Australasian Lichenology

Australasian Lichenology 

Number 53, July 2003 

5mm _______ 

The New Zealand crustose endemic Placopsis salaz ina in Southland. The species 

is saxicolous and has a determinate thallus which lacks isidia, soredia, or a prothallus, 

and contains salazinic acid. 

CONTENTS 

ARTICLES 

Archer, AW-Graphina hartmanniana Mull. Arg., an additional synonym for 

Dictyographa cinerea (C. Knight & Mitt.) Mull. Arg...................................... 3 

Elvebakk, A; Galloway, DJ-Notes on the heterogeneous genus Psoroma s. lat. 

in New Zealand .... .. .............. ......... .............................. ... ............. .... ............ ... ... 4 

Elix, JA; Jayanthi, VK-5-Methoxylecanoric acid, a new depside from Melanelia 

glabratula.................................. ... .. .................... .. ... ....... .............................. ... 10 

Elix, JA-New species and new records ofNeofuscelia (Parmeliaceae, lichenized 

Ascomycota) from Australia ... ... .. ....................... .... ... ....... .. ............. .. .... .. .... ... 14 

RECENT LITERATURE ON AUSTRALASIAN LICHENS 19 

ADDITIONAL LICHEN RECORDS FROM NEW ZEALAND 

Galloway, DJ (40)-Buellia aethalea (Ach.) Th. Fr., Catillaria contristans (Ny!.) 

Zahlbr., Frutidella caesioatra (Schaer') Kalb, Placynthium rosulans (Th. Fr.) 

Zahlbr. and Pseudocyphellaria mallota (Tuck.) H. Magn............................. 20 

I 

Australasian Lichenology 

Number 53, July 2003 ISSN 1328-4401

Graphina hartmanniana MillI. Arg., an additional synonym 

for Dictyographa cinerea (C. Knight & Mitt.) Miill. Arg. 

Alan W. Archer 

National Herbarium ofNew South Wales 

Mrs Macquaries Road, Sydney 2000, N.S.W., Australia 

The genus Dictyographa (Muller 1893) is characterized by sessile, simple, black 

lirellae, hyaline, muriform ascospores, and anastomosing paraphyses. Dictyo· 

grapha cinerea (C. Knight) Mull. Arg., described from New Zealand (Muller 1894), 

was recently reported from Australia (Archer 2000). A re-examination of a closeup 

photograph ofthe holotype ofGraph ina hartmanniana Mull. Arg. found it to be 

identical to a similar photograph ofD. cinerea. This superficial identity was confirmed 

by reference to Muller's original description of G. hartmanniana (Muller 

1882), and a later examination of the holotype ofG. hartmanniana (Archer 1999). 

In particular, both species have conspicuous, sessile, simple black lirellae with 

completely carbonized proper exciples, they lack lichen compounds, and their ascospores 

are similar in size, viz.: D. cinerea 24-28 x 10-12 11m, 6-7 x 2-4-locular; 

G. hartmanniana: 19-25 x 6-8(-11) Ilffi, 6-8 x 2-locular. 

In the protologue to G. hartmanniana, Muller commented that the species resembled 

no other known species. The inconspicuous nature of D. cinerea could 

explain why no other specimens of"'G. hartmanniana" have been collected since 

Millier's original publication. The species is now known from Queensland and New 

South Wales as well as New Zealand. 

Dictyographa cinerea (C. Knight & Mitt.) Mull. Arg., Bull. Herb. Boissier 2 (Appendix 

1), 78 (1894). 

=Opegrapha cinerea C. Knight & Mitt., Trans. Linn. &x:. London 23, 101 (1860). 

Type. New Zealand.Auckland, on trees, C. Knight. Lectotype: BM (Hayward 1977: 

576). 

=Graphina hartmanniana Mull. Arg., Flora 65, 503 (1882). 

Type. Australia. Queensland, 'Ibowoomba, C. Hartmann; holotype: G, isotype: MEL 

515669. 

References 

Archer, AW (1999): The lichen genera Graphis and Graphina (Graphidaceae) in 

Australia 1. Species based on Australian type specimens. Thlopea 8, 273-295. 

Archer, AW (2000): Additional lichen records from Australia 44. Dictyographa cin· 

erea (C. Knight & Mitt.) Milli. Arg.Australasian Lichenology 47,32. 

Hayward, GC (1977): Taxonomy ofthe lichen families Graphidaceae and Opegraphaceae 

in New Zealand. New Zealand Journal ofBotany 15, 565-584. 

Muller, J (1892): Lichenologische Beitriige XVI. Flora 65, 499-505. 

Muller, J (1893): Lichenes Arabici. Bulletin de l'Herbier Boissier 1. 130-131. 

Muller, J (1894): Conspectus systematicus Iichenum Novae Zelandiae. Bulletin de 

l'Herbier Boissier 2 (Appendix 1), 1-114. 

CORRIGENDA 

The specimen of Sclerophyton rostratum cited on page 19 ofArcher & Elix's paper 

in Volume 52 should read JAElix 22607, not 22606. 

The specimen of Dictyographa cinerea cited on page 32 ofArcher's paper in Volume 

47 should read A. WArcher G459, not 0498. 

AUSTRALASIAN LICHENOLOGY 63, July 2003 (3)

Notes on the heterogeneous genus Psoroma s. lat. in New Zealand 

Arve Elvebakk 

Department of Biology, University of Tromso, N-9037 Tromso, Norway 

e-mail: arve.elvebak.k@i.b.uit.no 

David J. Galloway 

Landcare Research, Private Bag 1930, Dunedin, New Zealand 

e-mail: gallowayd@LandcareResearch.co.nz 

Abstract: Recent changes in the circumscriptions of the genera Pannaria and 

Psoroma have led to the recognition ofan increasing number of"green" species of 

Pannaria formerly included in Psaroma. Comments are made on current knowledge 

of the heterogeneous genus Psoroma, with special reference to New Zealand 

species. Five new combinations in Pannaria are here proposed, viz. Pannaria 

allorhiza (Ny!.) Elvebakk & D.J. Galloway, Panna ria athroophylla (Stirt.) Elvebakk 

& D.J. Galloway. Pannaria durietzii (P. James & Henssen) Elvebakk & D.J. Galloway. 

Pannaria euphylla (Ny!.) Elvebakk & D.J. Galloway and Pannaria patagonica 

(Malme) Elvebakk & D.J. Galloway. 

Introduction 

In the major temperate biomes ofthe Southern Hemisphere, viz. southern South 

America, south-eastAustralia, Tasmania and especially New Zealand, the family 

Pannariaceae is both highly speciose (often at a local as well as a regional level), 

and of considerable biomass, in forest, scrub, and grassland landscapes, with the 

following genera being represented there: DegeliaArv. & D.J. Galloway, Erioderma 

Detailed studies of Pannaria s. lat. and Psoroma s. lat. in the Southern Hemisphere 

are currently in progress and will be reported in detail elsewhere (Elvebakk 

in prep.). However, as part ofrevisionary work for the forthcoming second edition 

ofFlora ofNew Zealand Lichens (Galloway in prep.), we here propose transferring 

five additional species of Psoroma to Pannaria, in line with present thinking. 

Psoroma s. str., as pointed out by J f/Jrgensen & Wedin (1999) and Jf/Jrgensen (2oo0a, 

2002a) refers to the Psoroma hypnorum group oftaxa, but there are at least two 

other groups of species within Psoroma that will eventually need formal recognition 

(Elvebakk, unpublished observations). As a first step towards clarifYing the 

limits of Psoroma s. lat., we here transfer five species currently in Psoroma to 

Panna ria, although Pannaria too might in the future also prove to be heterogeneous. 

Pannaria allorhiza (Nyl.) Elvebakk & D.J. Galloway, comb. nov. 

E Lecanora allorhiza Nyl., Flora 51: 373 (1868). 

ill Psoroma allorhizum (Nyl.) Hue, Nouv. Archs. Mus. Hist. Nat. Paris, ser. 3, 3: 45 

(1891). 

Type: New Zealand. Sine loco (probably Wellingtonl, Charles Knight s. n.; lectotype: 

H-NYL 30795 (fide Galloway (1985: (68)1; i80lectotypes: BM, WELT. 

=Ph;yscia regalis Zahlbr., Denkschr. Akad. Wiss. Wten Math.-Naturwiss. Kl. 104: 

379 (1941). 

Type: New Zealand. Auckland, Rangitoto Island, on Metrosideros tomentosa in 

light forest, H.H. Allan AIDS; holotype: W; isotype: CHR 379831. 

ILLUSTRATIONS: Malcolm & Galloway (1997: 107, 135 - as Psoroma allorhizum); 

Fee, Fuscoderma (D.J. Galloway & P.M. Jf/Jrg.) P.M. Jf/Jrg. & D.J. Galloway, Fusca Malcolm & Malcolm (2000: 40, 106 - as Psoroma allorhizum). 

pannaria P.M. Jf/Jrg., Leiodenna Nyl., Pannaria Delise ex Bory, Parmeliella Mull. 

Arg., Psoroma Ach. ex Michx., Psoromidium Stirt., Santessoniella Henssen and 

DESCRIPTION: Galloway (1985: 467-468 - as Psoroma allorhizum) 

Siphulastrum MUll. Arg. 

Chemistry: Several chemodemes reported, most commonly with vicanicin and oc 

Until relatively recently, the accepted circumscriptions ofthe genera Pannaria 

casionally with additional allorhizin (Ellx et al. 1982: 2328). 

and Psaroma in the Southern Hemisphere were those of Galloway (1985) and 

Jf/Jrgensen & Galloway (1992). Subsequent studies on the Pannariaceae in both 

DISTRIBUTION: New Zealand. Northland (Herekino, Waipoua, Omanaia, Thta· 

Northern and Southern Hemispheres have led to changes in these earlier views, 

moe, Hen &: Chickens Islands, Little Barrier Island, Great Barrier Island, Rakitu 

although the process of generic delimitation in the Pannariaceae is still far from 

Island. Whangarei, Tokatoka, Mahurangi River, Waiheke Island), Auckland 

settled (Ekman & Jf/Jrgensen 2002). 

(Anawahata, Waitakere Range, Rangitoto), South Auckland (Great Mercury Is 

Jf/Jrgensen (1994) defined Pannaria more narrowly as a mainly foliose and preland, 

Mt Maugatawhiri, Coromandel Peninsula, Kaimm Range, Slipper Island), 

dominantly tropical or warm-temperate genus, having pannarin and related com 

Wellington (Kapiti Island), Nelson (Mt Robert, Te Rata, S of Karamea, Maruia), 

pounds as majorsecondary metabolites, apothecia with thalline margins, asci with· 

Marlborough (D'Urville Island, Chetwode Islands, Resolution Bay, Queen Char 

out amyloid apical structures, and partly amyloid hymenia. He segregated from 

lotte Sound). 

Pannaria s. str. squamulose, cool-temperate taxa, usually having fatty acids and 

terpenoids as major secondary metabolites, and with variously marginate apoth 

NOTES: Pannaria allorhiza is characterized by the relatively long, parallel lobes, 

ecia, hemiamyloid hymenia and asci with amyloid apical structures, and placed 

with the central parts ofthe thallus having short, swollen, glomerulate isidia. It is 

these in a new genus Fuscopannaria (Jf/Jrgensen 1994, 2000a, 2000b, 2002b; 

a large species, endemic to northern New Zealand. 

Jf/Jrgensen & Zhurbenko 2002). Later, he suggested that the large, leafY, subtropical 

species of Psoroma should be regarded as "green" species of Pannaria, and 

Pannaria athroophylla (Stirt.) Elvebakk &: D.J. Galloway, comb. nov. 

assigned the Psoroma sphinctrinum group to Pannaria (Jf/Jrgensen 2001). Psoroma 

E Psoroma athroophyllum Stirt., Rep. 7rans. Glasgow Soc. Fld Nat. 1: 21 (1873). 

s. str., with P. hypnorum as generitype, refers to small-squamulose, terricolous, 

E Psoroma subpruinosum var. athroophyllum (Stirt.) C. Knight, 7rans. New Zealand 

muscicolous or saxicolous taxa, without any demonstrable secondary chemistry 

Inst. 7: 365 (1875). 

(with the exception ofP. buchananii and P. fruticulasum, always brownish through 

• Phloeopannaria athroophylla (Stirt.) Zahlbr., Denkschr. Akad. Wiss. Wten Math.deposition 

ofmelanins in the upper cortex), and with an apical amyloid ring struc 

Naturwiss. Kl. 104: 276 (1941). 

ture in the ascus apex (Jf/Jrgensen & Wedin 1994: 3(1). Until very recently, Psoroma 

Type: New Zealand. On bark oftrees, Tinakori Hills, Wellington, J. Buchanan 45; 

was interpreted much more broadly than this (see references above), and it is now 

lectotype: GLAM; isolectotype: WELT. 

recognized that Psoroma s. lat. comprises several discordant elements. 

o 

CD 

AUSTRALASIAN LICHENOLOGY 53, July 2003 

AUSTRALASIAN LICHENOLOGY 53, July 2003

colour when wet. It was reported to be quite common in New Zealand (Galloway 

1985: 479); however, current studies indicate that these records may refer to at 

least one undescribed species with deviating chemistry, but as yet only part of the 

material available for study has been analysed. Until this question is resolved, 

Pannaria patagonica is best considered a doubtful species in New Zealand. 

Acknowledgements 

We would both like to thank our friend Prof. Per Magnus J_rgensen (University of 

Bergen) for valuable discussions on generic delimitation in the Pannariaceae. The 

second author is grateful to Sue Gibb (Landcare Research, Lincoln) and to Mei 

Nee Lee (Auckland Institute and Museum) for information on collections ofPsor· 

oma I Pannaria in CHR and AI{ respectively. Funds to the second author were 

provided by the Foundation for Research Science and Thchnology (FRST Wellington, 

New Zealand) under Contract C09618. 

References 

Calvelo, S; Liberatore, S (2001): Checklist ofArgentinian lichens (Version 2). http:! 

lwww.biologie.uni-hamburg.delchecklistslargen_12.htm 

Ekman, S; Jlfrgensen, PM (2002): 1bwards a molecular phylogeny for the lichen 

family Pannariaceae (Lecanorales, Ascomycota). Canadian Journal ofBotany 

80,625-634. 

Elix, JA; Lajide, L; Galloway, DJ(1982): Metabolites from the lichen genusPsoroma. 

Australian Journal ofChemistry 35, 2325-2333. 

Galloway, DJ (1985): Flora ofNew Zealand Lichens. i-Ixxiii + 1-662 pp. P.D. Hasselberg, 

New Zealand Government Printer, Wellington. 

Galloway, DJ; QuiIhot, W (1999) ("1998"): Checklist ofChilean lichen-forming and 

lichenicolous fungi. Gayana (Botanica) 55, 111-185. 

James, PW; Henssen, A (1975): A new sorediate species ofPsoromo. with sorediate 

cephalodia. Lichenologist 7, 143-147. I 

Jlfrgensen, PM; Galloway, DJ (1992): Pannariaceae. Flora ofAustralia 54,246-293. 

Jlfrgensen, PM (1994): Studies in the lichen family Pannariaceae VI: The taxonomy 

and phytogeography of Pannaria Del. s. lat. Journal ofthe Hattori Bot· 

anical Laboratory 76, 197-206. 

J_rgensen, PM; Wedin, M (1999): On Psoroma, species from the Southern Hemisphere 

with cephalodia producing vegetative dispersal units. Lichenologist 31, 

341-347. 

J_rgensen, PM (2000a): Survey ofthe lichen family Pannariaceae on the American 

continent, north of Mexico. Bryologist 103, 670-704. 

Jlfrgensen, PM (2000b): Notes on some East-Asian species of the lichen genus 

Fuscopannaria. Journal ofthe Hattori Botanical Laboratory 89, 247-259. 

J_rgensen, PM (2001): New species and records ofthe lichen family Pannariaceae 

from Australia. Bibliotheca Lichenologica 78,109-139. 

Jlfrgensen, PM (2002a): Psoromo.. In: Nash III, TH; Ryan, BD; Gries, C; Bungartz, 

F (Eds) Lichen Flora ofthe Greater Sonoran Desert Region. Vol. I (the pyre nolichens 

and most of the squamulose and macrolichens). pp. 431-433. Lichens 

Unlimited, Arizona State University: 'Thmpe, Arizona. 

J,rgensen, PM (2002b): Further notes on Asian species ofthe lichen genus Fusco· 

pannaria. Journal ofthe Hattori Botanical Laboratory 92,225-229. 

J_rgensen, PM; Zhurbenko, M (2002): Two new, remarkable, arctic species in the 

lichen genus Fuscopannaria (Pannariaceae, Iichenized Ascomycetes). Bryologist 

105, 465-469. 

Kantvilas, J; James, PW; Jarman, SJ (1985): Macrolichens in Tasmanian rainforests. 

Lichenolol1ist 17, 67-83. 

Kantvilas, 

o 

G; James, PW (1987): The macrolichens of Tasmanian rainforest: key 

and notes. Lichenologist 19, 1-28. 

AUSTRALASIAN LICHENOLOGY ISS, July 2003 

Kantvilas, G (1989): A checklist ofTasmanian lichens. Papers and Proceedings of 

the Royal Society ofTasmania 123, 67-85. 

Kantvilas, G (1994): A revised checklistofthe Tasmanian lichen flora. Muelkria 8, 

155-175. 

Kantvilas, G; Jarman, SJ (1999): Lichens of rainforest in Tasmania and southeastern 

Australia. Flora ofAustralia SuppkTMntary Series 9, i-xi + 1-212. 

Malcolm, WM; Galloway, DJ (1997): New Zealand Lichens. Checklist, Key, and 

Glossary. i-vi + 192 pp. Museum ofNew Zealand 'Ie Papa 1bngarewa, Wellington. 

Malcolm, B (W.M.J; Malcolm, N (2000): New Zealand Lichens. i-ii + 134 pp. Micro 

Optics Press, Nelson. 

Malcolm, B [w.M.); Malcolm, N (2001): New Zealand's Leaf-Dwelling Lichens. i-vi 

+ 73 pp. Micro-Optics Press, Nelson. 

Malme, GOA (1925): Die Pannariazeen des Regnellschen Herbars.Arkiu fOr Bota.nik 

20A(6), 1-23. 

McCarthy, PM (2003): Catalogue ofAustralian lichens. FWra ofAustralia SuppleTMntary 

Series 19, 1-237. 

Purvis, OW (2000): Lichens. 1-112 pp. The Natural History Museum, London. 

Quilhot, W; Piovano, M; Arancibia, H; Garbarino, JA; Gambaro, V (1989): Studies 

on Chilean lichens, XII. Chemotaxonomyofthe genus Psoromo.. Journal ofNatural 

Products 52,191-192. 

Vezda, A(1997): Lichenes Rariores Exsiccati. Fasciculus undetricesimus (numeris 

281-290). 4 pp. Brno. 

AUSTRALASIAN LICHENOLOGY 53, July 2003 o

subsorediza, Brigantiaea fuscolutea, *Buellia adjuncta, B. aethalea, Caloplaca 

ammiospila, C. biatorina, C. caesiorufella, C. cerina, C. chrysodeta, C. 

chrysophthalma, C. concilians, C. crenulatella, C. rubelliana, C. saxicola, C. 

torwensis, C. xantholyta, Candelariella subde{lexa, Carbonea vitellinaria, C. 

vorticosa, Catapyrenium cinereum, C. daedalum, Catillaria contristans, 

*Cercidospora trypetheliza, Cetrariella delisei, Cladonia earneola, C. ecm.ocyna, C. 

gracilis subsp. vulnerata, C. sulphurina, C. uncialis, Clauzadea monticola, 

Clauzadeana macula, *Clypeococcum grossum, *Dactylospora acarosporae, *D. 

australis, *D. frigid4, *D. parasitica, Dermatocarpon luridum, Epigloea soleiformis, 

Frutidella caesioatra,lcmadophila ericetorum,lmmersaria athroocarpa, Lecanora 

bicincta, L. cavicola, L. intricata, L. swartzii, Lecidea diducens, L. lapicida subsp. 

lapicida, L. lapicida subsp. pantherina, L, swarlioidea, *L. verruca, Lecidella 

wulfenii, Lecido17UJ, demissum, Lepraria eburnea, L. membranacea, L. neglecta, L. 

vouauxii, Leptogium plicatile, *Lichenochora xanthoriae, Massalongia carnosa, 

Megaspora verrucosa, Miriquidica deusta, M. nigroleprosa, *Muellerella pygmaea, 

Mycobilimbia hypnorum, Myxobilimbia labulata, Ochrolechia xanthosto17UJ" 

Neofuscelia subhosseana, Pannaria hookeri, Peltigera lepidophora, P. malacea, P. 

neckeri, P. neopolydactyla, Perlusaria doctylina, Physcia semipinnata, Plac:ynthieUa 

oligotropha, Placynthium rosulans, *Polycoccum pulvinatum, *P. squa17UJ,rioides, 

Porpidia platycarpoides, P. superba, Pseudephebe minuscula' P. pubescens, Psoroma 

hypnorum, Racodium rupestre, Rhizocarpon copelandii, R. disporum, R. 

eupetraeum, R. geminatum, R. geographicum subsp. arcticum, R. grande, R. 

hochstetteri, R.lavatum, R.lecanorinum, R. polycarpum, *R. pusillum, R. reductum, 

R. submodestum, R. subpostumum, *Rimularia insularis, R. psephota, *Rinodina 

insularis, R. olivaceobrunnea, R. roscid4, Schaereria fabispora, S. fuscocinerea, 

Sclerophora a17UJ,bilis, Solorina crocea, S. spongiosa, Sporastatia testudinea, 

Staurothele {issa, Tetramelas papillata, *Thamnogalla crombei, Thelomma 

ocellatum, Thrombium epigaeum, Trapeliopsis pseudo granulosa, Tuckermannopsis 

chlorophylla, Umbilicaria grisea, U. krascheninnikovii, U. nylanderiana, U. 

subglabra, U. umbilicarioides, U. virginis, Wrrucaria aquatilis, V. ceuthocarpa, V. 

morgacea, V. mucosa, V. rheitrophila, V. striatula, Xanthoria elegans and x 

polycarpa. 

How are these bipolar lichen distributions explained? This was discussed in a 

preliminary way by Galloway & Aptroot (1995), and at present biogeographers 

suggest two major frameworks: (1) VlCariance models. Allopatric speciation resulting 

from some kind ofgeographical barrier separating a formerly continuous population. 

Rafting of fragments from earlier landmasses such as Gondwana, and/or 

the accretion of terranes ofexotic origin. There are several competing scenarios, 

but as yet none has been rigorously tested, partly because alpine lichens of the 

Southern Hemisphere and oftropical mountains are still not adequately and accurately 

known, nor is the geology ofcomposite present-day landmasses known with 

certainty. (2) Long-distance dispersal. This is now much more fashionable in discussions 

ofplant distributions in the Southern Hemisphere than it was a decade 

or so ago (see for example Pole 2001). The alpine vegetation for the South Island of 

New Zealand, although physically isolated in the southern ocean for some SOMY 

and with a high degree ofendemism, is nevertheless comparatively young, adaptive 

radiation having occurred after long-distance dispersal during the late Miocene 

to early Pleistocene (McGlone et al. 2001, Lee et al. 2001). Unlike the flowering 

plants of alpine New Zealand, the alpine lichens have apparently not speciated 

in situ at all, with only a few endemic taxa developed such as Labyrintha 

(Malcolm et al. 1995). Some undoubtedly have arrived in relatively recent times in 

the West Wind Drift, but others could well be truly Gondwanan or even earlier 

relicts. 

@ 

AUSTRALASIAN LlCHENOLOGY 63, July 2003 

The persistence oflichen communities in extreme, high-alpine environments at 

high latitudes in both hemispheres is a well-recognized phenomenon (Galloway & 

Aptroot, 1995, Adler & Calvelo 2002), but still without a single unifying explanation. 

Recent developments in phylogenetic systematics of fungi have shown that 

the lichenized state is indeed a very ancient symbiosis stretching back to the very 

beginnings ofthe colonization ofthe earth by living systems, and that lichen associations 

are the likely ancestors ofCree-living Ascomycete fungaIlines (Lutzoni et 

al. 2001). There is evidence that gains oflichenization often are followed by losses. 

It is axiomatic that earth and life evolved together, so in seeking an explanation 

for the phenomenon ofbipolarity in lichens, increasingly more sophisticated (and 

hopefully cheaper) molecular methods, plus advances in knowledge of composite 

geological areas, will be powerful tools to help provide some realistic answers. 

Acknowledgements 

I am grateful to my friend Prof. Per Magnus JfJrgensen (University of Bergen) 

for assistance with identification of Plac;ynthium rosuians, and to Ruth Lewis 

(Landcare Research Library, Lincoln), Dr Robin Craw (Otago Museum), Dr Peter 

Johnston (Landcare Research, Mt Albert, Auckland) and Priv. Doc. Dr Klaus Kalb 

(Neumarkt) for help with literature. Funds for this research were provided by the 

Marsden Fund administered br the Royal Society ofNew Zealand under Contract 

UOOS05, and by the Foundatlon for Research, Science and Technology (FRST, 

Wellington) under Contract C0961S. 

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