Keys for Lichens of North America
Third Edition
Irwin M. Brodo
Photographs by Sylvia Duran Sharnoff and Stephen Sharnoff
Drawings by Susan Laurie-Bourque
Draft reprint for private distribution only, with permission from Yale University Press,
New Haven, CT.
July 2006
1
Preface
Many users of Lichens of North America (LNA) have asked that the keys to groups,
genera, and species be reproduced as a separate publication bound in such a way as to make them
convenient to use in university courses and workshops. This would free the larger volume to be
used as more of a reference, somewhat removed from the ravages of splashed reagents and dusty
packets. The present compilation was therefore prepared to answer those requests. To facilitate
its use in a laboratory setting, the final version will have water-resistant covers, and the volume
will be spiral-bound.
The keys are reproduced with few changes or amendments, more or less as they appeared
in LNA, except they are grouped together in alphabetical order by genus. Some corrections and
nomenclatural updates, and a number of additional species, however, have been incorporated into
the text. The Third Edition includes a new key to the genus Biatora and similar lichens. (In this
draft version, these changes are in bold face in most cases.) Another addition is a key to lichen
photobionts, which is found following the keys to species. Comments on the efficacy of these
keys would be much appreciated by the author. Since the lichen keys were written to be used in
conjunction with the full descriptions, photographs, maps, and discussions found in the larger
work, they will seem short on detail to those attempting to use them alone.
All keys, even those stripped of unnecessary jargon, need a glossary to define technical
terms, and to explain how the author uses them. I have therefore included in this book the original
glossary found in LNA together with the figures and photographs that illustrate the terms.
Acknowledgements
I would like to thank Stephen Sharnoff for permission to use the photographs that
illustrate the glossary. I also thank Jean Thomson Black at Yale University Press for giving me
permission to try out this format on students during the summers of 2003 and 2004 to test its
usefulness as a separate publication. Michele LeBlanc assembled the figures and plates for these
editions, and I much appreciate her diligence and skill.
2
Table of Contents
Preface...............................................................................................................................................2
Acknowledgements ...........................................................................................................................2
Table of Contents ..............................................................................................................................3
Identification Keys to Genera and Major Groups .............................................................................5
Key to Keys.......................................................................................................................................5
Key A: Fruticose Lichens..................................................................................................................7
Key B: Dwarf fruticose Filamentous Lichens .................................................................................16
Key C: Sterile Crustose Lichens .....................................................................................................17
Key D: Crustose Perithecial lichens and Lichens with Ascomata Resembling Perithecia..............23
Key E: Crustose Script Lichens ......................................................................................................28
Key F: Crustose Disk Lichens.........................................................................................................30
Key G: Squamulose Lichens ...........................................................................................................49
Key H: Umbilicate and Fan-Shaped Lichens ..................................................................................56
Key I: Jelly Lichens.........................................................................................................................57
Key J: Yellow Foliose Lichens .......................................................................................................58
Key K: Foliose Lichens that Are Not Umbilicate, Jelly-Like or Yellow ........................................63
Identification Keys to Species (alphabetically by genus)................................................................78
Lichen Photobionts........................................................................................................................209
Glossary.........................................................................................................................................211
Figures...........................................................................................................................................224
Plates .............................................................................................................................................234
3
4
Identification Keys to Genera
KEY TO KEYS
1.
Fruiting body a mushroom or fleshy, ephemeral, club-shaped stalk with a mushroom-like
consistency..........................................................................................................................2
1.
Fruiting body a perennial ascoma (apothecium, perithecium, or lirella); or thallus without
any fruiting bodies ..............................................................................................................3
2.
Fruiting body a pale to dark yellow, gilled mushroom, growing out of a lichenized thallus
consisting of either dark green spherical granules, or green, lobed squamules .Omphalina
2.
Fruiting body club-shaped, usually unbranched, white to orange, growing out of a
gelatinous, dark green, barely lichenized algal film......................................... Multiclavula
3.
Thallus erect, shrubby, tufted, or pendent, at least in part (sometimes with a crust-like or
scaly basal thallus as well); branches or stalks round, angular, or flat in cross-section, but
if flattened, then both surfaces similar in color and texture ................................................4
3.
Thallus composed entirely of flat lobes, branches or scales, with clearly different upper
and lower surfaces, or crust-like and firmly attached to the substrate over the entire lower
surface, or growing within the substrate .............................................................................5
4.
Thallus relatively large, more than 5 mm long or high, with branches or stalks mostly
more than 0.2 mm in diameter (figs. 2h-j) .................................. Fruticose lichens (Key A)
4.
Thallus forming small tufts or clumps rarely exceeding 5 mm long or high, consisting of
fine, hair-like elements rarely more than 0.2 mm in diameter (see, for example,
Polychidium, Ephebe, or Coenogonium)
...................................................................... Dwarf fruticose filamentous lichens (Key B)
5.
Thallus consisting of a crust that grows with its entire lower surface in intimate contact
with the surface and cannot be removed from the substrate intact (i.e., without leaving
part of the lichen's lower surface behind); thallus can be smooth and continuous,
powdery, cracked into angular patches (areoles), extremely thin (even growing within the
substrate), or quite thick and warty, sometimes lobed at the margins (fig. 2a-d)6 (Crustose lichens)
5
5.
Thallus consisting of more or less flat dorsiventral lobes or scales, ascending or closely
appressed to the substrate, more or less easily removed from the substrate with the lower
surface intact (fig. 2e-g)......................................................................................................9
6.
Fruiting bodies absent ........................................................ Sterile crustose lichens (Key C)
6.
Fruiting bodies (ascomata) present .....................................................................................7
7.
Ascomata flask-shaped, perithecia or resembling perithecia (opening by a deeply
concave or pit-like ostiole) (figs. 17, 12h, 13d), superficial or buried in the thallus or in a
fertile wart (pseudostroma) with only the ostiole visible. [Note: Pycnidia can sometimes
resemble perithecia, but they contain hundreds of conidia not associated with asci; see
fig. 18.]………………………………………………….. Crustose perithecial lichens
(Key D)
7.
Ascomata apothecia: (1) disk- or cup-shaped (figs. 12a-c; 13a-c,f), sometimes raised on a
stubby or slender stalk (fig. 12f); (2) buried in thalline warts (as in Pertusaria subpertusa
or P. xanthodes) (fig. 12g); or (3) elongated and sometimes branched (lirellae)(figs. 12e;
13e) ....................................................................................................................................8
8.
Ascomata lirellae, short and ellipsoid to quite long, script-like, and sometimes branched ..
............................................................................................Crustose script lichens (Key E)
8.
Ascomata apothecia (disk- or cup-shaped or buried within thalline warts) ..........................
................................................................................ Crustose disk lichens (Key F)
9.(5)
Thallus composed of separate or overlapping scales (squamules) that are rarely more than
5 mm long or broad (fig. 2e)...................................................Squamulose lichens (Key G)
9.
Thallus composed of narrow or broad, flat lobes; thallus generally exceeding 5 mm long
or broad (fig. 2f-g) ................................................................................10 (Foliose lichens)
10.
Thallus attached to the substrate by a holdfast at a single point, either central, giving it
the shape of an umbrella (fig. 2g), or at one edge, giving it the shape of a fan. (Lobes
sometimes crowded and overlapping, obscuring the attachment.)........................................
...........................................................................Umbilicate or fan-shaped lichens (Key H)
10.
Thallus attached to the substrate at numerous points, either with hair-like attachment
6
organs (rhizines), a fuzzy tomentum, or directly by the lower surface (fig. 2f)................11
11.
Thallus, when wet, translucent and jelly-like, and typically black to very dark gray,
brown or olive (e.g., Collema); photobiont blue-green, usually distributed more or less
uniformly within the thallus, i.e., not confined to a definite layer (fig. 7)Jelly lichens (Key I)
11.
Thallus remaining opaque when wet, color various; photobiont green or blue-green,
confined to a definite layer within the thallus (fig. 4).......................................................12
12.
Dry thallus orange, yellow, greenish yellow, or yellowish green (see, for example,
Xanthoria, Candelaria, and Flavoparmelia) ………...........Yellow foliose lichens (Key
J)
12.
Dry thallus shades of white, gray, pale green, olive, brown, or black, without a yellowish
tint ....................................Non-umbilicate, non-jelly, non-yellow foliose lichens (Key K)
KEY A: FRUTICOSE LICHENS
1.
Thallus bright yellow, bright chartreuse, or orange ............................................................2
1.
Thallus pale greenish yellow, yellowish green, green, white, gray, brown, olive, or black
...........................................................................................................................................5
2.
Branches distinctly flattened; medulla yellow; cortex and medulla K–; on the ground in
arctic or alpine sites .................................................................................... Vulpicida tilesii
2.
Branches round to angular; medulla white; on trees, or, if on the ground, not arctic-alpine
............................................................................................................................................3
3.
Thallus bright yellow to chartreuse, K–; mostly in inland, montane localities....... Letharia
3.
Thallus orange, at least on exposed parts, K+ dark red-purple...........................................4
4.
Attached directly to rock along seashores in California .................... Caloplaca coralloides
4.
On trees, shrubs or soil .....................................................................................Teloschistes
5.(1)
Thallus pendent or almost pendent (i.e., growing downward or outward); mostly on trees,
shrubs, or vertical surfaces of rocks (includes hair and beard lichens)...............................6
7
5.
Thallus erect or prostrate (i.e., basically growing upward at least initially); mostly on the
8
ground or on horizontal surfaces of rocks (includes lichens with upright stalks or podetia)
..........................................................................................................................................28
6.
Thallus greenish yellow or yellowish green (containing usnic acid in the cortex) .............7
6.
Thallus shades of white, gray, brown, olive, or black (lacking usnic acid) ......................12
7.
Branches with a tough, single, central cord ................................................................Usnea
7.
Branches with a more or less uniform medulla, without a single, central
cord....................8
8.
Pseudocyphellae present .....................................................................................................9
8.
Pseudocyphellae absent ....................................................................................................10
9.
Branches flattened in cross section, at least at the base ....................................... Ramalina
9.
Branches round in cross section, except sometimes at the axils ............................Alectoria
10.
Thallus soft and pliable, with a thin cortex .............................................................. Evernia
10.
Thallus stiff because of a thick, tough cortex ...................................................................11
11.
Pycnidia black, abundant, appearing as black dots on the margins and sometimes the
surface of the thallus; restricted to coastal California ................................................ Niebla
11.
Pycnidia, when present, pale and inconspicuous; widespread.............................. Ramalina
12.(6)
Branches distinctly flattened, at least at the base or tips ...................................................13
12.
Branches round or angular in cross section throughout (sometimes grooved or twisted, or
flattened at the axils) .........................................................................................................20
13.
Thallus brown to greenish black .......................................................................Kaernefeltia
13.
Thallus pale gray to smoky brownish gray .......................................................................14
14.
Cilia present on the branches or apothecia........................................................................15
14.
Cilia absent........................................................................................................................17
9
15.
Branches 10-25 mm long; medulla C+ pink ...............................Everniastrum catawbiense
15.
Branches 40-150 mm long; medulla C–............................................................................16
16.
Cortex K+ yellow; medulla K+ red (atranorin and salazinic acid); sorediate on the lower
surface of the lobe tips .................................................................. Heterodermia leucomela
16.
Cortex and medulla K–; lobes without soredia ..................................... Anaptychia setifera
17.
Fruiting bodies almost spherical, black, powdery masses formed close to the tips of the
flattened branches; on mossy trees or rocks in Pacific Northwest coastal forests ...............
................................................................. [Bunodophoron melanocarpum; see couplet 34]
17.
Fruiting bodies round or elongate apothecia, never powdery; not found in Pacific
Northwest coastal forests ..................................................................................................18
18.
On trees or wood in the southwest or Appalachian-Great Lakes regions; cortex K+
yellow (atranorin); lower surface gray to black in part .................................. Pseudevernia
18.
On rocks and shrubs in coastal California; cortex K–; lower and upper surfaces the same
color ..................................................................................................................................19
19.
Cortex C+ red, PD– (lecanoric acid).......................................................................Roccella
19.
Cortex C–, PD+ red (protocetraric acid)........................................................Dendrographa
20.(12) Thallus almost white, sometimes darkening to dark yellowish gray; coastal rocks and
shrubs in California; cortex PD+ red .............................................................Dendrographa
20.
Thallus shades of gray or brown, or almost black; cortex PD– or PD+ yellow (except
rarely in some dark species of Bryoria); on bark or wood, rarely on rock........................21
21.
Thallus with thick main branches and much finer secondary and tertiary branches, graygreen, often browned at the tips; fruiting body a spherical mazaedium at the branch tips ...
...................................................................................................... Sphaerophorus globosus
21.
Thallus gradually tapering, mostly uniform in color; fruiting bodies
apothecia…………22
22.
Thallus shrubby, with divergent, stiff branches, usually with a spiny appearance ...........23
10
22.
Thallus mostly pendent with parallel branches .................................................................25
23.
Thallus dark greenish black, at least close to the branch tips; apothecia appearing
terminal on branches, almost black; epihymenium K+ violet.........Kaernefeltia californica
23.
Thallus and apothecia brown; epihymenium K– ..............................................................24
24.
Thallus distinctly reddish brown; cells of the cortex (viewed at 100-400X) interlocking
like a jigsaw puzzle; branches grooved and channeled throughout ......................................
........................................................................................................Nodobryoria abbreviata
24.
Thallus brown without a reddish tint; cells of the cortex elongate, parallel, not
interlocking; branches mostly uniformly terete except at the axils where they can be
flattened or pitted ..................................................................................................... Bryoria
25.
Strongly depressed pseudocyphellae forming long, deep, linear grooves down the
branches, sometimes twisting around the branch; thallus pale, dull, mottled with reddish
brown to gray-brown streaks; rare .................................................................Sulcaria badia
25.
Pseudocyphellae present or absent, linear or ellipsoid, but not forming deep grooves;
thallus usually more or less uniform in color....................................................................26
26.
Thallus gray to pale or dark brown or olive............................................................. Bryoria
26.
Thallus distinctly reddish brown.......................................................................................27
27.
Branches grooved and pitted, brittle; apothecia brown, without pruina; cortical cells in
surface view (viewed at 100-400X) resembling pieces of a jigsaw puzzle ..........................
............................................................................................................Nodobryoria oregana
27.
Branches more or less smooth except for the axils; apothecia, if present, with yellow
pruina; cortical cells straight, long, and parallel .......................................Bryoria fremontii
28.(5)
Stalks or branches solid ....................................................................................................29
28.
Stalks or branches hollow, at least in part.........................................................................63
29.
Stalks or branches distinctly flattened at least at the base or tips......................................30
29.
Stalks or branches mostly round or angular in cross-section (sometimes flattened at the
11
axils) .................................................................................................................................40
30.
Thallus yellow-green (containing usnic acid) ...................................................................31
30.
Thallus gray, brown, or black (lacking usnic acid) ..........................................................33
31.
Pseudocyphellae present, slightly depressed, on the lower surface of the branches; lobes
thin, entirely flattened and erect.......................................................................Flavocetraria
31.
Pseudocyphellae absent, although the branches sometimes have raised, white warts; lobes
relatively thick ..................................................................................................................32
32.
Lobes irregularly branched, strongly curled and twisted, usually with white warts on the
margins or tips of the branches; growing unattached over bare soils in the interior high
plateaus ................................................................................................ Rhizoplaca haydenii
32.
Lobes regularly dichotomous, not curled, generally round at least toward the base,
without white warts; on soil in arctic or alpine sites………...Allocetraria
madreporiformis
33.(30) Thallus gray or gray-green................................................................................................34
33.
Thallus very dark brown to black ....................................................................................35
34.
On limestone, in arid southcentral U.S; branches more or less uniform in width and very
regularly dichotomous. ................................................................. Speerschneidera euploca
34.
On mossy trees or rocks, in humid coastal forests of the Pacific Northwest; main
branches broad, giving rise to clusters of much narrower branches, irregularly
dichotomous..........................................................................................................................
................................................................. [Bunodophoron melanocarpum; see couplet 17]
35.
Directly on rock ................................................................................................................36
35.
On soil, heath, or mosses (sometimes over rock)..............................................................38
36.
Thallus abundantly branched, forming wool-like cushions….. Pseudephebe minuscula
36.
Thallus with relatively few strap-shaped branches; forming small clumps ......................37
12
37.
On limestone; photobiont blue-green; thallus jellylike when wet…....….Lichinella
nigritella
37.
On siliceous rock; photobiont green; thallus unchanged when wet......................................
.......................................................................................................Cornicularia normoerica
38.(33) Branches very flat, regularly dichotomous, with broad, pruinose patches (pseudocyphellae)
on lower surface; growing unattached (vagrant), rolling freely over the soil and heath
..................................................................................................................... Masonhalea richardsonii
38.
Branches usually grooved or inrolled at the margins; pseudocyphellae linear or irregular
in shape, not broad and pruinose; growing in more or less fixed clumps, not rolling freely
over the soil or heath.........................................................................................................39
39.
Medulla C+ pink (gyrophoric acid); lobe margins more or less even, without projections..
................................................................................................................. Cetrariella delisei
39.
Medulla C–; lobe margins with stalked pycnidia or cilia.........................................Cetraria
40.(29) Thallus yellowish green (containing usnic acid) ................................................................41
40.
Thallus white, gray, brown, olive, or black (lacking usnic acid) ......................................45
41.
Thallus prostrate, with long, slender branches, 0.5-3 mm in diameter, 30-50 mm long,
mostly irregularly branched and tangled................................................. Evernia divaricata
41.
Thallus at least partly erect; branches short, usually under 30 mm high ..........................42
42.
Branches very slender, less than 0.3 mm in diameter, covered with cottony granules
......................................................................................................................... Leprocaulon
42.
Branches thicker than 0.3 mm; surface relatively smooth at least at the
base…………...43
43.
In arctic or alpine sites in western mountains; not cushion-forming ....................................
...............................................................................................Allocetraria madreporiformis
43.
Not arctic-alpine; forming rounded cushions or clumps...................................................44
44.
On seashore rocks in California; branch tips becoming granular or sorediate, lacking
13
white warts; cortex C+ orange, KC+ red-orange (xanthones)….Lecanora phryganitis
44.
Rolling unattached over bare soils in the interior high plateaus; branches with white
warts but without granules or soredia; cortex C–, KC+ yellow (usnic acid alone)...............
............................................................................................................. Rhizoplaca haydenii
45.(40) Branching frequently ........................................................................................................46
45.
Unbranched, or branching at most once or twice..............................................................59
46.
Thallus black or very dark brown to almost black............................................................47
46.
Thallus reddish brown, yellowish brown, olive, gray, or white........................................49
47.
Attached directly to rock..................................................................................Pseudephebe
47.
On soil or heath.................................................................................................................48
48.
Pseudocyphellae pale, raised, usually conspicuous; thallus pale at the base; cortex and
usually medulla C+ pink, PD+ yellow ................................................... Alectoria nigricans
48.
Pseudocyphellae dark, level with surface or depressed, inconspicuous; thallus dark
throughout; cortex C–, PD–; medulla PD+ red……………………………Bryoria nitidula
49.(46) Thallus shades of brown or olive ......................................................................................50
49.
Thallus white to dark greenish gray..................................................................................53
50.
Photobiont blue-green........................................................... Dendriscocaulon intricatulum
50.
Photobiont green ...............................................................................................................51
51.
Pseudocyphellae absent ................................................................................Sphaerophorus
51.
Pseudocyphellae white, conspicuous ................................................................................52
52.
Pseudocyphellae usually raised like small bumps, C+ pink; medulla usually C+ pink as
well (olivetoric acid) ......................................................................... Bryocaulon divergens
52.
Pseudocyphellae depressed, C–; medulla C–.............................................Cetraria aculeata
53.(49) Photobiont blue-green........................................................... Dendriscocaulon intricatulum
14
53.
Photobiont green ...............................................................................................................54
54.
Branches very slender, less than 0.3 mm in diameter, covered with cottony granules
......................................................................................................................... Leprocaulon
54.
Branches thicker than 0.3 mm; surface relatively smooth at least at the base…………..55
55.
Thallus growing as prostrate cushions up to 30 mm in diameter on soil in the arid interior
of North America; round, depressed pseudocyphellae conspicuous on branches
...................................................................................................................Aspicilia hispida
55.
Thallus colonies usually larger than 30 mm, not on soil in the arid interior of North
America; pseudocyphellae, if present, not round and depressed ......................................56
56.
Cortex C+ red; on coastal rocks in southern California; fruiting bodies large, lecanorine,
with heavily pruinose disks.............................................................. Schizopelte californica
56.
Cortex C–; fruiting bodies not pruinose............................................................................57
57.
Stalks with a tough, cartilaginous central core, more or less covered with granular,
cylindrical, or scaly outgrowths (phyllocladia) (fig. 8g)..................................Stereocaulon
57.
Stalks filled with white medulla, without a cartilaginous central core; without
phyllocladia (fig. 8e).........................................................................................................58
58.
Branch tips blackened; pseudocyphellae usually conspicuous................Alectoria
nigricans
58.
Branch tips browned, not blackened; pseudocyphellae absent .....................Sphaerophorus
59.(45) On conifer branches in subalpine sites; fruiting body a mazaedium at the tip of a stubby,
barrel-shaped stalk .................................................................................Tholurna dissimilis
59.
On soil, rock, or moss; fruiting body not a mazaedium ....................................................60
60.
Stalks white, smooth or with longitudinal furrows, not growing from a primary thallus,
without fruiting bodies of any kind; cortex C+ violet (fading quickly) ................................
................................................................................................................... Siphula ceratites
60.
Stalks almost white to gray or olive-gray, without furrows, developing from a crustose or
granular primary thallus; cortex C– ..................................................................................61
15
61.
Cephalodia absent ................................................... Baeomyces key (Baeomyces, Dibaeis)
61.
Lumpy pink to dark gray cephalodia present on primary thallus, sometimes on stalks....62
62.
Apothecia usually present, black, cylindrical or almost spherical; stalks without soredia ...
.............................................................................................................. Pilophorus
62.
Apothecia extremely rare, brown; granular soredia at the tips of the stalks .........................
..........................................................................................................Stereocaulon pileatum
63.(28) Photobiont blue-green; thallus olive-brown; stalks cylindrical, mostly less than 2 mm
high; on rock .............................................................................................Peltula cylindrica
63.
Photobiont green; thallus not olive-brown; stalks taller than 2 mm .................................64
64.
Lacking a basal thallus......................................................................................................65
64.
With a scaly or crustose basal thallus ...............................................................................70
65.
Stalks abundantly branched ..............................................................................................66
65.
Stalks unbranched or branched only once or twice...........................................................67
66.
Thallus less than 20 mm tall; branches relatively stocky, usually violet or pinkish due to
pruina; interior cavity containing webby hyphae…………………….Dactylina ramulosa
66.
Thallus over 20 mm tall, without pruina; interior cavity empty, lined with a cartilaginous
stereome (fig.8f).............................................................. Cladonia key (Cladonia, Cladina)
67.
Thallus C+ red (erythrin); stalks inflated, brittle; on coastal rocks in southern California...
................................................................................................................... Hubbsia parishii
67.
Thallus C–, or, if C+ pink, then arctic-alpine ...................................................................68
68.
Stalks white, slender, 1-2.5 mm in diameter, usually partly or entirely prostrate.................
....................................................................................................... Thamnolia vermicularis
68.
Stalks yellowish, brown or greenish gray, erect ...............................................................69
69.
Stalks inflated, thin-walled and brittle, with no stereome; entirely without squamules
16
...................................................................................................................Dactylina arctica
69.
Stalks more or less slender, not inflated, stiff, supported by a cartilaginous stereome,
sometimes bearing squamules................................................................................ Cladonia
70.(65) Primary thallus crustose, granular, not at all squamulose…………...Pycnothelia
papillaria
70.
Primary thallus squamulose ................................................................................... Cladonia
KEY B: DWARF FRUTICOSE FILAMENTOUS LICHENS
1.
Thallus pale: gray, greenish, olive, or orange .....................................................................2
1.
Thallus dark: black, olive, or brown ..................................................................................6
2.
Thallus forming cottony tufts of fine, branching filaments or hairs; photobiont green or
blue-green ...........................................................................................................................3
2.
Thallus with more or less erect, fine stalks; photobiont green............................................4
3.
On subtropical and tropical trees and shrubs; tufts cottony, sometimes flattened, pale
green to orange, almost white when old; disk-shaped apothecia common, yellow to
orange; photobiont green (Trentepohlia) ...................................................... Coenogonium
3.
On twigs and branches on the very humid west coast; thallus forming small, foam-like
clumps, pale olive or olive-gray; photobiont blue-green (Hyphomorpha)............................
........................................................................................................Polychidium contortum
4.
Stalks branched, covered with cottony granules giving clumps the appearance of a pale
gray to whitish, leprose, crustose lichen ...................................... Leprocaulon gracilescens
4.
Stalks branched or unbranched, without granules or soredia; thallus thin and
membranous........................................................................................................................5
5.
Stalks like long, slender isidia, often branched, containing green algae; apothecia
sometimes produced on basal crust; on bark in the Pacific Northwest .................................
.....................................................................................................Loxosporopsis corallifera
5.
Stalks colorless, without algae, unbranched, tipped by starshaped, umbrella-like conidia-
17
containing structures; on mosses and tree bases in the southeast
........................................................................................................Gomphillus americanus
6.(1)
Thallus dark red-brown, with abundant, large, red-brown apothecia; growing in patches
of moss over rock; photobiont blue-green (Nostoc)………………….Polychidium
muscicola
6.
Thallus black to shades of olive or olive-gray; apothecia inconspicuous, rare, or absent;
photobiont blue-green or green
........................................................................7
7.
On bark or moss; thallus olive to olive-gray or brownish gray; photobiont blue-green .....8
7.
On rock; thallus mostly black to very dark olive, at least when dry ...................................9
8.
On moss or mossy trunks; branches thick, 0.1-0.6 mm in diameter, spiny, dark olive-gray
to brownish; photobiont Nostoc............................................ Dendriscocaulon intricatulum
8.
On twigs and branches in humid localities on the west coast; branches hair-like, less than
0.1 mm in diameter, not spiny; thallus forming small, foam-like clumps, pale olive or
olive-gray; photobiont Scytonema……………………………..Polychidium contortum
9.
Thallus like black wool, the hairlike filaments 10-20 μm thick containing the green alga
Trentepohlia; typically on shaded rock faces....................................................................10
9.
Thallus forming tufts or clumps of much thicker filaments (40-140 μm) containing
cyanobacteria (Stigonema); on dry or wet rocks, usually
exposed………………………11
10.
Cells enveloping algal filaments long and straight (Fig. 34) ............................... Racodium
10.
Cells enveloping algal filaments irregular and knobby.................................. [Cystocoleus]
11.
Thallus forming rounded clumps or cushions, 5-15 mm across and 6 mm high, on dry or
wet rock; blue-green, rhizine-like hyphae developing at the base of the cushions ...............
............................................................................................................. Spilonema revertens
11.
Thallus forming irregular patches or felt-like mats (when abundant), not rounded
cushions; usually on wet rocks near a stream or lake shore; lacking any blue-green,
rhizoid-like hyphae at the base of the clumps................................................ Ephebe lanata
18
KEY C: STERILE CRUSTOSE LICHENS
1.
Soredia and isidia absent.....................................................................................................2
1.
Soredia or isidia present....................................................................................................10
2.
On bark; older parts of thallus covered with hollow pustules that break up into granular
or soredia-like schizidia ....................................................................... Loxospora pustulata
2.
On wood, soil, mosses, dead vegetation, or rock................................................................3
3.
On rock; thallus cortex C+ red (lecanoric acid) .................................... Dirina catalinariae.
3.
On wood, soil, mosses, or dead vegetation. ........................................................................4
4.
Thallus yellow to orange, or greenish yellow to yellow-green ...........................................5
4.
Thallus without yellow or orange tint .................................................................................6
5.
K– ..........................................................................................................................Lecanora
5.
K+ purple; on soil ................................................................................................. Fulgensia
6.
Thallus pale.........................................................................................................................7
6.
Thallus dark ........................................................................................................................9
7.
Thallus cortex and medulla PD+ red (fumarprotocetraric acid); thallus almost white, with
tall verrucae; arctic-alpine..................................................................... Pertusaria dactylina
7.
Thallus cortex and medulla PD– or PD+ yellow (fumarprotocetraric acid absent); arcticalpine or temperate..............................................................................................................8
8.
Thallus C+ pink or red, PD– (gyrophoric acid; baeomycesic acid absent)
........................................................................................................ Trapeliopsis granulosa
8.
Thallus C–, PD+ yellow (baeomycesic acid; gyrophoric acid absent)
...............................................................................................................Dibaeis baeomyces
9.(4)
Thallus dark steel gray to dark greenish gray;
19
verrucose…………….Trapeliopsis flexuosa
9.
Thallus dark brown, granular...................................................... Placynthiella oligotropha
10.(1)
Isidia present.....................................................................................................................11
10.
Isidia absent ......................................................................................................................15
11.
Thallus olive, distinctly lobed; photobiont blue-green; directly on rock in and near
southeastern Alaska ...................................................................... Vestergrenopsis isidiata
11.
Thallus white to gray-green, or reddish brown, not lobed at the edge; photobiont green;
on bark, wood, mosses, or dead vegetation, temperate or tropical. .................................12
12.
On exposed wood, thallus reddish brown; boreal to north Temperate
.........................................................................................................[Placynthiella icmalea]
12.
On bark, rock, mosses, or dead vegetation; thallus pale, white to gray-green .................13
13.
Thick red or white prothallus present; isidia subspherical (granular) to cylindrical; coastal
plain of southeastern U.S. .............................................................................. Cryptothecia
13.
Prothallus present or absent, never red; isidia cylindrical; arctic-alpine, or temperate.....14
14.
On mosses or dead vegetation in arctic-alpine habitats; thallus white, verrucose; isidia
thick and crowded; medulla PD+ red................................................. Pertusaria dactylina
14.
On bark or rock; ; thallus yellowish gray to greenish gray, medulla PD- ………….
14a
14a.
On bark in the coastal Pacific Northwest; thallus more or less smooth; isidia long
and slender, often branched; medulla C……………..……Loxosporopsis corallifera
14a.
On bark or rock, Appalachians north to New England; thallus smooth to verrucose;
isidia short and stout; medulla C+ red ………………..……….. Ochrolechia yasudae
15.(10) Soredia comprising entire thallus .....................................................................................16
15.
Soredia in delimited soralia, or diffuse on older parts of the thallus. ...............................23
20
16.
Thallus pale green, gray, brown, or black, without yellow or orange tint ........................17
16.
Thallus yellowish green, greenish yellow, bright yellow, or orange ................................19
17.
Thallus forming round, shingled lobes with a basal cottony brown to white hypothallus;
thallus KC–, PD+ orange (pannaric and roccellic acids; usnic acid absent)
..................................................................................................Leproloma membranaceum
17.
Thallus thin, never forming shingled lobes; hypothallus absent, prothallus sometimes
present; pannaric acid absent ............................................................................................18
18.
Prothallus white, forming a fibrous fringe; thallus KC+ gold, K–, PD– (usnic acid and
zeorin present)……………………………………………………..Lecanora thysanophora
18.
Fibrous white prothallus absent; thallus KC–, K+ yellow or K–, PD+ yellow or red (usnic
acid absent) ............................................................................................................. Lepraria
19(16). Thallus greenish-yellow or yellowish green .....................................................................20
19.
Thallus yolk-yellow, lemon yellow, yellow-orange, or orange-yellow ............................21
20.
Prothallus absent, thallus thick, edge indefinite or weakly lobed; thallus PD+ orange,
KC– (stictic acid present, usnic acid absent) ......................................... Lepraria lobificans
20.
Prothallus present, white and fibrous; thallus thin, edge definite; thallus PD–, KC+ gold
(stictic acid absent, usnic acid
present)…………………………….Lecanora thysanophora
21.
Soredia K+ purple (anthraquinones present, calycin absent).................... Caloplaca citrina
21.
Soredia K– (anthraquinones absent, calycin present) .......................................................22
22.
Thallus uniformly leprose, the soredia not clustered or associated with granules; in
shaded, humid habitats .................................................................... Chrysothrix candelaris
22.
Thallus usually producing soredia in small clumps, originating from the breakup of tiny
areoles, but the soredia can become confluent in well-developed specimens; very
common on exposed deciduous trees even close to urban areas…………
Candelariella efflorescens
21
23.(15) Thallus distinctly bright yellow or orange, K+ deep purple or K–...................................24
23.
Thallus not distinctly yellow or bright orange, K– or K+ yellow to red
..........................................................................................................................................27
24.
On soil...............................................................................................................................25
24.
On bark, wood, or rock .....................................................................................................26
25.
Thallus K+ deep purple......................................................................................... Fulgensia
25.
Thallus K–...................................................................................................... Arthrorhaphis
26.
Thallus margin distinctly lobed, with a lower cortex, producing pustules and granular
schizidia
26.
.............................................. Xanthoria sorediata (see also Caloplaca)
Thallus not lobed, entirely crustose, lacking pustules or schizidia
.................................................................................................couplet 21 of Caloplaca key
27.(23) Cephalodia present. ..........................................................................................................28
27.
Cephalodia absent. ............................................................................................................29
28.
Thallus continuous, flat, with a distinctly lobed margin; cephalodia flat, disk-shaped, on
the thallus surface ...................................................................................... Placopsis lambii
28.
Thallus verrucose or dispersed areolate, not lobed at the margin; cephalodia convex,
cushion-shaped, between the areoles ................................................Amygdalaria panaeola
29.
On rock .............................................................................................................................30
29.
On bark, wood, or soil ......................................................................................................31
30.
Thallus thin, shiny, C–; soralia KC+ purple; common and widespread ...............................
................................................................................................................... Pertusaria amara
30.
Thallus thick, dull, C+ pink or red; soralia KC– or KC+ pink or red; restricted to coastal
rocks in California................................................................ Dirina catalinariae f. sorediata
31.
On trees in tropical and subtropical regions; prothallus thick and
conspicuous…………32
22
31.
In temperate to boreal regions; prothallus absent or thin and
inconspicuous……………33
32.
Prothallus red or white; thallus not lobed, more or less smooth, not cottony or webby
......................................................................................................................... Cryptothecia
32.
Prothallus black; thallus clearly lobed, cottony or
webby…………….Crocynia pyxinoides
33.
Soredia diffuse, not confined to discrete soralia even in young parts of the thallus, or
arising as schizidia from the breakup of small hollow pustules........................................34
33.
Soredia at first produced in discrete soralia, or from the disintegration of solid granules
or verrucae, sometimes becoming confluent in older parts of the thallus; pustules and
schizidia absent .................................................................................................................35
34.
Thallus PD+ yellow, K+ yellow becoming blood red (norstictic acid) ................................
..................................................................................................................... Phlyctis argena
34.
Thallus PD+ orange, K+ bright yellow (thamnolic acid)................................................34a
34a.
Soredia arising as schizidia from the breakup of small hollow pustules ....
........................................................................................................Loxospora pustulata
34a.
Soredia farinose to granular, not arising as schizidia from the breakup of small
hollow pustules ................................................................................[Loxospora elatina]
35.
Thallus cortex or soredia C+ pink or red, KC+ red (gyrophoric acid)..............................36
35.
Thallus cortex C–, KC– (gyrophoric acid absent); soredia KC– or KC+ purple ..............37
36.
Soredia white or yellowish; thallus continuous, verruculose or more or less smooth ..........
........................................................................................................ Ochrolechia androgyna
36.
Soredia pale greenish to dark green; thallus verrucose, areolate, or granular
..........................................................................................................................Trapeliopsis
37.
Thallus greenish or olive to brownish gray to dark greenish gray; soredia green or
yellow-green .....................................................................................................................38
23
37.
Thallus pale to dark slate gray or yellowish white; soredia white, pale to dark gray, or
brownish............................................................................................................................40
38.
Thallus distinctly lobed, appearing crustose but lower cortex present
.....................................................................................................Hyperphyscia adglutinata
38.
Thallus not at all lobed; truly crustose ..............................................................................39
39.
Soralia PD+ red (fumarprotocetraric acid); often with brown prothallus; common, boreal
to North Temperate .............................................................................[Fuscidea arboricola]
39.
Soralia PD– (contains perlatolic and hyperlatolic acids); with or without a brown or black
prothallus; rare, coastal British
Columbia……………………………[Ropalospora viridis]
40.
Soralia KC+ purple; thallus cortex PD– .................................................... Pertusaria amara
40.
Soralia medulla KC–; thallus cortex PD+ yellow, orange or red, or PD–. .......................41
41.
Thallus PD–, K–; thallus with conspicuous blue-black or dark gray prothallus. Soredia
usually grayish blue at least in part; all grayish blue tissue turning reddish purple in nitric
acid; common in Pacific Northwest on
alders………………………[Mycoblastus caesius]
41.
Thallus PD+ yellow or red; prothallus absent or pale.......................................................42
42.
Soredia olive-black, brown, or dark greenish gray, at least in part, PD+ yellow, K+ red
(norstictic acid); on bark and wood on the west and east
coasts…….[Buellia griseovirens]
42.
Soredia white, PD+ red or orange, K– or K+ yellow........................................................43
43.
Thallus very thin, whitish; soralia PD+ red, K– or brownish (fumarprotocetraric acid);
boreal forest, mostly on conifers and birch…………………….. Pyrrhospora cinnabarina
43.
Thallus thin or more frequently thick and verruculose; soralia (actually, sorediate fruiting
warts) PD+ orange, K+ deep yellow (thamnolic acid); temperate eastern regions, mainly
on deciduous trees………………………………………………..Pertusaria trachythallina
24
KEY D: CRUSTOSE PERITHECIAL LICHENS AND LICHENS WITH ASCOMATA RESEMBLING
PERITHECIA
1.
Spores 1-celled, ellipsoid and large (mostly 45-150 x 25-60 μm), most commonly with
thick walls; ascomata buried in thalline warts ....................................................................2
1.
Spores 1- celled, ellipsoid and small (less than 45 μm long), with thin walls; or 2- to
many-celled with either thin or thick walls; ascomata superficial or more or less buried in
warts, pseudostromata, or the thallus ..................................................................................4
2.
Spore walls conspicuously thick-walled, often layered (fig. 15-l); excipulum poorly
defined and always colorless; widespread ............................................................Pertusaria
2.
Spore walls relatively thin, never layered; excipulum colorless but distinct and welldeveloped; Pacific Northwest .............................................................................................3
3.
Ostiole deep and hole-like; hymenium and asci IKI– or orange; thallus cortex and
medulla K+ yellow, PD+ orange, UV– (stictic acid); on coastal maritime rocks; locally
common
....................................................................................................... Coccotrema maratimum
3.
Ostiole slightly depressed, not forming a hole; hymenium and asci IKI+ blue; thallus area
around ostiole C+ pink (gyrophoric acid), medulla UV+ white (alectoronic acid); on
maritime or alpine rocks near coast; rare………………………...[Ochrolechia
subplicans]
4.
Growing on rock, soil, mosses, dead vegetation, barnacle shells, lichens, or leaves..........5
4.
Growing on bark or wood.................................................................................................17
5.
Spores numerous in each ascus; inconspicuous, rather rare, temperate to boreal lichens...6
5.
Spores 8 or fewer per ascus ................................................................................................7
6.
Ascomata red-brown to black perithecia, without pruina; spores ellipsoid to fusiform, 4celled; on mosses and plant remains; asci cylindrical or club-shaped ............... [Thelopsis]
6.
Yellow pruina present on the thalline covering of the perithecium-like ascomata; spores
ellipsoid, one-celled; on soil or decaying lichens; asci pear-shaped (fig. 14o).....................
25
.......................................................................................................................[Thelocarpon]
7.
Spores 1-celled....................................................................................................................8
7.
Spores 2- or more-celled...................................................................................................10
8.
Photobiont blue-green; thallus reddish and gelatinous when wet; on rock
........................................................................................................... Pyrenopsis polycocca
8.
Photobiont green; thallus not reddish and gelatinous when wet .........................................9
9.
Paraphyses persistent; on soil. Thallus thin, continuous, greenish or greenish-brown;
widespread but overlooked ............................................................ [Thrombium epigaeum]
9.
Paraphyses absent, disappearing or not developing; on rock.............................. Verrucaria
10.(7)
Spores muriform, many-celled ..........................................................................................11
10.
Spores transversely septate; rarely submuriform ..............................................................13
11.
Ostioles deeply depressed (hole-like), sometimes encircled with radiating ridges;
paraphyses persistent; thallus thick, gray; medulla C+ red…………………..Diploschistes
11.
Ostioles level with perithecial surface, slightly depressed, or prominent, without radiating
ridges; paraphyses disintegrating or not developing; thallus thin or thick, pale gray to
dark brown; medulla C–....................................................................................................12
12.
Algae present in perithecial cavity .................................................................... Staurothele
12.
Algae absent from perithecial cavity............................................................... [Polyblastia]
13.(10) On mosses, dead vegetation, or leaves; spores 2- to 8-celled, rarely submuriform. Strigula
13.
On rock or barnacle shells.................................................................................................14
14.
Spores 4- to many-celled; paraphyses persistent ....................................................... Porina
14.
Spores 2-celled (rarely 4-celled); paraphyses persistent or disappearing .........................15
15.
On intertidal calcareous rocks or barnacle shells; thallus light brown to pale orange, often
only a membranous stain; photobiont blue-green; spores constricted at the septa; on
26
Atlantic and Pacific coasts…………………………………….[Pyrenocollema halodytes]
15.
On non-maritime, mostly calcareous, rocks; thallus pale gray or yellowish white;
photobiont green; spores not constricted at the septa; North Temperate to arctic-alpine .16
16.
Paraphyses disintegrating or not developing; excipulum carbonized or darkly pigmented;
periphyses conspicuous; spores narrow, length to width ratio mostly 2-3:1; mostly arcticalpine.................................................................................................................... Thelidium
16.
Paraphyses persistent; excipulum pale or colorless; periphyses absent; spores broadly
ellipsoid, length to width ratio mostly 2:1 or less; northeastern U.S. and Canada
..........................................................................................................[Acrocordia conoidea]
17.(2)
Spores very long and narrow, many-celled, not tapering, constricted at the septa giving
the spores a segmented, worm-like appearance ……………………. ……………
…………………Stictis urceolatum (Ach.) Gilenstam (syn. Conotrema urceolatum)
17.
Spores broadly to narrowly ellipsoid, fusiform, or needle-shaped, often tapering, never
worm-like..........................................................................................................................18
18.
Ostiole deep, forming a pit or hole; exciple pale or carbonized, well developed,
surrounded by a thalline envelope or covering giving the ascoma a double-walled
appearance, especially when the two walls are not fused ................................................19
18.
Ostiole not deep or forming a hole; exciple or excipulum dark or pale, surrounded by
thalline tissue or not ..........................................................................................................21
19
Short, hair-like hyphae resembling periphyses lining the inner face of exciple around the
ostiole; ascomata lacking a columella; northeastern and western coasts as well as
southeast............................................................................................................. Thelotrema
19.
Hair-like hyphae resembling periphyses absent; many species with a column of sterile
tissue (columella) developing in the ascomatal cavity; southeastern coastal plain...........20
20.
Exciple (inner wall of ascoma), and columella when present,
black……………Ocellularia
20.
Exciple, and columella when present, pale to reddish brown ............................Myriotrema
27
21.(18) Spores pale to dark brown when mature ..........................................................................22
21.
Spores colorless. ...............................................................................................................23
22.
Spores with unevenly thickened walls, central spore locules about equal in size, lensshaped; spores not constricted at the septa............................................................. Pyrenula
22.
Spores with uniformly thickened walls, spore locules not equal in size; spores constricted
at the septa. Thallus pale gray, endophloeodal; perithecia partly immersed, wall dark
above and pale below; spores fusiform, with pointed ends, 18-24 x 5-9 μm........................
.......................................................................................................[Eopyrenula intermedia]
23.
Spores 1-celled..................................................................................................................24
23.
Spores 2- or more-celled...................................................................................................25
24.
Asci containing numerous spores; thallus consisting of yellow pruinose warts containing
perithecium-like ascomata; on wood; spores smooth; paraphyses branched or
unbranched; northern temperate.....................................................................[Thelocarpon]
24.
Asci with 8 spores; thallus not at all yellow; perithecia black, not in thalline warts; on
bark; spores conspicuously warty on the surface; paraphyses branched; southeastern
coastal plain .................................................................................................. [Monoblastia]
25.
Asci containing numerous spores .............................................. [Thelopsis (see couplet 6)]
25.
Asci mostly 8-spored ........................................................................................................26
26.
Perithecia buried in a wartlike pseudostroma, often several perithecia per wart
............................................................................................................... Trypethelium key
26.
Perithecia discrete, not in a pseudostroma ........................................................................27
27.
Spores with unevenly thickened walls ..................................................... Trypethelium key
27.
Spores with uniformly thickened walls...........................................................................27a
27a.
Spores muriform; thallus endophloeodal ........................................................................
..........................................................[Julella fallaciosa (Stizenb. ex Arn.) R. C. Harris]
28
27a.
Spores transversely septate ...........................................................................................28
28.
Spores 6- or more-celled...................................................................................................29
28.
Spores 2- to 4-celled .........................................................................................................33
29.
Spores 18-27 x 4-7 μm, (5-)6-8(-9)-celled, constricted at the septa; perithecia black or
almost black; thallus whitish, within substrate, indistinct; southern coastal plain
........................................................................................ [Polymeridium quinqueseptatum]
29.
Spores 24-125 μm long, 8- to 14-celled, usually not, or only slightly constricted at the
septa; perithecia pale to dark brown or black; thallus yellowish gray to greenish gray, or
pale greenish, superficial or within
bark…………………………………………………30
30.
Thallus and sometimes perithecia with sparse to dense isidia. Spores 8-celled, 35-47(-57)
x 5.5-8 μm, tapered, not constricted at septa; southeastern coastal plain.. [Porina scabrida]
30.
Thallus and perithecia smooth, not isidiate.......................................................................31
31.
Spores 8-celled, fusiform (widest at or close to the center), often slightly constricted at
the septa, 24-42 x 5-7.5 μm; ascus thickened at tip but IKI–, with distinct ocular
chamber; northeastern to Great Lakes region…………………………………...[Strigula
stigmatella]
31.
Spores 8-14-celled, wider at one end than the other, not constricted at septa; ascus not
thickened at tip, IKI–, with no ocular chamber.................................................................32
32.
Spores 60-125 x 9-15 μm; perithecia covered with thalline tissue; inner wall pale brown
to yellowish brown; southeastern U.S.....................................................Porina heterospora
32.
Spores (32-)38-50 x 5.5-7.5 μm; perithecia exposed, brown to black, wall dark brown to
black; widespread, East Temperate..............................................[Trichothelium cestrense]
33.(28) Paraphyses abundantly branched and anastomosing; asci club-shaped; spores narrowly
ellipsoid to fusiform, length to width ratio mostly over 3:1, 2- to 4-celled, 14-20 x 4.5-6
μm; East Temperate……………………[Anisomeridium polypori (syn. A.
nyssaegenum)]
29
Note: Species of Arthopyrenia will also key here.]
33.
Paraphyses unbranched or slightly branched; asci narrowly cylindrical; spores ellipsoid,
length to width ratio mostly under 3:1 .............................................................................34
34.
Spores 4-celled, 18-27(-30) x 7-10(-12) μm; perithecia flask-shaped, with distinct,
usually off center to lateral neck; ostioles prominent; involucrellum brown; hymenium
IKI+ greenish-blue, paraphyses unbranched; northeastern…………[Lithothelium
hyalosporum]
34.
Spores 2-celled, 11-16.5 x 6-9.5 μm; perithecia hemispherical, ostioles level with
perithecial surface; involucrellum carbonized; perithecial cavity IKI–; paraphyses
slightly branched; northcentral U.S……………………………………………
[Acrocordia cavata]
KEY E: CRUSTOSE SCRIPT LICHENS
1.
On rock ...............................................................................................................................2
1.
On bark or wood .................................................................................................................4
2.
Spores 1-celled; thallus white, K+ red (norstictic acid); western coastal mountains and
Gaspé Peninsula (Quebec); very rare...............................................[Lithographa tesserata]
2.
Spores septate .....................................................................................................................3
3.
Hymenial surface ("disks" of ascomata) pruinose; exciple contains crystals
....................................................................................................................... Lecanographa
3.
Hymenial surface not pruinose; exciple lacks crystals........................................ Opegrapha
4.
Lirellae immersed in wart-like pseudostroma.....................................................................5
4.
Lirellae developing directly on the thallus..........................................................................6
5.
Lirellae black, opening with only a narrow fissure, pseudostroma heavily pruinose;
spores narrowly ellipsoid, 4- to 6-celled when mature .....................................Sarcographa
5.
Lirellae brown, flat, with a completely exposed hymenium; pseudostroma without pruina
or lightly pruinose; spores fusiform, 6- to 8 celled……………………..Glyphis
30
cicatricosa
6.
Spores one-celled ................................................................................................................7
6.
Spores 2- or more celled .....................................................................................................8
7.
Lirella wall brown, not carbonized; common and widespread ..........................Xylographa
7.
Lirella wall black, carbonized; rare, in Pacific Northwest
........................................................................................... [Ptychographa xylographoides]
8.
Spores muriform .................................................................................................................9
8.
Spores only transversely septate .......................................................................................11
9.
Spores brown to olive ...................................................................................Phaeographina
9.
Spores colorless ................................................................................................................10
10.
Lirellae with well-developed black wall (although sometimes immersed); asci cylindrical
to club-shaped ........................................................................................................Graphina
10.
Lirellae lacking any wall; asci balloon-shaped……………Arthothelium (key to
Arthonia)
11.
Spore walls evenly thickened, cells cylindrical (appearing square)..................................12
11.
Spore walls unevenly thickened, cells lens-shaped...........................................................14
12.
Ascomata appearing lecanorine (surrounded with thalline tissue); usually pruinose ...........
.................................................................................................................. [Schismatomma]
12.
Ascomata not at all lecanorine in appearance; pruinose or not.........................................13
13.
Lirellae with persistent carbonized walls (exciple)............................................. Opegrapha
13.
Lirellae without distinct walls (exciple absent) ..................................................... Arthonia
14.
Spores pale to dark brown when mature......................................................... Phaeographis
14.
Spores colorless .......................................................................................................Graphis
31
KEY F: CRUSTOSE DISK LICHENS
1.
Ascomata or pycnidia at the summit of a slender or stout stalk..........................................2
1.
Ascomata immersed to superficial, broadly attached or sometimes constricted at the base,
but not raised on a conspicuous stalk..................................................................................5
2.
Stalks stout; ascomata more than 0.5 mm in diameter, with flat to convex or
hemispherical disks; spores remain within asci at maturity ................................................3
2.
Stalks slender, hair-or stubble-like (fig. 12f); ascomata or pycnidia mostly under 0.4 mm
in diameter, irregular in shape or almost spherical; spores massed at the summit, loose or
remaining within asci at maturity .......................................................................................4
3.
Apothecia broader than stalk; disks pink to brown...............................................................
................................................................................ Baeomyces key (Baeomyces, Dibaeis)
3.
Apothecia immersed in the tip of the stalk; disks black........................ Pertusaria dactylina
4.
On moss; spores (conidia) in a gelatinized mass, thread-like, many-celled
........................................................................................................Gomphillus americanus
4.
On bark, wood, or rarely rock; spores (ascospores) usually in a dry mass, spherical to
ellipsoid, 1-2(-4)-celled................................................ key to Calicium and similar lichens
5.(1)
Apothecia deeply concave or opening by a pit-like hole; margin double, with a thalline
margin partially or entirely enclosing a well-developed excipulum (figs. 12h; 13d)
............................................................................................................................. see Key D
5.
Apothecia disk- or cup-shaped, or immersed in the thallus or in thalline warts, convex,
flat, or somewhat concave, not opening by a deep pitlike hole; margins various (figs. 12
a-c; 13 a-c)………………………. .....................................................................................6
6.
Apothecia buried in thalline warts, 1 or more per wart; spores very large and thickwalled, 1- to 2-celled, often with walls having 2 layers (fig. 15-l) .....................................7
6.
Apothecia disk- or cup-shaped, with the hymenium exposed; spores 1- to many-celled,
thin- or thick-walled........................................................................................................... 8
32
7.
Spores 1-celled; apothecia in fruiting warts opening to the surface with one or more
small ostioles with the hymenium largely enclosed (fig. 12g)..............................Pertusaria
7.
Spores 2-celled, constricted at the septum and easily breaking in two; apothecia in
fruiting warts broadly opened to the surface, pinkish or yellowish, sometimes with the
ascus tips showing as tiny glistening dots. Thallus white, thick or thin, sometimes
becoming sorediate; medulla K–, C+ red (lecanoric acid); spores 200-400 x 72-135 μm;
on moss, dead vegetation, wood, bark, or stones; arctic-alpine ... [Varicellaria rhodocarpa]
8.
Apothecia mazaedial: spores lying loose in a powdery mass within the exciple
............................................................................................................................................9
8.
Apothecia usually with a well-developed hymenium; spores remaining within the asci
until maturity, not forming a loose mass...........................................................................11
9.
Spores 4-celled, with unevenly thickened walls; ascomata with 2 chambers, one above
the other, the lower one immersed in the substrate, the upper one cylindrical, about 0.5
mm high; thallus UV+ yellow (lichexanthone), thin, continuous; southeast coastal plain ...
............................................................................................................ [Pyrgillus javanicus]
9.
Spores 1- to 2-celled, rarely submuriform; ascomata with one chamber; thallus UV– or
UV+ orange from yellow pigments (lichexanthone absent); mostly northern or western 10
10.
Exciple brown to black, well-developed; spores ellipsoid, 2-celled or submuriform in one
species (C. notarisii); thallus thin; apothecia immersed or superficial, not in thalline warts
............................................................................................................................Cyphelium
10.
Exciple pale and weakly developed; spores globose or broadly ellipsoid, 1- to 2-celled;
thallus thick; apothecia entirely immersed in thalline warts, disk level with surface ...........
....................................................................................................... Thelomma
11.(8)
Apothecia pale yellow, bright lemon- or yolk-yellow, orange, or red..............................12
11.
Apothecia green, gray, olive, brown, pink, or black.........................................................36
12.
Asci containing numerous spherical spores; apothecia tiny, 0.1-0.2 mm in diameter,
bright red, K–; rare................................................................... [Strangospora microhaema]
12.
Asci containing 1-32 spores; apothecia mostly larger than 0.2 mm in
33
diameter…………..13
13.
Apothecia blood red to cinnabar or pale red .....................................................................14
13.
Apothecia yellow or orange, not red.................................................................................17
14.
Spores polarilocular; apothecia pale red; California .............................................................
.....................................................................................Caloplaca luteominia var. bolanderi
14.
Spores thin-walled, not polarilocular ................................................................................15
15.
Apothecia biatorine; spores ellipsoid, 1-celled ................................................. Pyrrhospora
15.
Apothecia lecanorine; spores fusiform, 2- to 8-celled ......................................................16
16.
Southeastern and southwestern U.S.; thallus pale gray to greenish gray (lacking usnic
acid) ............................................................................................................... Haematomma
16.
Western and northern North America; thallus distinctly yellowish (usnic acid in the
cortex) ...............................................................................................................Ophioparma
17.(13) Pigmented tissues (epihymenium or thallus cortex) K+ deep purple or dark
red-purple (anthraquinones)...........................................................................................18
17.
Pigmented tissues K– or K+ pinkish (lacking anthraquinones) ........................................22
18.
Spores polarilocular, with a thickened septum .....................................................Caloplaca
18.
Spores 1- to several-celled or muriform, not polarilocular ...............................................19
19.
On dry soil; spores 1- to 2-celled, thin-walled...................................................... Fulgensia
19.
On rocks, bark, or bryophytes; spores 1- to several-celled or muriform ..........................20
20.
On rock, especially limestone; apothecia with thin, disappearing margins; spores 1-celled
.......................................................................................................................Protoblastenia
20.
On bark, bryophytes, or decaying vegetation; apothecia with prominent, persistent
margins; spores transversely septate or muriform.............................................................21
21.
Thallus olive to orange; spores with unevenly thickened walls, sometimes forming
34
spiralled locules ....................................................................................................Letrouitia
21.
Thallus white to pale gray; spores thin-walled, muriform, with many cells .........................
........................................................................................................................... Brigantiaea
22.(17) Apothecia biatorine, lacking algae in the margin .............................................................23
22.
Apothecia lecanorine (with algae in the margin) or cryptolecanorine (sunken into the
thallus, and lacking a recognizable exciple). ....................................................................28
23.
Spores 2- or more celled ...................................................................................................24
23.
Spores 1-celled..................................................................................................................27
24.
Apothecial margin fuzzy or tomentose (byssoid); paraphyses branched; asci club-shaped .
............................................................................................................................Byssoloma
24.
Apothecial margin smooth; paraphyses unbranched, slender; asci narrowly cylindrical .25
25.
Spores 4-8-celled, 16-48 per ascus ...................................................[Pachyphiale fagicola]
25.
Spores 2-celled, 8 per ascus ..............................................................................................26
26.
Thallus very thin, not gelatinous; photobiont Trentepohlia; hymenium K/I+ blue;
apothecia pale orange to pinkish orange, 0.2-1.5 mm in
diameter……………….Dimerella
26.
Thallus gelatinous; photobiont Chlorococcoid; hymenium K/I–; apothecia very pale
yellowish white, 0.3-0.4 mm in diameter.....................................................[Absconditella]
27.(23) Thallus leprose, bright yellow, UV+ orange (rhizocarpic acid); on rock
................................................................................................................. Psilolechia lucida
27.
Thallus smooth to granular, without soredia, greenish yellow, UV- (usnic acid); on bark
.............................................................................................................. Lecanora symmicta
28.(22) Apothecia cryptolecanorine, immersed in thallus, the disk flush with the thallus surface 29
28.
Apothecia lecanorine, broadly attached or constricted at the base; thallus pale yellow or
yellow-green or gray ........................................................................................................30
35
29.
Spores numerous in each ascus; thallus bright yellow................................ Acarospora key
29.
Spores 8 per ascus; thallus distinctly rusty orange or brownish gray
(shade forms) ............................................................................................ Ionaspis lacustris
30.
Apothecial disks bright yellow or bright yellow-orange...................................................31
30.
Apothecial disks pale greenish yellow, pale yellow, or pale pinkish orange....................34
31.
Thallus bright yolk-yellow (calycin).................................................................................32
31.
Thallus yellow-green or yellow-gray (usnic acid) ............................................................33
32.
Thallus distinctly lobed; lower surface corticate ................................................. Candelina
32.
Thallus edge indefinite, never lobed; lower surface without a
cortex………..Candelariella
33.
Apothecial disks not pruinose; apothecial margin flush with disk, becoming thin and
disappearing in maturity.........................................................................Lecanora strobilina
33.
Apothecial disks yellow pruinose; apothecial margin prominent………..Lecanora
cupressi
34.(30) Spores conspicuously thick-walled ......................................................................Pertusaria
34.
Spores thin-walled ............................................................................................................35
35.
Spores 10-20 μm long; paraphyses unbranched; apothecia generally less than 1.5 mm in
diameter..................................................................................................................Lecanora
35.
Spores over 25 μm long; paraphyses highly branched; apothecia frequently over 1.5 mm
in diameter ........................................................................................................ Ochrolechia
36.(11) Apothecia lacking algae in the margin: lecideine, biatorine, or without an
exciple; apothecia superficial, or sunken into thallus but retaining a distinct, usually
pigmented exciple .............................................................................................................37
36.
Apothecia with algae in the margin or in thalline tissue surrounding the sunken disk:
lecanorine (superficial) or cryptolecanorine (sunken into the thallus, usually without a
distinct exciple; but see couplets 107-110) ......................................................................98
36
37.
Spores one-celled, spherical to narrowly ellipsoid (if spores are fusiform with length to
width ratio more than 3:1, they may be species that become septate later in development;
see, for example, Icmadophila) .........................................................................................38
37.
Spores septate (2- or more celled), broadly ellipsoid to thread-shaped ............................62
38.
Apothecia biatorine, with a colorless to pigmented exciple that is soft rather than brittle,
with a clearly defined and usually radiating cellular structure throughout, or exciple very
reduced and indistinct .......................................................................................................39
38.
Apothecia lecideine, with an exciple that is very dark brown to black, usually somewhat
carbonized, occasionally brittle, with a poorly defined cellular structure, at least on the
outer part ...........................................................................................................................58
39.
Spores conspicuously thick-walled (more than 2.5 μm thick) ..........................................40
39.
Spores thin-walled relative to the spore size (usually less than 1.5 μm thick)..................41
40.
Apothecial disks black; thallus white to pale greenish gray; usually with a blood-red area
below the hypothecium; epihymenium green; spores 1or 2 per ascus; thallus cortex K+
yellow (atranorin)…………………………………………………………….Mycoblastus
40.
Apothecial disks dark reddish brown; thallus brown or pale; tissue below hypothecium
not red; epihymenium brown; spores 8 per ascus; thallus cortex K– (atranorin absent)
………………………………………………………………………….Japewia tornoensis
41.
Directly on rock ................................................................................................................42
41.
On bark, wood, soil, or mosses or dead vegetation...........................................................43
42.
Thallus pale gray, C+ pink (gyrophoric acid); apothecia pinkish or brown, with a rough
or ragged margin; asci cylindrical, usually with uniformly K/I+ pale blue walls including
the tip (fig. 14-l)...................................................................................................... Trapelia
42.
Thallus dark, brownish, olive, or dark gray, C–; apothecia dark brown to black, with a
smooth margin; ascus club-shaped, the tips staining as pale and dark blue layers in K/I
(fig. 14d) ................................................................................................................ Fuscidea
37
43.
On moss, soil, or dead vegetation .....................................................................................44
43.
On bark or wood ...............................................................................................................50
44.
Spores numerous within the ascus, 5-8 x 2-2.3 μm, colorless. Apothecia red-brown,
translucent when wet, 0.1-0.5 mm in diameter…………………..[Sarcosagium
campestre]
44.
Spores 8 per ascus.............................................................................................................45
45.
Hypothecium dark red-brown, brown, or black................................................................46
45.
Hypothecium colorless, yellowish, or very pale brown ....................................................48
46.
On moss; hypothecium distinct from exciple; paraphyses mostly unbranched; thallus pale
greenish gray to whitish; asci Porpidia-type .....................................................................47
46.
On soil or decayed peat; hypothecium merging with exciple; thallus dark brown or olivebrown; asci K/I–...............................................................................................Placynthiella
47.
Thallus thick, granular to warty or areolate; tips of paraphyses somewhat expanded and
brown; spores 10-16(-19) x 3.5-5(-6) μm, rarely 2-celled
............................................................................................["Mycobilimbia" berengeriana]
47.
Thallus thin, membranous; tips of paraphyses not expanded; spores 10-16(-19) x 4.5-6(7), 2-celled spores often mixed with 1-celled spores………...["Mycobilimbia"
hypnorum]
48.(45) Thallus cortex and medulla C+ pink, KC+ red (gyrophoric acid); epihymenium shades of
olive or green; paraphyses highly branched......................................................Trapeliopsis
48.
Thallus cortex or medulla C– or C+ orange, KC– or KC+ yellow to orange (gyrophoric
acid absent); epihymenium shades of yellow or brown, or colorless; paraphyses only
branched at tips .................................................................................................................49
49.
Thallus mostly pale green or gray-green; apothecia usually pale to dark yellowish brown,
but sometimes dark brown; exciple almost colorless or yellowish internally, sometimes
pigmented at the outer edge, with slender radiating hyphae .....................................Biatora
49.
Thallus dark brown or olive, thick, mostly continuous or dispersed areolate;; apothecia
38
dark brown to almost black; exciple distinct, dark brown, with large cells ..........................
............................................................................................................ Lecidoma demissum
50.(43) Spores numerous within the ascus. Apothecia black, 0.3-0.5 mm, convex; rare and
widely distributed in the west to the arctic and Great Lakes…………..[Strangospora
moriformis]
50.
Spores 8 per ascus.............................................................................................................51
51.
Thallus leprose, dark yellow, C+ orange (xanthones); epihymenium K+ deep purple
(anthraquinones) ................................................................................. Pyrrhospora quernea
51.
Thallus continuous to areolate or granular, C–, C+ orange, or C+ pink; epihymenium K–
..........................................................................................................................................52
52.
Spores almost spherical, 5-10(-13) x 4-8 μm; apothecia brown to black; margins PD+
red. Very common on conifers in western mountains………………….[Lecanora
fuscescens]
52.
Spores ellipsoid; apothecial margins PD– or rarely PD+ .................................................53
53.
Apothecia greenish to yellow-green; thallus cortex KC+ gold (usnic acid)
.............................................................................................................. Lecanora symmicta
53.
Apothecia pale beige, pinkish, brown, or black; thallus cortex KC– (usnic acid absent)....
..........................................................................................................................................54
54.
Paraphyses highly branched and anastomosing; thallus often C+ pink (gyrophoric acid)
..........................................................................................................................................55
54.
Paraphyses unbranched except for tips; thallus C–...........................................................56
55.
Thallus reddish brown to dark brown, granular to isidiate; hypothecium brown, merging
with exciple......................................................................................................Placynthiella
55.
Thallus pale gray to dark greenish gray; hypothecium pale to
colorless………Trapeliopsis
56.
Thallus a mass of isidia or coarse granules (squamulose in younger parts); spores 9-12 x
39
1.8-2.3 μm............................................................ granular forms of Phyllopsora parvifolia
56.
Thallus continuous, areolate to somewhat granular, never squamulose; spores broader
than 2.5 μm .......................................................................................................................57
57.
Thallus KC+ orange (somewhat faint); spores broad (length to width ratio 1.5-2:1).
Apothecia dark red-brown, 0.2-0.4 mm in diameter; very common on trees in the east
...........................................................................................................[Pyrrhospora varians]
57.
Thallus KC–; spores narrow (length to width ratio 2-3:1)........................................Biatora
58.(38) Spores 8 per ascus ............................................................................................................59
58.
Spores many more than 8 per ascus ..................................................................................60
59.
Spores brown, walls thickened at equator; prothallus thick, black. Arctic-alpine
.......................................................................................................Orphniospora moriopsis
59.
Spores colorless, walls unifornly thin; prothallus present or absent
................................................................................ key to Lecidea and Lecidea-like crusts
60.
Thallus well-developed, often lobed at the margin; medulla C+ pink (gyrophoric acid);
apothecia immersed in thallus between the areoles, disk flush with thallus; epihymenium
green or brown; spores globose or broadly ellipsoid…………………………. Sporastatia
60.
Thallus mostly within the substrate, absent from view, lacking gyrophoric acid; apothecia
sessile; epihymenium brown or black; spores narrowly ellipsoid.....................................61
61.
Apothecial disks rough, umbonate (with sterile columns in apothecial disk); apothecia
often concave or crushed due to crowding; apothecial margin rough or minutely fissured;
exciple dark brown to carbonized
throughout………………………………….Polysporina
61.
Apothecial disks smooth, without umbos; apothecia flat or soon convex; apothecial
margin smooth; exciple dark or carbonized only at edge, pale internally........... Sarcogyne
62.(37) Spores muriform...............................................................................................................63
62.
Spores only transversely septate .......................................................................................69
40
63.
Apothecia biatorine, concave, pale and waxy-looking, pink, yellow, or pale orange;
epihymenium and hypothecium colorless, paraphyses unbranched; asci narrowly
cylindrical .............................................................................................................. Gyalecta
63.
Apothecia biatorine or lecideine, flat to somewhat convex, brown or black, not waxy;
epihymenium pigmented; hypothecium colorless to yellowish, or brown to black;
paraphyses unbranched or abundantly branched or difficult to distinguish; asci clubshaped ...............................................................................................................................64
64.
Directly on rock; spores halonate or not ...........................................................................65
64.
On bark, soil, leaves, or mosses; spores not halonate .......................................................66
65.
Spores halonate ................................................................................................ Rhizocarpon
65.
Spores not halonate .................................................................. [Buellia key (Diplotomma)]
66.
On leaves; apothecia 0.2-0.5 mm in diameter, disks pale to dark brown; apothecial
margin even with disk.......................................................................................... Calopadia
66.
On bark, mosses, or soil; apothecia usually more than 0.5 mm in diameter, disks black or
almost black; apothecial margin prominent ......................................................................67
67.
Apothecial margin yellow or bright orange, paler than disk; exciple radiate, yellow;
epihymenium K+ red or deep purple-red (anthraquinones); on bark in tropical and
subtropical regions ................................................................................................Letrouitia
67.
Apothecial margin black, the same color as the disk; exciple dark brown at edge, pale
internally; epihymenium unchanged in K (anthraquinones absent); on bark, soil, or
mosses in northern regions................................................................................................68
68.
Spores 1 per ascus, over 50 μm long ................................................................... Lopadium
68.
Spores 8 per ascus, less than 30 μm long........................................[Buellia (Diplotomma)]
69.(62) Apothecia pink to yellowish pink, 1.5-4 mm in diameter; thallus green to greenish white,
continuous, sometimes verrucose; growing over well-rotted wood or peat......Icmadophila
69.
Apothecia pale brown to black (if pinkish, then under 1 mm in diameter); thallus and
habitat various...................................................................................................................70
41
70.
Spores 4- or more celled ...................................................................................................71
70.
Spores 2-celled..................................................................................................................82
71.
Photobiont blue-green; thallus with a conspicuous blue-black prothallus
......................................................................................................................... Placynthium
71.
Photobiont green; lacking a blue-black prothallus............................................................72
72.
Spores brown ............................................................................................................ Buellia
72.
Spores colorless ................................................................................................................73
73.
Spores slender and needle-shaped, some often curved or bent; length to width ratio 7:1 or
more ..................................................................................................................................74
73.
Spores ellipsoid, narrowly ellipsoid, or fusiform, straight; length to width ratio less than
7:1
74.
....................................................................................................................76
Thallus bright yellow; apothecia Arthonia-like; apothecial margin absent; growing on
soil or lichens ................................................................................................. Arthrorhaphis
74.
Thallus greenish, olive-gray, or brownish; apothecia biatorine, with a persistent margin;
on bark, mosses, or dead vegetation .................................................................................75
75.
Spores many per ascus; thallus edge definite, with a brown prothallus; exciple pale
internally, dark brown at edge; ascus Fuscidea-type, with a layered appearance when
stained with K/I...............................................................................Ropalospora chlorantha
75.
Spores 8 per ascus; thallus edge usually indefinite; prothallus present or absent; exciple
pigmented in various patterns, or sometimes colorless; ascus Bacidia-type, with a more
or less uniformly dark blue tholus in K/I
...........................................................Bacidia key (including Bacidina & Scoliciosporum)
76.(73) Thallus thick, areolate to squamulose; apothecia black; on soil or rocks, usually in arid or
alpine habitats………………………………………………………………………Toninia
76.
Thallus mostly thin, membranous to leprose or granular; apothecia very pale to black; on
bark, moss, or dead vegetation, occasionally on rock or soil............................................77
42
77.
Exciple absent; asci broad, balloon-shaped, thick at the tips but almost entirely K/I–.........
.............................................................................................................. Arthonia
77.
Exciple usually present, although sometimes indistinct, lecideine or biatorine; asci clubshaped, K/I+ or K/I– …………………. ..........................................................................78
78.
Exciple lecideine, dark brown to black.............................................................................79
78.
Exciple biatorine, usually pale; apothecia not, or lightly, pruinose ..................................80
79.
Apothecia usually pruinose or dull; margin not radially fissured .......................Lecanactis
79.
Apothecia black, shiny, never pruinose; margin broken up by radial fissures; on moss;
arctic-alpine..............................................................................[Sagiolechia rhexoblephara]
80.
Apothecial margin prominent or even with disk, fuzzy or tomentose (byssoid); apothecia
flat when mature; epihymenium colorless ..........................................................Byssoloma
80.
Apothecial margin becoming thin and disappearing in maturity, or absent; apothecia flat
to convex or hemispherical; epihymenium shades of yellow, brown, or green ................81
81.
Paraphyses unbranched or slightly branched at the tips; epihymenium always C–
......................................................................................................... Mycobilimbia
81.
Paraphyses highly branched and anastomosing; epihymenium C– or often C+ pink
(usually disappearing quickly) ................................................................................ Micarea
82.(70) Spores dark brown when mature ......................................................................................83
82.
Spores colorless ...............................................................................................................84
83.
Spores halonate; ascus tips K/I mostly negative; on rock ............................... Rhizocarpon
83.
Spores not halonate; ascus tips K/I+ dark blue, at least in part; on various substrates ......
...................................................................... Buellia key (Buellia, Amandinea, Diploicia)
84.
On rock .............................................................................................................................85
84.
On bark, wood, moss, leaves, or dead vegetation .............................................................89
43
85.
Hypothecium yellow-brown; asci balloon-shaped (fig. 14n); exciple absent (fig. 13f); on
maritime rocks along both coasts.......................................................[Arthonia phaeobaea]
85.
Hypothecium dark brown to black; asci club-shaped; exciple well-developed, biatorine or
lecideine ............................................................................................................................86
86.
Photobiont blue-green; prothallus conspicuous, blue-green; epihymenium emerald green..
......................................................................................................................... Placynthium
86.
Photobiont green; prothallus conspicuous or not, but not blue-green; epihymenium
brown, reddish, to olive or greenish..................................................................................87
87.
Spores small, 6-15 x 2.5-5; paraphyses expanded and pigmented at tips; widespread
temperate to boreal.............................................................................. [Catillaria chalybeia]
87.
Spores large 12-30 x 3-14.................................................................................................88
88.
Spores narrow, 3-5 μm wide, not halonate; or rock or soil; epihymenium gray to purplishToninia
88.
Spores broad, 8-12 μm wide, halonate; on rock; epihymenium brown or greenish, rarely
reddish.............................................................................................................. Rhizocarpon
89.(84) Apothecial margins fuzzy and tomentose (byssoid); thallus sometimes UV+ orange; on
bark or leaves on the southeastern coastal plain .................................................Byssoloma
89.
Apothecial margin, if present, smooth, not at all fuzzy; thallus always UV–; mostly East
Temperate, northern, or western, on various substrates.................................................90
90.
Hypothecium dark reddish brown to black .......................................................................91
90.
Hypothecium colorless to pale brown...............................................................................93
91.
Paraphyses much branched and anastomosing; exciple absent (fig. 13f); asci balloonshaped (fig. 14n). Spores 8-15 x 2.5-5; apothecia black, without a margin (fig. 12c);
epihymenium K–; forming white patches on poplar bark; common in boreal region from
maritimes to B.C. ............................................................................... [Arthonia patellulata]
91.
Paraphyses mostly unbranched except for tips; exciple lecideine or biatorine; asci usually
club-shaped or cylindrical.................................................................................................92
92.
Spores 6-12 x 2.5-4; epihymenium brown; hypothecium brown to black; widely
44
distributed on bark of all kinds, but especially conifers…………[Catillaria
glauconigrans]
92.
Spores 13-18(-24) x 5-7(-8); epihymenium black to greenish; hypothecium red-brown;
Appalachian-Great Lakes distribution, on hardwoods, especially beech and maple
.................................................................................................................. Megalaria laureri
93.(90) On leaves of conifers and broadleaf trees and shrubs, rarely on twigs; apothecia tiny, 0.10.3 mm in diameter, very pale beige; spores 9.5-14(-16) x 3.5-7 μm, tapered and
constricted at the septum; Pacific Northwest and southeastern U.S.
.......................................................................................................... [Fellhanera bouteillei]
93.
On bark, wood, or moss; apothecia (0.1-) 0.3-2 mm in diameter, pink to brown, gray, or
black..................................................................................................................................94
94.
Paraphyses much branched and anastomosing. On rotting wood and occasionally bark;
thallus dark green, granular; exciple poorly differentiated; apothecia pale gray to graybrown or black; epihymenium K+ violet; spores 1- to 2-celled; very widespread ...............
................................................................................................................. [Micarea prasina]
94.
Paraphyses mostly unbranched except at the tips. ............................................................95
95.
Spores 2-4 μm wide ..........................................................................................................96
95.
Spores 4-7 μm wide ..........................................................................................................97
96.
Apothecia pruinose, very pale pinkish to black or mottled; paraphyses barely expanded
and not pigmented at the tips; hypothecium colorless; spores 8-16 x 2.5-3.5 μm; on bark
of all kinds; coastal ............................................................................ Cliostomum griffithii
96.
Apothecia not pruinose, brown to black, not pinkish; paraphyses expanded and
pigmented at the tips; hypothecium pale brown; spores 8-10 x 2-4 μm; on bark,
especially poplars and white cedar, from North Dakota to southern Ontario and eastern
U.S.
....................................................................................................... [Catillaria nigroclavata]
97.
Apothecia black, mostly flat to lightly convex, with a persistent margin; spores all 2celled, 10-15 x 5-7 μm; on bark of different kinds, especially poplar and cedar; widely
45
distributed, especially in north temperate to boreal region and California ...........................
....................................................................................................... [Catinaria atropurpurea]
97.
Apothecia yellow- to red-brown, convex; spores 1- to 2-celled, 11.5-19 x 4-7 μm; on
moss, rarely bark; widespread..................................................................... Biatora vernalis
98.(36) Spores 2- or more celled...................................................................................................99
98.
Spores 1-celled................................................................................................................112
99.
Spores pale to dark brown when mature, mostly 2-celled ..............................................100
99.
Spores colorless, 2- or more celled .................................................................................102
100.
Spores 4 per ascus, mostly over 35 μm long; cyanobacteria in the continuous crustose
thallus, and green algae in lobes forming the apothecial rim; apothecia concave, cup-like,
reddish or orange-brown........................................................................ Solorina spongiosa
100.
Spores 8 or more per ascus, mostly under 35 μm long; photobiont green with no
cephalodia; apothecia flat or soon convex, black to dark brown ....................................101
101.
Thallus thin, areolate, forming round, often lobed rosettes on rock; spores 8-12 μm long,
with thin, uniformly thickened walls .................................................................. Dimelaena
101.
Thallus not forming rosettes; if lobed, then not on rock; on bark, soil, mosses, or rock;
spores 11-40 μm long, mostly with unevenly thickened walls at least when young
.............................. Rinodina key (Rinodina, Phaeorrhiza and Hyperphyscia)
102.(99) Spores muriform, 1 per ascus; apothecia buried in thallus ......................................Phlyctis
102.
Spores transversely septate, 2- to 8-celled, 8 per ascus ..................................................103
103.
Spores 2-celled, ellipsoid, straight or bean-shaped.........................................................104
103.
Spores 4- to 8-celled, fusiform........................................................................................105
104.
Spore walls evenly thickened, thin-walled; apothecial disks pale to dark brown,
frequently pruinose .................................................................................................. Lecania
104.
Spore walls with a thickened septum (polarilocular); apothecial disks brown to almost
black, sometimes pruinose ....................................................................................Caloplaca
46
105.
Hypothecium colorless; apothecia lightly pruinose or not pruinose, brown to black .....106
105.
Hypothecium brown to black; apothecia white because of a heavy pruina, black beneath.
Mainly in coastal California............................................................................................108
106.
Spores straight to slightly curved, with rounded ends, not twisted in the ascus; apothecia
pale to very dark brown or black, pruinose or not; on trees or
rocks………………Lecania
106.
Spores sinuous, curved and twisted in the ascus; apothecia pinkish brown to red-brown,
lightly pruinose or without pruina; on bark in temperate to boreal regions....................107
107.
Thallus producing long, slender, sometimes branched isidia; thallus K–, PD–; coastal
forests of Pacific northwest……………………………………Loxosporopsis corallifera
107.
Thallus lacking isidia; thallus K+ deep yellow, PD+ orange (thamnolic acid); eastern
temperate forests to boreal forest ........................................................................ Loxospora
108.(105) Paraphyses highly branched and anastomosing; spores halonate; apothecia appear
lecanorine when immersed in the thallus, but with a dark brown to black exciple; thallus
cortex and medulla usually C+ red, KC+ red; on
rocks......................………Lecanographa
108.
Paraphyses mostly unbranched or branched only at the tips; spores not halonate;
apothecia truly lecanorine or appear to be lecanorine; thallus cortex and medulla C+ or
C–, KC+ or KC–; on rocks or bark .................................................................................109
109.
Hypothecium extending like a peg to the substrate (seen in a section through the center of
the apothecium); thallus cortex of North American species C–, KC–; California..........110
109.
Hypothecium lens-shaped, not extending to the substrate; thallus cortex or medulla C+
red, KC+ red (erythrin, lecanoric acid); California or
Florida……………………….Dirina
110.
Thallus PD–; ascomatal disks irregular in shape, lobed or slightly elongate; on bark or
rock ................................................................................................[Roccellina franciscana]
110.
Thallus PD+ yellow or red-orange; ascomatal disks round; on bark or wood…………111
47
111.
Thallus creamy white when fresh, PD+ yellow (psoromic acid); on trees along coast and
into chaparral, San Francisco to Los Angeles
...................................................[Sigridea californica (syn. Schismatomma californicum)]
111.
Thallus tan to greenish brown, PD+ red-orange (unknown substance); on bark or wood,
Santa Catalina Island.......................................................................... Roccellina conformis
112.(98) Thallus gelatinous (and reddish) when wet; photobiont blue-green (Gloeocapsa);
apothecia opening by a deep pore; on streamside or lakeside rocks……………Pyrenopsis
112.
Thallus not gelatinous, most with green photobionts; apothecial disks broad; on various
substrates.........................................................................................................................113
113.
Cephalodia conspicuous on the thallus surface, pink to brown, disklike and more or less
lobed; apothecia usually pink................................................................................ Placopsis
113.
Cephalodia absent or very inconspicuous; apothecia pink, brown, or
black…………...114
114.
Apothecia more or less immersed in thallus, disk flush with thallus surface
(cryptolecanorine) ...........................................................................................................115
114.
Apothecia superficial: adnate or constricted at base .......................................................125
115.
Spores numerous in each ascus...................... Acarospora key (Acarospora, Pleopsidium)
115.
Spores 8 per ascus...........................................................................................................116
116.
Thallus distinctly lobed at the margin.............................................................................117
116.
Thallus margin without distinct lobes .............................................................................118
117.
Thallus and apothecia heavily pruinose, thin and clearly crustose; apothecia black
(beneath the pruina); thallus cortex and medulla K–, PD– .......................Aspicilia candida
117.
Thallus and apothecia without pruina, very thick and almost foliose; apothecia dark
brown; thallus cortex and medulla K+ yellow becoming blood red, PD+ yellow
(norstictic acid) .................................................................................................. Lobothallia
48
118.
Spores conspicuously thick-walled.................................................... Megaspora verrucosa
118.
Spores thin-walled ..........................................................................................................119
119.
Thallus pale yellow to yellowish white, thick, KC+ gold (usnic acid); epihymenium
green
..............................................................................................................Lecanora marginata
119.
Thallus not at all yellowish, KC– (lacking usnic acid); epihymenium shades of yellow,
brown, or green, or colorless ..........................................................................................120
120.
Apothecial disks C+ yellow, heavily pruinose; on siliceous rocks ..........Lecanora rupicola
120.
Apothecial disks C–, pruinose or not (if pruinose, then on calcareous
rock)…………..121
121.
Apothecial disks black; epihymenium shades of green ..................................................122
121.
Apothecial disks pink to brown or gray, rarely darkening to black; epihymenium shades
of gray, yellow, or brown................................................................................................124
122.
Epihymenium yellow-olive to olive-brown, unchanged or intensifying wiith nitric acid.
Ascus K/I –; widespread and common .................................................................. Aspicilia
122.
Epihymenium blue-green to olive-green, changing to wine-red with nitric acid, or
unchanged .......................................................................................................................123
123.
Mostly arctic-alpine species; thallus thin, continuous to rimose; ascus K/I –.. Ionaspis key
123.
East Temperate; thallus thick, white to pale gray, areolate; ascus with a K/I+ blue tip
............................................................................................................. Lecanora oreinoides
124.(121) Ascus tips with a IKI+ dark blue "plug" in a light blue tholus (Porpidia-type);
spores ...... always halonate; on dry rocks; apothecial disks very dark brown or red-brown
(especially when wet)......................................................................................... Bellemerea
124.
Ascus tips entirely IKI–; spores not, or sometimes vaguely halonate; on wet or
submerged rocks, or on dry rocks; apothecia pinkish to
brown…………………………...Ionaspis key
49
125. (114) Spores conspicuously thick-walled (more than 2.5 μm thick).....................................126
125.
Spores thin-walled (usually less than 1.5 μm) ................................................................128
126.
Apothecia pale, pruinose or not ........................................................................ Ochrolechia
126.
Apothecia black, sometimes whitened by pruina............................................................127
127.
Epihymenium olive to greenish, K+ more clearly green; spores 30-65 μm long, 8 per
ascus; boreal to arctic, on trees, bryophytes, and vegetation ............. Megaspora verrucosa
127.
Epihymenium brown, K– or K+ violet; spores usually longer than 60 μm, 1-8 per ascus;
widespread on various substrates..........................................................................Pertusaria
128.(125) Spores very numerous in the ascus, 3-6 x 1.5-2.5 μm. Asci similar to
Fuscidea-type; apothecia dark brown to almost black, 0.4-1 mm in diameter; thallus
brownish gray, bumpy; on trees; East Temperate ................................. [Maronea constans]
128.
Spores 1-32 per ascus, larger than 6 x 2.5 μm ................................................................129
129.
Hymenium purple; hypothecium brown; apothecia pitch black
................................................................................................................... Tephromela atra
129.
Hymenium colorless; hypothecium colorless to yellowish, rarely brown; apothecia
various shades of pink and brown, or black....................................................................130
130.
Spores with rough, bumpy, sculptured walls (clearly seen under 400X magnification);
thallus brownish to green, granular to, more commonly, squamulose; on mossy logs,
peat, and soil in boreal to arctic regions..........................................................................131
130.
Spore walls smooth; thallus rarely squamulose, pale to dark; on various substrates ......132
131.
Photobiont blue-green...........................................................................Pannaria pezizoides
131.
Photobiont green ................................................................................... Psoroma hypnorum
132.
Spores over 25 μm long; paraphyses highly branched; apothecia pale, usually pinkish to
yellow-pink ....................................................................................................... Ochrolechia
132.
Spores mostly 10-20 μm long; paraphyses mostly unbranched; apothecia pale to dark
brown or black ................................................................................................................133
50
133.
On dry soil; thallus clearly lobed at the margins, pale yellowish to chalky white and
pruinose, rarely brownish-green ........................................................................Squamarina
133.
On rock, wood, bark, dead vegetation, or peat; thallus not yellowish white and chalky
pruinose...........................................................................................................................134
134.
Thallus pale greenish gray to pinkish white, C+ pink (gyrophoric acid); apothecial
margin often with a frayed or ragged appearance; on rock, especially small stonesTrapelia
134.
Thallus pale or dark, C– in North American species (lacking gyrophoric acid); apothecial
margins smooth or bumpy but not ragged; on various substrates ...................................135
135.
On bark, wood, mosses, or dead vegetation...........................................................Lecanora
135.
On rock ...........................................................................................................................136
136.
Thallus distinctly lobed at the margin.............................................................................137
136.
Thallus not lobed ............................................................................................................138
137.
Thallus cortex K+ yellow becoming blood red (norstictic acid), rarely K-; tips of
paraphyses septate and constricted like a string of beads (moniliform); asci K/I –
........................................................................................................................... Lobothallia
137.
Thallus cortex K– or K+ yellow (norstictic acid absent); tips of paraphyses not appearing
"beaded;" asci with K/I+ blue tips ........................................................................Lecanora
138.
Ends of spores pointed; thallus grayish brown to reddish brown, shiny; apothecial disks
chocolate brown and shiny............................................................................. Protoparmelia
138.
Ends of spores rounded; thallus rarely brown and shiny; apothecia various colors
........................................................................Lecanora key (Lecanora, Rhizoplaca)
KEY G: SQUAMULOSE LICHENS
1.
Thallus gelatinous when wet...............................................................................................2
1.
Thallus dull or shiny, but not gelatinous when wet ............................................................3
2.
Thallus without a cortex...................................................................................... Collema
51
2.
Thallus with a cellular cortex.......................................................................... Leptogium
3.
Thallus greenish yellow, yellow-green, yellow, or orange .................................................4
3.
Thallus pale green, gray, brown, olive, or black, without a yellowish or orange tint.......17
4.
Thallus greenish yellow or yellowish-green .......................................................................5
4.
Thallus bright yolk-yellow, lemon yellow, or shades of orange.......................................10
5.
Directly on rock ..................................................................................................................6
5.
On soil................................................................................................................................ 8
6.
Apothecia entirely immersed in thallus, disk flush with thallus; asci containing numerous
spores .................................................................................................. Acarospora contigua
6.
Apothecia sessile or constricted at base; asci containing 8 spores......................................7
7.
Apothecia constricted at base; disks yellow-orange, lightly pruinose; thallus not forming
radiate, lobed rosettes.................................................................. Rhizoplaca subdiscrepans
7.
Apothecia sessile; disks pale greenish yellow, not pruinose; thallus usually forming
distinctly lobed rosettes.............................................................................Lecanora muralis
8.
Squamules ascending, with the lower surface easily seen; thallus cortex KC+ yellow,
UV– (usnic acid) .................................................. Cladonia species, especially C. robbinsii
8.
Squamules closely appressed or lifting only at the edges; apothecia frequently seen,
black, directly on the squamules; thallus cortex KC–, UV+ orange (rhizocarpic acid)......9
9.
Squamules flat, often turned up at the edges, sturdy; spores one-celled; southwestern and
south central USA ........................................................................................... Psora icterica
9.
Squamules closely appressed, convex, fragile; spores needle-shaped, 9- to 12-celled;
boreal to arctic................................................................................. Arthrorhaphis citrinella
10.(4)
Thallus yellow-orange or orange-yellow; thallus cortex K+ dark purplish
(anthraquinones) ...............................................................................................................11
10.
Thallus yolk-yellow or lemon yellow; thallus cortex K– (anthraquinones absent) ..........14
52
11.
Squamules ascending, with white lower surface easily seen; on bark or rock......................
..............................................................................................................................Xanthoria
11.
Squamules ascending or closely appressed; lower surface, if visible, yellow; on wood,
soil, or rocks......................................................................................................................12
12.
Squamules thick, yellow to orange, shiny, ascending, with deep yellow lower surface
clearly visible; apothecia black, hemispherical and marginless; spores spherical,
colorless, 1-celled, 3-4 μm in diameter; Texas to Mexico, on limestone[Xanthopsorella texana]
12.
Squamules mostly thin, appressed, lower surface not easily seen; apothecia orange,
clearly rimmed ..................................................................................................................13
13.
On soil in the interior of the continent north to the arctic; spore 1- or 2-celled, with uniformly
thin walls.............................................................................................................. Fulgensia
13.
On rock, often in maritime or nearby coastal localities; spores 2-celled, polarilocular ........
..............................................................................................................................Caloplaca
14.
Squamules ascending, with the pale lower surface easily seen; mainly on bark; thallus
thin
............................................................................................................................ Candelaria
14.
Squamules closely appressed; on rock or soil; thallus thick .............................................15
15.
Apothecial disks black, convex; spores needle-shaped, septate ...........................................
........................................................................................................ Arthrorhaphis citrinella
15.
Apothecial disks brown or yellow; spores globose or ellipsoid, one-celled. ....................16
16.
Apothecia sessile, lecanorine; apothecial disks yellow; spores 12.5-16.0 x 4.8-6.5 μm, 8
per ascus............................................................................................ Candelariella rosulans
16.
Apothecia entirely immersed in thallus, disk level with thallus, apothecial disks dark
brown, or reddish brown to orange-brown, spores 3-4
2-3 μm, many per ascus..............
........................................................................................................................... Acarospora
17.(3)
Squamules ascending or erect, flat or cylindrical, with the lower surface easily seen......18
53
17.
Squamules closely appressed, or lifting only at the edges ................................................23
18.
Thallus dwarf fruticose, consisting of erect cylindrical to somewhat flattened lobes.......19
18.
Thallus composed of flat, clearly dorsiventral squamules ................................................20
19.
Photobiont blue-green; spores globose, 1-celled, more than 8 per ascus..............................
..................................................................................................................Peltula cylindrica
19.
Photobiont green; spores ellipsoid, muriform, 2 per ascus ............ Endocarpon pulvinatum
20.
Squamules elongate and branched; apothecia clustered in groups on the squamule tips;
thallus cortex K+ yellow (atranorin); very rare………………………Gymnoderma
lineare
20.
Squamules not more than 3x longer than broad; apothecia occurring singly, thallus cortex
K– (atranorin absent); relatively common species............................................................21
21.
Aquatic or semiaquatic, growing on rocks in or near water (streams or lakeshore);
fruiting bodies are perithecia, appearing as black dots on the lobe surface
.................................................................................................... Dermatocarpon
luridum
21.
Not aquatic or semiaquatic; fruiting bodies, when present, are apothecia ............ 21a
21a.
Squamules lobed or finely divided, apothecia raised on a stalk or stipe; thallus cortex C–,
KC–, (gyrophoric acid absent as main compound)...Cladonia species (e.g., C. caespiticia)
21a.
Squamules round, or scalloped (with rounded lobes); apothecia sessile; thallus cortex C+
pink or red, KC+ pink or red (gyrophoric acid)................................................................22
22.
Thallus pale gray; squamules thick, distinctly lobed, with edges turned down; apothecia
flat or convex, pink to lead gray, with a prominent margin ............. Trapeliopsis wallrothii
22.
Thallus olive to greenish brown; squamules thin, round, with edges turned up, apothecia
becoming hemispherical, dark brown to black; apothecial margin absent.. Psora nipponica
23.(17) Photobiont (and exposed algal layer) blue-green ..............................................................24
23.
Photobiont (and algal layer) green ................................................................................... 29
54
24.
Soredia present..................................................................................................................25
24.
Soredia absent ...................................................................................................................26
25.
Thallus olive brown; on rocks in arid or damp habitats in the western interior; soredia
farinose; squamules round, with edges turned down or relatively flat......... Peltula euploca
25.
Thallus blue-gray to yellowish gray, on bark or sometimes rock in humid, boreal to
subarctic habitats; soredia coarsely granular; squamules scalloped (with rounded lobes)
or elongate, with edges turned up ........................................................... Pannaria conoplea
26.
Squamules extremely narrow-lobed (less than 0.3 mm wide), building into an areolate
crust with abundant isidia; apothecia black, lecideine………………..Placynthium nigrum
26.
Squamules mostly broader than 0.3 mm, isidiate or not; apothecia red-brown, lecanorine
or biatorine........................................................................................................................27
27.
Thallus olive or olive-brown; apothecia partly or entirely immersed in the thallus;
photobiont Scytonema or Anacystis .................................................................................28
27.
Thallus brownish or gray-brown; apothecia sessile or constricted at the base; photobiont
Nostoc ............................................... Pannaria key (Pannaria, Fuscopannaria, Parmeliella)
28.
Spores ellipsoid, 8 per ascus ...................................................................Heppia conchiloba
28.
Spores globose or broadly ellipsoid, up to 100 per ascus ......................................... Peltula
29.(23) Soredia present.................................................................................................................30
29.
Soredia absent ...................................................................................................................34
30.
On soil or rock, rarely wood; cortex and medulla K+ persistently yellow, PD+ orange
(stictic acid)............................................................................................... Baeomyces rufus
30.
On bark, wood, mosses, or lichens; thallus cortex and medulla K–, PD–, (stictic acid
absent)...............................................................................................................................31
31.
Soralia raised and finally cuplike; thallus olive-brown, with round, tiny squamules (less
than 0.5 mm in diameter); apothecia blue-gray, biatorine; spores 4-celled, fusiform;
55
California ............................................................................................[Waynea californica]
31.
Soralia on lobe surface or on margin of lower surface, not raised or
cuplike……………32
32.
Thallus pale green; squamules thin with a raised, thickened rim, separate, not
overlapping; on mosses and lichens (especially cyanobacterial lichens), also bark and
wood in humid habitats, thallus cortex and medulla C–, KC–.......... Normandina pulchella
32.
Thallus olive to brownish or gray-brown; squamules thick, commonly overlapping like
shingles; on wood (especially charred wood) or conifer
bark……………………………33
33.
Thallus olive; cortex and medulla C+ pink, KC+ pink or red), PD– (lecanoric acid)
........................................................................................................Hypocenomyce scalaris
33.
Thallus greenish brown to gray-brown; cortex and medulla C–, KC–, PD + red
(fumarprotocetraric acid) ...................................................... Hypocenomyce anthracophila
34.
Ascomata absent. Squamules green with raised or thickened rims...................................35
34.
Ascomata present, either perithecia or apothecia..............................................................36
35.
Squamules less than 2 mm in diameter; on bark, mosses, and lichens (see couplet 32)
.......................................................................................................... Normandina pulchella
35.
Squamules 2-5 mm broad, often lobed, closely appressed to soil or peat; arctic-alpine to
boreal. Fruiting bodies, when present, pale yellow mushrooms….. .Omphalina
hudsoniana
36.
On bark or wood ...............................................................................................................37
36.
On soil, mosses, dead vegetation, or rocks .......................................................................40
37.
Ascomata perithecia buried in the thallus, only the ostioles showing as dark spots at the
surface...............................................................................................................................38
37.
Ascomata reddish-brown apothecia on the thallus surface ...............................................39
38.
Spores muriform, brown, 2 per ascus; squamules thin, closely appressed, relatively flat
56
edges; perithecial cavity containing algae.......................................... Endocarpon pusillum
38.
Spores one-celled, colorless, 8 per ascus; squamules thick, free at the margins, with the
edges turned down; perithecial cavity containing no algae ............. Placidium tuckermanii
39.
Apothecia biatorine, 0.3-0.5 mm in diameter, flat to convex, margin becoming thin and
disappearing in maturity; spores narrowly ellipsoid,1.8-2.2 µm wide..................................
.......................................................................................................... Phyllopsora parvifolia
39.
Apothecia lecanorine, 0.5-3 mm in diameter, flat to concave, margin prominent; spores
ellipsoid, 6.5-10.5 µm wide .................................................................. Psoroma hypnorum
40.(36) Ascomata perithecia buried in the thallus with only the dot-like ostiole
showing......……41
40.
Ascomata apothecia, superficial or sunken into the thallus; or ascomata
absent………...42
41.
Spores muriform, brown, 2 per ascus; perithecial cavity containing algae...........................
........................................................................................................... Endocarpon pusillum
41.
Spores one-celled, colorless, 8 per ascus; perithecial cavity containing no algae ................
............................................................................................................................. Placidium
42.
Apothecia raised on a stalk or stipe ................................................................... Baeomyces
42.
Apothecia not raised on a stalk or stipe ............................................................................43
43.
Apothecia lecanorine or cryptolecanorine (with algae in the margin, or immersed in the
thallus) ..............................................................................................................................44
43.
Apothecia lecideine or biatorine (lacking algae in the margin) ........................................51
44.
Spores up to 100 per ascus................................................................................. Acarospora
44.
Spores 8 or fewer per ascus ..............................................................................................45
45.
Spores 2- or more celled ...................................................................................................46
45.
Spores one-celled ..............................................................................................................48
57
46.
Spores dark brown; growing on calcareous soil, mosses, or dead vegetation; arctic-alpine
............................................................................................................ Phaeorrhiza nimbosa
46.
Spores colorless; growing directly on non-calcareous rock..............................................47
47.
Spore walls evenly thickened; epihymenium K– (anthraquinones absent); in coastal
California .................................................................................................. Lecania brunonis
47.
Spores polarilocular; epihymenium K+ deep purple-red (anthraquinones); in the arid
southwest ............................................................................................. Caloplaca pellodella
48.
Apothecia entirely immersed in thallus, disks flush with thallus; apothecial disks black or
almost black, heavily pruinose; spores globose or broadly ellipsoid, 15-25 μm wide..........
................................................................................................................. Aspicilia contorta
48.
Apothecia sessile or somewhat raised; apothecial disks brown to yellowish orange, not or
lightly pruinose; spores ellipsoid, 4-10 μm wide ..............................................................49
49.
Directly on rocks; thallus shiny ................................................................Lecanora muralis
49.
On soil, mosses, or dead vegetation; thallus dull ..............................................................50
50.
Thallus thick, yellowish white, pruinose; apothecial margin flush with disk, or becoming
thin and disappearing in maturity; on soil in relatively dry habitats .....................................
............................................................................................................Squamarina lentigera
50.
Thallus thin, brown to gray-green; apothecial margin prominent; on mossy soil and peat
in relatively humid habitats................................................................... Psoroma hypnorum
51.(43) Spores 2- or more celled, spore length to width ratio mostly over 3.0 .................... Toninia
51.
Spores one-celled, spore length to width ratio mostly 2.0 or less, or mostly 2.0-3.0 .......52
52.
Squamules closely appressed ............................................................................................53
52.
Squamules lifting at least at the edges ..............................................................................54
53.
On siliceous rock; thallus shiny, with a black prothallus; apothecia lecideine, black;
apothecial margin prominent or flush with disk ..................................Lecidea atrobrunnea
53.
On calcareous soil; thallus dull, without a prothallus; apothecia biatorine, reddish brown
58
or orange-brown; apothecial margin absent……………………...Gypsoplaca
macrophylla
54.
Apothecial disks lead gray, pale brown, or pink.............................. Trapeliopsis wallrothii
54.
Apothecial disks black or almost black, dark brown, or reddish brown to orange-brown....
................................................................................................... Psora key (Psora, Psorula)
KEY H: UMBILICATE AND FAN-SHAPED LICHENS
1.
Thallus attached by a holdfast at one edge of the lobe and becoming fanlike……………2
1.
Thallus attached by a holdfast close to the center of the thallus ........................................ 5
2.
Thallus remaining opaque when wet; with a distinct grass-green algal layer (photobiont a
green alga), lower surface white, yellowish, or dark brown; on bark, or soil.................... 3
2.
Thallus jelly-like and translucent when wet; lacking a distinct algal layer (photobiont
cyanobacteria); lower surface black or gray; on rock. ........................................................4
3.
Thallus yellow-green, rather shiny; lobes with a network of depressions and sharp ridges,
wrinkled or bumpy (rugose), lower surface about the same color as the upper surface,
with a smooth, more or less uniform cortex; on bark ..............................Ramalina sinensis
3.
Thallus dark gray-green when dry and grass-green when wet, lobes smooth and even,
upper surface dull; lower surface distinctly different from the upper surface, entirely
without a cortex, webby or cottony, basically white, with conspicuous dark brown veins;
on soil........................................................................................................ Peltigera venosa
4.
On dry calcareous rocks in the open; lower surface more or less uniform; thallus black.....
.............................................................................................................. Lichinella nigritella
4.
Aquatic, on submerged rocks in mountain streams; lower surface with conspicuous veins;
thallus mineral gray, brown, or olive, at least when dry ....................... Hydrothyria venosa
59
5.(1)
Thallus yellow-green; cortex KC+ orange-yellow (usnic acid) ..........................................6
5.
Thallus gray, brown, or olive, without a yellow tint; cortex KC–, or KC+ red (lacking
usnic acid) ...........................................................................................................................7
6.
Rhizines abundant, stubby and peg-like; thalli large (up to 15 cm across); apothecia with
pale to dark red-brown disks............................................................Omphalora arizonicum
6.
Rhizines absent; thalli up to 6 cm across; apothecia various colors, but not clear redbrown. .................................................................................................................Rhizoplaca
7.
Black dots abundant all over upper surface caused by immersed perithecia ........................
.....................................................................................................................Dermatocarpon
7.
Black dots absent or rare, and then caused by scatted pycnidia; fruiting bodies, when
present, apothecia................................................................................................................8
8.
Apothecia brown, sometimes pruinose, broken up into segments by sterile tissue,
cryptolecanorine, sunken into thallus; thallus thick, surface areolate or cracked, chalky
white to pale gray; spores spherical, 3-4 μm in diameter, many per ascus; rare; on
limestone in the arid interior or arctic………………………………...[Glypholecia
scabra]
8.
Apothecia black, usually with concentric or radiating bands of sterile tissue, lecideine,
superficial or sunken; thallus thin or thick, very dark brown to gray; spores never
spherical, 8 or fewer per ascus; common and widespread, on siliceous rock
...............................................................................Umbilicaria key (Lasallia, Umbilicaria)
KEY I: JELLY LICHENS
1.
Thallus crustose ..................................................................................................................2
1.
Thallus foliose or almost fruticose......................................................................................3
2.
Thallus granular, reddish when wet; photobiont Gloeocapsa ............................. Pyrenopsis
2.
Thallus membranous, olive to black when wet; photobiont
Nostoc………[Lempholemma]
60
3.
Aquatic, on submerged rocks; lower surface with conspicuous veins..................................
.............................................................................................................. Hydrothyria venosa
3.
Not aquatic (i.e., not growing on submerged rocks); lower surface without veins.............4
4.
Lower surface tomentose (hairy or furry)
.................................................................. Leptogium key (Leptogium and Leptochidium)
4.
Lower surface smooth, or wrinkled ....................................................................................5
5.
Thallus pitch black when dry, attached by a single point with the lobes fanning out; on
limestone; photobiont Gloeocapsa ........................................................ Lichinella nigritella
5.
Thallus shades of gray, brown or olive, often dark but not pure black; broadly attached;
on various substrates; photobiont Nostoc ..........................................................................6
6.
Upper and lower surfaces with a cortex, usually a single layer of square to roundish cells;
thallus often shiny, gray to reddish brown ......................................................... Leptogium
6.
Upper and lower surfaces lacking a cortex; thallus always dull; olive to olive- brown or
almost black ........................................................................................................................7
7.
Spores transversely septate or muriform; very common lichens ............................ Collema
7.
Spores 1-celled; uncommon, inconspicuous lichens.................................. [Lempholemma]
KEY J: YELLOW FOLIOSE LICHENS
1.
Thallus orange to yellow-orange; cortex K+ dark purple (anthraquinones) .........................
.............................................................................................................................Xanthoria
1.
Thallus deep yolk-yellow, bright yellow, greenish yellow or yellowish green
("usnic-yellow"), yellowish olive, or yellowish gray; cortex K– or K+ yellow
(anthraquinones absent) ......................................................................................................2
2.
Thallus deep yolk-yellow or bright yellow .........................................................................3
2.
Thallus greenish yellow, yellowish green ("usnic-yellow"), yellowish olive, or yellowish
gray .....................................................................................................................................5
61
3.
Medulla white; calycin present, pinastric and vulpinic acids absent ..................................4
3.
Medulla bright yellow; calycin absent, pinastric and vulpinic acids present……
Vulpicida
4.
Thallus very closely attached, almost crustose (but with a lower cortex), lacking rhizines;
on rock ............................................................................................Candelina submexicana
4.
Thallus clearly foliose, with rhizines; on bark or wood, less frequently on rock
............................................................................................................................ Candelaria
5.(2)
Rhizines absent; lower surface with or without tomentum (see fig. 6)...............................6
5.
Rhizines abundant or sparse; lower surface without tomentum........................................13
6.
Thallus attached by a single central point (umbilicate), or not attached to the substrate
(vagrant); lower surface without tomentum........................................................Rhizoplaca
6.
Thallus not umbilicate or vagrant; lower surface with or without tomentum .....................7
7.
On bark ...............................................................................................................................8
7.
On rock, soil, or moss .......................................................................................................10
8.
Photobiont green; upper surface rather shiny; soredia absent but lobules present; medulla
PD+ orange .................................................................................................Lobaria oregana
8.
Photobiont blue-green; upper surface dull or scabrose; soredia present, lobules absent;
medulla PD– or PD+ yellow ...............................................................................................9
9.
Lobes with a network of depressions and sharp ridges; thallus greenish yellow; lower
surface smooth or wrinkled, dark in the center, pale to dark brown close to the margin;
medulla PD–, K–.................................................................................. Nephroma occultum
9.
Lobes smooth and even, or with rounded depressions; thallus yellowish olive or pale
green; lower surface pale or dark tomentose interrupted by small or large bald spots;
medulla PD+ yellow, K+ yellow to orange (stictic acid with some norstictic acid)
............................................................................................................. Lobaria scrobiculata
62
10.
Thallus of erect, elongated lobes, almost fruticose; lower surface yellow; pycnidia
abundant and conspicuous as black dots mostly along the lobe margins.........Flavocetraria
10.
Thallus prostrate, lobes rounded not erect, closely appressed or loosely attached over
entire surface; lower surface pale to dark brown or black; pycnidia absent or sparse and
very inconspicuous ...........................................................................................................11
11.
On mossy rock in humid, temperate regions; algal layer dark blue green (photobiont
cyanobacteria); soredia present; lower surface with tomentum interrupted by small or
large bald spots ................................................................................... Lobaria scrobiculata
11.
On the ground or over rocks in alpine or arctic regions; algal layer grass-green
(photobiont green algae); soredia absent ..........................................................................12
12.
Lobes smooth and even, or with shallow depressions or wrinkles; cephalodia producing
broad gray bumps on the thallus surface; pseudocyphellae absent; lower surface with a
tomentum; apothecia abundant, produced on the lower surface of lobe margins ……….
..............................................................................................................Nephroma arcticum
12.
Lobes with a network of depressions and sharp ridges; cephalodia absent;
pseudocyphellae usually abundant and conspicuous; lower surface smooth, lacking
tomentum; apothecia rare, on the upper surface of the
lobes………………………………. Asahinea chrysantha
13.
Cilia bulbous, common along lobe margins; small lichens (lobes less than 2 mm wide) of
the subtropics and tropics; rare………………[Relicina] (see comments under
Bulbothrix)
13.
Cilia, if present, not bulbous .............................................................................................14
14.(5)
On bark or wood ...............................................................................................................15
14.
On rock or soil ..................................................................................................................28
15.
Soredia present..................................................................................................................16
15.
Soredia absent ...................................................................................................................22
16.
Rhizines forked in regular dichotomies; soredia on upper surface of lobe tips; medulla
63
PD+ distinct yellow, K+ red (salazinic acid)………………………Hypotyrachyna
sinuosa
16.
Rhizines unbranched; soredia on the lobe margins or on the upper surface; medulla PD–
or PD+ red-orange, K– (salazinic acid absent) ................................................................17
17.
Tiny white dots (pseudocyphellae) present on the upper surface of the lobes; lower
surface dark brown to black; medulla C+ red or pink (lecanoric acid).............................18
17.
Pseudocyphellae absent; lower surface white, brown, or black; medulla C– or C+ red ..19
18.
Soralia almost entirely marginal; pseudocyphellae sparse and inconspicuous .....................
........................................................................................................Flavopunctelia soredica
18.
Soralia both on the thallus surface and along the margins; pseudocyphellae abundant and
conspicuous................................................................................. Flavopunctelia flaventior
19.
Lower surface black with a brown naked zone near margins; rhizines black; lobes 3-8
mm broad; medulla PD– or PD+, KC+ pink or red ........................................................20
19.
Lower surface white to pale brown or dark brown to almost black; rhizines pale or
dark; lobes 0.5-4 mm broad; medulla PD–, KC– ..........................................................21
20.
Soredia coarsely granular, entirely laminal; medulla PD+ red orange, KC+ pink, C–
(protocetraric acid)......................................................................... Flavoparmelia caperata
20.
Soredia both laminal and marginal; medulla PD–, KC+ red, C+ red (lecanoric acid)
........................................................................................................Flavopunctelia soredica
21.
Soredia in round soralia on thallus surface; upper surface dull; rhizines abundant, brown;
medulla UV+ blue-white (divaricatic acid)…………………………Parmeliopsis ambigua
21.
Soredia in elongate soralia along the lobe margins; upper surface rather shiny; rhizines
sparse, white or very pale tan; medulla UV– (divaricatic acid absent)
..............................................................................................Allocetraria oakesiana
22.(15) Lower surface dark brown or black ..................................................................................23
22.
Lower surface white to pale brown or yellow...................................................................26
64
23.
Pseudocyphellae abundant and conspicuous; medulla KC+ red, C+ red (lecanoric acid)
.....................................................................................................Flavopunctelia praesignis
23.
Pseudocyphellae absent; medulla KC–, C– ......................................................................24
24.
Thallus closely attached with abundant rhizines almost to the margin; cilia and isidia
absent; medulla PD+ red, south Texas, New Mexico, and Missouri
.......................................................................................................[Flavoparmelia rutidota]
24.
Thallus loosely attached; rhizines absent from a broad zone at the margin; isidia and cilia
present...............................................................................................................................25
25.
Medulla white; isidia often with short black cilia growing out of the tips; marginal cilia
common and abundant, lower surface smooth; cortex K–, KC+ orange-yellow (usnic acid
present, atranorin and vulpinic acid absent)..........................................................................
........................................................................................................ Parmotrema xanthinum
25.
Medulla yellow; isidia without cilia, marginal cilia sparse; lower surface wrinkled; cortex
K+ yellow, KC–, (atranorin and vulpinic acid present, usnic acid absent)...........................
..................................................................................................... Parmotrema sulphuratum
26.
Medulla bright yellow (pinastric and vulpinic acids); apothecia along the lobe margins
….
...................................................................................................................Vulpicida viridis
26.
Medulla white or pale yellowish orange; apothecia laminal, not marginal.......................27
27.
On southeastern coastal plain; medulla often becoming yellowish to orange in places, K+
yellow, PD+ orange (stictic acid).....................................................Pseudoparmelia uleana
27.
Western or Appalachian-Great Lakes region; medulla white, PD–, K– .................. Ahtiana
28.(14) Soredia present .................................................................................................................29
28.
Soredia absent ...................................................................................................................31
29.
Soredia on thallus surface; lower surface and rhizines black .......................................... 30
29.
Soredia along the lobe margins; lobes elongated; lower surface and rhizines white to pale
65
brown; medulla PD–, KC– ...............................................................Allocetraria oakesiana
30.
Widespread temperate; lobes rounded; medulla PD+ red, KC+ pink (protocetraric acid)
........................................................................................................ Flavoparmelia caperata
30.
Arctic-alpine; lobes narrow, under 1 mm wide, convex, medulla PD–, KC+ red
(alectoronic acid) ........................................................................... [Arctoparmelia incurva]
31.
Hollow pustules resembling isidia present in clumps on the upper surface, sometimes
breaking into granular fragments; medulla PD+ red, KC+ pink, K– (protocetraric acid)
................................................................................................ Flavoparmelia baltimorensis
31.
Hollow, isidia-like pustules absent; solid isidia present or absent; medulla with various
reactions ............................................................................................................................32
32.
Lobes 8-12 mm wide; medulla PD–, K–; marginal cilia common and abundant; rhizines
sparse, absent from a broad zone close to the margin……………..Parmotrema xanthinum
32.
Lobes less than 4 mm wide; marginal cilia absent; rhizines usually abundant to the
thallus edge ...................................................................................................................... 33
33.
Upper surface dull; lower surface white to dull black; medulla K–, PD–, KC+ red
(alectoronic acid); arctic-alpine to boreal…………………………………...Arctoparmelia
33.
Upper surface usually shiny; lower surface brown to pitch black, shiny; medulla usually
PD+ yellow to red, K+ yellow to red; widespread from arctic to southern temperate..........
.................................................................................................................... Xanthoparmelia
KEY K: FOLIOSE LICHENS THAT ARE NOT UMBILICATE, JELLY-LIKE, OR YELLOWISH
1.
Lichens on the ground (soil or mossy turf) .........................................................................2
1.
Lichens directly on bark, wood, or rock ..........................................................................17
2.
Thalli ascending, forming almost erect fruticose tufts; pseudocyphellae conspicuous on
the lower surface of the lobes or branches …………….Cetraria key (Cetraria,
Cetrariella)
2.
Thalli prostrate, not erect ...................................................................................................3
66
3.
Algal layer blue-green (photobiont cyanobacteria).............................................................4
3.
Algal layer green (photobiont green algae).........................................................................8
4.
Lobes less than 2 mm broad; on mossy or bare soil ........................... Massalongia carnosa
4.
Lobes usually more than 2 mm broad ................................................................................5
5.
Lower surface covered by a rather thick, blue-black tomentum; subtropics to eastern
coastal plain ......................................................................................................Coccocarpia
5.
Lower surface with a pale brown to brown-black tomentum, or veined ............................6
6.
Cyphellae present on lower surface; thallus dark brown; rare, arctic-alpine
...................................................................................................................... [Sticta arctica]
6.
Lacking cyphellae on lower surface....................................................................................7
7.
Lobes mostly under 4 mm broad; apothecia embedded in thallus lobes; lichens mainly of
arid localities; spores 1-celled; photobiont Scytonema...........................Heppia conchiloba
7.
Lobes mostly greater than 4 mm broad; apothecia marginal; spores septate; photobiont
Nostoc .................................................................................................................... Peltigera
8.(3)
Lobes mostly less than 3 mm wide .....................................................................................9
8.
Lobes mostly more than 3 mm wide .................................................................................12
9.
Rhizines absent; lobes generally appearing puffed ..........................................................10
9.
Rhizines present, sparse or abundant; lobes solid and flat, not appearing puffed……….
11
10.
Lobes tubelike, hollow.................................................................................... Hypogymnia
10.
Lobes solid...............................................................................................Brodoa oroarctica
11.
Lower surface mostly pale (but dark in the center) with sparse, unbranched rhizines;
lobes 0.2-0.5(-1) mm wide, not pruinose .......................................Phaeophyscia constipata
11.
Lower surface black with a dense mat of squarrose rhizines; lobes 1-3 mm wide, lightly
67
to heavily pruinose ............................................................................ Physconia muscigena
12.
Lower surface jet black, shiny; thallus lacking cephalodia; upper surface K+ yellow
(atranorin); western arctic-boreal......................................................................................13
12.
Lower surface pale tan to black, rarely shiny; cephalodia present as small warts on the
lower or upper surface or as internal patches; upper cortex K– (atranorin absent); arctic
to temperate.......................................................................................................................14
13.
Rhizines sparse but present, especially on older parts of thallus; isidia absent; rare,
Bering Sea coast of Alaska .....................................................................[Cetrelia alaskana]
13.
Rhizines entirely absent; isidia present on thallus surface; Alaska to Hudson Bay
.........................................................................................................[Asahinea scholanderi]
14.
Upper surface of lobes with a network of ridges and depressions; lower surface
uniformly pale brown, sparsely to clearly tomentose ..................................... Lobaria linita
14.
Upper surface of lobes smooth or wrinkled but not forming a network of ridges………15
15.
Cephalodia appearing as gray to brown scales on the upper surface; apothecia marginal
on small lobes ........................................................................................................ Peltigera
15.
Cephalodia hidden within the thallus, on the lower surface, or forming low bumps on the
upper surface.....................................................................................................................16
16.
Thallus lobes strongly crinkled and crisped, ascending; apothecia rare, on the lower side
of small marginal lobes .....................................................................Nephroma expallidum
16.
Thallus more or less smooth, prostrate; apothecia common, immersed in the thallus lobes,
often in depressions ............................................................................................... Solorina
17.(1)
Lobes mostly under 3 mm wide .......................................................................................18
17.
Lobes mostly over 3 mm wide ..........................................................................................87
18.
Lobes inflated and hollow; lower surface lacking rhizines or tomentum .............................
.................................................................... Hypogymnia key (Hypogymnia, Menegazzia)
18.
Lobes solid; rhizines or tomentum present or absent........................................................19
68
19.
Thallus shades of brown, olive or black ...........................................................................20
19.
Thallus shades of gray ......................................................................................................45
20.
Algal layer blue green ..........................................................................................................
................................. Pannaria key (Pannaria, Parmeliella, Massalongia, Vestergrenopsis)
20.
Algal layer green...............................................................................................................21
21.
Rhizines and tomentum absent, or thallus so tightly attached that it is hard to tell ..........22
21.
Rhizines or tomentum present, sparse or abundant...........................................................27
22.
Thallus ascending, very loosely attached at only a few points .........................................23
22.
Thallus appressed, closely attached at numerous points ...................................................26
23.
Lobes strap-shaped, squamule-like, dark olive-green; apothecia spherical; southern
Appalachian Mountains, rare and endangered
............Gymnoderma lineare
23.
Lobes almost fruticose, not flat and strap-shaped; apothecia flat; not from southeast......24
24.
Branches short, mostly up to 15 mm in length; thallus dark greenish brown to black
............................................................................................................ Kaernefeltia merrillii
24.
Branches quite long, over 20 mm in length; thallus brown to brownish gray, without a
greenish tint.......................................................................................................................25
25.
Lobe margins with long, branched, thallus-colored cilia; lower surface white, webby
.............................................................................................................. Anaptychia setifera
25.
Lobe margins often with short, unbranched, black projections but no true cilia; lower
surface brown and shiny like the upper surface ...................... Tuckermannopsis subalpina
26.(22) Lobes convex, appearing inflated but actually solid; on rock ..............................................
............................................ Allantoparmelia key (Allantoparmelia, Brodoa, Lobothallia)
26.
Lobes flat; on bark ......................................................... Hyperphyscia (Phaeophyscia key)
27.(21) Cortex K+ yellow (atranorin); thallus mainly brown on the lobe tips, gray in older parts
69
of the thallus......................................................................................................................28
27.
Cortex K–; thallus uniformly brown, olive or blackish ....................................................29
28.
Pseudocyphellae conspicuous, white, round to elliptical; medulla K–, C+ red (gyrophoric
acid) ...........................................................................................................Punctelia stictica
28.
Pseudocyphellae in a netlike pattern on ridges; medulla K+ red, C– (salazinic acid)
............................................................................................................................... Parmelia
29.
Lobes distinctly pruinose at least at tips; rhizines usually squarrose .................. Physconia
29.
Lobes without pruina ........................................................................................................30
30.
Soredia present..................................................................................................................31
30.
Soredia absent ...................................................................................................................36
31.
Soredia mainly in patches (soralia) on the thallus surface ................................................32
31.
Soredia mainly at the lobe tips or along the margins ........................................................34
32.
Medulla C+ pink or red.........................................................................................Melanelia
32.
Medulla C– .......................................................................................................................33
33.
Rhizines not visible from above; thallus very dark brown, often shiny; spores colorless,1celled
33.
................................................................................................Melanelia
Rhizines often protruding in a fringe around lobes; thallus dark to pale brown or dark
greenish gray, not shiny; spores brown, 2-celled............................................Phaeophyscia
34.
Medulla K+ red or C+ reddish ..............................................................................Melanelia
34.
Medulla K–, C–.................................................................................................................35
35.
Lower surface and rhizines pale brown; rhizines sparse, not extending out from margins
and visible from above; lobes chocolate brown to olive-brown, flat, crisped or undulating
............................................................................................ Tuckermannopsis chlorophylla
35.
Lower surface mostly black; rhizines mostly black (often with white tips), abundant and
extending out from the margins; lobes flat, not undulating or crisped ...........Phaeophyscia
70
36.(30) Isidia present.....................................................................................................................37
36.
Isidia absent ......................................................................................................................39
37.
Isidia mainly marginal, especially on young lobes ...........................................................38
37.
Isidia mainly on the upper surface of the lobes, even on young lobes
............................................................................. Melanelia key (Melanelia, Neofuscelia)
38.
On rock ..............................................................................................Phaeophyscia sciastra
38.
On wood, or sometimes bark ................................................. Tuckermannopsis coralligera
39.
Pycnidia prominent, black, along the lobe margins; pseudocyphellae often conspicuous
especially along the lobe margins .....................................................................................40
39.
Pycnidia absent or immersed in thallus with only a pale or dark ostiole showing, not
largely confined to the margins; pseudocyphellae present or absent, not especially
marginal ...........................................................................................................................41
40.
On bark or wood ..................................................................................... Tuckermannopsis
40.
On rock ................................................................................................ Melanelia hepatizon
41.
Lobes with conspicuous ridges and depressions................................. Kaernefeltia merrillii
41.
Lobes more or less flat, without ridges and depressions...................................................42
42.
Lower surface and rhizines pale brown ............................................................................43
42.
Lower surface and rhizines mostly dark brown or black ..................................................44
43.
On rock in arid habitats; lobules, if present, not strap-shaped; medulla K+ yellow, PD+
orange (stictic acid); spores colorless, 1-celled ; southwestern
U.S. ................................................................................................... Neofuscelia atticoides
43.
On rock or bark in mossy, forest habitats; lobules strap-shaped, marginal; medulla, K–,
PD–; spores brown, 2-celled; eastern North America……………….Anaptychia
palmulata
71
44.
Apothecia very dark brown to black, dull; spores brown, 2-celled; rhizines usually very
abundant, often visible from above as a cilia-like fringe and frequently growing on the
apothecial margins ..........................................................................................Phaeophyscia
44.
Apothecia yellowish brown to reddish brown, shiny; spores colorless, 1-celled; rhizines
abundant or sparse, not forming a fringe visible from above nor growing on the
apothecial margins ................................................................................................Melanelia
45.(19) Photobiont blue-green ....................................................................................Pannaria key
45.
Photobiont green ...............................................................................................................46
46.
Lower surface white, tan, yellow, or orange.....................................................................47
46.
Lower surface dark brown to black (except sometimes near lobe tips) ............................59
47.
Lobes thick and convex, appearing inflated (although solid); rhizines absent; PD+
yellow; on rock .................................. Allantoparmelia key (Allantoparmelia, Lobothallia)
47.
Lobes thin, convex or flat, not appearing inflated; rhizines present or absent; on various
substrates...........................................................................................................................48
48.
Rhizines and tomentum absent .........................................................................................49
48.
Rhizines or tomentum present, sometimes sparse.............................................................55
49.
Thallus very closely attached to substrate over the entire thallus surface (almost crustose
in appearance) ................................................................ Hyperphyscia (Phaeophyscia key)
49.
Thallus loosely attached and ascending ............................................................................50
50.
Marginal cilia absent.........................................................................................................51
50.
Marginal cilia present and conspicuous ............................................................................54
51.
On rock .............................................................................................................................52
51.
On trees .............................................................................................................................53
52.
Lobes convex, 0.1-0.3(-0.5) mm wide; on dry limestone or sandstone, southcentral U.S....
...................................................................................................... Speerschneidera euploca
72
52.
Lobes flat, 0.8-1.3 mm wide, strap-shaped; on wet rock walls in forests of the southern
Appalachian Mountains ......................................................................Gymnoderma lineare
53.
Lobes smooth to wrinkled, often strongly convex; eastern or southern ……Pseudevernia
53.
Lobes generally wrinkled and ridged, flat or concave; humid forests on the west coast
............................................................................................................................ Platismatia
54.(50) Soredia produced on the lower surface of the lobe tips; cortex K+ yellow (atranorin)
..................................................................................................................... Heterodermia leucomela
54.
Soredia absent; cortex K– ..................................................................... Anaptychia setifera
55.(48) Lower surface with a short, pale tomentum, sometimes very sparse and limited to the low
areas between raised bald spots; cephalodia visible on the lower surface as small bumps
that are dark blue-green inside ................................................................................. Lobaria
55.
Lower surface with distinct rhizines; cephalodia absent ..................................................56
56.
Soredia present...................................... Physcia key (Heterodermnia, Physcia, Physciella)
56.
Soredia absent ...................................................................................................................57
57.
Black cilia with bulbous bases fringing the lobe margins ................................. Bulbothrix
57.
Cilia, if present, not black and bulbous.............................................................................58
58.
Thallus cortex PD+ orange, K+ deep yellow (thamnolic acid); apothecia pale brown,
without pruina; spores colorless, 1-celled............................................................ Imshaugia
58.
Thallus cortex PD– or pale yellow, K+ yellow (atranorin); apothecia, if present, dark
brown to black, often pruinose; spores brown, 2-celled .......................................................
...................................................................................Physcia key (Heterodermia, Physcia)
59.(46) Distinct rhizines absent.....................................................................................................60
59.
Distinct rhizines present....................................................................................................65
60.
Lower surface with a spongy black hypothallus consisting of intricately interconnected
fibers (fig. 6g,h) ...........................................................................................Anzia colpodes
73
60.
Lower surface naked, without a spongy hypothallus ........................................................61
61.
Lower surface pitted with many tiny perforations; thallus small, usually forming rosettes
less than 2.5 cm across..................................................................................... Cavernularia
61.
Lower surface not perforated or pitted; thalli larger than 3 cm long or broad…………..62
62.
On alpine or arctic rocks; lobes thick, convex ......................................................................
..................................................................Allantoparmelia key (Allantoparmelia, Brodoa)
62.
On bark or wood ...............................................................................................................63
63.
Closely appressed over most of the thallus surface .................. Dirinaria (see Physcia key)
63.
Very loosely attached by relatively few points .................................................................64
64.
Lobes flat to concave, often wrinkled; pycnidia along the lobe margins; medulla C–; in
humid or boreal forests along the west coast ...................................................... Platismatia
64.
Lobes convex; pycnidia buried in lobe tips, not along the margins; medulla C+ red
(lecanoric acid); montane and interior sites .................................................... Pseudevernia
65.(59) Pseudocyphellae present on lobe surface .........................................................................66
65.
Pseudocyphellae absent ....................................................................................................67
66.
Pseudocyphellae dotlike, most easily seen on young lobes; medulla K–
…………Punctelia
66.
Pseudocyphellae netlike on reticulate ridges; most species with medulla K+ red (salazinic
acid) ...................................................................................................................... Parmelia
67.
Cilia present on lobe margins or in the axils of the lobes (sometimes
sparse!)………….68
67.
Cilia absent........................................................................................................................72
68.
True cilia arising from the margins or axils of the lobes ......................................................
.....................Parmelia key (Bulbothrix, Myelochroa, Parmelina, Parmelinopsis, Relicina)
68.
"Cilia" actually a fringe of rhizines extending beyond the lobe margins, not arising from
74
the lobe margins themselves .............................................................................................69
69.
Rhizines long and forked, uniformly black........................... Hypotrachyna (Parmelia key)
69.
Rhizines unbranched or squarrose, or frayed to brushlike at the tips, often with white tips70
70.
Cortex K+ yellow..............................................................................................................71
70.
Cortex K–........................................................................................................Phaeophyscia
71.
Lobe margins abundantly squamulose ........................................ Heterodermia squamulosa
71.
Lobe tips sorediate, not squamulose ........................................ [Heterodermia casarettiana]
72.(67) Rhizines forked in regular dichotomies; thallus without pruina. ................... Hypotrachyna
72.
Rhizines unbranched, squarrose, or brushlike, rarely forked; thallus with or without
pruina ....................................................................................................................................
..........................................................................................................................................73
73.
Thallus very loosely attached, strongly wrinkled, especially on the lower surface; on
conifers in the west ....................................................................... Esslingeriana idahoensis
73.
Thallus closely attached over most of its surface; smooth, or only slightly wrinkled, on
both surfaces .....................................................................................................................74
74.
Cortex K– (without atranorin); thallus usually dark greenish gray...................................75
74.
Cortex K+ yellow (atranorin); thallus usually pale gray...................................................77
75.
Thallus without pruina ....................................................................................Phaeophyscia
75.
Thallus pruinose at least at the lobe tips ...........................................................................76
76.
Rhizines squarrose (like bottle brushes); mostly northern ................................... Physconia
76.
Rhizines unbranched or rarely forked; northern to subtropical ............................................
......................................................................................................Pyxine (see Physcia key)
77.(74) Thallus pruinose at least at lobe tips.................................... Physcia key (Physcia, Pyxine)
77.
Thallus without pruina ......................................................................................................78
75
78.
Medulla pale yellow, at least close to the algal layer................Myelochroa (Parmelia key)
[Rare nonpruinose specimens of Pyxine eschweileri or adnate Esslingeriana will key out
here]
78.
Medulla white ...................................................................................................................79
79.
Soredia absent; squamulose lobules abundant on lobe margins ...........................................
.................................................................................................... Heterodermia squamulosa
79.
Soredia present..................................................................................................................80
80.
Soredia on or close to the lobe margins; medulla C– .......................................................81
80.
Soredia laminal or on the lobe tips....................................................................................82
81.
Medulla PD–, K+ yellow; rhizines sparse and distinct ............................ Physcia sorediosa
81.
Medulla PD+ orange, K–; rhizines abundant, forming an intricate mat ...............................
.............................................................................Pyxine eschweileri (nonpruinose morph)
82.
Lobes tips often curled into tubes; medulla C+ red (gyrophoric acid)..................................
......................................................................................................... Hypotrachyna revoluta
82.
Lobes more or less flat; medulla C– .................................................................................83
83.
Lobes 0.5-2 mm wide .......................................................................................................84
83.
Lobes 2-4 mm wide ..........................................................................................................86
84.
On rocks; medulla PD+ red-orange (protocetraric acid); southeastern U.S.
..................................................................................................[Paraparmelia alabamensis]
84.
On bark or wood; medulla PD– or PD+ orange................................................................85
85.
Rhizines black; southeastern coastal plain............................................. Pyxine eschweileri
85.
Rhizines pale tan to brown, not black; western and northern……...Parmeliopsis
hyperopta
86.
Medulla UV+ white (divaricatic acid); lower surface and rhizines dark brown ...................
76
............................................................................................................Canoparmelia texana
86.
Medulla UV-; lower surface and rhizines black ...................................................................
.............................................Myelochroa aurulenta (rare specimens with a white medulla)
87.(17) Lobes inflated and hollow ...............................................................................Hypogymnia
87.
Lobes solid........................................................................................................................88
88.
Thallus brown, brownish green, olive, or black when dry................................................89
88.
Thallus pale gray to greenish gray when dry ..................................................................103
89.
Algal layer blue-green.......................................................................................................90
89.
Algal layer green (with a lower, secondary, blue-green layer in Solorina) ......................95
90.
Lower surface smooth or with a short tomentum, usually pale brown (or gray to bluish in
Pannaria), lacking veins; mostly with a lower cortex; without discrete rhizines ..............91
90.
Lower surface cottony or webby, often with branching or interconnecting veins, lacking
a lower cortex; white to brown or black, with discrete or tufted rhizines.............................
................................................................. Peltigera key (Peltigera, Erioderma, Leioderma)
91.
Small round holes or pits (cyphellae) on the lower surface (plate 15)........................ Sticta
91.
Cyphellae absent ...............................................................................................................92
92.
Tiny, white or yellow, raised spots (pseudocyphellae) on the lower surface (plate 14)
................................................................................................................ Pseudocyphellaria
92.
Pseudocyphellae absent ....................................................................................................93
93.
Tomentum on lower surface mostly gray to bluish; apothecia on upper surface of lobes
................................................................................................................................Pannaria
93.
Tomentum on lower surface, or lower surface itself, pale brown.................................... 94
94.
Thallus olive-gray to brownish gray sorediate; apothecia infrequent, laminal
................................................................................................................................ Lobaria
94.
Thallus brown, sorediate or not; apothecia frequent, on lower surface of lobe margins;
77
medulla PD–.........................................................................................................Nephroma
95.(89) Lower surface tomentose or veined at least in part; cephalodia present (warts or
secondary algal layers containing cyanobacteria).............................................................96
95.
Lower surface smooth or wrinkled, not at all tomentose; cephalodia
absent…………….98
96.
Lower surface more or less veined or webby, without a cortex; apothecia immersed in
thallus, often in depressions.................................................................................... Solorina
96.
Lower surface uniform in color, pale brown, partly or entirely tomentose, often with
raised, smooth, naked areas, with a cortex; apothecia superficial or
raised……………...97
97.
Pseudocyphellae (yellow or white raised spots) on lower surface of lobes ..........................
................................................................................................................ Pseudocyphellaria
97.
Pseudocyphellae lacking on lower or upper surface ................................................ Lobaria
98.
Thallus loosely attached, ascending..................................................................................99
98.
Thallus more or less closely appressed to substrate, except at the lobe tips. ..................100
99.
Thallus greenish black or dark olive-brown; apothecia black or very dark brown;
pycnidia immersed in thallus; pseudocyphellae very inconspicuous or absentKaernefeltia merrillii
99.
Thallus brown to olive brown; apothecia red-brown; pycnidia black, prominent;
pseudocyphellae often conspicuous ......................................................... Tuckermannopsis
100.
Lobe margins fringed with black rhizines extending out from below and appearing like
cilia; rhizines forming an interwoven mat below………………….Phaeophyscia
hispidula
100.
Lobe margins without cilia-like rhizines; rhizines separate and distinct ........................101
101.
Lower surface and rhizines black; upper surface usually gray in part, K+ yellow (test the
parts that remain gray); surface of lobes usually with a network of ridges and
depressions, appearing like hammered metal..................................................................102
78
101.
Lower surface and rhizines pale to dark brown (rhizines rarely black); upper surface
uniformly brown or olive, K–; lobes smooth or rough, but rarely with a network of ridges
................................................ Melanelia key (Melanelia, Neofuscelia, Tuckermannopsis)
102.
Pseudocyphellae milk white, very conspicuous, round to elongated; medulla K–, C+ red
(gyrophoric acid)........................................................................................Punctelia stictica
102.
Pseudocyphellae pale, netlike or irregular in shape, occasionally round; medulla K+ red,
C– (salazinic acid).................................................................................................. Parmelia
103.(88) Lower surface tomentose, cottony, or veined, at least in part, rarely naked;
discrete rhizines sometimes present; photobiont blue-green or green ...........................104
103.
Lower surface smooth, wrinkled, or rough, but not tomentose or cottony; discrete
rhizines usually well developed; photobiont green.........................................................112
104.
Pseudocyphellae present on the upper or lower thallus surface......................................105
104.
Pseudocyphellae absent ..................................................................................................106
105.
Pseudocyphellae on the upper surface of the lobes; medulla C+red.....................................
............................................................................................................ Punctelia subrudecta
105.
Pseudocyphellae on the lower surface of the lobes; medulla C– ............ Pseudocyphellaria
106.
Soredia present................................................................................................................107
106.
Soredia absent .................................................................................................................109
107.
Lobes 2-5 mm wide; soredia marginal, coarsely granular; medulla PD+ orange (pannarin)
.............................................................................................................. Pannaria conoplaea
107.
Lobes 5-20 mm wide ......................................................................................................108
108.
Lower surface vaguely veined, cottony (not tomentose); soredia mostly marginal
.....................................................................................................Peltigera collina
108.
Lower surface uniformly brown, tomentose (or with scattered naked areas); soredia on
both the margins and upper surface of the lobes...................................................... Lobaria
79
109.
Algal layer blue-green.....................................................................................................110
109.
Algal layer green.............................................................................................................111
110.
Lower surface with a thick or thin gray to blue-black tomentum
................................................................................. Pannaria key (Coccocarpia, Pannaria)
110.
Lower surface with distinct or indistinct veins ...................................................... Peltigera
111.
Cephalodia in the form of brown or gray warts or lobed squamules on the upper surface
of the thallus; lower surface webby or cottony, white at the lobe edge and dark brown to
black in the center, sometimes with veins.............................................................. Peltigera
111.
Cephalodia in the form of small round warts or galls on the lower surface of the thallus;
lower surface without veins, abundantly or sparsely
tomentose……………………Lobaria
112.(103) Lower surface entirely white or very pale brown; rhizines white or pale brown .........113
112.
Lower surface pale to dark brown or black, sometimes blotched with white over small or
large areas, but in such cases, always brown to black in the oldest, central area; rhizines
black or brown ................................................................................................................114
113.
White dots (pseudocyphellae) present on upper surface of lobes ..........................Punctelia
113.
Pseudocyphellae absent ...............................................................................Physcia biziana
114.
Upper cortex K–, UV-; lobes fringed with black cilia-like rhizines .....................................
........................................................................................................ Phaeophyscia hispidula
114.
Upper cortex K+ yellow, UV-, or rarely K–, UV+ yellow; with or without cilia or cilialike rhizines.....................................................................................................................115
115.
Apothecia common, black; spores brown, 2-celled; closely adnate tropical species on
bark
..................................................................................................................Dirinaria confusa
115.
Apothecia, when present, brown; spores colorless, 1-celled ...........................Parmelia key
80
Identification Keys to Species
ACAROSPORA (including Pleopsidium)
1.
Thallus distinctly yellow .................................................................................................... 2
1.
Thallus brown or gray, without a yellowish or orange tint ................................................ 4
2.
Growing on soil................................................................................ Acarospora schleicheri
2.
Growing on rocks............................................................................................................... 3
3.
Thallus areolate, dispersed or contiguous, not conspicuously lobed at the margin; asci not
thickened at the apex, K/I- ................................................................. Acaropsora contigua
3.
Thallus somewhat lobed at the margin, otherwise areolate; asci with a distinctly thickened,
partly K/I+ blue tholus ......................................................................... Pleopsidium flavum
4.
Thallus brown, not pruinose; thallus cortex usually C+ pink, KC+ pink but sometimes hard to
demonstrate (gyrophoric acid present); on non-calcareous rock............Acarospora fuscata
4.
Thallus gray to gray-brown, lightly to heavily pruinose; thallus cortex C-, KC- (gyrophoric
acid absent); on calcareous rocks ...................................................................................... 5
5.
Thallus usually reduced to narrow, reddish brown, sometimes pruinose margins around the
apothecia, which become Lecanora-like; apothecial disks lightly pruinose
........................................................................................................ Acarospora glaucocarpa
5.
Thallus well-developed, areolate; one to several apothecia immersed in each areole, never
Lecanora-like ........................................................................................ Acaropsora strigata
AHTIANA
1.
Thallus pruinose, at least at lobe tips; lower surface pale yellow, wrinkled ……Ahtiana pallidula
1.
Thallus entirely without pruina; lower surface white to pale brown, smooth……………………2
2.
Thallus closely appressed; lower surface white or almost white; western montane.............
....................................................................................................... Ahtiana sphaerosporella
2.
Thallus loosely attached; lower surface pale brown; in Appalachian-Great Lakes region...
................................................................................................................. Ahtiana aurescens
ALECTORIA (including Bryocaulon, Bryoria, Nodobryoria, and Sulcaria)
1.
Thallus pale greenish yellow or yellowish green; cortex KC+ gold (usnic acid); pseudocyphellae
conspicuous, usually slightly raised ................................................................................... 2
78
1.
Thallus gray, olive, or shades of brown to almost black; if yellowish gray, then cortex KC+ pink,
not gold; pseudocyphellae conspicuous or inconspicuous, level with the surface or slightly to
deeply depressed ................................................................................................................ 6
2.
Thallus forming erect clumps, or sometimes prostrate, on the ground; tips of branches usually
becoming black or greenish black ....................................................... Alectoria ochroleuca
2.
Thallus shrubby to pendent, on trees or shrubs, rarely with blackened branch tips ........... 3
3.
Thallus forming bushy clumps, usually less than 10 cm long............................................ 4
3.
Thallus pendent to slightly pendent, usually 8-20 cm long when mature .......................... 5
4.
Thorny isidia and spinules developing in elongate pseudocyphellae and fissures; apothecia very
rare; cortex K+ bright yellow, PD+ orange (thamnolic acid) or K–, PD– (squamatic acid); medulla
KC–………………………………….Alectoria imshaugii
4.
Isidia and spinules absent; apothecia usually present, brown; cortex K–, PD–; medulla KC+ red
(alectoronic acid).............................................................................................Alectoria lata
5.
Branches very slender and pale, with tips curled up and granular or sorediate; medulla KC–
..................................................................................................................Ramalina thrausta
5.
Branches slender or thick, not curled up at the tips; without soredia; medulla KC+ red
(alectoronic acid), or infrequently KC– (usnic acid alone)…………….Alectoria sarmentosa
6.(1)
Thallus forming erect or prostrate clumps on soil, heath, or sometimes rock (rarely on shrubs)
............................................................................................................................................ 7
6.
Thallus bushy to pendent, on trees and shrubs, rarely on rock......................................... 10
7.
Thallus brown with a distinct reddish tint .......................................................................... 8
7.
Thallus black to very dark brown, not especially reddish .................................................. 9
8.
Pseudocyphellae dot-like, level to slightly raised, white, C+ red (olivetoric acid) ..............
........................................................................................................... Bryocaulon divergens
8.
Pseudocyphellae round to elliptical, depressed, white, C–…………………Cetraria aculeata
9.
Thallus dull, gray to yellowish gray at the base with blackened branch tips, occasionally entirely
black; pseudocyphellae raised, white; cortex PD+ yellow, KC+ pink (use the filter paper test)
................................................................................................................ Alectoria nigricans
9.
Thallus mostly shiny, uniformly dark brown to black; pseudocyphellae level with the surface,
79
brown and very inconspicuous (seen as dull, fusiform areas); outer cortex PD–; medulla and inner
cortex PD+ red (fumarprotocetraric acid; diffusing onto filter paper only where the branch is
broken)………….………………………………………….Bryoria nitidula
10.(6)
Thallus forming rounded or irregular bushy tufts or clumps with divergent branching... 11
10.
Thallus pendent or almost pendent when mature ............................................................. 14
11.
Branches without soredia or isidia ................................................................................... 12
11.
Branches with scattered soralia ........................................................................................ 13
12.
Thallus red-brown; apothecia shiny red-brown, decorated with spiny cilia on the margins;
epihymenium K–.............................................................................Nodobryoria abbreviata
12.
Thallus brown at base, greenish black at tips; apothecia greenish black; epihymenium K+ purple
.........................................................................................................Kaernefeltia californica
13.
Thallus shiny, uniformly brown, with fissural soralia that contain tiny, thornlike isidia; outer
cortex and medulla PD+ red, K–, KC– (fumarprotocetraric acid) ........... Bryoria furcellata
13.
Thallus dull, gray to pale or dark brown, with round, warty soralia that contain no isidia; cortex
PD+ orange-yellow, K+ bright yellow, KC+ pink (alectorialic acid) ..................................
...........................................................................................................Bryoria nadvornikiana
14.(10) Branches with soralia ....................................................................................................... 15
14.
Branches without soralia .................................................................................................. 19
15.
Soredia bright yellow; thallus with some stout main branches and slender, perpendicular side
branches; medulla and soralia PD– .................................................................................. 16
15.
Soredia white (sometimes flecked with black)................................................................. 17
16.
Pseudocyphellae yellow, twisting around branches………Bryoria tortuosa (rare sorediate form)
16.
Pseudocyphellae absent (although vague white cracks sometimes look like pseudocyphellae)
.................................................................................................................. Bryoria fremontii
17.
Pseudocyphellae present, rather long, but sometimes hard to see; cortex PD+ orange-yellow, K+
yellow, KC+ pink (alectorialic acid); main branches with short or long, perpendicular side
branches ...........................................................................................Bryoria nadvornikiana
17.
Pseudocyphellae absent; cortex PD– or PD+ red; soredia PD+ red (fumarprotocetraric acid);
perpendicular side branches present or absent ................................................................. 18
80
18.
Thallus very dark brown, almost black; branches very slender (less than 0.2 mm in diameter),
brittle; soralia typically abundant, often broader than the branch; cortex PD–
.................................................................................................................... Bryoria lanestris
18.
Thallus dark to medium brown to olive; branches 0.2-0.6 mm in diameter, not very brittle;
soralia relatively sparse, usually not broader than the branch; cortex PD+ red or PD–
........................................................................................................................................ 18a
18a.
Thallus dark to medium brown, typically dull, usually paler at the base; angles between the
branches (axils) acute, not rounded; soralia tuberculate or fissural; cortex PD+ red or PD.............................................................................................................. Bryoria fuscescens
18a.
Thallus olive to olive-brown, shiny, not paler at base; branch axils broad and rounded;
soralia always fissural; cortex PD-........................................................... Bryoria glabra
19.(14) Branches with deep, longitudinal grooves (special pseudocyphellae), often quite long; branches
commonly twisted; cortex K+ yellow, PD+ yellowish or brownish, KC– or KC+ yellow
(atranorin); very rare, California and Oregon.................................................Sulcaria badia
19.
Branches without deep grooves, with or without pseudocyphellae.................................. 20
20.
Pseudocyphellae absent; medulla PD–............................................................................. 21
20.
Pseudocyphellae present, usually conspicuous; medulla PD+ red or PD–………………22
21.
Thallus uniformly red-brown, with slender, brittle branches 0.1-0.2 mm in diameter; apothecia
fairly common, with red-brown, nonpruinose disks; cortex with jigsaw puzzle-shaped cells as
viewed at 100-400X magnification………………………….Nodobryoria oregana
21.
Thallus dark reddish brown to yellowish brown; thallus with stout, twisted and dented main
branches 0.4-1.5(-4) mm in diameter, and slender, perpendicular side branches; apothecia
uncommon, with yellow-pruinose disks; cortex with elongate cells ........ Bryoria fremontii
22.
Pseudocyphellae yellow, twisting around the branch in spirals; thallus reddish brown or often
yellowish (vulpinic acid)............................................................................ Bryoria tortuosa
22.
Pseudocyphellae white or pale brown, dot-like, fissural, or twisting; thallus never yellowish
.......................................................................................................................................... 23
23.
Thallus not brittle, pale to dark brown or red-brown; outer cortex PD– .......................... 24
23.
Thallus usually brittle, pale gray to gray brown or dark brown; outer cortex and sometimes
medulla PD+ yellow......................................................................................................... 25
81
24.
Thallus brown, without a reddish tint; pseudocyphellae elongate (fusiform or linear); inner cortex
and medulla usually PD+ red (fumarprotocetraric acid; seen through the cortex rendered
transparent in the filter paper test), C–; common ......................................Bryoria trichodes
24.
Thallus distinctly red-brown; pseudocyphellae dot-like, slightly raised; inner cortex and medulla
PD–, C+ red (olivetoric acid); rare, coastal Pacific Northwest…..[Bryocaulon pseudosatoanum]
25.
Cortex K+ bright yellow, KC+ red (alectorialic acid); thallus usually pale gray to brownish gray;
mostly in coastal or humid localities.........................................................Bryoria capillaris
25.
Cortex K+ brownish, KC– (norstictic acid); thallus usually dark brown, less frequently pale;
mostly in inland, montane forests………………………………….Bryoria pseudofuscescens
ALLANTOPARMELIA (including Brodoa and Lobothallia)
1.
Lobes long and divergent, not contiguous near the growing margin, gray to dark brown; upper
cortex K+ yellow on pale areas (atranorin); medulla PD– or rarely PD+ red
..................................................................................................................Brodoa oroarctica
1.
Lobes long or short, contiguous at the margin of the thallus, not divergent; cortex K– or K+ yellow
to red (atranorin absent); medulla PD+ yellow .................................................................. 2
2.
Cortex K–; medulla K–, KC+ red (alectorialic acid); apothecia always superficial, not immersed,
up to 7 mm across; spores 7.5-10 x 5-7 μm ……………………..Allantoparmelia alpicola
2.
Cortex and medulla K+red, KC– (norstictic acid); apothecia at first immersed, later superficial,
under 2.5 mm across; spores 10-14 x 6-10 μm................................................................... 3
3.
Thallus gray to gray-brown ..............................................................Lobothallia alphoplaca
3.
Thallus yellow brown to copper brown...........................................Lobothallia praeradiosa
ALLOCETRARIA: See Keys A and J, and Dactylina.
AMANDINEA: See Buellia and Rinodina.
AMYGDALARIA: See Keys C and F.
ANAPTYCHIA: See Keys A and K.
ANZIA: See Key K.
82
ARCTOPARMELIA: See Xanthoparmelia.
ARTHONIA (including Arthothelium)
1.
Spores muriform; ascomata lobed to starlike or almost round ........................................... 2
1.
Spores only transversely septate; ascomata round to scriptlike.......................................... 3
2.
Spores (15-)17-24(-26) x 7-9.5(-10.5) μm ...................................... [Arthothelium ruanum]
2.
Spores 26-36 x 12-15 μm .............................................................[Arthothelium spectabile]
3.
Ascomata round, with the appearance of a biatorine apothecium ...................................... 4
3.
Ascomata elongate or branched lirellae ............................................................................. 6
4.
On maritime rocks. Thallus brown, membranous; spores 3- to 6-celled, 15-23 x 5.5-8 μm; on both
Atlantic and Pacific coasts ................................................................. [Arthonia phaeobaea]
4.
On bark............................................................................................................................... 5
5.
Ascomata convex, bluish gray due to a heavy white pruina; spores 4-celled; on a variety of trees in
the northeast .................................................................................................Arthonia caesia
5.
Ascomata flat to slightly convex, black, not at all pruinose; spores 2-celled, strongly tapered, (8)10-12 x 3.5-5 μm; almost exclusively on trembling aspen trees, northern Temperate to southern
Boreal................................................................................................. [Arthonia patellulata]
6.(3)
Spores with 4 equal-sized cells .................................................................. Arthonia radiata
6.
Spores (4-)5- to 8-celled, tapered, with at least one end cell larger than the others ........... 7
7.
Ascomata ellipsoid to Y-shaped, convex, red pruinose on the margins or over the entire surface
(anthraquinones); spores 18-28 x 7-10 μm………………………….Arthonia cinnabarina
7.
Ascomata irregularly branched, flat or immersed, not pruinose (anthraquinones absent);
spores 26-35 x 10-15 μm.................................................................................................... 8
8.
Ascomata black, flat; spores tapered, with uppermost cell considerable larger than lower cells;
coastal Pacific Northwest ..........................................................................[Arthonia ilicina]
8.
Ascomata brown to reddish brown, usually immersed and narrowly or widely open, appearing like
irregular cracks in the bark; spores only slightly tapered if at all, both end cells larger than middle
cells; southeast .........................................................................................[Arthonia rubella]
ARTHRORHAPHIS: See Key G
83
ASAHINEA: See Keys J and K.
ASPICILIA
1.
Thallus fruticose, with clumps of tangled, terete branches; white depressed pseudocyphellae
abundant and conspicuous; apothecia rare………………………………..Aspicilia hispida
1.
Thallus crustose, with or without pseudocyphellae; apothecia abundant........................... 2
2.
Thallus areolate with areoles dispersed or contiguous, strongly convex or pyramidal, dark olive to
dark olive-gray, often with white pseudocyphellae; spores almost spherical, 15-25 μm wide
.................................................................................................................. Aspicilia contorta
2.
Thallus usually continuous, smooth or rimose-areolate, white to gray, rarely olive; lacking
pseudocyphellae; spores ellipsoid, 6-16 μm wide .............................................................. 3
3.
Thallus chalky white, often lobed at the periphery; on calcareous rock; all spot tests negative
...................................................................................................................Aspicilia candida
3.
Thallus creamy white or pale to dark gray, rarely chalky white, never lobed; on non-calcareous
rocks................................................................................................................................... 4
4.
Paraphyses septate and constricted at the tips, resembling a string of beads; asci K/I–; medulla K+
red or yellow, PD+ yellow or orange (norstictic acid or, occasionally, stictic acid) ...........
....................................................................................................................Aspicilia cinerea
4.
Paraphyses expanded at the tips, not constricted or beadlike; asci with a K/I + blue tip; medulla
PD–, K–................................................................................................ Lecanora oreinoides
ASTROTHELIUM: See Trypethelium.
BACIDIA (including Bacidina and Scoliciosporum)
1.
On rocks and occasionally wood, rarely tree bases ............................................................ 2
1.
On bark or mosses, very rarely on rock or wood ............................................................... 4
2.
In periodically submerged habitats such as streams and lake shores; epihymenium brown; thallus
greenish, usually with a white prothallus. Spores needle-shaped, 4(-8) celled, 24-43 x 2-3 μm;
apothecia black when wet, brown when dry; probably widespread temperate to
boreal…………………………………………………………………..[Bacidina inundata]
2.
In dry habitats; epihymenium greenish or brownish; thallus without a prothallus............. 3
3.
Spores needle-shaped, spirally twisted, (15-)20-30(-40) x 2-3 μm, 4- to 8-celled. Apothecia red-
84
brown to black; widespread temperate, especially in east……………[Scoliciosporum umbrinum]
3.
Spores fusiform, not spirally twisted, 5-8 μm wide, 4-to 6-celled……..
................................................................................ Myxobilimbia sabuletorum (syn. “Bacidia” sabuletorum)
4.
Apothecial disks black or almost black; epihymenium brown to green ............................. 5
4.
Apothecial disks pale brown, or reddish brown to orange-brown; epihymenium yellowish or
colorless ............................................................................................................................. 8
5.
Hypothecium dark orange-brown to red-brown ................................................................. 6
5.
Hypothecium colorless....................................................................................................... 7
6.
On moss on tree bases, rarely directly on bark; apothecia strongly convex to hemispherical; spores
fusiform, under 40 μm long, 4- to 6-celled……………Myxobilimbia (“Bacidia”) sabuletorum
6.
On bark; apothecia mostly flat; spores needle-shaped, mostly over 40 μm long, mostly 4- to 9celled .................................................................................................... Bacidia schweinitzii
7.
Spores rod- or club-shaped, mostly straight, 4- to 8-celled; thallus gray-green to pale gray,
areolate; apothecia mostly remaining flat with a persistent margin; spores 11-37(-45) x 1.6-3.7
μm; Pan-Temperate at low elevations……………………………….[Bacidia circumspecta]
7.
Spores comet-shaped, usually curved and tapering, 5- to 8-celled; thallus dark green to brownish
green, granular; apothecia convex and soon marginless; spores 3-5 μm wide
.............................................................................................. Scoliciosporum chlorococcum
8.(3)
Hypothecium and exciple below the hypothecium dark orange-brown or red-brown. Brown
pigments in apothecial tissues K+ purple-red; apothecia commonly pruinose at least on margins
when young; spores 4- to 12-celled, 31-74 x 2-5 μm; East Temperate…….[Bacidia polychroa]
8.
Hypothecium and exciple below the hypothecium colorless, pale yellowish, or pale brown………9
9.
Thallus consisting of large, round granules; apothecia orange-brown, sometimes pruinose on the
margins when young; all apothecial tissues negative or colors intensifying with K ............
...................................................................................................................... Bacidia rubella
9.
Thallus thin or thick, continuous or cracked, not consisting of large round granules; apothecia
pruinose or not; brown pigments in apothecial tissues K+ purple.................................... 10
10.
Exciple with radiating clusters of crystals; apothecia yellow-brown to purplish brown, rarely redbrown, usually pruinose in part; outermost 4-8 cell layers of exciple very large and distinct from
inner cells, which are much narrower; East Temperate............................. [Bacidia suffusa]
85
10.
Exciple normally without crystals; apothecia mainly orange-brown, not pruinose or pruinose; only
outermost 1-2 cell layers of exciple have enlarged cells .................................................. 11
11.
Brown pigment of the epihymenium deposited as distinct caps on the tips of the paraphyses (seen
best when the hymenium is squashed in K); spores 4- to 16-celled, 32-67(-73) x 2.5-4.5 μm; on
deciduous trees and shrubs; southeastern coastal plain and along the Pacific coast
………………………………………………………………………………[Bacidia heterochroa]
11.
Brown pigment of the epihymenium distributed uniformly in the upper hymenial jelly; spores 8- to
29-celled, (50-)57-96(-108) x 2-3.7 μm; on conifers and deciduous trees; Great Lakes to New
England, and Pacific Northwest .......................................................... [Bacidia laurocerasi]
BAEOMYCES (including Dibaeis)
1.
Apothecia pink; thallus almost white; medulla PD+ bright yellow, K– or K+ yellowish
(baeomycesic and squamatic acids) .......................................................Dibaeis baeomyces
1.
Apothecia brown; thallus pale green to gray green or brownish; medulla PD+ orange, K+ yellow
(stictic acid)........................................................................................................................ 2
2.
Thallus thin, edge indefinite or definite, not lobed at the margin……………….Baeomyces rufus
2.
Thallus thick, distinctly lobed at the margin .................................Baeomyces placophyllus
BELLEMEREA: See Key F.
BIATORA from eastern North America (including Cliostomum, Catillaria, Micarea and Mycobilimbia)
[New key for 3rd Edition]
[Based on “Preliminary key to Biatora species known to occur in Eastern North America” , C. Printzen & T. Tønsberg,
Symb. Bot. Ups. 34:1. 2004. New and interesting Biatora species. For identifying Biatora specimens from the west, see
C. Printzen & T. Tønsberg, The lichen genus Biatora in northwestern North America, Bryologist 102: 692-713. 1999; and
C. Printzen, Biatora, in Lichens of the Sonoran Desert Region, vol. 2, 2005.]
1. Thallus sorediate, often sterile .......................................................................…………….......................2
1. Thallus esorediate, fertile .………….........................................................................................................8
2. Spores one-celled or a few 2-celled .....….............................................……………................................ 3
2. Spores mostly 2- or more cells ..........................................................…………...................….............. 6
3. Thallus and soralia C+ orange, xanthones present, soralia usually pale green, flat to weakly convex,
punctiform, becoming confluent; apothecia greyish yellow to dark brownish grey, often with a dark
blue tinge. Southern Appalachians …………………………………………………….….... [B. pontica]
86
3. Thallus and soralia C-, but xanthones may be present; soralia yellowish green, convex and
well-defined, punctiform, remaining discrete; apothecia pale to dark brown; North Temperate
……………………………………………………………………………………. [Biatora sp.#4]
3!. Thallus and soralia C+ rose-red, gyrophoric acid present, soralia mostly thick, convex and elevated
above thallus level, apothecia without dark blue tinge ……………..……………………………...…...4
4. Thallus and soralia PD+ orange red, argopsin present..........................…………................[B. printzenii]
4. Thallus and soralia PD–, without argopsin ...........................................……………................................5
5.
Soralia mostly rounded and well delimited, apothecial sections C+ rose-red, spores 9.0–11.0 × 3.5–3.6
µm; southern Appalachians ….………………...................……………………...... [B. appalachensis]
5. Soralia mostly confluent, usually poorly delimited, apothecial sections C–, spores 11.9–15.6
× 3.8–5.9 µm; Great Lakes Region ……………………………………………............... [B. chrysantha]
6. Spores 4(-6) celled; apothecia red-brown to pale brown; pycnidia rare, pale; on bryophytes or mossy
bark; mostly boreal, Yukon to Maritimes.................…………………….. [Mycobilimbia epixanthoides]
6. Spores mainly 2-celled ……………………………………………………………………………….. 7
7. Apothecia pale yellow (containing usnic acid); pycnidia black, common and conspicuous; thallus
yellowish white, patchy soralia becoming confluent and leprose; Maritime Canada in old-growth
forests on bark …………………………………………….……………............. [Cliostomum leprosum]
7. Apothecia whitish to pale grey or brownish grey; pycnidia white, inconspicuous; thallus consisting of
fine, soredia-like, green goniocysts giving it a leprose appearance; widespread, boreal to temperate
……………………………………………………….……….[Micarea prasina]
8. Thallus and apothecial sections C+ orange, xanthones present. Spores 9.3–10.9 × 2.9–3.3 µm; so far
only known from Newfoundland ……………………….......……………………........[B. sphaeroidiza]
8. Thallus and apothecial sections C–, without xanthones. ….……………........….....................................9
9. Thallus PD+ orange red, argopsin present ...........................…………...................................................10
9. Thallus PD–, without major secondary substances ...............………….................................................11
10. Thallus terricolous, distinctly areolate, areoles 0.25–1 mm in diam., extremely hard; (sub)arctic species
..............................……………………………………….......………………………….......... [B. cuprea]
10. Thallus corticolous, rimose to areolate, areoles 0.15–0.35 mm in diam., more or less fragile;
species of boreal coniferous and mixed coniferous forests ..…………..………................ [B. pycnidiata]
87
11. Spores predominantly one-celled, occasionally 2-celled ....…...........…………...............................…..12
11. Spores predominantly with 2 or more cells ...................…………….................................................. 16
12. Hypothecium dark, red-brown to dark brown.……….…....……………... [Mycobilimbia berengeriana]
12. Hypothecium colorless or almost so (slightly yellowish or very pale brown) to yellow-orange .......... 13
13. Apothecia tiny, 0.15-0.3 mm in diam., white or grey, less frequently pale orange-brown;
thallus almost endophloeodal (within the bark); spores (8-)10-12(-17) x 2.0-3.5 μm
…………………………………….…………[”Biatora” albohyalina]
13. Apothecia mostly 0.3-0.7(-1.4) mm in diam., light brown to greyish brown; thallus always
evident, rugose to verruculose; spores over 3.5 μm wide ………………….……………………….... 14
14. Spores narrowly ellipsoid or bacilliform, usually 4–5 times as long as broad; corticolous on
deciduous trees; mainly Appalachian, into the Maritimes ….…………………………....[B. longispora]
14. Spores narrowly ellipsoid to ellipsoid, 3–4 times as long as broad; on bryophytes, peat or
twigs of shrubs near he ground, rarely directly on bark of trees ……………………….…….............. 15
15. Thallus whitish grey, exciple colourless outside, yellowish, orange- or dark brown within or
near the hymenium; hypothecium often orange- to dark brown;, spores 9.7–15.2 × 3.5–5.0
µm; (sub)arctic-(sub)alpine …..…………………………………………………..............[B. subduplex]
15. Thallus greenish grey; exciple and hypothecium colourless to brownish yellow, but without
markedly contrasting pigmentation; spores 12.5–19 × 4-6(-7) µm; in woodlands…….. .... B. vernalis
16. Spores predominantly 2-celled .............…..........…............................….......……………................... 17
16. Spores predominantly 4-celled or more ..........................................…...........……..........………........ 22
17. Thallus consisting of fine, soredia-like, green goniocysts giving it a leprose appearance; spores (7-)
8-14 x 2.5-4(-5) μm, ellipsoid; on bark and wood …………………………… .......... [Micarea prasina]
17. Thallus thin or thick, verruculose to rugose, never with powdery green goniocysts; spores 8-13(-17) x
2-5 μm, narrowly ellipsoid to fusiform; or on various substrates …………………….…………..... 18
18. Paraphyses capitate, with pigmented tips .................................................................………….............19
18. Paraphyses not conspicuously expanded at tips, not pigmented ....................…………...................... 20
19. Epihymenium greenish, intensifying with KOH ....………….......…………......... [Catillaria globulosa]
19. Epihymenium brownish, not changing with KOH ...............………………..... [Catillaria nigroclavata]
20. Apothecia with persistent, prominent margins; disks lightly pruinose, varying from pale
88
pinkish grey to black ; on bark or wood ..….........…………….............................. Cliostomum griffithii
20. Apothecia soon marginless, margins never prominent; disks epruinose, pale to dark brown;
on bryophytes, peat or bark .........................…………………………………………………………21
21. Hypothecium colorless; spores 9.5-11(-17) x 3.5-5 μm; apothecia pale brown to reddish brown; on
bryophytes, peat and bark ……..……………. [Mycobilimbia pilularis (syn. Catillaria sphaerioides)]
21. Hypothecium dark brown; spores 15-27 x (4.5-)5-7.5(-8) μm; apothecia black to very dark brown; on
bryophytes ………..………………………………………………………..[Mycobilimbia hypnorum]
22. Hypothecium dark brown…………………………………………………………………………….23
22. Hypothecium colorless or pale brown, especially in upper parts …………….…………………….. 24
23. Growing on bryophytes, less frequently on tree bases; apothecia 0.3-1.0 mm in diameter; hymenium
colorless, unchanged with KOH; spores (2-)4-6(-8) celled, 18-30(-40) x 5-8 μm
…………………..… Myxobilimbia sabuletorum (syn. “Bacidia” sabuletorum)
23. Growing on bark or wood; apothecia 0.15-0.4 mm in diameter; hymenium greenish, intensifying or
turning purple in KOH; spores (2-)4 celled (rarely 6), 12-21 x 4-5.5 μm ………… [Micarea melaena]
24. Growing on bark or wood; spores (2-)4(-6) celled; 15-23 x 3-6; apothecia pale grey to pale
brown or black……………....................................………….................................. Micarea peliocarpa
24. Growing on bryophytes .......................................……………............................................................. 25
25. Apothecia white or pale grey; spores 15-20 x 5-6 μm ..……………...........[Mycoblimbia carneoalbida]
25. Apothecia black to dark brown ; spores (14-)16-26(-30) x 5.5-8 …………...... Mycoblimbia tetramera
BRIGANTIAE: See Key F.
BRODOA: See Allantoparmelia.
BRYOCAULON: See Alectoria.
BRYORIA: See Alectoria.
BUELLIA (including Amandinea, Catolechia, and Diploicia)
1.
Spores 4-celled to muriform............................................................................................... 2
1.
Spores 2-celled ................................................................................................................... 3
89
2.
Spores muriform, (17-)21-34 x 10-17(-21) μm; hymenium filled with tiny oil drops; on wood and
bark along the west coast ................................................................................ [B. penichra]
2.
Spores 4-celled to few-celled muriform, (11-)15-20(-30) x (5.5-)8-10(-17) μm; hymenium without
oil drops; California to Northeastern U.S………………………………………[B. alboatra]
3.
Thallus yellow or with a yellow tint................................................................................... 4
3.
Thallus pale gray to greenish gray, cortex C–, KC– .......................................................... 5
4.
Thallus greenish yellow; cortex C+ orange, KC+ orange (xanthones); on maritime rocks .
...................................................................................................................... Buellia halonia
4.
Thallus bright lemon-yellow; cortex C–, KC– (containing pulvinic acid-type pigments); on soil
and peat. Spores (12-)13-17(-18) x 7-10 μm; rare but widespread arctic-alpine
..................................................................................................... [Catolechia wahlenbergii]
5.
Spores with angular locules (Physcia-type); thallus thick, distinctly lobed, pruinose and sorediate;
California, mostly coastal, on bark, wood, or rock………………..…...Diploicia canescens
5.
Spores with thin, uniformly thickened walls; thallus thin, not lobed, without pruina or soredia
............................................................................................................................................ 6
6.
Mature apothecia entirely, or almost entirely immersed in the thallus, the disk level with the
thallus surface or slightly depressed; thallus thick, rimose-areolate, with a black prothallus; on
rock, southern to eastern U.S……………………………………………………. B. spuria
6.
Mature apothecia superficial, not sunken into thallus; thallus thick or thin and disappearing,
lacking a black prothallus; on bark, wood, or rock ............................................................ 7
7.
Thallus composed of scattered or contiguous areoles, sometimes white pruinose; on siliceous rock
in California and the southern Rockies……………………………………..[B. retrovertens]
7.
Thallus thin, continuous or barely visible, not areolate or pruinose; mostly on bark or wood,
occasionally on rock........................................................................................................... 8
8.
Thallus almost imperceptible or very thin and dirty greenish gray to pale gray; apothecia less than
0.5 mm in diameter; very common and widespread on wood, but also sometimes on bark or rock
..............................................................................................................Amandinea punctata
8.
Thallus thin but clearly visible, pale gray; apothecia 0.4-1 mm in diameter; in temperate to boreal
forests, on bark ................................................................................................................... 9
90
9.
Thallus K+ becoming blood red (norstictic acid); hymenium yellowish; spores mostly 12-17 x 5-8
μm .......................................................................................................... Buellia stillingiana
9.
Thallus K– (although it contains atranorin); hymenium colorless; spores mostly 18-26 x 6-11 μm
................................................................................................................ Buellia disciformis
BULBOTHRIX (including Relicina)
1.
Thallus yellowish green ("usnic yellow")……[Relicina; see Comments under Bulbothrix]
1.
Thallus gray........................................................................................................................ 2
2.
Isidia abundant on the thallus surface ................................................................................ 3
2.
Isidia absent; apothecia common ...................................................Bulbothrix confoederata
3.
Lower surface and rhizines black; apothecia abundant; medulla C+ red (lecanoric acid present as
main compound)................................................................................Bulbothrix laevigatula
3.
Lower surface and rhizines pale beige to brown; apothecia rare; medulla C+ pink or C–, K+ red or
K–....................................................................................................................................... 4
4.
Medulla C+ pink, K– (gyrophoric acid present as main compound); common throughout
southeastern coastal plain...................................................................... Bulbothrix goebelii
4.
Medulla C–, K+ red (salazinic acid); common only in Florida
............................................................................................................... [Bulbothrix isidiza]
BYSSOLOMA: See Key F.
CALICIUM (including Chaenotheca, Chaenothecopsis, Mycocalicium, Phaeocalicium, Sphinctrina, and
Stenocybe)
1.
Spores ellipsoid, 1- to 4-celled, brown............................................................................... 2
1.
Spores spherical, 1-celled, colorless to brown ................................................................. 10
2.
Spores 4-celled. Thallus not lichenized; very common on the trunks and branches of balsam fir
................................................................................................................. [Stenocybe major]
2.
Spores 1- to 2-celled........................................................................................................... 3
3.
Spores 1-celled. Thallus not lichenized; very common on old stumps in forests
......................................................................................................... [Mycocalicium subtile]
3.
Spores 2-celled ................................................................................................................... 4
91
4.
Spores narrow, 5.5-7.5 x 2-2.3 μm; capitula without pruina, black; thallus not lichenized;
widespread boreal........................................................................ [Chaenothecopsis debilis]
4.
Spores broader than 3.5 μm; capitula often pruinose ......................................................... 5
5.
On living or dead twigs and branches of poplars; asci remaining intact when mature; spores 10-13
x 4-6 μm, smooth; thallus not lichenized…………………….…[Phaeocalicium populneum]
5.
On decaying wood (without bark); asci disintegrating leaving the spores in a powdery mass;
spores ornamented with ridges or warts; usually with algae in the thallus......................... 6
6.
Thallus granular, yellowish green; capitulum margin and base without pruina
......................................................................................................................Calicium viride
6.
Thallus whitish or gray, or within substrate and almost absent from view, sometimes forming a
whitish stain ....................................................................................................................... 7
7.
Capitulum with a yellow pruina on the margin and base; common in Great Lakes region, scattered
elsewhere.............................................................................................Calicium trabinellum
7.
Capitulum with a white or rusty pruina, or not pruinose.................................................... 8
8.
Thallus yellowish white, granular to leprose; stalks thick, unbranched; spores (9-)11-13(-18) x
(4.5-)6-7.5(-9.5) μm; capitulum often white pruinose; common along the coast in the Pacific
Northwest, occasional elsewhere……………………………….…..
8.
[Calicium lenticulare]
Thallus almost imperceptible, within the wood; stalks rather slender; spores mostly less than 6 μm
wide; capitulum white or brown pruinose; boreal species ................................................ 9
9.
Pruina white, on margin of the capitulum .........................................[Calicium glaucellum]
9.
Pruina rusty brown, on undersurface of capitulum .............................[Calicium salicinum]
10.(1)
Parasitic on lichens, especially Pertusaria; stalk extremely short or absent, never longer than
the capitulum, light to dark brown; spores 5-7 μm in diameter, almost spherical; widespread
………………………………………………………..[Sphinctrina turbinata (Pers.:Fr.) De Not.
]
10.
Not parasitic on lichens; stalks much longer than capitulum, black or almost black; spores less
than 5 μm in diameter....................................................................................................... 11
11.
Thallus leprose, with bright yellow-green powdery soredia; stalk covered with yellow pruina;
capitulum also yellow pruinose.......................................................Chaenotheca furfuracea
92
11.
Thallus within substrate, absent from view, or sometimes consisting of a very thin greenish
powder; stalk black, not pruinose; capitulum brown, without pruina ..................................
......................................................................................................... Chaenotheca brunneola
CALOPADIA: See Key F.
CALOPLACA
1.
Thallus fruticose, composed of a tangle of cylindrical branches; maritime California coast
............................................................................................................ Caloplaca coralloides
1.
Thallus crustose, or squamulose......................................................................................... 2
2.
Thallus within substrate and absent from view, or very thin and indistinct ....................... 3
2.
Thallus clearly visible and well-developed ...................................................................... 10
3.
Parasitic on a variety of saxicolous lichens.........................................Caloplaca epithallina
3.
Not parasitic ....................................................................................................................... 4
4.
Growing on bark or wood .................................................................................................. 5
4.
Growing on rock ................................................................................................................ 7
5.
In maritime habitats along the west coast, especially on beach logs
...........................................................................................................Caloplaca inconspecta
5.
In inland localities throughout the continent, on bark and wood ....................................... 6
6.
Apothecia dark orange to yellowish orange; spores 10-13 x 5-7 μm
..............................................................................................................Caloplaca holocarpa
6.
Apothecia rusty red-orange; spores 12-18(-20) x 6-10(-11) μm
...........................................................................................................[Caloplaca ferruginea]
7.(4)
Apothecia biatorine; hymenium containing abundant oil drops
.............................................................................................................Caloplaca luteominia
7.
Apothecia lecanorine; hymenium clear, without oil drops................................................. 8
8.
On shoreline rocks along the west coast; apothecial disks yellow-orange; spores with a broad
septum (over 2.5 μm) ........................................................................Caloplaca inconspecta
93
8.
On non-maritime rocks mostly east of California; apothecial disks dark orange or brownish
orange; spores with a very narrow septum (under 2 μm) ................................................... 9
9.
On calcareous rock and concrete; apothecial margin even with disk; epihymenium C........................................................................................................... Caloplaca feracissima
9.
On non-calcareous rock such as granite or gneiss; apothecial margin prominent; epihymenium C+
purple ......................................................................................................Caloplaca arenaria
10.(2)
Thallus distinctly lobed at the margin .............................................................................. 11
10.
Thallus not lobed at the margin (but sometimes with lobed squamules).......................... 14
11.
Round to irregular soralia containing granular soredia forming on upper surface of thallus;
growing on limestone, usually on dry overhangs ……… Caloplaca cirrochroa (Ach.) Th. Fr.
11.
Not sorediate, but sometimes granular or isidiate in part ……………………………11a
11a.
Older parts of thallus dissolving into granular isidia or granules, with a ring of narrow lobes at the
periphery; in marine habitats (salt spray zone)……………………….Caloplaca verruculifera
11a.
Older parts of thallus not dissolving into granules; in inland or coastal (but not marine) habitats
.......................................................................................................................................... 12
12.
Thallus thin, with rather flat lobes, often broadest at the tips; dark orange to red-orange; found
only in southern California.......................................................................... Caloplaca ignea
12.
Thallus with thick, convex lobes; western to widespread ................................................ 13
13.
Lobes relatively loosely attached; lower surface corticate; very widespread
..................................................................................................................Xanthoria elegans
13.
Lobes closely attached; lower surface without a cortex; western, centered in the Great Basin
..........................................................................................................Caloplaca trachyphylla
14 (10). Apothecial disks brown, K-, pruinose; spores 13-19 x 7-10 μm; eastern forests
............................................................................ [Caloplaca camptidia (Tuck.) Zahlbr.]
14.
Apothecial disks yellow to orange, K+ deep red-purple, pruinose or not ......................... 15
15.
Thallus pale to dark gray or pale brown, not yellowish ..................................................15a
15.
Thallus distinctly yellow, or rusty to bright orange ......................................................... 18
94
15a.
Growing on bark; thallus pale gray to blue-gray with similarly gray apothecial margins
....................................................................................................................Caloplaca cerina
15a.
Growing directly on rock ................................................................................................. 16
16.
Thallus pale beige, thin; apothecia biatorine with margins the same color as the disk (orange or
red); coastal localities, California to Vancouver Island………………Caloplaca luteominia
16.
Thallus gray; apothecial margins lecanorine, gray to black ............................................ 17
17.
Thallus pale gray, continuous to areolate, not squamulose or lobed; apothecia with a dark, narrow
ring of tissue between the disk and the margin; common from the southwest through central U.S.
to New England……………………………………………………..………[Caloplaca sideritis]
17.
Thallus dark olive gray or brownish gray, thick, consisting of lobed squamules; lacking a dark ring
between the apothecial disk and margin; restricted to the arid southwest............................
..............................................................................................................Caloplaca pellodella
18.(14) Soredia present ................................................................................................................. 19
18.
Soredia absent .................................................................................................................. 22
19.
Thallus continuous, with discrete round to irregular soralia on thallus surface; on poplar bark in
the boreal region.......................................................................[Caloplaca chrysophthalma]
19.
Thallus areolate; on rocks or wood, rarely on bark .......................................................... 20
20.
Thallus areoles usually on a conspicuous orange prothallus; apothecia common; on rocks in
strictly marine habitats (upper intertidal and salt spray zones); areoles often constricted at the base,
some dissolving into granular soredia ……………………….….Caloplaca flavogranulosa
20.
Prothallus absent; apothecia rare; on rock or wood, rarely bark, marine or not………..21
21.
Thallus and soredia yellow to orange-yellow; soredia developing first at the edges of the areoles,
but frequently taking over the thallus; on rock or wood in maritime or inland localities.....
................................................................................................................... Caloplaca citrina
21.
Thallus and soredia dark orange; soredia mostly remaining along the areole margins or in patches
on the surface; common on exposed wood or sometimes bark, never rock; from Colorado to the
northeast
…………………………………………………….[Caloplaca microphyllina]
22.(18) Thallus consisting of round or squamulose, convex areoles frequently constricted at the base, up to
2 mm in diameter, usually shiny or "waxy;" on rocks, coastal California ……..Caloplaca bolacina
95
22.
Thallus continuous or composed of small areoles (mostly less than 0.6 mm in diameter) or
granules; areoles not constricted at the base, not waxy in appearance; on rocks, bark, or wood
.......................................................................................................................................... 23
23.
On bark or wood............................................................................................................... 24
23.
On rock, rarely on wood................................................................................................... 25
24.
Thallus sparse, consisting of scattered small areoles; apothecial margin the same color as the disk;
on rocks or wood; maritime, along the west coast………………….….Caloplaca inconspecta
24.
Thallus continuous and rather smooth; apothecial margin paler than the disk; on bark or
sometimes wood; widespread in central and eastern regions………..Caloplaca flavorubescens
25.
Thallus continuous, mostly smooth to rimose or verruculose; widespread east of Arizona, Nevada,
and Idaho.......................................................................................Caloplaca flavovirescens
25.
Thallus mostly areolate or granular, the areoles contiguous or dispersed, sometimes becoming
continuous or thin and rimose .......................................................................................... 26
26.
Mostly northeastern, not maritime; thallus orange, areolate, the areoles often somewhat lobed; on
calcareous rock........................................................................................[Caloplaca velana]
26.
Along the extreme west coast, mostly maritime; thallus orange to yellowish; areoles lobed or not;
on siliceous or calcareous rock......................................................................................... 27
27.
Thallus well-developed, usually with an orange prothallus (see couplet 20)
...................................................................................................... Caloplaca flavogranulosa
27.
Thallus very thin; prothallus absent ................................................................................. 28
28.
Apothecia biatorine; apothecial disks dark orange (var. luteominia) or red (var. bolanderi);
hymenium containing abundant oil drops………………Caloplaca luteominia
28.
Apothecia lecanorine; apothecial disks yellow-orange; hymenium clear, without oil drops
...........................................................................................................Caloplaca inconspecta
CANDELARIA
1.
Soredia absent; apothecia abundant ........................................................ Candelaria fibrosa
1.
Soredia present; apothecia rare ............................................................ Candelaria concolor
CANDELARIELLA
1.
Thallus composed of tiny dispersed areoles breaking down into clumps of soredia, sometimes
96
coalescing into a leprose crust…………………………………... Candelariella efflorescens
1.
Thallus without soredia, but can be granular to areolate ............................................... 2
2.
Growing on soil.................................................................................................................. 3
2.
Growing on bark, wood, or rock ........................................................................................ 4
3.
On tundra soil; arctic-alpine; thallus granular to areolate, thin and scanty.
.......................................................................................................... Candelariella terrigena
3.
Mostly in sagebrush-juniper habitats; thallus composed of convex, often lobed squamules,
relatively thick................................................................................... Candelariella rosulans
4.
Sterile (without apothecia); thallus granular to areolate or verrucose ……………………..5
4.
With apothecia ………………………………………………………………………………… 6
5.
Granules rounded, 0.03-0.15 mm in diameter; on bark or wood
…………………………………………………………[Candelariella xanthostigma]
5.
Granules or areoles round or irregular in shape, larger than 0.15 mm in diameter; on noncalcareous rock or wood, rarely on bark ……………………………. Candelariella vitellina
6.
Spores 16-32 per ascus; on non-calcareous rock or wood………………….Candelariella vitellina
6.
Spores 8 per ascus .............................................................................................................. 7
7.
Thallus scanty, dispersed areolate; apothecial margin the same color as the disk; on noncalcareous
rock, wood, or bark ........................................................................... Candelariella rosulans
7.
Thallus usually within substrate, absent from view, or poorly developed, granular; apothecial
margin paler than the disk; on calcareous rock, occasionally wood.....................................
............................................................................................................. Candelariella aurella
CANDELINA: See Key J.
CANOMACULINA: See Rimelia.
CANOPARMELIA: See Parmelia.
CATILLARIA: See Key F and Biatora.
CATINARIA: See Key F.
97
CAVERNULARIA
1
Soredia present on upper surface of lobe tips; apothecia rare………….Cavernularia hultenii
1
Soredia absent; apothecia abundant, very broad compared to the width of the lobes ..........
........................................................................................................... Cavernularia lophyrea
CETRARIA (including Arctocetraria, Cetrariella, and Masonhalea)
1.
On trees and shrubs, subalpine; marginal projections very few; medulla PD–
.................................................................................................. Tuckermannopsis subalpina
1.
On the ground, sometimes mixed in moss mats or low heath; medulla PD– or PD+ red
(fumarprotocetraric acid).................................................................................................... 2
2.
Branches round in cross section; thallus entirely fruticose; with round, depressed pseudocyphellae
....................................................................................................................Cetraria aculeata
2.
Branches flat; thallus erect foliose; pseudocyphellae either irregular in shape or linear, depressed
or not .................................................................................................................................. 3
3.
Lower surface pruinose, with a regular pattern of white and dark areas; medulla UV+ blue-white
(alectoronic acid)........................................................................... Masonhalea richardsonii
3.
Lower surface not pruinose; medulla UV– (alectoronic acid absent) ................................ 4
4.
Medulla KC+ red, C+ red or pink (gyrophoric acid); lobe margins more or less even, not toothed
or ciliate................................................................................................... Cetrariella delisei
4.
Medulla KC– (gyrophoric acid absent); lobe margins even, or toothed, or ciliate ............ 5
5.
Medulla PD+ red ................................................................................................................ 6
5.
Medulla PD– ...................................................................................................................... 7
6.
Pseudocyphellae marginal, forming an almost unbroken line along the lobe margins
................................................................................................................. Cetraria laevigata
6.
Pseudocyphellae laminal and irregular in shape, but often also along the margins here and there
.................................................................................................................. Cetraria islandica
7.
Mainly temperate; thallus gray-olive to olive-brown when dry…………. Cetraria arenaria
7.
Mainly arctic-alpine; thallus yellowish brown to reddish brown when dry ....................... 8
8.
Pseudocyphellae mostly marginal ......................................................... Cetraria ericetorum
98
8.
Pseudocyphellae mostly laminal ........................................................................................ 9
9.
Pseudocyphellae broad and conspicuous; margins conspicuously fringed with projections;
widespread boreal to montane; contains protolichesterinic acid and (or) lichesterinic acid
..................................................................................... rare PD– strain of Cetraria islandica
9.
Pseudocyphellae small and dark, usually inconspicuous; marginal projections usually very short
and toothlike, rarely forming a noticeable fringe; arctic to northern boreal; contains rangiformic
acid............................................................................................... [Arctocetraria andrejevii]
CETRARIELLA: See Cetraria and Key A.
CETRELIA
1.
Soredia absent; on tundra in arctic Alaska ..............................................[Cetrelia alaskana]
1.
Soredia present along lobe margins; on trees or rocks ....................................................... 2
2.
Medulla C+ red or pink, UV– (olivetoric acid); pseudocyphellae sparse and small (up to 0.3 mm in
diameter) ............................................................................................... Cetrelia olivetorum
2.
Medulla C–......................................................................................................................... 3
3.
Medulla KC+ pink to red, UV+ blue-white (alectoronic acid); pseudocyphellae abundant and often
large (0.15-0.6 mm in diameter)................................................................ Cetrelia chicitae
3.
Medulla KC–, rarely KC+ faint pink (perlatolic or imbricaric acid); pseudocyphellae small and
inconspicuous.....................................................................................[Cetrelia cetrarioides]
CHAENOTHECA: See Calicium.
CHRYSOTHRIX: See Key C.
CLADINA: See Cladonia.
CLADONIA (including Cladina)
1.
Podetia much branched ...................................................................................................... 2
1.
Podetia unbranched, or branched only once or twice, or absent ..................................... 25
2.
Podetia without a cortex; surface dull and webby.............................................................. 3
2.
Podetia with a cortex; surface usually somewhat shiny ................................................... 13
99
3.
Podetia silver gray to pale greenish gray; thallus KC–, K– or K+ yellow ......................... 4
3.
Podetia pale yellow-green or greenish yellow (usnic-yellow); thallus KC+ yellow, K– .. 7
4.
Thalli forming tight, rounded tufts without clearly defined main stems; southeastern coastal plain
......................................................................................................................Cladina evansii
4.
Thalli not forming tight, rounded tufts; main stems usually obvious; mainly northern or western
............................................................................................................................................ 5
5.
Branching at tips divergent, not bent in one direction; thallus PD–, K–, UV+ bright blue-white
(perlatolic acid) .......................................................................................Cladina portentosa
5.
Branching at tips at least partly bent in one direction giving the thallus a combed appearance;
thallus PD+ red, K+ pale yellow, UV– (atranorin and fumarprotocetraric acid) .............. 6
6.
Basal part of podetia and stereome blackened (melanotic); pycnidial jelly red
........................................................................................................................Cladina stygia
6.
Basal part of podetia and stereome pale, more or less the same as upper portions; pycnidial jelly
hyaline....................................................................................................Cladina rangiferina
7.(3)
Podetia branching mostly in twos (dichotomies) .............................................................. 8
7.
Podetia branching mostly in threes and fours (trichotomies or tetrachotomies)…………..9
8.
Podetia thick, 0.7-2 mm wide; podetial axils often open; stereome composed of distinct strands;
thallus PD– or PD+ yellow (with or without psoromic acid; fumarprotocetraric acid absent)
........................................................................................................Cladonia pachycladodes
8.
Podetia slender, usually under 0.7 mm wide; podetial axils mostly closed; stereome continuous,
not broken into strands; thallus PD+ red (fumarprotocetraric acid) ......... Cladina subtenuis
9.
Thallus forming tight, rounded tufts without obvious main stems; podetial tips with radiating
branches around an open hole .................................................................... Cladina stellaris
9.
Thallus forming loosely organized cushions with clearly recognizable main stems; tips without
radiating branches around an open hole .......................................................................... 10
10.
Surface flocculent (with small fluffy clumps) except close to the branch tips; thallus UV+ bright
blue-white (perlatolic acid) .....................................................................Cladina portentosa
10.
Surface compact, smooth or bumpy, not flocculent; thallus UV– (lacking perlatolic acid)11
11.
Thallus PD+ red (fumarprotocetraric acid, lacking fatty acids)……….. Cladina arbuscula
100
11.
Thallus PD– (containing fatty acids)................................................................................ 12
12.
Branches very robust, 0.7-2 mm wide, often sprawling and strongly wrinkled; axils broadly open;
branching usually in fours; containing pseudonorangiformic acid; on northeastern coast...
....................................................................................................................Cladina submitis
12.
Branches usually slender, 0.5-0.8 mm wide, always erect, generally smooth; axils often closed or
only slightly open; branching usually in threes, usually containing rangiformic acid; widespread
boreal to north temperate……………………………………………………….Cladina mitis
13.(2)
Thallus gray to gray-green, often browned in sunny habitats, KC–, K– or K+ yellow (usnic acid
absent) .............................................................................................................................. 14
13.
Thallus yellowish green or greenish yellow, KC+ yellow (usnic acid present)………..…18
14.
Primary squamules large and persistent; podetia up to 25 mm tall; thallus K+ deep yellow
(thamnolic acid) ......................................................................................Cladonia floridana
14.
Primary squamules usually disappearing (although there may be few to many on the podetia);
podetia 20-60 (-120) mm tall; thallus K– ........................................................................ 15
15.
Podetia granular sorediate at the tips................................................. Cladonia scabriuscula
15.
Podetia without soredia or granules ................................................................................. 16
16.
Podetia lacking squamules, surface with flat areoles over a transluscent stereome; thallus very
Cladina-like, PD–, KC+ pinkish violet, rapidly disappearing (merochlorophaeic acid); uncommon,
arctic and west coast…………………………..[Cladonia wainioi (syn. C. pseudorangiformis)]
16.
Podetia usually having at least a few (often many) squamules, with a continuous cortex; thallus
stiff, PD+ red, KC– (fumarprotocetraric acid)…………………………………..17
17.
Podetia with unequal branches that are frequently split lengthwise
................................................................... uncupped morphotype of Cladonia multiformis
17.
Podetia with more or less equal, dichotomous branches, split or intact
.................................................................................................................... Cladonia furcata
18.(13) Apothecia red, almost spherical ............................................................... Cladonia leporina
18.
Apothecia brown or absent............................................................................................... 19
19.
Stereome rudimentary or absent, not forming a well-defined, cartilaginous cylinder or strands;
southeastern coastal plain................................................................................................. 20
101
19.
Stereome forming an intact, cartilaginous cylinder or network of cartilaginous cords or strands;
southeastern or widespread .............................................................................................. 21
20.
Podetial wall perforated with oval holes; podetial axils open; surface very shiny; very
rare……………………………………………………………………...Cladonia perforata
20.
Podetial wall not perforated or longitudinally split; podetial axils closed; surface smooth but
usually dull; common ............................................................................... Cladonia leporina
21.
Stereome cylindrical, smooth, not broken into strands .................................................... 22
21.
Stereome broken into broad or narrow strands ................................................................ 23
22.
Podetia tall and slender, pointed at the tips or forming narrow but distinct cups; axils often closed
or only partially open; barbatic acid present………………………….
22.
Cladonia amaurocraea
Podetia forming densely branched cushions, entirely without cups; axils wide open; barbatic acid
absent; with or without squamatic acid……………………….………………Cladonia uncialis
23.
Podetia slender, 0.5-1.5 mm in diameter, not appearing inflated, usually forming rather flattened
mats; stereome cords broad and flat……………………………………Cladonia dimorphoclada
23.
Podetia broad, mostly 3-11 mm in diameter, appearing inflated, usually erect and cushion forming;
stereome with broad or narrow cords ............................................................................... 24
24.
On rock, less frequently on soil; stereome barely broken into broad bands; podetial wall intact, not
perforated; southeastern U.S.. ............................................................ .Cladonia caroliniana
24.
On sandy soil; stereome broken into a meshwork of narrow cords; podetial wall perforated and
fissured; northeastern coastal plain ...............................................................Cladonia boryi
25.(1)
Main thallus consisting of primary squamules; podetia absent or extremely small (under 4 mm
high) [Note: Many young or poorly developed specimens of Cladonia may key out here]
.......................................................................................................................................... 26
25.
Main thallus consisting of many podetia, usually over 4 mm tall; with or without a basal thallus of
primary squamules ........................................................................................................... 33
26.
Lower surface of squamules pale yellow, upper surface yellow-green to olive; thallus KC+ yellow,
or orange (usnic and barbatic acids)........................................................ Cladonia robbinsii
26.
Lower surface of squamules white; upper surface grayish green, brownish, or olive; thallus KC–
(usnic and barbatic acids absent)...................................................................................... 27
102
27.
Primary thallus forming a lobed, almost foliose, closely attached rosette; medulla K– (atranorin
usually absent)......................................................................................... Cladonia pocillum
27.
Primary thallus composed of discrete squamules, or if forming a rosette, then loosely attached and
ascending; medulla K– or K+ pale yellow or red (atranorin, sometimes with norstictic acid)
.......................................................................................................................................... 28
28.
Primary squamules extremely large, 10-20 mm long, 2-8 mm wide................................ 29
28.
Primary squamules mostly less than 8 mm long and 4 mm wide..................................... 30
29.
Squamules usually forming a radiating rosette, curled up at the margins when dry; Florida and
vicinity .................................................................................................... Cladonia prostrata
29.
Squamules separate, not forming radiating rosettes; western and northern regions
......................................................................................................Cladonia macrophyllodes
30.
Thallus C+ green, KC+ green (strepsilin); thallus olive-green, sometimes forming almost
spherical, vagrant colonies ...................................................................... Cladonia strepsilis
30.
Thallus C–, KC–; thallus gray-green, never forming spherical colonies.......................... 31
31.
Thallus PD– or PD+ yellow (psoromic or norstictic acid often present)
.........................................................................................................Cladonia symphycarpia
31.
Thallus PD+ red (fumarprotocetraric acid) ...................................................................... 32
32.
Squamules finely divided; apothecia raised on short, non-corticate stalks 1-2 mm high; medulla
K–......................................................................................................... Cladonia caespiticia
32.
Squamules strap-shaped, lobed but not finely divided; apothecia produced directly on basal
squamules; medulla K+ yellow (atranorin)……………………………Cladonia apodocarpa
33.
Most or all podetia with cups or cup-like expansions ...................................................... 34
33.
Most or all podetia lacking cups ...................................................................................... 71
34.
Podetia sorediate, or with spherical, corticate granules. .................................................. 35
34.
Podetia without soredia or granules ................................................................................. 50
35.
Thallus distinctly yellow-green or greenish yellow, KC+ yellow (containing usnic acid); most
species have red apothecia ............................................................................................... 36
35.
Thallus gray, gray-green, pale green or olive, often becoming brown, KC– or KC+ pinkish orange
(usnic acid absent); most species have brown apothecia.................................................. 40
103
36.
Podetia mostly over 25 mm tall, with relatively narrow cups; soredia farinose; apothecia red
.......................................................................................................................................... 37
36.
Podetia 6-25(-30) mm tall, with narrow or broad cups; soredia farinose or granular; apothecia red
or pale brown ................................................................................................................... 38
37.
Cups malformed, with often irregularly dentate margins; podetia typically fissured; medulla (not
surface) UV+ blue-white (squamatic acid)…………………………………..…Cladonia sulphurina
37.
Cups usually well-formed, with even, often dentate or proliferating margins; podetial wall intact or
fissured; medulla UV– (zeorin)……………………………………….…………Cladonia deformis
38.
Primary squamules usually sorediate on the lower surface near the margins; cups very narrow;
apothecia red; thallus PD+ orange, K+ yellow, UV– (thamnolic acid) or PD–, K–, UV+ blue-white
(squamatic acid)…………………………………………………………………Cladonia umbricola
38.
Primary squamules never sorediate; cups goblet-shaped, narrow or broad; apothecia red or brown;
lacking both thamnolic and squamatic acids, containing zeorin ...................................... 39
39.
Apothecia and pycnidia bright red; soredia granular; cups relatively broad
.................................................................................................................. Cladonia pleurota
39.
Apothecia and pycnidia pale brown; soredia farinose; cups broad or narrow
..................................................................................................................Cladonia carneola
40.(35) Cups opening by a gaping hole; cup margins bent inward………………Cladonia cenotea
40.
Cups closed ...................................................................................................................... 41
41.
Apothecia or pycnidia red ...............................................................42 (also see couplet 37)
41.
Apothecia or pycnidia brown or absent............................................................................ 43
42.
Primary squamules finely divided, sorediate mostly near margins of lower surface; cups narrow,
not toothed; mostly coastal.................................................................... Cladonia umbricola
42.
Primary squamules large, rounded, lower surface uniformly sorediate; cups trumpet- to gobletshaped, often toothed at margin; mostly inland…………………………..Cladonia digitata
43.
Cups broad, goblet-shaped ............................................................................................... 44
43.
Cups narrow, trumpet-shaped or irregular........................................................................ 45
44.
Soredia granular; thallus gray-green or brownish, never gray; lacking fatty acids; very common
104
and widespread...................................................................................Cladonia chlorophaea
44.
Soredia farinose or, in part, granular; thallus gray to gray-green or very pale geen; containing fatty
acids; west coast...................................................................................... Cladonia asahinae
45.
Cups proliferating from the center as well as from the margins; podetia blackened at the base
(melanotic) with a spotty appearance; soredia coarsely granular
.............................................................................................................Cladonia verruculosa
45.
Cups proliferating from the margins (if at all); podetia not blackened at the base........... 46
46.
Cups well-formed, trumpet-shaped .................................................................................. 47
46.
Cups flat, shallow or irregular and aborted ...................................................................... 48
47.
Podetia with only farinose soredia; never with a bluish tint when wet; lacking fatty acids;
widespread ..............................................................................................Cladonia fimbriata
47.
Podetia with granular and farinose soredia; often with a bluish tint when wet; containing fatty
acids; west coast...................................................................................... Cladonia asahinae
48.
Cups very narrow and rather flat, with starlike proliferations; podetia brownish to olive, granular
sorediate; thallus PD– or PD+ red (homosekikaic acid, with or without fumarprotocetraric acid)
...........................................................................................................................Cladonia rei
48.
Cups poorly formed and irregular, without regular proliferations on the cup margins; podetia graygreen to pale green or brownish; soredia farinose or granular; thallus PD+ red (fumarprotocetraric
acid, lacking homosekikaic acid)…………………………………………….………………..49
49.
Soredia produced in irregular patches on the upper 1/2 to 3/4 of the podetia, often becoming
continuous at the tip; soredia usually granular; thallus greenish, often browned in part
............................................................................................................ Cladonia ochrochlora
49.
Soredia covering podetia, entirely farinose; thallus ashy gray to greenish gray
..................................................................................................................Cladonia subulata
50.(34) Thallus yellowish green or greenish yellow (containing usnic acid); apothecia red ........ 51
50.
Thallus not yellowish (usnic acid absent); apothecia brown ............................................ 52
51.
Podetia and cup interior covered with round areoles, not squamulose; cup margins smooth or
proliferating; thallus UV–, PD–, K– (barbatic acid present; squamatic and thamnolic acids absent)
...................................................................................................................Cladonia borealis
51.
Podetia and cup margins squamulose; thallus UV+ blue-white, PD–, K–, or UV–, PD+ orange, K+
105
deep yellow (barbatic acid absent; containing squamatic or thamnolic acid)….Cladonia bellidiflora
52.
Cups opening by a gaping hole; thallus PD–, UV+ blue-white, or PD+ orange, UV– (with
squamatic or thamnolic acid) ........................................................................................... 53
52.
Cups closed or partially perforate; thallus PD+ yellow or red (psoromic or fumarprotocetraric
acid).................................................................................................................................. 54
53.
Podetial cortex continuous; surface with or without squamules…………Cladonia crispata
53.
Podetial cortex absent; surface covered with large or extremely small squamules
............................................................................................................... Cladonia squamosa
54.
Podetia blackened at the base .......................................................................................... 55
54.
Podetial base more or less the same color as upper portions............................................ 57
55.
Cups proliferating from the margins; widespread ..............................Cladonia phyllophora
55.
Cups proliferating from the center ................................................................................... 56
56.
Arctic regions; thallus K+ pale yellow (atranorin); PD+ red (fumarprotocetraric acid); podetia with
pale areoles over a black stereome giving the upper part a mottled appearance
......................................................................................................................Cladonia trassii
56.
Eastern and southern coastal plain; thallus K–, PD+ red or yellow (fumarprotocetraric or psoromic
acid; atranorin absent); primary squamules gray-green; podetia more or less uniform in color, or
becoming brown at the tips ..........................................................................Cladonia rappii
57.(54) Cups proliferating from the center .................................................................................. 58
57.
Cups not proliferating, or proliferating from the margins ................................................ 61
58.
Thallus K+ pale yellow (atranorin) ..............................................Cladonia macrophyllodes
58.
Thallus K– (atranorin absent)........................................................................................... 59
59.
Cups poorly formed and asymmetrical, having both marginal and central proliferations
.......................................................................................................... Cladonia mateocyatha
59.
Cups round and symmetrical, prolferating almost exclusively from the center…………60
60.
Cups flaring abruptly, very flat ....................................................................Cladonia rappii
60.
Cups tapered, flaring gradually, remaining concave at least at the edges
.................................................................................... Cladonia cervicornis ssp. verticillata
106
61.(57) Primary thallus persistent ................................................................................................. 62
61.
Primary thallus evanescent............................................................................................... 67
62.
Cups poorly formed and asymmetrical; thallus K– (atranorin absent) …………………..63
62.
Cups broad and goblet-shaped, or narrow and trumpet-shaped; thallus K+ yellow or K–
.......................................................................................................................................... 64
63.
Podetia 10-60 mm tall; cups with only marginal proliferations, often squamulose at the margins
............................................................................................................Cladonia phyllophora
63.
Podetia under 20 mm tall; cups irregular, also having some central proliferations
........................................................................................................... Cladonia mateocyatha
64.
Thallus K+ yellow (atranorin); cups broad or very narrow…………..Cladonia ecmocyna
64.
Thallus K– (mostly lacking atranorin); cups broad, more or less goblet shaped.............. 65
65.
Primary thallus consisting of thick, radiating, closely appressed, almost foliose lobes; on
calcareous soil ......................................................................................... Cladonia pocillum
65.
Primary thallus consisting of distinct, ascending squamules; on acidic soils……………66
66.
Cortex broken into round areoles that cover the podetia and line the cups; podetia under 30 mm
tall ........................................................................................................... Cladonia pyxidata
66.
Cortex continuous, smooth; podetia 20-50 mm tall……….Cladonia gracilis ssp. turbinata
67.(61) Cups and (or) podetia perforated or fissured, usually irregular in form ........................... 68
67.
Cups and podetia closed (imperforate) ........................................................................... 69
68.
Cups with numerous perforations, like a sieve, cortex compact, rather smooth and shiny ..
............................................................................................................ Cladonia multiformis
68.
Cups without sieve-like performations, but podetia irregularly perforate or fissured; surface
dull, almost velvety in texture, especially at tips between areoles .....Cladonia phyllophora
69.
Thallus K+ pale yellow (atranorin) ....................................................... Cladonia ecmocyna
69.
Thallus K– (usually lacking atranorin)............................................................................. 70
70.
Podetia up to 150 mm tall and 3 mm in diameter..................................... Cladonia maxima
70.
Podetia 30-80 mm tall and 0.5-1.5 mm in diameter............................................................70a
107
70a.
Cups typically squamulose at the margins; surface dull, almost velvety in texture, especially
at tips between areoles .................................................................. Cladonia phyllophora
70a.
Cups without marginal squamules; podetia smooth and sometimes shiny throughout
..................................................................................................... Cladonia gracilis ssp. turbinata
71.(33) Podetia sorediate, or with spherical, corticate granules. .................................................. 72
71.
Podetia without soredia or granules ................................................................................. 85
72.
Apothecia and/or pycnidia red ......................................................................................... 73
72.
Apothecia and/or pycnidia brown, or absent .................................................................. 76
73.
Primary squamules with soredia on lower surface; thallus yellow-green or greenish yellow
(containing usnic acid) .......................................................................... Cladonia umbricola
73.
Primary squamules with marginal soredia; thallus gray, gray-green, or brown (usnic acid lacking)
.......................................................................................................................................... 74
74.
Soredia powdery, covering the podetia from top to bottom; apothecia present or very frequently
absent .....................................................................................................Cladonia macilenta
74.
Soredia coarsely granular, sometimes sparse; apothecia almost always present
.......................................................................................................................................... 75
75.
Primary squamules rounded, only slightly lobed; podetia corticate at least on the lower half;
thallus always PD–, K– (barbatic acid)…………………….………………Cladonia floerkeana
75.
Primary squamules finely lobed; podetia entirely without a cortex, revealing the cartilaginous
translucent stereome; thallus PD+ orange, K+ yellow (thamnolic acid) or, less commonly, PD–,
K– (barbatic acid)…………………………………………….……………….Cladonia didyma
76.(72) Soredia coarsely granular, confined to tip of podetia; podetia slender, tall, branched once or twice
........................................................................................................... Cladonia scabriuscula
76.
Soredia or granules covering at least the upper half of podetia........................................ 77
77.
Podetia with a continuous, more or less smooth cortex on the lower 1/3 to 2/3; soredia usually in
discrete patches ................................................................................................................ 78
77.
Podetial cortex more or less broken up with areoles, granules, or soredia, not smooth and
continuous except sometimes very close to the base; soredia, granules or squamules diffuse
.......................................................................................................................................... 80
108
78.
Podetia branched once or twice with mostly unequal branches, commonly split or fissured;
uncommon; northeastern ......................................................................[Cladonia farinacea]
78.
Podetia usually unbranched, rarely fissured; common and widespread ........................... 79
79.
Thallus with gray-green tones predominating; stereome relatively thick and tough
............................................................................................................ Cladonia ochrochlora
79.
Thallus mostly brownish, especially on upper half; stereome usually relatively thin ..........
................................................................................................................... Cladonia cornuta
80.(77) Primary thallus disappearing; podetia 30-100 mm tall, often branched once or twice near the tip,
covered with powdery soredia..................................................................Cladonia subulata
80.
Primary thallus persistent; podetia usually under 35 mm tall........................................... 81
81.
Primary squamules very finely divided, usually disintegrating into a granular crust; thallus K+
deep yellow, PD+ orange (thamnolic acid)…………………………………...Cladonia parasitica
81.
Primary squamules remaining discrete, not granular; thallus K–, PD– or PD+ red (thamnolic acid
absent) .............................................................................................................................. 82
82.
Podetia with a mixture of granules (microsquamules) and soredia on the lower half, frequently
with brown apothecia at the tips; southeastern coastal plain; thallus PD+ red, UV–
(fumarprotocetraric acid)……………………………………………………....Cladonia subradiata
82.
Podetia without a mixture of granules or microsquamules on the lower half, usually without
apothecia; mainly absent from coastal plain (but see C. coniocraea) .............................. 83
83.
Thallus greenish gray to brownish, mostly without a cortex; podetia 10-40 mm tall with only
scattered granular soredia or microsquamules; thallus PD–, UV+ white (perlatolic acid); boreal to
arctic.................................................................................................. [Cladonia decorticata]
83.
Thallus olive or dark green, either covered with cortex or soredia; thallus PD+ red, UV–
(fumarprotocetraric acid); temperate to boreal................................................................ 84.
84.
Soredia mostly granular; podetia corticate on lower 1/4; squamules finely divided
............................................................................................................ Cladonia ochrochlora
84.
Soredia entirely farinose, rarely granular in part, podetia covered with soredia except for a basal
area; squamules typically large and almost undivided………………..Cladonia coniocraea
85.(71) Thallus yellow-green or greenish yellow (usnic acid)...................................................... 86
109
85.
Thallus not yellowish (usnic acid absent) ........................................................................ 92
86.
Apothecia pale brown ...................................................................................................... 87
86.
Apothecia red ................................................................................................................... 88
87.
Growing on soil or rock; primary squamules dense, crenulate or deeply lobed
................................................................................................................. Cladonia robbinsii
87.
Growing on wood; primary squamules scattered, not lobed…………….Cladonia botrytes
88.
Primary squamules sorediate; medulla UV+ white (squamatic acid)............................... 89
88.
Primary squamules without soredia; medulla UV– ......................................................... 90
89.
Soredia on lower surface of squamules; contains didymic acid; common throughout the coastal
plain....................................................................................................... Cladonia incrassata
89.
Soredia on margins of squamules; contains grayanic acid; rare, North Carolina to Florida
................................................................................................................... [Cladonia anitae]
90.
Western and arctic regions; podetial surface areolate, abundantly squamulose
............................................................................................................. Cladonia bellidiflora
90.
Eastern; podetia with a smooth to areolate, cortex squamulose or without any squamules
.......................................................................................................................................... 91
91.
Thallus K–, PD– (barbatic acid); very common, East Temperate to boreal .........................
................................................................................................................Cladonia cristatella
91.
Thallus K+ deep yellow, PD+ orange (thamnolic acid); Florida
.........................................................................................................[Cladonia abbreviatula]
92.(85) Primary thallus disappearing............................................................................................ 93
92.
Primary thallus persistent ................................................................................................. 99
93.
Podetial axils closed......................................................................................................... 94
93.
Podetial axils open ........................................................................................................... 96
94.
Thallus K– ....................................................................................................................... 95
94.
Thallus K+ pale yellow ......................................................................... Cladonia ecmocyna
95.
Podetia up to 150 mm tall and 3 mm in diameter..................................... Cladonia maxima
110
95.
Podetia 30-80 mm tall and 0.5-1.5 mm in diameter………...Cladonia gracilis ssp. gracilis
96.(93) Podetial axils wide open, forming funnels; thallus PD–, or PD+ orange (squamatic or thamnolic
acid; fumarprotocetraric acid absent)………………………………..……Cladonia crispata
96.
Podetial axils partially open or wide open, not forming funnels; thallus PD+ red
(fumarprotocetraric acid).................................................................................................. 97
97.
Thallus K+ pale yellow (atranorin); podetia stout, 1-5 mm in diameter
....................................................................................................................Cladonia turgida
97.
Thallus K– (atranorin absent); podetia mostly slender, under 2 mm in diameter............. 98
98.
Branching unequal; cortex mostly continuous ................................... Cladonia multiformis
98.
Branching in more or less equal dichotomies; cortex usually breaking up into small irregular
patches especially on the upper half of the podetia
...............................................................................Cladonia furcata (western morphotype)
99.(92) Apothecia red; Florida.................................................................................................... 100
99.
Apothecia brown ............................................................................................................ 101
100.
Primary squamules small, white below, often dissolving into soredia
............................................................................................................. [Cladonia ravenellii]
100.
Primary squamules large, with a broad yellow band on lower surface, sometimes sorediate on the
margins.............................................................................................[Cladonia hypoxantha]
101.
Podetia mostly 1-3 mm tall, entirely without a cortex; thallus PD+ red, K– (fumarprotocetraric
acid)...................................................................................................... Cladonia caespiticia
101.
Podetia mostly 3-40 mm tall, with or without a cortex .................................................. 102
102.
Growing on wood or bark .............................................................................................. 103
102.
Growing on soil, moss, or rock ...................................................................................... 105
103.
Primary squamules granular at the margins often reducing the primary thallus to a granular crust;
thallus PD+ orange, K+ deep yellow (thamnolic acid)
................................................................................................................Cladonia parasitica
103.
Primary squamules finely divided but not granular........................................................ 104
104.
Thallus PD+ yellow, K– (baeomycesic acid); coastal plain………….Cladonia beaumontii
111
104.
Thallus usually PD–, K–, UV+ white (squamatic acid) or sometimes PD+ orange, K+ deep yellow
(thamnolic acid); widespread…………………………………………..….Cladonia squamosa
105.(102) Podetia without a cortex.............................................................................................. 106
105.
Podetia at least partly corticate, sometimes broken into areoles or flat squamules ........ 107
106.
Podetia abundantly lacerate, with scattered areoles and microsquamules; contains norstictic
acid ............................................................................................ Cladonia symphycarpia
106.
Podetia not or moderately lacerate; contains other compounds ........................... 106a
106a.
Podetia covered with abundant large to tiny, mostly shingled squamules; containing squamatic or
thamnolic acid (see couplet 104); podetial tips blunt, often with a gaping hole; very common and
widespread ............................................................................................ Cladonia squamosa
106a.
Podetial squamules small and scattered; podetia pointed at tips, sometimes fissured on the sides;
PD–, K–, UV+ white (perlatolic acid); boreal to arctic..................... [Cladonia decorticata]
107.
Medulla C+ green, KC+ green (strepsilin); thallus distinctly olive in color; podetia more or less
inflated .................................................................................................... Cladonia strepsilis
107.
Medulla C–, KC– ........................................................................................................... 108
108.
Thallus K+ yellow, orange, or red.................................................................................. 109
108.
Thallus K– (or brownish), PD+ red (fumarprotocetraric acid) or PD+ yellow .............. 112
109.
Thallus K+ red, PD+ yellow (norstictic acid) ................................................................ 110
109.
Thallus K+ pale yellow, PD– or PD+ yellow or red ...................................................... 111
110.
Podetia rare, stocky, 2-5 mm in diameter, 10-20 mm tall, walls abundantly lacerate and fissured;
containing atranorin ........................................................................Cladonia symphycarpia
110.
Podetia usually abundant, slender, 0.6-2.5 mm in diameter, 8-20 (-30) mm tall, walls intact or
sometimes split; lacking atranorin...................................................Cladonia polycarpoides
111.
Primary squamules large, 2-8 x 1-6 mm; podetia rare, 10-15 mm tall; thallus PD+ yellow
(psoromic acid) or PD– (atranorin alone)…………………………Cladonia symphycarpia
111.
Primary squamules small, thick, 1-3 x 0.5-2 mm; podetia usually abundant, 10-30 mm tall; thallus
PD+ red (fumarprotocetraric acid) or PD– (atranorin alone) .....................Cladonia cariosa
112.(108) Podetia covered with peltate areoles or squamules (attached in the center); thallus K–, PD+
112
bright yellow (psoromic acid); primary squamules 3-8 mm in diameter; arctic-alpine
........................................................................................................[Cladonia macrophylla]
112.
Podetial squamules and areoles not peltate; thallus PD+ red (fumarprotocetraric acid) 113
113.
Apothecia much broader than podetia; podetia minutely warty, slender, often twisted
........................................................................................................... Cladonia peziziformis
113.
Apothecia rarely broader than podetia; podetia smooth, stout, not twisted
............................................................................................................ Cladonia sobolescens
CLIOSTOMUM: See Key F and Biatora.
COCCOCARPIA: See Pannaria.
COCCOTREMA: See Key D.
COENOGONIUM: See Key B.
COLLEMA
1.
Growing on bark .............................................................................................................. 2
1.
Growing on rock, soil, or moss ........................................................................................ 8
2.
Lobes small, under 3 mm wide, crowded into small cushions; spores fusiform, 2-celled,
(13-)15-24(-26) x 3-4.5(-6) μm .................................................. [Collema conglomeratum]
2.
Lobes over 3 mm wide, not cushion-forming; spores fusiform to needle-shaped, 4- to 16-celled
............................................................................................................................................ 3
3.
Isidia absent ....................................................................................................................... 4
3.
Isidia present ..................................................................................................................... 6
4.
Apothecia pruinose.......................................................Collema pulcellum (some varieties)
4.
Apothecia without pruina ................................................................................................... 5
5.
Tissue below the hypothecium in mature apothecia composed of round to angular cells
................................................................. Collema pulcellum var. pulcellum
5.
Tissue below the hypothecium in mature apothecia composed of long, cylindrical cells
.................................................................................. Collema nigrescens
113
6.(3)
Isidia cylindrical in mature thalli, but granular and globose on young thalli, unbranched or
branched ........................................................................................ Collema furfuraceum
6.
Isidia spherical or globular in mature thalli .................................................................... 7
7.
Thallus relatively flat, although sometimes with folds, always isidiate
........................................................................................................... Collema subflaccidum
7.
Thallus with round to elongate pustules or blisters, rarely isidiate ...... Collema nigrescens
8.(1)
Lobes more or less uniform in thickness, not thicker at the margins; lobes mostly 2-6(-10) mm
wide, with or without erect marginal lobules ..................................................................... 9
8.
Lobe with distinctly thickened margins, often producing erect, swollen lobules; small species with
lobes mostly 0.5-3 mm wide
9.
.................................................... 15
Lobes erect and branched, finely divided to lobulate at the margins, under 3 mm wide; spores 4celled to muriform when mature ............................................................. Collema cristatum
9.
Lobes prostrate or on edge, branched or rounded, often over 3 mm wide; spores muriform or not
.......................................................................................................................................... 10
10.
Isidia absent; thallus pustulate; on siliceous rocks; mainly East Temperate
............................................................................................................. [Collema ryssoleum]
10.
Isidia present; thallus pustulate or not; on various kinds of rock ..................................... 11
11.
Isidia flattened like squamules, overlapping ....................................... [Collema flaccidum]
11.
Isidia cylindrical or globular, not flattened ...................................................................... 12
12.
Isidia cylindrical on mature specimens, unbranched or branched; lobes strongly ridged and
pustulate, 5-10 mm wide .................................................................. Collema furfuraceum
12.
Isidia spherical or globular, rarely cylindrical; lobes more or less smooth or folded, not pustulate,
(1-)2-6 mm wide ............................................................................................................. 13
13.
On siliceous rock; lobes mostly flat, with small, gobular isidia (rarely becoming cylindrical);
apothecia rare; spores 6- to 8-celled………………………………..…… Collema subflaccidum
13.
On calcareous rock; lobes crowded, often concave, with large globular isidia; apothecia relatively
abundant ........................................................................................................................... 14
14.
Spores fusiform, 4-celled .......................................... Collema undulatum var. granulosum
114
14.
Spores ellipsoid, muriform................................................................ [Collema fuscovirens]
15.(8)
On calcareous rock........................................................................................................... 16
15.
On soil.............................................................................................................................. 17
16.
Lobes flat, crowded, usually on one edge, without lobules or isidia; apothecia abundant; spores (2)4-celled, narrow, 6.5-8.5 μm wide ..................................................... Collema polycarpon
16.
Lobes with cylindrical, erect branches; globular isidia sometimes present; spores muriform to 4celled, ellipsoid, 8-13 μm wide ............................................................... Collema cristatum
17.
Lobes prostrate; lobules, when present, rounded, strap-shaped, or finely divided; globular isidia
sometimes present on lobe surface; spores mostly 4-celled and fusiform, sometimes submuriform
....................................................................................................................... Collema tenax
17.
Lobes erect, often with cylindrical lobules; spores 2- to 4-celled or submuriform ........ 18
18.
Spores 2-celled; lobules cylindrical, appearing globular from above, but true isidia absent
.......................................................................................................... Collema coccophorum
18.
Spores 4-celled to submuriform; lobules erect, sometimes branched; globular isidia sometimes
present ..................................................................................................... Collema cristatum
CONOTREMA: See Key D, under Stictis urceolatum .
CORNICULARIA: See Key A.
CROCYNIA: See Key C.
CRYPTOTHECIA: See Key C.
CYPHELIUM
1.
Thallus bright greenish yellow; margin of capitulum yellow pruinose
...............................................................................................................Cyphelium lucidum
1.
Thallus pale to dark gray; capitulum entirely black, without pruina
............................................................................................................ Cyphelium inquinans
DACTYLINA
1.
Stalks mostly unbranched, entirely hollow, 20-70 mm tall, 2-6(-14) mm in diameter
................................................................................................................... Dactylina arctica
115
1.
Stalks branched; medulla webby to dense, mostly under 20 mm tall and 2 mm in diameter
............................................................................................................................................ 2
2.
Stalks yellowish to pinkish violet, pruinose on young tips, with irregular, divergent branching,
almost hollow or partially filled with cobwebby hyphae; medulla KC+ pink, usually PD+ red
(physodic acid, usually with physodalic acid)....................................... Dactylina ramulosa
2.
Stalks greenish yellow, not pruinose, branching in regular dichotomies; medulla densely filled
with white hyphae; medulla KC–, PD– (fatty acids)..............Allocetraria madreporiformis
DENDRISCOCAULON: See Key B.
DENDROGRAPHA: See Key A.
DERMATOCARPON
1.
Lower surface covered with short stubby rhizines……………..[Dermatocarpon moulinsii]
1.
Lower surface lacking rhizines .......................................................................................... 2
2.
Thallus entirely without pruinose appearance.................................................................... 3
2.
Thallus appearing pruinose, at least at lobe tips................................................................. 4
3.
Thallus remaining brown when wet, not grass green; growing on dry rocks
......................................................................................................Dermatocarpon miniatum
3.
Thallus grass green when wet; growing on at least periodically submerged rocks ..............
........................................................................................................ Dermatocarpon luridum
4.
Lower surface smooth, without papillae, usually brown………..Dermatocarpon miniatum
4.
Lower surface roughly papillate like coarse sandpaper, usually black
..................................................................................................Dermatocarpon reticulatum
DIBAEIS: See Baeomyces.
DIMELAENA
1.
Thallus greenish yellow (usnic acid); widespread.................................... Dimelaena oreina
1.
Thallus white to brown; California to Washington ............................................................ 2
2.
Thallus creamy white to brownish gray; medulla PD–, K–……………..Dimelaena radiata
2.
Thallus dark brown ............................................................................................................ 3
116
3.
Hypothecium colorless; medulla PD–, K–; common………………[Dimelaena thysanota]
3.
Hypothecium dark brown; medulla PD+ orange, K+ yellow (stictic acid); rare
.........................................................................................................[Dimelaena californica]
DIMERELLA: See Key F.
DIPLOICIA: See Buellia.
DIPLOSCHISTES
1.
Thallus heavily pruinose, thick and areolate with areoles up to 3 mm across; spores 4-8 per ascus;
mainly on bare soil in the arid southwest………………………………….Diploschistes diacapsis
1.
Thallus without, or with little pruina, thin or thick, with areoles less than 1.5 mm across; spores 4or 4-8 per ascus; widely distributed ................................................................................... 2
2.
Spores constantly 4 per ascus; growing on soil, mosses, or lichens
....................................................................................................... Diploschistes muscorum
2.
Spores 4-8 per ascus; growing directly on rock ............................ Diploschistes scruposus
DIRINA: See Keys C and F.
DIRINARIA: See Physcia.
ENDOCARPON: See Key G.
EPHEBE: See Key B.
ERIODERMA: See Peltigera.
ESSLINGERIANA: See Platismatia.
EVERNIA
1.
Branches clearly flattened and dorsiventral throughout, with a pale, almost white lower surface;
round soralia on the lobe margins and upper surface, containing coarse, white to blue-gray soredia
................................................................................................................... Evernia prunastri
1.
Branches irregular or angular in cross section, mostly not flattened or dorsiventral; sorediate or not
............................................................................................................................................ 2
117
2.
Thallus sorediate on ridges along the branches; bushy to pendent………….Evernia mesomorpha
2.
Thallus without soredia, pendent or prostrate .................................................................... 3
3.
Branches stiff and brittle, with a tough, unbroken cortex; medulla dense; prostrate on calcareous
soil in western arctic............................................................................. [Evernia perfragilis]
3.
Branches usually soft, not brittle; cortex thin and cracked; medulla loose (in pendent specimens on
trees) or dense (in prostrate specimens on calcareous soil); Rocky Mountains…Evernia divaricata
EVERIASTRUM: See Key A.
FLAVOCETRARIA
1.
Lobes flat with a network of depressions and sharp ridges or at least wrinkled; base sometimes
turning dark yellow ..............................................................................Flavocetraria nivalis
1.
Lobes curled inward forming a channel, smooth, or undulating and crisped at the margins; base
often becoming blotched with red-violet………………………………..Flavocetraria cucullata
FLAVOPARMELIA: See Key J.
FLAVOPUNCTELIA: See Key J.
FULGENSIA
1.
Spores 2-celled ................................................................................... Fulgensia desertorum
1.
Spores 1-celled ................................................................................................................... 2
2.
Thallus areolate, not noticeably lobed at the margin; schizidia often present
................................................................................................................Fulgensia bracteata
2.
Thallus clearly lobed at the margin; schizidia present or absent ........................................ 3
3.
Lobes long and slightly ascending at the tips; schizidia absent; rare……[Fulgensia fulgens]
3.
Lobes short, entirely adnate; schizidia abundant on thallus surface; common
....................................................................................................... [Fulgensia subbracteata]
FUSCIDEA: See Lecidea.
FUSCOPANNARIA: See Pannaria.
118
GLYPHIS: See Key E.
GOMPHILLIS: See Keys B and F.
GRAPHINA (based on Harris, More Florida Lichens, 1995)
1.
Walls of the lirellae black. Spores 1 per ascus, 65-90 x 20-30 μm; lirellae partly immersed, with a
narrow disk........................................................................................ [Graphina xylophaga]
1.
Walls of lirellae pale to colorless ....................................................................................... 2
2.
Spores 1 or rarely 2 per ascus; lirellae crowded, short and irregular; spores 85-105 x 20-28 μm
............................................................................................................... [Graphina cypresii]
2.
Spores 2 to 8 per ascus ....................................................................................................... 3
3.
Spores small, 15-20(-35) x 6-9(-11) μm, 8 spores per ascus. Lirellae appearing like fissures in the
thallus, disk not exposed; thallus dark olive to yellow-brown, rather shiny, K–, PD–.........
............................................................................................................[Graphina incrustans]
3.
Spores large, over 35μm long, mostly 2-4 spores per ascus; thallus pale gray, K+ or K–, PD+
yellow to red; lirellae white; .............................................................................................. 4
4.
Spores ellipsoid, 35-78 x 18-36 μm; walls of lirellae layered and disintegrating; thallus PD+
orange, K+ yellow to red (constictic acid or salazinic acid); rare ......Graphina peplophora
4.
Spores fusiform, 110-140 x 15-17 μm; walls not disintegrating; thallus PD+ red-orange, K–
(protocetraric acid); common in Florida…………………………….[Graphina abaphoides]
GRAPHIS
1.
Lirellae white or thallus-colored; spores 4-celled .............................................................. 2
1.
Lirellae black, C–; spores 6- to 12-celled........................................................................... 3
2.
Lirellae prominent, thickly white pruinose, C+ red (lecanoric acid); walls of lirellae black; very
common in southeastern coastal plain…………………………………………. Graphis afzelii
2.
Lirellae not prominent, seen as raised fissures in the thallus, C–; walls of lirellae colorless or pale;
common on trees in southeast and in Pacific Northwest coastal forests…….[Graphis insidiosa]
3.
Walls of lirellae with long ridges, in layers like a french pastry; Florida........................... 4
3.
Walls of lirellae uniform, not ridged or layered ................................................................. 5
119
4.
Spores (6-)9- to 10-celled, 25-30 x 6-8(-10) μm; thallus UV– (lacking lichexanthone); rare
............................................................................................................... Graphis subelegans
4.
Spores 6- to 8-celled, 20-30 x 7-9 μm; thallus UV+ yellow (lichexanthone); common
.................................................................................................................. [Graphis lucifera]
5.
Spores 6- to 8-celled; thallus K+ red, PD+ yellow (norstictic acid); common only in Florida and
nearby coastal plain.................................................................................... [Graphis librata]
5.
Spores 8- to 11-celled......................................................................................................... 6
6.
Lirellae prominent or partly immersed; thallus K–, PD– (no lichen substances); extremely
common and widespread, but absent from Florida…………………………..Graphis scripta
6.
Lirellae mostly immersed; thallus K+ yellow to red, PD+ orange (stictic acid, sometimes with
norstictic acid); common in Florida and Louisiana……………………….[Graphis caesiella]
GYALECTA: See Key F.
GYMNODERMA: See Key G.
GYPSOPLACA: See Key G.
HAEMATOMMA
1.
Growing on rock ......................................................................... Haematomma fenzlianum
1.
Growing on bark ................................................................................................................ 2
2.
Thallus sorediate ................................................................................................................ 3
2.
Thallus not sorediate .......................................................................................................... 4
3.
Soralia irregular in shape, with greenish soredia; containing sphaerophorin in the thallus
and russulone in the rare apothecia................................. [Haematomma americanum]
3.
Soralia round and hemispherical, clearly delimited, with white soredia; containing
placodiolic acid derivatives in the thallus and haematommone in the very rare apothecia
...................................................................... [Haematomma guyanense Kalb & Staiger]
4.
Apothecial mostly sunken into thallus; conidia long and curved; epihymenium K+ red
........................................................................................................ Haematomma persoonii
4.
Apothecial superficial; conidia rod-shaped or very long ................................................... 5
120
5.
Spores over 50 μm long; conidia thread-like and twisted; epihymenium K+ red
.....................................................................................................[Haematomma rufidulum]
5.
Spores under 55 μm; conidia rod-shaped; epihymenium K+ violet or purple …………….6
6.
Conidia 5-7 x 0.8-1 μm; containg placodialic acid ......................... Haematomma accolens
6.
Conidia 7-8 x 1.5-2 μm; contains isoplacodialic and isopseudoplacodialic acids
...................................................................................................................... [H. flexuosum]
HEPPIA: See Key G.
HETERODERMIA: See Physcia.
HUBBSIA: See Key A.
HYDROTHYRIA: See Keys H and I, and Leptogium.
HYPERPHYSCIA: See Phaeophyscia.
HYPOCENOMYCE: See Key G.
HYPOGYMNIA (including Menegazzia)
1.
Lobes perforated on the upper surface with round holes; medulla PD+ orange, K+ yellow, AI+
blue (stictic acid) ................................................................................ Menegazzia terebrata
1.
Lobes perforated at tips, or not perforated at all; medulla PD– or PD+ red, K–, AI– (stictic acid
absent) ................................................................................................................................ 2
2.
Soredia present ................................................................................................................... 3
2.
Soredia absent .................................................................................................................... 7
3.
Soredia on expanded, turned back lobe tips (lip-shaped soralia) ....................................... 4
3.
Soredia on the thallus surface, sometimes close to the lobe tips (but not in lip-shaped soralia)
............................................................................................................................................ 5
4.
Medullary ceiling dark brown to grayish-black; lobules present on the lobe margins; medulla PD–
...............................................................................................................Hypogymnia vittata
4.
Medullary ceiling white; lobules absent; medulla PD+ red (physodalic and protocetraric acids)
..........................................................................................................Hypogymnia physodes
121
5.
Thallus ascending; lobes elongated, rarely perforated; soredia farinose, at tips of lobes
.......................................................................................................... Hypogymnia tubulosa
5.
Thallus closely appressed to substrate; lobes rounded, square, or truncated, perforated at tips;
soredia granular, on thallus surface or tips of lobes ........................................................... 6
6.
Soredia mostly on the upper surface of the lobe tips............................. Hypogymnia bitteri
6.
Soredia mostly on the older parts of the thallus surface…………..Hypogymnia austerodes
7.(2)
Eastern U.S. and Canada (Appalachians)............................................Hypogymnia krogiae
7.
Western or northern North America................................................................................... 8
8.
Medullary ceiling white ..................................................................................................... 9
8.
Medullary ceiling light to dark brown to grayish-black ................................................ 13
9.
On soil or rock (rarely wood)........................................................................................... 10
9.
On conifer bark or wood .................................................................................................. 11
10.
Lobules present along the lobe margins; pycnidia sparse or very inconspicuous; in alpine or arctic
tundra ...........................................................................................Hypogymnia subobscura
10.
Lobules absent, pycnidia abundant and conspicuous; mostly in the mountains
.................................................................................................. Hypogymnia metaphysodes
11.
Thallus loosely attached, ascending or pendent, sometimes forming round cushions of overlapping
lobes; lobes convex, perforated at tips; medulla PD+ red (diffractaic, physodalic, and protocetraric
acids present).................................................................................................................... 12
11.
Thallus closely appressed to substrate, forming round, flat rosettes; lobes flat or concave, rarely
perforated; medulla PD– or sometimes PD+ red (diffractaic and protocetraric acids absent,
physodalic acid sometimes present)………………………………..Hypogymnia metaphysodes
12.
Lobes forked in more or less regular dichotomies, more or less even, not constricted at intervals;
thallus rather stiff and shrubby; apothecia abundant……………………Hypogymnia imshaugii
12.
Lobes branching irregularly, constricted at irregular intervals; thallus usually somewhat pendent,
with upturned tips; apothecia rare .................................................... Hypogymnia duplicata
13.(8)
Lobules present along the lobe margins ........................................................................... 14
13.
Lobules absent.................................................................................................................. 16
122
14.
Medulla PD– (physodalic and protocetraric acids absent); lobes mostly less than 3 mm wide
......................................................................................................Hypogymnia occidentalis
14.
Medulla PD+ red (physodalic and protocetraric acids); lobes usually more than 3 mm wide at the
tips.................................................................................................................................... 15
15.
Lobules strap-shaped; diffractaic acid absent; rare, in coastal forests
.....................................................................................................Hypogymnia heterophylla
15.
Lobules rounded; diffractaic acid present; very common in Pacific Northwest
...................................................................................................Hypogymnia enteromorpha
16.(13) Lobes rarely perforated .................................................................................................... 17
16.
Lobes perforated at tips .................................................................................................... 18
17.
Older lobes becoming very wrinkled, often sinuous, convex throughout; apothecia abundant;
medulla PD– (hypoprotocetraric acid present)……………………….Hypogymnia rugosa
17.
Lobes relatively smooth; lobe tips concave; apothecia occasional; medulla PD– or PD+ red
(hypoprotocetraric acid absent)…………………………………Hypogymnia metaphysodes
18.
Lobes 1.5-2 mm wide, more or less even, not constricted at intervals; medulla KC+ red (diffractaic
and physodic acids) ............................................................................ Hypogymnia inactiva
18.
Lobes 3-4 mm wide, constricted at irregular intervals; medulla KC– (diffractaic and physodic
acids absent)...................................................................................... Hypogymnia apinnata
HYPOTRACHYNA
1.
Soredia, or soredia-like fragments originating from the breakdown of pustules or hollow warts
(schizidia), present on lobe surface .................................................................................... 2
1.
Soredia or soredia-like fragments absent ........................................................................... 6
2.
Rhizines largely unbranched, slender and short; soredia usually powdery, but sometimes from
schizidia, on upper surface of lobe tips; medulla C+ red……………..Hypotrachyna revoluta
2.
Rhizines dichotomously branched, sometimes more than once, robust; soredia powdery or from
schizidia; medulla C+ red or C–......................................................................................... 3
3.
Soredia powdery to granular, not originating from pustules ……………………………… 3a
3.
Soredia orginating from pustules (schizidia); medulla C+ red or C–; on bark .......... 4
123
3a.
Medulla C+ red (lecanoric and evernic acids); rare, Appalachian, mostly on
rocks…………………………………………………..…………………[Hypotrachyna rockii]
3a.
Medulla C- or C+ pale pink-orange (barbatic acid complex); Southwestern, on bark
…………………………………….…………………………….. [Hypotrachyna laevigata]
4.
Cortex K–, UV+ yellow (lichexanthone); medulla K+ brown-red, PD– (lividic acid)
....................................................................................................... Hypotrachyna osseoalba
4.
Cortex K+ yellow, UV– (atranorin); medulla K–, PD+ or PD–……………………………5
5.
Medulla white with orange patches, especially under the pustules, PD+ red-orange, K– where
white and K+ purple where pigmented (protocetraric acid and anthraquinone)
.............................................................................................. Hypotrachyna croceopustulata
5.
Medulla white throughout, PD–, K–; medulla C+ pink to red (lecanoric and evernic acids); fairly
common, Appalachians……………………………………………..[Hypotrachyna taylorensis]
6.(2)
Isidia present (sometimes sparse), apothecia infrequent; Appalachian, on bark…………..7
6.
Isidia absent; apothecia abundant and conspicuous ........................................................... 8
7.
Isidia cylindrical, laminal; medulla C– or C+ pale orange (barbatic acid complex)
.................................................................................................. [Hypotrachyna imbricatula]
7.
Isidia flattened like lobules, on the lobe margins and sometimes on the surface; medulla C+ red
(anziaic acid) ............................................................................. [Hypotrachyna prolongata]
8.
Thallus spotted with maculae; medulla K–, KC+ red, C+ red (evernic and lecanoric acids);
southwestern U.S.A........................................................................ Hypotrachyna pulvinata
8.
Thallus lacking maculae; medulla K+ pinkish brown, KC+ purple-brown, C– (lividic acid);
eastern U.S.A. .....................................................................................Hyptotrachyna livida
ICMADOPHILA: See Key F.
IMSHAUGIA
1.
Isidia absent; pycnidia abundant and conspicuous, scattered on lobe surface; apothecia abundant
............................................................................................................ Imshaugia placorodia
1.
Isidia abundant on thallus surface; pycnidia sparse and inconspicuous; apothecia rare
............................................................................................................... Imshaugia aleurites
124
IONASPIS (including Hymenelia and Eiglera)
1.
Apothecial disks black; epihymenium shades of green, changing to wine-red in nitric acid2
1.
Apothecia pink to brown or gray, rarely darkening to black; epihymenium shades of gray, yellow,
or brown, negative or changing to orange-yellow in nitric acid......................................... 3
2.
Ascus tip with an K/I+ blue tholus; thallus pale orange or yellowish when fresh; northern arctic
.................................................................................................................... [Eiglera flavida]
2.
Ascus tip K/I–; thallus whitish to pale yellow-white, gray, or pinkish, often endolithic; arcticalpine to temperate ........................................................................................... [Hymenelia]
3.
Thallus pale orange to pale brown; common, Appalachian-Great Lakes-Pacific Northwest
distribution ................................................................................................ Ionaspis lacustris
3.
Thallus pale pinkish (when fresh) to yellowish white........................................................ 4
4.
On dry rocks, often partly shaded boulders; temperate to southern boreal; apothecia almost
white ............................................................................................ [Ionaspis alba Lutzoni]
4.
On moist or submerged rocks; boreal to arctic-alpine; apothecia pink to dark, rarely white
............................................................................................................................................ 5
5.
Apothecia pale pinkish (when fresh); spores 12.5-17 x 7-10 μm; usually on calcareous (rarely
siliceous) wet rocks ..........................................................................[Hymenelia epulotica]
5.
Apothecia smoky gray-brown to brown; spores 8-15 x 5-8 μm; on siliceous rocks in mountain
streams; western mountains ..........................................................................Ionaspis lavata
JAPEWIA: See Key F.
KAERNEFELTIA: See Keys A and K.
LASALLIA: See Umbilicaria.
LAURERA: See Trypethelium.
LECANACTIS: See Key F.
LECANIA (including Solenopsora)
1.
On rock............................................................................................................................... 2
1.
On bark............................................................................................................................... 6
125
2.
Spores (2-)4-celled, 12-16(-18) x 4-6 μm; apothecia dark brown to black, always pruinose; on
limestone, widespread ......................................................................[Lecania nylanderiana]
2.
Spores 2-celled; apothecia pale to dark brown or black, sometimes pruinose…………....3
3.
Thallus areolate to almost squamulose, or clearly lobed at the margins; on calcareous or siliceous
rocks; California................................................................................................................. 4
3.
Thallus rimose-areolate, not at all squamulose or lobed, pale greenish to gray; on calcareous rocks
............................................................................................................................................ 5
4.
Thallus brown, areolate almost squamulose; apothecia dark brown, not pruinose; on siliceous
rocks.......................................................................................................... Lecania brunonis
4.
Thallus white, lobed at the margins, not truly squamulose; apothecia almost black, lightly
pruinose; usually on calcareous rocks ........................................... [Solenopsora candicans]
5.
Thallus not at all sorediate; mostly in east ......................................... [Lecania perproxima]
5.
Thallus granular to sorediate; mostly in west and north............................[Lecania erysibe]
6.(2)
Spores 4-celled, 12-23 x 4-6 μm, straight or slightly bent, 8-16 per ascus;
central parts of continent ..........................................................................[Lecania fuscella]
6.
Spores 2-celled; widespread............................................................................................... 7
7.
Spores straight.......................................................................................... [Lecania cyrtella]
7.
Spores bent or bean-shaped....................................................................... Lecania dubitans
LECANOGRAPHA: See Keys E and F.
LECANORA
1.
Growing on bark, wood, mosses, or dead vegetation......................................................... 2
1.
Growing directly on rock ................................................................................................. 24
2.
Thallus within substrate, absent from view, or indistinct................................................... 3
2.
Thallus clearly visible ........................................................................................................ 6
3.
Spores ellipsoid to broadly ellipsoid, ratio of length to width 1.8-2.2:1; apothecial disks pruinose;
amphithecium filled with opaque crystals…………………………………..Lecanora hagenii
3.
Spores narrowly ellipsoid, ratio of length to width 2.2-3.0:1; apothecial disks pruinose or not;
amphithecium without crystals........................................................................................... 4
126
4.
Epihymenium clear red-brown, not at all granular...................................Lecanora zosterae
4.
Epihymenium with granules on the surface or between the tips of the paraphyses ........... 5
5.
Apothecia pale yellowish brown or reddish brown to black; margin paler than disk, rough; algae
filling amphithecial medulla.................................................................. Lecanora piniperda
[Note: L. piniperda has been synonymized under L. albellula Nyl. by Printzen (2001)]
5.
Apothecia yellow; margin disk-colored, smooth; algae very sparse in the apothecial margin,
sometimes essentially absent................................................................. Lecanora symmicta
6.(2)
Thallus leprose, greenish yellow or yellow-green, with a well-developed white, fibrous prothallus;
apothecia rare .................................................................................. Lecanora thysanophora
6.
Thallus not leprose, yellowish or gray, lacking a fibrous prothallus; apothecia produced on all
mature thalli ....................................................................................................................... 7
7.
Apothecia essentially biatorine, almost without algae in margins (only a few in apothecial base)
............................................................................................................................................ 8
7.
Apothecia lecanorine, with algae relatively abundant in margins...................................... 9
8.
Apothecia yellowish green to vivid yellow (containing usnic acid); spores narrowly ellipsoid;
apothecial margin PD–; widespread on bark or wood of all kinds, boreal to temperate
............................................................................................................... Lecanora symmicta
8.
Apothecia pale to very dark brown to black; spores almost spherical, 6-8 x 5.5-8 μm; apothecial
margin PD+ red (fumarprotocetraric acid); common on conifers in western mountains
........................................................................................................... [Lecanora fuscescens]
9.
Thallus and(or) apothecial margin yellowish, KC+ gold (usnic acid); cortex K– (atranorin absent);
spores narrowly ellipsoid, length to width ratio 2.2-3.2:1................................................ 10
9.
Thallus and apothecial margin cortex KC– or KC+ yellow, K+ yellow (atranorin); spores usually
ellipsoid to broadly ellipsoid or spherical, length to width ratio 1.0-2.2:1....................... 13
10.
Apothecial disks with a light to heavy yellow pruina; apothecial margin prominent, flexuose
..................................................................................................................Lecanora cupressi
10.
Apothecial disks not pruinose or lightly gray pruinose; apothecial margin flush with disk or
becoming thin and disappearing in maturity, not flexuose............................................... 11
11.
Apothecia yellow-brown to black, never yellowish green or greenish yellow, thallus very thin, pale
yellowish gray; apothecial margin more or less smooth, not sorediate; amphithecium filled with
127
algae .......................................................................................................Lecanora piniperda
11.
Apothecia usually yellow-green or greenish yellow; thallus pale yellowish green; apothecial
margin sorediate or granular; algae sparse or relatively abundant in amphithecium ....... 12
12.
Apothecial margin smooth, containing very few algae, never sorediate or granular
............................................................................................................... Lecanora symmicta
12.
Apothecial margin verruculose, granular or sorediate, with algae abundant at least where thalline
tissues of the margin are well developed…………………………………….Lecanora strobilina
13.(9)
Apothecia heavily pruinose making them pinkish or violet; amphithecial cortex lacking with
medullary hyphae growing out to the margin; amphithecial medulla PD+ yellow or red, rarely PD–
(protocetraric or virensic acid)………………………………………… Lecanora caesiorubella
13.
Apothecia usually yellowish brown to reddish brown without pruina, but if pruinose, then not
pinkish or violet; amphithecial cortex present, indistinctly or distinctly delimited from the
medulla; amphithecial medulla PD– or PD+ pale yellow ................................................ 14
14.
Amphithecium containing numerous small crystals; epihymenium red-brown, not at all granular
.......................................................................................................................................... 15
14.
Amphithecium containing a few large, irregular crystals; epihymenium granular on the surface or
between the tips of the paraphyses ................................................................................... 16
15.
Growing on logs and fences near the sea; amphithecial cortex distinct from the medulla, which is
filled with crystals ................................................................................. Lecanora xylophila
15.
Growing on bark; amphithecial cortex indistinctly delimited from medulla with crystals extending
from one into the other .......................................................................... Lecanora allophana
16.
Tiny brown granules, insoluble in HNO3, distributed in upper third of hymenium; apothecia not
pruinose............................................................................................................................ 17
16.
Coarse granules, soluble in HNO3, limited to the surface of the epihymenium, not extending into
the hymenium; apothecia pruinose or not ..................................................................... 18
17.
Spores 10-13 x 6-7.5 μm; amphithecial cortex not much wider at the sides than at the base of the
amphithecium; apothecial disks pale yellow-brown to reddish brown; eastern temperate deciduous
forests .................................................................................................. Lecanora hybocarpa
17.
Spores 13-17.5 x 8-11 μm, amphithecial cortex much wider at the base than at the sides of the
amphithecium; apothecial disks dark reddish brown to black; boreal forests
...................................................................................................... Lecanora circumborealis
128
18.
Apothecial disks C+ orange; southeast coastal plain.......................... [Lecanora louisianae]
18.
Apothecial disks C– ......................................................................................................... 19
19.
Apothecia immersed in the thallus for a long time, finally superficial............................. 20
19.
Apothecia mostly superficial with well-developed margins ............................................ 21
20.
Spores narrowly ellipsoid, 9-13 x 5-7 μm; apothecia small and very crowded, 0.3-0.8 mm in
diameter; southeast coastal plain.............................................................[Lecanora leprosa]
20.
Spores broadly ellipsoid, mostly 10-14.5 x 7-8.5 μm, apothecia not usually crowded together, 0.71.5 mm in diameter ...........................................................................Lecanora cinereofusca
21.
Apothecia margin cortex less than 15 μm thick; apothecial margin strongly verrucose or "beaded";
apothecia never pruinose; epihymenium PD+ orange (with orange crystals slowly developing, as
seen under the microscope) ……………………………………………Lecanora cinereofusca
21.
Apothecia margin cortex 20-50 μm thick; apothecial margin even or slightly bumpy; apothecia
often pruinose; epihymenium PD–................................................................................... 22
22.
Thallus very thin; apothecia thin & flat; Pacific Northwest…………………Lecanora pacifica
22.
Thallus thick, verrucose to granular; apothecia thick, widespread................................... 23
23.
On bark, rarely wood........................................................................... [Lecanora rugosella]
23
On hard weathered wood, never bark.........................................................Lecanora cenisia
24.(1)
Thallus endolithic, within the rock and out of sight, with only the apothecia on the surface
.......................................................................................................................................... 25
24.
Thallus clearly visible ...................................................................................................... 26
25.
On calcareous rock (bubbling with HCl); apothecial margins white, K–, KC–
..................................................................................................................Lecanora dispersa
25.
On siliceous rock (unreactive with HCl); apothecial margins yellowish, K–, KC+ gold
(usnic acid)........................................................................................... Lecanora polytropa
26.
On shoreline rocks; thallus yellowish, often lobed at the margin, cortex C+ orange, KC+
orange (xanthones) .......................................................................................................... 27
26.
On non-maritime rocks; thallus yellowish or not, lobed or not, cortex C–, KC– or KC+ gold
(xanthones absent)............................................................................................................ 28
129
27.
In California; apothecia yellowish, heavily pruinose ................................ Lecanora pinguis
27.
On northern Pacific coast; apothecia red-brown, not or slightly pruinose
.............................................................................................................[Lecanora straminea]
28.
Thallus distinctly lobed .................................................................................................... 29
28.
Thallus not lobed.............................................................................................................. 36
29.
Thallus cortex K+ blood red (norstictic acid)..................................................... Lobothallia
29.
Thallus cortex K–, or K+ persistently yellow .................................................................. 30
30.
Apothecia bright red-brown; epihymenium entirely without granules; thallus pale yellow to gray
..............................................................................................................Lecanora argopholis
30.
Apothecia yellow-brown, greenish, or black; epihymenium usually granular on surface; thallus
yellow-green, greenish yellow, or honey-brown.............................................................. 31
31.
Thallus honey-brown, often shiny; apothecia without pruina but rarely seen…………….32
31.
Thallus yellowish green or greenish yellow; apothecia with or without pruina, usually present and
abundant ........................................................................................................................... 33
32.
Low elevation woodlands of California; contains rangiformic acid; upper cortex containing dead
algal cells................................................................................................. [Lecanora mellea]
32.
High elevations in California, and scrublands in Pacific Northwest and arid interior; contains
protolichesterinic acid; upper cortex purely fungal, lacking dead algal cells
...................................................................................................... [Lecanora pseudomellea]
33.
Thallus lobes mostly flat .................................................................................................. 34
33.
Thallus lobes usually somewhat to strongly convex; western.......................................... 35
34.
Apothecia not pruinose; lobes usually with thickened or slightly raised margins making the lobe
tips somewhat concave; lobe margins usually whitish or pruinose; very widespread, especially on
rocks frequented by birds; medulla PD– ...................................................Lecanora muralis
34.
Apothecia heavily yellowish pruinose; lobes without raised margins, not at all concave; lobe
margins not white or especially pruinose; western; medulla PD+ yellow (psoromic acid)
........................................................................................................... [Lecanora bipruinosa]
35.
Apothecia black to yellowish green, typically heavily yellowish pruinose but sometimes without
130
pruina; lobes convex, but usually not strongly sinuous or folded, rarely pruinose
...................................................................................................... Lecanora novomexicana
35.
Apothecia not pruinose; thallus verrucose or rugose, or with sinuous folds, often pruinose
...............................................................................................................Lecanora garovaglii
36.(28) Ends of spores pointed; thallus grayish brown to yellow-brown, shiny; apothecia shiny dark redbrown ............................................................................................................. Protoparmelia
36.
Ends of spores rounded .................................................................................................... 37
37.
On calcareous rock........................................................................................................... 38
37.
On non-calcareous rock.................................................................................................... 40
38.
Apothecial disks heavily pruinose; apothecia concave; ascus tip K/I–
...................................................................................................................Aspicilia candida
38.
Apothecial disks not, or lightly, pruinose; apothecia flat, convex, or becoming hemispherical;
ascus tip K/I+ dark blue (Lecanora-type)......................................................................... 39
39.
Apothecial disks black or almost black; epihymenium green…………Lecanora marginata
39.
Apothecial disks red-brown; epihymenium brown ..............................Lecanora argopholis
40.(37) Thallus leprose, with a conspicuous white, fibrous prothallus; apothecia very rare
......................................................................................................... Lecanora thysanophora
40.
Thallus continuous or dispersed areolate, not leprose; prothallus absent; apothecia abundant
.......................................................................................................................................... 41
41.
Apothecial disks shiny red-brown, never pruinose; epihymenium clear red-brown, not at all
granular ............................................................................................................................ 42
41.
Apothecia yellowish to brown or black, dull, pruinose or not; epihymenium granular on the
surface or not granular ..................................................................................................... 43
42.
Thallus thick, areolate; apothecia 1-3 mm in diameter, constricted at the base; western
..............................................................................................................Lecanora argopholis
42.
Thallus thin, continuous; apothecia less than 1 mm in diameter, more or less immersed in the
thallus; East Temperate ...............................................................[Lecanora subimmergens]
43.
Thallus white to pale gray or greenish gray (lacking usnic acid); apothecia typically pruinose
.......................................................................................................................................... 44
131
43.
Thallus greenish yellow or yellowish green (containing usnic acid); apothecia with or without
pruina ............................................................................................................................... 46
44.
Apothecial disks C+ yellow, heavily white pruinose ..............................Lecanora rupicola
44.
Apothecial disks C–, pruinose or not .............................................................................. 45
45.
Apothecia superficial with well-developed margins; disks brown to black, often pruinose
....................................................................................................................Lecanora cenisia
45.
Apothecia immersed in thallus (cryptolecanorine), lacking margins; disks black, never pruinose
.............................................................................................................. Lecanora oreinoides
46.(43) Apothecia 2-7 mm in diameter, constricted at the base; disks shades of orange or pink, lightly
pruinose; thallus well-developed, dispersed areolate to almost squamulose
..................................................................................................... Rhizoplaca subdiscrepans
46.
Apothecia 0.3-0.8(-1.3) mm in diameter, sessile; disks pale yellow to pale orange, not pruinose;
thallus composed of scattered, very small, flat, adnate areoles ............ Lecanora polytropa
LECIDEA (including Carbonea, Fuscidea, Lecidella, Schaereria, and Tremolecia)
1.
On bark; ascus Lecanora-type ............................................................................................ 2
1.
On rock, wood, soil or peat; asci various types.................................................................. 3
2.
Paraphyses easily separating when mounted in water; hypothecium yellow-brown to colorless,
clearly distinguished from exciple, which is dark at outer edge; widespread arctic to temperate,
from Lake Superior and James Bay westward .................................... [Lecidella euphorea]
2.
Paraphyses sticking together when mounted in water; hypothecium dark purplish brown to black,
merging with exciple which is dark internally and almost colorless externally; boreal forest region
and west coast…………………………………………………………..["Lecidea" albofuscescens]
3.
On wood, soil, or mosses ................................................................................................... 4
3.
Directly on rock.................................................................................................................. 6
4.
Thallus dark brown or olive brown, granular to almost isidiate, K–, PD–, C+ pink (gyrophoric
acid) or C–; apothecia dull red-brown to black, flat to strongly convex; paraphyses branched;
hypothecium merging with exciple, dark brown..............................................Placynthiella
4.
Thallus whitish, extremely thin and often within substrate, or more or less verrucose, K+ yellow,
C– (atranorin); apothecia black, margins thin to disappearing; paraphyses unbranched;
hypothecium colorless or brown ........................................................................................ 5
132
5.
On wood, temperate (mostly northeastern, Black Hills, Colorado); apothecia shiny, flat to convex;
exciple dark brown at edge, pale internally; hypothecium pale yellowish brown or colorless;
spores narrow, 8-10(-12) x 3-4 μm; thallus PD+ red (fumarprotocetraric acid)
.......................................................................................................... [Pyrrhospora elabens]
5.
On soil and moss, arctic and alpine tundra; apothecia dull, soon convex; exciple brown or greenish
at edge, brown internally; hypothecium red-brown; spores broad, 10-16 x 7-8 μm; thallus PD–
...............................................................................................................[Lecidella wulfenii]
6.
Cephalodia present ...........................................................................................Amygdalaria
6.
Cephalodia absent .............................................................................................................. 7
7.
Thallus distinctly rusty orange or bright orange................................................................. 8
7.
Thallus not bright orange ................................................................................................. 10
8.
Apothecia concave, immersed in thallus, usually less than 0.5 mm in diameter
...................................................................................................................Tremolecia atrata
8.
Apothecia flat when mature, sessile, usually more than 0.5 mm in diameter .................... 9
9.
Spores 14-24 x 6-11 μm, halonate (visible at least in ink preparation); epihymenium olive-brown;
paraphyses abundantly branched; medulla IKI–……………………Porpidia flavocaerulescens
9.
Spores 9-15 x 4.5-8 μm, without a halo; epihymenium green; paraphyses mostly unbranched
except for tips; medulla IKI+ blue………………………………orange form of Lecidea lapicida
10.(7)
Thallus red-brown to dark yellow (not rusty orange), shiny, thick, areolate to dispersed areolate
over the black prothallus, sometimes appearing almost squamulose ............................... 11
10.
Thallus gray to white or greenish gray to brownish gray, continuous or areolate, not appearing
squamulose, sometimes disappearing (endolithic) ........................................................... 12
11.
Thallus reddish brown; medulla IKI+ blue; apothecia immersed between the areoles; ascus tips
K/I- (Lecidea-type)...............................................................................Lecidea atrobrunnea
11.
Thallus dark yellow to bronze; medulla IKI-; apothecia superficial, not between areoles; ascus tips
K/I+ blue (Bacidia-type) .................................................................. Tephromela armeniaca
12.
Hypothecium brown to black, sometimes greenish black ................................................ 13
12.
Hypothecium colorless to yellowish ................................................................................ 18
133
13.
Spores under 3.5 μm wide; spores not halonate ............................................................... 14
13.
Spores over 3.5 μm wide; spores halonate or not............................................................. 15
14.
Apothecia 1-2.5 mm in diameter; thallus usually endolithic and not visible; hymenium under 50
μm high; on alpine and arctic rocks…………………………………..[Lecidea auriculata]
14.
Apothecia usually under 0.5 mm in diameter; thallus thin and indistinct or rimose areolate and
dark greenish gray; hymenium 50-60(-80) μm; typically on boulders, small stones, and pebbles;
widespread, East Temperate………………………………………………….[Micarea erratica]
15.
Spores narrow, 8.5-15 x 3.5-5.5 μm. Thallus thin, rimose, or endolithic; apothecia flat, 0.3-1 mm
in diameter, with thin, prominent rims; epihymenium and upper part of hymenium emerald green;
hypothecium and exciple black, not distinguishable; mostly western arctic and Rocky Mountains
............................................................................................................. [Carbonea vorticosa]
15.
Spores more than 5 μm wide............................................................................................ 16
16.
Paraphyses branched and anastomosing, coherent in a water mount; spores halonate, (10-)13-24 x
6-12 μm ............................................................................................................ Porpidia key
16.
Paraphyses mostly unbranched, easily separated in a water mount; spores not halonate, (8-)10-16 x
6-9 μm .............................................................................................................................. 17
17.
Thallus epilithic and visible or largely endolithic and hidden; medulla (between rock
crystals, if endolithic) IKI+ blue ......................................................... Lecidea lapicida
17.
Thallis epilithic, clearly visible; medulla IKI- .......................................................... 17a
17a.
Thallus white, areolate to verrucose, K+ yellow, C– (atranorin); widespread .....................
............................................................................................................[Lecidella carpathica]
17a.
Thallus pale yellowish to greenish yellow, rimose to areolate; on rocks along the Pacific coast
...................................................................................................[Lecidella eleochromoides]
18.(12) Epihymenium and exciple shades of brown; asci Fuscidea-type (fig. 14d); thallus gray-brown or
dark gray, often forming quilt-like patches outlined by black................................ Fuscidea
18.
Epihymenium and often the outer layer of the exciple shades of green; asci not Fuscidea-type;
thallus pale to dark gray, not brownish ............................................................................ 19
19.
Paraphyses easily separating in water or K; spores 5-9(-10) μm wide ............................ 20
19.
Paraphyses coherent in water and K; spores less than 7.5 μm wide................................. 22
20.
Apothecia remaining immersed between thallus areoles for a long time; thallus C+ pink
134
(gyrophoric acid); asci cylindrical, thin-walled including tip, K/I–
...................................................................................................... [Schaereria fuscocinerea]
20.
Apothecia superficial, not immersed; thalus C– or C+ orange; asci club-shaped with a thick tip
that is K/I+ dark blue........................................................................................................ 21
21.
Hymenium containing abundant tiny oil drops; western montane to arctic, on siliceous rocks
...............................................................................................................[Lecidella patavina]
21.
Hymenium without oil drops; widespread temperate to arctic, mostly on calcareous rocks
............................................................................................................... Lecidella stigmatea
22.(19) Prothallus lacking or inconspicuous; thallus gray to orange or mottled with orange areas,
continuous or, in part, cracked into areoles................................................ Lecidea lapicida
22.
Prothallus black, conspicuous .......................................................................................... 23
23.
Thallus white to pale gray, tile-like (areolate) with the black prothallus showing at the margin;
medulla IKI+ blue, K–; asci K/I– (Lecidea-type)…………………………..Lecidea tessellata
23.
Thallus whitish to pale yellow, continuous to areolate, but not tile-like; medulla IKI–, K+ yellow
(atranorin and sometimes usnic acid); asci K/I+ blue at tips (Lecanora-type)
...............................................................................................................Lecanora marginata
LECIDELLA: See Lecidea.
LECIDOMA: See Key F.
LEIODERMA: See Peltigera.
LEPRARIA (including Leprolomma)
1.
Thallus forming round, adnate or somewhat ascending lobes 3-6 mm across; brownish or white
cottony hypothallus present; containing pannaric and roccellic acids; on shaded rock faces
.............. Lepraria membranacea (Dickson) Vainio (syn. Leproloma membranaceum)
1.
Thallus not forming well developed, ascending, round lobes; hypothallus present or absent;
pannaric acid absent; on rock or other substrates ............................................................... 2
2.
Thallus coarsely granular sorediate, not cottony; margins irregular or, when on rocks, forming
distinct, often concentric rings about 20-40 mm in diameter, pale to dark gray; hypothallus entirely
absent; on siliceous rocks or moss; thallus K–, PD+ yellow or red (alectorialic, fumarprotocetraric,
or psoromic acid), lacking atranorin..........................................................Lepraria neglecta
135
2.
Thallus thin or thick, made up of fine, farinose soredia, never forming concentric rings; on
shaded rock, bark, or moss; containing atranorin and other compounds .................. 3
3.
Thallus thick and cottony, occasionally lobed, usually pale green to yellowish green, rarely
greenish gray; brownish hypothallus frequently present; thallus K+ yellow, PD+ orange
(atranorin, stictic acid, and zeorin). Widespread…………………………Lepraria lobificans
3.
Thallis thin, not at all cottony, gray to blue-gray, without a hypothallus; Thallus K+
yellowish, PD- (atranorin and zeorin). Eastern ........ [Lepraria caesiella R. C. Harris]
LEPROCAULON: See Key B.
LEPROLOMA: See Lepraria.
LEPTOCHIDIUM: See Leptogium.
LEPTOGIUM (including Leptochidium and Hydrothyrea)
1.
Aquatic, on submerged rocks in the western mountains .................................................... 2
1.
In dry or moist habitats, not submerged in water ............................................................... 3
2.
Lobes 3-10 mm broad, ascending and fan-like .................................... Hydrothyrea venosa
2.
Lobes 0.2-1.5 mm broad, closely attached to rocks ............................... [Leptogium rivale]
3.
Lower surface tomentose with a mat of white hairs........................................................... 4
3.
Lower surface smooth, or wrinkled, not hairy (except for scattered tufts in Leptogium platynum)
.......................................................................................................................................... 11
4.
Isidia and lobules absent; Arizona and New Mexico ........................................................ 5
4.
Isidia or lobules present on upper surface or margins of lobes .......................................... 6
5.
Surface conspicuously wrinkled........................................................... Leptogium rugosum
5.
Surface smooth.................................................................................. [Leptogium burgessii]
6.
Thallus lobulate, not isidiate .............................................................................................. 7
6.
Thallus isidiate, but isidia can be somewhat flattened in some species ............................. 8
7.
Lobules confined the lobe margins; tomentum white, conspicuous, well over 100 μm long,
composed of threads with long, cylindrical cells; southwestern
136
........................................................................................................... [Leptogium burgessii]
7.
Lobules on lobe surface as well as margins; tomentum consisting of a faint fuzz made up of
threads less than 100 μm long with spherical cells; Appalachians to Ontario and New Brunswick
....................................................................................................................... [L. laceroides]
8.
Fine, stiff, colorless hairs on the tips or upper surface of the lobes; isidia flattened and lobelike
when older, often bearing fine hairs…………………………………Leptochidium albociliatum
8.
Fine hairs absent or sparse and confined to lobe surface; isidia cylindrical to granular, never
flattened or hairy ................................................................................................................ 9
9.
Surface of lobes smooth, not wrinkled; isidia granular or cylindrical and branched ....... 10
9.
Surface of lobes clearly wrinkled; isidia cylindrical or slightly thicker at the tips
................................................................................................. Leptogium furfuraceum
10.
Thallus dark olive-brown or olive-gray when dry, never with fine colorless hairs; isidia mostly
granular; common ............................................................................ Leptogium saturninum
10.
Thallus slate gray, without an olive tint when dry, sometimes with scattered fine hairs on the
surface; isidia mostly cylindrical; uncommon………………………..[Leptogium burnetiae]
11.(3)
Thallus slate gray ........................................................................................................... 12
11.
Thallus brown at least when dry, or olive, black, or greenish gray.................................. 22
12.
Isidia present, cylindrical or, in part, flattened, resembling narrow lobules; apothecia abundant or
rare ................................................................................................................................... 13
12.
Isidia absent (lobules present or absent); apothecia usually present and abundant .......... 15
13.
Upper surface of lobes smooth and even ......................................... Leptogium cyanescens
13.
Upper surface of lobes wrinkled ..................................................................................... 14
14.
Southeastern; isidia flattened to cylindrical; thallus less than 200 μm thick
.............................................................................................. Leptogium austroamericanum
14.
Western montane; isidia cylindrical; thallus more than 200μm thick
.............................................................................................................. [Leptogium arsenei]
15.(12) Lobes curled inward and erect, forming tubelike tips……………. Leptogium corniculatum
15.
Lobes flat, convex, concave, or undulating and crisped at the margins, not forming tubes .. 16
137
16.
Apothecial margins with abundant lobules; apothecia restricted to the thallus margins ......
........................................................................................................Leptogium marginellum
16.
Apothecial margins smooth and even or thickly wrinkled, with or without lobules;
apothecia on the lobe surface .......................................................................................... 17
17.
Apothecia constricted at the base and somewhat raised .................................................. 18
17.
Apothecia broadly attached, adnate or sunken into depressions ..................................... 19
18.
Lobes strongly wrinkled ..................................................................... Leptogium corticola
18.
Lobes not wrinkled.............................................................................. Leptogium azureum
19.
Apothecia 2 - 7 mm in diameter, margins thickly wrinkled; southeastern USA
..................................................................................................... Leptogium phyllocarpum
19.
Apothecia 0.2 - 0.7 mm in diameter, margins smooth and even; western USA and Canada
.......................................................................................................................................... 20
20.
Rounded, overlapping lobules on thallus surface; thallus 200-500 μm thick, often attached to
substrate by tufts of pale hairs............................................................. Leptogium platynum
20.
Finely divided lobules on the lobe margins, or margins fairly even; thallus thin and
membranous, less than 200 μm thick, attached directly to substrate, lacking hairs ....... 21
21.
Ascospores 4 per ascus; on bark ..................................................... Leptogium polycarpum
21.
Ascospores 8 per ascus; on mossy rock, soil, or bark .....................Leptogium gelatinosum
22.(11) Isidia present .................................................................................................................... 23
22.
Isidia absent ..................................................................................................................... 24
23.
On mossy calcareous rock; lobe margins finely divided, resembling isidia (fig. 32); apothecia
often abundant................................................................................. Leptogium lichenoides
23.
On bark; lobe margins scalloped; granular to cylindrical isidia on lobe margins and ridges;
apothecia infrequent ...................................................................... Leptogium milligranum
24.
Lobes curled inward, forming a tube ............................................ Leptogium corniculatum
24.
Lobes flat, concave, or undulating and crisped at the margins......................................... 25
25.
Apothecia raised, especially on the crests of ridges and folds; mainly in the southeastern coastal
plain ..............................................................................................Leptogium chloromelum
138
25.
Apothecia adnate or sunken into depressions; mainly western and northern…………….26
26.
Rounded, overlapping lobules on thallus surface; thallus over 200 μm thick, often attached to
substrate by tufts of hairs .................................................................... Leptogium platynum
26.
Lobules, if present, marginal; thallus thin and membranous, less than 200 μm thick,
lacking hairs on the lower surface................................................................................... 27
27.
Margins very finely divided (fig. 32); thallus cushion forming; widespread, especially among
mosses over limestone......................................................................Leptogium lichenoides
27.
Margins usually smooth and even, or somewhat lobulate; Pacific Northwest…………. 28
28.
Ascospores 4 per ascus; on bark ..................................................... Leptogium polycarpum
28.
Ascospores 8 per ascus; on mossy rock, soil, or bark .....................Leptogium gelatinosum
LETHARIA
1.
Branches granular sorediate; apothecia rare………………………………………..Letharia vulpina
1.
Branches without soredia; apothecia abundant………………………………...Letharia columbiana
LETROUITIA: See Key F.
LICHINELLA: See Keys A and I.
LOBARIA
1.
Photobiont blue green; cephalodia absent ...................................................................................................... 2
1.
Photobiont green; cephalodia present.............................................................................................................. 3
2.
Stiff, tiny, colourless hairs on the lobe tips or on the upper surface of the lobes; rhizines rope-like,
covered with perpendicular hairs; medulla PD–, K– (norstictic and stictic acids absent); infrequent,
in Pacific Northwest.................................................................................................................... Lobaria hallii
2.
Stiff, tiny, colourless hairs absent; rhizines tufted; medulla PD+ yellow, K+ red (norstictic and
stictic acids present); widespread and frequent ................................................................ Lobaria scrobiculata
3.
Lobes with a network of depressions and ridges; thallus loosely attached; medulla KC–, C–
(gyrophoric acid absent).................................................................................................................................. 4
3.
Lobes relatively smooth and even, without a network of ridges and depressions; thallus closely
appressed to substrate; medulla KC+ red, C+ pink (gyrophoric acid)……………………………5
139
4.
Soredia, often mixed with isidia, present on ridges and lobe margins; medulla PD+ yellow to
orange, K+ yellow to red (norstictic and stictic acids) .......................................................Lobaria pulmonaria
4.
Soredia and isidia absent; medulla PD–, K– ............................................................................... Lobaria linita
5.
Lobes 5-20 mm broad; Appalachian-Great Lakes region ...................................................Lobaria quercizans
5.
Lobes 2-4(-6) mm broad; southeastern coastal plain....................................................................................... 6
6.
Lobules abundantly produced on upper surface and margins of lobes; cephalodia always small and
inconspicuous warts, seen best on lower thallus surface; pycnidia and apothecia rare
……………………………………………………………………………….Lobaria tenuis
6.
Lobules rarely present; cephalodia sometimes forming small branched outgrowths on the upper
thallus surface; pycnidia appearing as abundant black dots on the lobe surface; apothecia abundant
……………………………………………………………………………Lobaria ravenelii
LOBOTHALLIA
1.
Thallus yellowish to copper brown; lobes flat and appressed to rock……..Lobothallia praeradiosa
1.
Thallus shades of gray, or, less frequently, light brown; lobes very convex, almost foliose in
appearance…………………………………………………………………..Lobothallia alphoplaca
LOPADIUM: See Key F.
LOXOSPORA
1.
Apothecia rare; thallus sorediate or producing masses of small, hollow pustules that break down
into granular schizidia or soredia…………………………………………………2
1.
Apothecia common, lecanorine, often with a torn or ragged margin; thallus without hollow
pustules or soredia………………………………………………………………………………..3
2.
Thallus with abundant farinose to granular soredia in the older central parts; pustules small
if present, forming true soredia; almost exclusively on conifers; boreal… [Loxospora elatina]
2.
Thallus without true soredia, only with conspicuous pustules that break down into granular
schizidia; almost exclusively on decisduous trees; temperate ……………Loxospora pustulata
3.
Apothecial disks red-brown, not pruinose or lightly pruinose; thallus rather thick and irregular;
apothecia constricted at base………………………………………………Loxospora ochrophaea
3.
Apothecial disks purplish, heavily pruinose, thallus very thin, more or less smooth; apothecia
sessile………………………………………………………………………..Loxospora cismonica
140
LOXOSPOROPSIS: See Keys B, C, and F.
MASONHALEA: See Key A and Cetraria.
MASSALONGIA: See Pannaria.
MEGALARIA: See Key F.
MEGASPORA: See Key F.
MELANELIA (including Neofuscelia)
1.
Soredia present ................................................................................................................................................ 2
1.
Soredia absent ................................................................................................................................................. 9
2.
Medulla C+ red or pink .................................................................................................................................. 3
2.
Medulla C–...................................................................................................................................................... 6
3.
Pseudocyphellae abundant and conspicuous; lobes elongated; gyrophoric acid present; white or
brown soralia laminal or partly marginal; soredia granular...................................................Melanelia tominii
3.
Pseudocyphellae absent; lobes rounded; lecanoric acid present...................................................................... 4
4.
Soredia at least partly on thallus surface (laminal); soredia granular, often mixed with or arising
from isidia ....................................................................................................................................................... 5
4.
Soredia entirely marginal, farinose, forming powdery crescents; isidia absent
…………………………………………………………………………Melanelia albertana
5.
Soredia entirely laminal, arising from a disintegration of the cortex or isidia, leaving yellowish
green patches; thallus surface not pruinose, without cortical hairs…………..Melanelia subaurifera
5.
Soredia both laminal and marginal, arising from pustules or breakdown of the cortex, brown to
whitish; thallus surface commonly pruinose, and often with a fuzz of minute, colorless hairs on the
lobe tips …………………………………………………………………..Melanelia subargentifera
6.(2)
Soredia mostly on upper surface of lobes, dark brown (or whitish where abraded); rhizines
abundant, brown or black; lower surface dark brown or black; medulla UV+ blue-white (perlatolic
acid)................................................................................................................................................................. 7
6.
Soredia mostly marginal; rhizines sparse, very pale beige or tan; lower surface pale brown;
medulla UV– (perlatolic acid absent).............................................................................................................. 8
141
7.
Soredia fine, in rounded mounds mostly near the lobe tips............................................... Melanelia sorediata
7.
Soredia coarse, granular, in irregular patches on the older parts of the thallus…Melanelia disjuncta
8.
On bark and wood, rarely rock, in western North America; medulla K–, PD– (fatty acids); lobes
concave, undulating or crisped at the margins .................................................Tuckermannopsis chlorophylla
8.
On rock in the Appalachian region; medulla K+ red, PD+ yellow (norstictic and stictic acids);
lobes flat........................................................................................................................ Melanelia culbersonii
9.(1)
Isidia present ................................................................................................................................................. 10
9.
Isidia absent................................................................................................................................................... 18
10.
Lobes 0.4-0.7 mm wide; rhizines mostly marginal; on wood or bark .................Tuckermannopsis coralligera
10.
Lobes mostly more than 1 mm wide; rhizines all over lower surface; on rock, wood, or bark..................... 11
11.
Isidia spherical or globular............................................................................................................................ 12
11.
Isidia cylindrical, flattened, club-shaped, or conical, unbranched or branched ............................................ 14
12.
On rock; isidia hollow................................................................................................................................... 13
12.
On bark or wood, rarely rock; isidia solid....................................................................... Melanelia elegantula
13.
Medulla K–, PD–, KC+ pinkish violet (perlatolic acid)....................................................Neofuscelia loxodes
13.
Medulla K+ yellow or red, PD+ orange, KC– (unidentified substance)…..Neofuscelia subhosseana
14.(11) Medulla C+ red or pink (lecanoric acid) ....................................................................................................... 15
14.
Medulla C– (lecanoric acid absent)............................................................................................................... 16
15.
Isidia unbranched, mostly shorter than 0.2 mm; abrading to leave yellowish green patches
…………………………………………………………………………….Melanelia subaurifera
15.
Isidia frequently branched and longer than 0.2 mm; not forming yellowish patches where abraded
………………………………………………………………………..Melanelia fuliginosa
16.
Isidia cylindrical or conical when mature (Fig. 33a); upper surface dull .........................Melanelia elegantula
16.
Isidia flattened or club-shaped when mature; upper surface shiny or dull .................................................... 17
17.
Growing on rock; isidia solid; medulla UV+ blue-white (perlatolic acid) .....Melanelia panniformis
142
17.
Growing on bark or wood, rarely on rock; isidia solid or hollow; medulla UV–. ..……………17a
17a.
Isidia hollow, club-shaped to flattened, but rarely branched or cylindrical and never
bearing rhizines (Fig. 33b); growing on bark, rarely wood or rock; widespread in the west
………………………………………................………………………… Melanelia exasperatula
17a.
Isidia solid, cylindrical when young but soon becoming flattened and lobe-like, sometimes
branched, developing tiny rhizines on one side (Fig. 33d); growing exclusively on bark and
wood, central California to southern B.C. east to the Canadian Rockies
..................................................................................................................[Melanelia subelegantula]
18.(9)
Growing on bark ........................................................................................................................................... 19
18.
Growing on rock ........................................................................................................................................... 25
19.
Medulla C+ red, KC+ red (lecanoric acid); California........................................................... Melanelia glabra
19.
Medulla C–, KC– .......................................................................................................................................... 20
20.
Rhizines sparse, mostly marginal and resembling cilia; lobes 0.5-1.6 mm wide
................................................................................................................................. Tuckermannopsis fendleri
20.
Rhizines abundant all over lower surface, never resembling cilia; lobes (0.5-)1-5 mm wide....................... 21
21.
Medulla PD+ red (fumarprotocetraric acid) .................................................................................................. 22
21.
Medulla PD– ................................................................................................................................................ 23
22.
Pseudocyphellae sparse or inconspicuous on the thallus and apothecial margins; lobes smooth and
even ........................................................................................................................... Melanelia septentrionalis
22.
Pseudocyphellae abundant and conspicuous especially on the apothecial margins; lobes wrinkled
or bumpy (rugose) ...............................................................................................................Melanelia olivacea
23.
Spores 12-32 per ascus; lobes often with lobules, at least in center of thallus ............. Melanelia multispora
23.
Spores 8 per ascus; lobules absent ................................................................................................................ 24
24.
Pseudocyphellae absent or rare; western....................................................................... Melanelia subolivacea
24.
Pseudocyphellae on apothecial margins, lobes, and on thallus warts; East Temperate
.......................................................................................................................................[Melanelia exasperata]
25.(18) Pseudocyphellae abundant and conspicuous; rhizines mostly on or close to the margins………...26
25.
Pseudocyphellae absent or inconspicuous; rhizines more or less uniformly distributed………….28
143
26.
Pseudocyphellae marginal; pycnidia prominent, on lobe margins; medulla PD+ orange, K+ yellow
(stictic acid).......................................................................................................................Melanelia hepatizon
26.
Pseudocyphellae laminal; pycnidia scattered on lobe surface, entirely buried in thallus with just the
ostiole showing; medulla PD– or PD+ red, K– (stictic acid absent)………………………………27
27.
Medulla C+ pink, KC+ red, PD– (gyrophoric acid); thallus thick and stiff; pycnidia sparse or very
inconspicuous; conidia rod-shaped .......................................................................................Melanelia tominii
27.
Medulla C–, KC–, PD+ red or PD– (gyrophoric acid absent; fumarprotocetraric acid sometimes
present); thallus moderately thick; pycnidia abundant and conspicuous; conidia dumbbell-shaped
.................................................................................................................................................Melanelia stygia
28.(25) Thallus brown to olive-brown, usually producing abundant, overlapping, flattened lobules; lower
surface wrinkled; conidia rod-shaped............................................................................ Melanelia panniformis
28.
Thallus reddish brown or yellowish brown, without abundant, overlapping lobules; lower surface
smooth; conidia dumbbell-shaped................................................................................................................. 29
29.
Lower surface and rhizines dark brown or black; pycnidia sparse or very inconspicuous; medulla
K–, PD–, KC+ pinkish violet (perlatolic acid)…………………………………Neofuscelia loxodes
29.
Lower surface and rhizines pale brown; pycnidia abundant and conspicuous; medulla K+ yellow
or red-orange, PD+ orange, KC– (stictic and sometimes norstictic acids) …Neofuscelia atticoides
MENEGAZZIA: See Hypogymnia.
MICAREA: See Key F.
MULTICLAVULA: See Key to keys.
MYCOBILIMBIA: See Key F.
MYCOBLASTUS: See Keys B and F.
MYELOCHROA: See Parmelia.
MYRIOTREMA (based on Harris, More Florida Lichens, 1995)
1.
Spores colorless, muriform or transversely septate ...........................Myriotrema rufigerum
1.
Spores brown, muriform .................................................................................................... 2
144
2.
Spores 45-55 x 10-13 μm, many-celled. Ascomata round, not crowded; medulla PD+ orange, K+
yellow (stictic acid complex)………………………………[Myriotrema subcompunctum]
2.
Spores under 30 μm long, few-celled................................................................................. 3
3.
Ascomata round, with pore-like openings; thallus with a cortex; spores 14-28 x 9-14 μm; medulla
PD–, K–, but containing pockets of red crystalline pigment (K+ purple)
.............................................................................................................[Myriotrema wightii]
3.
Ascomata angular, with fissure-like openings, often in whitish areas without a cortex; spores 1316 x 7-10 μm; medulla PD+ orange, K+ yellow (stictic acid complex)
.....................................................................................................[Myriotrema glaucescens]
NEOFUSCELIA: See Melanelia.
NEPHROMA
1.
Photobiont green; cephalodia present; on the ground and mossy rocks and logs mostly in northern
boreal, arctic, and alpine habitats ....................................................................................... 2
1.
Photobiont blue green; cephalodia absent; on mossy bark or rock .................................... 3
2.
Thallus pale green, becoming browned especially at the margins; cortex KC– (usnic acid absent);
lobes undulating or crisped at the margins; cephalodia forming small, inconspicuous warts on
lower thallus surface; lobules present ...............................................Nephroma expallidum
2.
Thallus yellow-green; cortex KC+ orange-yellow (usnic acid); lobes flat; cephalodia forming
internal swellings visible externally as rounded dark bumps on the thallus surface; lobules absent
...............................................................................................................Nephroma arcticum
3.
Soredia present ................................................................................................................... 4
3.
Soredia absent .................................................................................................................... 5
4.
Thallus brown; soredia mainly on the lobe margins and sometimes on the surface; lobes mostly
smooth; widespread, common....................................................................Nephroma parile
4.
Thallus yellow-green (with usnic acid); soredia mainly on the thallus surface; lobes with a
network of depressions and sharp ridges; western, rare………………..Nephroma occultum
5.
Isidia present ...................................................................................................................... 6
5.
Isidia absent........................................................................................................................ 7
6.
Isidia cylindrical, unbranched or branched; lobes with a network of depressions and sharp ridges,
145
or wrinkled; lobules absent; apothecia absent……………………………Nephroma isidiosum
6.
Isidia flattened; lobes smooth and even; lobules present; apothecia typically abundant......
........................................................................................................... Nephroma helveticum
7.
Medulla yellow, K+ red or deep purple (anthraquinones)……………….Nephroma laevigatum
7.
Medulla white, K– (anthraquinones absent)....................................................................... 8
8.
Lower surface tomentose, with scattered pale bumps………………Nephroma resupinatum
8.
Lower surface smooth ..............................................................................Nephroma bellum
NIEBLA: See Ramalina.
NODOBRYORIA: See Alectoria.
NORMANDINA: See Key G.
OCELLULARIA: See Key D.
OCHROLECHIA
[Note: all spot tests with C must be done on thick sections of the apothecia or thallus, not on the thallus
surface.]
1.
Growing on rock, soil, peat, or moss.................................................................................. 2
1.
Growing on bark or wood .................................................................................................. 5
2.
On rock in temperate eastern United States; thallus thick, with thick isidia
............................................................................................................. Ochrolechia yasudae
2.
On soil, peat, or moss; western or arctic-alpine; isidia absent............................................ 3
3.
Thallus cortex and medulla C- (lacking gyrophoric acid); apothecia abundant, disks coarsely
pruinose-scabrose, C- or C+yellow ................................................Ochrolechia upsaliensis
3.
Thallus cortex and often medulla C+ pink (gyrophoric acid); apothecia abundant or rare, disks
smooth or cracked, not pruinose or scabrose, C+ pink…………………………………….4
4.
Thallus with round to irregular patches of yellowish granular soredia; thallus never
producing elongate, fruticose filaments at the edges; apothecia extremely rare
......................................................................................................... Ochrolechia androgyna
4.
Thallus without soredia, or rarely with irregular patches of whitish soredia; thallus frequently
producing elongate, fruticose filaments at the edges; apothecia frequent .......................
............................................................................................................... Ochrolechia frigida
146
5.(1)
Thallus sorediate ............................................................................ Ochrolechia androgyna
5.
Thallus without soredia ..................................................................................................... 6
6.
Eastern, but some species occasionally in the southwest ................................................... 7
6.
Western, California to Alaska, absent in the southwest ................................................... 11
7.
Thallus thick, more or less covered with thick isidia .......................... Ochrolechia yasudae
7.
Thallus thin or thick, smooth to verrucose, but not isidiate ............................................... 8
8.
Thallus and medulla C–; on conifer bark and wood; apothecia large and flat, 1.3-5 mm in
diameter, with pruinose or rough, scabrose disks; very common
............................................................................................[Ochrolechia pseudopallescens]
8.
Thallus cortex or medulla C+ pink or red; mostly on deciduous trees; apothecia 0.6-2(-3) mm
in diameter, pruinose or not, rough or smooth ................................................................... 9
9.
Spores 4-8 per ascus, under 70 μm long ...................................................................... 9a
9.
Spores 1 per ascus, over 70 μm long .................................................... Pertusaria velata
9a.
Cortex of apothecial margin C–, medulla C+ red; apothecia usually pruinose; apothecial
margins smooth and even.................................................................... Ochrolechia africana
9a.
Cortex of apothecial margin C+ pink, medulla C–; apothecia with or without pruina;
apothecial margins smooth or rough ................................................................................ 10
10.
Thallus UV– (lacking lichexanthone); apothecia usually without pruina; apothecial margins
verrucose .......................................................................................Ochrolechia trochophora
10.
Thallus UV+ yellow (lichexanthone); apothecia pruinose or without pruina; apothecial
margins usually smooth.................................................................. [Ochrolechia mexicana]
11.(6)
Cortex of thallus and apothecial margin C– (containing variolaric acid)………………..12
11.
Cortex of thallus and apothecial margin C+ pink to red (lacking variolaric acid) .......... 13
12.
Apothecial disks C+ pink (scratch the surface of the disk before testing), remaining small and
pore-like except in oldest apothecia, which become broad; thallus thin or well developed and
verruculose .........................................................................................Ochrolechia juvenalis
12.
Apothecial disks C–, broadening early in their development; thallus extremely thin
.....................................................................................................[Ochrolechia szatalaënsis]
13.
Thallus very thin and smooth; apothecial margins smooth, lacking a pinkish ring next to the
disk, containing few algae in the amphithecium; on deciduous trees, especially alder
............................................................................................................Ochrolechia laevigata
147
13.
Thallus thick and often verrucose, or rather thin and verruculose, rarely smooth; apothecial
margins smooth to verrucose, often or always with an inner ring of pinkish tissue next to the
disk................................................................................................................................... 14
14.
Algae abundant in the apothecial margin, but absent or spotty below the hypothecium; inner
ring always present on apothecial disks; hymenium 320-410 μm high; on conifer bark and
wood.............................................................................................. Ochrolechia oregonensis
14.
Algae forming a continuous layer under the hypothecium, and also present in the apothecial
margin; inner ring sometimes present on apothecial disks; hymenium 180-280 μm high;
mostly on deciduous trees, rarely on conifers………………….[Ochrolechia subpallescens]
OMPHALINA: See Key to keys, and Key G.
OMPHALORA: See Key H.
OPEGRAPHA
1.
On calcareous rock. Spores 4-celled, 22-29 x 5.5-8 μm; thallus thin or endolithic, often
parasitic on other lichens, C–; California, Alaska and Newfoundland
............................................................................................................[Opegrapha rupestris]
1.
On bark or wood................................................................................................................. 2
2.
Spores 4-celled ................................................................................................................... 3
2.
Spores 6- or more celled..................................................................................................... 5
3.
Spores (17.5-)19-25 x 5.5-10 μm. Lirellae fusiform, straight or curved, sometimes pruinose;
thallus creamy white to pale brown, forming small discrete patches; western, on bark and
wood of various kinds………………………………………………[Opegrapha herbarum]
3.
Spores under 5 μm wide; widely distributed ...................................................................... 4
4.
Thallus white in delimited patches; lirellae long and narrow with thick walls, opening by a
narrow slit; spores 13-18 (-20) μm long.................................................... [Opegrapha atra]
4.
Thallus brownish and indistinct, mostly within the bark; lirellae short, frequently branched or
star-shaped, scattered, with thin walls, opening broadly; spores 15-27 μm long
.......................................................................................................... [Opegrapha rufescens]
5.(2)
Spores 12- to 16-celled, 25-60 x 6-9 μm. Lirellae short and broad, elliptical, rarely forked;
widely distributed.................................................................................. [Opegrapha viridis]
5.
Spores 6- to 8-celled........................................................................................................... 6
148
6.
Lirellae with a red-brown surface, long, branched, opening broadly with a very thin black
margin that is sometimes hard to see; spores 16-21 x 4-6 μm; southeastern coastal plain
........................................................................................................ [Opegrapha longissima]
6.
Lirellae entirely black, sometimes pruinose, long or short; spores 20-40 μm long............ 7
7.
Spores 2.5-4.5 wide; lirellae long, very narrow (less than 0.25 mm wide), opening by a slit,
often branched; eastern …………………………….[Opegrapha vulgata (syn. O. cinerea]
7.
Spores 6-9 μm wide; lirellae fusiform, sometimes forked, opening broadly; very widely
distributed................................................................................................... Opegrapha varia
OPHIOPARMA
1.
On bark and wood, rarely rocks, in the Pacific Northwest south to California; spores straight
...........................................................................................................Ophioparma rubricosa
1.
On rocks in arctic-alpine regions........................................................................................ 2
2.
Spores curved, more than 30 μm long................................................. Ophioparma ventosa
2.
Spores straight, less than 30 μm long............................................ [Ophioparma lapponica]
ORPHNIOSPORA: See Key F.
PANNARIA (including Coccocarpia, Degelia, Fuscopannaria, Massalongia, Parmeliella, Psoroma, and
Vestergrenopsis)
1.
Soredia present along the lobe margins................................................... Pannaria conoplea
1.
Soredia absent .................................................................................................................... 2
2.
Thallus foliose, either minute (with lobes less than 1 mm wide), or relatively broad, generally
spreading to form rosettes, not shingled at the thallus margins……………………………3
2.
Thallus squamulose, with lobes about as long as they are broad, under 2 mm wide, commonly
overlapping at least in part1 ............................................................................................... 4
3.
Thallus olive to brown or yellow-brown; lower surface smooth........................................ 4
3.
Thallus gray to blue-gray or yellowish gray; lower surface blue-black, tomentose or with a
conspicuous prothallus ...................................................................................................... 7
4.
Lobes 0.1-1.5 mm wide, elongate or branching ................................................................. 5
4.
Lobes 2-4(-5) mm wide..................................................................... Nephroma helveticum
5.
Thallus chocoloate brown, with an upper cortex of pseudoparenchyma; lobes 0.5-1.5 mm
wide; on moss or mossy rock; apothecia frequent, biatorine, red-brown; photobiont Nostoc.
Humid western forests and northern boreal region……………………Massalongia carnosa
149
5.
Thallus olive to yellowish brown, lacking an upper cortex; lobes under 0.5 mm wide; on bare
rock; apothecia rare; photobiont Scytonema ...................................................................... 6
6.
Cylindrical isidia on thallus surface; on wet rocks in coastal mountains and foot of glaciers
........................................................................................................ Vestergrenopsis isidiata
6.
Cylindrical isidia rare, but margins divided into flattened lobules; on dry rocks mainly in
southwestern U.S., just reaching southern British Columbia; rare
........................................................................................................ [Koerberia sonomensis]
7.(3)
Isidia or lobules present ..................................................................................................... 8
7.
Isidia absent...................................................................................................................... 11
8.
Isidia on thallus surface; photobiont Scytonema; lower surface with a cortex, at least in part
.......................................................................................................... Coccocarpia palmicola
8.
Isidia or lobules marginal; photobiont Nostoc; lower surface entirely without a cortex.... 9
9.
Prothallus very thick, brown-black, projecting like a fringe all around the thallus; lobes 0.7-2
mm wide; coastal plain, Florida to North Carolina………………….[Parmeliella pannosa]
9.
Prothallus thin, conspicuous or not, or prothallus absent; mostly north of the southeastern
coastal plain...................................................................................................................... 10
10.
Lobes 0.3-1 mm wide, thin, flat, with a conspicuous blue-black prothallus; apothecia rare;
medulla PD–....................................................................................Parmeliella triptophylla
10.
Lobes 1-3(-4) mm wide, thick, slightly ascending, without a prothallus; apothecia abundant;
cortex and medulla PD+ orange (pannarin)……………………………….Pannaria tavaresii
11.(7)
Apothecia biatorine, convex with thin or disappearing margins ...................................... 12
11.
Apothecia lecanorine, more or less flat or concave, with persistent raised margins ........ 13
12.
Lower surface pale brown to black with a cortex; lobes smooth or with concentric ridges;
spores (6-) 7-14 (-16) x (2-)3-5 μm; mainly southern coastal plain; common
........................................................................................................ Coccocarpia erythroxyli
12.
Lower surface blue to black, lacking a cortex; lobes smooth with longitudinal radiate ridges;
spores 17-25 x 7-10 μm; northeastern maritime coast; rare……………[Degelia plumbea]
13.
Lobes mostly 0.7-2 mm wide, more or less smooth; apothecial margins smooth and even, or
toothed...................................................................................................Pannaria rubiginosa
13.
Lobes mostly 2-6 mm wide, with branching veins or ridges on the upper surface; apothecial
margins bumpy............................................................................................. Pannaria lurida
150
14.(2)
Thallus grass-green to brownish green when wet; photobiont green
............................................................................................................... Psoroma hypnorum
14.
Thallus dark gray to brownish gray when wet; photobiont blue-green ............................ 15
15.
Thallus with isidia or cylindrical, isidia-like outgrowths or lobules ................................ 16
15.
Thallus without isidia or isidia-like outgrowths............................................................... 17
16.
On wood or bark, rarely rock; black prothallus well-developed; edges of squamules not white
or felty; isidia slender, on the lobe margins and sometimes almost covering the entire thallus
.........................................................................................................Parmeliella triptophylla
16.
On soil or moss, rarely rocks; prothallus absent; edges of squamules gray to bluish white and
felty; thick, cylindrical to granular outgrowths on the lobe margins
............................................................................................... Fuscopannaria "praetermissa"
17.
Growing directly on rock, especially moist rock walls; edges of squamules not white
.................................................................................................... Fuscopannaria leucophaea
17.
Growing on bark, moss, or soil (sometimes on moss over rock) ..................................... 18
18.
Growing on moss, mossy rock or tree bases, or soil; spore walls conspicuously rough or
sculptured; prothallus absent; apothecia broad, lecanorine……………Pannaria pezizoides
18.
Growing directly on bark; spore walls smooth................................................................. 19
19.
Apothecia biatorine with smooth, pale brown margins; prothallus absent; squamules under 1
mm across, edges not or only slightly whitened…………………Fuscopannaria "saubinetii"
19.
Apothecia lecanorine with a bumpy, whitish margin; black prothallus usually conspicuous;
squamules 0.6-3 mm across, edges white and felty.......................................................... 20
20.
Spores tapering to a point at one or both ends; thallus reddish brown; without atranorin
..................................................................................................... Fuscopannaria leucosticta
20.
Spores ellipsoid, not pointed at the ends; thallus blue-gray to olive green; contains atranorin
(not detectable with a K test)………………………………….Fuscopannaria leucostictoides
PARMELIA (including Canoparmelia, Myelochroa, Paraparmelia, Parmelina, Parmelinopsis,
Pseudoparmelia, and leads to Bulbothrix, Canomaculina, Cetrelia, Hypotrachyna,
Parmotrema, Platismatia, and Rimelia)
1.
Marginal cilia or cilia-like rhizines present on lobe margins or in axils of lobes, often sparse
and visible only with a handlens ........................................................................................ 2
1.
Marginal cilia or cilia-like rhizines absent ....................................................................... 14
151
2.
Lobes broad, 4-20 mm wide; thallus usually loosely attached over entire surface, ascending
............................................................................................................................................ 3
2.
Lobes narrow to moderate, mostly under 5 mm wide; thallus closely appressed to substrate
............................................................................................................................................ 4
3.
Rhizines abundant to the edge of the lobes; white maculae usually conspicuous at least on lobe
tips in a reticulate pattern; medulla PD+ yellow, K+ red (salazinic acid)
............................................................................ Rimelia key (Rimelia and Canomaculina)
3.
Rhizines absent in a broad zone close to the lobe margin; maculae not common and usually
inconspicuous, never reticulate ......................................................................... Parmotrema
4.
Cilia bulbous at base, very short and stiff (noticeable only with a handlens)
............................................................................................................................. Bulbothrix
4.
Cilia not bulbous at base, short or long .............................................................................. 5
5.
Isidia present. ..................................................................................................................... 6
5.
Isidia absent........................................................................................................................ 9
6.
Rhizines regularly and abundantly forked......................................................Hypotrachyna
6.
Rhizines unbranched .......................................................................................................... 7
7.
Medulla C+ red or pink, KC+ red .....................................................................................7a
7.
Medulla C– or C+ pale pink, KC– or KC+ orange-yellow or pinkish violet. .................. 8
7a.
Isidia without black cilia growing from the tips; contains gyrophoric acid
..................................................................................................... Parmelinopsis minarum
7a.
Isidia with black cilia at the tips; contains 4,5 di-O-methylhiascic and 5-O-methylhiascic
acids)......................................................................................... Parmelinopsis horrescens
8.
Medulla white, PD–, K–, KC+ pinkish violet (4,5 di-O-methylhiascic acid and 5-Omethylhiascic acids); isidia becoming ciliate at the tips .............Parmelinopsis horrescens
8.
Medulla pale yellow, PD+ orange, K+ yellow to red, KC– or KC+ orange-yellow (secalonic
and galbinic acids); isidia without cilia at the tips………………………Myelochroa obsessa
9.(5)
Rhizines abundantly forked............................................................................Hypotrachyna
9.
Rhizines not branched ...................................................................................................... 10
10.
Soredia or schizidia present; medulla pale yellow, at least close to algal layer or below
pustules; apothecia absent or rare..................................................................................... 11
10.
Soredia or schizidia absent; medulla white or pale yellow; apothecia abundant…………12
152
11.
Medulla C+ pink, KC+ red, (gyrophoric acid); lobes square or truncated
........................................................................................................ Parmelinopsis spumosa
11.
Medulla C–, KC– or C+ yellow, KC+ yellow (secalonic acid derivatives); lobes rounded
........................................................................................................... Myelochroa aurulenta
12.
Rhizines absent from a broad or narrow zone close to the margin; medulla PD+ red
(protocetraric acid) .................................................................. Parmotrema michauxianum
12.
Rhizines more or less uniformly distributed; medulla PD– or PD+ orange………………13
13.
More or less restricted to California; medulla white, PD–, K–, C+ red (lecanoric acid)
................................................................................................................Parmelina quercina
13.
Eastern U.S. and adjacent Canada; medulla pale yellow, PD+ orange, K+ yellow or red, C+
yellow (galbinic acid)............................................................................Myelochroa galbina
14.(1)
Rhizines absent or sparse at least in a broad or narrow zone close to the lobe margin; thallus
usually loosely attached, and (or) with lobes 4-20 mm wide…………………………….15
14.
Rhizines abundant, uniformly distributed; thallus usually broadly attached, with lobes mostly
under 4 mm wide.............................................................................................................. 18
15.
Pseudocyphellae conspicuous on the lobe surface, appearing as white dots……………..16
15.
Pseudocyphellae absent or inconspicuous........................................................................ 17
16.
Thallus with marginal soredia; lobes with a smooth, even surface; medulla KC+ red and (or)
UV+ white................................................................................................................Cetrelia
16.
Thallus lacking soredia or isidia; lobes with a network of ridges and depressions; medulla
KC–, UV– ...........................................................................................................Platismatia
17.
Thallus surface usually very uneven, with ridges and depressions or otherwise wrinkled;
rhizines sparse throughout; pycnidia along the lobe margins…………………..Platismatia
17.
Thallus generally smooth and at most folded, without sharp ridges and depressions; rhizines
absent from a zone near the margin but usually abundant in the thallus center; pycnidia on the
lobe surface ....................................................................................................... Parmotrema
18.(14) Pseudocyphellae abundant and conspicuous, or sparse.................................................... 19
18.
Pseudocyphellae absent. .................................................................................................. 25
19.
Pseudocyphellae round or irregular, not forming a reticulate pattern of white markings;
medulla K–, often C+ red or pink .........................................................................Punctelia
19.
Pseudocyphellae irregular, forming a reticulate pattern of white markings; medulla
153
usually K+ red, never C+ pink or red ............................................................................. 20
20.
Isidia present. .................................................................................................................. 21
20.
Isidia absent. ................................................................................................................... 23
21.
Isidia globular, mostly without a cortex and dull, sometimes resembling soredia
.............................................................................................................. Parmelia hygrophila
21.
Isidia cylindrical or flattened, unbranched or branched, shiny, with a continuous cortex....
.......................................................................................................................................... 22
22.
Rhizines slender, squarrose except close to lobe margins; isidia especially abundant along
margins but also laminal ........................................................................ Parmelia squarrosa
22.
Rhizines thick, unbranched or rarely forked; isidia mostly laminal
.................................................................................................................. Parmelia saxatilis
23.(20) Soredia absent. On rock; montane, boreal and arctic. ....................... Parmelia omphalodes
23.
Soredia present ................................................................................................................. 24
24.
Soredia powdery, on lobe surface and margins; mature rhizines squarrose; on bark, wood,
rock, and sometimes soil; extremely common and widely distributed; lacking usnic acid
................................................................................................................... Parmelia sulcata
24.
Soredia coarsely granular, mostly limited to lobe margins; mature rhizines unbranched or
sometimes forked; on rock; uncommon, boreal to arctic; contains usnic acid in soralia
............................................................................................................... [Parmelia fraudens]
25(18). Lower surface white or pale to dark brown...................................................................... 26
25.
Lower surface black. ........................................................................................................ 29
26.
Medulla pale lemon yellow; thallus without soredia, isidia, or lobules; cortex and medulla
KC+ orange (secalonic acid). ..........................................................Pseudoparmelia uleana
26.
Medulla white, cortex KC– or difficult to interpret due to strong K+ yellow reaction .... 27
27.
Lower surface white or pale brown; cortex and medulla PD+ orange, K+ deep yellow
(thamnolic acid) ...................................................................................................Imshaugia
27.
Lower surface medium to dark brown; cortex PD–, K+ pale yellow (atranorin); medulla PD–,
K–..................................................................................................................................... 28
28.
Thallus surface with isidia; lobe tips with a network of white maculae
..................................................................................................... Canoparmelia caroliniana
154
28.
Thallus surface without isidia, but with granular soredia; maculae sparse and inconspicuous
.............................................................................................................Canoparmelia texana
29.(25) Rhizines dichotomously branched; medulla white,....................................... Hypotrachyna
29.
Rhizines unbranched or sometimes squarrose; medulla white or pale yellow ………….30
30.
Soredia or fragments originating from the breakdown of pustules or hollow warts present 31
30.
Soredia and soredia-like fragments absent. ...................................................................... 34
31.
Medulla pale yellow, at least close to algal layer, K–, or K+ yellow, PD–, UV–; common, East
Temperate.......................................................................................... Myelochroa aurulenta
31.
Medulla entirely white; southeastern U.S. ....................................................................... 32
32.
On rocks; lobes 1-2 mm wide, smooth; soralia discrete, in hemispherical mounds, containing
dark gray granular soredia; medulla PD+ red-orange, KC+ pink (protocetraric acid)
.................................................................................................. [Paraparmelia alabamensis]
32.
On bark of hardwoods; lobes 2-5 mm broad, wrinkled or ridged; soralia discrete or running
together, especially along ridges, remaining pale; medulla PD+ orange or PD–, KC– or KC+
faint purple ....................................................................................................................... 33
33.
Medulla K+ yellow, PD+ orange, UV– (stictic acid)…………….[Canoparmelia crozalsiana]
33..
Medulla K–, PD–, UV+ blue-white (divaricatic acid)………………….Canoparmelia texana
34.(30) Isidia absent; medulla pale yellow, at least near algal layer……………..Myelochroa galbina
34.
Isidia present; medulla white or pale yellow.................................................................... 35
35.
Lobes with a network of whitish maculae, wrinkled and rough; medulla white, PD–, K–, UV+
white (perlatolic acid) ................................................................. Canoparmelia caroliniana
35.
Lobes without maculae, smooth and uniform (where there are no isidia); medulla pale yellow,
at least close to algal layer, PD+ orange, K+ orange to red (galbinic acid)
.............................................................................................................. Myelochroa obsessa
PARMELIELLA: See Pannaria.
PARMELINA: See Parmelia
PARMELINOPSIS: See Parmelia.
PARMELIOPSIS: See Keys J and K.
155
PARMOTREMA
1.
Marginal cilia or cilia-like rhizines present (although sometimes sparse), on lobe margins or in
axils of lobes ...................................................................................................................... 2
1.
Marginal cilia or cilia-like rhizines absent ....................................................................... 17
2.
Soredia absent. ................................................................................................................... 3
2.
Soredia present. ................................................................................................................ 10
3.
Isidia present. ..................................................................................................................... 4
3.
Isidia absent........................................................................................................................ 8
4.
Medulla bright yellow .................................................................. Parmotrema sulphuratum
4.
Medulla white .................................................................................................................... 5
5.
Thallus yellowish green; cortex K–, KC+ yellow (usnic acid); medulla K–, PD–
......................................................................................................... Parmotrema xanthinum
5.
Thallus gray to yellowish gray; cortex K+ yellow, KC– (atranorin); medulla K+ yellow or red,
PD+ yellow or orange, or K–, PD– .................................................................................... 6
6.
Isidia without cilia; maculae present; medulla K+ red, UV+ yellow-orange (mostly in lower
half of medulla close to lower cortex) (salazinic acid and lichexanthone).
.........................................................................................................Parmotrema ultralucens
6.
Isidia often with short black cilia growing out of the tips; maculae absent; medulla K– or K+
yellow, UV+ white or UV– (lichexanthone absent) ........................................................... 7
7.
Isidia not breaking down into soredia; medulla K+ yellow, KC–, UV– (stictic acid)
.............................................................................................................Parmotrema crinitum
7.
Isidia partly breaking down into granular soredia; medulla K–, KC+ red, UV+ white
(alectoronic acid).............................................................................. [Parmotrema mellissii]
8.(3)
Apothecial disks not perforated; maculae absent; lower surface brown, never white at edge;
medulla PD+ red, K–, KC+ red (protocetraric acid)…………..Parmotrema michauxianum
8.
Apothecial disks perforated with an irregular hole through the center; maculae present on
upper surface; lower surface uniform or splotched with white; medulla PD+ orange, K+ red,
KC– ................................................................................................................................... 9
9.
Lower surface usually splotched with white near the margins; contains norstictic acid; mainly
southeastern coastal plain and Appalachians……………………….Parmotrema perforatum
9.
Lower surface pale to dark brown, not splotched with white; contains salazinic acid; arid
central states, Texas to North Dakota........................................... [Parmotrema eurysacum]
156
10.(2)
Medulla K–, PD–, KC+ red (alectoronic acid)................................................................. 11
10.
Medulla K+ yellow, orange, or red, PD+ yellow, orange, or red, KC– or KC+ pink
(alectoronic acid absent)................................................................................................... 13
11.
Soredia developing from short isidia, many of which are ciliate (with short black hairs)
.......................................................................................................... [Parmotrema mellissii]
11.
Soredia developing from breakdown of thallus cortex, not isidia .................................... 12
12.
Soredia along the lobe margins; southeastern coastal plain……..Parmotrema rampoddense
12.
Soredia on the upper surface of lobe tips; west coast, Appalachian and Great Lakes regions
............................................................................................................. Parmotrema arnoldii
13.(10) Cilia short, sparse, and usually confined to lobe axils; medulla PD+ red-orange (protocetraric
acid)................................................................................................... Parmotrema dilatatum
13.
Cilia long or short, often abundant, scattered along lobe margins; medulla PD+ yellow or
orange............................................................................................................................... 14
14.
Medulla K+ yellow or orange (stictic acid)........................................ Parmotrema chinense
14.
Medulla K+ red (salazinic or norstictic acid) .................................................................. 15
15.
Lower surface with blotches of ivory white close to the margins; medulla PD+ yellow
(norstictic acid)............................................................................. Parmotrema hypotropum
15.
Lower surface uniformly black to dark brown, rarely with white areas; medulla PD+ orange
(salazinic acid) ................................................................................................................. 16
16.
Soredia on or close to the margins; a network of white maculae and often white cracks usually
present at least on the lobe tips (rarely absent); rhizines abundant .......... Rimelia reticulata
16.
Soredia narrowly restricted to the lobe margins; maculae absent; rhizines sparse
............................................................................................Parmotrema stuppeum
17.(1)
Soredia absent, but sometimes isidiate........................................................................... 18
17.
Soredia present ................................................................................................................. 20
18.
Medulla pale orange-yellow, K+ yellow, C+ pink (gyrophoric acid); thallus isidiate
................................................................................................Parmotrema endosulphureum
18.
Medulla white, K–, C+ red or C– (gyrophoric acid absent). ........................................... 19
19.
Upper surface dull; medulla C+ red (lecanoric acid); thallus isidiate
..........................................................................................................Parmotrema tinctorum
157
19.
Upper surface rather shiny; medulla C–; thallus with or without isidia. ......... Platismatia
20.(17) Medulla K+ yellow or red................................................................................................ 21
20.
Medulla K–. .................................................................................................................... 22
21.
Soredia granular; medulla PD+ red, K+ yellow (protocetraric acid and echinocarpic acids)
........................................................................................................... Parmotrema dilatatum
21.
Soredia farinose; medulla PD+ orange, K+ red (salazinic acid)
........................................................................................................ Parmotrema cristiferum
22.
Soralia on the thallus surface; thallus closely appressed to substrate, lobes flat; lower surface
more or less uniform in color; medulla UV+ blue-white (divaricatic acid)
.............................................................................................................Canoparmelia texana
22.
Soralia marginal; thallus loosely attached over entire surface, ascending; lobes undulating or
crisped at the margins; lower surface pale to dark brown or blotched with white close to the
margin; medulla UV–....................................................................................................... 23
23.
Medulla C+ red or pink (lecanoric acid) .....................................Parmotrema austrosinense
23.
Medulla C–....................................................................................................................... 24
24.
Soredia in crescent-shaped soralia on older lobes; on Gulf coastal plain
.................................................................................................Parmotrema praesorediosum
24.
Soredia scattered over lobe margins, not in crescent-shaped soralia; not on Gulf coastal plain
................................................................................................................. Platismatia glauca
PELTIGERA (including Erioderma and Leioderma)
1.
Photobiont green; cephalodia present................................................................................. 2
1.
Photobiont blue-green; cephalodia absent.......................................................................... 5
2.
Thallus attached by a single point at one edge, fanning out; rhizines absent; cephalodia
forming tiny nodular lobules on the lower surface mostly on the veins; disks more or less flat
.................................................................................................................... Peltigera venosa
2.
Thallus attached by numerous rhizines except at the lobe margin; cephalodia forming scales
on the upper surface of the lobes; disks saddle-shaped ...................................................... 3
3.
Cephalodia in the form of lobed scales loosely attached to the upper thallus surface, easily
detached ................................................................................................ Peltigera britannica
158
3.
Cephalodia in the form of round to slightly scalloped scales closely appressed, firmly fixed to
the thallus surface............................................................................................................... 4
4.
Lower surface with conspicuous veins; lobes crisped and undulating at the margins; rhizines
separate and distinct; lower surface of apothecia with green, scale-like, discontinuous patches
........................................................................................................... Peltigera leucophlebia
4.
Lower surface black in the center changing abruptly to white at the margins, or with a regular
pattern of white and dark areas near the margins; lobes more or less flat, not crisped or
undulating at the margins; rhizines forming an intricately branched and anastomosing mat;
lower surface of apothecia with a continuous, bumpy cortex ................. Peltigera aphthosa
5.(1)
Soredia present ................................................................................................................... 6
5.
Soredia absent .................................................................................................................... 9
6.
Soredia in irregular gray patches on thallus surface................................Peltigera didactyla
6.
Soredia marginal or on the lower surface of the lobe tips .................................................. 7
7.
Upper surface smooth to scabrous, not tomentous; lower surface with conspicuous veins;
photobiont Nostoc ...................................................................................... Peltigera collina
7.
Upper surface tomentose at least near lobe margins; lower surface more or less uniform;
photobiont Scytonema........................................................................................................ 8
8.
Tomentum on upper surface made up of erect hairs; medulla PD+ orange (eriodermin)
........................................................................................................... Erioderma sorediatum
8.
Tomentum on upper surface webby; medulla PD–
...........................................................................................................Leioderma sorediatum
9.(5)
Isidia present on the thallus surface ................................................................................. 10
9.
Isidia absent; lobules present or absent ............................................................................ 11
10.
Lobes mostly 5-10 mm broad, concave; isidia flat and scale like, peltate
............................................................................................................ Peltigera lepidophora
10.
Lobes 7-25 mm broad, turned down at the margins; isidia granular to erect and flattened
................................................................................................................Peltigera evansiana
11.
Lobules present on margins and along cracks.................................................................. 12
11.
Lobules absent.................................................................................................................. 14
12.
Upper surface dull or tomentose, at least at lobe tips...........................Peltigera praetextata
12.
Upper surface rather shiny ............................................................................................... 13
159
13.
Lower surface with conspicuous veins; rhizines distributed along the veins; disks
saddle-shaped ............................................................................................Peltigera pacifica
13.
Lower surface with a regular pattern of white and dark areas; rhizines arising in more or less
concentric bands; disks rather flat ........................................................Peltigera elisabethae
14.
Lower surface without distinct veins or patterning .......................................................... 15
14.
Lower surface with distinct veins, or with a regular pattern of white and dark areas; rhizines
abundant ........................................................................................................................... 16
15.
Lower surface black in the center grading to white at the margins; rhizines sparse; thallus
thick, dark green when wet ...................................................................... Peltigera malacea
15.
Lower surface pale; thallus thin, not turning dark green when wet; east coast
............................................................................................................. [Peltigera hymenina]
16.
Upper surface rather shiny ............................................................................................... 17
16.
Upper surface dull, scabrose, or tomentose...................................................................... 21
17.
Apothecial disks black; maculae present on thallus surface.......................Peltigera neckeri
17.
Apothecial disks brown; maculae absent ......................................................................... 18
18.
Veins pale, narrow and conspicuously raised, distinctly fuzzy with an erect tomentum; thallus
thin and membranous, often with a blistered appearance……………. Peltigera membranacea
18.
Veins dark (at least in older parts of thallus), broad or narrow, relatively flat, fuzzy or smooth;
thallus not membranous and relatively smooth ................................................................ 19
19.
Rhizines arising in more-or-less concentric bands; apothecia flat; spores mostly under 45 μm
long .....................................................................................................Peltigera horizontalis
19.
Rhizines distributed along the veins; apothecia saddle-shaped; spores mostly over 45 μm long
.......................................................................................................................................... 20
20.
Lobes 7-15(-20) mm across, with crisped margins; upper surface shiny throughout...........
...........................................................................................................Peltigera polydactylon
20.
Lobes 20-40 mm across, round and not crisped; surface rather dull and often pruinose......
....................................................................................................... Peltigera neopolydactyla
21.(16) Lobes concave, or undulating at the margins, upper surface slightly to heavily tomentose
.......................................................................................................................................... 22
21.
Lobes mostly flat, not undulating at the margins; upper surface tomentose, smooth, or
scabrose............................................................................................................................ 24
160
22.
Lobe margins entire; lobes strongly erect ...............................................Peltigera didactyla
22.
Lobe margins scalloped; lobes mostly adnate .................................................................. 23
23.
Rhizines tufted, running together; lobe surface heavily tomentose
................................................................................................................ Peltigera rufescens
23.
Rhizines unbranched or brushlike, separate and distinct; lobe surface heavily tomentose or
dull but almost without tomentum........................................................ Peltigera ponojensis
24.(21) Rhizines rope-like, with a fuzzy surface, unbranched (see couplet 18); tomentum on upper
surface often confined to extreme lobe tips with the remainder of the surface rather shiny
......................................................................................................... Peltigera membranacea
24.
Rhizines fibrous and smooth, tufted, or brushlike; upper surface tomentose, dull, or scabrose
.......................................................................................................................................... 25
25.
Surface more or less smooth, not scabrose or pruinose, always tomentose at least near the lobe
tips; lower surface with raised, pale veins, sometimes dark in the center of the thallus ......
..................................................................................................................... Peltigera canina
25.
Surface scabrose or pruinose in part, with or without tomentum; lower surface with dark
veins, distinct or indistinct ............................................................................................... 26
26.
Upper surface dull, sometimes pruinose, but not scabrose; lobes very broad (20-40 mm)
....................................................................................................... Peltigera neopolydactyla
26.
Upper surface scabrose .................................................................................................... 27
27.
Lobes 20-40 mm across; veins distinct, more or less smooth; cortex usually C+ pink
(gyrophoric acid and tenuiorin)................................................................Peltigera scabrosa
27.
Lobes 7-15 mm across; veins very dark, contrasting with white spaces between them,
distinctly fuzzy with an erect tomentum; cortex always C– (gyrophoric acid and tenuiorin
absent) ................................................................................................Peltigera kristinssonii
PELTULA (including Heppia)
1.
Spores ellipsoid, 8 per ascus ...................................................................Heppia conchiloba
1.
Spores globose or broadly ellipsoid, up to 100 per ascus................................................... 2
2.
Spores broadly ellipsoid; growing directly on rock or soil ................................................ 3
2.
Spores globose; growing on soil. Squamules round; epihymenium turning red to violet or
purple with K ..................................................................................................................... 4
161
3.
Squamules scalloped (with rounded lobes); epihymenium K–
.................................................................................................Peltula obscurans var. hassei
3.
Squamules unlobed; epihymenium K+ red-purple……..[Peltula obscurans var. obscurans]
4.
Squamules with edges turned down, upper surface rough (rugose), lower surface blackish;
apothecial margin usually absent .............................................................. Peltula richardsii
4.
Squamules with edges turned up, upper surface smooth, lower surface pale brown; apothecial
margin prominent and usually persistent…………………………………. Peltula patellata
PERTUSARIA
1.
Apothecia buried in verrucae with straight or sloping sides, or that are slightly constricted at
the base, having small ostiole-like openings ...................................................................... 2
1.
Apothecia lecanorine, with narrow or broad disks, or becoming partly or entirely sorediate or
pruinose at the summit ..................................................................................................... 16
2.
Spores (4-)8 per ascus ........................................................................................................ 3
2.
Spores 2-4 per ascus........................................................................................................... 6
3.
Thallus yellowish green or sometimes yellow-gray; cortex C+ yellow-orange, KC+ orange,
UV+ dark orange (thiophanic acid).................................................................................... 4
3.
Thallus greenish gray, not yellowish; cortex C–, KC–, UV– or UV+ yellow.................... 5
4.
Ostiole area depressed, dark brown to black, often pruinose, C–; epihymenium K+ violet;
medulla K+ red (norstictic acid); Appalachian-Great Lakes distribution.............................
............................................................................................................ [Pertusaria rubefacta]
4.
Ostiole area raised and yellow, C+ orange; epihymenium K–; medulla K+ yellow (stictic
acid); southeastern..................................................................................... Pertusaria texana
5.
Spores in two irregular rows within the ascus. Thallus and verrucae UV– or orange-pink,
rather smooth; ostioles dot-like; Appalachian-Ozark distribution………..[Pertusaria ostiolata]
5.
Spores lined up in a single row within the ascus...............................................................5a
5a
Spores 50-130 x 25-45 μm; thallus and verrucae UV+ bright yellow (lichexanthone), smooth
or rough and warty; ostioles often with a whitish border; throughout the southeast
.................................................................................................. Pertusaria paratuberculifera
5a.
Spores 36-60 x 18-32 μm; Thallus and verrucae UV- or dull pinkish orange, smooth to
verruculose; ostioles without a white border; Great Lakes region east to Maritimes and
Massachusetts, also Alaska……………………………… [Pertusaria alpina Hepp ex Ahles]
6.(2)
Growing on bark, peat, or soil............................................................................................ 7
162
6.
Growing directly on rock ................................................................................................. 15
7.
On soil or peat, mostly arctic-alpine (rarely on hardwood bark on east coast). Spores 2 per
ascus; abundant black ostioles in separate or frequently confluent verrucae; epihymenium K+
violet; medulla K+ yellow (stictic acid)………………………….[Pertusaria subobducens]
7.
On bark............................................................................................................................... 8
8.
Spores mostly 4 per ascus .................................................................................................. 9
8.
Spores 2(-3) per ascus ...................................................................................................... 11
9.
Epihymenium K+ violet, fertile verrucae mostly with steep-sides. Southeastern coastal plain
.................................................................................................... [Pertusaria sinusmexicani]
9.
Epihymenium K–; fertile verrucae mostly with sloping sides.......................................... 10
10.
Spores 75-130 μm long, with rough inner walls; East Temperate
...................................................................................................... [Pertusaria tetrathalamia]
10.
Spores 50-100 μm long, with smooth inner walls; East Temperate and west coast
........................................................................... [Pertusaria leioplaca (syn. P. leucostoma)]
11(8).
Thallus pale greenish yellow to yellow-gray or greenish gray; cortex C+ yellow-orange, KC+
orange, UV+ orange-red; thallus medulla PD+ orange, K+ persistently yellow (stictic acid)
.......................................................................................................................................... 12
11.
Thallus gray; cortex C–, KC–, UV– or UV+ yellow or pink…………………………….13
12.
Inner spore walls rough and grooved when mature; ostioles pale brown, typically separate;
mainly on coastal plain........................................................................ Pertusaria xanthodes
12.
Inner spore walls smooth; ostioles black, sometimes clustered or fused; East Temperate
.............................................................................................................[Pertusaria pustulata]
13.
Ostioles black; epihymenium K+ violet. Medulla K+ yellow, PD+ orange (stictic acid); Great
Lakes to Maritime Provinces........................................................... [Pertusaria consocians]
13.
Ostioles pale to dark, sometimes black; epihymenium K– .............................................. 14
14.
Inner wall of spores with radiating channels; spores sometimes brownish and K+ dull violet;
medulla K+ yellow, PD+ orange (stictic acid); Appalachian-Great Lakes distribution
...............................................................................................................Pertusaria macounii
14.
Inner wall of spores smooth, not channelled, always colorless and K–; medulla PD+ red, K–
or brownish (fumarprotocetraric acid); southeastern …………………Pertusaria subpertusa
163
15.(6)
Thallus gray, thin; spores narrowly ellipsoid (75-)85-150(-245) x 25-70 μm, with rough inner
walls; thallus cortex C–, KC–; medulla K+ red (norstictic acid); eastern North America
.................................................................................................................Pertusaria plittiana
15.
Thallus sulphur yellow, thick; spores 60-90 x 30-50 μm, with smooth walls; thallus cortex C+
orange, KC+ orange (thiophanic acid); medulla K–; southern California
.............................................................................................................Pertusaria flavicunda
16.(1)
Growing on mosses or dead vegetation............................................................................ 17
16.
Growing on bark or rock .................................................................................................. 18
17.
Thallus white to bluish gray, verrucose, but verrucae not resembling isidia; black, pruinose
apothecia immersed in the verrucae; all spot tests negative………………..Pertusaria panyrga
17.
Thallus composed of white, cylindrical isidia with black, sometimes lightly pruinose apothecia
at the summits; thallus cortex and medulla PD+ red, K+ brownish (fumarprotocetraric acid)
............................................................................................................... Pertusaria dactylina
18.
On rock; thallus sulphur yellow, cortex C+ yellow-orange; spores 2(-4) per ascus (see couplet
15) .......................................................................................................Pertusaria flavicunda
18.
Usually on bark; thallus gray to yellowish green, not sulphur yellow, cortex C–, C+ yelloworange, or C+ red; spores 1-8 per ascus ........................................................................... 19
19.
Thallus UV+ yellow (lichexanthone); spores 1(-2) per ascus, but often absent; eastern.. 20
19.
Thallus UV–; spores 1, 2, or 8 per ascus.......................................................................... 22
20.
Verrucae becoming sorediate at the summit. Medulla KC+ pink becoming violet, PD–
(hypothamnolic acid); throughout southeast……………………..[Pertusaria hypothamnolica]
20.
Verrcucae not becoming sorediate, summit small or broad and lecanorine, white pruinose
.......................................................................................................................................... 21
21.
Medulla C–, KC–, PD+ yellow to orange (haemathamnolic acid); verrucae somewhat raised,
not expanded; southern coastal plain…………………………………….[Pertusaria copiosa]
21.
Medulla C+ red, KC+ red, PD– (lecanoric acid); verrucae usually expanded into lecanorine
apothecia; East Temperate ......................................................................... Pertusaria velata
22.
Apothecial margins smooth; disk pinkish orange, heavily pruinose but not at all sorediate;
thallus cortex and medulla C+ red, KC+ red (lecanoric acid)………………Pertusaria velata
22.
Apothecial margins or the entire verrucae rough or sorediate; thallus cortex C–, medulla KC–
or KC+ purple (lecanoric acid absent) ............................................................................ 23
164
23.
Soralia or sorediate verrucae KC+ purple (picrolichenic acid); spores 1 per ascus but rarely
found ..........................................................................................................Pertusaria amara
23.
Soralia (if present), verrucae, and thallus medulla KC–; spores 1-8 per ascus ................ 24
24.
Thallus medulla K+ bright yellow, PD+ orange (thamnolic acid); spores 2 per ascus but
sometimes hard to find ...................................................................Pertusaria trachythallina
24.
Thallus medulla K–, PD– or PD+ red; spores not 2 per ascus ......................................... 25
25.
Thallus medulla PD–, K–. Spores 1 per ascus; widely distributed
.......................................................................................................... Pertusaria ophthalmiza
25.
Thallus medulla PD+ red, K+ brown (fumarprotocetraric acid) ...................................... 26
26.
Apothecial disks usually distinct, various colors from yellow to green or black, usually
pruinose; apothecial margins ragged, often in concentric layers; spores 8 per ascus; Pacific
Northwest ........................................................................................Pertusaria subambigens
26.
Apothecial disks usually obscured by soredia and aborted, rarely black and pruinose with
sorediate margins; spores 1 per ascus; Appalachian-Great Lakes region
................................................................................................. [Pertusaria multipunctoides]
PHAEOGRAPHINA
1.
Lirellae branched; walls of lirellae entirely carbonized………Phaeographina quassiaecola
1.
Lirellae unbranched; walls of lirellae partially carbonized.
.............................................................................................. Phaeographina caesiopruinosa
PHAEOGRAPHIS: See Key E.
PHAEOPHYSCIA (including Hyperphyscia and Physciella)
1.
Soredia absent .................................................................................................................... 2
1.
Soredia present ................................................................................................................... 7
2.
Rhizines absent; thallus extremely closely appressed, almost crustose (but with a lower cortex)
.......................................................................................................... Hyperphyscia syncolla
2.
Rhizines present, sparse or abundant; thallus easily detached from substrate, clearly foliose
............................................................................................................................................ 3
3.
Thallus very loosely attached, growing over soil and moss; lower surface white to pale brown;
rhizines sparse, white to very pale brown…………………………..Phaeophyscia constipata
3.
Thallus closely to somewhat loosely attached, growing on bark or rock; lower surface black;
rhizines abundant, black..................................................................................................... 4
165
4.
Lobes 1-3(-4) mm across; cilia-like rhizines long........................... Phaeophyscia hispidula
4.
Lobes 0.2-1 mm across; cilia-like rhizines very short or absent ........................................ 5
5.
Growing on bark, moss, or wood ....................................................................................... 6
5.
Growing on rock ................................................................................Phaeophyscia decolor
6.
Erect lobules abundant on lobe margins.......................................[Phaeophyscia squarrosa]
6.
Lobules absent......................................................................................Phaeophyscia ciliata
7.(1)
Rhizines absent; thallus attached directly to substrate, almost crustose
......................................................................................................Hyperphyscia adglutinata
7.
Rhizines sparse or abundant; thallus attached by rhizines, clearly foliose......................... 8
8.
Lower surface and rhizines white to pale brown................................................................ 9
8.
Lower surface and rhizines black..................................................................................... 10
9.
Cortex K–, atranorin absent; rhizines abundant; conidia ellipsoid ....................................9a
9.
Cortex K+ yellow, atranorin present; rhizines sparse; conidia rod-shaped
........................................................................................................................ Physcia dubia
9a.
Soredia entirely marginal or on lobe tips ......................................Physciella chloantha
9a.
Soredia on lobe surface or lobe margins; northeastern to Montana and Colorado
........................................................................................................[Physciella melanchra]
10.
Medulla red-orange, sometimes only in spots........................... Phaeophyscia rubropulchra
10.
Medulla white .................................................................................................................. 11
11.
Stiff, tiny, colorless hairs on the upper surface of the lobes
...................................................................................................Phaeophyscia cernohorskyi
11.
Stiff, tiny, colorless hairs absent ...................................................................................... 12
12.
Soredia black, very coarse (almost isidia), along the lobe margins; on rock
............................................................................................................Phaeophyscia sciastra
12.
Soredia greenish, granular to farinose, on the margins and tips of the lobes; on bark or rock
.......................................................................................................................................... 13
13.
Lobes 1-3(-4) mm across; cilia-like rhizines long........................... Phaeophyscia hispidula
13.
Lobes 0.5-1.5(-2) mm across; cilia-like rhizines very short............................................. 14
166
14.
Soredia on the upper surface of the lobe tips, or on expanded, turned back lobe tips
(lip-shaped soralia).......................................................................................................... 15
14.
Soredia on the lobe margins, close to the margins, or on thallus surface......................... 16
15.
Soredia farinose, forming small hemispherical greenish mounds on the upper surface of the
lobe tips; lobes mostly less than 0.7 mm wide; on bark, or rarely on calcareous rock (e.g.,
limestone)..................................................................................... Phaeophyscia pusilloides
15.
Soredia granular, mostly in lip-shaped soralia at the lobe tips; lobes 0.5-1(-2) mm wide; on
siliceous rock (e.g., granite), or occasionally on bark……………..Phaeophyscia adiastola
16.
Soredia farinose, mainly on lobe surface in depressed patches, but some marginal
.......................................................................................................Phaeophyscia orbicularis
16.
Soredia granular, mainly on lobe tips, occasionally on surface in mounds
......................................................................................................... Phaeophyscia adiastola
PHAEORRHIZA: See Key G and Rinodina.
PHLYCTIS: Keys C and F.
PHYLLOPSORA: See Keys F and G.
PHYSCIA (including Dirinaria, Heterodermia, Imshaugia, and Pyxine)
1.
Soredia absent .................................................................................................................... 2
1.
Soredia present ................................................................................................................. 23
2.
Isidia present ...................................................................................................................... 3
2.
Isidia absent........................................................................................................................ 5
3.
Rhizines more or less uniformly distributed, short and mostly unbranched; cortex and medulla
K+ deep yellow, PD+ orange (thamnolic acid); on conifers and birch in northern and montane
regions................................................................................................... Imshaugia aleurites
3.
Rhizines mostly on or close to the margins; cortex K+ yellow; medulla PD– or PD+ yellow
(lacking thamnolic acid); on deciduous trees in southeastern U.S. .................................... 4
4.
Lower surface yellow or orange, webby; medulla K+ yellow (or deep purple where
pigmented) .......................................................................................... Heterodermia crocea
4.
Lower surface white to tan, smooth; medulla K+ red (salazinic acid)
....................................................................................................[Heterodermia granulifera]
5.(2)
Thallus attached by only a few points, almost fruticose, with long narrow lobes…………...6
167
5.
Thallus closely or loosely attached; lobes rarely long and narrow..................................... 7
6.
Apothecial margins smooth and even; southwestern U.S……………Heterodermia erinacea
6.
Apothecial margins toothed or lobulate; south central or southeastern U.S.
.......................................................................................................... Heterodermia echinata
7.
Medulla yellow; apothecia lecideine, black; cortex UV+ yellow-orange (lichexanthone)
...................................................................................................................Pyxine berteriana
7.
Medulla white; apothecia lecanorine, brown to black; cortex UV- (lichexanthone absent).
............................................................................................................................................ 8
8.
Cortex K– (atranorin absent)........................................................... Phaeophyscia hispidula
8.
Cortex K+ yellow (atranorin, except for Imshaugia) ........................................................ 9
9.
Medulla with patches of orange-yellow pigment (anthraquinones), K+ yellow (atranorin), or
K+ purple where pigmented; thallus heavily white pruinose. Uncommon, in extreme
southwest....................................................................................... [Heterodermia rugulosa]
9.
Medulla entirely white, K+ yellow (atranorin) or K-; thallus usually without pruina .... 10
10.
Lobules present at least on the inrolled apothecial margins and usually the lobe margins,
sometimes also on the thallus surface (occasionally absent in H. hypoleuca, which has brown
squarrose rhizines) ........................................................................................................... 11
10.
Lobules absent; rhizines never squarrose ......................................................................... 13
11.
Lobules finely divided, on margins and upper surface of lobes; lower surface becoming
purplish black in the center .........................................................Heterodermia squamulosa
11.
Lobules rounded or strap-shaped, on the margins of the lobes and apothecia; lower surface
white to gray throughout .................................................................................................. 12
12.
Lower surface cottony, entirely without a cortex; rhizines squarrose, well developed and often
forming a mat; thallus not pruinose; mainly eastern, although also in southwestern U.S.
....................................................................................................... Heterodermia hypoleuca
12.
Lower surface smooth, with a cortex, at least in part; rhizines unbranched or forked,
occasionally squarrose, not forming a mat; thallus somewhat pruinose on lobe tips;
southwestern U.S........................................................................... Heterodermia diademata
13.(10) On rock............................................................................................................................. 14
13.
On bark or wood............................................................................................................... 16
14.
Thallus heavily pruinose ..............................................................................Physcia biziana
168
14.
Thallus entirely without pruina ........................................................................................ 15
15.
Lobes 0.7-1.5(-2.5) mm wide; maculae present on upper surface; zeorin present
........................................................................................................................Physcia phaea
15.
Lobes 0.3-0.5 mm wide; maculae absent; zeorine absent ............................... Physcia halei
16.(13) Lower surface black; very tightly attached to substrate (almost crustose in appearance) 17
16.
Lower surface white to pale brown; thallus closely or loosely attached to substrate ....... 18
17.
Lobes 1-2(-4) mm wide; apothecial disks black, without pruina………….Dirinaria confusa
17.
Lobes 0.2-0.7 mm wide; apothecial disks reddish purple caused by a purple pruina
.......................................................................................................... Dirinaria purpurascens
18.
Spores colorless, single-celled; cortex and medulla PD+ orange (thamnolic acid; atranorin
absent) ................................................................................................ Imshaugia placorodia
18.
Spores brown, 2-celled; cortex and medulla PD– (atranorin present, thamnolic acid absent)
.......................................................................................................................................... 19
19.
Medulla K+ yellow (atranorin) ........................................................................................ 20
19.
Medulla K– ...................................................................................................................... 22
20.
Spores with rounded cells (Pachysporaria-type); apothecial margins sometimes with a fringe
of rhizines on the lower side; zeorin absent. .............................................. Physcia neogaea
20.
Spores with angular cells (Physcia-type); apothecial margins without rhizines on lower side;
zeorin present ................................................................................................................... 21
21.
Lobes convex, 0.3-0.5 mm wide ............................................................... Physcia pumilior
21.
Lobes flat or somewhat concave at the tips, 1-2(-3) mm wide
...................................................................................................................... Physcia aipolia
22.(19) Thallus entirely without pruina ...................................................................Physcia stellaris
22.
Thallus pruinose ..............................................................................................................22a
22a.
Lobes less than 1 mm across, convex; thallus under 3 mm in diameter; southern
Manitoba ............................................................................... [Physcia sp. (cfr. stellaris)]
22a.
Lobes more than 1 mm across, flat; thallus often more than 3 cm in diameter; western
.................................................................................................................... Physcia biziana
23.(1)
Lower surface black or purplish, at least in center of thallus .......................................... 24
23.
Lower surface white to dark brown, gray, yellow, or orange.......................................... 34
169
24.
Rhizines absent, thallus attached directly to substrate; medulla UV+ blue-white (divaricatic
acid).................................................................................................................................. 25
24.
Rhizines sparse or abundant; medulla UV- or UV+ yellow-orange or red (divaricatic acid
absent) .............................................................................................................................. 27
25.
Soredia granular, originating from the breakdown of pustules or hollow warts, covering the
thallus surface ..........................................................................................Dirinaria aegialita
25.
Soredia farinose, in hemispherical mounds...................................................................... 26
26.
On bark, southeastern coastal plain............................................................... Dirinaria picta
26.
On rock, East Temperate distribution........................................................ [Dirinaria frostii]
27.(24) Medulla yellow to pale orange ......................................................................................... 28
27.
Medulla white .................................................................................................................. 30
28.
Soredia entirely marginal, granular or coarse ……………………………………………28a
28.
Soredia mostly laminal (but see P. eschweileri below), farinose, greenish white or green;
thallus closely appressed; southeastern ............................................................................ 29
28a.
Thallus with ascending lobe tips that are mostly 1-2.5 mm wide; widely distributed, East
Temperate; soredia granular, bluish gray; thallus cortex K+ yellow, UV– (atranorin);
medulla mustard yellow, K –................................................................. Pyxine sorediata
28a.
Thallus appressed throughout; lobes under 1 mm wide; restricted to southeastern
coastal plain; soredia developing from fragments of pustules (schizidia), not bluish gray;
thallus cortex K–, UV+ yellow (lichexanthone) ; medulla salmon orange, K+ purple
(anthraquinone)........................................................[Pyxine caesiopruinosa]
29.
Thallus UV–, K+ yellow (atranorin); medulla PD+ orange; some forms with marginal isidialike granules (schizidia) from the breakdown of pustules
................................................................................................................ Pyxine eschweileri
29.
Thallus UV+ yellow, K– (lichexanthone); medulla PD–, never with marginal schizidia
.........................................................................................................................................29a
29a.
Medulla salmon orange, K+ purple (anthraquinone); restricted to southeastern coastal
plain ……………………………………………………….…….[Pyxine albovirens]
29a.
Medulla pale yellow, K–; widespread southeastern Temperate, rarely in
southwest………………………………………………….……..[Pyxine subcinerea]
170
30.(27) Thallus with crowded, overlapping lobes; cilia-like rhizines abundant along the margins
......................................................................................................... Phaeophyscia hispidula
30.
Thallus forming flat rosettes; marginal cilia or cilia-like rhizines present or absent........ 31
31.
Cilia-like rhizines present along the lobe margins……………..[Heterodermia casarettiana]
31.
Cilia-like rhizines or cilia absent...................................................................................... 32
32.
Medulla K+ yellow; rhizines sparse; zeorin present ................................ Physcia sorediosa
32.
Medulla K–; rhizines abundant; zeorin absent ................................................................. 33
33.
Cortex K+ yellow, UV- (atranorin); medulla PD+ orange (testacein); spores 4-celled
................................................................................................................ Pyxine eschweileri
33.
Cortex K–, UV+ yellow-orange (lichexanthone); medulla PD–; spores 2-celled ................
........................................................................................................................Pyxine cocoes
34.(23) Cortex K– (atranorin absent) ........................................................................................... 35
34.
Cortex K+ yellow (atranorin)........................................................................................... 36
35.
Lobes pruinose at least at the tips; western ......................................... Physconia thomsonii
35.
Lobes without pruina; mostly northeastern .....................................................................35a
35a.
Soredia entirely marginal or on lobe tips ......................................Physciella chloantha
35a.
Soredia on lobe surface or lobe margins; northeastern to Montana and Colorado
........................................................................................................[Physciella melanchra]
36.
Soredia laminal or on the upper surface of the lobe tips ................................................. 37
36.
Soredia mostly marginal, on the lower surface of the lobe tips, or on expanded lobe tips39
37.
Marginal cilia common and abundant; thallus loosely attached and often ascending; lobes 0.20.5 (-1) mm wide ......................................................................................... Physcia tenella
37.
Marginal cilia absent; thallus closely attached to substrate; lobes mostly 0.5-1.5(-2.5) mm
wide.................................................................................................................................. 38
38.
Lower surface and rhizines pale brown; soredia blue-gray to white; thallus moderately thick;
mainly on rock, rarely wood or bark ............................................................. Physcia caesia
38.
Lower surface and rhizines white or almost white; soredia greenish or white; thallus thin and
membranous; mainly on bark of deciduous trees, rarely on limestone ..Physcia americana
39.(36) Marginal cilia or cilia-like rhizines common and abundant ............................................. 40
39.
Marginal cilia or cilia-like rhizines absent or sparse........................................................ 45
171
40.
Thalli closely appressed to substrate; soralia crescent-shaped; marginal cilia or cilia-like
rhizines branched (fig. 10b) ............................................................................................. 41
40.
Thalli loosely attached over entire substrate, or ascending and almost fruticose; soralia not
crescent-shaped; cilia branched or unbranched................................................................ 42
41.
Lower surface white to pale brown, K– or K+ yellow, smooth to fibrous (with fibers running
in one direction), with a cortex, at least in part……………………….Heterodermia speciosa
41.
Lower surface pale orange, at least in spots (white to pale brown elsewhere), K+ purple where
pigmented (anthraquinones), entirely without a cortex……………..Heterodermia obscurata
42.
Cilia short, 1-2 mm long, unbranched; lobes relatively short, forming crowded clumps; lower
surface smooth, with a cortex; soralia on expansions of lobe tips………………………43
42.
Cilia 2-5 mm long, branched or unbranched; lobes very long and narrow; lower surface
cottony or webby, often pruinose, lacking a cortex; soredia on lower surface of lobe tips,
which do not widen .......................................................................................................... 44
43.
Soredia within hood-like expansions of lobe tips (between upper and lower cortices)........
............................................................................................................... Physcia adscendens
43.
Soredia on widened, turned-back lobe tips (lip-shaped soralia)……………Physcia tenella
44.
Lower surface usually pale sulphur yellow, at least close to lobe tips (K+ yellow, not an
anthraquinone); cilia white or brown, mostly unbranched; medulla PD–, K+ yellow (atranorin)
..................................................................................................Heterodermia appalachensis
44.
Lower surface white; cilia and rhizines white to pale brown, frequently branched; medulla
PD+ yellow, K+ yellow changing to red (salazinic acid)………..Heterodermia leucomelos
45.(39) Soredia produced all along lobe margins ......................................................................... 46
45.
Soredia confined to lobe tips or on crescent-shaped marginal lobes................................ 48
46.
Lower surface fibrous, more or less streaked with black or dark gray; thallus pruinose, at least
at lobe tips; rhizines brown to black; medulla K+ yellow………………….Physcia atrostriata
46.
Lower surface smooth, more or less uniformly white; thallus entirely without pruina; rhizines
white to pale brown, sometimes darkening at the tips; medulla K– or K+ red ................ 47
47.
Lobes rounded, thin, finely divided, the margins dissolving into granules (blastidia: fig. 19e)
or soredia; marginal cilia absent; medulla K–…………………………….Physcia millegrana
47.
Lobes elongated, moderately thick, not finely divided, with white granular soredia (not
blastidia) along the margins; marginal cilia sometimes present; medulla K+ yellow changing
to red (salazinic acid) ........................................................................Heterodermia albicans
172
48.(45) Rhizines black; upper cortex prosoplectenchyma (fig. 5e) .............................................. 49
48.
Rhizines white or pale tan, sometimes darkening at the tips, or rarely dark gray; upper cortex
pseudoparenchyma (fig. 5f).............................................................................................. 50
49.
Lower surface white to pale brown, smooth to fibrous, with a cortex (at least in part)
.......................................................................................................... Heterodermia speciosa
49.
Lower surface pale orange, at least in part (K+ purple, anthraquinones), webby, entirely
without a cortex.............................................................................. Heterodermia obscurata
50.
Lobes very narrow, 0.2-0.5 mm wide, hardly widening at the tips; soredia and blastidia very
coarse, forming under and at lobe tips; on siliceous rock……………………Physcia subtilis
50.
Lobes 0.3-1.2(-3) mm, usually widening or fanning out at the tips; soredia fine or coarse; on
various substrates ............................................................................................................. 51
51.
Lobes distinctly up-turned with soredia forming in lip-shaped soralia; soredia fine or coarse
.......................................................................................................................................... 52
51.
Lobe tips mostly flat or down-turned; soralia not lip-shaped; soredia coarsely granular. 53
52.
Soredia very fine, greenish; on rocks, wood, or sometimes bark, exposed or partly shaded;
very widespread and common........................................................................ Physcia dubia
52.
Soredia coarsely granular, blue-gray to white; on mossy rocks in forest habitats; Appalachian
Mountains.....................................................................................[Physcia pseudospeciosa]
53.
Lobes flat, finely divided and lacy, dissolving into granules (blastidia: fig. 19e) or soredia;
eastern temperate region; mostly on bark of various kinds, rarely on rock..........................
................................................................................................................ Physcia millegrana
53.
Lobes convex, clearly down-turned at the tips, with granular soredia forming on the lower
surface of the lobe tips; western; on rock, especially limestone and sandstone
...........................................................................................Physcia callosa sensu Thomson
[According to Moberg (Sonoran Flora), all specimens named as Ph. callosa sensu
Thomson are actually Physcia tribacia (Ach.) Nyl. The identity of Ph. callosa s. str. is still
unclear; it was described from California.]
PHYSCIELLA: See Phaeophyscia.
PHYSCONIA
1.
Soredia absent; apothecia common .................................................................................... 2
1.
Soredia present; apothecia rare .......................................................................................... 4
173
2.
On the ground or over rocks in arctic or alpine sites; thallus loosely attached over entire
surface ................................................................................................ Physconia muscigena
2.
On bark or rocks at low elevations; thallus closely appressed to substrate………………….3
3.
Isidia present, mostly cylindrical but sometimes flattened…………….Physconia elegantula
3.
Isidia absent, but often with lobules, especially on the apothecial margins
.............................................................................................................Physconia americana
4.(1)
Rhizines unbranched or dichotomously branched, pale tan to brown, mostly on or close to the
margins; lobes wrinkled or bumpy (rugose)……………………………..Physconia thomsonii
4.
Rhizines squarrose, black, more or less uniformly distributed; lobes smooth and even .... 5
5.
Soralia discrete, crescent-shaped or lip-shaped on the lobe margins, containing very coarse
soredia; lobe margins turned down ....................................................Physconia perisidiosa
5.
Soredia more or less continuous along the lobe margins; lobe margins turned up............. 6
6.
Medulla pale yellow, K+ yellow, KC+ orange-yellow (secalonic acid)
.........................................................................................................Physconia enteroxantha
6.
Medulla white, K–, KC– (secalonic acid absent) ............................................................... 7
7.
Upper cortex with thick-walled cells (scleroplectenchyma); mostly eastern North America
..................................................................................................................Physconia detersa
7.
Upper cortex with thin-walled cells (pseudoparenchyma); western North America............
.............................................................................................................. Physconia isidiigera
PILOPHORUS
1.
Stalks sorediate at the tip, under 5 mm tall ........................................... Pilophorus cereolus
1.
Stalks not sorediate, over 5 mm tall ................................................................................... 2
2.
Apothecia elongate, cylindrical; thallus olive to brown ........................ Pilophorus clavatus
2.
Apothecia spherical; thallus gray-green .............................................. Pilophorus acicularis
PLACIDIUM: See Key G.
PLACOPSIS: Keys C and F.
PLACYNTHIELLA: Keys C and F.
PLACYNTHIUM: See Keys F and G.
174
PLATISMATIA (including Esslingeriana)
1.
Lobes 0.5-5 mm wide, often elongate or finely divided and sometimes pendent…………2
1.
Lobes (3-)5-20 mm wide, rarely pendent........................................................................... 4
2.
Pycnidia on the thallus surface, common; thallus with a yellowish tint; medulla yellowish and
KC+ pink at least on lobe tips and apothecial margins…………..Esslingeriana idahoensis
2.
Pycnidia, when present, along the lobe margins; thallus without a yellowish cast, KC- ... 3
3.
Isidia absent..................................................................................... Platismatia stenophylla
3.
Isidia present along lobe margins............................................................. Platismatia herrei
4.(1)
Soredia and isidia absent.................................................................................................... 5
4.
Soredia or isidia present ..................................................................................................... 6
5.
Northeastern U.S., and southeastern Canada; medulla PD–……….Platismatia tuckermanii
5.
Western; medulla PD+ red (fumarprotocetraric acid) .........................Platismatia lacunosa
6.
Isidia and (or) soredia present on lobe margins; lobe surface relatively smooth, without
pseudocyphellae ...................................................................................... Platismatia glauca
6.
Isidia laminal; surface with a network of sharp ridges and depressions, with white
pseudocyphellae ................................................................................. Platismatia norvegica
PLEOPSIDIUM: See Acarospora.
POLYCHIDIUM: See Key B.
POLYSPORINA: See Key F.
PORINA: See Key D.
PORPIDIA (including Clauzadea and Melanolecia)
1.
Thallus orange............................................................................Porpidia flavocaerulescens
1.
Thallus gray........................................................................................................................ 2
2.
On limestone; arctic-alpine, rather rare; epihymenium brown or green............................. 3
2.
On noncalcareous rock; temperate to boreal, common; epihymenium brown to olive-brown
............................................................................................................................................ 4
175
3.
Apothecia red-brown when wet, very small (under 0.3 mm in diameter), flat, not pruinose;
margins sometimes prominent; hypothecium and exciple dark brown; epihymenium brown;
spores 6.5-12(-14) x 3.5-7 μm……………………………………….[Clauzadea monticola]
3.
Apothecia black when wet, 0.4-1 mm in diameter, flat, prominent margins; disk sometimes
lightly pruinose; epihymenium green to blue-green, at least in part; hypothecium and exciple
coal-black, not distinguishable; spores mostly 13-28 x 7-14 μm .. [Melanolecia transitoria]
4.
Apothecial disks gray due to a light or heavy pruina; apothecial margins black, contrasting
with the disk ............................................................................... Porpidia albocaerulescens
4.
Apothecial disks black, not pruinose; apothecial margin the same color as the disk ......... 5
5.
Apothecial margin brittle and radially cracked. Thallus endolithic; spores 12-18 x 6-8 μm;
Appalachian-Great Lakes-Ozarks distribution……………………..[Porpidia tahawasiana]
5
Apothecial margin not brittle or radially cracked............................................................... 6
6.
Apothecia usually less than 1.2 mm in diameter; hymenium 60-75(-100) μm high; spores 1017 x 5-9 μm; cells in the exciple about 5-8 μm in diameter (seen only in thin sections)
.................................................................................................................Porpidia crustulata
.
Apothecia commonly 1-2.5 mm in diameter; hymenium 80-120 μm high; spores mostly 13-23
x 7-10 μm; cells in the exciple mostly 3-6 μm in diameter………….[Porpidia macrocarpa]
PROTOBLASTENIA: See Key F.
PROTOPARMELIA: See Key F.
PSEUDEPHEBE
1.
Branches very slender and entirely terete, 0.1-0.2 mm in diameter, not flattening at the tips;
distance between axils mostly 1-3 mm………………………………Pseudephebe pubescens
1.
Branches 0.2-0.5 mm in diameter, usually becoming somewhat flattened at the tips; distance
between axils 0.2-0.5(-1) mm ........................................................ Pseudephebe minuscula
PSEUDEVERNIA
1.
Thallus abundantly isidiate on the upper surface ..........................Pseudevernia consocians
1.
Thallus not isidiate ............................................................................................................. 2
2.
Lobes mostly under 1 mm wide; lower surface white except at the base; in the Appalachian
mountains ..........................................................................................Pseudevernia cladonia
2.
Lobes 1-3 mm wide; lower surface dark gray to black except at the lobe tips; southwestern
............................................................................................................... Pseuevernia intensa
176
PSEUDOCYPHELLARIA
1.
Pseudocyphellae and soralia yellow (calycin)…………………………………………….2
1.
Pseudocyphellae and soralia (if present) white (calycin absent) ........................................ 3
2.
Photobiont green; medulla dark yellow throughout; lobes smooth and even; medulla
PD–, K–; southeastern ............................................................ Pseudocyphellaria aurata
2.
Photobiont blue green; medulla white or yellow; lobes with a network of depressions and
sharp ridges; medulla PD+ orange, K+ yellow (stictic acid?); Appalachian-Great Lakes
region and west coast……………………………………………………………….………..2a
2a.
Medulla mostly white, except yellow near soralia and pseudocyphellae; soralia laminal
and marginal...........................................................................Pseudocyphellaria crocata
2a.
Medulla yellow throughout; soralia entirely marginal in western populations, but
laminal and marginal in the east ……………………………………………………
…………………………... Pseudocyphellaria perpetua McCune & Miadlikowska
3.
Thallus greenish gray or pale brown, dull or scabrose; lobes smooth and even; algal layer
grass-green; isidia and lobules present along the lobe margins; medulla PD–, K–
.............................................................................................. Pseudocyphellaria rainierensis
3.
Thallus dark reddish brown, rather shiny; lobes with a network of depressions and sharp
ridges; algal layer dark blue green; isidia and lobules absent; medulla PD+ orange, K+ yellow
(stictic acid)........................................................................................................................ 4
4.
Soredia absent; apothecia abundant ....................................... Pseudocyphellaria anthraspis
4.
Soredia present, mainly on ridges; apothecia rare.....................Pseudocyphellaria anomala
PSEUDOPARMELIA: See Parmelia.
PSILOLECHIA: See Key F.
PSORA
1.
Apothecia mainly at the margins of the squamules; squamules pruinose, at least in part . 2.
1.
Apothecia mainly on the surface of the squamules; squamules pruinose or not………….4
2.
Squamules strongly convex and fissured, yellow-brown to olive-brown
............................................................................................................... Psora cerebriformis
2.
Squamules more or less flat, smooth or slightly fissured, reddish or orange-brown.......... 3
3.
Squamules brick red to orange-brown, flat or slightly convex, with edges turned up
177
......................................................................................................................Psora decipiens
3.
Squamules pink to pinkish orange, depressed in the center, with edges turned down
......................................................................................................................... Psora crenata
4(1).
Thallus bright yellow or yellow-green (rhizocarpic acid in the cortex)………Psora icterica
4.
Thallus brown, lacking yellow pigments in the cortex....................................................... 5
5.
Squamules 0.4-1.5(-3) mm wide ........................................................................................ 6
5.
Squamules 2 mm wide or more.......................................................................................... 9
6.
Parasitic on the cyanobacterial lichen, Spilonema;, lower surface and margins of squamules
blackish or blue-green ...............................................................................Psorula rufonigra
6.
Not parasitic; lower surface and margins of squamules white or pale brown, not blackish green
............................................................................................................................................ 7
7.
Squamules not lobulate, the edges conspicuously white pruinose; medulla C–
.................................................................................................................. Psora himalayana
7.
Squamule margins divided into tiny round lobules, sometimes lightly pruinose on the surface,
but not along the margins; medulla C+ pink (gyrophoric acid)…………………………….8
8.
Squamules loosely attached, often ascending; California .............................. Psora pacifica
8.
Squamules closely attached and flat against the substrate; mostly high elevations in the Rocky
Mountains................................................................................................... [Psora montana]
9(5).
Squamules pruinose or whitened on the edges and sometimes the surface ..................... 10
9.
Squamules almost entirely without pruina ....................................................................... 13
10.
Medulla K+ red (norstictic acid). Squamules usually concave; southwestern U.S, on soil.
..................................................................................................................... [Psora russellii]
10.
Medulla K– ...................................................................................................................... 11
11.
Found in central and eastern temperate North America on rock; apothecia rusty brown
typically with peristent margins ..........................................................Psora pseudorussellii
11.
Found in western or northern North America on rock or soil; apothecia red-brown to dark
brown or almost black, soon becoming convex and marginless…………………………..12
12.
Squamules pale yellowish brown to pale reddish brown, 2-5 mm wide, scattered or contiguous
and crowded, occasionally overlapping, lightly pruinose on the surface and margins
..................................................................................................................Psora tuckermanii
178
12.
Squamules dark reddish brown, 1-3 mm wide, overlapping like shingles, only the edges
conspicuously white pruinose ................................................................. Psora himalayana
13(9).
Medulla of thallus C+ pink or red .................................................................................... 14
13.
Medulla of thallus C–....................................................................................................... 16
14.
Squamules pale gray, thick and somewhat convex ......................... Trapeliopsis wallrothii
14.
Squamules greenish brown or red-brown, thin or thick ................................................... 15
15.
Squamules olive to greenish brown, often white on the margin, thin, flat or curled inward and
cup-like when dry, ascending or standing on edge, (2-)3-6(-10) mm wide… Psora nipponica
15.
Squamules (including the margin) red-brown, folded or flat, often shingled, adnate or
ascending, 2-4(-6) mm wide. Apothecia black or very dark brown; on soil or rock in
California ................................................................................................ [Psora californica]
16(13).Squamules uniformly reddish brown, shiny, smooth or fissured; apothecia very dark brown to
black............................................................................................................. Psora globifera
16.
Squamules pale to medium brown usually with whitish margins, dull, somewhat fissured;
apothecia typically red-brown.......................................................................................... 17
17.
Western, on rock or soil; apothecia soon convex and marginless……….Psora tuckermanii
17.
Central and eastern regions, on rock; apothecia mostly flat, with long persistent margins,
finally convex and marginless.............................................................Psora pseudorussellii
PSOROMA: See Key G and Pannaria.
PSORULA: See Psora.
PUNCTELIA
1.
Lower surface and rhizines mostly black. .......................................................................... 2
1.
Lower surface and rhizines pale to dark brown.................................................................. 3
2.
Growing on rock; lobules absent; medulla KC+ red, C+ pink (gyrophoric acid)
.................................................................................................................. Punctelia stictica
2.
Growing on bark or, rarely, rock.......................................................................................2a
2a.
Lobes covered with lobules; medulla KC–, C–; Appalachians, Minnesota
......................................................................................................Punctelia appalachensis
2a.
Lobes with soredia; medulla KC+ pink, C+ pink (gyrophoric acid); Appalachians and
west coast.................................................................................................... [Punctelia borreri]
179
3.
Soredia present; medulla C+ red (lecanoric acid) ………………………………………..3a
3.
Soredia absent .................................................................................................................... 4
3a.
Soredia powdery, along the lobe margins and on the surface, lobules absent.; pycnidia
sparse and inconspicuous; apothecia rare ..........................Punctelia subrudecta s. lat.
3a.
Soredia coarsely granular, corticate to almost lobulate, on the upper surface of the lobes
associated with pseudocyphellae and cracks; common in Ozarks
.........[Punctelia punctilla sensu Adler (North American material = P. missouriensis)]
4.
Isidia absent; lobules common; pycnidia and apothecia commonly seen; medulla C+ red or
C–....................................................................................................................................... 5
4.
Isidia present, cylindrical, branched or unbranched; lobules absent; pycnidia and apothecia
infrequent; medulla C+ red (lecanoric acid).............................................. Punctelia rudecta
5.
Medulla C–, KC–, lichesterinic and protolichesterinic acids present; conidia rod-shaped;
central and eastern U.S. and adjacent Canada……………………………Punctelia bolliana
5.
Medulla C+ red, medulla KC+ red (lecanoric acid); lichesterinic and protolichesterinic
acids absent ................................................................................................................................................... 6
6.
Conidia long, thread-shaped; southwestern U.S ................................................... .Punctelia hypoleucites
6.
Conidia relatively short, bacilliform; central and eastern U.S. ............................................
................................................................[Punctelia graminicola (B. de Lesd.) Egan (syn. P. semansiana)]
PYCNOTHELIA: See Key A.
PYRENOPSIS: See Keys D, F, and I.
PYRENULA (based on Harris, More Florida Lichens, 1995). All species from the southeastern coastal
plain, except as noted.
1.
Spores muriform................................................................................................................. 2
1.
Spores mostly transversely septate (occasionally with a few cells divided longitudinally) .
............................................................................................................................................ 6
2.
Spores (2-)4(-6) per ascus, 100-150(-190) x 30-41 μm; ostioles off-center (not at the
summits), sometimes fused ..................................................................... [Pyrenula falsaria]
2.
Spores 8 per ascus; ostioles at summit or off-center .......................................................... 3
3.
Thallus orange or yellow, K+ purple (anthraquinones); spores 23-35 x 11-17 μm
..................................................................................................... Pyrenula ochraceoflavens
180
3.
Thallus not orange or yellow; K–....................................................................................... 4
4.
Perithecia fused in groups by their off-center or lateral ostioles; spores 45-65 x 18-27 μm
..............................................................................................................[Pyrenula ravenellii]
4.
Perithecia solitary, ostioles at summit................................................................................ 5
5.
Spores 45-60 x 16-22(-25) μm ..........................................................[Pyrenula leucostoma]
5.
Spores 30-40(-53) x 11-15 μm ........................................................[Pyrenula thelomorpha]
6(1).
Spores 27-38 x 12-18 μm .................................................................................................. 7
6.
Spores less than 25 μm long............................................................................................... 9
7.
Thallus and perithecial warts dark red (the pigment sometimes confined to the perithecial
warts)..........................................................................................................Pyrenula cruenta
7.
Thallus and perithecial warts greenish when fresh, whitish to brownish in herbarium...... 8
8.
Overmature old spores containing a red oily substance; spores 4- to 6-celled
........................................................................................................ [Pyrenula concatervans]
8.
Overmature old spores simply empty, collapsing; spores 4-celled
.............................................................................................................. [Pyrenula punctella]
9.
Spore walls at the tips of the spores thin .......................................................................... 10
9.
Spore walls at the tips of the spores obviously thick........................................................ 11
10.
East Temperate; no orange pigment on summits of the perithecia; thallus UV+ yellow
(lichexanthone)............................................................................... Pyrenula pseudobufonia
10.
Pacific Northwest near coast; scant or abundant orange pigment on summits of the perithecia;
thallus UV– or dull whitish ..............................................................[Pyrenula occidentalis]
11.
Perithecia fused into an extensive raised pseudostroma; spores 17-21 x 7-9 μm
............................................................................................................... [Pyrenula anomala]
11.
Perithecia solitary, not in a pseudostroma........................................................................ 12
12.
Thallus scurfy, whitish, without a cortex; spores 19-25 x 8-12 μm; perithecia 0.3-0.4 mm in
diameter; hemispherical, relatively thin walled………[Pyrenula microcarpa (syn. P. cinerea)]
12.
Thallus smooth, with a cortex; spores 15-21 x 5.5-8; perithecia 1-1.5 mm in diameter,
flattened-conical, thick walled………………………[Pyrenula mamillana (syn. P. marginata)]
PYRRHOSPORA: See Keys C and F.
181
PYXINE: See Physcia.
RACODIUM: See Key B.
RAMALINA (including Niebla)
1.
Thallus pendent, 5-30 cm (2-20 inches) long, often with clustered or indistinct attachment
points.................................................................................................................................. 2
1.
Thallus growing in bushy or almost pendent tufts from a single point, generally less than 6 cm
long .................................................................................................................................... 6
2.
Branches round in cross-section, very fine (mostly under 0.5 mm wide), the tips curled up
often producing a few granules or soredia ...............................................Ramalina thrausta
2.
Branches or stalks distinctly flattened, over 0.5 mm wide ................................................. 3
3.
Branches with expanded netlike tips ..................................................... Ramalina menziesii
3.
Branches not producing netlike tips ................................................................................... 4
4.
Soredia abundant, in marginal and often laminal soralia………….Ramalina subleptocarpha
4.
Soredia absent .................................................................................................................... 5
5.
Branches 3-10 cm long, 2-6 mm broad, more or less flat; western
............................................................................................................Ramalina leptocarpha
5.
Branches 30-50 cm long, 0.2-1.3(-2) mm broad, twisted, with long white pseudocyphellae
giving the branches a striped appearance; subtropical, Florida, Texas, and California........
...................................................................................................................[Ramalina usnea]
6.(1)
Soredia present; apothecia rare .......................................................................................... 7
6.
Soredia absent; apothecia common .................................................................................. 15
7.
Soredia white to blue-gray; soralia in rounded mounds, becoming wooly in old specimens;
branches mostly round in cross section, often spotted with black bands or dots. On coastal
shrubs, trees, and rarely rocks along the west coast………………………..Niebla cephalota
7.
Soredia yellowish green; soralia of various shapes, never becoming wooly; branches usually
flattened, sometimes round in cross section, never spotted with black bands or dots ........ 8
8.
Branches at least partly hollow; medulla loose and webby................................................ 9
8.
Branches solid throughout; medulla dense and compact.................................................. 10
9.
Branches with round to oval perforations, smooth................................... Ramalina roesleri
9.
Branches not perforate or lacerate, but with depressions or ridges……..Ramalina obtusata
182
10.
Soralia very irregular in shape, at or close to the lobe tips, which often appear torn, frayed, or
hood-shaped; evernic acid present ................................................................................... 11
10.
Soralia round, elliptical, or elongate; branch tips tapering, sometimes finely divided but not
generally torn, frayed, or expanded; evernic acid absent ................................................. 12
11.
Branches narrow or broad, 0.5-3 mm wide; soredia mainly on the lower surface of the branch
tips, not developing within hood-shaped expansions …………………..Ramalina pollinaria
11.
Branches broad, 1.5-4 mm wide; soredia developing in inflated, hood-shaped expansions of
the branch tips ......................................................................................... Ramalina obtusata
12.
Soralia often producing tiny, isidia-like branchlets; branches angular or round in section, or
flattened in part; subtropical to tropical regions…………………………Ramalina peruviana
12.
Soralia lacking isidia-like branchlets; branches mostly flattened; not southern…………….13
13.
Soredia granular, usually concentrated at the branch tips; mainly on rocks in forest habitats;
sekikaic acid present ........................................................................... Ramalina intermedia
13.
Soredia farinose, marginal or laminal, not concentrated at the branch tips; mostly on trees;
sekikaic acid absent.......................................................................................................... 14
14.
Soralia mainly linear, along the branch margins, but also elliptical on the branch surface;
branches (1-)2-4(-10) mm broad; soralia always PD–, K–; zeorin present
...................................................................................................... Ramalina subleptocarpha
14.
Soralia mainly elliptical, on the branch margins, and occasionally on the surface; branches
mostly 0.5-3 mm broad; soralia PD– or PD+ yellow or red, K– or K+ red; zeorin absent
................................................................................................................Ramalina farinacea
15.(6)
On coastal rocks in California; pycnidia abundant and conspicuous, black……………..16
15.
On trees, or rarely rocks in interior regions; pycnidia inconspicuous, brown……………17
16.
Branches distinctly flattened or angular in cross-section; slender, cartilaginous cords present
in medulla.................................................................................................... Niebla homalea
16.
Branches round in cross-section; lacking cartilaginous cords in medulla
................................................................................................................Niebla combeoides
17.
Branches at least partly hollow, with many perforations; medulla loose and webby
...............................................................................................................Ramalina dilacerata
17.
Branches solid, without perforations; medulla dense and compact.................................. 18
18.
Branches smooth or with elongate striations.................................................................... 19
183
18.
Branches with depressions and ridges or long grooves, or with verrucae, tubercles, papillae
.......................................................................................................................................... 20
19.
Branches flattened throughout; spores fusiform, 16-25(-31) x 3-5 μm
............................................................................................................. Ramalina stenospora
19.
Branches almost round in cross-section except for the base; spores elongate ellipsoid to almost
fusiform, 11.5-22 x 3-5 μm ................................................................. Ramalina montagnei
20.
Apothecia on the branch surface or margins .................................................................... 21
20.
Apothecia on or close to the tips of the branches ............................................................. 23
21.
Warts or tubercles abundant on the branches; divaricatic acid present
............................................................................................................Ramalina complanata
21.
Warts and tubercles absent; divaricatic acid absent ......................................................... 22
22.
Western North America; branches more or less even in width, tapering only at the tip ......
............................................................................................................Ramalina leptocarpha
22.
Southcentral U.S.A.; branches tapering at the base and at the tips……….Ramalina celastri
23.(20) Branches with small white tubercles; medulla K+ red or purplish red............................. 24
23.
Branches with depressions and ridges, or with long grooves, but without tubercles; medulla
K–..................................................................................................................................... 25
24.
Spiny, perpendicular branches usually present, although sparse; spores ellipsoid, medulla PD+
red or yellow, C– (salazinic, norstictic, or protocetraric acids)................. Ramalina willeyi
24.
Spiny, perpendicular branches absent; spores fusiform; medulla PD–, C+ pink to red or violet
(rapidly disappearing) (cryptochlorophaeic and paludosic acids) ......... Ramalina paludosa
25.
Branches broad and fan-shaped; pseudocyphellae depressed, between the veinlike ridges on
the lower surface; southwestern or northcentral U.S……………………..Ramalina sinensis
25.
Branches divided and relatively narrow; pseudocyphellae raised or level with surface;
southcentral or eastern U.S. and adjacent Canada………………………Ramalina americana
RHIZOCARPON
1.
Thallus brown, gray, or white; medulla IKI– or rarely IKI+ blue ..................................... 2
1.
Thallus distinctly yellow; medulla IKI+ blue .................................................................. 13
2.
Spores 2-celled ................................................................................................................... 3
2.
Spores 4-celled to muriform............................................................................................... 5
184
3.
Spores becoming dark brown or greenish. Thallus thick, areolate, pale to dark brown or graybrown; widespread boreal ........................................................................... [R. badioatrum]
3.
Spores colorless.................................................................................................................. 4
4.
Thallus brownish, usually thin ......................................................Rhizocarpon hochstetteri
4.
Thallus thick, chalk-white, thick or thin. Rocky Mountains to arctic
......................................................................................................... Rhizocarpon chioneum
5.(2)
Spores colorless.................................................................................................................. 6
5.
Spores dark brown or dark green when mature.................................................................. 9
6.
Thallus rusty orange; spores 4-celled or with the occasional longitudinal septum; mostly
eastern montane...................................................................................[Rhizocarpon oederi]
6.
Thallus white, gray, or brownish, not rusty; spores muriform ........................................... 7
7.
Thallus dark gray to gray-brown, thin, verruculose; no lichen substances; very common and
widespread ................................................Rhizocarpon reductum (syn. R. obscuratum)
7.
Thallus pale brownish gray to gray; containing lichen substances; Great Lakes region to east
coast ................................................................................................................................... 8
8.
Thallus continuous, rimose; medulla K+ yellow or K–, PD+ orange or PD– (stictic acid,
sometimes in traces)................................................................[Rhizocarpon concentricum]
8.
Thallus contiguous or dispersed areolate; medulla K+ red, PD+ yellow (norstictic acid)
.........................................................................[Rhizocarpon rubescens (syn. ?R. plicatile)]
9.(5)
Spores 1-2 per ascus; thallus thick, areolate; medulla IKI– ............................................. 10
9.
Spores 8 per ascus; thallus areolate, often scattered over a black prothallus; medulla IKI+ blue
.......................................................................................................................................... 12
10.
Areoles peltate (free at the margins), brown; mainly western coastal mountains
....................................................................................................... [Rhizocarpon bolanderi)
10.
Areoles convex, contiguous or dispersed, pinkish brown to gray .................................... 11
11.
Spores mostly 1 per ascus; temperate to boreal................................ Rhizocarpon disporum
11.
Spores mostly 2 per ascus; widespread, temperate to arctic
.....................................................................................................[Rhizocarpon geminatum]
12.(9)
Thallus brown to gray-brown; medulla or cortex C+ pink, K+ yellow or K– (gyrophoric acid
and sometimes stictic acid); very common and widespread …………….[Rhizocarpon grande]
185
12.
Thallus pale brownish gray; medulla and cortex C–, medulla K+ red (norstictic acid); borealmontane and arctic ..................................................................... [Rhizocarpon eupetraeum]
13(1)
Spores 2-celled, dark, 18-32 x 10-15 μm; epihymenium intense blue-green. Arctic to eastern
montane...................................................................................[Rhizocarpon eupetraeoides]
13.
Spores muriform; epihymenium brown to olive-brown ................................................... 14
14(1).
Thallus greenish-yellow, with crescent-shaped areoles surrounding some apothecia;
epihymenium unchanged or becoming more intensely green with K
..................................................................................................... Rhizocarpon lecanorinum
14.
Thallus lemon yellow, with apothecia sunken between the areoles; epihymenium becoming
violet or purple with K ..................................................................................................... 15
15.
Spores 28-60(-70) x 15-25 μm; hymenium greenish……………Rhizocarpon macrosporum
15.
Spores 24-40 x 11-16 μm; hymenium colorless....................... Rhizocarpon geographicum
RHIZOPLACA
1.
Thallus attached directly to rock ........................................................................................ 2
1.
Thallus growing on soil, unattached (vagrant) ................................................................... 5
2.
Thallus crustose, dispersed areolate with very convex areoles ; apothecia orange, pruinose
..................................................................................................... Rhizoplaca subdiscrepans
2.
Thallus attached by a central holdfast (umbilicate)............................................................ 3
3.
Apothecial disks orange, pruinose...................................................Rhizoplaca chrysoleuca
3.
Apothecial disks yellow, brown, olive, or black, with or without pruina........................... 4
4.
Lower surface rough, broken into areoles with white cracks; apothecial disks yellowish
brown, not pruinose; medulla PD+ orange (pannarin)……………………Rhizoplaca peltata
4.
Lower surface smooth; apothecial disks yellow-brown to greenish or black, pruinose; medulla
PD+ bright yellow, or less frequently PD– (with or without psoromic acid)
....................................................... Rhizoplaca melanophthalma
5.(1)
Apothecia usually abundant, yellow-brown to greenish black; lobes lacking whitish warts;
medulla usually PD+ yellow (psoromic acid)………………….Rhizoplaca melanophthalma
5.
Apothecia lacking; small white warts produced especially on the margins or tips of the lobes;
medulla PD–.........................................................................................Rhizoplaca haydenii
186
RIMELIA (including Canomaculina)
1.
Maculae scattered, not reticulate nor associated with a network of cracks; isidia cylindrical,
developing on the lobe surface; medulla KC+ red (norlobaridone)
....................................................................................................Canomaculina subtinctoria
1.
Maculae creating a network of fine white cracks at least on lobe tips ............................... 2
2.
Soredia and isidia absent; pycnidia and apothecia abundant....................... Rimelia cetrata
2.
Isidia or soredia present ..................................................................................................... 3
3.
Lobes with cylindrical to branched, sometimes granular isidia, occasionally ciliate at the
tips, mainly developing close to the margins........................................ Rimelia subisidiosa
3.
Lobes with soredia on the upper surface, especially on the lobe tips and margins
.................................................................................................................. Rimelia reticulata
RINODINA
1.
On soil, mosses, or dead vegetation; arctic-alpine ............................................................. 2
1.
On bark, wood, or rock; mostly temperate (but see couplet 17)......................................... 4
2.
Thallus lobed at margins; spores with uniformly thin walled at maturity, mostly smaller than
23 x 11 μ; thallus yellowish, particularly at margins………………….. Phaeorrhiza nimbosa
2.
Thallus granular to verrucose or disappearing, not lobed; spores with unevenly thickened
walls (Physcia-type); thallus, if present, typically reddish-brown, never yellow............... 3
3.
Spores 22-35 μm long; apothecia 0.6-1.5 mm in diameter, margins typically prominent, with
an expanded lower cortex; thallus granular to verrucose; sphaerophorin present, zeorin absent
.................................................................................................................. Rinodina turfacea
3.
Spores 17-26 μm long; apothecia 0.4-0.8 mm in diameter, margins even with disk or
becoming thin or disappearing in maturity, without an expanded lower cortex; thallus
generally indistinct; sphaerophorin absent, zeorin present..................... [Rinodina archaea]
4.
On rock............................................................................................................................... 5
4.
On bark or wood............................................................................................................... 10
5.
Maritime, mostly on shoreline rocks in salt-spray zone. Spores inflated at septum, more so in
K, (10-)14-15 x (6-)7-8.5 μm................................................................ [Rinodina gennarii]
5.
On non-maritime rocks ...................................................................................................... 6
6.
Spores with a conspicuous pigmented band around the middle near septum, with walls thin at
tips, 15-20 x 9-13 μm; on calcareous rock.; widespread from arctic southwards.................
............................................................................................................ [Rinodina bischoffii]
187
6.
Spores lacking a pigmented band, with walls thickened at tips; on siliceous rock ............ 7
7.
Thallus brownish to olive, cortex K– (atranorin absent) .................................................... 8
7.
Thallus creamy white to pale brownish gray, dispersed areolate to verrucose, cortex K+
yellow (atranorin)............................................................................................................... 9
8.
Spores 17-24 x 8-14 μm, locules becoming round; thallus dull, verruculose to verrucose or
rimose-areolate; common, central plains to eastern seaboard, in moist habitats
............................................................................................................ [Rinodina tephraspis]
8.
Spores 23-41 x 11-17 μm, locules angular, walls remaining thickened only at cell tips; thallus
more or less smooth, shiny; rare on rock (see couplet 18); northeastern. Rinodina ascociscana
9.
Spores 17-22 x 8-11 μm, Physcia-type, retaining thick angular end-walls with age; apothecia
not pruinose; western mountains, Arizona and California to Alberta ..................................
...........................................................................................................[Rinodina confragosa]
9.
Spores 19-31 x 9-16 μm, with a thick septum, locules becoming round in older spores;
apothecia often pruinose; west coast, California to Vancouver Island
................................................................................................................Rinodina bolanderi
10.(4)
Thallus edge distinctly lobed; lower cortex present ........................ Hyperphyscia syncolla
10.
Thallus edge indefinite or definite, not lobed; lower cortex absent ................................. 11
11.
Spores 12-32 per ascus......................................................................... Rinodina populicola
11.
Spores 8 per ascus ............................................................................................................ 12
12.
Spores thin-walled, walls even in thickness ..................................................................... 13
12.
Spores conspicuously thick-walled, walls unevenly thickened ....................................... 14
13.
Spores even in outline, not constricted; thallus rugose, rimose, or more or less smooth; coastal
plain, Massachusetts to Gulf of Mexico…….[Amandinea milliaria (syn. Rinodina milliaria)]
13.
Spores becoming slightly constricted at septa; thallus verruculose or areolate; northcentral to
northeastern North America, not coastal plain
............................................................ [Amandinea dakotensis (syn. Rinodina dakotensis)]
14.
Thallus white or creamy, well developed, K+ yellow (atranorin); maritime California to
southern British Columbia ............................................................................................... 15
14.
Thallus brown, or olive, thin or thick, K– (lacking atranorin); not along west coast ....... 16
15.
Apothecia 0.4-0.7 mm in diameter; spores 15-20 x 8-11 μm; thallus thin, areolate to coarsely
granular; pannarin present in epihymenium (PD+ orange crystals)
..................................................................................................... [Rinodina marysvillensis]
188
15.
Apothecia 0.6-1.5 mm in diameter; spores 19-31 x 9-16 μm; thallus very thick, verrucose to
areolate; pannarin absent........................................................................Rinodina bolanderi
16.
Arctic to boreal and montane ............................................................17 (see also couplet 3)
16.
Temperate (except for R. archaea) ................................................................................... 18
17.
Thallus granular to verrucose; spores 22-35 μm long; sphaerophorin present, zeorin absent
.................................................................................................................. Rinodina turfacea
17.
Thallus largely within substrate and not seen; spores 17-26 μm long; sphaerophorin absent,
zeorin present ......................................................................................... [Rinodina archaea]
18.
Spores 30-40 x 11-16 μm ................................................................... Rinodina ascociscana
18.
Spores 12-25 x 6-10 μm ................................................................................................... 19
19.
Thallus membranous, very thin and indistinct, or mostly within substrate ...................... 20
19.
Thallus clearly visible, granular to verrucose or areolate. Southern boreal ..................... 22
20.
Arctic to boreal and northwestern montane; apothecia superficial with well developed thalline
margins, at least when young; zeorin present......................................... [Rinodina archaea]
20.
Temperate or southern boreal, apothecia broadly attached or immersed, typically with poorly
developed thalline margins when young ......................................................................... 21
21.
Spores with two round to almost triangular lumina creating an hour glass shape; zeorin absent;
coastal plain, Massachusetts to Texas ................................................. [Rinodina applanata]
21.
Spores with angular lumina, at least when young, Physcia-type; zeorin present; AppalachianGreat Lakes distribution. On deciduous trees, especially sugar maples….[Rinodina subminuta]
22.
Thallus bluish gray, dispersed areolate, or with darker granules; apothecial disks pitch black;
epihymenium gray-blue, turning purple with K and red with nitric acid
..............................................................................................................[Rinodina colobina]
22.
Thallus pale greenish gray to brown, not bluish gray; apothecial disks brown to black;
epihymenium red-brown, unreactive with nitric acid ...................................................... 23
23.
Mature spores with thin apical walls (Physconia-type); contains zeorin ............................
............................................................................................ [Rinodina archaea (Ach.) Arn.]
23.
Mature spores with thickened apical walls (Physcia-type); lacks zeorin ............................
............................................................................ Rinodina freyi H. Magn. (syn. R. glauca)
ROCCELLA: See Key A.
189
ROCCELLINA: See Key F.
ROPALOSPORA: See Keys C and F.
SARCOGRAPHA: See Key E.
SARCOGYNE: See Key F.
SCHAERERIA: See Lecidea.
SCHIZOPELTE: see Key A.
SCOLICIOSPORUM: See Bacidia.
SIPHULA: See Key A.
SOLORINA
1.
Thallus consisting of two parts: a brownish to gelatinous crustose layer containing
cyanobacteria (blue-green in section), and scattered red-brown apothecia with squamulose
margins containing green algae.............................................................. Solorina spongiosa
1.
Thallus foliose, with cephalodia in the form of a discontinuous blue-green layer of
cyanobacteria below the green algal layer, or warts on the lower surface ......................... 2
2.
Lower surface and medulla bright orange; upper surface not pruinose; arctic-alpine
......................................................................................................................Solorina crocea
2.
Lower surface pale brown; medulla white; upper surface white pruinose or scabrose, at least at
the lobe margins; widespread..................................................................... Solorina saccata
SPEERSCHNEIDERA: See Keys A and K.
SPHAEROPHORUS
1.
Thallus 30-80 mm long, with a stout main stem and finer side branches; fruiting bodies
common; medulla IKI+ blue ......................................................... Sphaerophorus globosus
1.
Thallus usually under 20 mm long, dichotomously branched; fruiting bodies rare; medulla
IKI– .................................................................................................. Sphaerophorus fragilis
SPILONEMA: See Key B.
SPORASTATIA: See Key F.
190
SQUAMARINA: See Key G.
STAUROTHELE
1.
Spores 8 per ascus ............................................................................Staurothele diffractella
1.
Spores 2 per ascus .............................................................................................................. 2
2.
Thallus continuous, rimose-areolate, without any lobes; perithecia buried in thallus, producing
low or tall bumps; prothallus absent........................................................... Staurothele fissa
2.
Thallus areolate, the perithecia buried in separate areoles ................................................. 3
3.
On wet or, more rarely, dry rock; areoles containing perithecia are larger than areoles that are
sterile; thallus usually with a lobed or fibrous prothallus...............Staurothele drummondii
3.
On dry rock; fertile and sterile areoles about the same size; thallus lacking a fibrous or lobed
prothallus............................................................................................ [Staurothele areolata]
STEREOCAULON
1.
Stalks sorediate .................................................................................................................. 2
1.
Stalks without soredia ........................................................................................................ 4
2.
Stalks 20-40 mm tall; side branches and phyllocladia sorediate; cephalodia on the stalks
...................................................................................................Stereocaulon coniophyllum
2.
Stalks under 5 mm tall; soredia at stalk summit; cephalodia on crustose primary thallus . 3
3.
Soredial mass KC+ pink to violet (lobaric acid); primary thallus mostly granular
...........................................................................................................Stereocaulon pileatum
3.
Soredial mass KC–; primary thallus areolate ........................................ Pilophorus cereolus
4.(1)
Growing attached directly to rock ...................................................................................... 5
4.
Growing on soil (sometimes soil over rock) .................................................................... 10
5.
Phyllocladia consisting of convex to flat, more or less round squamules, with dark green
centers and pale margins ..............................................................Stereocaulon vesuvianum
5.
Phyllocladia uniform in color............................................................................................. 6
6.
Tomentum on stalks thick or thin, usually pinkish; cephalodia abundant, consisting of large
spherical granules more or less buried in the tomentum, or lumpy galls; phyllocladia
squamulose, often deeply lobed .................. rare, saxicolous forms of Stereocaulon grande
6.
Tomentum usually thin, or if thick, then gray, not pinkish; cephalodia common, sparse, or
absent ................................................................................................................................. 7
191
7.
Phyllocladia flattened and lobed, rarely coralloid; thallus prostrate and clearly dorsiventral, at
least at the margins of the colony; cephalodia absent; containing lobaric acid
.............................................................................................................Stereocaulon saxatile
7.
Phyllocladia mostly coralloid or granular, not flattened; thallus prostrate or erect, dorsiventral
or not; cephalodia forming grapelike clusters when present .............................................. 8
8.
Phyllocladia mostly in warty or granular clusters at the stalk tips like cauliflower; cephalodia
rare; contains porphyrilic acid; arctic-alpine, especially in Alaska
..................................................................................................... [Stereocaulon botryosum]
8.
Phyllocladia mostly coralloid, not especially concentrated at stalk tips; cephalodia frequent or
rare; lacking porphyrilic acid ............................................................................................. 9
9.
Thallus dorsiventral in part; thallus PD+ orange (stictic acid)
...............................................................................................Stereocaulon dactylophyllum
9.
Thallus more or less erect, not dorsiventral; thallus PD– (lobaric acid)
.................................................................................................... Stereocaulon tennesseense
10 (4). Stalks usually under 2 cm high; phyllocladia warty, sometimes lobed ............................ 11
10.
Stalks mostly 2-8 cm long, erect or mat-forming, with distinct main stems; stems not woody
or brittle; phyllocladia granular or squamulose, rarely coralloid ..................................... 13
11.
Primary thallus disappearing; stalks mat-forming, without main stems; stems woody, brittle;
arctic-alpine.....................................................................................Stereocaulon rivulorum
11.
Primary thallus persistent, granular to squamulose; stalks usually erect and distinct ...... 12
12.
Mostly northeastern, Great Lakes to east coast, with scattered boreal localities farther north
and west; cephalodia black, with a fuzzy or grainy surface, containing Stigonema (fig. 1a)
................................................................................................. [Stereocaulon condensatum]
12.
Western montane and arctic; cephalodia brown, smooth and often fissured, containing Nostoc
(fig. 1d).........................................................................................[Stereocaulon glareosum]
13.(10) Phyllocladia mostly granular, clustered; cephalodia in fibrous tufts, dark brown or olive-black,
abundant ........................................................................................... Stereocaulon paschale
13.
Phyllocladia mostly flat, deeply or shallowly lobed; cephalodia not fibrous, often sparse..
.......................................................................................................................................... 14
14.
Tomentum thick or sometimes thin, often pinkish; cephalodia common and conspicuous,
coarsely granular and covered with tomentum or lumpy and bare ......Stereocaulon grande
14.
Tomentum thick, puffy, creamy white; cephalodia tiny, dark blue-green granules buried in the
tomentum ......................................................................................................................... 15
192
15.
Thallus usually prostrate, forming dorsiventral mats; common in Pacific Northwest,
uncommon elsewhere...........................................................................Stereocaulon sasakii
15.
Thallus erect, not forming dorsiventral mats; common in the northeast and into the northern
and western boreal regions.......................................................... Stereocaulon tomentosum
STICTA
1.
Soredia produced on or close to the lobe margins.......................................... Sticta limbata
1.
Soredia absent, isidia present ............................................................................................. 2
2.
Isidia along the lobe margins or cracks in the thallus; surface of the lobes smooth and often
shiny........................................................................................................... Sticta beauvoisii
2.
Isidia covering the thallus surface, which is dull and granular in appearance
.................................................................................................................... Sticta fuliginosa
SULCARIA: See Alectoria.
TELOSCHISTES
1.
Branches producing granular soredia; apothecia rare........................ Teloschistes flavicans
1.
Branches without soredia; apothecia common ................................................................... 2
2.
Thallus in tufts up to 2 cm high; apothecial margins abundantly ciliate
...............................................................................................Teloschistes chrysophthalmus
2.
Thallus in tufts 3-7 cm high; apothecial margins without cilia
................................................................................................................. Teloschistes exilis
TEPHROMELA: See Key F.
THAMNOLIA: See Key A.
THELIDIUM: See Key D.
THELOMMA
1.
Thallus areolate to verrucose, not lobed, shiny, never pruinose; spores 1-celled; on rocks
.........................................................................................................Thelomma mammosum
1.
Thallus distinctly lobed at the margin, verrucose-areolate in the center, dull, often pruinose;
spores 2-celled; on wood and sometimes rocks……………………Thelomma californicum
193
THELOTREMA (based on Harris, More Florida Lichens, 1995)
1.
Thallus with sparse, thick isidia; spores brown, submuriform, 16-28 x 7-12 μm; coastal plain
........................................................................................................... [Thelotrema santense]
1.
Thallus not isidiate; spores colorless.................................................................................. 2
2.
Spores muriform, over 50 μm long; mainly along the northeastern and northwestern coasts,
rare in Florida................................................................................... Thelotrema lepadinum
2.
Spores only transversely septate, 8- to 15-celled, 25-45 x 6-8 μm..................................... 3
3.
Apothecia 0.2-0.35 mm across; thallus well developed, with a thick medulla; coastal plain
.........................................................................................................[Thelotrema lathraeum]
3.
Apothecia 0.4-0.6 mm across; thallus thin, with an imperceptible medulla; Appalachian to
coastal plain..........................................................................................[Thelotrema subtile]
THOLURNA: See Key A.
THYREA: See Keys A and I.
TONINIA
1.
Spores 4- to 8-celled celled, 23-42 μm long;...................................................................... 2
1.
Spores 1- to 2-celled, 12-24 μm long ................................................................................. 4
2.
Thallus heavily and coarsely pruinose; epihymenium gray, K+ violet; Rocky Mountains,
Yukon to New Mexico ............................................................................... Toninia alutacea
2.
Thallus without pruina; epihymenium greenish to brown or red-brown, K– or K+ red;
widespread in western U.S. and adjacent southern Canada................................................ 3
3.
Epihymenium reddish brown, K+ red; thallus dark olive-brown, rarely reddish brown
.................................................................................................................[Toninia ruginosa]
3.
Epihymenium green, K–; thallus pale to dark brown, not olive…………[Toninia squalida]
4.(1)
Thallus crustose, areolate to rimose-areolate, without pruina; epihymenium greenish brown or
green, K–; Colorado ..............................................................................[Toninia phillippea]
4.
Thallus squamulose, pruinose or not.................................................................................. 5
5.
Pruina entirely absent from the thallus and apothecia; areoles and squamules flat to convex,
with deep depressions and pores; epihymenium brown to green, K–
......................................................................................... Toninia tristis ssp. asiae-centralis
194
5.
Pruina present on the thallus and often the apothecia, sometimes sparse or patchy;
epihymenium gray to violet, K+ violet .............................................................................. 6
6.
Squamules very small, 0.4-1(-2) mm across, almost granular in appearance, often scattered,
covered with coarse pruina; arctic and south into the Canadian Rockies ..[Toninia arctica]
6.
Squamules medium to large, 1-4(-5) mm across................................................................ 7
7.
Pruina coarse and granular; southern Rockies................................... [Toninia subdiffracta]
7.
Pruina fine and powdery .................................................................................................... 8
8.
Squamules strongly convex and folded throughout; thallus not lobed at the margins or forming
rosettes; pruina sometimes confined to most exposed surfaces of squamules and often lacking
on the apothecia; on soil; very widespread and common .......................... Toninia sedifolia
8.
Squamules flattened and somewhat lobed at the margins, convex in the center; thallus often
forming rosettes; pruina uniform and dense on the thallus as well as the apothecia; on
calcareous rock; southern Rockies…………………………………………[Toninia candida]
TRAPELIA: See Key F.
TRAPELIOPSIS
1.
Thallus squamulose, thick, not producing soredia, pale gray; contains lecanoric acid; on
rocks and soil, California to Oregon ....... Trapeliopsis californica McCune & Camacho
(= most N. Am. specimens named as T. walrothii)
1.
Thallus areolate to verrucose, not squamulose; usually sorediate in part; contains gyrophoric
acid; widespread ................................................................................................................ 2
2.
Thallus pale gray to greenish white or pinkish; apothecia pink, brown, or dark gray,
sometimes with all colors represented on the same thallus or even the same apothecium; on
soil or wood..................................................................................... Trapeliopsis granulosa
2.
Thallus dark gray to greenish black; apothecia almost always dark lead gray, but occasionally
pale; almost exclusively on wood ...................................................... Trapeliopsis flexuosa
TREMOLECIA: See Lecidea.
TRYPETHELIUM (including Laurera, Astrothelium, and Pseudopyrenula), based largely on Harris
(1995), "More Florida Lichens." With the exception of T. virens (East Temperate), all found
in tropical or subtropical areas, especially southeastern coastal plain.
1.
Perithecia buried in a wartlike pseudostroma or in thallus................................................. 2
1.
Perithecia discrete, not buried in a pseudostroma or in thallus .......................................... 9
195
2.
Spores thin-walled, muriform, 200-270 x 30-40 μm, 4 per ascus……..Laurera megasperma
2.
Spores thick-walled, unevenly thickened, muriform or only transversely septate, mostly less
than 75 x 30 μm; spores 8 per ascus................................................................................... 3
3.
Spores 6- to 14-celled......................................................................................................... 4
3.
Spores 4-celled ................................................................................................................... 5
4.
Medulla of pseudostromata yellow to tan, K+ purple (anthraquinones); spores 10- to 14celled, 40-50 x 9-12 μm ................................................................ [Trypethelium eluteriae]
4.
Medulla of pseudostromata not pigmented, K–; spores 6- to 12-celled, 38-52 x 7-10 μm
.............................................................................................................. Trypethelium virens
5.
Perithecia grouped two or three together sharing a single ostiole, pruinose at least around the
ostiole, dusted with yellow or orange anthraquinones (K+ purple); spores 28-35 x 10-13 μm
..................................................................................................... [Astrothelium versicolor]
5.
Perithecia not sharing an ostiole and not pruinose; anthraquinones absent or present; spores
18-28 x 6-10 μm................................................................................................................. 6
6.
Perithecia superficial; thallus whitish, without a cortex; hymenium containing many oil drops;
spore locules with angular walls…………………[Pseudopyrenula diluta (syn. P. subnudata)]
6.
Perithecia entirely immersed in pseudostromata with only the ostioles visible; thallus dark
brownish to olive, with a cortex or below the bark cells; spore locules lens-shaped ......... 7
7.
Pseudostromata brown ................................................................ [Bathelium carolinianum]
7.
Pseudostromata thallus-colored or black............................................................................ 8
8.
Thallus orange, K+ purple...............................................................[Trypethelium aeneum]
8.
Thallus olive to pale yellowish olive, K–….[Trypethelium variolosum (syn. T. ochroleucum)]
9.(1)
Perithecia conical; thallus light brown to pale orange; excipulum not distinguishable from
involucrellum; perithecial cavity colorless; spore locules lens shaped; spores with a gelatinous
halo................................................................................................. Trypethelium tropicum
9.
Perithecia hemispherical; thallus whitish; excipulum distinct from involucrellum at least at
base, perithecial cavity yellowish; spore locules with angular walls, spores without a halo
........................................................................................................ [Pseudopyrenula diluta]
TUCKERMANELLA: See Tuckermanopsis.
196
TUCKERMANOPSIS (= “TUCKERMANNOPSIS”)
1.
Soredia produced along the lobe margins ............................. Tuckermanopsis chlorophylla
1.
Soredia absent .................................................................................................................... 2
2.
Lobes long and strap-shaped, flat; dichotomously branched, forming shrubby clumps; linear
elongate pseudocyphellae sometimes present along the margins
.................................................................................................... Tuckermanopsis subalpina
2.
Lobes rounded or somewhat elongated, flat to crinkled and crisped; branching usually
irregular; pseudocyphellae present or absent, never linear-elongate……………………….3
3.
Isidia present on lobe margins and sometimes upper surface, cylindrical to spherical;
apothecia absent; pycnidia sparse or very inconspicuous; lobes elongated..........................
............Tuckermanella coralligera (W. A. Weber) Essl. (syn. Tuckermanopsis coralligera)
3.
Isidia absent, although marginal lobules or warty tubercles sometimes produced; apothecia
abundant; pycnidia abundant and conspicuous; lobes rounded.......................................... 4
4.
Lobes flat............................................................................................................................ 5
4.
Lobes undulating or crisped at the margins........................................................................ 6
5.
Thallus brown, at least when dry, brown to olive when wet; pseudocyphellae sparse; apothecia
on or close to the lobe margins................................................... Tuckermanopsis sepincola
5.
Thallus olive when dry, grass green when wet; pseudocyphellae abundant and conspicuous;
apothecia on the lobe surface ...............................................................................................
................................... Tuckermanella fendleri (Nyl.) Essl. (syn. Tuckermanopsis fendleri)
6.
Warty tubercles and (or) lobules frequent on the lobe surface and margins; pseudocyphellae
abundant and conspicuous, especially on the tubercles; medulla (especially in the apothecial
margin) usually yellow or orange in spots…………………… Tuckermanopsis platyphylla
6.
Warty tubercles absent; lobules present or absent, pseudocyphellae absent; medulla white
throughout .......................................................................................................................... 7
7.
Medulla C–, KC–, UV– (protolichesterinic acid present); lobules often present on the lobe
margins........................................................................................... Tuckermanopsis orbata
7.
Medulla C+ pink or KC+ pink to red (protolichesterinic acid absent); lobules absent ...... 8
8.
Medulla C–, KC+ pink, UV+ blue-white (alectoronic acid)….. Tuckermanopsis americana
8.
Medulla C+ pink, KC+ red, UV– (olivetoric acid)……………… [Tuckermanopsis ciliaris]
197
UMBILICARIA (including Lasallia)
1.
Lobes pustulate or blistered, with depressions on the lower surface corresponding to pustules
on the upper surface ........................................................................................................... 2
1.
Lobes not pustulate: smooth and even, wrinkled, bumpy or with a network of depressions and
sharp ridges; if warty, the warts do not have corresponding depressions on the lower surface
................................................................................. 4
2.
Rhizines sparse; lobes usually crowded and overlapping, convex or undulating and crisped at
the margins......................................................................................Umbilicaria caroliniana
2.
Rhizines absent; lobes usually solitary and flat.................................................................. 3
3.
Lower surface black and coarsely papillate like very rough sandpaper; upper surface rather
smooth, with low pustules having sloping sides……………………….Lasallia pensylvanica
3.
Lower surface pale to dark brown, rather smooth or slightly roughened; upper surface very
rough, with abundant pustules having almost vertical sides………………Lasallia papulosa
4.(1)
Rhizines present, sparse or abundant ................................................................................. 5
4.
Rhizines absent................................................................................................................. 13
5.
Lower surface black; upper surface smooth and even........................................................ 6
5.
Lower surface pale to dark brown, pink, or gray; upper surface rough ........................... 10
6.
Thallus with crowded, strongly folded, overlapping lobes; rhizines sparse, thick and irregular;
southern Appalachians and Alaska..................................................Umbilicaria caroliniana
6.
Thallus relatively round and flat; rhizines abundant .......................................................... 7
7.
Thallus upper surface dull or mat, never shiny; rhizines developing out of a granular lower
surface and forming a velvety mat ..................................................................................... 8
7.
Thallus upper surface shiny; lower surface not granular; rhizines not forming a short velvety
mat...................................................................................................................................... 9
8.
Thallus mostly brown, thin and membranous or moderately thick, rather fragile
........................................................................................................Umbilicaria mammulata
8.
Thallus mostly gray, thick and stiff like cardboard, rather tough
.......................................................................................................... Umbilicaria americana
9.
Lower surface covered with irregular plates of tissue with short, highly branched rhizines
developing from the plate margins; rhizines confined to lower surface; apothecia abundant,
flat, usually angular, with concentric ridges; western montane........ [Umbilicaria angulata]
198
9.
Lower surface rough, but plates confined to area immediately around attachment; rhizines
unbranched or forked, developing from the lower cortex (not plates), often forming clumps on
the upper surface; apothecia rare, convex, round, with radiating ridges; western North
America and Newfoundland.......................................................... [Umbilicaria polyrrhiza]
10.(5)
Thallus dull gray to gray-brown, 2-8 cm across, with coarse granular soredia developing from
the breakdown of the upper cortex close to the down-turned margins; lower surface rough,
with abundant, tapered, unbranched or forked rhizines; apothecia very rare; mainly
northeastern, scattered elsewhere………………………………………..[Umbilicaria hirsuta]
10.
Thallus gray or brown, surface without soredia, but covered with crystal-like deposits at least
close to the umbilicus; apothecia often present................................................................ 11
11.
Lower surface gray; rhizines unbranched, smooth; apothecial disks convex with thick
concentric ridges ........................................................................... Umbilicaria proboscidea
11.
Lower surface pink to pale brown; rhizines dichotomously branched; apothecial disks more or
less flat, with or without concentric ridges....................................................................... 12
12.
Marginal cilia or cilia-like rhizines absent, but rhizines abundant on lower surface; apothecia
broadly attached and adnate; apothecial disks with central buttons of sterile tissue or smooth
and even; cortex KC+ red, C+ pink, although the reactions are sometimes faint and difficult to
see (gyrophoric acid)……………………………………………………..Umbilicaria virginis
12.
Marginal cilia or cilia-like rhizines common and abundant; nonmarginal rhizines sparse;
apothecia constricted at the base and raised; apothecial disks with concentric ridges of sterile
tissue; cortex KC–, C– (gyrophoric acid absent)...............................Umbilicaria cylindrica
13.(4)
Isidia present on thallus surface, mostly granular .................................. Umbilicaria deusta
13.
Isidia absent ..................................................................................................................... 14
14.
Upper surface of thallus pruinose, or with coarse, crystal-like deposits .......................... 15
14.
Upper surface of thallus entirely without pruina or deposits ........................................... 18
15.
Lower surface pruinose at least close to margins ............................................................. 16
15.
Lower surface not pruinose .............................................................................................. 17
16.
Apothecia adnate; apothecial disks with concentric ridges of sterile tissue; thallus thin to
moderately thick............................................................................ Umbilicaria proboscidea
16.
Apothecia raised; apothecial disks with central buttons of sterile tissue; thallus thick and stiff
.............................................................................................. Umbilicaria krascheninnikovii
199
17.
Upper surface bumpy but without a reticulate pattern of ridges; apothecia abundant, raised,
with smooth and even disks; medulla KC–, C–…………………………..Umbilicaria rigida
17.
Upper surface with a network of depressions and sharp ridges, or wrinkled; apothecia not
common, adnate, with disks having central buttons of sterile tissue; medulla KC+ red, C+ pink
(gyrophoric acid)............................................................................... Umbilicaria decussata
18.(14) Lower surface with a network of rough and papillate membranes .................................. 19
18.
Lower surface smooth, papillate, or tuberculate, without membranes ............................. 20
19.
Margins of the thallus perforated with irregular holes and forming finely divided lobes;
apothecial disks with concentric ridges; medulla PD+ orange, K+ yellow (stictic acid)
........................................................................................................... Umbilicaria torrefacta
19.
Margins of the thallus not perforated and lobed; apothecial disks with radiating ridges;
medulla PD–, medulla K– ......................................................... Umbilicaria muehlenbergii
20.
Thallus round and flat; upper surface at least partly covered with convex warts or areoles over
a smooth, black, basal layer ........................................................... Umbilicaria hyperborea
20.
Thallus forming cushions of crowded, overlapping lobes; upper surface smooth, not covered
with warts or areoles ........................................................................................................ 21
21.
Lower surface black; lobes margins turned up; apothecia rare…………Umbilicaria polyphylla
21.
Lower surface pale or dark brown, lobe margins turned down; apothecia abundant
................................................................................................................. Umbilicaria phaea
USNEA
1.
Thallus pendent .................................................................................................................. 2
1.
Thallus growing in bushy tufts from a single point, or almost pendent ........................... 11
2.
Surface of main branches rough and eroded, with the cortex thin and crumbling; main
branches almost undivided, but with short to moderately long perpendicular side branches
abundant all along the filaments…………………………………………..Usnea longissima
2.
Surface of main branches with a continuous cortex, with depressions and ridges, or with
tubercles or papillae, not at all eroded................................................................................ 3
3.
Axis pink to brown; medulla white or reddish, usually CK+ orange (diffractaic acid)...... 4
3.
Axis and medulla white ..................................................................................................... 5
4.
Medulla usually pink to red, dense or lax, 20-30%; older branches with abundant whitish
warts; cortex very thick and hard .................................................................. Usnea ceratina
4.
Medulla white, dense, 16-21%; whitish warts absent; cortex thin ………..Usnea trichodea
200
5.
Isidia present ...................................................................................................................... 6
5.
Isidia absent........................................................................................................................ 9
6.
In Florida; isidia mostly developing within soralia on branch tips; medulla K+ red (norstictic
and galbinic acids)...................................................................................... Usnea dimorpha
6.
Boreal and western, rarely in the Appalachians; isidia not developing within soralia, although
sometimes in small clusters; medulla K– or K+ red (but norstictic acid absent) ............... 7
7.
Branches smooth, without papillae, with circular, thickened cracks (fig. 37e) at least at the
base; medulla KC+ pink, PD+ red, K– (protocetraric acid)…………………Usnea hesperina
7.
Branches with papillae, lacking circular, thickened cracks; medulla KC–; PD– or PD+
yellow, K– or K+ red (lacking protocetraric acid) ........................................................ 8
8.
Surface often dented or pitted; thallus more or less uniform in color; cortex very thin and
fragile (cortex 7-10%, medulla 22-26%, axis 28-38%); medulla PD–, K–, or PD + yellow, K+
red (salazinic acid)…………………………………………………………….Usnea scabrata
8.
Surface more or less even, not pitted or dented; main branches dark, especially at the base;
cortex moderately thick and tough (cortex 7-10%, medulla 14-21%, axis 36-43%); medulla
PD+ yellow, K+ red (salazinic acid)……………………………………….Usnea filipendula
9.(5)
Medulla loose and webby. Branches dented and pitted, not divided into segments by
circular cracks ........................................................................................ Usnea cavernosa
9.
Medulla thin and compact............................................................................................. 10
10.
Branches strongly ridged, often broken into small segments, but not bonelike (with swollen
circular cracks); medulla PD+ yellow, K+ red, KC–, CK– (norstictic acid); rare, AppalachianGreat Lakes distribution……………………………………………………..[Usnea angulata]
10.
Branches terete, not dented, pitted, or ridged, with swollen circular cracks and bonelike
segments (fig. 37e) at least at the base; medulla PD+ red, K–, KC+ pink, CK + orange
(protocetraric acid and diffractaic acids) ……………………………………Usnea hesperina
11.(1)
Thallus with apothecia ..................................................................................................... 12
11.
Thallus without apothecia ................................................................................................ 15
12.
In central to eastern North America; medulla rather dense .............................................. 13
12.
In southwestern U.S.; medulla loose, with white cottony hyphae.................................... 14
201
13.
Medulla usually red, at least in part, or sometimes white; base of thallus not blackened;
medulla K– or K+ red (norstictic acid); papillae, if present, wide at the base; very common
....................................................................................................................... Usnea strigosa
13.
Medulla white; base of thallus blackened; medulla K+ red (salazinic acid); papillae usually
present, cylindrical; infrequent.................................................................. [Usnea subfusca]
14.
Spiny fibrils abundant and crowded on the back of the apothecia; base of thallus narrowly or
irregularly blackened or not blackened at all; medulla K-, PD- (salazinic acid absent)
.........................................................................................................................Usnea cirrosa
14.
Spiny fibrils very sparse on the back of the apothecia; base of thallus extensively blackened;
medulla usually K+ red, PD+ orange or yellow (salazinic acid)................. Usnea arizonica
15.
On rock............................................................................................................................. 16
15.
On trees or wood .............................................................................................................. 17
16.
Branches spotted with black bands, coalescing into more or less blackened tips; round soralia
at the branch tips, lacking isidia; northern arctic with rare occurrences in the mountains of
Washington and Oregon………………….[Usnea sphacelata (syn. Neuropogon sulphureus)]
16.
Branches uniform in color, without black bands; isidia developing in soralia; temperate
............................................................................................................... Usnea amblyoclada
17.
Branches clearly constricted at base and usually the axils, giving the branches an inflated
appearance (fig. 37d)........................................................................................................ 18
17.
Branches not constricted at base, more or less uniform in diameter ............................... 20
18.
Isidia absent; soredia in excavate soralia mainly at the branch tips…………..Usnea glabrata
18.
Both isidia and soredia developing within soralia, concentrated at the branch tips or not...
.......................................................................................................................................... 19
19.
Soralia scattered over branches, remaining discrete; thallus black at the base for 1-2 mm..
................................................................................................................ Usnea fragilescens
19.
Soralia concentrated at branch tips, becoming confluent; thallus usually more or less uniform
in color, but sometimes darkening at the base................................................Usnea cornuta
20.(17)Thallus cortex distinctly reddish to red-brown or orange, sometimes mottled
....................................................................................................................Usnea rubicunda
20.
Thallus cortex shades of yellowish green (usnic yellow), not reddish ............................. 21
21.
Axis and medulla pink or red (rarely white in U. ceratina) .............................................. 22
21.
Axis and medulla white ................................................................................................... 24
202
22.
Axis broad and hollow; medulla PD+ yellow, K+ red (norstictic acid)………Usnea baileyi
22.
Axis solid throughout; medulla PD–, K– (norstictic acid absent)…………………………23
23.
Conspicuous whitish warts or tubercles present on branch surface; medulla thick, loose or
dense; contains diffractaic acid .................................................................... Usnea ceratina
23.
Whitish warts absent; medulla thin and dense; lacking diffractaic acid
.....................................................................................................................Usnea mutabilis
24.(21)Isidia absent, soralia abundant ............................................................................................ 25
24.
Isidia present, either directly on the branches or in soralia .............................................. 26
25
Soralia excavate, wider than half the branch diameter; medulla K– or K+ red (salazinic acid);
base of thallus pale or blackened................................................................ Usnea lapponica
25.
Soralia level with branch surface or mounded, round and relatively small; medulla usually K+
yellow (stictic acid complex); base of thallus blackened……………….[Usnea glabrescens]
26.
Branches ridged, at least partly angular in cross-section; isidia present, not arising from
soralia; cortex thin and fragile............................................................................ Usnea hirta
26.
Branches not ridged, uniformly round in cross-section; isidiate soralia present or absent;
cortex thick or moderate, not fragile ................................................................................ 27
27.
Thallus not blackened at base; branches rather smooth, with few or inconspicuous papillae;
cortex much thicker than medulla and very hard (cortex 18-22%, medulla 6-10%, axis 4058%); medulla PD+ red, KC+ pink to red (protocetraric acid) .............. Usnea subscabrosa
27.
Thallus blackened at the base; main branches bumpy with abundant papillae; cortex thinner
than medulla (cortex 7-14%, medulla 13-25%, axis 33-45%); medulla PD–, or PD+ orange or
yellow, KC– (protocetraric acid absent)........................................................................... 28
28.
Isidia scattered or clustered, abundant or sparse, not normally arising from soralia; branches
relatively soft and pliable ......................................................................... Usnea filipendula
28.
Isidia mostly clustered in round soralia (together with soredia); branches relatively stiff
.......................................................................................................................................... 29
29.
Main branches more or less equally dichotomous, tips not twisted or contorted; medulla
usually K-, PD–, UV+ (squamatic acid), sometimes K+ dark yellow, PD+ orange, UV–
(thamnolic acid), salazinic acid absent………………………………..[Usnea subfloridana]
29.
Main branches in unequal dichotomies, tips twisted or contorted; medulla K+ red, PD+ yellow
to orange (salazinic acid)................................................................ Usnea diplotypus group
203
VERRUCARIA
1.
On dry rocks....................................................................................................................... 2
1.
On at least periodically submerged rocks........................................................................... 6
2.
Thallus dark brown to black; lower half of medulla black. Perithecia half to entirely immersed
in thallus; spores 14-24 x 7-11 μm…………………………………..[Verrucaria nigrescens]
2.
Thallis white, gray, or barely perceptible; medulla, if discernable, entirely white............. 3
3.
Thallus very thick, composed of continguous or scattered areoles, pale gray to bluish gray,
surface rough and scabrose; in the arid southwest
........................................................[Verrucaria inficiens; see Comments under Placidium]
3.
Thallus very thin, never areolate; widespread.................................................................... 4
4.
Thallus largely endolithic, forming only a whitish stain; perithecia tiny, 0.15-0.3 mm in
diameter, immersed in tiny pits in the rock; spores (18-)23-31 x 9-14 μm
..............................................................................................................Verrucaria calciseda
4.
Thallus usually visible on the rock surface; perithecia 0.25-0.4 mm in diameter………….5
5.
Thallus white; perithecia not more than half immersed in thallus or rock; spores ellipsoid,
18-28 x 9-14 μm...................................................................................[Verrucaria muralis]
5.
Thallus greenish gray; perithecia 2/3 immersed; spores narrowly ellipsoid, 16-24 x 8-10 μm
........................................................................................................ [Verrucaria calkinsiana]
6.(1)
In streams or on lake shores. Thallus smooth to rimose; perithecia partially immersed; black
involucrellum distinct from pale excipulum; spores 25-30 x 10-12 μm; widespread
......................................................................................................... [Verrucaria aethiobola]
6.
In salt water habitats in the intertidal zone......................................................................... 7
7.
Thallus brown, very thin and membranous, smooth, without any black bumps or ridges; spores
8-11(-12) x 4.5-6.5 μm; on both coasts……………..[Verrucaria halizoa (syn. V. microspora)]
7.
Thallus black to dark olive-brown when dry, thick, smooth or rough................................ 8
8.
Thallus distinctly lobed at the margins, with radiating black ridges; perithecia entirely
immersed and showing at the surface only as pale ostioles; spores almost spherical, 7-10 x
6.3-8 μm; frequent, from Vancouver Island to southeastern Alaska…..[Verrucaria epimaura]
8.
Thallus not at all lobed at the margins; black ridges and bumps present or absent; perithecia
immersed or somewhat prominent; spores ellipsoid .......................................................... 9
9.
Thallus rimose-areolate when dry, mostly black when wet, normally with a rough upper
surface caused by abundant carbonized pegs or elongate ridges; perithecia immersed but
204
creating bumps on the thallus surface; spores 10-20 x 6-10 μm; extremely common on both
coasts........................................................................................................ Verrucaria maura
9.
Thallus smooth or with inconspicuous cracks when dry, never areolate; green to greenish
black when wet, lacking any carbonized pegs or ridges; perithecia entirely immersed, not
creating bumps on the surface; spores 8-10(-12) x 4-8 μm; fairly common on both coasts
..............................................................................................................[Verrucaria mucosa]
VESTERGRENOPSIS: See Pannaria.
VULPICIDA
1.
Soredia present on the lobe margins; pycnidia sparse or very inconspicuous
..................................................................................................................Vulpicida pinastri
1.
Soredia absent; pycnidia abundant and conspicuous.......................................................... 2
2.
Growing on soil or moss; thallus lacking rhizines; apothecia rare…………Vulpicida tilesii
2.
Growing on bark or wood; thallus with sparse rhizines; apothecia abundant……………..3
3.
Lobes smooth, with rounded depressions, or bumpy; pycnidia prominent; eastern U.S
..............................................................................................Vulpicida viridis
3.
Lobes with a network of depressions and sharp ridges or wrinkled; pycnidia entirely buried in
thallus with just the ostiole showing; western montane………………..Vulpicida canadensis
XANTHOPARMELIA (including Arctoparmelia)
1.
Soredia present on upper surface of thallus in hemispherical mounds............................... 2
1.
Soredia absent .................................................................................................................... 3
2.
Upper surface dull, grayish towards center of thallus; lower surface dull, dark gray to black;
medulla KC+ red, UV+ white (alectoronic acid); arctic and northern boreal ......................
........................................................................................................ [Arctoparmelia incurva]
2.
Upper surface shiny at lobe tips, brownish toward center of thallus; lower surface shiny black;
medulla K+ yellow to red, PD+ yellow-orange (stictic and norstictic acids); southwestern
montane habitats and coastal Nova Scotia; rare………………..[Xanthoparmelia mougeotii]
3.
Isidia present ..................................................................................................................... 4
3.
Isidia absent........................................................................................................................ 9
4.
Lower surface and rhizines black; medulla PD+ orange, K+ yellow to red (norstictic and
stictic acids) ............................................................................... Xanthoparmelia conspersa
4.
Lower surface and rhizines pale to dark brown.................................................................. 5
205
5.
Medulla K+ yellow, orange, or red .................................................................................... 6
5.
Medulla K– ........................................................................................................................ 7
6.
Isidia mainly cylindrical, often branched, not globular; widespread, especially in northeast,
also southwest; medulla AI+ blue, PD+ orange, K+ dark yellow (stictic acid)
........................................................................................................... Xanthoparmelia plittii
6.
Isidia globular at least at first, later cylindrical; western; medulla AI-, PD+ yellow-orange, K+
red (salazinic acid) ......................................................................Xanthoparmelia mexicana
7.
Medulla PD–, hypoprotocetraric acid present………………………Xanthoparmelia weberi
7.
Medulla PD+ distinct yellow or red, hypoprotocetraric acid absent .................................. 8
8.
Medulla PD+ bright yellow (psoromic acid); western……………Xanthoparmelia lavicola
8.
Medulla PD+ red (fumarprotocetraric acid); common in southeast, especially the Ozarks
..............................................................................................[Xanthoparmelia subramigera]
9.(3)
Lower surface dark gray to white or rarely pale brownish, dull and velvety; old thalli forming
full or partial concentric rings; medulla KC+ red, PD–, UV+ white (alectoronic acid); mainly
boreal to arctic ................................................................................................................. 10
9.
Lower surface brown or black, shiny, not velvety; old thalli not normally forming concentric
rings; medulla KC–, PD+ yellow, orange, or red, UV– (alectoronic acid absent) ........... 11
10.
Lower surface gray.......................................................................... Arctoparmelia separata
10.
Lower surface white or pale brown............................................. Arctoparmelia centrifuga
11.
Growing on soil ............................................................................................................... 12
11.
Growing on rock .............................................................................................................. 13
12.
Lobes 1-2 (-4) mm wide, convex, but not strongly inrolled; thallus attached to soil and pebbles
................................................................................................ Xanthoparmelia wyomingica
12.
Lobes 1.5-5 mm wide, strongly inrolled forming tubes; thallus not attached (vagrant), on arid
soils .........................................................................................Xanthoparmelia chlorochroa
13.
Lobes convex; thallus very loosely attached, ascending; pycnidia absent and apothecia rare
................................................................................................ Xanthoparmelia wyomingica
13.
Lobes flat; thallus closely or somewhat loosely attached to substrate; pycnidia and apothecia
abundant ........................................................................................................................... 14
14.
Lower surface and rhizines black..................................................................................... 15
14.
Lower surface and rhizines pale tan or brown.................................................................. 17
206
15.
Medulla K– or brownish, PD+ red (fumarprotocetraric acid)……………..X. hypomelaena
15.
Medulla K+ yellow or red, PD+ yellow-orange............................................................... 16
16.
Thallus closely attached to rock and difficult to remove intact; lobes short and crowded, often
overlapping and finely divided; contains stictic and norstictic acids ...................................
...........[Xanthoparmelia angustiphylla (previously called X. hypopsila in North America)]
16.
Thallus loosely attached and easily removed; lobes strap-shaped; contains salazinic acid
................................................................................................. [Xanthoparmelia tasmanica]
17.(14) Medulla K–, PD+ red (fumarprotocetraric acid)…………….Xanthoparmelia novomexicana
17.
Medulla K+ yellow or red, PD+ distinct yellow or orange (fumarprotocetraric acid absent)
.......................................................................................................................................... 18
18.
Thallus loosely attached to substrate over entire surface; lobes square to truncated or
elongated, usually somewhat constricted just behind the tips; often spotted with maculae;
medulla AI– (salazinic acid)…………………………………….Xanthoparmelia somloensis
18.
Thallus closely appressed to substrate at least over most of the surface; lobes rounded, not
constricted behind the tips; maculae absent ..................................................................... 19
19.
Medulla AI– (salazinic acid); thallus very closely attached to substrate; western
......................................................................................................... Xanthoparmelia lineola
19.
Medulla AI+ blue (stictic and norstictic acids); thallus closely to moderately attached to
substrate; widespread in northeast and parts of the west……Xanthoparmelia cumberlandia
XANTHORIA
1.
Soredia absent; apothecia abundant ................................................................................... 2
1.
Soredia present; apothecia absent or occasional ................................................................ 8
2.
Thallus loosely attached over entire surface, forming round cushions of overlapping lobes3
2.
Thallus closely appressed to substrate, forming round, flat rosettes .................................. 5
3.
Rhizines absent (thallus sometimes attached by stout holdfasts); conidia ellipsoid; on rocks,
bark, and wood .....................................................................................Xanthoria polycarpa
3.
Rhizines abundant, both attached to the substrate and unattached; conidia rod-shaped; on
trees, rarely on rock ........................................................................................................... 4
4.
Spores mostly 13-15.5 x 5-7.5 μm, septum 1.5-4 μm wide; arid western interior
............................................................................................................. [Xanthoria montana]
207
4.
Spores mostly 15.5-18 x 7.5-9.5, septum 5-8.5 μm wide; widespread, but not in western
interior region........................................................................................ Xanthoria hasseana
5.(2)
Lobes mostly wider than 1 mm, rounded, flat or concave, clearly foliose; lower surface
smooth, with scattered long rhizines ......................................................Xanthoria parietina
5.
Lobes mostly narrower than 1 mm, short or elongated to linear, appearing crustose at least in
center; rhizines absent or very rare..................................................................................... 6
6.
Lower surface with a cortex, wrinkled; lobes convex; on rock
..................................................................................................................Xanthoria elegans
6.
Lower surface mostly without a cortex and appearing crustose; lobes convex or flat ....... 7
7.
On bark or wood. Lobes often pruinose, rather thick; apothecia mostly under 1.5 mm in
diameter, concave to finally flattened; common in California…………….[Xanthoria tenax]
7.
On rock....................................................................................... lobed species of Caloplaca
8(1).
Thallus very closely appressed to substrate, almost crustose, without rhizines; coarse soredia
(schizidia) laminal, originating from the disintegration of pustules; on rock
.....................................................................................................................Xanthoria sorediata
8.
Thallus loosely attached to ascending, clearly foliose or squamulose; rhizines present or
absent; soredia rarely laminal; on bark, wood, or rock ..................................................... 9
9.
Rhizines abundant; pycnidia buried in the thallus, not conspicuous ................................ 10
9.
Rhizines absent or very sparse; pycnidia conspicuous or not........................................... 13
10.
Soredia mostly greenish yellow, within crescent-shaped hood- or lip-like expansions of the
lobe tips between the upper and lower cortices.......................................... Xanthoria fallax
10.
Soredia yellow to orange, on the lobe margins or sometimes laminal, or on the lower
surface of the lobe tips, not between the upper and lower cortices............................ 11
11.
Mostly northcentral to northeastern North America; soredia marginal at first, not produced in
hood-shaped expansions; conidia always rod-shaped………………….Xanthoria ulophyllodes
11.
Western montane and California; soredia produced on lower surface of lobes ............... 12
12.
Soredia frequently produced in hood-shaped expansions; conidia both ellipsoid and rodshaped.................................................................................................. [Xanthoria oregana]
12.
Soredia on the lower surface of the edges of the lobe tips, which are not expanded or
hood-shaped; conidia all rod-shaped
....................................................Xanthoria fulva (northern morphotype with rhizines)
208
13.(9)
Lobes flattened to almost cylindrical, branched, erect, sometimes almost fruticose in habit;
soredia produced on or just under the lobe tips; pycnidia immersed and inconspicuous; conidia
ellipsoid, 2-3.5 μm long…………………………………………………Xanthoria candelaria
13.
Lobes flat to strongly convex, rounded or divided; soredia on the lower surface or margins of
the lobe tips ...................................................................................................................... 14
14.,
On bark. Lobes finely divided, 0.2-0.4 mm broad, often in little rosettes; soredia on lower
surface of lobe tips close to the edge; pycnidia prominent, resembling orange pimples;
conidia rod-shaped, 3-4 μm long............................................................ Xanthoria fulva
14.
On calcareous rock. Lobes mostly 0.4-2 mm broad, very convex, almost hood-like;
soredia spreading over lower surface of lobes; pycnidia conspicuous or not; conidia rodshaped, 4-5(-6) μm long. In the western mountains. ................ [Xanthoria mendozae]
XYLOGRAPHA: See Key E.
Lichen Photobionts
Key to some important lichen photobionts as seen in thallus sections or squash preparations (based
largely on Poelt 1969). Many photobionts lose some of their distinctive characters when lichenized,
and so this key will only work for the most unaltered forms. A more complete and up-to-date key can
be found in Nash et al., Lichen Flora of the Greater Sonoran Desert Region, volume 1, pp 44-47. 2002.
1.
Cells grass-green or yellowish green; containing at least one green pigment-containing body
(chloroplast), making the cell wall distinct ........................................................................ 2
1.
Cells blue-green or dark green without a yellowish tint; without chloroplasts so that the green
pigments fill the cell and the cell wall (apart from any gelatinous outer envelope) is thin and
indistinct............................................................................................................................. 7
2.
Cells ellipsoid or cylindrical, not round ............................................................................. 3
2.
Cells basically round, although sometimes irregular in shape; chloroplasts filling the cell or
lying against one side ......................................................................................................... 5
3.
Cells with 2-7 chloroplasts; occasionally forming branched filaments; in a few species of
Verrucaria........................................................................................................Heterococcus
3.
Cells with a single chloroplast lying against one side of the cell ....................................... 4
4.
Cells ellipsoid, not curved; never forming filaments; in
Baeomyces, Icmadophila, Omphalina, Peltigera, among others ...................... Coccomyxa
4.
Cells squarish or cylindrical, sometimes slightly curved and bean-shaped; occasionally
forming filaments; in Chaenotheca ................................................................. Stichococcus
209
5.
Cells often bumpy in outline and not perfectly round, containing orange oil drops; sometimes
forming filaments of 2 to many cells; cell wall rather thick, refractive in polarized light; in
many lichens, e.g., Graphis, Pyrenula, Coenogonium, and Racodium ............ Trentepohlia
5.
Cells almost perfectly round, without orange oil drops; occuring singly or in clumps, not
forming filaments; cell walls not refractive in polarized light ........................................... 6
6.
Chloroplast central, with a lobed or scalloped margin; a relatively large seed-like pyrenoid
(rarely up to 3) clearly visible in the center of the chloroplast (appearing like a nucleus); the
most common lichen photobionts
...........................................Trebouxia or Pseudotrebouxia (distinguishable only in culture)
6.
Chloroplast central or on the side, not scalloped; pyrenoids small and inconspicuous
........................................................................"Chlorococcoid algae" or "unicellular algae"
(including many common photobionts such as Myrmecia, Dictyochloropsis, "Pleurococcus",
micareoid algae, etc., distinguishable with certainty only in pure culture).
7(1).
Cells solitary or grouped together in small packets or colonies, not forming filaments..... 8
7.
Cells forming distinct filaments ....................................................................................... 10
8.
Cells in groups of 2 to 4, 15-40 μm in diameter, surrounded by a rather thin and poorly
differentiated, gelatinous envelope that is single-layered or indistinctly layered; rather rare
photobiont in North America; in Phylliscum …………………………………Chroococcus
8.
Cells in groups of 2 to 8, mostly under 10 μm in diameter; surrounded by distinct, thick, often
pigmented and multi-layered, gelatinous envelopes........................................................... 9
9.
Gelatinous envelopes reddish or violet, K+ violet; common photobiont, especially in lichens
inhabiting non-calcareous rocks, e.g., Pyrenopsis, and in cephalodia
............................................................................................... Gloeocapsa in the strict sense
9.
Gelatinous envelopes yellowish brown, K-; common in lichens on calcareous substrates, e.g.,
Thyrea, Lichinella……………,…………"Xanthocapsa" (now included within Gloeocapsa)
10(7).
Filaments made up of cells in more than one row, commonly branching from the main
filament ("true branching"); in Ephebe, Spilonema, and the cephalodia of many lichens……
............................................................................................................................. Stigonema
10.
Filaments made up of cells in single rows, although sometimes clustered together ........ 11
11.
Filaments unbranched, not tapering, made up of round cells generally 3-6 μm in diameter,
appearing like a string of beads (as in Collema and Leptogium), or bunched up in tight coils
(as in Pannaria); heterocysts scattered along the filaments, not restricted to the tips; the most
common cyanobacterial photobiont…………………………………………………..Nostoc
210
11
.Filaments sometimes branched or tapering, made up of cells greater than 5 μm in diameter,
round or frequently squarish, broader than long; filaments commonly clustered into bundles
that split and branch off at different points ("false branching") ....................................... 12
12.
Heterocysts always at the tips of the filaments, which tend to be tapered (sometimes only very
slightly); mainly seen in cephalodia……………….Rivularia, Calothrix and related genera
12.
Heterocysts scattered along the filaments, which are not at all tapered; fairly common
photobiont, e.g., Coccocarpia and Polychidium..................................................Scytonema
Glossary
adnate. Tightly attached to the surface, like species of Dirinaria or Bulbothrix.
AI. A buffered iodine solution used for detecting stictic acid. Mix 1.5 ml of 20% Lugol's solution with
18.5 ml of pH 11.0 buffer.
alga (-gae). Green photosynthetic organism containing chloroplasts and nuclei and belonging to the
Kingdom Protoctista.
algal layer. Layer of algal cells in a stratified lichen thallus (fig. 4). Sometimes used synonymously
with "photobiont layer" (a layer of either green algae or cyanobacteria).
amphithecium. The portion of a lecanorine apothecium external to the exciple (fig. 13c), usually
containing algae, constituting the thalline margin.
amyloid. Containing carbohydrates that turn blue in an iodine solution (IKI).
anastomosing. Forming a netlike, interconnected growth (fig. 16c).
anisotomic. Dividing in unequal dichotomies to produce a distinguishable main branch with side
branches, as in Cladina rangiferina.
apical. At the tip or summit (i.e., the apex).
apothecium (-cia). A disk- or cup-shaped ascoma, usually with an exposed hymenium (figs. 12, 13).
appressed. Flattened and closely adnate; as in Xanthoparmelia cumberlandia.
areolate. Broken up into areoles, often appearing tile-like (fig. 2b; Lecidea tesselata).
areole. a) A small, irregular, often angular patch of thallus; b) a patch of vegetative tissue (containing
algae and cortex) on the podetial surface of some Cladonia species.
ascending. Lifting from the surface and becoming free from it, at least in part, like the lobes of species
of Tuckermanopsis.
ascocarp. See Ascoma.
ascohymenial. Pertaining to a type of ascoma having a hymenium with true paraphyses rather than
pseudoparaphyses (figs. 13a-e).
ascolocular. Pertaining to a type of ascoma in which the asci arise within a uniform mass of fungal
tissue and are separated in maturity, not by true paraphyses, but by highly branched
pseudoparaphyses (fig. 13f).
211
ascoma (-mata). The fruiting body of an Ascomycete; the structure that bears the asci, which in turn
contain the ascospores (figs 12, 17). Apothecia and perithecia are types of ascomata.
ascomycete. A fungus that produces its sexual spores within an ascus.
ascospore. A spore produced in an ascus (figs. 14, 15).
ascus (asci). The sac-like structure in Ascomycetes in which the ascospores are formed (fig. 14). The
sexual fusion of nuclei and reduction division occur within the ascus.
axial body. Conical, vertically oriented, nonamyloid structure in the tholus of certain types of asci such
as the Bacidia- and Biatora-types (figs. 14a, b); also called an apical cushion or masse axiale.
bacilliform. Stick- or rodshaped; cylindrical, usually straight.
basidiomycete. A fungus that produces its sexual spores as external buds on a clublike basal cell (the
basidium). Mushrooms, bracket fungi, and coral fungi, among others, belong to the
Basidiomycetes, one of the main classes in the Kingdom Fungi.
basidium. A clubshaped cell in the fruiting body of a Basidiomycete in which sexual fusion and
reduction division occurs, with the subsequent budding off of 2 to 4 spores (basidiospores).
biatorine. Referring to a type of apothecium having a relatively soft, clear or lightly pigmented (not
carbonized) margin containing no photobiont cells (fig. 13b).
blastidium (-dia). A granule-sized fragment of a lichen thallus that is formed by a budding off of the
thallus margin (fig. 19e).
blue-green algae. Cyanobacteria.
boreal. In reference to a northern region dominated by conifer forests; in North America, the region
forms a belt from the maritime provinces to Alaska.
bryophytes. Mosses, liverworts (hepatics), and hornworts.
C. Bleaching solution (sodium hypochlorite) or undiluted commercial bleach (e.g., Clorox
or
Javex ), used as a reagent in spot tests for revealing certain lichen substances (see table 2,
pages 106-107 in Lichens of North America).
calcareous. Containing lime or chalk (calcium carbonate), producing vigorous bubbling (CO2) in the
presence of a strong acid. Calcareous rocks include limestone, dolomite, and marble; some
sandstones and soils can also be calcareous.
campylidium (-dia). Helmet-shaped conidia-bearing structure found in many tropical, foliicolous
lichens.
capitate. Referring to a type of rounded, almost hemispherical structure (usually a soralium); see
Melanelia sorediata.
capitulum. The tiny spherical or cup-shaped apothecium formed at the summit of a slender stalk; found
in members Calicium, Chaeonotheca and related genera (fig. 12f).
carbonaceous. Opaque black, usually brittle, used in reference to tissue such as the exciple. It is hard to
distinguish individual cells in carbonaceous tissue.
cartilaginous. Tough, pliable cartilage- or sinew-like tissue. The term is usually used in reference to
supporting tissue (see fig. 8).
cephalodium (-dia). A small gall-like growth that contains cyanobacteria and occurs within the tissues
or on the surface of some lichens with green algal photobionts (plate 16; Placopsis lambii).
212
Chlorococcales. An order of green algae having taxa with individual spherical cells.
chloroplast. The structure in a green cell that contains chlorophyll, the substance responsible for
photosynthesis.
Chroococcus. A genus of cyanobacteria with large cells in colonies of 2 to 4 in a gelatinous, layered or
unlayered, relatively thin colorless envelope (fig. 1b). Individual cells do not have a clearly
defined gelatinous sheath as in Gloeocapsa.
cilia. Hair-like appendages on the margins of the thallus or apothecia of many foliose and fruticose
lichens (fig. 10; plate 13).
Coccomyxa. A genus of green algae with small, ellipsoid cells that contain a single chloroplast lying
against one wall.
columella. A vertical projection or axis of sterile tissue found in some ascomata.
conidium (-dia). An asexual spore usually formed in large numbers within special structures such as
pycnidia and campylidia. Conidia sometimes serve as male sexual cells (spermatia) (fig.18).
consoredium. A round cluster of tiny soredia that resembles a large soredium or granule.
continuous. Unbroken, or broken very little by cracks (fig. 2c; Lecanora caesiorubella).
coralloid. Composed of or having minutely branched cylindrical outgrowths, as in
Caloplaca coralloides or Sphaerophorus globosus.
cortex. The outer protective layers of a lichen thallus or apothecium, completely fungal in composition,
often composed of hyphae with thick, gelatinized walls (figs. 4, 13).
corticate (-ical). Having a cortex; pertaining to a cortex.
Corticolous. Growing on bark.
crenulate. Having a scalloped margin with rounded teeth or lobes (fig. 3).
crisped. Having a ruffled, wavy, or twisted margin, as in many species of Cetrelia and Parmotrema.
crustose. A thallus type which is generally in contact with the substratum at all points and lacks a lower
cortex; cannot be removed intact from its substrate without removing a portion of the substrate
as well (figs. 2a-d; plate 10).
cryptolecanorine. a) A kind of apothecium that is mostly sunken into the thallus and thereby lacks a
prominent margin; b) the partial, alga-containing margin in such apothecia (e.g., Aspicilia or
Ionaspis).
cudbear. The Scottish name for some lichens used for fermentation dyes, especially the mainly
European crustose species Ochrolechia tartarea.
cuff-shaped. In reference to laminal soralia that burst through the upper cortex of a hollow lobe,
leaving a hole in the center of the soralium (fig. 20f), seen in Menegazzia terebrata.
cyanobacteria. Photosynthetic, chlorophyll-containing organisms related to bacteria (in the Kingdom
Monera), without organized nuclei or chloroplasts; sometimes called blue-green algae.
cyanolichen. A lichen with cyanobacteria as the photobiont.
cyphella (-ae). A specialized, depressed pore in a lichen thallus, lined with small, loosely packed,
spherical cells (fig. 11c; plate 15); characteristic of the genus Sticta.
decorticate. Having had a cortex which has now fallen away or decomposed.
dichotomous. Branching into two equal parts, as in the letter "Y" (see, for example, Speerschneidera).
213
disjunct (-tion). A population that is geographically remote from other occurrences of the same species
(as in the distribution pattern of Umbilicaria caroliniana).
dorsiventral. With distinguishable upper and lower surfaces.
effigurate. Having a definite, usually somewhat lobed margin (fig. 2d), as in Dimelaena oreina.
ellipsoid. Oval in outline; more or less football-shaped (figs. 15a-d).
endemic. Found only in a certain, usually limited, region; in this book, used in reference to lichens
found only in North America or smaller regions, e.g., the Appalachian Mountains (see map of
Hypotrachyna croceopustulata).
endolithic. Growing within the upper layers of a rock, i.e., under and around the rock crystals, often
with little or no thallus visible on the outer rock surface (plate 32).
endophloeodal. Growing largely within the upper layers of bark tissue (fig. 9b), as in the thallus of
Graphis scripta.
epihymenium. The uppermost portion of the hymenium formed by the expanded tips of paraphyses;
often pigmented and sometimes containing tiny granules (fig. 13); considered here
synonymous with epithecium.
epispore. (see Perispore)
epithecium. See epihymenium.
excavate. Hollowed out or depressed; often in reference to a type of soralium (fig. 20h).
exciple. An area in an apothecium external to and below the hypothecium, forming the apothecial
margin in lecideine or biatorine apothecia (figs. 13a-b); much reduced in lecanorine apothecia
(fig. 13c).
excipulum. In this book, refers to the perithecial wall (fig. 17); often considered synonymous with
exciple.
f. Form; a formal subdivision of a species, subspecies, or variety usually applied to a relatively minor
morphological, chemical, or ecological variant.
family. A taxonomic category consisting of closely related genera.
farinose soredia. Very fine, powdery soredia, as on the podetial surface of Cladonia deformis.
fibril. A short branch, usually perpendicular to the main filament, in Usnea (fig. 37a; see U. diplotypus
U
or U. longissima).
U
filamentous. Hairlike.
fissural. Resembling a gaping slit, often oval or fusiform in shape; usually in reference to a type of
soralium (fig. 20d).
fixed nitrogen. Nitrogen from the air that has been chemically transformed into nitrogen-containing
compounds such as nitrates that plants can use for growth.
flexuose. Wavy.
flocculent. Having a loose, cottony, fibrous texture, like the nubby surface of an old sweater; as seen,
for example, on the surface of Cladina portentosa.
foliicolous. Growing on leaves of vascular plants (especially in the tropics; e.g., Calopadia fusca).
foliose. Pertaining to a more or less "leafy" lichen thallus, distinctly dorsiventral, and varying in its
attachment to the substrate from completely adnate to umbilicate (figs. 2f-g; plate 9).
214
fruiting body. The sexual reproductive structure of a lichen fungus (apothecium, perithecium,
mushroom, etc.); in most lichens, the ascoma.
fruticose. Pertaining to a lichen thallus that is stalked, pendent, or shrubby, and normally with no
clearly distinguishable upper and lower surfaces (figs. 2h-j; plate 9).
fusiform. Narrow, tapering at both ends, usually with pointed tips; spindle- or cigar-shaped (fig. 15f).
genus. A group of closely related species, presumably with the same ancestor; comprises the first word
of the two-word name of every organism.
Gloeocapsa. Cyanobacteria consisting of groups of 2 to 8 spherical cells enclosed within a thick
gelatinous matrix. Individual cells have their own, often colored, gelatinous sheaths that
usually are K+ purple (fig. 1c).
granular (-ose). (a) Having granules or granule-like particles (as in the thallus of Bacidia rubella). (b)
Pertaining to soredia that are large enough to be easily distinguished under a dissecting
microscope (as in Cladonia chlorophaea); see farinose.
granule. A spherical or nearly spherical corticate particle.
graphid. A lichen with a lirellum-type apothecium, resembling Graphis.
halo. A transparent gelatinous covering, often irregular in thickness, surrounding an ascospore (fig.
15k); technically called a perispore, and referred to as an epispore by some lichenologists;
revealed by an India ink preparation (see Microscopic Study, Chapter 13 in LNA).
halonate. Having a gelatinous perispore or "halo" (fig. 15k).
haustorium (-ria). A special branch of a mycobiont that penetrates or otherwise attaches itself to the
photobiont cell for the purpose of food absorption.
heath. A vegetation type consisting of extensive areas of low shrubs and few trees.
hemiamyloid. Turning dark blue with iodine (IKI) when pretreated with KOH, but negative or redorange with IKI in tissues not pretreated with KOH.
herbarium. A collection of dried plants (or lichens).
Heterococcus. A genus of yellow-green algae (Xanthophyceae) found in some species of Verrucaria as
single-celled or short filamentous units, but which in culture becomes filamentous and
branched (as in fig. 1i).
heterocyst. A specialized, thick-walled, colorless cell in certain cyanobacteria such as Nostoc, and the
site of most nitrogen fixation (fig. 1d).
holdfast. The relatively thick and, in many cases, only attachment point of some lichens, especially
Usnea and umbilicate lichens such as Umbilicaria.
hymenium. The spore-bearing layer of an ascoma, consisting of asci and paraphyses, paraphysoides, or
pseudoparaphyses (fig. 13).
hypha (-ae). The filamentous elements of a fungus, often modified and resembling round or angular
cells (fig. 5).
hyphophore. A stalked, often umbrella-shaped structure bearing thread-shaped conidia in certain
lichens, especially those growing on leaves. See Gomphillus.
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hypothallus. A specialized tissue developing as a basal layer on certain foliose and squamulose lichens
(e.g., Anzia, Coccocarpia; fig. 6g); sometimes considered synonymous with prothallus, but
not in this book.
hypothecium. The tissue just below the hymenium (and subhymenium) but above the exciple (fig. 13);
often with a distinctive color or texture as in Buellia, but sometimes merging with the exciple
as in Biatora vernalis.
IKI. A 1.5 per cent solution of iodine in 10 per cent potassium iodide; Lugol's solution (Chapter 13,
LNA). Lower percentages of IKI are sometimes used for special types of staining procedures.
imbricate. Overlapping in a shingle-like fashion; usually pertaining to scales or squamules, as in
Psora himalayana.
inflated. Swollen and hollow, like the lobes of Hypogymnia enteromorpha..
involucrellum. The covering or cap external to the excipulum and present on many perithecia; usually
black and carbonaceous (fig. 17).
isidium (-dia). A minute thalline outgrowth that is corticate and contains photobiont cells. Isidia are
easily detached from the thallus and serve as a vegetative reproductive units (fig. 19d; plate
18).
isotomic. Dividing in regular dichotomies so that a main stem is not easily distinguishable, as in
Cladina stellaris.
K. A 10 per cent solution of potassium hydroxide (KOH) used in various microscopic preparations, or
in spot tests for revealing certain lichen substances (see table 2, pages 106-107 in Lichens of
North America); can be substituted with household lye (sodium hydroxide, NaOH), 10 pellets
per 20-30 ml (1-1.5 ounces) of water.
KC. A spot test for revealing certain lichen substances, performed by wetting the tested area with K
followed by the application of C (see table 2, pages 106-107 in Lichens of North America).
K/I. An analytical staining procedure for the hymenium or asci performed by pretreating the tissue with
K, removing the K by replacing it with water, and then replacing the water with 1.5 per cent
IKI.
labriform. (a) Lip shaped. (b) Pertaining to soralia, generally formed by an upturned thallus margin or a
bursting hollow thallus lobe, sorediate on the lower or inside (i.e., exposed) surface, as in
Hypogymnia physodes (fig. 20e).
laminal. On the upper surface of a thallus (laminal soralium: fig. 20a).
lax. Loose, not compact; usually referring to the medulla.
lecanorine. Pertaining to an apothecium having a distinct thallus-like margin containing a photobiont,
as in the genus Lecanora (figs. 12b, 13c).
lecideine. Pertaining to an apothecium with no photobiont cells in the margin, and in which the exciple
is at least partially carbonized forming a black apothecial margin, as in the genus Lecidea
(figs. 12a, 13a); see also biatorine.
leprose. Composed entirely of soredia, in reference to a thallus surface or the thallus itself (fig. 2a;
plate 24); as in Lepraria.
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lichen. An association of a fungus and a photosynthetic symbiont (photobiont) resulting in a stable
vegetative body with a specific structure, in which the fungus encloses the phytobiont.
lichenicolous. Growing on or in a lichen, pertaining to fungi and lichens at various levels of parasitism
with respect to the host lichen (plates 4, 5).
lichenized. Pertaining to a fungus, alga, or cyanobacterium living within a lichen association.
lichenometry. A method of dating surfaces (usually rocks) using lichen growth rate and size. The size
of the oldest lichen on a particular surface allows us to estimate how long that surface has
been available for colonization (see fig. 24; plates 26A,B).
lignicolous. Growing on bare wood (lignum), as on a log or a wooden fence (plate 28).
lirella (-ae). An elongated, sometimes branched apothecium, as in Graphis (fig. 12e).
lobe. A rounded or somewhat elongated division or projection of a thallus margin; measured at its
widest point (fig. 3).
lobule. A small, often scale-like lobe growing from a foliose thallus either along its margin or from the
surface, sometimes also appearing along apothecial margins, generally of the same color and
character as the parent thallus (fig. 19f; plate 19). Lobules that are constricted at the base and
function as propagules are often called phyllidia.
lobulate. Having many lobules.
locule. The cell cavity in an ascospore; the locule sometimes developing a distinctive shape due to
unevenly thickened spore walls (figs. 15c, d, i, j, l).
macrolichen. A foliose or fruticose lichen.
maculate. Spotted or blotched by maculae, which are pale round or reticulate areas caused by gaps in
the photobiont layer below the cortex (fig. 4; plate 13).
marginal. Along the thallus margins (fig. 20b: marginal soralia).
mazaedium (-dia). A dry, powdery mass of ascospores and paraphyses formed by the disintegration of
the asci in the ascoma of lichens in the order Caliciales, such as Chaenotheca and Cyphelium
(fig. 12f).
mealy. Coarsely granular, like corn meal.
meiosis. A cell division in which the doubled chromosome number of a parent nucleus (the result of a
sexual union) is reduced to a single set in each daughter nucleus, while providing an
opportunity for the separating chromosomes to exchange parts, thereby increasing potential
genetic diversity in the progeny.
medulla. The internal layer in a thallus or lecanorine apothecium, generally composed of loosely
packed fungal hyphae (figs. 4, 13).
microlichen. A crustose or squamulose lichen. Dwarf fruticose lichens such as Polychidium and
Ephebe are often considered to be microlichens.
morphogenesis. A change in form or shape over time, usually triggered by some external or internal
event.
morphology. Physical characteristics, including external shape and internal anatomy.
muriform. Spores divided into several cells by both longitudinal walls and crosswalls; the cells
therefore look like stones or bricks in a wall (figs. 15g,h).
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muscicolous. Growing over bryophytes, especially moss (for example, like Biatora vernalis).
mutualism. A type of symbiosis in which both components benefit from the association.
mycobiont. The fungal symbiont or partner in a lichen.
Myrmecia. A genus of green algae with round to slightly pear-shaped single cells containing a flat or
(in lichen thalli) strongly folded and twisted chloroplast. Unlike Trebouxia, which it closely
resembles, the chloroplast has no central pyrenoid.
nitrogen fixation. The chemical transformation of atmospheric nitrogen, which cannot be used by
plants, into substances such as nitrates and ammonium compounds that can be used by plants.
nonamyloid. Not turning blue with iodine (IKI), whether or not the tissue has been pretreated with K.
(See also hemiamyloid.)
Nostoc. A genus of blue-green algae found in many lichens; producing bead-like chains or filaments of
cells including thick-walled, colorless heterocysts (fig. 1d); when lichenized, the filaments
may be few-celled.
nucleus (-clei). A spherical structure within the cells of fungi, protoctists, and plants, that contains
genetic material, i.e., the chromosomes and genes.
oceanic. Pertaining to a climate characterized by mild wet winters, cool moist summers, and frequent
fogs.
ocular chamber. A dome-shaped indentation of the lower edge of the tholus in certain types of ascus
tips such as those of Lecanora-type asci (fig. 14e).
orchil. A reddish purple dye derived by "fermentation" methods from lichens such as Roccella (plate
69, LNA).
ostiole. The small, usually round, apical pore in various types of ascomata, especially perithecia,
through which the ascospores escape (fig. 17).
papillae. Small rounded or cylindrical bumps like "goose pimples," on the cortex of certain lichens,
especially Usnea (fig. 37a). Papillae do not contain medullary tissue.
paraphysis (paraphyses). A sterile fungal filament, sometimes branched, attached at the base and free at
the summit, associated with asci in the hymenium (figs. 13, 16).
paraphysoids. Sterile hymenial tissue between the asci, usually abundantly branched with frequent
anastomoses (figs. 13f, 16c).
PD. Para-phenylenediamine, a reagent used in spot tests for revealing certain lichen substances (see
table 2, pages 106-107 in Lichens of North America).
peltate. Attached at the center of the lower surface; umbrella-like.
pendent. Hanging straight down and usually soft and pliable (fig. 2j; Usnea longissima); grades into
almost pendent forms that are bushy and relatively stiff at the base but pendent over most of
their length, as in Usnea diplotypus.
periphyses. Short, hairlike hyphae that sometimes line the inner walls of a perithecium (or an
apothecium opening by a pore) near the ostiole (fig. 17a).
perispore. See halo.
perithecium (-cia). A flask-shaped ascoma opening by a pore at the summit (fig. 17); may be
prominent, but is more often partially or completely embedded in the thallus tissue.
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photobiont. The photosynthetic component (symbiont) in a lichen thallus, either algae in the strict sense
(e.g., green algae) or cyanobacteria (blue-green algae) (fig. 1).
phyllidium (-dia). See lobule.
phyllocladium (-dia). A minute cylindrical, lobed or scalelike outgrowth on the stalks and branches of
species of Stereocaulon (fig. 8g).
podetium (-tia). A stalk formed by a vertical extension of lower apothecial tissues (usually the
hypothecium and stipe). The stalk can become secondarily invested with an algal layer and
cortex (as in Cladonia) or can remain almost free of vegetative tissue (as in Dibaeis); the
fertile tissue or apothecia can be present (as in the red-fruited Cladonia bellidiflora) or absent
(as in C. coniocraea); podetia can be either short and unbranched (fig. 2h) or quite tall and
highly branched (fig. 2i).
polarilocular. Pertaining to spores having two cell cavities separated by a relatively thick septum
through which a narrow canal passes (fig. 15d); characteristic of Caloplaca and related
genera.
polyphyllous. In species of Umbilicaria or Dermatocarpon: composed of numerous, crowded lobes
rather than a single, peltate or umbilicate thallus (see U. polyphylla).
U
primary squamules. Small, scale-like lobes forming the basal or primary thallus of Cladonia species.
primary thallus. A squamulose or crustose thallus from which fruticose stalks or podetia arise as
secondary components. Examples are found in Cladonia, Pilophorus, and Baeomyces.
propagule. A reproductive unit, either sexual (such as an ascospore) or vegetative (like a soredium or
isidium).
prosoplechtenchyma. Fungal tissue consisting of coalesced, rather elongate hyphal cells often with
thick, gelatinized walls (figs. 5e, g). (Compare with pseudoparenchyma.)
prothallus. The purely fungal, white or darkly pigmented border of many crustose thalli, often visible
as a fungal mat between the areoles or granules (fig. 2b; also see Lecanora thysanophora and
Rhizocarpon geographicum).
pruina. Powdery, frostlike deposit (usually white or gray, rarely yellow or reddish), typically composed
of calcium oxalate or pigment crystals, dead cortical tissue, or some mixture of them; often
occurs on a thallus (plate 11) or apothecial surface (see Lecanora rupicola).
pruinose. Having a frosted appearance due to a deposit of pruina.
pseudocyphella (-ae). A tiny white dot or pore caused by a break in the cortex and the extension of
medullary hyphae to the surface (figs. 4 and 11a,b).
pseudoparaphyses. The remains of fungal tissue found between the asci in ascolocular ascomata such
as those in Arthonia; generally highly branched and anastomosing (fig. 16c).
pseudoparenchyma. Fungal tissue that appears cellular in section due to short, rounded to almost square
cells of highly branched, irregularly oriented fungal filaments (figs. 5d, 5f); sometimes called
paraplectenchyma.
pustule. A more or less hollow wart or verruca, small and knobby as in Loxospora pustulata, or broad
and blisterlike as in Lasallia.
pustulate. Having many pustules.
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pycnidium (-dia). A small globular or flask-shaped body in which conidia are formed, embedded in a
thallus or entirely superficial, often closely resembling a perithecium (fig. 18).
pyrenocarp (-ous). A lichen in which the fruiting bodies are perithecia, e.g., Verrucaria or Pyrenula.
pyrenoid. A round, conspicuous body associated with the production and storage of starch and found
within or attached to the chloroplast of some green algae. In Trebouxia, it is in the center of
the chloroplast and is very conspicuous (often mistaken for the nucleus, which is almost
invisible without staining and is in the colorless cytoplasm).
refugium. A geographic region that escaped destruction from continental glaciers or inland seas and
thereby provided habitat for certain organisms during the ice ages or ancient periods of marine
incursions; examples are the Yukon-Alaska corridor and the southern Appalachian Mountains.
relict. A remnant population far removed or disjunct from the main population center, usually surviving
in a refugium.
reticulate. Netlike and interconnected, like the branches of Ramalina menziesii.
rhizine. A purely hyphal extension of the lower cortex that generally serves to attach a foliose thallus to
its substrate; of various lengths, thicknesses, colors, and degrees of branching (figs. 4 and 6).
rimose. Having a minutely cracked appearance (fig. 2c).
saprophyte. An organism that lives on decaying organic matter.
saxicolous. Growing on rock, stone, pebbles, concrete, or brick.
scabrose. Having a minutely roughened, almost crusty surface, generally caused by an accumulation of
dead cortical material (see Peltigera scabrosa); often intergrades with pruina, which is more
powdery than crusty.
schizidium. A lichen fragment consisting of the upper layers of a thallus (with the cortex and
photobiont) and serving as a vegetative reproductive unit; formed by a scaling off of the
thallus surface (as in Fulgensia) or a breakdown of pustules (as in some species of
Hypotrachyna or Neofuscelia) (fig. 19g).
Scytonema. A genus of cyanobacteria with a filamentous thallus (fig. 1e) that branches by breaking
through its gelatinous sheath (false branching).
sensu lato (s. lat.) Meaning, "in a broad sense;" usually used in reference to species or genera that are
somewhat heterogeneous and may include other taxa.
septum (-ta). A crosswall in a fungal hypha or spore (figs. 5, 15b-j).
sessile. Sitting on the surface, without a stalk of any kind, as with the apothecia of Lecanora pacifica.
siliceous. Pertaining to a rock or soil rich in silica and lacking calcium; examples are granite and
gneiss, and quartz sand.
soralium (-lia). An area of a thallus in which the cortex has broken down or cracked and soredia are
produced; can be in many forms (fig. 20; plate 17); sometimes contains isidia as well as
soredia, as in Lobaria pulmonaria and Usnea diplotypus.
soredium (-dia). A vegetative propagule of a lichen consisting of a few algal cells entwined and
surrounded by fungal filaments, and without a cortex; generally produced in localized masses
called soralia, or covering large diffuse areas on a thallus (figs. 19h-j).
sp. Abbreviation of "species," generally used where the species is unknown or unspecified.
220
species. The basic evolutionary unit of an organism; named with two words, the first being the genus to
which the species belongs, and the second the species' own name or "epithet." With respect to
lichens, the name of a species refers to its fungal component. Lichen species are defined by
discontinuities in various morphological, chemical, ecological, and geographic characteristics,
and more and more frequently in recent years, by analysis of the actual genetic material
(DNA) of the fungal component.
spore. A single- or multicelled reproductive body capable of giving rise to a new organism; as used
here, refers specifically to an ascospore (fig. 15).
spp. Several unspecified species.
squamule. A small, scalelike lobe or areole, lifting from the surface at least at the edges (as in
Acarospora fuscata or Peltula species), and sometimes strongly ascending and almost foliose
(as in some species of Cladonia).
squamulose. Composed of, or characterized by having squamules (fig. 2e).
squarrose. With short, stiff, perpendicular branches; having the general appearance of a bottle-brush, as
in certain types of rhizines (figs. 6d-e).
ssp. Subspecies, a formal subdivision of a species, used for important morphologically, chemically, or
ecologically distinct segregates, usually somewhat geographically isolated.
stereome. A tough, cartilagenous cylinder forming the supporting tissue for species of Cladonia and
Cladina (fig. 8f).
Stichococcus. A small, unicellular green alga having short, cylindrical (rod-shaped) cells (figure 84d).
Stigonema. A genus of filamentous cyanobacteria having "true branching" resulting from perpendicular
division of cells within the filament (fig. 1a). (Compare with Scytonema, fig. 1e). Lichens with
Stigonema (e.g., Ephebe) do not look very different from the free-living cyanobacteria.
stratified. Layered; in reference to lichen thalli having distinguishable layers of tissue including a
cortex, photobiont layer, medulla, and often, a lower cortex (fig. 4).
striate. With fine longitudinal lines, ridges, or grooves.
stroma (-ata). A wart-like mass of vegetative fungal tissue supporting fruiting bodies of various kinds,
as in Trypethelium species and Glyphis cicatricosa. The stromata seen in lichens almost
invariably contain some material from the substrate (usually bark) as well as fungal tissue and
are more precisely termed pseudostromata.
sub- (a) Partially. (b) Incompletely. (c) Approaching. (d) Under; as in subfoliose (not quite foliose),
submarginal (close to the margin), subpendent (almost pendent), or subhymenium (the layer
often distinguishable just below the hymenium and above the hypothecium).
substrate. The surface upon which a lichen grows; a nutritional relationship is not implied and rarely
occurs in lichens.
superficial. Used with reference to apothecia that sit on the surface of the thallus (i.e., sessile, not
immersed or stalked).
symbiosis. A long-term, usually physical association between at least two dissimilar organisms.
Mutualism and parasitism are types of symbiosis in which one or both organisms are changed
221
in some way, either beneficially or detrimentally. In commensalism, neither of the associated
organisms is affected.
taxon (-a). A unit in a classification scheme; most commonly used with reference to a genus, species,
or subdivision of a species (subspecies, variety, or form).
taxonomy. The study of taxa, especially with respect to their identification and classification, and the
correct application of their names.
terete. Round in cross section (cylindrical) like the branches of Bryoria species.
terricolous. Growing on soil, sand, or peat.
thalline. Pertaining to the lichen thallus; similar to the thallus in appearance or structure.
thalline margin. See amphithecium.
thallus. In lichens, the vegetative body consisting of both algal and fungal components (fig. 2).
thin-layer chromatography. A method used for the identification of chemical compounds, specifically
lichen substances (plate 89, LNA).
tier. A platform-like expansion or flat cup on the podetia of some species of Cladonia (e.g.,
Cladonia cervicornis ssp. verticillata), often proliferating from the center or margins with one
or more new branches.
tholus. The thickened tip of an ascus, frequently staining in iodine (IKI or K/I) in various ways (fig.
14).
TLC. Thin-layer chromatography.
tomentose. Having tomentum; with a downy or woolly appearance.
tomentum. A covering of fine hair or fuzz usually caused by a superficial growth of colorless hyphae
(plate 12).
Trebouxia. A genus of single-celled green algae with one distinctive, disk-shaped chloroplast almost
filling the cell. The chloroplast has a lobed or scalloped margin and a single round to oval
pyrenoid at the center. Trebouxia is the most common green photobiont in lichens. When
followed by a query (?) in the text, it is used in the broad sense, including Pseudotrebouxia
(fig. 1).
Trentepohlia. A genus of filamentous green algae found in many crustose lichens (fig 1j); when
lichenized, the alga often produces very short filaments or is single-celled. The orange-red
pigmented globules, common in the cells of unlichenized individuals, are infrequent or absent
in lichenized individuals.
tubercle. A wartlike protuberance that contains some medullary tissue; characteristic of Usnea ceratina
and Ramalina paludosa.
tuberculate. a) Having the general form of a tubercle, usually in reference to small, round soralia (as in
Bryoria fuscescens); b) having many tubercles, as in some species of Usnea.
umbilicate. Attached by a single, central holdfast (an umbilicus) on the lower surface of the thallus (fig.
2g).
umbilicus. A short, thick, purely fungal, central attachment organ present on certain foliose lichens
such as Umbilicaria and Dermatocarpon.
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umbo. The central bump (or depression) on the upper surface of an umbilicate lichen, corresponding to
the position of the umbilicus.
UV. Ultraviolet light. A number of lichen substances fluoresce and therefore can be detected in long
wave UV light (365 nm) (plate 90); short wave UV (254 nm) is used in thin layer
chromatography for analyzing plates made with gels containing certain fluorescent dyes.
vagrant. Growing unattached to rock or soil and therefore able to roll freely over the ground (plate 34).
var. Variety; a formal subdivision of a species or subspecies, usually used for recurring, genetically
based variants in morphology, chemistry, or habitat.
vein. In lichens, broad or narrow ridges or thickenings, often pigmented, on the lower surface of some
lichens such as Peltigera (e.g., P. kristinsonii), but not functioning as conducting tissue.
verruca (-cae). A conspicuous, wartlike, thalline protuberance (e.g., Pertusaria plittiana).
verrucose. With a rough, warty surface (e.g., Ochrolechia oregonensis).
verruculose. Minutely verrucose, like the thallus of Lecanora circumborealis.
wood. Lignum; trunks, logs, and stumps having no bark.
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Mari Aguilar