The Cladoniae of New Zealand
[Read before Otago Branch, October 8, 1957; received by Editor, October 15, 1957.]
Cladonia is a genus of lichens represented in New Zealand by more than seventy species. A very brief account of the morphology and ecology of the Cladoniae is followed by a short record of previous researches on New Zealand material. The place of chemistry in species diagnosis is discussed, and a criticism of the so-called “chemical species” is added. The main body of the paper is devoted to a key to the indigenous species, varieties, and commonest forms of Cladonia and to a detailed list of species with known local habitats. Mattick's system of classification, with modifications by Des Abbayes, has been followed in preference to that of Vainio.
The lichen genus Cladonia inhabits all parts of the earth, and is represented in New Zealand by possibly seventy or more species, only two of which, however, are endemic. Many of these are cosmopolitan or wide ranging in both hemispheres; some are confined to the southern hemisphere, and several are shared with Australia or with South America.
Lichens are commonly highly polymorphic, and no genus is more so than is Cladonia. Named forms are legion, but many of these have little or no taxonomic importance, having no clearly defined limits. Morphological convergence, presumably under the stimulus of environmental factors, makes the delimitation and determination of many species unusually difficult, often necessitating the aid and corroboration of microchemical tests. The modern practice of according specific status to plants morphologically inseparable but chemically distinct is referred to later. Recognition of these adds considerably to the total of indigenous species.
The present account follows some five years' study of these plants in the field and in the herbarium. Material help has been rendered by Dr. Alexander W. Evans, of Yale University, an acknowledged authority on the genus, both in the matter of species determination and in the provision of literature and specimens, which help is gratefully acknowledged.
The system of classification devised by Dr. E. Vainio, and published in his monumental “Monographia Cladoniarum Universalis” has till recently been followed by most students of Cladonia; but a revised system published in 1938 and modified in 1940 by Mattick is here followed as being more in line with the evolutionary development of the plants themselves. The subgenus Cladina has been incorporated with the Cenomyce to form the subgenus Eucladonia which now includes all New Zealand species save three comprising the subgenus Clathrina. The subgenus Eucladonia is subdivided into two sections—the Perviae and the Clausae. The Perviae are distinguished by the scyphi when present being open on perforate, while the axils between the main axis of the podetium and its branches are also commonly open, perforate, or gaping. In some species the axils may be either open or closed. In the Clausae neither axils nor scyphi are open, the latter being closed by a flat or concave membrane. All red-fruited Cladoniae belong to the Clausae. The subgenus Pycnothelia is absent from New Zealand, also the sub-section Leporinae of the Cocciferae, the series Tenues of the Cladinae, and possibly the series Macropus of the Podostelides.
General Morphology and Ecology
Cladonia as a genus is characterized by the presence of a dorsiventral primary thallus either crustose, foliose, or much more commonly squamulose in character, from the margin or surface of which one or more simple or branched, hollow podetia arise, variously terminated by saucer-shaped or goblet-shaped expansions known as scyphi, by apothecia, by spermagonia, or frequently by subulate, furcate or blunt, sterile apices. These podetia may exceed 15 cm in height, but more commonly are less than half of this.
The apothecia are turgid reproductive structures of variable size containing asci, each with eight fungal ascospores. The spermagonia or pycnidia are smaller, cylindrical or fusiform bodies containing what are believed to be non-motile male cells variously termed pycnidiospores, spermatia, or conidia, surrounded by a clear or reddish jelly-like matrix. Their true function is still not certainly known, but in any case reproduction in Cladonia is mainly vegetative, every part of the lichen being capable of initiating new individuals. Mealy or granular soredia and coralloid isidia are special structures containing both fungal hyphae and unicellular algae or gonidia, developed to ensure reproduction and distribution.
The podetial wall typically comprises three distinct layers—an outer cortex, an inner chondroid or cartilaginous layer, and an intermediate medullary layer—but any one or two of these may be absent from the whole or part of the podetium in a number of species. The cortex may be continuous or almost so; or it may be broken up into patches, into areolae, or into verruculae. The outer surface may be smooth, rugose, tomentose, squamulose, sorediose, or isidiose.
From the simple, cylindrical type of podetium an evolutionary advance leads to its upward expansion into a shallow or deep scyphus, or to a variously branched structure reaching its highest development in the Cladinae. Here the branching may be sympodial, dichotomous, or polytomous. The scyphi may be sterile, or may be margined with apothecia, spermagonia, or proliferations of various forms.
In the more primitive species, the primary thallus is generally well developed, large, and persistent, whereas the podetia are simple and short. On the other hand, the more advanced species have usually a small and often evanescent primary thallus, while the podetia are commonly tall and robust, scyphose, or much branched. The spermagonia and occasionally the apothecia in the more primitive members are usually borne on the primary squamules; but the more advanced species bear both apothecia and spermagonia on the podetia.
Most Cladoniae are highly xerophytic and grow on barren or acid soils, in heaths, or on peat, rotting logs, or decaying tussock bases. Some are frequent on roadside banks or on a thin soil veneer overlying rock. Though they occur commonly in areas both of high and of low rainfall, in general they avoid permanently saturated substrata. They are harmed neither by drought nor by frost. Most species are photophilous and none can grow in deep shade, nor do they thrive in a smoky atmosphere. Some species are mainly montane or subalpine; others are confined to the lowlands.
The greatest concentration of species or of individuals in New Zealand occurs in manuka heaths, on decaying logs on the forest margin, or on the dead bases of Danthonia tussocks margining areas of peat. As many as a score of species or even more may occur in an area twenty metres square. Podetial squamules are most strongly developed in damp areas screened from direct sunlight.
Chemistry in Cladonia Systematics
In 1866, Dr. W. Nylander published the results of experiments conducted by him on the reaction of various lichen species when a strong aqueous solution of KOH was applied to the primary thallus or podetium, pointing out that a colour change occurred in some species but not in others. He claimed that the same species
always gave the same reaction, and that such tests assisted him greatly “in not only the discrimination of many difficult and closely allied species, but in some instances in defining the affinities of genera”. (Translation.)
For a time Tuckerman challenged these conclusions, but later acknowledged that chemical tests might yield “clews to affinity”. Prof. Bruce Fink also observed, “as to chemical tests, it is extremely doubtful whether they are of diagnostic value in any lichen”. Most modern students find chemical tests of very considerable value as supplementary evidence, but opinion is strongly divided on the validity of species distinguished solely on chemical differences from plants morphologically indistinguishable.
The writer deems all plants morphologically indistinguishable as members of one and the same species even when variations occur in the products of the internal metabolism. Some twenty distinct “lichen substances” are produced by the various species of Cladonia. Some of these are invariably present in certain species; some are variously present or absent. The former may be deemed diagnostic and the latter accessory; but the practice of separating out plants that possess or lack a “lichen substance” otherwise lacking or present as the case may be in a majority of specimens of a single morphological entity, and according them specific status does not seem warranted. Rather should they be deemed strains of one common species as has already been done in the Taxaceae, Myrtaceae, and Labiatae where plants from different localities have yielded different chemical compounds, or where the same plant has yielded different products at different seasons of the year.
The absence of a “lichen substance” normally present may be due entirely to environmental factors as when usnic acid fails to form in the absence of good sunlight; or it may be due to an errant gene, or to some other undiscovered cause. The fact remains that “species” segregated purely on chemical grounds have almost invariably the same geographic range as the parent species which is strong presumptive evidence that a single species is involved. The distinction between “accessory” and “diagnostic” lichen substances is often somewhat arbitrary, for it is the removal of the non-conformist plants that leaves any substance diagnostic for the balance. Where, however, a particular substance is known in but a single species (e.g., divaricatic acid in C. sullivani) then that substance can be regarded as truly diagnostic.
Notwithstanding what has been said above, the writer finds it expedient to give tentative recognition to the so-called chemical species in the flora, leaving to someone with a wider experience to make any relevant changes. The reagents most in use for species determination or verification are caustic potash (KOH), chloride of lime (CaCl2O2) and p-phenylene-diamine usually indicated by the letters K, C, and P followed by a + or − sign according as a colour change results or fails to occur. A faint reaction is indicated by + f. (KC) indicates that the application of K is immediately followed by C. For microchemical tests for the various lichen substances see Asahina (1).
Short History of Previous Researches on New Zealand Cladoniae
Many of the earlier overseas botanists to visit New Zealand–Hooker, Raoul, Cunningham, Lyall, Lindsay, Berggren and others—included Cladoniae in their collections; while such resident botanists as Hochstetter, Haast, Sinclair, Colenso, Knight and Helms forwarded many gatherings to overseas specialists including Nylander, Arnold, Müller, Krempelhuber, Mitten and Leighton.
Hooker's collections went to Kew where they were, in Hooker's own words, “hastily named and published by Dr. Taylor”. Later these were reviewed by Babington, whose determinations were subsequently confirmed by Nylander, the leading lichenologist of the period. The first enumeration of New Zealand Cladoniae comprised a list of 16 species determined by Babington and recorded in Hooker's “Flora
Novae Zelandiae” (1855). Twelve years later Hooker, in the “Handbook of the New Zealand Flora”, published a revised list of 14 species. Dr. Lauder Lindsay spent three months in Otago in 1861–62 and subsequently (1866 and 1868) published a list of Cladoniae found by him in the environs of Dunedin.
These early botanists interpreted many species of Cladonia more widely than later specialists. All enumerated C. rangiferina and C. pycnoclada as New Zealand indigens; but, in the more restricted interpretation of the present day, neither species is now listed as occurring in this country. Lindsay (35, p. 533) even remarks, “I cannot distinguish it (i.e., C. pycnoclada) by any sufficient characters from the condition of C. rangiferina called alpestris”. Today, at least ten segregates are recognised as valid species. C. fimbriata has similarly been broken up into C. fimbriata (in restricted sense), C. Balfourii, C. Borbonica, C. coniocraea, C. cornutoradiata, C. cylindrica, C, major, C. ochrochlora, and C. nemoxyna, all present in New Zealand.
In 1865 Nylander published a paper on Lauder Lindsay's Otago collections, and in 1888 his “Lichenes Novae Zelandiae” appeared. Krempelhuber in 1876 published some new Cladoniae for New Zealand (32) as did Müller in 1892, 1894, and 1895, based largely on collections forwarded by Charles Knight and William Colenso.
During the first thirty years of the present century little or nothing was done save by Dr. E. du Rietz who, in 1926–27, made very extensive collections of New Zealand lichens, including some Cladoniae. In 1931, however, the late J. Scott-Thomson began a ten-year period of systematic collecting of the lichen flora of the South Island which included some 135 gatherings of Cladoniae. Duplicate specimens were forwarded to Dr. H. H. Allan, then Director of the Botany Division of the D.S.I.R., who in turn sent many specimens to overseas specialists for determination. The Cladoniae were examined by F. W. Gray and Dr. H. Sandstede and the latter's determinations appeared in Zahlbruckner's Lichenes Novae Zelandiae (62) in 1941. About the same period K. W. Allison collected Cladoniae in the Rotorua-Taupo area, of which several were new to New Zealand.
A short review of the Cladoniae of New Zealand by Dr. H. H. Allan was published in the 1937 report of the twenty-third meeting of A.N.Z.A.A.S. (1) in which 46 species comprising the known Cladonia flora of New Zealand were listed.
During the decades following 1940 the writer, while engaged in bryological field work, made a point of collecting associated lichens. For the past five years, only the genus Cladonia was systematically collected mainly in the South Island, some 800 gatherings resulting. On a study of these and of the plants in the herbaria of J. Scott-Thomson and of the D.S.I.R. the present paper is mainly based. At the present time Dr. James Murray, of Otago University, is engaged on systematic collecting and study of the indigenous lichen flora, and his help and collaboration have been much appreciated. The writer has also had great assistance from Dr. Alex. W. Evans, of Yale University, in the matter both of literature and of determination of specimens as well as in the provision of authentic material. Dr. W. Culberson has also without solicitation sent me valuable comparative material from U.S.A. It should be added that not all material has yet been determined, some being possibly new to New Zealand.
K = KOH in strong aqueous solution (KOH 1 part, H2O 5 parts).
C = CaO2Cl2 in saturated aqueous solution.
P = p-phenylene-diamine in alcoholic solution (P 1 part, Abs. Alcohol 10 parts), (faint reactions are indicated by + f, positive colour reactions by +, negative reactions by −).
H.H.A. = Dr. H. H. Allan; K.W.A. = K. W. Allison; J.S.T. = J. Scott-Thomson; T.W.A. = T. W. Attwood; J.M. = Dr. Jas. Murray; W.M. = W. Martin.
Cladoniae determined by Dr. A. W. Evans are indicated by an asterisk.
Previous records for Cladonia endoxantha, C. mitrula and C. japonica are here attributed to C. hypoxantha, C. capitata and C. carassensis respectively, of which the former are synonyms.
Records based on collections made by J. Scott-Thomson, K. W. Allison, and T. W. Attwood are listed on the basis of determinations by Dr. H. H. Allan and Dr. H. Sandstede respectively unless otherwise indicated. These plants are housed either at Otago University or in the D.S.I.R. herbarium in Christchurch.
Plants not attributed to named forms or varieties are not necessarily “type” forms, but include young plants or undetermined forms in addition, as well as early records lacking form names.
Species in square brackets have been recorded from New Zealand, but their presence lacks later confirmation.
Many named forms have small taxonomic value, being but stages in a more or less continuous series. Many forms of C. aggregata are of this character.
In the literature the distinction between varieties, forms, and modifications is often so vague that a plant diagnosed as a “form” by one author is listed as a variety by another, and even as a species by a third often without change of citation.
The writer considers that speciation in Cladonia has been greatly overdone and that a large reduction in species will eventually take place, especially by the elimination of those based solely on chemical distinctions.
In a genus so polymorphic as Cladonia where convergence in forms of two species is often very close, the preparation of a reliable key presents unusual difficulties. Final determinations should, therefore, rarely if ever be made with out reference to full diagnostic descriptions.
The subsection Cladniae in New Zealand requires complete overhaul in the light of recent researches.
Summary of Localities
As many of the localities listed do not appear on other than the most detailed maps, they are here located in their provincial districts.
Southland: Awarua, Bluff, Forest Hill, Fortrose, Greenhills, Kuriwao, Longwood Range, Lake Manapouri, Riverton, Tautuku, Wilderness.
Eastern Otago and Dunedin: Boyd's Bush, Flagstaff, Horse Range, Kakanui Mountains, Lamb Hill, Maungatua, Mihiwaka, Mt. Cargill, Otakou, Mt. Pisgah, Saddle Hill, Swampy Hill, Waitati, Mt. Watkins, Whare Flat, Verterburn.
West Otago: Cascade Creek, Cleddau Valley, Hollyford Valley, Homer Tunnel, Key Summit, Lake Howden.
Canterbury: Akaroa, Arthur's Pass, Lyttelton, Mt. Cook.
Marlborough: Dillon's Hill, Hundalee, Oaro, Puhipuhi Reserve, Picton, Wairau, Valley, Waihopai Valley.
Nelson: Aniseed Valley, Botanical Hill, Kaiteriteri, Pelorus Bridge, Lead Hills, Maruia, Rai Valley, Tophouse, Takaka Hill, Mt. Trovatore.
Westland: Greymouth, Hokitika, Kanieri, Kelly's Hill, Ross, Runanga.
I desire to acknowledge the material assistance rendered by Dr. Alex. W. Evans in determination of species, in supply of literature, and for specimens. Over 140 determinations, accompanied by notes on chemical reactions and lists of acids present have entailed much time and labour, for which I am indeed grateful. To Mr. T. Thomson I am also much indebted for permission to examine his father's lichen herbarium which contains duplicates of many plants determined by Sandstede, and also for permission to record the habitats of the Cladoniae present. Through the kindness and co-operation of Mr. C. M. Smith, the present Director, I have been privileged to examine the collections of New Zealand Cladoniae housed in the herbarium of the Botany Division of the D.S.I.R. Many of these had been determined by the late Dr. H. H. Allan, the former Director, who had an unrivalled knowledge of the lichen flora of this Dominion. For helpful criticism and for assistance in the preparation of the key I am much indebted to Dr. Jas. Murray, and for material assistance in securing the necessary literature my thanks are due both to Dr. Murray and to Mr. P. Havard-Williams, Librarian to the University of Otago.
|1. Podetia corticate or ecorticate, squamulose or esquamulose, often scyphiferous; commonly sorediose; inner cartilaginous layer seldom absent; walls imperforate or perforations few||Sub-genus Eucladonia (2–61).|
|Podetia always corticate, esorediose, esquamulose, and ascyphous; inner wall without cartilaginous lining; podetial walls with numerous and striking perforations, except in some forms of Cladonia aggregata||Sub-genus Clathrina (62–63).|
|2. Scyphi when present closed by a flat or concave membrane; axils never perforate or gaping; apothecia scarlet, pale, or brown||Section Clausae (3–41).|
|Scyphi when present open, or partially closed by a perforate or annular membrane; axils gaping, perforate, or closed; apothecia never scarlet||Section Perviae (42–61).|
|3. A. Lower surface of primary squamules cream, yellowish, or stramineous, sometimes white.|
|(a) Primary squamules medium to large, broad, persistent, bearing the spermagonia when present; podetia absent, or if present simple, short, occasionally scyphose. Apothecia pale||Subsection Foliosae (4).|
|(b) Primary squamules small to medium; podetia longer; ± stramineous; apothecia pale||Sub-section Ochroleucae (5).|
|B. Lower surface of primary squamules usually white, often tinged brown at base|
|(c) Apothecia always scarlet, or black by degeneration||Sub-section Cocciferae (6–15).|
|(d) Apothecia light or dark brown, or blackish; podetia thick-walled, usually short; central canal narrow; usually ascyphous and fertile; rarely sorediose; usually grey or brown||Sub-section Podostelides (16–22).|
|(e) Apothecia brown; podetia thin walled; central canal wide; short or long; simple or branched; with or without scyphi; frequently sterile; frequently sorediose||Sub-section Thallostelides (23–41).|
|4. Primary squamules large, crowded, ± ascending, cream or white beneath; laciniae linear, often fringed with rhizines. K −, (KG) + yellow||C. foliceae var. alcicornis.|
|5. Primary squamules 3–12 mm; podetia 6–40 mm, yellowish or straw-coloured; cortex basal or none; decorticate and farinose wholly, or above only; scyphose; scyphi regular or oblique, narrow or wide. K + f||C. carneola.|
|6. Primary squamules and podetia glaucescent or greenish grey; never yellowish and rarely scyphose Primary squamules and podetia tinged with yellow; usually scyphose||Series Stramineo-flavidae (14–15).|
|7. Podetia K+,P+||8.|
|Podetia K −, P −||10|
|8. Primary squamules elongate, glaucescent above; under surface with yellowish or brownish median bands margined with cream, large and intricately branched. Podetia 3–7 mm tall, corticate. Some times scyphose||C. hyporxantha.|
|Primary squamules chalky white beneath, small, often darkening towards the base. Podetia ascyphous||9.|
|9. Podetia simple or sparingly branched at apex, 10–25 mm tall, slender; mainly decorticate; cartilaginous layer often exposed, whitish or brown, and translucent. Soredia, squamules, and coarser granules often present||C. vulcanica.|
|Podetia generally rather robust, simple or sparsely branched, apices obtuse; often corticate at base and apex; soredia few or none; cartilaginous layer not exposed; esquamulose as a rule||C. macilenta.|
|10. Podetia usually under 30 mm tall, white, simple, slender, densely powdery sorediose; mainly decorticate, squamules none or basal, apothecia single, large but rare||C. bacillaris.|
|Podetia as in C. vulcantca; small patches of cortex often present, often decorticate and pellucid||C. didyma.|
|Podetia 6–50 mm tall but rarely over 30 mm; mainly corticate or areolate-corticate in patches; surface rough, soredia none, or few and granular; squamules present or absent; simple or sparingly branched, apothecia numerous, small||C. Floerkeana.|
|Podetia 10–35 mm, white, glaucous, or stramineous; simple or dichotomously branched; mainly decortirate, impellucid, and verrucose||C. oceanica.|
|11. Scyphi regular, ± irregular, deformed, or absent; podetia commonly over 30 mm tall. K + (except C. deformis)||12|
|Scyphi regular; plants usually less than 30 mm tall, K − (except C. digitata)||15|
|12. Podetia usually strongly verrucose-isidiose near base. K +,P+||13|
|Podetia not or not strongly verrucose near base||14|
|13. Podetia stout, mainly granulose-sorediose, sometimes squamulose, often branched near the apex. (KC) —||C. subdigitata.|
|Podetia slender, squamose or squamulose but not sorediose; rarely branched; (KC) +||C. metalepta.|
|14. Podetia scyphose, or escyphose and turgid; scyphi regular, deformed or absent; yellowish; esquamulose; farinose-sorediose wholly or upper part only. (KC) + lutescent||C. deformis.|
|Podetia slender, 10–50 mm tall; white, glaucous, or ashy, granulose-sorediose; base often corticate or verrucose; scyphi narrow and often coronate or proliferous; K + yellow, (KC) + fuscous red, P + red||C. flagelliformis|
|15. Podetia robust, large, scyphose, esorediose, verrucose, or areolate-verrucose above||C. pleurota.|
|Podetia scyphose but smaller; corticate at base, sorediose above. K −, P −||C. pleurota.|
|Podetia scyphose or ascyphous; corticate in lower part; farinose-sorediose above; scyphi simple or proliferate, narrow, margins ± incurved; K +, P +||C. digitata|
|16. Spermagonia when present mainly on primary thallus. Podetia usually short, terminated with apothecia wider than the podetia, and constricted at base||Series Helopodium (17–21).|
|Spermagonia usually on the podetia; podetia usually taller; with or without apothecia; apothecia not conspicuously wider than the podetia||Series Macropus (22).|
|(Note.—The only species of this series recorded from New Zealand is C. decorticata; which in any case seems inappropriately placed in Macropus. The record requires confirmation)|
|17. Podetia 12–15 mm tall, solid, branched above, central canal absent or basal only. Cortex ± verrucose||C. solida.|
|Podetia arising from apex or margins of primary squamules; central canal present but narrow; often exposed||18.|
|18. Podetial walls ± channelled or fluted but rarely fissured; inner walls not exposed and not or sparingly lacerate||19|
|Podetial walls split, exposing inner wall, which is fibrose or lacerate. All podetial branches bear apothecia as a rule||20|
|19. Primary squamules 3–17 mm long. Podetia present or absent, 5–15 mm tall, not or sparingly branched, areolate-corticate, soredia and squamules rare. K + yellow then red, P —||C. subcariosa.|
|Primary squamuies 1–4 mm. Podetia 3–10 mm, usually simple; cortex ± continuous or areolate, or scattered, often ± verrucose; sometimes sorediose K −, P +||C. capitata.|
|20. Primary squamules medium to large, 5–20 mm; some podetia often sterile, usually squamulose; irregularly branched K +, P + (yellow)||C. neozelandica.|
|Primary squamules 1–6 mm long; all podetial branches fertile; podetial squamules few or none||21.|
|21. Podetia simple or fastigiately branched, 3–12 mm tall, squamules few or none, corticate, cortex areolate verrucose; esorediose. K − or faintly + P + (red)||C. enaptia.|
|Podetia usually branched, 5–30 mm tall, squamules basal or none, cortex ± continuous, areolate ± verrucose; cartilaginous layer often exposed, fissured and torn. K + (yellow) P + (pale yellow ± golden)||C. cariosa.|
|22. Podetia ± granulose and squamose at base, squamulose above, often sorediose, decorticate areas chalky white; simple or branched; 9–40 mm tall; cartilaginous layer not much torn||[C. decorticata.]|
|23. Cortex ± continuous or areolate. Podetia not sorediose, commonly scyphose|
|Cortex ± continuous, in patches, or basal only; areolate or verrucose; podetia always ± sorediose and largely decorticate||32.|
|24. Primary squamules 1–4 mm of variable shape, finely incised; podetia 4–50 mm tall, scyphose, tubaeform. or ascyphous; scyphi not wider than 3 mm: sterile apices blunt or subulate. K −, P +||C. pityrea.|
|Primary squamules small, or medium to large; margins crenate, lobate, or digitately incised||25.|
|25. Scyphi 1–7 mm wide, regular and deep, or tubaeform, cavity always ± corticate' cortex smooth or verrucose; podetia often closely clustered; sometimes decorticate above and coarsely granular||C. pyxidata.|
|Scyphi shallow and regular, or irregular, or absent||26|
|26. Podetia ascyphous and slender, or scyphose and proliferous, corticate and esorediose; not blackening near base||27.|
|Scyphi cristate, irregular, or absent; cortex areolate, breaking up near the base into detached white flakes on dark, subtomentose under-layer||31.|
|27. Scyphose or ascyphose; scyphi large or small, regular. proliferating from cup-margins||C. gracilis.|
|Scyphi shallow, proliferating from cup centres, or more rarely from margins or even podetial walls as well, ranks few on many, each rank scyphose||28.|
|28. K −, P +||29.|
|K +, P +||30.|
|29. Scyphi ± gradually expanded; margins subentire or dentate; apothecia usually sessile; cortex 20–30μ thick; ranks 3–7; primary squamules few, small and narrow||C. verticillata.|
|Scyphi flaring more abruptly from slender base; margins thinner and more deeply incised; apothecia usually ± stipitate; cortex 10–20μ; ranks 3–7, primary squamules few and small||C. calycantha.|
|Scyphi similar but short and more irregular, 5–15 mm tall, sometimes squamulose; ranks 1–3, primary squamules large, ascending, caespitose||C. cervicornis.|
|30. Primary squamules small. Podetia repeatedly scyphose; ranks 3–7; K + (faint yellow)||C Krempelhuberi.|
|Primary squamules large; ranks 1–2, rarely 3; K + (strong yellow)||C subcervicornis.|
|(Note.—The last three species are sometimes listed as varieties of C. verticillata.)|
|31. Subcortical layer blackening near base and overlaid with white flecks of cortex.|
|(a) Primary squamules 3–12 mm long but usually rather small; cups usually irregular and often squamulose; proliferations central or marginal. Podetia 10–50 mm, K − or f, P +||C. degenerans.|
|(b) Primary squamules large, crenate; basal areoles often bearing large squamules; podetia short; scyphi difform or cristate. K +, P +||[C. lepidota.]|
|32. Scyphi regular and deep; verrucose or areolate near base. P +||33.|
|Scyphi small, irregular, obsolete, or absent. P +||35.|
|33. Podetia always scyphose, corticate or coarsely granular near base: ± decorticate and granular sorediose above; impellucid: cup cavitv granular or verrucose.|
|(a) K−, C −, P +||C. chlorophaea.|
|(b) K −, C −, P −||C. Grayi.|
|(c) K −. C +. P ±, (KC) + (purplish)||C. cryptochlorophaea.|
|(d) K + (red), C + (red), P ±, (KC) + (purplish) C. cryptochlorophaea.|
|Podetia corticate to cup base, never verrucose; smooth and areolate; soredia farinose; primary squamules robust; K −, P +||C. conista|
|Podetia mainly decorticate and densely farinose-sorediose; scyphi regular, slender, mouth narrow K − or f, P +||34.|
|34. Podetia short, scyphose, rather slender, under 20 mm tall, simple or proliferate; cartilaginous layer rarely well defined fimbriatic acid present||C.fimbriata.|
|Podetia usually taller, slender, but firmer; scyphi regular, often corticate to half height; fimbriatic acid absent||C. major.|
|35. Podetia mainly granular-sorediose; decorticate areas ± pellucid; scyphi rare; cups corticate within Car|
|tilaginous layer well defined. Primary squamules small, slender, finely incised, evanescent. K − or + f.||C. pityrea.|
|Soredia farinose; decorticate areas opaque or semipellucid||36|
|36. Lower part of podetia isidiose and sometimes squamulose also; 5–30 mm tall, scyphi present or absent, very narrow, shallow or deep; decorticate areas K + (pale yellow). Primary squamules broad, coarse, medium to large||C. Borbonica.|
|Podetia white, simple, 5–10 mm tall, mainly ecorticate and farinose sorediate; apex blunt or almost scyphose; base with coarse granules or minute squamules but scarcely isidiose; primary squamules 1 mm or less. Apothecia rare but large or small||C. cylindrica.|
|Lower part of podetia lacking isidia or large granules (except C. Balfouru)||37.|
|37. Podetia with continuous or areolate cortex almost to top, then sorediose; brown at base; 50–100 mm tall, cylindrical, subulate||C. cornuta.|
|Podetia wholly or mainly decorticate||38|
|38. Podetia usually taller than 25 mm||39.|
|Podetia usually shorter than 25 mm||39|
|39. Scyphi rare, regular and repeatedly proliferous. Podetia more commonly simple or sparingly branched above, 30–65 mm tall, ascyphous; mainly decorticate and farinose-sorediose. P +||C. cornutoradiata.|
|Scyphi narrow and shallow, abortive, or absent; podetia elongate, 25–90 mm tall; often areolate-corticate at base, granular-sorediose above, branched. P ±||C. nemoxyna.|
|40. Podetia decorticate save at base, densely sorediose, isidiose, ascyphous, unbranched; translucent cartilaginous layer often exposed; 5–25 mm tall.||C. Balfourii.|
|Podetia shortly corticate at base and apex, farinose-sorediate between, opaque||41|
|41. Corticate areas very short. Scyphi rare, small, narrow, or abortive; sorediose inside cups. Podetia usually simple, with truncate apices centrally depressed; 5–30 mm tall. Primary squamules rather large||C. coniocraea.|
|Corticate areas longer, with patches of cortex in sore diose area between; scyphi when present corticate within cup; podetia 5–40 mm tall; usually branched; apothecia usually yellowish-brown||C. ochrochlora.|
|42. Primary thallus squamulose, often persistent. Podetia white, green, glaucous, or brownish; not intricately branched; axils open or closed; scyphi when present never closed unless partially by an annular membrane||Subsection Chasmarae (43–51)|
|Primary thallus squamulose, soon disappearing. Podetia yellowish, white, greenish or stramineous, usually corticate, scyphi and squamules rare; axils open or closed; plants much branched, dying at base||Subsection Unciales (52–55).|
|Primary thallus crustaceous, evanescent. Podetia ecorticate, ascyphous and esquamulose, ± tomentose; densely and intricately branched, forming dense or open white, yellowish, or glaucous-green cushions||Subsection Cladinae (56–61)|
|43. Primary squamules medium to large, broad. K +, P +||Series Megaphyllae (44).|
|Primary squamules small, often narrow, evanescent||Series Microphyllae (45–51).|
|44. Podetia 20–75 mm long, stout, branching, sometimes obsoletely scyphose; cortex ± areolate; esorediose; walls often perforate or fissured. P ±, C +||C. turgida|
|Podetia 10–15 mm long, rather slender, usually un-branched or cornute; scyphi present or absent; granulose-furfuraceous; C –.||C. rigida.|
|45. Podetia smooth, cortex ± continuous; never sore diose||46.|
|Podetia sorediose, granulate, or ecorticate in patches||49.|
|46.K +, P + Podetia simple or shortly branched||47|
|K −, P ± Podetia rarely simple, branches long or short||48|
|47. Cortex continuous, smooth, branching by irregular dichotomies, axils open or closed; scyphi rare in New Zealand, opening gradually or abruptly. P + red||C. carassensis.|
|Cortex of close or scattered areoles; podetia often turning blue-black near base; brownish or white above; often fissured; 15–40 mm tall; axils open. P + yellow||C Aueri.|
|48. Axils, and scyphi when present, gaping and crispate-dentate, rarely squamulose; podetia 10–75 mm tall P||C. crispata.|
|Axils open but neither gaping nor crispate; podetia 15–85 mm tall, usually squamulose; P + red||C. furcata.|
|49. Podetia partly corticate, or decorticate and farinose-sorediose; often scyphose and proliferate, walls and apices often perforate; K −, P −||[C. cenotea.]|
|Podetia esorediose, or granular sorediose||50|
|50. Podetia elongate, 25–80 mm, branches rather long and widespread; cortex ± continuous but broken; usually squamulose; often sorediose or minutely squamulose near apex. K −, P +||G. scabriuscula.|
|Similar to G. scabriuscula but densely farinose-sorediate and without accompanying squamules.||G. farinacea (Vain) Evans.|
|Podetia long or more often short, subcorticate, areolate or verrucose; squamulose; simple or irregularly branched; axils usually open||51|
|51. Podetia ecorticate or sparsely areolate-corticate, or cortex subcontmuous at base and squamulose; scyphi present or absent, often proliferate; primary squamules 1–7 mm long, often persisting, K −, P −||G. squamosa.|
|Podetia similar but more granular or squamulose as a rule; cortex areolate or verrucose; K+, P +||C. subsquamosa|
|52. Podetia simple or sparingly branched, usually crowded and erect; cortex subcontinuous; areoles greenish; axils closed, or open near the base||C. capitellata.|
|Podetia much branched, but less complex than in the Cladinae, esquamulose; cortex smooth and glossy, (ecorticate in C. Boryi)||53.|
|53. Axils rarely perforate. Spermagonial jelly white||54.|
|Axils usually open; podetial walls often fissured;|
|spermagonial contents red||55|
|(Note.—The spermagonial jelly cannot be satisfactorily observed in either very young or in old spermagonia.)|
|54. Scyphi absent; branching dichotomous; cortex sub-continuous; axils closed; podetia yellowish stramineous, slender and smooth. P + (yellow)||[G. medusina.]|
|Scyphi sometimes present, cristate. P —||C. amaurocraea.|
|55. Podetia corticate, cortex ± continuous and areolate; neither scyphose nor squamulose; brittle when dry; axillary and internodal perforations conspicuous in older plants; KC− or f||C. uncialis.|
|Podetia ecorticate; cartilaginous layer absent or reticulate; sometimes obsoletely scyphose; walls pitted or perforate; (KG) +||C. Boryi.|
|56. Podetia robust; axis sympodial; branching polytom-ous; apical branchlets unilaterally deflexed||Series Rangiferinae (57–58).|
|Podetia less robust; axis sympodial near base only; branching polytomous, mainly trichotomous; branches subequal; ultimate branchlets straight or spreading, or deflexed in several directions||Series Impexae. (59–60).|
|Podetia with branches dichotomous and subequal; ultimate branchlets straight and spreading||Series Bicornutae. (61).|
|57. Ultimate branchlets deflexed in same direction||58|
|Ultimate branchlets divaricate or deflexed in several directions; P −||59|
|58. P +||G. sylvatica.|
|59. Sympodia obvious; axils open; ultimate branches forming a globose head; spermagonial contents red||G. alpestris|
|Branch tips deflexed on all sides. Spermagonial jelly white||60|
|60. Branching ± sympodial near base, mainly trichotom-ous above; branches subequal, usually grey-green; old podetia ± pellucid||C. impexa.|
|Sympodia more evident; branching polytomous; colour yellowish; axils open or closed||C. leptoclada.|
|61. Branching mainly dichotomous; axils usually per forate, colour yellowish or whitish; K −, P + (yellow)||[C. pycnoclada.]|
|(Note.—Santesson says C. pycnoclada is restricted to South America. The numerous records of f. flavida and f. exalbescens of this species from New Zealand require critical study in the light of recent research.)|
|62. Podetia coralloid, reticulate; white, grey, or rarely brown in New Zealand.||C. retipora.|
|Podetia stramineous, brown, or nigricant|
|63. Perforations in podetial walls numerous, revealing black lining of interior walls; sterile apices obtuse||C. Sullivanii.|
|Perforations many or few; interior walls never black; sterile apices subulate||C. aggregata.|
Key to Varieties and Forms
|Cladonia didyma (Fèe) Vainio|
|Podetia strongly squamose||f. squamulosa Robbins.|
|A. Podetia areolate-corticate near the apex||var. rugifera Vain.|
|B. Podetia decorticate||var. muscigena (Eschw.) Vain.|
|1. Branched||f. polydactyloides (Müll Arg.) Vain.|
|2. Simple and very short||f. pygmaea Vain.|
|Cladonia digitata Schaer. Podetia scyphose, sorediate near apex, esquamulose||f. monstrosa (Ach.) Vain.|
|Podetia scyphose, sorediate near apex, squamulose||f. phyllophora Anzi|
|Podetia without scyphi, moderately stout||f. cervchoides Vain.|
|Podetia without scyphi, slender||f. tenella Th. Fr.|
|Cladonia flabelliformis (Flk.) Vainio Podetia simple, tubaeform or scyphose||f. tubaeformis (Mudd.) Vain.|
|Podetia repeatedly proliferous and scyphose||var. polydactyla (Flk.) Vain.|
|Cladonia Floerkeana (Fr.) Sommerf. Podetia corticate, no soredia or squamuls||f. chloroides (Flk.) Vain.|
|Podetia corticate, sorediose, no squamules||f. intermedia Hepp.|
|Podetia moderately squamulose||f. carcata (Ach.) Nyl.|
|Cladonia macilenta (Hoffm.) Nyl. Podetia obtuse, farinose-sorediate, esquamulose, spermagonia on primary thallus||f. styracella (Ach.) Vain.|
|Podetia sorediate, squamulose, spermagonia on primary thallus if present||f. squamigera Vain.|
|Podetia corticate, soredia sparse||var. corticata Vain.|
|Cladonia coccifera (L). Willd.|
|Podetia without squamules, apothecia large, globose||var. stemmatina Arch.|
|Podetia squamose||f. phyllocoma Flk.|
|Cladonia subdigitata (Nyl.) Vain. Podetia proliferous, much digitately branched above||f. polydactyloides Nyl.|
|Cladonia pleurota (Flk.) Schaer. Podetia without squamules, robust, fertile, apothecia on robust marginal innovations. K —||f. extensa (Ach.) Sandst.|
|Podetia with wide, open cups, and conspicuous large marginal apothecia. K —||f. decorata Vainio|
|Podetia. K +||f. albida Vainio|
|Podetia squamulose. K —||f. phyllocoma (Flk.) Sandst.|
|Proliferations central.||f. centralis (Schaer) Oliv.|
|Cladonia deformis Hoffm. Podetia ascyphose, tapering to blunt points||f. cornuta Fors.|
|Podetia scyphose, not proliferous||f. cyathiformts Sandst.|
|Podetia scyphose, margins lacerate prolifei-ous||f. extensa (Hoffm.) Vain.|
|Cladonia Balfourii Cromb.|
|Podetia scyphose but cups gradually expanded to not more than 2 mm in width||f. chlorophaeoides (Vain.) Evans.|
|Cladonia Borbonica (Del.) Nyl. Scyphi absent or narrow. Podetia squamulose||f. squamulosa Robbins|
|Cladonia calycantha Del. Margins of scyphi squamulose||f. foliolosa Vain.|
|Cladonia chlorophaea (Flk.) Spreng. Podetia sterile, esquamulose, or few basal squamules. Scyphi without proliferations. Decoiticate surface of cups smooth and opaque||f. simplex (Hoffm.) Arn.|
|Decoiticate surface of cups ribbed, semi pellucid||f. costata (Flk.) Arn.|
|Scyphi proliferate, and proliferations mostly scyphose||f. prolifera (Wallr.) Arn.|
|Podetia fertile, esquamulose, cups distinct||f. carpophora (Flk.) Anders.|
|Cladonia comocraea (Flk.) Sandst. Podetia subulate, tips minutely centrally depressed||f. ceratodes (Flk.) Dalla Torre and Sarnth.|
|Podetia truncate or narrowly scyphose, tips conspicuously centrally depressed||f. truncata (Flk.) Dalla Torre and Sarnth.|
|Podetia squamose or squamulose||f. phyllostrota (Flk.) Vain.|
|Podetia scyphose, cups well expanded and proliferate||f. stenoscypha (Stuckenb) Sandst.|
|Cladonia cornuta (L.) Schaer. No scyphi or podetial squamules||f. cylindrica Schaer.|
|Scyphose||f. scyphosa Schaer.|
|Podential squamules present||f. phyllotoca (Flk.) Arn.|
|Cladonia cornutoradiata (Coem.) Sandst. Podetia elongate, branches few or none, not scyphose, almost wholly decorticate and sorediose||f. subulata (L.) Sandst.|
|Podetia elongate, scyphose, proliferous, wholly decorticate and sorediose||f. radiata (Schreb.) Coem|
|Podetia branched but not scyphose||f. furcellata Sandst|
|Podetia squamose or esquamulose||f. capreolata (Flk.) Flot.|
|Scyphose, repeatedly proliferous from margin of lower scyphi, scyphi large and regular||f. repetitoprolifera Sandst.|
|Cladonia degenerans (Flk.) Spreng. Podetia scyphose, squamules few or none.||f. repitoproifera Sandst.|
|Scyphi regular but sterile||f. euphorea (Ach.) Flot.|
|Scyphi irregular, with lacerate margins||f cladomorpha (Ach.) Vain.|
|Podetia ascyphous, irregularly branched, branches cornute or subulate||f. dilacerata Schaer.|
|Podetia with or without scyphi, squamulose||f. phyllophora (Ehrh.) Flot.|
|Primary thallus curling into loose balls which roll freely in the wind||f. erratica Lindsay|
|Cladoma gracilis (L) Willd Scyphi regular, sterile, proliferous, no squamules or soredia||var dilatata (Hoffm) Vain|
|Scyphi often irregular, podetia often squamulose||f. dilacerata Flk.|
|Scyphi if present minute. Podetia very slender||var gracillima Norrl,|
|Scyphi regular, deformed, or none Podetia rather slender and esquamulose. K −||var chordalis (Flk) Schaer.|
|Scyphi ± regular, narrow. Podetia stout, yellowish-biown. esquamulose K + (brown) atronorine absent.||var. elongata (Jacq.) Flk|
|Similar to last but glaucescent K + (strong yellow), atronorine present (Often treated as a valid species)||f ecmocyna (Ach) Vain|
|Podetia sparingly granulose, variegated||var Campbelliana Vain|
|Cladonia pyxidata (L) Hoffm Primary squamules adnate appiessed to substratum||val pocillum (Ach) Flot|
|Primary squamules free, ascending, podetia esquamulose||var neglecta (Flk.) Mass.|
|Podetia not proliferating; cups sterile or apothecia sterile||f simplex (Ach) Harm|
|Podetia proliferate from outer surface, these scyphose||f peutheta (Walh) Robbins|
|Podetia squamulose||f lophvra Ach|
|Cladonia pityrea (Flk.) Vainio Podetia rarely over 2.5 cm Scyphi present or absent||vai Zwackhu Vain|
|Podetia scyphose Podetia sorediose, squamules few or none||f cladomorpha Flk|
|Podetia sorediose, squamulose or isidiosquamulose||f hololepis (Flk) Vain|
|Podetia not sorediose, podetial squamules few or none||f scyphifera (Del.) Vain|
|Podetia ascyphous Squamules few or none, podetia sorediose||f subacuta Vain|
|Squamulose: podetia not sorediose||f phyllophora (Mudd.) Vain.|
|Podetia 3–6.5 cm high, ascyphous or scarcely scyphose, areolate-corticate near base, ± granulose-decorticate above||var subareolata Vain|
|Cladonia verticillata (Hoffm.) Schaer. Primary squamule. small. K−. Podetia usually esquamulose, proliferations central||var evoluta Th Fr|
|Podetia somewhat squamulose||f phyllocephala Flot.|
|Proliferations both central and marginal||f apoticta (Ach.) Vain.|
|C. furcata (Huds.) Schrad|
|Podetia without squamules or soredia||var racemosa (Hoffm) Flk.|
|Sterile and subulate||f furcatosubulata (Hoffm) Vain.|
|Fertile and subcorymbose||f corymbosa (Ach) Nyl|
|Podetia squamulose||var pinnata Flk|
|Sterile podetia subulate, fertile ± corymbose||f foltolosa Del|
|C. scabriuscula (Del.) Leight. Podetia whitish or glaucescent. Cortex subcontinuous or dispersed. Podetia|
|isidio-squamulose below, granular above. Sterile apices subulate. Repeatedly but sparingly dichotomous||f. surrecta (Flk.) Sandst.|
|Cortex as above. Podetia rather stout. Apex obtuse, becoming isidio-squamulose, and granular or verrucose||f. cancellata Mull. Arg.|
|Podetia mainly decorticate, but squamulose, and isidiose, granular, or verrucose||f. adspersa Flk.|
|Podetia dark, or pale brown. Mainly decorticate and squamulose||f. conspersa Vain.|
|(Forms sublevis Sandst and semipellucida Sandst are also recorded from New Zealand. See 52, p. 40.)|
|C. squomosa (Scop.) Hoffm. Podetia cup-forming. Podetia decorticate in upper part, sparingly granulose or esorediose. Podetia naked or finely but sparingly squamulose||f. denticollis (Hoffm.) Flk.|
|Podetia densely squamulose||f. squamosissima Flk.|
|Podetia corticate, white or ashy, squamose||f. phyllocoma (Rabenh.) Vain.|
|Podetia granulose, scyphose||f. gracilenta Mull. Arg.|
|Cladonia Boryi Tuck. Podetia blunt, scarcely scyphose||f. lacunosa (Bory) Tuck.|
|Podeti with ± distinct scyphi||f. reticulata (Russell) Merrill.|
|Cladonia capitellata (Tayl.) Bab Podetia aseyphous, primary thallus ± per-sistent. Podetial axils not dilated||f. fastigiata Vain.|
|Podetial axils dilated and branching||finterhtascents (Nyl.) Vain.|
|Podetia ascyphous, blackish at base with white spots, primary thallus soon vanishing||f. degenerata Vain.|
|Podetia scyphose or frequently so. Primary thallus ± persistent||f. amaurocraeiodes Vain.|
Genus Cladonia (Hill.)Vaini.
Subgenus 1. Eucladonia (Eschw.) Matttick.
To this subgenus belong all New Zealand species save three comprising the Clathrinae.
Section 1.Clausae Koerb.
Axils never perforate or gaping. Scyphi when present closed by a membrane.
Subsection 1. Cocciferae Del.
Apothecia scarlet, or blackish by degeneration.
Series 1. Subglaucescentes Vainio.
Podetia grey, white, glaucescent, or ash coloured, never yellowish.
Cladonia bacillaris (Ach.)Nyl.K —, C —,P—.
This species has the same chemical reactions as C. Floerkeana and C. didyma but is distinguished by the copious soredia into which the cortex breaks up. New Zealand (Jolliffe); South Island (Haast); Otago (Hector); Greymouth (Helms*, W.M. 2158), Mt.Cargill (J.S.T.571); Mihiwaka (J. S. T. 645); Saddle Hill (W. M. 5111); Maungatua (J. M.0541); Lake Howden (J.M. 0852); Greenhills (W.M. 3061).
Cladonia Floerkeana (Fr.) Sommerf.K —, C —, P —.
On earth, in heaths, on rotting wood, and in moss.
South Island (Lyall); New Zealand (Colenso); Swanson (T. C. Chambers in “Tane”, 1952.
f.carcata (Ach.) Vainio.
Wellington Botanic Gardens* (W.M. 2809); Lake Manapouri (W.M.5011); Leith Saddle (W.M 4391); Kuriwao (W.M.4308).
Wellington Botanic Gardens (W.M.* 2808,* 2827); Pohangina R. (H.H.A.); Lake Manapouri (W.M. 4623*, 5011); Kaiteriteri* (W.M. 3042); Wairau Valley (W. M.3037); Mt.Cargill (W.M. 4405); Dalefield (W.M. 4016); Lake Kanieri* (W.M. 4011).
Cladonia didyma (Fée) Vainio. K −, C −, P −.
Not very common in New Zealand; a plant of earth, peat, rock, dead timber. Kuriwao (W.M. 4416); Lake Kanieri (W.M. 4004); Forest Hill (J.M. 0905).
f.squamulosa Robbins. Feilding (H.H.A.).
var rugifera Vainio. Ohakune (T.W.A).
var. muscigena (Eschw.) Vainio. Greenhills (W.M.3061).
f.polydactyloides (Müll. Arg.) Vainio Wairau Valley (W.M. 4290).
Cladonia vulcanica Zoll. and Mor. K +, C−, P +.
A segregate from C. didyma occurring in similar habitats and having the same world distribution.
Whangarei (W.A. given in Herb D.S.I R.); Taupo (K.W.A. in Herb. D. S. I.R.); Lead Hills (J. S. T. 1973); Rangitoto Island (L.B. Moore in Herb. D. S. I. R.).
Cladonia oceanica Vain. K −, P −.
Near Atiamuri (Volcanic Plateau) K.W.A.; det. Dr. H. Sandstede.(L.32 in author's herbarium.)
Cladonia macilenta (Hoffm.) Nyl. K +, C −, P +.
A plant mainly of heath soils, decaying logs, and tussock stumps.North Island (Hooker, Cunningham, Colenso, Sinclair); Kaipara (W.L L.); Whangamoa Hill (W.M.3091); Lake Kanieri (W.M. 4011); Mt. Cargill (W.M.4422); Lake Manapouri (W.M. 4626); Longwood Summit (W.M 4538); Kuriwao (W. M.3092); Mihiwaka (J.S.T 624); Cascade Creek (W.M. 4593); Awarua (W M. 4530).
f.corticata Vain. Ruapehu (J. Jablonszky).
f. styiacella (Ach.) Vain. On peat near Westport* (W.M. 1338).
f.squamigera Vain Mt. Cargill (W. M.4415).
Cladonia digitata Schaer. K +, C −, (KG) +, P +.
On peaty and barren heath soils and decaying logs.
North Island (Cunningham, Colenso).
f.monstrosa (Ach.) Vain.* Kuriwao (W.M.2170);* Cascade Creek (W. M. 4497), Maungatua (J. M.0423).
f.phyllophora Anzi.Flagstaff (W. M. 5355, 6900).
Cladonia flabelliformis (Flk.) Vain. K +, (KC) + (rufous), P +.
[Sandstede (52, p. 26) lists this species from New Zealand as a synonym of C. polydactyla Flk.]
On tree stumps and decaying logs.
f.tubaeformis (Mudd) Vain.Peel Forest, Canterbury (H.H.A.); Tophouse (W.M.3028).
Cladonia hypoxantha Tuck. K +, P +.
[Evans (17, pp. 264–267) gives reasons for regarding C. endoxantha as a synonym Sandstede (52, p. 30) gives reactions for C. hypoxantha as K −, P −.]
On logs and caudices of tree-ferns. Podetia with isidioid granules.
On Dicksonia fibrosa, Raetihi (J. E. Attwood); Mihiwaka (J.S. T. 624); Cascade Creek (W.M 5138); Lake Howden track (J.M. 0247).
Series 2. Stramineo-Flavidae
Podetia yellowish or stramineous, never glaucescent; usually scyphose.
Cladonia coccifera (L.) Willd. K — or f., (KC) +, P —.
On heath soils and on tree stumps. Early records almost certainly included C. pleurota, then deemed a variety. Not common.
New Zealand (Babington, Hooker, Krempelhuber); North Island (Colenso); Lake Taupo (K. W. A); Mihiwaka (J.S.T 651); Mt. Peel (H.H.A.); Sandy Point Reserve, Southland (G. C. Martin)
f.phyllocoma Flk.Waipuna Bush, Napier (det. Sandstede).
Cladonia pleurota (Flk.) Schaer. K —, (KC) +, P —.
On peaty and heath soils, old logs, tussock bases, etc.Abundant. Here zeorine replaces the barbatic acid present in C. coccifera.
Ohakune (T. W. A); Mt Peel (H. H. A.); Maungatua (J.S. T. 8, 131, 1830); Verterburn (J. S. T. 348), Mihiwaka (J. S. T. 628); Stewart Island (W.M. 870); Lake Kanieri* (W. M.4008); Arthurs Pass (W.M.10); Lake Manapouri (W.M. 4622); Cascade Creek* (W. M.4519), Kuriwao* (W.M. 2137); Longwood Range (W.M. 4566). Greenhills (W.M.4347); Tophouse* (W. M.3015); Dillon's Hill* (W.M. 3010).
f.centralis. Kaiangaroa Plains (K.W.A.L.170)
f. extensa (Ach.) Sandst. Lake Manapouri* (W. M.4630).
f albida Vain.(K +).Pohangina River on tree ferns (H.H.A.); Bull Mound, Tararua Mts.(V. Zotov in Herb D. S.I.R.)
Cadonia deformis (L.) Hoff. K —, C +, P —.
On peaty upland soils, tussock bases, decaying logs, grassland. All present records are from South Island. Most of the following are f. cyathiformis.
New Zealand (Haast in 29); Mihiwaka (J.S.T.632, 673); Maungatua (J. S. T. 1352, 1833); Awarua (J.ST. 1659*; W.M.4516); Kuriwao (W.M. 4815); Swampy Hill (J. S. T. 1878): Longwood Range (W.M.4475, 4461); Flagstaff (W.M.6031): Cascade Creek (W. M.4541); Lake Manapouri (W. M.4625); Mt. Cargill (W.M.4405)
Cladonia subdigitata (Nyl.)Vain.K +.C —, P +.
Amongst tussocks or on their decaying bases. Usually montane or subalpine. Wainuiomata (H. H. A): Maungatua (J.S. T. 1832); Verterburn (J S.T. 349); Mihiwaka (J. S. T. 628). Mt.Pisgah (J. S. T. 1454, 1456, 1490): Mt Watkin (J S.T.): Lake Manapouri (W. M.4628); Longwood Range (W.M.4569); Kuriwao (W.M. 4288)
f.polydactyloides Nyl.Mt.Pisgah (J. S. T. 1450).
Cladonia metalepta Nyl K + f or —, C —, (KC) +, P—.
On earth and margins of dry peat bogs.Unlike typical plants from Cuba.
Mt.Hauhangatahi (H H.A.), Mt. Pisgah (J. S. T.); Homer Tunnel (W. M.4425)?
Subsection 2.Foliosat. Bagl.et Carest
Primary squamules large and laciniate, persistent, under surface ± stramineous Podetia simple, with or without scvphi. ± stramineous, never sorediose. often absent Apothecia brownish or pale.
Cladonia foliacea (Huds.) Schaer. var. alcicornis (Light f.) Schaer K —, (KC) +, C +, P + (yellow then red).
On moist banks. Primary squamules unusually small in the sole New Zealand collection to date. Podetia not seen. Plants forming dense colonies of squamules, yellowish to brownish on lower surface.
Greymouth (W. McKay) recorded by Zahlbruckner (62); Longwood Range (W.M. 4543); Strongman Mine, Greymouth (W.M. 7542).
Subsection 3. Ochroleucae Fr.
Primary squamules small to medium, under surface ± stramineous.Podetia usually simple, with or without scyphi, yellowish or stramineous. Apothecia pale.
Cladonia carneola Fr. K +, (KC) +, P —.
On bogs and in grassland. Known at present only from Otago and Southland.Maungatua (J.S. T. 1345); Fortrose (J.S. T. 1648).
Subsection 4. Podostelides (Wallr.) Vain.
Under surface of primary squamules white. Podetia glaucescent or brownish, esorediose, never scyphose, usually fertile. Apothecia brown.
Series 1. Helopodium (Ach.) Vain.
Spermagonia on primary squamules, rarely on podetia. Podetia almost always fertile, never sorediose.
Cladonia capitata (Michx.)Spreng. K + f.(greenish yellow turning brown), P +.
Heath soils, grasslands, and decaying logs.
Atiamuri (K.W A.); Rotorua (K.W.A); Cascade Creek (W.M. 4610, 5023, 5032); Lake Kanieri (W.M. 4005).
var. abbreviata Vain. Mt. Cargill (W.M.6444).
Cladonia cariosa (Ach.) Spreng. K +, C —, P + (red).
Heath soils and tussock grassland.
Mt. Cargill (J.S. T. 572; W.M. 3026); Flagstaff (W.M. 5350); New Zealand (J. D. Hooker 28: p. 297); Taieri Mouth(J.M. as W.M.6537); Hundalee (W. M.5611); Longwood Range (W.M. 4481a); Waitati (Hellbom).
f. minor Rabenh. New Zealand (Müll. Arg. 41, p. 24).
Cladonia subcariosa (Nyl.) Vain. K + (yellow then red),P + (yellow).
Allan (1) expresses the view that early records of C. cariosa and of C. subcariosa probably refer to C. neo-zelandica and C. enantia. The writer also finds difficulty in distinguishing C. cariosa and C. neo-zetandica, especially owing to the great variation in the shape and size of the primary squamules.
New Zealand (Vainio); Taupo(K.W.A. L.117 would appear to be this); Little Barrier Island (leg. J. N. Dingley, det W. M).
Cladonia enantia Nyl.K — (or f), C —, P +. Endemic.
Moist clay banks and soils, manuka heaths, in low bracken fern. Hellbom (31) says this hardly differs from C. cariosa.
Auckland (Knight); Waitakerei Hills (H. H. A); Greatford (H H.A.); Rangitikei (H.H.A.); Atiamuri (K.W.A.); Alexandra (leg. G. C. Martin); Saddle Hill (W.M 5102). (South Island records with reserve.)
Cladonia neozelandica Vain. K +, P +. Endemic.
In similar habitats to C. enantia. Not previously recorded from South Island. Scarcely differs from C. cariosa.
Auckland (Knight); Waitakerei Hills (H.H.A.); Greatford (H.H A.); Wainui-o-mata (H.H.A.); Taupo (K.W.A.); Kaiteriteri (W.M. 3057); Mt. Cargill (W.M. 4389), Trotter's Gorge (W.M. 6514); Saddle Hill (W.M 5100); Manapouri (W.M. 4513, 4584); Wairau Valley (W.M. 3045); Maungatua (J.M. 01115); Anawhata (H.H A); Wanaka (W.M 6516).
Cladonia solida Vain. K +, P +.
A Brazilian lichen determined from New Zealand by Sandstede.
Atiamuri(K.W.A.) recorded by Zahlbruckner (62). A Dunedin specimen (J. M.0929) may belong here.
Series 2 Macropus Vainio.
Under-surface of primary squamules white.Spermagonia confined to podetia Podetia sterile or fertile, soredia present or absent.
Cladonia decorticata (Flk.) Spreng. K —, P —.
This species was listed on the basis of a collection by Colenso. Hellbom (32) also records it from New Zealand.
Subsection Thallostelides Vainio.
Under surface of primary squamules white, sometimes tinged brown near the base, squamules usually small to medium. Podetia white, olive, brownish or glaucescent; commonly scyphose, often sterile and then subulate, cornute, or blunt with centrally depressed tips Apothecia brown.Spermagonia usually on the podetia or scyphi.
Cladonia Balfourii Cromb. K —, C —, P +.
Mainly on sandy or humous soils, or on soil overlying rock.
Mt. Cargill* (W.M. 825, 829); Greenhills (W.M. 4323); New Zealand (Knight).
f.chlorophaeoides (Vain.) Evans.New Zealand (Knight, Travers) (58, Vol 2, p. 337).
This is a scyphose form resembling a small C. chlorophaea.
Cladonia fimbriata (L.) Fr. K —, C —, P +.
On earth, logs, and dead tussock bases. Early records may sometimes refer to some of the species now segregated from C. fimbriata.
North Island (Colenso, Lindsay); South Island (Raoul); Lamb Hill (J.S.T. 293); Veterburn (J.S.T. 340), Black Head (J.S.T. 387); Mihiwaka (J.S.T. 672, 914, W.M 531, 541); Maungatua (J.S T. 1837); East Taieri (W.L.L.); Stewart Island (L.Cockayne); Whare Flat* (W.M 4407); Kuriwao (W.M. 5125); Roxburgh (W.M.4705), Nelson* (W.M.3041, 5026); Waihopai Valley* (W.M. 3009); Hundalee* (W. M.3052).
var.chondroidea Vain.North Island (Colenso);Waiheki Island (H.H.A.); Tiritea (H. H. A).
Probably synonymous with C. Balfourii f.chlorophaeoides (16, p. 304).
f. subprolifera Vain. Waikaremoana (J. Jablonszky).
f.chlorophaeoides Vain. Banks Peninsula (Raoul, Travers). Probably also a synonym of C. Balfourii f. chlorophaeoides.
f.radiata (Schreb.)Coem North Island (Colenso, Jolliffe).
f.subradiata Vain.Banks Peninsula (Raoul)
Cladonia major (Vain.) Zopf. K —, P +.
Grasslands, open ground, decaying logs.
Maussgatua (J. S. T.). Otakou Bush (J.S T. 612); Goat Island (J.S. T.1057); Mt.Pisgah (J.S. T.1487. 1492); Flagstaff (J.S T. 1720); Kakanui Mts.(J.S. T.1437); Leith Valley (J. S. T. 1746)
Cladonia cornutoradiata (Coem.) Sandst. K —, P +.
On peaty, clay and heath soils; and on decaying logs. Very abundant.
New Zealand (Krempelhuber (32) p. 447); Jolliffe, Milne, Haast, Leighton.
Waitakerei Hills (H.H.A.); Taupo (K.W.A.); Fortrose (J.S.T. 1645); Otakou Bush (J.S.T 579); Tautuku (J.S.T. 1665); Maungatua (J.S.T. 1829); Cascade Creek (W. M.4417); Greenhills (W.M. 4329); Awarua (W.M. 4529).
f. albissima Sandst.Verterburn (J.S T. 336).
f. tortuosa Del. Boyd's Bush, North Taieri (J.S.T.).
f. subulata (L.)Sandst Much the commonest form in New Zealand.
Feilding (H.H.A.); Wellington Botanic Gardens* (W. M.2815); Verterburn (J S.T. 337); Awarua* (W.M. 4523); Greenhills* (W.M. 1757, 1768); Kaiteriteri (W.M. 3054); Kuriwao* (W.M. 2130); Maungatua (W.M 949); Mihiwaka* (W.M. 830); Mt. Cargill* (W.M. 4401); Saddle Hill (leg. Miss D. L. Martin; W.M. 5112).
f. furcellata Sandst. Te Aroha(62) Zahlbruckner; Kuriwao (W.M. 5345); Awarua (W.M. 4441); Trotter's Gorge (W.M. 4801).
f. capreolata (Flk.) Flot. Mt.Cargill (W.M. 4379)?
f.radiata (Schreb.) Coem. Greenhills (W.M. 4312, 4326).
f. repititoprolifera Sandst. Greenhills* (W.M.* 1751, 1760, 538); Awarua (W. M.4448).
Cladonia nemoxyna (Ach.) Nyl. K—, P ±, usually —.
Allan (1, p. 338) observes that previous records probably belong for the most part to C. subnemoxyna, but it would appear this species has never been adequately published.
New Zealand (Knight in Herb. Arn.); Wairau Valley* (W.M. 3046, 5039); Cascade Creek (W.M. 4548).
Cladonia subnemoxyna Sandst.K—, P +.
Recorded by Sandstede in Zahlbruckner (62) but without diagnosis. Taupo (K.W.A.).
Cladonia coniocraea (Flk.) Sandst. K — (or dingy brown), P +.
On earth, or more commonly on logs and in moss. Tiritea (G. Chamberlain).
f. stenoscypha (Stuckenb) Sandst. Boyd's Bush, N. Taieri (J.S.T. 469); Green Island Bush (J.S.T. 422); Saddle Hill (W.M. 5132).
f. ceratodes (Flk.) Dalla Torre et Sarnth. Boyd's Bush (J.S.T. 496); Otakou (J.S.T 614); Pirongia Mt. recorded Zahlbruckner (62); Onamalutu (W.M. 4039a); Ohakune (J.E. Attwood).
f. truncata (Flk.) Dalla Torre et Sarnth. Saddle Hill (W.M. 5122).
Cladonia ochrochlora Flk. K — or faintly +, C —, P +.
On old wood or on mossy rocks in shady places. Very similar to C. coniocraea but somewhat larger, with larger corticate areas at base and apex and inside the scyphus.
Greymouth (Helms in Nylander 46, p. 17.)
f. flexuosa (Flk.) Sandst. Palmerston North (62).
f. phyllostrota Flk. Palmerston North (Zahlbruckner 62); Tophouse (W.M. 4056?).Waikaremoana (J. Jablonszky).
Cladonia Borbonica (Del) Nyl. K + f, P +.
On earth, logs and rocks.
New Zealand (Travers in Paris Museum Herbarium); Knight in Krempelhuber (32 p. 447); Waitakerei Dam (H.H.A.); Anawhata (H.H.A.); National Park (H.H.A.); Otakou Bush (J.S.T. 500); Mt. Cargill (J.S.T. 573); Bluff (J.S.T. 834); Flagstaff (J.S.T. 1722); Mt. Cargill (W.M. 4427, 5030); Riverton Rocks.
(W.M. 4703); Tophouse (W.M. 4029); Greenhills (W.M. 5029); Westport (W.M. 2161a).
Cladonia cylindrica (Evans) Evans. K —, P +.
(Synonym = C. Borbonica f. cylindrica Evans; see Evans (11)).
Ohakune (T.W.A.) on rotten logs; Otakou Bush (J.S.T. 588); Sandy Point (W.M. 6535).
Cladonia pityrea (Flk.) Vainio. K—, C —, P +.
A polymorphous species of earth, rocks, decaying wood, peat, and moss.
Stuart Mts. (W. Thomson); Flagstaff (J.S.T. 1115, 1387); Mt. Cargill (W.M. 4428); Otago (Hector); Greymouth (Helms); New Zealand (Müller J.).
f. phyllophora (Mudd.) Vain. Lee Stream (J.S.T. 220).
f. scyphifera (Del.) Vain. Fortrose (J.S.T. 1647).
f. sorediosa Vain. Feilding (H.H.A.).
f. cladomorpha Flk. Greenhills (W.M. 4338).
f. subacuta Vain. Green Island Bush (J.S.T. 434); Mihiwaka (J.S.T. 652); Riverton (J.S.T. 775); Catlins (W.M. 835).
f. subareolata Vain. Waiheki Island (H.H.A.); Taupo (K.W.A.); Mihiwaka (J.S.T. 641); Goat Island (J.S T. 1059).
Cladonia cornuta (L.) Schaer. Younger parts K + f; KC +; P +.
On earth, tussock grassland, bogs in upland areas.
New Zealand (Travers) in Herb. Paris Museum; Mercury Bay (Jolliffe); Mt. Pisgah (J.S.T 1482); Mt. Cargill (J.S.T. 574); Flagstaff (J.S.T. 1386). Manapouri (W.M. 4484).
f. phyllotoca (Flk.) Vain. Te Aroha, 2,500ft (62).
f. cylindrica Schaer. Mt. Cargill* (W.M. 826); Manapouri* (W.M. 4484); Cascade Creek (W.M. 4510,* 4551).
f. scyphosa Schaer. Kuriwao* (W.M. 2173).
Cladonia chlorophaea (Flk.) Spreng. K —, P +.
Similar to C. pyxidata but sorediose. The following three species are segregates based on the presence in each of differing “lichen substances”.
Greymouth (Helms, in 46, p. 17); New Zealand (Travers); Abbot's Hill (J S. T. 920); Bluff (W. M.4076).
f. simplex (Hoffm.) Arn Maungatua* (W.M. 924, 948), Mt. Cargill* (W.M. 828).
Bluff* (W.M. 4076).
f. prolifera (Wall.) Arn. Mt. Cargill* (W.M. 828a); Awarua* (W.M. 4532); Maungatua* (W.M. 950).
f. costata (Flk.) Arn. Flagstaff (J.S.T. 1701); Greymouth (W. McKay).
f. carpophora (Flk.) Anders. Takaka Hill* (W.M. 1207).
Claaonia cryptochlorophaea Asahina. K —, P +.
On earth, logs, or rocks.
Arthur's Pass* (W. M.524).
Cladonia Grayi Merrill. K—, P— (rarely + f.)
On earth, logs and rocks. Contains grayanic acid.
Flagstaff (J. S. T. 1724); Verterburn (J. S. T. 345, 346); Pohangina R. (H.H A).
f. squamulosa Sandst. Near Lake Taupo (K.W.A.).
Cladonia merochlorophaea Asahina. K—, KC +, P +.
Only distinguished from C. chlorophaea by presence of merochlorophaeic acid.
Greenhills* (W.M. 1751. 1764); Dillon's Hill* (W.M. 3011); Mt. Cargill* (W. M.827); Lake Kanieri* (W.M. 4006).
Cladonia conista (Ach.) Robbins f. simplex Robbins. K —, P +, H +.
This is another segregate from C. chorophaea distinguished by the presence of “substance H” of undetermined constitution; but is also morphologically distinct. Mt. Cargill* (W.M. 4414); New Zealand (Nylander in Lichenes Exotici 1892, p. 17).
Cladonia degenerans (Flk.) Spreng. K — (almost), P +.
On earth, rock and tree bases.
New Zealand (Colenso, Krempelhuber (29) p. 447, Nylander (36) p. 18); Saddle Hill (W.L.L.).
f. euphorea (Ach.) Flk. East Taieri (W.L.L. (31) p. 533).
f. aplotea Ach. East Taieri (W.L.L. (31) p. 533).
f. phyllophora (Ehrh.) Flot. Lee Stream (J.S.T. 219).
f. erratica Lindsay Saddle Hill on limestone (W.L.L.).
Cladonia lepidota Nyl. K +, C—, KC +, P +.
On barren soil in shade of tall manuka heath. Dunedin (W. Nylander). East Taieri and Saddle Hill (W.L.L.) as a form of C. degenerans.
Cladonia pyxidata (L.) Fr. K—, C —, P +.
On open ground, on banks, and on thin soils overlying rock.
Whare Flat (J.S.T. 136); Hindon (J.S.T. 248); Saddle Hill (W.L.L.); Dunedin (Travers); North Island (Colenso); South Island (Lyall).
f. neglecta (Flk.) Mass. Dunedin (Nylander 46, p. 17); New Zealand (Travers in Paris Museum).
Cladonia gracilis (L.) Willd. K ± (yellow turning brown), P +.
On earth in grass or moss, in manuka heath, on rotting logs, peat, and tussock bases.
New Zealand (Colenso, Knight); South Island (D'Urville).
var. dilitata (Hoffm.) Vainio. New Zealand (Knight); Arthurs Pass* (W.M. 8); Awarua (J.S.T.).
f. dilacerata (Flk.) Vainio. *Cascade Creek (W.M. 4522).
var. chordalis (Flk.) Schaer. New Zealand (Travers); Palmerston North (H.H.A.); Riverton (J.S.T. 782); Kaiteriteri (W.M. *3020, *3021, 4161); Wairau Valley (W.M. 4134, 4160); Tophouse (4055); Lake Manapouri (W.M. 4504); Cascade Creek (W.M. *4384, 4434, 4484); Taupo (K.W.A.; L. 210 det. W.M.).
var. Campbelliana Vainio. (?) Peel Forest (H.H.A.). Determined as C. pityrea by Du Rietz.; Campbell Island (H. Sandstede); New Zealand (H.S. 52, p. 64).
var. gracillima Norrl. Taupo, Rangitoto, Ruakura, and Stewart Island (Zahlbruckner (48); Chelsea (H.A.A.); Mt. Matthews (A. D. Beddie); Atiamuri (K.W.A.)*; Lake Manapouri (W. M. 4483); Taieri Mouth (W.M. 2570a)*; Greenhills (W.M. 1750); Cascade Creek (W.M. 4303, 4704).
var. elongata (Jacq.) Flk. Mt. Pisgah (J.S.T.); Kuriwao (W.M. 2168); Cascade Creek (W.M. 4510).
[f. ecmocyna (Nyl.) Vain.] (K +). New Zealand (Colenso); record requires confirmation.
[Cladonia subchordalis Evans]. K—, P —.
See Notes by Evans (18, pp. 133–135). Atiamuri (K.W.A.)?
Cladonia calycantha Del. K—, C —, P +.
Manuka heaths and clay soils. Scarcely distinguishable from C. verticillata.
Palmerston North (H.H.A.): Ohakune (J. E. Attwood); Goat Island (J.S.T. 1060): Wairoa (G. O. K. Sainsbury); Ruakura (H.H.A.); Saddle Hill (W. M.5104).
f. foliolosa Vain. Tokomairiro Mouth (J.S.T. 2297); Saddle Hill (W.M. 5105, 5113); Manapouri* (W.M. 4990).
Cladonia verticillata (Hoffm.) Schaer. K—, C—, P +.
A common species on all types of substratum.
North Island (J.D.H); Moehau (L. Cranwell); Flagstaff (J.S.T. 2049); Mt. Cargill (J.S T. 576. W.M. 4412); Glenledi (J.S.T. 452); Whare Flat (J.S.T. 499); Greenhills* (W.M. 1752); Horse Range (W.M. 4812); Saddle Hill (D. L. Martin); Cascade Creek (W. M. 5340). Ohakune (J. E. Attwood); Ruakura (H.H.A.).
var. evoluta Th. Fr. Kuriwao* (W.M. 2143, 2151, 2830); Mt. Cargill (W.M. 4377); Greenhills (W.M. 4344); Cascade Creek (W.M. 5342); Flagstaff (W.M. 4135).
f. apoticta (Ach.) Vain. Cascade Creek (W. M.4714). Mt Cargill (W.M. 4438); Dillon's Hill (W.M. 4367).
f. phyllophora Flk. Mt. Cargill (J. S. T. 576, W.M. 4350, 4446); Greenhills (W. M.4313, 4335); Cascade Creek (W.M. 4549a); Ruapehu (J. Jablonszky).
Cladonia cervicornis Schaer. K—, P +
New Zealand (J.D.H., W. L.L.), Porters Pass (Hellbom); Awarua (W.M. 4459, 4460); Kaiteriteri (W. M. 3048); Leith Saddle (W.M. 4396); Rai Valley (W.M. 5344); Cascade Creek (W.M 4719).
Cladonia subcervicornis Vain. K +, P +.
Kaikorai Hill (J.S.T. 897); Longwood Range (W.M. 4465); Kuriwao (W.M. 4120); Tophouse (W.M 7410); Lake Wanaka (W. M. 6551).
Cladonia Krempelhuberi Vain. K +, P +.
Manuka heath soils. Occurs elsewhere in Japan and in California.
Anawhata (H.H.A); New Zealand (Knight) in Herb. Arn.; Krempelhuber (32). Taupo in scrubland (K.W.A.). (See Sandstede 52, p. 70).
[Cladonia subgracilis Sandst]. No Latin diagnosis, but recorded by Sandstede as a spec. nov. from New Zealand. The type has since been destroyed (18).
Section 2. Perviae (Fr.) Mattick.
Primary thallus crustose or squamulose, well developed in the more primitive species, minute or absent in the more advanced. Podetia simple, scyphose, or branched, branching sometimes complicate; often dying at base but growth continuing at the apex. Axils typically open, but sometimes closed. Scyphi open or partially closed by a perforate or lacerate membiane.
Subsection 1. Chasmariae (Ach.) Flot.
Primary squamules well developed and persistent, or small and evanescent. Axils open or closed. Podetia simple or branched. Cups present or absent.
Series 1. Microphyllae Vain.
Primary squamules small and short lived.
Cladonia Aueri Rasän. K +, P + (yellow)
Subalpine bogs and grassland. A Chilian species known in New Zealand only from Otago.
Maungatua (J.S.T. 2028; J.M.: leg. A. L. Mark, W. M.7022); Longwood Range (W.M. 4485).
[Cladonia cenotea (Ach.) Schaer.] K —, P —.
New Zealand (Nylander). The record requires confirmation.
Cladonia carassensis Vain. K +, KC +, P + (yellow).
C. japonica is a synonym. (See Evans 11: 101–104). Grows in moss, on soil overlying rock, and on peaty soil; morphologically similar to C. crispata.
Mt. Watkins (J.S.T. 2065); Awarua (W.M. 4445); Mt. Cargill (W.M. 5108); Flagstaff (W. M. 5353); Horse Range (W.M. 4807); Cascade Creek (W.M. 4514a); Kuriwao (W.M. 4105); Flagstaff (W.M. 6044).
Cladonia squamosa (Scop.) Hoffm. K —, P —.
On earth, soil veneer verlying rocks, in moss, and on decaying logs.
North Island (Colenso); Mercury Bay (W.L.L.); Lamb Hill (J.S.T. 251); Cascade Creek (W.M. 5031); Wairau Valley (W. M. 3022).
f. denticollis (Hoffm.) Flk. Otakou Bush (J.S.T. 601); Cascade Creek (W.M. 6531).
f. densata Nyl. North Island (Müll. Arg.); Greymouth (Helms).
f. gracilenta Müll. Arg. New Zealand (Jelinck; Hochstetter).
f. minor Hoffm. New Zealand (Hochstetter).
[var. asperella Flk.] North Island (Colenso, 1580). Not since confirmed.
Cladonia subsquamosa Nyl. K +, P + (orange).
On thin soils overlying rocks.
Lamb Hill (J.S.T. 251a); Longwood Range (W.M. 4476); Otago only (H.H.A. 1./2338).
Flagstaff (W.M 6039).
Cladonia chondrotypa Vain. K +, P +.
A specimen from New Zealand forwarded to Dr. H. Sandstede by Dr. H. H. Allan was so named.
Cladonia crispata (Ach.) Flot. K—, P —.
On soil, rocks, peat, and decaying wood.
Verterburn (J.S.T. 321); Lamb Hill (J.S.T. 281); Cascade Creek (W.M. 4494, 4715); Longwood Range (W.M. 4567).
[Cladonia rangiformis Hoffm. f. pungens (Ach.) Vain.] K +, P —.
Recorded from New Zealand by Krempelhuber (32) p. 448; requires confirmation.
Cladonia furcata (Huds.) Schrad. K — or r., P +.
On soil, logs, and tussock bases. Not common in South Island.
New Zealand (Colenso, Gunn, Richard, Jolliffe). Some early records may refer to C. scabriuscula then deemed a form of C. furcata. Cave Hill (J S.T. 237)
var. pinnata Flk. f. foliolosa Del. Pohangina River (H.H.A.), Taupo (K.W.A.); Stewart Island (W.M. 1896).
var. racemosa (Hoffm.) Flk. New Zealand (Knight in 32, p. 447); Greymouth (Helms).
Müller (42) records also var. asperata Müll. Arg., var. farinacea Vain., and var. gracillima Müll. Arg.
Cladonia scabriuscula (Del.) Leight. K—, P +. On similar habitats to last. Ohakune (J. E. Attwood); Tiritea (H.H.A.); Mt. Egmont (H. H. A.); Port Hills (H. H. A.) Nelson (H H.A.); Flagstaff (J.S.T. 1702); Tautuku (J.S.T. 1664). Maungatua (W. M.945); Cascade Creek (W.M. 5035); Greenhills (W. M. 4340). f. adspersa (Flk.) Anders. Ohakune (J. E. A); Palmerston North (H. H. A.); Feilding.
(H. H. A.). Greymouth (Helms); Outram (W.M. 4608); Saddle Hill (W. M.5107); Mt. Cargill (W.M. 6530); Greenhills (W.M. 4461).
f. surrecta Flk. Port Hills (H. H. A.) New Zealand (Travers in Paris Museum).
f. sublevis Sandst. Cascade Creek* (W.M. 4495, 4575).
f. conspersa Vain. Doubtful Sound (J. S. T.)
f semipellucida Sandst. New Zealand. Sandstede (52, p. 40.)
f. cancellata (Müll. Arg.) Sandst. Palmerston North (H. H.A.); Waitakerci Hills (H.A.A.); Maungatua (J.S.T. 2028); Greenhills* (W.M. 1754); Kuriwao 4578, 4590); Butterfly Creek* (W.M. 2802); Ridley's Bush (J.S.T. 448); (W.M. 4310); Glenledi (J.S.T. 448); Cascade Creek* (W.M. 4500, 4518,
Awarua* (W.M. 4437, 4452); Mt. Cargill* (W.M. 4437); Manapouri (W.M. 4587); Longwood Range (W.M. 4452).
f. cancellata semi-pellucida New Zealand (L. Diels).
Cladonia farinacea (Vain.) Evans. K—, P +.
New Zealand (det. H. Sandstede).
Series 2. Megaphyllae Vain.
Primary squamules rather large and persistent.
Cladonia rigida (Tayl) Hpe. K +, C —, P +.
Soil-coated rocks, dry peat. Podetia very short and isidiose.
Rangitoto (L. B. Moore); Swampy Hill (J.S.T. 1876, with reserve).
Cladonia turgida (Ehrh.) Hoffm. K +, C +, P +.
On soil covered rocks, humic soil, peat, etc. Thomson's specimens determined by W. M.
Mt. Moehau, Coromandel Peninsula (L. B. Moore); Mt. Pisgah (J.S.T. 1457, 1913).
Subsection 2. Unciales (Del.) Vain.
Primary squamules foliose but minute and evanescent. Podetia repeatedly branched, rarely scyphose. Branching by dichotomies or whorls. Axils open or both open and closed. Cups open when present. Usnic acid always present.
[Cladonia medusina (Bon) Nyl.] K + f. or —, (KC) +, P +.
Recorded from New Zealand by Krempelhuber (32); Auckland (Knight).
Cladonia amaurocraea (Flk.) Schaer. K ±, (KC) +, C ±, P —.
On earth in grass or low vegetation.
Kuriwao (W.M. 4274, 4400); Otago (H.H.A.); Maruia (J.S.T. 2425); Palmerston North (H.H.A.); Ruakura (H. H. A); New Zealand (Knight in Krempelhuber 32, p. 447); New Zealand (Jolliffe).
Cladonia Boryi Tuck. f. lacunosa Tuck. K + f, (KC) +, P—.
On earth usually in montane or subalpine stations.
Mt. Technical (J.S.T. 2446); Lead Hills and Mt. Trovatore (H.H.A. 1 p. 338); Awarua Bay (W.M. 6570, 6581); Ross (W.M. 7620).
Cladonia capitellata (Tayl.) Bab. K ±, (KC) ±, P—.
On soils in manuka heaths, less often on logs or rocks.
North and South Islands (Hooker, Colenso, Lyall); Rangitoto (H.H.A.).
f. fastigiata Vain. Rangitoto (H.H A); Taupo (K.W.A.); Feilding (H.H.A.); Greymouth (W. McKay); Huka Falls (Zahlbruckner in 62); Whangarei (Zahlbruckner in 62); Maruia Saddle (J.S.T.); Waipori (J.S.T. 202); Stoneburn (J.S.T. 2544); Kaiteriteri (W.M. 3018, 4605); Wairau Valley (W. M.4063. 4103, 3047, 3058); Longwood Range (W.M. 4477); Kuriwao (W. M. 2167, 4159); Picton (W.M. 4600); Mt. Cargill (W.M. 4407); Greenhills (W.M. 4328); Cascade Creek (W.M. 4489, 5306); Murchison (W.M. 4404); Eastbourne (W.M. 2801).
f. amaurocraeoides Vain. Lamb Hill (J.S.T. 252); Verterburn (J.S.T. 341); Flagstaff (J. S. T. 1719; W. M.6034); Rangitoto (H.H.A.); Mt. Cargill (W.M. 4102, 4408); Lake Manapouri (W. M.5038); Lake Wanaka (W. M.6800–6803)
f. degentrata Vain. New Zealand (Hochstetter); Mt. Pisgah (J.S.T. 452); Veterburn (J. S. T. 374); Rangitoto (H.H A.); Cave Hill (J.S.T. 225); Mt. Pisgah (J. S. T. 2000).
Cladonia uncialis (L.) Web. K —, (KC) +, P—.
On earth or amongst low vegetation. Mainly montane or subalpine.
Otago only (H.H.A. 1. p. 338); Otago (Jacquinot)—specimens in Paris Museum listed by Hellbom as f. turgescens.
Subsection 3. Cladinae (Nyl.) Mattick.
Primary thallus crustaceous, soon evanescent, and rarely seen. Podetia repeatedly and intricately branched, dying at the base but continuing apical growth, slender; lacking squamules, scyphi, and soredia; no true cortex, the surface usually cobwebby; inner chondroid layer well developed. Axils open or closed. Apothecia small. pale or brown, in corymbose clusters, or lacking.
Des Abbayes (4) has divided the species into three and subsequently into four series.
Series 1. Rangiferinae
Podetia stout, comprising a sympodial axis with polytomous branching. Apical branchlets unilaterally deflexed.
Series 2. Impexae
Podetia more slender, sympodia not or scarcely evident, branching trichotomous or polytomous mainly, branches subequal, ultimate branchlets straight and spreading, or deflexed in various directions. Spermagonial jelly colourless.
Series 3. Bicornutae
Podetia dichotomously branched in equal divisions; ultimate branchlets straight and spreading. P —. Spermagonial jelly colourless. Not known from New Zealand.
Series 4. Tenues
Branching mainly dichotomous, branches commonly unequal, ultimate branchlets deflexed or spreading. P +. Spermagonial jelly red.
The subgenus Cladina in New Zealand is much in need of revision. Earlier botanists placed all of the numerous forms under C. rangiferina and C. pycnoclada, the latter comprising the two forms f. flavida and f. exalbescens. C. alpestris and its var. portentosa and C. sylvatica with its var. sylvestris were next segregated from C. rangiferina. Typical C. rangiferina does not occur in New Zealand and Santesson (55) states that C. pycnoclada is confined to South America, all other locality records being erroneous. C. fallax Des Abb. is stated by him to be a synonym of C. pycnoclada. Cladonia mitis is represented in this country by f. attenuata and many plants once recorded as C. pycnoclada are now listed as C. impexa and C. leptolada. Until further research has been conducted, it is not possible to state to what taxa many herbarium specimens belong. For a fuller discussion on the Cladinae see Evans (13 pp. 530–541); Evans (19 pp. 104–112); and Santesson (55).
Series 1. Rangiferinae
Cladonia alpestris (L.) Rabenh. K—, (KC) +, P—.
Branching here is by polychotomy mainly round open axils. Subalpine bogs.
North Island (Sinclair); South Island (Haast); Maungatua (J.S.T. 2030); Key Summit, Livingstone Range (J. S. T. 2927); Sandy Point, Southland, at sea level (G. C. Martin 5605).
var. portentosa (Duf.) Vain.
New Zealand (C. Knight). Recorded Müll. Arg. (40, p. 24); Maungatua (J. S. T. 2029, 2146); Sandy Point (G. C. Martin, 5606).
Cladonia sylvatica (L.) Hoffm. K—. (KC) +, P +.
Subalpine bogs and grasslands.
Kelly Range (J.S.T. 2396); Mt. Pisgah (J.S.T. 1899a).
var. sylvestris (Oed.) Vain.
Maungatua (J. S. T.) recorded by Allan (1. p. 337).
Cladonia mitis Sandst. f. attenuata Sandst. K—, (KC) +, P—.
Eglington Valley (J.S.T in Zahlbruckner 48); Atiamuri (K.W.A.) recorded by Evans (12, p. 434).
Series 2. Impexae
[Cladonia pycnoclada (Pers.) Nyl.]
This species is said by Santesson (55) to be restricted to South America. There are numerous records for New Zealand of f. flavida and of f. exalbescens, which Allan (1, p. 337) says are common on heath and boggy soils. In his monograph of the Cladinae (4) Des Abbayes split up this species as commonly understood into four segregates—viz., C. fallax K —, P +; C. Evansi K +, P—; C. impexa K—, P —; and C. Sandstedei K +, P + and proposed that C. pycnoclada, the type of which was lost, be deleted as a “nomen confusum”. Santesson (55) gave reasons for regarding C. fallax as a synonym of typical C. pycnoclada as it occurs in the Falkland Islands, the type locality. Most early records for this species in New Zealand would seem to come into C. impexa and C. leptoclada. The form f. flavida included plants now listed as C. flavida Sandstede, but the presence of this plant in New Zealand has yet to be confirmed.
Recorded areas include New Zealand (Hooker, Colenso, Jolliffe, Raoul, Lindsay, Hugel); Saddle Hill (W. L. L); Maungatua (J.S.T. 1818a); Lee Stream (J.S.T. 216); Verterburn (J.S.T. 376); Fortrose (J.S.T. 1660); Banks Peninsula (Raoul); Chatham Islands (Travers).
Cladonia leptoclada Des Abbayes. K—, P—.
Common in heaths and on bogs. Usnic and perlatolic acids are present.New Zealand (Des Abbayes) see Evans 19, p. 106; Kaiteriteri* (W.M. 4426); Head of Cleddau Valley* (W. M.4446); Awarua bog* (W.M. 4451); Lake Manapouri* (W. M.4619); Stewart Island (W M. 4451).
Cladonia impexa Harm. K —, (KC) + P —.
A segregate from C. pycnoclada f.exalbescens common on dry heath soils. Eastbourne, Wellington* (W. M.2804, 2829); New Zealand (Des Abbayes) see Evans 19, p. 106)
Series 3. Tenues
[Cladonia subtenuis (Des Abbayes) Evans.] P +.
Not yet recorded from New Zealand. Sandstede raised C. pycnoclada f. flavida to specific rank as C. flavida; but Des Abbayes showed this to be a mixed species including C. tenuis var. subtenuis since raised by Evans to specific status as C. subtenuis. It may be expected to occur in New Zealand.
Subgenus 2. Clathrina (Müll. Arg.) Vain.
Primary thallus unknown save in C. retipora, but consisting there of minute squamules.Podetia robust, forming compact or open clusters, dying at the base, but continuing apical growth; podetial walls with very numerous and striking perforations save in some forms of C. aggregata. This last always shows some shade of brown, a colour not seen in the Cladinae, which the Clathrinae most resemble. Always corticate, no chondroid layer.
Cladonia aggregata (Sw.) Ach.K ±, P —.
Found in abundance on most available habitats. Usually brown or stramineous, with smooth, glossy, cylindrical, much branched podetia. Perforations numerous or few, occasionally absent. Usually K —, but occasionally K +.
Stewart Island* (W. M.539, 538): Saddle Hill(W. L. L., W.M.); Kaiteriteri* (W. M.3035, 3038); Lake Kanieri* (W. M.4013); Westport* (W.M. 2160); Wairau.
Valley* (W.M. 3053); Mt. Cargill* (W.M. 2153); Mihiwaka* (W.M. 2163, 2166); Greenhills* (W.M. 1765, 1771); Kuriwao (W.M. 2130); Lake Manapouri (W.M. 4586, 4608); Atiamuri (K.W.A.); Bluff Hill (J.S.T. 835); Boulder Hill (J.S.T. 105); Waipori (J.S.T. 17); Fortrose (J.S.T. 1650); Greymouth (Fr. Müller).
f. imperforata Hellb. Sutherland Falls (J.S.T. 2916); Outram Glen (W.M. 4606).
f. pygmaea Müll Arg. Port Chalmers (J.S.T. 698); Mt. Watkins (J.S.T. 318); Kuriwao (W.M. 4292).
f. racemosa Gray. Kuriwao (W.M. 3068); Flagstaff (W.M. 1130); Glenledi (J.S T. 318); Saddle Hill (W.M. 5121).
f. trichophora Müll Arg. Glenledi (J.S.T 450); Auckland (Knight).
f. straminea Müll. Arg. Greymouth (W. McKay); Lamb Hill (J.S.T. 262).
Cladonia sullivani Müll. Arg. K +, P —.
The only Cladonia known to contain divaricatic acid. (Evans.) Colour usually dark brown or even blackish. Perforations copious, revealing a nigricant interior.On bogs and in grassland in montane areas. Plants 5–15 cm tall.
Volcanic Plateau (K.W.A. 3006 H. H. A); Lake Taupo (S Berggren); Kuriwao (W.M* 2134, 2144, 3079, 3081, 4000); Lead Hills (J.S.T. 1970); Denniston (V. Zotov); Arthurs Pass (W.M. 29, 126); Longwood Range (J.S.T. 2961; W.M. 4573); Maungatua (A L. Mark; W. M. 833; J.S. T.12); Stewart Island, leg Mrs. O. Sansom (W.M. 831).
Cladonia retipora Flk. K —, P —.
The finest and most handsome of all Cladonia species, coralloid and reticulate, white, grey, or rarely faintly brown, rarely dark brown as in Australia and Chatham Island. Sea-level to subalpine on bogs, manuka heaths, sand-dunes, grassland and shingly wastes.
New Zealand (La Billardier, Raoul, Jacquinot, Hombrom, Hooker); Auckland (Sinclair); Silverdale (W.M); Rotorua (W.M 4601); Kaiteriteri (W. M.3078) at sea level; Wairau Valley (W.M 3050); Arthurs Pass (W.M. 652); Whare Flat (J S.T 500); Kuriwao (W.M 540, 2136); The Wilderness, Otago* (W.M. 4482); Longwood Range (J.S. T.2962); Port Pegasus (W.M 728); Mason Bay dunes (L. Cockayne); Lake Moana, Westland (W.M.).
Specimens collected by me on Longwood Range (4480) show a primary thallus of minute squamules. This is probably the first certain discovery of a primary thallus on any member of the Clathrinae The discovery is corroborated by Dr. A. W. Evans.
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2. Asahina, Y., 1954. Microchemical Determination of Lichen Substances Jnl. Jap. Soc. Promot. Science, Ueno, Tokyo.
3. Des Abbayes, H., 1938. Some Cladoniae of the British Dominions. Jnl. of Botany LXXVI: 346–352.
4. ", 1939. Revision monographique des Cladonia du sous-genre Cladina. Bull. Soc. Sci. Bretagne 16: 2: 1–156.
5. Duvigneaud, P. and Bleret, L., 1940. Notes de microchemie lichénique II. Sur la valeur systematique de *Cladonia pycnoclada (Pers.) Nyl. emend Des Abbayes. Bull. Soc. Roy. Bot Belgique 72: 155–159.
6. Evans, Alexander W., 1930. The Cladoniae of Connecticut. Trans. Conn. Acad. Arts and Sci. 30:. 357–510.
7. ", 1944. Supplementary Report on Cladoniae of Connecticut Ibid. 35: 519–626.
8. " 1932. Notes on the Cladoniae of Connecticut. Rhodora 34: 121–142, 153–164.
9. " 1935. Ibid. Rhodora 37: 33–57.
10. " 1938. Ibid Rhodora 40: 4–26.
11. " 1950. Ibid. Rhodora 52: 77–123.
12. " 1943. Microchemical Studies of the genus Cladonia, subgenus Cladina. Rhodora 45: 417–438.
13. " 1943. Asahina's Microchemical studies on the Cladoniae. Torrey. Bot. Club. Bull. 70: 139–151.
14. " 1935. Cladoniae of New Jersey. Torreya 35: 81–109.
15. Evans, Alexander W., 1947. Cladoniae of Vermont. Bryologist 50: 221–246.
16. " 1947. A Study of Certain North American Cladoniae Ibid. 50: 14–51.
17. " 1952. Cladoniae of Florida. Trans. Conn. Acad. Arts and Sci. 38. 249–336.
18. " 1955. Three Specres of Cladonia from Patagonia. Revue Bryol. et Lichenol T. XXIV, fasc. 1–2, 132–137.
19. " 1955. Notes on North American Cladoniae. Bryologist. 58, No 2, 93–112.
20. Fink, Bruce, 1904. Further Notes on Cladonias. Bryologist 7: 21–27; 53–58.
21. " 1905. Id. Bryologist 8: 37–41.
22. " 1906. Id. Bryologist 9: 21–24; 57–60, 89–91.
23. " 1907. Id. Bryologist 10: 21–24, 41–45; 57–60; 77–79; 97–100.
24. " 1908. Id. Bryologist 11: 21–24.
25. " 1909. Lichen Notes No. 10. Bryologist 12: 43–46.
26. " 1935. Lichen Flora of U.S.A. Ann Arbor 1–436.
27. Hooker, J. D., 1847. Flora Antarctica. London.
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29. " 1859. Flora Tasmaniae. London.
30. " 1867. Handbook of the New Zealand Flora, Pt. 2. London.
31. Hellbom, P. J., 1896. Lichenae Neo-zelandica seu Lichenes Novae Zelandiae a Sv. Berggren annis 1874–1875 collecti. Bihang k. sw. Vet. Akad. Handlung 21: afd iii, No. 13, p. 150.
32. Kremfelhuber A. Von, 1876. Neue Beitr. zur Flecten Flora Neu Seelands. Verhandl. der. k.k. zool.-bot. Gesselsch. in Wien 26: 441.
33. "1870. Lichens from Reise der Fregatte Novara. Bot. Vol. 1, pp. 107–129.
34. Lindsay, Lauder, 1866. List of 62 species of Lichens from Otago, New Zealand. Trans. Bot Soc Edinburgh 8: 349.
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36. ", 1868. Contributions to New Zealand Botany. London and Edinburgh.
37. Mattick, F., 1938. Systembildung und Phylogenie der Gattung Cladonia. Bot. Centrbl. Beihefte 58b: 215–234.
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39. Merrill, G. K., 1910. Lichen Notes No. 15 Remarks on Some Cladonia species. Bryologist 13: 103–105.
40. Muller, J. (Argov). 1892. Lichenes Knightiana, in Nova Zelandia lecti. Soc. Roy. de Bot. de Belgique 31: 22–42.
41. " 1894. Conspectus Systematicus Lichenum Novae Zelandiae. Bull. del 'Herbier Boisster II: 2–16; 17–114.
42. " 1895. Lichenes Colensoani, a Reverendis Colenso in Nova Zelandia Jnl. Linn. Soc. (Bot.) 32: 197–208.
43. Nylander, W., 1858. Synopsis Methodica Lichenum, Paris.
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45. " 1867. Addenda quodam Lichenographiam Novae Zelandiae Flora 438–440.
46. " 1888. Lichenes Novae-Zelandiae. Paris.
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Mr. Wm. Martin, 27 Merchison Street, Dunedin, E.1, N.Z.