Bulletin of the

British Museum (Natural

BRITIS- JM

History)

Studies in Pseudocyphellaria (lichens)
I. The New Zealand species

David J. Galloway

Botany series Vol 17

25 February 1988

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World List abbreviation: Bull. Br. Mus. nat. Hist. (Bot.)

British Museum (Natural History), 1988

The Botany series is edited in the Museum's Department of Botany
Keeper of Botany: Mr J. F. M. Cannon

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ISBN 565 08018

ISSN 0068-2292 Botany series

Vol 17 complete

British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 25 February 1988

25FEBi988

Studies in Pseudocyphellaria (lichens)
I. The New Zealand species

BRITISI

PRESENTED
GENEHAL LIBRARY

David J. Galloway

Department of Botany, British Museum (Natural History), Cromwell Road,
London SW7 5BD

Contents

Synopsis 2

Introduction 2

Previous collections and research 3

Materials and methods 8

Results 9

Morphology 9

Anatomy 18

Photosymbiodemes 24

Lichenicolous fungi 25

Chemistry 26

Acetate-polymalonate pathway 26

Shikimic acid pathway 29

Mevalonic acid pathway 31

Role of secondary metabolites in Pseudocyphellaria 34

Ecophysiology 35

Nitrogen fixation 35

Growth 37

Geographical setting 38

Ecology 38

Habitats 38

Climate 40

Vegetation 41

Factors affecting distribution of Pseudocyphellaria 43

Biogeography 44

Endemic element 46

Australasian element 47

Austral element 48

Palaeotropical element 49

Cosmopolitan element 49

Discussion 50

Generic concept and infrageneric classification 50

The genus Pseudocyphellaria 53

Key to New Zealand species 55

The species 57

1. P. allanii. 57

2. P. ardesiaca 61

3. P. argyracea 64

4. P. auratd 68

5. P.bartlettii...:. 72

6. P. billardierei 75

7. P. carpoloma 80

8. P. chloroleuca 85

9. P. cinnamomea 91

10. P.colensoi 95

11. P.corbettii .. 99

Bull. Br. Mus. nat. Hist. (Bot.) 17: 1-267. Issued 25 February 1988

2 D. J. GALLOWAY

12. P.coriacea 102

13. P.coronata 106

14. P. crassa 108

15. P.crocata 113

16. P.degelii 118

17. P. dissimilis 122

18. P. durietzii 126

19. P.episticta 130

20. P.faveolata 134

21. P.fimbriata 139

22. P.fimbriatoides 142

23. P.glabra 146

24. P. granulata 152

25. P.gretae 156

26. P. haywardiorum 159

27. P. homoeophylla 162

28. P.hookeri 166

29. P.intricata 169

30. P.jamesii 174

31. P.knightii 177

32. P.lindsayi 180

33. P. lividofusca 183

34. P. maculata 187

35. P. margaretiae 191

36. P. montagnei 195

37. P.multifida 199

38. P.murrayi 204

39. P.neglecta 207

40. P.nermula 210

41 . P. physciospora 213

42. P. pickeringii 218

43. P.poculifera 224

44. P. pubescens 228

45. P. rubella 231

46. P. rufovirescens 235

47. P. sericeofulva 239

48. P.wilkinsii 242

Acknowledgements 245

References 246

Index 263

Synopsis

Forty-eight species of Pseudocyphellaria Vainio are recorded and described from New Zealand, and their
known distributions there are mapped. Details of the chemistry, ecology, ecophysiology, and bio-
geography of Pseudocyphellaria are presented, together with a key to species; the generic concept and
infrageneric classification are briefly discussed. Pseudocyphellaria corbettii, P. haywardiorum, P. jamesii,
P. lindsayi, P. nermula, and P. wilkinsii are newly described, and the new combination, P. pickeringii
(Tuck.) D. Galloway, is proposed.

Introduction

Species of Pseudocyphellaria* are, in the main, conspicuous, leafy, foliose lichens best de-
veloped and with richest species diversity in rain-forest, shrubland, and successional vegetation,
or subalpine and alpine grassland habitats of the Southern Hemisphere cool-temperate zone;

* In many early accounts of New Zealand lichens, species of Pseudocyphellaria were referred to Lobaria, Ricasolia,
Sticta, or Stictina. The genus name Pseudocyphellaria Vainio requires conservation, and a proposal to this effect is in
preparation.

PSEUDOCYPHELLARIA 3

only a few taxa extend into tropical and/or Northern Hemisphere temperate, oceanic habitats.
At present c. 110 species are known worldwide.

New Zealand is one of the two principal areas of endemism and speciation in the genus, the
other being southern South America. Because of their often large size, ease of collection, and
attractive colouring, species of Pseudocyphellaria were among the first lichens collected from
New Zealand by early botanical explorers. William Lauder Lindsay (1829-1880) the Scottish
doctor and lichenologist who botanized in the Otago settlement for several months in 1861
commented 'New Zealand is par excellence, the country of the Stictae. Not only do they there
occur in the greatest absolute as well as relative numbers, but they there attain their maximum
development, size and beauty . . . This preponderance of the Stictae, their frequently great size
and beauty of colouring, and the profusion of individuals give a sometimes peculiar character to
the foliaceous Lichen-flora of New Zealand' (Lindsay, 1869).

The luxuriance of growth and the species diversity of Pseudocyphellaria in New Zealand,
together with its novel and complex array of secondary metabolites (see below under Chemistry)
make it a genus of exceptional interest and significance. However, before species of Pseudo-
cyphellaria can be used in a variety of important, applied studies, an accurate recognition of the
taxa present in New Zealand is necessary, together with a stable system of nomenclature. In the
10 years that have elapsed since I and various collaborators first started work on Pseudo-
cyphellaria, emphasis has, of necessity, been placed on the evaluation and solution of the often
complex nomenclatural and taxonomic problems that the New Zealand and Southern Hemi-
sphere species presented (Galloway, 19836, 19850, 19856, 19860, 19866; Galloway & James,
1977, 1980, 1986; Galloway et al., 19836; Hawksworth & Galloway, 1984; Renner & Galloway,
1982). Recently, an introductory account of the New Zealand species appeared (Galloway,
19856) dealing with 42 species.

In anticipation of future advances in cool-temperate lichen biogeography, chemistry, eco-
physiology , and phytosociology, the present study of 48 species is offered as an introduction to
an important group of Southern Hemisphere lichens.

Previous collections and research

The 18th century

The New Zealand botanical record begins with the Endeavour voyage of Captain Cook
(1768-1771) and the collections of Joseph Banks and Daniel Carlsson Solander from various
coastal localities, between 8 October 1769 and 31 March 1770. Godley (1983) states 'During
these 174 days Endeavour was at anchor for 55 days and on 44 days the botanists went ashore.
Their collection of some 360 species [this does not include lichens] was the first ever made of New
Zealand plants. That it was such a comprehensive collection of the coastal and lowland plants of
northern New Zealand resulted from a happy chance of a visit during the spring and summer'.
From Cook's circumnavigation several coastal lichens were also collected, among the earliest
from the Southern Hemisphere (Galloway, 1985c) and these collections, now in the herbarium
of the British Museum (Natural History), contain four species of Pseudocyphellaria viz., P.
carpoloma, P. coronata, P. crocata [annotated in pencil in Banks's hand 'Lichen 4 nova'], and P.
dissimilis [annotated 'Lichen 5 nova'].

Cook's second voyage of 1772-1775 (David, 1981), subjected two main New Zealand
localities to greater botanical scrutiny viz., Queen Charlotte Sound in Cook Strait and Dusky
Sound in Fiordland, the latter named but not visited by Cook on the Endeavour voyage. Cook
had with him on his second great circumnavigation the biologists J. R. and J. G. A. Forster and
Anders Sparrmann (Du Rietz, 1981; Hoare, 1981, 1982; Lysaght, 1981) and from their
collections the first published description of a Southern Hemisphere species of Pseudo-
cyphellaria [as Lichen berberinus G. Forster] was made from material gathered in Tierra del
Fuego (Forster, 1789; Galloway & James, 1977). The Forsters also collected in Dusky Sound,
though in George Forster's Florulae Insularum Australium Prodromus (1786) the five lichens
listed from Australia and New Zealand are only given Roman numerals and remain unnamed.

D. J. GALLOWAY

From Forster's New Zealand lichen collections [preserved in BM, LINN-SM, S, and UPS-
THUNBERG- it is also possible that additional material may be found in Gottingen and Paris]
Swartz (1781) named Lichen filix (Stictafilix (Sw.) Nyl.), the first Southern Hemisphere lichen to
be described and illustrated and which was later depicted in colour engravings in both Hoffmann
(1801) and Delise (1825). Among Forster's known lichens (Galloway, 1981c), a specimen of P.
billardierei is preserved in BM though no specific locality is given for it. William Anderson and
Thomas Andrews also collected lichens at Dusky Sound (Galloway, 1981c) and in BM there is a
specimen of P. coronata which is annotated 'Dusky Bay, Capt. Cook 1775'. The date here is in
error, since Cook was in Dusky Sound from March to May, 1773.

The Scottish botanist Archibald Menzies (1754-1842), naturalist and later surgeon on
Vancouver's Discovery voyage of 1791-1795 (Lamb, 1984; Galloway & Groves, 1987) was in
Dusky Sound in 1791 , and during his time there made the most extensive 18th century collection
of New Zealand cryptogams. Among these were 16 lichen species in seven genera (Galloway,
unpublished), with six being species of Pseudocyphellaria (P. billardierei, P. cinnamomea, P.
coronata, P. faveolata, P. lividofusca, and P. multifida). A specimen of P. aurata (E) labelled by
Menzies in pencil, probably when he was an old man as the handwriting is unsteady [the other
collections are annotated by Menzies in ink], is most likely from a non-New Zealand tropical
collection, as P. aurata does not occur anywhere in Dusky Sound.

The 1 9th century

For the first half of the 19th century published records of New Zealand material referable to
Pseudocyphellaria (published initially under Lobaria, Ricasolia, Sticta, or Stictina) derive
mainly from visits of either British or French botanists to North Auckland and/or to the
subantarctic islands. The French were first on the scene and lichenological discoveries then
made (including species of Pseudocyphellaria) were closely linked with the name of Dumont-
D'Urville (Galloway, 1985c). On 11 August 1822 the corvette Coquille, under the command of
Louis Isidore Duperrey, left Toulon on a round-the-world voyage to study terrestrial magnet-
ism, meteorology, and natural history. Duperrey's second-in-command was J. S. C. Dumont-
D'Urville, a botanist of some eminence, who was assisted by R. P. Lesson, elder brother of P. A.
Lesson (botanist with D'Urville on the Astrolabe: 1826-1829). D'Urville and Lesson collected
lichens from the Bay of Islands, northern New Zealand between 3-17 April 1824 (Galloway &
James, 1986) and their collections are preserved in Paris (PC-LENORMAND and PC-
THURET) . Specimens of Pseudocyphellaria from the Coquille voyage were examined by Delise
shortly after the expedition's return to France and from New Zealand material he described
Sticta carpoloma Delise, and recorded S. aurata and S. angustata (Delise, 1825a, I825b). The
lichenological account of the Coquille voyage was prepared by Bory de St- Vincent (1829) who
recorded Sticta aurata and S. carpoloma from Lesson's Bay of Islands collections.

D'Urville's second expedition (1826-1829) in the Astrolabe (the renamed Coquille) produced
many important botanical discoveries in New Zealand, from the Thames estuary, and from
Astrolabe Harbour on the Nelson shores of Cook Strait. Achille Richard's account of the New
Zealand lichens discussed 27 taxa, among them four species of Pseudocyphellaria viz., Sticta
aurata, S. mougeotiana [= P. crocata], S. carpoloma, and a new species, 5. cinnamomea A.
Rich, (Richard, 1832). The atlas of plates (Richard, 1833) contained handsome coloured
engravings of Sticta cinnamomea and S. carpoloma. However, Sticta carpoloma sensu Richard is
an entity with white pseudocyphellae (and not yellow as in 5. carpoloma Delise), and is referable
to P. rufovirescens. Montagne (1835) recognized Richard's description and illustration of S.
carpoloma as a new species differing from S. carpoloma sens. str. when he noted 'S. carpoloma
Rich. . . . non Delise, quaetotohabitucyphellisquecitrinisdistinctissima'. This new species he
named Sticta richardi Mont., basing his description on material collected by Bertero from Juan
Fernandez. Later, Montagne (1845, 1856) referred all similar New Zealand material (including
the type of P. rufovirescens) to 5. richardi; however, Juan Fernandez material is an independent
taxon separable morphologically and chemically from the New Zealand P. rufovirescens.

Cunningham's (1836) account of the botany of northern New Zealand contains four species

PSEUDOCYPHELLARIA 5

referable to Pseudocyphellaria, viz., S. aurata, S. carpoloma, S. cinnamomea, and S.
mougeotiana, based on the reports of Delise (1825) and Richard (1832).

D'Urville's third Pacific expedition (1837-1840) visited the Auckland Islands and towards the
end of the voyage several anchorages were made in New Zealand, first in Hooper's Inlet, Otago
Peninsula, then in the wooded harbour of Akaroa (now completely deforested) where rich
gatherings of Pseudocyphellaria were made, and finally, in the Bay of Islands. Collecting was
under the charge of Jaques Hombron, senior surgeon of the Astrolabe, and Honore Jacquinot,
junior surgeon of the Zelee, with the senior surgeon of that vessel, E. Le Guillou, also collecting
lichens. Their lichens from this time are preserved in Paris, the Akaroa collections of P.
rufovirescens (including the type in BM) being particularly handsome and complete with elegant
printed labels giving details of habitat, locality, date, and collector. The lichens of the voyage
were published by Montagne (1845) who recorded Sticta delisea [= P. glabra], S. faveolata, S.
orygmaea [= P. coronata], and 5. richardi [= P. rufovirescens] from the Auckland Islands and
Akaroa. The Atlas contains a sumptuous plate devoted to lichens with a fine coloured engraving
of P. coronata (pi. 15, fig. 1).

However, it was the visit of J. D. Hooker to the Bay of Islands in 1841, during the Antarctic
voyage (1839-1843) of the ships Erebus and Terror commanded by Capt. James Clark Ross,
which gave lichenology in 19th century New Zealand its initial strong boost. Hooker, assistant-
surgeon of H.M.S. Erebus, stayed three months at the Bay of Islands, and in company with
David Lyall (assistant-surgeon of the Terror), William Colenso, and Andrew Sinclair, collected
many lichens from northern coastal forests, as well as making good collections from the
Auckland Islands and from Campbell Island. Hooker's Antarctic lichens which included those
from New Zealand and Tasmania were prepared for publication by Thomas Taylor (Hooker &
Taylor, 1844; Taylor & Hooker, 1845) and included 11 taxa referable to Pseudocyphellaria of
which Sticta glabr a J. D. Hook. & Taylor, S. celluliferaL D. Hook. & Taylor [= P. faveolata], S.
linearis J. D. Hook. & Taylor [= P. billardierei], S. impressa J. D. Hook. & Taylor [= P.
faveolata], S. flavicans J. D. Hook. & Taylor, S. coriacea J. D. Hook. & Taylor, and 5.
chloroleuca J. D. Hook. & Taylor, were new taxa.

Raoul's (1846) list of 11 names referable to Pseudocyphellaria was taken from the accounts of
Hooker & Taylor (1844) and Richard (1832), and added no new information.

The first detailed study of New Zealand species of Pseudocyphellaria [as the genus Sticta] was
that of Babington (1855), based on Hooker's New Zealand collections, supplemented with a
wide range of both historical and more recently gathered material including collections of Banks
& Solander, Forster, Anderson, Menzies, Allan and Richard Cunningham, Bidwill, W.
Stephenson, Samuel Mossman, Everard Home, Richard Taylor, Colenso, Lyall, Monro, and
Sinclair, as well as specimens from the Paris herbarium. Babington discussed 25 taxa in some
detail, provided a rudimentary infrageneric classification of Sticta, and described the following
new taxa: 5. colensoi and var. pinnatifida Church. Bab. , 5. richardi var. glauca Church. Bab. , 5.
richardi var. rufovirescens Church. Bab. , S. fragillima Church. Bab. , 5. granulata Church. Bab. ,
5. hookeri Church. Bab., S. limbata var. subflavida Church. Bab., and S. montagnei Church.
Bab. The account also provided excellent illustrations (by W. Fitch) of 5. colensoi, S.
carpoloma, S. faveolata, S. hookeri, and S. coriacea, and it stands as the major early account of
the genus after Delise (1825, 1825ft), with five of his names still in use today.

For the next 50 years relatively few new taxa referable to Pseudocyphellaria were described
from New Zealand collections despite extensive literature, viz. , Hooker, 1867; Hue, 1890, 1901;
Knight, 1871, 1880; Krempelhuber, 1868, 1870, 1876a, 1876b, 1881; Lindsay, 1859, 1866, 1867,
1869; Lojka, 1886; Miiller Argoviensis, 1883, 1887a, 1887ft, 1894, 1895; Nylander, 1858fl,
1858ft, 1860ft, 1865, 1866, 1868ft, 1876; Stizenberger, 1889, 1895; Tuckerman, 1874.

Subsequent to Babington's account, new taxa from New Zealand referable to Pseudocyphel-
laria appeared sporadically until the end of the century and included Sticta fossulata var.
physciospora Nyl. (Nylander, 1860ft); Sticta subcoriacea Nyl. [= P. coriacea] (Nylander, 1865);
5. episticta Nyl. (Nylander, 1865); Stictina fragillima f. lutescens Nyl. [= P. cinnamomea]
(Nylander, 1866); Sticta glaucolurida Nyl. [= P. carpoloma], S. homoeophylla Nyl., 5. sub-
variabilis Nyl. [= P. multifida] (Nylander, 1867); Sticta freycinetii var. isidioloma Nyl. [= P.

6 D. J. GALLOWAY

glabra], Ricasolia elaphocera Nyl. [= P. coriacea] (Nylander, 1868ft); 5. urvillei var.flavicans f.
laceratula Krempelh. [= P. pickeringii], S. freycinetii \ at. stauromatica Krempelh. [= P. glabra],
S. faveolata f. angustifolia Krempelh., nom. nud. [= P. billardierei] (Krempelhuber, 1870); Sticta
hirta Stirton [= P. coronata] (Stirton, 1873); Sticta pickeringii Tuck. (Tuckerman, 1874); Sticta
fossulata f. pallida Krempelh. [=P. mfovirescens] (Krempelhuber, 1876); 5. lividofusca
Krempelh. (Krempelhuber, 1876ft); 5. canaliculata Knight [=P. coriacea] (Knight, 1877); 5.
coronata Miill. Arg. (Miiller Argoviensis, 1879); S. amphisticta Knight [= P. lividofusca]
(Knight, 1880); 5. fossulata f. expallida Krempelh. [= P. rufovirescens], Parmelia isabellina
Krempelh. [= P. glabra] (Krempelhuber, 1881); 5. borneti Miill. Arg. [= P. carpoloma] (Miiller
Argoviensis, 1882); Stictina fragillima var. dissecta Miill. Arg. [= P. dissimilis], Sticta freycinetii
var. glabrescens Miill. Arg. [=P. glabra], S. freycinetii var. tenuis Miill. Arg. [= P. glabra]
(Miiller Argoviensis, 1883); 5. fossulata var. subcyphellata Nyl. [= P. rufovirescens] (Nylander,
1888ft); Stictina fragillima f. sublutescens Hue [= P. cinnamomea] (Hue, 1890); S. fragillima var.
myrioloba Miill. Arg. [= P. fimbriatoides], S. mougeotiana var. dissecta Miill. Arg. [= P.
neglecta], Sticta pubescens Miill. Arg., S. psilophylla Miill. Arg. [= P. chloroleuca], S. psilophyl-
la f. amphicarpa Miill. Arg. [= P. multifida], S. amphisticta \ar.platyloba Miill. Arg. [= P.
lividofusca] (Miiller Argoviensis, 1892); S. elatior Stirton [= P. faveolata], S. expansa Stirton
[= P. carpoloma], S. lorifera Stirton [= P. faveolata] (Stirton, 1900).

The last 15 years of the 19th century saw several detailed accounts or lists of New Zealand
Lobariaceae in which species of Pseudocyphellaria are recorded, viz., Nylander, 1888ft; Hue,
1890; Miiller Argoviensis, 1892, 1894, 1896, most based largely on collections of Charles Knight,
William Colenso, and the material from the Hookerian herbarium at Kew, with Hellbom's
(1896) account, where many taxa now in Pseudocyphellaria are placed in Lobaria, detailing
lichens collected in New Zealand in 1875-1875 by Sven Berggren. John Buchanan's lichens from
Otago and Wellington, and T. W. N. Beckett's collections from Canterbury and the Manawatu
were studied by James Stirton in Glasgow (Stirton, 1873, 1875, 1898, 1900), who described
several taxa from them referable to Pseudocyphellaria.

The major 19th century accounts record the following numbers of taxa of Pseudocyphellaria:
11 taxa (Hooker & Taylor, 1844), 24 taxa (Babington, 1855), 34 taxa (Nylander, 1888ft), 52 taxa
(Miiller Argoviensis, 1894), and 50 taxa (Hellbom, 1896); however, many of the names recorded
in these accounts were synonyms of widespread and polymorphic species. At the close of
the 19th century 27 species of Pseudocyphellaria accepted today were known from New Zeal-
and: P. argyracea, P. aurata, P. billardierei, P. carpoloma, P. chloroleuca, P. cinnamomea,
P. colensoi, P. coriacea, P. coronata, P. crocata, P. dissimilis, P. episticta, P. faveolata, P.
glabra, P. granulata, P. homoeophylla, P. hookeri, P. intricata, P. lividofusca, P. montagnei,
P. multifida, P. neglecta, P. physciospora, P. pickeringii, P. pubescens, P. rubella, and P.
rufovirescens.

The 20th century

In contrast to the taxonomic activity of the late 19th century, the first 40 years of the present
century saw very little new work on the New Zealand Lobariaceae apart from the discursive
account of Hue (1901) which was mainly a reworking of Nylander-annotated material in Paris,
supplemented with collections of Filhol from Campbell I. and New Zealand, H. L. Travers from
the Chatham Is and New Zealand, and a few specimens collected by Cockayne from Stewart I.
and identified by Lindau (Cockayne, 1910) ; and Cheel's two bibliographical treatments of South
Pacific Lobariaceae (Cheel, 1912, 1914).

The first lichenologist to undertake a comprehensive survey of Pseudocyphellaria in the sense
of Vainio (1890) was the Uppsala botanist G. E. Du Rietz. In the early 1920s he began serious
work on Southern Hemisphere species of Sticta and Pseudocyphellaria and annotations he made
on numerous specimens in H, S, UPS, and W, attest his knowledge of taxa in these genera.
However, he published only a brief note on the Southern Hemisphere species P. freycinetii and
P. chloroleuca (Du Rietz, 1924), his use of the latter name being sensu P. glabra. He made large
collections from New Zealand and Australia in 1925-1926, but these were never worked up for

PSEUDOCYPHELLARIA 7

publication, although a few of his New Zealand collections were referred to by Magnusson
(1940). Du Rietz was, however, instrumental in encouraging local New Zealand botanists to
collect lichens, especially H. H. Allan, the first Director of Botany Division, DSIR. Allan sent
New Zealand lichens to Wien to Zahlbruckner, and besides his own extensive gatherings of
Pseudocyphellaria, specimens from Lucy B. Moore and Lucy M. Cranwell from various North I.
localities, and of J. S. Thomson and G. Simpson from South I. habitats were also included.
Zahlbruckner's posthumous work Lichenes Novae Zelandiae . . . (1941) includes the first major
account of New Zealand Lobariaceae since the 19th century (pp. 279-293) with 50 taxa of
Pseudocyphellaria represented in Sticta (sections Eusticta and Stictina). Among these, Zahl-
bruckner described the following new taxa: Sticta condensata Zahlbr. [=F. faveolata], S.
coriacea f. vestitula Zahlbr., [= P. coriacea], S. amplificata Zahlbr. [=P. homoeophylla], S.
variabilis var. cinerata Zahlbr. [= P. chloroleuca], and S. intricata var . fimbriata Zahlbr. [= P.
dissimilis]. Zahlbruckner had a rather confused idea of some taxa, and in his list of 50 names,
only 24 are referable to Pseudocyphellaria, now accepted as occurring in New Zealand. He
apparently relied on published descriptions or earlier accounts of names reported from New
Zealand and obviously examined very few types. For instance, although he correctly identified
P. homoeophylla in a few cases, he also described Sticta amplificata for well-developed
specimens of this species, and placed other collections of it under S. dissimulata, S. sinuosa, and
even S. cinnamomea. His account is, however, an advance on earlier treatments, for it surveys a
much wider range of collections, from North Auckland, the Volcanic Plateau, Mt Egmont, and
Wellington in North I., and from Maryborough, Westland, Otago, Southland, and Fiordland in
South I. It must also be noted that Zahlbruckner died before a final revision of his manuscript
could be made and hence his published account contains a number of errors that may well have
been rectified had he lived. For the next 30 years, Zahlbruckner's treatment of taxa referable to
Pseudocyphellaria strongly influenced succeeding bibliographic, chemical, ecological, and
taxonomic accounts, notably Allan (1948, 1949), Martin (1965, 1966, 1968, 1969a, 19696), and
Martin & Child (1972).

Although species of Pseudocyphellaria are amongst the most prominent and widespread of
New Zealand's foliose lichens and constitute an appreciable biomass in forest ecosystems, they
were largely dismissed (together with bryophytes) from ecological studies until the 1960s. H. H.
Allan (1927) appealed for a greater awareness of lower plants in ecological studies 'it is now
realised that mosses, lichens, liverworts and fungi are of equal importance with the so-called
"higher" plants, and have equally important lessons to teach- economic no less than "purely
scientific" ' However, apart from a brief reference to lichens from the Antipodes (Cockayne,
1928) and Jablonszky's collections from Waikaremoana (Szatala, 1939), lichens were ignored in
floristic and ecological surveys until the posthumous work of Murray (1963) and the influence of
P. W. James (in Mark etal., 1964), who curated Murray's lichens (OTA) in 1962-1963.

Floristic and ecological work in which species of Pseudocyphellaria are recorded appeared
first in Otago (Martin, 1965, 19690, 19696, 1970; Galloway, 1966, 1968a, 19686; Mark & Bliss,
1970; Scott, 1970; Scott & Armstrong, 1966; Scott & Rowley, 1975) to be followed by
information from Canterbury (Dodge, 1971 [P. neozelandica Dodge = P. colensoi]; Galloway,
1976, 19786; J0rgensen & Galloway, 1983; Burrows, 1977), Westland (Wardle, 1977; Simpson
& Galloway, 1981), and Nelson (Galloway & Simpson, 1978). North I. floristic studies in the
1970s discussed species of Pseudocyphellaria from a number of offshore islands (B. W.
Hayward, 1973; B. W. & G. C. Hayward, 1974a, 19746; 1979, 1980, 19820; 19826; 1983; G. C. &
B. W. Hayward, 19730, 19736, 1978; Hayward etal., 1975, 1976, 1981, 1986; Dakin & Galloway,
1980); from the Urewera National Park (see work of Green, and of Snelgar under lichen
physiology below); and from Mt Tarawera (Clarkson & Clarkson, 1983). Species of Pseudo-
cyphellaria from the subantarctic islands (including the Auckland, Campbell, Macquarie, and
Snares groups) are discussed by Dodge (1948), Dodge & Rudolph (1955), Dodge (1971),
Fineran (1971), Imshaug [1977- a particularly important and overlooked paper giving a
convincing account of the distinction of the closely similar faveolate taxa P. billardierei (as P.
flotowiana), P. physciospora, and P. faveolata, using anatomical, chemical, and statistical
methods], and Filson, 19810, 19816.

8 D. J. GALLOWAY

Chemical work on New Zealand species of Pseudocyphellaria begun by Murray (1952) was
continued with great success by Corbett and his students at Otago University, and by Wilkins at
Waikato University (see below under Chemistry).

Taxonomic studies on New Zealand Pseudocyphellaria undertaken preparatory to the present
study include Galloway, 1983ft, 1985fl, 1985ft, 1986ft; Galloway & James 1977, 1980; Galloway
et al., 1983ft; Hawksworth & Galloway, 1984; and Renner & Galloway, 1982. During these
studies the following new taxa were established: P. allanii D. Galloway, P. ardesiaca D.
Galloway, P. bartlettii D. Galloway, P. crassa D. Galloway, P. degelii D. Galloway & P. James,
P. durietzii D. Galloway, P. fimbriata D. Galloway & P. James, P. fimbriatoides D. Galloway &
P. James, P. gretae D. Galloway, P. knightii D. Galloway, P. maculata D. Galloway, P.
margaretiae D. Galloway, P. murrayi D. Galloway, and P. sericeofulva D. Galloway.

Materials and methods

The following account is based on examination of an extensive range of herbarium material as
well as collection and examination of material in the field from 1963-1983. Type and representa-
tive material was obtained from the following herbaria: ABD, AK, AKU, BM, BP, CANU,
CHR, COLO, E, FH, FI, G, GB, GZU, H, H-NYL, L, LD, LINN-SM, M, MANCH, MB,
MEL, MSC, NY, OTA, PC-HUE, PC-LENORMAND, PC-MONTAGNE, PC-THURET, S,
U, UPS, UPS-THUNBERG, US, VER, W, WELT, WRSL, WU, ZT, and from private
herbaria of the following individuals: the late J. K. Bartlett (now in AK), G. Degelius (Askim),
J. A. Elix (Canberra), B. W. & G. C. Hayward (Lower Hutt), L. Tibell (Uppsala), and V.
Wirth (Stuttgart). A representative range of all taxa discussed in this paper is preserved in BM.

Field studies: With the exception of P. bartlettii, P. haywardiorum, P. physciospora, and P.
sericeofulva, all of the species discussed below were studied in their natural habitats in order to
assess the extent of environmentally induced variation, as well as to determine individual
ecological requirements. Field studies ranged from the Three Kings Is (Lat. 346'S) to southern
Stewart I. (Lat. 48S). Most of the major geographical regions of New Zealand were surveyed,
with the exception of North Auckland, the east coast of North I. and north-west Nelson,
Marlborough, and South Westland in South I. The subantarctic islands (Snares, Auckland,
Antipodes, Campbell, and Macquarie Is) were not visited, nor were the Chatham Is, although
collections from the Auckland, Antipodes, Campbell, Chatham, Macquarie, and Snares Is were
examined in the herbarium.

Light microscopy: Observations and measurements of external features were made at x 10 to
x 30 with an Olympus stereomicroscope. Anatomical features were investigated with a Leitz
Dialux 20 microscope. Sections were cut with a Kryomat freezing microtome giving section
thicknesses of 15-20 urn. Sections were normally mounted in water followed by (or directly in)
10% KOH(K), but cotton-blue in lactophenol was also sometimes used. Tests for amyloid
reactions were made by mounting directly in Lugol's iodine solution, or in this solution following
treatment with 10% K.

Scanning electron microscopy (SEM) : Material for SEM was air-dried, attached to aluminium
stubs with araldite and sputter-coated using a gold-palladium or a gold target. Stubs were
examined in an ISI 60A microscope using back-scatter detection.

Thin-layer chromatography (t.l.c.): Representative samples of all taxa referred to in this study
(including all type specimens) were analysed by t.l.c. using standard procedures (Culberson,
1972; Culberson & Johnson, 1976: Culberson et al., 1981; Wilkins & James, 1979; White &
James 1985).

Species descriptions are based on type material supplemented with a range of recently
collected, characteristic specimens. Details of distinguishing features, variation, chemistry,
ecology, and distribution are given for each species together with notes on typification and
nomenclature when applicable. All species have distribution maps, though full distribution lists
are given only in instances where critical or new taxa are involved.

PSEUDOCYPHELLARIA

Abbreviations of describing authors follow Laundon (1979); those of journals to the third
(1980) edition of Serial Publications in the British Museum (Natural History) Library; and those
of books to Hawksworth (1974) and to Stafleu & Cowan (1976-1986).

Results

Morphology

Species of Pseudocyphellaria discussed in this revision are conspicuous, foliose lichens growing
on twigs, bark, soil, or rock, often over or amongst bryophytes or other lichens in a wide variety
of habitats. Morphological features useful in species delimitation are discussed under sub-
headings below.

Habit: Thalli of Pseudocyphellaria are often very large, (2-)6-20(-50) cm diam. ; some of the
species found in New Zealand are among the largest of foliose lichens known, orbicular,
rosette-forming or irregularly laciniate, subpendulous to straplike, and are characterized by
scattered pseudocyphellae on the lower surface, occasionally also on the upper surface as well.
Lobes are very variable in size, shape, and branching, but even so it is possible to distinguish
three main morphologies although certain species may show a degree of overlap, depending on
local microclimatic and microhabitat parameters. (1) Monophyllous: no discernible branching
at the apices, with the lobes broadly rounded, often compact, and rosette-forming, e.g. P.
ardesiaca, P. bartlettii, P. coronata (twig forms), P. crocata, P. intricata, P. neglecta, and P.
rubella. (2) Dichotomous: lobes are narrow, often also somewhat straplike, and are
noticeably dichotomously branched at the apices. Dichotomous branching occurs at an early
stage in the growth of the lobe and is clearly visible in even very young plants. Further apical
growth of the bifurcating tips may be equal or unequal, and the tips themselves may be acute,
truncate or rounded. Lobes distal to the developing tips may be smooth, wrinkled or
faveolate, and variable in width. Dichotomously branching species include: P. allanii, P. billar-
dierei, P. carpoloma, P. chloroleuca, P. cinnamomea, P. coriacea, P. crassa, P. degelii,
P. dissimilis, P. durietzii, P. faveolata, P. glabra, P. homoeophylla, P. jamesii, P. lindsayi, P.
lividofusca, P. maculata, P. murrayi, P. physciospora, P. pubescens, P. rufovirescens . Species
with well-defined dichotomous lobe branching are generally rather loosely attached to the
substrate and may often be hanging. (3) Complex-imbricate: no regular discernible branching
pattern is visible, often overlapping and crowding of lobes obscures an initial unequal branching
pattern. A large proportion of the taxa discussed in this account have this lobe morphology and
generally speaking those that do are more closely appressed to the substrate than those having
dichotomously branching lobes. Complex-imbricate species include: P. argyracea, P. aurata, P.
chloroleuca, P. colensoi, P. corbettii, P. coronata, P. crocata, P. degelii, P. dissimilis, P.
durietzii, P. episticta, P. fimbriata, P. fimbriatoides , P. glabra, P. granulata, P. gretae,
P. haywardiorum, P. homoeophylla, P. hookeri, P. intricata, P. knightii, P. lindsayi, P. livido-
fusca, P. maculata, P. margaretiae, P. montagnei, P. multifida, P. neglecta, P. nermula,
P. physciospora, P. pickeringii, P. poculifera, P. pubescens, P. rubella, P. sericeofulva, and
P. wilkinsii.

The margins of lobes may be entire (e.g. P. billardierei, P. homoeophylla, P. physciospora) or
variously notched, incised, pectinate, crenate or ragged (e.g. P. chloroleuca, P. colensoi, P.
multifida, P. pickeringii, P. poculifera) plane to sinuous and ascendent, or involute,
furnished with isidia (e.g. P. dissimilis, P. glabra), phyllidia (e.g. P. fimbriata, P. neglecta), or
soredia (e.g. P. aurata, P. haywardiorum, P. intricata) in punctiform, erose or labriform soralia,
and occasionally with conspicuous, linear, punctate or verruciform pseudocyphellae (most
notably in P. carpoloma, P. crassa, P. faveolata, P. margaretiae, P. murrayi, P. poculifera, P.
pubescens, and P. rubella). Many species have conspicuously thickened margins, most notice-
able on the lower surface as in P. billardierei (x 10 lens), but occasionally also on the upper
surface as well, the thickening being in the form of a narrow, raised, rounded, continuous

10 D. J. GALLOWAY

ridge. Marginal thickening is much less apparent in species having prominent marginal
pseudocyphellae (e.g. P. faveolata). In P. crassa the thick tomentum of the lower surface often
appears at the lobe margins as a distinct fringe.

Lobes are attached to the substrate either (1) by tomental hyphae over the greater part of the
lower surface, or (2) by scattered fascicles or tufts of anchoring hyphae, or (3) directly to the
lower cortex in those species with poorly developed tomentum. In Pseudocyphellaria, the
degree of attachment of thalli to the substrate appears, in certain cases at least, to be determined
in part by ecological conditions obtaining at the substrate surface, as does the expression of lobe
morphology in some variable species. Light, and moisture appear to be of prime importance,
and Snelgar & Green (19810) have shown that populations of P. dissimilis from shade,
semi-shade, and full sunlight, have different morphologies and varying thicknesses of tomentum
on the lower surface. Shade populations are only loosely attached, while well-illuminated
populations are closely attached to the substrate with only the lobe tips free. Tomentum is longer
and thicker in the sun populations, c. four times thicker than in shade populations, and three
times thicker than in semi-shade populations. Such differences correlate with the evaporative
stress of the different environments, the sun population being most able to resist desiccation by
adapting in three ways, viz. (1) the thallus has the classic xerophyte characteristic of reduced
surface area to volume ratio, (2) increased water storage capacity is attained by increased thallus
thickness and a thicker layer of tomentum, and (3) thalli are more closely appressed to the
substrate, thereby aiding water storage by tomental hyphae and reducing the area exposed to
water loss by evaporation (Snelgar & Green, 1981a: 409-410).

Colour: Thallus colour is determined by two main factors: the nature of the photobiont, and
the degree of exposure of the habitat in which the lichen grows. When moist, thalli with green
photobionts are bright lettuce-green to dark olive-green or glaucous-green; species with usnic
acid (P. corbettii, P. glabra, P. homoeophylla) are pale yellowish green, the colour intensifying
in specimens from habitats exposed to direct sunlight. Thalli with cyanobionts, when moist, vary
from purplish blue (P. ardesiaca, P. nermula, P. jamesii) to livid leaden-grey or dark grey-black
(e.g. P. allanii, P. hookeri, P. knightii) to brownish or red-black (e.g. P. crocata, P. haywar-
diorum, P. neglectd). Thallus colour is also influenced in fresh material of yellow-medulla
species (e.g. P. colensoi, P. coronata, P. degelii) by pulvinic acid derivatives in the medulla
which impart an underlying yellow or golden hue. The colour of dried material, either in the field
or on storage in the herbarium, is pale greenish grey or whitish to fawnish, buff, brown or
red-brown. Material of P. gretae and P. rubella is often tinged reddish or pinkish when dry, and
on long storage these colours often intensify. P. coronata may also be suffused purplish, both in
the field and on storage in the herbarium. Plants exposed to sunlight, or from subalpine to
high-alpine grassland habitats are often dark red-brown with increased deposition of pigments in
the upper cortex (P. crocata, P. maculata, P. neglecta), or may even become superficially
blackened (e.g. P. faveolata, P. glabra).

Texture: Thallus texture may be smooth, matt, dull, shining, or glossy in parts and dull in
others, wrinkled or plicate, to shallowly or deeply faveolate (e.g. P. carpoloma, P. billardierei,
P. faveolata, P. physciospora, P. rufovirescens) , plane or undulate, with or without isidia,
maculae, papillae, phyllidia, pseudocyphellae, pseudoisidia, soredia, or tomentum.

Tomentum: Tomental hairs occur on the upper surface of a number of species (Fig. 1B,D)
although only in four do they cover the entire upper surface (P. gretae, P. margaretiae, P.
pubescens, P. rubella). In other species, hairs are most commonly seen at, or near, lobe apices or
margins, or on marginal phyllidia, or at the base of pedicellate apothecia (e.g. in P. allanii, P.
aurata, P. coriacea, P. crassa, P. fimbriata, P. fimbriatoides , P. intricata, P. knightii, P.
poculifera). Four species (P. coronata, P. jamesii, P. nermula, and P. sericeofulva) have
tomentum rather patchily developed on the upper surface. In all cases, tomentum is whitish or
pale buff and never dark brown, red-brown or blackened as it is when developed on the lower
surface. In general, the tomentum of the upper surface is thinner and more delicate than that of
the lower surface. Partially tomentose species are often also scabrid in parts and in P. gretae

PSEUDOCYPHELLARIA 11

and P. pubescens a scabrid surface is often seen beneath the tomentum. There seems to be a
relationship between scabrosity and tomentum and a scabrid upper surface may in fact represent
an arrested early stage in the development of tomentum (Galloway, 1986a:110).

Scabrosity: A distinctive, scabrid-areolate upper surface (x 10 lens) results when the upper
cortex becomes roughened-uneven and broken up into a mosaic of minute, shallowly convex
areolae (Fig. 1A,C). Scabrosity in New Zealand species of Pseudocyphellaria is rather variable
in its occurrence, often being best seen at lobe apices and margins, or on the thalline exciple of
apothecia (P. aurata, P. colensoi, P. poculifera) , and in irregular patches on the upper surface.
There are no distinctively scabrid-areolate species in New Zealand analogous to P. compar, P.
scabrosa, or P. vaccina from southern South America (Galloway, 1986a); however, in a number
of taxa both tomentum and a scabrid-areolate cortex occur together (P. coriacea, P. crassa, P.
fimbriata, P. gretae, P. margaretiae, and P. pubescens). Taxa with scabrid-areolate lobes
(margins, upper surface or thalline exciple) include P. allanii, P. ardesiaca, P. aurata, P. chlor-
oleuca, P. colensoi, P. coronata, P. crassa, P. degelii, P. fimbriata, P. fimbriatoides , P. gretae, P.
intricata, P. margaretiae, P. nermula, P. pickeringii, P. poculifera, and P. pubescens.

Faveolae: These are shallow to deep depressions, usually smoothly concave, formed between
a reticulate pattern of interconnecting ridges which may be broad to narrow, smoothly
rounded to abruptly and sharply defined (Fig. 2). Faveolae are especially characteristic of
many taxa in Pseudocyphellaria, being more common in narrow-lobed, dichotomously
branching species than in broad-lobed species. The following faveolate species are known from
New Zealand: P. bartlettii, P. billardierei, P. carpoloma, P. chloroleuca, P. colensoi, P. coronata,
P. crassa, P. crocata, P. degelii, P. durietzii, P.faveolata, P. granulata, P. hookeri, P. intricata,
P. jamesii, P. maculata, P. montagnei, P. murrayi, P. neglecta, P. physciospora, P. pickeringii,
P. pubescens, and P. rufovirescens . Green et al. (1985:68, tab. 3) on the basis of experiments

Fig. 1 Scabrosity and tomentum of upper surface. A. Pseudocyphellaria allanii x 400. B. P. margaretiae
x 200. C. P. nermula x 500. D. P. rubella x 500.

*?ff

Fig. 2 Faveolae on upper surface. A. Pseudocyphellaria crassa. B. P. durietzii. C. .P. faveolata. D. P.

with P. rufovirescens [cited as P. billardierei] suggest that faveolae of the upper surface may
serve as reservoirs to increase external water storage.

Papillae: Papillae are minute, pale, usually hemispherical swellings in the upper cortex (Fig. 3)
which are not cephalodia, pycnidia or apothecial initials (use x 10 lens). Their occurrence is
spasmodic and varies from sparse to frequent. They are commonly seen in P. cinnamomea and
P. dissimilis, but are also known from P. corbettii, P. fimbriatoides , P. glabra, P. knightii, P.
multifida, and P. wilkinsii.

Maculae: Maculae are macroscopic (x 10 lens) photobiont-free, spots or lines visible on the
upper surface of certain species as pale white or yellow areas, corresponding to discontinuities in
the photobiont layer and imparting a characteristic marbled appearance to the upper surface.
The spots or lines which may be arranged in a reticulate pattern are best seen in moist thalli
and are often more apparent towards the lobe margins than elsewhere on the surface. Maculae
are most often found in taxa with Nostoc as photobiont, e.g. P. ardesiaca, P. bartlettii, P.

Fig. 3 Papillae on upper surface. A. Pseudocyphellaria cinnamomea x 500. B. P.fimbriatoides x 500.

PSEUDOCYPHELLARIA

13

cinnamomea, P. crocata, P. dissimilis, P. hookeri, P. intricata, P. jamesii, P. knightii, P.
maculata, P. murrayi, P nermula, and P. sericeofulva. Occasionally they occur in species with
a green photobiont such as P. degelii and P. glabra, though they are much less common than
in species with cyanobionts. Certain species, e.g. P. dissimilis, sometimes develop extensive
photobiont-free patches, though the cause of these often extensive local decolorizations of the
photobiont cells is at present unknown.

Pseudocyphellae: These are pores in cortical tissue which are usually smaller and less
differentiated than cyphellae (Hale, 1981 ; Lawrey, 1984). Their occurrence on the lower surface
is a major feature of species of Pseudocyphellaria, although in a few taxa they may be rare or
absent (e.g. P. dubia, P. berteroana, P. epistictd) and in others they may in addition be present
on the upper surface or at the margins. The development of pseudocyphellae is discussed by
Hale (1981) and Lawrey (1984). Pseudocyphellae being plugged with medullary hyphae (Fig. 4),

Fig. 4 Pseudocyphellae on lower surface. A. Pseudocyphellaria crassa x 200. B. P. faveolata x 50. C. P.
fimbrlata x 30. D. P. lividofusca x 50. E. P. multifida x 250. F. P. physciospora x 95.

14 D. J. GALLOWAY

provide a pathway for diffusion of oxygen and carbon dioxide from both upper and lower lichen
surfaces to the interior and have, as well as cyphellae, been implicated in processes of gas
exchange (see Henssen & Jahns, 1973; Rundel et al., 1979; Hale, 1981; Green et al., 1985),
although this was only proved conclusively for cyphellae (Green et al., 1981). Hale (1981)
suggests that gas exchange structures such as pseudocyphellae evolved to overcome the cortical
resistances to gas diffusion in large foliose thalli with lobes 5-20 mm wide. However, of the 48
species of Pseudocyphellaria in New Zealand, only six have pseudocyphellae on the upper
surface, and of the remainder many reach a very large size with lobes in excess of 20 mm wide, so
either gas exchange must occur through the upper cortex of these species or else, and much more
likely, gas exchange is facilitated by pseudocyphellae at the lobe margins and on the lower
surface. Green et al. (1981) monitored carbon dioxide exchange in Sticta latifrons using a split
chamber which allowed separate analysis of gas exchange at the upper and lower surfaces of the
thallus. Their results showed that carbon dioxide exchange occurred largely through the lower
surface, providing strong circumstantial evidence for the notion that cyphellae have a central
role in carbon dioxide uptake during photosynthesis and oxygen uptake during respiration. It is
likely that similar results would be obtained with large species of Pseudocyphellaria. More
recently, Green et al. (1985) have suggested that pore systems (cyphellae and pseudocyphellae)
in members of the Lobariaceae have three possible adaptations which are of advantage to the
lichen system: (1) recycling of respired carbon dioxide (from both photobiont and mycobiont),
(2) maintaining a moist internal atmosphere and insulating cells of the photobiont from external
drying, and (3) surface water storage. Since pore systems confine carbon dioxide exchange to a
small proportion of the thallus surface, liquid water can then be stored over the remainder of the
surface without affecting carbon dioxide diffusion.

Pseudocyphellae are found on the upper surface of six species: P. argyracea, P. episticta, P.
knightii, P. lindsayi, P. lividofusca, and P. wilkinsii (Fig. 5), and are verruciform, margins raised
or not, round, ellipsoid to linear, to irregular, ulcerose, and becoming sorediate-
pseudoisidiate in P. argyracea (Fig. 6A) or surrounded by fragile isidia in P. wilkinsii (Fig. 6B).

Pseudocyphellae are present on the lower surface of all species of Pseudocyphellaria
investigated in this study, although in a few cases (e.g. P. intricata, P. episticta, P. montagnei, P.
wilkinsii) they may be inapparent or only very occasionally produced, and then are often most
readily seen at or near lobe margins. They are very variable in morphology and also in
occurrence, being widely scattered to crowded. Generally pseudocyphellae are smaller at the
margins and larger centrally, ranging in size from 0-l-2(-2-5) mm diam. , in shape from round to
sigmoid, irregular to linear. The decorticate area which may be flat, concave or convex, is
either yellow or white. Pseudocyphellae are either flat and flecklike as in P. chloroleuca, P.
cinnamomea, P. dissimilis, P. multifida, and P. rufovirescens , deeply urceolate, starkly white,
and often with raised, narrow to inapparent margins as in P. coriacea, punctiform to
verruciform to large-ulcerose with a prominent coarsely granular decorticate area as in
P. intricata, to conical, verruciform-papillate to pulverulent, or even pin-prick-like (P.
bartlettii). In some taxa pseudocyphellae have noticeable margins (x 10 lens) which may be thin
to thick, smooth and glossy to matt and puckered. Species with prominent marginal
pseudocyphellae include P. carpoloma, P. crassa, P.faveolata, P. maculata, P. margaretiae , P.
murrayi, P. pubescens, and P. rubella, and the pseudocyphellae range from scattered to
numerous, prominent and conical-verruciform to punctiform or short-linear.

Deposition of the yellow pulvinic acid derivatives (calycin , pulvinic acid , pulvinic dilactone) in
the exposed hyphae of pseudocyphellae is a good taxonomic character and one often used for the
separation of related species (e.g. P. carpoloma and P. faveolatd). In some cases (e.g. in P.
physciospora) the amount of yellow pigment present may be small and this, together with the
sometimes small decorticate area of pseudocyphellae, may necessitate a careful examination of
pseudocyphellae with a strong lens (x 10-X20). In a small number of species with yellow
pseudocyphellae, pigment deficient individuals may sometimes be found as reported for the
South American species P. hirsuta and P. vaccina (Galloway, 19860: 109). The following New
Zealand taxa have yellow pseudocyphellae: Pseudocyphellaria ardesiaca, P. aurata, P.
carpoloma, P. colensoi, P. coronata, P. crassa, P. crocata, P. degelii, P. jamesii, P. maculata, P.

PSEUDOCYPHELLARIA

15

A

Fig. 5 Pseudocyphellae on upper surface. A. Pseudocyphellaria knightii x 300. B. P. lindsayi x 400. C.
P. lividofusca x 300.

margaretiae, P. nermula, P. neglecta, P. physciospora, P. pickeringii, P. poculifera, P.
pubescens, P. rubella, and P. sericeofulva.

Lower surface: The lower surface, which may be smooth, wrinkled, ridged to bullate, is
thinly to densely tomentose, velvety-pubescent or glabrous and some species show all degrees
of hairiness from glabrous to densely and uniformly tomentose. Tomental hairs are strands of
hyphae (often in fascicles of 2-12 or more) which originate from the outermost cells of the lower
cortex and which anchor the thallus to the substrate. As Snelgar & Green (1981a) have shown,

Fig. 6 Pseudocyphellae associated with isidia-pseudoisidia on upper surface. A. Pseudocyphellaria
argyracea x 100. B. P. wilkinsii x 100.

16 D. J. GALLOWAY

the thickness of the tomentum of the lower surface in different populations is governed by the
evaporative demand of their environments, with populations exposed to full sunlight being more
closely appressed to the substrate and having a thicker tomentum, and hence increased water
storage capacity.

Species with a consistently glabrous lower surface include P. chloroleuca, P. rufovirescens ,
and P. glabra, although occasionally patchily tomentose specimens are found in all three
species. Tomentum varies in density from velvety-pubescent (e.g. in P. haywardiorurri) to thick
and woolly or shaggy (P. crassd) and may be uniform from margins to centre, or thickly
developed centrally with a wide, glabrous, smooth or wrinkled, glossy or matt, pale to dark
brown or black marginal zone. In P. dissimilis and P. cinnamomea (occasionally also in P.
multifidd) tomentum is restricted to a narrow, central, raised, somewhat prominent midrib.

Isidia: These are terete in cross-section and are small outgrowths of the thallus and as such
have a true cortex, internal to which is a photobiont layer and a central medulla. Isidia may be
simple, fingerlike, to coralloid-branched, rarely granular- verruciform (Fig. 7) and scattered
randomly over the thallus surface, to clustered at the margins, or associated with breaks in the
thallus surface (P. chloroleuca, P. dissimilis, P. glabra), or occasionally with pseudocyphellae
( P. argyracea, P. wilkinsii) , and sometimes thickly covering the upper surface as a diffract crust.
Isidia are also sometimes present on the margins and thalline exciple of apothecia. Isidia are
vegetative propagules and as such may be easily abraded by water, wind, foraging insects, and
molluscs, and possibly by birds and other animals. On removal of isidia from the thallus, small
pits are left on the surface which, however, should not be confused with pseudocyphellae. Isidia
may become eroded-sorediate in P. argyracea and P. poculifera, or phyllidiate as in P.
colensoi, P. dissimilis, P. gretae, and P. pickeringii.

Fig. 7 Marginal isidia on upper surface. A. Pseudocyphellaria chloroleuca x 100. B. P. episticta x 100. C.
P. glabra x 100.

PSEUDOCYPHELLARIA 17

Isidia are formed in P. argyracea, P. chloroleuca, P. colensol, P. dissimilis , P. glabra, P.
gretae, P. pickeringii, P. poculifera, and P. wilkinsii.

Pseudoisidia: These are small, delicate, terete, style-formed, simple to coralloid structures,
often somewhat darkened and eroding with age, usually 1 mm tall or less and to 0-1 mm diam.,
often developed at margins of pseudocyphellae (Fig. 6A) on the upper surface (in P. argyracea),
or at margins of clustered to erose soralia. In section, pseudoisidia are continuously thinly
corticate at first, the entire structure being covered by 1-2 rows of pigmented cortical cells. This
cortical covering is generally thinner then the normal cortex elsewhere in the thallus, and is
readily abraded at the base of the pseudoisidia on aging, with the formation of efflorescent
hyphal masses intermixed with intact tips of pseudoisidia. A feature of pseudoisidia is their
ability to reform, at least in part, a new cortex in sorediate areas. Indeed, pseudoisidia often start
off as soredia and then become secondarily, partially thinly corticate.

Unlike true isidia, pseudoisidia in Pseudocyphellaria appear not to contain photobiont cells
below the thinly developed cortical layer. Pseudoisidia are most commonly developed in P.
argyracea and P. poculifera, and are also occasionally found associated with soralia in P.
ardesiaca, P. bartlettii, P. crocata, P. granulata, P. haywardiorum, and P. intricata. A detailed
discussion of pseudoisidia in several other lichen genera is given in Walker (1985: 11).

Phyllidia: Phyllidia (Fig. 8A-D) are thallus outgrowths similar to isidia and like them, organs
of vegetative dispersal. They are distinctly foliose, flattened, dorsiventral structures, usually
constricted at the base, simple, expanded at tips, to subcoralloid-palmate or pectinate,
squamiform to richly divided, or straplike. As in the parent thallus they have a distinct upper
surface with photobiont layer, and a pale lower surface often with rudimentary tomentum
and pseudocyphellae. In two species (P. fimbriata, P. fimbriatoides) phyllidia have minute,

Fig. 8 Phyllidia on upper surface. A. Pseudocyphellaria corbettii x 25. B. P. coronata x 50. C. P.
fimbriata x 100. D. P. montagnei x 1000 (cross section).

18 D. J. GALLOWAY

marginal, glistening hairs (Fig. 8C) while those of P. gretae are densely and uniformly
tomentose. Phyllidia often cluster at lobe margins but are also found laminally especially
along breaks in the thallus; they are also developed on the thalline exciple of P. montagnei,
P. colensoi, and P. coronata (Fig. 8B). Phyllidia are known from P. corbettii, P. colensoi, P.
coronata, P. dissimilis, P. episticta, P. fimbriata, P. fimbriatoides , P. gretae, P. montagnei,
P. multifida, P. nermula, P. neglecta, and P. wilkinsii.

Soredia: Soredia are present in 10 species: P. ardesiaca, P. argyracea, P. aurata, P. bartlettii,
P. crocata, P. granulata, P. haywardiorum, P. intricata, P. poculifera, and P. rubella, and are
developed on the lobe margins and/or scattered over the upper surface in erumpent, pustular to
erose, rounded or irregular to linear soralia. Marginal soralia may be punctiform to
linear, or distinctly labriform and eroding areas of the lower surface (in P. aurata). Laminal
soralia are punctiform, pulverulent to erose, 0-1-3 mm diam., round to irregular, solitary to
confluent, sometimes forming reticulate patterns on thallus ridges (e.g. in P. crocata, P.
granulata, P. intricata) , or obscuring large areas of the upper surface ( P. bartlettii, P. granulata) .
Soredia are yellow, white, olive-greenish to somewhat blackened, farinose to coarsely granular,
gnarled-glomerulate (P. ardesiaca), to pseudoisidiate (P. aurata, P. crocata, P. intricata).
Occasionally in P. aurata and P. crocata (especially from humid habitats on Leptospermum)
development of minute, corticate, cochleate lobes from granular soredia may be seen.

Of the 10 sorediate species present in New Zealand all, except P. ardesiaca, P. argyracea, and
P. bartlettii, are known fertile and are not sorediate clones or forms of a non-sorediate fertile
parent species (for discussion of sorediate clones see Tehler, 1982). Pseudocyphellaria granulata
is the sorediate counterpart of P. faveolata, the two taxa constituting a species pair and it is
possible that P. ardesiaca is the sorediate counterpart of P. jamesii.

Ascomata: All species studied except P. allanii, P. ardesiaca, P. bartlettii, P. jamesii, P.
margaretiae, P. nermula, and P. sericeofulva are known fertile, although as a general rule species
with soredia only rarely produce ascomata. Two main types of ascomata are found in Pseudo-
cyphellaria: (1) Sessile, laminal, or marginal apothecia with a well-developed and often
scabrid-areolate-verrucose excipulum proprium lacking photobiont cells and often appearing
translucent when wet, characteristic of P. billardierei, P. carpoioma, P. chloroleuca, P. cinna-
momea, P. coriacea, P. crassa, P. degelii, P. dissimilis, P. episticta, P. faveolata, P.fimbriata, P.
fimbriatoides, P. glabra, P. granulata, P. haywardiorum, P. homoeophylla, P. intricata,
P. knightii, P. lindsayi, P. lividofusca, P. maculata, P. multifida, P. murrayi, P. neglecta,
P. pickeringii, P. rufovirescens , and P. wilkinsii, (2) Pedicellate marginal or submarginal
apothecia having a smooth to verrucose-areolate to tomentose excipulum thallinum contain-
ing photobiont cells, found in P. aurata, P. colensoi, P. coronata, P. durietzii, P. gretae, P.
hookeri, P. montagnei, P. physciospora, P. poculifera, P. pubescens, and P. rubella.

Apothecia are nodular-papillate at first with prominent, smooth to verrucose-scabrid
margins (pale whitish to red-brown) and frequently a plug of sterile tissue or a thick membrane
obscures the disc in early stages of its development (Fig. 9C). The disc is often deeply
cupuliform, concave at first, and expands at maturity to become plane or convex-undulate; the
surface is matt or shining, pale yellow-brown to dark red-brown to brown-black (the colour
darker in dried specimens), smooth to roughened-papillate and in P. faveolata, P. granulata,
and P. pubescens white-pruinose (Fig. 9A,B), the disc appearing greyish or grey-blue, the
pruinosity often best developed in young fruits and disappearing with age. Margins may be
entire, crenate, dentate-striate, isidiate, sorediate to eroded and exposing medullary hyphae,
scabrid, phyllidiate, pubescent, tomentose or excluded. The exciple is smooth, glossy, or matt
to tomentose, isidiate or phyllidiate, frequently scabrid-verrucose, pale flesh-pink, buff or
red-brown, usually translucent when wet (Fig. 9D).

Anatomy

Details of anatomical structure of thalline and apothecial tissues in Pseudocyphellaria are
discussed in some detail by Hue (1901), Magnusson (1940), and Galloway (19860). Thalli of all

PSEUDOCYPHELLARIA

Fig. 9 Ascomatal structures. A. Pseudocyphellaria granulata; young, hemiangiocarpic (right) and mature
(left) ascomata with pruinose discs x 30. B. P. granulata; surface of disc with pruina and ascospores x
500. C. P. pickeringii; young, hemiangiocarpic (right) and mature (left) ascomata x 30. D. P. degelii;
excipular tissue x 500.

species of Pseudocyphellaria are heteromerous and between 130-450 urn in thickness, with
well-developed specimens (e.g. P. coriacea) to 750 um. Thallus thickness varies considerably
within species (particularly in species with well-developed faveolae and interconnecting
ridges) and is mainly a consequence of the variation in thickness of the medulla and photobiont
layer (and to a lesser extent of the upper cortex) dependent on thallus age, and microecological
conditions obtaining at the thallus surface. All species have an upper and lower cortex, generally
of similar anatomical structure (Fig. 10), though the lower cortex (occasionally also the upper
cortex) is normally obscured by the development of tomental hairs from the outermost layer of
cortical cells.

Upper cortex: The upper cortex in all species is a paraplectenchyma (pseudoparenchyma) of
anticlinally arranged, round to irregular, isodiametric, angularly thickened cells (4-5-)7-
15-5(-18) um diam., with outermost cells rather smaller, compressed, and with somewhat
thicker walls than the larger, more loosely arranged cells adjacent to the photobiont layer. Cell
wall thickness is within the range 1-5-3-5 urn, with lumina round to irregular 2-5-4-4(-ll) um
diam. Cortical cells vary in colour from hyaline to pale straw-yellow to orange-brown or
red-brown, the colour being most intense at the upper surface and palest close to the photobiont
layer. Thickness of the cortex varies from 18-102 um, but within each species the variation is
much less, 4-12(-32) um. Scanning electron micrographs of the upper surface show the outlines
of individual hyphae comprising the underlying tissue, and occasional small, rather irregular
pores are seen (Fig. 11). In certain species tomental hairs may develop from cells of the upper
cortex (e.g. in P. gretae, P. pubescens) or the upper cortex is roughened-uneven, appearing
areolate-scabrid (Fig. 1A-D).

D. J. GALLOWAY

Fig. 10 Cross sections of thalli showing upper and lower cortices, medullary tissue, and tomentum of
lower surface. A. Pseudocyphellaria episticta x 200. B. P. fimbriatoides, lobe margin x 800. C. P.
intricata x 200. D. P. nermula x 200.

Photobiont layer: The photobiont layer lies between the upper cortex and the medulla and
varies considerably in thickness, between 11-140 um, with a majority of taxa having a
photobiont layer from 25-50 urn thick. Hyphae of similar structure to those found in the medulla
(though sometimes rather thicker), ramify throughout the photobiont cells. Species with a green
photobiont have the photobiont cells arranged in a continuous layer, while taxa with a
cyanobiont may have a continuous layer of photosynthetic cells, or the cyanobiont may be
present in discrete clusters or clumps with bands of hyphae separating the photosynthetic tissue.
Such discontinuities in the cyanobiont layer are visible macroscopically (x 10 lens) as maculae.
Thickness of the photobiont layer may vary within species and in certain taxa may even be
environmentally controlled; Snelgar & Green (1981a), for example, recording that populations
of P. dissimilis from sunny situations had a thicker cyanobiont layer (50 4 um) than
semi-shaded (48 1 urn), and deeply shaded (38 1 um) populations of the same species.

Photobionts in Pseudocyphellaria are either green algae or cyanobacteria. The cyanobiont in
all cases appears to be Nostoc, both as primary photosynthetic symbiont and in cephalodia,
while green photobionts in the genus are Chlorella-like (E. Tschermak-Woess, pers. comm.) or
Dictyochloropsis (Tschermak-Woess, 1984). The photobiont in New Zealand populations of
Pseudocyphellaria aurata is Dictyochloropsis symbiontica var. symbiontica, while other species
of Pseudocyophellaria examined (P. fimbriata, P. gretae, P. homoeophylld) have an undescribed
species of Dictyochloropsis as symbiont (Tschermak-Woess, 1984).

Green photobionts and cyanobionts are readily distinguished macroscopically (x 10 lens) in
dried and herbarium material. Green photobionts appear either bright green, greenish-yellow
or i colourless when the upper cortex is dissected from the underlying medulla to expose the
photobiont layer (seen also when the thallus is cracked or torn), whereas cyanobionts always

PSEUDOCYPHELLARIA

Fig. 11 Pored upper surface. A. Pseudocyphellaria ardesiaca x 1000. B. P. chloroleuca x 1000. C. P.
corbettii x 1000. D. P.fimbriata x 1000.

appear as a black layer, even in the oldest herbarium specimens and seem never to be
decolourized on long storage. Twenty-eight species of Pseu docyphellaria in New Zealand have a
green photobiont and 20 species have Nostoc cyanobionts.

Cephalodia: Cephalodia occur internally in all taxa having a green primary photobiont and
may be visible as hemispherical swellings of the upper or, more commonly, the lower surface.
In all cases the secondary photobiont is Nostoc which produces heterocysts capable of fixing
atmospheric nitrogen (Green et al., 1980). Cephalodia develop in the medulla just below the
photobiont layer (James & Henssen, 1976); a detailed investigation of the processes of
cyanobiont capture and subsequent cephalodial development in New Zealand species of
Pseudocyphellaria has not yet been made, although data is available for some South American
taxa (Renner, 1980). Cephalodia in Pseudocyphellaria are widely reported in earlier literature
(Nylander, 18600, 18606, 1867, 1868c, 1877, 18886; Lindsay, 1867; Forssell, 1883; Stirton, 1900;
Hue, 1901). Renner (19820, 19826) developed a cryptophotometric method for localizing
secondary metabolites (depsides, depsidones, dibenzofurans, etc.) in internal cephalodia (e.g.
in P. faveolata), and found such metabolites only in the outer enclosing envelope, the inner
fungal tissue of the cephalodium being without detectable amounts of secondary metabolites.
Earlier, Jordan (1972) recorded differences in chemistry between the vegetative thallus and
external cephalodia in Lobaria amplissima.

According to Renner (19826) the absence of secondary metabolites from the central parts of
internal cephalodia in the Peltigerineae may be the result of a locally different mycobiont
metabolism, such a difference possibly being connected with an adaptation of the mycobiont
towards uptake and transformation of reduced nitrogen compounds (e.g. ammonium ions)
released by nitrogen-fixing heterocysts. Since the rate of nitrogen fixation in Pseudocyphellaria
cephalodia is high (Green et al., 1980), and carbon dioxide assimilation is only of secondary

22 D. J. GALLOWAY

importance in supplying carbon skeletons to the mycobiont, it seems likely that cephalodial
mycobionts are exclusively supplied with reduced nitrogen compounds such as ammonia.
Complete uptake of fixed nitrogen may be important in avoiding nitrogen-limiting conditions for
its fixation, e.g. by a repression of enzyme synthesis in Nostoc heterocysts (Renner, 19826).

The formation of cephalodia is analogous to the formation of photosymbiodemes (see below),
except that in the latter case capture of a green photobiont by a cyanobiont-containing species
results in the formation of the external photosymbiodeme.

Medulla: The medulla is composed of loosely to compactly interwoven, unorientated
hyphae 2-5 um diam. In some species (e.g. P. crocata, P. faveolata) moniliform hyphae are
produced in the medulla (see Malme, 1899; Hue, 1901; Magnusson, 1940), such hyphae being
similar to the knobbly hyphae seen in the medulla of species of Leioderma (Galloway &
J0rgensen, 1987). In species with a yellow medulla, the hyphae are thickly encrusted with yellow
crystals (see Green et al., 1985:71, Fig. 7). The thickness of the medulla varies considerably
between species, within individuals of the same species, and even within a single thallus,
depending on the age and size of the thallus lobe, and on the particular microhabitat and
microclimate conditions prevailing at a given part of the thallus.

Lower cortex: The lower cortex is a pseudoparenchymatous tissue very similar in structure to
that of the upper cortex, although it is usually thinner than the upper cortex. Cells are colourless
to yellow-brown or red-brown, with the outer-most cells being most strongly pigmented and the
inner cell layers paler. The red-brown cortical pigment of P. granulata turns blue-green in K, a
reaction not seen in any other species. Cells of the lower cortex are 2-5-15-5 um diam., with
walls 1-5-2-5 um thick. The thickness of the lower cortex ranges from 9-60 um, but within each
species the variation observed within individual thalli is 4-14 um.

Tomentum: Tomental hairs develop from the outermost cells of the lower cortex (Fig. 10) and
also occasionally from those of the upper cortex (Fig. 1A, B). Hyphae are mainly simple, usually
thick-walled, and with a prominent central canal, rarely 2-3-branched at apices or occasionally
with short, lateral branchlets [analogous to structures seen on the tomental hairs of some species
oiSticta (see Harris, 1984)], septate (often with constrictions at septa, and rarely moniliform),
colourless, pale straw-yellow, yellow-brown to red-brown, solitary or more usually in conglutin-
ate fascicles, 2-10(-20)-together, from 4-5-9 urn diam. and (30-)45-230(-450) um long.

Ascomata: Ascomata in Pseudocyphellaria are hemiangiocarpic in development, arising from
a compact primordium which develops in the lower part of the photobiont layer. The ontogeny
of ascomata in Pseudocyphellaria is similar to that seen in Lobaria (Letrouit-Galinou, 1971) and
is discussed in detail in Keuck (1977) and Henssen (1981). An important, and also the earliest,
account of ascomatal development in Pseudocyphellaria (in P. anthraspis) is given by Sturgis
(1890).

Exciple: The exciple refers to the whole supporting structure of thecial tissues in ascomata and
may or may not contain photobiont cells. Species with a proper exciple (excipulum proprium),
lacking photobiont cells have sessile apothecia and include P. billardierei, P. carpoloma, P.
chloroleuca, P. cinnamomea, P. coriacea, P. crassa, P. crocata, P. degelii, P. dissimilis, P.
episticta, P. faveolata, P. fimbriata, P. fimbriatoides , P. glabra, P. granulata, P. haywardiorum,
P. homoeophylla, P. intricata, P. knightii, P. lindsayi, P. lividofusca, P. maculata, P. multifida,
P. murrayi, P. neglecta, P. pickeringii, P. rufovirescens , and P. wilkinsii. In these taxa the
proper exciple is a colourless to pale straw-yellow to yellow-brown cellular tissue 35-140(-320)
um thick, composed of rows of anticlinally arranged, radiating, isodiametric, round to oblong
to irregular cells, 4-5-13-5(-18) um diam., with walls 1-5-4-5 um thick, lumina round to
irregular (l-5-)3-7(-9) um diam. The outer parts of the proper exciple are often split into
pyramidal or wedge-shaped masses (Fig. 9D) giving a characteristic scabrid-verrucose macro-
scopic appearance (x 10 lens). Occasionally colourless, simple, septate hairs develop from outer
excipular cells at the base of the apothecia.
Species with a thalline exciple (excipulum thallinum) contain photobiont cells in the exciple,

PSEUDOCYPHELLARIA 23

the arrangement of tissues being similar to that found in the vegetative thallus. In addition,
apothecia with a thalline exciple are markedly stipitate. Species with stipitate fruits include
P. aurata, P. colensoi, P. coronata, P. durietzii, P. gretae, P. hookeri, P. montagnei,
P. physciospora, P. poculifera, P. pubescens, and P. rubella.

Hypothecium: The hypothecium is a compact zone of densely ramifying hyphae below the
hamathecium, usually pigmented (pale straw-yellow, yellow-brown, orange-brown to red-
brown), opaque, (30-)45-90(-145) um thick, mostly K- but occasionally becoming blue-green
in K (P. fimbriata, P. fimbriatoides , P. pickeringii) .

Thecium: The thecium is colourless to pale straw-yellow or pale brownish, to dilute orange-
brown, occasionally somewhat granular, 1+ blue, 45-200 um tall (most fertile species have a
thecium height of 80-150 um), measured from the upper edge of the hypothecium to the lower
edge of the epithecium, at the centre of the thecium. The thecium is in most cases K- but in the
lower part of P. faveolata and in P. fimbriatoides it stains greenish in K.

Hamathecium: The hamathecium is composed of numerous, markedly conglutinate para-
physes, separating in K, simple, straight, septate, 1-5-2-5 um diam., swollen, capitate at
apices (4-5-5 um), sometimes moniliform, with 3-5 apical vacuolate cells visible.

Epithecium: The epithecium, (6-5-)10-15(-22-5) um thick, consists of a pigmented,
granular gel external to and including the tips of the paraphyses, varying in colour from grey-
ish olivaceous to yellow-brown or red-brown. In a number of species a distinct colour change
in the epithecium pigments is noted when sections are irrigated with K. The epithecium of P.
billardierei turns dark greenish black, those of P. dureitzii, P. faveolata, P. granulata, P.
hookeri, P. montagnei, deep violet-purple, and those of P. episticta and P. lindsayi rose-pink.
The remaining taxa show either no change in epithecial pigments or else the epithecium slowly
decolorizes. The colour change of epithecial pigments of fertile species of Pseudocyphellaria
was earlier noted by Hue (1901), Magnusson (1940), Imshaug (1977), and Galloway (1986a).
See also White & James (1985: 13-15).

Asci: Asci in Pseudocyphellaria are always shorter than paraphyses, clavate with a tapering
foot, to cylindrical, bitunicate, I + blue, with thickened apices and an apical ring staining in
Lugol's iodine, dehiscence semifissitunicate, the ring everting (Keuck, 1977; Honegger, 1978;
Hawksworth et al. , 1983) , and producing eight ascospores, uniseriately or biseriately arranged in
the ascus.

Ascospores: Ascospores in Pseudocyphellaria are ellipsoid to fusiform-ellipsoid, rarely
acicular-fusiform, with broadly rounded or narrowly tapering apices. Immature spores are
simple, while mature spores are at first 1-septate, possibly the normal or general condition of the
mature spore as suggested by Lindsay (1869:495), and at length become 3-septate. Occasionally
over-mature spores with five or seven septa are seen. A number of species have 1-septate spores
with a conspicuously thickened septum traversed by a thin canal joining the two polar locules
(the so-called physcioid spores). Such spores, first noted by Nylander (18606) are found in P.
billardierei, P. carpoloma, P. durietzii, and together with 3-septate spores in P. crassa, P. crocata,
P. faveolata, P. granulata, P. gretae, P. hookeri, P. maculata, P. montagnei, P. neglecta, and P.
pubescens. Although called polarilocular in some earlier accounts (e.g. in Galloway, 19856)
these spores with thickened septa are not at all similar to true polarilocular spores as found in
Caloplaca for example, but are reminiscent of those otPhyscia (see Moberg, 1977) as claimed by
Nylander and later authors. In a few species (e.g. P. degelii, P. pickeringii) ascospores are
colourless even at maturity; however, normal spore colour is olivaceous to yellow-brown or dark
brown at maturity with quite often a range of colours seen in sections of thecium or even in a
single ascus.

The importance of spores of Pseudocyphellaria in the statistical analyses of local populations
of this genus is emphasised by Imshaug (1977).

Anamorphs: Adequate description of both morphology and anatomy of anamorphs (conidial

24 D. J. GALLOWAY

states) of lichen-forming fungi is increasingly required in modern taxonomic revisions and for
discussion of modern concepts and terminology followed in this account see Vobis (1980) , Vobis
& Hawksworth (1981), and Coppins (1983).

Anamorphs of Pseudocyphellaria were first described by Lindsay (1859: 191-205; 1869:
494-495) under the genera Ricasolia, Sticta, and Stictina. Lindsay's detailed observations on
conidiomata (pycnidia), conidiogenous cells, and conidia were supported by excellent
hand-coloured illustrations and are as pertinent today as they were over 100 years ago.
He drew attention to the uniformity of what he called spermatia (conidia) and sterigmata
(conidiophores) throughout each of these genera, a conclusion supported by Vainio (1890) for
Pseudocyphellaria, and by Hue (1901) for Sticta sens. lat. (including taxa referable to Lobaria,
Pseudocyphellaria, and Sticta).

Conidiomata: Conidiomata develop from primordia in the photobiont layer. They are
pycnidial (cited as spermogones by Lindsay and some later authors), immersed in the thallus,
globose to ovoid in shape, 0-l-0-5(-0-8) mm diam., often visible on both upper and lower
surfaces as shallow, hemispherical swellings, but most generally indicated by a dark brown,
red-brown or black ostiole, 0-05-0-1 mm diam., and which may be plane, slightly elevated, or
punctate-depressed. The ostiole is usually surrounded by a narrow, white or colourless halo
(x 10 lens). Mature conidiomata are filled with long, simple to occasionally branched con-
idiophores, (20)40-50(-70) um long, comprising rows of intercalary conidiogenous cells.
Conidiomata in Pseudocyphellaria are of the Lobaria type (Vobis & Hawksworth 1981:
252-253). They may be frequent to sparse, even on different parts of the same thallus, and
appear to be most common at lobe margins, on thalline ridges of faveolate species or towards the
apices of non-faveolate species, often arranged in lines at margins and on ridges or randomly
scattered, to crowded-confluent. Quite often conidiomata erode and fall out of the thallus,
leaving empty, gaping pits. Lindsay (1859: 191-192) first described this phenomenon 'With age it
[the ostiole] expands, and when the nucleus or body of the spermogone falls out, as frequently
happens, it may become saucer-shaped, with ragged, generally thickened and dark-coloured
margins. In this state, the ostioles of old spermogones sometimes have the aspect of black rings
or discs, as occasionally in S. aurata. However large or small the ostiole or external protuberance
of the spermogone, its body is almost in all cases a large white kernel of a dense horny tissue,
which becomes gelatinous and semipellucid in water. From its density, it can easily be
enucleated with the point of a needle.' Conidiomata in Pseudocyphellaria appear to be most
frequently developed in fertile species and are rare or absent in taxa having copious develop-
ment of asexual propagules, e.g. P. ardesiaca, P. argyracea, P. aurata, P. bartlettii, P. crocata, P.
granulata, P. gretae, P. haywardiorum, P. intricata, P. neglecta, and P. nermula.

Conidiogenous cells: The conidiogenous cells of Pseudocyphellaria (and also those of Lobaria
and Sticta) are all very similar in morphology and size, a fact noted first by Lindsay (1859: pi. X;
1869: tab. 60) and by Hue (1901). Cells are colourless, cuboid to somewhat irregular,
2-5-5 um diam. They are phialidic and type VII in the arrangement of Vobis & Hawksworth
(1981:257).

Conidia: These are very uniform throughout Pseudocyphellaria, both in New Zealand and in
other parts of its range, and also in Lobaria and Sticta. Conidia are colourless, short, bacilliform
to slightly dumbell-shaped (the two poles being only slightly swollen and usually more intensely
birefringent than the remainder of the cell), (2-)3-5(-7) x 0-5-1 urn.

Photosymbiodemes

In Pseudocyphellaria (also in Lobaria, Nephroma, Peltigera, Solorina, and Sticta) distinctive yet
obviously closely related pairs of taxa exist, the major apparent difference between them being
in the nature of the photobiont present. Recently, in Pseudocyphellaria particularly, joined
thalli comprising a single mycobiont with two different photobionts were recorded (James &
Henssen, 1976; Renner & Galloway, 1982; Galloway, 1985ft), the green photobiont- and the
cyanobiont-containing demes being given separate names for their free-living representatives

PSEUDOCYPHELLARIA 25

which are usually in this genus subtly distinct when growing independently. The term photo-
symbiodeme is used for such combined thalli in preference to phycosymbiodeme as originally
proposed (Renner & Galloway, 1982), being a more neutral term and not carrying with it any
specific reference to algae since blue-green algae are now increasingly accepted as
cyanobacteria.

In New Zealand both free-living green photobiont- and cyanobiont-containing species may
exist in similar habitats alongside examples of joined thalli; P. allanii-P. coriacea, P. hookeri-P.
durietzii, P. knightii-P. lividofusca, P. murrayi-P. rufovirescens , P. margaretiae-P . pubescens are
presently known (the cyanosymbiodeme preceding the chlorosymbiodeme) and there seems, on
available evidence, to be no particular microclimatic factor promoting formation of photo-
symbiodemes, though this has not yet been closely studied in the field. Presently, joined thalli
are known from moist, humid habitats in moderate shade, either close to sheltered streams, or in
deep gorges, or in areas of wet forest on slopes often shrouded in mist or cloud. As both
chloro- and cyanosymbiodemes appear to be capable of independent existence and have
characters which permit their taxonomic separation, they are given independent species names.
As James & Henssen (1976) point out, the International Code of Botanical Nomenclature makes
no allowance for the existence of joined thalli differing only in the nature of the photobiont
present.

Renner & Galloway (1982) have shown that the development of photosymbiodemes follows a
uniform pattern in Pseudocyphellaria, and that chlorosymbiodemes are produced secondarily
by the mycobiont of the cyanosymbiodeme. Whenever a chlorosymbiodeme is formed it is of
secondary origin and is characterized by an heteromerous arrangement of tissues. A comparison
between development of internal cephalodia (see above) and of chlorosymbiodeme formation
shows certain parallels. Formation of cephalodia may be regarded as the secondary lichenisation
of a cyanobacterium (Nostoc), whereas the formation of a chlorosymbiodeme involves capture
of a green photobiont by a species having a cyanobiont as its primary photobiont.

Of the known photosymbiodemes in New Zealand, only two cyanosymbiodemes are known in
the fertile state (P. hookeri and P. murrayi}. In all the cases mentioned above, the chlorosym-
biodemes and cyanosymbiodemes are mirror images of each other morphologically speaking,
neither symbiodeme produces asexual propagules, nor is there any major chemical difference
between chlorosymbiodeme and cyanosymbiodeme. Since photosymbiodemes are known only
from forest or scrub vegetation in humid habitats, photosymbiodeme formation may be a
mechanism which is evolving to allow mycobionts to exploit a wider range of macrohabitat
and/or microclimate conditions (see also James & Henssen, 1976).

Lichenicolous fungi

On many New Zealand species of Pseudocyphellaria distinctive black or blackish brown spots,
stains or apothecia-like galls are produced on the upper (rarely on the lower) surface, the result
of attack by one or more genera of lichenicolous fungi (Hawksworth, 1982b). Such parasitic
fungi were first recorded from Otago collections of Pseudocyphellaria faveolata, P. granulata,
and P. rubella by Lindsay (1867: 448-453 ; 1869: 496) under the name Celidium dubium. Peculiar
apothecia-like structures on what is now known as Pseudocyphellaria glabra led Fee (1824-
1825), who first investigated this Southern Hemisphere lichen, to make it the type of a new
genus, Delisea Fee. However, since Delisea was first described in 1819 by Lamouroux for a genus
of red algae, Fee renamed his new lichen Plectocarpon (Fee, 1824-1825; 1828). Hawksworth &
Galloway (1984) showed the lichenicolous fungus Lichenomyces Trevisan, to be a synonym of
Plectocarpon, and that the fungus causing apothecia-like galls on Pseudocyphellaria glabra must
be called Plectocarpon pseudosticta. It is likely that several unidentified species of Plectocarpon
are associated with New Zealand species of Pseudocyphellaria, and these are presently under
investigation. Pseudocyphellaria is a member of the Peltigerales, of which a number of genera
(e.g. Lobaria, Nephroma, and especially Peltigera) are rich in lichenicolous fungi (Hawksworth,
1982fl). A study of the lichenicolous fungi occurring in Southern Hemisphere species of
Pseudocyphellaria, together with comparative chemical studies, should reveal something of the

26 D. J. GALLOWAY

past and present relationships within the genus as it is at present represented in cool-temperate
southern regions.

Chemistry

In lichens, primary metabolites such as lipids and carbohydrates are important to the lichen
association both metabolically and structurally. Carbohydrates, for example, occur widely in
lichens and probably constitute the major part of the dry weight of the thallus in most lichens.
Lichen carbohydrates can be conveniently divided into the following major groups: (1)
polysaccharides, (2) sugars and oligosaccharides, and (3) polyhydric alcohols (for a recent
review of relevant literature see Elix etal., 1984). Polyhydric alcohols (polyols) are implicated in
the transfer of carbon skeletons (probably as their phosphorylated derivatives) from autotrophic
to heterotrophic partners in a variety of symbiotic associations including the lichens (Smith,
1975, 1981; Hill, 1976). In species of Pseudocyphellaria in New Zealand, the polyols D-
arabinitol and mannitol are the major water-soluble compounds present and often occur in
considerable abundance (unpublished observations).

However it is the complex array of secondary metabolites produced by lichens which are of the
greatest utility in chemosystematic studies. Chemotaxonomy has a wide application in contem-
porary lichenology (Hawksworth, 1976) and is especially useful in helping to separate taxa at the
species level in Pseudocyphellaria (Wilkins & James, 1979; Galloway et al., 19836; Galloway,
1986a). Substances from all three major pathways of secondary metabolism in lichens are found
in New Zealand species viz., the acetate-polymalonate pathway, the mevalonic acid pathway,
and the shikimic acid pathway (see Culberson, 1969; 1970; Culberson & Culberson, 1970;
Culberson et al, 1977; Santesson, 1973; Huneck, 1974; Mosbach, 1974; Elix et al., 1984;
Huneck, 19846).

The richness and diversity of secondary metabolites found in species of Pseudocyphellaria
(Wilkins & James, 1979; Galloway et al., 19836), and especially of the four series of tri-
terpenoids discovered in New Zealand species (see below) has proved to be of great utility in
the unravelling of species taxonomy. For instance, the very closely related faveolate species
P. billardierei, P. carpoloma, P. faveolata, and P. physciospora all have a closely similar
morphology and may often be difficult to distinguish on morphological grounds alone. How-
ever, they each have a distinctive chemistry and are readily separable chemically. Thus chemical
characters reinforce differential morphological characters in closely related taxa and frequently
allow the accurate determination of atypical material which otherwise might be difficult to
differentiate on morphological grounds alone.

Within the family Lobariaceae there is a dramatic increase in the number and complexity of
secondary metabolites as one moves from Sticta, through Lobaria, to Pseudocyphellaria.
Species of Sticta have few or no secondary products soluble in acetone, instead they produce
large quantities of water-soluble simple carbohydrates and methylamine; species of Lobaria
have an intermediate range of secondary compounds, while species in Pseudocyphellaria have
the most diverse secondary chemistry of the family, and one of the most complex chemistries of
any lichen genus known. This chemical diversity is persuasive evidence for considering
Pseudocyphellaria to be taxonomically remote from Sticta and more closely related to Lobaria.
Indeed, there are probably good reasons for including Sticta in a family of its own (Stictaceae
Zahlbr.) and keeping Lobaria and Pseudocyphellaria together in the family Lobariaceae Chev.

Compounds formed in each of the three pathways of secondary metabolism in Pseudo-
cyphellaria are reviewed briefly below.

A cetate-polymalonate path way

The most common phenolic acid units synthesized via the acetate-polymalonate pathway are of
two kinds: (1) orcinol-type compounds based on the skeleton

PSEUDOCYPHELLARIA 27

and (3-orcinol-type compounds which have an extra Ci substituent at the 3-position
CH -

oH ] (Culberson, 1969).

A. Orcinol series
i) Monocyclic derivatives: methyl orsellinate

j, 4-0-methyl-orsellinate

, ofi^^OH

01 J and the parent orsellinic acid [ fof COOH ] isolated from Pseudocyphellaria
crocata by preparative two-way TLC (Maass, 19756).
ii) para-depsides: The major structural variations seen in the orcinol /?0ra-depsides

R, O

[ ..jQjLlOjC^ 1 ] are the length of the polyketide derived side chains (R 1 , R 2 ), the degree

of oxidation of these side chains (CH 2 COR or CH 2 CH 2 R), and the degree of methylation
of the phenolic and carboxyl groups (R 3 , R 4 , R 5 , R 6 = CH 3 or H) (Elix et al. , 1984).

1. Evernicacid

f r^r TO-Or

First isolated from P. crocata (Maass, 19756)

2. Methyl evernate

3. Lecanoric acid

4. Methyl lecanorate

Isolated from P. crocata (Maass, 19756) and from
Lobaria linita (Maass, 1975c), and Peltigera
aphthosa (Maass, 19750).

First isolated from P. crocata (Maass, 19756)
First isolated from P. crocata (Maass, 19756).

iii) tridepsides: Besides variation in the degree of methylation of the phenolic and carboxyl
groups (R 1 , R 2 , R 3 , R 4 , R 5 = H or CH 3 ), the orcinol tridepsides commonly have an
additional OH (or OCH 3 ) group in the A-ring (Elix et al., 1984).

1. Gyrophoric acid

2. Methyl gyrophorate

3 . 4-0-methyl gyrophorate

4. Tenuiorin

5. 2-0-acetyltenuiorin

First isolated from P. crocata
(Maass, 19756) and from P.
sulphurea (as P. quercifolia} by
Maass (1975d).

Isolated from P. crocata, P. sul-
phurea, and Lobaria linita by Maass
(19756, 1975c, 1975d).
First isolated in Pseudocyphellaria
from P. crocata and P. sulphurea
(Maass, 19756, 1975d).
First isolated in P. crocata (Corbett
& Gumming, 1971; Maass, 19756).

First isolated from Pseudocy-
phellaria neglecta by Bryan & Elix
(1976). Also from P. dissimilis, Pel-
tigera dolichorhiza, and P. spuria.

28

6. 2' -0-methyltenuiorin
1. 2" -0-methyltenuiorin
8. 2',2"-Di-O-methyltenuiorin

B. $-Orcinol Series

i) /?0r0-depsides

1. Atranorin

2. Chloratranorin

ii) Fully substituted depsides
1. Pseudocyphellarin A CH , f"

D. J. GALLOWAY

,OCH, _Av ,COOCH,

ror rcj

Isolated from Pseudocyphellaria
faveolata (Elix & Lajide, 1981).

Isolated from Pseudocyphellaria
jo faveolata (Elix & Lajide, 1981).

Also in P. billardierei (Elix, 1986).
, Isolated from Pseudocyphellaria
faveolata (Elix & Lajide, 1981).
Also in P. billardierei (Elix, 1986) .

First isolated from Pseudocyphellaria crocata
(Maass, 19756).

First isolated from Pseudocyphellaria crocata
(Maass, 19750).

Isolated from Pseudocyphellaria endochrysa
gX Jo_ (Huneck, 19840; Huneck ef 0/., 1984). See also

r "' Elix & Lajide (1984).

2. Pseudocyphellarin B CH, rjL rV OH Isolated from Pseudocyphellaria endochrysa (see

Oi loll above).

OH ctTY^ 000 "'

I CH,

iii) Depsidones

1. Methyl virensate (granulatin)

O

First isolated from Pseudocyphellaria faveolata
and P. granulata (Renner ef a/., 1978; Goh &
Wilkins, 1979).

2. Physciosporin (5-chloro methyl virensate, chlorogranulatih)

^ First isolated from Pseudocyphellaria faveolata

and P. granulata (Renner e? a/., 1978; Goh &
' Wilkins, 1979).

3. Norstictic acid

4. Salazinic acid

5. Consalazinic acid

6. Galbinicacid f- g

7. Sticticacid

PSEUDOCYPHELLARIA

29

8. Cryptostictic acid

9. Constictic acid

10. Hypos tictic acid

11. Hyposalazinic acid

12. Connorstictic acid

13. Virensicacid

14. Hypoconstictic acid

A scheme showing the interrelationships of stictic acid and related depsidones is given in Elix ef
a/. (1984: 194-195).

iv) Usnicacid s "'oc H . Usnic acid occurs as a cortical pigment in

Pseudocyphellaria corbettii, P. freycinetii, P.
glabra, and P. homoeophylla.

Shikimic acid pathway

This pathway is responsible for the synthesis of terphenylquinones and pulvinic acid (4-
Ylidenetetronic acid) derivatives. This biosynthetic route, accounting for a vast array of
products in non lichen-forming fungi, leads to only a few of the compounds known in lichens
(Culberson & Culberson, 1970). Terphenylquinones and 4-Ylidenetetronic acid derivatives are
all pigments.

i) Terphenylquinones. Polyporic acid:

Zopf (1901, 1907) obtained from Sticta orygmaea (Pseudocyphellaria coronatd) [collected by
Helms, number 1214 in Arnold's exsiccata], a brown, sparingly soluble pigment 'orygmaeic
acid', m.p.310, which gave purple salts with alkalis. Zopf thought it might be an anthraquinone.
Fifty years later, James Murray (1952) of the University of Otago (Dunedin, New Zealand),
Chemistry Department, isolated orygmaeic acid from the New Zealand lichens Pseudo-
cyphellaria colensoi and P. coronata, and showed that it was identical in structure to polyporic
acid, discovered originally in the fungus Polyporus nidulans. Although able to distinguish
chemically P. coronata and P. colensoi, Murray found some difficulty at first in separating these
taxa solely on morphological criteria, and this led to his subsequent important taxonomic studies
on New Zealand lichens. Sadly these were cut short by his tragically early death in 1961 at the age
of 37. However, his initial chemical work formed the basis of three different and independent
lines of study in New Zealand, Canada, and Germany.
The antileukaemic activity of polyporic acid was discovered by Burton & Cain (1959). In the

30 D. J. GALLOWAY

course of screening indigenous New Zealand plant products for experimental anti-cancer
activity, a preparation of the lichen P. coronata was found to prolong significantly the lifespan of
mice previously inoculated with an acute lymphocytic leukaemia. Polyporic acid isolated from
the lichen was shown to possess antileukaemic activity and to account completely for that of the
original crude extract.

The Rf 's of polyporic acid isolated from P. colensoi, on t.l.c. in three solvent systems were
recorded by Santesson (1967). Polyporic acid is known from P. colensoi, P. coronata, and P.
pickeringii (Murray, 1952; Chin etal., 1973).

ii) 4-Ylidenetetronic acids

Tetronic acid is a trivial name for 3-hydroxy-but-2-enolide . \A,

The corresponding 4-ylidene derivatives contain an additional exocyclic C-C double bond at C 4 .

Natural products incorporating a 4-ylidenetetronic ring system have been known for almost a
century, The ring system is a characteristic structural feature in the pulvinic acid group of yellow
and red pigments found in lichens and known since 1879. These compounds also occur in the
fungal families Boletaceae and Gomphidiaceae.
1. Calycin

Its structure was accurately established by Akermark (1961).
2. Calycinic acid

Isolated from P. crocata by Maass (1975 or b).
3. Pulvinic acid

4. Pulvinamide

Isolated from P. crocata by Maass (19706).
5. Pulvinic dilactone

The common pulvinic acid derivatives (pulvinic acid, pulvinic dilactone, and calycin) which
nearly always occur together were first extracted from species of Candelariella and 'Leprarid 1 in
the late 19th century [for references to pre-1950 literature see Culberson (1969)]. These pulvinic
acid derivatives are known from yellow-medulla species of Pseudocyphellaria particularly, and
from those species having yellow pseudocyphellae (Murray, 1952; Maass et al., 1964; Maass &
Neish, 1967; Huneck & Follmann 19670, 19676; Maass, 19700, 19706; Corbett & Gumming,
1971; Chin etal., 1973; Huneck etal., 1973; Maass, 19756; Wilkins & James, 1979; Galloway &
James, 1980; Galloway etal., 19836; Galloway, 19856, 19860; Corbett etal., 1987).

The yellow-medulla species of Pseudocyphellaria found most widely in New Zealand, P.
colensoi, P. coronata, P. pickeringii, are commonly used as sources of dyes by spinners and
weavers. The various yellow and orange pigments found in these lichens also attracted the
interest of chemists in the late 19th century and Zopf (1892, 1899, 1901, 1907) isolated stictaurin,
a 1:1 molecular complex of pulvinic dilactone and calycin, from a New Zealand specimen of
what was then called Sticta orygmaea (P. coronata); he also obtained the compound from P.

PSEUDOCYPHELLARIA 31

aurata, P. pickeringii (as Sticta flavicans) , P. carpoloma (his specimens of Sticta glaucolurida
were from Arnold's Exsiccata 1199, the material being collected by Charles Knight), P. crocata,
and P. gilva.

Biosynthesis of 4-Ylidenetetronic acids in P. crocata using C 14 -labelled precursors was studied
by Maass (Maass et al., 1964; Maass & Neish, 1967; Maass 19700, 19706) [see also review of
Pattenden (1978)], who showed that the basic pulvinic acid skeleton is derived via a phenylpro-
panoid precursor involving polyporic acid as a key intermediate.

Murray's work on P. coronata (Murray, 1952), also demonstrated distinctive red pigments
easily extracted in organic solvents but proving unstable on purification. This pigment complex
is not present in other related yellow-medulla species of Pseudocyphellaria and is therefore a
useful screening test for P. coronata.

Mevalonic acid pathway

The mevalonic acid pathway produces sterols and triterpenoid skeletons, many of which occur in
genera of the Peltigerineae. The exciting chapter of triterpenoid chemistry which has proved to
be of such taxonomic usefulness in Pseudocyphellaria was anticipated by Murray (1952) who
wrote 'Compound D has properties typical of a group of about 20 neutral unreactive substances
which have been isolated from lichens . . . It is a colourless, highly crystalline substance, slightly
soluble in most solvents, and optically active. It contains no methoxyl groups, shows no carbonyl
or hydroxyl group reactions, and is unaffected by acid or alkali. Despite this lack of reactivity,
the infra-red absorption spectrum has a strong carbonyl band at 1742 cm" 1 . There is no
absorption corresponding to hydroxyl groups. Although there is a weak absorption band in the
UV below 220 m(i the substance appears to be saturated and the infra-red spectrum shows no
ethylenic absorption. Compound D may be a triterpenoid, but owing to our inability to prepare
any derivatives of it, we have not yet been able to define accurately its molecular formula.'

Murray's was the first in a series of chemical studies on New Zealand lichens, primarily species
of Pseudocyphellaria, carried out by Corbett and his students at Otago University between 1966
and 1985, and by Wilkins and his collaborators at the University of Waikato. This work, one of
the most productive and taxonomically important fields in New Zealand natural product
chemistry, established the nature and identity of four series of triterpenoids based on hopane,
stictane, fernene, and lupane skeletons, and has had profound consequences on taxonomic
judgements in Nephroma, Peltigera, and especially in Pseudocyphellaria (Corbett & Young,
19660, 19666; Corbett & Smith, 1967; Corbett etal., 1968; Corbett & Smith, 1969; Corbett &
Gumming, 1971; Corbett & Heng, 1971; Corbett et al., 1972; Chin et al., 1973; Corbett &
Wilkins, 1976; Corbett et al., 1976; Corbett & Wilkins, 1977; Wilkins, 19770, 19776; Goh et al,
1978; Ronaldson & Wilkins, 1978; Holland & Wilkins, 1979; Wilkins & James, 1979; Corbett et
al., 1982; Corbett etal., 1985; Corbett et al. , 1987).

Terpenoids represent one of the largest and biologically most important classes of natural
products, exhibiting a remarkable structural and functional diversity especially in view of their
common origin from the same C 5 isopentenoid units. Although acetyl Co A is the basic building
block of all terpenoids, mevalonic acid is the branch point separating terpenoid synthesis from
other metabolic pathways. The term triterpenoid refers to a group of natural products
containing 30 carbon atoms based on six isoprenoid units (Kulshreshtha et al., 1972; Pant &
Rastogi, 1979) produced from mevalonic acid and proceeding via the cyclisation of squalene
from the intermediates geranyl pyrophosphate and farnesyl pyrophosphate in a scheme now
universally accepted (Mabry & Gill, 1979). The polymerisation of six isopentenoid pyrophos-
phate units gives the 30C triterpenoid skeleton, and once the carbon-carbon bonds between the
C 5 units are formed, they are remarkably stable. Subsequent to polymerisation and cyclisation
with the formation of the triterpenoid skeleton, inumerable rearrangements, oxidations, etc.
are possible, with the formation of thousands of structurally unique terpenoids possible.

In Pseudocyphellaria four major triterpenoid series are known to date. Fernene triterpe-
noids were recently discovered in specimens of Pseudocyphellaria aurata (A. L. Wilkins, pers.
comm.).

32

1. Hopanes

D. J. GALLOWAY

Corbett & Young, 19660, 19666; Corbett & Smith, 1967; Corbett etal., 1968; Corbett & Smith,
1969; Corbett & Gumming, 1971; Corbett & Heng, 1971; Corbett et al., 1972; Corbett &
Wilkins, 1976, 1977; Ronaldson & Wilkins, 1978; Wilkins & James, 1979.

2. Stictanes

Chin et al., 1973; Corbett et al., 1976; Wilkins, 19770, 19776; Goh et al., 1978; Holland &
Wilkins, 1979; Corbett etal., 1982.

3. Lupanes R! J_,

Corbett etal., 1985, 1987.

1. Hopane series: Hopanoid triterpenoids, which are thought to be 'primitive' phylogenetic
precursors of sterols (Ourisson et al. , 1979), were first detected in species of Pseudocyphellaria
by Corbett & Young (19660, 19666) who isolated and determined the structure of 76-acetoxy-
22-hydroxyhopane (7(3-acetoxyhopan-22-ol) (1) and 15a, 22-dihydroxyhopane (hopane-15a,
22-diol) (3) from Sticta billardierii [sic] (= Pseudocyphellaria rufovirescens) . Since then, some 12
hopane triterpenoids have been detected in white-medulla species of Pseudocyphellaria (see
above, and also Huneck & Follmann, 19670, 19676; Follmann & Huneck, 1972; Huneck etal.,
1973; Wilkins & James, 1979; Renner, 1980, 19826; Renner & Galloway, 1982; Galloway etal.,
19836; Huneck, 19846; Galloway, 19866). The recent report of these two hopanes in extracts of
Sphaerophorus scrobiculatus (Huneck & Tibell, 1985) is presumed to be because of contamina-
tion with material of Pseudocyphellaria present in the collection.

Ri

R 2

R 3

R 4

R 5

1

CH 3

H

OAc

H

H

2

CH 3

H

OH

H

H

3

CH 3

H

H

OH

H

4

CH 3

OH

OH

H

H

5

CH 3

OH

H

H

H

6

CH 3

OAc

H

H

OAc

7

CH 3

OAc

H

H

OH

8

COOH

OAc

H

H

H

9

COOH

OH

H

H

H

10

CH 3

OH

OAc

H

H

11

CH 3

OAc

OH

H

H

12

COOH

H

H

OAc

7|3-acetoxyhopan-22-ol (peltidactylin)
hopane-7(3, 22-diol
hopane-15a, 22-diol
hopane-6a, 7(3, 22-triol
hopane-6a, 22-diol (zeorin)
6a, 16|3-diacetoxyhopan-22-ol
6ct-acetoxyhopane- 16(3-22-diol
6a-axetoxy-22-hydroxyhopan-23-oicacid
6a-22-dihydroxyhopan-23-oic acid
7|3-acetoxyhopane-6a, 22-diol
6a-acetoxyhopan-7|3, 22-diol
15a-acetoxy-22-hydroxyhopan-24-oicacid

Hopanes 5-11 (above) are often found in species of Heterodermia. The hopanes zeorin,
pentadactylin, dolichorrhizin (15a-acetoxyhopan-22-ol), phlebic acid A (28-acetoxy-22-
hydroxyhopan-23-oic acid), phlebic acid B(22-hydroxyhopan-23-oic acid), and hopane-

PSEUDOCYPHELLARIA 33

15a, 22-diol are known from Northern Hemisphere, and New Zealand species of Peltigera
(Kurokawa et al., 1966; Takahashi et al., 1969, 1970; T0nsberg & Holtan-Hartwig, 1983;
Galloway, 19856; Vitikainen, 1985), and the hopanes zeorin, hopane-7|3, 22-diol, 7(3-
acetoxyhopane-22-ol, hopane-15a, 22-diol, from species of Nephroma (Galloway, 19856). Four
hopane triterpenoids were isolated from Physcia aipolia by Elix et al. (1982) and were shown to
be 20a-acetoxyhopane-6a, 22-diol, 16(3-acetoxyhopane-6a, 22-diol, 6a-acetoxy hopane- 16(3,
22-diol, and 6a, 16|3-diacetoxyhopan-22-ol.

2. Stictane series: A chair, boat, chair, chair, chair conformational sequence was proposed for
stictane, the parent of 10 triterpenoids isolated by Chin et al. (1973) using column chromato-
graphy and multiple, preparative t.l.c. from the neutral fraction of hexane extractives of
yellow-medulla species of Pseudocyphellaria from New Zealand (P. colensoi, P. coronata, and
P. pickeringii).

stictane-2a, 30, 22a-triol
2a, 3(3, 22a-triacetoxystictane
2a, 3|3-diacetoxystictan-22a-ol
2a-acetoxystictane-3(3, 22a-diol
3(3-acetoxystictane-2a, 22a-diol
2a, 3(3-diacetoxystictan-22-one
stictane-3|3, 22a-diol
3(3, 22a-diacetoxystictane
3(3-acetoxystictan-22a-ol

O H OH 22a-hydroxystictane-3-one

H H H H stictane

Three new seco-stictane triterpenoids were isolated from the New Zealand species P. degelii by
Goh et al. (1978), possessing the novel 3,22-disubstituted 3,4-secostict-4(23)-ene structure

Ri

1 COOH 22a-hydroxy-3,4,-secostict-4(23)-ene-3-oicacid

2 CHO 22ct-hydroxy-3,4,-secostict-4(23)-en-3-ol

3 CH 2 OAc 3-acetoxy-3,4,-secostict-4(23)-en-22a-ol

3 . Lupane series: Twenty lupane triterpenoids including five (20RS)-epimeric pairs have been
isolated and identified in the chloroform extractives of the yellow-medulla species Pseudo-
cyphellaria rubella (Corbett et al., 1985; 1987). All fractions obtained were complex mixtures,
and multiple preparative layer chromatography (x 3) led to isolation of pure compounds. The
following lupane triterpenoids were characterized:

34

D. J. GALLOWAY

Rl

OAc

OAc

OH

OH

OAc

OH

OH

OH

OH

OAC

CHO

CH 3

CHO

COOH

COOH

CH 3

CH 2 OH

CH 3

CH 2 OH

CH 2 OH

R 3 R 4
CH 3 H
CH 3 OH
CH 3 H
CH 3 H
CH 3 H
CH 3 OH
CH 3 H
H OH
CH 3 OH
CH 3 OH

3(3-acetoxylupan-29-al
3(3-acetoxylupan-20-ol
(20RS)-3|3-hydroxylupan-29-al
(20RS)-3|3-hydroxylupan-29-oicacid
(20RS)-3(3-acetoxylupan-20-oicacid
lupan-3(3, 20-diol
(20RS)-lupan-3(3, 29-diol
(20RS)-30-norlupane-3p\ 20-diol
(20RS)-lupane-3|3, 20, 29-triol
(20RS)-3(3-acetoxylupane-20, 29-diol

OAc CH 2 CH 3
OAc O CH 3
OAc CH 2 CH 2 OH
OH O
OH CH 2
OH CH 2

CH 3
CH 3
CH 2 OH

3p-acetoxylup-20(29)-ene

3(3-acetoxy-30-norlupan-20-one

3|3-acetoxylup-20(29)-en-30-ol

3|3-hydroxy-30-norlupan-20-one

3(3-hydroxylup-20(29)-ene(lupeol)

Lup-20(29)-en-3p, 30-diol

RI R 2

OH OH 20,29-30-trinorlupane-3|3, 19-diol

OAc OAc 3(3, 19a-diacetoxy-20,29,30-trinorlupane

R

OAC

OH

3(3-acetoxylupan-20(29)-epoxide
3 |3-hydroxylupan-20(29)-epoxide

Role of secondary metabolites in Pseudocyphellaria

Secondary metabolites are defined as those compounds having no recognized role in the
maintenance of fundamental life processes in the organisms synthesizing them (Bell, 1981), a
definition which excludes intermediate and/or end products of primary metabolic pathways, and
photosynthetic pigments. The synthesis of secondary metabolites is complex, requiring multi-
step reactions in a sequential pathway mediated by precisely regulated enzymes or multi-enzyme
complexes (Luckner, 1980). Although there is conjecture over the adaptive significance of
secondary plant metabolites (Siegler, 1981) there is now a large body of evidence supporting a
primarily ecological role for these compounds, a proposition discussed particularly for lichens by

PSEUDOCYPHELLARIA 35

Rundel (1978). Secondary metabolites are implicated in a variety of ecological roles, as
defensive agents for plants in plant-plant (allelopathic), plant-pathogen, and plant-herbivore
interactions (Rhoades, 1979; Bell, 1980, 1981; Beart etal., 1985).

In Southern Hemisphere, cool-temperate, rain-forests a diverse and luxuriant epiphytic
lichen flora is well developed, with large, leafy species of Pseudocyphellaria, Menegazzia,
Nephroma, and Sticta constituting a rich potential food supply for vertebrate or invertebrate
herbivores. However, in Chilean Nothofagus forests Rundel (1978) records a surprising lack of
predation on large, nutrient-rich lichens in the above mentioned genera, and suggests that the
presence of terpenes may protect species producing them from herbivore predation. As with
bryophytes, in New Zealand, Tasmania, and south-east Australia species of Pseudocyphellaria
often reach a very great size (30-50 cm diam.) and normally show no evidence of attack by
herbivores. It is possible that they are protected against herbivore predation by medullary
and/or cortical secondary metabolites, such as pulvinic acid derivatives and triterpenoids. Thus,
apart from their importance in species-level taxonomy, triterpenoids in Pseudocyphellaria may
well prove to be effective anti-herbivore substances, an hypothesis ripe for experimental proof.

Green et al. (1985) suggest that in some species of Pseudocyphellaria both pulvinic acid
derivatives and triterpenoids can provide a water-proofed CO 2 diffusion pathway from external
pseudocyphellae, through the medulla to the photobiont layer, citing evidence from P. colensoi
(p. 70, fig. 7).

Ecophysiology

Over the past 15 years there has been considerable progress made in the understanding of lichen
ecophysiology, especially of water relations, gas exchange, photosynthetic rates, and respir-
atory rates (Rundel, 1982; Matthes & Feige, 1983; Kershaw, 1985; Lange & Ziegler, 1986). The
main bulk of ecophysiological data on lichens comes from species present in relatively open
habitats such as open woodland, tundra, deserts, and from the maritime Antarctic, with until
recently, little data on lichens in dense forests, especially from rain-forest in cool-temperate
and/or tropical areas (Green et al. , 1980).

A preliminary report on the physiological responses of Pseudocyphellaria glabra (and Sticta
stipitatd) in Tasmanian forest (Rundel et al., 1979) was followed by a series of detailed
ecophysiological studies [investigating CO 2 exchange, photosynthetic rate, respiratory rate, and
water relations] on species of Pseudocyphellaria [P. chloroleuca, P. colensoi, P. dissimilis, P.
glabra, P. homoeophylla, P. hookeri, P. lividofusca, P. rufovirescens] growing as conspicuous
epiphytes in North Island Nothofagus forest (Green et al., 1980, 1985; Snelgar et al., 1980;
Snelgar & Green, 1981a, 1981ft; Brown etal, 1981; Green & Snelgar, 1981; Snelgar, Green &
Beltz, 1981; Snelgar, Green & Wilkins, 1981). These studies of Green and his colleagues
conclusively demonstrate the suitability of species of Pseudocyphellaria for ecophysiological
studies in rain-forest, and point to such dominant epiphytes as important components in the
nutrient cycles of forest ecosystems.

Nitrogen fixation

The cyanobacterium Nostoc occurs widely in Pseudocyphellaria, both as a primary photobiont
as well as in internal (rarely in external) cephalodia in species having green primary photobionts
(James & Henssen, 1976). Cyanobacteria (blue-green algae) are microbial prokaryotes which
show characters both of gram-negative bacteria and of eukaryotic algae. Those features of
gram-negative bacteria are a prokaryotic cell structure with no membrane-bound organelles
(e.g. chloroplasts), no distinct nucleus, 70S ribosomes, and a glycoprotein cell wall, while
features characteristic of algae and higher plants include the capacity for oxygenic photosyn-
thesis, possession of a non-cyclic photosynthetic electron-transport chain and presence of
chlorophyll a (Stewart et al., 1980). Nitrogen fixation in Nostoc and other cyanobacterial
symbionts in lichens takes place in heterocysts, the frequency (20-25%) of heterocysts in
cephalodia (Green et al., 1980; Matthes & Feige, 1983) being higher than in vegetative tissue
(5%) with a cyanobiont as primary photobiont. Cyanobacteria form associative symbioses

36 D. J. GALLOWAY

(Stewart etal., 1979, 1985) with certain eukaryotic fungi, liverworts, ferns, gymnosperms, and
angiosperms. Associations are uncommon or rare and have limited economic or ecological
significance on a world basis, though nitrogen fixation by lichens is widespread in many
ecosystems, and in several may be of considerable importance (Stewart etal., 1979, 1980, 1985;
Green et al., 1980; Gibson & Jordan, 1983; Millbank, 19850, 19856; Sprent & Raven, 1985;
Rowell et al., 1985). Nitrogenous compounds (fixed from atmospheric nitrogen utilizing
nitrogenase and other nitrogen-metabolising enzymes, such as glutamine synthetase and
glutamate dehydrogenase), are lost from thalli of nitrogen-fixing lichens by leaching as a normal
feature of their existence (Millbank, 1985a, 19856) . The loss of carbon compounds from lichens
as a result of wetting after desiccation is now well documented (see Millbank, 19856), and a
similar situation exists with soluble nitrogen compounds (Crittenden, 1983) although the exact
nature of the combined nitrogen lost from the thallus is not known.

In arctic tundra and taiga vegetation the nitrogen input into the ecosystem from lichen fixation
is estimated at 0-5-1-5 kg hectare" 1 , annum" (Alexander, 1981) which accounts for c. 50%
of total nitrogen input, and in Colombian rain-forest, canopy lichens with cyanobionts
contribute 1-8 kg hectare" 1 , annum" 1 to the forest ecosystem (Forman, 1975).

In a preliminary survey of 44 New Zealand lichens for nitrogenase activity (estimated by the
acetylene-reduction technique), Green et al. (1980) surveyed 21 species of Pseudocyphellaria.
Fixation rates were found to be species specific ranging from 0-2-32 nmol g" 1 , min" 1 . Rates of
reduction were generally higher in species having a cyanobacterial symbiont (P. crocata, P.
dissimilis, P. hookeri, P. intricatd), although taxa with a green photobiont as primary symbiont
also exhibited a range of reduction rates since all of these taxa have internal cephalodia
containing Nostoc. Digestion of cephalodia with 10% chromium trioxide solution released
filaments of cyanobiont with a frequency of heterocysts of 20-25%. The observed variation of
nitrogen reduction rates in the different species of Pseudocyphellaria is not completely under-
stood, but may be due to frequency of cephalodia in green photobiont species, and to climatic
factors. Among species of Pseudocyphellaria with green photobionts, P. homoeophylla from
sites in both North and South Is consistently reduced acetylene at low rates (2-1 nmols C 2 H 2 g" 1
min." 1 ), P. colensoi consistently showed intermediate rates of reduction (4-6), and P. rubella
consistently exhibited high rates of reduction (12-5). Higher reduction rates are found in species
with high nitrogen contents. Nitrogen content is generally higher in Sticta (S. fuliginosa, S.
limbata) than in Pseudocyphellaria, also in both genera cyanobiont-containing species have the
higher reduction rates.

In Pseudocyphellaria, taxa with yellow medulla and pseudocyphellae tend to have lower
reduction rates than species with white medulla and pseudocyphellae and in addition species
with identical chemistry grouped together when acetylene reduction rates were plotted against
nitrogen content (% dry wt). Thus taxa with a two-hopane chemistry had higher rates of
reduction and higher nitrogen contents than taxa with two hopanes and stictic acid metabolites,
and taxa with pulvinic acid derivatives. The results of Green et al. (1980), although still only
preliminary, indicate that species of Pseudocyphellaria (and of Sticta) provide significant inputs
of nitrogen into native forests in New Zealand, and are a good basis for further, more detailed
work on the role of lichens in the nitrogen economy of rain-forest ecosy terns. Species of
Pseudocyphellaria 'are strongly represented in both species and quantity particularly in rain or
cloud forests (above 700 m altitude) and must represent a large biomass. Environmental
conditions are such, with heavy rainfall, and high insolation, that lichens could be metabolically
active and fixing nitrogen for a considerable proportion of each year . . . Preliminary estimates
of biomass indicate about 100 kg dry wt hectare" 1 in the lowest two metres of beech forest of the
Urewera National Park. This suggest a possible nitrogen contribution of between 1 and 10 kg N
hectare" 1 year" 1 by the lichens, a significant input in these rain forests where the alternative
nitrogen source would be rainfall averaging 1 to 2 kg N hectare" 1 year" 1 ' (Green et al., 1980:
348).

Snelgar & Green (1981fl) found a positive linear relationship between rate of acetylene
reduction and thallus water content. In a study of populations of the nitrogen-fixing species P.
dissimilis, shade, mesic, and sun populations were compared. It was found that the upper cortex

PSEUDOCYPHELLARIA 37

of all three populations were of equal thickness but that mesic and sun populations had a thicker
photobiont layer and lower cortex, while the sun population had a thicker medulla and thicker
and denser rhizines. These observed differences in thallus morphology correlated with evapor-
ative demand of their environments. Sun (least shaded) populations are closely appressed to the
substrate whereas shaded populations are only loosely attached and project from the substrate.
Sun populations also show three adaptations which help maintain the lichen in a moist condition:
(1) thalli have the classical xerophyte characteristic of reduced surface area to volume ratio
(accounted for in the increase in thallus thickness); (2) an increased water storage capacity is
attained by increased thallus thickness and a more developed rhizine layer; and (3) thalli are
closely appressed to the substrate thereby assisting water storage by rhizines and reducing the
area exposed to water loss by evaporation. Rates of acetylene reduction on an area basis
(expressed as nmol cm~ 2 min~ f x 10 5 ) for the three populations of P. dissimilis were: shade (164
16), mesic (193 10), and sun (262 20), with the least shaded populations showing a
decidedly increased rate of acetylene reduction as a result of the increased depth of cyanobiont
layer.

Growth

New Zealand's forests [beech (Nothofagus) forest, beech-podocarp forest, and podocarp-mixed
broadleaf (hardwood) forest] have a richly developed epiphytic lichen flora that is dominated by
large, foliose species which form often visually striking clones (20-50 cm diam.), where
conditions of light, humidity, and precipitation are optimal. Dominant epiphytes of tree trunks
in such situations include: Lobaria adscripta (Galloway, 19816), Parmelia tenuirima (Galloway
& Elix, 1983), Sticta subcaperata, S. latifrons, and several species of Pseudocyphellaria, the most
common being P. billardierei, P. colensoi, P. coriacea, P. coronata, P. dissimilis, P. faveolata, P.
fimbriata, P. glabra, P. gretae, P. homoeophylla, P. lividofusca, and P. rufovirescens . Rundel
(1980) speculates that the diversity and biomass of corticolous lichen communities in southern
Chile, Tasmania, and New Zealand probably exceed those of corticolous communities any-
where in the Northern Hemisphere, and that the cool, humid conditions found in forested areas
in these southern regions provide near ideal growing conditions for lichens. For a recent review
of lichen growth and productivity see Matthes & Feige (1983).

The forested areas of New Zealand have a climate favourable for optimal lichen growth,
rainfall being high [c. 150-300(-400) cm annually] , and mist or cloud cover common throughout
the year. Large foliose lichens are best developed on tree trunks (from below the canopy
branches to near the ground) where there is a low evaporative stress as the closed canopy is
10-25 m above the ground. In closed forest, light is a limiting factor and only those species of
Pseudocyphellaria with a cyanobiont as primary photosynthetic symbiont (usually P. dissimilis
and P. fimbriatoides or locally P. cinnamomea and P. hookeri) are able to compete successfully
with the bryophyte cover. The most spectacular growth of Pseudocyphellaria is to be found in
the Nothofagus forests close to the Main Divide in South Island where P. homoeophylla is
especially prolific, covering tree trunks up to 6 m above the ground and spreading over the forest
floor in large swards. As a general rule, the richest development of epiphytic species occurs at
forest margins, and near edges of lakes rivers, and streams where humidity is high and light is
sufficient.

The large size and luxuriant habit of the species of Pseudocyphellaria mentioned above, as
well as of various fruticose taxa (e.g. Sphaerophorus , Thysanophoron) implies high growth rates
and this is confirmed in the relatively few measurements made to date (Snelgar & Green, 1982).
Growth rates of several species of Pseudocyphellaria and of the fruticose Sticta filix were
calculated from sequential photographs taken over a two-year period in Nothofagus forest in the
Urewera National Park, North Island, New Zealand. Mean annual increases in P. homoeo-
phylla and in Sticta subcaperata were linearly related to thallus diameter and ranged from
2-0-27-0 mm and 3-0-16-7 mm respectively. Growth rate estimates for 32 individual thalli were
taken over periods of 386 to 733 days. Both taxa showed a dependence of growth rate on thallus
size (not constant radial growth), a situation generally found only in small thalli (Topham,

38 D. J. GALLOWAY

1977). Other species examined had the following mean annual growth rates: P. faveolata
(3-7-13-1 mm), P. lividofusca (4-6-9-4 mm), and Stictafilix, a fruticose, stalked species but
lobate above the stalk (9-3-20-0 mm). Useful information on lichen biomass in Tasmanian rain
forests is given in Kantvilas (1985).

Relatively rapid growth rates in New Zealand lichens are not only restricted to foliose and
fruiticose species in forested areas. In high-alpine regions and on glacial moraines the growth
rates of the crustose lichen Rhizocarpon geographicum have had some application in dating
Holocene glacial deposits. The growth rate of this species in the Mt Cook area is one of the most
rapid recorded anywhere in the world for this lichen, being 62-5 mm per 100 years (Burrows" &
Orwin, 1971; Birkeland, 1981). A high growth rate is to be expected for the Mt Cook area
because of the high annual precipitation there (over 4 m per year).

The high growth rate of many epiphytic species of Pseudocyphellaria in New Zealand,
together with their production of medullary triterpenoids which inhibit predation by herbivores,
make these plants an important component of the forest biomass. Green et al. (1980) have
published a preliminary estimate of the biomass of epiphytic lichens in the lowest two metres of
the beech forest of Urewera National Park amounting to 100 Kg dry weight per hectare.

Geographical setting

New Zealand is an island archipelago and has been so throughout the greater part of its
geological history (Fleming, 1979a). It consists of three main islands (North, South, and
Stewart) and numerous smaller islands or island groups (Antipodes, Auckland, Bounty,
Campbell, Chatham, Kermadec, Snares, and Three Kings), extending over c. 23 of latitude or
some 2500 Km, in the South Pacific Ocean, from the subtropical Kermadec Islands in the north,
to the subantarctic Campbell Island in the south. The New Zealand landmass is surrounded by a
shallow, well-defined submarine platform, the continental shelf, sloping gently downwards from
the intertidal zone to 150-180 m. Below this level, the seafloor gradient steepens abruptly and
passes into the continental slope which plunges to depths usually greater than 1000 m.

The islands of New Zealand lie at the hub of a complex of submarine plateaux and ridges with
intervening basins and associated deep trenches (Fig. 12). To the south-east and north-west of
New Zealand, the Campbell Plateau (Adams, 1983; Adams et al., 1979), Chatham Rise, Lord
Howe Rise, Norfolk Ridge are presumed remnants of the former continental margin and
peripheral geosynclinal belt of the Pacific (Panthalassic) margin of Gondwanaland. The
submarine ridges and plateaus represent crustal strips separated from each other, and from
Australia and Antarctica by rifting, with formation of new sea floor below the Tasman Sea, and
the south-western Pacific Ocean continuing since late Cretaceous times. In contrast, the
Kermadec-Colville ridge system to the north, north-east, and the Macquarie Ridge to the
south-west of New Zealand, are seismically active, volcanic island arcs formed concurrently with
new sea floor during the Cenozoic (Cullen, 1976, 1978).

New Zealand lies at the boundary of the India- Australian Plate and the Pacific Plate; the deep
Kermadec (north-east) and Puysegur (south-west) trenches denote subduction zones, where the
edge of one crustal plate is forced under its neighbour. In the north-east, Pacific Ocean floor is
being subducted under the edge of the India- Australian Plate, while to the south-west, the
Pacific Plate is being pushed over the India- Australian Plate. The Alpine Fault, a transcurrent
fault of considerable tectonic activity, links these two subduction zones (Stevens, 19806).

Ecology

Habitats

Habitats available to lichens in New Zealand are particularly various over a limited area, a
consequence of diverse geology, vegetation, and climate. Conditions range from subtropical to
alpine within relatively short distances, precipitation varies from 350 to 12,000 mm per annum,
and the three main islands cover 13 of latitude, comprising between them a widely varying
topography from rocky shore to glaciated summit. The lichen flora of New Zealand is extremely

PSEUDOCYPHELLARIA

39

'~ / \^*~ I ^"^

Fig. 12 New Zealand and the surrounding ocean floor showing submarine plateaux and ridges with
intervening basins and deep trenches (Cullen, 1976).

diverse, reflecting this rich array of habitats, and comprises in excess of 200 genera (Galloway,
19856).

Species of Pseudocyphellaria are primarily epiphytes in forest and scrub vegetation (see
below), although a few species also grow on soil, viz. , P. dissimilis in low-light environments on
the forest floor or over rocks, in warm, moist, temperate rain-forest [especially in podocarp or
podocarp-broadleaved, hardwood forest in south-eastern and western South Island, and
throughout North Island]; P. crocata, P. neglecta, and P. pickeringii on stones or boulders
(rarely on walls) in dry, sunny, open sites in short grassland; and P. crocata (occasional), P.
degelii, P. glabra, P. maculata, and P. pickeringii, and occasionally P. neglecta, on soil or on

40 D. J. GALLOWAY

rocks in subalpine tussock grassland dominated by species of Celmisia and Chionochloa
(snowgrass), or at the base of subalpine scrub such as Cassinia, Dracophyllum, or Olearia.

Climate

New Zealand lies across the mid latitude zone of westerly winds, and its climate is greatly
influenced by this predominant wind flow (Coulter, 1973; Tomlinson, 1976; Fleming 19796).
New Zealand occupies an isolated position in a vast domain of ocean (Fig. 13) which ensures that
the westerly winds reaching it are moisture-laden; hence in many parts and especially on western
coasts and in mountainous regions, humid conditions are common. The passage of winds over
great expanses of ocean has a moderating effect on the air temperature. The main mountain
ranges of New Zealand are aligned north-east/south-west, and offer a major barrier to the
prevailing moisture-laden westerlies, and in South Island especially, western slopes of the Main
Divide receive very high rainfalls (to 12,000 mm per annum). East of the Main Divide, rainfall
drops progressively in a rapid gradient, with the intermontane basins of Central Otago and
Canterbury receiving the lowest rainfalls of any region in the country (to 350 mm per annum).

Species of Pseudocyphellaria are found in New Zealand in five bioclimatic zones viz.,
subtropical, warm temperate, cool temperate, subantarctic, and low antarctic (nomenclature

ThtScum
Auckland!*'
C.mpbellld
Kbopurtt kL<

ERA/

Fig. 13 New Zealand and the southern oceans and landmasses.

PSEUDOCYPHELLARIA 41

follows Meurk, 1984) with most taxa represented in warm-temperate and cool-temperate zones
between lats 35S and 48S.

Vegetation

New Zealand's native vegetation derives from plants which have survived from Gondwanan
times when New Zealand was part of the Panthalassic margin of that great southern landmass , as
well as from those which arrived by long-distance dispersal during some 60 million years of
isolation. The New Zealand flora comprises c. 2000 species of vascular plants of which about 40
genera and 85% of the species are endemic. This high degree of endemism at species and generic
levels (there are no endemic families), and the noteworthy development of certain genera such
as Hebe, is sufficient to give New Zealand the status of a distinct botanical region (Good, 1974;
Godley, 1976). The vegetation is extremely varied since habitat conditions range from sub-
tropical to alpine, precipitation is from 350 to 12,000 mm per annum, and there is a great
diversity of sites. Species of Pseudocyphellaria are found mainly in two principal vegetation
types, forest communities from sea-level to tree-line and in subalpine-alpine scrub and/or
grassland communities above tree-line.

Forest: It is generally accepted that at the advent of Polynesian settlement, c. 1000 years ago,
some 75% of the total land surface had a continuous forest cover, amounting to c. 20,000,000
hectares. Forest was absent from the 15% of the country lying within the alpine zone, and from a
further 2% of mostly lowland swamps or bogs, gravel floodplains, and coastal sand dunes. The
remaining 8% of New Zealand comprises areas of very low rainfall, <650 mm per annum, and in
these areas was probably a mosaic of forest, scrub, and tussock-grassland, subject to occasional
lightning fires, but not true savannah. These irregularly forested districts were mainly in the
eastern parts of South I. , but were locally significant also in parts of North I. (Nicholls, 1980). As
a result of Polynesian and natural fires (McGlone, 1983), and later logging and land clearance
subsequent to European settlement from 1840 to the present, natural forests cover
c. 23% of the land surface (a little over 6,000,000 hectares), mainly in upland or mountainous
areas often with high rainfall.

Main canopy species of New Zealand forests are generally characterized as 'softwoods'
(coniferous species) or 'hardwoods' (dicotyledonous species), and four main physiognomic
elements are recognized viz., kauri (Agathis austmlis) forest; beech (Nothofagus) forest;
softwood forest; and broadleaf hardwood (hardwoods other than beech) forest (Robbins, 1962;
Dansereau, 1964; Poole & Adams, 1980; J. Wardle, 1984).

46% of New Zealand's indigenous forests are 'pure beech forest' with a canopy dominated by
species of Nothofagus, and 22% of the forests are mixtures of beech with conifers or broadleaf
hardwoods (J. Wardle, 1984). Of the beech species, silver beech (Nothofagus menziesii) is most
common when the climate is cool and wet, and large silver beech forests are found in montane
and subalpine areas of the western parts of South I. Mountain beech (N. solandri var.
diffortioides) occurs in montane and subalpine zones of drier areas and forms extensive natural
monocultures in the central parts of South I. Red beech (N. fusca) is primarily confined to
lowland or montane forests occupying deep, fertile soils. Black beech (N. solandri var. solandri)
and hard beech (N. truncatd) usually occur at lower altitudes in forests mixed with conifers
and/ or with other hardwood species.

The remaining 32% of New Zealand's indigenous forests are dominated by conifers and
hardwoods other than Nothofagus. Kauri forest occurs in the northern parts of North I., with
Agathis australis emerging from a canopy of other conifers such as Phyllocladus trichomanoides ,
and hardwoods e.g., Beilschmiedia tawa, Knightia excelsa, Weinmannia silvicola, and other
trees.

Forests dominated by other softwoods (mainly podocarps such as Dacrydium cupressinum,
Dacrycarpus dacrydioid.es, Prumnopitysferruginea, etc.) are found mainly on moderately fertile
soils in North I. (e.g. in Urewera National Park), and at low altitudes (e.g. in Westland).

Conifer-broadleaf hardwood forest with a mixed canopy of conifers and hardwoods other
than beeches, are found mainly in steepland areas with wet climates.

42 D. J. GALLOWAY

Conifer-broadleaf hardwood forests occur especially in montane areas with a mild climate and
relatively high rainfall. They are especially well-developed in central Westland. The steepland
montane conifer-broadleaf hardwood forests are commonly characterized by few dominant
canopy species, mainly Metrosideros umbellata, Weinmannia racemosa, and Quintinia acuti-
folia. At lower altitudes (600 m) these angiosperms are associated with the conifers Dacrydium
cupressinum and Prumnopitys ferruginea, and at higher altitudes (800 m), Libocedrus bidwillii
and Podocarpus hallii.

In beech forests and in mixed beech-podocarp forests especially, species of Pseudocyphellaria
reach their maximum diversity, development, and abundance, sometimes as ground cover (P.
homoeophylla),but more commonly as epiphytes, with extensive clones of the following species
often present: P. colensoi, P. coronata, P. degelii, P. faveolata, P. homoeophylla, P. multifida
and P. mfovirescens .

At altitudes above c. 1000 m, forest gives way to alpine vegetation (see below), with or
without a zone of scrub at tree-line separating the forest from the tussock-grasslands and fellfield
of the alpine zone. Tree-line, a major ecological boundary (see P. Wardle, 1985) represents the
altitude at which climate becomes too cold to support growth of trees. In New Zealand two
species of Nothofagus form most tree-lines; on drier mountains N. solandri var. cliffortioides
grows as pure stands, and on wetter mountains both this species and N. menziesii occur together
with a much richer accompanying flora. On the latter species at tree-line, a rich selection of
Pseudocyphellaria is met with, including P. ardesiaca, P. corbettii, P. episticta, P. glabra,
P. granulata, P. gretae, P. lividofusca, P. pickeringii, P. pubescens, and P. rubella.

However, Nothofagus is widely absent in areas deforested during Pleistocene glaciation, or
volcanism (P. Wardle, 1964; J. Wardle, 1984), and instead there are floristically complex
treelines of small trees and large shrubs belonging to genera such as Podocarpus, Dacrydium
(Podocarpaceae), Hoheria (Malvaceae), Senecio and Olearia (Compositae), Dracophyllum and
Archeria (Epacridaceae) , and Hebe (Scrophulariaceae) . Species of these genera often form a
dense, interlocking subalpine scrub zone which can extend for 100-300 m above the upper limit
of tall forest. Species of Pseudocyphellaria found in subalpine scrub include P. crocata,
P. degelii, P. faveolata, P. glabra, and P. maculata. .

Alpine vegetation: The alpine zone in New Zealand (of some 1000 m elevation) extends from
the tree-line, where present, upwards to the summer snowline, the level to which snow thaws
during most summers (Mark, 1979). Above tree-line there is generally a belt of grassland
dominated by several tall species of snow tussock (Chionochloa) , sometimes also with an
important co-dominant shrub component such as Aciphylla or Dracophyllum. This zone is
termed the low-alpine zone (P. Wardle, 1964; Mark, 1979), and comprises mixed snow tussock
scrub (Mark & Adams, 1973) including species of Chionochloa together with scrub species such
as Dracophyllum uniflorum, Hebe odor a, Phyllodadus alpinus, Podocarpus nivalis, etc., and
also snow tussock herbfield with Chionochloa and species of Aciphylla, Anisotome, Celmisia,
and occasionally Dracophyllum and Hebe. In snow tussock herbfield which Mark & Bliss (1970)
regard as being the lower level of the high-alpine zone, the following species of Pseudocyphel-
laria are found: P. degelii, P. glabra, P. maculata, and P. pickeringii, either among tussocks or
herbs such as Celmisia viscosa, or at the bases of low scrub (Aciphylla or Dracophyllum). In the
high-alpine zone (P. Wardle, 1964; Mark & Bliss, 1970; Mark & Adams, 1973; Bliss & Mark,
1974; Mark, 1979; Gibson & Kirkpatrick, 1986) vegetation is low and less continuous and
consists of fellfield, cushion vegetation, snowbank vegetation, and scree. The only species of
Pseudocyphellaria which penetrate into the high-alpine zone are P. degelii, P. glabra, and P.
maculata, some or all being found in fellfield on the Central Otago mountains (Mark & Bliss,
1970), and on mountain ranges of Canterbury from Mt Technical to the Kirkliston Range.

The tussock grasslands and tundras of the subantarctic shelf islands (Antipodes, Auckland,
Campbell, and Macquarie Is) are closest ecologically to the snow tussock grasslands and
high-alpine tundras of the central and eastern mountains of South I. (Bliss, 1979), and in these
exposed subantarctic habitats, P. degelii, P. glabra, and occasionally P. physciospora occur. In
contrast, the Low Subantarctic closed forest (Metrosideros) of the Auckland Is and the High

PSEUDOCYPHELLARIA 43

Subantarctic scrub (Dracophyllum) of the Auckland and Campbell Is (Bliss, 1979) supports a
more diverse assemblage of Pseudocyphellaria including P. billardierei, P. coronata, P. faveolata,
P. glabra, P. multifida, P. physciospora, and P. rubella.

Factors affecting distribution of Pseudocyphellaria

Detailed studies on the ecology and phytosociology of cool-temperate South Pacific lichens are
still in their infancy, the most comprehensive account to date being that of Kantvilas (1985) on
Tasmanian rain forest macrolichens. Apart from a preliminary phytosociological study of
cryptogamic epiphytes in the Camp Creek catchment of central Westland (Reif & Allen,
unpublished), nothing is known of the phytosociology of such prominent lichen genera as
Pseudocyphellaria, Psoroma, Menegazzia, Lobaria, Sphaerophorus, Sticta, and Usnea in a New
Zealand setting. Several ecological factors, either singly or in combination, such as moisture,
light, temperature, altitude, substrate, etc., may influence the distribution of species of
Pseudocyphellaria in New Zealand, though detailed studies of these parameters have yet to be
made.

A major factor controlling the distribution of species of Pseudocyphellaria is rainfall, with the
richest diversity of taxa being found in forested areas having an annual average rainfall in the
range 600-8000 mm. In the very driest areas of New Zealand, such as Central Otago, which has
an annual rainfall of c. 400 mm or less, species of Pseudocyphellaria are uncommon, with only
occasional specimens of P. crocata, P. glabra, and P. pickeringii found among rocks in dry
grassland or induced steppe. As a general rule, species of Pseudocyphellaria prefer mild
temperatures and cloudy, foggy, drizzling conditions with high humidity, such as are found in
areas of medium to high rainfall, or where the number of rain days is high.

Light is another important factor, with the richest diversity of species found in habitats of
moderate to strong illumination, especially at forest margins, at tree-line, and on larger
branches of the forest canopy. Species with a cyanobiont (e.g. P. allanii, P. cinnamomea, P.
dissimilis, P. hookeri) are much more tolerant of shade than species with a green photobiont,
indeed P. cinnamomea and P. dissimilis often comprise the main ground cover in deeply shaded
interiors of lowland forests where the ambient light level is very low and the humidity high.
These two species, together with Lobaria retigera and Peltigera dolichorhiza, are the main (often
the sole) foliose lichens able to compete successfully with bryophytes under conditions of low
illumination in forest interiors. The green photobiont species P. multifida also appears to be able
to grow in moderately shaded habitats and is often found at the base of forest trees together with
P. billardierei, P. glabra, P. homoeophylla, and P. pickeringii. The vertical distribution of species
Pseudocyphellaria on the trunks of forest trees is discussed by Kantvilas (1985) for Tasmanian
rain-forest, and a similar and noteworthy gradient of species is seen in many New Zealand
forests with often large clones of P. colensoi, P. coriacea, P. coronata, P. faveolata, P. gretae, P.
lividofusca, and P. rufovirescens augmenting or displacing the basal species mentioned above.

Besides moisture and light, other factors affecting the distribution of Pseudocyphellaria in
New Zealand are latitude, altitude, and mean annual temperature, with groupings of species
being influenced by all of these parameters in varying degrees. For instance, several species are
found in mainly northern coastal habitats (e.g. P. aurata, P. carpoloma, P. chloroleuca, P.
montagnei, P. poculifera) which have a mean annual temperature of c. 12-17C, while other
species are found in subalpine to high alpine grasslands exposed to a high number of ground
frosts and to freezing or subzero temperatures for long periods.

The nature of the substrate, and also the floristic composition of forests may also influence the
species of Pseudocyphellaria present in particular habitats. For example, species of Pseudocy-
phellaria (and of many other associating lichens) are usually poorly represented on the bark of
Nothofagus fusca, especially in pure stands of this tree, whereas the species diversity of
Pseudocyphellaria on the bark of N. menziesii and, to a lesser extent, N. solandri var.
cliff ortioides , is much greater. The moisture-retaining bark of Leptospermum also seems to be a
very favourable substrate for species of Pseudocyphellaria, especially when illumination and
humidity are optimal.

44 D. J. GALLOWAY

Biogeography

Pseudocyphellaria in the family Lobariaceae Chev., is a primarily Southern Hemisphere
cool-temperate genus of c. 110 species, with New Zealand (48 species) and South America (c. 46
species) being the main present-day areas of specification (Fig. 14).

Biogeography concerns itself with present and past patterns of spatial distribution of life on
earth and the means whereby such distributions have come about. Problems of Southern
Hemisphere biogeography have concerned botanists, zoologists, and palaeontologists for many
years, with interpretations of past and present distribution patterns of certain southern genera,
for example Nothofagus, attracting considerable discussion and controversy (Craw, 1982, 1985;
Heads, 1985; Humphries, 1981, 1983, 1985; Humphries & Parenti, 1986; Melville, 1982; van
Steenis, 1971). In lichen-forming fungi, disjunct distributions in many genera parallel dis-
junctions in other plant and animal groups, and in spite of their polyphyletic origins and dual
nature, lichens and their distribution patterns seem capable of conventional biogeographic
analysis.

Pseudocyphellaria is one of several lichen genera which have a rich development of species in
the Southern Hemisphere but with only one or a few widespread species present in the Northern
Hemisphere. Other mainly southern lichen genera with which Pseudocyphellaria can be
compared include Menegazzia (Santesson, 1942; James in Galloway, 19856); Neuropogon
(Lamb, 1939, 1948, 1964; Walker, 1985); Placopsis (Lamb, 1947); Psoroma (J0rgensen, 1978;
Henssen & Renner, 1981; Henssen et al., 1983; Henssen, 1983; Galloway, 19856); Siphula
(Galloway, 19856); and Sphaerophorus (Sato, 1968; Ohlsson, 1974; Tibell, 1984; Galloway,
19856). Explanations for such southern distributions found in these various genera must be
looked for in terms of accepted biogeographical frameworks, with the concepts of plate
tectonics, and the accretion and dispersal of terranes being relevant to most arguments.

The idea that the earth is made up of a series of interlocking plates whose geometry has been
subject to cyclic geographical rearrangements through geological time is the central hypothesis
of plate tectonics. The earth appears to have consisted of a single 'supercontinent' Pangaea,
surrounded by 'superocean' Panthalassa in the late Palaeozoic. In the early Mesozoic two major
components of this single landmass, Gondwanaland in the south, and Laurasia in the north,
were separated by a triangular re-entrant of Panthalassa, the Tethys Sea, of which today's

Fig. 14 World distribution of Pseudocyphellaria with numbers of species recorded from the cool-
temperate Southern Hemisphere (Australia-Tasmania, New Zealand, Juan Fernandez, southern South
America), from the tropics, and from the Northern Hemisphere cool-temperate zone.

PSEUDOCYPHELLARIA 45

Mediterranean Sea is the remnant. Warm-temperate, palaeotropical taxa present in Southern
Hemisphere biotas derive from organisms that originally populated the Tethyan frontage of
Gondwanaland, while cool-temperate taxa were present along the Panthalassic margins of
Gondwanaland.

The fragmentation of Gondwanaland and the subsequent movement of its constituent plates
is of central importance in discussions of Southern Hemisphere biogeography, and details of the
timing of fragmentation and subsequent continental drift of tectonic plates are now fairly well
known (e.g. Molnar etal., 1975; Barker & Griffiths, 1977; Weissel etal., 1977; Coleman, 1980;
Craddock, 1982; Stevens, 1983; Kamp, 1986). Initial rifting led to the separation of West (South
America- Africa), and East Gondwanaland (Antarctica-India- Australia-New Zealand); then
followed the separation of South America from Africa with the opening of the southern Atlantic
Ocean and the separation of India from Antarctic- Australia.

The separation of the New Zealand microcontinent (including New Zealand, the Campbell
Plateau, the Chatham Rise, the Norfolk Ridge, and the Lord Howe Rise - see Fig. 12), itself
thought to be of polyphyletic origin (see Howell, 1980), from the Panthalassic margin of
Gondwanaland began c. 81 million years ago and was completed with the opening of the Tasman
Sea (Griffiths, 1971; Hayes & Ringis, 1973; Weissel et al., 1977; Crook & Belbin, 1978;
Coleman, 1980; Grindley & Davey, 1982; Cawood, 1984). The final stages in the fragmentation
of Gondwanaland were the separation of Australia from East Antarctica (Mutter et aL, 1985),
the opening of the Drake Passage between Antarctica and South America (Lawver et al. , 1985) ,
and the initiation of the circum- Antarctic current (Jenkins, 1964; Kennett etal., 1974; Kennett,
1977, 1979, 1980;Berggren&Hollister, 1977), and the West Wind Drift (Fell, 1962; Close etal.,
1978; McDowall, 1978; Fleming, 19790, 19796; Knox, 1979, 1980). Plate tectonics in relation to
Southern Hemisphere biogeography is discussed by Melville (1966, 1967); Keast (1971, 1973);
Raven & Axelrod (1972); Schuster (1976); and Raven (1979).

Recently the concept of accretion and dispersal of terranes (Nur & Ben-Avraham, 1977, 1982;
Kamp, 1980; Howell, 1980, 1985) was used in Southern Hemisphere biogeographical debate
(see Craw, 1982, 1983, 1985). Terranes are crustal blocks bounded by faults and not necessarily
of uniform composition, which may accrete onto existing continental crust thus introducing new
geological (and ? biological) material into a pre-existing formation. New Zealand is thought to
be composed of at least four terranes of different origin (Howell, 1980) and it seems that many,
if not most, of the circum-Pacific landmasses are also assemblages of unrelated geological
taxa (Nur & Ben-Avraham, 1982; Craw, 1985). The terrane concept has thus important
consequences for future arguments concerning lichen biogeography (see also Galloway,
1988).

Explanations of Southern Hemisphere biotic distribution patterns fall into two main classes,
(1) Dispersal explanations, and (2) vicariance explanations. Dispersal models postulate spread
from centres of origin (e.g. Darlington, 1965), a view deprecated by many recent bio-
geographers (e.g. Croizat et al., 1974; Craw, 1978; Cracraft, 1980; Humphries, 1981, 1983).
Vicariance models involve fragmentation of an originally widespread biota into two or more
descendant biotas, resulting in the formation of allopatric species (vicariants), the fragmenta-
tion involving the formation of a barrier preventing gene flow between the component parts of
the originally continuous population (e.g. Rosen, 1978; Nelson & Platnick, 1980a, 19806).
There is now widespread acceptance of vicariance explanations in the interpretation of Southern
Hemisphere biotic distribution patterns (see Brundin, 1966, 1975; Croizat et al., 1974; Cracraft,
1975; 1980; Humphries, 1983), although Craw (1985) strongly advocates use of Croizat's (1958)
panbiogeographic methods.

Lichens are an ancient plant group with very slow rates of evolution (Hawksworth, 1982a) and
were undoubtedly well-developed in cool-temperate Panthalassic Gondwanaland before frag-
mentation and continental drift. Pseudocyphellaria or its progenitors would be present on the
cool-temperate vegetation of Panthalassic Gondwanaland, as the high number of vicariant
species between New Zealand and South America shows; however, until species from Australia
and South America are studied in more detail a biogeographic analysis of the Southern
Hemisphere taxa will not be given here.

46 D. J. GALLOWAY

Five biogeographical groupings or elements are recognized in the New Zealand Pseudo-
cyphellaria flora and comprise endemic, australasian, austral, palaeotropical, and cosmopolitan
taxa. They are each discussed separately below.

Endemic element

Just over half of the species of Pseudocyphellaria found in New Zealand (25 species or c. 52%)
are endemic and include: P. allanii, P. carpoloma, P. corbettii, P. coriacea, P. crassa, P. degelii,
P. durietzii, P. episticta, P.fimbriata, P.fimbriatoides, P. gretae, P. homoeophylla, P. hookeri,
P. knightii, P. lindsayi, P. lividofusca, P. maculata, P. margaretiae, P. montagnei, P. murrayi, P.
nermula, P. pubescens, P. rufovirescens , P. sericeofulva, and P. wilkinsii. Of these taxa, 17 are
fertile, primary species without any vegetative propagules, and nine have either isidia or
phyllidia as vegetative propagules. Although some taxa are widespread in New Zealand (e.g.,
P. coriacea, P. homoeophylla, P. rufovirescens), several of the endemics have restricted
geographical ranges such as P. carpoloma, P. hookeri, P. montagnei, and P. sericeofulva from
North I. habitats, while P. corbettii, P. crassa, P. maculata, and P. nermula are South I. species.
Three species, P. crassa, P. corbettii, and P. margaretiae are best developed in north-west
Nelson while the remaining species are more widely scattered.

There is a strong correlation between the distribution of Pseudocyphellaria and of Nothofagus
in New Zealand and most species of Pseudocyphellaria are found in lowland to montane beech
or beech-podocarp or mixed beech-hardwood forests, with only P. degelii and P. maculata
(together with the austral P. glabra) penetrating above tree-line into subalpine to high-alpine
grassland. Pseudocyphellaria is linked ecologically and evolutionarily with angiosperm phoro-
phytes, especially with Nothofagus, and would no doubt have been present in New Zealand,
Australia, and South America in the Late Cretaceous and Early Palaeocene at the time of the
emergence of the three major groups of beeches, Nothofagus brassii, N. fusca, and N. menziesii
(Mildenhall, 1980).

New Zealand's isolation, and the persistence of moist, cool-temperate conditions in both
forest and shrubland habitats since the Cretaceous has given rise to many relict plants, from
angiosperms to mosses and hepatics (Mildenhall, 1980; J. Wardle, 1984), as well as to many
lichens. Although the level of endemism at the generic level is small (Galloway, 1979, 1985b)
[only Thysanophoron in the Caliciales is endemic] , at the species level in lichens of Southern
Hemisphere genera found in New Zealand, the incidence of endemic taxa is much higher. The
most important of these endemic lichens (after species of Pseudocyphellaria) are: Anziajamesii
(Galloway, 1978); Argopsis megalospora (Galloway, 1980/>); Austroblastenia pupa (Sipman,
1983); Brigantiaea lobulata, B. tabacodes (Hafellner, 1983); Caloplaca homologa; Chaenotheca
degelii (Tibell, 1983); Collema novozelandicum (Degelius, 1974); Cryptolechia myriadella;
Dendriscocaulon dendroides; Haematomma babingtonii, H. hilare; Hyperphyscia plinthiza;
Leproplaca lutea; Lobaria adscripta, L. asperula, L. dictyophora; Megaloblastenia flavidoatra
(Sipman, 1983); Megalospora bartlettii, M. disjuncta, M. gompholoma, M. knightii (Sipman,
1983, 1986); Melanelia calva, M. glabratuloides (Esslinger, 1977, 1978); Menegazzia aucklandi-
ca, M. dielsii, M. foraminulosa, M. inflata, M. lucens, M. pertransita, M. pulchra; Mycoblastus
campbellianus; Neofuscelia adpicta, N. brattii, N. epheboides, N. martinii, N. melanobarbatica,
N. minuta, N. peloloba, N. petriseda, N. pictada, N. plana, N. waiporiensis (Esslinger, 1977,
1978); Pannariaperiptera, P. subimmixta; Parmeliella apiculata, P. gymnocheila, P. mucorina,
P. neozelandica (Galloway & James, 1984), P. thysanota, P. variegata; Peltularia crassa
(J0rgensen & Galloway, 1984); Pertusaria allanii, P. amaurospora, P. circumcincta, P. erumpes-
cens, P. graphica, P. laevis, P. lavata, P. leucodes, P. leucodioides , P. melaleucoides , P.
novaezelandiae, P. obvelata, P. perfida, P. perrimosa, P. subverrucosa, P. theochroa, P.
truncata; Phlyctella longifera, P. megalospora, P. oleosa, P. sordida, P. uncinata; Phylisciella
aotearoa (Henssen & Biidel, 1984); Physcia tenuisecta; Placopsis gelidioides, P. illita, P.
lateritioides , P. salazina, P. subgelida, P. subparellina; Polychidium contortum; Psoroma
allorhizum, P. athroophyllum, P. buchananii, P. coralloideum, P. euphyllum, P. implexum, P.
melanizum, P. xanthomelanum; Rinodina cacaotina, R. gallowayi, R. jamesii, R. murrayi, R.

PSEUDOCYPHELLARIA 47

nigricans, R. otagensis, R. peloleuca, R. proprior, R. subtubulata, R. tibellii, R. tubulata
(Mayrhofer, 1983); Sagenidium citrinum (Follmann, 1975); Scleromorpha amabilis (Tibell,
1984); Siphula dissoluta; Sphaerophoms microspoms , S. notatus; Steinera polymorpha,
S. radiata, S. sorediata (Henssen & James, 1982); Stereocaulon colensoi, S. fronduliferum, S.
gregarium, S. loricatum (Galloway, 19806); Sticta babingtonii, S. cinereoglauca, S. filix,
S. lacera, S. latifrons, S. martinii; Thelotrema circumscriptum, T. farinaceum, T. obovatum,
T. saxatile; Thysanophoron stereocauloides; Trapeliopsis colensoi; Usnea oncodes , U. tenerior,
U. xanthophana, U. xanthopoga; andXanthoria novozelandica.

The 54% endemism of New Zealand's Pseudocyphellaria flora compares with Australia (35
species- endemism 17%), South America (46 species- endemism c. 65%), and Juan Fernandez
(22 species - endemism 30%) and shows the relatively high level of endemism found in austral
populations (Galloway, 1987). In these austral populations and areas of speciation, similar
habitats in New Zealand and in South America have a number of vicariant species (i.e. species
which are closely related but taxonomically separable) which indicate subsequent evolution
after fragmentation and isolation from an originally widely distributed ancestral stock. Thus in
subalpine grasslands in New Zealand and in South America, P. maculata and P. lechleri
respectively occupy similar habitats. Other vicariants between New Zealand and South America
include P. degelii, P. pickeringii, and P. rufovirescens (New Zealand) with P. berberina,
P. flavicans, and P. divulsa in South America.

Australasian element

Twelve species of the New Zealand Pseudocyphellaria flora (25%) also occur in Australia, with
most shared species being found in south-east Australia and Tasmania, indicating a close floristic
and biogeographical relationship of this area with New Zealand. These Australasian taxa are: P.
ardesiaca, P. bartlettii, P. billardierei, P. chloroleuca, P. cinnamomea, P. colensoi, P. coronata,
P. haywardiorum, P. jamesii, P. multifida, P. neglecta, and P. rubella. Of these taxa, five
(P. ardesiaca, P. bartlettii, P. colensoi, P. haywardiorum, and P. rubella) have soredia, and four
(P. chloroleuca, P. coronata, P. multifida and P. neglecta) have isidia or phyllidia.

Apart from P. bartlettii, P. chloroleuca, and P. haywardiorum which are shared between
northern New Zealand and north-eastern Australia, including Lord Howe I., the bulk of the
Australasian species of Pseudocyphellaria are found in rain-forest in Tasmania (Kantvilas et al. ,
1985; Kantvilas & James, 1987) and in Victoria, and in rain-forest and on successional
vegetation in New Zealand, although Tasmania has a more depauperate lichen flora at the
species level than New Zealand (Kantvilas et al. , 1985). For instance, in Pseudocyphellaria only
15 species are known from Tasmania (Kantvilas, 1985; Kantvilas & James, 1987), in comparison
with 48 species from New Zealand.

There are many similarities in the cool-temperate vegetation of New Zealand and ancient
Tasmania, with Gondwanan elements present in both lowland and alpine habitats in both areas
(e.g. Burbidge, 1960; Barlow, 1981; Nelson, 1981; Page & Clifford, 1981; Smith, 1981;
Kirkpatrick & Brown, 1984). Among the cryptogams, Vitt (1979) records five species of moss
common to the Auckland Islands and Tasmania, and Schuster (1982) shows that eight families,
and eight genera of hepatics are shared by New Zealand and Tasmania. Numerous lichens
(Martin, 1965; Bratt, 19760, 19766, 1976c; Bratt & Cashin, 1975, 1976; Galloway, 19856;
Kantvilas et al. , 1985; Kantvilas, 1985) are also common to New Zealand and Tasmania although
the degree of endemism in Tasmanian lichens is low, c. 5% (Kantvilas, 1985; Kantvilas & James,
1987). Many of these disjunct taxa are ancient relicts which evolved in Cretaceous times in
cool-temperate habitats around the Panthalassic margins of Gondwanaland and which became
isolated in cool-temperate habitats on the New Zealand microcontinent and in Tasmania after
rifting and drift consequent upon the opening of the Tasman Sea and Bass Strait. New Zealand
being isolated from West Antarctica for a greater period of time than Tasmania, possibly
experienced a greater variety of more favourable conditions, with opportunities for increased
speciation in cool-temperate habitats, with the result that genera such as Pseudocyphellaria,
Placopsis, and Sphaerophorus evolved more taxa in New Zealand than in Tasmania. In addition

48 D. J. GALLOWAY

to the relict or palaeoaustral element in the lichen floras of Tasmania and New Zealand
(Galloway, 1987; Galloway & J0rgensen, 1987), a distinctive and more recent relation-
ship exists between Australian and New Zealand lichen floras (Galloway, 1979, 1985ft) with
many taxa being transported from sources in Australia to habitats in New Zealand by the
prevailing westerly winds (Raven, 1973; Raven & Raven, 1976; Close et al., 1978; Wardle,
1978).

Lichens disjunct between Tasmania, south-eastern Australia, and New Zealand besides the
species of Pseudocyphellaria already mentioned include Austroblastenia pauciseptata (Sipman,
1983); Bacidia buchananii; Baeomyces arcuatus, B. heteromorphus (Galloway, 1980; Kant-
vilas & James, 1987); Caloplaca cribrosa (Poelt & Pelleter, 1984); Chiodecton colensoi; Cladia
fulginosa, C. inflata, C. retipora, C. sullivanii (Galloway, 1977a; Filson, 1981c); Cladonia
bimberiensis, C. murrayii, C. neozelandica, C. subsubulata, C. weymouthii (Archer, 1985);
Conotremopsis weberiana; Degelia durietzii (Arvidsson & Galloway, 1981); Dendriscocaulon
dendriothamnodes; Ephebe fruticosa; Gymnoderma melacarpum; Hypogymnia billardierei, H.
kosciuskoensis, H. mundata, H. turgidula (Elix, 1979); Knightiella splachnirima (Galloway &
Elix, 1981); Lecanoraflavopallida; Lecidea coromandelica, L. laeta; Megalopsora atrombicans ,
M. campylospora, M. subtuberculosa (Sipman, 1983, 1986); Leioderma amphibolum (Galloway
& J0rgensen, 1987); Menegazzia aeneofusca, M. caliginosa, M. castanea, M. nothofagi, M.
testacea, M. ultralucens; Metus conglomerates, Miltidea ceroplasta (Hafellner, 1984); Neofusce-
lia loxodella; Nephroma australe, N. rufum; Neuropogon acromelanus, N. ciliatus, N. subcapil-
laris (Walker, 1985); Pannaria crenulata (Galloway etal., 1983); Pannoparmelia angustata, P.
wilsonii (Galloway, 1978a); Parmelia signifera, P. tenuirima, P. testacea (Galloway & Elix,
1983); Pertusaria nothofagi; Phyllopsora congregans; Placopsis trachy derma; Protoparmelia
petraeoides (Hertel, 1985); Psoroma asperellum, P. caliginosum, P. durietzii, P. soccatum\
Psoromidium aleuroides (Galloway & James, 1985); Ramalina inflata; Roccellinastrum neglec-
tum (Henssen et al., 1982); Sagenidium molle; Siphula foliacea, S. fragilis; Sphaerophorus
insignis, S. macrocarpus; Stereocaulon caespitosum, S. trachyphloeum (Galloway, 1980ft);
Teloschistes fasciculatus , T. sieberianus, T. velifer, T. xanthorioides; Thelotrema decorticans;
Usnea capillacea, U. contexta, U. torulosa; Wawea fruticulosa (Henssen & Kantvilas, 1985), and
see also Galloway (1985ft).

Austral element

Four species, or c. 8% of the Pseudocyphellaria flora of New Zealand, belong to this element
viz. , P. faveolata, P. glabra, P. granulata, and P. physciospora. The first three taxa occur widely
in southern circumpolar, cool-temperate regions including New Zealand, Tasmania, south-
eastern Australia, southern South America, and the various subantarctic island groups scattered
in the southern oceans, at latitudes of 45S or higher; P. physciospora also occurs in Juan
Fernandez.

Biotas of austral regions have many similarities as well as many disjunct distribution patterns
(e.g. Barlow, 1981; Brundin, 1966, 1975; Cracraft, 1975, 1980; Craw, 1984; Darlington, 1965;
Dawson, 1958; Fleming, 19790, 1979ft; Frankel, 1984; Galloway, 1979, 1987a; Galloway &
J0rgensen, 1987; Godley, 1960, 1975; J0rgensen, 1983; Keast, 1973; Knox, 1979, 1980;
McDowall, 1980; Miller, 1982, Moore, 1972, 1979a, 1979ft; Parsons, 1985; Raven, 1979; Raven
& Axelrod, 1972; Rich, 1975; Schuster, 1969, 1979; South, 1979; Stevens, 1977, 1980a; Vitt,
1979, 1983). Joseph Hooker (1817-1911) was the first to record similarities between lichen floras
of New Zealand, Tasmania, the subantarctic islands, and South America (Hooker & Taylor,
1844; Taylor & Hooker, 1845; Hooker, 1847; Galloway, 1985ft, 1985c), and he was also the
first to offer a possible explanation of disjunct distributions in southern floras. 'I was led to
speculate on the possibility of plants of the Southern Ocean being the remains of a flora that had
once spread over a larger and more continuous tract of land than now exists in that ocean; and
that the peculiar Antarctic genera and species may be vestiges of a flora characterised by the
predominance of plants which are now scattered throughout the southern islands' (Hooker,
1853: xxi). The floristic affinities may be explained in terms of vicariance biogeography (e.g.

PSEUDOCYPHELLARIA 49

Brundin, 1966; Croizat etal., 1974; Craw, 1978; Cracraft, 1975, 1980; Humphries, 1983), or by
panbiogeographic analysis (e.g. Croizat, 1958; Craw, 1985).

All three species of Pseudocyphellaria are epiphytic on trees or shrubs in lowland to montane
forests (P. glabra also occurs widely in subalpine scrub and subalpine to high-alpine grasslands)
characterized by cool, moist, humid conditions such as may well have existed in cool-temperate
regions of the Panthalassic margin of Gondwanaland (Brundin, 1966; Raven & Axelrod, 1972;
Arvidsson & Galloway, 1981; Schuster, 1982; Galloway, 19870; Galloway & J0rgensen, 1987).

Two major groupings are found in austral lichen floras (Galloway, 19870), (1) Palaeoaustral
lichens of which Pseudocyphellaria faveolata is an example. These are thought to represent
primitive Gondwanan groups poorly adapted for transoceanic dispersal, and which derived from
the Cretaceous (or earlier) when the Panthalassic margin of Gondwanaland was available for
colonization, where cool-temperate conditions prevailed, and where a vegetated west Antarctic
could link South American and South Pacific landmasses. Paleaoaustral lichens show a number
of shared features; they grow in cool-temperate environments, often in forests dominated by
species of Nothofagus, or else in subalpine shrubland or grassland habitats; they are often fertile
and generally lack vegetative diaspores; they have been flexible enough to colonize or recolonize
cool-temperate habitats during and after periods of climatic deterioration; and they show
disjunct distributions. (2) Neoaustral lichens , such as Pseudocyphellaria glabra and P. granulata,
are taxa dispersed after the fragmentation of Gondwanaland, mainly post-Pleistocene to the
present. They are generally richly provided with vegetative propagules, (isidia in P. glabra and
soredia in P. granulata} which allow long distance transport via, birds, ocean currents or in the
West Wind Drift (Raven, 1973; Raven & Raven, 1976; McDowall, 1978; Moore, 19796; Close
etal. 1978; Galloway, 1979, 19870).

Palaeotropical element

Palaeotropical taxa, in contrast to southern, cool-temperate taxa, have warm-temperate or
Tethyan affinities and comprise those organisms shared with ancient Tethys (i.e. the vicinity of
the modern Mediterranean) and with its seaway extensions via the Arabian Peninsula, Hima-
layas, south-west Asia, Indonesia, and New Guinea to the islands of the Pacific (Melville, 1966,
1967). The palaeotropical species Pseudocyphellaria argyracea, P. pickeringii, P. dissimilis, and
P. poculifera constitute c. 9% of the New Zealand Pseudocyphellaria flora and range from
Madagascar to New Zealand and the islands of the Pacific with an Indo-Pacific, Pacific Plate
distribution, analogous to some 50 phanerogamic genera (Good, 1974) including Cymbidium
and Nepenthes, and the hepatic Paraschistochila (Schuster, 1979). In New Zealand, palaeo-
tropical species of Pseudocyphellaria are associated throughout parts of their range with an
assemblage of palaeotropical and amphi-Pacific lichens including: Anzia madagascarensis
(Galloway, 19856); Bryoria indonesica (J0rgensen & Galloway, 1983); Cetrelia braunsiana;
Cladonia krempelhuberi; Coccotrema cucurbitula, C. porinopsis; Collema japonicum, C. sub-
conveniens, C. subfragrans; Everniastrum sorocheilum; Heterodermiajaponica, H. microphyl-
la; Hypotrachyna ensifolia, H. thysanota; Laurera cumingii; Leioderma duplicatum, L. erythro-
carpum, L. sorediatum (Galloway & J0rgensen, 1987); Leprocaulon arbuscula; Lobaria retigera
(Galloway, 19816); Lopadium subcoerulescens; Menegazzia eperforata; Pannaria gemmascens\
Peltigera dolichorhiza, P. nana\ Physma byrsaeum; Placopsis cribellans; Pseudocyphellaria
sulphurea (Galloway, 19850); Ramalina celastri; Sphaerophorus murrayi; Sticta sublimbata;
Thelotrema weberi; Thysanothecium scutellatum (Galloway & Bartlett, 1983); Turgidosculum
complicatulum; Usnea maculata, U. societatis; Xanthoparmelia scabrosa (Elix etal. , 1986).

Cosmopolitan element

Three species (P. aurata, P. crocata, P. intricata) or c. 6% of the New Zealand Pseudocyphellar-
ia flora have a cosmopolitan distribution, being found on all major landmasses, as well as on
oceanic islands in both Northern and Southern Hemispheres. All three species are sorediate and
although fertile specimens of all taxa are known, apothecia in P. aurata and P. intricata are
exceedingly rare, and are only occasionally seen in P. crocata. While both P. crocata and P.

50 D. J. GALLOWAY

intricata have a wide distribution in New Zealand, being known from both cool, moist, shaded,
woodland habitats to dry, open, sunny, grassland sites, P. aurata is restricted to northern
coastal habitats north of lat. 37S.

Discussion

Generic concept and infrageneric classification

Between 1790 and 1890, when Vainio described Pseudocyphellaria (Vainio, 1890), many species
now recognized in that genus were originally placed in Sticta or in several other genera including
Crocodia, Nephroma, Parmelia, Parmosticta, Parmostictina, Phaeosticta, Platysma, Pulmonar-
ia, Ricasolia, Saccardoa, or Stictina. Vainio described Pseudocyphellaria for species having
pseudocyphellae on the lower surface of the thallus, a distinction which earlier generic
descriptions had not emphasized. In his descriptions of Pseudocyphellaria and of Sticta, Vainio
(1890) distinguished anatomical structures of cyphellae and pseudocyphellae, differences which
were strengthened over 70 years later by the discovery of a complex medullary chemistry in
species of Pseudocyphellaria, correlating with the presence of pseudocyphellae. Species of Sticta
on the other hand, have a notably depauperate medullary chemistry. For most of the 19th
century and for at least half of the present century, taxa in Pseudocyphellaria were included in
Sticta (see below), but as we have seen, there are fundamental reasons (viz., the presence of
pseudocyphellae, and a richly complex medullary chemistry) for maintaining Pseudocyphellaria
as an independent genus.

Acharius (1803, 1810, 1814) had a wide generic concept of Sticta, including within it species
belonging to Sticta, Pseudocyphellaria, and Lobaria 'subtus liberum villosum: cyphellis, sorediis
vel maculis interspersis', cyphellis being present in Sticta sens, str., sorediis referring to
pseudocyphellae (in Pseudocyphellaria}, and maculis to naked patches on the lower surface
often seen in Lobaria. Acharius included the following taxa referable to Pseudocyphellaria:
Sticta anthraspis, S. aurata, S. crocata, S. obvoluta, and S. orygmaea [=P. berberina] (see
Galloway & James, 1977; Galloway 1986a).

Delise (1825a, b ) also had a wide genus concept of Sticta, his monograph of 78 taxa including
species now accommodated in Erioderma, Lobaria, Pseudocyphellaria, Sticta, and Xanthoma-
culina (Hale, 1985). In his account he included 28 species of Pseudocyphellaria, 23 of which he
described and figured for the first time (Galloway & James, 1986). Fee (1824-1825) described a
new genus, Delisea, from King I., Australia, but later changed this name to Plectocarpon (Fee,
1824-1825; 1828) , since Lamouroux had earlier described Delisea in the Rhodophyta. However,
it was recently shown that Plectocarpon refers to a lichenicolous fungus growing on Pseudocy-
phellaria glabra (Hawksworth & Galloway, 1984).

Link (1833) described a new genus, Crocodia for Sticta aurata, based on the yellow medulla
and pedicellate fruits of that species, and his genus was later taken up by Trevisan (1869) to
accommodate species of Sticta having a green photobiont and brown, 3-septate spores; however,
since then, Crocodia was never used at the generic level by any lichenologist. In a study of
Brazilian lichens, Martius (1833) made Sticta a section of Parmelia; however, numerous new
species of Pseudocyphellaria from explorations in the Pacific and in South America were first
described as species of Sticta (Montagne, 1835, 1845, 1854, 1856; Meyen & Flotow, 1843;
Hooker & Taylor, 1844; Schaerer in Moritzi, 1846; Montagne & v. d. Bosch, 1857).

De Notaris (1851) divided Sticta into three unnamed sections based on colour, structure, and
presence or absence of cyphellae 'cyphellis urceolatis vel soredii-formibus vel papillaribus,
rarius deficientibus praeditus' , his concept encompassing Sticta, Pseudocyphellaria, and Lobaria
respectively, and being followed by Massalongo (1853). Babington (1855) in his treatment of
New Zealand species of Sticta proposed three groupings or sections within the genus: 1.
Chrysosticta Church. Bab., 'cyphellae punctiform, bright golden-yellow' for species of
Pseudocyphellaria, 2. Leucosticta Church. Bab., 'cyphellae white or pale yellow; either punc-
tiform, or explanate, or urceolate', for species of Pseudocyphellaria and Sticta, and 3. Pseudos-

PSEUDOCYPHELLARIA 51

ticta Church. Bab., 'cyphellae wanting altogether, or only occasional', for species of Pseudocy-
phellaria [P. montagnei] and Lobaria.

Nylander (1854) placed Sticta with Parmelia and Physcia in the tribe Parmeliae but the
following year (Nylander, 1855) he refined his concept of the genus to include Sticta and
Ricasolia De Not. (= Lobaria), and within Sticta he proposed a sectional division viz. , 'A. Frons
subtus inaequalibus gibberosus, subnudus' [= Lobaria], 'B. Frons infra cyphellis praedita albis
vel albicantibus' [= Sticta pr. p., Pseudocyphellaria pr. p.], and 'C. Frons cyphellis citrinis'
[= Pseudocyphellaria]. Later (Nylander, 18580: 101-103; 18586: 333) he emended his section
containing white or whitish cyphellae, to comprise two races viz., 'Stirps Stictae fuliginosae' and
'Stirps Stictae damaecornis ' , the former comprising only species of Sticta and the latter species of
Sticta and Pseudocyphellaria. In this account, Nylander gave the geographical distribution of
taxa and also placed Sticta coriacea in Ricasolia, a procedure he adopted in all his subsequent
accounts of New Zealand lichens (Nylander, 18606; 1866; 18886). In 1860, Nylander's most
ambitious and certainly his most detailed work on Sticta appeared in Part II of his Synopsis
(Nylander, 18606). Earlier the same year he described Stictina Nyl., to accommodate species
with a cyanobiont, his sub-tribe Stictei now consisting of three genera, Stictina, Sticta, and
Ricasolia, an arrangement he adopted in all subsequent publications dealing with the Lobar-
iaceae (Nylander, 18610; 18616; 1865; 1866; 18680; 18686; 18880; 18886 ) and which was widely
used by later authors (e.g., Hooker, 1867; Hue, 1890, 1901; Jatta, 1890; Stizenberger, 1895;
Reinke, 1896). Nylander (18600) first introduced the term pseudocyphellae, and he also
described two new genera, Parmosticta Nyl. (Nylander, 1875a), and Parmostictina Nyl.
(Nylander, 18756) for taxa with a thalline exciple and pedicellate fruits, and a green
photobiont, or a cyanobiont respectively. In his Conspectus synopticus Sticteorum (18686)
Nylander proposed sections Pseudocyphellatae Nyl. and Cyphellatae Nyl., in both Sticta and
Stictina, a procedure later followed by Hue (1890, 1901), and by Stizenberger (1895).

Muller Argoviensis (1862) divided Sticta into two sections, Eusticta Mull. Arg. and Stictina
(Nyl.) Mull. Arg., based on the nature of the photobiont, but in his later accounts of New
Zealand lichens (Muller Argoviensis, 1892; 1894, 1896) he followed Nylander's (18600; 18606)
use of Sticta and Stictina as generic names.

Stizenberger (1862) recognized Sticta, Eusticta, and Ricasolia as subgenera of Sticta sens. lat. ,
but in his later account of the worldwide distribution of the family Lobariaceae (Stizenberger,
1895) he adopted Nylander's genera based on the nature of the photobiont, retaining Ricasolia
for species of Lobaria, and with species referable to Pseudocyphellaria divided between Stictina
and Sticta.

Trevisan (1869) used both spore characters, and the nature of the photobiont to divide Sticta
into eight genera; Sticta, Stictina, Saccardoa Trevisan, and Diclasmia Trevisan all having a
cyanobiont, and Ricasolia, Phaeosticta Trevisan, Crocodia, and Lobaria having a green
photobiont. Species now referred to Pseudocyphellaria he included in the following genera:
Stictina pr. p. (spores brown, 3-septate); Saccardoa (spores brown, 1-septate, always pseudo-
cyphellate), Ricasolia pr. p. (spores colourless, 1-septate); Phaeosticta (spores brown, 1-
septate); Crocodia pr. p. (spores brown, 3-septate); or Lobaria pr. p. (spores colourless,
3-9-septate), and although his generic segregates were adopted in the fungal classifications of
Clements (1909) and Clements & Shear (1931), and in the invalid infrageneric classification of
the 'order' Stictales by Choisy (19606), they have not been subsequently used by any other
lichenologists. The use of spore and photobiont characters alone for the delimitation of genera
in the Lobariaceae is unsatisfactory and reflects no basic biological differences, since spore
colour and septation varies with age of thallus and degree of maturation of the spore.

Vainio (1890) proposed a new classification of the Lobariaceae, comprising the genera Sticta,
Lobaria, and a new genus, Pseudocyphellaria Vainio, to accommodate all pseudocyphellate
taxa. Vainio placed as synonyms of Pseudocyphellaria, Nylander's (18686) Sticta sect.
Pseudocyphellatae and Stictina sect. Pseudocyphellatae, and Babington's Sticta sect. Chryso-
sticta and sect. Leucosticta. Within his new genus Vainio recognized four sections, depending on
the nature of the photobiont and also on the presence or absence of a thalline exciple (i.e.
photobiont cells in the excipular tissue). Sects Parmosticta (Nyl.) Vainio and Parmostictina

52 D. J. GALLOWAY

(Nyl.) Vainio, for taxa having a cyanobiont and a green photobiont and a thalline exciple, and
sects Lecidosticta Vainio and Lecidostictina Vainio, for taxa without a thalline exciple and with a
green photobiont and a cyanobiont respectively. Vainio's treatment of the Lobariaceae was
roundly condemned by Miiller Argoviensis (1891: 387) who wrote 'Eigentlich misshandelt und
zerfetzt sind die Sticteen', although he allowed that Vainio was correct in synonymising
Ricasolia with Lobaria. Miiller stressed that all other taxa in the Lobariaceae should be defined
in one of two genera, Sicta or Stictina, depending on the photobiont present.

Stizenberger (1895), although subscribing to Nylander's arrangement of genera in the
Lobariaceae, proposed two new sections, Leucopseudocyphellatae Stizenb. and Xantho-
pseudocyphellatae Stinzenb., within the subgen. Pseudocyphellatae of both Sticta and Stictina,
the sections being defined by the colour of the pseudocyphellae present.

Hellbom (1896) in his account of the New Zealand lichens collected by Sven Berggren in
1874-75, included species of Pseudocyphellaria either in Sticta or Lobaria, the cyanobiont-
containing taxa being placed in Sticta, and green photobiont species in Lobaria, a rather curious
arrangement not subsequently followed by any later author.

In the present century Zahlbruckner, whose views on lichen taxonomy influenced systematic
lichenology for nearly 50 years, reverted to Miiller Argoviensis's (1862) arrangement of genera
in the Lobariaceae, with Sticta including both cyphellate and pseudocyphellate taxa, and being
divided into sections Eusticta and Stictina according to the nature of the photobiont
(Zahlbruckner 1907; 1925; 1926, 1932, 1941). Pseudocyphellaria, although only rarely used in
the late 19th century (Malme, 1899; Vainio, 18980, 18986, 1899), was widely used in the present
century (see for example Degelius, 1935, 1941; Dodge, 1948, 1964, 1965, 1971; Du Rietz, 1924,
1926; Follmann, 1966; Galloway, 1979, 19836, 19850, 19856, 19860, 19866; Galloway & James,
1977, 1980, 1986; Galloway et al., 19836; Gyelnik, 19310, 19316, 1933, 1938; Huneck etal.,
1973; Imshaug, 1950, 1977; Lamb, 1955, 1959; Magnusson, 1940, 1956; Malme, 1934; Magnus-
son & Zahlbruckner, 1943; Martin, 1965, 1966, 19690, 19696, 1970; Martin & Child, 1972;
Rasanen, 1932, 1936, 1937, 1939, 1943, 1944; Renner & Galloway, 1982; Szatala, 1956; Tavares,
1952; Vainio, 1903; Wilkins & James, 1979).

Clements (1909) on the basis of spore colour and septation, and the nature of the photobiont,
extended Trevisan's classification of genera within the Lobariaceae to include the following
additional pseudocyphellate genera: Diphaeosticta Clem., Diphanosticta Clem., Merostictina
Clem., and Podostictina Clem. These genera, although recorded in Clements & Shear (1931),
were never subsequently used by later lichenologists.

Gyelnik (19310) emended Pseudocyphellaria to refer only to taxa with a green photobiont,
and he proposed two new sections, Albidopseudocyphellaria Gyelnik and Flavopseudocyphel-
laria Gyelnik, for species with white, and with yellow pseudocyphellae respectively. At the same
time, cyanobiont-containing taxa formerly included in Pseudocyphellaria were referred to a new
genus, Cyanisticta Gyelnik, comprising sections Eucyanosticta Gyelnik, for taxa with white
pseudocyphellae, and Cyanostictina Gyelnik, for taxa with yellow pseudocyphellae. Cyanisticta,
although used by several authors (Dodge, 1964, 1965, 1971; Gyelnik, 19316, 1931c, 1932, 1938;
Rasanen, 1932, 1933, 1936, 1937, 1940, 1943, 1944; Szatala, 1956) is correctly included as a
synonym of Pseudocyphellaria.

Rasanen (1943) proposed a classification of pseudocyphellate taxa in the Lobariaceae
comprising Pseudocyphellaria (Vainio) Gyelnik, with two sections, Parmosticta (Nyl.)
Rasanen, and Lecidosticta (Nyl.) Rasanen for species with a green photobiont and with or
without a thalline exciple, and Cyanisticta and the two corresponding sections Parmostictina
(Nyl.) Rasanen and Lecidostictina (Nyl.) Rasanen, for species with cyanobionts, and with or
without a thalline exciple.

Magnusson divided Pseudocyphellaria into just two sections, Leucopseudocyphellatae
(Stizenb.) Magnusson and Xanthopseudocyphellatae (Stizenb.) Magnusson, on the basis of the
colour of pseudocyphellae (Magnusson & Zahlbruckner, 1943; Magnusson, 1956).

The most recent infrageneric arrangement of Pseudocyphellaria appears as part of a rather
curious classification of what is called 'Les Stictales' (Choisy, 19600, 19606). Choisy's order
Stictales contains four genera, Heterodea Nyl., Sticta, Knightiella Mull. Arg., and Nephroma

PSEUDOCYPHELLARIA 53

Ach. The genus Sticta is divided into five subgenera viz., Podosticta Choisy, Crocodia (Link.)
Choisy, Cyphellaria Choisy, Pseudocyphellaria (Vainio) Choisy, and Lobaria (Schreber)
Choisy, with taxa referable to Pseudocyphellaria occurring in subgenera Podosticta, Crocodia,
and Pseudocyphellaria. Subgen. Podosticta accommodates stalked taxa of both Sticta and
Pseudocyphellaria and has two sections distinguished by the photobiont present, sect. Eupodo-
sticta Choisy, having a green photobiont, and sect. Podostictina (Clem.) Choisy, based on Sticta
endochrysoides (Mull. Arg.) Hue [= Pseudocyphellaria compar (Nyl.) Magnusson)] with a
cyanobiont. Subgen. Crocodia, for yellow-medulla species has three sections, sect. Flavo-
pseudocyphellaria (Gyelnik) Choisy, and sect. Hypocrocodia Choisy [for specimens of Lobaria]
having a green photobiont, and sect. Saccardoa (Trevisan) Choisy with a cyanobiont. Subgen.
Pseudocyphellaria, Choisy divided into two sections; sect. Albidopseudocyphellaria (Gyelnik)
Choisy, without a thalline exciple [farther divided into subsect. Lecidosticta (Vainio) Choisy,
with a green photobiont, and subsect. Lecidostictina with a cyanobiont] and Sect. Parmosticta
(Nyl.) Choisy with a thalline exciple and comprising two subsections; subsect. Delisea (Fee)
Choisy with a green photobiont [further divided into Stirps Phaeosticta (Trevisan) Choisy, with
brown spores, and Stirps Phanosticta (Clem.) Choisy, with colourless or pale spores] and
subsect. Parmostictina (Nyl.) Choisy with a cyanobiont. All of Choisy 's new names are invalid
since they are not provided with Latin descriptions, and his new combinations are also invalid
(Art. 33.2) since no basionyms are cited.

Presently it is not considered opportune to elaborate a satisfactory modern infrageneric
classification of Pseudocyphellaria reflecting evolutionary trends in the genus based on detailed
comparisons of anatomy, morphology, chemistry, and biogeography, and pending further
studies on Australian, South American, palaeotropical, and tropical taxa, no formal infragen-
eric arrangement is adopted in the present work. It is hoped, however, that in the near future an
acceptable infrageneric classification will be formulated, together with a synoptic key to all
known species of the genus.

The genus Pseudocyphellaria

Pseudocyphellaria Vainio

in Acta Soc. Fauna Flora fenn. 7: 182 (1890). - Sticta subgen. Pseudocyphellaria (Vainio) M. Choisy in

Bull. mens. Soc. linn. Lyon 29: 124 (1960) comb, inval. (Art. 33.2). Type: Pseudocyphellaria aurata

(Ach.) Vainio.
Crocodia Link, Handbuch 3: 177 (1833). - Sticta subgen. Crocodia (Link) M. Choisy in Bull. mens. Soc.

linn. Lyon 29: 123 (1960) comb, inval. (Art 33.2). Type: Crocodia aurata (Ach.) Link.
Stictina Nyl., Syn. meth. lich. 1(2): 333 (1860). - Sticta sect. Stictina (Nyl.) Mull. Arg. in Mem. Soc. Phys.

Hist. not. Geneve 16: 371 (1862). Lectotype: Stictina crocata (L.) Nyl. [selected by Clements & Shear

(1931: 322)].
Phaeosticta Trevisan, Lichenotheca veneta exs. 75 (1869). - Sticta subgen. Pseudocyphellaria sect.

Parmosticta, subsect. Delisea stirps Phaeosticta (Trevisan) M. Choisy in Bull. mens. Soc. linn. Lyon 29:

125 (1960) comb, inval. (Art. 33.2). Lectotype: Phaeosticta physciospora (Nyl.) Trevisan [selected by

Choisy (1960: 125)].
Saccardoa Trevisan, Lichenotheca veneta exs. 75 (1869). - Sticta subgen. Crocodia sect. Saccardoa

(Trevisan) M. Choisy in Bull. mens. Soc. linn. Lyon 29: 123 (1960) comb, inval. (Art. 33.2). Lectotype:

Saccardoa crocata (L.) Trevisan [selected by Choisy (1960: 123)].
Parmosticta Nyl. in Flora, Jena 58: 303 (1875), nom. superfl. (Art. 63.1). - Pseudocyphellaria sect.

Parmosticta (Nyl.) Vainio in Acta Soc. Fauna Flora fenn. 7: 183 (1890). -St icta sect. Parmosticta (Nyl.)

M. Choisy in Bull. mens. Soc. linn. Lyon 29: 125 (1960) comb, inval. (Art. 33.2). Type: Parmosticta

aurata (Ach.) Nyl.
Parmostictina Nyl. in Flora, Jena 58: 363 (1875). - Pseudocyphellaria sect. Parmostictina (Nyl.) Vainio in

Acta Soc. Fauna Flora fenn. 7: 183 (1890). - Cyanisticta sect. Parmostictina (Nyl.) Rasanen in Acta hot.

fenn. 33: 12 (1943). - Sticta subgen. Pseudocyphellaria sect. Parmosticta subsect. Parmostictina (Nyl.)

M. Choisy in Bull. mens. Soc. linn. Lyon 29: 125 (1960) comb, inval. (Art. 33.2). Type: Sticta hirsuta

Mont.
Diphaeosticta Clements, Gen.fung.: 81 (1909). Type: Diphaeosticta physciospora (Nyl.) Clements (1909:

175).

54 D. J. GALLOWAY

Diphanosticta Clements, Gen. fung.: 81 (1909). Type: Diphanosticta cellulifera (J. D. Hook. & Taylor)

Clements (1909: 175).
Merostictina Clements, Gen. fung.: 82 (1909). Type: Merostictina mougeotiana (Delise) Clements (1909:

175).
Phanosticta Clements, Gen. fung. : 81 (1909). - Sticta subgen. Pseudocyphellaria sect. Parmosticta subsect.

Delisea stirps Phanosticta (Clements) M. Choisy in Bull. mens. Soc. linn. Lyon 29: 125 (1960) comb.

inval. (Art. 33.2). Type: Phanosticta freycinetii (Delise) Clements (1909: 175).
Podostictina Clements, Gen. fung. : 82 (1909). -Sticta subgen. Podosticta sect. Podostictina (Clements) M.

Choisy in Bull. mens. Soc. linn. Lyon 29: 126 (1960) comb, inval. (Art. 33.2). Type: Podostictina

endochrysoides (Mull. Arg.) Clements (1909: 175).
Cyanisticta Gyelnik in Feddes Reprium. Spec. nov. veg. 29: 1 (1931). - Pseudocyphellaria sect. Cyanisticta

(Gyelnik) Lamb, in An. Parq. nac. B. Aires 7: 67 (1959) ['1958']. Lectotype: C. argyracea (Delise)

Gyelnik [selected by Dodge (1964: 168)].

Thallus heteromerous, foliose, dorsiventral, lobate, often very large, orbicular, rosette-forming
or irregularly laciniate, or dichotomously branched, to complexly entangled, rarely
monophyllous, loosely to closely attached. Lobes very variable, narrow, strap-shaped,
dichotomously branching with distinctly bifurcating apices which may be pointed, blunt, or
rounded, to complex, imbricate. Margins entire or variously incised, notched or folded, often
conspicuously thickened especially below, often with conspicuous, elongate or verruciform-
conical or glomerulate pseudocyphellae, often free and ascending, thick and coriaceous to thin
and fragile. Upper surface smooth, wrinkled, scabrid or hairy, often deeply or shallowly
faveolate, with faint or strongly marked reticulum of interconnecting ridges, ridges shallowly
rounded to sharply delimited, shining, matt or dull, often conspicuously maculate (x 10 lens,
best seen when moist), with or without isidia, pseudoisidia, phyllidia, pseudocyphellae or
soredia. Medulla white or yellow. Photobiont green (Dictyochloropsis or C7i/ore//0-like) or
blue-green (Nostoc). Internal cephalodia containing Nostoc present in taxa with a green
photobiont. Lower surface glabrous in some taxa but usually tomentose, tomentum pale to
dark, thick and felted to indistinctly pubescent. Pseudocyphellae always present, white or
yellow, sparse to frequent, immersed in tomentum, plane and fleck-like to raised-conical,
verruciform or glomerulate, round to irregular, margins distinct or indistinct.

Conidiomata pycnidial (Lobaria-type) , immersed, globose, visible as slight swellings on
upper surface and occasionally on lower surface, ostiole dark red-brown to black, plane,
elevated or punctate-depressed, randomly scattered or in lines at margins and on thalline ridges,
most common towards lobe apices, often eroding and leaving gaping pits. Conidiogenous cells
colourless, cuboid, phialidic, on simple to slightly branched conidiophores, bearing conidia
laterally and terminally. Conidia colourless, short, baccilliform to slightly dumbbell-shaped.

Ascomata apothecioid, hemiangiocarpic, sessile to pedicellate, laminal or marginal, rounded,
margins entire or crenate-striate to stellate, with or without phyllidia, isidia or soredia, disc
matt or shining, sometimes white-pruinose, thalline exciple in taxa with pedicellate ascomata,
smooth to verrucose or areolate-scabrid, hairy, sorediate or maculate; proper exciple in taxa
with sessile ascomata, smooth to verrucose or areolate-scabrid or tomentose. Ascomata
ascohymenial, hymenium 1+ blue, hamathecium of simple, septate, filiform paraphyses,
swollen, occasionally moniliform, and often pigmented at apices, epithecium present or
absent, often pigmented and becoming coloured in K. Hypothecium usually pigmented,
sometimes changing colour in K. Asci shorter than paraphyses, cylindrical or clavate, bitunicate,
apex with an 1+ blue ring, dehiscence semifissitunicate, the ring everting. Ascospores 8 per
ascus, uniseriately or biseriately arranged in ascus, colourless or yellow-brown to brown,
fusiform-ellipsoid to broadly ellipsoid, rarely acicular-fusiform, apices rounded or pointed,
simple at first becoming 1-3 (rarely to 5- or 7-)-septate at maturity, sometimes 2-celled spores
have strongly thickened walls with a narrow central canal connecting the locules.

Chemistry: diverse, containing depsides, depsidones, terphenylquinones, pulvinic acid
derivatives and triterpenoids of hopane, fernene, stictane or lupane series.

PSEUDOCYPHELLARIA 55

Key to New Zealand species

la Medulla white 2

Ib Medulla yellow 39

2a (la) Photobiont green 3

2b Photobiont blue-green 22

3a (2a) Upper surface pseudocyphellate 4

3b Upper surface without pseudocyphellae 7

4a (3a) Lobe margins isidiate or phyllidiate 5

4b Lobe margins entire, isidia or phyllidia absent 16

5a (4a) Phyllidia mainly marginal, rarely or never associated with laminal pseudocyphellae;

epithecium granular, K + rose pink; spores (27-)32-36(-43) x 7-9 um

20. P. episticta (p. 130)

5b Isidia terete, marginal and laminal, often associated with laminal pseudocyphellae, at
length forming phyllidia at margins; epithecium opaque, yellow-brown, unchanged in
K;spores27-34 x l-9\im 48. P. wilkinsii (p. 242)

6a (4b) Lower surface pale buff or whitish; epithecium granular, K -I- rose pink; spores 30-40 x

7-9 urn 32. P. lindsayi (p. 180)

6b Lower surface dark brown; epithecium opaque, yellow-brown, unchanged in K; spores

25-30 x 7-9nm 33. P. lividofusca (p. 183)

7a (3b) Sorediate 24. P. granulata (p. 152)

7b Esorediate 8

8a (7a) Upper surface tomentose or pubescent, or at least at margins 9

8b Upper surface and margins glabrous 12

9a (8a) Tomentum on upper surface continuous from margins to centre 10

9b Tomentum or pubescence marginal 11

lOa (9a) Lobe margins entire ; pseudocyphellae yellow ; with a well-defined chemistry

44. P. pubescens (p. 228)

lOb Lobe margins phyllidiate or granular-isidiate; pseudocyphellae white; without demon-
strable chemistry 25. P. gretae (p. 156)

lla (9b) Lobe margins entire 12. P. coriacae (p. 102)

lib Lobe margins with minutely pubescent phyllidia 21. P. fimbriata (p. 139)

12a (8b) Lobe margins phyllidiate or isidiate 13

12b Lobe margins entire 17

13a(12a) Lobe margins phyllidiate 14

13b Lobe margins isidiate 16

14a(13a) Lobes rounded; phyllidia dentate-squamiform 15

14b Lobes entangled-imbricate, narrow, linear-laciniate; phyllidia finger-like to coralloid

37. P. multifida (p. 199)

15a (14a) Lower surface wrinkled-bullate or papillate, thinly tomentose; pseudocyphellae incon-
spicuous ; cortex C -I- red (fugitive) 36. P. montagnei (p. 195)

15b Lower surface plane or shallowly wrinkled, densely tomentose; pseudocyphellae

conspicuous; cortex C- 11. P. corbettii (p. 99)

16a(13b) Lower surface glabrous, shining, pale pinkish white; isidia coralloid; medulla C + rose

(fugitive) 8. P. chloroleuca (p. 85)

16b Lower surface densely tomentose, chocolate brown or blackish; isidia simple; medulla

C- 23. P. glabra (p. 146)

17a(12b) Upper surface plane to undulate 27. P. homoeophylla (p. 162)

17b Upper surface faveolate-impressed 18

18a(17a) Pseudocyphellae yellow

18b Pseudocyphellae white 20

56

D. J. GALLOWAY

19a(18a) Lobes elongate; margins with prominent pseudocyphellae 7. P. carpoloma (p. 80)

19b Lobes short, rounded; margins entire, without pseudocyphellae

41. P. physciospora (p. 213)

20a (18b) Lower surface pale , tomentum thin or sparse 21

20b Lower surface dark, tomentum thick 22

21a(20a) Lobes elongate, dichotomously branched, shallowly faveolate; apothecial disc red-
brown; cortex C- 46. P. rufovirescens (p. 235)

21b Lobes rounded, deeply faveolate ; apothecial disc black ; cortex C + red

18. P. durietzii (p. 126)

22a(20b) Lobes variable, often irregular; margins often indented, with projecting verruciform
white pseudocyphellae, not or rarely ridged below; apothecial disc often white-
pruinose; medulla containing physciosporin 20. P. faveolata (p. 134)

22b Lobes rather narrow, dichotomously branched; margins entire, smoothly rounded,
ridged below, without pseudocyphellae; apothecial disc never pruinose; medulla
without physciosporin 6. P. billardierei (p. 75)

23a (2b) Upper surface pseudocyphellate 24

23b Upper surface without pseudocyphellae 25

24a(23a) Pseudocyphellae developing isidia at margins 3. P. argyracea(p. 64)

24b Pseudocyphellae not isidiate 31. P. knightii (p. 177)

25a(23b) Pseudocyphellae white 26

25b Pseudocyphellae yellow 34

26a(25a) Sorediate 27

26b Esorediate 29

27a(26a) Upper surface plane or undulate, emaculate; pseudocyphellae of lower surface rare or

absent 29. P. intricata (p. 169)

27b Upper surface faveolate-reticulate or punctate-impressed, white-maculate (x 10

lens) ; pseudocyphellae prominent below 28

28a(27b) Cortex C + red; medulla K + yellow; upper surface reticulate-faveolate; pseudocyphel-
lae below minute, punctiform 5. P. bartlettii (p. 72)

28b Cortex C ; medulla K ; upper surface punctate-impressed; pseudocyphellae below

conspicuous, large, plane 26. P. haywardiorum (p. 159)

29a(26b) Lobe margins tomentose or pubescent , 1. P. ullunii (p. 57)

29b Lobe margins glabrous 30

30a(29b) Isidiate or phyllidiate 31

30b Without isidia or phyllidia 32

31a(30a) Isidia mainly terete, glabrous, occasionally dorsiventral 17. P. dissimilis (p. 122)

31b Phyllidia pubescent 22. P. fimbriatoides (p. 142)

32a(30b) Upper surface plane ; lobes narrow , subcanaliculate ; lower surface strongly costate

9. P. cinnamomea (p. 91)
32b Upper surface f aveolate-impressed ; lobes not subcanaliculate ; lower surface not costate 33

33a(32b) Lobes rounded, deeply faveolate; cortex C + red 28. P. hookeri (p. 166)

33b Lobes elongate , dichotomously branched , shallowly faveolate ; cortex C

38. P. murrayi (p. 204)

34a (25b) Upper surface tomentose 35

34b Upper surface glabrous 36

35a(34a) Upper surface densely and uniformly tomentose; margins tomentose; pseudocyphellae

of lower surface sunk in tomentum 35. P. margaretiae (p. 191)

35b Upper surface tomentose centrally; margins glabrous, shining; pseudocyphellae of lower

surface conical-verruciform, projecting above tomentum 47. P. sericeofulva (p. 239)

36a(34b) Lobe margins phyllidiate 39. P. neglecta (p. 207)

36b Lobe margins entire 37

PSEUDOCYPHELLARIA

57

37a(36b) Sorediate 15. P. crocata (p. 113)

37b Esorediate 38

38a(37b) Upper surface maculate (x 10 lens); lobes thin; tomentum pale; mainly terricolous in

subalpine grassland, occasionally corticolous 34. P. maculata (p. 187)

38b Upper surface emaculate ; lobes thick ; tomentum dark ; corticolous , never terricolous ....

14. P. crassa (p. 108)

39a (Ib) Photobiont green 40

39b Photobiont blue-green 46

40a (39a) Without soredia, isidia, or phyllidia 16. P. degelii (p. 1 18)

40b Sorediate, isidiate, or phyllidiate 41

41a(40b) Soredia well-defined 42

41b Soredia absent, isidiate or phyllidiate 44

42a(41a) Soralia marginal; upper surface mainly glabrous 43

42b Soralia laminal; upper surface tomentose 45. P. rubella (p. 231)

43a(42a) Soralia linear, confluent, labriform 4. P. aurata (p. 68)

43b Soralia derived from small, marginal, crowded isidia 43. P. poculifera (p. 224)

44a(41b) Phyllidiate 45

44b Isidiate 10. P. colensoi (p. 95)

45a(44a) Lobes linear-laciniate to irregular; upper surface smooth, plane; apothecial exciple

translucent , verrucose-scabrid ; acetone extract yellow 42. P. pickeringii (p . 218)

45b Lobes broadly rounded; upper surface faveolate; apothecial exciple concolorous with

thallus, coronate, dentate-phyllidiate; acetone extract red-purple... 13. P. coronata (p. 106)

46a(39b) Sorediate or phyllidiate 47

46b Without soredia or phyllidia 30. P. jamesii (p. 174)

47a(46a) Sorediate 2. P. ardesiaca (p. 61)

47b Phyllidiate 40. P. nermula (p. 210)

The species

1. PseudocyphellariaallaniiD. Galloway

Fig. 15

in Mycotaxon 16: 202 (1982). Type: New Zealand, South I., Canterbury, Mt Peel, on track to Emily
Falls, Peel Forest, on fallen Pseudowintera on bank of stream, 20 March 1979, D. J. Galloway (CHR
343256! - holotype; BM! - isotype).

Morphology: Thallus orbicular to spreading, in entangled clones, 8-15(-20) cm diam.,
corticolous, loosely to closely adnate centrally, margins free. Lobes linear-elongate, (3-)5-
8(-14) mm wide, 1-5-6 cm long, attenuating at apices, apices blunt, rounded or furcate, plane
or subconvex centrally, often canaliculate towards apices, subdichotomously branched,
discrete, sinuses prominent, semicircular. Margins entire, sinuous, conspicuously ridged and
thickened below, occasionally with conspicuous, round, white pseudocyphellae, often
tomentose. Upper surface dark leaden grey or grey-blue suffused brownish at margins when
moist, pale greyish fawn tinged brownish or reddish at margins when dry, coriaceous, smooth,
matt or minutely wrinkled or pitted in parts, minutely areolate-scabrid near margins, also
occasionally patchily white-tomentose towards lobe apices, soredia, isidia, maculae, phyllidia,
and pseudocyphellae absent. Medulla white. Photobiont Nostoc. Lower surface uniformly
thickly tomentose from margins to centre, tomentum dense, silky, entangled, occasion-
ally patchily glabrous centrally, pale yellowish buff at margins, dark chocolate brown, grey-
brown to blackened centrally. Pseudocyphellae white, conspicuous, scattered, round
to irregular, 0-5-2-5 mm diam., plane to concave with a distinctive raised, narrow rim,

58

D. J. GALLOWAY

Fig. 15 Pseudocyphellaria allanii. Holotype (CHR). A. Scale = 1 cm. B. Scale = 1 mm.

concolorous with lower surface, often sunk in tomentum. Apothecia not seen. Pycnidia
occasional to frequent, at margins and scattered over thallus, singly or in groups, ostiole
punctate-depressed, dark red-brown to black, visible on lower surface as hemispherical
swellings, 1-1-5 mm diam.

Anatomy: Thallus 230-410 |im thick. Upper cortex 65-80 |im thick, cells 9-14 ^im diam. , upper
15 um straw-yellow, cells compressed, inner 50-65 um more loosely arranged, cells colourless.
Photobiont layer 90-140 um thick, photobiont Nostoc, in chains, tightly to loosely clustered,
surrounded by a thin gelatinous envelope. Medulla 60-150 urn thick, hyphae 3-4 um diam.
Lower cortex 30-40 um thick, colourless to pale straw at outer surface, cells thin-walled 2-5-11-5
um diam. Tomental hairs colourless, 30-230 um long, in dense clusters.

PSEUDOCYPHELLARIA 59

Chemistry: 7(3-acetoxyhopan-22-ol, hopane-7|3, 22-diol(tr.), hopane-15a, 22-diol.

Pseudocyphellaria allanii is named for Dr H. H. B. Allan (1882-1957), first Director of Botany
Division, D.S.I.R., who in the 1920s described the vegetation of Mt Peel and later promoted
interest in New Zealand lichens (Galloway, 1976).

Distinguishing features: Pseudocyphellaria allanii is an endemic species having linear-elongate
lobes, and a coriaceous to patchily tomentose upper surface, especially towards apices and
margins, that is devoid of isidia, phyllidia, pseudocyphellae or soredia. It has a white medulla
and a blue-green photobiont, and a densely tomentose lower surface with conspicuous white
pseudocyphellae with a distinct, raised, narrow margin at maturity and which resemble true
cyphellae. It has a two-hopane chemistry [code A of Wilkins & James (1979)]. It is not known
fertile, though small, laminal pycnidia are occasionally produced.

Variation: Pseudocyphellaria allanii shows little variation in lobe morphology and appears in
the field to be a rather constant species. It occurs rather spasmodically throughout both main
islands of New Zealand and is nowhere a common species. It is most closely related to P.
coriacea and in several localities forms photosymbiodemes with this species (Renner &
Galloway, 1982), the chlorosymbiodeme P. coriacea developing only on the lower surface and
along the margins of the parent cyanosymbiodeme. P. allanii is distinguished from P. coriacea, a
much more widespread and common lichen in New Zealand, only in the nature of the
photobiont.

Pseudocyphellaria allanii is readily distinguished from two other endemic non-glabrous
species of Pseudocyphellaria with blue-green photobionts: (i) P. margaretiae D. Galloway, has a
completely tomentose upper surface, broader, more rounded, and imbricate lobes, yellow
pseudocyphellae on the lower surface and a more complex chemistry (Galloway et al., 19836),
(ii) P. fimbriatoides D. Galloway & P. James, has lobes with marginal, tomentose phyllidia, and
a much less thickly tomentose lower surface; indeed in many specimens tomentum in this species
is greatly reduced or may be often absent altogether. It does however have the same chemistry
as P. allanii.

Distribution: (Fig. 16) Pseudocyphellaria allanii is nowhere a common species in New Zealand
and is known from Great Barrier I., (Hayward et al., 1986), South Auckland (Kuratau, West
Taupo) eastwards to Kuripapango and Lake Waikaremoana, and south to north-west Nelson
(Mt Glasgow), South Westland, and South Canterbury (Peel Forest). It has an altitudinal range
from near sea-level to subalpine forest at 550 m.

Habitat ecology: Pseudocyphellaria allanii is a corticolous species found on the bark of the
following phorophytes: Griselinia littoralis, Kunzea ericoides, Myrsine australis, and Pseudo-
wintera colorata, in lowland to subalpine forest. It is a species of medium to high-light regimes in
standing forest and is found on tree trunks, but not on canopy branches, growing with other
lichens and with bryophytes in cool, moist, humid habitats. It is still a rather poorly collected
species and is rarely a common or obvious plant, although on a sheltered bench on the eastern
slopes of Mt Peel traversed by Allan's Track, a stand of Myrsine australis has photosymbiodemes
of P. allanii and P. coriacea as common epiphytes. The two taxa are indistinguishable in the
dried state, but when wet the dark grey-blue thalli of P. allanii are strikingly apparent. It has also
been collected from roadside scrub in South Westland (Fox Bridge).

Specimens examined: North Island. South Auckland: Kuratau, West Taupo, /. K. Bartlett
(BM, Herb. Bartlett). Gisborne: Lake House, Waikaremoana, 3-4 May 1966, A. E. Wade
(BM). Hawkes Bay: Kuripapango, 23 August 1981, /. K. Bartlett (BM, Herb. Bartlett). South
Island. Nelson: Mt Glasgow track, /. K. Bartlett 245l4d (Herb. Bartlett). Westland: Fox
Bridge, 7 September 1982, /. K. Bartlett 19708 (Herb. Bartlett). Canterbury: Mt Peel, 20, 21
March 1979, D. J. Galloway (CHR 343256, CHR 343125).

60

D. J. GALLOWAY

Lord Howe I

Norfolk I

Three Kings Is

Kermadec 30-
Is

The Snares

Auckland Is

\

Campbell

N

Macquarie I

160

-40

-42

-44

46

166

i

170

Bounty Is

\

Antipodes Is

168

50 l

180

172

174

176

178

170

L_

172

174

176

I

34-

40-

42"

Fig. 16 Distribution of Pseudocyphellaria allanii.

PSEUDOCYPHELLARIA 61

2. Pseudocyphellaria ardesiaca D . Galloway Fig. 17

in Lichenologist 15: 141 (1983). Type: New Zealand, South I., Canterbury, Boyle River near Lewis
Pass, on bark ofLeptospermum, on terrace overlooking Boyle Lodge, 9 January 1979, D. J. Galloway
(CHR 343237! - holotype; BM! - isotype).

Morphology: Thallus orbicular, rosette-forming to irregularly spreading, 2-8(-10) cm
diam., closely attached over whole of lower surface, free at margins, corticolous. Lobes
rounded, imbricate-convolute, 4-10 mm wide, rarely to 20 mm wide at centre, usually rather
short, convex, rarely elongate-laciniate and discrete from margins to centre. Margins entire or
shallowly notched or incised, slightly thickened below, sinuous, sometimes downrolled,
usually conspicuously sorediate, soralia linear to irregular, sometimes eroding back part of
lower surface. Upper surface dark slate blue to navy blue tinged yellowish at margins and ridges
when wet, glaucous-yellow or pale yellowish buff with a pale grey cast when dry, undulate,
smooth, pitted or furrowed to plicate-faveolate in parts, shining to minutely scabrid-areolate
in parts, occasionally sorediate. Soralia linear to irregular at margins, scattered, rounded,
erose on lamina, soredia yellow to blackened, coarsely granular to pseudoisidiate and
somewhat gnarled-coralloid, densely clustered in marginal soralia, sometimes parasitized by a
fungus and appearing pale to dark red-brown and corticate. Maculae common on upper
surface, best seen when thallus is wet (x 10 lens) forming a mottled or marbled pattern of
yellow photobiont-free areas on blue upper surface, conspicuous at margins and near thalline
ridges. Medulla yellow. Photobiont Nostoc. Lower surface pale yellow-brown at margins,
darkening and often blackened centrally, tomentose to margins, tomentum short, velvety
at margins, woolly, densely squarrose-entangled centrally, pale buff or whitish at margins,
brown-black to black centrally, central areas sometimes glabrous. Pseudocyphellae yellow,
scattered, rather sparse at margins, more common centrally, sunk in tomentum near margins,
very slightly raised above tomentum centrally, round, ulcerose, 0-05-0-3 mm diam. , decorticate
area flat or very slightly convex. Apothecia. Not seen.

Anatomy: Thallus 180-230 um thick. Upper cortex 34-45(-50) um thick, uppermost 10-15 um
(2-3 rows of cells) pale straw-coloured, cells round to irregular, 4-7 um diam. , rather flattened
compressed, 2-3 rows of cells abutting photobiont layer formed of much larger, more loosely
arranged colourless cells 9-15-5 um diam. , with prominent air spaces between cells. Photobiont
layer (25-)45-60(-70) um thick, photobiont Nostoc. Medulla 45-75 um thick, hyphae to 4-5 um
thick, encrusted with yellow crystals. Lower cortex (15-)20-27 um thick, cells 4-7 um diam.
Tomental hairs developing as budding extensions of cells of lower cortex, at first 5-7 (-12) cells,
then in fascicles of hyphae (10-12 together) 158-230(-270) um long, hyphae 5-7 um thick, walls
1-1-5 um thick, conspicuously septate, colourless.

Chemistry: pulvinic acid, pulvinic dilactone, calycin, 2a, 3|3, 22a-triacetoxystictane, 2a,
3p-diacetoxystictane-22-ol, stictane-3(3, 22a-diol(tr.), 2a-acetoxystictane-3(3, 22a-diol, 3(3-
acetoxystictane-2a, 22a-diol, stictane-2a, 3(3, 22a-triol.

Distinguishing features: Pseudocyphellaria ardesiaca is an Australasian species having
closely attached, orbicular or rosette-shaped thalli with short, rounded, convex, somewhat
scalloped lobes with sinuous, slightly ascending margins which are often conspicuously soredi-
ate. The upper surface is smooth, occasionally areolate-scabrid at the margins, to often
complexly rugose centrally and characteristically dark slate blue when moist, often with a
conspicuous reticulum of pale yellowish maculae (xlO lens). Yellow granular soredia are
present in linear, sinuous, marginal soralia or scattered on the upper surface (especially towards
the lobe apices) as small, ulcerose spots. The medulla is golden yellow with no tendency towards
pale or whitish areas, and the photobiont blue-green. The lower surface is pale yellow or
brownish with a short, even, rather velvety tomentum, pale yellow or buff at margins, greyish
centrally and has scattered yellow pseudocyphellae, either sunk in the tomentum or convex
and raised above the tomentum. Apothecia are not known.

Variation: Pseudocyphellaria ardesiaca shows rather little morphological variation and is

62

D. J. GALLOWAY

Fig. 17 Pseudocyphellaria ardesiaca. Holotype (CHR). A. Scale = 1 cm. B. Scale = 2 mm.

usually a tightly adnate, rosette-forming, corticolous species with a degree of variation in the
position and density of soredial development. Certain specimens seen have only rather small
and widely scattered, discrete marginal soredia, while others may have much or all of the lobe
margins copiously sorediate. In most specimens seen, laminal soralia are not especially
common, being most obvious towards apices of lobes and only rarely and exceptionally is the
upper surface densely covered in soredia. The presence of soredia in P. ardesiaca distinguishes it
from two related species endemic to New Zealand, P. jamesii which is non-sorediate and P.
nermula which has marginal phyllidia. P. jamesii has, in addition, rather narrower, more linear
lobes than either P. ardesiaca or P. nermula, and the surface of its lobes is often reticulate-
faveolate.

Three southern South American taxa are similar to P. ardesiaca but are all taxonomically
distinct from it. A vicariant South American sorediate species is very similar in many respects to
P. ardesiaca but it has thinner, more papery lobes which are larger (8-15 cm diam.), more
loosely attached to the substrate, irregularly spreading, linear-laciniate to rather ragged, and not

PSEUDOCYPHELLARIA

63

Fig. 18 Distribution of Pseudocyphellaria ardesiaca.

64 D. J. GALLOWAY

rosette-forming, and with more highly dissected margins. Its upper surface tends to be more
distinctly scrobiculate-faveolate, not merely irregularly rugose as in P. ardesiaca, and it has
prominent and often thickly scattered laminal soralia, with linear, marginal soralia either absent
or poorly developed. Its medulla is pale yellow-white rather than uniformly golden yellow as in
P. ardesiaca and its lower surface is distinctly wrinkled-subbullate with a thin, fine, rather sparse
tomentum, and rather sparse, inconspicuous yellow pseudocyphellae (punctate to plane, 0-1
mm diam. or less). P. scabrosa R. Sant. is distinguished from P. ardesiaca by its characteristic
scabrid-areolate upper surface and the somewhat glomerulate laminal soralia, and P.
piloselloides (Rasanen) H. Magnusson differs from P. ardesiaca in the longer, ragged, linear-
lacinate lobes, the upper surface of which are distinctly reticulate-faveolate, and continuously
to patchily tomentose (Galloway, 1986a).

Distribution: (Fig. 18) From the Mangaotaki River catchment near Pio Pio (South Auckland)
eastwards to Lake Waikaremoana, and south to the Tararua Range in North I., and from the
mountains of north-west Nelson and northern Westland, and sourthwards mainly east of
(though close to) the Main Divide from Lewis Pass, to Mt Peel in south Canterbury, South I. It
has an altitudinal range from 200 m to 1350 m. It is known also from south eastern Australia and
Tasmania (Kantvilas etal., 1985).

Habitat ecology: Pseudocyphellaria ardesiaca occurs on bark or amongst mosses on trunks of
the following phorophytes: Kunzea ericoides, Leptospermum scoparium, Nothofagus menziesii,
N. solandri var. cliff ortioides , and Phyllocladus alpinus. It prefers cool, moist, humid conditions
in montane to subalpine habitats and although having a blue-green photobiont it is not tolerant
of low-light intensities and is most frequently found at tree-line (Galloway & Simpson, 1978) or
lower margins of Nothofagus forest (it is most commonly collected from mountain beech, N.
solandri var. cliff 'ortioides) , or on the bark of Leptospermum in humid scrub associations on river
flats, in gullies, or on disturbed ground near recently fired beech forest.

It most frequently associates with other species of Pseudocyphellaria: P. bartlettii (in the
northern part of its range), P. crocata, P. crassa, P. degelii, (usually at tree-line in the mist zone
with Parmelia testacea and Usnea capillacea), P. fimbriata, P. pickeringii, P. glabra, P.
granulata, P. gretae, P. jamesii, P. margaretiae, P. nermula, and P. rubella. Elsewhere it
associates with the following lichens, most of which are indicators of cool, moist, humid
conditions of moderate to high light intensity: Hypogymnia turgidula, Hypotrachyna sinuosa,
Leio derma applanatum, L. sorediatum, Menegazzia circumsorediata, M. dielsii, M. pertransita,
Nephroma cellulosum, N. rufum, Normandina pulchella, Parmotrema chinense, Psoroma
durietzii, P. leprolomum, P. sphinctrinum, P. microphyllizans , Sphaerophorus ramulifer, S.
tener, and Stictafuliginosa.

Specimens examined: 30.

3. Pseudocyphellaria argyracea (Delise) Vainio Fig. 19

in Hedwigia 37: 35 (1898). - Sticta argyracea Delise in Mem. Soc. linn. Normandie 2: 91 pi. 7, fig. 30
(1825). For typification and additional synonymy see Galloway & James (1986).

Morphology: Thallus rosette-forming, orbicular, l-5(-10) cm diam. , corticolous or saxico-
lous, closely attached to bark or rocks, loosely attached and free when on twigs or amongst
moss. Lobes irregularly laciniate to broadly rounded, 3-10(-20) mm diam., 1-3 cm long,
discrete to imbricate. Margins sinuous, often slightly re volute and very slightly ridged below,
entire or minutely crenate-incised, usually delicately isidiate. Upper surface livid grey-blue or
grey-black, very minutely mottled (x 10 lens) when moist, pale olivaceous-grey to red-brown
when dry, plane, undulate or minutely and irregularly wrinkled, smooth or minutely roughened,
fragile, sorediate and isidiate, without maculae or phyllidia. Pseudocyphellae laminal, scat-
tered, punctiform, becoming sorediate, to 3 mm diam., ulcerose, round to irregular, flat to
convex, soredia coarsely granular, white. Pseudoisidia minute, style-formed, terete, simple
to coralloid, at first clustered at margins of pseudocyphellae (Fig. 6A), and at apices and margins
of lobes, in some specimens spreading more widely to form a diffract-coralloid crust, dark

PSEUDOCYPHELLARIA

65

Fig. 19 Pseudocyphellarla argyracea. Lake Rotoiti, Adams (WELT). A. Scale = 1 cm. B. Scale = 2 mm.

brown, mainly corticate, rarely sorediate below or occasionally forming expanded and flattened
regenerating lobules. Medulla white. Photobiont Nostoc. Lower surface pale fawnish or pinkish
white, shallowly wrinkled or pitted, densely tomentose from centre to margins, tomentum
whitish, silky, entangled, rather shaggy. Pseudocyphellae very rare or absent, minute, white,
punctiform, best seen near margins. Apothecia not seen.

Anatomy: Thallus 230-320 ^m thick. Upper cortex 40-50 um thick, uppermost 10-12 |im dark
brown, remainder straw yellow often with a clear yellow 10 |im zone immediately above
photobiont layer, cells thick-walled 4-5-13-5 urn diam. Photobiont layer 45-65 um thick,
photobiont Nostoc, cells to 4-5 ^im diam. Medulla 75-140 um thick, colourless, hyphae 4-5 um
diam. Lower cortex 34-45 um thick, pale straw yellow, cells thicker walled than those of upper
cortex, 4-5-14 um diam, outermost row of cells yellow-brown. Tomental hairs to 7\im diam.,
yellow-brown to red-brown, 100-450 um long, in clustered fascicles.

Chemistry: 7|3-acetoxyhopan-22-ol, hopane-7|3, 22-diol(tr.), hopane-15a, 22-diol, methyl
gyrophorate, and gyrophoric acid.

Distinguishing features: Pseudocyphellaria argyracea, a widespread, palaeotropical species
has a white medulla, a blue-green photobiont, and white pseudocyphellae on the upper and

66 D. J. GALLOWAY

lower surfaces. It is characterized by numerous, laminal, white pseudocyphellae which at
maturity become marginally pseudoisidiate with corticate, very delicate, finger-like
pseudoisidia, concolorous with thallus or darker (often dark brown to brown-black) developed
in clusters. In section the pseudoisidia are corticate in their upper parts, being covered by
brownish cortical cells continuous with the upper cortex. In some specimens, isidia persist as
integral cortical structures and may become coralloid, especially at the lobe margins where
they are not associated with laminal pseudocyphellae. Very often, isidia associated with laminal
pseudocyphellae eventually lose the cortex at the base and become sorediate. Such sorediate-
pseudoisidiate forms are then often very difficult to separate from specimens of P. intricata
where occasionally the laminal soredia may become pseudoisidiate and it is possible that a
continuum of forms exist between strictly sorediate P. intricata and strictly isidiate and laminally
pseudocyphellate F. argyracea. It has a characteristic two-hopane chemistry [Code A of Wilkins
& James (1979)], together with methyl gyrophorate and gyrophoric acid. It is not known fertile.

Variation: In the type locality and throughout the tropics where it is most often collected, P.
argyracea shows two main growth forms, with lobes being either broadly rounded and
imbricate, or elongate-laciniate and rather narrow, and sometimes with abraded, sorediate
margins (Galloway & James, 1986). New Zealand populations are rather similar to the type of
Sticta argyracea Delise from Reunion, but tend to be rather thinner and more fragile than
Mascarene specimens, and rarely in New Zealand does the species form luxuriant colonies such
as one finds in early tropical collections, especially those of Bory de St- Vincent made from the
type locality.

P. argyracea is related to P. boryana, another tropical species with which it is often sympatric,
but this latter species differs from it in having narrower, subdichotomously branching lobes
which are distinctly marginally phyllidiate or isidiate. The laminal pseudocyphellae of this
species are also not associated with isidia or pseudoisidia as they are in P. argyracea.

Pseudocyphellaria argyracea is distinguished from P. intricata and P. norvegica by the delicate
isidia or pseudoisidia which are developed both at the margins of the lobes and of the laminal
pseudocyphellae. Erosion of isidia associated with the margins of pseudocyphellae in P.
argyracea can occasionally give rise to pseudoisidiate-sorediate structures which may be very
difficult to distinguish from coarsely sorediate strains of P. intricata [see above, and also Coppins
& James (1979)].

The isidiate Macaronesian-western European species P. lacerata differs from P. argyracea in
several important respects. In P. lacerata there are no laminal pseudocyphellae and the isidia are
mainly marginal, though occasionally they regenerate from tears or cracks in the upper surface.
Isidia are rather nodular, and terete at first with a conspicuous apical white-frosted tomentum
(use x 10 lens), later isidia become glomerulate-coralloid-branched, and may even be somewhat
flattened and phyllidiate with a pale whitish lower surface and a dark brownish upper surface,
also usually with a characteristic whitish tomentum. Densely clustered imbricate marginal isidia
in P. lacerata appear grey-blue [see also Coppins & James (1979)].

The Australasian species P. haywardiorum differs from P. argyracea in its characteristic
punctate-impressed upper surface and the bullate lower surface with its prominent, large,
raised pseudocyphellae.

Distribution: (Fig. 20) Pseudocyphellaria argyracea is still poorly collected in New Zealand,
though it appears to be genuinely rare there and apparently does not have the same pattern of
distribution as the other widespread, primarily tropical species, P. aurata. So far it is known
from lat. 39S southwards to Stewart I. , in both lowland and subalpine habitats from sea-level to
600m.

Habitat ecology: Pseudocyphellaria argyracea in New Zealand is a species of medium - to
low-light regimes being found in cool, moist, semi-shaded to deeply shaded habitats on stones or
rock, among bryophytes and ferns ( Phymatodes diversifolium) on the ground, and at or near the
bases of forest trees. It is known from the following phorophytes: Dracophyllum, Kunzea
ericoides, Melicytus ramiflorus, and Nothofagus solandri var. cliff ortioides . On rock in subalpine

PSEUDOCYPHELLARIA

67

-40

172

Norfolk I

Kermadec 30-
Is

Lord Howe I

Three Kings Is

The Snares

Auckland Is
\

Campbell I
\

Macquarie I

160

170

-42

-44

174 C

176

36-

38-

Fig. 20 Distribution of Pseudocyphellaria argyracea.

68

D. J. GALLOWAY

habitats it associates with Parmelia signifera, and elsewhere in its range in New Zealand it grows
with the lichens Normandina pulchella, Pseudocyphellaria dissimilis, P. glabra, P. intricata,
Sticta fuliginosa, and 5. limbata. Its detailed ecological requirements in New Zealand are still
poorly understood however. It is a rather thin, fragile species, often growing amongst mosses,
with thalli being rather flattened, and when growing on bark, closely appressed.

Specimens examined:

14.

4. Pseudocyphellaria aurata (Ach.) Vainio

Fig. 21

in Acta Soc. Fauna Flora fenn. 7: 183 (1890). - Sticta aurata Ach., Methodus: 277 (1803). - Lichen
auratus (Ach.) Sm. & Sowerby, Engl. Bot. 33: tab. 2359 (1812). - Nephroma aurata (Ach.) Pers. in
Gaudichaud, Voy. Uranie Bot. : 202 (1827). - Crocodia aurata (Ach.) Link, Handbuch 3: 177 (1833). -
Parmelia aurata (Ach.) Eschw. in Martius, Iconespl. crypt. 2: 34 (1834). - Parmosticta aurata (Ach.)
Nyl. in Flora, Jena 58: 303 (1875). - Lobaria aurata (Ach.) Kuntze, Revis. gen. pi. 2: 876 (1891). Type:
? England, Devon, sine loco, ex Herb. Hudson - label incomplete (H-ACH 1534! - holotype).

Morphology: Thallus in neat rosettes to irregularly spreading, 5-10(-15) cm diam., loosely
attached, subascendent, undulate at margins, more tightly adnate centrally, corticolous,
occasionally terricolous or saxicolous. Lobes 3-12(-20) mm wide, 25-40 mm long, irregularly
laciniate, discrete at margins, becoming imbricate-confluent centrally to discrete from
margins to centre, occasionally with small, epiphytic thalli of Normandina pulchella and species
of Pannaria. Margins sinuous, undulate, subascendent, entire to slightly notched or
scalloped or crenate, slightly thickened below, with punctiform to linear, yellow pseudocyphel-

Fig. 21 Pseudocyphellaria aurata. Great Mercury Island, Hayward H 40.87 (AK 154839). A. Scale = 1
cm. B. Scale = 2 mm.

PSEUDOCYPHELLARIA 69

lae, or more often becoming sorediate, occasionally minutely lobulate with small, regenerating
thalline leaflets. Upper surface bright lettuce green to a dark, fresh green when wet, pale
glaucous-grey, olive-brown or buff when dry, often noticeably reddening on storage and
becoming brick red to red-brown in the herbarium; undulate, occasionally shallowly to
obscurely ridged or pitted in parts, though never truly faveolate, internal cephalodia visible as
hemispherical swellings; matt, minutely scabrid-areolate in parts, especially near margins which
may also be occasionally very finely short-tomentose, to distinctly or delicately pruinose, pruina
patchy, thin, white. Soredia yellow, coarse, granular often blackened and appearing
pseudoisidiate, in primarily marginal, sinuous, labriform soralia which erode conspicuously on
the lower surface, also widely scattered at times laminally, in rounded, erose soralia (in
favourable conditions a developing sequence from granular soralium to small, corticate lobules
differentiating both upper and lower surfaces may be observed). Isidia, maculae, phyllidia, and
pseudocyphellae absent. Medulla yellow. Photobiont green, Dictyochloropsis . Lower surface
yellow when wet, rufous, brownish red when dry, dark brown centrally; tomentum rather thin
and variable, silky, pinkish buff or whitish at margins, thicker and darker to chocolate brown
centrally to rather poorly developed and often lacking centrally; shallowly wrinkled-puckered at
margins, smooth elsewhere. Pseudocyphellae yellow, numerous, conspicuous, round to
irregular, minute, punctiform especially at margins, 0-5-l-5(-2) mm diam., often confluent
centrally, margins very slightly raised, decorticate area flat to shallowly concave, rarely
somewhat convex, occasionally with central corticate areas. Apothecia rather rare, submarginal
to occasionally laminal, solitary, l-3(-5) mm diam., appearing at first as rounded swellings on
upper surface, at length fissuring at the apex to expose concave disc, margins, ragged,
yellow-sorediate, distinctly pedicellate, pedicel 1-5-2-5 mm diam. , 0-5-l-5(-2) mm tall, slightly
narrower than disc and giving fruits a 'waisted' appearance, exciple concolorous with thallus,
minutely areolate-scabrid, not hirsute; disc matt, red-brown, imperf orate, epruinose, often
obscured by overarching sorediate margins; insertion of pedicel distinctly excavate-invaginate
on lower surface. Pycnidia scattered, occasional, minute, punctiform to raised papillate, to
0.1 mm diam., red-brown when wet, black when dry.

Anatomy: Thallus (200-)250-350(-450) um thick. Upper cortex colourless, irregularly lumpy,
uneven, (40-)45-70(-80) um thick, cells 5-9 um diam. Medulla loosely interwoven, 85-120 um
thick, brownish yellow, hyphae 4-5 urn thick, densely encrusted with yellow-brown crystals.
Photobiont layer (45-)55-75(-90) um thick, cells of Dictyochloropsis round to slightly irregu-
lar, densely packed, 4-5-7 um diam. , chloroplast occupying most of cell. Cephalodia developed
between photobiont layer and lower cortex, broady ovoid to hemispherical, 100-140 x
90-110 um, comprising an outer sheath (10-14 um thick) of periclinal hyphae enclosing a core
80-110 um thick of thick-walled cells enclosing packets ofNostoc. Lower cortex 25-37(-45) um
thick, pale straw to hyaline, 3-6 rows of cells 6-5-11 um diam. Tomental hairs thick-walled, to
7 um thick, central canal conspicuous, simple, solitary or in fascicles of 3-6 hyphae together,
45-150 um long. Apothecia: Exciple containing photobiont cells and comprising cortex,
photobiont layer, and medulla as in vegetative thallus. Hypothecium 45-75 um thick, opaque,
yel|ow-brown to red-brown, unchanged in K. Thecium colourless to pale straw, 55-75 um tall;
epityiecium red-brown, minutely granular, 7-11-5 um thick, colour external to tips of paraph-
yses, unchanged in K. Asci very rare, broadly clavate, tapering narrowly at foot, 50-65 um tall,
13-5-U8 um at widest part. Ascospores broadly fusiform-ellipsoid, pointed at apices, 3-septate at
maturity, central two locules larger, end locules small and often irregular, distinctly vacuolate,
brown, (25-)30-32 x 6-7 um.

Chemistry: Pulvinic acid, pulvinic dilactone, calycin, fernene triterpenoids (A. L. Wilkins,
pers. comm.) and unidentified substances.

Distinguishing features: Pseudocyphellaria aurata is a cosmopolitan species characterized by a
yellow medulla, a green photobiont, yellow pseudocyphellae on the lower surface, and
prominent, marginal, labriform, linear, yellow soralia eroding back the lower surface and
containing coarse, granular soredia.

70 D. J. GALLOWAY

Variation: Pseudocyphellaria aurata, an epiphyte of northern coastal forest trees and shrubs, is
a fairly constant species showing rather little morphological variation other than differences in
lobe dimensions often encountered in rosette-forming taxa. There is some variation in the
texture of the upper surface especially of the lobe margins where, pruina, scabrosity or laminal
cracking may be most evident in some collections, though these features are not always constant
in local populations and appear to be influenced by particular microclimatic factors. The
laminal, erose soralia are also highly variable in their occurrence and may be common to
absent altogether. The marginal, sinuous, linear, labriform, yellow soralia are probably the
most characteristic and distinguishing feature of this species and are never pectinate-isidiate as
they are in the related species P. poculifera, which is also much more commonly fertile than P.
aurata. Occasionally small lobules developing from soralia may be seen in the marginal soralia,
less commonly in the laminal soralia, but these are easily distinguished from the coralloid isidia
at the margins of P. poculifera. The lobes of P. aurata are generally broader, thicker, and less
incised than those of P. poculifera, and there are also differences between the two species in
apothecial anatomy (see below under P. poculifera}.

P. pickeringii is distinguished from P. aurata by its copious laminal and marginal isidia, its lack
of soredia, the massive, corrugate-scabrid apothecial exciple, colourless spores, and a chemistry
dominated by stictane triterpenoids. The montane-subalpine P. rubella has certain similarities
with both P. aurata and P. poculifera but, although sorediate, it has a distinctive non-glabrous
upper surface with development of a characteristic laminal tomentum. Its chemistry is distinct,
containing triterpenoids of the lupane series. Specimens of P. aurata from some 19th century
tropical collections were sometimes misidentified as Sticta crocata, this error no doubt deriving
from Hoffmann's erroneous use of the name Platisma crocatum (L.) Hoffm. [Descr. pi. cl. crypt.
2: 52, tab. XXXVIII figs 1-3 (1794)] for material of P. aurata collected by Olof Swartz in the
West Indies.

Distribution: (Fig. 22) In mainly coastal habitats from Three Kings Is to Wellington in North
I., and a solitary collection from Nelson in South I., sea-level to 500 m. Known from the
following offshore islands north of lat. 38S; Three Kings Is (Great I. and South West I.), Cavalli
Is, Poor Knights Is, Hen and Chickens, Rakitu I., Cuvier I., Rangitoto I., Mokohinau I., Little
Barrier I., Great Barrier I., Great Mercury I., Red Mercury I., Shoe I., Slipper I., and the
Aldermen Is, Tiri Tiri I. , Whale I. , Penguin I. , and Rabbit I.

Habitat ecology: Pseudocyphellaria aurata is widespread in both cool-temperate and tropical
regions of the world and in New Zealand is characteristic of lowland coastal forest north of lat.
38S. It will colonize both trunks and twigs of the following phorophytes: Avicennia marina var.
resinifera, Coprosoma macrocarpa, C. rhamnoides, Cordyline australis, C. kaspar, Corynocar-
pus laevigatus, Gaultheria antipoda, Griselinia littoralis, Kunzea ericoides, Leptospermum
scoparium, Litsaea calicaris, Melicytus ramiflorus, Metrosideros excelsa, Myrtus bullata, and
Sphaeropteris medullaris. It has also been collected from boulders and obsidian outcrops in
grassland, from soil below Leptospermum in coastal scrub, and from maritime rocks. It is a
photophilous species in areas having high summer temperatures and medium to low rainfall
where a proportion of the precipitation is as mist or fog. Although primarily a northern species
and best represented in North Auckland habitats, P. aurata has been able to extend its
range beyond the biogeographical barrier of the Auckland isthmus (during higher sea levels
of interglacials the isthmus became a strait, isolating North Auckland as an island), through
production of soredia.

Pseudocyphellaria aurata associates in northern habitats with the following lichens: Cocco-
carpia erythroxyli, C. pellita, Leioderma duplicatum, L. pycnophorum, L. sorediatum,
Normandina pulchella, Pannaria elatior, P. fulvescens, P. periptera, Parmotrema crinitum, P.
cristiferum, P. reticulatum, P. tinctorum, Pseudocyphellaria carpoloma, P. chloroleuca,
Ramalina australiensis , Sticta squamata, and Teloschistes flavicans .

Specimens examined: 65 .

PSEUDOCYPHELLARIA

71

Fig. 22 Distribution of Pseudocyphellaria aurata.

72 D. J. GALLOWAY

5. Pseudocyphellaria bartlettii D. Galloway Fig. 23

in Lichenologist 17: 303 (1985). - Stictina mougeotiana f. isidiosa Mull. Arg. in Bull. Herb. Boissier4:
89 (1896). - Sticta mougeotiana f. isidiosa (Mull. Arg.) Zahlbr., Car. L/c/i. t/mV. 3: 393 (1925). Type:
Australia, Queensland, Mr Bailey 739 (G 002234! - lectotype).

Morphology: Thallus orbicular, 3-6(-12) cm diam. , loosely attached centrally, margins free,
corticolous. Lobes broadly rounded, discrete to subimbricate, 1-3 cm diam. Margins entire to
minutely crenate, notched or incised, often copiously sorediate and eroding back onto lower
surface, sometimes inrolled. Upper surface dark leaden or slate blue when wet, with white
maculae in well-defined reticulate patterns giving a marbled appearance to thallus (x 10 lens),
pale olivaceous-grey, grey-buff or brownish red when dry (maculae much less obvious in dry
thalli), weakly to strongly reticulate-faveolate or wrinkled-plicate towards margins, undulate
to shallowly pitted or wrinkled centrally, matt, rather fragile and papery when dry, sorediate,
pseudocyphellate, without isidia or phyllidia. Soredia in scattered, laminal erose soralia, 0-2-2
mm diam. , becoming confluent and linear at margins and along reticulate ridges, brownish or
greyish, gnarled-glomerulate, minutely style-formed, pseudoisidiate, coarsely granular,
becoming eroded-white at maturity, often densely clustered near margins and obscuring lamina.
Pseudocyphellae occasional, scattered, inconstant, minute, pinprick-like, yellow, on ridges near
margins of lobes. Medulla white. PhotobiontNostoc. Lower surface noticeably bullate, minutely
wrinkled or pitted below tomentum, pale yellowish buff and glabrous in a narrow zone at
margins, or uniformly buffer dark red-brown to black, tomentum tufted, rather short, thin or
thickly developed, occasionally pale buff, more often brown or black. Pseudocyphellae scat-
tered, sparse to moderately numerous, eroded-papillate appearing verruciform, 0-1 mm diam.
or less, decorticate area flat, white or very pale cream, level with or slightly embedded in
tomentum. Lichenicolous fungus rarely present, ascomata discoid, translucent, red-brown,
developed among soredia 0-5 mm diam. or less. Apothecia not seen.

Anatomy: Thallus 130-200(-275 at soredia), um thick. Upper cortex 18-24 um thick, upper
9-11 um dilute orange-brown or red-brown, cells of upper surface compressed forming a
necrotic layer, lower 9-13 um colourless, cells larger, pseudoparenchymatous, of round to
irregular isodiametric, thick-walled cells 4-6 um diam. Photobiont layer (34-)45(-65) um
thick, photobiont Nostoc in irregular clumps 11-22 um diam. , with interstitial hyphae ramifying
in all directions. Medulla 45-90(-135) um thick, erupting through upper cortex as erumpent
soralia and carrying aloft scattered clumps of photobiont, hyphae dilute yellow-brown, peri-
clinally arranged, loosely interwoven, encrusted with small crystals. Lower cortex (15-)20
(-34) um thick, dilute yellow-brown, 2-3 rows of cells 4-5-9 um diam. , a narrow zone abutting
medulla (4-9 um) of dark brown, stretched cells appearing periclinally arranged, visible in
section as a continuous narrow, brown-black line. Tomental hairs pale to dark brown, simple,
septate, 4-5-6-5 um diam., 60-180(-300) um long, in fascicles, 3-10(-20). Ascospores of
parasitic fungus colourless, simple, oval-ellipsoid, uniseriate in ascus, 6-8-9-1 x 2-3 um.

Chemistry: Tenuiorin, methyl gyrophorate, hopane-6a, 7(3, 22-triol, stictic, constictic,
cryptostictic, and norstictic (tr) acids.

Distinguishing features: Pseudocyphellaria bartlettii has a white medulla, a blue-green photo-
biont, white pseudocyphellae on the lower surface, and marginal and laminal erose soralia
containing brownish or greyish, pseudoisidiate to coarsely granular soredia. The lobes are
broadly rounded and on the upper surface photobiont-free areas are present as well-defined
reticulate maculae (x 10 lens). It has a complex chemistry related to that of P. crocata, but
lacking the yellow pigments calycin, pulvinic dilactone, and pulvinic acid from the maculae,
soralia, and pseudocyphellae. Pseudocyphellaria bartlettii is named for the late J. K. Bartlett
(Galloway, 19876), a distinguished collector of New Zealand lichens.

Variation: In the eight specimens seen, little morphological variation was observed, all
individuals from South Auckland to South Canterbury appearing remarkably uniform in gross
morphology.

PSEUDOCYPHELLARIA

73

Fig. 23 Pseudocyphellaria bartlettii. Rangipo, Bartlett 18865 (BM). Scale = 1 cm.

Pseudocyphellaria bartlettii differs from P. crocata in lacking yellow pigments in the upper
cortex, soralia, and pseudocyphellae, and also in its broad-lobed rosette-forming thallus with a
wrinkled-plicate and rather papery upper surface. It differs from P. neglecta for the same
reasons as well as lacking marginal and/or laminal phyllidia; and it is distinguished from P.
haywardiomm chemically and morphologically (P. haywardiorum has laciniate lobes, with a
strongly punctate-impressed upper surface, a bullate lower surface with conspicuous, raised
pseudocyphellae well delimited from an evenly red-brown to brown-black tomentum; it also has
a two-hopane chemistry).

Distribution: (Fig. 24) North I. , South of lat. 38S from near Te Awamutu (South Auckland)
to Wellington (Rangipo) and Hawkes Bay (Kuripapango and Kaweka Range). South I. , Nelson
(Whangapeka Track) to South Canterbury (Mt Peel), 500-1100 m. Known also in eastern
Australia (Queensland).

Habitat ecology: Pseudocyphellaria bartlettii is still an undercollected species in New Zealand.
It forms neat rosettes on the bark of Leptospermum in cool, humid, shaded sites and associates
with Heterodermia speciosa, Normandina pulchella, Pseudocyphellaria ardesiaca, P. gretae,
Parmotrema chinense, P. reticulatum, and Psoromasphinctrinum.

Specimens examined: North Island. South Auckland: Ngaroma, S. of Te Awamutu, 14 July
1979,7. K. Bartlett (EM); Waipapa, Tea Tree Bush N. of Mangakino, 12 May 1981,7. K. Bartlett
(BM). Wellington: Snake Valley S. of Turangi, 11 May 1981, 7. K. Bartlett (BM); Rangipo, 12
May 1981, 7. K. Bartlett (BM). Hawkes Bay: Kaweka Range, headwaters of Ngaruroro River,
11 July 1982, 7. K. Bartlett (BM); Kuripapango, 13 July 1981,7. K. Bartlett (BM). South Island.
Nelson: Whangapeka Track, Tasman Mountains, 26 January 1980, 7. K. Bartlett (BM).
Canterbury: Deer Spur Track, Mt Peel, 21 March 1979, D. J. Galloway (CHR 267113).

74

D. J. GALLOWAY

Lord Howe I

Norfolk I

Three Kings Is

Kermadec 30-
Is

The Snares

Auckland Is
\

Campbell I
\

Macquarie I

Bounty Is
Antipodes Is

160

170 C

40

46

166

i

168

50'

180

170

172 C

172

174

176

174

176

178

34-

36-

40-

42

Fig. 24 Distribution of Pseudocyphellarla bartlettii.

PSEUDOCYPHELLARIA 75

6. Pseudocyphellaria billardierei (Delise) Rasanen Figs 25, 26

in Annls Bot. Soc. zool. -hot. fenn. Vanamo 2 (1): 39 (1932). - Sticta billardierei ['Billardierf] Delise in

Mem. Soc. linn. Normandie 2: 99, pi. 8 fig. 35 (1825). - Lobaria billardierei (Delise) Hellbom in Bih.

K. svenska Vetensk. - Akad. Handl. 21 (3/13): 41 (1896). - Sticta faveolata var. billardierei (Delise)

Church. Bab. in J. D. Hook., Fl. nov.-zel. 2: 278 (1855). - S. cellulifera var. billardierei (Delise) Hue

\nNouv. ArchsMus. Hist. nat. Paris III, 2: 306 (1890). Type: Cap de Van-Diemen (Tasmania), J.J-H.

de la Billardiere (PC-LENORMAND! - lectotype; FI!-isolectotype) [see Galloway et al. (19836);

Galloway & James (1986)].
Sticta fossulata Dufour in Delise in Mem. Soc. linn. Normandie 2: 99 (1825), nom. nud. (Art. 32-1). -

Crocodia fossulata Trevisan, Lichenotheca veneta exs. 75 (1869) nom, nud. (Art. 32.1). - Lobaria

fossulata Kuntze, Revis. gen. pi. 2: 876 (1891) nom. nud. (Art. 32.1).
Sticta flotowiana Laurer in Linnaea 2: 40 (1827). - Pseudocyphellaria flotowiana (Laurer) Malme in Bih.

K. Svenska Vetensk. -Akad. Handl. 25 (3/6) :22 (1899). Type: Nova Hollandia (Australia), Sieber

(BM! - lectotype; M! - isotype).
Sticta linearis J. D. Hook. & Taylor in Hook. Lond. J. Bot. 3: 648 (1844). - Pseudocyphellaria linearis (J.

D. Hook. & Taylor) Dodge in Nova Hedwigia 19: 489 (1971). - Sticta fossulata var. linearis (J. D.

Hook. & Taylor) Lindsay in Trans. Linn. Soc. Lond. 25: 499 (1866). - Lobaria fossulata var. linearis

(J. D. Hook. & Taylor) Hellbom in Bih. K. svenska Vetnsk. -Akad. Handl. 21 (3/13): 41 (1896).

Type: Lord Auckland's Group (New Zealand), /. D. Hooker (BM! - lectotype).
Sticta fossulata f. lacinulata Krempelh., Reise Novara Bot. 1: 120 (1870). - S. billardierei var. lacinulata

(Krempelh.) Mull. Arg. in Bull. Herb. Boissier 2, App. 1: 36 (1894). - Lobaria billardierei var.

lacinulata (Krempelh.) Hellbom in Bih. K. svenska Vetensk. -Akad. Handl. 21 (3/13): 41 (1896).

Type: New Zealand, sine loco, Jelinek (M - not seen).

Morphology: Thallus irregularly spreading, somewhat entangled, 5-15(-30) cm diam.,
corticolous, loosely attached centrally, margins free, often ascending. Lobes regularly dichot-
omously to subdichotomously branched, discrete at margins, imbricate centrally, (l-)3-
15(-30) mm wide, (l-)3-6(-10) cm long, rather flat, main lobes canaliculate, apices truncate,
acute or bifurcate. Margins entire, slightly to markedly thickened above and below, forming a
noticeable ridge on the lower surface, smoothly rounded, without (or very rarely with)
projecting pseudocyphellae. Upper surface bright lettuce green to glaucous-green when moist,
pale grey-green to olive brownish when dry, shallowly to deeply faveolate, main interconnecting
ridges well-defined, arcuate, 0-l-0-5(-l-5) mm wide, flat, extending across width of lobe, and

It

,*-* /f/f -/if-

Fig. 25 Sticta billardierei. Lectotype (PC-LENORMAND). Scale = 2 cm.

76

D. J. GALLOWAY

Fig. 26 Pseudocyphellaria billardierei. North Egmont, Tibell 14967 (UPS). Scale = 1 cm.

dividing lobes into rather large, shallow or deep faveolae, within these secondary, intricating
ridges form smaller secondary faveolae, without soredia, isidia, phyllidia, maculae or pseudo-
cyphellae. Medulla white. Photobiont green. Lower surface white or pale buff at margins, darker
centrally, conspicuously wrinkled to bullate, prominently ridged at margins, glabrous in a
narrow to wide marginal zone, tomentose centrally or from margins to centre, tomentum thin
and scattered to dense, wholly, pale whitish or buff and velvety at margins, thicker and
entangled, black or brown centrally. Pseudocyphellae white, scattered to frequent, minute,
flecklike at margins, 0-05 mm diam. or less, elsewhere to 0-2 mm diam., on interconnecting
ridges, raised, verruciform, decorticate area small, depressed-punctate or intented below
surrounding margins, rarely flat, margins thin, puckered. Pycnidia common, in rows or
clustered at margins and on laminal ridges, minute, punctiform, 0-1 mm or less, red-brown to
black, often eroding and leaving small pits. Apothecia sparse to moderately common, rather
scattered, often more common towards lobe apices, marginal or submarginal, rarely laminal on
ridges, solitary to clustered, sessile, constricted at base to subpedicellate, 0-5-5 mm diam.,
rounded or contorted through mutual pressure, disc dark red-brown to black, shining at first
becoming matt, slightly papillate or sooty, epruinose, exciple pale yellow-buff to red-brown,
translucent and colourless when moist, corrugate-scabrid to occasionally smooth and
shining, rarely pubescent or tomentose towards base, obscuring disc in young fruits, restricted
to a thin, crenulate or scabrid margin at maturity.

PSEUDOCYPHELLARIA 77

Anatomy: Thallus 150-300(-450-500) um thick. Upper cortex 35-45 um thick, upper parts (9
to 20 urn) yellow-brown, remainder colourless, of densely compacted thick-walled cells, 5-7 um
diam. Photobiont layer 22-35 um thick, photobiont green, cells round to irregular 4-5-7 um
diam, closely compacted. Medulla 100-300 um thick, colourless, hyphae to 4-5 um diam.,
encrusted with small crystals. Lower cortex 20-30(-34) um thick, uniformly dilute yellow-
brown, colour most intense near medulla, cells similar to those of upper cortex, tomental hairs
pale yellow-brown, 4-5 um thick, 20-60(-120) um long, in fascicles. Apothecia: Exciple
90-120(-180) urn thick, cellular, outer 20-30 um dilute yellow-brown otherwise colourless, cells
7-11 um diam, thick-walled. Hypothecium 54-82 um thick, dilute orange-brown or yellow-
brown, opaque. Thecium 90-140 um tall, colourless; epithecium dilute red-brown or yellow-
brown, dark greenish black in K, 7-13 um thick, granular in parts, colour external to apices of
paraphyses;para/?/ryses simple, apices (2-4 cells) swollen, moniliform. Asci 70-75 x 15-18
[Am. Ascospores ellipsoid, thickened 1-septate, brown or grey-brown 25-32(-38) x 9-11.5 um,
septum 4-5-5-5 um thick.

Chemistry: Tenuiorin, methyl evernate, methyl lecanorate, methyl gyrophorate, evernic and
gyrophoric acids, 2"-O-methyl tenuiorin, 2', 2"-di-O-methyltenuiorin, hopane-6a, 22-diol
(zeorin), 6a, 16(3-diacetoxyhopan-22-ol, 6a-acetoxyhopan-16(3, 22-diol, 6a-acetoxyhopan-23-
oic acid, 6a, 22-dihydroxyhopan-23-oic acid, norstictic, stictic, cryptostictic (tr.) and 9-a-
acetylconstictic acid (tr.) [Code C of Wilkins & James (1979), see also Galloway et al. (1983ft),
Galloway & James (1986), Elix (1986)].

Distinguishing features: Pseudocyphellaria billardierei is an Australasian species having linear-
elongate, dichotomously branching lobes, often widely divergent at their apices and which are
conspicuously reticulate-faveolate. Faveolae may be shallow or deep, and the main intercon-
necting ridges separating them are strongly defined, arcuate, flat, and extending across the
width of the lobe. Lobe margins are entire, smoothly rounded, without projecting pseudocy-
phellae and thickened both above and especially below, forming a distinct marginal ridge. The
upper surface lacks soredia, isidia, phyllidia, maculae or pseudocyphellae. The species has a
white medulla and a green photobiont. The lower surface is wrinkled to bullate, pale to brown
or blackened, with moderate to dense tomentum and a narrow to broad, glabrous, marginal
zone. Pseudocyphellae are white, elevated, verruciform, sometimes with puckered margins,
and an impressed-punctate decorticate area. Apothecia are primarily marginal, the disc dark
red-brown to black and never pruinose. The epithecium turns dark greenish black in K, a
characteristic reaction of the species, and ascospores are thickened 1-septate, grey-brown to
brown.

Variation: Pseudocyphellaria billardierei shows considerable variation in lobe width and length
and in complexity of branching, depending on local ecological conditions, though, in general,
narrow-lobed specimens are more commonly encountered than broad-lobed ones. Specimens
with broad lobes tend to be more commonly collected in North Island (especially from Mt
Egmont National Park) habitats. The great diversity of lobe form found in P. faveolata is not
encountered in P. billardierei. Very occasionally, depauperate or damaged specimens may
develop gnarled-glomerulate to obovoid phyllidia at the margins, but this seems a rare and
inconstant condition and is probably a response to particular local adverse conditions. The
colour of the upper surface varies with degree of exposure to wind, light or salt-spray, with both
coastal scrub and subalpine scrub forms having a noticeably darkened to blackened upper
(and often also lower) surface. The tomentum of the lower surface also varies from thin,
scattered, and poorly developed to thick and entangled, being best developed centrally, with the
margins of most specimens examined glabrous.

This species is most often confused with P. faveolata, both in the field and in the herbarium,
however, there are a number of important characters which distinguish the two species. P.
billardierei has entire, smoothly rounded lobe margins, noticeably ridged below, and without
the projecting pseudocyphellae which are characteristic of P. faveolata. The apothecia are never

78 D. J. GALLOWAY

pruinose in P. billardierei as they are in P. faveolata (at least in the youngest fruits) and are usually
also less commonly developed. Epithecial pigments in P. billardierei turn greenish black in K
whereas in P. faveolata epithecial pigments turn rose-purple in K. The chemistries of the two
taxa are also different, P. billardierei being Code C of Wilkins & James (1979) and lacking
physciosporin, a metabolite characteristic of P. faveolata [Code B of Wilkins & James (1979)].
Pseudocyphellaria billardierei is distinguished from P. physciospora by differences in epithecium
reaction in K, chemistry, and colour of the pseudocyphellae.

Pseudocyphellaria billardierei is distinguished from P. carpoloma which has yellow pseudo-
cyphellae (which project at the lobe margins) and a differing chemistry [Code D of Wilkins &
James (1979)], and also from P. rufovirescens which has a pale, glabrous lower surface and a
simple, 2-hopane chemistry [Code A of Wilkins & James (1979)]. Epithecial pigments in both P.
carpoloma and in P. rufovirescens remain unchanged on addition of K to optical sections. See
also Table 2 (p. 216).

Ascospores in P. billardierei remain thickened 1-septate at maturity, whereas in both P.
carpoloma and P. faveolata both thickened 1-septate and 3-septate spores are routinely found,
while in P. rufovirescens the spores are never thickened 1-septate, and are pale yellow-brown
and not the smoky grey-brown or dark brown of the three other species mentioned (see also
Galloway etal, 19836; Galloway, 19856).

Distribution: (Fig. 27) Auckland, Rangitoto I., South Auckland south of lat. 37S to South-
land and also Stewart I., and the Auckland Is (lat. 53S). Most common in rain-forest areas,
especially in South I. , both east and west of the Main Divide. Sea-level to 1080 m.

Habitat ecology: Pseudocyphellaria billardierei is a large, conspicuous, reticulate-faveolate,
dichotomously branching lichen, most commonly collected in rain-forest areas of both North
and South Islands and on Stewart Island. It is found in beech (Nothofagus) forest, conifer
(Dacrydium, Dacry carpus, Libocedrus, Phyllocladus, Podocarpus, and Prumnopitys) forest,
and conifer-mixed broadleaf hardwood forest, also in successional stands of Leptospermum in
fire-altered habitats, and in subalpine and coastal scrub or shrublands (Dracophyllum, Hebe,
Olearia, Senecio). It is also known from lava blocks on Rangitoto I. Although it reaches an
altitudinal limit of 1080 m on Phyllocladus alpinus on the Kaimanawa Range (Lat. 40S) near the
most northern part of its distribution in New Zealand, in general it is most richly developed in
low- to mid-altitude, open forest between 200 and 800 m where it often forms distinctive,,
large, shelf-like clones with the divergent, dichotomously branching lobes projecting from tree
boles. It is an epiphyte of the following phorophytes: Beilschmiedia tawa, Carpodetus serratus,
Coprosma linearifolia, C. lucida, C. propinqua, Dacrydium biforme, D. cupressinum, Dacry-
carpus dacrydioides, Dracophyllum longifolium, D. uniflorum, Griselinia littoralis, Hebe salici-
folia, Hoheria glabrata, Fuchsia excorticata, Kunzea ericoides, Leptospermum scoparium,
Libocedrus bidwillii, Melicytus ramiflorus, Metrosideros robusta, M.umbellata, My r sine divari-
cata, Myrtus bullata, Nothofagus fusca, N. menziesii, N. solandrivar. cliff ortioides , N. truncata,
Olearia, Phyllocladus alpinus, Pittosporum tenuifolium, Podocarpus hallii, P. totara, Prumnop-
itys ferruginea, Pseudopanax crassifolius , Senecio eleagnifolius , and Weinmannia racemosa.

In many habitats it is one of the major lichen epiphytes, frequently co-dominant with
Pseudocyphellaria multifida and Sticta subcaperata, especially in mid-altitude forests west of the
Main Divide in South Island. It is also a common component of the rich lichen assemblages
found on tree bark in moderate to high light habitats, associating with the following lichens:
Coccotrema cucurbitula, Lobaria adscripta, Menegazzia pertransita, Metus conglomeratus ,
Nephroma australe, Pannaria immixta, Parmeliella nigrocincta, Phylctella sordida, P. sub-
uncinata, Pseudocyphellaria colensoi, P. dissimilis, P. faveolata, P. glabra, P. homoeophylla,
P. intricata, P. lividofusca, P. multifida, P. rubella, P. rufovirescens, Psoroma contextum,
P. microphyllizans, P. pallidum, P. pholidotoides , P. sphinctrinum, Sphaerophorus melano-
carpus, S. notatus, S. patagonicus, S. scrobiculatus , S. tener, Sticta filix, S. lacera, S. latifrons, S.
subcaperata, and Usnea capillacea.

Specimens examined: 130.

PSEUDOCYPHELLARIA

79

-40

172

174

176

178

Norfolk I

Lord Howe I

Three Kings Is

The Snares

Auckland Is

Campbell I
Macquarie I

160

-42

-44

46

166

i

168

170

172

174 C

1

176 C
I

34_

40-

42-

44 u

Fig. 27 Distribution of Pseudocyphellaria billardierei.

80 D. J. GALLOWAY

7. Pseudocyphellaria carpoloma (Delise) Vainio

Figs 28, 29

in Hedwigia 37: 34 (1898). - Sticta carpoloma Delise in Mem. Soc. linn. Normandie 2: 159, pi. 19, right
hand figure (1825). - Stictina carpoloma (Delise) Nyl. , Syn. meth. lich. 1 (2) : 339 (1860). - Saccardoa
carpoloma (Delise) Trevisan, Lichenotheca veneta exs. 75 (1869). - Cyanisticta carpoloma (Delise)
Gyelnik in Feddes Reprium Spec. Nov. reg. veg. 29: 2 (1931). Type: New Zealand, 'Sur les vieux arbres
a la Nouvelle Zelande', Bay of Islands, 1824, ?fl. P. Lesson (PC-LENORMAND! - holotype) [see
note 1].

Sticta impressa J. D. Hook. & Taylor in Hook. Lond. J. Bot. 3: 648 (1844). - Pseudocyphellaria impressa
(J. D. Hook. & Taylor) Vainio in Hedwigia 38: 187 (1899). Type: New Zealand, sine loco (prob. Bay
of Islands), /. D. Hooker (FH! - lectotype [fide Muller Argoviensis, (1888:13)]: BM! isotypes) [see
note 2] .

Sticta glaucolurida Nyl. in Flora, Jena 50: 438 (1867). - Lobaria glaucolurida (Nyl.) Hellbom in Bih. K.

i/it-'tfo +njCf6u,t<

Fig. 28 Pseudocyphellaria carpoloma. New Zealand, Hooker (BM). Note Joseph Hooker's pencil
sketches of anatomical details. Scale = 2 cm.

PSEUDOCYPHELLARIA

81

Fig. 29 W. H. Fitch's plate (of Pseudocyphellaria carpoloma) in Flora Novae Zelandiae based on
specimen in Fig. 28. (BM).

svenska Vetensk. - Akad. Handl. 21 (3/13): 37 (1896). Type: Nova Zelandia, sine loco (prob.

Wellington), 1867, C. Knight 10 (H-NYL 33587! - holotype). [see note 3].
Sticta borneti Mull. Arg. in Flora, Jena 65: 204 (1882). Type: New Zealand, D'Urville (G! - holotype)

[see note 4]. Fig. 30

Sticta expansa Stirton in Trans. N. Z. Inst. 32: 72 (1900). Type: New Zealand, near Wellington,

J. Buchanan (BM! -lectotype).

Note 1 : Sticta carpoloma Delise

In 1980 the name Sticta carpoloma was lectotypified on material from Montagne's herbarium now in BM
(Galloway & James, 1980:293); however, in 1982 Delise's original material was located in PC-
LENORMAND (Galloway et al., 1983ft; Galloway & James, 1986). The holotype cited above represents
material of this species from New Zealand that is named S. carpoloma in Delise's own handwriting
(Galloway & James, 1986 fig. 5). A specimen in G! is annotated by Delise 'Sticta carpoloma D. Delise Stict.
Specs de la Nouvelle Zelande (Donne a Delise par Bory de St- Vincent 1825)', and another in PC-HUE
labelled by Delise 'Sticta carpoloma Delise', has the annotation '206 Baie des iles-Nouv. Zeelande. D.
Lesson 1825'.

Sticta carpoloma is figured by Delise in plate 19 which is often missing from copies of the atlas illustrating
his monograph on Sticta. In a copy of the atlas presented by Delise to the Linnean Society of London on 25
August 1828, both the plate number and the names of the two species depicted (5. endochrysa and 5.

82

D. J. GALLOWAY

002064

-

Fig. 30 Sf/cto borneti. Holotype (G). Scale = 1 cm.

carpoloma) are marked on the page in Delise's hand (Galloway & James, 1986: fig. 5). 5. carpoloma was
collected in the Bay of Islands (North Auckland) in 1824. The corvette Coquille, captained by Louis-
Isidore Duperrey anchored in the Bay of Islands from 3-17 April 1824 (Lesson, 1829, 1839) and
both Lesson and D'Urville, botanists on the expedition, would have had opportunities to gather this
conspicuous and usually well-developed lichen which is still prominent today in coastal vegetation in
northern New Zealand. Bory de St- Vincent (1829: 236) records the species as Sticta corpoloma [sic] '6.
Sticte corpolome, Sticta corpoloma Delise, Stict. suppl. (non figure). M. Lesson a rapporte de nombreux
echantillons de cette belle espece, qui croit sur les vieux arbres autour de la Baie des lies de la
Nouvelle-Zelande', showing that although he was familiar with the description of the new species from
New Zealand and had seen Lesson's collections of it, he was unaware of its having been engraved and
figured in some copies of Delise's monograph.

After Delise's publication of 5. carpoloma, the concept of the species changed in several accounts,
introducing confusion to the correct circumscription of the taxon. Richard (1832) gave the name 5.
carpoloma to specimens of Pseudocyphellaria rufovirescens collected from Nelson, New Zealand (see
Galloway et al., 19836: 140), ignoring the yellow pseudocyphellae alluded to in Delise's description as a

PSEUDOCYPHELLARIA 83

good character for species separation. Hooker & Taylor (1844: 649) followed Richard's interpretation of 5.
carpoloma. Babington (1855: 276) gives a correct account of 5. carpoloma and makes the following
observations 'having exactly the habit of S. foveolata Del . , from which it can scarcely be distinguished when
the cyphellae are obsolete ... In spite of a certain amount of variation in the length and breadth of the
segments of the thallus, and of the more or less tomentose or quite naked under side, S. carpoloma is in
general easily recognised by its deeply lacunose and forked lobes: the apothecia and medullary stratum
agree with S. crocata . . . This plant does not bear soredia or coralline processes' . A fine coloured plate of a
large specimen, complete with anatomical drawings of apothecia, hymenium, and spores (Plate CXXVI)
accompanies Babington's account (see Fig. 29).

Both Nylander (18606: 339-40) and Hue (1890: 297, 1901: 83-84) introduced further confusion when
they recorded the photobiont of 5. carpoloma as being blue-green, and the species also having soredia or
isidia. Nylander recorded its range as being Chile, New Zealand, Tasmania, Polynesia, Java, and Reunion,
to which Hue added Japan. Such a circumscription refers not to P. carpoloma sens, str., but to P.
desfontainii (Delise) Vainio, and in part to P. lechleri and P. gilva from Chile.

Note 2: Sticta impressa J. D. Hook. & Taylor

Sticta impressa is one of a group of linear-laciniate, reticulate-faveolate species of Pseudocyphellaria, the
protologue stating in addition 'sorediis [=pseudocyphellae] pallide flavis punctiformibus convexis apice
impressis' (Hooker & Taylor, 1844: 648). Further, the species was reported by its authors as occurring in
'Lord Auckland's group, Campbell Island, Falkland Islands and New Zealand'. In choosing a lectotype of
material labelled S. impressa by Thomas Taylor (such material exists in BM, FH, and M), Galloway &
James (1980: 298) erroneously chose amply fertile material so labelled by Taylor from the Auckland Is
having white pseudocyphellae and containing physciosporin. Consequently, S. impressa was judged to be
Pseudocyphellaria faveolata and was accordingly incorporated in the synonymy of that species. However,
such a procedure overlooks an earlier typification of S. impressa by Miiller Argoviensis (1888: 137) who
chose material from New Zealand (collected by Joseph Hooker) and present in Taylor's personal
herbarium (FH). Miiller Argoviensis (loc. cit) states of 5. impressa 'simillima est Stictae Billardieri Del.,
sed pseudocyphellae definite sulphureo-flavae v. demum pallide flavae et minores, caeterum similiter
hemispherico-v. conico-prominentes.' The type of 5. impressa in Taylor's herbarium is annotated by
Taylor 'Sticta impressa Tayl. sorediis flavis (Sticta carpoloma Hook. Herb.) (Sticta ochracea Tayl.) New
Zealand, J. D. Hooker, 1844' (material collected at the Bay of Islands in 1841 by Hooker, was sent to
Taylor in 1844). In Taylor's herbarium other material labelled S. impressa comes from New Zealand,
collected by Dr Stanger, and by David Lyall (both are P. carpoloma); a specimen from New Zealand
collected by Dr Stanger is referable to P. billardieri and one collected by Hooker from Campbell Island is
P. faveolata. In Hooker's herbarium (BM) there are nine different specimens labelled 5. impressa by
Taylor; three are specimens of P. faveolata (J. D. Hooker collections from Auckland I., and from
Campbell I. , and a New Zealand collection of Dieffenbach), and six are of P. carpoloma (three collections
of Hooker, one of Stanger, and one of Andrew Sinclair from Great Barrier I. One is without collector's
name). A specimen of P. gilva (Ach.) Malme, collected by Lyall in the Falkland Is, is annotated S. impressa
by Taylor (BM!). Of this Taylor wrote to Hooker 'S. impressa H.f & T. 'of Falklands' I happen not to have
retained any such specimen - 1 suppose from New Zealand is not S. gilva Ach . . . S. ochracea Tayl. MSS I
know to be S. carpoloma' (Taylor, Correspondence, Royal Botanic Gardens, Kew Director's Correspond-
ence, 103 (98), 23 April, 1847).

Note 3: Sticta glaucolurida Nyl.

This taxon was lectotypified (Galloway & James, 1980: 293) on material collected by Knight and labelled
by him (although not annotated by Nylander) 5. glaucolurida Nyl. , from Leighton's herbarium now in BM.
Authentic material annotated by Nylander exists (H-NYL 33587) and consists of three minute scraps with
three apothecia loose in the packet. Nylander (1867: 438-9) noted of S. glaucolurida 'prope Stictam
granulatam in serie systematica locum habeat, sed affinis est Stictae physciosporae , a qua differt thallo
sublaevi latius diviso et pseudocyphellis determinatae citrinis'.

Note 4: Sticta borneti Mull. Arg.

Type material of Sticta borneti (Fig. 30) is labelled by Miiller Argoviensis as 5. impressa, no doubt after
1882, for six years later Miiller comments on New Zealand material of P. carpoloma (G 002600) under the
name Sticta impressa (1888: 137). Miiller Argoviensis (1894) lists S. borneti as a synonym of 5. impressa,
although he maintains S. carpoloma as a good species but following Nylander's interpretation has it with a
blue-green photobiont in the genus Stictina.

Morphology: Thallus irregularly spreading, loosely entangled in often extensive festoons,
corticolous, rarely saxicolous, 5-20(-50) cm diam. Lobes laciniate-elongate, plane to subcon-

84 D. J. GALLOWAY

cave-subcanaliculate, discrete to entangled, dichotomously branching, 3-8 mm wide and 1-7
cm long, tapering from centre to periphery, apices blunt or pointed, often furcate, or
shallowly notched or crenate. Margins entire, thickened below, with sparse to frequent,
verruciform, conical, yellow pseudocyphellae projecting, decorticate area strongly to weakly
pigmented. Upper surface bright lettuce green to olive green when wet, suffused brownish in
parts especially towards apices of lobes, pale greyish green, buff or glaucous-brownish, suffused
red-brown on storage in herbarium; strongly to obscurely faveolate, ridges rather smooth,
faveolae deep to shallow and somewhat inapparent, matt, smooth, isidia, phyllidia, maculae,
pseudocyphellae, and soredia absent, though in insect damaged specimens small irregular
regenerating lobules can develop at margins and from edges of damaged lamina. Medulla white.
Photobiont green. Lower surface, pale whitish or yellowish at margins, buff, brownish to dark
red-brown or blackened centrally, margins with a conspicuously and continuously raised
rim, wrinkled, bullate, glabrous to sparsely or densely tomentose, tomentum pale, silky at
margins, brown or black at centre. Pseudocyphellae scattered, very small. 0-1 mm diam. or less,
on ridges, verruciform, decorticate area flat, yellow, conspicuous. Apothecia marginal or
submarginal, subpedicellate, narrowly attached centrally, 0-5-3 mm diam., round to irregular
to somewhat compressed, disc matt, pale to dark red-brown when wet, black when dry,
imperf orate, epruinose, exciple pale flesh pink or whitish, pubescent at first, then roughened-
verrucose to scabrid, margins conspicuously dentate-striate often obscuring disc. Pycnidia
sparse to frequent, on upper surface, small hemispherical swellings 0-5 mm diam. or less, apical
pore minute, red-brown or black, surrounded by a pale marginal zone.

Anatomy: Thallus 120-200 um thick (to 350 um at margins). Upper cortex 25-30(-32) um
thick, dilute red-brown or yellow-brown in outermost 11-15 um, zone abutting photobiont layer
colourless, outer 2-4 um of cells necrotic, pigment dissolving in K, cells 4-5-7 urn diam.
Photobiont layer 18-22 um thick, densely compacted, photobiont green ? Dictyochloropsis , cells
rounded 2-5-3-5 um diam. Medulla 50-75 (-90) um thick, appearing grey-black and granular
in optical section (colour not changed in K), of loosely interwoven hyphae, to 4-5 um diam.,
encrusted with crystals. Pycnidia oval or rounded, 180 um diam. to 330 x 230 um. Lower cortex
18-23 um thick, lower 9-11 um pale yellow-brown, inner 9-12 um colourless, cells, 5-7 um
diam. Tomental hairs septate, simple, to 5 um thick, in fascicles, (50-)65-140(-210) um long,
often with spores entangled at apices. Apothecia: Exciple without photobiont, of colourless,
isodiametric, thick-walled cells arranged in parallel, radiating rows from centre to periphery, in
basal parts cell rows elongating to form short tomental hairs, cells round to irregular 4-5-9 um
diam., walls to 2-5 um thick, tissue 25-45 um thick at margins, 70-100 um thick near pedicel,
sometimes formed into coarsely irregular clumps (scabrosity visible in x 10 lens) Hypothecium
dilute yellow-brown, opaque, 45-65 um thick. Hymenium 80-90(-100) um tall, colourless;
epithecium pale red-brown or yellow-brown, 6-5-11-5 um thick paler in K. Asci cylindrical to
clavate, 70-78(-85) x 12-16 um. Ascospores grey-brown, oval-ellipsoid, thickened 1-septate,
septum 4-5-7 um thick, at maturity 3-septate (20-)22-25(-27) x 7-11 um.

Chemistry: Methyl evernate, tenuiorin, methyl lecanorate, methyl gyrophorate, evernic acid
(tr.), gyrophoric acid (tr.), hopane-7(3, 22-diol, hopane-6a, 7(3, 22-triol, 7|3-acetoxyhopane-6a,
22-diol (tr.), 6a, acetoxyhopane-7(3, 22-diol (tr.), norstictic acid (tr.), stictic acid, cryptostictic
acid, constictic acid, pulvinic dilactone, pulvinic acid, calycin, and several unidentified
compounds.

Distinguishing features: Pseudocyphellaria carpoloma is one of a group of dichotomously
branching, faveolate species well-developed in New Zealand. It has a white medulla, green
photobiont, and prominent yellow pseudocyphellae on the lower surface and projecting from
the margins. It has a distinctive chemistry [Code D of Wilkins & James (1979)].

Variation: In northern coastal forest where P. carpoloma is most commonly collected,
individuals are characteristically narrow-lobed with branching of lobes, at least at the
margins, dichotomous or subdichotomous, the whole thallus being somewhat open or loosely
entangled. In southern populations (from Banks Peninsula and South Westland) lobes are much

PSEUDOCYPHELLARIA 85

wider (1-2 cm wide in older, central parts) and much less obviously dichotomously branching,
although the apical lobes are furcate to subdichotomously branched.

Occasionally, a distinctive closely attached rosette-forming ecotype of P. carpoloma is
collected in which the lobes are attenuated and of equivalent length and breadth spreading
into rosettes 10-15 cm diam. In this form individual lobes are hard to distinguish except at the
margins which are sinuous-crenate and ascending and commonly also densely covered in small
squamiform-dentate phyllidia which seem not to be developed as a response to injury, but rather
to be induced by a particular microclimate and microhabitat. Such rosette forms are often also
densely fertile, and may have a distinctive white pruinosity developed on the apices of some
lobes. However, in all other respects (anatomically and chemically) the more compact,
rosette-forming ecotype is exactly similar to the normal dichotomously branching form of P.
carpoloma. An analogous situation exists in populations of P. faveolata (see below) with the
compact, rosette-forming ecotype formerly being designated as P. condensata. Separate species
designations for such ecotypes are not regarded as being either appropriate or necessary.

The particular combination of chemistry [Code D of Wilkins & James (1979)] and yellow
pseudocyphellae distinguishes P. carpoloma from P. faveolata [Code B of Wilkins & James
(1979), white pseudocyphellae, and apothecial discs which are white-pruinose at first and which
lack a scabrid exciple], from P. rufovirescens [Code A of Wilkins & James (1979), white,
punctate pseudocyphellae on a glabrous lower surface, and pale red-brown, epruinose
apothecial discs], and from P. billardierei [Code C of Wilkins & James (1979), white pseudocy-
phellae (not present at the lobe margins)] . P. physciospora is distinguished from P. carpoloma
by the entire margins which lack pseudocyphellae, the broader, and somewhat thinner lobes,
and by differences in chemistry (most notably the production of appreciable amounts of
norstictic and salazinic acids). See also Table 2 (p. 216).

Distribution: (Fig. 31) From the Three Kings Is (lat. 34S) southwards in mainly coastal
habitats from North Auckland to Kapiti I., in North I., and from Karamea Bluff (Nelson) to
Milford Sound west of the Main Divide, and Arthur's Pass, Banks Peninsula and Taieri Mouth
east of the Main Divide in South I. It occurs also on the Chatham Is. It has an altitudinal range
from sea level to 300 m. Known from the following offshore islands: Three Kings Is, Cavalli Is,
Poor Knights Is, Hen and Chickens Is, Little Barrier I., Great Barrier I., Rakitu I., Cuvier I.,
Rangitoto I., Kapiti I., and Chatham Is.

Habitat ecology: Pseudocyphellaria carpoloma is a common epiphyte on both bark and canopy
twigs of the following photophytes in northern coastal forest: Avicennia marina var. resinifera,
Beilschmiedia tar air e, Cordyline australis, C. kaspar, Dysoxylumspectabile, Elaeocarpus denta-
tus, Griselinia lucida, Kunzea ericoides, Leptospermum scoparium, Metrosideros excelsa, M.
robusta, Pittosporum eugenioides, Planchonella costata, Podocarpus totara, Rhopalostylis
sapida, Vitex lucens, and Weinmannia racemosa. In South I. localities it grows on Nothofagus
solandri var. cliffortioides and Podocarpus hallii. It has also been collected from basalt boulders
under Metrosideros excelsa, from andesite boulders, and from rocks in open-canopied coastal
forest. It favours similar ecological conditions to those in which P. aurata flourishes, and
associates with the same group of lichens in these habitats (see under P. aurata).

Specimens examined: 105.

Exsiccatae seen: Arnold (1894) Lichenes exsiccati 1199 [As Sticta glaucolurida Nyl., Neuseeland, Knight
(1886)]; Zahlbruckner & Redinger (1936) Lichenes Rariores Exsiccati 371. [As Sticta impressa Tayl. -
Nova Zelandia: Ins. septentrionalis, in montibus Tararua, ad corticem Nothofagi cliffortioidis et Weinman-
niae racemosae. Leg. H. H. Allan]. Material of this exsiccata was used by Magnusson (1940: 28-30) in his
account of P. carpoloma.

8. Pseudocyphellaria chloroleuca (J. D. Hook. & Taylor) Du Rietz Figs 32, 33

in Bot. Notiser 1924: 53 (1924). - Sticta chloroleuca^. D. Hook. & Taylor in Hook. Land. J. Bot. 3: 649
(1844). - Pseudocyphellaria freycinetii var. chloroleuca (J. D. Hook. & Taylor) Vainio in Hedwigia 38:
187 (1899). - Sticta freycinetii var. chloroleuca (J. D. Hook. & Taylor) Zahlbr. in Bih. K. svenska

86

D. J. GALLOWAY

-40

Norfolk I

Lord Howe I

Three Kings Is

The Snares

Auckland Is
\

Campbell I

X

Macquarie I
160

-42

44"

46'

166

i

168

170

172 C

I

174

'

176

178

100

34-

36-

40-

42-

Fig. 31 Distribution of Pseudocyphellaria carpoloma.

PSEUDOCYPHELLARIA 87

Vetensk. -Akad. Handl. 57(6): 18 (1917). Type: New Zealand, sine loco, No 2 ho, J. D. Hooker (FH!

- holotype; HM! - isotype) [see note 1].
Sticta psilophylla Mull Arg. in Bull. Soc. r. Bot. Belg. 31: 29 (1892). - Pseudocyphellaria psilophylla

(Mull. Arg.) D. Galloway & P. James in Lichenologist 12: 301 (1980). Type: New Zealand, sine loco,

C. Knight (G! - holotype).
Sticta variabilis var. cinerata Zahlbr. in Denkschr. Akad. Wiss. Wien math. - naturwiss. Kl. 104: 288

(1941). Type: New Zealand, North Island, Northland, Anawhata, in open Leptospermum shrubland

on L. scoparium at c. 120 m, March 1935, H. H. Allan ZA 513 (W! - lectotype).

Note 1: Pseudocyphellaria chloroleuca (J. D. Hook. & Taylor) Du Rietz

When Du Rietz combined Sticta chloroleuca J. D. Hook. & Taylor in Pseudocyphellaria, he did so without
having seen type material annotated by Taylor, and instead referred to material collected in New Zealand
by Berggren, Helms, Jelinek and Knight, the isidiate austral species P. glabra (J. D. Hook. & Taylor)
Dodge. Du Rietz (1924) correctly separated the isidiate P. glabra from the related non-isidiate P.
freycinetii, but mistakenly placed Sticta glabra as a synonym of S. chloroleuca, with the result that for many
years P. glabra was recorded in the literature as P. chloroleuca.

Galloway & James (1980) reinstated P. delisea (Delise) D. Galloway & P. James \Sticta delisea Delise is a
superfluous name (see p. 149) and hence Pseudocyphellaria delisea becomes a synonym of P. glabra} as a
good species [see also Hawksworth & Galloway (1984)] and included P. chloroleuca as a synonym,
lectotypifying in error Sticta chloroleuca from New Zealand material collected and labelled by Joseph
Hooker, but referable to P. glabra and not seen by Thomas Taylor who proposed the name chloroleuca. A
recent examination of the holotype of Sticta chloroleuca from Taylor's herbarium (FH) shows this (Fig. 32)
to be an independent Australasian species, quite distinct from P. glabra. The lectotypification of Galloway
& James (1980: 297) must therefore be rejected (Art. 8) in favour of the holotype material annotated by
Taylor. The taxon P. psilophylla (Mull Arg.) D. Galloway & P. James, is a later name for P. chloroleuca
and is reduced to synonymy with it (Galloway, 1986fo).

Morphology: Thallus orbicular to irregular spreading, 5-20(-30) cm diam., corticolous,
loosely attached centrally, margins and apices free. Lobes linear-elongate, very variable,
2-8( 12) mm wide, l-4(-8) cm long, rarely to 15 cm long, apices discrete, complex-imbricate
centrally, slightly convex to strongly canaliculate. Margins entire to ragged, lobulate-incised,
sparsely to densely isidiate, slightly to strongly revolute, ascending. Upper surface bright lettuce
green when moist, pale greenish grey often suffused red-brown towards apices when dry,
becoming yellow-brown on storage, irregularly wrinkled to shallowly faveolate, matt, minutely
scabrid-areolate (x 10 lens) in parts, also here and there minutely granular-papillate, isidiate,
without soredia, maculae, phyllidia or pseudocyphellae. Isidia mainly marginal and there often
densely developed, in small clumps laminally or originating from breaks and fissures, sometimes
densely covering thallus in a thick crust, simple, fingerlike at first often in groups of 3-5, then
branching and becoming coralloid, slender, fragile, 0-5-2 mm tall and 0-2 mm wide, terete or
somewhat flattened, eroding and leaving small pits. Medulla white. Photobiont green. Lower
surface white or pale buff-pink, glabrous, wrinkled-plicate to bullate, rarely very thinly and
patchily white-tomentose centrally, distinctly ridged at margins. Pseudocyphellae white,
minute, 0-1 mm diam. or less, numerous, inapparent, punctiform, flecklike, flat or slightly
punctate. Pycnidia rare to occasional, marginal and on thallus ridges, minute, punctiform,
red-brown, inapparent, not papillate. Apothecia sessile to subpedicellate, sparse to lacking,
marginal and laminal, margins obscuring disc at first, excluded at maturity, 0-5-3-5 mm diam.,
cupuliform, concave at first becoming subconvex at maturity, disc pale yellow-brown to dark
red-brown, matt to slightly glossy, epruinose, exciple white to pale flesh-coloured, translucent
when moist, delicately scabrid-verrucose.

Anatomy: Thallus 120-180(-210) um thick. Upper cortex 27-34 um thick, upper 7 um pale
yellow-brown, remainder colourless, cells 7 um to 9 um diam. Photobiont layer 22-27 (-34) um
thick, photobiont green, cells closely packed, to 7 um diam. Medulla 65-160 um thick,
colourless, hyphae 2-5 um thick. Lower cortex 15-20 um thick, pale yellow-brown, cells 2-7 um
diam. Apothecia: Exciple colourless, 90-135 um thick, cells 7-13-5(-18) um diam, in irregular
clusters towards margins. Hypothecium dilute yellow-brown to pale straw-yellow, 64-74 um
thick. Thecium 64-74 um tall, colourless; epithecium 7-14 urn thick, pale yellow-brown,
unchanged in K, tips of paraphyses overtopped by 4-5-9 um of colourless, amorphous gel,

D. J. GALLOWAY

Fig. 32 Sticta chloroleuca. Lectotype (FH). Scale = 1 cm.

Fig. 33 Pseudocyphellaria chloroleuca. North of Westport, Galloway (BM). Scale = 2 cm.

PSEUDOCYPHELLARIA 89

paraphyses 2 urn diam., apices swollen, moniliform, upper 10-12 um surrounded by a
yellow-brown gel. Asci 78-83 x 13-5-15-5 ^m. Ascospores brown, broadly ellipsoid-fusiform,
apices rounded or pointed, 1-3-septate, 25-30 x 6-8-9-1 um.

Chemistry: 7(3-acetoxyhopan-22-ol, hopane-7p\ 22-diol (tr.), hopane-15a, 22-diol, methyl
gyrophorate, and gyrophoric acid (C -I- red, often fugitive).

Distinguishing features: Pseudocyphellaria chloroleuca is an Australasian species characterized
by its pale, glabrous lower surface, a white medulla, green photobiont, fragile, marginal and/or
laminal, terete isidia (simple to coralloid), and a distinct but often fleeting C + red (gyrophoric
acid) reaction of the cortex. The lobes are linear-elongate, often somewhat ragged, and convex
to strongly canaliculate.

Variation: Pseudocyphellaria chloroleuca is a rather variable species, considerable differences
in lobe width and length, and in manner of branching being encountered. Isidia are primarily
marginal and at first are simple and finger-like, but later become coralloid-branched and
somewhat flattened. Being rather fragile they are readily fractured or eroded, and leave small
laminal pits which may be confused with pseudocyphellae. The pale, glabrous lower surface with
inapparent, white pseudocyphellae, and the C+ red cortical reaction readily distinguish P.
chloroleuca from several other similar species which lack these characters (Table 1), viz. P.
glabra, simple, terete isidia, a dark tomentose lower surface with prominent white pseudocy-
phellae, and a distinctive chemistry containing usnic acid and the stictic acid series of metabolites
accompanying Code A hopanes; P. corbettii, as for P. glabra but with marginal and occasionally
laminal, palmate phyllidia; P. episticta, laminal, white, scattered pseudocyphellae not derived
from isidia, marginal to laminal, simple to coralloid-branched, phyllidia, and a tomentose
lower surface, and with two hopane acids as medullary constituents; P. fimbriata, marginal
phyllidia which are apically white-tomentose, thicker, more coriaceous lobes which are densely
tomentose below; P. wilkinsii, as for P. episticta, but with two hopanes, and with fragile, terete,
simple to coralloid isidia and marginal and laminal phyllidia.

Distribution: (Fig. 34) Three Kings Islands (lat. 346'S) to Wellington in North I. , and Nelson
to Westland (lat. 4230'S) and Southland (lat. 4630'S) in South I., mainly northern coastal,
sea-level to 270 m. Also known from Lord Howe Island.

Table 1 Comparison of species similar to Pseudocyphellaria chloroleuca.

Character chloroleuca corbettii episticta fimbriata glabra wilkinsii

epithecium K + rose +

Thallus isidiate + +

Thallus phyllidiate + + + +

Margins tomentose +

Upper surface

pseudocyphellate + +

Cortex C + red +

Lower surface glabrous +

Lower surface tomentose + + + + +

15-acetoxy-22-hydroxyhopan-

24-oic acid +

15-22-dihydroxy-24-hopanoic

acid +

7-acetoxyhopan-22-ol + +

hopane-15a, 22-diol + + + +

Methyl gyrophorate +

Gyrophoric acid +

Stictic acid complex +

Usnic acid + +

90

D. J. GALLOWAY

Norfolk I

Lord Howe I

The Snares

Auckland Is

\

Campbell I

X

Macquarie I

160

170

40

-42

46

166

168 C

180

172

170 C

172

174

176

174 C

L_

176

I

178

36-

40-

42-

-44"

Fig. 34 Distribution of Pseudocyphellaria chloroleuca.

PSEUDOCYPHELLARIA

91

Habitat ecology: Pseudocyphellaria chloroleuca is a species of lowland, coastal forest and
occurs in partially shaded habitats to dry, open situation in full sunlight. Although mainly
epiphytic on trees or shrubs, it is also occasionally found on rocks in partial shade. It is known
from the following phorophytes: Beilschmiedia tawa, Dacry carpus dacrydioides, Elaeocarpus
hookerianus, Kunzea ericoides, Leptospermum scoparium, Metrosideros excelsa, Myrsine
aus tralis, Phyllocladus trichomanoides , Rhipogonum scandens , Rhopalostylissapida, and Wein-
mannia racemosa. It is commonly associated in such habitats with the following lichens:
Pseudocyphellaria aurata, P. poculifera, P. wilkinsii, Sticta squamata, S. latifrons, Usnea arida,
and U. rubicunda.

Specimens examined: 55.

9. Pseudocyphellaria cinnamomea (A. Rich.) Vainio Figs 35, 36

in Philipp. J. Sci. C. 8: 120 (1913). - Sticta cinnamomea A. Rich. , Voy. de I' Astrolabe, Bot. : 28, tab. 8
fig. 3 (1832). Type: New Zealand. South Island, Nelson [Novae-Zelandiae, Havre de V Astrolabe].
D'Urville, ex Herb. Bory de St- Vincent (PC-THURET! - holotype).
Sticta fragillima Church. Bab. in J. D. Hook., Fl. nov.-zel. 2: 279 (1855). - Stictina fragillima (Church.

HEBBI1R

Fig. 35 Sticta cinnamomea. Holotype (PC-THURET), and plate from Richard (1833). Scale = 1 cm.

92

D. J. GALLOWAY

Fig. 36 Pseudocyphellaria cinnamomea. New Zealand, J. D. Hooker (NY). Scale = 1 cm.

Bab.) Nyl., Syn. meth. lich. 1(2): 335 (1860). Type: New Zealand, sine loco, J. D. Hooker (BM! -

lectotype) [see note 1].
Sticta fragillima var. glaberrima (Laurer) Church. Bab. in J. D. Hook., Fl. nov.-zel. 2: 279 (1855).

Authentic material not seen.
Stictina fragillima f. lutescens Krempelh. in Nyl. in J. Linn. Soc. Lond. Bot. 9: 246 (1866). - Stictina

fragillima f. sublutescens (Krempelh.) Nyl. in Hue in Nouv. Archs Mus. Hist. nat. Paris, III, 2: 296

(1890). - Sticta fragillima f. lutescens (Krempelh.) Hellbom in Bih. K. svenska Vetensk.-Akad. Handl.

21 (3/13): 30 (1896). Type: New Zealand, Nelson, Tarndale, Dr A. Sinclair, ex Herb. W. L. Lindsay

(BM ! - lectotype ; BM ! , E ! - isotypes) .
Icon: Krempelhuber 1870. Tab XIV. 2. [as Sticta fragillima].

Note 1: Sticta fragillima Church. Bab.

Babington (1855: 279) remarks of his Sticta fragillima and its variety (3 glaberrima 'Very similar to S.
damaecornis in its ramification, and, like it, prone to considerable variation, as is apparent from the few
specimens which I have examined. The consistency of the thallus, however, is widely different, and is so
extremely brittle that most of the specimens are more or less mutilated. The central costa being more or less
distinct, is likewise a character of importance. The var. p cannot be satisfactorily separated; the lobes are
broader and shorter, and the colour is different; the under side likewise is sometimes, but not always, quite
smooth ... In all likelihood S. cinnamomea A. Rich. Fl. de la N. Zel. p. 28. pi. 8, f. 3. is a fine state of this
form. It does not seem, however, to be costate; the apothecia are represented as very dark, and having a
tolerably broad margin. I have seen no authentic specimens.'

In Babington's description of var. glaberrima which is based on Laurer's (1827) Sticta glaberrima
(a tropical species of Sticta which has nothing whatever to do with P. cinnamomea), he mentions
'cyphellis urceolatis' which seems to refer to species of Sticta. However, there is no blue-green
species of Sticta known from New Zealand remotely resembling P. cinnamomea, and it is likely that he
confused marginate pseudocyphellae with true cyphellae.

Thomas Taylor annotated New Zealand specimens of P. cinnamomea collected by J. D. Hooker and by
Dieffenbach with the unpublished name Sticta digitata Tayl. One Hooker specimen bears the following
note in Taylor's hand 'Sticta digitata n. sp. this occurred in Dr Hooker's Collection. I know not how its
character and description were omitted in the printing [obviously referring to the joint paper with Hooker,
Lichenes Antarctici of 1844]. I shall send them again.'

Morphology: Thallus irregularly spreading, 5-10(-15) cm diam., corticolous, muscicolous,
rarely terricolous, loosely attached centrally, margins and lobe apices free. Lobes linear-
elongate, straplike, 2-6(-8) cm long, usually rather narrow, 2-5(-8) mm wide, rarely to 15 mm
wide, subcanaliculate, irregularly to subdichotomously branching, often somewhat constricted

PSEUDOCYPHELLARIA 93

at branch points, sinuses wide-angled. Margins entire, noticeably ridged below, very rarely
secondarily lobulate or with scattered, white, punctiform pseudocyphellae. Upper surface dark
slate blue or yellowish blue, often with white or buff blotches or spots (photobiont-free zones),
pale grey to cinnamon-brown or yellowish when dry, plane, shallowly undulate to convex at
apices, centrally shallowly canaliculate, not cracked or wrinkled, often minutely white- or
buff -warty-papillate, smooth, matt or slightly glossy in parts, rather coriaceous, without soredia,
isidia, phyllidia or pseudocyphellae. Maculae minute (use x 10 lens) best seen when moist.
Medulla white. Photobiont Nostoc. Lower surface white to pale yellow-buff at margins, pale to
dark brown centrally, short, velvety tomentose from margins to centre, or with a narrow,
glabrous marginal zone, smooth to minutely wrinkled at margins, often with a prominent,
slightly darkened, raised midrib. Pseudocyphellae white, sparse to numerous, scattered,
minute, 0-05 to 0-2 mm diam., flecklike, very shallowly verruciform, margins slightly raised,
apparent only in larger pseudocyphellae, decorticate area flat to concave. Pycnidia occasional to
moderately common, scattered, laminal, often common towards lobe apices, shallowly
papillate, to 0-2 mm diam., apical ostiole red-brown to black, punctate-depressed. Apothecia
marginal, sparse or absent to occasional, sessile, constricted at base, 0-4-4 mm diam. , rounded,
concave at first, becoming plane to subconvex, disc pale to dark red-brown, smooth, matt,
epruinose, exciple whitish to pale flesh-coloured, translucent when wet, minutely verrucose-
areolate or granular, not tomentose, crenate at margins of disc.

Anatomy: Thallus 230-370 um thick. Upper cortex 34-45 um thick, outer 10-12 um pale
straw-yellow, remainder colourless, central row of cells to 14 um diam, outer and inner rows of
cells 4 5 -6 5 um diam. Photobiont layer 27-46 um thick , photobiont Nostoc in irregular clusters ,
cells to 7 um diam. Medulla 110-230 um thick, hyphae to 6 um diam. Lower cortex 34-40 urn
thick, outermost cells pale straw-yellow, remainder colourless, cells adjoining medulla com-
pressed, inner rows of cells 7-13-5 um diam. Tomental hairs 7-9 um diam., 45-275 um long,
single or in fascicles. Apothecia: Exciple 70-110(-140) urn thick. Hypothecium 45-55(-73) um
thick, dilute yellow-brown to hyaline, opaque, unchanged in K. Thecium 110-137 um tall,
colourless; epithecium 11-5-15 um thick, pale to dark yellow-brown or olive-brown, slightly
granular, unchanged in K. Asci 76-92 x 11-5-15-5 um. Ascospores pale yellow-brown,
fusiform-ellipsoid, 1-3-septate, apices rounded or pointed, straight or slightly curved,
25-32(-34) x 7-9 urn.

Chemistry: Gyrophoric acid (tr.), 7|3-acetoxyhopan-22-ol, hopane-7p, 22-diol (tr.), hopane-
15a, 22-diol.

Distinguishing features: Pseudocyphellaria cinnamomea is an Australasian species having
linear-elongate, straplike, rather narrow lobes, which are subcanaliculate, subdichotomously
branching, and with entire, non-isidiate margins. It has a white medulla, a blue-green photo-
biont, and the lower surface is distinctly costate. Apothecia occasional to rare or absent,
marginal, sessile, exciple whitish, minutely verrucose-areolate. Spores pale yellow-brown,
1-3-septate. It has a two-hopane chemistry [Code A of Wilkins & James (1979)]. It is a papery,
rather brittle species and in older parts gives the impression of being stalked. On prolonged
storage, specimens stain herbarium paper reddish brown.

Variation: Pseudocyphellaria cinnamomea is a rather uniform species, well-developed speci-
mens having wider (to 15 mm wide) lobes than the normal 2-5(-8 mm). It is closely related to P.
dissimilis but is distinguished from it by the entire, non-isidiate margins and the canaliculate
lobes having a well-defined midrib on the lower surface. P. allanii differs from P. cinnamomea
in having thick, coriaceous, sometimes minutely areolate-scabrid lobes with tomentose
margins and/or upper surface, and the densely tomentose lower surface which is non-costate.
Pseudocyphellaria insculpta, an Australian taxon differs from P. cinnamomea is having richly
divided to phyllidiate lobes, a punctate-impressed upper surface, and broader and longer
spores (27-)29-5-34(-36) x 9-13-5 um. Pseudocyphellaria fimbriatoides is distinguished from
P. cinnamomea by its marginal, hirsute phyllidia; P. murrayi by its dichotomously branched,
reticulate-faveolate lobes, with a pale sometimes glabrous lower surface and no midrib; and

94

D. J. GALLOWAY

Fig. 37 Distribution of Pseudocyphellaria cinnamomea.

PSEUDOCYPHELLARIA 95

P. junghuhniana, a widely distributed palaeotropical species, by the subdichotomously branch-
ing lobes which have a distinctive, dimpled, punctate-impressed upper surface.

Distribution: (Fig. 37) From North Auckland (lat. 35S) to Wellington in North I., and from
Nelson to Fiordland in South I., where it is most common west of the Main Divide; sea-level to
920m.

Habitat ecology: Pseudocyphellaria cinnamomea is a species of deep shade, most commonly
found on the bases of forest trees or shrubs, or on stones or boulders among mosses and ferns on
the forest floor or even growing directly on soil. It is best developed in densely shaded, moist,
humid habitats in high rainfall areas (it is especially rich in Westland and Fiordland), particularly
on rocks alongside streams, or in waterfall spray. It is epiphytic on the following phorophytes in
mainly lowland or coastal forest: Beilschmiedia tawa, Dacrycarpus dacrydioides, Dacrydium
cupressinum, Freycinetia baueriana ssp. banksii, Metrosideros excelsa, M. umbellata, Notho-
fagus menziesii, N. solandri var. cliff ortioides , and Rhipogonum scandens.

Specimens examined: 68.

10. Pseudocyphellaria colensoi (Church. Bab.) Vainio Fig. 38

in Result. Voy. S. Y. Belgica, Bot. Lich. : 28 (1903). - Sticta colensoi Church. Bab. in J. D. Hook. , Fl.
nov.-zel. 2: 274 (1855). - Sticta urvillei var. colensoi (Church. Bab.) Nyl., Syn. meth. lich. 1 (2): 360
(1860). - Lobaria colensoi (Church. Bab.) Trevisan, Lichenotheca veneta exs. 75 (1869). Type: New
Zealand, sine loco, J. D. Hooker (BM! - lectotype) [see note 1].

Sticta colensoi var. pinnatifida Church. Bab. in J. D. Hook., Fl. nov.-zel. 2: 274 (1855) PI. CXXIII. 5.
urvillei f. pinnatifida (Church. Bab.) Stizenb. in Flora, Jena 81: 177 (1895). - Lobaria colensoi f.
pinnatifida (Church. Bab.) Hellbom in Bih. K. svenska Vetensk.-Akad. Handl. 21 (3/13): 41 (1896).
Type: New Zealand, sine loco, W. Colenso (BM! - holotype) [see note 2].

Pseudocyphellaria neozelandica Dodge in Nova Hedwigia 19: 468 (1971). Type: New Zealand, Canter-
bury, road to Arthurs Pass, on Nothofagus, C. W. Dodge 61-B10 (? FH - not seen).

Note 1: Sticta colensoi Church. Bab.

Babington (1855: 274) writes of this conspicuous plant 'A perplexing and variable species, but I do not see
to what other it can be safely united: its nearest ally is perhaps 5. orygmaea (Raoul's specimen is named 5.
orygmaed) [= P. coronata], but in S. Colensoi the thallus is usually thicker, less minutely scrobiculated,
more divided, and differently coloured; the beautiful violet tint is wanting, though in decay the plant is
sometimes obscurely blotched with purple; the under side is less brightly-coloured and more even;
coral-like processes yellowish (rising from the gonimical stratum). The apothecia, however, perhaps afford
the best marks, as above indicated'.

Note 2: Sticta colensoi var. pinnatifida Church. Bab.

This name refers to highly divided, ragged forms of P. colensoi which are common throughout the range of
the species and without any taxonomic significance. A fine coloured engraving of this taxon (PI. CXXIII) is
given in Babington's account.

Morphology: Thallus orbicular to spreading, often pendulous, in irregular clustered clones,
8-20(-40) cm diam., corticolous, rarely saxicolous, loosely attached centrally, margins
free, ascending. Lobes linear-elongate to somewhat broadly rounded, 10-25(-30) mm wide,
5-12(-15) cm long, irregularly incised or notched, discrete at margins to complex-imbricate
centrally. Margins crenulate, notched, often torn or rather ragged, ascending to revolute,
slightly thickened and often eroded and yellow or orange-red pseudocyphellate below,
sparsely to densely isidiate. Upper surface bright lettuce green, yellow-green or tinged pale
bluish green when moist, pale green-grey becoming tawny yellow or golden-brown when dry
and on long storage, undulate, pitted, wrinkled-ridged or shallowly to deeply faveolate, often
cracked or deeply fissured, matt or glossy, minutely scabrid in part (use x 10 lens), without
soredia, maculae or pseudocyphellae, consistently isidiate. Isidia marginal and laminal,
scattered in small random groups or erupting in clusters or lines on ridges and along breaks in
thallus, minute, nodular or delicately fingerlike and fragile at first, soon becoming flattened,
bifurcating to coralloid, inflated at apices, verruciform-granular, often densely clustered on

96

D. J. GALLOWAY

Fig. 38 Pseudocyphellaria colensoi. A. New Zealand, Knight (WELT). Scale = 1 cm. B. Lake Howden,
Scott (COLO). Scale = 1 mm.

laminal ridges, at lobe margins or on apothecial exciple, often eroding apically and appearing
yellow, normally concolorous with thallus, sometimes becoming dorsiventral with distinct upper
and lower surface as in parent thallus. Medulla yellow, often appearing orange-red in thallus
breaks or cracks. Photobiont green. Lower surface pale yellow, yellow-grey to tawny at margins,
darkening to yellow-brown, red-brown to blackened centrally, thinly tomentose from margins
to centre, tomentum short, uniform to scattered arachnoid in parts, pale buff to brownish,
shallowly wrinkled to bullate. Cephalodia scattered in prominent, rounded warts, glabrous to
tomentose, 1-2 mm diam. Pseudocyphellae conspicuous, numerous, rounded, on prominent
verrucae raised above tomentum, 0-05-0-2(-0-3) mm diam., margins prominent, shining,
brown to red-brown, opaque, decorticate area flat to convex, yellow. Pycnidia scattered,

PSEUDOCYPHELLARIA 97

laminal or submarginal, punctiform, 0-1 mm diam., ostiole red-brown to black, often moribund
and excavate, appearing as small, scattered pinholes in rows or clusters. Apothecia rare to
numerous, pedicellate, submarginal or laminal, rounded, (l-)2-10 mm diam., often contorted
through mutual pressure, disc pale to dark red-brown or orange-brown, matt, epruinose,
margins sparsely to densely isidiate often also eroded and appearing yellow to orange-red,
exciple concolorous with thallus or slightly paler, sometimes appearing pale flesh-coloured,
conspicuously scabrid-areolate, often eroded-isidiate in well-developed specimens and
appearing yellow-pseudocyphellate .

Anatomy: Thallus 250-410(-540) um thick. Upper cortex uneven, 45-68 um thick, upper
10-18 um pale yellow-brown, remainder colourless, of thick-walled cells 1-5-3 um diam.
Photobiont layer 20-45 um thick , photobiont green , cells rounded , closely packed 3-7 um diam .
Medulla 155-300(-450) um thick, of loosely to compactly interwoven hyphae 4-5 um diam. , two
dense bands 90 um thick (200 um thick at pseudocyphellae) of hyphae encrusted with dark
red-brown crystals immediately below photobiont layer and above lower cortex. Lower cortex
27-35 um thick, layer (8-10 um) close to medulla pale yellow-brown, remainder colourless, of
thick-walled cells 1-5-3 um diam. Tomental hairs 5-7 um thick, 25-185 um long, pale
yellow-brown, in fascicles. Apothecia: Exciple 100 um thick at margins, 135-200 um thick below
disc, splitting away in ragged, pyramidal clusters (scabrosity), cells 4-5-9(-ll) um diam.,
thick-walled. Hypothecium 45-65 um thick, hyaline to pale straw-yellow or yellow-brown,
opaque, appearing slightly granular. Thecium 105-155 um tall, colourless; epithecium 11-16 um
thick, dilute yellow-brown, unchanged in K; paraphyses colourless to pale straw-yellow, apices
swollen, not moniliform. Asci 60-100 x 15-18 um. Ascospores hyaline to pale yellow-brown,
1-3-septate, oval-ellipsoid or broadly fusiform, apices pointed, straight or slightly curved,
(27-)32-36-5(-41) X9-11 urn.

Chemistry: Pulvinic acid, pulvinic dilactone, calycin, polyporic acid, 2a, 3(3, 22a-
triacetoxystictane, 22a-hydroxystictane-3-one, 2cx, 3|3-diacetoxystictane-22-ol, stictane-3(3,
22a-diol, 2a-acetoxystictane-3(3, 22a-diol, 3(3-acetoxystictane-2a, 22a-diol, stictane-2a, 3(3,
22a-triol.

Distinguishing characters: Pseudocyphellaria colensoi is a large, conspicuous Australasian
species having linear-elongate to broadly rounded lobes, with irregular notched, ragged, and
often deeply incised margins. The upper surface is undulate, pitted to reticulate-faveolate,
with marginal and laminal isidia scattered in groups or arranged in lines on ridges or along thallus
breaks. Isidia flattened, bifurcate to coralloid, verruciform-granular, often abraded apically
and appearing yellow. It has a yellow medulla, a green photobiont, and a wrinkled to bullate,
thinly tomentose lower surface. Apothecia are common (often numerous), large, to 1 cm diam. ,
pedicellate, with eroded-isidiate margins. Spores are colourless to pale yellow-brown, 1-3-
septate. The chemistry is dominated by stictanes (Chin et al., 1973) and the acetone extract is
yellow.

Variation: The major morphological variation seen in this species is in the size of the lobes
which may vary from 1-3 cm wide, and from 5-15 cm long. All other characters are constant,
although some variation is seen in the numbers of isidia and/or apothecia. P. colensoi is most
closely related to P. coronata; however, P. coronata differs in having broadly rounded, not
ragged-indented lobes, numerous, scattered, semi-immersed to sessile apothecia with dentate-
coronate margins, and a characteristic red-magenta extract in organic solvents.

Distribution: (Fig. 39) From Rangitoto I. , (lat. 3645'S) to Cook Strait in North I. , and from
Nelson to Southland in South I., also in Stewart I., coastal and inland, lowland to subalpine,
sea-level to 1500m.

Habitat ecology: Pseudocyphellaria colensoi is an epiphyte of trees and shrubs in a variety of
habitats, e.g. northern coastal forest, lowland podocarp-hardwood forest, upland Nothofagus
forest, Leptospermum scrub, and lowland to subalpine scrub. Although most luxuriantly
developed on tree boles, it also colonizes twigs and small branches. It prefers medium to high

98

D. J. GALLOWAY

-40 C

Norfolk I

Kermadec 30'
Is

Lord Howe I

Three Kings Is

The Snares

Auckland Is
\

Campbell I
Macquarie I

160

170

-42

-44 C

46

166

i

168 C

40-

180

172

176

170

L_

172 C

176 C

178

100

34 l:

36-

38-

40"-

,42-

Fig. 39 Distribution of Pseudocyphellaria colensoi.

PSEUDOCYPHELLARIA

99

light intensities and is thus best developed in open forests, at forest margins, and in the forest
canopy. In areas receiving frequent light rain or where mist or cloud cover is common,
specimens of P. colensoi may reach a great size and contribute a substantial amount to the forest
biomass. In humid, partly shaded sites, P. colensoi will grow on rocks below shrubs or small
trees. It is known from the following phorophytes: Avicennia marina var. resinifera, Buddleia
sp., Dacry carpus dacrydioides , Dacrydium cupressinum, Griselinia littoralis, Kunzea ericoides,
Leptospermum scoparium, Melicytus ramiflorus, Metrosideros robusta, Nothofagus fusca, N.
menziesii, N. solandri var. cliff ortioides , N. truncata, Phyllocladus alpinus, Podocarpus hallii, P.
totara, Pseudopanax, and Weinmannia racemosa. Often a dominant epiphyte, it associates with
species ofMenegazzia, Psoroma, Parmelia, Pseudocyphellaria, Sphaerophorus, and Usnea.

Specimens examined: 400.

11. Pseudocyphellaria corbettii D. Galloway, sp. nov.

Fig. 40

Diagnosis: Pseudocyphellariae homoeophyllae similis sed lobis rotundatis, 0-5-2-5 cm latis, margine dense
phyllidiatis, phyllidiis teretibus, coralloideus vel squamiformibus, cuneatus, flabellatus vel pectinatus.
Apothecia ignota. Typus: New Zealand, South Island, Nelson, Red Hills, 1050 m, on Leptospermum
scoparium, 30 December 1978, /. K. Bartlett (BM! - holotype; CHR- isotype). [TLC: 7(3-acetoxyhopan-
22-ol, hopane-7(3-22-diol (tr.), hopane-15a, 22-diol, usnic acid (tr.), hyperstictic (tr.), norstictic, stictic,
cryptostictic, constictic acids, and unidentified accessory compounds].

Fig. 40 Pseudocyphellaria corbettii. Holotype (CHR). A. Scale = 1 cm. B. Scale = 1 mm.

100 D. J. GALLOWAY

Morphology: Thallus orbicular to irregularly spreading, 5-10 cm diam. , corticolous or musci-
colous, closely attached centrally, free and ascending at apices. Lobes broadly rounded,
0-5-2-5 cm diam., discrete at margins, imbricate centrally, plane to irregularly undulate-
concave and appearing somewhat cochleate. Margins entire in part, to irregularly and
shallowly crenate or lacerate, becoming phyllidiate, sinuous, ascending in parts, in others
re volute, slightly thickened, sinuses prominent, semicircular. Upper surf ace lettuce green with a
yellowish tinge when moist, pale greenish or greyish yellow or buff, apices suffused red-brown
when dry, smooth or minutely wrinkled or impressed, matt, in places with minute solitary or
clustered pale fawnish spots or papillae, somewhat coriaceous, without soredia, isidia, maculae
or pseudocyphellae. Phyllidia mainly marginal, rarely laminal along breaks or tears in thallus or
in areas of insect damage, 0-5-1 mm tall, solitary to densely clustered, dorsiventral, rarely
terete, simple, bifurcating to coralloid, cuneate, flabellate to pectinate, upper surface
concolorous with thallus, apices often inflated, brownish, lower surface pale buff to brown,
glabrous or occasionally with small, white, flecklike pseudocyphellae. Medulla white. Photo-
biont green. Lower surface pale whitish or yellow-buff, glabrous in a wide marginal zone,
darkening towards centre and there densely tomentose, tomentum pale buff to brown, thick,
woolly, ragged, entangled, cortex smooth or slightly wrinkled-ridged. Pseudocyphellae white,
scattered, minute, flecklike close to margins, prominent, excavate 0-5-1-5 mm diam., round to
irregular centrally, often with a prominent margin, sunk in tomentum, decorticate area plane
or concave, white, granular. Apothecia not seen.

Anatomy: Thallus 180-270(-325) |im thick. Upper cortex 56-68 um thick, uppermost 2-3
rows of cells pale straw-yellow, compressed, cells 4-6 urn diam. , inner rows of cells larger, walls
thinner, 7-15-5 um diam. , more loosely arranged. Photobiont layer 34-45 um thick, photobiont
green, cells rounded with a prominent gelatinous envelope, to 5 ^im diam. Medulla 45-150 um
thick, colourless, hyphae 2-5-4 \im diam. Lower cortex 34-40 um thick, outermost cells pale
yellow-brown or straw-yellow, 4-7 urn thick, inner cells larger, thin-walled to 14 um diam.
Tomental hairs to 5 um diam. and to 270 um long, colourless.

Chemistry : as above .

This species is named for Prof R. E. Corbett, Emeritus Professor of Chemistry in the University
of Otago , Dudedin , New Zealand , for his researches into the chemistry of New Zealand lichens ,
and especially of the structural elucidation of hopane, stictane, and lupane triterpenoids in
species of Pseudocyphellaria from New Zealand.

Distinguishing features: Pseudocyphellaria corbettii is a distinctive, endemic phyllidiate spe-
cies in the P. freycinetii group having broadly rounded lobes, a smooth, somewhat coriaceous
upper surface with characteristic marginal phyllidia (simple, bifurcate to coralloid), a white
medulla, green photobiont, and a densely tomentose lower surface with conspicuous, white
pseudocyphellae. It has a distinctive chemistry of two hopanes [Code A of Wilkins & James
(1979)], the stictic acid assemblage of metabolites, and variable (often trace) amounts of usnic
acid in the cortex, the latter imparting a characteristic yellow-green colour to the upper surface.
Morphologically and chemically the species is related to P. glabra, P. freycinetii, and P.
homoeophylla. It is not known fertile.

Variation: At present the species is known only from two South I. , subalpine collections, and
so variation in lobe morphology and the development of phyllidia cannot be assessed. Pseudo-
cyphellaria corbettii is distinguished from the related South American taxon P. freycinetti
[larger, more papery lobes, with irregularly indented-lacerate, sinuous margins (see Galloway &
James, 1986)], the austral P. glabra (marginal and laminal, simple to furcate, terete isidia), and
the New Zealand endemic P. homoeophylla (longer and narrower lobes with entire, sinuous
margins lacking either isidia or phyllidia) by the development of marginal, dorsiventral phyllidia
(Fig. 8B).

Distribution: (Fig. 41) At present P. corbettii is known only from two localities in Nelson and
Canterbury in South I., but is undoubtedly of wider occurrence.

PSEUDOCYPHELLARIA

101

Lord Howe I

Norfolk I

Three Kings Is

Kermadec 30-
Is

The Snares

160

170 C

40

-42

46 c

166

i

168

i

180

174

176

178

170

172"

174 C

I

176

1

34-

36-

40-

42-

100

Fig. 41 Distribution of Pseudocyphellaria corbettii.

102

D. J. GALLOWAY

Habitat ecology: Pseudocyphellaria corbettii is a corticolous species known from the bark of
Leptospermum scopariwn (manuka) and Nothofagus solandri var. cliff ortioides (mountain
beech) in moderate to high-light habitats at forest margins, where it associates with the lichens
Leptogium azureum, Pseudocyphellaria homoeophylla, Sticta subcaperata, Usnea capillacea,
and U. xanthophana. Its detailed ecological requirements are still incompletely known.

Specimens examined: South Island. Nelson: Red Hills, 30 December 1978, /. K. Bartlett (BM,
CHR). Canterbury: Mt Faust above Boyle River, 11 January 1979, D. J. Galloway (BM, CHR).

12. Pseudocyphellaria coriacea (J. D. Hook & Taylor) D. Galloway & P. James

Fig. 42

in Lichenologist 12: 295 (1980). - Sticta coriacea J. D. Hook. & Taylor in Hook. Lond. J. Bot. 3: 648
(1844). -Ricasolia coriacea (J. D. Hook. & Taylor) Nyl. in Mem. Soc. Imp. Sci. not. Cherbourg 5: 103
(1857). - Lobaria coriacea (J. D. Hook. & Taylor) Trevisan, Lichenotheca veneta exs. 75 (1869).
Type: New Zealand, sine loco, W. Colenso 3754 (BM! - lectotype).

Sticta subcoriacea Nyl. Flora, Jena 48: 298 (1865). - Phaeosticta subcoriacea (Nyl.) Trevisan, Lichen-
otheca Veneta exs. 75 (1869). - Lobaria subcoriacea (Nyl.) Hellbom in BUi. K. svenska Vetensk. -

Fig. 42 Pseudocyphellaria coriacea. Holotype of Sticta canaliculata (WELT). Scale = 1 cm.

PSEUDOCYPHELLARIA 103

Akad. Handl. 21 (3/13): 41 (1896). Type: New Zealand, Otago, Saddle Hill Bush, on trunks of dead

trees, 26 October 1861, W. L. Lindsay (H-NYL 33660! - holotype: E! - isotype) [see note 1].
Ricasolia coriacea var. elaphocera Nyl. in Bull. Soc. linn. Normandie 2: 504 (1868). - Sticta elaphocera

(Nyl.) Stizenb. in Flora, Jena 81: 113 (1895). - Lobaria coriacea var. elaphocera (Nyl.) Hellbom in

Bih. K. svenska Vetensk. -Akad. Handl. 21 (3/13): 41 (1896). Type: New Zealand, sine loco, F. V.

Hochstetter 1859 (H-NYL 33376! - holotype) [see note 2].
Sticta canaliculata Knight in Trans. Linn. Soc. Lond. Bot. II 1: 282 (1877). Type: New Zealand, sine loco

(? Wellington), C. Knight (WELT Herb. Knight Vol. 13A p.ll! -holotype; BM!, H-NYL 33659!-?

isotypes) [see note 3].
Sticta coriacea f. vestitula Zahlbr. Denkschr. Akad. Wiss. Wien math.-naturwiss. Kl. 104: 282 (1941).

Type: New Zealand, North Auckland, Whangarei, on tree trunks, November 1935, W. A. Given

V166(W!-lectotype).

Note 1: Sticta subcoriacea Nyl.

This name, given by Nylander to narrow-lobed, elongated specimens of P. coriacea first appeared in Flora
(Nylander, 1865) and not in the paper on Lauder Lindsay's Otago lichens (Nylander, 1866) as recorded in
Galloway & James (1980: 295). Nylander's specimen has the following annotation 'On trees: Bush on top
of Saddlehill near Dunedin Otago. -New Zealand. Oct. 26, 1861. Dr Lauder Lindsay'.

Note 2: Ricasolia coriacea var. elaphocera Nyl.

This taxon was described in a footnote (p. 504) to Nylander's Conspectus synopticus Sticteorum (Nylander,
18686) where he states that variety elaphocera has a thallus branched as in Sticta damaecornis, i.e.
narrow-lobed and dichotomously branching. Galloway & James (1980: 295) are in error in assigning the
name to Nylander (1869).

Note 3: Sticta canaliculata Knight

Holotype material of this taxon (Fig. 42) was recently found in Knight's herbarium in New Zealand

(WELT), thus making Galloway & James's (1980: 295) typification of the name on BM material incorrect.

Morphology: Thallus orbicular to irregularly spreading, often pendulous in well-developed
specimens, corticolous, 5-15(-35) cm diam., loosely to closely adnate centrally, free and
ascending at margins and apices. Lobes linear-elongate and rather narrow, straplike, to broad
and rounded, shallowly concave to markedly canaliculate, 5-15(-20) mm wide, 3-5(-15) cm
long, subdichotomously to irregularly branching, discrete or contiguous from margins to
centre, to complex-imbricate, apices rounded, pointed or furcate. Margins entire, thickened-
ridged below, sinuous, ascendent to plane, conspicuously white-tomentose. Upper surface
bright lettuce green, suffused brownish at apices and margins when moist, pale grey-green to
buff when dry, becoming brown yellowish- or red-brown on storage, smooth, coriaceous,
scabrid-areolate in patches, shallowly undulate, occasionally minutely white-pubescent
especially towards apices and margins, without soredia, isidia, maculae, phyllidia or pseudo-
cyphellae. Cephalodia immersed, visible as small (0-1-1 mm diam.) swellings on upper and
lower surfaces. Medulla white. Photobiont green. Lower surface pale buff to chocolate brown,
tomentose from margins to centre, tomentum short and velvety to long, silky, entangled-
arachnoid, white and glistening to grey-buff. Pseudocyphellae numerous, starkly white, con-
spicuous, minute, flecklike, 0-1 mm diam., at margins, centrally 0-4-4 mm diam., flat to
deeply urceolate, margins distinctly raised, projecting above, or slightly sunk in tomentum.
Pycnidia sparse to frequent, mainly laminal, scattered, papillate, 0-1-1 mm diam. , apical ostiole
red-brown to black, punctiform to irregularly fissured. Apothecia sessile to subpedicellate,
sparse to frequent, mainly marginal or submarginal, occasionally laminal, 0-5-4 mm diam.,
rounded to irregular, often contorted through mutual pressure, disc pale to dark red-brown,
matt, epruinose, exciple pale buff or flesh-coloured, translucent when wet, coarsely corrugate-
scabrid, appearing dentate-coronate at margins, thinly pubescent to densely white-tomentose.

Anatomy: Thallus 300-500(-750) um thick. Upper cortex 56-76(-90) um thick, uppermost
11-14 um pale red-brown, cells compressed, surface roughened-uneven, remainder colourless
or pale straw-yellow, cells 4-5-11-5 um diam., walls thick. Photobiont layer 28-55 um thick, in
irregular, scalloped clumps of densely packed cells, photobiont green, cells round, to 7 um diam.
Medulla 130-300(-600) um thick, hyphae 4-5 um diam., encrusted with colourless, granular
crystals. Lower cortex 45-50 um thick, outermost 10-12 um dilute yellow-brown or straw-

104 D. J. GALLOWAY

yellow, remainder colourless, cells thick-walled, 6-9 \im diam. Tomental hairs 45-185 um long,
colourless, single or in fascicles, densely developed. Apothecia: Exciple 110-140(-230) um
thick, colourless, cells 4-5-14 urn diam., splitting into pyramidal groups in outer parts (scabros-
ity). Hypothecium pale straw-yellow, 45-65 um thick, opaque. Thecium 90-110 um tall,
colourless; epithedum 11-5-18 um thick, red-brown to yellow-brown overlain by a colourless,
amorphous gelatinous layer 9-11-5 um thick, epithecial pigments turning olive-grey in K,
persisting in a band below tips of paraphyses. Asci 70-75 x 18-22 um. Ascospores pale
yellow-brown to red-brown, broadly ellipsoid, apices pointed, 1-3-septate, locules irregular,
middle locule largest, remainder often irregularly lozenge-shaped, 27-30 x 7-9(-ll) um.

Chemistry: 7(3-acetoxyhopan-22-ol, hopane-7(3, 22-diol (tr.), hopane-15a, 22-diol.

Distinguishing features: Pseudocyphellaria coriacea is an endemic species having mainly
linear-elongate, rather narrow, straplike lobes which are subdichotomously to irregularly
branched, shallowly concave to strongly canaliculate, with enire margins, thickened below and
conspicuously white-tomentose. The upper surface is coriaceous, scabrid-areolate in parts
and occasionally white-tomentose towards apices and margins. It has a white medulla, a green
photobiont, and a pale buff to chocolate brown, densely tomentose lower surface with
prominent white, abruptly margined pseudocyphellae superficially resembling cyphellae.
Apothecia are sessile to subpedicellate, marginal, the exciple corrugate-scabrid, thinly
pubescent to densely white-tomentose. Epithecium red-brown or yellow-brown turning olive-
grey in K. Spores pale yellow-brown to red-brown, 1-3-septate, middle locule largest. The
chemistry is two hopanes [Code A of Wilkins & James (1979)].

Variation: There is considerable variation in lobe size in individuals of P. coriacea throughout
its range in New Zealand, and relatively broad-lobed specimens are collected, as well as long,
pendulous, narrow-lobed specimens. Lobes may be plane to markedly canaliculate, but in all
cases the coriaceous, scabrid-areolate upper surface and the marginal and laminal tomentum
are immediately recognizable. P. coriacea is one of a small group of endemic species charac-
terized by lobes with white marginal (and/or laminal) pubescence or tomentum. It is distin-
guished from P. allanii which has a blue-green photobiont (see Renner & Galloway, 1982), from
P. fimbriata which has marginal and occasionally laminal tomentose phyllidia, and from P.
pubescens which has a well-developed laminal tomentum, yellow pseudocyphellae below,
white-pruinose apothecial discs, and a differing and distinctive chemistry.

Distribution: (Fig. 43) North Auckland (lat. 3445'S to Cook Strait in North I., in South I.,
from Nelson to Southland both east and west of the Main Divide, and in Stewart I. , sea-level to
900m.

Habitat ecology: In the northern part of its range, P. coriacea is a common coastal species,
being found on tree bark in lowland forest, and on rocks and soil in habitats subject to moderate
to high light intensity. Elsewhere, especially in southern beech (Nothofagus) forests, it is a
prominent epiphyte of forest trees (mainly on trunks, rarely on twigs or in the canopy) and
shrubs, most noticeably where light is sufficient, at the forest margin. Also on successional
shrubs (especially Leptospermuni) . In Fiordland specimens often reach a great size, probably in
response to the favourable growth conditions, especially frequent precipitation, and a high
percentage of mist or cloud days throughout the year. Primarily a species of open situations, it
will also tolerate moderate shade and in partially shaded habitats it forms photosymbiodemes
with P. allanii. It is known from the following phorophytes: Avicennia marina var. resinifera,
Beilschmiedia tawa, Cordyline australis, Corynocarpus laevigatus, Dacrycarpus dacrydioides,
Dacrydium cupressinum, Griselinia littoralis, Kunzea ericoides, Leptospermum scoparium,
Metrosideros excelsa, M. robusta, Myrsine australis, Nothofagus fusca, N. menziesii, N. solandri
var. cliffortioides , Pinus radiata, Pittosporum eugenioides, Podocarpus hallii, Prumnopitys
ferruginea, P. taxifolia, and Weinmannia racemosa.

In northern coastal habitats P. coriacea associates with the following lichens: Heterodermia
japonica, H. obscurata, Psoroma allorhizum, P. araneosum, Pseudocyphellaria aurata, P.

PSEUDOCYPHELLARIA

105

Fig. 43 Distribution of Pseudocyphellaria coriacea.

106

D. J. GALLOWAY

montagnei, P. carpoloma, Parmotrema chinense, P. reticulatum, and Usnea rubicunda.

In southern lowland to subalpine habitats P. coriacea associates with: Anzia jamesii,
Brigantiaea chrysosticta, Lobaria adscripta, species of Menegazzia, Pseudocyphellaria colensoi,
P. episticta, P. glabra, P. homoeophylla, P. pickeringii, P. pubescens, and P. mfovirescens ,
species of Psoroma and Sphaerophorus, Usnea capillacea, and U. xanthophana.

Specimens examined: 165.

13. Pseudocyphellaria coronata (Mull. Arg.) Malme

Fig. 44

in Bih. K. svenska Vetensk. - Akad. Handl. 25 (6): 29 (1899). - Sticta coronata Mull. Arg. in Flora,
Jena 62: 163 (1879). Type: New Zealand, Greymouth, comm. F.v. Mueller (G! - lectotype).
Sticta hirta Stirton in Kept Trans. Glasgow Soc. Fid. Nat. 1: 23 (1873), non 5. hirta (Nyl.) Trevisan [1869]

Slicta orygmaea i.Wh iXyl. Syn. I p. Jtio,
ll.ilv l-'lura X. /.. \VI. 3 p. J-o.

Wellington t

Fig. 44 Pseudocyphellaria coronata. New Zealand, Knight (W). Scale = 2 cm.

PSEUDOCYPHELLARIA 107

(Art. 64-1). - Pseudocyphellaria hirta D. Galloway & P. James in Lichenologist 9: 95 (1977). Type:
New Zealand, Westland, Okarito, January 1872, /. Buchanan (OTA! - lectotype).
Sticta orygmaea var. calvescens Stirton in Trans. N.Z. Inst. 32: 73 (1900). Type: New Zealand,

Wellington, /. Buchanan (BM! - lectotype).

In several 19th century accounts this species from New Zealand and the subantarctic islands was known
as Sticta orygmaea Ach. (see Montagne, 1845; Taylor & Hooker, 1845; Babington, 1855), but this name
is a synonym of the South American Pseudocyphellaria berberina (Galloway & James, 1977). For a
discussion of the synonymy of P. coronata see Galloway & James (1977; 1980) and Galloway (19856).

Morphology: Thallus orbicular, rosette-forming, 5-12(-35) cm diam., loosely attached cen-
trally, margins free and often subascendent, corticolous, saxicolous. Lobes large, rounded,
(0-5-)l-5-3(-5) cm wide, discrete at apices, complex-imbricate centrally. Margins minutely
indented, crenate, becoming minutely lobulate to phyllidiate in parts, often densely so in centre
of thallus in well-developed specimens. Upper surface bright lettuce green, glaucous green with
a bluish tinge to yellow-green when moist, pale grey-green or yellow-green when dry becoming
cinnamon-brown to tawny and sometimes suffused red-purple on storage, wrinkled-faveolate,
faveolae rather small, numerous, shallow or deep, interconnecting ridges short, contorted,
sharply defined to smoothly rounded, matt or shining, occasionally minutely scabrid-areolate,
often irregularly cracked, without soredia, isidia, maculae or pseudocyphellae, consistently
phyllidiate. Phyllidia occasional to frequent at lobe margins and scattered over, or erupting from
lamina of large specimens, also on margins of mature apothecia, delicate, fragile, minute, to 0-2
mm tall, constricted at base, simple to subcoralloid, corticate on upper surface, pale yellow
below. Medulla yellow, often appearing orange-red in places. Photobiont green. Lower surface
yellow to pale yellow-buff, blackened centrally, complexly wrinkled-verrucose to bullate,
not or sparingly tomentose, tomentum sparse, scattered, in tufts, buff to dark brown, base of
apothecia visible as buff brown, glabrous, wrinkled protuberances often sunk below level of
lower surface. Pseudocyphellae yellow, numerous, 0-1-0-6 mm diam., verruciform-papillate,
round to irregular, flat to convex. Pycnidia occasional to frequent, scattered or clustered on
laminal ridges, minute, 0-1 mm or less diam. , papillate to punctiform, black. Apothecia sparse to
frequent, semi-immersed to sessile, scattered to tightly clustered, 0-1-3 mm diam., disc
red-brown, matt, epruinose, older fruits occasionally perforate, exciple concolorous with thallus
or suffused brownish, margins smooth at first and obscuring disc then becoming coronate-
dentate-phyllidiate, phyllidia often eroding and exposing yellow medulla.

Anatomy: Thallus 135-260(-450) urn thick. Upper cortex 26-36(-40) urn thick, pale straw-
yellow to yellow-brown, uppermost 7 um of stretched, compressed, necrotic cells, overlying
densely compacted thick-walled cells to 4-5 urn diam. Photobiont layer 18-22 um thick, lower
margin obscured by granular crystals of medullary hyphae,photobiont green, cells rounded, 5-7
um diam., chloroplast filling most of cell. Medulla 65-110(-270) um thick, hyphae to 4-5 um
diam., densely encrusted with granular red-brown crystals. Lower cortex 11-15 um thick,
colourless to pale straw-yellow, cells thick- walled, 3-5 um diam. Tomental hairs colourless, 9
um thick, 75-200 um long, rather sparsely scattered. Apothecia: Exciple cellular, colourless,
64-90(-185) um thick, cells 7-9 um diam., splitting away at margins to form ragged, pyramidal
clumps. Hypothecium 45-64 um thick, dilute orange-brown or yellow-brown. Thecium colour-
less to pale straw-yellow, 135-175(-200) um tall; epithecium 10-18 um thick, minutely granular,
red-brown or yellow-brown; paraphyses swollen at apices, 4-5 um thick, apical 3-5 cells
moniliform. Asci 76-90 x 15-5-23 um. Ascospores pale yellow-brown, 1-3-septate, ellipsoid,
apices rounded or pointed, straight or curved, cell contents distinctly vacuolate, 32-38-5(-43)
x9um.

Chemistry: Pulvinic acid, pulvinic dilactone, calycin, polyporic acid, 2a, 3(3, 22a-
triacetoxystictane, 22a, hydroxystictane-3-one, 2a, 3(3-diacetoxystictane-22-ol, stictane-3(3,
22cc-diol, 2a-acetoxystictane-3|3, 22a-diol, 3(3-acetoxystictane-2a, 22a-diol, stictane-2a, 3(3,
22a-triol, and a range of labile purple or purple-red acidic pigments.

Distinguishing features: Pseudocyphellaria coronata is a large, Australasian species having
broadly rounded lobes with minutely indented margins which are often also minutely phyllidi-

108 D. J. GALLOWAY

ate. The upper surface is markedly reticulate-faveolate with mature specimens having
scattered, often erumpent phyllidia. It has a yellow medulla, a green photobiont, and a pale
yellow to blackened lower surface which is mainly glabrous and only sparingly tomentose.
Apothecia are common, small (to 3 mm diam.), semi-immersed to sessile with dentate-
phyllidiate margins. Spores are pale yellow-brown, 1-3-septate. The chemistry is characteristic,
containing polyporic acid, unidentified pigments (red-magenta in organic solvents), and several
stictanes (Murray, 1952; Chin et al., 1973). The pigments in P. coronata occasionally impart a
purplish colour to thalli in the field, and specimens on long storage in the herbarium may also
become suffused purple. This phenomenon was observed by Babington (1855: 276) who
noted 'Here and there the surface (especially after being wetted) has a delicate purple-violet
tint, by which character the species may usually be recognized'; and by Lindsay (1869: 501) who
records 'The thallus is frequently stained in patches a beautiful purple, as I have noticed also in
various alpine lichens, such as Platysma cucullatum, Hoffm., which have a lemon-yellow thallus.
I attribute this coloration to the action of the ammonia produced by the decay of the droppings of
birds on the colorific principles of the lichen-thallus' .

Variation: Pseudocyphellaria coronata is a uniform species and, apart from a wide amplitude
of thallus size, the only variables appear to be in the numbers of apothecia and the quantities of
phyllidia developed. Pseudocyphellaria coronata is most closely related to P. colensoi, but is
distinguished from it by the rounded, regularly reticulate-faveolate lobes, the glabrous
lower surface, the semi-immersed, small apothecia with coronate-dentate margins, and the
red-magenta acetone extract. These characters also differentiate it from P. degelii, a non-
phyllidiate yellow-medulla species which has a markedly tomentose lower surface, entire lobe
margins, sessile to pedicellate apothecia with a well-developed, coarsely-scabrid exciple, and
colourless fusiform spores. Its acetone extract is yellow-orange and the medulla contains
seco-stictane triterpenoids (Goh etal., 1978).

Distribution: (Fig. 45) North Auckland (lat. 3520'S) to Cook Strait in North I., and from
Nelson to Foveaux Strait in South I. Also on the Chathams, Stewart I. , the Snares, the Auckland
Is, Antipodes I., and Campbell I., lowland to subalpine, sea-level to 850 m.

Habitat ecology: Pseudocyphellaria coronata is an epiphyte of trees and shrubs (it colonizes
both twigs and trunks) occurring in a wide variety of habitats from northern coastal forest
through lowland beech-podocarp, mixed podocarp-hardwood, to beech (Nothofagus) forest to
subantarctic lowland forest and subalpine scrub. It is a photophilous species and so reaches its
best development in the canopy, on tree trunks in open forest, and at the forest margins in areas
of moderate to high rainfall. It is probably the most common and widespread of the corticolous
species of Pseudocyphellaria in New Zealand, and because of its often great size and beauty (and
more recently of its potential as a source of dyestuffs), it has attracted much attention from
collectors over the past 150 years. In many forested areas, P. coronata is a significant component
of the forest ecosytem and contributes substantially to the forest biomass. In humid, semi-
shaded habitats P. coronata will successfully colonize rocks and stones on the forest floor where
there is sufficient light. It is known from the following phorophytes: Agathis australis, Coprosma
polymorpha, C. pseudocuneata, Dacrydium biforme, D. cupressinum, D. intermedium, Dacry-
carpus dacrydioides, Dracophyllum longifolium, D. subulatum, D. uniflorum, Fuchsia excorti-
cata, Griselinia littoralis, Kunzea ericoides, Leptospermum scoparium, Metrosideros excelsa, M.
umbellata, Myrsine australis, Nothofagus fusca, N. menziesii, N. solandri var. cliffortioides ,
Phyllocladus alpinus, Podocarpus hallii, P. totara, Rubus australis, and Weinmannia racemosa.

Specimens examined: 325.

14. Pseudocyphellaria crassaD. Galloway Fig. 46

in Lichenologist 15: 308 (1983). Type: New Zealand, South I., Nelson, West Bay, Lake Rotoiti, on
bark of Kunzea ericoides, 15 November 1977, D. J. Galloway (CHR 381001! - holotype, BM! -
isotype).

PSEUDOCYPHELLARIA

109

Fig. 45 Distribution of Pseudocyphellaria coronata.

110

D. J. GALLOWAY

Fig. 46 Pseudocyphellaria crassa. Isotype (BM). A. Scale = 1 cm. B. Scale. = 1 mm.

Morphology: Thallus orbicular to irregularly spreading, 6-12(-20) cm diam. , loosely to tightly
adnate centrally, margins free and spreading, ascending, corticolous. Lobes linear-elongate,
4-8(-15) mm wide, 2-5(-10) cm long, subdichotomously to irregularly branching, apices
discrete, contiguous from margins to centre or imbricate centrally, convex to plane, apices
acute, truncate or furcate, sinuses semicircular. Margins entire to minutely notched or incised
in parts, often minutely papillate and patchily tomentose, sinuous, subascendent, markedly
thickened above and below, with occasional, punctate, yellow pseudocyphellae, projecting dark
tomentum of lower surface often visible as a fringe. Upper surface livid greyish blue, suffused
reddish or red-brown when moist, pale grey, grey-buff, brownish or red-brown when dry,
irregularly reticulate-faveolate, uneven, lacunae deep or shallow, interconnecting ridges
sharply defined to smoothly rounded, coriaceous, glossy in parts, matt in others, rarely minutely
scabrid-areolate or patchily tomentose towards apices, soredia, isidia, maculae, phyllidia, and
pseudocyphellae absent. Medulla white. Photobiont Nostoc. Lower surface densely and thickly
tomentose from margins to centre, tomentum woolly, entangled, red-brown to dark chocolate
brown, apices whitish. Pseudocyphellae vivid yellow, conspicuous, moderately common to

PSEUDOCYPHELLARIA 111

frequent, raised above or slightly sunk in tomentum, papillate-verruciform, round to irregular,
0-1-1 mm diam., decorticate area flat or convex, margins conspicuous, slightly raised, red-
brown to black. Pycnidia sparse to frequent, mainly laminal, hemispherical, 0-2-0-5 mm
diam. , central ostiole black, minute, punctate-impressed. Apothecia sessile, constricted at base,
sparse to frequent, marginal or submarginal, concave at first with disc entirely obscured by
exciple, plane to convex at maturity, 0-5-3 mm diam. , solitary or crowded and slightly contorted
through mutual pressure, disc dark red-brown to black, matt, smooth to slightly papillate,
epruinose, exciple pale flesh-coloured, translucent when moist, coarsely verrucose-scabrid,
often slightly tomentose, markedly dentate-coronate at margins of disc.

Anatomy: Thallus 180-370(-460) ^m thick. Upper cortex 45-50 um thick, upper surface very
uneven, with scattered short hairs to 4-5 um diam. and 11-25 um tall, upper 11-18 um dilute
orange-brown, remainder colourless, cells thick-walled to 7 um diam. Photobiont layer 45-75
(-82) um thick, photobiont Nostoc, cells 4-5-7 ^m diam. Medulla 36-73(-230) um thick,
colourless. Lower cortex 23-30 um thick, outermost 7-9 um dark red-brown, inner zone
colourless, cells as in upper cortex. Tomental hairs pale straw-yellow to dark brown, to 7 um
thick and 70-350 um long. Apothecia: Exciple 80-150 ^m thick, colourless, cells thick-walled,
7-18 um diam., splitting away at margins into irregular, pyramidal clumps (scabrosity).
Hypothedum 45-55 um thick, pale yellow-brown, opaque. Thecium 100-120 um tall, colourless
to pale straw-yellow; epithecium 11-14 um thick, yellow-brown to olive-brown, unchanged in K,
overlain by a thin (4-5-7 um thick), colourless, amorphous gel; paraphyses colourless, capitate
at apices which are covered with an olive-brown gel, not dissolving in K. Asci 86-108 x
18-20 um. Ascospores olive-brown to dark red-brown, thickened 1 -septate to 3-septate, oval-
ellipsoid, apices rounded or pointed, septum of 1-septate spores 2-3 um thick, (23-)25-30
(-32) x 9-11 urn.

Chemistry: Methyl evernate (tr.), tenuiorin, methyl lecanorate (tr.), evernic acid (tr.),
gyrophoric acid (tr.), four unidentified depsides (tr.), hopane-7(3, 22-diol, hopane-6cx, 7(3,
22-triol, 7(3-acetoxyhopane-6a, 22-diol (tr.), 6a-acetoxyhopane-7|3, 22-diol (tr.), five un-
identified triterpenoids, norstictic (tr.), stictic, cryptostictic, and constictic acids, pulvinic acid,
pulvinic dilactone, and calycin.

Distinguishing features: Pseudocyphellaria crassa is an endemic species having linear-
elongate, subdichotomously to irregularly branching lobes with entire to minutely notched,
thickened margins with projecting, punctate, yellow pseudocyphellae; the upper surface is
noticeably reticulate-faveolate, the faveolae round to irregular, shallow to deep, without
soredia, isidia, maculae, phyllidia or pseudocyphellae. It has a white medulla and a blue-green
photobiont. The lower surface is thickly felted-tomentose, red-brown to dark chocolate brown,
with conspicuously vivid, yellow pseudocyphellae. Apothecia are marginal, occasional to
frequent, the disc dark red-brown to black, epruinose, with a coarsely verrucose-scabrid,
dentate-coronate exciple. Spores are olive-brown to dark red-brown, thickened, 1-septate to
3-septate. It has a complex chemistry [Code D of Wilkins & James (1979)].

Variation: This species shows surprisingly little morphological variation throughout its known
range in New Zealand, although the production of apothecia seems not especially common, with
the majority of specimens examined being sterile. It is one of a group of species having
linear-elongate lobes and a reticulate-faveolate upper surface, though its thick, dark, felted
tomentum, yellow pseudocyphellae, and blue-green photobiont clearly set it apart from other
taxa in the group (P. billardierei, P. carpoloma, P. faveolata, P. maculata, P. physciospora, and
P. rufovirescens). It has a chemistry identical to that of P. carpoloma, a green photobiont species
most common in northern coastal forest which has thinner lobes and a much less thickly
developed tomentum. Pseudocyphellaria crassa is distinguished from P. maculata (primarily a
terricolous, subalpine grassland species) by the thickness of the lobes, the absence of maculae on
the upper surface, the dense tomentum of the lower surface, and by the chemistry. P. crassa is
distinguished from P. physciospora by the narrower, thicker lobes, prominent, marginal
pseudocyphellae, a thicker, darker tomentum on the lower surface, and a different chemistry. In

D. J. GALLOWAY

Fig. 47 Distribution of Pseudocyphellaria crassa.

PSEUDOCYPHELLARIA 113

earlier accounts (Galloway & Simpson, 1978; Galloway & James, 1980; Galloway etal., 19836)
P. crassa was incorrectly recorded as P. neglecta (see also Galloway, 19836).

Distribution: (Fig. 47) Most commonly collected in New Zealand in North Canterbury and
Nelson between lats 41S and 42S, but recorded once from Wellington (lat. 39S) and from Peel
Forest, South Canterbury, and from localities near Haast Pass in Otago, from 300-1050 m.

Habitat ecology: Pseudocyphellaria crassa is a subalpine species growing as an epiphyte on
successional shrubs and trees (especially Kunzea and Leptospermum) in mountainous areas of
Nelson where it is best developed. It colonizes both bark and twigs of Kunzea ericoides and at
West Bay, Lake Rotoiti, it is commonly found at the base of this tree and also overgrowing rocks
and soil in the vicinity. Apart from Kunzea and Leptospermum, which seems its preferred
substrates, P. crassa is known from the following phorophytes: Aristoteliafruticosa, Nothofagus
fusca, N. menziesii, Plagianthus regius, and Populus (Hanmer Springs in an exotic plantation).

Specimens examined: 25.

15. Pseudocyphellaria crocata (L.) Vainio Figs 48, 49

in Hedwigia 37: 34 (1898). - Lichen crocatus L., Mantissa Alter.: 310 (1771). - Lobaria crocata (L.)
Rauschel, Nomenclat. Bot. 3rd ed.: 330 (1797). - Sticta crocata (L.) Ach., Meth. Lich.: 211 (1803). -
Stictina crocata (L.) Nyl., Syn. meth. lich. 1(2): 338 (1860). - Saccardoa crocata (L.) Trevisan,
Lichenotheca veneta exs. 75 (1869). - Cyanisticta crocata (L.) Gyelnik in Feddes Reprium Spec. nov.
veg. 29: 7 (1931). Type: India, sine loco, Konig (LINN 1273. 137! - holotype) [see note 1].

Pulmonaria aurigera Bory in Floerke in Magazin Ges. naturf. Fr. Berl. 2: 126 (1809). - Sticta aurigera
(Bory) Delise in Mem. Soc. linn. Normandie 2: 54 (1825). - S. mougeotiana var. aurigera (Bory) Nyl. ,
Syn. meth. lich. 1 (2): 341 (1860). - Cyanisticta mougeotiana var. aurigera (Bory) Szatala in Annls
hist.-nat. Mus. natn. Hung. 7: 41 (1956) nom. inval. (Art. 33.2). - Cyanisticta aurigera (Bory) Dodge
in Beih. Nova Hedwigia 12: 171 (1964). Type: Les arbres des forets montagneuses des lies de France
(Mauritius) et de Mascareigne, Herb. Bory de St- Vincent (PC-THURET! - lectotype) [see note 2].

Sticta aurigera var. nuda Delise in Mem. Soc. linn. Normandie 2: 55 pi. 3 fig. 9 (1825)^. - Cyanisticta
aurigera var. nuda (Delise) Dodge in Beih. Nova Hedwigia 12: 172 (1964). Type: lies de France
(Mauritius) et Mascareigne, IBory de St-Vincent, ex Herb. Bory (PC-THURET! - lectotype).

Sticta mougeotiana Delise in Mem. Soc. linn. Normandie 2: 62 pi. 5 fig. 13 (1825). - Stictina mougeotiana
(Delise) Nyl., Syn. meth. lich. 1(2): 340 (1860). - Pseudocyphellaria mougeotiana (Delise) Vainio in
Hedwigia 37: 36 (1898). - Merosticta mougeotiana (Delise) Clements, Gen. Fungi: 175 (1909). -
Cyanisticta mougeotiana (Delise) Dodge in Beih. Nova Hedwigia 12: 177 (1964). Type: lie de France
(Mauritius), Bory de St-Vincent (PC-THURET! - lectotype) [see note 3].

Sticta mougeotiana var. xantholoma Delise in Mem. Soc. linn. Normandie 2: 63 pi. 5 fig. 14 (1825). -
Stictina mougeotiana var. xantholoma (Delise) Nyl. Syn. meth. lich. 1(2): 340 (1860). - Pseudocyphel-
laria xantholoma (Delise) Dodge, Hancock Pacific Expeditions 3: 37 (1936). - Cyanisticta xantholoma
(Delise) Dodge in Beih. Nova Hedwigia 12: 178 (1964). Type: Ex Insulae Borboniae ou Franciae, sine
collectoribus nomine (PC-LENORMAND! - lectotype).

Sticta leucosticta Pers. in Gaudichaud, Voy. Uranie Bot.: 200 (1827). - Stictina crocata f. leucosticta
(Pers.) Nyl. in Hue in Nouv. Archs Mus. Hist. nat. Paris. 3,2: 296 (1890). - Sticta crocata f. leucosticta
(Pers.) Zahlbr., Cat. Lich. Univ. 3: 378 (1925). Type: In insulis Sandwicensibus [Hawaii] (adtruncos),
Gaudichaud (not seen).

Sticta xanthosticta Pers. in Gaudichaud, Voy. Uranie Bot.: 201 (1827). Type: In insulis Sandwicensibus
(ad arborum truncos), Gaudichaud (not seen).

Sticta citrina Pers. in Gaudichaud, Voy. Uranie Bot. : 201 (1827). - S. crocata f. citrina (Pers.) Zahlbr.,
Cat. Lich. Univ. 3: 377 (1925). - Cyanisticta citrina (Pers.) Gyelnik in Feddes Reprium Spec. nov. veg.
29: 1 (1931). Type: In insulis Maclovinus [Falkland Islands] (obtegit rupes), Gaudichaud (not seen).

Cyanisticta subcrocata Gyelnik in Feddes Reprium Spec. nov. veg. 29: 7 (1931). Type: Africa australis
(Drege, sub St. crocata v. gilva, pr. p. Spec. orig. in herb. mus. nat. hist. Wien) (not seen).

Cyanisticta epiflavoides Gyelnik in Annls Cryptog. exot. 4: 170 (1931). Type: U.S.A., Maine, Mt Desert
Island, Witch Hole, on maple, C. C. Plitt 7-20-28. (BP- not seen).

Cyanisticta positiva Gyelnik in Rev. Bryol. Lichenol. 5: 31 (1932). Type: U.S.A., Maine, Mt Desert
Island, Witch Hole, on tree trunk, Plitt, sub Sticta crocata (BP herb. Szatala - not seen).

114

D. J. GALLOWAY

""*..

Fig. 48 Pseudocyphellaria crocata. A. New Zealand, Knight (WELT). Scale = 1 cm. B. Rangitoto Island,
Allan W1(W). Scale = 2 mm.

Note 1: Pseudocyphellaria crocata (L.) Vainio

Pseudocyphellaria crocata is based on Linnaeus's Lichen crocatus, a name given to an Indian plant
collected by Konig (see Galloway & James, 1980: 296, fig. 2). The type has a smooth, shining surface with
only slight faveolation visible; the lobes are narrow and much divided, rather sinuous with irregularly
raised sorediate margins; the soralia are golden yellow, farinose, and are also sparingly present on the
upper surf ace.

Note 2: Pulmonaria aurigera Bory

Pulmonaria aurigera is a synonym of Pseudocyphellaria crocata. It has a relatively broad-lobed, undulat-
ing, but not faveolate-ridged thallus (Fig. 49) with prominent, yellow, notably delimited (i.e. not in any
reticulate or confluent pattern), marginal and laminal soralia. In var. nuda Delise, the upper surface is
slightly ridged-reticulate and the soralia are primarily marginal. The type specimens of both taxa are
illustrated and discussed in Galloway & James (1986: figs 11, 12). Delise (18250) is in error in ascribing the
basionym of this species to Lichen aurigerus Bory (Bory, 1804), and appears not to have seen Floerke's
(1809) work where Bory's description of Pulmonarea aurigera is validated (see Galloway & James, 1986).
Bory's description states 'frondibus laciniato-lobatis submembranaceis, verrucis aureis subtus, supra et in
marginibus sparsis'.

PSEUDOCYPHELLARIA

115

Fig. 49 Pseudocyphellaria crocata. North branch of Routeburn, Galloway (CHR 367876). Scale = 5 mm.

Note 3: Sticta mougeotiana Delise

Sticta mougeotiana closely resembles the holotype of Lichen crocatus L. , in the Linnean Herbarium (LINN
1273.137). It has an undulate to occasionally ridged, subfaveolate upper surface with mainly marginal
yellow soralia, though occasional laminal soralia are found as well. In earlier accounts of New Zealand
lichens (e.g. Allan, 1949; Martin & Child, 1972) specimens with long, sinuous, marginally sorediate lobes
were referred to this taxon (see also Magnusson, 1940: 19). The types of Sticta mougeotiana and the var.
xantholoma are discussed and illustrated in Galloway & James (1986: figs 26, 27).

Morphology: Thallus very variable in size , shape , and thickness , rosette-forming to irregularly
spreading, corticolous, muscicolous, terricolous, and saxicolous, loosely to closely attached
centrally, margins free. Lobes (0-2-)0-5-l(-3-5) cm wide and (l-)3-5(-7) cm long, broad
and rounded, plane, to narrow, canaliculate, deeply laciniate and discrete from margins
to centre, to imbricate-complex. Margins entire, sinuous, subascending to indented, ragged,
incised, crenulate, slightly thickened below, sorediate, soralia punctiform to linear, often
distinctly labriform and eroding lower surface, soredia yellow, farinose to coarsely granular.
Upper surface pale to dark slate blue, often suffused brownish or reddish when moist, pale
fawnish grey to glaucous grey or brownish yellow suffused red or brown in parts or completely
brownish or red-brown when dry, plane, undulate, shallowly and irregularly wrinkled to densely
faveolate, ridges sharply defined to smoothly rounded, smooth, slightly coriaceous, matt to
shining in parts, occasionally slightly and irregularly short tomentose, without isidia, phyllidia or
pseudocyphellae; maculate and sorediate. Maculae occasional to frequent, best seen in
faveolate specimens, delicate, white, distinctly reticulate, following ridges and in faveolae (use
x 10 lens). Soredia common, laminal and marginal, very variable, randomly scattered in erose,
rounded soralia to 3 mm diam. , densely farinose yellow, or linear-elongate at lobe margins,
or erupting from pseudoisidiate warts on laminal ridges, central parts eroding yellow-granular,
marginal pseudoisidia granular, brownish, not truly corticate as in P. desfontainii, often
coalescing in lines along ridges, rarely developing into small, corticate, cochleate lobes. Medulla
white. Photobiont Nostoc. Lower surface pale yellow-white to buff at margins, red-brown to
black centrally, wrinkled-undulate to +bullate, tomentose from margins to centre, or with a
narrow, glabrous marginal zone, often erose-sorediate, tomentum short, velvety, thin and
scattered to thick and woolly, pale buff to dark red-brown or black. Pseudocyphellae scattered,
occasional to frequent, 0-1-1-2 mm diam., flat to convex or very shallowly papillate, sunk in

116 D. J. GALLOWAY

tomentum or slightly projecting above. Apotheda occasional to rare or absent, marginal and
laminal, solitary or clustered (2-5 together), 0-3-2-5 mm diam., round to irregular through
mutual pressure, shallowly to deeply cupuliform, undulate, subconcave at maturity, disc pale
to dark red-brown, matt, smooth, epruinose, exciple pale flesh pink, translucent when moist,
obscuring disc at first, rupturing and leaving an irregular, dentate margin, conspicuously
verrucose-areolate, occasionally minutely tomentose towards base.

Anatomy: Thallus 185-310(-400) urn thick. Upper cortex 34-40 um thick, pale straw-yellow,
outermost cells slightly darker, cells thick-walled, 3-7 urn diam. Photobiont layer 45-73 um
thick in irregular clumps, photobiont Nostoc. Medulla 64-230 um thick, colourless. Lower
cortex 23-28 um thick, pale straw-yellow, outermost layer red-brown, cells 7-11 um diam.
Tomental hairs red-brown, 4-5 um thick, to 140 um long. Apotheda: Exciple colourless, 80-140
um thick, cells thick-walled, 4-5-15 um diam. Hypothedum 45-65 um thick, opaque yellow-
brown to red-brown, intensifying in K producing sheafs of red crystals (?norstictic acid).
Thedum colourless to pale straw-yellow, 80-100 um tall; epithedum 11-23 um thick, dark
yellow-brown to olive-brown, pale or dissolving in K. Asci 77-82 x 13-15 um. Ascospores
broadly ellipsoid, smoky olive-brown to dark brown, thickened 1-septate to irregularly 3-
septate, straight or slightly curved, septum of thickened spores variable, 6 um thick or less,
22-5-27(-29-5) x 9 um.

Chemistry: A-Tenuiorin, methyl gyrophorate, gyrophoric acid (tr.), hopane-7|3, 22-diol (tr.),
hopane-6a, 7(3, 22-triol, 7|3-acetoxyhopane-6a, 22-diol (tr.), 6ct-acetoxyhopane-7(3, 22-diol
(tr.), physciosporin (tr.), norstictic (tr.), stictic, cryptostictic (tr.), and constictic acids, pulvinic
acid, pulvinic dilactone, and calycin (see Maass, 19756 ). B - as above but with the replacement
of stictic, cryptostictic, and constictic acids by consalazinic and salazinic acids. This latter
chemistry occurs in about 20% of specimens examined. These two chemodemes also occur in P.
neglecta.

Distinguishing features: Pseudocyphellaria crocata is a cosmopolitan species having a white
medulla, a blue-green photobiont, yellow soralia (both laminal and marginal) on the upper
surface, and yellow pseudocyphellae on the lower surface. It has a well-defined chemistry (see
above) with two chemodemes present in the total population.

Variation: Pseudocyphellaria crocata is probably the most common species of Pseudocyphel-
laria in New Zealand and has a very variable morphology. The two extremes appear to be
individuals with broadly rounded lobes (Fig. 49), a smooth upper surface, and with marginal
and/or scattered, rounded laminal soralia (aurigera-typQ, see Galloway & James, 1986), to
individuals with linear-elongate, narrow lobes (Fig. 48) and a reticulate-faveolate upper
surface, with scattered, laminal and/or marginal soralia (mougeotiana-type, see Galloway &
James, 1986). A complete intergradation exists between these two extremes, and it is often
difficult to make a strict segregation of individuals with a even, plane surface from those
having shallowly to deeply defined faveolae. Soralia are also very variable in both position and
density, ranging from exclusively marginal, either labriform, granular-subcorticate or powdery
erose-sorediate, to laminal, especially in faveolate forms where soralia are confined to ridges
and also to inter-ridge lacunae, to smooth-surfaced forms where soralia are randomly distri-
buted on the upper surface. In contrast to this considerable variation in thallus morphology, the
chemistry of the species is remarkably uniform. Colour of the upper surface varies from
glaucous-grey in shaded populations to dark red-brown or blackened in forms exposed to full
sunlight. A large number of the specimens examined (and it is true also for the species in the
field) were sterile, though apothecia are occasionally found. As a general rule apothecia in P.
crocata are produced much less frequently than in the closely related P. neglecta.

Pseudocyphellaria crocata is one of a primarily palaeotropical group of species loosely defined
by the characters white medulla, blue-green photobiont, and yellow pseudocyphellae (see
Magnusson, 1940) but is distinguished from the related endemic species P. sericeofulva by
lacking the small, imbricate cochleate lobes and the tomentum of the upper surface. Two other
taxa may be confused with P. crocata: (i) P. desfontainii which is similar in all respects to P.

PSEUDOCYPHELLARIA

111

176

Norfolk I

Lord Howe I

Three Kings Is

The Snares

Auckland Is

Campbell I
Macquarie I

160

-40 C

-42

-44 C

46

166 C
I

168

170

172 C

174 C

i

176 C

I

178

34-

36-

100

40-

42-

44-

Fig. 50 Distribution of Pseudocyphellaria crocata.

118 D. J. GALLOWAY

crocata but has, instead of soralia, fragile, terete isidia. It is known from South Africa, the
islands of the Indian Ocean, Sri Lanka (it is conspecific with P. ceylonensis Magnusson), New
Caledonia, and north-eastern Australia; (ii) P . neglecta (referred to as P. australiensis in earlier
accounts) which is marginally, occasionally laminally phyllidiate, lacks soralia, and as a general
rule tends to form larger, more robust, often copiously fertile, thalli than P. crocata. Like this
latter species a small proportion of P. neglecta have stictic acid replaced by salazinic acid (see
Galloway etal., 19836).

Distribution: (Fig. 50) Widespread and common throughout, from the Three Kings Is (lat.
356'S) to Campbell I., coastal and inland, sea-level to 1950 m.

Habitat ecology: Pseudocyphellaria crocata is a common species of open grasslands and heaths
where it is found on rocks or on decorticated wood. It is known from maritime rocks (especially
from the offshore islands north of Auckland), from basalt, lava, serpentine, clay banks, and
stone walls in drier parts of the country . It is not seen in areas of high rainfall or high cloud cover,
preferring drier, open habitats exposed to full sunlight. Although a common coastal species it
ascends into high-alpine fellfield on the Remarkables (1950 m), though it is rare or absent from
subalpine localities west of the Main Divide. As an epiphyte of trees and shrubs it is most
commonly collected from Cordyline australis or especially Leptospermum. Besides these it is
known from the following phorophytes: Coprosma spp, D aery carpus dacrydioides, Dracophyl-
lum subulatum, Metrosideros excelsa, Populus, Nothofagus solandri var. cliffortioides , Prum-
nopitys ferruginea, Pseudopanax crassifolius , Salix, and Weinmannia racemosa. In Central
Otago it is common on schist rocks with Parmelia signifera, P. sulcata, and species of Umbilicaria
and Xanthoparmelia. Elsewhere in its range it associates with Everniastrum soroch'eilum,
species ofMenegazzia, Psoroma, and Usnea, Sticta fuliginosa and species of Pseudocyphellaria,
notably P. colensoi, P. coronata, P. dissimilis, P. granulata, P. neglecta, P. pickeringii, and P.
poculifera.

Specimens examined: 450.

16. Pseudocyphellaria degelii D. Galloway & P. James Figs 51, 52

in Lichenologist 15: 141 (1983). Type: New Zealand, South I., Nelson, Mt Haast, on Nothofagus
solandri var. cliffortioides at tree-line, 28 December 1978, D. J. Galloway (CHR 381020! - holotype,
BM!-isotype).

Morphology: Thallus orbicular to irregularly spreading, loosely attached, margins free and
ascending to unattached, corticolous, musciolous or terricolous, rarely saxicolous. Lobes
variable, broadly rounded to elongate-laciniate, 0-5-1 -5(-3) cm wide and l-3(-5) cm long,
plane to canaliculate (especially in grassland forms), discrete to complex-imbricate. Margins
entire, sinuous, shining, thickened or slightly inflated, with scattered, punctate to sublinear,
yellow pseudocyphellae, slightly notched in parts, sinuses deeply to shallowly rounded. Upper
surface bright emerald green, glaucous green or yellow-green occasionally suffused brownish at
apices when moist, pale yellow-green or yellow-grey when dry, becoming tawny or ochre-brown
on storage, undulate, weakly to strongly wrinkled, pitted, scroliculate or faveolate, intercon-
necting ridges low, rounded, texture rather thin and papery (grassland forms) to thick,
coriaceous (bark forms), matt, occasionally minutely scabrid-areolate (use x 10 lens) at lobe
apices, occasionally minutely maculate (xlO lens), without soredia, isidia, phyllidia or
pseudocyphellae. Medulla yellow. Photobiont green. Lower surface irregularly wrinkled or
puckered, undulate to shallowly bullate, pale yellow to ochre brownish at margins, dark
red-brown to blackened centrally, tomentose to margins or with a narrow to broad, glabrous,
marginal zone, tomentum often sparse and patchy, ragged to dense and woolly, pale whitish
grey to brownish, silky. Pseudocyphellae yellow, scattered, minute, often somewhat inapparent,
0-05-0-15 mm diam., punctiform to minutely papillate, decorticate area flat to subconvex.
Apothecia sessile, constricted at base, becoming pedicellate at maturity, pedicel smooth,
concolorous with thallus, sparse, often absent from well-developed thalli, (0-5-)l-5(-7) mm

PSEUDOCYPHELLARIA

119

Fig. 51 Pseudocyphellaria degelli. Holotype (CHR) A. Scale = 1 cm. B. Scale = 5 mm.

diam., rounded to irregular, concave to plane or undulate or irregularly flaring, disc pale to
dark red-brown, shining or matt, epruinose, exciple coarsely corrugate-scabrid, areolae deeply
delimited, pyramidal, whitish buff to red-brown, translucent when wet.

Anatomy: Thallus 130-200(-250) um thick. Upper cortex 22-27 um think, upper half dilute
yellow-brown to orange-brown, remainder colourless, uppermost layers flattened-compressed,
appearing periclinal, innermost cells thick-walled, 4-6 urn diam. Photobiont layer 18-30 um
thick, cells densely packed, photobiont green, cells to 5 um diam. Medulla 90-140(-170) um
thick, hyphae to 4-5 um diam., encrusted with yellow crystals especially near photobiont zone
and near lower cortex. Lower cortex ll-18(-22) um thick, pale straw-yellow, cells 5-9 um diam.
Tomental hairs septate, constricted at septa, pale straw-yellow, to 7 um diam. , 30-120 um long.
Apothecia: Exciple 130-200 um thick, cells 6-8-15-5 um diam., separating in rows near margins
as short tomental hairs, or into irregular pyramidal clumps (scabrosity). Hypothecium 64-83 um
thick, pale orange-brown, opaque, unchanged in K. Thecium 100-110 um tall, colourless;

120

D. J. GALLOWAY

epithecium 7-11-5 urn thick, yellow-brown to pale red-brown, granular, colour dissolving in K.
Asci 85-110 x 15-18 urn. Ascospores colourless, fusiform, 1-3-septate, apices pointed, straight
or curved, contents of cells often granular-vacuolate, (27-)29-5-36(-40) x (4-5-7-9 um.

Chemistry: Pulvinic acid, pulvinic dilactone, calycin, 22a-hydroxy-3, 4,-secostict-4(23)-ene-3-
oic acid, 22a-hydroxy-3, 4-secostict-4(23)-en-3-ol, and 3-acetoxy-3, 4-secostict-4(23)-en-22a-ol
(Goh etal., 1978).

Distinguishing features: Pseudocyphellaria degelii is a large, endemic species having broadly
rounded to linear-elongate lobes with mainly entire margins, lacking isidia, soredia or phyllidia.
The upper surface is undulate to scrobiculate or strongly reticulate-faveolate, occasionally
maculate and scabrid-areolate (x lO lens). It has a yellow medulla, a green photobiont, and a
pale yellow to red-brown or black lower surface with a continuous to patchy tomentum.
Apothecia sparse, often absent, sessile to pedicellate, exciple well-developed, coarsely
corrugate-scabrid. Spores colourless, fusiform, 1-3-septate. The medulla contains pigments and
seco-stictane triterpenoids (Goh etal., 1978).

Variation: Pseudocyphellaria degelii has two main growth forms. A corticolous form from near
tree-line or at forest margins, has thicker, narrower lobes which are normally wrinkled-
scrobiculate to faveolate and which are sparsely fertile (Fig. 51). Subalpine grassland forms, on
the other hand, have thinner, broader, and more papery lobes (Fig. 52) which are undulate,
smooth, and never or rarely scrobiculate or faveolate, and very rarely fertile. This latter form
was recorded in earlier accounts as Sticta durvillei (Zahlbruckner, 1941; Allan, 1949), as Sticta

Fig. 52 Pseudocyphellaria degelii. Mt Arthur, Nelson, Bartlett (CHR 375875). Scale = 1 cm.

PSEUDOCYPHELLARIA

121

Norfolk I

Kermadec 30-

Is

Lord Howe I

Three Kings Is

The Snares

Auckland Is
\

Campbell I

Macquarie I

160

170

40 c

-42

-44

-46

Bounty Is
Antipodes Is

50'

180

172 C

174

176

178

166

168

170 C

172 C

174 C

176

34-

42 o

-44

Fig. 53 Distribution of Pseudocyphellaria degelii.

122 D. J. GALLOWAY

endochrysa (Martin, 1966, 1969; Martin & Child, 1972), and as P. berberina (Galloway &
James, 1980).

Pseudocyphellaria degelii is readily distinguished from both P. colensoi and P. coronata on
morphological and chemical grounds; it lacks the lacerate margins and marginal and laminal
verrucose isidia of P. colensoi which has also a yellow acetone extract, and it does not have the
broadly rounded lobes of P. coronata which are glabrous below, or the semi-immersed to
sessile and coronate-dentate margined apothecia, or the distinctive red-magenta acetone extract
of this species. It appears to be the South Pacific vicariant of the South American taxon P.
berberina (Galloway & James, 1977), but differs from it in several respects. It is distinctive
chemically (Wilkins, 1977 'b; Goh et al., 1978), has narrower lobes which are not as broadly
rounded, margins are entire or notched but not conspicuously indented, folded, incised or
phyllidiate as in P. berberina. In P. degelii the upper surface is smoother, less deeply faveolate
and more rarely scabrid-areolate, and the exciple is also much more massively developed even at
an early stage, and is more coarsely areolate-scabrid. Spores in P. degelii, (27-)29-5-36(-40) x
(4-5-)7-9 urn, are shorter and broader than those of P. berberina, (20-)30-50(-55) x
(5-)6-7(-8) urn.

Pseudocyphellaria degelii is also distinct from the South American subalpine, subantarctic
species P. endochrysa and P. vaccina (see Galloway, 1986a). P. endochrysa has a finely
pubescent upper surface, broad, papery lobes with linear, marginal pseudocyphellae below, and
P. vaccina has a distinctive scabrid-areolate upper surface with thickened, inflexed margins
without linear marginal pseudocyphellae below. Both these latter species have colourless,
acicular spores c. 20-32 x 2-5-4 urn.

Distribution: (Fig. 53) From lat. 3740'S in North I., (Raukumara Range, Huiarau Range,
Kaimanawa Range, Tararua Range), and from Nelson to Fiordland in South I., close to the
Main Divide and eastwards on foothill ranges of Marlborough, Canterbury, and Otago to Silver
Peaks and Mt Maungatua near Dunedin, the Blue Mountains, Forest Hill, and Waihopai Scenic
Reserve (Invercargill). Also on Stewart I. (Mt Anglem) and Campbell I. (Mt Azimuth, 395 m),
rarely at sea-level, most often from 450-1800 m.

Habitat ecology: Pseudocyphellaria degelii occurs in canopy branches and on trunks of
Nothofagus solandri var. cliffortioides and Podocarpus hallii, and is most noticeable at, or near,
treeline. In subalpine grassland dominated by species of Chionochloa, P. degelii is found at the
bases of low shrubs (species of Cassinia, Coprosma, Dracophyllum, and Hebe) and around
tussock bases, and among alpine herbs, predominantly species otAciphylla and Celmisia. It is
also known from damp soil among rocks or boulders in fellfield. In subalpine grassland habitats
and in the higher, more restricted alpine tundra environments (see Mark & Bliss 1970; Bliss &
Mark, 1974) , P. degelii associates with the following lichens: Alectoria nigricans, Cetraria delis ei,
C. islandica ssp. antarctica, Coelocaulon aculeatum, Haematomma babingtonii, Hypogymnia
lugubris, Lecanora broccha, L. flavopallida, Menegazzia aeneofusca, M. castanea, M. inflata,
M. nothofagi, M. testacea, Pseudocyphellaria maculata, Psoroma hirsutulum, Siphula decum-
bens, S. foliacea, S. fragilis, Sphaerophorus tener, Sticta martinii, Thamnolia vermicularis , and
Usnea contexta.

Corticolous associates of P. degelii include: Degelia duplomarginata, Parmelia testacea,
Lecanora atra, Haematomma Mare, Pseudocyphellaria ardesiaca, P. faveolata, P. glabra, P.
homoeophylla, Menegazzia pertransita, M. dielsii, Usnea capillacea, and U. xanthophana.

Specimens examined: 80.

17. Pseudocyphellaria dissi mills (Nyl.) D. Galloway & P. James Fig. 54

in Lichenologist 12: 297 (1980). - Stictina fragillima var. dissimilis Nyl., Syn. meth. lich. 1 (2): 336
(1860). - Stictina dissimilis (Nyl.) Nyl. in Lindsay in Trans. Linn. Soc. Land. 25: 506 (1866). - Sticta
fragillima var. dissimilis (Nyl.) Krempelh., Reise Oest. Freg. Novara Bot. 1: 119 (1870). - Cyanisticta
dissimilis (Nyl.) Rasanen in/. Jap. Bot. 16: 143 (1940). Type: Australia, sine loco, Hampe (H-NYL
34103! - lectotype) [see note 1].

PSEUDOCYPHELLARIA

123

HERBARIUM
LOJKANTJM

&:,/ u^

nrp*. ifm <f*?n.v-t*~rn. -

Fig. 54 Pseudocyphellaria dissimilis. New Zealand, Helms [as Sticta filix] (W). Scale = 2 cm.

Stictina fragillima var. dissecta Mull. Arg. in Flora, Jena 66: 22 (1883). - Sticta fragillima var. dissecta

(Mull. Arg.) Hellbom in fl//i. K. svercsfcfl Vetensfc. -/Uad. /fand/. 21(3/13): 32 (1896). Type: Norfolk

Island, Dr Woolls (G! - lectotype).
Sricta intricata var. fimbriata Zahlbr. in Denkschr. Akad. Wiss. Wien math.-naturwiss. Kl. 104: 290

(1941). Type: New Zealand, Wellington, Totara Reserve, Pohangina River, on Podocarpus totara, 31

January 1935, H. H. Allan v 186 (W! - lectotype).

Note 1: Stictina fragillima var. dissimilis Nyl.

In proposing a variety (dissimilis) of Stictina fragillima, Nylander (1860ft 336) wrote '-Varietatem hujus (ni
fallor), in hb. Hampeano vidi, laciniis thallinis supra medio canaliculato-impressis et margine hinc inde
minute laciniato-dissectis; ex Australia Felici et Tasmania. Dicatur var. dissimilis. Plerumque lacinias
thallinas typo breviores latiores et magis irregulares habet; margines saepe adscendentes'.

The lectotype, H-NYL 34103, chosen by Galloway & James (1980: 297) is only a fragment, as are many
of Nylander's specimens retained from larger collections sent to him for determination. The remainder of
the Hampe collection has not been located. The type, which is sufficient for identification, is a single sterile
lobe (3-5 x 0-5 cm) with small, terete isidia developed at the margins. It is attached to a sheet annotated by
Nylander' Sticta fragillima var. Austr. fel. Hb. Hampe'. The packet containing the specimen is labelled by
Nylander 'Stictina fragillima Nyl. f . dissimilis Nyl. Australia'.

Morphology: Thallus very variable, orbicular to irregularly spreading, 5-12(-25) cm diam.,
loosely to closely attached centrally, margins free often ascending, terricolous, saxicolous or
corticolous. Lobes very variable, linear-elongate to shallowly rounded, (l-)3-7(-12) mm
wide, (l-)2-5(-10) cm long, branching subdichotomous to irregular, discrete at apices,
discrete or contiguous from margins to centre, to complex-imbricate centrally, often
canaliculate, sometimes divided into numerous, strap-like phyllidia. Margins entire, sinuous,
smoothly rounded or ridged above, distinctly thickened below, sinuses rounded to acute,
prominent, becoming isidiate and/or phyllidiate, without soredia or pseudocyphellae. Upper
surface dark leaden-grey to blue-black at margins, paler centrally, with minute, white, marbled
maculae when moist (x 10 lens), pale grey-blue, or glaucous-grey when dry, becoming cinna-

124 D. J. GALLOWAY

mon-brown on storage and often staining herbarium paper orange-red or brown, smooth or
shallowly wrinkled, often with minute, pale buffer whitish, scattered to crowded papillae (x 10
lens), glossy to matt. Isidia marginal then laminal, simple, terete, 0-5-2-5(-3-5) mm tall, 0-1 mm
diam., becoming densely crowded, coralloid branched and eventually flattened and phyllidi-
ate. Phyllidia mainly marginal, constricted at base, simple, strap-like to palmate-squamiform.
Maculae laminal, minute, marbled, scattered, irregular, pale buff or brown photobiont-free
zones often present as often extensive blotches or patches. Medulla white. Photobiont Nostoc.
Lower surface white to pale buff at margins, brownish centrally, glabrous, shining at margins,
distinctly costate and tomentose centrally, tomentum short, velvety, often rather sparse, white,
glistening. Pseudocyphellae white, widely scattered to numerous, flat, fleck-like, rather indis-
tinct at margins, 0-01-0- 1 mm diam. , rarely to 0-3 mm diam. , centrally, round to oval or sigmoid,
decorticate area convex or plane, a thin margin often apparent, level with, or slightly projecting
above, tomentum. Apothecia occasional to rare or absent, scattered, solitary to crowded (2-9
together), rounded to irregular-contorted through mutual pressure, marginal and laminal,
sessile, constricted at base, 0-1-2-5 mm diam., shallowly concave to plane or undulate-convex,
disc pale yellow-brown to dark red-brown, smooth to minutely granular, occasionally with
central plugs of sterile tissue, epruinose, exciple pale whitish-buff to flesh-coloured, translu-
cent when wet, scabrid-aereolate, crenulate at margins of disc.

Anatomy: Thallus 170-250 urn thick. Upper cortex 39-47 um thick, pale straw-yellow, upper
cells compressed, inner cells more loosely arranged, 6-5-18 um diam. Photobiont layer 27-45
um thick photobiont Nostoc. Medulla 73-110 um, colourless, hyphae 5 um diam. Lower cortex
27-34 um thick, outermost row of cells yellow-brown to red-brown, cells 9-12 urn diam., inner
rows of cells larger and thinner-walled, 11-15(-20) um diam., more loosely arranged. Tomental
hairs yellow-brown, slightly constricted at septa, 4-5-7 um diam., 34-112 um long, scattered,
single. Apothecia: Exciple cellular, cells thick-walled, colourless to pale straw-yellow, 8-27 um
diam. Hypothecium 55-62 um thick, pale yellow-brown to pale straw-yellow, not changing in K.
Thecium colourless, 90-102 um tall; epithecium yellow-brown to pale red-brown, unchanged
in K, to 12 um thick. Asci 77-95 x 11-15 um. Ascospores pale yellow-brown, fusiform-
ellipsoid, 1-3-septate, straight or slightly curved, apices rounded or pointed, 20-5-29-5 x
7-9 um.

Chemistry: gyrophoric acid (tr.), 7|3-acetoxyhopan-22-ol, hopane-7(3 22-diol (tr.), hopane-
15a,22-diol.

Distinguishing features: Pseudocyphellaria dissimilis is a highly variable australasian or,
possibly palaeotropical, species having linear-elongate to shallowly rounded, subdichotomously
to irregularly branched lobes, often canaliculate and with entire margins becoming isidiate or
occasionally also phyllidiate. The upper surface is smooth or shallowly wrinkled, glossy or matt,
minutely maculate and papillate (xlO lens), and with marginal and laminal isidia, and/or
phyllidia (Fig. 55) Isidia are simple, terete at first and later may become coralloid-branched,
or flattened and phyllidiate, though simple, terete isidia are most common. It has a white
medulla, a blue-green photobiont, and a pale to brownish often costate lower surface with
rather sparse, short, central tomentum and scattered, white, fleck-like pseudocyphellae.
Apothecia are rare, sessile, with a pale, scabrid-areolate exciple. Spores are pale yellow-brown,
1-3-septate. It has a two-hopane chemistry [Code A of Wilkins & James (1979)], gyrophoric
acid.

Variation: Pseudocyphellaria dissimilis is an extremely variable species having a highly plastic
morphology, with lobes varying from narrow, strap-like with only a few marginal isidia and
only 1-2-branched, to highly divided, imbricate, densely isidiate or phyllidiate structures. Isidia
in P. dissimilis in the majority of cases are terete but occasionally terete isidia and flattened
dorsiventral phyllidia occur on the same plant, which suggests a rather broad spectrum of
variation in these structures in this species, possibly related to specific microclimate or
microhabitat conditions. The diversity in lobe morphology parallels that seen in P. multifida and
it seems likely that this latter taxon is the green-photobiont counterpart species of P. dissimilis.

PSEUDOCYPHELLARIA

125

Fig. 55 Pseudocyphellaria dissimilis. New Zealand, Helms (W). A. Showing apothecia. B. Showing
profuse development of phyllidia. Scale = 5 mm.

Photosymbiodemes of the two species are known from near the Lewis Pass in South Island, but
are extremely rare.

Pseudocyphellaria dissimilis is distinct from P. cinnamomea, which is closely related, differing
in the narrow, straplike, strongly canaliculate lobes with entire margins which are never
isidiate or phyllidiate, and its lower surface is always distinctly costate-ridged. Pseudo-
cyphellaria fimbriatoides is also similar to P. dissimilis but its phyllidia are noticeably pubescent
and its lobes are broader, undulate, and shining, resembling Peltigera. An Australian-?
palaeotropical species P. insculpta (not known from New Zealand) resembles P. dissimilis but
has a distinctive punctate-impressed upper surface (reminiscent of P. haywardiorum, P.
junghuhniana, P. rigida, and P. sulphured) and very narrow sublinear lobes which are very richly
divided and may even be phyllidiate but which are never furnished with terete isidia.

Dried specimens of P. dissimilis (and also P. cinnamomea) stain herbarium paper reddish
brown after long storage.

126 D. J. GALLOWAY

Distribution: (Fig. 56) From North Auckland (lat. 35S) to Wellington in North I., and from
Nelson to Fiordland and eastwards from Marlborough to Invercargill in South I. Also in Stewart
I. , from Mt Anglem to Port Pegasus, mainly lowland, both coastal and inland, occasionally to
subalpine, sea-level to 700 m.

Habitat ecology: Pseudocyphellaria dissimilis is tolerant of low light intensities and is the most
common (often the sole) species of the genus to be found in deeply shaded habitats in New
Zealand. Very often it is the only major lichen epiphyte in dark forest interiors, especially in
North I. forests where it is very common, often in association with Clathroporina exocha and
Metus conglomerates. It grows readily on damp soil, amongst mosses or on rocks on the forest
floor, on tree roots, and on tree trunks in moderate to dense shade, but is rare on twigs in these
habitats. It prefers damp, cool, humid habitats and in these habitats it is an efficient fixer or
atmospheric nitrogen (Green et al., 1980). As such, it makes a significant contribution to the
nitrogen balance of northern rain forests.

It is known from the following phorophytes: Agathis australis, Ascarina lucida, Beilschmiedia
taraire, B. tawa, Carpodetus serratus, Cordyline australis, Cyathodes, Dacrycarpus dacry-
dioides, Dacrydium cupressinum, D. intermedium, Dysoxylum spectabile, Griselinia littoralis,
Knightia excelsa, Melicytus ramiflorus, Metrosideros excelsa, M. lucida, Nothofagus fusca,
Podocarpus hallii, P. totara, Prumnopitys ferruginea, Pseudopanax anomalus, P. arboreus,
Pseudowintera colorata, Rhipogonum scandens, Rhopalostylis sapida, Sphaeropteris medullar-
is, and Weinmannia racemosa.

Pseudocyphellaria dissimilis associates with the following lichens: Calycidium cuneatum,
Clathroporina exocha, Letogium azureum, L. cyanescens, Lobaria adscripta, L. retigera,
L. scrobiculata, Metus conglomerates, Nephroma cellulosum, N. plumbeum, N. helveticum,
Normandina pulchella, Pannaria crenulata, P. immixta, Parmeliella nigrocincta, Peltigera
dolichorhiza, Polychidium contortum, Pseudocyphellaria billardierei, P. colensoi, P. faveolata,
P. glabra, P. fimbriatoides , P. homoeophylla, P. hooked, P. multifida, P. murrayii, P.
rufovirescens , Psoroma sphinctrinum, Sphaerophorus insignis, S. notatus, S. tener, Sticta
caliginosa, S. filix, SI. lacera, and 5. subcaperata.

Specimens examined: 205.

18. Pseudocyphellaria durietzii D. Galloway Fig. 57

in Lichenologist 15: 142 (1983). Type: New Zealand, Stewart I., Lower Kopeka River, on Coprosma
foetidissima roots on rata, in riverine rata-kamahi-totara-miro forest, c. 10 m, 12 February 1980,
C. D. Meurk (CHR 375960! - holotype, BM! - isotype).

Morphology: Thallus rosette-forming to irregularly spreading, 10-15 cm diam., adnate cen-
trally, rather loosely attached at margins, corticolous. Lobes subdichotomously to irregularly
branching, 0-5-1-5 cm wide, 2-6 cm long, discrete or subimbricate at margins, often discrete
or contiguous from margins to centre. Margins entire, occasionally slightly notched, shallowly
sinuous, without projecting pseudocyphellae, not noticeably thickened below. Upper surface
olive or yellowish green to fresh green when wet, pale olivaceous or grey-buff when dry, strongly
reticulate-faveolate, ridges prominent, to 1 mm wide, smoothly rounded, furnished with
minute, punctiform, red-brown pycnidia, faveolae prominent, deeply lacunose, 2-5 mm wide;
smooth to very faintly wrinkled or pitted, matt or slightly shining, soredia, isidia, maculae,
phyllidia, and pseudocyphellae absent. Medulla white. Photobiont green. Lower surface pale
yellowish buff or whitish at margins, yellow-brown centrally, strongly wrinkled-bullate,
glabrous in patches or with scattered tomentum, tomentum very thin, patchy, pale buff or
yellow-brown. Pseudocyphellae common, scattered, minute, papillate, hemispherical, 0-1
mm diam. or less, decorticate area white, flat, minute, 0-03 mm diam. Apothecia marginal or
submarginal, never laminal, 1-5 (-8) mm diam., sessile to subpedicellate, pedicel best seen in
young fruits, 0-5-1-5 mm wide, concave, margins inrolling and often obscuring disc, often also
lacerate-striate or scalloped, without phyllidia, disc matt or shining to minutely granular, dark
red-brown to black, epruinose, imperf orate, exciple concolorous with thallus or suffused

PSEUDOCYPHELLARIA

127

Fig. 56 Distribution of Pseudocyphellaria dissimilis.

128

D. J. GALLOWAY

Fig. 57 Pseudocyphellaria durietzii. Holotype (CHR). Scale = 1 cm.

brownish at margins, smooth and shining at margins, minutely areolate to delicately
pilose-scabrid towards pedicel, smooth or occasionally coarsely rugose-plicate towards base.
Pycnidia common, mainly on reticulate ridges of upper surface, minute, papillate, apical ostiole
red-brown to black. 0-2 mm diam., or less.

Anatomy: Thallus 110-150(-200) um thick (360-600 urn thick at ridges). Upper cortex 18-23
(-27) urn thick, outer 7-9 um, pale orange- or yellow-brown, cells stretched and compressed,
inner 11-14 um colourless, cells to 5 um diam. Photobiont layer 11-22 um thick, photobiont
yellow-green, cells round to irregular, 3-5 um diam. Medulla 27-110(-500 at ridges) um thick,
hyphae loosely interwoven, encrusted with granular crystals. Lower cortex 14-22 um thick,
colourless to pale straw, cells (3-5 rows), 4-5 um diam. Tomental hairs sparse, pale straw,
simple, septate, to 5 um thick and 20-90(-140) um long, in fascicles. Pycnidia hemispherical
to oval, 170-210 um diam. Cephalodia forming in medulla below photobiont layer and bursting
through lower cortex, ovoid, 135-165 x 90 um, Nostoc in a layer (35-45 um thick) in uppermost
part of structure. Apothecia: Exciple containing photobiont layer and medulla as in thallus,
35-45(-60) um thick, cells pale straw, walls 2-3 um thick, lumina very small, 2 um or less, tissue
rather irregular and ragged below, splitting off into irregular, pyramidal groups of cells.
Hypothecium 35-45(-50) um thick, dilute orange-brown, granular, opaque. Thecium colourless
to pale straw, 70-90 um tall; epithecium 7-11 um thick, smoky grey-brown, turning vinous-
purple in K, minutely granular. Asci 60-80 x 18-21 \nm.Ascospores oval-ellipsoid with pointed
ends, straight or slightly curved, brown, thickened 1 -septate, septum variable 2-4-5 um thick,
narrower at maturity, 23-25 x 9-11 um.

Chemistry: Methyl evernate, tenuiorin, methyl lecanorate, evernic (tr.) and gyrophoric acids,
and four unidentified depsides, hopane-7|3, 22-diol (tr.), hopane-6ct, 7(3, 22-triol, 7(3-
acetoxyphopane-aa, 22-diol (tr.), 6a-acetoxyhopane-7|3, 22-diol, norstictic (tr.), stictic,
hypostictic, cryptostictic, and constictic acids.

Pseudocyphellaria durietzii is named for G. Einar Du Rietz, formerly Professor of Ecological
Botany and Director of the Vaxtbiologiska Institutionen, Uppsala University, who, with his first

PSEUDOCYPHELLARIA

129

Norfolk I

Lord Howe I

Three Kings Is

172

174

176

178

-40

-42

166
_!

168

170

I

172

40-

42-

-44-

100

176

Fig. 58 Distribution of Pseudocyphellaria durietzii.

130 D. J. GALLOWAY

wife Greta Sernander-Du Rietz, first collected the species from rimu (Dacrydium cupressinum)
bark in mixed rain forest on Seymour I. , Doubtful Sound, Fiordland in 1927.

Distinguishing features: Pseudocyphellaria durietzii is an endemic species having dichoto-
mously branching, linear-elongate to somewhat imbricate lobes with entire to crenulate
margins. The upper surface is deeply and regularly reticulate-faveolate, without soredia, isidia,
phyllidia, or pseudocyphellae. It has a white medulla, a green photobiont, and a pale buff,
wrinkled-bullate, glabrous to scattered-tomentose lower surface. Apothecia marginal, sub-
pedicellate, disc dark red-brown to black, exciple concolorous with thallus. Epithecium
granular, smoky grey-brown, becoming vinous-purple in K. Spores brown, thickened 1-septate.
It has a complex chemistry containing gyrophoric acid (cortex C+ red, fading fast) and other
depsidones, hopane - 6a, 7(3, 22-triol and metabolites of the stictic acid complex.

Variation: Pseudocyphellaria durietzii shows little morphological variation throughout its
known range. It is related to P. hooker i (the two species have the same chemistry and epithecial
colour reactions) but this species has a blue-green photobiont and a dark brown lower surface. In
the Urewera National Park near the northern part of the range of P. durietzii, photo-
symbiodemes of two species are found. It is similar to P. physciospora, but this species has
yellow pseudocyphelne and a different chemistry. See also Table 2 (p. 216).

Distribution: (Fig. 58) In North I. , from Mt Hikurangi (lat. 3755'S) and the Urewera National
Park to Wellington, and from Nelson to Fiordland and Mt Peel (South Canterbury) in South I.
Also in Stewart I., from the north coast to Port Pegasus and in Campbell I., mainly lowland to
subalpine, sea level to 800 m.

Habitat ecology: Pseudocyphellaria durietzii is a species of lowland forest and scrub , favouring
habitats of moderate shade, high humidity, and high rainfall. It is found in subalpine scrub on Mt
Hikurangi, but in the Urewera National Park and on Stewart I. , the two areas where it has until
now been most commonly collected (scattered occurrences on the western mountains of North
I., and from Nelson to Fiordland connect these two major population centres), it is found in
open areas of lowland forest and/or scrub. It is an epiphyte of both bark and twigs of the
following phorophytes: Coprosma rhamnoides, D aery carpus dacrydioides , Dacrydium cupres-
sinum, Ixerba brexioides, Metrosideros umbellata, Podocarpus totara, Prumnopitysferruginea,
and Weinmannia racemosa.

Specimens examined: 30.

19. Pseudocyphellaria episticta(Nyl.) Vainio Fig. 59

in Philipp. J. Sci. C, 8: 117 (1913). - Sticta episticta Nyl. in Flora, Jena 48: 299 (1865). - Lobaria
episticta (Nyl.) Trevisan, Lichenotheca Veneta exs. 75 (1869). Type: New Zealand, Otago, on trees,
top of Saddlehill near Dunedin, 26 October 1861, W. L. Lindsay (H-NYL 33433! - holotype; E! -
isotype).

The citation of the basionym (Sticta episticta) as Nylander, 1866: 248, in Galloway & James (1980: 298), and
Galloway (19856: 440) is incorrect. Even though Nylander (1866) gives a good description of the taxon in
his discussion of Lindsay's Otago lichens, a valid description of the species appeared the previous year as
shown above.

Morphology: Thallus rosette-forming to irregularly spreading, 5-10(-20) cm diam., closely
attached centrally and to margins, apices slightly free and subascendent, corticolous, rarely
saxicolous. Lobes linear-elongate, deeply indented, rather ragged, 0-5-1-5 cm diam., 2-5 cm
long, apices and central parts complex-imbricate, folded. Margins lacerate-indented, ragged,
rarely entire and shallowly rounded, usually sinuous and ascending, crisped, crenate to
densely coralloid-phyllidiate, very slightly thickened below. Upper surface bright lettuce-green
when moist, pale grey-green to fawnish or glaucous-yellow when dry, undulate, convex to
subcanaliculate or plane, minutely wrinkled and/or pitted, not faveolate, matt or shining, rather
coriaceous, without soredia, isidia or maculae, pseudocyphellate. Pseudocyphellae laminal,

PSEUDOCYPHELLARIA

131

Fig. 59 Pseudocyphellaria episticta. Isotype oiSticta episticta (E). A. Scale = 1 cm. B. Scale = 2 mm.

scattered, white, punctiform, fleck-like to occasionally linear or effigurate, 0-05 mm diam. or
less, rarely in older parts, regenerating phyllidia at margins. Phyllidia mainly marginal, simple at
first and constricted at base, flattened, becoming coralloid-branched, 0-05-0-5 mm tall, often
clustered, imbricate or forming extensive crusts, very occasionally on lamina and then rather
scattered and irregular in appearance, not, or rarely, associated with pseudocyphellae. Medulla

132 D. J. GALLOWAY

white. Photobiont green. Lower surface whitish to pale buff, darker centrally, tomentose from
margins to centre or occasionally with a narrow, glabrous, slightly wrinkled, shining marginal
zone, tomentum short, even, velvety to arachnoid-woolly, pale whitish-buff, glistening, to
brownish or grey-brown. Pseudocyphellae white, sparse, occasional to rare, sunk in tomentum,
or scattered, fleck-like at or near margins, decorticate area flat, 0-05-0-2 mm diam. Apothecia
sessile, constricted at base, sparse to common (in favourable sites), solitary to crowded,
laminal, 0-5-3 mm diam., round to irregular, constricted through mutual pressure, shallowly
concave at first, plane, undulate at maturity, disc pale to dark red-brown, matt, minutely
granular, epruinose, exciple pale whitish buff, translucent when moist, corrugate-scabrid,
becoming irregularly phyllidiate in older fruits, margins often also excluded in older fruits,
disc completely obscured by exciple in young, emergent fruits.

Anatomy: Thallus 230-275(-320) um thick. Upper cortex 55-68 um thick, uppermost 7-1 1 urn
brown or yellow-brown, somewhat amorphous, remainder colourless to pale straw-yellow, cells
7-15 urn diam. Photobiont layer 34-45 um thick, photobiont cells green, densely packed, 4-5-7
urn diam. Medulla 100-135(-220) um thick, colourless, hyphae 3-4 um diam. Lower cortex
36-45 um thick, pale straw-yellow to yellow-brown, cells 4-5-15 um diam. Tomental hairs
colourless, simple, 5 um thick, 60-110(-180) um long, solitary or in small fascicles. Apothecia:
Exciple colourless, 90 um thick at margins, to 140 um thick at base, outer surface splitting away
into ragged clumps of cells (scabrosity), cells thick-walled, 4-5-13-5 um diam. Hypothecium
dilute yellow-brown, opaque, 45-64 um thick. Thecium 90-100 um tall, colourless to pale
straw-yellow; epithecium red-brown, 18-22 um thick, turning rose-pink in K and slowly
dissolving; paraphyses strongly conglutinate, apices swollen, not moniliform. Asci 85-95 x
14-18 um. Ascspores pale yellow-brown, fusiform, apices pointed, l-3(rarely -7)-septate,
slightly curved, cell contents distinctly vaculate, (27-)32-36-5(-41) x 7-9 [Am.

Chemistry: 15a-acetoxy-22-hydroxyhopan-24-oic acid, 15a-22-dihydroxy-24-hopanoic acid.

Distinguishing features: Pseudocyphellaria episticta is an endemic species having linear-
elongate, deeply indented, rather ragged lobes, with lacerate-crenate, cripsed margins which
are densely coralloid-isidiate. The upper surface is pseudocyphellate, the pseudocyphellae
occasionally associated at their margins with regenerating phyllidia, never with isidia. It has a
white medulla, a green photobiont, and a pale buff or darker lower surface which is tomentose
from margins to centre. Apothecia are sparse to common, the corrugate-scabrid exciple being
irregularly phyllidiate at maturity. The epithecium is red-brown, turning rose-pink and dissolv-
ing in K, the spores pale yellow-brown, l-3(rarely -7)-septate. It has two hopane acids present in
the medulla (Ronaldson & Wilkins, 1978; Galloway etal., 1983ft; Galloway, 1985ft).

Variation: Pseudocyphellaria episticta is a rather uniform species, though individual thalli may
vary from neat rosettes to irregularly spreading colonies. Fertile specimens are reasonably
common and in favourable sites apothecia are often densely crowded. It is most closely related
to P. wilkinsii but has a differing chemistry, does not produce true isidia, and has a distinctive
epithecial colour change in K. It is distinguished from P. chloroleuca by its tomentose lower
surface, the different chemistry (its cortex is C-), and lack of true isidia. P. multifida lacks the
laminal and marginal phyllidia, and the laminal pseudocyphellae characteristic of P. episticta
and also has a different chemistry.

Distribution: (Fig. 60) North Auckland (lat. 36S) to Cook Strait in North I., and spasmodi-
cally in South I., from Westland (Greymouth), Canterbury (Lewis Pass, Banks Peninsula,
Mt Peel), and eastern Otago (Waikouaiti, Green Island, Saddle Hill, Lamb Hill, Flagstaff),
mainly lowland and coastal, sea-level to 800 m.

Habitat ecology: Pseudocyphellaria episticta is found in lowland to montane forest in cool,
moist, humid habitats of moderate illumination close to the forest margins. Mainly collected
from tree trunks (not from twigs) it will also occasionally colonize coastal rocks and also rocks in
tussock grassland. It is known from the following phorophytes: Coprosma spp., Dacrycarpus
dacrydioides , Dysoxylum spectabile, Griselinia littoralis, Knightia excelsa, Leptospermum

PSEUDOCYPHELLARIA

133

Fig. 60 Distribution of Pseudocyphellaria episticta.

134

D. J. GALLOWAY

scoparium, Nothofagus menziesii, Podocarpus hallii, and P. totara. It associates with the
following lichens: Lobaria adscripta, Parmelia tenuirima, Parmelia testacea, Pseudocyphellaria
coriacea. P. pickeringii, P. rufovirescens, Sticta filix, S. latifrons, and 5. subcaperata.

Specimens examined: 70.

20. Pseudocyphellaria faveolata (Delise) Malme Figs 61, 62

in Bih. K. svenska Vetensk.-Akad. Handl. 25(3/6): 23 (1899). - Sticta faveolata Delise in Mem. Soc.
linn. Normandie 2: 102 (1825). - Stictina faveolata (Delise) Nyl., Syn. Meth. Lich. 1(2): 337 (1860). -
Cyanisticta faveolata (Delise) Rasanen in Annls Bot. Soc. zool.-bot. fenn. Vanamo 2(1): 42 (1932).
Type: Straits of Magellan, sine collectoribus nomine (BM! - lectotype) [see Galloway et al. (19836:
140); Galloway & James (1986)].

Sticta cellulifera J. D. Hook. & Taylor in Hook. Lond. J. Bot. 3: 647 (1844). - Crocodia cellulifera (J. D.
Hook. & Taylor) Trevisan, Lichenotheca veneta exs. 75 (1869). - Diphanosticta cellulifera
(J. D. Hook. & Taylor) Clements, Gen. fung.: 175 (1909). - Pseudocyphellaria cellulifera (J. D.
Hook. & Taylor) Gyelnik in Feddes Reprium Spec. nov. veg. 29: 2 (1931). - Sticta foveolata var.
cellulifera (J. D. Hook. & Taylor) Church. Bab. in J. D. Hook., Fl. nov.-zel. 2: 278 (1855). - S.
fossulata var. cellulifera (J. D. Hook. & Taylor) Nyl. , Syn. meth. lich. 1(2): 364 (1860). - 5. billardierei

Fig. 61 Pseudocyphellaria faveolata. Lectotype of Sticta cellulifera (BM). Scale = 2 cm.

PSEUDOCYPHELLARIA

135

Fig. 62 Pseudocyphellariafaveolata. Lectotype of Sticta lorifera (BM). Scale = 2 cm.

var. cellulifera (J. D. Hook. & Taylor) Mull. Arg. in Flora, Jena 71: 135 (1888). - Lobaria billardierei

var. cellulifera (J. D. Hook. & Taylor) Hellbom in Bih. K. svenska Vetensk. -Akad. Handl. 21(3/13):

41 (1896). Type: Campbell Island, /. D. Hooker (BM! - lectotype) [see note 1]. Fig. 61
Sticta lorifera Stirton in Trans. N.Z. Inst. 32: 71 (1900). Type. New Zealand, near Wellington,

/. Buchanan (BM! - lectotype; BM! - isotype). Fig. 62
Sticta elatior Stirton in Trans. N.Z. Inst. 32: 73 (1900). Type: New Zealand, near Wellington,

/. Buchanan (BM! - lectotype).
Sticta condensata Zahlbr. in Denkschr. Akad. Wiss, Wien math.-naturwiss. Kl. 104: 280 (1941). -

Pseudocyphellaria condensata (Zahlbr.) D. Galloway & P. James in Lichenologist 12: 294 (1980).

Type: New Zealand, Southland, Riverton Bush, on rotten log,/. 5. Thomson ZA 541 (W!-holotype;

BM! CHR 317808! -isotypes) [see Galloway, 1986&].

In a preliminary account of the confused nomenclature of this species and its synonyms (Galloway &
James, 1980: 298) several taxa were recorded incorrectly as synonyms and are now placed elsewhere viz.,

136 D. J. GALLOWAY

Sticta fossulata [= P. billardierei]; S. flotowiana [= P. billardierei]; S. linearis [= P. billardierei]; S.
impressa [ P. carpoloma\;S. physciospora [= P. physciospora].

Note 1: Sticta cellulifera J. D. Hook. & Taylor

Thomas Taylor based his name Sticta cellulifera on broadly lobed, deeply faveolate specimens from
Campbell Island, the Auckland Islands, and New Zealand. Material annotated S. cellulifera by Taylor in
the BM is referable to Pseudocyphellariafaveolata, the specimens containing the characteristic marker
compound for this species, physciosporin. Of the three specimens (all collected by J. D. Hooker)
annotated 5. cellulifera by Taylor in his own herbarium (FH), one from Campbell I. is P. faveolata,
one from the Auckland Is is referable to P. billardierei, and the specimen from New Zealand is referable
to P. hookeri. Hooker's interpretation of S. cellulifera (Taylor & Hooker, 1845: 198) records yellow
pseudocyphellae in the material discussed and refers to P. physciospora (see under that species), while
Babington (1855: 278) made 5. cellulifera a variety of 5. foveolata [sic] . Fitch's illustration (plate CXXIV in
Babington, 1855) shows pseudocyphellae strongly reminiscent of those of P. faveolata, and the deeply
faveolate, broadly lobed form of this species.

Morphology: Thallus orbicular to spreading, often forming extensive entangled clones,
loosely attached centrally, margins free, 5-15(25) cm diam., corticolous. Lobes linear-
elongate, very variable in width, 20-40 mm wide at centre, 2-15 mm wide towards apices,
l-5(-10) cm long, discrete or contiguous to subimbricate at margins, complex-imbricate
centrally, broad and somewhat rounded or blunt at apices, with branching, rather irregular or
rather narrow and subdichotomously branching with apices furcate. Margins entire,
shallowly sinuous or scalloped, suffused brownish at apices, slightly to markedly thickened
above, with prominent, raised, conical-verrucose, white pseudocyphellae, rarely with irregular-
ly coralloid isidia or lobules, sometimes ragged-uneven and subascendent. Upper surface
bright lettuce-green to olive-green, suffused red-brown at apices when wet, pale olivaceous,
buff, to dark green or green-black or olivaceous-brown when dry, coriaceous, matt or shining,
rather waxy, strongly reticulate-faveolate, ridges mainly smoothly rounded, 0-1-1 mm wide,
often copiously wrinkled-plicate, often with numerous pycnidia or pycnidial cavities arranged
along ridges, faveolae very variable, shallow to deep, without isidia, maculae, pseudocyphellae
or soredia, occasionally phyllidia develop at margins in response to insect damage. Medulla
white. Photobiont green. Lower surface sometimes with a narrow, glabrous, wrinkled marginal
zone at lobe apices, densely and uniformly tomentose to margins in older lobes, tomentum
felted, brown or black, rarely pale buffer whitish. Pseudocyphellae conical-verruciform, raised
above tomentum or slightly sunken below tomentum, numerous, prominent, 0-1-1 mm diam.,
with a conspicuous, inflated, smooth, fawn or buff to dark brown-black, waxy margin,
decorticate area white, flat or slightly excavate. Apothecia mainly marginal or submarginal,
rarely laminal, most commonly towards lobe apices, l-4(-6) mm diam., shallowly concave at
first, becoming plane to shallowly convex-undulate at maturity, subpedicellate, disc matt to
slightly roughened, pale to dark red-brown when wet, black when dry, white-pruinose when
young, epruinose at maturity, exciple whitish-arachnoid-scabrid at first and then completely
obscuring disc, remaining for a time as a whitish to buff or pale red-brown, ragged-arachnoid
margin, becoming excluded in mature fruits, buff-brown or whitish-arachnoid-scabrid below
disc. Pycnidia occasional to frequent, mainly on reticulate ridges or scattered on floors of
faveolae, minute, punctiform to hemispherical, to 0-5 mm diam., often completely eroded
leaving rows of shallow pits along ridges, ostiole red-brown to black.

Anatomy: Thallus 180-250 (-600 at ridges and pycnidia) um thick. Upper cortex 25-36 um
thick, outer zone (7-11 um) dilute yellow-brown, of necrotic or tightly compressed cells, layer
abutting photobiont zone colourless, cells 4-5-7 um diam. Photobiont layer 35-45(-55) um
thick, photobiont green, cells round to irregular, 4-5-9 urn diam. , chloroplast occupying most of
cell. Medulla 70-150(-400) um thick, colourless, of loosely interwoven hyphae to 4-5 um diam. ,
covered with small, granular crystals. Lower cortex 35 um thick, comprising 4-5 rows of cells,
similar in size and structure to those of upper cortex, outermost layer pale yellow-brown, inner
zone colourless. Tomental hairs 4-5-7 um thick, septate, simple, pale straw to red-brown,
densely conglutinate in fascicles, 90-150(-250) um long. Apothecia: Exciple colourless, cellular,
45 um thick at margins, 90-100 um thick below disc, of parallel, anticlinally arranged hyphae,

PSEUDOCYPHELLARIA 137

cells 4-5-9 x 0-5-2-5 um. Hypothecium 65-90(-112) um thick, dense, opaque, upper parts
orange-brown, lower half olive-brown to dark brown, granular. Thecium hyaline to pale
straw, or dilute yellow-brown, rather granular, 100-120(-135) um tall (lower parts occasionally
greenish in K); epithedum dark grey-brown or olivaceous brown, 11-13-5 um thick, turning
vinous-purple in K. Asci 75-90 x 15-20 urn. Ascospores smoky grey to brown, oval-ellipsoid,
apices pointed, thickened, 1-septate at first then 3-septate, septum 2-5-7 um thick, 25-35 x
9-11 um.

Chemistry: Methyl virensate, physciosporin, hopane-6a, 7|3, 22-triol, stictic, norstictic (tr.),
cryptostictic, and constictic acids [Code B of Wilkins & James (1979)].

Distinguishing features: Pseudocyphellaria faveolata is an austral species having linear-
elongate, narrow to broad lobes, dichotomously to irregularly branched, apices divergent, blunt
or acute, margins entire, thickened, with prominent, raised, verruciform, white pseudocyphel-
lae, rarely also irregularly lobulate or isidiate (response to insect damage?). The upper surface
coriaceous, waxy, reticulate-faveolate, without isidia, maculae, phyllidia, pseudocyphellae or
soredia. It has a white medulla, a green photobiont, and a dark brown to black, often thickly
tomentose lower surface, with scattered white or creamish, verruciform pseudocyphellae.
Apothecia are marginal, subpedicellate, the disc dark red-brown to black, often white-pruinose
when young, exciple whitish or buff-brown, arachnoid-scabrid, obscuring disc at first. Epithe-
cium dark grey-brown, turning vinous-purple in K. Spores grey-brown, thickened 1-septate to
3-septate. It has a complex chemistry containing physciosporin and stictic acid metabolites
[Code B of Wilkins & James (1979)].

Variation: Pseudocyphellaria faveolata is a widespread and very polymorphic lichen. Lowland
forms tend to be narrow-lobed (Fig. 66) and regularly dichotomously branching, whereas
subalpine forms close to tree-line, and in southern Stewart I., and the subantarctic islands
(Auckland and Campbell Is) have broader, less regularly divided lobes (Fig. 65). The tomentum
of the lower surface also varies from a thick, dense, dark matt from margins to centre, to a more
open-textured covering mainly at centre leaving a conspicuous glabrous zone at the margins.
Thalli exposed to full sun or to high light intensities especially at forest margins, or in open scrub
communities, are noticeably darkened, either suffused brownish or even totally blackened.
Shade forms on the other hand are without cortical darkening.

In the northern part of its range, P. faveolata in many habitats appears to be replaced by P.
carpoloma, a superficially similar species but with vivid yellow pseudocyphellae below and a
different chemistry [Code D of Wilkins & James (1979)], and lacking physciosporin. P. faveolata
is distinguished from P. rufovirescens which has entire lobe margins without projecting
pseudocyphellae, a pale, glabrous lower surface, red-brown marginal apothecia which are never
pruinose, epithecial pigments giving no reaction in K, and a different chemistry [Code A of
Wilkins & James (1979)]; and from P. billardierei which also has entire margins without
pseudocyphellae, a sparse dark tomentum on the lower surface, marginal black, epruinose
apothecia, epithecial pigments which turn green-black in K, and a different chemistry deficient
in physciosporin [Code B of Wilkins & James (1979)]. Pseudocyphellaria physciospora,
although similar in many respects to broad-lobed, subantarctic forms of P. faveolata, is
distinguished from it in the entire lobe margins which lack pseudocyphellae, the minute yellow
pseudocyphellae on the lower surface, and the stripitate, epruinose fruits which have a
subdenticulate margin. The chemistry is also distinctive, lacking physciosporin, and containing
tenuiorin, several triterpenoids not present in P. faveolata, plus norstictic and salazinic acids.
See also Table 2 (p. 216).

Distribution: (Fig. 63) Throughout from lat. 35S in North Auckland, though more commonly
collected from lat. 38S southwards in both main islands, also in Stewart I., the Snares, the
Auckland, and Campbell Is; sea-level to 1200 m.

Habitat ecology: Pseudocyphellaria faveolata is a polymorphic, wide-ranging species occupy-
ing many habitat niches from windswept coastal scrub, lowland coastal forest, beech forest,

138

D. J. GALLOWAY

Lord Howe I

Norfolk I

Three Kings Is

Kermadec 30-
Is

The Snares

Auckland Is

Campbell I

Macquarie I

160

40

-42

-44

166

i

170

168

180

170

172 C

174 C

176

172

174

I

176

I

178

34-

36-

100

42'

-44'

Fig. 63 Distribution of Pseudocyphellariafaveolata.

PSEUDOCYPHELLARIA 139

mixed beech-podocarp forest, subalpine scrub close to or above tree-line, to successional
vegetation in altered sites. It is common on both bark and twigs, being especially well-developed
in open forest habitats (close to margins of standing forest) in high-rainfall areas, and is more
commonly collected from southern and western localities (Fig. 67). It is a moderately photo-
philous species and does not tolerate deep shade. It appears to be an obligate epiphyte and is not
known from rocks or soil although it will attach to ferns. It is known from the following
phorophytes: Coprosma propinqua, C. pseudocuneata, Dacrydium cupressinum, Draco-
phyllum filifolium, D. longifolium, Griselinia littoralis, Ixerba brexioides, Kunzea ericoides,
Leptospermum scoparium, Metrosideros robusta, Myrsine divaricata, Myrtus obcordata,
Nothofagus menziesii, N. solandri var. cliffortioides , Olearia colensoi, Pennantia corymbosa,
Phyllodadus trichomanoides , Podocarpus hallii, P. nivalis, P. totara, Pseudopanax simplex,
Pseudowintera colorata, Senecio eleagnifolius , 5. reinoldii, Weinmannia racemosa, and W.
silvicola.

Specimens examined: 180.

21. Pseudocyphellaria fimbriata D. Galloway & P. James Fig. 64

in Lichenologist 15: 142 (1983). Type: New Zealand, South I., Nelson, Lake Rotoiti (Nelson Lakes
National Park), Peninsula, on bark of Nothofagus menziesii, 28 February 1980, D. J. Galloway (CHR
381021! -holotype, BM! -isotype).
Pseudocyphellaria hamata D. Galloway & P. James in NZJl Bot. 16: 521 (1978), nom. nud. (Art. 32.1).

Morphology: Thallus orbicular to irregularly spreading, 6-12(-25) cm diam. , loosely attached
centrally, margins and apices free and ascending, corticolous and occasionally terricolous
(among tussock bases). Lobes irregularly to complexly dissected, elongate-laciniate and
contiguous or discrete from margins to centre, to very richly and complexly divided-imbricate,
branching subdichotomous to very irregular, sinuses well-defined, semicircular, plane
towards apices, becoming canaliculate centrally. Margins entire in parts to richly dissected,
becoming copiously phyllidiate, often ascending, white-tomentose, tomentum short, glisten-
ing, with white, scattered pseudocyphellae or tomentum of lower surface visible. Upper surface
bright lettuce-green to glaucous olive-green, suffused red-brown at apices when moist, pale
greyish yellow to brownish or red-brown when dry, coriaceous, plane or canaliculate centrally
or weakly wrinkled in parts, minutely scabrid-areolate (xlO lens), without soredia, isidia,
maculae or pseudocyphellae. Phyllidia mainly marginal, rarely laminal (and then regenerating
from damaged cortex), fragile, flattened, dorsiventral, rarely simple, furcate to palmately
divided to complex-imbricate to coralloid, 0-5-2-5 mm tall, margins glistening, delicately
white, pubescent, lower surface with minute, white pseudocyphellae. Medulla white. Photo-
biont green. Lower surface whitish or pale buff, in a narrow marginal zone, pale red-brown to
dark-brown or black centrally, tomentose from margins to centre, or with a narrow, glabrous,
marginal zone, tomentum short, velvety at margins, thick, woolly centrally, pale red-brown
marginally, dark-brown to black centrally. Pseudocyphellae white, conspicuous, numerous,
0-05-l(-2) mm diam., round to irregular, minute, flecklike at margins, larger, prominent
centrally, decorticate area brilliant white, shallowly to deeply concave, margins narrow, slightly
raised, inapparent. Pycnidia sparse to frequent, scattered, laminal, minute, punctate-
depressed, ostiole red-brown. Apothecia sessile, constricted at base to subpedicellate, sparse,
when present often towards lobe apices, marginal or submarginal, 1-3 (-4) mm diam., concave
to plane, round to slightly irregular, sinuous, margins pale whitish buff, disc pale to dark
red-brown, smooth, matt or shining, epruinose, exciple pale whitish buff to red-brown,
translucent when moist, coarsely verrucose-areolate, occasionally patchily tomentose, well-
developed in young fruits and then often obscuring disc.

Anatomy: Thallus 180-275(-325) um thick. Upper cortex 45-57(-68) ^im thick, upper 18-27
um pale straw-yellow, cells compressed, flattened, upper surface very uneven-irregular, remain-
ing cells colourless, loosely arranged, 4-5-15(-20) urn diam. Photobiont layer 34-45 (im thick,
photobiont green, cells rounded, 8-13-5 urn diam., chloroplast not filling whole of cell,

140

D. J. GALLOWAY

Fig. 64 Pseudocyphellariafimbriata. Holotype (CHR). A. Scale = 1 cm. B. Scale = 1 mm.

IDictyochloropsis. Medulla 90-140(-185) um thick, hyphae 4-5 ^im diam. Lower cortex 28-34
(j,m thick, outer 7-10 \im dark red-brown, remainder pale straw-yellow, cells thick-walled, 4-5-9
um diam. Tomental hairs colourless to pale red-brown, single or in fascicles 4-5 ^im diam.,
30-140 [im long. Apothecia: Exciple colourless, 110-200 (im thick, in ragged, pyramidal clusters
at margins, cells thick-walled, 4-5-ll(-15) ^im diam. Hypothecium 45-58 (im thick, dilute
yellow-brown, opaque, overlying a compact, colourless (greenish blue on addition of K),
periclinal layer 30-45 um which merges into the medulla proper. Thecium dilute yellow-brown
130-145 |im tall, with conspicuous oil droplets or granules investing paraphyses, colour
dissolving in K or becoming greenish in parts; epithecium not apparent, concolorous with
thecium. Asci 95-102 x 11-14 ^im. Ascospores pale yellow-brown, broadly ellipsoid, apices
rounded or pointed, 3-septate at maturity, locules irregularly vacuolate, 25-32 x 9 x 11 \im.

Chemistry: 7(3-acetoxyhopan-22-ol, hopane-7(3, 22-diol (tr.), hopane-15a, 22-diol.

Distinguishing features: Pseudocyphellaria fimbriata is an endemic species having elongate-
laciniate, complexly dissected lobes with conspicuously phyllidiate margins (Fig. 64B). The

PSEUDOCYPHELLARJA

141

Fig. 65 Distribution of Pseudocyphellariafimbriata.

142 D. J. GALLOWAY

phyllidia are generally margined with glistening, minute tomental hairs. The upper surface is
coriaceous, minutely areolate-scabrid (x 10 lens), often suffused red-brown at apices. P.
fimbriata has a white medulla, a green photobiont, and a pale red-brown to dark brown velvety
tomentose lower surface, with numerous, prominent, white pseudocyphellae. Apothecia sessile
to subpedicellate, the exciple well-developed, coarsely verrucose-areolate, patchily tomen-
tose. Spores pale yellow-brown, broadly ellipsoid, 3-septate. It has a two-hopane chemistry
[Code A of Wilkins & James (1979)]. Individuals are rather brittle and fragile when dry.

Variation: The species shows rather little variation in morphological characters throughout its
range. It is distinguished from P. coriacea by the marginal and laminal phyllidia, and from P.
pubescens which has entire - margined, uniformly tomentose lobes, yellow pseudocyphellae
below, black apothecial discs which are normally grey- white pruinose, and a chemistry
containing different hopanes, plus several depsides, pigments, and metabolites of the stictic acid
complex. P. fimbriata differs also from P. gretae, a phyllidiate species having densely tomentose
lobes and no detectable chemistry, and from the blue-green P. fimbriatoides which also has
marginally pubescent phyllidia, but thinner, broader lobes which are glossy and much remi-
niscent of Peltigera.

Distribution: (Fig. 65) Rather rare and restricted in North I., from Mt Hikurangi (lat.
3750'S), Mt Egmont, Kaimanawa Range, and the southern Ruahine Range. In South I., from
the mountains of north-west Nelson to Fiordland, close to or west of the Main Divide, and on the
east coast from Banks Peninsula to Invercargill, sea-level to 1500 m.

Habitat ecology: Pseudocyphellaria fimbriata is mainly a rain-forest species, constantly found
in areas with a rainfall of 1600-6500 mm annually, and with spasmodic occurrence in drier areas
(800-1600 mm annually). It is most commonly collected in Nothofagus forests close to and west
of the Main Divide in South I. , from areas having both high precipitation and a high number of
rain days (100-250 days per annum). In the northern part of its range (Mt Hikurangi in the
Raukumara Range), P. fimbriata occurs on exposed rocks with Toninia bullata in subalpine
scrub, but elsewhere it grows on mossy stumps, mossy rocks, and most commonly on tree trunks
in Nothofagus forest, or more rarely in the southern part of its range in lowland coastal forest.

Well-developed specimens form large, often irregular, rosettes on tree trunks in medium to
well-lit habitats with sufficient moisture. Prominent colonies are seen near forest margins or in
rather open forest, and the species is not found in low-light habitats. It is an epiphyte of the
following phorophytes: Dracophyllum traversii, Griselinia littoralis, Nothofagus fusca, N.
menziesii, N. solandri var. cliffortioides , Phyllodadus alpinus, and Weinmannia racemosa.

P. fimbriata associates mainly with other large species of Pseudocyphellaria including P.
colensoi, P. coriacea, P. degelii, P. gretae, P. homoeophylla, P. pickeringii, and P. pubescens,
and species oiMenegazzia, Psoroma, Sphaerophorus, and Usnea.

Specimens examined: 60.

22. Pseudocyphellaria fimbriatoides D. Galloway & P. James Fig. 66

in Lichenologist 15: 142 (1983). Type: New Zealand, South I., Canterbury, Banks Peninsula, below

summit of Mt Sinclair, 670 m, on rocks on forest floor in dense bush, 14 February 1980, J. A. Elix

(CHR 381117! -holotype).
Stictina fragillima var. myrioloba Mull. Arg. in Bull. Soc. r. Bot. Belg. 31: 27 (1892). - Sticta fragillima

var. myrioloba (Mull. Arg.) Hellbom in Bih. K. svenska Vetensk.-Akad. Handl. 21(3/13): 32 (1896).

Type: New Zealand, sine loco, C. Knight 323 (G! - holotype).
Pseudocyphellaria hamataoides D. Galloway & P. James in NZJl Bot. 16: 521 (1978), nom. nud. (Art.

32.1).

Morphology: Thallus 5-10(-15) cm diam., irregularly spreading, closely attached, cortico-
lous, saxicolous, terricolous. Lobes variable, broad and elongate (1x5 cm) to short and
rounded (2-8 mm diam.), apices discrete, contiguous or imbricate, complex-imbricate cen-
trally, plane or canaliculate. Margins entire, sinuous, ascending, rounded or shallowly

PSEUDOCYPHELLARIA

143

Fig. 66 Pseudocyphellaria fimbriatoides . Nelson, Rough Creek, Maruia Valley, Galloway (CHR
379945). A. Scale = 1 cm. B. Scale = 5 mm.

incised, sinuses semi-circular, crenate, ragged, lacerate, and densely phyllidiate, glabrous or
minutely glistening-pubescent, slightly thickened below. Upper surface dark slate blue, suffused
red-brown at apices when moist, pale greyish fawn, dark brownish or livid at apices and margins
when dry, coriaceous, smooth or shallowly dimpled, glossy in parts or matt, often minutely
granular white-papillate (xlO lens), without soredia, isidia, maculae or pseudocyphellae.
Phyllidia marginal, lobulate, dorsiventral, simple, bifurcate to coralloid-branched, con-
stricted at base, scattered and solitary to densely crowded 0-1-2 mm tall, marginally
pubescent-tomentose, glistening. Medulla white. Photobiont Nostoc. Lower surface white and
glabrous, shining, smooth to minutely wrinkled or pitted at margins, tomentose, pale to
dark brown or blackened centrally, tomentum thin, short, and velvety to woolly-arachnoid,
entangled. Pseudocyphellae white, scattered, rather sparse to moderately common, 0-05-0-2
mm diam., slightly punctate-impressed, flat or very shallowly convex, sunk in tomentum at
centre. Apothecia sessile, constricted at base, rather rare scattered, submarginal to laminal,
0-2-2-5 mm diam., shallowly concave at first, becoming plane and at length convex, disc pale
orange-yellow to pale red-brown, matt, epruinose, smooth, margins pale, persistent, exciple

144 D. J. GALLOWAY

pale whitish buff, translucent when moist, delicately arachnoid-scabrid, margins darker than
disc when moist.

Anatomy: Thallus 120-185(-240) um thick. Upper cortex 27-34 um thick, outermost 7-11 um
pale straw-yellow, cells compressed, internally cells thin-walled, colourless, rather loosely
arranged, 4-5-11-5 um diam. Photobiont layer 45-57 um thick, photobiontNostoc, cells to 7 um
diam. Medulla 30-75(-120) um thick, colourless, hyphae 4-5 um diam. Lower cortex 25-32 um
thick, lowest 8-10 um dark red-brown, the remainder colourless, cells thin-walled, similar to
those of upper cortex. Tomental hairs 4-5 um thick, colourless to pale yellow-brown, 30-80 um
long, single, rarely in fascicles. Apothecia: Exciple 90-185 um thick, outermost 10 um pale
yellow-brown, the remainder colourless, cells densely conglutinate, thick-walled at margins,
thinner and more loosely arranged internally, ll-15(-22) um diam. Hypothecium 54-75(-90)
um thick, pale straw-yellow to dilute yellow-brown, changing blue-green on addition of K.
Thecium colourless, 110-120 um tall, without granules or oil droplets; epithecium 18-22 ^m
thick, dilute yellow-brown, unchanged in K. Asci 68-80 x 13-5-18 um. Ascospores fusiform-
ellipsoid, pale brown, 1-3-septate, straight or curved, 27-32 x 7-9 um.

Chemistry: 7(3-acetoxyhopan-22-ol, hopane-7(3, 22-diol (tr.), hopane-15a, 22-diol.

Distinguishing features: Pseudocyphellaria fimbriatoides is an endemic species having rather
variable, though broad lobes, with conspicuously phyllidiate margins (Fig. 70B), the phyllidia
having marginal, glistening tomental hairs. The upper surface is coriaceous, glossy in parts
(reminiscent of some species of Peltigera), and minutely granular, white papillate (x 10 lens). It
has a white medulla, a blue-green photobiont, and a lower surface that is glabrous marginally,
and pale to dark brown tomentose centrally with scattered, small, white pseudocyphellae.
Apothecia are rare, sessile, the exciple pale whitish buff, delicately arachnoid-scabrid. Spores
pale brown, 1-3-septate. It has a two-hopane chemistry [Code A of Wilkins & James (1979)].

Variation: The lobes of P. fimbriatoides are rather variable in shape and in width and length,
and also the phyllidia vary in numbers from sparse to densely crowded. This species is closely
related to P. dissimilis and in many situations the two taxa are sympatric. They are distinguished
by the tomentose phyllidia which are characteristic of P. fimbriatoides. Occasionally, P.
dissimilis produces laminal and/or marginal phyllidia (normally it has terete, simple to branched
isidia) but these are never marginally tomentose, and true isidia are never formed in P.
fimbriatoides. P. fimbriatoides is also distinct from the australian species P. insculpta which has
an identical chemistry and highly divided lobes which at times appear phyllidiate, but the
margins of the lobes are never tomentose, and the upper surface is distinctly punctate-
impressed. The spores of P. insculpta are also longer and broader than those of P. fimbriatoides.
P. fimbriatoides is distinct from P. fimbiata which has a green photobiant, and thicker, more
coriaceous lobes with a distinctive areolate-scabrid upper surface; it is not a photosymbiodeme
of this species.

Distribution: (Fig. 67) North I. , from Coromandel Peninsula (lat. 37S) to Cook Strait, and in
South I. , from north-west Nelson to Southland, both east and west of the Main Divide, sea-level
to 1200 m.

Habitat ecology: Pseudocyphellaria fimbriatoides along with P. dissimilis is, in the main, a
shade-loving species and is most commonly collected from low-light, moist, cool habitats, on the
forest floor (on both rocks and soil), at the base of tree trunks, and on tree roots deep in the
forest interior, or along stream beds. It also grows on damp soil, on shaded rock faces, and can
even withstand periodic inundation on stream bed boulders. It is known from the following
phorophytes: Cordyline australis, Beilschmiedia tawa, Melicytus ramiflorus, Metrosideros
lucida, Nothofagus fusca, N. solandri var. cliff ortioides , Podocarpus totara, Pseudopanax
crass if olius , Pseudowintera color ata, and Senecio eleagnifolius.

Other lichens growing in deeply shaded situations with P. fimbriatoides include Lobaria
retigera, Pseudocyphellaria cinnamomea, P. dissimilis, and Sticta caliginosa.

Specimens examined: 67.

PSEUDOCYPHELLARIA

145

Fig. 67 Distribution of Pseudocyphellariafimbriatoides.

146 D. J. GALLOWAY

23. Pseudocyphellaria glabra (J. D. Hook. & Taylor) Dodge

Figs 68, 69, 70

inB.A.N.Z.R.E. 1929-31 ReptsBJ: 19 (1948) .- Sticta glabra J. D. Hook. & Taylor in Hook. Lond.

J. Bot. 3: 647 (1844). - 5. freycinetii var. glabra (J. D. Hook. & Taylor) Zahlbr. , Cat. Lich. Univ. 3: 347

(1925). - P. freycinetii \ar.glabra (J. D. Hook. & Taylor) Rasanen in AnnlnBot. Soc. zool.-bot. fenn.

Vanamo 2 (1): 35 (1932). Type: Auckland Islands, /. D. Hooker (FH! - lectotype, fide Dodge (1948:

80); BM! - isotype) [see note 1].
Sticta delisea Delise in Mem. Soc. linn. Normandie 2: 94 pi. 9, fig. 32 (1825) nom. superfl. (Art. 63.1). -

Pseudocyphellaria delisea (Delise) D. Galloway & P. James in Lichenologist 12: 297 (1980). - S.

freycinetii var. delisea (Delise) Church Bab. in J. D. Hook., Fl. nov.-zel. 2: 281 (1855). Type: 'He de

King. Nouvelle Hollande, ex Herbar Commerson. Don de mon estimable ami A. L. A. Fee',

ILeschenault de la Tour (PC-LENORMAND! - lectotype; BM!, G!, L 2091-393!, PC-THURET!,

VER! - isotypes) [see note 2].
Sticta freycinetii var. isidioloma Nyl. in Bull. Soc. linn. Normandie II, 2: 504 (1868). Type: New Zealand,

sine loco, D'Urville (H-NYL 33454! - holotype).
Parmelia isabellina Krempelh. in Verhandl. zool.-bot. Ges. Wien 30: 338 (1881). Type: Australia, Mt

Ellery, Gippsland, 1870, Walter (M! -holotype).

Fig. 68 Pseudocyphellaria glabra. New Zealand, Helms 77 (W). Scale = 2 cm.

PSEUDOCYPHELLARIA

147

Fig. 69 Pseudocyphellaria glabra. Denniston, Wa/fcer (BM). Scale = 1 cm.

Sft'cta freycinetii var. glabrescens Mull. Arg. in F/ora, 7ena 66: 23 (1883). - Lobaria freycinetii var.

glabrescens (MM. Arg.) Hellbom in B//I. A:. sve/wfca Vetensfc.-yifcfld. Handl. 21(3/13): 41 (1896).

Type: New Zealand, Middle Island [South !.],>. Lya// (G 003023! - lectotype).
Sticta freycinetii var. conjungens Mull. Arg. in F/ora, /ena 66: 24 (1883). Type: Australia, Moe,

Gippsland, Webb ex F. v. Mueller (G 002071! - holotype; MEL! - isotype).
Sticta freycinetii var. prolifera Mull. Arg. in Flora, /ena 66: 24 (1883). Type: Australia, Moe, Gippsland,

Webb ex F. v. Mueller (G 003054! - lectotype; MEL! - isotype).
Sticta freycinetii var. tenuis Mull. Arg. in Flora, Jena 66: 24 (1883). Type: New Zealand, Kaipara,

S. Mossmann ex herb. Babington, 1879 (G 002096! - lectotype).

Note 1: Sticta glabra}. D. Hook. & Taylor

Thomas Taylor's description of Sticta glabra (Hooker & Taylor, 1844: 647) gives an accurate account of the
isidiate upper surface 'gemmis marginalibus minutis planis oblongis' and also records true apothecia for the
first time 'apotheciis marginalibus subsessilibus concavis extus villosis disco fusco-olivaceo demum nigro
margine lacero gemmifero'. The new species was recorded from 'Lord Auckland's group, Campbell's
Island, Falkland Islands, Cape Horn and Van Diemen's Land' with the comment 'A span or more wide;
unaltered by moisture, apothecia a little larger than turnip seed. Margins of the lobes raised and much
waved. Falkland Island specimens have the lobes very wide'.

While helping Hooker with the initial publication of the lichens of the Antarctic voyage (1839-1843),
Taylor wrote several times of his difficulties with the species of Sticta that Hooker had collected. 'The
Stictae form a prominent feature of lichenological life in Antarctic regions. I am almost afraid that I have
made too many new species. I know not how to help it. Montagne incidentally mentions 'Sticta Delisei'
without any reference . . . , perhaps it is one of our new ones, I have no means of finding out what
Montagne means' (Taylor Correspondence, Royal Botanic Gardens Kew, Director's Correspondence 103
(50), 31 October 1844). Further, after having submitted to Hooker the manuscript of Lichenes Antarctici
[on 2 November 1844] in which nine new species of Sticta were described, Taylor observed 'I find it most
difficult without seeing the coloured plates to compare the descriptions of Antarctic Lichens which you
have kindly copied out with your collected species. And I have the most lively apprehensions lest some of
the species I have believed to be new may turn out to be described already. It is for you my dear Sir, after
receiving back your lichens to compare the new ones, especially the Stictae ... I have not Delise's
Monograph' (Taylor Correspondence ... 103 (53), 15 November 1844).

Joseph Hooker visited Montagne in Paris in 1845 (Galloway, 19776) and saw the extensive collections of
French, Southern Hemisphere, and Antarctic lichens in the herbaria of Montagne and Webb and possibly

148

D. J. GALLOWAY

Fig. 70 Sticta glabra. Isotype (BM). Scale = 1 cm.

also material cited by Delise (\S25a) in his monograph of Sticta. He undoubtedly saw Sticta freycinetii and
realized the similarity of this South American species to the recently described S. glabra, for in the first part
of Flora Antarctica he made S. glabra a synonym of 5. freycinetii with the comment 'An exceedingly
variable plant, of which we have added a character, that of Delise being imperfect [a recognition that the
apothecia in the type of 5. delisea were not true apothecia but a fungal parasite]. The most obvious specific
distinction lies in the pubescent apothecia with fimbriated margins to the cups, to which may be added, the
pale colour, and the wrinkled margins of the lobes, which are sometimes extremely concave. In alpine
specimens the thallus is often quite smooth underneath, with the margins singularly crumpled. The colour
varies, underneath it is of all shades, from black to a dirty yellow. Cape Horn and Falkland Island
specimens are more plane, with the lobes and apothecia larger' (Taylor & Hooker, 1845: 198).

Examination of authentic material of S. glabra annotated by Taylor (BM, FH - 17 specimens seen from
Auckland and Campbell Is, Tasmania, Hermite I., Falkland Is) shows that material from Hermite I. and
the Falklands is Pseudocyphellaria freycinetii while the South Pacific material is P. glabra. Hooker
(1847:528) realized that his earlier circumscription of S. freycinetii included two different taxa and provided
a coloured plate (pi. CXCVI) of true P. freycinetii with the note 'We have added a figure of this
much-disputed species, concerning which we have fallen into an error in the previous part of this work,
having regarded it as synonymous with the 5. glabra of Lord Auckland's group and Tasmania (probably the
5. delisea Fee), and which differs from the S. freycinetii principally in the very shallow, not deeply cupped
apothecia'. Babington (1855:281) in recording S. freycinetii var. delisea from New Zealand ignored the
presence of isidia as a distinguishing character and declared 'A careful consideration of a large number of

PSEUDOCYPHELLARIA 149

specimens has convinced me that the Sticta freydnetii of the two parts of the 'Flora Antarctica' are only
forms of one and the same species'.

Du Rietz (1924), in emending Pseudocyphdlaria freydnetii to include only non-isidiate taxa, included
material referable to P. glabra in the synonymy of P. chloroleuca not having seen authentic material of
Sticta chloroleuca which is an isidiate species quite distinct from P. glabra. Galloway & James (1980:297)
followed Du Rietz in making P. chloroleuca a synonym of P. glabra, an error which is discussed further in
Galloway (19866), and in this paper (see above under P. chloroleuca, p. 85).

Dodge (1948) discusses the typification of Sticta glabra from material in Taylor's herbarium (FH) and
chooses a fertile, Auckland I. specimen as lectotype (Dodge, 1948:80), a treatment which is adopted here
in accordance with Art. 8.1 since it predates the lectotypification of 5. glabra made by Galloway & James
(1980: 297).

Note 2: Sticta delisea Delise

A. L. A. Fee described the monotypic genus Delisea for a curious Southern Hemisphere lichen differing
from any species of Sticta then known, by the singular nature of the apothecia. It was collected from King
Island in Bass Strait separating Australia and Tasmania. Fee (1824-1825: xcv, ci, tab. 2, fig. 15) named the
species pseudosticta and illustrated it with a life-like coloured engraving. Delisea Fee, a later homonym of
Delisea Lamouroux (Rhodophyta), was renamed Plectocarpon by Fee (1824-1825: 151). Later in the same
year the material was sent to Delise who included it in his Sticta monograph as 5. delisea (Delise 1825a),
attributing the epithet to Fee. The material examined and described by Delise was recently discovered in
PC-LENORMAND (Hawksworth & Galloway, 1984; Galloway & James, 1986) and is labelled in Delise's
hand 'He de King. Nouvelle Hollande. Ex herbar. Commerson. Don de mon estimable ami A. L. A. Fee'.
Philibert Commerson (1727-1773), naturalist with L. A. de Bougainville on La Boudeuse and I'Etoile
during the circumnavigation of 1766-1769, never visited King Island and it is likely that the Australian
material incorrectly ascribed by Fee and Delise to Commerson came from a later French expedition to
Australia. Nelson (1974, 1975) has shown that several plant records mistakenly attributed to La Billardiere
[the first French botanist to collect in Australia, Filson, 1976; Kantvilas, 1983; Galloway, 1985a] are
properly assigned to Leschenault de la Tour, botanist to the expedition commanded by Nicolas Baudin
which visited King L, in December 1801. It seems probable that Leschenault was the collector of the
material described by Fee, and Delise (Galloway & James, 1986).

The type material from Delise's herbarium (now in PC-LENORMAND) shows the specimen to be a
sterile, isidiate species of Pseudocyphellaria attacked by a lichenicolous fungus causing apothecia- like galls
to develop on the thallus. Hawksworth & Galloway (1984) typified the genus Plectocarpon on the
lichenicolous fungus element and designated the host lichen element as lectotype of the name Sticta delisea.
However, Sticta delisea is a superfluous name for Delisea pseudosticta, since this latter name [erroneously
printed as Delisea sticticoides (Delise 1825:94)] is cited as a synonym by Delise (Art. 63.1). The use of the
name Pseudocyphellaria delisea in several recent accounts is therefore erroneous (e.g. Galloway, 19856,
19866; Galloway & James, 1980, 1986; Galloway etal. , 1983; Hawksworth & Galloway, 1984). The earliest
valid name for this Southern Hemisphere isidiate species is therefore Sticta glabra.

Morphology: Thallus rosette-forming to irregularly spreading, 5-10(-25 +) cm diam. , loosely
adnate centrally, margins closely attached to ascending, corticolous, terricolous, saxicolous.
Lobes very variable, narrowly laciniate and almost straplike and discrete from margins to
centre, 2-6 mm wide and 15-4 mm long, to 10-17 mm wide and imbricate-complex from near
margins to centre. Margins entire, sinuous to crenulate or shallowly or deeply notched or
incised becoming isidiate with age to somewhat erose-glomerulate. Upper Surface fresh lettuce
green to pale greenish yellow when moist, pale greenish to yellowish fawn often suffused
brownish at margins when dry (alpine forms whitish to red-brown or blackened in exposed
sites), undulate, shallowly wrinkled or obscurely ridged in parts, never faveolate, alpine forms
very coriaceous often conspicuously wrinkled and deeply cracked, cracking occasional to
extensive, random, not in any pattern, matt to glossy, occasionally with small dimples,
without soredia, maculae, phyllidia or pseudocyphellae. Isidia sparse to copiously developed,
mainly marginal or from surface cracks, terete, fingerlike, mainly simple, occasionally furcate,
0-5-1 mm tall, often crowded-congested. Medulla white. Photobiont green. Lower surface
smooth to irregularly wrinkled-shining, very variable, glabrous, pale buff or whitish to brown or
brown-black in a narrow to wide marginal zone, tomentose centrally, or completely glabrous, or
entirely tomentose from margins to centre, tomentum thin to thick and woolly, even, not
tufted, pale brown to black, occasionally completely glabrous and black. Pseudocyphellae

150 D. J. GALLOWAY

common, prominent, white, round to irregular, 0-1-1 mm diam., raised, verruciform-papillate,
margins of papillae free from tomentum, decorticate area flat or shallowly excavate. Pycnidia
occasional to rare, mainly marginal or submarginal, punctate-impressed, rather inconspicuous,
0-1 mm diam. or less, central ostiole pale red-brown, minute, surrounded by a slightly raised
margin concolorous with thallus, often eroding and leaving small pits. Apothecia sessile to
subpedicellate, sparse to occasionally crowded and deformed through mutual pressure, often
absent, deeply to shallowly concave, plane to shallowly undulate and subconvex with age, round
to irregular, 0-5-7 mm diam. , margins entire to crenate-striate or occasionally stellate-ragged,
formed from coarsely splitting edges of exciple, disc pale to dark red-brown, obscured at first by
exciple, matt, epruinose, exciple translucent when moist, whitish to buff-pink when dry,
brownish on storage in herbarium, arachnoid-tomentose at first and obscuring disc with a
thick plug of tomentum, coarsely scabrid-areolate with age.

Anatomy: Thallus 135-200(-370) um thick. Upper cortex 45-56 um thick, upper 11-12 urn
straw yellow, of necrotic or compressed cells appearing fibrous and periclinal, lower 30-40 um
colourless, thin-walled, 5-13 um diam. Photobiont layer 35-45 um thick, photobiont green, cells
round, 9 um diam. Medulla 90-250 urn thick, hyphae periclinally arranged, compact towards
lower cortex, more loosely arranged near photobiont layer. Lower cortex 22-27 um thick,
outermost 11-12 urn dark red-brown (dissolving in K), inner zone straw-yellow to hyaline,
cells thick-walled, 5-7 um diam. Tomental hairs strongly septate, locules short, 4-5-7 um long,
slightly constricted at septa, red-brown, solitary or in fascicles, rather short, 35-110 um long.
Apothecia: Exciple 45-90 um thick at margins, to 180 um below disc, cells 7-ll(-15) um'diam. ,
colourless, in rows splitting away as pyramidal clumps or as thick- walled, short tomental hairs.
Hypothecium 45-75 um thick, dilute orange brownish or pale straw yellow. Thecium 65-80 um
tall, colourless; epithecium 11-13 um thick, dilute yellow-brown to red-brown, dissolving in K,
granular, colour external to tips of paraphyses. Asci 65-81 x ll-15um. Ascospores pale
yellow-brown, fusiform, apices pointed, mainly 1-septate, 3-septate at maturity, not thickened,
(13-5-)15-5-20-5(-22-5) x 4-5-7 urn.

Chemistry: 7(3-acetoxyhopan-22-ol, hopane-7[3, 22-diol (tr.), hopane-15a, 22-diol, norstictic,
stictic, cryptostictic, constictic, hypostictic (tr.), hyposalazinic (tr.), and usnic acids.

Distinguishing features: Pseudocyphellaria glabra is a widespread austral lichen having nar-
rowly laciniate to broad, imbricate-complex lobes with entire, sinuous to crenulate or incised
margins, often isidiate. Upper surface glossy, undulate, shallowly wrinkled, never faveolate,
isidiate. Isidia marginal or laminal, often from surface cracks, terete, finger-like, mainly simple
to occasionally furcate or palmate-flattened. It has a white medulla, a green photobiont, and a
pale to dark brown or black, glabrous to tomentose lower surface with prominent white
pseudocyphellae. Apothecia occasional, sessile to subpedicellate, exciple whitish to buff-pink,
prominent, coarsely scabrid-areolate. It has a two-hopane chemistry [Code A of Wilkins &
James (1979)] with the addition of the stictic acid group of metabolites and usnic acid (), which
confers a yellow-green colour to the upper surface, especially in habitats exposed to full sunlight.

Variation: Pseudocyphellaria glabra is a very polymorphic species colonizing a wide variety of
habitats in New Zealand from coastal forest and/or scrub, to Nothofagus forest, and mixed
hardv/ood-Nothofagus forest, subalpine scrub at and above tree-line, and subalpine to alpine
grasslands and herbfield. It shows a wide range of thallus morphology throughout these different
habitats, lobes varying from narrow, discrete, and straplike, to broad, rounded, imbricate,
and with a lower surface that may be entirely glabrous to centrally or entirely tomentose, pale
buff or brown to entirely black. Thallus thickness varies a good deal depending on the degree of
exposure to sun, frost, and wind. Shaded specimens from forest and scrub communities are pale
green, deficient in usnic acid, and rather thinner and less coriaceous than exposed subalpine
forms which tend also to be less isidiate, are more yellow, and often partially or totally
blackened, or suffused red-brown. Some alpine forms are whitish or greyish and are generally
very thick and coriaceous. All forms have characteristic terete (usually simple), finger-like isidia;
these are often abraded, and tend to lie scattered on the upper surface. Forest forms are usually

PSEUDOCYPHELLARIA

151

Norfolk I

Lord Howe I

Three Kings Is

172

174

176

178

The Snares

40 c

-42

46

166

168

I

170

172

174

40-

42 '

176

Fig. 71 Distribution of Pseudocyphellaria glabra.

152 D. J. GALLOWAY

more richly isidiate (especially at the margins) than those from above tree-line. Although a wide
diversity of thallus form exists in this species, there is no chemical variation observed.

Pseudocyphellaria glabra is the isidiate counterpart of the New Zealand endemic P.
homoeophylla. It is also closely related to P.. corbettii, a species having marginal phyllidia and
broad lobes. The South American P. freycinetii is related to P. glabra and has a similar
chemistry, but it lacks isidia and has larger, thinner, more folded, and papery lobes.

Distribution: (Fig. 71) Widely distributed from North Auckland (lat. 35S) southwards to
Stewart I., and the subantarctic islands (Snares, Auckland, Campbell, Antipodes, and
Macquarie), coastal to alpine, sea-level to 2000 m.

Habitat ecology: Pseudocyphellaria glabra has the widest range and ecological tolerance of any
species of Pseudocyphellaria in New Zealand, being known from all forest types, coastal and
subalpine scrub, subalpine to high-alpine grassland and herbfield. It grows luxuriantly in the
wettest areas of the country, in forests west of the Main Divide and especially in Fiordland, and
is also found in the very driest areas, in Central Otago and eastern parts of South I. In forest
and/or scrub associations it is an epiphyte of branches and trunks, and in some drier areas will
grow on soil, on rocks, and amongst bryophytes. It has been collected from the following
phorophytes: Agathis australis, Aristotelia serrata, Coprosmafoetidissima, Dacrycarpus dacry-
dioides, Dacrydium bidwillii, D. cupressinum, Dracophyllum latifolia, D. longifolium, Fuchsia
excorticata, Griselinia littoralis, Hebe spp., Kunzea ericoides, Leptospermum scoparium,
Libocedrus bidwillii, Melicytus ramiflorus, Metrosideros lucida, M. umbellata, Myrsine austra-
lis, M. divaricata, Myrtus bullata, Nothofagus menziesii, M. solandri var. cliff ortioides , Olearia
colensoi, Pimelea spp., Pittosporum eugenioides, P. tenuifolium, Podocarpus hallii, P. nivalis,
P. totara, Phyllocladus alpinus, Prumnopitys ferruginea, Pseudopanax edgerleyi, P. simplex,
Pseudowintera color ata, Quintinia acutifolia, Rhopalostylis sapida, Senecio eleagnifolius , S.
reinoldii, and Weinmannia racemosa.

Pseudocyphellaria glabra is also found in subalpine bogs, straggling amongst tussocks, or on
rock outcrops in subalpine grassland, and in herbfield at the lower limit of the high-alpine zone.

Specimens examined: 507.

24. Pseudocyphellaria granulata (Church. Bab.) Malme Fig. 72

in Bih. K. svenska Vetensk. -Akad. Handl. 25 (3/6): 21 (1899). - Sticta granulata Church. Bab. in J. D.
Hook.,F/. nov.-zel. 2: 281 (1855). -5. richardi var. granulata (Church. Bab.)Nyl. in Mem. Soc. Imp.
Sci. nat. Cherbourg 5: 335 (1857). - Stictina carpoloma subsp. granulata (Church. Bab.) Nyl., Syn.
meth. lich. 1 (2): 338 (1860). - Sticta carpoloma var. granulata (Church. Bab.) J. D. Hook. , Handb. N.
Zeal. fl. 2: 568 (1867). - Phaeosticta granulata (Church. Bab.) Tevisan, Lichenotheca veneta exs. 75
(1869). - Lobaria granulata (Church. Bab.) Kuntze, Revis. gen. pi. 3: 384 (1893). Type: New Zealand,
Middle Island [South l.],D. Lyall (BM! - holotype).

Babington (1855:281) says of Sticta granulata, 'Our Lichen resembles S. pulmonacea a good deal in its
ample size and general mode of division, but seems to be as nearly allied to the preceding as to any other.
The olive-green colour, the scrobiculated thallus, and, above all, the tendency of the plant to produce
copious dirty coralline pulvinate soredia, often covering the centre, and the irregular, ill-developed, dirty
yellow cyphellae, are its most obvious characters. Found also in Tasmania by Gunn and by Hooker . . . The
imperfectly-formed cyphellae show a transition to the structure of the simple naked spots of 5. scrobiculata
and its allies, which constitute a section (Lobaria) not yet found in New Zealand'.

Robert Gunn's Tasmanian specimen, originally in Taylor's herbarium is labelled 'Sticta granulata Tayl.
MSS' in Thomas Taylor's hand. In Taylor's herbarium (FH) four Gunn specimens so named by Taylor are
preserved, together with one from Swan River [Western Australia] collected by James Drummond which is
also named 5. granulata Tayl., but which is referable to Pseudocyphellaria neglecta. In Babington's
herbarium (BM), the holotype specimen is labelled by Babington 'S. granulata Tayl ! Bab. in Hook. Fl.
N.Z.', so he obviously used Taylor's name. No New Zealand material of P. granulata is preserved in
Taylor's (FH) herbarium.

Morphology: Thallus orbicular to irregularly spreading, 5-12(-25) cm diam., corticolous or
saxicolous, loosely to closely attached centrally, margins free. Lobes deeply incised,

PSEUDOCYPHELLARIA

153

Fig. 72 Pseudocyphellaria granulata. Lake Te Anau, Hale 65085 (US). Scale = 5 mm.

linear-elongate to broadly rounded, 0-5-2-5 cm diam. and 1-5 cm long, discrete or contiguous at
margins, complex-imbricate centrally. Margins very irregular, rarely entire, more often
ragged, notched, incised, crenulate-lacerate, flat to sinuous and ascending, coarsely and
densely sorediate, often eroded-white. Upper surface glaucous-green to olivaceous, often
suffused greyish or blackened when moist, grey-fawn or glaucous yellowish, often blackened
in exposed sites when dry, plane to sub-convex, undulate, dimpled, punctate-impressed,
wrinkled-plicate to faveolate-reticulate, with erumpent, pustular soralia, often coalescing
into lines or patches, often cracked or fissured, coriaceous, matt, without isidia, maculae,
phyllidia or pseudocyphellae. Soredia eroding lobe margins and erupting in rounded, pustular
soralia on lamina, also developing from breaks or fissures, coarse, granular-glomerulate, in
dense clusters, pale olive greenish eroding whitish, often blackened, sometimes appearing
pseudoisidiate. Medulla white. Photobiont green. Lower surface pale buff or whitish at lobe
margins, black centrally, tomentose from margins to centre, or with a narrow, glabrous, pale
buff, delicately wrinkled, shining marginal zone, tomentum thick, woolly. Pseudocyphellae
frequent, conspicuous, white, round to irregular 0-1-1-5 mm diam., in shallow to well-
developed papillae, margins black, slightly raised, prominent, often fissured, decorticate area
flat, coarsely granular, sunk in tomentum. Pycnidia occasional, hemispherical, laminal, 0-2-0-5
mm diam., ostiole punctate, black, minute. Apothecia sessile to subpedicellate, rather rare,
marginal and laminal, often most common towards lobe apices, 1-3 mm diam., round to
irregular, contorted through mutual pressure, margins pale to dark red-brown, crenulate-
roughened to tomentose, massive at first and completely obscuring disc in young fruits, disc
black, matt or granular, coarsely white-pruinose at first, pruina uneven, areolate (x 10 lens),
lost with age, exciple pale buff to red-brown, translucent when moist, roughened, granular-
verrucose.

Anatomy: Thallus 165-280(360) urn thick, to 540 um thick at cephalodia. Upper cortex 27-34
urn thick, upper 10-12 um pale yellow-brown becoming faint violet in K, lower zone colourless,

154 D. J. GALLOWAY

cells 6-5-13-5 um diam. Photobiont layer 34-56 um thick, photobiont green, cells rounded, to
11-5 um diam. Medulla 90-200 um thick, colourless, hyphae to 5 um diam. Lower cortex 27-35
um thick, red-brown, dark blue-green in K, densely conglutinate, cells 5-9 um diam. Tomental
hairs to 5 um diam. , in dense fascicles, red-brown, to 230 um long. Apothecia:Exciple colourless,
120-200 um thick, yellow-brown towards margins, cells 714 um diam. Hypothecium 82-95 um
thick, pale yellow-brown to straw-yellow, unchanged in K. Thecium colourless, 100-110 um tall,
granular or with numerous oil droplets; epithecium 7-14 um thick, smoky olive-grey to
olive-brown, purple-violet in K. Asci 76-80x13-15 um. Ascospores smoky grey to olive-
brown, oval-ellipsoid, thickened 1-septate to 3-septate, apices pointed, sometimes papillate,
septum variable in thickened 1-septate spores 1-5-4-5 um thick, locules uneven, irregular-
deformed, 22-5-32 x 9-11-5 um.

Chemistry: Hopane-7(3, 22-diol (tr.), hopane-6ct, 7(3, 22-triol, 7p-acetoxyhopane-6a, 22-diol
(tr.), 6a, acetoxyhopane-7(3, 22-diol (tr.), methyl virensate, physciosporin, norstictic (tr.),
stictic, cryptostictic (tr.), and constictic acids [Code B of Wilkins & James (1979)].

Distinguishing features: Pseudocyphellaria granulata is an austral species having deeply in-
cised, linear-elongate to broadly rounded lobes, with irregular, ragged or incised-crenulate,
sorediate-eroded margins. The upper surface is plane to punctate-impressed to faveolate in
parts, with erumpent, pustular soralia at margins, and scattered over, to thickly covering the
upper surface, or restricted to reticulate ridges. The medulla is white, the photobiont green,
and the lower surface uniformly black, or with a pale buff to whitish glabrous marginal zone,
thickly tomentose and with conspicuous, white pseudocyphellae. Apothecia are sessile to
subpedicellate, rather rare, the disc black, coarsely white-pruinose at first, the exciple pale buff
to red-brown, roughened-granular-verrucose. The epithecium is smoky grey to olive-brown,
becoming purple-violet in K. Spores are grey to olive-brown, thickened 1-septate to 3-septate.
The chemistry is distinctive, containing physciosporin as a major metabolite [Code B of Wilkins
& James (1979)].

Variation: Pseudocyphellaria granulata is a rather variable species, showing a wide diversity of
lobe shape and size, and with a surface texture varying from plane to reticulate-faveolate,
depending on local microclimate and microhabitat conditions. The numbers and position of
soralia also show a wide amplitude of variation from marginally sorediate forms to forms
totally sorediate in a dense laminal crust. Soredia are always present and are the major
taxonomic character differentiating this species from all other related white-medulla species.
The non-sorediate species P. faveolata has an identical chemistry and in many habitats is
sympatric with P. granulata. The two species constitute a species pair, and occur widely to-
gether throughout the austral zone. In some recent accounts, records of P. granulata from
northern, offshore islands (Hayward & Hayward, 1978, 1980, 1982, 19826, 1984) refer to
P. haywardiorum.

Distribution: (Fig. 73) North I., south of lat. 3650'S, to Cook Strait, and in South I., from
Nelson to Fiordland, also in Stewart I., lowland to subalpine, sea-level to 1000 m.

Habitat ecology: Pseudocyphellaria granulata is a common epiphyte of lowland coastal forest
and scrub, and especially of successional vegetation (notably Fuchsia and Leptospermum) in
altered sites, though it is also often found near treeline and in subalpine scrub. It is a moderately
photophilous species and will not tolerate deep shade, and most commonly associates with other
species of Pseudocyphellaria in areas of moderate to high rainfall. It will also colonize rocks in
tussock grassland and grows on dead stumps as well as the following phorophytes: Coprosma
spp. , Cordyline australis, Dacrycarpus dacrydioides, Dracophyllum subulatum, Fuchsia excorti-
cata, Griselinia littoralis, Kunzea ericoides, Leptospermum scoparium, Metrosideros excelsa,
Myrsine australis, Nothofagus menziesii, N. solandri var. cliff ortioides , Picea abies, and Wein-
mannia racemosa.

Specimens examined: 65.

PSEUDOCYPHELLARIA

155

Fig. 73 Distribution of Pseudocyphellaria granulata.

156 D. J. GALLOWAY

25. PseudocyphellariagretaeD. Galloway

Fig. 74

in Lichenologist 15: 143 (1983). Type: New Zealand, South I., Cantebury, Boyle River near Lewis
Pass, on bark ofNothofagus menziesii, 13 September 1981, D. /. Galloway (CHR 381118! -holotype,
BM!-isotype).
Pseudocyphellaria hirsutula D. Galloway & P. James in NZJl Bot. 16: 521 (1978), nom. nud. (Art. 32.1)

Morphology: Thallus orbicular or spreading in irregular rosettes, 5-10(-18) cm diam.,
rather loosely attached from margins to centre, corticolous. Lobes sublinear-laciniate to broadly
rounded, apices discrete to subimbricate, usually densely isidiate centrally, 0-5-1 -5(-2-5) cm
wide and l-2(-5) cm long. Margins entire, delicately notched or incised to crenulate, often
densely isidiate, slightly thickened below, markedly re volute at lobe apices. Upper surface
bright lettuce-green, occasionally suffused brownish at margins when fresh, pale grey-green to
glaucous-buff, yellowish olive or pinkish in parts when dry, becoming dark red-buff to brownish
on storage, undulate, mainly convex, rarely plane to subconcave, shallowly and irregularly
wrinkled or dimpled, never faveolate, internal cephalodia visible as small, rounded papillae
0-2-0-5 mm diam., texture rather delicate, densely to sparsely tomentose, tomentum white,
silky, often abraded or absent in older parts, cortex below tomentum smooth to minutely
scabrid-verrucose and reticulate-cracked, soredia, maculae, phyllidia, and pseudocyphellae
absent. Isidia marginal and laminal, delicate, fragile, brittle, granular to coralloid, 0-2-0-5 mm
tall, flattened, appearing squamiform, in rows or dense clusters, often spreading over surface

Fig. 74 Pseudocyphellaria gretae. Holotype (CHR). A. Scale = 1 cm. B. Scale = 5 mm.

PSEUDOCYPHELLARIA 157

as a thick coralloid crust, uniformly tomentose. Medulla white. Photobiont green. Lower
surface white, pale buff to pinkish or brownish centrally, densely tomentose from margins to
centre, or with extensive glabrous patches centrally in older specimens, shallowly wrinkled or
dimpled in parts, tomentum thick, silky, white, rather short. Pseudocyphellae minute, often
inapparent, scattered, sparse to moderately common, obscured by tomentum, flat or slightly
papillate, 0-05-0-3(-0-8) mm diam., round to irregular, mainly white, occasionally pale
yellowish. Pycnidia rather rare, laminal, punctate to papillate, 0-1 mm diam. or less, ostiole
red-brown. Apothecia very rare, marginal and laminal, sessile at first, becoming distinctly
pedicellate, 0-5-2(-3-5) mm diam., shallowly to deeply concave, margins involute,
isidiate-ragged often obscuring disc, disc dark red-brown to brown-black, matt, epruinose,
exciple delicately scabrid-verrucose, tomentose, concolorous with thallus.

Anatomy: Thallus 105-202 um thick. Upper cortex 34-45(-50) um thick, upper surface very
uneven splitting into irregular pyramidal areas (scabrosity), colourless to pale straw-yellow,
cells 4-5-11-5 um diam. Tomental hairs 45-140(-230) um long, 7-11 um thick, sparsely septate.
Photobiont layer 23-27 (-36) um thick, densely compacted, photobiont green, 7-9 um diam.,
Dictyochloropsis. Medulla 55-92(-120) um thick, colourless, compact, hyphae 2-5-3-5 um
diam. Lower cortex 27-34(-45) um thick, colourless or pale straw-yellow, cells 7-14 um diam.
Tomental hairs to Sum diam. , 45-135(-180) um long, colourless. Apothecia: Exciple colourless,
90-140 um thick, cells thick-walled, 9-15 um diam. Hypothecium 45-73 um thick, upper 20 um
colourless, remainder dilute orange-brown, unchanged in K. Thecium colourless, 100-120 um
tall; epithecium 7-21 um thick, yellow-brown to red-brown, colour in gel external to tips of
paraphyses, dissolving in K; paraphyses distinctly vacuolate (? oil droplets), moniliform at
apices. Asci 78-90 x 11-5-15-5 um. Ascospores brown, broadly ellipsoid, thickened 1-septate to
3-septate, locules lozenge-shaped, irregular, apices rounded or pointed 23-29-5 x 11-5-13-5
um.

Chemistry: t.l.c. negative.

Pseudocyphellaria gretae is named for the late Greta Du Rietz who collected lichens widely
throughout New Zealand in 1926-1927 with her husband G. Einar Du Rietz.

Distinguishing features: Pseudocyphellaria gretae is an endemic species forming irregular
rosettes on bark and having broadly rounded to sublinear lobes with entire to incised, crenulate,
often densely isidiate margins. The upper surface is densely to sparsely tomentose (Fig. 74B),
often with granular to coralloid tomentose isidia spreading as a thick, diffract crust. It has a white
medulla, a green photobiont, and whitish, densely tomentose lower surface with minute,
scattered, white to pale yellow pseudocyphellae. Apothecia are very rare, marginal, and
laminal, distinctly pedicellate with ragged, isidiate margins and a tomentose, areolate-scabrid
exciple, concolorous with thallus. Spores are brown, thickened 1-septate to 3-septate. There is
no detectable chemistry.

Variation: Pseudocyphellaria gretae shows some variation in lobe morphology in the degree of
development of tomentum, and in the numbers and position of the isidia. All specimens
examined were tomentose to some extent and all were isidiate. Well-developed specimens from
beech forest are usually thickly tomentose and richly isidiate, and these characters also seem to
correlate with medium to high rainfall. Specimens from drier areas are often thinner, cracked or
torn and of much smaller size. Pseudocyphellaria gretae is distinguished from P. pubescens by
the marginal and laminal isidia, the epruinose apothecial discs which are red-brown and not
black, by the white pseudocyphellae, and by the lack of acetone-soluble metabolites detectable
on t.l.c. These characters also distinguish P. gretae from P. fimbriata which has marginal,
tomentose phyllidia, coriaceous, irregularly elongate-laciniate lobes which are not tomentose, a
dark brown lower surface with large white pseudocyphellae, and a two-hopane chemistry.

Specimens collected by Richard Helms (1842-1914) from near Greymouth (H-NYL, W) are
annotated in Nylander's hand 'Sticta obvoluta Ach.' under which name it was recorded in
Nylander (1888^:36-7), Miiller Argoviensis (1894), and Hellbom (1896- as Lobaria obvoluta}.

158

D. J. GALLOWAY

-40

Norfolk I

Lord Howe I

Three Kings Is

The Snares

Auckland Is
\

Campbell I
Macquarie I

160

-42

-44

46

174 C

176

166

_J

168

170

I

172

L_

174

176

I

178

34-

40-

42-

44 c

Fig. 75 Distribution of Pseudocyphellaria gretae.

PSEUDOCYPHELLARIA 159

As Pseudocyphellaria obvoluta it is recorded by Martin (1966, 1969a) and Martin & Child
(1972), though in this last account, the plant referred to is obviously P. pubescens.

Distribution: (Fig. 75) North I., from Raglan Harbour (Lat. 3650'S) to Wellington, and in
South I. , from Nelson to Fiordland and eastwards from north Canterbury near the Lewis Pass to
Invercargill. Also on Stewart I., from Oban to Port Pegasus, mainly lowland to subalpine,
sea-level to 1200 m.

Habitat ecology: Pseudocyphellaria gretae is best developed in areas of high rainfall and is
especially well-developed on beech trees near the Lewis Pass where very large rosettes are often
found on trunks. It appears to prefer moist, cool, humid conditions in moderate but not dense
shade, though smallish specimens on scrub can withstand full sunlight. P. gretae is known from
the following phorophytes: Dacry carpus dacrydioides, Griselinia littoralis, Leptospermum
scoparium, Nothofagus fusca, N. solandri var. cliffortioides , N. menziesii, Podocarpus totara,
Pseudopanax crassifolius , and Sphaeropteris medullaris. P. gretae associates with the following
lichens: Coccocarpia palmicola, Degelia durietzii, Leioderma amphibolum, L. sorediatum,
Lobaria adscripta, Menegazzia caliginosa, M. lucens, Nephroma australe, Phlyctella megalo-
spora, Parmelia testacea, P. tenuirima, Pseudocyphellaria billardierei, P. fimbriata, P. glabra, P.
pickeringii, P. pubescens, P. multifida, P. homoeophylla, Sphaerophorus melanocarpus , 5.
ramulifer, S. tener, Usnea capillacea, and U. xanthophana.

Specimens examined: 40.

26. Pseudocyphellaria haywardiorum D. Galloway, sp. nov. Fig. 76

Diagnosis: Pseudocyphellariae bartlettii similis sed thallus supra punctulatus non faveolatus, subtus
pseudocyphellis niveis, 0-1-2 mm latis. Sporae dilute fuscescentes, (27 -5-) 30- 32 (-34) x 6-7 umm.
Medulla K-, KC-, C-, Pd-. Typus: New Zealand, North Island, South Auckland, Red Mercury Island, on
tea tree (Leptospermum) bark, August 1971, B. W. & G. C. Hayward H 40-4 (AK 161261! - holotype).
[t.l.c.: 7(3-acetoxyhopan-22-ol, hopane-7(3, 22-diol (tr.), hopane-15a, 22-diol].

Named for Bruce and Glenys Hayward (Lower Hutt) for their collections of, and researches into, the
lichens of the offshore islands of northern New Zealand.

Morphology: Thallus orbicular, 3-7 (-12) cm diam., corticolous, loosely to firmly attached
from margins to centre, or margins ascending. Lobes rounded to irregularly laciniate,
2-10(-20) mm wide, 4-25 mm long, discrete, contiguous or complex-imbricate and clustered.
Margins sinuous, ascending, very irregular, delicately to coarsely crenate-lacerate, slightly
thickened below, or with clustered soredia and grey-white, erose below. Upper surface dark
grey-blue or glaucous-blue to blue-black, darker at lobe apices or with a livid brownish tinge
when moist, olivaceous-brownish to glaucous-yellowish or isabelline, dark greyish at margins
when dry, plane or shallowly undulate \ wrinkled-faveolate at apices, elsewhere conspicuously
punctate-impressed, here and there with minute, pale buff papillae (x 10 lens), matt, smooth or
appearing minutely arachnoid in parts, rather coriaceous when dry, soft and flabby when moist,
sorediate, maculate, without phyllidia or pseudocyphellae. Maculae minute, scattered, irregu-
lar, whitish buff, most noticeable and in reticulate patterns towards lobe apices, occasionally
more extensive photobiont-free patches centrally. Soredia coarsely granular to crowded-
pseudoisidiate (x 10 lens), scattered in efflorescent clusters, 0-5 mm diam., often dense and
spreading, delimited in round to irregular, laminal soralia, or in sinuous, linear, marginal
soralia, or coalescing and forming broad areas of sorediate to pseudoisidiate crust, soredia
dark brownish blue, glaucous-greyish, often eroding white. Medulla white. PhotobiontNostoc.
Lower surface bullate or irregularly undulate-wrinkled, pale buffer brownish and glabrous
in a narrow, marginal zone, elsewhere evenly and often densely tomentose, red-brown to
brown-black, tomentum short and velvety near margins, long and woolly centrally. Pseudocy-
phellae white, conspicuous, well delimited from tomentum, common centrally, rare marginally,
round to irregular, 0-1-2 mm diam., with a conspicuously raised, pale-buff, glabrous margin,
decorticate area concave to convex, granular. Apothecia rare or absent, marginal and laminal,
solitary or in groups (2-7), sessile, constricted at base to subpedicellate, rounded, irregular or

160 D. J. GALLOWAY

Fig. 76 Pseudocyphellaria haywardiorum. Holotype (AK). Scale = 1 cm.

compressed-distorted through mutual pressure, (0-l-)0-5-2-5(-3) mm diam, concave at first,
becoming plane to convex at maturity, disc pale yellow or orange-brown to dark red-brown,
shining at first, then matt and slightly roughened with age, epruinose, exciple pale-buff to
brownish, translucent when moist, finely to coarsely scabrid-verrucose.

Anatomy: Thallus 220-460(-550) um thick. Upper cortex 34-45 um thick, cells 4-5-9 urn
diam., outermost 10-12 um yellow-brown to red-brown, remainder hyaline to pale straw.
Photobiont layer 45-75 um thick, with prominent dark red-brown hyphae between bundles of
photobiont, and running from cortex to medulla, photobiont Nostoc. Medulla colourless,
90-360 um thick, hyphae to 4-5 um diam. Lower cortex 2-3 rows of round to irregular
thick- walled cells, outermost layer dark red-brown, inner two rows colourless to pale straw,
4-5-11-5 um diam. Tomental hairs yellow-brown to red-brown, 45-230 um long, in fascicles.
Apothecia: Exciple cellular, 45-110 um thick, without photobiont, outermost layers pale
red-brown, remainder colourless to straw-yellow. Hypotheciwn (30-)45-65 um thick, dilute
yellow-brown, unchanged in K. Thecium colourless, 75-95 um tall, epithecium 18-23 um
thick, clear yellow-brown to red-brown, unchanged in K. Asci 55-70(-80) x 13-15(-20)
um. Ascospores fusiform-ellipsoid, apices pointed, yellow-brown, 1-septate, unthickened,
(27-5-)30-32(-34) x 6-7 um [spore measurements taken from a specimen from Rangitoto I.,
collected by H. H. Allan (CHR)].

Chemistry: 7|3-acetoxyhopan-22-ol, hopane-7|3, 22-diol (tr.), hopane-15a, 22-diol. [Code A of
Wilkins& James (1979)].

Distinguishing features: Pseudocyphellaria haywardiorum is an Australasian sorediate species
having rounded to irregularly laciniate lobes with coarsely granular to pseudoisidiate (x 10
lens) laminal and marginal soralia, and a conspicuously punctate-impressed upper surface.
Neither surface depressions nor soralia are arranged in a reticulate pattern. It has a white

PSEUDOCYPHELLARIA

161

Lord Howe I

Norfolk I

Three Kings Is

Kermadec 30-
Is

The Snares

Auckland Is
\

Campbell I
\

Macquarie I

Bounty Is

\

Antipodes Is

160 C

170 C

40

42

-44

166

168

50 l

180

172

170

172

174

176

176
1

178

42-

Fig. 77 Distribution of Pseudocyphellaria haywardiorum.

162 D. J. GALLOWAY

medulla, a blue-green photobiont, and prominent large, white pseudocyphellae, well delimited
from the densely and evenly tomentose, red-brown to brown-black lower surface. It has a
two-hopane chemistry [code A of Wilkins & James (1979)].

Variation: This characteristic, northern coastal forest species is still rather poorly known and
still rather seldom collected, and the 15 specimens examined in this study show little variation in
most external characters except lobe configuration (broadly rounded to irregularly laciniate). I
have not studied the species in the field so can report little about either its morphological
variation or its habitat ecology. P. haywardiorum is distinguished from P. intricata by the
punctate-impressed upper surface and the bullate lower surface with its prominent, large,
raised pseudocyphellae, characters which also separate it well from P. argyracea, a related
isidiate species. It is also distinct from P. bartlettii in lobe configuration (P. bartlettii has broad,
rounded lobes and always forms neat rosettes), soralium morphology, and in chemistry, P.
bartlettii being C+ rose (cortex) and K + yellow (medulla) while P. haywardiorum is negative to
both spot tests. P. bartlettii, which is also much thinner in texture, also lacks the conspicuous
punctate-impressed upper surface of P. haywardiorum and is instead, weakly to strongly
reticulate-faveolate. P. haywardiorum is also superficially similar to Lobaria scrobiculata, but
the latter does not have pseudocyphellae and it has a distinctive and different chemistry [medulla
K+ orange-red, C rose, Pd+ orange. Constictic, norstictic, stictic, and usnic acids, and
scrobiculin] and, moreover, is scabrid-areolate near the margins which are never maculate.
Apothecia appear to be rather rarely developed in P. haywardiorum, only two of the specimens
examined were fertile and two others were infected with a lichenicolous fungus.

Distribution: (Fig. 77) In coastal forest from Radar Bush, North Auckland to Rangitoto I.,
near Auckland city. Absent on the Three Kings Is, but present on the following northern
offshore islands: Poor Knights Is, Lady Alice I., Hen I., Little Barrier I., Rakitu I., and Red
Mercury I. It has an altitudinal range from sea-level to 240 m. In some previous accounts
(Hayward & Hayward, 1978, 1980, 19820, 19826, 1984) the species is recorded as P. granulata.

Habitat ecology: Pseudocyphellaria haywardiorum is an epiphyte of both bark and twigs of the
following phorophytes in northern coastal forest: Cor dy line australis, Kunzea ericoides, and
Metrosideros excelsa. It is known also from K. ericoides in grassland. It is one of a group of
northern species (P. aurata, P. bartlettii, P. carpoloma, and P. poculifera are others) closely
associated with Cordyline, Metrosideros, and Leptospermum in lowland and primarily coastal
habitats north of lat. 37S.

Specimens examined: North Island. North Auckland: Radar Bush, 7 May 1976, /. K. Bartlett
24641 (Herb. Bartlett, BM) - also collections (unnumbered) from 1 & 2 January 1980, /. K.
Bartlett (Herb. Bartlett, BM); Bay of Islands, Waiparo Bay, B. W. & G. C. Hayward, January
1980 (AK 161552); Poor Knights Islands. Tawhiti Rahi, September 1980, B. W. Hayward (AK
164496, BM); Lady Alice Island, B. W. & G. C. Hayward (AK); Little Barrier Island, B. W. &
G. C. Hayward L33, L20 (AK); Rakitu Island, January 1981, B. W. Hayward (AK 164490);
Rangitoto Island, 24 April 1927, G. E. & Greta Du Rietz 2673: 3 (UPS); 13 April 1936, H. H.
Allan (BM, CHR); January 1935, H. H. Allan (BM, CHR); L. H. Millener (BM, CHR).

27. Pseudocyphellaria homoeophylla (Nyl.) Dodge Fig 78

in Nova Hedwigia 19: 489 (1971). - Sticta homoeophylla Nyl. in Flora, Jena 50: 439 (1867). - Lobaria
homoeophylla (Nyl.) Hellbom in Bih. K. svenska Vetensk.-Akad. Handl. 21(3/13): 39 (1896). Type:
New Zealand, sine loco (prob. Wellington), 1867, C. Knight (H-NYL 33464! - holotype; WELT,
Herb. Knight Vol. 36 A, p. 9! -isotype).

Sticta amplificata Zahlbr. in Denkschr. Akad. Wiss. Wien math.-naturwiss. Kl. 104: 287 (1941). Type:
New Zealand, Ohakune, L. B. Moore ZA 167 (W 2506! - holotype; CHR 374654! - isotype).

Morphology: Thallus orbicular to spreading, often forming extensive swards, 10-20(-50) cm
diam., loosely adnate centrally, margins free, projecting outwards or ascending, corticolous,
saxicolous or terricolous. Lobes very variable, linear-laciniate, di- or trichotomously

PSEUDOCYPHELLARIA

163

Fig. 78 Pseudocyphellaria homoeophylla. Nina Valley, Galloway (CHR 374645). A. Scale = 1 cm. B.
Scale = 5 mm.

branched at apices, complex-imbricate centrally, 2-5(-10) mm wide, occasionally to 25 mm
wide, and rarely to 40 mm wide centrally (in S. amplificatd), and to 12 cm long. Margins entire,
sinuous, shallowly scalloped or notched at apices, smoothly rounded and markedly thickened
and sometimes ridged below, without pseudocyphellae, sinuses semicircular. Upper surface
bright lettuce-green when wet, pale greenish grey when dry, becoming yellowish buff on
prolonged storage in herbarium, smooth, matt, coriaceous, undulate, shallowly wrinkled or
pitted, occasionally with a crazing of fine black lines, or obscurely ridged, never reticulate-
faveolate, without isidia, maculae, pseudocyphellae or soredia. Medulla white. Photobiont
green. Lower surface pale yellowish buff to brown at margins, red-brown to black centrally,

164 D. J. GALLOWAY

shallowly wrinkled-plicate or pitted, internal cephalodia visible as scattered, hemispherical,
wrinkled swellings 1-2 mm diam., patchily tomentose from margins to centre or with extensive
marginal and central glabrous areas, tomentum short, entangled, white or buff to blackened.
Pseudocyphellae common, conspicuous, white, round to irregular, 0-1-1 mm diam. , strongly
excavate, crateriform, with strongly defined, raised margins. Apothecia marginal or submar-
ginal, most common towards lobe apices, l-4(-6) mm diam., rounded, concave at first
becoming shallowly undulate with age, pedicellate to sessile, disc pale to dark red-brown,
blackened with age, matt, epruinose, margins conspicuously denticulate or striate to stellate,
pale buff, translucent when wet, exciple translucent when wet, pale buff or pinkish or creamish
when dry, thickly arachnoid at first and obscuring disc for a time after emergence with a thick
plug, becoming coarsely areolate-scabrid with age and projecting from margins at maturity as a
stellate fringe. Pycnidia submarginal, marginal (often in lines) or scattered on upper surface,
hemispherical, minute, 0-2 mm diam. or less, often eroded and leaving shallow pits, commonly
clustered near lobe apices.

Anatomy: Thallus 180-260 (to 360 at margins) urn thick. Upper cortex 45-54(-64) urn thick,
upper zone of 15-20 um appearing fibrous, pale straw-coloured or yellow-brown, of necrotic,
compressed cells, lower zone abutting photobiont layer, colourless, of round to irregular,
thin-walled cells in a pseudoparenchyma, cells 4-5-15 um diam. Photobiont layer 18-27 um
thick, photobiont green, cells rounded, 7-9 um diam. Medulla 90-200 um thick, hyphae loosely
interwoven, 2-3 um diam., encrusted with granular crystals and appearing greyish olive.
Lower cortex 27-34(-45) um thick, outer 10-12 um pale to dark red-brown, inner zone
adjoining medulla, hyaline to pale straw-coloured of thick-walled round to irregular,
isodiametric cells in 6-8 rows, walls 2-2-5 um, lumina 2-3 um diam. Tomental hairs septate,
red-brown, 7-9 um thick, septate, constricted at septa, locules short, single or in scattered
fascicles 4-8-together, 45-140(-230) um long. Apothecia: Exciple cellular, colourless, 90 um
thick at margins, 180-230 um thick below disc, cells 10-15 um diam., thicker in outer parts of
tissue, splitting below into large, pyramidal clumps (surface scabrosity) with 1-3 rows of cells
extending at margins as short, tomental hairs. Hypothecium 45-75 um thick, very pale yellow or
orange-brown, dense. Thecium 80-90(-100) um tall, colourless, yellow-brown towards tips of
paraphyses; epithecium 11-22 um thick, pale yellow-brown, slightly granular, colour dissolving
in K. Medulla and a narrow photobiont zone (30 um) between hypothecium and exciple. Asci
78-85 x 18-20-5 um. Ascospores fusiform-ellipsoid, apices rounded or pointed, straight or
curved, pale straw, 1-3-septate, not thickened, 25-32 x 7-9 um.

Chemistry: 7(3-acetoxyhopan-22-ol, hopane-7(5, 22-diol (tr.), hopane-15a, 22-diol, norstictic,
stictic, cryptostictic, constictic, methylstictic, hypostictic, and hyposalazinic (tr.) acids, and
usnic acid (Elix, 1986).

Distinguishing features: Pseudocyphellaria homoeophylla is an endemic species having di- or
trichotomously branching, linear-laciniate lobes, with entire, sinuous margins (Fig. 78B),
lacking soredia, isidia, phyllidia, or pseudocyphellae. The upper surface is glabrous, smooth to
shallowly wrinkled or pitted, never faveolate, coriaceous. It has a white medulla, a green
photobiont, and a red-brown to black lower surface with rather patchy tomentum and with
numerous, conspicuous, white pseudocyphellae. Apothecia are sparse to moderately common,
mainly marginal and towards apices, the exciple prominent, coarsely areolate-scabrid, buff to
pinkish, often stellate-fimbriate at margins. Spores are fusiform-ellipsoid, pale straw-
coloured, 1-3-septate. The chemistry comprises two main hopanes [Code A of Wilkins & James
(1979)] with the addition of stictic acid metabolites and usnic acid, the last of which gives the
thallus a yellowish tinge.

Variation: Pseudocyphellaria homoeophylla varies most widely in the width of its lobes,
particularly exuberant forms, especially in the northern part of its range, having very broad
lobes [25-40 mm wide as in Sticta amplificata] while most southern collections are narrower 2-10
mm wide. The entire, sinuous, somewhat scalloped margins, which are devoid of asexual
propagules, distinguish this species from the two most closely related taxa, the isidiate P. glabra

PSEUDOCYPHELLARIA

165

Fig. 79 Distribution of Pseudocyphellaria homoeophylla.

166 D. J. GALLOWAY

and the marginally phyllidiate P. corbettii, both of which have the same chemistry as P.
homoeophylla. It is distinct from the South American subantarctic species P. freycinetii which
has more irregular, indented-lacerate margins and much larger, broader (0-5-4-5 cm wide),
shorter, thinner, and more papery lobes (Galloway & James, 1986).

Specimens recorded in Zahlbruckner (1941) as Sticta dissimulata (p. 283) are referable to
P. homoeophylla, as are several specimens recorded as Sticta freycinetii (ZA 407 & 667; V 206;
V 60; V 61 -W!), or as Sticta sinuosa (W 10; ZA 161 & 666).

Pseudocyphellaria homoeophylla is the counterpart primary species to the isidiate P. glabra,
the two taxa forming a species pair. It is, however, known only from New Zealand and does not
share the same austral range of distribution as P. glabra.

Distribution: (Fig. 79) North I. , south of lat. 37S on most mountain ranges, on both east and
west coast to Wellington (Akatarawa Range), and in South I., from Nelson and Marlborough
Sounds close to and west of the Main Divide to Fiordland, also in the Catlins (Southland) and in
Stewart I., lowland, montane to subalpine, sea-level to 1440 m.

Habitat ecology: Pseudocyphellaria homoeophylla is primarily a rain-forest species and is best
developed in beech (Nothofagus) forest close to the Main Divide in South I. It is especially
common both as an epiphyte and as a forest floor species where it competes successfully with
bryophytes (in some areas it may form extensive swards) in mid-altitude forests (200-600 m) and
in areas dominated by mountain beech (Nothofagus solandri var. cliffortioides) it is the
dominant epiphyte up to treeline where increased light and exposure allows other species of
Pseudocyphellaria to proliferate. Its often great size and luxuriant development, along with P.
colensoi and P. coronata, make it one of the most prominent of cryptogamic epiphytes in New
Zealand beech forest, and it undoubtedly contributes substantially to the forest biomass. Apart
from species of beech (TV. fusca, N. menziesii, N. solandri), it is known from the following
phorophytes: Agathis australis, Dacrycarpus dacrydioides , Libocedrus bidwillii, Myrsine
divaricata, Pseudopanax, Quintiniaserrata, and Weinmannia racemosa.

Specimens examined: 140.

28. Pseudocyphellaria hooker! (Church. Bab.) D. Galloway & P. James Fig. 80

in Lichenologist 12: 299 (1980). - Sticta hookeri Church. Bab. in J. D. Hook., Fl. nov.-zel. 2: 282
(1855). - Stictina hookeri (Church. Bab.) Nyl. , Syn. meth. lich. 1(2): 336 (1860). - Saccardoa hookeri
(Church. Bab.) Trevisan, Lichenotheca veneta exs. 75 (1869). - Cyanisticta hookeri (Church. Bab.)
Gyelnik in Feddes Reprium Spec. nov. veg. 29: 2 (1931). Type: New Zealand, Bay of Islands, /. D.
Hooker (BM! - lectotype).

Morphology: Thallus orbicular, rosette-forming, loosely attached centrally, margins free,
ascending, 5-10(-15, rarely to 22) cm diam., corticolous. Lobes broadly laciniate, rounded at
apices, 0-5-2-5 cm diam., discrete or subimbricate at apices, complex-imbricate centrally,
plane or shallowly convex towards apices, shallowly concave centrally. Margins entire,
sinuous, shallowly crenate to irregularly convolute or sublobulate, shining, often suffused
brownish, slightly thickened below or inflated, with occasional minute, punctiform, pale
yellow pseudocyphellae. Upper surface dark slate-blue or blue-grey, suffused red-brown in
parts especially towards margins when moist, glaucous-grey to red-brown when dry, strongly
reticulate-faveolate, faveolae weakly to strongly defined, shallow or deep, interconnecting
ridges rather short, smoothly rounded to sharply delimited, matt to glossy, smooth or wrinkled,
without soredia, isidia, phyllidia or pseudocyphellae. Maculae common, effigurate, whitish to
pale buff, scattered, or forming an irregular reticulum, larger photobiont-free areas visible as
whitish or pale buff spots or blotches. Medulla white. Photobiont Nostoc. Lower surface pale
buff or whitish at margins, darker centrally, irregularly wrinkled-bullate especially at lobe
apices, glabrous at margins, elsewhere uniformly short, stiff, tomentose, tomentum
brownish often in discrete bundles. Pseudocyphellae minute, common, on raised, nodular to
elongate verrucae, margins concolorous with lower cortex, conspicuous, decorticate area often
reduced to a thin slit or scarcely apparent, pinprick-like, very pale yellow or whitish. Pycnidia

PSEUDOCYPHELLARIA

167

Fig. 80 Pseudocyphellaria hookeri. Mangaotaki Valley, Pio Pio, Galloway (CHR 375877). Scale = 1 cm.

numerous, scattered, marginal and laminal, or in rows on thalline ridges, papillate, 0-1-04 mm
diam., ostiole red-brown, punctate. Apothecia sessile to pedicellate, sparse to frequent, round
to irregular, 1-8 mm diam. , concave at first becoming plane to irregularly convex-undulate with
age, disc dark brown to black, matt or minutely granular-papillate, epruinose, margins
concolorous with thallus, crenate-striate, involute when young, exciple concolorous with
thallus, smooth and shining to wrinkled or faveolate, occasionally scabrid-areolate (x 10
lens), in places minutely tomentose, tomentum thin, brownish.

Anatomy: Thallus 100-230(-320) urn thick. Upper cortex 22-27 um thick, dilute straw-yellow
above, colourless below, cells 4 urn diam. or less. Photobiont layer 36-55 (-64) um thick, in
irregular clumps, photobiontNostoc, cells to 11 um diam. Medulla 45-200 um thick, colourless,
hyphae to 4-5 um diam. Lower cortex 11-5-15-5 um thick, dilute yellow-brown, dissolving in K,
cells 4-5 um diam. or less. Tomental hairs colourless to pale brown, 45-230 um long, in scattered
fascicles. Apothecia: Exciple colourless, 73-110 um thick, irregularly scalloped at margins, cells
thick-walled, 4-5-12 um diam. Hypothecium dilute yellow-brown, intensifying in K, 40-45 um
thick. Thecium colourless, 100-112 um tall; epithecium granular, not dissolving in K, 7-14 um
thick, dark olive brownish turning violet-purple in K. Asci 63-76 x 15-5-18 um. Ascospores
smoky olive-brown, thickened 1-septate to 3-septate, broadly fusiform, straight or curved,
septum of thickened 1-septate spores 4-5-7 um thick, contents granular, (18-)23-27 x 7-9 um.

Chemistry: Methyl evernate, tenuiorin, two unidentified depsides (tr.), methyl lecanorate,
methyl gyrophorate, evernic acid (tr.), gyrophoric acid, hopane-6a, 7(3, 22-triol, 7(3-
acetoxyhopane-6a, 22-diol (tr.), 6a-acetoxyhopane-7(3, 22-diol (tr.), norstictic (tr.), stictic,
cryptostictic, constictic, and hypostictic acids (tr.), and traces of pulvinic acid, pulvinic
dilactone, and calycin.

168

D. J. GALLOWAY

-40

Lord Howe I

Norfolk I

Three Kings Is

Kermadec 30-
Is

The Snares

Auckland Is
\

Campbell
\

Macquarle I

Chatham Is

Bounty Is
Antipodes Is

160 C

170

-42

-44

166

i

168

BO-

172

170

L_

172

174

176

174

176

I

178

34-

36-

38-

40-

42"

Fig. 81 Distribution of Pseudocyphellaria hookeri.

PSEUDOCYPHELLARIA 169

Distinguishing features: Pseudocyphellaria hookeri is an endemic, rosette-forming species
having broadly laciniate lobes with entire, sinuous, crenate to irregularly convolute-sublobulate
margins. The upper surface is strongly reticulate-faveolate, with whitish or buff maculae but
without soredia, isidia, phyllidia or pseudocyphellae. It has a white medulla, a blue-green
photobiont, and a wrinkled-bullate, buff or brown, tomentose lower surface with scattered,
minute, whitish or pale yellow pseudocyphellae. Apothecia common, sessile to subpedicellate,
disc black, granular-roughened, epruinose, exciple concolorous with thallus. Epithecium
granular, olive, turning violet-purple in K. Spores olive-brown, thickened 1 -septate to 3-
septate. It has a complex chemistry containing gyrophoric acid (cortex C+ red), depsidones,
hopane-6a, 7(3, 22-triol, and stictic acid metabolites.

Variation: Pseudocyphellaria hookeri is a species of uniform morphology throughout its range .
It is distinguished from P. durietzii by the blue-green photobiont and the brown tomentum of the
lower surface; however, the two species are closely similar morphologically and chemically and
photosymbiodemes of the two species are known. P. montagnei is similar chemically, but differs
in the thinner, more fragile, less faveolate lobes, in the closely appressed habit (P. hookeri is
usually only centrally attached and is free and subascendent at the margins), the pale buff
sparsely tomentose lower surface with pale yellowish insignificant sparse pseudocyphellae, and
the laminal phyllidia and phyllidiate apothecial margins.

Distribution: (Fig. 81) Mainly a northern species in North I., from lat. 3450'S at Cape
Karikari, to the Manawatu, throughout. Rare in South I. , (north-west Nelson and Jackson Bay),
mainly lowland to subalpine, sea-level to 400 m.

Habitat ecology: Pseudocyphellaria hookeri is primarily a northern species, characteristic of
cool, moist, humid habitats (gullies, stream sides, on successional shrubs, or in standing forest or
forest remnants), in moderate to deep shade. It is mainly a twig species and is often richly
developed on young regenerating Dacry carpus dacrydioides. It is also known from the following
phorophytes: Coprosma spp., Leptospermum scoparium, Myrtus bullata, Podocarpus hallii,
Salix, Weinmannia racemosa, and W. silvicola.

Specimens examined: 30.

29. Pseudocyphellaria intricata (Delise) Vainio Fig. 82A, B

in Hedwigia 37: 35 (1898) . - Sticta intricata Delise in Mem. Soc. linn. Normandie2: 96 pi. 7 fig. 33 (1825) .
- Stictina intricata (Delise) Nyl., Syn. meth. Lich. 1(2): 334 (1860). - Cyanisticta intricata (Delise)
Gyelnik in Lilloa 3: 76 (1938). Type: lie de Bourbon (Reunion), Bory de Si-Vincent (PC-
LENORMAND! - lectotype) [see note 1].

Sticta thouarsii Delise in Mem. Soc. linn. Normandie 2: 90 pi. 8 fig. 29 (1825). - Stictina intricata var.
thouarsii (Delise) Nyl., Syn. meth. lich. 1(2): 335 (1860). - Pseudocyphellaria intricata var. thouarsii
(Delise) Vainio in Hedwigia 37: 35 (1898). - Stictina thouarsii (Delise) Hav., Lich. Exs. Norveg. 430
(1913). - Sticta intricata f. thouarsii (Delise) Zahlbr., Cat. Lich. Univ. 3: 388 (1925). - Cyanisticta
thouarsii (Delise) Rasanen, Die Flechten Estlands 1: 161 (1931). - Pseudocyphellaria thouarsii Degel.
inActa Phytogeogr. Suec. 7: 150 (1935). - Cyanisticta intricata var. thouarsii (Delise) Rasanen in Revta
Univ. Santiago 21: 144 (1936). Type: Tristan da Cunha, Du Petit Thouars (PC-LENORMAND! -
holotype) [see note 1].

Sticta limbata var. subflavida Church. Bab. in J. D. Hook., Fl. nov.-zel. 2: 283 (1855). - Stictina limbata
var. subflavida (Church. Bab.) Nyl. in Hue in Nouv. Archs Mus. Hist. not. Paris III, 2: 300 (1890).
Type: New Zealand, Middle Island [South Island], sine loco, D. Lyall (BM! -lectotype) [see note 2].
Fig. 82 A.

Stictina intricata var. subargyracea Nyl., Lich. Nov. Zel. : 29 (1888) [qua forma nomen Nyl., Flora, Jena
69: 172 (1886)] . - Sticta intricata f . subargyracea (Nyl.) Hue in Nouv. Archs Mus. Hist. not. Paris IV, 3:
90 (1901). Type: New Zealand, sine loco (prob. near Greymouth, Westland), Helms 35 (W! -
lectotype) [see note 3] .

Stictina subcrocea Stirton in Trans. N.Z. Inst. 32: 74 (1900). - Sticta subcrocea (Stirton) Zahlbr., Cat.
Lich. Univ. 3: 399 (1925). Type: Australia, Queensland, Brisbane, F. M. Bailey (BM! -lectotype).

Note 1: Discussing the use of the names intricata and thouarsii, J0rgensen (1977: 23) states: 'when the two

170

D. J. GALLOWAY

Fig. 82 Pseudocyphellaria intricata. A. Lectotype of Sricta limbata var. subflavida (BM- left). Scale = 1
cm. B. New Zealand, Helms 93 (W). Scale = 5 mm.

species are united the correct name is Pseudocyphellaria intricata . . . Nylander (1860) united them
choosing intricata as the species epithet, and he must be followed'.

Note 2: Sticta limbata var. subflavida Church. Bab.

Babington (1855: 283) states that his taxon scarcely differs from Delise's Sticta thouarsii, the colour of the
upper surface being 'plus minus fuscoflavescentibus', and further, 'This can hardly be distinguished with
safety from the European Lichen. Delise's figure of 5. thouarsii well represents the ramification, and differs
in nothing material except in the white soredia, which are decidedly lead-coloured in the New Zealand
plant.' The lectotype of 5. limbata var. subflavida is the left hand specimen on a sheet (BM) containing
collections of Lyall, and Colenso (Fig. 82 A).

Note 3: Stictina intricata var. subargyracea Nyl.

The lectotype of this taxon (W) was collected by Richard Helms and is in Herbarium Lojkanum. It consists
of three specimens having a mustard-yellow upper surface (of the subflavida-type see above), broadly
rounded lobes, and mainly laminal soralia, although towards the centre, lobes have linear, marginal
soralia. A sheet in WU labelled Nymphaea intricata f. subargyracea Nyl., has seven specimens of P.

PSEUDOCYPHELLARIA 171

intricata collected in New Zealand (probably from near Grey mouth in Westland) by Richard Helms.
Specimens from both W and WU are growing on Leptospermum bark and seem to be from the same
collection, having associated with the dominant P. intricata, and other lichens, including Leioderma
sorediatum, Leptogium cyanescens, Pannaria fulvescens , and Psoroma leprolomum.

Morphology: Thallus very variable, from narrow-lobed, furcate, to irregularly divided lacini-
ate demes to broad-lobed, rounded demes, 2-10(-20) cm diam., rosette-forming to irregu-
larly spreading, closely attached centrally, margins free and often ascendent, corticolous,
rarely saxicolous. Lobes irregularly linear-laciniate, rather ragged, or furcate or straplike, or
broadly rounded, apices discrete, contiguous or imbricate, 3-10(-25) mm wide, 1-5 cm long,
discrete or separate in part from margins to centre, or complexly folded centrally with apices
only discrete. Margins rarely entire, usually delicately crenulate to ragged-lacerate,
ascending, sinuous, often becoming coarsely sorediate and eroding both upper and lower
surfaces. Upper surface dark glaucous blue-green, olivaceous-glaucous or blue-black suffused
brownish or reddish when moist, pale olivaceous greyish, mustard-yellow, grey-brown, or
reddish brown sometimes with a lilac tinge when dry, plane, undulate, minutely wrinkled or
occasionally subfaveolate to punctate-impressed towards lobe apices, smooth to minutely
scabrid in parts (x 10 lens), rarely shining, coriaceous, rarely minutely tomentose in scattered
patches near lobe apices, sorediate, without isidia, phyllidia or pseudocyphellae. Soralia laminal
and marginal, Soredia farinose to coarsely granular, often appearing pseudoisidiate or with
clustered, corticate, regenerating lobules, laminal soralia punctiform, or eroding large areas,
round to irregular 0-1-3 mm diam., flat to distinctly raised, pulverulent, widely scattered,
crowded or in lines on shallow ridges near margins, marginal soralia linear, sinuous, eroding
lower surface, often labriform, soredia white, occasionally brownish or greyish. Maculae
minute, scattered, irregular, whitish or buff patches (x 10 lens), not in any reticulate pattern,
only seen when moist. Medulla white. Lower surf ace plane, undulate or irregularly to complexly
wrinkled-ridged, pale yellowish or whitish-buff to grey-brown, red-brown or chocolate-brown,
rarely with a narrow, glabrous, marginal zone, tomentose from margins to centre, tomentum
stiff, silky, even, velvety to woolly-entangled. Pseudocyphellae white, rather rare, widely
scattered, submerged in tomentum, often absent, round to irregular, 0-2-1-5 mm diam.,
margins thin, raised, concolorous with lower cortex, decorticate area plane to deeply concave,
granular. Apothecia very rare, normally absent, laminal, sessile to subpedicellate, 0-2-2 mm
diam., rounded, concave at first becoming plane or undulate at maturity, disc pale orange-
brown to red-brown, matt or glossy, epruinose, immarginate at maturity, exciple slightly
roughened, pale buffer brownish, concealing disc at first, excluded at maturity.

Anatomy: Thallus 200-370 urn thick. Upper cortex 34-40 \im thick, outermost 10 um pale
straw-yellow, cells compressed, remainder colourless, cells 4-5-9 urn diam. Photobiont layer
50-90 um thick, rather variable, photobiont Nostoc. Medulla 100-220 um thick, colourless,
hyphae 4-5 um diam. Lower cortex (13-)18-22 um thick, pale yellow-brown, cells 4-5-9 um
diam., Tomental hairs pale yellow-brown to red-brown, to 5 um diam. and 45-220 um long, in
dense fascicles. Apothecia: Exciple colourless, 40-90 um thick, cells 9-15 um diam. Hypo-
thecium pale yellow-brown or olivaceous-brown 28-45 um thick. Thecium pale straw-
yellow, 90-110 um tall; epithecium absent \paraphyses densely conglutinate, not much swollen
at apices ( to 3 um diam.). Asci 76-86 x 15-18 \im. Ascospores pale yellow-brown, fusiform-
ellipsoid, straight or curved, 3-septate, slightly constricted at septa, contents distinctly
vacuolate, (27-)29-5-36-5(-38-5) x 7-9 um.

Chemistry: Tenuiorin (tr.), methyl gyrophorate (tr.), 7p-acetoxyhopan-22-ol, hopane-7|3,
22-diol (tr.), hopane 15a, 22-diol.

Distinguishing features: Pseudocyphellaria intricata is a cosmopolitan species having irregu-
larly laciniate to broadly rounded lobes with sinuous, incised to entire, often sorediate
margins. The upper surface is plane, undulate to minutely wrinkled or subfaveolate in parts,
coriaceous with scattered, erose, laminal and marginal soralia, containing coarsely granular,
often pseudoisidiate soredia. It has a white medulla, a blue-green photobiont, and a pale

172 D. J. GALLOWAY

yellow-buff to chocolate-brown tomentose lower surface with occasional to rare, widely
scattered, white pseudocyphellae submerged in tomentum. Apothecia generally absent, when
present sessile to subpedicellate, the disc pale orange-brown to red-brown, exciple slightly
roughened, pale brownish, excluded at maturity. It has a two-hopane chemistry [Code A of
Wilkins & James (1979)] with traces of tenuiorin and methyl gyrophorate.

Variation: Pseudocyphellaria intricata is a widespread and rather variable species, and lobe
morphology, colour of the upper surface, and numbers and position of the soralia all vary,
depending on local microhabitat and microclimate conditions. Thick, broad-lobed individuals
having a coriaceous, mustard-yellow upper surface and coarse grey-blue, often marginal soralia
are common in drier areas and in habitats exposed to relatively high light intensities (Fig. 82 A).
Specimens from rocks in rough pasture land or near bush margins are most commonly of this
form and were referred by Babington (1855) to Sticta limbata var. subflavida. In wetter, more
humid, and more shaded habitats, the upper surface is red-brown to pale grey-brown often with
a bluish or lilac tinge, and with linear, marginal, and scattered laminal white soralia, often
pseudoisidiate at the margins (Fig. 82B). Such forms are typical in forested areas, or from damp,
humid scrub in partial shade, and are frequently found growing amongst moss. In some northern
collections (especially from the Waiotapu Valley near Lake Rotorua, and also from near Lake
Taupo) specimens of P. intricata have thin, papery lobes with mainly marginal, sinuous soralia,
and are strongly reminiscent of material of this species from Tristan da Cunha, the typical
locality of Sticta thouarsii Delise (Galloway & James, 1986).

The New Zealand populations, although of variable morphology depending on local ecologi-
cal conditions, all have a uniform chemistry and are presently regarded as a single variable
species. The sorediate P. crocata also exhibits a variety of ecotypic forms; in fact morphological
diversity in this taxon is somewhat greater than that seen in P. intricata. Accessory depsides and
depsidones occur in P. intricata (Galloway & James, 1986), although stictic, salazinic, and
consalazinic acids which are recorded from Indian Ocean collections are not found in specimens
from New Zealand. Variation in P. intricata is also discussed by Coppins & James (1979) and by
Galloway & James (1980).

Pseudocyphellaria intricata is distinguished from the related species P. argyracea which has
delicate, simple to coralloid isidia at the lobe margins and also at the margins of laminal
pseudocyphellae which do not form soredia. In a number of cases pseudoisidia are formed at the
margins of laminal soralia and are then difficult to separate from forms of P. argyracea where the
laminal isidia may lose their cortex and become sorediate. It is possible that a continuum
of forms exist between strictly sorediate P. intricata and strictly isidiate, laminally pseudo-
cyphellate P. argyracea. Detailed field work should elucidate this point. P. intricata is distinct
from the sorediate P. haywardiorum (from northern New Zealand and north eastern Australia)
which has a strongly punctate-impressed upper surface and a dark red-brown, bullate lower
surface, and prominent numerous white pseudocyphellae.

Distribution: (Fig. 83) Throughout both main islands from Radar Bush, North Auckland (lat.
3425'S), to Invercargill, and on Stewart I., from Oban to Port Pegasus, sea-level to 1150 m.

Habitat ecology: Pseudocyphellaria intricata is primarily a forest species, growing on bark,
amongst other lichens, and in very damp, humid, shaded habitats amongst mosses on twigs of
shrubs. It is tolerant of a wide range of light regimes, being found in dense shade, and from rocks
exposed to full sunlight at forest and scrub margins in areas of moderate to high rainfall.
Generally, it is a good indicator of moist, humid habitats and associates with the following
lichens: Coccocarpia palmicola, Coenogonium implexum, Collema spp., Degelia gay ana,
Hypotrachyna sinuosa, Hypogymnia subphysodes, Heterodermia podocarpa, Leioderma
amphibolum, L. pycnophorum, L. sorediatum, Leptogium brebissonii, Nephroma rufum, N.
cellulosum, Normandina pulchella, Pseudocyphellaria argyracea, P. crocata, P. bartlettii, P.
fimbriatoides , P. glabra, P. hookeri, Psoroma leprolomum, P. sphinctrinum, Sticta fuliginosa,
S. limbata, S. martinii, S. subcaperata, Usnea arida, and U. torquescens.

It is an epiphyte of the following phorophytes: Beilschmiedia tawa, Carpodetus serratus,

PSEUDOCYPHELLARIA

173

Norfolk I

Kermadec 30-
Is

Lord Howe I

Three Kings Is

The Snares

Auckland Is
\

Campbell I
\

Macquarie I
160

40

-44

166
_J

168

170

172

174 C

178 C

36-

40-

42-

44.

176 C

I

Fig. 83 Distribution of Pseudocyphellaria intricata.

174

D. J. GALLOWAY

Dacry carpus dacrydioides, Dodonaea viscosa, Griselinia littoralis, Kunzea ericoides, Leptosper-
mum scoparium, Metrosideros excelsa, Myrtus bullata, Nothofagus menziesii, N. solandri var.
cliff ortioides , Pennantia corymbosa, Rhopalostylissapida, and Weinmannia racemosa.

Specimens examined: 1 06 .

30. Pseudocyphellaria jamesii D. Galloway, sp. nov.

Fig 84

Diagnosis: Pseudocyphellariae coerulescentis similis sed lobis parvis, angustis, pagina superior passim
tomentosus, subtus luteolus, pseudocyphelliis sparsis, minutis, immarginatis. Apothecia ignota. Typus:
New Zealand, South I., Canterbury, Boyle River near Lewis Pass, on twigs of Leptospermum scoparium
on terraces behind Boyle Lodge growing with P. maculata, September 1981. P. W. James (BM! -
holotype). [t.l.c.: pulvinic acid, pulvinic dilactone, calycin, and a mixture of stictane triterpenoids similar
to those found in P. ardesiaca.}

Morphology: Thallus rosette-forming, 2-5(-8) cm diam. , loosely attached centrally, free at
margins, corticolous. Lobes rather short and narrow, 1-2 mm, rarely to 8-10 mm wide, 0-5-2-5
cm long, concave to plane or shallowly convex, subdichotomously branching and discrete at
margins to rounded, imbricate-folded marginally, complexly folded, imbricate centrally.

Fig. 84 Pseudocyphellaria jamesii. Boyle River, Galloway (CHR 374943). Scale = 1 cm. B. Scale = 1

mm.

PSEUDOCYPHELLARIA 175

Margins slightly thickened below, slightly ascending, sinuous above, rarely entire, more usually
crenate, notched or occasionally minutely lobulate, with conspicuous, raised verruciform,
yellow pseudocyphellae. Upper surface dark grey-blue suffused red-brown at apices when moist,
pale grey-blue or glaucous-blue when dry, cyanobiont-free areas yellow-buff, shallowly to
strongly reticulate faveolate at margins, ridges less obvious centrally, or irregularly wrinkled-
plicate, here and there patchily tomentose, tomentum thin, pale buff to whitish, marginal and
laminal, matt, without soredia, isidia, phyllidia or pseudocyphellae. Maculae prominent, yellow
or whitish, laminal and marginal, effigurate to reticulate, most obvious when moist. Medulla
yellow. Photobiont Nostoc. Lower surface pale yellow to buff, distinctly wrinkled or
irregularly ridged, papillate or bullate, tomentose from margins to centre, tomentum thin to
thick, white, silky, woolly entangled. Pseudocyphellae rather sparse, scattered, minute 0-1 mm
diam. or less, on raised conical verrucae, decorticate area flat, immarginate, sunk in
tomentum. Pycnidia scattered, laminal, rather sparse, 0-05-0-15 mm diam., slightly papillate,
ostiole dark red-brown to black, punctate-depressed. Apothecia not seen.

Anatomy: Thallus 135-185(230-310) um thick. Upper cortex 25-30 um thick, upper 10-12 um
pale straw-yellow to yellow-brown, cells compressed, remainder colourless, cells 2-5-9 um
diam. Tomental hairs occasional, colourless, to 7 um diam. and to 50 um long. Photobiont layer
36-90(-130) urn thick, photobiont Nostoc, cells 4-5-7 urn diam. Medulla 34-140 um thick,
hyphae to 4-5 |im diam., thickly encrusted with yellow crystals. Lower cortex 13-5-22 urn thick,
colourless to pale straw-yellow, cells 4-5-9 urn diam. Tomental hairs colourless, septate, locules
short, constricted at septa appearing moniliform, 4-6 um diam., 45-140(-185) urn long, singly
or in fascicles.

Chemistry : As above .

Pseudocyphellaria jamesii is named in honour of Peter W. James (BM) in recognition of his
collections of, and researches into Southern Hemisphere lichens; most notably of the chemistry
and taxonomy of Menegazzia, Nephroma, and Pseudocyphellaria. He has made fine collections
of Pseudocyphellaria from New Zealand, Tasmania, and Patagonia and in 1962-63, while on
leave in Dunedin, New Zealand, he discovered photosymbiodemes of Dendriscocaulon and
Sticta filix at Lake Thompson in Fiordland National Park, a discovery which led to a later
important monograph (James & Henssen, 1976) on lichen cephalodia.

Distinguishing features: Pseudocyphellaria jamesii is a small, australasian, rosette-forming
species characterized by a yellow medulla and blue-green photobiont, and a maculate upper
surface lacking either soredia or phyllidia. It is not known fertile and it has a chemistry similar to
that of P. ardesiaca and P. nermula.

Variation: P. jamesii is a species of constant morphology, showing only a slight variation in
the width of lobes and also some variation in the extent and density of tomentum (Fig. 84B) on
the upper surface (from completely glabrous to tomentum scattered in ragged patches).
Marginal lobules are sometimes developed but their appearance is rare and spasmodic and it is
not known what ecological conditions influence their formation. P. jamesii is distinguished from
two related yellow-medulla species with which it is sometimes sympatric; from P. ardesiaca in
lacking soredia and from P. nermula in lacking phyllidia.

In several earlier accounts (Galloway & James, 1980; Arvidsson & Galloway, 1981 ; Galloway
et al., 19836; Galloway, 19856) P. jamesii was recorded as P. coerulescens , a vicariant South
American species not known from New Zealand. P. coerulescens differs from P. jamesii in
several respects: its lobes are wider and longer (0-5-1-5 cm x 5-10(-15) cm), plane or convex
and not so strongly faveolate; the upper surface is smooth, shining, here and there minutely
papillate but not tomentose (use x 10 lens); the lower surface is pale yellow and glabrous at
margins, dark grey to blackened tomentose centrally; and the pseudocyphellae are larger
(0-1-0-8 mm diam.) with conspicuous, raised margins.

Distribution: (Fig. 85) Pseudocyphellaria jamesii is a rare and local species known only from
montane to subalpine habitats in South I., east of the Main Divide from Nelson (near Lake

176

D. J. GALLOWAY

Norfolk I

Kermadec 30'
Is

Lord Howe I

Three Kings Is

The Snares

Auckland Is
\

Campbell I
v

Macquarie I

160

170 C

40

-42

-44

40-

Chatham Is

Bounty Is
Antipodes Is

46 <

166

168

50'

180

172

170

172

174 C

176

174

176

I

178

34-

42'

Fig. 85 Distribution of Pseudocyphellaria jamesii.

PSEUDOCYPHELLARIA 111

Rotoiti) to Central Otago, 200-900 m. It also occurs in South-eastern Australia where it appears
to be rare.

Habitat ecology: Pseudocyphellaria jamesii occurs in successional habitats altered by fire in the
northern part of its range, on twigs and/or bark of Leptospermum in river flats, terraces, and on
the shores of Lake Rotoiti. In Central Otago it is found on soil overlying schist rock in much
drier, open, Leptospermum-shmbland habitats. In more humid habitats it associates with the
following lichens: Anzia jamesii, Coccocarpia erythroxyli, C. palmicola, Degelia duplomargi-
nata, Erioderma sorediatum, Hypogymnia lugubris, H. mundata, H. subphysodes, Lobaria
scrobiculata, Leioderma amphibolum, L. pycnophorum, Menegazzia dielsii, M. pertransita, M.
subpertusa, Normandina pulchella, Nephroma australe, N. cellulosum, N. rufum, Pannaria
fulvescens, Pannoparmelia angustata, P. wilsonii, Pseudocyphellaria ardesiaca, P. coronata, P.
crocata, P. dissimilis, P. faveolata, P. glabra, P. maculata, P. nermula, P. pickeringii, P. rubella,
Psoroma euphyllum, P. durietzii, P. leprolomum, P. sphinctrinum, Sticta limbata, Usnea
capillacea, and U. pusilla.

Specimens examined: South Island. Nelson: Takaka Hill, /. K. Bartlett (Herb. Bartlett); St
Arnaud, Lake Rotoiti, Black Hill, 19 November 1977, D. J. Galloway (CHR); West Bay, Lake
Rotoiti, 15 November 1977, D. J. Galloway (CHR) Canterbury: Boyle River, 11 January 1979,
D. J. Galloway (CHR); September 1981, P. W. James (BM); Otago: Tarras, 14 May 1978,7. K.
Bartlett 30478 (Herb. Bartlett); Arrowtown, 3 September 1981, /. K. Bartlett (Herb. Bartlett).

31. Pseudocyphellaria knightii D . Galloway Fig. 86

in Lichenologist 15: 143 (1983). Type: New Zealand, South I., Nelson, five miles north of Westport,
on bark of Dacrycarpus dacrydioides on side of small stream, March 1980, D. J. Galloway (CHR
375745! - holotype, BM! - isotype).

Morphology: Thallus irregularly spreading, 3-10(-15) cm diam., uniformly closely attached
or loosely attached centrally, marginal lobes free, flat or ascending, corticolous. Lobes
linear-laciniate, 2-5(-12) mm wide, 10-30(-70) mm long, discrete from margins to centre, or
more often imbricate-complex, apices rounded, truncate or shallowly notched or incised,
canaliculate to flat. Margins entire to shallowly notched or incised, sinuous, often ascending
and lobes appearing canaliculate, slightly thickened, and inrolled below, sometimes beset with
short tufts of tomentum or with tomentum projecting from lower surface, rarely with scattered,
white, punctiform pseudocyphellae. Upper surface dark grey-black or bluish black, suffused
red-brown at apices when moist, pale greyish brown or fawn, suffused red-brown or cinnamon in
parts when dry, undulate, smooth to minutely wrinkled, pitted or papillate, matt, somewhat
coriaceous, without soredia, isidia or phyllidia. Maculae minute, white, effigurate, most
noticeable at lobe margins (x 10 lens) but also giving lamina a mottled appearance when moist,
larger areas of photobiont-free tissue often present as irregular, buff, pink or brown patches.
Pseudocyphellae frequent, scattered, white, 0-1 mm diam. or less, rarely to 0-2 mm diam.,
fleck-like, decorticate area flat, shallowly punctiform to very slightly raised. Medulla white.
Photobiont Nostoc. Lower surface pale buff to grey-brown, densely tomentose to margins,
tomentum brown or brown-black or red-brown, densely entangled, woolly, often projecting
beyond lobe margins. Pseudocyphellae rather sparse, often difficult to see in thick tomentum,
rarely conspicuous, raised, to 0-5 mm diam. (normally 0-1 mm) occasionally with prominent,
white decorticate area, more usually decorticate area flat, buff, brown or dirty white, very
inconspicuous. Pycnidia scattered, marginal, submarginal to laminal, solitary to 3-4-together,
hemispherical, 0- 1-0-3(0-5) mm diam., ostiole red-brown to black, punctate-depressed.
Apothecia very rare (only seen once), sessile, rounded 2-3 mm diam., plane to subconvex, or
shallowly undulate, disc pale red-brown, matt, smooth, epruinose, exciple translucent when
moist, pale flesh-coloured, restricted to a narrow, irregular margin, scabrid-areolate or
roughened below disc.

Anatomy: Thallus 130-230(-280) urn thick. Upper cortex 40-45 urn thick, upper 10-12 um
straw-yellow, cells tightly compacted, to 4-5 um diam., remainder colourless, to 9 urn diam.

178 D. J. GALLOWAY

Fig. 86 Pseudocyphellaria knightii. Holotype (CHR). Scale = 1 cm.

Photobiont layer 34-45 um thick, photobiont Nostoc. Medulla 45-90(-180) um thick. Lower
cortex 27-32 um thick, colourless, cells 6-13 |im diam. Tomental hairs colourless, 45-140 um
long, single or in sparse fascicles. Apothecia: Exciple colourless, 90-140 um thick, poorly
developed at margins of disc, cells 5-18 um diam. Hypothecium pale straw-yellow unchanged in
K, 46-75 um thick. Thecium colourless, 90-100 um tall: epithecium 11-15 um thick, pale
yellow-brown to red-brown, translucent, dissolving in K. Asci 58-82 x 13-5-15 um. Ascospores
pale yellow-brown to brown, fusiform-ellipsoid, apices pointed or one end rounded, 1-3-
septate, (25-)27-32 x 9-11-5 urn.

Chemistry: 7|3-acetoxyhopan-22-ol, hopane-7|3, 22-diol (tr.), hopane-15a, 22-diol.

Pseudocyphellaria knightii is named for Charles Knight (1808-1891), the distinguished 19th
century New Zealand lichenologist whose herbarium contains many well-annotated and beauti-
fully illustrated species of Pseudocyphellaria. He collected the genus widely, sending material of
it to Kew, Paris, Geneva, Munich, and Vienna, with selected specimens distributed in the
exsiccati of Arnold and of Lojka.

Distinguishing features: Pseudocyphellaria knightii is an endemic species having linear-
laciniate to somewhat rounded-flabellate lobes, with entire to shallowly incised margins. The
upper surface is undulate, coriaceous, smooth or slightly wrinkled, with minute, white maculae
visible (x 10 lens) near margins, and with frequent, scattered, white pseudocyphellae. It has a
white medulla, a blue-green photobiont, and a pale buffer grey-brown, thickly tomentose lower
surface with rather sparse, white pseudocyphellae. It has a two-hopane chemistry [Code A of
Wilkins & James (1979)], and is only very rarely fertile.

Variation: Pseudocyphellaria knightii is still a rather seldom collected species and such

PSEUDOCYPHELLARIA

179

Norfolk I

Lord Howe I

Three Kings Is

The Snares

Auckland Is
\

Campbell I
Macquarie I

160

170

-40

-42 C

-44

46

172

166

168

170

172 C

174 C

176

174

176
_J

178

36-

42-

-44 c

Fig. 87 Distribution of Pseudocyphellaria knightii.

180 D. J. GALLOWAY

specimens as have been examined show a rather uniform morphology. It is related to P.
lividofusca (both have the same chemistry and occasionally form photosymbiodemes) but it is
distinct from it in having a blue-green photobiont and rather thinner, more fragile lobes. It is
similar to a palaeotropical species P. semilanata, known from Fiji and Samoa, but this latter
species has a punctate-impressed upper surface, narrower lobes, and it is besides often fertile.
An Australian species, frequent in the rain forests of Queensland, P. punctillaris , is similar to P.
knightii but has a thicker, more coriaceous thallus and has distinctly tomentose-pubescent lobe
margins and apices.

Distribution: (Fig. 87) North I., from lat. 3520'S, to West of Lake Taupo, and the Kaweka
Range and in South I. , from Nelson (Kaihoka Lakes) to Westland (Lake Mahinapua), also from
the Waihopai Scenic Reserve (Invercargill), and from Stewart I., sea-level to 1410 m.

Habitat ecology: Pseudocyphellaria knightii is still very incompletely known. It is a species of
moderate to deep shade, being found on logs on the forest floor, at the bases of forest trees, and
on the bark and twigs of trees and and shrubs, and is known from the following phorophytes:
Agathis australis, Dacrycarpus dacrydioides, Dacrydium cupressinum, Dracophyllum traversii,
Hoheria glabrata, Leptospermum scoparium, and Nothofagus solandri var. cliff ortioides.

Specimens examined: 16.

32. Pseudocyphellaria lindsayi D. Galloway, sp. nov. Fig. 88

Diagnosis: Pseudocyphellariae lividofuscae similis sed pagina inferior pallidus vel bubalinus, epithecio
granulatus, K+ roseus, sporae 1-3-septatae, fuscae, 32-41 X7-9 urn. Typus: New Zealand, South I.,
Nelson, Denniston Plateau north of Westport, on Nothofagus menziesii near road, 500 m, 21 September
1981, D. J. Galloway (BM! - holotype, CHR-isotype). [t.l.c.: 15a-22-dihydroxy-24-hopanoic acid,
15a-acetoxy-22-hydroxyhopan-24-oic acid ()].

Morphology: Thallus orbicular to irregularly spreading, 5-12(-20) cm diam. , closely attached
centrally, margins free, often ascending, corticolous. Lobes linear-elongate or elongate-
flabellate, apices rounded, discrete, occasionally contiguous to centre or more often complex-
imbricate centrally, sinuses circular. Margins entire, sinuous, occasionally delicately and
shallowly notched or incised, slightly thickened below. Upper surface bright lettuce-green,
suffused brown or red-brown in parts and at margins, with occasional to frequent, minute, white
to pale yellow-brown or red-brown papillae when moist, pale grey-green to pale fawn, suffused
red-brown in parts when dry, undulate, plane or shallowly ridged or wrinkled, not faveolate,
matt or shining, smooth, coriaceous, without soredia, isidia, maculae, or phyllidia. Pseudo-
cyphellae occasional to frequent, white, conspicuous, 0-3 mm diam., generally much less, flat,
fleck-like, shallowly to deeply punctate, verruciform, margins raised, prominent. Medulla
white. Photobiont green. Lower surface smooth to wrinkled-plicate, tomentose from margins to
centre or occasionally with a narrow, glabrous, marginal zone, tomentum pale whitish buff to
chocolate brown centrally, thin, scattered, to thick woolly-entangled centrally. Pseudocyphellae
numerous, prominent, small at margins, larger centrally, round to irregular, 0-05-1-2 mm
diam., shallowly verruciform or papillate, margins distinct, concolorous with lower cortex, free
of tomentum, decorticate area flat or convex or concave, white. Pycnidia sparse to frequent,
often common in clustered groups at margins or scattered over lamina, 0-1 mm diam. or less,
slightly raised, red-brown to black. Apothecia sessile, occasional to frequent, submarginal to
laminal, rounded, 0-5-3 mm diam., concave at first then plane to shallowly convex, disc
red-brown turning dark brown or black on drying, matt, epruinose, margins crenate-stellate or
dentate, obscuring disc when young, exciple pale flesh-coloured or whitish, translucent when
moist, conspicuously corrugate-scabrid.

Anatomy: Thallus 200-300 um thick. Upper cortex pale straw-yellow to hyaline, cells 4-
13-5 um diam. Photobiont layer 35-55 um thick, cells densely packed, photobiont green, cellular
envelope prominent, 11 um diam. Medulla 110-150 um thick, colourless. Lower cortex 22-34

PSEUDOCYPHELLARIA

181

Fig. 88 Pseudocyphellaria lindsayi. Holotype (BM). A. Scale = 5 mm. B. Scale = 1 mm.

|im thick, colourless to pale straw-yellow, cells 4-5-12 ^im diam., tomental hairs colourless,
rather sparse, 45-150 um long. Apothecia: Exciple colourless, 90-180(-230) \im thick, cells 7-15
\im diam. Hypothecium 45-60 ^im thick, dilute red-brown or pale orange-brown, opaque.
Thecium colourless, 80-105 \im tall; epithecium densely conglutinate, dark red-brown, densely
granular, 11-22-5 um thick, granules not dissolving in K, turning rose pink in K; paraphyses
submoniliform at apices, granules external to tips. Asci 86-90 x 18-20 um. Ascospores brown,
fusiform, 1-3-septate, apices pointed, straight or curved, contents distinctly vacuolate, 32-41 x
7-9 um.

Chemistry: as above.

Pseudocyphellaria lindsayi is named for the Scottish cryptogamist William Lauder Lindsay
(1829-1880) who visited New Zealand in 1861 and who botanized extensively in the Otago
Province. From these collections, both Lindsay and also William Nylander published several

182

D. J. GALLOWAY

Fig. 89 Distribution of Pseudocyphellaria lindsayi.

PSEUDOCYPHELLARIA 183

important papers, including many original observations on species of Pseudocyphellaria, then
still included in Sticta.

Distinguishing features: Pseudocyphellaria lindsayi is an endemic species having linear-
elongate or elongate-flabellate lobes, with entire, rounded margins, the upper surface has
scattered, white pseudocyphellae, the medulla is white, the photobiont green, and the lower
surface is pale whitish buff to brownish with numerous, white pseudocyphellae. Apothecia are
submarginal to laminal, the disc red-brown to black and epruinose. Paraphyses are submonili-
form at apices, the epithecium densely granular, the granules external to the paraphyses tips and
dark red-brown, dissolving in K and turning rose-pink. The ascospores are fusiform, brown,
1-3-septate. The chemistry is characteristic containing 15a-acetoxy-22-hydroxyhopan-24-oic
acid () and 15ct-22-dihydroxy-24-hopanoic acid.

Variation: Pseudocyphellaria lindsayi is still a poorly collected species, the 10 specimens
examined all having a similar and rather constant morphology. The species is closely similar to
P. lividofusca and in several earlier accounts (Ronaldson & Wilkins, 1978; Galloway & James,
1980; Galloway et a/., 1983ft; Galloway, 19850) was included in the circumscription (as a
chemodeme) of this latter species. However, it differs from it in several respects: the lower
surface of P. lindsayi is pale whitish buff and not the uniform dark-brown of P. lividofusca, the
epithecium is granular, red-brown, turning rose-pink in K (that of P. lividofusca is opaque,
yellow-brown, and is unchanged in K), and the spores of P. lindsayi are longer (32-41 urn) than
those of P. lividofusca (25-29-5 um). The two species also each have a distinctive chemistry,
with P. lindsayi containing hopane acids as primary constituents (amphistictinic acid of
Ronaldson & Wilkins (1978)), whereas P. lividofusca has the normal two-hopane chemistry
[Code A of Wilkins & James (1979)]. The entire, non-phyllidiate lobe margins distinguish
P. lindsayi from P. episticta (the chemistry of the two species is the same).

Distribution: (Fig. 89) Pseudocyphellaria lindsayi appears to have a somewhat restricted
distribution in New Zealand, being confined to high-rainfall areas west of the Main Divide in
South I. , from Nelson to Fiordland, and from Mt Egmont National Park, Taranaki in North I. It
has an altitudinal range from sea-level to 990 m.

Habitat ecology: Little is known of the detailed habitat requirements, though it appears to
favour high rainfall as evidenced by its collection from Westland and Fiordland rain-forest. It is
an epiphyte of shrubs and trees being found amongst moss on bark in medium to low-light
habitats, and associating with the following species of Pseudocyphellaria: P. dissimilis, P.
episticta, P. glabra, P. homoeophylla, and P. lividofusca.

Specimens examined: North Island. Taranaki: North Egmont, along Veronica Walk, 26
November 1983, L. Tibell 15039 (UPS). South Island. Nelson: Denniston Plateau, 21 Septem-
ber 1981, D. J. Galloway (BM); sine loco (near Westport? Paparoa Range), R. Helms 90, 102
(W). Westland: Kelly's Hill, W. Martin (CHR); Otira, 17 September 1981, H. J. M. Sipman
(BM). Southland: Mt Burnett, Dusky Sound, 15 February 1969, R. M. Carter (CHR); Oak
Island, Wet Jacket Arm, Dusky Sound, 15 February 1969, G. A. M. Scott (CHR); Cook
Stream, Pickersgill Harbour, Dusky Sound, 18 February 1969, D. J. Galloway (CHR).

33. Pseudocyphellaria lividofusca (Krempelh.) D. Galloway & P. James Fig. 90

in Lichenologist 12: 300 (1980). - Sticta lividofusca Krempelh. in Verh. zool.-bot. Ges. Wien 26: 448
(1876). Type: New Zealand, sine loco (prob. Wellington), C. Knight (M! - lectotype).
Sticta amphisticta Knight in Trans. N.Z. Inst. 12: 267 (1888). Type: New Zealand, sine loco (prob.
Wellington), C. Knight (WELT - Herb. Knight Vol. 16A p. 23! - lectotype; BM! WELT - Herb.
Knight Vol. 36 p. 14! - isotypes). Fig. 90A

Morphology: Thallus orbicular to spreading, 5-10(-20) cm diam. , loosely to closely attached
centrally, margins free and ascending, corticolous. Lobes elongate-laciniate or flabellate,
discrete from apices to centre, contiguous to imbricate, subdichotomously to complexly and
irregularly branched, 0-4-1-5 cm wide, 1-5 cm long. Margins entire, rounded, shallowly

184

D. J. GALLOWAY

Fig. 90 Pseudocyphellaria lividofusca. A. Lectotype of Sticta amphisticta (WELT) Scale = 2 cm. B.
Helms 28 ( W) . Scale = 5 mm .

scalloped, notched or incised, sinuous, subascendent in parts, markedly thickened or slightly
ridged above and below, sinuses circular, prominent, often with brown tomentum projecting,
without pseudocyphellae. Upper surface bright lettuce green to glaucous green suffused

PSEUDOCYPHELLARIA 185

red-brown at apices when moist, pale greenish grey to brownish or red-brown when dry, plane or
convex, shallowly and irregularly wrinkled or ridged, matt or shining, smooth, coriaceous,
without soredia, isidia, phyllidia or maculae. Pseudocyphellae white, prominent, numerous,
0-1-1 mm diam., round to ellipsoid or linear, margins noticeably