Flora of New Zealand Series

PREFACE

"Every botanist who prepares a Flora starts from the same standpoint reached by his predecessors in the same field," so wrote Thomas Cheeseman in 1906 in the preface to his Manual of the New Zealand Flora. Some 50 years later, in the Preface to the first volume of the current Flora of New Zealand series, H.H. Allan (1961) defined a Flora as "…a compilation, a drawing together by a competent taxonomist of information already published." Edgar & Connor (2000) remark that the modern requirements of a Flora are: "… descriptions, keys, uncertainties, modern classification, notes, references to allied studies, and to the relevant revisions". A "lichen Flora" of a region, to plagiarise Edgar & Connor, "…should inform the region about the [lichens] resident in that region; it should tell the rest of the world about the [lichens] of that region, especially of those with which they are unacquainted; and it should tell the world about the variation, behaviour, ecology and distribution of the [lichens] in that regional environment". To this of course we must now add molecular systematics and what it can tell us of the major fungal lineages and their ability to form obligate (or optional) symbiotic associations with green algae, cyanobacteria, or with both photobionts, in the formation of the mixed associations that we call lichens. Recent results suggest that "…lichens evolved earlier than believed, and that gains of lichenization have been infrequent during Ascomycota evolution, but have followed multiple independent losses of the lichen symbiosis..." (Lutzoni et al. 2001). In a fascinating recent paper, Wedin et al. (2004) have shown that in the case of the saprophytic (non-lichenised) fungus Stictis and the closely related lichenised genus Conotrema, the same taxon may exist in both saprophytic and lichenised states. Thus saprotrophy and optional lichenization allow separate individuals of the same taxon to exploit different niches (bark or decorticated wood) in the landscape. It is postulated that environmental plasticity is an overlooked strategy in fungi adapted to unpredictable successional ecosystems. This has implications for many lichen groups, especially those in the families Lobariaceae and Pannariaceae where photobiont versatility is a well-known phenomenon. Phylogenetic relationships within the Ascomycota, as part of progress towards assembling the "Fungal Tree of Life", are discussed in Lutzoni et al. (2004).

Lichens continue to be misunderstood (Hawksworth 1997). Theoretically the entities that we recognise and name as lichens are not organisms at all, and they have no name – the name that we give them referring correctly to the fungal component of the symbiotic association, with the photosynthetic organism(s) with which the lichenised fungus is associated having its own name, and belonging moreover to a different biological kingdom! Yet lichens are, in many ways, plant-like (Sanders 2001) and have been most commonly studied by botanists (rather than mycologists) in botanical departments of museums and/or universities for many generations, and there is a strong and long "botanical" tradition in lichenology (Hawksworth 1997; Rambold & Triebel 1999). Communities of lichens, from a fencepost to a regional descriptive catalogue, tend to be still given the name lichen flora, when the correct term is lichen mycobiota, and regional Floras (including the present one!) perpetuate this "plant-based" view. But knowledge of lichenised fungi, and especially of the generic concepts in lichenised and lichenicolous ascomycetes, has advanced dramatically in the last two decades (Hawksworth 1999; Rambold & Triebel 1999; Grube 2004; Lutzoni et al. 2004; Will-Wolf et al. 2004), and it now seems appropriate that future, online versions of this lichen "Flora" should be accommodated in the series Fungi of New Zealand/Ngā Harore o Aotearoa. Indeed, the current checklist of New Zealand Ascomycete names in the first volume of this series contains most of our lichen names (Pennycook & Galloway 2004).

In the current series of Flora of New Zealand, which comprises five volumes devoted to flowering plants, conifers and ferns, two to desmids and one to lichens (Galloway 1985) none, except the lichens, has yet appeared in a revised version. The first edition of Flora of New Zealand Lichens was primarily a herbarium-based compilation, prepared during my time at the Natural History Museum in London. The text for this work was completed in 1983, just when Northern Hemisphere lichenologists were starting to question generic limits of some of the large and unwieldy genera such as Lecidea and Parmelia. Its publication acted as a spur to further investigations, not only of New Zealand lichenology, but also of Southern Hemisphere lichenology, and in the years since its publication, knowledge of Southern Hemisphere and of New Zealand lichens has increased dramatically. Subsequent to publication of Flora of New Zealand Lichens, a series of lichen workshops were held in various parts of the country, a number of enthusiastic local lichenologists emerged, closer linkages with lichenologists in Australia were forged, and the number and standard of local and regional lichenological publications increased dramatically (Johnson & Galloway 2002). It was in fact obvious by the early 1990s that Flora of New Zealand Lichens was increasingly out of date and in need of revision, to accommodate both the major changes in lichen taxonomy and the increasing numbers being added to the New Zealand lichen mycobiota, either as additions of existing genera and species or as newly described taxa. I returned to New Zealand to live in November 1994, and in 1995 started contract work for Manaaki Whenua – Landcare Research on a variety of applied projects. In 1996 I began work on a second edition of Flora of New Zealand Lichens, supported by 60% funding from the Foundation for Research, Science and Technology (FRST) as part of the Plant Systematics Programme of Landcare Research. This period coincided with a dramatic upturn in interest in South Pacific lichenology stimulated by the appearance of regional lichen Floras, checklists, keys, and popular works on lichenology (see for example Grgurinovic 1992, 1994; Malcolm & Galloway 1997; Kantvilas & Jarman 1999; Gilbert 2000; Malcolm & Malcolm 2000, 2001; Lumbsch et al. 2001; McCarthy 2001; Kantvilas et al. 2002; McCarthy 2003, 2005; Galloway 2004a; McCarthy & Malcolm 2004).

Although the 1985 Flora must now be considered as pretty much a "trial run", in many instances it is still a useful compilation and does show where we have advanced from in recent years. Twenty years on from there, the present Flora is still just a snapshot of New Zealand lichenology, though now one from near the beginning of the 21st century. Like its predecessor, it has its share of uncertainties and conjectures, but it now signals, rather more strongly than heretofore, where we might usefully travel in several areas in the future (see below).

The layout of this Lichen Flora follows that of the first edition with the genera being arranged alphabetically and incorporating the following descriptive elements: (1) A genus description in which essential characters are outlined. Where the existing, firstedition genus description is still applicable it is referred to, together with reference to a more recent complete description where it is thought necessary or helpful, e.g. for Brigantiaea ["… Flora (1985: 40). See also Hafellner (1997: 40)."]. (2) A discussion of the genus with references to the literature pertinent to it, together with the currently accepted ordinal and/or family affiliation (Eriksson et al. 2004; Pennycook & Galloway 2004; Eriksson 2005), and an indication of the species diversity both within New Zealand and worldwide. (3) A dichotomous key to species in the genus known to occur in New Zealand.

A standard arrangement of species description is adopted (as in the first edition) and includes thalline characters, apothecial characters, pycnidial characters (when known), and chemical characters (when known). For useful general technical accounts of lichen morphology and anatomy see Purvis et al. (1992: 4–7), Büdel & Scheidegger (1996) and Malcolm & Galloway (1997: 183–189), with a recently published compendium of techniques for handling lichens experimentally (Kranner et al. 2002) catering for lichenological laboratories needing a functional working manual of techniques. As in the previous edition of the Flora, both thalline colour tests (see Malcolm & Galloway 1997) and thin-layer chromatography of acetone extracts were routinely used, following standardised methods (Culberson 1972) with later modifications (see Orange et al. 2001). An overview is available (Elix 1996), and a comprehensive compendium of chemical data on lichens is given in Huneck & Yoshimura (1996).

Full bibliographic citations are listed for all currently accepted taxa and their basionyms, and for all heterotypic, and homotypic, synonyms based on taxa originally described from New Zealand. Every effort has been made to make the listing of synonyms as complete as possible to assist future studies. Synonyms are printed in italics both in the text and index. Citation of genera and dates of publication follow Index Nominum Genericorum (Farr et al. 1979a, 1979b, 1979c) and the 9th edition of Ainsworth & Bisby's Dictionary of the Fungi (Kirk et al. 2001). Author abbreviations accord with Kirk & Ansell (1992).

Holotypes and/or lectotypes of names based on New Zealand material are cited, together with all known provenances of isotype, isolectotype or syntype material, since much of New Zealand's extant lichen types are widely scattered, and it was considered important to have available in one place as comprehensive a listing of type material as possible to assist future systematic work. Where New Zealand lichen taxa are already lectotypified in the literature, reference to the typification is given thus "…[fide Galloway (1985a: 71)]". In a number of cases, earlier lectotypifications are overturned where authentic holotype collections have subsequently come to light after the original lectotypification was made. These cases are appropriately noted in the text. Conserved names are in accord with the list published in the Saint Louis Code (Greuter et al. 2000).

Biogeographical elements recognised in the New Zealand lichen mycobiota follow Galloway (1985, 1996).

An indication of the known distributions of lichen (and lichenicolous fungus) taxa in New Zealand is given after the description, chemistry, and biogeographical designation. Provincial names used are those shown on the endpaper maps at the end of the volumes. Habitat notes are given when such information is available, together with a brief indication of the known world distribution of the taxon under discussion.

Lichenicolous fungi are included in this Flora for the first time, since these specialised fungi, adapted or co-evolved to grow on the thallus or apothecia of lichens, are traditionally studied by lichenologists (Clauzade et al. 1989; Hawksworth 2003; Lawrey & Diederich 2003; Diederich 2004, Gams et al. 2004). Many taxa are now known from New Zealand, but the number of genera and species can be expected to increase dramatically once serious studies on these fascinating fungi are undertaken here.

The Lichen Flora is supported by a comprehensive bibliography, a glossary of lichenological terms, and an index of accepted taxa and synonyms.

Conservation of lichen diversity in New Zealand is still very much a neglected field. In 1985 I wrote "…The astonishing diversity of habitats that exist between western and eastern coasts, and between the northern and southern extremities of New Zealand will ensure many lichenological discoveries for generations to come. However, this remarkable richness and diversity of New Zealand's lichens is an easily destroyed phenomenon and already atmospheric and terrestrial pollution, hydro-electric power development, changes in land use and especially in management practices in native forests have placed, (and will continue to do so) many lichen communities at risk" (Galloway 1985: xxiii). Unfortunately this situation still holds today. The rich, lowland cryptogamic communities developed on Sphagnum bogs on the Southland Plains, that first attracted me to lichens in the 1950s, have largely disappeared as wetlands have been drained for farmland, and the diverse, lichen-dominated crusts of the dry, inland lake basins of Canterbury have been dramatically reduced by the rapid growth of Hieracium in these areas in recent years. Further threats to lichen communities are discussed in Johnson & Galloway (2002) in a general overview of lichens and their conservation needs in New Zealand, a treatment soon to be published by the New Zealand Department of Conservation. However, and with considerable cause for optimism, the resilience of lichens in New Zealand habitats seriously affected by major pollution events is remarkable. For example, monitoring of individual lichens at the edge of a "lichen desert" surrounding a North Island metal-processing plant, over three consecutive years after installation of emission controls, showed a rapid response to improved air quality. Total area of lichen thalli doubled or trebled (depending on the species), numbers of newly established lichen individuals doubled, and species numbers increased by 30–40% within 3 years. Some improvements were even detected within a year! Within the area previously devoid of lichens, re-establishment has begun with up to eight species recorded (Johnson et al. 1998). For a wider, global perspective on conservation of lichens see Scheidegger et al. (1995).

In the present work, 310 lichen-forming and 44 lichenicolous genera (a total of 354 genera) are treated, supported by descriptions of 1706 taxa. This figure, although c.70% greater than the number of taxa reported from New Zealand in 1985, still represents probably only about 75–80% of the lichens to be eventually found here. It compares, for example, with 303 genera and 1743 taxa recorded from Great Britain and Ireland (Coppins 2002); 307 genera and 1608 taxa recorded from Japan (Kurokawa 2003); 396 genera and 3227 taxa from Australia (McCarthy 2005); and 449 genera and 2968 taxa known from Fennoscandia (Santesson et al. 2004). With the currently estimated number of lichens known worldwide standing at 18 000 species (Sipman & Aptroot 2001), New Zealand has about 10% of the world's lichen mycobiota – a significant diversity for its geographical size. A current source of real concern worldwide is the decline in numbers of professional taxonomists in general (see for example Godfray & Knapp 2004; Raven 2004), and of lichenologists in particular (Smith 1997; Hawksworth 1999). Given New Zealand's rich lichen biodiversity, and four decades of steady growth in knowledge of that biodiversity, it is imperative that lichen-based studies in this country are both encouraged and sustained into the future.

Thirty years ago, when I was collecting lichens in the King Country, a farmer remarked to me, "It's all very well collecting these things, but what use are they to anyone?" I have pondered that question many times since, even though as a systematist I have a vested interest in providing correct names for taxa in our lichen mycobiota before suggesting possible or probable uses of lichens. However, we have now reached a stage in lichenology where the supremacy of taxonomy must yield to other, more applied disciplines, as we seek to find out exactly where lichens "fit in" to our landscapes and ecosystems. We can now answer the King Country farmer with the rejoinder that nowadays lichens have immediate and potential application in a variety of fields, including: (1) Lichens as biomonitors of terrestrial and atmospheric pollution [there is a huge literature on this subject, but for data on New Zealand environments see Johnson et al. (1998)]. (2) Lichens as monitors of global and environmental change and especially of the importance of lichen secondary compounds in photoprotection, light-screening and amelioration of the effects of changes in UVB consequent upon ozone depletion (Galloway 1993; Quilhot et al. 1994, 1998; Bjerke et al. 2003, 2005). (3) Lichens as nitrogenfixers. Diazotrophic (nitrogen-fixing) lichens are of major importance in N-cycling in forest and grassland biomes (Galloway 1995) with 235 species in 41 genera being potential diazotrophs or biological fertilisers. (4) Lichens as sources of bioactive compounds having anti-microbial, anti-viral, cytotoxic, or anti-carcinogenic activity (Perry et al. 1999). (5) Use of lichens as monitors in environmental impact assessment (Seaward 2004) These and other avenues are exciting new directions, along with molecular approaches to systematics, for lichenology in the future in a New Zealand context.

Since this lichen Flora is, in one sense, a great "telephone book" of names (to use Pier Luigi Nimis's telling term for floras and checklists!), Marie Taylor's compendium of fascinating and useful information on the origins and meanings of botanical names of New Zealand plants (including lichens) and their collectors (Taylor 2002) is warmly recommended to all users of this Flora, as also are the books of Feige (1996), and of Wagenitz (1996), dealing with a wider lichenological, and botanical canvas, respectively.

At the time of publication, we are witnessing many reassessments of generic limits as more molecular data become widely available (see for example Lumbsch 2002; Blanco et al. 2004a, 2004b, 2005; Wedin et al. 2004, 2005a, 2005b; Andersen & Ekman 2005; Hawksworth 2005), and undoubtedly this trend will continue with increasing tempo and momentum, requiring at times major adjustments to taxonomy and nomenclature. Fortunately, it is the intention of soon having the present lichen Flora available on the World Wide Web, with the potential for regular updates of systematic, chemical and distributional data, as well as the provision of interactive, illustrated keys, and eventually high-quality illustrations of all lichenised and lichenicolous taxa. This is in line with current thinking on the availability of electronic Floras, keys, checklists, databases, computerised herbaria and their collections, and a host of lichen websites via the Internet (for further information see Nimis & Martellos 2003; Feuerer 2004; Grube 2004; Rambold 2004; Wilkinson & Foster 2004; Will-Wolf et al. 2004). With lichens now emerging as biomonitors of choice in studies of atmospheric and terrestrial pollution and of global environmental change, and considering their importance in nutrient cycling in forests and grasslands and their potential value in habitat restoration studies, it is rapidly becoming clear just how important lichens really are in our New Zealand landscape (Galloway 2004b).

REFERENCES

Allan, H.H. 1961: Flora of New Zealand. Volume I Indigenous Tracheophyta, Psilopsida, Lycopsida, Filicopsida, Gymnospermae, Dicotyledones. Wellington, Government Printer.

Andersen, H.L.; Ekman, S. 2005: Disintegration of the Micareaceae (lichenized Ascomycota): a molecular phylogeny based on mitochondrial rDNA sequences. Mycological Research 109 (1): 21–30.

Bjerke, J.W.; Zielke, M.; Solheim, B. 2003: Long-term impacts of simulated climatic change on secondary metabolism, thallus structure and nitrogen fixation in two cyanolichens from the Arctic. New Phytologist 159 : 361–367.

Bjerke, J.W.; Elvebakk, A.; Domínguez, E.; Dahlback, A. 2005: Seasonal trends in usnic acid concentrations of Arctic, alpine and Patagonian populations of the lichen Flavocetraria nivalis. Phytochemistry 66 : 337–344.

Blanco, O.; Crespo, A.; Divakar, P.K.; Esslinger, T.L.; Hawksworth, D.L.; Lumbsch, H.T. 2004a: Melanelixia and Melanohalea, two new genera segregated from Melanelia (Parmeliaceae) based on molecular and morphological data. Mycological Research 108 : 873–884.

Blanco, O.; Crespo, A.; Elix, J.A.; Hawksworth, D.L.; Lumbsch, H.T. 2004b: A molecular phylogeny and new classification of parmelioid lichens containing Xanthoparmelia -type lichenan (Ascomycota: Lecanorales). Taxon 53 : 959–975.

Blanco, O.; Crespo, A.; Divakar, P.K.; Elix, J.A.; Lumbsch, H.T. 2005: Molecular phylogeny of parmotremoid lichens (Ascomycota, Parmeliaceae). Mycologia 97 : 150–159.

Büdel, B.; Scheidegger, C. 1996: Thallus morphology and anatomy. In : Nash III, T.H. (Ed.) Lichen Biology. Cambridge, Cambridge University Press. Pp. 37–64.

Cheeseman, T.F. 1906: Manual of the New Zealand Flora. Wellington, Government Printer.

Clauzade, G.; Diederich, P.; Roux, C. 1989: Nelikeniğintaj fungoj likenloğaj. Ilustrita determinlibro. Bulletin de la Société linnéenne de Provence. Numéro Spécial 1 : 1–142.

Coppins, B.J. 2002: Checklist of Lichens of Great Britain and Ireland. London, British Lichen Society.

Culberson, C.F. 1972: Improved conditions and new data for the identification of lichen products by a standardized thin-layer chromatographic method. Journal of Chromatography 72 : 113–125.

Diederich, P. 2004: Lichenicolous fungi [including Introduction; Terminology used in lichenicolous fungi; Outline of keys; Key 1 – lichenicolous lichens; Key 2 –lichenicolous basidomycetes; Key 3 – lichenicolous anamorphic ascomycetes; Key 4 – lichenicolous calicioid ascomycetes; Key 5 – lichenicolous perithecioid ascomycetes; Key 6 – lichenicolous apothecioid ascomycetes. In : Nash III, T.H.; Ryan, B.D.; Diederich, P.; Gries, C.; Bungartz, F. (Eds) Lichen Flora of the Greater Sonoran Desert Region. Vol. II (most of the microlichens, balance of the macrolichens, and the lichenicolous fungi). Tempe, Lichens Unlimited, Arizona State University. Pp. 617–625.

Edgar, E.; Connor, H.E. 2000: Flora of New Zealand. Volume V Gramineae. Lincoln, Manaaki Whenua Press.

Elix, J.A. 1996: Biochemistry and secondary metabolites. In : Nash III, T.H. (Ed.) Lichen Biology. Cambridge, Cambridge University Press. Pp. 154–180.

Eriksson, O.E. (Ed.) 2005: Outline of Ascomycota – 2005. Myconet 11 : 1–113.

Eriksson, O.E.; Baral, H.-O.; Currah, R.S.; Hansen, K.; Kurtzman, C.P.; Rambold, G.; Laessøe, T. 2004: Outline of Ascomycota – 2004. Myconet 10 : 1–99.

Farr, E.M.; Leussink, J.A.; Stafleu, F.A. 1979a: Index Nominum Genericorum (Plantarum). Regnum Vegetabile 100 : 1–630.

Farr, E.M.; Leussink, J.A.; Stafleu, F.A. 1979b: Index Nominum Genericorum (Plantarum). Regnum Vegetabile 101 : 631–1276.

Farr, E.M.; Leussink, J.A.; Stafleu, F.A. 1979c: Index Nominum Genericorum (Plantarum). Regnum Vegetabile 102 : 1277–1896.

Feuerer, T. (Ed.) 2004: Checklists of lichens and lichenicolous fungi of the world. Version 1. June 2004. [http://www.checklists.de]

Feige, G.B. 1996: Etymologie der wissenschaftlichen Gattungsnamen der Flechten europas. Essen, Botanisches Institut und Botanischer Garten der Universität Essen – Gesamthochschule.

Galloway, D.J. 1985: Flora of New Zealand Lichens. Wellington, Government Printer.

Galloway, D.J. 1993: Global environmental change: Lichens and chemistry. Bibliotheca Lichenologica 53 : 87–95.

Galloway, D.J. 1995: Lichens in Southern Hemisphere temperate rainforests and their role in the maintenance of biodiversity. In : Alsopp, D.; Hawksworth, D.L.; Colwell, R. (Eds) Microbial Diversity and Ecosystem Function. Wallingford, CABI International. Pp. 125–135.

Galloway, D.J. 1996: Lichen biogeography. In : Nash III, T.H. (Ed.) Lichen Biology. Cambridge, Cambridge University Press. Pp. 199–216.

Galloway, D.J. 2004a: New lichen taxa and names in the New Zealand mycobiota. New Zealand Journal of Botany 42 : 105–120.

Galloway, D.J. 2004b: Lichens in our landscape. Te Taiao 3 : 8–9.

Gams, W.; Diederich, P.; Pöldmaa, K. 2004: Fungicolous fungi. In : Mueller, G.M.; Bills, G.F.; Foster, M.S. (Eds) Biodiversity of Fungi. Inventory and Monitoring Methods. San Diego, Elsevier Academic Press. Pp. 343–392.

Gilbert, O. 2000: Lichens. London, Harper Collins [The New Naturalist No. 86].

Godfray, H.C.J.; Knapp, S.J. 2004: Introduction. Taxonomy for the twentyfirst century. Philosophical Transactions of the Royal Society of London, B, 359: 559–569.

Greuter, W.; McNeill, J.; Barrie, F.R.; Burdet, H.M.; Demoulin, V.; Filgueiras, T.S.; Nicolson, D.H.; Silva, P.C.; Skog, J.E.; Trehane, P.; Turland, N.J.; Hawksworth, D.L. (Eds) 2000: International Code of Botanical Nomenclature (Saint Louis Code). Regnum Vegetabile 138. Königstein, Koeltz Scientific Books.

Grgurinovic, C. (Ed.) 1992: Lichens – Introduction, Lecanorales 1. Flora of Australia 54 : i–xviii + 1–349.

Grgurinovic, C. (Ed.) 1994: Lichens – Lecanorales 2, Parmeliaceae. Flora of Australia 55 : i–xviii + 1–360.

Grube, M. 2004: Lichenology since the turn of the millennium. International Lichenological Newsletter 37 (1): 15–22.

Hawksworth, D.L. 1997: Orphans in "Botanical" diversity. Muelleria 10 : 111–123.

Hawksworth, D.L. 1999: Visions of systematic and organismal lichenology in the next century. British Lichen Society Bulletin 84 : 1–4.

Hawksworth. D.L. 2003: The lichenicolous fungi of Great Britain and Ireland: an overview and annotated checklist. Lichenologist 35 : 191–232.

Hawksworth, D.L. 2005: Life-style choices in lichen-forming and lichendwelling fungi. Mycological Research 109 : 135–136.

Huneck, S.; Yoshimura, I. 1996: Identification of Lichen Substances. Berlin, Springer-Verlag.

Johnson, P.N.; Galloway, D.J. 2002: Lichens and their conservation needs in New Zealand. Landcare Research Contract Report LC0102/132, prepared for the Department of Conservation, Wellington. Pp. 1–69.

Johnson, P.N.; Burrows, L.E.; Galloway, D.J. 1998: Air pollution indicators – summary report of main findings. Sustainable Management Fund Project No. 5003. Landcare Research Contract Report LC9899/004, prepared for the Ministry for the Environment, PO Box 10-362, Wellington. Pp. 1–42.

Kantvilas, G.; Jarman, S.J. 1999: Lichens of rainforest in Tasmania and southeastern Australia. Flora of Australia Supplementary Series 9 : 1–212.

Kantvilas, G.; Elix, J.A.; Jarman, S.J. 2002: Tasmanian lichens, identification, distribution and conservation status. I. Parmeliaceae. Flora of Australia Supplementary Series 15 : 1–274.

Kirk, P.M.; Ansell, A.E. 1992: Authors of fungal names. A list of authors of scientific names of fungi, with recommended standard forms of their names, including abbreviations. Index of Fungi Supplement. Wallingford, CAB International.

Kirk, P.M.; Cannon, P.F.; David, J.C.; Stalpers, J.A. 2001: Ainsworth & Bisby's Dictionary of the Fungi, 9th ed. Wallingford, CABI Bioscience, CABI International.

Kranner, I.; Beckett, R.P.; Varma, A.K. (Eds) 2002: Protocols in Lichenology. Culturing, Biochemistry, Ecophysiology and Use in Biomonitoring. Berlin, Springer-Verlag.

Kurokawa S. (Ed.) 2003: Checklist of Japanese Lichens. Tokyo, National Science Museum. 128 p.

Lawrey, J.D.; Diederich, P. 2003: Lichenicolous fungi: interactions, evolution and biodiversity. Bryologist 106 : 80–120.

Lumbsch, H.T. 2002: How objective are genera in euascomycetes? Perspectives in Plant Ecology, Evolution and Systematics 5 : 91–101.

Lumbsch, H.T.; McCarthy, P.M.; Malcolm, W.M. 2001: Key to the genera of Australian lichens. Apothecial crusts. Flora of Australia Supplementary Series 11 : 1–64.

Lutzoni, F.; Pagel, M.; Reeb, V. 2001: Major fungal lineages are derived from lichen symbiotic ancestors. Nature 411 : 937–940.

Lutzoni, F.; Kauff, F.; Cox, C.J.; McLaughlin, D.; Celio, G.; Dentinger, B.; Padamsee, M.; Hibbett, D.; James, T.Y.; Baloch, E.; Grube, M.; Reeb, V.; Hofstetter, V.; Schoch, C.; Arnold, A.E.; Miadlikowska, J.; Spatafora, J.; Johnson, D.; Hambleton, S.; Crockett, M.; Shoemaker, R.; Sung, G.-H.; Lücking, R.; Lumbsch, H.T.; O'Donnell, K.; Binder, M.; Diederich, P.; Ertz, D.; Gueidan, C.; Hansen, K.; Harris, R.C.; Hosaka, K.; Lim, Y.-W.; Matheny, B.; Nishida, H.; Pfister, D.; Rogers, J.; Rossman, A.; Schmitt, I.; Sipman, H.; Stone, J.; Sugiyama, J.; Yahr, R.; Vilgalys, R. 2004: Assembling the fungal tree of life: progress, classification, and evolution of subcellular traits. American Journal of Botany 91 : 1446–1480.

Malcolm, W.M.; Galloway, D.J. 1997: New Zealand Lichens. Checklist, Key, and Glossary. Wellington, Museum of New Zealand Te Papa Tongarewa.

Malcolm, B. [W.M.]; Malcolm, N. 2000: New Zealand Lichens. Nelson, Micro-Optics Press.

Malcolm, B. [W.M.]; Malcolm, N. 2001: New Zealand's leaf-dwelling lichens. Nelson, Micro-optics Press.

McCarthy, P.M. (Ed.) 2001: Lichens 3. Flora of Australia 58A : i–xviii + 1–242.

McCarthy, P.M. 2003: Catalogue of Australian lichens. Flora of Australia Supplementary Series 19 : 1–237.

McCarthy, P.M. 2005: Checklist of the lichens of Australia and its island territories. http://www.anbg.gov.au/abrs/lichenlist/introduction.html

McCarthy, P.M.; Malcolm, W.M. 2004: Key to the genera of Australian macrolichens. Flora of Australia Supplementary Series 23 : i–viii + 1–63.

Nimis, P.L.; Martellos, S. 2003: A second checklist of the lichens of Italy with a thesaurus of synonyms. Museo Regionale di Scienze Naturali Saint-Pierre, Valle d-Aosta, Monografie 4 : 1–191.

Orange, A.; James, P.W.; White, F.J. 2001: Microchemical Methods for the Identification of Lichens. London, British Lichen Society.

Pennycook, S.R.; Galloway, D.J. 2004: Checklist of New Zealand "Fungi". In : McKenzie, E.H.C. (Ed.) Introduction to Fungi of New Zealand. Fungi of New Zealand/Ngā Harore o Aotearoa Vol. 1. Fungal Diversity Research Series 14 : 401–488.

Perry, N.B.; Benn, M.H.; Brennan, N.J.; Burgess, E.J.; Ellis, G.; Galloway, D.J.; Lorimer, S.D.; Tangney, R.S. 1999: Antimicrobial, antiviral and cytotoxic activity of New Zealand lichens 1. Lichenologist 31 : 627–636.

Purvis, O.W.; Coppins, B.J.; Hawksworth, D.L.; James, P.W.; Moore, D.M. (Eds) 1992: The Lichen Flora of Great Britain and Ireland. London, Natural History Museum Publications.

Quilhot, W.; Fernández, E.; Hidalgo, M.E. 1994: Photoprotection mechanisms in lichens against UV radiation. British Lichen Society Bulletin 75 : 1–5.

Quilhot, W.; Fernández, E.; Rubio, C.; Godard, M.; Hidalgo, M.E. 1998: Lichen secondary products and their importance in environmental studies. In: Marcelli, M.P.; Seaward, M.R.D. (Eds) Lichenology in Latin America: History, Current Knowledge and Applications. São Paulo, CETESB. Pp. 171–179.

Rambold, G. (Ed.) 2004: LIAS – a global information system for lichenized and non-lichenized Ascomycetes. A multi-authored distributed Internet project containing information about phylogeny and biodiversity. [http:// checklists.lias.net]

Rambold, G.; Triebel, D. 1999: Generic concepts in lichenized and lichenicolous ascomycetes since 1950 – a historical approach. Symbolae Botanicae Upsalienses 32 (2): 123–164.

Raven, P.H. 2004: Taxonomy: where are we now? Philosophical Transactions of the Royal Society of London, B, 359: 729–730.

Sanders, W.B. 2001: Lichens: the interface between mycology and plant morphology. Bioscience 51 : 1025–1035.

Santesson, R.; Moberg, R.; Nordin, A.; Tønsberg, T.; Vitikainen, O. 2004: Lichen-forming and Lichenicolous Fungi of Fennoscandia. Uppsala, Museum of Evolution.

Scheidegger, C.; Wolseley, P.A.; Thor, G. (Eds) 1995: Conservation biology of lichenised fungi. Mitteilungen der Eidgenössichen Forschungsanstalt für Wald, Schnee und Landschaft 70 (1): 1–173.

Seaward, M.R.D. 2004: The use of lichens for environmental impact assessment. Symbiosis 37 : 293–305.

Sipman, H.J.M.; Aptroot, A. 2001: Where are the missing lichens? Mycological Research 105 : 1433–1439.

Smith, D.C. 1997: What future for lichenology? British Lichen Society Bulletin 80 : 1–5.

Taylor, M. 2002: Meanings and origins of botanical names of New Zealand plants. Auckland Botanical Society Bulletin 26 : 1–201.

Wagenitz, G. 1996: Wörterbuch der Botanik. Morphologie, Anatomie, Taxonomie, Evolution. Die Termini in ihhrem historischen Zusammenhang. Jena, Stuttgart, Lübeck, Ulm, Gustav Fischer.

Wedin, M.; Döring, H.; Gilenstam, G. 2004: Saprotrophy and lichenization as options for the same fungal species on different substrata: environmental plasticity and fungal lifestyles in the Stictis-Conotrema complex. New Phyologist 164 : 459–465.

Wedin, M.; Döring, H.; Könberg, K.; Gilenstam, G. 2005a: Generic delimitations in the family Stictidaceae (Ostropales, Ascomycota): the Stictis - Conotrema problem. Lichenologist 37 : 67–75.

Wedin, M.; Wiklund, E.; Crewe, A.; Döring, H.; Ekman, S.; Nyberg, Å.; Schmitt, I.; Lumbsch, H.T. 2005b: Phylogenetic relationships of Lecanoromycetes (Ascomycota) as revealed by analysis of mtSSU and nLSU rDNA sequence data. Mycological Research 109 : 159–172.

Wilkinson, F.A.; Foster, M.S. 2004: Appendix III. Institutions with significant collections of fungi or fungal allies and fungus-related websites. In : Mueller, G.M.; Bills, G.F.; Foster, M.S. (Eds) Biodiversity of Fungi. Inventory and Monitoring Methods. San Diego, Elsevier Academic Press. Pp. 619–626.

Will-Wolf, S.; Hawksworth, D.L.; McCune, B.; Rosentreter, R.; Sipman, H.J.M. 2004: Lichenized fungi. In : Mueller, G.M.; Foster, M.S.; Bills, G.F. (Eds) Biodiversity of Fungi. Inventory and Monitoring Methods. San Diego, Elsevier Academic Press. Pp. 173–195.