This study revises the cosmopolitan saxicolous lichen species of Rinodina (Ascomycetes) in Australia and New Zealand, including maritime southern South America and subantarctic islands, based on morphological, anatomical and molecular data. The cosmopolitan genus Rinodina used to comprise 200 species (Hawksworth et al. 1995). In the meantime, approximately 250 species are known (Kirk et al. 2001), which are widely distributed in both hemispheres, from (ant)arctic to tropical regions. 29 saxicolous species occur in Australia and New Zealand. The author expands on the revision of saxicolous species of Rinodina (with exception of the xanthone containing species R. thiomela and R. xanthomelana with yellow thalli) in southern Africa, carried out by Matzer & Mayrhofer (1996).
The genus Rinodina belongs to the Ascomycetes, the biggest fungus group worldwide, where the ascus represents the most important character. The usually lichen-forming order of the Lecanorales includes 40 families, being characterised by apothecial ascomata and very variable ascospores. Within the family of the Physciaceae nom. cons. (Eriksson et al. 2003), the large genus Rinodina is heterogeneous (e.g. Hafellner et al. 1979, Mayrhofer 1982) because of a multitude of ascospore types. Different ascus types were described by Rambold et al. (1994). In recent years these results are confirmed by molecular phylogenetic studies by Grube & Arup (2001) and Helms et al. (2003).
Preface 5
I General information 7
1. Areas of investigation 7
2. Geological history of Australia and New Zealand 8
3. Climate 9
4. Ecology 10
II Anatomical and morphological studies: basics for the traditional species concept 11
1. Introduction 11
2. Material and methods l2
3. Morphology, anatomy and chemistry of the taxa l4
3.1 Thallus 14
3.2 Apothecia 15
3.3 Ascospores 17
3.4 Chemistry l9
4. Keys to all known saxicolous taxa of Rinodina in Australasia 20
4.1 Total key - mainly Australia and New Zealand 20
4.2 Partial key - Australia and Tasmania 23
4.3 Partial key - New Zealand 25
5. The taxa of the genus Rinodina in Australia and New Zealand 26
Rinodina bischoffii (Hepp) A.Massal 26
Rinodina blastidiata Matzer & H. Mayrhofer 32
Rinodina cacootina Zahlbr 39
Rinodina confragosa (Ach.) Körb 45
Rinodina confragosula (Nyl. in Cromb.) Müll.Arg 47
Rinodina gallowayii H. Mayrhofer 53
Rinodina gyrophorica Kaschik sp. nov 55
Rinodina herteliana Kaschik sp. nov 57
Rinodina immersa (Körb.) Arnold 59
Rinodina jamesii H.Mayrhofer 62
Rinodina luridata (Körb.) H. Mayrhofer, Scheid. & Sheard 65
Rinodina milvinodes H. Mayrhofer 68
Rinodina moziana (Nyl.) Zahlbr. var. moziana 70
Rinodina moziana var. parasitica Kaschik & H. Mayrhofer var. nov 76
Rinodina murrayii H.Mayrhofer 78
Rinodina nigricans H.Mayrhofer 82
Rinodina occulta (Körb.) Sheard 84
Rinodina oleae Bagl 86
Rinodina oxydata (A.Massal.) A.Massal 94
Rinodina peloleuca (Nyl.) Müll.Arg 101
Rinodina ramboldii Kaschik sp. nov 105
Rinodina reagens Matzer & H. Mayrhofer 108
Rinodina striatitunicata Matzer & H. Mayrhofer 111
Rinodina subcrustacea (Müll.Arg.) Zahlbr 114
Rinodina substellulata Müll.Arg 116
Rinodina subtubulata (C.Knight) Zahlbr 117
Rinodina williamsii H.Mayrhofer 119
Excluded species 122
'Rinodina' brattii H.Mayrhofer 122
Rinodina compensata (Nyl.) Zahlbr 125
'Rinodina' insularis (Arnold) Hafellner 126
Rinodina kozakensis (Vain.) Zahlbr 129
'Rinodina' lecideina H.Mayrhofer & Poelt 129
'Rinodina' microspora Dodge 130
'Rinodina' otagensis (Zahlbr.) H. Mayrhofer 130
'Rinodina' procellarum (Massal.) H. Mayrhofer 130
'Rinodina' tubulata (Knight) Zahlbr 130
6. Discussion 131
6.1 Conclusions 131
6.2 New characters l32
6.3 Sociology 133
6.4 Distribution 135
III Phylogenetic analysis based on nuclear ITS rDNA 141
1. Introduction 141
2. Material and methods 142
2.1 Material 142
2.2 Molecular methods 144
2.3 Phylogenetic analyses 149
3. Results 150
4. Discussion 152
Summary 155
References 157
"The latest monograph of Rinodina from the group of lichen taxonomists under the direction of H. Mayerhofer at Graz University concentrates on the saxicolous species of Australia and New Zealand and includes all known taxa from these areas. In addition to the 29 species previously known, 6 new species had to be described (2 from Australia, 4 from New Zealand); 9 species, excluded from the genus but cannot be transferred to other genera at this time, are also treated fully. Each species has a full-length description which also includes a discussion and a series of photographs illustrating the spore ontogeny, but there are no photographs of the habitus. A full list of specimens examined is given and distributions are illustrated by maps. The lists of studied specimens also include material from other parts of the World, especially the Southern Hemisphere (e.g. Chile, South Africa), but also Austria etc. The classical treatment of taxa based mainly on spore ontogeny is supported by some new characters and by phylogenetic analysis based on nuclear ITS rDNA. The new characters take account of ascospore types and pigments. Some of the classical spore types in Rinodina grade into other types. Regarding pigments, it was found that Bagliettoana-green is relatively common but at least 3 other pigments occur, one identified as Arceutine-yellow and two as yet unidentified. The phylogentic analysis is based on the alignment of 71 taxa (40 from GenBank and 31 newly generated). For the convenience of users three keys are given, one for all saxicolous species of Rinodina s.l. in the studied area and two for Australia and New Zealand exclusively. Some minor criticisms might be added. Some more biogeographical discussion would be of interest even at this stage of knowledge. In the introduction on page 7 it is noted that according to a paper by McCarthy one third of Australian lichen taxa are considered to be endemic - but there is no discussion of this for the species under study, nor is a comparison made between Australia and New Zealand. The first sentences in the chapter on materials and methods giving figures on studied material and the storage of voucher specimens (p. 12) are identically repeated in the summary (p. 155). With this body of information, the monograph will certainly serve its purpose and is a further important step forward in the knowledge of this large genus."
– International Lichenological Newsletter 39(2), page 7-8
"This study, which has been undertaken with the supervision of the world’s leading authority on the genus, Helmut Mayrhofer (Graz), takes a traditional approach, but also includes some molecular phylogenetic information. The bulk of the work, based on 814 collections many of which were made by Mayrhofer, comprises descriptions, distribution maps, photomicrographs of the ascospores that are so important in the genus, and lists of specimens examined. Full bibliographic and type information is supplied. In all 21 taxa are recognized in the region, of which four are described as new to science; some new synonymies are proposed, and some names are excluded for different reasons. Particular attention was paid to the pigments, using HPLC and HPTLC as well as traditional TLC. The molecular studies, based on freshly sequenced material and also data downloaded from GenBank, used ITS rRNA sequences showed ascus types and hypothecium pigmentation to be highly conserved features, corroborating the recognition of “Physcia-“ and “Buellia-type” groups within the “genus”. The genus is paraphyletic with the sister groups often foliose or fruticose genera, but the ascospore types do not support larger monophyletic groups. In due course it is inevitable that formal taxonomic changes will be made amongst these lichens, but that should await the analysis of many more specimens on a worldwide scale. For the moment, however, the author has produced a revision that will enable these taxa to be reliably identified for the first time."
– Mycotaxon vol. 102, 2007