Ocean view: A first assessment of the littoral, crustose lichen biota of south Brazil

Abstract

The crustose lichen biota of coastal rocks in South Brazil was investigated. Several distinct lichen zones were found: a littoral black zone, a supralittoral yellow zone and a grey zone with species restricted to either exposed granite, dry overhangs, damp overhangs or places subjected to run-off. Dendrographa austrosorediata is newly described, with a crustose, flat to partly curling up or blister-like thallus which is much dissected, whitish grey, 0·1–0·2 mm thick; surface minutely densely cracked, rimose, with cracks intersecting at each c. 0·1 mm; soredia whitish to bluish grey but asymmetrically blackened in the direction facing the light, originating on the thallus surface, in initially discrete convex soralia. The phylogenetic position of this new species was traced by molecular methods. Stigmidium marinum , generally regarded as a lichenicolous fungus, was found as a free-living lichen, thousands of kilometres away from the nearest known occurrence of any purported host.

Full text

The Lichenologist 49(6): 597–605 (2017) © British Lichen Society, 2017
doi:10.1017/S0024282917000512
Ocean view: a first assessment of the littoral, crustose lichen biota
of south Brazil
André APTROOT, Emerson Luiz GUMBOSKI and
Marcela Eugenia da Silva CÁCERES
Abstract: The crustose lichen biota of coastal rocks in South Brazil was investigated. Several distinct
lichen zones were found: a littoral black zone, a supralittoral yellow zone and a grey zone with species
restricted to either exposed granite, dry overhangs, damp overhangs or places subjected to run-off.
Dendrographa austrosorediata is newly described, with a crustose, flat to partly curling up or blister-like
thallus which is much dissected, whitish grey, 0·1–0·2 mm thick; surface minutely densely cracked,
rimose, with cracks intersecting at each c. 0·1 mm; soredia whitish to bluish grey but asymmetrically
blackened in the direction facing the light, originating on the thallus surface, in initially discrete convex
soralia. The phylogenetic position of this new species was traced by molecular methods. Stigmidium
marinum, generally regarded as a lichenicolous fungus, was found as a free-living lichen, thousands of
kilometres away from the nearest known occurrence of any purported host.
Key words: Dendrographa, maritime, saxicolous, Stigmidium
Accepted for publication 9May 2017
Introduction
The lichen biota of Brazil is still incompletely
known. However, with over 4750 species
recorded to date, it is already a country con-
taining a relatively high number of lichen
species. As every year around 200 species are
added to this list, about half of which are
newly described, it will soon become the
country with the highest number of lichen
species. Yet there are still whole states and
habitats that have not been sampled at all.
The coast of Brazil is mostly known for its
wide and long sandy beaches, only inter-
rupted by river estuaries with mangrove
forest. In the southern states of Paraná, Santa
Catarina and Rio Grande do Sul, these
beaches are interspersed with coastal granite
rocks that often rise as steep cliffs directly
from the sea. The targeted study of the lichen
mycobiota on these coastal cliffs began in
1890 (Vainio 1890), to be taken up again in
1984 with the report of some macrolichens
(Osorio & Fleig 1984a,b), and then subse-
quently discontinued for many years but
taken up again recently, mainly with the
study of Parmeliaceae and Cladoniaceae
(Benatti et al. 2008; Benatti & Marcelli
2009a,b; Gumboski & Eliasaro 2011, 2012a,b;
Gerlach & Eliasaro 2012, 2014a,b).
All over the Northern Hemisphere, as well
as in Australia and New Zealand, specialized
lichen biota of coastal rocks have been repor-
ted, consisting partly of littoral species that are
restricted to those coastal rocks (e.g. Arup
1995; Gilbert 2000). Usually, different zones
with characteristic lichens can be recognized: a
littoral black zone, a supralittoral yellow zone
and a grey zone, the latter with some micro-
habitats such as dry and wet overhangs,
exposed rocks and places with run-off. Species
from the lower zones and the dry overhangs
are generally restricted to maritime habitats.
The lichen biota of coastal rocks have rarely
been reported from tropical regions.
A. Aptroot: ABL Herbarium, G.v.d.Veenstraat 107,
NL-3762 XK Soest, The Netherlands. Email: andreapt
root@gmail.com
E. L. Gumboski: Departamento de Ciências Biológicas,
Universidade da Região de Joinville, CEP: 89219-710,
Joinville, Santa Catarina, Brazil.
M. E. S. Cáceres: Departamento de Biociências,
Universidade Federal de Sergipe, CEP: 49500-000,
Itabaiana, Sergipe, Brazil.
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In 2015, the second author organized
an excursion of the Reunião Brasileira de
Estudos Liquenológicos in Santa Catarina
which allowed the authors to study the
lichens on coastal cliffs in the area.
Material and Methods
Identification and descriptive work was carried out in
Soest using an Olympus SZX7 stereomicroscope and an
Olympus BX50 compound microscope with interference
contrast, connected to a Nikon Coolpix digital camera.
Sections were mounted in tap water, in which all mea-
surements were also taken. The specimens from this study
are preserved in JOI and ABL. The chemistry of selected
specimens was investigated by thin-layer chromatography
(TLC) using solvent A (Orange et al. 2001).
DNA extraction, amplification and sequencing were
carried out in Santander (ALVALAB): total DNA was
extracted from dry specimens using a modified protocol
based on Murray & Thompson (1980). A portion of each
sample was blended with the aid of a micropestle in
600 µl CTAB buffer (CTAB 2%, NaCl 1·4 M, EDTA
(pH 8·0) 20 mM, Tris-HCl (pH 8·0) 100 mM). The
resulting mixture was incubated for 15 min at 65 ºC.
A similar volume of chloroform:isoamilalcohol (24:1)
was added and carefully mixed with the samples until an
emulsion formed. It was then centrifuged for 10 min at
13 000 g, and the DNA in the supernatant was pre-
cipitated with a volume of isopropanol. After a new
centrifugation of 15 min at the same speed, the pellet was
washed in 70% cold ethanol, centrifuged again for 2 min
and dried. It was finally resuspended in 200 µl of ddH
2
O.
PCR amplification was performed with the primers
mrSSU1 and mrSSU3R (Zoller et al. 1999) for the
mtSSU region, and LR0R and LR5 (Vilgalys & Hester
1990; Cubeta et al. 1991) for 28S nrDNA. PCR
reactions were performed under a program consisting of
a hot start at 95 °C for 5 min, followed by 35 cycles at
94 °C, 54 °C and 72 °C (for 45, 30 and 45 s respectively)
and a final 72 °C step for 10 min. PCR products
were checked in 1% agarose gels and positive reactions
were sequenced with one of the PCR primers. Chroma-
tograms were checked searching for putative reading
errors, and these were corrected.
Phylogenetic analyses
BLAST (Altschul et al. 1990) was used to check the
most closely related sequences of the family Roccellaceae in
INSD public databases and a combined 28S rDNA- RPB2
dataset of this family, similar to that in Ertz et al. (2015),
was built. Species of Opegraphaceae were used as out-
groups. Sequences were firstalignedinMEGA5.0
(Tamura et al. 2011) software with its ClustalW application
and then corrected manually. The final alignment included
405/826 (28S rDNA) and 484/844 (RPB2) variable sites.
The aligned loci were loaded in PAUP* 4.0b10 (Swofford
2001) and subjected to MrModeltest v.2.3 (Nylander
2004). The model GTR+Γ+I was implemented in
MrBayes v.3.1 (Ronquist & Huelsenbeck 2003) where a
Bayesian analysis was performed (two simultaneous runs,
six chains, temperature set to 0·2, sampling every 100th
generation) until convergence parameters were met after
c. 7 000 000 generations (after which the Bayesian analyses
were carried out), the standard deviation having fallen
AB
FIG. 1. Coastal rocks in Santa Catarina, Brazil. A, north of Prainha, showing a lower supralittoral zone with algae
where the undescribed Teloschistaceae are growing (arrow); B, south of Prainha, showing grey zone, including
overhanging parts (left arrow) and some yellow supralittoral Teloschistaceae (right arrow). In colour online.
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below 0·01. Finally, a full search for the best-scoring
maximum likelihood tree was performed in RAxML
(Stamatakis 2006) using the standard search algorithm
(data partitioned, 2000 bootstrap replications).
Significance threshold was set above 0·95 for posterior
probability (PP) and 70% bootstrap proportions (BP).
Results
Crustose lichens were found to be abun-
dantly present on the rock faces investigated.
Three distinct lichen zones were found: a
littoral black zone, a supralittoral yellow zone
and a grey zone with species restricted to
either exposed granite, dry overhangs, damp
overhangs or places subjected to run-off.
Therefore the whole range of variation in
microhabitat was found to be present.
In the littoral black zone, only one species
was found, viz. Stigmidium marinum (Deakin)
Swinscow.
Just above the black zone (Fig. 1A), in the
lower supralittoral, there is a zone containing a
number of Teloschistaceae with thallus colours
predominantly yellow to orange, hence the
name “yellow zone”. Several different addi-
tional species were found that are yet to be
determined. The yellow Teloschistaceae that
occur slightly higher up in the intermediate
supralittoral belt could all be identified as
species which are known from other areas in
Brazil.
In the grey zone (Fig. 1B), the higher
supralittoral belt, several microhabitats can
be distinguished such as exposed granite, dry
overhangs, damp overhangs or places sub-
jected to run-off. Dry overhangs of maritime
rocks in at least warm-temperate regions
usually abound with Arthoniales, especially
Roccellaceae. The overhangs in southern
Brazil are no exception to this, with species of
Dendrographa,Enterographa,Lecanographa,
Opegrapha s. lat. and Roccellographa. Fruticose
Roccellaceae, which can be abundant in
such places, were absent but Roccellographa
circumscripta (Leight.) Erzt & Tehler (reported
by Gumboski & Eliasaro 2012b)hasa
partly lobate thallus. Several species from
Opegrapha s. lat. were present, of which only
O. lithyrga Ach. could be identified with some
certainty. The taxonomy of the group is still
too poorly known to warrant the description of
the other species. The species of Enterographa
has only pycnidia present on the thalli
and cannot be identified with certainty. The
Dendrographa is different from all other species
TABLE 1. Crustose and microfoliose maritime lichens on the
Santa Catarina coast, Brazil.
Microhabitat Lichen species
Upper grey zone
exposed Diploschistes euganeus (A. Massal.)
Zahlbr.
Dirinaria aegialita (Afzel. ex Ach.)
B. J. Moore
Hyperphyscia adglutinata (Flörke)
H. Mayrh. & Poelt
Lecanora sulfurescens Fée
L. wilsonii Müll. Arg.
Lecidella asema (Nyl.) Knoph
& Hertel
L. carpathica Körb.
L. chodatii (Samp.) Knoph &
Leuckert
Physcia convexa Müll. Arg.
P. erumpens Moberg
Pyxine albovirens (G. Mey.) Aptroot
P. cocoes (Sw.) Nyl.
Rinodina oleae Bagl.
R. oxydata (A. Massal.)
A. Massal.
Sarcogyne privigna (Ach.)
A. Massal.
Scoliciosporum camptosporum
(Vain.) Aptroot
S. umbrinum (Ach.) Arnold
Tephromela atra Fée
Trapelia coarctata (Turner)
M. Choisy
damp overhanging Agonimia opuntiella (Buschardt &
Poelt) Vězda
Endocarpon pallidulum (Nyl.) Nyl.
Flakea papillata O.E. Erikss.
Physcia atrostriata Moberg
Porina chlorotica (Ach.) Müll. Arg.
Rinodina oleae Bagl.
R. oxydata (A. Massal.)
A. Massal.
influenced by run-off Peltula bolanderi (Tuck.) Wetmore
P. clavata (Kremp.) Wetmore
Leptogium isidiosellum (Riddle)
Sierk
dry overhanging Dendrographa austrosorediata
Aptroot & Gumboski
Lecanographa farinulenta (Müll.
Arg.) Egea & Torrente
Opegrapha lithyrga Ach.
Intermediate upper yellow
supralittoral zone
Caloplaca brouardii (B. de Lesd.)
Zahlbr.
C. isidiosa (Vain.) Zahlbr.
C. subvitellina (Müll. Arg.) Zahlbr.
Roccellographa circumscripta
(Leight.) Ertz & Tehler
Lower littoral zone Stigmidium marinum (Deakin)
Swinscow
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Ertz 7564 (BR) Opegrapha vulgata
Tehler 9346 (S) Dictyographa varians
Ertz 12651 (BR) Sparria endlicheri
1.00/100
1.00/100
1.00/100
1.00/100
1.00/100 1.00/100
1.00/100
1.00/100
1.00/100
1.00/100
1.00/100
1.00/100
1.00/86
1.00/83
1.00/100
1.00/99
0.98/69
1.00/99
1.00/96
0.99/80
1.00/99
0.92/77
1.00/100
1.00/100
1.00/100
1.00/100
1.00/100
1.00/100
1.00/99
1.00/99
1.00/100
1.00/100
1.00/100
1.00/100
1.00/98
1.00/90
1.00/83
0.99/92
1.00/95
1.00/100 1.00/100
1.00/100
0.99/100
1.00/75
1.00/71
1.00/71
1.00/96
1.00/100
1.00/100
9817 Dendrographa austrosorediata sp. nov.
Ertz 17343 (BR) Cresponea proximata
Frisch 13Jp291 (TNS) Cresponea japonica
0.98/72
Cresponea
Dichosporidium
Erythrodecton
Enterographa
Gyrographa
Psoronactis Austroroccella
Mazosia
Sigridea
Chiodecton
Sagenidiopsis
Isolanactis
Lecanactis
Crocellina
Dendrographa
Ertz 17968 (BR) Dichosporidium brunnthaleri
Frisch 11Ug8 (UPS) Dichosporidium brunnthaleri
Ertz 15330 (BR) Dichosporidium nigrocinctum
Ertz 15378 (BR) Dichosporidium nigrocinctum
Ertz 15920 (BR) Dichosporidium nigrocinctum
Ertz 9908 (BR) Erythrodecton granulatum
Ertz 5041 (BR) Enterographa crassa
Vigneron 104 (BR) Enterographa zonata
Ertz 13875 (BR) Enterographa pitardii
Ertz 14043 (BR) Enterographa pitardii
Ertz 16854 (BR) Enterographa pitardii
van den Boom 46943 (h) Enterographa aff. diaderichiana
Ertz 17330 (BR) Enterographa incognita
Ertz 17350 (BR) Enterographa cf. tropica
Ertz 17351 (BR) Enterographa cf. tropica
Ertz 14196 (BR) Enterographa cf. tropica
Ertz 15684 (BR) Mazosia carnea
Ertz 15686 (BR) Mazosia carnea
Ertz 18554 (BR) Mazosia dispersa
Ertz 9264 (BR) Mazosia paupercula
Frisch 13Jp126 (TNS) Mazosia sp.
Tehler 9110 (S) Sigridea californica
Ertz 16549 (BR) Gyrographa gyrocarpa
Ertz 17434 (BR) Gyrographa gyrocarpa
Ertz 17533 (BR) Gyrographa gyrocarpa
Ertz 17545 (BR) Gyrographa gyrocarpa
Ertz 17584 (BR) Gyrographa gyrocarpa
Ertz 18102 (BR) Gyrographa gyrocarpa
Ertz 16120 (BR) Gyrographa saxigena
Ertz 16853 (BR) Gyrographa saxigena
Ertz 8899 (BR) Psoronactis dilleniana
Ertz 8900 (BR) Psoronactis dilleniana
Tehler 9852 (S) Austroroccella gayana
OTA61795 Chiodecton colensoi
Frisch 12Jp76 (TNS) Chiodecton leptosporum
Ertz 17886 (BR) Chiodecton leptosporum
Ertz 6576 (BR) Chiodecton natalense
Ertz 17986 (BR) Chiodecton sp.
Ertz 18026 (BR) Chiodecton sp.
van den Boom 46932 (h) Chiodecton sp.
van den Boom 46948 (h) Chiodecton sp.
Ertz 18264 (BR) Sagenidiopsis isidiata
Ertz 13021 (BR) Isalonactis madagascariensis
Ertz 13024 (BR) Isalonactis madagascariensis
Ertz 5068 (DUKE) Lecanactis abietina
Tehler 8550 (S) Lecanactis abietina
Tehler 9926 (S) Lecanactis luteola
Ertz 4780 (BR) Lecanactis borbonica
Ertz 12962 (BR) Lecanactis borbonica
Ertz 17876 (BR) Lecanactis borbonica
Ertz 7995 (BR) Lecanactis submollis
Frisch 11Ug37 (TNS) Lecanactis submollis
Kantvilas 156/11 (UPS) Lecanactis mollis
Ertz 15561 (BR) Lecanactis epileuca
Ertz 15922 (BR) Lecanactis epileuca
Ertz 15911 (BR) Lecanactis epileuca
Tehler 9310 (S) Crocellina cinerea
Tehler 9342 (S) Crocellina cinerea
Tehler 9353 (S) Crocellina cinerea
Rammeloo s.n. (BR) Gyronactis asiatica
Robertson 8970 (S) Dendrographa alectoroides
Tehler 9104 (S) Dendrographa leucophaea
Tehler 9131 (S) Dendrographa conformis
Ertz 12413 (BR) Dendrographa franciscana
Ertz 13556 (BR) Dendrographa decolora
Frisch 12Jp11 (TNS) ‘Roccellina nipponica’
Ertz 13826 (BR) Dendrographa decolorans
Ertz 14008 (BR) Dendrographa decolorans
Ertz 14063 (BR) Dendrographa decolorans
Tehler 9048 (S) Dendrographa decolorans
Ertz 14059 (BR) Dendrographa latebrarum
FIG. 2. Phylogenetic position of Dendrographa austrosorediata (shown in bold). Consensus phylogram obtained
in MrBayes after the analysis of a combined 28S rDNA and RPB2 dataset. Bayesian PP (set above 0.95) and
ML BP (set at 70%) are given adjacent to nodes. Only nodes supported by both analyses are shown.
Continued on following page.
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Ertz 18179 (BR) Syncesia byssina
van den Boom 43811 (h) Syncesia depressa
Tehler 8720 (S) Syncesia farinacea
van den Boom 43836 (h) Syncesia farinacea
van den Boom 43837 (h) Syncesia farinacea
Tehler 10139_19 (S) Syncesia hawaiiensis
Tehler 10154_25 (S) Syncesia hawaiiensis
Ertz 15904 (BR) Syncesia glyphosoides
Ertz 15905 (BR) Syncesia glyphosoides
van den Boom 44378 (h) Syncesia psaroleuca
van den Boom 44526 (h) Syncesia sp.
van den Boom 44529 (h) Syncesia sp.
Ertz 11059 (BR) Syncesia intercedens
Ertz 12966 (BR) Syncesia madagascariensis
Tehler 9727 (S) Syncesia aff. palmensis
Ertz 10000 (BR) Syncesia aff. palmensis
Ertrz 14230 (BR) Syncesia aff. palmensis
Ertz 14766 (BR) Syncesia aff. palmensis
Ertz 17987 (BR) Syncesia mascarena
Ertz 18052 (BR) Syncesia mascarena
Tehler 9533 (S) Syncesia myrticola
Ertz 16273 (BR) Syncesia myrticola
Ertz 10555 (BR) Syncesia myrticola
Ertz 16274 (BR) Syncesia myrticola
Ertz 16659 (BR) Syncesia myrticola
Ertz 16857 (BR) Syncesia myrticola
Ertz 12059 (BR) Syncesia graphica
Tehler 8622 (S) Syncesia leprobola
Ertz 12054 (BR) Syncesia leprobola
van den Boom 43790 (h) Syncesia leprobola
van den Boom 43792 (h) Syncesia leprobola
van den Boom 43818 (h) Syncesia leprobola
Tehler 9333 (S) Syncesia socotrana
Tehler 9347 (S) Syncesia socotrana
Tehler 8451 (S) Roccellina accedens
Tehler 8432 (S) Roccellina suffruticosa
Tehler 8408 (S) Roccellina cerebriformis
Tehler 8355 (S) Roccellina chalybea
Tehler 9922 (S) Roccellina flavida
Tehler 8375 (S) Roccellina limitata
Tehler 8452 (S) Roccellina mahuiana
Tehler 8372 (S) Roccellaria mollis
Tehler 8430 (S) Roccellina terrestris
Tehler 8431 (S) Roccellina nigricans
Tehler 7721 (S) Roccella hypomecha
Tehler 8368 (S) Roccellina inaequabilis
Tehler 8369 (S) Roccellina lutosa
Tehler 8360 (S) Roccella tinctoria
Tehler 8350 (S) Roccella portentosa
Ertz 15695 (BR) Roccellina leptothalla
Ertz 18586 (BR) Roccellina leptothalla
Ertz 18616 (BR) Roccellina leptothalla
Ertz 15656 (BR) Vigneronia cypressi
Ertz 15677 (BR) Vigneronia cypressi
Ertz 18136 (BR) Vigneronia cypressi
Ertz 18320 (BR) Vigneronia cypressi
Ertz 12060 (BR) Vigneronia spieri
Ertz 18394 (BR) Vigneronia spieri
Tehler 9051 (S) Ocellomma picconianum
Tehler 9815_11 (S) Ocellomma picconianum
Diederich 16390 (BR) Ocellomma picconianum
Ertz 17132 (BR) Ocellomma picconianum
Ertz 17143 (BR) Ocellomma picconianum
Vigneron 75 BR Pseudoschismatomma rufescens
Ertz 17388 (BR) Pseudoschismatomma rufescens
Ertz 17397 (BR) Pseudoschismatomma rufescens
Tehler 9703 (S) Dirina angolana
Tehler 8524 (S) Dirina astridae
Tehler 9614 (S) Dirina cretacea
Tehler 8671 (S) Dirina approximata
Tehler 9151 (S) Dirina catalinariae
Ertz 13215 (BR) Dirina madagascariensis
Tehler 10219 (S) Dirina insulana
Tehler 8888 (S) Dirina badia
Tehler 9301 (S) Dirina arabica
Tehler 9300 (S) Dirina immersa
Tehler 10307 (S) Dirina canariensis
Tehler 9021 (S) Dirina ceratoniae
Tehler 9608 (S) Dirina massiliensis
Tehler 9015 (S) Dirina candida
Tehler 8653 (S) Roccella albida
Tehler 8667 (S) Roccella galapagoensis
Tehler 8656 (S) Roccella margaritifera
Tehler 8655 (S) Roccella nigerrima
Tehler 9190 (S) Roccella gracilis
Tehler 8272 (S) Roccella elisabethae
Tehler 8309 (S) Roccella canariensis
Tehler 8144 (S) Roccella tuberculata
Tehler 8221 (S) Roccella phycopsis
Tehler 8163 (S) Roccella allorgei
Tehler 8255 (S) Roccella fuciformis
Tehler 8140 (S) Roccella maderensis
Tehler 8568 (S) Roccella sinensis
Tehler 8506 (S) Roccella boryi
Tehler 8870 (S) Roccella montagnei
Tehler 9039 (S) Diromma dirinellum
Tehler 9053 (S) Diromma dirinellum
Ertz 14011 (BR) Diromma dirinellum
Tehler 8551 (S) Schismatomma pericleum
Tehler 10660 (S) Schismatomma pericleum
Vondrák 8455 (BR) Schismatomma cf. umbrinum
Tehler 10652 (S) Schismatomma cf. umbrinum
1.00/100
1.00/100
1.00/100
1.00/100
0.99/85
1.00/100
1.00/100
1.00/97
0.95/71
1.00/98
1.00/89
1.00/99
1.00/100
1.00/100
1.00/98
1.00/98
1.00/100
1.00/97
1.00/83
1.00/87
1.00/83
0.81
1.00/100
1.00/81 0.98/100
1.00/100
1.00/100
1.00/99
1.00/100
1.00/100
1.00/100
0.99/88
1.00/100
1.00/88
1.00/94
1.00/91
1.00/97
1.00/72
1.00/95
1.00/98
1.00/100
0.96/89
1.00/100
1.00/99
0.98/93
1.00/100
1.00/93
0.98/80
0.99/71
1.00/100
1.00/100
1.00/95
1.00/95
1.00/100
1.00/100
1.00/88
1.00/100
1.00/100
1.00/94
1.00/99
1.00/97
1.00/100
0.5
Schismatomma pericleum
Syncesia
Roccelina
Vigneronia
Ocellomma
Pseudoschismatomma
Dirina
Roccella
Schismatomma cf. umbrinum
FIG.2.Continued from facing page.
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accepted in the genus and is therefore descri-
bed here as new. It is one of the few crustose
species in the genus.
Places that can be classified as wet over-
hangs are generally shaded and damp.
Lichens growing here are usually only partly
maritime. In southern Brazil, mostly ubiqui-
tous cosmopolitan species such as Agonimia
opuntiella (Poelt & Buschardt) Vězda,
Endocarpon pallidulum (Nyl.) Nyl., Flakea
papillata O. E. Erikss., Physcia atrostriata
Moberg, Porina chlorotica (Ach.) Müll.
Arg., Rinodina oleae Bagl. and R. oxydata
(A. Massal.) A. Massal. were found in this
microhabitat.
In places with frequent run-off, lichens
with cyanobacteria were dominant. The
pantropical Peltula bolanderi (Tuck.) Wetm.,
P. clavata (Kremp.) Wetm. and Leptogium
isidiosellum (Riddle) Sierk were present,
as well as several so far unidentified Lichinaceae
(i.e. probably one or more Pterygiopsis species).
The exposed granitic rocks of the grey zone
are the richest in lichen species, just as else-
where in the world. Here the majority of the
macrolichens of the coastal rock also occur.
The simple explanation is that the environ-
ment is less extreme and harbours many
species that can also grow inland. Most of the
crustose species identified so far seem to be
cosmopolitan, while a considerable number
of the reported macrolichens had a more
restricted distribution. One Ramalina was
present, viz. R. gracilis (Pers.) Nyl. Ramalina
is often the dominant lichen on coastal rocks
around the world and the species are often
partly endemic, just as here, which is rather
unusual among lichens. The crustose and
microfoliose species found by us are listed in
Table 1. The list is far from complete as only
a small number of coastal cliffs have been
systematically sampled and not all species
could be identified.
Some of the lichens encountered, including
all species in the lower yellow (lower supra-
littoral) zone, could not be identified as any
described species. One undescribed species
was found in overhangs. It was sequenced as
it was sterile. ITS (KY986702), 28S rDNA
(KY986703) and mtSSU (KY986704)
sequences were obtained, but we failed to
obtain an RPB2 sequence. In the analysis
only 28S rDNA was used for our specimen,
but RPB2 was used for other taxa from which
it was available. Two analyses were carried
out: a Bayesian simulation in MrBayes and a
ML analysis in RAxML. The tree depicted
(Fig. 2) is the Bayesian tree, with nodes
annotated with PP (Bayesian) and BP (ML)
values.
The new species clustered with one of the
few other known crustose Dendrographa
species, D. latebrarum (Ach.) Ertz & Tehler
(Fig. 2), and is described here in that genus.
Dendrographa austrosorediata Aptroot
& Gumboski sp. nov.
MycoBank No.: MB 820870
Saxicolous Dendrographa from overhanging coastal rock,
with a crustose, flat to partly curling or blister-like thallus
which is much dissected, whitish grey, 0·1–0·2 mm thick;
surface minutely densely cracked, rimose, with cracks
intersecting every c. 0·1 mm; soredia whitish to bluish
grey but asymmetrically blackened in the direction facing
the light, originating on the thallus surface, in initially
discrete convex soralia.
Type: Brazil, Santa Catarina State, municipality of São
Francisco do Sul, S of Prainha, 26°14'06''S, 48°30'05''W, on
coastal granite, c. 10 m alt., 8 October 2015, M. E. S. Cáceres
& A. Aptroot 27936 (JOI—holotype; ABL—isotype).
(Fig. 3)
Thallus crustose, individual thalli not dis-
tinguishable, colonies indeterminate and cov-
ering large areas, flat to partly curling up or
blister-like, much dissected, epruinose or
slightly pruinose, whitish grey, 0·1–0·2mm
thick; surface minutely densely cracked, rimose
with cracks intersecting at each c. 0·1mm;
cortex 20–40 μmthick;medulla not differ-
entiated, sometimes surrounded by a thin,
black prothalline line; alga trentepohlioid,
c.6–11 μmdiam.Soredia present, whitish to
bluish grey, but asymmetrically blackened in
the direction facing the light, originating on the
thallus surface in initially discrete convex sor-
alia of c.0·2–0·4mm diam. and up to 0·3mm
high, later covering most of the thallus; soredia
round to ellipsoid, c.18–32 μmdiam.,surface
hyphae partly blackened.
Apothecia unknown.
Pycnidia not observed.
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Chemistry. Thallus and soredia C+ red.
TLC: erythrin & lecanoric acid.
Ecology and distribution. On maritime
granite, on overhanging rockfaces. Known
only from Brazil.
Discussion
In the new phylogenetic classification
of the Roccellaceae (Ertz et al. 2015), some
crustose species are classified in the genus
Dendrographa Darb. which traditionally
contained only a few fruticose species. The
available material of the new species is sterile
but characteristic enough to be described and
recognized, even in the field. Its placement in
Dendrographa is based on a phylogenetic
analysis (Fig. 2) using two genes. We present
here an analysis of a larger group because
ITS produced no matches, with the genera
Roccellina and Dendrographa as top results,
while LSU was only 90% similar to both
A
B
C
FIG.3.Dendrographa austrosorediata (holotype). A, thallus; B, detail of thallus; C, soredia. Scales: A =1 mm;
B=0·5 mm; C =10 μm. In colour online.
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genera. Without a broader analysis, it was
not possible to decide whether it was a
Roccelina or a Dendrographa after the BLAST
results (or whether it even belonged to
another genus). Moreover, the noise in the
LSU sequence could alter BLAST results
significantly, making it too risky to decide if it
was a Dendrographa without conducting an
analysis.
Stigmidium marinum (Fig. 4) is a somewhat
surprising find, especially as it is generally
regarded as a lichenicolous fungus. It is
usually reported as an obligate parasite on
some littoral Verrucaria species. However,
it has been repeatedly observed occurring as a
free-living lichen and is treated as such by van
Herk & Aptroot (2004). Its occurrence in the
littoral zone of southern Brazil, thousands of
kilometres away from the nearest known
occurrence of any purported host, proves
once more that this is a lichenized fungus.
It is lichenized with Dilabofilum.
MESC thanks CNPq (Conselho Nacional de Desen-
volvimento Científico e Tecnológico) for a research
grant (Processo 311706/2012-6); AA thanks the Sticht-
ing Hugo de Vries-fonds for a travel grant; ELG thanks
the Universidade da Região de Joinville –UNIVILLE for
support provided for the event. We are grateful to Pablo
Alvarado (ALVALAB, Santander) for carrying out the
sequencing and phylogenetic analysis.
A
BC
FIG.4.Stigmidium marinum (Cáceres & Aptroot 27943). A, thallus; B, flattened perithecium in surface view revealing
Dilabofilum algal cells around the margin; C, ascospore. Scales: A =0·1 mm; B =25 μm; C =5μm. In colour online.
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