Alectoria and allied genera in North America
Irwin M. Brodo and
OPERA
BOTANICA
Scope. Opera Botanica is an international mono-
graph series, issued as a supplement to Bota-
niska Notiser. It is open for members of the
Lund Botanical Society for the publication of
papers on plant taxonomy and morphology and
related subjects, such as phytogeography, cyto-
logy, anatomy, embryology and phytochemistry
in so far as they have a bearing on taxonomy.
The series is primarily intended for comprehen-
sive articles exceeding 32 printed pages. The
numbers are issued at irregular intervals.
Owner. The Lund Botanical Society.
Director. Ingemar Björkqvist, Lund.
Editor. Thomas Karlsson. Assistant Editor:
Anna Weimarck. Manuscripts should be
addressed to: The Editor, Department of Plant
Taxonomy, Ö. Vallgatan 20, S-223 61 Lund,
Sweden.
Editorial board. Stig Falk and Gunnar Harling,
Göteborg, Olov Hedberg and Nils Lundqvist,
Uppsala, Bengt Pettersson, Umeâ, Rolf Santes-
son, Stockholm, Anders Kylin, Sven Snogerup
and Gunnar Weimarck, Lund.
Business address. Requests on prices, discounts,
subscriptions as well as orders for single
numbers should be sent to the Editorial Service,
Swedish Natural Science Research Council,
P.O. Box 23136, S- 1 04 35 Stockholm, Sweden.
Reprints. Authors will receive 50 copies free of
charge. A limited number of additional copies for
non-commercial circulation may be purchased
at a 50% discount.
Copyright. © 1977, The Lund Botanical Society.
Distribution. The Editorial Service of the
Swedish Natural Science Research Council.
Botaniska Notiser
Botaniska Notiser aims at the speedy publica-
tion of papers of high quality that are of
distinctly international interest. It covers the
same general field as Opera Botanica, but
publishes short papers only. The journal appears
quarterly, each volume of four issues comprising
about 480 pages. Requests on subscriptions,
prices and discounts should be sent to the
Swedish Natural Science Research Council. The
staff, editorial board and addresses are as for
Opera Botanica.
Svensk Botanisk Tidskrift
Svensk Botanisk Tidskrift is a journal with
articles in Swedish, but internationally
comprehensible by abstracts and summaries in
English. This journal publishes papers of
national interest on phytotaxonomy, phytogeo-
graphy, phytosociology, and floristic notes as
well as review articles. A yearly bibliography of
Swedish botanical literature is also published
and reprints are available. The journal appears
quarterly, each volume of four issues comprising
about 320 pages. Requests on subscriptions,
prices and discounts should be sent to the
Swedish Natural Science Research Council. The
staff, editorial board and addresses are as for
Opera Botanica.
Flora of Ecuador
Flora of Ecuador (excluding the Galapagos
Islands) is published in parts which are issued at
irregular intervals. Subscriptions and orders for
single parts should be sent to the Swedish
Natural Science Research Council (address, see
above).
International editorial board: Misael Acosta
Soil's, Quito; Angel L. Cabrera, La Plata;
Lincoln Constance, Berkeley; Richard S.
Cowan, Washington; Lauritz Holm-Nielsen,
Aarhus; Alicia Lourteig, Paris; Bassett Maguire,
New York; Peter H. Raven, Saint Louis;
Rolf Santesson, Stockholm; Richard E.
Schultes, Cambridge, Mass, and Frans A.
Stafleu, Utrecht.
Editors: Gunnar Harling, Department of
Systematic Botany, University of Göteborg,
Carl Skottsbergs Gata 22, S-413 19 Göteborg,
Sweden ; Benkt Sparre, Curator of the Regnellian
Herbarium, Section for Botany, Riksmuseum.
S- 104 05 Stockholm 50, Sweden and Uno Elias-
son (assistant Editor), Department of Systematic
Botany , Göteborg.
Opera Botanica 42
Alectoria and allied genera in North America
Irwin M. Brodo and David L. Hawksworth
Publications in Botany, No. 6
National Museum of Natural Sciences
National Museums of Canada
ISSN 0078-5237
ISBN 91-546-0211-4
Berlingska Boktryckeriet, Lund 1977
Frontispiece. Bryoria nadvornikianu draping the branches of Picea mariana in Parc des Laurentides, north of
Quebec City, Quebec.
I
Alectoria and allied genera in North America
Irwin M. Brodo and David L. Hawksworth
Brodo, I. M. & Hawksworth, D. L. 1977 02 25: Alectoria and allied genera in North
America. Opera Bot. 42: 1—164. Stockholm. ISSN 0078-5237. ISBN 91-546-0211-4.
In the present monographic study of the North American taxa generally referred to the
genus Alec toria, four genera and 38 species were recognized: Alectoria Ach. (8 species),
Bryoria nom. nov. (27), Pseudephebe Choisy (2), and Sukaria Bystr. (1). Nine new
species are described: Alectoria imshaugii, A. mexicana, Bryoria cariottae, B. friabilis,
B. pikei, B. pseudocapillaris, B. salazinica. B. spiralifera, and Sulcaria hadia. One
species name is validated and transferred to Bryoria: B. cervinula. Combinations
transfering 26 North American and 29 extra-North American taxa into the accepted genera
are made. Some aspects of the cortical structure are useful in characterizing the genera
and the sections within Bryoria. Secondary chemical products have been studied in
all taxa. Olivetoric and gyrophoric acid were previously unknown for this group.
Salazinic acid was definitely established as occurring in Bryoria. Atranorin and
chloroatranorin are absent in Alectoria s. str. but present in Bryoria. Thamnolic acid
appears to be absent in Bryoria. but fairly common in Alectoria s. str. The group as a
whole is northern in distribution in North America extending southward in the cordillera.
Only Sulcaria appears to be essentially temperate. The corticolous species are mainly
confined to trees with acid bark, and those on rock and soil seem to be more character-
istic of acid, non-calcareous substrates.
Irwin M. Brodo, National Museum of Natural Sciences. National Museums of Canada.
Ottawa, Ontario KIA OM8, Canada.
David L. Hawksworth, Commonwealth Mycological Institute, Kew, Surrey TW9 3AF,
England.
Acknowledgements
Introduction
Anatomy and morphology
Chemistry
T axonomic concepts
Ecology
Distribution
Importance to man and nature
Taxonomic survey
Key to the genera of Alectorioid lichens .
Check-list of the North American species
Key to the North American species
Alectoria Ach
Bryoria Brodo & D. Hawksw
4 sect. Bryoria 79
5 sect. Divaricatae (DR.) Brodo & D. Hawksw. 98
6 sect. Implexae (Gyeln.) Brodo & D. Hawksw. 1 14
21 sect. Subdivergentes (Mot.) Brodo & D.
30 Hawksw 132
35 sect. Torluosae (Bystr.) Brodo & D. Hawksw. 135
39 Pseudephebe Choisy 140
44 Sukaria Bystr 146
48 Excluded taxa 149
48 New combinations for non-North American and
48 some infraspecific taxa 155
49 References 157
56 Index 162
78
4 Alectoria in North America
OPERA BOT. 42 (1977)
Acknowledgements
The production of a major taxonomic revision of a
group of lichens by aulhors on different sides of the
Atlantic presented us with a number of problems.
The file of correspondence between the junior and
senior authors stands some 5 cm high. The authors
have been able to work together jointly on this
project in May 1970 (in Kew). March and April 1973
(in Ottawa), and September 1973 (in Kew); most final
points in the manuscript were discussed together at
the XII International Botanical Congress in Leningrad
in July 1975. We are indebted to the Commonwealth
Agricultural Bureaux, the National Museums of
Canada, and the Royal Society of London for making
these meetings possible.
This study, made over a period of eight years,
has profited trom the help of many individuals and
institutions. We would particularly like to thank the
curators of all the herbaria from which we borrowed
specimens, especially as w*e often had to keep the
material longer than the allotted time, and frequently
reborrowed certain specimens.
Certain individuals merit special thanks. We bene-
fited from the experience and comments of Hildur
Krog and Teuvo Ahti from the beginning. Their deep
interest in our work and invaluable field observations
of many species are most warmly appreciated.
Theodore Esslinger generously made his extensive
collections of A. imshaugii available to us, together
with detailed comments on the chemistry of his
material. Gary Schroeder also sent us many specimens
of that species, including a large collection that we
have designated as the type. E. D. Greenwood made
special trips inlo the mountains of Oaxaca. Mexico,
to search out and collect material of A. mexicana,
A. lata, and Oropogon spp. Per Magnus Jorgensen
gave us many valuable comments on species of the
section Divaricatae ot Bryoria. We also profited
trom discussions of certain taxonomic and nomen-
clatural problems with Peter James and Rolf
Santesson.
We are most grateful for the time spent by John
Thomson in testing our keys and compiling comments
and corrections on them. We should especially like
to thank Henry Imshaug for locating the type
specimen of Setaria trichodes at the Paris Museum.
Ove A I inborn generously gave of his time in the
revision of all the Latin diagnoses, and for this
we are very grateful. Finally, we would like to thank
Pak Yau Wong at the National Herbarium of Canada
for his careful attention to the chromatographic runs,
for testing our keys, and for painstakingly typing
thousands of specimen citation cards. The drawings
were made by C. H. Douglas of the National
Museum's Zoology Division.
All the maps were prepared using the Goode Base
Map Series, No. 102. copyrighted by the University
of Chicago and used with their permission.
Introduction
The lichen genus Alectoria in North America,
although containing some conspicuous and
frequently collected species, has persistently
proved to be a source of confusion and
frustration among both amateur and profes-
sional lichenologists.
The first attempt to provide a monographic
treatment of the North American species of
this genus was that of Howe (1911), who
accepted ten species. Howe's work was not
sufficiently critical, however, and he misapplied
several names. A number of new taxa were
described from North America by Gyelnik in
the 1930s (see Sjödin 1954), but most of these
have been poorly understood by subsequent
workers. More recently Motyka (1960) de-
scribed three new species from North American
material, and later (1964) published a revision
of the thirty species he considered to occur
in North America. Unfortunately, his paper
was based on a study of very few North
American specimens, and suffered from
numerous errors and omissions. Attempts to
follow Motyka’s treatment have led to some
confusions in the application of many names,
and it became apparent that a thorough re-
examination of the North American Alectoriae,
in the broadest sense, was imperative.
The present paper provides a revision of the
North American species (and those of Mexico)
of Alectoria and its segregate genera Bryoria,
Sulcaria, and Pseudephebe . Oropogon Th. Fr.
is an essentially Asiatic genus (Sato 1967)
also occurring in the Central and South Amer-
ican highlands. We have seen no North Ameri-
can specimens, and thus it has been excluded
from the present study. It is much in need of
a generic revision.
The introductory sections discuss the ana-
tomical, morphological and chemical charac-
ters of these genera only so far as they affect
the taxa treated here. This paper constitutes
“Regional Studies in Alectoria (Lichenes) IV“
of the junior author; part III appeared in
Annales Botanici Fennici 1 1: 189-196 (1974).
Material
The information presented here is based on a study
of nearly 9,000 specimens from North America.
Besides studying the extensive material housed at
the National Herbarium of Canada (CANL), we
borrowed and revised material from the following
herbaria (abbreviations follow Holmgren & Keuken
1974): ALA. AR, BM. COLO, DUKE, F. FH. H.
LAM. MSC, NYBG, O, QFA. QUE. S. TRTC, UAC.
UPS, US. WIS. In addition, the following private
herbaria were consulted: DEGEL1US, ESSL1NGER,
PIKE. SCHROEDER, THOMSON. Type specimens
and related materials were also examined from BP,
FI. LBL, LE. M, MW. ORE. P, PC, and ZT.
Some collections were made in Oaxaca, Mexico, by
Mr E. D. Greenwood especially for this study.
A number of special field studies were made by
the senior author to investigate the habitat ecology
of some of the more puzzling or rare taxa, and one
excursion was taken by the two authors together.
Among the areas visited were the California and
Oregon coasts; the Corvallis, Oregon, region:
coastal British Columbia; some of the western
mountains; and boreal forest segments near
Moosonee, Ontario, and in Parc de la Verendrye,
the Gaspé Peninsula, and Parc des Laurentides,
all in Quebec.
Anatomy and morphology
Habit
The terms caespitose, subpendent, pendent,
and decumbent are frequently used to describe
the various growth forms of species of Alectoria
s. lat. and they are used in the keys that fol-
low. Our usage of these terms is as follows:
caespitose: forming shrubby tufts, generally
producing a thallus not more than twice as long
as it is broad; a number of thick branches
generally arise together at the base of the
plant. Subpendent: producing many thick
branches at the base of the thallus, but
acquiring a more slender and pendulose appear-
ance at the tip; generally several times as
long as it is broad. Pendent: hanging vertically
almost from the base; becoming very elongate.
Decumbent: growing along the surface of the
substrate and forming low cushions or rosettes.
Erect: becoming more-or-less arborescent,
usually with a main “stem” or branch with
subordinate lateral branches arising from it.
Most species show only one type of growth
form: caespitose: Bryoriu furcellata, B.
abbreviata, Alectoria imshaugii; subpendent:
Bryoria glabra, B. tenuis, B. nadvornikiana;
pendent. B. fremontii, B. trichodes subsp.
trichodes, B. capillaris. Others can vary some-
what, such as Alectoria sannentosa subsp.
sarmentosa which can be subpendent when
young, or Bryoria simplicior which can be
caespitose or subpendent.
Branching
Terminology of branching basically follows
Hawksworth (1972). Various types and some
examples are illustrated in Fig. I. Among the
most clearly isotomic dichotomous types in
North America is Bryoria glabra with even,
regular, and broad dichotomies occurring
over half the length of the thallus. Bryoria
lanestris has clearly anisotomic dichotomies
over almost its entire length; B. trichodes
varies from isotomic in subsp. americana to
anisotomic in subsp. trichodes. In B.fuscescens
considerable variation in types of branching
occurs even in the same thallus. The erect
species are all clearly anisotomic.
Isidia
As pointed out by Hawksworth (1972), true
isidia do not occur in the genus Alectoria s. lat.
Isidioid branches and spinules do frequently
appear, usually associated with soralia (as in
Bryoria furcellata, Fig. 2 A) or pseudocyphellae
(as in Alectoria imshaugii (Fig. 2 B) or, less
frequently, in A. sarmentosa and A. vancou-
verensis). Spinulose branches characterize such
species as Bryoria nadvornikiana (Fig. 3) and
some forms of B. nitidula.
Pike 843 B (CANL) - C-V"^ nfsotomic dfchnf ^ ** ' ~ A ’ B „ lsotomic dichotomous: Bryoria glabra. Washington,
*■ nadvornikiana. Quehec Bi l T ip'cAN, 0 ;"' E n Zd 19753 < CAN ‘-> -D:
Aiectoria nigricans. N. W. T.. Cody & ÄoV lol!o£<£^ " F =
/lv
OPERA BOT. 42 (1977)
Anatomy and morphology 7
8 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 2. Isidioid spinules. - A: Bryoria furcellata. Cummings, Decades No. 53 (CANL). - B: Alectoria imshaugii
(holotype). - Scale 0.5 mm.
Soredia
Soredia always arise within discrete soralia of
one of two types. Fissurai soralia originate as a
longitudinal split in the cortex and are gener-
ally elliptical and flat or concave (Fig. 4 A).
Tuberculate soralia are round or irregular in
outline, conspicuously project above the sur-
face of the filament, and often display a
narrow cortical rim (Fig. 4 B). Bryoria glabra
and B. simplicior have only fissurai soralia;
B. nadvornikiana and B. fuscescens var. posi-
tiva have tuberculate soralia predominating.
The presence of soralia is constant in some
species, but not in others. North American
specimens of B. capillaris and B. implexa do
not have soredia, although European specimens
often do. Soralia range from abundant to
Fig. 3. Spinulose branches. Bryoria nadvornikiana.
Quebec, Guimont 4442 (CANL). Scale 0.5 mm.
absent in B. glabra, although the latter situation
is very rare. Both subspecies of B. trichodes
have sorediate morphotypes. There is a rather
rare, abundantly sorediate morphotype of
Alectoria sarmentosa subsp. sarmentosu with
tuberculate soralia, and rare specimens of A.
fallacina and A . mexicana also show soralia.
The soralia in all four Alectorioid genera
tend to be white, but in Bryoria fremontii
(and in the rare sorediate morphotype of B.
tortuosa), they are bright yellow due to the
presence of concentrations of vulpinic acid.
In Bryoria lanestris and especially B. simplicior,
the soralia are black to greenish flecked,
probably due to the remnants of cortical and
algal tissues on their surface.
Pseudocyphellae
Pseudocyphellae are constant in Alectoria and
Sulcaria, but are very variable in Biyoria. In
the section Divaricatae, pseudocyphellae are
generally dark and very inconspicuous. In
taxa such as B. trichodes subsp. trichodes,
they are white and are often somewhat raised
(Fig. 5 A); in other taxa, they are plane
(Fig. 5 B). Pseudocyphellae are yellow in
B. tortuosa (Fig. 5 C), and are sunken into
spiraled channels in members of the genus
Sulcaria such as S. badia (Fig. 5 D). In
Bryoria pseudocapillaris and B. spiralifera,
the pseudocyphellae are linear, white, and
very long (Fig. 5 E).
OPERA BOT. 42 (1977)
Anatomy and morphology 9
Fig. 4. Soralia. - A: Fissurai soralium. Bryoria glabra. Washington, Pike 843 (CANL). - B: Tuberculate
soralia. Bryoria fuscescens var. positiva. Newfoundland. Ahti 560 (CANL). Scale 0.5 mm.
Contrasting sharply with the flat or chan-
nelled pseudocyphellae described above,
those of the genus Alectoria s. str. are con-
spicuously raised and striking (Fig. 5 F).
Most are somewhat elongate, but in A. falla-
cina most pseudocyphellae are punctiform.
Colour
The use of colour causes particular difficulties
which arise from differences in interpretation
by individual observers. For this reason, we
have endeavoured not to place too much
emphasis on it. However, we found that after
looking at many examples of various species,
their characteristic colour tones began to
become clear, and it seems evident that species
do have characteristic colours, although these
can be subtle and difficult to interpret.
It is most difficult to interpret colour changes
due to the age of a specimen. For one thing,
we have noticed that species which some-
times have a distinctly greenish appearance in
the field or in fresh specimens gradually lose
this tone in the herbarium with the death and
bleaching of the algal layer. This is obviously
most serious with the paler species such as
Bryoria capillaris, B. fuscescens. B. trichodes
subsp. trichodes, and Alectoria sarmentosa.
Species with barbatolic acid or alectorialic
acid (e.g., Bryoria capillaris and Alectoria
nigricans ) gradually become pinkish or even
reddish and may even stain their packets
reddish in time. Bryoria friabilis, normally
very pale olivaceous brown, turns to dark
brown with age. Most other species retain
their colour, however.
There are three basic hues in Bryoria:
(1) the olivaceous or olive-black of species such
as B. glabra, B. carlottae, B. pikei, B.
chalybeiformis, and B. vrangiana; (2) the
‘‘cervine brown” (i.e., deer brown) of B.
trichodes, B. furcellata, and B. simplicior;
and (3) the white to grey-fuscous of B. subcana,
B. capillaris and B. nadvornikiana. One might
call the clear red-brown of abbreviata distinct
as well as the unique chestnut brown of
Sulcaria badia and Bryoria spiralifera, but
it is difficult or impossible to distinguish
them adequately from cervine brown using a
verbal description.
Even the species with usnic acid show some
constant variation in tone, although it is
difficult to describe. Alectoria sarmentosa is
extremely variable in colour, but is generally
a very “yellow” yellow-green (i.e., straw-
coloured) especially in the herbarium. A. falla-
cina has a much darker, greyer tone as does
A. vancouverensis. There is a great deal of
variation in the amount of green or green-
black on A. sarmentosa and A. ochroleuca
10 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 5. Pseudocyphellae. - A: Fissurai, raised. Bryoria trichodes subsp. trichodes. Ontario, Brodo 13737 (CANL).
Scale 0.25 mm. - B: Fissurai, plane. Bryoria carlottae. Holotype. Scale 0.25 mm. -C: Twisted, yellow. Bryoria
tortuosa. Rel. Tuck. No. 2 (CANL). Scale 0.5 mm. -D: Sulcate. Sulcaria badia. Oregon, Hall (FH-Tuck. 504).
Scale 0.4 mm. - E: In long spiral. Bryoria spiralifera. Holotype. Scale 0.5 mm. - F: Raised on ridges. Alectoria
vancouverensis. B. C., Bird 28472 (CANL). Scale 1.0 mm.
OPERA BOT. 42 (1977)
Anatomy and morphology 1 1
apparently depending on the degree of exposure
and moisture-availability.
Unhappily, there are a number of species
which vary a great deal in colour. Bryoria
pseudofuscescens can be pale brown, dark
brown, or greenish-black with every possible
intermediate. The possibility that the norstictic
acid-producing species may actually consist
of several taxa has been raised by Hawksworth
(1972) (see Discussion under B. pseudo-
fuscescens). B. duplexa varies from pale brown
to pale olivaceous, but so few fresh specimens
have been seen, this difference may be due to
ageing. The same is true of B. salazinica
which does seem to have a pale olivaceous
caste when fresh. Alectoria nigricans varies
from pale grey with black tips through yellowish
grey with black tips, to entirely black with
grey basal areas; rare specimens are a shiny
brownish grey and blackened at the tips.
True black thalli are rarely found in any
species, although some specimens of Alectoria
nigricans and Bryoria nitidula are almost
uniformly black. The bicolourous thalli of
B. bicolor and B. tenuis have truely black
main stems with only the secondary or tertiary
branches remaining olivaceous brown. Some
forms of B. trichodes subsp. americana
have extensive blackened areas and in some
cases appear somewhat like B. tenuis. These
blackened areas are a type of fragmentation
area referred to by Hawksworth (1972).
Only one colour is due to something other
than the presence or absence of pigments in
the cortex. In B. tortuosa, a thallus may
acquire a bright emerald green colour by
virtue of an accumulation of vulpinic acid on
the surface of the cortex. Often thalli apparently
collected on the same branch (being in the
same packet) vary from red-brown to bright
emerald green.
Shininess, or dullness, is tempting to use
as a character, but it is usually too variable
to be reliable. Certain species do. however,
show clear tendencies. B. glabra is almost al-
ways very shiny; B. abbreviata is virtually al-
ways dull; B. trichodes can be very shiny or sa-
tiny; B. lanestris tends to be satiny; B. capillaris
is almost always dull; B. pseudofuscescens can
be shiny or dull.
Friability
Friability or brittleness was mentioned by
Hawksworth (1972) as a reliable characteristic
of B. lanestris in Great Britain, and this has
proved to be the case in North America as
well. B. friabilis is also extremely brittle
and the same is true of many other species in
the section Implexae. B. abbreviata and B.
oregana in the section Subdivergentes are also
very easily fragmented. The basis for this
character seems to lie in the cortical structure
(see below).
Ascocarps
Apothecia are very rare in the Alectorioid
genera as a whole and many species are
known only in a sterile condition, their classi-
fication within a certain genus being based
on a combination of vegetative characteristics.
In the case of the North American species,
we have seen apothecia in North American
collections of only 18 of the 38 accepted
species. Even in those species able to produce
apothecia, only a few regularly do so (e.g.,
Bryoria abbreviata. Alectoria lata. Pseudephebe
minuscula, P. puhescens ) and in some very
widely distributed species, apothecia appear to
be produced often in localized populations
(e.g., Bryoria trichodes subsp. americana, B.
fremontii, Alectoria nigricans, A. sarmentosa).
The frequency of ascocarp production in some
species is exceedingly low. In the present study,
for example, out of 247 collections of Bryoria
capillaris and 66 of B. tenuis examined,
apothecia were seen in only one specimen of
each (i.e., 0.40% and 1.52% of the collections
examined, respectively). Even when an apothe-
cium is produced, ascospores are often formed
only sparingly and mature ascospores free
from their asci are often difficult to observe.
Despite the wide distribution and size of
these genera, there has been no detailed study
published of ascocarp structure and ontogeny
in any species. It seems probable that the
asci in all these genera are of the “arrested
bitunicate” type, that is, having inseparable
exo- and endoascus walls, as is the case in
almost all the lichenized ascohymenial fungi
(Letrouit-Galinou 1974, Henssen & Jahns 1973).
12 Alectoria in North America
OPERA BOT. 42 (1977)
Awasthi (1970) considered the presence of
anastomosing paraphysoid tissue (which he
called pseudoparaphyses), particularly at an
early stage of development, to indicate a
possible affinity between the genus Alectoria
s. lat. and the family Roccellaceae Chev.
This family was treated by Henssen & Jahns
(1973) as belonging to the order Arthoniales,
a “Zwischengruppe'' between the ascohymenial
and ascolocular groups. The presence of
anastomosing paraphysoid tissue in ascocarps
which seem to be ascohymenial in all other
respects has long been a source of confusion
to lichen systematists. Henssen & Jahns (1973)
through an investigation of many types of
ascocarps have done much to clear up this
difficulty. They found that one must distin-
guish between paraphysoid tissue (originating
from the generative tissue of many ascohymenial
lichen fungi and often, but not always, gradually
changing into true paraphyses with the matu-
ration of the ascocarp), and pseudoparaphyses
(which have an entirely different origin and
development and are characteristic of the
ascolocular fungi) (see Henssen & Jahns 1973
pp. 90, 116-117).
Our observations of the hymenial tissue of
Alectoria mexicana, A. lata, Bryoria pseudo-
fuscescens, B. furcellata. and B. oregana
have shown that the cells of the paraphyses
of all these species are short and tend to
branch and even anastomose to some extent.
This is particularly evident in A. lata and
A. mexicana and is perhaps true of all members
of the genus Alectoria. The short cells and
branching are well illustrated by Galloe (1950)
for A. sarmentosa and A. ochroleuca. In these
species, one could well speak of paraphysoid
tissue in the sense of Henssen & Jahns (1973)
rather than paraphyses. These species would be
represented by the type of ascocarp illustrated
in Figure 4.19 d of Henssen & Jahns (1973 p
118).
In all species of Alectoria, Bryoria and
Pseudephebe that we have examined, the
excipulum proprium is well developed. Although
Awasthi (1970) regards this as a character
unique to Alectoria s. lat. and Oropogon,
we have seen a distinct excipulum proprium
in the closely related Cornicularia californica
(Tuck.) DR. Gallpe (1947) has illustrated the
same condition in other species of the Parme-
liaceae (e.g., Cetraria nivalis (L.) Ach. and
Evernia prunastri (L.) Ach.).
Thus, there seems to be no reason to regard
any of the Alectorioid genera as a possible
member of the Arthoniales, and indeed, little
reason to isolate these genera from other
closely related genera such as Cornicularia
and Parmelia, as was suggested by Awasthi
(1970).
The numbers of spores per ascus (i.e.,
2-4 vs. 8), their pigmentation when mature
(i.e., brownish vs. hyaline), and their septation
and size, are of importance in the recognition
of the four genera accepted here (see Table 2).
Within the genera, however, ascospore size
appears to be of very limited taxonomic
value for the characterisation of species to
judge from the information so far available.
In view of this and the practical difficulty
of finding mature apothecia in many species,
only scant attention has been paid to these
characters in the present survey.
Anatomy of the vegetative tissues
The anatomy of the vegetative tissues was
studied in some detail because of its potential
value in the delimitation of species. Our
intention was merely to survey the anatomical
features of a majority of the species to see if
trends could be noted and useful characters
discovered. No attempt was made to examine
large numbers of specimens within each species
although this would obviously be desirable,
as pointed out below. Wherever possible,
type specimens were examined.
Our methods for preparing the microtome sections
were almost precisely those employed by Wetmore
(1960 p. 375). Sections were made at 12 pm using
a freezing microtome after the material had been
soaked overnight in an alcohol-detergent solution.
The material was then mounted in lactophenol
and cotton blue. This stain colours the cytoplasm
within the cell lumina making the lumina easy to
see and measure. It also stains the chloroplasts of
the algal cells dark blue. The lactophenol served
as a semi-permanent mounting medium when the
coverslips were sealed with two coats of clear nail
polish, lo see finer characters of the conglutinated
cortical hyphae and the extent of the matrix material,
all the sections were also examined in phase-
contrast illumination.
OPERA BOT. 42 (1977)
Anatomy and morphology 13
* J 1 fg {
V
mmJÊÊ- ç. - 'y .T
BWE* : > Vo : * 1
..V f* »*: »1 * *v>> a/ -1* 5
“a* 4 V* *' ,
r
»* # Ü. >o
J*PtS A ,
V;
_r , ,^. •
4' 7 -»
* £
v y
w
Fig. 6. Cortex of Ramaiina thrausta. On-
tario, Garton 6697 (CANL). Phase photo-
micrographs; scale 20 pm. Photos by Peter
Bowler. - A, B: Longitudinal section. -C:
Cross section.
Cortex
Characters of the cortex have been used by
many authors to distinguish Alectoria s. lat.
from related (or at least superficially similar)
genera, especially Cornicularia, Parmelia, and
Ramaiina. For example, Hawksworth (1969 a)
showed that characteristics of the surface of
the filaments as revealed by scanning electron
microscopy (SEM) for the most part do correlate
well with classical generic concepts.
Studies carried out in connection with the
North American species have shown that there
is much more variation in cortical tissue than
previously suspected. This is especially true
in the extent of septation and emergence
of the superficial layer of cortical hyphae.
However, we must hasten to add that we have
found even the most “Alectoria” - like species
of Cornicularia (e.g., C. pseudosatoana Asah.
and C. californica ) and Ramaiina (i.e., R.
thrausta ) to be quite distinct from any species
in any genus studied here. The cortex of
Cornicularia pseudosatoana has clearly
distinguishable periclinal and netlike layers
of prosoplectenchyma. C. californica almost
develops a paraplectenchymatous outer layer.
Ramaiina thrausta has a fibrous, periclinal
cortex similar in some respects to that of
species of Alectoria but with a noticeable
increase in the branching of the superficial
hyphae (Fig. 6). This, however, is not the
two-layered cortex considered by many authors
to be typical of Ramaiina and illustrated by
Jahns (1974) in his figures 30 and 31. The
surface features of R. thrausta (Hawksworth
14 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 7. Decomposed cortex of Alec-
toria vaneouverensis. California,
Tavares 371 (CANL), cross section.
Scale 20 gm.
1969 a pi. IV c) obviously would not reveal
this.
Basically, the Alectorioid cortex is made up
of periclinal hyphae fused into a chondroid
mass to varying degrees. In some species,
it is composed of fairly distinct interwoven
hyphae and in others, it is clearly proso-
plectenchymatous with only the lumina dis-
tinguishable (a “skleroplektenchym” in the
terminology of Frey 1936). This structure
is particularly well seen in scanning electron
micrographs of cut ends and longitudinal slices
of branches (Hawksworth 1972, Hale 1976,
Jorgensen 1975). The cortex in Alectoria s.
lat. has been called a stereome by a number
of recent authors (Ozenda 1963, Hawksworth
1969 a, 1972) in the context of its being a
supporting tissue. This term has been used by
lichenologists more frequently to refer to
chondroid medullary tissue (as in Cladonia
and Usnea) (e.g., Galloe 1947, 1954) and so
we will simply refer to this tissue in the
Alectorioid genera as the cortex. (It should
be noted that all these tissues are remarkably
analogous in anatomy as well as function.)
Internally, the cortex is generally very distinct
from the medulla. The surface of the cortex
is relatively rough and sometimes ridged (see
Hawksworth 1969 a), but sometimes this
roughness is due to a heavy deposition of
pigmented material, and in other cases, the
superficial hyphae themselves protrude to
varying degrees. A non-cellular epicortex, as
seen in Parmelia and some allied genera
(Hale 1973, 1976), is not found in any of the
Alectoria -like genera. In many species of
Alectoria s. str., especially A. vaneouverensis,
the surface of the cortex seems to decompose
to some extent giving rise to a highly irregular
surface (Fig. 7) (Hawksworth 1969 a pis. 1 f,
111 a).
As mentioned above a number of species
are noticeably more brittle than others. The
basis for this friability apparently lies in the
quantity of matrix material in which the cortical
hyphae are embedded, i.e. , the ratio of lumen
area to wall-matrix area in the cortex. Brittle
species such as Bryoria friahilis and B. implexa
have little matrix material apparent (Fig. 8)
whereas strong, resilient species such as
Alectoria fallacina and Bryoria glabra have
a great deal (Fig. 9). Species such as B.
trichodes and B. fuscescens are more or less
intermediate (Fig. 10).
We have implied above that there are certain
cortical characters which are correlated with
certain genera, and with sections within Bryoria,
and this indeed appears to be true in certain
cases.
In Alectoria s. str., the cortex is generally
thick, averaging about 50-80 /am and commonly
going as high as 120 p m; in the other genera,
the cortex is generally between 30 and 40
pm rarely exceeding 50 pm. In addition, the
cortex in Alectoria is highly cartilaginous with
a great deal of matrix material. As mentioned
OPERA BOT. 42 (1977)
Anatomy and morphology 15
B
0
Fig. 8. Cortex with little matrix. Scale 20 pm. -
A, B: Bryoria friahilis . Oregon, Hawkins (WIS). -
A: Longitudinal section, phase contrast. - B:
Cross section, light microscopy. - C: Bryoria
implexa. Quebec, Macoun? (CANL 16294), longi-
tudinal section, phase contrast.
Rg. 9. Cortex, much matrix. Scale 20 pm. - A, B:
Alectoria fatlacina. Isotype (US). - A: Longitudinal
section, light microscopy. - B: Longitudinal section.
Phase contrast. -C: Bryoria glabra. Isotype (CANL),
cross section, light microscopy.
16 Alectoria in North America
OPERA BOT. 42 (1977)
in A te ™ ediate j matrix- Scale 20 pm. - A, B: Bryoria trichodes subsp. americana. B. C„ Brodo
1 1668 (CAND - A: Longitudinal section, phase contrast. - B: Cross section, light microscopy. -C, D: Bryoria
fuscescens. Isotype (CANL). -C: Longitudinal section, phase contrast. -D: Cross section, light microscopy.
Fig. 11. Cortex with hyphae arranged in fascicles. Scale 20 pm.
(QUE), cross section. - B: Bryoria cervinula. B. C.. Brodo 17769
- A: Bryoria furcellata.
(CANL), cross section.
Quebec, Masson s.n.
OPERA BOT. 42 (1977)
Anatomy and morphology 17
Fig. 12. A-E: Surface cortical features of Bryoria sect. Subdivergentes. Scale 20 - A: Bryoria oregana.
B. C., Brodo 8178 (CANL). surface view of cortex showing jigsaw pattern of superficial cortical cells. Eight,
transmission microscopy. — B— F: Scanning electron micrographs (SEM). — B, C. Biyotia abbreviate i. Alberta,
Bird & Lakusta 16703 (CANL). - B: Negative frame 290 x. -C: Negative frame 580 x. -D, E: Bryoria oregana.
Washington, Imshaug 6237 (CANL). - D: Negative frame 290 x. - E: Negative frame 580 x. - F: Bryoria /re-
monta. Alberta, Bird & Glenn 14214 (CANL), negative frame 580 x . There are no projecting superficial cells of
the cortex in this species which is in the section Tortuosae.
2 - Opera Botanica nr 42
18 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 13. Anatomy of Pseudephehe. - A: Pseudephehe pubescens. B. C., Brodo 14209 (CANL), longitudinal
section of the cortex showing superficial cellular layer. Scale 20 pm. - B-D: Pseudephehe minuscula. Colorado,
Shushan 8323 (CANL), longitudinal sections of the cortex, phase contrast. Scale = 20 pm. - B: “Lower” cortex
of a flattened branch. - C: Upper ’ cortex of a flattened branch. - D: Cortex of a terete branch
above, the surface is also somewhat decom-
posed.
In a number of species of the section
Divaricatae of Bryoria (e.g., B. furcellata,
B. nitidula, B. cervinula ), the cortical hyphae
Fig. 14. Lax, arachnoid medulla in Bryoria implexa.
Quebec, (Macoun?) s.n. (CANL 16294). Longitudinal
section. Scale 60 pm.
are, at least in part, distinct with conglutinate
thick-walled filaments often in fascicle-like
groups (Fig. 11). Most have superficial cells
protruding above the surface giving it a some-
what knobby or ridged appearance, and most
are also encrusted with pigmented material.
Bryoria oregana, B. ahhreviata, and B.
subdivergens have a great many similarities
in habit, colour, spores, etc. and are obviously
closely related. They are treated here as
comprising the section Subdivergentes.
It was very interesting to discover, therefore,
that the cortices of all three are very similar
and very unusual. In these species (and only
these species) the outermost cells of cortical
hyphae become short, irregular, and knobby
forming a jigsaw-like pattern on the thallus
surface (Fig. 12 A). They also extend above
the surface giving the cortex a very rough,
bumpy appearance with SEM or under the
light microscope (Fig. 12 B-E) and a dull or
matt texture to the eye. (Compare this cortical
OPERA BOT. 42 (1977)
Anatomy and morphology 19
Fig. 15. Ornamented medullary hyphae in Alectoria sarmentosa, phase contrast. Scale 20 /im. - A: Subsp.
sarmentosa B. C., Brodo 12792 (CANL). - B: Subsp. vexillifera. Quebec, Guimont 4446 (CANL).
surface with the relatively smooth, encrusted
surface of B. fremontii. Fig. 12 F.) The SEM
photograph of the surface features of Bryoria
divergescens (syn. Alectoria divergescens) in
Hawksworth (1969 a pi. 11 d) as well as the
comments of Dahl (1950) and Awasthi (1970)
lead us to believe that this mainly Himalayan
species may belong in this section as well.
It is not surprising that Pseudephebe minus-
cula and P. pubescens have similar cortices.
The cortical anatomy reflects the general habit
of the plant closely, especially in P. minuscula.
In the more “foliose" parts of the thallus,
there is a net-like prosoplectenchyma (as in
Jahns 1974 fig. 12) forming the "upper" cortex,
and a fibrous, periclinal prosoplectenchyma
forming the "lower” cortex (Fig. 13 A-C).
In the terete branches, the cortex is more
uniformly periclinal, but one can still make
out a distinct shortening of the superficial
cells which results in a kind of false pseudo-
parenchyma at the surface (Fig. 13 D). We
noted no encrusting pigmented material on
the surface; all the pigment (and there is a
great deal) is within the cell walls of the
superficial cells.
Algal layer
As stated by Hawksworth (1972) and illustrated
by Gallpe (1950), the algal layer of almost all
species of the Alectorioid genera is located
just inside the cortex. If the medulla fills the
filament cavity (as in Alectoria vancouverensis
and extreme morphotypes of A. sarmentosa
subsp. vexillifera), one may find algal cells
even in the centre of the filament, but in a
much reduced number. In addition, the distri-
bution of algal cells along the length of the
filament is often very discontinuous and
clumped (Galloe 1950 figs. 573, 570, 575).
For example, there are no algae in the blackened
“fragmentation areas” of e.g. B. trichodes
subsp. americana (see Hawksworth 1972).
No studies on the taxonomy of the algal
components have been carried out in connection
with this revision. This would have necessitated
investigations of the algae from pure cultures.
Further studies on the systematics of lichen
phycobionts at the species level are desirable
in view of their possible role in influencing
thallus form and chemistry (see the section
Taxonomic concepts: species).
20 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 16. Cortical material invading the medulla. - A. B:
Alectoria vaneouverensis. B. C.. Bird 28472 (CANL).
- A: Longitudinal section, phase contrast. Scale 60
(‘m. - B: Cross section, light microscopy. Scale 20
pm. - C: Bryoria pseudocapillaris. Holotype, cross
section, phase contrast. Scale 20 pm.
Medulla
The medulla in most Alectorioid species is
very lax and “arachnoid" (Hawksworth 1972
pi. I c; Fig. 14). Most species have medullary
hyphae between 3.5 and 5.0 /urn in diameter.
The hyphae are generally smooth, but in most
species of Alectoria s. str. the medullary
hyphae are “ornamented” with knobs and
projections (Fig. 15). The only exceptions
were A. ochroleuca and A. nigricans. A similar
ornamentation occurs rarely in Bryoria
fuscescens (M. Skytte Christiansen, in litt.).
Among those species with a consistently
dense medulla (Alectoria vaneouverensis, A.
mexicana, A. fallacina, Bryoria pseudocapil-
laris), a few have bundles of cortical-like
hyphae running through the medulla, joining
and then leaving the cortex proper. Most
noteworthy in this regard is A. vaneouverensis
where the fibrous medulla apparently can be
used as a species character (Fig. 16 A, B).
According to the drawings of Galloe (1950
figs. 603, 604, 607), the same is true of A.
sarmentosa subsp. vexillifera. An anastomosing
network of conglutinate fibres was also seen
in the medulla of Bryoria pseudocapillaris
(Fig. 16 C).
Chemistry
Methods
Spot tests
A great many misidentifications exist in herbaria
and even in the literature due to errors in
interpreting gross colour tests with the standard
PD. KOH, and C reagents. With few exceptions
(mentioned below) we believe it is possible
to use these reagents with reliability if certain
procedures are followed.
For our tests, “K” is a 10% aqueous
solution of KOH, “C” is fresh, undiluted
household bleaching solution (such as Clorox
or Javex in North America and Parazone in
the United Kingdom) which is a concentrated
sodium hypochlorite solution, and "PD” is
either a solution made from a few crystals of
dry p-phenylenediamine in one or two drops
of 70% ethanol, or Steiner's solution, a
saturated aqueous solution of PD in 10%
sodium sulphite to which a few drops of a
strong wetting agent (detergent) has been added.
For a “KC” test, a small quantity of C is
added to a spot still moist from the application
of K. A “CK” test is made by reversing the
order of application of these two reagents.
“IKI” solution (Melzer's reagent, undiluted
or diluted 1:1) was tested on all species of
Bryoria , section Divaricatae , and on a number
of other species from other sections and
genera. No cortical or medullary reaction such
as that reported by Bystrek (1969) for “Alectoria
Perspinosa" Bystr. was seen in our material.
The reagents are applied in various ways
depending on the tissues being tested or the
substances expected. In most cases, tests of
medullary reactions are most satisfactorily made
with melting point capillary pipettes of about
1-6-1. 8 mm outside diameter drawn to a fine
tip. The reactions are observed under a dis-
secting microscope. As is the case with most
lichen spot tests, the smaller the quantity
of reagent, the more sensitive the test. The
most sensitive PD tests of soralium reactions
are made with a capillary tube and fresh
alcoholic PD.
Almost all other PD tests and most K tests
are made in the following manner. About 5 to
15 filaments of the specimen, including some of
the finest tips and growing portions, are put
on a small piece of white filter paper (about
one cm 2 ). The paper squares are placed on a
microscope slide which, in turn, is on a piece
of cardboard to prevent the table surface from
becoming stained (Fig. 17). The filaments are
then flooded with two or three drops of
reagent (K or Steiner’s solution). There must
be sufficient liquid reagent remaining around
the filaments to permit the diffusion of any
coloured material out of the thallus and onto
the paper. On the other hand, if too much
reagent is used, the chemical reaction will be
obscured by excess dilution.
A positive reaction might be indicated in
one of two ways. The more obvious one is the
oozing out of colour onto the filter paper.
This will occur with all species having PD +
yellow reactions and with thalli containing
fumarprotocetraric acid in the outermost portion
of the cortex (Fig. 18 A). Equally dependable
is the PD + red reaction of the inner portion
of the cortex together with the medulla which
produces no colour at all on the filter paper
(Fig. 18 B, C). The tissues turn deep red and
the reaction is visible through the pigmented
outer cortex which is rendered semi-transparent
by Steiner’s PD solution itself. If the fumar-
protocetraric acid is abundant, permeating
almost the entire cortex as well as the medulla,
the entire filament will seem to develop a red
22 Alectoria in North America
Fig. 17. Filter paper method for colour reactions.
Three squares of white filter paper have been placed
on a microscope slide. The piece in the center with
some filaments of Bryoria sp. has been flooded with
Steiner’s solution; the piece on the right is about to be
treated .
colour (Fig. 18 C). However, unless the surface
of the cortex contains the substance, no pigment
will ooze out (except at breaks in the cortex,
e.g., at soralia or pseudocyphellae). More
frequently, the red reaction is indicated by a
thin red "line” within the filaments (Fig. 18 B).
The test is extremely sensitive, sometimes
detecting the presence of fumarprotocetraric
acid in small portions of a filament even
when TLC techniques fail.
The position of a positive PD reaction appears
to be relatively species-specific. As seen in
Fig. 19, one can classify the reactions into five
categories: (A) occurring in the outer cortex
alone; (B) occurring in the inner cortex alone
(sometimes expanding to include the entire
cortex, as in B ); (C) including all tissues
except the outermost part of the cortex; (D)
occurring both in the outer and inner portions
of the cortex (and usually the medulla as
well); (E) occurring in the medulla alone.
Type E reactions are encountered most
frequently in the genus Alectoria, as in the
thamnolic acid test with A. imshaugii (see
comments below). It never has been seen
as a fumarprotocetraric acid reaction. Type A
reactions are due to the presence of the acid
on or just below the thallus surface. As a
PD+ red reaction, it is seen only in Bryoria
subcana and rarely in B. fuscescens. On the
other hand, it is the characteristic reaction
OPERA BOT. 42 (1977)
Fig. 18. Types of reactions on filter paper with
Steiner's solution. - A: If there is a positive cortical
reaction, the resulting colour diffuses out onto the
filter paper. A positive KOH reaction will have the
same effect. - B; The Steiner’s solution "clears" the
filament and a positive reaction of the inner cortex or
medulla is visible through the cortex as a red, cen-
trally located line. - C: If the inner tissues react
strongly, the entire filament will turn red, but the
colour will not diffuse out.
for alectorialic, barbatolic, norstictic and
salazinic acids. Specimens producing a strong
PD+ red oozing reaction are most frequently
of type D.
The most commonly seen reactions, however,
are those of type B and C. American Bryoriae
of the section Divaricatae which contain fumar-
protocetraric acid (e.g., B. furcellata, B. nitidu-
la) are always type C. Bryoria trichodes (both
subspecies) usually give a type B reaction
(occasionally developing into B'), but thalli
with type C reactions have also been seen.
After the addition of PD solution, Bryoria
cervinula starts as either type B or C, but
generally finally oozes some red coloration
(thus resembling a type D reaction) apparently
due to the sheer quantity of acid in the cortex.
In some species fumarprotocetraric acid
appears to be largely restricted to the soralia
(e.g. Bryoria chalybeiformis, B. glabra, B.
lanestris) so that an intense PD+ reaction is
obtained from these structures which is very
strictly localized (a phenomenon apparently
first recorded and figured by Frey 1952, for
"A. jubata var. prolixa”). When testing speci-
mens for cortical and medullary reactions
OPERA BOT. 42 (1977)
Chemistry 23
B 1
Fig. 19. A classification of possible colour reactions on filament cross sections. See text for explanation.
with this reagent, it is consequently important
to test portions of the thallus which lack not
only mature but also developing soralia (these
appear as somewhat swollen areas or breaks
on the branches). In a few species PD reactions
are characteristically rather ‘‘patchy " and it may
be necessary to test several branches before
concluding a specimen is really PD— (this is
especially true for B. tenuis; see Discussion
under this species). Blackened basal stems of
some species are often apparently devoid of
algal cells and habitually fail to provide PD +
reactions even when these are obtainable in
other parts of the thalli (e.g., B. bicolor, B.
cervinula, B. nadvornikiana, B. tenuis).
The K. test conspicuously fails with species
containing the K+ red substances, norstictic and
salazinic acids. With those species, a positive
K test is typically indicated on the filter paper
by a flash of yellow followed by the formation
of a brownish red, orange-brown, or “dirty”
brown ring at the edge of the reagent front.
The yellow reaction is easily missed, and unless
the substance is very abundant, the brownish
colour may be very diffuse. The PD+ yellow
(to yellow-orange) reaction of norstictic and
salazinic acids is constant, however. With a K
reaction which appears to be negative, one
might assume the presence of psoromic acid
and thus be in error. Fortunately all of these
substances are very easily demonstrated by
microchemical crystal tests (see below).
The KC medullary reactions in the genus
Alectoria are generally without problems,
although it is well to test a portion of the
filament having well-developed medullary
hyphae rather than a portion which is virtually
hollow. It is interesting to note, in this connec-
tion, that except for usnic acid, alectorialic
acid, and one K+ red unknown, all the sec-
ondary metabolic substances found in the genus
Alectoria s. str. are apparently restricted to the
medulla (e.g.. alectoronic, olivetoric. thamnolic,
diffractaic, and presumably barbatic and
squamatic acids). (An exception worth noting
is the thamnolic acid-containing chemotypes
24 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 20. The KC test using a square
of filter paper. See test for descrip-
tion.
of A. sarmentosa and A. vancouverensis from
California, some of which produce this acid
in the outer cortex and not in the medulla
at all.)
The KC test (K followed by C) for barbatolic
acid is often most easily demonstrated by
adding a small drop of K to a concentration
of filaments and absorbing this onto a piece
of filter paper as the drop turns deep yellow
(Fig. 20). If a spot of C is placed next to the
yellow spot formed by reaction with K and
allowed to run into it, a pinkish red colouration
is produced where the spots overlap which is
easily seen against the white of the filter paper.
Care should be taken not to employ too much
C since excess C will itself rapidly bleach
the pinkish red colour.
A positive CK test (C followed by KOH)
was often useful in establishing the presence of
diffractaic acid in Alectoria ochroleuca. If
the acid is present in fairly high quantities,
one gets a strong yellow-orange or gold reaction
in the medulla. If the acid is in low concen-
tration, the reaction is correspondingly weaker.
The same test for diffractaic acid was dis-
covered independently on Parmelia diffractaica
Essl. and was reported by Dey (1974).
The general rule that the best reactions can
be obtained from rapidly growing tissues such
as soralia, pseudocyphellae, filament tips and
apothecial margins applies to Alectoria and its
related genera as it does to other lichens.
Microchemical tests (MCT)
Crystal tests were made most frequently in the
identification of norstictic, salazinic and
psoromic acids. The former two were deter-
mined from acetone extracts using “double-K”
(KK, K 2 ; 5% potassium hydroxide and 5%
potassium carbonate. 1:1). Psoromic acid was
demonstrated using GE solution (glycerine and
acetic acid, 1:3; Fig. 21 D). Gyrophoric and
olivetoric acids were also verified using micro-
chemical tests, but thin layer chromatography
proved more reliable. GE, or GAQ (glycerine,
ethanol, quinoline, 2:2:1) was used for gyro-
phoric acid tests, and GAW solution (glycerine,
ethanol, water, 1:1:1) was used to detect
olivetoric acid. Crystal tests are necessary to
distinguish diffractaic acid from barbatic acid
since the two substances run together in TLC,
but, as Kurokawa (1967) points out, diffractaic
acid is often very difficult to demonstrate.
With adequate concentrations of the substances,
however, they can readily be crystalized from
GE solution. Diffractaic acid produces colour-
less to very slightly yellowish lamellae, with
pointed blade-shaped crystals. They are usually
abundantly overlapping and become fanned out
producing fasciculate clusters (Fig. 21 A, B).
In concentrated preparations, the clusters
become arborescent and curved (see Asahina
1936 p. 871, Kurokawa 1967), and some
crystals curve strongly to one side or produce
a longitudinal split and side branches, but this
curving and “curly” form is relatively rare.
On the other hand, the characteristic “splitting”
can often be found.
OPERA BOT. 42 (1977)
Chemistry 25
Fig. 21 . Crystals produced by lichen substances in GE solution. - A, B: Diffractaic acid from Alectoria
ochroleuca. - C : Barbatic acid from A. vancouverensis. -D: Psoromic acid from Bryoria implexa.
Usnic acid crystals are easily distinguished
by the fact that they are always truncate at
the ends and, of course, they are generally
yellow. Barbatic acid in GE produces short,
Prismatic or rhomboid colourless crystals (Fig.
21 C). Alectoronic acid crystals are colourless
lamellae, always straight and truncate, and
occurring in radiating clusters. Alectorialic
acid could often be demonstrated with GAoT
solution (glycerine, ethanol, o-toluidine, 2:2:1)
(see Krog 1968 pp. 103-104) or GAQ solution
(Hawksworth 1972 pi. 3 A) but again, TLC
Proved to be more sensitive.
Thin layer chromatography (TLC)
The techniques used in our thin layer chromato-
graphic analyses were largely those recom-
mended by Culberson & Kristinsson (1970)
and later expanded by C. Culberson (1972).
Acetone extractions made in small culture
tubes were spotted on 10x20 cm Merck pre-
coated aluminum sheets with silica gel F-254,
or 60F-254. The latter sheets were equilibrated
in an acidic atmosphere as recommended by
C. Culberson (1974). Runs were made to a
height of approximately 60-65 mm. Spots were
examined in long and short wave ultraviolet
light, outlined in pencil, and then sprayed with
either 10% sulphuric acid, Steiner’s solution
or Mr-diazotized benzidene depending on the
substances being identified. The resulting spots
were then related to the norstictic and
atranorin standards as recommended in Culber-
son & Kristinsson (1970) and C. Culberson
(1972) or compared to standards run alongside
the unknowns. The results were recorded both
within the specimen packets and in a log book.
The chromatogram sheets, being thin and light,
were retained for future reference.
Table 1. Secondary chemical products of North American species of Alectoria, Bryoria, Pseudephebe, and
Sulcaria. Filled circle: present in all specimens examined. Half-filled circle: present in most. Unfilled circle:
present in some. Question mark: doubtfully present in some. Literature reports, not necessarily based on
North American specimens: (DC. Culberson 1969. (2) C. Culberson 1970. (3) Hawksworth 1972. (4) Huneck &
Follmann 1970. (5)Solberg 1967. (6) Solberg 1975.
Orcinol series
^-orcinol series
Depsides Uepsidones Depsides
Depsidones
Taxa
ALECTORIA
fallacina
imshaugii
O
O
3
lata
O
3
3
mexicana
. •
nigricans
4 5
ochroleuca
O
1 2
3
4
sarmentosa
3
O
3 4 0
3
O
subsp. vexillifera
3
o
.24.
vancouverensis
3
O
O
O
O
BRYORIA
sect. BRYORIA
chalybeiformis
fuscescens
glabra
lanestris
subcana
trichodes
subsp. americana
vrangiana
sect. DIVARICATAE
bicolor
carlottae
cervinula
furcellata
nitidula
simpl icior
tenuis
O
sect. SUBDIVERGENTES
abbreviata
oregana
sect. T0RTU0SAE
fremonti i
tortuosa
sect. IMPLEXAE
capil laris
friabil is
implexa
nadvornikiana
pi kei
pseudocapillaris
pseudofuscescens
salazinica
spiralifera
4
4 4
4
4 1 4
• •
• 9
• •
6 • • • • 4 . . 4
4
PSEUDEPHEBE
minuscula
pubescens
O
O
3
3
O
O
o
o
o
?
o
o
o
O 3
SULCARIA
no substances
OPERA BOT. 42 (1977)
Chemistry 27
We had some difficulty with resolution in
solvents “B" and “C” until it was discovered
that the spot crowding was due to our use
of an Eastman Chromagram '■Sandwich”
Chamber Plate set. From that point on, standard
chromatogram tanks (c. 25 cm high, 10 cm
deep, 22 cm wide) were used for those solvents.
The tanks did not require filter paper liners.
Solvent "A” runs were no different in the
tank, and so the Sandwich Chamber was
used for those.
There was no difficulty in identifying the
majority of compounds, although a number
of spots of uncertain identity which appeared
in some specimens of a species and not others
were ignored. It is possible that some of these
compounds are of importance and should be
studied further. Of course, there were some
unknowns of obvious significance, and these
were always recorded (see Table 1 ).
Other compounds, particularly some of the
/3-orcinol depsides, run at similar Rfs in all
commonly used solvents. It was therefore
necessary to perform microhydrolyses on them
as described by C. Culberson (1972) to verify
their identity. The technique worked extremely
well as long as Culberson's instructions were
followed carefully.
Since chloroatranorin and atranorin are
virtually indistinguishable with solvents A, B.
and C (C. Culberson 1972), a separation of
these two compounds was made by using
chloroform-acetone, 1:1 as a solvent. In this
system, most other PD compounds found in
our species run close to the source. Chloro-
atranorin and atranorin are then revealed,
using a spray of Steiner’s solution, at Rfs of
31-57, and 75-95, respectively. It is apparent
that the ranges are very broad. In this solvent,
it seems the more concentrated the substance,
the higher it migrates.
Results
An unexpected development in this study
was the discovery of a number of lichen
substances new for Alectoria s. lat. in the North
American species: two orcinol depsides,
gyrophoric acid and olivetoric acid; and one
3-orcinol depsidone, salazinic acid. Thamnolic
acid was reported for A. nigricans by Solberg
(1967) and by Huneck & Follmann (1970) for
a number of other species (see Table 1) but
this substance could not be confirmed in our
work on North American material of those
species. (In a more recent study of A. nigricans,
Solberg (1975) found only alectorialic acid,
a mixture of tetrahydroxy fatty acids, and
5,7-dihydroxy-6-methyl phthalide; no mention
was made of the presence of thamnolic acid.)
On the other hand, several other species do
indeed contain thamnolic acid, as will be pointed
out below. There are a number of unknown
substances which apparently are of some tax-
onomic importance as well.
As might be expected from a reading of the
recent literature (especially W. Culberson 1969,
and W. Culberson & C. Culberson 1970),
the substances which we found seem to fit tax-
onomic groups distinguishable by traditional
morphologic criteria. We should add that this is
particularly true when we disregard the some-
times conflicting literature reports and rely
only on our own investigations.
For example, orcinol derivatives are abun-
dantly represented in Alectoria s. str., the only
genus to produce olivetoric and alectoronic
acids. (A specimen of Sulcaria sulcata from
Japan (Yoshimura 1680, Shikoku [BM]) appar-
ently contains some alectoronic acid, besides
psoromic acid and atranorin, as evidenced by
repeated TLC tests). In the olivetoric acid
containing specimens, in all but a few rare
cases, the olivetoric acid entirely replaces
the alectoronic acid and characterizes a mor-
phologically distinguishable species. The only
other orcinol derivative, the tridepside gyro-
phoric acid, is found in two species of Bryoria.
All the /3-orcinol depsides with the exceptions
of atranorin. chloroatranorin and barbatolic
acid are restricted to Alectoria s. str.; atranorin
and chloroatranorin appear to be entirely absent
from that genus. We have examined the North
American specimens of A. sarmentosa reported
by Hawksworth (1972) as containing atranorin,
and no trace of atranorin could be found in
them by TLC. The identity of the high Rf
compound which he detected by paper chro-
matography remains unknown. The reports of
Huneck & Follmann (1970) should be regarded
with some scepticism until they are confirmed.
Usnic acid is restricted to this genus. On the
other hand, /3-orcinol depsidones, so common
28 Alectoria in North America
OPERA BOT. 42 (1977)
in Bryoria, are entirely absent from Alectoria
s. str.
Fumarprotocetraric acid is by far the most
common chemical in the species under study,
having been found in 14 out of 38 species. It
is apparently a constant constituent of Bryoria
nadvornikiana occurring along with alectorialic
and barbatolic acids and atranorin and is
quite possibly the basis for the report of
thamnolic acid in that species by Huneck &
Follmann (1970) (see comments by Jorgensen
1972 p. 196).
Although Fluneck & Follmann (1970) reported
atranorin in a number of ‘Alectoria” species,
it appears that this compound and the related
chloroatranorin have relatively restricted
occurrences. Sulcaria hadia and some other
non-American Sulcaria species contain
atranorin alone; Bryoria trichodes and B.
fuscescens have chloroatranorin alone; B.
nadvornikiana in almost all cases has both
atranorin (in traces) and chloroatranorin. In
the specimens tested so far (several dozen)
chloroatranorin has been restricted to the
sorediate morphotypes and the subsp. trichodes
of B. trichodes although not all specimens of
this taxon contain the substance. It also appears
that only specimens of B. fuscescens east of
the Rocky Mountains contain chloroatranorin,
but again, the presence of this substance is
not constant. Solberg (1975) reported the
presence of atranorin in one specimen (out of 4)
of Bryoria fremontii, and in a 19.6 gm sample
of Bryoria bicolor. These reports should be
verified.
Barbatolic acid, a /3-orcinol depside best
classified as a benzyl ester (C. Culberson 1969,
Persson & Santesson 1970), is found in B.
capillaris, B. pseudocapillaris, and B. nadvorni-
kiana as characteristic components, and occa-
sionally in Alectoria nigricans.
Alectorialic acid is a very similar benzyl
ester (Persson & Santesson 1970). It is the
main constituent of Alectoria nigricans and
Bryoria pikei. It is always present at least in
trace amounts along with barbatolic acid in
B. capillaris and B. nadvornikiana, a fact
not surprising in view of the similarity of the
two compounds in chemical structure. Speci-
mens in the B. capillaris- aggregate with large
amounts of alectorialic and trace amounts of
barbatolic acid seem to be morphologically
distinguishable and have generally been re-
ferred to B. pikei (see Discussion under that
species). The depsidone salazinic acid occurs
in a species closely related to B. implexa and
B. capillaris (named B. salazinica here).
Norstictic acid has so far been found only in
the very variable species, B. pseudofuscescens,
and in the new species B. spiralifera. In a
puzzling specimen from Dungeness, Washington
(Foster 2552 a [FF1]), both norstictic and barba-
tolic acids were found, pointing out how
closely related B. pseudofuscescens and B.
capillaris are. If lichen-forming fungi hybridize,
this specimen could be thought of as a hybrid.
The presence of barbatic acid in many
specimens could neither be confirmed nor
denied due to the possible confusion of that
substance with diffractaic acid in TLC. The
substance turned out to be diffractaic acid in
Alectoria ochroleuca in all cases in which it
was checked using the microhydrolysis methods
ot C. Culberson (1972), or crystal tests in
GE solution. Spots corresponding to these
substances were seen with several specimens
ot A. vancouverensis and with A. sarmentosa.
Since microchemical crystal tests conclusively
established the presence of barbatic acid in an
aberrant specimen of A. vancouverensis it is
likely that barbatic acid also occurs in some
other specimens of vancouverensis and
sarmentosa. Krog (1968) reported having
demonstrated good barbatic acid crystals in
GE solution for A. “vexillifera" as well as
A. ochroleuca, and she reported barbatic acid
from several ‘‘varieties'’ of sarmentosa.
Squamatic acid has been reported as frequent
in A. sarmentosa (Ftawksworth 1972), but
although this is true for Europe, we could
not detect this substance in the North American
material. It is interesting that the corresponding
metadepside, thamnolic acid, was seen in
several west coast specimens. This may be
another case ot vicarious chemical races within
the well-documented west coast — west Euro-
pean disjunct distribution pattern. Alectoria
imshaugii demonstrates the interchange-
ability of squamatic and thamnolic acids within
a taxon or closely related taxa, as in Cladonia,
sect. Cladonia (e.g., C. squamosa (Scop.)
Hoffm. and C. subsquamosa (Nyl. ex Leight.)
Vain.). Thamnolic acid occurs as a main
constituent only in A. imshaugii .
OPERA BOT. 42 (1977)
Chemistry 29
One of the more interesting unknown sub-
stances, again occurring only in Alectoria
s. str., produces K+ red (persistent) and
C + green-black (ephemeral) reactions on the
thallus cortex. It is generally restricted to the
older parts of the plants, especially at the bases
of terricolous species, but occasionally can
be demonstrated throughout the plant, as in a
remarkable specimen of Alectoria nigricans
from Southampton Island (Parker SP-70-14
[CANL]). Chromatographic results confirm
that the substance is often present above the
base in many species, but in very small
amounts. The substance has relative Rf position
values of 3 or 3-4, 3, 2 in solutions A, B & C
of Culberson & Kristinsson (1970), is strong
UV+ blue-white under both long and short
wave illumination, and is PD-. In sulphuric
acid charring, it is yellowish to pale brown.
This is almost surely the substance responsible
for the K test referred to by Räsänen (1939)
for his “f. rnhropunctata Räs." of A . sarmen-
tosa, and by Krog (1968) for A. “vexillifera" .
Solberg (1975) recently reported a PD— substan-
ce with low Rf values in the same solvents from
Norwegian material of A. nigricans. This
substance, named “AN-X”, was identified
by Solberg as 5,7-dihydroxy-6-methyl phthalide.
It is conceivable that our unknown K+ red,
C+ green-black substance may be Solberg's
AN-X, but we have not had an opportunity
to test this possibility.
Other unknown substances and chemical
problems specific to certain taxa will be dis-
cussed later in the treatments of those taxa.
Certain substances presented particular prob-
lems in either TLC or microchemistry. These
Problems may be the cause of some of the
unconfirmed reports of substances in the genus.
For example a spot which appeared to corre-
spond to a-collatolic acid often, but not always,
occurs as an accessory substance with alecto-
ronic acid. This joint occurrence is well docu-
mented (see Culberson & Culberson 1968)
but we feel that the presence of a-collatolic
acid requires verification in Alectoria. However,
alectoronic acid apparently degrades in sol-
vents B & C resulting in several spots, one
highly fluorescent and the other not.
Alectorialic and barbatolic acids often occur
as mixtures, but not always. If barbatolic
acid is absent, the cortex of the specimen
is often K- or K+ very light yellow; if it is
present, the specimen has a distinct K +
yellow reaction. Both alectorialic and barbatolic
acids are C+ and KC+ pink. The taxonomic
significance in the variation of this chemical
pair is still not clear (see Discussion under
Bryoria capillaris).
The appearance of olivetoric acid in a group
already known to contain alectoronic acid
was not extraordinary, since these substances
occur in closely related taxa in Cetrelia and
are biogenetically related (Culberson & Culber-
son 1968 p. 472). Gyrophoric acid is bioge-
netically isolated from the other orcinol com-
pounds, and it is therefore perhaps not sur-
prising that it occurs in a different genus.
Besides the compounds discovered and re-
ported here, it is conceivable that through
the use of specialized techniques such as mass-
spectrophotometry, other compounds may be
found in these genera, or that certain compounds
will be found in trace amounts in species
other than those for which they were reported
here.
Taxonomie concepts
Genera
Hawksworth (1972 p. 199) emphasized that a
case could be made for recognizing the four
subgenera of Alectoria as distinct genera.
Subsequent investigations reported here have
revealed additional chemical and anatomical
characters supporting this separation. In view
of the now overwhelming evidence (Table 2)
it is clear that the maintenance of Alectoria as
a single genus cannot be justified and the four
subgenera of Hawksworth ( 1972) are thus treated
as genera here. The characters distinguishing
the genera are mostly independent of one
another and clearly are of considerable taxo-
nomic importance (i.e., ascospore colour, size
and septation; the major groups of lichen
products present; cortical structure).
The suggestion that Alectoria be subdivided
is by no means new. Fries (1860) was the first
author to clearly distinguish units at the rank
of genus, although he later modified his views
(Fries 1871). Fries' earlier views were, how-
ever, followed by a few authors (e.g., Stein
1879) but have not been generally accepted
by most specialists concerned with the genus
(e.g.. Du Rietz 1926, Motyka 1962, 1964,
Stizenberger 1892). Bystrek (1971) proposed
that Alectoria should be divided into three
genera but presented no new information
supporting his treatment. Data which have come
to light in the course of the present study,
however, support Bystrek's conclusions, and
his generic concepts are basically those adopted
here (with the addition of Pseudephebe).
Unfortunately this taxonomic decision means
that a new generic name is required for the
bulk of the species formerly placed in Alectoria.
Bryopogon Link cannot be taken up in the
manner proposed by Bystrek (1971) on nomen-
clatural grounds (see Discussion under Bryoria).
We were first reluctant to take this step in
the interests of nomenclatural stability but
following discussions with various colleagues,
especially during the XII International Botanical
Congress in Leningrad, we came to the con-
clusion that this should be done.
In the last decade there has been an in-
creasing tendency to recognize subdivisions
of well-established genera at the rank of genus
in the lichen-forming fungi (e.g., Poelt 1965,
Hale & Culberson 1970, Hale 1974). In most
instances these changes in taxonomic opinion
reflect a better understanding of the species
concerned leading to more natural groupings.
This is also true for Alectoria s. lat. We might
add that there is now probably more justi-
fication for subdividing Alectoria into several
genera than there is for the recognition of some
segregates now becoming widely accepted from
Cetraria. Cladonia, Parmelia and Physcia.
The characters distinguishing the genera
Alectoria. Bryoria. Oropogon, Pseudephebe and
Sulcaria are indicated in Table 2 and will be
readily apparent from both this and the Key
to the genera; they will not be discussed in
further detail here.
Species pairs
The concept of species pairs (“Artenpaare”)
has come to assume an increasingly important
role in lichen systematics below the generic
level following the important papers of Poelt
(1970, 1972). The essential thesis of the concept
is that existing lichen species which only very
rarely or never form ascocarps and reproduce
by vegetative methods (secondary species)
have been derived from extant or extinct species
reproducing by means of ascospores with
OPERA BOT. 42 (1977) Taxonomie concepts 31
Table 2. Diagnostic characters separating the five Alectorioid genera. Included are characters of non-North
American species.
Character
Alectoria
Bryoria
Pseudephebe
Sulcaria
Oropogon
Ascospores
No. per ascus
2-4
8
8
(6— )8
i
Colour when mature
Brown
Colourless
Colourless
Yellow to
brownish
Brown
Septation
Simple
Simple
Simple
l(-3)-septate
Muriform
Length (gm)
20-45
4-15
7-12
22-44
75-100
Width (gm)
12-25
3-7
6-8
8-15
25-38
Excipulum
Persistent and
Becoming
Becoming
Persistent and
Persistent; incurved at
thallinum
incurved
excluded; not
excluded; not
incurved
least when young
incurved
incurved
Pycnidia
Usually rare
Rare
Common, opening
by a broad ostiole
Unknown
Unknown
Vegetative structure
Cortex
Surface texture
Decomposing
Usually smooth
Smooth
Uneven
Uneven to knobby
Differentiation of
superficial cells
Absent
Present in a few
Present in all
Absent
Usually present
species
species
Thickness (yxm)
Matrix, relative to
50-110
25-45
25-75
15-40
25-60
hyphae
Abundant
Little to abundant
Moderate
Moderate
Little to moderate
Medullary hyphae
Usually
ornamented
Not ornamented
Not ornamented
Not ornamented
Ornamented
Cyphellae-like
perforations
Absent
Absent
Absent
Absent
Always present
Pseudocyphellae
Always present.
Variable, when
Absent
Present in long
Depressed, developing
scattered, con-
present adpressed
longitudinal
into perforations
spicuous, raised
or rarely slightly
raised
furrows
Pigmentation
Usnic acid crystals
Mostly in a super-
In cortical cells
Granular on
Mostly in superficial
in outer cortical
ficial crust
surface
cortical cells
Colour of thallus
layers
Usually yellow
Brown to oliva-
Dark brown to
Grey to badious,
Fuscous to grey; rarely
ceous, rarely grey
almost black
or bright yellow
red-brown
Habit
Pendent to erect
Pendent to erect
± Appressed,
Pendent to
Pendent to
rosette-forming
caespitose
caespitose
Chemical
components
Csnic acids
Present in all but
one species
Absent
Absent
Absent
Absent
Atranorin and (or)
chloroatranorin
Absent
Frequent
Absent
Frequent
Absent
Drcinol depsides
Frequent
Rare (tridepside)
Absent
Absent
Absent
Qrcinol depsidones
ß-orcinol depsides
Frequent
Absent
Absent
Rare
Absent
•cxcl. atranorin s.
lat.)
ß-orcinol benzyl
Frequent
Absent
Absent
Absent
Rare
esters
Rare
Frequent
Absent
Absent
Absent
ß-orcinol depsidones
Pulvinic acid
Absent
Very frequent
Absent
Present
Frequent
derivatives
Hydroxyanthra-
Absent
Rare
Absent
Present
Absent
Quinones
Absent
Absent
Absent
Absent
Present
Unknown: K+ red,
green
Very frequent
Absent
Absent
Absent
Absent
Centre of diversity
Temperate
Boreal
Arctic-alpine
Temperate
Temperate-
subtropical
Number of species
known
8
48
2
3+?
4+?
32 Alectoria in North America
OPERA BOT. 42 (1977)
little or no alternative method of reproduction
(primary species). Primary and secondary
species normally have identical (or sometimes
closely related) chemical components (Poelt
1972, Culberson & Culberson 1973, C.
Culberson & Hale 1973, Hawksworth 1976)
and secondary species have a tendency to
have wider distributions and ecological ampli-
tudes than the primary species if the latter
is extant.
As emphasized above, ascocarps are very
rare in most Alectorioid genera and it is clear
that, for most species, vegetative methods of
propagation are of paramount importance. This
suggests that the bulk of the species, whether
reproducing by thallus fragmentation, spinule
abscission, soredia, or combinations of these,
are to be viewed as secondary. A few species,
however, are clearly primary (e.g., Pseudephebe
minuscula, P. pubescens, Bryoria abbreviata,
B. divergescens, B. subdivergens, Sulcaria
sulcata) but, except in the case of Bryoria
abbreviata of which B. oregana may perhaps
be the secondary counterpart, secondary species
derived from these are unknown. A further
group of species may be primary but currently
in the process of spéciation to form secondary
species since some populations are commonly
fertile, others lack ascocarps and soredia, and
yet others lack ascocarps and form soredia
(e.g., Bryoria trichodes, B. fremontii, Alectoria
sarmentosa). A. sarmentosa is of particular
interest in this respect as A . imshaugii might
have been secondarily derived from some
chemotypes of A. sarmentosa (see Discussion
under A. sarmentosa subsp .sarmentosa).
Many of these secondary species are very
widely distributed on a global scale indicating
that the group is of considerable antiquity.
(The only Alectorioid fossil specimen known to
us is Alectoria succini Mägd. represented by
a single plant embedded in amber discovered
in Tertiary (?) deposits in the Baltic region
(Mägdefrau 1957). This material (examined by
D.L.H.) has an Alectorioid cortex, a single
ascocarp, and a structure that might be a
soralium; superficially it is strongly reminiscent
of Bryoria subcana.)
Species
The production of what is termed a lichen
“species” now appears to result from the
interplay of a fungal partner with a particular
algal one which together form a single biological
unit with a characteristic anatomy, morphology,
and chemistry, and with certain distributional
and ecological requirements. The extent to
which the algal partner can be important in
this connection is only just becoming to be
appreciated (James & Henssen 1976); the im-
portance of its role in genera such as those
treated here, presumably with a single algal
genus as phycobiont, remains unknown. Some,
but not all, chemical components used in
lichen taxonomy may be affected in a complex
manner by the alga present (James & Henssen
1976, Hawksworth 1976). As a result of these
fundamental questions which must necessarily
remain largely unanswered for most lichens for
many decades to come, it is evident that for
practical purposes when we use the term
“species" in the lichen-forming fungi we are
applying it to the dual organism and not only
to the fungal partner. (That for nomenclatural
purposes the names of lichens refer to the
fungal partner alone is immaterial in this
connection.) In the case of the species studied
here, it is of interest to note that with regard
to the ascocarps and ascospores, the mycobionts
involved seem to have undergone very little
divergent evolution (a phenomenon rather
unusual in the non-lichen-forming Ascomycotina
where in the larger species, differences in
vegetative tissues tend to be correlated with
differences in ascocarps and ascospores).
The lichen taxonomist has to delimit his
taxa on the basis of examinations of large
numbers of herbarium specimens and studies
of populations in the field. If too few herbarium
collections are studied and sufficient field work
is not carried out, there will be a tendency
to recognize too many taxa. This is due to
an incomplete sampling of populations which
precludes any reliable assessment of the extent
of environmental impact on the genotype.
(See the discussion of Hawksworth 1973 on this
topic.) In the case of Alectoria, environmental
factors exert profound effects on thallus colour,
contortion, and the production of secondary
branchlets and spinules. On the basis of our
OPERA BOT. 42 (1977)
Taxonomie concepts 33
field and herbarium studies we have, in most
instances, been able to arrive at what we
consider to be clearly delimited taxa. Most
prove to be readily recognizable in the field
(indeed some appear more distinctive there
than when pressed in the herbarium) but a few
difficult groups nevertheless remain. In an
attempt to introduce some degree of con-
sistency in the application of taxonomic ranks
in the lichen-forming and non-lichenized fungi.
Hawksworth (1974) proposed that individual
species should be based on well-marked
discontinuities in several apparently unrelated
characters and we have endeavoured to adhere
to this tenet in the present revision.
The taxonomy of Alectoria has been regarded
as exceptionally difficult by many contemporary
lichenologists. While many of the difficulties
prove to be man-made and due to inadequate
study and errors in descriptions which appear
in the literature, our present investigations
have demonstrated that a great deal of work
is still required in some groups of species.
Certainly, we are only likely to arrive at an
entirely satisfactory understanding of the taxa
when more information on the group in Europe
and Asia is available. These problems are
discussed in some detail under the taxa con-
cerned but it is pertinent to refer to two
Particular areas of difficulty here.
Firstly, the Bryoria chalybeiformis-B. fus-
cesccns-t B. glabra)-B. Ianestris-B. vrangia-
na group. (The non-North American B. intri-
cans also belongs here.) While, in our view,
these species are distinct, in a significant
number of cases we have had some difficulty
>n deciding to which of two (or sometimes
three) species a particular specimen belongs.
This problem is discussed in detail under B.
fasces cens. In view of the characteristic
habitats, distributions and distinctness of the
bulk of collections, we are inclined to the
opinion that phenotypic parallelism arising from
environmental stress is of paramount importance
here, particularly as some other species are
a ble to produce morphotypes not dissimilar
from these (these being readily separable,
however, by virtue of distinctive chemistries
0r other characters, e.g., Bryoria trichodes
su bsp. trichodes, B. pseudofuscescens).
Secondly, during the course of the present
survey, a number of chemotypes came to light
3 " Opera Botanica nr 42
which had been previously entirely unknown.
In most cases, species delimited here on the
basis of morphological criteria have both
distinctive distributions and constant (or ±
constant) chemistries. The difficulties for the
taxonomist arise from the association of two
or more chemotypes with a single morphotype.
The employment of chemical criteria in lichen
taxonomy has been a source of controversy
for over a century. In a recent review of
this subject Hawksworth (1976) proposed some
guidelines for their taxonomic treatment which
have been adhered to here except that in-
fraspecific taxa have not generally been
employed (see below). Attention is also drawn
here to the problems in the Bryoria capillaris,
B. fuscescens, B. pseudofuscescens and
Alectoria sarmentosa groups. In addition, a few
apparently very rare chemotypes have come
to light which might conceivably be hybrids
(pp. 71, 116, 119). The phenomenon of hy-
bridization has only recently been suggested as
an important consideration in lichen evolution
(C. Culberson & Hale 1973). If it occurred in
the evolutionary past in one group, it presum-
ably may well be occurring in others at the
present time. A discussion of the possible
mechanisms involved can be found in con-
nection with the chemical variation in Alectoria
sarmentosa (see Discussion under this species).
Infraspecific taxa
In the early stages of the present work we
were inclined to recognize a number of infra-
specific taxa within certain species but in the
final analysis decided to refrain from doing this
except in two cases where subspecific rank
seemed particularly appropriate ( Alectoria
sarmentosa subsp. vexilUfera, Bryoria trichodes
subsp. americana ). This decision is not to be
taken as indicating that no further taxa meriting
infraspecific ranks occur among the North
American species studied here. It was simply
done because we felt that our primary task
in undertaking the present survey was to
endeavour to provide a satisfactory delimitation
of the species. In the future more detailed
studies of the species accepted here should
be carried out with a view to analyzing some
of the variations in chemical components and
morphology noted in the discussions of the
34 Alectoria in North America
OPERA BOT. 42 (1977)
various species. In this way, one may be able
to arrive at soundly based conclusions as to
the taxonomic importance of these variations.
In many instances such studies will also have
to consider non-North American material.
Within the discussions of individual species
(and occasionally in the descriptions of them)
we have, however, noted the names of some
of the more distinctive infraspecific taxa
accepted by European authors which also occur
in North America.
Ecology
Although it is admittedly hazardous to gen-
eralize about the ecology of a series of four
genera of lichens, in the case of Alectoria,
Bryoria, Pseudephebe, and Sulcaria, certain
general statements can be made which will
indeed characterize the majority of the species.
With regard to the corticolous species,
coniferous trees and trees with similar bark
characteristics such as birch are by far the most
frequently encountered substrates. However,
it is not at all clear whether the species
actually prefer acidic bark (in a statistical
sense) due to some physiological requirements,
thus explaining their abundance in coniferous
forests, or whether the species require the
climate and general environment of coniferous
forests and simply happen to be found on the
most available substrate there, namely conif-
erous trees. It seems the latter is more likely,
to judge by the wide variety of dominant tree
species used as substrates by the lichens in
each forest type.
Rock-dwelling species include those enduring
highly exposed and well-illuminated surfaces
(e.g., Pseudephebe pubeseens) and those
frequenting more shaded rock walls (e.g.,
Bryoria nadvornikiana). The rock substrates,
however, are almost invariably non-calcareous.
Soil-dwelling species generally can tolerate a
wide variety of soil types and pH levels.
It would probably be instructive to examine
various habitat types with the aim of determining
which species share the same habitats and are
likely to be found together. It is one of the
burdens of would-be “Alectoriologists" that
these species very frequently grow mixed
together and are collected that way. Knowing
which species are likely to be found together
may be a help in their determination.
Since the western mountains of North
America show a great diversity of species,
we will begin the discussion by trying to dissect
out the various montane forest types and
describe the species most characteristic of
each. At the outset, it should be pointed out
that other species may be present in these
communities; we are merely listing the most
characteristic ones.
The mountains we are mainly concerned with
here are centered in the Canadian Rockies
(basically along the British Columbia-Alberta
border) and Columbia Mountains, extending
southward to include most of the Idaho and
Montana mountains north of the Columbia
plateau. Most of the ecological information in
the discussions pertaining to western North
America is based on Krajina (1965).
An examination of the altitudinal distribution
of the 12 most common montane taxa (com-
prising 11 species) (Fig. 22) is interesting but
not very helpful. As one might expect, a
forest type at one altitude on a south-facing
slope may be several thousand feet higher
than the same forest type a few hundred
miles north and on a north-facing slope. On
the other hand, certain species are closely
associated with certain forest types.
The Interior Douglas Fir zone, a rather dry
zone with only about 16-19 inches (410-480
mm) of precipitation per year, is the preferred
habitat of Alectoria imshaugii, Bryoria abbre-
viata, and B. fremontii. The most common
trees in this zone, Pseudotsuga menziesii, Pinus
ponderosa, P. contorta var. latifolia and
Larix occidentalis are also the most frequent
substrates.
The Interior Western Hemlock zone, having
many of the same tree species as the Coastal
Western Hemlock forest (e.g., Tsuga hetero-
phylla, Thuja plicata, Pseudotsuga menziesii).
36 Alectoria in North America
OPERA BOT. 42 (1977)
3.000
o
d
CO
n
<0
as o>
« / ) >
<
Fig. 22. Altitudinal distributions of several species found in the western mountains. Solid line = principal
range; broken line = aberrant extension of the range.
also resembles that forest in being relatively
moist. Montane populations of Alectoria
sarment osa are most frequently found here.
The Sub-boreal Spruce zone is basically an
extension of the northern boreal forest, or at
least, its ecological affinities are with that
forest region. The dominant trees are Picea
glauca, P. engelmannii, and Pinas conforta
var. latifolia. Bryoria lanes tris, a very con-
spicuous boreal forest species, is, not un-
expectedly, also found in this zone.
The Subalpine Abies lasiocarpa-Picea engel-
mannii-Larix lyallii zone found on the higher
mountains, is the characteristic habitat of B.
oregana, B. pseudofuscescens, and B.
vrangiana.
On the alpine heath and rocky ground, one
can find mixtures of Alectoria sarmentosa
subsp. vex illifera, B. chalybeiformis, Pseude-
phebe pubescens, P. minuscula, and, especially
in the northern mountains, Alectoria ochroleuca.
The Pacific Coast Mesothermal Forest is
OPERA BOT. 42 (1977)
Ecology 37
composed of two parts: a relatively dry segment
(26-65 inches (660-1650 mm) precipitation per
year) called the Coastal Douglas Fir zone by
Krajina (1965), and a wetter zone with an
annual precipitation ranging from about 70 to
over 260 inches (1780-6600 mm). The wetter
zone, the Coastal Western Hemlock Zone,
is a veritable rain forest in places. It is
dominated by Tsuga heterophylla, Thuja plicata,
Picea sitchensis, and Acer macrophyllum.
Inhabiting this zone are: Bryoria trichodes
(both subspecies), B. hicolor, B. tenuis, B.
capillaris, B. glabra (which is also montane),
and Alectoria sarmentosa.
The Coastal Douglas Fir zone, w'ith Pseudo-
tsuga, Arbutus menziesii, and Quere us gar-
ryana, often contains the following species:
Alectoria vancouverensis, Bryoria fuscescens,
B. fremontii, B.friabilis, and B. tortuosa.
The Boreal Forest of North America stretches
from Alaska southeastward to Ontario, Quebec
and the maritime provinces with very little
basic change in tree dominance: Picea mariana
and Larix laricina in the wetter sites, and Pin us
banksiana. Picea glauca and Populus tremu-
loides in the drier sites. In the southern and
eastern boreal forest, Abies balsamea is very
important. Throughout this entire region (with
the few exceptions noted below) the following
Bryoriae are found: B. simplicior, B. lanestris,
B. furcellata, B. fuscescens, B. nadvornikiana,
B. trichodes subsp. trichodes, and B. capillaris
(especially in the humid southern boreal section;
see Ahti 1964).
In the Abies fraseri forests of the southern
Appalachian Mountains, one can find Alectoria
fallacina, Bryoria tenuis, and B. bicolor. Bryoria
furcellata is more characteristicly found in
the pine forests of the Appalachian chain.
The coniferous associations of the Acadian
forest on the east coast are indicated by the
presence of Picea rubens. In other respects
they are very similar to parts of the southern
boreal forest but with many more deciduous
species. Often found in mixed collections,
especially from exposed, humid, coastal
localities, are: Bryoria fuscescens, B. trichodes
(especially subsp. americana and its sorediate
morphotypes), and B. capillaris.
In the arctic tundra, Alectoria ochroleuca,
A. nigricans, A. sarmentosa subsp. vexillifera
and Bryoria chalybeiformis are found over
calcareous or non-calcareous soil and rock.
On non-calcareous rock and soil, one also
may find B. nitidula in great abundance.
In the British Isles, the concept of lichen
species which are “old forest indicators” (i.e.,
species which are restricted to very old,
undisturbed sites) has recently been extended
to consider taxa of coniferous woodlands (Rose
1976): Bryoria capillaris and Alectoria sarmen-
tosa subsp. sarmentosa both fall into this
category in that country (Rose 1976). This
phenomenon may be an important aspect of
the local distribution of some species in North
America (e.g., Bryoria abbreviata, B. fremontii,
Alectoria imshaugii) and requires further study.
In Europe Bryoria fuscescens has a tendency
to be abundant in localities affected by man
(Hawksworth 1972) and this also appears to
be true for North America, at least in New-
foundland (Ahti & Hawksworth 1974). B.
furcellata is also an “opportunist” in the east
becoming not uncommon on fence-rails. This
species, together with B. trichodes subsp.
trichodes, is apparently a rapid colonizer of
young coniferous trees and can be found
recolonizing clear-felled areas in Quebec.
Air pollution is considered to be an im-
portant factor in the extant distribution of
several species of the genus in Europe (e.g.,
Barkman 1958, Hawksworth 1972) and this,
together with other effects of urbanization,
certainly limits the occurrence of Alectorioid
species in major conurbations (e.g., Brodo
1968) and in the vicinity of industrial plants
(e.g., Rao & LeBlanc 1967, LeBlanc et al.
1972, 1974). The information currently avail-
able indicates that species of these genera differ
in their sensitivity to air pollutants but is
insufficient to clearly indicate relative sensi-
tivities in north America at the present time;
indeed these might be expected to vary accord-
ing to climate and other factors in such a diverse
area for a variety of reasons (Hawksworth &
Rose 1976).
As noted above, Alectorioid species may
prefer relatively acidic barks in North America
as they do in Europe (Barkman 1958, Hawks-
worth 1972). Sulphur dioxide in the air leads
to bark acidification and thus can enable spe-
cies to occur on phorophytes from which they
are normally absent as long as conditions are
not too severe. In a site in Sweden, Skye &
38 Alectoria in North America
OPERA BOT. 42 (1977)
Hallberg (1969) noted an increase of “A.jubata”
following a prolonged pollution episode,
apparently due to bark acidification. This
phenomenon would also be expected to occur
in North America, enabling species to colonize
deciduous trees under moderate pollution stress.
Distribution
Although there are exceptions, Alectoria,
Bryoria and Pseudephebe as a whole can be
thought of as cold climate genera, with most
species either decidedly northern or associated
with mountains. This is well documented in
North America, where two-thirds of the species
have the major portion of their range within
Canada, and only 4 are found entirely south
of the Canadian border. Sulcaria appears to
be more temperate both in North America
and in Asia. The four genera are apparently
unrepresented in Florida (Moore 1968). or for
that matter, in any of the southeastern states
(South Carolina, Georgia, Mississippi. Alabama,
Louisiana, and Arkansas). They are apparently
also absent from the central plains, probably
due to the hot, dry summers and lack of
coniferous forest. There are no records of
Alectorioid species from Texas, Oklahoma,
Kansas, Nebraska, or North Dakota. The only
species of this group from South Dakota are
from the Black Hills (Wetmore 1968).
With the exception of Bryoria furcellata,
extensions into the southeastern mountains
are restricted to occurrences in the “south-
eastern spruce-fir forest" (Kiichler 1964). The
map of Bryoria trichodes subsp. trichodes
shows this well. The absence of these genera
in the Ozark region (northern Arkansas and
southern Missouri) correlates well with the lack
of any northern forest remnants there.
Other dry, warm areas such as southern
California and the Ponderosa Pine forests of
Montana and Arizona largely lack species trom
this group.
Porsild's (1958) treatment of some geographic
elements in the Canadian flora is one of the
most useful in a discussion of the distribution
of species covered here. Since lichens, like
mosses, have relatively few North American
endemics (see Crum 1966), the same degree
of representation in his “North American
Species” group will not be seen here. In fact,
"wide-ranging" species endemic to North
America, while comprising the bulk of the
Canadian vascular plant flora (Porsild 1958),
are virtually nonexistent in lichens.
Indeed, circumboreal taxa are well-repre-
sented among the North American species of
Alectoria, Bryoria and Pseudephebe. They
are found in Europe in climatologically and
vegetationally equivalent areas, especially in
Scandinavia and the U.S.S.R. but also in the
central and eastern European mountains, and
sometimes in mountainous areas of Asia,
Australasia, and Africa as well.
Circumboreal-circumpolar species
All the arctic Alectorioid species appear to be
wide-ranging in the tundra region. The zonations
with regard to northern distributional limits of
vascular plants which are so well demonstrated
and discussed by Young (1971) apparently do
not apply to lichens. Young showed that
these limits correspond closely with the amount
of available warmth during the growing season.
Since lichens have a “growing season" quite
different from that of flowering plants, and their
microenvironment is even more restricted and
critical than that of cushion plants, it is easy
to see why the same factors would not apply.
Young (1971) also points out that this warmth
is most critical for the flowering and fruiting
process in the vascular plants, something which
would have no bearing on the presence or
absence of asexually reproducing lichens,
although the positive effect of warmth on
ascocarp production in some lichens has been
documented (Sernander-Du Rietz 1957).
40 Alectoria in North America
OPERA BOT. 42 (1977)
To a limited extent, probably due as much
to historical as to climatic factors, one can
recognize some of Porsild’s (1958) floristic
elements among the North American species.
(1) Arctic-alpine element: Pseudephebe minus-
cula, P. pubescens, Bryoria chalybeiformis ,
Alectoria nigricans, A. ochroleuca.
Different species have southward extensions
into the western Cordillera to various degrees.
The alpine range of A. ochroleuca stops close
to the Canadian border (although A. ochroleuca
is seen again in the Mexican highlands and in
South America). Pseudephebe rninuscula is
more abundant in the Rocky Mountain system,
with P. pubescens being most common in the
Cascade ranges.
(2) Low-arctic element: Bryoria nitidula.
Although B. nitidula ranges well into the
Queen Elizabeth Islands, its main area of
distribution is clearly the low arctic. Other
species extending into the low arctic and even
the northern boreal zones (e.g., Alectoria
nigricans and A. ochroleuca ) seem to be
better characterized by their northern limits
rather than their southern limits. Like the
arctic knotweed Polygonum viviparum (see
Porsild 1958), they are best classified with the
wide-ranging arctic-alpine species.
(3) Amphi-atlantic, southern element: Alectoria
sarmentosa subsp. vexillifera.
Although this taxon has disjunct populations
in the western mountains and in the Aleutians,
it is placed here due to its apparent amphi-
atlantic character. Its interesting distribution
is discussed more fully in the taxonomic
part.
(4) Boreal forest element: Bryoria capillaris, B.
fuscescens, B. implexa, B. lanestris, B. nadvor-
nikiana, B. simplicior, ( Alectoria sarmentosa
subsp. sarmentosa).
This element is difficult to interpret on a
North American scale since it has obviously
been dissected during glaciation and reas-
sembled depending upon rates of dispersal,
edaphic factors, and historic factors which
differed from one species to the next. However,
one can conveniently apply the zonations
suggested by Sjörs (1963) and modified by
Ahti (1964) to eastern North America. The
coniferous forests of the western mountains
are not entirely equivalent to those of the
Boreal Forest Region, although, by virtue of
the general similarities of these forests, many
boreal species, especially those of the middle
and northern boreal forest zones, have been
able to survive there.
Bryoria simplicior is a narrowly restricted
Hemiarctic to northern boreal zone plant (see
Ahti 1964). B. lanestris is also found in these
zones, but is more common in the western
mountains. B. fuscescens and B. implexa appear
to be middle boreal zone species, although
the former is much more common and widely
distributed than the latter both in Europe
and North America.
The southern boreal zone is represented by
two species: B. nadvornikiana and B. capillaris.
Both show preferences for strongly humid
locations, as was mentioned by Ahti (1964)
for B. nadvornikiana.
Alectoria sarmentosa seems to have a
severely dissected middle boreal forest distri-
bution, and might better be described simply as
having a bicoastal distribution, although it
is not at all "coastal - ' in the strict sense.
(5) Appalachian-Great Lakes, temperate:
Bryoria furcellata.
Bryoria furcellata is surely the most temperate
ot the North American Bryoriae or Alectoriae.
Having originated as a circumboreal species,
it may well have been displaced southwards
onto the Appalachian chain during glaciation,
and only gradually attained the more northern
boreal localities. In Canada, its northern
limit roughly coincides with the 13°C July
isotherm according to Ahti & Hawksworth
(1974), in agreement with Ahlner’s (1948)
observations for Scandinavia.
(6) Oceanic-suboceanic species: Bryoria
bicolor, B. tenuis.
Bryoria bicolor and B. tenuis have almost
identical distribution patterns in North America,
i.e., they are largely restricted to the humid
and mild coastlines and foggy Appalachian
mountain tops.
The marked disjunction between the highest
■peaks” of the southern Appalachians (espe-
cially in Great Smoky Mountain National Park)
OPERA BOT. 42 (1977)
Distribution 41
and the high mountains of the northeast (e.g.,
the Adirondacks, the White Mountains and Mt.
Katahdin in Maine) is repeated in the distrib-
ution of many lichen taxa including Nephroma
parile (Ach.) Ach. (Wetmore 1960), Pseude-
vernia cladonia (Tuck.) Hale & Culb. (Hale
1955), and Parmelia halei Ahti ( 1966) in addition
to Bryoria bicolor, B. tenuis and Alectoria
fallacina. It is likely that this disjunction
resulted from the gradual reinvasion of popu-
lations towards the north with the retreat
of the last major ice sheets. Presumably,
the intervening areas became too dry or warm
to support these species, some of which, like
those listed above, are clearly suboceanic.
Thus, a careful search of suitable microclimatic
habitats in other parts of the Appalachians
may well turn up some of these species which
require cool temperatures and high humidities.
Widely distributed circumboreal oceanic
distributions of this type are generally
thought of as representing remnants of the
ancient Arcto-Tertiary flora (Schofield 1969).
(7) Western American-European disjuncts:
Bryoria fremontii , B. pseudofuscescens, B.
subcana, (B. tortuosa).
It is widely accepted that this disjunct
Pattern results from a circumboreal distribution
(Schofield 1969), probably eliminated from
eastern North America by continental glacia-
tion. Eastward migration in post-Pleistocene
times was prevented by the expanse of the
Great Basin.
Although B. subcana is the only one of this
group which might possibly be called “oceanic”
(see Hawksworth 1972), moisture requirements
might have limited several species in this
element to the west coast (see Schofield
1969). Alectoria sarmentosa might well have had
an identical history in North America, but
with the survival of an additional remnant on
the east coast as suggested for Platismatia
norvegica (Lynge) Culb. & C. Culb. (W.
C ulberson 1972) which has a similar distribution.
Bryoria tortuosa is tentatively included here
Pending a reexamination of the single European
Plant reported (see under B. tortuosa, Distri-
bution).
This element is very well known among
flowering plants and bryophytes (Schofield
'969, Schofield & Crum 1972).
Asian affinities
(8) Bicoastal, Appalachian-West Coast dis-
juncts: Bryoria trichodes.
Asian-Appalachian disjuncts are generally
thought of as relicts of an ancient Arcto-
Tertiary flora which at one time extended
across Eurasia and later became reduced
through extinctions (W. Culberson 1972). The
presence of the species on the west coast
could have come about from a pre- or post-
Pleistocene establishment from Asia via the
Aleutian route, or the species may have per-
sisted south of the ice along the west coast
(Schofield 1969). The subspeciation of Bryoria
trichodes is even more difficult to deal with,
but morphological evidence points to subspecies
trichodes as being the older of the two, having
more similarities with the Asian populations.
This would be supported by its presence in the
southern Appalachians. Subspecies americana
presumably became distributed across North
America, possibly in a boreal forest environment
that existed along the ice front, becoming
disjunct and restricted to available humid zones
after glacial retreat. (See the discussion under
category 11 below. See also category 15 for a
discussion of Alectoria lata, another Asian-
North American disjunct.)
North American endemics
(9) North Pacific: Bryoria cervinula, B. car-
lottae.
The close relationship between these species
and some Asian species in the "Alectoria
asiatica group” (see also Motyka 1964) suggests
that they may have been pre-Pleistocene, amphi-
Beringian or North Pacific arrivals which be-
came isolated in certain nonglaciated headlands
and slopes along the north Pacific coast during
glaciation (seeCalder & Taylor 1968). Schofield
(1969) elaborates on this element, giving many
examples of species found on both sides of
the Pacific. Other lichens, such as Coccotrema
maritimum Brodo, have similar distributions
and probably had comparable histories (Brodo
1973).
(10) Coastal lowland: Sttlcaria badia, Alectoria
vancouverensis, Bryoria spiralifera, B. pseudo-
capillaris.
42 Alectoria in North America
OPERA BOT. 42 (1977)
All of the species in this category are found
basically between southern Vancouver Island
and northern California either corresponding to
the “cool, moist (conifer) forest" (Daubenmire
1969), or, as in the case of Sulcaria badia,
to the drier Pseudotsuga forest. The evolu-
tionary relationships are varied: Alectoria
vancouverensis is clearly closely related to
A. sarmentosa (a boreal species); Sulcaria
hadia has Asian affinities; Bryoria spiralifera
and B. pseudocapillaris are members of the
section Implexae which seems to have more
boreal-montane, if not oceanic tendencies. All
these species would be included in the “Cali-
fornian” element of Schofield (1969). Species
of this element such as A . vancouverensis
may have spread northward into British Co-
lumbia during the Hypsithermal Interval
(Schofield 1969).
Most California endemics occupy, and ap-
parently developed, in a region with a Medi-
terranean climate (moderate to heavy winter
precipitation and extreme summer drought
(Stebbins & Major 1965). Since California
has never been connected with another area
of Mediterranean climate, the plants there had
to adapt in situ from a flora adapted to other
kinds of climatic regimes. This would have
occurred towards the latter part of the Tertiary
when California developed its present climate.
Stebbins & Major point out that this is why
California has endemic elements from the
northern flora whereas the Mediterranean region
itself, by and large, does not. Alectoria, and
Bryoria of course, would be a good example
of this.
If Bryoria pseudocapillaris and B. spiralifera
are ever proved to be members of the genus
Sulcaria (which is possible, to judge from
their chemistry and morphology) this Californian
element (together with S. badia) would be
represented by three endemic species with
Asian affinities. The presence of many such
“relict endemics” in the coastal northern
California-Oregon area is well documented,
corresponding, for example, to the distributions
of Sequoia, Sequoiadendron, and Torreya
(Stebbins & Major 1965). These trees apparently
represent remnants of the Arcto-Tertiary flora
of the mesic coniferous and deciduous forests.
Alectoria lata is a more montane species
which shows a present-day relict disjunction
with eastern Asia. Alectoria vancouverensis
perhaps evolved somewhere along the coast
and succeeded more than the other species
in invading more northern as well as southern
coastal localities. At the southern edge of its
range, where the climate is marginally accept-
able and the forests and topography are very
mixed, the development of chemical races and
perhaps hybridization would be more likely.
This would be in close accordance with the
thoughts of Stebbins & Major (1965) on the
origin of the California vascular plant endemics.
According to such a hypothesis one would
have to assume that physiological adaptations
in the lichens, rendering them better suited to
survive in these less climatologically stable,
marginal regions, accompanied the changes in
external form and secondary metabolic
products.
(11) Lowland to western montane (humid
forests): Bryoria glabra, B. friabilis, B. pikei.
These species occur from open lowland
to humid montane localities mainly on the
Coast and Cascade Ranges, but often (as with
B. glabra) extending to the humid Interior
Western Hemlock zones in southeastern
British Columbia (see Daubenmire 1969 pp.
120-123). Interestingly, all three species have
disjunctions with humid localities on the east
coast. Packer (1971) lists a number of Canadian
endemic flowering plants with similar bicoastal
distributions.
The question of the origin of such striking
east-west disjunctions has been discussed by
many authors and reviewed by Morisset (1971).
Morisset begins with the assumption that such
disjunctions represent relict populations of
distributions which were at one time continuous.
(Thus he quickly dispenses with long-distance
dispersal.) The possibility of migrations from
the western cordillera eastward along the re-
ceding ice front (the so-called “sidewalk"
hypothesis of Stebbins & Major 1965) presents
some problems for arctic-alpine species. The
presumed tundra belt was apparently extremely
narrow in the west, which would interfere
with any eastward migration, and fossil evidence
is lacking. Boreal forest species, however,
may well have been able to use this route.
On the other hand, Morisset points out that
the exposed coastal plain would have been not
OPERA BOT. 42 (1977)
Distribution 43
only available but ideal as a refugium south
of the ice front, and would have permitted
an easy reinvasion of coastal and even montane
localities as the ice retreated. In the absence
of localities of these bicoastal disjuncts in
suitable microhabitats within the intervening
area, one would have to say that at least
for these lichens, the coastal plain refugium
hypothesis seems the most plausible.
(12) Western montane and intermontane (dry
forests): Alectoria imshaugii, Bryoria abbre-
viata, B. oregano.
The main ranges of these species occur in
the relatively dry, montane Pin us ponderosa-
Pseudotsuga forests which extend from the
Idaho-Washington border (and parts of Mon-
tana) through central Oregon south to the
northern California highlands (Daubenmire
1969).
Alectoria imshaugii may have evolved as an
isolated population in or bordering the dry
interior plains as have numerous other endemic
flowering plants and bryophytes (Schofield
1969). It apparently extended its range to the
California and Oregon coasts in open stands of
Pinus conforta or Pseudotsuga. Bryoria oregana
and B. abbreviata probably had their origin
as more montane endemics.
(13) Appalachians: Alectoria fatlacina.
The Appalachian mountains, due to their
age and long isolation, have been the site of
spéciation in many groups of plants. Even with
the relatively slow rate of evolution in lichenized
fungi, endemism can be seen in the Appalachian
lichen flora, for example, in Cladonia caro-
liniana Schwein, ex Tuck. (Ahti 1973) and
Lecanora caesiorubella Ach. subsp. caesio-
rubella (lmshaug & Brodo 1966) in addition
to Alectoria fallacina. However, many species
regarded as endemics at one time are now
known from other areas of the world, especially
from eastern Asia and the Alps (Schauer &
Brodo 1966), and others have closely related
vicariad species elsewhere (Yoshimura 1968).
(14) Northeastern coastal plain: Bryoria
salazinica.
Many coastal plain endemics are known
in the lichen flora of eastern United States
and adjacent Canada (Brodo 1968). Those with
restricted distributions in New England and the
maritime provinces such as B. salazinica,
and perhaps Cladonia boryi Tuck, (also known
in Japan; see Ahti 1973), probably occupied
the expanded coastal plain as the Pleistocene
ice sheet receded to the north, adapting to,
and then becoming restricted to coastal localities
(see Morisset 1971). Whether the endemic
species evolved here or simply represent
fragments of a once world-wide population is
usually impossible to determine in lichens,
which have virtually no fossil record.
(15) Central American highlands: Alectoria
mexicana, A. lata.
Alectoria mexicana is a rare plant, and its
origins and relationships are still very unclear.
A. lata, on the other hand, is a species of
cool, high montane pine forests in subtropical
and tropical latitudes and occurs in similar
forests in the Pacific Northwest from California
to Washington, as well as in Asia (Hawksworth
1972).
Asian affinities with plants of Mexico have
been the subject of numerous papers (see
especially Sharp 1966). Some Asian-Mexican
disjuncts are known to occur in western Amer-
ican mountains as well. These most probably
represent ancient Arcto-Tertiary distributions,
disrupted by “the rigors of the late Tertiary
and Pleistocene" (Sharp 1966 p. 231).
Importance to man and nature
Few groups of lichens are considered to be
of major importance to man. Cladina, com-
prising the reindeer lichens, is certainly the
most outstanding example of an ■‘economically”
important lichen genus. Alectoria s. lat., as a
“genus”, has probably been used by man
directly and indirectly more than any other
genus except Cladina. Since one of these
uses, that of caribou forage, partly led us to
undertake this study, it seems appropriate to
review the subject briefly, as regards the species
in North America.
Forage
Although the terricolous reindeer lichens are
the most abundant and important winter food
for caribou, arboreal lichens are generally
plentiful enough in the winter range of the
barren-ground and woodland caribou ( Rangifer
tarandus groenlandicus, and R. t. caribou,
respectively; Banfield 1974) to be of major
importance. In some regions such as Wells
Gray Park in eastern British Columbia, arboreal
lichens, especially species of Alectoria s. lat.,
provide caribou with most of their winter
food (Edwards et al. I960). Edwards and his
coworkers go so far as to say that the survival
of caribou in the region depends on the
availability of Alectoria s. lat. In other areas
such as northern Saskatchewan (Scotter 1962)
and northern Ontario (Cringan 1957, Ahti &
Hepburn 1967), Alectoria s. lat. forms a
significant, although not dominant, role in winter
food requirements, especially when the snow
is exceptionally deep or encrusted.
The quantities of arboreal lichens produced
in these forests are impressive. In northern
Saskatchewan, one can find 359-572 kg/acre
in a black spruce forest, and 217-1167 kg/acre
in mature jack pine stands (Scotter 1962).
Maxima of 114 kg/acre and 1332 kg/acre of
dried Alectoria s. lat. can be produced in
lodgepole pine forests and timberline spruce-
fir forests, respectively, of Wells Gray Park
(Edwards et al. I960).
In Saskatchewan, caribou can sometimes
reach branches 3.3 m from the ground giving
them access to an average of 154-275 kg/acre
(Scotter 1960). In Wells Gray Park, where
the snow is often much deeper and the caribou
can reach up to 5 m above ground level, the
animals can find 52-329 (with an average of
116) kg/acre (Edwards et al. 1960 Table I;
the figures given in their summary for a reach
of 2.6-5 m are 5-128 kg/acre).
One cannot compare these productivity data
with too much confidence since the samples
in both studies were relatively small, but it
seems safe to say that the heavier the snow
cover, the more difficult it is for caribou to
find and utilize ground lichens (e.g., Cladina
spp.), and the easier it is for them to reach and
utilize arboreal lichens (e.g., species of Alec-
toria and Bryoria ).
Caribou in northern Saskatchewan prefer
Bryoria species ("Alectoria jubata") to leaves,
according to Scotter (1962). In Ontario fallen
trees laden with Alectoria s. lat. are quickly
eaten bare of lichens by caribou (Cringan 1957).
Ahti (1959) reported that “Alectoria jubata”
ranks with Cladina mitis and C. rangiferina
as a preferred winter food eaten on a large
scale in Newfoundland; Alectoria sarmentosa
was only eaten in smaller quantities. In their
studies in Ontario, Ahti & Hepburn (1967)
describe “A. jubata”, Bryoria furcellata (sub
A. nidulifera) and B. simplicior as “highly
palatable.” Experiments carried out in Quebec
(DesMeules & Heyland 1969) demonstrated
OPERA BOT. 42 (1977)
Importance 45
that while Alectoria s. lat. (in a mixture with
Evernia and Usnea spp.) was not the favourite
food when compared with some Cladinae, it
was preferred to Cetraria islandica and
Stereocaulon spp. Apparently the terricolous
Alectoria ochroleuca is avoided by caribou,
according to papers cited by Ahti & Hepburn
(1967). Whether this is due to a quantitative
or qualitative difference in lichen substances
in this species as suggested by Ahti & Hepburn
(1967) requires investigation.
The food value of Alectoria s. lat. seems
to be rather high in comparison to that of other
lichens eaten by caribou, especially with regard
to protein. All lichens are high in carbohydrate
but are low in both fat and protein (Scotter
■ 965). Karev (1956) gives the protein and fat
content of “Alectoria implexa s. lat.” as
7.3% and 1.3%, respectively, compared with
2. 6-2. 9% protein and 1.8-2. 2% fat for the
Cladinae. Many Alectoriae s. lat., especially
A. sarmentosa, are digested very rapidly by
the caribou (Bergerud & Russell 1964) and so
are presumably utilized efficiently.
From the foregoing discussion, it is evident
that the majority of those working with the
brown “Alectoriae’’ (i.e., Bryoria) were
generally not certain as to the specific identity
°f their material. Although it would be im-
possible to ‘'translate" reports of “Alectoria
jubata ” into a list of known Bryoria species
in any precise way, our revision of the vouchers
for many of these studies has enabled us to
specify some of the species involved.
Ahti & Hepburn ( 1967) probably came closest
to defining the species they treated under “A.
jubata s. lat.” Based on identifications by
Motyka, they reported “A. americana”, “A.
canadensis” , “A. glabra ” and “A. tenerrima”
for northern Ontario. According to our exam-
inations of their material, Ahti & Hepburn were
working mainly with Bryoria trichodes subsp.
trichodes with smaller amounts of B.fuscescens
a nd B. lanestris. Scotter's material from
northern Saskatchewan (Scotter 1962, 1964.
■ 965) was a mixture of B. lanestris and B.
ximpUcior. The Newfoundland “A. jubata ”
reported by Ahti (1959) and Bergerud (1972)
ls mainly B. trichodes subsp. americana (Ahti &
Hawksworth 1974, sub A. americana). but some
Bryoria trichodes subsp. trichodes and perhaps
also B. fuscescens were probably also included.
The “Alectoria jubata” of Wells Gray Park
(Edwards et al. 1960) included “A. oregana ”,
“A. chalybeiformis" , and “A. jubata” accord-
ing to the authors themselves. Based on col-
lections made by Teuvo Ahti in the park in
1961, it would appear that their “A. jubata”
consisted of Bryoria pseudofuscescens, B.
fuscescens, B. glabra, and B. lanestris. Since
B. furcellata is virtually absent in the mountains,
their “A. chalybeiformis ” was probably a
spinulose form of B. fuscescens (such as f.
pallida). Since we have seen no specimens of
B. fremontii from that park, although the species
does occur in the Selkirk Mountains and in
Yoho National Park, it is likely that the
“ Alectoria fremontii ” referred to by Edwards
et al. (1960) was largely B. pseudofuscescens.
The “Alectoria spp.” used by DesMeules &
Hey land (1969) in Parc des Laurentides was
not segregated according to colour group,
and probably consisted largely of Bryoria
trichodes subsp. trichodes, B. nadvornikiana,
B. fuscescens, and Alectoria sarmentosa, to
judge from observations made in the park by
one of us (I.M.B.).
Food for man
In parts of interior British Columbia, some
tribes of Indians have long been using certain
species of Bryoria as a traditional food. Some
fascinating folk legends have been built up
around its use (e.g., “How the Coyote
Happened to Make the Black Moss Food”,
Dove 1933). Nancy Turner of the British
Columbia Provincial Museum reports (in press)
that members of the Lillooet tribe still use
Bryoria to make a sort of baked pancake
(Fig. 23), often eaten sweetened with sugar.
Richardson (1974) describes its preparation
in some detail based on the account by Dove
(1933).
Samples of the lichens used by the Indians
were sent to us by Turner and all are B.
fremontii. We have also seen herbarium speci-
mens off?, tortuosa collected by J. T. Rothrock
in ‘‘New Caledonia, northern British Columbia”
annotated by the collector as ‘‘used by Indians
and cattle for food in hard winters.” It is
very unlikely that ‘‘New Caledonia” is in the
northern part of the province in view of the
known distribution of the species. In any case,
46 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 23. A type of "pancake” made
from Bryoria fremontii by Lilloet
Indians in British Columbia. Scale
25 mm. Specimen from the B. C.
Provincial Museum; photo by R.
Turner.
the possible use of B. tortuosa as a food is
intriguing in view of the fact that it contains
vulpinic acid, a substance which appears to be
toxic (Llano 1948). Bryoria fremontii also can
contain vulpinic acid but only on apothecia or
in soralia, which are seldom present. If B.
tortuosa is actually capable of producing illness
in man, which has not been established, one
wonders if the limits of its use as a food
might follow the distribution limits of B.
tortuosa, since in southwestern British Co-
lumbia, the two species grow together and some
forms can easily be confused. Fortunately,
vulpinic acid is bitter and the Indians always
tasted samples of the lichen before collecting
large quantities for consumption (Turner in
press). They therefore probably never made
serious mistakes. The use of Bryoria by the
Interior British Columbia Indians will be
described in detail by Turner (in press).
Insect pests
Recently, a relationship between species of
Alectoria and Bryoria and the life cycle of an
extremely destructive insect pest has been
discovered. In studying outbreaks of the
Hemlock Looper ( Lambdina fiscellaria fiscel-
laria [Guen.]) in eastern Canada, Jobin (1973)
discovered that this small moth lays its eggs,
90% of the time, on pendent lichens hanging
from the branches of Abies balsamea. Almost
all these lichens are species of Alectoria or
Bryoria, including B. trichodes (both sub-
species), B. fuscescens, B. nadvornikiana, B.
glabra, and Alectoria sarmentosa. When the
larvae emerge from their eggs, they crawl
over the lichen to the branch, and then crawl
to the new leaves where they begin to feed.
The Hemlock Looper is so successful that in
times of outbreaks, it can completely defoliate
vast areas of fir forest. On lie Anticosti in
the Gulf of St. Lawrence, this insect destroyed
210,000 acres of timber, with a loss of 3,000,000
cords of wood, within the single year 1971.
Jobin (1973) found that the outbreaks centre
around islands, lake shores, or along rivers.
Whether the insect is responding to the micro-
climate characteristic of these localities or to
the presence of certain lichen species which
in turn require these habitats is still not known.
Nothing is known concerning any possible
specificity for certain species of Alectoria
or Bryoria, nor concerning its response to
certain lichen substances, colours, thallus
characteristics, branching patterns, age etc.
What role the lichen might play in the control
of the insect is under study, but wholesale
destruction of the lichen vegetation has been
ruled out (Jobin in litt.), at least at this stage.
It is interesting that the western Hemlock
Looper (Lambdina fiscellaria lugubrosa
OPERA BOT. 42 (1977)
Importance 47
[Hulst]) also lays its eggs on lichens (Thompson
1958). Thompson found that on Vancouver
Island, both “a fine hair-like black lichen”
(probably a species of Bryoria) as well as
“a pale yellow-green branching lichen” (the
latter possibly Alectorici sarmentosa or A.
vancouverensis or a mixture of the two)
harboured many eggs, although bark crevices
were still the preferred site. He also found that
along the coast, where lichens are largely
replaced by epiphytic mosses, the mosses are
the favoured substrate for oviposition. Thus,
in the case of this insect, specificity for lichen
species or even lichens in general seems to be
low.
Lichenometry
Pseudephebe minuscula and P. pubescens are
rapid colonizers of bare rock following the
retreat of glaciers and so have been used in the
lichenometric dating of glacial moraines. (For
a synopsis and extensive bibliography on this
topic see Webber & Andrews 1973.) They have
proved particularly valuable on Baffin Island
where isophyses (lines of equal growth) based
on them have been constructed (Andrews &
Webber 1964, 1969). Because of the taxonomic
problems involved in the separation of these
two species, Andrews & Webber (1964)
combined data from both and used the epithet
minuscula as that species was the more frequent
in the areas they studied. These authors cal-
culated an optimal annual growth rate of 0.52
mm and a mean of about 0.40 mm in diameter
assuming a 10 year initial colonization lag;
these data are consistent with the 0.56 mm
per annum rate recorded on Disko, Greenland,
by Beschel (1963). Tracings of individual
thalli on Baffin Island over the years 1963-67
revealed a tendency for larger thalli to grow
more quickly, but, over a period of up to
300 years, a mean annual growth rate of 0.40
mm appears to hold (Andrews & Webber
1969). A more recent study of P. minuscula
on Baffin Island, using photographic tech-
niques, showed that the growth rate of this
species decreases with increases in altitude
(Miller 1973).
Taxonomie survey
Key to the genera of Alectorioid lichens
1 Ascospores under 16 /nm long, colourless at maturity 2
- Ascospores over 19 p m long, yellowish to brown at maturity 3
2(1) Thallus closely adpressed to the substrate, attached over almost the whole thallus area,
rosette-forming, under 1 cm tall; cortex cellular in part, ± differentiated at the surface which
is smooth or rough; pycnidia common; lichen products absent 4 . Pseudephebe
Thallus eiect, caespitose to pendent, attached at the base or sometimes secondarily by
branches but never over the whole thallus area, not forming rosettes, over 1 cm tall; cortex not
distinctly cellular, not or little differentiated at the surface; pycnidia rare; atranorin and/or
chloroatranorm, oremol tridepsides, /3-orcinol depsidones and pulvinic acid derivatives
known Bryoria
3(1) Ascospores becoming transversely septate or muriform at maturity ' 4
- Ascospores remaining simple at maturity, 2-4 per ascus, 2(M5 pm long; cortex with much
matrix, not differentiated but often disintegrating at the surface; medullary hyphae usually
ornamented; usme acid, and orcinol depsides known 1 Alectoria
4(3) Ascospores I (-3 (-septate at maturity, 22^10 pm long. 6-S per ascus;' 'cyphella-like
perforations absent; pseudocyphellae present as a single continuous longitudinal furrow;
atranorin, /3-orcinol depsidones and pulvinic acid derivatives known 5 Sulcaria
- Ascospores muriform at maturity, 76-96 pm long, 1 per ascus; cyphella-like perforations
present, pseudocyphellae absent; /3-orcinol depsides (inch atranorin) and depsidones, as
well as hydroxyanthraquinones known 3 0ropogonTh Fr
Check-list of the North American species
Accepted taxa
Selected synonyms
Alectoria Ach.
1. A. fallacina Mot.
2. A. imshaugii Brodo & D. Hawksw.
3. A. lata (Tayl.) Linds.
4. A. mexicana Brodo & D. Hawksw.
5. A. nigricans (Ach.)Nyl.
6. A. ochroleuca (Hoffm.) Massai.
7a. A. sarmentosa (Ach.) Ach. subsp. sarmentosa
7b. A. sarmentosa subsp. vexillifera (Nyl.)D. Hawksw.
8. A. vancouverensis (Gyeln.)Gyeln. ex Brodo & D.
Hawksw.
Bryopogon Link
A. osteina Nyl.
A. stigmata Bystr.,T. subsarmentosa Stirt.
A. vexillifera (Nyl.) Stiz.
OPERA BOT. 42 (1977)
Taxonomie survey 49
Accepted taxa
Bryoria Brodo & D. Hawksw.
sect. Bryoria
1 . B. chalybeiformis (L.) Brodo & D. Hawksw.
2. B. fuscescens (Gyeln.) Brodo &D. Hawksw.
3. B. glabra (Mot.) Brodo & D. Hawksw.
4. B. lanestris (Ach.) Brodo & D. Hawksw.
5. B. subcana (Nyl. ex Stiz.) Brodo & D. Hawksw.
6a. B. trichodes (Michx.) Brodo & D. Hawksw.
subsp. trichodes
6b. B. trichodes subsp. americana (Mot.) Brodo &
D. Hawksw.
7. B. vrangiana (Gyeln.) Brodo & D. Hawksw.
sect. Divaricatac (DR.) Brodo & D. Hawksw.
8. B. bicolor (Ehrh.) Brodo & D. Hawksw.
9. B. carlottae Brodo & D. Hawksw.
10. B. cervinula Mot. ex Brodo & D. Hawksw.
1 1. B. furcellata (Fr. ) Brodo & D. Hawksw.
12. B. nitidula (Th. Fr.) Brodo & D. Hawksw.
■ 3. B. simpiicior (Vain.) Brodo & D. Hawksw.
14. B. tenuis (Dahl) Brodo & D. Hawksw.
sect. Implexae (Gyeln.) Brodo & D. Hawksw.
13. B. capillaris (Ach.) Brodo & D. Hawksw.
16. B.friabilis Brodo & D. Hawksw.
17. B. implexa (Hoffm.) Brodo & D. Hawksw.
18. B. nadvornikiana (Gyeln.) Brodo & D. Hawksw.
10. B. pikei Brodo & D. Hawksw.
20. B. pseudocapiiiaris Brodo & D. Hawksw.
21 B. pseudofuscescens (Gyeln.) Brodo &D.
Hawksw.
22. B. satazinica Brodo & D. Hawksw.
23. B. spiralifera Brodo & D. Hawksw.
sect. Suhdivergentes (Mot.) Brodo & D. Hawksw.
24. B. abbreviata (Müll. Arg.) Brodo & D. Hawksw.
25. B. oregana (Tuck.) Brodo & D. Hawksw.
sect. Tortuosae (Mot.) Brodo & D. Hawksw.
26. B. fremontii (Tuck.) Brodo & D. Hawksw.
27. B. tortuosa (Merr. ) Brodo & D. Hawksw.
Pseudephebe Choisy
1- P ■ minuscula (Nyl. ex Arnold) Brodo &
D. Hawksw.
2- P . pubescens (L.) Choisy
Sulcaria Bystr.
1- 5. badia Brodo & D. Hawksw.
Selected synonyms
Setaria Michx., Bryopogon Th. Fr., non Link
A. positiva (Gyeln.) Mot.
A. tenerrima Mot.
A. haynaldii Gyeln.
A. canadensis Mot., A. deiicata Mot.
A. americana Mot., A. ambigua Mot.,
A. sepiacea Mot.
A . nidulifera N orrl .
A. lanea auct., non (Ehrh. ex Hoffm.) Vain.,
A. irvingii Llano
A. nana Mot.
A. bicolor var. subbicolor Mot.
A. implexa auct., non (Hoffm.) Nyl. p.p.
A. norstictica Mot., A. subtilis Mot.
A. oregana auct. p.p.
A. virens sensu R. Howe, nonTayl.
Parmelia minuscula (Nyl. ex Arnold) Nyl.
Parmeiia pubescens (L.) Vain.
Key to the North American species
1 Thallus yellow or yellowish-green; conspicuous pseudocyphellae present (occasional in
Ramalina thrausta, couplet 7); cortex KC + yellow (usnic acid) 2
' Opera Botanica nr 42
50 Alectoria in North America
OPERA BOT. 42 (1977)
- Thallus not yellow or yellowish-green; pseudocyphellate or not; cortex KC- or KC+ reddish
(usnic acid absent) 10
2( 1) Medulla C+ red (olivetoric acid) 3
- Medulla C— (olivetoric acid absent) 4
3(2) Thallus subpendent, bright yellow throughout; cortex clearly delimited, not becoming
fibrous internally; known only from the highlands of Mexico and Guatemala .... 4. A. mexicana
- Thallus pendent, often very long, yellow to greyish-green; cortex joined with a fibrous
network within the medulla; medulla dense and compact; southern California to British
Columbia close to sea level 8. A . vancouverensis
4(2) Thallus caespitose to subpendent or pendent, usually corticolous 5
- Thallus erect to prostrate, usually terricolous 9
5(4) Thallus caespitose, rarely subpendent, usually less than 10 cm long 6
- Thallus pendent to subpendent, 8-20 cm long or longer 7
6(5) Isidioid spinules arising from the pseudocyphellae in tufts or along the branches; ascocarps
rare; medulla PD+ orange-yellow or PD— (thamnolic or squamatic acid present), medulla
KC— , exceptionally KC + red (alectoronic acid very rare); Idaho, west to B.C. coast, south
to central California; locally frequent 2. A. imshaugii
- Isidioid spinules absent; ascocarps not uncommon; medulla KC+ red (alectoronic acid);
from Washington southwards in mountains to Guatemala 3. A. lata
7(5) Apices of branches sorediate or granular, often hooked; medulla KC -
Ramalina thrausta (Ach.) Nyl.
- Apices of branches not sorediate or granular, never hooked; medulla KC + red or KC - ... 8
8(7) Branches more or less regular, pendent; medulla usually lax, KC + red or rarely KC-;
cortex relatively uniform throughout; a very common boreal-montane species
7 a. A. sarmentosa subsp. sarmentosa
- Branches recurved and tangled, subpendent; medulla dense, KC - or rarely KC + red; cortex
very uneven in thickness, giving thallus a knobby appearance; Appalachian Mountains,
Tennessee to Nova Scotia; rare 1. A. fallacina
9(4) Thallus usually erect, rarely subdecumbent; main stems more or less terete, usually less
than 2 mm diam.; medulla KC - , CK+ golden-yellow (diffractaic acid), or rarely KC +
red (alectoronic acid) especially in the Aleutians and in Central America; a frequent to abun-
dant species in the tundra 6. A. ochroleuca
- Thallus prostrate to decumbent; main stems becoming dorsiventrally compressed, expanded
and foveolate, usually exceeding 2 mm diam. (to 4 cm wide); medulla KC + red (alectoronic
acid) or intrequently KC — ; an occasional subspecies of western alpine and eastern arctic
tundra 7b. A. sarmentosa subsp. vexillifera
10(1) Thallus closely adpressed to rock, usually less than 1 cm tall, forming rosettes attached
over most parts of the thallus; medulla K-, C-, KC-, PD- 11
- Thallus not closely adpressed to rock, usually exceeding 1 cm in height or length, not
forming rosettes attached over most parts of the thallus; medulla PD 4- or PD- 12
1 1(10) Branches usually terete, even; internodes long; adpressed only at the margins, becoming
bushy, prostrate or erect towards the centre of the thallus; arctic-alpine 2.P. puhescens
- Branches becoming dorsiventrally compressed, uneven; internodes short; closely attached to
the rock over almost the whole thallus area, becoming closely adpressed to almost crustose or
foliose towards the centre of the thallus; arctic-alpine \. P. minuscula
12(10) Thallus erect or prostrate to decumbent; on rocks or soil (exceptionally on bark or
wood) 13
- Thallus caespitose or pendent; on bark or wood (rarely on rock) 26
13(12) Thallus PD + yellow-orange (diffusing onto filter paper), surface C + pink to red on pale
areas (alectorialic and/or barbatolic acid); thallus some shade of grey, white, yellowish grey
or black; branching anisotomic dichotomous 14
- Thallus PD — or PD + red 15
OPERA BOT. 42 (1977)
Taxonomie survey 5 1
14( 13) Pseudocyphellae white, raised and conspicuous, soralia absent; thallus usually pale at the
base with at least the tips blackened; apices usually reflexed; common on the ground in
arctic-alpine areas (very rarely on bark or wood) 5 .A. nigricans
- Pseudocyphellae small, dark, plane, generally inconspicuous; soralia usually present; thallus
either pale at tips or entirely dark brown to black, at least black at the base; apices pointed
and straight, with abundant, divergent, spinulose lateral branches (see also couplet
28) 18. B. nadvornikiana
15(13) Medulla or only pseudocyphellae C + red or pink (gyrophoric acid); thallus bright to dark
red-brown, usually shiny, concolourous; branching isotomic dichotomous, broad, divergent;
subarctic to arctic-alpine; common Cornicularia divergeas Ach.
- Medulla and pseudocyphellae C — 16
16(15) Thallus with conspicuous, broadly fusiform, cyphella-like perforations into the hollow
medulla; grey to dark red-brown; known from the high mountains of Central
America Oropogon loxensis (Fée) Th. Fr. s. lat.
- Thallus without cyphella-like perforations 17
17(16) Soralia present (sometimes sparse) 18
- Soralia absent 21
18(17) Soralia bearing tufts of isidioid spinules; PD + red; branching regularly isotomic
dichotomous; branches smooth, not twisted or foveolate, shiny, pale to cervine brown;
occasionally on rocks but also known from submaritime heathland in Newfound-
land 11. B. furcellata
~ Soralia not bearing tufts of isidioid spinules; branching often irregular 19
19(18) At least main branches very thick, twisted and often foveolate, generally over
0.5 mm diam 20
- Branches very slender, pendent or subpendent, usually less than 0.2 mm diam.; soralia
numerous, virtually all fissurai 4. B. lanestris
20( 19) Thallus dark brown to olivaceous black, often shiny: soralia sparse, usually tuberculate,
PD + red; medulla and cortex usually PD - ; arctic-alpine 1 . B. chalybeiformis
~ Thallus fuscous brown to brown, usually matt; soralia frequent or sparse, fissurai
and tuberculate types usually present, PD+ red; cortex often PD + red; usually
coastal 2. B. fuscescens var. positiva
21(17) Thallus uniformly red-brown, small, prostrate, under 3 cm long; main branches
conspicuously longitudinally foveolate and flattened; medulla PD-; ascocarps to 2.5 mm
diam., red-brown; known with certainty only from tundra in southwest Greenland
B. subdivergens (Dahl) Brodo & D. Hawksw. (p. 153)
~ Thallus not red-brown and foveolate 22
22(21) Main branches shiny, black, with numerous grey to pale brown perpendicular side
branches and apices; spinules abundant; medulla PD + red at least in parts 23
~ Main branches not black with pale side branches and apices; thallus more or less uniformly
cervine brown to dark brown or black; medulla PD + red or PD - 24
23(22) Thallus forming dense tufts; third order branches common and arising at right
angles; coastal or montane 8. B. bicolor
~ Thallus not forming dense tufts; third order branches sparse and usually arising at acute
angles; mainly coastal and montane, very rare in the tundra 14. B. tenuis
24(22) Thallus erect or frequently decumbent, usually forming compact tufts, uniformly dark
cervine brown to black; branching anisotomic dichotomous with lateral branches arising
perpendicularly or at acute angles: medulla PD + red (exceptionally PD — ); mainly restricted
to the tundra where it is often common 12. B. nitidula
~ Thallus prostrate to decumbent, not forming compact tufts, cervine brown to dark brown,
never uniformly brown-black 25
25(24) Branching anisotomic dichotomous; conspicuous, stout, lateral spinules present; pale
52 Alectoria in North America
OPERA BOT. 42 (1977)
fuscous brown to dark brown, or blackened at the base, often appearing variegated; medulla
PD+ red; pseudocyphellae absent to sparse; ascocarps unknown; known only from Alaska
and coastal British Columbia 10. B. cervinula
- Branching isotomic dichotomous; lateral spinules, if present, not stout; uniformly cervine
brown, often shiny; medulla PD + red or PD-; ascocarps not infrequent; pseudocyphellae
often abundant; mainly coastal 6b. B. trichodes subsp. americana
26(12) Thallus K+ persistently bright yellow, KC + red (fading) and often C + pink (fading),
PD + orange-yellow (barbatolic and/or alectorialic acid) 27
- Thallus K. — , or if K+ yellow, the yellow is very pale or quickly darkens to brown, orange
or red and largely disappears; PD + or — 30
27(26) Thallus caespitose to subpendent, often with perpendicular lateral spinulose branches . . 28
- Thallus pendent, without perpendicular lateral spinulose branches; soralia unknown in
North American material; basal branches rarely blackened, apices usually concolourous or
darker than the basal parts, thallus greyish to fuscous brown or rarely dark brown; widely
distributed in Canada and the northern United States (if K+ reaction weak, see B. pikei,
couplet 60) 15. B. capillaris
28(27) Soralia present; basal branches blackened, apical branches greyish to fuscous brown;
pseudocyphellae short, fusiform; boreal forests in Canada and Alaska 18. B. nadvornikiana
- Soralia absent 29
29(28) Tips of branches and sometimes most of thallus blackened; pseudocyphellae white,
short, raised; thallus much branched, caespitose (see couplet 14) 5. A. nigricans
- Basal branches and tips concolourous with main branches, not blackened; thallus grey to dull
badia-brown, especially on lateral branches; pseudocyphellae elongate, linear, becoming
canaliculate; thallus tangled, subpendent; on trees along sea coast, northern California to
Oregon 20. B. pseudocapillaris
30(26) Thallus with soredia 31
- Thallus without soredia 40
31(30) Thallus caespitose, pale to dark cervine brown, usually shiny 32
- Thallus subpendent to pendent 33
32(31) Soralia white, bearing tufts of isidioid spinules; inner cortex, medulla and soredia PD +
red (fumarprotocetraric acid); widely distributed in eastern temperate and boreal regions,
infrequent in the west 1 1 . B. furcellata
- Soralia greenish black, not bearing tufts of isidioid spinules; all tissues PD — ; common in
northern boreal forests \7>.B. simplicior
33(31) Soralia bright yellow or greenish yellow (vulpinic acid), tuberculate; medulla and soralia
PD - (see couplet 51) 26. B. fremontii
- Soralia white, sometimes flecked with black (not bright yellow or greenish yellow), PD + red
(fumarprotocetraric acid), tuberculate or fissurai; cortex or medulla PD - or PD + red 34
34(33) Pseudocyphellae conspicuous; branches usually uneven in diameter, cervine brown
throughout; inner cortex PD + red or PD — a morphotype of B. trichodes
- Pseudocyphellae absent or very inconspicuous; branches even or uneven in diameter;
colour various; cortex or medulla PD + red or PD- 35
35(34) Main blanches (not base) usually less than 0.2 mm diam.; thallus dark brown to dark
olivaceous or black throughout; soralia fissurai or very rarely tuberculate, often speckled with
black; branches very uneven in diameter, brittle, tending to fragment in herbarium packets;
cortex and medulla PD -; common in the northern boreal forests, extending southwards in
the western mountains B i anestr is
- Main branches (not base) exceeding 0.2 mm diam.; thallus brown or olivaceous, but rarely
darkening to black; soralia almost always uniformly white 36
36(35) Outer and sometimes inner cortex of branches PD + intense red (fumarprotocetraric
acid) ^7
- Medulla and cortex of branches PD- 38
OPERA BOT. 42 (1977)
Taxonomie survey 53
37(36) Thallus uniformly grey, sometimes becoming pale fuscous brown towards the apices (not
pale at the base); soralia tuberculate; west coast, very rare 5. B. subcana
- Thallus fuscous brown to brown, usually pale at the base; fissurai and tuberculate soralia
usually present; scattered throughout Canada and the northern U.S 2. B.fuscescens
38(36) Branches even in diameter throughout; thallus shiny, olivaceous to olive-brown,
concolourous ; angles between the main dichotomies obtuse and rounded; soialia fissurai,
rarely tuberculate; mainly in the west where it is locally abundant; very rare on the east
coast 3- B. glabra
- Branches uneven in diameter, at least in part, or if even, not olivaceous; angles between the
main dichotomies, if obtuse, then generally not rounded ; soralia tuberculate or fissurai 39
39(38) Thallus shiny olivaceous brown to olive black, concolourous; branching irregular
anisotomic dichotomous especially towards the apices; main branches becoming contorted
and foveolate, often becoming spinulose; soralia rare 1 . B. vrangiana
- Thallus usually dull, fuscous brown to dark brown, usually paler at the base; branching
isotomic dichotomous; main branches rarely becoming contorted and foveolate; soralia
usually abundant, usually fissurai and tuberculate 2 .B.fuscescens
40(30) Thallus with conspicuous broadly fusiform cyphella-like perforations opening into the
hollow medulla; grey to dark red-brown, shiny (see couplet 16)
Oropogon bxensis (Fée)Th. Fr, s. lat.
- Thallus without cyphella-like perforations 41
41(40) Main branches shiny, black, with numerous, obviously paler, olivaceous grey to light
brown, perpendicular side branches and apices; spinules abundant; inner cortex and medulla
PD + red at least in apical parts; mainly coastal and montane 23
- Main branches not black with pale olivaceous perpendicular side branches and apices;
cortex and/or medulla PD + red or PD — 42
42(41)Thallus caespitose to erect, very small, usually less than 3 cm tall; branches longitudinally
furrowed, dull and flattened; almost always abundantly fertile 43
- Thallus pendent or subpendent, usually exceeding 3 cm long 44
43(42) Thallus dull red-brown; epithecium red-brown, K-; mountainous areas from the
Rockies westwards 24 B - abbreviata
- Thallus olive brown to greenish black; epithecium greenish to brown black, usually K +
violet; California to the Queen Charlotte Islands .Cornicularia californica ( I uck. ) DR.
44(42) Scattered elongate yellow pseudocypheilae present; thallus dark red-brown to dusky
yellow-brown, occasionally becoming bright greenish yellow in parts; long, pendent; main
branches becoming twisted and foveolate; thallus PD — or faintly PD + yellow on filtei paper,
mainly along the west coast from California to southern British Columbia -7. B. tortuosa
- Pseudocypheilae present or absent, but if present, white or brown (never yellow) 45
45(44) Branches with very long (at least 4 mm) deep furrows or grooves (sulci) breaking into
the medulla; thallus dull chestnut brown to almost yellowish brown in parts; outer cortex
K + faintly yellow (at least on filter paper). PD + brownish or PD- (or pale yellow on
filter paper) (atranorin present); California and Oregon 1-5. badia
- Branches without very long, deep furrows or grooves breaking into the medulla 46
46(45) Cortex and medulla PD - (and not producing any colouration on filter paper) 47
- Cortex and/or medulla PD + yellow or red (or producing a colouration on filter paper) 52
47(46) Medulla and/or pseudocypheilae C+ pink to red, KC + pink (olivetoric acid), thallus
red-brown, with punctiform, almost tuberculate, white pseudocypheilae, branching mostly
isotomic dichotomous; infrequent, along B.C. coast Cornicularia pseudosatoana Asah.
- Medulla C - , KC - ; cortex C - or C + pink (gy rophoric acid) 48
48(47) Cortex C+ pink, KC+ pink (gyrophoric acid); branches very brittle; thallus pale
greyish brown, or rarely pale olive-brown; usually very uneven in diameter and twisted,
infrequent on the west coast, and rare in the western mountains and along the east
16. B. friabilis
coast
54 Alectoria in North America
OPERA BOT. 42 (1977)
- Cortex C , KC ; thallus usually dark red-brown or yellowish brown to dark brown or
olive-black 49
49(48) Thallus shiny olivaceous brown to olive-black; branching anisotomic dichotomous
especially towards the apices; main branches becoming contorted and foveolate, often
spinulose (see couplet 39) 7 . b. vrangiana
- Thallus dull or shiny, yellow-brown to red-brown or dark red-brown 50
50(49) Main branches even in diameter, not foveolate or channelled, cervine brown throughout
(see couplet 55) 6b. B. trichodes subsp. americana
- Main branches foveolate or channelled 51
51(50) Thallus very dull, distinctly red-brown; branches foveolate or longitudinally ridged,
channelled but not twisted, very brittle, to 0.3 mm diam.; branching irregularly anisotomic
dichotomous; ascocarps, if present, red-brown, epruinose, sometimes “ciliate" at margins;
hyphae at the surface of the cortex consisting of short, knobby hyphal cells (jigsaw-like
in surface view, see Fig. 12 A); in the western mountains 25. B. oregana
- Thallus shiny or less frequently dull, yellow-brown to red-brown or dark red-brown; main
branches toveolate, twisted, not brittle, mainly 0.4-1. 5 mm diam.; branching isotomic
dichotomous at the base, with slender perpendicular lateral branches; ascocarps, if present,
yellow pruinose, not ciliate; cortex in surface view consisting of long, slender, parallel
hyphae; mainly along the west coast 26. ß. fremontii
52(46) Cortex and/or medulla PD + red (fumarprotocetraric acid) ! 53
- Cortex and/or medulla PD + yellow (at least on filter paper) 56
53(52) Thallus olive; branching mostly anisotomic dichotomous to submonopodial, with many
short perpendicular side branches along the main axes; pseudocyphellae abundant; PD + red
in inner cortex only (not diffusing onto filter paper); rare, known only from the Queen
Charlotte Islands 9. B. carlottae
- Thallus pale greyish brown to cervine brown (not olive) 54
54(53) Thallus pale grey to greyish brown; outer cortex PD + intensely red (diffusing out onto
filter paper); rare (see couplet 38) 5 ß subcana
- Thallus pale to dark cervine brown; inner cortex and sometimes medulla PD + red (rarely
producing a red colouration of filter paper) 55
55(54) Branches even in diameter throughout; pseudocyphellae usually sparse, dark and
depressed; thallus usually dark; mainly along east and west coasts
."T b. B. trichodes subsp. americana
- Branches very uneven in diameter; pseudocyphellae usually abundant, white and raised;
thallus often pale; mainly with an Appalachian-Great Lakes distribution, but also on the
6a. ß. trichodes subsp. trichodes
56(5..) Thallus dull red-brown or badious, often becoming paler in parts; pseudocyphellae
linear, white, often exceeding 4 mm long and spiraling around the main branches;
perpendicular side branches frequent; contains abundant norstictic acid; known only from
coastal northern California 23 . B. spiralifera
- Thallus grey to pale olive-brown or pale to dark brown to black, not red-brown or badious;
pseudocyphellae never exceeding 1.5 mm in length; perpendicular side branches absent or
very sparse, mainly north of California 57
57(56) Thallus dark olive-brown to black, often shiny; pseudocyphellae fusiform, white or dark!
to 1 mm long, sometimes sparse; contains norstictic acid; common in Rocky and Selkirk
Mountains southward into Oregon 2 1 . ß. pseudofuscescens
hallus pale olive-brown to olive-grey, often variegated; pseudocyphellae abundant and
obvious
58(57) Norstictic acid present Infrequent morphotype of B. pseudofuscescens
- Norstictic acid absent 59
59(58) Salazimc acid present; thallus pale olive to olive-brown, loosely branched and entangled-
rare, known only from the northeast coast 22. ß. salazinica
OPERA BOT. 42 (1977)
Taxonomie survey 55
- Salazinic acid absent
60(59) Thallus generally smooth, even, shiny, olive to olive-grey or pale olive-brown; cortex
sometimes KC + pink (contains alectorialic acid); infrequent along the west coast and rare
on the east coast 19. B. pikei
- Thallus generally uneven, dull, twisted, pale fuscous to pale olive-brown; cortex always
KC- (contains psoromic acid); rare, scattered throughout the northern boreal forest
17. B. implexa
region
Notes on the presentation of the species
(1) In the case of taxa for which a detailed account
of the synonymy has been presented elsewhere by one
of us (D.L.H.), the lists of synonyms included in
this section have been limited to names based on
North American collections. The identity of unlisted
synonyms not included here, which have appeared in
the North American literature, can be traced through
the Index, and full particulars of them located
through the literature references appearing under the
accepted name. Where a detailed account of the
taxon has not been provided elsewhere, an attempt
has been made to determine the full synonymy
(i.e., including names not based on North American
material).
(2) Descriptions are based only on North American
material examined in the course of this study. Any
major departures from previously published descrip-
tions are mentioned under "Discussion .
(3) The observations on "Ecology" similarly refer
only to the autecology of the species in North
America unless otherwise indicated.
(4) In the course of the present survey we have
not endeavoured to trace collections supporting
literature records except in the case of a few extremely
rare species and some species reported as important
to man or caribou. Comments on known and suspected
erroneous reports in the literature have been confined
to ones from well outside the range of the species.
(5) In the case of species known from a relatively
small number of localities, all collections from
North America examined during this study are
listed under “Specimens". Where species are ex-
tremely common, because of limitations of space, it
has not proved possible to cite all those studied
and only "Selected specimens" are listed. This selec-
tion has been designed to cite at least one specimen
from each state or province. A full listing of the
specimens examined (with abbreviated localities)
is available at a nominal charge, from the Depository
of Unpublished Data, CIST1. National Research
Council of Canada. Ottawa. Ontario K 1 A OS2,
Canada. Individual record cards prepared in the course
of the production of the distribution maps are pre-
served in the Botany Division of the Museum of
Natural Sciences, National Museums of Canada
(CANL) in Ottawa and are available for consultation
by those working on the floristics of a particular
region.
Alectoria Ach.
in Luyken, Tent. Hist. Lieh.: 95 (1809). - Evernia
sect. Alectoria (Ach.) Fr., Syst. Orb. 1: 237 (1825). -
Cornicularia sect. Alectoria (Ach.) Duby, Bot.
Gall. 2: 616 (1830). - Parmelia sect. Alectoria
(Ach.) Spreng., Fl. Halen., ed. 2, 2: 521 (1832). -
Bryopogon Link, Grund. Kraut. 3: 164 (1833);
nom. illegit. (Art. 63). - Alectoria subgen . Eualectoria
Th. Fr., Lieh. Scand. I: 19 (1871); nom. illegit.
(Art. 63). - Eualectoria Gyeln., Ann. Mus. Nat.
Hung. 28: 283 (1934); nom. inval. (Art. 34), nom.
illegit. (Art. 63). — Alectoria sect. Phaeosporae Hue,
Nouv. Arch. Mus. Paris, Sér. 4, 1: 93 (1899); nom.
illegit. (Art. 63). - Alectoriomyces Cif. & Tom..
Atti 1st. Bot. Lab. Critt. Pavia, Ser. 5. 10: 44, 70
(1953); nom. illegit. (Art. 63).
Ceratocladia Del., in Schwendener, in Naegeli,
Beitr. Bot. 2: 149 (1860); nom. inval. (Art. 34);
type: C. luteola (Mont, ex de Not.) Schwend.
L -Alectoria sarmentosa (Ach.) Ach.], holotype.
Alectoria sect. Ochroleuca Gyeln., Feddes Repert.
38: 243 (1935); type: A. ochroleuca (Hoffm.) Massai.,
holotype.
Non Alectoria Link, Grund. Kraut. 3: 163 (1833);
nom. illegit. (Arts. 63. 64); type: A. articulata (L.)
Link L= Usnea articulata (L.) Hoffm.], holotype.
Type species: Alectoria sarmentosa (Ach.) Ach.
(syn. Lichen sarmentosus Ach.; lectotype designated
by Clements & Shear 1931 p. 322).
Thallus fruticose, erect, caespitose, decumbent,
subpendent or pendent; branching variable,
branches generally terete but becoming com-
pressed and angular to foveolate near the
base and axils, in a few taxa becoming markedly
expanded and dorsiventrally compressed;
greenish-yellow to yellow in most species,
fuscous black to black and pinkish below in
one.
True lateral spinules absent, spinules arising
through the pseudocyphellae in some species.
Isidia absent. Soralia large and tuberculate.
very rare, unknown in most species. Pseudo-
cyphellae always present, abundant, conspic-
uous, fusiform, white, markedly raised, some-
times becoming sorediate and bearing spinules.
Cortex composed of periclinal, arachnoid, con-
glutinate hyphae, the hyphae immersed in a
relatively large amount of matrix, not disin-
tegrating and not differentiated at the surface.
Medullary hyphae usually ornamented.
Apothecia lateral, sometimes appearing ge-
niculate owing to the death of the branch
distal to them, rare and unknown in some
species, common in others; excipulum thalli-
num concolourous with the thallus, persistent
and often markedly incurved, not ciliate; disc
brown to dark brown or black. Asci clavate,
thick-walled, arrested bitunicate, 2-4-spored.
Ascospores ellipsoid, with a distinct hyaline
epispore, brown at maturity, simple, 20-45
/am long. Pycnidia usually rare.
Usnic acid present in all but one species,
usually abundant; atranorin and chloroatranorin
absent; orcinol depsides abundant; /3-orcinol
depsides (excluding atranorin and chloroatra-
norin) frequent; orcinol tridepsides. /3-orcinol
depsidones, pulvinic acid derivatives and
hydroxyanthraquinones absent. An unidenti-
tied K + red, C + green-black compound occurs
in the cortex of the basal parts of the thalli
of most species.
Discussion
As interpreted here Alectoria comprises eight
species all of which are known from North
America. The genus shows its greatest diversity
in western North America where all but one
species (A. fallacina) occur. Species of this
genus are widely distributed throughout the
world and occur in both hemispheres; three
have bipolar tendencies and three are circum-
boreal and widely distributed in the Northern
Hemisphere.
Alectoria is readily separable from the other
genera accepted here, even in the absence of
OPERA BOT. 42 (1977)
Ale ct or ia Ach. 57
apothecia, by virtue of the characteristic
chemistry, cortical structure and pseudo-
cyphellae.
The nomenclature and typification of Alec-
toria are discussed under Bryoria.
I . Alectoria fallacina Mot.
Fragm. Florist. Geobot. 6: 447 (1960); type: U.S.A.,
Tennessee, Great Smoky Mountains, near Alum
Cave, on Picea rubra, alt. 1575 m, G. Degelius,
13 September 1939 (US holotype! herb. Degelius,
US isotypes!).
Thallus subpendent to almost caespitose, rarely
pendent, having a tangled appearance, 6-8(-20)
cm long; branching variable, broad and obtuse
to narrowly acute, with perpendicular lateral
branches not uncommon, basically isotomic
dichotomous; branches uneven in thickness,
knobby due to large differences in cortical
thickness, usually 0.2-0. 4 mm diam.; grey-
green to greenish yellow, rarely yellow; medulla
dense, filling medullary cavity; cortex becoming
extremely thick in places, commonly up to
120 /xm and reaching 160 /am in thickness.
True lateral spinules absent; soralia absent,
or very rarely present (see discussion below).
Pseudocyphellae punctiform to short fusiform,
often inconspicuous, commonly raised, white,
without spinules.
Apothecia not seen. Pycnidia common,
usually in raised, darkened tubercles; conidia
bacillariform, c. 2.0 x 0.8 /urn.
Cortex K - , C - . KC + yellow, PD - ; medulla
K-, KC - or KC+ red. C— , PD-. Contains
usnic acid, and sometimes alectoronic acid.
Plus an unknown substance which is barely
visible in thin layer chromatography. The
unknown is not visible in UV light and produces
only a light yellow colour after development
with H 2 S0 4 . It has relative positions of 1.
T 3, in the standard solvents (see C. Culberson
1972).
Ecology. On Picea rubens, Abies fraseri, and
Quercus spp. on mountain trails and along the
seashore.
Distribution. Along the Appalachian mountains
(Fig. 24A). It might well be considered as a
vicariad of A. sarmentosa, having speciated
in the southern Appalachians. The two species
ar e almost entirely allopatric.
The report of A. fallacina from Alaska (Krog
1968) is erroneous being based on a specimen
of A. sarmentosa.
Discussion
This Appalachian population, very closely
related to A. sarmentosa, is being tentatively
recognized at the species level following Motyka
(1960), but a study of more material may
prove it to be more properly ranked as a
subspecies. Its knobby, uneven appearance,
sparse, punctiform pseudocyphellae, con-
spicuous raised pycnidial tuberculae, variable
but generally very thick cortex, dense medulla,
grey-green colour and tangled habit all serve
to distinguish it from A. sarmentosa. Specimens
showing all these characters most distinctly
are invariably deficient in alectoronic acid
(i.e., medulla KC-). Specimens which are
KC + red, including the holotype, are some-
what yellower, and their cortices are less
variable and slightly thinner; the other distin-
guishing characters usually remain. The pres-
ence of a few soralia on a single specimen
from Tennessee (litis & Sharp, 1942 [CANL])
is not regarded as taxonomically important.
Although the holotype is entirely KC + red,
the isotypes are either KC — (US) or a mixture
of KC - and KC + red plants (herb. Degelius).
The description of A. fallacina as given by
Motyka (1964) agrees with our observations in
most points (e.g.. the irregular appearance,
the presence of “swellings", the indistinct
pseudocyphellae). However, Motyka surpris-
ingly found the cortex to be exceptionally
thin ("only 35 jim thick”).
Motyka (1964) compared this species with
Alectoria pellucida Mot., a lichen first de-
scribed from Bulgaria, and which Bystrek placed
in Bryopogon. Motyka (1962) and Bystrek (1971)
placed B. pellucidus in its own section
(Pellucidae) within the subgenus and genus
Bryopogon, respectively. Later, Bystrek (1974)
again referred it to the genus Alectoria, com-
paring it this time with A. stigmata Bystr.,
which appears to be a morphotype of A.
sarmentosa (see Discussion under A. sarmen-
tosa subsp. sarmentosa). Bystrek, in the latter
paper, mentioned that the soralia of A. pellucida
are PD + red, a very unusual characteristic for
an Alectoria s. str. and one which persuaded
58 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 24. A: Alectoria fallacina. Known distribution. - B: A. imshaugii. Known distribution. - C: A. lata. N.
American distribution. A uncertain locality.
OPERA BOT. 42 (1977)
Alectoria Ach. 59
us to request the type from Lublin thinking
that the name might refer to one of our
thamnolic-containing Alectoriae. Bystrek very
kindly sent us a Polish specimen named as
A. pellucida (Montes Tatry, Wotoszyn Mt.,
ad ramos Picea in Piceeto primitivo, J. Bystrek,
12 June 1964 [LBL]), which he said was
identical with the holotype of that species. This
specimen is clearly a Bryoria, with conspicuous
fissurai pseudocyphellae and very sparse
fissurai soralia. The thallus is entirely devoid
of cortical pigments leaving it pale greenish and
translucent, as described by Bystrek (1971).
A chemical analysis of the thallus by TLC
showed it to have trace amounts of psoromic
acid. Thus it would seem that at least this
specimen is very close to, if not conspecific
with, Bryoria osteola, as suggested by Hawks-
worth (1972 p. 250). The soralia of this thallus
give a PD + yellow rather than PD + red
reaction, as reported by Bystrek, and so the
holotype should be reexamined by TLC.
Specimens
Canada. New Brunswick. Charlotte County: Grand
Minan (sic) (Manan), Willey s.n.. 1879 (US).
U.S.A. New Hampshire. Coos County: Mt.
Washington. Burkholder 43 (CANL) - North
Carolina. Swain County: Great Smoky Mountains,
Mt. Kephart, Degelius s.n.. 15 Sept. 1939 (DEGE-
L1US) — Tennessee. Sevier County: Great Smoky
Mountains, Alum Cave, Degelius s.n., 13 Sept.
1939 (US. DEGELIUS) - Clingman's Dome. Dege-
üus s.n., 12 Sept. 1939 (DEGELIUS) - Great Smoky
Mountains National Park, litis & Sharp s.n.. 16
May 1942 (CANL) - West Virginia. Randolph County:
Winchester, Millspaugh 769 (NYBG) - Locality
unknown. Hale 10998 (US) - Pendleton County:
Panther Knob, Hale 14386 (US).
2. Alectoria imshaugii Brodo & D. Hawksw.,
S P- nov.
Thallus caespitosus vel subpendulus, rigidulus,
Plerumque 5-8 cm longus; rami aniso-dichotomiter
Ve l iso-dichotomiter fastigiati, frequentes e baso
emergen tes, uno loco ad substratum affixi. in angulos
acutos divergentes, irregulariter compress! et
angulares, diametro inaequales, 0.4-0. 7 mm diam.
basi; tota planta viridi-fulva vel straminea.
Rami laterales spiniformes veri et soralia desunt.
Pseudocyphellae copiosae, conspicuae, albae,
elongato-fusiformes, planiusculae vel convexae,
amplitudine variabites, spinulis isidiiformibus eis
hryoriae furcellatae similibus, copiose instructae,
'nterdum leviter sorediosae.
Fig. 25. Alectoria imshaugii. Holotype. Scale 0.5-1 .0
mm.
Apothecia rarissima. lateralia, ad circa 3 mm diam.,
excipulum thallinum thallo concolor et incurvatum.
discus concavus, brunneus. Ascosporae non visae.
Pycnidia ignota.
Cortex K- vel leviter flavescens, C-, CK-,
KC+ flavescens, PD-; medulla K+ flavescens
vel -, C-, CK-, KC— vel rarissime rubescens,
PD - vel flavescens-aurantiacus. Thallus acidum
usneicum continens una cum acido thamnolico vel
squamatico et duas substantias chemicas ignotas
(rarissime etiam acidum alectoronicum).
Holotypus: America septentrionalis, U.S.A.,
Idaho, circa 4.5 mis ad boreo-occidentalem a St
Joe, in summo monte St Joe Baldy. 116° 25’ W
et 47° 22' N, G. L. Schroeder L 1944, 19 July 1971
(CANL 38827). Fig. 25.
Thallus caespitose to subpendent, rigid, usually
5-8 cm long; branching anisotomic to isotomic
dichotomous, frequent from the base, attached
to the substrate at a single point, angles between
the dichotomies acute, branches irregularly
flattened and angular, uneven in diameter,
0.4-0. 7 mm diam. at the base; greenish
yellow to straw yellow throughout.
True lateral spinules and soralia absent.
Pseudocyphellae abundant, conspicuous,
white, elongate fusiform, plane to convex,
variable in size, bearing numerous isidiiform
spinules similar to those of Bryoria furcellata,
sometimes becoming sparsely sorediate
(Fig. 2 B ).
Apothecia very rare, lateral, to c. 3 mm
diam.; excipulum thallinum concolourous with
the thallus, incurved, disc concave, brown.
Ascospores not seen. Pycnidia unknown.
60 Alectoria in North America
OPERA BOT. 42 (1977)
Cortex K- or slightly yellow, C — , CK — ,
KC+ yellow, PD — ; medulla K+ yellow or — ,
C — , CK — , KC— or very rarely KC+ red,
PD— or PD + yellow to orange. Contains usnic
acid and two unidentified compounds together
with either thamnolic or squamatic acid (plus
accessory alectoronic acid in a small population
in California).
Ecology. Mostly on coniferous trees (especially
Pseudotsuga and Pinus ) and lignum, rarely
on rock; either in plateau or mountain areas
(750-1750 m elevation) or close to sea level
(2-85 m elevation); usually in dry, well-lighted
conifer stands. On the coast it occurs in
“pine barrens” of Pinus contorta.
Distribution. Western intermontane, centered
to the east of the Columbia Plateau in eastern
Washington, northern Idaho, and north-
western Montana with a westward extension
along the montane Ponderosa pine - Douglas
fir forest zones to the mixed pine forests of
central Oregon and northern California (Fig.
24 B). A specimen morphologically identical to
A. imshaugii but containing alectoronic rather
than thamnolic or squamatic acid was seen
from the mountains of Venezuela (Hale 43,198
[US]) and may be distinct. The disjunct speci-
men from the Queen Charlotte Islands (see
below) is not morphologically identical to the
others and may represent a distinct taxon.
Discussion
This extremely well-defined species has been
recognised as distinct by some students of
western North American lichens for many
years (notably, H. A. Imshaug in whose
honour it is named) but does not appear to
have been described previously. Its characters
are generally very uniform throughout its
range but with two interesting exceptions.
First, the species is capable of producing
either thamnolic acid or its corresponding
para-depside, squamatic acid; the thamnolic
acid chemotype is the more frequently en-
countered. No morphological differences are
correlated with the differences in chemistry
and in many instances both chemotypes have
been found to occur mixed in the same
herbarium packet. We detected, however, no
thallus containing both thamnolic and squamatic
acids.
Secondly, a departure from the uniform
morphology is seen in a specimen of the
disjunct population from the Queen Charlotte
Islands (Brodo 18224 [CAN L]). In this collection
the spinules are rather long and scattered along
the branches or arising from short raised
pseudocyphellae in contradistinction to plants
from its main distributional area which have
rather short and clustered spinules arising from
elongate and generally sorediate pseudo-
cyphellae. It is possible that the Queen
Charlotte population merits taxonomic recogni-
tion, perhaps at the rank of subspecies, but
with only a single collection at hand we are
reluctant to accord it any status. This specimen
belongs to the thamnolic acid chemotype.
Selected specimens
Canada. British Columbia. Kootenay District: Moyie
Lake Area, Schroeder LI 864 (SCHROEDER) -
Alberta. Waterton Lakes National Park, Imshaug
6088 (MSC).
U.S.A. California. Humbolt County: Manila,
Brodo 20502 (CANL) - Mendocino County: 8.4
miles W of junction Hwy I and 101. Tavares 345A
(UC) - Mendocino, Brodo 20489 (CANL) - Idaho.
Benewah County: Plummer. Schroeder LI324
(SCHROEDER) - Bonner County: Priest River Exp.
Forest, Esslinger 1157 (ESSLINGER) - Boundary
County: Kaniksu National Forest, Schroeder &
Anderegg L.2104 (SCHROEDER) - Latah County:
Moscow. Schroeder LI 157 (SCHROEDER) - Sho-
shone County: Calder, Schroeder LI 160
(SCHROEDER) - Montana. Glacier County: Glacier
National Park, Imshaug 5966 (MSC) - Oregon.
Douglas County: Leuthner 175 (US) -Grant County:
Malheur National Forest. Schroeder & Anderegg
L2102 (SCHROEDER) - Josephine County: Selma.
Becking 6909108 (BECKING) - Lane County:
Florence. Pike 1475 (CANL, DUKE) - Union
County: Camp Elkanah, Schroeder & Anderegg
L2109 (SCHROEDER) - Wheeler County: Ochoco
National Forest. Schroeder & Anderegg L2I11
(SCHROEDER) - Washington. Asotin County:
Fields Spring State Park, Imshaug 16480 (MSC) -
Chelan County: Wenatchee National Forest, Hoffman
OP-26 (US) - Pend Oreille County: Newport.
Schroeder L1065 (SCHROEDER) - Spokane County:
Mt. Spokane State Park. Esslinger 2555 (ESSLINGER)
- Stevens County: Colville National Forest, Schroeder
L21I8 (SCHROEDER) - Whitman County: Kamiak
Butte, Imshaug 16444 (MSC) - Yakima County:
Deadhorse Hill, Howard 982 (COLO).
Venezuela. Merida: Above La Aquada, Sierra
Nevada de Merida, alt. ca. 3500 m. Hale 43,198 (US).
OPERA BOT. 42 (1977)
Alectoria Ach. 61
3. Alectoria lata (Tayl.) Linds.
Trans. Roy. Soc. Edinb. 22: 135 (1859). - Corni-
cularia lata Tayl., Hook. Lond. J. Bot. 6: 190
(1847); type: Mexico, ex herb. Hooker (FH holotype!).
- Bn/opogon latus (Tayl.) Müll. Arg., Flora, Jena
71: 130 (1888).
Alectoria osteina Nyl., Flora, Jena 41: 378 (1858);
type: Mexique, Pic Orizaba, 10,000 ft, Galeotti 6947
(H-Nyl. 35983 holotype!). - Alectoria ochroleuca
(.osteina (Nyl.)Nyl., Syn. Lieh. I: 282(1860).
Alectoria japonica Tuck., Am. J. Arts Sei., ser. 2,
28: 202 (1859); type: Japan, Ayan, hillsides, on dead
pine trees, C. Wright (FH holotype!).
Alectoria ochroleuca f. gracilis Kremp.. Verh.
Zool.-Bot. Ges. Wien 18: 314 (1868); type: Mexico,
Vulcan Toluca, 10,000 ft, C. Heller 325 (M holotype!
ZT isotype!).
Alectoria lata f. suhfihriltosa Gyeln., in Sato,
J. Jap. Bot. 10: 19 (1934); type; Saghalien, Sisuka,
on the bark of Larix dahurica var. japonica. M. Sato
23 (not traced; no material with Prof. M. Sato
and none located in BP). - Bryopogon latus f.
subfibrillosus (Gyeln.) Gyeln., Feddes Repert. 38:
249 (1935).
Thallus caespitose to subpendent, occasionally
prostrate and entangled, stiff, usually 5— 8(— 1 5)
cm long; branching mainly isotomic dicho-
tomous from the base, sometimes becoming
anisotomic dichotomous towards the apices,
angles between the dichotomies mainly acute,
sometimes with short perpendicular lateral
branches; branches usually somewhat angular,
even to uneven in diameter, 0.5—1 ,5(— 2.0) mm
diam.; yellow to yellowish green, sometimes
becoming blackened in parts, especially towards
the base, so as to assume a striately blackened
appearance.
True lateral spinules and soralia absent.
Pseudocyphellae abundant, conspicuous, raised,
elongate fusiform, white, mainly 0. 5-1.0 mm
long, not bearing isidiiform spinules.
Apothecia common, occasionally absent,
lateral but sometimes appearing geniculate due
to the death or disintegration of the branch
distal to the ascocarps; excipulum thallinum
concolourous with the thallus, incurved; disc
concave, brown to black, 2 — 4(— 10) mm diam.
Spores 2(-?4) per ascus, ellipsoid, with a
hyaline epispore. becoming brown at maturity,
simple 35 — 45 x 18-24 gm. Pycnidia not seen.
Cortex K — or sometimes K.+ red, C— or
sometimes C + green-black, KC + yellow,
p D-; medulla K-, C- or slowly becoming
yellow, KC+ red or -, PD-. Contains
usnic and alectoronic acids, and occasionally,
the K + red unknown referred to on p. 29.
Ecology. On conifer branches, soil, and rocks,
usually at high altitudes.
Distribution. Central Mexican highlands north-
ward along the Sierra Madre Occidental, then
disjunct to the Cascade range in northern
California, Oregon and Washington (Fig. 24C).
Having seen the luxuriant and typical growth
of this species in northern California in stands
of Pinus jeffrey i which probably resemble
the montane Mexican pine forests closely, it
seems probable to us that the species occurs
elsewhere in the California mountains in suitable
habitats. It should be noted that the range of
Pinus jeffreyi extends without interruption into
northern Baja California.
The record of this species from British
Columbia (Räsänen 1933, Gyelnik 1935) was
based on a fragmentary specimen in H filed
under this name; it is most probably A. sarmen-
tosa subsp. vexillifera (Hazelton, ad terram,
8-10 August 1931, V. Kujala).
Outside North America, A. lata is essentially
a species of eastern Asia but also occurs in
Central America in Guatemala (Huehuetenango,
summit of Cuchumatanes Mts. above Chiantla,
3090 m, corticole, C. W. Dodge, 13 May
1941, herb. Dodge). A map of its known world
distribution has been published by Hawksworth
(1972 p. 242). The possible origin of such a
disjunction is discussed in the general section
Distribution, category 15.
Discussion
Alectoria lata is a well-defined species which
may be distinguished from other yellow species
of the genus by the more caespitose habit
and generally abundant ascocarps. On Horse
Mountain (California) this species has been
seen by one of us (l.M.B.) growing with A.
sarmentosa subsp. sarmentosa where their
habits render them easily separable in the field.
Specimens at the northern edge of the range
of the species in North America tend to be
somewhat more slender than those in the
central parts of its range. A sterile collection
from the Queen Charlotte Islands (Graham
L, Newton Point, R. Taylor 126 [COLO])
may also belong here, but it is unusually
long in part.
62 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 26. Alectoria mexicana. Holotype. Scale 0. 5-1.0 mm.
Selected specimens
Exsiccatae. Weber. Lieh. Exs. 417: California,
Humboldt County: summit of Horse Mountain,
Pike & Becking (CANL).
U.S.A. California. Del Norte County: Bear Basin,
Tucker 6433 (MSC) - Oregon. Josephine County:
Selma, Becking 6908108 (BECKING) - Benton
County: Corvallis, Mary's Peak, Brodo 20952
(CANL) - Clatsop County: Saddle Mtn. State Park,
Pike 3817 (OSC) - Washington. Pacific County:
South Bend, Herre s.n.. 20 Aug. 1939 (F). - Pierce
County: Mt. Rainier National Park. Van Trump
Peak, Howard 4428 (LAM) - Skamania County:
Gifford PinchotNat. Forest, Pechanec 1023 (COLO)-
Kittitas County: Stampede Pass, Thiers 35180
(SFSU).
Mexico. Chihuahua. San Pedro Springs, Gooding
s.n., Jan. 1907 (F, US) - Durango. El Salto, Bell
s.n., June 1965 (DXJKE)-Guerrero. Mt. Popocatepetl,
Moore 4942 (US) - Mexico. Nevado di Tolucco
(sic), Pringle s.n., 25 Sept. 1892 (FH, COLO.
FH:Riddle) - Michoacan. Tancitaro region. Leaven-
worth 9 (US) - Oaxaca. Cerro San Felipe, Beharre!
s.n., 14 Aug. 1975 (CANL 55341) - Vera Cruz.
Pic Orizaba, Galeotti 6947 (H:Nyl).
Guatemala. Sacatepéquez. Volcan de Agua,
Kellerman s.n,, 15 Feb. 1905 (US) - Huehuetenango,
summit of Chucumatanes Mts., C. W. Dodge s.n.,
13 May 1941 (DODGE).
4. Alectoria mexicana Brodo & D. Hawksw.,
sp. nov.
Thallus subpendulus vel pendulus, 10—1 2(— 22) cm
longus; rami plerumque e baso iso-dichotomiter
fastigiati. praecipue in angulos obtusos divergentes,
teretiusculi vel leviter compressi et angulares,
aequales vel inaequales diametro, 0.5-1 .5 mm diam.;
tota planta straminea.
Rami laterales spiniformes veri desunt. Soralia
aliquando producta, conspicua, alba, tuberculata,
latiora quam rami ex quibus orientia, ad 1-1.5 mm
diam. Pseudocyphellae copiosae, conspicuae, albae,
depressae vel elevatae. elongato-fusiformes vel ovales,
plerumque circa 0.5-0. 7 mm longae, rarissimo
spinulis isidiiformibus instructae. Medulla alba, lata,
ubique densa.
Apothecia vulgaria, lateralia sed interdum ap-
parenter geniculata, plerumque 3^1 mm diam.,
excipulum thallinum thallo concolor, primo in-
curvatum sed gradatim fere exclusum, discus
planiusculus vel convexus, brunneus vel niger. Asci
2(-4)-spori. Ascosporae ellipsoideae, episporio
OPERA BOT. 42 (1977)
Alectoria Ach. 63
hyalino, gradatim brunneae, simplices, 35 — 43 .5 x 21-
27 ;u.m. Pycnidia ignota.
Cortex K-, C— , CK-, KC+ flavescens, PD-;
medulla et soralia K-, C+ rubescentes, CK +
rubescentes, KC -t- rubescentes, PD - . Thallus acidum
usneicum et acidum olivetoricum continens.
Holotypus: Nova Hispania. provincia Oaxacae,
Sierra de San Felipe, in pino, alt. 10,500 ft. C. G.
Pringle 195. 3 June 1894 (CANL 16582). Isotypi:
ALA. BM. COLO. DUKE. LE, MINN. MSC.
NYBG.UC.W1S. Fig. 26.
Thallus subpendent to pendent, 10— 12(— 22) cm
long; branching mainly isotomic dichotomous
from the base, angles between the dichotomies
mainly obtuse, branches terete to slightly
flattened and angular, even to uneven in
diameter, 0.5-1. 5 mm diam.; straw yellow
throughout.
True lateral spinules absent. Soralia rare,
conspicuous when present, white, tuberculate,
broader than the branches on which they arise,
to 1-1.5 mm diam. Pseudocyphellae abundant,
conspicuous, white, depressed to elevated,
elongate-fusiform to oval, mainly c. 0.5-0. 7
mm long, rarely bearing isidiiform spinules.
Medulla white, thick and dense throughout.
Apothecia common, lateral but sometimes
appearing geniculate (due to the death and
disintegration of the branch distal to the
apothecium), mainly 3—4 mm diam., excipulum
thallinum concolourous with the thallus, at
first incurved but tending to become excluded
with age. disc plane to convex, brown to black.
Asci 2(^t)-spored. Ascospores ellipsoid, with a
hyaline epispore, brown at maturity, simple,
35-43.5 x 21-27 /am. Pycnidia unknown.
Cortex K -, C — , CK-, KC + yellow, PD-;
medulla and soralia K — , C + red, CK+ red,
KC + red, PD - , Contains usnic and olivetoric
acids.
Ecology. On trees at high altitudes, probably
in pine forests (2850-3500 m).
Distribution. Known only from two localities
■n the Central American highlands (Fig. 27),
and possibly limited to the high altitude pine
forests.
Discussion
This previously unrecognized species, which is
most closely related to A. vancouverensis in
•Is chemical components, differs from that
species in its habit, colour, ability to produce
Fig. 27. Alectoria mexicana. Known distribution.
soralia, and frequent apothecia. Soralia have
only been seen in a few of the duplicates of
Pringle 195 examined and so cannot be regarded
as a diagnostic character for this species;
they are probably of sporadic occurrence here
as they are in other members of this genus.
Studies of populations of this species in the
field are needed before a more detailed account
of its variability and affinities can be produced.
Specimens
Mexico. Oaxaca. Sierra de San Felipe, Pringle 195
(ALA, NYBG, W1S, MSC, CANL, COLO, UC,
DUKE), S 21,157 (DUKE) - Cerro San Felipe,
Beharrel s.n., 14 Aug. 1975 (CANL 55334, 55342,
55335).
Guatemala. Totonicapân. Santa Maria Tecum,
30 km from Totonicapân, C. W. Dodge 11551
(DODGE).
5. Alectoria nigricans (Ach.) Ny I .
Lieh. Scand.: 71 (1861). - Cornicularia ochroleuca
ß. nigricans Ach., Lieh. Univ.: 615 (1810); type:
Lapponia (H-Ach. lectotype!).
Alectoria boryana Del., in Th. Fr., Nova Acta
Reg. Soc. Sei. Upsal., Ser. 3, 3: 28 (1860)
[Reprint]; type; Newfoundland, 18278, Despréaux,
ex herb. Delise (BM lectotype! FH isolectotypel);
nom. inval. (Art. 34).
For further synonyms see Flawksworth (1972 p.
224). Alectoria arctica Elenk. & Saviez (Acta Florti
Petrop. 32: 73, 1912) was tentatively placed as a
64 Alectoria in North America
OPERA BOT. 42 (1977)
synonym of A. nigricans by Hawksworth (1972 p.
224) on the basis of the original description and
illustration. We examined the holotype in 1975
(Novaya Zemlya, simus Krestovaja, in terra inter
alios lichenes. J. V. Palikin, 1901 [LE]) and con-
cluded that it is a species of Evernia. Its medulla
is more-or-less dense throughout, and the cortex
is scarcely differentiated; TLC of a fragment showed
it to contain usnic and what appears to be divaricatic
acid. We are thus in agreement with Lynge (1928),
who transferred the species into Evernia (as E.
arctica (Elenk. & Saviez) Lynge), and Llano (1951),
who, because E. arctica is a later homonym of
E. arctica (Hook.) Tuck. ( =Dactylina arctica
(Hook.) Nyl.), created the new name, Evernia
perfragilis Llano, for the taxon.
Thallus erect, sometimes becoming decumbent
to prostrate (in f. subchatybeiformis Räs.),
usually 3-8 cm tall; branching anisotomic
dichotomous from the base, often appearing sub-
monopodial towards the apices; angles between
the dichotomies variable but usually mainly
acute; branches terete at first but becoming
compressed and sometimes slightly foveolate
towards the base, to 2.5 mm diam.; thallus
dull, rarely shining in parts, mottled grey-
fuscous to black or entirely black, often pale
brown or pinkish below, discolouring herbarium
paper reddish after about 10 years; dying
from the base.
True lateral spinules absent. Soralia (f.
sorediata Dahl) unknown in North America.
Pseudocyphellae abundant, conspicuous,
usually slightly raised, fusiform to fissurai,
mainly 0. 5-1.0 mm long.
Apothecia infrequent (most commonly seen
on corticolous specimens), lateral but sometimes
appearing geniculate; excipulum thallinum thin,
concolourous with the thallus, becoming ex-
cluded; disc concave at first, becoming plane
and then convex with age, yellowish brown to
pale reddish brown, ( 1 .0—) 1 .5— 4.0(— 9.0) mm
diam.; thecium 80-90 pm tall. Spores 2(— 4)
per ascus, ellipsoid, hyaline at first but becoming
reddish brown when mature, with an hyaline
epispore, simple, (21-)30— 39(-4 1 ) x (12-)20-23
Aim. Pycnidia not seen.
Cortex and medulla K faintly + yellowish
(rarely K+ red), C + rose-red, KC + red,
PD+ yellowish. Contains alectorialic acid,
usually traces of barbatolic acid, and some-
times an unidentified K+ red (PD-) substance.
Ecology. On rocks and soil, frequently calcar-
eous, and over ground vegetation; often also
on the low branches of spruce or fir trees.
In highly exposed habitats, this lichen is often
restricted to cracks and spaces between rock
fragments.
Distribution. Circumpolar arctic-alpine to
northern boreal, especially common in the
Ungava Peninsula (Fig. 28). It is apparently
rare in the Rocky Mountains, and more common
in the Coast Range. A. nigricans has a classical
bipolar distribution (see Hawksworth 1972 p.
227 for a map of its world distribution).
Discussion
The identification of this frequently abundant
species generally presents few problems.
Particularly dark specimens have sometimes
been confused with Bryoria chalybeiformis
or B. nitidula in herbaria but these can be
readily separated on the basis of chemical
criteria.
The frequent occurrence in the genus Alec-
toria of a K + red, C+ green-black substance
in the dying basal portions of the thallus has
already been referred to in the general section
on Chemistry. In one specimen from South-
ampton Island (Parker SP-70-14 [CANL]). how-
ever, the whole thallus contained this substance.
The thallus of this specimen was also shiny
rather than dull and more divergently branched
than is usual of this species. Having seen
only a single specimen of this chemotype.
we have been unable to make a reliable
assessment of its taxonomic importance.
Despite the very broad world distribution
of this species, apothecia are only known
from North American specimens. Apothecia are
especially frequent in the eastern part of the
species' range.
Selected specimens
Canada. British Columbia. Chilkat River Basin:
Haines Road, mile 60, Szczawinski 1259/5 (US) -
Coast District: Terrace, Ohlsson 2817 (CANL) -
Fraser River Basin: Wells Gray Provincial Park.
Ahti 14278 (H) - Islands North District: Moresby
Island: Takakia Lake, Brodo 10967 (CANL) - Koo-
tenay River Basin: Gold Range, Macoun 4, Aug. 1889
(CANL)- Skeena River Basin: E of Hazelton, Ohlsson
2986 (MSC) - Alberta . Banff National Park. Silvertip
Mountain, Lamb 6444 (MSC) - Manitoba. Churchill, J.
W. Thomson 3712 (CANL, DUKE) -Ontario. Kenora
District: Cape Henrietta Maria, Ahti 4995 (H) - Fort
OPERA BOT. 42 (1977)
A le et or ia Ac h. 65
>
1 ‘"■ysA/’l y r
T^— s
Rg. 28 . Alectoria nigricans. N. American distribution.
Severn, Ahti 4220 (H) - Pen Island, Kershaw s.n.
(CANL 36003) - Quebec. Cte Gaspe-Ouest: Parc de
Gaspesie, Mont Albert. Brodo 18537 (CANL) -
Territoire de la Baie de James: Goose Bay. Kucyniak
& Tuomikoski 361 (CANL) - Territoire de Mistassini:
Otish Mountains, Moni du Lagopède, Shchepanek
' l-L-50 (CANL) - Nova Scotia. Victoria County:
Tape Breton Highlands National Park. Scotter 6280
TH) - Newfoundland. Labrador South District: Battle
Harbor, Waghom s.n., 25 Aug. 1891 (NYBG).
iSee also Ahti & Hawksworth 1974.) - Yukon.
British Mountains. Trout Lake, J. W. Thomson &
Larsen 14756 (p.p.) (WIS) - Northwest Territories.
* ' Opera Botanica nr 42
District of Franklin: Baffin Island: Barnes Ice Cap,
Haie 480 (CANL) - District of Keewatin: Coral
Harbor, Southampton Island, Weber S 23,628
(CANL, COLO, NYBG) - Mackenzie District:
Artillery Lake, J. W. Thomson & Larsen 1 1334 (WIS).
U. S. A. Alaska. Aleutian Islands: Agattu Island,
Townsend 78 (US) - Unalaska. Setchell s.n., 1899.
June-Aug. (FH) - Alaska Range District: Mt.
McKinley National Park, Mt. Eielson, Weber &
Viereck S7084 (CANL, COLO, DUKE, US, FH) -
Bering Sea District: St. George Island. J. M. Macoun
38 (CANL) - St. Lawrence Island, Geist s.n.,
1933 (US) - St. Paul Island, J. M. Macoun 39
66 Alectoria in North America
OPERA BOT. 42 (1977)
(CANL) - Eastern Pacific Coast District: Mt. Juneau,
Imshaug 28714 (MSC) - Colorado. Summit County:
North Star Mountain. Weber 6531 (COLO) - New
Mexico. Otero County: Sierra Blanca Peak,
Sacramento Mountains, Egan El-1560 (MSC) -
Washington. Clallam County: Olympic National Park,
Obstruction Peak, Brodo 14409 (CANL) - Jefferson
County: Moose Lake Trail, Imshaug 17588 (WIS) -
Pierce County: Mt, Rainier National Park, Burroughs
Mountain, Imshaug 17354 (MSC).
6. Alectoria ochroleuca (Hoffm.) Massai.
Sched. Crit. Lieh, Ital . : 47 (1855). - Usnea ochroleuca
Hoffm., Descr. Adumbr. PI. Lieh. 2(1): 7 (1791);
type: Funck, Crypt. Gew. Ficht, no. 420 (MW-Hoffm.
8562 neotype!).
For a detailed treatment of the nomenclature
of this epithet see Laundon (1970) and for further
synonyms see Hawksworth (1972 p.p. 231-233).
Thallus erect, usually 5— 8(— 12) cm tall or,
less frequently, decumbent; branching aniso-
tomic dichotomous appearing submonopodial
and of the Cladina type, apices often falcate
and drooping; branches terete, often straight,
even to uneven in diameter, sometimes be-
coming compressed towards the base, usually
0. 5-2.0 mm diam.; yellow to yellowish green,
the apices often becoming striately blackened
and the blackening sometimes extending down
the main stems; dying from the base.
True lateral spinules absent. Spinules rarely
arising from old pseudocyphellae. Soralia
absent. Pseudocyphellae abundant, con-
spicuous, raised, elongate fusiform to ovoid,
white, mainly about I mm long.
Apothecia occasional, sometimes abundant,
lateral, sometimes appearing geniculate; ex-
cipulum thallinum concolourous with the thallus,
thick, often incurved and persistent; disc
usually plane to concave, rarely becoming
convex, orange-yellow to reddish brown or
black, 3-6 mm diam. Spores 2(— 4) per ascus,
ellipsoid, with a hyaline epispore, becoming
dark brown when mature, simple, (26— >28—35
(-42) x ( 1 2—) 1 6— 22(— 28 ) /am. Pycnidia occasional,
apical or subapical, minute, black and shining,
to 0. 15 mm diam.; conidia 7-8 x 0.8 gm.
Cortex K- (sometimes K + reddish at the
base), C— , KC + yellow, PD—; medulla K— ,
C — , PD — , KC — , CK + yellow-orange; rarely
KC + red, CK-; or very rarely KC-, CK-.
Contains diffractaic or alectoronic acid, and
usnic acid and sometimes an unidentified K +
reddish substance in the base. The diffractaic
race (med. KC-, CK+ yellow-orange) is
much more widespread than the alectoronic
race (KC+ red, CK— ). Acid deficient
specimens are also known (KC-, CK-) but
are extremely rare.
Ecology. On dry, or less frequently, wet
tundra soil; characteristically found on or
between non-calcareous or calcareous rock
fragments on talus and fel-fields. It is infre-
quently found on the lower branches of spruce
or willow in the northern subarctic region.
Distribution. Arctic to northern boreal, with
scattered alpine localities in the western
Canadian mountains, and in Quebec's Gaspé
Peninsula (Fig. 29). It appears again in the
very high Mexican ranges, the Andes of South
America, and Tierra del Fuego. In view of
their isolation it is interesting that the Mexican
populations tend to contain alectoronic rather
than diffractaic acid. The plants that we ana-
lyzed from Tierra del Fuego, on the other
hand, contained diffractaic acid. A. ochroleuca
is much more widespread in North America
than is Bryoria nitidula, being fairly common
in the northern boreal zone, especially on
exposed, alpine-like ridges. In fact, it is the
most widespread North American Alectoria.
This species is widely distributed in other
parts of the world as well (e.g., all the European
mountain ranges, Japan, China, India, Nepal,
New Zealand) although it is not very common
in the British Isles (Hawksworth 1972).
Discussion
Several collections from the Mexican highlands
referred to this species were unusually slender
and delicate and provided a strong KC + red
medullary reaction (shown to be due to
alectoronic acid by TLC). It is possible that
such plants belong to the taxon termed var.
variegata (Samp.) Zahlbr. (syn. Bryopogon
guatemalensis Gyeln.) by Hawksworth & Moore
(1969). In many cases, it proved difficult to
distinguish this morphotype from slender,
terricolous specimens of A. lata. A rare
alectoronic acid race with a typical morphology
is also present in the American arctic (e.g.,
Aleutian Islands, Alaska; Churchill, Manitoba).
OPERA BOT. 42 (1977)
Alectoria Ach. 67
Pig. 29. Alectoria ochroleuca. Central and N . American distribution.
68 Alectoria in North America
OPERA BOT. 42 (1977)
Specimens with alectoronic acid were also seen
from Uruguay and Venezuela.
In general the North American material of
this species contains diffractaic acid as does
all European and Asian material so far ex-
amined. It can thus be distinguished from
rather similar morphotypes of A. sarmentosa
subsp. vexillifera lacking alectoronic acid
(medulla KC-) by TLC (and also the CK.+
test noted under Spot tests in the general
section Chemistry ).
The neotype in the Hoffmann herbarium
has a typical erect habit and apparently
contains only diffractaic and usnic acids as
determined by TLC (although we were unable
to demonstrate good diffractaic crystals in GE
solution).
Motyka’s (1964) reference to the occurrence
of a sorediate morphotype of A. ochroleuca
in North America (treated as “f. sorediosa
Lynge”) appears to be erroneous; Lynge’s
epithet belongs to A. sarmentosa, as pointed
out by Hawksworth (1972 p. 240).
Selected specimens
Exsiccatae. Merrill, Lieh. Exs. Ser. 11, 106: Alaska.
Cantwell, Palmer (ALA, BM, COLO, DUKE, F,
MSC, NYBG, US, W1S) - Thomson: Lieh. Arct.
59: N.W.T., District of Keewatin: Rossby Lake.
J. W. Thomson (CANL, COLO, DUKE, LAM.
MSC) - Thomson: Lieh. Arct. 90: N.W.T., District
of Mackenzie: Artillery Lake, J. W. Thomson &
Larsen (CANL, COLO, DUKE) - Weber: Lieh.
Exs. 220: N.W.T., District of Mackenzie: Mackenzie
Mts., O'Grady Lake, Cody 16815 (BM, CANL,
COLO, US).
Canada. British Columbia. Chilkat River Basin:
Mile 77, Haines Road, Szczawinski 28/62 (US) -
Coast District: Mt. Walker, E of Bella Coola.
Ohlsson 2240 (CANL) - Fraser River Basin: Wells
Gray Provincial Park, Battle Mountain, Ahti 14277
(H) - Liard River Basin: Summit pass, Alaska
Highway, Szczawinski 177/1 (US) - Skeena Rivet-
Basin: E of Hazelton on Nine Mile Mountain, Ohlsson
2979 (CANL) - Yukon River Basin: Altin Lake Re-
gion. Anderson 925 (MSC) - Alberta. Banff National
Park. Snow Creek Pass, Weber S23.486 (COLO.
CANL, FH, DUKE) - Manitoba. Fort Churchill,
J. W. Thomson 3589 (CANL) — Ontario. Kenora
District: Cape Henrietta Maria, Courtin 7 (CANL) -
Quebec. Clé Gaspé-Nord: Mont Jacques-Cartier,
Gauthier 2578 (CANL) - Territoire du Nouveau
Québec: Gérin Mountain, Viereck 702 (BM, ALA,
CANL, COLO, DUKE) - Newfoundland. St. Barbe
North District: Highlands of St. John, Femald 2271
(FH, US) - Labrador North District: Crater Lake
vicinity, Gillett 8765 (CANL) - Labrador South
District: Battle Harbor, Waghome s.n., 25 Aug.
1891 (NYBG) - Yukon. Stewart Plateau, Porsild
5B (CANL) - Northwest Territories. Mackenzie
District: Vicinity of Lake-on-the-Mountain, Thieret &
Reich 5910 (CANL, FH, NYBG, W1S, US) -
Keewatin District: Parsons Lake, J. W. Thomson &
Larsen 5933 (COLO. US) - Franklin District:
Strathcona Sound, Admiralty Inlet, Soper 381 (CANL.
DUKE. NYBG, US).
U. S. A. Alaska. Aleutian Islands: Amchitka Island.
Reich & McCann 60 (F, CANL. WIS) - Amlia
Island, Eyerdam s.n., 10 July 1932 (NYBG) - Attu
Island, Hardy 14 (COLO) - Kiska Island. McCarthy
& Kellas 5 (CANL) - Unalaska, Bean 3 (US) -
Alaska Range District: Mt. McKinley National Park.
Mt. Eielson, Weber S7061 (ALA, BM. CANL, COLO.
DUKE, FH).
Mexico. Mexico. Nevado de Toluca, Kiener 30894
(WIS, MIN, COLO) - Volcan de Toluca, litis
3127 (WIS) - Puebla. Mt. Popocatepetl, Kiener 18585
(WIS, COLO, MIN) - Vera Cruz. Orizaba, J. G.
Smith 1? (US, MIN).
7a. Alectoria sarmentosa (Ach.) Ach. subsp.
sarmentosa
Lieh. Univ.: 595 (1810). - Lichen sarmentosus
Ach., K. Vet. Acad. Handl. 16: 212 (1795); type:
Suecia (H-Ach.. upper specimen, lectotype!).
Alectoria luteola Mont, ex de Not., Giorn. Bot.
ltal. 1: 206 (1846); type: Newfoundland, 1828.
Depréaux, herb. Montagne ex herb. Delise (RO
lectotype! FH isotype!).
Alectoria subsarmentosa Stirt . , Trans. Proc. Bot.
Soc. F.dinb. 14: 358 (1883); type: Newfoundland,
trom rocks near Brigus, A. Gray, August 1878
(E holotype! BM isotype!).
Alectoria sarmentosa var. gigantea Räs., Ann.
Missouri Bot. Gard. 20: 10 (1933); type: British
Columbia. Longworth, ad corticem Thuja occiden-
talis, V. Kujala, September 1931 (H holotype! H
two isotypes !).
Alectoria sarmentosa var. hypocyphellata Gyeln.,
Ann. Mus. Nat. Hung. 28: 283 (1934); type: Oregon,
Crater Lake, on limbs [of trees], F. P. Sipe 680,
August 1930, (ORE lectotype!); nom. inval. (Art. 32). -
Eualectoria sarmentosa var. hypocyphellata Gyeln..
Ann. Mus. Nat. Hung. 28: 283 (1934); nom. inval.
(Arts. 32, 34).
Alectoria sarmentosa var. alaskana Mot., in Krog.
Norsk Polarinst. Skr. 144: 138 (1968); type: Alaska,
Eastern Pacific Coast District, Sitka. Harbour Mt..
H. Krog 6255, 31 August 1957 (O lectotype!); nom-
inval. (Arts. 32, 36, 37).
Alectoria sarmentosa var. brevior Mot., in Krog.
Norsk Polarinst. Skr. 144: 139 (1968); type: Alaska.
Eastern Pacific Coast District, Ketchikan, Deer
Mt., H. Krog 6263, 3 September 1957 (O lectotype!):
nom. inval. (Arts. 32, 36, 37).
Alectoria sarmentosa var. macra Mot., in Krog.
Norsk Polarinst. Skr. 144: 139 (1968); type: Alaska,
Eastern Pacific Coast District, Juneau, Loops Road.
OPERA BOT. 42 (1977)
Alectoria Ach. 69
H. Krog 5565, 24 August 1957 (O lectotype!); nom.
inval. (Arts. 32, 36, 37).
Alectoria stigmata Bystr., Frag. Flor. Geobot.
22: 255 (1973); type: Alaska, Central Pacific Coast
District. Kenai Peninsula, Creek Ridge, H. Krog
s n., 28 May 1957 (LBL holotype!).
For further synonyms see Hawksworth (1972 p.p.
238-239).
Thallus pendent, flaccid except at the base,
commonly 20^)0 cm long, exceptionally to
80 cm; branching mainly isotomic dichotomous,
frequent from the base, angles between the
dichotomies variable, acute or obtuse, often
rounded; branches terete often becoming com-
pressed and angular to foveolate towards the
base and at the axils, 0.5— 2.0(— 2.5 ) mm diam.;
greenish grey to bright golden yellow, some-
times becoming striately blackened in parts;
base of the thallus persistent.
Lateral spinules absent. Soralia irregularly
tuberculate, rare in North American material
(see Discussion). Pseudocyphellae abundant,
conspicuous, raised, elongate fusiform to ovoid
and tuberculate, clearly delimited, white, usually
c. 1 mm long.
Apothecia often abundant (especially in the
w est), lateral; excipulum thallinum con-
colourous with the thallus, usually persistent;
disc orange-yellow to dark brown or black,
2 -3(-5) mm diam. Spores 2-3 H 0 per ascus,
dlipsoid, with a hyaline epispore, becoming
dark brown at maturity, simple, 23— 35(— 48) x
(12-)15_20(-25) (im. Pycnidia sometimes
frequent, mainly apical, to c. 2.0 mm diam.,
black and shining; conidia not seen.
Cortex K -, C — , KC + yellow, PD - (some-
times K+ red, C + green-black near the
base); medulla K- (rarely K+ yellow), C— or
slowly becoming yellow, KC+ red or -,
TK— , PD— (rarely PD + yellow). Contains
hsnic acid, ± alectoronic acid, rarely thamnolic,
[squamatic,] or barbatic acids, and occasionally
an unidentified K+ red, C+ green-black sub-
stance.
Ecology. On a variety of trees, but especially
conifers, in Picea-Tsuga heterophylla forests.
Most collections are from Picea (especially
T. sitchensis ), Tsuga heterophylla, Abies spp.,
Pseudotsuga menziesii, and Pinus contorta.
Rarely, collections are made from the ground
0r rocks. The species grows most luxuriantly
along the edges, or in glades, of the tall,
tfoist, lowland forests along the west coast.
In Wells Gray Park, British Columbia, A.
sarmentosa was found to be largely confined
to lower tree branches within 30 feet of the
ground, apparently requiring more moderate
levels of light, moisture, and temperature than
the brown species (Edwards et al. 1960).
Distribution. Mainly along the west and east
coasts in humid coniferous forests (Fig. 30).
Its close association with the more mesic
western hemlock forests both along the coast
and in the Rocky and Selkirk Mountains,
avoiding the drier inter-montane regions, is
apparent in the distribution map. Ahti (1964)
suggests that, rather than lower humidity, the
colder winters of the interior (e.g., northern
Ontario) may be a critical factor in largely
restricting it to the coasts. In the Appalachian
mountains, A. sarmentosa is replaced by its
vicariad. A. fallacina. The arctic localities may
be based on unusually slender forms of subsp.
vexillifera; the two taxa intergrade to some
extent (see Discussion under the latter).
The alectoronic acid-deficient race can be
found scattered throughout the range of the
species and appears to be of no taxonomic
importance, although its particularly high
frequency in Newfoundland should be noted
(Ahti & Hawksworth 1974). The world distri-
bution of this circumboreal species has been
mapped by Hawksworth (1972 p. 242). Poelt
(1969) reports that it is common in the boreal
and subalpine forests in mountainous regions of
northern, southern and central Europe. Al-
though A. lata appears to replace/!, sarmentosa
in Japan (loc. cit.) the two species grow side
by side in the montane Jeffrey pine forests
of northern California.
Discussion
Specimens of this subspecies containing
abundant quantities of thamnolic acid in addition
to alectoronic acid, were seen from California.
These specimens were often robust, having a
dense medulla very much like that of A.
vancouverensis. Less robust specimens with
small amounts of thamnolic acid were also
seen from this area and from the Queen
Charlotte Islands. We regard these collections
as representing only a chemical race of A.
sarmentosa, although further collecting and
70 Alectoria in North America
OPERA BOT. 42 (1977)
OPERA BOT. 42 (1977)
Alectoria Ach. 71
more careful study may result in a réévaluation
of their taxonomy.
A reading of the descriptions of A. imshaugii,
A. vancouverensis, and A. sarmentosa will
reveal the immense complexity of the morpho-
logical and chemical variation in this species
group, especially along the west coast, and in
Particular, in California. No explanation for
this complexity can be offered at this time.
One simply must accept the fact that the
variation exists and hope that closer attention
to the Alectoriae of California will clarify the
situation.
While the distinctiveness of these three
taxa, at some taxonomic level, can hardly be
questioned, one must grapple with the un-
deniable fact that clear morphological and
chemical intermediates exist between all pos-
sible combinations of them. There is even
one excellent specimen (Tavares 371) which
combines the chemical characters and many
of the morphological features of all three taxa
(see also Discussion under A. vancouverensis).
If one may put aside for the moment the
Possibility that the variation might be produced
by environmental factors (since no obvious
ecological or distributional correlations were
apparent), one is left with two possible expla-
nations: (a) the enzyme systems leading to the
production of lichen substances in the genus
Alectoria are somewhat plastic, and that a
certain amount of variation should be expected
*n all the taxa, and (b) that some sort of genetic
recombination or "hybridization” is occurring
among the taxa.
If we were dealing with possibility “a”,
We would expect a certain amount of chemical
Variation in plants showing little or no morpho-
logical intermediacy. Such might be the case
f°r the many “typical” A. sarmentosa speci-
me ns found to contain traces or large amounts
°( thamnolic acid in addition to alectoronic
a cid. It should be noted that these thamnolic-
ulectoronic-containing A. sarmentosa plants
can be found over a very large range: from
l be Queen Charlotte Islands to central Cali-
fornia. As stated previously (see p. 28), the
ability for squamatic acid production has been
well-documented in A. sarmentosa (although
not the North American specimens). The
occasional appearance of the corresponding
meta-depside, thamnolic acid, would therefore
not be altogether surprising and probably has
nothing to do with the presence of this sub-
stance in A. imshaugii.
On the other hand, the very complex com-
bination of chemical and morphological char-
acters, especially in central and northern Cali-
fornia where all three taxa converge in the
same forests, can best be explained by pos-
tulating some kind of genetic exchanges (i.e.,
possibility “b"). The genetic exchanges may
be occurring through sexual mechanisms as
species in the genus Alectoria are frequently
fertile. Kristinsson (1969) suggested such an
explanation for the morphological and chemical
variation in Cetraria islandica s. lat. in Iceland.
In that case, one is dealing with a simpler
system: the presence or absence of a single
chemical substance and correlating morpho-
logical features. Even in the absence of sexual
reproductive bodies, however, it is possible to
have genetic exchange in certain non-lichenized
fungi by parasexual mechanisms such as
somatic heterokaryon formation. It may be
that under environmental stress, there is more
of a tendency for such a phenomenon to
occur.
Perhaps it is significant that northern Cali-
fornia is close to the distributional limits for
all three Alectoria species while at the same
time, they grow close together.
A true heterokaryon, moreover, although
haploid, might produce a phenotypic expression
of both parents, acting almost like a hétéro-
zygotie diploid. Thus, while an exchange of
characters might best be explained by sexual
reproduction and crossing-over, the addition
of certain chemical or morphological characters
might suggest heterokaryosis. C. Culberson &
Hale (1973) found that they could account
for the occurrence of compounds in addition
to the compounds found in one parental species
by postulating a hybridization which involved
("'g- 30. N. American distribution. -A: Alectoria sarmentosa subsp. sarmentosa, KC+ red chemotype. - B: AT.
sarmentosa subsp. sarmentosa, KC- chemotype. • A. sarmentosa subsp. vexiUifera. O A. sarmentosa,
s °rediate morphotype.
72 Alectoria in North America
OPERA BOT. 42 (1977)
a partial exchange of enzyme systems such
that new compounds were formed. It would
be extremely interesting to see a detailed
analysis of the chemical, morphological, and
geographical correlations of this complex similar
to that of Culberson & Hale for Parmelia
sect. Hypotrachyna.
There is, however, another possible mecha-
nism for the addition of chemical constituents.
Henssen & Jahns (1973 p. 221) recently sug-
gested the possibility that two closely related
mycobionts might grow together and form
a single lichen thallus, sharing the phycobiont.
Presumably, such a chimera would produce
the chemical products of both mycobiont races.
This is an attractive hypothesis and deserves
considerable attention. As discussed in the
section Taxonomic concepts: Species, any
possible role of the phycobiont is difficult
to assess at the present time.
How to deal with this variation taxonom-
ically is another problem. Kristinsson (1969)
concluded, in agreement with Imshaug (1957),
that perhaps using the rank of subspecies for
C. er ice tor am was the most realistic solution
to the C. ishindica problem. We decided,
however, to regard A. vancouverensis, A.
imshaugii. and A. sarmentosa all as equivalent
species, at least for the present. A. vancouveren-
sis maintains its distinctiveness more than
90 % of the time even in areas where it grows
very closely associated with A. sarmentosa:
in the field, we have had no difficulty in
distinguishing one from another. A. imshaugii
has such a remarkably distinct morphology
it would be clearly incorrect to consider it
merely as a subspecies. At first we were
tempted to recognize the Tavares specimen
(see Discussion under A. vancouverensis) as
nomenclaturally distinct, but, in the final
analysis, we decided to consider it as just
another strange variation of A. vancouverensis .
The variability in the morphology of A.
sarmentosa has led to the recognition of
numerous infraspecific taxa by Motyka (in Krog
1968). We have examined Krog’s collections
studied by Motyka and feel these are of little
or no taxonomic importance since inter-
gradations between all of them occur. Motyka’s
names are typified here by the packets which
he annotated "typus varietas”, etc. Two
collections (Krog 2212 and 6255) were labelled
as the type of v.ar. alaskana by Motyka; the
latter is designated as lectotype here as it is
larger and better developed.
Alectoria stigmata Bystr. is another species
based on one of Krog's specimens from Alaska
which we feel should be placed into synonymy
with A. sarmentosa subsp. sarmentosa. The
holotype, a portion of which was generously
sent to us by Dr. Bystrek, appears to be a
broadly pseudocyphellate morphotype; it
contains usnic and alectoronic acids.
An abundantly sorediate morphotype of this
species, var. sorediosa (Läng ex Ras.) DR.
(see Hawksworth 1972 p. 240), occurs rarely
in North America (Fig. 30 B) as it does in
Europe (Ahlner 1948). It seems to be of rela-
tively minor taxonomic importance.
Selected specimens (KC +, C —)
Exsiccatae. Brodo: Lieh. Can. Exs. 1: British
Columbia. S of Princeton, Brodo 8508 (CANL) -
Hale: Lieh. Amer. Exs. 126: Oregon, Lane County:
Willamette National Forests, Shushan sl-1876
(CANL, WIS) - Lojka: Lieh. Univers. 218: Insula
Miquelon. Delamare (H) - Merrill: Lieh. Exs., Ser.
L 13: Idaho, (no locality mentioned). Tower 1902
(BM [KC-], CANL, DUKE, NYBG. US) - Rel.
Tuck. 4: Oregon (no locality), E. Hall (DUKE, F,
FH, LAM, NYBG) - Univ. California: Lieh. Oregon
1126: Oregon, Linn County: Tombstone Pass,
Tavares & Chisaki 125 (BM . COLO, DUKE, F, MSC,
US) - Weber: Lieh. Exs. 132: California, Humboldt
County: Hoopa. Weber (CANL, COLO, DUKE, US).
Canada. British Columbia. Coastal District: Burke
Channel in Crayden Bay, Ohlsson 2034 (MSC.
CANL) - Columbia River Basin: Revelstoke, Shaw
851 (NYBG, US). Deen River Basin: Tweedsmuir
Provincial Park, Ohlsson 2207 (CANL) - Fraser
River Basin: Blue River, Brodo 15792 (CANL) -
Islands North District: Graham Island: Boulton Lake,
Brodo 9954 (CANL) - Islands South District: Tofino,
Ohlsson 1080-A (MSC) - Kootenay River Basin:
Glacier National Park. Ohlsson 310 (MSC) -
Okanagan District: Rock Creek, Brodo 15127
(CANL) - Skagit River Basin: Manning Provincial
Park, Bird 15417 (CANL) - Skeena River Basin:
W of New Hazelton. Ohlsson 2893A (CANL) -
Alberta. Oldman River Watershed. Waterton
Lakes National Park. Bird 16925 (CANL) - Quebec.
Cté Gaspé-Nord: Mont Albert Nord, Gauthier 2640
(CANL)- Parc des Laurentides: Junction of Highway
54A and 54 B. Scotter 6872B (H) - Territoire de
Mistassini: Otish Mountains on Mont du Lagopède.
Shchepanek 7I-L-17 (CANL) -Territoire du Nouveau
Quebec: Poste-de-la-Baleine, Brisson & Forest 20485
(CANL) - Schefferville. Brodo 9239 (CANL) -
Netv Brunswick. Albert County: Fundy National
Park, Ireland 10979B (CANL) - Charlotte County:
Campobello, collector unknown, July 1902 (FH) -
OPERA BOT. 42 (1977)
Alectoria Ach. 73
Nova Scotia. Inverness County: French Mountain,
Brodo 19002 (CANL) - Victoria County: St. Paul Is-
land. Erskine 53C2402 (CANL) - Prince Edward Is-
land. Prince County: Richmond, Ireland 10297D
(CANL) - Newfoundland. Labrador North District:
Goose Bay. Judd GB-52 (CANL) - Labrador South
District: Backway, Tanner s.n., 5 July 1937 (H). (See
also Ahti & Hawksworth 1974.) - Northwest Terri-
tories. District of Keewatin: Coral Harbor, Southamp-
ton Island, Brown 33 (CANL).
O.S.A. Alaska. Aleutian Islands: Nazan Bay,
Atka, Van Dyke 232 (FH) - Amchitka Island,
Reich 201 (CANL) - Salteny Cove, US Bureau
Soils Kelps Expedition s.n. (F) - Eastern Pacific
Coast District: Juneau, Auke Lake, Imshaug 28505
(MSC) - Central Pacific Coast District: Evans
Island, Port San Juan, Eyerdam 560 (LAM, ALA) -
California. Humboldt County: Horse Mountain, Brodo
-0515 (CANL) - San Bernardino County: San
Bernardino Mountains, Arrowhead Lake. Bonnie &
Templeton s.n., 29 Dec. 1929 (LAM) - Siskiyou
County: Sissou, region of the Upper Sacramento.
Howe 139 (NYBG) - Colorado. County unknown:
Justin s.n., no date (FH) - Idaho. Idaho County:
Sei way Falls, Selway River, Imshaug 16408 (CANL,
MSC) - Maine. Washington County: Head Harbor.
Norton s.n., 3 May 1921 (FH) - Piscataquis County:
"JL Katahdin, Degelius s.n.. 31 Aug. 1939
(DEGELIUS) - Montana. Glacier County: Avaleneh
Lake, Glacier National Park, Mains 6036 (FH) -
Nevada. Humboldt County: Paradise Valley, Britton
* n., 10 Aug. 1904 (FH) - New Hampshire. Coos
County: Mt. Washington, Merrill s.n.. Sept. 1904
(CANL, FH) - Oregon. Deschutes County: Doris
Lake in the Deschutes National Forest, Pike L-390
'DSC) - Washington. Clallam County: Hurricane
R'dge, Olympic National Park. Brodo 13213 (CANL).
Selected specimens (KC-)
Canada. British Columbia. Columbia River Basin:
r\, irit Mountains near Emerald Lake, Petersen 65
(NYBG)-Dean River Basin: Young Creek, Tweeds-
a>uir Provincial Park. Ohlsson 1899 (MSC) - Fraser
Biver Basin: Wells Gray Provincial Park, Battle
7*ountain, Ahti 6529 (H) - Islands North District:
Graham Island: Dinan Bay, Brodo 18387 (CANL) -
Hands South District: Vancouver Island: Mount
Arrowsmith, Shchepanek 312A (CANL) - Kootenay
■strict: East Kootenny (sic) (Kootenay), from Fort
ol ville to Rocky Mountains, Lyall s.n., 1861 (FH) -
- keena River Basin: Dorren (sic) (Dorreen), Kujala
I' n -' Aug. 1931 (H) - Nova Scotia. Annapolis
l °unty: Albany Crossing, Darrow s.n., 30 Aug.
v, (LAM) - Newfoundland. Lewisporte District:
1 ;° tr e Dame Bay, Pyke Island, Riewe P-50 (CANL,
ALA).
IY*n’^"^' /4 ^ as ^“- Eastern Pacific Coast District:
ah Island, Augustine Bay, Foster s.n., 7 July
I .B (LAM) - Central Pacific Coast District: Evans
j-. and, Prince William Sound, Eyerdam 560 (F) -
ahfornia. County unknown: Sacramento Canyon,
i n ea c r . Sima, Sisson (collector? or “Sissou”, locality
■Siskiyou County), July 1902 (F) - Idaho. Idaho
County: Selway Falls, Imshaug 16408 (CANL) -
Oregon. County unknown: Cascade Mountains, Hogg
Pass, Britton s.n., 19June 1952 (LAM)- Washington.
Clallam County: Olympic National Park, Hurricane
Trail, Herre 5171 (F).
Specimens ( sorediate morphotype)
Canada. Nova Scotia. Victoria county: Cape Breton
Highlands National Park, Scotter6212B (H).
U.S.A. Alaska. Eastern Pacific Coast District:
Windham Bay, Culbertson 118 (F, FH) - Aleutian
Islands: Amlia Island, Eyerdam s.n., 10 July 1932
(FH).
7b. Alectoria sarmentosa subsp. vexillifera
(Nyl.) D. Hawksw.
Taxon 19: 241 (1970). - Alectoria ochroleuca
[subsp.] vexillifera Nyl., in Kihlman, Medd. Soc.
Fauna FI. Fenn. 18: 48 (1891); type: U.S.S.R.,
Murmansk District, Lapponia ponojensis, ad
promentorium Orlow, in apertis glareosis, A. O.
Kihlman, 15 May 1889 (H-Nyl. 35997 holotype!
ZT isotype!).
For further synonyms see Hawksworth (1972
p. 243).
Plants differing from subsp. sarmentosa
in having prostrate thalli to 8-15 cm long which
are rigid, more sparsely branched; branching
becoming anisotomic dichotomous towards the
base, main stems becoming dorsiventrally
compressed, expanded, and foveolate, some-
times with spinulose side branches, usually
2— 4(— 40 ) mm wide (Fig. 31); basal parts dying,
attached to the surrounding vegetation or sub-
strates by hapters.
Apothecia very rare (see Discussion).
Chemical reactions and components as in
subsp. sarmentosa but thamnolic acid is always
absent.
Ecology. On well-drained rocky or gravelly
soil at various pH levels; rarely on peaty
soils; usually on rocky ridges or mounds;
high alpine (2150-2800 m) in the Rocky Moun-
tains to sea level in the arctic. However,
one coastal rock outcrop on Vancouver Island
had what appears to be this subspecies.
Distribution. Eastern Canadian arctic, Aleutian
Islands, and the Canadian Rocky Mountains,
with scattered high alpine localities along the
west coast, and in the White, Shickshock
and Otish Mountains of the east (Fig. 30 B).
The strikingly dissected distribution of this
74 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 31. Alectoria sarmentosa subsp. vexillifera. An unusually broad specimen. Quebec, Iles Dormeuses,
Payette s.n. (CANL). Scale 0.5-1 .0 mm.
subspecies requires some comment. A. sarmen-
tosa subsp. vexillifera is a common plant in
southwest Greenland, the mountains of Scan-
dinavia, and the Scottish highlands (Dahl 1950,
Hawksworth 1972), but is virtually absent
from eastern Greenland (Dahl et al. 1937)
and is very uncommon in the central European
mountains (Poelt 1969). It is apparently absent
from Asia and the Southern Hemisphere
(Hawksworth 1972). Such a distribution pattern
probably had its origin as a pre-glacial,
continuous, amphi-atlantic distribution dissected
during the Pleistocene. Remnant populations
could then have radiated from coastal réfugia
(as in Greenland) and from réfugia south or
north of the ice sheets as evidenced, for
example, by the isolated localities in northern
Ellesmere Island (see the comments of Dahl
1950). The eastern population corresponds almost
exactly with Porsild's (1958) "Amphi-atlantic
southern element” matching the distribution
of the willow Salix herbacea very closely.
The western localities also suggest a pushing
of the population southward during the Ice
Age with limited subsequent reinvasion north-
ward. The persistence of the subspecies in the
Aleutians and coastal mountains almost certainly
is due to the presence of coastal réfugia
along the headlands and fiords (see Calder &
Taylor 1968). The entire absence of the sub-
species from the northwestern arctic despite
the many well-documented réfugia of that
area, together with its absence from Asia,
would argue strongly against its ever having
had an amphi-Beringian distribution.
The distinctive distribution of A. sarmentosa
subsp. vexillifera is good evidence for its
genetic basis.
Discussion
The present taxon has been treated by recent
authors as a variety (James 1965), a subspecies
OPERA BOT. 42 (1977)
Alectoria Ach. 75
(Hawksworth 1970, 1972, 1973), and a species
(Krog 1968); Motyka (1964) regarded it as having
no taxonomic importance. The situation here
,s certainly complex since while transitional
morphotypes can be found at least in parts of
Europe (Hawksworth 1973) and Newfoundland
(Ahti & Hawksworth 1974), they appear to be
äbsent in subarctic regions. Indeed a comparison
°f the distribution of vexillifera- morphotypes
with that of sarmentosu -morphotypes lends
considerable support to the distinctness of the
two taxa (see above). The changes in thallus
morphology seen in this subspecies are clearly
adaptive in serving to present the maximum
thallus area to the incident sunlight when
growing with a prostrate habit, but it seems
difficult to determine with any high degree of
certainty that this is always genotypic in origin
f ather than phenotypic. In North America
Particularly, the two taxa are generally well
defined and only very exceptionally show
■my tendency to intergrade. These exceptions
might be due to (a) gene exchange taking place
where the morphotypes are sympatric thus
Producing large series of intermediates, or (b)
s annentosa -genotypes exhibiting phenotypic
convergence with vexillifera -ge not y pe s , thus
forming thalli morphologically similar to them.
* n the absence of transplant experiments it
ls not possible to say which of these ex-
planations is the more nearly correct, but
•he evidence currently available tends to support
•he thesis of two genetically distinct populations
°ccasionally intergrading where they are
s ympatric. As these populations have different
distributional tendencies, the rank of subspecies
appears to be the most appropriate at the
Present time.
As pointed out by Hawksworth (1973), a
lumber of other Alectorioid species occasionally
Produce vexillifera- like morphotypes although
' n all but a few cases (e.g., A. ochroleuca
v ar. ecuadorensis Zahlbr., Sulcaria virens var.
rest'd (D. Hawksw.) D. Hawksw.) the
e gree of flattening and expansion is much
ess than is seen in subsp. vexillifera.
Apothecia are exceedingly rare in subsp.
'ex illifera and have been noted only in a
jangle North American collection (Newfound-
und, Despreaux, 1828 [FH]) which was given
an herbarium name by Delise. In contrast,
Su bsp. sarmentosu is frequently found fertile
and some populations have numerous apothecia
on individual thalli.
Selected specimens
Canada. British Columbia. Flat-Head River Basin:
Flat-Head Provincial Forest, NW of Mt. Rowe,
Bird & Lakusta 17257 (UAC) - Islands South
District: Vancouver Island: Victoria, Ahti 15134
(H) - Alberta. Waterton Lakes National Park,
Bird & Lakusta 16751 (CANL, US, UAC) - Quebec.
Cté Gaspé-Ouest: Mt. Albert, Collins 2656 (US) -
Cté lles-de-la-Madeleine: Grande Entrée, Le Gallo
2999 (US) - Territoire du Nouveau Quebec: 53°20’N,
69°54'W, Guimont QFB-E 4446 (CANL) - Deception
Bay, Eskimo Hill, Gardner 281 (WIS) - lies
Donneuses, Payette s.n., Aug. 1970 (CANL) -
Newfoundland. Labrador North District: Hebron,
Oldenburg 30A (WIS). (See also Ahti & Hawks-
worth 1974) - Northwest Territories. Franklin
District: Diggs (sic) (Digges) Island, Hudson Strait,
Bell 342 (CANL, FH) - Ellesmere Island, 82°24-
32'N, 62°00-32'W, Schuster 35523 (CANL) - Keewa-
tin District: Dubawnt Lake, J. W. Thomson, Larsen &
Foote 14395 (WIS).
U.S.A. Alaska. Aleutian Islands: Amchitka
Island, Reich 111 (CANL, F) - Attu Island, Van
Schaack 293 (US) - Oumak (sic) (Umak Island),
Stevenson s.n.. 24 Nov. 1920 (US) - Unalaska Island,
Eyerdam s.n., 24 May 1932 (ALA. FH. LAM) -
Eastern Pacific Coast District: Sitka, Harbour Moun-
tain. Krog 6265 (O) - Western Pacific Coast District:
Kadiak Island (sic) (Kodiak Island), Trelease s.n.,
2 July 1899 (US) - New Hampshire. Pierce County:
Mt. Rainier National Park, Burroughs Mt.. lmshaug
17371 (US).
8. Alectoria vancouverensis (Gyeln.) Gyeln. ex
Brodo & D. Hawksw., comb. nov.
Bryopogon vancouverensis Gyeln., Feddes Repert.
38: 245 (1935); type: Canada, Vancouver Island.
J. Macoun, 22 July 1897 (UPS holotype!); basionym.
- Alectoria vancouverensis Gyeln., Feddes Repert.
38: 245 (1935); nom. inval. (Art. 34).
Thallus pendent, commonly 15-20 cm, but
sometimes to 1-2 m long (Pike in litt.);
branching mainly isotomic dichotomous, angles
between the dichotomies mainly acute; branches
very uneven in diameter, often flattened
(especially at the axils) and becoming very
twisted and foveolate, 0.4-0. 9 mm diam.; straw-
yellow to yellowish green or greyish green,
usually concolourous.
Medulla compact and dense.
True lateral spinules and soralia absent.
Pseudocyphellae abundant, conspicuous, mostly
raised, often elongate fusiform and twisting
around the branches to give them a conspicuous
76 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 32. Alectoria vancouverensis. Known distribu-
tion.
striped appearance, sometimes confluent, white,
rarely with spinules arising from them.
Apothecia not uncommon, lateral, to 2.6 mm
diam.; excipulum thallinum concolourous with
the thallus, persistent, incurved; discs concave,
dark brown to black. Spores 2-4 per ascus,
ellipsoid, with a hyaline epispore, becoming
brown at maturity, simple, 23-35 x 16-23 /urn.
Pycnidia not seen.
Cortex K — , C-, K.C+ yellow, PD — ;
medulla K-, C + red or rarely C-, KC + red,
PD — . Contains olivetoric and usnic acids and
some unidentified accessory substances; rarely
contains alectoronic acid or barbatic acid in
addition to, or instead of, olivetoric acid.
Ecology. On well-lighted tree limbs, especially
Pseudotsuga menziesii and Tsuga heterophylla,
from an elevation of 30 to 1000 m along the
coast, particularly in humid zones.
Distribution. West coast, lowland, mainly from
southern British Columbia to northern Cali-
fornia (Fig. 32). Its centre of distribution
appears to be the moderately humid to dry
Pseudotsuga-Quercus forests in the southern
Vancouver Island and northern Washington
region corresponding to a ‘‘dry summer, sub-
tropical or mediterranean” climate ("Csb”,
of Köppen's classification; Atlas of Canada
1958 map 30), although it was found in some
localities in the very moist coastal Thuja-
Tsuga forests as well.
Alectoria vancouverensis is endemic to North
America.
Discussion
This species closely resembles A. sarmentosa
in several respects, but differs from the latter
in chemistry, thallus anatomy, colour and size.
Rare specimens can be found which are morpho-
logically more like A. sarmentosa s. str. (i.e. ,
with even, straw-coloured branches having
a more-or-less lax medulla). Specimens also
occur which appear to be A. vancouverensis
due to a dense medulla and twisted branches,
but which have alectoronic rather than olive-
toric acid.
We saw two very odd chemotypes from Cali-
fornia. In addition to alectoronic acid, one
contained abundant thamnolic acid and the
other contained barbatic acid. We named these
specimens A. vancouverensis on morphological
grounds, and because other specimens of A.
vancouverensis exist which have olivetoric-
barbatic, olivetoric-alectoronic, and olivetoric-
thamnolic combinations, giving one the im-
pression that the chemistry is more plastic
than one would expect.
The vast majority of specimens, however,
are easily separated on chemical as well as
morphological grounds. In the field the species
is usually easily spotted as being distinct from
A. sarmentosa.
Two rather odd forms are worth mentioning.
A small caespitose and strigose specimen in
FH collected by John Macoun in June 1915
“on old fences" was given a herbarium name
by G. K. Merrill but the name was apparently
never published. Its medulla was very dense
and C + red and we are regarding it as an
aberrant form of A. vancouverensis.
A specimen which presented us with a more
serious problem was Tavares 371 collected in
Humboldt Co., California. This robust specimen
was so clearly intermediate between A. van-
couverensis and A. imshaugii that for a long
OPERA BOT. 42 (1977)
Alectoria Ach. 77
time we considered naming it as new. The
discussion under A. sarmentosa subsp. sarmen-
tosa will explain why we did not take this
step here. A more complete description of the
specimen is worthwhile, however, in the
possibility that it may be rediscovered else-
where.
Thallus subpendent, 8-20 cm long, stiff,
branching mostly anisotomic dichotomous, with
frequent perpendicular side branches giving
parts of the thallus a “strigose” appearance;
main branches very uneven and twisted, with
an irregular surface, 0.3-0.45 mm diam.
True lateral spinules, isidia, and soredia
absent; pseudocyphellae white, raised, very
conspicuous, often twisting around the filaments
on ridges.
Apothecia and pycnidia unknown.
Cortex K+ yellow, C-, KC + yellow, PD +
orange -yellow; medulla K+ yellow or K — ,
C+ red, KC+ red, PD- or PD + yellow in
part. Contains alectoronic, olivetoric, tham-
nolic, and usnic acids, the first two in the
medulla and the latter two mainly in the cortex.
Selected specimens
Canada. British Columbia. Coastal District: Earl's
Cove, Brodo 8320 (CANL) - Islands South District:
Saltspring Island, Bird 23770 (UAC) - Skeena River
Basin: Skeena River. J. M. Macoun 164 (CANL).
U.S.A. California. Del Norte County: 2.4 miles E
of Gasquet, Tucker 6426 (UC) - Humboldt County:
Near Manila, Stone 37 (HSC) - Pepperwood, Tavares
371 (FH) - Oregon. Josephine County: Between
Cave Junction and O'Brien, Weher 8032 (COLO) -
Lane County: W of Creswell, Pike L-453 (OSC) -
Washington County: Forest Grove. Lloyd s.n., 1894
(F)- Washington. Jefferson County: Olympic National
Park, Hoh Rain Forest, Brodo 13222A (CANL).
Bryoria Brodo & D. Hawksw., nom. nov
Nom. nov. for Setaria Michx., FI. Bor.-Amer.
2: 331 (1803); nom. rejic. (Art. 14). Etymology:
/irv(-opogon) and (Alect-Jor/'a,
Non Setaria Beauvois, Agrost.; 51, 178 (1812);
nom. cons.; type: S. viridis (L.) Beauvois [Gramineae],
Bryopogon Th. Fr., Nova Acta Reg. Soc. Sei.
Upsal., Ser. 3, 3; 25 [repr.J (1860); nom. illegit.
(Art. 64); type: B. jubatum (L.) Th. Fr., lectotype. -
Alectoria subgen. Bryopogon Th. Fr. , Lieh. Scand.
1: 23 (1871). - Alectoria sect. Bryopogon (Th. Fr.)
Zahlbr., Cat. Lieh. Univ. 6: 375 (1930). - Alectoria
sect. Hyalosporae Hue, Nouv. Archs. Mus. Paris,
Sér. 4, i: 86 (1899).
Non Bryopogon Link, Grund. Kraut. 3: 164 (1833);
nom. illegit. (Art. 63); type: B. sarmentosus (Ach.)
Link [=Alectoria sarmentosa (Ach.) Ach.], holotype
(Art. 63).
Type species: Bryoria trichodes (Michx.) Brodo &
D. Hawksw. (syn. Setaria trichodes Michx.; holo-
type).
Thallus fruticose, erect, caespitose, decumbent,
subpendent or pendent; branching variable,
branches generally terete but becoming some-
what compressed, angular and foveolate in a
few species, not becoming markedly expanded
and dorsiventrally compressed; greenish grey
to brown, dark brown or black (and shades of
these colours), some species characteristically
bicoloured.
True lateral spinules with constricted bases
arising at right angles to the main stems in
some species, absent in others. Isidia absent
but isidiiform spinules arising in tufts from
soralia in a few species. Soralia absent to
abundant, tuberculate or fissurai, sometimes
bearing sprouts of secondary branches or tufts
of isidiiform spinules. Pseudocyphellae absent
to abundant, usually rather sparse and in-
conspicuous, elongated and spiralling around
the main stems in a few species, fusiform,
white, yellow or brownish, depressed in most
species but somewhat raised in a few, not
becoming sorediate or bearing spinules. Cortex
composed of periclinal conglutinate hyphae
immersed in moderate or relatively little matrix,
not disintegrating, and smooth or knobby at
the surface. Medullary hyphae not ornamented.
Apothecia lateral, sometimes appearing
geniculate owing to the death of the branch
distal to them, rare or unknown in many
species, frequent to common in others; ex-
cipulum thallinum concolourous with the thallus,
becoming excluded in most species, or if
persistent, not markedly incurved; margin
ciliate in a few species; disc reddish brown
to dark brown, rarely white, never black,
sometimes yellow-pruinose. Asci clavate,
thick-walled, arrested-bitunicate, 8-spored.
Ascospores ellipsoid, lacking a distinct hyaline
epispore, hyaline at maturity, simple, 4.2-15
Atm long. Pycnidia usually rare.
IJsnic acid absent; atranorin and chloro-
atranorin frequent; orcinol tridepsides rare;
/3-orcinol depsidones abundant; pulvinic acid
derivatives rare; orcinol depsides, /3-orcinol
depsides (excluding atranorin and chloro-
atranorin) and hydroxyanthraquinones absent.
Discussion
With some 46 accepted species, Bryoria is
by far the largest segregate of Alectoria s. lat.
and includes species occurring on all continents
of the world in montane, north temperate
and boreal to arctic regions. Some individual
species are very widely distributed and common
whilst others are more localized and rare.
As pointed out above, it is with some
regret that we are introducing here a new
generic name for so many species. As the
original account of Bryopogon Link included
both A. jubata and A. sarmentosa, whether
the earlier name Alectoria Ach. is lectotypified
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 79
by A. jubata or A. sarmentosa. Link’s name
is superfluous and illegitimate under Art. 63.
That is, Bryopogon Link must be lectotypified
by one of these two species selected as the
lectotype of Alectoria Ach. as prescribed in
Art. 7. This would be, in fact, A. sarmentosa,
as pointed out by Hawksworth (1972 p. 197).
As Th. Fries (1860) specifically excluded A.
sarmentosa from Bryopogon. Bryopogon Th.
Fr. has to be lectotypified by one of the
I species he included in it, and A. jubata has
been selected by almost all subsequent authors.
Bryopogon Link and Bryopogon Th. Fr. are
thus nomenclaturally distinct as they are based
on different type species. The latter name is a
later homonym of Link’s and thus illegitimate
k under Art. 64. The employment of Bryopogon
Th. Fr. for the genus named Bryoria here in
the manner proposed by Bystrek (1971) is thus
contrary to the Code and unacceptable.
In our view, it would not be possible to
make a sound case for conserving Bryopogon
Th. Fr. against Bryopogon Link since it could
not be argued that "Bryopogon", at the generic
rank, had actually been widely used in the
sense of Bryoria. Most authors in the past
using this name as a genus have applied the
name to Alectoria s. lat. (including A . sannen-
tosa)', Rabenhorst (1845), Korber (1855) and
Gyelnik (1935) are examples. Some Soviet
authors, however, have followed the usage of
Fries (1860), e.g., Oxner 1956, and Elenkin
and Saviez in many papers. Even if Bryopogon
Th. Fr. was eventually conserved, a large
number of new combinations would still have to
be made since many of the transfers made by
Bystrek (1971) into that genus are invalid
under Article 33.
We decided that the species formerly called
Alectoria americana Mot. would be most ap-
propriate as the type species of the new genus
>n that it exhibits most features of it well
and is not uncommonly fertile. The earliest
epithet available for this species, in fact.
Proves to be Setaria trichodes Michx., the
type species of Setaria Michx. nom. rej.
As pointed out by Hawksworth (1972 p.
197), although Michaux’s name was based on
Lichen tribus Setaria Ach. (Acharius 1798 pp. 4,
219) it must not be cited as “ Setaria (Ach.)
Michx.”, as was done, for example, by Lanjouw
U966 p. 259), since it has been typified by
S. trichodes Michx., a species quite unknown
to Acharius in 1798. We are introducing the
name Bryoria as a nomen novum for Setaria
Michx. nom. rej. typified by the holotype
(monotype) of Michaux’s name, as authorized
by Art. 72.
Setaria Beauvois was first added to the list of con-
served generic names after the Cambridge Congress
of 1930 (Briquet 1935 p. 131) but no type was
designated there for the rejected Setaria “(Ach.)”
Michx. The first selection of a type for the latter
name appears to be that of Rickett & Stafleu (1959 p.
224) whose choice of S. trichodes Michx. has
been adopted in all subsequent editions of the Code.
Acharius (1798 pp. 219-221) definitely accepted
three species in his tribe Setaria, viz. Lichen
jubatus L., L. chalybeiformis L. and L. hippotrichocles
G. Web. As the first two are definitely Bryoria
species (the third probably being a non-lichenized
fungus), even if the selection of S. trichodes were
to be considered a “lapsus” and “corrected" in a
future edition of the Code, the name would still
conform to our concept of Bryoria although the type
would require “correction”.
Most authors since Du Rietz (1926) have
endeavoured to recognize a number of in-
frageneric units within Bryoria as we have
circumscribed it. On the basis of morphological,
anatomical and chemical characteristics, we
are accepting five groups at the rank of section.
In general these sections are quite distinct,
and are based on several unrelated characters
(Table 3). That some of these sections have
geographically separate regions of maximum
diversity suggests that they are indeed natural
groupings. We do not feel that there are, as
yet, adequate distinctions within the sections to
merit their further subdivision.
Bryoria sect. Bryoria
Bryopogon sect. Eujubatae subsect. Lanestres Gyeln.,
Feddes Repert. 38: 223 (1935); type: B. lanestris
(Ach.) Gyeln. [= Bryoria lanestris (Ach.) Brodo &
D. Hawksw.], holotype.
Bryopogon sect. Subfibrillosae subsect. Suh-
fibriUosae Bystr., Ann. Univ. Mariae Curie-
Sklodowska, C. 26: 271 (1971); type: B. fuscescens
(Gyeln.) Gyeln. [= Bryoria fuscescens (Gyeln.)
Brodo & D. Hawksw.], holotype; nom. illegit.
(Art. 66).
? Alectoria subgen. Bryopogon sect. Juhatae DR.,
Ark. Bot. 20A (11): 5 (1926); type: A. jubata (L.)
Ach. [ = B. sp., see Excluded taxa], holotype. -
Alectoria sect. Bryopogon subsect. Jubatae (DR.)
Keissl., Rabenh. Krypt.-FI. 9. 5(4): 97 (1958). -
Bryopogon sect. Eujubatae Gyeln., Feddes Repert.
38: 223 (1935); nom. illegit. (Art. 63).
80 Alectoria in North America
OPERA BOT. 42 (1977)
Table 3. Diagnostic characters separating the five sections of Bryoria. Included are characters of non-North
American species.
Character
Bryoria
Divaricatae
Implexae
Subdivergentes
Tortuosae
Apothecia
Presence
Occasional
Rare
Rare
Common
Occasional
Margins
Yellow pruinose
Eciliate
Eciliate
Eciliate
Ciliate
Eciliate
discs
Absent
Absent
Absent
Absent
Present
Vegetative structure
Cortex
Surface texture
Smooth
Smooth to ±
Smooth
Rough and
Smooth to ±
rough
knobby
rough
Differentiated
superficial layer
Absent
Absent or
virtually so
Absent
Present
Absent
Matrix, relative
to hyphae
Moderate to
Moderate to
Very little to
Moderate
Moderate
abundant
abundant
moderate
Lateral spinules or
spinulose branches
Rare
Present and
Rare
Present but not
Absent
constricted basally
constricted basally
Pseudocyphellae
Present or absent
Present or absent
Always present
Absent
Present or absent
Soralia
Frequent
Occasional
Occasional
Absent
Occasional
Pigmentation
Superficial crust
Superficial crust
Superficial crust
In tips of cortical
cells
Superficial crust
Chemical
components
Atranorin and (or)
chloroatranorin
Occasional
Usually absent
Rare
Absent
Absent
Fumarprotocetraric
acid
Present
Usually present
Very rare
Absent
Absent
Other /3-orcinol
depsidones
Absent
Absent
Rare
Absent
Absent
Orcinol tridepsides
Absent
Absent
Rare
Absent
Absent
Pulvinic acid
derivatives
Absent
Absent
Absent
Absent
Present
Number of species
10
20
12
4
2
Type species: Bryoria trichodes (Michx.) Brodo &
D. Hawksw. (holotype species of Bryoria Brodo &
D. Hawksw.).
Thallus decumbent, subpendent or pendent;
greyish, fuscous, brown to dark brown, not
becoming blackened except in fragmentation
areas. True lateral spinules absent. Isidia absent.
Soralia frequent, fissurai or tuberculate, white
or brownish. Pseudocyphellae present or absent,
usually inconspicuous, fusiform, depressed or
elevated, brown or white. Cortex with a
moderate quantity of matrix material, the
hyphae not markedly fasciculate. Apothecia
frequent in some species, unknown in others;
margins not ciliate; discs orange-brown to
reddish brown. Atranorin and/or chloroatranorin
occasionally present in some species; fumar-
protocetraric acid present in almost all species;
other ß-orcinol depsidones and pulvinic acid
derivatives absent.
Discussion
Bryoria sect. Bryoria comprises eight currently
described species and at least two non-North
American species awaiting description. One may
recognize two distinct elements within it, one
represented by B. trichodes and the other by
the B. fuscescens group. The differences
between these groups are much less than those
separating sections but a treatment in the rank
of subsection perhaps could be justified on
the basis of the presence or absence of
pseudocyphellae.
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 81
Alectoria subgen. Bryopogon sect. Jubatae
1 DR. has been included as a synonym of
this section with some hesitation in view of
the uncertain identity of its type species.
1. Bryoria chalybeiformis (L.) Brodo &
D. Hawksw., comb. nov.
Lichen chalybeiformis L., Sp. PI. 2: 1153 (1753);
type: sine loc., herb. C. Linnaeus (LINN 1273.90
lectotype!); basionym. - Alectoria chalybeiformis
(L.)Gray, Nat. Arr. Brit. PI. 1: 408 (1821).
For further synonyms see Hawksworth (1972 pp.
V 213-216).
Thallus prostrate or decumbent, usually
8—1 5(— 20) cm long; branching irregular, often
) sparse towards the base, isotomic dichotomous
towards the base but becoming anisotomic
dichotomous towards the apices, often en-
tangled, angles between the dichotomies mainly
obtuse; main branches even in diameter, some-
times straight but usually becoming twisted
and foveolate and sometimes channelled,
(0.3— )0.5— 1 ,0(— 2.0) mm diam.; brown to dark
brown or olivaceous-black, usually shiny, some-
times paler in colour towards the base.
True lateral spinules absent but irregular
spinulose branches sometimes present. Soralia
absent or sparse, usually tuberculate, occa-
sionally becoming spinulose, usually narrower
than the branches on which they occur. Pseudo-
cyphellae absent.
Apothecia and pycnidia unknown.
Cortex K — , C — , KC — , PD — ; soralia K. — ,
C — , KC — , PD + red; medulla K — , C - , KC - ,
PD — (or exceptionally PD + red in parts).
Contains fumarprotocetraric acid.
Ecology. Growing over calcareous or non-
c alcareous rocks and soil, and on tundra
vegetation. It is occasionally found on lignum,
and rarely on trees.
Distribution. Hemiarctic to arctic tundra west
and north of Hudson’s Bay; rare in the east,
mainly in exposed and alpine localities (Fig.
33). The occurrence of the species on the Lake
Superior shore is still doubtful, and may be
based on an exceptionally olivaceous speci-
men of Bryoria fuscescens var. positiva which
occurs in the same area. Disjunct alpine
occurrences are common in the western moun-
tains at high altitudes. Hawksworth (1972)
-Opera Botanica nr 42
comments on the world distribution of this
circumboreal and apparently bipolar, arctic-
alpine species.
Discussion
Bryoria chalybeiformis in North America can
usually be reliably recognized on the basis
of its stout, very dark and often almost
olivaceous black, shiny main stems, and the
failure of the medulla to react with PD whilst
the soralia are generally PD + red. At least
some PD + soralia are generally present on a
thallus but in some plants a few may fail to
provide a positive reaction. Some morpho-
types of B. fuscescens var. positiva may
approach it in habit but can in most instances
be distinguished by the duller and paler colour
of the thallus, and by the fact that quite
often, both the medulla and the soralia react
readily with PD. The separation of these taxa is
discussed further by Hawksworth (1972 pp.
214-215).
A further difficulty has arisen in Europe
over the separation of this species and Bryoria
intricans, a taxon of somewhat doubtful validity
(see Excluded taxa). Material corresponding
to B. intricans does not seem to occur in
North America.
In the past North American authors have
widely misapplied the name B. chalybeiformis
when referring to B. furcellata, a very different
species.
Selected specimens
Exsiccatae. Weber: Lieh. Exs. 125: Colorado.
Larimer County: Rocky Mt. National Park, Mount
Chapin. R. A. Anderson 3523 (CANL, COLO, DUKE,
US. WIS).
Canada. British Columbia. Islands South District:
Vancouver Island: Mt. Benson, Macoun 207 (NYBG)
-Coast Area: Garibaldi Park, Krajina 351-4 (CANL) -
Fraser River Basin: Spences Bridge, Macoun s.n.,
28 May 1889 (CANL) - Skeena River Basin:
Hazelton, Ohlsson 2936 (MSC) - Alberta. Morely.
Bird 18510 (CANL) - Saskatchewan. Cypress Hills,
Macoun 448 (CANL) - Manitoba. Kasmere Lake.
Scotter 2857A (WIS) - Ontario. Kenora District:
Pen L, Kershaw s.n., July 1971 (CANL) - Quebec.
Cté lles-de-la-Madeleine: Grand Entrée, Le Gallo
3004 (US) - Cté de Matapédia: Lac au Saumon,
Le Gallo 2157A (US) - Nova Scotia. Halifax
County: Halifax, Taylor 1118 (MSC)- Newfoundland.
Humber East District: Sheffield Lake, Ahti 558
(H) - Labrador North District: Chidley Peninsula,
82 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 33. Bryoria chalybeiformis . N. American distribution.
MacGregor s.n., 1905 (BM) - Yukon. British Moun-
tains: Trout Lake. J. W. Thomson 14782 (W1S) -
Northwest Territories. District of Franklin: Victoria
Island: Cambridge Bay, Weber S 23,810 (COLO,
CANL) - District of Keewatin: Aberdeen Lake,
Scotter 3973 (WIS).
U.S.A. Alaska. Aleutian Islands: Amchitka
Island, Schacklette 8013 (COLO) - Unalaska, J. M.
Macoun 202 (CANL) - Arctic Coast District:
Colville R. at Umiat, J. W. Thomson & Shushan
6198 (WIS) -Arizona. Graham County: Mt. Graham,
Nash 4211 (NASH) - California. County unknown:
Pak?low Hills, Blander 161 (US) - Colorado.
Boulder County: Summit of Rollins Pass, Shushan
& Anderson S 25,716 (COLO) - Michigan.
Keweenaw County: Isle Royale, Raspberry Island.
J. W. Thomson 17233 (WIS) - Washington. Clallam
County: Hurricane Ridge, Olympic National Park.
Brodo 13197 (CANL).
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 83
Fig. 34. Bryoria fuscescens var. positiva showing the pale basal area. Newfoundland, Ahti 560 (CANL). Scale
0. 5-1.0 mm.
2. Bryoria fuscescens (Gyeln.) Brodo & D.
Hawksw., comb. nov.
A lectoria fuscescens Gyeln., Nyt Mag. Naturvid. 70:
55 (1932); type: Finland, Tavastia austr., Hollola,
ad truncos Pini locis apricioribus in silva, J. P . Norrlin,
September 1882, Nyl. & Norrl.. Lieh. Fenn. Exs. no.
466 (BP 33.947 leclotype! BM. CANL, H isolecto-
lypes!); basionym.
Bryopogon paciftcus Gyeln., Acta Geobot. Hung.
2: 166 (1937); type: Oregon. Benton Co., on apple
trees in old orchard, F. P. Sipe (ORE lectotype!).
For further synonyms see Hawksworth (1972 pp.
217-219).
Thallus pendent (to prostrate in some morpho-
types of var. positiva), usually 5— 15(— 30) cm
long; branching anisotomic or isotomic dichoto-
mic at the base, often becoming anisotomic
dichotomous towards the apices, frequent from
the base, angles between the dichotomies
acute or obtuse, sometimes with lateral spinu-
lose branches; branches even to uneven in
diameter, usually straight but sometimes becom-
ing twisted and occasionally foveolate (in var.
positiva), (0.2— )0.3— 0.4(— 0.6) mm diam. ; pale fus-
cous to brown or blackish, the basal parts
usually much paler in colour than the apices
(Fig. 34); black fragmentation regions often
present.
True lateral spinules and pseudocyphellae
absent (note that young fissurai soralia may
resemble pseudocyphellae). Soralia abundant to
sparse, tuberculate or fissurai, some fissurai
soralia always present, usually broader than the
branches on which they occur, sometimes be-
coming spinulose.
Apothecia and pycnidia not seen in North
American material.
Cortex K-, C-, KC-, PD+ red or -;
soralia K-, C-, KC-, PD + red; medulla K-,
C— , KC-, PD+ red at least in parts or -, Con-
tains fumarprotocetraric acid, with chloroatra-
norin as an accessory substance (only in speci-
mens east of the Rocky Mountains). The
84 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 35. Bryoria fuscescens s. lat. N. American distribution.
isotype in CANL contained fumarprotocetraric
acid and atranorin s. lat., and the cortex of the
filaments was PD+ red in part.
Ecology. On trees, mostly coniferous, in Picea
glauca-Abies balsamea forests in the east;
Picea mariana forests in the north; Pinus
contorta, P. ponde rosa, and Pseudotsuga
forests in the west. Well developed on Betula
papyrifera in northern Michigan.
The var. positiva occurs on rocks, lignum, and
trees, generally in highly exposed habitats.
Distribution. Throughout the boreal forest zones
of North America from Newfoundland to
Alaska, north to the tree-line (Fig. 35). It ranges
southward in the west along the Rocky Moun-
tains to New Mexico and into the Arizona moun-
tain systems, and, to some extent, along the
Coast Range. In the east, it shows a preference
for the more humid coastal localities and milder
central Newfoundland forests.
B. fuscescens is widespread in the Northern
Hemisphere, and is the most common Bryoria
in the lowland areas of northern Europe and the
British Isles (Hawksworth 1972). It also occurs
in the East African mountains (Hawksworth
1971 a, Krog & Swinscow 1975).
Discussion
There is no species group in which there are
more problems in delimiting taxa than those
OPERA BOT. 42 (1977)
B ry or ici Brodo & D. Hawksw. 85
species of the section Bryoria which generally
have PD+ red soralia. “Intermediate” speci-
mens can be demonstrated from almost any pair
within the group, although the distinctness of
the taxa is unquestioned in most cases. Few
characters of any of the taxa are constant for
all specimens and variability is exceptionally
broad. Environmental modifications may be the
cause of at least some of this variability.
Most plastic in morphology and chemistry is
Bryoria fuscescens (Bystrek 1963. Hawksworth
1972, 1973). Its “forms” and "varieties" are so
abundant that there is a danger of circumscribing
it in a way similar to the former use of the
name “Alectoria jubata" , with almost every
puzzling Bryoria being conveniently placed
within its all-encompassing boundaries.
The most easily identified specimens of B. fus-
cescens, those having all the most characteristic
features of the species, are as follows: pendent;
dark fuscous brown except at the base which is
much paler, often almost white; having a mixture
of fissurai and tuberculate soralia; the outer
portion of the cortex PD+ red; the branches
about 0.2-0. 4 mm in diameter, more or less even
except for the basal branches which can be
flattened and somewhat twisted. Bryoria glabra
is “typically” dark olivaceous and shiny with
very even, broadly angled, and strictly dichoto-
mous branches (especially at the base), about
0.2-0. 4 mm in diameter: soralia are small, oval,
and fissurai. In southern Alberta and southern
British Columbia, the two seem to intergrade to
some extent.
In the western mountains, B. fuscescens
develops a morphotype with abundant fis-
surai, often quite spinulose, soralia. It is usually
pale fuscous, although sometimes extremely
dark, and has a paler base. Otherwise, it resem-
bles B. vrangiana in many ways. This pale,
spinulose B. fuscescens probably can be as-
signed to f. pallida (Saviez) D. Hawksw.
(Hawksworth 1972 p. 218). We are tentatively
regarding B. vrangiana as a distinctly dark,
olivaceous species with sparse fissurai soralia
(which often are absent altogether). The plant
is quite thick at the base (often over 0.5 mm
diam.) and is strongly foveolate, twisted, and
spinulose. Räsänen’s Lieh. Fenn. Exs. no. 23 is
slightly more pendent and slender than the North
American specimens, but is a good match in
other respects. Brown (non-olivaceous) speci-
mens named as B. vrangiana in Europe as well
as North America due to their foveolate, twisted
branches and spinulose soralia are usually either
B. fuscescens f. pallida or var. positiva.
As can be seen from the above circumscrip-
tion of B. vrangiana, the species comes very
close to resembling B. chalybeiformis. In fact,
B. chalybeiformis and B. vrangiana differ
largely in their habitat and distribution with
predictable differences in growth form and habit.
B. vrangiana is found on trees and stumps in
subalpine and subarctic localities, and is pendent
with branches forming acute angles; B. chaly-
beiformis grows over soil, vegetation, and rocks
mainly in alpine and arctic localities, and is, not
surprisingly, prostrate with branches forming
broad, obtuse angles. The only other distinction
worthy of note is that the soralia of vrangiana
are largely or entirely fissurai, and those of
B. chalybeiformis are commonly tuberculate.
In any discussion of B. chalybeiformis, one
must return to B. fuscescens for a consideration
of its variety positiva. These two taxa have
already been compared in Britain by Hawks-
worth (1972) but a reexamination of the problem
is warranted in view of the distinctive and abun-
dant North American material. Whereas in the
British Isles, var. positiva could be distinguished
from B. chalybeiformis by basal colour, branch
diameter, and type of soralia, this cannot be
done with the North American specimens. Both
taxa here commonly have pale bases, thick (over
0.5 mm) branches and both fissurai and tuber-
culate soralia. B. fuscescens var. positiva is
largely a temperate taxon occurring on rocks
and frequently on trees. It is always brown to
pale fuscous (especially the pendent form) and
commonly has a PD+ red cortex or medulla.
B. chalybeiformis is almost always olive-black
to olive-brown, at least in part, and is a terri-
colous and saxicolous species of arctic-alpine
situations. The cortex and medulla is PD- with
very rare exceptions.
For discussions of the distinctions between B.
fuscescens, B. lanestris and B. subcana, see the
comments under the latter two species.
As regards its chemistry, B. fuscescens is not
entirely uniform throughout its range. Chloro-
atranorin occurs fairly regularly in material east
of the Rocky Mountains but appears very
rarely in Europe and in the west. On the other
hand, neither the intensity nor the anatomical
86 Alectoria in North America
OPERA BOT. 42 (1977)
position of the PD reaction has any geographi-
cal correlation.
Unfortunately, it appears that the taxonomic
problems in this group cannot be resolved on the
basis of the North American material alone and
must await a critical study of the group on a
world scale.
Selected specimens ( var. fuscescens)
Canada. British Columbia. Columbia River Basin:
Revelstoke, Shaw 977 (US) - Coastal Region: Alice
Lake Provincial Park, Ohlsson 740B (MSC) - Fraser
River Basin: S of Clearwater, Brodo 15767 (CANL)-
Islands North District: Graham Island: Marian Lake,
Brodo 18474 (CANL) - Islands South District: Van-
couver Island: Ladysmith, Ahti 15384 (H) - Kootenay
River Basin: Fernie, Brodo 15076 (CANL) -Okanagan
District: Princeton, Brodo 7815 (p.p.) (CANL) -
Skeena River Basin: 30 miles W of New Hazelton,
Ohlsson 2887B (MSC) - Alberta. Bow River Water-
shed, Wildcat Hills, Bird 9924 (WIS) -Saskatchewan.
Cypress Hills, Lamb 6128 (DUKE, CANL, WIS,
NYBG) - Waskesiu, Jesberger 745 (SASK) - Mani-
toba. Zed Lake, Larsen s.n., 10 Aug. 1959 (CANL) -
Ontario. Cochrane District: Moosonee, Brodo 14784
(CANL)- Winisk, Webber W 22B (CANL) -Thunder
Bay District: Sibley Provincial Park, Denison 645 (US,
CANL) - Quebec. Bonaventure Island, Brodo 1 869 1C
(CANL) - Cté Pontiac: Parc de la Verendrye, Brodo
17028 (CANL) - New Brunswick. Albert County:
Fundy National Park. Ireland 11332 (p.p.) (CANL) -
Nova Scotia. Halifax County: Peggy's Cove, Taylor
1487 (MSC) - Prince Edward Island. Queens County:
West Barkley Beach, Fabiszewski s.n., 20 July 1970
(CANL) - Newfoundland. Humber East District:
Upper Humber Big Falls, Ahti 572 (H) - Labrador
West District: Menihek Lake, Brodo 8825-5B (WIS,
CANL) - Yukon. Dawson, Williams 71 (NYBG) -
Northwest Territories. Mackenzie District: Mosquito
Lake. Oldenburg 46-1534 (WIS).
U.S.A. Alaska. Central Yukon River District: White
Mts., Krog 2767 (O) - Arizona. Apache County:
Trail to Mt. Baldy, Nash 7838 (p.p.) (CANL) -
Coconino County: San Francisco Peaks, Nash 7489
(CANL) - Graham County: Pinatino Mts., Darrow
1780 (WIS) - Pima County: Santa Catalina Mountains,
Nash 4010 (CANL, ASU) - California. San Mateo
County: Pilarcitos Creek Canyon, Herre & Doty 3348
(NYBG, F) - Santa Cruz County: Santa Cruz Mts.,
Black Mt., Herre s.n. (NYBG) - Colorado. Latimer
County: Rocky Mt. National Park, Bierstadt Lake,
Anderson 2507 (COLO) - Idaho. Latah County: N of
State Route 8, Cooke 23047 (WIS) — Maine. Washing-
ton County: Quody Head near Lubec, Taylor I073B
(MSC) - Massachusetts. Hampden County: Spring-
field, Farlow s.n.. May 1878 (NYBG) - Michigan.
Chippewa County: 1 mile E of Makina County,
Imshaug 3351 (MSC) - Montana. Granite County:
Clinton, Norris 2072C (HSC) — New Hampshire .
Crafton County: White Mts., Faxton s.n., 12 June
1879 (FH) - White Mt. Region, Tuckerman s.n.
(NYBG) - New Mexico. Sandoval County: Sandia
Mt., Ellis s.n., 1914 (FH) - Taos County: Très
Piedras, Shushan & Weber S-6533 (COLO, ABSL) -
Oregon. Josephine County: Rough and Ready Creek
Botanical Wayside, Pike L-234 (PIKE) - South
Dakota. Lawrence County: Bower Intake, Wetmore
9020 (MSC) - Washington . Pierce County: Mt Rainier
National Park, Burroughs Mt., Imshaug 170C (LAM)
- Wyoming. Pennington County: Black Hills, 4 miles
W of US 385, Wetmore 8865 (BM).
Selected specimens (var. positiva)
Canada. Alberta. Cypress Hills Provincial Paris,
MacNeil 1274 (CANL) - Manitoba. Fort Churchill,
J. W. Thomson 3822 (CANL) - Ontario. Kenora
District: Cape Henrietta Maria, Cowell 1287 (CANL)-
Thunder Bay District: Longcroft I. , Garton 625 1 (WIS)
- Quebec. Cté Gaspé-Est: Bonaventure Island, Cain
27403 (US) - New Brunswick. Charlotte County: Cam-
pobello, collector unknown, July 1902 (FH) - New-
foundland. Grand Falls District: Great Rattling Brook
road, Ahti 578 (H) - Labrador South District: Red
Bay, Waghome s.n., 6 Aug. 1894 (BM).
U.S.A. Alaska. Bering Sea District: St. Paul Island.
J. M. Macoun 447? (624?) (CANL) - Bering Strait
District: King Island, Krog s.n., 27 Aug. 1969 (O) -
Arizona. Coconino County: Mt. Agassiz, Brodo
20350 (CANL) - Maine. Hancock County: Salisbury
Cove (sic) (Salsburg Cove), Mount Desert Isl., Plitt
26a (US) - Massachusetts. Hampden County: Spring-
field, Herb. W. G. Farlow s.n.. May 1878 (FH) -
Michigan. Keweenaw County: Sand Bay, W of Eagle
Harbour, J. W. Thomson 2799 (WIS) - Oregon. Ben-
ton County: Corvallis. Plitt? 19 (US) - Washington.
Kittitas County: Easton, Howard 837 (FH).
3. Bryoria glabra (Mot.) Brodo & D. Hawksw.,
comb. nov.
Alectoria glabra Mot., Fragm. Florist. Geobot. 6: 448
(1960): type: U.S.A.. Washington, Olympic Peninsula,
Clallam Co., Hurricane Ridge, alt. 5,800 ft, on trunk
of Abies lasiocarpa, B. I. Brown & W. C. Muenscher
129, 24 July 1950 (US holotype!); basionym. -
Bryopogon glaher (Mot.) Bystr., Ann. Univ. Mariae
Curie-Skfodowska, C, 26: 271 (1971; nom. inval.
(Art. 33).
Thallus pendent, usually 10-15 cm long; branch-
ing regularly isotomic dichotomous, angles
between the dichotomies usually obtuse and
rounded; branches even in diameter, straight,
not becoming twisted or foveolate, 0.2-0. 4 mm
diam.; olive brown to greenish black throughout,
shiny, often with a translucent appearance.
True lateral spinules and pseudocyphellae
absent. Soralia present, fissurai or very rarely
tuberculate, sparse to abundant, usually nar-
rower than the branches on which they occur,
white, to 1 .0 mm long.
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 87
Fig. 36. Bryoria glabra. Known distribution.
Apothecia rare, lateral 0.65-1.15 mm diam.;
excipulum thallinum concolourous with the
thallus or darkening to disc colour, persistent,
even, smooth, becoming somewhat reflexed in
age; disc concave to plane, becoming convex
when old, pale red-brown to dark fuscous.
Asci 27-30x9-10 p,m; no mature spores seen.
Pycnidia unknown.
Cortex K— , C-, KC-, PD — ; soralia K-,
C— , KC-, PD+ red; medulla K-, C-, KC-,
PD — . Contains fumarprotocetraric acid.
Ecology. On coniferous trees, especially Abies
iasiocarpa and Picea in the subalpine zone of
niountains; also on Picea sitchensis, Tsuga
heterophylla , and Pinus contorta and associated
trees on the coast, typically around bogs.
Distribution. Along the west coast from northern
California to Alaska, extending eastward into the
northern Rocky Mountains (Fig. 36). Disjunct
populations exist in the Gaspé area and central
Newfoundland (see general section on Distribu-
tion, category 11). Like B. friabilis and B. pikei,
with bicoastal disjunctions, B. glabra is a North
American endemic.
Discussion
Although only proposed relatively recently,
this name has been used in a rather broad
sense by North American lichenologists, and has
often included plants clearly referrable to B.
fuscescens. In the course of the present in-
vestigation we have restricted the name
“glabra" to specimens with shiny, smooth
branches which have regularly isotomic dichoto-
mous branching with broadly rounded axils,
and have fissurai soralia. Such specimens are
88 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 37. Bryoria tanes-
tris on a spruce twig,
showing rather typical
“wool-like" appearance.
Quebec, Schefferville,
Brodo 9125a (CANL).
Scale 0. 5-1.0 mm.
mainly confined to an area from the Rocky
Mountains westwards, with their greatest
abundance in the Coast Range and along the
British Columbia coast; they are extremely
rare on the east coast.
Bryoria glabra is usually fairly regularly sore-
diate but rare specimens have been found with
very few or no soralia. Such specimens super-
ficially resemble B. trichodes subsp. americana
except in colour. They can be easily distin-
guished, however, by the consistently PD -
medulla ofß. glabra and the presence of pseudo-
cyphellae in subsp. americana (although these
may be rare in some specimens and easily over-
looked). The separation of B. glabra from B.
fuscescens can present a number of difficulties;
these are discussed with the latter species.
A peculiar specimen found in Washington,
containing gyrophoric acid but with some
characters of B. glabra, is discussed under
B. friabilis.
Selected specimens
Canada. British Columbia. Columbia River Basin:
Columbia R. below Revelstoke, Macoun 53 (CANL)
- Coastal Region: Garibaldi, Brodo 8182 (CANL) -
Fraser River Basin: Wells Gray Provincial Park, Ahti
13044 (H) - Islands North Region: Port Clements,
Brodo 9780 (CANL) - Islands South District: Powell
River, Tusko 8015 (H)- Okanagan Region: Tulameen,
Ahti 10307 < H ) - Skagit River Basin: Manning Park,
Ahti 15387 (H) - Skeena River Basin: 9 Mile Moun-
tain, east of Hazelton , Ohlsson 2962A (MSC.CANL)-
Alberta. Banff National Park, trail to Lake Agnes,
Imshaug 6866 (MSC) - Quebec. Cté Gaspé-Est:
Bonaventure Island, Shushan S 23,039 (NYBG) -
New Brunswick. Restigouche County: Dalhousie,
Reijonen s.n., 16 Nov. 1930 (H) - Newfoundland.
(See Ahti & Hawksworth 1974.)
U.S.A. Alaska. Eastern Pacific Coast District:
Juneau, Mt. Roberts, Krog 5530 (O) - Central Yukon
River District: White Mts., Lion Peak, Krog 2758 (O)
- Indian Village, Krog s.n., 17 June 1969 (O) -
California. Del Norte County: Near Bear Basin,
Tucker 6435 (COLO) - Idaho. Bonner County: Gis-
borne Mt., Priest R. Experimental Forest, Esslinger
377B (COLO) - Montana. Glacier County: Glacier
National Park, near Anaconda Creek, Imshaug 7591
(WIS) - Oregon. Klamath County: O’Dell Lake,
Howard 4733 (COLO) - Washington. Clallam County:
Hurricane Ridge, Muenscher 129 (NYBG, US).
4. Bryoria lanestris (Ach.) Brodo & D.
Hawksw., comb. nov.
Alectoria jubata £. lanestris Ach., Lieh. Univ.:
593 (1810); type: Helvetia (H-Ach. holotypel); basio-
nym. - Alectoria lanestris (Ach.) Gyeln., Nyt Mag.
Naturvid. 70: 58 (1932).
Bryopogon negativus Gyeln., Acta Geobot. Hung.
2: 164 (1937); type: Oregon, Crater Lake, on trees,
F. P. Sipe 682, August 1930 (ORE lectotype!).
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 89
Alectoria tenerrima Mot., Bryologist 67: 31 (1964);
type: British Columbia, Southern Caribou Mts.,
Wells Gray Park, south-west end of Turtle Lake, on
old branches of Picea engelmannii in swamp, T. Ahti
6512, 25 June 1961 (H holotype!). - Bryopogon
tenerrimus (Mot.) Bystr.. Ann. Univ. Mariae Curie-
Skfodowska, C, 26: 274 (1971); nom. inval. (Art.
33).
For further synonyms see Hawksworth (1972 p.
222 ).
Thallus pendent to subpendent, 5—1 0(— 1 5) cm
long; branching irregularly isotomic dichoto-
mous to anisotomic dichotomous, frequent from
the base, angles between the dichotomies
usually acute; branches usually very uneven in
diameter, straight, sometimes becoming com-
pressed toward the base, very brittle and tending
to fragment in herbarium packets, 0. 1— 0.25(— 3.0)
mm diam.; brown-black to olivaceous black
or black, usually concolourous, dull or rarely
becoming shiny (often in parts). Fig. 37.
True lateral spinules and pseudocyphellae
absent. Soralia sparse to abundant, fissurai,
white or white speckled with black, to 0.3 mm
long.
Apothecia and pycnidia unknown.
Cortex and medulla K— , C — , KC-, PD— ;
soralia K-, C-, KC-, PD+ red. Contains
fumarprotocetraric acid.
Ecology. On coniferous trees, especially Picea
mariana, P. glauca, Pinus banksiana, and
Larix laricina (rarely on willows and birches) in
boreal black spruce forests, fens and bogs, and
Pinus conforta or Picea engelmanii in the
western mountains. (See comments under
Biyoria simplicior.) Rarely, it is found on rocks
or even soil.
Distribution. Mainly in the northern boreal zone
from Alaska to Labrador, and in the western
mountain ranges south to Colorado, New
Mexico and Arizona (Fig. 38). B. lanestris seems
to prefer the more continental forest regions,
although there are a number of occurrences in
the coastal ranges, especially in British Colum-
bia. This is evidently a circumboreal species.
Although not confirmed for Asia by Hawks-
Worth (1972), we have now seen it from several
sites in northern Mongolia collected by Nina
Golubkova in 1972 and 1974 (e.g., Golubkova
138, 787, 789 [LE]) and it is undoubtedly also
Present in Kamchatka and Siberia. We have seen
a collection from Hawaii (Maui, Haleakala
National Park, Bowler 1645 [BM, CANL, 1MI])
which appears to belong to B. lanestris. Gyel-
nik’s (1935) reports of this species from Algeria
and Mexico must be treated as dubious;
Japanese references to B. lanestris mainly refer
to B. trichodes subsp. trichodes.
Discussion
Bryoria lanestris is normally a well-marked
species which can be separated from B. fusees-
cens by its very dark colour, narrower branches
which are characteristically uneven in diameter
and extremely brittle (readily fragmenting in
herbarium packets); the cortex in B. lanestris
is also always PD-, whilst in B. fuscescens,
the outer cortex sometimes reacts PD+ red.
B. lanestris often grows mixed with B. simpli-
cior and the two thalli can be intricately en-
tangled. B. simplicior is readily separable by the
greenish black colour of the soralia, more even
branches, and the failure of the soralia to react
with PD.
In North America B. lanestris usually has fre-
quent fissurai soralia. Tuberculate soralia, which
occur in some European collections, appear to
be absent here. Specimens which are almost
esorediate also sometimes occur. These varia-
tions appear to be of little taxonomic im-
portance.
Selected specimens
Exsiccatae. Merrill: Lieh. Exs., Ser. II, 129 (mixed
collection): Alaska, Fairbanks, Palmer (US) - Thom-
son: Lieh. Arct. 88: Northwest Territories, District
of Mackenzie: Artillery Lake, J. W. Thomson &
Larsen (ALA. CANL. COLO, DUKE) - Weber:
Lieh. Exs. 431: Wyoming, Park County: SilverGate,
Weber (CANL).
Canada. British Columbia. Columbia River Basin:
Glacier National Park, Ohlsson 395 (MSC, CANL) -
Coastal Region: Pemberton, Garibaldi Mountains,
Brodo 8269 (CANL) - Dean River Basin: Tweedsmuir
Provincial Park. Ohlsson 1899 (MSC, CANL) - Fraser
River Basin: Kamloops Tearsdale 3 (W1S) - Okanagan
District: Okanagan Lake, Maslin s.n., 14 Aug. 1952
(COLO) - Skagit River Basin: Manning Park, Otto
105A (H) - Skeena River Basin: 30 miles W of New
Hazelton, Ohlsson 2887B (MSC) - Alberta. Bragg
Creek, Bird 10015 WIS) - Saskatchewan . South Black
Lake, Scotter 66 (WIS) - Charcoal Lake, Scotter
2684 (WIS) - Prince Albert National Park, Jesberger
1171 (SASK) - Manitoba. Churchill, Masson 9854
(QUE, WIS) - Ontario. Cochrane District: Gardiner,
Wetmore 5765 (US) - Thunder Bay District: Niblock.
Cain 26417 (CANL, TRTC) - Quebec. Parc des
90 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 38. Bryoria lanestris. N. American distribution.
Laurentides: Riv. Montmorency. Masson 9982 (WIS)-
Territoire de Mistassini: Rupert House, Kucyniak &
Tuomikoski 51A (H) - Newfoundland. Central
Labrador: Mary River, Hustich s.n., 26 Aug. 1937
(H) - Labrador North District: Goose Bay, Murray
& Pruitt 1652 (ALA, H) - Yukon. Mt. Sheldon,
Porsild & Breitung 2297 (DUKE. W1S, US, NYBG.
CANL) - Semenof Hills, Scotter 19744 (CANL) -
Northwest Territories. District of Mackenzie: W shore
of Liard R., Fort Liard, Cody 11617 (CANL) -
Keewatin District: Coral Harbor, Southampton Island,
Weber S23, 687 (p.p.) (US) - Thelon River, Aberdeen
Lake, Scotter 4236 (W1S).
U.S.A. Alaska. Alaska Range District: Wonder
Lake, Mt. McKinley Nat. Park, Weber & Viereck
S 7259 (DUKE, COLO, MSC, ORE. WIS) - Arctic
Coast District: Valley of Mancha Creek and Firth
River, Sharp 6532 (DUKE, US, MSC, WIS )- Arizona.
Coconino County: San Francisco Peaks, Nash 7525
(CANL, NASH) - Colorado. Grand County: Fraser
Exp. Forest, Weber & Dahl s.n., I Aug. 1953 (COLO)
- Hinsdale County: trail to Upper Powderhom Lakes,
Weber s.n., 28 July 1964 (COLO) - Larimer County:
Rocky Mt. Nat. Park, Wild Basin, Kiener 1695
(COLO) - Idaho. Fremont County: Targhee Pass,
Cain 26415 (VS) -Maine. Penobscot County: Bangor,
Merrill s.n., 23 May 1896 (US) - Montana. Gallatin
County: Above Gallatin R. on US 191, Norris 2054D
(HSC) - New Mexico. Taos County: Très Piedras,
Shushan S6533 (US) - Oregon. Benton County:
Near Corvallis, Plitt s.n.. Mar. 1925 (US) - Klamath
County: Crater Lake, Sipe 682 (ORE) - Lane County:
Salt Creek Falls, Doty 5271 (NYBG) -South Dakota.
Pennington County: 24 miles SW of Lead, Wetmore
1 1702 (MSC)- Washington. Whitman County : Steptoe
Butte, Esslinger 1737b (WIS) - Wyoming. Teton
County: Moran, Cain 26034 (US).
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 91
5. Bryoria subcana (Nyl. ex Stiz.) Brodo &
D. Hawksw., comb. nov.
Alectoria prolixa var. subcana Nyl. ex Stiz., Ann.
Naturhist. Mus. Wien 7: 129 (1892); type: Scotland,
J. M. Crombie. 1875 (H-Nyl. 35835 lectotype! BM
isolectotype!); basionym. - Alectoria subcana (Nyl.
ex Stiz.)Gyeln., Magy. Bot. Lapok 30: 54 (1931).
For further synonyms see Hawksworth (1972 pp.
249-250).
Thallus pendent to subpendent, to 5 cm long
(in North American material); branching iso-
tomic dichotomous, angles between the dicho-
tomies mainly rounded, 80-90°; branches terete,
even in diameter, straight, often brittle, 0.15-
0.3 mm diam.; basal parts pale fuscous brown,
usually matt but occasionally slightly shiny,
apical parts very pale fuscous to greenish white
or whitish, sometimes becoming variegated
(probably an insolation effect); base persistent.
True lateral spinules absent. Pseudocyphellae
often present, sparse, inconspicuous, fusiform,
white. Soralia usually abundant, tuberculate, as
wide as or slightly broader than the branches on
which they occur, occasionally becoming
spinulose, to 0.8 mm diam.
Apothecia and pycnidia not seen in North
American material.
Outer cortex, medulla and soralia K— , C — ,
KC-, PD+ bright red (rapid). Contains large
amounts of fumarprotocetraric acid.
Ecology. On Picea along coastal bays and
breams in the north, and on conifers and lignum
on ridges and sand hills in California.
Distribution. Along the coasts of Alaska and
British Columbia, with a disjunct population in
the Santa Cruz area of California (Fig. 39).
This basically oceanic species has a European
distribution very similar to that of B. smithii
(see maps in Hawksworth 1972 p. 248). The
species is unknown outside North America and
Europe.
Discussion
Eryoria subcana is most closely allied to B.
fuscescens from which it can be distinguished
hy its paler colour and generally extremely
strong PD+ cortical reaction. These two species
"'ere also found to differ by Hawksworth
((972) in that B. subcana never tends to be paler
at the base, often has blackened fragmentation
Fig. 39. % Bryoria subcana. N. American distribution.
▲ Bryoria spiralifera. Known distribution.
regions, has only tuberculate soralia, and is
more regularly fertile. As this species is known
in North America only from a few specimens
several of which are poorly developed, it would
be premature to comment on its variability
on this continent.
Atranorin s. lat. occurs together with fumar-
protocetraric acid in small amounts in at least
some European material of this species.
B. subcana has generally been a poorly
understood species in Europe although it has
been accepted in the British Isles for over a
century. It has most frequently been subsumed
92 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 40. Irregular soralia found on the sorediate
morphotype of Bryoria trichodes showing the charac-
teristically reflexed filaments. Quebec, Gaspé, Brodo
18694 (CANL). Scale 0.5 mm.
into B. capillaris by Scandinavian lichenologists.
More recently Motyka and Bystrek have been
applying this name to B. implexa. See the com-
ments in Hawksworth (1972 p. 250).
Several specimens of a very odd morphotype
(virtually caespitose, with large, empty
soralium-like fissures; somewhat spinulose)
were seen from the San Francisco area of Cali-
fornia and Oregon. They are only provisionally
referred to B. suhcana based on their very pale
colour and strong PD+ red outer cortex (see
Discussion under B. furcellata). More typical
B. suhcana is known from the same area.
Reports of#, suhcana from British Columbia
(Motyka 1964), Ontario (Ahti 1964) and Alaska
(Krog 1968, sub Alectoria haynaldii) based on
specimens determined by Motyka have proved,
on reexamination, to be errors for other species,
most commonly B. capillaris.
Specimens
Canada. British Columbia. Coastal District: Namu
Harbor at the west end of Burke Channel, Ohlsson
2082 (MSC) - Islands North District: Langara Island,
Persson 34, 38A (S) - Lyell Island, Brodo 11800
(CANL).
U.S.A. Alaska. Central Pacific Coast District:
Knight Island, Prince William Sound, Eyerdam (244 )A
(FH) - Eastern Pacific Coast District: Washington
Bay, Kuiu Island, Eyerdam 856 (W1S) - California.
San Mateo County: San Francisco Watershed, Cahill
Ridge, Jordan WJ956B (WIS. SFS) - Santa Clara
County: Santa Cruz Mountains above Saratoga, Herre
491 (FH) - Santa Cruz, Anderson 530 (FH) - Near
Ben Lomond, Thiers et al. 9139 (WIS) - Oregon.
Clatsop County: Saddle Mtn. State Park, summit.
Pike 3818 (OSC).
6 a. Bryoria trichodes (Michx.) Brodo &
D. Hawksw., comb. nov. subsp. trichodes
Setaria trichodes Michx., FI. Bor. -Am. 2: 331 (1803);
type: Canada, ad ramulos putridos (PC holotype!);
basionym. -Alectoria trichodes (Michx. )D. Hawksw..
Lichenologist 5: 252 (1972).
Alectoria jubata f. minuscula Merr., Bryologist
14: 36 (1911); type; Nova Scotia, near Digby, on
spruces, J. Macoun, 20 May 1910 (FH lectotype!).
Alectoria canadensis Mot.. Bryologist 67: 35 (1964);
type: Ontario, Thunder Bay District, Slate Islands on
north shore of Lake Superior, small island on south-
east side of Edmonds Island, 48°40' N, 87°01' W,
on Abies balsatnea in rich forest, abundant, T. Ahti
4369, 10 July 1958 (H lectotype!); nom. inval. (Art.
37). - Bryopogon canadensis (Mot.) Bystr., Ann.
Univ. Mariae Curie-Sklodowska, C, 26: 271 (1971);
nom. inval. (Arts. 33, 37).
Alectoria delicata Mot., Bryologist 67: 34 (1964);
type: Alaska, Sitka, Sitka National Monument, H.
Krog 6254. 1 September 1957 (O lectotype!); nom.
inval. (Arts. 36, 37). - Bryopogon delicatus (Mot.)
Bystr., Ann. Univ. Mariae Curie-Sklodowska, C,
26: 274 (1971); nom. inval. (Arts. 33, 36, 37).
Alectoria cervinula var. eciliata Mot., in Krog.
Norsk Polarinst. Skr. 144: 135 (1968); type: Alaska.
Central Pacific Coast District, Richardson Highway,
mile 9, H. Krog 2176, 18 August 1957 (O lectotype!);
nom. inval. (Arts. 32, 36, 37).
Thallus pendent, subpendent, or prostrate,
usually 7— 15(— 20) cm long; branching anisotomic
dichotomous (to isotomic dichotomous in subsp.
americana), frequent from the base, angles
between the dichotomies mainly acute (to obtuse
and rounded in subsp. americana), occasionally
with short perpendicular lateral spinulose
branches; branches terete, uneven (to even in
subsp. americana), 0.2-0. 3 mm diam. at the
base, main branches 0. 1— 0.2(— 0.45) mm diam.:
translucent to pale cervine brown or dark
brown, often with blackened stems and frag-
mentation regions, consistently shiny.
True lateral spinules absent. Soralia rare,
modified fissurai, in which the filaments are
strongly split, opening flat and recurving the
branch, exposing an irregular mass of white
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 93
farinose soredia (Fig. 40). Pseudocyphellae
abundant in subsp. trichodes, where they are
white, oval and raised; sparser in subsp. ameri-
cana where they are fusiform to elongate-fusi-
form, brownish, depressed and 0.5-1 mm long.
Apothecia occasional, sometimes abundant,
lateral; excipulum thallinum concolorous with
the thallus, thin, becoming excluded; disc
becoming convex, reddish brown, 0.5-l(-2)
mm diam. Spores 8 per ascus, ellipsoid, hyaline,
simple, 5.5-6(-7)x4-5 fcm. Pycnidia unknown.
Outer cortex K-, C— , KC — , PD-; inner
cortex and sometimes medulla K. — , C-, KC-,
PD+ red at least in parts (very rarely PD- in
subsp. americanci). Contains fumarprotocetraric
acid and usually chloroatranorin (the latter
absent in subsp. americanci).
Ecology. Mainly on coniferous trees, especially
spruce and fir, usually in open, wet habitats
such as bogs, swamps, lake and river edges;
sometimes in spruce-fir or beech-maple forests.
One specimen (Michigan. Isle Royale. Wetmore
2135A [MSC]) was collected on rock. The
sorediate morphotype is not associated with
bogs but rather appears to be found in open
or closed woodlands.
distribution. Subsp. trichodes occurs in the
Great Lake-St. Lawrence forest region in
the east, extending southward in the coniferous
forests of the Appalachians and northwards into
the southern boreal zone where it can be locally
abundant (Fig. 41 A). In the west it occurs
tn the coastal forest region from the Olympic
Peninsula to the Central Pacific Coast district of
Alaska.
B. trichodes subsp. trichodes is not uncom-
mon in Japan (Kurokawa in litt., Hawksworth
1972, Yoshimura 1974 sub Alectoria americana ).
discussion
Eryoria trichodes is readily separated from other
brownish pseudocyphellate members of the
Senus by its chemical reactions. Most speci-
mens of subsp. trichodes can be easily separated
from those of subsp. americana on the basis
°f the former’s uneven stems, raised pseudocy-
Phellae, generally paler thallus, and the regular
°ccurrence of chloroatranorin. Some intergrada-
h°n does occur (see below).
Fumarprotocetraric acid is almost invariably
present in subsp. trichodes and occurs in much
larger amounts than is generally true for subsp.
americana in which this compound can be
difficult to detect. The presence of chloro-
atranorin in this subspecies but not in subsp.
americana is difficult to explain, although it
must be stressed that it is not always detectable
in subsp. trichodes and it is also at low con-
centrations. West coast specimens of subsp.
trichodes are often extremely pale and may be
nearly translucent due to the almost complete
absence of any cortical pigments. This morpho-
type, which is represented by the type specimen
of "Alectoria delicata Mot.’’, may perhaps
represent an infraspecific taxon, but its status
is in need of further investigation.
Were it not for the large numbers of inter-
mediate specimens encountered in areas where
the two subspecies are sympatric (e.g., New-
foundland; Ahti & Hawksworth 1974), we would
have been inclined to recognize two species
in view of the several differentiating characters
and different distributions in North America.
Because of the intergradation, the rank of sub-
species, as interpreted by Hawksworth (1974),
appears to be particularly appropriate. Because
it is frequently fertile and is widely distributed,
subsp. trichodes is perhaps of greater antiquity
than subsp. americana.
Over the major part of its range, subsp.
trichodes is a rather uniform taxon. However,
one interesting ecological modification is charac-
terized by being quite small, often with some-
what contorted branches, and an abundance of
spinulose side branches. This morphotype is
generally found growing on coniferous trees in
Picea mariana bogs and peatlands. (We have
seen specimens from Connecticut, Michigan,
Quebec, and other areas.) A complete inter-
gradation of the spinulose morphotype and typi-
cal specimens of the subspecies has been
examined by both of us in the field, in La
Verendrye Park (Quebec). In this area, the
spinulose morphotype occurred in exposed
habitats on isolated trees and those at forest
margins, whilst normally developed plants were
encounted as one moved into more protected
(sheltered) parts of the adjoining forest. The
spinulose morphotype thus appears to be merely
an ecad.
The epithet trichodes has long been over-
looked by North American lichenologists. Fol-
94 Alectoria in North America
OPERA BOT. 42 (1977)
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 95
lowing the rediscovery of the holotype specimen
in PC by Dr. H. A. Imshaug in 1971, this was
sent on loan to us. After we examined the
specimen, the epithet trichodes was taken up as
the correct name for the species previously
called Alectoria pseudofuscescens Gyeln.
(Hawksworth 1972) and treated as Bryoria
pseudofuscescens here. The holotype was re-
examined since Michaux (1803) reported apo-
thecia in his species and since our studies showed
these to be absent in North American material
of B. pseudofuscescens . Also, it became clear
that Michaux's specimen was undoubtedly
gathered from eastern America (probably
Quebec; see below), quite outside the range
of B. pseudofuscescens. The holotype actually
consists of two Bryoria species on a spruce
twig; small, typical, fertile plants of B.furcellata,
together with abundant material of a very dark,
pendent, esorediate, sparsely pseudocyphellate
Bryoria with uneven branches which is also
fertile. TLC had earlier revealed a substance
which appeared to represent norstictic acid in
the blackish pendent specimen and it was for
this reason that the epithet was employed for
B. pseudofuscescens by Hawksworth (1972).
B. furcellata and B. pseudofuscescens are not,
however, sympatric in North America (com-
pare Figs. 48 and 60 A). In re-examining the
material we also discovered that neither B.
furcellata nor fragments of Hypogymnia
physodes (L.) Nyl. present in the packet ap-
peared to contain any PD + compounds although
these taxa normally react readily with this
reagent. It therefore seems likely that the whole
collection may have been treated in some way.
Perhaps by immersion in alcohol, which removed
the original substances. The PD + compound
originally determined by TLC as norstictic acid
might have been some unrelated compound.
Possibly resulting from a reaction of a thallus
substance with the preservative.
These data forced us to make some reluctant
decisions with respect to the epithet trichodes.
At first we considered the possibility of rejecting
the name as one based on discordant elements
(Art. 70). Since the original description, “S.
tenuissime capilliformis, prolixa. fuliginoso-
otra, laevigata; scutellis concoloribus, peltatis,
convexis” (Michaux 1803 pp. 331-332), could
be interpreted as referring to the blackish pen-
dent thallus alone, this procedure could not be
justified. In fact, the diagnosis supports the
treatment of this element as the holotype, and
this is the conclusion reluctantly reached
here. Furthermore, this interpretation of the
name is in agreement with the early usage of
it by at least one early author: Bachelot de la
Pylaie (see Alectoria trichodes under Excluded
taxa). Apart from the dark colour and the lack
of demonstrable fumarprotocetraric acid, the
holotype conforms to the present subspecies
which we termed canadensis, following Motyka,
earlier in our investigations. We are thus forced
to take up Michaux's name for this taxon and
reject our first interpretation of it.
Field work in the spruce and fir forests north
of Quebec City by one of us (l.M.B.) in the
autumn of 1974 and spring of 1975 uncovered
the fact that a fine, very dark, frequently fertile
morphotype of this taxon is extremely common
in that region and almost invariably grows
with B. furcellata. The holotype of Setaria
trichodes almost certainly came from this
area, since Michaux's travels took him from the
Lake Champlain area through the Richelieu
Valley and then northward through the southern
and middle boreal forests near Lac Saint-Jean
and Lac Mistassini (Marie-Victorin 1964).
Acharius (1810) was uncertain of the identity
of Michaux's Setaria trichodes but doesn't
appear to have examined it. Muller Argoviensis
(1878), however, did examine the holotype and
stated that it was identical to “A. jubata var.
lanestris Ach.”, also evidently regarding the
darkish pendent plant as the portion to be
treated as the holotype for Michaux's name.
Selected specimens
Exsiccatae. Howe: Lieh. Novae Angl. 70 (mixed):
Maine, Hancock County: Bar Harbor, Hopkins (FH,
NYBG) - Thomson: Lieh. Wise. Exs. 25: Wisconsin,
Ashland County: Glidden. J. W. Thomson (CANL,
COLO, DUKE, LAM, US. WIS) -Tuckerman: Lieh.
Amer. Septen. 2: New Hampshire. County unknown:
White Mountains, Tuckerman (BM).
Canada. British Columbia. Islands North Region:
Graham Island, Juskatla. Brodo 1 1675 (CANL) -
Islands South Region: Vancouver Island: Long Beach
Campground. Tibell 5132 (UPS) - Saskatchewan.
Waskesiu, Prince Albert Provincial Park, Jesberger
1240 (SASK) - Ontario. Algoma District: N Aubina-
Fig. 4L N. American distribution. - A: Bryoria trichodes subsp. trichodes. - B: B. trichodes subsp. americana.
96 Alectoria in North America
OPERA BOT. 42 (1977)
dong R., Cain 26009 (US, CANL, TRTC) - Cochrane
District: Moosonee area, Wetmore 5967A (MSC) -
Quebec. Territoire d Abitibi: Guyenne, Gaudreau 8213
(CANL) - Cté Frontenac: N-D des Bois, Masson
6046B (W1S) - Territoire du Nouveau Québec:
Schefferville. Brodo 9125D (p.p.) (CANL) - New
Brunswick. Albert County: Fundy National Park.
Ireland 11370 (CANL) - Prince Edward Island.
Queens District: West Barkley Beach, Charlotte-
town, Fabiszewski s.n., 21 July 1970 (CANL) -
Nova Scotia. Annapolis County: Kejimkujik National
Park, Ireland 12448 (CANL) - Victoria County:
Ingonish, Brodo 19069 (CANL) - Newfoundland.
Humber East District: NE of Deer Lake, Taylor
1736B (MSC) - Labrador North District: North
West River Twp., Coleman 28 (CANL) - Labrador
West District: Faden, Brodo 8837-5B (CANL).
U.S.A. Alaska. Eastern Pacific Coast Region:
Ketchikan, Krog 6225 (O) - Central Pacific Coast
Region: Kenai Peninsula, Marathon Mt., Krog 2180
(O) - Connecticut. Hartford County: Suffield, Evans
s.n., !9Aug. 1923 (FH)- Maine. Hancock County : Mt.
Desert L, Davis 32 (US) - Massachusetts. Plymouth
County: Plymouth, Faxon s.n., I Mar. 1884 (FH) -
Michigan. Chippewa County: Lower Tahquamenon
Falls, Imshaug 19944 (MSC) - Minnesota. Cook
County: Horseshoe L, Lake Saganaga. Hale 33526
(US) - New Hampshire. Cheshire County: Jaffrey,
Riddle 1714 (FH-Riddle) - New Jersey. Morris
County: Austin, Herb, of P.V. Le Roy s.n., Oct 1871
(NYBG) - New York. Hamilton County: Adirondack
Mts., Hermann 13851 (p.p.) (CANL) -North Carolina.
Macon County: Fodderstack Mt., Moore 1954
(DUKE) - Tennessee. Sevier County: Great Smoky
Mts. Nat. Park, Mt. Leconte, Dey 3357A (DUKE) -
Vermont. Addison County: Goshen (Corners), Dutton
1850 (FH) - Washington. Clallam County: Olympic
National Park, Brodo 14396 (CANL) - West Virginia.
Pocahontas County: Cranberry Glades, Gray 211 (FH)
- Wisconsin. Douglas County: Brule River Valley,
J.W. Thomson 2014 (WIS).
6b. Bryoria trichodes subsp. americana (Mot.)
Brodo & D. Hawksw., comb. nov.
Alectoria americana Mot., Fragm. Florist. Geobot.
6 : 449 (1960): type: America borealis. New Bruns-
wick, Grand Manon, H. Willey, 1879 (US holotype!);
basionym. - Bryopogon americunus (Mot.) Bystr.,
Ann. Univ. Mariae Curie-Sktodowska. C, 26: 274
(1971); nom. inval. (Art. 33).
Alectoria ambigua Mot., Bryologist 67: 17 (1964);
type: Newfoundland, Ferryland, 6 miles north-west
of Cape Broyle. east side of Mt. Carmel Pond, boulder
in upland heath, abundant, T. Ahti 582, 26 May 1956
(H holotype!). - Bryopogon ambiguus (Mot.) Bystr.,
Ann. Univ. Mariae Curie-Skiodowska, C, 26: 270
(1971); nom.inval. (Art. 33).
Alectoria sepiacea Mot., in Krog, Norsk Polarinst.
Skr. 144: 139 (1968); type: Alaska, Central Pacific
Coast Distr. , Kenai Peninsula. Marathon Mt., lower
part, H. Krog 2197, I June 1957 (O lectotypel); nom.
inval. (Arts. 32, 36, 37).
Plants differing from subsp. trichodes in the
generally darker brown colour of the thallus,
consistently isotomic dichotomous branching
pattern, with branches which are even in dia-
meter and often have blackened fragmentation
regions. The pseudocyphellae are often sparse,
fusiform, depressed, brownish and longer than
those of subsp. trichodes measuring 0.5-1 mm
in length. Contains fumarprotocetraric acid, but
in this subspecies, it is often in low concen-
trations, and a PD+ red reaction is often diffi-
cult to demonstrate; some plants may lack this
acid. Chloroatranorin appears to be absent
in subsp. americana.
Ecology. On trees, especially conifers; most
abundant in bogs or swamps or along rivers or
lake edges; sometimes on trees in meadows.
In the west it is found equally on Picea sitchen-
sis, Pinas contorta, Thuja plicata and Tsuga
heterophylla; in the east it is usually found on
Picea glauca. The “a mbigua ’’-morphotype
(see below) usually occurs on rocks and dry
soil, often in heathland areas, but is also rarely
present on trees.
Distribution. Restricted to lowland coastal
forests along both the east and west coasts
corresponding closely with areas having a high
annual rainfall (Fig. 41 B). The Mt. Washington
and Isle Royale outlying localities are well
known for their "oceanic” character with fre-
quent mists. The disjunct Californian population
is difficult to explain. This subspecies appears
to be endemic to North America to judge from
material we have examined, although it should
be pointed out that Motyka (I960) also reported
it from eastern Asia (Saghalien). Records of
“Alectoria americana" in Japan refer to subsp.
trichodes.
Discussion
This subspecies is rather variable with respect
to the thickness of the main stems, the fre-
quency of blackened regions on the thallus,
and the abundance of pseudocyphellae. This
subspecies, like subsp. trichodes, has a tend-
ency to produce short, pointed, lateral branches
recalling spinules but differing from them in
being broad at the base. Soralia are very rarely
found in subsp. americana and occur mainly
in the west. The extent of fumarprotocetraric
OPERA BOT. 42 (1977)
Brvoria Brodo & D. Hawksw. 97
acid production is also somewhat variable, and
in some cases (particularly terricolous popula-
tions in the east) this acid may be entirely
absent: PD + reactions are thus sometimes diffi-
cult to demonstrate.
In his accounts of this taxon Motyka (1960,
1964) indicated that the holotype was PD- and
made no reference to the occurrence of pseudo-
cyphellae. In fact the holotype reacts readily
with PD and has pseudocyphellae, although
these are of the slightly depressed, dark and
easily overlooked type characteristic of this
subspecies.
Specimens from the oceanic heaths of New-
foundland were treated as a distinct species,
A lectoria ambigua Mot., by Motyka (1964).
This morphotype commonly has apothecia. is
often PD-, and frequently has short, perpen-
dicular lateral, spinulose branches, but other-
wise appears identical to many of the corticolous
specimens examined. On the basis of extensive
field observations in Newfoundland, Dr. T. Ahti
(in litt.) considers this morphotype to belong to
subsp. americana and we concur with this view.
Selected specimens
Exsiccatae. Howe: Lieh. Novae Angl. 70 (mixed):
Maine, Hancock County: Bar Harbor, Hopkins (FH-
Riddle); Merrill: Lieh. Exs.. Ser. II, 129 (mixed):
Alaska, Fairbanks, Palmer (COLO).
Canada. British Columbia. Islands North District:
Graham Island, head of Awun Lake, Brodo 18485
(CANL) - Islands South District: Mt. Cain, N of
Schoen Lake, Ohlsson I I99B (MSC) - Coast District:
*9 miles E of Bella Coola, Ohlsson 2273 (CANL) -
Skeena River District: 21 miles E of Terrace, Ohlsson
-835 (CANL) - Quebec. Cté Gaspé-Est: Ile Bona-
''enture, Beaudoin 86 (CANL) - Cté Montmorency:
Laurentide Park, junction Hwy 54A and 54B. Scotter
6880 (H) - New Brunswick. Charlotte County: Grand
^lanon (sic) (Manan) Island. Willey s.n., 1879 (US) -
J^estigouche County: Dalhousie, Reijonen s.n., 16
Nov. 1930 (H) - Nova Scotia. Victoria County?:
8l- Paul Island, Erskine 53C2403 (NSPM) - New-
foundland. Bonavista S. District: Charlottetown,
Taylor 2110 (MSC) - Isle Miquelon: Delamare
s -n.,Sept. 1886 (COLO).
C.S.A. Alaska. Central Pacific Coast District: 1 mile
from Sterling Hwy. on Mt. Aleyska Rd., J. W.
Thomson 17817 (CANL, WIS) - East Pacific Coast
I 'strict: Juneau, Krog 5529 (O) - Bering Strait of
Lower Yukon River District: Darby Mountains. Pegau
T* (WIS, ALA) - California. San Benito County:
Look’s Bear Valley, Hannibal s.n., 16 Feb. 1907
* LAM ) - San Mateo County: Pilarcitos Creek Canyon,
Gerre & Doty 3348 (p.p.) (NYBG)-Mume. Aroostook
County: Portage, Riddle s.n., Aug. 1907 (LAM) -
~ Opera Botanica nr 42
Massachusetts . Bristol County: New Bedford, Willey
s.n., 1862-1898 (US) - Michigan. Keweenaw County:
Isle Royale National Park, Lowe 937J (FH) - Forbes
Lake, Wetmore 41 16A (MSC) -Oregon. Lane County:
Florence, Pike L-414 (PIKE) - Vermont. Rutland
County: Sherburne. Dutton 2410 (FH) - Washington.
Clallam County: Olympic Peninsula, Hurricane Ridge,
Brown and Muenscher 129 (FH).
7. Bryoria vrangiana (Gyeln.) Brodo & D.
Hawksw., comb. nov.
Alectoria vrangiana Gyeln., Magy. Bot. Lapok 31:
46 (1932); type: Sweden, Prov. Dalecarlia, par. Lok-
sund, Laknäs, E. P. Vrang 236, 18 July 1931 (BP
33.967 holotype!); basionym.
For further synonyms see Hawksworth (1972 pp
253-254).
Thallus pendent, to 10(— 12) cm long; branching
isotomic dichotomous towards the base but be-
coming anisotomic dichotomous towards the
apices so as to appear submonopodial, angles
between the branches mainly acute; branches
even to uneven in diameter, becoming spirally
twisted and contorted to foveolate, main
branches (0.3-)0.5-l .0 mm diam.; thallus often
bearing irregular contorted spinules not con-
stricted at the bases and which usually arise at
acute angles; thallus usually concolor-
ous, matt, dark olive brown to blackish.
True lateral spinules and pseudocyphellae
absent. Soralia absent to sparse or more rarely
abundant, mainly tuberculate on the main
branches, some fissurai soralia often also pre-
sent. white to brownish, to 1.0 mm long.
Apothecia and pycnidia unknown.
Cortex and medulla K— , C — , KC-, PD-;
soralia K — , C— , KC— , PD+ red. Contains
fumarprotocetraric acid.
Ecology. On conifers in the subalpine Abies
lasiocarpa, Larix lyalii-Pinus albicaulis forests
( 1650-2250 m elevation in the Canadian Rockies;
2950-3250 m in the southern Rockies). We saw
one collection on lignum from the arctic coastal
strip of Ontario.
Distribution. Because we are still not sure about
the taxonomic limits of this species, it is diffi-
cult to be certain of its distribution. It would
appear that B. vrangiana is a montane forest
species in North America (Fig. 42), as it is in
Europe (Hawksworth 1972).
This species appears to have been first report-
ed from North America by Räsänen (1933), but
98 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 42. Bryoria vrangiana. N. American distribution.
the specimen from Hazelton, British Columbia,
upon which this report was based (H !) was found
to be a mixture of B. fuscescens and B. pseudo-
fuscescens.
Discussion
Circumscription of B. vrangiana has proved
difficult and is discussed together with B. fusces-
cens. Rather few collections of the species have
been seen, and its limits are likely to remain
somewhat uncertain until more collections of it
are available and it has been studied carefully
in the field.
A specimen superficially extremely similar to
B. vrangiana but containing norstictic acid is
referred to B. pseudofuscescens (see the discus-
sion under the latter species).
Specimens
Canada. British Columbia. Coastal District: Garibaldi
Park, ridge to Little Diamond Head. Otto 8041 (p.p.)
(CANL, H) -Garibaldi Mountains, WhistlerMountain,
Brodo 13939 (CANL)- LakeTatleh (sic) (Tatla Lake),
Rothrock s.n., 21 Aug. 1863 ( 1865?) (US) - Columbia
River Basin: Revelstoke National Park, ridge between
Millar and Upper Jade Lake, Otto 3 1 3 1 B (UBC) -
22 miles N of Rossland, Brodo 15125B (CANL) -
Fraser River Basin: 28.9 miles S of Merritt, Brodo
8500 (CANL)- Kootenay District: Rogers Pass, Brodo
7661B (CANL) - Alberta. Banff, Sanson S-27 (FH) -
Waterton Lakes National Park, Bird & Lakusta 16836,
16841 (CANL) - Ontario. Kenora District: Cape
Henrietta Maria, Courtin 28 (CANL).
U.S.A. Arizona. Apache County: Trail toMt. Baldy,
Nash 7838 (p.p.) (CANL) -Graham County: Pinaleno
Mountains, top of Mt. Graham, Nash 4223, 4236
(NASH) - Montana. Glacier County: Glacier National
Park, Ptarmigan Creek, Imshaug 6036 (CANL)- South
Dakota. Custer County: Cicero Peak, Custer. Wet-
more 6993 (MSC) - Utah. San Juan County: Navaho
Mountain. Nash 5185 (CANL).
Bryoria sect. Divaricatae (DR.) Brodo & D.
Hawksw., comb. nov.
Alectoria subgen. Bryopogon sect. Divaricatae DR.,
Ark. Bot. 20A (II): 10 (1926); basionym. - Alectoria
sect. Bryopogon subsect. Divaricatae (DR.) Keissl.,
Rabenh. Krypt.-Fl. 9, 5(4): 138 (1958).
Alectoria subgen. Bryopogon sect. Divaricatae
subsect. Suhfihrillosae DR.. Ark. Bot. 20A (11): 10
(1926); type: A. bicolor (Ehrh.) Nyl. [=B. bicolor
(Ehrh.) Brodo & D. Hawksw.], lectotype; nom.
illegit. (Art. 63). - Bryopogon sect. Eujubalae subsect.
Suhfihrillosae (DR.) Gyeln., Feddes Repert. 38: 223
(1935). - Alectoria subgen. Bryopogon sect. Suhfihril-
losae (DR.) Mot., FI. Polska, Porosty 5(2): 58 (1962).
- Bryopogon sect. Subfibrillosae (DR.) Bystr., Ann.
Univ. Mariae Curie-Sktodowska, C, 26: 270 (1971).
Bryopogon sect. Fuscidulae subsect. Divaricatae
Bystr., Ann. Univ. Mariae Curie-Sktodowska, C, 26:
272 (1971), ‘‘(DR.)”; type: B. altaicus Gyeln.
[ = Bryoria nadvornikiana (Gyeln.) Brodo & D.
Hawksw.], holotype; nom. illegit. (Arts. 48, 64).
Bryopogon sect. Perspinosae Bystr., Ann. Univ.
Mariae Curie-Sktodowska, C, 26: 273 (1971); type:
B. perspinosus (Bystr.) Bystr. [=Bryoria perspinosa
(Bystr.) Brodo &D. Hawksw.], holotype; nom. inval.
(Art. 36).
Type species: Bryoria bicolor (Ehrh.) Brodo & D
Hawksw. (syn. Lichen bicolor Ehrh., Alectoria bicolor
(Ehrh.) Nyl.; lectotype).
Thallus erect, decumbent, caespitose or sub-
pendent to pendent; olivaceous-grey, brownish
or black, frequently with extensive blackened
areas, particularly towards the base, which are
devoid of algal cells. Lateral spinules generally
with somewhat constricted bases frequent to
numerous, arising either perpendicularly or at
slightly acute angles to the main stems, usually
rather short, straight or arcuate. Soralia present
in only a few species, fissurai, white or brownish,
bearing tufts of isidiiform spinules in some
species. Pseudocyphellae present or absent, in-
conspicuous, fusiform, depressed, often very
sparse. Cortex tending to become fasciculate.
Apothecia frequent in some species, very rare or
unknown in most; margins not ciliate, becoming
sorediate in one species; discs reddish brown.
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 99
Atranorin and chloroatranorin usually absent;
fumarprotocetraric acid present in most species;
other /3-orcinoI depsidones and pulvinic acid
derivatives absent.
Discussion
Bryoria sect. Divaricatae is the largest section
of the genus recognized here comprising about
20 species of which only 7 are known from
North America. This section shows its maximum
diversity in the mountains of south-east Asia.
The chemical components of the species in
this section are extremely uniform. Of those
species in which lichen products have been
demonstrated almost all have fumarprotocetraric
acid (sometimes in very low concentrations)
alone. B. nadvornikiana, although it contains
fumarprotocetraric acid, also has compounds
otherwise confined in Bryoria to sect. Implexae
(i.e., alectorialic and barbatolic acids) and is
placed in that section here. It is tempting to
speculate that this aberrant chemistry may have
been derived from an ancient hybridization
between members of these two different sections
(see also Discussion under Alectoria sarmentosa
subsp. sarmentosa).
Many species in this section tend to have erect
to caespitose habits and to occur predominately
on the ground or on moss-covered boulders
while others are strictly corticolous.
The species of the sect. Divaricatae are gen-
erally very sharply delimited from one another
and present few difficulties in determination.
This suggests that the group is particularly
ancient and not undergoing active spéciation at
the present time.
Bryoria bicolor (Ehrh.) Brodo & D. Hawksw.,
comb. nov.
Lichen bicolor Ehrh., Beitr. Naturk. 3: 82 (1788);
type: Hercynia. Ehrhart Crypt. Exs. no. 40 (LINN-
Sui. 1712.2(3) lectotypel); basionym. - Alectoria bi-
color (Ehrh.) Nyl., Actes Soc. Linn. Bordeaux 21:
291 (1856).
For further synonyms see Hawksworth (1972 p.
204-206).
Thallus erect or caespitose, rigid, usually
2-3.5(-4) cm tall; branching isotomic dichoto-
mous, angles between the dichotomies variable,
secondary and tertiary branches usually at right
angles present, often obtuse at the base;
branches terete, even in diameter, sometimes
becoming slightly compressed towards the base,
usually 0.2-0.3(-0.5) mm diam.; basal parts
black, apices olivaceous grey to cervine brown,
shiny.
Numerous perpendicular lateral spinules with
somewhat constricted bases present. Soralia
absent. Pseudocyphellae usually sparse, incon-
spicuous, fusiform, dark to pale brown, plane or
slightly raised, to 0.2-3. 5 mm long.
Apothecia and pycnidia not seen.
Inner cortex and medulla K-, C-, KC-,
PD+ red at least in parts (never PD-). Con-
tains fumarprotocetraric acid.
Ecology. Mainly on rock and heath, but often on
trees, in subalpine or coastal habitats. Degelius
(1941), describing the ecology of the southern
Appalachian population, said the species is
found above 1540 m, mainly in the Abies fraseri
forest. Some of Degelius’ plants, however, have
been identified here as B. tenuis, and so his
comments may apply in part to both species.
Distribution. In oceanic and suboceanic areas
along the west coast from Vancouver Island to
south-east Alaska, in the Great Smoky Moun-
tains, the White Mountains, and a few other
mountain or heath areas (Fig. 43). Reports of
this species in Mexico (see Imshaug 1956 p. 366)
appear to be erroneous and probably refer to
Oropogon spp.
The tendency of this species to occupy
“oceanic” habitats in the northern Alps
(Schauer 1965), Scandinavia (Degelius 1935)
and in the British Isles (Hawksworth 1972) has
been well documented. Hawksworth (1972)
mapped the world distribution of B. bicolor
noting its occurrence in montane subalpine and
subarctic areas of Europe, Himalaya. Japan, and
Malaysia. Krog & Swinscow ( 1975) consider the
montane East African species Bryoria ruwen-
zoriensis to be conspecific with B. bicolor,
but in view of its very robust habit and charac-
teristic habitat we accept it as a species as did
Jprgensen (1972).
Discussion
Bryoria bicolor is a clearly circumscribed
species in North America and is only likely to
be confused with B. nitidula (from which it is
easily distinguished by the black main stems and
100 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 43. Bryoria bicolor. N. American distribution.
pale perpendicular lateral spinules), and B.
tenuis (from which it can be distinguished in
that B. hicolor forms dense tufts with abundant
third-order branches arising at right angles).
The separation of B. bicolor from B. tenuis
has been discussed in some detail by Jprgensen
& Ryvarden (1970).
Motyka (1964) reported that the “f. melaneira
(Ach.) Nyl.” (see Hawksworth 1972 pp. 205-206
for the nomenclature of this taxon) occurred in
North America. This morphotype, characterized
by concolourous apices (and lateral spinules as
well, on occasion), does not appear to be well-
marked in North America.
Apothecia have been described for this
species by a number of workers but it seems
that these reports have been based on examina-
tions of incorrectly determined (mainly Asian)
material: the reports of them in Wade (1959),
for example, are known to have been based on
nineteenth century sources (A. E. Wade in litt.).
As far as we are aware, Motyka's (1964) state-
ment that apothecia are unknown in this species
is correct.
Selected specimens
Canada. British Columbia. Islands North District:
Graham Island: Juskatla, Brodo 1 1676 (CANL) -Coast
District: Banks Isles, Menzies s.n., 1785 (BM)- Island
South District: Vancouver I., summit of Mt. Benson,
Macoun 202 (CANL) - Quebec. Cté Charlevoix
Ouest: Laurentide Park, Scotter 6693 (H) - Gaspé-
Nord: Mt. Albert. Gauthier 2638 (CANL) -Territoire
de Mistassini: Otish Mts., Shchepanek 71-L-I3
(CANL) - Newfoundland. Labrador West District:
Churchill Falls, Brassard 5326 (CANL) - Ferryland
District: Biscay Bay, Tuomikoski 393 (H) - Gander
District: Mt. Peyton, Ahti 8961 (H).
V .S. A .Alaska. Central Pacific Coast District: Prince
William Sound. Pigot Bay, Krog 2203 (O) - Eastern
Pacific Coast District: Sitka, Krog 6241 (O) - New
Hampshire. County unknown: White Mts., Tucker-
man s.n., 1840 (US) -North Carolina. Swain County:
Great Smoky Mts., Clingman’s Dome, Degelius s.n.,
12 Sept. 1939 (DEGELIUS) -County unknown: Great
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 101
Fig. 44. Bryoria carlottae . Holotype. Scale
0.5-1 .0 mm.
Smoky Mts., Forney Ridge, Degelius s.n., 12 Sept.
1939 (DEGELIUS) - Yancey County: Mt. Mitchell,
Black Mts.. Sierk 784 (DUKE) - Oregon. Clatsop
County: Saddle Mountain State Park. Pike 3812A
(OSC) - Tennessee. Carter County: Roan Mountain,
Dey 2125 (DUKE) - Sevier County: Mt. LeConte,
Great Smoky Mts., Degelius s.n., 13-14 Sept. 1939
(DEGELIUS) - Vermont. Lamoille County: Mt.
Mansfield, Cain 26002 (TRTC).
9. Bryoria carlottae Brodo & D. Flawksw., sp.
nov.
Thallus pendulus vel subpendulus, 7-8 cm longus;
rami praecipue aniso-dichotomiter vel fere submono-
podialiter sed basi iso-dichotomiter fastigiati et in
angulos obtusos divergentes; rami principales basi
0.2-0.25 mm diam., ramulis numerosis brevibus
praecipue 2-4 mm longis perpendiculariter instructi;
Iota planta olivacea sed interdum basi brunneo-
olivacea.
Rami laterales spiniformes veri, isidia et soralia
desunt. Pseudocyphellae copiosae, albae. fusiformes,
aliquando elongatae, planiusculae vel interdum leviter
elevatae.
Apothecia et pycnidia ignota.
Cortex K— , C — , KC — , PD— ; medulla K— , C— ,
KC — , PD+ rubescens. Thallus acidum fumarprotoce-
traricum continens.
Holotypus: America septentrionalis. Canada. British
Columbia, Insulae Queen Charlotte, lnsula Graham.
2mls austro-orientalem versus a Port Clements, austra-
lem versus a Tlell-Port Clements Road, 53°40N.
132°09'W. in aperto turbario Pinus contortae et
Proxima sylva Tsugae-Thujae, corticola, 1. M. Brodo
>8096, 11 July 1971 (CANL 38266). Isotypus: BM.
Fig. 44.
Thallus pendent to subpendent, 7-8 cm long;
branching mostly anisotomic dichotomous to
almost submonopodial except at the base where
the dichotomies are isotomic and the angles
between the branches obtuse; main branches
0.2-0.25 mm diam. at the base, with numerous
short perpendicular side branches 2-4 mm long;
uniformly olivaceous but sometimes brownish-
olivaceous at the base.
True lateral spinules, isidia and soralia absent.
Pseudocyphellae abundant, white, fusiform,
sometimes elongated, plane or sometimes
slightly raised.
Apothecia and pycnidia unknown.
Outer cortex K-, C-, KC-, PD-; inner
cortex and medulla K-, C — , KC-, PD+ red.
Contains fumarprotocetraric acid.
Ecology. On trees at the edge of bogs, or on
exposed bluffs.
Distribution. Endemic to Queen Charlotte
Islands (Fig. 45). Plants which are restricted to
the Queen Charlotte Islands or which have
their centres of distribution there are not un-
common (see general section Distribution, cate-
gory 9).
Discussion
The broadly divergent habit and frequently per-
pendicular side branches together with the
distinctive olive colour of B. carlottae readily
distinguish it from B. trichodes subsp. ameri-
cana, the only other uniformly coloured non-
sorediate species containing fumarprotocetraric
102 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 45. Bryoria cariottae. Known distribution.
acid in the medulla and found in the same region.
Its relationships to other species are unclear
from the relatively few specimens examined so
far but its affinities appear to lie with the sec-
tion Divaricatae which has its centre in Asia.
Specimens
Canada. British Columbia. Islands North District:
Graham Island: Tow Hill, Brodo 9900 (CANL) -
Port Clements, Brodo 18093, 18096 (CANL)- Langara
Island: McPherson Point, Brodo I0657B (CANL) -
Moresby Island: Between Sandspit and Copper Bay,
Brodo 12879 (CANL) - Skidegate Lake. Brodo 18499,
18503B (CANL).
10. Bryoria cervinula Mot. ex Brodo & D.
Hawksw., sp. nov.
Alectoria cervinula Mot., Bryologist 67: 19 (1964);
type: Alaska, East Pacific Coast District, Sitka,
Harbour Mt., lower part. H. Krog 6222, 31 August
1957 (O lectotypel): nom. inval. (Arts. 36, 37). -
Bryopogon cervinutus (Mot.) Bystr., Ann. Univ.
Mariae Curie-Skfodowska, C, 26: 271 (1971); nom.
inval. (Arts. 33, 36. 37).
Thallus decumbens vel suberectus, 6-10 cm longus;
rami praecipue monopodialiter fastigiati, ramulis
numerosis brevibus et crassis perpendiculariter vel
arcuate instructi, quare thallus spinosus videtur,
ramis diametro aequalibus, basi 0.25-0.5 mm diam.;
planta pallide brunnea vel atrobrunnea, non concolor,
aspectu variegato, opaca vel bombycina. plerumque
non nitida.
Rami laterales spiniformes veri, isidia et soralia
desunt. Pseudocyphellae praesentes sed plerumque
rarissimae, albae vel brunneae, fusiformes, depressae.
Apothecia et pycnidia ignota.
Cortex K-, C — , KC-, PD — ; medulla K- vel +
brunnea, C - , KC - , PD + rubescens. Thallus acidum
fumarprotocetraricum continens.
Holotypus: America septentrionalis, U.S.A.,
Alaska, East Pacific Coast District, Sitka, ad pedes
montis Harbour Mountain. H. Krog 6222, 31 August
1957 (O).
Thallus decumbent to somewhat erect, 6-10 cm
long; branching mostly monopodial, with
numerous short stout perpendicular to arcuate
side branches giving the thallus a very spiny
appearance; branches even in diameter, 0.25-0.5
mm diam. at the base; pale brown to dark brown,
not concolourous, appearing variegated, matt to
satiny, not generally shiny. Fig. 46.
True lateral spinules, isidia and soralia absent.
Pseudocyphellae present but often very sparse,
white or brown, fusiform, depressed.
Apothecia and pycnidia unknown.
Outer cortex K-, C — , KC-, PD — ; inner
cortex and medulla K— or + brown, C— ,
KC — , PD+ red. Contains fumarprotocetraric
acid.
Ecology. On soil and vegetation between rocks,
on subalpine slopes, and on coniferous tree
limbs in subalpine or subarctic localities.
Distribution. Southeastern Alaska and the
Queen Charlotte Islands, with a single locality
in the Bering Sea region (Fig. 47). This North
American endemic should be sought on the
Aleutian Islands.
Discussion
Bryoria cervinula is a very distinctive species
characterized by extremely stout, often some-
what arcuate, lateral, secondary branches. It
appears to show affinities with a number of
species in the “A. asiatica group” known only
from Asia (Bystrek 1969, Jprgensen 1972, Nuno
1971) but to differ from all those described in
many features. (The types of all pertinent taxa
have been examined by us.)
Specimens
Canada. British Columbia. Islands North District:
Moresby Island: Laing Point, Brodo 10786 (CANL) -
SW of Skaat Harbour. Brodo 17769 (CANL).
U.S.A. Alaska. Central Pacific Coast District: Cor-
OPERA BOT. 42 (1977)
Brxoria Brodo & D. Hawksw. 103
Fig. 46. Bryoria cervinula. B. C., Queen Charlotte Islands. Brodo 17769 (CANL). Scale 0.5-1. 0 mm.
dova, Auliot 1550 (DUKE) - Eastern Pacific Coast
District: Baranoffl., Trelease s.n.. 15 June 1899 (US)
- Juneau, Krog 551 1 (O) - Sitka, Harbour Mt.. Krog
6222. 6223 (O) - Sitka. Indian River. Cooley s.n.,
12 Aug. 1891 (NYBG) - Kuiu !.. Washington Bay,
Eyerdam 1117 (COLO) - Lower Yukon River District:
6Ö miles NW ofGolovin, Pegau 43 (W1S).
1 1 . Bryoria furcellata (Fr.) Brodo & D.
Hawksw., comb. nov.
Cetraria furcellata Fr., Syst. Orb. Veg. 1: 283 (1825),
type: America borealis (UPS-Fr. holotype!); basio-
nym. - Evernia furcellata (Fr.) Fr., Lieh. Eur.: 468
(1831).
Non Alectoria furcellata R. Sant.. Svensk Bot.
Tidskr. 62:489(1968).
Alectoria nidulifera Norrl., in Nylander, Flora
58: 8 (1875); type: Fennia, Tavastia australis, kor-
pilahti, super corticem Pini, E. Lang, 1873 (H-Nyl.
35979 holotype!). - Bryopogon niduliferus (Norrl.)
Elenk., Lieh. Fl. Ross. I: 84 (1906). - Alectoria
chalybeiformis f. nidulifera (Norrl.) Merr., Bryo-
logist 14: 37 (191 1).
? Alectoria simplicior v. alba Gyeln., Nyt Mag.
Naturvid. 70: 62 (1932); type: Fennia, Karelia ladogen-
sis, Sortavala. Kotiluoto. ad scopulos, 1. inkola (holo-
type; not traced). - Bryopogon niduliferus f. albus
(Gyeln.) Gyeln.. ActaGeobot. Hung. 2: 166 (1937).
Bryopogon niduliferus f. compactus Gyeln., Acta
Geobot. Hung. 2: 166 (1937): type: U.S.A.. New York.
Essex Co.. Minerva, on twigs. H. D. House. II
August 1927 (US lectotype! "portion of specimen
sent toGyelnik").
Bryopogon niduliferus f. isidialius Gyeln., Acta
Geobot. Hung. 2: 166 (1937); type: U.S.A., New
Jersey, Point Pleasant. C.C. Plitt (holotype not traced;
?isotypes! "Plitt 405. 29 Aug. 1909" FH, US).
Alectoria carta Ostm., in Hasselr., Ark. Bot. 30A
(13): 3 (1943); nom. inval. (Arts. 32, 34. 36, 37).
Thallus caespitose. usually 3— 5(— 12) cm long;
branching regularly isotomic dichotomous,
angles between the basal dichotomies usually
broad, those between the apical dichotomies
usually acute; branches even in diameter, some-
times becoming slightly compressed towards
the base, 0.3— 0.4(— 0.5) mm diam.; very pale
brown to cervine brown or rarely dark brown
104 Alectoria in North America
OPERA BOT. 42 (1977)
to black, often darker towards the base; base
persistent.
Short perpendicular lateral spinulose branches
sometimes present, occasionally abundant.
Pseudocyphellae absent. Soralia usually abun-
dant, fissurai, sometimes slightly raised, nar-
rower than the branches on which they occur,
white. 0. 3-1.0 mm long, becoming covered with
tufts of isidiiform spinules 0.5-2. 5 mm long.
Apothecia very rare, lateral; excipulum thalli-
num concolourous with the thallus, thick,
usually becoming sorediate; disc concave at first
but becoming convex and excluding the margin
with age, light brown to reddish brown,
0-8— 2.0(— 4.0) mm diam. Spores 8 per ascus, sub-
globose, hyaline, simple, 6-7 x 3-4 /am.
Outermost portion of cortex K-, C-, KC-,
PD— ; inner cortex and medulla K-, C-, KC-,
PD+ red or very rarely PD— ; soralia K-, C-,
KC-, PD+ red. Contains fumarprotocetraric
acid.
Ecology. Mainly on coniferous trees, especially
Pinus spp. and Picea mariana, but also found
on rock faces, boulders, coastal heaths (espe-
cially in Newfoundland; Ahti & Hawksworth
1974), lignum (especially fence rails), and,
rarely, soil. In the north, it is a characteristic
species of the more open conifer forests such as
those dominated by jack pine and black spruce.
In northeastern United States, it is frequently
associated with bogs and pine stands (see Brodo
1968). In the southeastern United States, it
seems to be largely restricted to pine forests at
high elevations.
Distribution. Bryoria furcellata is the most wide-
spread species treated in this paper. It is basi-
cally found in the Appalachian-Great Lakes
region, ranging northward into the middle boreal
and even the northern boreal zones (Fig. 48),
but its "front of abundance" ends in the southern
half of the latter zone (Ahti 1964 p. 193). Its
restriction to the mountains and its absence in
the Piedmont in southeastern United States has
been well documented by W. Culberson (1958).
The occurrence of disjunct populations in the
southern Rocky Mountains and in the Mexican
highlands is not unusual for this kind of distri-
bution, and many examples can be cited (see
Brodo 1968 pp. 82, 73). The disjunct west coast
localities are more noteworthy, probably demon-
strating the great age of the North American
population. In Europe, where it appears to have
a continental distribution (Ahlner 1948), the
species is found mainly in the boreal forest
region (Poelt 1969), especially associated with
pine forests (Ahlner 1948), but it also occurs in
Madeira (Jorgensen 1972) and has recently been
discovered in Scotland (Hawksworth & Coppins
1976). B. furcellata (sub Alectoria nidulifera)
has been reported from Himalaya by Bystrek
(1969), and from Japan by Kurokawa (1959) who
indicated that it was not rare in inland forests
in central and northern Japan; this can therefore
be considered a circumboreal species. B. furcel-
lata also occurs as a disjunct in Central America
in the Costa Rican highlands (Cartago Prov., S
slope of Volcan Irazu, c. 3000 m, R. W. Holm
& H. H. Iltis 1415 [herb. Dodge]), but is so
far unknown from South America.
Discussion
The identification of this very clearly delimited
species rarely presents much difficulty. Very
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 105
Fig. 48. Bryoria furcellata. N. American distribution.
ra re specimens lacking sprouts of spinules in
the soralia might possibly be confused with
simplicior but these two species can be
e asily distinguished in that B. simplicior has
§ r eenish-black soralia broader than the branches
° n which they occur (a most useful character
apparently first noted by Degelius (1934) and is
always PD — .
Motyka (1964 p. 15) considered that the North
American material of this species tended to
differ from that in Europe in having soralia
located in minute tubercles rather than in deeply
106 Alectoria in North America
OPERA BOT. 42 (1977)
incised fissures. However, we have found that
many North American specimens have deeply
incised soralia, and European material can have
somewhat raised soralia. Furthermore, these
types intergrade, frequently on the same thallus.
An extremely pale, grey-brown specimen with
many spinulose branches and adventitious
spinules was seen from Oregon (Pike 3818
[OSU]). Although resembling B. furcellata
in branching type, it lacks true soralia and is
probably conspecific with the material ten-
tatively called B. cfr. subcana (see Discussion
under that species) from California. A decision
on its true affinities will have to await further
studies.
The name Alectoria chalybeiformis has been
incorrectly applied to this species by a number
of early North American authors (e.g., Howe
1911).
Hawksworth (1969 p. 397) pointed out that the
earliest name for this taxon at the rank of species
was Cetraria furcellata Fr., something Ahlner
had been aware of in 1941. In view of the
existence of the name Alectoria furcellata R.
Sant., which was treated as a synonym of A.
nigricans by Hawksworth (1970. 1972), the
epithet was preoccupied in Alectoria and so
could not be taken up for this species then.
When A. nidulifera is considered to belong to a
genus other than Alectoria, however. Fries'
name cannot be rejected and must be taken up
in accordance with Art. 11. It is with some
reluctance that we introduce this change here
since the epithet nidulifera is so well-known on
both sides of the Atlantic. Two methods of
saving this epithet were considered, (a) to trans-
fer Santesson's epithet into Bryoria and (b) to
describe one of our new species of Bryoria
under this epithet. The first alternative is un-
tenable as Santesson's “species" does not be-
long to Bryoria but to Alectoria s. str. as under-
stood here. To implement the second would
establish three “ furcellata ” epithets in this
group of genera all based on different types -
something contrary to Rec. 23B(h).
Selected specimens
Exsiccatae. Brodo: Lieh. Can. Exs. 82: Ontario,
Cochrane District: Moosonee, Brodo 14779 (CANE) -
Cummings, Seymour & Williams: Dec. N. Amer.
Lieh. 53: Virginia. Norton, Seymour (BM. CANL,
DUKE, MSC, NYBG. US, WIS) - Cummings, Wil-
liams & Seymour: Lieh. Bor. Amer. 16: Maine,
Washington County: Lubec, Cummings & Teller
(NYBG, US) - Hale: Lieh. Amer. Exs. 74: West
Virginia, Randolph County: Spruce Knob Lake. Hale
(BM. CANL, COLO, DUKE, LAM, MSC) - Merrill:
Lieh. Exs., Ser. I, 211: Maine, Knox County: War-
ren, G. K. Merrill (BM, CANL, FH. NYBG. US) -
Rel. Tuck. 3: Massachusetts , Hampshire County:
Amherst, Tuckerman (BM, CANL, COLO, DUKE.
FH, LAM, MSC, NYBG, US, WIS).
Canada. Alberta. Waterton Lakes National Park,
Redrock Canyon, Imshaug 6112 (MSC) - Saskatche-
wan. Lake Athabasca, Uranium City, Scotter 3785
(WIS)- La Ronge, Jesberger 1281 (SASK )-Manitoba.
Lac Brochet, Scotter 2800 (WIS) - Ontario. Algoma
District: Mississagi R., Twp. 196. Cain 2601 1 (CANL.
UBC. TRTC) - Haliburton County: W of Algonquin
Park Gate. Cain 27295 (US) - Quebec. Cté Dor-
chester: Morisset, Masson 5632 (WIS) - Cté Gaspé-
Est: Cap-des-Rosiers-Est. Brodo 18674 (CANL)-Ter-
ritoire de Mistassini: Otish Mts.. Mont du Lagopède.
Shchepanek 7I-L-59 (p.p. ) (CANL) - Territoire de la
Baie de James: Rupert House, Kucyniak & Tuomi-
koski 51A (p.p.) (CANL) - Territoire du Nouveau
Québec: George River. Hustich s.n. (H) -New Bruns-
wick. Albert County: Fundy National Park, Ireland
11332 (p.p.) (CANL) - Nova Scotia. Halifax County:
Lake Chalotte. Taschereau B-8 (NSPM) - Prince
Edward Island: Kings County: Montague, Ireland
10208A (p.p.) (CANL) - Newfoundland. Labrador
West District: Menihek Lake. Brodo 8853-5B (CANL)
(See Ahti & Hawksworth 1974.) - Yukon. Hunker
Creek. Mascoun s.n., 26 July 1902 (FH) - Northwest
Territories. Mackenzie District: Middle Ross Lake,
Ahti 10271 (H) - Keewatin District: Ennadai Lake.
Brown 1330 (CANL).
U.S.A. Alaska. Alaska Range District: McKinley
Park. Hanson 461 (COLO)- Arizona. CochiseCounty:
Chiricahua Mts.. Weber & Shushan s.n., 18 April 1957
(COLO) - Greenlee County: Hannaga Meadows
campground, Nash 7710 (CANL, NASH) - Pima
County: Mt. Lemmon. Darrow 774 (COLO) - Cali-
fornia. Humbolt County: Areata, North Spit. Becking
6104017 (US) - Marin County: Inverness, Tavares 561
(COLO) - Santa Clara County: Santa Cruz Mountains,
Herre 798 (F ) -Colorado. Costilla County: Purgatory.
Faxon s.n., 6 April 1882 (FH) - Connecticut. New
London County: Old Lyme, Evans 754 (FH) - Illinois.
County unknown: Calkins (107) 491 (NYBG) -
Indiana. St. Joseph County: South Bend. Bosch s.n..
no date (FH-Tuck.) - Iowa. Fayette County: Locality
unknown, Fink s.n., 1895 (WIS. MIN. US. NYBG).
Fink s.n.. 1899 (H) - Maine. Oxford County: Buck-
field, Parlin 7438 (FH ) - Maryland. Alleghany County:
Oldtown, Hale 14483 (US) - Massachusetts. Barn-
stable County: Cape Cod, Nicholson State Park.
Brodo 4209 (CANL, MSC) - Michigan. Alger County:
NE of Kingston Lake, Harris 6039 (MSC) - Minne-
sota. St. Louis County: Near Bear Lake State Park.
Hale 23220 (US) - Missouri. St. Louis, Engelmann
s.n.. 18 Aug. 1878 (US) - New Hampshire. Grafton
County: Hanover. Croasdale 14 (US) - New Jersey-
Sussex County: Highview Mt. State Park. Hale I73Ö2
(US) - New Mexico. San Miguel County: Las Vegas,
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 107
Arsène 20086 (FH) - New York. Franklin County:
Adirondack Mts., Hermann 15643 (US) - North
Carolina. Macon County: Bearpen Mt.. Moore 1418
(DUKE) - Ohio. Ashtabula County: Orwell, Bogue
864 (NYBG) - Oregon. Lane County: Florence, Pike
L-564 (CANL) - Pennsylvania. Luzerne County: Red
Rock, Hale 16168 (US) - Rhode Island. County un-
known: Gilliman s.n.. no date (NYBG) - Tennessee.
Knox County: House Mt.. Cain 132 (FH) - Vermont.
Rockbridge County: Goshen. Dutton 2292 (FH) -
Virginia. Floyd County: Near Rocky Knob, Blue
Ridge Parkway, Hale 12833 (US) - Washington. Sno-
homish County: Near Big Four Inn. Schallert 3210
(US) - West Virginia. Nicholas County: Birch River.
Haie 11562 (US) - Wisconsin. Ashland County:
Apostle Islands, Outer Island, Tans 4 (WIS).
Mexico. Oaxaca. Sierra de San Felipe, Pringle
195 (p.p.) (DUKE) - 53 km NW of Oaxaca, Hale
20805 (p.p.) (US).
12. Bryoria nitidula (Th. Fr.) Brodo &
D. Hawksw., comb. nov.
Bryopogon jubatum var. nitidulum Th. Fr., Nova
Acta Reg. Soc. Sei. Upsal., ser. 3, 3: 25 (I860)
[Reprint]; type: Norway, Finnmark, Varanger,
Klubben. T. M. Fries, 15 August 1857 (UPS lecto-
typel); basionym. - Alectoria nitidula (Th. Fr.)
Vain., Medd. Soc. Fauna Fl. Fenn. 6: 1 16 (1881).
Alectoria irvingii Llano, J. Wash. Acad. Sei.
41 : 198 (1951); type: Alaska, Anatuvuk Pass, alt.
100 m, 151°32'W. 69°20’N. on ground talus slopes,
west wall, G. A. Llano 307b, 15 July 1949 (US
holotypel). — Bryopogon irvingii (Llano) Bystr.,
Ann. Univ. Mariae Curie-Sklodowska, C, 26: 275
(1971); nom. inval. (Art. 33).
For further synonyms see Hawksworth (1972 p.
228).
Thallus erect to decumbent, 5-8 cm tall;
branching isotomic dichotomous at the base,
becoming anisotomic dichotomous towards the
apices, angles between the dichotomies mainly
acute; branches even in diameter, straight,
0-5— 0.6(— 0.8) mm diam.; cervine brown to dark
brown or black, usually concolourous; dying
Irom the base.
Lateral spinules usually frequent, slightly
constricted basally, arising over the whole
lhallus. Pseudocyphellae usually present,
sparse, inconspicuous to conspicuous, elongate
fusiform, plane to slightly raised, c. 0.5 mm
long, brownish to dark brown or black. Soralia
absent.
Apothecia and pycnidia unknown.
Inner cortex and medulla K-, C— , KC— ,
PD+ red at least in parts. Contains fumar-
Protocetraric acid.
Ecology. On dry or wet tundra, or occasionally,
in pine stands in the northern hemiarctic
region; generally associated with rocks,
especially non-calcareous rocks, although we
have seen some specimens from calcareous
tundra. There is one specimen from the low
branches of a white spruce (Ontario, Fort
Severn, Ahti s.n., 2 Aug. 1958 [O]).
Recent research by K. A. Kershaw and
D. W. Larson (Kershaw & Larson 1974.
Larson & Kershaw 1975, Kershaw 1975) has
helped to explain the physiological bases for
the ecological and geographic distribution of
B. nitidula. They have shown that this species
is an extreme xerophile at cold temperatures,
explaining its tendency to be found on ridge
tops where air flow and drainage are at a
maximum and where both moisture levels and
temperatures are low (Kershaw & Larson 1974).
Further work provided evidence for the adaptive
value of dark colour for lichens such as B.
nitidula which are found in habitats largely
swept clean of snow during the winter (Larson &
Kershaw 1975. Kershaw 1975). Whereas B.
nitidula would seem to have a net carbon
deficit during the summer months due to high
respiration rates at high temperatures, in early
spring its dark cortical pigmentation enables
the thallus temperature to rise significantly
above the air temperature and greatly increase
its available growth period. In small "melt-
pockets" caused by tufts of these dark plants,
assimilation may even occur underneath thin
layers of snow (Larson & Kershaw 1975).
Distribution. Arctic to hemiarctic; entirely
absent from the western mountains with the
exception of one collection from the Garibaldi
Mountains north of Vancouver (Fig. 49). There
are other populations in the Otish and
Shickshock Mountains of Quebec. These peaks
are all well known for their arctic disjuncts.
The absence of B. nitidula in the forested
interior of Newfoundland is striking but not
unexpected. Its occurrence over granite on the
wind-swept coastal heaths near Halifax is also
not entirely surprising. These coastal localities
have numerous northern relicts (Roland &
Smith 1969 p. 296). B. nitidula is circumpolar,
but is of very scattered occurrence in northern
Europe and Asia (Hawksworth 1972).
It has recently been recognized in Japan as
108 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 49. Bryoria nitidula. N. American distribution.
part of the orohemiarctic zone flora (Jprgensen
1975).
Discussion
Bryoria nitidula has often been confused with
B. bicolor, a species from which it is readily
separated by its coarser habit, branching
pattern, and concolourous darker thallus. In
addition B. nitidula is essentially a species of
the tundra in North America whilst B. bicolor
is a coastal species restricted to more temperate
parts of the continent. A more frequent cause of
misidentification aries from the superficial
similarities of B. nitidula and Cornicularia
divergens, species which are often associated
with one another. C. divergens is, however,
easily distinguished by its white and more
discrete oval pseudocyphellae which react C +
rose-red (olivetoric acid), and its shiny, dis-
tinctly red-brown colour.
Ahti (1964) noted that Ontario material of
B. nitidula tended to be coarser than the
Scandinavian specimens available to him. This
species does indeed tend to be more luxuriantly
developed in North America than either Europe
or Japan, but as specimens similar to those in
Europe occur in North America and intergrade
with the coarser plants, the degree of luxuriance
appears to be of little taxonomic importance.
This species varies considerably in colour
from dark, dull red-brown to almost black,
and the habit and branching vary from erect
and almost strictly monopodia! to decumbent
and very irregularly branched (Fig. 50). While
extreme morphotypes are often very distinctive,
a complete series of intergradations occurs,
indicating that these types of variations are of
little taxonomic significance.
Motyka (1964) considered Alectoria irvingii
Llano as a good species closely allied to “A",
subdivergens Dahl, and possibly belonging in
Oropogon Th. Fr. We agree with Krog (1968)
that this taxon comes within the range of
variation of B. nitidula although the thalli of
the type specimen are rather coarse.
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 109
Fig. 50. Bryoria nitidula. - A: The common erect, robust morphotype. Ontario, Cape Henrietta Maria, Neal 019
(CANL). Scale 0. 5-1.0 mm. - B: The small, prostrate morphotype. Quebec, Gaspé, Mt. Albert, Brodo 18538A
(CANL). Scale 0.5-1. 0 mm.
Selected specimens
Exsiccatae. Thomson: Lieh. Arct. 84: N.W.T..
District of Keewatin: Rossby Lake, J. W. Thomson &
Larsen (CANL, COLO, DUKE, US).
Canada. British Columbia. Coast Area: Garibaldi
Mountains (other data unknown) - Manitoba. Fort
Churchill, Gillett 1608 (CANL) - Ontario. Kenora
District: Cape Henrietta Maria, Cowell 1286A (CANL)
- Quebec. Cté Gaspé-Nord: Mont Albert-Nord.
Gauthier 2061 (CANL) - Territoire du Nouveau
Québec: Great Whale River. L. Thomson 21 (US) -
Nova Scotia. Halifax County: Duncan's Cove.
Taschereau B-4 (NSPM ) - Newfoundland. Labrador
North District: Hopedale, Perrott 1-L (FH) (See also
Ahti & Hawksworth 1974.) - Yukon. British Moun-
tains, Trout Lake, J. W. Thomson & Larsen 14756
tP-P ) (WIS) - Northwest Territories. Franklin District:
Baffin Island, Frobisher Bay, Hale 344 (CANL,
WIS). - Keewatin District: Parsons Lake, J. W.
Thomson & Larsen 5878 (WIS) - Mackenzie District:
Crossley Lakes, Scotter 8058 (H).
U.S.A. Alaska. Aleutian Islands: Attu Island,
Man Schaack 292 (US) - Amchitka Island, Reich &
McCann 77 (F, WIS) - Arctic Coast District:
Franklin Bluffs, J. W. Thomson 6474 (WIS).
13. Bryoria simplicior (Vain.) Brodo &
D. Hawksw., comb. nov.
Alectoria nidulifera f. simplicior Vain., Medd. Soc.
Fauna FI. Fenn. 6: 115 (1881); type: Finlandia,
Lapponia, Paatsjoki. E. A. Vainio, 1878, herb.
Vainio no. 01067 (TUR 001403 lectotype!); basionym.
- Alectoria simplicior (Vain.) Lynge. Norske Vid.-
Akad. Oslo. Mat. -Nat. Kl. 1921 (7): 212 (1921). -
Bryopogon simplicior (Vain.) Gyeln., Feddes Repert.
38: 233 (1935). - Alectoria simplicior f. typica Vain.,
Ann. Univ. Turku., ser. A, 7(1): 8 (1940); nom.
illegit. (Art. 63).
Bryopogon nitidulum f. caespitosa Saviez, Trudy
Student. Nautsch. Fis. Mat. Fak. Univ. Jurjew 3:
40 (1911); type; U.S.S.R., Prov. Archangel, R. Pohle
W538, 1904 (LE lectotype!). - Alectoria nitidula f.
caespitosa (Saviez) Zahlbr., Cat. Lieh. Univ. 6: 396
(1930). - Alectoria lanea f. caespitosa (Saviez)
Gyeln., Nyt Mag. Naturvid. 70; 60 (1932). - Bryo-
pogon bicolor var. nitidulum f. caespitosus (Saviez)
Gyeln., Feddes Repert. 38: 237 (1935).
Alectoria simplicior f. subintricans Vain., Ann.
Univ. Turku., Ser. A, 7(1); 8 (1940); type: Finlandia
ladogensis, Sortavala, Kotiluoto, E. Vainio, 14 July
1926, herb. Vainio no. 01073-5 (TUR 001250 lecto-
type!).
110 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 5 1 . Bryoria simpiicior. Quebec, Upper Seal Lake,
Campbell s.n. (CANL 49618). Scale 0.5—1 .0 mm.
Bryopogon simpiicior f. aibidosorediosus Gyeln.,
Acta Geobot. Hung. 2: 166 (1937); type: U.S.S.R.,
Sibirien. Jakutskaja A.S.S.R., Uralgebirge. Elbyn-
torsee, W. Safrasknikow. 9 September 1919 (BP
33.975 holotype!).
Alectoria simpiicior var. lapponica Gyeln., Nyt
Mag. Naturvid. 70: 62 (1932): type: Suecia. Lapponia
tomeensis, par. Jukkasjärvi, prope lacum Aptasjärvi,
ad ramulos Pini desiccatos, G. Läng, Krypt. Exs.
Vindob. no. 1977 (BP 33.962 holotype!). - Bryopogon
simpiicior f. lapponicus (Gyeln.) Gyeln., Feddes
Repert. 38: 234 (1935).
Alectoria nana Mot., Bryologist 67: 16 (1964);
type: Alaska, Alaska Range District. Mt. McKinley
National Park, Wonder Lake, H. Krog 4779, 14
August 1957 (O lectotype!); nom. inval. (Arts. 36,
37). - Bryopogon nanus (Mot.) Bystr., Ann. Univ.
Mariae Curie-Sklodowska, C, 26: 371 (1971); nom.
inval. (Arts. 33, 36, 37).
Thallus caespitose, sometimes becoming
decumbent, 2^1(-5) cm long; branching mainly
isotomic dichotomous from the base, angles
between the dichotomies usually acute;
branches even in diameter, usually straight,
rarely becoming slightly foveolate, 0.2-0. 4 mm
diam.; cervine brown to dark brown or almost
black, concolourous; base persistent. Fig. 51.
Lateral spinules with slightly constricted
bases, sparse to frequent, becoming abundant
in some forms. Pseudocyphellae absent. Soralia
abundant, fissurai, usually broader than the
branches on which they occur, white to
brownish black or more usually greenish black,
normally without any spinules but sometimes
bearing irregular, often contorted, non-isidiiform
spinules.
Apothecia and pycnidia unknown.
Cortex, medulla and soralia K — , C — , KC — ,
PD -. Contains no lichen products.
Ecology. Characteristically on well-lighted twigs
and small branches of Picea mariana and Larix
laricina in the open black spruce muskeg of
the northern boreal forest, but occasionally on
other trees in that community (e.g., Picea
glauca. Betula spp., Populus balsamifera). We
have seen no specimens from Abies balsamea,
a tree very common in the southern boreal
forest. In the west, Bryoria simpiicior is often
found on Pinas contorta. Rarely, the species is
collected from the ground or rock (on several
occasions, associated with B. c Italy beiformis),
but it is often found on lignum. Krusenstjerna &
Santesson (1950) have remarked that, in the
subalpine birch forests of Jämtland (Sweden),
this species and "A. jubata ” are clearly
restricted to the portions of the birch trees
above the highest level of winter snow.
This species almost invariably grows mixed
with B. lanestris and frequently with B.
furcellata as well.
Distribution. Northern boreal to hemiarctic
zone from Newfoundland to Alaska, with
scattered occurrences in the western mountains
south to Colorado, and in the Shickshock
Mountains of Quebec (Fig. 52). The other
Quebec localities (e.g., in Parc des Laurentides)
undoubtedly represent edaphically northern
boreal outliers. The species can occur in arctic
localities on rock, as it does in southwest
Greenland (Dahl 1950), but its main distribution
even in Europe is clearly in the northern
boreal forests (Ahlner 1948. Poelt 1969). Ahlner
(1948) cites published reports from Siberia,
Kamchatka and Japan, and we have also seen
collections from Mongolia (Golubkova [LE]);
the species can thus be regarded as circum-
boreal, apparently with continental rather than
oceanic tendencies (Ahlner 1948, Ahti & Hawks-
worth 1974).
Discussion
Bryoria simpiicior is a clearly defined species
which is only likely to be confused with
unusually dark specimens of B. furcellata
lacking their characteristic tufts of spinules in
the soralia, or with young plants of B. lanestris,
a species with which B. simpiicior is frequently
entangled. B. simpiicior is always PD — through-
out. {B. furcellata is PD + red throughout.
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 1 1 1
F-ig. 52. Bryoria simplicior. N. American distribution.
and the soralia of B. lanestris are PD + red
whilst its thallus is PD-.) The greenish black
soralia so characteristic of B. simplicior are
unknown in any other North American species.
(Soralia in both B. furcellata and B. lanestris
are white or flecked with black.)
Attention is drawn to Ahlner's (1948) detailed
discussion of this species.
Motyka (1964) indicated that the soralia of
“A. nana", treated as a synonym of B.
s implicior here, were PD + red but this was
due to some of the material he annotated as
(his being mixed with B. lanestris.
Selected specimens
Exsiccatae. Brodo: Lieh. Can. Exs. 2: Ontario ,
Cochrane District: Moosonee, Fabiszewski M26/127
(CANL).
Canada. British Columbia. Coastal District:
Garibaldi Mountains, Whistler Mountain, Brodo 13939
(CANL) - Fraser River Basin: W of Marquart Lake,
Scotter 9848 (W1S) - Liard River Basin: Cassiar,
Szczawinski 684-8 (US, O) - Okanagan District:
Princeton, Brodo 7815 (p.p.) (CANL) - Alberta.
Banff National Park, canyon of Johnson Creek.
Weber S23.592 (CANL. COLO) - 125 miles W of
Edmonton. Turner s.n., I Aug. 1959 (LAM) -
Waterton Lakes National Park, Redrock Canyon,
lmshaug 6103 (W1S) - Saskatchewan. Hasbala Lake
Region, Argus 953-62 (COLO) - Prince Albert
National Park, Waskesiu, Jesberger 1171 (SASK) -
Manitoba. Brochet, Scotter 3031 (W1S) - Fort
Churchill, Crum & Schofield 6889 (CANL) -
Ontario. Cochrane District: Mouth of the Moose
River, Brodo 14748 (CANL) - Kenora District:
SE of Fort Severn. Ahti 3726 (H) - Quebec. Cté
Frontenac: Woburn, Masson 6030B (QUE) - Cté
Gaspé-Est: Penouillc Point, Gaspé. Brodo 18706
(p.p.) (CANL) - Parc des Laurentides: 70°47'W,
47°39'N, Scotter 6515 (H) - Cté Témiscouata: St-
Juste-du-Lac, near Lac Témiscouata, Masson 1 1403B,
(WIS) - Territoire de Mistassini: Otish Mountains,
Mont du Lagopède, Shchepanek 7I-L-58 (p.p.)
112 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 53. Bryoria tenuis. A specimen from the tundra
showing some pale tips. N. W. T., Spence Bay,
McGrath 18 (CANL). Scale 0. 5-1.0 mm.
(CANL) - Territoire du Nouveau Québec: Chimo
Air Base, Marr 5 (COLO) - Newfoundland. Central
Labrador: Goose Bay, Haufe 25098 (US) - Knob
Lake, J. W. Thomson 13730 (WIS). (See also Ahti &
Hawksworth 1974). - Yukon. 40 miles W of Watson
Lake, Ahti 23492 (H) - British Mountains, Trout
Lake, J. W. Thomson & Larsen 14758 (WIS) -
Mountains N of the McQuesten River, Christie s.n.
(CANL, LAM) - Northwest Territories. Mackenzie
District: 3 miles SE of Indian Mountain Lake.
Cody 16024 (CANL, WIS) - Keewatin District:
Ennadai Lake, J. W. Thomson 11803 (CANL, US.
WIS).
U.S.A. Alaska. Alaska Range District: Dry Creek,
63°53-59'N, 147 0 20-35'W, Viereck 6000 (ALA,
COLO, CANL, WIS) - Arctic Coast District: Umiat,
Llano 5 1 1 x (US) - Bering Sea District: Safety
Lagoon, Krog s.n., 22 June 1969 (O) - Central
Yukon River District: Tanana River at mouth of
Delta Creek, Viereck 7567F (ALA. WIS) - Western
Pacific Coast: Nanwhyenuk Lake, Muller 855 (p.p.)
(US) - Colorado. Larrimer or Middle Grand County:
Rocky Mountain Park, Summit Simpson Pass, Sanson
18 (FH) - Idaho. Payette County: Payette Lakes,
A. H. Smith I6076C (WIS) - Montana. Sanders
County: Fishtrap Creek, Taylor 7071 (p.p.) (WIS).
14. Bryoria tenuis (Dahl) Brodo &
D. Hawksw., comb. nov.
Alectoria tenuis Dahl, Meddl. Gronl. 150(2): 144
(1950); type: Greenland, Julianehaab District, Igalik-
fjord, Eqaluit, E. Dahl, 9 August 1937 (O holotypel);
basionym. - Bryopogon tenuis (Dahl) Bystr., Ann.
Univ. Mariae Curie-Skfodowska, C, 26: 271 (1971);
nom. inval. (Art. 33).
Alectoria bicolor var. subbicolor Mot., Bryologist
67 : 9 (1964); type: Alaska, Eastern Pacific Coast
District, Ketchikan. Deer Mt., lower part, H. Krog
6252, 3 September 1957 (O lectotypel); nom. inval.
(Arts. 36, 37).
Thallus erect to decumbent, or rarely sub-
pendent, 4-6(-12) cm long; branching isotomic
dichotomous towards the base but often
tending to become anisotomic dichotomous
towards the apices, angles between the basal
dichotomies usually obtuse, angles between
the apical dichotomies usually acute, third
order branches usually absent; branches terete,
even in diameter, sometimes becoming slightly
compressed towards the base, 0.3— 0.4(— 0.5)
mm diam.; basal branches becoming black,
apical branches pale brown to brown (always
paler than the basal parts); base often dying.
Fig. 53.
Perpendicular lateral spinules with slightly
constricted bases present, sometimes poorly
developed. Soralia absent.
Pseudocyphellae abundant to scanty, fissurai,
plane to slightly raised, usually dark and in-
conspicuous.
Apothecia very rare (known from one
collection from Alaska, Krog 6252), lateral,
appearing geniculate; excipulum thallinum con-
colourous with the thallus, thin, becoming
excluded as the ascocarps mature; disc concave
at first, becoming convex when mature,
yellowish brown to reddish brown, to 1.5 mm
diam. Thecium c. 70 gm high, tall, asci c.
45 x 15 gm, epithecium I K1 + aeruginose. Spores
8 per ascus, subglobose to ellipsoid, hyaline,
simple, with a rather thick (1 /urn) wall, (6-)
7-9.5 x 5-7 /am. Pycnidia not seen.
Inner cortex and medulla K-, C-, KC-,
PD + red at least in parts (perhaps rarely PD — ).
Contains fumarprotocetraric acid.
Ecology. Characteristically on mossy trees and
rocks in moist conifer forests, but also found
in rock crevices and on mossy rock surfaces
in more exposed subalpine sites. In the southern
Appalachian mountains, it is apparently re-
stricted to the fir forests above 1 800 m. All
the specimens we have seen from that area
are from Abies (presumably A. fraseri). The
arctic collections (e.g., McGrath, Spence Bay.
OPERA BOT. 42 (1977)
Brvoria Brodo & D. Hawksw. 113
Fig. 54. Bryoria tenuis. N. American distribution.
N.W.T.) were generally growing between rock
fragments much as Alectoria ochroleuca. They
are fairly typical in morphology, although
sometimes lacking in the long, curled tips
described by Dahl (1950) in the original Green-
land material. In the alpine areas of the Tyrolian
Alps, the species grows on boulder surfaces
mixed with other saxicolous foliose and fruticose
lichens.
Distribution. Oceanic localities along the
Appalachian Mountains, and along the coasts
°f British Columbia and Alaska, with scattered
arctic populations in northern Alaska, Keewatin
(N.W.T.) and southwestern Greenland (Fig.
^4). In Europe, B. tenuis does not seem to be
8 - Opera Botanica nr 42
as clearly oceanic as it does in North America.
Although it is known from western Norway,
Jprgensen & Ryvarden (1970) point out that the
localities are in the mountains. The species
is also known from a few localities in Sweden
(Jorgensen 1972). One of us (l.M.B.) has seen
and collected it on rocks near tree-line in the
Austrian Alps. Its occurrence in the American
arctic is not entirely surprising since the species
was described from the tundra of southwest
Greenland (Dahl 1950). A very similar distri-
bution is shown by the clearly oceanic crustose
lichen, Placopsis gelida (L.) Linds. (Thomson
1972). We have not seen the specimen from
the Aleutian Islands (Adiak Island) reported
by Jorgensen ( 1972) but presume it is correct.
114 Alectoria in North America
OPERA BOT. 42 (1977)
Discussion
Bryoria tenuis is most closely allied to B.
bicolor. The decumbent habit and the third-
order perpendicular branches in B. bicolor
serve to distinguish the two species. The
branching pattern of B. tenuis is not unlike
that of B. nadvornikiana which is easily
distinguished by its chemical reactions (K +
bright yellow. KC + red, PD+ orange-yellow)
and much paler apical parts. More detailed
information on the differences among these
three species may be found in Dahl (1950)
and Jqrgensen & Ryvarden (1970).
Although some specimens of B. tenuis
appear to be PD — on the basis of routine
testing, a careful re-examination using the
methods described under Chemistry (Figs. 17,
18) shows them all to be PD + red at least in
some of their pale, apical branches. This is
also true of the type specimen of B. tenuis
which, contrary to the findings of earlier
authors, proves to be PD + red in part.
Consequently, we do not believe there to be a
truly “inactive strain" of this species as was
suggested by Krog (1968), but would stress
that the amount of fumarprotocetraric acid in
this species may sometimes be so small as to be
barely detectable in routine testing.
Motyka (1964) treated only specimens he
found to be PD— as "Alectoria tenuis",
referring PD + plants to "A. bicolor var.
subbicolor” . As pointed out by Krog (1968)
and Jqrgensen (1972), such a view is in any
event entirely unacceptable as it ignores the
morphological differences which readily serve to
distinguish these two species.
Apothecia have not previously been reported
in this species and were seen by us only in
the single collection cited above.
One specimen of B. tenuis, collected in the
White Mountains of New Hampshire by
Tuckerman (FH-Tuck.!), was superficially much
like B. smithii but lacked soralia and the typical
clusters of isidioid spinules. Per M. Jorgensen
(in litt.) examined the material at our request
and concluded that it did not seem to be B.
smithii, but was more likely a rather coarse
B. tenuis. The thallus is completely PD- and
revealed nothing in TLC; it is therefore the
only specimen of B. tenuis we have seen totally
lacking in fumarprotocetraric acid.
Selected specimens
Exsiccatae. Brodo: Lieh. Can. Exs. 83: British
Columbia. Moresby Island, Brodo 18507 (CANL).
Canada. British Columbia. Chilkat River Basin:
Near Chilcoot Pass, Williams 69 (NYBG) - Islands
North District: Graham Island: Awun Lake. Brodo
18478 (CANL) - Moresby Island: Between Sandspit
and Copper Bay, Brodo 12880 (CANL) - Quebec.
Cté Gaspé-Nord: Mont Albert Nord. Gauthier 2637
(CANL) - Cté Matane: Between Mount Mattaouisse
and Mount Collins, Collins 2155 (FH)-Cté Saguenay:
Archipel de Mingan, Marie-Victorin & Rolland-
Germain 18,298 (p.p.) (US) - Newfoundland.
Placentia East District: St. Bride's, Ahti 8867 (H) -
lie Miquelon. Delamar S211 (S) - Northwest Terri-
tories. Keewalin District: Spence Bay, McGrath 18
(CANL) - Thelon River, Aberdeen Lake. Scotter
4233 (W1S).
U.S.A. Alaska. Arctic Coast District: Pitmegea
River, Cape Sabine, J. W. Thomson 5530 (WIS) -
Central Pacific Coast District: Prince William Sound.
Pigot Bay. Krog 2206 (O) - Eastern Pacific Coast
District: Sitka, Sitka National Monument, Krog
6245 (O) - New Hampshire. County unknown: White
Mountains, Herb. E. Tuckerman 676 (EH) - North
Carolina. Yancey County: Black Mountains, Mt.
Mitchell State Park, Dey 1197 (DUKE) - Oregon.
Clatsop County: Saddle Mountain State Park, Pike
3763 (OSC) - Tennessee. Sevier & Cocke Counties:
Old Black in the Great Smoky Mountains National
Park, Dey 6628 (DUKE).
Bryoria sect. Implexae (Gyeln.) Brodo &
D. Hawksw., comb. nov.
Bryopogon sect. Implexae Gyeln., Feddes Repert.
38: 223 (1935); basionym. - Bryopogon sect. Bryo-
pogon subsect. Implexae (Gyeln.) Bystr., Ann. Univ.
Mariae Curie-Sklodowska, C, 26: 274 (1971).
Bryopogon sect. Fuscidulae Bystr., Ann. Univ.
Mariae Curie-Sklodowska, C, 26: 271 (1971); type:
B. fuscidulus (Arnold) Bystr. nom. inval. [= Bryoria
capillaris (Ach.) Brodo & D. Hawksw.]. holotype.
Bryopogon sect. Flexuosae Bystr., Ann. Univ.
Mariae Curie-Sktodowska, C. 26: 272 (1971); type:
B. flexuosus (Mot.) Bystr. nom. inval. [1= Bryoria
pseudofuscescens (Gyeln.) Brodo & D. Hawksw.
aggr., see Hawksworth 1972], holotype.
? Alectoria subgen. Bryopogon sect. Pellucidae
Mot., FI. Polska, Porosty 5(2): 39 (1962); type:
A. pellucida Mot. [=Bryoria sp., see p. 59], holotype.
- Bryopogon sect. Pellucidae (Mot.) Bystr., Ann.
Univ. Mariae Curie-Sklodowska, C, 26: 275 (1971),
as “Mot."
Type species: Bryoria implexa (Hoffm.) Brodo &
D. Hawksw. (syn. Usnea tax. vag. implexa Hoffm..
Alectoria implexa (Hoffm.) Nyl.; lectotype designated
by Bystrek 1971 p. 274).
Thallus decumbent to subpendent or pendent;
greyish green, fuscous or brown to dark brown,
generally not becoming blackened except in
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 115
fragmentation regions. True lateral spinules and
isidia absent. Soralia absent in most species,
frequent in others, mainly tuberculate, white.
Pseudocyphellae always present, very variable
in size, fusiform and inconspicuous to spiralled
and readily visible, depressed or very slightly
elevated, usually whitish. Cortex composed of
broad hyphae with very little matrix. Apothecia
rare, unknown in many species; margins not
ciliate; discs orange-brown to reddish brown
or rarely white. /3-orcinol depsidones other
than fumarprotocetraric acid frequent; atranorin
and/or chloroatranorin, and fumarprotocetraric
acid sometimes present in traces; pulvinic
acid derivatives absent.
Discussion
This section comprises about 12 species
characterized principally by the occurence of
/3-orcinol depsidones other than fumarproto-
cetraric acid, small pseudocyphellae, and a
characteristic cortical structure which tends to
make the branches more friable than usual in
the genus. Two species with conspicuous spiral
pseudocyphellae (B. pseudocapillaris and B.
spiralifera ) are being included here with some
hesitation.
As a result of our studies it has become
clear that this group is much more varied than
previously supposed. While most of the taxa
accepted here are readily recognizable,
taxonomic problems exist with several of them
as will be evident from the discussions following
individual species.
This section is alone amongst Alectoria and
its allied genera in including species able to
Produce a tridepside (gyrophoric acid): B.
friabilis (consistently), and B. pikei (some
specimens).
15. Bryoria capillaris (Ach.) Brodo &
D. Hawksw., comb. nov.
Parmelia jubata ß. capillaris Ach., Meth. Lieh.:
273 (1803); type: Suecia (H-Ach., upper specimen,
lectotype!); basionym. - Alectoria capillaris (Ach.)
Cromb., J. Bot., Lond. 9: 177 (1871).
For further synonyms see Hawksworth (1969 pp.
393-394, 1972 pp. 209-211).
Thallus pendent, commonly 10— 15(— 30) cm long;
branching irregular, isotomic dichotomous to
anisotomic dichotomous, angles between the
main branches usually acute; branches even to
uneven in diameter, sometimes becoming
slightly twisted or compressed especially
towards the base, capillaceous. 0. 1— 0.3(— 0.5)
mm diam.; greenish grey to grey, sometimes
becoming brown to dark brown, matt to slightly
shiny.
True lateral spinules absent. Soralia absent
in North American material. Pseudocyphellae
usually present, inconspicuous on pale morpho-
types, fusiform, white, 0.1-0.25 mm long.
Apothecia rare in North American material,
seen in a single collection (Montana, Glacier
National Park, H. & V. Bailey 17 p.p. [H]),
lateral, up to 1.5 mm in diameter; discs
becoming strongly convex, light brown,
epruinose; excipulum thallinum concolourous
with thallus, soon becoming reflexed and
disappearing. Spores 8 per ascus, hyaline,
ellipsoid to subglobose, simple, 5. 3-6.8 x 4. 0-4. 5
pm. Pycnidia not seen.
Cortex, medulla and soralia K + bright yellow,
C+ pink or -, KC+ red, PD+ yellow.
Contains barbatolic acid, ± alectorialic acid.
Ecology. On trees of all kinds, especially
Picea, Pseudotsuga, Abies and Tsuga species,
mainly in well-lighted portions of mature
coniferous forests. Collected once from rock
(Michigan, Isle Royale, Wetmore 1597 [MSC]).
In oceanic or suboceanic forests, such as those
on the Queen Charlotte Islands (British
Columbia) the species grows in great abundance
forming large, grey clumps hanging from low
branches.
Distribution. Southern boreal to north temperate
(hemiboreal) distribution in the east; Rocky
Mountains of Alberta, British Columbia, Idaho
and Montana; and coastal forests from California
to southern Alaska (Fig. 55). The North
American distribution shows distinct clumping
in the more humid forest zones: coastal and
interior Tsuga-Thuja forests; Lake Superior
region corresponding to the “strongly humid
southern boreal zone” (Ahti 1964); the St.
Lawrence region; and the Atlantic coastal
region. Scattered middle and northern boreal
forest localities are also known. Hawksworth
(1972) comments on the world distribution of
B. capillaris, citing it from northern and
southern Europe, the Canary Islands, and Japan.
116 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 55. Bryoria capillaris: N. American distribution. A determination uncertain.
The species is probably circumboreal in the
Northern Hemisphere.
Discussion
Our study has revealed a number of problems
in the systematics of this species arising from
both morphological and chemical variability.
Firstly, while B. capillaris is frequently (but
by no means always) sored iate in Europe, no
sorediate North American specimens have been
found by us; this character is therefore region-
ally useful for the separation of B. capillaris
from exceptionally pendent specimens of B.
nadvornikiana which is almost invariably
sorediate in North America. Secondly, there are
variations in the degree of shininess, the
thickness of the main stems, and colour.
Thirdly, these latter variations may perhaps be
related to differences in the ratio of alectorialic
to barbatolic acid and thus to differences in the
intensity of the K+ reaction. (In the presence
of alectorialic acid without barbatolic acid,
a K + very pale yellow or K - reaction is
obtained.) The species is clearly very closely
allied to B. pikei (see Discussion under that
species). It also appears to be related to B-
pseudofuscescens, to judge from the discovery
of a collection with both barbatolic and
norstictic acids (Washington, near Dungeness,
A. S. Forster 2552a [FH !] ; mixed with typical
B. capillaris and Sulcaria badia ) which might
even be considered a hybrid (see pp. 33, 71).
When the chemistry of some European
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 117
collections was examined by TLC, further
difficulties came to light. It now appears that
many European specimens (those with soredia)
contain chloroatranorin or atranorin, or both,
and often traces of fumarprotocetraric acid in
addition to the usual American components.
We have not found any atranorin or chloro-
atranorin in any North American material of
this species. The lectotype specimen of B.
capillaris is mainly esorediate but has a few
basal sorediate branches. Through the courtesy
of Mr. O. Vitikainen. we have been able to
examine both sorediate and esorediate portions
of it by TLC. These prove to have similar
chemistries with chloroatranorin, possibly a
trace of atranorin (a weak spot evident in
PD but not under UV alone), alectorialic acid,
barbatolic acid, and a spot which may be
fumarprotocetraric acid (very faint).
Material identical to the North American
B. capillaris (i.e., esorediate, containing
alectorialic and barbatolic acids and lacking
atranorin and chloroatranorin) does, however,
occur in parts of Europe (e.g., Arnold, Lieh.
Exs. no. 914a, sub A. cana f. fuscidula [M !]).
If this population later proves to be of taxonomic
significance at the species level, type specimens
of the numerous synonyms of B. capillaris
will require examination by TLC before the
appropriate name can be determined.
Specimens corresponding to Hawksworth s
(1972) concept of A. capillaris f. fuscidula
occur in North America. We have reinvestigated
the status of this name, however, since Motyka
has applied the epithet in herbaria to specimens
more properly identified as B. kuemmerleana
(Gyeln.) Brodo & D. Hawksw. or. more
rarely, B. capillaris and B. pseudofuscescens
as understood by us. This confusion appears to
have arisen primarily from Motyka (1960) con-
sidering the epithet to have been published in
1884 (when it in fact was validated in 1882)
and to his treating as the holotype a specimen
in M which was collected in the type locality
in 1896 (M!) (rather than in 1880 as stated
by Motyka I960). The lectotype designated
by Hawksworth (1972) was at least an isotype
and could have been considered the holotype
apart from the fact that Arnold did not annotate
any one collection “typus”. This specimen,
and duplicates of it, consist of mixture of
B- capillaris and fragments of B. kuemmerleana.
in which B. capillaris clearly predominates.
An examination of all available material in M
which was collected by Arnold and was
annotated by him as fuscidula, showed that he
had not applied this name consistently. In view
of this confusion, it is probably in the best
interests of taxonomic stability to consider
this epithet a nomen ambiguum to be rejected
in accordance with the provisions of Art. 69.
It will be evident from the above comments
that many critical studies considering both
morphological and chemical characters on a
world scale need to be carried out in order to
ascertain the taxonomic importance of the
findings reported here. We consequently feel
that it would be premature for us to propose
any changes in the nomenclature or circum-
scription of B. capillaris and its allied chemo-
and morphotypes here.
Selected specimens
Exsiccatae. Brodo: Lieh. Can. Exs. 78: British
Columbia, Moresby Island, Brodo 12247A (CANL) -
79: British Columbia, Creston, Brodo 15110 (CANL) -
?80: British Columbia, Earl's Cove, Brodo 8330
(CANL) - Lieh. California Exs. (no number): Cali-
fornia, Marin County Sausalito. M. A. Howe 79
(COLO. DUKE, FH. NYBG).
Canada. British Columbia. Islands North District:
Moresby Island: Harriet Harbour. Brodo 12551
(CANL) - Islands South District: Little Qualicum
Falls Provincial Park, Ahti 6528 (H) - Coastal Area:
Alice Lake. Ohlsson 436A (CANL) - Columbia River
Basin: Valley of Illecillewaet R.. Brodo 7668 (CANL)
- Dean River Basin: Stuie. Tweedsmuir Lodge,
Ohlsson 2165 (CANL) - Fraser River Basin: Near
Malakwa, Bird 15275 (p.p.) (WIS) - Kootenay
River Basin: Creston, Brodo I5123A (CANL) -
Okanagan: 34.6 miles S of Princeton. Brodo 8503B
(CANL) - Peace River Basin: Alaza (sic) (Aleza)
Lake, Kujala s.n., 24 July 1931 (H) - Skagit River
Basin: Manning Park. Ahti 15399 (H) - Skeena River
Basin: Hazelton. Ohlsson 2962B (MSC) - Alberta.
Bragg Creek. Bird 10966 (p.p.) (CANL) - Saskatche-
wan. Cypress Hills Park, Looman s.n., 7 April 1959
(LAM) - Ontario. Algoma District: Near Brule,
Grassl 1674 (MSC) - Cochrane District: Near Moose
Factory, Brodo 14694 (CANL) - Kenora District:
Fort Severn, Ahti 3950 (H) - Thunder Bay District:
Shebandowan, Brodo 13632 (CANL) - Quebec.
Cté Abitibi Ouest: Guyenne. Brodo 16889 (CANL) -
Cté Gaspé Est: Cap-des-Rosiers-Est, Brodo 18668
(CANL) - Cté Rimouski: Ile au Massacre, Masson
14162 (WIS) - Prince Edward Island. Kings County:
Montague, Ireland 10207B (CANL) - Queens County:
14 miles NW of Charlottetown, Fabiszewski s.n.,
22 July 1970 (CANL) - Nova Scotia. Colchester
County: Macoun s.n., 12 June 1883 (CANL). Kings
118 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 56. Bryoria friabilis. Note the
irregularly broken end of the fila-
ments. B. C., Skeena River,
Macoun s.n. (CANL 16156). Scale
0.5 mm.
County: Wolfville, Gillett 15683C (CANL) - New-
foundland. Labrador West District: Schefferville,
Brodo 9343B (CANL) - Humber East District:
Cormack, Ahti 5930 (H) - Northwest Territories.
Mackenzie District: Artillery Lake, J. W. Thomson &
Larsen 12623 (WIS).
U.S.A. Alaska. Central Pacific Coast District:
Kenai Peninsula, Seward, Krog 2195 (O) - Eastern
Pacific Coast District: Auke Lake, Imshaug 28490
(MSC) - California. Humbolt County: Horse
Mountain, Brodo 20516 (CANL) - Colorado.
Chaffee County: Near Buena Vista, Imshaug 11679
(MSC) - Idaho. Latah County: Bovill, Schroeder
L367, 13 Sept. 1969 (SCHROEDER) - Maine.
Hancock County: Salisbury Cove (sic) (Salsbury
Cove), Plitt 167 (US) - Michigan. Keweenaw County:
Isle Royale National Park, Capt. Kidd L, Wetmore
4674 (MSC) - Minnesota. Cook County: Grand
Portage L, Fink 17 (H, US, FH) - Montana. Flathead
County: Glacier National Park. Grinnell Creek,
Imshaug 5547 (CANL, LAM, WIS, US) - Oregon.
Lane County: Willamette Pass, Becking 6308194
(BECKING) - Vermont. Windham County: Stratton,
Moore 1532 (US) - Washington. Kittitas County:
Wenatchee National Forest, Hoffman OP-27 (US) -
Whitman County: Kamiak Butte, Esslinger 1774C
(WIS).
16. Bryoria friabilis Brodo & D. Hawksw.,
sp. nov.
Thallus pendulus, vel raro ubi saxicola decumbens,
friabilis, ad tactum disrumpens, ad 15 cm longus;
rami praecipue aniso-dichotomiter fastigiati et in
angulos obtusos ad proxime 90° divergentes,
plerumque diametro inaequales, valde contorti,
rugosi et foveolati, ramis principalibus crassis
plerumque ramis brevibus gracilibus perpendiculariter
instructis (similibus iis Bryoriae fremontii); planta
pallidissime cervineo-brunnea, in herbario deni-
gricans.
Rami laterales spiniformes veri, isidia et soralia
desunt. Pseudocyphellae copiosae, brunneolae vel
albae, late fusiformes, circa ramos tortuosae. Cortex
tenuis, e hyphis distincte undulatis compositus,
parietibus cellularum subtilibus.
Apothecia rarissima, lateralia, 0.35-0.5(-0.8) mm
diametro, saepe aliquantum elongata: discus convexus
et reflexus, albus vel dilute fuscus, epruinosus;
excipulum proprium hyalinum vel brunneum. Asco-
sporae incolores, ellipsoideae, simplices, parvae,
4. 2-5.0 x 2.4-3.0 gm. Pycnidia ignota.
Cortex K - , C + rubescens vel roseus, KC + roseus,
PD-; medulla K — , C — , KC — , PD— ; apothecia
PD + lutea, K-, C-, KC + rosea. Cortex
thalli acidum gyrophoricum continens; epithecium.
hyménium et excipulum proprium apothecii acidum
alectorialicum continentia.
Holotypus: America septentrionalis, U.S.A.,
Washington, Gate, in Quercu, rara, A. S. Foster
s.n., 14 October 1911 (NYBG).
Thallus pendent, or rarely decumbent when
on rocks, brittle, breaking when touched, to
15 cm long; branching mainly isotomic dicho-
tomous, angles between the dichotomies obtuse
to almost 90°, branches usually uneven, very
contorted, wrinkled and foveolate, main
branches often with short, slender, perpendic-
ular side branches (similar to those of B.
fremontii ); very pale cervine brown, becoming
darker in the herbarium. Fig. 56.
True lateral spinules, isidia, and soralia
absent. Pseudocyphellae abundant, brownish to
white, broadly fusiform, twisting around the
branches. Cortex thin, composed of distinctly
undulating thin-walled hyphae.
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 119
Apothecia rare (seen only in a single speci-
men: British Columbia, Vancouver Island,
Nimpkish Valley, Stevenson 747a, 2 Nov. 1975
[UBC]), lateral. 0.35-0.5(— 0.8) mm in diameter,
often somewhat elongated along the lilament;
discs strongly convex and reflexed, white to
very pale buff (or rarely brown in apothecia
having a pigmented excipulum), epruinose;
excipulum thallinum concolourous with disc and
hardly distinguishable from it; excipulum
proprium hyaline, or brown in some apothecia.
Spores eight per ascus, hyaline, ellipsoid to
subglobose, simple, very small, 4.2-5.0X2.4-
3.0 /am. Pycnidia not seen.
Cortex K-, C+ red or pink, KC+ pink,
PD - ; medulla K - , C - , KC - , PD - ; apothecia
PD + deep yellow, K — , C — , KC + pink.
Contains gyrophoric acid in the thallus cortex
and alectorialic acid in the epithecium, hy-
ménium and excipulum proprium of the apoth-
ecia.
Ecology. On trees in exposed habitats, from
sea level to I 500 m elevation; very rarely on
rocks or shrubs.
Distribution. West coast-east coast disjunct
distribution, with a few isolated localities in
the interior montane forests (Fig. 57 A). It is
likely that this North American endemic was
once widespread (see general section on
Distribution, category 1 1).
Discussion
Bryoria friabilis is almost unique in the Alec-
torioid genera in producing gyrophoric acid.
Its relationships are obscure, but on the basis
of external morphology it is not unlike B.
bnplexa in many respects. Some difficulty
may be encountered in distinguishing B.
friabilis from some morphotypes of B. fre-
rnontii and B. pikei in the absence of chemical
examinations; this latter species sometimes has
trace amounts of gyrophoric acid in addition
to its main component (alectorialic acid).
In the single fertile specimen examined (cited
above), gyrophoric acid was produced in the
thallus filaments and alectorialic acid was
produced in certain tissues of the apothecia.
The filaments were entirely devoid of the latter
substance (verified by TLC). Assuming that this
chemical distribution is characteristic of the
species as a whole, this would provide additional
evidence for the close relationship between
B. friabilis and B. pikei. The restriction of
lichen substances to apothecial tissues is well
known in many genera of lichens (Lecanora:
Imshaug & Brodo 1966, Letharia: W. Culberson
1969a, Ramalina: Krog & Swinscow 1974,
etc.) and should not be regarded as very unusual.
It should also be noted that B. friabilis is
the only species of Bryoria with white
apothecial discs; its spores are the smallest
seen in the genus.
An extremely perplexing specimen from the
Olympic Mountains in Washington (Thomson
16261 [WIS]), which we are referring to B.
friabilis with some hesitation, gives the im-
pression of being a hybrid between this species
and B. glabra. Although this collection has the
very fragile branches and gyrophoric acid of
B. friabilis, at the same time it possesses the
olivaceous and regularly dichotomous, smooth,
even, and very shiny branches and white
soralia of B. glabra. The soralia are, however,
tuberculate rather than fissurai (as is usual for
B. glabra ) and react C+ red and PD - ,
Gyrophoric acid could be detected by TLC
examination only from branches bearing
soralia; esorediate branches contained no lichen
products. The colour and evenness argue
against this being a “white-sorediate” B. fre-
montii (see Discussion under that species).
B. friabilis is normally very uneven and twisted,
only rarely shiny, with a quite different
branching system, and it normally lacks soralia.
Furthermore, the gyrophoric acid in B. friabilis
occurs in the cortex of the whole thallus.
The area from which this specimen was collected
is remarkable for its abundance of B. glabra
(it is also the type locality for that species)
and is also within the distributional range of
B. friabilis. B. glabra is very rarely fertile and
B. friabilis has only once been found with
ascocarps, but, on the basis of the above
information, it is conceivable that this specimen
represents some sort of hybridization between
these two species. It may prove to represent
a further undescribed gyrophoric acid-con-
taining Bryoria but with only a single collection
available, we consider it premature to describe
this as a new taxon despite its high degree of
distinctness.
120 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 57. A: Bryoria friabilis. Known distribution, A determination uncertain. O locality uncertain. - B:
B. implexa. N. American distribution. A determination not confirmed chemically.
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 121
Specimens
Canada. British Columbia. Coastal Area: Garibaldi,
Brodo 8179 (CANL) - Fraser River Basin: Wells
Gray Provincial Park. Clearwater Lake, Ahti 7332
(H) - Islands South District: Vancouver Island,
Little Qualicum Falls Campground. Tibell 5076 (UPS)
- Sidney, Macoun 77 (CANL. FH) - Salt Spring
Island. Bird 25604 (UAC) - Victoria, Macoun s.n.,
6 June 1908 (NYBG) - Skeena River Basin: Skeena
River, Macoun. 2 Oct. 1891 (CANL) - Quebec.
Cté Gaspé-Est: Cap-des-Rosiers-Est, Brodo 18678
(CANL)- Bonaventure Island. Brodo 18691D (CANL)
- New Brunswick. Charlotte County: Campobello, no
collector, s.n., July 1902 (FH) - Newfoundland.
District unknown: Middle Bay, A. Hodgkins s.n.,
1897 (M).
U.S.A. California. Mendocino County: Ukiah,
McClain 39 (US) - Del Norte County: Near Bear
Basin, Tucker 6434 (p.p.) (US) - San Mateo County:
Cahill Ridge, Jordan WJ956A (WIS) - Pilarcitos
Creek Canyon, J. W. Thomson 4754 (WIS) - Santa
Clara County: Devil’s Canyon, J. W. Thomson
4700 (WIS) - Santa Cruz County: Santa Cruz
Mountains, above Saratoga, Herre 491 (590) (p.p.)
(FH) - Santa Cruz Mountains, Herre 491 (425)
(NYBG, US) - Idaho. Bonner County: Priest River
Experimental Forest, Esslinger 282 (WIS). 318 (H) -
Latah County: Dry Creek, Cooke 23060 (WIS) -
Maine. Sagadahoc County: Reid State Park, Taylor
487 (MSC) - Montana. Lincoln County: Yaak River
campsite, J. W. Thomson 16308 (p.p.) (WIS) -
Oregon. Benton County: Corvallis, Hawkins s.n.,
Aug. 1921 (CANL, WIS) - 13 miles W of Corvallis,
Brodo 20923 (CANL) - Lane County: Willamette
National Forest, Pike L-494 (PIKE)- Willamette Pass,
Becking 6308194 (BECKING) - Andrews Exp.
Forest. 44°N, 122°W, Pike s.n. (OSC 31,421) -
Washington. Clallam County: Olympic Peninsula,
Hurricane Ridge, J. W. Thomson 16261 (WIS) -
Klickitat County: Goldendale, Foster 1154 (FH) -
County unknown: Gate, Foster 1977 (NYBG).
'7. Bryoria implexa (Hoffm.) Brodo &
D. Hawksw., comb. nov.
Osnea [tax. vag.] implexa Hoffm., Deutsch. FI.
T 134 (1796); type: sine loc. (MW-Hoffm. 8569
neotype!); basionym. - Usnea implexa (Hoffm.)
Hoffm., Herb. Viv. 2: 453 (1825). - Alectoria implexa
(Hoffm.) Nyl., Lieh. Scand.: 72 (1861).
For further synonyms see Hawksworth (1969 pp.
395-396).
Thallus pendent, 6-15 cm long; branching
irregularly isotomic dichotomous, angles
between the dichotomies acute or obtuse,
(ending to be acute towards the apices; branches
ev en to uneven in diameter, often becoming
(wisted, compressed and slightly foveolate
Specially towards the base, 0. 1-0.3 mm diam.;
pale brown to greenish brown or dark brown,
concolourous or slightly paler towards the base.
True lateral spinules and soralia absent in
North American material. Pseudocyphellae
sparse to abundant, conspicuous, elongate
fusiform, usually depressed, often becoming
twisted around the branches, white, to 1.5 mm
long.
Apothecia and pycnidia unknown.
Outer cortex K — , C-, KC-, PD+ yellow;
medulla K— , C — , KC— , PD+ yellow or — .
Contains psoromic acid.
Ecology. On conifers and lignum in the boreal
spruce-fir forest.
Distribution. Widely scattered in the middle
boreal to southern boreal zones, and in the
western mountains (Fig. 57 B). We have seen
specimens from central, northern, and eastern
Europe (Hawksworth 1969), but as the taxon-
omy of this species is still not entirely settled
(see below), it would be premature to comment
further on its extra-American distribution.
It should be noted that most literature
reports of this species refer to B. capillaris
but that the name has also been applied to
B. trichodes by some early North American
authors.
Discussion
This species is clearly closely allied to B.
pseudofttscescens from which it is distinguish-
able by its chemical components and to some
extent by its colour. The North American
collections of B. implexa tend to be somewhat
more delicate and paler than those from Europe
and always lack soralia (as is often the case
in Europe as well). Extremely pale almost
pigment-deficient plants corresponding to the
European B. osteol'a (see Hawksworth 1972 p.
253 for probable synonyms of this name) have
not been noted amongst the North American
material examined. Heavily sorediate, rather
pale and coarse specimens from central Europe
may be specifically distinct, but their nomen-
clature is obscure at the present time.
Specimens
Exsiccatae. Merrill: Lieh. Exs., Ser. II, 129 (mixed):
Alaska, Fairbanks, Palmer (NYBG).
122 Alectoria in North America
OPERA BOT. 42 (1977)
Canada. British Columbia. Dean River Basin:
Along Mosher Creek, Tweedsmuir Provincial Park,
Ohlsson 1932 (MSC) - Liard River Basin: Fort
Newson, Gillett 2995 (CANL) - Saskatchewan.
Prince Albert Nat. Park. Waskesiu, Jesberger 1224.
733 (WIS), 737 (SASK) - Manitoba. Porcupine
Mountains, Macoun 463? (CANL) -Ontario. Thunder
Bay District: Nipigon River, Macoun s.n., 2 July
1884 (CANL) - Edmondson Lake, N of Port Arthur,
Garton 10027 (CANL) - Kenora District: Attawapsi-
kat, Ahti 3942 (H) - Quebec. Cté Gaspé-Est:
Penouille Point, Brodo 18706 (p.p.) (CANL) - Cté
de Québec/Montmorency: Montmorency River,
Macoun s.n., 30 June 1905 (CANL) - Nova Scotia.
Halifax County: Upper Brookside, East Mt.,
Prince 6240 (FH).
U.S.A. Alaska. Alaska Range District: Cantwell,
Krog s.n., 10 June 1969 (O) - Central Yukon River
District: Cheena Ridge near Fairbanks, J. W.
Thomson 5078 (p.p.) (WIS) - Idaho. Bonner County:
Minar's Bay, Priest Lake, MacDougal 267 (NYBG) -
Maine. Knox County: Camden, Merrill s.n., Oct.
1904 (FH) - Montana. County unknown: John's
Lake, Vreeland 1123 (NYBG) (Specimen not tested
for psoromic acid). - New York. Cattaraugus
County: Lime Lake (Machias?), Clinton s.n., no
date (US).
18. Bryoria nadvornikiana (Gyeln.) Brodo &
D. Hawksw., comb. nov.
Alectoria nadvornikiana Gyeln., Acta Fauna FI.
Univ., ser. 2, I: 6 (1932); type: Rossia carp.,
Turkul, alt. 1850 m. G. Nâdvornik, 1930, herb.
Servit 17.653 (BP 167267 holotype!): basionym. -
Bryopogon imptexus var. nadvornikianus (Gyeln.)
Gyeln., Feddes Repert. 38: 242 (1935). - Bryopogon
nadvornikianus (Gyeln.) Gyeln., Ann. Mus. Nat.
Hung., Bot. 32: 154 (1939). - Alectoria implexa
var. nadvornikiana (Gyeln.) Zahlbr., Cat. Lieh. Univ.
10: 555 (1940).
Bryopogon altaicus Gyeln.. Acta Geobot. Hung.
2: 166 (1937); type: U.S.S.R., Sibérien, Altaigebirge,
N. N. Lawrow, 1927, ex herb. Tomin 20 (H
holotype!). - Alectoria altaica (Gyeln.) Räs., Ann.
Bot. Soc. Zool.-Bot. Fenn. Vanamo 12(1): 34 (1939).
Alectoria altaica var. spinulosa Räs.. Ann. Bot.
Soc. Zool.-Bot. Fenn. Vanamo 12(1): 34 (1939),
as "(Ahln.) Räs.”; type: Suecia, Helsingia, in par.
Alfta, Grannäs, Jägtjäm, ad ramum Piceae, S. Ahlner,
3 July 1936 (S holotype; not seen). - Alectoria
nadvornikiana var. spinulosa (Räs.) Mot., Fragm.
Florist. Geobot. 4: 235 (1958); nom. inval. (Art. 33).
Alectoria karelica Räs., Ann. Bot. Soc. Zool.-
Bot. Fenn. Vanamo 12(1): 34 (1939); type: U.S.S.R.,
Karelian A.S.S.R.. Karelia ladogensis, Kurkijoki,
Lapinlahti. Ristissaari, ad rupicariam, V. Räsänen,
21 May 1937 (H holotype!); nom. inval. (Art. 34).
Alectoria curta f. pallidior Östm., in Hasselr.,
Ark. Bot. 30A (13): 3 (1943); nom. inval. (Arts.
32, 34, 36).
Alectoria nadvornikiana var. extensa Mot., Fragm.
Florist. Geobot. 4: 236 (1958); type: In Tartorum
monte dicta Lysanki, alt. 1300 m, in silva Picearum
fere primitiva Augusto 1925 (holotype; not seen).
Alectoria nadvornikiana var. eciliata Mot., Fragm.
Florist. Geobot. 4: 237 (1958); nom. inval. (Art. 37). -
Bryopogon eciliatus (Mot.) Bystr., Ann. Univ. Mariae
Curie-Skfodowska, C, 26: 271 (1971); nom. inval.
(Arts. 33, 37).
Thallus caespitose to subpendent (or rarely
pendent in unusually long corticolous speci-
mens) usually 4—7 (—9 ) cm long; branching
isotomic dichotomous towards the base but
strictly anisotomic dichotomous towards the
apices, angles between the basal dichotomies
obtuse, angles between the apical dichotomies
acute (Fig. 1 D, E); branches terete, even in
diameter, straight, or more rarely flattened at
the axils and somewhat twisted; (0. 1— )0.2— 0. 3
(-0.4) mm diam.; basal branches becoming
black, apical branches pale greyish green to
pale brown or olivaceous brown, or rarely
dark brown to almost black (always paler
than the basal parts); base persistent.
Lateral, perpendicular, spinulose branches
present, sparse to frequent (Fig. 3). Pseudo-
cyphellae absent or sparse, to abundant, plane,
0.25-0.50 p. m long. Soralia abundant to sparse,
or rarely absent, tuberculate or sometimes
fissurai, usually broader than the branches on
which they occur, white or greenish white,
to 1 mm long.
Apothecia and pycnidia not seen.
Cortex, medulla, and soralia K + bright
yellow, C - or + pink, KC+ red, PD + deep
orange to reddish. Contains barbatolic acid and
small amounts of alectorialic acid, fumarpro-
tocetraric acid, chloroatranorin, and traces
of atranorin (in most specimens).
Ecology. On Abies balsamea, Picea spp.,
Larix laricina, Betula papyrifera, and occa-
sionally other trees in the southern boreal
spruce-fir forest; also frequently on vertical
rock faces and cliffs, especially in ravines or
along lake shores, and on rock in subarctic
and subalpine sites; on A Intts rubra in British
Columbia (Brodo 13881 [CANL]).
Bryoria nadvornikiana can be found in great
abundance in fairly closed, almost pure stands
of Abies balsamea in the southern boreal
zone north of Quebec City, Quebec. Its tol-
erance of the shade and its absence in more
open habitats are very striking.
Distribution. Mainly in the strongly humid
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 123
Fig. 58. N. American distribution. - A: • Bryoria nadvornikiana. OB. pseudocapillaris. - B: B. pikei.
southern boreal forest zone, but with scattered
occurrences in the northern boreal zones of
Lingava (Quebec-Labrador) and Keewatin
(N.W.T.), the southern Appalachians, the
Rocky Mountains, the Gulf Islands near
Vancouver (B.C.), and southeastern Alaska
(Fig. 58 A). The same kind of clumping noted
' n the distribution pattern of B. capillaris
ls seen here, again, mainly corresponding to
areas of high precipitation. It is of scattered
distribution in northern and middle Europe
(Poelt 1969). Ahlner (1948), interpreting the
Fennoscandian distribution of B. nadvornikiana
(sub Alectoria altaica ), suggested that the
species survived the Pleistocene glaciation as a
saxicolous plant, reinvading coniferous sub-
strates with the reappearance of the forests.
The tendency of this species in North America
124 Alectoria in North America
OPERA BOT. 42 (1977)
to be saxicolous in cool ravines and exposed
headlands in the southern boreal forest, and to
appear on boulders and rock faces again
in the most northern subarctic localities would
support this hypothesis.
In addition to its European and North
American localities, this species has also been
reported from the Hawaiian Islands (Magnusson
1955), the East African mountains (Krog &
Swinscow 1975), Himalaya (Bystrek 1969), and
Japan (Yoshimura 1974).
Discussion
Bryoria nadvornikiana, although easy to identify
by virtue of its chemistry and soralia, is
nevertheless quite variable in appearance. Speci-
mens which are basically caespitose can produce
virtually pendent branches with occasional,
spinulose laterals. Some specimens are almost
entirely pendent except for the base, and can
only be distinguished from B. capillaris by the
short, pointed, lateral branches mentioned
above, and the presence of soredia (in the
North American material). B. capillaris also
lacks atranorin, chloroatranorin, and fumar-
protocetraric acid in North America and has
a quite different branching pattern.
Chloroatranorin was present in all specimens
of B. nadvornikiana tested by us (including
some from Europe), and while fumarprotoce-
traric acid may also be a constant together
with alectorialic and barbatolic acids, it some-
times occurs in such low concentrations as to
be almost undetectable byTLC.
The soredial production is also variable.
Some specimens are covered with abundant
fissurai or tuberculate soralia or occasionally
both, but some are virtually esorediate, and a
few are entirely without soralia. Colour varies
from almost white to almost black, although
most specimens are very pale fuscous to grey-
brown except for the blackened basal region
which may be extensive.
One group of specimens shares enough
characters to warrant some sort of taxonomic
recognition, but we are refraining from giving
it a name here in the absence of a thorough
study of the northern and eastern European
material. These specimens are generally from the
northern limits of the range and almost always
are saxicolous. They are characterized by
being extremely dark in colour (dark brown
to almost black), growing in dense, caespitose
clumps, with abundant spinulose branches as
in B. furcellata or B. nitidula, and having few
soralia with those present being all fissurai.
Although a few of these specimens apparently
lack fumarprotocetraric acid and/or atranorin
(s. lat .), for the most part they are chemically
indistinguishable from B. nadvornikiana s. str.
In addition, one can find morphological inter-
mediates with more typical B. nadvornikiana.
Thus we are tentatively treating these morpho-
types as conspecific.
Bryoria pseudocapillaris, found along the
Oregon and California coasts, is also very similar
in some respects to certain morphotypes of
B. nadvornikiana.
There remains some question as to which
section of the genus Bryoria is most appropriate
for B. nadvornikiana. Bystrek (1971) placed
the species in his subsection Divaricatae (of
the section Fuscidulae), allied to other Divari-
catae such as B. asiatica. Indeed, B. nadvorni-
kiana often closely resembles members of the
section Divaricatae, especially in the more
northern morphotypes described above. How-
ever, its chemistry is clearly more closely
allied to that of species in the Implexae,
especially B. capillaris, in which the sorediate
morphotypes are virtually identical chemically
to B. nadvornikiana. Other characteristics such
as colour, soralium type, and cortex are much
more characteristic of species in the Implexae
than the Divaricatae.
Selected specimens (*dark, caespitose
morphotype on rock )
Exsiccatae. Brodo: Lieh. Can. Exs. 81: Quebec,
Montmorency County: Parc des Laurentides, Brodo
20595 (CANL).
Canada. British Columbia. Islands South District:
Saltspring Island, Brodo 13881 (CANL) - Saskatche-
wan. *W. Dodge Lake, Scotter 259 (WIS) -Ontario.
Algoma District: 30 miles NW of Wawa, Brodo
6850 (CANL) - Thunder Bay District: St. Ignace
L, Garton 6251 (US) - Quebec. Cté Charlevoix:
Parc des Laurentides, Gauthier 2216B (CANL) -
Cté Frontenac: Mont Mégantic, Masson 6067 (WIS)-
Territoire de Mistassini: Otish Mts., Mont du
Lagopède, Shchepanek 71-L-59 (p.p.) (CANL) -
Territoire de la Baie de James: 54°00'N, 69°56'W,
Gérardin QFB-E 4401 (CANL) - Territoire du
Nouveau Québec: Fort-Rupert, (collector unknown)
12059 (CANL) - Nova Scotia. Halifax County:
OPERA BOT. 42 (1977)
B ry or ici Brodo & D. Hawksw. 125
Prospect Road, Brown s.n., 4 Aug. 1949 (NSPM) -
New Brunswick. Albert County: Fundy National
Park, Ireland 11332 (p.p.) (CANL) - Newfoundland.
Placentia West District: Boat Harbor, Ahti 565 (H) -
Labrador West District: Menihek Lake, Brodo
8837A-5B (CANL) - Central Labrador: Lake St.
John, Lake Ducharme, Hustich s.n., 7 July 1948 (H) -
Northwest Territories. District of Mackenzie: *North
Talson R., Scotter 2333 (W1S) - Gordon Lake
Area, Scotter 1552 (W1S) - *Thekulthili Lake,
Scotter 2471 (W1S).
U.S.A. Alaska. Eastern Pacific Coast District:
Juneau, Loop Road, Krog 5522 (O) - Maine.
Knox County: Camden, Mt. Battie, Merrill s.n.,
Aug. 1904 (FH) - Michigan. Keweenaw County:
W of Agate Harbour, Imshaug 4955 (MSC) - Min-
nesota. Cook County: Susie Islands. J. W. Thomson
3935 (COLO, DUKE, US, DEGELIUS) - Montana.
Glacier County: Glacier National Park. Snyder Creek,
Imshaug 7309 (S) - New Hampshire. Coos County:
Mt. Washington, Burkholder 38A (p.p.) (US) -
North Carolina. Avery County: Roane Mt., Hale
18043 (US) - Swain County: Mt. Kephart, Great
Smoky Mts., Degelius s.n., 15 Sept. 1939 (US,
DEGELIUS) - Yancey County: Mt. Mitchell,
Becking 5907139 (BECKING) - Tennessee. Blount
County: Mt. Le Conte, Great Smoky Mts.. Degelius
s.n., 14 Sept. 1939 (DEGELIUS).
19. Bryoria pikei Brodo & D. Hawksw.,
S P- nov.
Thallus pendulus, 10-15 cm longus; rami iso-di-
chotomiter ramosi, in angulos acutos et obtusos
divergentes, aequales vel leviter inaequales diametro,
ton tortuosi, basi 0.2-0.35 mm diam.; planta
olivacea vel parum olivaceo-brunnea sed basi
Pallidior.
Rami laterales spiniformes veri, isidia et soralia
desunt. Pseudocyphellae copiosae, albae vel
hrunneae, fusiformes vel late fusiformes, saepe in
s Piram circa ramos contortae.
Apothecia et pycnidia ignota.
Cortex K- vel leviter flavescens, C+ roseus,
BD+ flavescens; medulla K-, C-, KC-, PD-.
Thallus acidum alectorialicum et interdum minime
acidum gyrophoricum vel minime acidum barbato-
hcum continens.
Holotypus: America septentrionalis. U.S.A. ,
Dregon, Marion County, Silver Creek Falls State
fhtrk, prope Winter Falls, orientent versus a Salem,
I22°39'W, 44°53'N , alt. 1,450 ft, in Pseudotsuga
p* enziesii juvenili, corticola, L. Pike 2475, 9 April
19 ?2 (CANL 38271).
A duplicate (arithmotype) of Pike 2475 at OSC was
*°und to be entirely B. pseudofuscescens. The
holotype packet is a mixture of B. pikei and B.
Ps eudofuscescens.
Phallus pendent, 10-15 cm long; branching
ls otomic dichotomous, angles between
dichotomies acute and obtuse, branches even to
lightly uneven in diameter, not twisted, 0.2-
0.35 mm diam. at the base; uniformly olivaceous
or occasionally becoming olivaceous brown
and somewhat paler at the base.
True lateral spinules, isidia, and soralia
absent. Pseudocyphellae abundant, white to
brown, fusiform to broadly fusiform, often
spirally twisting around the branches.
Apothecia and pycnidia unknown.
Cortex K- or + weakly yellow, C+ pink,
KC+ pink, PD+ yellow; medulla K-, C-,
KC-, PD-. Contains alectorialic acid and
sometimes also traces of gyrophoric acid or
traces of barbatolic acid.
Ecology. On conifer branches and lignum,
usually in humid forests close to the coast or a
lake, at 0-1250 m elevation.
Distribution. West coast, from the Queen
Charlotte Islands to Oregon, with a single
disjunct locality on Cape Breton Island, Nova
Scotia (Fig. 58 B). The comments for B.
friabilis also apply to this North American
endemic.
Discussion
Bryoria pikei seems to be most closely related
to B. capillaris but can usually be distinguished
from that species by its dark olivaceous colour;
the shiny, smooth, and very even surface of its
branches; the production of very distinct, long,
white, fissurai pseudocyphellae; and by its
characteristic lack of barbatolic acid. We have
seen specimens which are olivaceous only in
part, with most of the thallus a variegated
grey-brown. Some of these specimens also
produce trace amounts of barbatolic acid.
However, even dark specimens of B. capillaris
(which are not rare) are usually quite dull
and uneven, and tend to be smoky brown or
brownish grey rather than olivaceous although
the conspicuous white pseudocyphellae in
these thalli are often quite similar of those of
B. pikei. Despite the convergence of some of
these characters, we believe that the two taxa
are distinct. We have seen several mixed packets
of B. capillaris and B. pikei in which it was
easy to distinguish the two morphologically.
Bryoria pikei also has a much more limited
distribution.
This species is also closely related to B.
friabilis, with some specimens being almost
126 Alectoria in North America
OPERA BOT. 42 (1977)
intermediate in colour; B. friahiiis is generally
pale buff. The presence of trace amounts of
gyrophoric acid in the holotype of B. pikei
and the production of alectorialic acid in the
apothecia of B. friabilis confirm their close
relationship. B. pikei, however, is not nearly
as friable, has a characteristic olive colour,
and has alectorialic rather than gyrophoric acid
as its principal constituent. Gyrophoric acid
could not be detected at all in several specimens
of B. pikei.
It should be noted that there is a distinct
possibility that in our study some old and
blanched specimens of B. pikei have been
misidentified as B. capillaris. Almost all the
material of B. pikei traced has been collected
recently, and we do not know how the char-
acteristic olivaceous colour of the thallus stands
up to ageing in herbarium packets. A number
of specimens listed under B. capillaris here,
which were examined early in our study,
were rather shiny and therefore might con-
ceivably represent B. pikei. If this proves to
be the case B. pikei is a more common species
than we have suggested.
Specimens
Canada. British Columbia. Islands North District:
Graham Island: Ian Lake, Brodo 18435B (CANL) -
Kumdis Bay, Port Clements, Brodo 18144A (p.p.)
(CANL) - Moresby Island: Between Sandspit and
Copper Bay, Brodo I2874A (CANL) - Skidegate
Lake, Brodo I8513B (p.p.) (CANL) - Wathus Island:
Masset Inlet, Brodo I8314A (CANL) - Islands South
District: Saltspring Island. Brodo 13839 (CANL) -
Nova Scotia. Victoria County: Cape Breton High-
lands National Park, Speer 73-24 (CANL).
U.S.A. Oregon. Benton County: Corvallis, Mary's
Peak, Brodo 20954 (CANL) - Linn County: Mount
Jefferson National Wilderness, Hank Lake, Pike
1110 (CANL) - Marion County: Silver Creek Falls
State Park E of Salem, Pike & Rossman 2475 (L.P.)
(CANL, OSC) - Washington. Clallam County:
Hurricane Hill, Pike 844 (CANL) - Pierce County:
Mt. Rainier National Park near Paradise Valley,
Brodo 15438B (p.p.) (CANL).
20. Bryoria pseudocapillaris Brodo &
D. Hawksw., sp. nov.
Thallus subpendulus, 5-7 cm longus, parum rigidus;
rami plerumque iso-dichotomiter fastigiati, sed ramulis
copiosis brevibus acutis vel perpendicularibus
instructi; rami principales plerumque omnino
teretes, non compressi vel tortuosi, diametro
aequales, laeves, 0.25-0.33 mm diam.; planta
pallidissime brunnea vel badia, opaca.
Rami laterales spiniformes veri, isidia et soralia
desunt. Pseudocyphellae longae, lineares, albae,
depressae, plerumque rectae sed interdum circa
ramos leviter fortae, 1. 2-3.0 mm longae.
Apothecia et pycnidia ignota.
Cortex K+ flavus, C + roseus, KC+ roseus,
PD + aureus; medulla K — , C - , KC — , PD -. Thallus
acidum alectorialicum, acidum barbatolicum et sub-
stantiam ignotum continens.
Holotypus: America septentrionalis, U.S.A. ,
Oregon, Curry County, Cape Blanco, 8 mis boreo-
orientem versus a Port Orford, in arboribus sejunctis
summi ventosissimi, I. M. Brodo 20539, 28 June
1974 (CANL 50596). Isotypus: BM. Fig. 59.
Thallus subpendent, 5-7 cm long, rather stiff;
branching mainly isotomic dichotomous but with
frequent acute to perpendicular short side
branches; main branches mostly terete through-
out, not flattened or twisted, even, smooth,
0.25-0.33 mm diam.; very pale brown to
badious, matt.
True lateral spinules, isidia and soralia absent.
Pseudocyphellae long, linear, white, depressed,
usually straight but sometimes slightly twisted
around the branches, 1. 2-3.0 mm long.
Apothecia and pycnidia unknown.
Cortex K+ yellow, C+ pink, KC+ pink,
PD+ deep yellow; medulla K-, C-, KC-,
PD— . Contains alectorialic and barbatolic acids
together with an unidentified substance.
Ecology. Coastal trees and rocks in exposed
situations.
Distribution. Rare, from coastal northern Cali-
fornia to Oregon (Fig. 58 A).
Discussion
This species is being placed in the genus
Bryoria due to its resemblance to certain
forms of B. capillaris and B. kuemmerleana,
although its pseudocyphellae are so striking
that we cannot entirely exclude the possibility
that it should be included in the genus Sulcaria.
Its branching type and vaguely badious colour
suggest a close relationship with B. spiralifera,
which shares the same habitat.
Chemically, it resembles B. nadvornikiana
and B. capillaris, but differs from the former in
lacking soralia and chloroatranorin, and from the
latter in being stiff and having perpendicular
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 127
Fig. 59. Bryoria pseudocapillaris. Holo-
( ype. Scale 0. 5-1.0 mm.
branches. It differs from both in the unusually
long pseudocyphellae.
Specimens
U-S.A. California . Humbolt County: Manila, Brodo
2051 IB (CANL) - Oregon. Curry County: Cape
Blanco, near Port Orford, Imshaug 17660 (MSC),
Brodo 20539 (CANL).
21. Bryoria pseudofusceseens (Gyeln.) Brodo &
D. Hawksw., comb. nov.
4 leetoria pseudofusceseens Gyeln., Ann. Mus. Nat.
Hung., Bot. 28: 283 (1934) et Revue Bryol. Lichén.
51 (1934); type: “Oregon, Corvallis, on old apple
trees, F. P. Sipe" (BP 33.958 holotype!); basionym.
Alectoria norstictica Mot., Bryologist 67: 33 (1964);
type: Alaska, East Pacific Coast District, Juneau,
Mindenhall Glacier Trail, H. Krog 5507, 27 August
1^57 (O lectotype!): nom. inval. (Arts. 36, 37). -
jpO ’opogon norsticticus (Mot.) Bystr., Ann. Univ.
Mariae Curie-Sktodowska. C, 26: 274 (1971); nom.
‘nval. (Arts. 33, 36, 37).
Alectoria subtilis Mot., Bryologist 67: 32 (1964);
type: Alaska. Central Pacific Coast District, Kenai
Peninsula, H. Krog 2164, 7 June 1957 (O lectotype!);
nom. inval. (Arts. 36, 37). - Bryopogon subtilis
(Mot.) Bystr., Ann. Univ. Mariae Curie-Sklodowska,
C, 26: 274 (1971); nom. inval. (Arts. 33, 36, 37).
For further synonyms see Hawksworth (1972 p.
Thallus pendent, usually 5— 10(— 1 5) cm long;
branching isotomic dichotomous at least at the
base but sometimes becoming submonopodial
towards the apices, rarely with sparse, short,
perpendicular, lateral branches, angles between
the main dichotomies variable, branches even
to uneven in diameter, straight or twisted,
mainly 0. 1 5— 0.3(— 0.35) mm diam., basal
branches usually 0.20-0.35 mm diam.; pale
brown to greyish in some specimens, darkening
to dark brown or black, often olivaceous
especially towards the base, dull or shiny.
Soralia and true lateral spinules absent.
Pseudocyphellae usually abundant, white, short
to elongate fusiform, depressed, often forming
partial spirals around the filaments, often to
0.5-1 .0 mm long.
Apothecia very rare, lateral, c. 0.4-0. 5 mm
diam., excipulum thallinum concolourous with
the thallus, incurved, generally excluded at
maturity; disc reddish brown, concave at first,
sometimes becoming convex. Hymenium 36-40
gm high. Ascospores 8 per ascus, hyaline,
nonseptate, broadly ellipsoid to subspherical,
5. 4-7. 5 x 4. 0^1. 5 gm.
Pycnidia unknown.
Cortex and medulla K + yellow changing
128 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 60. A: Bryoria pseudofuscescens. N. American distribution. A determination uncertain. - B: B. salazinica.
Known distribution.
to red, or faintly yellow, C— , PD + yellow.
Contains norstictic acid, and rarely, traces of
connorstictic acid.
Ecology. On coniferous trees, especially Picea
glauca, Abies lasiocarpa, and Pinus aibicaulis
in montane conifer forests, mainly at 1100-2350
m. In some subalpine forests, B. pseudo-
fuscescens drapes well-exposed fir trees from
top to bottom. The holotype, however, is from
apple trees close to sea level. One specimen
was seen from rock (California, Santa Cruz
Co., Becking 6607721 [BECKING]).
Distribution. West coast from southern Alaska
to California, eastward into the northern Rocky
Mountains and Douglas fir forests of northern
Idaho and adjacent Washington and Montana,
but avoiding the Columbia plateau (Fig. 60 A).
Several of the Oregon localities, including that
of the holotype, are in the more lowland oak
forest region which, however, also contains
Pseudotsuga as a dominant tree species
(Kiichler 1964) (see below). The specimen from
the Keewatin District west of Hudson Bay
was reexamined and apparently is correct. Its
occurrence there is extremely puzzling.
The taxonomy of this species with its many
forms is still unsettled. It certainly occurs in
the boreal forests of Europe and has been
reported from a few localities in Scotland
(Hawksworth 1972, sub “Alectoria trichodes”).
Discussion
Bryoria pseudofuscescens appears to be most
closely allied to B. implexa from which it
differs primarily in having norstictic rather
than psoromic acid. (See also the discussion
under B. implexa.) The two species sometimes
occur mixed together in Alaska (H. Krog pers.
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 129
comm.). B. pseudofuscescens is basically a
slender, extremely dark plant with uneven
filaments and scattered, dark or white pseudo-
cyphellae. Like B. implexa, B. pseudofuscescens
is esorediate in North America.
The morphology of B. pseudofuscescens is
somewhat variable, and marked divergences
from the above traits are sometimes seen.
The type specimen of "Alectoria subtilis”
is unusually pale in colour and has rather few
pseudocyphellae, for example. A particularly
distinct morphotype with extremely thick,
twisted and foveolate main branches, super-
ficially not unlike B. vrangiana except that it
lacks soralia and has norstictic instead of
fumarprotocetraric acid, has been encountered
several times (Sansom, 1913 [FH]; Otto 3131b
[UBC]). At first we were inclined to regard
this morphotype as an infraspecific taxon of
B. vrangiana but as the main branches of
generally very slender specimens can become
somewhat thickened and contorted, it seems that
these specimens merely represent environmental
modifications of B. pseudofuscescens.
Norstictic acid is only known otherwise
amongst the North American Bryoriae in B.
spiralifera, which is easily distinguished by its
colour, elongate pseudocyphellae, and branch-
ing pattern. In addition, B. spiralifera always
contains abundant connorstictic acid. A peculiar
specimen of B. pseudofuscescens from Washing-
ton contains norstictic as well as barbatolic
acid and is discussed under/?, capillaris.
In Europe there may well be several norstictic
acid-containing species in addition to B. pseudo-
fuscescens, but they remain poorly known;
for this reason Hawksworth (1972) treated
them as a species aggregate (sub Alectoria
trichodes (Michx.) D. Hawksw., a name which
must now be used in a different sense; see
Discussion under Bryoria trichodes subsp.
trichodes). The pale, often variegated, grey-
brown robust species of central and eastern
Europe, B. kuemmerleana (Gyeln.) Brodo &
D. Hawksw., is almost certainly worthy of
this rank; we have found that at least some
specimens (all which have been tested so far)
of this species contain atranorin and connor-
stictic acid in addition to norstictic acid. No
specimens referrable to B. kuemmerleana have
been seen by us from North America.
The amount of norstictic acid in this species
9 - Opera Botanica nr 42
is rather variable, and for this reason, a positive
K reaction is sometimes difficult to obtain
(see p. 23). It may be that the apparent lack
of connorstictic acid in all but two or three
specimens is tied to the low concentrations
of norstictic acid.
A certain amount of confusion surrounds the
type material of B. pseudofuscescens. To begin
with, an “isotype” in ORE (i.e., bearing the
same number as that designated by Gyelnik
as the holotype) is entirely different from the
holotype in morphology and chemistry, and,
in fact, represents a new species which we
are calling Sulcaria badia. In addition, although
the “isotype" packet is marked “No. 669,
on old apple trees”, it is labelled as being
from “Philomath", while in the original
description, “Corvallis” is given. This is not as
serious as it might first appear since Philomath
is a small village about seven miles west of
Corvallis. It is understandable that Dr. Sipe
would mark a specimen sent to a foreign
colleague with the name of the closest city
likely to be found on an ordinary map. Of
course, it would have been easy to have
collected a mixture of two species and sent
only one of them away for determination.
More disturbing, however, is the fact that
the Philomath-Corvallis area and most of the
surrounding valley are quite unlike the “typical”
habitat of B. pseudofuscescens as we have
come to understand it, i.e., mountainous
coniferous forests. The trees in the valley are
mainly deciduous, occurring in open stands of
Quercus garryana with occasional individuals or
even small stands of Pseudotsuga menziesii.
The area is dotted with old and recent apple
orchards. One of us (I.M.B.) searched for both
B. pseudofuscescens and Sulcaria badia on
five separate occasions in old apple orchards,
oak stands, Pseudotsuga forests, and pine
stands, both in open valleys and on hills,
without finding either of the lichens. There are
some localities in the general area which may
be sufficiently mountain-like to serve as a site
for B. pseudofuscescens, for example Mary's
Peak, several miles to the west of Philomath,
but a short visit to that locality proved fruitless.
It seems impossible to definitely determine
whether (a) the localities marked on the isotype
and given in the original description were
incorrect (i.e., mislabelled in some way), (b)
130 Alectoria in North America
OPERA BOT. 42 (1977)
Gyelnik mislabelled the holotype, confusing it
with a completely different collection, or (c)
the two species in question are simply too
rare in the Corvallis area to be found easily.
In the latter connection, since we have seen
several specimens conforming to our concept of
B. pseudofuscescens collected in the oak-
Douglas fir forest region of western Oregon, it
would seem that even the unlikely locality
can be accepted as probable.
As Gyelnik's descriptions are so brief and
often unreliable, they help little in the eluci-
dation of this problem. However, the epithet
pseudofuscescens would have been an ex-
tremely inappropriate choice for Sulcaria badia.
Even if the specimen in the “typus” packet
in BP were not collected by Sipe at all but
came from another locality, it is not nomen-
claturally important since there can be no
doubt that Gyelnik regarded that packet as the
nomenclatural type. That Gyelnik intended his
name to refer to the species treated here
under this epithet is also supported by his
application of the name to other collections
(e.g., Lichenotheca no. 16 [BM]). If the holo-
type actually proved to be something other than
Sipe’s collection, the stated locality in the
original description could simply be treated
as a lapsus.
Selected specimens
Exsiccatae. Merrill: Lieh. Exs., Ser. II, 129 (mixed):
Alaska, Fairbanks, Palmer (MSC, NYBG, US, WIS).
Canada. British Columbia. Coastal District: E of
Pemberton, near Lillooet Lake, Brodo 8313D (CANL)
- Columbia River Basin: Big Bend District, Shaw
977 (NYBG, BM)-Dean River Basin: Mt. Walker,
40 miles E of Bella Coola, Ohlsson 2259 (CANL) -
Islands South District: Vancouver Island: “on Lea’s
Farm”, (Macoun?), Herb. G. K. Merrill, 17 June
1908 (FH) - Kootenay River Basin: Glacier National
Park, Glacier Crest Trail, Ohlsson 395 (MSC, CANL)
- Okanagan District: Princeton, Brodo 7811 (CANL)
- Skagit River Basin: Manning Park, Blackwall
Peak, Brodo 20565 (CANL) - Skeena River Basin:
South Hazelton, Ohlsson 3020 (MSC, CANL) -
Alberta. Banff National Park, trail from Lake
Louise to Lake Agnes, Weber S23, 544 (CANL,
NYBG) - Northwest Territories. Keewatin District:
Dubawnt Lake, J. W. Thomson 14299 (p.p.) (WIS?).
U.S.A. Alaska. Alaska Range District: Cantwell,
Krog s.n., 8 June 1969 (O) - Central Pacific Coast
District: Anchorage Area, Krog 1420 (O) - Eastern
Pacific Coast District: Juneau, Mindenhall Glacier
Trail. Krog 5507 (O) - Western Pacific Coast
District: Dillingham Quadrangle, Wood River Lakes,
Viereck 8964 (p.p.) (CANL, ALA) - California.
Del Norte County: Near Bear Basin, Tucker 6434
(COLO) - S of Takilma, Oregon, Pike L-325A
(CANL) - Santa Cruz County: Butano State Park.
Becking 6607721 (BECKING) - Idaho. Bonner
County: Priest River Experimental Forest, Esslinger
1580B (WIS) - Nez Perces County: Lake Waha,
Heller 3364 (MSC) - Montana. Mineral County:
Donna Morra campsite, J. W. Thomson 16300
(CANL) - Ravalli County: Bass Creek, Stevensville,
Stickney 1581 (WIS) - Oregon. Benton County:
Corvallis, Sipe s.n., no date (BP) - Deschutes
County: Doris Lake in the Deschutes National
Forest, Pike L-392 (PIKE) - Klamath County:
Crater Lake, Kraus s.n., Aug. 1921 (WIS) - Lane
County: Eugene, Doty 3099 (p.p.) (NYBG) - Union
County: Blue Mountain, Summit Pass (US 30),
Imshaug 21 (MSC) - Washington. Pierce County:
Mt. Rainier National Park, trail to Glacier Basin,
Imshaug 315 (F, US) - Wyoming. Teton County:
Cascade Canyon, Grand Teton National Park,
Alexander s.n., 29 Aug. 1955 (COLO).
22. Bryoria salazinica Brodo &
D. Hawksw., sp. nov.
Thallus pendulus vel subpendulus, plerumque
crispatus et implexus, 5-9 cm longus; rami praeter
basim praecipue aniso-dichotomiter fastigiati, in
angulos obtusos vel perpendiculares valde diver-
gentes, diametro aequales vel inaequales, recti vel
leviter torti, in axillis saepe foveolati et complanati,
0.16-0.35 mm diam.; tota planta obscure olivaceo-
brunnea vel pallide brunnea, opaca vel bombycina.
Rami laterales spiniformes veri, isidia et soralia
desunt. Pseudocypheiiae plerumque copiosae, im-
polite albae, fusiformes vel elongato-fusiformes,
0.08-0.4(-0.65) mm longae.
Apothecia et pycnidia ignota.
Cortex (et forsan medulla) K+ flavescens, C-,
KC+ flavescens. PD+ aurantiaco-flavescens. Thallus
acidum salazinicum continens.
Holotypus: America septentrionalis, U.S.A. ,
Massachusetts, Springfield, W. G. Farlow, May 1878
(NYBG). Fig. 61.
Thallus pendent to subpendent, often curled
and entangled, 5-9 cm long: branching mainly
anisotomic dichotomous except at the base,
branches broadly divergent at obtuse to per-
pendicular angles, even to uneven, straight to
somewhat twisted, often foveolate and com-
pressed at the axils, 0.16-0.35 mm diam.;
dusky olivaceous brown to pale brown through-
out, dull to satiny.
True lateral spinules, isidia, and soralia
absent. Pseudocypheiiae usually abundant, dull
white, fusiform to elongate-fusiform, 0.08-0.4
(-0.65) mm long.
Apothecia and pycnidia unknown.
Cortex (and perhaps medulla) K + yellow.
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 131
C-, KC+ yellow, PD + orange-yellow. Con-
tains salazinic acid.
Ecology. On coniferous trees along the coast
or on the coastal plain.
Distribution. Northeastern coastal plain; North
American endemic (Fig. 60 B).
Discussion
This species appears to be closely related to
the B. implexa - B. friabilis - B. pikei group,
all of which have long twisting pseudocyphellae,
an olivaceous caste, and a similar uneven
branching. B. pseudofuscescens is also very
similar in many respects. We considered the
possibility that B. salazinica might merely
represent a chemical race of one of these
four species but its remarkable chemistry and
well-defined and restricted distribution
prompted us to regard it as a species.
Alectoria angolensis Dodge, described from
Africa, is reported as having salazinic acid,
but that taxon is sorediate and, to judge from
an authentic specimen examined by Hawks-
worth (1971 a), is almost certainly a species
of Ramalina on the basis of its cortical
structure. (Microtome sections are preserved in
IMl.)The only other previous report of salazinic
acid in Alectoria s. lat. is Solberg's (1956)
mention of it in European material named as
"A. cana (Ach.) Leight.” (a synonym of
B. capillaris ); this report should be verified
since no other workers have found this com-
pound in any European Alectorioid species.
Specimens
Canada. Nova Scotia. Halifax County: Ketch
Harbour, Taschereau B-14 (NSPM) - Prince Edward
Eland. Prince County: Richmond, Ireland 10298
(CANL) - Queens County: West Barkley Beach,
H miles NW of Charlottetown, Fabiszewski s.n.,
20 July 1970 (CANL) - Newfoundland. Trinity
North District: Bay Bull's Ann, Trinity Bay, collector
unknown, April 1897 (M).
U.S.A. Maine. Washington County: Eastport. Herb.
"L G. Farlow s.n., no date (FH) - Jonesport, Cheever
1314 (US) - Massachusetts. Hampden County:
Springfield, Farlow s.n., May 1898 (NYBG).
9 *
Fig. 61. Bryoria salazinica. Holotype. Scale 0. 5-1.0
mm.
23. Bryoria spiralifera Brodo & D. Hawksw.,
sp. nov.
Thallus pendulus, 6-7 cm longus; rami basi iso-
dichotomiter sed in ramis principalibus praecipue
aniso-dichotomiter fastigiati; rami principales 0.2-
0.25 mm diam.. diametro inaequales, recti vel
tortuosi, ramulis numerosis brevibus, gracilibus
perpendiculariter instruct!; planta atrobadia vel pallide
badia, opaca, ramis lateralibus saepe dilutioribus.
Rami laterales spiniformes, isidia et soralia desunt.
Pseudocyphellae copiosae, conspicuae, albae,
lineatae, in spiras longas circa ramos contortae vel
partim rectae. ad 4 mm longae.
Apothecia et pycnidia ignota.
Cortex K+ rubescens, C— , KC— , PD+ flavescens;
medulla K-, C-, KC-, PD-, Thallus abundanter
acidum norsticticum sed minime acidum connorsticti-
cum et atranorinum (non chloroatranorinum) con-
tinens.
Holotypus: America septentrionalis, U.S.A., Cali-
fornia. Humboldt County, in pineto prope Manila, in
Pino contorta, S. Dowty 137, 22 January 1972
(CANL 38403). Fig. 62.
Thallus pendent, 6-7 cm long; branching isoto-
mic dichotomous at the base but becoming
mainly anisotomic dichotomous on the main
branches; main branches 0.2-0.25 mm diam.,
uneven, straight to twisted, with numerous,
short, slender, perpendicular branches; dark to
pale badious, matt, lateral branches paler.
True lateral spinules, isidia, and soralia
absent. Pseudocyphellae abundant, con-
spicuous, white, linear, twisted in long spirals
around the branches or straight in parts, to 4
mm long (Fig. 5 E).
-Opera Botanica nr 42
132 Alectoria in North America
OPERA BOT. 42 (1977)
V
Fig. 62. Bryoria spiralifera. Holotype. Scale 0. 5-1.0
mm.
Apothecia and pycnidia unknown.
Cortex K+ red, C— , K.C+ red, PD+ yellow;
medulla K— , C-, KC— , PD — . Contains large
amounts of norstictic acid together with smaller
quantities of connorstictic acid and atranorin
(not chloroatranorin).
Ecology. On Pinus conforta, Picea sitchensis,
and shrubs in scrub forests near the coast; found
in shaded or sunny habitats generally mixed with
Ramalina menziesii.
Distribution. Known only from the Samoa
peninsula in Humboldt County, California,
where it is very scattered, but not uncommon
(Fig. 39).
Discussion
The badious colour, and extremely long pseudo-
cyphellae which sometimes become somewhat
furrow-like, at first suggested that this species
might belong to the genus Sulcaria. We now feel
it more properly belongs in the genus Bryoria,
closely allied to B. pseudocapillar is (see Dis-
cussion under this species). In some respects
this species resembles the European B. kuem-
merleana (see Discussion under B. pseudofus-
cescens) but that species is generally much
larger and has a different branching type.
Specimens
U.S.A. California. Humboldt County: Near Manila,
Dowty 137 (HSC, CANL) - Manila (W of Areata),
Brodo 20501, 20509, 20511 A (CANL) - Samoa
Peninsula, Lindsay 782, 926 (W1S) - Locality un-
known: J. Tallant s.n., 1856 (US).
Bryoria sect. Subdivergentes (Mot.) Brodo &
D. Hawksw., comb. nov.
Alectoria subgen. Bryopogon sect. Subdivergentes
Mot., Bryologist 67: 36 (1964); basionym. -Bryopogon
sect. Subdivergentes (Mot.) Bystr., Ann. Univ. Mariae
Curie-Sktodowska, C, 26: 275 (1971), as "Mot.”
Type species: Bryoria subdivergens (Dahl) Brodo &
D. Hawksw. (syn. Alectoria subdivergens Dahl;
holotype).
Thallus erect and caespitose to decumbent or
pendent, reddish brown, matt, not becoming
blackened. Lateral spinulose branches often
present, rarely abundant, not constricted at the
base. Isidia, soralia, and pseudocyphellae
absent. Cortex distinctly cellular, knobby at the
surface. Apothecia common in most species;
margins ciliate; discs reddish brown, sometimes
markedly convex. No lichen products known.
Discussion
This section comprises four species of which
only two are recorded in North America with
certainty; both of these are endemic here ( B .
abbreviata and B. oregana). The other two
species of this section are B. subdivergens
(only reliably recorded from southwest Green-
land; see Excluded taxa) and B. divergescens
(a southeast Asian species). The section is well
characterized by the colour of the thallus, the
structure of the cortex, the ciliate apothecia,
and the absence of lichen products. It appears to
occupy a rather isolated position within Bryoria.
24. Bryoria abbreviata (Müll. Arg.) Brodo &
D. Hawksw., comb. nov.
Alectoria divergens f. abbreviata Müll. Arg., Flora,
Jena 72: 362 (1889); type: Oregon, Mt. Hood, Boell
111, 1889 (G holotype!); basionym. - Alectoria abb-
reviata (Müll. Arg.) R. Howe, Class. Fam. Usnea-
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 133
ceae: 22 (1912). - Bryopogon abbreviatus (Müll.
Arg.)Gyeln., Feddes Repert. 38: 235 (1935).
Thallus caespitose, up to 2.5 cm in length;
branching anisotomic dichotomous, broadly
divergent at the base, angles between dichoto-
mies acute or obtuse, branches uneven in dia-
meter, 0. 3-1.0 mm diam. at base, 0. 2-0.4 mm
diam. on main stem, foveolate and channeled;
generally very angular or flattened; occasionally
anastomosing, red-brown, matt. Fig. 63.
True lateral spinules, soralia, and pseudocy-
phellae absent; branches sometimes appearing
spinulose due to short pointed lateral branches.
Apothecia common, subterminal or rarely
lateral; excipulum thallinum concolourous with
thallus, persistent and ciliate to disappearing,
flat to recurved, red-brown darkening to almost
black with age, 2.0 — 3 .0( — 3 .5) mm diam. Spores 8
per ascus, broadly ellipsoid, c. 6. 5x4. 5 /Am.
Pycnidia not seen.
Cortical hyphae as in B. oregana (Fig. 12
A-E).
Cortex and medulla K-, KC — , C— , PD — ;
lichen substances absent.
Ecology. On coniferous trees, mainly Pinus pon-
derosa and Pseudotsuga menziesii in the rather
dry Interior Douglas Fir forests, c. 700-1400 m
in elevation.
Distribution. North American endemic of
western Douglas fir forests from southern British
Columbia and Alberta to Baja California (Fig.
64 A). It has the most southerly continuous
distribution among the North American Alec-
toriae s. lat. and clearly avoids the more humid
coasts of Oregon, Washington, and British
Columbia, showing a preference for dry forest
regions.
Discussion
This species has normally been included in B.
oregana but appears to be distinct from it. The
separation of these two species is discussed
under B. oregana.
Bryoria abbreviata shows a strong super-
ficial resemblance to Cornicularia californien
(Tuck.) DR. and is often confused with it in
herbaria. C. californica has an olive brown to
greenish black thallus and a greenish to brown-
black epithecium usually reacting K + violet,
whereas B. abbreviata is always dull red-brown
Fig. 63. Bryoria abbreviata. Note the “ciliate”
apothecia. B. C., Creston, Brodo 15120 (CANL).
Scale 0.5-1 .0 mm.
and has a red-brown epithecium not reacting
with K. The cortical structures in these two
species also differ (see general section on
Anatomy and morphology: Cortex).
Selected specimens
Exsiccatae. Cummings, Williams & Seymour: Lieh.
Bor. Amer. 18: Montana, Flathead County: Columbia
Falls, Williams (FH, H. NYBG) - Cummings, Sey-
mour & Williams: Dec. N. Amer. Lieh. 96: Montana,
Flathead County: Columbia Falls, Williams (CANL,
DUKE, MSC, NYBG, US) - Krypt. Vind. 3711:
Washington. Whitman County: Pullman, Herre
(COLO, F, LAM. US) - Merrill: Lieh. Exs., Ser. II,
36 (some mixed with B. efr . fuscescens). Washington,
Ferry County: Republic, Foster (F, MSC, NYBG,
US. WIS).
Canada. British Columbia. Columbia River Basin:
Kootanie Lake (sic), Macoun (440?) (CANL) - Fraser
River Basin: Wells Gray Provincial Park, Ahti 14269
(H) - Kootenay River Basin: East of Creston, Brodo
15120 (CANL) -Okanagan River Basin: Mt. Anarchist
near Osoyoos, Krajina 645-124 (CANL) - Alberta.
Waterton Lakes National Park, Scotter4901 (WIS).
U.S.A. California. Del None County: Near Fair-
view Peak in the Siskiyou National Forest, Pike
L-329A (OSC) - Los Angeles County: San Gabriel
Mts.. Hasse 1485 (NYBG) - Tulare County: Sequoia
National Park. Rundel 2569 (UC1) - Idaho: Bonner
County: Priest R. Experimental Forest, on Gisborne
Mt., Esslinger 392 (COLO) - Montana: Missoula
County: S of Alberton. J. W. Thomson 16328 (WIS) -
Oregon. BakerCounty: Unity, Shushan SL-3961 (BM,
CANL, US, WIS) -Union County: Summit Pass, Blue
Mt., Imshaug 19 (COLO) -Washington. Ferry County:
Republic, Foster 36 (COLO) - Pierce County: Mt.
Rainier National Park, Brodo 15475 (CANL) -
134 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 64. Known distribution. - A: Bryoria abbreviata. - B: B. oregana.
Wyoming. County unknown: Belt Mts., Willey s.n.,
no date (US).
Mexico. Baja California. Cape San Quentin (sic)
(San Quintin), Greene s.n., Feb. 1906 (FH), Greene
189? (NYBG ).
25. Bryoria oregana (Tuck, ex Nyl.) Brodo &
D. Hawksw., comb. nov.
Alectoria oregana Tuck, ex Nyl., Lieh. Jap.: 104
(1890); type: Union, Oregon, W. C. Cusick (FH-Tuck.
472 holotype! H-Nyl. 35971 isotype!); basionym. -
Bryopogon oreganus (Tuck, ex Nyl.) Schneid., Guide
Stud. Lieh.: 166 (1898). - Coelocaulon oreganum
(Tuck, ex Nyl.) Choisy, Icon. Lieh. Univ., ser. II,
fasc. 2, sine pagin. (1931).
Thallus pendent, up to 17 cm in length; branch-
ing anisotomic dichotomous, angles between the
branches acute or obtuse; branches very uneven
in diameter, 0.25-0.40 mm diam. at the base,
0.10-0.20 mm on pendent branches; branches
longitudinally foveolate and channeled, gener-
ally very angular; red-brown, matt (never shiny).
True lateral spinules, soredia, and pseudocy-
phellae absent; branches sometimes appearing
spinulose due to short, pointed, lateral branches.
Apothecia infrequent, lateral; excipulum thalli-
num concolourous with thallus, smooth or
sometimes ciliate, disappearing in age: to 2.0
mm diam.
Spores 8 per ascus, broadly ellipsoid, thin-
walled, c. 6.5 x 4.5 /urn. Pycnidia not seen.
Cortical hyphae in surface view short,
knobbed, thin-walled, appearing like a jigsaw
puzzle (Fig. 12 A).
Cortex and medulla K-, C-, KC-, PD-;
contains no lichen substances.
Ecology. On conifers, especially Abies lasio-
carpa, Picea, and Pinas species, usually be-
tween 1500 and 2300 m in elevation, charac-
teristically in subalpine Abies lasiocarpa-Picea
engelmannii stands, but also in dry or exposed
montane pine stands.
Distribution. Western montane, from south-
western Alberta and neighbouring Montana
westward to the coast and southward through
California (Fig. 64 B). Apparently, B. oregana
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 135
requires, or is tolerent of, cooler, more humid
habitats than is B. abbreviate!, and is therefore
more commonly found in the coastal mountains.
B. oregana, like B. abbreviata, is a North
American endemic.
Discussion
Bryoria abbreviata and B. oregana have gener-
ally been regarded as synonymous (Howe 191 1,
Motyka 1964). While the two taxa are clearly
very closely allied, intergrades appear to be rare.
The two species are able to grow side by side in
nature and maintain their identities. B. abbrevia-
ta is invariably fertile and has large, ciliate,
subterminal apothecia and a distinctly caespitose
habit (although one or two of the marginal cilia
may become elongate and develop into pen-
dulose branches). In contrast, B. oregana only
rarely forms apothecia. When they do occur they
are almost eciliate and always clearly lateral.
Furthermore, the thallus is pendent rather than
caespitose.
These two species frequently occupy the same
habitat in regions where they are sympatric and
their thalli may be intricately entangled. It is
undoubtedly this frequent admixture which has
led to their being treated as conspecific in the
past.
Sterile plants of B. oregana may be con-
fused with some slender sterile and esorediate
morphotypes of B. fremontii, but these taxa can
usually be separated reliably on the basis ot
thallus colour and cortical features (see Fig. 12).
The holotype collection of B. oregana is
mixed and comprises intricately entangled (a)
typical elongate and fertile thalli of B. oregana
s. str., (b) a large amount of B. fuscescens,
(c) some B. fremontii, (d) Letharia Columbiana,
and (e) at the base of the specimen, what appears
to be B. abbreviata with broader more divergent
branches. A dissection of the collection would
have resulted in the destruction ot part of the
type and so was not attempted. Isotype material
in Nylander's herbarium (H !). undoubtedly sent
to Nylander by Tuckerman, is B. oregana as
interpreted here. The epithet oregana thus refers
to the pendent rather than the caespitose
elements of "Alectoria oregana auct ." and we
have adopted it in this sense.
Selected specimens
Canada. British Columbia. Coastal District: Gari-
baldi Mountains, Garibaldi. Brodo 8178 (CANL) -
Columbia River Basin: GlacierNational Park, Ohlsson
395 (MSC) - Fraser River Basin: Southern Cariboo
Mountains. Wells Gray Park, Ahti 6517 (H) - Islands
South District: Between Mt. Thornhill and Mt. Attree,
Ohlsson 2568 (MSC, CANL) - Skagit River Basin:
Manning Park. Ahti 15388 (H) - Alberta. Waterton
Lakes National Park. W of Avion Ridge, Bird 16836
(CANL).
U.S.A. California. Del Norte County: Near Bear
Basin. Tucker 6434 (US, COLO) - Riverside County:
San Jacunta Mountain (sic) (Jacinto), Hasse s.n., 1904
(US) - San Diego County: San Diego, Orcutt s.n.. no
date (F) - Santa Cruz County?: Santa Cruz Mountains,
Castle Rock, Herre 1 144 (F) - Idaho. Payette County:
Payette Lakes, A. H. Smith I6076C (DUKE.COLO)-
Montana. Glacier County: Glacier National Park.
Sprague Creek. Imshaug 7341 (LAM, US) - Sanders
County: Fishtrap Creek, Taylor 7071 (p.p.) (Herb, not
indicated) - Oregon. Benton County: Mary’s Peak W
of Corvallis. Pike 3106 (CANL) - Klamath County:
Crater Lake National Park. Sipe 1152 (ORE) -
Washington. Clallam County: Olympic National Park,
near Mt. Angeles, Ireland 6898 (US) - Pend Oreille
County: Colville National Forest, Layser 1460 (p.p.)
(US).
Bryoria sect. Tortuosae (Bystr.) Brodo & D.
Hawksw., comb. nov.
Bryopogon sect. Tortuosae Bystr., Ann. Univ. Mariae
Curie-Sklodowska, C, 26: 273 (1971), as "Mot.";
basionym. - Alectoria subgen. Bryopogon sect. Tor-
tuosae Mot., FI. Polska, Porosty 5(2): 38 (1962); nom.
inval. (Art. 37).
Type species: Bryoria tortuosa (Merr.) Brodo &
D. Hawksw. (syn. Alectoria tortuosa Merr., Bryo-
pogon tortuosus (Merr.) Bystr., nom. inval.; holo-
type).
Thallus subpendent to pendent; emerald-green-
ish to cervine brown, not blackened. True lateral
spinules and isidia absent. Soralia rather rare,
tuberculate, emerald-green to greenish yellow.
Pseudocyphellae present and abundant or sparse
to almost absent, conspicuous, depressed,
elongate-fusiform, greenish yellow in one
species. Cortex with a moderate amount of
matrix, longitudinally orientated (not cellular
and knobby) at the surface. Apothecia occa-
sional; margins not ciliate; discs yellowish to
emerald-green, often markedly convex. Vulpinic
acid known from all species; atranorin and/or
chloroatranorin, fumarprotocetraric acid, and
other /S-orcinol depsidones absent.
136 Alectoria in North America
OPERA BOT. 42 (1977)
Discussion
This section comprises two species, the circum-
boreal B. fremontii and the western North
American B. tortuosa, which are unique in
Bryoria in having pulvinic acid derivatives. (It
should be noted that in B. fremontii these are
restricted to the soralia and apothecia.)
26. Bryoria fremontii (Tuck.) Brodo & D.
Hawksw., comb. nov.
Alectoria fremontii Tuck., Am. J. Arts Sei., ser. 2,
25: 422 (1858); type: Sierra Nevada, camp of Decem-
ber 5-6, 1845, a piece of moss which is abundant in
this region, Fremont (FH-Tuck. 498 holotype!);
basionym. - Bryopogon fremontii (Tuck.) Rabenh.,
Flecht. Eur. no. 918 (1871). - Alectoria jubata var.
fremontii (Tuck.) Boist., Nouv. Fl. Lieh. 2: 38 (1903).
Alectoria olivacea Räs., Medd. Soc. Fauna FI.
Fenn. 43:4(1917); type: Fennia, Ostrobottnia borealis,
Simo, V. Räsänen, 6 June 1916 (H lectotype!). -
Bryopogon olivaceus (Räs.)Choisy, Icon. Lieh. Univ.,
ser. 2, 2: sine pagin. (1930). - Alectoria fremontii var.
olivacea (Räs.) H. Magn., FI. Skand. Busk. Bladl. 100
(1929). - Alectoria fremontii subsp. olivacea (Räs.)
Räs., Lieh. Fenn. Exs. no. 162 (1936). - Alectoria
fremontii f. olivacea (Räs.) Mot., FI. Polska, Porosty
5(2): 76 (1962).
Alectoria corneliae Gyeln., Ann. Crypt. Exot. 4:
171 (1931); type: Oregon, near Brownsville, on pine,
L. L. Haskin 1031, 25 May 1930 (US holotype!). -
Bryopogon olivaceus f. corneliae (Gyeln.) Gyeln.,
Feddes Repert. 38: 230 (1935). - Alectoria olivacea f.
corneliae (Gyeln.) Zahlbr., Cat. Lieh. Univ. 10: 559
(1940).
Alectoria fremontii subsp. erikssonii DR.. Ark. Bot.
20A (11): 8 (1926); nom. nov. for A. fremontii sensu
Räs., non Tuck. - Bryopogon fremontii f. erikssonii
(DR.) Saviez, Lieh. Ross. 4 no. 34 (1935). - Alectoria
fremontii f. erikssonii (DR.) Mot., FI. Polska, Porosty
5(2): 77 (1962); nom. inval. (Art. 33).
Alectoria fremontii var. perfertiUs Räs., Ann. Mis-
souri Bot. Gard. 20: 10(1933); type: British Columbia,
Golden, ad corticem arboris ( Pinus ), V. Kujala, Sep-
tember 1931 (H holotype!). - Bryopogon olivaceus
var. perfertiUs (Räs.) Gyeln., Feddes Repert. 38: 230
(1935). - Alectoria olivacea var. perfertiUs (Räs.)
Zahlbr., Cat. Lieh. Univ. 10: 559 (1940).
? Alectoria fremontii f. dealbata Mot., FI. Polska.
Porosty 5 (2): 77 (1962); nom. inval. (Art. 37).
Thallus pendent, often very long, to 45 cm (or
exceptionally 90 cm, fide Turner in litt.);
branching anisotomic dichotomous to submono-
podial, usually frequent from the base; branches
uneven in diameter, becoming twisted and
foveolate, sometimes becoming dorsivcntrally
compressed and expanded towards the base,
(0.2-) 0.4-1. 5(^4. 0) mm diam.; yellowish brown
to reddish brown or dark brown, colour often
variable within the same specimen, usually
shiny.
True lateral spinules absent. Soralia absent or
sparse, tuberculate, pale yellow to bright yellow,
usually as wide as the branches on which they
occur, to 2.0( — 4.0) mm long. Pseudocyphellae
absent or sparse (perhaps only torsion cracks),
depressed and elongate fusiform, white to pale
brown.
Apothecia absent or sparse, lateral; excipulum
thallinum concolourous with the thallus, thin and
soon becoming excluded; disc becoming convex
at maturity, 1.0-2.0(-4.0) mm diam., yellow
pruinose. Spores 8 per ascus, hyaline, sub-
globose, simple, 5-8 x 4-5 /zm. Pycnidia not
seen.
Cortex, medulla, apothecia, and soralia K— ,
C — , KC-, PD-. Contains vulpinic acid in the
soralia and apothecia.
Ecology. Mainly on Pinus ponderosa, P. con-
forta, Larix occidentalis, and Pseudotsuga
menziesii in open pine or Douglas fir forests
(at elevations of 700-2500 m in the Canadian
Rockies and 3000-3400 m in the southern
Rockies); also on oaks closer to the coast. It
seems to prefer somewhat open and rather dry
sites (Szczawinski 1953).
Distribution. Western conifer forests from mid-
Alberta and British Columbia west to the coast
and south throughout California to Baja Cali-
fornia, with disjunct populations in Colorado and
the Black Hills (Fig. 65A). Wetmore (1968)
regards the presence of B. fremontii in the Black
Hills as a relatively recent (i.e., post-Pleisto-
cene) occurrence due to its rarity. B. fremontii
shows a classical western American-western
European disjunct distribution. In Europe it is
known from Fennoscandia and the Caucasus
(Krog 1968), and Schwarzwald (Schindler 1974),
but it is especially abundant in northern Fin-
land (Ahlner 1948).
Bryoria fremontii was determined from eastern
Asia (Saghalien) by Gyelnik (1935, Sato 1936)
and has also been reported from Mexico
(Williams 1895) and Ellesmereland (Darbishire
1909). These three records require confirmation
but we have seen one specimen which may be
this species from Ecuador (Gualaquiza [^Guaya-
quil], 1200 m, Abate M. Allioni, 1910, comm.
Sbarbaro [KGY]). Its absence in the British
Isles and distribution in the drier Douglas fir
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 137
Fig. 65. N. American distribution. -A: Bryoria fremontii. - B: B. tortuosa.
forests of western America would tend to
support Ahlner’s (1948) assessment of the
species as “northern continental’' although he
does point to its occasional occurrence in
somewhat oceanic localities.
The presence of a northern species like B.
fremontii in Baja California is apparently not
unusual since Stebbins & Major (1965 p. 5)
point out that “Boreal plants of holarctic distri-
bution extend southward along the California
coast to Monterey, and in the mountains
throughout the Sierra Nevada and even into
southern California." The jump between
southern California and northern Baja California
is not very far with regard to distance or climate.
Discussion
Bryoria fremontii is an extremely variable
species in some respects but is generally un-
likely to be misidentified, even when sterile,
owing to its shiny, broad, and foveolate main
stems and its characteristic yellowish brown to
dark reddish brown colour. Some confusion
can occur with unusually large and acid-deficient
plants of B. trichodes subsp. americana. Very
slender, dark specimens can be distinguished
from B. oregana by examinations of the cortical
structure (Fig. 12). Specimens appearing similar
to B. fremontii but differing in being unusually
pale, dull, and quite brittle often prove to be
B. friabilis on examination by TLC. (The latter
species contains gyrophoric acid.)
Specimens of B. fremontii with both yellow
soralia and yellow-pruinose apothecia are
extremely rare in North America, as they are in
Europe, though they are more frequent in Cali-
fornia than elsewhere. Specimens with only
soralia or only apothecia are almost as rare.
The most commonly encountered morphotype is
dark red-brown to dark brown in colour, with
very thick and foveolate, shiny, main branches
138 Alectoria in North America
OPERA BOT. 42 (1977)
and numerous slender side branches. Various
infraspecific taxa have been proposed to accom-
modate the morphotypes with and without
soralia, etc. (see Motyka 1962), but these
variations seem to be without any taxonomic-
significance. Material with white soralia (e.g.,
“f. dealbata Mot.”) is mentioned in the litera-
ture but we have seen none in our investiga-
tions and are sceptical as to its existence; such
reports may well be based on misidentifica-
tions although it is occasionally possible to find
old soralia rubbed free of their coating of vul-
pinic acid so as to appear almost white. (In
such cases, typical yellow soralia are invariably
present elsewhere on the same specimen.)
On the west coast, there occurs a population
of this species with an unusually pale, almost
yellow-brown thallus and white incipient
pseudocyphellae (perhaps actually ”torsion”
cracks). Plants belonging to this morphotype,
which includes the type specimen of Alectoria
corneliae Gyeln., are frequently extremely
robust and have very thick shiny branches.
This morphotype might at first appear to merit
recognition at some infraspecific rank, but since
a continuous series of intermediate plants exists,
we do not consider it to be taxonomically
important.
Because the vulpinic acid in this species is
restricted to soralia and apothecial discs, it
cannot be demonstrated by TLC in material
lacking these structures.
Attention is drawn to Ahlner’s (1948) impor-
tant discussion of the variability of this species
in Europe.
Selected specimens
Exsiccatae. Cummings, Seymour & Williams: Dec.
N. Amer. Lieh. 54: Montana, Flathead County:
Columbia Falls, Williams (CANL, DUKE, MSC,
NYBG. US, WIS) -Cummings. Williams & Seymour:
Lieh. Bor. Amer. 17: (same as above) (COLO) -
Merrill: Lieh. Exs., Ser. II, 47: Washington, Ferry
County: Republic, Foster (COLO, F, MSC, NYBG,
WIS) - Rel. Tuck, (no number): Wyoming, County
unknown. Wind River Mountains, Hayden (LAM) -
Tuckerman: Lieh. Amer. Septentr. Exs. 52: Califor-
nia, no specific locality. Fremont (NYBG) - Weber:
Lieh. Exs. 418 (p.p.): California, Shasta County:
Burney, Pike & Jackman (CANL).
Canada. British Columbia. Islands South District:
Vancouver Islands: Little Qualicum Falls Provincial
Park, Becking s.n., 25 Sept. 1953 (LAM) - Coast
Area: Garibaldi Park, Krajina 351-4 (CANL) -Colum-
bia River Basin: Deer Park, Macoun 55 (NYBG, US)
- Fraser River Basin: Blue River, Brodo 15791
CANL. WIS) - Okanagan River Basin: Princeton,
Brodo 7812 (CANL) - Peace River Basin: Mcleod
Lake, Leechman s.n., July 1954 (FH) - Skagit River
Basin: Manning Park, Ahti 15389 (H) - Skeena River
Basin: E of Hazelton, Ohlsson 2962A (MSC) -
Alberta. Banff National Park: Trail to Lake Agnes,
Imshaug 6866 (WIS).
U.S.A. California. Eldorado County: near Kyburz,
EldoradoNat. Forest, Belshaw6005 (DUKE, COLO)-
Los Angeles County: St. Antonio Mts., Johnston
3085 (FH, F) - Colorado. Routt County: Seedhouse
Guard Station, Weber 6835 (COLO) - Idaho. Latah
County: Kendrick, Esslinger 248B (COLO) - Mon-
tana. Missoula County: S of Alberton, J. W. Thomson
16319 (US, WIS, CANL) - Oregon. Klamath County:
Crater Lake, Sipe 990 (ORE, F) — South Dakota.
Custer County: Custer, Wetmore 10053 (MSC) -
Washington. Asotin County: Fields Spring State Park,
Imshaug 16483 (MSC, CANL) - Wyoming. Yellow-
stone National Park: Near Canyon Junction, Imshaug
9441 (WIS, LAM).
Mexico. Baja California: Cape San Quentin (sic)
(San Quintin), Greene s.n., Feb. 1906 (FH) -Nacho-
guero Valley, Schoenfeldt 3427 (US).
27. Bryoria tortuosa (Merr.) Brodo & D.
Hawksw., comb. nov.
Alectoria tortuosa Merr., Bryologist 12: 5 (1909); type:
British Columbia, near Westminster, A. J. Hill. July
1904 (FH holotype!); basionym. - Eualectoria tor-
tuosa (Merr.) Gyeln., Ann. Mus. Nat. Hung., Bot. 28:
283 (1934); nom. inval. (Art. 34). — Bryopogon
tortuosus (Merr.) Gyeln.. Feddes Repert. 38: 229
(1935).
rhallus pendent, 10-30 cm long; branching
mainly anisotomic dichotomous; angles between
the dichotomies acute, with frequent slender,
perpendicular side branches arising from the
axes; branches uneven in diameter, strongly
twisted and tortuous, foveolate and often flat-
tened. 0.4 — 0.5(— 1 .0) mm in diam.; dull, dark red-
brown to dusky yellow-brown, occasionally be-
coming bright yellow in thalli having heavy con-
centrations of vulpinic acid; pale and dark speci-
mens often grow mixed.
Spinules and isidia absent; soredia exceed-
ingly rare. Pseudocyphellae usually abundant,
occasionally rare, bright yellow, linear or some-
times short fusiform, slightly raised, twisting
around filaments in long yellow spirals (Fig. 5C).
Apothecia rare, lateral, 2-3 mm diam., excipu-
lum thallinum raised, persistent, disc heavily
yellow pruinose. Spores 7. 5-8. 7 x 4. 7-5.0 /urn,
8 per ascus, hyaline, ellipsoid. Pycnidia un-
known.
OPERA BOT. 42 (1977)
Bryoria Brodo & D. Hawksw. 139
Cortex and medulla K— , C— , KC— , PD — ,
but thallus producing a yellowish stain on filter
paper with PD in most specimens. Contains
vulpinic acid.
Ecology. On trees in well-lighted, open stands.
Most frequently on oaks and pines, although it
has been collected on a large variety of trees
and shrubs.
Distribution. Along the west coast, mainly from
southern British Columbia to central California;
with scattered localities in central British Colum-
bia, mainly on exposed hillsides or in pine
stands (Fig. 65 B). The Idaho localities are also
in ponderosa pine stands fitting in well with its
known ecology (see above). The only reported
locality outside of North America for B. tortuosa
is in the Carpathian Mountains in Europe
(Motyka 1958), but the specimen upon which
this record is based has not been seen by us.
The distribution of the moss Crummia latifolia
(common in western North America with a
single occurrence in the Caucasus Mountains)
is very similar (Schofield & Crum 1972 p. 175,
map 26).
Discussion
Bryoria tortuosa is readily recognizable by its
red-brown to dusty yellow-brown to almost
greenish yellow thallus, and yellow, generally
conspicuous, pseudocyphellae. Specimens with
very few pseudocyphellae are likely to be con-
fused with B. fretnontii, so a careful examina-
tion of all material of the latter is advisable:
in critical cases, TLC always reveals vulpinic
acid in B. tortuosa while in B. fretnontii it is
only found associated with soralia and apothe-
cia.
The ability to produce vulpinic acid varies
considerably in this species and is responsible
for the marked variations in the colour of the
thallus. Numerous packets contain mixtures of
both greenish yellow thalli with abundant vul-
pinic acid and dark brown thalli showing only
yellow pseudocyphellae; all intergradations
occur.
The name Alectoria virens Tayl. was incor-
rectly applied to this species by Howe (1911).
A. virens is a very different species unknown
outside Asia (see Excluded taxa).
A single specimen from Vancouver Island,
B. C. (Tibell 5083 [UPS]), was seen with scat-
tered, yellow, tuberculate soralia. It was bright
yellow with conspicuous yellow pseudocy-
phellae and was therefore definitely not B.
fretnontii. Soralia have not previously been
reported from B. tortuosa.
Selected specimens
Exsiccatae. Brodo: Lieh. Can. Exs. 3: British Colum-
bia, Vancouver Island, Saanich, Brodo 8096 (CANL)
- Brodo: Lieh. Can. Exs. 4: British Columbia, 29
miles S of Merritt. Brodo 8496 (CANL) - Merrill:
Lieh. Exs., Ser. I, 160: Oregon. Marion County:
Silverton. Foster (BM, CANL. DUKE. MSC.NYBG,
US) - Weber: Lieh. Exs. 418 (p.p.): California,
Shasta County: Burney, Pike & Jackman (CANL).
Canada. British Columbia. Coastal District: Earl's
Cove, Brodo 8343 (CANL) -Dean River Basin: Along
Mosher Creek, Tweedsmuir Provincial Park, Ohlsson
1932A (p.p.) (MSC) - Fraser River Basin: Beaver
Lake, Macoun 417 (CANL) - Islands South District:
Between Parkville and Port Alberni, Tibell 5083 (UPS)
- Skeena River Basin: 30 miles W of New Hazelton.
Ohlsson 2887 A (MSC).
U.S.A. California. Humboldt County: Near Manila,
Lindsay 90 (HSC) - Santa Cruz County: Boulder
Creek, Hesse 3056 (CANL. COLO, F, NYBG) -
Trinity County: Weaverville, Richards & Drouet
1315 (FH, F, US) - Idaho. Idaho County: Stites,
Cooke 22967 (W1S) - Latah County: Kendrick,
Esslinger 232B (WIS?) - Oregon. Jackson County:
Eagle Point, Sipe 640 (MSC, ORE) - Washington.
Clallam County: Dungeness, Foster 12 (US) - Okano-
gan County: Winthrop, 8 mile Creek, Hasles s.n.
(p.p.), 1950 (WIS).
Pseudephebe Choisy
Icon. Lieh. Univ., Ser. 2, Fase. 1: sine pagin. (1930).
- Alectoria sect. Pseudephebe (Choisy) Choisy, Bull.
Mens. Soc. Linn. Lyon 24: 26 (1955). -Parmelia sect.
Teretiuscula Hillm.. Rabenh. Krypt.-Fl. 9, 5(3): 104
(1936). - Alectoria sect. Teretiuscula (Hillm.) Lamb,
Br. Antarct. Surv. Sei. Rep. 38: 26 (1964). - Alectoria
subgen. Teretiuscula (Hillm.) D. Hawksw.. Licheno-
logist 5: 201 (1972). - Alectoria sect. Subparmelia
Degel., Nyt Mag. Naturvid. 70: 286 (1938).
Type species: Pseudephebe pubescens (L.) Choisy
(syn. Lichen pubescens L.; holotype).
Thallus fruticose, sometimes becoming compact-
ed and suberustose centrally, prostrate, closely
adpressed to the substrate (under 1 cm tall),
attached to the substrate by hapters over the
whole thallus area and not merely at the base;
base sometimes dying so that the thalli form
rosettes; branching isotomic-dichotomous, often
strictly so, branches terete bul tending to
become dorsiventrally compressed in one
species, minute, even to uneven; brown to dark
or black, matt to slightly shiny, usually con-
colourous except where attached to the sub-
strate.
True lateral spinules, isidia, soralia, and
pseudocyphellae absent. Cortex composed of
longitudinally orientated hyphae which become
prosoplechtenchymatous to almost paraplecten-
chymatous at the surface. The surface itself
is smooth or slightly rough. Medullary hyphae
not ornamented.
Apothecia lateral, frequent to abundant in
both species; excipulum thallinum concolourous
with the thallus, becoming excluded or, if per-
sistent, not incurved, sometimes ciliate; disc
brown to almost black. Asci clavate, thick-
walled, arrested bitunicate, 8-spored. Asco-
spores ellipsoid, lacking a distinct epispore,
hyaline at maturity, simple, 7-12x6-8 pm.
Pycnidia common.
Lichen products absent.
Discussion
The systematic position of the species compris-
ing the genus Pseudephebe has long been a
cause of controversy, with some authors placing
them in Alectoria, and others classifying them
under Parmelia. They cannot really be satisfac-
torily placed in either of these two genera and
are most appropriately regarded as constituting
a distinct genus. With regard to Parmelia,
Pseudephebe is clearly allied to the subgenus
Melanoparmelia Hue, but differs from most
species of that subgenus in the fruticose rather
than foliose habit and in the absence of lichen
substances. It is probable that some species with
almost fruticose "lobes” currently generally
placed in Melanoparmelia may eventually prove
to be additional members of Pseudephebe. We
have not investigated those species in view of
work in progress elsewhere.
In Pseudephebe the cortex is composed of
longitudinally orientated hyphae as reported by
Degelius (1937). However, where the hyphae
reach the surface, a “layer” of short almost
isodiametric cells is formed (Fig. 13) quite unlike
that seen in other taxa treated here, with the
possible exception of species placed in Bryoria
sect. Subdivergentes. In that section of Bryoria,
the superficial cortical cells are extremely
irregular in shape and project above the surface
to produce a dull, matt texture and appearance.
In Pseudephebe the surface of the cortex,
although “cellular”, is only very slightly
roughened by unevenness of the cells.
1. Pseudephebe minuscula (Nyl. ex Arnold)
Brodo & D. Hawksw., comb. nov.
Imbricaria lanata var. minuscula Nyl. ex Arnold,
Ver. Zool.-Bot. Ges. Wien 28: 293 (1878); type:
Fennia, Lapponia enontekiensis. Enontekio, in alpe
OPERA BOT. 42 (1977)
Pseudephebe Choisy 141
Pietsovaara prope Kilpisjärvi, J. P. Norrlin, 1867
(H-Nyl. 34255 lectotype!); basionym. - Parmelia
minuscula (Nyl. ex Arnold) Nyl., Bull. Soc. Linn.
Normand., sér. 4, 1: 205 (1887). -Cornicularia lanata
var. minuscula (Nyl. ex Arnold) Hue. Deux. Exped.
Antarct. Fr., Lieh.: 41 (1915). - Alectoria minuscula
(Nyl. ex Arnold) Degel., Nyt Mag. Naturvid. 78:
286(1938).
Parmelia pubescens var. congesta Zahlbr., Dtsch.
Südpol. Exped. 8: 52 (1928); type: Antarctica. Kaiser
Wilhelm II Land. Gaussberg. Drygalskey's Expedi-
tion, 1901-03 (BM lectotype!). - Alectoria congesta
(Zahlbr.) Dodge, B.A.N.Z. Antarct. Res. Exped.
1929-31, B, 7: 195 (1948). - Alectoria minuscula f.
congesta (Zahlbr.) Lamb, l.illoa 14: 244 (1948).
Parmelia minuscula subsp. minutissima Räs.,
Medd. Soc. Fauna Fl. Fenn. 43: 118 (1919); type:
Finland, Ostrobottniae borealis. Simo. Onkalo, kivia-
dala. V. Räsänen. 26June 1915 (H holotype!).
Alectoria antarctica Dodge & Baker, Ann. Missouri
Bot. Gard. 25: 599 (1938) p.p.; type: Antarctica, Marie
Byrd Land, Edsel Ford Range. Mt. Rea-Cooper. P.
Siple et al. R-l (herb. Dodge, holotype slide!); nom.
illegit. (Art. 70).
Parmelia minuscula f. aperta Lynge, in Lynge &
Scholander, Skr. Svalbard ishavet 41: 71 (1932): type:
Greenland, Kapp Humboldt, B. Lynge, 3 August 1929
(O lectotype !).
Parmelia minuscula f. applanata Lynge, in Lynge
& Scholander, Skr. Svalbard Ishavet 41: 71 (1932);
type: Greenland, Ostgronland, Kjerulffjorden, B.
Lynge. 13 August 1929 (O lectotype!). - Alectoria
minuscula f. applanata (Lynge) Lamb, Nyt Mag.
Naturvid. 80: 264(1939).
Parmelia minuscula f. crustacea Lynge. in Lynge &
Scholander, Skr. Svalbard Ishavet 41: 71 (1932); type:
Greenland. 0stgrpnland, Jacksonoya. B. Lynge, 31
July— 1 August 1929 (O lectotype!). - Alectoria
minuscula f. crustacea (Lynge) Degel. , Bot. Notiser
1943: 108 (1943).
Parmelia minuscula f. luxurious Lynge, in Lynge
& Scholander. Skr. Svalbard Ishavet 41: 71 (1932);
type: Greenland. Ostgronland, Kjerulffjorden, B.
Lynge, 13 August 1929 (O holotype!).
Parmelia pubescens f. biformis Vain., Res. Voy.
S. V. Belgica 1897-99. Lieh.: 14 (1903); type: An-
tarctica (TUR syntypes: not seen; see Lamb, 1948:
243). - Alectoria minuscula f. biformis (Vain.) Lamb,
Lilloa 14: 243 (1948). - Alectoria biformis (Vain.)
Dodge, Lieh. FL Antarct.: 219 (1973).
Parmelia almquistii f. opaca Lynge. Rep. Scient.
Results Norw. Exped. Nova Zemlya 43: 197 (1928):
type: Nova Zemlya, Belushi Bay, B. Lynge, 1921 (O
holotype; not seen).
For further information on the synonymy of this
species, see Lamb (1948, 1964).
Thallus prostrate and decumbent, in extreme
forms becoming almost subfoliose or even
crustose and pulverulent, generally forming
more or less circular adpressed thalli up to 7 cm
diam.; branches always somewhat flattened,
especially on older portions, up to 1 .0 mm broad.
tapering irregularly to 0.2-0. 5 mm on main stems
and 0. 1 mm at tips where the branches are very
uneven, but more or less terete; branching ir-
regularly isotomic dichotomous with numerous
short lateral branches often present giving
thallus a “ciliate” appearance; internodal
distance short, usually c. 0.2-0. 5(— 1 .0) mm; dark
brown to black, shiny or dull, occasionally
pale brown at the base.
True lateral spinules absent, but simple or
branched isidioid adventitious branches are not
uncommon in one morphotype. Soralia and
pseudocypheliae absent (or brown and incon-
spicuous in one specimen?).
Apothecia frequent, disc dark red-brown to
black, flat, to 3 mm diam.; margins persistent,
smooth or verrucose; spores 8 per ascus, 7.7-
8.0x6. 7-9.0 ju.m, hyaline. Pycnidia very abun-
dant, embedded in the thallus and opening to
the surface through broad ostioles; conidia
6-8 x 2-1 pm.
All tissues of thallus K-, C— , KC-, PD-;
no lichen substances present.
Ecology. On siliceous, or sometimes lime-
containing rocks in alpine or arctic habitats.
John Thomson (in litt.) considers it to be charac-
teristic of windswept gravels and rock outcrops.
Imshaug (1957) pointed out that P. minuscula
is more abundant in drier, more exposed local-
ities than is P. pubescens.
Distribution. Circumpolar arctic-alpine (Fig. 66).
There seems to be a tendency for P. minuscula
to be more continental in its distribution than
P. pubescens, although the two are often found
together. Weber (in litt.) has observed the same
thing. P. minuscula is almost entirely absent
from the mountains on the British Columbia
coast and is fairly common in the southern
Rockies. It was growing luxuriantly on one of
the dry subalpine summits of the San Francisco
Mountains in Arizona. The species is also known
from a number of localities in the Southern
Hemisphere. In Antarctica, it is circumpolar,
whereas P. pubescens is restricted to the milder
Antarctic Peninsula no further south than the
Marguerite Bay area (Lamb 1964).
Discussion
Pseudephebe minuscula, like other lichens
growing under extremely rigorous conditions, is
142 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 66. Pseudephebe minuscula. N. American distribution.
very variable in its superficial appearance,
depending on the protection and moisture avail-
able to the thallus. Typically, the thallus is
conspicuously flattened and very minute, but
robust specimens can become almost ascendent
and develop terete branches which appear very
like those of P. pubescens. Similarly, under
extremely severe conditions, P. pubescens can
become very adpressed and approach P. minus-
cula. As a result of environmental pressures
tending to induce a similar phenotype, as
Imshaug (1957) has already pointed out, some
thalli will always remain difficult to refer to one
species or the other. Imshaug (1957) accurately
described the situation with regard to the diffi-
culties in the taxonomy of this pair of species.
The environmental modifications in both
species have been given a large number of infra-
specific names, mainly at the rank of variety or
form (e.g., Lynge & Scholander 1932, Hillman
1936, Lamb 1948, 1964, Motyka 1960a). Most
of these are probably of little taxonomic im-
portance and we have not attempted to differen-
tiate one from another and assess their relative
OPERA BOT. 42 (1977)
Pseudephebe Choisy 143
importance in this study. One morphotype of
P. pubescens, f. subciliata (see p. 156), may,
however, be genotypically distinct (see Hawks-
worth 1972) but that form appears to be ex-
tremely rare in North America.
Dodge (1973 p. 218) endeavoured to separate
five species in this group on the basis of
differences in habit, branching, and measure-
ments of various anatomical characters. In our
view, his system is impractical, being mainly
based on characters of dubious taxonomic value.
Selected specimens
Canada. British Columbia. Fraser River Basin: Wells
Gray Provincial Park, Ahti 14158 (H) - Kootenay
River Basin: Lake Oesa, MacFadden 62 (US) -
Alberta. Jasper National Park, Queen Elizabeth
Ranges, Scotter 7797 (CANL) - Manitoba. Churchill.
J. W. Thomson 3772 (CANL) - Quebec. Territoire
du Nouveau Québec: Fort Chimo, J. W. Thomson
13778 (CANL, US) - Newfoundland. Grand Falls
District: Summit of Hodges Hill. Ahti 7740 (H) -
Yukon. 62°39'N , 134°45'W, Bostock 34 (CANL) -
Kluane Lake, Murray L-41336 (FHl - Northwest
Territories. District of Franklin: Axel Heiberg Island:
White Glacier, Kuc 85 (CANL) - Devon Island:
Dundas Harbour, Innes-Taylor 11 (CANL) - Melville
Island: Bridport Inlet, Tener 395 (CANL)- Mackenzie
District: Bernard Harbour, Johansen s.n.. June 1914
(FH) - Keewatin District: Southampton Island,
Salmon Pond, Parker SP-70- 1 6C (CANL).
V.S.A. Alaska. Alaska Range District: E fork of
the Kuskokwim River, Viereck 5117A (ALA)- Arctic
Coast District: Okpilak Lake, J. W. Thomson &
Shushan 10159 (US) - Eastern Pacific Coast District:
Muir Glacier, Kincaid s.n., 13 July 1897 (CANL) -
Arizona. Coconino County: San Francisco Peaks,
Nash 7561 (CANL, MIN) - California. Amador
County: Silver Lake, Thiers 17241 (HSC) - Calaveras
County: Ebetts Pass, Thiers 17664 (CANL) -Riverside
County: San Jacinto Mts., Darrow 305 (WIS) -
Siskiyou County: Mt. Shasta, Cooke 25701 (WIS) -
Colorado. Summit County-Clear Creek County: NE
of Loveland Pass, Shushan SL-1593 (US) - Idaho.
Owyhee County: Silver City, Anderegg 1437 (WIS) -
Montana. Glacier County: Glacier National Park,
Altyn Peak, Imshaug 5488 (CANL) - New Mexico.
Santa Fe County: Santa Fe, Egan EL-1652 (US, WIS)
- Oregon. Hood River County: Mt. Hood, Shushan
SL-2500 (WIS) - Washington. Clallam County:
Olympic National Park, Obstruction Peak. Brodo
14401 (CANL) - Pend Oreille County: Newport,
Schroeder L-305 (WIS) - Wyoming. Albany County:
Medicine Bow Mts., Cain 25736 (WIS).
2. Pseudephebe pubescens (L.) Choisy
Icon. Lieh. Univ., ser. 2, 1: sine pagin. (1930). -
Lichen pubescens L., Sp. PI. 2: 1 155 (1753); type: sine
loc.. herb. C. Linnaeus (LINN 1273.286 lectotypel).
- Alectoria pubescens (L.) R. Howe, Class. Fam.
Usneaceae: 23 (1912) et Bull. Torrey Bot. Club 39:
201 (1912).
Alectoria pacifica Stiz.. Proc. Calif. Acad. Sei. 5(2):
537 (1895); type: Mexico. Island of Guadalupe, E.
Palmer. 1875 (ZT holotype! US isotype!).
For further synonyms see Hawicsworth (1972 pp.
235-236).
Thallus decumbent to ascendent and shrubby;
branches generally very uneven in diameter but
more or less terete throughout their length,
with diameters near the base 0. 1-0.2 mm and at
the tips 0.05-0.1 mm; branching isotomic to
anisotomic dichotomous with short lateral
branches rare; internodal distance generally
long, (0.6—) 1— 3(— 5) mm; dark red-brown to black,
satiny to shiny.
True lateral spinules, soredia, and isidia
absent, although adventitious outgrowths similar
to those on P. minuscula are found on rare
occasions. Pseudocyphellae absent. Pycnidia
abundant and conspicuous, as in P. minuscula.
Apothecia infrequent, disc black, flat, 1.2-1. 6
mm diam.; margin thick, red-brown when
young, excluded when old; spores 8 per ascus,
broadly ellipsoid, c. 10 x 7.5 /am, hyaline.
All thallus tissues K-, C-, K.C — , PD-; no
lichen substances present.
Ecology. On dry, sandy, or stony ground and
acid rocks, especially igneous rocks, sometimes
shale; usually on exposed rocky ridges or
boulders soon free of snow in the spring. The
species tends to be found in moister sites in
regions having a continental climate (Weber in
litt.).
Distribution. Low to high arctic, subalpine to
alpine; frequent on exposed habitats along the
west coast and in the Districts of Keewatin
and Mackenzie (N.W.T.) as well as on the arctic
islands; many more occurrences in the humid
coastal ranges than the similar P. minuscula. It
ranges as far south as Guadalupe (type of
Alectoria pacifica). It is circumpolar in both the
Northern and Southern Hemispheres (Hawks-
worth 1972).
Discussion. See under/*, minuscula.
Selected specimens
Exsiccatae. Merrill: Lieh. Exs., Ser. I, 88: Washing-
ton, Spokane County: Crater Basin, Bosner (BM,
CANL, DUKE) - Merrill: Lieh. Exs., Ser. II, 135:
144 Alectoria in North America
OPERA BOT. 42 (1977)
Fig. 67. Pseudephebe puhescens . N. American distribution.
Washington, Klickitat County. Goldendale, Foster
(BM, DUKE, US) - Thomson: Lieh. Arct. 55:
N.W.T., District of Keewatin: Rossby Lake. J. W.
Thomson (CANL. DUKE, MSC) - Weber: Lieh.
Exs. 219: N.W.T., District of Mackenzie: Mackenzie
Mountains, O’Grady Lake, Cody 16816 (CANL,
DUKE).
Canada. British Columbia. Coastal District: W of
Hagensburg, Ohlsson 2289 (CANL) - Fraser River
Basin: E of Hazelton, Ohlsson 2973 (CANL) - Islands
North District: Moresby Island: Laing Point, Brodo
10795 (CANL) - Islands South District: Vancouver
Island: Mount Benson. Nanaimo, Macoun 49 (US) -
Saltspring Island, Brodo 13890 (CANL) - Skeena
River Basin: Terrace, Ohlsson 2798 (CANL) -
Alberta. Oldham River Watershed, Beauvais Lake,
Bird 14754 (CANL) - Quebec. Territoire du Nouveau
Quebec: Porvungnituk (sic) (Povungnituk), Thompson
30 (CANL) - Poste-de-la-Baleine. Brisson & Forest
20342 (CANL) - Newfoundland. Labrador North
District: Crater Lake vicinity, 52 miles WSW of
Hebron, Gillett 9415A (CANL) - Labrador South
District: Pack’s Harbour, Waghorne 25 (US) -
Yukon. Dezadeash Lake, Douglas 5355 (CANL) -
OPERA BOT. 42 (1977)
Pseudephebe Choisy 145
34 miles from West Dawson, Calder & Billard 3320
(CANL) - Northwest Territories. Mackenzie District:
NE corner of Great Bear Lake, Wisensel JW18B
(SASK1- District of Keewatin: Ennadai Lake. Brown
1327 (CANL) -Franklin District: Baffin Island, Dutch
Polar Station, Soper 152 (CANL. DUKE. NYBG).
U.S.A. Alaska. Alaska Range District: Denali
Highway, mile 28, Krog 3805 (O) - Aleutian Islands:
Unalaska, Captain's Bay, Eyerdam s.n., 1 May 1932
(FH) - Bering Strait District: Ogotoruk Creek
Drainage, Johnson, Viereck & Melchior 106 (ALA.
US) - Central Yukon River District: Steese Highway,
Eagle Summit, Krog 2546 (O) - Eastern Pacific
Coast District: Sitka, Harbour Mountain, Krog 6266
(O) - Western Pacific Coast District: Cold Bay,
55°12'N, 162°40'W, MacCartney s.n.. 6 Aug. 1971
(WIS) - California. Mariposa County: Yosemite
Valley, Herb. H. E. Hasse 571 (US) - Sierra County:
E of Yuba Pass, Tavares 621 (US) - Siskiyou County:
Trail to Sugar Lake, Salmon Mountains, Largent 4495
(HSC, WIS) -Colorado. Clear Creek County: Summit
Lake, Mount Evans. Sushan & Weber S I598A
(CANL, FH) - Montana. Ravalli County: St. Mary
Peak, Stevensville, Stickney 1559 (WIS) - Oregon.
Deschutes County: Lava Butte, near Bend, Pojar
s.n.. 8 Aug. 1970 (UBC) - Hood River County:
Mt. Hood, Foster 571 (FH) - Lake County: Near
Fort Rock, Horse Ranch, Pike L-106 (OSC) -
Washington. Okanogan County: Slate Peak, near
Winthrop, Imshaug 18568 (CANL) - Whatcom
County: Mt. Baker, Herre s.n., 14 Aug. 1914 (US) -
Wyoming. Yellowstone National Park: Shoshone
Lake. Coulter 3 (US).
Mexico. Baja California. Guadalupe Island, Palmer
s.n., 1875 (US, ZT).
Sulcaria Bystr
Ann. Univ. Mariae Curie-Sktodowska, C, 26: 275
(1971). - Alectoria sect. Sulcaria Mot., FI. Polska,
Porosty 5(2): 39 (1962); nom. inval. (Art. 37). -
Alectoria subsect. Sulcata DR., Ark. Bot. 20A (II):
5, 19 (1926). - Alectoria subgen. Sulcata (DR.)
D. Hawksw., J. Jap. Bot. 46: 335 (1971). - Alectoria
sect. Coelocaulopsis Choisy, Bull. Mens. Soc. Linn.
Lyon 24: 26 (1955); nom. inval. (Art. 36).
Type species: Sulcaria sulcata (Lév.) Bystr. ex
Brodo & D. Hawksw. (syn. Cornicularia sulcata
Lév.; holotype).
Thallus fruticose. erect and caespitose to decum-
bent or pendent; branching variable; branches
generally terete, but in one species bursting
open longitudinally and forming dorsiventrally
compressed and expanded flattened areas;
greyish-brown, badious, or emerald green.
Lateral spinules present consistently in some
species, rarer or absent in others. Isidia and
soralia absent. Pseudocyphellae present in the
form of a single longitudinal furrow (Fig. 5D),
± continuous from the base to the apex of the
thallus, deeply depressed, sometimes appearing
almost scalariform, white or emerald green,
sometimes bursting open to expose the medulla,
occasionally tending to become weakly sore-
diate. Cortex not disintegrating, not differen-
tiated and rough at the surface. Medullary
hyphae not ornamented.
Apothecia lateral, often appearing geniculate
owing to the death of the branch distal to them,
common in one species, unknown in the
others; excipulum thallinum concolourous with
the thallus, persistent and incurved, sometimes
becoming ciliate; disc brownish-black, often
distinctly blue-grey pruinose. Asci clavate,
thick-walled, arrested bitunicate, 6-8-spored.
Ascospores ellipsoid, yellowish to brown at
maturity, I- or later 3-septate when mature,
22-40 x 8-15 /am. Pycnidia unknown.
Usnic acids absent; atranorin abundant in two
species, chloroatranorin absent; /3-orcino! dep-
sidone present in two species; pulvinic acid
derivatives known in two species; orcinol
depsidone known in one specimen of one
species; orcinol depsides, tridepsides, /3-orcinol
depsides and hydroxyanthraquinones absent.
Discussion
Of the three species we are accepting in this
genus, only one (S. baclia) is known from North
America; the others (S. sulcata and S. virens)
are confined to southeast Asia. Only S. sulcata
is known fertile but this species is regularly so.
The distinctive pseudocyphellate furrow serves
to separate sterile members of this genus from
the other genera treated here. Two species
which we have placed in Bryoria (B. pseudoca-
pillaris and B. spiralifera ), and which are only
known sterile, may be allied to it. In fact, the
section Implexae of Bryoria seems to be closely
allied to the genus Sulcaria to judge by their
similarities in chemistry (the frequency of ß-
orcinol depsidones), colour (as in B. spiralifera
and the European B. kuemmerleana ), and long
pseudocyphellae.
It is of interest to note that, whereas atranorin
is usually present in only trace amounts in
Bryoria so that it is only detectable by TLC,
in two of the species of Sulcaria it is so abundant
that a yellow reaction with K is readily obtained
and the compound easily demonstrated by
microcrystal tests.
I . Sulcaria badia Brodo & D. Hawksw., sp. nov.
Thallus pendulus, 20-50 cm longus, flaccidus; rami
praecipue iso-dichotomiter fastigiati, frequentes e
basi emergentes, in angulos praecipue acutos rotundos
divergentes, valde compressi et toiluosi, conspicue
sulcati, diametro aequales, basi 0.25— 0.4(— 1 .0) mm
diam., ramis lateralibus brevibus et gracilibus fere
perpendiculariter frequenter instruct]'; tota planta
OPERA BOT. 42 (1977)
Sulcaria Bystr. 147
opaca castanea vel badia vel partim flavo-brunnea.
Rami laterales spiniformes veri, isidia, et soralia
desunt. Pseudocyphellae conspicuae. albae. lineatae,
longissimae, in sulcos profundos immersae.
Apothecia et pycnidia ignota.
Cortex K+ flavescens, C-. KC+ flavescens,
PD+ brunneolus; medulla K-. C-, KC-. PD ; in
Charta separata K+ flavescens et PD+ flavescens
reagens. Thallus solum atranorium continens.
Holotypus: America septentrionalis, U.S.A.,
Oregon, Philomath, in Malis veteribus, F. P. Sipe
669 p.p. (ORE). Fig. 68.
The holotype is a portion of the same number
designated by Gyelnik as the holotype of B. pseudo-
fuscescens (Gyeln.) Brodo & D. Hawksw. in BP (i.e.,
an arithmotype). The specimen in BP is, however,
quite distinct in both morphology and chemistry from
that in ORE. See also the discussion under B.
pseudof usees cens .
Thallus pendent, 20-50 cm long, flaccid;
branching mainly isotomic dichotomous,
frequent from the base, angles between the
dichotomies mainly acute and rounded; branches
markedly flattened and twisted, conspicuously
sulcate, even, 0.25— 0.4(— 1 .0) mm diam. at the
base, with short, slender almost perpendicular
lateral branches; very dull chestnut-brown to
almost badious or yellowish brown in parts.
True lateral spinules, isidia and soralia absent.
Pseudocyphellae conspicuous, white, linear,
extremely long, most developing into deep
furrows (Fig. 5D).
Apothecia and pycnidia unknown.
Cortex K+ yellow, C— , KC+ yellow, PD +
brownish; medulla K — , C— , KC— , PD — ; on
filter paper K+ yellow and PD+ yellow. Con-
tains atranorin alone. A specimen from Cali-
fornia (Pike & Rhoades 2245 [OSU]) contains,
in addition, an unidentified PD- substance
having relative Rf classes (C. Culberson 1972)
of 5-5-5.
Ecology. On trees, especially apple and oak
trees, in well-lighted Quercus garryana com-
munities.
Distribution. Along the west coast from
Washington to Northern California (Fig. 69).
Endemic to North America.
Discussion
This previously undescribed species is easily
recognized by its very distinctive colour. The
colour of the thallus is not unlike that ot the
Asian S. sulcata which is almost always fertile,
Fig. 68. Sulcaria badia. Oregon, Hall, 1871 (FH-
Tuck. 504). Scale 0.5 mm.
caespitose, and contains psoromic acid in addi-
tion to atranorin.
A morphologically typical specimen of Sulcaria sul-
cata var. sulcata from China (Krypt. Exs. Vindobon.,
No. 2766, Handel-Mazzetti. prov. Yiinnan [CANL])
was found to contain virensic acid instead of psoromic
acid with TLC and microchemical tests (Hawksworth
1971). The frequency of this chemical strain is
unknown at present, it is intriguing that the locality
of this specimen is the same as the type locality of
S. sulcata f. vidpinoides, a form differing from f.
sulcata in being yellow-green in colour, due to vulpinic
acid (Hawksworth 1971).
148 Alectoria in North America
OPERA BOT. 42 (1977)
Large amounts of atranorin occur in this species:
a K+ yellow reaction is easily obtained, and this
compound can be readily confirmed by micro-
crystal tests in GAoT.
We are referring this species to the genus
Sulcaria because of the characteristic sulci,
although this feature is perhaps less well devel-
oped in S. badia than it is in either S. sulcata
or S. virens. In the absence of apothecia, the
placing of this North American species in the
genus Sulcaria should be regarded as tenta-
tive.
Specimens
U.S.A. California. Mendocino County: Round Valley,
Chesnut s.n., 20 July-3 Aug. 1897 (US) - 2 miles W
of Willits, Pike & Rhoades 2245 (OSC) - Oregon.
Benton County: near Philomath, Sipe s.n., no date
(ORE, F) - Philomath, Sipe 669 (CANL) - Locality
unknown: Hall s.n., no date (FH-Tuck., US) -
Washington. Clallam County: nearDungeness, Foster
2552A (FH).
Excluded taxa
Alectoria acharianaGyeln.
Nyt Mag. Naturvid. 70: 54 (1932).
This species was reported by Gyelnik (1931)
prior to the valid publication of the name from
“Canada, Lake Louise, alt. 5665 ft. (SE1GFRID,
sub Alectoria Fremontii. A 1-9)“. Gyelnik,
however, later reconsidered his determination
and treated this collection as the holotype of
Bryopogon negativus f. canadensis Gyeln.,
which is discussed separately below.
The identity of A. achariana itself remains
uncertain in the absence of the holotype collec-
tion which is not in BP (Verseghy in litt.).
Hawksworth (1972 p. 219) noted that an iso-
paratype of this taxon in BM was Bryoria
fuscescens var. positiva.
Alectoria californica (Tuck.) Merrill
Lieh. Exs. no. 82 ( 1908). - Cetraria californica Tuck.,
Am. J. Arts Sei., ser. 2, 28: 203 (1859).
Although this taxon shows a strong resemblance
to some morphotypes of Bryoria abbreviate!,
it is more appropriately placed in the genus
Cornicularia (as C. californica (Tuck.) DR.)
on the basis of the anatomy of the cortex (see
general section Anatomy and morphology:
Cortex). The characters most useful for the
separation of these two species are discussed
under B. abbreviata.
Alectoria cetrariza Nyl.
Bull. Soc. Linn. Normand., sér. 4, 1: 270 (1887);
type: U.S.A., Oregon, Millamak, on Pinas contorta,
Herb. Nylander no. 35973 (H holotype!). — Coeloc 'an-
ion cetrariza (Nyl.) Gyeln., Ann. Mus. Nat. Hungar..
Bot. 28: 282 (1934).
An examination of the holotype of this taxon
showed it to be conspecific with Cornicularia
californica.
Alectoria crinalis Ach.
Lieh. Univ.: 594(1810).
This epithet was taken up by Hale (1969 p.
191) for the species treated under the name
Ramalina crinalis (Ach.) Gyeln. by Motyka
(1964). As pointed out below under Alectoria
tlirausta, R. thrausta (Ach.) Nyl. is the nomen-
claturally correct combination for this species.
Although we have not seen the material under
this name in Acharius' main herbarium in H.
authentic material of it in BM (!) is R. thrausta.
Alectoria divergens (Ach.) Nyl.
Mém. Soc. Imp. Sei. Nat. Cherb. 3: 171 (1855); et
Bot. Notiser 1855: 136 (1855). - Cornicularia diver-
gens Ach., Meth. Lieh.: 305 (1803); type: Sweden.
Lapponia. Herb. Acharius (H lectotype!): basionym.
This species was accepted in Alectoria by Howe
(191 1), but on the basis of its cortical structure
must be placed in Cornicularia at the present
time (Hawksworth 1972). The separation of C.
divergens from Bryoria nitidula, a species with
which it has sometimes been confused, is
discussed under the latter species.
Cornicularia fibrillosa (Ach.) Halsey
Ann. Lyceum Nat. Hist. N. Y. 1: 20 (1823). -Corni-
cularia puhescens y. fibrillosa Ach.. Syn. Lieh.: 303
(1814); type: America] borfealis], Muhlenberg (H-Ach
holotype); basionym.
This name was applied by Halsey (1823) to a
species "Common on fences" in the vicinity
of New York. He clearly based his name on
1 0 - Opera Botnnica nr 42
150 Alectoria in North America
OPERA BOT. 42 (1977)
that of Acharius (1814) and intended to treat it
as distinct species (“Certainly a distinct
species”). Acharius, however, gave the habitat
as “ad saxa" and an examination of the only
specimen under this name in his main herbarium
(presumed to be the holotype) by Dr. T. Ahti (in
litt.) revealed that this was a member of the
genus Ephehe Fr. Its small size renders a deter-
mination to the species level difficult. The
epithet fibrillosa was not mentioned by Henssen
(1963). From Flalsey’s description and com-
ments there can be little doubt that he misap-
plied the Acharian name to Bryoria furcellata.
A specimen under this name from Tuckerman’s
herbarium (New England, "ad septimenta
lignea, vulg., sterilis” [BM]) is also B. furcellata.
It should also be pointed out that Zahlbruck-
ner (1930 p. 604) erroneously attributed the
name Cornicularia fibrillosa to Eaton (1829 p.
188); Eaton did employ this name and even
provided a description at least five years earlier
(Eaton 1824 p. 273; editions of this work prior
to the fourth have not been seen by us). Since
Eaton did not cite the names of authors of
lichen taxa in the fourth edition, and as C.
fibrillosa was listed after C. pubescens and
stated to occur “on rocks”, it seems that he was
simply following Acharius but adopting specific
rank as proposed by Flalsey in the previous year.
Bryoria intricans (Vain.) Brodo & D. Hawksw.,
comb. nov.
Alectoria chalybeiformis f. intricans Vain., Terme-
szetr. Fuzetek 22: 276 (1899); type: Caucasus, in rupe
ad Adisch, 1885. H. Lojka, Herb. Vainio no. 01010
(TUR no. 001244 holotype!); basionym. - Alectoria
intricans (Vain.) Mot., Fragm. Florist. Geobot. 3:
228 (1958).
Gyelnik (1931) reported this species from Idaho
on the basis of a collection from the “Upper
Priest River, on Pinas (STILLING ER, sub
Alectoria jubata, sorediata. AI. -II)" which we
have not been able to locate. Motyka later
tentatively referred one of Krog's Alaskan
collections to this taxon but Krog (1968) did
not accept his determination; in our view,
Krog's specimen is a slender plant of B. chaly-
beiformis .
The holotype collection of this taxon is poorly
developed but there seems little doubt that the
interpretation of it adopted by Krog (1971), and
Dahl & Krog (1973), is correct. We have not
seen any North American material definitely
referrable to this taxon although young thalli
and fragmentary specimens of B. chalybeiformis
resemble it in part but differ in being prostrate
to decumbent rather than pendent. It is possible
that B. intricans is a saxicolous morphotype of
B. lanestris and it is of interest to note in this
respect that Motyka (1958 a) applied the epithet
lanestris to a saxicolous taxon apparently iden-
tical to B. intricans.
Alectoria jubata (L.) Ach.
Lieh. Univ.: 592 (1810). - Lichen jubatus L., Sp.
Plant. 2: 1155 (1753); type: sine loc., Herb. Linnaeus
no. 1273.281 p.p. (LINN lectotype!).
The name Alectoria jubata has been applied by
both North American and European lichenol-
gists at various times to numerous pendent
species of Bryoria and has tended to be used as
a "blanket" name for this taxonomically diffi-
cult group. As the name has been taken up in
different senses, its rejection under Art. 69 now
appears to be justifiable.
The lectotype specimen is extremely squashed
and is in a poor state of preservation. It
may either correspond to the concept of "A.
prolixa " (an illegitimate name; see below)
adopted by Motyka (1958 a), as pointed out by
Hawksworth (1970), or even a very poor and
maltreated specimen of A. fremontii. “A.
prolixa", as interpreted by Motyka (1958a),
appears to be a taxon restricted to eastern
Europe and of dubious taxonomic validity. In
view of the state of the type specimen and
the uncertainty surrounding it, and since the
name has been used in different senses, it seems
preferrable to reject this name altogether as a
nomen confusum.
Alectoria jubata var. philadelphiae Gyeln.
Nyt Mag. Naturvid. 70: 58 (1932). - Bryopogon
jubatus var. philadelphiae (Gyeln.) Gyeln.. Feddes
Repert. 38: 225 (1935). - Bryopogon fuscescens var.
philadelphiae (Gyeln.) Gyeln., Ann. Mus. Nat.
Hungar.. Bot. 32: 153 (1939).
This variety was described on the basis of a
specimen from British Columbia (Selkirk. Shaw
977) but this collection has not been located in
BP (Verseghy in litt.). Gyelnik’s diagnosis of
OPERA BOT. 42 (1977)
Excluded taxa 151
this taxon was “ Thallus pendulus, opacus,
levis, 4-8 cm longus, olivaceofuscus, sorediis
lateralibus maculiformibus, fuscocaesiis,
granuloso-pulverulentis instructus" . From
Gyelnik (1935) it appears that the taxon must
have been K-. KC- and C-. While these data
suggest that this variety may well belong to B.
fuscescens, Räsänen. who corresponded with
and sent material to Gyelnik. determined some
North American collections of#, pseudofusces-
cens in H as var. philadelphiae. The true ap-
plication of this epithet must consequently re-
main in doubt until the holotype collection is
found.
Alectoria loxensis (Fée) Nyl.
Mém. Soc. Imp. Sei. Nat. Cherb. 5: 98 (1857). -
Cornicutaria loxensis Fée, Essai Crypt.: 137 (1825):
lectotype: loc. cit. PI. 31 fig. 7.
This species is most satisfactorily accom-
modated in the genus Oropogon Th. Fr., of
which it is the holotype species (i.e., as O.
loxensis (Fée) Th. Fr.). Oropogon differs from
the other Alectorioid genera in having single-
spored asci, massive brown submuriform
ascospores, "cyphella-like" cortical perfora-
tions, and a characteristic chemistry. Consider-
able morphological and chemical variation
occurs in what has been called O. loxensis and,
as pointed out by Jorgensen (1972), the group
is in need of a thorough taxonomic revision.
O. loxensis s. lat. has not been correctly report-
ed from either Canada or the United States but
is very common on trees and rocks in the moist
mountains of Central and South America. The
thalli of O. loxensis are chestnut brown and
shiny to very pale whitish-grey (var. atroalbi-
cans (Nyl.) DR.; see Dodge & Vareschi 1956)
and are very frequently referred to Alectoria
s. lat. by collectors. Sterile plants of O. loxensis
are, however, easily separated from similar
Bryoria and Sulcaria species by the conspi-
cuous fusiform “cyphella-like"' perforations in
the cortex (illustrated by Hawksworth 1972).
The world distribution of Oropogon was dis-
cussed and mapped by Sato (1967).
Bryopogon negativus f. canadensis Gyeln.
Acta Geobot. Hungar. 2: 165 (1937). - Bryopogon
lanestris f. canadensis Gyeln.. Acta Geobot. Hungar.
2: 165 (1937); nom. inval. (Art. 34).
This form was described on the basis of a collec-
tion from Lake Louise (near Banff, Alberta)
which Gyelnik (1932) had determined as Alec-
toria achariana (see this name). Gyelnik's origi-
nal diagnosis of this form was “Similis f.
barbato Gyeln. sed rami primarii thalli crassi
( usque ad 1 mm), minus contorti, angulosi,
irrégularité r lacunosi. Thallus K— , C— , KC — ,
P— ”. Although Gyelnik indicated that the type
material was preserved in BP. it cannot be
located there now (Verseghy in litt.), and so the
identity of this taxon must remain in doubt.
Alectoria prolixa (Ach.) Nyl.
Medd. Soc. Fauna FI. Fenn. 2: 14 (1878). - Alectoria
jubata a. prolixa Ach., Lieh. Univ.: 592 (1810).
The name Alectoria prolixa, which was applied
to North American collections by several early
authors (e.g., Nylander 1885, Hutting 1896.
Macoun 1902), was used by Motyka (1958 a,
1962) for a species he considered to be essen-
tially eastern European. As pointed out by
Hawksworth (1970), however, a number of
nomenclatural difficulties surround this name. It
can be treated as (a) illegitimate under Art.
63 as it was a superfluous name for [var.]
jubata, and (b) a name to be rejected under
Art. 70 as based on discordant elements. In
fact, as Art. 63 applies, this epithet is automa-
tically typified by the type of [var.] jubata
under the provisions of Art. 7. Thus the name
“A. prolixa ” is an obligate synonym of A.
jubata, a name which itself is best rejected (see
this name). As is the case with A. jubata, this
name has been applied to diverse dark-coloured
species of the genus by North American authors.
Alectoria sarmentosa var. dichotoma (Hoffm.)
Gyeln.
Ann. Mus. Nat. Hung., Bot. 28: 283 (1934). - Vsnea
dichotoma Hoffm., Deutsch. FI. 2: 134 (1796); type:
Sine loc.. Herb. Hoffmann no 8573 (MW neotype!).
This taxon was reported from North America by
Gyelnik (1934) but, as pointed out by Hawks-
worth (1970 p. 240, 1972 p. 239), Hoffmann's
only extant collection of this proves to be
Ramalina thrausta (Ach.) Nyl. In the absence of
further evidence, this epithet is thus most
appropriately treated as a synonym of R.
thrausta. This does not affect the nomenclature
152 Alectoria in North America
OPERA BOT. 42 (1977)
of/?, thrausta as there is already a R. dichotoma
Hepp ex Zoll, validly published in 1854 (fide
Zahlbruckner 1930 p. 524). We have not seen the
material referred to this taxon by Gyelnik, so
the sense in which he used the name remains
uncertain.
Bryoria setacea (Ach.) Brodo & D. Hawksw.,
comb. nov.
Alectoria jubata ft. setacea Ach., Lieh. Univ.: 594
(1810); type: sine loc. (H-Ach., upper left-hand
specimen, lectotype!); basionym. - Alectoria setacea
(Ach.) Mot., Fragm. Florist. Geobot. 3: 216 (1958).
This species was recorded for North America
by both Ahti (1964) and Krog (1968) on the
basis of collections determined by Motyka which
we have reexamined and consider to be con-
specific with Bryoria capillaris. Motyka (1964),
however, indicated that this species did not
occur in North America and so appears to have
reconsidered these determinations himself. As
pointed out by Hawksworth (1973), B. setacea
seems to represent a “vexitlifera"-\ike morpho-
type of B. capillaris with robust main stems
which are often markedly foveolate and exceed
1 mm in diameter, sparse branching, and a
prostrate habit. The true B. setacea appears to
be an exclusively European species most fre-
quent in upland areas of Scandinavia, although
it is occasionally encountered in the moun-
tainous regions of central Europe.
Motyka has tended to use the name in a
rather broad sense applying it also to large
pendent specimens of B. capillaris in which the
basal branches are unusually stout, tending to
become somewhat foveolate. Such a difference
appears to be mainly environmentally controlled
and thus of no taxonomic importance. Collec-
tions from North America falling within this
broadened concept of B. setacea have been
found during the present studies, and the follow-
ing may be cited as representative (all on trees
and treated as belonging to B. capillaris):
British Columbia, Victoria, 1908, Macoun
(CAN L 16194): Ontario, Geraldon District,
1958, Ahti 3927, 3932 (H); Quebec, Gaspé
Peninsula, 1907, Macoun (CANL 16181). In
addition we examined one specimen from an
uncertain locality (“on rocks by the Rim”),
collected by Merrill (FH) and containing alec-
torialic and barbatolic acids together with
atranorin (?) and some unknown compounds,
but this is perhaps merely an abnormal specimen
of 5. nadvornikiana.
Bryoria smithii (DR.) Brodo & D. Hawksw.,
comb. nov.
Alectoria smithii DR., Ark. Bot. 20A (11): 15 (1926);
basionym.
Alectoria bicolor var. berengeriana Massai, ex
Stiz. , Ann. Naturh. Mus. Wien 7: 127 (1892).
For further synonyms and typifications see Hawks-
worth (1972 p. 246).
This species was reported for North America
by Stizenberger (1892) on the basis of a collec-
tion made in Newfoundland (ZT!) which has been
redetermined as B. fuscescens var. positiva
(Jorgensen & Ryvarden 1970, Ahti & Hawks-
worth 1974). Keissler (1958 p. 151) also con-
sidered this species to occur in North America
on the basis of material distributed by Merrill,
Lieh. Exs. no. 211 (sub Alectoria chalybeifor-
mis), but, as pointed out by Jorgensen & Ryvar-
den (1970), material under this number proves
to be B. furcellata.
The distribution of B. smithii is centered in
Asia, but extends through the moister parts of
Europe and onto the Pacific Islands. Although
we have seen no North American material
of this distinctive species, it is conceivable that
it may occur at least near the east or west
coasts.
B. smithii is perhaps most similar to B. bicolor
from which it differs in the presence of minute
soralia bearing tufts of isidioid spinules (not
unlike those seen in B. furcellata), the complete
absence of fumarprotocetraric acid (i.e., PD —
throughout), and a decumbent rather than ±
erect habit. For further information on this
species see Jorgensen & Ryvarden (1970) and
Hawksworth (1972).
Bryopogon spadiceus Choisy
Icon. Lieh. Univ., Ser. II, Fasc. 2, sine pagin. (1931).
This is a superfluous name for Alectoria jubata
(L.) Ach., as that epithet was placed as a
synonym of B. spadiceus by Choisy (1931).
The latter is therefore typified by the type of
Lichen jubatus L. (see under Alectoria jubata),
and so is illegitimate and must be rejected (Art.
63).
OPERA BOT. 42 (1977)
Excluded taxa 153
The name Bryopogon spadiceus was applied
by Choisy (1931) to two North American
collections: one from Alberta, Cypress Hills,
J. Macoun, 1880 (Macoun, Lieh. Exs. no. 19
[herb. Choisy. herb. GandogerJ) and one from
the Bering Sea, St. Paul Island, J. (M.'?)Macoun,
June-July 1897 [herb. Gandoger], As Macoun's
exsiccata no. 19 consists of material from several
localities and the St. Paul Island material has
not been located, the identity of these speci-
mens remains uncertain.
Bryoria subdivergens (Dahl) Brodo & D.
Hawksw., comb. nov.
Alectoria subdivergens Dahl. Medd. Gronl. 150(2):
145 (1950): type: Greenland, Julianehaab District.
Quagssimuit, E. Dahl, 4 July 1937 (O holotype! O
isotype!); basionym. - Bryopogon subdivergens (Dahl)
Bystr., Ann. Univ. Mariae Curie-Sklodowska, C.
26: 271 (1971); nom. inval. (Art. 33).
Thallus subcaespitose to decumbent or trailing,
to 25-30 mm long; branching anisotomic dicho-
tomous, angles between the branches mainly
obtuse; main branches longitudinally foveolate
and pitted, angular in cross section, mainly even
in diameter, to 0.8 mm diam. (1-2 mm diam.
fide Dahl 1950); red-brown throughout, matt.
True lateral spinules, soralia and pseudocy-
phellae absent; branches sometimes appearing
spinulose due to the occurrence of short,
pointed, lateral branches.
Apothecia infrequent, lateral; excipulum
thallinum concolourous with the thallus, be-
coming excluded as the ascocarps mature,
eciliate; disc red-brown becoming markedly
convex with age, 1.5-2. 5 mm diam. Ascospores
8 per ascus, subglobose, thin-walled, hyaline.
6-8 /am diam. Pycnidia not seen.
Cortical hyphae as in B. oregana.
Cortex and medulla K — , C— , KC— , PD-;
contains no lichen substances.
Ecology. In its type locality B. subdivergens
occurred growing amongst mosses and lichens
in a tundra lichen-heath.
Distribution. This species was reported from
four additional sites in Greenland and one in
Labrador by Dahl (1950). Dahl considered the
species as easily mistaken for Cornicularia
divergeas but this appears to have been due to
his referring material of the latter to B. sub-
divergens. All Greenland material apart from
that from the type locality proves to be C.
divergeas. The report from Labrador was based
on material collected by Tanner at Hebron but
this collection has not been located in H. O,
S or UPS. B. subdivergens was also recorded
for North America by Thomson (1960) on the
basis of a collection from Baffin Island (Pang-
nirtung Fjord, Soper 74 [CAN L]) which has been
redetermined as Cornicularia divergeas. Motyka
(1964) stated that he had seen a collection of
this species from Cornwallis Island (Resolute
Bay, Collins [ K]). but this specimen has not been
located amongst the lichen collections transfer-
red to BM from K. We have seen some unusually
dark specimens of Alectoria nigricans collected
by Collins at Resolute Bay (in herb. US), and
these may be duplicates of the specimen seen
by Motyka at Kew.
This species is thus only known with certainty
from the type locality but should be searched
for elsewhere in Greenland and in arctic North
America.
Discussion
Bryoria subdivergens is extremely similar to B.
abbreviata, a corticolous western species, and
were it not for their very disjunct localities and
different habitats we would have been inclined
to treat them as conspecific. The type material of
B. subdivergens, upon which the above descrip-
tion is based, is rather sparse and additional
collections of this species are urgently required.
The similarity of the cortices in these two
species. B. oregana. and probably also B. diver-
gescens is discussed in the general section
Anatomy and morphology.
Alectoria thrausta Ach.
Lieh. Univ.: 596 (1810); type: Helvetia (H-Ach.
lectotype!).
This species, which is often confused with A.
sarmentosa, has been treated as belonging to the
genus Ramalina (R. tlirausta (Ach.) Nyi.) by
most recent authors. Its retention in Ramalina
is justified, in the absence of apothecia, by (a)
the occurrence of apical soralia, (b) the presence
of only (-l-)-usnic acid, (c) the presence of occa-
sional net-like tips (as in young R. menziesii),
and (d) the presence of a partially prosoplec-
154 Alectoria in North America
OPERA BOT. 42 (1977)
tenchymatous cortex (see general section on
Anatomy and morphology). Interestingly, the
very similar Ramalina chondrina Steiner, a
species described from the Canary Islands,
produces apothecia with the I -septate hyaline
ascospores characteristic of that genus.
The most useful characters for the separation
of R. thrausta from Alectoria sarmentosa are
its generally more delicate habit, the infre-
quency of convex pseudocyphellae, its tendency
to assume a characteristic orange-yellow colour
in the herbarium, the presence of minute
apical soralia on short and often hook-like
branchlets, and the consistently C-, K.C-,
PD— medulla (but note that some chemotypes of
A. sarmentosa have these reactions).
Motyka (1960, 1962. 1964) adopted the name
R. crinalis (Ach.)Gyeln. (syn. A. crinalis Ach.)
for this species but. as pointed out by Ahti
(1964), this is nomenclaturally incorrect; the
name R. thrausta is the one which must be
used for this taxon.
Alectoria trichodes Bach. Pyl.
Mém. Soc. Linn. Paris 4: 432. 505 (1826).
This name was applied by Bachelot de la Pylaie
(1826) to some pendent Bryoria he saw in New-
foundland. No description was provided and
there was no indication that he intended to
introduce a new combination based on
Michaux ’s name (see Discussion under Bryoria
trichodes subsp. trichodes ), so this name was
not validly published by him (Arts. 32-34).
In a resumé of Bachelot de la Pylaie's paper,
Arnold (1826 p. 128) also referred to this name
but failed to validate it either as a new species
or a new combination. The nomenclatural
situation here is comparable to that of Du
Rietz’s use of the name “ Haematomma ochro-
leucum" discussed by Laundon (1970).
Some of Bachelot de la Pylaie’s material is
now preserved in FH and there is one specimen
from “E. Newfoundland” under the name
“Setaria trichodes Michx." mounted on a sheet
including a collection made by Despréaux, pro-
bably in 1828, which might have been collected
by Bachelot de la Pylaie; this belongs to
Bryoria trichodes subsp. americana (see Ahti
& Hawks worth 1974).
Alectoria tristis (Web.)Th. Fr.
Lieh. Scand. 1: 28 (1871). - Lichen tristis Web.,
Spicil. Fl.Goett.: 209(1778).
This name has been treated as a synonym of
Cornicularia normoerica (Gunn) DR. by most
authors since Du Rietz (1926). This species is
clearly not an Alectoria on the basis of its ana-
tomical structure. It does occupy a very iso-
lated position within the genus Cornicularia
(Schreb.) Ach. which, in any case, seems to
be a heterogeneous group.
Alectoria usneoides (Ach.) Ach.
Lieh. Univ.: 594 (1810). - Parmelia usneoides Ach.,
Meth. Lieh.: 270 (1803).
Parmelia usneoides is a superfluous name intro-
duced by Acharius for Lichen usnea L., the
basionym of Ramalina usnea (L.) R. Howe,
which is a true species of Ramalina with
uniseptate ascospores. The typification of L.
usnea is discussed by Imshaug (1972).
Sulcaria virens (Tayl.) Bystr. ex Brodo & D.
Hawskw., comb. nov.
Alectoria virens Tayl., Hook. Lond. J. Bot. 6: 188
(1847); type: India, Sheopore, Wallich, January 1821
(FH holotype! BM isotype!); basionym. - Sulcaria
virens (Tayl.) Bystr., Ann. Univ. Mariae Curie-
Sktodowska. C, 26: 276 (1971); nom. inval. (Art. 33).
This name was incorrectly taken up by Howe
(1911) for A. tortuosa Merr. ( =Bryoria ), and he
has been followed by several later North
American authors. Sulcaria virens , however, is
a very distinctive species which is restricted to
the Himalayan region (including parts of India,
Nepal and China) and Taiwan. Bryoria tortuosa
and Sulcaria virens are perhaps most easily
separated on the basis of the single longitudinal
furrow which characterizes all branches of the
latter, but they also differ in several other
respects (see Hawksworth 1971).
New combinations for non-North American and
some infraspecific taxa
Bryoria asiatica (DR.) Brodo & D. Hawksw., comb,
nov. - Alectoria asiatica DR., Ark. Bot. 20A (11):
18 (1926); type: China, Prov. Sze-ch'uan, reg. bor.-
occid., mellan Tsagogamba och Tamba, alt. 4000
m.s.m., p à Juniperus, Picea eher Rhododendron, H.
Smith 5018, 2 October 1922 (UPS holotype!);
basionym.
Bryoria chalybeiformis f. prostratosteola (Gyeln.)
D. Hawksw., comb. nov. - Alectoria prostratosteola
Gyeln.. Nyt Mag. Naturvid. 70: 58 (1932); type:
U. S.S.R., Kasachskaja S.S.R.. Altanigeb rupe.
Koton. Karagny. P. Kwaszmikowa. 1926 (BP 48575
lectotypel); basionym. - For further synonyms see
Hawksworth (1972 p. 215).
Bryoria confusa (Awasthi) Brodo & D. Hawksw.,
comb. nov. — Alectoria confusa Awasthi, Proc.
Indian Acad. Sei., B. 72: 152 (1970): type: E. Nepal.
Mewakhola valley, alt. c. 11,500 ft, D. D. Awasthi
2278, 28 May 1953 (herb. Awasthi. holotype; not
seen); basionym.
Bryoria cornicularioides (P. Jorg.) Brodo & D.
Hawksw., comb. nov. - Alectoria cornicularioides
P. Jorg., Bryologist 78: 77 (1975); type: China. Prov.
Shensi Kuan-tou-san, among mosses, G. Giraldi.
1896 (FI holotype!).
Bryoria dahlii (P. Jorg.) Brodo & D. Hawksw.,
comb. nov. - Alectoria dahlii P. Jorg., Svensk Bot.
Tidskr. 66: 192 (1972); type: New Guinea. Chimbu
District, Mt. Wilhelm. Pinaunde Valley, E. Dahl,
1970 (BM isotype!).
Bryoria divergescens (Nyl.) Brodo & D. Hawksw..
comb. nov. - Alectoria divergescens Nyl., Flora,
Jena 69: 466 (1886); type: China, Prov. Yunnan, in
monte Tsang-chan, supra Ta-li, supra ramulos, alt.
4,000 m, R. P. Delavay, 1885 (H-Nyl. 35972 holo-
type!): basionym. - For further synonyms see Hawks-
worth (1970 p. 237).
Bryoria fuscescens f. pallida (Saviez) D. Hawksw.,
comb. nov. - Bryopogon chalybeiform f. pallidum
Saviez, Not. Syst. Inst. Crypt. Horti Petrop. 1: 64
(1922); type: Poland, Prov. Suwalki, distr. Augustov,
V. P. Saviez, 9 May 1914 (LE holotype!); basionym. -
For further synonyms see Hawksworth (1972 p. 218).
Bryoria fuscescens var. positiva (Gyeln.) Brodo &
D. Hawksw., comb. nov. - Bryopogon positivus
Gyeln., Acta Geobot. Hung. 2: 164 (1937): type:
U.S.S.R., Siberia, Altaigebirge, Dorf Tscherga, N.
Lawrow, 18 June 1927 (BP 33.974 holotype!);
basionym. - For further synonyms see Hawksworth
(1972 p. 219).
Bryoria himalayensis (Mot.) Brodo & D. Hawksw.,
comb. nov. - Alectoria himalayensis Mot., Fragm.
Florist. Geobot. 6: 450 (I960); type: Himalaya,
Sikkim, T. Thomson 299 (BM isotype!); basionym.
Bryoria indonésien (P. Jorg.) Brodo & D. Hawksw.,
comb. nov. - Alectoria indonésien P. Jorg., Svensk
Bot. Tidskr. 66: 194 (1972): type: Indonesia, Celebes,
south-west slope of Mt. Rautemario, on large Quercus,
alt. 3,100 m, A. C. Jermy 7448, 1969 (BM holotype!);
basionym.
Bryoria kuemmerleana (Gyeln.) Brodo & D. Hawksw.,
comb. nov. - Alectoria kuemmerleana Gyeln., Nyt
Mag. Naturvid. 70: 49 (1932); type: Hungary. Comit.
Szepes, Fenyveserdd alomjdn a “Stösschen" lejtôjén
a M. Tâtrâban, c. 1380 m, Timkô Györgye, 17 July
1917 (BP 33.952 holotype!); basionym.
Bryoria lactinea (Nyl.) Brodo & D. Hawksw.. comb,
nov. - Alectoria lactinea Nyl.. Lieh. Jap.: 23 (1890);
type: Japonia, ltjigome. E. Almqvist, 1879 (H-Nyl.
35882 holotype!); basionym.
Bryoria motykae (D. Hawksw.) Brodo & D. Hawksw..
comb. nov. - Alectoria motykae D. Hawksw., Bot.
Notiser 124: 124 (1971); type: Kenya. Mt. Kenya,
Teleki Valley, on vertical surface of boulders, alt,
4,200 m. O. Hedberg 1720b, 28 July 1948 (UPS
holotype!); basionym.
Bryoria osteola (Gyeln.) Brodo & D. Hawksw., comb,
nov. - Alectoria osteola Gyeln., Nyt Mag. Naturvid.
70: 49 (1932); type: Hungary, Comit. Gömör, in monte
Kiralyhegy, Filarszky & Kümmerte, 3 July 1906
(BP 33.955 holotype!); basionym.
Bryoria perspinosa (Bystr.) Brodo & D. Hawksw..
comb. nov. - Alectoria perspinosa Bystr., Khumbu
Himal. 6: 21 (1969): type: E. Nepal. Vorhimalaya,
Abies-Rhododendron-BergwM, östlich Junbesi, J.
Poelt L778. 9 October 1962 (M isotype!); basionym.
Bryoria poeltii (Bystr.) Brodo & D. Hawksw., comb,
nov. - Alectoria poeltii Bystr.. Khumbu Himal. 6:
20 (1969); type: E. Nepal, Himalaya, Mahalangur
Himal, Khumbu, bei Bibre, J. Poelt L805, September
1962 (M isotype!); basionym.
156 Alectoria in North America
OPERA BOT. 42 (1977)
Bryoria ruwenzoriensis (D. Hawksw.) Brodo & D.
Hawksw., comb. nov. - Alectoria ruwenzoriensis
D. Hawksw., Bot. Notiser 124: 124 (1971); type:
Uganda, Ruwenzori. Mijusi Valley, alt. 3,900 m, O.
Hedberg 52 1 f, 27 March 1948 (UPS holotype!);
basionym.
Bryoria variabilis (Bystr.) Brodo & D. Hawksw.,
comb. nov. - Alectoria variabilis Bystr., Khumbu
Himal. 6: 22 (1969); type: E. Nepal, Himalaya,
Khumbu, Abies-Rhododendron-'Na\A, J. Poelt L798,
9 October 1962 (M isotype!); basionym.
Pseudephebe pubescens f. subciliata (Nyl.) D.
Hawksw., comb. nov. - Parmelia lanata var. suh-
ciliata Nyl., Flora, Jena 51: 346 (1868); type: Scotland,
ad rupes calcareosus monti Morrone in regione
Braemar, J. M. Crombie, 1867 (H-Nyl. 34350 holo-
type!; BM isotype!): basionym. -For further synonyms
see Hawksworth (1972 p. 236).
Sulcaria sulcata (Lév.) Bystr. ex Brodo & D.
Hawksw., comb. nov. —Cornicularia sulcata Lév., in
Jacquin, Fr.-Voy. Inde, Descr. Coll. 4: 179 (1844);
type: loc. cit. 2: pi. 180 fig. 3(2) lectotype!; basionym. -
Sulcaria sulcata (Lév.) Bystr., Ann. Univ. Mariae
Curie-Skfodowska, C, 26: 276 (1971); nom. inval.
(Art. 33).
Sulcaria sulcata f. vulpinoides (Zahlbr.)D. Hawksw.,
comb. nov. - Alectoria sulcata var. vulpinoides
Zahlbr., in Handel-Mazzetti, Symb. Bot. Sin. 3: 202
(1930); type: China, Prov. Yunnan, prope urbem
Lidjiang [Likiang], imprimis in monte YOlung-schan,
“von Einheimischen" 3606, 1914-18 (W holotype!);
basionym.
Sulcaria sulcata var. barbata (D. Hawksw.) D.
Hawksw., comb. nov. -Alectoria sulcata var. barbata
D. Hawksw., Taxon 19: 242 (1970); type: Formosa,
Mt. Shi-San, Mt. Shin-Kao-San, alt. 3,300-3,600 m,
S. Kurokawa 337, 1 January 1964 (TNS holotype!);
basionym.
Sulcaria virens f. decolorans (Asah.) D. Hawksw.,
comb. nov. - Alectoria virens var. decolorans Asah.,
in Hara, Fl. E. Himal.: 604 (1966); type: India,
Sikkim, Darjeeling, Phalut, M, Togashi, 9 May 1960
(TNS holotype!); basionym.
Sulcaria virens var. forrestii (D. Hawksw.) D.
Hawksw., comb. nov. - Alectoria virens var. forrestii
D. Hawksw., Mise. Bryol. Lichen., Nichinan 5:
1 (1969): type: China, Prov. Yunnan, Jhi Shan east of
Tali Lake, 25°48' N, alt. 10,000 ft, on rocks in dry open
situations, G. Forrest 13471, August 1914 (E holo-
type!); basionym.
References
Acharius, E. 1798: Lichenographiae Svecicae Pro-
dromus. Linköping.
- 1810: Lichenographia Universalis. Göttingen.
- 1814: Synopsis Methodica Lichenum. Lund.
Ahlner, S. 1948: Utbredningstyper bland nordiska
barrträdslavar. Ada Phytogeogr. Suec. 22: 1-257.
Ahti, T. 1959: Studies on the caribou lichen stands on
Newfoundland. Ann. Bot. Soc. Vanamo 30(4):
1-44.
- 1964: Macrolichens and their zonal distribution in
boreal and arctic Ontario. Canada. Ann. Bot. Fenn.
1: 1-35.
- 1966: Parmelia olivacea and the allied non-isidiate
and non-sorediate corticolous lichens in the North-
ern Hemisphere. Acta Bot. Fenn. 70: 1-68.
- 1973: Taxonomic notes on some species of Cladonia
subsect. Unciales. Ann. Bot. Fenn. 10: 163-184.
Ahti, T. & Hawksworth, D. L. 1974: Notes on the
lichens of Newfoundland. 4. Alectoria. Ann. Bot.
Fenn. II: 189-1%.
Ahti, T. & Hepburn. R. L. 1967: Preliminary studies
on woodland caribou range, especially on lichen
stands in Ontario. Ont. Dep't Lands Forest
Research Report (Wildlife) No. 74: 1-134.
Andrews, J. T. & Webber. P. J. 1964: A licheno-
metrical study of the north-western margin of the
Barnes ice cap: a geomorphological technique.
Geogr. Bull. 22: 80-104.
- 1969: Lichenometry to evaluate changes in glacial
mass budgets: as illustrated from north-central
Baffin Island, N.W.T. Arctic Alp. Res. 1 : 188-194.
Arnold, F. G. C. 1896: Lichenologische Fragmente.
35. Neufundland. Österr. Bot. Z. 46: 128-131,
176-182, 213-220, 245-251. 286-292, 326-332,
359-363.
Asahina, Y. 1936: Mikrochemischer Nachweis der
Flechtenstoffe IL J. Jap. Bot. 12: 25-38.
Atlas of Canada 1958: Geographic Branch. Ministry
of Mines and Tech. Services. Ottawa.
Awasthi, D. D. 1970: On Alectoria acanthodes Hue,
Alectoria confusa sp. nov. and systematic position
of Alectoria. Proc. Indian Acad. Sei.. B, 72: 149—
155.
Bachelot de la Pylaie, A. J. M. 1826: Notice sur
Tîle de Terre Neuve et quelques îles voisines. Mém.
Soc. Linn. Paris 4: 417-547.
Banfield, A. W. F. 1974: The mammals of Canada.
Toronto.
Barkman, J. J. 1958: Phytosociology and ecology of
cryptogamie epiphytes. Assen.
Bergerud, A. T. 1972: Food habits of Newfoundland
caribou. J. Wildl. Manag. 36: 913-923.
Bergerud. A. T. & Russell, L. 1964: Evaluation of
rumen food analysis for Newfoundland caribou
J. Wildl. Manag. 28: 809-814.
Beschel, R. E. 1963: Geobotanical studies on Axel
Heiberg Island in 1962. In E. Muller (ed.). Axel
Heiberg Island Preliminary Report, pp. 1-18.
Montreal. (Original volume not seen.)
Briquet, J. 1935: International rules of botanical
nomenclature adopted by the International Botani-
cal Congresses of Vienna. 1905, and Brussels, 1910,
revised by the International Botanical Congress of
Cambridge, 1930. Jena.
Brodo, I. M. 1968: The lichens of Long Island, New
York: A vegetational and floristic analysis. N. Y.
State Museum & Science Service Bull. 410: 1-330.
- 1973: The lichen genus Coccotrema in North
America. Bryologist 76: 260-270.
Bystrek, J. 1963: De variabilitate Alectoriae fusces-
centis Gyel. Ann. Univ. Mariae Curie-Sktodowska
C, 18: 41 1-418.
- 1969: Die Gattung Alectoria. Khumbu Himal. 6:
17-24.
- 1971: Taxonomic studies on the genus Alectoria.
Ann. Univ. Mariae Curie-Sktodowska C, 26:
265-279.
- 1974: Alectoria stigmata, lichenum nova species in
Alaska inventa. Frag. Flor. Geobot. 20: 255-256.
Calder, J. A. & Taylor, R. L. 1968: Flora of the
Queen Charlotte Islands. Part I . [Research Branch,
Canada, Dept, of Agr. Monogr. 4(1): 1-659.]
Ottawa.
Choisy, M. 1931: leones lichenum universalis, Ser. II,
Fasc. 2. Oullins.
Clements, F. E. & Shear, C. L. 1931: The genera of
fungi. New York.
Cringan, A. T. 1957: History, food habits and range
requirements of the woodland caribou of continental
North America. Trans. N. Amer. Wildl. Conf. 22:
485-501.
Crum, H. 1966: Evolutionary and phytogeographic
patterns in the Canadian moss flora. In R. L. Taylor
& R. A. Ludwig (ed.). The evolution of Canada's
flora, pp. 28-42. Toronto.
Culberson, C. F. 1969: Chemical and botanical guide
to lichen products. Chapel Hill.
- 1970: Supplement to "Chemical and botanical
guide to lichen products". Bryologist 73: 177-377.
- 1972: Improved conditions and new data for the
158 Alectoria in North America
OPERA BOT. 42 (1977)
identification of lichen products by a standardized
thin-layer chromotography method. J. Chromatogr.
72: 113-125.
- 1974: Conditions for the use of Merck silica gel
60 F 25j plates in the standardized thin-layer chroma-
tographic technique for lichen products. J. Chroma-
togr. 97: 107-108.
Culberson. C. F. & Hale, M. E., Jr. 1973: Chemical
and morphological evolution in Parmelia sect.
Hypotrachyna: Product of ancient hybridization?
Brittonia 25: 162-173.
Culberson, C. F. & Kristinsson, H. 1970: A standard-
ized method for the identification of lichen products.
J. Chromatogr. 46: 85-93.
Culberson, W. L. 1958: Variation in the pine-inhab-
iting vegetation of North Carolina. Ecology 39:
23-28.
- 1969: The use of chemistry in the systematics of
the lichens. Taxon 18: 152-166.
- 1969 a: Norstictic acid as a hymenial constituant of
Letharia. Mycologia 6!: 731-736.
- 1972: Disjunctive distributions in the lichen-forming
fungi. Ann. Missouri Bot. Gard. 59: 165-173.
Culberson, W. L. & Culberson, C. F. 1968: The
lichen genera Cetrelia and Platismatia (Parme-
liaceae). Contr. U.S. Nat. Herbarium 37(7): 449-
558.
- 1970: A phylogenetic view of chemical evolution
in the lichens. Bryologist 73: 1-31.
1973: Parallel evolution in lichen-forming fungi.
Science 180 : 196-198.
Dahl, E. 1950: Studies in the macrolichen flora of
South West Greenland. Meddel. Grant. 150(2):
1-176.
Dahl, E. & Krog, H. 1973: Macrolichens of Denmark,
Finland, Norway and Sweden. Oslo, Bergen and
Tromsp.
Dahl, E., Lynge. B. & Scholander, P. F. 1937: Lichens
from southeast Greenland. Skr. om Svalbard og
Ishavet 70: 1-76.
Darbishire, O. V. 1909: Lichens collected during the
2nd Norwegian Polar Expedition in 1898-1902.
Rept. 2nd Norw. A ret. Exped. 21 : 1-68.
Daubenmire, R. 1969: Ecologie plant geography of the
Pacific Northwest. Madrono 20: 1 1 1-128.
Degelius, G. 1934: Flechten aus Nordfjord und Sunn-
fjord (Norwegen). Bergens Mus. Ärbok 1934(3):
1-31.
- 1935: Das ozeanische Element der Strauch- und
Laubflechtenflora von Skandinavien. Acta Phyto-
geogr. Suec. 7: 1-411.
- 1937: Lichens from southern Alaska and the Aleutian
Islands, collected by Dr. E. Hultén. Acta Horti
Gothob. 12: 105-144.'
- 1941: Contributions to the lichen flora of North
America. 2. The lichen flora of the Great Smoky
Mountains. Ark. Bot. 30 A (3): 1-80.
DesMeules, P. & Heyland, J. 1969: Contribution to
the study of the food habits of caribou. Part 1. Lichen
preferences. Naturaliste Can. 96: 317-331.
Dey, J. P. 1974: Parmelia commensurata, a lichen new
to North America. Bryologist 77: 250-252.
Dodge, C. W. 1973: Lichen flora of the Antarctic
continent and adjacent islands. Canaan (New
Hampshire).
Dodge, C. W. & Vareschi, V. 1956: Resultados
liquenologicos de excursiones efectuadas en
VenezuelaNo. 1 . Acta Biol. Venez. 2(1): 1-12.
Dove, M. 1933: Coyote stories. Caldwell, Idaho.
Du Rietz, G. E. 1926: Vorarbeiten zu einer “Synop-
sis lichenum ", 1. Die Gattungen Alectoria, Oro-
pogon und Comicularia. Ark. Bot. 20A (11): 1—43.
Eaton, A. 1824: A manual of botany , for the northern
and middle states of America. Fourth edition.
Albany.
- 1829: A manual of botany, for the northern and
middle states of America. Fifth edition. Albany.
Edwards, R. Y„ Soos. J. & Ritcey, R. W. 1960:
Quantitative observations on epidendric lichens
used as food by caribou. Ecology 41: 425-43 1 .
Frey, E. 1936: Vorarbeiten zu einer Monographie der
L) mbilicariaceen. Her. Schweiz ■ Bot. Gesell. 44:
198-230.
- 1952: Die Flechtenflora und -Vegetation des Natio-
nalparks im Unterengadin. I. Die diskokarpen
Blatt- und Strauchflechten. Ergehn. Wiss. Unter-
such. Schweiz. Nationalparks 3: 361-503.
Fries, Th. M. 1860: Lichenes Arctoi Europae Groen-
landiaeque hactenus cogniti. Nova Acta Soc. Regiae
Sei. Upsal., ser. 3, 3: 103-398. [Reprint issued 1860;
journal issued 1861. J
- 1871: Lichenographia Scandinavica 1 . Uppsala.
Galloe, O. 1947: Natural history of the Danish lichens
7. Copenhagen.
- 1950: Natural history of the Danish lichens 8. Copen-
hagen.
- 1954: Natural history of the Danish lichens 9. Copen-
hagen.
Gyelnik, V. 1931: Additamenta ad cognitionem liche-
num extraeuropeorum. Ann. Crypt. Exot. 4:
166-174.
- 1932: Alectoria Studien. Nyt Mag. Naturvid. 70:
35-62.
- 1934: Lichenes Sipeani ex Oregon. Ann. Mus. Nat.
Hung. 28: 278-284.
- 1935: Conspectus Bryopogonum. Feddes Repert. 38:
219-255.
Hale, M. E., Jr. 1955: Studies on the chemistry and
distribution of North American lichens. Bryologist
58: 242-246.
- 1969: How to know the lichens. Dubuque, Iowa.
- 1973: Fine structure of the cortex in the lichen
family Parmeliaceae viewed with the scanning
electron microscope. Smithsonian Contr. Bot. 10:
i-iii, 1-92.
- 1974: Bulbothrix, Parmelina. Relicina, and Xantho-
parmelia, four new genera in the Parmeliaceae
(Lichens). Phytologia 28: 479-490.
- 1976: The structure of lichens viewed with the
scanning electron microscope. In D. H. Brown,
D. L. Hawksworth & R. H. Bailey (eds.), Lichen-
ology: progress and problems, pp. 1-15. London,
New York and San Francisco.
Hale, M. E., Jr. & Culberson, W. L. 1970: A fourth
checklist of the lichens of the continental United
States and Canada. Bryologist 73: 499-543.
Halsey, A. 1823: Synoptical view of the lichens
growing in the vicinity of the City of New York.
Ann. Lyceum Nat. Hist. N. Y. 1: 1-21.
Hawksworth. D. L. 1969: Chemical and nomencla-
OPERA BOT. 42 (1977)
tural notes on Alectoria (Lichenes) I. Taxon 18:
393-399.
- 1969 a: The scanning electron microscope. An aid
to the study of cortical hyphal orientation in the
lichen genera Alectoria and Comicularia. J.
Microscopie 8: 753-760.
- 1970: Chemical and nomenclatural notes on Alec-
toria (Lichenes) II. Taxon 19: 237-243.
- 1971: Chemical and nomenclatural notes on Alec-
toria (Lichenes) III. The chemistry, morphology and
distribution of Alectoria virens Tayl. J. Jap. Bot.
46: 335-342.
- 1971 a: Regional studies in Alectoria (Lichenes) 1.
The central and southern African species. Bot.
Notiser 124: 122-128.
- 1972: Regional studies in Alectoria (Lichenes) II.
The British species. Lichenologist 5: 181-261.
- 1973: Ecological factors and species delimitation in
the lichens. In V. H. Heywood (ed . ). Taxonomy
and ecology, pp. 31-69. London and New York.
- 1974: Mycologist's handbook. Kew.
- 1976: Lichen chemotaxonomy . In D. H. Brown. D.
L. Hawksworth & R. H. Bailey (eds.). Lichenology:
progress and problems, pp. 139-184. London.
New York and San Francisco.
Hawksworth, D. L. & Coppins, B. J. 1976: Alectoria
nidulifera Norrl. discovered in Scotland. Lichen-
ologist 8: 95-96.
Hawksworth, D. L. & Moore, D. M. 1969: Some
lichens from Tierra del Fuego with notes on
their chemical constituents. Bryologist 72: 247-251.
Hawksworth. D. L. & Rose, F. 1976: Lichens as
pollution monitors. London.
Henssen, A. 1963: Eine Revision der Flechten-
familien Lichinaceae und Ephebaceae. Symb. Bot.
Upsal. 18(1).
Henssen, A. & Jahns, H. M. 1973 (1974): Lichenes.
Eine Einführung in die Flechtenkunde. Stuttgart.
Hillmann, J. 1936: Parmeliaceae. Rabenh. Krypt.-FL
9, 5(3): 1-309, 1-10.
Holmgren, P. K. & Keuken, W. 1974: Index herbario-
rum. The herbaria of the world. Ed. 6. Regnum
Veg. 92: 1-397.
Howe, R. H. 1911: American species of Alectoria
occuring north of the fifteenth parallel. Mycologia
3: 106-150.
Hulting, J. 1896: Beiträge zur Flechtenflora Nord-
amerikas. Hedwigia 35: 186-193.
Huneck, S. & Follmann. G. 1970: Mitteilungen über
Flechteninhaltsstoffe. LXXXI. Zur Phytochemie
und Chemotaxonomie der Usneaceengattung Alec-
toria. Österr. Bot. Z. 118: 486-498.
lmshaug, H. A. 1956: Catalogue of Mexican lichens.
Rev. Bryol. Lichénol. 25: 321-385.
- 1957: Alpine lichens of western United States and
adjacent Canada. I. The macrolichens. Bryologist
60: 177-272.
- 1972: Typification of Ramalina usnea (L.) R. H.
Howe. Lichenologist 5: 317-318.
Imshaug, H. A. & Brodo, I. M. 1966: Biosystematic
studies on Lecanora pallida and some related lichens
in the Americas. Nova Hedwigia 12: 1-59.
Jahns, H. M. 1974 (1973): Anatomy, morphology,
and development. In V. Ahmadjian & M. E. Hale
(eds.), The lichens, pp. 3-58. New York.
References \ 59
James, P. W. 1965: A new check-list of British lichens.
Lichenologist 3: 95-153.
- 1973: The effect of air pollutants other than hydrogen
fluoride and sulphur dioxide on lichens. In B. W.
Ferry, M. S. Baddely & D. L. Hawksworth (eds.).
Air pollution and lichens, pp. 143-175. Toronto,
London.
James, P. W. & Henssen, A. 1976: The morphologi-
cal and taxonomic significance of cephalodia. In
D. H. Brown, D. L. Hawksworth & R. H. Bailey
(eds.), Lichenology: progress and problems, pp. 27-
76. London, New York, and San Francisco.
Jobin, L. 1973: L’arpenteuse de la pruche. Environne-
ment Canada, Centre de Recherches Forestières des
Laurentides, Feuillet d' Information CRFL4: 1-6.
Jorgensen. P. M. 1972: Further studies in Alectoria
sect. Divaricatae DR. Svensk Bot. Tidskr. 66-
191-201.
- 1975: Further notes on Asian Alectoria. Bryologist
78: 77-80.
Jorgensen, P. M. & Ryvarden, L. 1970: Contribution
to the lichen flora of Norway. Ârbok Univ. Bergen.
Mat.-Naturv. Ser. 10(1969): 1-24.
Karev, T. 1. 1956: Korma i pasthishchu severnogo
olenya. (Forage and pastures of the reindeer.)
Gosud. Izdat. Sel’skohoz. Liter., Moskva and
Leningrad. 100 pp. (Not seen: cited from Ahti &
Hepburn 1967).
Keissler, K. von 1958-60: Usneaceae. Rabenh. Krvpt.-
Fl. 9, 5(4): i-xiii, 1-755.
Kershaw. K. A. 1975: Studies on lichen-dominated
systems. XII. The ecological significance of thallus
color. Can. J. Bot. 53: 660-667.
Kershaw, K. A. & Larson, D. W. 1974: Studies on
lichen-dominated systems. IX. Topographic in-
fluences on microclimate and species distribution.
Can. J. Bot. 52: 1935-1945.
Körber, G. W. 1855: Systema Lichenum Germaniae.
Breslau.
Krajina, V. J. 1965: Biogeoclimatic zones and biogeo-
coenoses of British Columbia. In V. Krajina (ed.).
Ecology of western North America I. pp. 1-17.
Vancouver.
Kristinsson, H. 1969: Chemical and morphological
variation in the Cetraria islandica complex in
Iceland. Bryologist 72: 344-357.
Krog, H. 1968: The macrolichens of Alaska. Norsk
Polarinst. Skr. 144: 1-180.
- 1971: En lavekskursjon til Rogaland. Blyttia 29: 161—
168.
Krog, H. & Swinscow, T. D. V. 1974: Ramalina
species with a hollow thallus (Fistularia) in East
Africa. Nor w. J. Bot. 21: 1 1 1-124.
- 1975: Parmeliaceae, with the exclusion of Parmelia
and Usnea. in East Africa. Norw. J. Bot. 22:
115-123.
Krusenstjema, E. von & Santesson, R. 1950: Bryolo-
gical, lichenological and mycological excursion to
Jümtlund. (VU Internat. Bot. Congr. Excursion
Guide 104.) Stockholm.
Küchler, A. W. 1964: Potential natural vegetation of
the conterminous United States. Amer. Geogr.
Soc. Special Pub!. 36: 1-1 16.
Kurokawa, S. 1959: Notulae miscellaneae lichenum
japonicorum (6). J. Jap. Bot. 34: 23-24.
160 Alectoria in North America
OPERA BOT. 42 (1977)
- 1967: On the occurrence of diffractaic, physodalic,
and psoromic acids in Parmeliae. Bull. Nat. Sei.
Mus. (Tokyo) 10: 369-376.
Lamb, 1. M. 1948: New, rare or interesting lichens
from the Southern Hemisphere. Lilloa 14: 203-251.
- 1964: Antarctic lichens I. The genera Usnea. Rama-
lina, Himantormia, Alectoria, Comicularia. Brit.
Antarctic Survey Sei. Rep. 38: 1-34.
Lanjouw, J. (ed.) 1966: International code of botanical
nomenclature. Regnum Veg. 46.
Larson, D. W. & Kershaw, K. A. 1975: Studies on
lichen-dominated systems. XL Lichen-heath and
winter snow cover. Can. J. Bot. 53: 621-626.
Laundon. J. R. 1970: Lichens new to the British
flora: 4. Lichenologist 4: 297-308.
LeBlanc, F., Rao, D. N. & Comeau, G. 1972: The
epiphytic vegetation of Populus balsamifera and its
significance as an air pollution indicator in Sudbury,
Ontario. Can. J. Bot. 50: 519-528.
LeBlanc, F.. Robitaille, G. & Rao, D. N. 1974:
Biological response of lichens and bryophytes to
environmental pollution in the Murdoch ville Copper
Mine area, Québec. J. Hattori Bot. Lab. 38: 405-
433.
Letrouit-Galinou. M. A. 1974 (1973): Sexual reproduc-
tion. In V. Ahmadjian & M. E. Hale (eds.). The
lichens, pp. 59-90. New York and London.
Llano, G. A. 1948: Economic uses of lichens. Econ.
Bot. 2: 15-45.
- 1951: A contribution to the lichen flora of Alaska.
J. Wash. Acad. Sei. 41: 196-200.
Lynge, B. 1928: Lichens from Novaya Zemlya. Rep.
Scient. Results None. Exped. Nova Zemlya 43.
Lynge, B. & Scholander, P. F. 1932: Lichens from
North East Greenland collected on the Norwegian
Scientific Expeditions in 1929 and 1930. 1. Skr. ont
Svalbard og Ishavet 41: l-l 16, 7 pi., 1 map.
Macoun, J. 1902: Catalogue of Canadian plants. Part
VII. Lichenes and Hepaticae. Ottawa.
Miigdefrau, K. 1957: Flechten und Moose im baltischen
Bernstein. Ber. Deutsch. Bot. Ges. 70: 433-435.
Magnusson, A. H. 1955: A catalogue of the Hawaiian
lichens. Ark. Bot., Ser. 2, 3: 223-402.
Marie-Victorin, Fr. 1964: Flore Laurentienne .
Montreal.
Michaux, A. 1803: Flora Boreati-Americana 1-2.
Paris and Strasbourg.
Miller, G. H. 1973: Variations in lichen growth from
direct measurements: preliminary curves for
Alectoria minuscula from eastern Baffin Island,
N. W. T., Canada. Arctic Alp. Res. 5: 333-339.
Moore, B. J. 1968: The macrolichen flora of Florida.
Bryologist 71: 161-266.
Morisset, P. 1971: Endemism in the vascular plants in
the Gulf of St. Lawrence region. Naturaliste Can.
98: 167-177.
Motyka, J. 1958: Odkrycie Alectoria tortuosa Merrill
w Karpatach Wschodnich. - Alectoria tortuosa
Merrill in Carpatis Orientalibus inventa. Fragm.
Florist. Geobot. 3: 201-203.
- 1958 a: Lichenum generis Alectoria Ach., subgenus
Bryopogon (L.) Th. Fr. in Europa media (Descrip-
tiones specierum). Fragm. Florist. Geobot. 3: 205-
231.
- 1960: De speciebus generis Alectoria Ach. minus
cognitis et novis. Fragm. Florist. Geobot. 6: 441—
452.
- 1960 a: Flora Polska. Porosty (Lichenes), 5(1).
Warsaw.
- 1962: Flora Polska, Porosty (Lichenes), 5(2).
Warsaw.
- 1964: The North American species of Alectoria.
Bryologist 67: 1—44.
Müller Argoviensis. J. 1878: Lichenologische Beiträge
VIL Flora, Jena 61 : 481-496.
Nuno. M. 1971: A preliminary note on the Alectoria
asiatica-group in Asia. Mise. Brvol. Lichenol. 5:
157-158.
Nylander, W. 1885: Liste de quelques lichens récoltés
à Pile Miquelon par le Compté de Saint-Phalle,
Commandant des îles et parent due président.
Rev. Hort. Bouches-du-Rhône 1885: 132-133.
Oxner, A. M. 1956: Flora Lishainikiv Ukrdini (Lichen
flora of the Ukraine) I . Kiev.
Ozenda, P. 1963: Lichens. Handbuch der Pflanzen-
anatomie 6(9). Berlin-Nikolassee.
Packer, J. G. 1971: Endemism in the flora of western
Canada. Naturaliste Can. 98: 131-144.
Persson, B. & Santesson, J. 1970: The structure of the
depside alectorialic acid. Acta Chetn. Scand. 24:
345-346.
Poelt, J. 1965: Zur Systematik der Flechtenfamilie
Physciaceae. Nova Hedwigia 9: 21-32.
- 1969: Bestimmungsschlüssel europäischer Flech-
ten. Lehre.
- 1970: Das Konzept der Artenpaare bei den Flechten.
Vortr. Gesamtgeb. But., N. F. 4: 187-198.
- 1972: Die taxonomische Behandlung von Arten-
paaren bei den Flechten. Bot. Notiser 125: 77-81.
Porsild, A. E. 1958: Geographical distribution of some
elements in the flora of Canada. Geogr. Bull. II:
57-77.
Rabenhorst, L. 1845: Deutschlands Kryptogamen-
Flora 2. Leipzig.
Rao, D. N. & LeBlanc, F. 1967: Influence of an iron-
sintering plant on corticolous epiphytes in Wawa,
Ontario. Bryologist 70: 141-157.
Räsänen, V. 1933: Contribution to the lichen flora
of North America. Ann. Missouri Bot. Gard. 20:
7-21.
- 1939: Die Flechtenflora der nördlichen Küstenge-
gend am Laatokka-See. Ann. Bot. Soc. Vanamo 12:
1-240.
Richardson, D. H. S. 1974: The vanishing lichens.
New York.
Rickett, H . W. & Stafleu, F. A. 1959: Nomina generica
conservanda et rejicienda Spermatophytorum.
Taxon 8: 213-243.
Roland. A. E. & Smith, E. C. 1969: The flora of
Nova Scotia. Part II. The Dicotyledons. Proc. Nova
Scotia Inst. Sei. 26: 277-743.
Rose, F. 1976: Lichenological indicators of age and
continuity in woodlands. In D. H. Brown. D. L.
Hawksworth & R. H. Bailey (eds.), Lichenology:
progress and problems, pp. 279-307. London,
New York, and San Francisco.
Sato, M. 1936: Notes on the lichen flora of Minami-
Karafuto, or the Japanese Saghalien. Bull. Biogeogr.
Soc. Japan 6: 97-121 .
OPERA BOT. 42 (1977)
References 161
- 1965: Catalogus Lichenum Japonicorum (ed. 2).
Japan.
- 1967: Distribution and substratum of the Japanese
lichens. 3. The genus Oropogon Th. Fr. Mise.
Bryol. Lichenol. 4: 129-131.
Schauer, T. 1965: Ozeanische Flechten im Nordal-
penraum. Portugaliae Ada Biol. <B) 8: 17-229.
Schauer, T. & Brodo, I. M. 1966: Lecanora insignis
und L. degelii. Zwei verwandte Flechten der Alpen
und der Appalachen aus der Lecanora-subfusca-
Gruppe. Nova Hedwigia 9: 527-533.
Schindler, H. 1974: Die Flechte Alectoria fremontii
Tuck, im Nordschwarzwald. Beilr. Nalnrk. Forsch.
Südwestdeutschi. 33: 103-106.
Schofield. W. B. 1969: Phytogeography of north-
eastern North America: bryophytes and vascular
plants. Madrono 20: 155-207.
Schofield, W. B. & Crum, H. A. 1972: Disjunctions
in bryophytes. Ann. Missouri Bot. Gard. 59:
174-202,
Scotter, G. W. 1962: Productivity of arboreal lichens
and their possible importance to barren-ground
caribou (Rangifer arcticus). Arch. Soc. Zool. Bot.
Fenn. Vanamo 16: 155-161.
- 1964: Effects of forest fires on the winter range of
barren-ground caribou in northern Saskatchewan.
Can. Wildl. Serv., WiidL Manag. Bull., Ser. I, 18:
1-109.
- 1965: Chemical composition of forage lichens from
northern Saskatchewan as related to use by barren-
ground caribou. Can. J. Plant Sei. 45: 246-250.
Semander-Du Rietz, G. 1957: Om yttre faktorers in-
verkan pâ apotheciebildningen hos Parmelia tiliacea.
Svens k Bot. Tidskr. 51: 454-488.
Sharp, A. J. 1966: Some aspects of Mexican phyto-
geography. Ciencia (Mexico) 24: 229-232.
Sjödin, A. 1954: Gyelnik's new lichen names. Acta
HortiGothoh. 19: 1 13-156.
Sjörs, H. 1963: Amphi-Atlantic zonation, nemoral to
arctic. In A. Löve & D. Love (eds.). North Atlantic
biota and their history, pp. 109-125. Oxford.
Skye, E. & Hallberg, I. 1969: Changes in the lichen
flora following air pollution. Oikos 20: 547-552.
Solberg, Y. J. 1956: Dyeing of wool with lichens and
lichen substances. Ada Client. Stand. 10: 1 1 16—
1123.
- 1967: Studies on the chemistry of lichens. VI.
Chemical investigations of the lichen species Alec-
toria nigricans (Ach.) Nyl. and Parmelia alpicola
Th. Fr. Z. Naturf. 22 b: 777-783.
- 1975: Studies on the chemistry of lichens. XL
Chemical investigations of five Norwegian Alectoria
species. Acta Chem. Scand. B 29: 145-152.
Stebbins, G. L. & Major, J. 1965: Endemism and
spéciation in the California flora. Ecolog. Monogr.
35: 1-35.
Stein, B. 1879: Flechten. In F. Cohn (ed.), Krypto-
gamen-Ftora von Schlesien 2(2): 1—400. Breslau.
Stizenberger, E. 1892: Die Alectorienarten und ihre
geographische Verbreitung. Ann. Naturhist. Mus.
Wien 7: 1 17-134.
Szczawinski, A. 1953: Corticolous and lignicolous
plant communities in the forest associations of the
Douglas fir forest on Vancouver Islands. Unpubl.
Ph. D. thesis, Univ. of British Columbia, Van-
couver (Not seen: cited from Edwards et al. 1960).
Thompson, M. G. 1958: Egg sampling for the western
hemlock looper. Forest Chron. 34: 248-256.
Thomson, J. W. 1960: Lichens of arctic America.
IV. Lichens collected mainly by A. Innes-Taylor in
Greenland and the Canadian Archipelago. Brvolo-
gist 63: 181-188.
- 1972: Distribution patterns of American arctic
lichens. Can. J. Bot. 50: 1 135-1 156.
Turner, N. L. (in press): Food plants of British Colum-
bia Indians. Part II. Interior peoples. British
Columbia Prov. Mus. Handh., Victoria.
Wade, A. E. 1959: The British species of Alectoria.
Lichenologist 1 : 89-97.
Webber. P. J. & Andrews, J. T. 1973: Lichenometry: a
commentary. Arctic Alp. Res. 5: 295-302.
Wetmore. C. M. I960: The lichen genus Nephroma in
North and Middle America. Publ. Museum, Michi-
gan State Univ., Biol. Ser.. 1(11): 369-452.
- 1968: Lichens of the Black Hills of South Dakota
and Wyoming. Publ. Museum, Michigan State
Univ., Biol. Ser. 3(4): 209-464.
Williams, T. A. 1895: Notes on Mexican lichens. I.
Amer. Naturalist 29: 480-485.
Yoshimura, I. 1968: The phytogeographical relation-
ships between the Japanese and North American
species of Cladonia. J. Hattori Bot. Lab. 31: 227-
246.
- 1974: Lichen fora of Japan in colour. Osaka.
Young, S. B. 1971: The vascular flora of St. Lawrence
Island with special reference to floristic zonation
in the arctic regions. Contr. Gray Herb. Harvard
Univ. 201: 11-115.
Zahlbruckner, A. 1929-30: Catalogus Lichenum Uni-
versalis 6. Leipzig.
Index
This index comprises, in addition to epithets of accepted North American Alectorioid taxa (bold type) and ones
listed as synonyms of these here, other synonyms mentioned in North American publications (with author and
their accepted name prefixed by " = mostly discussed in detail by Hawksworth 1972), epithets of extra-
North American taxa mentioned, and those excluded from the Alectorioid genera. Pages where treatments
of accepted North American species and subspecies start are indicated in bold type, and ones with figures
indicated by an asterisk (*).
abbreviata 6 9 1 1 17* 18 26 32 35 36* 37 43 49 53
132 133* 134* 135 149 153
achariana 149 151
alaskana 68 72
alba 103
albidosorediosus 1 10
Alectoria Link, non Ach. 56
Alectoriomyces 56
altaicus 98 122 123
ambigua 49 96 97
americana 6 11 16* 19 26 33 37 41 45 49 52 54 79 88 92
93 94* 96 97 101 137 154
angolensis 131
antarctica 141
aperta 141
applanata 141
arctica 63 64
articulatus 56
asiatica 41 102 124 155
atroalbicans 151
badia 8 9 10* 26 28 41 42 49 53 1 16 129 130 146 147*
148*
barbata 156
berengeriana 152
bicolor 1 1 23 26 28 37 40 41 49 51 98 99 100* 108 1 14
152
bicolorata Mot. ?=implexa
biformis 141
boryana 63
brevior 68
Bryopogon Link 30 48 56 78 79
BryopogonTh. Fr. 49 78 79
caespitosus 109
californica 12 133 149
canaAch. 1 17 131 = capillaris
canadensis (Bryopogon negativus f.) 149 151
canadensis (Alectoria) 45 49 92 95
capillaris 6 8 9 1 1 26 28 29 33 37 40 49 52 92 1 14 115
116* 117 121 123-126 129 131 152
carlottae 9 10* 26 41 49 54 101* 102
catharinae Räs. = implexa
Ceratocladia 56
cervinula 16* 18 22 23 26 41 49 52 102 103* 104*
cetrariza 149
chalybciformis L. 9 22 26 33 36* 37 40 45 49 51 64 79
81 82* 85 106 110 150 152
cincinnata Fr. = vexillifera
“circinata” = vexillifera
Coelocaulopsis 146
compactus 103
confusa 155
congesta 141
comeliae 136 138
cornicularioides 155
crinalis 149 154
crispa Mot. = pallidum
Crustacea 141
curta 103
dahlii 155
dealbata 136 138
decolorans 156
delicata 49 92 93
dichotoma 151
Divaricatac 8 18 22 26 49 98 99 102 124
divergens 51 108 132 149 153
divergescens 19 32 132 153 155
eciliata (cervinula var.) 92
eciliata (nadvomikiana var.) 122
ecuadorensis 75
eriksonii 136
Eualectoria 56
Eujubatae 79
extensa 122
fallacina 8 9 14 15* 20 26 37 41 43 48 50 56 57 58* 69
fibrillosa 149 150
firmior Mot. = ochroleuca
flexuosa 114
Flexuosae 114
forrestii 75 156
fremontii 6 8 11 17* 19 26 28 32 35 36* 37 41 45 46* 49
52 54 1 18 1 19 135 136 137* 139 149 150
friabilis 9 11 14 15* 26 37 42 49 53 87 88 1 15 118* 1 19
120* 125 126 131 137
furcellata (Bryoria) 6 8* 9 12 16* 18 22 26 37 39 40 44
45 49 51 52 81 92 95 103 104 105* 106 1 10 1 1 1 124 150
152
furcellata (Alectoria) 103 106
fuscescens 6 8 9* 14 16* 20 22 26 28 33 37 40 45 46 49
51 53 79-81 83* 84* 85-89 91 97 98 135 149 151 152
fuscidula 114 117
Fuscidulae 114 124
genuinum Körb. = sarmentosa
gigantea 68
glabra 6 7* 8 9* 1 1 14 15* 22 26 33 37 42 45 46 49 53 85
86 87* 88 119
gracilis 61
guatemalensis 66
OPERA BOT. 42 (1977)
Index 163
haynaldii 49 92
himalayensis 155
Hyalosporae 78
hypocyphellata 68
implexa 8 11 14 15* 18* 25* 26 28 40 45 49 55 92 114
119 120* 121 128 129 131
Impie xae 1126 42 49 99 114 124 146
imshaugii 6 8* 22 26 28 32 35 36* 37 43 48 50 58* 59*
60 71 72 76
indonesica 155
infumata Mot. ? = capillaris
intricans 33 81 150
irvingii 49 107 108
isidialius 103
japonica 61
Jubatae 79 81
jubatus 22 38 44 45 78 79 85 88 1 10 150-152
karelica 122
kuemmerleana 1 17 126 129 132 146 155
lactinea 155
"laeta" = lata
lanata auct. = pubescens
lanea auct. 49 109
lanea Ehrh. ex Hoffm. = pubescens
Lanestres 79
lanestris 6 7* 8 1 1 22 26 33 36* 37 40 45 49 51 52 79 85
88* 89 90* 95 110 111 150
lapponica 110
lata Tayl. 11 12 26 41-43 48 50 58* 61 66 69
loxensis 51 53 151
luteola 56 68
luxurians 141
macra 68
melaneira 100
mexicana 8 12 20 26 43 48 50 62* 63*
minuscula (Alectoria jubata f.) 92
minuscula (Pseudephebe) 11 18* 19 26 32 36* 40 47 49
50 140 141 142* 143
minutissima 141
motykae 155
nadvornikiana 2* 6 7* 8* 9 23 26 28 35 37 40 45 46 49
51 52 98 99 114 116 122 123* 124 126 152
nana 49 1 10
negativus 88
nidulifera 44 49 103 104 106
nigricans Ach. 7* 9 1 1 20 26-29 37 40 48 51 52 63 64
65* 106 153
nitidulum 6 11 18 22 26 37 40 49 51 64 66 99 107 108*
109* 124 149
norstictica 49 127
obscurata Mot. ? = subcana
Ochroleuca 56
ochroleuca 9 12 20 24 25* 26 28 36 37 40 45 48 50 56 66
67* 68 75 1 13
olivacea 136
opaca 141
oregana 11 12 17* 18 26 32 36* 43 45 49 54 132 133
134* 135 137 153
Oropogon 5 12 30 31 99 108 151
osteina 48 61
osteola 59 121 155
pacifica (Alectoria) 143
pacificus (Bryopogon) 83
pallidior 122
pallidum 45 85 155
pellucida 57 59 114
Pellucidae 57 114
perfertilis 136
perspinosa 21 98 155
Perspinosae 98
Phaeosporae 56
philadelphiae 150 151
pikei 9 26 28 42 49 52 55 87 115 116 119 123* 125 126
131
poeltii 155
positivus 8 9* 49 51 81 83* 84-86 149 152 155
prolixa 22 150 151
prostratosteola 155
pseudocapillaris 8 20* 26 28 41 42 49 52 1 15 123* 124
126 127* 132 146
pseudofuscescens 11 12 26 28 33 36* 41 45 49 54 95 97
98 114 116 117 121 125 127 128* 129-132 147 151
pubescens 1 1 18* 19 26 32 35 36* 47 49 50 140-142 143
144* 150
rigida Fr. = ochroleuca
rubropunctata 29
ruwenzoriensis 99 156
salazinica 1 1 26 28 43 49 54 128* 130 131*
sarmentosus 6 8 9 11 12 19* 20 24 26-29 32 33 36* 37
40-42 44-48 50 56 57 61 68 69 70* 71-73 74* 75-79 99
151 153 154
sepiacea 49 96
setacea 152
Setaria 49 78 79
simplicior 6 8 9 26 37 40 44 45 49 52 89 105 109 1 10*
111 *
smithii 91 114 152
sörediata 64
sorediosa (sarmentosa f.) 72
"sorediosa" (ochroleuca f.) 68
spadiceus 152
spinulosa 122
spiralifera 8 9 10* 26 28 41 42 49 54 91* 115 126 129
131 132* 146
stigmata 48 57 69 72
stricta Ach. = vrangiana
subbicolor 49 112 114
subcana 9 22 26 32 41 49 53 54 85 91* 92 106
subchalybeiformis 64
subciliata 143 156
subdivergens 18 32 51 108 132 153
Subdivergentes 11 17* 18 26 49 132 140
subfibrillosa 61
Subfibrillosae 79 98
subfuscaGyeln. ? = capillaris
subintricans 109
subolivacea Gyeln. & För. = bicolor
subosteolus Gyeln. = subcana
Subparmelia 140
subsarmentosa 48 68
subtilis 49 127 129
succini 32
Sulcata 146
sulcata 27 32 146-148 156
“sulphurea" = ochroleuca
164 Alectoria in North America
OPERA BOT. 42 (1977)
tenerrima 45 49 89
tenuior Cromb. = ochroleuca
tenuis 6 1 1 23 26 37 40 41 49 51 99 100 112* 113* 114
tenuissimaGyeln. = bicolor
Teretiuscula 140
thrausta 13* 49 50 149 151-154
thulensisTh. Fr. = nigricans
tortuosa 8 10* 1 1 26 37 41 45 46 49 53 135 136 137* 138
139 154
Tortuosae 17* 26 49 135
trichodes Bach. Pyl. 154
trichodes 68 9 10* 11 14 16* 19 22 26 28 32 33 37 39 41
45 46 49 52 54 78-80 88 89 92* 93 94* 95 96 101 121
128 129 137 154
tristis 154
typica DR. = sarmentosa
typica 109
usneoides 154
vancouverensis 6 9 10* 14* 19 20* 24 25* 26 28 37 41
42 47 48 50 63 69 71 72 75 76*
variabilis 156
variegata 66
vexillifera 19* 20 26 28 29 33 36* 37 40 48 50 61 68 69
70* 73 74* 75
virens 49 75 139 146 148 154
vrangiana 9 26 33 36* 49 53 54 85 97 98* 129
vulpinoides 147 156
Information to contributors
Manuscripts for Opera Botanica should be written in
English (preferably), German or French. Before being
accepted all articles will be examined by one or more
referees selected by the Editor. Authors using a lan-
guage foreign to them are responsible for having their
language checked, but the Editor is entitled to submit
manuscripts to an additional linguistic examination.
Manuscripts. Manuscripts should be typed (double-
spaced, with a wide left-hand margin). The author is
requested to retain a complete copy. An abstract,
always in English, not exceeding 200 words should
appear at the beginning of the paper. The author’s
name and address should be given at the end of the
abstract.
The title should be informative but as short as
possible. A translation of non-English titles should be
given in the abstract.
Within each chapter, use no more than three grades
of headings. These should be written in lower case
throughout, and should not be underlined. A table of
contents giving a complete hierarchy of headings
should be submitted for editorial purposes. Headings
should be as short as possible - a single key-word will
often do.
Instead of footnotes, use parentheses in the main
text. Cross-references by page number must be kept to
an absolute minimum.
The desired position of all figures and tables may
be indicated in the left-hand margin. Text references to
the figures should be given as ‘Fig. T, ‘Fig. 2D, E‘,
Fig. 2D-F', or ‘Figs. IA, 2A-C’, text reference to
the tables as ‘Table 2‘ ( not abbreviated).
Scientific names of genera, species, and subspecific
categories should be underlined once to indicate
italics. Names of taxa above generic level are not
italicized. Names of persons are set in lower case and
should not be underlined or otherwise marked.
References. Text references should be given as
Hjelmqvist & Grazi (1965)' or ‘(Hjelmqvist & Grazi
1965)'. If reference to exact page(s) is necessary, it
should be made as ‘Böcher (1949 p. 287)' or ‘Bocher
(1949 pp. 287-289)’.
The List of References should include all publica-
tions cited in the text, but no others. Abbreviations
Used should be consistent: necessary editorial changes
will adhere to usage in International list of periodical
title word abbreviations (ISO 833-1974). It is essential
that punctuation conforms exactly with the examples
given below. Note especially that scientific names are
not italicized in the References and should not be
Underlined.
Rocher, T. W. 1949: Racial divergencies in Prunella
vulgaris in relation to habitat and climate. New
Phyto! . 48: 285-314.
Veo, P. F. 1972: Euphrasia L. In T. G. Tutin et al.
(eds.). Flora Europaea 3: 257-266. Cambridge.
Robinson, B. L. & Fernald, M. L. 1908: Gray's new
manual of botany. Ed. 7. New York.
Tables. Tables must be written on separate sheets.
They should be accompanied by a brief explanatory
text. The tables are usually reproduced by type-
setting; elaborate tables are therefore to be avoided.
Vertical lines should not be used, horizontal lines in
the table heads and at the base only.
Lists of collections. Extensive lists of collections will
not be published. They should be preserved at an
official institute (specified in the text), from which
copies should be obtainable.
Keys. Taxonomic keys should conform to the example
given below. Indented keys will not be accepted,
since they do not make optimal use of the page.
Illustrations. All half-tones will be placed in their
proper place in the text. All illustrations should thus
be termed figures, even if taking up an entire page.
Figures may be composed of several units separated
by I mm broad spaces. The parts of composite figures
should be designated A, B, C, etc., not a, b, c, or I, 2,
3, etc.
Illustrations must be planned for publication. They
should be reducible to a size of 140x 195 mm or less.
If possible, construct the figures either for page width
(140 mm) or column width (68 mm).
Originals of drawings should preferably be 2-3 times
the intended size; they should be in Indian ink on
paper of good quality or on plastic film. Photographs
should preferably be about the intended size; they
must be distinct and glossy. The parts of composite
figures should be similar in tone and be mounted on
completely white paper with a minimum amount of
adhesive.
Each separate illustration should be marked (outside
the area to be reproduced) with the author's name and
the number of the figure. The legends should be
written on separate sheets; in non-English articles
a translation should be given.
Great care should be taken in stating magnifications
of illustrations, making it clear whether numbers refer
to size before or after reduction. A scale line in-
corporated in the figure is the most satisfactory method
of indicating magnification.
Colour plates will be printed only if paid for by
the author.
Manuscripts and illustrations not conforming to the
above requirements will be refused or returned for
technical adjustment. If there are many handwritten
corrections in the manuscript the author will be
charged the extra expenses for type-setting.
Proofs. Proofs will be sent to the author once; the
author is requested to read them carefully and to
return them without delay. The author will be charged
for changes against the manuscript made by him in
the proofs. The number of reprints desired should be
indicated on an order form supplied with the proofs.
Original illustrations and manuscripts will be
destroyed after publication unless requested by the
author on the order form for reprints.
Key to the species of Bobartia
E Inflorescence lax, paniculate. Stem subterete, leaves flat 2
Inflorescence dense, more or less capitate. Stem and leaves externally very similar 3
2- Flowers mauve 15. B. lilacina
Flowers yellow 14 . B. paniculata
Opera Botanica 42: 1-164
ISSN 0078-5237
ISBN 91-546-0211-4
Stockholm February 25th 1977
TI 1 Statens naturvetenskapliga forskningsräd
I ^kl Swedish Natural Science Research Council
-L * J- X V Redaktionstjänsten Hditorial Service
WemterGren Center Box 23136 S-104 35 Stockholm Sweden
Tel 08-151580 Telex 13599 Rescoun S
Burling«ka Boktryckee**'
Live Music Archive
Librivox Free Audio
Metropolitan Museum
Cleveland Museum of Art
Internet Arcade
Console Living Room
Books to Borrow
Open Library
TV News
Understanding 9/11