Haematomma and Ophioparma: Two Superficially Similar Genera of Lichenized Fungi

Abstract

The genus Haematomma Massal. (type species: H. ochroleucum (Necker) Laundon) includes the pantropical and subtropical H. puniceum group, but differs in a range of characters including ascus structure, paraphyses, and chemistry from Ophioparma Norman (type species: Ophioparma ventosa (L.) Norman). The differences are such that a new family Ophioparmaceae Rogers & Hafellner is described with Ophioparma as its type genus, and the circumscription of the family Haematommaceae Hafellner is emended to take these changes into account. The combination Rosellinula kalbii (Hafellner) comb. nov. is proposed.

Full text

Lichenologist 20(2): 167-174 (1988)
HAEMATOMMA
AND OPHIOPARMA:
TWO SUPERFICIALLY SIMILAR GENERA
OF
LICHENIZED FUNGI
R.
W.
ROGERS*
and J.
HAFELLNERJ
Abstract: The genus
Haematomma
Massal. (type species:
H.
ochroleucum
(Necker)
Laundon) includes the pantropical and subtropical H.
pumceum
group, but differs in
a range
of
characters including ascus structure, paraphyses,
and
chemistry from
Ophioparma
Norman (type species: Ophioparma ventosa
(L.)
Norman).
The
differences
are
such that
a new
family Ophioparmaceae Rogers
&
Hafellner
is
described with
Ophioparma
as its type genus, and the circumscription
of
the family
Haematommaceae Hafellner
is
emended
to
take these changes into account.
The
combination
Rosellinula kalbii
(Hafellner) comb. nov. is proposed.
Introduction
The generic name
Haematomma
Massal. was proposed originally to house two
species,
Haematomma vulgare
Massal. (i.e.
H.
ochroleucum
(Necker) Laundon)
and H.
ventosum
(L.) Massal. (Massalongo 1852). Since that time, many more
species have been included
in
Haematomma
without
any
detailed consider-
ation
of the
circumscription
of the
genus.
The
great diversity
of
species
in
Haematomma
was
appreciated
by
Culberson (1964)
who
recognized four
sections in the genus.
It
has
become clear that
Haematomma
as
currently applied
is a
collective
genus consisting
of
species with some morphological similarities which ought
to be
segregated
(Rogers &
Bartlett
1986).
When discussing
this
problem Rogers
&
Bartlett (1986) erroneously associated
the
type species
of
Haematomma,
H.
vulgare
Massal,
with the H.
ventosum
group.
On this
basis,
after considering the
characterization
of
Haematomma
and the
family Haematommaceae
in
Hafellner (1984), they suggested that
the
subtropical species
of
Haematomma
(the H.
puniceum
group) should
be
transferred
out of
the genus
Haematomma
and the family Haematommaceae, perhaps into the family Scoliciosporaceae.
To clarify the situation
we
have examined material from three species groups
within the
genus,
H.
ochroleucum
(Necker) Laundon representing the holarctic
ochroleucum
group,
H.
puniceum representing
the
widespread subtropical
puniceum group,
and H.
ventosum representing
the
arctic-boreal ventosum
group.
From these observations (Table
1,
Figs
1-3), it
became apparent that
the
puniceum
group was indeed closely related to the
ochroleucum
group, and could
be retained in the genus
Haematomma.
The
ventosum
group, however, was very
different,
and
must
be
segregated into
its own
genus,
for
which
the
name
Ophioparma
Norman is available. Because of its distinctive ascus structure, the
•Botany Department, University of Queensland, St Lucia, Queensland
4067,
Australia.
JInstitut
fur
Botanik, Karl-Franzens Universitat, Holteigasse
6,
Graz
A-8010,
Austria.
0024-2829/88/020167 +
08
$03.00/0
© 1988
British Lichen Society

TABLE
1.
Comparison
of
the
H. ochroleucum £ro«p, H. puniceum
group and
Ophioparma
Attribute
Ecology
Distribution
Substrate preference
Reproduction
Sexual
Vegetative diaspores
Excipulum type
Proper exciple
Thalline exciple
Hymenial characters
Asci
Paraphyses
Spores
K reaction of epihymenium
Red pigment
Hypothecium
Pycnidia
Ostiole of pycnidium
Conidiophores
Chemistry
Atranorin
Thamnolic acid
Lichenicolous fungi
H.
ochroleucum
group
Holarctic
Most species corticolous, a few
saxicolous
Apothecia rarely developed
Soredia
Thin
Thick and permanent or aspicilioid
Lecanora-type, usually poorly
developed
Thin, richly branched and anasto-
mosing, not thickened at the tip
Often helically twisted
K + magenta in hymenial gel
Acetone insoluble
Hyaline
Without green pigment
Vobis type
V
Present
Absent
Rosellinula
kalbii*
H.
puniceum
group
Tropical-subtropical
Most species corticolous, a few
saxicolous
Apothecia commonly developed
Soredia
Thin
Thick and permanent or aspicilioid
Lecanora-type, well developed
Thin, richly branched and anasto-
mosing, not thickened at the tip
Often helically twisted
K + magenta in hymenial gel,
or K + purple-grey crystals
rapidly fading in solution
Acetone insoluble
Hyaline
Without green pigment
Vobis type
V
Present
Absent
Rosellinula
kalbii*
Ophioparma
Arctic-boreal
Saxicolous, mostly on siliceous rocks
Apothecia commonly developed
Absent
Thick
Thick or absent
Ophioparma-type
Thick, with few branches and anasto-
moses, thickened at the tip
Sometimes twisted
K + blue crystals fading to magenta
and dispersing in solution
Acetone soluble
Pigmented
With green pigment
Vobis type VI
Absent
Present or absent
Muellerella
pygmaea
•Rosellinula kalbii (Hafellner) Hafellner
&
Rogers comb. nov. Basionym:
Roselliniella kalbii
Hafellner,
Herzogia
7:
152
(1985).

1988Haematomma
&
Ophioparma—Rogers
&
Hafellner
169
FIG.
1.
Haematomma puniceum (Rogers
884,
BRIU).
A,
young ascus
and
paraphyses;
B,
mature
ascospore;
C,
conidiophores
and
conidia
[see
also
Fig.
3E].
Amyloid parts
of
the
ascus stippled.
Scale
=
10 \xm.
genus Ophioparma
is
best placed
in a new
family Ophioparmaceae Rogers
&
Hafellner. Consequent upon these decisions, emended descriptions
of the
genera Haematomma
and
Ophioparma
are
necessary.
Emended descriptions
Haematommaceae Hafellner
Beih.
Nova Hedwigia 79: 281 (1984).

170THE LICHENOLOGIST
Vol.
20
FIG.
2.
Gphioparma ventosa (Santesson
31089, BRIU). A, young ascus; B, paraphyses; C, mature
ascospore; D, conidiophores and conidia. Amyloid parts of the ascus stippled. Scale=
10
um.
Lichenized ascomycetes containing a coccoid green algal symbiont. Thallus
crustose. Apothecia sessile to immersed, proper margin thin, thalline margin
usually well developed, disc pigmented red. Asci of the
Lecanora-type
with an
1+ tholus, distinct ocular chamber and axial mass, encased in 1+ blue gel;
eight-spored. Paraphyses richly branched and anastomosing. Ascospores
hyaline, transversely septate, without a halo.

1988Haematomma & Ophioparma—Rogers & Hafellner 171
FIG.
3. Haematomma ochroleuceum. A, young ascus B, paraphyses; C, mature ascospore
(Magnusson, Lich. Set.
Scand.
no. 74, GZU). D, H. ochroleucum var. porphyrium, conidia
(Buschardt et al. s.n., GZU). E, H. puniceum conidia (Kalb, Lich. Neotrop. no. 17, GZU); see also
Fig. 1C. Scale =10(im.
Haematomma Massal.
Ric. Lich. Crost.: 32 (1852)
Type species: Haematomma vulgare Massal. (lectotype, selected by Fink 1910:
186).
[
= Haematomma
coccineum
(Dickson) Korber, Haematomma ochroleucum
(Necker) Laundon.]
Haematomma sect. Haematomma
sensu
Culb., Bryologist 66:226 (1964).

172 THELICHENOLOGIST Vol.20
(Figs
1
&
3).
Lichenized ascomycetes. Thallus crustose, corticate, with or without soredia.
Apothecia with an open disc, sessile or adnate to immersed in the thallus; the
disc red, round or somewhat irregular; proper exciple thin, thalline margin well
developed and coloured like the thallus in species with sessile apothecia, less
developed to absent in species with adnate to immersed apothecia; hymenial
layers with a red acetone insoluble pigment mostly concentrated above the
ascus tips; hypothecium hyaline. Paraphyses less than 2 um thick, richly
branched and anastomosing, not thickened at the apices. Asci of the
Lecanora
type,
tholus I + blue with an ocular chamber and faintly I + blue axial mass.
Ascospores
hyaline, transversely 3—25-septate, thin-walled usually helically
coiled in the
ascus.
Pycnidia immersed with ostiole region red (K + magenta) or
hyaline, never with a green pigment; conidiophores type V (Vobis 1980).
Ophioparmaceae
R.
W. Rogers
&
Hafellner fam.
nov.
Ascomycetes lichenisati algas coccales continentes. Thallus crustaceus. Apothecia sessilia; discus
sanguineus
vel
castaneus; margine proprio crassa; margine thallino presentia
vel
absentia. Asci
tholis amyloideis, sine
'
masse axiale'
et '
chambre oculaire', gelatina amyloidea circumdati,
octospori. Paraphyses crassae, parce ramosae
et
anastomosantes. Ascosporae hyalinae, aciculares,
transversaliter septatae, non halonatae.
Typus familiae:
Ophioparma
Norman.
Lichenized ascomycetes with coccoid green algal symbiont.
Thallus
crustose.
Apothecia dark blood red to chestnut, sessile, proper margin thick, thalline
margin present or absent* Ascus with an I + blue tholus, without an ocular
chamber, axial plug or ring structure, encased in 1+ blue gel, eight-spored.
Paraphyses thick, little branched, with thickened apices.
Ascospores
hyaline,
broadly acicular with transverse septa, helically contorted in the ascus, without
a halo.
Ophioparma Norman
NyttMag.
Naturv.
7:230
(1853);
typespecies-.Ophioparma
ventosa
(L.)Norman (lectotypeselected
by Hafellner
1984:
283).
Lepadolemma
Trevisan,
Riv.
period
Lav. I. R. Acad.
Padua
1: 267
(1853); type species:
Lepadolemma ventosum
(L.) Trevisan (lectotype selected by Hafellner 1984).
Haematomma
sect.
Ventosa
Culb,
Bryologist
66:229
(1964);
type
species:
Haematomma ventosum
(L.) Massal. (holotype).
(Fig. 2)
Lichenized ascomycetes. Thallus crustose, corticate esorediate. Apothecia
sessile, disc round to irregular, blood red to chestnut brown, proper margin
thick, concolorous with the disc, thalloid margin present or absent, con-
colorous with the thallus; hymenium pigmented orange to umber throughout;
hypothecium hyaline in the upper parts but pigmented rose to buff in the lower
parts.
Paraphyses
1-5-2-5
urn thick, rarely branched and anastomosing, some-
what thickened at the tip. Asci with an 1+ blue tholus, but without an ocular
chamber or axial mass. Ascospores hyaline, thin-walled, transversely 3-7-
septate. Pycnidia immersed; ostiole region pigmented dark green (pigment
K-,HNO3+ purple-red); conidiophores type VI (Vobis 1980).

1988 Haematomma &
Ophioparma—Rogers
& Hafellner 173
Additional European taxa requiring inclusion are:
Ophioparma lapponica (Rasanen) Hafellner
& R. W.
Rogers comb,
nov.
Haematomma lapponicum Rasanen,
Ann. Acad.
Sci.fenn.,
A,
34(4):
67
(1931).
Ophioparma ventosa var. cuprigena (Poelt) Hafellner
&
R. W. Rogers
comb.
nov.
Haematomma ventosum var. euprigenum Poelt, Verh. zool.-bot.
Ges.
Wien 95:
112
(1955).
Discussion
It is apparent from Table 1 that Haematomma as treated here (including the
H.
ochroleucum
group and the H. puniceum group) is an homogeneous genus,
although the two groups differ somewhat in distribution pattern and in the
degree to which reproductive effort is concentrated in vegetative propagules.
The H.
puniceum
group is found in both Hemispheres, mostly in the tropics and
subtropics, but is known (e.g. H.
infuscum,
H.
sorediatum)
from cool temperate
areas in both Australia and New Zealand (Rogers
&
Bartlett 1986, Rogers 1982,
1985),
and from alpine areas of New Zealand {H. alpinum, Rogers & Bartlett
1986).
The H. ochroleucum group is found in cool, coastal holarctic
regions, and can be considered ecologically to be a northward extension of the
H. puniceum group, occupying essentially similar habitats. The variation in
conidial morphology in material determined as H. puniceum (Figs lc & 3e)
is puzzling: it may be that these are actually different species, or that
Haematomma
has dimorphic conidia in at least some species.
Some species of the H.
ochroleucum
group have extensive development of
soredia, the whole thallus effectively dissolving into soredia in H.
ochroleucum.
Apothecia are normally rare in this group. In the H.
puniceum
group soredia are
rare,
occurring in small punctiform soralia on a few species; apothecia are
common in all species.
In contrast to Haematomma, Ophioparma is restricted to the Northern
Hemisphere where it is found in arctic-boreal regions and, whereas most
Haematomma species occur only on bark, all species of Ophioparma are
restricted to rocks. In Ophioparma there are no vegetative diaspores, and
apothecia are common.
All species of
Haematomma
examined have a thin proper exciple and either
have a well-developed thalline exciple or are aspicilioid. This contrasts with
Ophioparma,
in which a thick proper exciple is always present, and in which a
thalline exciple may or may not be present.
Hymenial characters are critical for the demonstration of differences
between the genera. Haematomma has a Lecanora-type ascus, although in
the H.
ochroleucum
group the apical apparatus of the ascus may be poorly
developed. Such poor development is not uncommon in the asci of species
which rely primarily on vegetative diaspores for their propagation (e.g.
Hypocenomyce). The ascus of Ophioparma is quite different, having well-
developed amyloid structures in
a
continuous dome over the apical region (Fig.

174 THELICHENOLOGIST Vol.20
2),
with no ocular chamber or axial mass visible under
a
light
microscope.
This
is a distinct and different ascus type, here named the
Ophioparma
type.
When comparisons are made between these two genera it is apparent that
they have very little in common besides a more or less red apothecial disc (but
with different pigments) and acicular, transversely septate spores. It is likely
that the brown-fruited European species until now referred to
Haematomma
ought to be placed in the genus
Loxospora
Massal., and that the east Asian
species with brown fruits ought also to be transferred to another genus.
Specimens
examined:
Haematomma ochroleucum (Necker) Laundon:
Sweden:
Bohuslan,
par.
Stala, Varekil, on perpendicular or slightly overhanging rocks, 17 July 1928, A. H.
Magnusson
[Lich. Sel. Scand. no. 74, as H.
coccineum
var.
ochroleucum],
(GZU). H. ochroleucum var.
porphyrium (Persoon) Laundon: Norway: Hordaland, Insel Stord, Sandvikag, Gem. Fitjar,
Waldund Felshange aus teilweisemetamorphen Gestein,
10
September
1976,
A.
Buschardt,
P.
M.
Jorgensen
& J. Poelt,
(GZU).—H.
puniceum (Ach.) Massal.: Australia: Queensland, Keppel
Sands on bark of
Rhizophora,
10
July
1975,
R. W.
Rogers 884
(BRIU);
Brazil: Sao Paulo, Serra da
Mantiqueira, bei der Stadt Campos do Jordao, etwa
150
km NE von Sao Paulo,
1700
m, am Rand
eines feuchten Cerrado, 25 May 1978, K. Kalb & G. Plobst [Kalb, Lich.
Neotropici
no. 17]
(GZU).—Ophioparma ventosa (L.) Norman: Norway: Harjedalen, Tannas parish, between
Giertebaune and Rossvalen,
(c.
6
km
W
of Mt Stora Mittaklappen)
c.
980
m, on rocks on an alpine
heath,
1
August
1984,
R.
Santesson 31089
(BRIU).
Professor
R.
Santesson kindly provided the material of
Ophioparma ventosa
used in this study. The
differences in ascus structure between
Ophioparma
and
Haematomma
first
became apparent when
R. Rogers
was a
guest
on
a Deutscher Akademischer Austauschdienst fellowship
in
Professor Dr A.
Henssen's Laboratory in Marburg. J. H.'s contribution to this work was aided by a University of
Queensland travelling fellowship.
REFERENCES
Culberson,
W.
L. (1964) [' 1963
'] A
summary of the lichen genus
Haematomma
in North America.
Bryologist
66:224-236.
Fink, B. (1910) The Lichens of Minnesota.
Contributions
from the United States National
Herbarium
14(1):
1-269.
Hafellner, J. (1984) Studien in Richtung einer naturlicheren Gliederung der Sammelfamilien
Lecanoraceae und Lecideaceae.
Beiheft
zur Nova
Hedwigia
79:241-372.
Massalongo,
A.
B. (1852)
Ricerche sull'Autonomia dei Licheni
Crostosi.
Verona: Friziero.
Rogers,
R.
W.
(1982) The corticolous
species
of
Haematomma in
Australia.
Bryologist
14:115-129.
Rogers, R. W. (1985) Additional notes on
Haematomma
in Australia.
Lichenologist
17:
307-309.
Rogers,
R.
W.
&
Bartlett,
J.
K.
(1986)
The lichen
genus Haematomma
in New
Zealand.
Lichenologist
18:247-255.
Vobis, G. (1980) Bau und Entwicklung der Flechten-Pycnidien und ihrer Conidien.
Bibliotheca
Lichenologica
14:1—141.
Accepted
for
publication
25
September
1987