A Synopsis of the Lichen Genus Psiloparmelia (Ascomycotina, Parmeliaceae)

Author(s): John A. Elix, Thomas H. Nash and III

Source: The Bryologist, Vol. 95, No. 4 (Winter, 1992), pp. 377-391
Published by: American Bryological and Lichenological Society
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 THE BRYOLOGIST
QUARTERLY JOURNAL OF THE AMERICAN BRYOLOGICAL AND LICHENOLOGICAL SOCIETY

VOLUME
95 WINTER1992 4
NUMBER

The Bryologist 95(4), 1992, pp. 377-391

Copyright ? 1992 by the American Bryological and Lichenological Society, Inc.

A Synopsis of the Lichen Genus Psiloparmelia

(Ascomycotina, Parmeliaceae)

JOHNA. ELIX
Chemistry Department, The Faculties, AustralianNational University, GPO Box 4, Canberra,A.C.T. 2601,
Australia

THOMASH. NASH, III

Departmentof Botany,Arizona State University, Tempe, AZ 85287-1601

Abstract. The genus Psiloparmelia Hale has its center of diversity in the high Andes Mountains
of South America, but also occurs in the mountains of southern Africa. The genus is characterized
by a velvety lower surface without rhizines, isolichenan in the cell walls, a rudimentary pored
epicortex, unusual distribution of secondary metabolites, and significant concentrations of both
atranorin and usnic acid in the upper cortex. Psiloparmelia has secondary-product chemistry char-
acterized by O-orcinol depsides and depsidones and fatty acids. A total of 24 known compounds is
reportedfor the genus, based upon thin-layer and high-performance liquid chromatographic analyses
of 189 specimens. Psiloparmelia is now considered to comprise 12 species; nine are here described
as new and one is a new combination.

The generic concepts in several groups of lichens Hale (1989a) considered that the following char-
are currently in a state of flux, especially in the acters were most significant in distinguishing this
crustose genera Lecanora Ach. and Lecidea Ach. new genus: the velvety lower surface without rhi-
and the macrolichen family Parmeliaceae (Hale zines, a negative test for lichenan, and a high con-
1984b). While special emphasis is accorded the as- centration of usnic acid and atranorin in the cortex.
cus structure, apothecial ontogeny, spore develop- We have reevaluated these characters and, although
ment, and paraphysis morphology within the crus- we are substantially in agreement, a more detailed
tose groups, these characters are relatively uniform understanding of several unique features has been
within the larger foliose and fruticose families. uncovered--namely the rudimentary nature of the
Within Parmelia s.l. a number of workers (Culber- pored epicortex and the unique distribution of the
son & Culberson 1968, 1981; Esslinger 1978; Hale secondary metabolites within the thallus.
1974a,b,c, 1976; Krog 1982; Sipman 1980) segre- Upon undertaking our present study we soon re-
gated groups from Parmelia s.s. during the period alized that the morphological and chemical varia-
1968-1982, based on morphological, chemical, and tion within the genus was much more extensive than
geographical, as well as anatomical characters. had previously been appreciated, and we now con-
However, further investigations have shown that sider that a number of new species should be rec-
even these segregates were heterogeneous and con- ognized. These variations and the new species are
sequently further new genera have been proposed described here.
by Elix et al. (1986b), Elix and Hale (1987), Hale
(1984a, 1985, 1986a,b, 1988, 1989a,b), Hale and MATERIALS AND METHODS
Fletcher (1990), and Lumbsch et al. (1988), one of This studywas basedprimarilyupon collectionsin ASU,
which was Psiloparmelia (Hale 1989a). s, TNs,and ups, the herbariawith the best representation
0007-2745/92/377-391 $1.65/0

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378 THE BRYOLOGIST
of materialsfrom the parts of South America in which
Psiloparmeliais most abundant. Other specimens were
borrowed from BER, CANL, GB, GZU, HBG, KOELN,LILLO, M,
and w, and the privateherbariaof J. Redon and K. Kalb.
In all, 189 herbariumspecimenswerestudied,all of which
were analyzed by thin-layerchromatography(TLC) and
approximately50%by high-performanceliquid chroma-
tography(HPLC). For each species we cite at least one
specimen from every province from which materialwas
seen.
All the major phenolic metabolites encounteredwere
readilyidentifiedby the TLCmethod describedelsewhere
(Elix et al. 1987, 1988), as were the most common fatty
acids, constipatic acid, protoconstipaticacid, dehydro-
constipaticacid, and pertusaricacid (Chester& Elix 1979;
Elix et al. 1986a). The contrastingchemistryof medullary
and corticaltissue was determinedby extractionof tissues
after microdissectionand subsequentTLC analysis, for
several representativecollections of each species (when
available).However,it provedextremelydifficultto com-
pletely segregatethe uppermostmedullarylayer from the
cortical tissue, so these are treatedjointly. Further,it is
possible that traces of fatty acids could have been over-
looked using this procedure.Substancesgiving positive
spot tests (e.g.,fumarprotocetraricacid, salazinicacid, and
norsticticacid) were also detecteddirectlyby application
of PD or K.
In addition a numberof specimenscontainedtracesof
fatty acids which were less well resolved and for which
criticalcontrolswere not available.TLC analysesof these
substances was difficult, and as our analyses were per-
formed over a long period of time, plate-to-platecom-
parisonswere not possible. A detailed chemical study of
these compounds would exceed the scope of the present
research;consequentlythese compounds are simply re-
corded as "unknown"fatty acids.
CURRENTCONCEPTS
According to our current concepts there are four
generic segregates ofParmelia s.l. which are broadly
lobate, have a pored epicortex, are yellow-green in
color (with usnic acid being predominant in the up-
per cortex), and lack pseudocyphellae. These in-
clude Arctoparmelia Hale [typified by A. centrifuga
(L.) Hale], Flavoparmelia Hale [typified by F. cape-
rata (L.) Hale], Psiloparmelia Hale [typified by P.
distincta (Nyl.) Hale], and Xanthoparmelia Hale
[typified by X. conspersa (L.) Hale].
The genera Arctoparmelia and Psiloparmelia are
obligately saxicolous as are the large majority of
species of Xanthoparmelia, but a significant mi-
nority of the Xanthoparmeliae are terricolous. Fla-
voparmelia species by contrast are primarily cortic-
olous, and although several widespread species (F.
baltimorensis and F. haysomii) are commonly found
on rocks they are rarely confused with the former.
Indeed, in his original circumscription of the genus
Xanthoparmelia, Hale (1974c) included several het-
erogeneous elements within it. He subsequently seg-
regated five well-known arctic species-P. aleuritica
Nyl., P. centrifuga (L.) Ach., P. incurva (Pers.) Ach.,
P. separata Th. Fr., and P. subcentrifuga Oxner-
and accommodated them in a new genus, Arcto-
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[VOL.95
parmelia Hale (Hale 1986a). The main distinguish-
ing characters of Arctoparmelia were the presence
of high concentrations of both usnic acid and atra-
norin in the cortex, presence of Cetraria-type li-
chenan in the cell walls (Common 1991), medullary
alectoronic acid (only one of 406 species of Xan-
thoparmelia contains alectoronic acid as the major
medullary component, i.e., X. alectoronica), and the
velvety lower surface. Xanthoparmelia, by contrast,
rarely contains cortical atranorin (and then only in
traces), the cell walls contain Xanthoparmelia-type
lichenan (Common 1991), and the lower surface is
smooth and shiny.
Hale (1974c) also placed Parmelia distincta in
Xanthoparmelia. This high Andean species is su-
perficially very similar to Arctoparmelia, having a
velvety lower surface, analogous cortical chemistry,
and similar lobe configuration. However, it differs
in several important characters, notably the total
lack of rhizines and the presence of isolichenan in
the cell walls. Furthermore, the common occurrence
of fumarprotocetraric acid in the cortex (rather than
the medulla) and the absence of alectoronic acid
distinguish this species from Arctoparmelia, indeed
from all other Parmeliaceae. Following the discov-
ery of a second species in Lesotho closely related to
P. distincta, Hale (1989a) erected the new genus
Psiloparmelia to accommodate these species.
Like Psiloparmelia, Flavoparmelia contains pre-
dominantly isolichenan in the cell walls but is dis-
tinguished by the broad subrotund lobes, much larg-
er spores (14-20 x 7-10 -,m), and lower surface
with a naked, erhizinate region along the margins
but rhizinate abundantly within (Arctoparmelia and
Xanthoparmelia are effectively rhizinate to the mar-
gins). The major distinguishing features of these four
genera are summarized in Table 1.
MORPHOLOGY
GENERICCHARACTERS
The morphologicalfeaturesof Psiloparmeliaare
typically parmeliaceousbut for two unusual fea-
tures:the velvety lowersurfacewithoutrhizinesand
a rudimentaryepicortex. The upper cortex of this
genusis unique.AlthoughHale (1973) reportedthis
to consist of a typical palisade plectenchymawith
a thin, pored epicortex this was an oversimplifica-
tion. For the most partthe upper surfaceis nonepi-
corticate,but in some areashas a rudimentarypored
epicortex(Lumbschet al., in prep.).
The lower surface of Psiloparmeliais also very
unusual, being minutely papillate and appearing
velvety and completely lacking rhizines. The color
of the lower surface is darker (or black in most
species)towardthe centerand palerat the margins.
The majority of species have a finer, velvety yel-
1992] ELIX & NASH: PSILOPARMELIA 379

TABLE1. Comparisonof the chemistry,distribution,ecology, and morphologyof Arctoparmelia,Flavoparmelia,

Psiloparmelia,and Xanthoparmelia.

Arctoparmelia Flavoparmelia Psiloparmelia Xanthoparmelia

Epicortex Pored Pored Subpored Pored
Cell walls contain Cetraria-type Isolichenan Isolichenan Xanthoparmelia-type
lichenan lichenan
Corticalsubstances Usnic acid*, atra- Usnic acid (major), Usnic acid, atra- Usnic acid, atranorin(?
norin atranorin(minor) norin, f-orcin- traces)
ol depsidones,
and depsides
Lower surface Velvety Smooth, shiny Velvety Smooth, shiny
Rhizines Present Present Absent Present**
Spores 10-12 x 4-6 Atm 14-20 x 7-10 /tm 10-12 x 6-8 /tm 6-10 x 4-6 /tm
Conidia Bifusiform Bifusiform-fusiform Bifusiform Bifusiform-bacilliform
Substrate Saxicolous Corticolous,saxico- Saxicolous Saxicolous
lous
Geography Arctic Cosmopolitan Southernhemi- Cosmopolitan
sphere
Ecology Arctic-boreal Temperate High montane Temperate-subarid
Medullarychemistry Alectoronicacid Fatty acids,3-orcinol Fatty acids Varied
depsides, and
depsidones
* Lackingin A. aleuritica.
** Rhizines absent in 6 of 406 species.

lowish-gray (or mouse-gray) marginal zone which
is particularly characteristic. This minutely papil-
late lower surface structure is well depicted in the
SEM photographs published by Hale (1989a).
SPECIES CHARACTERS
Vegetative propagules. --Isidia are unknown but
soredia are relatively common in Psiloparmelia, and
are an important character for distinguishing taxa
at the species level.
Three sorediate species have been recognized: P.
arhizinosa, P. pustulata, and P. sorediosa. All three
become sparsely to moderately pustulate, and at
length the pustules erupt into coarsely sorediate
masses up to 1 mm in diameter.
The concept of pairs of species that are chemically
identical and morphologically similar-except for
the production of vegetative propagules by one-
has received considerable attention (Poelt 1970,
1972; Tehler 1982). Within Psiloparmelia there are
three chemically identical pairs of species differing
primarily by the presence or absence of soredia: P.
arhizinosa and P. flavobrunnea, P. denotata and P.
pustulata, and P. norstictica and P. sorediosa. How-
ever, it should be emphasized that these are not all
morphologically identical but show various degrees
of differentiation in addition to the presence or ab-
sence of soredia.
Color of lower surface.- The lower surface of the
thallus in Psiloparmelia varies from tan to jet black
and such color differences may be a significant tax-
onomic character. The color of the lower surface is
usually invariant toward the center of the thallus;
of the 11 species, nine have a black lower surface
and two have a pale-tan or brown lower surface.
Lobe configuration. - Lobe configuration and the
degree of adnation are important characters for sep-
arating several species of Psiloparmelia. Extremes
of development may be observed by comparing P.
dichotoma (with narrow, dichotomously divided,
linear-elongate lobes ca. 1 mm wide and up to 5
mm between lobe branches) and P. distincta (with
broad, irregularly branched, rotund lobes up to 10
mm wide). However, a number of species do have
a highly plastic morphology and their lobe config-
uration may be significantly influenced by environ-
mental factors. Even P. distincta may develop la-
ciniae within the thallus that are narrower and more
elongate, and ultimately become densely imbricate,
building up the thallus into a thick mat, and forming
an apparently different overall thalline morphology.
In summary the majority of species described in
the present revision are clearly distinguished by two
or more independent characters (e.g., the presence
or absence of soredia, medullary chemistry, lobe
configuration, adnation of the thallus, and color of
the lower surface).
CHEMISTRY
Cell- wall polysaccharides. --While secondary
products are often useful taxonomically at the spe-
cific and generic levels in lichens, polysaccharide
content is often diagnostic for larger phylogenetic
units (Common 1991; Shibata 1973a,b). Polysac-
charides have a fundamental role in the biochem-
istry of fungi, and tend to be conservative features

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380 THE BRYOLOGIST
in their evolution. Some polysaccharides are taxo-
nomically significant at the highest levels of clas-
sification. For example the presence of chitin, chi-
tosan, or cellulose in the cell wall is a feature which
helps define the classes of fungi (Bartnicki-Garcia
1968). Similarly, in the class Oomycetes, Aronson
and coworkers have shown that a biochemical di-
chotomy exists with respect to hyphal wall com-
position between Rhipidiaceae and Leptomitaceae,
the two families comprising the order Leptomitales
(Aronson 1977; Aronson et al. 1967). Furthermore,
they found that these biochemical differences par-
alleled the traditionally accepted morphological and
anatomical differences between these families
(Aronson 1977). In a similar vein, Shibata and co-
workers (Shibata 1973b) showed that pustulan is a
characteristic polysaccharide in lichens of the Um-
bilicariaceae (Umbilicaria and Lasallia), and that
glycopeptides are important cell wall components
of the Lobariaceae. The taxonomic utility of this
criterion in the Parmeliaceae was developed by Im-
shaug and Common (Common 1991; Imshaug
1981), who recognized four major groups - contain-
ing either isolichenan, Xanthoparmelia-type lichen-
an, Cetraria-type lichenan, or an intermediate-type
lichenan. Chemically these polysaccharides differ pri-
marily in the stereochemistry of the glycosidic bonds,
being largely f3 in lichenan and a in isolichenan.
Xanthoparmelia-type lichenan, Cetraria-type lich-
enan, and the intermediate-type lichenan differ pri-
marily in their staining properties with various io-
dine reagents; the structural features responsible for
these differences have yet to be elucidated (Com-
mon 1991). The utility of this character is readily
demonstrated by application to related but well ac-
cepted genera; for example Hypogymnia contains
Cetraria-type lichenan whereas Menegazzia con-
tains isolichenan.
In the present context this character can be used
as one of the primary discriminators to differentiate
Psiloparmelia and Flavoparmelia (containing iso-
lichenan) from Arctoparmelia (Cetraria-type lichen-
an) and Xanthoparmelia (Xanthoparmelia -type
lichenan) (see Table 1).
Chemotaxonomy within Psiloparmelia.--Given
the number of species involved, the chemistry of
Psiloparmelia is quite diverse with eight combina-
tions of taxonomically important secondary prod-
ucts, and has proved to be of value here as in many
other groups of parmelioid lichens. As most chem-
ically characterized populations have morphologi-
cal or distributional characteristics that justify their
ranking as species, a chemotaxonomic approach has
been adopted. The eight diagnostic combinations of
secondary metabolites, and the species of Psilopar-
melia belonging to each combination, are: 1). Con-
stipatic acid (major), protoconstipatic acid (major),
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[VOL. 95
dehydroconstipatic acid (?), pertusaric acid (?),
usnic acid, atranorin-P. arhizinosa, P. dichotoma,
P. flavobrunnea. 2). Diffractaic acid (major), bar-
batic acid (minor-trace), obtusatic acid (minor-
trace), 4-O-demethylbarbatic acid (minor-trace),
usnic acid, atranorin-P. diffractaica. 3). Fumar-
protocetraric acid (major), protocetraric acid (trace),
usnic acid, atranorin-P. distincta. 4). Hypostictic
acid (major), hypoconstictic acid (minor-trace),
hyposalazinic acid (minor-trace), usnic acid, atra-
norin-P. hypostictica. 5). Isousnic acid, atranorin,
unknown fatty acids-P. subcrustosa. 6). 4-0-
Methylhypoprotocetraric acid (major), notatic acid
(minor-trace), isonotatic acid (minor), hypoproto-
cetraric acid (trace), usnic acid, atranorin-P. den-
otata, P. pustulata. 7). Norstictic acid (major), con-
norstictic acid (minor), usnic acid, atranorin-P.
norstictica, P. sorediosa. 8). Salazinic acid (major),
consalazinic acid (minor-trace), usnic acid, atra-
norin-P. salazinica.
The secondary metabolite chemistry of Psilopar-
melia is characterized by a series of /-orcinol de-
rivatives and by fatty acids. However, the distri-
bution of the former compounds appears to be
unusual in this genus-although the fatty acids ap-
pear to be confined to the medulla, the phenolic
substances appear to originate in the uppermost
medullary layer and to diffuse into the upper cortex.
A total of 20 phenolic substances has been detected;
the large majority are f-orcinol depsidones while
f-orcinol depsides are characteristic of one species.
Four species lack f-orcinol derivatives and contain
only fatty acids.
The aliphatic acids--constipatic acid and proto-
constipatic acid-are also very common in Psilo-
parmelia, but in a number of species they appear
to be accessory in nature, that is they are substances
that occur sporadically in these species (e.g., P. dis-
tincta), usually in addition to other invariant con-
stituents and have no correlation with morpholog-
ical or distributional variations. Other related but
less common fatty acids include dehydroconstipatic
acid and pertusaric acid as well as a number of
unidentified aliphatic acids. These substances com-
monly occur as accessory compounds in a number
of species and vary in quantity from deficiency to
abundance. They exhibit similar accessory occur-
rence in a number of Xanthoparmeliae (Elix et al.
1986a).
However, these compounds are the only sub-
stances present in P. arhizinosa, P. flavobrunnea, P.
dichotoma, and P. subcrustosa, so in a formal man-
ner these species could be considered "acid defi-
cient" species containing only accessory metabo-
lites. Although they appear to show minor but
consistent morphological differences, P. denotata,
P. flavobrunnea, P. hypostictica, P. norstictica, and
1992] ELIX& NASH:PSILOPARMELIA 381

Progenitor

O-methylation
AcidDeficient acid
4-O-methylbarbatic

diffractaicacid

O-methylation
acid
hypoprotocetraric

oxidationat C9 4-O-methylhypo-
acid
protocetraric

norsticticacid acid
Protocetraric

oxidationat C6
oxidation esterification
oxidationat C9 at C6C4
hyposticticacid

salazinicacid acid
fumarprotocetraric

FIGURE 1. Postulatedbiosequentialrelationshipsof the secondarymetabolitesdetected in the lichen genus Psilo-

parmelia.

P. salazinica could be considered chemotypes of P.
distincta in a broad sense. Similarly P. sorediosa
could be considered a chemotype of P. arhizinosa.
Biosequential relationships. -There have been
relatively few biosynthetic studies on lichens, but
evidence from those that have been studied and
pathways to similar products in nonlichen-forming
fungi and higher plants allow us to construct a pre-
liminary biogenetic hypothesis. From such simple
hypotheses it is possible in some cases to postulate
biosynthetic sequences (Elix 1991).
Some biosequential relationships are self evident;
thus 1). O-methylated derivatives are considered
more highly derived than the corresponding hy-
droxy compounds, e.g., methylation of norstictic
acid will give stictic acid. 2). C-hydroxylated deriv-
atives are considered more highly derived than the
corresponding parent compounds, e.g., hydroxyl-
ation of lecanoric acid will give diploschistesic acid.
3). Esterified products are considered more highly
derived than the parent hydroxy compounds, e.g.,
fumarprotocetraric acid is considered more highly
derived than protocetraric acid. 4). Sequential ox-
idations are also common in lichens, with the more
highly oxidized substrate representing the more
highly derived secondary metabolite, e.g., oxidation
of norstictic acid will give the more highly derived
salazinic acid. 5). C-methylated derivatives are con-
sidered more highly derived than the corresponding
parent compounds, e.g., 4-O-demethylbarbatic acid
is considered more highly derived than lecanoric
acid.
If one accepts the validity of such biosequential
relationships, it is then possible to postulate prim-
itive and derived chemical characters. Biogeneti-
cally, all these fl-orcinol derivatives are closely re-
lated; the biosynthetic sequence to the major
secondary metabolites can be rationalized in the
manner depicted in Figure 1.
BIOGEOGRAPHY
Eleven of 12 species of Psiloparmelia recognized
in this revision are confined to high elevations of
the Andes Mountains of South America from Ec-
uador and Peru, south to Bolivia and into northern
Argentina. Indeed they constitute the most com-
monly collected foliose lichens on rocks at high al-
titudes in this region. The twelfth species, P. arhi-
zinosa, occurs in alpine habitats at high elevation
(3,000-3,300 m) in Lesotho, southern Africa. It is
possible that the latter sorediate species may have
originated from South America via readily dissem-
inated asexual propagules (Culberson 1972). Al-
though P. arhizinosa has yet to be found in South
America, subsequent chemical evolution on either
continent could explain the chemical disparity of
the morphologically very similar species P. arhizi-
nosa and P. sorediosa.

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382 THEBRYOLOGIST [VOL.95

THE GENUSPSILOPARMELIA 11. Lobes narrow,0.8-1.2 mm wide, containingdif-

fractaic acid ............................ 4. P. DIFFRACTAICA
Hale, Mycotaxon 35: 42. 1989.
PSILOPARMELIA 11. Lobesbroader,1.0-2.5 mm wide,containing4-0-
acid .........- 2. P. DENOTATA
methylhypoprotocetraric
Type species:Psiloparmeliadistincta(Nyl.) Hale.
Parmelia sect. Xanthoparmeliasubsect. Endocoerulea 1. PSILOPARMELIA ARHIZINOSAHale, Mycotaxon 35:
Gyelnik, Feddes Repert. Sp. Nov. Reg. Veg. 29: 282. 43. 1989. (FIG. 2)
1931.
TYPE:LESOTHO.On dolerite ledges 16.6 km north-
Type species of subsection:Parmeliaboulyde lesdainii west of Sani Pass at the west side of KotisepholaPass (on
["lesdaini"]Gyel. roadto MotengPass),3100 m, Grid2929 AC,Hale 81411,
Thallus foliose, dorsiventral, heteromerous, lo- 5 May 1988 (us, holotype; LD, PRE,isotypes).
bate, orbicular, 3-15 cm diam., tightly to loosely Thallus tightly adnate to adnate on rock, 3-7 cm
adnate, saxicolous; lobes subirregular to sublinear- broad, pale yellow-green; lobes subirregular, 0.8-
elongate, apically rotund to narrow and incised, 2.0 mm wide, irregularly dichotomously branched,
margins without cilia; upper cortex of palisade plec- subimbricate; upper surface continuous, dull, emac-
tenchyma covered in part by a rudimentary pored ulate, faintly to strongly white pruinose, rugulose
epicortex; upper cortex smooth or wrinkled, pru- with age, sparsely to moderately pustulate, the pus-
inose, pustules or soralia present or absent, pseu- tules large, erupting into coarsely sorediate masses
docyphellae and isidia absent; cell walls containing to 1 mm diam.; medulla white; lower surface plane,
isolichenan; medulla white; photobiont green; lower finely felted-velvety, dull, brown to dull mineral-
surface black, velvety, minutely papillate but lack- gray at center, pale to ivory-colored at tips, rhizines
ing rhizines. Apothecia adnate to substipitate, 2-5 lacking. Pycnidia and apothecia not seen.
mm diam., spores 8 per ascus, spherical to elliptical, Chemistry. -Cortex K+ pale yellow, C-, PD-;
9-12 x 5-9 4m. Pycnidia immersed, conidia bi- medulla K-, C-, KC-, PD-. Atranorin (major)
fusiform, 5-7 x 0.5 inm. and usnic acid (major) in the cortex and upper me-
dulla, protoconstipatic acid (major) and constipatic
KEY TO THE SPECIES OF PSILOPARMELIA acid (major) in the medulla.
1. Upper surfacesorediate-pustulate......................... 2 Additionalspecimenexamined.- LESOTHO.56.1 km
1. Upper surfacelackingsorediaand pustules 4 N of Mapholanengon Sani-MotengPassroad,Hale 81440
2. Lower surfacebrown, lacking4-O-methylhy- ...............
(ASU).
poprotocetraricacid ........................................ .... 3
2. Lower surface black, with 4-O-methylhypo- This unusual species occurs in high-elevation al-
protocetraric acid ........................ 9. P. PUSTULATA pine habitats (3,000-3,300 m) in Lesotho, usually
3. Lobes sublinear,with norsticticacid (cortex K+ on the sheltered lower sides of dolerite ledges which
red) 11. P. SOREDIOSA
3. Lobes...................................................................................
subirregular,lackingnorsticticacid (cortex receive little if any direct rainfall. It has a peculiar
K-)
.....................
......... 1..... . P. ARHIZINOSA grayish velvety lower surface without rhizines, a
4. Thallus loosely adnate, lobes linear-elongate, rugose, pruinose surface, and coarse soralia.
dichotomously divided 3. P. DICHOTOMA
4. Thallusadnateto tightly..................
adnate,lobes sublin-
ear, subirregularlydivided ........................... 5 2. PSILOPARMELIA DENOTATA Elix & Nash sp.
5. Cortex PD+ orange-red.................................... .... 6 nov. (FIG. 3)
5. Cortex PD - ............ 8
........................................
6. CortexK-, containingfumarprotocetraric acid Species cum thallo ut in Psiloparmelia distincta sed ab
5 . P . DISTINCTA hac specie acidum 4-O-methylhypoprotocetraricum con-
............................................................................................
6. CortexK+ yellow-red,containingsalazinicor tinente differt.
norsticticacid ... 7
7...... HOLOTYPE: - PERU. DEPT.LIMA.PROV.HUAROCHIRI: On
7. Lobes broad, 1.5-2.5 ........................................................................
mm wide, usuallylobulate- rocks, valley of rio Santa Eulalia, NE of Carampoma, c.
laciniate,containingnorsticticacid ......................... 3700 m, 11038'S, 76027'W, Santesson & Moberg P24:4A,
8 . P . NORSTICTICA 12th RegnellanExpedition, 15 February1981 (s).
.............................................................................................
7. Lobes 0.7-1.5 mm lobulae
narrow, wide, lacking
and lacinae, containingsalazinicacid Thallus adnate to loosely adnate, 5-10 cm diam.,
10.............................
P . SALAZINICA pale yellow-green; lobes subirregular, irregularly
..............................................................................................
8. CortexK+ palered,containinghyposticticacid 1.0-2.0 mm wide, occasionally becoming
7 . P . HYPOSTICT ICA branched,
..................................................................................
8. Cortex K-, lackinghyposticticacid .......................... 9 densely laciniate, laciniae imbricate, building up the
9. Thallus subcrustose,containingisousnic acid ........ thallus into a thick mat or very rarely becoming
12. P. SUBCRUSTOSA lobulate, lobules leaf-like, 0.1-0.4 mm wide, con-
9. Thallus foliose, containingusnic acid .......................... 10 vex-distorted; upper surface dull, emaculate, sub-
10. Lobes subirregular,containing 4-O-methyl-
isidia and soredia lacking; medulla
hypoprotocetraricor diffractaicacid .................... 11 apically shiny,
to
10. Lobessublinear linear,containingfatty ac- white, dense and waxy in texture; lower surface plane,
ids ........................................
.. 6. P. FLAVOBRUNNEAdull, velvety, yellow to pale gray at margins but

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1992] ELIX& NASH: PSILOPARMELIA 383

23
boA-

of

q 10,~
I rdo 4A
-At'
J4
A~~p? I
V ~
AW. ?

,Ile

.0

?~J 4

i """"""""""""""O"

C,?101 1~

FIGURES2-5. Psiloparmelia. - 2. P. arhizinosa, Hale 81440 (ASU). - 3. P. denotata, type, R. Santesson & R.
Moberg P24:4A (s). - 4. P. dichotoma, type, H. Kashiwadani 21630 (TNs). - 5. P. diffractaica, type, T. H. Nash
27856A (ASU).Scale bars = 5 mm.

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384 THEBRYOLOGIST [VOL.95

black within, rhizines lacking. Pycnidia immersed, This species is distinguished by the loosely adnate
commonly produced; conidia bifusiform, 0.5 x 5- thalli with very narrow, linear-elongate, dichoto-
6 tm. Apothecia adnate, 1.0-3.0 mm diam.; spores mously divided lobes, black margins of the lobes,
subspherical to elliptical, 5-6 x 7-9 qm. and lack of cortical depsidones (PD-).
Chemistry.-Cortex K-, C-, PD-; medulla K-,
C-, KC-, PD-. Usnic acid (major), atranorin (mi- 4. PSILOPARMELIA Elix & Nash sp.
DIFFRACTAICA
nor), chloroatranorin (minor), 4-O-methylhypopro- nov. (FIG. 5)
tocetraric acid (major), isonotatic acid (minor), no-
Species cum thallo ut in Psiloparmeliadistinctased ab
tatic acid (trace), hypoprotocetraric acid (minor- hac specie thallo diminuto et acidum diffractaicumcon-
trace), unknown (minor) in the cortex and upper tinente differt.
medulla, fatty acids (+ trace) in the medulla. HOLOTYPE:ARGENTINA.SALTAPROVINCE. On acidic
nine specimensexamined.-AR- rocks, 14 km W of pass throughCumbres de Obispo along
Representativesof the
GENTINA. SALTA.14 km W of pass through Cumbres route 33, 65056'W,25014'S,c. 3000 m, T. H. Nash27856A,
de Obispoalongroute13,Nash27856 (ASU). PERU.Cuzco. 25 May 1989 (ASU).
Canchis,aroundAbraLa Raya,betweenSicuaniand Aya- Thallus adnate, forming small rosettes 1-2 cm
viri, Kashiwadani22158 (TNs);Quispicanchis,between
Abra Hualla Hualla and Mercapata,Kashiwadani21560 diam., pale yellowish-green; lobes subirregular, ir-
(TNs).PUNo. Azaingaro,Jarjani,about 7 km SW of Asillo, regularly branched, narrow, 0.5-1.0 mm wide (to
Kashiwadani22360 (TNs);Lampa,CaraCara,nearPucarai, 2.0 mm at apices), contiguous but hardly imbricate;
Kashiwadani22324 (TNs);Jarjani,aroundLagunaUmayo, upper surface dull, emaculate, becoming white pru-
Sillustani, c. 20 km NW of Puno, Kashiwadani22213 inose near
apices, isidia and soredia lacking; me-
(TNS).
dulla white; lower surface plane, dull, velvety, gray-
This species is distinguished by thalli composed black to black, mouse-gray to dark-gray at margins,
of densely imbricate lobes which form a thick mat, rhizines lacking. Pycnidia immersed, common; co-
and the presence of 4-O-methylhypoprotocetraric nidia bifusiform, 0.5 x 5-6 gm. Apothecia not seen.
acid and isonotatic acid in the cortex. Chemistry. -Cortex K-, C-, PD-; medulla K-,
C-, KC-, PD-. Usnic acid (major), atranorin
(trace), diffractaic acid (major), obtusatic acid (mi-
3. PSILOPARMELIA DICHOTOMA Elix & Nash sp.
nor), and 4-O-demethylbarbatic acid (trace) in the
nov. (FIG. 4) cortex and upper medulla; an unknown (trace) in
Speciescum thallo ut in Psiloparmeliaflavobrunnea sed the medulla.
ab hac specie thallo laxe adnato, lobis lineariter-elonga- Distinguished by its small adnate rosette-forming
tibus, separatiset dichotome ramosis. thalli and the medullary diffractaic acid, at present
HOLOTYPE: PERU. DEPT.CUZCO. PROV. QUISPICANCHIS: P. diffractaica is known only from the type collec-
Abra HuallaHualla,in upperpartof "Puna"zone, 4500- tion.
4600 m, H. Kashiwadani21630, 14 September1984 (TNs).
Thallus loosely adnate, 4-8 cm diam., pale yel- 5. PSILOPARMELIA
DISTINCTA(Nyl.) Hale, Mycotax-
low-green but blackening toward center; lobes nar- on 35: 42. 1989. (FIG. 6)
row, linear-elongate, dichotomously divided, 0.4-
1.0 mm wide, not at all imbricate; upper surface Parmelia distinctaNyl., Ann. Sci. Nat. Bot., ser. 4, 15:
dull, emaculate, with conspicuous black margins, 374. 1861. Xanthoparmeliadistincta(Nyl.) Hale, Phy-
isidia and soredia lacking; medulla white, dense and tologia 28: 487. 1974. TYPE:BOLIVIA. PROV.YUNGAS.
Weddell s.n. (H-NYL,Nyl. Herb. No. 34819, lectotype
waxy in texture; lower surface plane, dull, velvety, selected by Gyelnik 1938).
mouse-gray at tips but black within, rhizines lack- Parmelia subcongruensMiill. Arg., Flora 72: 64. 1889.
ing. Pycnidia immersed, commonly produced; co- Parmelia distinctafo. subcongruens(Miull.Arg.)Gyel.,
nidia bifusiform, 0.5 x 5-6 gm. Apothecia not seen. Ann. Hist.-Nat. Mus. Nat. Hung. 31: 26. 1938. TYPE:
ARGENTINA. Lorentz & Hieronymus s.n. (M, lecto-
Chemistry. -Cortex K+ pale yellow, C-, PD-;
type; G, w, isolectotypesselected by Hale 1989a).
medulla K-, C-, KC-, PD-. Atranorin (major) Parmeliaboulyde lesdainiiGyel., FeddesRepert.Sp. Nov.
and usnic acid (major) in the cortex and upper me- Reg. Veg. 29: 281. 1931. Parmelia subcongruens var.
dulla, protoconstipatic acid, constipatic acid, and bouly de lesdainii(Gyel.) Gyel., Ann. Hist.-Nat. Mus.
an unknown fatty acid in the medulla. Natl. Hung. 29: 21. 1935. Parmelia distincta fo. bouly
de lesdainii (Gyel.) Gyel., Ann. Hist.-Nat. Mus. Natl.
Representativesofthe six specimensexamined.- PERU. Hung. 31: 26. 1938. TYPE:PERU. Haciendad'Angas-
Cuzco. Quispicanchis,Abra HuallaHualla,Kashiwadani murca, 160 km from Trujillo, Libertad,Standaerts.n.
21491 (TNs);Paucartambo,Oropesa-Paucartamboroad, (BP,lectotypeselected by Hale 1989a).
betweenHuancaraneand Sayllapata,Santessonet al. P95: Thallus adnate to loosely adnate, leathery, 5-10
53 (ups). HuANUCO.Dos de Mayo, valley of rio Marafion,
c. 58 km (roaddistance)WNW of Huanuco,Santesson& cm diam., light yellowish-green; lobes subirregular
Moberg P50:42 (s). to sublinear, irregularly branched, 0.8-3 mm wide,

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1992] ELIX& NASH: PSILOPARMELIA 385

t. I-
d•

6
f-
-ioi
_• 440
itt'I c- ,3

PI . '4"

'rI
' "•, t
r,
bt.••"
40 1;3? ,.iS
....~a
.• '"',"?
A'w &; 1 A

r-fo

/ M6-W

*OWN

"i'Opt. 10t
) logo
r0
"- Ole.
•.- .k ._ t,

FIGURES6-9. Psiloparmelia. - 6. P. distincta, T. H. Nash 27835 (Asu). - 7. P. flavobrunnea, H. Kashiwadani

21491 (TNs). - 8. P. hypostictica, type, T. H. Nash 27941 (ASU). - 9. P. norstictica, type, T. H. Nash 27930A (Asu).
Scale bars = 5 mm.

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386 THE BRYOLOGIST [VOL. 95

imbricate, rarely becoming laciniate or lobulate type of P. distincta. Hale subsequently followed this
within, lobules leaf-like, 0.1-0.4 mm wide, plane; lectotypification (Hale 1989a) as we have done. Like
upper surface dull, emaculate, becoming coarsely that of other Psiloparmelia species, the chemistry
white pruinose with age, transversely cracked and of P. distincta is highly unusual, not so much in the
rugulose at center, isidia and soredia lacking; me- contents but in their distribution within the thallus.
dulla white or darkening somewhat, dense and waxy Thus P. distincta contains fumarprotocetraric acid,
in texture; lower surface plane, dull, velvety, yel- and this depsidone appears to be confined largely
lowish-gray at margins but turning jet black toward to the cortex which reacts PD+ orange, while the
center, rhizines lacking. Pycnidia immersed, com- medulla is P-. The closely related species P. fla-
mon; conidia bifusiform, 0.5 x 5-6 gm. Apothecia vobrunnea, however, is PD- in the cortex and con-
adnate to substipitate, 2.0-5.0 mm diam.; spores tains only the fatty acids. The types of Parmelia
subspherical to elliptical, 6-9 x 10-12 ,m. distincta (nos. 34819 and 34820), P. bouly de les-
Chemistry. -Cortex K+ pale yellow or K-, C-, dainii, and P. subcongruens all react PD+ red and
PD-; medulla K-, C-, KC-, PD+ orange-red. contain fumarprotocetraric acid; the third syntype
Usnic acid (major), atranorin (minor), fumarpro- of P. distincta (no. 34818) is PD- and is chemically
tocetraric acid (minor), succinprotocetraric acid (? identical with the type of P. flavobrunnea. Gyelnik
minor-trace), malonprotocetraric acid (? minor- (1931) also used P. bouly de lesdainii as the type for
trace), and protocetraraic acid (? trace) in the cortex his Parmelia sect. Xanthoparmelia subsect. Endo-
and upper medulla; protoconstipatic acid (? ma- coerulea Gyeln. The description of the subsection,
jor), constipatic acid (? minor), unknown fatty ac- "medulla thalli coerulea," was apparently based on
ids (? trace), and ursolic acid (? trace) in the me- the waxy, sometimes darkening medulla in some
dulla. specimens, but he later (Gyelnik 193 5) amended the
description of this subsection to read "erhizinosus."
Representativesof the 110 specimensexamined.-AR-
GENTINA. CATAMARCA. Nevados de Ancoqija, valley of Psiloparmelia distincta is probably the most com-
rio Pisavil, Lamb 5515 (LILLO).CORDOBA.14 km S of monly collected foliose lichen at high elevations in
Candelariaalong route 15, Nash 28152 (ASU).Jujuv. 28 the Andes Mountains from Ecuador and Peru, south
km N of Humahuacaalong route 9, Nash 27929 (ASU). to Bolivia and into northern Argentina. It is not
SALTA.Valle Encantadoin the Sierrade Candado,Nash
the most common but also morphologically
27835 (Asu).TUCuMAN. 2 km NW ofTafitdel Valle along only
route 307, Nash 28057 (ASU). the most variable species of Psiloparmelia--from
BOLIVIA.LAPAZ.Aroma,on the highwayfromLa Paz specimens with tightly adnate large-foliose to al-
to Oruro, 6 km N of Calamarca,Puna, Feuerer(HBG); most subcrustose thalli, and from specimens with
BautistaSaavedra,ChacabayanearAmarete,Feuerer(HBG); imbricate lobes to thalli dominated by highly
Los Andes, main road from La Paz to Tiquina, Feuerer barely
15091 (HBG);Murillo, Zongotal, Feuerer (HBG). imbricate secondary lacinae. Despite its morpho-
CHILE. ANTOFAGASTA.Lachimba, Follmann 7735 logical variability this species can be distinguished
(KOELN). chemically by the presence of cortical usnic acid,
ECUADOR. AZUAY.ParqueNacional Las Cajas, 37.5 atranorin, and fumarprotocetraric acid, and med-
km W of Cuenca on road to Malleturo,Arvidssonet al.
(GB). CHIMBORAZO. 20 km NNE of Tixan, Nash 23799 ullary fatty acids.
(ASU).COTOPAXI. W slope of VulcanCotopaxi,Nash 23848
(ASU). 6. PSILOPARMELIA FLAVOBRUNNEA (Miill. Arg.) Elix
PERU. ANCASH. Huaraz,LagunaLlaca,NE of Huaraz,
Santesson&MobergP60:64 (s);Yungay-Llanganuco road, & Nash comb. nov. (FIG. 7)
22 km NE of Yungay, Santesson & P55:101
Moberg (s). Parmelia
Cuzco. Anta, 2-3 km N of Izcuchuca,N of Anta, Santes- flavobrunneaMll. Arg., Flora 74: 379. 1891.
son et al. P81:9 (s); Canchis, around Abra La Raya, be- Xanthoparmeliaflavobrunnea(Mfll. Arg.) Hale, Phy-
tween Sicuani and Ayaviri, Kashiwadani22153 (iNS); tologia 28: 487. 1974. TYPE:PERU. Azangaro,Lechler
1766 (G, lectotype; BM,isolectotype, designated here).
Quispicanchis,Abra Hualla Hualla, Kashiwadani21479
(TNS); Urubamba, valley of rio Piri, 28 km from Ollan- Thallus adnate to loosely adnate, leathery, 5-10
taytambo, Santesson et al. P90:42 & 43 (s). HuANco. cm
Dos de Mayo, valley of rio Marafion,58 km WNW of diam., light yellowish green; lobes sublinear to
Huanuco,Santesson & MobergP50:15 & 43 (s). JUNN. linear or rarely subirregular, subdichotomously to
Tarma, Pampa Huicushcancha, Hegewald (Gzu). LIMA. irregularly branched, 0.8-3 mm wide, imbricate,
Huarochiri,valley of rio Santa Eulalia, NW of Caram- rarely becoming lobulate, lobes bearing marginal
poma, Santesson & MobergP24:4 (ups). Puto. Lampa, lobules and becoming laciniate or lobulate within,
CaraCara,nearPucara,Kashiwadani22329 (TNs);around
mm wide; upper
LagunaUmayo, Sillustani,c. 20 km NW of Puno, Kashi- lobules rounded, leaf-like, 0.1-0.4
wadani22411 (TNS). surface dull, emaculate, becoming coarsely white
pruinose with age, transversely cracked and rugu-
Nylander's herbarium contains three collections lose at center, isidia and soredia lacking; medulla
from Bolivia. Gyelnik (1938) apparently saw only white or darkening somewhat, dense and waxy in
one of these (no. 34819) and designated it as the texture; lower surface plane, dull, velvety, yellowish

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1992] ELIX & NASH: PSILOPARMELIA 387

to mouse-gray at margins but turning jet black to- isidia and soredia lacking; medulla white; lower sur-
ward center, rhizines lacking. Pycnidia immersed, face plane, dull, velvety, pale tan to black, mouse-
commonly produced; conidia bifusiform, 0.5 x 5- gray to dark-gray at margins, rhizines lacking. Pyc-
6 Am. Apothecia adnate to substipitate, 2.0-5.0 mm nidia immersed, common; conidia bifusiform, 0.5
diam.; spores subspherical to elliptical, 6-9 x 10- x 5-6 Am. Apothecia adnate, 1.0-1.5 mm diam.;
12 am. spores subspherical to elliptical, 5-7 x 8-10 Am.
K-, C-, PD-; medulla K-, Chemistry. -Cortex K+ yellow, C-, PD-; me-
Chemistry.--Cortex dulla K-, C-, KC-, PD-. Usnic acid (major),
C-, KC-, PD-. Usnic acid (or rarely isousnic acid)
(major) and atranorin (minor-trace) in the cortex atranorin (minor), hypostictic acid (major), hypo-
and upper medulla; protoconstipatic acid (major), salazinic acid (minor-trace), hypoconstictic acid
constipatic acid (? minor) and unknown fatty acids (minor-trace), and hypoprotocetraric acid (? trace)
(trace) in the medulla. in the cortex and upper medulla; protoconstipatic
acid (?), constipatic acid (?) and unknowns (mi-
Representativesof the 37 specimens examined.-AR-
Nevados de Ancoqija, Quebra-
GENTINA. CATAMARCA. nor) in the medulla.
da de los Cazadores,E of Nevado Pabell6n de la Abra
Representativesof the 10 specimens examined.-AR-
Grande, Lamb 5589 (LILLo).CORDOBA.Partido de Pun- GENTINA. JujuY. W slope of Sierrade Santa Victoria,
tilla, 3 km W of El Durazno,Nash 23972 (ASU).MENDOZA.c. 51 km E of La Quiacaalong route 5, Nash 27973 (Asu).
70 km NW of Mendoza,Nash 23928 (ASU).NEUQUEN. 10 BOLIVIA.LAPAz. Los Andes, 1.6 km past the turnoff
km SE of Caviahue along route 70, Nash 27531 (ASU). to Pefiasand Mina Fabulosaalongthe main highwayfrom
SALTA. 46 km W of top of escarpmentalong route 33, La Paz to Tiquina,Feuerer(HBG); Omasuyos,Sorejapaon
Nash27869 (Asu).TUCUMAN. 17 km NNW of passthrough Lake
Sierra del Aconquija and Cumbres Calchaquies,Nash Feuerer Titicaca, along the road from Huarinato Tiquina,
(HBG).ORURO.Sajama, Nevado Sajama, lado Sur,
28023 (Asu). en laderaN del rio Sururia,Liberman337 (M).
BOLIVIA.LAPAZ.BautistaSaavedra,Pumazani,near
Charazani,Feuerer(HBG); Murillo,Zongo-Tal,Feuerer&
Preiss (HBG). This is a species which is distinguished by the
Lago Chungerdi,Redon 2474 (herb.
CHILE. TARAPACA.
Redon). tightly adnate to adnate thalli and the cortical hy-
Above San Juan, 27 km W
ECUADOR. CHIMBORAZO. postictic acid.
of Riobambaon road to Guaranda,Arvidssonet al. (GB).
COTOPAXI.Rio Saquimale, 3 km along road to Vulcan 8. PSILOPARMELIA NORSTICTICA Elix & Nash sp.
Cotopaxi, Nash 23841 (ASU).NAPO.Between Quito and nov.
Baeza, Kalb (herb. Kalb, ANUC).PICHINCHA.E slopes of (FIG. 9)
CerroIliniza,Arvidsson& Nilson (GB).
PERU. Cuzco. Quispicanchis, Abra Hualla Hualla, hac Speciescum thallo ut in Psiloparmeliadistinctased ab
Kashiwadani21477 (TNs).JUNIN.Yauli, aroundAbraAn- acidum specie lobi plerumquelaciniatibusvel lobulatibuset
norsticticumcontinentediffert.
ticona, Ticlio, Kashiwadani22761 (TNs).LIMA.Huaro-
chiri,betweenHuarochiriand Chumpicocha,Cerrate2036 HOLOTYPE:ARGENTINA. JUJUY PROVINCE. On acidic
(TNs). rocks,28 km northof Humahuacaalongroute9, 65022'W,
23000'S, c. 3300 m, T. H. Nash 27930A, 28 May 1989
Morphologically this species is similar to P. dis- (ASU).
tincta, but differs in being more loosely adnate and
Thallus adnate, 3-4 cm diam., pale yellowish-
developing more elongated lobes. It also differs
chemically in that the upper cortex is PD- because green; lobes sublinear, irregularly branched, 0.5-1.5
mm wide, contiguous but hardly imbricate, very
it lacks the fumarprotocetraric acid typical of the
former taxon. often becoming laciniate or lobulate within, lobules
leaf-like, 0.1-0.4 mm wide, often convex-distorted;
upper surface dull, emaculate, becoming white pru-
7. PSILOPARMELIAHYPOSTICTICAElix & Nash sp. inose near apices, isidia and soredia lacking; me-
nov. (FIG. 8) dulla white; lower surface plane, dull, velvety, gray-
Species cum thallo ut in Psiloparmeliadistinctased ab black to black, mouse-gray to dark-gray at margins,
hac specie thallo arcte adnato et acidum hyposticticum rhizines lacking. Pycnidia immersed, common; co-
continentediffert. nidia bifusiform, 0.5 x 5-6 Am. Apothecia adnate,
HOLOTYPE:ARGENTINA. JUJUYPROVINCE. On acidic 1.0-1.5 mm diam.; spores subspherical to elliptical,
rocks, 36 km north-west of Humahuacaalong route 9, 4.5-5.5 x 7-10 Am.
65023'W,22057'S,c. 3600 m, T. H. Nash 27941, 28 May Chemistry. -Cortex K+ yellow-pale red, C-,
1989 (ASU).
PD+ yellow-orange; medulla K-, C-, PD-. Us-
Thallus adnate to tightly adnate, 4-5 cm diam., nic acid (major), atranorin (minor-trace), norstictic
pale yellowish-green; lobes subirregular to sublin- acid (major), connorstictic acid (trace), and salazinic
ear, irregularly branched, narrow, 0.3-1.0 mm wide, acid (? trace) in the cortex and upper medulla; pro-
contiguous but hardly imbricate; upper surface dull, toconstipatic acid (trace), constipatic acid (? trace),
emaculate, becoming white pruinose near apices, and an unknown fatty acid (? trace) in the medulla.

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388 THEBRYOLOGIST [VOL.95

Additionalspecimensexamined.-ARGENTINA. Jujuv. Species cum thallo ut in Psiloparmeliadistinctased ab

43 km E of La Quiaca,lower part of Sierrade SantaVic- hac specie acidum salazinicumcontinentediffert.
toria, along route 5, Nash 27950 (Asu). HOLOTYPE:ARGENTINA. JuJuY PROVINCE.On acidic
ECUADOR. 17 km N of Riobamba,Nash
23793 (Asu). CI-nmoRAzo. rocks, east slope of the Sierrade SantaVictoria,53 km E
of La Quiacaalong route 5, 65016'W,22008'S,c. 4400 m,
PERU. AroundLagunaUmayo, Sillustani,c. 20
km NW ofPU•o. T. H. Nash 28193, 29 May 1989 (Asu).
Puno, Kashiwadani22193 (TNs).
Thallus adnate, 4-5 cm diam., pale yellowish-
This species is distinguished by the adnate to
green; lobes sublinear, irregularly branched, narrow,
tightly adnate thalli which often become laciniate
0.8-2.0 mm wide, contiguous but hardly imbricate;
or lobulate towards the center, and the cortical nor-
stictic acid. The related species P. salazinica has upper surface dull, emaculate, becoming white pru-
inose near apices, isidia and soredia lacking; me-
broader lobes (0.8-2.0 mm cf. 0.5-1.5 mm), lacks
dulla white; lower surface plane, dull, velvety, brown-
laciniae, and contains salazinic acid.
black to black in center, mouse-gray to dark-gray at
margins, rhizines lacking. Pycnidia immersed, com-
9. PSILOPARMELIA PUSTULATAElix & Nash sp. mon; conidia bifusiform, 0.5 x 5-6 Am. Apothecia
nov. (FIG. 10) adnate, 2.0-5.0 mm diam.; spores subspherical to
elliptical, 4.5-5.5 x 7-9 Am.
Species cum thallo ut in Psiloparmeliaarhizinosased
ab hac specie facie inferiore nigricanteet acidum 4-0- Chemistry. -Cortex K+ yellow-pale red, C-,
methylhypoprotocetraricum continente differt. PD+ yellow-orange; medulla K-, C-, KC-, PD-.
HOLOTYPE: ARGENTINA. MENDOZAPROVINCE.On Usnic acid (major), atranorin (minor), salazinic acid
acidic rocks, Partidode Tunuyan,3 km east of Manzano (minor), consalazinic acid (minor-trace), norstictic
Historico, S side of isolated hill, 33?37'S,69015'W, 1070 acid (? trace), and protocetraric acid (? trace) in
m, T. H. Nash 23949, 7 October 1985 (Asu). the cortex and upper medulla; an unknown fatty
Thallus tightly adnate to adnate on rock, 3-4 cm acid (trace) in the medulla.
broad, pale yellow-green; lobes subirregular, 0.8- Additionalspecimensexamined.--ARGENTINA. CA-
2.0 mm wide, irregularly dichotomously branched, TAMARCA. Nevados de Anconquija,Quebradade los Pa-
subimbricate; upper surface continuous, dull, emac- zadores, Lamb 5552 (LILLO). JUJUY.43 km E of La Quiuca,
to white along route5 to SantaVictoria,Nash27950A(ASU). CHILE.
ulate, faintly strongly pruinose, rugulose TARAPACA.
Chapiquifiaa Murmuntani,Redons.n. (herb.
with age, sparsely to moderately grossly pustulate, Redon).
pustules erupting into coarsely sorediate masses to
1 mm diam.; medulla white; lower surface plane, This species is distinguished by the adnate to
finely felted-velvety, dull, dark brown to black, tightly adnate thalli and the cortical salazinic acid.
mouse-gray at tips, rhizines lacking. Pycnidia and
apothecia not seen. 11. PSILOPARMELIA SOREDIOSA Elix & Nash sp.
nov. (FIG. 12)
Chemistry.--Cortex K+ pale yellow or K-, C-,
PD-; medulla K-, C-, KC-, PD-. Usnic acid Species cum thallo ut in Psiloparmeliaarhizinosased
(major), atranorin (minor), chloroatranorin (mi- ab hac specie thallo laxe adnato, lobi angustioribussub-
nor), 4-O-methylhypoprotocetraric acid (major), linearibuset acidum norsticticumcontinentediffert.
hypoprotocetraric acid (minor-trace), and ? an un- HOLOTYPE:ARGENTINA.NEUQUtN. On acidic rocks,
known orange pigment in the cortex and upper me- Partido de PicuinLeufu, along route 237 above Embalse
dulla; protoconstipatic acid (?) and constipatic acid Ezequiel Ramos Mexia, c. 10 km SSW of Picfin Leufu,
40027'S,69019'W,460 m, T. H. Nash 24180, 2 October
(?) in the medulla. 1985 (ASU).
Additional specimen examined. -ARGENTINA. Thallus loosely adnate on rock, 2-4 cm wide, pale
TUCUMAN. 14 km W of Tafi del Valle along route 307, lobes sublinear, 0.2-1.0(-2.0) mm
Nash 28070 (ASU). yellow-green;
wide, irregularly dichotomously branched, subim-
This new species resembles the southern African bricate; upper surface continuous, dull, emaculate,
species, P. arhizinosa, in the production of pustulate faintly to strongly white pruinose, rugulose with age,
soralia and in configuration of the lobes. However, sparsely to moderately pustulate, pustules erupting
these species differ in the color of the lower surface, into coarsely sorediate masses to 1 mm diam.; me-
which is brown-black to black in P. pustulata but dulla white; lower surface plane, finely felted-vel-
brown to dull mineral-gray in P. arhizinosa. These vety, dull brown to dull mineral-gray at center, pale
species also differ chemically as P. pustulata con- mouse-gray to ivory at tips, rhizines lacking. Pyc-
tains 4-O-methylhypoprotocetraric acid. nidia immersed, common; conidia bifusiform, 0.5
x 5-6 Am. Apothecia not seen.
10. PSILOPARMELIA SALAZINICA Elix & Nash sp. Chemistry. -Cortex K+ yellow-pale red, C-,
nov. (FIG. 11) PD+ yellow-orange; medulla K-, C-, KC-, PD-.

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1992] ELIX& NASH: PSILOPARMELIA 389

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FIGURES10-13. Psiloparmelia. - 10. P. pustulata, type, T. H. Nash 23949 (ASU).- 11. P. salazinica, type, T. H.
Nash 28193 (Asu). - 12. P. sorediosa, type, T. H. Nash 24180 (ASU). - 13. P. subcrustosa, type, Lentner (herb.
Feuerer). Scale bars = 5 mm.

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390 THE BRYOLOGIST
Atranorin (major), usnic acid (major), norstictic acid
(minor), and connorstictic acid (? trace) in the cor-
tex and upper medulla; protoconstipatic acid and
constipatic acid in the medulla.
This new species resembles the southern African
species, P. arhizinosa, in the production ofpustulate
soralia and in the brown lower surface. However,
the thallus of P. sorediosa differs in being loosely
adnate with separate, narrow (0.2-1.0 mm), sublin-
ear lobes rather than being adnate to tightly adnate
with irregular, broader (0.8-2.0 mm), contiguous
lobes as in P. arhizinosa. These species also differ
chemically as P. sorediosa contains norstictic acid
which is lacking in P. arhizinosa. At present P. so-
rediosa is known only from the type collection.
12. PSILOPARMELIA SUBCRUSTOSA Elix & Nash sp. CULBERSON,
nov. (FIG. 13)
Speciescum thallo ut in Psiloparmeliaflavobrunnea sed -
ab hac specie thallo arcteadnato,diminuto, subcrustaceo
et acidum isousnicumcontinentediffert.
PERU. In the of Chacha-
HOLOTYPE: vicinity Arequipa,
ni, 4400-4600 m, A. Lentner,30 August 1978 (HBG).
Thallus subcrustose, tightly adnate, 1-3 cm diam.,
pale yellow-green; lobes congested to subirregular,
0.4-0.8(-1.0) mm wide, plane or convex above,
forming convex areolae in center of thallus; upper
surface dull, emaculate, subapically pruinose, isidia -
and soredia lacking; medulla white, dense and waxy
in texture; lower surface plane, dull, velvety, dark
gray at margins but black within, rhizines lacking.
Pycnidia immersed, commonly produced; conidia
bifusiform, 0.5 x 5-6 qm. Apothecia adnate,' 1.0-
1.5 mm diam.; spores subspherical to elliptical, 4.5-
6 x 7-8 Aim.
Chemistry. -Cortex K-, C-, PD-; medulla K-,
C-, KC-, PD-. Isousnic acid (major), usnic acid
(trace), and atranorin (minor) in the cortex and up-
per medulla; traces of unknown fatty acids in the
medulla.
This species is distinguished by the small sub-
crustose thalli with isousnic acid in the cortex and
the fatty acids in the medulla. At present this species
is only known from the type collection.
ACKNOWLEDGMENTS
We gratefullyacknowledgefinancialsupport from the -
U.S. National Science Foundation grants BSR 8820016
and BSR 8943558 to Arizona State University. The cu-
ratorsof the followingherbariaare thankedfor theirkind
cooperation: BER, CANL,GB, GZU, HBG, LILLO, M, S, TNS,
uPs, w, as are the followinglichenologistswho loaned us
material from their private herbaria: K. Kalb and J. Re- HALE,M. E., JR. 1973. Fine structure of the cortex in
don. We thank W. L. Culberson, Duke University,. and
H. T. Lumbsch, University of Essen, for helpful sugges-
tions on this manuscript.
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