Pseudocyphellaria Vain.
Thallus heteromerous, foliose, dorsiventral, lobate, often very large, orbicular, rosette-forming or irregularly laciniate or ± dichotomously branched or polyphyllous, 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, complex, imbricate, margins entire or variously incised orfolded often with conspicuous, elongate or verruciform pseudocyphellae, often free and ascending, tough, thick and coriaceous to thin and fragile. Upper surface smooth or wrinkled, or ± scabrid or hairy, often deeply or shallowly faveolate, with faint or marked reticulum of interconnecting ridges, shining, matt or dull, often conspicuously maculate, with or without pseudocyphellae, isidia, phyllidia or soredia. Medulla white or yellow. Photobiont green (Dictyochloropsis or Chlorella -like) or blue-green (Nostoc). Internal cephalodia containing Nostoc present in species with a green photobiont. Lower surface glabrous in some species 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 to raised-conical, round to irregular, margins distinct or indistinct. Apothecia hemiangiocarpic, emergent, sessile to pedicellate, laminal or marginal, rounded, margins entire or crenate-striate, phyllidiate, isidiate or sorediate in some species, disc matt or shining, sometimes white-pruinose, thalline exciple well-developed, smooth to verrucose or areolate-scabrid, hairy, sorediate or maculate. Ascospores 8 per ascus at maturity, colourless or brown, simple at first, becoming polaribilocular to 3-septate.
Key
Pseudocyphellaria is a genus of temperate to tropical lichens included in the family Lobariaceae and appears to be more closely related to Lobaria than it is to Sticta. Species are characterised by: pseudocyphellae on the lower surface (in some species these are also present on the upper surface or at the margins), emergent apothecia containing ascospores which are colourless or brown, 1-3-septate or polaribilocular, and a diverse chemistry consisting of orcinol deriv atives, β-orcinol derivatives, triterpenoids, terphenylquinones and pulvinic acid derivatives. [Murray J. Chem. Soc. 1952: 1345-1350 (1952); Corbett and Young J. chem. Soc. 1966c: 1556-1563, 1564-1567 (1966); Corbett and Cumming J. chem. Soc. 1971c: 955-960 (1971); Chin et al., J. chem. Soc. Perkin Trans. 1: 1437-1446 (1973); Huneck et al., J. Hattori bot. Lab. 37: 539-562 (1973); Maass, Can. J. Bot. 53: 1031-1039 (1975); Maass et al., Can. J. Chem. 55: 2839-2844 (1977); Corbett and Wilkins Aust. J. Chem. 30: 2329-2332 (1977); Wilkins Phytochem. 16: 2031-2032 (1977); Ronaldson et al., Aust. J. Chem. 31: 215-219 (1978); Goh et al., J. chem. Soc. Perkin Trans. 1 (12): 1560-1564 (1978); Renner et al., Z. Naturf. 33c: 826-830 (1979); Wilkins and James Lichenologist 11: 271-281 (1979); Goh and Wilkins J. chem. Soc. Perkin Trans. 1 (6): 1656-1658 (1979); Green et al., New Phytol. 84: 339-348 (1980); Elix and Lajide Aust. J. Chem. 34: 2005-2011 (1981); Renner and Galloway, Mycotaxon 16: 197-231 (1982); Galloway et al., Lichenologist 15: 135-145 (1983)].
New Zealand is especially rich in species of Pseudocyphellaria (in earlier accounts of New Zealand lichens the genus was referred either to Sticta, Lobaria or to Ricasolia), and is one of the two great centres of speciation, the other being southern South America. Since species are on the wholelarge and easily collected and occupy a wide range of habitat niches, they are among the most frequently gathered of New Zealand lichens and were early represented in European herbaria. W. Lauder lindsay, a Scottish doctor and lichenologist who visited the Otago settlement in 1861 wrote "...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... They are also numerous in relation to the Parmeliae, whose place they to a great extent occupy in New Zealand, especially in the forests. While in Britain the proportion of the Stictae to the Parmeliae is as 100 to 366, and throughout the world generally as 100 to 154, in New Zealand it is as 100 to 84. 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 Trans. Linn. Soc. Lond. 25: 494 (1869)].
Nineteenth century collections led to a profusion of names [J.D. Hooker and Taylor Hook. Lond. J. Bot. 3: 646-650 (1844); Babington Fl. N.Z. 2: 273-284 (1855); Nylander J. Linn. Soc. Lond. Bot. 9: 246-248 (1866); Hooker Handbk N.Z. Fl.: 566-570 (1867); Lindsay Trans. Linn. Soc. Lond. 25: 494-508 (1869); Knight J. Linn. Soc. Lond. Bot. 11: 244 (1871); Krempel-huber Verhandl. zool.-bot. Ges. Wien 26: 448-449 (1876); Knight T.N.Z.I. 12: 267 (1888); Nylander Lich. N.Z.: 29-41 (1888); Müller Argoviensis Bull. Herb. Boissier 2, App. 1: 32-37 (1894); Hellbom Bihang K. Sv. Vet.-Akad. Handl. 21 (3) 13: 29-42 (1896)], and in the present century the confusion surrounding names of New Zealand taxa persisted [Zahlbruckner Denkschr. Akad. Wiss. Wien math.-naturwiss. Kl. 104: 279-293 (1941); Allan Tuatara 4: 97-101 (1949); Martin Tuatara 17: 106-117 (1969); Martin and Child "Lichens of New Zealand", pp. 127-140 (1972); Imshaug "Adaptation within Antarctic Ecosystems", pp. 955-965 (1977)].
In New Zealand populations of Pseudocyphellaria, it is often possible to find pairs of species growing together as single joined thalli, the two species differing apparently only in the nature of the photobiont present, one part with a green photobiont, the other with a blue-green photobiont. Such pairs of species are termed photosymbiodemes [phycosymbiodemes of Renner and Galloway Mycotaxon 16: 197-231 (1982)]. The following pairs of photosymbiodemes are known at present and doubtless others will be discovered*: "P. allanii" -"P. coriacea"; "P. margaretae"-"P. pubescens"; "P. murrayi"-"P. rufovirescens", with the blue-green photosymbiodeme being listed first in each case. Until more is known of the influence of photobionts on the expression of mycobiont mor- phology in Pseudocyphellaria both component symbiodemes are given an independent species name since they are often found free-living. In additon to photosymbiodemes, there are several species-pairs represented in the New Zealand with a fertile species and a corresponding asexual species possessing either isidia or soredia but rarely or never fertile. Examples of such species-pairs are P. homoeophylla-P. delisea (isidiate) and P. faveolata-P. granulata (sorediate). It is thought that the asexual species (which is normally more widely distributed) is derived from the sexual species, and from an evolutionary point of view the asexual species is regarded as arising subsequent to the emergence of the fertile parent species. Interestingly enough in the case of P. delisea which is widely distributed in New Zealand, south eastern Australia and southern South America, the fertile parent species P. homoeophylla is known only from New Zealand.
The present account recognises 42 species in New Zealand but several undescribed species are also known and are the subject of further study. Synonyms of accepted species and their typification are in Galloway and James [ Lichenologist 12: 291-303 (1980); Galloway et al. Lichenologist 15: 135-145 (1983)].
--------------------
* Dr A.L. Wilkins and Dr T.G.A. Green reported to a meeting of A.N.Z.A.A.S. at Adelaide in May 1980, their discovery of photosymbiodemes of "P. hookeri"-"P. durietzii" and "P. knightii"-"P. lividofusca" subsequent to those photosymbiodemes discussed above.
