The CONTRIBUTIONS TOWARDS A NEW SYSTEMATICS OF THE LICHEN FAMILY
THELOTREMATACEAE is a three-part monograph based on the Ph.D. thesis
of Andreas Frisch. The main part of the book deals with THE LICHEN
FAMILY THELOTREMATACEAE IN AFRICA (370 pages), authored by Frisch,
completed by A MONOGRAPH OF THELOTREMATACEAE WITH A COMPLEX STRUCTURE
OF THE COLUMELLA (with Kalb; 146 pages), a MOLECULAR PHYLOGENY OF THE
THELOTREMATACEAE (with Kalb and Grube; 23 pages), and a taxonomic
index. This monumental work is another milestone, which significantly
improves our knowledge of tropical crustose lichens. The numerous
photographic plates and line drawings are of excellent quality. In the
main part, African Thelotremataceae are critically revised, completed
by genera with complex columella treated in the second part. The total
number of accepted species for Africa is 94, including 17 new to
science and 47 new records for the continent, while 35 names are
synonymized. This is an important achievement and an invaluable
contribution to the African lichen biota, especially considering that
most of the relevant collections were made by the author himself. Many
types were studied and new characters explored, closing substantial
gaps in our knowledge of the Thelotremataceae at the world level.
Besides the taxonomic treatment, the profound generic revision makes
this work outstanding. Sixteen genera are accepted, compared to the
five ascospore genera distinguished by Müller Argoviensis in 1887 and
the three excipular genera from Hale’s revision in 1980. Besides
Chroodiscus, Leptotrema, Myriotrema, Ocellularia, and Thelotrema, the
genera treated in this monograph include three reinstated names
(Chapsa, Leucodecton, Stegobolus), three recently established taxa
(Ampliotrema, Reimnitzia, Topeliopsis), and five new genera introduced
by the author: Acanthotrema, Fibrillithecis, Gyrotrema, Melanotrema,
and Redingeria. This work is thus a very important step towards a
natural generic delimitation in the family and confirms other recent
studies that ascoma morphology, wall structures, and ascospore types
(but not necessarily septation and color) are important characters to
distinguish genera. In some instances it is even shown that species
with pale and others with carbonized excipulum, previously separated
in Myriotrema and Ocellularia, are closely related. The separation of
four genera with chroodiscoid apothecia, traditionally included in
Thelotrema s.lat., comes as a surprise, but is backed by morphological
and molecular data. Chroodiscus is now restricted to foliicolous taxa,
while the non-foliicolous species are assigned to Chapsa. Reimnitzia
is certainly an oddball deserving its own genus, but the new
Acanthotrema for the single species A. brasilianum needs further
study. The diagnostic character are the spiny paraphyses and
periphysoids, but the author presumes that certain species with smooth
paraphyses might also belong in Acanthotrema. In the phylogenetic
analysis, the genus falls outside Chapsa and Chroodiscus, but its
position is unresolved and hypothesis testing to reject a monophyletic
Chapsa that includes Acanthotrema would have been appropriate.
In the second part, the authors show that taxa with complex columella
structures evolved independently and make sense of this by
establishing three new genera, Gyrotrema, Melanotrema, and Redingeria,
and reinstating the old name Stegobolus, while retaining a couple of
species in Myriotrema and Ocellularia. Except for Redingeria, for
which no sequences were available, and the apparently polyphyletic
Stegobolus, the concept is confirmed by molecular data. However, one
might have problems to distinguish Redingeria and Stegobolus. In the
discussion of Redingeria, it is stated that Stegobolus is "... well
distinguished ... in morphology and might be only distantly related."
(p. 406), but the picture of Redingeria glyphica (Pl. 20) closely
resembles Stegobolus species (e.g., Pl. 21, 23). According to the key,
the diagnostic differences are the Redingeria-type ascospores and the
inclusion of ascospore remnants in the excipulum, but it takes a while
to find out what exactly Redingeria-type ascospores are. The
difference in ornamentation, depicted in Fig. 121 (Redingeria) versus
129b (Stegobolus), does not appear significant, especially as the
ascospore in Fig. 121 is fully turgescent, while the one in Fig. 129b
appears collapsed. The structure of the ornamentation itself is
similar and could even be interpreted as a synapomorphy. As for the
inclusion of old ascospores in the excipulum and columella, the
authors themselves state that this character is "... of limited
taxonomic relevance." (p. 406). The remaining differences are the
thick-walled ascospores (especially when young) with irregular lumina
in Redingeria, but it remains to be tested whether these hold as
diagnostic characters at genus level.
One also wonders about the separation of Melanotrema versus the
Ocellularia auberianoides group, including the common neotropical O.
bahiana (formerly in Myriotrema). In the key (p. 378), the main
difference is the carbonized versus pale to brown excipulum, but
species with different excipular pigmentation are accepted in
Ocellularia. The Ocellularia auberianoides group is said to have an
excipulum with "... clearly radiating hyphae ..."; the corresponding
character is not mentioned for Melanotrema, but in the discussion the
latter genus is said to have an excipulum with "... clearly radiating
hyphae ..." (p. 383). The phenocortex of Melanotrema is said to be
"... loosely organised or cartilaginous ...", while the
corresponding alternative for the Ocellularia auberianoides group is
not given. In the descriptions, the phenocortex is given as formed
from irregular hyphae for both groups. At the species level, some
conclusions are difficult to follow. The African material of
Acanthotrema brasilianum differs from neotropical populations by
submuriform versus 3-septate ascospores and might warrant recognition
at the species level. Specimens from Africa identified as Chapsa
alborosella differ from neotropical material by the presence of
stictic acid, but it is argued that the TLC results were contaminated
by dead thalli of Graphis growing under the Chapsa specimens. It is
highly unlikely that the two widely separated African collections from
Tanzania and Réunion grew on dead thalli of Graphis species that both
produced stictic acid, a substance which is comparatively rarer in
Graphis than in Chapsa. Thelotrema platycarpoides is listed as synonym
of Chapsa platycarpa and said to represent a juvenile form of the
latter because of the small apothecia. However, the ascospores of the
latter are mostly 5-septate, while typical C. platycarpa has 3-septate
ascospores, which contradicts the hypothesis of the former being a
juvenile form of the latter.
The genus keys work relatively well, but a good knowledge of the
morphological concepts outlined throughout the book are required to
use the keys, since terms such as Ocellularia-type excipulum or
Redingeria-type ascospores are frequently used. A quick reference as
to where these terms are defined, or perhaps a short, illustrated
glossary, would have been helpful. The lengthy couples in the genus
keys make it sometimes difficult to filter the characters that
separate a genus from another; it might have been better to exclude
most of the non-differential information. The species keys work well,
although in some instances (e.g., Redingeria), they strongly rely on
chemistry, and one is required to do TLC before even starting to work
with the key, especially in cases where spot tests do not work.
The third part of the work is a first attempt to test the generic
rearrangement with DNA data. The number of newly produced sequences is
impressive and many of the genera are supported, such as Chapsa,
Fibrillithecis, Leucodecton, and Melanotrema, while a few need further
study. The inclusion of the molecular phylogeny in this work supports
the systematic changes, but as a stand-alone paper, one would have
wished for more rigorous analyses of the data, including maximum
parsimony and alternative hypothesis testing. The placement of the
figures as appendix is odd, and it also takes some time to realize
that the four trees do not represent different analyses, but map
different taxonomic groups and characters on the same tree. Otherwise,
this analysis is an excellent base for further studies and identifies
the problematic genera and species groups.
The taxonomic index at the end of the book is a bit difficult to use
since the epithets are listed under the genera in which they were
described. In order to find an epithet, one has to know the original
genus or scroll through the entire index. Alphabetical arrangement of
epithets, with corresponding genera in brackets, would have been a
more practical solution. A few other inconsistencies might also be
mentioned. The concept of "phenocortex" does not conform to its
original definition, according to which a phenocortex is a cortex-like
structure formed by dead algal cells, hyphal fragments, and other
materials. From the descriptions and illustrations (p. 37ff), it is
obvious that Thelotremataceae largely feature a true cortex formed 552
by fungal hyphae (e.g., Pl. 1d). Naming the perfectly conical
columella shape in Fig. 5c "truncated conical", while the shape that
resembles a skittle (Fig. 5a) is termed "conical", will cause
confusion. It probably stems from the notion that in German, skittles
are named "Kegel" ("cones"), even if those "Kegel" are not
conical in the technical sense. The species descriptions are reversed,
mentioning ascospore characters before asci and paraphyses, which
makes them more difficult to read. In some places, data on
distributions are inconsistent. Chapsa indica is given as ´Africa and
tropical Asia¡, while the pantropical Thelotrema pycnophragmium is
listed as a synonym. On page 328, is is stated that Thelotrema
glaucopallens "... might be rare or even absent in some areas for no
obvious reasons ..." (citing Hale’s Sri Lanka paper from 1981), but it
is difficult to understand how the absence of that species in East
Africa "... can be explained by this ...". A few Latin epithets are
incorrect. Chapsa cinchonara should read cinchonarum, as it represents
the genitive plural of the tree genus Cinchona. Ocellularia polydisca
should read polydiscus, the epithet being a noun. In the citations of
Leucodecton subcompunctum (p. 163) and Reimnitzia santensis (p. 272),
the term "lectoparatype" is used, suggested some time ago to denote
those syntypes or paratypes remaining after lectotypification.
However, it was argued (Vorster, Taxon 35: 316-317, 1986) that such a
designation has no meaning, since syntypes and paratypes are not types
and, with the selection of a lectotype, cease to have value for
typification, except for the original protologue in which they
continue to be syntypes or paratypes. Thus, there is no provision in
the Code for the term "lectoparatype" and such material cannot be
cited as type. The (≡) sign is applied against current usage to
denote heterotypic taxonomic synonyms, for which the (=) sign should
have been used instead.
Overall, the CONTRIBUTIONS TOWARDS A NEW SYSTEMATICS OF THE LICHEN
FAMILY THELOTREMATACEAE is an important work, indispensable for
colleagues and students dealing with tropical lichens. It represents
an outstanding treatment of African Thelo-tremataceae and an important
step towards a generic rearrangement of the family, including a wealth
of data and new ideas which will stimulate many further studies in
this diverse group of lichens. The book invites for comparison with
Staiger’s DIE FLECHTENFAMILIE GRAPHIDACEAE, and while both have a
similar underlying philosophy, the CONTRIBUTIONS TOWARDS A NEW
SYSTEMATICS OF THE LICHEN FAMILY THELOTREMATACEAE has the advantage of
being entirely in English and thus accessible for a wide audience. The
taxonomic treatment, even if geographically restricted to Africa or
taxonomically to the species with complex columella, is very useful
for identifying species at the world level, especially as many
non-African taxa are discussed in the first part. Students will be
happy to see that, while Staiger’s Graphidaceae from 2002 is priced at
128,- (approx. $ 165,-), the 88,- ((approx. $ 113,-) for the present
book are much more reasonable, especially considering that it has 40
pages more and twice as many photographic plates, all integrated in
the text.
ROBERT LÜCKING, Chicago
Nova Hedwigia 84 3-4, p. 549-552