A Lichenological Legacy – Festschrift Thomas H

CBibliotheca Lichenologica 106, 319-351
Bates et al. (eds.), Biomonitoring, Ecology, and Systematics of Lichens
Festschrift Thomas H. Nash III
J. Cramer in der Gebr. Borntraeger Verlagsbuchhandlung, Stuttgart, April 2011
The lichen genus Phyllopsora
(Ramalinaceae) in the West Indies
EINAR TIMDAL
Natural History Museum, University of Oslo, P.O. Box 1172 Blindern, N-0318 Oslo, Norway
(Einar.Timdal@nhm.uio.no)
Abstract: 34 species of Phyllopsora are accepted in the West Indies.
New species are: P. hispaniolae, P. imshaugii, P. teretiuscula, and P.
tobagensis. New combinations are: P. cognata (Nyl.) Timdal and P.
phaeobyssina (Vain.) Timdal. New to the West Indies: P. byssiseda,
P. labriformis, and P. lacerata. The names P. microsperma Müll. Arg.
and P. porphyromelaena (Vain.) Zahlbr. are proposed for P. glabella
(Nyl.) Brako, nom. illeg., and P. albicans sensu Swinscow & Krog,
respectively. The secondary compounds hyperlatolic acid, methyl
barbarate, norsolorinic acid, norstictic acid, perlatolic acid, and
superlatolic acid are reported from the genus for the first time.
Keywords: Lecanorales, lichenized ascomycetes, Neotropics,
taxonomy
Introduction
This study began as a revision of the material of Phyllopsora Müll. Arg. collected
in the West Indies by Henry A. Imshaug and stored in MSC. The material was
collected in the period 1952–1963 and consists of 159 collections (including
immixtures) from 10 countries or islands. Imshaug did not publish on the material
and it was identified only to the genus level.
Phyllopsora is mainly a tropical and subtropical genus consisting of crustose to
squamulose species growing primarily on bark in humid forests. More than 100
species names exist in the literature; 15 of them, plus 8 varieties and forms, are
based on material from the West Indies (Tab. 1). Many of these names are old,
e.g. introduced by E. Tuckerman or W. Nylander based on C. Wright’s collections
from Cuba, which makes it particularly important to understand the taxonomy of
Phyllopsora in this area for the nomenclatural stability in the genus.
In IMSHAUG’s (1957) catalogue of the West Indian lichens, 26 species, varieties
and forms which in this study are regarded as belonging in Phyllopsora were
listed under the genera Lecidea Ach., Phyllopsora, and Psorella Müll. Arg. (Tab.
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2). IMSHAUG (1957) listed one additional Phyllopsora species, P. cryptocarpa
Riddle, which was transferred to Fellhanera Vĕzda by BRAKO (1989) and is not
treated here, and one additional Psorella species, P. triptophyllina (Nyl.) Zahlbr.,
which is here regarded as belonging in Eschatogonia Trevis. (cf. TIMDAL 2008a).
Table 1. Phyllopsora taxa described from the West Indies.
Basionym Country/Island Current name
Biatora pyrrhomelaena Tuck. Cuba P. pyrrhomelaena
Lecidea breviuscula Nyl. Cuba P. breviuscula
L. breviuscula var. phaeobyssina Nyl. Guadeloupe P. phaeobyssina
L. canoumbrina Vain. Trinidad P. canoumbrina
L. cognata Nyl. Cuba P. cognata
L. corallina var. schizophylloides Vain. St. Vincent P. minor
L. furfuracea var. biatorina Vain. Guadeloupe unknown
L. glabella Nyl. Cuba P. microsperma
L. glabriuscula Nyl. Cuba P. orchroxantha
L. granulifera Fink Puerto Rico P. canoumbrina
L. intermediella Nyl. Cuba P. intermediella
L. leucophyllina Nyl. Cuba P. leucophyllina
L. longiuscula Nyl. Cuba P. longiuscula
L. microphyllina Nyl. Cuba P. microphyllina
L. microphyllina var. subgranulosa Tuck. Cuba P. pertexta
L. parvifolia f. corallina Tuck. Cuba P. furfuracea chemotype 2
L. parvifolia f. subgranulosa Tuck. Cuba P. canoumbrina
L. parvifoliella Nyl. Cuba P. parvifoliella
L. pertexta Nyl. Cuba P. pertexta
L. subbreviuscula Nyl. Cuba P. parvifolia
Phyllopsora cryptocarpa Riddle Cuba Fellhanera cryptocarpa
P. parvifolia var. glauca de Lesd. Cuba P. porphyromelaena
P. parvifolia var. hirtella de Lesd. Cuba P. parvifolia
In BRAKO’s (1989, 1991) revision of the neotropical species of Phyllopsora, 18
species and 8 varieties were accepted, of which 13 species and 6 varieties were
recorded from the West Indies (Tab. 2). My own study of Phyllopsora in Peru
(TIMDAL 2008b), however, revealed 20 species in that country alone. Although
my concept of the genus is broader than that of Brako’s, e.g., including most of
the species placed in Psorella in ZAHLBRUCKNER (1926–1927), it soon became
apparent that an examination of Brako's material was needed in order to reevaluate
the circumscription of the West Indian taxa. Hence additional material
was borrowed from several herbaria and I have now studied most of the
specimens of Phyllopsora reported from the West Indies. Several recent
collections not examined by Brako were also included in this study.
320
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Material and methods
Imshaug collected Phyllopsora in Jamaica in 1952 (11 collections, including
immixtures), 1953 (18), and 1958 (3), Grenada in 1953 (6), the Dominican
Republic in 1958 (7), Haiti in 1958 (33), Cuba in 1959 (15), Dominica in 1963
(4), Guadeloupe in 1963 (5), Saint Lucia in 1963 (9), Saint Vincent in 1963 (2),
and Trinidad and Tobago in 1963 (46). The collections from the Dominican
Republic and Haiti were made together with C. M. Wetmore and those in 1963 with
Table 2. Comparison of the taxonomy and nomenclature of IMSHAUG (1957) with that of
BRAKO (1991) and the present study.
Imshaug (1957) Brako (1991) Present study
Lecidea canoumbrina Phyllopsora canoumbrina Phyllopsora canoumbrina
L. furfuracea P. furfuracea P. furfuracea
L. furfuracea var. biatorina unknown unknown
L. glabella P. glabella P. microsperma
L. granulifera P. canoumbrina P. canoumbrina
L. pyrrhomelaena excluded P. pyrrhomelaena
Phyllopsora breviuscula P. parvifolia var. breviuscula P. breviuscula
P. breviuscula var. phaeobyssina P. corallina var. phaeobyssina P. phaeobyssina
P. corallina P. corallina var. corallina P. corallina
P. corallina var. schizophylloides P. minor P. minor
P. intermediella P. intermediella P. intermediella
P. longiuscula P. longiuscula P. longiuscula
P. parvifolia P. parvifolia var. parvifolia P. parvifolia
P. parvifolia f. corallina P. corallina var. corallina P. furfuracea chemotype 2
P. parvifolia var. glauca P. buettneri var. glauca P. porphyromelaena
P. parvifolia var. hirtella not identified P. parvifolia
P. parvifolia var. subgranulosa P. canoumbrina P. canoumbrina
P. parvifoliella P. parvifoliella P. parvifoliella
P. subbreviuscula P. parvifolia var. parvifolia P. parvifolia
Psorella cognata not treated P. cognata
Ps. glabriuscula P. corallina var. ochroxantha P. ochroxantha
Ps. janeirensis excluded P. cinchonarum
Ps. leucophyllina not treated P. leucophyllina
Ps. microphyllina not treated P. microphyllina
Ps. microphyllina var. subgranulosa not treated P. pertexta
Ps. pertexta not treated P. pertexta
P. buettneri var. munda P. buettneri chemotype 2
P. chlorophaea P. chlorophaea
P. confusa P. confusa
P. corallina var. glaucella P. glaucella
P. corallina var. santensis P. santensis
P. kalbii P. kalbii
321
eschweizerbart_xxx

F.A. Imshaug. See FRYDAY & PRATHER (2001) for a general overview on
Imshaug's lichen collections.
Additional material from the West Indies were borrowed from or examined
during visits to: B (17 collections), BG (16), BM (23), FH (3), H (17), NY (171),
TUR (5), UPS (15), and the private herbarium of Sergio Pérez-Ortega (5). My
own collections from a trip to Tobago in 2008 are stored in O (90). Type material
from outside the West Indies were borrowed from or examined in BM, G, H, O,
and S.
Microscopical examinations were carried out on microtome sections cut at 16
μm or on squash preparations. The preparations were observed in distilled water,
lactophenol cotton blue, 10 % KOH (K), and a modified Lugol’s solution (water
replaced by 50 % lactic acid). Polarized light was used for locating crystals in the
microtome sections. The terminology of cortex types follows SWINSCOW & KROG
(1981). Spore measurements are given as X ± 1.5 × SD rounded to 0.5 μm, where
X is the arithmetic mean and SD the standard deviation. With the exception of the
material in UPS and some specimens where otherwise is stated, all cited
specimens were subjected to thin-layer chromatography (TLC) using the standard
methods of CULBERSON & KRISTINSSON (1970) and CULBERSON (1972), modified
by MENLOVE (1974) and CULBERSON & JOHNSON (1982). Routine examinations
were made in solvent systems B and C; for critical studies all three systems were
used.
Results
The secondary chemistry of the West Indian species and chemotypes is given in
Table 3.
The West Indian species
1. Phyllopsora breviuscula (Nyl.) Müll. Arg.
Bull. Herb. Boissier 2, App. 1: 45 (1894). – Lecidea breviuscula Nyl., Annls Sci. nat., Bot., sér. 4,
19: 339 (1863). – Type: CUBA (see SWINSCOW & KROG 1981). – Descriptions: TIMDAL & KROG
(2001), ELIX (2009).
Chemistry: no lichen substances (by TLC).
Notes: The species is recognized by the ellipsoid to fusiform ascospores (9.5–13
× 3.5–5 μm), well developed, reddish brown prothallus and large, elongate,
convex, adnate squamules lacking vegetative dispersal units and lichen
substances. BRAKO (1991) treated it as P. parvifolia var. breviuscula (Nyl.) Brako,
but I regard the taxon as a distinct species. Phyllopsora parvifolia forms smaller,
more plane squamules with phyllidia on the lobe margins in the inner part of the
thallus. It may also be confused with P. microphyllina, but that species forms
smaller squamules and has acicular ascospores, 27–35 ca. 2.5 μm. See also P.
byssiseda and P. microsperma for discussions.
322
eschweizerbart_xxx

Table 3. Secondary compounds occurring in the West Indian species and chemotypes of Phyllopsora. Atr: atranorin, arg: argopsin, narg:
norargopsin, pan: pannarin, vic: vicanicin, nvic: norvicanicin, phy: phyllopsorin, cphy: chlorophyllopsorin, mps: methyl 2,7dichloropsoromate,
mnps: methyl 2,7-dichloronorpsoromate, parv: parvifoliellin, fur: furfuraceic acid.
fur
·
·
parv
·
·
mnps
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
M
·
·
·
·
·
·
·
·
·
·
·
·
·
·
mps
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
cphy
·
·
·
·
·
·
·
·
·
·
·
·
·
·
m
·
·
·
phy
·
M
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
nvic
vic
·
·
t/m
·
·
·
·
·
·
·
·
·
·
M
·
·
·
·
·
·
M
·
·
·
·
·
·
·
·
·
·
M
·
·
·
·
pan
·
M
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
narg
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
arg
·
·
·
·
·
·
t/M
·
·
·
·
·
·
·
M
·
·
·
atr
·
·
·
·/m
·
·/m
·
·/M
M
·
·
·
·
·
·
·
·
·
None
x
·
·
x
x
·
·
·
·
x
x
·
x
·
·
·
x
x
Species/Chemotype
Phyllopsora breviuscula
Phyllopsora buettneri 2
Phyllopsora buettneri 3
Phyllopsora byssiseda
Phyllopsora canoumbrina
Phyllopsora chlorophaea 1
Phyllopsora chodatinica
Phyllopsora cinchonarum
323
eschweizerbart_xxx
Phyllopsora cognata
Phyllopsora confusa
Phyllopsora corallina
Phyllopsora furfuracea 1
Phyllopsora furfuracea 2
Phyllopsora glaucella
Phyllopsora hispaniolae
Phyllopsora imshaugii
Phyllopsora intermediella
Phyllopsora kalbii

fur
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
parv
·
·
·
·
·
·
·
·
·
M
·
·
·
·
·
·
·
·
mnps
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
M
·
·
mps
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
m/M
·
·
cphy
·
·
·
·
·
·
·
S/M
·
·
·
·
·
·
·
·
·/m
·
phy
·
·
·
·
·
·
·
M
·
·
·
·
·
·
·
·
·
·
nvic
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
vic
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
pan
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
narg
·
·
·
·
·
·
·
·/t
·
·
·
·/m
m/M
·
m/S
·
m
·
arg
·
·
·
·
·
·
·
·/t
·
·
·
M
M
·
M
·
M
·
atr
·
·
·
·
·
·
·
·
·
·/m
·
·
·
·
·
·
·
·
None
·
x
·
x
x
x
x
·
x
·
x
·
·
·
·
·
·
·
Table 3. Secondary compounds (continued)....
Species/Chemotype
Phyllopsora labriformis
Phyllopsora lacerata
Phyllopsora leucophyllina
Phyllopsora longiuscula
Phyllopsora microphyllina
Phyllopsora microsperma
Phyllopsora minor
Phyllopsora ochroxantha
Phyllopsora parvifolia
Phyllopsora parvifoliella
Phyllopsora pertexta
Phyllopsora phaeobyssina
Phyllopsora porphyromelaena 1
Phyllopsora pyrrhomelaena
Phyllopsora santensis
Phyllopsora swinscowii
Phyllopsora teretiuscula
Phyllopsora tobagensis
324
eschweizerbart_xxx

Table 3. Secondary compounds occurring in the West Indian species and chemotypes of Phyllopsora. Fpc: fumarprotocetraric acid, lob:
lobaric acid, sek: sekikaic acid, hsek: homosekikaic acid, per: perlatolic acid, hper: hyperlatolic acid, sper: superlatolic acid, nor: norstictic
acid, mbar: methyl barbarate, zeo: zeorin, xan: xanthones, nsol: norsolorinic acid, unk: unknown compound. M: major, m: minor, S:
submajor, t: trace.
unk
·
·
·
·
·
·
·
M
·
·
·
·
·
·
·
·
·
·
nsol
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
xan
·
·
·
·
·
·
M
·
·
·
·
·
·
·
·
·
·
·
zeo
·
M
M
·
·
·
t/M
·
·
·
·
·
·
·
·
·
·
·
mbar
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
nor
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
M
·
·
sper
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
hper
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
per
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
hsek
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
sek
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
lob
·
·
·
·
·
·
·
M
·
·
·
·
·
·
·
·
·
·
fpc
·
·
·
·
·
·
·
·/M
·
·
·
·
·
·
·
·
·
·
Species/Chemotype
Phyllopsora breviuscula
Phyllopsora buettneri 2
Phyllopsora buettneri 3
Phyllopsora byssiseda
Phyllopsora canoumbrina
Phyllopsora chlorophaea 1
Phyllopsora chodatinica
Phyllopsora cinchonarum
325
eschweizerbart_xxx
Phyllopsora cognata
Phyllopsora confusa
Phyllopsora corallina
Phyllopsora furfuracea 1
Phyllopsora furfuracea 2
Phyllopsora glaucella
Phyllopsora hispaniolae
Phyllopsora imshaugii
Phyllopsora intermediella
Phyllopsora kalbii

unk
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
nsol
·
·
·
·
·
·
·
·
·
·
·
·
·
M
·
·
·
·
xan
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
zeo
·
·
·
·
·
·
·
·
·
·
·
·
m/M
·
·
·
·
·
mbar
M
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
nor
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
sper
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
m
hper
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
M
per
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
m
hsek
·
·
M
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
sek
·
·
S
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
lob
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
fpc
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
·
Table 3. Secondary compounds (continued)....
Species/Chemotype
Phyllopsora labriformis
Phyllopsora lacerata
Phyllopsora leucophyllina
Phyllopsora longiuscula
Phyllopsora microphyllina
Phyllopsora microsperma
Phyllopsora minor
Phyllopsora ochroxantha
Phyllopsora parvifolia
Phyllopsora parvifoliella
Phyllopsora pertexta
Phyllopsora phaeobyssina
Phyllopsora porphyromelaena 1
Phyllopsora pyrrhomelaena
Phyllopsora santensis
Phyllopsora swinscowii
Phyllopsora teretiuscula
Phyllopsora tobagensis
326
eschweizerbart_xxx

Phyllopsora breviuscula is pantropical, but in the West Indies only known
from Wright’s material from Cuba and Duss’ material from Guadeloupe (not
examined by me). The specimen from Saint Vincent reported by VAINIO (1896)
belongs in P. chodatinica (Elliot 135, BM).
Specimens examined: CUBA. Wright s.n. (H-NYL 20557, holotype, TLC not performed), Wright,
Lich. Cub. 181 (B, BM, UPS).
2. Phyllopsora buettneri (Müll. Arg.) Zahlbr.
Catal. Lich. Univers. 4 (25): 396 (1926). – Psora buettneri Müll. Arg., Engler Bot. Jahrb. 15: 506
(1893). – Type: TOGO (see SWINSCOW & KROG 1981). – Descriptions: TIMDAL & KROG (2001),
TIMDAL (2008b), ELIX (2009).
Chemistry: chemotype 1 (paleotropical): pannarin (major) and zeorin (major);
chemotype 2 (neotropical): pannarin (major), phyllopsorin (major) and zeorin
(major); chemotype 3 (neotropical): vicanicin (major), norvicanicin (trace to
minor) and zeorin (major); chemotype 4 (paleotropical): dechloropannarin (major)
and zeorin (major); chemotype 5 (Norfolk Island, ELIX 2006a, not examined by
me): argopsin, norargopsin and zeorin.
Notes: The species is recognized by the large, elongated, pruinose, lacinulate
squamules always containing zeorin and in addition pannarin (plus phyllopsorin in
the Neotropics) or the related compounds argopsin (plus norargopsin),
dechloropannarin or vicanicin. Both neotropical chemotypes occur in the West
Indies. If regarded as distinct species, they should be named P. munda (Malme)
Zahlbr. (chemotype 2) and P. melanoglauca Zahlbr. (chemotype 3). There are no
known morphological or anatomical differences between the five chemotypes of
P. buettneri, however. The holotype of P. munda (Brazil, Malme, Lich. Regnell
617b, S) contains phyllopsorin (major), pannarin (submajor), and zeorin (minor),
and an isotype of P. melanoglauca (Brazil, Schiffner s.n., BM) contains vicanicin
and zeorin (both major).
BRAKO's (1991) varieties and chemical strains of P. buettneri correspond to the
following species and chemotypes in the present work: var. buettneri = P.
buettneri chemotype 1; var. glauca (B. de Lesd.) Brako chemical strain I = P.
porphyromelaena; strain II = P. chodatinica; strain III = P. porphyromelaena; var.
munda (Malme) Brako = P. buettneri chemotype 2. See also P. hispaniolae for
discussion.
The species is pantropical. New to Haiti, Jamaica and Trinidad and Tobago.
Specimens examined: – Chemotype 2: CUBA. Harris 14444, 14559, 14592 (NY). DOMINICAN
REPUBLIC. Harris 20493, 20495 (NY). – Chemotype 3: CUBA. Harris 14555 (NY), Imshaug
24839, 24865 (MSC), Pérez-Ortega s.n. (hb Pérez-Ortega), Wright, Lich. Cub. 180 (BM, UPS).
HAITI. Imshaug & Wetmore 2883, 3145, 3157- immixture, 22701 (MSC). DOMINICAN REPUBLIC.
Buck 8348 (NY), Harris 15563 (NY), Imshaug & Wetmore 23662, 23669 (MSC). JAMAICA.
Imshaug 13166, 14617 (MSC). TRINIDAD & TOBAGO. Imshaug & Imshaug 31665, 32146 (MSC).
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3. Phyllopsora byssiseda (Nyl.) Zahlbr.
Catal. Lich. Univers. 4 (25): 396 (1926). – Lecidea byssiseda Nyl. in Hue, Nouv. Arch. Mus. Hist.
Nat., sér. 3, 3: 103 (1891). – Type: MEXICO, s. loc., Fr. Müller (H-NYL 20517, holotype) [no
lichen substances (by TLC)].
(Colour Plate 14A)
Description: Prothallus well developed, white. Thallus effuse or incompletely
circular, squamulose, with radiating marginal squamules. Squamules medium
sized, up to 0.8 mm diam., adnate, elongated, deeply divided or lobed, weakly to
moderately convex; upper side pale brown (old herbarium material), epruinose,
glabrous; margin concolorous with upper side, pubescent. Isidia numerous,
developing from the tips of the squamules or often directly from the prothallus,
cylindrical, long, sometimes branched. Upper cortex of type 1, 40–50 μm thick,
containing scattered crystals, these dissolving in K. Medulla not containing
crystals, PD–, K–. Apothecia not common, up to 1.5 mm diam., weakly convex,
medium brown, rounded, simple or more rarely conglomerate, with a paler, thin,
more or less persistent margin which is pubescent on the lower side. Excipulum
pale brown to colourless, K–. Hypothecium dark yellowish brown, K–.
Epithecium pale brown to colourless, K–. Crystals present only in the hymenium,
dissolving in K. Ascospores narrowly ellipsoid to fusiform, simple, 9–13 3–4
μm (20 spores from one apothecium).
Chemistry: no lichen substances (by TLC) or atranorin (trace to minor).
Notes: The type material was studied by BRAKO (1991) but left uninterpreted due
to its poor condition. The West Indian material now available shows a
characteristic species similar to P. breviuscula but differing in having a thick,
white prothallus, long, thick isidia, and in the presence of small amounts of
atranorin in the thallus. In P. breviuscula, the prothallus is reddish brown,
vegetative dispersal units are lacking, and no lichen substances occur. I failed to
observe atranorin in the holotype of P. byssiseda (TLC), but a possible trace of
this compound was reported from the same specimen by SWINSCOW & KROG
(1981). In Imshaug & Wetmore 3144 and 22726 (no atranorin by TLC), crystals
dissolving in K were seen in the upper cortex (polarized light), indicating that
atranorin occurs in these specimens in a concentration too low for detection by
standard TLC.
Phyllopsora byssiseda was previously known only from the type locality in
Mexico (near Orizaba, according to NYLANDER 1858).
Specimens examined: HAITI. Dept. de L'Ouest, road from Foret des Pins to Savane-Zombie, 5300
ft, 1958, Imshaug & Wetmore 3060-immixture in P. ochroxantha (MSC); summit of small peak at
N end of ridge above (E of) Foret des Pins, 5800–5900 ft, 1958, Imshaug & Wetmore 3144, 3157
(MSC); ridge N of Foret des Pins (SHADA station), near border of Dominican Republic, 5800 ft,
1958, Imshaug & Wetmore 22726 (MSC). DOMINICAN REPUBLIC. Independencia, Sierra de
Baoruco, 23.5 km S of Puerto Escondido, then 9 km E along road to Charco Colorado, 1816'N,
7131'W, 1800 m, 1987, Harris 20547 (NY). JAMAICA. Parish of Portland or St. Andrew, Dicks
Pond Trail near Hardwar Gap, Blue Mts, 3800 ft, 1952, Imshaug 13112 (MSC); Mount Horeb
ridge, Blue Mts, 4400–4600 ft, 1952, Imshaug 13171 (MSC); Parish of St. Thomas, S slope of
Mossmans Peak, Blue Mts., 5000 ft, 1953, Imshaug, 14671-immixture in Krogia antillarum
(MSC).
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4. Phyllopsora canoumbrina (Vain.) Brako
Mycotaxon 35: 12 (1989). – Lecidea canoumbrina Vain, Proc. Am. Acad. Arts Sci. 58: 135
(1923). – Type: TRINIDAD & TOBAGO (see BRAKO 1991). – Description: BRAKO (1991).
Chemistry: no lichen substances (by TLC).
Notes: This poorly understood species is characterized by the crustose thallus on a
thin, white or partly reddish brown prothallus, the cylindrical isidia, the medium
brown apothecia, the dark yellowish brown hypothecium, and the ellipsoid
ascospores (5–7.5 2.5–3 μm, n = 20, my measurements in TUR-V 23680). It
may be conspecific with the acid deficient chemotype of P. furfuracea, but differs
apparently in having a yellowish brown hypothecium and shorter ascospores. In
P. furfuracea the hypothecium is reddish brown and the ascospores 7–13 2–3
μm (TIMDAL 2008b).
In addition to the West Indian material, the species is known from a few
collections from Guatemala, Venezuela, and Brazil (BRAKO 1991). The report
from Jamaica [RIDDLE 1912, as Bacidia subgranulosa (Tuck.) Riddle], was based
on a specimen of P. pertexta (Cummings 118, NY).
Specimens examined: CUBA. Wright, Lich. Cub. 185 (BM, UPS, isolectotypes of Lecidea
parvifolia f. subgranulosa Tuck.). PUERTO RICO. Britton & Cowell 4235 (NY, isotype of Lecidea
granulifera Fink, TLC not performed). TRINIDAD & TOBAGO. Thaxter 19 (TUR-V 23680, isotype
of L. canoumbrina).
5. Phyllopsora chlorophaea (Müll. Arg.) Zahlbr.
Denkschr. Kaiserl. Akad. Wissensch. 83: 133 (1909). – Psora chlorophaea Müll. Arg., Flora 70:
320 (1887). – Type: BRAZIL (see SWINSCOW & KROG 1981). – Descriptions: TIMDAL & KROG
(2001), TIMDAL (2008b).
Chemistry: chemotype 1 (pantropical): no lichen substances or atranorin (trace to
minor); chemotype 2 (paleotropical): furfuraceic acid and sometimes atranorin
(trace).
Notes: The species is recognized by the ascending, lacinulate squamules attached
to a well developed, reddish brown prothallus, the dark brown apothecia, the dark
reddish brown (K+ purple) hypothecium, and the narrowly ellipsoid to fusiform
ascospores, 7–12.5 × 2.5–3 μm. All examined specimens from the West Indies
belong in chemotype 1, as all other examined specimens from the Neotropics (cf.
BRAKO 1991, TIMDAL 2008b).
When sterile, it may be confused with P. longiuscula, but differs in forming a
thallus consisting mainly of ascending squamules; areolae are few and mostly
elongated. In P. longiuscula the thallus is dominated by isodiametrical areolae and
ascending squamules are scattered. Admittedly, sterile, poorly developed material
may be difficult to identify. When fertile, the two species differ in the colour of
the apothecium, the colour of the hypothecium, and in the spore size. See also P.
confusa for discussion.
Phyllopsora chlorophaea is pantropical. New to Trinidad and Tobago.
Specimens examined: CUBA. Britton et al. 15491 (B), Buck 7693 (NY), Imshaug 24749 (MSC),
Johnson s.n. (NY), Pérez-Ortega s.n. (hb Pérez-Ortega). HAITI. Buck 9083 (NY), Imshaug &
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Wetmore 2943, 3011, 3183, 3187, 22719, 23068 (MSC). DOMINICAN REPUBLIC. Buck 4361, 4376,
8362, 14499, 14503 (NY), Harris 15367, 15367, 20473, 20480, 20487, 20489, 20670, 20671
(NY), Imshaug & Wetmore 3788, 23680, 23700 (MSC). JAMAICA. Hart 34 (NY), Imshaug 13163,
13301, 13758, 15610 (MSC). TRINIDAD & TOBAGO. Imshaug & Imshaug 32307-immixture
(MSC).
6. Phyllopsora chodatinica Elix
Australasian Lichenology 59: 23 (2006). – Type: AUSTRALIA (see ELIX 2006b). – Descriptions:
ELIX (2006b, 2009), TIMDAL (2008b).
Chemistry: a series of xanthones (major), argopsin (trace to major) and zeorin
(trace to major).
Notes: The species resembles P. porphyromelaena and differs mainly in the
secondary chemistry, i.e. in the presence of a series of xanthones. No attempt was
made to identify the xanthones in this study. The main xanthone(s) corresponds to
BRAKO's (1991) unknown B, and the species P. chodatinica corresponds to
Brako’s P. buettneri var. glauca chemical strain II.
Phyllopsora chodatinica is known from Australia and New Guinea, and is
widely distributed in the Neotropics. New to Saint Lucia.
Specimens examined: CUBA. Harris 14279, 14290, 14379 (NY), Tønsberg 37724, 37923 (BG).
DOMINICA. Imshaug & Imshaug 33186 (MSC). DOMINICAN REPUBLIC. Harris 15005, 26683
(NY). PUERTO RICO. Buck 4195, 4090A (NY), Harris 22248, 22447 (NY), Heller 4764 (NY),
Landrón 394, 491, 502 (MSC). JAMAICA. Cummings 44 (NY), Imshaug 13101 (MSC). SAINT
LUCIA. Imshaug & Imshaug 29810 (MSC). SAINT VINCENT AND THE GRENADINES. Elliott 135
(BM). TRINIDAD & TOBAGO. Britton & Hazen 390 (NY), Fleming s.n. (B, NY), Imshaug &
Imshaug 31472, 31664, 31689, 31710, 31892, 31982, 32338 (MSC), Rui & Timdal: 39 collections
(O).
7. Phyllopsora cinchonarum (Fée) Timdal
The Lichenologist 40: 346 (2008). – Triclinum cinchonarum Fée, Essai crypt. écorc.: 147 (1825)
– Type: PERU (see TIMDAL 2008b). – Descriptions: TIMDAL (2008b), ELIX (2009, as Triclinum
cinchonarum).
Chemistry: West Indian material: lobaric acid (major), unknown compound
(major), atranorin (absent to major) and fumarprotocetraric acid (absent to major).
See TIMDAL (2008b) for a summary of the chemical variation elsewhere.
Notes: The species was treated as P. janeirensis (Müll. Arg.) Swinscow & Krog
by SWINSCOW & KROG (1981). BRAKO (1989) included it in the monotypic genus
Squamacidia Brako, as Squamacidia janeirensis (Müll. Arg.) Brako. TIMDAL
(2008b) reduced Squamacidia to synonymy with Phyllopsora. The species may be
confused mainly with P. imshaugii, see that species for discussion.
Phyllopsora cinchonarum is pantropical. New to Trinidad and Tobago.
Specimens examined: CUBA. Wright, Lich. Cub. 179b-immixture (BM). JAMAICA. Cummings 48
(NY), Hart 320 (NY), Imshaug 13151 (MSC). TRINIDAD & TOBAGO. Fleming s.n. (NY), Imshaug
& Imshaug 31726-immixture (MSC).
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8. Phyllopsora cognata (Nyl.) Timdal comb. nova
Lecidea cognata Nyl., Annls Sci. nat., Bot., sér 4, 19: 347 (1863). – Psorella cognata (Nyl.)
Zahlbr., Catal. Lich. Univers. 4 (26): 402 (1926). – Type: CUBA, s. loc., Wright, Lich. Cub. 218
(BM, lectotype selected here; BM, UPS, isolectotypes) [atranorin (major)].
(Colour Plate 14B)
Description: Prothallus thin, white. Thallus effuse, crustose, not radiating at the
margin. Areolae small, up to 0.1 mm wide, discrete when young, soon adjoining
to form a more or less continuous crust, pale green (old herbarium material), dull,
epruinose, glabrous. Isidia and soredia lacking (but see below). Upper cortex
poorly defined, 5–10 μm thick, containing crystals dissolving in K. Medulla
containing crystals dissolving in K, K+ faintly yellow. Apothecia common, up to
1.5 mm diam., plane when young, becoming weakly to moderately convex,
yellowish brown, rounded to irregular, simple to conglomerate, with a thin, more
or less persistent, paler, glabrous margin. Excipulum pale brown to colourless,
containing crystals dissolving in K, K–. Hypothecium pale brown, not containing
crystals, K–. Epithecium colourless, K–. Ascospores acicular, simple, with up to 3
pseudosepta, 19–30 × ca. 1.5 μm (n = 20).
Chemistry: atranorin (major).
Notes: The species is known only from Wright’s collections in Cuba. No original
specimens were located in H-NYL, but No. 218 in Wright’s exsiccata is assumed
to be a part of the original material and a specimen in BM is here selected as the
lectotype. Wright’s exsiccata No. 217 (BM, UPS) may belong in the same species
but is sorediate; more material is needed to decide if the presence of punctiform
soralia is a part of the variation of P. cognata. See also P. pertexta for discussion.
BRAKO (1989) did not include the species in Phyllopsora because it was said to
possess different tissue types in the exciple and hypothecium; thinner, less
gelatinized paraphyses; and long, septate, filiform ascospores. It fits Phyllopsora
in the broader concept of the genus adopted here, however (see discussion on the
circumscription of Phyllopsora in TIMDAL 2008b).
Specimens examined: CUBA. Wright, Lich. Cub. 218 (BM, lectotype; BM, UPS isolectotypes).
9. Phyllopsora confusa Swinscow & Krog
The Lichenologist 13: 229 (1981) – Type: KENYA (see SWINSCOW & KROG 1981). – Descriptions:
TIMDAL & KROG (2001), ELIX (2009).
(Colour Plate 14C)
Chemistry: no lichen substances (by TLC).
Notes: The species is characterized by the thin, white or sometimes partly reddish
brown prothallus, the small to medium sized, narrow (up to 0.3 mm wide),
ascending, more or less imbricate squamules which easily break off diaspores
terminally (lacinules), the lack of lichen substances, the pale apothecia with pale
exciple and hypothecium, and the narrowly ellipsoid to shortly bacilliform
ascospores. Phyllopsora chlorophaea differs mainly in having a thicker, always
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eddish brown prothallus and darker brown apothecia with a reddish brown inner
part of the exciple and hypothecium.
Phyllopsora confusa is pantropical. New to the Dominican Republic, Haiti and
Jamaica.
Specimens examined: CUBA. Britton et al. 15491 (NY), Buck 23422 (NY), Harris 14400, 14424
(NY), Johnson s.n. (NY), Pérez-Ortega s.n. (hb Pérez-Ortega), Tønsberg 37813, 37816, 37818
(BG). DOMINICAN REPUBLIC. Buck 14196 (NY), Harris 15750, 19737, 15679a, 26476 (NY).
HAITI. Imshaug & Wetmore 3147 (MSC). JAMAICA. s. loc., II or III. 1905, Cummings 44 p.p. (NY,
immixture with P. chodatinica in two envelopes). SAINT VINCENT & THE GRENADINES. Elliot 264
(TUR-V 22605), Imshaug & Imshaug 30336, 30637 (MSC). TRINIDAD & TOBAGO. Britton &
Britton 684 (NY), Imshaug & Imshaug 31159, 31160 (MSC), Rui & Timdal 10736, 10788, 10789,
10790, 10797, 10819, 10826, 10863 (O).
10. Phyllopsora corallina (Eschw.) Müll. Arg.
Engler Bot. Jahrb. 20: 264 (1894). – Lecidea corallina Eschw. in Martius, Fl. Bras. 1: 256 (1833).
– Type: BRAZIL (see SWINSCOW & KROG 1981). – Descriptions: TIMDAL & KROG (2001), ELIX
(2009).
Chemistry: no lichen substances (by TLC).
Notes: The current circumscription of P. corallina corresponds to P. corallina
var. corallina in BRAKO (1991). Brako’s other varieties are here regarded as
distinct species: P. glaucella, P. ochroxantha, P. phaeobyssina, P. rappiana
(Brako) Elix (not known from the West Indies), and P. santensis; see each of them
(except P. rappiana), and also P. intermediella and P. kalbii, for discussions.
Phyllopsora corallina is mainly characterised by the medium sized squamules
forming marginal, long, cylindrical isidia and the lack of lichen substances.
BRAKO’s (1991) four specimens of P. corallina var. corallina from the West
Indies were found to belong in P. furfuracea chemotype 2 (SIPMAN 14953,
Underwood & Earle 1004, Wright Lich. Cub. 184) and P. confusa (Harris 14400).
However, I have examined a specimen of P. corallina from the Dominican
Republic. One of the redetermined specimens, Wright’s Lich. Cub. 184 (BM,
UPS), is an isolectotype for Lecidea parvifolia var. corallina Tuck.; this name is
hence a synonym of P. furfuracea, not of P. corallina as stated by BRAKO (1991).
The species is pantropical. New to the Dominican Republic.
Specimens examined: DOMINICAN REPUBLIC. Harris 26562 (NY).
11. Phyllopsora furfuracea (Pers.) Zahlbr.
Flechten. Naturl. Pflanzenfam. T. 1, Abt. 1 * , Lief. 225: 138 (1906). – Lecidea furfuracea Pers. in
Gaudichaud, Voy. Uranie (5): 192 (1827). – Type: MARIANA ISLANDS (see BRAKO 1991). –
Lecidea parvifolia var. corallina Tuck., Proc. Am. Acad. Arts Sci.6: 273 (1864). – Descriptions:
TIMDAL & KROG (2001), TIMDAL (2008b), ELIX (2009).
Chemistry: chemotype 1: furfuraceic acid (major); chemotype 2: no lichen
substances (by TLC).
Notes: The species is mainly characterised by the crustose thallus and the long,
cylindrical, often dominating isidia, the reddish brown hypothecium, the narrowly
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ellipsoid ascospores (7–13 2–3 μm, TIMDAL 2008b), and the presence of
furfuraceic acid (chemotype 1) or no lichen substances (chemotype 2). There are
several morphologically similar species, see P. canoumbrina, P. teretiuscula, and
P. tobagensis for discussions. Lecidea parvifolia var. corallina Tuck. is here
synonymised with P. furfuracea chemotype 2 (see P. corallina).
Phyllopsora furfuracea is pantropical. New to Haiti, Netherlands Antilles (St.
Eustatius), and Trinidad and Tobago. The specimen from Guadeloupe (VAINIO
1915, type material of Lecidea furfuracea var. biatorina Vain.) was not examined
by me (nor BRAKO 1991) as the material was not found in TUR.
Specimens examined: – Chemotype 1: CUBA. Buck 23401 (NY), Harris 14453 (NY), 14547 (B,
NY), 14548, 14657 (NY), Imshaug 24632, 25074 (MSC). HAITI. Imshaug & Wetmore 3199,
22719-immixture (MSC). DOMINICAN REPUBLIC. Buck 14493, 14688 (NY), Harris 15009, 15345,
15679, 15754, 19751, 20079, 20109, 20471, 20481, 20677, 26725 (NY), Imshaug & Wetmore
3786 (MSC). JAMAICA. Buck 5646, Cummings 49, s.n. (NY), Hart 59 (NY), Imshaug 15799,
15803 (MSC). TRINIDAD & TOBAGO. Imshaug & Imshaug 31256 (MSC), Rui & Timdal 10787,
10794, 10795, 10799, 10809, 10852, 10859, 10866, 10869 (O). – Chemotype 2: CUBA. Imshaug
24817 (MSC), Underwood & Earle 1004 (NY), Wright, Lich. Cub. 184 (BM, UPS, isolectotypes
of Lecidea parvifolia var. corallina Tuck). NETHERLANDS ANTILLES. Sipman 14953, 56733 (B).
TRINIDAD & TOBAGO. Fleming s.n. (NY), Imshaug & Imshaug 31169, 31327, 31765 (MSC), Rui
& Timdal 10732, 10792 (O).
12. Phyllopsora glaucella (Vain.) Timdal
The Lichenologist 40: 349 (2008). – Lecidea breviuscula var. glaucella Vain., Dansk Bot. Arkiv 4,
11: 21 (1926); – Phyllopsora corallina var. glaucella (Vain.) Brako, Mycotaxon 35: 13 (1989). –
Type: MEXICO (see TIMDAL 2008b). – Description: TIMDAL (2008b).
(Colour Plate 14D)
Chemistry: vicanicin (major) and norvicanicin (major).
Notes: The species was treated as P. corallina var. glaucella in BRAKO (1991). In
addition to the chemical difference (vicanicin and norvicanicin vs. no lichen
substances), it differs from P. corallina in having generally larger, more adnate
squamules attached to a thicker, always reddish brown prothallus.
The species is neotropical. New to Cuba and Haiti.
Specimens examined: CUBA. Imshaug 25061, 25064 (MSC), Johnson s.n. (NY). HAITI. Imshaug
& Wetmore 22929, 22936 (MSC). DOMINICAN REPUBLIC. Buck 14757, 19349 (NY), Harris 20779
(BM, NY), 20794, 26773 (NY). PUERTO RICO. Britton et al. 4398 (NY), Buck 15983 (NY), Harris
22028 (NY).
13. Phyllopsora hispaniolae Timdal sp. nova
Phyllopsorae buettnero similis, sed squamis epruinosis, minoribus, angustioribus,
ascosporis angustioribus, et thallo zeorinum destituto. – Type: DOMINICAN REPUBLIC.
Prov. Independencia, Sierra de Baoruco, Charco de la Paloma, 48.4 km S of Puerto
Escondito, 1800 m, ca. 18°15’N, 71°36’W, humid hardwoods around waterhole, I.
1987, Harris 20672 (NY, holotype) [argopsin (major) and chlor-ophyllopsorin
(minor)]; 23.5 km S of Puerto Escondido at intersection of road to Charco Colorado,
ca. 18°16'N, 71°33'W, 1750 m, Harris 20455; 30.5 km S of Puerto Escondido, ca.
18°14'N, 71°33'W, 1940 m, Buck 14618, 14634 (NY, paratypes).
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(Colour Plate 14E)
Description: Prothallus well developed, reddish brown. Thallus effuse,
squamulose, not radiating at the margin. Squamules medium sized, adnate or
ascending, consisting mainly of deeply divided or coralloid, ca. 0.1 mm wide, up
to 1 mm long, convex to almost terete, more or less geotropically arranged lobes;
upper side pale green, slightly shiny, epruinose, finely tomentose; lobe tips paler
than inner part, pubescent. Vegetative dispersal units lacking but lobe parts
probably acting as lacinules. Upper cortex of type 1–2, 30–40 μm thick, not
containing crystals. Medulla containing crystals not dissolving in K, PD+ orange,
K–. Apothecia common, up to 1 mm diam., weakly to moderately convex,
medium brown, rounded, simple to conglomerate, with an indistinct, finely
pubescent margin when young. Excipulum pale brown in inner part, darker in the
rim, K–. Hypothecium pale brown, K–. Epithecium pale brown to colourless, K–.
Scattered crystals, not dissolving in K, occurring in the inner part of the exciple
and hymenium. Ascospores narrowly ellipsoid, simple, 8–11 2.5–3 μm (n = 50).
Chemistry: argopsin (major) and chlorophyllopsorin (minor).
Notes: The species resembles P. buettneri but the squamules are epruinose and
smaller, more narrow-lobed to almost terete, the ascospores are smaller, and the
thallus lacks zeorin. In P. buettneri the lobes are up to 1 mm wide, the ascospores
11–14 × 3–3.5 μm (TIMDAL 2008b), and the thallus contains other combinations
of compounds, always including zeorin. The paleotropical P. africana Timdal &
Krog and some specimens of P. teretiuscula contain the same combination of
compounds as P. hispaniolae. The former is an isidiate, broad-lobed species with
a glabrous upper side, the latter forms more densely coralloid squamules with
glabrous, or at most finely pubescent, tips.
Phyllopsora hispaniolae is known from three localities in the Sierra de
Baoruco in the Dominican Republic, collected in humid forests from 1740 to 1990
m altitude. The species is corticolous.
14. Phyllopsora imshaugii Timdal sp. nova
Phyllopsorae cinchonaris similis, sed squamis minoribus, cortici tenuiori,
ascosporis brevioribus, et thallo acidum norsticticum continenti. – Type: JAMAICA,
Parish of Portland or St. Thomas, summit of Blue Mountain Peak, 7400 ft, X. 1952,
Imshaug 13037 (MSC-25550, holotype) [norstictic acid (major)]; Parish of St.
Thomas, summit of Mossmans Peak, Blue Mts, 6600 ft, Imshaug 14711 (MSC-
25546, paratype).
(Colour Plate 14F)
Description: Prothallus thin, white. Thallus effuse, crustose to squamulose.
Squamules small, up to 0.2(–0.3) mm wide, adnate, isodiametrical, more or less
scattered when young, later contiguous, more or less crenulate, plane to weakly
convex; upper side medium green, somewhat shiny, epruinose, glabrous; margin
concolorous with upper side, glabrous to pubescent. Isidia common, cylindrical,
long, simple or branched. Upper cortex of type 1–2, 15–30 μm thick, containing a
few scattered crystals dissolving in K. Medulla containing crystals dissolving in K
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and recrystallizing by forming acicular, red crystals, PD+ yellow, K+ red.
Apothecia common, up to 2 mm diam., plane when young, soon becoming weakly
to moderately convex, medium brown to dark brown, rounded to irregular, simple
to conglomerate, when young with a rather thick, paler, often weakly pubescent
margin. Excipulum pale brown, darker in inner part, containing crystals dissolving
in K and recrystallizing by forming acicular, red crystals. Hypothecium dark
yellowish brown in upper part, paler below, not containing crystals, K–.
Epithecium pale brown to colourless, K–. Ascospores narrowly ellipsoid to shortly
bacilliform, simple, 10.5–14.5 3–4 μm (n = 40).
Chemistry: norstictic acid (major).
Notes: The isidiate thallus, white prothallus, distinctly marginate apothecia (at
least when young), and dark brown hypothecium make the species most similar to
P. cinchonarum. It differs, however, in having smaller squamules (often just
areoles), a thinner upper cortex, shorter ascospores, and in containing norstictic
acid. This compound is not previously reported from Phyllopsora.
The species is known from two collections from the summit area of the Blue
Mountains in Jamaica (2010–2260 m). The specimens were corticolous, partly
also growing over corticolous bryophytes.
15. Phyllopsora intermediella (Nyl.) Zahlbr.
Catal. Lich. Univers. 4 (25): 398 (1926). – Lecidea intermediella Nyl., Annls Sci. Nat., Bot., sér.
4, 19: 339 (1863). – Type: CUBA (see BRAKO 1991). – Description: BRAKO (1991).
(Colour Plate 15A)
Chemistry: no lichen substances (by TLC).
Notes: The species is characterized by the pale green, closely adnate or
sometimes slightly ascending, mainly isodiametrical squamules attached to a
thick, reddish brown prothallus, and the long, cylindrical to partly flattened,
sometimes branched isidia. Apothecia are rare and poorly developed in the West
Indian material, but according to BRAKO (1991) they are yellow-brown. The
ascospores are 7–9 2.5–3.5 μm in the type material (BRAKO 1991). It may be
confused mainly with P. corallina, which differs in forming darker (often
brownish) green, more ascending, elongated and imbricate squamules on a
thinner, often white prothallus, and in having exclusively cylindrical isidia.
The species is neotropical (BRAKO 1991). New to Haiti, Netherlands Antilles,
and Puerto Rico. With the exception of the material from Cuba, all West Indian
specimens were saxicolous.
Specimens examined: CUBA. Wright s.n. (H-NYL 20558, holotype, TLC not performed), Wright,
Lich. Cub. 183 (BM, UPS). HAITI. Imshaug & Wetmore 3026, 22511 (MSC). DOMINICAN
REPUBLIC. Buck 8305 (NY). JAMAICA. Buck 5883 (NY). PUERTO RICO. Harris 23942 (NY).
NETHERLANDS ANTILLES. Buck 50750 (B), Sipman 14819 (B).
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16. Phyllopsora kalbii Brako
Fl. Neotropica 55: 51 (1991). – Type: BRAZIL (see BRAKO 1991). – Description: TIMDAL & KROG
(2001).
Chemistry: no lichen substances (by TLC).
Notes: The species is recognized by the pale green, squamulose thallus on a thin,
white prothallus, the globose to shortly cylindrical isidia, the medium brown
apothecia, the dark yellowish brown hypothecium, and the ellipsoid to bacilliform
ascospores (6–10 2–3 μm). It may be confused with P. corallina, which differs
in having long, cylindrical isidia and a pale brown to colourless hypothecium.
The species is known from USA, the West Indies, Brazil, Kenya, and Tanzania
(BRAKO 1991). New to Cuba.
Specimens examined: CUBA. Pérez-Ortega s.n. (hb Pérez-Ortega). DOMINICAN REPUBLIC. Buck
4631, 4634 (NY).
17. Phyllopsora labriformis Timdal
The Lichenologist 40: 350 (2008). – Type: PERU (see TIMDAL 2008b). – Description: TIMDAL
(2008b).
Chemistry: methyl barbarate (major).
Notes: The unknown substance reported from P. glaucescens Timdal and P.
labriformis by TIMDAL (2008b) was identified as methyl barbarate by ELIX (in
litt.) based on paratype material of P. glaucescens. Phyllopsora labriformis was
previously known from a single locality in the Peruvian Amazonas. New to
Trinidad and Tobago.
Specimens examined: TRINIDAD & TOBAGO. Parish of St. Paul, along Roxborough – Parlatuvier
Road, 1117.07'N, 6035.70'W, 400–450 m, 2008, Rui & Timdal 10771 (O); Parish of St. George,
along road between Mason Hall and Hillsborough Dam, 1113.81'N, 6041.07'W, 230 m, 2008,
Rui & Timdal 10786 (O).
18. Phyllopsora lacerata Timdal
The Lichenologist 40: 352 (2008). – Type: PERU (see TIMDAL 2008b). – Description: TIMDAL
(2008b).
Chemistry: no lichen substances (by TLC).
Notes: Previously known from Ecuador and Peru. New to Cuba and Trinidad and
Tobago. The Cuban specimens and one from Tobago (Rui & Timdal 10805) are
rather poorly developed, lacking the vein-like tissue at the base of the squamules.
Specimens examined: CUBA. s.loc., Wright, Lich. Cub. 212 (BM, UPS); s. loc., Wright s.n. (H-
NYL 17345b, d, TLC not performed). TRINIDAD & TOBAGO. Parish of St. Paul, along Roxborough
– Parlatuvier Road, 1116.82'N, 6036.60 W, 500–520 m, 2008, Rui & Timdal 10759 (O);
1117.01'N, 6036.39'W, 500 m, Rui & Timdal 10805 (O); 1116.84'N, 6037.32'W, 500–550 m,
Rui & Timdal 10829 (O); 1116.77'N, 6037.44'W, 500–550 m, Rui & Timdal 10845 (O).
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19. Phyllopsora leucophyllina (Nyl.) Timdal
The Lichenologist 40: 352 (2008). – Lecidea leucophyllina Nyl., Annls Sci. Nat., Bot., sér. 4, 19:
347 (1863). – Type: CUBA (see TIMDAL 2008b). – Description: TIMDAL (2008b).
Chemistry: homosekikaic acid (major) and sekikaic acid (submajor).
Notes: The species is known from a few collections from Cuba, Guyana and Peru
(TIMDAL 2008b).
Specimens examined: CUBA. s. loc., Wright s.n. (H-NYL 17345c & e, holotype); s. loc., Wright,
Lich. Cub. 213 (BM, UPS).
20. Phyllopsora longiuscula (Nyl.) Zahlbr.
Catal. Lich. Univers. 4 (25): 398 (1926). – Lecidea longiuscula Nyl., Annls Sci. Nat., Bot., sér. 4,
19: 339 (1863). – Type: CUBA (see SWINSCOW & KROG 1981). – Description: BRAKO (1991).
(Colour Plate 15B)
Chemistry: no lichen substances (by TLC).
Notes: The species is characterized by the reddish brown prothallus, the more or
less continuous crust of small (0.1–0.3 mm) isodiametrical areolae from which
develop scattered, ascending, elongated squamules, the medium brown apothecia,
the yellowish brown (K–) hypothecium, the large ascospores (12.5–17.5 × 3–4
μm, n = 20, from two collections), and the absence of lichen substances. BRAKO
(1991) regarded the ascending squamules as isidia and discussed the species’
delimitation against the isidiate P. intermediella. I regard the squamules as
lacinules, and find sterile material difficult to distinguish especially from P.
chlorophaea, see that species for discussion.
Phyllopsora longiuscula is known from few collections. Only Wright’s
material from Cuba and Imshaug 32292 from Trinidad are fertile and the basis for
the spore measurements given above. In addition to the West Indian material, the
species is known from Venezuela (BRAKO 1991). New to Trinidad and Tobago.
Specimens examined: CUBA. Wright s.n. (H-NYL 20537, holotype, TLC not performed), Wright
Lich. Cub. 179[a] (BM, UPS), Ser. 2, 121 (H-NYL 20535, TLC not performed), Wright s.n. (BM).
PUERTO RICO. Britton et al. 4509 (NY). TRINIDAD & TOBAGO. Imshaug & Imshaug 31363, 31528,
31606, 32292 (MSC), Rui & Timdal 10730, 10800, 10865 (O).
21. Phyllopsora microphyllina (Nyl.) Swinscow & Krog
The Lichenologist 13: 243 (1981). – Lecidea microphyllina Nyl., Annls Sci. Nat., Bot., sér. 4, 19:
347 (1863). – Type: CUBA, “in ins. Cuba”, C. Wright s.n. (H-NYL 17345a, holotype) [too scanty
for TLC].
(Colour Plate 15C)
Description: Prothallus indistinct. Thallus effuse, squamulose, without radiating
marginal squamules. Squamules medium sized, up to 0.5 mm wide, mostly
adnate, isodiametrical to elongated, plane to weakly convex, crenulate to lobed,
geotropically arranged; upper side brownish green (old herbarium material),
337
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slightly shiny, epruinose, glabrous; margin concolorous with upper side, not
pubescent. Vegetative dispersal units lacking. Upper cortex of type 1–2, 30–50
μm thick, not containing crystals. Medulla not containing crystals, PD–, K–.
Apothecia common, up to 1 mm diam., plane to weakly convex, medium brown,
rounded, mostly simple, with an indistinct, slightly paler, glabrous, more or less
persistent margin. Excipulum colourless, K–. Hypothecium colourless in lower
part, pale brown in upper part, K–. Epithecium colourless, K–. No crystals in the
apothecium. Ascospores acicular, simple, with up to 3 pseudosepta, 27–35 ca.
2.5 μm (n = 27, from a single apothecium).
Chemistry: no lichen substances (by TLC).
Notes: The species is characterized by the squamulose thallus without vegetative
dispersal units, the acicular ascospores, and the lack of lichen substances. See P.
breviuscula for discussion.
Phyllopsora microphyllina is known only from Wright’s material from Cuba
and from two collections from Isle of Pines (RIDDLE 1923, material not examined
by me).
Specimens examined: CUBA. Wright s.n. (H-NYL 17345a, holotype, TLC not performed),
Wright, Lich. Cub. 211 (BM, UPS).
22. Phyllopsora microsperma Müll. Arg.
Bull. Herb. Boissier 2: 89 (1894). – Type: MEXICO (see BRAKO 1991). – Phyllopsora glabella
(Nyl.) Gotth. Schneid. ex Brako, Fl. Neotropica 55: 48 (1991), nom. illeg. – Lecidea glabella Nyl.,
Sert. Lich. Trop.: 37 (1891), non Kremp., Verh. zool.-bot. Ges. Wien 26: 457 (1876). –
Description: BRAKO (1991, as P. glabella).
(Colour Plate 15D)
Chemistry: no lichen substances (by TLC).
Notes: The species is characterized by the mostly adnate, medium sized (up to 0.6
mm wide), rather thick, shiny squamules, thin, reddish brown prothallus, absence
of vegetative dispersal units, short ellipsoid ascospores (4.5–6.5 2.5–3.5 μm,
BRAKO 1991), and absence of lichen substances. It may be confused mainly with
P. breviuscula and P. minor, but the former differs in having larger squamules (up
to 1 mm wide) and the latter in having smaller squamules (up to 0.1 mm wide);
both differ in having larger ascospores (9.5–13 3.5–5 μm and 7–12 × 2.5–5 μm,
respectively).
The species was treated as P. glabella by SWINSCOW & KROG (1981) and
BRAKO (1991), but the basionym Lecidea glabella Nyl. is illegitimate, being a
later homonym of Lecidea glabella Kremp. [syn. Cryptolechia myriadella (Nyl.)
D. Hawksw. & Dibben].
The species is known from Mexico, Honduras, Brazil, and the West Indies
(BRAKO 1991). New to Haiti.
Specimens examined: CUBA. Wright, Lich. Cub., Ser. 2, 142 (H-NYL 20518, holotype, TLC not
performed). HAITI. Dept. de L'Ouest, summit of small peak at N end of ridge above (E of) Foret
des Pins, 5800–5900 ft, 1958, Imshaug & Wetmore 23050 (MSC).
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23. Phyllopsora minor Brako
Mycotaxon 35: 15 (1989). – Lecidea corallina var. schizophylloides Vain., J. Bot. 34: 106 (1896).
– Type: SAINT VINCENT & THE GRENADINES (see SWINSCOW & KROG 1981). – Description:
BRAKO (1991).
(Colour Plates 15E)
Chemistry: no lichen substances (by TLC).
Notes: This poorly known species is characterized by the effuse thallus consisting
of irregularly oriented, narrow (up to 0.1 m wide) squamules which are closely
adnate to a well developed, reddish brown prothallus, the lack of vegetative
dispersal units, the medium to dark brown apothecia, the ellipsoid (7–12 × 2.5–5
μm, BRAKO 1991) ascospores, and the lack of lichen substances. It is known only
from the type material. See P. microsperma for discussion.
Specimens examined: SAINT VINCENT & THE GRENADINES. Elliott 261[a] (TUR-V 22612,
lectotype; BM, isolectotype).
24. Phyllopsora ochroxantha (Nyl.) Zahlbr.
Catal. Lich. Univers. 10 (24): 377 (1939). – Lecidea ochroxantha Nyl., Annls Sci. Nat., Bot., sér.
4, 11: 223 (1859). – Type: BOLIVIA (see SWINSCOW & KROG 1981). – Lecidea glabriuscula Nyl.,
Sertum Lich. Trop.: 48 (1891). – Descriptions: TIMDAL (2008b), ELIX (2009).
Chemistry: phyllopsorin (major), chlorophyllopsorin (minor to major), argopsin
(occasional trace), norargopsin (occasional trace), and unknown compounds
(occasional traces).
Notes: The present circumscription of P. ochroxantha corresponds to P. corallina
var. ochroxantha chemical strains I and II in BRAKO (1991). Strain III is here
regarded as a distinct species, P. swinscowii, see that species for discussion.
Phyllopsora ochroxantha differs from P. corallina mainly in the presence of
phyllopsorin and chlorophyllopsorin, but also in having a thicker, always reddish
brown prothallus, a thicker (20–30 μm vs. 15–20 μm) upper cortex which is
composed of more thick-walled lumina (type 1–2 vs. type 2), and in having darker
brown apothecia.
In some specimens there are traces of unknown compounds below phyllopsorin
and chlorophyllopsorin on the chromatograms. They are possibly identical with
BRAKO’s (1991) methyl-phyllopsorate and methyl-chlorophyllopsorate. The
difference between Brako’s strain I and II was based on the presence of those two
compounds, but I found the occasional traces insufficient for distinguishing two
chemotypes. Atranorin, methyl 2,7-dichloropsoromate, pannarin, and zeorin
(reported by BRAKO 1991 and/or ELIX 2009) were not found in the West Indian
material.
The species is widely distributed in the Neotropics (BRAKO 1991) and also
occurs in Australia (ELIX 2009). New to Guadeloupe, Haiti, and Trinidad and
Tobago.
Specimens examined: CUBA. Buck 7620, 7684 (NY), Harris 14181, 14190, 14282, 14285, 14350,
14351, 14359, 14410, 14445, 14523, 14644, 14654 (NY), Imshaug 24843, 25111 (MSC), Wright,
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Lich. Cub., Ser. 2, 105 (H-NYL 20534, holotype of Lecidea glabriuscula Nyl.), Ser. 2, 726 (H-
NYL 20533, TLC not performed). HAITI. Imshaug & Wetmore 3060, 22719, 22919 (MSC).
DOMINICAN REPUBLIC. Buck 14509 (NY). GRENADA: Imshaug 16268 (MSC). JAMAICA.
Cummings 44 (MSC, NY), Imshaug 12974, 12982, 13389, 13769, 14099, 14213, 14976, 15515
(MSC). TRINIDAD & TOBAGO. Imshaug & Imshaug 31260, 31657-immixture, 31707, 32018
(MSC), Rui & Timdal 10772, 10833, 10849 (O).
25. Phyllopsora parvifolia (Pers.) Müll. Arg.
Bull. Herb. Boissier 2 (2): 90 (1894). – Lecidea parvifolia Pers. in Gaudichaud, Voy. Uranie (5):
192 (1827). – Type: BRAZIL (see BRAKO 1991). – Phyllopsora parvifolia var. hirtella B. de Lesd.,
Revue Bryol. Lichénol. 7: 60 (1934). – Description: ELIX (2009).
(Colour Plate 15F)
Chemistry: no lichen substances (by TLC).
Notes: This species corresponds to P. parvifolia var. parvifolia in BRAKO (1991).
Var. breviuscula is here treated as a distinct species, P. breviuscula, see that
species for discussion.
Phyllopsora parvifolia is a common neotropical species and also occurs in
Africa (Tanzania; BRAKO 1991), Australia and in islands in the Pacific Ocean
(ELIX 2009). The previous record from Jamaica (RIDDLE 1912, i.e., Cummings 44,
five envelopes in NY) is a mixture of P. chodatinica, P. confusa, P. ochroxantha,
and P. phaeobyssina. I have not examined material from the Bahamas,
Guadeloupe and Martinique (cf. IMSHAUG 1957). New to Haiti, Jamaica,
Netherlands Antilles (St. Eustatius), Saint Kitts and Nevis, Saint Lucia, and
Trinidad and Tobago.
Specimens examined: CUBA. Harris 14410a (NY), Hioram 9545 (NY, isolectotype of P.
parvifolia var. hirtella B. de Lesd., TLC not performed) Imshaug 24603, 24734, 24907 (MSC),
Johnson s.n. (NY). HAITI. Imshaug & Wetmore 3187-immixture, 22948 (MSC). DOMINICAN
REPUBLIC. Buck 14655 (NY), Harris 15677, 15683, 15774, 15861, 15870, 20496, 20587, 26575
(NY), Liogier 15725 (NY). PUERTO RICO. Harris 22005, 22032, 24334 (NY), Heller 448 (NY).
JAMAICA. Imshaug 14105, 14205, 14380 (MSC). SAINT KITTS & NEVIS. Harris 37916, s.n. (NY).
SAINT LUCIA. Imshaug & Imshaug 29624 (MSC). NETHERLANDS ANTILLES. Sipman 56790 (B).
TRINIDAD & TOBAGO. Imshaug & Imshaug 31151, 31399, 31751, 31757, 31842, 32033 (MSC),
Rui & Timdal 10785, 10798, 10860, 10867, 10870 (O).
26. Phyllopsora parvifoliella (Nyl.) Müll. Arg.
In Durand, T. & Pittier, H., Prim. Fl. Costaric., Lich., Fasc. 2: 10 (1894). – Lecidea parvifoliella
Nyl., Annls Sci. Nat., Bot., sér. 4, 19: 339 (1863). – Type: CUBA (see SWINSCOW & KROG 1981,
BRAKO 1991). – Description: TIMDAL (2008b).
(Colour Plate 16A)
Chemistry: parvifoliellin (major) and atranorin (absent to minor).
Notes: The species is recognized by the medium-sized, elongated squamules, the
terete to flattened isidia attached mainly terminally on the squamules, the short
ascospores (5–6.5 × 2–2.5 μm), and by containing atranorin and parvifoliellin.
The latter compound is elsewhere known only from P. rappiana, a species which
340
eschweizerbart_xxx

seems to differ in forming marginal and laminal, globose to terete isidia and in
having longer ascospores (6.5–10 × 2.5–3.5 μm; cf. ELIX 2009). See also P.
phaeobyssina for discussion.
Phyllopsora parvifoliella is neotropical, occurring from Florida to Brazil
(BRAKO 1991). New to Haiti.
Specimens examined: CUBA. Wright, Lich. Cub. 182 (H-NYL 20545, holotype; BM, UPS
isotypes). HAITI. Imshaug & Wetmore 2873, 23051, 23059 (MSC). NETHERLANDS ANTILLES.
Sipman 14820, 14833, 14852, 15069, 56742a (B).
27. Phyllopsora pertexta (Nyl.) Swinscow & Krog
The Lichenologist 13: 244 (1981). – Lecidea pertexta Nyl., Annls Sci. Nat., Bot., sér. 4, 19: 347
(1863). – Type: CUBA, “in ins. Cuba”, Wright s.n. (H-NYL 17344, holotype) [no lichen
substances]. – Lecidea microphyllina var. subgranulosa Tuck., Proc. Am. Acad. Arts. Sci. 6: 278
(1864).
(Colour Plate 16B)
Description: Prothallus often well developed, white to dark brown. Thallus
effuse, crustose, not radiating at the margin. Areolae small, up to 0.06 mm wide,
discrete when young, soon adjoining to form a more or less continuous crust,
brownish green, dull, epruinose, pubescent along the margin. Isidia and soredia
lacking. Upper cortex poorly defined, resembles type 2, 5–10 μm thick, not
containing crystals. Medulla not containing crystals, PD–, K–. Apothecia
common, up to 1.5 mm diam., plane when young, becoming weakly to moderately
convex, reddish brown, rounded to irregular, simple to conglomerate, with a thin,
more or less excluding, paler, sometimes finely pubescent margin. Excipulum
pale brown to colourless, patchily reddish brown, K+ purple (red pigment).
Hypothecium pale brown or patchily reddish brown, K+ purple (red pigment) in
upper part, pale brown to colourless in lower part. Epithecium colourless, K–. No
crystals in apothecium. Ascospores acicular, simple, with up to 3 pseudosepta,
28–41 × ca. 1.5–2 μm (n = 22).
Chemistry: no lichen substances (by TLC).
Notes: Phyllopsora cognata differs in having yellowish brown apothecia, shorter
ascospores (19–30 μm long), and in containing atranorin. The paleotropical P.
borbonica Timdal & Krog is closely related or perhaps synonymous with P.
pertexta, but differs in having a deep reddish brown, K+ purple hypothecium.
According to SWINSCOW & KROG (1981), the hypothecium of P. pertexta is
colourless, but the present examination of the West Indian material showed patchy
occurrences of a red, K+ purple pigment in the hypothecium of older apothecia of
P. pertexta.
According to EKMAN (1996), Bacidia prasinata (Tuck.) Coppins is a later
synonym of P. pertexta. That species was described on material from Venezuela
(not seen by me) and is the holotype of the genus Sporacestra A. Massal. If the
current broad concept of Phyllopsora is adopted, the name Phyllopsora (priority
1894) should be conserved against both Sporacestra (priority 1860) and Triclinum
Fée (priority 1825; see TIMDAL 2008b).
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The species was previously known from Wright’s material from Cuba,
Venezuela (see above), and Samoa [MÜLLER 1896, as Psorella pertexta (Nyl.)
Müll. Arg.; not examined by me]. New to Jamaica and Puerto Rico.
Specimens examined: CUBA. Buck 23433 (NY), Harris 14514, 14632 (NY), Hioram 12577 (NY,
TLC not performed), Imshaug 24853 (MSC), Pérez-Ortega s.n. (hb Pérez-Ortega), Shafer 3359
(NY), Wright s.n. (H-NYL 17344, holotype), Wright, Lich. Cub. 214, 215, 216 (BM, UPS,
isotypes of Lecidea microphyllina var. subgranulosa Tuck.). PUERTO RICO. Buck 3516 (NY).
JAMAICA. Buck 5697, Cummings 118 (NY), Imshaug 13800, 14202 (MSC).
28. Phyllopsora phaeobyssina (Vain.) Timdal comb. et stat. nov.
Lecidea breviuscula var. phaeobyssina Vain., Ann. Acad. Sci. Fenn., ser. A, 6, 7: 127 (1915). –
Phyllopsora corallina var. phaeobyssina (Vain.) Brako, Mycotaxon 35: 14 (1989). – Type:
GUADELOUPE, Houelmont, sur un Coffea arabica, Duss 481 (TUR-V 22602, holotype) [argopsin
(major)].
(Colour Plate 16C)
Description: Prothallus well developed, dark brown. Thallus effuse, squamulose,
without radiating marginal squamules. Squamules medium sized, up to 0.5(–1)
mm wide, adnate, isodiametrical to elongate, more or less scattered, plane to
weakly convex, crenulate to deeply divided; upper side medium brown to dark
brown, shiny, epruinose, glabrous; margin concolorous with the upper side,
glabrous. Isidia common, more or less flattened, long, usually branched,
sometimes coralloid. Upper cortex of type (1–)2, 40–60 μm thick, not containing
crystals. Medulla containing crystals not dissolving in K, PD+ orange, K–.
Apothecia common, up to 0.8(–1.2) mm diam., weakly to moderately convex,
medium brown, mostly rounded and simple, with a paler, non-pubescent margin.
Excipulum and hypothecium pale brown throughout, K–. Epithecium dark brown,
K–. No crystals in apothecium. Ascospores ellipsoid, simple, 5.5–7.5 2.5–3 μm
(n = 50).
Chemistry: argopsin (major) and norargopsin (absent to minor).
Notes: This species was treated as P. corallina var. phaeobyssina by BRAKO
(1991), but differs from P. corallina in having darker brown, more shiny
squamules with more or less flattened isidia, shorter ascospores, and in containing
argopsin and often norargopsin. In P. corallina, the isidia are terete, the
ascospores narrowly ellipsoid to shortly bacilliform, 6.5–14.5 2–2.5 μm, and the
squamules do not contain lichen substances.
Phyllopsora santensis, treated as P. corallina var. santensis (Tuck.) Brako by
BRAKO (1991), differs in forming more greenish brown, often pale-edged, less
shiny, more isodiametrical squamules with shorter, cylindrical isidia. The upper
cortex of P. santensis is thinner (20–30 μm) and the ascospores are longer, 7.5–
10.5 2.5–3 μm. Phyllopsora santensis always contains norargopsin in addition
to argopsin, but this compound is often missing in P. phaeobyssina.
Phyllopsora parvifoliella has mostly flattened isidia and short ascospores like
P. phaeobyssina, but differs in being green to greenish brown and less shiny,
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eschweizerbart_xxx

having a thinner upper cortex (15–30 μm) of type 1, and in containing atranorin
and parvifoliellin.
The species is neotropical. New to Dominica, the Dominican Republic,
Grenada, Haiti, Jamaica, Saint Lucia, and Trinidad and Tobago.
Specimens examined: HAITI. Imshaug & Wetmore 2945-immixture, 3179 (MSC). DOMINICAN
REPUBLIC. Imshaug & Wetmore 3732 (MSC). PUERTO RICO. Harris 21987 (NY), 27320 (NY, O).
JAMAICA. Cummings 44 (NY), Imshaug 14030, 14647 (MSC). GUADELOUPE. Duss 481 (TUR-V
22602, holotype). DOMINICA. Imshaug & Imshaug 33192 (MSC). SAINT LUCIA. Imshaug &
Imshaug 29802, 29857, 30025, 30181 (MSC). SAINT VINCENT & THE GRENADINES. Elliott 261bimmixture
(BM, TUR-V 22612). GRENADA. Imshaug 16081, 16265 (MSC). TRINIDAD & TOBAGO.
Imshaug & Imshaug 31441, 31452, 31657, 32021-immixture (MSC), Rui & Timdal 10756, 10872,
10876, 10882 (O).
29. Phyllopsora porphyromelaena (Vain.) Zahlbr.
Catal. Lich. Univers. 4 (26): 401 (1926). – Lecidea porphyromelaena Vain., Acta Soc. Sci.
Fennicae 15: 113 (1920). – Type: THE PHILIPPINES, prov. Bataan, Luzon, Mount Mariveles, XII.
1908, Merrill 6273 (TUR-V 22619, lectotype selected by SWINSCOW & KROG 1981, not seen),
6256 (syntype, BM) [TLC not performed]. – Phyllopsora albicans sensu SWINSCOW & KROG
(1981), TIMDAL & KROG (2001), ELIX (2009), non Müll. Arg. – Phyllopsora buettneri var. glauca
B. de. Lesd., Revue Bryol. Lichénol. 7: 60 (1934). – Descriptions: TIMDAL & KROG (2001), ELIX
(2009), both as P. albicans.
Chemistry: chemotype 1 (pantropical): argopsin (major), norargopsin (minor to
major) and zeorin (absent to major, always present in the West Indies); chemotype
2 (paleotropical): argopsin (major) and pannarin (major).
Notes: The species is recognized by the large, elongated, ascending, epruinose
squamules breaking off lacinules at the tips and by containing argopsin and
additional compounds.
This species was called P. albicans Müll. Arg. by SWINSCOW & KROG (1981),
TIMDAL & KROG (2001), and ELIX (2009). BRAKO (1991), however, placed P.
albicans in synonymy with P. santensis, and my recent examination of the
holotype of P. albicans (Costa Rica, Pitt 5474, G) showed that this is correct.
BRAKO (1991) treated P. porphyromelaena as P. buettneri var. glauca (B. de
Lesd.) Brako chemical strains I and III. Strain I was characterized by argopsin,
norargopsin, vicanicin, norvicanicin, and ± zeorin, and strain III by argopsin and ±
zeorin. My re-examination of the West Indian material showed a constant
presence of norargopsin but did not confirm the presence of vicanicin or
norvicanicin in any specimen of P. porphyromelaena (including the isolectotype
of P. parvifolia var. glauca B. de Lesd., Hioram 9098, NY). A distinction
between two chemotypes of P. porphyromelaena in the Neotropics is hence not
supported here. Probably the report of vicanicin and norvicanicin in this species
was based on erroneously identified material of the taxon here called P. buettneri
chemotype 3.
Brako's chemical strain II is here treated as P. chodatinica, see that species for
discussion. For P. melanoglauca, listed by BRAKO (1991) as a synonym of P.
buettneri var. glauca, see P. buettneri.
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The species is pantropical and here reported as new to Dominica, Guadeloupe
and Saint Lucia. BRAKO's (1991) six specimens from the Dominican Republic
belong in P. buettneri (Buck 8348, Harris 15563) and P. hispaniolae (Buck
14618, Harris 14634, 20455, 20672).
Specimens examined: – Chemotype 1: CUBA. Harris 14325, 14379, 14656 (NY), Hioram 9098
(NY, isolectotype of P. parvifolia var. glauca B. de Lesd.), Imshaug 24680 (MSC), Leon et al.
10232 (NY). PUERTO RICO. Harris 22217, 22234, 22495 (NY). JAMAICA. Imshaug 14051 (MSC).
GUADELOUPE. Imshaug & Imshaug 33592, 33593, 33594, 33609, 33613 (MSC). DOMINICA.
Imshaug & Imshaug 32953, 33086 (MSC). SAINT LUCIA. Imshaug & Imshaug 29832, 29926,
30250 (MSC). GRENADA. Broadway s.n. (NY), Imshaug 16101, 16121, 16263 (MSC). TRINIDAD
& TOBAGO. Fleming s.n. (NY), Imshaug & Imshaug 31709, 31726, 31887, 31890, 31918, 32171
(MSC), Rui & Timdal 10753 (O).
30. Phyllopsora pyrrhomelaena (Tuck.) Swinscow & Krog
The Lichenologist 13: 244 (1981). – Biatora pyrrhomelaena Tuck., Am. J. Sci. Arts, ser. 2, 28:
205 (1859). – Type: CUBA (see SWINSCOW & KROG 1981).
(Colour Plate 16C)
Description: Prothallus thick, dark brown at the periphery, reddish brown in
inner part. Thallus circular to effuse, crustose to almost squamulose. Areolae/
squamules small, up to 0.2 mm wide, adnate, scattered to contiguous or fusing,
isodiametrical, rounded to crenulate; upper side pale green (old herbarium
material), dull, epruinose, glabrous; margin paler than or concolorous with the
upper side, pubescent. Vegetative dispersal units absent. Upper cortex of type 2,
5–10 μm thick, not containing crystals. Medulla (mainly lower part) containing
orange crystals dissolving in K, PD–, K+ purple. Apothecia common, up to 0.8
mm diam., plane to weakly convex, brownish black, rounded to crenulate or
lobate, simple to conglomerate, with a persistent, shiny, castaneous brown,
pubescent margin. Excipulum dark reddish brown (dark olivaceous brown after
orange pigments are dissolved), containing orange crystals dissolving in K with a
strong K+ purple effusion. Hypothecium olivaceous brown, not containing
crystals, K–. Epithecium colourless, K–. Ascospores narrowly ellipsoid, simple,
6–8 2.5–3 μm (n = 20).
Chemistry: norsolorinic acid (major) and at least three additional pink pigments
(minor to trace).
Notes: Phyllopsora nigrocincta Timdal, described from Peru (TIMDAL 2008b), is
closely related or perhaps a chemical strain of this species. It differs in containing
fumarprotocetraric acid in addition to norsolorinic acid. The latter compound,
reported as an unknown orange pigment by TIMDAL (2008b), was identified by
ELIX (in litt.) in paratype material of P. nigrocincta.
Phyllopsora pyrrhomelaena is known only from Wright's material from Cuba
and from one collection from Isle of Pines, Cuba (RIDDLE 1923, material not
examined by me). According to ZAHLBRUCKNER [1924–1925, as Lecidea
pyrrhomelaena (Tuck.) Tuck.], Lecidea circumpurpurans Nyl. is synonymous
with this species (described from Brazil, not examined by me).
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Specimens examined: CUBA. Monte Verde Woods, on trunks of trees near the ground, Wright,
Lich. Cub. 178 (FH 286104, holotype; FH 197468, UPS isotypes, TLC not performed); s. loc.,
Wright, Lich. Cub. Ser. 2: 124 (FH 28102, H-NYL 20498); ad truncos in montib. Cubae, Wright
s.n. (H-NYL 20500, TLC not performed).
31. Phyllopsora santensis (Tuck.) Swinscow & Krog
The Lichenologist 13: 236 (1981). – Lecidea santensis Tuck., Am. J. Sci. Arts, ser. 2, 25: 428
(1858). – Type: U.S.A., South Carolina (see SWINSCOW & KROG 1981). – Phyllopsora albicans
Müll. Arg., Bull. Soc. R. Bot. Belg. 32: 132 (1893). – Descriptions: TIMDAL (2008b), ELIX (2009).
Chemistry: argopsin (major) and norargopsin (minor to submajor).
Notes: This species was treated as P. corallina var. santensis (Tuck.) Brako by
BRAKO (1991). She accepted two chemical strains, one with argopsin,
norargopsin, ±atranorin, and ±zeorin, and one with argopsin only. I found
argopsin and norargopsin in the three West Indian specimens, but no atranorin or
zeorin.
The species differs from P. corallina mainly in the presence of argopsin and
norargopsin. Phyllopsora santensis generally forms rather discrete, pale-edged
squamules, often with only a few isidia, and medium brown to dark brown
apothecia with a darker margin, whereas P. corallina generally forms more
contiguous or imbricate squamules with a concolorous margin and numerous
isidia, and pale to medium brown apothecia with a paler margin. See also P.
phaeobyssina for taxonomic discussion. BRAKO (1991) correctly included
Phyllopsora albicans in P. santensis, see discussion under P. porphyromelaena.
Phyllopsora santensis is widely distributed in the Americas (BRAKO 1991) and
also occurs in Asia and Australia (ELIX 2009). BRAKO’s (1991) specimen from the
Dominican Republic (Harris 20779) belongs in P. glaucella, whereas that from
Jamaica (Wight 10, FH) was not examined by me. New to Cuba.
Specimens examined: BAHAMAS. Britton 6633 (NY). CUBA. Buck 23555 (NY), Ranker &
Lemieux 1702 (NY).
32. Phyllopsora swinscowii Timdal & Krog
Mycotaxon 77: 88 (2001). – Type: MAURITIUS (see TIMDAL & KROG 2001). – Descriptions:
TIMDAL & KROG (2001), TIMDAL (2008b), ELIX (2009).
Chemistry: methyl 2,7-dichloronorpsoromate (major) and 2,7-dichloropsoromate
(minor to major).
Notes: This species corresponds to P. corallina var. ochroxantha chemical strain
III in BRAKO (1991). It differs from P. ochroxantha apparently only in the
secondary chemistry and is perhaps merely a chemotype of that species.
Phyllopsora swinscowii is pantropical (BRAKO 1991, TIMDAL & KROG 2001, ELIX
2009).
Specimens examined: BAHAMAS. Brace 6881 (NY). CUBA. Britton et al. 15586 (NY), Tønsberg
37719, 37746, 37815, 37817, 37831, 37848, 37947 (BG). DOMINICAN REPUBLIC. Harris 15383
(BM, NY). TRINIDAD & TOBAGO. Fleming s.n. (NY), Imshaug & Imshaug 31449, 31450, 31488,
32021 (MSC), Rui & Timdal 10766 (O).
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33. Phyllopsora teretiuscula Timdal sp. nova
Phyllopsorae furfuraceae et P. tobagensi similis, sed squamis majoribus,
teretiusculis, hypothecio pallidiori, et thallo argopsinum, norargopsinum et
interdum chlorophyllopsorinum continenti. – Type: CUBA, Pinar del Río, Reserva de
la Biosfera Sierra del Rosario, N of and near lake “La Palma”, near river,
downstream from the path/road, 22°51.31’N, 82°56.25’W, 140−145 m, over mosses
on trunk of Roystonea regia in mixed hardwood forest, III. 2007, Tønsberg 37814
(BG-L-87831, holotype); Tønsberg 37848b, 37850, 37980 (BG, paratypes)
[argopsin (major) and norargopsin (minor)].
(Colour Plate 16E)
Description: Prothallus thin, reddish brown. Thallus effuse, crustose to
squamulose, without radiating marginal squamules. Areolae small, up to 0.2(–0.3)
mm diam., adnate and circular when young, soon proliferating into a mat of richly
branched, narrow to almost terete, coralloid, up to 0.1 mm wide squamules; upper
side pale green, patchily stained reddish brown, slightly shiny, epruinose,
glabrous; margin concolorous with upper side, glabrous or finely pubescent. Isidia
and soredia lacking. Upper cortex of type 2, 30–40 μm thick, not containing
crystals. Medulla containing crystals which are not dissolving in K, PD+ orange,
K–. Apothecia not common (only immature seen), up to 0.5 mm diam., weakly
convex, medium brown to dark brown, rounded, simple, with a darker, thin, finely
pubescent, soon excluded margin. Excipulum reddish brown, K+ purple in the
rim, pale brown, K– in inner part. Hypothecium pale brown, K–. Epithecium pale
brown, K–. No crystals in the apothecium. Asci and ascospores not seen.
Chemistry: argopsin (major), norargopsin (minor), and chlorophyllopsorin
(absent to minor).
Notes: The species is recognized by the coralloid thallus containing argopsin,
norargopsin, and sometimes low amounts of chlorophyllopsorin. It may be
confused with P. furfuracea and P. tobagensis, but differs in the secondary
chemistry, in having a pale (not reddish brown) hypothecium, and in forming
larger areolae and thicker, not fully terete squamules. The terete thallus parts are
called isidia in P. furfuracea and P. tobagensis and squamules in P. teretiuscula,
but this does not infer a basic structural difference. It seems likely that thallus
fragments function as vegetative diaspores in P. teretiuscula. See also P.
hispaniolae for discussion.
The species is known from three sites within 350 m in one locality in Cuba.
The species is corticolous or growing over corticolous bryophytes.
34. Phyllopsora tobagensis Timdal sp. nova
Phyllopsora furfuracea et speciebus affinibus similis, sed ascosporis acicularibus et
thallo acidum hyperlatolicum, acidum perlatolicum et acidum superlatolicum
continenti. – Type: TRINIDAD & TOBAGO, Tobago, Parish of St. Paul, along
Roxborough – Parlatuvier Road, 11°16.80’N, 60°36.66’W, 500–520 m, on tree
trunk in rainforest, III. 2008, Rui & Timdal 10764 (O-L152061, holotype; CANB,
isotype); Rui & Timdal 10747, 10757, 10768, 10839, 10873, 10877 (O-L152044,
L152054, L152065, L152136, L152170, L152174). [hyperlatolic acid (major),
perlatolic acid (minor), and superlatolic acid (minor); according to ELIX in litt.].
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(Colour Plate 16F)
Description: Prothallus thin or indistinct, white to dark brown. Thallus effuse,
formed partly by minute areolae and partly by isidia developing directly from the
prothallus, not radiating at the margin; isidia often dominating. Areolae up to 0.06
mm wide, adnate, isodiametrical, plane to weakly convex, medium to dark green,
more or less shiny, epruinose, glabrous. Isidia abundant, cylindrical, long, 0.03–
0.05 mm wide, medium to dark green, simple to slightly branched. Upper cortex
poorly defined, 5–10 μm thick, not containing crystals. Medulla containing
crystals dissolving in K; PD–, K–. Apothecia not common (seen in the type only),
up to 1 mm diam., plane when young, later weakly to moderately convex, dark
brown, shiny, rounded, mostly simple, with a distinct, brownish black, not
pubescent margin when young, later more or less immarginate. Excipulum dark
reddish brown in inner part, paler in the rim, containing colourless crystals dissolving
in K. Hypothecium reddish brown in lower part, deeper pigmented in upper
part, containing some scattered colourless crystals dissolving in K. Epithecium
colourless. Reddish brown pigment in apothecium K+ purple. Ascospores
acicular, simple, usually with 3 pseudosepta, 27–42 ca. 1.5 μm (n = 20).
Chemistry: hyperlatolic acid (major), perlatolic acid (minor) and superlatolic acid
(minor).
Notes: The species is morphologically and anatomically similar to P. furfuracea
and the paleotropical P. dolichospora Timdal & Krog, P. foliatella Elix, P.
homosekikaica Elix, and P. methoxymicareica Elix, but differs from them all both
in chemistry and in ascospore size. In P. furfuracea, the ascospores are narrowly
ellipsoid, 7–13 2–3 μm, and the thallus contains furfuraceic acid or no lichen
substances; in P. dolichospora the spores are bacilliform, 16–25 2–3 μm, and
the thallus contains methyl furfuraceiate (major), methyl homofurfuraceiate
(major), and furfuraceic acid (minor); in P. foliatella the spores are 11–18 μm
long and the thallus does not contain lichen substances; in P. homosekikaica the
spores are narrow-ellipsoid, 7–11 2–3 μm, and the thallus contains
homosekikaic acid (major) and hyperhomosekikaic acid (major); and in P.
methoxymicareica the spores are ellipsoid, 10–14 3–5 μm, and the thallus
contains methoxymicareic acid (major) and hydroxymicareic acid (trace) (TIMDAL
& KROG 2001, ELIX 2006b, 2006c, 2008). See also P. teretiuscula for discussion.
Phyllopsora tobagensis is known from seven collections along three trails in
the Main Ridge rainforest in Tobago, collected at altitudes of about 400–520 m.
The species is corticolous or growing over corticolous bryophytes.
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Key to the West Indian species of Phyllopsora
(Chemical character states are restricted to the West Indian chemotypes)
1 Thallus sorediate .............................................................................................. 2
1' Thallus not sorediate ........................................................................................ 3
2 Soralia labriform; squamules rounded, containing methyl barbarate .................
...................................................................................................... P. labriformis
2' Soralia starting from the lower side near the tip of ascending squamules, later
spreading to the upper side; squamules elongated, containing homosekikaic
and sekikaic acids .................................................................... P. leucophyllina
3 Thallus with globose, cylindrical or slightly flattened isidia ........................... 4
3' Thallus with lacinules or phyllidia, or vegetative diaspores lacking ............. 18
4 Thallus crustose, often consisting only of isidia on the prothallus .................. 5
4' Thallus squamulose .......................................................................................... 7
5 Ascospores acicular, 27–42 ca. 1.5 μm; thallus containing hyperlatolic acid
and related compounds ................................................................. P. tobagensis
5' Ascospores ellipsoid, up 13 μm long; hyperlatolic acid lacking ..................... 6
6 Ascospores 7–13 × 2–3 μm; hypothecium dark reddish brown; thallus
containing furfuraceic acid or lacking lichen substances ............. P. furfuracea
6' Ascospores 5–7.5 2.5–3 μm; hypothecium dark yellowish brown; thallus
lacking lichen substances ........................................................ P. canoumbrina
7 Thallus effuse or incompletely circular, with radiating marginal squamules on
a thick, white prothallus ................................................................. P. byssiseda
7' Thallus effuse; squamules not radiating; prothallus white or reddish brown .. 8
8 Thallus lacking lichen substances .................................................................... 9
8' Thallus containing lichen substances ............................................................. 11
9 Isidia globose to shortly cylindrical; hypothecium dark yellowish brown;
prothallus thin, white ............................................................................ P. kalbii
9' Isidia long, cylindrical to partly flattened; hypothecium medium brown to
colourless; prothallus variable ........................................................................ 10
10 Squamules pale green, closely adnate or sometimes slightly ascending, mainly
isodiametrical; prothallus thick, reddish brown; isidia cylindrical to partly
flattened .................................................................................... P. intermediella
10' Squamules medium to brownish green, mainly ascending, elongated and partly
imbricate; prothallus variable; isidia cylindrical ............................. P. corallina
11 Ascospores bacilliform to acicular (more than 25 μm long); thallus containing
i.a. lobaric acid ........................................................................ P. cinchonarum
11' Ascospores ellipsoid to shortly bacilliform (up to15 μm long); thallus not con-
taining lobaric acid) ........................................................................................ 12
12 Medulla K+ red (norstictic acid) ................................................... P. imshaugii
12' Medulla K– ..................................................................................................... 13
13 Medulla PD– .................................................................................................. 14
13' Medulla PD+ yellow or orange ...................................................................... 15
14 Thallus containing vicanicin and norvicanicin; isidia cylindrical; squamules
mostly adnate; ascospores narrowly ellipsoid, 6.5–9 μm long ...... P. glaucella
14' Thallus containing atranorin and parvifoliellin; isidia cylindrical to flattened;
squamules adnate to ascending; ascospores broadly ellipsoid, 5–6.5 μm long ..
.................................................................................................... P. parvifoliella
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15 Medulla PD+ yellow (2,7-dichloropsoromate and 2,7-dichloronorpsoromate) ..
....................................................................................................... P. swinscowii
15' Medulla PD+ orange ...................................................................................... 16
16 Thallus containing mainly phyllopsorin and chlorophyllopsorin .......................
................................................................................................... P. ochroxantha
16' Thallus containing argopsin and often norargopsin ....................................... 17
17 Squamules medium brown to dark brown, shiny, with a concolorous margin;
isidia more or less flattened; ascospores 5.5–7.5 μm long; norargopsin
sometimes absent .................................................................... P. phaeobyssina
17' Squamules greenish brown, often with a paler margin; isidia cylindrical;
ascospores 7.5–10.5 μm long; norargopsin always present ........... P. santensis
18 Thallus crustose, apothecia common, vegetative diaspores absent ............... 19
18' Thallus squamulose, apothecia and vegetative diaspores present or absent .......
......................................................................................................................... 21
19 Ascospores narrowly ellipsoid, 6–8 2.5–3 μm; exciple and lower part of
medulla containing orange crystals dissolving in K, K+ purple (norsolorinic
acid) ...................................................................................... P. pyrrhomelaena
19' Ascospores acicular, more than 19 μm long; exciple and medulla not
containing orange crystals .............................................................................. 20
20 Apothecia yellowish brown; ascospores 19–30 μm long; thallus containing
atranorin ........................................................................................... P. cognata
20' Apothecia reddish brown; ascospores 28–41 μm long; thallus lacking lichen
substances ......................................................................................... P. pertexta
21 Thallus lacking lichen substances .................................................................. 22
21' Thallus containing lichen substances ............................................................. 30
22 Ascospores acicular, more than 19 µm long .................................................. 23
22' Ascospores ellipsoid to bacilliform, up to 18 µm long .................................. 24
23 Squamules ascending, deeply incised or divided, often with vein-like
supportive tissue ............................................................................... P. lacerata
23' Squamules mostly adnate, crenulate to lobed, without vein-like tissue ..............
................................................................................................. P. microphyllina
24 Thallus with phyllidia in inner part and elongated, more less radiating
marginal squamules ....................................................................... P. parvifolia
24' Thallus without phyllidia (but sometimes with lacinules, i.e. apical sections of
squamules breaking off), marginal squamules of various shapes .................. 25
25 Squamules mostly adnate, without lacinules ................................................. 26
25' Squamules mostly ascending, lacinulate ........................................................ 28
26 Ascospores ellipsoid, up to 6.5 µm long .................................. P. microsperma
26' Ascospores ellipsoid to fusiform, more than 7 µm long ................................ 27
27 Thallus composed of large, up 0.5(–1) mm wide squamules ..... P. breviuscula
27' Thallus composed of small, up to 0.1 mm wide squamules ................ P. minor
28 Hypothecium evenly dark reddish brown, K+ purple ............... P. chlorophaea
28' Hypothecium colourless to yellowish brown, K– .......................................... 29
29 Thallus consisting of squamules ascending from a more or less continuous
crust of adnate areolae; prothallus thick, reddish brown ............ P. longiuscula
29' Thallus consisting of ascending squamules only; prothallus white or reddish
brown, thin or thick .......................................................................... P. confusa
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30 Thallus pruinose, containing i.a. pannarin or vicanicin .................. P. buettneri
30' Thallus not pruinose, not containing pannarin and vicanicin ........................ 31
31 Thallus containing i.a. xanthones ............................................... P. chodatinica
31' Thallus not containing xanthones ................................................................... 32
32 Squamules plane, containing i.a. zeorin ........................... P. porphyromelaena
32' Squamules convex to terete, not containing zeorin ........................................ 33
33 Tips of squamules paler than inner part, roughly pubescent; thallus containing
argopsin and chlorophyllopsorin ................................................ P. hispaniolae
33' Tips of squamules concolorous with inner part, glabrous or finely pubescent;
thallus containing argopsin, norargopsin and sometimes chlorophyllopsorin …
..................................................................................................... P. teretiuscula
Acknowledgements
I am grateful to Alan M. Fryday for suggesting the study of Imshaug’s Phyllopsora
material and for arranging the loan from MSC, to John A. Elix for identification of lichen
substances in selected specimens, and to Sergio Pérez-Ortega and the curators of the
herbaria B, BG, BM, FH, G, H, NY, S, TUR, and UPS for loan of material or for access
to the collections at visits.
References
BRAKO, L. (1989): Reevaluation of the genus Phyllopsora with taxonomic notes and
introduction of Squamacidia, gen. nov. – Mycotaxon 35: 1–19.
BRAKO, L. (1991): Phyllopsora (Bacidiaceae). – Flora Neotropica Monograph 55. – New
York Botanical Garden, New York.
CULBERSON, C.F. (1972): Improved conditions and new data for the identification of
lichen products by a standardized thin-layer chromatographic method. – Journal of
Chromatography 72: 113–125.
CULBERSON, C.F. & JOHNSON, A. (1982): Substitution of methyl tert.-butyl ether for
diethyl ether in the standardized thin-layer chromatographic method for lichen
products. – Journal of Chromatography 238: 483–487.
CULBERSON, C.F. & KRISTINSSON, H. (1970): A standardized method for the
identification of lichen products. – Journal of Chromatography 46: 85–93.
EKMAN, S. (1996): The corticolous and lignicolous species of Bacidia and Bacidina in
North America. – Opera Botanica 127: 1–148.
ELIX, J.A. (2006a): Additional lichen records from Australia 58. New records from
Norfolk Island. – Australasian Lichenology 59: 12–15.
ELIX, J.A. (2006b): Five new species of Phyllopsora (lichenized Ascomycota) from
Australia. – Australasian Lichenology 59: 23–29.
ELIX, J.A. (2006c): Additional lichen records from Australia 56. – Australasian
Lichenology 58: 4–13.
ELIX, J.A. (2008): Lichen phytochemistry: additions and amendments I. – Australasian
Lichenology 63: 20–25.
ELIX, J. A. (2009): Phyllopsoraceae. – Flora of Australia 57: 41–59.
FRYDAY, A.M. & PRATHER, L.A. (2001): The lichen collection of Henry Imshaug at the
Michigan State University Herbarium (MSC). – The Bryologist 104: 464–467.
IMSHAUG, H.A. (1957): Catalogue of West Indian lichens. – Bulletin of the Institute of
Jamaica. Science series 6: 1–153
350
eschweizerbart_xxx

MENLOVE, J.E. (1974): Thin-layer chromatography for the identification of lichen
products. – Bulletin of the British Lichen Society 3: 3–5.
MÜLLER, J. (1896): Lichenes. – In: REINECKE, F., Die Flora der Samoa-Inseln. – Engler's
Botanische Jahrbücher für Systematik, Pflanzen-Geschichte und Pflanzen-Geographie
23: 291–299.
NYLANDER, W. (1858): Lichenes collecti in Mexico a Fr. Müller. – Flora 41: 377–381.
RIDDLE, L.W. (1912): An enumeration of lichens collected by Clara Eaton Cummings in
Jamaica I. – Mycologia 4: 125–140.
RIDDLE, L.W. (1923): The lichens of the Isle of Pines. – Mycologia 15: 68–88.
SCHNEIDER, G. (1980): Die Flechtengattung Psora sensu Zahlbruckner. – Bibliotheca
Lichenologica 13: 1–291.
SWINSCOW, T.D.V. & KROG, H. (1981): The genus Phyllopsora, with a report on the East
African species. – The Lichenologist 12: 203–247.
TIMDAL, E. & KROG, H. (2001): Further studies on African species of the lichen genus
Phyllopsora (Lecanorales). – Mycotaxon 77: 57–89.
TIMDAL, E. (2008a): Studies on Eschatogonia (Ramalinaceae) in Peru. – The
Lichenologist 40: 31–38.
TIMDAL, E. (2008b): Studies on Phyllopsora (Ramalinaceae) in Peru. – The
Lichenologist 40: 337–362.
VAINIO, E.A. (1896): Lichenes Antillarum a W.R. Elliot collecti. – The Journal of Botany
34: 31–36, 66–72, 100–107, 204–210, 258–266, 292–297.
VAINIO, E.A. (1915): Additamentum ad lichenographiam Antillarum illustrandam. –
Annales Academiae Scientiarum Fennicae. Serie A. 6, 7: 1–226.
ZAHLBRUCKNER, A. (1924–1925): Catalogus lichenum universalis. Band III. – Gebrüder
Borntraeger, Leipzig.
ZAHLBRUCKNER, A. (1926–1927): Catalogus lichenum universalis. Band IV. – Gebrüder
Borntraeger, Leipzig.
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Colour Plate 14. AF, Thalli of Phyllopsora species from the West Indies (all
scalebars = 1 mm). A, Phyllopsora byssiseda (Imshaug 3157 & Wetmore, MSC).
B, P. cognata (Wright, Lich. Cub. 218; BM lectotype). C, P. confusa (Cummings 44,
NY). D, P. glaucella (Imshaug 25064, MSC). E, P. hispaniolae (Harris 20672,
NY holotype). F, P. imshaugii (Imshaug 13037, MSC holotype).
Colour Plate 15. AF, Thalli of Phyllopsora species from the West Indies (all scalebars
= 1 mm). A, Phyllopsora intermediella (Wright s.n., H–NYL 20558 holotype).
B, P. longiuscula (Timdal 10730 & Rui s.n., O). C, P. microphyllina (Wright, Lichenes
Cubensis 211, BM). D, P. microsperma (Wright, Lichenes Cubensis Ser. 2, 142, H–
NYL 20518, holotype of Lecidea glabella Nyl.). E, P. minor (Elliott 261, TUR–V
22612[a], holotype). F, P. parvifolia (Timdal 10867 & Rui, O).
Colour Plate 16. AF, Thalli of Phyllopsora species from the West Indies (all
scalebars = 1 mm). A, Phyllopsora parvifoliella (Sipman 14852, B). B, P. pertexta
(Wright s.n., H–NYL 17344 holotype). C, P. phaeobyssina (Duss 481, TUR–
V 22602 holotype). D, P. pyrrhomelaena (Wright, Lichenes Cubensis Ser. 2, 124, H–
NYL 20498). E, P. teretiuscula (Tønsberg 37814, BG holotype). F, P. tobagensis
(Timdal 10764 & Rui, O holotype).
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