Lichens A-Pac (2007) - Flora of New Zealand Lichens - Revised Second Edition A-Pac
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Cladonia P.Browne

CLADONIA P.Browne, 1756  nom. cons. 

Type : Cladonia subulata (L.) F.H.Wigg. [=Lichen subulatus L.]

Description : Flora (1985: 100). See also Archer (1992b: 111), Ahti (2000: 82) and Ahti & Hammer (2002).

Key

1
Podetia with prominent cups, unbranched or with proliferations from cup margins; cups equal to or wider than supporting podetium, usually much wider
2
Podetia without prominent cups, branched or unbranched; cups usually narrower than supporting podetium
19
2
Podetia yellow
3
Podetia not yellow
8
3
Podetia with soredia
4
Podetia without soredia
7
4
Podetia partially sorediate; soredia farinose or granular; lowland
5
Podetia entirely covered with farinose soredia; alpine
5
Podetia and cups ±symmetrical
6
Podetia and cups asymmetrical
6
Base of podetia distinctly corticate; apothecia, if present, red; common
Base of podetia indistinctly corticate; apothecia, if present, beige; uncommon
7
Podetia and cups more or less symmetrical, untorn, proliferations absent; apothecia, if present, red
Podetia and cups asymmetrical, often torn longitudinally, proliferations common; apothecia, if present, brown
8
Podetia without soredia
9
Podetia at least partly sorediate
11
9
Primary thallus unfused, squamulose, not resembling a foliose lichen; lower surface not cottony; podetia ≥ 1 cm tall
10
Primary thallus fused, resembling a foliose lichen; lower surface cottony; podetia < 1 cm tall
10
Cup interiors with corticate granules; Pd+, UV−
Cup interiors without corticate granules; Pd+ or −, UV+ blue
11
Base of podetia distinctly corticate
12
Base of podetia sorediate or indistinctly corticate
14
12
Podetia UV−; apothecia if present, brown
13
Podetia UV+ ice-blue (squamatic acid): apothecia if present, red
13
Podetia containing fumarprotocetraric acid and accessory atranorin
Podetia containing fumarprotocetraric acid, atranorin and bourgeanic acid
14
Cups no wider than supporting podetia
15
At least some cups in clump wider than supporting podetia
16
15
Cups without proliferations or with blunt proliferations
Cups with subulate proliferations
16
Podetia with fumarprotocetraric acid and other compounds
17
Podetia with fumarprotocetraric acid only
17
Podetia with merochlorophaeic acid
18
Podetia with cryptochlorophaeic acid
18
Podetia with homosekikaic acid
Podetia without homosekikaic acid
19
Surface of podetia arachnoid; without soredia
20
Surface of podetia not arachnoid: with or without soredia
21
20
Podetia in a single thallus generally of variable widths; tips downturned, whitish
Podetia in a single thallus generally of similar widths; tips ± randomly oriented, greyish
21
Thallus with soredia
22
Thallus without soredia
39
22
Thallus K+ yellow
23
Thallus K− or K+ dingy yellow turning to brownish
26
23
Podetia unbranched or rarely, sparsely branched
24
Podetia branched
25
24
Basal squamules persistent; podetia K+ yellow weak (atranorin)
Basal squamules persistent or absent, podetia K+ yellow, strong (thamnolic acid)
25
Podetia with cups bearing twisted proliferations
Podetia without proliferations on cups
26
Podetia not yellow (usnic acid absent)
27
Podetia yellow (usnic acid present)
27
Podetia ≥1 cm tall; not hood-shaped
28
Podetia <1 cm tall, hood-shaped
28
Podetial squamules absent or sparse
29
Podetial squamules present, often abundant
31
29
Podetia forming narrow cups
Podetia not cup-forming
30
30
Podetia >2 cm tall, Pd+ red, often curved and decurved; apothecia, when present brown
Podetia <2 cm tall, Pd−, uncurved; apothecia, when present, red
31
Soredia or soredium-like granules not restricted to tips of podetia
32
Soredia restricted to tips of podetia
32
Tips of mature podetia brown, inflated, deformed
33
Tips of mature podetia not inflated or deformed
34
33
Bases of podetia brown; top 1/4 of podetium similar to rest of podetium; soredia farinose
Bases of podetia concolorous with rest of podetium, top 1/4 of podetium smooth or with scattered microsquamules, soredia granular
34
Soredia farinose
35
Soredia granular
36
35
Podetia Pd+ yellow (psoromic acid)
Podetia Pd+ red (fumarprotocetraric acid)
36
Podetia not cup-forming
37
Podetia cup-forming
38
37
Podetia straight, whitish, soredium-like granules under imbricating squamules; infrequent
Podetia slightly curved, especially toward tips, greenish-brownish, soredium-like granules under squamules and along surface of podetium: common
38
Cups with brown meristematic tissue on margins, proliferations or brown tissue always present: common
Cups without brown meristematic tissue or proliferations, rare
39
Podetia cup-forming; cups in one-to several verticillate tiers
40
Podetia cup-forming or not cup-forming; tiers absent
43
40
Bases of podetia not black
41
Bases of podetia black
41
1–2 tiers of cups present
More than 2 tiers of cups present
42
42
Cortex K+ (atranorin)
Cortex K− or K+ yellowish to dingy brown
43
Some part of podetia appearing torn or fissured
44
No part of podetia appearing torn or fissured
53
44
Podetia tubular
45
Podetia not tubular
48
45
Podetia branched or unbranched, fissures and tears not restricted to branches
46
Podetia branched; longitudinal fissures restricted to branches
46
Podetia K− or + yellowish to dingy brown
47
Podetia K+ yellow (atranorin)
47
Podetia without cups, rarely branched, discrete longitudinal vulnerae on podetia
Podetia with or without cups, usually branched, irregularly torn and shredded
48
Basal squamules generally distinguishable from podetia
49
Basal squamules barely distinguishable from podetia, usually similar height or longer
49
Podetia K− or +, Pd− or +
50
Podetia K−, P+ red (fumarprotocetraric acid only)
50
Basal squamules persistent or absent; branching frequent; axils open
51
Basal squamules persistent, branching infrequent; axils closed
51
Pd + weak yellow (psoromic acid absent)
52
Pd+ strong yellow (psoromic acid present)
52
Stictic acid major compound
Norstictic acid major compound
53
Cortex K+ yellow
54
Cortex K− or K+ dingy yellowish to brownish
63
54
Podetia usually in thick, grass-like masses
55
Podetia usually distinct, not massed
58
55
Podetia greenish, whitish, brownish or yellowish: common
56
Podetia ashy-greyish: infrequent
56
Podetia instantly K+ yellow on most of podetial surface
57
Podetia slowly K+, reaction strongest at tips
57
Podetia yellowish (usnic acid); base concolorous with rest of podetium
Podetia not yellowish (usnic acid absent); base concolorous or bluish to blackened
58
Podetia with prominent, often abundant podetial squamules
59
Podetia without prominent, abundant podetial squamules
61
59
Podetial squamules >0.3 mm: on various substrata
60
Podetial squamules <0.3 mm: on rotting decorticated wood
60
Squamules orange toward base; apothecia, when present, red
Squamules not orange toward base; apothecia, when present, brown
61
Tips of podetia not decumbent
62
Tips of podetium decumbent (dissecting microscope needed)
62
Outer surface appearing thick, relatively smooth to chinky-areolate, with bluish tinge; apothecia, when present, brown; alpine
Outer surface rough, scurfy-subsquamulose, podetia greenish to yellowish; apothecia, when present, red; lowland
63
Podetia usually in thick, grass-like masses
64
Podetia usually distinct, not massed
67
64
Podetia brownish
65
Podetia yellowish
66
65
Podetia Pd+ red, UV− (fumarprotocetraric acid)
Podetia Pd−, UV+ ice-blue (squamatic acid)
66
Podetia UV+ ice-blue (squamatic acid)
Podetia UV− (usnic acid only)
67
Podetia with prominent, often abundant podetial squamules
68
Podetia without prominent, abundant podetial squamules
73
68
Podetia Pd+ red
69
Podetia Pd− or + yellow
72
69
Outer surface of podetia scurfy, appearing sorediate
70
Outer surface of podetia smooth, without soredia
70
Podetia greenish to brownish
71
Podetia yellowish
71
Podetia with straight tips; squamules + equally distributed along length of podetia
Podetia with curved tips, squamules more frequent toward base and often more abundant on one side of the podetia
72
Podetia instantly K+ yellow over entire surface
Podetia slowly K+, reaction strongest toward tips
73
Podetia Pd+ red (fumarprotocetraric acid)
74
Podetia Pd+ yellowish or −
80
74
Podetia >2.5 cm tall; branches if present occurring lower on podetium
75
Podetia <2.5 cm. tall, 1–3× branched near apex; surface tessellate-areolate
75
Podetia + smooth and shiny, brownish to greenish
76
Podetia + rough, matt, with yellowish tinge
76
Podetia without swollen apices
77
Podetia with swollen apices
77
Podetia irregulary branched or unbranched
78
Podetia + regularly, abundantly, dichotomously branched
78
Podetial branches not candelabrum-like
79
Podetial branched candelabrum-like; (uncommon)
79
Podetia K− or + dingy yellowish to brownish
Podetia briefly K+ yellow (atranorin)
80
Outer surface smooth; apothecia, when present, brown
81
Outer surface scurfy, appearing sorediate; apothecia, when present, red
81
Podetia UV+
82
Podetia UV−
82
Podetia mostly recumbent, yellowish, highly branched; alpine
Podetia mostly erect, greenish, not highly branched

Cladonia is a genus of c. 450 known species in the family Cladoniaceae (Kirk et al. 2001; Stenroos et al. 2002a; Eriksson et al. 2004; Pennycook & Galloway 2004), widely distributed in both Northern and Southern hemispheres. Cladonia is the most highly speciose lichen genus in New Zealand with some 70 taxa currently recognised. The first relatively modern treatment of New Zealand species of Cladonia was that of Sandstede in Zahlbruckner (1941: 324–328) based largely on collections made by H.H.Allan and J.S. Thomson in the 1930s. Carroll Dodge's account of subantarctic taxa in the Cladoniaceae (Dodge 1948) is a neglected but still useful treatment, dealing with taxa from the subantarctic islands. In the 1950s William Martin took up Cladonia in earnest, making extensive collections from many parts of the South I. (Martin 1958: 607). His identifications (Martin 1958) were based in large part on the work and assistance of the Yale lichenologist, A.W. Evans (who placed New Zealand material sent to him by Martin, in mainly Northern Hemisphere names), but he later added several new taxa of his own (Martin 1962). These initial accounts, Dodge aside, failed to recognise New Zealand as an area of much endemism or speciation in Cladonia, the observed variation in the large collections made between 1926 and the 1960s resulting in recognition of many infraspecific taxa of mainly Northern Hemisphere names. For additional notes on the genus in New Zealand see Galloway (1985a: 100–124), but this account too has many deficiencies and is not a reliable guide to species identification of taxa now known to be present here.

Recent taxonomic work on Cladonia from the Northern Hemisphere, Australia, North and South America, Antarctica and the subantarctic shelf islands has clarified the identity and distributions of many taxa that also occur in New Zealand and the following accounts are of relevance to any study of New Zealand populations (Ahti & Kashiwadani 1984; Archer 1985a, 1985b, 1986a, 1986b, 1988, 1989, 1992b; Archer & Bartlett 1986; Stenroos 1986, 1988a, 1989, 1993; Ahti et al. 1990; Stenroos & Ahti 1991; Stenroos & Smith 1992; Hammer 1993b, 1995a, 1995b, 1996a, 1996b, 1996c, 1996d, 1996e, 1997a, 1997b, 1998a, 1998b, 1999, 2000, 2002, 2003a, 2003b; Hyvönen et al. 1995; Ahti 1997, 2000; Stenroos et al. 1997; Øvstedal & Lewis Smith 2001; Ahti & Hammer 2002; Stenroos et al. 2002b, 2002c; Litterski & Ahti 2004; Ahti & De Priest 2005). Chemistry in Cladonia is reviewed in Huovinen et al. (1990).

Molecular studies (Hyvönen et al. 1995; Stenroos et al. 1997, 2002b; Stenroos & De Priest 1998a; De Priest et al. 2000) indicate that taxa formerly included in the genus Cladina Nyl. (e.g. Hale & Culberson 1970; Brodo 1976; Ahti & Lai 1979; Follmann 1979; Ahti 1984, 2000; Ahti & Kashiwadani 1984; Galloway 1985a; Stenroos 1988b, 1993; Beard & De Priest 1996; Ahti & De Priest 2001) are best kept within Cladonia, a procedure followed in this account. Recent molecular studies (Stenroos et al. 2002a, 2002b, 2002c) have led to a reappraisal of infrageneric taxonomy within Cladonia, their results not supporting current sectional classification. Stenroos et al. (2002a) propose a new preliminary classification of Cladonia and distinguish three informal subdivisions. Within these subdivisions four supergroups and, further, seven groups and two subgroups are distinguished (Stenroos et al. 2002a: 251), and no doubt this arrangement will change again once the remaining 300 taxa are analysed and a world monograph is achieved. In anticipation of this, parts of the proposed monograph are being made available online as regional treatments are consolidated (see Stenroos et al. 2002c).

What has severely frustrated local studies of Cladonia, to a very large extent, is a comprehensive knowledge of the variation of Northern Hemisphere taxa in order to be sure which of the species present in New Zealand are truly cosmopolitan, and which are more regional or local in their distribution. We have been singularly fortunate in recent years to have a Northern Hemisphere Cladoniologist take a detailed interest in species of Cladonia in New Zealand. Since January 1999, Prof. Samuel Hammer (Boston University) has made several visits to New Zealand (supported by grants from the National Science Foundation PEET Programme, and from the National Geographic Society) specifically to study the family Cladoniaceae, in the field and in the herbarium, as part of a wider study of morphological development, speciation and phylogeny of the family in the Southern Hemisphere. It is more highly diversified in the Southern Hemisphere than it is in the Northern Hemisphere, and may reflect a Gondwanan origin of the group. He has patiently worked through large unidentified Cladonia collections of H.H. Allan, J.K. Bartlett, J. Child, P. Child, D.J. Galloway, W. Martin and J.S. Thomson and revised the named collections in AK, BM, CHR, OTA and WELT and this, together with his extensive field knowledge of the genus in New Zealand from Fiordland to Northland, has given him an unrivalled overview of this genus in a local context. The present account is based on his published accounts as well as on collaborative work with him in the field and the herbarium, and he also prepared the revised key to species.

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