Key - Phyllopsora Neotropical

Organization for Flora Neotropica
Phyllopsora (Bacidiaceae)
Author(s): Lois Brako
Source: Flora Neotropica, Vol. 55, Phyllopsora (Bacidiaceae) (Mar. 5, 1991), pp. 1-66
Published by: New York Botanical Garden Press on behalf of Organization for Flora Neotropica
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FLORA NEOTROPIC
MONOGRAPH 55
PHYLLOPSORA (BACIDIACEAE)
by
Lois Brako
C* \ -'? TROPI'C Of CANCER
FLORA:
NEOTROPICAi
IOPIC OF CAPRICOIN
Publishedfor
Organization for Flora Neotropica
by
The New York Botanical Garden
New York
Issued 5 March 1991
OANICgAL
mX
ULLRS O[NCO

Copyright ? 1991
Published by
Bronx, New York 10458
International Standard Serial Number 0071-5794
Library of Congress Cataloging in Publication Data

Flora neotropica. - Monograph no. 1 - New York: Published
for Organization for Flora Neotropica by the New York
Botanical Garden, 1968-
v.: ill.; 26 cm.
Irregular.
Each issue has distinctive title.
Separatelycatalogedand classifiedin LC before monographno. 40.
ISSN 0071-5794 = Flora neotropica.
1. Botany-Latin America-Classification-Collected works. 2. Botany-
Tropics-Classification-Collected works. 3. Botany-Classification-Col-
lected works. I. Organizationfor Flora Neotropica. II. New York Botanical
Garden.
QK205.F58 581.98'012-dcl9 85-647083
AACR 2 MARC-S
Libraryof Congress [8508]
ISBN 0-89327-364-3

PHYLLOPSORA (BACIDIACEAE)
LOISBRAKO1
TABLE OF CONTENTS
Abstract .. . .... .... .. ... ..... .. ... .... .. .... ...... .. ... ....... .. .... ...... . . .. .. .. .... . 2
Resumen ......................................... ....................................... 2
Introduction ........................................ ....................................... 2
HistoricalOutline ........................................................................ 3
Materialsand Methods ........................................................................ 4
Morphologyand Anatomy ..................................................................... 7
Chemistry ...................................................................... ...... 15
Distributionand Ecology ...................................................................... 19
PhylogeneticRelationshipsand SystematicPosition of Phyllospora................................. 21
Delimitation of Species and InfraspecificTaxa ................................... ................ 23
Taxonomic Treatment ........................................................................ 26
Key ..........2................ ......................................................26
Descriptionof the Genus .................................................................. 28
Descriptionsof the NeotropicalSpecies ...................................... ............... 29
1. Phyllopsorabibula(Taylor)Swinscow& Krog ................... .................... 29
2. P. buettneri(MiillerArgoviensis)Zahlbruckner....................................... 29
a. var. buettneri.................................................................. 30
b. var. glauca (Bouly de Lesdain)Brako ............................................ 30
c. var. munda(Malme)Brako ..................................................... 33
3. P. canoumbrina(Vainio) Brako .................................................... 33
4. P. chlorophaea(MiillerArgoviensis)Zahlbruckner.................................... 34
5. P. confusaSwinscow& Krog....................................................... 35
6. P. corallina(Eschweiler)MiillerArgoviensis ................... ...................... 37
a. var. corallina .................................................................. 38
b. var. glaucella (Vainio) Brako ................................................... 39
c. var. ochroxantha(Nylander)Brako .............................................. 39
d. var. phaeobyssina(Vainio) Brako ................................................ 42
e. var. rappianaBrako......................... ................... ..... ....... 42
f. var. santensis(Tuckerman) Brako................................................ 43
7. P. cuyabensis(Malme)Zahlbruckner................................................ 44
8. P. fendleri (Tuckerman& Montagne)MiillerArgoviensis.............................. 44
9. P. furfuracea(Persoon)Zahlbruckner............................................... 46
10. P. glabella (Nylander)G. Schneider................................................. 48
11. P. intermediella(Nylander)Zahlbruckner............................................ 49
12. P. isidiotyla(Vainio) Riddle ....................................................... 50
13. P. kalbii Brako ................................................................... 51
14. P. longiuscula(Nylander)Zahlbruckner............................................. 51
15. P. minor Brako.................................................................. 52
16. P. parvifolia(Persoon)MiillerArgoviensis........................................... 52
a. var. parvifolia ................................................................. 53
b. var. breviuscula(Nylander)Brako................................................ 56
17. P. parvifoliella(Nylander)MiillerArgoviensis........................................ 56
18. P. subcrustacea(Malme)Brako..................................................... 57
ConfusedNames, Nomina Dubia, and ExcludedSpecies ................... ................... 57
Types Not Seen ................ ................... ............................ 60
Acknowledgments ............................................................................ 60
LiteratureCited ............................................................................ 60
NumericalList of Taxa ............................................................... ........ 61
List of Exsiccatae............................................................................ 62
Index to ScientificNames ........................................ ........................... 65
Presentaddress:MissouriBotanicalGarden,P.O. Box 299, St. Louis, Missouri63166, U.S.A.

1

2 Flora Neotropica
ABSTRACT
Brako, L. (The New York Botanical Garden, Bronx, New York, 10458). Phyllopsora
(Bacidiaceae).Flora Neotropica55: 1-67. 1991.
A taxonomic monographis presented for the neotropical species of the lichen genus
PhyllopsoraMiill. Arg. (Bacidiaceae).The genus is distinguishedby a combinationof char-
acters:a squamuloseto subfoliose thallus with an obvious prothallus;an ascal type with
an amyloid hemisphericdome and narrow,conical masse axiale;an apothecialtype com-
posed of highly gelatinizedhyphae with no clear distinction between the exciple and the
hypotheciumand with the same texturefound in the centerand marginof the apothecium
as well as in the paraphyses;and by small, thin-walled, rarely septate ascospores. The
relationshipof Phyllopsorawith the generaBacidia, Bacidiopsora,Biatora,Eschatogonia,
Physcidia,and Squamacidiais discussed.Type specimenshave been studied for nearlyall
names includedin Phyllopsora.Eighteenspecies and eleven varietiesare recognizedin the
neotropics, including one species (P. kalbii) and one variety (P. corallina var. rappiana)
describedas new.
The speciesconceptin Phyllopsorais reassessed.Most specieswerefound to exhibitgreat
morphologicalvariation.New chemicaldataaregiven for a numberof species,and chemical
strains are recognizedfor P. buettneriand P. corallina and their varieties. Eight of the
eighteenspecieshave a pantropicaldistributionand most of the remainingspeciesarewidely
distributedin the New World.
RESUMEN
Brako,L. (TheNew YorkBotanicalGarden,Bronx,New York 10458,U.S.A.) Phyllopsora
(Bacidiaceae).Flora Neotropica 55: 1-67. 1991. Se presentauna monografiataxon6mica
de las especies neotropicalesdel genero de liquenes PhyllosporaMull. Arg. (Bacidiaceae).
El genero se distingue por una combinaci6n de caracteres:un talo escuamuloso con un
protaloevidente;un tipo de ascus con un domo hemisfericoamiloide y un "masse axiale"
angosto y c6nico; un tipo de apotecio compuesto de hifas altamentegelatinizadassin una
distinci6nclaraentreel paratecioy el hipotecio y con la misma texturahalladaen el centro
y margendel apotecio como en las parafisis;y por asc6sporaspequefias,de pareddelgada,
y raras veces septadas. Se discute el parentescode Phyllopsoracon los generos Bacidia,
Bacidiopsora,Biatora,Eschatagonia,Physcidia,y Squamacidia.Especimenestipo se estu-
diaron paracasi todos los nombresincluidos dentro de Phyllospsora.Dieciocho especies y
once variedadesson reconocidasen el area neotropical,incluyendouna especie (P. kalbii)
y una variedad nueva (P. corallina var. rappiana).Tambien se re-evalu6 el concepto de
especie en Phyllopsora,encontrandosemucha variaci6nmorfol6gicaen la mayoria de las
especies. Se dan nuevos datos quimicos para ciertas especies, y razas qimicas fueron re-
conocidas en P. buettneriy P. corallinay sus variedades.Ocho de las dieciocho especies
tienen una distribuci6npantropical,y la mayoriade las especies restantesson distribuidas
ampliamenteen el Nuevo Mundo.
INTRODUCTION spread in tropical and subtropicalareas of the

world, with one extendinginto westernEurope.
This study provides a systematictreatmentof All have an obvious prothallus,and thallusform
the neotropicalspecies of the lichen genus Phyl- varies from squamulose to subfoliose or gran-
lopsora.It is also one of the firstapplicationsof ular-crustose.Most species are corticolous, and
modem taxonomic concepts to the systematic some also grow on rock or on bryophytes.The
treatmentof a predominantlytropical group of genus rangesaltitudinallyfrom sea level to 3500
microlichens. m. Althoughover 90 names have been published
Species of Phyllopsoraare common and wide- in the genus, with 51 species describedfrom the

Introduction3
neotropics, this is the first monographof Phyl- rella with tranversely septate, hyaline spores. Four
lopsora.The only modem referenceto the genus species (one considered doubtful) and one vari-
is the regionaltreatmentof the eleven East Af- ety were assigned to Phyllopsora. Zahlbruckner
ricanspeciesby Swinscowand Krog(1981). They (1903-1908) presented Miller Argoviensis's tribe
presented additional notes on the non-African as the family Phyllopsoraceae. Between Zahl-
material they examined, including many type bruckner (1926-1927) and Lamb (1963) 40 spe-
specimens. cies and numerous varieties were described to
As definedby Brako(1987, 1989), Phyllopsora Phyllopsora. Phyllopsora breviuscula (Nyl.) Mill.
is placed in the Bacidiaceaeand is distinguished Arg. was selected as the lectotype of the genus
from other generain that family by the combi- by Clements and Shear (1931). Poelt (1973)
nation of a squamuloseto subfoliosethalluswith treated Phyllopsora as a member of the Lecidea-
an obvious prothallus;an ascal type with an am- ceae s.l. Riedl (1973) added a new species from
yloid hemispheric dome and narrow, conical Surinam, and Coppins and James (1979) de-
masse axiale, the masse axiale not surrounded scribed the first species of the genus from Europe.
by a darkenedarea;an apothecialtype composed In a revision of the genus Psora Schneider (1979)
of highly gelatinized hyphae with no clear dis- made 18 new combinations in Phyllopsora and
tinction between the exciple and the hypothe- placed it in the Cladoniaceae. For their revision
cium, with the same texturefound in the center of East African Phyllopsora Swinscow and Krog
and marginof the apotheciumas in the paraph- (1981), added three new species and made seven
yses and by small, thin-walled,rarelyseptateas- new combinations. After Ferraro's (1983) de-
cospores. scription of a new species from Argentina, and
Species delimitations have been reassessedin Swinscow and Krog's (1985) description of an-
Phyllopsoraafter examining the type specimens other new species from Africa, the total of pub-
of nearly all taxa described in the genus. Re- lished names in Phyllopsora stood at 93.
evaluationof their taxonomy has been made us- In his reassessment of the formerly broadly
ing comparativeanatomy,analysisof lichen sub- circumscribed families Lecanoraceae and Lecid-
stances, and by ecological observations. Most eaceae, Hafellner (1984) segregated Phyllopsora
Phyllopsora species were previously distin- and Physcidia in the Phyllopsoraceae, and sug-
guished by thalline characterswithout any idea gested a possible relationship with Hypoceno-
of the extentof environmentalvariabilityin these myce. Psorella, on the other hand, was placed in
traits. Extensive field studies were conductedto the Bacidiaceae. This was emended in Eriksson
determine variability, and most species were and Hawksworth (1986) where Phyllopsora is
found to exhibit great morphologicalvariation. placed in the Bacidiaceae. Finally, Brako (1989)
Taxa are separatedat the species level on the presented a series of taxonomic and nomencla-
basis of two or more independent characters, tural changes resulting from her revision of Phyl-
while the rank of variety is used for taxa which lopsora.
differ by a single characterand which are not The best previous descriptions of Phyllopsora
fully allopatric.As a result of this study, the 51 were by Swinscow and Krog (1981) and Hafell-
previously described taxa from the neotropics ner (1984) and included: 1) squamulose thallus
have been reducedto eighteenspeciesand eleven with an obvious prothallus, 2) ascal apex com-
varieties. posed of a hyaline hemispheric dome staining
blue with iodine surrounding a narrow, conical,
Historical Outline lighter staining masse axiale, 3) paraphyses with
colorless unswollen apices, and 4) biatorine apo-
Miller Argoviensis(1894) describedthe tribe thecia with a hypothecium composed of highly
Phyllopsoraein his survey of New Zealand li- gelatinized hyphae. Genera such as Bacidia, Bia-
chens, which he includedin his "SeriesII Tham- tora, Physcidia, Psorella and Eschatogonia, how-
no-Phylloblastae." It was differentiated by a ever, have some or all members that share many
loosely or moderatelyfixed squamulosethallus, of these characters with Phyllopsora and are thus
"gonidia palmellaceae"and biatorine-lecideine thought to be closely related to it. These genera
apothecia. Two genera comprised this tribe, have been separated mostly by vegetative char-
Phyllopsorawith simple hyalinespores,and Pso- acters, such as thallus type and its asexual propa-

4 Flora Neotropica
gules. For vegetative charactersto be meaning- of duplicatesare not indicatedin the specimens
ful, phenotypic variabilityand ecology must be cited.
understood, especially with consideration of Field studiesweremade in Brazil(1983, 1987),
thallus age and exposure. But except for tem- Ecuador(1981-1983, 1988), Fiji (1981), Florida,
peratetaxa, most membersof these generahave U.S.A. (1985), Panama(1985), Venezuela(1984-
not been examined since their original descrip- 1988), New Zealand (1988), Tahiti (1988), Co-
tions. They are poorly representedin herbaria, lombia, Peru and Chile (1988). Most of the ma-
often by incomplete or badly mixed collections. terial collected duringthese field trips is depos-
Field studies conductedduringthis study made ited at NY, withunicatesandduplicatesdeposited
possible a critical interpretationof thallus mor- in host countries.
phologyand providedsufficientmaterialfor lab- For citation of type specimenssquarebrackets
oratorystudies. Over 500 collectionsof Phyllop- have been used to indicate information which
sora and related genera were made to better was not in the protologue or, which was only
delimit Phyllopsoraand for furtherassesmentof found on isotypes or syntypes.For all specimens
intergenericrelationships. parenthesesare used to correctspellings,partic-
Field studieswere also importantfor detecting ularlyof place names. All informationhas been
intraspecificvariation. Many of the 93 taxa de- translatedinto English,exceptpropernames,and
scribedin Phyllopsorawere only minor variants dates and numbers have been converted to a
of other species, while several species were im- standardform. To indicateduplicatespecimens,
properlyplaced in this genus. A more coherent an equals sign has been used afterthe herbarium
species concept has been employed here, requir- acronym,followed by the number of duplicates
ing two independentcharactersfor species level and an "x" (e.g., L = 2 x indicates there are 2
separation,to reevaluatethe status of all names duplicatesat Leiden).
and to excludetaxa which clearlybelong outside Sectionsof thalli and apotheciawere made by
Phyllopsora. hand with a razor-blade.A freezingmicrotome
was used to cut 10-20 um thick sections of type
materialand for observation of detailed anato-
MATERIALSAND METHODS my. Asci, ascospores and conidia were studied
in squashpreparations.Materialwas firstmount-
Over 3000 collectionswere examined, included in water, followed by applicationof 10%po-
ing materialon loan from the followingherbaria: tassium hydroxide (KOH). The iodine reaction
AAU, AK, B, BG, BM, BUF, CAS,CHR,COLO, of the asci was studiedin a solution of potassium
CTES, F, FH, FLAS, FLOR, G, GB, GLAM, iodide (IKI) and in modified Lugol's solution
GOET, GZU, H, INPA, L, LD, LG, LSU, M, (MLS: 1 g iodine, 2 g potassium iodide in 300
MBM, MERF, MICH, MIN, NY, 0, PC, QCA, ml of 100%lactic acid). The iodine reactionwas
RB, S, TO, TUR, U, UPS, US, VEN, W, WELT, studied with and without pretreatmentwith po-
ZT. Abbreviations follow Index Herbariorum tassium hydroxide.
(Holmgrenet al., 1981). Materialsfrom the pri- Ascospores, conidia, and paraphyses were
vate herbariaof K. Kalb, Neumarkt,West Ger- measuredwith an accuracyof 0.5 /m. The mean
many, (Hb. Kalb), A. Aptroot, Utrecht, The value (X) and the standarddeviation (SD) were
Netherlands, (Hb. Aptroot), and H. Osorio, calculated(Table I) for comparisonof measure-
Montevideo, Uruguay (Hb. Osorio), were also ments of ascospores.The mean value is given as
examined. For the Kalb collections that lack the middle number of the ascospore size range
unique collecting numbers, most of the one- to in the descriptions.
three-digitnumbersused for specimenscited re- For detectionof lichen secondarycompounds,
fer to locality codes as indicatedon cardsinside thin-layerchromatography (TLC)was carriedout
the specimen packets. The same Kalb number, in accordancewith Menlove's (1974) modifica-
therefore,may be used for severaltaxa and "p.p." tion of standardproceduresas describedby Cul-
is used to indicate portions sharing the same berson and Kristinsson (1970) and Culberson
number. Many of the Kalb specimens studied (1972) for most specimenscited. The thalluswas
had duplicates,but as these were not distributed spot-testedwith paraphenylenediamine (Pd).Elix
at the time of this study,the numberand location and co-workersat the AustralianNational Uni-

Materials and Methods 5
Table I
Ascospore lengths (left) and widths (right)given in um, showing the standarddeviation (SD) from
the mean value (X)
Ascospore Length Taxon Ascospore Width

4 5 6 7 8 9 10 11 12 13 14 15 1 2 3 4 5
P. parvifoliella
P. glahella
~P. coralina var.phaohbyssina "

-4--*_~~~~
*? ,-P. __ canoumbrina H
_^*^-^-I~~~~~ _ ~P. intermediella

P. corallinavar.rappiana H
P. cuyabensis
P. parvifoliavar.hreviuscula
-' -^ /*~~~P.
corallinavar.ochroxantha
-iîM^
P. corallinavar.glaucella
-*w4iii.i--~ IP. kalbii
_ _ P. fufurracea
P. buectnerivar.glauca
P. corallinavar.corallina
P. minor --
, ! ,, P. chlorophaea
P. isidiotyla
P. confusa -t4
,,I
^^- ,, P. corallinavar.santensi
B---~~I ~ P. bibula
P. parvifoliavar.parvifolia ,
_--*^*- ~ P. buettnerivar.munda
'I~~
,*-,,- i ~ P. fenderi
__*?ti"?M" P. subcrustacea
P. longiuscula
16 17 18
versity(pers.comm.) providedadditionalchem- tures were made on the day of collection and to

icalinformationbasedon highperformanceliquid two weeksaftercollection.Culturesand air-dried
chromatography(HPLC) and mass spectrosco- material were kept at room temperatureor re-
py. frigeratedat 17?C.Individual cells or groups of
The proceduresoutlinedby Ahmadjian(1967, algal cells were extractedfrom intact squamules
1973) were followedin an attemptto cultureand and placed on Bold's MineralMedium and -Tre-
identifythe phycobiontof Phyllopsoraand some bouxia OrganicNutrient Media I after a series
relatedtaxa, and for study of growthand lichen- of cleansing transfers.Only a few attempts to
ization. Selected material was initially isolated culturethe algal cells were successful,however.
and culturedon artificialmedia in Caracas,Ven- These cultureswere maintained for a period of
ezuela, in 1985, wherefreshmaterialwas readily severalmonths with weeklytransferto freshme-
available. The photobiont was isolated and cul- dia to avoid bacterialcontaminants.Fresh ma-

k I V 46
1hk ~ Es
~ ~ V ;
ir
,~ ~ ~., , t, '
?~~~~~~~~~~~~~~~~1 ~ .. .o
~~~~
-*91-i --Op~~~~~~~~~
4'
'C1~
-_%,~~~~~~~~~~~~~l~
. 4
4- -? 1 ..
I .... ;'.' ,',%? .~ " 3~~~~~~~~~~

& 4? * F ?~~
Z, o
'-.
,~.,~... .. .~YZ
canoumb, , ( T'H :
.P
Tc. .:,
o c o (L p
]FG ] hylpsr bbl (~vtyc ]ua Fenade Isad,[Bneo14] H1:x.bPhlp
caoubin,
(Isolectotype: B Hloyp:TriiaT
W hatr1,FHsae=Im.
,1 ? c. m mPhlosra choohe Loye
Brail Pigar
121G)
12x.d.Phllpsracofua Kogan SwnsowK
(oltye:Keya 8/77 O 1x
e.Phllpor cralnava.coalin
BrziKab 59(b.Kab) 2 . .Phllpsr
cralnava.croanh
(Isolectotype:Bolivia,Weddell s~~~~~~~~~~n.,
PC),
sc a l e = 1 m m .~I rl
(Iolcttpe Blii, Wedl .n.,PC,scl=Im

, .
Phipsr
FI. corallonasvr.crinabul
(Brzile Kuaib25 (Hb.nde eo 12 x.
68] H
KsandiB). fPhlosrcralnvab.ochroxantha

Morphologyand Anatomy 7
terial was sent to Dr. V. Ahmadjian who also dividual squamuleswhich grow in a fan-shaped
made isolations and cultures.The photobiont is pattern,elongatingand dividing to take the form
discussed on page 10. shown in Figure 4a. These are considered sub-
Single ascospores, conidia and hyphae were foliose as they may reach a centimeteracross.
placed on 2%wateragarand Malt Yeast Extract Young, fresh thalli are bright green, bluish-
Medium and watched for germination and grey, yellow-brown, or reddish-brown.Thallus
growth. Experimentsto isolate the mycobiont color varies with age, condition of the thallus
were unsuccessful. and ecologicalconditionsand is thereforea poor
taxonomic character.
The upper surface of the thallus is generally
MORPHOLOGY AND ANATOMY glabrousand shiny. However, Phyllopsoracuy-
abensishas a rough,dull uppersurface;the con-
The thallus of Phyllopsoravaries from very vex squamulesof P. bibulaare fibrillose;and the
small patches of a few centimetersin diameter upper surface of the thallus of P. buettneriis
to extensivecolonies completelyencirclingwhole commonly pruinose (the whitish pruina often
tree trunks.Maturethalli are usually circularor gives the yellow-brown colored squamules a
oval in outline. Thallus form is squamulose,or pinkishcolorin olderherbariumspecimens).The
rarelysubfolioseor granular-crustose.Most spe- marginof the squamulesis usuallypubescentor
cies have a thallus composed of discrete to ad- fibrillose,or rarelyglabrous.The marginalfibrils
joined and overlapping squamules. The squa- are usuallyfine and pale, but in P. parvifoliavar.
mules are round to irregularand measure from breviuscula(Fig. 4c) they are coarseand reddish.
0.1 to 1.0 mm in diameter or lobe width; they A prothallus composed of superficial, unli-
may also be elongate,lobulate, or digitate.They chenized hyphae is always present in Phyllop-
are often bifurcateat the apex owing to terminal sora. It is either web-like, consistingof fine pale
formationof new lobes. The squamulesareeither hyphae between the thalline elements, or well
adnate to the prothallusor substrate,or ascend- developed, forminga distinct fringe around the
ing. Smallglobose squamulesaregenerallyclose- peripheryof the thallus.The prothallusmay also
ly adnate,while the largeror longer-lobedforms form an extensive mat of thick-walled, highly
are more loosely adnateand are often ascending. branched and interwoven hyphae located be-
All squamulesareconvex or complanate.In most tween, under and/or around the thalline ele-
species the squamules adjoin and overlap. In ments. The thallineelementshave a layerof pale
Phyllopsoracamoumbrinathe squamulesare so or reddish-brownloosely woven hyphae on the
closely adjoined that they form a nearlycontin- lower surfacewhich either interweaveswith the
uous crust (Fig. Ib), while in P. parvifoliellathe hyphaeof the prothallusor attachesand anchors
squamulesare mostly discrete (Fig. 3e). directly to the substrate. Prothallus color is a
The squamules may be placed in three size poor taxonomiccharactersince it varieswith age
categories:ca. 0.1 mm wide, 0.1-0.3 mm wide and light exposure.
and (0.3-)0.5-1.0 mm wide. Measurementswere Isidia are common in the genus. The term is-
made behind the apex, just behind the bifurca- idium is used here for all non-dorsiventral,fin-
tion point or across the diameter of unlobed ger-likeprojectionsthat have a continuous cor-
squamulesat their widest point. In the smallest tex, formed either as appendages from the
size categorythe squamulesare often mere gran- squamules or independentlyon the prothallus.
ules which are globose or complanate, elongate They aregloboseto cylindricalor coralloid.Isidia
or digitate.Examplesof this groupincludePhyl- usually arise from the lamina or margin of the
lopsorafurfuracea(Fig. 2), with globose granular squamules,but they are also found to occur in-
squamules;P. minor (Fig. 3b) with complanate dependentlyon the prothallus,as in Phyllopsora
squamules;and P. confusa (Fig. Id) with small furfuracea(Fig. 2c, d). Phyllopsoracorallinavar.
digitate squamules. Within this size group it is phaeobyssinahas unusual flattened isidia (Fig.
particularlydifficultto distinguishdigitatesqua- 5c, d). Eleven of the 18 neotropical species of
mules from isidia. Phyllopsoraare isidiate, but the presenceor ab-
The species that produce a subfoliose type of sence of isidia is an unreliablecharacterin most
thallus usually begin their development as in- species. Field observations have shown that a

8 Flora Neotropica
A. B.
mm.
C. D.
FIG.2. Phyllopsorafurfuracea (DominicanRepublic,Harris19751, NY). Linedrawing.a & b. Youngapothecia

and squamules,side view (upper)and top view (lower).c. Apothecia and developing isidia, top view (upper)
and side view (lower).d. Elongatedand deflatedisidia, top view (upper)and side view (lower).See page 48 for
discussionof developmentalsequence.
singlecolony of P. corallinavar. santensisvaries although soredia-like granules have been ob-

from non-isidiateto densely isidiate on different served on a few thalli.
portionsof the same thallus. A distinct upper cortex is always present. As
Lobulesare common in the genus and are ex- observed in cross-section, it is generally com-
tremelyvariablein shape.Both isidia and lobules posed of a colorless or pale layer of randomly
arepotentialdiaspores,since they are easily bro- oriented or anticlinally arranged,hyphae. Two
ken off and are found on neighboringmosses, extremetypes of cortex tissue can be recognized,
otherlichensand baresubstrate.Sorediaformed designatedtype 1 and type 2 by Swinscow and
in distinct soralia are not known in the genus, Krog (1981). Type 1 (Fig. 6a, 7a) has heavily

Morphologyand Anatomy 9
w
~
r~~~~~~~1.; ~ ~ ~ ~ ~ ~
.B, .aOT =
r~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~e,.
', ,.~
-. ?i" '., .,-L
: ~~'--)~ rr,: :i-

'i aC #'r'" '~ ; D
e__ ..X <
FIG.3. a. Phyllopsorakalbii(Holotype:Brazil,Kalb258, NY), scale= 1 mm. b. Phyllopsoraminor(Lectotype:

St. Vincent, Elliott 261, TUR-VAIN 22612A), scale = 1 mm. c. Phyllopsora parvifolia var. parvifolia (Brazil,
Malme s.n., Lich. Regnell. 1227, S), 12 x d. Phyllopsora parvifolia var. breviuscula (Isotype: Cuba, Wright s.n.,
Lich. Cub. 181, UPS), scale = 1 mm. e. Phyllopsora parvifoliella (Lectotype: Wright s.n., Lich. Cub. 182, BM),
scale = 1 mm. f. Phyllopsora subcrustacea (Holotype: Paraguay, Malme s.n., Lich. Regnell. 1612B, S) 12 x.

10 Flora Neotropica
A'
( 5,
[123~~:. .
1 A r~~~~~~~~~~"'"
"r~
%,~ ~ ~~~~~+~?
'~'~ " L- .
~, ~L'L :
',..~ ~ ~ ~~~~~~~~~~~~~~~?
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~??
:~~'.b'., ~;
FIG 4.a. h
brvucl parifi ylopsr(~aE var (Ioye
Cuba,
Wrigtsn,Lc.Cu.11
P)E
FIG. 4. a. Phylopsora parvifoia var. breviuscula (Isotype: Cuba, Wright s.n., Lich. Cub. 181, UPS). SEM
micrographshowingfan-shapedsquamulesand pycnidium(arrow),40 x. b. Phyllopsoracorallinavar. santensis
(USA, Florida, Brako 8227, NY). SEM micrographshowing smooth surfaceof the apothecia and cylindrical
isidia, 100x. c. Phyllopsoraparvifoliavar. breviuscula(Isotype:Cuba, Wrights.n., Lich. Cub. 181, UPS). SEM
micrographof uppersurfaceof squamuleswith coarseprojectingfibrils,150 x. d. Eschatogoniaprolifera(Guyana,
Sipman & Aptroot 18812, B). SEM micrograph of cortex, 3000 x.
gelatinizedhyphaewith narrow,net-likelumina. PseudochlorellaLund(Ahmadjian,pers.comm.).

In type 2 (Fig. 6b, 7b), the lumina are much The algalcells are often penetratedby numerous
broaderand rarelyappearnet-like. Intermediate fungalhaustoria.A high frequencyof haustorial
typesarecommon, however.In specieswith very penetration has been found to be common in
small thalline elements the cortex may appearas structurallyprimitive lichens (Plessl, 1963).
a thin gelatinouslayer (Fig. 6c). The cortex also Hyphae from the cortex often extend through
varies with developmental stage and ecological the algal layer to the medulla, a layer of loosely
conditions;it often contains small granules,per- woven pale or reddish-brownhyphae that are
haps atranorincrystals,which dissolve in KOH. sometimes covered with granulesof lichen sub-
The photobiont occurs in a more or less con- stances. The lower surfaceconsists of randomly
tinuous layer below the upper cortex. The algae oriented, loosely woven pale or reddish-brown
are unicellularand green, with cells 10-15(-20) hyphae that may interweavewith the hyphae of
Am in diam. (Fig. 7c), and belong to the genus the prothallus. The surface of these hyphae is

Morphology and Anatomy 11
*****P^^
v^lrss:
C2B 1i
HflJr'
KwikKA
^^
^L^4 ^!a u.9
sB^^tî^.
^^^WSs^^-' I^W
i^^L^ '^^^J(L:
^^v-UsL-3^s1 "<<fc^<
~.d'- ? .. ..
* N
'e'r
FIG. 5. Phyllopsoracorallinavar. santensis(USA, Florida,Brako 8227, NY). a. SEM micrographshowing

smooth surfaceof the apothecia and globose to cylindricalisidia, 50 x. b. Habit, 12 x. Phyllopsoracorallina
var.phaeobyssina(Holotype:Guadeloupe,Duss 481, TUR-VAIN 22602). c. SEM micrograph,isidia, 50 x. d.
SEMmicrograph,
SEM micrograph,cross-section flattenedisidium,
of flattened
cross-sectionof isidium, 2000 x.
2000 x.
sometimes papillose (Fig. 6d), which might be distinctupperand lowercortex(Fig. 4d). Besides
associatedwith water or nutrientabsorption. the Bacidiaceae,members of the Pannariaceae,
The characteristicsquamulose thallus form Pilocarpaceae,Lecanoraceae,Psoraceae,and Le-
withan obvious prothallusas seen in Phyllopsora cideaceaes. lat., also have a similarthallus mor-
is also foundin othermembersof the Bacidiaceae phology. Most species in the Pannariaceaeare
andeven in membersof otherfamilies.The ques- squamulosewith a thick prothallus,but all have
tion naturallyarises whether this phenomenon Nostocas a photobiont(Jorgensen,1978), which
is due to true relationshipor parallelevolution. gives the squamulesa distinctivegraycoloration.
Within the Bacidiaceae,several tropical species The squamules in some members of the Pilo-
with the fundamentalcharacteristicsof Bacidia carpaceaeand Lecanoraceaeare very similar to
and some taxapreviouslyplacedin Psorellahave those of Phyllopsora,and the taxa can only be
a thallus morphologyidentical to that of Phyl- separated by apothecial characters. Primary
lopsora.PhyscidiasquamulosaTuck., with leca- squamulesof some speciesofCladonia (Cladoni-
norine apothecia, has a similar thallus mor- aceae)also resemblePhyllopsorasquamules,but
phology to Phyllopsora,and Eschatogonia has they have a differentcortex and usually distinc-
squamuleswhich are similar in size, but have a tive chemistry.

12 Flora Neotropica
i"'"
FIG. 6. a. Cross-sectionsof thallus-Type 1 cortex. Phyllopsoraparvifoliavar. breviuscula(Isotype:Cuba,

Wrights.n., Lich. Cub. 181, UPS). SEM micrograph,450x. b. Cross-sectionsofthallus-Type 2 cortex. Phyl-
lopsoraparvifoliavar. parvifolia(Brazil,Malme s.n., Lich. Regnell. 1227, S). SEM micrograph,450 x. c. Cross-
sections of thallus-thin gelatinous cortex. Phyllopsoraintermediella(Isotype:Cuba, Wrights.n., Lich. Cub.
183, BM). SEMmicrograph,900 x. d. Hyphaeof lower surfaces.Phyllopsoraparvifoliavar. breviuscula(Isotype:
Cuba, Wright sn., Lich. Cub. 181, UPS). SEM micrograph, 1500 x.
Crocyniapyxinoides Nyl. (Crocyniaceae)has sora, but they lack a distinct prothallus.Timdal

a squamulose-crustose thallus form with an (1984a) described three types of squamules in
abundantprothallus,but it differsfrom Phyllop- Hypocenomyce:adnate,bullateor ascending.He
sora in its lack of an upper cortex and green- recognizedseveral species groupswithin the ge-
black pigmentation in the prothallus and apo- nus where anatomical and chemical characters
thecia. are constant within each group, but the thallus
The squamulesof Psora (Psoraceae)are some- form varies. Timdal (1984a) suggestedthat the
what largerand have a thicker cortex (80-200 morphological similarities in squamule types
,m; Timdal, 1984b) than those of Phyllopsora found in species formerlyplaced in Psora have
(5-60 ,m), and the "Scheinrindentype"of cortex evolved independentlyand are not an indication
found in Psora species is not found in Phyllop- of close geneticaffinityof the speciesgroups.The
sora.Severalspecies ofHypocenomyce(Lecidea- widespreadoccurrenceof the squamulosethallus
ceae s. lat.) have squamules similar to Phyllop- form in the broad array of families mentioned

Morphology and Anatomy 13
' I
FIG.
7.a. C ss-sec~~tioso
r o hlu-Tp otxhlosravflavr rvuclIoyeua
rs-etoso
Wright 8,US.Lgtmcocp,40.b
s..Lc.Cb Bai,Mlesn,Lc.Rgel hlu-Tp otx hl
1osoapavioi a27 ) ih va.irsoe 0prvioi
.cla
c e l l pavflavr
s . ~ avfla(rzl
~ ~ ~ ~~; am.. ic.Rgel
Phlosr 27) ih irsoe00x
d.scs trctreshwig ase xilean rstat dhicecePyloporprvfoiava. arifli (rail
Mames..
Lch
egel.1227 t a
S) outdinMSprete KO,sae=1m.eaahss wi
e
Phyllopsora parv~~~~ifiavtpavfia(rzlMams..LihRenl.12
g
Im.?.
1 0~~~~Pd'* h
FIG. 7. a. Cross-sectionsof thallus-Type 1 cortex. Phyllopsoraparvifoliavar. breviuscula(Isotype:Cuba,

Wright s.n., Lich. Cub. 181, UPS). Light microscope, 400 x.b.bCross-sections of thallus-Type 2 cortex. Phyl-
lopsoraparvifoliavar. parvifolia(Brazil,Malme s.n., Lich. Regnell. 1227, S). Light microscope,400 x. c. Algal
cells.Phyllopsoraparvifoliavar.parvifolia(Brazil,Malmes.n., Lich. Regnell. 1227, S). Lightmicroscope,1000 x.
d. Ascus structureshowing masse axiale and rostratedehiscence,Phyllopsoraparvifoliavar. parvifolia(Brazil,
Malme s.n., Lich. Regnell. 1227, S), mounted in MLS, pretreatedwith KOH, scale = 10 Am.e. Paraphyses.
Phyllopsoraparvifoliavar. parvifolia(Brazil,Malme s.n., Lich. Regnell. 1227, S), mountedin phloxine, scale =
10 lm.
above suggeststhat it has developed repeatedly may also arisedirectlyfromthe hymeniaof older
and independentlyin differentlineages. apothecia.The disc is commonly plane and mar-
Apothecia are common in most species of ginate when young, but usuallybecomes convex
Phyllopsora,even in denselyisidiate speciessuch with age.It is tan,orange,yellow-brown,or brown
as P. isidiotyla.They arebiatorine,but may have to darkred with a slightlypaleror darkermargin.
a dark reddish, yellow-brownor orange colora- The surfaceof the disc is smooth (Fig. 4b). Hy-
tion in the exciple and/or hypothecium(Fig. 8b- phae extendingfreely from the exciple may cre-
d). Apotheciaare usuallyround to slightlyirreg- ate a byssoid fringearound the apothecia.
ular, 1.0 to 1.5 mm diam., sessile, and constrict- The internal organizationof the apothecia is
ed at the base. They are usually anchoredat the relatively uniform. Figure 8 shows the typical
margin with fibrils extending from the exciple tightlycoherent,highlygelatinizedhyphaeof the
and occur singly or aggregated.New apothecia exciple and hypothecium which have the same

14 Flora Neotropica
textureas the paraphyses.The textureof the cen- The hymenium varies in height from 20-60
ter and the marginis the same. The development um.It is generallycolorlessbut may have orange,
ofapothecia was firstdescribedby Swinscowand yellow-brownor brown pigmentation.
Krog (1981), and my own observationsconfirm Paraphysesin Phyllopsora(Fig. 7e) are rela-
theirfindings.Apothecialinitials arefirstnoticed tively broad compared to those of other mem-
as small nodules on the surface of the thallus. bers of the Lecanoraleswith the exception of
Generative hyphae within the nodule can be Crocyniapyxinoides.Wall thicknessvaries from
identifiedby their amyloid reactionin iodine so- 0.2-0.5 um diam. In length the paraphysesvary
lution. The generativehyphaeare surroundedby with hymenial height, the length to width ratio
thalline cortex. As development proceeds, the is 6:1. They are cellular,repeatedlyseptate and
hymenium begins to take shape, expanding at sparselybranchedand anastomosing.The apical
the margin,formingthe disc. The hypothecium cells are at most only slightlythickened,usually
then enlargesand gelatinizes.Swinscowand Krog colorless,but may be coveredwith yellow-brown,
suggest that in some cases the hypothecium is orange or brown pigments. The paraphysesare
derivedmainly from the ingrowthof corticaltis- stronglyconglutinated,and thus not easily sep-
sue and in other cases mainly from the gelati- aratedin squash preparations.
nization of medullarytissue alreadypresent.The The structureof the ascus is uniform in the
hypothecium continues to spread laterallywith genus (Fig. 7d). It is best observed by applying
the outer layers forming the exciple. The exci- 10%KOH to a thin section followed by treat-
pular hyphae may continue to grow outwardto ment with MLS. Young asci are cylindricalwith
become the fibrils which attach to the thalline thick side walls and a thickenedapicaldome that
cortex or form a byssoid fringe.Thereis no clear stainsblue in MLS.The asci areusuallyelongate-
separationbetweenthe hypotheciumand the ex- clavate at maturitywith a deeply stainingapical
ciple, and the hyphae of both the hypothecium dome and distinct non-stainingconical shaped
and exciple and the paraphyseshave the same area(the"masseaxiale"),at the base of the dome.
texture. Between the hymenium and hypothe- The asci commonlyhave a blue-stainingcap over
cium is generativetissue, but it does not form a the exterior of the apex. This ascus type agrees
distinct subhymenium as seen in apothecia of with type "L" of Hafellner (1984). The asco-
Bacidia rosella (Pers.)De Not. spores are liberatedby rostratedehiscence. The
There is some interspecific variation in the asci and paraphysesaregenerallythe same height.
dimensions and number of the hyphae present Ascospores vary in size and shape as shown
within the apotheciaof differentspecies.The hy- in Figure 9a-d and Table I. They are usually
phae of Phyllopsoraparvifolia(Fig. 8a) have rel- simple, rarelyuniseptate,ovoid, ellipsoid or fu-
ativelynarrowluminawhencomparedwith those siform. Ascosporesgerminatewithin or outside
of P. intermediella(Fig. 8b), and the apothecia the ascus.
of P. parvifoliaare usually convex with a thick Pycnidia are common in the genus, but are
hypothecium.The apotheciaof P. intermediella often difficult to find since they mostly appear
are more complanate with fewer hyphae in the as slightly raised areas on the squamule surface
hypothecium.The apothecialtype foundin Phyl- (Fig. 4a) and can be confused with apothecial
lopsorais foundelsewherein Biatoravernalis(L.) initials. They are globose, 0.1-0.2 mm in diam.,
Fr. and generally partly immersed in the thallus,
Pigmentsarecommon in the hypotheciumand rarely superficial.The pycnidial wall is tan or
exciple and they can be seen to vary within a reddish-brown,KOH-, and the ostiolar region
species. In Phyllopsoracorallina var. santensis may be more darklypigmentedthan rest of the
the exciple is irregularlyred pigmented(Fig. 8c). wall. The outer wall is paraplectenchymatous.
The apothecia of P. furfuraceaoften contain an The innersurfaceis lined with irregularlyshaped
orangecrystallinematerialthatdissolves and dif- branchedor unbranchedconidiophores(Fig. 10).
fusespurplish-redin KOH. Apotheciaof P. chlo- Conidiaareproducedapicallyfrom elongate,en-
rophaeaare suffusedwith a darkreddishor pur- teroblasticconidiogenouscells. The conidia are
plish-brown pigment which is non-reactive in rod-shaped,9.0-15.0 x 0.5-1.0 um, straightor
KOH (Fig. 8d). slightlybent.

Chemistry 15
Lig't m 12 .
. '..'' '.' ,
*ip . - ' .
LA
FIG.8. Cross-sectionsofapothecia.a. Phyllopsoraparvifolia

var.parvifolia(Brazil,Malmes.n., Lich.Regnell.
1227, S). Light microscope, 160x. b. Phyllopsoraintermediella(Isotype:Cuba, Wrights.n., Lich. Cub. 183,
BM).Lightmicroscope,160 x. c. Cross-sectionof apotheciumshowingpigmentedexciple.Phyllopsoracorallina
var.santensis(Florida,USA, Brako8227, NY). Lightmicroscope,120 x. d. Cross-sectionof apotheciumshowing
pigmentthroughoutexciple and hypothecium.Phyllopsorachlorophaea(Lectotype:Brazil, Puiggari 1721 G).
Lightmicroscope,120x.
CHEMISTRY were often, but not consistently, detected, in-

cluding ursolic acid, and it is possible they may
Usingthin-layerchromatographic analysis,five have originatedin the bark or from overgrown
of the 18 neotropicalspecies of Phyllopsorahave bryophytesratherthan from the lichen.
readilydetectablelichen substances.The occur- A numberof unknownsubstancesoccurin the
rence of these substancesis shown in Table II. genus, but their structuralanalysis and identifi-
J. Elix (pers. comm.) has begun examination of cation are a subject of continuing study by Elix
the secondary compounds in the genus using and co-workers.The relative positions of some
HPLCand mass spectrometryand has confirmed of these compoundsare shown in Figure 11. Sev-
the presence of the following substances:atra- eralof the unknowncompoundshave been given
norin, pannarin,vicanicin, norvicanicin,argop- names by Elix and co-workers, and are listed
sin, norargopsin,and zeorin. Zeorin is the only below togetherwith names used by Brako(1987)
terpene recognizedby TLC. Other tri-terpenes and Swinscow and Krog (1981).

16 Flora Neotropica
Li
E. FI
V. V
~~oo D.i~~~~ EG.
D.
IODm H.
FIG. 9. Ascospores.a. Phyllopsorachlorophaea,b. Phyllopsoraglabella,c. Phyllopsoraparvifoliavar. brev-
iuscula,d. Phyllopsoralongiuscula,e. Squamacidia,f. Taxa previouslyplaced in Psorella(excludingthe type.)
g. Bacidiopsora,and h. Bacidia rosella.
treatmentthe spot turns bluish-gray.It is Pd+

Compounds Found in Phyllopsora yellow-orange.
Atranorin,Rf. classes 7:7:7, appears as a dark Vicanicin, 'Unknown G1' of Brako (1987), Rf.
spot in short wave UV light. After sulfuricacid classes7:7:7,appearsas a darkspot in shortwave
treatmentthe spot turnsdull yellow-brown.It is UV light. After sulfuric acid treatment it turns
Pd+ yellow. Small yellowish granularcrystalsof brown. It is Pd-.
atranorincan be observed in the cortex on the
outer walls of the hyphae. Norvicanicin, 'Unknown G2' of Brako (1987),
Rf. classes 6-7:6-7:?, appearsas a dark spot in
Pannarin, Rf. classes in
7:7:7, long wave UV light short wave UV light. After sulfuric acid treat-
producesa white fluorescence.Aftersulfuricacid ment it turns brown. It is Pd-.

Chemistry 17
Argopsin(1'chloropannarin),'albicansunknown
2' of Swinscow and Krog, Rf. 6-7:7:7, in long
wave UV lightproducesa yellowishfluorescence.
Aftersulfuricacidtreatmentthe spot turnsbrown.
It is Pd+ orange.
Norargopsin,'Unknown A' of Brako(1987), 'al-
bicans unknown 1' of Swinscow and Krog, Rf.
classes 6:6:6, in long wave UV light producesa
yellowfluorescence.Aftersulfuricacid treatment
the spot is colorless or gray. It is Pd+ yellow.
Unknown B, Rf. classes 6:6:6, in long wave UV
light producesa dull orange fluorescence.After
sulfuric acid treatment the spot is colorless or
faint orange. It always occurs with argopsin, so
the Pd reaction is not known.
Parvifoliin, 'Unknown C' of Brako (1987), Rf. l0pm
IOjum
classes 5-6:6:6, in long wave UV light produces
a faint fluorescence.Aftersulfuricacid treatment cells and conidia.Phyl-
FIG. 10. Conidiogenous
(DominicanRepublic,Harris20481,
lopsorafurfuracea
the spot turns brown. It is Pd-. NY).Linedrawing.
Furfuracein,'Unknown HI' of Brako (1987),
'haemophaeaunknown'of Swinscow and Krog,
Rf. classes 5:5:3,in long wave UV light produces study. The substancefound in the apothecia of
a white fluorescenceAftersulfuricacid treatment Phyllopsorafurfuraceamay be parietin.
There is a long history of controversy con-
the spot turns gray. It is Pd-.
cerningthe use of naturalproduct chemistry in
Phyllopsorin, 'Unknown 01' of Brako (1989), the systematics of lichenized fungi. Recently, a
'ochroxanthaunknown1'of Swinscowand Krog, symposium (W. Culberson, 1986) was devoted
Rf. classes6:5:6, in long wave UV light produces to this subject and addressedthe biogenetic re-
a yellow fluorescence.After sulfuric acid treat- lationships of the lichen substances,correlation
ment it turns bright yellow-orange. It is Pd+ and non-correlationsof chemical variation pat-
orange. ternswith lichenmorphologyand geography,and
biological and ecological considerations of li-
Chlorophyllopsorin,'Unknown 02' of Brako chen substances. In Phyllopsora,consideration
(1989), Rf. classes 7-8:7:8, in long wave UV pro- of chemical substancesas taxonomic characters
duces a yellow fluorescence.After sulfuric acid
is complicatedby the fact that there are so many
treatmentit turns yellow or green. The Pd re-
unknowncompounds.
action is not known.
Upon examiningthe spectrumof lichen com-
Methyl-phyllopsorateand methyl-chlorophyl- pounds found in Phyllopsora,atranorin,the only
lopsorate, 'Unknown 03' of Brako (1987) but para-depsidefound in the genus, is not found to
subsequently found to be two distinct com- be useful as a taxonomic character,as it occurs
pounds,Rf. classes 6-7:6-7, appearas darkspots in the cortex of hundredsof species of lichens in
in shortwave UV light. After sulfuricacid treat- many families. Likewise, zeorin, a triterpenoid
ment they turn brown. They are Pd- or weakly found in threeof the neotropicalspecies of Phyl-
Pd+ yellow. lopsora,is also widespreadin many lichen fam-
ilies, thus giving no clues to relationships.The
Zeorin,Rf. classes 4:4:4, not visible in UV light. most interestingcompounds found in the genus
After sulfuric acid treatment it turns brownish
are the biogeneticallyclosely relateddepsidones
to purplishgray. It is Pd-.
pannarinand argopsin.Pannarinis reportedelse-
Solubleand non-solublepigmentsoccurin the wherefrom species of unrelatedgeneraincluding
thallusand apotheciaand are in need of further Pannaria,Lecanora,Megalospora,Psoromaand

Ao 0? +0 8
)000
o o
1 2 3 4 5 6 7 8
?
o00 3
0?
1 2 3 4 5 6 7 8
o Substances which turn brightyellow after treatment with

dilute sulfuric acid and heat.
* Indicates the darkblue-grayspot characteristic of pannarin.
* Indicates zeorin.
* Implies an orange fluorescence under long wave UV light.
+ Implies an yellow fluorescence under long wave UV light.

FIG. 11. Diagramof chromatograms.

Distribution and Ecology 19
Pseudocyphellaria.Argopsinis reportedfromAr- widely separatedlocalities in the West Indies,

gopsis, Biatora, Erioderma and Micarea. Pan- South America, and southern North America,
narin and argopsinboth occur at Rf. classes 7, but the gaps in the species rangemay well be an
and are distinguishedby the color of their spot artifactof the few collectionsof such uncommon
after sulfuricacid treatment,but it could not be taxa. Two species of limited range are Phyllop-
determined if the substances occur together. sora subcrustacea,known only from Paraguay,
Phyllopsorin,chlorophyllopsorin,methyl-phyl- and P. bibula,which has only been collected on
lopsorate, and methyl-chlorophyllopsorateare the JuanFernandezIslandsoff the Chileancoast.
structurallyrelateddepsidones, but requirefur- There is no clear biogeographicalpattern to
ther study for full elucidation, as does the dep- the chemicalvarietiesdetectedin the genus(Figs.
sidone furfuracein.Vicanicin and norvicanicin, 14, 15 & 16). Although the greatest chemical
also depsidones, are found elsewherein the Te- diversityis foundin the New Worldspecies,these
loschistaceae.Parvifoliinis not a depsidone,but have been muchbettercollectedand studiedthan
Elix and co-workers(pers. comm.) have not yet the Old Worldtaxa. Also, differentchemicalva-
determinedto which class of compoundsit does rieties often occur sympatrically.
belong. Phyllopsoraspecies are generallycorticolous,
In Phyllopsorabuettnerithere are three chem- mostly on the basal portion of the main trunks
ical races (Table II), which seem to follow the of living trees, and two or three species often
replacement pattern of Hawksworth (1976). I occuron the same trunk.Thereseems to be little
have recognizedthese at the level of variety as host specificityfor Phyllopsora;species occuron
they correlatewith only slightdifferencesin mor- a broad range of woody angiosperms. Several
phologyand the distributionpatternsarenot cer- taxa were found to occur on trunksof Araucaria
tain. In P. buettnerivar. glauca and P. corallina angustifoliain southernBrazil, but Phyllopsora
vars. ochroxantha and santensis, I recognize is rarely found on other gymnosperms. Some
chemical strainsbecause the variation seems to species may also growon rock or on bryophytes,
be an additive patternof accessorycompounds yet they arerarelyfound on leaves or dead wood.
which may prove to be closely related,and there The altitudinalrangeof Phyllopsorais sea level
are no correlating differences in morphology, to 3500 m. Phyllopsoraparvifoliahas an unusu-
ecology or distribution.In P. corallinas. lat. thisally broad altitudinal distributionfrom coastal
could also be interpretedas a case of deletion forests to 3200 m. Phyllopsorafendleri, on the
chemistry(Lamb, 1977), with the full potential other hand, is apparentlyrestrictedto montane
chemistry found in P. corallina var. ochroxan- habitats between 1200 and 2800 m. Species of
tha, but P. corallina var. corallina having lost Phyllopsoraare most common in humid, sub-
the potential for productionof medullarycom- tropicalmontane forests between 500 and 2000
pounds. m, as in the family Megalosporaceae(Sipman,
1983). Both groups are also found at lower ele-
vations on islands such as the Galapagos,or in
DISTRIBUTION AND coastalareaswherelocalcloudcondensationleads
ECOLOGY to high humidity. In drier savannasareas,Phyl-
lopsorais restrictedto the gallery forests. Both
Phyllopsora has a largely pantropical distri- Phyllopsoraand the Megalosporaceaeare rarein
bution (Fig. 12), similar to that of Megalospora tropicallowlandrainforests,whichSipman(1983)
(Sipman, 1983) and Coccocarpia (Arvidsson, attributedto high temperatureintolerancein the
1982). A single species, P. rosei, is found well case of Megalosporaceae.Phyllopsoramay also
into westernEurope.Of the 18 species found in be intolerantof very low light intensitiesin high
the neotropics, nine are pantropical,and those canopiedforestswherethe Thelotremataceaeare
that are restrictedto the neotropics almost al- the dominant corticolous epiphytes as seen in
ways are broadly distributed. Phyllopsoracor- centralAmazonia (pers.obser).Phyllopsoraspe-
allina var. corallina occurs north into Illinois, cies prefer partially shaded habitats, and indi-
while P. confusa extends to 39?S in Chile. Ap- viduals that occur in full sunlight often show
parent disjunctions occur in several species in- some bleachingor browningof the thallus.
cludingP. glabella(Fig. 13) and P. isidiotyla(Fig. Manylichens commonly found growingin the
20). Phyllopsorakalbii is currentlyknown from same habitat with Phyllopsorashare a similar

Table II
Lichen substances
yrC _
.'c^
- C. CS ^ ^ ECi-E:3^ ^ -
-
| i | | - |
Taxon / Chemical strain _ i -
'
- t
-
2^ |? C
= -
< < Z C-'
P. bi ula ..................................................
P. buettnerivar.glauca, strain .............
P. huettneri var. glauca, strain II ............. . E

P. buettnerivar.glauca, strain III...........
P. buettneri var munda ......................... ..
P. canoumbrina ...................................
P. chlorophaea............................ .
.
P. confusa..................................
P. corallinavar.corallina........................ C
P. corallinavar.glaucella........................ o * o
P. corallinavar.ochroxantha,strain .... * * * * S
P. corallinavar.ochroxantha,strain I!..
P. corallinavar.ochroxantha,strain III.
!
P. corallinavar.phaeohyssina.................
P. corallinavar.rappiana........................ 0
P. corallinavar.santensis,strain I......... 0
P. corallinavar.santensis,strain I!........ a
P. c ahensis............................................
P. fendleri................................. ...
P. furfuracea................................. 0
P. glahella .......... ..... ............
P. intermediella.....................................
P. isidio la........ ...................................
P. kalhii.................................. ...........
P. longiuscula..................................
.
.
P. minor....... . .........
...................
P. parvifoliavar.parvifolia.....................
P. parvifoliavar.hreviuscula...................
P. parvifoliella....................... ....
P. subcrustacea........ ..............
* present
] sometimespresent

Phylogenetic Relationships 21
.... -
op~~?-VC -,'-o-. .-
Zp^
-~j '-'l
,~ I
_Tr,lopc o_f Cncer
.__.E E 1o.
tror- r- -
FIG. 2. rangeofhyllopsora.on
Worldwide
FIG. 12. Worldwide range of Phyllopsora.
squamulosethallus form with obvious prothal- paraphyses,thick walled, I+ blue asci and squa-
lus. These include members of Eschatogonia, mulose thallus. Following reassessmentand re-
Physcidia, species with the fundamental char- alignmentof the broadlydefinedfamiliesLecide-
acters of Bacidia, as well as some members of aceae and Lecanoraceaeby Hafellner(1984) and
the Pilocarpaceaeand Lecanoraceae.Othergen- subsequentmodificationsin Erikssonand Hawk-
era commonly found growing with Phyllopsora sworth(1986), Phyllopsoraseems well placed in
are Catinaria,Crocynia,Coenogonium,Chiodec- the familyBacidiaceae.The Bacidiaceaeare con-
ton, Megalosporaand Porina. sidereda monophyleticgroupbased on the syn-
All species of Phyllopsorahave the potential apomorphyof asci with an amyloid hemispheric
for sexualreproductionas all formapotheciawith dome containinga narrow,conical non-amyloid
ascosporesat some point in their development, masse axiale. More than twenty generaare cur-
andthereareno strictlysterilespecies.Ascospore rently included in the family. The characters
size rangesfrom 4 to 20 ,m, which is normalfor which best separatethe generaare basic apothe-
wind dispersal(van Zanten & P6cs, 1981: 505). cial form, types of paraphyses,and ascospores.
Thallus fragments can also be transportedby Littleis knownaboutthe ontogenyof these struc-
wind. Asexual reproduction occurs partly by tures, so discussion must be limited to general
isidia, which occur in 61% of the neotropical morphology.The genera which share the most
taxa,and can act as short-distancedispersalunits characterswith PhyllopsoraareBacidia,Biatora,
(Henssen& Jahns, 1973). Ascospores,isidia and Eschatogonia and two additional newly de-
thallus fragments may also be transported by scribed genera,Bacidiopsoraand Squamacidia,
rainwateror by animals, particularlyinsects. and species which previously were assigned to
Psorella.
Some species of the genera discussed above
PHYLOGENETIC RELATIONSHIPS havebeenconfusedwithPhyllopsorabecausethey
AND SYSTEMATIC POSITION also have a squamulosethallus with an obvious
OF PHYLLOPSORA prothallusand biatorine apothecia. Phyllopsora
can be separatedfrom these generaby the com-
Phyllopsorahas been placedin the orderLeca- bined charactersof its ascus type, its apothecial
norales(Poelt, 1973) because of its open, round, type composed of highlygelatinizedhyphaewith
disk-shapedapotheciawith radiateexciple, true no clear distinction between the exciple and hy-

22 Flora Neotropica
TROPICAL AMERICA Flora Neotropica base map no 1
-..'?.-?'
._ .......... . ... -- ,----'_
-.....- .--'_' -...................... -. - -----
-2-
>
't~zeC~ .m-...X^ ^
20. ..3. . . ...
soraglabella (squares).
FIG. 13.
FIG. 13. Neotropical
Neotropical distribution
distribution of Phyllopsora fendleri (stars),
Phyllopsora fendleri Phyllopsora furfuracea
(stars), Phyllopsora furfuracea (dots),
(dots), and
and Phyllop-
Phyllop-
sora glabella(squares).
pothecium, with a similar texture in the center familywith the additionof a darkerstainingarea
and at the margin,and by the small, thin walled, aroundthe masse axiale. Biatora vernalishas the
rarelyseptate ascospores(Fig. 9a-d, Table III). same apothecialpatternas Phyllopsora,but with
BacidiaDe Not., as typifiedby B. rosella(Pers.) larger, thicker-walledascospores. It also has a
De Not., has an apothecial pattern with a cup- crustose thallus and is apparentlya primarily
shaped exciple and distinct hypothecium and temperate genus. Biatora efflorescensNyl., an
subhymenium of different textures. The asco- unusualsorediatespecies, resemblesPhyllopsora
sporesaremultiseptateand thick-walled(Fig.9h). in apothecial anatomy and has argopsin,which
As currently recognized, Bacidia is heteroge- may supporta close relationshipbetweenBiatora
neous. It is a very large genus, widespread in and Phyllopsora.
tropicaland temperateareas,and in need of sys- Eschatogonia Trev., a genus comprised of a
tematic revision. few tropicalspecies that have squamulosethalli,
BiatoraTh. Fr., as typifiedby Biatora vernalis is often collected togetherwith Phyllopsora.It is
(L.) Fr., is distinguishedfrom Phyllopsoraby its distinguishedfrom Phyllopsoraby its apothecial
ascus type. It has the basic type found in the type with a distinct subhymenium, and by the

Delimitation of Species 23
r.............>. -~----
----- ?;----^=^
-
\ Vs C->";L
0
Tropc_ ot Cancer
'
-- \ -
_-__ TropicVof_ _Caopricorn t
_I
FIG. 14. Worldwidedistributionof Phyllopsorabuettnerivar. buettneri(triangles),var. glauca (stars),var.

munda(dots).
presence of a lower cortex. The cortex has a spore dimensions (Fig. 9f) as discussed in Brako
distinct outer layer of uniformly arrangedcells (1989).
that is unique in the Bacidiaceae(Fig. 4d). The Within the Bacidiaceae,Phyllopsorahas a rel-
ascosporesare slightlylargerthan those found in atively richchemistryand lichen substancesmay
Phyllopsora,and are usually septate. also prove to be importantfor genericseparation
BacidiopsoraKalb has a similar thallus mor- in the family.
phology to that of Phyllopsora,but has an apo-
thecial pattern with an obvious subhymenium
and longer,multiseptateascospores(Fig. 9g). Al- DELIMITATION OF SPECIES
though the ascospores have thinner walls than AND INFRASPECIFIC TAXA
do ascosporesof Bacidia rosella,this genusis not
clearlydistinct from Bacidia. Delimitation of species and infraspecifictaxa
Squamacidia Brako can be separated from in lichens is notoriouslydifficultas their biology
Phyllopsoraand other generain the Bacidiaceae is so poorly understood (Karnefelt, 1979; Sip-
by its thick, pale apothecial margin and its dis- man, 1983; Tehler, 1983). Accordingto Article
tinct chemistry of lobaric and/or fumarproto- 13 of the InternationalCode of Botanical No-
cetraricacid. Apothecial anatomy is closest to menclaturea lichen "species"refers only to the
thatof Phyllopsora,but the hyphaeof the exciple fungalcomponent.This treatmentfollows Grad-
are expanded at the margin and covered with stein (1975) and Sipman(1983) in requiringtwo
granules. The ascospores are broader than in or more independent charactersfor separation
Phyllopsoraand somewhatirregularin shape(Fig. of taxa at the species level. The rank of variety
9e), and the thallus is squamulose, often with a is used for taxa that differ from their nearest
brightscarletpigment in the medulla. relative by only one characterand are not fully
PsorellaMull. Arg.,typifiedby P. pannarioides separatedgeographically.Varietiesrecognizedin
(Knight)Miill. Arg., is probably a synonym of this treatmentare distinguishedby chemicaldif-
Bacidiaas it agreeswith Bacidia in its apothecial ferences, mostly additive chemistry, and slight
anatomyand long, thick-walledascospores.The morphologicaldifferences.
other species previouslyplaced in Psorelladiffer Most Phyllopsoraspecies were previouslydis-
from Phyllopsora in apothecial anatomy and tinguishedby singlecharacterdifferences,usually

24 Flora Neotropica
Table III
Comparisonof genera
Genus Phyllopsora Bacidiopsora Squamacidia
Ascal type Bacidia-type Bacidia-type Bacidia-type
Apothecial type Biatorine Biatorine Biatorine
Phyllopsora-type Non-Phyllopsora Non-Phyllopsora
type type
Spores (um) thin-walled thin-walled thin-walled
simple-uni- uni-multi- simple-uni-
septate septate septate

7-20 x 2-5 20-4( x 2.5-4 25-40 x 2.5-4
Conidia rod-shaped ? ?
Thallus squamulose- squamulose squamulose

subfoliose
Vegetative ? isidia ? lobules isidia

propagules
? lobules
Thallus cortex Type 1, 2, or 1-2 Type I Type 1
Phycobiont Pseudochlorella Pseudochlorella ?
Lichen substances atranorin, atranorin fumar-
argopsin, pannarin, protocetraric
vicanicin, zeorin, & lobaric acid
unknowns
Distribution subtropical- tropical tropical

tropical

Delimitation of Species 25
Table III
Continued
Physcidia Eschatogonia Bacidia rosella Biatora vernalis
Bacidia-type Bacidia-type Bacidia-type Biatora-type
Lecanorine Biatorine Biatorine Biatorine
Non-Phyllopsora Non-Phyllopsora Phyllopsora-type
type type
thin-walled thin-walled thick-walled thick-walled
uni-multi- uni-multi- multi- simple-uni-
septate septate septate septate

12-25 x 2-3 12-20 x 2-3 60-65 x 3-5 12-15x 3-5
rod-shaped rod-shaped ?
squamulose- squamulose crustose crustose

foliose
? isidia ? lobules none none
Type 1 single layer Type I Type I

of cells prosoplect. prosoplect.
Pseudochlorella Pseudochlorella
atranorin, unknown atranorin none

divaricatic
acid
tropical tropical temperate temperate

26 Flora Neotropica
TROPICAL AMERICA FloraNeotropicabase map no 1
" 2o- :;
'-,et,, ? -- ' 3 '
~
R~~~~~~~prr~~~~~~~~~~~~~d~
.. .~drl
'~' . t
,'
=- 10 s0 .0 80 '70 60 *0...
Preparedby HcndrikR Rypk\*.
FIG, 5, Notropcal istriutio of Pyllosorabuetteri ar. gauca(star), vr. muda .i (o
0 1 979 by the Univrsity of Utrecht Publishod by the State Univrsity of Utrecht, th Notherl.nds Deprtment of Systematic Botny
FIG. 15. Neotropical distribution of Phyllopsora buettneri var. glauca (stars), var. munda (dots).
in squamule morphology, color of the thallus, highly variable in most species. Apothecial pig-
prothallus,or apothecia,secondarychemistryor ments are useful charactersfor distinguishinga
ascosporedimensions. Species in this study are few species. Chemical charactersare used for
separatedby a combination of charactesbased separation of species only when they correlate
mostly on mycological characters. Squamule with other differences. Ascospores sizes are
morphologyis useful in some species. Color of broadlyoverlappingin many species.
the thallus and prothallus is not useful as it is
TAXONOMIC TREATMENT
Key to the Neotropical Species of Phyllopsora
la. Upper surfaceof thallus roughand fibrillose,white. ................................ 7. P. cuyabensis.
lb. Upper surfaceof thallus smooth or pruinose,greenishto brown.
2a. Thalluspruinose,squamulesgenerallyascendingto expose a white, cottony undersurface. .....
............................................................................ 2. P. buettneri.

Taxonomic Treatment 27
TROPICALAMERICA FloraNeotropicabase map no 1

P
I11 0
9I 70 60 so 0
II I d
'* O ---- _ .
2?0.1.
too
oao
Zoo
Zoo
zoo ^
|
metes~~~~~~~~~
ti-oo \ ;~<t~ ^^ y l ~ -
ep.W.~a~b,
H4.wxk*~
FtRyp&. ,
0 1979 by th Unnerty of Utrecht Published by the State University of Utrecht. the Netherlands Department of Systematic Botany
FIG. 16. Neotropical distribution of Phyllopsora corallina var. corallina (closed stars), var. glaucella (tri-
angles),var. ochroxantha(dots),var.phaeobyssina(squares),var. rappiana(asterisks),var. santensis(open stars).
2b. Thallusnot pruinose,squamulesgenerallyadnate.

3a. Squamuleslarge,lobes more than 0.5 mm wide.
4a. Thalluscomplanate,squamuloseto subfoliose, marginsdeeply incised;ascospores7-13
x 2-4 m. ........................................................ 16. P. parvifolia.
3-4 m. ..... . ...................... ...................... 8. P. fendleri.

3b. Squamulesminute to medium-sized,lobes less than 0.5 mm wide.
5a. Squamulessmall to medium-sized,lobes 0.1-0.5 mmwide.
6a. Thallus isidiate.
7a. Isidia all globose and proliferatingby budding;apotheciabrown. ... 13. P. kalbii.
7b. Isidia variable,globose but not proliferating,cylindricalor irregular;apothecia
tan or dark red.
8a. Ascosporesovoid, 4-6.5 x 2-3 ,m. .................... 17. P. parvifoliella.
8b. Ascosporesellipsoid, more than 6.5 Mmlong.
9a. Ascospores 12.5-19 Mmlong. ....................... 14. P. longiuscula.
9b. Ascospores6.5-13.0 mmlong.
lOa. Cortex type 1 or 1-2, more than 20 Mmthick, not extendingto
the lower surface. ........................ .... 6. P. corallina.

28 Flora Neotropica
lOb. Cortexa thin gelatinouslayer, less than 20 ,tm thick, extending

to the lower surface. ....................... 11. P. intermediella.
6b. Thalluslackingisidia.
1la. Apotheciasuffusedwith a darkred pigment. ................. 4. P. chlorophaea.
1lb. Apothecianot suffusedwith a darkred pigment.
12a. Squamules translucent;cortex with an orange pigment; conidia bent;
restrictedto the Juan FernandezIslands. .................... 1. P. bibula.
12b. Squamulesnot translucent;cortextan or colorless;conidia straight;other
areas.
13a. Ascosporesovoid, less than 6.5 Amlong. ........... 10. P. glabella.
13b. Ascosporesellipsoid, more than 6.5 ,tm long.
14a. Squamulesmostly ascendingand exposing a cottony white
underside. ..................... 2b. P. buettneri.var. glauca.
14b. Squamulesadnate.
15a. Pd+ yellow-orange........ 6f. P. corallinavar. santensis.
15b. Pd-.
16a. Squamulesdeeply incised; cortex type 2, more
than 20 m thick. ................. 5. P. confusa.
16b. Squamulesnot deeply incised; cortex a thin ge-
latinous layer less than 10 Mmthick. . 15. P. minor.
5b. Squamulesminute, ca. 0.1 Mum wide.
17a. Thallus containingfurfuracein;apothecia containingan orange-brownsubstance.
.............................................................. 9. P. furfuracea.
17b. Thallusnot containingfurfuracein;apothecianot containingan orange-brownsub-
stance.
18a. Isidia always present,finely branchedwith a roughsurface. ... 12. P. isidiotyla.
18b. Isidia present or lacking, when present smooth and cylindrical,not finely
branched.
19a. Cortexwell developed, type 2, ca. 20 Mrm thick; matureapotheciatan
with a red margin;ascospores 11-15.5 x 2.5-3 Am. . 18. P. subcrustacea.
19b. Cortexa thin gelatinouslayerless than 10 ,m thick;apothecialmargin
tan to brown;ascospores6.5-9.5 x 2.5-3 um. ...... 3. P. canoumbrina.
pannarin,vicanicin, zeorin and several new or

DESCRIPTION OF unknowncompounds.
THE GENUS Apotheciato 1.5 mm diam., simple or aggre-
gated,attachedlaminallyto the squamules.Disc
Phyllopsora Miiller Argoviensis. Bull. Herb. plane to convex, tan to dark reddish-brown,
Boissier 2(Appendix 1): 11. 1894. Type spe-
cies. Phyllopsorabreviuscula(Nylander)Miil- epruinose, margin often slightly raised. Exciple
ler Argoviensis(lectotypedesignatedby Clem- composed of radiating hyphae, tan to yellow-
brown or dark red, sometimes containingcrys-
ents & Shear, 1931). tals or pigmentsreactingKOH- or KOH+ red
Thallus squamulose, rarely appearingsubfo- or purplish-brown.Hypotheciumtan to yellow-
liose or crustose. Squamules0.1-1.0 mm wide. brown, or dark red, sometimes containingcrys-
Isidia common, sometimes dominatingthe thal- tals or pigmentsreactingKOH- or KOH+ red
lus. Upper cortex 5-60 Aimthick, consisting of or purplish-brown.Hymenium 20-60 im tall,
anticlinally oriented, thin- to thick-walled hy- amyloid, epihymenium indistinct or a thin ge-
phae with roundto narrowlycylindricallumina, lantinouslayerwith slightpigmentation,KOH-.
commonly containing lichen substances. Pho- Paraphysescellular,straight,sparinglybranched
tobiontin a continuous layer, green, unicellular, and anastomosing, apical cell slightly swollen,
5-15 ,umdiam., genuswhereknown,Pseudochlo- length to width ratio 6:1. Asci elongate-clavate,
rella.Medullausuallypoorlydeveloped,of loosely with a well developedtholuswith a paler,conical
woven and non-amyloidhyphae,frequentlycon- masse axiale.Ascosporeseight in the ascus, sim-
taining lichen substances.Lower cortex absent. ple to uniseptate, colorless, ovoid, ellipsoid to
Prothalluspale to dark red. Lichen substances: fusiform, smooth-walled,without a halo, (4.5-)
argopsin, atranorin, norargopsin,norvicanicin, 5.0-20.0 x 2.0-5.0 Am.

Description of Neotropical Species 29
Pycnidia spherical,immersedin the thallus to of Phyllopsoraparvifolia,but the squamulesare

superficial,outer wall tan to reddish-brown,os- much smallerin P. bibula.Phyllopsorabibulais
tiole pale to brown pigmented. Conidiophores closest to P. corallinavar. corallina,but P. bibula
irregularin shape, sometimes branched.Conid- has smaller, more fibrillose squamules, it lacks
iogenous cells elongate, enteroblastic. Conidia isidia, and has an orangecolor in the cortex.The
rod-shaped,straightor bent, 7.0-15.0 x 0.5-1.0 conidia are distinctly bent. Additional material
Jim. is need for clarificationof this species.
Distribution:Pantropicaland subtropicalwith
rareextensions into the temperatezone. 2. Phyllopsora buettneri (Muller Argoviensis)
Habitat: Primarily corticolous, sometimes Zahlbruckner,Cat. lich. univ. 4: 396. 1926.
growing on rock or over bryophytes,rarely on Figs. 14, 15, 17a-d.
leaves or dead wood; from sea level to 3500 m,
Psorabuettneri
MiillerArgoviensis,
Bot.Jahrb.Syst.
most common in humid subtropical montane 15: 506. 1893. Lecidea buettneri(Miller Argovien-
forest between 500 and 2000 m. Ber.Thatigk.St. GallischenNa-
sis) Stitzenberger,
turwiss.Ges. 1893-1894:246. 1895. Type.Togo.
Bismarksburg, Bittners.n. (holotype,G, pannarin
DESCRIPTION OF andzeorin;isotype,BM).
NEOTROPICAL SPECIES
Thallus squamulose, squamules elongate, in-
OF PHYLLOPSORA cised or lobulate,0.3-1.0 mm wide, complanate
to thick and convex, adjoined and overlapping,
1. Phyllopsorabibula(Taylor)Swinscow& Krog, adnateor often ascendingand exposinga cottony
Lichenologist13: 239. 1981. Fig. la. white lower surface,Pd+ orange.Upper surface
LecanorabibulaTaylor,Lond.J. Bot. 6: 160. 1847. pubescentor obviously pruinose,fibrilloseat the
Type.Chile.JuanFernandezIslands,[Apr 1830, margin.Isidia cylindrical,usuallylacking.Cortex
Bertero1648] (holotypeFH-TAYLOR684; isotype 1-2, 35-50 Amthick. Prothallusabundant,
types,BM,fattyacidsreported
bySwinscow &Krog, red. Lichen substances:argopsin, norargopsin,
1981, G = 2 x, H-NYL 20540, H-NYL P.M. 4109,
PC-MONTAGNE). pannarin,phyllopsorin,chlorophyllopsorin,un-
known compound B, and zeorin.
Thallus squamulose, squamules translucent, Apotheciacommon. Disc plane to convex, tan
round to irregular-elongate,0.1-0.3 mm wide, to darkred. Marginnot raised,concolorouswith
convex, closely adjoined, adnate to slightly as- the disc or darker.Exciple and hypotheciumtan
cending,Pd-. Upper surfacefibrillose,fibrillose to partlyred pigmented,KOH- or the pigment
at the margin. Isidia lacking. Cortex type 1-2,
reacting KOH+ purplish-brown. Hymenium
30 jLmthick, cortical gelatin orange. Prothallus colorless, to 30 ,m tall. Epihymeniumnot ob-
pale. Lichen substances:fatty acids reportedby vious. Ascosporessimple,shortfusiform6.0-13.0
Swinscow and Krog (1981). x 2.0-3.0,m.
Apotheciacommon. Disc planeto slightlycon- Pycnidia common, dark reddish brown, par-
vex, tan to reddish-brown.Marginslightlyraised tially immersed in the thallus. Conidia rod-
and darkerthan the disc. Exciple and hypothe- shaped, straight,9.0-10.0 x 0.5-1.0 ,m.
cium partlyred pigmented,KOH-. Hymenium Discussion.This species is distinguishedby its
pale, to 35 ,m tall. Epihymeniumnot obvious. pubescent or pruinose, incised, lobulate or in-
Ascosporessimple, ellipsoid to short fusiform, frequentlyisidiate squamulesand distinctchem-
9.0-10.5-12.0 x 2.5-3.0(-3.5) im. istry. The isidia may be heavily pruinose and
Pycnidia tan to reddish-brown, mostly im- fibrillose.Three varieties are recognizedby dif-
mersedin the thallus. Conidiarod-shaped,bent, ferencesin chemistrycorrelatedwith differences
9.0-14.0 x 0.5-1.0 ,m. in thallus morphology and distribution. The
Distribution.Juan FernandezIslands. squamulemorphologyvaries greatly.
Discussion. The above description is tenta-
tive, as this species is only known from the orig- Key to Varieties of
inal material, which is in poor condition. The
Phyllopsora buettneri
squamulesare very shiny and translucent.Zahl-
bruckner(1926-1927) listed this as a synonym la. Thalluscontaining
phyllopsorin.. var.munda.

30 Flora Neotropica
lb. Thalluslackingphyllopsorin. Distribution (Figs. 14, 15). Pantropical, sea

2a. Thalluscontainingpannarin ......... level-3500 m.
......................... var. buettneri.
2b. Thalluscontainingargopsin. . var. glauca. Discussion. This variety differs from var.
buettneri by its chemistry of argopsin, norargop-
2a. Phyllopsora buettneri (Muller Argoviensis) sin, vicanicin, norvicanicin, ? unknown B, and
Zahlbruckner var. buettneri. +zeorin, and its squamules which are usually
Fig. 17a.
smaller (ca. 0.3 mm wide), more ascending, and
The type variety is distinguished by its chem- less frequently pruinose than those of the other
istry of pannarin and zeorin. The squamules are P. buettneri varieties. The ascospores are 7.C-
0.3-1.0 mm wide and heavily pruinose. 9.5-12.0 x 2.5-2.0-3.0 jm.
Distribution (Fig. 14). Tropical Africa; north- Chemical strains: Chem. strain I: argopsin,
ern Australia, Papua-New Guinea, Phillipines, norargopsin, vicanicin, norvicanicin, and +zeo-
Society Islands, 900-2800 m. rin. Chem. strain II: argopsin, unknown B, and
?zeorin. Chem. strain III: argopsin and +zeorin.
Representativespecimensexamined.AUSTRALIA.
QUEENSLAND: Six km from Yabba Creek, no. 2 turnoff,
Sep 1981, KrogAU 27/12 (0). Chemical Strain I
KENYA.WESTERN PROV.: KakamegaDistrict,Kak-
amegaForest,nearforeststation, 1700 m, 20 Jan 1970, Specimens examined. MEXICO. GUERRERO:
Mun.
R. Santesson21688 (UPS). GeneralHeliodoroCastillo, 31 km N of El Paraisoon
UGANDA. BUGISUDISTRICT.:North Bugisu, one roadto Puertodel Gallo, 2000 m, 9 Jun 1985, Thomas
half km N of ford over Sasa River, 2800 m, Jun 1970, & Contreras3766A (NY).
Swinscow 2U 45/49B (BM). MASAKADISTRICT:Bu- PANAMA. PANAMA:Along trail to Cerro Brewster
koto, N edge of MalabigamboForest, 1100 m, Dec from Rio PacouraValley, 670 m, 18-21 Nov 1985,
1971, Swinscow 3 U25/8 (BM). WESTMENGODISTRICT: Brako 8258, 8295B (NY). Along the Llano-CartiRd,
Mawakota,MpangaForestReserve,1500m, June1970, 400 m, 24 Nov 1985, Brako 8371 (NY).
Swinscow2U 41/4 (BM). CUBA. SANTIAGO DE CUBA:La Gran Piedra, SE of
TANZANIA. TANGA PROV.: Usambara Prov., peak, 1000 m, 2 Apr 1982, Harris 14216 (HAC, NY),
Amani, 900 m, 9 Jan 1971, Moberg 1480D, 1464A 4 Apr 1982, Harris 14325, 14379 (HAC, NY), 6 Apr
(UPS), Vitikainen9292 (H). 1982, Harris 14555 (HAC, NY), 7 Apr 1982, Harris
MADAGASCAR. TAMATAVE PROV.:Perinet = An- 14656 (HAC, NY). Without prov. Wrights.n., Lich.
desibe, E slope of Mt. Maromiza, 1150 m, 12 May Cub. 180 (BM = 2x,G = 2x,L = 2x, M, UPS, US),
1984, Aptroot13589 (Hb. Aptroot). 186 p.p. (BUF, FH-TUCK 2919, 2920 p.p., M).
PHILLIPINES. LUZON.BENGUET PROV.:Mt. Santa JAMAICA. 1885, Hart s.n. (FH-TUCK 2921)
Tomas, 2200 m, Feb 1987, Aptroot20350 (Hb. Ap- DOMINICAN REPUBLIC. INDEPENDENCIA: Sierra
troot). de Baoruco,30 km S of PuertoEscondito, 1940 m, 25
PAPUA NEW GUINEA. MOROBEPROV.:Wau area, Jan 1987, Buck 14618 (NY). PEDERNALES:
Las Abejas,
SW slope of Mt. Missim, Poverty CreekValley, 1350 1150 m, 7 May 1982, Buck 8348 (NY), Harris 15563
m, 6 Aug 1981, Sipman 15692 (U). (NY).
TAHITI. H. Smith BotanicalGarden,Papeari,sea PUERTO RICO. HUMACAO:Caribbean National
level-10 m, 3 Feb 1988, Brako 9383 (NY). Forest Luquillo Division, Mt. El Toro, trail from El
Verde side on Hwy 186, 1000-1074 m, 4 Jun 1988,
Harris22217, 22234 (NY),ElYunqueRecreationArea,
2b. Phyllopsora buettneri (Muller Argoviensis) trail up Monte Britton,ca. 850--950 m, 9 June 1988,
Harris22495 (NY).
Zahlbruckner var. glauca (Bouly de Lesdain) GRENADA. 17 Oct 1905, Broadways.n. (NY).
Brako, Mycotaxon 35(1): 12. 1989. VENEZUELA. ANZOATEGUI: CerroPeonia, above
Fig. 17c, d. Los Pajaritos,2-3 Dec 1981,Davidse&Gonzdles19971
(VEN). ARAGUA:
6.5 km from Colonia Tovar along the
Phyllopsoraparvifolia(Persoon)MiillerArgoviensisvar. La Victoria-Tovarroad,6 Apr 1986,Brako8610 (NY).
glauca Bouly de Lesdain, Rev. Bryol. Lichenol. 7: BOLIVAR:
Parque Nacional Canaima, La Gran Sabana,
60. 1934. Type. Cuba. [Oriente:]Loma del Gato, on roadfromAeropuertoLuepato Kavanayen,10 Apr
1100 m, Jul 1931, Hioram 9098 (lectotype,HAC, 1985, Brako 8164 (NY). DISTRITOFEDERAL:
Caracas,
designated by Vezda, 1969 in hb., fragmentNY, trackfromAltamirato La Selva, 1900 m, 13 Jan 1979,
argopsin, vicanicin, norvicanicin, zeorin); isosyn- Sipman 10691 (B). MERIDA:Sierra Nevada de Santo
type: Cuba. Scala Victoria, 800 m, Hioram 8909 Domingo, Paramode Mucabaji,3500 m, 11 Oct 1981,
(UPS). L6pez-Figueiras& Lindstrom 26986 (MERF). MI-
Phyllopsora melanoglauca Zahlbruckner,Denkschr. RANDA: El Volcan, above Baruta, 1200-1400 m, 4 May
Kaiserl.Akad. Wiss., Math.-Naturwiss.K1.83: 133. 1986, Brako 8674, 8678 (NY). TRUJILLO:Near the
1909. Type. Brazil.Sao Paulo:Near Iguape,20-100 T.V. tower, at the base of Paramo la Nariz, on the
m, Aug 1901, Schiffners.n. (holotype, W; isotype, Carache-LasPalmasroad,2400 m, 24 Jan 1980,L6pez-
BM, argopsin,vicanicin, norvicanicin,zeorin). Figueiras22033 (MERF).

FIG. 17. a. Phyllopsora buettneri (Holotype: Togo, Bismarksberg, Dr. Buttner s.n., G) 12 x. b. Phyllopsora
buettneri var. munda (Holotype: Brazil, Hamburgerburg, Malmes.n., Lich. Regnell. 617b, S) 12 x. c. Phyllopsora
buettnerivar. glauca (Holotype of PhyllopsoramelanoglaucaBrazil, Schiffners.n. (W) 12x. d. Phyllopsora
buettneri var.
buettneri var. glauca Ilha Comprida,
(Brazil, Ilha
glauca (Brazil, Kalb 313,
Comprida, Kalb Hb. Kalb)
313, Hb. 12 x.
Kalb) 12 x.
TRINIDAD. 10 mile track,Arima Road, Jan 1958, m, 20 Aug 1982, Aptroot& Hensen 10726 (Hb. Ap-
Flemings.n. (NY). troot).
SURINAM. Brokopondo,Brownsberg,450 m, 25- PERU. SAN MARTIN:Lamas, Cerro Blanco (ca. 58
29 Jan 1985, Aptroot14909 (U), 500 m, 24 Mar 1978, km W-NW of Tarapoto), 1000 m, 14 Mar 1981, R.
v.Looy 16 (U); 25 Feb 1961,Kramer&Hekking3037a Santesson& ThorP73:4 (S).
(U). BRAZIL. PARANA: Prainhas,near Porto de Cima,
FRENCHGUIANA. Mountainridge 40 km SW of 100 m, 16 Jan 1987,Hatschbach&Brako8845 (MBM,
Cayenne,100 m, Mar 1985, Aptroot15595 (U); Saiil, NY). Rio GRANDE DOSUL:South of Torres, on road
primaryforest near Roche Bateau, 200-300 m, Mar to CampingItapeba, 10 m, 9 Jan 1987, Falkenberg&
1985,Aptroot15416 (U), 2 km SW of the village,"sen- Brako3604, 3605, 3606, 3608, 3609, 3614, 3615, 3616
tier Limonade,"180-210 m, 20 Jun 1986, Montfoort (FLOR,NY). SXo PAULO: Camposdo Jordao, 150 km
&Ek 100, 101 (U), 15 Aug 1986, Montfoort&Ek 127, NE of Sao Paulo, 1700 m, 25 May 1978, Kalb&Plobst
129, 147 (U). 20 p.p. (Hb. Kalb);Ilha Comprida,3 m, 15 Jul 1979,
ECUADOR. AZUAY: W slopes of CordilleraOcci- Kalb 163 p.p. (Hb. Kalb), 2 m, 1 Nov 1980, Kalb 313
dental, 55.5 km W of Cuenca on road to Molleturo, (Hb. Kalb.),nearCanan6ia,15 Jul 1979, Kalb& Poelt
3100 m, 2 Mar 1985, Arvidssonet al. 7132 (GB). s.n. (GZU); Ilha de Sao Sabastiao(Ilha Bela), ca. 250
Banos, N-slope of Tungurahua, 2300
TUNGURAHUA: m, 6 Jul 1979, Kalb & Poelt s.n. p.p. (GZU); Morro

32 Flora Neotropica
GrandenearCotia, ca. 25 km from Sio Paulo, 850 m, Sierrade San Luis, Sabanade Paraguariba,1400 m,
27 Sep 1980, Kalb 309 p.p. (Hb. Kalb);Rio Itaguare, 13 Oct 1984, L6pez-Figueiras31083 (MERF). MI-
ca. 20 km E of Bertioga,13 Apr 1980, Kalb325, Lich. RANDA: El Volcan, above Baruta, 1200-1400 m, 1-4
Neotropici 291 (BM, COLO,GZU, H, M, US). May 1986, Brako 8668, 8686B (NY).
KENYA. CENTRAL PROV.:KirinyagaDistrict, Mt. TRINIDAD. Mora Forest, east of SangreGrande,
Kenya, S of CastleForest Station, 2000 m, Feb 1972, 8-9 Mar 1920, Britton & Hazen 390 (NY); 10 mile
Krog48/176 (0). COAST PROV.: KwaleDistrict,Shim- track,ArimaRoad,Jan 1958,Flemings.n. (NY);Guia-
ba Hills, 480 m, Feb 1972, SwinscowK 43/1 (BM). guiare,20 May 1957, Fleming s.n. (NY).
EASTERNPROV.:Meru District, Mt. Kenya, 2100 m, GUYANA.EastDemerara,Timehri,DakaraCreek,
Feb 1974, Swinscow3K 16/19 (BM). Thompson'sfarm, 10 m, 2 Feb 1985, Sipman & Ap-
TANZANIA.TANGA PROV.: Usambara,1894, Holst troot 17947 (B). Upper Mazaruni,trail from Kamar-
1431 (G);LushotoDistrict,LutindiForest,Hoist 3330 ang river to PwiPwi mountain, ca. 10 km N of War-
p.p. (G). amadan,ca. 800 m, 28 Feb 1985, Sipman & Aptroot
UGANDA. MASAKA DISTRICT: Bukoto Co., Mala- 19392, 19411 (B).
bigambo Forest, 1100 m, 1980, Swinscow 3U 25/9 SURINAM. Brokopondo,Brownsberg,trailto Witti
(BM). Creek,ca. 500 m, 24 Oct 1981, Bekker1050 (U), Nat.
MADAGASCAR. FIANARANTSOA PROV.:Ambala- Res. Brownsberg,ca. 500 m, 23 Oct 1984, Zielman
manakara,30 km S of Ambositra, 1800 m, 1 May 1339A, 1359A (U).
1984, Aptroot& Hensen 12833 (Hb. Aptroot). FRENCHGUIANA. Cayenne,ex hb. Montagne(H-
NEW ZEALAND. North Island,Waima State For- NYL 20552, L); 1878, Leprieur754 (G); Saul, 2 km
est, WaitemaraGorgeTrack,28 Jan 1988,Brako9306, SW of the village,"sentierLimonade,"180-210 m, 22
9324 (AK, NY). Jul 1986,Montfoort&Ek 115 (U), 24 Aug 1986, Mont-
foort &Ek 148 (U), 21 Sep 1989, Montfoort&Ek 176,
24 Sep 1986, Montfoort& Ek 182 (U).
Chemical Strain II PERU. Cuzco: Ca. 11 km S of Pillcopata,850 m,
28 Mar 1981, R. Santesson & Thor P102:21 (UPS).
Specimens examined. COSTA RICA. CARTAGO: 20 km road distance
Turrialba,650 m, 21-22 Feb 1986, Berry4571 (NY). SAN MARTIN:Cerro Escalera (ca.
PANAMA. PANAMA: NE of Tarapoto), 900-1100 m, 13 Mar 1981, R.
Along trail to CerroBrewster
from Rio Pacoura valley, 670 m, 18-21 Nov 1985, Santesson & Thor P72:20, P72:53, P72:59, P72:73
Brako 8267, 8277, 8289 (NY); along the Llano-Carti (UPS).
BRAZIL. ESPIRITO SANTO:Mun. Guarapiri, 0.3 km
Road, 400 m, 24 Nov 1985, Brako8360, 8370, 8372, E
8383, 8385, 8390 (NY);CerroJefe-Alto Pacouraroad, on side road to Praia Setiba, 24 Feb 1988, Thomas
28 Nov 1985, Brako 8497 (NY). et al. 6157 (NY). PARANA: ReservaEcol6gicaSaptian-
CUBA. SANTIAGO DECUBA:La Gran Piedra, 1.5 km duva between Morretesand Antonina, 100 m, 17-18
SE of peak, 1000 m, 3 Apr 1982, Harris 14279 (HAC, Jan 1987, Hatschbach& Brako8860, 8899 (NY). Sio
PAULO: Praiade Pernubenear Itanha6m,5 m, 30 Mar
NY). Without prov.: La Prenda,Hioram 5314 (US);
1980, Kalb229 (Hb. Kalb);alongthe Rio Itaguar6,ca.
Wrights.n., Lich. Cub. 186 p.p. (BM, G, L = 2 x, M, 20km
UPS); Wrights.n., Lich. Cub. ser. 2, 728 (G). EofBertioga, 13 Apr 1980,Kalb231 (Hb. Kalb);
JAMAICA. Feb & Mar 1905, Cummings 44 p.p. Ilhabela, along the road from Ilhabela to Praia dos
(COLO, NY = 2x, 0). Castelhanos,250 m, 7 Jul 1979, Sipman et al. 13444
DOMINICAN REPUBLIC. LA VEGA:La Sal, 1100- (B);mangroveswampnearPraiade Pernube,nearItan-
1200 m, 29 Apr 1982, Harris 15005 (NY). ha6m,sea level, 9 Jul 1979, Kalb& Poelt s.n. (GZU =
PUERTO RICO. CAGUAS: Reserva ForestalCarite, 2x), Sipman et al. 13548 (B), 13553 (B, NY).
at intersectionof Hwys 184 and 179, ca. 900 m, 9 Jun KENYA. CENTRAL PRov.: Kiriyaga District, Mt.
1988, Harris 22447 (NY). HUMACAO: Caribbean Na- Kenya, S of Castle Forest Station, 1800 m, Feb 1977,
tional Forest,LuquilloDivision, northslope, 2000 m, Swinscow 5K 5/12 (BM).
27 Feb 1900, Heller4764 (NY), El YunqueRecreation TANZANIA. TANGA PROV.: Usambara,Hoist 3330
Area, 500 m, 5 Mar 1981, Buck 4090A (NY), Mt. El p.p. (G); Kwambuga,1894.
Toro, trail from El Verde side on Hwy 186, ca. 850 NEW GUINEA. CHIMBU PROV.: Mt. WilhelmArea
m, 4 Jun 1988, Harris 22248 (NY).; trail from Hwy Keglsugl,in the village, 2400 m, Mar 1987, Aptroot
988 to Rio Mameyes, 100-150 m, 6 Mar 1981, Buck 18669 (Hb. Aptroot).
4195 (NY).
DOMINICA. Dom-Can LoggingArea, Newfound- Chemical Strain III
land, 240 m, Jan 1969, Hale 35230 (US).
ST. VINCENT. Bow Woods, 240 m, Elliott 135 Specimens examined. CUBA. ORIENTE:Loma del
(BM). Gato, 900 m, 11 Jul-14 Aug 1921, Le6n et al. 10232
COLOMBIA. HUILA:Mun. La Plata, Vereda La (NY), Wrights.n. Lich. Cub. 186 (FH-TUCK 2920
Candelaria,E-slope, headwatersof the Rio La Can- p.p.).
delaria,2300 m, 1 Oct 1984, Aguirre& Sipman 6142 DOMINICAN REPUBLIC. INDEPENDENCIA: Sierra
(B). de Baoruco,30.5 km S of Puerto Escondito, 1940 m,
VENEZUELA. BOLiVAR:Parque Nacional Canai- 25 Jan 1987, Buck 14634 (NY); 23.5 km S of Puerto
ma, La GranSabana,on road from AeropuertoLuepa Esconditoat intersectionof road to CharcoColorado,
to Kavanayen,11 Apr 1985,Brako8183(NY).FALC6N: 1750 m, 24 Jan 1987, Harris20455 (NY); Charcode

la Paloma, 48.4 km S of Puerto Escondito, 1800 m, GUYANA. Upper Mazaruni,Mount Latipu, ca. 8
25 Jan 1987, Harris 20672 (NY). km N of Kamarang,ca. 800 m, 25 Feb 1985, Sipman
COLOMBIA.CUNDINAMARCA: BogotaD.E. Regi6n & Aptroot19165 (B).
de Monserrate,Acueductode Bogota, 3150 m, 4 Mar PERU. SANMARTIN: CerroEscalera(NE of Tara-
1980, Vargas236 (U); Choachi, 2600 m, Lindig 706 poto), NW of tunnel, 1000 m, 15 Mar 1981, R. Santes-
(FH-TUCK 2921, H-NYL P.M. 4113, M), 803 (M), son & ThorP74:122 (S).
2791 (H-NYL 20548), Lindig s.n. (H-NYL 20541). BRAZIL.Rio DEJANEIRO: Itatiaia,1850 m, 28 July
TRINIDAD. 10 mile track,Arima Road, Jan 1958, 1966, Eiten & Eiten 7495 (NY); between Registrodo
Fleming s.n (NY = 2x). Picfiand AgulhasNegras, + 1650 m, 10 Jul 1979, Kalb
ECUADOR. TUNGURAHUA: Banios, N slope of Tun- & Poelt "b"(GZU), 1800 m, 14 Mar 1980, Kalb 221
gurahua,2300 m, Aptroot& Hensen 10726 (Hb. Ap- (Hb. Kalb). SANTACATARINA: Between Uribici and
troot). CapivaraAlta on road to Bracodo Norte, 1040 m, 12
BRAZIL. PARA: 1849, Spruce s.n. (BM). PARANA: Jan 1987, Falkenberg & Brako 3708, 3731, 3798
ReservaEcologicaSapitanduva,betweenMorretesand (FLOR,NY), SAOPAULO: Camposdo Jordao,at base
Antonina,100 m, 17-18 Jan 1987,Hatschbach&Brako of Pico de Itapeva, 1760 m, 25 Jan 1987, Brako8925
8885-8888, 8898 (MBM, NY). (NY).
Discussion. Similar in morphology to var.
2c. Phyllopsora buettneri(Miller Argoviensis)
Zahlbruckner var. munda (Malme) Brako, My- buettneri, this variety differs by its chemistry of
cotaxon 35(1): 12. 1989. pannarin, phyllopsorin and zeorin. The asco-
Figs. 14, 15, 17b.
spores are 10.0-11.0-12.0 x 2.5-2.8-3.0 ,m.
Lecidea munda Malme, Ark. Bot. 28A(7): 49. 1936.
Phyllopsoramunda(Malme)Zahlbruckner, Cat. lich. 3. Phyllopsora canoumbrina (Vainio) Brako,
univ. 10: 377. 1939. Type. Brazil. Rio Grande do
Sul:Hamburgerberg, nearSaoLeopoldo,18Oct 1892, Mycotaxon 35(1): 12. 1989. Fig. Ib.
Malme s.n., Lich. Regnell. 617b (holotype, S, pan- Lecideacanoumbrina
Vainio, Proc. Amer. Acad. Arts
narin, phyllopsorin,zeorin). 58: 135. 1923. Type:Trinidad.MaravalValley,Jan-
LecideaschizophylloidesMalme,Ark.Bot. 28A(7):45.
1936. Phyllopsoraschizophylloides(Malme)Schnei- LecideaApr 1913, R. Thaxter19 (holotype,FH).
Biblioth. Lich. 13:178. 1979. parvifoliaPersoon var. subgranulosaTucker-
der, Type. Brazil.Rio man, Proc. Amer. Acad. Arts 6: 273. 1866. Biatora
Grandedo Sul:SilveiraMartins,7 Mar 1893,Malme
parvifolia (Persoon) Montagne var. subgranulosa
s.n., Lich. Regnell. 1251B (holotype, S, pannarin, (Tuckerman)Tuckerman,Syn. N. Amer. lich. 2: 8.
phyllopsorin,zeorin). 1888. Psora parvifolia (Persoon) Massalongo var.
Distribution (Figs. 14, 15). West Indies, South subgranulosa(Tuckerman)Muller Argoviensis, J.
Linn.Soc., Bot. 29:219. 1893.Phyllopsoraparvifolia
America and southern North America, 1000- (Persoon) Miiller Argoviensis var. subgranulosa
2200 m. (Tuckerman)Miiller Argoviensis,Bot. Jahrb.Syst.
20: 264. 1894. Type. Cuba.[Oriente:]Monte Verde,
Specimens examined. UNITED STATES. LOUI- 6 Apr, Wrights.n., Lich. Cub. 185 (lectotypehere
SIANA:East Baton Rouge Parish, Burden Research designated,FH-TUCK2923, piecemarked"w,"also
Plantation, of Essen Lane, SE Baton Rouge, 19 Feb in this packetare P. furfuraceaand P. intermediella;
1974, Tucker11956 (LSU). isolectotypes, BM, BUF, G = 3 x, L = 2 x, L-6889,
CUBA. SANTIAGO DECUBA:La Gran Piedra, 1000- no lichen subtances,M, UPS, US).
1200 m, 6 Apr 1982,Harris14559 (HAC,NY);Arroyo Lecidea granuliferaFink in Hedrick, Mycologia 22:
Negro, ca. 4 km SE of La GranPiedra, 1000 m, 5 Apr 252. 1930. Type. Puerto Rico. Rio Maricao, 600-
1982,Harris14444 (HAC,NY); vic. Museode la Fran- 720 m, 14 Feb 1915, Britton& Cowell4235 (holo-
cesca "La Isabelita,"ca. 2 km SE of La Gran Piedra, type, MICH;isotype, NY).
1000 m, 7 Apr 1982, Harris 14592 (HAC, NY).
DOMINICAN REPUBLIC. INDEPENDENCIA: Sierra Thallus squamulose, squamules minute, close-
de Baoruco, 23.5 km S of Puerto Escondito at inter- ly adnate, initially granular, irregularly crenate
section of road to CharcoColorado, 1750 m, 24 Jan to lobed, ca. 0.1 mm wide, complanate to
1987, Harris20493, 20495 (NY). convex, adjoined and closely adnate, Pd-.
VENEZUELA. ARAGUA:6.5 km from Colonia To-
var along the La Victoria-Tovarroad, 6 Apr 1986, Upper surface glabrous, fibrillose at the margin.
Brako 8605, 8617, 8619(NY). MRIDA: La Carbonera, Isidia cylindrical. Cortex a thin gelatinous layer,
SW of Merida towards La Azulita, 2399 m, 22 Jan 5-10 ,Amthick. Prothallus abundant, pale to red-
1979, Sipman & L6pez Figueiras 11028 (B, MERF), dish. Lichen substances: none detected.
2100-2200 m, 17 Dec 1984, Brako 8110 (NY); La
Apothecia common. Disc plane to convex, tan
Valle,NE of M6rida,ca. 2700 m, 28 Mar-7 May 1969, to brown.
Hertel&Oberwinkler10528 (M).MIRANDA: El Volcan, Margin not raised, concolorous or
above Baruta,1200-1400 m, 1 May 1986, Brako8662 darker than the disc. Exciple tan to partly red
(NY), 4 May 1986, Brako 8688 (NY). pigmented, KOH-. Hypothecium tan to yellow-

34 Flora Neotropica
brown, KOH-. Epihymenium not obvious. Hy- glabrous,fibrilloseat the margin.Isidia lacking.
menium tan, to 40 um tall. Ascospores simple, Cortex type 1-2, to 20 gm thick. Prothallus abun-
ellipsoid to short fusiform, 6.5-8.0-9.5 x 2.5- dant, red. Lichen substances:none detected.
2.7-3.0 ,m Apothecia common. Disc plane to convex, tan
Pycnidia common, tan, superficial on the pro- to darkreddish-brown.Marginslightlyraisedand
thallus. Conidia rod-shaped, straight, 8.0-10.0 darkerthan disc. Exciple and hypotheciumdark
x 0.5-1.0 am. red, KOH-. Hymeniumcolorless,to 35 Amtall.
Distribution (Fig. 18). Central America, West Epihymeniumnot obvious. Ascosporessimple or
Indies and South America, 140-300 m. uniseptate,ellipsoid to shortfusiform,8.5-10.0-
11.5 x 2.5-2.9-3.0 um.
Specimens examined. GUATEMALA. PETEN:Near
Tikal, 300 m, 9-10 Jan 1979, Kalb & Plobst 73 p.p. Pycnidia common, slightly emergent from
(Hb. Kalb). thallus,brown.Conidiarod-shaped,straight,9.0-
CUBA. Wrights.n. (FH-TUCK 2922). 12.0 x 0.5-1.0 um.
VENEZUELA. AMAZONAS:
Rio Negro, base of Cer- Distribution(Fig. 18). CentralAmerica, West
ro de la Neblina, 140 m, 20 Feb 1984, Buck 11028
(NY, VEN).
Indies, South America,Hawaii and Africa, 140-
BRAZIL. ACRE:Vizinhancade Sena Madureria,1 2300 m.
Nov 1980, Nelson et al. 491 (INPA, NY). AMAZONAS:
Specimensexamined.UNITED STATES. Hawaii:
Along IgarapeSanta Luzia,just off Rio Uatuma, 16, Forbess.n. (US).
17 Aug 1979, Buck 2882 (NY). SANTACATARINA:
Blu-
COSTA RICA. SANJOSE:La Divisi6n, 2000 m, 25
menau, 1891, Ule 96 (G). Feb 1986, Berry4576, 4577, 4578A (NY).
Discussion. This species is distinguished by its CUBA. SANTIAGO
DECUBA:Arroyo Negro, 1000 m,
5 Apr 1982, Buck 7693 (HAC, NY). Without prov.:
granular, minute squamules, small to medium-
Wrights.n., Lich. Cub. ser. 2, 107 (H-NYL 20546).
size ascospores and lack of chemistry. A collec- JAMAICA. Hart 34 (NY), 1884, Hart 153 (FH-
tion from Saul, French Guiana (Aptroot 15322 TUCK 2923).
(U)) is closest to this species, but has a smooth HAITI. Dept. de la Grand'anse:Massifde la Hotte,
710 m, 13 Nov 1982, Buck 9083 (NY).
crustose, non-isidiate thallus, tan apothecia and DOMINICAN REPUBLIC. PEDERNALES:
Las Abe-
ascospores 8.0-13.0 x 2.5-3.0 ,im. jas, 3900 ft, 13 Mar 1980,Buck4361, 4376 (NY), 3800
ft, 7 May 1982, Buck8362 (NY), 3600 ft, 5 May 1982,
Harris 15367 (NY). INDEPENDENCIA:
Sierra de Bao-
4. Phyllopsora chlorophaea (Miller Argovien- ruco, 23.5 km S of PuertoEscondito, 1750 m, 24 Jan
sis) Zahlbruckner, Denkschr. Kaiserl. Akad. 1987, Harris20473, 20480, 20487, 20489 (NY);Char-
Wiss., Math.-Naturwiss. K1. 83: 133. 1909. co de la Paloma, 48 km S of Puerto Escondito, 1800
m, 25 Jan 1987, Harris 20670, 20671 (NY).
Figs. Ic, 8d, 9a. VENEZUELA. AMAZONAS:
Rio Negro, base of Cer-
Psora chlorophaeaMullerArgoviensis,Flora 70: 320. rode la Neblina,140m, 25-28 Nov 1984,Brako7521B,
1887. Type. Brazil.Sao Paulo:Apiahy (Apiai),Jun 7526 (NY, VEN). BOLIVAR: La Gran Sabana, 11 Apr
1881, Puiggari 1721 (holotype, G, no lichen sub- 1985, Brako 8179A, 8181A (NY). DISTRITOFEDERAL:
stances;isotype, W). Caracas,1878, Ernst 38, 45 (G). MERIDA: La Carbo-
LecideahaemophaeaNylandervar.subparvifolia Miiller nera,Finca San Eusebio,along the Merida-LaAzulita
Argoviensis,Flora 60: 473. 1877. Phyllopsorasub- road, 2200-2300 m, 13 Apr 1976, L6pez-Figueiras
parvifolia(Miller Argoviensis)Miiller Argoviensis, 13714 (MERF),8 Oct 1977, L6pez-Figueiras& Keogh
Hedwigia34: 114. 1895. Type. Venezuela.Distrito 14099 (MERF).MIRANDA: El Volcan, above Baruta,
Federal:Caracas,Ernst 114 (holotype,G, no lichen 1200-1400 m, 4 May 1986, Brako 8685, 8687 (NY).
substances). TACHIRA: Patade Gallina,900 m, 28 Feb 1981, L6pez-
LecideafurfuraceaPersoon in Gaudichaudf. schizo- Figueiras& Rodrigues25645B (MERF).
phylla Vainio, Acta Soc. Fauna Fl. Fenn. 7(2): 47. PERU. PUNO:Carabaya,Hb. Nylander (H-NYL-
1890.Lecideaschizophylla(Vainio)Malme,Ark.Bot. 20502).
28A(7): 43. 1936. Type. Brazil. Minas Gerais: La- BRAZIL. MINASGERAIS:Between Vila Monte Ver-
fayette, 1885, Vainios.n., Lich. Bras. Exs. 335 (ho- de and Camanducaia,1300 m, 28-29 Feb 1980, Kalb
lotype, TUR-VAIN 22641, terpenes reported by 314 p.p. (Hb. Kalb);Lafayette,1885, Vainios.n., Lich.
Swinscow& Krog, 1981; isotype, ZT). Bras.Exs. 318 (TUR-VAIN 22640), 366 (TUR-VAIN
22642). Rio GRANDEDOSUL: Hamburgerberg, 10 Oct
Thallus squamulose, squamules round to elon- 1892, Malme s.n., Lich. Regnell.615 (UPS);Serrados
PadresnearCruzAlta, 21 Apr 1893, Malmes.n., Lich.
gate, lobed and incised, 0.1-0.3 mm diam., com-
Regnell. 1275 (S);SantoAngelonearcachoeira,30 Jan
planate to convex, closely adjoined and overlap- 1892, Malme s.n., Lich. Regnell. 236A (S); Serrados
ping, adnate to ascending, Pd-. Upper surface VallesnearCruzAlta, Malme s.n., Lich. Regnell.236b


lO II lOG 90 80 70 60 SC Go
?
X _ f
;"~ I:' Y!1^ :----
21
-JJCC
......~~~~~~~~~~~~~~~~~~~~~~~~~~r
-~~~-~~~-~
....
-'
'"'%. ................ i: .....,
r 2.
~~
"~p .. 'edt '1 ,./'e
C 179by heUnierity ofUtrch PulihedbytheStte nier,tyof trcht te Nthrl?dr DeprtentofSyre.
..;oan
"',1.Norpcldsrbto fPylpoacnumrn tinls,Pylpoaclrpaa(tr)

! Prepared by Hendnk R Rypkema \ 8 70 6 50
___
'
Phylopsoa coSusa(dot), ad Phll~pora uyabns- ........)
FIG. 18. Neotropical distribution of Phyllopsora canoumbrina (triangles), Phyllopsora chlorophaea (stars),
Phyllopsoraconfusa(dots), and Phyllopsoracuyabensis(squares).
(G, US, BM); Sao Franciscode Paula, Lago Sao Ber- cium, a colorless hymenium, and medium size
nardo, 900 m, 18, 19 Oct 1987, Osorio & Fleig 2SF/ ascospores.It is closest to non-isidiate forms of
61 (Hb. Osorio). SAo PAULO:Campos do Jordao, 150
km NE of Sao Paulo, 1700 m, 25 May 1978, Kalb & Phyllopsoraconfusa which usually have more
Plibst 20 p.p. (Hb. Kalb);Morro Grande near Cotia, narrow squamules and are never pigmented
25 km W of Sao Paulo, 850 m, 27 Aug 1980, Kalb309 throughoutthe exciple and hypothecium.
p.p. (Hb. Kalb).
KENYA. CENTRALPROV.:Kirinyaga District, Mt.
Kenya, 2 km NW of Irangi Forest station, 2000 m,
Feb 1977, Swinscow5K 4/53 (BM). 5. Phyllopsora confusa Swinscow & Krog, Li-
TANZANIA. ARUSHAPROV.:Mt. Meru, E slope, chenologist13: 229. 1981. Type. Kenya. Cen-
road to the crater,JekukumiaRiver, 2100-2200 m, 7 tral Prov.: Kirinyaga District, Mt. Kenya, 2
Jan 1971, R. Santesson23030 (GZU).
km N of IrangiForestStation, in damp decid-
Discussion. This species is distinguished by its uous forest near River Ena, 0?20'S, 37?28'E,
small to medium size, non-isidiate, thin squa- 2000 m, Feb 1972, Krog & Swinscow K 48/
mules, lack of chemistry, a dark red, KOH- pig- 177 (holotype, O, no lichen substances).
mentation throughout the exciple and hypothe- Fig. Id.

36 Flora Neotropica
Thallus squamulose, squamules elongate to FALCON: Top of mountain ridge 25 km NE of Coro

flabellate,deeply incised, 0.1-0.3 mm diam., ad- along La Negrita-Curimaguaroad, above La Chapa,
1250 m, 18 Jan 1979, Sipman & Vander Werff10899
joined and overlapping,complanate to convex, (B, MERF).MERIDA: Sierrade St. Domingo, Paramo
adnate to ascending, Pd-. Upper surface gla- de Mucubaji,3500 m, 28 Mar-7 May 1969, Hertel &
brous,occasionallyfibrilloseat the margin.Isidia Oberwinkler10474 (M); El Paramito,near El Morro,
not well developed, never cylindrical or erect. 2650 m, 21 Dec 1977,L6pez-Figueiras14704 (MERF);
Cortextype 2, 25 um thick, cortex often extend- Monte Zerpa,above La Hechicera,nearMerida,2000
m, 2 Apr 1980,L6pez-Figueiras22733 (MERF),4 Dec
ing to the undersideof the squamules.Prothallus 1980, L6pez-Figueiras25009 (MERF);Valley of the
thin, pale to reddish. Lichen substances:none Rio Chama, 1600 m, 21 Jan 1979, Sipman 10977 (B,
detected. MERF).SUCRE: Peninsulade Paria,Cerrode Rio Ar-
Apotheciacommon. Disc planeto slightlycon- riba, 700 m, 10-12 Aug 1966, Steyermark& Rabe
96279 (US).
vex, tan, orangeor yellow-brown.Marginslightly TRINIDAD. St. Ann's Heights, 17 Mar 1920, E.
raised,concolorous,paleror darkerthanthe disc. Britton684 (NY).
Excipletan to partlyredpigmented,KOH-. Hy- GUYANA. Upper Mazaruni:Trail from Kamarang
pothecium tan to yellow-brown, KOH-. Hy- riverto Mt. PwiPwi, 10 km N of Waramadan,ca. 800
menium pale tan, to 40 um tall. Epihymenium m, Sipman & Aptroot19489 (B).
SURINAM.Paramaribo,Palmgarden,sea level, Mar
not obvious. Ascosporessimple, short fusiform, 1985, Aptroot14840 (U).
8.5-10.0-12.0 x 2.5-2.7-3.0 Am. ECUADOR.AZUAY:Ca. 35 km S of Cuenca, 3200
Pycnidia not seen. m, 24-26 Aug 1987, Kalb&A. Kalb17576 (Hb. Kalb).
Distribution (Fig. 18). Pantropical, 100- BRAZIL. MATOGROSSO: 35 km S of Cuiaba, 120
3500 m. m, 4 Jul 1980, Kalb273 p.p. (Hb. Kalb);Chapadados
Guimaraes, 800 m, 7 Jul 1980, Kalb 277 p.p. (Hb.
Kalb), 9 Jul 1980, Kalb281 (Hb. Kalb); 10 km NE of
Specimens examined. UNITED STATES (Repre- Chapadados Guimaraes,680 m, 9 Jul 1980, Kalb282
sentative specimens). FLORIDA:Hillsborough Co., p.p. (Hb. Kalb);Buriti, 19 Jan 1894, Malme s.n., Lich.
HillsboroughState Park, 8 Mar 1947, Schallert1145 Regnell. 2268 (FH, S, US), 20 Jan 1894, Malme s.n.,
(US); Seminole Co., Sanford, 5 Jul 1911. LOUISIANA: Lich.Regnell.2267C (S),4 Jun 1894,Malmes.n., Lich.
IberiaParish,Weeks Island, 14 mi S of New Iberia,4 Regnell.s.n., 2737E (S);Rio Vermelho, 10 Nov 1894,
Aug 1958, Pursell3368 (COLO);St. MartinParish,St. Malmes.n., Lich. Regnell.s.n. (S);SantaAnnade Cha-
Martinville, 17 Oct 1894, Langlois s.n. (NY). pada, 7 Mar 1894, Malme s.n., Lich. Regnell. 2515B
HONDURAS. Quebradoel Gallo,NE slope of Cerro (S).MATOGROSSO DOSUL:30 km S of CampoGrande,
de Uyuca, 900 m, 14 Mar 1951, Morton 7045 (US). 550 m, 14 Nov 1979, Kalb199 p.p (Hb. Kalb).MINAS
NICARAGUA. ZELAYA:Vic. Bluefields, 21 Apr-23 GERAIS: Vila Monte Verde, 30 km E of Camanducaia,
May 1949, Standley20125 (F). 1500 m, 8 Aug 1978, Kalb & Plobst 95, Lich. Neotro-
PANAMA. PANAMA:Along the Llano-Carti road, pici 342 (GZU, M, US), 1500-1800 m, 7-11 Aug 1978,
400 m, 24 Nov 1985, Brako 8358 (NY). Kalb & Plobst 44 p.p. (Hb. Kalb); Between Caman-
CUBA. Camaguey:Cubito Prov., midway between ducaia and Monte Verde, 1300 m, 28-29 Nov 1980,
Camagueyand La Gloria, 19-21 Feb 1941, Johnson Kalb 314 p.p. (Hb. Kalb); Fazenda Sao Mateus E of
s.n. (NY = 2x). ISLE OF PINES: San Juan, 15-17 Mar Camanducaia,1800 m, 30 Nov 1980, Kalb 315 p.p.
1916, Britton et al. 15491 (NY = 2x). SANTIAGO DE (Hb. Kalb);MonteVerde, 1750 m, 3 Jul 1979, Sipman
CUBA:"Monte Kentucky,"ca. 5 km SE of La Gran 13053 (B, NY), 13068 (B). PARA:Serrado Cachimbo,
Piedra,ca. 1000 m, 5 Apr 1982, Harris 14424 (HAC, AeroportoCachimbo, ca. 430-480 m, 27 Apr 1983,
NY). Brako & Dibben 6241 (INPA, NY); Rio Jamanxim,
DOMINICAN REPUBLIC. LA VEGA:Arroyo Pied- 974-1024 km N of Cuiabf, ca. 300 m, 10-15 May
rosa, 1.2 km from Monabaoon road to Jarabacoa,11 1983, Brako & Dibben 7031 (INPA, NY); Santarein,
May 1982, Harris15679A(NY); km 17 on Jarabacoa- Spruces.n., Lich.Amaz.And. 210 (BM= 3 x). PARANA:
La Vega road, 11 May 1982, Harris 15750 (NY); San- ReserveEcol6gicaSapitanduva,betweenMorretesand
tiago, ParqueNacional J. Armando Bermudez,along Antonina,100m, 17-18 Jan 1987,Hatschbach&Brako
trail along Rio Los Guanos, near La Ci6naga,3500- 8854, 8866, 8870, 8879, 8883B, 8896 (MBM, NY);
4000 ft, 12 Jan 1987, Harris 19737 (NY). SAN JUAN: near Cacatu, 17 Jan 1987, Hatschbach& Brako 8907
Along streamjust E of "LaCompartici6n,"along trail (NY). Rio DEJANEIRO: Itatiaia,between Registrodo
up Pico Duarte, 2350 m, 14 Jan 1987, Buck 14196 Pici and AgulhasNegras,850 m, 22 Jul 1978, Kalb&
(NY). Plobst 40 (Hb. Kalb), 1800 m, 23 Jul 1978, Kalb &
ST. VINCENT. RichmondPeak, 800 ft, Elliott 264 Plobst 37, 161 (Hb. Kalb). SANTACATARINA: 13 km
(BM). above Timbe do Sul, on road to SerraRochina, 650
COLOMBIA. TOLIMA: Mun. Santa Isabel, El m, 11 Jan 1987, Falkenberg& Brako 3655 (FLOR,
Ochoral,3130 m, 1980, Valencia&BoekhoutVB177J NY); Between Uribici and CapivaraAlta on road to
(B). BraSodo Norte, 1040 m, 12 Jan 1987, Falkenberg&
VENEZUELA. ARAGUA: La Mora, W of Colonia Brako 3723 (NY); Pinheiral and Ruderal Riozinho,
Tovar, 1200 m, Dec 1986, Brako 8510, 8511 (NY). 1000 m, 22 Jan 1957, Smith & Reitz 10287 (US =

2 x). SAOPAULO: Serrade Paranapiacaba,80 km SW cies is closest to Phyllopsorachlorophaeawhich

of Sao Paulo, 700 m, 24 Mar 1978, Kalb & Plobst 4 is distinguishedby its completelyred pigmented
(Hb. Kalb);Camposdo Jordao, 1700 m, 25 May 1978,
Kalb & Plobst 20B p.p. (Hb. Kalb); Pico de Itapeva, apotheciaand its broadersquamules,and P. cor-
above Camposdo Jordio, 1820 m, 14 Nov 1978, Kalb allina var. corallina which has broader squa-
& Plobst 50 p.p. (Hb. Kalb);near Socorro, 130 km N mules and erect isidia.
of Sao Paulo, 700 m, 14 Apr 1979, Kalb & Assis 174,
Lich. Neotropici 241 (BM, COLO,GZU, H, M, US);
near Cachoeirasde Emas, ca. 15 km NE of PiraCu- 6. Phyllopsoracorallina(Eschweiler)MiillerAr-
nunga, 550 m, 14 Jun 1979, Kalb & Plobst 145 p.p. goviensis, Bot. Jahrb.Syst. 20: 264. 1894.
(Hb. Kalb); Sao Sebastiao, 400 m, 6 Jul 1979, Kalb Figs. le, f, 5, 8c, 16.
156 (Hb. Kalb),600 m, 7 Jul 1979, Sipman 13408 (B);
Serrado Garraozinho,between Moji das Cruzesand LecideacorallinaEschweilerinMartius,Fl.bras.enum.
Bertioga,850 m, 29 Mar 1980, Kalb 227 (Hb. Kalb); pi. 1(1):256. 1833.Parmeliacorallina(Eschweiler)
Serrade Paranapiacaba,60 km SWof Sao Paulo,above Montagne in Sagra,Hist.fis.Cub.,Bot.4:216. 1838.
Juquitiba,550 m, 27 Apr 1980, Kalb233 (Hb. Kalb); Biatoracorallina(Eschweiler) Hookerf. &Taylorin
FazendaSao JoaonearRio Claro,600 m, 16 Aug 1980, Hookerf., Fl.Antarct.2: 537. 1847.Psoracorallina
Kalb 302 p.p., (Hb. Kalb);MorroGrandenear Cotia, (Eschweiler)MillerArgoviensis,Flora71:527.1888.
850 m, 27 Aug 1980, Kalb 309 p.p. (Hb. Kalb). Phyllopsoraparvifolia(Persoon)MullerArgoviensis
CHILE. CAUTiN: Villa Rica, 12 Jan 1953, Thomas- var.corallina(Eschweiler) Boulyde Lesdain,Lich.
son s.n. (UPS). Mex. 20. 1914.Type.Brazil.Bahia,Martiuss.n..
PARAGUAY. ALTO PARANA:Reserva Biologica del (holotype,M, no lichensubstances; fragments, H-
Itab6, ca. 37 km NNW of Herandarias, 250 m, 15 NYL20483,G).
Oct 1984, Buck 12408 (NY). AMAMBAY:
ParqueNa-
cional Cerro Cora, along trail up Cerro Muralla, 300 Thallussquamulose,squamulesroundor elon-
m, 19 Oct 1984, Buck 12507 (NY). CENTRAL: Asun-gate and digitate-lobulate, 0.1-0.5 mm wide,
ci6n, Villa Morra, 14 Aug 1893, Malme s.n., Lich.
Regnell. 1471 (S, UPS), 157 (S). PARAGUARi:
complanateto convex, adnate to somewhat as-
Parque
Nacional Ybycui, trail along Arroyo Mina, ca. 200 m, cending,Pd- or Pd+ orange.Upper surfacegla-
5 & 7 Oct 1984, Buck 11981 (NY). SAN LORENZO: brous, pubescent and fibrillose at the margin.
Ciudad Universitaria, 1 Nov 1984, Bordas 152A (NY). Isidia rare to abundant,globose to cylindrical.
Without prov.: Colonia Risso, 28 Sep 1893, Malme Cortextype 1, 1-2, or 2, 25-45 ,im thick, con-
s.n., Lich. Regnell. 1864F (S), 7 Oct 1893, Malme s.n.,
Lich. Regnell. 1897A, 1897B (S), 20 Oct 1893, Malme tainingcrystals.Prothallusscant, pale to red. Li-
s.n., Lich. Regnell. 1944B (S); Gran Chaco on the Rio chen substances:atranorin,argopsin,norargop-
Negro, 14 Sep 1893, Malme s.n., Lich. Regnell. s.n. sin, vicanicin, norvicanicin, phyllopsorin,
(S). chlorophyllopsorin, parvifoliin, zeorin, addi-
KENYA (Representative specimens). RIFTVALLEY
PROV.:Kajiado, Ngong Hills, 2360 m, 3 Jan 1971,
tional unknownsor none detected.
Moberg1411 (UPS). WESTERN PROV.: Near Yala Riv- Apotheciacommon. Disc planeto convex with
er, 5-7 km S of Kakamega Forest Station, 1500-1700 age, orange-brownto dark red. Margin slightly
m, 21 Jan 1970, R. Santesson 21810 (UPS). raised,concolorousor darkerthan the disc. Ex-
MADAGASCAR. ANTANARIVEPROV.: Ambohi-
ciple tan, commonly red pigmentedat the mar-
manga, 1550 m, 17 Apr 1984, Aptroot 12506 (Hb.
Aptroot). gin, KOH- or the pigmentreactingKOH+ pur-
TANZANIA (Representativespecimens). ARUSHA plish-brown. Hypothecium tan or partly red
PROV.:Mt. Meru, E slope, road to the crater,2000 m, pigmented, KOH- or the pigment reacting
17 Jan 1970, R. Santesson21490 (UPS).KILIMANJARO KOH+ purplishbrown.Hymeniumtan with oc-
PROV.: Mt. Kilimanjaro,W slope, 1700 m, 21 Jan 1970,
R. Santesson20978 (UPS). TANGA PROV.:Usambara
casional red-brownpigmentation,to 45 ,m tall.
Mts., 1894,Holst 3328 (G);Amani,900 m, 9 Jan 1971, Epihymeniumnot obvious. Ascosporessimple or
Moberg1481 (UPS). uniseptate,ellipsoid to short fusiform, 6.5-13.0
x 2.0-3.5 um.
Discussion. This species may prove to be a Pycnidia common, tan, yellow-brownor or-
heterogeneous group. Individuals with the above ange-brown,partlyimmersedin the thallus. Co-
morphology are united by the characters: small nidia rod-shaped, straight, 8.0-11.0 x 0.5-1.0
thalline elements without erect isidia, a thin up- /um.
per cortex, medium ascospore size and no de- Discussion.This species is distinguishedby its
tectable chemistry. The shape of the squamules small to medium-size squamules,abundantisi-
is very variable as is pigmentation in the apo- dia and orange-brownor reddishapothecia.Six
thecia. These differences may prove to be im- varietiesare recognizedbased on chemistrycor-
portant and need further investigation. This spe- relatedwith some morphologicalvariation.

38 Flora Neotropica
Key to Varieties of Phyllopsoracorallina

la. ThallusPd-.
2a. Thalluscontainingphyllopsorin. ........... ........................ ......... var. rappiana.
2b. Thallus lackingphyllopsorin. ................................................. var. corallina.
Ib. Thallus Pd+.
3a. Isidia elongate,marginal,flattened,horizontal;containingonly argopsin. ....... var. phaeobyssina.
3b. Isidia globose to cylindrical,laminal or marginal,erect, or isidia lacking.
4a. Thalluscontainingphyllopsorinor chlorophyllopsorin. .................... var. ochroxantha.
4b. Thalluslackingabove compounds.
5a. Thallus containingvicanicin, argopsinonly in trace quantities;isidia marginal,globose
or bulbous based ..................................................... var. glaucella.
5b. Thallus containingargopsinas a major component;isidia mostly laminal, thin and cy-
lindrical ........................... .................................. var. santensis.
6a. Phyllopsora corallina (Eschweiler) Muller ArroyoNegro,ca. 4 km SEof La GranPiedra,ca. 1000

Argoviensis var. corallina. Fig. e. m, 5 Apr 1982, Harris 14400 (HAC, NY).
ST. EUSTATIUS. On the rim of the Quill Volcano,
Lecideaparvifolia(Persoon) Miiller Argoviensis var. 500 m, 2 Aug 1980, Sipman 14953 (B).
corallinaTuckerman,Proc.Amer.Acad.Arts6: 273. VENEZUELA. LARA:Sierra Portuguesa, Parque
1864. Phyllopsoraparvifolia(Persoon) Muller Ar- Nacional"Yacambfi,"1400-1900 m, 20-21 May 1978,
goviensis var. corallina(Tuckerman)Merrill,Bryol- L6pez-Figueiras15974 (MERF). MERIDA: Valley of
ogist30: 85. 1927, hom. illeg.,non (Eschweiler)Bou- the Rio Chama, 1600 m, 21 Jan 1979, Sipman 10984
ly de Lesdain, 1914. Type. Cuba. Wrights.n., Lich. (B, MERF).MIRANDA: El Volcfn, above Baruta,1200-
Cub. 184e(lectotypedesignatedby Swinscow& Krog, 1300 m, 1 May 1986, Brako 8659, 8671, 8672 (NY);
1981, FH-TUCK 2923, no lichen substances;iso- along road from Hoya de la Puertato San Jos6 de los
lectotypes,BM, G, L, M, UPS). Altos, 1100-1250 m, 9 May 1986, Brako 8694 (NY).
LecideadensifloraeVainio, Bot. Mag. (Tokyo)35: 67. GUYANA. Rupununi, savanna between Takutu
1921. Phyllopsoradensiflorae(Vainio)G. Schneider, River and KanukuMts., Smith 3340 p.p. (FH, U).
Biblioth.Lich. 13: 175. 1979. Type. Japan.Kozuke, FRENCH GUIANA. Cayenne, Montagne du Ma-
25 Feb 1918, Yasuda 350 (holotype, TUR-VAIN hury, 10 km SE of Cayenne,100 m, Mar 1985, Aptroot
22632, no lichen substances). 15481 (U). Saul, 2 km SW of the village, "sentierLi-
monade," 180-210 m, 18 Jul 1986, Montfoort& Ek
Distribution (Fig. 16). Pantropical, 500- 110 (U); 1859, Leprieur509 (G).
1900 m. ECUADOR.NAPO:Tena, alongthe Rio Misahualli,
700 m, 8 Aug 1987, Kalb&A. Kalb 17569 (Hb. Kalb);
Specimens examined. UNITED STATES (Repre- betweenTenaandPuertoMisahualli,9 Aug 1987,Kalb
sentative specimens). ALABAMA: Monroe Co., Brook- & A. Kalb 17084 (Hb. Kalb). TUNGURAHUA:
South slope
lyn, 1853, Beaumont188 (FH-TUCK2828). FLORIDA:of Mt. Tungurahua,N ofBafios, 1850 m, 16 Aug 1987,
AlachuaCo., Gainesville,Ravenel11 (BM);Duval Co.; Kalb & A. Kalb 17566 (Hb. Kalb).
St. Johns Co., 6 Mile Creek,Calkins93 (G, NY=2x). PERU. HUANUCO:Ca. 18 km NNE of Tingo Maria,
GEORGIA: De Kalb Co., Stone Mt., Ravenel s.n. (BM, PuentePumahuasi,20 Feb 1981, R. Santesson& Mo-
FH-TUCK 2821). ILLINOIS:Menard Co., Athens, 10 berg P38:11 (S). SAN MARTIN:Tarapoto, NE of Hotel
Feb 1880, Hall s.n. (BM, FH-TUCK 2821). LouI- de Turistas,350 m, 12 Mar 1981, R. Santesson& Thor
SIANA:East Baton Rouge Parish, Burden Research P71:53, P71:92 (S).
Plantation,W of EssenLane,SE BatonRouge, 19 Mar BRAZIL. BAHIA:Serra das Mangabeiras, ca. 8 km
1974, Tucker12019A(M).MISSOURI: ButlerCo., Ned- before Seabra, 1050 m, 17 Jul 1980, Kalb 290 (Hb.
yville, Nov 1898, Russell 90 (NY). NORTHCAROLINA: Kalb); ca. 10 km N of Rui Barbosa, 400 m, 18 Jul
PolkCo., Tryon,20 Jan 1928,Evans412 (UPS).SOUTH 1980, Kalb293 (Hb. Kalb);ChapadaDiamantina,be-
CAROLINA:
Aiken Co., Aiken, Jan 1885, Ravenel s.n. tween MundoNovo and Morrodo Chapeu,800 m, 20
(BM=2x); BERKELEY CO., Santee Canal, Ravenel s.n. Jul 1980, Kalb295 (Hb. Kalb).MATOGROSSO: Ca. 35
(BM, F, UPS, US). TEXAS:Liberty Co., 14 m E of km SE of Cuiaba, 120 m, 3-4 Jul 1980, Kalb269 p.p.
Cleveland, 9 Jan 1976, Egan EL-7762 (O). VIRGINIA: (Hb. Kalb);Santo Ant6nio de Leverger,ca. 40 km S
SussexCo., TowerHill, Jun 1854, Tuckermans.n. (FH- of Cuiaba,100 m, 5 Jul 1980, Kalb275 p.p. (Hb. Kalb);
TUCK 2821). Serrados Coroados,6 km SW of Buriti, 600 m, 8 Jul
MEXICO. MICHOACAN:
Cerro Azul, Morelia, 10 Mar 1980, Kalb280 (Hb. Kalb);ca. 10 km NE of Chapada
1890, Arsene 3802 (COLO,H, S, UPS). NUEVO LEON: dos Guimaraes, 680 m, 9 Jul 1980, Kalb 282 (Hb.
Sierrade la Silla, Monterrey,10 Jun 1889, Pringle40 Kalb);betweenSao Vicenteand AguasQuentes,90 km
(FH=4x). VERACRUZ:
Cordoba, 1876, Duby s.n. (G); ESEof Cuiaba,650 m, 2 Jul 1980,Kalb286 (Hb. Kalb);
Veracruz,520 m, 17 Mar 1960, Hale & Soderstrom between Santo Ant6nio and Coxip6s, 22 Dec 1893,
19841 (US). Malme s.n., Lich. Regnell. 2170C, 2178C (S). MATO
CUBA.ORIENTE: ElYunque,Mt. Baracoa,Mar 1908, GRosso DOSUL:Estrada do Pantanal, E of Coxim, 300
Underwood & Earle 1004 (NY). SANTIAGODE CUBA: m, 26 Jun 1980, Kalb259 (Hb. Kalb);ca. 30 km S of

Campo Grande, 550 m, 14 Nov 1979, Kalb & Plobst Specimens examined. UNITED STATES. FLORIDA:
266, Lich. Neotropici 343 (M, US). MINASGERAIS: Dade Co., Coconut Grove, 1897, Thaxter 177 (FH);
1885, Glazious.n. (G, PC);Serrado Espinhaco,Serra Seminole Co., Oviedo, Mena s.n. (FH = 2 x).
do Caraca,8 Jul 1978, Kalb & Plobsts.n. (Hb. Kalb). DOMINICAN REPUBLIC.PEDERNALES: LasAbe-
Rio DEJANEIRO:1878, Glaziou s.n. (G), 1885, Vainio jas, S slope of Sierra de Baoruca, ca. 40 km N of Caba
s.n., Lich.Bras.Exs. 145 (BM,M, TUR-VAIN-22606). Rojo, 1750 m, 26 Jan 1987, Buck 14757 (NY).
Rio GRANDE DOSUL:SantoAngelo,7 Feb 1893,Malme PUERTO RICO. PONCE:Cordillera Central, above
s.n., Lich. Regnell. 1013 (BM, G, H, US, W); ColoniaVillalba, Donia Juana Recreation Area, 800-1000 m,
Ijui, 30 Mar 1893, Malme s.n., Lich. Regnell. 1227B 2 Jun 1988, Buck 15983, Harris 22028 (NY).
(S). SANTACATARINA: 13 km above Timb6 do Sul on MAYAGUEZ: Indiera Fria, near Maricao, 430-800 m,
roadto SerraRochina,650 m, 11 Jan 1987, Falkenberg 19-22 Feb 1915, Britton et al. 4398 (NY).
& Brako 3655B (NY). SAo PAULO: FazendaSao Joao BRAZIL. ESPiRITOSANTO:20 km E of Linhares, 30
near Rio Claro, 600 m, 16 Aug 1980, Kalb 302 p.p. m, 25 Jul 1980, Kalb 300. (Hb. Kalb). GoIAs: Between
(Hb. Kalb);Apiahy (Apiai),21 Sep 1878, Puiggaris.n. Jatai and EstAncia, 510 m, 11 Jul 1980, Kalb 283 p.p.
(G);Mun. EldoradoPaulista,Cavernado Diablo, 200 (Hb. Kalb).SAo PAULO: FazendaSao Joao on the Rio
m, 29 Sep 1984, Vital& Buck 12495 (NY, SP). Claro,600 m, 16 Aug 1980, Kalb302 p.p. (Hb. Kalb).
ARGENTINA. MISIONES:Cataracts of Iguazfu, 150 PARAGUAY. ALTOPARANA:Centro Florestal Alto
m, 29 Apr 1979, Kalb 143 (Hb. Kalb). Parana, 4.5 km W of Pto. Pte. Stroessner on Ruta 7,
AUSTRALIA. QUEENSLAND:
Conway State Forest, ca. 250 m, 15 Oct 1984, Buck 12360 (NY).
18 km ENE of Proserpine,180 m, 28 Jun 1986, Strei- ARGENTINA.CORRIENTES: Pase de la Patria,Ar-
mann 37394 (B, CANB) royo San Juan, 9 Sep 1985, Ferraro& Nash 3220 p.p.
SOUTH AFRICA. CAPEPROV.:Div. Albany (near (CTES).
Grahamstown),Fern Kloof, 25 Nov 1953, Almborn
10814 (LD). Discussion. This variety differs from other va-
rieties of P. corallina by its chemistry of ? atra-
Discussion. Variety corallina has a variable norin, vicanicin and traces of norvicanicin and
morphology with atranorin as the only lichen argopsin. The squamules are 0.1-0.3 mm diam.
substance detected. The type specimen has well with a type 2 cortex, 25 gm thick. The isidia are
developed squamules 0.1-0.5 mm diam., a type primarily marginal, globose to short cylindrical
1 cortex 30 ,m thick, and both laminal and mar- and commonly bulbous at the base. The asco-
ginal cylindrical isidia. Isidia are also formed spores are 7.0-8.5-11.0 x 2.5-2.7-3.0 ,m.
from very tiny squamules. Additional specimens The original description says "non isidiosis,"
but the type collection is clearly isidiate. Mis-
may have only this latter form so the thallus is
completely dominated by isidia. The ascospores quoted in Zahlbruckner's catalogue (1926-1927)
are 7.5-9.5-11.5 x 2.0-2.5-3.0 ,m. as f. glaucina Vainio, it is closest to var. ochro-
xantha in morphology.
Variety corallina is distinguished from all oth-
er varieties of Phyllopsora corallina by its chem-
6c. Phyllopsora corallina (Eschweiler) Muller
istry of ? atranorin and non-flattened isidia. In
morphology it is closest to P. isidiotyla which is Argoviensis var. ochroxantha (Nylander)
distinguished by its finely branched isidia and Brako, Mycotaxon 35(1): 13. 1989. Fig. If.
the abundant pale hyphae extending from the Lecidea ochroxanthaNylander, Ann. Sci. Nat. Bot.,
exciple. ser. 4, 11: 223. 1859. Phyllopsoraochroxantha(Ny-
lander)Zahlbruckner,Cat.lich. univ. 10:377. 1939.
Type. Bolivia. Camplolicans,[1847,] Weddells.n.
6b. Phyllopsora corallina (Eschweiler) Muller (lectotypedesignatedby Swinscow& Krog, 1981, H-
NYL 20489, atranorin,phyllopsorin,chlorophyllop-
Argoviensis var. glaucella (Vainio) Brako, My- sorin, isolectotypes,H, PC).
cotaxon 35(1): 13. 1989. LecideasubvirescensNylander,Ann. Sci. Nat. Bot., ser.
Lecidea breviusculaNylander var. glaucella Vainio, 5, 7: 321. 1867. Phyllopsorasubvirescens(Nylander)
G. Schneider,Biblioth. Lich. 13: 179. 1979. Type.
DanskBot. Arkiv4(11):21. 1926. Phyllopsorabrevi- Colombia. Rio Negro, 1200 m, 1863, Lindig s.n.
uscula(Nylander)MullerArgoviensisvar. glaucella
(holotype, H-NYL 20492, phyllopsorin, chloro-
(Vainio)Zahlbruckner,Cat.lich. univ. 10:426. 1939 phyllopsorin).
"fo. glaucina."Type. Mexico. Veracruz:Mirador, Lecidea ernstianaMullerArgoviensis,Flora 60: 473.
Liebmann s.n., PI. Mex. 7381a (holotype, TUR- 1877. Psora ernstiana(MullerArgoviensis)Miiller
VAIN 34026, argopsin,vicanicin, norvicanicin).
Argoviensis,Flora70: 320. 1887. Phyllopsoraernsti-
Distribution (Fig. 16). Mexico, West Indies, ana (MullerArgoviensis)Muller Argoviensis, Bot.
Jahrb.Syst.20:265. 1894.Type.Venezuela.Distrito
South America, and southern North America, Federal:Caracas,Ernst 190 (holotype, G, phyllop-
250-1750 m. sorin, chlorophyllopsorin).

40 Flora Neotropica
Psora polydactylaMiiller Argoviensis,Flora 70: 320. nulosa. The type specimen of L. polydactyla has
1887. Phyllopsorapolydactyla(MiillerArgoviensis) longlobes with abundantmarginal,elongateisid-
Zahlbruckner,Cat. lich. univ. 4: 400. 1926. Type. ia. This seems to be an extreme form. The type
Brazil.Sao Paulo: Apiahy (Apiai), Apr 1882, Puig-
gari 2156 (holotype,G, argopsin,phyllopsorin,chlo- ofPhyllopsora martinii has a type 2 cortex. Thal-
rophyllopsorin). lus morphologymay be similarto var. santensis,
Lecidea spinulosa Vainio, Acta Soc. Fauna Fl. Fenn. but var. oxchroxanthahas slightly smaller as-
2: 46. 1890. Phyllopsoraspinulosa (Vainio) Zahl-
cospores than var. santensis.
bruckner,Cat. lich. univ. 4: 401. 1926. Type. Brazil.
Minas Gerais: Sitio, 1885, Vainio s.n., Lich. Bras.
Exs.993 (holotype,TUR-VAIN22627;isotypes,BM Chemical StrainI
= 2 x, phyllopsorin,chlorophyllopsorin,M, TUR-
VAIN 22626, ZT). Specimensexamined.UNITED STATES. FLORIDA:
LecideaglabriusculaNylander,Ser.lich. trop.40. 1891. Faxon s.n. (FH).
Phyllopsora glabriuscula (Nylander) Swinscow & MEXICO. MORELOS: Cuernavaca, 6500 ft, 31 Oct
Krog,Lichenologist13:241. 1981. Type. Cuba.[Re- 1908, Pringle 10742 (COLO).SANLUISPOTOSi: Ta-
tiro, Mar 15,] Wrights.n., Lich. Cub. ser. 2, 105 masopo, 15 Jun 1892, Pringle36A (FH = 2x).
(holotype,H-NYL 20534, unknowns;isotype, FH- GUATEMALA. PETEN:Near Tikal, 300 m, 9-10
TUCK 2922). Jan 1979, Kalb & Plobst 73 p.p. (Hb. Kalb).
PhyllopsoracinerellaZahlbrucknerin H. Magnusson COSTA RICA. PUNTARENAS:
Monteverde, 1450 m,
& Zahlbruckner,Ark. Bot. 3IA(6): 18. 1944. Type. 4 Jan 1979, Sipman 12098 (B).
U.S.A. Hawaii: Kauai, near Robinson's summer- PANAMA: COCLE: Near El Valle de Anton, 550 m,
house, 1000 m, Feb 1910, Faurie308 (lectotypehere 25-26 Nov 1985, Brako 8428 (NY).
designated,PC; syntypes, Faurie 307, UPS, phyl- DECUBA:La Gran Piedra, 1.5 km
CUBA. SANTIAGO
lopsorin, chlorophyllopsorin,Maui, 350 m, Faurie SE of peak,ca. 1000 m, 4 Apr 1982, Buck7620 (HAC,
539, n.v.). NY), 2 Apr 1982, Harris 14181 (HAC, NY), 3 Apr
PhyllopsoracinerellaZahlbrucknerf. virescensZahl- 1982, Harris 14282, 14285, 14290 (HAC, NY), 4 Apr
brucknerin H. Magnusson & Zahlbruckner,Ark. 1982, Harris 14350, 14351, 14359 (HAC,NY), 4 Apr
Bot. 31A(6): 19. 1944. Type. U.S.A. Hawaii:Oahu,
1982, ca. 1 km W of peak, 1000 m, 6 Apr 1982, Harris
near Honolulu, 400 m, Faurie 413 (lectotype here 14523 (HAC, NY); "Monte Kentucky,"ca. 5 km SE
designated, PC, phyllopsorin, chlorophyllopsorin; of La Gran Piedra, 1000 m, 5 Apr 1982, Buck 7684
isolectotype,BM). (HAC,NY), Harris14410 (HAC,NY). Withoutprov.:
PhyllopsoramartiniiSwinscow& Krog, Lichenologist La Prenda,Dec 1921, Hioram 5306 (US); Wrights.n.,
13: 232. 1981. Type. Kenya. Coast Prov.: Kwale Lich. Cub. ser. 2, 727 (FH-TUCK 2919, G, H-NYL
District, Shimba Hills, Kivumoni Forest, 350 m, 20532).
1972, Krog& SwinscowK 42/3 (holotypeBM, phyl- DOMINICAN REPUBLIC. PEDERNALES:
Las Abe-
lopsorin,chlorophyllopsorin; isotypes,BM, O, UPS). jas, S slope of Sierrade Baoruco,ca. 40 km N of Cabo
Distribution (Fig. 16). Pantropical, sea level- Rojo, 1750 m, 26 Jan 1987, Harris20794 (NY).
COLOMBIA. 1869, Lindigs.n. (PC).
2000 m. VENEZUELA.ARAGUA: 6.5 km from Colonia To-
Discussion. Variety ochroxantha is distin- var along the La Victoria-Tovar road, 6 Apr 1986,
guished by the presence of phyllopsorin or chlo- Brako 8607 (NY). BOLiVAR: ParqueNacional Canai-
rophyllopsorin. The full chemistry is best de- ma, on road from AeropuertoLuepa to Kavanayen,
10 Apr 1985, Brako 8176 (NY), 11 Apr 1985, Brako
tected in solvent B. The squamules are 0.3-1.0 8181 (NY, VEN). DISTRITO
FEDERAL:
Parque Nacional
mm diam. and the isidia are often very long, El Avila, Boca de Tigre,Jun 1985, Brako8189 (NY);
cylindrical and mostly marginal. The ascospores Caracas, 1900 m, 13 Jan 1979, Sipman 10687 (B,
are 6.5-8.5-12.0 x 2.5-3.0-3.5 uim. MERF).MERIDA: Road from Estanquezto Canagua,
2200 m, 13 Dec 1984, Brako 7934A (NY); La Car-
Chemical strains: Chem. strain I: atranorin,
bonera,2100-2200 m, 17 Dec 1984,Brako8124, 8129,
phyllopsorin, chlorophyllopsorin, methyl-phyl- 8131, 8132, 8160 (MERF, NY); along the El Morro
lopsorate, methyl-chlorophyllopsorate, and road, 2000 m, 18 Feb 1976, L6pez-Figueiras& Keogh
?zeorin. Chem. strain II: phyllopsorin, -+chlo- 12362 (MERF);MountZerpa,nearLaHechicera,2000
rophyllopsorin and ?zeorin. Chem. strain III: m, 3 Mar 1978, L6pez-Figueiras& Keogh 15495
(MERF).MIRANDA: Above El Hatillo, 27 May 1986,
chlorophyllopsorin and an unknown compound. Berrys.n. (NY); El Volcin, above Baruta,1200-1400
The type specimen has Chem. strain I. The m, 4 May 1986, Brako 8691 (NY).
thallus morphology is variable. The isidia are SURINAM. Nickerie, area of the Kabalebo Dam
especially variable in occurrence and morphol- project,line W from roadkm 34, 2 Nov 1981, Bekker
1203a (U), 100-200 m, 30 Oct 1981, Zielman 1364
ogy. The type specimen has small lobes with only (U).
few cylindrical isidia. It is identical in morphol- FRENCHGUIANA.Saul,primaryforestnearRoche
ogy and chemistry with the type of Lecidea spi- Bateau,200-300 m, Mar 1985, Aptroot15365 (U), 2

km SW of the village, "sentierLimonade," 180-210 m, 6 Feb 1985, Sipman & Aptroot18459 (B);Mt. La-
m, 22 Jul 1986,Montfoort&Ek 114, (U), 20 Aug 1986, tipu, ca. 8 km N of Kamarang,ca. 600 m, 24 Feb 1985,
Montfoort& Ek 137, 138 (U). Sipman & Aptroot18998 (B).
PERU. SAN MARTIN:Lamas, Cerro Blanco (ca. 63 ECUADOR. PASTAZA:Mera, at cemetery, 1100 m,
km W-WNW of Tarapoto),1200 m, 17 Mar 1981, R. 26 Nov 1972, Arvidsson& Nilson 414 (GB).
Santesson& ThorP77:18 (S). BRAZIL.BAHIA: Itanagra,8 km W ofItanagraalong
BRAZIL. GOIAS:Between Jatai and Estancia,510 road to Subauima,50 m, 27 May 1981, Boom & Mori
m, 11 Jul 1980, Kalb 283 p.p. (Hb. Kalb). MINAS 978 (NY). MINASGERAIS: Vila Monte Verde, 30 km
GERAIS: Tiradentesnear Sao Joao del Rei, 900 m, 6 E of Camanducaia,1500 m, 7-11 Sep 1978, Kalb &
Jul 1978,Kalb&Plobst29 (Hb. Kalb);Lafayette,1885, Plobst 44 p.p. (Hb. Kalb). PARA:Serrado Cachimbo,
Vainios.n., Lich. Bras.Exs. 275 (TUR-VAIN 22600), 774 km N of Cuiaba,400 m, 22 Apr 1983, Brako &
338 (BM, TUR-VAIN 22601); Caraca, 1885, Vainio Dibben 5523 (INPA, NY); Aeroporto Cachimbo, 20
s.n., Lich. Bras.Exs. 1356 (TUR-VAIN 22628), 1443 km N of the borderwith Mato Grosso, 430-480 m,
(TUR-VAIN 22629), s.n. (TUR-VAIN 22630). PARA: 27 Apr 1983,Brako&Dibben6141, 6168, 6170 (INPA,
Serrado Cachimbo,842 km N of Cuiaba,ca. 350-500 NY); 842 km N of Cuiaba,350-500 m, 5 May 1983,
m, 5 May 1983, Brako & Dibben 6758 (INPA, NY); Brako & Dibben 6748 (INPA, NY); cataractson the
Rio Jamanxim, 974-1024 km N of Cuiabd, 300 m, Rio Curua,877 km N of Cuiaba,350-500 m, 2 May
10-15 May 1983, Brako&Dibben7006A(INPA,NY); 1983, Brako&Dibben6527 (INPA,NY), 7 May 1983,
SerraMaze, 1208-1229 km N of Cuiaba, 100-200 m, Brako & Dibben6909 (INPA, NY). PARANA: Reserva
18-22 May 1983, Brako & Dibben7331 (INPA, NY). Ecol6gicaSapitanduva,between Morretesand Anto-
Rio DEJANEIRO:Glaziou 1938B (M). SAOPAULO:Api- nina, 100 m, 17-18 Jan 1987, Hatschbach& Brako
ahy (Apiai), 1890, Puiggari1721 (G);Camposdo Jor- 8873 (NY). Rio GRANDEDOSUL:Hamburgerberg, near
dao, 1700 m, 25 May 1978, Kalb&Plobst20 p.p. (Hb. Sao Leopoldo, 18 Nov 1892, Malme s.n., Lich. Reg-
Kalb); near Cachoeirade Emas, 15 km NE of Pira- nell. 599 (S, UPS). SANTACATARINA:
Between Uribici
cununga,550 m, 14 Jun 1979, Kalb& Plobst 145 (Hb. and CapivaraAlta on road to Braqodo Norte, 1040
Kalb);Serrado Itapeti,betweenMogi das Cruzesand m, 12 Jan 1987, Falkenberg& Brako 3731A (NY).
Aruja, 500 m, 20 Apr 1980, Kalb 232 (Hb. Kalb); URUGUAY. TACUAREMBO:
Rinc6n da Vissoura, 16
MorroGrandenearCotia, 25 km W of Sao Paulo, 850 Dec 1965, San Martins.n. (Hb. Osorio 4737).
m, 27 Sep 1980, Kalb309 p.p. (Hb. Kalb);Ilha de Sao
Sebastiao(Ihla Bela), ca. 250 m, 6 Jul 1979, Kalb & Chemical Strain III
Poelt s.n. p.p. (GZU).
PARAGUAY. PARAGUARI:
Parque Nacional Yby- Specimensexamined.UNITED STATES. FLORIDA:
cui, trailalongArroyoMina, ca. 200 m, 5-6 Oct 1984, 27 Dec 1898, Patillo 165 (FLAS).
Buck 11935 (NY). MEXICO. OAXACA: In hills between Arroyo Hu-
San Cosme, Paso de la
ARGENTINA. CORRIENTES: maca and Rio Verde,300 m, 20 Apr 1985, Thomaset
Patria,ArroyoSan Juan, 9 Sep 1985, Ferraro& Nash al. 3574 (NY).
3220 p.p. (CTES). COSTA RICA. PUNTARENAS:
Near La Cruzes Gar-
den, ca. 4 km SSE of San Vito, 1330 m, 1 Jan 1979,
Chemical Strain II Sipman 11989 (B).
PANAMA. BOCADELTORO:Fortuna Dam region,
Specimens examined. COSTA RICA. PUNTARENAS: 1000 m, 8 Dec 1985, McPherson7847 (NY). COCLE:
Valle de Coto Brus,ca. 350 m, 30 Dec 1978, Hafellner Near El Valle de Anton, 550 m, 25-26 Nov 1985,
7434 (GZU). Brako8417, 8421, 8430 (NY). PANAMA:Along trailto
PANAMA. DARIEN:W ridge of Cerro Tacarcuna CerroBrewster,670 m, 18-21 Nov 1985, Brako8256
massif, 4100-4800 ft, 26 Jan 1975, Mori & Gentry (NY);alongEl Llano-Cartiroad,400 m, 24 Nov 1985,
4420 (US). Brako 8386 (NY); along the CerroJefe-Alto Pacoura
CUBA. SANTIAGO DE CUBA: La Gran Piedra, NE road, 28 Nov 1985, Brako 8499 (NY).
slope, 1100m, 7 Apr 1982,Harris14564, 14644 (HAC, BAHAMAS. ANDROS: Nicholl's Town and vicinity,
NY); Arroyo Negra, ca. 4 km SE of La Gran Piedra, 13-15 Mar 1907, Brace 6881 (NY).
1000 m, 5 Apr 1982, Harris14445 (HAC, NY). With- CUBA. ISLEOF PINES:San Juan, 15, 17 Mar 1916,
out prov.: Wrights.n., Lich. Cub. ser. 2, 726 (H-NYL Britton et al. 15586 (FH, NY). ORIENTE:Bayate, Ek-
20533). man 37 (S, TUR-VAIN 22611);NoguerasHill, 27 Oct
JAMAICA. 1884. Hart 104 (FH-TUCK 2923). 1921, Hioram 5690 (US).
DOMINICAN REPUBLIC. INDEPENDENCIA:
Sierra DOMINICAN REPUBLIC. PEDERNALES:
Las Abe-
de Baoruco, 25.5 km S of Puerto Escondito at inter- jas, 55 km N of portofCabo Rojo on Alcoa road,3600
section of road to CharcoColorado, 1750 m, 14 Jan ft, 5 May 1982, Harris 15383 (NY).
1987, Buck 14509 (NY). VENEZUELA. AMAZONAS: Rio Negro, Cerro de La
VENEZUELA. MERIDA:Finca "Buenos Aires," near Neblina, along the Rio Mawarinuma,just outside Ca-
Aricagua,1700 m, 8 Mar 1976, L6pez-Figueiras12649 nionGrande, 140 m, 5 Feb 1985, Buck 12827 (NY,
(MERF).MIRANDA: El Volcan, above Baruta, 1200- VEN).MIRANDA: El Volcan,above Baruta,1200-1400
1400 m, 1 May 1986, Brako 8663 (NY). m, 4 May 1976, Brako 8682 (NY).
GUYANA. Upper Mazaruini, Jawalla village, at TRINIDAD. Rio Grande Forest, S of Tamana, 5
confluenceof Kukuiriver and Mazaruniriver,ca. 500 May 1957, Fleming s.n. (NY).

42 Flora Neotropica
SURINAM. Paramaribo,50 m, Narain 144 (B). VENEZUELA.AMAZONAS: San Carlos,Spruces.n.,

FRENCHGUIANA.Saiil,primaryforestnearRoche Lich. Amaz. And. 225 (BM).
Bateau,200-300 m, Mar 1985, Aptroot15417 (U), 2 GUYANA. Upper Mazaruni,Mt. Latipu, ca. 8 km
km SW of the village, "sentierLimonade," 180-210 N of Kamarang,24 Feb 1985, Sipman&Aptroot18982
m, 4 Jul 1986, Montfoort& Ek 102 (U), 18 Jul 1986, (B).
Montfoort& Ek 109, 111 (U), 22 Jul 1986, Montfoort SURINAM. Pleike village at Hawa River, 10 Nov
& Ek 112 (U), 3 Sep 1986, Montfoort& Ek 159 (U), 1971, MacGillavrys.n. (U)
21 Sep 1986, Montfoort& Ek 174, 177 (U), 24 Sep FRENCHGUIANA.Saiil,primaryforestnearRoche
1983,Montfoort&Ek 183 (U), 30 Sep 1986, Montfoort Bateau,200-300 m, Mar 1985, Aptroot15420 (U).
& Ek 191 (U). ECUADOR.NAPO:Tena, alongthe Rio Misahualli,
ECUADOR. NAPO:Santa Rosa, km 5 North, 400 700 m, 8 Aug 1987, Kalb&A. Kalb17568, 17570 (Hb.
m, 20 Aug 1982, Aptroot& Hensen 10650 (Hb. Ap- Kalb).
troot);Auca Oil Fields, 56 km S of Coca, 17 Jun 1983, BRAZIL.SAOPAULO: Serrade Boissocanga,above
Brako 7459, 7460, 7463A (NY, QCA). PICHINCHA:
Ca. Maresias,330 m, 18 Feb 1980, Kalb 212 (Hb. Kalb);
18 km NW of San Miguel de los Bancos, 800 m, 27- FazendaSaoJoaonearRio Claro,600 m, 16 Aug 1980,
29 Dec 1983, Buck 10334 (NY). Kalb 302 p.p. (Hb. Kalb).
PERU. SAN MARTIN:Lamas, Cerro Blanco (ca. 63
km W-WNWof Tarapoto),1200 m, 17 Mar 1981, R. Discussion. This variety is distinguished by its
Santesson& ThorP77:23 (S). medium-size squamules with abundant, flat-
BRAZIL. PARA: Serra do Cachimbo, Aeroporto tened isidia and its unusual chemistry of only
Cachimbo,20 km N of the Mato Grossoborder,430-
480 m, 27 Apr 1983, Brako & Dibben 6169 (INPA, argopsin. The spores are 6.0-7.3-10.0 x 2.5-
NY); 842 km N of Cuiaba, 350-500 m, 5 May 1983, 2.9-3.5 /um. It most closely resembles Phyllop-
Brako & Dibben 6752 (INPA, NY); Rio Jamanxim, sora corallina var. ochroxantha which has cylin-
974-1024 km N of Cuiaba,300 m, 10-15 May 1983, drical isidia.
Brako & Dibben 7080 (INPA, NY); SerraMaze and
vicinity, 1208-1229 km N ofCuiaba, 100-200 m, 18- 6e.
22 May 1983, Brako & Dibben 7267 (INPA, NY). Phyllopsora corallina (Eschweiler) Muller
PARANA:Prainhas, near Porto de Cima, 100 m, 16 Jan Argoviensis var. rappiana Brako, var. nov.
1987, Hatschbach&Brako8839 (NY);ReservaEcol6-
gicaSapitanduva,betweenMorretesandAntonina,100 P. corallinae var. santensi similis sed atranorino
m, 17-18 Jan 1987, Hatschbach& Brako 8856 (NY). et parvifoliino diversa. Type. United States.
RORAIMA:Alto Alegre, Ilha de Maraca, 9 Jun 1986, Florida: Sarasota Co., Myakka River State Park,
Rodrigueset al. 847 (INPA, NY), 12 Jun 1986, Ro- 16 Aug 1985, Brako 8229 (holotype, NY).
drigueset al. 933 (INPA,NY). Sio PAULO: Ilhade Sao
Sebastiao,200 m, 6 Jul 1979,Kalb155, 157 (Hb.Kalb), Distribution (Fig. 16). Central America, South
+ 1-500 m, 6 Jul 1979, Kalb & Poelt s.n. (GZU);Fa-
zenda Sao Joao near Rio Claro,600 m, 16 Aug 1980, America, and southern North America, sea level-
Kalb302 p.p. (Hb. Kalb);PraiaGrande,S of Sao Paulo, 850 m.
2 m, 18 Dec 1978, Kalb & Hannack 59 (Hb. Kalb).
AUSTRALIA. QUEENSLAND:Trinity Bay, 1886, Specimens examined. UNITED STATES. FLORIDA:
Miillers.n. (G). SarasotaCo., MyakkaRiver State Park, 16 Aug 1985,
Brako8223 (NY); SeminoleCo., Sanford,9 Apr 1923,
Rapps.n. (FLAS).
6d. Phyllopsora corallina (Eschweiler) Miller MEXICO. VERACRUZ: Cordoba, [1876], Duby s.n.
Argoviensis var. phaeobyssina (Vainio) Brako, (US).
Mycotaxon 35(1): 14. 1989. Fig. 5c, d.
GUATEMALA. IZABAL: Quirigua,Jul 1976, Hale
50881, 50974 (US = 2x).
Lecidea breviusculaNylander var. phaeobyssinaVai- HONDURAS.CoPAN:Copin, Jul 1976,Hale 50964
nio, Ann. Acad. Sci. Fenn., ser. A, 6(7): 127. 1915. (US).MORAZAN: Suyatillo,northof ElZamorano,800-
Phyllopsora breviuscula(Nylander) Miiller Argo- 850 m, 18 Jul 1949, Standley 21360 (F). OLANCHIO:
viensisvar.phaeobyssina(Vainio)Zahlbruckner, Cat. BetweenJuticalpaand El Asilo, 380-400 m, 10 Mar
lich. univ. 4: 396. 1926. Type: Guadeloupe.Houel- 1949, Standley 17763 (F); between Juticalpaand La
mont, [1897,] Duss 481 (holotype, TUR-VAIN Concepci6n,400 m, 12 Mar 1949, Standley17916 (F).
22602, argopsin;isotype, NY). Near bridge on Rio
COSTA RICA. PUNTARENAS:
Brujo,450 m, 2 Jan 1979, Sipman 12061 (B).
Distribution (Fig. 16). West Indies and South GUYANA. Rupununi, savanna between Takutu
America, 200-2000 m. River and the KanukuMts, 12-22 Mar 1938, Smith
3340 (NY).
Specimens examined. PUERTO RICO. PONCE:Cor- BRAZIL.MATOGROSSO: Ca. 35 km SE of Cuiaba,
dilleraCentral,above Villalba,Doia JuanaRecreation 120 m, 3-4 Jul 1980, Kalb 269 p.p. (Hb. Kalb).
Area, 800-1000 m, 2 Jun 1988, Harris21987 (NY).
ST. VINCENT. RichmondPeak, 1000-2000 m, El- Discussion. This variety is distinguished by its
liott 261 p.p. (TUR-VAIN 22612A). chemistry of atranorin and parvifoliin. It is clos-

est to var. santensis in morphology. The asco- enels.n., ReliquiaeTuckermanianae15 (B, BM, COLO,
spores are 6.5-8.0-10.0 x 2.5-3.0-3.5 gm. F, FH, L, NY = 2x, O, S, US). TEXAS:Jasper Co., 6
mi S of Everdale,19 Apr 1951, Whitehouse25059 (US).
COSTA RICA. PUNTARENAS: Valle de Coto Brus,
6f. Phyllopsora corallina (Eschweiler) Miiller 350 m, 30 Dec 1978, Hafellner 7465, 7477 (GZU);
Argoviensis var. santensis (Tuckerman) Brako, Valle General,near Terraba,ca. 250 m, 2 Jan 1979,
Mycotaxon 35(1): 14. 1989. Fig. 5a, b. Hafellner6514 (GZU); RestaurantRio Brujo, ca. 10
km from Rio Grandede Terraba,350 m, 2 Jan 1979,
LecideasantensisTuckerman,Amer. J. Sci. Arts, ser. Kalb & Plobst 71 (Hb. Kalb);Monteverde, 1450 m, 4
2,25:428. 1858. Phyllopsorasantensis(Tuckerman) Jan 1979, Sipman 12088 (B, NY).
Swinscow& Krog,Lichenologist13:236. 1981.Type. BAHAMAS. NEW PROVIDENCE: Waterloo, 12-24
U.S.A. SouthCarolina:[BerkeleyCo.,] SanteeCanal, Mar 1907, E. Britton6633 (NY, US).
1849, Ravenel 182 (holotype,FH-TUCK 2822, ar- DOMINICAN REPUBLIC. PEDERNALES: Las Abe-
gopsin, norargopsin). jas, S slope of Sierrade Baoruco, 40 km N of Cabo
PhyllopsoraalbicansMiillerArgoviensis,Bull.Soc.Roy. Rojo, 1750 m, 26 Jan 1987, Harris 20779 (NY).
Bot. Belgique32: 132. 1893. Type. Costa Rica. Car- COLOMBIA. VAUPES: Rio Apaporis, Mar 1951,
tago: Terraba(Turrialba),1893, Tonduzs.n. (holo- Schultes11753 (FH), s.n. (NY).
type, G; isotypes, US = 2 x, argopsin,norargopsin, SURINAM. Paramaribo,50 m, 1976,Narain55 (B).
as Pittier & Durands.n., PI. Costar.Exs. 5474). FRENCH GUIANA. Saiil, in and near the village,
Lecidea porphyromelaenaVainio, Ann. Acad. Sci. 200 m, Mar 1985, Aptroot15155 (U), 2 km SW of the
Fenn., ser. A, 15(6): 113. 1921. Phyllopsoraporphy- village,"sentierLimonade,"180-210 m, 20 Aug 1986,
romelaena(Vainio)Zahlbruckner,Cat. lich. univ. 4: Montfoort& Ek 136 (U).
401. 1926. Type. Philippines.Luzon:BataanProv., ECUADOR. NAPO:Aiangu, 70 km E of Coca on
Mount Marivales, Dec 1908, Merrills.n., Bur. Sci the Rio Napo, 1 Mar 1983, Brako 5311 (NY, QCA).
6273 (lectotype designated by Swinscow & Krog, PERU. SAN MARTIN:Tarapoto, NE of Hotel Turis-
1981, TUR-VAIN 22619; syntype, Bur. Sci. 6256, tas, 350-500 m, 11 Mar 1981, R. Santesson & Thor
TUR-VAIN 22620, argopsin,norargopsin;isosyn- P70:47 (S), 16 Mar 1981, R. Santesson & ThorP76:
types, BM, US). 14, P76:18, P76:19 (S).
Lecidea miradorensisVainio, Dansk Bot. Ark. 4(11): BRAZIL. AMAZONAS: 0.5 km E of Borba, 22 Jun
22. 1926. Phyllopsora miradorensis(Vainio) G. 1983, Nelson et al. 1311 (INPA, NY). MATOGROSSO:
Schneider,Biblioth.Lich. 13:177. 1979. Type.Mex- Mun. Terezinha,SerraCobrinha,ca. 10 km W of BR-
ico. [Veracruz:]Mirador, 18 Mar 1842, Liebmann 158 and 17 km N of junction of BR-158 and MT-413,
s.n., P1. Mex. 7373 (lectotype designatedby Swin- 14 Oct 1985, Thomas et al. 4392 (INPA, NY); San
scow & Krog, 1981, TUR-VAIN 34034; isolecto- Ant6nio do Leverger,40 km S of Cuiaba,100 m, 5 Jul
type, FH; syntype,Liebmanns.n., P1.Mex. 7372a, 1980, Kalb274 (Hb. Kalb);Chapadados Guimaraes,
TUR-VAIN 34035, argopsin,norargopsin,reported 800 m, 6 July 1980, Kalb 277 p.p. (Hb. Kalb);Serra
by Swinscow& Krog, 1981). dos Coroados,Buriti,600 m, 8 Jul 1980, Kalb280 p.p.
Phyllopsoraformosana Zahlbruckner,Repert. Spec. (Hb. Kalb); Santa Anna da Chapada, 2 Mar 1894,
Nov. Regni.Veg. 33: 43. 1933. Type. Taiwan.Rais- Malme s.n., Lich. Regnell. 2481 (S). MATOGROSSODO
ha, 5 Jan 1925, Asahinas.n. (holotype,W, argopsin, SUL:Estradado Pantanal,E of Coxim, 270 m, 29 Jun
norargopsin,reportedby Swinscow& Krog, 1981). 1980, Kalb256 p.p. (Hb. Kalb),300 m, Kalb259 (Hb.
Lecidea corallina Eschweiler in Martius f. saxicola Kalb). PARA:Serrado Cachimbo,Base Aerea do Ca-
Malme, Ark. Bot. 28A(7): 47. 1936. Type. Brazil. chimbo, 430-480 m, 26 Apr 1983, Brako & Dibben
Mato Grosso:Santo Ant6nio near Cuiaba,Morrin- 6036 (INPA, NY). PARANA:Guaira, 200 m, 9 Aug
ho, 24 Apr 1894, Malme s.n., Lich. Regnell.2607B 1980, Kalb 301 p.p. (Hb. Kalb). SAo PAULO: Serrade
(holotype,S, argopsin,norargopsin). Paranapiacaba,40 km SW of Sao Paulo, 800 m, 19
Mar 1978, Kalb & Plobst 1 p.p. (Hb. Kalb);Praia de
Distribution (Fig. 16). Pantropical, sea level- Peruibe near Itanhaem, 1 m, 23 Aug 1978, Kalb &
1750 m. Plbst 47, 132 (Hb. Kalb); Ilha da Cananeia, 16 Jul
1979, Kalb 168 (Hb. Kalb);Ilha Comprida,sea level,
15-16 Jul 1979, Sipman 14088 (B).
PARAGUAY.CENTRAL: Villa Elisa, ca. 20 km S of
Chemical Strain I Asunci6n, 15 Oct-10 Nov 1947, Olrogs.n. (UPS).
PHILIPPINES LUZON: Bontoc Prov., Nov-Dec
Specimens examined. UNITED STATES (Repre- 1910, Vanoverbergh,Bur. Sci. 1023 (TUR-VAIN
sentative specimens). ALABAMA:Baldwin Co., Fish 22621).
River, 29 Nov 1924, Evans 133 (FH, NY, US). FLORI-
DA: Alachua Co., Gainesville, 14 Mar 1938, Murrill
s.n. (NY), SarasotaCo., MyakkaRiver State Park, 7
May 1967, Harris 2614 (MIN), 16 Aug 1985, Brako Chemical Strain II
8224, (NY). GEORGIA: Camden Co., ca. 5 mi S of
WoodbinealongUS Route 17, 28 Feb 1975, Egan EL- Specimensexamined.UNITED STATES. FLORIDA:
6751 (0). LOUISIANA: St. Tammany Parish, Abita, 27 Lake Co., Lake Norris, 6 May 1921, Kelly s.n. (US);
Nov. 1891, Langlois 847 (US). SOUTHCAROLINA: Rav- SeminoleCo., Sanford,Nov 1910,Rapp89 (FH = 2 x).

44 Flora Neotropica
ILLINOIS:MenardCo., Athens, 1878, Halls.n. (S).Mis- Distribution (Fig. 18). Guatemala, Paraguay
sissippi: 1845, Dr. Vetschs.n. (FH-TUCK 2822). and Brazil, 100-300 m.
JAMAICA.Mandeville,28 Jan 1909, Wight10 (FH).
PERU LORETO:Iquitos, Explorama Lodge, Lake Specimens examined. GUATEMALA. ALTAVER-
Trail, 100 m, 23 Jan 1981, R. SantessonP7:19 (S). APAZ:Chama-Chichoob,1000 ft, 23 Jul 1920, Johnson
355 (US). PETEN: Near Tikal, 300 m, 9-10 Jan 1979,
Discussion. This variety differs from var. cor- Kalb & Plobst 73 p.p. (Hb. Kalb).
allina by its chemistry of ?atranorin, argopsin, BRAZIL. AMAZONAS: Along road to Balbina Hy-
+norargopsin and zeorin. Variety santensis gen- droelectricDam ProjectfromManaus-Caracarai Road,
8 & 11 Aug 1979, Buck 2709A (INPA, NY). MATO
erally has well developed squamules with nu- GRosso: Sao Vicente, 20 Oct 1974, Freire46 (INPA,
merous globose to cylindrical, erect, isidia. The
NY); ca. 35 km SE of Cuiaba, 120 m, 3-4 Jul 1980,
apothecial margin is usually darker than the disc. Kalb269 p.p. (Hb. Kalb);Santo Ant6nio de Leverger,
The ascospores are 7.0-10.3-13.0 x 2.5-2.8-3.0 ca. 40 km S of Cuiaba, 100 m, 5 Jul 1980, Kalb 275
A.m. (Hb. Kalb);Cuiaba,25 May 1894, Malme s.n., Lich.
Regnell.s.n. (S). RORAIMA: Alto Alegre,Ilha de Mara-
Most easily distinguished by their chemistries,
ca, 11 Jun 1986, Rodrigues 908 (INPA, NY). SXo
var. santensis and var. ochroxantha are often PAULO:Salto Grande de Paranapanema,Jul 1901,
similar in morphology. Variety santensis usually Wettstein& Schiffners.n. (W).
has abundant laminal, globose isidia where var. PARAGUAY. ALTO PARANA:Colonia 13 Tuyuti, 31
ochroxantha usually has mostly marginal, elon- km N of Herandarias on CarreteraAkataete,ca. 250
m, 14 Oct. 1984, Buck 12350 (NY).
gate-cylindrical isidia. The spores in var. santen-
sis tend to be slightly longer than those of var. Discussion. The large squamules remind one
ochroxantha. Collections distributed as Reliqui- of the P. parvifolia group, but the species is dis-
ae Tuckermanianae no. 15, are often mixed col- tinguished by its rough surface (Fig. 19b), deeply
lections of var. santensis and var. corallina. incised lobes and abundant isidia. All herbarium
Chemical strains: Chem. strain I: +atranorin, specimens have white squamules. Malme (1936)
argopsin, norargopsin, and ?zeorin. Chem. strain confused small squamule forms of P. cuyabensis
II: argopsin. with P. furfuracea, which is distinguished by its
chemistry of furfuracein.
7. Phyllopsora cuyabensis (Malme) Zahlbruck-
ner, Cat. lich. univ. 10: 377. 1939. 8. Phyllopsora fendleri (Tuckerman & Mon-
Fig. 19a, b.
tagne) Miller Argoviensis, Bot. Jahrb. Syst.
LecideacuyabensisMalme,Ark.Bot. 28A(7):48. 1936. 20: 264. 1894. Fig. 19c.
Type.Brazil.MatoGrosso:Serrada Chapada,Buriti,
26 Jun 1894, Malme s.n., Lich. Regnell. s.n. (holo- BiatorafendleriTuckerman& Montagnein Montagne,
type, S, no lichen substances;possibleisotype,UPS). Ann. Sci. Nat. Bot., s6r. 4, 8: 296. 1857. Lecidea
fendleri (Tuckerman& Montagne)Nylander, Ann.
Thallus squamulose, squamules round to ir- Sci. Nat. Bot., s6r. 4, 19: 339. 1863, s6r. 4, 20: 262.
regular or elongate, lobes deeply incised, 0.3-1.0 1863. Type. VenezuelaFendlers.n. (holotype,FH-
mm wide, thick and convex, adjoined and over- TUCK 2923, no lichen substances;isotype, H-NYL
20523).
lapping, adnate to ascending, Pd-. Upper sur-
face rough and fibrillose, chalky white; margin Thallus squamulose, squamules short-lobed,
with dense fine white fibrils. Isidia common, glo- 0.5-1.0 mm wide, strongly convex, adjoined and
bose or branched. Cortex type 2, with an irregular overlapping, adnate, Pd-. Upper surface gla-
upper surface, 15-25 Am thick. Prothallus pale, brous, pubescent and fibrillose at the margin.
thick. Lichen substances: +atranorin. Isidia lacking. Cortex type 1-2, 50-60 um thick.
Apothecia common. Disc convex, tan. Margin Prothallus abundant, pale. Lichen substances:
slightly raised, tan. Exciple and hypothecium none detected.
yellowish, KOH+ slightly brighter. Hymenium Apotheciacommon. Disc plane to convex, tan
colorless, to 50 ,um thick. Epihymenium color- to brown. Marginslightly raised, paler than the
less. Ascospores simple, ellipsoid to short fusi- disc with white fibrils.Exciple tan, KOH-. Hy-
form, 6.5-8.3-13.0 x 2.5-2.8-3.0 ,m. pothecium yellow-brown, KOH-. Hymenium
Pycnidia abundant, tan, ostiole darker, partly tan, to 55 ,m tall. Epihymeniumnot obvious.
immersed in the thallus. Conidia rod-shaped, Ascospores simple, ellipsoid to short fusiform,
straight, 7.0-9.0 x 0.5-1.0 ,m. 9.0-11.3-15.0 x 3.0-3.5-4.0 ,m.

vC~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~*C
'~l~'.f L
AN,~~~~~~
~~~~* ?~~'~l, '.'.l :q
4p
4b
4~
....,
: ~~~~~~~~~~~~~~~~~~~~~~~.-.
I -'
q q~~~~~~~~~~~~~~~~~~~~~q
FIG. 19. a. PhyllopsoracuyabensisHolotype: Brazil, Malme s.n., Lich. Regnell. s.n., S). Habit 12 x. b.
Phyllopsoracuyabensis(Holotype:Brazil,Malme s.n., Lich. Regnell. s.n., S). SEM micrographshowing rough
uppercortex, 50 x. c. Phyllopsorafendleri(Venezuela,M6rida,L6pez-Figueiras& Sipman 18483, MERF),scale
= I mm. d. Phyllopsorafurfuracea (Dominican Republic,Harris 19751, NY) 12x. e. Phyllopsoraintermediella
(Ecuador,Galipagos Islands,Isla SantaCruz, WeberL-40262, COLO)12 x. f. Phyllopsoraisidiotyla(Holotype:
Brazil, Vainios.n., Lich. Bras.Exs. 222, TUR-VAIN 22634) 12 x.

46 Flora Neotropica
Pycnidia common, tan to yellow-brown, os- Thallus squamulose or dominated by isidia,

tiole darker,partially immersed in the thallus. squamulesroundto irregular,ca. 0.1 mm diam.,
Conidia rod-shaped,straight, 15.0-16.0 x 0.5- mostly slightly convex, closely adjoined and
1.0 inm. sometimes appearingas a continuous crust, ad-
Distribution (Fig. 13). Costa Rica, Venezuela nate, Pd-. Upper surfaceglabrose, fibrilloseat
and Brazil, above 1200 m. the margin.Isidiacylindricalto coralloid.Cortex
Specimens examined. COSTA RICA. SAN JOSE:
type 2, 10 ,m thick. Prothallussparseto abun-
Cordillerade Talamanca,ca. 5 km W of Cerrodel la dant,reddish.Lichensubstances:furfuraceinand
Muerte,San Gerardode Dota, 2600 m, 21 Mar 1985, +unknown compounds.
Sipman & Chaverri20806 (B). Apotheciacommon. Disc plane to convex, tan
VENEZUELA. DISTRITOFEDERAL: Caracas, Ernst to red-brown.Marginslightly raisedwith abun-
s.n. (US); trackfrom Altamirato La Silla, 1900 m, 13
Jan 1979, Sipman 10688 (B, MERF).MERIDA: La Car- dant pale fibrils,palerthan the disc. Exciple pale
bonera, 2400 m, 22 Jan 1979, L6pez-Figueiras& Sip- tan to partlyredpigmented,containingan orange
man 18483 (MERF);MorroNegro, nearPico de Hor- crystallinematerialdissolvingand diffusingpale-
ma, 2400 m, 12 Apr 1980,L6pez-Figueiras&Rodrigues yellow to red or purplishin KOH. Hypothecium
22887, 22951 (MERF);Paramo de Mucuchies, 3500 a to
m, Mar 1969, Oberwinkleret al. s.n. (GZU).TRUJILLO:containing yellow-brown orangecrystalline
Paramode Cende,2800-3100 m, 30 Mar 1976,L6pez- material dissolving and diffusingpale-yellow to
Figueiras 13036 (MERF);Paramo de La Nariz, near red or purplish in KOH. Hymenium yellow-
the T.V. tower,2500 m, 28 Mar 1979,L6pez-Figueiras brown, to 60 ,Lm.Epihymeniumyellow-brown,
& Hale 19474 (MERF),2400 m, 24 Jan 1980, L6pez- 5 ,m thick. Ascosporessimple or uniseptate,cy-
Figueiras 22026 (MERF); Las Palmas, 2300 m, 20 lindricalto short
&
Aug 1981,L6pez-Figueiras Rodrigues 26280
(MERF).
fusiform, 6.0-9.0-12.0 x 2.0-
BRAZIL AMAZONAS: Pico Rondon, 1250 m, 4 Feb 2.3-2.5 Am.
1984, Samuels et al. 185 (INPA, NY). SAOPAULO: Pycnidia common, tan to yellow-brown, os-
Campos do Jord3o, 1700 m, 25 May 1978, Kalb & tiole darker.Conidia rod-shaped,straight,8.0-
Pl6bst20 p.p. (Hb. Kalb). 10.0 x 0.5-1.0 ,m.
Discussion. This species is closest to Phyllop- Distribution (Fig. 13). Pantropical, sea level-
sora parvifolia var. breviuscula, but is distin- 2200 m.
guished by its shorter lobes, yellow-brown hy-
pothecium and longer ascosporesand conidia. Specimensexamined.UNITED STATES. FLORIDA:
Dade Co., Hattie Bauer Hammock, EvergladeKeys,
12 Jul 1915, Small et al. 6235 (NY); Seminole Co.,
9. Phyllopsorafurfuracea(Persoon)Zahlbruck- Sanford,Mar 1926, Rapp s.n. (FLAS).
ner in Engler & Prantl, Nat. Pflanzenfam. 1(1*): MEXICO. CHIAPAS:Mun. La Trinitaria, Lagos de
138. 1905. Figs. 2a-d, 10, 19d. MontebelloNational Park,near Cinco Lagunas,1600
m, 14 May 1988, Breedlove& Bourell 67771 (CAS).
Lecideafurfuracea Persoonin Gaudichaud.,Voy. Uran. VERACRUZ: Mirador,Aug 1841,Liebmanns.n., PI.Mex.
192. 1827. Type. "MariannaIsland." Gaudichaud 7381B (TUR-VAIN 34030).
s.n. (lectotype here designated, PC; frag. H-NYL BELIZE. EL CAYO: Valentin, 27 Jun 1936, Mains
20507). 3653 (F, US).
Lecidea haemophaeaNylander, Flora 52: 122. 1869. PANAMA. COLON:Near Fort Davis, 1 m, 23 Nov
Phyllopsorahaemophaea(Nylander)Muller Argo- 1985, Brako8355 (NY).
viensis, Hedwigia34: 141. 1895. Type. Peru. [Lore- CUBA. ISLE OF PINES:La Cunagua, 19 Feb 1918,
to:] Yurimaguas,Spruces.n., Lich. Amaz. And. 185 Brittonet al. 14596 (FH, NY, US = 2x). Santiagode
(holotype,H-NYL 20520; isotypes,BM, furfuracein, Cuba:ArroyoNegro, ca. 4 km SE of La Gran Piedra,
G= 2x). 1000 m, 5 Apr 1982, Harris 14453 (HAC, NY); La
Lecidea schizophylla (Vainio) Malme var. isidiata GranPiedra,ca. 1 km W of peak, 1000 m, 6 Apr 1982,
Malme, Ark. Bot. 28A(7): 44. 1936. Type. Brazil. Harris 14547, 14548 (HAC, NY), NE slope, 1100 m,
Rio Grande do Sul: Colonia Ijuhy, 30 Mar 1893, 7 Apr 1982, Harris 14657 (HAC, NY). Withoutprov.:
Malme s.n., Lich. Regnell. 1226B p.max.p. (holo- La Prenda, Dec 1921, Hioram 5335 (NY, S), 5345,
type, S). 5407 p.p. (US); Wrights.n., Lich. Cub.ser. 2, 119 (FH-
Lecidea hypochryseaVainio, Ann. Acad. Sci. Fenn., TUCK 2922, H-NYL 20579, US).
ser. A., 15(6): 114. 1921. Phyllopsorahypochrysea JAMAICA. PORTLAND PARISH:7 mi NW of Muir-
(Vainio)G. Schneider,Biblioth.Lich. 13:176. 1979. ton on roadto Ecclesdown,alongtrailinto John Crow
Type. Philippines.Mindanao:Subprov.Butuan,320 Mts., 11 Apr 1981, Buck 5646 (NY). Withoutparish:
m, Mar-Jul 1911, Weber 1393 (holotype, TUR- 3 Mar 1905, Cummings37 (COLO= 2x, F, O, NY
VAIN 22622, furfuracein). = 2 x, US = 2 x), Feb & Mar 1905, Cummings49 (FH
= 2x, NY); Hart 59(NY).

DOMINICAN REPUBLIC. INDEPENDENCIA:

Sierra FRENCH GUIANA. Saul, in and near the village,
de Baoruco,25.5 km S of Puerto Escondito, 1750 m, 200 m, Mar 1985, Aptroot15196 (U), primaryforest
24 Jan 1987, Buck 14493 (NY), Harris20471, 20481 nearBoeufMort,200-300 m, Mar 1985,Aptroot15331
(NY); Charcode La Paloma, 48.4 km S of PuertoEs- (U), 2 km SW of the village,"sentierLimonade,"180-
condito, 1800 m, 25 Jan 1987, Buck 14688 (NY), Har- 210 m, 15 Aug 1986, Montfoort& Ek 128 (U), 20 Aug
ris 20677 (NY). EL SEIBO:19 km E of Sabana de La 1986,Montfoort&Ek 139(U), 24 Aug 1986,Montfoort
Mar on the road to Miches, sea level, 28 Mar 1981, & Ek 144, 146 (U), 3 Sep 1986, Montfoort& Ek 160
Buck 5003 (NY). LA VEGA:La Sal, 13.3 km N of El (U), 15 Sep 1986, Montfoort& Ek 173 (U), 21 Sep
Rio then 10 km E of PassssBajitoon roadto Casabito, 1986, Montfoort& Ek 178 (U).
3500-3600 ft, 29 Apr 1982, Harris 15009 (NY); Ar- ECUADOR. ESMERALDAS: San Lorenzo, 30 m, 12
royo Piedrosa,1.2 km from Monabaoon roadto Jara- Jul 1982, Aptroot & Hensen 11026 (Hb. Aptroot).
bacoa, 11 May 1982, Harris 15679 (NY); km 17 on MORONA-SANTIAGO:Los Encuentros, km 12 East Cor-
Jarabacoa-LaVega road, 11 May 1982, Harris 15754 dillera del Condor, 1200 m, 26 Jul 1982, Aptroot&
(NY); "La Cotorra"along trail to Pico Duarte, 5000 Hensen 10450 (Hb. Aptroot);Palora, 900 m, 13 Aug
ft, 16 Jan 1987, Harris20079, 20109 (NY). LA VEGA/ 1982, Aptroot& Hensen 10362 (Hb. Aptroot);Guala-
SANTIAGO: Parque Nacional J. Armando Bermudez, quiza, 900 m, 28 Aug 1982, Aptroot& Hensen 10453,
along trail along Rio Los Guanos near La Cienaga, 10456 (Hb. Aptroot).NAPO:Sta. Rosa, km 5 N, 400
3500-4000 ft, 12 Jan 1987, Harris 19751 (NY). PE- m, 20 Aug 1982, Aptroot& Hensen 10608 (Hb. Ap-
DERNALES: Las Abejas, 55 km N of port of Cabo Rojo troot).Los Rios: Quevedo,km 54,200 m, 15 Jul 1982,
on Alcoa road, 3600 ft, 5 May 1982, Harris 15345 Aptroot& Hensen 11229 (Hb. Aptroot). PICHINCHA:
(NY). Ca. 18 km W of San Miguel de Los Bancos, 800 m,
COLOMBIA: CUNDINAMARCA: Between San Jose 27-29 Dec 1983, Harris 17928 (NY). ZAMO-
and Lago Pedro Paulo (ca. 32 km from Mosquera,on RA-CHINCHIPE:Zamora, along the Rio Zamora, 1200
the Mosquera-LaMesa road), 2000-2250 m, 1 Aug m, 25 Jul 1982, Aptroot& Hensen 10536 (Hb. Ap-
1976, DumontCO-5433, CO-5441(NY);Villeta,2100 troot).
m, 186?, Lindig 840 (H-NYL 20506, H-NYL P.M. PERU. SAN MARTIN:Tarapoto, NE of Hotel de
4116), 2696 (BM, H-NYL 20509, H-NYL P.M. 4107, Turistas,350 m, 11 Mar 1981, R. Santesson & Thor
UPS). RISARALDA: Mun. de Sta. Rosa de Cabal, Vereda P70:82 (S), 12 Mar 1981, R. Santesson & ThorP71:
La Colina, 1725 m, 19 Aug 1986, Wolf 1228 (B). 15, P71:42 (S). Cuzco: Paucartambo,ca. 3 km S of
VENEZUELA. ARAGUA: Rancho Grande, Parque Pillcopata, 750 m, 28 Mar 1981, R. Santesson et al.
NacionalHenriPittier,1200m, Dec 1986,Brako8507A P101:29 (S);ca. 11 km S of Pillcopata,850 m, 28 Mar
(NY), 1080 m, Mar 1969, Oberwinkler et al. s.n. (GZU), 1981, R. Santessonet al. P102:22 (S).
La Mora, W of Colonia Tovar, 1200 m, Dec 1986, BRAZIL.ACRE: Alongthe Rio Iaco, a few km above
Brako 8510, 8517 (NY, VEN); 6.5 km from Colonia Sena Madureira, 4 Oct 1980, Nelson 572 (INPA). MATO
Tovaralongthe La Victoria-Tovar,6 Apr 1986,Brako GROSSO: Sierrados Coroados,6 km SW of Buriti,600
8608, 8612 (NY). BOLIVAR: ParqueNacionalCanaima, m, 8 Jul 1980, Kalb280 p.p. (Hb. Kalb);Buriti, 19 Jan
on roadfromAeropuertoLuepato Kavanayen,10 Apr 1894, Malme s.n., Lich. Regnell. 2268B (S), 20 Jun
1985,Brako8165 (NY). CARABOBO: Alongthe Rio San 1894, Malmes.n., Lich. Regnell.s.n. (S), 21 Jun 1894,
Gian betweenthe top of the electricalplant and 2 km Maimes.n., Lich.Regnell.s.n. (S).MINASGERAIS: Ser-
below the plant, 350-550 m, 2 Apr 1966, Steyermark ra do Espinhaco, Serra do Caraca, 1200 m, 8 Jul 1978,
& Steyermark 95589 (NY). DISTRITOFEDERAL:Ca- Kalb&Plobst31 p.p. (Hb. Kalb), 1500 m, 12 Jul 1978,
racas,Ernst 143 (G); trackfrom Altamirato La Silla, Kalb & Plobst 34 (Hb. Kalb); Vila Monte Verde, ca.
1900 m, 13 Jan 1979, Sipman 10690 (B, MERF). 30 km E of Camanducaia,1500 m, 7-11 Sep 1978,
MERIDA:Road from Estanquezto Canagua,2200 m, Kalb & Plobst 44 p.p. (Hb. Kalb), 2 Jul 1979, Kalb &
13 Dec 1984, Brako 7943 (NY); La Carbonera,2100- Plobst 150 (Hb. Kalb), 1600 m, 2 Jul 1979, Sipman
2200 m, 17 Dec 1984, Brako8111, 8117, 8130A, 8158 12959 (B); Fazenda Sao Mateus E of Camanducaia,
(NY), 2300 m, 22 Jan 1979,Sipman&L6pez-Figueiras 1800 m, 30 Nov 1980, Kalb315 p.p. (Hb. Kalb);Sitio,
11029 (B, MERF);El Maciegal,near Merida, 1800- 1885, Vainio s.n., Lich. Bras. Exs. 719 (BM = 2x,
2100 m, 10Jul 1975, L6pez-Figueiras&R. Terdn9931 TUR-VAIN 22658); Caraca, 1895, Vainios.n., Lich.
(MERF);Mount Zerpa,near La Hechicera,2000 m, 3 Bras. Exs. 1451 (BM, TUR-VAIN 22607, ZT), 1469
Mar 1978, L6pez-Figueiras& Keogh 15464, 15481, (TUR-VAIN 22639). PARA:Serra do Cachimbo, 780
15498, 15502 (MERF),28 Dec 1978, L6pez-Figueiras km N of Cuiaba,430-480 m, 29 Apr 1983, Brako &
17289, 17290 (MERF),4 Dec 1980, L6pez-Figueiras Dibben 6428 (INPA, NY). PARANA:Reserva Ecol6gia
25003 (MERF);La Mucuy,above Tabay NE Merida, SapitanduvabetweenMorretesand Antonina, 100 m,
2200-2500 m, Mar 1969, Oberwinkler et al. s.n. (GZU). 17-18 Jan 1987,Hatschbach&Brako8862, 8865, 8878,
MIRANDA:El Volcan, above Baruta, 1200-1400 m, 1 8892 (MBM, NY). Rio DE JANEIRO:1878, Glaziou
May 1986,Brako8655, 8657, 8664, 8665B, 8666 (NY), 1938 (G, NY); Itatiaia,22 Jul 1978, Kalb& Plobst53,
4 May 1986, Brako 8677, 8680 (NY) NUEVAESPARTA: Lich.Neotropici341 (M,US);Corcovado,15 Aug 1892,
Isla Margarita,CerroMatasiete, 11 Apr 1986, Brako Malme s.n., Lich. Regnell.82, 83 (S). Rio GRANDE DO
8622, 8628, 8630, 8631A (NY). SUL: S of Torreson the road to CampingItapeba, 10
GUYANA. East Demerara,Timehri,OakaraCreek, m, 9 Jan 1987, Falkenberg& Brako 3603, 3612B,
10 m, 2 Feb 1985, Sipman&Aptroot17966 (B).Upper (FLOR, NY); Pirating near Pelatas, 19 Dec 1892,
Mazaruni,Mt. Latipu, ca. 8 km N of Kamarang,ca. Malme s.n., Lich. Regnell. 851 (S); 20 km S of Julio
600 m, 24 Feb 1985, Sipman & Aptroot19003 (B). de Castilhos,Taquarembo,550 m, 13 Nov 1989, Oso-

48 Flora Neotropica
rio & Fleig 89/139 (Hb. Osorio). RONDONIA: 120 km irregular,shiny and adnate to the substrate(Fig.
SW of Porto Velho, 20 km NW of Mibrasa,25 May 2a, b). The prothallusis pale and web-like, apo-
1982, McFarland 266 (INPA, NY). SANTACATARINA:
SerraRio do Rastro, ca. 12 km W of Bom Jardimda theciaand pycnidiaarecommon. The squamules
Serra,1470 m, 27 Sep 1984, Vital& Buck 12361 (NY, expand and produceisidia which are globose at
SP); between Uribi9i and CapivaraAlta on road to onset and become cylindricalwith age (Fig. 2c).
Bravodo Norte, 1040 m, 12 Jan 1987, Falkenberg& Whenyoungthe surfaceof eachisidiumis smooth
Brako 3722 (NY). Sio PAULO: Campos do Jordao, and shiny, as they elongate, the isidia may be-
1700 m, 25 May 1978, Kalb & Plobst20 p.p., 22 (Hb.
Kalb), 1520 m, 25 Jan 1987, Brako 8933 (NY); Serra come swollen. With age, the isidia deflate, ap-
de Paranapiacaba,80 km SW of Sao Paulo, 700 m, 24 pearingirregularin shape and with a rough sur-
Mar 1978, Kalb& Plobst5 p.p. (Hb. Kalb);60 km SW face (Fig. 2d);the prothallusdarkensand appears
of Sao Pauloabove Juquitiba,660-800 m, 13-14 May carbonized(Fig. 19d). The apothecia found on
1978, Kalb & Plobst 18 (Hb. Kalb), 550 m, 27 Apr the darkenedprothallusare old, often growing
1980, Kalb233 (Hb. Kalb);Serrado PeruibenearAna
Dias, 120 km SW of Sao Paulo, 50 m, 2 Apr 1978, from the hymenia of expired apothecia. The
Kalb& Plobst 9 p.p., 58 (Hb. Kalb);betweenTaubat6 young,healthyapotheciacontainan orangecrys-
and Ubatuba, 800 m, 18 Jun 1978, Kalb & Plobst 26 talline materialwhich readilydissolves and dif-
p.p. (Hb. Kalb);Serrada Cantareira,ca. 30 km N of fuses red or purplishin KOH. Older apothecia
Sao Paulo, 950 m, 5 Aug 1978, Kalb & Hannack 42
(Hb. Kalb);Serrado Garraozinho,between Moji das may have orange crystalline material, but this
Cruzesand Bertioga,850 m, 30 Jun 1979, Kalb 148 dissolves and diffusespale yellow in KOH.
(Hb. Kalb), Sipman 12675, 12721 (B), 29 Mar 1980,
Kalb 227 p.p. (Hb. Kalb);Ilha da Cananeia,5 m, 16
Jul 1979, Kalb 166 (Hb. Kalb); 3 km E of Botukatu, 10. Phyllopsoraglabella (Nylander)G. Schnei-
850 m, 11 Nov 1979, Kalb & Plobst 197 p.p. (Hb. der, Biblioth. Lich. 13: 176. 1979.
Kalb);Serrade Botukatu,Pardinho,800 m, 11 Nov Figs. 9b.
1979, Kalb&Plobst198 p.p. (Hb. Kalb);Serrade Bois-
socangaabove Maresias,30 km W of Sao Sebastiao, LecideaglabellaNylander,Sert.lich.trop.37. 1891.
330 m, 18 Feb 1980, Kalb 212 (Hb. Kalb); Morro Type.Cuba.Wright s.n.,Lich.Cub.ser.2, 142(ho-
GrandenearCotia, 850 m, 27 Sep 1980, Kalb309 p.p. lotype,H-NYL20534,nolichensubstances; isotype,
(Hb. Kalb); Ilha Comprida, 2 m, 1 Nov 1980, Kalb FH-TUCK2922).Phyllopsora microsperma Muller
313 p.p. (Hb. Kalb); Eldorado Paulista, Cavera do Argoviensis,Bull.Herb.Boissier89. 1894.Type.
Diablo, 200 m, 29 Sep 1984, Vital&Buck 12505 (NY, Mexico.Jalisco,1890,sinecoll.[misit]Eckfeldt190
SP). (holotype,G).
IVORY COAST. NANANE: N'zo (near N'Zerekore, Lecidea Malme,Ark.Bot.28A(7):41. 1936.
subglabella
base of Mt. Nimba), 550 m, 1951, Des Abbayess.n. Phyllopsora (Malme)G. Schneider,
subglabella Bib-
(UPS). lioth.Lich.13:179.1979.Type.Brazil.MatoGros-
KENYA.COASTPROV.: KwaleDistrict,ShimaHills, so: "Guiapr.Cuyaba,"14 May 1894,Malmes.n.,
350 m, 1-2 Sep 1985, Kalb & Schrigl 13235 (Hb. Lich. Regnell.2547 (holotype,S, no lichen sub-
Kalb). stances;isotype,UPS).
TANZANIA. ARUSHA PROv.: Mt. Meru, E slope,
roadto crater,2100-2200 m, 7 Jan 1971, R. Santesson Thallus squamulose, squamules round to ir-
23030 (S). regularlylobed, 0.2-0.35 mm diam., convex, ad-
PROV.:Perinet = An-
MADAGASCAR. TAMATAVE
joined and overlapping,adnate,Pd-. Uppersur-
dasibe,950 m, 10 May 1984, Aptroot& Hensen 13342
(Hb. Aptroot).
faceglabrous,shortfinepale fibrilsat the margin.
JAVA. Junghuhns.n., LichenJavan N. 38 (L). Isidia lacking. Cortextype 1-2, to 60 ,m thick,
AUSTRALIA. QUEENSLAND: Mt. Spec State Forest, containinggranules.Prothallusscant,paleto red-
Paluma Range, 6 km W of Paluma, 920 m, 18 Jun dish. Lichen substances:none detected.
1986, Streimann 36972, 37019 (CANB). NEW CALE-
DONIA.Jun 1886, Saves 21 (M). Apothecia common. Disc convex, yellow-
brown. Margin slightly raised, darkerthan the
Discussion. This species is distinguished by its disc. Exciple yellow-brown,partlyred pigment-
small insidiate squamules, apothecia with a yel- ed. Hypothecium yellow-brown, KOH-. Hy-
low-brown to orange crystalline material, me- menium colorless, to 50 j,m. Epihymeniumnot
dium size ascospores and furfuracein. It is closest obvious. Ascosporessimple, ovoid, 4.5-5.0-6.5
to Phyllopsora corallina var. corallina, but differs x 2.5-2.8-3.5 um.
by its smaller squamules and chemistry. Pycnidia common, yellow-brown, immersed
The thallus morphology varies with age. In in the thallus. Conidiarod-shaped,straight,6.0-
young thalli, the squamules are small, round to 9.0 x 0.5-1.0 ,m.

Distribution(Fig. 13).Mexico,Honduras,Cuba Specimens examined. COSTA RICA. CARTAGO:

and Brazil, 100-500 m. Turrialba,650 m, 21-22 Feb 1986, Berry4566, 4567,
4568, 4569, 4570 (NY).
Specimens examined. MEXICO. SAN LUIS POTOSi: CUBA. ORIENTE:Bayate, 4 Apr 1917, Ekman s.n.
Las Palmas, 300 ft, 6 Jun 1890, Pringle s.n. (COLO, (S).
FH = 3 x, UPS, US). JAMAICA. TRELAWNY PARISH:Cockpit Country,
HONDURAS. OLANCO:Vic. Juticalpa, 380-480 m, 500 m, 18 Apr 1981, Buck 5883 (NY).
5-16 Mar 1949, Standley 17941 (F). DOMINICAN REPUBLIC.PEDERNALES: 6 km N
CUBA. PINARDELRIO: San Crist6bal, Wright s.n., of Los Arroyos,5200 ft, 6 May 1982, Buck8305 (NY);
Lich. Cub.ser. 2, 102 (FH-TUCK2922). Las Abejas,S slope of Sierrade Baoruco,ca. 40 km N
of Cabo Rojo, 1750 m, 26 Jan 1987, Buck 24720,
Discussion.This species is distinguishedby its 24728 (NY).
medium-sized,thick squamules,small ovoid as- VENEZUELA.AMAZONAS: Rio Negro,base of Cer-
ro de la Neblina, 140 m, 25-28 Nov 1984, Brako7531
cosporesand lack of lichen substances.It is clos-
est to Phyllopsoraparvifoliellawhich differs in (NY, VEN), 23 Feb 1984, Buck 11104 (NY, VEN).
MERIDA: La Carbonera,2100-2200 m, 17 Dec 1984,
having a thinner,type 1 cortex, abundantisidia, Brako8100, 8123 (NY, MERF),28 Jun 1978, Lopez-
and a trace of what may prove to be argopsin. Figueiras 16238 (MERF). MIRANDA:Along road from
Hoya de la Puertato San Jose de los Altos, 1100-1250
m, 9 May 1986, Brako 8694B, 8696 (NY). TACHIRA:
Near Betania, 2400 m, 30 Jul 1975, L6pez-Figueiras
11. Phyllopsoraintermediella(Nylander)Zahl- 10014 (MERF).
FRENCH GUIANA. Saul, 2 km SW of the village,
bruckner,Cat. lich. univ. 4: 398. 1926. "sentierLimonade,"180-210 m, 15 Jul 1986, Mont-
Figs. 6c, 19e. foort &Ek 107 (U), 19 Aug 1986, Montfoort&Ek 133,
20 Aug 1986, Montfoort& Ek 135 (U), 22 Aug
Lecidea intermediellaNylander, Ann. Sci. Nat. Bot., (U),
ser. 4, 19: 339. 1863. Psora intermediella(Nylander) 1986, Montfoort& Ek 140, (U), 27 Aug 1986, Mont-
foort & Ek 150, 151 (U), 29 Aug 1986, Montfoort&
MillerArgoviensis, Flora71:528.1888.Type.Cuba. Ek 154
(U), 30 Sep 1986, Montfoort& Ek 192, 193,
[Oriente:Lomo del Gato,] Wrights.n. (holotype,H- 196
NYL 20558; isotypes,as Wrights.n., Lich.Cub. 183, (U), 1 Oct 1986, Montfoort& Ek 199, 200 (U).
ECUADOR. GALAPAGOS ISLANDS:Santa Cruz Is-
BM = 2x, FH-TUCK 2921, G, L = 2x, M, UPS,
no lichensubstances, land, vic. Academy Bay, 21 Jan-9 Mar 1964, Weber
W). L-40262 (COLO,FH, UPS, US); FloreanaIsland,trail
Thallus squamulose, squamules granularor from Black Beach to Highlands,25 Apr 1976, Weber
& Lanier L-62909 (COLO). NAPO:25-31 km E of
elongate-flabellate,0.1-0.3 mm diam., mostly Papallactaon Quito-Baezaroad,2000-2250 m, 22 Feb
discrete,less frequentlyadjoinedor overlapping, 1985, Arvidssonet al. 6684 (GB).
adnate,complanateto convex, Pd-. Upper sur- PERU. LORETO: Iquitos, ExploramaLodge, ca. 50
faceglabrous,shortfibrilloseat the margin.Isidia km NE of Iquitos, 100 m, 24 Jan 1981, R. Santesson
P9:8 (UPS).
variable, irregular,globose, cylindrical with a BRAZIL. PARA:Serra do cataractson
bulbousbase or slightlyflattened,oftenbranched, the Rio Curua, 877 km N ofCachimbo, Cuiaba, 350-500 m, 2
to 1 mm long, erect. Cortex a thin gelatinous May 1983, Brako & Dibben6614a (INPA, NY). SXo
layer,to 20 ,im thick, extendingto the lower side PAULO: EldoradoPaulista,Cavera do Diablo, ca. 200
of the squamules.Prothallusabundant,reddish. m, 29 Sep 1984, Vital& Buck 12497 (NY, SP).
Lichen substances:none detected.
Apotheciacommon. Disc planeto slightlycon- Discussion. This species is closest to Phyllop-
vex, yellow-brown.Marginslightlyraised,darker sora longiuscula from which it differs by its glo-
than the disc, with abundantred fibrils.Exciple bose or elongate, often swollen isidia, darker
reddish-brown, KOH-. Hypothecium yellow- margined apothecia and shorter ascospores. It is
brown to partly red pigmented, KOH-. Hy- treated in the broad sense. The type material of
menium tan, to 50 ,im tall. Epihymeniumnot P. intermediella has elongate, flattened isidia and
obvious. Ascosporessimple, ovoid or fusiform, ascospores 7-9 x 2-3 ,im. Only a few of the
7.0-9.0 x 2.5-3.5 j,m (type only). specimens cited below are fertile. Buck 24720
Pycnidiacommon, mostly superficial,tan with differs from type material by having more erect
a darker ostiole. Conidia rod-shaped, straight, isidia and ascospores 10-15 x 3-4 ,um. Speci-
12.0-15.0 x 0.5-1.0 ,m. mens from the Galapagos Islands, have larger,
Distribution(Fig. 20). CentralAmerica, West more regular squamules than other collections,
Indies and South America, 100-2400 m. and have isidia that are swollen at the base and

50 Flora Neotropica
:;
-- -
.....- ; --y-
- ..o"
-- 0- "- -3
0
'0 20 30 *0 50 60- . . . . . . . . . :
.01 -.
,'I,G. 20. .-- .'.- -io . .... a), Phyllop-
sora
kalbii andPhylopsora
(squares), longiuscula
(triangles).
conical at the tip, and ascosporesfrom 11-15 x TUR-VAIN 22634, atranorin,zeorin;isotypes, BM

2.5-3.5 Am. Sterile collections from Venezuela = 2 x, M, UPS, ZT).
are close in thallusmorphologyto the Galapagos Thallus squamulose or dominated by isidia,
specimens. Other sterile specimens including squamulesgranularto elongate,ca. 0.1 mm wide,
Brako 8694b, 8696, Lopez-Figueiras 10014 and Pd-. Upper surfacedull to
delicately fibrillose.
Ekman s.n., have very long cylindrical, some- Isidia irregular-shapedand finelybranched.Up-
times branchedisidia. The isidia on Berry4566- per cortex heavily gelatinized, to 10 uimthick.
4570 are very small and crowded, This species Prothallus abundant, pale. Lichen substances:
has been found growingon bark and rock. ?atranorin and ?zeorin.
Apotheciacommon. Disc concave to slightly
12. Phyllopsoraisidiotyla (Vainio) Riddle, My- convex, yellow-brown.Marginraised, pale with
cologia 15: 81. 1923. Fig. 19f. abundantpale projectingfibrils.Exciple yellow-
LecideaisidiotylaVainio,Etud.class.lich.Bresil2:49. brown to pale at the margin, KOH-. Hypothe-
1890.Type.Brazil.MinasGerais:Lafayette,1885, cium yellow-brown to red-brown, KOH- or
1000 m, Vainios.n., Lich. Bras.Exs. 222 (holotype, KOH+ brightening.Hymenium yellow-brown,

to 45 jumtall. Epihymenium yellow-brown to septate, ellipsoid to short fusiform, 6.5-8.9-12.0

red-brown. Ascospores ellipsoid to short fusi- x 2.5-2.9-3.5 ,m.
form, 7.5-10.0-12.5 x 2.0-2.8-3.5 j,m. Pycnidia common, tan, ostiole orange or
Pycnidia not common, tan to yellow-brown, brown, immersed in the thallus. Conidia rod-
superficial.Conidia rod-shaped, straight, 11.0- shaped, straight, 9.0 x 1.0 ,m.
12.0 x 0.5-1.0 um. Distribution (Fig. 20). Pantropical, 300-
Distribution (Fig. 20). Costa Rica, South 2400 m.
America, and southern North America, to Specimens examined. UNITED STATES. LOUI-
3000 m. SIANA: RapidesParish,at intersectionof La. 1 and La.
8 nearZimmerman,22 Apr 1978, Tucker17566 (LSU).
Specimensexamined.UNITED STATES.LOUI- OKLAHOMA: Cherokeecounty,alongTerapinCreekca.
SIANA:NatchitochesParish,KisatchieNationalForest,
6 mi. N of St. Rd. 82 on St. Rd. 100, 23 Apr 1988,
ca. 35 m W-NWof Alexandria,
9 Apr 1978, Tucker Harris 21382
17538 (LSU). TEXAS: (NY).
NacogdochesCo., "Tick Hill," DOMINICAN REPUBLIC. DISTRITONACIONAL:
9 m W of Nacogdoches,27 Aug 1967, Tucker7144a SantoDomingo,JardinBotanicoNacional"Dr.Rafael
(LSU). M. Moscoso," 19 Mar 1981, Buck 4631, 4634 (NY),
COSTA RICA. 1893, Tonduzs.n. (G). Reese 15195A (NY).
VENEZUELA. MERIDA: Paramo de Marinio,near
BRAZIL. MATOGRoSSO:Cuiaba, 2 Dec 1893,
Tovar, 3000 m, 25 Nov 1980, L6pez-Figueiras24739 Malme s.n., Lich. Regnell.2056A, Lich. Austroameri-
(MERF). cani 235a (BM, UPS), 11 Dec 1893, Malme s.n., Lich.
BRAZIL. MINASGERAIS:Vila Monte Verde, 30 km
E of Camanducaia,1800 m, 7-11 Sep 1978, Kalb & Regnell.2097C (S), 3 Jan 1894, Malmes.n., Lich.Reg-
Plobst44 p.p. (Hb. Kalb).Rio GRANDE DOSUL:Santo nell. 2208 (S), 9 Jan 1894, Malme s.n., Lich. Regnell.
near 6 Malme 2231 (BM, FH, S), 2233C, 2234B (S), 25 May 1894,
Angelo, Cachoeira, Jan 1893, s.n., Lich. Malme
s.n., Lich. Regnell. s.n. (S); Coxip6, near Cu-
Regnell. 861X (S), 10 Jan 1893, Malme s.n., Lich. 27 Dec 1893, Malme s.n., Lich. Regnell.2187B,
Regnell.887B (S), 7 Feb 1893, Malme s.n., Lich. Reg- iaba,
nell. 1013 (S);SilveiraMartins,20 Mar 1893, Malme Lich.Austroamericani235b (BM,S, UPS), 9 Feb 1894,
Malme s.n., Lich. Regnell. 2349 (S, UPS), 2349B (S);
s.n., Lich. Regnell. 1208B (S).
Serrada Chapada,Buriti, 19 Jan 1894, Malme s.n.,
Discussion.This species is distinguishedby its Lich.Regnell.2271 (S);SantaAnnade Chapada,4 Feb
1894, Malme s.n., Lich. Regnell. 2339B (S). MATO
granular, finely isidiate thallus and the abundant GROssoDOSUL:Estradado Pantanal,E of Coxim, 300
pale fibrilson the marginof the apothecia. m, 29 Jun 1980, Kalb 258 (Hb. Kalb).
KENYA. CENTRAL PROV.:Nanyuki District, be-
tween Naro Moru and Nanyuki, 2050 m, 17-18 Aug
1985,Kalb&Schr6gl13520, 13551, 13562(Hb. Kalb).
13. Phyllopsorakalbii Brako, sp. nov. TANZANIA.ArushaProv.:Mt. Meru,E slope,2100-
Squamulismediocribusconvexis proximead- 2200 m, 16 Jan 1970, R. Santesson21525 (UPS). KIL-
IMANJARO PROV.:Mt. W slope, Shima
natis, isidiis globoso-moniliformibus,necnon Farm, 1700 m, 12 JanKilimanjaro,1970, R. Santesson 21001A
apotheciis brunneis distincta. Type. Brazil. (UPS);E of LemoshoGlades,ca. 2400 m, 13 Jan 1970,
MatoGrossodo Sul:Estradado Pantanal,some R. Santesson21050 (UPS).
kms E of Coxim, 270 m, 29 Jun 1980, Kalb
Discussion. This species is distinguished by its
256 p.p. (holotype, NY; isotypes, Hb. Kalb,
medium-sized, convex, closely adnate squa-
M). Fig. 3a.
mules, globose-moniliform isidia and brown
Thallussquamulose,squamulesroundto elon- apothecia.
gate, 0.1-0.3 mm diam., discreteto adjoinedand
overlapping,convex, closely adnate, Pd-. Up-
per surface glabrous, fibrillose at the margin. 14. Phyllopsora longiuscula (Nylander) Zahl-
Isidiaglobose,elongatingby budding.Cortextype bruckner, Cat. lich. univ. 4: 398. 1926.
2, 20 jumthick. Prothallusthin, pale. Lichensub- Lecidea
longiusculaNylander,Ann. Sci. Nat. Bot., s6r.
stances:none detected. 4, 19: 339. 1863. Psoralongiuscula(Nylander)Miil-
Apotheciacommon. Disc planeto slightlycon- ler Argoviensis, Flora 65: 483. 1882. Type. Cuba.
vex, pale to dark brown. Marginslightly raised, [Oriente: Monte Verde,] Wright s.n. (holotype,
darker or paler than the disc. Exciple yellow- H-NYL 20537; isotypes, as Lich. Cub. 179, often
mixed with or replacedby Squamacidiajaneirensis
brown, KOH-. Hypothecium yellow-brown, (MullerArgoviensis)Brako,FH-TUCK2919, BUF,
KOH-. Hymenium tan, to 40 ,umtall. Epihy- G = 2 x, H-NYL 20535, L = 2 x, M = 2 x, no lichen
menium not obvious. Ascospores simple or uni- substances,UPS, W).

52 Flora Neotropica
Thallus squamulose, squamules round or elon- fibrilloseat the margin. Isidia lacking. Cortexa
gate-flabellate, 0.1-0.3 mm diam., discrete, ad- thin gelatinous layer, to 10 ,umthick, in part
joined or overlapping, adnate to ascending, com- extendingto the lower surface.Prothallusabun-
planate to convex, Pd-. Upper surface glabrous, dant, red. Lichen substances:none detected.
short fibrillose at the margin. Isidia flattened, Apotheciacommon. Disc plane to convex, tan
incised and often branched. Cortex a thin gelat- to yellow-brown. Margin plane, dark red to
inous layer, to 20 um thick, extending to the brown. Exciple yellow-brown,red pigmentedat
lower side of the squamules. Prothallus abun- the margin,KOH-. Hypotheciumyellow-brown,
dant, reddish. Lichen substances: none detected. KOH-. Hymenium pale yellow-brown, to 45
Apothecia common. Disc plane to slightly con- Am thick. Epihymenium yellow-brown. Asco-
vex, yellow-brown. Margin slightly raised, paler spores simple, short-ellipsoid, 7.0-9.7-12.0 x
than the disc, with abundant red fibrils. Exciple 2.5-2.7-3.0 ,um.
tan, KOH-. Hypothecium yellow-brown, Pycnidianot seen.
KOH-. Hymenium tan, to 50 um tall. Epihy- Distribution.St. Vincent, 1000-2000 m.
menium not obvious. Ascospores simple, ovoid Discussion. Known only from the type speci-
or fusiform, 12.5-15.9-19.0 x 3.0-3.5-4.0 ,m. men, this species is distinguishedby its small,
Pycnidia not seen. elongate and closely adnate thin squamules,its
Distribution (Fig. 20). Cuba, Puerto Rico and medium-size ascospores,and the lack of lichen
Venezuela, 400-1000 m. substances.The type specimen includes 2 spe-
cies. The piece designated"a" is closest to the
Specimens examined. CUBA. ORIENTE:Bayate, 4
Apr 1917,Ekmans.n. (S);MonteVerde,14 Sep, Wright original description and has been selected by
s.n. (M-KREMPELHUBER). Sineloc., Wrights.n. (M, Swinscowand Krog(1981) as the lectotype.It is
UPS). closest to Phyllopsora canoumbrina,which is
PUERTO RICO. MAYAGUEZ: Indiera Fria, near isidiate and has more
granularsquamules and
Maricao,430-800 m, 19-22 Mar 1915, N. Brittonet smaller and P. confusa, which has
al. 4509 (NY). ascospores,
VENEZUELA. NUEVA ESPARTA:Isla Margarita, thicker,more divided and ascendingsquamules.
CerroMatasiete,11 Apr 1986,Brako8631 (NY, VEN), Additionalmaterialis needed for clarificationof
8632, 8635, 8636, 8637 (NY). this species. Specimen "b" is P. corallina var.
Discussion. This species is closest to Phyllop- phaeobyssina.
sora intermediella from which it differs by its
isidial morphology, pale-margined apothecia and 16. Phyllopsoraparvifolia(Persoon)Miller Ar-
by its longer ascospores. Only the type material goviensis, Bull. Herb. Boissier 2(Appendix1):
and the Wright collection from the Krempel- 90. 1894.
huber hb. are fertile. Figs. 3c, d, 4a, c, 6a, b, d, 7a-e, 8a, 9c.
LecideaparvifoliaPersoon in Gaudichaud,Voy. Ura-
15. Phyllopsora minor Brako, Mycotaxon 35(1): nie 192. 1827. Biatora parvifolia (Persoon) Mon-
15, 1989. Fig. 3b. tagne, Ann. Sci. Nat. Bot., ser. 2, 4: 92. 1835. Par-
melia parvifolia(Persoon)Montagnein Sagra,Hist.
Lecideacorallina(Eschweiler)MullerArgoviensisvar. fis. Cuba, Bot. 214, tab. 10, fig. 3. 1838. Psorapar-
schizophylloidesVainio, J. Bot. 34: 106. 1896. Phyl- vifolia(Persoon)Massalongo,Framm.lichenogr.25.
lopsoracorallina(Eschweiler)Miller Argoviensisvar. 1855. Zeoraparvifolia(Persoon)C. Miiller,Bot. Zei-
schizophylloides(Vainio) Zahlbruckner,Cat. lich. tung (Regensburg)15: 386. 1857. Type. Brazil. Rio
univ. 4: 397. 1926 (non Phyllopsoraschizophylloides de Janeiro,Gaudichauds.n. (holotype,PC;isotypes,
(Malme)Schneider= Phyllopsorabuettneri(Muller BG, G).
Argoviensis) Zahlbrucknervar. munda (Malme)
Brako).Type. St. Vincent. Richmond Peak, 1000- Thallus squamulose to subfoliose, subfoliose
2000 m, Elliott 261 p.p. (lectotype designated by thalli
Swinscow & Krog, 1981, TUR-VAIN 22612A, no
reachingover 1 cm wide, squamuleselon-
lichen substances;isolectotype,BM). gate, often denselymicrophylline-lobulate,lobes
deeply incised, 0.5-1.0 mm wide, complanateto
Thallus squamulose, squamules round to ir- convex, adnate to slightly ascending,Pd-. Up-
regular or elongate, lobes 0.1-0.3 mm wide, dis- per surfaceglabrous,pubescent and long fibril-
crete to adjoined and overlapping, complanate lose at the margin.Isidia presentor lacking.Cor-
to convex, adnate, Pd-. Upper surface glabrous, tex type 1, 1-2 or 2, 30-60 .amthick. Prothallus

abundant, pale to reddish. Lichen substances: Phyllopsora weberi Ferraro, Bol. Soc. Argent. Bot. 24:
179. 1985. Type. Argentina. Misiones: Dept. San
parvifoliindetected by HPLC.
Ignacio, 8 Dec 1981, Ferraro et al. 2231 (holotype,
Apotheciacommon. Disc plane to convex, tan CTES; isotype, COLO, no lichen substances).
to dark reddish-brown.Margin slightly raised,
concolorouswith the disc or darker,smooth or Distribution (Fig. 21). Pantropical, sea level-
with abundant white projectingfibrils. Exciple 3200 m.
and hypotheciumtan, KOH-. Hymenium col- Specimens examined. UNITED STATES. (Repre-
orless to tan, to 55 ,tm tall. Epihymeniumnot sentative specimens).ALABAMA:Mobile Co., Mobile,
obvious. Ascosporessimple to uniseptate,ovoid, 1901, Mohr s.n. (US). FLORIDA:Duval Co., Jackson-
ellipsoidor shortfusiform,(7.0-)8.0-13.0 x 2.5- ville, Calkins 433 (NY); Highlands Co., Lake Placid,
Archbold Biological Station, 9 Oct 1985, Thor 4516
5.0 ,um.
(S). HAWAII:Hawaii Volcanos National Park, Kipuka
Pycnidiacommon, tan, partiallyimmersed in Pua'ulu on Mauna Loa strip road, 4000 ft, 2 Apr 1983,
the thallus. Conidia rod-shaped,straight,9.0 x Weber& BujakiewiczL-73252 (COLO,GZU). LOUI-
1.0 Am. SIANA:Livingston Parish, Livingston, 26 Sep 1968,
Discussion.This speciesis distinguishedby its Tucker 7637 (LSU). MISSISSIPPI:
Wilkinson Co., Clark
CreekNaturalArea, ca. 1 mi. S of pond, Boullions.n.
large,elongate, +lobulate squamulescontaining (LSU). TEXAS:Harris Co., Houston, 1872, Hall s.n.
parvifoliin. It is closest to Phyllopsorafendleri (FH-TUCK 2828), 1869.
which has smallersquamuleswith short lobes, a MEXICO. TAMAULIPAS: Las Palmas, Pringle 20 (FH),
brownhypotheciumand longerconidia.Two va- 6 Jun 1890, Pringle 219 (FH).
COSTA RICA. PUNTARENAS: Monteverde, 1450 m,
rietiesarerecognizedin the neotropicswhich are 4 Jan 1979, Hafellner 7552 (GZU), Sipman 12099,
distinguished by different cortex types and by 12201 (B).
differencesin spore dimensions. Phyllopsoraja- PANAMA PANAMA:Along trail to Cerro Brewster
vanica, known from the Old World, may prove from Rio Pacora valley, 670 m, 18-21 Nov 1985, Brako
to be an isidiate variety of P. parvifolia. 8270 (NY).
CUBA. SANTIAGO DECUBA:"Monte Kentucky," ca.
5 km SE of La Gran Piedra, 1000 m, 5 Apr 1982,
Harris 14410A (HAC, NY). Without prov.: Wright
s.n., Lich. Cub. ser. 2, 105 (UPS).
Key to the Varieties of DOMINICAN REPUBLIC. INDEPENDENCIA: Sierra
de Baoruca, 30.5 km S of Puerto Escondito, 1940 m,
Phyllopsora parvifolia 25 Jan 1987, Buck 14655 (NY), Harris 20587 (NY);
la. Thallusuppercortextype 1-2, greaterthan50 23.5 km S of Puerto Escondito at intersection of road
gm thick;ascospores to Charco Colorado, 1750 m, 24 Jan 1987, Harris
ovoid. ................
................. P. parvifolia var.breviuscula.W 20496, (NY). LA VEGA:East part of La Cienaga, 6 km
lb. Thallusuppercortextype2, less than50 um; of Monabao, 11 May 1982, Harris 15677 (NY);
Arroyo Piedrosa, 1.2 km from Monabao on road to
ascospores ellipsoidto shortfusiform ........
11 May 1982, Harris 15683 (NY); Piedra
................. P. parvifolia var.parvifolia. Jarabacoa,
Blanca, 4 km from Jarabacoa then 5.7 km on road to
Piedra Blanca, 12 May 1982, Harris 15774 (NY); Salto
de Jimenoa, 7.5 km from Jarabacoa on road to El Rio
16a. Phyllopsora parvifolia (Persoon) Miller de Constanza from Pueblo Salto de Jimenoa, 12 May
Argoviensisvar. parvifolia. 1982, Harris 15861, 15870 (NY); Loma del Puerto,
Figs. 3c, 6b, 7b-e, 8a. 400 m, 18 Jan 1969, Liogier 15725 (NY).
PUERTO RICO. MAYAGUEZ: Three miles E of San-
Lecidea subbreviusculaNylander, Sert. lich. trop. 40. turce, 1899, Heller 448 (NY); Maricao Forest Reserve,
1891. Phyllopsorasubbreviuscula(Nylander)Zahl- ca. 5 m S of Maricao, 18 Jun 1970, Tucker 8682a
bruckner,Cat. lich. univ. 4: 401. 1926. Type. Cuba. (LSU). PONCE:Cordillera Central, above Villalba, Dofia
Wrights.n., Lich.Cub.ser. 2, 120 (holotype,H-NYL Juana Recreation Area, 800-1000 m, 2 Jun 1988, Har-
20524; isotype, FH-TUCK 2922). ris 22005, 22032 (NY).
Lecideaparvifolia(Persoon)Miller Argoviensisvar. COLOMBIA. CUNDINAMARCA: Choachi, 2000 m,
concrescensMalme,Ark. Bot. 28A(7):51. 1936. Lindig 803 p.p. (H-NYL P.M. 4112). TOLIMA: Mun.
Phyllopsoraparvifolia(Persoon)Miller Argoviensis Santa Isabel, El Ochoral, 3130 m, 1980, Valencia &
var. concrescens(Malme)Zahlbruckner, Cat. lich. Boekhout VB177h (B). RISARALDA:
Hda. Perias, Vereda
univ. 10: 377. 1939.Type.Brazil.Rio Grandedo Puerto Caldas, 1000 m, 25 Nov 1985, Wolf 427 (B),
Sul:Col.IjuhynearCruzAlta,30 Mar1893,Malme 29 Nov 1985, Wolf 452 (B).
s.n., Lich. Regnell.1227 (lectotypedesignatedby VENEZUELA. ARAGUA:Agua Fria, 17 Apr 1987,
Swinscow& Krog,1981,S, no lichensubstances; Brako 8949 (NY). BOLiVAR:Parque Nacional Canai-
syntype,Malme s.n., Lich. Regnell. 1235, S). ma, on road from Aeropuerto Luepa to Kavanayen,

54 Flora Neotropica
TROPICAL AMERICA Flora Neotropica base map no 1
?C
"'\?
^ê '^ 1X9
c:. so 70 60 S0 C
'
;-_ X%
i 9~ '0 '. a.0 ,.s-....:-o
c---~~~~~~~~..
?0 200
300 *00 5 ....000. . . . . ..
i. i: ~~~~~~~~~~~i
~
'
F _ 0 .0 0 00101
10 0 00*,~0 0 ~?
R?0?r?d~~~~~~~~~~~.
............... -' '
Prepred by HendrkR Rypkema s e 6
FIG. 21. Neotropicaldistributionof Phyllopsoraparvifoliavar. parvifolia(dots), var. breviuscula(stars),and

Phyllopsoraparvifoliella(triangles).
10 Apr 1985, Brako 8168 (NY). DISTRITO FEDERAL: (Hb. Kalb); BOLIVAR:15 km E of Guaranda on road
Caracas,1887, Ernst 34, 84 (G), 1895, Ernst 188 (G), to Riobamba, 3200 m, 14 Feb 1985, Arvidssonet al.
s.n. (US). MERIDA: El Paramito,near El Morro, 2650 6324 (GB). MORONA-SANTIAGO: Pachicutza at "Es-
m, 21 Dec 1977, L6pez-Figueiras14699 (MERF);at cuela Fiscomisional Cardinal Dopfner," km 140 on
the lake of the Paramode los Granates,2600-2800 m, roadLoja-Gualaquiza,900-1000 m, 26-27 Apr 1973,
28 Dec 1977, L6pez-Figueiras 15160 (MERF). MI- Holm-Nielsonet al. 4559D (AAU).
El Volcan, above Baruta, 1200-1400 m, 1 May
RANDA: BRAZIL. BAHIA:Ca. 10 km N of Rui Barbosa, 400
1986, Brako 8656, 8660, 8665 (NY), 4 May 1986, m, 18 Jul 1980, Kalbs.n. (Hb. Kalb).GOIAS: Between
Brako8692 (NY); along road from Hoya de la Puerta Jatai and Estdncia,510 m, 11 Jul 1980, Kalb 283B
to San Jose de los Altos, 1100-1250 m, 9 May 1986, (Hb. Kalb). MATOGROSSO: Ca. 35 km SE of Cuiaba,
Brako 8695 (NY). TRUJILLO:Hoya del Carruzo, be- 120 m, 4 Jul 1980, Kalb273 p.p. (Hb. Kalb);Serrados
tween El Paramo de Turnal and El Paramo de Cende, Coroados, between Cuiaba and Buriti, 500 m, 6 Jul
2800-2900 m, 1 Apr 1976, L6pez-Figueiras 13211 1980, Kalb276 (Hb. Kalb);Buriti,600 m, 8 Jul 1980,
(MERF);La Cava, between La Hoya del Carruzoand Kalb 280 p.p. (Hb. Kalb), 20 Jan 1894, Malme s.n.,
El Paramo de Cende, 3000 m, 1 Apr 1976, L6pez- Lich. Regnell. 2261 (S, UPS); 10 km NE of Chapada
Figueiras 13240 (MERF). dos Guimaraes,680 m, 9 Jul 1980, Kalb282 p.p. (Hb.
Ca. 35 km S of Cuenca, 3200
ECUADOR.AZUAY: Kalb). MATOGROSSODOSUL: Ca. 30 km S of Campo
m, 24-26 1987, Kalb & A. Kalb 17572, 17573, 17574 Grande,550 m, 14 Nov 1979,Kalb199 p.p. (Hb. Kalb).

MINASGERAIS: Mountainslope above Tiradentesnear caba, 40 km SW of Sao Paulo, 800 m, 19 Mar 1978,
Sao Joao del Rei, 900 m, 6 Jul 1978, Kalb& Plobst29 Kalb & Plobst 1 p.p. (Hb. Kalb); 80 km SW of Sao
p.p. (Hb. Kalb);Serrado Caraca,8 Jul 1978, 1200 m, Paulo, on the Rio Juquia, 700 m, 24 Mar 1978, Kalb
Kalb & Plobst 31 p.p. (Hb. Kalb);above Vila Monte & Plobst5 p.p. (Hb. Kalb);ca. 60 km SW of Sao Paulo
Verde, ca. 30 km E of Camanducaia,1800 m, 3 Jul above Juquitiba,550 m, 14 May 1978, Kalb & Plobst
1979, Kalb 151 (Hb. Kalb);betweenVila Monte Verde 29 p.p., Lich. Neotropici 293 (COLO, GZU, H, M,
and Camanducaia,1300 m, 28-29 Nov 1980, Kalb US); Ilha de Sao Sebastiao,400 m, 22 Apr 1978, Kalb
314 (Hb. Kalb);Lafayette,1885 Vainios.n., Lich.Bras. & Plobst 12 (Hb. Kalb), 500 m, 22 Apr 1978, Kalb &
Exs. 337 (BM);LagoaSanta, Warming133 (M = 2x), Pl1bst13 (Hb. Kalb),400 m, 2 Apr 1979,Kalb&Plobst
s.n. (NY, UPS). PARA:Rio Jamanxim,974-1024 km 137 (Hb. Kalb), 500 m, 6 Jul 1979, Kalb & Plobst157
N ofCuiaba, 300m, 10-15 May 1983, Brako&Dibben (Hb. Kalb);Camposdo Jordao,Pico de Itapeva, 1730
6990 (INPA,NY). PARANA: Guarapuava,17Apr 1956, m, 25 Jan 1987, Brako 8917 (NY), 1850 m, 28 May
Montes 10121F (Hb. Osorio);Prainhas,near Port de 1978, Kalb& Plobst23 (Hb. Kalb), 14 Oct 1978, Kalb
Cima, 100 m, 16 Jan 1987, Hatschbach& Brako8838, &Plobst50p.p.(Hb.Kalb),Lich.Neotropici292 (GZU);
8850 (MBM,NY); ReservaEcologicaSapitanduvabe- betweenTaubat6Ubatuba,800 m, 18 Jun 1978, Kalb
tweenMorretesandAntonina,100 m, 17-18 Jan 1987, 26 p.p. (Hb. Kalb);Serrade Sao Lourenco,NW of Sao
Hatschbach& Brako 8858, 8861, 8872, 8874, 8882, Lourencoda Serra,850 m, 3 Sep 1978, Kalb & Plobst
8884, 8894, 8895 (MBM, NY); near Bateias, W of 45 (Hb. Kalb);above Campos do Jordao,ca. 45 km
Curitiba,19Jan 1987,Hatschbach&Brako8910 (NY); N of Taubat6,14 Oct 1978, 1950 m, Kalb& Plobst49
Guaira,200 m, 9 Aug 1980, Kalb301 (Hb. Kalb);Foz (Hb. Kalb);Ilha Comprida,2 m, 18 Mar 1979, Kalb
do Iguacu, 100-200 m, 22-23 Sep 1984, Vital& Buck & Plobst 136 (Hb. Kalb), 3 m, 15 Jul 1979, Kalb 163
11955 (NY, SP). Rio DE JANEIRO:Itatiaia, between p.p. (Hb. Kalb);near Cachoeirasde Emas, ca. 15 km
Registro do Picii and Agulhas Negras, 10 Jul 1979, NE of Piraqununga,550 m, 14 Jun 1979, Kalb&Plobst
Kalb& Poelt s.n. (GZU = 2x), 23 Jul 1979, Kalb 161 145 p.p. (Hb. Kalb); Anhemi District, Fazenda Bar-
(Hb. Kalb);between Paratiand Cunha, 2 Nov 1979, renco Rico, 450 m, 10 Nov 1979, Kalb 192 p.p. (Hb.
Kalb&P6lbsts.n. (Hb.Kalb);Corcovado,15 Aug 1892, Kalb);3 km E of Botukatu,850 m, 11 Nov 1979, Kalb
Malme s.n., Lich. Regnell. 56, 68 (S); Rio de Janeiro, & Plobst 197 p.p. (Hb. Kalb);Serrade Botukatu,Par-
1885, Vainios.n., Lich. Bras. Exs. 101 (TUR-VAIN dinho, 800 m, 11 Nov 1979, Kalb & Pr6bst 198 p.p.
22610). Rio GRANDEDO SUL: 1 km E de Gramado, (Hb. Kalb);above Maresias,ca. 30 km W of Sao Se-
Av. CasteloBranco,850 m, 10, 12 May 1981, Osorio bastiao, 330 m, 18 Feb 1980, Kalb 212 (Hb. Kalb);
7909 (Hb. Osorio), Lago Negro, 850 m, 10, 12, May between Moji das Cruzesand Aruja, 500 m, 20 Apr
1981, Osorio 7964 (Hb. Osorio);7 km SE of Julio de 1980, Kalb232 (Hb. Kalb);FazendaSaoJoao nearRio
Castilhos, Rio Soturo, Passo de Felicio, 650 m, 14 Claro,600 m, 16 Aug 1980, Kalb302 p.p. (Hb. Kalb);
Oct 1989, Osorio&Fleig89/194, 89/203 (Hb. Osorio); MorroGrandenear Cotia, 850 m, 27 Sep 1980, Kalb
SantaMaria,Repressade CORSAN, 5 km SW of the 309 p.p. (Hb. Kalb).
village Val de Serra,500 m, 13 May 1989, Osorio & PARAGUAY. ALTO PARANA:Centro Forestal Alto
Fleig 89/73 (Hb. Osorio), Vale do Diabo, 400 m, 15 Parana,4.5 km W of Pto. Pte. Stroessneron Ruta 7,
Oct 1989, Osorio & Fleig 89/244 (Hb. Osorio); San 250 m, 15 Oct 1984, Buck 12361 (NY). AMAMBAY:
Franciscode Paula,LagoSio Berardo, 900 m, 18/19 ParqueNacionalCerraCora,alongtrailup CerroMur-
May 1987, Osorio& Fleig 2SF/31, 2SF/60 (Hb. Oso- alla, 300 m, 19 Oct 1984, Buck 12530 (NY). CENTRAL:
rio);PortoAlegre,29 Sep 1892, Malme s.n., Lich.Reg- Asunci6n, 1878, Balansa 16 (G);Villa Morra, 14 Aug
nell. 503 (S, US), 3 Oct 1892, Malme s.n., Lich. Reg- 1893, Malme s.n., Lich. Regnell. s.n. (S). PARAGUARi:
nell. 526 (S); Hamburgerberg,near Sao Leopoldo, 18 ParqueNacional Ybycui, along trail to Mirador,200
Oct 1892, Malme s.n., Lich. Regnell. 617B (S); Santo m, 4 Oct 1984, Buck 11769 (NY), along trail to Salto
Angelo near cachoeira,8 Feb 1893, Malme s.n., Lich. Mbocaruzfion Rio Corrientes,ca. 200 m, 6 Oct 1984,
Regenell. 1021 (S); Silveira Martins, 28 Feb 1893, Buck 12093 (NY). Guarapi,Apr 1881, Balansa 4201
Malmes.n., Lich.Regnell.1092 (S),8 Feb 1893,Malme (BM, G = 3 x, H-NYL P.M. 4111, L, M, W), 30 Jul
309A, Lich. Regnell. 1021A (W);ColoniaIjuhy,2 Apr 1881, Balansa 4202 (BM, G = 3 x, M, W). CerroCu-
1893, Malme s.n., Lich. Regnell. 1229 (S), 6 Apr 1893, rupatiti,Apr 1879,Balansa 4199 (BM,G = 2 x), 1887,
Malme s.n., Lich. Regnell. 1235B (S), Malme 309B, Balansas.n. (G); 1878-1884, Balansa s.n. (G, H-NYL
Lich. Regnell. 1235 (W); South of Torres,on road to P.M. 4110). Cerro Le6n, 23 Jul 1881, Balansa 4207
Camping Itapeba, 9 Jan 1987, Falkenberg& Brako (G = 2x). CerroYaguaron,1887, Balansa.4150 (G).
3603, 3612 (NY); 10 km E of Tainhas, on road from ARGENTINACHACO: Lero.de Mayo, ColoniaBe-
Terrade Areia to Tainhas, 850 m, 10 Jan 1987, Fal- nitez, Reservadel INTA, 16 May 1979, Ferraroet al.
kenberg& Brako 3633 (NY); Itaimbezinho, 10 Jan 1820 (CTES);Gran Chaco, 15 Sep 1893, Malme s.n.
1987, Falkenberg& Brako 3645 (NY), 1100 m, Grii- (S). CORRIENTES: Espedrado, El Sombrerito, Estaci6n
ninger 3018 (BG). SANTA CATARINA:13 km above ExperimentalINTA, Costa del Rio Parana,26 Nov
Timbe do Sul, on road to Serrada Rochina, 650 m, 1978, Ferraroet al. 1410 (BG, CTES);Saladas,Ruta
11 Jan 1987, Falkenberg& Brako3653, 3657 (FLOR, 12, 27 Dec 1983,Ferraro2830 (CTES);Ituzaing6,Rin-
NY); between Uribici and CapivaraAlta on road to can de Santa Maria, 22 Nov 1977, Neiffs.n. (CTES);
Bracodo Norte, 1040 m, 12 Jan 1987, Falkenberg& Rio Uruguay, Isla Itacumbfi,21 Aug 1977, Achaval
Brako3706, 3720 (FLOR,NY). SAOPAULO: Sao Pau- s.n. (Hb. Osorio). ITAPUA:Cantera, 19 Jul 1957, Mon-
lo, 6 Oct 1922, Hoehne s.n. (H); Serrade Paranapia- tes 12042F (NY); Colonia Fram, 25 Jul 1957, Montes

56 Flora Neotropica
12051F, 12052D (NY). MISIONES:Candelaria,Arroyo Distribution. (Fig. 21). Pantropical, 650-

Yabebiry and Ruta 12, 18 Dec 1981, Ferraroet al. 1600 m.
2316 (CTES);Puertodo Iguaz6,200 m, 31 Aug 1980,
Kalb 306 p.p. (Hb. Kalb). SANTAFE: Rosario, 13 Sep Specimens examined: UNITED STATES. ALA-
1893, Malme s.n., Lich. Regnell.s.n. (S);Colonia Ris- BAMA: BaldwinCo., Yupon Point, 1 Mar 1925, Evans
so, 23 Sep 1893, Malmes.n., Lich. Regnell. 1817B (S), 202 (NY).
28 Sep 1893, Malme s.n., Lich. Regnell 1862A (S = VENEZUELA.AMAZONAS: San Carlos,Rio Negro,
3 x), 1864C(S), 7 Oct 1893, Malmes.n., Lich. Regnell. Spruces.n., Lich. Amaz. And. 142 (G = 3 x, H-NYL
1897A & B (S), 8 Oct 1893, Malme s.n., Lich. Regnell. 20526,20527, PC = 2 x, W = 2 x). DISTRITO FEDERAL:
1911A (S), 8 Oct 1893, Malme s.n., Lich. Regnell. Caracas,Ernst 5 (US), 95, 139, 723, s.n. (G).
1912C(S, UPS); 1882, Spegazzinis.n. (H-NYL 20544, BRAZIL.MATOGROSSO: Chapadados Guimaraes,
W= 2x). 800 m, 6 Jul 1980, Kalb 277 p.p. (Hb. Kalb);ca. 20
URUGUAY. ARTIGAS: ArroyoEl Tigre y Rio Uru- km NE of Chapadados Guimaraes,670 m, 9 Sep 1980,
guay, 14 km W de ColoniaPalma,5-8 Jul 1975, Osorio Kalb 281 p.p. (Hb. Kalb);Santa Anna de Chapada,2
7261 (Hb. Osorio). DURAZNO: El Cordobes,Estancia Mar 1894, Malmes.n., Lich.Regnell.2485, Lich.Aus-
Las Pitangas,El Prado, 8-10 Feb 1953, Osorio 2950 troamericani233 (G, H, S, US = 2x, W). MINAS
(Hb. Osorio). ROCHA: Ruta 9 y Arroyo Chafalote, 17 GERAIS: Caldas,Malmes.n., Lich.Regnell.s.n. (COLO,
Feb 1967, Osorio5487, 5510 (Hb. Osorio) S); 5 km W of Monte Verde, 1400 m, 2 Jul 1979,
TANZANIA. TANGAPROv.:Usambara,Hoist 3330 Sipman 12821 (B);Monte Verde, 1600 m, 2 Jul 1979,
p.p. (G). Sipman 12909 (B). SXo PAULO: Itapira,16 May 1927,
AUSTRALIA. NEW SOUTH WALES: Braidwood Hoehne s.n. (W);Apiahi, Puiggaris.n. (G).
District, between Majors Creekand Araluen, 18 Oct ARGENTINA.MISIONES: Iguazfi,roadto Garganta
1967, Weber & McVean L-47095 (COLO). South del Diablo, 2 Mar 1982, Ferraro2579 (CTES).
Queensland.Lowood, 1890, Wilsons.n. (H). AUSTRALIA. QUEENSLAND:Brisbane, 1887, Bailey
s.n. (G); Rosewood Margins,Bailey s.n. (B), 13 (G);
Discussion. Variety parvifolia has a type 2 cor- Toowoomba,Hartmans.n. (FH = 2 x, FH-TAYLOR
tex, to 30 ,um thick, and ellipsoid to short fusi- 684, G = 5x, M, UPS = 3x, W = 2x).
form ascospores, 8.0-10.9-12.5 x 2.0-2.7-3.0
Discussion. This variety differs from var. par-
,im. The squamule lobes are thinner than in var.
breviuscula. Lobules may completely dominate vifolia by its type 1-2 cortex that is more than
50 Am thick, and by its ovoid ascospores, 7.0-
the thallus. Malme (1936) recognized the highly
8.4-10.5 x 2.5-3.8-4.0 Am. The thicker cortex
lobulate forms as var. concrescens. Collections
from Andean Ecuador (Kalb & A. Kalb 17572- may give the thallus a slightly convex appear-
ance. The thallus is rarely lobulate. It is closest
17574 (HB. Kalb) and Arvidson et al. 6324 (GB))
to Phyllopsorafendleri, which has shorter thallus
have smaller squamules and/or more divided
a and longer
lobes than most collections of Phyllopora par- lobes, yellow-brown hypothecium
ascospores. Some specimens of Wright s. n., Lich.
vifolia.
Cub. 181 are labeled Lecidea parvifolia var. fus-
cescens, apparently a herbarium name used by
Tuckerman, but never published.
16b. Phyllopsora parvifolia (Persoon) Muller
17. Phyllopsora parvifoliella (Nylander) Miller
Argoviensis var. breviuscula (Nylander) Brako,
Mycotaxon 35(1): 15. 1989. Argoviensis, Bull. Soc. Roy. Bot. Belgique 32:
131. 1893. Fig. 3e.
Figs. Id, 4c, 6a, d, 7a, 9c, 21.
LecideabreviusculaNylander,Ann. Sci. Nat. Bot., ser. LecideaparvifoliellaNylander,Ann. Sci. Nat. Bot., s6r.
4, 19: 339. 1863. LecideaparvifoliaPersoonin Gau- 4,19:339. 1863. Type.Cuba.Wrights.n., Lich.Cub.
dichaudvar. breviuscula(Nylander)Nylander,Ann. 182 (holotype,H-NYL 20545, no lichen substances;
Sci. Nat. Bot., ser. 5, 7: 321. 1867. Psorabreviuscula isotypes, BM, BUF, trace of argopsin?,FH-TUCK
(Nylander)Miller Argoviensis,Flora65:483. 1882. 2919, G = 3x, L = 2x, M, UPS, W = 2x).
Phyllopsora breviuscula(Nylander) Miller Argo- Thallus squamulose, squamules elongate and
viensis,Herb.Boisser2(Appendix1):45. 1894.Type.
Cuba.[Oriente:]MonteVerde, Wrights.n. (holotype, lobed, lobes 0.1-0.3 mm diam., complanate to
H-NYL 20557, no lichen substances;isotypes as slightly convex, mostly discrete, loosely adnate
Wrights.n., Lich.Cub. 181, B, BM, BUF, FH-TUCK to ascending, Pd-. Upper surface glabrous,
2921, G=3x, L = 2x, UPS). slightly pubescent or fibrillose at the margin. Isidia
Phyllopsorabrachyspora MiillerArgoviensis,Bot.Jahrb.
Syst. 20: 264. 1894. Type. Tanzania."Hochwaldob common, cylindrical. Cortex type 1, 40 um thick.
Kwa Mstufain Usambara,D.O. Afr.," 1894, Hoist Prothallus abundant, red. Lichen substances: trace
9181 p.p. (holotype,G; isotypes, BM, M, W). argopsin?

Confused Names 57
Apotheciacommon. Disc plane to convex, yel- Thallus squamulose, squamules granular to

low-brown to dark red. Margin slightly raised, elongate,0.1 mm wide, closely adjoinedto form
concolorousor darkerthan the disc. Exciple and an almost continuouscrust,adnateto ascending,
hypotheciumpartlyred pigmented,KOH-. Hy- Pd-. Upper surfaceglabrous,slightly fibrillose
menium colorless, to 40 ,m tall. Epihymenium at the margin. Isidia cylindrical.Cortextype 2,
red.Ascosporessimple,ovoid, 4.0-6.0-6.5 x 2.0- 20 Am thick, in part extending over the entire
2.5-3.0 ,um. squamule. Prothallus abundant, pale. Lichen
Pycnidia not seen. substances:none detected.
Distribution(Fig. 21). CentralAmerica, West Apotheciacommon. Disc planeto convex, tan.
Indies, South America and southern North Marginslightlyraisedand wavy, brightred, with
America, sea level-550 m. abundant white projecting fibrils. Exciple red
pigmented, KOH-. Hypotheciumtan, KOH-.
Specimensexamined.UNITED STATES. FLORIDA:
Lake Co., Lake Norris, 5 Jun 1921, Kelly s.n. (NY, Hymeniumtan, to 40 timtall. Epihymeniumcol-
US). orless. Ascosporessimple, ellipsoid to short fu-
COSTA RICA. 1893, Tonduzs.n. (G). siform, 11.0-12.9-15.5 x 2.5-2.7-3.0 tm.
EUSTATIUS. Vulcano Quill, 450-500 m, 30 Jul
1980,Sipman 14819, 14820 (B), 14833 (B, NY), 14852 Pycnidianot seen.
(B), 540 6
m, Aug 1980, Sipman 15069 (B).
Distribution.Paraguay.
VENEZUELA. AMAZONAS: Rio Negro, at base of Discussion. This species is only known from
Cerrode la Neblina, 140 m, 23 Feb 1894, Buck 11118 type material. It is distinguishedby its closely
(NY, VEN);San Carlos,Spruces.n., Lich. Amaz. And. adjoined squamules, its bright red-marginate
221 (BM).NUEVA ESPARTA: Isla Margarita,CerroMa-
tasiete, 11 Apr 1986, Brako 8624, 8625 (NY). apotheciaand long ascospores.
GUYANA.UpperMazaruni,PakaraimaMountains,
ca. 2 km NW of Kamarang,ca. 500 m, 4 Feb 1985,
Sipman & Aptroot18279 (B).
CONFUSED NAMES, NOMINA
FRENCH GUIANA. Saul, 2 km SW of the village, DUBIA, AND EXCLUDED
"sentierLimonade," 180-200 m, 4 Jul 1986, Mont- SPECIES
foort &Ek 103, (U), 18 Jul 1986, Montfoort&Ek 108,
(U), 11 Aug 1986, Montfoort& Ek 122, 123 (U), 29 Biatora
Aug 1986,Montfoort&Ek 152 (U), 6 Sep 1986, Mont- pyrrhomelaenaTuckerman,Amer.J. Sci.
foort & Ek 165 (U). Arts, ser. 2, 28: 205. 1859. Phyllopsorapyr-
PERU. LORETO:Iquitos, Explorama Lodge, 100 m, rhomelaena(Tuckerman)Swinscow & Krog,
23 Jan 1981, R. SantessonP7:3 (UPS). Lichenologist13: 244. 1981. Type. Cuba. [Or-
BRAZIL.AMAZONAS: Rio Uatuma,8 & 11Aug 1979,
Buck 2737C (INPA, NY). MINASGERAIS: 1885, Gla-
iente:] Monte Verde Woods, 7 Apr, Wright
ziou s. n. (G). RONDONIA:
Vicinity of first rapids on the s.n., Lich. Cub. 178 (holotype, FH; isotypes,
Rio Pacfs Novos, sea level-2 m, 24-28 Mar 1978, BM, BUF, L = 2x, UPS, US = 2x).
Reese 13570 (INPA, NY).
This species belongs in an undescribedgenus
Discussion.This speciesis distinguishedby the in the Bacidiaceae.It differsfromPhyllopsoraby
combination of its medium-size, elongate, isid- its apothecialanatomy and pigmentation.
iate squamules, red apothecial pigment and small
ovoid ascospores. Phyllopsora corallina var. san- LecideaaleuroidesStirton,J. Linn. Soc., Bot. 14:
tensis has more adnate squamules and longer as- 469. 1875. Phyllopsora aleuroides (Stirton)
Miller Argoviensis, Bull. Herb. Boissier
cospores.
2(Appendix 1):45. 1894. Type. New Zealand.
Near Wellington, Buchanan s.n. (lectotype
designated by Galloway, 1983, GLAM; iso-
18. Phyllopsora subcrustacea (Malme) Brako, lectotypes, BM, WELT).
Mycotaxon 35(1): 15, 1989. Fig. 3f. =Psoromidiumaleuroides(Stirton) D. Gallo-
LecideacorallinaEschweilervar. subcrustaceaMalme, way.
Ark. Bot. 28A(7): 47. 1936. Phyllopsoracorallina Lecidea congregansZahlbruckner,Akad. Wiss.
(Eschweiler) Miller Argoviensis var. subcrustacea Wien, Math.-Naturwiss.K1., Denkschr. 104:
(Malme)Zahlbruckner, Cat.lich. univ. 10: 377. 1939.
Type. Paraguay.Asunci6n, 18 Aug 1893, Malme
305. 1941. Phyllopsora congregans (Zahl-
s.n., Lich. Regnell. 1612b (holotype, S, no lichen bruckner)D. Galloway, New ZealandJ. Bot.
substances;isotype, UPS). 21: 196. 1983. Type. New Zealand.Otago,Mt.

58 Flora Neotropica
Cargillnear Dunedin, 400 m, May 1935, J. S. its pseudoparenchymatous hypothecium, its

ThomsonA96 (lectotype designatedby Swin- crustosethallusand its algaltype, which does not
scow & Krog, 1981, W3424). agreewell with Pseudochlorella.
=Trapeliopsis congregans (Zahlbruckner)
Brako Phyllopsorabyssiseda(Nylander ex Hue) Zahl-
bruckner,Cat.lich. univ. 4:396.1926. Lecidea
LecideacoroniformisKrempelhuber,Verh.Zool.-
Bot. Ges. Wien 18: 327. 1868. Phyllopsoraco- byssisedaNylander,Flora41: 380. 1859, nom.
nud.LecideabyssisedaNylanderex Hue,Nouv.
roniformis (Krempelhuber)Zahlbrucknerin Arch. Mus. Hist. Nat., Paris, ser. 3, 3: 103.
Engler& Prantl,Nat. Pflanzenfam.1(1*): 139. 1891. Type. Mexico. Miller s.n. (holotype,H-
1905. Biatora coroniformis (Krempelhuber)
NYL 20517).
Jatta, Flora Ital. Cryptog.3: 520. 1911. Type.
United States. Texas: ex. hb. Endlicher(ho- The type collection is sterile and in poor con-
lotype, M). dition and cannot be identified.
=Psora crenata(Taylor)Reinke
LecideacuratellaeMalme, Ark. Bot. 28A(7):42. PhyllopsoracryptocarpaRiddle, Mycologia 15:
1936. Phyllopsora curatellae (Malme) G. 80. 1923. Type. Cuba.Isle of Pines:San Juan,
Schneider,Biblioth.Lich. 13:175.1979. Type. 15, 17 Mar 1916, Britton et al. 15588 (holo-
Brazil. Mato Grosso: Cuiaba, 27 Nov 1893, type, FH; isotypes, FH, NY).
Malme, Lich. Regnell. 2038 (lectotype desig- =Fellhaneracryptocarpa(Riddle) Brako
nated by Swinscow & Krog, 1981, S; isolec-
totype, UPS). Phyllopsora halei (Tuckerman)Zahlbruckner,
This species belongs to an undescribedgenus Cat. lich. univ. 4: 398. 1926. Pannaria halei
in the Lecanoraceaebased on its ascal type, dark Tuckerman,Amer. J. Sci. Arts, ser. 2, 25: 424.
hypothecium and small ascospores. 1858. Lecidea halei Nylander,Mem. Soc. Sci.
Nat. Cherbourg5: 337. 1857, nom. nud. Le-
LecideastylophoraMalme,Ark. Bot. 28A(7):40. cidea halei (Tuckerman) Tuckerman, Proc.
1936. Phyllopsora stylophora (Malme) G. Amer. Acad. Arts Sci. 6: 274. 1866. Type.
Schneider,Biblioth.Lich. 13:179. 1979. Type. U.S.A. Louisiana: 1853, Hale [58] (holotype,
Brazil.Mato Grosso:Serrada Chapada,Buriti, FH-TUCK 2828, atranorin;isotypes H-NYL
27 Jun 1894, Malme s.n. (holotype, S, atra- 20521, 20522).
norin, terpenes;isotypes, G, H, US).
This species belongs to an undescribedgenus pannosaMiillerArgoviensis,Bot.Jahrb.
Phyllopsora
in the Lecanoraceaebased on its ascal type and Syst.20: 265. 1894.Lecideapannosa(MillerAr-
Ber.Tatigk.St. Gallischen
goviensis)Stitzenberger,
simple, hyaline, broad ascospores. Naturwiss.Ges. 1893-1894:246. 1895.Type.Tan-
zania.[TangaProv:]Usambara,"Kwambugu-Hoch-
Lecidea thaleriza Stirton, Trans. Glasgow Soc. walderin D.O. Africa,"Hoist1432(lectotypedes-
Field Nat. 5: 217. 1877. Phyllopsorathaleriza ignatedby Swinscow& Krog,1981,G, atranorin;
(Stirton)G. Schneider,Biblioth.Lich. 13:180. isolectotype,
BM).
1979. Type. South Africa. Somerset East,
Boschberg,1874, McOwans.n. (holotype,BM; This species is distinguished by its unusual
not found in GLAM). thallustype. The centerof the thallusis more or
less crustose,with discreteto ajoinedsquamules
Psoracompaginata MiillerArgoviensis,Rev. Mycol. at the margin.It has been confusedwith Crocynia
10: 60. 1888. Phyllopsoracompaginata(MiillerAr- pyxinoidesNyl. whichlacksa cortex,has a
G.Schneider, green-
goviensis) Biblioth.
Lich.13:175.1979. ish-black in the and
Type.Paraguay. CerroSanThomas,Jun 1881,Ba- pigmentation apothecia pro-
lansa4134(holotype,G, no lichensubstances; iso- thallus,and may also contain norsticticand stic-
types,M, W;possibleisotype,H-NYL4132). tic acid aggregates.Phyllopsorahalei is known
from the type materialfrom Louisianaand Tan-
This species belongs to an undescribedgenus zania,and fromnumerousrecentcollectionsfrom
in the Bacidiaceae.It differsfrom Phyllopsoraby east Africa.

Confused Names 59
PhyllopsoraleprosaW. Riedl, Osterr.Bot. Z. 121: Mycol. 33: 44. 1935. Type. U.S.A. Florida:
145. 1973. Type. Surinam. 1827, Weigels.n. Sanford,Rapp 62 (holotype, W).
(holotype, W). =Fuscidea subfilamentosa (Zahlbruckner)
=Crocyniagossypina(Swartz)Massalongo Brako
Phyllopsora melanocarpa Miiller Argoviensis, Phyllopsorasubhyalina (Stirton) Zahlbruckner,
Hedwigia34: 28. 1895. Type. Australia.Vic- Cat. lich. univ. 4: 401. 1926. Lecidea subhy-
toria, Wilson 150 (holotype, G; isotype, W). alina Stirton,Trans. Proc. Roy. Soc. Victoria
=Neophyllispachyphylla(MiillerArgovien- 17:77. 1881. Type. Australia.Victoria:Gipps-
sis) G. Schneider land, Waterloo,Stirton 8662 (holotype, BM).
Phyllopsoraparvifolia (Persoon) Miiller Argo- The type collection is a small fragment,too
viensis var. fibrillifera(Nylander)Miiller Ar- scrappyfor full description. The apothecia are
goviensis, Bull. Soc. Roy. Bot. Belgique 32: highlygelatinized,but not typicalof Phyllopsora.
131.1893, Bull. Herb.Boissier2(Appendix1): The ascosporesare simple and ovate, 13-15 x
45. 1894. Lecidea parvifolia Persoon var. fi- 7-9 ,m.
brilliferaNylander, Ann. Sci. Nat. Bot., ser 4,
15: 47. 1861. Psora parvifolia(Persoon)Mas- Phyllopsorasubparvifolia(Persoon) Muller Ar-
goviensis var. dactyligeraMiller Argoviensis,
salongo var.fibrillifera(Nylander)MiillerAr-
goviensis, J. Bot. (Morot) 7: 55. 1893. Type. Hedwigia 34: 141. 1895. Type. Venezuela.
Distrito Federal:Caracas,Ernst 43 (holotype,
New Caledonia. Vieillard1790 (holotype, H-
NYL 20528). G).
The type is squamulose with coralloid isidia
The type is only a small fragment,which can-
to 1 mm long, lackinglichen substancesand ster-
not be identified.
ile.
Phyllopsoraparvifolia (Persoon) Miiller Argo- PhyllopsoraviridisPaulson, J. Siam Soc. (Nat.
viensis var. granulosa (Miiller Argoviensis) Hist.) 8:101. 1930. Type. Thailand.Kaw Tao,
MiillerArgoviensis,Bot. Jahrb.Syst. 20: 264. 22 Sep 1918, ca. 100 m, Paulson29 (holotype,
1894. Psora parvifolia(Persoon) Massalongo BM).
var. granulosa Miiller Argoviensis, Flora 65:
327. 1882. Lecidea parvifolia Persoon var. The type is a small scrap on rock and is too
granulosa(MiillerArgoviensis)Shirley, Proc. small for full study.
Roy. Soc. Queensland6:166.1889. Type.Java. PsoromidiumwellingtoniiStirton,Proc.Phil. Soc.
Junghuhns.n. (holotype, G; isotype, L). Glasgow 10: 304. 1877. Phyllopsorawelling-
The type material is in poor condition and tonii (Stirton)Miller Argoviensis,Bull. Herb.
cannot be identified. Boissier 2(Appendix 1): 45. 1894. Type. New
Zealand. Near Wellington, J. Buchanan s.n.
Phyllopsoraparvifolia (Persoon) Miiller Argo- (lectotype designatedby D. Galloway, 1983,
viensis var. hirtella Bouly de Lesdain, Rev. BM).
Bryol. Lichenol.7: 60. 1934. Type. Cuba. Ar- =Psoromidiumaleuroides(Stirton) D. Gallo-
menia: 700 m, Hioram s.n. (lectotype HAC, way
designatedby Vezda, 1969, in hb.; frag.NY). Thalloidima
janeirensis Miller Argoviensis,
I have only seen the NY fragment,which is Hedwigia 31: 280. 1892. Psorellajaneirensis
sterile and cannot be identified. (Miller Argoviensis)Zahlbruckner,Cat. lich.
univ. 4: 402. 1926. Phyllopsorajaneirensis
Phyllopsora subcorallina Zahlbruckner, Ann. (Miller Argoviensis) Swinscow & Krog, Li-
Mycol. 33: 43. 1935. Type. U.S.A. Florida: chenologist3: 242. 1981. Type. Brazil.Rio de
Sanford, Mar 1928, Rapp 70 (lectotype here Janeiro:Portella s.n. (holotype, BM, fumar-
designated,W; syntype, Rapp 69, FH). protocetraricacid, lobaric acid; isotype, G).
=Catinariasubcorallina(Zahlbruckner) Brako
Phyllopsora stenosporaZahlbruckner, Repert.Spec.
PhyllopsorasubfilamentosaZahlbruckner,Ann. Nov. Regni Veg. 33: 44. 1933. Type: Taiwan. Mt.

60 Flora Neotropica
Arisan,Toroyen,Asahina170(holotype,W,lobaric CAS,CHR, COLO,CTES,F, FH, FLAS,FLOR,

acid;i3otype,NY). G, GB, GLAM, GOET, GZU, H, INPA, L, LD,
=Squamacidia janeirensis (MiullerArgoviensis) LG, LSU, M, MBM, MERF, MIN, NY, O, PC,
Brako
QCA, RB, S, TO, TUR, U, UPS, US, VEN, W,
WELTand ZT. K. Kalb,A. Aptrootand H. Oso-
TYPES NOT SEEN rio generouslyloaned collections from their per-
sonal herbaria.I am grateful to M. Lopez-Fi-
Type materialfor the following taxa were re-
guieras(MERF),G. Hatschbach(MBM), G. D.
quested from, but not located in the herbaria McPherson
listed in the methods section. Type materialmay (MO)and D. Falkenberg(FLOR)and
the members of the 1983 Projeto Flora Expe-
exist in other herbariaand will be searchedfor
dition for assistance with fieldwork. Financial
duringpreparationof a worldwide study of the
supportfor this study was provided by the New
genus. York Botanical Garden, a stipend from the
CUNY StudentTravel and ResearchFund, and
Phyllopsorafoliata (Stirton) Zahlbruckervar. a
atrovirens(Knight) Zahlbruckner,Cat. lich. Sigma Xi Grant-in-Aidof Research.
I thank Amy Rossman and her staff at the
univ. 4: 397. 1926. Lecideafoliata var. atro-
National Fungus Collections, Beltsville, Mary-
virens Knight in Shirley, Proc. Roy. Soc.
land, particularlyJ. Plaskowitz,who allowedme
Queensland 6: 166. 1889. Type: Australia. the use of their facilitiesand
helped in the prep-
Queensland:Fassifern& Milora. aration of photographs.Bobbi Angell skillfully
Lecideafurfuracea Persoon var. biatorina Vai- preparedall line drawingsand Rupert Barneby
nio, Ann. Acad. Sci. Fenn., ser. A, 6(7): 127. assisted with the Latin descriptions.I especially
1915. Type: West Indies. Guadeloupe, Duss thank Paul Berryfor field assistanceand help in
1498. final preparationof the manuscript.
Phyllopsoraglaucoprasina(Sprengel)G. Schnei-
der, Biblioth. Lich. 13: 176. 1979. Lecidea
glaucoprasinaSprengel,Mant.prim.fl. hal. 57. LITERATURECITED
1807. Type: not mentioned in protologue.
Ahmadjian,V. 1967. Lichen symbiosis. Blaisdell
Phyllopsoraparvifolia (Persoon) Miiller Argo- PublishingCo., Massachusetts.
viensis var. coralloides(Montagne)Zahlbruck- .1973. Resynthesisof lichens.Pages 565-579
ner, Cat. lich. univ. 4: 399. 1926. Parmelia in V. Ahmadjian& M. E. Hale (eds.),The lichens.
AcademicPress, London and New York.
parvifolia(Persoon)Montagnevar. coralloides Arvidsson,L. 1982. A monographof the lichengenus
Montagnein Gay, Fl. chil. 8: 145. 1852. Type: Coccocarpia.OperaBot. 67: 1-96.
not mentioned in protologue. Brako,L. 1987. The lichen genus Phyllopsorain the
neotropics.UnpublishedPh.D. Dissertation.City
University of New York, New York.
ACKNOWLEDGMENTS . 1989. Reevaluationof the genusPhyllopsora
with taxonomic notes and introductionof Squa-
I am very grateful to Richard Harris for his macidia,gen. nov. Mycotaxon35(1): 1-19.
advice, guidance and constructivecriticisms of Clements,F. E. & C. L. Shear. 1931. The generaof
the dissertation on which this study is based. fungi. H. W. Wilson, New York.
Coppins,B. J. & P. W. James. 1979. New or inter-
William Buck firstsuggestedthis project,served esting British lichens IV. Lichenologist11: 139-
as my graduatestudent advisor, and has been 179.
helpfulin many ways. I thank G. J. Samuelsand Culberson,C. F. 1972. Improvedconditionsandnew
J. Hafellnerfor comments on an earlierdraftof data for the identificationof lichen productsby a
standardizedthin-layerchromatographicmethod.
the manuscript. Helpful comments by A. Ap- J. Chromatogr.72: 113-125.
troot, S. R. Gradstein, the late M. E. Hale, H. & H. Kristensson. 1970. A standardized
D. Hammond, H. Hertel and H. J. M. Sipman method for the identificationof lichen products.
are also acknowledged. J. Chromatogr.46: 85-93.
I thank the curatorsand staff of the following Culberson,W. C. (ed.). 1986. Natural-productchem-
istry in the systematics of the lichenized fungi.
herbariafor providingloans and assistancewith Bryologist89: 89-138.
herbarium material:AAU, AK, B, BM, BUF, Eriksson,O. & D. H. Hawksworth. 1986. Notes on

Numerical List of Taxa 61
ascomycetesystematics.Nos. 1-224. SystemaAs- the identificationof lichen substances.Bull. Brit.

comycetum5(1): 113-174. Lich. Soc. 34: 3, 4.
Ferraro,L. I. 1983. Una nueva especie de Phyllop- Muller,J. 1894. Conspectussystematicusspecierum
soraceae(liquen).Bol. Soc. Argent.Bot. 24: 179- lichenum Novae Zelandiae.Bull. Herb. Boissier
182. 2(Appendix 1): 1-114.
Galloway,D. 1983. New taxa in the New Zealand Plessl, A. 1963. Uber die Beziehungenvon Hauto-
lichen flora.New ZealandJ. Bot. 21: 191-206. rientype und Organisatiorischihebei Flechten.
Gradstein,S. R. 1975. A taxonomic monographof Osterr.Bot. Z. 110: 194-269.
the genus Acrolejeunea(Hepaticae) with an ar- Poelt,J. 1973. AppendixA. Classification.Pages565-
rangementof the generaofPtychanthoideae.Bryo- 579 in V. Ahmadjian& M. E. Hale (eds.). The
phyt. Biblioth. 4: 1-162. lichens. AcademicPress, London and New York.
Hafellner,J. 1984. Studienin Richtungeiner natiir- Riedl,H. 1973. Phyllopsoraleprosa,eine neue Flech-
licheren Gliederung der Sammelfamilien Leca- ten-Speciesaus Surinam.Osterr.Bot. Z. 121:145-
noraceae und Lecideaceae.Nova Hedwigia 79: 149.
241-371. Schneider,G. 1979. Die FlechtenGattungPsorasen-
Hawksworth,D. H. 1976. Lichen chemotaxonomy. su Zahlbruckner.Biblioth. Lich. 13: 1-291.
Pages 139-184 in D. H. Brown, D. L. Hawks- Sipman,H. J. M. 1983. A monographof the lichen
worth,& R. H. Bailey(eds.),Lichenologyprogress familyMegalosporaceae. Biblioth.Lich. 18: 1-241.
and problems. Academic Press, London, New Swinscow, T. D. V. & H. Krog. 1981. The genus
York, San Francisco. Phyllopsora,with a reporton EastAfricanspecies.
Henssen, A. & H. M. Jahns. 1973 ("1974"). Li- Lichenologist13: 203-247.
chenes. Eine Einfiihrungin die Flechtenkunde. & . 1985. Phyllopsoralongisporasp.
GeorgeThieme, Stuttgart. nov. Nord. J. Bot. 5: 493-495.
Holmgren,P. K., W. Keuken& E. K. Schofield. 1981. Tehler, A. 1983. The generaDirina and Roccellina
Index Herbariorum.Ed. 7. Bohn, Scheltemaand (Roccellaceae).OperaBot. 70: 1-86.
Holkema, Utrecht/Antwerpen,Dr. W. Junk B.V. Timdal,E. 1984a. The genus Hypocenomyce(Leca-
Publishers,The Hague/Boston. norales, Lecideaceae),with special emphasis on
J0rgensen,P. M. 1978. The lichen family Pannari- the Norwegianand Swedishspecies.Nord. J. Bot.
aceae in Europe.OperaBot. 45: 1-123. 4: 83-108.
Karnefelt,I. 1979. The brown fruticose species of . 1984b. The delimitationof Psora (Lecidea-
Cetraria.OperaBot. 46: 1-150. ceae) and relatedgenera,with notes on some spe-
Lamb, I. M. 1963. Index nominum lichenum inter cies. Nord. J. Bot. 4: 525-540.
annos 1932 et 1960 divulgatorum.Ronald Press, Zahlbruckner,A. 1903-1908. Lichens.In A. Engler
New York. & K. Prantl., Die natiirlichenPlanzenfamilien.
.1977. A conspectusof the lichen genusSter- 1(1*):49-249.
eocaulon(Schreb.)Hoffm.J. HattoriBot. Lab.43: .1926-1927. CatalogusLichenumUniversal-
191-355. is 4. Bortraeger, Leipzig.
Malme, G. O. A. 1936. Lecideae:ExpeditionisReg- Zanten,B. 0. van & T. Pocs. 1981. Distributionand
nellianaeprimae.Ark. Bot. 28A(7): 1-53. dispersalof bryophytes.Adv. Bryol. 1: 479-562.
Menlove,J. E. 1974. Thin-layerchromatographyfor
NUMERICALLIST OF TAXA
1. Phyllopsorabibula(Taylor)Swinscow& Krog 7. P. cuyabensis(Malme)Zahlbruckner
2. P. buettneri(MiillerArgoviensis)Zahlbruckner 8. P. fendleri (Tuckerman& Montagne)MullerAr-
a. var. buettneri goviensis
b. var.glauca (Bouly de Lesdain)Brako 9. P. furfuracea(Persoon)Zahlbruckner
c. var. munda(Malme)Brako 10. P. glabella (Nylander)G. Schneider
3. P. canoumbrina(Vainio) Brako 11. P. intermediella(Nylander)Zahlbruckner
4. P. chlorophaea(MiillerArgoviensis)Zahlbruckner 12. P. isidiotyla(Vainio) Riddle
5. P. confusaSwinscow& Krog 13. P. kalbii Brako
6. P. corallina(Eschweiler)MiillerArgoviensis 14. P. longiuscula(Nylander)Zahlbruckner
a. var. corallina 15. P. minor Brako
b. var. glaucella(Vainio) Brako 16. P. parvifolia(Persoon)Miller Argoviensis
c. var. ochroxantha(Nylander)Brako a. var. parvifolia
d. var. phaeobyssina(Vainio) Brako b. var. breviuscula(Nylander)Brako
e. var. rappianaBrako 17. P. parvifoliella(Nylander)Miller Argoviensis
f. var. santensis(Tuckerman)Brako 18. P. subcrustacea(Malme)Brako

62 FloraNeotropica
LIST OF EXSICCATAE
Achaval, F., s.n. (16a). 12408, 12507, 14196, 15489 (5); 12360, 14757,
AguirreC., J. & H. Sipman, 6142 (2b). 15983(6b);7620, 7684, 10334, 11935, 12827, 14509
Allen, C., 14582, 14650 (16a). (6c); 2709A, 12350 (7); 5003, 5646, 14493, 14688
Almborn,O., 10814 (6a). (9); 5883, 8305, 11104, 24720, 24728 (11); 4631,
Aptroot,A., 13589, 20350 (2a); 13338, 14909, 15416, 4634 (13); 11769, 12093, 12361, 12530, 14655,
15595, 18669 (2b); 12506, 14840 (5); 15481 (6a); 15474, 20587 (16a); 2737C, 11118 (17).
15365, 15417 (6c); 15420 (6d); 15155 (6f); 15196, Biittner,Dr., s.n. (2a).
15331 (9). Calkins,W. W., 40 (5); 27, 47, 93, 447 (6a); 138, 139,
Aptroot,A. & R. Hensen, 10726, 12833, 13589 (2b); 149 (6f); 108, 433 (16a).
10650, 15417 (6c); 10362, 10450, 10453, 10456, Cummings,C., 44 p.p. (2b); 37, 49 (9); 44 p.p. (6c).
10536, 10608, 11026, 11229, 12826, 13342, 13344 Davidse, G. & A. Gonzales, 19971 (2b).
(9). Davoli, Fr., 4 (5).
Arsene,Br. G., 3802 (6a). Des Abbayes,H., s.n. (9).
Arvidsson, L. & D. Nilson, 414 (6c). Duby, J., s.n. (6a), s.n. (6d).
Arvidsson,L. et al., 7132 (2b); 6684 (11); 6324 (16a). Dumont, K., CO-5433, CO-5441 (9).
Asahina,J., s.n. (6f). Duss, R.-P., 481 (6d).
Austin, C., s.n. (6a). Eckfeldt,J., 190 (9).
Austin, C. & J. Donnell-Smith,48 (6a). Eiton, G. & L. Eiton, 7495 (2c).
Bailey, F. M., s.n. (16b). Egan,R., EL-7762 (6a); EL-6751 (6f).
Balansa, B., 16, 4150, 4199, 4201, 4202, 4207, s.n. Ekman,E., 37 (6c); s.n. (11), s.n. (14).
(16a). Elliot, W., 135 (2b); 264 (5); 261 p.p. (6d); 261 p.p.
Barbour,P., 1888 (16a). (15).
Beaumont,J. F., 94, 104, 149, 188 (6a). Ernst,A., 38, 45, 114 (4); 190 (6c); s.n. (8); 143 (9);
Bekker,J., 1050 (2b); 1203A (6c). 34, 84, 188, s.n. (16a); 5, 95, 139, 723, s.n. (16b).
Berry,P., 4571 (2b); 4576-4578A (4); s.n. (6c); 4566- Evans, A., 412 (6a); 133 (6f); 202 (16b).
4570 (11). Falkenberg,D. & L. Brako, 3604-3606, 3608, 3609,
Bertero,C., 1648 (1). 3614-3616 (2b); 3708, 3731, 3798 (2c); 3655, 3723
Boom, B. & S. Mori, 978 (6c). (5); 3655B (6a); 3731A (6c); 3603, 3612B, 3722 (9);
Bordas,E., 152A (5). 3603, 3612, 3633, 3645, 3653, 3657, 3706, 3720
Boullion, K., s.n. (16a). (16a).
Brace,L., 6881 (6c). Faurie,A., 307, 308, 413, 539 (6c); 46 (6f).
Brako,L., 9383 (2a); 8164, 8183, 8258, 8267, 8277, Faxon, C., 310, s.n. (6a); s.n. (6c).
8289, 8295B, 8360, 8370, 8371, 8372, 8383, 8385, Fendler,A., s.n. (8).
8390, 8497, 8610, 8668, 8674, 8678, 8686B, 9306, Ferraro,L. 2579 (16b).
9324 (2b);8110, 8605, 8617, 8619, 8662, 8688, 8925 Ferraro,L. & T. Nash, 3220 p.p. (6b); 3220 p.p. (6c).
(2c);7521B, 7526, 8685, 8687 (4);8358, 8510, 8511, Ferraro,L. et al., 1410, 1820, 2231, 2316, 2830 (16a).
8922 (5); 8659, 8667A, 8671, 8672, 8694 (6a);7459, Fleming,H., s.n. (2b); s.n. (6c).
7460, 7463A, 7934A, 8124, 8129, 8131, 8132, 8160, Forbes,C., s.n. (4).
8176, 8181, 8189, 8256, 8386, 8417, 8421, 8428, Freire,46 (7).
8430,8499,8607,8663, 8682,8691 (6c);8223, 8229 Gaudichaud,C., s.n. (9); s.n. (16a).
(6e); 5311, 8220, 8224-8228, 8230, 8231 (6f);6428, Glaziou, G., s.n. (6a); 1867, 1938B (6c); 1938 (9); s.n.
7943, 8111,8117, 8130A, 8158, 8165, 8355, 8507A, (17).
8510, 8517, 8608, 8612, 8622, 8628, 8630, 8631, Griininger,3018 (16a).
8655, 8657, 8664, 8665B, 8666, 8677, 8680, 8933 Hafellner,J., 7434 (6c); 6514, 7465, 7477 (6f); 7552
(9);7531, 8100, 8123, 8694B, 8696 (11); 8635-8637 (16a).
(14); 8168, 8270, 8656, 8660, 8665, 8692, 8695, Hale, M., 35230 (2b); 50881, 50964, 50974 (6e).
8917, 8949 (16a); 8624, 8625 (17). Hale, M. & T. Soderstrom,19841 (5).
Brako,L. & M. Dibben, 6241, 7031 (5); 5523, 6141, Hall, E., 153, s.n. (6a); s.n. (6f); s.n. (16a).
6168-6170, 6527, 6748, 6752, 6758, 6909, 7006A, Hall, E. & H. Ravenel, 16 (6a).
7080, 7267, 7331 (6c); 6036 (6f); 6428 (9); 6614A Harris,R., 14216, 14279, 14325, 14379, 14555, 14656,
(11); 6990, 8917 (16a). 15005, 15563, 20455, 20672, 22217,22234, 22495,
Breedlove,D. & M. Bourbell,67771 (9). 22248, 22447 (2b); 14444, 14559, 14592, 20493,
Britton,E., 684 (5); 6633 (6f). 20495 (2c); 15367, 20473, 20480, 20487, 20489,
Britton,N. & J. Cowell, 4235 (3). 20670,20671 (4); 14424, 15679B, 15750, 19737 (5);
Britton,N. & T. Hazen, 390 (2b). 21137,21138; 14400(6a);22028 (6b);14181, 14282,
Britton, N. et al., 15491 (5); 4398 (6b); 15586 (6c); 14285, 14290, 14350, 14351, 14359,14410, 14445,
14596 (9); 4509 (14). 14523, 14564, 14644, 15383,20794 (6c);21987 (6d);
Broadway,W., s.n. (2b). 2614, 18088, 18113, 18114, 18151, 20779, 21033
Buck,W., 4090A,4195,8348,14618,14634 (2b);2882, (6f); 14453, 14547, 14548, 14657, 15009, 15345,
11028 (3);4361, 4376, 7693, 8362, 9083 (4); 11981, 15679, 15754, 17928, 19751, 20079, 20109, 20471,

List of Exsiccatae 63
20481,20677 (9);21382(13); 14410B,15677, 15683, Krog, H., AU 27/12 (2a); 48/176 (2b).
15774, 15861, 15870, 20496, 20587, 22005, 22032 Krog, H. & T. Swinscow,K48/177 (5); K42/3 (6c).
(16a). Landry,G., s.n. (16a).
Hart, J., s.n. (2b); 34, 153 (4); 104 (6c); 59 (9). Langlois,A., s.n. (5); 846, 848 p.p., "bd," "az" (6a);
Hartman,s.n. (16b). 847 (6f).
Hatschbach,G. & L. Brako,8845, 8860, 8899, 8885- Leon, Fre. et al., 10232 (2b).
8888, 8899 (2b); 8854, 8866, 8870, 8873, 8879, Leprieur,M., 754 (2b); 509 (6a).
8883B, 8896, 8907 (5); 8839, 8856 (6c);8862, 8865, Liebmann,F., P1.Mex. 7381A (6b); PI. Mex. 7372A,
8878, 8892 (9);8838, 8850, 8858, 8861, 8872, 8874, 7373 (6f); P1.Mex. 7381B (9).
8882, 8884, 8894, 8895, 8910 (16a). Lindig,A., 706, 803 p.p., 2791, s.n. (2b);s.n. (6c);840,
Heller, A., 4764 (2b); 448 (16a). 2696 (9); 803 p.p. (16a).
Hertel, H. & B. Oberwinkler,10528 (2c); 10474 (5). Liogier,A., 15725 (16a).
Hioram, Br., 5314, 8909, 9098 (2b); 5306, 5690 (6c); Lopez-Figuieras,M., 22033, 31083 (2b); 13714 (4);
5335, 5345, 5407 p.p. (9). 14704, 22773, 25009 (5); 15974 (6a); 12649 (6c);
Hoehne, F. C., s.n. (16a); s.n. (16b). 13036, 22026 (8); 17289, 17290, 25003 (9); 10014,
Holm-Nielson, L. et al., 4559D (16a). 16238(11);24739(12); 13211,13240,14699,15160
Hoist, C., 1431, 3330 p.p. (2b); 3328 (5); 3330 p.p. (16a).
(16a); 9181 p.p. (16b). L6pez-Figuieras,M. & M. Hale, 19474 (8).
Husnot, P.-T., 455 (5). Lopez-Figuieras,M. & M. Keogh, 14099 (4); 12362,
Johnson, G., s.n. (5); 7080, 7082, 7085, 7088, 7100, 15495 (6c); 15464, 15481, 15498, 15502 (9).
7105, 7110, 7143, 7151, 7158, 7218, 7221, 7226, L6pez-Figuieras,M. & M. Lindstrom,26986 (2b).
7228, 7241, 7263, 7280, 7310, s.n. (6a). Lopez-Figuieras,M. & H. Rodrigues, 25645B (4);
Johnson, H., 355 (7). 22887, 22951, 26280 (8).
Junghuhn,F., s.n. as Lich. Javan. 38 (9). L6pez-Figuieras,M. & H. Sipman, 18483 (8).
Kalb, K., 163 p.p., 215, 229, 231, 309 p.p., 313, 325 L6pez-Figuieras,M. & R. Teran, 9931 (9).
as Lich. Neotrop. 291 (2b); 221 (2c); 309 p.p., 314 MacGillavry,C., s.n. (6d).
pp. (4); 156, 199 p.p., 227, 233, 273 p.p., 277 p.p., McFarland,K. et al., 266 (9).
281, 282 p.p., 302 p.p., 309 p.p., 314 p.p., 315 p.p. McPherson,G., 7847 (6c).
(5); 143, 259, 269 p.p., 275 p.p., 280, 282 p.p., 286, Mains, E., 3653 (9).
290, 293, 295, 302 p.p., 314 p.p. (6a);283 p.p., 300, Malme, G., s.n. as Lich. Reg. 617B, 1251B (2c); s.n.
302 p.p. (6b); 155, 157, 232, 283 p.p., 302 p.p., 309 as Lich.Reg. 236A, 236B, 615, 1275 (4);s.n. as Lich.
p.p. (6c); 212, 302 p.p. (6d); 269 p.p. (6e); 168, 256 Reg. 1471, 1571, 1864F, 1897A, 1944B, 2267C,
p.p., 259, 274, 277 p.p., 280 p.p., 301 p.p. (6f); 269 2268, 2515B, 2737E, s.n. (5);s.n. as Lich.Reg. 1013,
p.p.,275 p.p. (7); 26, 148, 161 p.p., 166, 192, 212, 1227B, 2170C, 2178C (6a); s.n. as Lich. Reg. 599
227 p.p., 232, 233, 280 p.p., 302 p.p., 309 p.p., 313 (6c);s.n. as Lich. Reg. 2481, 2607B (6f);s.n. as Lich.
p.p., 315 p.p. (9); 256 p.p., 258 (13); 151, 161 p.p., Reg. s.n. (7); s.n. as Lich. Reg. 82, 83, 341, 851,
163 p.p., 192 p.p., 199 p.p., 212, 232, 273 p.p., 276, 1226B, 2268B, s.n. (9); s.n. as Lich. Reg. 2547 (10);
277 p.p., 280 p.p., 282 p.p., 283 p.p., 301 p.p., 302 s.n. as Lich. Reg. 861, 887B, 1013, 1208B; s.n. as
p.p. 306 p.p., 309 p.p., 314 p.p., s.n. (16a);277 p.p., Lich. Reg. 2056A and Lich. Austroamericani235A,
281 p.p. (16b). Lich. Reg. 2097C, 2187B as Lich. Austroamericani
Kalb, K. & L. Assis, 174 as Lich. Neotrop. 241 (5). 235A, 2208, 2231, 2233C, 2234B, 2271, 2339B,
Kalb, K. & D. Hannack,59 (6c); 42 (9). 2349, 2349B (13); 309A as Lich. Reg. 1021A, 309B
Kalb, K. & A. Kalb, 17576 (5); 194, 17084, 17566, as Lich. Reg. 1235, s.n. as Lich. Reg. 56, 68, 503,
17569(6a);17568, 17570(6d); 17572, 17573, 17574, 526, 617B, 1021, 1092, 1227, 1229, 1235B, 1817B,
17579 (16a). 1862A, 1864C, 1897A & B, 1911A, 1912C, 2261,
Kalb, K. & G. Plobst, 9 p.p., 20 p.p. (2b), 73 p.p. (3); s.n. (16a);s.n. as Lich. Reg. 2484, s.n. as Lich. Aus-
20 p.p., 309 p.p. (4); 4, 20 p.p., 37, 40, 44 p.p., 50 troamericani233, s.n. (16b);s.n. as Lich.Reg. 1612B
p.p., 95 as Lich.Neotrop. 342, 145 p.p., 161 (5); 266 (18).
as Lich. Neotrop. 343, s.n. (6a);20 p.p., 29, 44 p.p., Magnum,S., s.n. (2a).
73 p.p., 145 p.p. (6c); 1 p.p., 47, 71, 73 p.p., 132 Martius,C., s.n. (6a).
(6f); 73 p.p. (7); 20 p.p. (8); 5 p.p., 9 p.p., 18, 20 Mayrhofer,H. & R. Rogers, 2260 (6c).
p.p., 22, 26 p.p., 31 p.p., 34, 44 p.p., 53 as Lich. Mena, T., s.n. (6b).
Neotrop. 341, 58, 150, 197 p.p., 198 p.p. (9);44 p.p. Merrill,E., s.n. as Bur. Sci. 6256, 6273 (6f).
(12); 1 p.p., 5 p.p., 12, 13, 23, 26 p.p., 29 p.p. as Moberg,R., 1480D, 1464A(2a); 1411, 1481 (5).
Lich. Neotrop. 293, 31 p.p., 45, 49, 50 p.p. as Lich. Mohr, C., s.n. (16a).
Neotrop. 292, 136, 137, 145 p.p., 157, 197 p.p., 198 Montagne,Hb., s.n. (2b); 569 (6a).
p.p., s.n. (16a). Montes, J., 10121F, 12042, 12051F, 12052D(16a).
Kalb, K. & J. Poelt, s.n. (2b); "b" (2c); s.n. (6c); s.n. Montfoort,D. & R. Ek, 100, 101, 115, 127, 129, 147,
(16a). 148, 176, 182 (2b); 110 (6a); 102, 109, 111, 112,
Kalb,K. &A. Schr6gl,13235 (9); 13520,13551,13562 114,137,138,159,174,177,183,191 (6c); 136(6f);
(13). 128,139,144,146, 160,173, 178 (9); 107,133,135,
Kelly, H. A., s.n. (6f); s.n. (17). 140, 150, 151, 154, 192, 193, 196, 199, 200 (11);
Kramer,K. & W. Hekking,3037A (2b). 103, 108, 122, 123, 152, 165 (17).

64 Flora Neotropica
Mori, S. & A. Gentry,4420 (6c). Small, J. et al., 6235 (9).

Morton, C., 7045 (5). Smith, A. C., 3340 p.p. (6a), 3340 p.p. (6e).
Muller,Fr., s.n. (6c). Smith, L. B. & P. Reitz, 10287 (5).
Murrill,W., s.n. (6f). Spegazzini,C., s.n. (16).
Narain, T., 144 (6c); 55 (6f). Spruce,R., s.n. (2b);s.n. as Lich. Amaz. And. 210 (5);
Neiff, s.n. (16a). s.n. as Lich. Amaz. And. 225 (6d);s.n. Lich. Amaz.
Nelson, B., 572 (9). And. 185 (9); s.n. as Lich. Amaz. And. 142 (16b);
Nelson, et al., 491 (3); 1311 (6f). s.n. as Lich. Amaz. And. 221 (17).
NylanderHb., s.n. (4). Standley,P. 20125 (5); 17763, 17916,21360 (6e);17941
Oberwinkleret al., s.n. (8); s.n. (9). (10).
Olrog,C., s.n. (6). Steyermark,J. & M. Rabe, 96279 (5).
Osorio,H., 2950, 5487, 5510, 7909, 7961, 7964 (16a). Steyermark,J. & C. Steyermark,95589 (9).
Osorio, H. & M. Fleig, 2SF/61 (4); 89/139 (9); 2SF/ Streimann,H., 37456, 37501 (2a); 37394 (6a); 36972,
31, 2SF/60, 89/73, 89/194, 89/203, 89/244 (16a). 37019, 37337 (9).
Patillo, 165 (6c). Swinscow,T., 2U 41/4, 2U 45/49B, 3U 25/8, 3U 28/
Peters,T., s.n. (6a). 16 (2a); 3K 16/19, 5K 5/12, K 43/1, 3U 25/9 (2b);
Pittier, H. & T. Durand, s.n. as PI. Costar Exs. 5474 5K 4/53, K 48/7 (4).
(60. Swinscow,T. & H. Krog, K48/177 (5).
Preston-Mafham,16 (5). Thaxter,R., 19 (3); 177 (6b).
Pringle,C., 40 (6a);36A, 10742, s.n. (6c);s.n. (10); 20, Thomas, W. & J. Contreras,3766A (2b).
219 (16a). Thomas, W. et al., 6157 (2b); 3574 (6c); 4392 (6f).
Pruski,J., 2342 (5). Thomasson,K., s.n. (5).
Puiggari,1721 (1881) (4); s.n. (6a); 1721 (1890), 2156 Thor, G., 4516 (16a).
(6c); s.n. (16b). Thwaites,G., 1876 as Leight.Lich. Ceyl. 110 (9).
Pursell,R. 3368 (5). Tonduz, A., s.n. (6f); s.n. (12); s.n. (17).
Rapp, S., 34, 59, 160, 4818, s.n. (5); s.n. (6e); 89 (60; Troxell,C., s.n. (12).
s.n. (9). Tucker,S., 11956 (2c); 7454B, 9045, 9156B, 15286B
Ravenel, H., 11, 101, s.n. (6a); 182, 674, s.n. as Relq. (5);7486,7629B, 9183,9474,11889,12019,12019A,
Tuckermanianae15 (6f). 16044, 17430, 18104, 18152,21638A, 25107, 25402,
Reese, W., 15195A (13); 13570 (17). 25965 (6a); 7144A, 17538 (12); 17566 (13); 6762A,
Rodrigues,K., 847, 933 (6c); 908 (7). 6762C, 7454A, 7637, 7779A, 8447, 8548, 8682A,
Russell, C., 90 (6a). 9911, 10156, 10230, 11815, 12683, 12828, 12891,
Samuels,G. et al., 185 (8). 13249,13884, 14189, 15904, 15668, 15890, 16188,
San Martin,P., s.n. (6c). 17322,18094,18581,21587,21823B, 25110,26654,
Santesson,R., 21688, 21833 (2a); 23030 (4); 20978, 26185, 27132 (16a).
21490, 21560, 21810, 21824, 21826, 21836A & B, Tucker,S. & R. Jones, 11346 (6a).
22983, 23276 (5); P7:19 (60; 23030 (9); P9:8 (11); Tuckerman,E., s.n. (6a); s.n. (6f).
21001A, 21050, 21525 (13); P7:3 (17). Ule, E., 96 (3).
Santesson,R. & R. Moberg,P38:11 (6a). Underwood,L. & F. Earle, 1004 (6a).
Santesson,R. & G. Thor,P72:20, P72:53,P72:59,P72: Vainio, E., s.n. as Lich. Bras. Exs. 318, 335, 366 (4);
73, P73:4 (2b); P74:122 (2c); P71:53, P71:92 (6a); s.n. as Lich. Bras. Exs. 145 (6a); s.n. as Lich. Bras.
P77:18, P77:23 (6c); P70:47, P76:14, P76:18, P76: Exs. 275, 338, 993, 1356, 1443 (6c); s.n. as Lich.
19 (6f); P70:82, P71:15, P71:42 (9). Bras. Exs. 719, 1451, 1469 (9); s.n. as Lich. Bras.
Santesson,R. et al., P102:21 (2b); P101:29, P102:22 Exs. 222 (12); s.n. as Lich. Bras.Exs. 101, 337 (16).
(9). Valencia, H. & T. Boekhout, VB177J (5); VB177H
Saves, T., 21 (9). (16a).
Schallert,P., 1145, 3007 (5). Vanoverbergh,M., Bur. Sci. 1023 (6f).
Schiffner,V., s.n. (2b). van Looy, C., 16 (2b).
Schultes,R., 11753 (6). Vargas,0., 236 (2b).
Sipman, H., 15692 (2a); 10691 (2b); 10899, 10977, Vetch, Dr., s.n. (60).
13053, 13068, 13408 (5); 10984, 14953 (6a); 10687, Vital, D. & W. Buck, 12495 (6a); 12361, 12505 (9);
11989, 12098 (6c); 12061 (6e); 12088, 14088 (6f); 12497 (11); 11955 (16).
10688 (8); 10690, 12098, 12675, 12721, 12959 (9); Vitikainen,0., 9292 (2a).
12099, 12201 (16a); 12821, 12909 (16b); 14819, Warming,J., 133 (16).
14820, 14833, 14852, 15069(17). Weber,C., 1393 (9).
Sipman,H. & A. Aptroot, 17947, 19392, 19411 (2b); Weber,W., L-40262 (11).
19165 (2c); 19489 (5); 18459, 18998 (6c); 18982 Weber,W. & Bujakiewicz,L-73242 (16a).
(6d); 17966, 19003 (9); 18279 (17). Weber,W. & J. Lanier,L-62909 (11).
Sipman, H. & A. Chaverri,20806 (8). Weber,W. & D. McVean,L-47095 (16a).
Sipman, H. et al., 13444, 13548 13553 (2b). Weddell,H., s.n. (6c).
Sipman, H. & M. L6pez-Figuieras,11028 (2c); 11029 Wettstein,R. & V. Schiffher,s.n. (7).
(9). Whitehouse,E., 22632 (6a); 25059 (6f).
Sipman, H. & H. van der Werff, 10899 (5). Wight,A., 10 (6f).

List of Exsiccatae 65
Willey, H., s.n. (5). 2, 119 (9); s.n. as Lich. Cub. 102, 142 (10); s.n. as
Wilson, F. R., s.n. (16a). Lich. Cub. 183 (11); s.n. as Lich. Cub. 179, s.n. (14);
Wolf, J., s.n. (6a); 1228 (9); 427, 452 (16a). s.n. as Lich. Cub. ser. 2, 105, 120 (16a);s.n. as Lich.
Wright,C., s.n. as Lich. Cub. 180, 186, ser. 2, 728 (2b); Cub. 181, 182 (17).
s.n. as Lich. Cub. 185, s.n. (3); s.n. as Lich. Cub. ser Yasuda,A., 350 (6a).
2, 107 (4); s.n. as Lich. Cub. 184E (6a);s.n. as Lich. Zielman,R., 1339A, 1359A (2b); 1364 (2c).
Cub. ser. 2, 105, 726, 727 (6c);s.n. as Lich. Cub. ser
INDEX TO SCIENTIFIC NAMES

Taxa currentlyacceptedin Phyllopsoraarein boldfaceand synonymsarein italics. Genericnames
for species not belonging to Phyllopsoraare in parenthesis following the specific epithet. Page
numbers in boldface indicate primarypage numbers. Page numbers with an asterisk (*) indicate
pages with illustrationsor maps.
albicans 43 crenata(Psora)58
aleuroides(Psoromidium)57, 59 Crocynia12, 14, 21, 58, 59
Argopsis 19 Crocyniaceae12
Bacidia 3, 11, 21, 22, 23 cryptocarpa(Fellhanera)58
Bacidiaceae3, 11, 21, 23, 58 curatella(Lecidea)58
Bacidiopsora16*, 21, 23 cuyabensis7, 26, 35*, 44, 45*
Biatora3, 21, 22 densiflorae38
bibula 6*, 7, 19, 28, 29 efflorescens(Biatora)22
brachyspora56 Erioderma19
breviuscula3, 28, 56 ernstiana39
breviusculaf. glaucina 39 Eschatogonia3, 10*, 11, 21, 22
breviusculavar. glaucella 39 Fellhanera58
breviusculavar. phaeobyssina42 fendleri 19, 22*, 27, 44, 45*, 46, 53, 56
buettneri7, 19, 26, 29-33* foliata var. atrovirens60
buettnerivar. buettneri23*, 29, 30, 31*, 33 formosana 43
buettnerivar. glauca 19, 23*, 26*, 28, 29, 30, 31*-33 furfuracea7, 8*, 14, 17*, 22*, 28, 33, 44, 45*, 4648
buettnerivar. munda23*, 26*, 29, 31*, 33, 52 furfuraceaf. schizophylla34
byssiseda 58 furfuraceavar. biatorina60
canoumbrina6*, 7, 28, 33-34, 35*, 52 Fuscidea59
Catinaria21, 59 glabella 16*, 19, 22*, 28, 48-49
Chiodecton21 glabriuscula40
chlorophaea6*, 14, 15*, 16*, 28, 34-35, 37 glaucoprasina60
cinerella40 gossypina(Crocynia)59
cinerellaf. virescens40 granulifera(Lecidea)33
Cladonia 11 haemophaea46
Cladoniaceae3, 11 haemophaeavar. subparvifolia34
Coccocarpia19 halei 58
Coenogonium21 Hypocenomyce3, 12
compaginata58 hypocrysea46
confusa 6*, 7, 19, 28, 35*-37, 52 intermediella12*, 14, 15*, 28, 33, 45*, 49-50*, 52
congregans(Trapeliopsis)57, 58 isidiotyla 13, 19, 28, 39, 45*, 50*-51
corallina 19, 27, 37-44 janeirensis(Squamacidia)51, 59
corallinaf. glaucina 39 javanica 53
corallinaf. saxicola 43 kalbii 9*, 19, 27, 50, 51*
corallinavar. corallina6*, 19, 27*, 38-39, 44, 48 Lecanora17
corallinavar. glaucella 27*, 38, 39 Lecanoraceae3, 11, 21, 58
corallinavar. ochroxantha6*, 19, 27*, 38, 39-2, 44 Lecanorales14, 21
corallinavar. phaeobyssina7, 11*, 27*, 38, 42, 52 Lecideaceae3, 11, 21
corallinavar. rappiana27*, 38, 42-43 leprosa59
corallinavar. santensis 8, 10*, 11*, 14, 15*, 19, 27*, longiuscula16*, 27, 49, 50*, 51-52
28, 38, 40, 43-44, 57 martinii40
corallinavar. schizophylloides52, 57 Megalospora17, 19, 21, 23, 25
corallinavar. subcrustacea57 Megalosporaceae19
coroniformis58 melanocarpa59

66 Flora Neotropica
melanoglauca30, 31 Pseudocyphellaria19
Micarea 19 Psora 3, 12
microsperma48 Psoraceae11, 12
minor 7, 9*, 28, 52 Psorella3, 11, 16*, 21, 23
miradorensis43 Psoroma 17, 22
munda 33 Psoromidium57, 59
Neophyllis 59 pyrrhomelaena(Biatora)57
Nostoc 11, 13 pyxinoides(Crocynia)12, 14, 58
ochroxantha39 rosei 19
pachyphylla(Neophyllis)59 rosella(Bacidia)14, 16*, 22, 23
Pannaria17 santensis43
Pannariaceae11 schizophylla34
pannarioides(Psorella)23 schizophyllavar. isidiata 46
pannosa 58 schizophylloides33
parvifolia14, 19, 27, 30, 44, 52-56 spinulosa40
parvifoliavar. breviuscula7, 9*, 10*, 12*, 13*, 16*, squamulosa(Physcidia)11
46, 53, 54*, 56 Squamacidia16*, 21, 23
parvifoliavar. concrescens53 stenospora59
parvifoliavar. corallina 37, 38 stylophora(Lecidea)58
parvifoliavar. coralloides60 subbreviuscula53
parvifoliavar. fibrillifera59 subcorallina(Catinaria)59
parvifoliavar. fuscescens56 subcrustacea9*, 19*, 28, 57
parvifoliavar. glauca 30 subfilamentosa(Fuscidea)59
parvifoliavar. granulosa59 subglabella48
parvifoliavar. hirtella 59 subhyalina59
parvifoliavar. parvifolia9*, 12*, 13*, 15*, 53, 54*-56 subparvifolia34
parvifoliavar. subgranulosa33 subparvifoliavar. dactyligera59
parvifoliella7, 9*, 27, 49, 54*, 56-57 subvirescens39
Phyllopsorae3 thaleriza(Lecidea)58
Phyllopsoraceae3 Thalloidima59
Physcidia3, 21 Thelotremataceae19
Pilocarpaceae11, 21 Trapeliopsis58
polydactyla40 Trebouxia5
Porina 21 veralis (Biatora)14, 22
porphyromelaena43 viridis 59
prolifera(Eschatogonia)10* weberi53
Pseudochlorella10, 28, 58 wellingtonii59


Key - Phyllopsora Neotropical

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Organization for Flora Neotropica

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C* \ -'? TROPI'C Of CANCER

Organization for Flora Neotropica

Issued 5 March 1991

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Library of Congress Cataloging in Publication Data

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Presentaddress:MissouriBotanicalGarden,P.O. Box 299, St. Louis, Missouri63166, U.S.A.

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INTRODUCTION spread in tropical and subtropicalareas of the

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Ascospore Length Taxon Ascospore Width

~P. coralina var.phaohbyssina "

_^*^-^-I~~~~~ _ ~P. intermediella

-*w4iii.i--~ IP. kalbii

versity(pers.comm.) providedadditionalchem- tures were made on the day of collection and to

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I .... ;'.' ,',%? .~ " 3~~~~~~~~~~

(Iolcttpe Blii, Wedl .n.,PC,scl=Im

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FIG.2. Phyllopsorafurfuracea (DominicanRepublic,Harris19751, NY). Linedrawing.a & b. Youngapothecia

singlecolony of P. corallinavar. santensisvaries although soredia-like granules have been ob-

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-. ?i" '., .,-L

: ~~'--)~ rr,: :i-

e__ ..X <

FIG.3. a. Phyllopsorakalbii(Holotype:Brazil,Kalb258, NY), scale= 1 mm. b. Phyllopsoraminor(Lectotype:

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gelatinizedhyphaewith narrow,net-likelumina. PseudochlorellaLund(Ahmadjian,pers.comm.).

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FIG. 5. Phyllopsoracorallinavar. santensis(USA, Florida,Brako 8227, NY). a. SEM micrographshowing

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FIG. 6. a. Cross-sectionsof thallus-Type 1 cortex. Phyllopsoraparvifoliavar. breviuscula(Isotype:Cuba,

Crocyniapyxinoides Nyl. (Crocyniaceae)has sora, but they lack a distinct prothallus.Timdal

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FIG. 7. a. Cross-sectionsof thallus-Type 1 cortex. Phyllopsoraparvifoliavar. breviuscula(Isotype:Cuba,

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FIG.8. Cross-sectionsofapothecia.a. Phyllopsoraparvifolia

CHEMISTRY were often, but not consistently, detected, in-

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treatmentthe spot turns bluish-gray.It is Pd+

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o Substances which turn brightyellow after treatment with

* Indicates the darkblue-grayspot characteristic of pannarin.

* Implies an orange fluorescence under long wave UV light.

+ Implies an yellow fluorescence under long wave UV light.

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Pseudocyphellaria.Argopsinis reportedfromAr- widely separatedlocalities in the West Indies,

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P. buettnerivar.glauca, strain .............

P. huettneri var. glauca, strain II ............. . E

P. buettneri var munda ......................... ..

P. glahella .......... ..... ............

P. isidio la........ ...................................

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_Tr,lopc o_f Cncer

FIG. 12. Worldwide range of Phyllopsora.