Notes on the genus Usnea (lichenized Ascomycota,

Parmeliaceae) IV
Author: Clerc, Philippe
Source: Herzogia, 29(2) : 403-411
Published By: Bryological and Lichenological Association for Central
Europe
URL: https://doi.org/10.13158/heia.29.2.2016.403

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 Herzogia 29 (2) Teil 1, 2016: 403 – 411 403

Notes on the genus Usnea (lichenized Ascomycota,

Parmeliaceae) IV

Philippe Clerc

Abstract: Clerc, P. 2016. Notes on the genus Usnea (lichenized Ascomycota, Parmeliaceae) IV. – Herzogia 29:
403 – 411.
Several species of the genus Usnea occurring supra-continentally are treated. Usnea barbata is new to the United
Kingdom. Usnea entoviolata is new to Europe and a map of its known worldwide occurrence is given. Usnea per-
hispidella (syn. nov: U. eumitrioides), U. poliotrix, and U. subdasaea are new to North America. Usnea perplexans,
a Himalayan taxon, is the correct name for U. lapponica, a widely occurring species in Europe and North America.
Usnea aciculifera, U. hondoensis and U. pangiana are also discussed.

Zusammenfassung: Clerc, P. 2016. Anmerkungen zur Gattung Usnea (lichenisierte Ascomyceten, Parmeliaceae)
IV. – Herzogia 29: 403 – 411.
Einige Arten der Gattung Usnea mit zum Teil über mehrere Kontinente hinweggehender Verbreitung werden behan-
delt. Usnea barbata ist neu für die Britischen Inseln, Usnea entoviolata ist neu für Europa. Usnea perhispidella (Syn.
nov: U. eumitrioides), U. poliotrix und U. subdasaea sind neu für Nordamerika. Usnea perplexans, aus dem Himalaya
beschrieben, ist der gültige Name für U. lapponica, eine weit verbreitete Art aus Europa und Nordamerika. Usnea
aciculifera, U. hondoensis und U. pangiana werden diskutiert.

Key words: Systematics, taxonomy, lichens.

Introduction
In terms of species number, Usnea Adans. is the largest genus of fruticose lichens worldwide
and the second largest genus in the Parmeliaceae; some 1134 names are cited in the literature
(Clerc et al. unpublished). The exact number of well-defined and accepted species is still un-
known, but it is most probably around 400. Usnea is recognized by the presence of usnic acid
in the cortex, an elastic central cord in the medulla and the production of various depsides and
depsidones but never divaricatic and sekikaic acids. Most species occur on the bark or twigs
of trees in well-lit situations; the genus is found in polar, temperate and tropical regions and
the center of distribution is in the Neotropics. Many species seem to have a worldwide or at
least supracontinental distribution range (Clerc 2011a); this together with the high number
of published names makes the search for correct and stable names difficult. The present paper
is the result of an ongoing process aiming at studying herbarium specimens and type material
from taxa occurring all over the world.

Material and methods

The present notes are based on herbarium specimens deposited in the following herbaria: BM,
CDS, DUKE, G, H, M, TNS, TUR, UPS, W and WU, as well as in the private herbaria of Otto

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 404 Herzogia 29 (2) Teil 1, 2016

Gockman (Saint Louis, Minnesota, USA). All material was morphologically, anatomically
and chemically examined. Morphological and anatomical observations were made using a
dissecting microscope Leica MZ6. Anatomical measurements of the cortex (C), medulla (M)
and central axis (A) were recorded according to the method given in Clerc (1984, 1987).
Morphological characters used here are described in Clerc (2011b). Secondary chemistry
was analyzed with thin-layer chromatography (Culberson & Ammann 1979) with solvent B
modified (Culberson & Johnson 1982).

Taxonomy
Usnea barbata (L.) F.H.Wigg. (Fig. 1)
Prim. Fl. Holsat.: 91 (1780). – Lichen barbatus L., Sp. Pl. 2: 1155 (1753). Type: Icon in Dillenius, Hist. Musc.: t.
12, f. 6, r (1742), right-hand specimen (lectotype, Jørgensen et al., Bot. J. Linn. Soc. 115: 372 (1994)); Sweden,
Västmanland, Kila, SO om Granmuren, 1962, Nordin 1449 (epitype: UPS[!], Jørgensen et al., Bot. J. Linn. Soc.
115: 372 (1994)). %C/M/A: 3.5/35/24. Chemistry: usnic, salazinic acids (isoepitype: G).
Notes: So far none of the European species with irregular branches and salazinic acid in the medulla
[U. barbata, U. cavernosa Tuck., U. intermedia (A.Massal.) Jatta, U. perplexans Stirt. (syn. U. lappo-
nica Vain., see below) and U. substerilis Motyka] were known to occur in the United Kingdom (James
et al. 2009). During herbarium studies at the BM several years ago, I was able to find a few specimens
collected in Scotland corresponding well to Usnea barbata (= U. chaetophora sensu auct. p. p.). This
sorediate pendulous species is sometimes difficult to separate from the other sorediate pendulous spe-
cies Usnea dasopoga (Ach.) Nyl. (= U. chaetophora Stirt.) occurring in Europe with a frequent occur-
rence in the United Kingdom. It differs from U. dasopoga mainly by the distinctly irregular branches
with slightly to strongly inflated segments (Fig. 1), a thinner cortex and a rather large and dense to lax
medulla. See Clerc (2011b) for a full description and illustrations of both taxa. My taxonomic fin-
dings, however, do not accord with those published in Mark et al. (2015), where I suspect specimens
may have been misidentified. Usnea barbata is new to the United Kingdom. The species key in James
et al. (2009: 919) should be modified in the following way:
12 (10) Main branches usually distinctly irregular with the largest diameter not close to the basal
part; cortex thin to moderately thick (3 –9 %); medulla moderately thick to thick (16 –36 %), lax to
dense ................................................................................................................................... U. barbata
Main branches tapering or slightly irregular but then with largest diameter situated close to the basal
part; cortex moderately thick to thick (7–16 %); medulla thin to moderately thick (10 –26 %), dense
to compact ........................................................................................................................ U. dasopoga

Fig. 1: Usnea barbata. Longitudinal section of an irregular main branch. – Scale = 1 mm.

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 Clerc: Notes on the genus Usnea IV. 405

Additional specimens examined: UNITED KINGDOM. Scotland, Easterness (VC 96), Daviot, on larch, 1958,
J. K. Duncan (BM–733832, as U. dasopoga); Daviot, on larch, viii.1958, U. K. Duncan (BM–733830, as U. mu-
ricata); Angus (VC 90), Glen Clova, larch trunk, ix.1967, T. D. V. Swinscow (BM–733826, as U. filipendula).

Usnea entoviolata Motyka (Fig. 2)

Lich. Gen. Usnea Stud. Monogr., Pars Syst. 2: 411 (1938). Type: Ins. Sandwich [Hawaii]. D. D. Bailey s.n. (holoty-
pe: W[!]), %C/M/A: 9.5/28.5/24, chemistry: usnic, diffractaic, barbatic and squamatic (trace) acids].
Notes: This species is morphologically, anatomically and chemically close to Usnea ceratina Ach.
differing only in the development and the morphology of the soralia (Clerc 2004). It is known to
occur in North America, the Carribean region, Hawaii, Africa and South America (Truong & Clerc
2012). During an excursion in western France organized by the Association française de lichénologie
(AFL) in the Landes department near Soustons in a Quercus and Salix forest with Osmunda, I was
able to collect several specimens corresponding perfectly well to U. entoviolata. The same species was
collected on Pinus maritima in a Pine forest with Quercus suber by J. Asta (Grenoble). Furthermore I
received later from J.-L. Farou, a member of the AFL, a specimen of the same species collected in the
Pyrenees in the Ariège department at an altitude of around 1700 m, which is surprisingly high for a spe-
cies known to occur in Hawaii and the Carribean region. However, Truong & Clerc (2012) mention
several localities above 2000 m in Bolivia, Colombia, Peru, and Venezuela. Usnea entoviolata is new
to Europe. Fig. 2 gives the so far known world distribution of this species.
Additional specimens examined: FRANCE. Département des Landes, Marensin, région de Soustons-Vieux
Boucau, entre Pontolé et Piquère, à l’ouest de l’étang de Soustons, sur Pinus maritimus, 22.v.2010, P. Clerc (G–
260860); Département de l’Ariège, Plateau de Beille, Les Cabanes, c. 1700 m, 2010, J.-L. Farou (G–260917).

Fig. 2: Usnea entoviolata. Known world distribution.

Usnea perhispidella J.Steiner (Figs 3 & 4)

Sitzungsberichte Akad. Wiss. Wien, Math.-Natur. Kl. 106: 210 (1897). Type: [Kenya], Matchakos, 1896,
Liechtenstein & Pospischill s.n. (holotype: WU[!]; isotypes: W[!], G[!]), %C/M/A (holotype): 13/18.5/37, che-
mistry (holotype): usnic, stictic, constictic, cryptostictic, menegazziaic, and norstictic acids.
= Usnea eumitrioides Motyka, Lich. Gen. Usnea Stud. Monogr., Pars Syst. 2: 322 (1938). Syn. Nov. Type: [China],
Prov. Fukien occid., In monte Tienwa-Tschan ad occid. urbis Dingdschou (Tingchow), substr. arenaceo, in lapidibus

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 406 Herzogia 29 (2) Teil 1, 2016

in cacumine, ca. 1100 m, vi-vii.1921, Wang-Te-Hui s.n. (holotype: W[!]), %C/M/A: 7.5/20.5/44, chemistry: usnic,
stictic, constictic, cryptostictic, menegazziaic, and norstictic acids.
Notes: In 2015 Otto Gockman (Saint Louis, MIN, USA) sent me a few Usnea specimens collected in
Minnesota on a sandstone cliff (Fig. 3). These specimens correspond morphologically and chemically
well to U. perhispidella [see Truong et al. (2013b) for a full description]. Beside the chemistry (stictic
acid complex), the main characters of this species are the subpendulous thallus, the pale basal part, the
irregular main branches with non inflated segments and with lateral branches that are not constricted at
attachment points, the minute, ± plane soralia of the cornuta-type covering densely and ± regularly the
branches from above the basal part to the apices, and the numerous isidiomorphs developing into isi-
diofibrils often covering almost completely at least some branches or parts of them (Figs. 4A and 4B).
The morphology and the identity of the East African specimen with galbinic, salazinic and psoromic
acids mentioned by Swinscow & Krog (1975) need careful reexamination.
The holotype of Usnea eumitrioides consists of small, young, erect-shrubby thalli sharing the same
characters mentioned above and the same chemistry. It is considered here as a synonym of U. per-
hispidella.
Usnea perhispidella is mainly a corticolous species that can secondarily grow on rocks (Truong et
al. 2013). It has so far been mentioned from Africa (Swinscow & Krog 1975), Asia (Wei 1991, as U.
eumitrioides; Shukla et al. 2014, as U. eumitrioides) and South America (Truong et al. 2013b). It is
new to North America.
Usnea aciculifera Vain. [Bot. Mag. Tokyo 35: 45 (1921). Type: Japan, [Honshu], Kozuke, in arboribus,
4 viii 1913, A. Yasuda 195 [TUR–V00620! – lectotype, designated by Ohmura (2001)] is a closely
related Asiatic species with the same chemistry (Ohmura 2001, 2012). However the minute soralia

Fig. 3: Usnea perhispidella. Habitat (sandstone cliffs in Minnesota). Photograph by John Thayer (Minnesota).

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 Clerc: Notes on the genus Usnea IV. 407

Fig. 4: Usnea perhispidella. A – Soralia of the cornuta-type. B – Isidiomorphs and isidiofibrils. – Scales = 0.5 mm.

are not regularly but unevenly distributed on the branches, in irregular patches of ± aggregated soralia
(Fig. 5). Furthermore, the isidiomorphs of U. aciculifera remains short and rarely develop into isidio-
fibrils. So far U. aciculifera is known to occur only in Asia: China, India, Nepal and Taiwan (Asahina
1972, Awasthi 1986, Ohmura 2001, 2012, Shukla et al. 2014, Wei 1991).
Usnea hondoensis Asahina [Lich. Jap. 3: 87 (1956). Type: Japan, Honshu, prov. Musashi, Nippara,
10.1922, Y. Asahina 115 (TNS! – holotype)] has the same morphology with the numerous cornuta-
type soralia covering evenly the branches (Fig. 6A). However the branches are distinctly segmented
with annular cracks and isidiofibrils are lacking. The chemistry is different with salazinic, ± norsticitic,
± barbatic acids. I disagree with Ohmura (2001, 2012) who synonymized this species with U. pangia-
na Stirt. [Scott. Natur. 7: 77 (1883). Type: India, Himalaya, Lingalelah Range, 2000 –2300 m, G. Watt
7051 (BM! – holotype)]. The chemistry is the same but the holotype of U. pangiana has much larger
and stronger, stipitate, not cornuta-like ‘soralia’ (Fig. 6B) that are probably growing out of fibercles.
Additional specimen examined: U.S.A. Minnesota, Houston Co: ca. 5.7 km north of Eitzen and 1.1 km southeast
of Camp Winnebago, along Winnebago Creek, saxicolous high up on a large NE facing sandstone cliff; 20.i.2015,
O. Gockman & J. Thayer (OTG05199a).

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 408 Herzogia 29 (2) Teil 1, 2016

Fig. 5: Usnea aciculifera. Soralia of the cornuta-type in patches. – Scale = 0.5 mm.

Usnea perplexans Stirt.

Scott. Naturalist (Perth) 6: 103 (1881). Type: India, Himalayas, [Himachal Pradesh], Pangi, Dr. G. Watt s.n., (iso-
type: BM[!]), %C/M/A (isotype, specimen A): 4/34/24, chemistry (isotype, specimens A, B & C): usnic, salazinic
acids.
= Usnea lapponica Vain., Meddeland. Soc. Fauna Fl. Fenn. 48: 173 (1925, ‘1924'). Type: Russia, Murmansk
Region, Lapponia Imadrae, Lovozero [Lowosersk], ad ramulos Piceae, 25.v.1887, Kihlman s.n. (lectotype: H[!],
Clerc, Nord. J. Bot. 7: 494, 1987; isolectotype: TUR[!]), %C/M/A (isolectotype): 6/28/32, chemistry (lectotype,
isolectotype): usnic, salazinic acids.
Usnea perplexans was misspelled “U. perplectans” by Motyka (1936: 293).
Figures: Clerc (2011b: 169, as U. lapponica).
Because of the presence of a slightly orange pigmentation of the central axis in U. perplexans, Clerc
(1987) disagreed with Carlin & Swahn (1977) who reduced U. lapponica to synonymy with U.
perplexans. It is a fact that in old specimens collected in tropical areas, the surface of the axis and the
medulla close to the axis can become slightly orangish pigmented when either salazinic or norstictic
or galbinic acids are present in the medulla. This might be the result of a chemical alteration of the
depsidones over time, without any taxonomic value. Both species are morphologically and chemi-
cally identical, sharing the erect-shrubby thallus, the irregular main branches with lateral branches not
constricted at attachment points, the large and deeply excavate soralia without isidiomorphs and the
presence of salazinic acid in the medulla. Anatomically, if we consider the mean values of the %C/M/A
of European specimens of U. lapponica (n = 122) and the %C/M/A of the isotype of U. perplexans
(Tab. 1), we can see that, although somewhat extreme, all the values of the latter species fit in the
variability of U. lapponica.
Usnea perplexans is a corticolous Northern Hemisphere taxon known to occur in cold, mountainous are-
as of Europe [Wirth (2013) as U. lapponica, Clerc (2011b) as U. lapponica], North America [Brodo et
al. (2001) as U. lapponica] and Asia [Awasthi (1986), Shukla et al. (2014)]. Swinscow & Krog (1979,
1988) mention this species for Africa including it in the U. abissinica group. However, this group of non-

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 Clerc: Notes on the genus Usnea IV. 409

Fig. 6: A – Usnea hondoensis. Soralia of the cornuta-type. B – Usnea pangiana. Fibercles turning into stipitate
soralia. – Scales = 0.5 mm.

Table 1: Usnea lapponica and U. perplexans. Comparison of  CMA values (C = cortex, M = medulla, A = central axis).

  %C (cortex) %M (medulla) %A (central axis)

U. lapponica (n= 122) (3 –)5.3 – 6.8 – 8.3(–11.5) (8 –)21.2–25.5 –29.8(–35.5) (21–)28.1–35.5 – 42.9(–71)

U. perplexans (isotype) 4 34 24

pigmented tropical species with large and excavate soralia has ± swollen, fusiform branches with lateral
branches constricted at attachment points, and a cornuta-like cortex and CMA type. Usnea perplexans
should therefore be excluded from the U. abissinica group and is so far not known to occur in Africa.

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 410 Herzogia 29 (2) Teil 1, 2016

Usnea poliotrix Kremp.

Vidensk. Meddel. Naturh. Foren. Klöbenhavn. 5: 4 (1873). Type: Brazil, Minas Gerais, Warming s.n. (holotype:
M[!]), %C/M/A: 6.5/32.5/22, chemistry: usnic, protocetraric and lobaric acids.
Figures: Truong et al. (2011: 480, 492).
Note: In 1989, when working on the genus Usnea in eastern North America in the herbarium of Duke
University (DUKE), I came upon a strange collection made by Barbara Joe Moore in Florida (Moore 1968):
a sorediate subpendulous to pendulous species with a subcortical pigment and salazinic, galbinic, norstictic
and lobaric acids in the medulla. This collection remained a mystery until 22 years later Camille Truong and
I were studying type material from South America. Its identity as Usnea poliotrix is beyond doubt.
The diagnostic features of Usnea poliotrix are the subpendulous to pendulous thallus, the irregular
branches often with ridges and foveoles, the numerous isidiofibrils covering the branches growing
from the minute soralia, the orange subcortical medulla pigmentation and the chemistry [see Truong
et al. (2011) for a complete description]. Usnea poliotrix is a mainly corticolous, very common spe-
cies in the Galapagos Islands where it is found in the arid and transition zones, more rarely in the
humid zone (Truong et al. 2011). The specimens found in the Galapagos Islands have the same che-
mistry (salazinic, galbinic, norstictic, and lobaric acids) as the specimen collected in Florida by Moore.
However the type of this species collected in Brazil contains protocetraric and lobaric acids. Usnea
poliotrix seems to be rare on the South American continent since I don’t know of any other specimen
collected there. Usnea poliotrix is new to North America.
Additional specimens examined: U.S.A. Florida, Charlotte Co., Englewood, scrub pine, 16.v.1964, B. J. Moore
1352a, b (DUKE), %C/M/A (1352b): 8/29.5/25, chemistry (1352a, b): usnic, salazinic, galbinic, norstictic, & lo-
baric acids.

Usnea subdasaea Truong & P.Clerc

Bryologist 114: 499 (2011). Type: Ecuador, Galapagos Islands, Isabella, road to Sierra Negra crater, close to la
Esperanza, 306 m, farming areas in the humid zone, living fence posts, Truong 1194 (holotype: CDS39505[!]; iso-
types: G[!], UPS[!]), %C/M/A: 5/30/30, chemistry: usnic, salazinic, galbinic, and norstictic acids.
Figures: Truong et al. (2011: 492).
Notes: In 1989, when studying the genus Usnea at Duke University (DUKE), I came upon a second
strange specimen collected by B. Moore in Florida. The morphology and the chemistry of this specimen
correspond well to Usnea dasaea Stirt. [see Clerc & Herrera Campos (1997) for a full description of
this taxon], however with the presence of a subcortical red pigment. It remained labeled as such until we
discovered further material from the Galapagos Islands that was described as Usnea subdasaea (Truong
et al. 2011). The presence of the pigmentation was later confirmed as being an important specific charac-
ter since U. subdasaea and U. dasaea are not closely related (Truong et al. 2013a). Usnea subdasaea is
a frequent taxon in the Galapagos, usually in the farming areas of the humid zone. It seems to be rare in
the Andes at low altitude (Truong et al. 2011). Usnea subdasaea is new to North America.
Specimen examined: U.S.A. Florida, Lake Co., S-433, 2,5 mi. s. of 46 Bay-head, twigs of Magnolia virginiana,
10.iv.1964, B. J. Moore 799 (DUKE), %C/M/A: 7.5/29.5/27, chemistry: usnic, salazinic, galbinic and norstictic acids.

Acknowledgements
I am thankful to the curators of the herbaria cited above, to Otto Gockman (Saint Louis, Minnesota, USA) for send-
ing me many interesting specimens he collected in the state of Minnesota, and to J.-L. Farou (Barsac, France) for
sending me the U. entoviolata specimen from the Ariège department. Many thanks to the Association Française de
Lichénologie (AFL) for organizing the course on the genus Usnea and the excursions around Souston (F, Les Landes).
Finally, thank you to Camille Truong for many fruitful discussions and sharing with me her knowledge of the South
American species of the genus Usnea.

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Manuscript accepted: 30 July 2016.

Communicated by: Philipp Resl

Address of the author

Philippe Clerc, Conservatoire et Jardin botaniques de la Ville de Genève, 1, ch. de l’Impératrice,
Case postale 60, 1292 Chambésy, Switzerland. E-mail: philippe.clerc@ville-ge.ch

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