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Author(s): W. Watson
Source: The New Phytologist, Vol. 28, No. 1 (Mar. 11, 1929), pp. 1-36
Published by: Wiley on behalf of the New Phytologist Trust
Stable URL: https://www.jstor.org/stable/2428040
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THE
NEW PHYTOLOGIST
VOL. XXVIII, No. I II MARCH, I929
PART I
GENERAL DISCUSSION
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2 W. WATSON
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The Classification of Lichens 3
by him to lichenology, his scheme has few sympathisers to-day.
His system is of interest to British lichenologists as it was followed
by Leighton(17b) and Crombie(6) in their works on British lichens.
It is only in recent years that the shackles of this slavish attachment
to Nylanderian conceptions have been broken by British licheno-
logists(32a). The importance he attached to the spermogones and
spermatia is not warranted by the present-day knowledge of these
structures, and their occurrence is too uncertain and limited for
their characters to be considered important for taxonomic purposes.
If one considers them as male gametangia and gametes, which are
sometimes functional though usually functionless, they have a
certain value, especially in doubtful cases of affinity, but they are
too frequently absent in many lichens to be of general taxonomic
value. On the theory that they are pycnidia and pycnidiospores
their value is lessened when one considers the variability of such
accessory asexual structures in the fungi. Nylander's work on the
spermogonia was supplemented in this country by that of Lindsay (20),
but, as Lorrain Smith remarks((32c) p. 205), "in many instances he
must have been dealing with species of the 'Fungi imperfecti' that
were growing in association with the scattered granules of crus-
taceous lichens." This proneness to inaccuracy when dealing with
spermogonia seriously discounts their use in a general scheme of
classification.
Leighton, in i85I, was one of the first lichenologists to pay
particular attention to the spore, as in his account of the British
angiocarpous lichens (17 a). A little later Massalongo and Koerber (16),
among continental lichenologists, evoked a large amount of hostile
criticism by using the spore as a generic distinction. They were
followed by Mudd, who in his Manual of British Lichens(25), anti-
cipated many of the generic names which are in general use to-day.
The classification proposed by Mifller-Argau(26) in I862, and the
important contributions of Reinke(30)1 in I894-6 made some ad-
vances towards the development of a natural system of classification
of lichens, but the most successful attempts have been made by
Zahlbruckner whose system has, to a large extent, been followed by
British lichenologists. It is based on all the characters of the dual
plant though the thecia and the spores contained in them have a
dominating influence. Instead of the old divisions founded upon the
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4 W. WATSON
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The Classification of Lichens 5
tion are better understood and sufficient skill in manipulation of
cultures obtained, to produce an entirely new lichen by the synthesis
of an alga and a fungus. So far as is known a completely new syn-
thesis has not been accomplished between a fungus and another
plant in the numerous experiments on mycorrhizas. Bachmann has
recently given an account of the establishment of mutualistic rela-
tions between a foreign fungus and the algal cells in the thallus
(podetial) of a Cladonia. The algal cells multiplied to a great extent,
the fungal hyphae were stimulated and a gall-like body was formed (2).
The probability that there have been many independent origins
of the consortia, together with the relationship of the spore to the
fungal constituent, has led some botanists to discard lichens as a
class, and some attempts have been made to relegate them to appro-
priate positions amongst the fungi. Lichenologists are not prepared
to admit this general su-bordination. The mutualistic relations be-
tween the fungus and the alga result in what is usually a complex
and entirely different type of plant, whose life depends as much on
the alga as the fungus, except during the formation of the thecium. In
this new plant, evolutionary development has occurred. Many lichens
are also partially or altogether dependent for their continuance on
their vegetative methods of reproduction, and in these the two
symbionts are of similar importance.
An order of lichens then, is merely a group in which families
sufficiently similar can be arranged together. Lichenologists accept
the thecia as the taxonomic bases of these " orders," which are mostly
considered as polyphyletic. When the fungal constituents of two
lichens (containing the same alga) are distinctly different, one can
generally assume that two independent associations of alga and
fungus have originated the consortia. It must not be overlooked,
however, that variation and evolution may go on in the fungal partner
after symbiotic union has been established. The likeness between the
fungal symbiont of a lichen and a free-living fungus of the present
day does not necessarily mean that the latter is the representative
of the former, since the resemblance may be due to convergence.
When the algal constituents of two lichens (containing the same
fungus) are different, it is often assumed that the consortia have
had quite independent origins from the association of free-living
algae and fungi. This may not be exactly correct, since there is a
possibility that the germinating spore of the first lichen may have
been able to enter into partnership with another alga so as to produce
the second lichen.
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6 W. WATSON
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The Classification of Lichens 7
Weber gave Cladonia as one of the i8 genera amongst which he
distributed lichens, whilst Miiller-Argau in I862 recognised the
Cladonia group (Capitularieae) as one of the three groups in his
Eulichens.
On account of these considerations and others which will be
mentioned later, it is proposed to divide Ascolichens as follows. The
Pyrenocarpeae, Coniocarpineae and Graphidineae constitute the
orders Pyrenocarpales, Coniocarpales and Graphidales respectively.
The group Cyclocarpineae is such a large and varied one that it is
more convenient to divide it into smaller groups, which may be
considered as follows:
(I) Ectolechiales. Apothecia unstalked and of simple structure;
thallus primitive and usually homoiomerous; algal cells belonging to
Chlorophyceae.
(2) Collemales. Apothecia unstalked; thallus homoiomerous,
usually more or less gelatinous; algal cells belonging to Cyanophyceae.
(3) Peltigerales. Apothecia unstalked; thallus heteromerous; algal
cells normally belonging to Cyanophyceae.
(4) Parmeliales. Apothecia unstalked (or almost so); thallus
heteromerous; algal cells belonging to Chlorophyceae.
(5) Cladoniales. Thallus heteromerous and of two kinds, the
ordinary or primary and the podetial or secondary; algal cells
belonging to Chlorophyceae. The podetial thallus is usually more or
less erect, often has a radiate structure and, on it, the apothecia are
borne.
Reinke's Cyanophila (30) includes Collemales and Peltigerales,
Ectolechiales consist of those lichens arranged by Zahlbruckner
before the cyanophyceous ones whilst Parmeliales and Cladoniales
follow them.
It must again be noted that the division of Cyclocarpineae into
five groups does not necessarily imply a separate monophyletic
origin for each group, though such a desirable objective is brought
appreciably nearer. The five groups must be regarded as convenient
for placing together families possessing similar characters.
Ectolechiales are certainly a primitive group in regard to the
thallus but the possession of a homoiomerous thallus cannot be
taken as a rigid character for purposes of classification. Some
Ectolechiale families may possess members in which the algal cells
have become arranged in a definite layer whilst some members of
the Parmeliale group may remain in a primitive thalline condition.
Froin the Ectolechiale families the Parmeliale families were probably
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8 W. WATSON
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The Classification of Lichens 9
orders or as orders. The chief objection to the use of the latter term
(in addition to the phylogenetic implication previously mentioned)
is that thalline characters, as well as thecial ones, are used as taxo-
nomic bases, so that these "orders" are not commensurate with
Pyrenocarpales, Coniocarpales and Graphidales in which thecial
characters form the main bases of classification. As a matter of
convenience their consideration as orders is justifiable.
In the further separation into families the taxonomic importance
of the apothecia and the spores are again emphasised by Zahlbruckner.
As regards the importance attached to spore characters the most
striking example is afforded by the treatment of the present-day
genera of Teloschistes, Xanthoria, Caloplaca (Placodium and Callo-
pisma), Blastenia, Anaptychia, Physcia, Rinodina and Buellia. In
many lichenological works the thalline characters of these plants
largely determine their positions. The fruticose Teloschistes, the
more or less fruticose Anaptychia and the foliose Xanthoria were
included together under the foliose Physcia. Because their thalli
were + crustaceous Caloplaca, Blastenia and Rinodina were put
under Lecanora, whilst Buellia was placed with Lecidea. In these
lichens Zahlbruckner seems to consider the colour and septation of
the spore as of great importance. Those having colourless and
polarilocular spores are placed in the neighbouring families of Telo-
schistaceae (Theloschistaceae) and Caloplacaceae, the former con-
taining the fruticose Teloschistes and Lethariopsis, and the foliose
Xanthoria, whilst the latter contains the ? crustaceous Caloplaca
and Blastenial. Those having one-septate and brown spores constitute
the allied families Physciaceae and Buelliaceae, the former con-
taining the fruticose or foliose Anaptychia, Physcia and Pyxine,
whilst the latter contains the crustaceous Rinodina and Buellia.
This consideration of the apothecium and its contained spores
as of great importance seems to be in accordance with the habits of
crustaceous lichens. In many of these the thallus is either evanescent
or poorly developed, and in many others in which a thallus is normally
present, there are varieties or forms in which the thalli are so poorly
developed that they render no aid to their identification. This is
especially the case with lichens occurring in the arctic or alpine
regions, the thallus often being so scanty that the characters of the
apothecium form the only means of determination. "Some lichens
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IO W. WATSON
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The Classification of Lichens II
Bilimbia do not show more than one septum except when full
development has occurred. Some species of Biatorina also appear
to belong to Lecidea (with unseptate spores) as a septum is not
formed till the spores are completely mature. The appearance of
longitudinal divisions in transversely-septate spores so as to give
rise to muriform spores is inconstant in some cases (e.g. species of
Rhizocarpon), many apothecia having spores with transverse septa
only; full development so as to give rise to muriform spores only
occur in some of the apothecia. On the other hand, an extra oblique
or longitudinal septum occasionally occurs in species (e.g. Thelidium
cataractarum) which are considered to have spores which are merely
transversely 3-septate. In the same plant some perithecia may
contain spores which are merely i-septate or even unseptate. Many
of these irregularities can be shown to be more apparent than real.
In Rinodina conradi and Callopisma tetrastichum the 3-septate
appearance of the spore may be considered as due to the constriction
of the lumina, whilst some other irregularities are due to incomplete
development. A sufficient number of real irregularities occur to show
that one or more extra divisions may be formed. This is especially
the case with spores having three or more septa, one or more extra
longitudinal divisions frequently occurring. The easy nature of the
passage from spores with many transverse septa to spores which are
more or less muriform, suggests that it is unnecessary to attach much
taxonomic value to such additional septation. It may be sufficient to
put the species possessing longitudinal as well as transverse division
into a different genus from those with transverse septation only,
but it seems of doubtful value for placing them in different families.
It should also be noted that Zahlbruckner's classification some-
times cuts across the old line of demarcation between lecanorine and
lecideine apothecia, as in Buelliaceae he includes the lecanorine
Rinodina and the lecideine Buellia. In Caloplacaceae the species
having a thalline margin to their apothecium are placed in Calooplaca,
whilst those without are put in Blastenia. The presence or absence
of a thalline margin to the apothecium does not deserve to have such
a weighty significance attached to it as is done by some licheno-
logists, who, on this account, place in a particular family genera
with various sporal characters. Zahlbruckner himself, in his families
Lecanoraceae and Lecideaceae, seems to attach great importance to
the presence or absence of a thalline margin. The apothecium has a
thalline margin in his family Lecanoraceae which includes the simple-
spored genera of Lecanora, Harpidium and Ochrolechia with the
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I2 W. WATSON
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The Classification of Lichens
Teloschistaceae (put under Caloplacaceae by Zahlbruckner), attempts
to apportion them to genera with or without a thalline margin lead
to much confusion. When both the thallus and apothecium are yellow
the concolorous margin may belong to either, and there is so much
uncertainty in deciding as to the presence or absence of algal cells,
around or immediately under the apothecium, that it seems advisable
to place all those which possess yellowish apothecia under one genus.
This course was practically adopted by Lorrain Smith (32 b) when
she used Placodium, though this was used in a wider sense still,
including the radiately-lobed species as well as the truly crustaceous
ones'. Malme adopts a similar course but uses Callopisma instead
of Placodium as the generic name (22). For many years Callopisma
has been used by me for all the truly crustaceous species (as in (35 e)).
When Blastenia is used as the generic name for those without thalline
margins there is much divergence in the nomenclature, e.g. both
citrinum and phloginum are put in the Callopisma section (or its
equivalent) by Lorrain Smith and Zahlbruckner, whilst Lesdain puts
them under Blastenia (18). The difficulty of deciding whether the
margin is a thalline or proper one when these are concolorous, is
often so great that it counterbalances the convenience of reducing
the size of the genus Callopisma.
Zahlbruckner further emphasises the taxonomic importance of
the sporal reproductive bodies in another way. His family Acaro-
sporaceae is founded chiefly on the presence of many spores in the
ascus. The spores, with the possible exception of Maronea, are simple.
The apothecium of A carospora may be regarded as possessing a
thalline margin whilst in Biatorella a thalline margin is never present.
Glypholechia is more or less foliose whilst the other members are
crustaceous or, at most, squamulose. Some lichens with polyspored
asci are not included under Acarosporaceae because their structure
in other respects suggests a different phylogenetic relationship. This
applies to the foliose Anzia which was included by Reinke in this
family.
Zahlbruckner therefore takes account of all the characters and
considers the probable phylogeny in his attempts to found a natural
classification, but is not altogether consistent in his methods. There
seems to be no real reason why the method of treatment adopted for
Teloschistaceae, Caloplacaceae, Physciaceae and Buelliaceae should
not be extended in regard to other lichens. For family characters
1 These are placed under the sub-genera Callopisma, with a thalline margin,
and Blastenia, without a thalline margin.
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I4 W. WATSON
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Thne Classification of Lichens I
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i6 W. WATSON
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The Classification of Lichens
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i8 W. WATSON
Class LICHENS
Family TELOSCHISTACEAE
Thallus fruticose.
Medulla of loose tissue ... ... ... ... Teloschistes
Medulla of closely-packed hyphae ... ... Lethariopsis
Thallus i foliose; horizontal or partly ascending ... Xanthoria
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The Classification of Lichens I9
Family CALOPLACACEAE
Characters as given by Zahlbruckner (36).
Ascus 8-spored. Thallus and apothecia usually K + purple
Thallus squamulose (usually placodioid) ... Placodium
Thallus entirely crustaceous
Apothecia normally yellowish or reddish ... Callopisma
Apothecia dark-coloured, K - ... ... Pyrenodesmia
Ascus many-spored. Thallus and apothecia K -
Thallus squamulose ... ... ... Candelaria
Thallus crustaceous ... ... ... Candelariella
Placodium is used here in the same way as Lorrain Smith uses
Euflacodium. Like so many other names of lichen genera it owes it
name to vegetative characters. It was first used to denote those
lichens having a radiate-squamulose (placodioid) thallus and in-
cluded some plants in which the spores were not polarilocular. In
consequence it has been rejected by some lichenologists, whilst
some use it for the placodioid lichens with simple spores. Amphiloma
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20 W. WATSON
Xanthon'a
I
Placodium
Pyrenodesmia
Callopisma Candelaria
I ~~~~~~~~~~I
(Blastenia) Candelariella
I/
(Protoblastenia)
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The Classification of Lichens 2I
lariella is truly crustaceous in its common and original species,
C. vitellina. If it is desirable to separate the 8-spored species without
parietin from Placodium and Callopisma, Candelariella may be
extended to include those which are crustaceous, whilst a similar
course may be followed for Candelaria in regard to the squamulose
species. This does not seem desirable and Candelariella should be
restricted to crustaceous species containing many spores in the
ascus. When used in this restricted sense it occupies a similar position
to Callopisma as Candelaria does to Placodium. It seems better
to retain the name Placodium medians, to put Lecanora crenata Nyl.
under Candelaria and L. epixantha Nyl. under Callopisma. The
absence of parietin is to be noted in other plants placed under
Placodium and Callopisma. It is absent in the apothecia of C.
refellens and in the thallus of several other species.
Both Smith and Zahlbruckner recognise the relationship of
Candelariella to Caloplacaceae though the latter puts the genus in
the Lecanoraceae. He, however, says that its inclusion with Lecanora
is doubtful, as it " zeigt zweifellos Beziehungen zur Gattung Caloplaca
und ist entweder der Ausgangpunkt der letzteren oder eine reduzierte
Form derselben. Ein Zusammenfassen der Gattung, lediglich nach
der Sporenform, mit Lecanora oder Lecania, von welchen sie einzeln
genommen allerdings durch geringe, in ihrer Gesamtheit jedoch
bemerkenswerte Merkmale abweicht, wiirde den phylogenetischen
Verhaltnissen kaum entsprechen " ((36) p. 207). Lorrain Smith also
states that "there is also affinity with the genus Candelaria " (32 b).
The two polar loculi are usually evident in the spore but are
sometimes absent or indistinct, as in Placodium fulgens, Callopisma
nivale, and C. rupestre (Protoblastenia r.). A i-septate appearance
may be shown in the spores of C. luteo-album and C. nivale, whilst
C. tetrastichum has the spore-contents so arranged as to give a 3-
septate appearance. C. cerinellum often has more than eight spores
in the ascus, but, like the other exceptions, should be considered as
a variant within the genus.
Family PHYSCIACEAE
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22 W. WATSON
Physcia
Pyxine
Placothallia
Diploicia Rinodina
? Rhizocarpon
Buellia
Apothecia lecanorine.
Thallus foliose or fruticose.
Cortical hyphae parallel to the surface of
the thallus ... ... ... ... ... Physcia
Cortical hyphae perpendicular to the surface of thallus.
Thallus fruticose or sub-fruticose ... Anaptychia
Thallus horizontal ... ... ... Pseudophyscia
Thallus placodioid ... ... ... Placothallia (Dimelaena)
Thallus crustaceous ... ... ... ... Rinodina
Apothecia lecideine.
Thallus foliose ... ... ... ... Pyxine
Thallus placodioid or squamulose ... ... Diploicia
Thallus crustaceous ... ... ... Buellia
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The Classification of Lichens 23
Family RAMALINACEAE
Family LECANIACEAE
Thallus crustaceous, sometimes placodioid or ? squamulose,
corticate or non-corticate above, attached by hyphae to the sub-
stratum, with heteromerous structure and green algal cells, exception-
ally orange or reddish. Apothecia usually superficial and sessile, with
or without a thalline margin; spores eight or fewer in the ascus (excep-
tionally more), colourless, i-septate, occasionally with three septa.
Ranialina
\ Thamnolecania
Solenopsora Trichoplacia
Icmadophila |
\Lecania
Lecania Lecaniopsis <-
I | / Catillaria
Biatorina "
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24 W. WATSON
Algal cells green.
Thallus ? squamulose ... ... ... Thalloidima
Thallus crustaceous.
Spores large (over 40,u long) with thick
walls; spermatia pleurogenous ... ... Megalospora
Spores under 30,u with thin walls; spermatia acrogenous.
Apothecium dark; hypothecium usually
dark ... ... ... ... ... Catillaria
Apothecium and hypothecium pale or
bright coloured ... ... ... ... Biatorina
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The Classification of Lichens
their apothecia are unknown or doubtful. Massalongo and Th. Fries
considered Cerania to have apothecia similar to Cladonia and there-
fore it has been placed in Cladoniaceae by some authors. Har-
mand(13) and Crombie(6) place Cerania and Sipfhula together as a
family (tribe) of their own. The descriptions of the apothecia remain
unconfirmed, and therefore Cerania is included with Sipfhula and
Endocena in Usneaceae, but as of doubtful place.
The spermatia in Usnea and Dufourea are acrogenous, whilst in
the other genera of Usneaceae they are pleurogenous or unknown.
There is a possibility that these two genera have evolved from
Parmeliopsis. In that case they would constitute the family Usne-
aceae, the remaining genera forming another family.
Usnea
Letharia Everniopsis
Alectoria ? Siphula
Cerania?
Evernia
Heterodea
Cetraria 7/ Omphalodium
Parmelia
Pseudoparmelia
\ / Menegazzia
Platysma
Nephromopsis\ Parmeliopsis
\ Hypogymnia
Pannoparmelia
Squamaria Jonaspis
Ochrolechia Aspicilia
\ / / Harpidiuni
Lecanora
Phyllopsora
Psora Mycoblastus
Orphniospora
-. Pseudolecanacti Biatora
" Lecidea
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W. WATSON
Family PARMELIACEAE
Family LECANORACEAE
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The Classzflcation of Lichens 27
Family LECIDEACEAE
Family BACIDIACEAE
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28 W. WATSON
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The Classification of Lichens 29
were septate, Oropogon is allied to Mvxodictyon. The large muriform
and sometimes brown spore which is single in the ascus suggests
affinity with Myxodictyon, but there are no certain links, in regard
to thalline development, connecting the two genera.
Megalospora, because of its acicular spores which may become
3-septate, is transferred from Parmeliaceae.
Psorella is like a foliose or squamose Bilimbia or Bacidia and may
differ from other members of the family in possessing rhizinae.
Zahlbruckner placed it in the family Phyllopsoraceae with Phyllopsora
and the doubtful Trichoplacia. Phyllopsora, as its name implies, is
similar to a foliose Psora and is here included in Lecideaceae, whilst
Trichoplacia is here regarded as akin to Solenopsora. The family
Phyllopsoraceae therefore disappears, the three genera which con-
stituted it being distributed to three different families.
Some species of Rhizocarpon appear to be related to Buellia.
R. alboatrum is usually quite distinct from B. myriocarpa, but
occasionally there is some difficulty in distinguishing them. In both
the paraphyses are capitate and dark-tipped. The spores become
dark in both plants, but in the former become muriform, whereas
in the latter the septation does not proceed further than the develop-
ment of one transverse septum. In the formation of the spores
R. alboatrum partially "recapitulates" the mature B. mvriocarpa.
The spores are i-septate, become dark, and then extra transverse
and longitudinal divisions occur. In R. oederi the septation seldom
proceeds further than the development of two additional transverse
septa. There are some reasons for regarding both Rhizocarpon and
Bilimbia as mixed, i.e. as having species relegated to them which
have had an origin different from the others.
Family ACAROSPORACEAE
Acarospora
Biatorella
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30 W. WATSON
Family PERTUSARIACEAE
Family GYROPHORACEAE
Family THELOTREMACEAE
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The Classification of Lichens 3I
Family CLADONIACEAE
The spores are usually simple but occasionally i- or more-septa
are developed in Pycnothelia, Thysanothecium and Baeomyces. In
some species of Baeomyces the podetium originates in the same way
as an apothecial stalk whilst in others it is formed by the extension
upwards of the primary granule. The latter method of podetial
development also occurs in Pilophorus.
Podetia usually well developed and often widening upwards.
Primary thallus squamulose and ? persistent;
podetia often scyphiferous ... ... ... Cladonia
Primary thallus evanescent or obsolete. Podetia rarely
scyphiferous.
Podetia perforated by many pores ... Clathrina
Podetia not perforated ... ... ... Cladina
Primary thallus crustaceous and persistent;
podetia short and not scyphiferous ... ... Pycnothelia
Podetia short and not widening upwards
Podetia clothed with granules or squamules Pilophorus
Podetia not clothed with granules or squamules.
Primary thallus foliose with marginal podetia Gymnoderma
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32 W. WATSON
Family STEREOCAULONACEAE
Podetia usually well developed, solid, clothed with granules or
squamules.
Spores elongate, 3- to many-septate ... Stereocaulon
Spores muriform ... ... ... ... ArgoPis
Podetia short, ? hollow and naked. Spores i-3-septate.
Primary thallus foliose. Spores cylindrical Heteromyces
Primary thallus crustaceous. Podetia widened
above with apothecia on one side. Spores fusi-
form ... ... ... ... ... ... Glossodium
Clathrina Cladina
\ Pycnothelia
Cladonia
Thysanothecium
Gymnoderma Stereocaulon
Heteromyces
Gomphillus / s/
\ / / ~~~~~Glossodi
Family GOMPHILLACEAE
Order ECTOLECHIALES
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The Classification of Lichens 33
Order PELTIGERALES
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34 W. WATSON
rules unless Peltidea, as used by Nylander, is accepted as a nomen
conservandum. The following tabular arrangement shows how
Nylander named the genera:
Order COLLEMALES
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The Classification of Lichens 35
Family COLLEMACEAE
Leptogiu n
Latzelia?
Collemna
Synechoblastus
Arctomia
Homotheciu m
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36 W. WATSON
Family PHYSMACEAE
Ramalodium
I
Lemmopsis
I Koerberia
Lempholemma
Physma I
Leciophysma
Leprocollema
(To be continued)
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