WJO *<

" GENERAL

Bulletin of the l-m**

British Museum (Natural His

LIBRARY

Botany series Vol 9 1981

British Museum (Natural History)
London 1982

Dates of publication of the parts

No 1 30 July 1981

No 2 26 November 1981

No 3 26 November 1981

No 4 17 December 1981

ISSN 0068-2292

Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset

Contents
Botany Volume 9

Page
No 1 The lichenicolous Coelomycetes

D. L. Hawksworth ......... 1

No 2 The genus Callithamnion (Rhodophyta: Ceramiaceae) in the British
Isles
P. S. Dixon & J. H. Price 99

No 3 Parmelia subgenus Amphigymnia (lichens) in East Africa

H. Krog & T. D. V. Swinscow 143

No 4 The genus Selaginella in tropical South America

A. H. G. Alston, A. C. Jermy & J. M. Rankin . . . .233

Bulletin of the ( 4 **

\ *

British Museum (Natural His

The lichenicolous Coelomycetes

D. L. Hawksworth

Botany series Vol 9 No 1 30 July 1981

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World List abbreviation: Bull Br. Mus. nat. Hist. (Bot.)

Trustees of the British Museum (Natural History), 1981

The Botany Series is edited in the Museum's Department of Botany
Keeper of Botany: Mr J. F. M. Cannon

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Assistant Editor: Mr J. R. Laundon

ISSN 0068-2292 Botany series

Vol 9 No 1 pp 1-98
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 30 July 1981

The lichenicolous Coelomycetes

D. L. Hawks worth

Commonwealth Mycological Institute, Ferry Lane, Kew, Surrey TW9 3AF*

> GENERAL

4 AUG 198!

Contents

Synopsis 1

Introduction 2

Biological and taxonomic concepts 2

Methods 5

Key to the genera 6

Accepted species 7

Ascochyta 7

Asterophoma 8

Bachmanniomyces 10

Cornutispora 14

Epicladonia 15

Karsteniomyces 22

Keissleriomyces 25

Laeviomyces 26

Libertiella 30

Lichenoconium 33

Lichenodiplis 37

Lichenosticta 38

Microcalicium [anamorphs] 41

Minutophoma 44

Nigropuncta 46

Phoma 49

Pseudoseptoria 57

Pyrenotrichum 59

Vouauxiella 64

Vouauxiomyces 67

Additional reports of anamorphs 71

Excluded species 74

Acknowledgements 89

References 89

Host index 92

Fungus index ... 95

Synopsis

A revision of all Coelomycetes so far described from lichens is presented. Twenty genera are accepted,
of which 16 are exclusively lichenicolous; seven of these are described as new and an additional new
genus is introduced for the lignicolous Levieuxia borealis P. Karsten. Forty-two species are treated,
including seven new species and seven new combinations; two additional new combinations are made
for excluded taxa. A summary of the information on the anamorphs of 21 lichenicolous ascomycetes
not considered in detail is included. Fifty epithets, including the type species often genera, are excluded

*Present address: Commonwealth Agricultural Bureaux, Farnham House, Farnham Royal, Slough SL2 3BN.

Bull. Br. Mus. not. Hist. (Bot.)9 (1) : 1-98

Issued 30 July 1981

2 D. L. HAWKSWORTH

for a variety of reasons. Keys to the accepted genera and species are provided, as are indexes to both the
lichen hosts and fungi reported as growing on them.

Introduction

Remarkable progress has been made in the last three years in synthesizing the available
information on the Coelomycetes, fungi producing conidia within flask-like (pycnidia) or
disc-like (acervuli) sporocarps (conidiomata). Sutton (1977) assessed the status of the 1336
generic names proposed for Coelomycetes up to that time and was able to reject 720 for a
variety of reasons. Synoptic keys to 200 genera were provided by Michaelides et al. (1979),
and Sutton (1980) has prepared an exhaustive treatment of 375 accepted genera, including
numerous line drawings and keys. Despite uncertainty in the application of 226 generic
names, it is now possible to approach the Coelomycetes from a firmer taxonomic base than
has hitherto been possible.

It is against this background that the present revision of the Coelomycetes now known
from lichens is presented. As in the case of my companion treatment of the lichenicolous
Hyphomycetes (Hawksworth, 1979), this work is not to be interpreted as a definitive
monograph. Rather, it should be viewed as a summation and re-assessment of the available
information to provide a starting point for future work on this group of lichenicolous fungi.

Vouaux (1914: 191-198, 281-301) treated 16 genera of lichenicolous Coelomycetes,
comprising 52 species, world-wide. Keissler (1930: 534-589) accepted 42 species in central
Europe distributed through 19 genera. In the compilation of Clauzade & Roux (1976:
77-89), 54 species were distributed through 19 genera, although it should be stressed that
their publication overlooked some previously published generic and specific names. Even at
this late date, arbitrary nineteenth century generic concepts, ignoring methods of
conidiogenesis and critical study of conidiomata, were still being employed. Furthermore,
the identities of all but a handful of taxa had never been illustrated nor re-examined since
their original description.

In the course of the last few years, I have investigated a number of lichenicolous
Coelomycetes (Hawksworth, 19750, 1976, 1977, 1978a, 1980a; Hawksworth & Dyko,
1979; Hawksworth & Punithalingam, 1973). The present work represents a culmination of
my studies on these fungi by considering all names previously described as lichenicolous
Coelomycetes and, further, by taking the opportunity to describe some hitherto
unrecognized genera and species.

Biological and taxonomic concepts
Biological concepts

Very few, if any, lichens appear to have anamorphs (imperfect states) referrable to the
Hyphomycetes (Vobis & Hawksworth, 1981), so the separation of the normal anamorphs of
the host from Hyphomycetes also present does not create any special problems. In the case of
the Coelomycetes the situation is very different. Pycnidial anamorphs are known in perhaps
as many as 8,000 lichens and these have repeatedly been misinterpreted as lichenicolous
fungi; six generic and 19 species names are excluded here as definitely or probably based on
the normal pycnidia of the host (pp. 74-89).

There is a considerable degree of variation between the coelomycete anamorphs of lichens
depending on the order to which they belong (Vobis & Hawksworth, 1981; Vobis, 1980).
However, these structures have been little studied by lichenologists and only in the last few
years have they started to be described more fully in species descriptions. Fortunately there
are a few early papers which include careful drawings of a wide range of lichen pycnidia
(Tulasne, 1852; Lindsay, 1859, 1872; Gliick, 1899; Gallee, 1927-1972); it is essential when

LICHENICOLOUS COELOMYCETES 3

working on lichenicolous Coelomycetes to have these works to hand, and also to check
herbarium specimens in the case of undocumented taxa. This is especially important when
the invading fungus does not cause any visible damage to the host (i.e. is parasymbiotic). If
necrotic patches, gall-like growths, or discolourations occur, or the pycnidia are within the
hymenium or discrete local patches, it is fortunately normally safe to assume that a
lichenicolous fungus is involved.

In some cases the normal pycnidia may function as spermatia, but at least in some groups
the conidia are evidently able to germinate to form extensive mycelia (Vobis, 1977). For
descriptive purposes it is, following usage in other ascomycetes, now usual to use the terms
'pycnidia' and 'conidia' as morphological ones not implying function. There seems to be
no reason to depart from this procedure when considering lichen-forming fungi.

Some lichens regularly occur in the pycnidial state (e.g. Cliostomum griffithii (Sm.)
Coppins) and there are examples where only pycnidia and no ascocarps are known (e.g.
Lecanactis subabietina Coppins & P. James; Coppins & James, 1979); in the latter case the
species are referred to genera defined on the ascosporic state only on the basis of similarities
in the pycnidia and chemical products. Some knowledge of these cases is prerequisite to
investigations into the lichenicolous Coelomycetes if unfortunate errors are not to be
introduced. The situation is further complicated by the recent report of a probably dioecious
lichen, Lecidea verruca Poelt (Poelt, 1980), and the probable existence of some true
lichen-forming Coelomycetes. Many genera of the latter have been described, particularly
from South America, but most merit re-investigation (Vobis & Hawksworth, 1981); one is
treated as a synonym of Vouauxiella here (p. 64) and it is possible that some others also
really represent lichenicolous fungi. One taxon described in the present revision,
Nigropuncta rugulosa (p. 46), might conceivably be a lichen-forming coelomycete.

Additional complexities arise from the occurrence of pycnidial anamorphs in
lichenicolous ascomycetes. These have so far been described for very few such fungi, and
usually very inadequately. Many more are to be expected to be recognized as the
lichenicolous ascomycetes become better known. In the course of this revision I have not
made a critical search for these and describe in full only a few which can be found without
the teleomorph (ascosporic phase). The results of a preliminary literature survey are,
however, included for completeness (pp. 71-74) as some of these reputed anamorphs can
be expected to occasionally occur alone.

The above considerations will make it clear that the production of a definitive account of
obligately lichenicolous Coelomycetes will probably not be possible until (1) much more is
known of the anamorphs of lichen-forming ascomycetes, (2) those of lichenicolous
ascomycetes, and (3) an unequivocable basis for separating lichen-forming Coelomycetes
from parasymbiotic lichenicolous taxa on 'unknown' thalli emerges. I doubt that the latter
will ever be achieved in view of the complexity of fungus-alga interrelationships
(Hawksworth, 19786). Fortunately, as stated at the start of this section, it is only where the
invading fungus does not cause necrotic patches, discolourations, or gall-like growths that
such biological demarcation problems arise.

The lichenicolous Coelomycetes causing damage to the host can clearly be interpreted as
parasitic (e.g. Lichenoconium erodens), but when the host is little affected with no
discolouration or damage to the algal cells the situation is better referred to as parasymbiotic.
In some instances (e.g. Pyrenotrichum splitgerberi) the invading fungus may depress
ascocarp production in the host, have evenly scattered conidiomata, and appear lichenized
itself; such instances may perhaps be compared to the production of one lichen by the
usurping of the algae from a pre-existing lichen colony (Hawksworth et al, 1 980).

Gall formation, especially well marked in the case of some lichenicolous Coelomycetes on
Cladonia (e.g. Bachmanniomyces uncialicola, Epicladonia sandstedei), may be viewed as an
example of parasymbiosis in which the fungus has an important role in morphogenesis. Galls
appear to be particularly common in Cladonia where they can be induced by a wide range of
fungi and perhaps also other organisms. Painstaking surveys of Cladonia galls were prepared
by Bachmann (19270, 19276, 19280, 19286, 1929), but they occur more rarely in many

4 D. L. HAWKSWORTH

other genera also (Grummann, 1960). The causal agents of many lichen galls currently
remains obscure; as knowledge of the lichenicolous fungi increases more may be related to
individual species, but in some cases it may be a host reaction to invasion by disparate taxa
which is being observed (cf. Hawksworth, 19806: 174).

In my revision of the obligately lichenicolous Hyphomycetes (Hawksworth, 1979: 186), 18
species were excluded as saprophytes found on dead or decaying thalli; several of these
were ubiquitous fungi known from an extremely wide range of substrates. In the case of the
Coelomycetes found on lichens this does not at present appear to produce any difficulties. As
far as I am aware no coelomycete species also known from other substrates has yet been
found on either healthy or decaying lichen thalli.

Taxonomic concepts

The characters of value for the delimitation of genera in Coelomycetes have already been
discussed in detail by Sutton (1973, 1980), whose works should be consulted for full
information on this aspect. It is, however, necessary to draw attention to two particular areas
of difficulty. During the last 1 5 years there has been an increasing emphasis on the method of
conidiogenesis in the delimitation of genera of Coelomycetes. However, while the increased
precision in the description of taxa is welcome and long overdue, in practice conidiogenesis
in Coelomycetes is often much more difficult to study than in many Hyphomycetes as the
conidiogenous cells tend to be very much smaller. When working close to the limits of
resolution of light microscopy, it is all too easy to be misled; ultrastructural studies have
demonstrated this in several cases. It is further becoming clear that some differences which
have been given considerable taxonomic weight are not as clearly demarcated at the ultra-
structural level as was at first presumed; the outstanding example in this case is the annellide/
phialide separation (Sutton, 1980: 20; Vobis & Hawksworth, 1981). Similar difficulties in
the Hyphomycetes led Malloch (1979) and Kendrick (1980) to assert that too much
emphasis should not be accorded to minor differences in conidiogenesis in otherwise very
similar taxa. In the case of the Coelomycetes such caution is also required and I have
endeavoured to bear this in mind in the present revision, as in my earlier account of
Lichenoconium where the problem was also recognized (Hawksworth, 1977: 161).

Sutton (1980) endeavoured to separate pycnidia-like conidiomata into two main types:
simple (unilocular) eustroma when composed of a mass of fungal cells or closely interwoven
hyphae, reserving 'pycnidia' for ostiolate, brown, thin-walled structures with walls
composed of angular pseudoparenchymatous cells (textura angularis). Wall structure is an
extremely important, but until recently largely overlooked, character in the Coelomycetes as
in many Pyrenomycetes and Loculoascomycetes. Particular attention has consequently been
accorded to this feature in the present study. The introduction of increasing numbers of
terms for superficially similar but ontogenetically or structurally distinct sporocarps
in the ascomycetes is now starting to be rejected, and it therefore seems unwise for
coelomycete terminology to pass a similar course. In this account, 'pycnidium' is
consequently retained for all globose or flask shaped conidiomata regardless of the wall
structure; the nature of the walls is of course indicated in the generic and species
descriptions.

Conidial shape, colour, septation, ornamentation, and whether extruded in mucous or
dry, are all important characters which, taken in conjunction with others, support generic
distinctions. It is, however, becoming clear that the abilities of conidia to develop additional
septa or become slightly pigmented do not provide a sound basis for generic separations on
their own. In general, such features are rather of specific importance and these, taken
together with their size and any differences in pycnidial sizes or aggregation, provide the
species concept in most genera treated here.

The ability of a fungus to occur on a particular host does not provide a specific criterion if
all other features are similar. For example, Cornutispora lichenicola was originally described
from Parmelia sulcata but is now also known from Lobaria pulmonaria, P. borreri, P.
glabratula subsp. fuliginosa, Platismatia glauca, and Rhizoplaca chrysoleuca (see p. 14);

LICHENICOLOUS COELOMYCETES 5

Vouauxiomyces truncatus was thought to be restricted to P. caperata but has since been
found on P. perlata (see p. 71). Where occurrences on different hosts are related to
differences in conidial or pycnidial size or other features they can of course support species
separations, as in the distinction of V. santessonii from V. truncatus.

Reaction of the host to infection was recognized by Santesson (1949) as an important
supporting feature at the species level in the taxonomy of lichenicolous fungi. This has
already proved of value in Lichenoconium where species differing in pycnidial, conidio-
genous cell, or conidial sizes give rise to dissimilar symptoms on the same host lichen
(Hawksworth, 1977; Christiansen, 1980).

It is with some hesitation that I introduce eight new generic names in this work, especially
as I have already described three additional genera of lichenicolous Coelomycetes and
resurrect two (Libertiella and Lichenosticta] not treated in recent reviews of coelomycete
genera. However, this was to have been expected as (1) most lichenicolous fungi in other
groups belong to obligately lichenicolous genera, and (2) the lichenicolous Coelomycetes
have been largely ignored by earlier students of both lichenicolous fungi and other
Coelomycetes. Additional obligately lichenicolous coelomycete genera certainly remain to
be collected and described. Indeed, I have several unnamed specimens which probably
represent new genera but are too scant to permit their thorough examination and
description. At the same time it is possible that 'intermediate' species may be discovered
necessitating the union of some genera accepted here (e.g. Laeviomyces and Lichenodiplis)
and others may merit transfer to independent genera (e.g. Pseudoseptoria usneae,
Vouauxiella uniseptatd).

In the case of discovery of a lichenicolous coelomycete which does not appear in this
revision, it must be stressed that before deciding it is undescribed it is of paramount
importance to establish (a) that it is not the normal pycnidia of the host, and (b) examine the
literature on other Coelomycetes (especially Sutton, 1973, 1980) to see if it can be
accommodated into another not obligately lichenicolous genus (as is the case with e.g.
Ascochyta lichenoides, Cornutispora lichenicola, Phoma species).

Glossary

Most terms which may be unfamiliar to lichenologists are defined in Hawksworth (1979:
289-290); this glossary should be supplemented by that of Sutton (1980: 642-645) for
additional terms peculiar to the Coelomycetes.

Methods

In order to ascertain the names of Coelomycetes described from lichens, the literature
searches carried out for my survey of the lichenicolous Hyphomycetes were repeated (see
Hawksworth, 1979: 189). Undoubtedly some taxa have been inadvertently overlooked, but I
endeavoured to make this aspect of the work as thorough as possible so that the present
revision can serve as a starting-point for further investigations on these fungi.

Specimens supporting the various published reports were traced wherever possible and the
material re-examined, supplemented by other collections available. Type or other material
was obtained from the following herbaria in the course of this study: Angers, B, BM, E, GZU,
H, IMI, K, LPS, M, PC, S, UPS, URM, SIENA, W, herb. Hafellner, herb. Poelt, and herb.
Vouaux (the latter with Prof. Y. Rondon at Marseille). Herbarium abbreviations follow
Holmgren & Keuken (1974). An exclamation mark (!) indicates that I have examined the
specimen cited.

The microscopic techniques used for the lichenicolous Hyphomycetes (see Hawksworth,
1 979: 1 89-1 90) were employed; the temporary erythrosin mountant was especially useful for
the study of conidiogenous cells. Sections cut at 10 //m with a freezing microtome were
examined for almost all taxa discussed. During the latter part of this work I have been
fortunate in being able to use an Olympus BH microscope fitted with Nomarski differential

6 D. L. HAWKSWORTH

interference contrast, drawing tube, and camera. The need for good optical equipment in
studies on Coelomycetes cannot be over-emphasised.

The scanning electron microscope has already proved a valuable aid in the study of those
lichenicolous Coelomycetes with ornamented conidia (Hawksworth, 1977; Hawksworth &
Dyko, 1979), but no new scanning electron micrographs are included here.

Key to the genera

This is an entirely artificial key to the genera accepted in this revision. It does not include
anamorphs of lichenicolous ascomycetes which normally occur in association with the
teleomorph (perfect state), information on which is summarized separately (pp. 71-74). In
the case of genera which are not obligately lichenicolous, the key is designed only to allow for
the known lichenicolous taxa within them.

1 Conidiomata pycnidial 2

Conidiomata acervular or superficial and helmet-shaped 21

2(1) Conidia shades of brown 3

Conidia hyaline 7

3(2) Conidia arising singly 4

Conidia arising in long chains 6

4(3) Conidia unicellular 5

Conidia 1 -septate Lichenodiplis (p. 37)

5(4) Conidia verruculose or echinulate; conidiogenous cells hyaline; pycnidial walls dark

brown Lichenoconium (p. 33)

Conidia smooth-walled; conidiogenous cells brownish; pycnidial walls subhyaline

Laeviomyces (p. 26)

6(3) Conidia unicellular and smooth-walled, or 1 -septate and rough-walled Vouauxiella (p. 64)

Conidia multicellular, muriform, rough-walled Nigropuncta (p. 46)

7(2) Conidia unbranched 8

Conidia with a distinct stem and two apical arms Cornutispora (p. 14)

8(7) Conidia septate 9

Conidia unicellular 12

9(8) Conidia 1 -septate 10

Conidia 3-septate; conidiogenous cells subglobose to short-ampulliform, polyphialidic

Keissleriomyces (p. 25)

10(9) Pycnidial wall hyphal or composed of sclerenchymatous cells 11

Pycnidial wall pseudoparenchymatous; conidiogenous cells phialidic, 4-8 x 2 -5-3 //m;
conidia rounded at both ends, narrowly ellipsoid, 8' 5-10 x 3-4 um

Ascochyta lichenoides (p. 8)
11(10) Pycnidia superficial, shades of orange, with sclerenchymatous walls; conidiophores

elongate and branched Karsteniomyces (p. 22)

Pycnidia immersed to erumpent, brown to black, with hyphal walls; conidiophores
short, usually absent Epicladonia (p. 15)

12(8) Conidiogenous cells arising singly, conidia acrogenous (borne only at the apex) . 13

Conidiogenous cells arising in irregularly branched chains, conidia acro-pleurogenous
(borne both apically and laterally); conidia lacriform . . Lichenosticta (p. 38)

13(12) Conidia bacilliform, ellipsoid or obpyriform, rounded or truncated at the base . 14

Conidia lens-shaped to pyriform, pointed at the ends, adhering in mucilage

Bachmanniomyces (p. 10)

LICHENICOLOUS COELOMYCETES 7

14(13) Pycnidia brown or black, superficial or immersed 15

Pycnidia whitish to pale orange, with a deep reddish-brown ring around the ostiole,
superficial Libertiella (p. 30)

15(14) Conidia distinctly truncated at the base; conidiogenous cells holoblastic .... 16
Conidia with a rounded base; conidiogenous cells enteroblastic (phialidic) ... 18

16(15) Conidia elongate-ellipsoid or obpyriform 17

Conidia cymbiform Pseudoseptoria (p. 57)

17(16) Conidia obpyriform; pycnidial wall pseudoparenchymatous . . Vouauxiomyces (p. 67)
Conidia elongate-ellipsoid; pycnidial wall hyphal Epiciadonia (p. 15)

18(15) Pycnidial wall entire, cells around the ostiole not separating 19

Pycnidial base formed by the conidiogenous cells alone, cells around the ostiole readily
separating Minutophoma (p. 44)

19(18) Conidiogenous cells subcylindrical or elongate-ampulliform 20

Conidiogenous cells subglobose to short-ampulliform Phoma(p. 49)

20(19) Pycnidial wall smooth; conidia subglobose or narrowly ellipsoid

Microcalicium [anamorphs] (p. 4 1 )
Pycnidial wall with pointed cells projecting from it; conidia bacilliform

Asterophoma (p. 8)

21(1) Conidiomata helmet-shaped, superficial; conidia simple, multiseptate, or to 4-branched

Pyrenotrichum (p. 59)

Conidiomata scarcely developed, acervular; conidia with c. 50 radiating multiseptate
arms Psammina (see Hawksworth, 1979: 244-246)

Accepted species

The accepted genera and species have been arranged alphabetically for ease of reference. In
the case of genera including more than a single obligately lichenicolous species, keys to the
species are included after the account of the genus itself. Descriptions and full synonymies
for taxa I have previously treated elsewhere are generally omitted, but references to full
descriptions, illustrations, etc., are provided. Synonyms not repeated from these works have
been included in the Fungus index (pp. 95-98). Where genera include both lichenicolous
and non-lichenicolous species, the generic diagnoses cover the genus as a whole and are
partly based on those provided by Sutton ( 1 980).

I. ASCOCHYTA Lib.
PL Crypt. Ard.Jasc. I : 8 (1830).

See Sutton (1980 : 408) for generic nomenclature.

Conidiomata pycnidial, arising singly or in small groups, globose, immersed, brown to
dark brown, ostiolate; walls composed of a few layers (often 1-3) of cells, cells pseudo-
parenchymatous, pale to dark brown, rather thin-walled, textura angularis. Conidiophores
absent. Conidiogneous cells enteroblastic, lining the pycnidial cavity, acrogenous, short-
cylindrical to doliiform, phialidic, not proliferating, hyaline. Conidia arising singly, not
catenate, narrowly ellipsoid or subcylindrical, rounded at the apex but often with a some-
what truncated base, hyaline, 1 -septate when mature (exceptionally becoming 2-septate),
sometimes distinctly guttulate, thin-walled, smooth-walled.

Type species: Ascochyta pisi Lib.

Number of species: About 350 species are currently accepted but there are many more
names in the literature. Taxa in the genus have generally been separated on a host basis in
the past, and thorough revisions are required on the lines of that of Punithalingam (1979)

8 D. L. HAWKSWORTH

concerning the graminicolous species. Most species occur as leaf and stem-spotting fungi;
only a single lichenicolous species is referred to this genus here.

Observations: For practical purposes this genus may be viewed as a 1 -septate counterpart
of Phoma (see p. 49), which it resembles closely in all other respects. As noted by Sutton
(loc. cit.), attempts to separate these two genera on the basis of ultrastructural differences in
conidiogenesis are difficult to relate to light-microscopical observations.

1. Ascochyta lichenodes (A. L. Sm.) D. Hawksw., Notes R. hot. Gdn Edinb. 36 : 1 82 (1978).
For illustration and further discussion of this species see Hawksworth (loc. cit.).
Host: Indeterminate sterile crustose lichen on bark.
Distribution: British Isles (England). Known only from the original collection.

II. ASTEROPHOMA D. Hawksw. gen. nov.

Genus lichenicola ad Coelomycetes Sphaeropsidales pertinens. Conidiomata singularia, uniloculata,
dispersa vel laxe aggregata, globosa, superficialia vel hemi-superficialia, atrobrunnea vel nigra,
ostiolata, cum muris atrobrunneis e cellulis elongatis pseudoparenchymaticis externe attenuatis et
erumpescentis. Conidiophora desunt. Cellulae conidiogenae enteroblasticae, acrogenae, anguste
ampulliformes vel late subcylindrices, phialidicae, interdum biphialidicae, non prolifericae, hyalinae.
Conidia subcylindrica, hyalina, simplicia, laevia, minutissima, in muco aggregata.

Conidiomata pycnidial, arising singly, uniloculate, scattered or loosely aggregated,
globose, superficial or half-immersed, dark brown to black, ostiolate; walls composed of
radially orientated elongated dark brown pseudoparenchymatous cells becoming attenuated
and projecting slightly at the exterior to give the whole a star-like appearance. Conidiophores
absent. Conidiogenous cells enteroblastic, interdigitated with the inner wall cells and lining
the pycnidial cavity, acrogenous, narrowly ampulliform to broadly subcylindrical, phialidic,
occasionally biphialidic, not proliferating, hyaline. Conidia subcylindrical, hyaline, simple,
smooth-walled, minute, aggregated into a slimy mass which is extruded through the ostiole
as a whitish drop.

Type species: Asterophoma mazaediicola D. Hawksw. (holotypus).
Number of species: Monotypic.

Observations: This new genus recalls Minutophoma (see p. 44) in the minute
Conidiomata and subcylindrical conidia, but differs in the entire conidiomatal wall, presence
of an ostiole, the externally attenuated wall cells, the hyaline conidiogenous cells that are
also occasionally biphialidic, and the conidia being extruded in a mucilaginous whitish drop.
It does not appear to have any other close allies.

1. Asterophoma mazaediicola D. Hawksw. sp. nov.
(Fig. 1A-D)

Fungus lichenicola. Conidiomata singularia, uniloculata, dispersa vel laxe aggregata, globosa, super-
ficialia vel hemi-superficialia, atrobrunnea vel nigra, 30-42 /zm diam, ostiolata, cum muris textura
angulari 7-10 jim latis e cellulis atrobrunneis elongatis et externe attenuatis et erumpescentis
5-8 x 2-4 nm. Conidiophora desunt. Cellulae conidiogenae enteroblasticae, acrogenae, anguste
ampulliformiae vel late subcylindrices, phialidicae, interdum biphialidicae, non prolifericae, hyalinae,
5-7x2-5-3-5 fim. Conidia subcylindrica, hyalina, simplicia, laevia, l-5-3(-3'5)xO-5-l-5 //m, in muco
aggregata.

Typus: Italia, Mendelgebirge, Nordhange des Roen, inter Mendel-Pass et Marga di Romeno, in
mazaediis Caliciis trabinellae Ach., 24.x. 1976, J. Hafellner 1 822 (herb. Hafellner holotypus!).

Conidiomata pycnidial, superficial or partly immersed, arising in the mazaedial ascospore
mass of the host, arising singly but sometimes loosely aggregated and juxtaposed, to 12 in a
single mazaedium, dark brown to black, 3(M2 //m diam, ostiolate; wall 7-10/zm thick,
composed of a layer of radially elongated dark brown pseudoparenchymatous cells becoming

LICHENICOLOUS COELOMYCETES

lO/^rn

Fig. 1 Asterophoma mazaediicola (herb. Hafellner holotype). A, Infected mazaedium with
pycnidia extruding conidia in mucilaginous drops. B, Vertical section of pycnidium. C,
Conidiogenous cells. D, Conidia.

attenuated and projecting slightly at the exterior to give the whole a star-like appearance,
5-8 x 2-4 //m, lined internally with smaller subhyaline to hyaline subglobose to polyhedral
pseudoparenchymatous cells 1'5-3'5/zm diam; mycelium ramifying through the mazae-
dium, hyphae hyaline, uneven in diameter, septate, irregularly branched, mainly
l'5-2'5 //m thick. Conidiogenous cells enteroblastic, interdigitated with the inner wall cells
lining the pycnidial cavity, difficult to distinguish, acrogenous, narrowly ampulliform to
broadly subcylindrical, rather variable in shape, phialidic, occasionally biphialidic with two
Conidiogenous loci per cell, not proliferating, hyaline, 5-7 x2'5-3'5 //m. Conidia
abundantly produced, subcylindrical, rounded at the apices, hyaline, simple, smooth-walled,
not distinctly guttulate, l'5-3(-3-5)xO'5-l-5 //m, aggregated into a slimy mass which is
extruded through the ostiole as a whitish drop.

Hosts: Calicium glaucellum Ach. and C. trabinellum Ach., apparently confined to
mazaedial ascospore masses. Especially frequent on C. trabinellum. Evidently a mild
pathogen as diseased fruits often appear stunted and bent; the amount of vulpinic acid
pigment produced by C. trabinellum also appears to be reduced in diseased specimens so that
such collections can easily be misidentified as C. glaucellum. The absence of records from
other Calicium species suggests that the fungus has a very narrow host range; indeed C.
glaucellum and C. trabinellum are extermely close to one another and conceivably

10 D. L. HAWKSWORTH

chemotypes of a single taxon (Tibell, 1975 : 106). Infected fruits are easily recognized by the
verrucose surface that derives from the pycnidia themselves and particularly by the extruded
white mucilaginous drops containing the conidia.

Distribution: Canada, France, Italy, Norway, Rwanda, Sweden and the U.S.A. Probably
scattered throughout the range of its host species (i.e. circumboreal), to judge from material
so far available, and locally abundant.

Observations: A distinctive species easily recognized by its very specialized habitat. The
pycnidia of Microcalicium subpedicellatum (p. 42) are larger (40-60 //m diam), with a
totally different wall structure, longer phialides, broadly ellipsoid to subglobose conidia, and
occur only on the thalli (not in the mazaedium); M. subpedicellatum is known from
Calicium glaucellum (Tibell, 1978 : 244) but has not been reported from C. trabinellum. The
pycnidia of these Calicium species, when present, only occur on the thallus, not the
mazaedium, and produce larger conidia c. 4-5 x 1 -5 //m (fide L. Tibell, in litt.).

Additional specimens: Canada: British Columbia: Glacier National Park, Highway 1, 7 km NE of
Rogers Pass, on Calicium glaucellum, 6 September 1972, L. Tibell 4959 (UPS!); Wells Gray National
Park, Helmcken Falls, on C. glaucellum, 19 September 1972, L. Tibell 5185 (UPS!); Pemberton Valley,
c. 20 miles N of Pemberton, on C. glaucellum, 17 September 1972, L. Tibell 5180 (UPS!). France:
Haute Savoie : Chamonix, - 5 km S of Planpra, Plan des Chablettes, on C. trabinellum, 27 August

1970, L. Tibell 4375 (UPS!); 1-5 km SSE of Chamonix, on C. trabinellum, 26 August 1970, L. Tibell
4383 (UPS!). Italy: Trentino-Alto Adige : Ortlegruppe, Martelltal, on C. trabinellum, 2 October

1971, L. Tibell 4466 (UPS!). Norway: Hedmark, Engerdal par., 3-5 km SW of Hovdroseter, on C.
trabinellum, 29 July 1973, L. Tibell 5548 (UPS!). Rwanda: Chaine des Birunga, Versant S of Bisoke,
on C. trabinellum, 25 February 1972, J. Lambinon 72/RW/883 (LG n.v., fide Tibell, in litt.).
Sweden: Asele Lappmark : Frederika par., Vargan, 1'8 km SW of Flakatrask, on C. trabinellum, 5
August 1972, L. Tibell 4612 (UPS!). Dalarna : Hamra par., Hamra National Park, on C. trabinellum,
20 July 1973, L. Tibell 5433 (UPS!), 5448 (UPS!); loc. cit., on C. glaucellum, 20 July 1973, L. Tibell
5424 (UPS!). Harjedalen: Ljusnedal par., 6 km NNE of Ljusnedal, Ormberget, on C. trabinellum, 25
July 1973, L. Tibell 5503 (UPS!). Jamtland: Fors par., 10km NNW of Fors, on C. trabinellum, 2
August 1977, L. Tibell 7262 (UPS!). Norbotten: Overkalix par., 39 km NW of Overkalix, mire between
Svartberget and O. Naverberget, on C. trabinellum, 14 July 1977, L. Tibell 6762 (UPS!). Lule
Lappmark: Muddas National Park, 5 km E of Porjus, on C. glaucellum, 21 July 1977, L. Tibell 6962
(UPS!); loc. cit., at the little stream N of Muddusfallet, on C. trabinellum, 21 July 1977, L. Tibell 7026
(UPS!). Uppland: Vange par., Fiby urskog, on C. glaucellum 22 March 1964, L. Tibell 2044a (UPS!).
U.S.A.: Michigan : Keweenaw Co., Isle Royale, Rasberry Island, outside Rock Harbor, on C.
trabinellum, 24 August 1972, L. Tibell 4656 (UPS!).

III. BACHMANNIOMYCES D. Hawskw. gen. nov.

Genus lichenicola ad Coelomycetes Sphaeropsidales pertinens. Conidiomata singularia, uniloculata,
dispersa, globosa vel subglobosa, immersa, hyalina sed ostiolis aurantiacis vel brunneis, cum muris
textura intricatis, ostiolata. Conidiophora desunt. Cellulae conidiogenae holoblasticae, acrogenae,
elongato-ampulliformes, prolifericae, annellatae, hyalinae. Conidia late fusiformia ad pyriformia,
plerumque asymmetrica, hyalina, simplicia, laevia, in mucosa.

Conidiomata pycnidial (eustromatic), arising singly, scattered, globose to subglobose,
immersed, hyaline but orange to brownish at the ostiole; walls composed of moderately
thick-walled compactly entwined hyphae forming a textura intricata, hyaline except near the
ostiole. Conidiophores absent. Conidiogenous cells holoblastic, lining the inner wall of the
pycnidial cavity, acrogenous, elongate-ampulliform, somewhat broader at the base and
tapering above, proliferating, annellate, hyaline, unbranched, occasionally septate. Conidia
slimy and adhering together, not catenate, lens-shaped to pyriform, attenuated at the apices,
the base narrowly truncate at the point of secession, often asymmetric, hyaline, simple,
smooth-walled.

Type species: Bachmanniomyces uncialicola (Zopf) D. Hawksw. (syn. Phyllosticta
uncialicola Zopf holotypus).

LICHENICOLOUS COELOMYCETES

11

Fig. 2 Bachmanniomyces uncialicola (W 1929/2 holotype of Sirococcus lichenicola). A,
Gall with pycnidia close to the apex of a branch, note the thinner stem above the gall
(x 12-5). B, Gall with pycnidia arising laterally, the branch similar in thickness above
and below the gall (x 12'5).

Number of species: Monotypic.

Observations: Bachmanniomyces has no very close allies amongst the non-lichenicolous
Coelomycetes. The conidiogenous cells in Discosporium Hohnel (type species D. populeum
(Sacc.) B. Sutton) are rather similar but the pycnidia in that genus are dark brown and
aggregated into stromata. Sirococcus Preuss (type species S. stobilinus Preuss) was probably
selected by Keissler (in Bachmann, 1929) for this fungus as he interpreted the conidia as
catenate, but in Sirococcus the pycnidia are carbonaceous, lack a distinct ostiole, and the
conidiogenous cells are branched (see Sutton, 1980).

This new genus is named in honour of Edwald Th. Bachmann (1850-1937) in recognition
of his pioneering work on gall-like structures in lichens, including investigations of the type
species of Bachmanniomyces (Bachmann, 1929).

1. Bachmanniomyces uncialicola (Zopf) D. Hawksw. comb. nov.
(Figs. 2A-B, 3A-E)

Phyllosticta uncialicola Zopf, Hedwigia 35 : 324 (1896).
type: British Isles, Scotland, Perthshire, Dunkeld, summit of Birnam Hill, on Cladonia uncialis,

September 1858, W. L. Lindsay (E lectotype!).

Phoma uncialicola (Zopf) Vouaux, Bull, trimest. Soc. mycol Fr. 30 : 198 (1914).
Sirococcus lichenicola Keissler, in Bachmann, Arch. Protistenk. 66 : 469 (1929).
Type: Sweden, Asele Lappmark, Kirchspiel Vilhelmina, auf Cladonia amaurocraea, Sommer
1928, G. NilssonfG. Degelius](W 1929/2 holotype!).

14 D. L. HAWKSWORTH

lectotype for Zopfs name here, but the possibility that it was the holotype should not be
discounted.

Vouaux (1914 : 198) indicated 'sans doute' that Spilomium epicladonia H. Olivier was a
synonym of the present species. However, as the conidia of Olivier's fungus were given as
6-8 x 3-4 um it seems unlikely that this interpretation is correct.

Additional specimens: Canada: British Columbia, Queen Charlotte Islands, Graham Island, central
part between Boulton Lake and Watun River, 5349'N, 132 4' W, on Cladonia uncialis, 3 August
1966, H. Sjors (H!). Newfoundland, St. Mary's District, 2 mi E of Peter's River, on C. terrae-novae, 1
June 1956, T. Ahli 9327 (H!). Germany: Oldenburg, Kaihausermoor in Moorboden, on C. uncialis,
April 1918, H. Sandstede [Clad. Exs. no. 162.] (BM!, UPS!); loc. cit., October 1917, H. Sandstede
[Clad. Exs. no. 161.] (BM!). Italy: Hohe zwischen Glashutten und Volsbach, siidlich von Bayreuth,
auf Sandboden des braunen Jura, on C. uncialis, 6 September 1884, F. G. C. Arnold [Lich. Exs. no.
102 la.] (BM!). Sweden: Uppland, Uppsala, Witulfsberg, on C. uncialis, 1852, Th. M. Fries (UPS!).

IV. CORNUTISPORA Piroz.

Myeologia 65 -.163(1913).

Conidiomata pycnidial, arising singly, subgloboge, alrriQM translucent to pale brown,
immersed to almost superficial, opening by an irregular disintegration pf the upper part of
the pycnidial wall; walls composed of thin-walled ' pseudoparenchyniatous cells, textura
angularis. Conidiophores arising from the inner wa-11 of the pycnidial cavity, irregularly
branched, septate, smooth-walled, hyaline, conidia forming acropleurogenously. Conidio-
genous cells polyblastic, integrated, indeterminate,- broader towards the apices, hyaline, with
1-4 apical denticles. Conidia arising singly, triradiate with a subcylindrical stem, two apical
branches diverging at an obtuse angle, and a distinctly narrowed foot-cell, hyaline, simple,
smooth-walled.

Type species: Cornutispora limaciformis Piroz.

Number of species: Two; C. limaciformis, which occurs in the apothecia of Therrya
fuckelli (Rehm) Kujala (Phacidiales), and a single lichenicolous species distinguished by its
much shorter conidia.

Observations: The conidia of this genus are especially characteristic and it appears to have
no close relatives in the Coelomycetes.

1. Cornutispora lichenicola D. Hawksw. & B. Sutton, in Hawksworth, Trans. Br. mycol. Soc.

67:51(1976).
(Fig. 4A-B)

This fungus is described and illustrated in Hawksworth (1976:51-53), and further
drawings are given by Sutton (1980 : 90), so it is not discussed in detail here.

Hosts: Lobaria pulmonaria (L.) Hoffm. (thallus), Parmelia borreri (Sm.) Turner, P.
glabratula subsp. fuliginosa (Fr. ex Duby) Laundon (thallus), P. sulcata Taylor (thallus),
Platismatia glauca (L.) Culb. & C. Culb. (thallus), and Rhizoplaca chrysoleuca (Sm.) Zopf
(syn. Lecanora rubina (Vill.) Ach.) (apothecia). Commonly in mixed infections with other
lichenicolous fungi. The host tissues are sometimes blackened, decolourized, or otherwise
discoloured, but it is often not clear whether this is due to the Cornutispora or the other fungi
present.

Distribution: Austria, British Isles (England, Scotland), Italy, and Switzerland.

Specimens (additional to Hawksworth, loc. cit): Austria: Burgenland: Bez. Oberwart, SW of Bad
Tatzmannsdorf, on Parmelia sulcata, 1 April 1980, J. Poelt (GZU!; with Lichenoconium usneae).
British Isles: England, S. Devon, Slapton, Slapton Ley Nature Reserve, Peasdish, on P. borreri, 28
August 1980, D. L. Hawksworth 5064 (IMI 251276!; with L. lecanorae)\ S. Somerset, Porlock, Horner
Combe, on Platismatia glauca, 21 September 1980, D. L. Hawksworth 5072 (IMI 251490!); S.

LICHENICOLOUS COELOMYCETES 1 3

1 00-200 //m diam when mature; pycnidial wall translucent, hyaline, mainly 12-20//m
thick, becoming slightly more swollen and orange to brownish near the ostiole, the pigmen-
tation due to interstitial mainly extra-hyphal deposition, only the limiting membrane of the
hyphae in the ostiolar region coloured, wall throughout formed of irregularly orientated
entwined and compacted moderately thick-walled hyphae mainly 2-3 um wide.
Conidiogenous cells holoblastic, arising from the inner wall of the pycnidium, completely
lining the pycnidial cavity, elongate-ampulliform, the bases often expanded and tapering
markedly towards the apex, hyaline, smooth-walled, annellidic with to at least 4
annellations, unbranched, simple or occasionally 1 -septate, 7-1 2 //m tall, mainly 3-4 um
wide at the base and tapering to 2-3 um at the apex. Conidia abundantly produced, not
catenate, slimy and adhering together in large groups in aqueous preparations, lens-shaped
to pyriform, often asymmetrical, apices often somewhat attenuated, the basal one more
markedly so and also abruptly truncate when about 1 um thick, hyaline, simple, often with
a large guttule, smooth-walled, (7-)8-10(-10'5) x 4-5'5(-6)//m.

Hosts: Cladonia amaurocraea (Florke) Schaerer, C. terrae-novae Ahti, and C. uncialis (L.)
Wigg., thalli. The infection leads to the formation of distinctive galls (see above; Figs 2, 3).
The galls are of the same colour as the healthy thallus and healthy algal cells consistently
occur in the close proximity of the pycnidia; this fungus consequently appears to form a
stable relationship with the host and not to be a pathogen. Vouaux's (1914 : 198) mention of
its occurrence on C. bellidiflora (Ach.) Schaerer is due to confusion with ' Abrothallus'
moorei Lindsay, and his mention of it on C. stellaris Opiz may also be incorrect. Keissler's
(1930 : 540) reports from C. ochrochlora Florke, 'C. silvatica' and C. surrecta Florke, and
Sandstede's (193 1 : 3 17) from C. turgida (Ehrh.) Vainio, require confirmation.

Distribution: British Isles (Scotland), Canada, France, Germany, Italy, Sweden and
Switzerland.

Observations: A very detailed account of the galls produced by Bachmanniomyces
uncialicola is given by Bachmann (1929 : 460^469) who did, however, describe the conidia
as forming in chains. I was unable to see any evidence of this, though the conidia strongly
adhere in irregular slimy masses which presumably swell when moist and ooze through the
rather wide ostioles. If the conidia were catenate, it would be anticipated that the mature
ones would have a minute truncate point of secession at both ends when in fact this only
occurs at their bases. The first-formed conidium arises holoblastically and if a young
pycnidium is studied no or few conidiogenous cells with annellations will be found. The
annellations are not easy to see in older conidiogenous cells unless studied in erythrosin-
ammonia or a similar mountant.

Lindsay (1859 : pi. 7 figs 1 5-16) was evidently the first author to recognize this fungus. He
originally considered it to be a pycnidial state of Lee idea cladoniaria Nyl. (syn. Nesolechia
cladoniaria (Nyl.) Arnold) but later realized that this was not so (Lindsay, 1871 : 28-29). He
also discovered an ascomycete on this host, named ' ' Abrothallus' moorei Lindsay (Lindsay,
1869<2 : 545), and described its conidia which were, however, much longer and narrower
than those of Bachmanniomyces uncialicola. Vainio (1887:265) recognized that defor-
mations of Cladonia uncialis named as var. leprosa Delise were caused by a lichenicolous
fungus, and Harmand (1907 : 245) ascribed this to Phyllosticta uncialicola Zopf.

Zopf (1896:324) gave an extremely short diagnosis of Phyllosticta uncialicola:
'Anschwellung an der Podetien bewirkend. Conidien einzellig, farblos, ei- oder birnformig,
8, 4 : 5' and did not cite any specimens. No material of this taxon could be located in B (B.
Hein, in litt.} but the original description is so characteristic of the present species that there
can be no doubt as to the taxon he intended to describe. However, as Zopf introduced the
name in his compendium of lichenicolous fungi then known, it is most unlikely that he
overlooked Lindsay's (1859) description and comments on this fungus; Zopf cites these
works in the introduction to his list and it is probable that he introduced the name with
Lindsay's then unnamed fungus in mind. Lindsay's material is consequently designated

14 D. L. HAWKSWORTH

lectotype for ZopPs name here, but the possibility that it was the holotype should not be
discounted.

Vouaux (1914 : 198) indicated 'sans doute' that Spilomium epicladonia H. Olivier was a
synonym of the present species. However, as the conidia of Olivier's fungus were given as
6-8 x 3-4 /zm it seems unlikely that this interpretation is correct.

Additional specimens: Canada: British Columbia, Queen Charlotte Islands, Graham Island, central
part between Boulton Lake and Watun River, 53 49' N, 132 4' W, on Cladonia uncialis, 3 August
1966, H. Sjors (H!). Newfoundland, St. Mary's District, 2 mi E of Peter's River, on C. terrae-novae, 1
June 1956, T. Ahti 9327 (H!). Germany: Oldenburg, Kaihausermoor in Moorboden, on C. uncialis,
April 1918, H. Sandstede [Clad. Exs. no. 162.] (BM!, UPS!); loc. cit., October 1917, H. Sandstede
[Clad. Exs. no. 161.] (BM!). Italy: Hone zwischen Glashiitten und Volsbach, sudlich von Bayreuth,
auf Sandboden des braunen Jura, on C. uncialis, 6 September 1884, F. G. C. Arnold [Lich. Exs. no.
102 la.] (BM!). Sweden: Uppland, Uppsala, Witulfsberg, on C. uncialis, 1 852, Th. M. Fries (UPS!).

IV. CORNUTISPORA Piroz.
Mycologia 65 -.163(1913).

Conidiomata pycnidial, arising singly, subglobose, almQst translucent to pale brown,
immersed to almost superficial, opening by an irregular disintegration pf the upper part of
the pycnidial wall; walls composed of thin-walled ' pseudoparenchyniatous cells, textura
angularis. Conidiophores arising from the inner wall of the pycnidial cavity, irregularly
branched, septate, smooth-walled, hyaline, conidia forming acropleurogenously. Conidio-
genous cells polyblastic, integrated, indeterminate,- broader towards the apices, hyaline, with
1-4 apical denticles. Conidia arising singly, triradiate with a subcylindrical stem, two apical
branches diverging at an obtuse angle, and a distinctly narrowed foot-cell, hyaline, simple,
smooth-walled.

Type species: Cornutispora limaciformis Piroz.

Number of species: Two; C. limaciformis, which occurs in the apothecia of Therrya
fuckelli (Rehm) Kujala (Phacidiales), and a single lichenicolous species distinguished by its
much shorter conidia.

Observations: The conidia of this genus are especially characteristic and it appears to have
no close relatives in the Coelomycetes.

1. Cornutispora lichenicola D. Hawksw. & B. Sutton, in Hawksworth, Trans. Br. mycol. Soc.

67:51(1976).
(Fig. 4A-B)

This fungus is described and illustrated in Hawksworth (1976:51-53), and further
drawings are given by Sutton ( 1 980 : 90), so it is not discussed in detail here.

Hosts: Lobaria pulmonaria (L.) Hoffm. (thallus), Parmelia borreri (Sm.) Turner, P.
glabratula subsp. fuliginosa (Fr. ex Duby) Laundon (thallus), P. sulcata Taylor (thallus),
Platismatia glauca (L.) Culb. & C. Culb. (thallus), and Rhizoplaca chrysoleuca (Sm.) Zopf
(syn. Lecanora rubina (Vill.) Ach.) (apothecia). Commonly in mixed infections with other
lichenicolous fungi. The host tissues are sometimes blackened, decolourized, or otherwise
discoloured, but it is often not clear whether this is due to the Cornutispora or the other fungi
present.

Distribution: Austria, British Isles (England, Scotland), Italy, and Switzerland.

Specimens (additional to Hawksworth, loc. cit}: Austria: Burgenland: Bez. Oberwart, SW of Bad
Tatzmannsdorf, on Parmelia sulcata, 1 April 1980, J. Poelt (GZU!; with Lichenoconium usneae).
British Isles: England, S. Devon, Slapton, Slapton Ley Nature Reserve, Peasdish, on P. borreri, 28
August 1980, D. L. Hawksworth 5064 (IMI 251276!; with L. lecanorae); S. Somerset, Porlock, Horner
Combe, on Platismatia glauca, 21 September 1980, D. L. Hawksworth 5072 (IMI 251490!); S.

LICHENICOLOUS COELOMYCETES

15

lOjim

Fig. 4 Cornutispora lichenicola (IMI 186831 holotype). A, Conidiogenous cells. B, Conidia.

Reproduced from Hawksworth (1976 : 52).

Somerset, Nettlecombe Park, on Parmelia sulcata, 20 September 1980, D. L. Hawksworth 5069 (IMI
251487!). Scotland, Kintyre, c. 7km SW of Crinan, on islet N of Carsaig Island, on Lobaria
pulmonaria, 14 August 1977, N. Brandt (E!, IMI 224707!; with Endophragmiella hughesii). Italy:
Siidtirol, Otztaler Alpen, Vinschgau, on Rhizoplaca chrysoleuca, 18 October 1975,/. Hafellner (GZU!;
with Lichenoconium lecanorae). Switzerland: Wallis, Aletschwald ob Brig, on P. glabratula subsp.
fuliginosa, 1 1 September 1968, J. Poelt (hb. Poelt 6359!; with L. usneae).

V. EPICLADONIA D. Hawksw. gen. nov.

Genus lichenicola ad Coelomycetes Sphaeropsidales pertinens. Conidiomata singularia, uniloculata,
subglobosa, ad cupuliformia, atrobrunnea immersa ad erumpescentia, cum muris textura intricatis,
subhyalinis sed atrobrunneis in parte exposito, ostiolata. Conidiophora plerumque desunt. Cellulae
conidiogenae holoblasticae, acrogenae, subcylindrices ad elongato-ampulliformes, prolifericae,
annellatae, hyalinae. Conidia subcylindrica ad anguste-ellipsoidea vel cymbiformia, hyalina, 0-1
septata, laevia.

Conidiomata pycnidial (eustromatic), arising singly, scattered, subglobose to cupuliform,
immersed to erumpent, dark brown; walls composed of compacted entwined short-celled
hyphae forming a textura intricata, subhyaline except in the exposed parts, ostiolate.
Conidiophores usually absent. Conidiogenous cells holoblastic, lining the inner wall of the
pycnidial cavity, acrogenous, subcylindrical to elongate-ampulliform, proliferating,
annellate, usually unbranched, hyaline. Conidia arising singly, not catenate, subcylindrical
to narrowly ellipsoid or cymbiform, hyaline, 0-1 septate, smooth-walled.

Type species: Epicladonia sandstedei (Zopf) D. Hawksw. (syn. Diplodina sandstedei
Zopf holotypus).

Number of species: Three, all of which are only known from Cladonia species on which
they commonly form galls.

16 D. L. HAWKSWORTH

Observations: Epidadonia is distinguished from Bachmanniomyces, which also forms
galls on Cladonia species (p. 10), in that the exposed parts of the pycnidia are dark brown to
black, never shades of orange. Furthermore, the conidia are never pyriform nor strongly
adhering in mucilage, 1 -septate in two species, and have a much broader scar than is seen in
B. uncialicola. In Pseudoseptoria usneae (p. 57), the walls of the pycnidia are of
greenish-brown pseudoparenchymatous cells (textura angularis) and the conidiogenous cells
tend to be relatively short and stout.

The three species referred to this new genus here are separated primarily on the basis of
conidial size and septation and further, for E. simplex, the degree of emergence and extent of
consequent pigmentation of the pycnidial wall.

It is conceivable that Verrucaster Tobler might provide an earlier name for this genus, but
in the absence of further information on the type species (see p. 88), I feel it is not possible
to take it up here with any degree of confidence.

Key to the species

1 Pycnidia mainly less than 100//m diam, immersed, brown only immediately around the

ostiole 2

Pycnidia 100-175 //m diam, erumpent and becoming almost superficial, the exposed parts
all brown; forming galls; conidia simple, subcylindrical, rounded at the apex, 8'5-l 1(-12)
x2-5-3(-3'5)//m Epicladonia simplex (p. 19)

2 Conidia predominantly 1 -septate, subcylindrical to narrowly ellipsoid, apex always

rounded, (7'5-)9-12(-14) x (2'5-)3-4 //m; usually forming galls Epicladonia sandstedei (p. 16)

- Conidia predominantly simple, subcylindrical to almost cymbiform, apex often attenuated,

7*5-1 1 x 3-3'5//m; not forming galls Epicladonia stenospora (p. 20)

1. Epicladonia sandstedei (Zopf) D. Hawksw. comb. nov.
(Figs. 5A-B, 6A-E)

Diplodina sandstedei Zopf, in Sandstede, Abh. naturw. Ver. Bremen 18 : 429 (1906).

Type: Germany, Oldenburg, Westerstede, 'Erdwalle in der Waldung "Hassen" bei Keilburg', on

Cladonia caespiticia (Pers.) Florke [?], H. Sandstede.
? Diplodina claudelii Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 289 (19 14); as ' 'claudeli '.

Type: France, Vosges, on Cladonia sp., V. Claudel.

Exsiccatae*: Krypt. Exs. Vindob. no. 13306 (sub Diplodina sandstedei; B!, K!). [The fungus was not
found on copies of no. 1 330[<a] studied (B!, K!).]

Icones: Keissler, Rabenh. Krypt.-Fl. 8 : 18 fig. 1 1, 19 fig. 13, 25 fig. 20, 570 figs. 109-1 1 1 (1930).

Conidiomata pycnidial, normally immersed in galls; galls convex, verruciform, to c. 1 mm
diam, often somewhat constricted at the base, 5-10(-15) pycnidia in each gall, galls
concolorous with the host tissue, not developed in early stages of infection when mature
pycnidia can arise directly on the podetia; pycnidia arising singly, not aggregated into
stromata, evenly scattered over the galls, immersed, the ostiole and upper part of the
pycnidium slightly erumpent, translucent brown to almost black when mature, subglobose at
first but becoming gaping and almost cupuliform with age, 50-80(-125)//m diam; ostiole
clearly delimited in younger pycnidia, depressed, circular, to c. 50 //m diam, centre
appearing subhyaline due to the conidial mass, formed by a breakdown of the upper wall of
the pycnidium, irregular in shape in older pycnidia; pycnidial walls subhyaline except in
the vicinity of the ostiole, there olivaceous brown to dark brown, mainly 8-15 //m thick, to
20 //m thick near the ostiole, wall throughout composed of densely interwoven moderately
thick-walled hyphae, these short-celled and mainly 2-5-4-5 //m wide, textura intricata but

*Sandstede (1931) lists 24 exsiccatae numbers from seven sets on which he found this fungus. These are excluded as
they were issued for the host Cladonia arid the fungus is probably absent from some copies of these numbers.

LICHENICOLOUS COELOMYCETES
'41

17

Fig. 5A-B, Epicladonia sandstedei (IMI 240228); A, infected podetium with galls (x 12'5);
B, vertical section of pycnidium (x 500). C, E. stenospora (Dobbeler 1827), infected
squamule showing decolorization and pycnidia (x 20).

appearing almost cellular in parts. Conidiophores absent or short-cylindrical, simple or
sparsely branched at the base, hyaline, variable in length, to 25 /um tall, 3-4 /am wide.
Conidiogenous cells holoblastic, arising directly from the pycnidial wall or integrated into
the conidiophores, subcylindrical to elongate-ampulliform, tapering towards the apex,
hyaline, smooth-walled, annellidic, with 0-3 annexations, mainly 10-20//m tall, 3-4 /zm
wide at the base, mainly 1 -5-2-5 //m wide at the apex. Conidia abundantly produced,
arising singly, not catenate, subcylindrical to narrowly ellipsoid, rounded at the apex,
truncated at the base with a scar l - 5-2 /urn wide (remnants of wall tissue sometimes evident
at high magnifications), hyaline, (0-)1 septate when mature, 2-3 guttulate, thin-walled,
smooth-walled, (7'5-)9-12(-14) x (2'5-)3-4 /zm.

Hosts: Cladonia species, squamules and podetia, usually forming irregularly lobed bullate
galls which may be hollow inside. Probably parasymbiotic. I have not made an exhaustive
study of Cladonia specimens in search of the fungus but according to Sandstede (1931) it has
a very wide host range. Sandstede mentioned it on C. anomaea (Ach.) Ahti & P. James, C.
carneola (Fr.) Fr., C. coniocraea auct., C. chlorophaea (Florke ex Sommerf.) Sprengel 5. lat.,

18

D. L. HAWKSWORTH

lOjim

Fig. 6 Epicladonia sandstedei. A-C (IMI 240228); A, vertical section of pycnidial wall; B,
conidiogenous cells; C, conidia. D (Krypt. Exs. Vindob. no. 13306, K), conidia. E-F
(1893, Harmand, Angers); E, conidiogenous cell; F, conidia.

C. cornuta (L.) Hoffrn., C. cyanipes (Sommerf.) Vainio, C. deformis (L.) Hoffrn., C.fimbriata
(L.) Fr. (incl. C. major (Hagen) Sandst.), C. grayi G. K. Merrill ex Sandst., C. ochrochlora
Florke, C. pleurota (Florke) Schaerer, C. pyxidata (L.) Hoffm., C. subulata (L.) Wigg., and C.
turgida Ehrh. ex Hoffm.

His report from C. caespitica is probably an error for C. glauca Florke (see below), and
at least one on C. pyxidata probably refers to C. merochlorophaea Asah. (see below); some

LICHENICOLOUS COELOMYCETES 19

other reports may also be due to mis-identified hosts or lichenicolous fungi. However, in
addition, the fungus occurs also on at least C. conoidea Ahti (see below).

Distribution: Evidently widespread in Europe. Reliably reported from at least the British
Isles (England), France, Germany, Sweden and Switzerland.

Observations: This fungus is easily distinguished from the other two species referred to the
genus here on the basis of the predominantly 1 -septate conidia.

The original material studied by Zopf could not be located in B (B. Hein, in litt.) but
specimens from the type locality were distributed as Krypt. Exs. Vindob. no. 1330[a] (on
Cladonia glauca Florke*; B!, H!, K!). Specimens on C. cf. merochlorophaea Asah., collected
by Sandstede, were also issued as no. 13306*. No copy of 1330[a] seen had good material of
the fungus, but as this exsiccatum would be ideal for neotypification, and the possibility that
another copy with it well-developed exists, no neotypification is proposed here. The fungus
on no. 13306, however, conforms to the usual concept oi Diplodina sandstedei and so there
is little doubt as to the correct application of the epithet.

Diplodina claudelii is not present amongst the remnants of Vouaux's herbarium (Rondon,
1970), nor could it be found in Claudel and Harmand's material in Angers (M. Guerlesquin,
in litt.}. This name was based on a fungus on the squamules of an unidentified Cladonia with
pycnidia 90-1 50 //m diam, branched conidiophores with terminal conidia, conidiogenous
cells 14-20 x 3 //m and 1 -septate conidia truncated at the base and rounded at the apex
which measured 10-13 x3-3'5 /^m. There is consequently little doubt that this fungus
should be placed here. The absence of galls does not preclude this possibility as these are not
formed in early stages of infection; pycnidia in galls and directly on podetia can occur in the
same collection (e.g. IMI 240228). As 1 -septate conidia were reported, it is unlikely that
Vouaux was dealing with Epicladonia stenospora but that possibility cannot be
entirely excluded as 2-guttulate conidia may appear septate if not studied carefully.

I have not seen conidia over 1 1-5 //m in E. sandstedei, but Zopf (loc. cit.) originally gave
them as 9-14 x 3-5-4 //m.

Keissler (1930 : 571) placed Diplodina peltigerae as a synonym of D. sandstedei, perhaps
because of the similar conidial dimensions, but in the absence of material of the former and
the disparate hosts such a treatment cannot be accepted here (see p. 76).

Specimens: British Isles: England, W. Sussex, Pagham Harbour Nature Reserve, on Cladonia
conoidea on soil amongst shingle, 30 August 1972, B. J. Coppins (IMI 240228!). France: Docelles, on
C. chlorophaea s. lat., 1893, J. Harmand (Angers!). Germany: See under Exsiccatae.

2. Epicladonia simplex D. Hawksw. sp. nov.
(Figs. 7A-B, 8A-B)

Similis Epicladonia stenospora (Harmand) D. Hawksw. sed difiert in pycnidiis erumpescentibus vel
superficialbus, 100-1 75 //m diam, et conidiis semper simpliciibus, 8'5-l 1(-12) x 2'5-3(-3'5) //m.

Typus: Finlandia,? Lapponia, in Cladoniae bacilliformes (Nyl.) Dalla Torre & Sarnth., c. 1925, V.
Kujala (H holotypus!).

Conidiomata pycnidial, usually on galls formed on squamules or podetia of the host, galls
convex, tuberculate, to c. 1 mm diam; pycnidia immersed at first but erumpent and almost
superficial at maturity, arising singly, not aggregated into stromata, scattered, dark brown,
subglobose to cupuliform, 100-175 //m diam; ostiole clearly delimited, depressed, gaping to
reveal the subhyaline conidia inside and so appearing whitish centrally; pycnidial wall
subhyaline in the parts sunken into the galls, but brown to dark brown in the exposed parts,
mainly \0-\5 urn thick, composed of densely interwoven short-celled moderately thick-
walled hyphae mainly 2'5-3'5 /*m wide, textura intricata, but becoming almost cellular in
parts. Conidiogenous cells holoblastic, arising from the pycnidial wall, subcylindrical

*Det. T. Ahti; the original host and material in no. 1330[a] was erroneously named as C. caespiticia (Pers.) Florke,
and in no. 1 3306 as C. pyxidaia (L.) Hoffm., by Sandstede.

D. L. HAWKSWORTH

B

*

f

41

: t i

1

"+~ t *

, '**

f

-*.

- * *

-J-

f

^^ .*

|

X

"

/ V V

1 4

a^^-'*

ii

Fig. 7 Epicladonia simplex (H holotype). A, Infected podetia showing pycnidia arising
from galls and directly on the podetia (x 12'5). B, Vertical section of pycnidium (x 500).

to elongate ampulliform, only slightly tapered towards the apex, annellate with to 3
annellations, hyaline, smooth-walled, 7-15 x 2-5-3-5 //m. Conidia arising singly, not
catenate, subcylindrical, always rounded at the apex, truncated at the base with a scar c.
1'5-2/im wide, always simple, 2 or more guttulate, thin-walled, smooth-walled, 8'5-
ll(-12)x2-5-3(-3-5)//m.

Host: Cladonia bacilliformis, usually forming galls on both the primary squamules and
podetia of the host. Probably parasymbiotic.

Distribution: Finland. Known only from the original collection.

Observations: This species is most similar to Epicladonia stenospora (see below) from
which it is distinguished by the generally larger and erumpent to almost superficial pycnidia,
the more extensively pigmented and more cellular appearance of the pycnidial wall, the
shape of the conidia, and further in giving rise to the formation of galls, although these are
not formed immediately (Fig. 7A).

The annellations on the conidiogenous cells are more distinct in this species than in E.
sandstedei and none were noted in E. stenospora. Indeed, one might interpret the
conidiogenous cells of E. simplex as proliferating phialides rather than annellides. However,
I am sceptical as to the merits of such characters at the genus level (see p. 4) and so place this
species here.

3. Epicladonia stenospora (Harmand) D. Hawksw. comb. nov.
(Figs. 5C, 9A-B)

Aposphaeria stenospora Harmand, Lich. Fr. 3 : 308 (1907).

Conidiomata pycnidial, immersed in squamules, not associated with galls, arising singly,
not aggregated into stromata, scattered, the ostiole slightly erumpent or depressed, brown,
subglobose, 75-1 20 /zm diam; ostiole clearly delimited at first but later tending to gape;
pycnidial wall hyaline except in the vicinity of the ostiole where it is olivaceous brown to
dark brown, mainly 7-1 5 //m thick but to 25 //m thick near the ostiole, wall mainly
composed of densely interwoven hyphae mainly 2-3 /zm wide, textura intricata but
becoming almost cellular in the vicinity of the ostiole and there 4-6 um diam.
Conidiogenous cells holoblastic, arising from the pycnidial wall, elongate-ampulliform,

LICHENICOLOUS COELOMYCETES

21

10/im

Fig. 8 Epidadonia simplex (H holotype). A, Vertical section of pycnidial wall. B, Conidiogenous

cells. C, Conidia.

strongly tapered towards the apex, hyaline, smooth-walled, no annellations seen, mainly
7-12 x 3-3*5 //m. Conidia arising singly, not catenate, subcylindrical to almost cymbiform,
often rather irregular in shape, the apex rounded to slightly attenuated and the base
narrowed and abruptly truncated by a scar c. 1-1 -5 //m wide, simple or exceptionally
1 -septate, frequently 2 or more guttulate, thin-walled, smooth-walled, 7 - 5-l 1 x 3-3'5 //m.

Hosts: Cladonia cf. coniocraea auct. and C. pyxidata (L.) Hoffm. [?] (see below). Occurring
on the primary squamules in the specimen from Austria examined; no gall-like deformations
are produced, but the infected tissue eventually becomes decolourized to give circular pale
brownish necrotic infection spots on the host squamules (Fig. 5C). Probably parasitic.

Distribution: Austria and France.

Observations: Despite the lack of annellations this fungus is referred to Epidadonia in
view of the very similar structure of the pycnidia and the occasional presence of 1 -septate
conidia. Further, the presence of distinct annellations in E. simplex which consistently has
simple conidia and further forms galls on Cladonia adds weight to this taxonomy. The
separation of E. stenospora from E. simplex is discussed under the latter species (p. 20).

In the original description of Aposphaeria stenospora, the host was named as Cladonia
pyxidata but no locality details were provided. The original material could not be located in
Angers (M. Guerlesquin, in lift.), but as the conidia were described as 8-9 x2-2'5//m it is
probable that the name referred to a fungus such as that of Dobbeler's (see below), on which
the above description is based. The Austrian specimen is not designated as a neotype for
Harmand's name as it is rather sparse.

Specimen: Austria: Tirol, Ha'nge unmittelbar ostlich des Achensees in der Nahe des Achenseehofes,
on Cladonia cf. coniocraea, 13 October 1976, P. Dobbeler 1827 (GZU!).

22

D. L. HAWKSWORTH

10/im

Fig. 9 Epicladonia stenospora (Dobbeler 1827). A, Vertical section of pycnidial wall with

conidiogenous cells. B, Conidia.

VI. KARSTENIOMYCES D. Hawksw.
Trans. Br. mycol. Soc. 74 : 371 (1980).

Conidiomata pycnidial, arising singly, scattered or loosely aggregated, subglobose,
superficial, nectrioid, translucent pale orange to deep red, ostiole irregular and forming
schizogenously; walls thick, composed of thick-walled pseudosclerenchymatous cells with
much-reduced lumina. Conidiophores cylindrical, sympodially branched. Conidiogenous
cells holoblastic, arising acrogenously or pleurogenously, cylindrical, hyaline. Conidia
probably dry, elongate-ellipsoid, the apex rounded and the base slightly to clearly truncate,
hyaline, 1 -septate, smooth-walled, guttulate.

Type species: Karsteniomyces peltigerae (P. Karsten) D. Hawksw. (syn. Stagonopsis
peltigerae P. Karsten).

Number of species: Monotypic.

Observations: This genus appears to occupy an isolated position in the Sphaeropsidales
and is unlikely to be confused with any other. Stagonopsis Sacc. (type species S. pallida
(Berk. & M. A. Curtis) Sacc.) is of somewhat uncertain application, but as the type species
was described as forming arcuate 7-8 septate conidia, it is most unlikely that it has anything
to do with Karsteniomyces.

This new genus is named in honour of Petter A. Karsten (1834-1917) who made very
substantial contributions to our knowledge of microfungi in Finland, including the discovery
of several previously unrecognized lichenicolous species.

1. Karsteniomyces peltigerae (P. Karsten) D. Hawksw., Trans. Br. mycol. Soc. 74:371

(1980).
(Figs. 10A-D, 11A-E)

Stagonopsis peltigerae P. Karsten, Meddel. Soc. Fauna Flora fenn. 14 : 106 (1 887).
Type: Finland, Tavastia australis, Tammela, Mustiala, on Peltigera rufescens, July 1868, H. A.
Kullhem (H-KARST 2041 holotype!).

Conidiomata pycnidial, almost entirely superficial with only the base entering the host
tissues, arising singly but often numerous and sometimes crowded together, globose,
translucent pale orange at first, becoming a dark orange red with age especially around

LICHENICOLOUS COELOMYCETES

B ; Af

Fig. 10 Karsteniomyces peltigerae (H-KARST 2041 holotype). A, Pycnidia on the host
thallus (x25). B, As A showing the wide ostioles (x 50). C, Vertical section of pycnidium
(x 250). D, Vertical section of pycnidial wall showing the thick-walled almost sclerenchymatous
cells (x 1020).

the ostiole, mainly 75-200 //m diam but exceptionally larger, appearing nectriaceous,
opening by an irregular schizogenously formed ostiole; pycnidial wall very variable in thick-
ness, mainly 20-50 //m thick, composed of two layers, the outer comprising hyaline very
thick-walled pseudosclerenchymatous cells with extremely narrow compressed lumina, the
outer layer sometimes almost absent, very uneven externally, 5-20 /am thick, the inner
layer also pseudosclerenchymatous but formed of cells with larger lumina and pigmented
pale orange, 10-25 jum thick. Conidiophores arising from the inner walls of the pycnidial
cavity, cylindrical, irregularly sympodially branched, hyaline, smooth-walled, 5-15 jum tall
and 2-3(-4)//m wide. Conidiogenous cells arising terminally and laterally from the
conidiophores, indistinguishable from the conidiophores except in the absence of septa.
Conidia holoblastic, abundant, dry or at least not adhering, elongate-ellipsoid, the apex
rounded and the base slightly to clearly truncate, hyaline, 1 -septate when mature, the
septum thin and sometimes undulate, 1^ guttulate, smooth-walled, (12-)15-22(-24)x
3-5-5(-6)/zm.

24

D. L. HAWKSWORTH

Fig. 11 Karsteniomyces peltigerae (H-KARST 2041 holotype). A, Vertical section of
pycnidium. B, Cells from the inner part of the pycnidial wall. C, Cells from the outer
part of the pycnidial wall. D, Conidiophores and conidiogenous cells. E, Conidia.

Host: Peltigera canina (L.) Willd. and P. rufescens (Weis) Humb., thalli. The type
specimen was originally stated to be on P. canina but that host has been redetermined by Mr
O. Vitikainen as P. rufescens. The species is, however, correctly reported here from P.
canina (see Additional specimens). The infected thalli do not appear to be very significantly
damaged by Karsteniomyces peltigerae and no clearly demarcated decolourized patches are
produced.

LICHENICOLOUS COELOMYCETES 25

Distribution: Finland and Norway. Keissler (1930 : 585) mentions two dubious records
from Germany and Italy but these are in need of confirmation.

Observations: The conidia have been described as becoming up to 4-septate (Keissler,
1930:585) but I have only seen 1 -septate conidia. I strongly suspect that reports of
additional septa are due to misinterpretations of the guttules, which are often conspicuous.

Additional specimens: Finland: Regio aboensis, Kakskerta Kokkila, on Peltigera canina and P.
rufescens, 4 August 1934, L. E. Kari (H two packets!). Norway: Akershus, Oslo (Christiania),
Disen, on P. canina, 25 March 1869, N. G. Moe 145 (UPS!).

VII. KEISSLERIOMYCES D. Hawksw. gen. nov.

Genus lichenicola ad Coelomycetes Sphaeropsidales pertinens. Conidomata singularia, uniloculata,
dispersa, subglobosa, immersa, nigra, ostiolata, cum muris tenuibus et textura angulari compressis.
Conidiophora desunt. Cellulae conidiogenae enteroblasticae, subglobosae vel breve ampulliformes,
polyphialidicae, hyalinae. Conidia probaliter sicca, late fusiformia, basi leviter arcuata, hyalina,
3-septata, laevia.

Conidiomata pycnidial arising singly, scattered, subglobose, immersed, black, ostiolate;
walls thin, composed of radially compressed polyhedral dark brown cells, textura angularis.
Conidiophores absent. Conidiogenous cells enteroblastic, subglobose or short-ampulliform,
polyphialidic, each giving rise to up to 3 conidia, hyaline. Conidia probably dry, broadly
fusiform, the base weakly curved, hyaline, 3-septate, smooth-walled.

Type species: Keissleriomyces sandstedeanus (Keissler) D. Hawksw. (syn. Stagonospora
sandstedeana Keissler holotypus).

Number of species: Monotypic.

Observations: The genus Stagonospora (Sacc.) Sacc. nom. cons, (type species S. paludosa
(Sacc. & Speg.) Sacc.) is characterized by conidiogenous cells forming conidia holo-
blastically and not polyphialides; S. sandstedeana consequently does not belong there.
Keissler's choice of Stagonospora was most probably due to the rather similar conidia seen
in some species of that genus. Polyphialidic conidiogenous cells are rather rare in
Coelomycetes and Keissleriomyces has little else in common with other genera exhibiting
this feature.

Although with hindsight many of Karl von Keissler's (1872-1965) taxonomic decisions
amongst the lichenicolous fungi prove to be unsatisfactory, his compendium of the central
European species (Keissler, 1930) remains an essential reference work to all students of
lichenicolous fungi today; it is in recognition of this debt to him that the new generic name
Keissleriomyces has been introduced here.

1. Keissleriomyces sandstedeanus (Keissler) D. Hawksw. comb. nov.
(Figs. 12, 13A-E)

Stagonospora sandstedeana, Keissler, Annln naturhist. Mus. Wein 38 : 166 (1925).
Type: Czechoslovakia (Bohemia), Riesengebirge, Schneekoppe, on Cladonia furcata, O. L. A.
Ohlert(VJ 1921/185 holotype!).

Conidiomata pycnidial, almost entirely immersed in the host tissues, arising singly,
scattered, subglobose but often radially compressed, black, 40-75(-80)^m diam, ostiolate;
pycnidial wall 7-10/zm thick, composed of usually two layers of radially compressed
polyhedral pseudoparenchymatous cells, cells dark brown, moderately thick-walled,
10-14 x 3-4 //m in vertical section and mainly 6-9 //m diam in surface view. Conidio-
genous cells enteroblastic, lining the inner wall of the pycnidial cavity except near the apex,
subglobose or shortly ampulliform with swollen bases, 5-8 x 5-6 //m, polyphialidic, each
cell with 1-3 phialides, hyaline. Conidia abundant, probably dry, broadly fusiform with the
base generally more attenuated and frequently weakly curved, hyaline, 3-septate when

26

D. L. HAWKSWORTH

i <*:

Fig. 12 Keissleriomyces sandstedeanus (W 1921/185 holotype), vertical section of pycnidium

(x 1020).

mature, not markedly guttulate, contents rather granular, smooth-walled, 18-22x
3-5-5 //m.

Host: Cladonia furcata (Huds.) Schrader, podetia. In the type material the pycnidia are
mainly located on the upper part of the podetia, the infected part of which is slightly
decolourized but not deformed. Keissleriomyces sandstedeanus consequently appears to be
only a weak parasite or opportunist.

Distribution: Czechoslovakia. Only known from the type collection.

Observations: The characteristic polyphialides are otherwise unknown amongst the
lichenicolous Coelomycetes, and no other fungus forming similar spores is recorded from
any Cladonia species.

VIII. LAEVIOMYCES D. Hawksw. gen. nov.

Genus lichenicola ad Coelomycetes Sphaeropsidales pertinens. Conidiomata singularia, pycnidiifor-
mia, uniloculata, dispersa, subglobosa vel irregularia, erumpescentia, nigra, cum muris plusminusve
hyalinis e cellulis pseudoparenchymaticis (sed in unica textura intricatis) et ostiolis irregulariter
dehiscentibus. Conidiophora desunt. Cellulae conidiogenae holoblasticae, acrogenae, subcylindrices
vel elongato-ampulliformes, prolifericae, annellatae, fuscae vel brunneae, laeves. Conidia subglobosa
ad ellipsoidea, basi truncata, pallide brunnea, simplicia, laevia.

Conidiomata pycnidial, arising singly, uniloculate, scattered, subglobose or irregular with
somewhat convoluted internal walls, erumpent, black; walls composed of several layers of
irregularly polyhedral to subglobose hyaline cells forming a textura angularis (but in one
species reduced and hyphal, textura intricata); ostiole irregular in shape and gaping in age.
Conidiophores absent. Conidiogenous cells holoblastic, lining the inner wall of the pycnidial
cavity, acrogenous, subcylindrical to elongate-ampulliform, proliferating, annellate, fuscous
brown to brown, smooth-walled. Conidia subglobose to ellipsoid, abruptly truncated at the
base, a small basal frill sometimes discernible, not catenate, brown, simple, smooth-walled.

LICHENICOLOUS COELOMYCETES

27

10/im

Fig. 13 Keissleriomyces sandstedeanus (W 1921/185 holotype). A, Vertical section of
pycnidium. B, Vertical section of pycnidium wall. C, Surface view of pycnidial wall. D,
Conidiogenous cells. E, Conidia.

Type species: Laeviomyces pertusariicola (Nyl.) D. Hawksw. (syn. Spilomium pertusarii-
cola Nyl. holotypus).

Number of species: Two, both lichenicolous.

Observations: The type species of this genus was retained in Lichenoconium as an interim
measure by Hawksworth (1977) because a suitable generic name was considered to possibly
exist amongst the non-lichenicolous Coelomycetes. Subsequent studies have failed to reveal
any suitable generic name and so the genus Laeviomyces is introduced here to accommodate
it. The generic name recalls the smooth-walled conidia (x 8000), but L. pertusariicola further
differs from Lichenoconium in the massive pycnidia, hyaline and relatively flaccid pycnidial
wall, the consistently brown and smooth-walled conidiogenous cells, and individually much
less strongly pigmented conidia, sometimes with a recognizable basal frill.

Farr et al. (1979 : 1649) listed Spilomium pertusariicola Nyl. as the type species of
Spilomium Nyl. However, they give the place of publication of the latter as Nylander
(1858 : 91) when in fact the genus must be considered as already validly published earlier by
Nylander (1856 : 337), who made no mention of S. pertusariicola although referring three
other species to Spilomium. Nylander's generic name is correctly interpreted as a
superfluous name for the hyphomycete genus Sclerococcum Fr. ex Fr., as already pointed out
by Hawksworth (197 5a : 222).

Laeviomyces could be viewed as a counterpart of Lichenodiplis with simple as opposed to
1 -septate conidia. In view of the more complex wall structure of Laeviomyces pertusariicola,

28 D. L. HAWKSWORTH

'*"i

Fig. 14 Laeviomyces opegraphae (E holotype), vertical section of Opegrapha niveoatra
lirellum with the hymenium partly replaced by a pycnidium of the Laeviomyces (x 500).

and the emphasis placed on spore septation in traditional coelomycete systematics, it seems
preferrable to retain two genera while intermediate taxa are unknown.

Key to the species

1 Pycnidia 40-60 ^m wide; pycnidial wall poorly developed and hyphal; conidia olivaceous

brown, 3-4*5 x 1 -5-2 /zm Laeviomyces opegraphae (p. 28)

Pycnidia 100-250(-325)//m diam; pycnidial wall well developed and pseudoparen-
chymatous; conidia reddish brown, 3*5-6 x 2*5-3*5 /zm . Laeviomyces pertusariicola (p. 29)

1. Laeviomyces opegraphae D. Hawksw. sp. nov.
(Figs. 14, 15A-B)

Similis Laeviomycei pertusariicola (Nyl.) D. Hawksw. sed differt in pycnidiis 40-60 //m latis, muris
rudimentalibus e textura intricata, et conidiis olivaceo-brunneis, 3^-5 x 1-5-2 jj.m.

Typus: Magna Britannica, Scotia, Dumbarton, Loch Lomond, Gartocharn, Claddochside, in thallo
Opegraphae niveoatrae (Borrer) Laundon, 3.iii. 1980, B. J. Coppins 4675 (E holotypus!).

Conidiomata pycnidial, immersed in the thallus or lirellae of the host, arising singly,
scattered, sometimes becoming confluent in heavy infections, brown, 40-60 /zm wide, to
100/zm tall when in lirellae, at first globose but becoming very irregular in shape, ostiole
forming by an irregular breakdown of the upper wall of the pycnidium, becoming gaping and
the pycnidia sometimes almost cupuliform or melanconiaceous with age; pycnidial wall
poorly delimited, composed of 2-3 layers of intertwined short-celled hyphae forming a layer
only 3-5 //m thick, hyphae olivaceous, sometimes becoming almost cellular just below the
conidiogenous cells, subhyaline to pale olivaceous brown, 1*5-2*5 //m thick. Conidio-
genous cells holoblastic, acrogenous, subcylindrical, proliferating, distinctly annellate with
to 6 annellations, olivaceous brown, smooth-walled, 4-7(-9)x I '5-2*5 /*m. Conidia dry,
ellipsoid, narrowed to an abruptly truncated base, basal frill sometimes visible, olivaceous
brown, simple, not conspicuously guttulate, smooth-walled, 3-4*5 x 1-5-2 //m.

Host: Opegrapha niveoatra (Borrer) Laundon, thallus and hymenium of the lirellae. When
on the thallus little damage occurs, but when the lirellae become infected almost the whole
of the hymenium may eventually be occupied by the fungus (Fig. 14), thus precluding the
possibility of ascospore formation.

LICHENICOLOUS COELOMYCETES

B

29

lOfim

Fig. 15A-B, Laeviomyces opegraphae (E holotype); A, conidiogenous cells; B, conidia.

C, L. pertusariicola (IMI 1 86240), conidia.

Distribution: British Isles (Scotland). Evidently not uncommon in Scotland on this host
and to be expected throughout the range of the host lichen.

Observations: This new species is included within Laeviomyces with some hesitation as
the pycnidial wall is so poorly developed and almost hyphal, rather than pseudoparen-
chymatous as in L. pertusariicola. However, the pigmented and annellate conidiogenous
cells and shape of the conidia are so similar in these two fungi that it seems more appropriate
to view L. opegraphae as a species in which the wall has been reduced. It would not be
prudent to describe a new genus based only on this single feature in our present state of
knowledge of the lichenicolous Coelomycetes.

Additional specimens: British Isles: Scotland, Midlothian, Dalkeith Oak Wood, on Opegrapha
niveoatra on Acer, 25 February 1977, B. J. Coppins & C. Pope 4696 (E!); Peebles, Dawyck Arboretum,
on O. cf. niveoatra on A. pseudoplatanus, 10 May 1980, B. J. Coppins 4860 (E!).

2. Laeviomyces pertusariicola (Nyl.) D. Hawksw. comb. nov.
(Fig. 15C)

Spilomium pertusariicola Nyl., Mem. Soc. Imp. Sci. nat. Cherbourg 5 : 91 (1858); as "pertusaricola\
Lichenoconium pertusariicola (Nyl.) D. Hawksw., Trans. Br. mycol. Soc. 65 : 233 (1975).

For description, fuller illustrations, and further synonyms see Hawksworth (\915a:
233-234, 1977 : 182-183).

Hosts: Buellia disciformis (Fr.) Mudd, Pertusaria leioplaca DC. and P. pertusa (Weigel)
Tuck., thalli. Apparently parasymbiotic, not damaging the host thalli.

Distribution: British Isles, Canada, Denmark, France, Germany and Spain. There is also
an unconfirmed report from New Caledonia (Keissler, 1933 : 387).

Observations: In the Canadian collection on Buellia disciformis, a host now confirmed for
this fungus, the conidia tend to be slightly longer (to 7*5 //m) and rather more deeply
pigmented than is usual for the species; further collections on that host may show that it
merits recognition as a separate taxon. Muellerella lichenicola (Sommerf. ex Fr.) D.
Hawksw. is also present in the collection on this Buellia and intimately mixed with the
Laeviomyces.

Lichenodiplis lecanorae, which can also occur on corticolous Pertusaria species, is easily
separated from Laeviomyces pertusariicola by the consistently 1 -septate and generally paler
rather more cylindrical and longer conidia, which have an even more strongly truncated
base, as well as by the much smaller pycnidia.

Specimens (additional to those listed by Hawksworth, 1975a, 1977): British Isles: Scotland,
Dunbartonshire, Loch Lomond, Ross Priory, on Pertusaria leioplaca on Acer, 13 March 1980, B. J.
Coppins 4664 (E, IMI 247022!); Wales, Breconshire, Erw Fawr, above Elan Valley, on P. cf. leioplaca

30 D. L. HAWKSWORTH

on Quercus, 20 October 1979, F. Rose (IMI 244540!). Canada: Nova Scotia, Halifax, South Park, on
Buellia disciformis on Acer, April 1979, H. J. M. Bowen (E, IMI 244538!).

IX. LIBERTIELLA Speg. & Roum.

in Spegazzini, Revue mycol. 2 : 22 (1880).
Nicholsoniella Kuntze, Rev. Gen. PI. 2 : 862 (1891).
Shecutia Nieuwl., 4m. Midi Nat. 4 : 379 (1916).

Conidiomata pycnidial, arising singly or in small groups, often aggregated, subglobose,
superficial, whitish to pale orange, ostiole forming by a disintegration of the upper wall within
an area delimited by a reddish-brown ring of tissue; walls composed of almost hyaline very
thick-walled pseudoparenchymatous cells, often interspersed with orange granules. Conidio-
phores hyaline, irregularly sympodially branched, short, cylindrical. Conidiogenous
cells enteroblastic, phialidic, arising laterally or terminally from the conidiophores, rarely
proliferating. Conidia adhering in gelatinous masses, ellipsoid to obovoid with a slightly
truncated base, hyaline, simple, smooth-walled.

Type species: Libertiella malmedyensis Speg. & Roum.

Number of species: Monotypic. Other species previously referred to this genus are now
placed in different genera (see Index).

Observations: Both Nicholsoniella Kuntze and Shecutia Nieuwl. were introduced as new
names for Libertiella in the belief that Libertiella was a homonym of the earlier Libertella
Desm.; Nieuwland (1916) was evidently unaware of the new name introduced by Kuntze.
However, as pointed out by Sutton (1977 : 113), in coining the new generic name Libertiella,
Spegazzini and Roumeguere deliberately inserted the T and made it clear in a footnote that
they were doing this to prevent confusion with Libertella Desm. The generic name
Libertiella is consequently not to be treated as a later homonym of Libertella and can be
retained.

1. Libertiella malmedyensis Speg. & Roum., in Spegazzini, Revue mycol. 2 : 22 (January

1880).
(Figs. 16A-B, 17A-E)

Type: Belgium, 'prope malmedyanum', on Peltigera spuria (Ach.) DC., 'Hieme', M.-A. Libert

(BR holotype, non vidi; K isotype!).

Nicholsoniella malmedyensis (Speg. & Roum.) Kuntze, Rev. Gen. PI. 2 : 863 (1891).
Shecutia malmedyensis (Speg. & Roum.) Nieuwl. Am. Midi. Nat. 4 : 379 (1916).
Zythia peltigerae Lib. ex Cooke, Grevillea 8 : 83 (March 1 880), nom. illegit. (Art. 63).
Type: Belgium, [Malmedy], on Peltigera spuria (Ach.) DC., M.-A. Libert (BR holotype, non vidi;

K isotype!).
Libertiella peltigerae (Lib. ex Cooke) Keissler, Rabenh. Krypt.-Fl. 8 : 582 (1930).

Exsiccatae: Roumeguere, Fungi set. Gall. no. 671 (K!). Thiimen, Mycoth. univ. no. 1775 (K!).

Conidiomata pycnidial, superficial, with only the base immersed in the host, arising
singly or more commonly in groups, often aggregated, subglobose, whitish to pale orange,
mainly (100-)150-200(-300),um diam, becoming flattened at the apex with the develop-
ment of the ostiole; ostiole forming after the pycnidial cavity is full of spores, demarcated by
a limiting reddish-brown ring of tissue about 25 um wide encircling the whitish 50-75 //m
diam pore; pycnidial wall mainly 10-12 //m thick, composed of hyaline translucent
thick-walled pseudosclerenchymatous cells, cells mainly 3-4*5 //m thick, walls of the inner
layers of cells thinner and these cells often orientated perpendicularly to the outermost
layer(s) which are often interspersed with minute orange granules also present on the wall
surface. Conidiophores arising from the inner wall of the pycnidial cavity, cylindrical,
irregularly sympodially branched, hyaline, smooth-walled, 2-8 jum tall and 3-3*5 //m

LICHENICOLOUS COELOMYCETES

wm

31

\\

l$ife^ ; : '

., . , -j ,/ *

^f&J l rfK?Jfsj>Ct~ yr ' * , *

sS^SS^? /;s *^ J>

Fig. 16 Libertiella malmeydensis (K isotype). A, Vertical section of pycnidium (x 500).
B, Vertical section of pycnidial wall with conidiogenous cells and conidia (x 1020).

wide. Conidiogenous cells enteroblastic, arising terminally and laterally from short conidio-
phores or directly from the inner wall of the pycnidial cavity, cylindrical, phialidic with a
short collarette, rarely proliferating, hyaline, 10-12 x2-3'5 //m. Conidia abundant,
adhering together in gelatinous masses, ellipsoid to obovoid with a slightly truncated base,
rather irregular in outline, hyaline, simple, rarely guttulate, smooth-walled, (5-)6-8 x (2-)3-

Host: Peltigera spuria (Ach.) DC., thallus. The pycnidia arise most abundantly on the
underside of the thallus but are also present in groups on the upper cortex, especially where
soredia were formerly produced in the P. erumpens (Taylor) Vainio morph of this species.
Libertiella malmedyensis causes relatively minor damage to infected thalli and does not
lead to the formation of necrotic patches. The host of this fungus has consistently been cited
as P. polydactyla (Necker) Hoffm., but Libert's material is P. spuria and not P. polydactyla.

Distribution: Belgium and Poland. It was evidently very abundant in the type locality as in
addition to sending material to several herbaria, Libert left enough in Brussels for the
exsiccatae of both Roumeguere and de Thiimen.

Observations: Libertiella malmedyensis shows some similarity to Karsteniomyces
peltigerae (p. 22) but differs in both the method of conidiogenesis and the non-septate and
differently shaped ascospores; in addition the pycnidial wall in K. peltigerae has much more
thickened pseudosclerenchymatous cells than L. malmedyensis and the conidia are not
markedly gelatinous.

Keissler (1930 : 582) took up the epithet 'peltigerae' for this species as he believed that was
published in March 1880 while 'malmedyensis' came out in April 1880. The issue of the
Revue mycologique including Spegazzini's paper was volume 2 part 1 as this has the title of
his paper printed on the front wrapper together with 'Janvier 1880'. In Grevillea volume 8
number 47, dated March 1 880 on the first page, and which included Cooke's article 'Relique
Libertianae' on pp. 81-87, on p. 120 of this same number appears under 'Cryptogamic
Literature' the entry 'ROUMEGUERE, C. Reliquae Libertianae, in "Revue Mycologique". Jan.,

32

D. L. HAWKSWORTH

Fig. 17 Libertiella malmeydensis (K isotype). A, Pycnidia on the host thallus showing
the deeply pigmented ring around the ostiole. B, Vertical section of pycnidium. C,
Vertical section of pycnidial wall with granular encrustations on the outer cells. D,
Conidiophores and conidiogenous cells. E, Conidia.

1880.' There can consequently be no doubt that Spegazzini's article, which was an appendix
to that of Roumeguere, appeared before the printing and distribution of the part ofGrevillea
including Cooke's paper. The epithet 'malmedyensis* must consequently be reinstated for
this species because it predates "peltigerae\ and in the absence of evidence to the contrary
'malmedyensis' is taken to have been validly published in January 1880. Keissler's date for
'malmedyensis' might have arisen from a hasty confusion of the wrappers of the Revue;
volume 2 part 2 has 'Avril 1 880' printed on it.

Roumeguere's exsiccatum included a description of Libertiella malmedyensis on the
printed label. The fascicle including no. 671, fasc. VII, was also issued in January 1880
according to a mention in Revue mycologique 2(1): 27. I have not been able to sequence the

LICHENICOLOUS COELOMYCETES 33

exsiccata and journal article within January 1880; it is even possible that they were
distributed simultaneously to save postage as Roumeguere then edited the journal!

Additional specimen: Poland: Puszcza Piska, on Peltigera spuria, June 1958, J. Zielinska (K!).

X. LICHENOCONIUM Petrak & H. Sydow
Beih. Repert. Spec. nov. Regni veg. 42 : 432 (1927).

See Hawksworth (1977: 170) and Sutton (1980: 118) for modern descriptions of the
genus.

Type species: Lichenoconium lichenicola (P. Karsten) Petrak & H. Sydow.
Number of species: Eight, all obligately lichenicolous.

Observations: Two additional species, Lichenoconium boreale (P. Karsten) Petrak & H.
Sydow and L. pertusariicola (Nyl.) D. Hawksw., were only retained in Lichenoconium by
Hawksworth (loc. cit.) pending further research on the generic names proposed for
Coelomycetes. From the major studies of Sutton (1977, 1980) it is evident that there is no
currently acceptable genus available for either of these two fungi. L. pertusariicola is
therefore referred to the new genus Laeviomyces above, and its separation from
Lichenoconium is discussed under that name (p. 27). L. boreale is not a lichenicolous
fungus but occurs on conifer lignum; that fungus differs from Lichenoconium in having a
pycnidial wall which consists of interwoven hyphae (textura intricata), very slender
conidiogenous cells, and almost colourless to pale olivaceous conidia which are ellipsoid to
almost allantoid, clearly guttulate, with smooth walls when studied by light microscopy
(although with a delicate close-packed verruculose ornamentation in SEM). The new generic
name Xeroconium D. Hawksw.* is therefore introduced for this species here; a detailed
description and illustrations of Xeroconium boreale (under the name Lichenoconium
boreale} were presented in Hawksworth (1977 : 171-172, fig. 1A, pi. 22 A-E) and further
drawings are given by Sutton (1980 : 1 19, fig. 56 A-C).

Key to the species

1 Pycnidia mainly exceeding 100 //mdiam

- Pycnidia mainly less than 80 //m diam

2 Conidia mainly exceeding 4 /um in length, distinctly echinulate or verruculose

- Conidia (2-)2-5-3'5(-4)x2-3/mi, rather smooth-walled; pycnidia (60-)80-120(-150)x

50-80(-100)yum Lichenoconium pyxidatae (p. 36)

3 Conidia globose to subglobose, sometimes rather angular

Conidia ellipsoid with a tapering and abruptly truncated base, (4-)6-8(-9) x l-4(-6) //m;

pycnidia 100-200 /*m diam; conidiogenous cells (6-)8-l 3(-l 5) x 2-3'5M'5) //m

Lichenoconium lichenicola (p. 36)

4 Conidia (2'5-)3^'5(-6)//m diam; pycnidia (80-)100-175(-200)//m diam; conidiogenous

cells (5-)6-8(-ll)x(2-)2-5^/im Lichenoconium xanthoriae (p. 37)

- Conidia 5-7(-7'5)x3-5-5(-6)//m; pycnidia 100-175(-200)//m diam; conidiogenous cells

(7-)8-10(-12)x(2-5-)3-3-5//m Lichenoconium cargillianum (p. 35)

5 Pycnidia mainly exceeding 50 //m diam; conidia mainly exceeding 3'5 //m diam .

- Pycnidia (20-)30-50(-60) //m diam; conidiogenous cells (3'5-)4-5(-6)x(2-)3-3'5(-4)//m;

conidia 2-3 -5Hl)//m Lichenoconium erodens (p. 35)

*Xeroconium D. Hawksw. gen. nov.

Genus ad Coelomycetes Sphaeropsidales pertinens. Similis generi Lichenoconio Petrak & H. Sydow sed ditlert in
muris pycnidiis textura intricata, cellulis conidiogenis gracilioribus, et conidiis ellipsoideis ad plusminusve
allantoideis, subhyalinis ad pallide olivaceis, gutullatis, laevibus (sed per microscopium electromcum mmuto-
verruculosis). Species holotypica, adhuc unica, est Xeroconium boreale (P. 'Karsten) D. Hawksw. comb. nov.
(basionym: Levieuxia borealis P. Karsten, Hedwigia 26 : 126, 1887).

34

D. L. HAWKSWORTH

B

10/im

Fig. 18 Lichenoconium species, conidiogenous cells and conidia. A, L. echinosporum
(UPS holotype). B, L. erodens (herb. Christiansen holotype). C, L. cargillianum
(E holotype). D, L. lichenicola (H-KARST 1246 holotype). E, L. lecanorae (IMI 192264). F,
L. pyxidatae (L holotype). G, L. usneae (K. isotype). H, L. xanthoriae (C holotype).
Reproduced from Hawksworth (1977 : 162, 164).

LICHENICOLOUS COELOMYCETES 35

6 Conidiogenous cells mainly exceeding 7 jum tall 7

- Conidiogenous cells (4-)5-7(-8)x(2-)3-3'5(-4)//m; conidia '(2-5-)3-4-5(-5-5)/n;' pyc-

nidia (30-)40-80(- 100) //m diam Lichenoconium lecanorae (p. 36)

7 Conidia verruculose, (2'5-)3-4(-5)//m diam; pycnidia (40-)50-80(-100)//m diam;

Conidiogenous cells (5-)7-9(-l 1 ) x (2-)2-5-3'5(-4) fim . . Lichenoconium usneae (p. 37)

- Conidia sparsely echinulate, 4- 5-5 x 3^4 //m; pycnidia 65-125 /zm diam; Conidiogenous

cells (7-)8-10(-ll)x 2-3 jam Lichenoconium echinosporum (p. 35)

1. Lichenoconium cargillianum (Lindsay) D. Hawksw., Persoonia 9 : 172 (1977)
(Fig. 18C)

See Hawksworth (1977 : 1 72-3) for description, illustrations and synonymy.

Hosts: Parmelia perforata Ach., Ramalina yemensis (Ach.) Nyl. and Usnea jlorida (L.)
Wigg., predominantly in apothecia.

Distribution: British Isles, Mexico and New Zealand.

2. Lichenoconium echinosporum D. Hawksw., Persoonia 9 : 173 (1977)
(Fig. 18 A)

See Hawksworth (1977: 1 73-1 74) for description and illustrations.
Host: Heterodea muelleri (Hampe) Nyl., a pathogen on the thallus.

Distribution: Australia. I have only seen the holotype but Filson (1978 : 17) reports seeing
pale brown necrotic patches like those due to this fungus from a wide range of Australian
localities.

3. Lichenoconium erodens M. S. Christ. & D. Hawksw., in Hawksworth, Persoonia 9 : 174

(1977).
(Fig. 18B)

See Hawksworth (1977: 1 74-1 77) for description and illustrations.

Hosts: Cladonia coniocraea (Florke) Sprengel, C. digitata (L.) Hoffim., Evernia prunastri
(L.) Ach., Hypogymnia bitteriana (Zahlbr.) Krog, H. physodes (L.) Nyl., Lecanora chlarona
(Ach.) Nyl., L. conizaeoides Nyl. ex Crombie, Parmelia afrorevoluta Krog & Swinscow (syn.
P. revoluta auct. p.p.), P. caperata (L.) Ach., P. crinita Ach., P. galbina Ach., P. laevigata
(Sm.) Ach., P. perlata (Huds.) Ach., P. saxatilis (L.) Ach., Parmeliopsis ambigua (Wulfen)
Nyl., Pertusaria hymenea (Ach.) Schaerer and P. pertusa (Weigel) Tuck.

Distribution: Austria, British Isles, Denmark, France, Germany, Sweden and the U.S.A.

Observations: A wide-ranging species which has been found on seven additional hosts
since its original description (Hawksworth, 1977). By far the most pathogenic species of the
genus, typically producing bleached areas on thalli which subsequently die. In cases where
this and other species of the genus are able to occur on the same host species, the symptoms
generally differ, Lichenoconium erodens always causing the most damage. This is also true
for the additional hosts included in the list above, including the Cladonia species and
Lecanora conizaeoides; its effects on the latter are illustrated and compared with those due
to L. lecanorae by Christiansen (1980).

Specimens (additional to those listed in Christiansen, 1980; Hawksworth, 1977, 1980c; and
Hawksworth & Minter, 1980): Austria: Steiermark, Grazer Bergland, Graz, Maria Trost, on
Hypogymnia physodes, 24 March 1974, J. Hafellner 568 (GZU!); Steiermark, Frauenalm S. of Musau,
on Cladonia digitata, 18 June 1978, /. Hafellner 3897 (GZU!). British Isles: England, Middlesex,
Ruislip, Park Wood, on Cladonia coniocraea on Quercus, 9 December 1979, D. L. Hawksworth 5009
(IMI 2445440!); S. Devon, Cornwood, Dendles Wood, on Parmelia laevigata, 29 February 1976, D. L.
Hawksworth 4293b (IMI 20 166 1/?!); S. Devon, Okehampton, nr Fatherford Viaduct, on Hypogymnia

36 D. L. HAWKSWORTH

physodes on Quercus, 30 March 1979, D. L. Hawksworth 4903 (IMI 236902!); Dorset, Studland Heath,
on H. physodes on Betula, 25 June 1979, D. L. Hawksworth 4952 (IMI 240076!); Hampshire, New
Forest, Stubbs Wood, on H. physodes, 13 May 1979, D. L. Hawksworth 4936 (IMI 238699!); S.
Somerset, nr Stoke Perol, on H. physodes on Quercus, 6 October 1979, D. L. Hawksworth 5002 (IMI
243338!); S. Somerset, Porlock, Horner Combe, on P. laevigata on Betula, 21 September 1980, D. L.
Hawksworth 5073 (IMI 251491!); Scotland, Dunbartonshire, Loch Lomond, Ross Park, on P. caperata
on Quercus, 9 October 1978, B. J. Coppins 3688 (E, IMI 232973!); Wales, Anglesey, Abberfraw sand
dunes, on Evernia prunastri, 1 7 October 1 977, D. L. Hawksworth 465 1 (IMI 2 1 7423!).

4. Lichenoconium lecanorae (Jaap) D. Hawksw., Bull. Br. Mus. nat. Hist. (Bot.) 6: 183

(1979).
(Fig. 18E)

See Hawksworth (1977 : 178-182, sub L. parasiticum D. Hawksw.) for description and
illustrations.

Hosts: Evernia prunastri, Lecanora admontensis Zahlbr., L. carpinea (L.) Vainio, L.
chlarotera Nyl., L. conizaeoides Nyl. ex Crombie, L. pallida (Schreber) Rabenh., L.
polytropa (Hoffm.) Rabenh., L. subfuscata Magnusson, L. superfluens Magnusson, Parmelia
borreri (Sm.) Turner, P. galbina Ach., P. pastillifera (Harm.) R. Schub. & Klem., P. saxatilis
(L.) Ach., P. sulcata Taylor, Rhizoplaca chrysoleuca (Sm.) Zopf and Squamarina lentigera
(Weber) Poelt. There are also some unconfirmed literature reports (see Hawksworth
1977: 179).

Distribution: Austria, ?Belgium, British Isles, Canada, Czechoslovakia, Denmark, France,
Germany, Hungary, Italy, Netherlands, Spain, Sweden, Switzerland and the U.S.A.

Observations: The symptoms of this species on Lecanora conizaeoides are illustrated and
compared with those due to Lichenoconium erodens by Christiansen (1980).

Specimens (additional to those listed by Hawksworth, 1977): Austria: Steiermark, Schladminger
Tauren, Kleinsolk-Obertal, on Lecanora polytropa, 12 August 1974, J. Hafellner 3932 (GZU!).
British Isles: England, Hampshire, New Forest, Stubbs Wood, on Parmelia sulcata, 13 May 1979, D.
L. Hawksworth 4937 (IMI 238700!); S. Devon, Slapton Ley Nature Reserve, Peasdish, on P. sulcata, 23
August 1980, D. L. Hawksworth 5044b (IMI 251 1262!); loc. cit., on Lecanora sp., 23 August 1980, D. L.
Hawksworth 5044a (IMI 251261!). France: Dept. Isere, Rhone-Tal E. Lyon, on Squamarina
lentigera, 13 July 1975, J. Hafellner 1575 (GZU!). Italy: Siidtirol, Mendelgebirge, Penegal SW. von
Bozen, on Lecanora carpinea, 20 October 1975, J. Hafellner 1092 (GZU!); Otztaler Alpen, Vinschgau,
on Rhizoplaca chrysoleuca, 1 8 October 1975,7. Hafellner 997 p.p. (GZU!).

5. Lichenoconium lichenicola (P. Karsten) Petrak & H. Sydow, Beih. Repert. nov. Spec. Regni

veg. 42: 432(1927).
(Fig. 18D)

See Hawksworth (1977 : 177-178) for description and illustrations. Further drawings are
provided by Sutton (1980 : 1 19 fig. 55a-c).

Host: Physcia aipolia (Ehrh. ex Humb.) Fiirnrohr, apothecia.
Distribution: Finland. Known only from the type collection.

6. Lichenoconium pyxidatae (Oudem.) Petrak & H. Sydow, Beih. Repert. nov. Spec. Regni

veg. 42: 435 (1927).
(Fig. 18F)

See Hawksworth (1977 : 184-185) for description and illustrations.

Hosts: Cladonia arbuscula (Wallr.) Rabenh., C. cenotea (Ach.) Schaerer, C.fimbriata (L.)
Fr., C. incrassata Florke, C. pocillum (Ach.) O.-J. Rich., and C. pyxidata (L.) Hoffm.;
usually on the podetia but occasionally on squamules. There are also unconfirmed reports
from additional hosts in Cladonia and other genera, the latter certainly erroneous (see
Hawksworth, loc. cit.).

LICHENICOLOUS COELOMYCETES 37

Distribution: Austria, British Isles, Denmark, France, Germany, the Netherlands and
Sweden.

Observations: Harmand (1907 : 226) stated that Cladonia rangiferina f. moribunda Harm,
was based on deformed podetia with verrucae caused by a fungus close to Lichenoconium
pyxidatae but 'il se distingue par des spores franchement brunes et ovoi'des' according to
Vouaux who Harmand indicates studied the material. I have not seen type or authentic
material ofthistaxon.

7. Lichenoconium usneae (Anzi) D. Hawksw., Persoonia 9 : 185 (1977).
(Fig. 18G)

See Hawksworth (1977 : 1 85-1 90) for description, illustrations and synonymy.

Hosts: Anaptychia ciliaris (L.) Korber, Bryoriafuscescens (Gyelnik) Brodo & D. Hawksw.,
Cladonia arbuscula (Wallr.) Rabenh., C. cariosa (Ach.) Sprengel, Hypogymnia physodes (L.)
Nyl., Lecanora pacifica Tuck., Letharia vulpina (L.) Hue, Parmelia conspersa (Ehrh. ex
Ach.) Sprengel, P. exasperata (Ach.) de Not., P. glabratula (Lamy) Nyl., P. olivacea (L.)
Ach., P. pulla Ach., P. rudecta Ach., P. saxatilis (L.) Ach., P. verruculifera Nyl., Physcia
aipolia (Ehrh. ex Humb.) Fiirnrohr, P. stellaris (L.) Nyl., Physconia pulverulacea Moberg,
Ramalina cf. baltica Lettau, R. calicaris (L.) Fr., R.fraxinea (L.) Ach., R. siliquosa (Huds.)
A. L. Sm., R. subgeniculata Nyl., Usneafilipendula Stirton aggr., and U. tf.jlorida (L.) Wigg.
For unconfirmed literature reports see Hawksworth (loc. cit.).

Distribution: Austria, British Isles, Canada, Canary Islands, Czechoslovakia, Denmark,
France, Germany, Italy, Norway, Spain, Sweden, Switzerland, Yugoslavia (Keissler,
1933: 388) and the U.S.A.

Specimens (additional to those listed in Hawksworth, 1977): Austria: Steiermark, Stubalpe, Gaberl,
on Usnea cf. florida, 19 December 1976, J. Poelt & J. Hafellner 1825 (GZU!); Steiermark, Kreuzberg,
on Cladoniasp., 16 March 1975, W. Moschl&H. Pittoni [herb. Hafellner 437] (GZU!); Nockgruppe,
Afritzer Berge, Karten, on Usnea sp., 27 August 1974, J. Poelt (GZU!). British Isles: England, S.
Devon, Slapton, Duck Marsh, on Ramalina fraxinea, 5 September 1978, D. L. Hawksworth 4869 (IMI
231752!); Scotland, Midlothian, Carlops, Hobbies House, on Bryoriafuscescens, 29 March 1977, B. J.
Coppins 2727 (E!). Canada: Alberta, Kananaskis Range, between Wind Ridge and Mount Longhead,
on Letharia vulpina, 22 June 1978, D. C. Lindsay (IMI 2298696!). France: Dept. Vas, He de Port
Cros, on Ramalina calicaris, 17 July 1973, Y. Rondon [herb. Hafellner 1826] (GZU!).

8. Lichenoconium xanthoriae M. S. Christ., Friesia 5:212 (1956).
(Fig. 18H)

See Hawksworth (1977 : 190-192) for description and illustrations.

Hosts: Cetraria sepincola (Ehrh.) Ach., Cetrelia olivetorum (Nyl.) Culb. & C. Culb. s. lat.,
Xanthoria parietina (L.) Th. Fr., X. polycarpa (Hoffm.) Rieber. The collections on Cetrelia
have been reported on by Hawksworth & Minter ( 1 980).

Distribution: British Isles, Czechoslovakia, Denmark and Sweden.
XI. LICHENODIPLIS Dyko & D. Hawksw.

in Hawksworth & Dyko, Lichenologistll : 5 1 (1979).
See Hawksworth & Dyko (1979) for further information on this genus.
Type species: Lichenodiplis lecanorae (Vouaux) Dyko & D. Hawksw.
Number of species: Two, both obligately lichenicolous.

Key to the species

1 Conidiomata 50-1 20 //m diam; conidia 4-7-5 x 2-3 ^m . - Lichenodiplis lecanorae (p. 38)
- Conidiomata 50-70 //mdiam;conidia 9-5-1 3 x 4-4-5 //m . . Lichenodiplis hchemcola (p. 38)

38 D. L. HAWKSWORTH

1. Lichenodiplis lecanorae (Vouaux) Dyko & D. Hawksw., in Hawksworth & Dyko,

Lichenologist II : 52(1979).

See Hawksworth & Dyko (1979) for description, illustrations and synonymy.

Hosts: Caloplaca caesiorufa (Wibel) Flagey, C. cerina (Ehrh. ex Hedw.) Th. Fr., C.
ferruginea (Huds.) Th. Fr., C. flavovirescens (Wulfen) Dalla* Torre & Sarnth., Evernia
prunastri (L.) Ach., Lecanora confusa Almb., L. dispersa (Pers.) Sommerf., L. pallida
(Schreber) Rabenh., L. polytropa (Hoffm.) Rabenh., L. saligna (Schrader) Zahlbr., Pertusaria
albescens (Huds.) M. Choisy & Werner and P. cf. leioplaca DC. Hawksworth & Dyko (1979)
also note that there are literature records from Caloplaca lactea (Massal.) Zahlbr., Lecanora
expallens Ach., L. varia (Hoffm.) Ach., Diploschistes scruposus (Schreber) Norman,
Lecidella elaeochroma (Ach.) M. Choisy, Lecidea enteroleuca Ach., Micarea nitschkeana
(Lahm ex Rabenh.) Harm, and unidentified Lecidea and Pertusaria species.

Distribution: Hawksworth & Dyko (1979) saw material from the British Isles, Canary
Islands, Denmark, France, Germany, Spain and Sweden; they also noted a literature report
from the U.S.S.R. (Novaya Zemyla). Also mentioned from Morocco by Werner (1972 : 95)
on Pertusaria pustulata f. glabrata (Anzi) Hue.

Observations: While examining material in H-NYL in August 1979, I discovered that
Nylander had given this species the herbarium name 'Dichaena pertusariae" 1 ; this name does
not, however, ever appear to have been used in print.

Specimens (additional to those listed by Hawksworth & Dyko, 1979): British Isles: England, S.
Devon, Torquay, on Pertusaria cf. leioplaca, [?R. Deakin] [Leighton, Lich. exs. no. 245; sub
Opegrapha atra var. parella.] (E!); Westmorland, Brathay church, on Lecanora cf. saligna, 22
September 1979, A. Henderson (herb. Henderson!). Scotland, W. Inverness, Strontian, Horsley Hall, on
Pertusaria albescens on Ulmus, 14 July 1966, P. W. James (IMI 237277!). France: Brittany, Brest, on
Pertusaria cf. leioplaca, Crouan 135 (H-NYL p.m. 7683!).

2. Lichenodiplis lichenicola Dyko & D. Hawksw., in Hawksworth & Dyko, Lichenologist

11:56(1979).

See Hawksworth & Dyko (1979) for description, illustrations and a possible synonym.
Host: Rinodina septentrionalis Malme, apothecia.

Distribution: Hawksworth & Dyko (1979) indicated that the type and only known
collection of this species came from Finnmark. However, Dr P. M. Jorgensen (in litt.)
has kindly pointed out that the type almost certainly came from the Yenisey region of the
U.S.S.R.

Observations: I have also seen a further collection that may belong to this species (France:
Vosges, Docelles, on Pertusaria cf. leioplaca, J. Harmand, herb. Vouaux!), with conidia
9-12 x4-5 jam, but hesitate to refer it definitely to Lichenodiplis lichenicola as the conidia
are rather thick-walled, much darker brown, and tend to be somewhat broader in relation to
their length. Until more collections are available which enable the variability of L.
lichenicola to be determined, it would be premature to either describe this material as new or
state that it definitely belonged here.

XII. LICHENOSTICTA Zopf

Nova Acta Acad. Caesar. Leap. Carol. 70(4) : 263 (1898).

Conidiomata pycnidial, arising singly, uniloculate, subglobose to broadly pyriform,
erumpent to largely superficial, translucent brown to dark brown or black; walls composed
of a few layers of intertwined brownish irregular hyphae forming a textura intricata, lined by
a hyaline almost pseudoparenchymatous layer of cells, thickened near the ostiole.
Conidiophores hyaline, irregularly branched, flexuous, multi-septate, arising from the inner

LICHENICOLOUS COELOMYCETES 39

cells of the pycnidial wall. Conidiogenous cells enteroblastic, phialidic, sometimes
proliferating, integrated into chains, acro-pleurogenous, conidia arising from the apex of a
chain and (for other conidiogenous cells) laterally, sometimes polyphialidic, hyaline,
cylindrical to doliiform, channel and collarette distinct, narrow proliferating tissue often
evident. Conidia adhering in slime and extruded as a gelatinous drop, lacriform, distinctly
attenuated at the base, hyaline, simple, smooth-walled.

Type species: Lichenosticta alcicornaria (Lindsay) D. Hawksw. (syn. L. podetiicola Zopf).
Number of species: Monotypic.

Observations: The generic name Dendrophoma Sacc. cannot be used for this species as it is
a synonym of Dine masporium Lev., a very different fungus (Sutton, 1968). The catenate
arrangement of the conidiogenous cells and details of conidiogenesis show a strong similarity
to those of the pycnidia of some lichenized members of the Lecanorales (see Vobis &
Hawksworth, 1981). The normal pycnidia of Cladonia species have much more regularly
branched conidiophores and mainly ellipsoid to bacillariform conidia.

1. Lichenosticta alcicornaria (Lindsay) D. Hawksw., in Hawksworth et al, Lichenologist

12: 107(1980).
(Fig. 19A-G)

Microthelia alcicornaria Lindsay, Q. Jl microsc. Sci. II, 9 : 349 ( 1 869).

Type: British Isles, England, Shropshire, Haughmond Hill, on Cladonia foliacea (Huds.) Willd.,

W. A. Leighton [Lich. exs. no. 1 5] (E lectotype!; BM, E isolectotypes!).
Dendrophoma alcicornaria (Lindsay) Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 283 (1914).
Lichenosticta podetiicola Zopf, Nova Acta Acad. Caesar. Leop. Carol. 70(4) : 263 ( 1 898).
Dendrophoma podetiicola (Zopf) Keissler, Ost. hot. Z. 60 : 62 ( 1 9 1 0).
Aposphaeria cladoniae Allescher & Schnabl, in Allescher, Ber. Bayer, hot. Ges. 4 : 32 (1 896).

Type: Germany, Munich, Pullach, on Cladonia fimbriata (L.) Fr., October 1894, J. N. Schnabl

(M holotype!).

Phoma cladoniae (Allescher & Schnabl) Keissler, Annln naturh. Mus. Wien 34 : 77 (1 92 1 ).
1 "Aposphaeria cladoniae var.floerkeanae Vouaux, in Harmand, Lich. Fr. 3 : 335 (1907).

Exsiccatae: Leighton, Lich. exs. no. 1 5 (sub Cladonia alcicornis; BM!, E!). Rasanen, Lich.fenn. exs.
no. 600 (sub Dendrophoma alcicornaria*; BM!).

Icones: Lindsay, Trans. R. Soc. Edinb. 22: pi. 8 fig. 3 ( 1859). Zopf, Nova Acta Acad. Caesar. Leop.
Carol. 70(4) : 264 figs 22-23, 265 figs 24-25 (1898).

Conidiomata pycnidial, immersed or sometimes becoming to } erumpent, arising singly,
often numerous, scattered, translucent brown to dark brown or black, sometimes shiny,
(50-)80-120(-150)//m, subglobose to broadly pyriform, ostiolate; pycnidial wall irregular
in outline, mainly 5-10 //m thick but to 20-25 /urn thick around the ostiole, composed of
1-3 (or more near the ostiole) irregular layers of intertwined hyphae, hyphae olivaceous
brown to dark brown, uneven in diameter, very variable in shape, sometimes strongly in-
flated between the septa, 3-7 um wide, thick-walled near the ostiole, inner cells subhyaline
and almost pseudoparenchymatous in parts. Conidiophores hyaline, irregularly branched,
occupying the whole of the pycnidial cavity in young pycnidia, flexuous, multi-septate,
arising from the inner cells of the pycnidial wall. Conidiogenous cells enteroblastic,
phialidic, sometimes percurrently proliferating, integrated into chains, acro-pleurogenous
with conidia arising from the apex of a chain and (for other conidiogenous cells) laterally or
near a transverse septum, sometimes polyphialidic, hyaline, cylindrical to doliiform, mainly
2-5-4 um wide and 4-6 um long, channel and collarette distinct, this (and often

*No. 350, distributed by Rasanen under this name, is not this species but probably an unknown pycnothyriaceous
fungus. Mentioned by Sandstede (1931) as present in his Clad. Exs. nos. 601, 870-2, and 1574 but these numbers
have not been studied by me.

40

D. L. HAWKSWORTH

Fig. 19 Lichenoslicta alcicornaria. A-E (Rasanen, Lich. Exs. no. 600, K); A, pycnidia on
podetia of the host; B, vertical section of pycnidium; C, cells from the pycnidial wall; D,
chains of conidiogenous cells; E, conidia. F (Hafellner 1834), Conidiogenous cells and
conidia. G (BM isotype), Conidiogenous cells and conidia.

conspicuous narrow proliferations) 1-1*5 //m wide. Conidia adhering in slime and extruded
as a gelatinous drop, lacriform, plano-convex to concavo-convex, rounded at the apex and
distinctly attenuated at the base, hyaline, simple, often 1 or more guttulate, smooth-walled,
(6-)6-5-10(-l 1 ) x (2-)3^-5(-6) //m.

LICHENICOLOUS COELOMYCETES 41

Hosts: I have examined material on Cladonia arbuscula (Wallr.) Rabenh. (C. sylvatica
auct.), C. fimbriata (L.) Fr., C. foliacea (Huds.) Willd. and C. gracilis (L.) Willd. Also
mentioned from C. cornuta (L.) Hoffm. by Keissler (1910:2), C. pyxidata (L.) HofTm. by
Keissler (1930 : 551); from C. acuminata (Ach.) Norrlin (C. norrlinii Vainio), C. portentosa
(Dufour) Coem. (C. impexa Harm.), and C. ciliata var. tennis (Florke) Ahti by Sandstede
(1931 : 52, 76, 340); and from C. coniocraea auct. by Folan & Mitchell (1970 : 169). The
fungus occurs on both podetia and squamules (especially the lower surfaces) but causes little
damage so is easily overlooked. Probably parasymbiotic.

Distribution: Austria (Keissler, 1930 : 551), the British Isles (England; Ireland, Folan &
Mitchell, 1970 : 169), Canada (Newfoundland; Keissler, 1910:2), Finland, Germany, Italy
and Sweden (Keissler, 1910 : 2;Santesson, 1949 : 142).

Observations: The fungus is easily distinguished from other Coelomycetes on Cladonia by
the characteristic lacriform conidia and catenately arranged acro-pleurogenous conidio-
genous cells. Further, it never appears to induce the formation of gall-like growths.

The conidial shape is rather constant in this species but there is a considerable degree of
variation in the sizes of conidia between different collections. For example, on a specimen of
C. arbuscula (IMI 232629!) they measured (8-)9-10(-l l)x 4-5-6 //m while on one of C.
gracilis (Lich. Fenn. Exs. no. 600, BM!) they were (6-)6'5-8(-8'5) x 3^4(-5)//m (see Fig.
19E-G). It is thus conceivable that more than a single species might be involved, but in the
current state of our knowledge of the variability of Lichenosticta alcicornaria it would be
imprudent to attempt to recognise two taxa at the present time. The types of both
Aposphaeria cladoniae and Microthelia alcicornaria have conidia in the lower size range, as
did that of Lichenosticta podetiicola according to the measurements given by Zopf
(1898:264).

Lindsay (1869& : 349) did not provide a description of this species but the name is validly
published because he refers to his earlier work (Lindsay 1 859 : 161), where a description and
illustrations were provided; his epithet must therefore be adopted as it pre-dates that of Zopf.
No authentic material of Lichenosticta podetiicola could be traced in B (B. Hein, in litt.) or
M (H. Hertel, in litt.}, but the figures published by Zopf (1898 : 264-5) leave little doubt the
taxa are conspecific, even though Zopf did not show any conidiogenous cells, as the conidia
are also of the correct shape and size.

Aposphaeria cladoniae \ar.floerkeanae could not be located amongst Harmand's material
in Angers (M. Guerlesquin, in litt.) and is not represented in the remnants of Vouaux's
herbarium (Rondon, 1970). It was distinguished by conidia 8 x 3 //m so is tentatively
retained here as those dimensions are within the range of Lichenosticta alcicornaria as
interpreted here.

Additional specimen: Italy: Siidtirol, Siidtiroler Dolomiten, Passo di Rolle, S Abhange des Mt
Castellazzo, on Cladonia arbuscula, 23 October 1976, J. Hafellner 1834 (herb. Hafellner!, IMI
232629!).

XIII. MICROCALICIUM Vainio [ANAMORPHS]

Ada Soc. Fauna Flora fenn. 57(1) : 77 (1927).

Conidiomata pycnidial, arising singly or in small groups on the thallus of the host,
subglobose, erumpent to almost sessile, with the base to half immersed, dark brown to almost
black, ostiolate; walls composed of several layers of pseudoparenchymatous cells, cells in the
upper parts of the pycnidium very thick-walled, olivaceous, aeruginose or brown, almost
sclerenchymatous, cells in the lower parts of the pycnidium similar or sometimes subhyaline
to pale brown, thin-walled, irregular in shape and often almost hyphal in parts.
Conidiophores absent. Conidiogenous cells enteroblastic, lining the pycnidial cavity,
acrogenous, subcylindrical to cylindrical, phialidic, with a particularly distinctive often
elongate canal, not proliferating, hyaline, smooth-walled. Conidia arising singly, not

42 D. L. HAWKSWORTH

catenate, subglobose to narrowly ellipsoid, sometimes slightly apiculate at the base, hyaline,
simple, thin-walled, smooth-walled, extruded through the ostiole in a mucilaginous drop.

Type species: Microcalicium subpedicellatum (Schaerer) Tibell (syn. M. disseminatum
(Ach. ex Fr.) Vainio).

Number of species: Four species of Microcalicium (Caliciales) were accepted by Tibell
(1978) in a critical revision of the genus. Anamorphs are known for two of the species and
only these are treated here. Tibell's work should be consulted for information on the
teleomorphs.

Observations: No suitable coelomycete generic name could be found for the anamorphs of
Microcalicium species. I have not introduced a separate generic name for them here because
they often (though not exclusively) occur together with the teleomorph, and, when found
separately, 'anamorph' can easily be added after the name. Further, as the genus includes
lichenized and not only lichenicolous fungi, Art. 59.1 should perhaps be invoked to prevent
the introduction of an anamorphic name, as this rule, which permits a dual nomenclature in
fungi with pleomorphic life-cycles, specifically excludes (correctly in my view; cf.
Hawksworth, 1978&: 236-237) lichen-forming taxa.

Key to the species

1 Conidia subglobose, 2-3 x 1-5-2 //m; conidiogenous cells 6-9 x 2-3 um

Microcalicium subpedicellatum [anamorph] (p. 42)
- Conidia narrowly ellipsoid, 2-3 (-4) x 1 -5-2 um; conidiogenous cells 3-5(-6)x 1-5-2-5

(-3)/zm Microcalicium conversum [anamorph] (p. 42)

1. Microcalicium conversum Tibell, Bot. Notiser 131 : 237 (1978) [anamorph].
(Fig. 20G-H)

Icones: Tibell, Bot. Notiser 131 : 242 fig. 7B (1978) [anamorph].

Conidiomata pycnidial, arising singly, subglobose, erumpent to almost sessile or partly
immersed, dark brown to black, 40-60(-80) um wide and 50-80 um tall; walls 8-1 2 um
thick, composed of several layers of reddish brown very thick-walled almost scleren-
chymatous pseudoparenchymatous cells, becoming somewhat olivaceous in postassium
hydroxide, 3-5 um diam, basal wall apparently similar to the exposed parts at least in
almost superficial pycnidia. Conidiophores absent. Conidiogenous cells enteroblastic, lining
the pycnidial cavity, acrogenous, subcylindrical, phialidic, with a distinct and often elongate
canal, not proliferating, hyaline, smooth-walled, 3 -5 (-6) x l'5-2'5(-3)//m. Conidia arising
singly, not catenate, narrowly ellipsoid, rounded at the ends, hyaline, simple, thin-walled,
smooth-walled, sometimes guttulate, 2-3(-4) x (1-) 1*5-2 um, extruded through the ostiole
in a mucilaginous drop.

Hosts: Calicium viride Pers. and Chaenotheca subroscida (Eitner) Zahlbr., thalli.

Distribution: The species was recorded by Tibell (loc. cit.) only from Argentina (Tierra del
Fuego), Australia (Tasmania) and Chile (Magallanes).

Observations: The anamorph of Microcalicium conversum is clearly separated from that of
M. subpedicellatum on the basis of the more complete pycnidial wall, its reddish brown
colour, the length of the conidiogenous cells, and the quite different shape of the conidia.
The teleomorph also differs from that of M. subpedicellatum in having a reddish brown (not
greenish) excipulum, and 1 (not 3-7)-septate ascospores (fide Tibell, loc. cit.).

Specimen: Chile: Terr. Magallanes: Rio Rubens, near Hotel Rio Rubens (about 50 km SE of
Natales), on Calicium viride on Nothofagus pumila, 1 3 January 1 94 1 , R. Santesson 56 1 8 p.p. (S!).

2. Microcalicium subpedicellatum (Schaerer) Tibell, Bot. Notiser 131:240 (1978)

[anamorph].
(Fig. 20A-F)

LICHENICOLOUS COELOMYCETES

43

Fig. 20A-F, Microcalicium subpedicellatum [anamorph] (Hajellner 1095); A, pycnidia on
the host thallus; B, vertical section of pycnidium; C, cells from the upper part of the
pycnidial wall; D, cells from the lower part of the pycnidial wall; E, conidiogenous cells;
F, conidia. G-H, M. conversum [anamorph] (Santesson 5618 p.p); G, conidiogenous cells; H,
conidia.

44 D. L. HAWKSWORTH

Icones: Tibell, Bot. Notiser 131 : 245 fig. 10B, 246 fig. 1 1 (1978) [anamorph].

Conidiomata pycnidial, subglobose, arising singly or in small groups, erumpent with the
lower \-\ remaining immersed, black, 40-60 /zm diam; walls in the upper parts mainly
10-1 5 //m thick, composed of several layers of somewhat aeruginose very thick-walled
almost sclerenchymatous pseudoparenchymatous cells, becoming brownish in potassium
hydroxide, 3-5 //m diam, walls in the lower parts 5-8 //m thick, composed of 2-3 layers of
thinner-walled subhyaline cells forming an irregularly pseudoparenchymatous to in places
almost hyphal tissue. Conidiophores absent. Conidiogenous cells enteroblastic, lining the
pycnidial cavity, acrogenous, subcylindrical to cylindrical, phialidic, with a distinctly
thickened and elongate canal, not proliferating, hyaline, smooth-walled, 6-9 x 2-3 //m.
Conidia arising singly, not catenate, subglobose, sometimes slightly pointed at the base,
hyaline, simple, thin-walled, smooth-walled, 2-3 x l-5-2//m, extruded through the ostiole
in a milky-white mucilaginous drop.

Hosts: Calicium glaucellum Ach., C. viride Pers., Chaenotheca brunneola (Ach.) Mull.
Arg., C. chrysocephala (Turner ex Ach.) Th. Fr., C. subroscida (Eitner) Zahlbr., C. trichialis
(Ach.) Th. Fr., and undetermined Caliciales, thalli. A parasitic species in which the pycnidia
colonize at an early stage with to 15 per host areole. Infected thalli are eventually
decolourized and may turn brownish with the formation of extensive lesions in the colonies.
Sometimes apparently growing directly on lignum or with other algae, perhaps persisting in
situ after the loss of the host in such cases.

Distribution: Circumboreal. Recorded by Tibell (loc. cit.) from Austria, the British Isles
(Scotland), Canada, France, Italy, Norway, Sweden, Switzerland and the USSR.

Observations: The separation of this species from the anamorph of M. conversum is
discussed under that species above.

Specimens: Austria: Steiermark, Ostalpen, Gurktaler Alpen, Frauenalm S von Murau, on
Chaenotheca trichialis, 18 June 1978, J. Hafellner 3899 (GZU!). Italy: Sudtirol, Mendelgebirge,
Penegal SW von Bozen, on C. chrysocephala, 20 October 1975, J. Hafellner 1095 (GZU!); loc. cit., on
Chaenotheca sp., 20 October 1975, J. Poelt & J. Hafellner 1094 (GZU!); loc. cit., on Chaenotheca sp.,
20 October 1975,7. Hafellner 1084 (GZU!).

XIV. MINUTOPHOMA D. Hawksw. gen. nov.

Genus lichenicola ad Coelomycetes Sphaeropsidales pertinens. Conidiomata singularia, uniloculata,
dispersa, globosa vel subglobosa, semi-immersa, nigra, cum muris textura angulari sed inferiore solum
ex cellulis conidiogenis, ostiolata. Conidiophora desunt. Cellulae conidiogenae enteroblasticae,
acrogenae, ampulliformes vel breve ampulliformes, phialidicae, non prolifericae, hyalinae sed basi
plerumque pallide brunneae. Conidia subcylindrica, hyalina, simplicia, laevia, minutissima.

Conidiomata pycnidial, arising singly, scattered, globose to subglobose, half-immersed,
black; walls composed of thick-walled dark brown cells forming a textura angularis above,
these cells readily separating, the lower part consisting only of the conidiogenous cells, the
upper cells separating to form an irregular ostiole. Conidiophores absent. Conidiogenous
cells enteroblastic, forming the lower part of the pycnidial wall, acrogenous, ampulliform to
shortly ampulliform, phialidic, not proliferating, hyaline but with the base often pale brown.
Conidia not adhering, not catenate, subcylindrical, rounded at the apices, hyaline, simple,
smooth-walled, minute.

Type species: Minutophoma chrysophthalmae D. Hawksw. (holotypus).
Number of species: Monotypic.

Observations: Minutophoma is most closely allied to Phoma (see p. 49), but differs in the
extremely small size of the pycnidia and some other important respects: the thick-walled
angular cells tending to separate around an irregular ostiole, the absence of a differentiated
wall in the immersed part of the pycnidium, the conidiogenous cells tending to be more

LICHENICOLOUS COELOMYCETES

45

lOjim

Fig. 21 Minutophoma chrysophthalmae (IMI 237276 holotype). A, Vertical section of
pycnidium. B, Surface view of cells from the upper part of the pycnidium which easily
separate. C, Conidia. D, Conidiogenous cells.

elongated and also pigmented below, and the extremely small conidia. An additional feature
might be the absence of a very deeply pigmented area around the ostiole, but this is not
emphasized here because the whole of the exposed part of the pycnidium in M.
chrysophthalmae might be considered as equivalent to this zone.
The separation ofAsterophoma from Minutophoma is discussed above (p. 8).

1. Minutophoma chrysophthalmae D. Hawksw. sp. nov.
(Fig.21A-D)

Fungus lichenicola. Conidiomata singularia, uniloculata, dispersa, globosa vel subglobosa, semi-
immersa, nigra, 20-40 ^m diam, cum muris textura angulari 5-8 /urn latis e cellulis atrobrunneis
3-4 //m diam sed inferiore solum ex cellulis conidiogenis; ostiolata. Conidiophora desunt. Cellulae
conidiogenae enteroblasticae, acrogenae, ampulliformes vel breve ampulliformes, phialidicae, non
prolifericae, hyalinae sed basi plerumque pallide brunneae, 3 '5-6 x2-3 - 5yum. Conidia subcylindrica,
hyalina, simplicia, laevia, 2-3 '5 x 1 -5 jum.

Typus: Magna Britannica, Scotia, E. Inverness, Glen Affric, Coille Ruigh na Cuileige, in apotheciis
Chrysothrix chrvsophthalmae (P. James) P. James & Laundon e ligno Pint, 18. vi. 1976, B. J. Coppins
3196 p.p. (IMI 237276 holotypus!; E isotypus!).

Conidiomata pycnidial, semi-immersed, sometimes becoming erumpent and almost
superficial, scattered over the surface of the host apothecia, arising singly, black, 20-40 //m
diam; wall in the exposed part of the pycnidium composed of 2-3 layers of cells, 5-8 jam
thick, dark brown, cells thick-walled, pseudoparenchymatous, tending to be angular, not
radially compressed, readily separating in squash preparations, 3-4 jam diam, the upper
cells separating to form an irregular ostiole, the lower part of the pycnidium composed
almost exclusively of conidiogenous cells, the basal parts of which become slightly pig-
mented. Conidiogenous cells enteroblastic, forming the lower part of the pycnidial wall,
arranged in a single layer, acrogenous, ampulliform to shortly ampulliform, phialidic, not
proliferating, mainly hyaline to subhyaline but with the base often pale brown,
3'5-6 x 2-3'5 //m. Conidia abundantly produced, subcylindrical, rounded at the apices, not
adhering, not catenate, hyaline, simple, not distinctly guttulate, smooth-walled,
2-3-5 x 1-1-5 //m.

Host: Chrysothrix chrysophthalma (P. James) P. James & Laundon (syn. Micarea
chrysophthalma P. James), apothecia. Infected apothecia are often richly covered by

46 D. L. HAWKSWORTH

pycnidia but otherwise retain their characteristic bright yellow-green appearance and
continue to produce ascospores. The species consequently has to be interpreted as a
parasymbiont rather than a parasite.

Distribution: British Isles (Scotland).

Observations: A distinctive species which, despite the minute size of the pycnidia, is easily
detected by their black colour contrasting markedly with that of the host's apothecia. As a
result of its small size and the minute conidia, Minutophoma chrysophthalmae is unlikely to
be confused with the other taxa treated in the present revision.

Additional specimens: British Isles: E. Inverness, Guisachan Forest, south-west ofGarve Bridge, on
Chrysothrix chrysophthalma on Pinus lignum, 26 May 1975, B. J. Coppins et al. 1955 (E!); E. Ross,
Amat Forest, on Chrysothrix chrysophthalma on Pinus lignum, 28 May 1975, B. J. Coppins (& F.
Rose) 2225 (E\).

XV. NIGROPUNCTA D. Hawksw. gen. nov.

Genus lichenicola ad Coelomycetes Sphaeropsidales pertinens. Conidiomata singularia, uniloculata,
dispersa, subglobosa ad cupuliformia, immersa, plusminusve hyalina, cum muris textura intricata, late
ostiolata. Conidiophora non vel sparse ramosa, hyalina, septata, e cellulis plusminusve isodiametricis.
Cellulae conidiogenae holothallicae, acrogenae, integratae, terminales, determinatae, subglobosae ad
subcylindrices, hyalinae vel ad apicem olivaceae. Conidia catenata, in cirrhum accumulata,
irregulariter, e cellulis numerosis composita, cellulis subglobosis vel angularibus, atro-olivacea vel
nigra, plerumque simplicibus, rugulosa.

Conidiomata pycnidial, arising singly, subglobose at first but becoming cupuliform with
age, immersed, hyaline, largely obscured by the mass of conidia; walls composed of loosely
interwoven hyaline hyphae forming a textura intricata. Conidiophores forming a compact
layer lining the inner wall of the pycnidial cavity, not or sparsely branched, hyaline, septate,
composed of isodiametric cells. Conidiogenous cells holothallic, acrogenous, integrated,
terminal, determinate, subglobose to subcylindrical, hyaline but becoming olivaceous at the
apex. Conidia catenate at first, accumulating in a cirrhus, irregular, composed of numerous
subglobose to angular cells, dark olivaceous or black, individual cells mainly simple,
rough-walled.

Type species: Nigropuncta rugulosa D. Hawksw. (holotypus).
Number of species: Monotypic.

Observations: This genus appears to have no close allies amongst the non-lichenicolous
fungi. The method of conidiogenesis and the aggregation of cells to form multicellular
propagules recalls that seen in the lichenicolous hyphomycete genus Sclerococcum Fr. ex Fr.
(Hawksworth, 19750, 1979), but there the conidiogenous cells are arranged in superficial
sporodochia and the conidia are smooth-walled. Amongst the lichenicolous Coelomycetes,
there is some slight similarity to Vouauxiella uniseptata (see p. 66) but in that taxon the
conidiogenous cells are much longer and pigmented, the conidia are consistently 1 -septate,
not aggregating into multi-cellular clumps, have a different type of ornamentation, and are
produced in a pycnidium with a pigmented and cellular wall.

The generic name proposed recalls the black-spotted appearance imparted to the host
thallus.

1. Nigropuncta rugulosa D. Hawksw. sp. nov.
(Figs.22A-D,23A-C)

Fungus lichenicola. Conidiomata singularia, uniloculata, dispersa, subglobosa ad cupuliformia,
immersa, plusminusve hyalina, 1 00^150 //m lata, cum muris textura intricata 10-20/zm latis, e hyphis
hyalinis 2'5-3^m latis, late ostiolata. Conidiophora non vel sparse ramosa, hyalina, septata,
10-15 x 2-3-5 //m, e cellulis isodiametricis. Cellulae conidiogenae holothallicae, acrogenae, integratae,
terminales, determinatae, subglobosae vel subcylindrices, hyalinae vel ad apicem interdum olivaceae,

LICHENICOLOUS COELOMYCETES

47

Fig. 22 Nigropuncta rugulosa (IMI 241409 holotype). A, Infected areolae (x 30). B, Infected
areolae showing the emergent conidial masses (x 80). C, Vertical section of pycnidium (lower
part) to show the pycnidial wall and conidiogenous cells (x 1020). D, Conidia (x 1020).

3-4 yum latae, Conidia catenata, in cirrhum accumulata, irregulariter, e cellulis numerosis composita,
plerumque 20^0 /urn diam, atro-olivacea vel nigra, cellulis subglobosis vel angularibus, plerumque
simplicibus, rugulosis, plerumque 6-8 //m diam.

Typus: Austria, Steiermark, Schladminger Tauren, Kleinsolk-Obertal, west. Abhange der Kessel-
spitze, alt. 1700m, in thallo lichenis ignoti e saxis schistosis, 12. viii. 1974, /. Hafellner 3929 (IMI
241409 holotypus!; herb. Hafellner isotypus!).

Conidiomata pycnidial, arising singly, immersed in areolae of the host, the adjacent
thalline tissue sometimes proliferating to form a distinct raised margin around the spore
mass which recalls a thalline exciple, only exceptionally more that one pycnidium in each
areole, hyaline but black when viewed from above as a result of the dense spore mass,
1 00-450 //m wide; ostiolate, the ostiole equal to the width of the pycnidium in older
conidiomata, mainly circular but the margin sometimes invaginated or angular, not
clearly differentiated but forming by an irregular opening caused by an expansion of the
pycnidium, the pycnidia are at first subglobose or flask-shaped but become cupulate to
almost acervular with age; walls poorly demarcated from the host tissues, mainly hyaline but
sometimes becoming olive-tinged, especially towards the upper parts, 10-20/zm thick, the

48

D. L. HAWKSWORTH

10/im

Fig. 23 Nigropuncta rugulosa (IMI 241409 holotype). A, Conidiogenous cells and detail
of pycnidial wall. B, Conidiogenous chain with developing conidia. C, Cells squashed
from the irregular multicellular conidia.

outer part composed of 2-5 irregular layers of loosely to moderately compacted interwoven
hyphae, hyphae thick-walled, hyaline, 2*5-3 //m wide, inner part comprising thinner-walled
mainly isodiametric loosely packed pseudoparenchymatous cells, 2-3*5 /um diam, these
scarcely distinguishable from conidiophores. Conidiophores forming a compact layer lining
the inner wall of the pycnidial cavity, simple or sparsely branched, arranged parallel to
one another, hyaline, septate, 10-15 x2-3 - 5 //m, composed of almost isodiametric cells,
somewhat constricted at the septa. Conidiogenous cells holothallic, integrated, terminal,
determinate, subglobose to subcylindrical, resembling the conidiophore cells but sometimes
becoming olivaceous at the apex, 3^ /urn wide. Conidia arising in short basipetal chains,
tightly adhering to those produced from adjacent Conidiogenous cells to form multicellular
propagules, these very irregular in shape but mainly 20-40 //m overall, deep olive to almost
black, extruded through the ostiolar opening as a dense black cirrhus which is clearly seen
with a x 10 lens, individual cells mainly simple, thick- walled, subglobose to angular through
compression by adjacent cells, walls roughened with a very irregular granular-lacerate
ornamentation, this perhaps originating from exfoliating layers of wall tissue, cells mainly
6-8 jum diam.

Host: On an unidentified sterile grey areolate crustose lichen thallus. The areolae are not
discoloured in any way and algal cells close to the pycnidia retain their usual colour also.
The species is evidently parasymbiotic rather than parasitic, but could even be lichenized
(see below).

Distribution: Austria. Known only from the type collection with certainty, but perhaps
also present in Germany (see below).

Observations: This is a most distinctive species macroscopically and superficially recalling
Thelomma siliceum (Fee) Tibell (see Tibell, 1976: figs. 15-16); however, microscopic
examination soon reveals that it is not an ascomycete.

Within the host thallus, it seem impossible to distinguish hyphae of the pycnidium from

LICHENICOLOUS COELOMYCETES 49

those forming the tissues of the host. Further, most areolae have only a single pycnidium and
the pycnidia are evenly distributed over the material, and not in any way localized. These
facts, together with the consideration that the thallus and its algae are maintained in a
healthy condition, makes it necessary to speculate whether the present fungus is actually an
independent lichenized coelomycete. In the absence of additional material for study and
detailed ultrastructural investigations, it seems impossible to resolve this question at the
present time.

One factor which may support the parasymbiotic thesis is a report of what may be the
same fungus on Lecanora alphoplaca (Wahlenb.) Ach. by Gerber (1931 : 481 figs. 14-15).
This author illustrated an unnamed fungus on this host collected by Zopf at 'Oberwinkel in
Groden' which has some important similarities to Nigropuncta rugulosa, particularly in the
macroscopic appearance of the infection and the formation of dark-coloured multicellular
conidia which were very irregular in shape and composed of subglobose to angular cells.
Interestingly, he appears to have considered the structure referred to above called the
pycnidial wall as the invaginated cortical layer of the host. I have not been able to locate the
material Gerber studied, and from his paper alone cannot confidently assert that he had
the same taxon because sufficient information on dimensions and conidium formation was
not provided. If the two are conspecific, however, as Gerber had a host he could determine,
then N. rugulosa must be considered a parasymbiont and not an independent lichen.

The dark olive colour of the conidia is particularly characteristic and must be due to a
pigment (or pigments) different from those present in other dark-spored lichenicolous
Coelomycetes. The lacerate-granulose roughening of the conidia is also an unusual feature.
This may conceivably originate either from an exfoliation of the conidial wall layers, or be
wall remnants produced during conidiogenesis. The pigmentation precludes a more detailed
light microscopic study, and ultrastructural investigations are needed to firmly determine the
nature of this roughening and any relationship it might have to the process of conidiogenesis.

XVI. PHOMA Sacc. nom. cons.
Af/c/M?/ia2:4(1880).
See Sutton( 1980 : 3 78) for generic synonyms.

Conidiomata pycnidial, arising singly or aggregated, globose, immersed or partly
erumpent, brown to dark brown, ostiolate; walls usually thin, composed of a few layers (often
1-3) of cells, cells pseudoparenchymatous, pale brown to brown, more deeply pigmented
around the ostiole in many species, rather thin-walled, textura angularis. Conidiophores
normally absent. Conidiogenous cells enteroblastic, lining the pycnidial cavity, short-
cylindrical to doliiform, phialidic, not proliferating, hyaline. Conidia arising singly, not
catenate, variable in shape in different species, ellipsoid, subcylindrical, fusiform, pyriform
or globose, rounded at the apex and base or with the base slightly truncated, hyaline, simple
when mature (exceptionally becoming 1 -septate), sometimes distinctly guttulate, thin-
walled, smooth-walled.

Type species: Phoma herbarum Westend.

Teleomorph (perfect states): A wide range of loculoascomycetes are described as having
anamorphs referrable to Phoma, including species of Leptosphaeria Ces. & de Not. and
Pleospora Rabenh. ex Ces. & de Not. Most Phoma species, however, remain unconnected to
teleomorphs.

Number of species: According to Sutton (loc. cit.) many more than 2000 species have
been described in this genus, which was traditionally employed for all caulinicolous
Coelomycetes with small, hyaline, simple conidia. The genus is now employed for species
from many habitats (and includes taxa from a very wide range of substrates) and perhaps
really should only include about 50 species; a key to 27 is included in Sutton (loc. cit.). Five
lichenicolous species are now known.

50

D. L. HAWKSWORTH

10/im

Fig. 24 Phoma caloplacae (UPS holotype). A, Surface view of pycnidium. B, Surface
view of pycnidial wall. C, Conidiogenous cells and pycnidial wall. D, Conidia.

Observations: Phoma-\\ke fungi on lichens have often been placed in Phyllosticta Pers. ex
Desm., a genus formerly adopted for similar fungi on leaves as opposed to stems, presumably
in the belief that lichen thalli approximated more closely to 'leaves' than 'stems'. The type
species of Phyllosticta, however, is in any case very different, conforming to the concept of
Phyllostictina Sydow, and can not be used for such fungi (Punithalingam, 1974 : 60-6 1). The
lichenicolous species referred to Phoma here agree closely with the concept of Phoma
adopted by Sutton (loc. cit.}.

Modern work on species concepts in Phoma has emphasized studies of their behaviour in
pure culture and it would obviously be of interest to obtain the lichenicolous species in
culture and then compare the isolates with the known non-lichenicolous taxa.

Key to the lichenicolous species

1 Conidia broadly to narrowly ellipsoid 2

Conidia subglobose, (4-)5-6(-7) //m diam; on Caloplaca cerina . Phoma caloplacae (p. 50)

2(1) Conidia narrowly ellipsoid, usually less than 2-5 //m wide 3

Conidia broadly ellipsoid, 4-5 - 5(-6)x2'5-3'5(-4) J um; on Physcia aipolia and Physconia
pulverulacea Phoma physciicola (p. 56)

3(2) Conidia to 2 //m wide 4

Conidia (4-)4'5-6(-7) x 2-2'5(-3) /im; on Peltigera malacea .

4(3) Conidia 5-7 x 1-5-2 /zm; on Parmelia species

Conidia 3 "5-5 x 1 -5-2 //m; on an Vsnea species

Phoma peltigerae (p. 54)

Phoma cytospora (p. 5 1 )
Phoma dubia (p. 53)

1. Phoma caloplacae D. Hawksw. sp. nov.
(Fig. 24A-D)

Fungus lichenicola. Conidiomata singularia, uniloculata, dispersa, subglobosa, immersa sed
erumpescentia, atrobrunnea, (50-)70-100(-130)/^m lata, cum muris textura angulari usque 10-15 /im

LICHENICOLOUS COELOMYCETES 51

latis, e cellulis 5-8 x3-4//m. Cellulae conidiogenae enteroblasticae, acrogenae, subglobosae vel late
obpyriformes, phialidicae, non prolifericae, hyalinae, 5-6 //m diam. Conidia subglobosa, hyalina,
simplicia, laevia, (4-)5-6(-7) jum diam.

Typus: U.S.S.R., Guv. Jenisejsk, Stolba, lat. bor. 6020'N, in apotheciis Caloplacae cerinae (Ehrh. ex
Hedw.) Th. Fr., 1 . vii. 1 876, M. Brenner 1027b p.p. (UPS holotypus!).

Conidiomata pycnidial, immersed at first, the upper \ erumpent at maturity, arising
singly, scattered, dark brown, subglobose, (50-)70-100(-130)//m diam, ostiolate, cells
surrounding the ostiole dark brown, darker than the remaining wall which is subhyaline to
pale brown; wall 3-5 layers of cells thick, 10-1 5 um thick, cells pseudoparenchymatous, the
outermost brown to dark brown with the walls thickened, the inner subhyaline with thinner
walls, generally somewhat radially compressed and angular, 5-8 //m diam in surface view
and mainly 3-4 //m thick in section. Conidiogenous cells arising from the inner wall of the
pycnidium, lining the pycnidial cavity, subglobose to broadly obpyriform, hyaline,
smooth-walled, phialidic, not proliferating, 5-6 um diam. Conidia abundant, subglobose
but often somewhat angular due to mutual compression within the pycnidial cavity, hyaline,
simple, sometimes guttulate, smooth-walled, (4-)5-6(-7) um diam.

Host: Caloplaca cerina, apothecia. Infected discs become discoloured dark brown to
blackish and ascospore production is inhibited by the presence of the fungus.

Distribution: U.S.S.R. Known only from the type collection.

Observations: Readily separated from the other species accepted in the genus by the
subglobose to angular, and not ellipsoid, conidia.

2. Phoma cytospora (Vouaux) D. Hawksw., Trans. Br. mycol. Soc. 67 : 56 (1976).
(Fig. 25A-C)

Phyllosticta cytospora Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 193 (1914).
Type: France, Blainville, on Parmelia caperata, 4 March 1906, M. Missie (herb. Vouaux
neotype!).

Icones: Hawksworth & Punithalingam, Trans. Br. mycol. Soc. 61 : 62 fig. 4 (1973).

Conidiomata pycnidial, immersed, arising singly or more commonly in groups, scattered,
black, subglobose, 40-80 //m diam, ostiolate; wall 2-3 layers of cells thick, mainly
7-10/zm thick but to about \5 um near the ostiole, brown to dark brown, cells
pseudoparenchymatous, not markedly radially compressed, 5-1 um long and 3^ um tall,
the inner cells subhyaline to pale brown and with somewhat thinner walls. Conidiogenous
cells arising from the inner wall of the pycnidium, lining the pycnidial cavity, subglobose to
broadly obpyriform, hyaline, smooth- walled, phialidic, not proliferating, 2-3xl-2/zm.
Conidia abundant, narrowly ellipsoid to slightly curved, one apex slightly truncated,
hyaline, simple, sometimes 1-2 guttulate, smooth-walled, 5-7 x 1*5-2 um.

Hosts: Parmelia caperata (L.) Ach., P. laevigata (Sm.) Ach., P. perlata (Huds.) Ach., P.
reticulata Taylor, and P. sulcata Taylor, thalli. Necrotic decolourized patches 1-4 mm diam
are formed, each of which includes 5-18 pycnidia, the boundary with healthy tissue being
marked by a blackened zone of tissue 100-250 /^m wide; the patches are usually discrete
and damage localized but occasionally they may coalesce, particularly on P. reticulata,
killing large areas of the thallus. Reported by Vouaux (1914) and Keissler (1930) only on P.
caperata.

Distribution: British Isles, France, and possibly Germany (see p. 83).

Observations: The decolourized necrotic patches produced by this species recall those
formed on the same or allied hosts by Cornutispora lichenicola and Lichenoconium erodens.

Phyllosticta cytospora was originally described from material of Parmelia caperata
collected in the Toret de Vitrimont en Meurthe-et-Moselle' by Vouaux. This material is no
longer present in Vouaux's herbarium (Rondon, 1970) but there is a further collection made

Fig. 25 Phoma cytospora (IMI 164975). A, Vertical section of pycnidium. B, Conidio-
genous cells and pycnidial wall. C, Conidia. Reproduced from Hawksworth & Punithalingam
(1973:62).

LICHENICOLOUS COELOMYCETES

53

25/im

Fig. 26 Phoma dubia (E holotype). A, Vertical section of pycnidium. B, Conidiogenous
cells and pycnidial wall. C, Conidiogenous cells. D, Conidia.

at Blainville in 1906 labelled 'Phyllosticta cytospora mihi' present which agrees in all details
with Vouaux's original description and later collections; the Blainville specimen is
consequently designated as neotype for this name here.

The name Phyllosticta lichenicola Allescher may have been partly based on a fungus very
close to Phoma cytospora (see p. 83). Phoma caperatae Vouaux, however, seems unlikely
to represent P. cytospora (see p. 80).

Additional specimens: British Isles: England, S. Devon, Slapton Ley Nature Reserve, on Parmelia
perlata on Quercus, 30 March 1972, D. L. Hawksworth 2695 (IMI 164975!), loc. cit., on P. reticulata on
Acer pseudoplatanus, 25 August 1978, D. L. Hawksworth 4772 (IMI 231655!), loc. cit., on P. sulcata, 25
August 1976, D. L. Hawksworth 4322 (IMI 206364!); S. Devon, Killerton Park, on P. reticulata on
Quercus, 3 September 1978, D. L. Hawksworth 4849 (IMI 231732!); S. Devon, Start Point, nr
Langerstone Point, on P. perlata on Salix, 27 August 1972, D. L. Hawksworth 3075 (IMI 168590!); S.
Devon, Cornwood, Dendles Wood, on P. laevigata, 29 February 1976, D. L. Hawksworth 4293a (IMI
20 166 la). Scotland, Dumfriesshire, nr Gretna, on P. sulcata on Ulmus, 6 August 1972, B. J. Coppins
(E!); Argyll, S side Loch Sunart, on P. laevigata, 14 August 1972, S. R. Davey(lMl 250032b).

3. Phoma dubia (Lindsay) Sacc. & A. Trotter, in Saccardo, Syll. Fung. 22 : 897 ( 1 9 1 3)*.
(Fig. 26A-D)

*Phoma dubia Rupprecht (Sydowia 11 : 128, 1958), described from Stellaria graminea in Hungary, is a later
homonym which must be rejected under Art. 64. 1 .

54 D. L. HAWKSWORTH

Phymatopsis dubia Lindsay, Trans. R. Soc. Edinb. 24 : 442 (1866).

Type: New Zealand, Otago, Pelichet Bay, in the bush, on Usnea sp. (apothecia), 17 December
1861, W. L. Lindsay (E holotype!).

Icones: Lindsay, Trans. R. Soc. Edinb. 24 : pi. 30 figs. 36^1 (1 866).

Conidiomata pycnidial, immersed in the hymenium of the host, only the ostiole
sometimes slightly protruding above the level of the epithecium, scattered to loosely
aggregated, sometimes almost confluent, subglobose, 25-60 um diam, ostiolate; wall mainly
1-3 layers of cells thick, mostly 3'5-5/zm thick but to 8 jam near the ostiole, cells
pseudoparenchymatous, subglobose to somewhat polyhedral, subhyaline except in the
vicinity of the ostiole and there thicker-walled and golden-brown, mainly 2-3 um diam.
Conidiogenous cells arising from the inner wall of the pycnidium, short-ampulliform to
obpyriform, hyaline, smooth-walled, phialidic, not proliferating, 4-8 x 3-5 um. Conidia
abundant, narrowly ellipsoid to almost bacilliform, sometimes slightly truncated at the base,
hyaline, simple, 0-2 guttulate, smooth-walled, 3'5 x 1*5-2 um.

Host: Usnea sp., apothecia. The host was originally named as U. barbata var.florida (L.)
Fr. (i.e. U.florida (L.) Wigg.) by Lindsay but I am not sure that it really belongs there; certain
identification must await a re-evaluation of that genus in New Zealand. The infected parts of
the apothecial disc become discoloured and brownish suggesting that Phoma dubia is a mild
pathogen.

Distribution: New Zealand. Known only from the original collection.

Observations: This fungus is clearly separated from the other lichenicolous Phoma species
treated here by the minute conidia.

4. Phoma peltigerae (P. Karsten) D. Hawksw., Trans. Br. mycol. Soc. 74 : 381 (1980).
(Fig. 27A-E)

Phyllosticta peltigerae P. Karsten, Hedwigia 23 : 62 (1884).

Type: Finland, Tavastia australis, Tammela, Kyto, on Peltigera malacea, 4 May 1869, P. A.
Karsten (H-KARST 2040 holotype!).

Conidiomata pycnidial, completely immersed at first, the upper | erumpent at
maturity, arising singly, scattered, black, subglobose, (75-)100-150(-200)//m diam,
ostiolate, cells surrounding the ostiole dark brown to black and darker than the remainder of
the wall; wall 1-2 layers of cells thick, 5-8 //m thick, cells pseudoparenchymatous, the
outermost dark brown with the external wall thickened, not markedly radially compressed,
5-9 um long and 4-7 um tall, the inner cells subhyaline to pale brown and with thinner
walls. Conidiogenous cells arising from the inner wall of the pycnidium, lining the pycnidial
cavity, subglobose to broadly obpyriform, hyaline, smooth-walled, phialidic, not pro-
liferating, 4-6 um diam. Conidia abundant, narrowly ellipsoid, rounded at the apices,
hyaline, simple, sometimes guttulate, smooth-walled, (4-)4'5-6(-7) x 2-2'5(-3) um.

Host: Peltigera malacea (Ach.) Funck, thallus. The host in the type collection was
originally determined as P. canina (L.) Willd., and this has been given as the host in all
previous references to this species. On P. malacea the infected area becomes entirely
decolourized, the whitened patches being 5-10 mm across and very sharply delimited from
the healthy tissues; a slight blackening of the host occurs just at the margin of the infection
spot. Phoma peltigerae consequently appears to be pathogenic to this species. This fungus
has also been reported from P. horizontalis (Huds.) Baumg., P. polydactyla (Necker) Hoffm.
and P. rufescens(Weis)Humb. by Vouaux(1914 : 193), and Keissler(1930 : 536) mentions a
report on Platismatia glauca (L.) Culb. & C. Culb.; these four host records require
substantiation.

Distribution: Finland. Known to me only from the type collection but mentioned as
present in France by Bouly de Lesdain (1910:276) and Vouaux (1914: 193), and in
Germany by Keissler(1930 : 536, 1933 : 392).

LICHEN1COLOUS COELOMYCETES

55

Fig. 27 Phoma peltigerae (H-KARST 2041 holotype). A, Surface view of pycnidium. B,
Vertical section of pycnidium. C, Surface view of pycnidial wall. D, Conidiogenous
cells and pycnidial wall. E, Conidia.

Observations: Phoma peltigerae recalls P. cytospora but differs in the broader conidia,
thickening of the pycnidial wall cells, and larger pycnidia, as well as in occurring on quite
different hosts.

Phyllosticta peltigerae was originally described from 'Mustiala' but no material under this
name from that locality could be found in H-KARST (O. Vitikainen, in litt.}. However, in a
packet incorrectly labelled as "Stagonopsis peltigerae' in W. Nyberg's hand (an amateur
mycologist who once helped in H) was a slip of paper with the herbarium name 'Phoma
peltigerae' in Karsten's writing, conidial characters and notes conforming to the published
description of Phyllosticta peltigerae, and the locality name Kyto. This is a place name in
Tammela parish, situated west of Mustiala (O. Vitikainen, in lift.), and it appears that when
publishing this fungus Karsten (a) used a nearby better-known locality name, and (b) as
Phoma and Phyllosticta were then separated according to whether the fungus was on a stem
or leaf, decided that the thallus of Peltigera was leaf-like not stem-like. H-KARST 2040 is
consequently considered to be the holotype of Phyllosticta peltigerae.

56

D. L. HAWKSWORTH

10/mi

Fig. 28 Phoma physciicola (W 1910/609 holotype). A, Vertical section of pycnidium. B,
Conidiogenous cells and pycnidial wall. C, Surface view of pycnidial wall. D, Conidi-
genous cells. E, Conidia.

5. Phoma physciicola Keissler, Hedwigia 50 : 294 (1911).
(Fig. 28A-E)

Type: Austria, Steiermark, Gams bei Hieflau, on Physcia aipolia, June 1910, K. von Keissler (W
1910/609 holotype!).

Icones: Keissler, Hedwigia 50:295 fig. la-c (191 1). Keissler, Rabenh. Krypt.-Fl. 8:539 figs.
102-103(1930).

Conidiomata pycnidial, immersed at first, with the upper \ erumpent at maturity, arising
singly or in small groups, black, subglobose, 100-150 jum diam, ostiolate, cells surrounding
the ostiole dark brown to black and darker than the remainder of the wall; wall 1-3 layers of
cells thick, mainly 7-10 /zm thick but to 15 /urn near the ostiole, cells pseudoparenchy-
matous, the outermost brown with moderately thickened walls, not markedly radially com-
pressed, 5-7 /i m diam in side view, 5-10/zm diam and rather angular in surface view, the
inner cells pale brown to subhyaline and with thinner walls. Conidiogenous cells arising
from the inner wall of the pycnidium, lining the pycnidial cavity, subglobose, hyaline,
smooth-walled, phialidic, very rarely proliferating, 4-7 //m diam. Conidia abundant,
broadly ellipsoid, rounded at the apices, hyaline, simple, frequently with 2 or more guttules,
smooth-walled, 4-5'5(-6) x 2-5-3'5(-4) //m.

Host: Physcia aipolia (Humb.) Fiirnrohr and Physconia pulverulacea Moberg, apothecia.
Also cited by Keissler (1930 : 542) from Baeomyces rufus (Huds.) Rebent. (thallus),
Parmelia caperata (L.) Ach. (thallus) and P. glabratula (Lamy) Nyl. (thallus) but these
reports are dubious and require confirmation. In the holotype, as many as 1 5 pycnidia may
occur in a single apothecium of the host in which they are rather strictly confined to the
thecium, scarcely extending down into the hypothecium. Infected apothecia retain their
black colour but lose their characteristic whitish pruina; ascospores still formed in the most
heavily infected apothecia studied.

LICHENICOLOUS COELOMYCETES 57

Distribution: Austria and Sweden.

Observations: Phoma physciicola is separated from other species of the genus accepted
here by the shape and size of the conidia. Phoma caperatae Vouaux was treated first as a
variety of P. physciicola and later subsumed under this species by Keissler (1930 : 542), but it
is most unlikely that these taxa are conspecific (see p. 80).

Additional specimens: Sweden: Smaland, Aneboda s:n, Aneboda limnologiska laboratorium, pa
lonn, i alle, on Physconia pulverulacea, 27 July 1947, R. Santesson (UPS!); Sodermanland, Bankyska
s:n, Sturehav, alle trad, on P. pulverulacea, 21 July 1944, R. Santesson (UPS!).

XVII. PSEUDOSEPTORIA Speg.

An. Mus. nac. B. Aires 20 : 388 (1910).
Lunospora Frandsen, Meddr Kgl. Veter. Landboh. Kobenh. 26 : 70 (1943).

Conidiomata pycnidial, arising singly or aggregated into short rows, globose, immersed to
erumpent, brown to black, ostiolate; walls composed of a few layers of cells, cells
pseudoparenchymatous, thin- to moderately thick-walled, pale to greenish or dark brown,
textura angularis. Conidiophores absent. Conidiogenous cells holoblastic, lining the inner
wall of the pycnidial cavity, acrogenous or sympodial, subcylindrical to ampulliform, often
proliferating, then with distinct annellations, hyaline. Conidia not catenate, falcate, fusiform
or cymbiform, attenuated at the apex, attenuated to truncate at the base, hyaline, simple,
smooth-walled.

Type species: Pseudoseptoria donacis (Pass.) B. Sutton (syn. P. donacicola Speg.).

Number of species: Four species are treated by Sutton (1980), all of which are
graminicolous. A fifth lichenicolous species is added to the genus here.

Observations: The genus Pseudoseptoria has been revived by Sutton (1980) for several
graminicolous species hitherto referred to Selenophoma Maire, a genus that differs in both
pycnidium structure and the method of conidiogenesis. The inclusion of Phoma usneae
Vouaux in the genus expands the generic concept slightly to embrace species in which the
conidia are not strongly falcate. It is probable that some additional non-lichenicolous and
non-graminicolous will eventually be found to require transfer here.

1. Pseudoseptoria usneae (Vouaux) D. Hawksw. comb. nov.
(Fig. 29A-B)

Phoma usneae Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 196 (1914).

Type: Italy, Tirol, Bolzano [Bolzen], Mendola [Mendel], wood by Mt Roen [Roen], on Usnea
filipendula aggr. (thallus), 13 August 1896, F. G. C. Arnold [Lich. Exs. no. 1718 p.p.]
(K isotype!).

Exsiccatae: Arnold, Lich. Exs. no. 1718 p.p. (K!; sub Epicoccum usneae Anzi).

Conidiomata pycnidial, immersed at first but becoming erumpent with age, arising singly,
scattered, black, subglobose, ostiolate, 75-100(-200)//m diam when mature; wall about 3-4
layers of cells thick, cells moderately thick-walled, pseudoparenchymatous, forming a
textura angularis, subglobose to polyhedral, greenish brown, 4-7 um diam. Conidiogenous
cells holoblastic, lining the inner wall of the pycnidial cavity, subcylindrical to ampulli-
form, percurrently proliferating, to 4 annellations seen, hyaline, smooth-walled, rarely
proliferating sympodially, 6-12x3-5/zm. Conidia cymbiform, tapering to the rounded
apex but with an abruptly truncated base, base 2-3 urn wide, hyaline, smooth-walled,
simple, often with numerous small guttules, (10-)12-14(-16) x (3-)3'5^ urn.

Host: Usnea filipendula Stirton aggr., thallus. The fungus is present on some of the smaller
branches but very few pycnidia were found on the isotype studied. Infection, at least in its
early stages, does not appear to damage the host significantly. The apothecia in the type

58

D. L. HAWKSWORTH

Fig. 29 Pseudoseptoria usneae (K isotype). A, Conidiogenous cells. B, Conidia. C, Surface view

of cells of pycnidial wall.

collection are infected with Lichenoconium usneae (Anzi) D. Hawksw., and some dark
brown torulose mycelium and galls that may be young Abrothallus usneae Rabenh. are
present on other parts of the thallus.

Distribution: Italy and Spain (fide Santesson, 1 960 : 519).

Observations: Vouaux described this taxon on the basis of the example of Arnold's
exsiccatum in Harmand's herbarium. This could not be located amongst Harmand's
herbarium in Angers (M. Guerlesquin, in lift.) but one collection of this number in K was
found to have a few pycnidia producing conidia conforming so precisely to Vouaux's
description that there can be no doubt they belong to the same fungus. The only
discrepancies noted were that Vouaux indicated the pycnidia were 1 00-200 //m diam and
that he gave the conidiogenous cells as only 3-6 x 2 um. So few pycnidia were present on
the specimen in K that no microtome sections were prepared; a more detailed account of the
pycnidium structure must consequently await the availability of further material of this
taxon.

This species occupies a rather isolated position in Pseudoseptoria as currently
circumscribed in view of the relatively broad and straight, rather than curved or falcate,
conidia. Such a difference scarcely merits a generic separation, however, as it seem to agree
with the type species of the genus in all other details.

LICHENICOLOUS COELOMYCETES 59

Keissler (1960 : 416) indicated that this fungus could be distinguished from the normal
pycnidia of Usnea species in that the conidiogenous cells were simple, but there are many
other differences also. For example, the conidia of Usnea are, contrary to some earlier
reports, bacillariform with a marked swelling just above the base so that they appear
pendulum-shaped (Swinscow & Krog, 1976 : 264-5).

XVIII. PYRENOTRICHUM Mont.

Annls Sci. nat. (Bot.) II 20 : 376 (1843).

Chlorocyphella Speg., An. Mus. nac. B. Aires 19 : 279 (1909).

Conidiomata ('campylidia', 'orthidia') helmet-shaped, erect, convoluted or becoming
adpressed, superficial, arising singly, not aggregated into stromata, pale fawn to shades of
blue-green or blue-black, wall thick at the base but tapering towards the tip, outer convex
surface composed of arachnoid hyphae, central tissue and inner concave surface of hyaline to
pigmented thick-walled pseudoparenchymatous cells, textura angularis. Conidiophores
mainly in the lower part of the concave surface, simple or branched near the base, septate or
unicellular, conidiogenous cells arising at the same level, ellipsoid to subcylindrical.
Conidiogenous cells probably enteroblastic, phialidic, not proliferating, lacking distinct
collarettes, hyaline, smooth-walled, subcylindrical. Conidia arising singly, ellipsoid,
pyriform, clavate, filiform or scolecosporous, sometimes variously curved, 0-multiseptate,
not or distinctly branched, hyaline, smooth-walled.

Type species: Pyrenotrichum splitgerberi Mont.

Number of species: Three species have validly published names in Pyrenotrichum, and
Santesson (1952 : 41) recognized six additional species which have not yet been formally
published. All these nine species are parasymbionts on tropical foliicolous lichens and it is
probable that there are additional species awaiting description on other crustose lichens on
bark in the tropics (see below).

Observations: Pyrenotrichum species are evidently widespread in the tropics and have
been studied in some detail by Santesson; unfortunately only a brief account of his important
investigations has so far appeared (Santesson, 1952:40^11). I have not attempted to
duplicate his research and consequently describe in full only the type species of the genus
and note one of the other species with a validly published name represented in IMI. In
addition to these two species, P.foliicola mentioned under P. staurosporum, and P.filiferum
mentioned under P. splitgerberi, Santesson accepts five species with simple or 1 -septate
conidia; ellipsoid conidia occur in P. bicolor R. Sant. ined. (on Lopadium flammeum Mull.
Arg.), P. mirum R. Sant. ined. (on L. tayabasense (Vainio) Zahlbr.), and P. atrocyaneum R.
Sant. ined. (on Lopadium sp.); and pyriform to clavate conidia in P. irregulare R. Sant.
ined. (on four Sporopodium species), and P. podosphaera R. Sant. ined (on Sporopodium
sp.).

The genus is evidently not restricted to foliicolous lichens as I have seen two collections
referrable to it on sterile crustose lichens on bark from the tropics (Ecuador: Galapagos
Island, Isla Pinta, near volcano in south-central part, on Tournefortia sp., 8 July 1976, H.
Sipman, COLO-L63608, IMI 214571!; St Helena: Ascension Island, Green Mountain, nr
summit, on Podocarpus sp. bark, 30 October 1976, P. W. James, IMI 251280!). Whether
these represent additional new species is unclear, resultant upon the publication of
San lesson's revision.

It is possible that some of the allegedly lichenized foliicolous Coelomycetes described by
Brazilian workers in recent years also belong to Pyrenotrichum as interpreted by Santesson,
for example Acleistomyces rionegrensis Bat. & Maia with ellipsoid, simple, conidia (Batista,
1961). The status of many of the taxa introduced by these workers requires a critical
re-evaluation; the generic names involved are compiled by Vobis & Hawksworth (1981).

Keissler (1927: 159) listed generic names 'Campylidium Mull. Arg.' and 'Orthidium
Mull. Arg.' as synonyms of Chlorocyphella; this is quite incorrect as Muller (1881 : 111,

60 D. L. HAWKSWORTH

1 890 : 202) introduced these as anatomical terms for conidiomata of different shapes and not
as generic names. Neither of these names is validly published as a genus.

I found the method of conidiogenesis in P. splitgerberi difficult to ascertain conclusively,
even by differential interference contrast. Although the conidia are often large, at the point of
secession a structure only about 1*5 //m wide must be resolved; ultrastructural studies will
be necessary to conclusively determine this matter.

Key to the species treated

1 Conidia unbranched, filiform, multi-septate, arcuate to C- or S-shaped or variously

convoluted, 55-80 x 2-3 //m Pyrenotrichum splitgerberi (p. 60)

- Conidia with a main stem and 1-&t branches arising at a single locus about halfway along the
conidum, arms at acute angles, straight to slightly flexous, 60-85 x 2'5-3'5 um

Pyrenotrichum staurosporum (p. 63)

1. Pyrenotrichum splitgerberi Mont., Annls Sci. nat. (Bot.) II, 20 : 377 (1843).
(Figs. 30A-F,31A-B)

Lee idea irregular is Fee, Bull. Soc. hot. Fr. 20 : 3 1 8 ( 1 873).

Melophia woodsiana Sacc. & Bed., in Saccardo, Syll. Fung. 3 : 659 (1884)*.

Cyphella subcyanea Ell. & Ev., J. Mycol. 2 : 37 (1886).

Cyphella aeruginascens P. Karsten, Hedwigia 28 : 191 (1889).

Chlorocyphella aeruginascens (P. Karsten) Keissler, Annln naturh. Mm. Wien 41 : 1 59 ( 1 927).

? Trichosperma cyphelloidea Hohnel, Sber. Akad. Wiss. Wien 1, 116 : 145 (1907)*.

ITrichosperma aeruginosa Hohnel, Sber. Akad. Wiss. Wien 1, 118 : 1530 (1909)*.

Chlorocyphella subtropica Speg., An. Mus. nac. B. Aires 19 : 279 (1909).

Chlorocyphella aeruginascens var. convoluta Keissler, Annln naturh. Mus. Wien 41 : 161 (1927).

Chlorocyphella lichenicola Keissler, in Zahlbruckner et al, Trans. Proc. N.Z. Inst. 59 : 3 1 3 (1928).

Icones: Keissler, Annln naturh. Mus. Wien 41 : 161 fig. \a-b (1927). Mameli-Calvino, Nuovo G.
hot. Hal II, 37: pi. 19 figs. 1-16 (1930). Montagne, Annls Sci. nat. (Bot.) II, 20 : pi. 16 fig.
2a-m (1 843). Rizzini, Archos Jard. hot., RiodeJ. 12 : figs. 1-7(1952).

Conidiomata helmet-shaped, erect, convoluted, or geniculately adpressed, entirely
superficial, almost fawn to blue-green or blue-black, mat, arising singly, scattered, often
rather evenly and sparsely distributed over the host thallus, very variable in size and shape,
mainly 300-800 //m long and 250-600 /zm wide when mature; wall 40-60 /zm thick at the
base but tapering to about 10 //m thick at the tip, in vertical section seen to be
composed of two layers, an irregular outer layer of arachnoidly branched thick-walled
hyphae 3-3 '5 jam wide with granular incrustations often evident on their walls, and air-
spaces present in the gaps formed by the hyphae, inner layer of thick- walled pseudoparen-
chymatous cells, textura angularis, cells irregularly subglobose to ellipsoid or polyhedral,
mainly 4-9 //m diam, without air spaces, subhyaline to olivaceous, the cells towards the
inner concave side adjacent to the conidiogenous layer most deeply pigmented, those above
the conidiogenous region often with papilliform projections. Conidiophores simple or
branched near the base, septate or unicellular, hyaline, ellipsoid to subcylindrical, with 1-3
conidiogenous cells arising at about the same level, to 10x3 //m, restricted to the concave
surface of the conidiomata and not extending more than about j of the length of the
conidiomata in the specimens sectioned. Conidiogenous cells probably enteroblastic,
phialidic with minute apical thickenings, arising from the conidiophores or sometimes
apparently directly from the innermost wall cells, hyaline, smooth-walled, subcylindrical,
slightly wider at the base but not ampulliform, mainly 10-12 x 2'5-3-5(-4)//m. Conidia
abundantly produced, arising singly, filiform, scolecosporous, arcuate to C- or S-shaped or

*Santesson (1952:52) attributes combinations of these three epithets into Pyrenotrichum to Hohnel; Hohnel
(1910 : 652) suggested these species should be placed in that genus but did not validly publish the transfers (they were
also not accepted in his 'Index', loc. cit.: 678).

LICHENICOLOUS COELOMYCETES

v , : V"A- - T > :

v -u: p- V^*^

Fig. 30 Pyrenotrichum splitgerberi (A-B, D-F, IMI 44258;; C, IMI 246129). A-B, Dark
conidiomata (x. 16). C, Light conMiomata (x25). D, Vertical section of conidoma showing
portion with conidiogenous cells (left) and papillae not bearing conidia (centre to right) (x 1020).
E, Vertical section of conidioma (x 250). F, Vertical section of lower part of conidioma showing
the branched conidiophores (x 1020).

62

D. L. HAWKSWORTH

Fig. 31 Pyrenotrichum splitgerberi (IMI 44258;). A, Conidiophores, conidiogenous cells

and developing conidia. B, Conidia.

otherwise convoluted, slightly thicker at the rounded apex, truncated and to about 1 //m
wide at the extreme base, hyaline, multiseptate, with to 8 septa when mature,
often with numerous guttules, smooth-walled, 55-80 x 2-3 //m.

Hosts: According to Santesson (1952:40) this species is restricted to the thalli of
foliicolous lichens belong to the genera Lopadium Korber (9 species) and Tapellaria Mull.
Arg. (6 species). Parasymbiotic, sometimes apparently depressing the formation of apothecia

LICHENICOLOUS COELOMYCETES 63

in the host and then appearing as if a lichen-forming coelomycete. This species has been
considered to be a lichen by several authors (e.g. Mameli-Calvino, 1930; Rizzini, 1952) and
it may be that in some cases the fungus can take over the algae from the original lichen
thallus it invades, so becoming lichenized (cf. Blarneya hibernica D. Hawksw. et al;
Hawksworth et al., 1980). A critical anatomical investigation of the fungus-alga relationship
in material of this species on 'sterile' thalli lacking apothecia would be of interest.

Distribution: Probably pantropical. There are reliable reports at least from Brazil, China,
Cuba, Ghana, New Zealand, Surinam, and the U.S.A. (Louisiana).

Observations: I have not carried out a critical investigation into the range of variation of
this species in view of Santesson's unpublished studies; the above description is mainly based
on IMI 44258) (determined by Santesson in 1951). The synonymy presented above is derived
from Keissler (1927, 1933), Santesson (1952 : 49-53), and a few minor sources; I have not
studied any of the relevant type collections. Santesson (1952 : 41) mentions only one other
species of the genus as having filiform unbranched and multiseptate conidia, P.filiferum
R. Sant. ined., known from two species of Lopadium; the characters separating this species
from P. splitgerberi were not indicated.

Specimens: Brazil: Rio de Janeiro, Jardin Botanico, on Buxus sempivirens, 1947, C. T. Rizzini 11
(PC!); Parana, Desvio Ypiranga, 26 September 1909, P. Dusen 8822 (PC!); sine loc., comm. March
1951, comm. R. Santesson (IMI 45098!). Ghana: Pokoasi, on Diospyros sp., 1 June 1949, S. J.
Hughes 1040 (IMI 43725c); loc. cit., on Lopadium sp. on Pancovia bijuga, 25 May 1949, 5 1 . J. Hughes
834 (IMI 44258J!); Aburi, on Trycalysia cf. pallens, 24 May 1949, S. J. Hughes 807 (IMI 441 70d!);
near Nsuaem, 8 May 1949, S. J. Hughes 219 (IMI 442 196!); Togoland, Jasikan, on Cola caricifolia, 27
May 1949, S. J. Hughes 957 (IMI 43662g!); Togoland, Hobae, on Culcasia scandens, 28 May 1949,
5. J. Hughes 934b (IMI 442 13a!). U.S.A.: Louisiana, Assumption Parish, 2-3 mis W of Grand Bayou,
on Sabal palmetto, 8 February 1976, S. C. Tucker 1 5448 (IMI 246129!).

2. Pyrenotrichum staurosporum (Keissler) R. Sant., Symb. hot. upsal. 12(1): 41 (1952).
(Fig. 32A-B)

Chlorocyphella aeruginascens var. staurospora Keissler, Annln naturh. Mus. Wien 41 : 162 (1927).
Icones:KQiss\Qr, Annln naturh. Mus. Wien4\ : 161 fig. lc(1927).

This fungus is very similar in habit to Pyrenotrichum splitgerberi but is clearly separated
from that species in that the conidia are branched with 2-4 arms arising at a single locus from
a point about halfway along the total length of the conidium. In the specimens studied, the
conidia were (overall) 60-85 x 2'5-3'5 //m.

Hosts: According to Santesson (1952 : 40) this species is restricted to foliicolous lichens
belong to the genus Lasioloma R. Sant. (4 species). Parasymbiotic.

Distribution: This taxon was first described from Guam but I have seen specimens also
from Ghana and Malaysia.

Observations: This species is clearly separated from Pyrenotrichum splitgerberi on the
basis of its branched conidia as well as the disparate hosts. Santesson (1952 : 4 1 ) transferred a
second species with branched conidia to Pyrenotrichum, P. foliicola (Vainio) R. Sant.*; this
was first described from the Phillipine Islands and occurs on foliicolous species ofBacidia de
Not. (5 listed by Santesson, loc. cit.: 40). I have not seen any material of P. foliicola. Although
Santesson did not give the basionyms of P. foliicola nor P. staurospoum, his combinations
are treated as validly published here as these taxa were clearly accepted as distinct, albeit in a
Table. As Santessons's monograph appeared before 1 January 1953, Art. 33.2 does not
apply.

*Symb. hot. upsal. 12(1): 41 (1952). Cyphella foliicola Vainio, Annls Acad. Sci. fenn. 15(6): 83 (1921).
Chlorocyphella foliicola (Vainio) Keissler, Annln naturh. Mus. Wien4\ : 159(1927).

64

D. L. HAWKSWORTH

Fig. 32 Pyrenotrichum staurosporum (IMI 444 \Qd). A, Conidiogenous cells and developing

conidia. B, Conidia.

Specimens: Ghana: Aburi, on Landolphia owariensis, 24 May 1949, S. J. Hughes 791 (IMI
444 \0d). Malaysia: Cameron Highlands, on Lasioloma arachnoideum on Xanthophvllum affma, 6
September 1953, W. J. Cherwick [A. Johnston no. 1113] (IMI 54915 f-g\).

XIX. VOUAUXIELLA Petrak & H. Sydow

Beih. Repert. nov. Spec. Regni veg.42 : 482 (1927).
Alysia Cavalc. & Silva, Publcoes Inst. Micol. Recife 647 : 32 (1972).

LICHENICOLOUS COELOMYCETES 65

Conidiomata pycnidial, arising singly, subglobose to almost cupuliform, immersed or
semi-immersed, dark brown to black; walls composed of thick-walled dark brown to
olivaceous green cells forming a textura angularis but sometimes almost hyphal below, the
upper cells separating to form an irregularly delimited ostiole. Conidiophores short,
branched only at the base, septate, pale brown, smooth or verruculose, lining the inner wall
of the pycnidium. Conidiogenous cells integrated or discrete, terminal, holothallic,
subhyaline to pale brown or olivaceous green, smooth or verruculose, a basal thicker-walled
'mother' cell present in some species and becoming flared at the onset of conidiogenesis.
Conidia catenate, forming in acropetal chains, chains simple, with up to 8 or more adhering
conidia, ellipsoid with both ends abruptly truncated (except in the apical conidium and in V.
uniseptata), usually simple, rarely 1 -septate, olivaceous green, brown or dark brown,
smooth or verruculose, liberated by the breakup of the chains, dry.

Type species: Vouauxiella verrucosa (Vouaux) Petrak & H. Sydow (lectotype; selected by
Clements & Shear, 1931 : 378).

Number of species: Four, all obligately lichenicolous, and one of which may eventually
prove not to be congeneric.

Observations: Vouauxiella has an especially interesting method of conidiogenesis
(Morgan-Jones, 1971; Sutton, 1980); indeed it is the only non-stromatic coelomycete with
holothallic conidiogenesis recognized, with the possible exception of the new genus
Nigropuncta described above (p. 46). As noted by Hawksworth (1978a : 185) and Sutton
(1980 : 24), Vouauxiella uniseptata may well merit recognition as a distinct genus on the
basis of the absence of conidiophore mother cells, much more deeply pigmented and
1 -septate conidia, and the broadly opening pycnidia. Vouauxiella does, however, provide a
convenient temporary niche for that fungus pending more detailed studies on conidiogenesis.

Key to the species

1 Conidia simple

- Conidia 1-septate, (10-)12-15(-17)x 5-7(-8)//m, dark brown, smooth to weakly

verruculose; pycnidia almost cupuliform, 40-70 //m diam . Vouauxiella uniseptata (p. 66)

2 Conidia smooth-walled

Conidia distinctly verruculose, 6-9 x 3 -5-5 //m .... Vouauxiella verrucosa (p. 67)

3 Conidia (5-)6-8(-9) x 3-4 //m; pycnidia mainly 60-90 /zm diam

Vouauxiella lichenicola (p. 65)
Conidia 3-3*5 x 2'5//m [but see below]; pycnidia 70-1 60//m diam

Vouauxiella pithospora (p. 66)

1. Vouauxiella lichenicola (Lindsay) Petrak & H. Sydow, Beih. Repert. nov. Spec. Regni veg.
42:484(1927).

This species was described and illustrated in colour in great detail by Lindsay (18690 : pi.
23 figs. 1-18), although he also included some other taxa in his concept. Its conidiogenesis is
treated by Morgan-Jones (1971), and illustrations have been published more recently by
Hawksworth (1976 : 58) and Sutton (1980 : 25). This well-known fungus is not therefore
described in detail here because its diagnostic features will be apparent from the generic
description and key.

The typification and synonymy of Lindsay's epithet is discussed by Hawksworth
1979:287).

Hosts: Lecanora species, apothecia. Especially frequent on L. chlarotera Nyl. but reported
from a range of other red-fruited species of the genus. The pycnidia of this species and
Vouauxiella verrucosa are characteristically found at the edges of the disc adjoining the
thalline margins; infected discs are commonly paler in colour than is usual for the lichen and
piebald in appearance. Reports from other host genera are dubious, partly due to confusion

66 D. L. HAWKSWORTH

with Lichenodiplis lecanorae, a very different fungus (p. 38), to which taxon references to it
in Keissler (1930 : 568) on at least Calopaca cerina (Ehrh. ex Hedw.) Th. Fr., Lecanora
dispersa (Pers.) Sommerf, and Pertusaria leioplaca DC., should be referred. The report of V.
lichenicola on Caloplaca holocarpa (Hoffm.) Wade by Werner (1969 : 204) may also belong
to that fungus.

Distribution: Probably widespread, at least in Europe. Reliably reported from the British
Isles (England, Ireland, Scotland), France (Vouaux, 1914:297), Germany, Morocco
(Werner, 1970, 1972), Spain (Santesson, 1960 : 519), Sweden (Santesson, 1949 : 143) and
the U.S.A. (Mass.; Santesson, 1960 : 519). Material from Finland distributed under the name
V. lichenicola by Rasanen (Lich. Fenn. Exs. no. 1 50, BM!) is a Taeniolella species.

Specimens (all on Lecanora chlarotera apothecia): British Isles: S. Devon, Slapton, Slapton Ley
Nature Reserve, 30 July 1973, D. L. Hawksworth 3395 (IMI 178325!); loc. cit., 10 October 1974, D. L.
Hawksworth 3930 (IMI 188759!); loc. cit., Peasdish, 23 August 1980, D. L. Hawksworth 5044c (IMI
251263!); Oxfordshire, Hook Norton, January 1973, H. J. M. Bowen (IMI 185887!); Sussex, Eridge Old
Park, 20 May 1972, F. Rose (E!). Scotland, Kincardineshire, The Burn, Fettercairn, 26 July 1972, B. J.
Coppins et al. (E!); Perthshire, Loch Tay, June 1856, W. L. Lindsay (E lectotype!); E. Lothian, near
Dunglass, Cockburnpath, June 1 856, /. M. Lindsay (E!).

2. Vouauxiella pithospora (Cavalc. & Silva) B. Sutton, The Coelomycetes: 24 (1980).
Alysia pithospora Cavalc. & Silva, in Cavalcante et al., Publcoes Inst. Micol. Recife 647 : 34 ( 1 972).

For description and illustrations see Cavalcante et al. (1972 : 36 fig. 7).

Host: Originally described as a lichenized coelomycete with a Phycopeltis phycobiont, but
almost certainly a lichenicolous species on an unknown foliicolous lichen.

Distribution: Brazil (Rondonia). Known only from the original locality.

Observations: Neither Sutton (loc. cit.} nor myself have seen material of this taxon but the
published figures leave little doubt that a species of Vouauxiella was involved. The pycnidia
and conidia were originally given as 70-1 60 /zm diam and 2-6 x l-5-2'5//m, respectively.
These measurements are not in accord with the scale on the figures which suggest the
pycnidia are to about 60 //m wide and the conidia 3-3*5 x 2'5 /im; these latter figures were
used by Sutton. It seems impossible to resolve this discrepancy in the absence of the type
material. If the scale on the figure is correct, the species is clearly separated from the others
treated here by the small conidia; if the text is correct it will merit a critical comparison with
V. lichenicola.

3. Vouauxiella uniseptata D. Hawksw., Notes R. hot. Gdn Edinb. 36 : 195 (1978).

For description and illustrations see Hawksworth (1978^:195-196), and Sutton
(1980:24-25).

Host: Parmelia laevigata (Sm.) Ach., thallus. Forming distinctive raised pustular black
patches, mainly 1-2 mm diam, on the lobes.

Distribution: British Isles (Scotland). Three collections were reported by Hawksworth (loc.
cit.} and four others have since been seen (see below). Not yet seen by me on this host in
Devon despite careful searches.

Observations: This species may well eventually require transfer to an independent genus
for the reasons indicated above (p. 65).

Specimens (additional to those listed by Hawksworth, loc. cit.): British Isles: Scotland, Argyll, S side
Loch Sunart, 14 August 1972, S. R. Davey (IMI 250032a!), Benderloch, grounds of Lochnell House, 4
August 1980, B. J. Coppins 8052 (E, IMI 251027!), N side Loch Etive, 8 km E of Connel Burn, 7
August 1980, B. J. Coppins 8083 (E, IMI 251025!); Kirkudbright, Kintyre, Tarbert, Wilhashemraig, on
Alnus, 29 July 1973,7. E. Menlove(lM\ 228667!).

LICHENICOLOUS COELOMYCETES 67

4. Vouauxiella verrucosa (Vouaux) Petrak & H. Sydow, Beih. Repert. nov. Spec. Regni veg
42:483(1927).

For illustrations of this species and discussion of its separation from V. lichenicola see
Hawksworth (1976 : 58). Further illustrations are included in Sutton (1980 : 25).

Host: Lecanora chlarotera Nyl. and L. laevis Poelt, apothecia and rarely thallus. The
pycnidia have a similar habit to those of Vouauxiella lichenicola as described above and
microscopic examination is therefore needed to separate these two species.

Distribution: British Isles (England, Ireland, Scotland), France and Pakistan. Perhaps often
overlooked as V. lichenicola and perhaps much under-recorded, at least in Europe.

Observations: Lindsay (18690) included material of this taxon within his concept of V.
lichenicola but, in accordance with later usage, a smooth-spored specimen was selected as
lectotype for his epithet (see above).

Specimens (additional to those listed by Hawksworth, loc. cit.): British Isles: England, Wiltshire,
Imber, on Lecanora chlarotera, 1979, A. M. Burnet (IMI 237278!). Ireland, Co. Cork, near Cork, on L.
chlarotera, March 1858, /. Carroll (E!). Pakistan: Khanaspur, on L. laevis, 25 April 1975, S. Ahmad
1059 (IMI 199857c!).

XX. VOUAUXIOMYCES Dyko & D. Hawksw.
in Hawksworth & Dyko, Lichenologist 11 : 57 (1979).

For a description and discussion of the affinities of this genus see Hawksworth & Dyko
(1979).

Type species: Vouauxiomyces truncatus (B. de Lesd.) Dyko & D. Hawksw.

Teleomorph (perfect state): Abrothallus de Not. In the course of investigations into the
anamorphs of lichenicolous ascomycetes it became clear that those of Abrothallus were
referrable to Vouauxiomyces. All connections are based on strong circumstantial evidence,
the concurrence of ascomata and conidiomata, and have not been proved by ascospore
cultures.

Number of species: Originally described as monotypic, but two additional species are
added here. These three anamorphs all regularly occur without the teleomorph and it is
consequently useful to have binomials available for these phases. The genus Abrothallus has
been studied intensively by Santesson but his results still await publication; according to
Santesson (in Nordin, 1964 : 226) the genus comprises about 15 species and it is probable
that others have anamorphs in Vouauxiomyces also. Published accounts of anamorphs in
four species of Abrothallus in addition to the three treated here are noted on p. 72.

Key to the species

1 Conidia exceeding 6 /um in length .............

- Conidia 3-5(-6) x 2-3-5(-4) /an; on Ramalina species . . Vouauxiomyces ramalinae (p. 67)

2(1 ) Conidia 6' 5-8 x 4-5- 5 //m; on Parmelia caperata and P. perlata

Vouauxiomyces truncatus (p. 70)

- Conidia (7-)7'5-10'5(-ll'5)x(5-)5'5-7(-7'5)/im; on Parmelia saxatilis and Platismatia

........... Vouauxiomyces santessonii (p. 69)

1. Vouauxiomyces ramalinae (Nordin) D. Hawksw. comb. nov.
(Fig. 33A-B)

Phoma ramalinae Nordin, Svensk hot. Tidskr. 58 : 227 (1964).
Type: Sweden, Uppland, Malsta par., on Ramalina fraxinea (L.) Ach. (apothecia), 23 May 1

Nordin 563a (UPS holotype n.v.).
Icones: Nordin, Svensk hot. Tidskr. 58 : pi. I figs 1-4, pi. II figs. 5 and 7 (1964).

68

D. L. HAWKSWORTH

B

10/im

Fig. 33A-B, Vouauxiomyces ramalinae (IMI 194098); A, conidiogenous cells; B, conidia.
C-D, V. santessonii (UPS holotype); C, conidiogenous cells; D, conidia. E, V. truncatus
(Savoie, Santesson, UPS), conidia.

Conidiomata pycnidial, immersed at first but becoming erumpent, to about half exposed
at maturity, arising singly, scattered or sometimes in a ring in young infections, black,
ostiolate, 55-75 /im diam; pycnidial wall mainly 3-5 cells thick, thickest in the exposed
part adjacent to the ostiole, 5-1 2 //m thick, dark brown, pseudoparenchymatous, textura
angularis, cells thick-walled, tending to be polyhedral, mainly 3-5 jum diam.
Conidiogenous cells holoblastic, ampulliform to lageniform, lining the pycnidial cavity,

LICHENICOLOUS COELOMYCETES 69

percurrently proliferating, to 3 annellations seen, hyaline, smooth-walled (4-)6-10x
3-5 um. Conidia arising singly, obpyriform, hyaline, collecting in a mucilaginous mass in
the pycnidial cavity and extruded as a drop, simple, irregularly guttulate, apex rounded, the
base abruptly truncate, smooth-walled by light microscopy but indications of a sparse
perhaps echinulate ornamation are sometimes visible by interference contrast, 3-5(-6)x
2-3-5H)//m.

Teleomorph (perfect state): Abrothallus suecicus (Kirschst.) Nordin. The connection is
based on the repeated occurrence of the ascomata in assocation with the anamorph.

Hosts: Ramalina calicaris (L.) Fr., R. dilacerata (Hoffim.) Hoffm., R.fastigiata (Pers.) Ach.
and R. fraxinea (L.) Ach., thallus and especially apothecia. These hosts are taken from
Nordin (1964 : 228-229) who also indicated that the species might well be present on R.
sinensis Jatta. The symptoms and course of infection were studied carefully by Nordin, who
found the pycnidia to start at one point near the margin of an apothecial disc and then extend
over the disc forming pycnidia concentrically, the discs become greyish and the surface
layers extremely brittle.

Distribution: British Isles (England, Ireland, Scotland), France, Finland, Norway and
Sweden. Nordin (1964) reported Abrothallus suecicus from 104 localities, 50 of these as the
anamorph alone. Evidently widespread where the host lichens are well-developed.

Observations: Vouauxiomyces ramalinae is distinguished from the other species of the
genus accepted here by the smaller pycnidia, the shorter and narrower conidia, and in being
associated with a different teleomorph. As the pycnidia are not uncommonly found without
the ascomata, it is clearly desirable that the anamorph should have its own binomial;
Nordin's epithet is consequently recombined into Vouauxiomyces here. The anamorph
cannot be accommodated in Phoma, as currently circumscribed, because the conidiogenous
cells are holoblastic and annellate and the conidia are not regularly bacilliform, ellipsoid or
subglobose.

Specimens (see Nordin, 1964:229-230 for numerous other specimens he studied): British Isles:
England, S. Devon, Slapton, Duck Marsh, on Ramalina fastigiata on Salix, 21 May 1975, D. L.
Hawkswonh 4137 (IMI 194098!). Ireland, Co. Galway, Connemara, Kilkieran, Flannery Bridge, on R.
fastigiata, 1 August 1966, G. Morgan-Jones (IMI 1214426!).

2. Vouauxiomyces santessonii D. Hawksw. sp. nov.
(Fig. 33C-D)

Similis Vouauxiomvcei truncato (B. de Lesd.) Dyko & D. Hawksw. sed differt in conidiis latioribus,
(7-)7-5-10-5(-ll-5)x"(5-)5-5-7(-7-5)/n.

Typus: Suecica, Uppland, Alsike s:n, Lunsen, vid akertegen SV om Hjortronmossen, pa grenar pa
marken i granskog, in thallo Platismatiae glaucae (L.) Culb. & C. Culb., 23. ix. 1947, R. Santesson
(UPS holotypus!).

Icones: Kotte, Zentralbl. Bakt. ParasitKde II, 24 : pi. 3 fig. 32 (1909). Lindsay, Q. Jl microsc. Sci.
II, 7 : pi. 5 figs. 5-7 (1857). Schaechtelin & Werner, Bull, trimest. Soc. mycol. Fr. 42 : pi. 3 figs.
11-12(1926).

Conidiomata pycnidial, immersed at first but becoming erumpent through the surface of
the host, about \ exposed at maturity, arising singly, scattered, sometimes arising on the site
of a previous pycnidium and then assuming a rimmed appearance from above and a
double-walled structure in vertical section, black, ostiolate, the ostiole often conspicuous
and c. 25 um diam, 125-225 urn diam; wall of varying numbers of cell layers, mainly 4-8
15-25//m thick, thickest around the ostiole in the erumpent part of the pycnidium, dark
brown, pseudoparenchymatous, textura angularis, cells thick- walled, tending to be poly-
hedral, mainly 3^4 um diam. Conidiogenous cells holoblastic, ampulliform to lageniform,
lining the pycnidial cavity, percurrently proliferating, annellate with to 4 annellations seen,
hyaline, smooth-walled to weakly echinulate, 8-12 x 3-4 //m. Conidia obpyriform, hyaline,

70 D. L. HAWKSWORTH

collecting in a mucilaginous mass in the pycnidial cavity and extruded as a drop, simple,
sometimes 1-3 guttulate, apex rounded, the base abruptly truncated, thin-walled, smooth-
walled at lower magnifications but with a widely spaced echinulate ornamentation just
discernible by interference contrast, (?-)? 5-1 0'5(-1 1 *5) x (5-)5'5-7(-7'5) /zm.

Teleomorph (perfect state): Abrothallus parmeliarum (Sommerf.) Arnold. The conidia of
this species were well-described in some of the earliest papers on lichenicolous fungi, notably
those of Tulasne (1852 : 1 14*) and Lindsay (1857*), and the connection has been accepted
by subsequent students of Abrothallus (e.g. Kotte, 1909 : 87; Schaechtelin & Werner,
1926:238-239).

Hosts: Parmelia saxatilis (L.) Ach. and Platismatia glauca (L.) Culb. & C. Culb., thalli.
The teleomorph is reported from a large number of species of Parmelia, many of which must
also support the anamorph. In early infections the pycnidia arise on almost healthy lobes,
with a circle of brownish discoloured tissue around the pycnidia (rarely over 25 um thick).
In more advanced stages of infection the pycnidia occur on gall-like deformations of the host
thalli typical of infections due to Abrothallus parmeliarum.

Distribution: I have studied material of the anamorph from the British Isles and Sweden,
but it can be expected to occur throughout the range of the teleomorph. The distribution of
the teleomorph is difficult to ascertain from the literature due to taxonomic uncertainties but
Abrothallus parmeliarum s. lat., in addition to being widespread in Europe, is mentioned
from at least Canada, Chile, Bolivia, Ecuador, Hawaii, Tenerife, Kenya, Tasmania,
Morocco, New Zealand, South Africa, Mongolia, the U.S.A., Siberia and Venezuela (Hertel,
1971 : 228-229).

Observations: This species differs from Vouauxiomyces truncatus in the broader, and
often also longer, conidia; the taxonomic importance of this difference is substantiated by the
different teleomorphs and host ranges.

The species is named in honour of Prof. Dr R. Santesson in recognition of his important,
yet unpublished, studies on Abrothallus, Pyrenotrichum, and other lichenicolous fungi.

The name Epicoccum parmeliarum H. Olivier may provide an earlier epithet for this
species but is not taken up here in view of some uncertainties surrounding its application (see
p. 76).

Additional specimens: British Isles: Scotland, Argyll, 4 km E of Dalmally, wood on N side of River
Lochy, on Parmelia saxatilis on Quercus, \ August 1980, P. W. James & B. J. Coppins 4961 (E, IMI
251031!). Wales, Montgomery, Gregynog Hall, on P. saxatilis on Fagus sylvatica, 25 May 1979, K. L.
Brady (IMI 239067!). Sweden (all on Platismatia glauca): Uppland, Vange s:n, Fiby urskog (S om
Fibysjon), i granskogen O om Getryggen, pa grankvistar, 28 April 1948, R. Santesson 4859 (UPS!);
Viksta s:n, Djupdalen, c. % km V om Nyboda, pa gran i tat barrskog, 21 May 1947, R. Santesson
(UPS!); Rasbo parish, c. 1 km SSE of the lake Rokarn, on twigs of Picea abies in a rather dark forest, 23
October 1955, R. Santesson 11139b (UPS!); Skuttunge, Norra myren, 3 May 1939, R. Sernander
(UPS!); Uppsala, Vange socken, Fiby urskog, 15 November 1945, B. H. Svenonius (UPS!).
Vastergotland, Halleberg, Vanersnas s:n, Hallesnipan, pa ek, 1 5 June 1944, R. Santesson (UPS!).

3. Vouauxiomyces truncatus (B. de Lesd.) Dyko & D. Hawksw., in Hawksworth & Dyko,

Lichenologist 11 : 58 (1979).
(Fig. 33E, 34)

For description, illustrations and synonymy see Hawksworth & Dyko ( 1979).

Teleomorph (perfect state): Abrothallus microspermus Tul. The anamorph of this species
was illustrated and described by Tulasne (1852:116, pi. 16 figs. 22-26) in his
original account of the fungus, giving the conidia as 6'5 x 3'5 um. Hepp (Flecht. Eur. no.

These reports were given under the name Abrothallus smithii Tul., a superfluous name for A. bertianus de Not.
(which has smaller conidia; see p. 72) misapplied by these authors.

LICHENICOLOUS COELOMYCETES

Fig. 34 Vouauxiomyces truncatus (Savoie, Santesson, UPS), conidia (x 2500).

471, 1857; K!) later distributed and illustrated material of both states. Pycnidia commonly
occur without the teleomorph but are regularly encountered mixed amongst the ascomata.
Abrothallus microspermus differs from A. parmeliarum, with which it has often been
confused, in the consistently epruinose ascomata which only reach 300 um diam; according
to Keissler ( 1930) the ascospores are also paler brown and the epithecium K-.

Hosts: Parmelia caperata (L.) Ach., and more rarely P. perlata (Huds.) Ach., thalli.
Little damage is usually encountered and the fungus is probably parasymbiotic.

Distribution: Austria, the British Isles (England, Ireland, Scotland, Wales), France,
Germany and Switzerland. The report of the occurrence of this species in Sweden
(Santesson, 1949 : 142) is based on material now referred to Vouauxiomyces santessonii.

Specimens (additional to those listed by Hawksworth & Dyko, 1979): British Isles: England,
S. Devon, Slapton Ley Nature Reserve, The Causeway, on Parmelia perlata on Salix, 30 March
1972, D. L. Hawksworth 2696 p.p. (IMI 164972 p.p.l, UPS!); he. cit., Peasdish, on P. caperata
on Ulmus, 23 August 1980, D. L. Hawksworth 5043 (IMI 251260!); Dorset, Wareham, on P.
caperata, 1978, V. Giavarini (El). Scotland, Dunbarton, Loch Lomond, Aber Island, on P.
perlata, 10 October 1979, B. J. Coppins 4482 (E, IMI 243340!). Wales, Merioneth, Barmouth,
nr Arthog, Fegla Fawr, on P. caperata. 12 December 1972, B. J. Coppins (E!). France:
Savoie, Dept. Haute-Savoie, N of Sciez, Chateau Coudree, on P. caperata on Tilia, 9 June
1946, R. Santesson (UPS!). Switzerland: sine loc., P. Hepp [Flecht. Eur. no. 471] (K!).

Additional reports of anamorphs

In addition to the anamorphs of Abrothallus and Microcalicium species treated in the
preceding section, which can occur without their teleomorphs, there are literature
reports of pycnidial anamorphs in a number of other lichenicolous ascomycetes. As
many of these may be merely spermatial, the conidia not functioning as diaspores but
rather as gametes, and consequently are normally to be found in association with

72 D. L. HAWKSWORTH

ascomata, it seems superfluous to provide them with independent binomials. No
compilation of the lichenicolous ascomycetes in which such anamorphs have been
reported appears to be available. As these states may cause confusion when studying
lichenicolous Coelomycetes, or sometimes be found alone, reports of those I have
been able to discover are listed here. The location of all such reports would be a major
bibliographical exercise involving the study of the whole literature on lichenicolous
fungi; the following should consequently not be viewed as an exhaustive list.

Abrothallus bertianus de Not. (syn. A. glabratulae Kotte, A. smithii Tul.)

Kotte (1909:81) gave the conidia in this species as 5'46-6'5 x3'9-4-42/zm, but I
have not found a pycnidial state in the few specimens of it I have seen. This fungus
appears to be confined to brown species of Parmelia and is especially frequent on P.
glabratula (Lamy) Nyl. (Santesson, 1960; Hawksworth & Minter, 1980).

Abrothallus cetrariae Kotte

In the original account of this species, described from Platismatia glauca, the
conidia were given as 4-94-6-5 x 3 - 9-4'9 //m (Kotte, 1909:84). These measurements
are too small for Vouauxiomyces santessonii (p. 69), which occurs on the same host.

Abrothallus peyritischii (B. Stein) Kotte

The pycnidia of this species, which occurs on Cetraria nivalis (L.) Ach., were
illustrated by Kotte (1909 : pi. 1 figs. 12-17) and appear to belong to Vouauxiomyces;
the conidia were said to measure 5'2-6'76 x 3'9-5'2 um. One of Kotte's figures is
reproduced by Keissler(1930 : 205 fig. 46).

Abrothallus usneae Rabenh.

This fungus, which has often been united with A. parmeliarum, was investigated
anatomically by Gallee (1950:73, pis. 113-115). His drawings of vertical sections of
pycnidia, conidiogenous cells and conidia leave no doubt that the pycnidia of this
species belong to Vouauxiomyces, but no conidial dimensions were provided.

Arthonia clemens (Tul.) Th. Fr. (syn. Conidia clemens (Tul.) Massal., Phacopsis clemens

Tul.)

In the original description of this species, based on material from species of
Lecanora and Rhizoplaca chrysoleuca, Tulasne (1852:125) described the pycnidia
as 70-80 //m tall, thick- and brown-walled, with straight, bacilliform conidia 3-4 //m
long. They were also reported from this fungus in the hymenium of Lecanora atra
(Huds.) Ach. by Werner & Deschatres (1974 : 3 1 7-8) who gave them as 4 x 0'5 um.

Arthonia fuscopurpurea (Tul.) R. Sant. (syn. Celidiumfuscopurpureum Tul.)

This species, which occurs on Peltigera thalli, is described as forming bacilliform
conidia which are hyaline and to 3'5 /^m long (Tulasne, 1852 : 121).

Arthonia varia (Tul.) Jatta (syn. Phacopsis varia Tul.)

Tulasne (1852 : 126, pi. 14 fig. 3) described straight bacilliform conidia about
4-5 um long in this species on Xanthoria parietina (L.) Th. Fr. The relationship of
this taxon to both A. epiphyscia Tul. and A. destruens Rehm, which also attack
Xanthoria species, requires further study; contrary to Keissler's (1930 : 98) treatment, it is
distinct from A. glaucomaria (Nyl.) Nyl. which is found on Lecanora rupicola (L.) Zahlbr.

'Celidium' insidens Vouaux

The pycnidia of this fungus were rather fully described by Vouaux (1914:170).
They arose in groups of 5-6 and were lenticular-convex, immersed only at the base,
mm diam, black, and produced cuneiform hyaline 1 -septate conidia 9-12 x4 g 5 jum

LICHENICOLOUS COELOMYCETES 73

from simple to 2-septate conidiogenous cells measuring 9-20 x 2-3 //m. The species was
found on material ofRamalinafraxinea collected in Tenerife.

Dacampiosphaeria rivana (de Not.) D. Hawksw. (syn. Homostegia lichenum Fuckel)

According to Fuckel (1870:224) this fungus produces 1 -septate, cylindrical, curved
conidia which are 16x2-5/^m. In my investigations on this fungus I did not
find any pycnidial state (Hawksworth, 19800:367-371), and it seems probable that
Fuckel's report is based on an unidentified lichenicolous coelomycete on Peltigera.

Dichosporium glomerata Pat.

This species, which occurred on an undetermined corticolous lichen from Guadaloupe, is
described as producing pycnidia on a superficial mycelium which give rise to hyaline,
simple, subcylindrical conidia, measuring 5-6 x 2 //m (Vouaux, 1912 : 199).

Didymella parvispora Henssen

Henssen (1977:44, pi. SK), in the original description of this fungus from
Zahlbrucknerella africana Henssen, reported the pycnidia as partially immersed,
elongate, 30-50 /zm broad, with a dark brown ostiole, and elongate conidiogenous cells
giving rise to bacilliform, hyaline conidia 1*5-2 x O'5-l //m.

Guignardia olivieri (Vouaux) Sacc.

Pycnidia resembling those of Leptodothiorella, scattered in thalline warts, immersed,
globose, 75-100 //m diam; pycnidial wall subhyaline, pseudoparenchymatous, 8-12 //m
thick; conidiogenous cells phialidic, short-ampulliform, 7-10 x l'5-2-5//m; conidia arising
in short chains, bacilliform, hyaline, simple, often slightly compressed medianly, 3'5-5 x
1'5-2/zm. The anamorph of this species was described and illustrated by Hawksworth
(19756: 190-191, fig, 5o-j) on Xanthoria parietina. The conidiogenous cells were then
described as holoblastic, but improved microscopic techniques indicate that they are almost
certainly enteroblastic-phialidic.

Homostegia piggotii (Berk. & Broome) P. Karsten

This species was studied in some detail by Schaechtelin & Werner (1928) who
noted pycnidia in developing stromata on Parmelia saxatilis (L.) Ach. These were
pyriform, ostiolate, with slender conidiogenous cells radially orientated. Their illustration
(loc. cit.: pi. 1 1 fig. 17) suggests that the conidiogenous cells may sometimes be multicellular
and branched, but the hyaline, simple, subglobose conidia were reported to be formed
apically. No measurements were provided.

Lethariicola sipei Grumm.

In the original account of this monotypic genus of odontotremoid fungi on Letharia
vulpina (L.) Hue, Grummann (1969: pi. XIV figs. 19-20) provided photographs of
rather thick-walled erumpent pycnidia, but did not mention them in the written
account of the species. The nature of the conidiogenous cells and the conidia cannot
be deduced from the low-power photographs provided.

Lichenomyces lichenum (Sommerf. ex Fr.) R. Sant. (syn. Celidium stictarum Tul.)

Pycnidia have been noted in this species by several workers, notably Tulasne
(1852: 123, pi. 14 fig. 8), Werner (1928 : pi. IX figs. 2-4) and Vezda (1970:222, 225
fig. 4). The pycnidia are immersed to slightly erumpent at the ostiole in young
stromata, and contain filiform, simple conidiogenous cells to 30 //m tall, which in
turn produce bacilliform, hyaline, simple conidia 3^ x 1 //m singly from their apices. This
fungus is especially frequent on Lobaria pulmonaria (L.) Hoffm., and is known from a wide
range of genera in the Stictaceae.

74 D. L. HAWKSWORTH

Merismatium lecanorae (H. Olivier) Vouaux

Vouaux (1913:78) discovered material of this fungus, on Ochrolechia parella (L.)
Massal., with pycnidia which were black, 250-350 //m diam, and formed ovoid
conidia which were hyaline and simple at first, but later became brown and 3-septate
to muriform when mature, and 1 1-15 x 6-7 //m. He was not, however, convinced of
the relationship between the pycnidia and the putative teleomorph.

Muellerella lichenicola (Sommerf. ex Fr.) D. Hawksw. (syn. Tichothecium erraticum

Massal.)

Vouaux (1913:63) found pycnidia in a collection of Caloplaca vitellinula auct.
associated with the species now called Muellerella lichenicola. These gave rise to
brown, 1 -septate, ellipsoid conidia measuring 6-7 x 2'5-3 /im. Vouaux had little doubt that
he was dealing with an anamorph of M. lichenicola, but the details he provides show that his
material belonged to Lichenodiplis lecanorae (see p. 38).

Nesolechia oxyspora (Tul.) Massal.

Kotte (1909:93, pi. 3 figs. 29-30) noted pycnidia in the thecium of this species,
which normally occurs on thalli of Parmelia saxatilis. These produced rather irregularly
shaped broadly ellipsoid to subglobose hyaline conidia, but no further details were provided.
He may have been dealing with an unknown fungicolous coelomycete on the Nesolechia,
rather than an anamorph of N. oxyspora.

Scutula epiblastematica (Wallr.) Rehm (syn. S. wallrothii Tul.)

According to Tulasne (1852: 119-120, pi. 14 figs. 18-24) and Vouaux (1913:424),
this fungus, which occurs on Peltigera thalli, has two types of pycnidia. One, which
they considered to be spermagonial, had pycnidia 60-90 /zm diam with hyphal walls
and formed cylindrical, slightly curved, simple, hyaline conidia, 8-12 xO'5-1 /^m;
the second type of pycnidium was larger, 100-1 50 //m diam, with the walls composed
of thick-walled pseudoparenchymatous cells, and gave rise to narrowly ellipsoid,
often slightly curved, simple or rarely 1-septate, hyaline conidia, 12-16 x 2 - 5-4 ^m,
from conidiophores measuring 6-10 x 2 //m. There is a strong resemblance between
the reported macroconidial phase and the description of Diplodina peltigerae (see
p. 76). The pycnidial wall structure for this assumed state, as illustrated by Tulasne
(loc. cit.), recalls that of Karsteniomyces peltigerae (see .p. 22) and it is conceivable that he
had a mixed collection in which the conidia of the Karsteniomyces had not attained their
normal length and most not their final septum.

Stigmidium dispersum (Lahm ex Korber) D. Hawksw. (syn. Pharcidia dispersa (Lahm

ex Korber) Winter)

Pycnidia were apparently first reported for this species by Vezda (1963:151-153,
fig. 4), who described them as semi-immersed, 80-1 20 /zm diam, and forming bacilli-
form, simple or 1 -septate, hyaline conidia, 1 0-1 5x1-5

'Telimena' foreaui F. Moreau

In the original account of this species, described from specimens of Heterodermia
speciosa (Wulfen) Trevisan collected in India, Moreau (1951:209) reported that
pycnidia were present in the stroma. These were c. 50 //m diam, immersed, and
formed bacilliform, hyaline conidia, 4 x 1 //m.

Excluded species

All taxa referred to coelomycete genera which have been considered as lichenicolous
by their original or later authors, and which have not been treated in the preceding

LICHENICOLOUS COELOMYCETES 75

sections of this contribution, are compiled here alphabetically; obligate synonyms
are listed by their basionyms and not separately.

Aposphaeria ramalinae Vouaux, in Pitard & Harmand, Bull. Soc. hot. Fr., Mem.
22:72(1912)['19H'].

Type: Canary Islands, Tenerife, Vueltas de Taganana, 'sur le Ramalina implectens, dans
la zone sylvestre', alt. 700 m, 1 907, J. Pitard.

This fungus was originally described as forming a brownish superficial reticulum of
hyphae 1'5-3/zm wide, and from which elongate fusiform brownish l(-3) septate
conidia 9-12 x2-3'5 //m were formed in short chains. In the centre of the mycelium
pycnidia 80-1 30 /zm were produced which contained hyaline, simple, broadly ovate
conidia, measuring 3-3 '5 x2'5 //m. No material of this taxon could be located amongst
Harmand's herbarium in Angers (M. Guerlesquin, in litt.), but according to the original
description the name seems to have been based on two different fungi: an unidentifiable
dematiaceous hyphomycete and a coelomycete. The conidia of the latter recall those of
Vouauxiomyces ramalinae (see p. 67), but the pycnidia were larger than usual for that
species.

Chaetothyriolum puiggarii Speg., Boln Acad. nac. Cienc. Cordoba 23 : 522 (1919).

This monotypic genus was considered by Petrak & Sydow (1935 : 192) to be based
on the pycnidia of a lichen. The type material was re-examined by Santesson
(1952 : 53) and found to be too fragmentary for reliable determination.

Chlorocyphella aeruginascens var. cystidiifera Keissler, Annln naturh. Mus. Wien 41 : 162

(1952).

This fungus, described from material of Lopadium phyllogenum (Mull. Arg.) Zahlbr.
collected in the Solomon Islands, is almost certainly a species of Pyrenotrichum. According
to Keissler (loc. cit.), it differs from the species now known as P. splitgerberi in having sterile
cystidia-like structures between the conidiogenous cells, and much longer conidia,
120-140 x 1-5 //m. I have not examined the type material and the name was not discussed
by Santesson (1952 :41, 49-53).

Coniothyrium epiphyllum Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 296 (1914).

This name is of uncertain application, no fungus recalling the original description
being present on the holotype (Hawksworth, 1977 : 193).

Coniothyrium harmandii Zopfex Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 295 (1914).

This name is of uncertain application in the absence of type material (Hawksworth
1977: 193-194).

Coniscosolen mirabilis Schilling, Hedwigia 67 : 297 (1927).

This monotypic genus was based on the normal pycnidia of an unidentified
Psorotheciopsis Rehm species according to Santesson (1952 : 331) who examined the type
material.

Cristidium pallidum R. Sant., Symb. hot. upsal. 12(1) : 357 (1952); nom. inval. (Art. 32).

This name was mentioned as an undescribed genus and species of imperfect fungi -
on Gyalectidium filicinum Mull. Arg., but no further details were given and it is thus
uncertain if this was a coelomycete or hyphomycete. It is conceivable that this might
be Hansfordiellopsis lichenicola (Bat. & Maia) Deighton, which is well known on
Gyalectidium (Hawksworth, 1979 : 224-227).

Dendrophoma lecanorae Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 284 (1914).

This species was described on the basis of two different French collections, neither
present among the remnants of Vouaux's herbarium (Rondon, 1970); on Lecanora
piniperda Korber (syn. L. glaucella (Flotow) Nyl.) from Meyrueis, Lozere, and on

76 D. L. HAWKSWORTH

Lecania fuscella (Schaerer) Korber (syn. L. syringed (Ach.) Th. Fr.) from Malo-
Terminus. The erumpent pycnidia were described as 100-1 72 //m, with a hyphal
wall, 2-3-irregularly branched conidiophores 12-25x2//m, and ovoid hyaline simple
conidia 3-5 x 1-5-2 um. In the absence of material this name remains of uncertain appli-
cation; the conidiophore description and conidium shape indicates that Lecania conidia
were probably not involved (Eigler, 1969 : pi. 5).

Diplodia parmeliae Berk. & M. A. Curtis, in Berkeley, Grevillea 3 : 3 (1874).

Type: U.S.A., South Carolina, on Parmelia cf. rudecta Ach., M. A. Curtis (K holotype!).
Diplodina parmeliae (Berk. & M. A. Curtis) Sacc., Syll Fung. 3:413(1 884).

This species was introduced with an extremely brief description which included no
information on the sizes of spores or other features. The perithecia were reported to
have depressed ostioles and to be loosely clothed with flexuous threads, but their
colour was not indicated, and the spores stated to be hyaline with 1 septum and
unequal cells. Examination of the holotype established that the perithecia were translucent-
orange, about 200 /zm wide, with whitish lax hairs, and that ascospores in two size ranges
were formed, macroascospores about 51x23//m and microascospores 9-llx4-6//m.
The fungus agrees in all details with Nectria heterospora Speg., a species described and
discussed by Hawksworth & Booth (1976), but as the epithet heterospora dates from 1889,
the new combination Nectria parmeliae (Berk. & M. A. Curtis) D. Hawksw. comb. nov. must
be adopted for this species.

Diplodina peltigerae Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 289 (1914).

Type: France, Vosges, Plainfaing, on Peltigera canina (L.) Willd., J. Harmand & L.

Vouaux.

This species is not represented in the remnants of Vouaux's herbarium (Rondon,
1970) and it could not be located amongst Harmand's collections in Angers (M.
Guerlesquin, in lilt.). It was described as forming black, immersed pycnidia 60-1 00 //m
diam, which produced hyaline, ellipsoid conidia with a single septum often located below
the centre and 10-14 x 3-4 um. If it is re-collected it should be possible to recognize it from
these details and then re-assess its position. No similar fungus is known to occur on Peltigera
(Hawksworth, 1980a); the measurements are suggestive of the reported macroconidial
anamorph ofScutula epiblastematica (p. 74).

Epicoccum parmeliarum H. Olivier, Bull, internal, geogr. Bot. 17 : 232 (1907).

Type: France, Orne, on Parmelia caperata and P. sulcata (thalli), H. Olivier.
Phoma parmeliarum (H. Olivier) Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 196 (1914).

Vouaux (loc. cit.) considered that this name was based on the anamorph of Abrothallus
parmeliarum. There can be little doubt Olivier was dealing with a Vouauxiomyces species
since the conidia were originally described by him as 'terminees en pedoncule a la base'; the
pycnidial size quoted (0-2-0-4 mm diam) and the lengths and breadths of the conidia
(6-9 x 5-7 um) certainly recall the measurements for V. santessonii (teleomorph A.
parmeliarum), but the mention of Parmelia caperata suggests that he may really have been
dealing with either V. truncatus (teleomorph A. microspermus), or a mixture of both these
anamorphs on different hosts. As Olivier's epithet would pre-date both ''santessonii'' and
'truncatus' at species level it seems unwise to take it up here in either sense in the absence of
Olivier's material.

Libertiella obscurior Gerber, Arch. Protistenk. 74 : 485 (193 1).

Type: Austria, Hohen Tauren [Hohentauren], Naturschutzpark, on Hypogymnia intestiniformis
(Vill.) Rasanen (syn. H. encausta (Sm.) W. Watson), 1929, F. Mattick'.

The holotype was formerly preserved in the herbarium at DR but the lichen
material there was destroyed in 1945 and no duplicate appears to have existed (F.

LICHENICOLOUS COELOMYCETES

'?l ..^'

77

^'M. . r ' s . . fc . / ^*\ O.

Fig. 35 Lichenophoma haematommatis. A, Pycnidia on the host. B, Vertical section of
pycnidium. Scale not indicated. Reproduced from Keissler (1911 : 297).

Mattick, in litt.). This taxon was described as producing numerous immersed pycnidia,
O 1 1 5-0'2 mm diam, pale at the base but brownish above, delicate and hyphal, and producing
ellipsoid, hyaline conidia measuring 2'5-3'4 x 1-7-2 /zm. No conidiophores were recog-
nized. Pycnidia are often numerous in Hypogymnia intestiniformis but the conidia measure
5-6-5 xO'8-1 um (Hillmann, 1936 : 80). If Gerber's figures are reliable, there can conse-
quently be little doubt that his assertion was correct in that he was not dealing with the
normal pycnidia of the host. The name must, however, remain of uncertain application until
fresh material that might be referrable to Gerber's taxon is discovered, and the method of
conidiogenesis determined.

Libertiella xanthoriae Keissler, Annln naturh. Mus. Wien 38 : 166 (1925).

Type: Sweden, Vestmanland, Vesteras, on Xanthoria fallax (Hepp) Arnold [?], 2 August
1924,y. A. Nannfeldt(W 1925/10417 holotype!).

The Xanthoria thalli appear to be quite healthy and are their normal colour, except
for some deeper orange convex swellings (200-600 //m wide). In vertical section these
were seen to comprise a very irregular compact conidiiferous structure and to produce
numerous hyaline subcylindrical to rather ellipsoid conidia 3^xl-2//m which arose
laterally on integrated doliiform conidiogenous cells. This area was not sharply delimited
from the host tissues internally and the hyphae merged indistinguishably with the rest of the
thallus, which included healthy algal cells in the immediate vicinity of the pycnidia.

Pycnidia (spermogonia) are not common in X. fallax but were figured by Galloe
(1948: pi. 98 fig. 641) as deeper orange convex swellings superficially identical to
those on Nannfeldt's specimen. As the conidiogenous cells are similar to those of
other Xanthoria species (see e.g. Galloe, op. cit.) it is evident that Keissler's name
was based on the normal pycnidia of the host; the species of Xanthoria may not have
been X. fallax, however, as the conidia in that species are reported to be a rather
more elongate (Moberg, 1977 : 13).

Lichenophoma haematommatis Keissler, Hedwigia 50 : 296 (1911).

(Fig.35A-B)

Type: Austria, 'in valle flumius "Enns", dicto "Gesause", ad Gesause-Eingang', on Haematomma
elatinum (Ach.) Massal. thallus, July 19 10, A". Keissler.

The new genus Lichenophoma Keissler was introduced by Keissler (loc. cit.) for this
single species. Unfortunately the type material could not be located in W (U.
Passauer, in litt.) and, as noted by Sutton (1977: 113-114), the structure of the wall,
conidiogenesis, and relationship of the sterile paraphyses-like filaments figured (Fig.

78 D. L. HAWKSWORTH

35e) to the conidiophores all require clarification before any firm opinion on the
genus can be reached. The pycnidia were described as immersed, c. 120-150x60-
80 jam, with conidiogenous cells 12-15 x 1 //m (!) forming oblong conidia which
were hyaline and c. 5x2 //m; the sterile filaments, c. 60-70 x 1 //m, branched and
approached the ostiole. I have not seen any fungus on Haematomma elatinum that
could be identical with Keissler's taxon. There is a second report of this species by
Keissler (1933 : 390) from a sterile lichen thallus on Quercus in France (Oise, Foret
de Compiegne, au mont St Marc, November 1923, E. Cottereau} but that collection also has
not been located.

Lichenophoma opegraphae Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 285 (1914).

This taxon was based on two collections of Bouly de Lesdain from northern France:
on Opegrapha sp. at Merckeghem, and on O. niveoatra (Borrer) Laundon (syn. O.
subsiderella (Nyl.) Arnold) from Ghyvelde. The pycnidia were described as immersed,
100-1 50 //m diam, with pseudoparenchymatous brownish walls composed of polyhedral
cells 3-6 //m thick. Oblong conidia somewhat narrowed at the base, often unequal-sided,
simple, hyaline, and 3*5-5 x 1-5-2 /zm, were given as produced from simple conidiophores,
10-22 x 1*5 /urn. Sterile branched and anastomosing hyphae 2-2-5 ^m thick formed
'un reseau irregulier'. This taxon is not represented in the remnants of Vouaux's herbarium
(Rondon, 1970) and as that of Bouly de Lesdain was destroyed in 1940 the application of this
name must remain uncertain. It is, however, probable that Vouaux was dealing with the
microconidial state of the host which forms unequal-sided conidia mainly 4-6(-7) x
1-1 '5 /zm. It is most unlikely that he had the fungus treated here as Laeviomyces
opegraphae (p. 28) as that has much shorter conidiogenous cells, pale brown conidia with
equal sides and a distinctly truncated base, and no network of hyphae in the pycnidial cavity.

Microthelia solorinaria Lindsay, Q. Jlmicrosc. Sci. II, 9 : 350 (1869).

Type: Ireland, Co. Kerry, Brandon Mountain, on Solorina crocea (L.) Ach., /. Carroll

(E holotype!).
Diplodina solorinaria (Lindsay) Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 289 (1914).

In the original place of publication of this name, Lindsay (loc. cit.) stated merely
'M. solorinaria, Linds., Spermog., 175. On thallus of Solorina crocea. Pycnidia only.'
Although no description was provided here, as the earlier publication of Lindsay
(1859 : 175) is cited, and a description was given there, the name must be accepted as
validly published. Lindsay (1859 : 175) cited two specimens 'Brandon Mountain, Kerry,
Ireland, CARROLL; Wicklow, D. MOORE, in Herb. Carroll' but only one of these
(unspecified) had the fungus concerned. Both were thought to be lost (Hawksworth,
1980& : 174) but the former was recently re-discovered misfiled under Pleospora hookeri
(Borrer) Keissler in E; as this has a fungus conforming to Lindsay's description and drawings
it must be the holotype. This collection supports what is certainly Rhagadostoma lichenicola
(de Not.) Keissler; although no intact asci were seen the ascospores were typical for that
species and 'Munk-pores' were present on the peridial cells. Dacampiosphaeria rivana (de
Not.) D. Hawksw. is also present on the specimen but does not predominate nor conform to
Lindsay's account. The name M. solorinaria is consequently correctly interpreted as a later
synonym ofR. lichenicola.

Phaeoantenariella lichenicola Cavalcante, Publcdes Inst. Micol. Recife 633 : 3 (1969).

Type: Brazil, Maranhao, alto Turi, on Mazosia melanophthalma (Mull. Arg.) R. Sant. on
indet. leaves, 27 December 1967,7. de Anchieta (URM 80709 n.v.).

The monotypic new genus Phaeoantenariella Cavalcante was introduced for a
fungus with broad, torulose, dark brown hyphae spreading over the surface of the
host, and which was reported to produce minute intercalary pycnidia, only 1-3 times
the width of the hyphae, and conidia which were brown, simple, and 2^4 x 1'5-2/zm. A

LICHENICOLOUS COELOMYCETES 79

specimen received from Recife and labelled 'tipo' from the type locality was examined but
this bore the date 8 April 1967 and was numbered URM 71963 (exs. 41664!), contrary to the
type description. This material had, however, been studied by Cavalcante in July 1968, prior
to the publication of the name, and supported hyphae certainly identical to those figured by
Cavalcante (loc. cit.: 4-5). The 'pycnidia' described appeared to be only compact dividing
regions of cells, and the 'conidia' must be assumed to be merely cells that had become
dissociated by pressure during slide preparation.

This name is consequently rejected as based on sterile mycelium.

Phoma abietinae Vouaux, Bull, trimest. Soc. mycol. Fr, 30 : 282 (1914).

Type: Switzerland, 'ad truncos abietinos in m. Gurnigel', L. E. Schaerer [Lich Exs no
534] (E holotype!).

Exsiccata: Schaerer, Lich. Exs. no. 534 (E!; sub Lecidea leucocephala C. patelliformis).

Vouaux (1914:282) based his account of Phoma abietinae on Lindsay's (1872:259)
description of pycnidia in the above exsiccatum; he did not indicate that he had studied any
examples for himself of this number. Pycnidia are now rather sparse in the holotype,
scattered over the thallus of Lecanactis abietina (Ach.) Korber, which has numerous
apothecia, and agree in all details with the pycnidial state of this lichen (see Coppins &
James, 1979); the C+ red reaction obtained from the pruina covering the aging pycnidia, the
shape of the pycnidia, and size of the conidia, all show that this is not L. subabietina Coppins
& P. James, a species with C pycnidial pruina not known to form apothecia. The name
Phoma abietinae should consequently be added to the synonymy of Lecanactis abietina, and
excluded from the lichenicolous fungi.

Phoma alectoriae Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 197 (19 14).

Type: Switzerland, 'ad Abietum truncos in m. Gurnigel', on Bryoria capillaris (Ach.)
Brodo & D. Hawksw., L. E. Schaerer [Lich. Exs. no. 496] (E lectotype!).

Exsiccata: Schaerer, Lich. Exs. no. 496 (E!; sub Parmelia jubata d. cana).
Icones: Lindsay, Trans. R. Soc. Edinb. 22 : pi. IV figs. 17-18 (1859).

Vouaux's (1914: 197) tentative introduction of this name for a species he considered allied
to Phoma usneae (see p. 57) was based entirely on Lindsay's (1859 : 134) description and
illustrations. The flattened blackened warts mentioned by Lindsay now occur rather sparsely
on the lectotype and in none could any pycnidia or conidia be found. These warts do, how-
ever, closely resemble the young ascocarps of Phacopsis huuskonenii Rasanan, which is
well-known on the same host (Hawksworth, 1978a). The 'sterigmata' figured recall the
paraphyses of P. huuskonenii, and furthermore the spore size given (15-23x3//m) is
reminiscent of that of this same species ((12-)14-16(-18) x2-3'5 um,fide Hawksworth, loc.
cit.), although the conidia in Lindsay's figure do not belong here. Lindsay's figure
was, however, also based on Schaerer's Lich. Exs. no. 392 (sub Parmelia mollis y.
divaricata; E!), but this packet no longer contains any intermixed Bryoria.

This name, which in any case might be interpreted as not validly published under
Art. 34, is considered to have been based on Phacopsis huuskonenii and some other
fungus. As neither of these elements constitutes a satisfactory type, Vouaux's name
must be rejected.

Phoma arachnoidea Gerber, Arch. Protistenk. 74 : 483 (193 1).

Type: Austria, Solden, on Hypogymnia intestiniformis (Vill.) Rasanen (syn. H. encausta
(Sm.) W. Watson), W. Zopf.

No material under this name could be located in B (B. Hein, in litt.) or DR (see
p. 88). The pycnidia were described as about half-immersed and producing ellipsoid,
hyaline, and often pointed conidia 3'5^'25 x 1 -75-2-5 urn, formed apically on filiform,
non-septate, smooth conidiophores, measuring 12x2'5//m. If Gerber's measurements are

80 D. L. HAWKSWORTH

correct and his description of the conidiophores accurate, he must have been dealing with a
lichenicolous fungus and not the normal pycnidia of the host (see p. 77). Gerber's name
consequently remains of uncertain application, but the long conidiophores certainly exclude
it from Phoma.

Phoma biformis Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 282 (1914).

Type: France, Moeres, on Anisomeridium biforme (Borrer) R. C. Harris thallus on Salix
alba wood, M. Bouly de Lesdain.

No material of this taxon is present amongst the remnants of Vouaux's herbarium
(Rondon, 1970), and that of Bouly de Lesdain was destroyed in 1940. The pycnidia
were described as 50-1 40 //m wide, and as producing ovoid to globose conidia
1-1-75 x 1-1-5 um on conidiogenous cells 5-\2 um long. Normal pycnidia of this
lichen are not uncommon but may be of two types: microconidia which are orbicular
and 2-3 um, or macroconidia which are orbicular to elliptical or ovate and 3-5 um
diam or 3*5-5 x2'5-3 um (Harris, 1975:104). Vouaux's measurements suggest that
he was not therefore dealing with the pycnidia of the reported host, but it is quite
possible mistakes were made in measuring such small conidia. The name remains of
uncertain application.

Phoma caperatae Vouaux, in Bouly de Lesdain, Bull. Soc. hot. Fr. 59 : 16(1912).

Type: France, Versailles, Fausses-Reposes wood, on Parmelia caperata (L.) Ach. on oak,

M. Bouly de Lesdain.
Phoma physciicola var. caperatae (Vouaux) Keissler, Beih. Bot. Zbl. II. 37 : 273 (1920).

This species is not represented in the remnants of Vouaux's herbarium (Rondon,
1970), and that of Bouly de Lesdain was destroyed in 1940. The description of the
pycnidia as superfical, black, subglobose and 0- 1-0-25 mm diam is suggestive of
Vouauxiomyces truncatus (see p. 70), but the ovoid conidia were described as 'basi
saepe angustatis atque acutiusculis' most inappropriate for that species. The conidia were
given as 3-6 x 2'5-3 um, and the conidiogenous cells as 3-4 x 2 um. Bouly de Lesdain (loc.
cit.) mentions a second collection on the sterile thallus of a Calicium species on oak at
Rodez, Aveyron; these dimensions are suggestive of the normal conidia of Calicium rather
than any lichenicolous fungus known from this host. The collection on Parmelia at least
must have been a lichenicolous fungus, but its position remains uncertain in the absence of
the original material.

Phoma curvispora Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 28 1 (19 14).

In the original description of this species Vouaux cited three specimens, all collected by
Bouly de Lesdain: on Lecanora saligna (Schrader) Zahlbr. from Versailles, on Lecania sp.
from Zuydcoote, and on Lecidella elaeochroma (Ach.) M. Choisy from Ghyvelde. Unfortu-
nately Bouly de Lesdain's herbarium was destroyed in 1940, and this taxon is not represented
amongst the remnants of Vouaux's herbarium (Rondon, 1970). Keissler (1930 : 547) states
that he saw the 'Originalexamplar' himself, and that it represents only pycnidia of the lichen
in question but does not indicate which of the three original specimens he was able to see.
The description of the conidia as 'tres courbes, souvent en demi-cercle', and 8-1 5 x 2-3 um,
leaves little doubt that the name Phoma curvispora was partly based on the pycnidia of
Lecanora saligna.

Phoma epiparmelia Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 282 (1914).
Type: Yugoslavia, Nadin, Gradina, on Peltigera pusilla Korber, Baumgartner.

This fungus was described as not damaging the host thallus and comprising erumpent,
black, globose pycnidia 60-1 30 //m diam, with pseudoparenchymatous walls made up of
brown cells 7-10 x 4-6 um. Unbranched conidiophores, simple or rarely 1 -septate, sinuous,
and 6-10 x 1*5-2 um gave rise to rather irregularly shaped ellipsoid to frequently unequal-

LICHENICOLOUS COELOMYCETES 8 1

sided simple hyaline conidia measuring 3'5^'5 x 2-3 //m. Unfortunately the original
material could not be located in W (H. Riedl, in //.), and is not present amongst the
remnants of Vouaux's herbarium (Rondon, 1970). The long conidiogenous cells, smaller
pycnidia, and lack of damage to the host, separate this taxon from Phoma peltigerae (see
p. 54). In the absence of the original material the name must be treated as of uncertain
application until a fungus conforming to Vouaux's description is re-discovered.

Phoma epiphyscia Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 197 (19 14).

Type: France, Ghyvelde, on Phaeophyscia orbicularis (Necker) Moberg and Xanthoria
parietina (L.) Th. Fr., M. Boulyde Lesdain.

This taxon is not represented amongst the remnants of Vouaux's herbarium (Rondon,
1970), and as that of Bouly de Lesdain was destroyed in 1940, the type material is presumed
to be lost. The species was described as having immersed black pycnidia 80-1 50 //m, with
pseudoparenchymatous walls, and conidia hyaline, simple, 2-guttulate, 4-6 x 2-3 /zm,
formed from short conidiogenous cells. The conidial dimensions given are too large for
Phaeophyscia orbicularis and Xanthoria parietina. Vouaux (loc. cit.) suggested that it might
represent an anamorph of the species now called Arthonia epiphyscia Nyl.; I have not seen
pycnidia associated with A. epiphyscia, but they may occur (see p. 72). A further possibility
is that Vouaux was dealing with the anamorph of Guignardia olivieri (see p. 73), as
suggested by Hawksworth (19756: 191).

Phoma fusispora Vouaux, in Bouly de Lesdain, Bull. Soc. hot. Fr. 59:215 (1912);

nom. illegit. (Art. 64).
Non Phoma fusispora Rostrup, Bot. Faeroes 1 : 3 1 3 ( 1 90 1 ).

This species was originally described from specimens of Lecanora saligna (Schrader)
Zahlbr. and L. dispersa (Pers. ex Sommerf.) Nyl. collected by Bouly de Lesdain l aux environs
de Dunkerque'. It was also mentioned by Bouly de Lesdain (loc. cit.) as on Buellia punctata
(HofTm.) Massal. Vouaux (1914 : 196) added Bacidia spododes (Nyl.) Zahlbr.* as a host and
Bouly de Lesdain (1914 : 165) Opegrapha niveoatra (Borrer) Laundon. Bouly de Lesdain's
herbarium was destroyed in 1940, and this species is not represented in the remnants of
Vouaux's herbarium (Rondon, 1970). However, Keissler (1930 : 547) reports that he was
sent the 'Originalexemplar' by Bouly de Lesdain and that the name was based on the
pycnidia of the host, but does not indicate which host he received. The conidia were origin-
ally described as 'fusiformibus, paulum arcuatis, continuis, hyalinis' and 7-8 x 2 //m,
characters seen in numerous lichen pycnidia. In the absence of material some doubt must
remain, but the evidence suggests that this name was based on the pycnidia of several
different lichens.

Phoma glaucella Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 198 (19 14).

Three French collections were mentioned in the original account of this species: on
Bacidia inundata (Fr.) Korber at Malo-Terminus, on Lecanora sp. at Ghyvelde, and
on L. piniperda Korber (syn. L. glaucella (Flotow) Nyl.) at Meyrueis, Lozere; the first
two collections were made by Bouly de Lesdain and the last by Marc. The species was
described as having mainly immersed, black pycnidia 60-1 40 //m diam, simple or
rarely bifurcate conidiogenous cells 5-10 x 1-5-2 //m, and ellipsoid, hyaline conidia
l'5-3 x 1-1 -5 //m. As no material is present in the remnants of Vouaux's herbarium
(Rondon, 1970), and that of Bouly de Lesdain was destroyed in 1940, the application
of this name remains uncertain.

Phoma lecanorae Vouaux, in Bouly de Lesdain, Rech. Lich. Dunk.: 277 (19 10).

Type: France, Dunkerque, dunes pres le Phare, 'sur Lecanora umbrina var.\ 18 June
1904, M. Bouly de Lesdain (herb. Vouaux syntype!).

* = Micarea nitschkeana (Lahm ex Rabenh.) Harm.

82 D. L. HAWKSWORTH

Two specimens were mentioned in the original description of this taxon, the above
and one from Malo-Terminus reported as on Rinodina exigua (Ach.) Gray; this latter
collection is not represented in Vouaux's herbarium (Rondon, 1970), and as Bouly de
Lesdain's herbarium was also destroyed, must be presumed lost. The collection from
near Le Phare comprises a blackish-brown thallus on flint over which the pycnidia
are evenly scattered; fruits of Lecanora polytropa are scattered amongst the dark
thallus and were presumably mistaken for L. dispersa (Pers.) Sommerf. (syn. L.
umbrina Nyl.) by Vouaux, but appear to be quite unrelated to the pycnidia. The
evenly distributed pycnidia and healthy thallus leave no doubt that this material is
the pycnidial state of a lichenized species; the thallus colour and short conidia
(3-4 x 1-1-5 //m) are suggestive ofOpegrapha lithyrga Ach.

Bouly de Lesdain (1914) and Vouaux (1914) reported eleven further hosts from a
wide range of genera (interestingly, these included two Opegrapha species); these
cannot be said to have comprised only lichen pycnidia in the absence of the
collections cited.

Phoma lichenis Pass., Atti Accad. naz. Lincei Re. 7 : 48 ( 1 89 1 ).

Type: Italy, prov. Parma, VighefTio, Frassino, 'sul tallo di un Lichene sterile, fosse Parmelia
pulverulenta ' [Physconia pulverulacea Moberg], G. Passerini.

The type material could not be located in either PAD (S. Chiesa, in litt.} or
PARMA (F. Lona, in litt.). The black pycnidia were described as having mem-
branaceous pseudoparenchymatous walls and producing narrow bacilliform conidia
5 jum long. Conidia 4-7xl-5/zm are, however, characteristic of the pycnidia in both
Physcia (Schreber) Michaux and Physconia Poelt (Moberg, 1977), and it therefore
seems probable that this name was merely based on the pycnidia of the host. If this is
so, as Art. 59 does not apply to lichen-forming fungi, Passerini's name may provide
an earlier epithet for Physconia pulverulacea.

There are reports of this taxon from a wide range of hosts, some of which may refer
to true lichenicolous fungi. Keissler (1930:541-2) mentions alsp Cladonia rangi-
ferina (L.) Wigg., Lecanora pallida (Schreber) Rabenh., Physcia stellaris (L.) Nyl.,
and Xanthoria parietina (L.) Th. Fr.; Keissler (1933:386) adds Cladonia rangiformis
Hoffm. and Parmelia laevior Nyl.

Phoma lichenis f. immersa B. de Lesd., Rech. Lich. Dunk. : 277 (1910).

Type: France, Dunkirk, on Hypogymnia physodes (L.) Nyl., M. Bouly de Lesdain.

As Bouly de Lesdain's herbarium was destroyed in 1940, the type collection is
presumably lost. The only description provided was 'Les peritheces sont enfonces au
lieu d'etre superficial'. Assuming that the conidia were roughly the same size as those
given for Phoma lichenis, there can be little doubt that this form was based on normal
pycnidia of the host.

Phoma verrucariae Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 28 1 (19 14).
Type: France, Bergues, on Verrucaria sp. [?], M. Bouly de Lesdain.

This species was described as producing 2-3 pycnidia per areole of the sterile host,
the pycnidia as immersed to almost superfical, black, 60-1 10 um diam, with conidiophores
10-20 //m long, and conidia which were cylindrical, attenuated at the ends, slightly or
strongly curved, hyaline, simple, and 12-14 x 1-8-2-5 um. The host may not have been a
Verrucaria as it was sterile and so could not have been checked microscopically; the name
could therefore have been based on pycnidia of the 'host' and not a lichenicolous fungus. As
this taxon is not represented in the remnants of Vouaux's herbarium (Rondon, 1970), and
that of Bouly de Lesdain was destroyed in 1940, the application of this name remains
uncertain.

LICHENICOLOUS COELOMYCETES 83

Phoma versoniana Sacc., Michelia 2 : 342 (1 88 1 ); nom. inval. (Art. 32).

Keissler (1930: 582) lists the name as a possible synonym of the species now called
Libertiella malmedyensis, but it does not appear to have been ever validly published
and the only mention of it I have been able to locate is the reference cited above
where the phrase 'Phomae Versonianae meae analogo videtur' appears. It is not even
known whether P. versoniana was considered to be lichenicolous.

Phyllosticta galligena F. Moreau, Bull. Soc. hot. Fr. 98 : 102 (195 1 ).

Type: India, Palni Hills, Shembaganur, on Parmelia perforata Taylor, 1933, G. Foreau.
Icones: Moreau, Bull. Soc. hot. Fr. 98 : 1 1 ( 1 95 1 ).

The type material of this gall-forming species could not be located in PC (J.
Mouchacca, in litt.), nor was it available from Dr C. Moreau. Convex galls 0'25-1'5 mm
were formed each including several black pycnidia (to 150//m diam) forming simple,
ellipsoid, hyaline conidia 1 -5-3-5 x 1-2 um. The conidiophores were described as
unbranched and filiform, so it is clear that this species cannot be referred to Phoma (see
p. 49). In the absence of further information it is impossible to suggest an appropriate genus
for this species, but it should be possible to recognize it if re-collected and then re-assess its
position.

Phyllosticta lichenicola Allescher, Ber. bayer. hot. Ges. 4 : 32 (1896).

Type: Germany, Miinchen, Ebenhausen, on Parmelia perlata (Huds.) Ach., 1 1 April 1895, J. N.
Schnabl (M holotype!).

This fungus was originally described as forming in necrotic patches about 2 mm
diam, and having globose, black pycnidia with conidia hyaline, oblong, biguttulate,
and 2-4 x 0'5-1'5/zm. Keissler (1930:535) studied the type material superficially
but made no microscopic preparation because it was so small; no later authors appear
to have found any fungus matching Allescher's description exactly, so a careful study
of this specimen was needed. Only a single infection spot 2 mm diam remained on the
thallus, this was necrotic with a black margin about 1 25-200 //m wide and contained
six small black immersed sporocarps. Three of these proved to be ascomata of a
Polycoccum or Endococcus species (paraphyses doubtfully present and not clearly
demonstrable) with asci about 30 x 1 um and 1 -septate, pale brown, echinulate ascospores
about 8x3 um; the asci were 6-8 spored. A fourth was immature and without spores, and
the fifth a pycnidium with subglobose to broadly ellipsoid, hyaline conidia 3'5^ - 5 x
2-3 '5 //m; no conidiogenous cells could be found. The sixth sporocarp was left un-
examined for future investigators. Representative slides from the sporocarps examined are
preserved in IMI.

In view of these observations, it is clear that Allescher's taxon was based on
discordant elements, the two fungi found by me and a third, perhaps Phoma cytospora, with
conidia approximating to the shape and width he indicated. This name cannot be satis-
factorily typified by these disparate elements and must consequently be rejected.

Pleurosticta lichenicola Petrak, Kryptog. Forsch. 2(2) : 190 (193 1).

This new genus and species was introduced by Petrak (loc. cit.) for what he inter-
preted as a lichenicolous coelomycete growing on Parmelia olivacea (L.) Ach. in
Germany. The identity of this taxon has been investigated by Santesson (1949: 142)
who found it to be based on the normal pycnidia of a different lichen, P. acetabulum
(Necker) Duby. Unfortunately the type collection could not be re-located in W (H.
Riedl, in litt.), but there seems to be no reason to question Santesson 's interpretation.

Pleurosticta Petrak must consequently be treated as a further synonym of Parmelia
Ach. nom. cons. Further, as Art. 59 specifically excludes lichenized fungi, if the

84 D. L. HAWKSWORTH

group of brown Parmelia species for which Esslinger (1978) introduced the new genus
Melanelia Essl. are regarded as generically distinct, then Pleurosticta would be an
earlier name for that genus unless conserved, as proposed by Esslinger ( 1 980).

Pseudodiplodia lichenis Vouaux, in Bouly de Lesdain, Rech. Lich. Dunk., Suppl. : 167
(1914).
Type: France, Malo Terminus, on a telegraph pole, 28 August 1910, M. Bouly de Lesdain

(herb. Vouaux holotype!).
Stagonopsis lichenis (Vouaux) Keissler, Rabenh. Krypt.-Fl. 8 : 586 (1930).

This species was described as a fungus occurring on Lecanora saligna (Schrader)
Zahlbr. (syn. L. effusa (Hoffm.) Ach.), and has pale orange, cupuliform pycnidia
1 00-200 //m wide evenly distributed over a greyish-green crustose thallus; the hyaline
conidia are curved with rounded ends, 0-1 septate, and 8-10 x 2-4 jam. No Lecanora
apothecia were found on the specimen. The lack of damage to the host as well as the even
distribution of the pycnidia on the thallus leave no doubt that this name was based on the
pycnidial state of an unidentified lichenized fungus. The shape of the conidia recalls that
characteristic of the L. dispersa (Pers.) Sommerf. group within Lecanora, but the orange
pycnidia are atypical for that facies.

Vouaux's name was transferred to Stagonopsis by Keissler (1930:586) primarily
because the conidia could be 1 -septate.

Pyrenochaeta collematis Vouaux, in Bouly de Lesdain, Rech. Lich. Dunk. : 275 (1910).

Type: France, Dunkerque, Bray Dunes, on apothecia and thalli of Collema tenax (Swartz)
Ach., M. Bouly de Lesdain.

This fungus was originally described as forming black, almost superficial pycnidia
100-1 50 um diam, with 3-9 setae 40-60 x 3-5 um arranged around the ostiole, and
producing hyaline simple conidia 5-6 x 2 um on conidiogenous cells 6-1 x \-5um.
Vouaux (1912:185, 1914:288) suggested that it might represent an anamorph of
Nectria tenacis (Vouaux) Vouaux, but it would be remarkable for any Nectria to have
a coelomycete state; Pyrenochaeta de Not. species have teleomorphs in Herpotrichia
Fuckel. As Bouly de Lesdain's herbarium was destroyed in 1940, and this taxon is not
represented amongst the remnants of Vouaux's herbarium (Rondon, 1970), it has not
been possible to examine any material of this taxon. According to Vouaux's descrip-
tion this fungus cannot have been a Pyrenochaeta because the conidiogenous cells in
this genus are arranged in long acropleurogenous chains (Schneider, 1979).

Rhabdospora antarctica Speg., An. Mus. nac. B. Aires 20 : 390 (1910); as 'n.f.'

Type: Antarctic Islands, South Orkney Islands ['Nuevas Orcadas'], on Caloplaca regalis
(Vainio) Zahlbr., February 1908, C. Spegazzini (LPS 1 1231 holotype!).

The host, originally indicated to be a Teloschistes Norman species, proves to be the
ubiquitous antarctic polymorphic Caloplaca regalis (syn. Polycauliona regalis (Vainio)
Hue). In the original description of this fungus the 'perithecia' were given as black and
75-90 um diam, and the spores as hyaline, 1 -septate or simple, 'rotundato-subtruncatae'
and 12-16 x 2-3 um. The holotype is very small, the host fragment measuring only
3x2 mm, but two sporocarps recalling Spegazzini's description were present; one was
examined and found to be the pyrenomycete Polycoccum rugulosaria (Lindsay) D. Hawksw.,
which is confined to this host and described and illustrated elsewhere (Hawksworth in Pegler
et al, 1980). A few of the normal pycnidia of the Caloplaca were also found but they form
conidia quite unlike those described by Spegazzini (loc. cit.). It is consequently clear that
Rhabdospora antarctica should be treated as a synonym of P. rugulosaria. That Spegazzini
gave the spores as hyaline and sometimes simple, and also only 2-3 um wide,

LICHENICOLOUS COELOMYCETES 85

indicates that he based his description on some immature ascospores. Neither asci
nor conidiogenous structures were mentioned by Spegazzini.

The name Rhabdospora antarctica was evidently missed by Vouaux (1914); as a
result it seems to have been overlooked by subsequent students of the lichenicolous
fungi (e.g. Clauzade & Roux, 1976).

Rhabdospora lecanorae Vouaux, in Bouly de Lesdain, Reck. Lich. Dunk., Suppl. : 167
(1914).

Type: Spain, Catalogne, Hostaleto, nr Figueras, on Lecanora chlarona (Ach.) Nyl. on
olives, 1 1 April 1905, F. Heribaud (herb. Vouaux lectotype!).

In Bouly de Lesdain's publication three collections were cited, on Opegrapha vulgata
(Ach.) Ach. (syn. O. cinerea Chev.) from Dunkirk, on Micarea nitschkeana (syn. Bilimbia
spododes (Nyl.) Arnold) thalli from Bergues, and on Lecidella elaeochroma (Ach.) M. Choisy
(syn. Lecidea parasema auct.) from Ghyvelde. Vouaux's selection of the epithet 'lecanorae',
however, implied that he had a fourth collection in mind when providing his description for
Bouly de Lesdain, on Lecanora. Vouaux (1914 : 290) cites a collection on L. chlarona first,
before the Bouly de Lesdain material, and this must be regarded as a syntype for this name.
As only this latter collection is extant it is here designated as the lectotype.

The type is in a poor condition and is overgrown by superficial moulds; in some
apothecial margins acicular to arcuate or sigmoid hyaline conidia 20-30x1-1*5
(-2) um, produced on pyriform conidiogenous cells 4-6 x 3-5 /zm, were found. Identical
conidia were also produced in black pycnidia directly on the thallus, something overlooked
by Vouaux. These correspond to the pycnidia (spermogonia) usually produced by this
lichen, and the name Rhabdospora lecanorae should consequently be treated as a synonym
of Lecanora chlarona.

Rhabdospora lesdainii Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 290 (1914).

Type: France, Malo-Terminus, on Lecidella elaeochroma (Ach.) M. Choisy (syn. Lecidea
parasema auct.) on Populus monilifera, M. Bouly de Lesdain.

The type material is not amongst the remnants of Vouaux's herbarium (Rondon,
1970), and as that of Bouly de Lesdain was destroyed in 1940, must be presumed lost.
The fusiform, straight to slightly curved conidia were described as with 5-7 'plus ou
moins distinctes' septa and given as 38-60 x 3-4-5 um. The normal conidia of this
species recall Vouaux's description of their shape but are not septate (only guttulate)
and only \l-\8um long (Galloe, 1927:92). The name is consequently of uncertain
application.

Rhabdospora thallicola Tassi, Bull. Lab. Orto Bot. Reale Univ. Siena 3 : 129 (1900).
Type: Brazil, 'corteccie di Cinchona', September 1900, herb. F. Tassi (SIENA holotype!).

The type collection comprises two pieces of bark each covered with the creamy-
white thallus of a sterile crustaceous lichen. The pycnidia are scattered rather regularly over
the material and not associated with any signs of damage. The filiform conidia and arrange-
ment of the conidiogenous cells is similar to that seen in the pycnidia of numerous lichens,
particularly the Lecanora chlarotera Nyl. group. This name is consequently excluded as
based on the pycnidia of an unknown lichen.

Sphaeromma mazosiae Upadhyay, Publcoes Inst. Micol. Recife 402 : 4 (1964).

Type: Brazil, Rondonia, Porto Velho, Perfil AR-4, on Mazosia melanophthalma (Mull.
Arg.) R. Sant. on Orbigyna martiana, January 1962, O. M. Fonseca (URM 38672
[19.945 p.p.] holotype!).

86 D. L. HAWKSWORTH

The herbarium specimen URM 19.945 comprises twelve portions of leaf, mostly
about 10-12 cm long, which support a wide range of lichens and associated lichenicolous
fungi constituting 'Fungos do Brasil' nos. 38670-38682 (i.e. including the holotypes of both
Sphaeromma mazosiae and Sporhaplus rondoniensis Upadhyay). Sphaeromma mazosiae
and Sporhaplus rondoniensis were both described as having superficial, dark brown pycnidia
(33-45 um diam and 30-35 um diam, respectively) with stout, dark brown setae (given as
6-12-5 x 3*4 um for each species); the Sphaeromma was stated to have 2-4 septate conidia
7-10'5 x 1-5-2 //m, and the Sporhaplus simple conidia l'5-2 xO'75-1 um. Only after very
extensive searching did I manage to locate any Mazosia thalli with setose sporocarps
recalling those of the original description; several of these were squashed and found
to contain pseudoparenchymatous tissue, young asci, or asci with 1 -septate hyaline
immature spores about 8x4 um. It seems probable that the ascomycete is identical to
the monotypic Keratosphaera batistae Upadhyay, the holotype of which (URM 38673)
should be on this specimen, although its spores were indicated to become 3-septate when
mature and measure 6-10 x 2-2*5 um, and no evidence for the existence of paraphyses
figured by Upadhyay (he. cit. : 10) was obtained.

The host lichen was originally referred to Mazosia phyllosema (Nyl.) Zahlbr. by
Upadhyay, but the thallus is not smooth but has the minute whitish verrucae
characteristic of M. melanophthalma (Mull. Arg.) R. Sant., although they are rather
poorly developed in this material. According to Santesson (1952 : 1 18), M. melanophthalma
has macroconidia 5-8 x 2-3 um and microconidia 2-3 x 1 um; the correspondence be-
tween the sizes of these and those given for Sphaeromma mazosiae and Sporhaplus
rondoniensis is striking. In my opinion descriptions of both these fungi were probably based
on the conidia of M. melanophthalma (the 'septation' of the larger conidia being due to
guttulation) and sporocarps of the Keratosphaera. These two names cannot consequently be
satisfactorily typified and must be rejected.

Sphaeronema lichenophilum Durieu & Mont., in Montagne, Syll. gen. crypt. : 249 (1856).

Type: Algeria, 'sur les troncs du vieux Chamaeropis, du Marabout du sommet du Bouzareah', 8
April 1844, M. C. Durieu de Maisonneuve (PC holotype!).

This name has generally been cited as first published in Botanique I. Cryptogamiae
of Durieu's Exploration scientifique de I'Algerie, which is referred to with correct
page numbers in Montagne's book cited above. However, it is now clear that the part
of Durieu's work including Sphaeronema lichenophilum lay in proof for at least 13
and perhaps as many as 23 years before they were finally published in October 1869
(Hawksworth & Booth, 1974: 12); only the first parts of this volume were issued in
1846-50, the dates usually given for S. lichenophilum. Montagne cited only the single
collection detailed above, which is consequently the holotype; Durieu mentioned the
same collection, but also a further one made on Chamaeropis in February 1840
which is also now preserved in PC (!).

Durieu gave the host as 'Lecideae canescentis aut Dirinae Ceratoniae\ but in the
holotype, the 1840 specimen, and further two small undated specimens in PC (!), the
pycnidia are distributed over a whitish-grey, thin, crustose thallus forming circular
patches mainly 5-1 Omm diam. Eleven fragments of bark comprise the holotype
sheet, ten of which have pycnidia conforming to the original description, but the
other is Diploicia canescens (Dickson) Massal. unaffected by any pycnidia. In vertical
section the pycnidia were found to extend down through the thallus to the bark and this,
together with their even distribution and no evidence of damage to the thallus, strongly
suggest that they are the pycnidia of a lichenized fungus, and not an independent
coelomycete invading a lichen. The pycnidia are thick-walled, dark brown and
carbonaceous, perennial with new pycnidia arising within effete ones, phialides mainly
about 10 x 1-5 um, and numerous, minute, simple, cylindrical, guttulate conidia mainly

LICHENICOLOUS COELOMYCETES 87

5-6 x 2-3 urn. Its identity is uncertain, although the pycnidia suggest Arthoniales or
Hysteriales; it is not Dirina ceratoniae Fr., the pycnidia of which were illustrated by Vobis &
Hawksworth(1981).

Sphaeronema lichenophilum was also very tentatively reported from a Parmelia
collected in 1921 in Brazil (Sao Paulo, Premembe) by Keissler (1933 : 385), but as the
material was stated to be in Bouly de Lesdain's herbarium it was presumably
destroyed in 1940.

Sphaeropsis cladoniae Ell. & Ev., /. Mycol. 5 : 149 (1889).
Coniothyrium cladoniae (Ell. & Ev.) Sacc., Syll. Fung. 10 : 268 (1 892).

Lichenoconium cladoniae (Ell. & Ev.) Petrak & H. Sydow, Beih. Rep. nov. Spec. Regni veg.
42:433(1927).

This name was based on a collection including an Abrothallus species and Lichenoconium
usneae, but is considered correctly used for the former element (Hawksworth,
1977: 192-193).

Sphaeropsis scripta Schwein. ex Berk. & M. A. Curtis, J. Acad. nat. Sci. Philad.
2:280(1853).

Dothiorella erumpens Sacc., Syll. Fung. 3:242 (1884); as '(Berk. & C.)', nom. illegit. (Art.
63.2).

This fungus was listed as a lichenicolous species by Viegas (1961 : 869, as 'Dothiorella
erupens' [sic]) but Berkeley & Curtis (loc. cit., pi. 25 fig. 3a-c) illustrated a black, stromatic,
pulvinate, multilocular conidioma bursting through the bark from below, and were evidently
not dealing with a lichenicolous fungus; the fungus was erumpent through unidentified
lichens on the bark, not on the lichens themselves. The conidia of the fungus were broadly
ellipsoid, hyaline, simple, with a short persistent pedicel.

Spi lorn in m epicladonia H. Olivier, Paras. Lich. Fr., Suppl. : 22 (1907) [not seen].
Type: France, Herault, Mons-la-Frivola, on Cladonia sp., A. de Crozals.

No type or authentic material of this taxon has been located. Vouaux (1914: 198)
considered it might be identical to the species now called Bachmanniomyces uncialicola, but
this seems unlikely to be correct (see p. 14).

Sporhaplus rondoniensis Upadhyay, Pubic, oes Inst. Micol. Recife 402 : 7 (1964).

Type: Brazil, Rondonia, Porto Vehlo, Perfil AR-4, on Mazosia melanophthalma (Mull.
Arg.) R. Sant. on Orbigyna martiana, January 1962, O. M. Fonseca (URM 38674 [URM
19.945 p.p.] holotype!).

The holotype of this monotypic genus is present on the same collection as that of
Sphaeromma mazosiae and some other fungi, and is discussed under S. mazosiae
above (p. 86).

Sporocadus lichenicola Corda, Icones Fung. 3 : 24 (1839).

Type: Czechoslovakia, Prague, Fiirstl. Lobkowitzischen Garten, on Rosa sp., 1838, A. J.

C. Corda (PRM 1 55664 holotype n.v.).

Mender sonia lichenicola (Corda) Fr., Summ. veg. Scand. 2 : 416 (1849); as \Sommerf)\
Seimatosporium lichenicola (Corda) Shoem. & E. Muller, Can. J. Bot. 42 : 405 (1964).

This taxon, the lectotype species of the generic name Sporocadus Corda (Hughes,
1958:810), was thought by Corda to be associated with some unidentified crustose
lichen because algal cells were associated with the pycnidia of the fungus below the
surface of the host. The type material was examined by Hughes (loc. cit.) and his
slides were also studied by Shoemaker & Muller (1964); these, together with numerous later
collections identical to Corda's species, demonstrate conclusively that this is not a
lichenicolous fungus, but rather a common saprophyte of Rosa stems. The algae present in

88 D. L. HAWKSWORTH

Fig. 36 Verrucaster lichenicola. A, Infected podetium (x2). B, Gall with pycnidia (x 12).
C, Conidiophores (x 1000). D, Conidiogenous cells (x 1000). E, Conidia (x 1000). Reproduced
from Tobler (1913: 383).

Corda's collection occurred only fortuitously. Corda's fungus is correctly placed in
Seimatosporium and has an extensive synonymy (Sutton, 1975 : 136); it is the anamorph of
Clathridium corticola (Fuckel) Shoem. & E. Miiller (syn. Griphosphaeria corticola (Fuckel)
Hohnel) the connection between the two having been demonstrated by cultures of the
ascospores giving rise to the Seimatosporium state (Shoemaker & Miiller, 1964).

In accepting "Hendersonia lichenicola' as a lichenicolous species, Clauzade & Roux,
1976 : 89) were presumably simply following Keissler (1930 : 578) who listed it from several
lichen hosts; these must be presumed to be cases where the fungus grew through overlying
lichen thalli or otherwise they were misidentifications.

Sporocadus rosicola Rabenh., Bot. Ztg6 : 294 (1848); as "rosaecola\

Type: Germany, prope Leipzig, 'ad Rosarum ramulos', L. Rabenhorst [Fungi Exs. no. 66]
(not seen).

Rabenhorst's fungus occurred on Rosa branches and there can be little doubt that
its usual treatment as a synonym of Seimatosporium lichenicola (Corda) Shoem.
& E. Miiller is correct (Sutton, 1975:136). However, Saccardo (1884:439) stated
that it had been found '. . . in apotheciis Lecideae luteolae [i.e. Bacidia rubella
(Hoffrn.) Massal.], Gand in Belgio'. This later report must be treated as a misidentification
based on an unknown lichenicolous fungus.

Verrucaster lichenicola Tobler, Abh. naturw. Ver. Bremen 21 : 384 (1913) ['1912'].
(Fig. 36A-E)
Type: Germany, Oldenburg, Kehnmoor, on Cladonia bacillaris auct., H. Sandstede.

This new genus and species was introduced by Tobler (1913) for a fungus forming
gall-like growths on podetia of the host. The pycnidia were described as superficial,
and opening irregularly, waxy, forming simple, hyaline, narrowly ellipsoid 2-guttulate
conidia 3'6-7'6 xO'8-1'6 um from elongate conidiogenous cells apparently arranged on
short conidiophores (Fig. 36C-D). The material on which this taxon was based has not been
located inB (B. Hein, in litt.)or MSTR (B. Gries, in litt.)\ cryptogam collections in BRM are
currently unavailable during re-housing (H. Kuhbier, in ////.), and any at DR would have
been destroyed in 1945 (F. Mattick, in litt.}.

LICHENICOLOUS COELOMYCETES 89

The galls produced by this fungus recall those of Epicladonia sandstedei (see
p. 16), but the waxy, superficial pycnidia, and the simple and much narrower
conidia, make it unlikely to belong there. Bachmanniomyces uncialicola is easily
excluded as a possibility on the basis of conidial shape (p. 11), and E. simplex (p. 19) by the
much shorter and narrower conidia.

As I have seen no fungus recalling Tobler's description in all details, the applica-
tion of his name must currently remain uncertain. However, the possibility that
Tobler was dealing with an Epicladonia species cannot be entirely discounted.

Acknowledgements

Without the important studies of Dr B. C. Sutton on generic names and generic concepts in the
Coelomycetes as a whole the present investigation would have been impossible. I have also
benefited from his comments as to the placement of various taxa treated in this work, but stress
that the decisions taken are my own. I have also continued to receive a steady stream of enig-
matic specimens from Mr B. J. Coppins, which not only led to the discovery of some hitherto
unrecognized taxa, but further enabled species concepts to be clarified through the study of
more than single collections.

Any revision of this type is heavily dependent on the availability of material from other herbaria. I
am indebted to the curators and directors of all herbaria cited in the text (see p. 5)
for enabling me to study collections in their care, and further to the following for assistance in
trying to locate particular specimens: T. Ahti, S. Chiesa, B. J. Coppins, B. Gries, M. Guerlesquin, B.
Hein, H. Hertel, H. Kuhbier, F. Lona, R. Moberg, J. Mouchacca, U. Passauer, H. Riedl, Y. Rondon, L.
Tibell and O. Vitikainen.

Miss F. J. Walker was of particular help in tracking down elusive collections at the British
Museum (Natural History) and obtaining copies of some papers. Most of the sections illustrated were
cut by Mrs C. Beer, and Mr D. W. Fry prepared the habit photographs included.

Mr P. W. James and J. R. Laundon are thanked for their continuing interest in my studies on
lichenicolous fungi and for their assistance in the publication of the present revision.

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Host index

Names of lichen-forming fungi mentioned in the introductory sections and elsewhere are also included
here. Accepted names are in roman and synonyms in italic.

Anaptychia ciliaris 37 Bryoria capillaris 79

Anisomeridium biforme 80 fuscescens 37

Buellia disciformis 29-30

punctata 8 1
Bacidia 63
inundata 8 1

rubella 88 Calicium9,80

spododes%\ glaucellum9-10,44

Baeomyces rufus 56 trabinellum 8-10

Bilimbia spododes 85 viride 42, 44

Blarneya hibernica 63 Caloplaca caesiorufa 38

LICHENICOLOUS COELOMYCETES

93

cerina 38, 50-5 1,66
ferruginea 38
flavovirescens 38
holocarpa 66
lactea 38
regalis 84
vitellinula 74
Cetraria nivalis 72

sepincola 37
Cetrelia olivetorum 37
Chaenotheca 44
brunneola 44
chrysocephala 44
subroscida42,44
trichialis 44

Chrysothrix chrysophthalma 45-46
Cladonia3, 15-17, 19,26,36-37,39,41,87
acuminata41
alcicornis 39
amaurocraea 11,13
anomaea 1 7
arbuscula 36-37, 41
bacillaris 88
bacilliformis 19-20
bellidiflora 13
caespiticia 15, 18-19
cariosa 37
carneola 17
cenotea 36
chlorophaea 17, 19
ciliata var. tenuis41
coniocraea 17,21,35,41
conoidea 19
cornuta 18,41
cyanipes 18
deformis 18
digitata 35

fimbriata!8,36,39,41
foliacea39,41
furcata25-26
glaucal8-19
gracilis41
grayi 18
impexa 4 1
incrassata 36
major 1 8

merochlorophaea 18-19
norrlinii 4 1
ochrochlora 13, 18
pleurota 1 8
pocillum 36
portentosa 12,41
pyxidata 18-19,21,36,41
rangiferina 82

f. moribunda 37
rangiformis 82
silvatica 13
stellaris 13
stellata f. adusta 1 2

subulata 18

surrecta 13

sylvatica 4 1

terrae-novae 13-14

turgida 13, 18

uncialis 1 1-14
f. biuncialis 12
f. leprosa 1 2
var. leprosa 1 3
Cliostomum griffithii 3
Collema tenax 84

Diploicia canescens 86
Diploschistes scruposus 38
Dirina ceratoniae 86-87

Evernia prunastri 35-36, 38

Gyalectidium 75
filicinum 75

Haematomma elatinum 77-78
Heterodea muelleri 35
Heterodermia speciosa 74
Hypogymnia bitteriana 35

encausta 76, 79

intestiiniformis 76-77, 79

physodes 35-37, 83

Lasioloma 63

arachnoideum 64
Lecanactis abietina 79

subabietina 3, 79
Lecania 80

fuscella 76

syringe a 76
Lecanora36,65,72,84-85

admontensis 36

alphoplaca 49

atra 72

carpi nea 36

chlarona35, 85

chlarotera36,65-67,85

confusa 38

conizaeoides 35-36

dispersa38,66,81-82,84

e/fusa 84

expallens38

glaucella!5,S\

laevis 67

pacifica37

pallida36,38,82

piniperda 75, 81

polytropa36,38,82

94

D. L. HAWKSWORTH

rubina 14

rupicola 72

saligna 38, 80-8 1,84

subfuscata 36

superfluens 36

umbrina 81-82

varia 38
Lecidea 38

canescens 86

enteroleuca 38

leucocephala C. patelliformis 79

luteola 88

parasema 85

verruca 3

Lecidella elaeochroma 38, 80, 85
Letharia vulpina 37, 73
Lobaria pulmonaria 4, 14-15,73
Lopadium 59, 62-63

flammeum 59

phyllogenum 75

tayabasense 59

Mazosia 86

melanophthalma 78, 85-87

phyllosema 86
Melanelia 84
Micarea chrysophthalma 45

nitschkeana38,81,85

Ochrolechia parella 74
Opegrapha 78, 82

atra var. parella 38

cinerea 85

lithyrga 82

niveoatra28-29,78,81

subsiderella 78

vulgata 85

Parmelia 50, 70,72,83-84,87
acetabulum 83
afrorevoluta35
borreri4, 14, 36

caperata 5, 35-36, 5 1 , 56, 67, 7 1 , 76, 80
conspersa 37
crinita35
exasperata37
galbina 35-36
glabratula37,56,72

subsp. fuliginosa4, 14-15
jubata 6. cana 79
laevigata35-36,51-52,66
laevior 82

mollis y. divaricata 79
olivacea 37, 83
pastillifera 36

perforata35, 83

perlata 5, 35-36, 51-52, 67, 71 , 83

pulla37

pulverulenta 82

reticulata 51-52

revoluta 35

rudecta 37, 76

saxatilis 35-37, 67, 70, 73-74

sulcata 4, 14-15,36,51-52,76

verruculifera 37
Parmeliopsis ambigua 35
Peltigera 55, 72-74, 76

canina 24-25, 54,76

erumpens 3 1

horizontalis 54

malacea 50, 54

polydactyla 31, 54

pusillaSO

rufescens 22, 24-25, 54

spuria30-31,33
Pertusaria 29, 38

albescens 38

hymenea 35

Ieioplaca29,38,66

pertusa29,35

pustulata f. glabrata 38
Phaeophyscia orbicularis 81
Physcia 82

aipolia 36-37, 50, 56

stellaris 37, 82
Physconia 82

pulverulacea 37, 50, 56-57, 82
Platismatia glauca 4, 14, 54, 67, 69-70, 72
Polycauliona regalis 84
Psorotheciopsis 75

Ramalina67

baltica 37

calicaris37, 69

dilacerata 69

fastigiata 69

fraxinea37,67,69,73

implectens 75

siliquosa 37

sinensis 69

subgeniculata 37

yemensis 35

Rhizoplaca chrysoleuca 4, 14-15, 36, 72
Rinodina exigua 82

septentrionalis 38

Solorina crocea 78
Sporopodium 59
Squamarina lentigera 36

Tapellaria 62

Teloschistes 84
Thelomma siliceum 48

LICHENICOLOUS COELOMYCETES

Verrucaria 82

95

unidentified corticolous 8, 59, 73

foliicolous 66

saxicolous 47^8
Usnea37, 50, 54, 59

barbata var.florida 54

filipendula37, 54

fiorida35,37,54

Xanthoria 72, 77
fallax 77

parietina37,72-73,81-82
polycarpa 37

Zahlbrucknerella africana 73

Fungus index

In addition to synonyms cited in this revision, this index contains some synonyms of accepted
species for which further information is provided in papers cited in the text. New taxa, and the
page numbers of the principal references to accepted species, are printed in bold, accepted
names in roman, and synonyms in italic type. An asterisk (*) indicates pages on which there is a
figure.

Abrothallus 67, 70-7 1,87

bertianus 70, 72

cetrariae 72

glabralulae 72

microspermus 70, 76

moorei 13

parmeliarum 70-72, 76

peyritischii 72

smithii 70, 72

suecicus 69

usneae 58, 72

Acleistomyces rionegrensis 59
Alysia 64

pithospora 66

Aposphaeria cladoniae 39, 4 1
var. floerkeanae 39,41

ramalinae 75

stenospora 20-2 1
Arthonia clemens 72

destruens 72

epiphyscia72, 81

fuscopurpurea 72

glaucomaria 72

varia 72
Ascochyta 7-8

lichenoides 5-6,8

pisi 7
Ascochytula lecanorae = Lichenodiplis

lecanorae
Asterophoma 7, 8-10, 45

mazaediicola 8, 9*

Bachmanniomyces 6, 10-14, 16

uncialicola3,10, 11-12*, 13-14, 16,87,89

Campylidium 59
Celidiumfuscopurpureum 72

insidens 72

stictarum 73

Chaetothyriolum puiggarii 75
Chlorocyphella 59

aeruginascens 60
var. convoluta6Q
var. cystidiifera 75
var. staurospora 63

foliicola 63

lichenicola 60

subtropica 60
Clathridium corticola 88
Conida clemens 72
Coniosporium pertusariicola = Laeviomyces

pertusariicola
Coniothyrium cladoniae 87

epiphyllum 75

harmandii 75

imbricariae = Lichenoconium usneae

jaapii = Lichenoconium usneae

lecanoracearum s. str. = Lichenoconium
usneae

lecanoracearum auct. = Lichenoconium
lecanorae

lichenicola = Lichenoconium lichenicola
var. buelliae = Laeviomyces pertusariicola

pyxidatae = Lichenoconium pyxidatae

ramalinae = Lichenoconium cargillianum

usneae = Lichenoconium usneae
Coniscosolen mirabilis 75
Cornutispora 6, 14-15

lichenicola 4-5, 14, 15*, 51

limaciformis 14

96

D. L. HAWKSWORTH

Cristidium pallidum 75
Cyphella aeruginascens 60

foliicola 63

subcyanea 60

Dacampiosphaeria rivana 73, 78
Dendrophoma 39

alcicornaria 39

lecanorae 75

podetiicola 39
Dichaena pertusariae 38
Dichosporium glomerata 73
Didymella parvispora 73
Dinemasporium 39
Diplodia lecanorae = Lichenodiplis lecanorae

parmeliae 76
Diplodina claudelii 16, 19

lecanorae = Lichenodiplis lecanorae

lichenoides = Ascochyta lichenoides

parmeliae 76

peltigerae 19, 74, 76

sands tedei 15-16, 19

solorinaria 78

vouauxii ? = Lichenodiplis lecanorae
Discosporium 1 1

populeum 1 1
Dothiorella erumpens 87

Endococcus 83

Endophragmiella hughesii 15

Epicladonia 6-7, 15-22, 89
sandstedei 3, 15,16, 17-18*, 19-20,89
simplex 16, 19,20-21*, 89
stenospora 16, 17*, 19,20,21,22*

Epicoccum parmeliarum 70, 76

Griphosphaeria corticola 88
Guignardia olivieri 73, 8 1

Hansfordiellopsis lichenicola 75
Render sonia lichenicola 87-88
Herpotrichia 84
Homostegia lichenum 73
piggotii 73

Karsteniomyces 6, 22-25
peltigerae 22, 23-24*, 3 1 , 74

Keissleriomyces 6, 25-26
sandstedeanus 25, 26-27*

Keratosphaera batistae 86

Laeviomyces 5-6, 26-30, 33 >
opegraphae 28*, 29*, 78
pertusariicola 27-28, 29*

Lecidea cladoniaria 1 3

irregularis 60
Leptodothiorella 73
Leptosphaeria 49
Lethariicola sipei 73
Levieuxia borealis 1,33
Libertella 30
Libertiella5,7,30-33

malmedyensis 30, 3 1-32*, 83

obscurior 76

peltigerae 30-32

xanthoriae 77
Lichenoconium 5-6, 27, 33-37

boreale33

cladoniae%l

cargillianum 33, 34*, 35

echinosporum 34*, 35

erodens3,33,34*,35,36,51

imbricariae = Lichenoconium usneae

jaapii = Lichenoconium usneae

lecanorae 14-1 5, 34*, 35, 36

lecanoracearum auct. = Lichenoconium
lecanorae

lecanoracearum^. str. = Lichenoconium
usneae

lichenicola 33, 34*, 36
var. buelliae = Laeviomyces pertusariicola

parasiticum 36 = Lichenoconium lecanorae

pertusariicola 29, 33

pyxidatae33,34*,36, 37

usneae 14-15, 34*, 35, 37, 58, 87

xanthoriae 33, 34*, 37
Lichenodiplis 5-6, 27, 37-38

lecanorae 29, 37, 38, 66, 74

lichenicola 37,38
Lichenomyces lichenum 73
Lichenophoma 77

haematommatis 77*

opegraphae 78
Lichenosticta 5-6, 38-41

alcicornaria 39, 40*, 41

podetiicola 39,41

Melophia woodsiana 60
Merismatium lecanorae 74
Microcalicium 7, 41-44, 7 1

conversum 42, 43*, 44

disseminatum 42

subpedicellatum 10,42, 43*, 44
Microdiplodia effiisae = Lichenodiplis lecanorae

ferrugineae = Lichenodiplis lecanorae

lecanorae = Lichenodiplis lecanorae

lichenicola^ = Lichenodiplis lichenicola
Microthelia alcicornaria 39,41

cargillianum = Lichenoconium cargillianum

solorinaria 78
Minutophoma 7-8, 44-46

chrysophthalmae 44, 45*, 46

LICHENICOLOUS COELOMYCETES

97

Muellerella lichenicola 29, 74

Nectria heterospora 76

parmeliae 76

tenacis 84
Nesolechia cladoniaria 1 3

oxyspora 74
Nicholsonietta 30

malmedyensis 30
Nigropuncta 46-49

rugulosa 3, 6, 46, 47^8*, 49

Orthidium 59

Phacopsis clemens 72

huuskonenii 79

varia 72
Phaeoantenariella 78

lichenicola 78
Pharcidia dispersa 74
Phoma 5, 7-8, 1 1 , 44, 49-57, 69, 80, 83

abietinae 79

alectoriae 79

arachnoidea 79

biformis 80

caloplacae 50*, 5 1

caperatae53,57,80

cladoniae 39

curvispora 80

cytospora50,51,52*, 53,55

dubia50,53*,54

epiparmelia 80

epiphyscia 8 1

fusispora 8 1

glaucellaSl

herbarum 49

lecanorae 8 1

lichenis 82
f. immersa 82

parmeliarum 76

peltigerae 50, 54, 55*, 81

physciicola 50, 56*, 57
var. caperatae 80

ramalinaebl

truncata = Vouauxiomyces truncatus

uncialicola 1 1

usneae51,19

verrucariae 82

versoniana 83
Phyllosticta 50, 55

cytospora 51,83

galligena 83

lichenicola 53, 83

peltigerae 54-55

uncialicola 10-1 1, 13
Phyllostictina 50

Phymatopsis dubia 54
Pleospora 49

hooked 78
Pleurosticta 83-84

lichenicola 83
Polycoccum 83

rugulosaria 84
Psammina 7

Pseudodiplodia lichenis 84
Pseudoseptoria 7, 57-59

donacicola 57

donacis 57

usneae5, 16,57,58*
Pyrenochaeta 84

collematis 84
Pyrenotrichum 7, 59-64, 70, 75

atrocyaneum 59

bicolor 59

filiferum59,63

foliicola 59, 63

irregulare 59

mirum 59

podosphaera 59

splitgerberi 3, 59, 60, 61-62*, 63, 75

staurosporum 59, 60, 63, 64*

Rhabdospora antarctica 84-85

lecanorae 85

lesdainii 85

thallicola 85
Rhagadostoma lichenicola 78

Schecutia 30

malmedyensis 30
Sclerococcum 27, 46
Scutula epiblastematica 74, 76

wallrothii 74
Seimatosporium 88

lichenicola 87-88
Selenophoma 57
Sirococcus 1 1

lichenicola 1 1

strobilinus 1 1
Sirothecium lichenicola = Vouauxiella

lichenicola

f. bispora = Lichenodiplis lecanorae
f. cerinae = Lichenodiplis lecanorae
Sphaeromma mazosiae 85-87
Sphaeronema lichenophilum 86-87
Sphaeropsis cladoniae 87

scripta 87
Spilomium 27

epicladonia 14,87

pertusariicola 27, 29
Sporhaplus rondoniensis 86-87
Sporocadus 87

lichenicola 87

98

D. L. HAWKSWORTH

rosicola 88
Stagonopsis22, 84

lichenis 84

pallida22

peltigerae22, 55
Stagonospora25

paludosa25

sandstedeana 25
Stigmidium dispersum 74

Taeniolella66

Telimena foreaui 74

Tichothecium erraticum 74

Torula lichenicola = Vouauxiella lichenicola

verrucosa = Vouauxiella verrucosa
Trichosperma aeruginosa 60

cyphelloidea 60

Verrucaster 16

lichenicola 88*
Vouauxiella 3, 6, 64-67

lichenicola 65, 66-67

pithospora 65,66

uniseptata 5, 46, 65, 66

verrucosa 65,67
Vouauxiomyces 7, 67-71, 72

ramalinae67,68*,69,75

santessonii 5, 67, 68*, 69, 7 1-72, 76

truncatus 5,67, 68*, 69, 70, 7 1 *, 76, 80

Xeroconium 33
boreale 33

Zythia peltigerae 30

Manuscript received 25 September 1980

\

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Titles to be published in Volume 9

The lichen icolous Coelomycetes. By D. L. Hawksworth

The genus Callithamnion (Rhodophyta : Ceramiaceae) in the
British Isles. By P. S. Dixon and J. H. Price

Parmelia subgenus Amphigymnia (lichens) in East Africa.

By Hildur Krog and T. D. V. Swinscow

The genus Selaginella in tropical South America.

By A. H. G. Alston, A. C. Jermy and J. M. Rankin

Printed by Henry Ling Ltd, Dorchester

f A Mi

Bulletin of the

British Museum (Natural History)

The genus Callithamnion (Rhodophyta:
Ceramiaceae) in the British Isles

Peter S. Dixon & James H. Price

Botany series Vol 9 No 2 26 November 1981

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World List abbreviation: Bull. Br. Mus. nat. Hist. (Bot.)

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The Botany Series is edited in the Museum's Department of Botany
Keeper of Botany: Mr J. F. M. Cannon

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Assistant Editor: Mr J. R. Laundon

ISSN 0068-2292 Botany series

Vol 9 No 2 pp 99-141

British Museum (Natural History)

Cromwell Road

London SW7 5BD Issued 26 November 1981

r f3f

*> GENER

The genus Callithamnion (Rhodophyta:
Ceramiaceae) in the British Isles

Peter S. Dixon

Department of Ecology and Evolutionary Biology, University of California, Irvine,
California 927 17, U.S.A.

James H. Price

Department of Botany, British Museum (Natural History), Cromwell Road, London
SW7 5BD

Contents

Synopsis 100

Introduction 100

Aspects of structure and reproduction 100

Analysis of character validity: adult material 101

1 . Main axis characters 101

(a) Integrity of a main axis throughout the thallus 102

(b) Degree of cortication of the main axis and major lateral axes . . . 102

(c) Development of lateral filaments from the cortication .... 104

(d) Cell dimensions 104

2. Characteristics of major laterals 105

(a) Branching patterns of major laterals 105

(b) Number of orders of laterals distinguishable 105

(c) Reflexion of major laterals 105

3. Characteristics of peripheral laterals 105

(a) Branching patterns of peripheral laterals 105

(b) Occurrence of 'secund processes' 106

(c) Overall shape and size 106

(d) Hyaline hairs 106

4. Reproductive characteristics 106

(a) Sporangia 106

(b) The gametangial phase 107

(c) The carposporophyte

5 Gross criteria of species and groups

The genus Callithamnion Lyngbye

General key to British species

Accepted species

1 . Callithamnion byssoides Arnott ex Harvey

2. Callithamnion corymbosum (Smith) Lyngbye

3. Callithamnion decompositumL Agardh

4. Callithamnion granulatum (Ducluzeau) C. Agardh

5. Callithamnion hookeri(Di\lv/yn)S. F. Gray

6. Callithamnion roseum (Roth) Lyngbye

7. Callithamnion sepositum (Gunnerus) P. Dixon & J. Price ....

8. Callithamnion spongiosum Harvey

9. Callithamnion tetragonum (Withering) S. F. Gray

10. Callithamnion tetricum(Dil\wyn)S. F. Gray

Rejected species

1 . Rejections from the British flora

2. Entities of uncertain attribution

3. Rejections from the genus

Typifications

Acknowledgements

References

Bull. Br. Mus. nat. Hist. (Bot.) 9 (2): 99-1 4 1 Issued 26 November 1 98 1

99

100 P. S. DIXON&J. H. PRICE

Synopsis

The algal genus Callithamnion is represented in the British Isles by 10 species: C. byssoides, C. corym-
bosum, C. decomposition, C. granulatum, C. hookeri, C. roseum, C. sepositum, C. spongiosum, C.
tetragonum and C. tetricum. There is one new combination: C. sepositum (Gunnerus) P. Dixon & J.
Price.

A critical description is provided for the genus and for each species, together with comments on those
features of structure and reproduction of systematic importance, detailed analyses of the validity of
these features for taxonomic distinction, and the range of variation encountered in the British Isles and
in adjacent regions. A general key to the British species is provided, together with details of the
typification of all entities relevant to a consideration of the genus in the British Isles.

Introduction

The genus Callithamnion presents a supreme example of the taxonomic confusion in
red algae.^The following are the principal reasons for this:

1 . A lack of understanding of the effects of the environment on an extremely 'plastic'
phenotype, capable of variation at many different levels.

2. The lack of data on the extent of that morphological variation and its relation to
the limits of subsidiary taxa in different geographical areas.

3. The misunderstanding, misinterpretation and confusion in the application of names at
different times, by different workers, in different countries.

The present paper is the beginning of an attempt to rationalize treatment of this group of
algae, in terms of morphology, development, variation and systematics.

Our basic orientation has been towards those entities occurring on or reported from British
shores. As a result of carrying out typifications and dealing with doubtful reports and
records, it became necessary to refer to entities from elsewhere, mostly in western Europe but
also in other parts of the Atlantic and Mediterranean. Since the geographical confines of this
study were envisaged originally as being from North Cape to the Algarve, with treatment of
the genus in full to be undertaken only for the British Isles, data on Callithamnion in areas
outside Great Britain and Ireland have been considered only in so far as they affect the
situation in the British Isles. Treatment of the genus in those areas outside the British Isles is,
therefore, not complete.

Aspects of structure and reproduction

The thallus in species of Callithamnion is of uniseriate filamentous construction, although
this may become obscured through the development of 'cortication'. Each uniseriate
filament results from the activity of a single apical cell which divides transversely or
obliquely. Once formed, a product cell does not divide again in the plane in which it was
produced, so that the number of cells in an undamaged filament is a reflection of the number
of divisions of the apical cell. Cells may be uninucleate or multinucleate when mature and,
apart from rhizoidal cells, each contains several pigmented chloroplasts. The shape and
number of chloroplasts are modified by age or the conditions in which the thallus is growing.
Although a product cell does not divide again in the plane of division in which it was
formed, it frequently does divide once more in a different plane to form the primordium of a
lateral branch. Each primordium is clearly of lateral origin, although the final disposition of
branches may appear dichotomous if evection occurs.

Each thallus in Callithamnion is thus a complex pattern of filaments. A few filaments are
of unlimited or unrestricted growth, whereas most are of limited or restricted growth. The
difference between these two types is of a physiological nature, with few easily-definable
morphological differences. Interconversions can occur, although for many species there are
characteristic arrangements of filaments of limited and unlimited growth. In certain species,
other lateral filaments may form which grow downward, investing axes, to form the
'cortication'. These 'corticating' filaments are composed of pigmented cells and they arise

CALLITHAMNION W THE BRITISH ISLES 101

from the basal poles of axial cells or from the lowermost cells of lateral filaments. The latter
cannot be clearly distinguished from 'corticating' filaments as either can readily give rise to
the other. Nevertheless, the term 'corticating filament' has utility in indicating function and
position at that time. The extent of cortication varies considerably, even within and between
individuals of the same species.

Following the formation of lateral primordia, cells undergo considerable enlargement so
that the volume of mature cells may be as much as 50,000 times the original volume. This
enlargement is almost entirely in the basal region of a cell, so that in the mature state the
lateral filament is attached at the apical pole. Enlargement is associated with development of
a large central vacuole and as a result the cytoplasm becomes a peripheral lining within the
wall. Cell size varies considerably, not only with respect to age but also to environmental
modifications caused by temperature, light intensity, light quality and photoregime. There
are characteristic patterns of variation in cell size due to interactions between laterals and the
axes on which they are borne. In some species of Callithamnion this is shown most clearly in
cells at the base of lateral filaments. These cells are older than cells further along the lateral
filaments so that they should be larger, but their smaller size indicates that some retardation
of enlargement has occurred, possibly through proximity to the axial filament. There may be
other factors involved, since a similar phenomenon occurs in main axes (cf. Dixon, 1971;
Price, 1978).

A thallus is attached to the substrate by multicellular rhizoidal filaments which resemble
'cortication' but differ in that pigmented chloroplasts are absent. Such filaments also
occasionally develop from the cortication as adventitious outgrowths (Price, 1978). When
spores germinate, they divide into an upper and a lower cell, the latter forming only the
primary rhizoid. Subsequently, additional rhizoids are formed, initially from the lowermost
cells of the axial filament and then from the lowermost cells of lateral filaments.

This discussion of the developmental morphology of Callithamnion may at first sight
appear to differ markedly from the descriptions given in recent years by various French
workers. These use an elaborate system of nomenclature first proposed by Chadefaud
(1954; see also 1979). In its original form, this system carried the implication that morpho-
logical categories were rigidly distinct one from another, although in recent publications by
L'Hardy-Halos (1970, 197 la, b, c) a more flexible interpretation has been adopted which we
consider relates better to the realities of morphogenetic plasticity in Florideophyceae. This is
not the place for a detailed analysis of the French system, and the reader is referred to the
publications cited for its details. It seems to us that, with few exceptions, the differences of
opinion between the system which we have used and the French system are now largely
semantic.

In its vegetative morphology, Callithamnion resembles some other genera of the
Ceramiaceae, from which it differs in reproductive details. Callithamnion and its immediate
allies, the 'Callithamnion-Gruppe'' of Kylin (1956), are characterized by the lateral
intercalary position of procarps. After fertilization, the support cell divides to produce an
auxiliary cell. Once contact has been established with the carpogonium, the auxiliary cell
gives rise to gonimoblast initials. These form the filaments of the carposporophyte; the latter
may be single and entire or deeply lobed. In either event, each cell of the gonimoblast
develops into a carposporangium which produces a single carpospore.

Various other types of spores occur in the genus. Tetrasporangia are probably of the widest
occurrence. Structures producing more than four spores have been described and these
appear to be parasporangia (cf. Drew, 1939), while structures producing only two spores
have been observed. Claims for the occurrence of mature, functional monsporangia cannot
be substantiated. For comments on polysporangia, see p. 106.

Analysis of character validity: adult material
1. Main axis characters

The characteristics of main axes demand careful examination to ensure that the structure

102

P. S. DIXON & J. H. PRICE

under investigation is the main axis. Major branches may resemble the primary axis but the
identity should not be assumed without question. Analysis of main axis characteristics
requires a whole thallus, but herbarium specimens of Callithamnion are not always
complete. Thalli were often 'pruned' to produce a specimen with a greater aesthetic appeal,
while thalli occurring on animals are usually stripped from the substrate during either
collection or preservation and thereby often damaged.

(a) Integrity of a main axis throughout the thallus

Many authors have characterized species of Callithamnion by the degree of distinctness of
the main axis, differentiating between species in which a main axis is easily distinguishable
and those in which it is not. This feature depends in part on whether branching is obviously
lateral, and on the extent to which evection occurs so that the branching is of a more
dichotomous aspect. From personal observations, the degree to which a main axis remains
identifiable in a species is highly variable and the feature is of little value of taxonomic
discrimination.

(b) Degree ofcortication of the main axis and major lateral axes

Virtually all treatments of the genus Callithamnion have made reference to cortication. Only
rarely has this feature been discussed with any degree of precision or with reference to its
variability. Appreciation of the relationship between degree of cortication and environ-
mental factors is a relatively recent development (Boddeke, 1958; Harris, 1962; Price, 1978).

B

Fig. 1 A, Callithamnion tetragonum: a well -corticated axis showing a primary lateral and two
adventitious axes arising from the cortication (x 70). B, C. sepositum: transverse section of a
well-corticated axis (x 35). C, C. byssoides: origin ofcortication by down-growth from the basal
cell of a primary lateral (x 250).

CALLITHAMNIONM THE BRITISH ISLES

103

CO

o
o

CO

0<

(LU//UI) 1130 1VIXV dO H19N31

104

P. S. DIXON & J. H. PRICE

These authors comment that cortical and adventitious development is greatest in entities
tolerating or exposed to wave action. This increase in cortication density in such entities
appears to result from two causes. There is an increase in the formation of the downward-
growing filaments which constitute cortication. Also, without exception, entities occurring
in conditions of strong wave action display cells (in the main axis and major laterals) which
are short compared with their diameter. As a result, the amount of cortication is increased
while the surface area to be covered is less than in an entity where cell enlargement is greater
(Fig. 1).

(c) Development of lateral filaments from the cortication

The down-growing corticating filaments can give rise to adventitious lateral filaments and
the occurrence of these has been used by some authors (e.g. Rosenvinge, 1924) to
characterize certain entities. Maximum expression of this feature occurs in those entities
tolerant of or exposed to severe wave action, where cortication development is greatest. The
feature is of little general use as a strictly taxonomic criterion in the genus Callithamnion
(Price, 1978).

(d) Cell dimensions

The dimensions or length-to-breadth ratios of cells have been used extensively for discrimi-
nating between entities in the genus Callithamnion (e.g. Kylin, 1907; Newton, 1931;
Boddeke, 1958) even though some authors (e.g. Rosenvinge, 1924) have questioned the value
of this feature. It has been demonstrated (Dixon, 1971) that there are elaborate patterns of
cell enlargement in species of Callithamnion, with regard to both main axes and principal
laterals (Fig. 2). Patterns of variation in length-to-breadth ratios are even more complex
(Fig. 3). In Pleonosporium, which is of similar morphology to Callithamnion, variation in
cell size has been related to a complex interaction of light intensity and duration (Murray &
Dixon, 1972, 1975). There are indications that cell size variation in Callithamnion is
controlled by similar environmental factors.

The degree of variation in cell size makes it impractical except in certain instances to use
this feature for taxonomic discrimination. A further implication of the available studies is
that if cell sizes or ratios are quoted for any entity, the positions of the cells measured must be
precisely cited. For further detailed consideration, see Price (1978).

APEX

10

20 30 40 50

SEGMENT NUMBER

BASE

Fig. 3 Callithamnion corymbosum: histogram showing the length/breadth ratio of every cell

along the principal axis of a small thallus.

CALLITHAMNION IN THE BRITISH ISLES 1 05

2. Characteristics of major laterals

The most important characteristics of the major laterals in the genus Callithamnion, from
the taxonomic standpoint, are in their branching patterns and arrangement. As previous
authors (e.g. Boddeke, 1958; Harris, 1962; Price, 1978) have noted, differences in branching
patterns may occur in the same entity in different reproductive states so that the condition of
material must always be indicated in discussion of this feature. The arrangement of major
laterals in Callithamnion is frequently described as 'phyllotaxis', a term not strictly
applicable to the algae in the absence of leaves. The alternatives are clumsy or even more
inaccurate so that, despite the reservations on its use, we employ 'phyllotaxis' for the
arrangement of major laterals.

(a) Branching patterns of major laterals

Of the several arrangements of filaments that are related to patterns of branching and have
been employed in taxonomic discrimination within the genus, two are most significant.

1. The relationship between two individual filaments derived from a common cell lineage;
the arrangement may be alternate or pseudodichotomous.

2. The relationship between successive lateral filaments along an axis; these may lie in a
single plane (giving a distichous or biseriate arrangement) or be distributed radially
(giving a polystichous or spiral arrangement).

Both are potentially useful criteria, although there can be variations in different parts even of
a single specimen. Some diagrams illustrating 'phyllotaxis' have been published by L'Hardy-
Halos (1970) and more are being prepared for use elsewhere.

(b) Number of orders of laterals distinguishable

With few exceptions (e.g. Callithamnion caudatum), the number of orders of laterals is
highly variable depending on a complex of factors, such as the species, the age of the thallus,
its vigour and the environment in which it occurs. This feature is therefore of little
taxonomic value for specific discrimination.

(c) Reflexion of major laterals

The tendency of major laterals, particularly of older basal laterals, to be reflexed has been
used as a characteristic of certain entities (e.g. Westbrook, 1927). The feature is highly
variable and of little taxonomic value.

3. Characteristics of peripheral laterals

The thalli of species of Callithamnion are richly branched and the overall form is a reflection
of the branching pattern of the constituent filaments. The form of the thallus is determined
not only by the arrangement of major laterals, discussed in the preceding section, but also by
the arrangements which prevail in the ultimate portions of the constituent filaments. It has
become apparent from our investigations that considerable taxonomic dependence can be
placed on the morphology, branching pattern and spatial arrangements of what we term the
peripheral laterals, so-called because they contribute to the periphery of the thallus.
Previous workers with the genus have made extensive use of this feature but, in almost every
case, each has used a different term by which to refer to it. A cursory survey of the literature
disclosed more than 50 terms which had been applied, of which the most widely used were
probably ramuli, ramelli, branchlets, pinnae, and pinnules. The use of different terms, often
with variant applications, has produced an impossibly complex terminological situation and
it is beyond the scope of the present paper to discuss every usage.

(a) Branching patterns of peripheral laterals

The branching patterns of major axes can differ markedly from those of peripheral laterals,

106 P. S. DIXON & J. H. PRICE

such differences often being associated with the formation of reproductive structures. Never-
theless, branching patterns at the peripheral level are usually sufficiently distinct and
characteristic for taxonomic discrimination.

(b) Occurrence of secund processes'

One particular aspect of the branching pattern of peripheral laterals relates to the occurrence
of ultimate lateral filaments which consist only of a single cell. These have been termed
'secund processes' (e.g. Newton, 1931) and used for taxonomic discrimination. Our investi-
gations show that these structures are nothing more than early developmental stages of other
organs or laterals.

(c) Overall shape and size

The overall shapes and sizes of peripheral laterals and of their constituent cells have been
used by many authors (e.g. Rosenvinge, 1924; Newton, 1931; Feldmann-Mazoyer, 1941) in
diagnoses and for discrimination. In our experience, the sizes of entire peripheral laterals are
very variable although the shapes and relative sizes of constituent cells are more constant. In
addition, other features of a peripheral lateral, such as the number of cells which it contains
and its pattern of branching, are particularly significant.

(d) Hyaline hairs

Apical or sub-apical hyaline hairs occur frequently on peripheral laterals in some species of
the genus. The function of these structures is unknown and, although widespread, their
occurrence and abundance are subject to great variation. A further difficulty preventing the
use of these structures for taxonomic discrimination is that they may be damaged or
destroyed by fixation or the preparation of herbarium specimens. For further detailed
considerations, see Price (1978). Culture conditions may induce or permit the formation of
such hairs where field conditions would not. Rosenvinge (1924) found hairs so rarely in
Callithamnion byssoides (as C. furcellariae) that he suggested so distinguishing the taxon
from C. corymbosum; Rueness & Rueness (1980), by contrast, frequently observed
terminal hyaline hairs on female gametophytes of C. byssoides in culture.

4. Reproductive characteristics

Reproductive characteristics tend to be relatively stable but the structures are ephemeral and
their use as a source of characteristics has been kept to the minimum in this general
treatment.

(a) Sporangia

Monosporangia have been reported in various species of Callithamnion. Field reports are of
immature tetrasporangia, while reports in culture must be dismissed as artifacts so that we
have ignored monosporangia for taxonomic purposes.

Bisporangia are produced inconsistently, even in entities where they have been reported,
sometimes alone or mixed with tetrasporangia. Their occurrence cannot be relied upon as an
absolute criterion. For their significance, see Guiry (1978).

Tetrasporangia are very common and their position, size and shape have been much
employed taxonomically in the genus Callithamnion, although these characteristics are
minimally useful.

A few instances of sporangia containing more than four spores have been detected in
British species of Callithamnion. These structures can be regarded as parasporangia rather
than polysporangia (sensu Drew, 1939). The structures are of rare occurrence and the
position is confused because C. decompositum (which here seems habitually to produce
parasporangia as the consistent reproductive structure) has previously been overlooked in

CALLITHAMNIONIN THE BRITISH ISLES

107

the British Isles (Price & Tittley, 1978). Until it can be confirmed that British C. hookeridoes
produce parasporangia, as in Scandinavia, or does not, their use as a taxonomic distinction is
best avoided.

(b) The gametangial phase

Previous authors have made considerable use of structures of the gametangial phase in
distinguishing between species. The principal difficulty is that these structures are com-
paratively ephemeral. For this reason, no attempt has been made to utilize data from the
male and female structures or carposporophyte in the present general key. Keys for morpho-
logical states other than the vegetative and for ecological characteristics are certainly useful.
Keys involving distinctions between the reproductive states of all accepted species will be
presented elsewhere (Price, in preparation) although available descriptions are offered in
species characterization. As otherwise indicated, periodicity of reproductive structures is
less useful and is discussed but not employed.

In the male organs, group distinctions based on the degree of branching in the sperma-
tangiophore system and the positions of spermatangia are reasonably reliable. Positions of
the spermatangial clusters and their mature form have also been employed. This is not a new
approach; many authors have attempted to utilize these features.

ABC

Fig. 4 Cell patterns in the carpogonial branches of different species of Callithamnion. A, C.
corymbosum, B, C. byssoides. C, Material identified as "C. tripinnatum\ After Miranda (1934).
Diagrammatic, not scaled.

The employment of cell spatial arrangements and alignment in the carpogonial branch
owes much to observations by Miranda (1934), but has been subsequently developed by
other authors. Apart from the ephemeral nature of the material, difficulties of accurate
observation of the arrangement of cells precludes use of the data at a principal distinguishing
level, but we have employed these data in other contributory capacities. The original
concepts put forward by Miranda are shown diagrammatically in Fig. 4; our suggested
modifications of or additions to these, with terminological clarification, appear as Fig. 5.
Since there is some variation in arrangement and cell relationship, it is highly likely that
these aspects of the carpogonial branch depend at least on the relative position of the branch
on the thallus, on the bearing support cell, and on the bearing axis. The general position
away from functioning apices may reflect the level at which vegetative apical dominance is
no longer inhibitory.

Sizes and shapes of carpogonia, trichogynes, and other cells of the procarp all appear too
variable to be of taxonomic value at the species level. Trichogyne ramification, not often
reported, has been noted by us in a few entities, but its inconsistency suggests aberrations
and/or environmental influences. It is possible that different patterns of cell size relationship
would emerge for different species from a statistical survey of the carpogonial branch, but the
taxonomic utility of this is doubtful.

108 P. S. DIXON& J. H. PRICE

Fig. 5 Suggested modifications and additions to cell spatial arrangements and alignment types
recognized in carpogonial branches by Miranda (1934): variations especially emphasized. (All
measurements in um.) Pp. 109-1 12.

5A, Callithamnion tetragonum, straight form usual to species: Carpogonium width (c/w) 15.

Trichogyne length (t/1) 195.

5B, C. tetragonum, intermediate form near zigzag; c/w 7; t/1 99.
5C, C. tetragonum, intermediate form near curved: (i) c/w 1 1 ; t/1 48; (ii) c/w 9; t/1 64.
5D, C. granulatum, zigzag form usual to species: (i) c/w 1 8; t/1 239; (ii) c/w 1 8; t/1 7 1 .
5E, C. sepositum, curved form usual to species: (i)c/w 18; t/1 75;(ii)c/w 17; t/1-; (iii) c/w 26; t/1 106.
5F, C. tetragonum, straight form with extruded 2: c/w 14; t/l*l 7 1 .

Variations perceived probably depend on relative positions of branch on the axis, on the support cell,
and on the overall thallus. They are therefore a reflection of the balance between constraints and
overall space available. Types recognized certainly represent arbitrary divisions of a continuum of
variation. All figures presented are taken from actual carpogonial branches; dimensions given for the
carpogonial width and trichogyne length. Arrows and curves = space relations of cells determined by
focal-planing. Symbols: R = bearing axis cells or line of them; S = support cells (S,; S 2 ); 1 =cell 1 of
carpogonial branch; 2 = cell 2 of c/b; 3 = cell 3 of c/b; 4 = carpogonium with trichogyne.

Straightform

(Callithamnion corymbosum-type of Miranda, 1934; 'straight'-type of Harris, 1959, 1962, 1966).
Orientation of cells 1 - +2 usually at right angles to long axis of bearing axial cell (walls of 1/2
therefore parallel to that axis), as in Fig. 5A (example) and Fig. 4A (diagrammatic). Cells 1 - K2
and 2 - -*-3 are in linearly parallel arrangement, carpogonium (4) being variously arranged on 3
(Fig. 5A). Form occurs usually in C. tetricum, C. sepositum, C. corymbosum, C. roseum, and C.

tetragonum (Fig. 5 A), sometimes in C. granulatum, C. hookeri, C. byssoides and C. spongiosum.

Zigzagform

(Callithamnion byssoides-type of Miranda, 1934; "zigzag'-type of Harris, 1959, 1962, 1966).
Compound curve or zigzag (Fig. 4B, diagrammatic). Orientation 1 - +*2 inclined upwards or at
outward angle to long axis of bearing axial cell (Fig. 5D). Form occurs usually in C. byssoides, and C.
granulatum (Fig. 5D), sometimes in C. hookeri, C. spongiosum, C. roseum, C. tetricum, C.
sepositum, C. tetragonum and C. corymbosum.

Curved form

(Callithamnion 'tripinnatum'-type of Miranda, 1 934)

Single curve ('U'-shaped), often in more than one plane by lateral curvature (Figs 4C/5E). Form

occurs usually (?) in 1C. tripinnatum, often in C. sepositum (Fig. 5E), and probably sometimes in all

known species.

Variations of standard' forms

(i). Parallel status of 1 -- *-2/2 - >3 depends on position of pit connexion between cells 2 and 3.
If the connexion is high on 3 and 4 (carpogonium) is large or in cramped position, depression or offset
position of 3 will cause the relationship

4

to occur (as in Fig. 5B, where 4 (carpogonium) rises towards viewer out of the paper, but would
obscure detail if drawn fully so). This is very close to the form attributed by Miranda (1934) to C.
byssoides-type (Fig. 5D; here in examples of C. granulatum).

(ii). If cell 2 is depressed from the 'normal' C. corymbosum-type position, and pit connexions 1/2 and
2/3 are high on 2, then the 'U'-shape of Miranda's (1934) 'tripinnatum' -type is approached, as is
usual in C. sepositum (Fig. 5E) and occasional in C. tetragonum (Fig. 5C).

(iii). Arrangement of the carpogonial branch is not always planar, as is clear from the above and from
curvature diagrams accompanying parts of Fig. 5. Angling to the vertical or into more than one plane
is common to usual, and can confuse the pattern as, e.g. in Fig. 5F. There, a C. tetragonum branch
with 1 - *>2 - > 3 all still parallel has 2 extruded by pressure to jut out from the usual line.
Otherwise, the branch is clearly of C corymbosum-type (straight), the periphery of the lateral branch
system bearing the carpogonial branch being basal to the figure. Harris's (1959, his text-figure 13C)
intermediate type may have been due to this non-planar situation.

CALLITHAMN1ONM THE BRITISH ISLES

109

5A

5B

5C

ii

110

P. S. DIXON & J. H. PRICE

Support cell
not visible

5D ii

Fig. 5 Suggested modifications and additions to cell spatial arrangements and alignment types
recognized in carpogonial branches by Miranda (1934): variations especially emphasized. (All
measurements in urn.) Full explanation on p. 108.

CALLITHAMNIONW THE BRITISH ISLES

111

21 4

If flattened
probably c

5E i

Trichogyne
fractured in
position shown

\ s

Line of

peripheral

lateral

112

P. S. DIXON&J. H. PRICE

5F

Fig. 5 Suggested modifications and additions to cell spatial arrangements and alignment types
recognized in carpogonial branches by Miranda (1934): variations especially emphasized. (All
measurements in urn.) Full explanation on p. 108.

CALLITHAMNION IN THE BRITISH ISLES 1 1 3

(c) The carposporophyte

Events occurring after fertilization have a standard sequence. It appears that the extent of
development and number of cells involved in primary and secondary gonimoblast have a
relative constancy that is useful at a contributory level in species distinction. To some extent,
basic differences in the early carposporophyte development are reflected in lobing of the
carposporophyte at early maturity (carposporangial production). However, subsequent
environmental and morphogenetic effects obscure or override the earlier patterns and
produce variations in form so that species patterns cannot be distinguished. We have
employed such criteria with circumspection in the following treatment. Sizes and shapes of
carposporangia and carpospores are of little taxonomic value.

Development of a rudimentary vegetative 'involucre' around the carposporophyte occurs
in some of the more delicate species. It is referred to as necessary in the treatments that
follow, but it is inadvisable to employ this essentially ephemeral phenomenon as a
fundamental criterion.

5. Gross criteria of species and groups

In addition to particular features used as systematic criteria, numerous other gross criteria
are cited in diagnoses. Generally, these gross criteria are of little importance and we do not
employ them for taxonomic distinction.

Periodicity of upright growth is similar in all species, with one possible exception. It is
probable that the less robust forms rarely persist as recognizable basal fragments through the
winter, although it is difficult to be sure of that. Robust, exposed shore forms show rather
abrupt dieback to variable basal portions in the usual conditions of late autumn or early
winter. Those thalli which do persist have lost peripheral or even whole branch systems and,
although still recognizable, have a very different appearance from the luxuriant summer
growth. One exception, Callithamnion tetragonum, tends to persist as recognizable growths
on laminae of the Laminariales in some parts of its range. In some years and locations, C.
tetricum behaves similarly. Although contributory, this behavioural difference is not of
prime taxonomic importance.

Statistically, characteristic growth forms can be recognized for most species of
Callithamnion. Unreliable as primary taxonomic criteria, these are nevertheless very useful.
The growth forms are a consequence of growth vigour, robustness, texture, frond outline,
branching pattern, and other unquantifiable phenomena. In some species, external form is
almost indescribable but still highly characteristic. Species of Callithamnion rarely form
homogeneous populations with individuals closely-knit enough to possess a recognizable,
distinct population morphology. Two species form occasional exceptions to this. C.
roseum populations mainly manifest what is merely a multiplication of the individual
plant characteristics; C. scopulorum (now C. hookeri) has the most recognizable population
morphology that is rather more than merely the sum of many individuals. Many authors
have employed the term 'turf for this. The terms is not entirely satisfactory but it is at least
evocative. In northern parts of the British Isles, exposed shore species (principally C.
sepositum) sometimes grow so densely as to produce what has been equally inaccurately
described as 'spongy turf.

The genus Callithamnion Lyngbye

Lyngbye, Tent. Hydr. Dan.: 123 (1819).

Phlebothamnion Kutzing, Phyc. Gen.: 374 (1843).

Poecilothamnion Nageli in Neue Denkschr. Allg. schweiz. Ges. ges. Naturw. 9 (un-nod. Art.

2): 202 (1847).
Leptothamnion Kutzing, Sp. Alg.: 896 (1849).

114 P. S. DIXON & J. H. PRICE

Dorythamnion Nageli in Sber. bayer. Akad. Wiss. 1861 (2) : 344 (1862).
Ceratothamnion J. Agardh in Acta Univ. lund. 28 (6) : 35 (1892).
Aglaothamnion Feldmann-Mazoyer, Rec. Ceram. Med. Occident.: 451 (1941).

Thallus of variable shape, colour and size; colour rose-pink to muddy yellow-brown, varying
with species, vigour and habitat in natural conditions, between specimens and within a single
specimen; size and shape vary with species, specimen and habitat conditions; filamentous,
axes uniseriate, to 400 mm in length, variably but always copiously branched; branching
monopodial, although some species are peripherally flabellate from pseudodichotomous
evection; sometimes corticate by down-growing filaments derived from cells of axis or basal
cells of primary laterals, less often from subsequent order laterals; cortical development and
extent variable according to species, specimen and habitat conditions, involving main axis
only, or all orders of major axes; adventitious branching from cortication in a few species,
most dense in exposed shore specimens where a densely interwoven continuous thick matrix
is formed over the cortication; axial widths over cortication reaching 5 mm; normal
branching alternate, spiral or distichous, or approaching dichotomy in aspect; irregularities
occur occasionally in all species, with some variation with reproductive state; characteristic
form of the peripheral laterals occurs in both determinate smaller apical or peripheral
laterals (commonly referred to as 'ramuli') and indeterminate higher order laterals (in most
species), where the latter levels represent the current apices of the branching system and
contribute to the form of the thallus periphery; active main apices (main axes and major
laterals) either buried within local peripheral lateral development or clearly emerging above
that development according to species; apical cells of peripheral laterals gradually- to
abruptly-domed cylinders, of variable length according to species, or cones; apical (or
subsequently sub-apical) unicellular hyaline hairs irregularly present in active (younger)
parts of frond in many species, hairs commonly and often suddenly deciduous; cells length-
to-breadth ratios variable within the range 16 : 1 to 1 : 1 for all parts of the vegetative thallus;
cells multinucleate with uninucleate immediate products of apical division, or uninucleate
throughout, according to species; chloroplasts irregular and/or discoid in young cells,
forming irregular masses or ribbons by division and coalescence when older; chromosome
numbers commonly variable in the range n = 28-33, with extremes to n = 90-100.

Gametangial thalli commonly dioecious, occasionally monoecious in many species,
habitually in one. Spermatangia arise either in separate cushions, formed on cells of
outermost peripheral laterals, one or more per cell, cushions sometimes coalescing; or in
secund rows on adaxial surfaces of peripheral laterals, spreading when luxuriant to sides and
sometimes abaxial surfaces, finally encircling more mature, usually basal, parts; vigorous
production of spermatangia may lead to additional development from cells of major
branches of all orders in lateral systems, as in Callithamnion corymbosum or C. hookeri.

Procarps formed normally just behind or near apices of active axes and major indeter-
minate laterals, but may occur on lower branches in thalli with more vigorous reproductive
development; early procarp development with paired support cells, exceptionally with only
one, formed laterally on bearing axis; carpogonial branch developing on one support cell or
on both support cells, four-celled, including carpogonium; cell arrangement straight,
U-shaped, zigzag, or any intermediate of these; carpogonium relatively small, 6*5-26 um in
breadth, but with trichogyne of very variable dimensions and shape, curved, ramified,
nodular, swollen, uneven-sided, or bulbous-tipped, 47-242 urn in length; trichogyne
degenerating after fertilization, the carpogonium transferring a derivative of the fertilization
nucleus by a connecting cell, or directly, to an auxiliary cell formed by division from the
original support cell; following nuclear transfer the auxiliary cell dividing to produce a
central cell and a foot cell, central cell developing two, less often three, gonimoblast initials,
all producing few-celled gonimoblast filaments; one or two lobes of the gonimocarp
developing on the ultimate cell of the gonimoblast, one only on the penultimate cell of the
gonimoblast and on all subsequent cells; gonimolobes rounded, conical or uneven, varying
according to species, environmental factors and state of maturity; carposporophyte very

CALLITHAMN10N IN THE BRITISH ISLES 1 1 5

rarely sheathed by enveloping vegetative filaments; carposporangia of variable dimensions at
maturity, commonly 25-70 um in diameter.

Tetrasporangia ellipsoid or ovoid, less often subspherical to spherical, variable in size and
shape to 130 x 120 um, formed adaxially on cells of peripheral laterals, usually one or two
per cell; bisporangia usually exclusively on particular plants (Callithamnion byssoides) but
sometimes intermixed with tetrasporangia in C. byssoides and C. hookeri; parasporangia
reported elsewhere, not often authenticated in Britain, except in C. decomposition.

General key to British species

Price (1978) has correctly indicated that ecology should be the initial approach to a
utilitarian key aimed at field use with fresh material. Here, with all the complications of
older herbarium or preserved material and sparse ecological data, this is not practicable.
Nevertheless in the present key an attempt is made to integrate thallus or population
characteristics likely to have been noted or obvious from morphological state even in
individuals out of their normal environment. Use is also made of some data of reasonable
reliability derived from customary habitat preferences. All these aspects have been utilized
by past authors but an integrated key which does not place too much dependence on any one
criterion has not been attempted previously. Keys specific to ecology and to reproductive
states are being published elsewhere (Price, 1978; Price, in preparation).

1 Laterals of limited growth nor overtopping bearing axis 2

Laterals of limited growth overtopping the axis on which they arise 3

2 Cells uninucleate; cortication not commonly (although potentially) giving rise to an inter-

woven matrix of adventitious laterals; thallus almost never coarse, shaggy, or rope-like
C. hookeri (p. 122)

- Cells largely multinucleate; interwoven matrix of adventitious laterals arising from

cortication in at least the lower 7/8ths of the thallus; thallus always coarse, shaggy, and
rope-like in mature specimens, although less so if sublittoral . . C. tetricum(p. 128)

3 Lateral rudiments arising from the first product of a principal apical cell 4

Lateral rudiments arising on or after the second product of a principal apical cell ... 8

4 Apical cells of limited growth peripheral laterals rounded terminally into a dome, sometimes

strongly tapered 5

- Apical cells of limited growth peripheral laterals very small (12 3 urn long), conical

C. tetragonum (p. 127)

5 Cell lengths not exceeding 5 x breadth in any part of the thallus; ramification below lateral

branch system level strictly pseudodichotomous or not at all so, never mixed ... 6

- Cell lengths exceeding 5 x breadth in older axes and major laterals; ramification below

lateral branch system level strictly pseudodichotomous ... C. corymbosum (p. 1 1 8)

6 Cells uninucleate; interwoven matrix of adventitious laterals from cortication usually absent

C. hookeri (p. 122)

Cell multinucleate; interwoven matrix of adventitious laterals from cortication develops
over at least most of main axes and principal laterals

7 Apical clusters of lateral systems > 700 urn across; peripheral laterals (containing more than

7 cells) with their greatest width well above their base or parallel-sided for much of their
length; ramification pinnate; common on exposed shores in the sector from south Ireland

and south Wales northwards round to Yorkshire C. sepositum (p. 124)

Apical clusters of lateral systems < 700 /^m across; peripheral laterals (containing fewer
than 6 cells) with greatest width at or near their base; ramification pseudodichotomous;
either rarely recorded anywhere on exposed shores or rare in the north of the British Isles 9

8 Cells multinucleate; cells near apices > 30 urn diameter . . . C. roseum(p. 123)
Cells uninucleate; cells near apices < 20 urn diameter . . . . C. byssoides (p. 1 1 6)

9 Peripheral laterals with more than three cells above last ramification present in most lateral

systems; common on exposed shores in the southern half of the British Isles

C. granulatum (p. 120)

Peripheral laterals with never more than 2 cells, most usually only one cell, per lateral

116 P. S. DIXON & J. H. PRICE

above last ramification anywhere in the periphery, detailed peripheral branching pattern
'aping' C. corymbosum', probably widespread on exposed shores but sparse and rarely
authentically recorded C. spongiosm(p. 126)

Note: C. decompositum (p. 1 19) is omitted from this key, although described as fully as possible below.
Comparative information does not permit effective keying with so little material available. The
presence of parasporangia and strictly distichous apical arrangement indicate that confusion is
possible only with C. hookeri or Compsothamnion. In the forms ofCallithamnion hookeri in which the
indeterminate apices are not or only just overtopped by determinate laterals, there is commonly
discernible variation somewhere on the plant from the distichy by which many indeterminate branch
system apices may resemble those of C. decompositum. The tetrasporangia in C. decompositum are a
mixture of sessile (predominantly) and pedicellate (very occasionally), differing from the consistently
terminal tetrasporangia on pedicellate laterals in Compsothamnion. The latter is also more delicate and
less luxuriantly branched than Callithamnion decomposition.

Accepted species

1. Callithamnion byssoides Arnott ex Harvey in Hooker, Eng. Fl. Sir J. E. Smith,
5:342(1833).

Callithamnion arnottii Trevisan, Nomen. Alg. ... 1 : 77 (1845)
Callithamnion furcellariae J. Agardh, Sp. Gen. Ord. Alg. 2 (1) : 37 (1851).
Callithamnion hiemale Kjellman ex Kylin, Stud. Alg. schwed. Westk.: 170 (1907).

The accepted binomial, Callithamnion byssoides, may by antedated by two binomials. Callithamnion
arachnoideum C. Agardh, Sp. Alg. 2(1): 181 (1828), was described from the American coast of the
north Atlantic. Although some American material resembles closely the specimens from the eastern
shore of the north Atlantic, further detailed study is needed before conspecificity is accepted without
question. Ceramium tenuissimum Bonnemaison in Mem. Mus. Hist. nat. Paris 16 : 132 (1828), was
described from European material, but exact typification of this binomial is not possible (see p. 1 36).

Thallus of wide colour range, from pink to brick red; to 70 (-120) mm in height, bushy,
flaccid, rarely with easily recognizable main axis, although usually only one exists, less often
several more or less equivalent major axial branches diverge just above attachment; main
axis to 120(-180)um in breadth, commonly ecorticate throughout, less often with a few
basal, down-growing, corticating filaments from lower branches; axial cells narrowly
elongate, length-to-breadth ratios commonly 5 (or more) to 1 with detected range 3'00-13'19
to 1; branching alternate, monopodial throughout, with laterals arising irregularly in
direction and number; laterals of variable length in lower thallus, of more even length in
upper; uppermost larger laterals and peripheral laterals sparsely or not branched, long,
flaccid, flagelliform; insertion appearing distichous in lower regions, but becoming clearly
spiral towards apices of branching system; peripheral laterals usually unbranched, borne in
proximal 3/4 of bearing penultimate laterals but forming small corymbose or uneven clusters
near apices in distal 1/4 of bearing laterals, commonly straight, rarely incurved; apical
development in lateral systems not pseudodichotomous, but apical cell surpassed by
peripheral laterals; cells of laterals cylindrical, of similar length on the same lateral; apical
cells of peripheral laterals elongate domed cylinders, 52-59 17-17um in length,
12-2 3-33 um in basal breadth and 5*49 2-24 urn in apical breadth, with length-to-
breadth ratio of (2-5-) 3'5-5'5 to 1; peripheral laterals contain (1-) 2-5 (-7) cells, 5 1-430 urn
in length, 7-26 um in greatest breadth, with position of maximum breadth usually above
base.

Gametangial thalli dioecious. Spermatangiophores once-branched, sometimes with
rudiments of second lateral, often several on each cell of peripheral laterals; spermatangio-
phore branches 4 to 5 cells in length; 1-5 spermatangia on each cell, 6x8 um, but variable in
size.

Procarps formed behind apices of major axes and branches, also spreading towards the
base in very fertile material, with two support cells but sometimes only one; carpogonial

CALL1THAMNION IN THE BRITISH ISLES 1 1 7

branches zigzag or straight, on each support cell, but fertilization usually only unilateral;
two gominoblast initials from each auxiliary cell, first-developed producing two-celled
primary gonimoblast, second-developed forming one-celled primary gonimoblast, two
gonimolobes developing from the apical cell of first-developed primary gonimoblast, with
one gonimolobe forming from the subapical cell of first-developed and second-developed
primary gonimoblast; gonimolobes not lobed, narrow to pointed when young, becoming
rounded when mature, carposporangia 20-35 (-40) um in length.

Sporangial thalli may bear tetrasporangia or bisporangia (cf. Kylin, 1907; Rosenvinge,
1924), but usually not both, although this was reported by Harris (1959, 1962); bisporangia
ellipsoid, 62-4 5*7 x 33*4 5*6 jim, with transverse division; tetrasporangia sub-ellipsoid
to sub-spheroid, 55*2 4'6x40'9 7*4 um, with tetrahedral spore arrangement; both
usually sessile, tetrasporangia occasionally with 1 -celled pedicel, borne adaxially on
peripheral laterals of higher order but not prolifically on the smaller laterals of subsidiary
axes, usually 1 (-2) per cell.

Thalli with tetrasporangia or gametangia from March to November in the British Isles.
The presence of tetrasporangia on gametangial plants (Feldmann-Mazoyer, 1941; Foldvik,
1963; comments in Rueness & Rueness, 1980) or of bisporangia amongst gametangia
(Rosenvinge, 1 924) has not been detected in British field material so far.

Ecology and distribution*

Occurs in pools or damp sheltered areas through much of the littoral, extending to the
sublittoral; epiphytic, commonly on Fucus, Saccorhiza, Laminaria and larger Rhodophyta.

BRITISH ISLES: Channel Islands, Cornwall, Devon, Dorset, Isle of Wight, Sussex, Kent, Norfolk,
Yorkshire, Cheshire, Isle of Man; Caernarvonshire, Angelsey; Ayr, Bute, Argyll, Orkney, Aberdeen;
Dublin, Cork, Galway, Mayo, Down.

WORLD: Eastern Atlantic from western Norway to Canaries and Morocco; western Mediterranean;
western Atlantic from southern Massachusetts to Florida; Texas, West Indies; reported for New
Zealand and Tasmania.

Observations

Callithamnion byssoides has been confused with C. corymbosum and C. roseum, but the
species has a consistent ecological and geographical distribution. Critical cases are few in the
British Isles, but this is not necessarily true for the western part of the north Atlantic. C.
byssoides and C. corymbosum are occasionally confusable where the former lacks bispor-
angia. C. byssoides usually lacks the apical pseudodichotomy at unlimited functional apices,
the pseudodichotomies at peripheral lateral apices, and the hyaline hairs typical of C.
corymbosum. C. corymbosum has much coarser growth and all cells are multinucleate. C.
byssoides is finer than C. roseum, another multinucleate species. Mature cells near the
functional apices of C. byssoides are less than 20 (im maximum breadth, whereas in C.
roseum they exceed 30 um. The often greater length of cells of C. byssoides emphasizes
this distinction.

The sequence of morphological phases in the life history has been established for material
identified as Callithamnion byssoides from Port Aransas, Texas (Edwards, 1969) and from
Wrightsville Beach, North Carolina (Kapraun, 1978). These materials may not be con-
specific with British C. byssoides, a point already correctly made by Rueness & Rueness
(1980). Chromosome counts of n = 28-33 were reported in thalli from an undefined location
in the British Isles (Harris, 1962). The recent study (Rueness & Rueness, 1980) on
Norwegian material of C. byssoides established both a Polysiphonia-type life-history for the

*For this and all subsequent species, distribution is stated on the basis of the old county boundaries. The
newly-formed counties are both too large in area (particularly in Wales and Scotland) for effective discussion, and
not yet well enough comprehended in geographical terms.

118 P. S. DIXON & J. H. PRICE

European taxon and an interesting mode of perennation. It is curious that although the alga
is normally epiphytic (so reported by most authors, including this study), the perennating
creeping filaments that subsequently gave rise to C. byssoides plants were borne on the
ascidian Ciona intestinalis. Further field studies of this phenomenon elsewhere are
required.

2. Callithamnion corymbosum (Smith) Lyngbye, Tent. Hydr. Dan. : 125 (1819).

Conferva corymbosa Smith in Smith & Sowerby, Engl. Bot.: pi. 2352 (1811).
Ceramium versicolorC. Agardh, Syst. Alg.: 140 (1824).

Thallus pinkish-red, or brownish to deep red in colour; to 55 mm in height; bushy, flaccid,
usually with a single recognizable main axis; branching commonly pseudodichotomous
throughout, but with all gradations from m@nopodial to complete pseudodichotomy;
principal laterals of unlimited growth irregular in position and number; growth form
irregular to more or less regular; main axes 40-270 (-300) um in breadth; commonly
corticate but sometimes ecorticate"; occasional adventitious branching from the corticating
filaments, but lacking formation of adventitious matrix; axial cells in straight lines, some
curvature but no zigzags; branching often straggly and inconsistent in older parts towards
base, younger branches show spiral pseudodichotomous arrangement, divergence angles
usually about 30, rarely more; peripheral laterals borne toward apices on axes of various
orders, simple and spiral, or pseudodichotomous; apical development pseudodichotomous
in peripheral laterals axes; apical cells of axes surpassed by peripheral laterals; apical cells of
peripheral laterals 5-1 um in diameter, axial apical cells often smaller; hairs apical or
subapical, often present on all apical cells but equally often absent; cells at base of thallus
170-250 um in breadth, 25-35 um in mature cells near apices; length-to-breadth ratios of
cells 6- 1 2 to 1 ; cells multinucleate, except for a few cells near apices.

Gametangial thalli dioecious. Spermatangiophores multi-branched, one to two cells in
length, often several on each cell of peripheral or high order laterals; 2 to 4 spermatangia on
ultimate cells of Spermatangiophores; spermatangia 5-8 um.

Procarps normally formed just behind apices of principal axes and branches; usually 2

4

support cells, occasionally only one; carpogonial branch usually in form 4

less often completely straight; two one-celled gonimoblasts from each central cell,
commonly two gonimolobes on primary gonimoblast, only one on secondary gonimoblast,
but one or more gonimolobes occasionally missing; gonimolobes rounded, usually finally
lacking subsidiary lobing; carposporangia 25-35 um in diameter.

Tetrasporangia globose, 45-60 x 55-65 um, sessile, one or more on each cell of peripheral
laterals; often appearing to be axillary to pseudodichotomies.

The growth form of Callithamnion corymbosum can be very variable. Thalli may possess
either distinct axes bearing spirally arranged branches and simple peripheral laterals or with
branches and peripheral laterals developing at same rate as principal axes, producing a
pseudodichotomous aspect. Pseudodichotomies may occur throughout a plant, but are most
common only in the upper parts. The number of hairs is variable; some thalli have hairs on
almost all apical cells, others a few scattered hairs, some totally devoid of them. Harris (1959)
noted in Galway that plants of greater shelter had a few spirally-arranged hairs in the upper
parts while thalli of greater exposure showed pseudodichotomies and hairs on nearly all
apical cells. This is not a difference that is consistent in such conditions (see Price, 1978).

Thalli with gametangia reported between April and November; tetrasporangia occur
throughout the year.

Ecology and distribution

Usually epiphytic, on C odium fragile, Fucus vesiculosus, or Zostera; occasionally epilithic
in lower littoral, usually in standing water, pools and channels; known in the sublittoral to at
least 22 m in sheltered bays.

CALLITHAMNION IN THE BRITISH ISLES 1 1 9

BRITISH ISLES: Channel Islands, Cornwall, Devon, Dorset, Hampshire, Isle of Wight, Sussex, Somerset,
Cheshire, Isle of Man, Suffolk, Northumberland; Pembroke, Cardigan, Merioneth, Caernarvonshire,
Anglesey; Ayr, Bute, Argyll, Ross & Cromarty, Orkney, Aberdeen, Fife, Midlothian; Dublin, Wexford,
Cork, Galway, Mayo, Down.

WORLD: Eastern Atlantic, Norway (Nordland) to Canaries; western Baltic, Mediterranean and Black
Sea; Western Atlantic, Newfoundland to New Jersey; West Indies; Australia; Japan.

Observations

Callithamnion corymbosum has been confused with C. byssoides and C. roseum.
Pseudodichotomies at apices are absent in C. roseum and the hairs characteristic of
C. corymbosum are also lacking in that species. These features are also absent in C. byssoides
but C. corymbosum has much coarser growth and multinucleate cells.

The sequence of phases in the life history of Callithamnion corymbosum was established
in culture for material from Helgoland (Hassinger-Huizinga, 1952). In the present study, a
few plants have been observed with both tetrasporangia and carposporophytes and some
with tetrasporangia and alleged bisporangia. Hassinger-Huizinga reported the occurrence in
Helgoland material of thalli with tetrasporangia and either spermatangia or procarps in her
cultural studies. She claimed that the expression of gametangia and tetrasporangia is
governed by the addition or deletion of a chromosome. These data must be regarded with
caution since neither figures nor photographs of the chromosome complements are given
and the results necessitate absolute accuracy in counting chromosomes with a complement
of about 60 in a nucleus 5 urri in diameter.

In Newfoundland, Callithamnion corymbosum occurs as tetrasporangial thalli which
appear to reproduce only by fragmentation (Whittick, 1978).

Chromosome counts in Callithamnion corymbosum have been reported in material from
various locations as follows: Helgoland: = 30-32; 2n = 60-64 (Hassinger-Huizinga, 1952),
British Isles: n = 27-29 (Harris, 1959), British Isles: n = 28-33 (Harrris, 1962), British Isles:
n = c. 28 (Harris, 1966), Newfoundland: 2n = 57-65 (Whittick, 1973).

3. Callithamnion decompositum J. Agardh, Sp. Gen. Ord. Alg. 2 (1) : 45 (185 1).

Thallus light red in colour; somewhat rounded pyramidal in outline (see Price, 1978: 283, fig.
1 5b); rarely attaining more than 20 mm in length; main axis little corticate in British
specimens, although dense cortication reported elsewhere; branching usually strictly
distichous, commonly commencing at the second cell of major axes, branches restricted to
the adaxial side of the lower two cells of primary laterals giving a slightly secund appearance
in a few specimens; peripheral laterals attenuate and much ramified, causing flabellate
spreading distal effect that rounds off the pyramidal outline; hyaline hair points to 60 jim in
length reported in southern collections, but not noted in northern British material; cells of
main axes relatively large, 130-550 urn in length.,70-200 urn in breadth in mature material;
chloroplasts said to be polygonal-discoid when young, becoming elongate in older cells (in
British material the variation between variable discoid and elongate does not appear to be
correlated with age); masses of proteinaceous material present.

Gametangial thalli unknown for the British Isles; in non-British material dioecious;
spermatangiophores 2-4 celled, occurring on the first or second cells of laterals, each cell of
the spermatangiophore bearing 3 or 4 spermatangia; spermatangia c. 5 x 8 urn.

Procarps formed behind apices of lateral axes, commonly with two support cells;
carpogonial branch 'U '-shaped, trichogyne to 9 5 urn in length; gonimolobes cordate,
41-70 x 35^7 um; carposporangia 10x12 urn in diameter.

Tetrasporangia uncommon in British material; rounded or ovoid, to 55x65 jam in
diameter, usually sessile, occasionally pedicellate, occurring adaxially on peripheral laterals;
spore arrangement usually tetrahedral, occasionally irregular.

120 P. S. DIXON & J. H. PRICE

Parasporangia the most usual reproductive structure formed in British material, lobed or
irregular, sometimes linear, depending on state of development, 22-56 x 1 1-22 urn,
usually at the apices of adaxial products of peripheral lateral cells; individual paraspores
9-1 Sum.

The form range of Callithamnion decompositum is not fully elucidated, either in the
British Isles or elsewhere (Price, in preparation). This is partly due to uncertainties regarding
the status of material referred by other workers to C. bipinnatum, C. tripinnatum, C.
gallicum or C. caudatum. For a recent study of C. bipinnatum, see Rueness & Rueness
(1980).

Ecology and distribution

Epiphytic on other algae, but also epizoic and epilithic; usually sublittoral, to 30 m, but also
reported elsewhere in the lower littoral. Usually rather few thalli present.

BRITISH ISLES: Known only for Argyll (Mull), Inverness (Rhum) and Zetland, but probably occurs more
widely.

WORLD: Eastern Atlantic from the Faeroes to Senegal. The species may also occur in Denmark and
Norway, for which areas the relationships with material referred to Callithamnion bipinnatum require
elucidation. Recent work on the latter species by Rueness & Rueness (1980) suggests that, whatever the
application by some authors of that name, there is a good taxon concealed amongst the doubtful
applications.

Observations

Callithamnion decompositum may be confused with C. hookeri in the lower littoral and
shallow sublittoral, where both species have similar appearance and dimensions. The apical
organization of the two species is, however, usually quite distinct with virtually absolute
distichy in C. decompositum and some spiral arrangement of the ultimate laterals
somewhere on the plant in C. hookeri. Below the shallow sublittoral, C. hookeri becomes
more lax and less branched, whereas C. decompositum retains its characteristic growth-form.

Although parasporangia are the most reproductive structures found in British material,
there is insufficient information to comment on their phenology.

Nothing is known of the life history of Callithamnion decompositum. These northern
populations are parasporangial, although gametangia and/or tetrasporangia are produced by
material from areas south of the British Isles. In Finistere, parasporangia have been reported
mixed with tetrasporangia. Data on the life history in northern material will, it is hoped, be
more fully available when the Faeroe Islands specimens at present in culture at the
University of Oslo (J. and M. Rueness) have developed beyond vegetative growth from the
originally-released tetrasporangia.

4. Callithamnion granulatum (Ducluzeau) C. Agardh, Sp. Alg. 2 (1) : 1 77 (1 828).

Ceramium granulatum Ducluzeau, Essai Hist. Nat. Conf. Montpellier. 72 (1 806).
Callithamnion harveyanum J. Agardh in Linnaea 15 : 45 (1841).
Callithamnion grande J. Agardh, Alg. Mar. Med. Adr.: 73 ( 1 842).

Thallus pale straw, brownish, rust, to purple-red or dark red in colour; to 150 mm in height;
robust but highly flexible, flaccid in water; spongy in texture and appearance, but less so than
in C. sepositum; appearing as regular or irregular, simple or compound pine-tree like
pyramids; main axes rarely bifurcate near base, to 3 mm or more in breadth, but probably
greater in larger specimens; strongly corticate throughout; cortication develops dense matrix
of short tightly-woven adventitious laterals, giving rise occasionally to peripheral laterals of
normal form; main axial cells almost invisible without dissection; axial cells in straight lines,
length-to-breadth ratios of any visible upper axial cells usually 1-5 to 3 : 1; branching
monopodial, associated with pseudo-dichotomy at all levels and especially obvious at

CALLITHAMNION IN THE BRITISH ISLES 121

peripheries of thallus; divergence angles greater than 45, sometimes 90, unless strongly
appressed in corymbose clusters; apical cells of indeterminate laterals buried in the
corymbose clusters of peripheral laterals, normally needing dissection for examination;
peripheral laterals often spreading, but also frequently grouped in corymbose clusters, that
represent individual lateral systems, to 700 um in breadth (Price, 1978 : Fig. 11A-C);
individual peripheral laterals often straight although occasionally curved in the corymbose
clusters; greatest width of peripheral lateral at or near base; cells (1-) 2-4 (-6) per peripheral
lateral, usually more or less rectangular, with some apical narrowing; peripheral lateral
apical cell a short cylinder to roundly-truncated cone, 17'6 6*6 um in length; with
occasional terminal or subterminal hyaline hairs.

Gametangial thalli dioecious or (less often) monoecious. Spermatangial groups hemi-
spherical, rarely coalescent; spermatangiophores multibranched, branches 3-5 cells long;
two to four spermatangia on peripheral cell of each branch; spermatangia ovate to ellipsoid,
6x8 um.

Procarps formed just behind functional apices of penultimate and lower order laterals;
usually two support cells, but occasionally one; carpogonial branch form difficult to
interpret, but straight, with minor variations, and zigzag (the more usual) noted; carpo-
gonial maximum breadth 1 8 urn; trichogyne to 70-240 um in length; gonimoblasts single-
celled, two from each central cell; usually one gonimoblast giving rise to one gonimolobe,
occasionally gonimolobes missing; gonimolobes rounded without subsidiary lobing: carpo-
sporangia 25-35 um in diameter.

Tetrasporangia initially apical on cell below peripheral dichotomies, spreading to cells of
peripheral laterals, large, globose to ellipsoid, 45-60 x 50-60 um, never more than one per
bearing cell; polysporangia reported but never observed in the present study.

Detailed consideration of some aspects of growth form variation in Callithamnion
granulatum is given in Price ( 1 978).

Reproductive phenology in Callithamnion granulatum is variable with location, year and
individual thalli, although patterns can be perceived. Detailed information is being
published elsewhere.

Ecology and distribution

Occurs in rock pools and on emersed damp surfaces in the lower littoral of wave-exposed
shores; commonly epiphytic, very rarely epilithic, frequently epizoic on Mytilus, Patella and
Balanus, especially in blanket populations; reported from the sublittoral, but all such reports
erroneous except in very shallow conditions (to 0'5 m).

BRITISH ISLES: Channel Islands, Cornwall, Devon, Dorset, Isle of Wight, Yorkshire, Isle of Man;
Pembroke, Caernarvonshire, Anglesey; Ayr, Bute, Argyll, Inverness, Ross & Cromarty, Orkney,
Zetland, Angus, Fife, Midlothian, East Lothian; Antrim, Mayo, Galway, Clare, Cork, Wexford,
Wicklow, Dublin, Down.

WORLD: North-eastern Atlantic from Shetlands to Morocco, perhaps to the Gulf of Guinea, locally
(Liberia, Cote d'lvoire, Ghana); Mediterranean. Material reported from the Faeroes all appears
referable to the exposed-water form of Callithamnion hookeri, growing epiphytically on C. sepositum.

Observations

The only possible confusions are with Callithamnion sepositum or C. spongiosum. The
principal criterion for discrimination is the most usual number of cells in the highest order
peripheral laterals, with 1 or 2 cells in C. spongiosum, 3 to 4 in C. granulatum and 7 to 9 in
C. sepositum. C. granulatum may also be distinguished from C. sepositum in its consistent
pseudodichotomy and the basal, or near basal, maximum width of the peripheral laterals.
The greatest difficulty is in discriminating from C. spongiosum, with which it tends to occur
in mixed populations although intermediate forms are lacking. Cotton (1912) distinguished

122 P. S. DIXON & J. H. PRICE

between the two species, although the ecological differences which he noted do not hold
consistently.

Little published information is available on the life history of Callithamnion granulatum\
all exposed shore forms are difficult to culture. Harris (1962) reported a chromosome
number of n = 28-33 in material identified by him as C. purpurascens and possibly referable
to this entity.

5. Callithamnion hookeri (Dillwyn) S. F. Gray, Nat. Arr. Br. PI. 1 : 324 (1821).

Conferva hookeri Dillwyn, Br. Conf.: pi. 106 (1809).
Callithamnion roseum /? tenue Lyngbye, Tent. Hydr. Dan.: 126(1819).
Ceramium scopulorum C. Agardh, Syst. Alg.: 132 (1824).
Callithamnion scopulorum C. Agardh, Sp. Alg. 2 (1) : 166 (1828).
Callithamnion polyspermum C. Agardh, Sp. Alg. 2 (1) : 169 (1828).
Callithamnion brodiaei Harvey in Hooker, Eng. Fl. SirJ. E. Smith 5 : 340 (1 833).
Callithamnion grevillei Harvey in Hooker, Eng. Fl. SirJ. E. Smith 5 : 345 (1833).
Callithamnion spinosum Harvey in Hooker, Eng. Fl. SirJ. E. Smith 5 : 345 (1833).
Callithamnion acrospermum J. Agardh, Sp. Gen. Ord. Alg. 2(1): 52(1851).

Thallus of widely different colours but usually dark to brownish red; to 70 mm or more in
height; bushy, straggly, or sparsely branched, sometimes even reduced to a single main axis;
main axis or axes 300-500 jim in breadth, commonly corticate, often with reduced
cortication, or even none in younger material; cortication often producing adventitious
laterals, when extreme forming a matrix that may be over 2 mm overall breadth; branching
very variable, from regular alternate-spiral to highly irregular; peripheral laterals distichous
or spiral, but not usually overtopping the apex of main axis, borne on penultimate laterals
with strong axial zigzag; apical cells rounded, basal breadth 5-1 Oum; cell numbers and
dimensions in peripheral laterals highly variable; cells of peripheral laterals cylindrical or
slightly tapered; mature cells near base ofthallus 50 to 100 um maximum diameter, reducing
to 40 to 80 um near apex, length-to-breadth ratios varying between 0'5 to 1 and 5 to 1 .

Gametangial thalli dioecious. Spermatangiophores one, rarely more, per cell of peripheral
laterals, variable in form even in same thallus; most Spermatangiophores consisting of a
single cell with a single secund branchlet, 3 to 4 cells in length, others with a series of secund
branchlets; ultimate cells of branchlets bearing 2 to 4 spermatangia, usually secund but
sometimes distichous; spermatangia 4x8 um.

Procarps formed just behind apices of axes and indeterminate laterals; carpogonial branch
of variable form, from straight to zigzag; usually developing two auxiliary cells after
fertilization, but occasionally only one; each auxiliary cell producing two single-celled
gonimoblasts, with one or two gonimolobes, although the second-developed filament may
produce only one or even none; gonimolobes lobed or simple, usually rounded, but
sometimes pointed or even conical; carposporangia at maturity 35-50 um in diameter.

Tetrasporangia globose or nearly so, sessile, 40-60 x 45-60 um, borne adaxially on
peripheral laterals, usually one per cell, but occasionally with a second tetrasporangium
borne abaxially. Bisporangia occur rarely, mixed with tetrasporangia, and may represent
only an aberrant or developmental stage of the latter. Parasporangia replacing vegetative
lateral filaments reported widely, but not authenticated in the British Isles. Monosporangia
reported, but probably nothing more than tetrasporangial initials.

The overall thallus form is highly variable under different environmental conditions and
in different locations. So far as can be determined, there are no consistent morphological
distinctions between Callithamnion hookeri and the entities usually treated under the names
C. scopulorum, C. brodiaei and C. polyspermum. The thalli which have been referred to C.
scopulorum occur in the upper littoral in damp locations as clumps or local turfs. The
customary epiphytic littoral populations tend to be lax whereas in situations exposed to
wave-action the thalli are robust, short, stubby and much branched. The sublittoral

CALLITHAMNION IN THE BRITISH ISLES 1 23

populations tend to be extremely lax. Analysis of this variation in respect to previous
nomenclature and taxonomy is given in Price ( 1978).

Reproductive phenology in Callithamnion hookeri is variable with location, year and
individual thalli, although patterns can be perceived. Detailed information is being
published elsewhere.

Ecology and distribution

Occurs in rock pools and on emersed damp surfaces in the lower littoral of wave-exposed
shores; commonly epiphytic, rather rarely epilithic, frequently epizoic on Mytilus, Patella
and Balanus.

BRITISH ISLES: Callithamnion hookeri is probably present throughout the British Isles, occurring under
appropriate circumstances. Positive identifications are, however, available only for the following
counties; Channel Islands, Cornwall, Devon, Dorset, Hampshire, Sussex, Kent, Essex, Suffolk,
Norfolk, Lincolnshire, Yorkshire, Durham, Northumberland, Somerset, Cheshire, Cumberland, Isle of
Man; Glamorgan, Pembroke, Cardigan, Caernarvonshire, Anglesey; Dumbarton, Wigtown, Ayr,
Bute, Renfrew, Argyll, Inverness, Ross & Cromarty, Orkney, Zetland, Sutherland, Moray, Aberdeen,
Kincardine, Angus, Fife, Midlothian, East Lothian, Berwick; Antrim, Londonderry, Donegal, Leitrim,
Mayo, Galway, Clare, Limerick, Kerry, Cork, Waterford, Wexford, Wicklow, Dublin, Down.

WORLD: Callithamnion hookeri is extremely widespread but a precise statement of its total distribution
is difficult in view of its great variability and the need to clarify the status of material referred to such
entities as C. bipinnatum, C. tripinnatum, C. pusillum, C. hirtellum, C. truncatum, etc. The species
occurs in the north Atlantic in Iceland and between northern Norway and the Canary Islands and
Morocco, occurring also (Price, in press) in areas of cold upwelling in the Gulf of Guinea
(and perhaps sparsely in Angola), on the Baltic Sea coast of Germany, and from Newfoundland (South
& Hooper, 1980) southward. The reported occurrence of various entities possibly referable to this
species in the Mediterranean must await taxonomic clarification; the same comment applies to reports
from the Pacific Ocean.

Observations

The life history of Callithamnion hookeri has been shown by culture studies of British
material to consist of a sequence of gametangial, carposporangial and tetrasporangial
phases (Edwards, 1979), although nothing is known of the chromosome counts. In
populations from the west coast of Sweden, it has been shown that paraspores develop into
parasporangium-bearing thalli for several successive generations (Rueness & Rueness, 1978).
The role of the alleged monosporangia has not been established. On the relations between
life-history, tolerance levels, and distribution pattern, see van den Hoek (in press).

6. Callithamnion roseum (Roth) Lyngbye, Tent. Hydr. Dan.: 126, tab. 39 (1819).

Ceramium roseum Roth, Archiv Bot. (Romer) 1 (3): 47 (1798).

Conferva rosea (Roth) Smith in Smith & Sowerby, Engl. Bot.: pi. 966 (1802).

Callithamnion octosporum C. Agardh, Sp. Alg. 2(1): 177 (1828).

Thallus pinkish to purple-red; to 80 or more mm in height; bushy and fairly robust, less
delicate or flouncy than Callithamnion byssoides or C. corymbosum, but still flaccid; usually
with one main axis; main axis to I'O mm in breadth, sometimes ecorticate but commonly
with at least a few cortical filaments from lower branches, occasionally moderately corticate
but no development of adventitious matrix from cortication; branching monopodial but
sometimes with pseudodichotomous aspect below carposporophyte; axial (indeterminate)
branches irregular in position and number, variable in length, especially at base of plant,
spirally arranged in upper parts; peripheral laterals usually simple, upsweeping and
incurving, spiral; apical cells of peripheral laterals 5-10 urn in breadth; apical cells of main
axis and main branches surpassed by subordinate peripheral laterals; cells at bases of plants
130-225 jim in breadth; tapering; mature cells, near apices, 30-35 um in breadth; length-to-
breadth ratio 5-12 to 1 but cells usually shorter, more robust, than C. byssoides; cells usually

124 P. S. DIXON & J. H. PRICE

multinucleate except immediately behind apices; chloroplasts irregularly discoid in young
cells, forming continuous ribbons with age.

Gametangial thalli dioecious. Spermatangiophores two cells in length, multibranched,
often several on each cell of peripheral laterals; two to four spermatangia on ultimate cells of
spermatangiophore cluster; spermatangia 5x8 urn.

Procarps formed behind apices of main axes and branches, commonly with two support
cells, rarely only one; carpogonial branch straight, occasionally zigzag; two gonimolobes
from each central cell, each one-celled; commonly two gonimolobes on first gonimoblast,
but only one on secondary gonimoblast; one or more gonimolobes occasionally missing;
gonimolobes usually lacking subsidiary lobing, rounded; carposporangia 20-35 um in
diameter.

Tetrasporangia globose, 40-50 x 45-55 um, sessile, often abundant, one or more, usually
adaxial, on each mature cell of peripheral laterals.

Spermatangial thalli of Callithamnion roseum tend to be more delicate than tetra-
sporangial or sterile specimens, but the degree of robustness varies enormously.

Fertile thalli appear to occur at all times of the year, but there is considerable variation in
the reproductive organs present according to individual thallus, year, time and location.

Ecology and distribution

Epiphytic on other algae in lower littoral in sheltered conditions or sheltered niches on more
wave-exposed shores; can extend into the sublittoral to at least 20 m. A characteristic habitat
is just below water level on floating structures in areas of somewhat fluctuating salinity.

BRITISH ISLES: Channel Islands, Cornwall, Devon, Hampshire, Sussex, Kent, Suffolk, Norfolk,
Lincolnshire, Isle of Man; Caernarvonshire, Anglesey; Cork, Galway, Wexford, Dublin. Probably
widespread.

WORLD: Eastern Atlantic from Norway to Portugal; Baltic; eastern coast of the United States.

Observations

Callithamnion roseum may be confused with C. corymbosum and C. byssoides, but lacks
pseudodichotomous apical development and the hairs which occur quite commonly in
British C. corymbosum. C. roseum has coarser growth than C. byssoides, with multinucleate
cells.

Tetrasporangia occur in all months of year in most areas of the British Isles. Gametangial
plants and carposporophytes detected April-November. Little published information exists
on the life history of Callithamnion roseum. Bisporangia or carposporophytes have been
reported on a few tetrasporangial plants. Chromosome number reported, doubtfully, as n = c.
39 (Harris, 1962, 1966), but no other data available.

7. Callithamnion sepositum (Gunnerus) P. Dixon & J. Price, comb. nov.

Conferva seposita Gunnerus, Fl. Norv.: 116(1 772).
Conferva purpurascens Hudson, Fl. Angl., 2nd ed. 2 : 600 (1 778).
Conferva arbuscula Dillwyn, Br. Conf: pi. 85 (1807).
Ellisius glaber S. F. Gray, Nat. Arr. Br. PI. 1:333(1821).
Phlebothamnionfaroense Kiitzing, Tab. Phyc. 14 : 30 (1864).
Phlebothamnion scoticum Kiitzing, Tab. Phyc. 14 : 30 (1864).

Thallus dark red to dark purple in colour, distally bleaching to straw-coloured; robust and
spongy but highly flexible, the texture resembling wet wool; main axes and laterals strongly
corticate, covered throughout by a matrix of short, adventitious laterals obscuring the axes;
axes from 310 um to more than 5 mm in breadth; branching alternate-pinnate throughout,
with occasional apparent aberrant pseudodichotomies; intermediate laterals arising in all
directions; peripheral laterals grouped into tightly-packed, incurved, corymbose clusters,

CALLITHAMNION IN THE BRITISH ISLES 1 25

700 to 1400 jim in breadth (Price, 1978 : Fig. 10A-C); apical cells truncated cones, length
frequently greater than breadth, length 20'5 5'0 Jim, basal breadth 16'604'2um,
apical breadth 5'50 2'5 um; cell numbers in peripheral laterals (2-) 7-9 (-16); peripheral
laterals 1 25-539 um in length x 24-60 um in breadth, position of the maximum breadth
usually well above the base or lower half parallel-sided; cells of peripheral laterals
cylindrical, the maximum breadth equal to the length.

Gametangial thalli monecious or dioecious, predominance apparently varying with
region. Spermatangial thalli with dense spermatangia within the apical corymbose clusters,
sometimes in localized areas, sometimes widespread throughout; initial stages adaxial along
incurved peripheral laterals, but subsequently spreading to form coalescent cushions along
the lengths of whole laterals.

Carpogonial thalli usually bearing abundant procarps; position of procarps variable,
above or below outer forkings on peripheral laterals in heavy-cropped specimens; two
support cells developed, frequently only one visible; carpogonial branch apparently
U-shaped but sometimes with possible zigzag; carpogonia 15-26 um in breadth, trichogyne
to 240 jim in length, always twisted, occasionally with swellings or bifurcations; carpo-
sporophytes large and conspicuous when fully grown, normally paired on opposing support
cells, occasionally unilateral; in luxuriant growth, carposporangia becoming angular through
lateral pressure; rounded-off carposporangia 70 x 70 urn.

Tetrasporangia sub-spherical to spherical, sessile; (42-) 59-77 (-110)x(37-) 55-70
(-100) um; adaxial, occasionally abaxial on cells of outer peripheral laterals.

There are correlations between the form of mature plants and reproductive state.
Tetrasporangial specimens show luxuriant clumped and bushy branching, while carpo-
sporangial thalli show sparser, more open, branching with longer laterals, which however do
not contain more cells. Spermatangial specimens are intermediate between these two.
Hyaline hair points can occur on any material but particularly on carpogonial plants.
During winter and early spring, thalli of Callithamnion sepositum are very straggly and tend
to lack the peripheral corymbose clusters of laterals.

Reproductive phenology in Callithamnion sepositum is variable with location, year and
individual thalli, but patterns can be perceived. Detailed information is being published
elsewhere.

Ecology and distribution

Commonly epilithic on steep slopes or epizoic on shells of Patella, Balanus and Mytilus; in
blanket populations, occasionally epiphytic on Corallina or on the bases of exposed Fucus
vesiculosus; very rarely in pools except in blanket populations.

BRITISH ISLES: Cornwall, Devon, Yorkshire, Durham, Northumberland, Isle of Man; Pembroke,
Caernarvonshire, Anglesey; Wigton, Ayr, Bute, Argyll, Inverness, Ross & Cromarty, Sutherland,
Caithness, Orkney, Zetland, Banff, Aberdeen, Angus, Fife, Midlothian, East Lothian, Berwick;
Waterford, Cork, Kerry, Clare, Galway, Mayo, Donegal.

WORLD: Norway, Faeroes, and Iceland.

Observations

Callithamnion sepositum is similar in size, appearance and growth in blanket populations
on the shore to C. granulatum, but mature thalli of C. sepositum lack the symmetrical
appearance of those of C. granulatum. See pp. 121-2; 126-7 where the differences are
discussed in detail.

Little published information is available on the life history of Callithamnion sepositum;
all exposed shore forms are difficult to culture. Harris (1962) reported a chromosome
number of n = 28-33 in C. purpurascens, possibly referable to this entity.

126 P. S. DIXON & J. H. PRICE

8. Callithamnion spongiosum Harvey in Hooker, Engl. Fl. SirJ. E. Smith 5 : 346 (1833).

Thallus through pale straw or brownish or rust, to purple-red or dark red in colour according
to season and environment; to 50 mm in height; robust but highly flexible, flaccid in water;
spongy in texture and appearance, but (like Callithamnion granulatum) less so than in C.
sepositum, appearing as regular or irregular, simple or compound pine-tree like pyramids;
main axes rarely bifurcate near base; to 3 - mm in breadth, but probably greater in larger
specimens; strongly corticate throughout; cortication developing dense matrix of short
tightly woven adventitious laterals, giving rise occasionally to peripheral laterals of normal
form; main axial cells almost invisible without dissection; axial cells in straight lines, length-
to-breadth ratios of any visible upper axial cells usually 1 '5 -3 to 1; branching monopodial,
associated with pseudodichotomy at all levels but especially obvious at periphery of thallus;
divergence angles greater than 45, sometimes 90; indeterminate apical cells buried in
corymbose clusters (representing individual lateral systems) of peripheral laterals, normally
needing dissection for examination; peripheral laterals often spreading, frequently grouped
in corymbose clusters to 700 jim in breadth; individual peripheral laterals usually straight
(although occasionally curved in corymbose clusters) consisting of one, less often two, cells,
greatest width at or near base; cells usually more or less cylindrical, with some apical
narrowing; apical cell short cylinder or rounded, 17*6 6'6 urn in length; occasional
terminal or subterminal hyaline hairs.

Gametangial thalli dioecious or monoecious. Spermatangial cluster hemispherical,
rarely coalescent; spermatangiophores multibranched, branches 3-5 cells long; two to four
spermatangia on peripheral cell of each branch; spermatangia ovate to ellipsoid, 6x8 urn.

Procarps formed just behind functional apices of penultimate and lower order laterals;
usually two support cells, but occasionally one; carpogonial branch form difficult to
interpret, but straight (with minor variations) and zigzag noted; carpogonial maximum
breadth 18um, trichogyne to 70-240 um in length; gonimoblasts single-celled, two from
each central cell; usually one gonimoblast giving rise to one gonimolobe, but occasionally
gonimolobes missing, mature gonimolobes rounded without subsidiary lobing; carpospor-
angia 25-35 um in diameter.

Tetrasporangia initially apical on cell below peripheral dichotomies, spreading to cells of
peripheral laterals, large globose to ellipsoid, 45-60 x 50-60 um, never more than one per
bearing cell; polysporangia reported but never observed in the present study.

So far as can be determined from the sparse material available, the growth form variation
in Callithamnion spongiosum is similar to that of C. granulatum, discussed on p. 120 et seq.

Callithamnion spongiosum occurs too sparsely for any meaningful comments to be made
on the reproductive phenology of this species.

Ecology and distribution

Occurs in rock pools and on emersed damp surfaces in the lower littoral, commonly
epiphytic, less frequently epizoic on Mytilus, Patella and Balanus. Very rarely in large
populations; usually mixed as sporadic thalli with Callithamnion granulatum.

BRITISH ISLES: Cornwall, Devon, Isle of Man; Pembroke, Caernarvonshire, Anglesey; Ayr, Bute;
Antrim, Dublin.

Other records (Sussex, Yorkshire, Fife, Zetland, Angus, Channel Islands) have proved on
re-examination to be erroneous, although there is no reason to doubt the sporadic occurrence of the
entity in western areas.

WORLD: No verifiable records are known, but on the basis of present knowledge the presence of this
species in the least north-west France is probable.

Observations

The only possible confusions are with Callithamnion sepositum or C. granulatum. The most
usual number of cells in the higher order peripheral laterals is the prime characteristic for

CALLITHAMNION W THE BRITISH ISLES 127

discrimination, with 1 or 2 cells in C. spongiosum, 3 to 4 in C. granulatum and 7 to 9 in C.
sepositum. C. spongiosum may also be distinguished from C. sepositum in its consistent
pseudodichotomy and the basal maximum width of the peripheral laterals. The greatest
difficulty is in discriminating from C. granulatum, with which it tends to occur in mixed
populations, although intermediate forms are lacking. Cotton (1912) distinguished between
the two species, although the ecological differences which he noted do not consistently hold.

It may be that the rarity of occurrence of specimens amongst Callithamnion granulatum
populations represents the detected manifestation of tendencies towards speciation in a
genome currently still interfertile with the C. granulatum from which it arises. If this is so,
the expected intermediates must be being dismissed as plants of C. granulatum. The
inconsistency and sparseness of appearance in particular C. granulatum populations may
indicate a present lack of success in establishment as a separate and defined breeding
population (eventually, species). It will be interesting to see if specimens of C. spongiosum
become more frequent in the field in the (imponderable) future, with some gradual emphasis
of ecological tolerance differences, at present not recognizable to us.

No published information is available on the life history of Callithamnion spongiosum; all
exposed shore forms are difficult to culture. Harris (1962) reported a chromosome number of
n = 28-33 in material referred by him to C. purpurascens, possibly referable to this entity.

9. Callithamnion tetragonum (Withering) S. F. Gray, Nat. An. Br. PI. 1 : 329 (1821).

Conferva tetragona Withering, An. Br. PL, 3rd ed. 4 : 405 (1 796).

Ceramiumfruticulosum Roth, Cat. Bot. 2 : 183 (1800).

Conferva fruticulosa Wulfen, Crypt. Aquat. : 26 (1803).

Ceramium brachiatum Bonnemaison in Mem. Mus. Hist. nat. Paris 16 : 136 (1828).

Ceramium dudresnayi Bonnemaison in Mem. Mus. Hist. nat. Paris 16 : 125 (1828).

Ceramium guttatum Bonnemaison in Mem. Mus. Hist. nat. Paris 16 : 127 (1828).

Callithamnion fruticulosum J. Agardh in Linnaea 15 : 46 (1841).

Thallus red, brownish-red to rust in colour; robust, coarser than all other species except
Callithamnion tetricum, irregularly pyramidal in outline, to 110mm in height; usually a
single main axis in a thallus, apparent basal division into several principal axes often
resulting from separate development of adjacent spores; branching regularly spiral,
occasionally irregular towards base of main axis, larger laterals similar to main axis; main
axis strongly corticate, 233 to 550 um (to 1 mm) in breadth; adventitious branching from
the cortication absent or sparse; axial cells in linear rows, rarely even distal axial zigzag;
length-to-breadth ratios of axial cells easily seen through cortication, usually less than 3 to 1 ;
cell breadth tapering from axial base to apex, from 200-300 to 90-1 10 um; apices surpassed
by clustered peripheral laterals, latter without the corymbose grouping that occurs in many
other exposed shore species; older lower order laterals easily confused with main axes when
detached; peripheral laterals incurved, often strongly so giving 'awl'-shape, occasionally
straighter, longer and narrower, appearing almost parallel-sided ('brachiatum'); cells of
peripheral laterals barrel-shaped ('tetragonum'') or cylindrical ('brachiatum''); apical cells of
peripheral laterals small in relation to products, more or less conical, with pointed apices,
12 3 um, in length, 10 4 um in breadth at base; cells below determinate lateral apices
truncated cones; breadth of immediate derivatives equal to or greater than length; cells
usually two or more times as long as apical cell; cells in peripheral laterals 1 50 to 700 um in
length, maximum breadth, basal to median in position, (11-) 40-165 um; hairs absent from
British material; cells multinucleate, except near apices; chloroplasts irregularly discoid,
occasionally becoming long thin ribbons or irregular masses.

Gametangial thalli usually monoecious, habitually so in some populations. Spermatangio-
phores many-branched, branches 3 to 5 cells long, forming unilateral adaxial tufts,
60 6um in length, 30 7 um in height, one per cell of peripheral laterals, borne near
apical septum, rarely coalescent; ultimate cells bearing two to four spermatangia per cell;
spermatangia 5x8 um.

128 P. S. DIXON & J. H. PRICE

Procarps formed behind indeterminate apices of peripheral branch systems, often basal to
clusters of peripheral laterals; normally two support cells, occasionally only one;
carpogonial branch of variable form, but usually straight; trichogyne shape and size variable,
with various swellings and usually divided distally, 50 to 200 ju,m in length; each auxiliary
cell producing two single-celled gonimoblasts and a single gonimolobe from each gonimo-
blast cell; mature gonimolobes rounded, not lobed; carposporangia variable in shape, due to
appression, 25-45 um in diameter.

Tetrasporangia spherical to ellipsoid, sessile, 60 8 x 52 7 um, borne adaxially on
peripheral laterals, rarely more than one per cell.

The overall thallus form of Callithamnion tetragonum is usually pyramidal and quite
characteristic, only rarely appearing straggly and irregular. The peripheral laterals exhibit
varying degrees of curvature and cell form, the extremes having been sufficiently distinct for
two species to have been recognized formerly, although now treated merely as forms. In
some thalli, the laterals are strongly incurved (tetragonum form; Price, 1978 : Fig. 13), whilst
in others they are rather straighter and much narrower (brachiatum form; Price, 1978 : Fig.
14A): mixtures of the two types are also known.

Reproductive phenology in Callithamnion tetragonum is variable with location, year and
individual thalli, but patterns can be perceived. Detailed information is being published
elsewhere.

Ecology and distribution

Occurs in rock pools or open damp conditions at all levels, except high littoral and supra-
littoral, but best developed in the lower littoral under conditions of considerable, but not
extreme, wave action, and in the sublittoral to 24 m; predominantly epiphytic, more rarely
epilithic. Usually more abundant and luxuriant from the shallow sublittoral, especially on
the laminae ofLaminaria spp.

BRITISH ISLES: Channel Islands, Cornwall, Devon, Somerset, Dorset, Hampshire, Isle of Wight, Sussex,
Kent, Yorkshire, Durham, Northumberland, Isle of Man; Glamorgan, Pembroke, Cardigan,
Caernarvonshire, Anglesey, Denbigh; Bute, Argyll, Ross & Cromarty, Orkney, Aberdeen, Angus, Fife,
East Lothian, Berwick; Derry, Down, Antrim, Dublin, Wexford, Waterford, Cork, Kerry, Clare,
Galway, Mayo.

WORLD: North-eastern Atlantic from Iceland and Norway to the Algarve; Mediterranean; eastern coast
of U.S.A. Recorded, possibly in error, from the Canaries, Cape Verde Islands, and the Cape of Good
Hope.

Observations

Callithamnion tetragonum is not easy to mistake; even so, there have been numerous
misidentifications.

Little information is available on the life history of Callithamnion tetragonum. Life
history of the American C. baileyi, considered by some to be conspecific with C. tetragonum,
has recently been elucidated by Whittick & West (1979). Chromosome counts n = 9-10,
2n= 18-20 (Mathias 1927, 1928, 1932, 1935) were shown to be erroneous by Westbrook
(1930a, 1935); n = 28-33 (Harris, 1962).

10. Callithamnion tetricum (Dillwyn) S. F. Gray, Nat. An. Br. PL 1 : 324 (1821).

Conferva tetrica Dillwyn, Br. Conf. : pi. 8 1 ( 1 806).

Ceramium congestum Bonnemaison in Mem. Mus. Hist. nat. Paris 16 : 125 (1828).

Thallus red to brown in colour, rarely purple; rope-like, coarse and straggly, to 250 mm in
height; main axes and major laterals densely corticate, cortication giving rise to very thick
matrix of adventitious laterals; main axes 2-4 mm in breadth; branching strictly mono-
podial, indeterminate branches irregular in position and development, resembling main

CALLITHAMNION IN THE BRITISH ISLES 1 29

axes; distal middle order laterals bearing stiff, shorter but progressively lengthening
peripheral laterals giving a 'conical' (narrow pyramidal) outline to the individual branching
system, the lower few cells of these distal and larger peripheral laterals occasionally bare;
higher (later) order laterals distichous, angle of insertion acute, 50-60; apical cells of bearing
indeterminate laterals never over-topped by peripheral laterals (see Price, 1978 : Fig. 12);
apical cells 125-175 urn near base of thallus, 75-100 um behind distal apices of thallus; cell
length-to-breadth ratios 1-3 (-5) to 1 in apical parts, (1-) 3-5 to 1 in basal parts of thallus;
cells lengths and breadths increasing rapidly for about 4 to 6 cells behind the apical cell and
immediate derivatives; chloroplasts irregular, discoid, or ribbon-shaped.

Gametangial phase dioecious. Spermatangiophores branched, branches 3-5 celled, borne
on every cell of peripheral lateral, in luxuriant development forming large coalescent masses;
2-4 spermatangia from ultimate cells of Spermatangiophores; spermatangia 4-8 urn

Procarps formed behind apices, especially of peripheral laterals; usually two bilateral
support cells, occasionally only one; carpogonial branch straight, less often zigzag; each
central cell developing two 1 -celled gonimoblast filaments which bear 1 gonimolobe each;
mature gonimolobes with no subsidiary lobing; carposporangia 8-12 um in diameter.

Tetrasporangia globose, 65-90 x 75-90 um, sessile, adaxial on peripheral laterals, one per
cell; bi- and tri-sporangia reported (Westbrook, 1930).

Habit varies little. Specimens more straggly over the winter and branching sparser and
more open in sublittoral material. Perennial at least at times in Britain and probably
throughout range. Branching pattern correlates with reproductive state, but seasonal effects
probably equally as great.

Reproductive phenology in Callithamnion tetricum is variable with location, year and
individual thalli, but patterns can be perceived. Detailed information is being published
elsewhere.

Ecology and distribution

Usually epilithic or less commonly epiphytic on shores exposed to strong wave-beat, but
never on the fully exposed positions that encourage especially Callithamnion sepositum and
to a lesser extent C. granulatum on very exposed shores; C. tetricum occurs only in shore
conditions offering some protection from direct wave impact (see Price, 1978).

BRITISH ISLES: Channel Islands, Cornwall, Devon, Dorset, Hampshire (Isle of Wight), Kent, Norfolk
(doubtful), Isle of Man; Glamorgan, Pembroke, Anglesey; Down, Dublin, Wexford, Waterford, Cork,
Clare, Galway.

WORLD: Known from north Ireland to Morocco.

Observations

Callithamnion tetricum, wherever it occurs, is one of the most distinctive species of the genus
and has been involved in few confusions.

Little is known of the life history of Callithamnion tetricum. Reported chromosome
numbers show curious disagreements. Westbrook (1930/7) reported ?n = c.25 and ?2n = c.50,
while Harris (1959) reported counts of 92, 93, 94 and 98 in sporangia, later cited (Harris,
1962) as '790-100'. The latter counts are three times greater than those obtained in any
other species of Callithamnion and approximately twice the number reported by Westbrook.

Rejected species
1. Rejections from the British flora

Two species of Callithamnion accepted in the British flora for many years must be rejected,
either on the grounds of taxonomic uncertainty or because of misidentifications. These are:

130 P. S. DIXON&J. H. PRICE

Callithamnion rabenhorstii (Kiitzing) P. Crouan & H. Crouan.

Described originally from the eastern Mediterranean (Ionian Sea) by Kiitzing (1849 : 896),
the species was added to the British flora on the basis of material collected by E. M. Holmes
at Studland (Dorset). The status of the entity is questionable, even at its original location, and
there is no valid reason to accept the species in the British flora. Harris (1966) has suggested
that the Holmes material resembles C. byssoides, and this attribution appears to be correct.

Callithamnion tripinnatum C. Agardh

This entity has been accepted in the British flora for many years, although it has been the
subject of much confusion. The initial record for the British Isles, by Harvey (in Hooker,
1833 : 346), is based on material of Compsothamnion sp. Subsequently, Harvey (1847, pi.
LXXVII) corrected this initial error, although the plants to which the name was then applied
are referable to Callithamnion hookeri. Whatever the status of the original material of C.
tripinnatum might be, as stated on p. 123 it would seem to form part of the C. hookeri
assemblage; there is no valid reason for accepting C. tripinnatum in the British flora.

2. Entities of uncertain attribution

Several species, attributable to the genus Callithamnion and described from the shores of the
British Isles and adjacent regions, cannot be referred with certainty to any of the previously
mentioned species. There are two principal reasons for this uncertainty:

1 . The type material cannot now be located, if it ever was preserved.

2. The type material has been located, but is inadequate in quality or quantity for
determination.

The entities of uncertain attribution are:

Callithamnion arachnoideum Griffiths ex Moore in Larcom, Ord. Surv. Londonderry 1:11
(1837).
Callithamnion bipinnatum P. Crouan & H. Crouan, Fl. Finistere : 136 (1867).

See comments on pp. 123; 131.

Callithamnion caudatum J. Agardh in Linnaea 15 : 46 (1841).
Ceramiumfelicii Bonnemaison in Mem. Mus. Hist. nat. Paris 16 : 129 (1828).
Ceramium tetricum ft pectinatumC. Agardh, Syst. Alg.: 141 (1824).

3. Rejections from the genus

In a few cases, examination of type material has disclosed that the entities in question must
be rejected from the genus Callithamnion. For each, the reason for rejection is stated:

Callithamnion fallax P. Crouan & H. Crouan

Type locality: rade de Brest (Crouan & Crouan, 1 867 : 135).

Holotype: CO*, Brest, 27 December 1856.

Comments: the holotype consists of depauperate scraps ofAntithamnion sp.

Callithamnion parvulum P. Crouan & H. Crouan

Type locality: 'Sur le Codium spongiosum . . . rade de Brest' (Crouan & Crouan,
1867: 135).

Holotype: CO, Crouan, Brest, 4 March 1 858.

Other material: LD, Herb. Alg. Agardh 1 7973, possibly a duplicate of the holotype.

Comments: the holotype has the aspect of Spermothamnion, rather than Callithamnion,
and terminal procarps are present.

*Abbreviations relating to herbaria are listed in the acknowledgements (pp. 1 38-1 39).

CALLITHAMNION W THE BRITISH ISLES 131

Callithamnion tetricum y minus C. Agardh

Type locality: Pornic, coll. Desvaux (C. Agardh, 1828 : 180)
Holotype: LD, Herb. Alg. Agardh 18514.

Comments: the specimen is of Pleonosporium borreri, as was suggested by J. Agardh
(1851 :53).

Typifications

An attempt has been made to typify every name relevant to the present study. The results are
listed in alphabetical order, initially by genus and then by species.

Callithamnion acrospermum J. Agardh

Type locality: 'in Oceano atlantico ad oras Galliae' (J. Agardh, 1851: 52).
Lectotype: LD, Herb, Alg. Agardh 18295.
Other material: LD, Herb. Alg. Agardh 18296.,

Callithamnion affme Harvey

Type locality: Bute, coll. Greville (Harvey in Hooker, 1833 : 344).

Lectotype specimen: TCD, Greville, Bute, undated, annotated 'Plant of Br. Flora H' in
Harvey's hand.
Other material: TCD, three specimens. BM(K), one specimen.

Callithamnion arachnoideum C. Agardh

Type locality: 'in mari Atlantico ad Noveboracum, unde misit Torry' (C. Agardh,
1828: 181).

Lectotype: LD, Herb. Alg. Agardh 18435.

Other material: LD, Herb. Alg. Agardh 1 8436 [mica preparation, probably of 1 8435],
18443.

Callithamnion arachnoideum Griffiths ex Moore, nom. illeg.

Type locality: 'near the mouth of the Bann' (Moore, 1837 : 1 1).
Type material: it has not been possible to locate any original material.

Callithamnion arnottii Trevisan, nom. illeg.

Comments: C. arnottii (as 'C. Arnott?} was proposed by Trevisan (1 845 : 77) as a substitute
name for C. byssoides. It is, therefore, a superfluous and illegitimate name which must be
typified by the type of C. byssoides (q.v.).

Callithamnion bipinnatum P. Crouan & H. Crouan

Type locality: rade de Brest (Crouan & Crouan, 1 867 : 1 36).
Lectotype: CO, Crouan, rade de Brest, Aug. 1 850.
Other material: CO, 3 specimens.

Comments: for discussion of the possible status of this taxon in different parts of Europe,
see Price ( 1 978) and Rueness & Rueness ( 1980).

Callithamnion brodiaei Harvey

Type localities: Forres, Coll. Brodie; Northumberland, coll. Robertson; Torquay, coll.
Griffiths & Cutler (Harvey in Hooker, 1833 : 340).

132 P. S. DIXON & J. H. PRICE

Lectotype: TCD, Brodie, Lossiemouth, undated, annotated as 'authentic' in Harvey's
hand.

Other material: TCD, three specimens. LD, Herb. Alg. Agardh 19018, 19025. GL, two
specimens.

Comments: the original treatment contains two curious misstatements. The Brodie
specimens were collected at Lossiemouth according to Brodie's annotations, not at Forres,
which was where he lived and is a few kilometres from the sea. The Northumberland
specimen is annotated as having been collected by 'Mr. Robinson', whereas the collector
was quoted as 'Mr. Robertson' in the published text.

Callithamnion byssoides Araott ex Harvey

Type localities: 'Whitsand bay, G.A.W. Arnott, Esq. Devonshire, Mrs. Griffiths' (Harvey
in Hooker, 1833:342).

Lectotype: TCD, Arnott, Whitsand Bay, 1829, annotated 'original plant of Br. Flora
WHH' (in Harvey's hand).

Other material: TCD, one specimen. LD, Herb. Alg. Agardh 18407. GL, seven
specimens, possibly duplicates.

Callithamnion caudatum J. Agardh

Type locality: 'in mari mediterraneo ad littora Galloprovinciae; rupibus insulae St.
Marguerite' (J. Agardh, 1841 : 46).

Lectotype: LD, Herb. Alg. Agardh 18386.

Other material: LD, Herb. Alg. Agardh 18390. Numerous other specimens exist (BM,
TCD, etc.), distributed by J. Agardh, but it would seem best to exclude these because of
doubts as to their provenance.

Callithamnion decompositum J. Agardh

Type locality: 'in oceano atlantico ad oras Galliae' (J. Agardh, 1851: 45).

Holotype: LD, Herb. Alg. Agardh 1 8964.

Comments: this entity is often attributed, partly or completely, to Grateloup, due to a
misinterpretation of the comment by J. G. Agardh that Grateloup had given the manuscript
name Mertensia decomposita to the specimen used as the basis for the original treatment.

Callithamnion fruticulosum J. Agardh

Type locality: 'in sinu Codano' (J. Agardh, 1841 : 46).

Lectotype: LD, Alg. Agardh 19047, bearing TYPUS slip attributed to T. Levring.

Other material: LD, Herb, Alg. Agardh 19046, 19048. TCD, J. Agardh, undated, a mica
preparation in a packet annotated "Callith. fruticulosum Ag. MSCR e sinu Codano J.
Agdh.' in J. Agardh's hand.

Callithamnion furcellariae J. Agardh

Type locality: 'in Furcellaria parasiticum ad oras Bahusia' (J. Agardh, 1851 : 37).

Lectotype: LD, Herb. Alg. Agardh 19087.

Other material: LD, Herb. Alg. Agardh 19088, 19089.

Comments: a TYPUS label has been attached to Herb. Alg. Agardh 19085, but it is not
clear why this was selected because it makes reference to Solenia, not Furcellaria, which is
cited on the specimen cited above as lectotype.

Callithamnion grande J. Agardh

Type locality: 'ad rupes insularum Stoechadum' (J. Agardh, 1842 : 73).
Holotype: LD, Herb. Alg. Agardh 19277.

CALLITHAMNION W THE BRITISH ISLES 133

Callithamnion grevillei Harvey

Type localities: 'Firth of Forth, Dr. Greville, Berwick, Dr. Johnston' (Harvey in Hooker,
1833:345).

Lectotype: TCD, Greville, no locality, undated, a packet containing a mica preparation,
annotated 'Plant of Br.Fl.' in Harvey's hand.

Callithamnion harvcyanuni J. Agardh

Type localities: Callithamnion harveyanum was first used by J. Agardh (1841 : 45), who
claimed that the plant treated by Harvey under the name Callithamnion granulatum was
distinct from that taxon. As J. Agardh did not provide a formal diagnosis, the original
description must be that given by Harvey, and the type localities those quoted by the latter,
viz. Appin [in error, cf. comment below], coll. Carmichael; Whitsand Bay, coll. Arnott.

Lectotype: TCD, Carmichael, Staffa, July 1826, annotated 'Specimen described in British
Flora as C. granulatum is not that plant WHH' in Harvey's hand.

Other material: LD, Herb. Alg. Agardh 18339.

Comments: the lectotype and the LD specimen are both annotated to indicate that they
were collected by Carmichael at Staffa, not Appin, as indicated in the Harvey treatment.

Callithamnion hiemale Kjellman ex Kyi in

Type locality: Kristineberg (Kylin, 1907 : 170).
Holotype: LD, Kylin, Kristineberg, 9 December 1905.

Callithamnion lanosum Harvey

Type localities: 'Tor Abbey, Mrs. Griffiths. Killiney, W. H. Harvey' (Harvey in Hooker,
1833:341).

Lectotype: TCD, Harvey, Killiney, undated.

Other material: TCD, four specimens. BM, one specimen. LD, Herb. Alg. Agardh 18257,
18258.

Comments: Harris (1966) cited the type locality for this entity as 'Beach below Tor Abbey,
Menai Straits, N. Wales', a geographically meaningless aggregation of localities that bears no
relation to the original treatment.

Callithamnion octosporum C. Agardh

Type locality: 'Ad litora Galliae maris Atlantici, ut ad Port Louis' (C. Agardh,
1828: 177).

Holotype: LD, Herb. Alg. Agardh 18493.

Other material: LD, Herb. Alg. Agardh 18492, a mica preparation apparently derived
from 18493.

Callithamnion polyspermum C. Agardh

Type locality: 'Ad Finisterre Galliae, unde misit Bonnemaison' (C. Agardh, 1828 : 169).

Holotype: LD, Herb. Alg. Agardh 18236.

Other material; LD, Herb. Alg. Agardh 18235, a mica preparation apparently derived
from 18236. PC, F, QUI, several specimens occur which may possibly represent duplicates
of the holotype indicated above, since they were received from Bonnemaison and referred
by him, as was the holotype, to "Lamourouxia polysperma' (a nomen nudum).

Callithamnion pseudobyssoides P. Crouan & H. Crouan

Type locality: 'sur le Plocaria confervoides . . . Kervallon' (Crouan & Crouan, 1867 : 136).

Lectotype: CO, Crouan, Kervallon, undated.

Other material: CO, one specimen. LD, Herb. Alg. Agardh 18426.

134 P. S. DIXON & J. H. PRICE

Callithamnion roseum ft tenue Lyngbye

Type locality: Faeroe (Lyngbye, 1819 : 126).

Lectotype: LD, Herb. Alg. Agardh 18997.

Other material: LD, Herb. Alg. Agardh 18994. C, Lyngbye, 'Hoyviig prope Thorshavn
Faeroe', 11 June 1817. PC, Lyngbye, Faeroe, undated, labelled 'Callithamnion roseum
ft tenue', in Lyngbye's hand.

Comments: Lyngbye and C. Agardh obviously exchanged material extensively; C. Agardh
made use of exchange material as the basis for both Ceramium scopulorum and
Callithamnion scopulorum (q.v.).

Callithamnion scopulorum C. Agardh

Type locality: Faeroe (C. Agardh, 1828 : 166).

Lectotype: LD, Herb. Alg. Agardh 18997.

Other material: LD, Herb. Alg. Agardh 18994. C, Lyngbye, 'Hoyviig prope Thorshavn
Faeroe', 11 June 1817. PC, Lyngbye, Faeroe, undated, labelled 'Callithamnion roseum ft
tenue 'in Lyngbye's hand.

Comments: as discussed under Callithamnion roseum ft tenue (q.v.), C. Agardh made use
of Lyngbye exchange material of the latter entity as the basis for Callithamnion scopulorum.
The original treatment of the latter quoted 'Callithamnion roseum tenue Lyngb.' in its
synonymy, although there is no mention of Ceramium scopulorum (q.v.), which had already
been based (C. Agardh, 1 824) on the same material.

Callithamnion spiniferum Kylin

Type localities: Hogardsgrund, Varberg, Morup (Kylin, 1907 : 159).
Lectotype: LD, Kylin, Morup, 21 February 1903.
Other material: LD, Kylin, Hogardsgrund, 6 July 1907.

Callithamnion spinosum Harvey

Type locality: Sidmouth, coll. Griffiths (Harvey in Hooker, 1833 : 345).

Holotype: TCD, Griffiths, Sidmouth, undated, annotated 'Cal. spinosum H. in Br. Flora.
Sent by Mrs. Griffiths to Dr. Greville from Sidmouth my only specimen' in Harvey's hand.

Comments: Harris (1966) cited as holotype a specimen at TCD, undefined as to locality
and lacking further identifying data. That specimen is of no importance to the correct
typification.

Callithamnion spongiosum Harvey

Type localities: Torquay, coll. Griffiths; Kingston, coll. Harvey; Killiney, coll. Harvey
(Harvey in Hooker, 1833 : 346).

Lectotype: TCD, Griffiths, Meadfoot (Torquay), June 1 832.

Other material: TCD, five specimens. BM, two specimens. BM(K), one specimen. GL, one
specimen. LD, Herb. Alg. Agardh 19249.

Callithamnion tetragonum ft firmius C. Agardh

Type locality: 'Frishwater [sic] Bay' (C. Agardh, 1828 : 176).
Holotype: LD, Herb. Alg. Agardh 1 834 1 .

Other material: LD, Herb. Alg. Agardh 18342, a mica preparation apparently prepared
from 18341.

Callithamnion tripinnatum C. Agardh

Type locality: 'Mertensia tripinnata Gratel. Msct. In Mari Atlantico, unde specimen misit
Grateloup' (C. Agardh, 1828 : 168).

CALLITHAMNIONM THE BRITISH ISLES 135

Holotype: LD, Herb. Alg. Agardh 18975.

Comments: citation of the authority is confused. Direct attribution to Grateloup is
incorrect as his name is a nomen nudum, while citation in the form 'Gratel. ex. C. Ag.' is also
incorrect in that C. Agardh took up only the epithet and not the whole binomial from
Grateloup (see International Code of Botanical Nomenclature, Recommendation 46C. 1).

Ceramium brachiatum Bonnemaison, nom. illeg.

Comments: the original treatment (Bonnemaison, 1828:136) made reference to
collections from three localities on the Atlantic or Channel coasts of France, to another from
the Mediterranean, and to two specimens collected in the Canary Islands and at the Cape of
Good Hope. More significantly, it placed in synonymy binomials which lead back to
Conferva tetragona Withering and made no mention of the latter elsewhere in the text, so
that these synonyms must be taken as references to type. There is no option but to dismiss
Ceramium brachiatum as a superfluous and illegitimate name which must be typified by the
type of Conferva tetragona.

Ceramium congest urn Bonnemaison, nom. illeg.

Comments: the original treatment (Bonnemaison, 1828 : 125) made reference to
collections from nine localities on the Channel and Atlantic coasts of France. The critical
feature of the treatment is that it placed in the synonymy Conferva tetrica Dillwyn (1806).
On the basis of that citation, one must dismiss Ceramium congestum as a superfluous and
illegitimate name, to be typified by the type of Conferva tetrica.

Ceramium dudresnayi Bonnemaison, nom. illeg.

Comments: the original treatment (Bonnemaison, 1828 : 125) made reference to various
specimens but also to several previously-published binomials, of which the oldest is
Conferva purpurascens Hudson. Therefore Ceramium dudresnayi is a superfluous and
illegitimate name which must be typified by the type of Conferva purpurascens.

Ceramium felicii Bonnemaison

Type locality: 'Dieppe' (Bonnemaison, 1828 : 129).

Lectotype: PC, Gaillon, Dieppe, undated, annotated as 'Ceramium felicii Gaillon mnscr'
in Bonnemaison's hand.

Other material: specimens were distributed widely by Gaillon, although he never
provided a description. The greatest number were issued as specimen 203 of Desmaziere's
'Plantes Cryptogames' but others, not from this source, occur at CO (ex Crouan) and at LD
(Herb. Alg. Agardh 19016, also ex Crouan).

Ceramium fruticulosum Roth

Type locality: not stated (Roth, 1800 : 183), but as the original material was collected by
Wulfen it was probably from the Adriatic.

Type: typification is not currently possible as there is no evidence at either B or W to
indicate that the original material is still in existence and no illustration is provided.

Comments: Roth (1800) compared his alga with Conferva marina nodosa of Dillenius
(1741), the type of Conferva purpurascens Huds., and in a later treatment (Roth, 1806 : 147)
actually cited the latter in his treatment of Ceramium fruticulosum. Wulfen (1803)
independently described Conferva fruticulosa (q.v.), and it is probable that the two entities
are based on fragments of the same original gathering.

Ceramium granulatum Ducluzeau

Type locality: Sette (Ducluzeau, 1806 : 72).

136 P. S. DIXON & J. H. PRICE

Type: typification is not currently possible as nothing is known of the location of any
Ducluzeau material and no illustration, although announced, is provided with the original
treatment.

Ceramium guttatum Bonnemaison

Type localities: Penmark, Cherbourg (Bonnemaison, 1828 : 137).

Lectotype: PC, locality not indicated, Bonnemaison, undated, labelled 'Ceramium
guttatum B n sp. nov.', in Bonnemaison's hand, and filed under Callithamnion tetragonum.

Ceramium roseum Roth

Type locality: Bayonne (Roth, 1798 : 47).

Type: indisputable type material not located. Neotype designated below.

Comments: the existence of Roth's herbarium is at present a matter of doubt. Usually, it is
assumed to have been lost in the bombing of the Berlin-Dahlem Museum (Pilger, 1953); this
seems true of specimens of Ceramium roseum. There is a specimen at LD (Herb. Alg.
Agardh 18466), labelled 'Ceramium roseum Roth Bayonne dedit Roth ipse non rosea Dillw'
in C. Agardh's hand, but the status of this is uncertain. Subsequent interpretations (Dillwyn,
1802; Smith, 1802) were based on a specimen collected in 1797 by Sowerby at Yarmouth
and sent to Roth for comment by Dawson Turner. This specimen, which Roth identified as
'Ceramium roseum' , passed to K (now at BM), with a duplicate of the original Sowerby
collection at BM. Although not part of the type collection, these specimens help to unravel
subsequent interpretations of this taxon, of which there have been several. This K(BM)
specimen is here designated as of neotype status.

Ceramium scopulorum C. Agardh

Type locality: Faeroe (C. Agardh, 1824 : 132).

Lectotype: LD, Herb. Alg. Agardh 18997.

Other material: C, Lyngbye, 'Hoyviig prope Thorshavn Faeroe', 11 June 1817. PC,
Lyngbye, Faeroe, undated, labelled 'Callithamnion roseum tenue" in Lyngbye's hand.

Comments: as discussed under Callithamnion roseum ft tenue (q.v.), Lyngbye and C.
Agardh exchanged material and C. Agardh made use of material of the latter entity as the
basis for Ceramium scopulorum. The original treatment of the latter quoted 'Callithamnion
roseum tenue, Lyngb.' in its synonymy. The same material was later used by C. Agardh as
the original material for Callithamnion scopulorum (q.v.), although the latter description
makes no reference to the earlier Ceramium scopulorum.

Ceramium tenuissimum Bonnemaison

Type localities: St-Pol-de-Leon, Brest, St. Malo (Bonnemaison, 1828 : 132).

Provisional lectotype: PC, Bonnemaison, no locality, undated, annotated 'Ceramium
tenuissimum B"' in Bonnemaison's hand, filed as Seirospora tenuissimum.

Comments: despite widespread search at QUI, it was not possible to locate any authentic
material in what remains of Bonnemaison's herbarium. Presumably the specimen cited
above as provisional lectotype is that referred to by Harris (1966).

Ceramium tetricum ft pectinatum C. Agardh

Type locality: 'Ad oras Galliae, Africae Borealis' (C. Agardh, 1 824 : 141).
Lectotype: LD, Herb. Alg. Agardh 18307.
Other material: LD, Herb. Alg. Agardh 18309.

CALLITHAMNIONW THE BRITISH ISLES 137

Ceramium versicolor C. Agardh nom. illeg.

Comments: the original treatment (C. Agardh, 1824:140) cites three previously
published binomials, one identification, and the manuscript source of the epithet. Despite
widespread search, we have been unable to locate the type material of any of the previous
binomials. Ceramium versicolor is a superfluous and illegitimate name, incapable of typifi-
cation, which must be dismissed from consideration.

Conferva arbuscula Dillwyn

Type localities: Ballycastle, coll. Brown; Bantry Bay, coll. Hutchins (Dillwyn, 1807, pi.
85).

Lectotype: BM(K), ? Brown, North coast of Ireland, undated.

Other material: BM, 2 specimens. GL, 2 specimens.

Comments: the typification has been discussed in detail by Dixon (1960, 1964). The
choice of lectotype was made on the basis of the statement by Dillwyn (1809 : 35) that
Dawson Turner provided the original description of the entity. The position became
confused as a consequence of a later figure (Dillwyn, 1809, pi. 6G) based on a specimen of
Dasya, so that the epithet arbuscula came to be applied in both Callithamnion and Dasya.
Despite previous resolution of this problem (Dixon, 1960, 1964), Harris (1966) stated that he
had been unable to locate original material.

Conferva corymbosa Smith

Type localities: Brighthelmstone [=Brighton], coll. Borrer; Kingsbridge, coll. Gibbs;
Bantry Bay, coll. Hutchins (Smith, 1 8 1 1 , pi. 2352).

Lectotype: BM, Borrer, Brighton, July 1811.

Comments: Harris (1966) quoted the type locality as Kingsbridge and cited as holotype a
specimen 'in the British Museum of Natural History, no. 271'. The original treatment is
based on three collections, so that holotype status for any specimen is impossible. It would
appear that this author was misled by the seven irrelevant specimens, in the type folder at
BM, mounted on the same sheet as the Borrer specimen. The specimen cited by Harris is an
example from 'Algarum Fasciculi' distributed by J. Cocks, and has no relevance whatsoever
to the typification of Conferva corymbosa.

Conferva fruticulosa Wulfen

Type locality: 'Insulae Gradensis' (Wulfen, 1803 : 26).

Type: typification is not currently possible as the original material cannot be located, and
no illustration is provided with the original treatment.

Comments: Wulfen stated that his material was similar to the illustration of Conferva
marina nodosa of Dillenius (1742). Roth (1800) described Ceramium fruticulosum on the
basis of material supplied by Wulfen and also commented on the relationship to the same
Dillenian entity. Conferva fruticulosa Wulfen and Ceramium fruticulosum Roth are
independent names, based on different types, although these are probably fragments of the
same original gathering.

Conferva hookeri Dillwyn

Type localities: Cawsie, coll. Hooker and Borrer; Holyhead, coll. Davies; Bantry Bay, coll.
Hutchins (Dillwyn, 1809 : pi. 106).

Lectotype: TCD, Hooker and Borrer, Cawsie, undated, labelled 'Br. Flora C. Hookeri
Cawsie Hook. Borrer', in Harvey's hand.

Other material: BM(K), three specimens.

Comments: Harris (1966) cited a K specimen, supposedly from Holyhead, Anglesey.
None of the K material mentioned above is so annotated and the specimen at TCD is the
legitimate choice of lectotype.

138 P. S. DIXON&J. H. PRICE

Conferva purpurascens Hudson

Type locality: unspecified (Hudson, 1778 : 600).

Holotype: OXF, Dillenian material of Conferva marina nodosa.

Comments: Hudson's description is based exclusively on the plant described previously by
Dillenius (1742) as Conferva marina nodosa . . . Harris (1966) came to a similar conclusion,
although he considered the Dillenian material as of neotype status. Although Hudson cited
no precise locality in his treatment of Conferva purpurascens, Dillenius (1742 : 36) indicates
that his material was collected near 'Lhanfaethly', Anglesey, which is therefore the type
locality for the Hudson taxon.

Conferva seposita Gunnerus

Type localities: 'ad Hoburgam Gotlandiae; in man sundmorico [near Alesund, Norway];
prope Lhanfaethly in Mona Insula [Anglesey]' (Gunnerus, 1772 : 116).

Lectotype: OXF, Green, Lhanfaethly, undated, in Dillenius's Historia Muscorum
herbarium.

Comments: the lectotype selected is also the lectotype of Conferva purpurascens Hudson
(q.v.).

Conferva tetragona Withering

Type locality: 'found by Major Velley and Mr. Stackhouse at the Bill of Portland'
(Withering, 1796:405).
Provisional lectotype: GL, Velley, Isle of Portland, undated.

Conferva tetrica Dillwyn

Type locality: 'Common about the Mumbles and in other parts of the Peninsula of Gower'
(Dillwyn, 1806, pi. 81).
Lectotype: WELT, Herb. Silvanus P. Thompson A 4502, Dillwyn, Swansea, 1806.

Ellisius glaber S. F. Gray, nom. illeg.

Comments: E. glaber is a superfluous and illegitimate name, in that Conferva arbuscula
Dillw. is cited as a synonym (Gray, 1821 : 333). The genus Ellisius was established with two
species, E. glaber and E. coccineus, the latter based on Conferva coccinea Hudson
[= Heterosiphonia plumosa (Ellis) Batters]. Heterosiphonia Montagne (1842) has been
conserved against Ellisius.

Phlebothamnion faroense Kiitzing

Type locality: 'Faeroe' (Kiitzing, 1864 : 30).

Holotype: L, Herb. Lugd. Bat. 940.237.1 16, Lyngbye, Faeroe, undated.

Phlebothamnion scoticum Kiitzing

Type locality: Aberdeen (Kiitzing, 1864 : 30).

Holotype: L, Herb. Lugd. Bat. 940.237.1 18, Dickie, Aberdeen, June 1843.

Acknowledgements

A work of this magnitude could not have been conducted without the active support and co-operation
of many institutions and collections. We wish to express our indebtedness to the Directors and
Curators of the following institutions: School of Plant Biology, University College of North Wales,
Bangor (UCNW); Laboratoire de Biologic Marine, Concarneau (CO); Botanical Museum and
Herbarium, University of Copenhagen (C); University College, Cork; School of Botany, Trinity

CALLITHAMNION IN THE BRITISH ISLES 1 39

College, Dublin (TCD); Herbarium, Royal Botanic Garden, Edinburgh (E); Istituto Botanico, Firenze
(FL); Department of Botany, University of Glasgow, Glasgow (GL); Royal Botanic Gardens, Kew (K)
[algal material now at BM]; Rijksherbarium, Leiden (L); Merseyside County Museum, Liverpool (LIV);
British Museum (Natural History), London (BM); Botanical Museum, Lund (LD); The Manchester
Museum, The University of Manchester, Manchester (MANCH); Hancock Museum, Newcastle-upon-
Tyne (HAMU); Botanical Museum, Oslo (O); Fielding-Druce Herbarium, Oxford (OXF); Laboratoire
de Cryptogamie, Museum National d'Histoire Naturelle, Paris (PC); Eastgate House Museum,
Rochester; Station Biologique, RoscofT; Sunderland Museum, Sunderland; County Museum, Truro;
Institute of Systematic Botany, University of Uppsala, Uppsala (UPS); Museo Civico di Storia
Naturale, Venezia; National Museum, Wellington (WELT); Naturhistorisches Museum, Wien (W).

Financial support has been received from various sources over the past 20 years, including the Joint
Committee on Research of the University of Liverpool; the Browne Fund of the Royal Society, London;
the British Council; the Faculty Research and Travel Fund of the University of California; the
National Science Foundation; and the British Museum (Natural History).

Many individuals have assisted with discussions and with the provision of information and
specimens. We would like to thank P. Edwards, W. F. Farnham, M. D. Guiry, J. D. Guiterman,
C. E. L. Hepton, D. E. G. & L. M. Irvine, D. M. John, T. Levring, C. Maggs, O. Morton, W. Prud'
homme van Reine, J. and M. Rueness, P. C. Silva, I. Tittley, J. A. West & A. Whittick for this
assistance.

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Hildur Krog & T. D. V. Swinscow

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/<$" GENERAL

Parmelia subgenus Amphigymnia (lichens) in East ^f
Africa N ^C^

Hildur Krog

Botanical museum, Trondheimsveien 23 B, Oslo 5, Norway
T. D. V. Swinscow

24 Monmouth Street, Topsham, Exeter, England

Contents

Page

Synopsis 143

Introduction 144

Previous collections and research 144

Taxonomic review 144

Materials and methods 145

Results 145

Morphology 145

Chemistry 148

Ecology and distribution 150

Discussion 153

Circumscription of the species 153

Generic concept and infrageneric classification 158

The East African Amphigymnia species 159

Key to the species 159

Descriptions of the species 165

Acknowledgements 227

References 227

Index 229

Synopsis

Seventy species of Parmelia subgenus Amphigymnia (Vainio) Dodge are reported from East Africa,
their ecology and distribution discussed, and a key and descriptions to the species are provided. The
following new species are described: Parmelia aprica, P. durumae, P. jacarandicola, P. kwalensis, P.
leonis, P. pardii, P. pigmentifera, P. rava, P. taitae, P. tsavoensis, P. umbrosa, and P. vivida. The new
name Parmelia indoafra is proposed for Parmotrema indicum Hale, non Parmelia indica Sprengel.
The following taxa are reduced to synonymy: Parmelia aberrans (Vainio) des Abb., P.
madagascariacea (Hue) des Abb., and P. nyasensis Dodge with P. xanthina (Mull. Arg.) Vainio;
Parmelia ajjluens Hale with P. dilatata Vainio; Parmelia balensis Winnem, P. neolobulascens
Winnem, and Parmotrema subcompositum Hale with Parmelia subschimperi Hale; Parmelia
breviciliata Hale and P. procera Steiner & Zahlbr. with P. maclayana Mull. Arg.; Parmotrema
conferendum Hale and Parmelia leucosemotheta Hue with P. subsumpta Nyl.; Parmelia composita
Hale and P. spilota Hale with P. euneta Stirton; Parmelia deflectens Kurok. with P. subarnoldii des
Abb.; Parmelia diversa Hale with P. nilgherrensis Nyl.; Parmelia modesta Hue with P. andina Mull.
Arg.; Parmelia mordenii Hale with P. praesorediosa Nyl.; Parmelia natalensis Steiner & Zahlbr. and P.
pseudonilgherrensis Asah. with P. lobulascens Steiner; Parmelia pachyspora Hale with P. amaniensis
Steiner & Zahlbr.; Parmelia paradoxa Hale with P. inexpectata des Abb.; Parmelia pseudoreticulata
Tavares and P. simulans Hale with P. reticulata Taylor; Parmelia pseudotinctorum des Abb. with P.
tinctorum Nyl.; and Parmelia virens Mull. Arg. with P. erubescens Stirton. Parmelia consors Nyl. and
P. pilosa Stizenb., previously classified in genus Parmelina Hale, are here included in subgenus

Bull. Br. Mus. not. Hist. (Bot.) 9 (3) : 1 43-23 1 Issued 26 November 1 98 1

143

144 H. KROG & T. D. V. SWINSCOW

Amphigymnia (genus Parmotrema Massal.). The generic concept and infrageneric classification are
briefly discussed.

Introduction
Previous collections and research

Species of Parmelia subgenus Amphigymnia (Vainio) Dodge form one of the major
dominant groups of lichens in East Africa. Being large and conspicuous they were frequently
collected by both botanists and non-botanists on various expeditions during the 19th
century. New species and infraspecific taxa now classified in Amphigymnia were
described by contemporary lichenologists (names in parentheses) from material brought
back to Europe by the following collectors: Hannington (Muller, 1890), Hildebrandt
(Krempelhuber, 1877, Miiller, 1884, 1885), Hoist (Muller, 1894), Liechtenstein &
Pospischill (Steiner, 1897), Maclay (Muller, 1891), Meyer (Stein, 1888; Muller, 1890),
Schimper (Muller, 1892), Scott Elliot (Vainio, 1898), Stuhlmann (Muller, 1893, 1894),
Thomson (Stirton, 1877-78), and Volkens (Muller, 1894). Most of these new taxa, together
with records of other lichens occurring in our area, were included in an account by
Stizenberger(1890, 1891, 1893, 1895) of lichens known from Africa at that time.

In this century Jatta (1909) published an account of lichens from the Ruwenzori
mountains, and Hue (1916) reported on lichens collected in Kenya by Vicomte de Poncins.
Steiner & Zahlbruckner (in Zahlbruckner, 1 926), Zahlbruckner ( 1 932), and Zahlbruckner &
Hauman (1936) published on various collections of lichens from the African continent.
Several of them are pertinent to our area, especially those collected by Brunnthaler, as well
as those by Fincke, Schroder, Tobler, and F. & R. Wettstein. Cengia Sambo (19370, b, 1938,
1940) reported on lichens mainly from Ethiopia, but also from Kenya and Tanzania, and
Klement (1962) published an account of a small collection of lichens from Tanzania made
by Drs E. and R. Schiiz.

Dodge (1953, 1959) studied those East African Parmelias lodged in the herbaria of EA,
FH, and K (now in BM), describing a number of new species. His accounts include many
species in Amphigymnia which are now accommodated elsewhere. Hale (1965) published a
world monograph of the subgenus, reducing numerous taxa to synonymy and describing
many new species. Nearly 30 of the species he treated were then known to occur in East
Africa. In subsequent publications Hale (1972, 1973, 1977) described several additional
species from our area. Winnem (1975) treated the 3 1 species of the subgenus Amphigymnia
occurring in Ethiopia. A further two species have previously been discussed by the present
authors (Krog, 1974; Krog & Swinscow, 1975). It is noteworthy that most of the authors cited
were not familiar with the African lichen flora in the field. The main exception is Winnem
(1975), who based her study of Ethiopian lichens on her own collections and a first-hand
knowledge of the physiography of the country.

The aim of the present study is to give a comprehensive account of the species belonging to
Parmelia subgenus Amphigymnia in East Africa. Special emphasis is placed on the circum-
scription of the species in the light of characters of the reproductive organs, new chemical
data, ecological observations, and a knowledge of the variation within local populations.

Taxonomic review

Massalongo (1860) segregated the genus Parmotrema Massal., based on Parmelia perforata
(Jacq.) Ach., from Parmelia Ach. Vainio (1890) introduced Parmelia subgenus Euparmelia
sect. Amphigymnia Vainio with 1 1 species, but did not designate a type. Dodge (1959) raised
the section to subgeneric level, noting that a type had not been designated. Hale (1965)
treated the subgenus monographically and selected Parmelia tinctorum Nyl. as the lectotype
for it. Hale (19746) reintroduced genus Parmotrema Massal., including in it not only most of

P ARM ELI A (AMPHIGYMNIA) IN EAST AFRICA 145

the species which had previously been accommodated in subgenus Amphigymnia (Vainio)
Dodge, but also the P. reticulata group, previously included in subgenus Parmelia.

Materials and methods

The following account is based mainly on collections made by the authors as follows:
Uganda (T. D. V. S. 1969, 1970, 1971), southern Ethiopia (H. K. 1972), Kenya (H. K. & T.
D. V. S. 1972, 1973, 1974, H. K. 1976, T. D. V. S. 1977), and northern Tanzania (H. K. & T.
D. V. S. 1974). In addition to specimens in institutional herbaria we have been permitted to
examine material collected in recent years by the following botanists: Lye (Uganda), Moberg
(Kenya, Tanzania), Pocs (Tanzania), Santesson (Kenya, Tanzania), Sharp (Tanzania),
Sipman (Kenya, Tanzania), and Vitikainen (Kenya). Specimens cited with a number but
without a collector's name were collected by one of the authors. Specimens have been
deposited in O, BM, and East African herbaria.

The material referred to in this study has been subjected to thin-layer chromatography
(TLC) by means of standard methods (C. Culberson, 1972; Menlove, 1974). However, type
specimens which have recently been tested by TLC have not been retested unless it was
specially indicated. All specimens cited, including types, have been personally studied
unless otherwise noted. In the text + denotes a low concentration of a given substance, whilst
denotes its sporadic occurrence (accessory substance).

The descriptions of the species are based on East African material, and comprise only
those morphotypes and chemotypes which occur in our area. Only selected East African
records are given for the commoner species, but in such cases at least one record is chosen
from each district (or, for Uganda, each county) in which the species has been collected. Only
new or otherwise interesting synonyms are given. For a more complete citation see Hale
(1965).

Results
Morphology

The apothecia of Parmelia tinctorum, the type species of the subgenus, are substipitate and
laminal. But in the majority of East African species they were found to be submarginal to
marginal, with swollen and inflated or more or less constricted but well developed stipes. At
or near the lobe margin the stipe was sometimes formed by a semitubular fold of a
convoluted lobe, and in such cases the disc might be eccentrically orientated in relation to its
stipe.

In contrast to those of other subgenera in Parmelia, Amphigymnia species often have
perforated apothecial discs. In some species the perforations were found to have developed at
an early stage in nearly all the apothecia of a thallus, while in others they were present only
in some of the apothecia of a given thallus or occurred sporadically and seemingly accident-
ally in old apothecia.

The spores were simple and ellipsoid and varied in length from 10 to 40 //m. With some
individual variation, spore size in the subgenus could be roughly divided into three fairly
well defined categories: (a) small, thin-walled, 10-20 (22) //m; (b) intermediate, usually
relatively thick-walled, 20-28 /*m; and (c) large, thick-walled, 25-40 //m. In the first group it
was, to a certain degree, even possible to distinguish between species with spores less than c.
16 jum and those 16-20 (22) //m in length. Our own observations and previous reports show
that the East African species could be divided by spore size into the four groups shown in
Table 1 . In sorediate species the spores were sometimes poorly developed and on the average
smaller than in their non-sorediate counterparts.

The pycnidia were situated in a broad submarginal zone in all primary species, and were

146 H. KROG & T. D. V. SWINSCOW

Table 1 Approximate spore sizes in the East African species of Parmelia subgenus
Amphigymnia.

10-16 (18) //m

16-20 (22) urn

20-28 (30) fim

25^0 urn

P. aldabrensis

P. abessinica

P. dilatata

P. amaniensis

P. cetrata

P. andina

P. durumae

P. bangii

P. erubescens

P. austrosinensis

P. euneta

P. crinita

P. eurysaca
P. grayana

P. cooperi
P. gardneri

P. inexpectata
P. lobulascens

P. cristifera
P. cryptoxantha

P. jacarandicola
P. maclayana

P. hababiana
P. hanningtoniana

P. nilgherrensis
P. perlata

P. eciliata
P. subarnoldii

P. pardii

P. hololoba

P. subschimperi

P. reticulata

P. leonis

P. sancti-angelii

P. mellissii

P. soyauxii

P. pigmentifera

P. stuhlmannii

P. praesorediosa

P. subcolorata

P. pseudocrinita

P. subisidiosa

P. rava

P. subsumpta

P. rimulosa

P. subtinctoria

P. uberrima

P. taitae

P. vivida

P. tinctorum

P. zollingeri

P. ultralucens

P. xanthina

also found in this position in a number of secondary species. In modern taxonomic treat-
ments of parmelioid lichens little emphasis has been placed on the conidia, although their
shape and size were faithfully recorded by most earlier lichenologists as an important part of
the descriptions of new species. Within some lichen genera there is obviously little variation
between the species, but in Parmelia s. lat. the conidia were very variable and are therefore
treated here in some detail.

In the Parmeliaceae the conidia are produced endobasidially on conidiophores of the
bayonet type (Gliick, 1899) (Fig. la). The development of the conidiophores, which has been
discussed by Vobis (1980), is of little interest in the present study and is not treated here. As
for the conidia, however, we have found that five common types can be recognized within
the Parmeliaceae:

(a) Bifusiform conidia. The term has been applied by, for example, Esslinger (1977) and
Vobis (1980), to short conidia, usually 4-6 (7) yum long, which are constricted in the middle
with a swelling near but not at both ends (Fig. Ib). Other terms for this type are
'hantelformig' and 'bisquitfdrmig' (Gliick, 1899) and 'dumb-bell-shaped' (Karnefelt, 1979).
Within the Parmeliaceae we have observed bifusiform conidia in Cetraria, Cetrelia,
Hypogymnia, ;>id Parmelia subgenus Hypotrachyna, but never in subgenus Amphigymnia.

(b) Sublageniform conidia. The term has been used to denote short, rod-shaped conidia,
5-8 //m long, with a slight swelling near one end (Fig. Ic). Nylander (1885) applied the term
to the conidia of Parmelia corniculans, and Steiner & Zahlbruckner (in Zahlbruckner, 1926)
to those of P. procera. The term lageniform means flask-shaped, but since the swelling in this
type of conidia is subterminal rather than terminal, the term sublageniform is adopted. We
find this term more appropriate than Gliick's choice 'keulig' or clavate (Gliick, 1899). In
some species the swelling is indistinct and may be discernible only in a few of the many
conidia from a pycnidium, while the rest may appear rod-shaped. In the Parmeliaceae sensu
Henssen & Jahns (1973) sublageniform conidia occur in, for example, Usnea (Swinscow &
Krog, 1976, Fig. 7), Letharia, Platismatia, and Parmelia subgenus Amphigymnia pro parte.

PARMELIA (AMPHIGYMNIA) IN EAST AFRICA

147

Fig. 1 Conidia in genus Parmelia s. lat. a: conidiophore with filiform conidia, P. euneta Stirton,
3K 16/122. b: bifusiform conidium, P. ducalis Jatta, E 30/12. c: sublageniform conidium, P.
maclayana Mull. Arg., Ryvarden 9032. d: unciform conidium, P. subpraesignis Nyl., T 16/137.
e: rod-shaped conidium, P. eciliata (Nyl.) Nyl., 4 May 1978, Dahls.n.

Sublageniform conidia with one or two darker 'dots' (see below under rod-shaped conidia)
have been seen in the subgenus Everniiformes.

(c) Unciform conidia. We have chosen this term to denote rod-shaped conidia c. 5 //m long,
with a single hook-shaped end (Fig. Id), a type not discussed by Gliick (1899) or Vobis
(1980). They are particularly characteristic for Parmelia subgenus Parmelia sect. Simplices,
occurring in all the species examined by us, with the exception of P. microsticta Mull. Arg.
and P. hypoleucites Nyl s. str. in which they were filiform (see also W. Culberson & C.
Culberson, 1980). The presence of unciform conidia in Amphigymnia has not been con-
clusively proved.

(d) Rod-shaped (bacilliform) conidia. This is a rather heterogeneous group, consisting of
rods mainly 8-1 0(1 2) //m long and about 10-12 times as long as broad (Fig. le). They have
no swellings, but in some cases one or two darker 'dots' are situated in either a central,
subapical, or apical position. We have made no effort to distinguish between the various
types of 'dotted rods' in this work. They are of little interest in Amphigymnia but occur in
the subgenus Cyclocheila and subgenus Parmelia sections Parmelia and Bicornuta. Plain
(i.e. non-'dotted') rods, on the other hand, occur in a few Amphigymnia species, such as
Parmelia eciliata and P. mesotropa. They form a transition between weakly sublageniform
and filiform conidia.

(e} Filiform conidia. Filamentous conidia have been described as filiform by, for example,
Vainio (1890), Moberg (1977), and Vobis (1980). Their diameter is usually about 1 jum or
less, so that the conidia are at least 15-20 times as long as broad (Fig. la). Plain rod-shaped
conidia mainly differ from filiform conidia in being broader in relation to their length and

148 H. KROG & T. D. V. SWINSCOW

usually less than 10 /zm long. In the Parmeliaceae longly filiform, curved conidia, about
30 //m long, are known in Parmeliopsis. In Parmelia s. lat. filiform conidia (usually less than
20 /zm long) occur rarely in Simplices and Cyclocheila but are common in Amphigymnia.
They are mainly straight, but occasionally a conidium may be bent or curved.

The shape and size of most forms of conidia were found to show very little variation within
a single species. Filiform conidia were a notable exception, varying considerably in length
from specimen to specimen of a species, although they were often of fairly uniform size
within individual pycnidia.

Important thalline characters in the subgenus Amphigymnia are the presence or absence
of vegetative propagules, cilia, and maculae, and the colour of the lower cortex. These
characters have been discussed in detail by Hale (1965). For further comments, see below
under 'Circumscription of the species' (p. 1 53).

A number of more or less well established species pairs were present in our area. Table 2
shows the primary species of such pairs and their sorediate and isidiate counterparts.

Table 2 Species counterparts in East African representatives of Parmelia subgenus
Amphigymnia.

No soralia or isidia

Sorediate

Isidiate

P. abessinica

P. hababiana

P. aldabrensis

P. kwalensis

P. amaniensis

P. subarnoldii

P. andina

P. austrosinensis

P. cetrata

P. reticulata

P. subisidiosa

P. eciliata

P. perlata (?)

P. crinita

P. erubescens

P. subsumpta

P. subtinctoria

P. euneta

P. subschimperi

P. hololoba

P. cooperi

P. pseudocrinita

P.jacarandicola

P. direagens (?)

P. maclayana

P. poolii

P. nilgherrensis

P. lobulascens

P. soyauxii

P. defecta

P. stuhlmannii

P. subcolorata

P. permutata

P. taitae

P. pseudograyana

P. uberrima

P. louisianae (?)

P. vivida

P. aprica

P. zollingeri

P. gardneri

Chemistry

Atranorin and usnic acid are present either separately or jointly in the upper cortex. The
/?-orcinol depsidone psoromic acid is restricted to the soralia in Parmelia direagens, and to
the cortex near the apothecia in P. jacarandicola. In the medulla the following substances
occur: orcinol depsides (lecanoric and gyrophoric acids), orcinol depsidones (alectoronic,
a-collatolic, and physodic acids, norlobaridone, loxodine, and the lividic acid complex),
/?-orcinol depsides (generally lacking in the species of our area except that the mainly cortical
substance atranorin may be responsible for the K+ yellow reaction in the medulla of P.
praesorediosa), /?-orcinol depsidones (stictic, norstictic, salazinic, galbinic, protocetraric,
fumarprotocetraric, and echinocarpic acids), fatty acids (caperatic and protolichesterinic
acids, the reddenda fatty acid complex, and other unknowns), a xanthone (lichexanthone),
and various pigments. The substances, with the species in which they occur, are presented in
Tables 3-5. Only those of diagnostic value are listed, while substances which in our

FARM ELI A (AMPHIGYMN1A) IN EAST AFRICA 149

Table 3 Chemical properties of species without soralia and isidia.

lee gyr nib ale liv stc nst sal pso pc fpc fat pgm ind atr usn

P. abessinica

. . x.

P. aldabrensis

XX . .

P. amaniensis

X.. . X.

P. andina

X X

P. cetrata

X +

P. durumae

X X

P. eciliata

X X

P. erubescens

X....X + .

P. euneta

... X.

P. eurysaca

X X .

P. hanningtoniana

X X. . X.

P. hololoba

X X .

P. inexpectata

X X .

P. jacarandicola

X +.... . X.

P. leonis

X . X

P. maclayana

X X.

P. nilgherrensis

.X X.

P. pigmentifera

XXX.

P. soyauxii

X X

P. subcolorata

X X . X.

P. taitae

XX.. . X .

P. uberrima

X X .

P. vivida

X ... . .X

P. zollingeri

X . . . X .

Only substances of diagnostic importance are included, lee = lecanoric acid, gyr = gyrophoric acid, nlb =
norlobaridone, alc = alectoronic acid (including or-collatolic acid), liv = the lividic acid complex, stc = stictic acid,
nst = norstictic acid, sal = salazinic acid, pso = psoromic acid, pc = protocetraric acid, fpc = fumarprotocetraric acid,
fat = fatty acids, pgm = pigments, ind = undetermined substances, atr = atranorin, usn = usnic acid. Symbols:
x = constant; + = low concentrations; = occasionally present (accessory substance) or present in a chemical strain.

specimens always occur in conjunction with one of the main compounds, such as loxodine,
oxyphysodic acid, connorstictic acid, constictic acid, unknown with alectoronic acid, and
unknowns with stictic acid, are omitted from the tables.

In P. pigmentifera we found two undetermined main substances in the following Rf
classes: in IDA 2-3 : 4, in HEF 4-5 : 6, and in TA 2-3 : 5-6. The medulla reacted PD+
ochraceous, K+ deep yellow, C , KC+ red. The reactions with K and PD may have been
caused either by the above mentioned unknowns or by an unknown medullary pigment,
while the KC reaction points to the presence of one or two orcinol depsidones. Both
substances gave a bright bluish white fluorescence under long-wave U V light.

Among the fatty acids of diagnostic importance is an array of substances which occur
together, having Rf values intermediate between those of caperatic and protolichesterinic
acids; they are referred to as the 'reddenda fatty acid complex'. The name is derived from
their presence in, for example, P. reddenda Stirton (Krog & Swinscow, 1977). Unidentified
fatty acids, usually with rather high Rf values, are accessory in some species.

In the P. dilatata group (including, for example, P. affluent and P. progenes) atranorin and
usnic acids are produced together in considerable amounts. The proximity of other
substances sometimes caused the spots indicating usnic acid to take on a dark sordid grey
colour on the TLC plates. The presence of usnic acid in this species group has gone partly

1 50 H. KROG & T. D. V. SWINSCOW

Table 4 Chemical properties of species with isidia, sorediate isidia, or dactyls.

lee gyr nib ale phy stc nst sal pc ech fat pgm lex ind atr usn

P. crinita
P. cryptoxantha
P. kwalensis
P. lophogena

P. mellissii
P plcincitilobcitci

x x

XXX.. X
XX X

x x. . . x.

x x
x x

P pseudocrinitci

x . x .

P stuhlffidfinii

x x

P. subisidiosa
P. subtinctoria
P sulphurttta

x +

X....X.... .. +.
X + .

P tinctorum

X . ...... X .

P. tsavoensis
P. ultralucens
P. xanthina

x x

X .... X X .
X . .X .X

Only substances of diagnostic importance are included, lee = lecanoric acid, gyr = gyrophoric acid, nlb =
norlobaridone, ale = alectoronic acid (including ct-collatolic acid), phy = physodic acid, stc = stictic acid, nst =
norstictic acid, sal = salazinic acid, pc = protocetraric acid, ech = echinocarpic acid, fat = fatty acids, pgm = pigments
(including vulpinic acid), lex = lichexanthone, ind = undetermined substances, atr = atranorin, usn = usnic acid.
Symbols: x = constant; + = low concentrations.

unnoticed, and the grey spots were thought by Hale (1971c, 1977), for instance, to represent
an unknown substance.

Usually the chemical properties of lichens remain unchanged after extended storage in the
herbarium. However, we noticed that in old specimens with alectoronic and a-collatolic
acids we often obtained additional spots below those of the main substances on the TLC
plates, correlated with an ochraceous reaction with C in the medulla. They were especially
prevalent in the P. nilgherrensis group, of which many old collections exist. We found no
correlation between morphological characters and the presence or absence of such
substances, and are inclined to regard them as artefacts. On the other hand, an unknown with
low Rf values, occurring with alectoronic acid in a chemical strain of P. maclayana, appears
to be of diagnostic value. It will be referred to below as 'unknown with alectoronic acid'.

The African chemotype of P. xanthina contained an undetermined C , KC+ red
substance which is probably close to gyrophoric acid. It ran slightly higher than gyrophoric
acid on the TLC plates in TDA and HEF.

Ecology and distribution

Tropical East Africa offers a wide range of habitats, from the sea coast to more than 5000 m
altitude, and from subdeserts and savannas to montane rain forests. Very few lichens are
found in the large subdesert areas of northern Kenya and Uganda and southern Ethiopia, and
also in the dry thornbush savannas which lie below 1000 m altitude and have an annual
rainfall of less than 500 mm.

While species of Parmelia subgenus Hypotrachyna are absent from the East African sea
coast and lowlands below c. 800m altitude (Krog & Swinscow, 1979), the subgenus
Amphigymnia is represented there by more than a dozen species. They grow partly in
mangroves at sea level and partly in the coastal forest and low coastal hills up to c. 500 m. All
are drought tolerant, and several of them are found also in the upland savannas. The coastal

PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 151

Table 5 Chemical properties of species with soralia.

lee gyr nib ale stc nst sal pso gib pc fpc ech fat pgm atr usn

P. aprica

x . . .x

P. araucariarum

XXX.

P. austrosinensis

x x

P. bangii

x x

P. cooperi

x x

P. cristifera

x x

P. defecta

x x .

P. dilatata

x . x x xx

P. direagens

....x x.

P. gardneri

X ~+~ X

P. grayana

X . X

P. hababiana

-+- . x.

P. indoafra

XX X.

P. lobulascens

. X X.

P. louisianae

X X

P. parahypotropa

X X . X . . . . X

P. pardii

XX.. . X .

P. perlata

X X

P. permutata

X X X.

P. pilosa

X X .

P.poolii ... x x .

P. praesorediosa x . x

P. pseudo grayana xx.. . x.

P. rava x x x

P. reticulata

-H -4- +

P. rimulosa
P. sancti-angelii

X X .
X X .

P. subarnoldii

P. subschimperi
P. subsumpta
P. umbrosa

X... X.

. . . X.
... 4- .
X X... . X.

Only substances of diagnostic importance are included, lee = lecanoric acid, gyr = gyrophoric acid, nib =
norlobaridone, ale = alectoronic acid (including a-collatolic acid), stc = stictic acid, nst = norstictic acid, sal =
salazinic acid, pso = psoromic acid, glb = galbinic acid, pc = protocetraric acid, fpc = fumarprotocetraric acid,
ech = echinocarpic acid, fat = fatty acids, pgm = pigments, atr = atranorin, usn = usnic acid. Symbols: x = constant; + =
low concentrations; = occasionally present (accessory substance) or present in a chemical strain.

Table 6 Coastal species of Parmelia subgenus Amphigymnia in East Africa, arranged by the
lowest altitude at which they occur, and showing the altitudinal range in metres.

P. kwalensis P. gardneri 0-2000

P. pigmentifera 0-70 P. uberrima 0-2000

P. parahypotropa 0-300 P.tinctorum 0-2700

P. aldabrensis 0-1000 P. reticulata 0-3000

P.zollingeri 0-1000 P. dilatata 300-900

P.durumae 0-1100 P. cristifera 300-1450

P.hololoba 0-1800 P. sulphurata 480

152 H. KROG & T. D. V. SWINSCOW

Table 7 Photophilous, drought tolerant inland species of Parmelia subgenus Ampkigymnia
in East Africa, arranged by the lowest altitude at which they occur, and showing the altitudinal
range in metres. See Table 6 for additional species.

P. leonis

700-1400

P. grayana

1160-1500

P. praesorediosa

700-1800

P. rava

1250-1800

P. hababiana

800-2650

P. pilosa

1 300-2000

P. andina

900-2400

P. subtinctoria

1 300-2300

P. maclayana

900-2200

P. pseudograyana

1400-1900

P. pool ii

900-2300

P. indoqfra

1450-2600

P. abessinica

900-2600

P. xanthina

1500

P. vivida

920

P. stuhlmannii

1500-2100

P. aprica

1000

P. cooperi

1 500-3000

P. louisianae

1000

P. pardii

1650-1750

P. tsavoensis

1000

P. hanningtoniana

1700

P. soyauxii

1000-2100

P. defecta

1750-2100

P. sancti-angelii

1000-2800

P. jacarandicola

1900

P. austrosinensis

1000-3000

P. taitae

1980-2050

species, with their altitudinal ranges, are shown in Table 6; they include, for example, the
widespread, photophilous P. tinctorum and the ubiquitous P. reticulata.

Drought tolerant photophilous species which do not reach the coast are found in savannas,
bushed grassland, well lit sites in the montane forests, and artificial habitats such as parks,
plantations, town avenues, and roadside trees and gardens (Table 7).

Species with a somewhat higher moisture requirement are found in the more shaded sites
in the lower montane forests and those parts of the inselbergs which are regularly influenced
by mist. They often grow together with savanna species, but are here included among those
of the montane forests when they are largely absent from drier habitats. Species that are more
exacting in their moisture requirements are often best developed in the upper montane forest
where the rainfall is higher and more equally distributed throughout the year, especially on
the wetter slopes of the mountains. In the lower part of the alpine zone, above c. 3500 m,
there is still ample moisture, combined with high light intensity and low night temperatures.
Only four of our species, P. euneta, P. lobulascens, P. nilgherrensis, and P. subschimperi,
extend through the ericaceous zone and into the low alpine zone.

Table 8 Montane and low alpine species of Parmelia subgenus Amphigymnia in East Africa,
arranged by the lowest altitude at which they occur, and showing the altitudinal range in
metres. Parmelia reticulata is an additional species, included in Table 6.

P. subarnoldii

500-3000

P. crinita

1400-3400

P. pseudocrinita

700-2600

P. direagens

1 500-2600

P. planatilobata

800-1000

P. bangii

1 500-3200

P. amaniensi.s

800-1150

P. cryptoxantha

1550-2100

P. eciliata

800-1200

P. euneta

1600-3750

P. ultralucens

900

P. subcolorata

1700-2500

P. inexpectata

1100

P. rimulosa

1 800-2900

P. permutata

1100-2400

P. subisidiosa

1800-3200

.-*. umbrosa

1300-2100

P. lobulascens

1800-3600

P. erubescens

1400

P. subschimperi

1800-3600

P. subsumpta

1400-2100

P. araucariarum

2000-2100

P. lophogena

1400-2400

P. mellissii

2000-2400

P. cetrata

1400-2600

P. nilgherrensis

2000-3600

P. perlata

1400-3100

PARMEL1A (AMPHIGYMNIA) IN EAST AFRICA 153

Table 9 Species of Parmelia subgenus Amphigymnia which are not known outside
Africa.

P. aldabrensis

P. kwalensis

P. stuhlmannii

P. amaniensis

P. leonis

P. subcolorata

P. aprica
P. cryptoxantha
P. defecta
P. durumae
P. hololoba

P. lophogena
P. pardii
P. pigmentifera

P. pseudograyana
P. rava

P. subschimperi
P. taitae
P. tsavoensis
P. uberrima
P. umbrosa

P. inexpectata
P. jacarandicola

P. rimulosa
P. soyauxii

P. vivida

The species of the montane forests and the lower part of the alpine zone are shown in
Table 8. (In addition comes P. reticulata, included in Table 6 as one of the species that reach
the coast.) The subgenus Amphigymnia is not represented in the upper parts of the alpine
zone, above c. 4000 m altitude.

Of the 70 Amphigymnia species here recorded for East Africa, 26 species, or nearly 40 per
cent, are not yet known outside Africa (Table 9). Among the remaining species, 35 are found
also in the Americas; some of them are so far known only from Africa and the Americas,
namely Parmelia abessinica, P. araucariarum, P. bangii, IP. eurysaca, P. hanningtoniana,
IP. louisianae, P. pilosa, and P. subsumpta. Thirty-three are known from Asia, and of these
species the following are known only from Africa and Asia: Parmelia cooperi, P. direagens,
IP. euneta, P. indoafra, P. lobulascens, P. nilgherrensis, P. planatilobata, and P.
pseudocrinita. Twenty-three of the East African species occur in the Australian region; they
are all known also from other regions. Only five cosmopolitan, mainly temperate species are
common to East Africa and Europe; they are P. austrosinensis, P. cetrata, P. crinita, P.
perlata, and P. reticulata.

Discussion
Circumscription of the species

In his world monograph Hale (1965) discussed the relative taxonomic value of various
morphological and chemical characters on the basis of their degree of association with each
other. He found that cilia had a positive or negative correlation with more than half of the
characters considered, and deduced that they were of importance at species level. Similarly,
maculae, a white marginal zone on the under side, an entirely brown under side, and a
perforated apothecial disc were found to be valid characters, whereas spore size and
ornamentation of the thalline exciple were not. Conidia were assumed to have no
importance and were not investigated. Chemical strains were accepted in a limited way;
many of them have been regarded as species in subsequent publications by the same author.
It should be borne in mind that Hale's monograph appeared before the advent of TLC, so
that knowledge of the chemical properties of the species was then in many cases inferior to
that reached during the last decade.

We have had the advantage of studying the East African Amphigymnias in the field and of
collecting numerous specimens of the commoner species from sites with differing ecology.
This has enabled us to evaluate the morphological variation within the species in the light of
environmental factors, and to form an opinion on the relative taxonomic value of the various
characters as expressed in the species of our area. As might be expected, we have observed
that species with a great ecological amplitude usually show a much greater morphological
variation than species with a limited range. In a moist, shady habitat in an area with favour-
able temperatures the thallus is often membranaceous, greenish in colour, has long, slender

1 54 H. KROG & T. D. V. SWINSCOW

cilia and distinct maculae, whereas plants growing in a well lit site, subjected to drought or
low temperatures conducive to slow growth, in general have a coriaceous thallus, a grey
colour more or less tinged with brown, short, stunted and coarse cilia, and less conspicuous
maculae. At their extremes such plants look very different and have often been interpreted as
distinct species.

Taking the environmentally induced variation into account, we find that most of the
characters listed by Hale (1965) are of value at species level, but with certain reservations. A
normally ciliate species, such as Parmelia nilgherrensis, may rarely lack cilia, while a few
normally eciliate species, such as P. cristifera and P. gardneri, may produce rudimentary
cilia in the lobe axils. A strongly maculate species clearly differs from a species in which
maculae are always indistinct or absent, but in a normally maculate species it is possible for
saxicolous specimens with a coriaceous thallus to develop very few maculate areas. Presence
or absence of a perforated apothecial disc is fairly constant in most species but varies greatly
in some. In P. euneta and P. maclayana, for example, the apothecia may all be perforate, or
all imperforate, or both types may be present on a single specimen, so that the character is of
limited diagnostic value in these two species. A white marginal zone on the under side is
characteristic of some species, such as P. leonis and P. uberrima, but occurs only in some
specimens of, for example, P. abessinica and P. andina. In the P. erubescens species
complex, as here delimited, the colour of the under side varies from black through brown to
ivory.

The various types of vegetative propagules are usually considered to be species specific,
but in some species there is a complete intergradation between granular soredia, sorediate
isidia, non-sorediate isidia, and dactyls. This is the case in, for example, P. lophogena and P.
mellissii. We have also found that P. tinctorum is subject to variation in its vegetative
propagules, which range from simple or branched, slender, cylindrical isidia to coarse,
branched isidia and closed or open dactyls. Both isidia and dactyls may at times disintegrate
partly or wholly into granular soredia. We have chosen to regard P. pseudotinctorum (coarse
isidia and dactyls) as conspecific with P. tinctorum (slender isidia), since the variation is
largely continuous and both morphotypes have the same type of spores and conidia, and
mainly the same ecological requirements. If a distinction were to be maintained, we believe
that it should be at infraspecific level.

We do not agree with Hale (1965) that spore size alone is not a valid character. In our
experience there is little variation in spore length within species whose spores are below
20 um long, so that two subgroups whose spores are respectively above and below about
16 yum can be recognized (Table 1). In species with medium sized spores, on the other hand,
the spores may vary between 20 and 30 um, often with a maximum around 22-25 //m, while
in those with large spores, the spores may in extreme cases show variation between 25 and
40 //m, usually with a maximum around 28-30 //m. There is in our area a clear discontinuity
between species with small spores and those with large spores, and we regard this character as
species specific.

There is at present no common agreement on the function of the lichen conidia. A sexual
function for them has not been conclusively proved, but nor has their asexual nature been
firmly established. Regardless of their function, their constancy within morphologically well
delimited species and partly also within supraspecific units show that a taxonomic value
should be assigned to them that would normally not be bestowed upon vegetative
propagules. In agreement with, for example, Henssen & Jahns (1973) we consider the shape
of the conidia to be species specific, and like Moberg (1977) we believe that they are of
taxonomic significance also at the supraspecific level.

In Amphigymnia we have noted a strong correlation between imperforate apothecia and
sublageniform conidia on the one hand and perforate apothecia and filiform conidia on the
other, although there are exceptions to this rule (Table 10). Parmelia abessinica, for
example, has perforate apothecia but weakly sublageniform conidia. In P. maclayana, in
which the apothecia may be perforate or not, the conidia are sublageniform, whereas in P.
euneta, with similarly variable apothecia, they are filiform.

PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 155

Table 10 Number of species with either filiform or
sublageniform conidia, arranged by presence or absence
of perforate apothecial discs.

Conidia

Apothecia filiform sublageniform

perforate 24 4

imperforate 1 1 3

Table 11 Epithets applied to various chemotypes in the Parmelia
euneta Parmelia subschimperi species pair.

Substances No soralia or isidia Soralia

gyrophoric acid euneta neolobulascens

(1877-78) (1975)

gyrophoric acid composita subcompositum

norstictic acid (1972) (1977)

gyrophoric acid unnamed balensis

norlobaridone (1975)
norstictic acid ()

norlobaridone unnamed subschimperi

(1972)

norlobaridone spilota unnamed

norstictic acid (1973)

norstictic acid unnamed

In the delimitation of the species we assign greater taxonomic value to characters of spores
and conidia than we do to characters of the thallus. Many secondary species without
apothecia still produce pycnidia, and if conidia can be found they offer a valuable guide to
the correct interpretation of the species.

Chemical characters show no continuous variation, with the exception of accessory
substances which vary from abundance to absence within a given species and are commonly
regarded as of no taxonomic value. All other cases of chemical variation represent easily
registered discontinuities, and have therefore been regarded as species specific by some
authors. However, as with the morphological variation we have attempted to evaluate each
case of chemical variation individually on the basis of trends observed in our material.

Winnem (1975) clarified the chemical properties of the sorediate species P. direagens. She
found that it had three chemical strains: (1) psoromic acid, gyrophoric acid, and alectoronic
acid; (2) psoromic acid and gyrophoric acid; and (3) psoromic acid and alectoronic acid. The
psoromic acid, an extremely rare substance in Amphigymnia, is mainly connected with the
soralia in P. direagens, which is an unusual situation in Parmelia but known in certain other
genera, for example Usnea (Swinscow & Krog, 1979). In our opinion the presence of this
common denominator unites the morphologically uniform chemotypes and shows beyond
doubt that they merely represent chemical strains within a species. Apart from the presence
of psoromic acid, a depside and a depsidone of the orcinol series occur separately or jointly.

A parallel to this example is found in the chemical variation of the P. euneta P.
subschimperi species pair, although there is no common denominator present (Table 1 1 ).

156

H. KROG & T. D. V. SWINSCOW

The species comprise strongly maculate, ciliate plants with intermediate spores, filiform
conidia, and a montane to low alpine distribution. Winnem (1975) showed that norstictic
acid was accessory in this group and of no taxonomic importance. The remaining substances,
gyrophoric acid and norlobaridone, may occur separately or jointly. As in P. direagens this is
also a situation involving a depside and a depsidone of the orcinol series.

The P. amaniensis P. subarnoldii pair are ciliate, emaculate species with imperforate
apothecia, a strongly dentate-ciliate thalline margin, large, thick-walled spores, and
sublageniform conidia. The following chemical strains occur in each species: (1)
protocetraric acid and alectoronic acid; (2) protocetraric acid and a-collatolic acid,
alectoronic acid; and (3) protocetraric acid and protolichesterinic acid (Table 12). Spores,
conidia, and thallus morphology afford several valid characters by which to define this
species pair, and chemically protocetraric acid is a common denominator. We have here a
substitution situation involving two closely related orcinol depsidones and a fatty acid.

Another example involving an orcinol depsidone and a fatty acid is that of the P.
abessinica P. hababiana species pair, each of which is known in the following chemical
strains: (1) norlobaridone; (2) norlobaridone and protolichesterinic acid; and (3) proto-
lichesterinic acid.

A group of mainly Australian and South American species poses problems related to
variation in both cortical and medullary substances. All specimens have imperforate or
narrowly perforate apothecia, small spores, and a distinctly maculate upper side. The under-
side varies from black in the centre with a broad, brown or mottled marginal zone through
uniformly brown to ivory. The rhizines are usually dimorphous, and rhizines and papillae
often, but not always, reach the lobe margins. The majority of species have negligible
quantities of atranorin in the cortex, often in too small amounts to be detected with TLC, but

Table 12 Epithets applied to various chemotypes in the Parmelia
amaniensis Parmelia subarnoldii species pair.

Substances

No soralia or isidia

Soralia

protocetraric acid
alectoronic acid

amaniensis
(1926)

deflectens
(1979)

protocetraric acid
a-collatolicacid

pachvspora
(1965)

unnamed

protocetraric acid
protolichesterinic acid

unnamed

subarnoldii
(1961)

Table 13 The oldest epithets for the various chemotypes in the Parmelia
erubescens species complex.

Substances

No soralia or isidia

Sorediate

Isidiate

salazinicacid

erubescens
(1877-78)

subsumpta
(1869)

unnamed

norlobaridone

recipienda
(1885)

conferendum
(1977)

haitiensis
(1959)

salazinicacid
norlobaridone

unnamed

reitzii
(1977)

subtinctoria
(1930)

PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 157

some Central and South American species, such as P. subcaperata Krempelh. and
Parmotrema neotropicum Kurok., have copious amounts of usnic acid together with
medium amounts of atranorin. In the Parmeliaceae there is no precedent for accepting such
large variation in cortical substances within a species. The presence of usnic acid in the
cortex is here correlated with salazinic acid in the medulla, while species with only traces of
atranorin in the cortex may produce salazinic acid and norlobaridone separately or jointly.
That the variation in cortical substances does not parallel that of the medulla is a further
reason for regarding the species that produce usnic acid as distinct from those with only
traces of atranorin (referred to below as the P. erubescens complex).

The oldest epithets available for each chemotype within the P. erubescens species
complex are listed in Table 13, providing an example of chemical variation involving an
orcinol depsidone and a /?-orcinol depsidone. As in the examples mentioned above we
believe that these chemotypes also should be regarded as chemical strains. However, since
the relevant primary species are poorly represented in our area, they are in need of further
study, especially with regard to their conidial characters, in order to ascertain their relation-
ship with each other. We have therefore chosen to treat the chemotypes occurring in our area
as chemical strains, but do not formally reduce the extra-African chemotypes to synonymy
at this point.

When the chemical relationship seemed more complicated than in the examples given
above, or we felt that we had studied insufficient material, we have hesitated to apply the
theory of chemical strains until further evidence can be produced. Some examples follow.

The morphological differences between the two species pairs P. euneta P. subschimperi
and P. nilgherrensis P. lobulascens are few and insignificant; they are mainly that the P.
nilgherrensis pair has a higher frequency of perforate apothecia and on the whole a more
coriaceous thallus and more irregularly incised lobes. However, the two species pairs differ
both in basic chemistry (gyrophoric acid and norlobaridone versus alectoronic acid) and in
accessory substances (norstictic acid versus gyrophoric acid), while we have not yet come
upon specimens with a combination of the two orcinol depsidones alectoronic acid and
norlobaridone. In view of these considerable chemical differences, which are correlated with
a few weak morphological traits, we regard the two species pairs as distinct from each other.

Parmelia inexpectata differs from P. amaniensis in that it lacks protocetraric acid (the
common denominator in the P. amaniensis complex), while it produces an array of orcinol
depsidones (lividic acid and associated substances), none of which occur in P. amaniensis.
Since P. inexpectata is known from very few collections, we feel that we have insufficient
grounds for including the species in P. amaniensis for the time being, especially since it
differs in several chemical characters.

The two sorediate species P. grayana (protolichesterinic acid) and P. pseudograyana
(fumarprotocetraric and protocetraric acids) are morphologically virtually indistinguishable.
The supposed parent morph of P. pseudograyana, P. taitae, is known only with the /?-orcinol
depsidones. This species pair appears to be restricted to Africa, while P. grayana is widely
distributed. We should like to see much more material of/ 3 , taitae, and preferably also fruit
bodies in P. pseudograyana, before we make a final decision on the relationship between the
two secondary species.

It is difficult to assess the relationship between chemically discordant secondary species
when the corresponding primary species are not known. In such cases it is preferable to
regard the species as distinct until further evidence becomes available.

The underlying causes of chemical diversity in lichens are not known. Theoretically, a
single mutation influencing the biogenetic pathway at an early point may change the
chemical end product considerably. If sexual reproduction is assumed to take place among
lichens, cross fertilization between two individuals with different chemical properties might
result in the spores from one apothecium giving rise to plants of different chemotypes. If one
further assumes that some gene exchange takes place, a new chemotype characterized by a
combination of substances may result. Chemotypes originating in that manner would
probably not be regarded as distinct species.

158 H. KROG & T. D. V. SWINSCOW

It is possible that conidia rarely function as spermatia in lichens, or, if they do, that
fertilization usually takes place between conidia and ascogones produced on the same
thallus. However, if cross fertilization does occur under certain circumstances, the
conditions in the tropical rain-forests would be conducive to it. There the various lichens are
crowded and often multilayered on tree trunks and branches, and they are subjected almost
daily to tropical showers and water condensed from heavy mist, which could carry conidia
from one plant to another with minimal risk of desiccation.

Generic concept and infrageneric classification

Genus Parmelia s. lat. has in recent years been divided into a number of genera. As we have
stated earlier (Krog & Swinscow, 1979), we believe some of these genera to be based on
characters of little importance at the generic level, the result being obscure delimitation
between some neighbouring genera. We have therefore been reluctant to accept the new
genera proposed, although we believe that a satisfactory division of Parmelia s. lat. into
several genera will eventually be reached.

According to Hale's main division of Parmelia s. lat., Parmelia s. str. has a paraplec-
tenchymatous upper cortex and a non-pored epicortex frequently bearing pseudocyphellae,
while the other segregates possess a palisade plectenchymatous upper cortex and a pored
epicortex without pseudocyphellae (Hale, 19760). We agree in this major division, but have
not yet reached a decision on the interpretation of the residual genera in Parmelia s. lat.
Although the generic name Parmotrema primarily applies to the group of lichens that are
dealt with here, we hesitate to adopt it until the genus has been satisfactorily circumscribed,
notably in relation to Parmelia subgenus Cydocheila sensu Krog & Swinscow (1979). The
conidia may prove to be of considerable value in the circumscription of the parmelioid
genera.

For a subdivision of Amphigymnia, various characters have been employed. Vainio (1890)
relied on the colour of the upper cortex (presence or absence of usnic acid) when he proposed
the subsections Subflavescens Vainio and Subglaucescens Vainio. Gyelnik (1932) regarded
Vainio's subsections as sections and further subdivided them into each two subsections
(Eciliatae Ciliatae, Eciliolae Ornaticolae) on the basis of presence or absence of cilia.

Hale (1965) found that the presence or absence of cilia was a character of greater
importance at section level than the colour of the upper cortex. He proposed the following
scheme:

Sect. Amphigymnia [lobe margins eciliate or rarely with sparse cilia in the axils].
Sect. Subflavescentes (Vainio) Gyelnik [margins and apices of lobes distinctly ciliate].
Subsect. Subflavescentes [upper cortex emaculate].
Ser. Subflavescentes [usnic acid present],
Ser. Emaculatae Hale [usnic acid absent].
Subsect. Ornaticolae Gyelnik [upper cortex maculate].
Ser. Subpallidae Hale [under side brown, rhizinate to the margins].
Ser. Ornaticolae [under side black in the centre, with a commonly white or mottled,
naked marginal zone].

We are undecided whether to regard the presence of cilia as a valid character for infrageneric
classification in Amphigymnia, since its correlation with other characters is not yet clear. In
ser. Subpallidae there appears to be correlation between a strongly maculate upper cortex,
rhizine morphology, and chemical traits, whilst in most other cases the presence of maculae
seems uncorrelated with other characters. We find the presence or absence of usnic acid in
the cortex to be of no taxonomic value above species level. However, a classification of the
amphigymnioid species can only be settled on the basis of a world wide study. Before that is
possible, the delimitation of the genus Parmotrema must be conclusively agreed upon.

PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 159

The East African Amphigymnia species

Key to the species

la Soralia and isidia absent 2 (p. 159)

Ib Soralia or isidia present 46

46a Isidia (including sorediate isidia and dactyls) present , 47 (p. 161)

46b Isidia absent. Soralia present 62 (p. 162)

Soralia and isidia absent

2a (la) Marginal cilia absent 3

2b Marginal cilia present 6

3a (2a) Thallus yellow (usnic acid present) 68. P. vivida(p. 223)

3b Thallus grey (usnic acid absent) 4

4a (3b) Apothecia imperforate. Medulla C , PD+ orange-red (protocetraric acid)

70. P. zollingeri (p. 225)
4b Apothecia perforate. Medulla C+ red, PD (lecanoric acid) 5

5a (4b) Plant saxicolous, strongly attached. Lobes less than 1 cm broad 53. P. soyauxii (p. 2 1 0)
5b Plant normally corticolous, loosely attached. Lobes more than 1 cm broad 4. P. andina (p. 168)

6a (2b) Medulla pigmented pale yellow, ochraceous, or salmon pink, pigment K ... 7
6b Medulla white, at most with patches of an ochraceous, K+ purple pigment near the

lower cortex 8

7a (6a) Apothecia imperforate, with a dentate-ciliate thalline margin. Medulla UV+

(undetermined substances). Coastal species .... 42. P. pigmentifera (p. 201)

7b Apothecia perforate, with a smooth, eciliate thalline margin. Medulla UV

(gyrophoric acid). Upland species 56. P. subcolorata (p. 2 1 3)

8a (6b) Upper cortex with a reticulate pattern of maculae and cracks. Rhizines in part

squarrose. Salazinic acid present 9. P. cetrata (p. 1 72)

8b Upper cortex without a reticulate pattern of maculae and cracks. Rhizines not

squarrose. Salazinic acid present or absent . 9

9a (8b) Underside with a distinct, white marginal zone 10

9b Underside with a brown or mottled marginal zone 14

lOa (9a) Apothecia perforate. Medulla UV- 11

lOb Apothecia mainly imperforate. Medulla UV+ 13

lla (lOa) Underside almost entirely white. Medulla C-, KC (protolichesterinic acid)

31. P. leonis(p. 191)
lib Underside black in the centre, white peripherally. Medulla C+ or C , KC+ or KC . 12

1 2a (lib) Lobe margins flat or revolute. Medulla C+ red (lecanoric acid) 26. P. hololoba (p. 1 86)
12b Lobe margins ascending. Medulla C (norlobaridone and/or protolichesterinic acid)

1 . P. abessinica (p. 165)

1 3a (1 Ob) Spores less than 20 //m long. Only alectoronic acid present 65. P. uberrima (p. 220)

1 3b Spores more than 20 //m long. Both alectoronic and or-collatolic acids present

17. P. durumae (p. 178)

14a (9b) Apothecia present 15

14b Apothecia absent 30

160 H. KROG & T. D. V. SWINSCOW

15a ( 14a) Spores more than 20 //m long 16

1 5b Spores less than 20 //m long 21

16a ( 15a) Upper cortex emaculate. Apothecia imperforate 17

16b Upper cortex distinctly maculate. Apothecia perforate or imperforate 20

1 7a (1 6a) Apothecia with an even, eciliate thalline margin. Stictic acid present

18. P. eciliata (p. 180)
1 7b Apothecia normally with a dentate-lobulate and/or ciliate thalline margin. Stictic acid

absent -18

1 8a (1 7b) Spores 22-26 /^m long. Alectoronic and a-collatolic acids present 17. P. durumae (p. 1 78)
18b Spores 2 5-2 8 (40) //m long. Chemistry various 19

1 9a (1 8b) Medulla with substances in the lividic acid complex . . . 28. P inexpectata (p. 1 88)
19b Medulla with protocetraric acid (usually in combination with alectoronic, a-collatolic,

or protolichesterinic acid) 3. P. amaniensis (p. 167)

20a ( 1 6b) Medulla U V+ (alectoronic acid, a-collatolic acid, gyrophoric acid)

37. P. nilgherrensis (p. 197)
20b Medulla UV (gyrophoric acid and/or norlobaridone, norstictic acid) 20. P. euneta (p. 1 8 1 )

2 la (15b) Upper cortex distinctly maculate 22

21b Upper cortex emaculate or faintly maculate 25

22a (2 la) Thalline exciple with coarse, isidioid protuberances. Medulla C+ rose, PD

(gyrophoric acid) 25. P. hanningtoniana (p. 185)

22b Thalline exciple smooth to rugose, without isidioid protuberances. Medulla C , PD+

orPD- 23

23a (22b) Rhizines dimorphous. Salazinic acid and norlobaridone present

19. P. erubescens (p. 180)
23b Rhizines uniform. Chemical properties otherwise 24

24a (23b) Underside brown. Medulla PD+ orange (stictic and norstictic acids). Coast and

lowland species 2. P. aldabrensis (p. 166)

24b Underside black in the centre, white, mottled, or brown peripherally. Medulla PD

(norlobaridone and/or protolichesterinic acid). Upland species 1. P. abessinica (p. 165)

25a (21b) Cortex PD+ sulphur yellow near the apothecia (psoromic acid present). Medulla

C+ rose (gyrophoric acid) 29. P. jacarandicola (p. 190)

25b Cortex at most PD+ pale yellow (psoromic acid absent). Medulla C+ or C . ... 26

26a (2 5b) Medulla UV+ (alectoronic acid) 35. P. maclayana (p. 195)

26b Medulla UV (alectoronic acid absent) 27

27a (26b) Plant saxicolous. Apothecia imperforate. Medulla PD+ orange-red (fumarproto-

cetraric acid) 62. P. taitae (p. 2 1 7)

27b Plant normally corticolous. Apothecia perforate. Medulla PD+ or PD (fumarproto-

cetraric acid absent) 28

28a (27b) Central lobes laciniate. Medulla PD+ orange (salazinic acid) 21 . P. eurysaca (p. 1 82)
28b Central lobes not laciniate. Medulla PD- 29

29a (28b) Lobe margins flat or revolute. Medulla C+ red (lecanoric acid) 26. P. hololoba (p. 1 86)
29b Lobe margins ascending. Medulla C (norlobaridone and/or protolichesterinic acid)

(norlobaridone and/or protolichesterinic acid). Upland species 1 . P. abessinica (p. 1 65)

30a (14b) Medulla PD+ orange or red - 31

PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 161

30b Medulla PD (but cortex may be PD+ sulphur yellow, see 42b) 37

3 la (30a) Plant saxicolous. Fumarprotocetraric acid present . . 62. P. taitae(p. 217)
31b Plant normally corticolous. Fumarprotocetraric acid absent 32

32a (3 Ib) Upper cortex distinctly maculate 33

32b Upper cortex emaculate or faintly maculate 35

33a (32a) Salazinic acid and norlobaridone present ... 19. P. erubescens (p. 1 80)
33b Salazinic acid absent, norlobaridone present or absent . 34

34a (33b) Montane forest species. Norstictic acid present in combination with gyrophoric

acid and/or norlobaridone 20. P. euneta(p. 181)

34b Coastal species. Norstictic acid present in combination with stictic acid

2. P. aldabrensis (p. 166)

35a (32b) Protocetraric acid present (usually in combination with alectoronic, or-collatolic,

or protolichesterinic acid) 3. P. amaniensis (p. 167)

35b Protocetraric acid absent 36

36a (3 5b) Salazinic acid present 21. P. eurysaca(p. 182)

36b Stictic acid present 18. P. eciliata (p. 1 80)

37a (30b) Upper cortex distinctly maculate 38

37b Upper cortex emaculate or faintly maculate 41

38a (3 7a) Species of dry, well lit upland habitats 39

38b Species of the montane forests and the alpine zone . 40

39a (38a) Medulla C+ rose (gyrophoric acid) .... 25. P. hanningtoniana (p. 185)

39b Medulla C (norlobaridone and/or protolichesterinic acid) 1. P. abessinica (p. 165)

40a (38b) Medulla U V+ (alectoronic acid, <ar-collatolic acid, gyrophoric acid)

37. P. nilgherrensis (p. 197)
40b Medulla UV (gyrophoric acid and/or norlobaridone) . . . 20. P. euneta(p. 181)

41a (37b) Medulla C+ rose or red - 42

41b Medulla C- 43

42a (4 1 a) Lecanoric acid present 26. P. hololoba(p. 186)

42b Gyrophoric acid present (combined with psoromic acid in the cortex of some lobes)

29. P.jacarandicola(p. 190)

43a (4 1 b) Substances in the lividic acid complex present . . 28. P. inexpectata (p. 1 88)
43b Substances in the lividic acid complex absent 44

44a (43b) Medulla UV (norlobaridone and/or protolichesterinic acid) 1. P. abessinica (p. 165)
44b Medulla UV+ (alectoronic acid, a-collatolic acid) 45

45a (44b) Plant corticolous or saxicolous. Upland species above c. 1 000 m altitude

35. P. maclayana (p. 195)
45b Plant corticolous. Coastal and lowland species below c. 1 000 m altitude

17. P. durumae (p. 178)

46a (Ib) Isidia (including sorediate isidia) or dactyls present . 47

46b Isidia and dactyls absent. Soralia present 62

Isidia (including sorediate isidia and dactyls) present

47a (46a) Marginal cilia absent 48

162 H. KROG & T. D. V. SWINSCOW

47b Marginal cilia present 51

48a (47a) Medulla PD+ orange-red (protocetraric acid) 71. P. sp. A (p. 226)

48b Medulla PD- 49

49a (48b) Plant large, loosely fastened to tree bark or rock. Cylindrical isidia, granular

isidia, or dactyls present. Medulla C+ red (lecanoric acid) 63. P. tinctorum (p. 2 1 8)

49b Plant small, strongly fastened to rock. Only dactyls or coarse isidia present. Medulla C+

orC- 50

50a (49b) Medulla C+ red (lecanoric acid) 54. P. stuhlmannii (p. 21 1)

50b Medulla C- (physodic acid) 64. P. tsavoensis (p. 220)

5 la (47b) Thallus yellow (usnic acid present) 69. P. xanthina (p. 224)

51b Thallus grey (usnic acid absent) 52

52a (51b) Upper cortex with a reticulate pattern of maculae and cracks. Rhizines in part

squarrose 57. P. subisidiosa (p. 214)

52b Upper cortex without a reticulate pattern of maculae and cracks. Rhizines not squarrose 53

53a (52b) Medulla pigmented pale to bright yellow or orange 54

53b Medulla white, at most with patches of an ochraceous, K+ purple pigment near the

lower cortex 55

54a (53a) Medulla pigmented bright yellow to orange throughout. Cylindrical isidia present.

Coastal species 61. P. sulphurata (p. 217)

54b Medulla pigmented pale yellow, pigment sometimes inapparent. Open dactyls present.

Lower montane forest species 13. P. cryptoxantha (p. 175)

55a (53b) Isidia flattened, dorsiventral. Gyrophoric acid present 44. P. planatilobata (p. 203)
55b Isidia cylindrical. Chemistry various 56

56a (5 5b) Upper side distinctly maculate 57

56b Upper side emaculate or faintly maculate 58

57a (56a) Rhizines dimorphous. Salazinic acid and norlobaridone present. Upland species

60. P. subtinctoria (p. 2 1 6)
57b Rhizines uniform. Stictic and norstictic acids present. Coastal species 30. P. kwalensis (p. 191)

58a (56b) Thallus coriaceous. Isidia mainly laminal, never becoming sorediate . . . 59

58b Thallus usually membranaceous. Isidia mainly submarginal, often becoming sorediate-

granular 60

59a (58a) Medulla C+ rose, PD , UV- (gyrophoric acid) . 47. P. pseudocrinita (p. 205)

59b Medulla C , PD+ orange, UV+ intensely yellow (salazinic acid, lichexanthone)

66. P. ultralucens (p. 221)

60a (58b) Upper cortex usually continuous. Isidia rarely sorediate. Medulla PD+ orange

(stictic acid) v 11. P. crinita(p. 173)

60b Upper cortex fragile and flaking. Isidia often sorediate. Medulla PD 61

61a (60b) Medulla C+ rose, UV (gyrophoric acid) 33. P. lophogena(p. 194)

61b Medulla C ,UV+(alectoronic acid) 36. P. mellissii (p. 197)

Isidia absent. Soralia present

62a (46b) Marginal cilia absent 63

62b Marginal cilia present 73

PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 163

63a (62a) Plant saxicolous 64

63b Plant corticolous .67

64a (63a) Medulla PD+ orange-red (protocetraric acid) 39. P. pardii (p. 199)

64b Medulla PD- .65

65a (64b) Medulla C (fatty acids) 46. P. praesorediosa (p. 205)

65b Medulla C+ red (lecanoric acid) 66

66a (65b) Thallus small, coriaceous, strongly attached. Upper cortex emaculate, shiny

14. P. defecta (p. 176)
66b Thallus usually large, relatively thin, loosely attached. Upper cortex faintly maculate,

often matt 7. P. austrosinensis (p. 171)

67a (63b) Thallus yellow or yellowish grey (usnic acid present). Medulla PD+ orange-red

(protocetraric acid) 68

67b Thallus pale grey (usnic acid absent). Medulla PD+ or PD 70

68a (67a) Thallus bright yellow to yellow-green. Atranorin absent . . . 5. P. aprica (p. 1 69)
68b Thallus yellowish grey. Atranorin present - 69

69a (68b) Echinocarpic acid and various unknowns present ... 15. P. dilatata (p. 1 77)
69b Echinocarpic acid and unknowns absent 49. P. rava (p. 207)

70a (67b) Medulla C+ red (lecanoric acid) .... 7. P. austrosinensis (p. 171)
70b Medulla C- 71

7 la (70b) Medulla PD- (fatty acids) 46. P. praesorediosa (p. 205)

71b Medulla PD+ orange or red 72

72a (7 Ib) Medulla K+ red (salazinic acid) 12. P. cristifera (p. 173)

72b Medulla K+ sordid brown (protocetraric acid) ..... 22. P. gardneri(p. 183)

73a (62b) Upper cortex fragile and flaking. Soralia erupting in a pustular fashion or more or

less mixed with isidia 74

73b Upper cortex continuous. Soralia rarely pustular, isidia absent 77

74a (73a) Medulla pigmented pale yellow. Echinocarpic acid and fatty acids present

13. P. cryptoxantha (p. 175)
74b Medulla white. Chemistry otherwise 75

75a (74b) Medulla PD+ orange, UV- (stictic acid) ....-- 8. P. bangii (p. 1 7 1 )
75b Medulla PD-, UV+(alectoronic acid) 76

76a (75b) Soralia pustular, without isidia 51. P. rimulosa(p. 209)

76b Soralia not pustular, but here and there interspersed with isidia . 36. P. mellissii (p. 197)

77a (73b) Medulla pigmented yellow, salmon pink, or ochraceous, pigment K . ... 78
77b Medulla white, at most with patches of an ochraceous, K+ purple pigment near the

lower cortex 79

78a (77a) Medulla C+ red in upper parts (gyrophoric acid) . . . . 41. P. permutata (p. 201)
78b Medulla C- (fatty acids) 6. P. araucariarum (p. 170)

79a (77b) Upper cortex with a reticulate pattern of maculae and cracks. Rhizines in part

squarrose 50. P. reticulata (p. 208)

79b Upper cortex without a reticulate pattern of maculae and cracks. Rhizines not squarrose 80

80a (79b) Underside with a distinct, white marginal zone 81

164 H. KROG & T. D. V. SWINSCOW

80b Underside with a brown or mottled marginal zone 84

8 la (80a) Lobes deeply divided, with sublinear laciniae. Medulla K+ red, PD+ orange

(norstictic, galbinic, and salazinic acids). Coastal species . 38. P. parahypotropa (p. 1 98)

81b Lobes more or less rounded, sublinear laciniae absent. Medulla K , PD . Inland

species 82

82a (81b) Underside black in the centre, white peripherally. Medulla UV (norlobaridone

and/or protolichesterinic acid) * 24. P. hababiana (p. 1 84)

82b Underside almost entirely white. Medulla UV+ or UV 83

83a (82b) Medulla UV+(alectoronic acid) 34. P. louisianae (p. 195)

83b Medulla UV- (protolichesterinic acid) 72. P. sp.B (p. 226)

84a (80b) Rhizines dimorphous, often extending to the margins 85

84b Rhizines uniform, rarely extending to the margins 86

85a (84a) Soralia marginal. Medulla PD+ orange, KC (salazinic acid) or PD , KC+ red

(norlobaridone) 59. P. subsumpta (p. 2 1 5)

85b Soralia laminal. Medulla PD-,KC- (fatty acids) .... . 43. P. pilosa (p. 202)

86a (84b) Upper cortex distinctly maculate - 87

86b Upper cortex emaculate or faintly maculate 88

87a (86a) Medulla UV+(alectoronic acid, or-collatolic acid, gyrophoric acid)

32. P. lobulascens (p. 192)
87b Medulla UV (gyrophoric acid and/or norlobaridone, norstictic acid)

58. P. subschimperi (p. 2 14)

88a (86b) Plant saxicolous 89

88b Plant normally corticolous 91

89a (88a) Medulla UV+(alectoronic acid) 45. P. poolii (p. 203)

89b Medulla UV- . 90

90a (89b) Medulla PD- (protolichesterinic acid) 23. P.grayana(p. 184)

90b Medulla PD+ orange-red (fumarprotocetraric and protocetraric acids)

48. P. pseudograyana (p. 206)

91a (88b) Medulla PD+ orange to red 92

91b Medulla PD (but soralia may be PD+ sulphur yellow) .96

92a (9 1 a) Protocetraric acid present 93

92b Protocetraric acid absent 95

93a (92a) Medulla C+ rose. Gyrophoric acid present 67. P. umbrosa (p. 222)

93b Medulla C-. Gyrophoric acid absent 94

94a (93b) Cilia well developed. Medulla UV+ (alectoronic or or-collatolic acid) or UV

(protolichesterinic acid) 55. P. subarnoldii (p. 212)

94b Cilia poorly developed, present only in the lobe axils. Medulla UV ( undetermined

fatty acids) 22. P. gardneri (p. 1 83)

95a (92b) Stictic acid present 40. P. perlata (p. 200)

95b Salazinic acid present 12. P. cristifera (p. 173)

96a (9 Ib) Soralia PD+ sulphur yellow (psoromic acid) . . . . 16. P. direagens (p. 178)

PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 165

96b Soralia PD or at most PD+ pale yellow (psoromic acid absent) .97

97a (96b) Medulla UV+(alectoronic acid) .... 45 p poolii(p 203)
97b Medulla UV- ' 98

98a (97b) Medulla C (norlobaridone and/or protolichesterinic acid) 24. P. hababiana (p 1 84)
98b Medulla C+ red 99

99a (98b) Lecanoric acid present 10. P. cooper! (p. 1 72)

99b Gyrophoric acid present 100

1 OOa (99b) Soralia often ciliate, soredia granular. Fatty acids present . . 33. P. lophogena (p. 1 94)
lOOb Soralia eciliate, soredia farinose. Fatty acids absent .101

lOla (lOOb) Norlobaridone present 27. P. indoafra(p. 187)

101 b Norlobaridone absent 52. P. sancti-angelii (p. 210)

Descriptions of the species

1. Parmelia abessinica Nyl. ex Krempelh.

Linnaea 41 : 140 (\%ll).Parmotrema abessinicum (Nyl. ex Krempelh.) Hale, Phytologia 28 : 334
(1974). Type: Abessinia, leg. Hildebrandt (G-holotype). [TLC (Winnem 1974) : norlobaridone,
loxodin, protolichesterinic acid, atranorin.]

Thallus corticolous, more or less coriaceous, adnate to loosely attached, pale grey. Lobes
0-5-1 cm broad, margins ascending, crenate, ciliate, cilia 0-5-1-5 (2) mm long. Upper side
faintly to distinctly maculate, rugose and cracked with age. Medulla white. Underside
rugose, sparsely rhizinate, black in the centre, with a brown, mottled, or white marginal
zone. Soralia and isidia absent. Apothecia common, often crowded, stipitate, up to 1 cm in
diameter, thalline margin smooth to crenulate, eciliate (or very rarely ciliate), disc perforate,
spores 1 5-18 (20) x 8-10 /zm. Conidia weakly sublageniform, 6-8 /zm long.

TLC: (1) norlobaridone, loxodin, protolichesterinic acid, atranorin, (2) norlobaridone,
loxodin, atranorin, (3) protolichesterinic acid, atranorin, (4) fatty acids of the reddenda
type, atranorin.

Two specimens, 3K 23/124 and Nordal INB 820, from Kenya and Tanzania respectively,
belong to a hitherto undescribed chemotype. They have the same general habit as P.
abessinica, spores in the same size range, and similar conidia, and differ only in producing
unnamed fatty acids of the reddenda type in the medulla. An additional specimen of this
chemotype has been seen from Zaire (LD). The same chemical properties are found in P.
mesotropa Mull. Arg., described from Paraguay, but that species lacks cilia and has
imperforate apothecia.

Parmelia glaucocarpoides Zahlbr., based on P. glaucocarpa Mull. Arg., non Ach., and
described on material from Madagascar (G holotype), was regarded by Hale (1965) as
synonymous with P. abessinica. Its morphology is similar, it has sublageniform conidia
5-6 /zm long, and contains protolichesterinic acid, but the spores are 25-27 (30) x 12-15 /im
and fairly thick-walled, showing that P. glaucocarpoides should be regarded as a species
distinct from P. abessinica.

Parmelia abessinica is a common and widespread species of dry, well lit sites at inter-
mediate altitudes, such as bushed grassland, artificial habitats, open hillsides, and the edge of
forests. It has been collected between 900 and 2600 m, but is most common below 2000 m.
Outside our area it is known from South and West Africa and Mexico (Hale, 1965).

Selected East African records

Ethiopia. Sidamo Province, Yirga Alem, Winnem 477/1 (O). Kenya. Eastern Province, Marsabit
District, Mt Marsabit, vicinity of Lake Paradise, 4K6/124: Kitui District, Mutomo, 50km NE of
Kibwezi, K21/2; Machakos District, Ol Doinyo Sapuk, 20 km SE of Thika, 2K 1/101, lava flow 5 km

166

H. KROG & T. D. V. SWINSCOW

NW of Kibwezi, 2K 22/101, 5K.2/31; Embu District, Izaak Walton Inn, K 53/8; Meru District,
between Chogoria and South Mara River, 3K 1 1/104. Rift Valley Province, Kajiado District, Chyulu
Hills, K 39/13; Samburu District, WSW slope of Warges, 4K 3/101; Laikipia District, Burguret,
4K 25/108; Nakuru District, Eastern Mau Forest 8 km ENE of Mau Narok, 4K 32/109, Lake Naivasha
Hotel, on east shore of lake, 3K 21/14. Coast Province, Taita District, near school W of Wundanyi,
2K 27/101. Western Province, Kakamega District, Kakamega Forest by River Ikuyawa, 4K 10/101.
Tanzania. Lake Province, Mosuma District, near Fort Ikomo Lodge, Sipman 6357 (herb. Sipman).
Northern Province, Arusha District, Arusha National Park, Juniper Hill, T 3/1 14, Mt Meru Crater,
T 5/162; Mbulu District, Babati, Sitari 541 (TUR). Southern Highlands Province, Njombe District,
12km SE of Njombe, Nordal INB 820 (O). Uganda. Kigezi District, Bufumbira County, Kisoro,
Travellers' Rest Hotel, U 19/3, 3U 59/10; Ruzhumbura County, W edge of Maramagambo Forest,
2U 1 1/6. Karamoja District, Matheniko County, near Sogolomon on Mt Moroto, 2U 36/53, NW of Mt
Moroto, 4 km SW of Nakiloro, 2U 38/6.

2. Parmelia aldabrensis Dodge.

Fig. 2

Ann. Mo. hot. Gdn 46 : 160 (\959).Parmotrema aldabrense (Dodge) Hale, Phytologia 28 : 334
(1974). Type: Aldabra Group, coll. Walter Fox 220 (BM holotype). [TLC (Winnem 1974):
norstictic acid.]

Thallus corticolous, adnate, pale grey to grey-green. Lobes 0-5-0*8 cm broad, deeply
divided, margins ascending, crenate, ciliate, cilia up to 2- 5 mm long. Upper side distinctly
maculate. Medulla white. Underside smooth or somewhat rugose, medium to dark brown,
rarely with a white mottled marginal zone, rhizines slender, scattered almost to the margins.
Soralia and isidia absent. Apothecia numerous, marginal and submarginal, often
eccentrically orientated on semitubular stipes, disc O'5-l (1*5) cm in diameter, with a small
perforation, thalline margin smooth or weakly crenate, occasionally ciliate, spores
1 3-1 7 x 5-8 ^m. Conidia filiform, 1 2-1 5 //m long.
TLC: norstictic acid, stictic acid.

All East African specimens contained stictic and norstictic acids, but the type specimen, as
well as two other specimens from the Aldabra Islands, Stoddart 992 (BM) and Fosberg 493 1 9
(S), had only norstictic acid. Morphologically the two populations are the same. No
apothecia or conidia were found in the holotype, but the collection Stoddart 992 had spores

Fig. 2 Parmelia aldabrensis Dodge, 3 K 30/ 1 1 (O). Rule = 1 cm.

PARMELIA (AMPHIGYMNIA) IN EAST AFRICA

167

12-16 x 6-7 //m and filiform conidia 10-12 /^m, which agrees well with the East African
material.

Parmelia preperforata W. Culb., described from Texas (US holotype), is chemically
concordant with the East African specimens of P. aldabrensis, has a maculate upper cortex,
marginal to submarginal aoothecia, and similar spores and conidia. It differs in having
broadly rounded lobes which are not particularly deeply divided, and an underside which is
black in the centre with a wide, pale ivory marginal zone.

Parmelia aldabrensis is locally abundant in mangroves by the Indian Ocean, where it
often grows convoluted around thinner branches. It has also been collected in the lowlands of
Tanzania up to 1000m. Outside our area it is known from the Aldabra Islands and
Madagascar (Hale, 1965).

East African records

Kenya. Coast Province, Kwale District, 2 km N of Gazi, K44/2, 1 1 1, 3K 30/45, 1 10; Kilifi District,
Mida Creek, 3K 27/104, 3K 29/106, Gedi ruins, 3K 25/104. Tanzania. [?Tanga Province]
Masaihochland, Wadiboma, Fischer 703 (G, type of P. hildebrandtii var. ciliata Mull. Arg.). Eastern
Province, Mzizima [Uzaramo] District, Kunduchi, Bjornstad AB 1942 (O); Morogoro District, Mindu
Forest Reserve, 4 May 1978, Dahl(O).

3. Parmelia amaniensis Steiner & Zahlbr.

Fig. 3

Bot. Jb. 60:526 (1926). [Tanzania] Im Regenwald bei Amani, Ost-Usambara, 800m, July
1909, leg. Brunnthaler (W holotype). [TLC: alectoronic acid, protocetraric acid, atranorin.]
Parmelia pachyspora Hale, Contr. U.S. natn. Herb. 36 (5) : 299 (1965). Parmotrema pachysporum
(Hale) Hale, Phytologia 28:338 (1974). Type: Angola, Huila, 10km N of Sa de Bandeira, 3
February 1960, leg. G. Degelius (herb. Degelius holotype). [TLC: a-collatolic acid, alectoronic acid
(+), protocetraric acid, atranorin.]

Thallus corticolous, membranaceous to more or less coriaceous, loosely attached, pale grey.
Lobes 0'5-l'5cm broad, crenate, ciliate, cilia 3-5 (7) mm long. Upper side emaculate.

Fig. 3 Parmelia amaniensis Steiner & Zahlbr., Ryvarden \ 1 632 (O). Rule = 1 cm.

168 H. KROG & T. D. V. SWINSCOW

Medulla white. Underside black, with a narrow brown or rarely white mottled marginal
zone, rhizines sparse. Soralia and isidia absent. Apothecia up to l'7cm in diameter,
submarginal on swollen stipes which become somewhat constricted with age, or marginal
and eccentrically situated on semitubular, convoluted lobes, thaline margin normally
lobulate-dentate and ciliate, disc imperforate, spores thick-walled, 25^0 x 15-18/^m.
Conidia weakly sublageniform, 5-8 //m long.

TLC: (1) alectoronic acid, protocetraric acid, atranorin, (2) cr-collatolic acid,
alectoronic acid (trace), protocetraric acid, atranorin, (3) protolichesterinic acid,
protocetraric acid, atranorin.

Overlooking the presence of protocetraric acid, Hale (1965) reduced P. amaniensis to
synonymy with P. subrugata Krempelh., a South American species with alectoronic and
a-collatolic acids. In the same work he described the new species P. pachyspora, with
protocetraric acid as its diagnostic medullary substance. Later, under the description of the
new species P. paradoxa (Hale, 1973), he noted that P. pachyspora was now known to
contain alectoronic acid in addition to protocetraric acid. When subjecting the type speci-
mens to TLC we found that P. amaniensis contained alectoronic acid and P. pachyspora
mainly a-collatolic acid in addition to protocetraric acid. Since anatomy, morphology, and
ecological requirements are the same for the two species, we here reduce P. pachyspora to
synonymy with P. amaniensis. We also include in the species some specimens which
produced protocetraric acid protolichesterinic acid in the medulla (Table 12).

In our area the typical strain has been collected in Tanzania, the pachyspora strain in
Uganda, and strain (3) in both countries.

Parmelia amaniensis has been found in the lower montane forests between 800 and
1 150 m altitude, with the exception of one collection from 2000 m. In East Africa we have
seen specimens from Tanzania and Uganda. Outside our area the species occurs in Angola,
Malawi, Mozambique, Sierra Leone, Zambia and Zimbabwe. It seems to be restricted to
Africa.

East African records

Tanzania. Eastern Province, Morogoro District, Nguru Mountains, E slope above Kwamanga village
near Mhonda Mission, Pocs & Mabberley 6397/H (herb. Pocs), Uluguru Mountains, N slope of
Bondwa above Morogoro, Pocs 6732 (herb. Pocs). Tanga Province, Tanga District, East Usambara
Mountains, Amani Forest Reserve, T. & S. Pocs 6100/AD-B (herb. Pocs); Lushoto District, in rain
forest by Amani, East Usambara, Brunnthaler 7/1909 (W holotype of P. amaniensis), Usambara Mts,
Amani, Karimi Estate (road towards Monga), Moberg 1485b (UPS). Uganda. Masaka District, Bukoto
County, Jubiya Forest, 3U 28/18, 3U 32/3, Lye L 604A (herb. Lye), N edge of Malabigambo Forest,
3U 25/4.

4. Parmelia andina Mull. Arg.

Revue mycol. 1 : 169 (\&19).Parmotrema andinum (Mull. Arg.) Hale, Phytologia 28 : 334 (1974).
Type: prope Cisne, Ecuador, 2800 m, leg. Ed. Andre 4324 bis (G holotype, BM isotype). [TLC
(Winnem 1975): lecanoric acid, atranorin.]

Parmelia modesta Hue, Bull. Soc. Bot. France 63, Mem. 6 (28) : 6 (1916). Type: Afrique equatoriale
anglaise, 1912, leg. Vie de Poncins, (PC holotype). [TLC: lecanoric acid, atranorin.]

Thallus corticolous or rarely saxicolous, coriaceous, loosely attached, pale grey, medium
grey, or grey-green. Lobes up to 2 cm broad, rounded, entire or crenate, eciliate. Upper side
usually matt, faintly to distinctly maculate (especially in the vicinity of the apothecia),
pitted, folded, or rugose towards the centre. Medulla white. Underside black, matt and
rugose, with a white, brown, or mottled marginal zone, rhizines short, black in central parts,
pale brown or white peripherally, unevenly distributed. Soralia and isidia absent. Apothecia
submarginal to laminal, often numerous, sometimes crowded and obscuring the thallus,
1-3 (4) cm in diameter, thalline exciple rugose, strongly maculate, disc widely perforate,
spores 13-18 x 8-10 //m. Conidia filiform, 10-16 (20) //m long.
TLC: lecanoric acid, atranorin.

PARMEL1A (AMPHIGYMNIA) IN EAST AFRICA 169

Parmelia andina differs from P. hololoba mainly in the absence of cilia. When P. hololoba
extends inland and occurs in the same habitats as P. andina, its cilia sometimes become
sparse, short, and stunted, so that the two species are almost indistinguishable.

Parmelia andina is common and widespread in dry, well lit sites, such as savannas, open
hillsides, and artificial habitats. It has been collected between 900 and 2400 m altitude in all
four countries of our studies. Outside our area it is known from numerous localities in
continental Africa as well as from Madagascar, India, Thailand, Tahiti, and South America
(Hale, 1965).

Selected East African records

Ethiopia. See Winnem (1975). Kenya. Eastern Province, Isiolo District, 8 km S of Isiolo, 4K 28/101;
Machakos District, Kilima Kiu, 5K3/2; Embu District, Embu, Izaak Walton Inn, K 53/109; Meru
District, vicinity of Chogoria, 3 K 9/1 04; Marsabit District, c. 1 km N of the mountain Ajaumarka,
Moberg 3864 (UPS). Rift Valley Province, Elgeyo Marakwet District, 8 km NE of Kapcherop,
2K 1 5/1 14; Nakuru District, E of road by Lake Elmenteita, 2 K 20/101 ; Samburu District, WSW slope
of Warges, 4K3/106. Western Province, Kakamega District, Kakamega Forest by River Ikuyawa,
4K 10/105. Nyanza Province, Kisii District, 1 km SE of Kisii, 4K 12/114. Central Province, Nyeri
District, Mt Kenya Safari Club, 4K 19/112. Tanzania. Northern Province, Arusha District, Arusha
National Park, Kusare Forest, T 2/120. Eastern Province, Morogoro District, above Morningside in
the Uluguru Mts, 6 May 1978, Dahl(O). Southern Highlands Province, Iringa District, Ruaha National
Park, summit of Magangwe Hill. Bjornstad 1780-a (O). Southern Province, Songea District, Gumbiro,
Nordal INB 849 (O). Uganda. W. Mengo District, Kyadondo County, Makerere University Campus,
U l/7b, Manum 1/1968 (O); Busiro County, Entebbe Botanical Garden, U 9/56b. Toro District,
Burahya County, Fort Portal, Dale 144 (BM type of P. dalei Dodge); Busongora County, 10 km NW
of Kilembe, 2U 12/22. Karamoja District, Matheniko County, south side of Mt Moroto, 2U 35/4. West
Nile District, Mt Otse, 1962, Thomas (BM type of P. thomasii Dodge).

5. Parmelia aprica Krog & Swinscow sp. nov. Fig. 4

Thallus corticola, coriaceus, adnatus, flavus ad flavo-viridis. Lobi O'5-l cm lati, eciliati, marginibus

Fig. 4 Parmelia aprica Krog & Swinscow, holotype (O). Rule = 1 cm.

170 H. KROG & T. D. V. SWINSCOW

integris vel leniter crenatis, superne emaculati, rugescentes, rimosi, Soralia marginalia, ad lobos
peripherales linearia, ad lobos ascendentes laterales subcapitata, aliquando ad laminam extensa.
Apothecia et pycnidia ignota. Acidum protocetraricum et acidum usnicum continens.

Thallus corticolous, coriaceous, adnate, bright yellow to yellow-green. Lobes O'5-l cm
broad, eciliate, with entire or weakly crenate margins. Upper side emaculate, becoming
rugose, irregularly cracked and sometimes lobulate towards the centre. Medulla white.
Underside black in the centre, with a broad, brown, shiny marginal zone, rhizines
sparsely developed. Soralia marginal, linear on peripheral lobes and subcapitate on
ascending lateral lobes, sometimes spreading diffusely on to the lamina. Soredia granular,
pale green to yellowish. Apothecia and pycnidia unknown.

TLC: protocetraric acid, traces of fatty acids, usnic acid.

Type: Kenya. Eastern Province, Machakos District, lava flow 5 km NW of Kibwezi,
235'S, 3751'E, 1000m altitude, on shrubs, January 1972, coll. H. Krog & T. D. V.
Swinscowno. K 20/103 (O holotype; BM, UPS isotypes).

Parmelia aprica differs from both P. rava and P. dilatata in its bright yellow colour, yellow-
green soredia, and the absence of atranorin in the cortex; it differs from P. dilatata also in the
absence of echinocarpic acid and associated substances.

Parmelia vivida, newly described in this work, appears to be the corresponding primary
species. It has spores 20-22 x 8-10 jam and sublageniform conidia 6-7 //m long.

Parmelia aprica is known only from the type locality, where it grew on shrubs in a rather
dry site exposed to high light intensity. Further collection numbers are 2K 22/105 and
3K23/1,129.

6. Parmelia araucariarum Zahlbr.

Denkschr. Akad. Wiss., Wien, math.-nat. Kl. 83 : 179 (1909). Parmotrema araucariarum (Zahlbr.)
Hale, Phytologia 28 : 334 (1974). Type: Brasilia. Prov. Sao Paulo. Prope S. Amaro in circuitu urbis
S. Paulo, 800 m, May 1901, leg. V. Schiffner (W holotype, O isotype). [TLC: pigment, fatty acids
in the reddenda complex, atranorin.]

Thallus corticolous, membranaceous, loosely attached, green-grey. Lobes O'5-l - 5 cm broad,
ciliate, cilia 1-2 (3) mm long. Upper side emaculate, more or less irregularly cra