Preface
PREFACE
New Zealand has a richly diverse and well-developed lichen flora, arguably one of the most interesting and best preserved in the world today. Many species are of great size and beauty, and in wetter, forested areas, are often dominant components of the epiphytic and ground vegetation. For all that, New Zealand's lichens have long been the "Cinderella" group of its flora, disregarded by professional botanists and, except in a few cases, interested amateurs alike. However, over the past 10-15 years there has been an awakening of interest in New Zealand's lichens and in anticipation of future research into lichen ecology, ecophysiology, sociology, pollution studies and biogeography, the present lichen flora is offered as a working introduction to a complex and still poorly known plant group. The luxuriance of New Zealand's lichen vegetation was early remarked upon by visiting botanists, and, with few exceptions, that impression still obtains today. In view of the considerable extinction of lichens through atmospheric and terrestrial pollution in Britain, Europe and parts of North America, New Zealand's astonishing lichen flora must gain a wider interest and importance in the international lichenological community, containing as it does centres of speciation in such genera as: Megalospora, Menegazzia, Neuropogon, Placopsis, Pseudocyphellaria, Psoroma, Sphaerophorus and Sticta, genera which are often represented in the Northern Hemisphere by only one or a few species.
Churchill Babington wrote in his introduction to the account of lichens in Hooker's Flora Novae Zelandiae "... the lichenological flora of a country requires to be studied by a native botanist, who may recognize a protean species by a knowledge of its living forms, or, as Fries expresses it, of its history. Without this knowledge, especially in the case of crustaceous Lichens, it is oftentimes hopeless to attempt to arrive at a clear result; and I am confident that the more difficult species of the southern hemisphere, even of the higher genera (of Sticta, for example) will never be properly elucidated, till they have been described and limited by some one who is familiar with them in the living state". Babington wrote this in 1855 after having worked on New Zealand lichens for over two years in his rooms in Cambridge, never having seen either the South Pacific or New Zealand.
The Scottish physician W. Lauder Lindsay, on the other hand, spent several months in Otago in 1861 collecting lichens and making notes on the vegetation and geology. In 1867 ( Trans. Roy. Soc. Edin. 24: 407-8 ) he expressed a completely opposite view, "...I hold that he only is fully competent to determine and describe species from new countries, who, in addition to the requisite analytical and descriptive power, has, on the one hand, constant access to, and an intimate knowledge of, the now overwhelming and ever-increasing mass of Botanical Literature in all the principal European languages; and, on the other, equally habitual access to Herbaria which contain the largest collections of specimens from all parts of the world, such, for instance, as those of Kew or Paris. By no other means does it appear possible now-a-days accurately to ascertain or distinguish what is new from what is already known in the plant world. This virtually restricts systematic and descriptive botany to the Naturalists of London or Paris, or of similar centres of botanical knowledge; and as virtually excludes Provincial Botanists, who are isolated from the sources of the necessary fundamental information. It were easy for a collector or observer in a new field to name and describe, what to himself, according to his limited opportunities for judging, appears to be new. But if he do so, however otherwise qualified, without that knowledge, which can, generally speaking, only be acquired in the Botanical Libraries, and from the Herbaria of the largest European cities, he cannot fail to add to the confusion of synonyms, and impede the true progress of botanical discovery and science, by publishing as new, and under new names, species, which a wider experience speedily proves to be identical with, or mere forms of, other plants already known as natives of other parts of the world. Collectors are, as a body, and as a rule, naturally desirous of naming and describing their own collections; and in certain respects, no other Botanists can be so well equipped to do so. Nor is it always possible to secure the co-operation of those overworked, eminent authorities, who have the largest knowledge of the special departments of botanical science which they respectively cultivate and adorn...".
The basic raw materials of any flora are adequate regional collections (with associated ecological and habitat data), and a detailed knowledge of the taxonomic literature (supported by examination of type specimens) relating to that flora. For a New Zealand lichen flora the first requirement demands local field knowledge, while the second means herbarium and library work in Britain and Europe, 12000 miles distant from where the lichens actually grow. Thus, both Babington's and Lindsay's injunctions apply, if one is to build securely on the work of earlier botanists who studied and described New Zealand lichens. Having the best of both worlds is of course an enviable solution, and to a considerable extent in the 10 years that is has taken to prepare this flora, this I have had. The bulk of my time was spent at the British Museum with visits to other important European herbaria to study New Zealand lichens, interspersed with reasonably frequent visits back to New Zealand for field and herbarium work. My own field studies have alas been relatively superficial, except in a few cases, but it was considered more important to have available quickly, a working flora, warts and all, as a basis for informed future fieldwork, without postponing the work indefinitely until New Zealand's lichens were more completely known.
In the present work 210 lichen genera are discussed, supported by descriptions of 966 taxa. This figure represents possibly about 60% of the lichens to be found in New Zealand and it is expected that many major discoveries remain to be made, especially in the numerous microlichen genera.
This lichen flora would not have been completed within 10 years were it not for the great help and encouragement given to me by very many people who are mentioned in the Acknowledgments (see below).
Generic affinities when known with certainty are indicated in the appropriate generic discussions. A formal suprageneric classification is not adopted in this account as relationships, and even exact limits, of many lichen genera remain unresolved and controversial. Until a satisfactory integration of lichens into the general system of fungal classification is achieved, it has been thought better to avoid adherence to a particular arrangement such as those suggested by Henssen and Jahns ["Lichenes. Eine Einführung in die Flechtenkunde". Georg Thieme: Stuttgart. Pp. 252-418 (1973)], Poelt [In Ahmadjian, V. and Hale, M.E. (Eds) "The Lichens", Academic Press: London and New York. Pp. 599-632 (1974)], Barr [ Mem. N.Y. Bot. Gard. 28: 1-8 (1976); Mycologia 75: 1-13 (1983)] or Eriksson [ Opera Bot. 60: 1-220 (1981); Mycotaxon 15: 203-248 (1982)]. Since the lichen floras of many tropical and southern temperate regions are still only incompletely known, lichen taxonomy in these regions will of necessity for some time yet concentrate on the delimitation of species and genera.
The descriptive part of the flora in which genera are arranged alphabetically incorporates the following: A genus description in which essential characters are outlined accompanied by a discussion citing relevant literature. Descriptions of species are arranged alphabetically following a key, and are based on personal examination of New Zealand material. A standard arrangement of species description is adopted and includes thalline characters, apothecial characters (including dimensions of ascospores and in some cases a more detailed account of apothecial anatomy), and chemical characters (when known). For general accounts of lichen anatomy and morphology see e.g., Henssen and Jahns ( loc. cit. ), Hale ["The Biology of lichens", 2nd ed. Edward Arnold: London (1974)] and Jahns (In Ahmadjian, V. and Hale, M.E. (Eds) "The Lichens". loc. cit., pp. 3-58].
The use of chemotaxonomy in lichenology is now universally accepted, see Hale [ loc. cit., (1974) pp. 103-129] and Hawksworth [In Brown et al., (Eds) "Lichenology: Progress and problems", Academic Press: London and New York. Pp. 139-184 (1976)]. In this flora, both thalline colour tests [Hale loc. cit., (1974) pp. 119-121; Walker and James Bull. Br. Lich. Soc. 46 (Suppl.): 13-29 (1980)] and thin-layer chromatography (TLC) of acetone extracts were routinely used.
The standardised methods of Culberson [ J. Chromatogr. 72: 113-125 (1972); Bryologist 84: 16-29 (1981)] were used in TLC analyses. Comprehensive chemical data on lichens is given in Culberson ["Chemical and botanical guide to lichen products", Univ. N. Carolina Press: Chapel Hill (1969); Bryologist 73: 177-377 (1970)] and in Culberson et al., ["Second supplement to chemical and botanical guide to lichen products", American Bryological and Lichenological Society: St Louis (1977)]. In order to avoid long lists of synonyms, full bibliographic citations are given only for currently accepted taxa in the New Zealand lichen flora, for appropriate basionyms and for heterotypic synonyms based on taxa originally described from New Zealand. In most cases homotypic synonyms are given in Zahlbruckner ["Catalogus Lichenum Universalis" Vols 1-10, Bornträger: Leipzig (1921-1940)]; Lamb ["Index Norminum Lichenum inter annos 1932 et 1960 divulgatorum", Ronald Press: New York (1963)]; Hawksworth [ Index of Fungi Supplement. Lichens 1961-1969" (1972)] and, since 1970, in the twice-yearly Index of Fungi. References to relevant taxonomic works giving synonymy in particular genera are cited in the appropriate generic discussions. Since type material of all lichen taxa described from New Zealand was not available for examinations, synonymies are not always complete. Synonyms are printed in italics both in the text and index.
Holotypes and/or lectotypes of names based on material from New Zealand are cited when authentic material seen by the describing author was examined. Where New Zealand lichen taxa are already typified in the literature, the citation of the type is given thus: [fide Martin 1962 p. 19]; in all other cases typifications given are those of the present author. In some cases authentic material of taxa described from New Zealand was not located and such taxa still require typification. Type citations of certain species not originally described from New Zealand, but which are significant in the New Zealand flora are also given when appropriate. Isotype and/or syntype material is not specifically cited except in certain special instances where a holotype or lectotype may be endangered by further examination. For location of herbaria, or specimens of individual collectors in New Zealand see chapter on collectors (p. xxiv).
An indication of the known distibutions of lichen 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 this volume. Terms illustrating altitudinal zonation are those of Wardle [ N.Z.J. Bot. 2: 352-366 (1964)]. Abbreviated habitat notes are given when such information is available.
Biogeographical elements recognised in the New Zealand lichen flora are briefly defined as follows:
Endemic: taxa confined to New Zealand only, and including the subantarctic islands. Only two endemic genera (Calycidium and Thysanophoron) are known at present.
Australasian: taxa occurring in New Zealand and Australia. Three subgroups may be discerned in this element: (1) New Zealand, Tasmania and the subantarctic islands. (2) New Zealand, Australia and the subantarctic islands. (3) The subantarctic islands and Tasmania.
Austral: taxa occurring in New Zealand, the subantarctic islands and southern South America.
Cosmopolitan: taxa occurring on all known landmasses.
Bipolar: taxa occurring in New Zealand (usually in alpine localities) and in boreal localities in the Northern Hemisphere.
Western Pacific: taxa occurring on the Asiatic seaboard, Japan, China, Malesia, New Guinea, New Caledonia, the east coast of Australia and New Zealand.
Pantropical: taxa occurring in all major tropical areas of the world.
Palaeotropical: taxa occurring east of and including Easter I., to St Helena-most of the Pacific islands are included here.
Circum-Pacific: taxa occurring on the seaboards of North and South America, and of Asia and including Australia and New Zealand.
Southern xeric: taxa occurring in South Africa, Western Australia, South Australia, ?Tasmania and New Zealand, usually defined by winter-rain-fall-summer drought conditions. This term, suggested to me by Dr P. Wardle, is an alternative to mediterranean [see Johnson and Briggs Bot. J. Linn. Soc. Lond. 70: 149-153 (1975)].
A preliminary discussion of biogeographical elements in the New Zealand lichen flora with examples is given in Galloway [In Bramwell, D. (Ed.) "Plants and Islands", Academic Press: London and New York. Pp. 201-224 (1979)]. For a more general account of distribution patterns of lichens in the Pacific region see Jørgensen. [ Aust. J. Bot. Suppl. ser. 10: 43-66 (1983)].
For additional general information on lichens and the lichen symbiosis, and for accounts of Australian lichens (useful for comparative purposes), the reader is referred to the following: Richardson, D.H.S., "The vanishing lichens", David and Charles: Newton Abbot (1975); D.H. Brown, D.L. Hawksworth and R.H. Bailey (Eds), "Lichenology: progress and problems", Academic Press: London and New York (1976); M.R.D. Seaward (Ed.), "Lichen ecology", Academic Press: London and New York (1977); R.B. Filson and R.W. Rogers, "Lichens of South Australia", D.J. Woolmann, Government Printer: South Australia (1979); R.W. Rogers, "The genera of Australian lichens (lichenized fungi)", University of Queensland Press: St Lucia (1981); Ahmadjian, Progr. Phycol. Res. 1: 179-233 (1982) and D.L. Hawksworth, B.C. Sutton and G.C. Ainsworth (Eds), "Ainsworth & Bisby's Dictionary of the Fungi", 7th ed., C.M.I.: Kew (1983).
ACKNOWLEDGMENTS
This flora was begun in January 1973, at the instigation of Dr E.J. Godley, former Director of Botany Division, DSIR, and concluded in 1983 under the aegis of his successors Dr H.E. Connor and Dr W. Harris. All three Directors showed much interest in the project and took great pains to facilitate its trouble-free execution; their encouragement and advice have been mainstays of my work. Special thanks are due to Dr Godley and to Dr G.W. Butler (Head Office, DSIR) who successfully negotiated with the State Services Commission for extended periods of study at the British Museum (Natural History), without which the lichen flora could not have been completed within ten years. Mr R. Ross, and Mr J.F.M. Cannon, successive Keepers of Botany at the BM offered me extensive use of herbarium and library facilities between 1973 and 1983 and I am much indebted to them, and to Mr P.W. James, Deputy Keeper of Botany, for their continuing interest in my work.
Colleagues at Botany Division were unfailingly helpful and contributed in many different ways: Dr Elizabeth Edgar oversaw the final editing of the text, and checking of printouts from the word processor, a task in which she was assisted by Miss A. Miles and Mr A. Dawson who read the final proofs. Dr Edgar also made many helpful suggestions on the final form of the flora and assisted with descriptions, validations of new taxa and discussed endless taxonomic and nomenclatural matters in her usual, cheerfully selfless manner - my debt to her is considerable. I am grateful to Dr Lucy B. Moore, Dr P.J. Garnock-Jones, Dr M.J. Parsons and Mr K.R. West for fruitful discussions on taxonomic matters; Dr P. Wardle and Mr W. Sykes for help with biogeographical details; Mrs M.J.A. Bulfin (née Simpson), Mr I. Brown and Dr B.P.J. Molloy for valued field assistance; Mrs M.E. Blackmore and Mr J. Knox and their library staffs for assistance with literature and photocopying; Miss Bryony Macmillan and her preparation room staff for unstinting help with the mailing of herbarium specimens and typescript between Lincoln and London, and also in the abstracting of distributions and of habitat information from specimens in CHR - this generous assistance has played no small part in the successful preparation of the flora. To Miss Audrey White sincere thanks for the efficient and faultless manner with which she dealt with several series of typescript. The cover design was prepared by Mrs Patricia Brooke.
It is a pleasure to acknowledge the cheerful assistance of Mr and Mrs Michael and Diana Young and their staff at Wordcom, Christchurch, who entered the text on disc, and of Mrs Christine McDonald, (and her assistants, Shirley Mendoza and Christine McNeill) Science Information Publishing Centre, DSIR, Wellington, who oversaw the book through all stages of printing, with meticulous care.
I am also grateful to many other New Zealand colleagues and friends. In Dunedin between 1961 and 1969 I received help and encouragement from Prof. G.T.S. Baylis, the late Dr Elizabeth J. Batham, the late Dr J. Murray, Dr G.A.M. Scott, Mr W. Tubman (skipper R.V. Munida) and Prof. A.P. Wylie. In Christchurch Prof. J.D. Lovis, Dr C.J. Burrows and Dr B.A. Fineran (University of Canterbury, Botany Department) provided access to lichen collections housed at CANU, and the members of the Canterbury Botanical Society provided many useful additions to the lichen flora on several interesting excursions. In Wellington I was constantly helped by the botanists at WELT, Miss Nancy Adams, Dr P.J. Brownsey and the late Mr B.W. Hamlin, who sent critical material from the Buchanan, Colenso and Knight herbaria.
I am particularly grateful to the following for their assistance in the field, or for allowing me access to their lichen collections, or for specifically collecting for me in various parts of New Zealand: Dr I.A.E. Atkinson, Mr I.J. Barton, Dr J.E. Braggins, Dr W.T. Bussell, Dr J. Child, Mr P. Child, Mr A.J. Dakin, Mrs Mavis Davidson, Mr A.A. Deans, Dr B.W. and Mrs Glenys Hayward, Dr P.N. Johnson, Dr L.D. Kennedy, Mr T.H. Kennedy, Dr L.F. Molloy, Dr C.D. Meurk, Miss J. Singleton and Mr H.D. Wilson.
Special thanks are due to Mr J.K. Bartlett for furnishing rich collections for study from North Auckland, central and eastern North I., north-west Nelson and Westland. His serious collecting began only in 1976 but since then he has disclosed many new records for New Zealand and discovered several new taxa, and were it not for his diligent endeavours the present lichen flora would have been much poorer in genera than it is now.
During this study it was necessary to examine many early collections (mainly nineteenth century but a few from the eighteenth century), from herbaria in Britain and Europe. Apart from the important collections in BM, specimens were received from the following herbaria whose curators are thanked for their prompt and courteous attention: ABD, BP, CANU, CHR, E, FH, FI, G, GB, GLAM, H, H-NYL, L, LD, LINN, LINN-SM, M, MANCH, MEL, NSW, OTA, PAD, PC-HUE, PC-LENORMAND, PC-MONTAGNE, PC-THURET, PD, S, U, UPS, UPS-THUNBERG, UPSV, US, VER, W, WU, WELT and WRSL.
The Department of Scientific and Industrial Research provided travel monies for me to visit lichen herbaria in Scandinavia [Helsinki, Uppsala (including cataloguing the Du Rietz lichens collected in New Zealand in 1926-27), Stockholm, Lund and Bergen] in 1974 and Melbourne in 1980, and for this assistance I record my gratitude. I have made several private visits to Sweden to study New Zealand lichens held in GB, UPS and S, and my wife's operatic commitments have allowed me to work on lichen collections in Florence, Geneva, Genoa, Padua, Paris and Verona. A Royal Society - Nuffield Foundation, Commonwealth Bursary awarded to me in 1972 allowed me to travel to London to commence work at the BM and I am grateful to the administrators of that fund for their material help.
I gratefully acknowledge the assistance of the following friends and colleagues for specific help on many topics; Prof. T. Ahti (University of Helsinki - Cladonia), Dr O. Almborn (University of Lund - checking bibliographical and biographical data, provision of literature), Dr A. Archer (New South Wales - chemistry of Thysanothecium, Cladonia), Dr L. Arvidsson (University of Göteborg - Coccocarpia, Degelia, assistance with literature), Dr I.M. Brodo (National Museum of Canada, Ottawa - Lecanora); Dr B.J. Coppins (Royal Botanic Gardens, Edinburgh - Bacidia, Cliostomum, Lecideaceae, Micarea); Prof. G. Degelius (Universty of Göteborg - Collema, assistance with literature); Mrs Greta Du Rietz and Mr Rolf Du Rietz (Uppsala - access to the lichen collections, notes and records of Prof. G. Einar Du Rietz, assistance with literature); Dr J.A. Elix (Australian National University, Canberra - Hypogymnia, Parmelia, Psoroma, Xanthoparmelia, assistance with chemistry); Mr R.B. Filson (Australian National Herbarium, Melbourne - Cladia, Australian lichens); Dr A. Fletcher (Leicester - Lichina); Dr J. Hafellner (University of Graz - Brigantiaea); Dr M.E. Hale (Smithsonian Institution, Washington - Parmelia, Parmelina, Parmotrema, Pseudoparmelia, Xanthoparmelia); Dr D.L. Hawksworth (Commonwealth Mycological Institute, Ferry Lane, Kew, Richmond, Surrey - Anisomeridium, Mycomicrothelia, assistance with nomenclatural problems); Prof. A. Henssen (University of Marburg - Lichinaceae, Pannariaceae, Psoroma, Roccellinastrum, Steinera, access to unpublished results); Dr H. Hertel (Botanische Staatssammlung, Munich - Lecidea, Lecidella); Mr P.W. James (British Museum (Natural History) - all groups, assistance with chemistry, editing and checking final MS); Prof. P.M. Jørgensen (University of Bergen - Bryoria, Erioderma, Leioderma, Leptogium, Pannariaceae, advice on nomenclatural problems); Dr H. Krog (University of Oslo - Heterodermia, Parmeliaceae, Punctelia, Phyllopsora, Sticta, access to unpublished results); Mr J.R. Laundon (British Museum (Natural History) - Caloplaca, Chrysothrix, Lecanora, Lepraria, Leproplaca, Xanthoria, assistance with biographical, bibliographical and nomenclatural problems); Dr E. Mackenzie (Cambridge, Mass. - Stereocaulon); Dr H. Mayrhofer (University of Graz - Rinodina); Dr R. Moberg (University of Uppsala - Hyperphyscia, Phaeophyscia, Physcia, Physconia, assistance with literature); Dr A.O. Orchard (University of Hobart - access to G.C. Bratt lichen collection); Prof. J. Poelt (University of Graz - discussion and advice on nomenclatural problems); Dr B. Renner (University of Marburg - Pseudocyphellaria, Psoroma, assistance with chemistry, discussion on photosymbiodemes); Prof. R. Santesson (University of Uppsala, and Botanical Museum, Stockholm - Siphula, Pseudocyphellaria and many other groups, assistance with literature and advice on nomenclatural problems); Mr N. Sammy (University of Western Australia - Xanthoparmelia); Dr M.R.D. Seaward (University of Bradford - advice on historical matters and on 19th century lichen collectors and collections); Dr J.W. Sheard (University of Saskatchewan - Buellia); Prof. H. Sjörs (University of Uppsala - access to the lichen collections of Prof. G.E. Du Rietz); Dr H. Sipman (University of Utrecht - Austroblastenia, Megaloblastenia, Megalospora); Mrs G.N. Stevens (University of Queensland - Ramalina); Dr T.D.V. Swinscow (Topsham, Devon - Coccocarpia, Phyllopsora, Sticta); Dr L. Tibell (University of Uppsala - Caliciales); Dr P.B. Topham (Dundee - Umbilicaria); Mr O. Vitikainen (University of Helsinki - Peltigera, assistance with literature, 19th century archival material and access to Acharius and Nylander collections); Dr A.L. Wilkins (University of Waikato - Nephroma, Pseudocyphellaria chemistry); Miss F.J. Walker (British Museum (Natural History) - Neuropogon). Any errors remaining in the text after this considerable catalogue of assistance, willingly given, must remain my responsibility.
My debt to Peter James (BM) is inestimable - we first worked together in 1962-63 when I was his assistant in the curating of James Murray's lichen collection (OTA). From that time, through a regular exchange of correspondence he actively encouraged my interest in lichens as he has done for so many other beginners in lichenology. Since 1973 he has taken a vital interest in the progress of the present flora and at the start determined how the problems should best be approached. His considerable knowledge of Southern Hemisphere lichens has continuously and unstintingly been placed at my disposal, he patiently undertook many of the chemical analyses recorded herein, and he also prepared the account of Menegazzia. In most other genera he offered advice, criticism and encouragement. Finally he helped to read, check and edit the entire manuscript through its many drafts, so that the final work contains considerably fewer errors of fact and judgement than it otherwise would have. Such a contribution should not go unmarked, for without his help this flora which is, in its way, a testament to his interest in the lichens of New Zealand (fostered through his earlier association with James Murray), would never have been attempted or written.
Lastly, my profound thanks, gratitude and love to my wife Patricia, for her encouragement, optimism, strength and unfailing support. Despite the demands of an international operatic career, she bravely accepted the additional burden of the many months of separation that my botanical work in New Zealand imposed, and still cheerfully urged me on to the flora's successful conclusion.
