chrysophthalma
3. Thallus distinctly aerolate . . . . . . . C. decipiens

1. Thallus esorediate
4. Thallus lobed at the margins

5. Exciple moderately thick and somewhat elevated. . . .
C. murorum

5. Exciple thin and not elevated. . . . . . . C. lobulata
4. Thallus not lobed at the margins

6. Apothecia yellow, exciples whitish . . . . C. ulmorum
6. Apothecia orange, exciple dull gray to yellow-green or

orange
7. Exciple distinctly yellowish-green. . C. aurantiaca
7. Exciple gray or orange

8. Exciple gray, thallus well developed, areolate. .
C. arizonica

8. Exciple orange, thallus scanty
9. Apothecia crowded, angular, spores slightly

curved. . . . . . . . . . . . C. flavovirescens
9. Apothecia scattered, circular, spores not

curved. . . . . . . . . . . . . . . . C. cerina

Teloschistaceae
1. Thallus fruticose. . . . . . . . . . . . . 1. Teloschistes
1. Thallus foliose. . . . . . . . . . . . . . . . 2. Xanthoria

1. Teloschistes
The only species representing this genus is T.chrysophthalmus.

2. Xanthoria
The only species representing this genus is X. candelaria.



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Buelliaceae
1. Apothecia with proper exciples . . . . . . . . . 1. Buellia
1. Apothecia with thalloid exciples . . . . . . . 2. Rinodina

1. Buellia
1. Spores three-septate . . . . . . . . . . . . . B. alboatra
1. Spores one-septate

2. Thallus of well-developed areoles
3. Areoles olive-brown. . . . . . . . . .B. novomexicana
3. Areoles greenish-gray to ashy

4. Hypothallus scanty or obsolete. . . B. retrovertens
4. Hypothallus black, prominent

5. Exciple white initially, then turning darker. . .
B. stigmaea

5. Exciple black. . . . . . . . . . . . . .B. spuria
2. Thallus not areolate

6. Hypothecium colorless. . . . . . . . . . . . B. vilis
6. Hypothecium brownish

7. Hymenium colorless throughout. . . . . B. schaereri
7. Hymenium brown above. . . . . . . . . . B. punctata

2. Rinodina
The only species representing this genus is R. oreina.

Physciaceae
1. Medulla yellow to salmon-colored. . . . . . . . . . .Pyxine
1. Medulla white

2. Lower cortex well developed. . . . . . . . . . . Physcia
2. Lower cortex poorly developed or wanting. . . Anaptychia

1. Pyxine
The only species representing this genus is P. caesiopruinosa.

2. Physcia
1. Thallus esorediate

2. Thallus tightly adnate throughout. . . . . P. syncolla
2. Thallus loosely adnate

3. Lobe tips thin and somewhat elevated . . P. aipolia
3. Lobe tips thickened and turned down

4. Whitish to gray, apothecia usually pruinose,
on trees. . . . . . . . . . . . . . P. stellaris

4. Dark gray to blackish, apothecia not pruinose,
on rocks. . . . . . . . . . . . . . . . P. halei

1. Thallus sorediate
5. Soredia in capitate patches on the upper surface

6. Thallus under two cm. across, brownish, appearing
crustose. . . . . . . . . . . . . . . . . . P. elaeina

6. Thallus up to five cm. across, grayish, definitely
foliose
7. Soredia white to pale blue . . . . . P. tribacoides
7. Soredia grayish-green. . . . . . . . P. orbicularis



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5. Soredia along margins, and sometimes on the upper
surface, but not in capitate patches
8. Thallus pruinose . . . . . . . . . . . . . . P. grisea
8. Thallus not pruinose

9. Lobes thin, flat, and often raised at the margins
P. millegrana

9. Lobes more or less rounded over the top with tips
touching the substrate . . . . . . . . P. subtilis

3. Anaptychia
1. Thallus sorediate

2. Hypothallus distinctly yellow. . . . . . A. heterochroa
2. Hypothallus gray to blackish . . . . . . . . A. speciosa

1. Thallus esorediate
3. Thallus isidiate . . . . . . . . . . . . A. granulifera
3. Thallus without isidia. . . . . . . . . . . A. hypoleuca

Lichenes Imperfecti
Leprariaceae

1. Thallus often zonate, grayish-green . . . . . . 1. Crocynia
1. Thallus never zonate, bright yellow . . . . . . 2. Lepraria

1. Crocynia
The only species representing this genus is C. membranacea.

2. Lepraria
The only species representing this genus is L. chlorine.