Lichens Pan-Z (2007) - Flora of New Zealand Lichens - Revised Second Edition Pan-Z
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Usnea Dill. ex Adans.

USNEA Dill. ex Adans., 1763

=NEUROPOGON Nees & Flot., 1835

=EUMITRIA Stirt., 1881

Type : Usnea plicata (L.) A.L.Sm. [=Lichen plicatus L.]

Type : Neuropogon antennarius Nees & Flot. [Usnea aurantiaco-ater (Jacq.) Bory]

Type : Eumitria baileyi Stirt. [=Usnea baileyi (Stirt.) Zahlbr.]

Description : Flora (1985: 596).

Key

1
Saxicolous or terricolous
2
Corticolous or lignicolous
13
2
Saxicolous
3
Terricolous or straggling over low, subalpine vegetation
3
Black surface annulation present; apothecial discs dark-coloured (black, brown or greenish grey)
4
Black surface annulation absent; apothecial discs pale yellow-green
9
4
Soredia absent; apothecia usually present
5
Soredia present; apothecia rare or absent
6
5
Thallus entirely erect to 9 cm long; branches stout; apothecia subterminal, rarely lateral, frequent
Thallus pendulous, straggling, to 15 cm long; branches filiform, capillary; apothecia lateral, rare
6
Medulla ±densely interwoven, axis thick, more than half diameter of branch
7
Medulla lax, axis thin, less than half diameter of branch; cortex smooth to minutely papillate; soralia small, inconspicuous, ±blackened, ±convex-pulvinate
7
Cortex smooth, wavy, ±black-edged, annulate-cracked
8
Cortex matt, ±coarsely papillate, not annulate-cracked
8
Thallus ±subdecumbent, to 3 cm long; richly branched from a delimited holdfast, with numerous, fragile, capillary, divergent secondary branches; soralia pale, punctiform
Thallus erect, to 6 cm long; ±monopodial or moderately branched, often from a proliferating holdfast; branches stout; soralia plane, pale, rarely nodular, corticate then pigmented
9
Sorediate or isidiate, without red pigments
10
Without soredia; red-pigmented
12
10
Sorediate
11
Isidiate, pseudocyphellate; containing norstictic, salazinic and galbinic acids
11
Pendulous, to 40 cm long; soredia becoming isidiate; salazinic acid present (K+ yellow→red) [northern coastal]
Erect to partly decumbent, to 5 cm long; soredia in granular masses, not becoming isidiate; salazinic acid absent, squamatic acid generally present [alpine–subantarctic]
12
Thallus shrubby, erect, to 6 cm long
Thallus subpendulous, to 10(–30) cm long
13
Thallus with a solid axis
14
Thallus with a hollow axis
14
Soredia present
15
Soredia absent
22
15
Medulla not yellow; red cortical spots absent
16
Medulla yellow; red cortical spots present (×10 lens)
16
K+ yellow→red (salazinic acid present)
17
Thallus K+ yellow or brownish (salazinic acid absent; psoromic or fumarprotocetraric acids present)
21
17
Soralia pseudoisidiate, spinulose; secondary branches numerous
18
Soralia efflorescent, confluent, without spinules or pseudoisidia; secondary branches rare
20
18
Thallus pale yellow-green, occasionally brownish, never reddish
19
Thallus distinctly reddish
19
Thallus pendulous, to 40 cm long; northern coastal, rare
Thallus shrubby, 3–10 cm long; widespread, common
20
Soralia apical and subapical on terminal branches and branchlets; papillae, if present, sparse to moderate
Soralia on secondary, tertiary and terminal branches, not present on primary branches; papillae dense
21
Thallus K+ deep-yellow (psoromic acid present)
Thallus K− or + brownish (fumarprotocetraric acid present)
22
Branches papillate
23
Branches without papillae
26
23
Thallus 4–6(–10) cm tall
24
Thallus to 2.5 cm tall
25
24
Thallus K+ yellow→red (salazinic acid present); lateral fibrils frequent
Thallus K− or + brownish (fumarprotocetraric acid present)
25
Branches 1 mm thick and 1–3-branched at base; cortex distinctly papillate
Branches 0.5–0.8 mm thick, simple to 1–2-branched at base; cortex indistinctly papillate
26
Thallus pendulous (5–)30–60(–200) cm long; branches fine
27
Thallus shrubby, 2–3(–5) cm tall; branches inflated
27
Thallus branches soft, pliant, green, to pale greenish yellow or whitish, not longitudinally ridged or winged
28
Thallus branches stiff, olivaceous, tinged reddish, distinctly longitudinally ridged or winged
28
Branches terete; cortex not eroded; fumarprotocetraric acid present; a canopy "mist" species at or near treeline in Nothofagus forest
Branches terete to flattened and ribbon-like; cortex partly eroding; protocetraric acid present; on trees in northern coastal forest

Usnea, included in the family Parmeliaceae (Eriksson et al. 2004; Pennycook & Galloway 2004; Eriksson 2005), comprises c. 300 species worldwide (Kirk et al. 2001). Although Usnea is one of the most readily recognised and easily collected of fruticose lichen genera, it still remains poorly understood at the species level, because of the often extreme plasticity of such characters as colour, length of thallus branches, density of branching, development and density of fibrils, shape of papillae, presence of faveolae, and thickness of growth apices (Clerc 1991). According to Clerc (1984a, 1987a, 1987b, 1998) the best diagnostic morphological characters in Usnea are: (1) colour and frequency of segementation of the basal portions of the thallus, (2) shape of the secondary branches, (3) dominant branching patterns, (4) type of soralia, and (5) details of the internal structure. The shape and density of papillae and the density of fibrils are also useful but appear to be modified to a large extent by environmental factors. The colour of the living thallus has only minimal diagnostic value, except when other pigments in addition to usnic acid are present. A ±well-developed internal prosoplectenchymatous medullary cord is a characteristic feature of the genus (see Wright 1999). Apothecia, when present, are zeorine, characterised by a double-layered exciple. Asci are Melanelia -type. Pycnidia are laminal and immersed. Conidia are sublageniform (Kärnefelt et al. 1998: 87, fig. 5F). Recent molecular work on the Usnea florida – U. subfloridana species-pair, distinguished mainly by their dispersal strategies, with U. florida being the fertile, primary species and U. subfloridana the derived, sterile, secondary species, has shown them to be one monophyletic group of intermixed specimens (Articus et al. 2002). This result has major consequences for our understanding of "species-pairs" in both Usnea and other lichen genera.

Although the Usnea monograph of Motyka (1936, 1937, 1938, 1947) is still an important, though outdated, resource for many areas of the world, including New Zealand, in recent years taxonomic work on Usnea has revived in the Northern Hemisphere and modern methods have been used in several important regional studies, viz. Swinscow & Krog (1974, 1975c, 1976a, 1976c, 1978b, 1979a, 1986b, 1988) for East Africa; Clerc (1984a, 1987b, 1991, 1992, 1994, 1997), Clerc & Diederich (1991), and Fos & Clerc (2000) for Europe; Clerc (1987a, 2004), Esseen et al. (1981), Myllys (1994), Halonen & Puolasmaa (1995), and Halonen (1997) for Scandinavia; Tavares (1987, 1997), Clerc & Herrera-Campos (1997), McCune & Geiser (1997), Halonen et al. (1998), Halonen (2000), Herrera-Campos et al. (1998, 2001), and Clerc (2004) for North America including Mexico; Awasthi (1986) for India; Ohmura & Kashiwadani (2000), Ohmura et al. (2000), and Ohmura (2001, 2002) for Japan; and Rogers & Stevens (1988), Stevens (1990, 1991a, 1991b, 1992, 1999, 2004b) and Clerc (2004) for Australia. James (2003) has recently compiled an aide mémoire to the British species of Usnea, an account having useful information on species discrimination in the genus.

The Neuropogon"group" in Usnea is a bipolar assemblage of saxicolous taxa, with all but one of its 15 species confined to high-altitude rock outcrops in Arctic, Antarctic or southern cool-temperate biomes (Lamb 1939, 1948b, 1964; Galloway 1985a; Walker 1985; Stevens 1999, 2004a; Øvstedal & Lewis Smith 2001). Recent molecular studies (Ohmura & Kanda 2004) place Neuropogon as a section within subgen. Usnea. Another molecular study (Articus 2004) reached a different conclusion and confirmed that Dolichousnea, Eumitria and Neuropogon are all monophyletic, and should be treated as independent genera, but that Usnea as presently conceived is clearly non-monophyletic and requires further analysis using larger data sets. However, recently published molecular work on 12 of the 15 species known from Neuropogon (including the generitype) has indicated that Neuropogon is polyphyletic, with a core group nested within Usnea subgen. Usnea as sister-group to sect. Usnea (Wirtz et al. 2006). Accordingly, they synonymise Neuropogon with Usnea, a position followed here.

In New Zealand, Usnea is a common and widespread genus, occurring in a variety of habitats from coastal to subalpine and even high-alpine biomes. Preliminary accounts of New Zealand species are given in Galloway (1985a) and Stevens (1999), but the genus is still far from understood here and is much in need of serious revisionary study. Although large collections of Usnea exist in several major herbaria in New Zealand, it is surprising that until very recently no serious attempt has been made at a regional study of the genus, a study that would amply repay the effort invested in it. Darrell Wright (Wellington) has recently begun investigations of lower North I. populations (Wright 2003, 2004) and has already added useful information to our knowledge of New Zealand's Usnea mycobiota. The present treatment discusses 28 species, but it is still only a very preliminary account.

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