Bull. Natn. Sci . Mus., Tokyo, Ser. B, 27( I), pp. 1- 10, March 22, 200 I

Taxonomic Notes on Parmelia reparata (Parmeliaceae, Lichenes)
and the Related Species
Syo Kurokawa
Botanic Gardens ofToyama,
42 Kamikutsuwada, Fuchu-machi, Nei-gun, Toyama, 939- 2713 Japan

Abstract Parm elia reparata and the related species were revised taxonomically.
As a result, two new combinations, Canomacu /ina reparata (Stirt. ) Kurok. andParmotrema permacu/atum (Hale) Kurok., are proposed and three new species,
Parmotrema acutatum , P cornigerum and P despectum, are described. Parmotrema
eurysacum (Hue) Hale, which has been variously applied, is clearly characterized.
Key words: Canomaculina, new combinations, new species, Parme/ia reparata,

Pannotrema

Parmelia reparata was described by Stirton ( 1877-1878) on the basis of two
specimens collected in Australia and New Zealand. Hale ( 1965) in 'A monograph of
Parmelia subgenus Amphigymnia' classified this species under series Ornaticolae,
subsection Ornaticolae, section Subftavescentes in the subgenus Amphigymnia andtypified the species by the designation of a lectotype. Nine specimens collected in
Alabama in the United States, Mexico , Australia and New Zealand were cited. Subsequently, Hale (1971) described a new spec ies, Parmelia permaculata, designating as
the holotype one of the specimens cited in Hale ( 1965) as P reparata. When Hale
( 1974) transferred most species of the sub genus Amphigymnia to the resurrected
genus Parmotrema, no combination was proposed for Parmelia reparata.
Recently B. Hale & De Priest ( 1999) treated P reparata as a synonym of
Canomaculina subcaperata (Kremp.) Elix and on the basis of Hale's unpublishednotes reduced Parmelia permaculata to a synonym of Parmotrema eurysacum (Hue)Hale. Specimens reported as Parmelia reparata by Hale in 1965 were filed under P
eurysacum in US (P. T. DePriest pers. comm.). Thus, apparently there has been confusion in understanding Parmelia reparata and Parmotrema eurysacum. In the present study, I attempt to clarify the taxonomic features of Parmelia reparata,
Parmotrema eurysacum and their related species. Based on new examinations ofspecimens of these species, two new combinations, Canomaculina reparata (Stirt.)
Kurok. and Parmotrema permaculatum (Hale) Kurok., are proposed and three new
species, Parmotrema acutatum, P cornigentm and P despectum , are described. In addition, taxonomic features of Parmotrema eurysacum are clarified.
The present study is based principally on speci mens filed under Parmotrema eu-

2

Syo Kurokawa

rysacum in US and those of Parmelia reparata and the related species preserved in
TNS.
Canomaculina reparata (Stirt.) Kurok., comb. nov.
Parmelia reparata Stirt. , Scott. Natural. 4: 201. 1877-78 . Lectotype: Cave
Mountain near Brisbane, Queensland, Australia, F. M. Bailey s.n. (BM).
Parmelia virens Mull. Arg., Flora 69: 255 . 1886. Holotype: Toowoomba,
Queensland, Australia, Hartmann in 1882 (G).
Thallus loosely adnate, mineral gray, coriaceous, I 0- 20 cm broad; lobes irregularly divided, rotund at the apices , broadly crenate and ciliate at the margins, cilia
abundant and dense, 1.5- 2.5 mm long; upper surface glossy, effigurate-maculate,more or less cracked with age, soredia and isidia lacking; lower surface black except
for very narrow blackish brown apical zone, minutely wrinkled, short-rhizinate orpapillate even near the apices, short rhizines less than 0.1 mm long, long rhizines
formed ± in group, thick, about I mm long. Thallus 160- 220 J..lm thick; upper cortex
21-26 J..lm thick, gonidial layer subdiscontinuous, 8- 20 J..lm thick, medulla 100- 140
J..lm thick, lower cortex brownish, ea 13 J..lm thick. Apothecia common, stipitate,
10- 20 mm in diameter, amphithecium rugose, effigurate-maculate, disk brown todark brown, perforate; hymenium 75- 85 J..lm high, asci ea 26 X70 J..lm in size, spores
6-8 X 15-18 J..lm .
Chemistry. Atranorin, salazinic acid and consalazinic acid.
Since the lectotype of Parmelia reparata has an effigurate-maculate upper surface and dimorphic rhizines , the species can be placed in the genus Canomaculina .
Previously, the species was considered as a synonym of C. subcaperata (Kremp.)
Elix by B. Hale & DePriest (1999). However, the lower surface of P reparata is jet
black except for the very narrow dark brown apical zone (for 2- 5 mm from the apex),
whereas it is pale to dark brown for the most part, and blackish brown to black only
near the center on very old lobes in C. subcaperata. In addition, dimorphic rhizines
are more common in C. subcaperata than in C. reparata , in which they are observed
only on certain lobes. Therefore, the present species can be considered as a distinct
species differentiated in Australia and New Zealand, although it is closely related toC. subcaperata.
In 1991 , Kurokawa ( 1991) reported the occurrence of Rimeliella ( Canomaculina) subcaperata from Australia citing two specimens preserved in MEL.
These specimens should be reexamined on the basis of the distinctions between thespecies described above. Thus, C. subcaperata is considered provisionally to be endemic to South America (Brazil, Paraguay and Urugay).
Specimens examined. Australia. Queensland: Brisbane, F. M. Bailey s.n. in
1882 (BM); Maiala National Park, Mt. Glorious, elevation about 700 m, S. Kurokawa
5466 (TNS, US) and 5469, 5473 (TNS); O'Reilly's Place, Lamington National Park,
E. Dahl s.n. (TNS). New South Wales: Tropical rain forest , Dorrigo Mt. , Dorrigo Na-

Taxonomic Notes on Parmelia reparata (Parme li aceae, Lichenes) and the Related Species

3Fig. I.

Holotype of Parmotrema acutatum Kurok. (Bar= 1 cm).

tional Park, elevation about 800 m, S.Kurokawa 5244 (TNS). New Zealand. N. Island,
R. Gobby s.n. (BM). Ashburton , S. Island, H. H . Allan s.n. in 1920 (BM).

Parmotrema acutatum Kurok ., sp. nov.
Species habitu cum Parmotremate eurysaco optime congruens sed differt lobis
eciliatis et superficie superiore distincte maculata.
Chemistry. Atranorin, salazinic acid and consalazinic acid.
Type. Mexico. Nayarit: Near Tepic, Rt. 15, km 865, on tree, M. Wirth 26 holotype in US and isotypes in US and TNS.
Thallus loosely adnate, membranaceous, grayish glaucous, 6-10 cm broad; lobes
irregularly divided, eciliate, 5- 10 mm wide, conspicuously laciniate at the margin,
laciniae 5- 15 mm long, often subpalmately divided, acuminate towards the apices,pale ochre and erhizinate on the lower surface; upper surface glossy, distinctly maculate, soredia and isidia lacking; lower surface naked and pale brown near the apices(for 3- 10 mm) and black and very sparsely rhizinate near the center, rhizines black,coarse and short (less than I mm) , papillae commonly formed on black lower surface.
Thallus 180-240 Jlm thick; upper cortex 20 Jlm thick, gonidial layer continuous,
20- 30 Jlm thick, medulla 110-160 Jlm thick, lower cortex brownish, ea 25 Jlm thick.
Apothecia not seen.

4

Syo Kurokawa

The present new species closely resembles P eurysacum in that they both haveloosely adnate membranaceous thalli with subpalmately divided laciniae and produce
atranorin, salazinic acid and consalazinic acid. However, this species entirely lacks
cilia, whilst P eurysacum has long cilia, although sometimes they are quite rare.
In the present species the maculae on the upper surface are very distinct andoften connected to each other and appears reticulate. Thus, the present species might
be regarded as a member of the genus Rimelia. However, its rhizines are very sparse,coarse and thick. Such rhizines are typical of species in the genus Parmotrema. In
contrast, rhizines are rather dense even near the lobe tips and only moderate thick in
Rimelia . In addition, the present species lacks cilia, while all species of Rimelia havecilia, although they are sometimes rare. Thus, the present species is classified in thegenus Parmotrema here.
The present species may be related to P latissimum (Fee) Hale, since both
species lack cilia and produce atranorin and salazinic acid. However, P acutatum isclearly distinguished from the latter by the presence of subpalmately divided laciniae.
The species is known only from the Pacific side of Mexico at present.
Specimen examined. Mexico. Jalisco : Rt. 15, km 839, in lava bed, on tree,
M.Wirth 70 (US).

Parmotrema cornigerum Kurok ., sp. nov.
Species cum thallo ut in Parmotremate eurysaco sed ab hac species lobis eciliatis et thallo acidum protocetraricum continenti differt.
Chemistry. Atranorin and protocetraric acid.
Type. Mexico. Chiapas: Scrub deciduous forest bordering rim of El Sumidero,
El Sumidero canyon, north of Textula Gutierrez, on deciduous trees , elevation
1220 m, M. E. Hale 20078 & T. R. Soderstrom - holotype in US .
Thallus loosely adnate on bark of trees, membranaceous, glaucous gray, 8- 15cm broad; lobes irregularly divided, eciliate, 5- 12 mm wide, conspicuously laciniate,
laciniae usually dichotomously or subpalmately divided, 6- 14 mm long, canaliculate
below and becoming terete near the apices ; upper surface glossy, emaculate, rugose
or more or less foveolate on older lobes, soredia and isidia lacking; lower surface distinctly wrinkled, pale buff or pale brown in quite wide zone (for about 1.5- 2.5 andoften more than 3 cm), black only on older lobes; rhizines rather rare, formed only on
black lower surface, coarse and short (less than 0.5 mm long). Thallus 140- 190 ,urn
thick; upper cortex 16- 25 ,urn thick, gonidial layer continuous, 13- 15 ,urn thick,medulla 130-150 ,urn thick, lower cortex pale brown, ea 15 ,urn thick. Apothecia stipitate, I 0- 16 mm in diameter, amphitheci urn distinctly rugose, emaculate, disk pale
brown to brown, perforate; hymenium 90- 100 pm high, asci subcylindrical ,22- 24X 50- 70 ,urn in size, spores ellipsoidal or more less ovoidal, 9- 10 X20- 21 ,urn.
The present species may be confused with P eurysacum because they both form
similar subpalmate laciniae. However, its lobes lack cilia and their laciniae becomeTaxonomic Notes on Parmelia reparata (Parmeliaceae, Lichenes) and the Related Species

Fig. 2.

5

Ho lotype of Parmotrema cornigerum Kurok . (Bar = I cm)

terete towards the apices. In addition, the medulla is P +orange red, containing protocetraric acid rather than sa lazi nic and consalazinic acids. Spores are much larger(9- IO X20- 21 Jl.m) than those of P eurysacum (7- 9 X 14- 16 Jl.m).
This species resembles Rimelia cetrata (Ach.) Hale & A. Fletcher, which also
forms distinct laciniae. However, R. cetrata is sparsely to densely ciliate and has areticulately maculate upper surface . The main secondary product of R. cetrata is
sa lazinic acid.
This new species is known only from Mexico at present.
Specimen examined. Mexico. Chiapas: Scrubby deciduous pasture, mist forest, Road to El Suspiro, 5-7 km north of Berriozabal , on deciduous trees , elevation
920 m, M. E. Hale 20115 & T. R. Soderstrom (US).

Parmotrema despectum Kurok. , sp. nov.
Species cum thallo ut in Parmotremate eurysaco sed ab hac species thallo plus
minusve coriaceo, laciniis aubpalmatim ramosis destituto et ciliis raris sparsisve et
brevibus (1- 3 mm longi s) differt.
Chemistry. Atranorin , salazinic acid and consalazinic acid.
Type. U.S.A. Virginia: Along Rt. 623 near Rt. 30 l , King George Co., C. F.
Reed 59408- holotype in US.
Thallus adnate, coriaceous, mineral gray to glaucous gray, about 10 cm broad, irregularly divided; lobes 2.5- 10 mm wide, crenate to gnawed at the margin, sometimes with rotund lobules I X 1 to 2X3 mm in size; upper surface glossy, sometimes
faintly maculate, roughly rugose on older lobes, soredia and isidia lacking; cilia rare

6

Syo Kurokawa

Fig. 3.

Holotype of Parmotrema despectum Kurok. (Bar= I cm ).

or sparse, simple or very rarely branched, 1- 3 mm long; lower surface black, sometimes ivory-mottled near the margin; rhizines simple, dense, 1- 2 mm long. Thallus
260- 300 f-Lm thick; upper cortex ea 50 f-Lm thick, gonidial layer subcontinuous, ea
25 f-Lm thick, medulla 190- 300 f-Lm thick, lower cortex dark brown, ea 20 f-Lm thick.
Apothecia sessile, 3- 10 mm in diameter, amphithecium more or less rugulose, whitemaculate, disk dark brown to blackish brown, perforate, often fissured in mature
stage; hymenium ea 55 f-Lm high , asci subclavate, ea 33 X 53 f-Lm in size, spores

6- IO X 10- 14 f-Lm.
The present species has been confused with P eurysacum , probably because
both species have ciliate lobes, produce atranorin , salazinic acid and consalazinicacid and show similar distribution pattern in southeastern United States and Mexico.
However, it can be distinguished from the latter by I) the::t:coriaceous adnate thallus ,2) the crenate to gnawed margins of lobes, 3) the rare to sparse short cilia (1 - 3 mmlong) and 4) the absence of subpalmately divided lacinjae. The thallus of this species
is rather coriaceous, reflecting thicker lobes (260-300 f-Lm thick) compared with P
eurysacum (150-160 f-Lm thick) . Spores are also somewhat smaller (6- 10 X 10- 14
f..Lm) than in P eurysacum (7-9 X 14- 16 f..Lm) .
The present species is very common in the southern and southeastern UnitedStates, with its range extending to northern Mexico where P eurysacum is muchTaxonomic Notes on Parmelia reparata (Parmeli aceae, Lichenes) and the Related Species

7

more common.
Specimens examined. U.S.A. Maryland: Baltimore, on tree trunk, C. C. Plitt
300 (US). West Virginia: Near town of Cucumber, McDowell Co., on red oak, M. E.
Hale 11505 (US). South Carolina: Near town of Spartanburg, Spartanburg Co., on
hickory, M. E. Hale 7685 (US). Ohio: Springfield, ex Willey (US). Illinois: Bell
Smith Springs, Pope Co. , on Quercus stellata in dry woods, A. C. Skorepa 4256
(US); Canton, 1. Wolf s.n. (US). Iowa: On trees, Fayette Co. , B. Fink s.n. (three packets, US). Kentucky: Olympia, Bath Co., on living oak, M. 1. Alien 212 (US). Missouri: Near town of Rayborn, Wright Co., on black oak, M. E. Hale 4259 (US); Near
town of Viola, Barry Co., on elms, M. E. Hale 2580 (US). Arkansas: Ozark National
Forest, Blanchard Springs, Stone Co., elevation 750ft, D. Demaree 28860 (US); Near
town of White Rock Mtn. , Franklin Co. , on Quercus alba, M. E. Hale 3674 (US);Near town of Winslow, Washington Co., on oak branches, M. E. Hale 2659 (US). Oklahoma: Near town of Ardmore, Carter Co. , on post oak, M. E. Hale 4892 (US).
Texas: Gillepse Co., G. Jermy 152456 (US); On state 1604, 4 miles west of the oldBlanco Road, Burnet Co., live oak and mesquite, C. Fox T-42 (US) . Arizona: Chiricahua Mts. Cave Creek, near headquarters, Southwestern Research Station, limestone
outcrops and floodplain, Cochise Co. , 6000 ft, Weber, Shushan & Anderson S24741(US). Mexico. Zacatecas: On El Bote that overlooks Zacatecas and the new technological college, C. Fox M-314 (US). Michoacan: Mal Pais de Curucu, old lava flowbadlands 12 km west of Uruapan, on lava, M. Wirth 340 (US). Baja California Sur:Top of the ridge of Sierra Laguna and east of Cerro Valde, Quercus tuberculata forest, on granite, T. H. Nash Ill 39614 (distributed as Nash: Lich. Exs., no. 278) (US).
Michoacan: Dry pine-oak pasture on rocky hillside, km 223 on highway 190, about
61 km northwest of Zitacuaro, on rocks, elevation 2131 m, M. E. Hale 20861 &Soderstrom (US).
Parmotrema eurysacum (Hue) Hale, Phytologia 28: 336. 1974.
Parmelia eurysaca Hue, Nuov. Arch. Mus. Paris, ser. 4, 1:194. 1899.
Chemistry. Atranorin, salazinic acid and consalazinic acid.
The present species is characterized by 1) loosely adnate membranaceous thalli( 150- 160 .urn thick), 2) emaculate upper surface, 3) lobes subentire or broadly crenate at the margins, 4) subpalmately divided long laciniae, 5) long, often branchedcilia (1.5- 5 mm long), which may be sparse to dense, 6) medium-sized spores(7-9 X 14- 16 .urn) and 7) the production of atranorin, salazinic acid and consalazinicacid. Lobes are irregularly divided but elongated and the margin may be subentire or
somewhat crenate. Usually the cilia are quite long and are highly variable in density.
The thallus typically forms subpalmately-divided, long laciniae. Occasionally thelaciniae are short lobules with acuminate tips along the lobe margins.
The lower surface of lobes is black with a brown naked zone (for 3- 5 cm) near
the apices and, often, white-mottled along the margin. The laciniae are broadly white

8

Syo Kurokawa

or white mottled and naked on the lower surface. Rhizines are formed on the black
lower surfaces and are moderate to dense. They are mostly simple and 1- 3 mm long.
This species is quite common in Mexico with its range extending north to southeastern United States. It is reported from Brazil for the first time.
Representative specimens examined. U.S.A. Alabama: 5 miles W of Okatchie,Calhoun Co., sandstone ridge in open oak woods, M. E. Hale 33848 (US). Arkansas:
MacNab. , Hemmpstead Co. , old tree tops, elevation 320, D. Demare 44674 (US).
Texas: Austin, Cuero Co. , E. Whitehouse 2286 (US); Near town of Tyler, Smith Co.,
M. E. Hale 5444 (US); Frio Canyon, 11 miles north of Leaky, Acal Co. , L. Hubricht
13175 (US). Mexico. Tamaulipas: Vicinity of El Milagro, Cerro Zamora, Sierra de
San Carlos, H. H. Bartlett 11098 (US); Near Rancho de Cielo above Gomez Farias,
A. J. Sharp 507 (US). Veracruz: Scattered shade trees in coffee plantation, 64 km
southwest of junction of highways 140 and 155, northeast of Huatusco, elevation
1310 m, M. E. Hale 19535 & Soderstrom (US); Obere Barranea de Fenampa, 9001000 m, J. A. Purpus 55 (US). San Luis Potosi : Alvarez, alt. 8000 ft. , E. Pal mer 466(US). Hidalgo: Cuyamaloya, an Quercus, C. G. Pringle (distributed as Cryptog. Exs.
ed. Mus. Hist. Natur. Yindobonensi, no . 4447) (TNS , US). Sierra de Pachuca, alt.
2520 m, X. Madrigel 145 (US); El Hi! oche, M. Martinez s.n. & 71 (US); Cuyamaloya, on rocks, C. G. Springle 10753 (US). Oaxaca: Mature pine-oak mist forest ,Trail leading to Cerro San Felipe, overlooking Oaxaca, on deciduous trees, M. E.
Hale 20692 & T. R. Soderstrom (US , TNS); The same locality, M. E. Hale 20709,
20785 & Soderstrom (US); Open dry deciduous forest on mountainside, 132 km
northwest of Oaxaca on highway 190, elevation 2380 m, M. E. Hale 20829 & T. R.
Soderstrom (US). Mexico: Sierra de Las Cruces, C. G . Pringle 168 (US); North ofAcambay, R. F. Cain 27597a (US). Jalisco: San Sebastian, Sierra Madre Occidental ,on living tree trunk, Y. Mexia 1904 a (US). Michoacan: Mature pine forest in mountains, km 248 on highway 190, 65 km east of Morelia, on deciduous trees, elevation
2560 m, M. E. Hale 20988 & T. R. Soderstrom (US); On conifers, the same locality,
M. E. Hale 21033 & T. R. Soderstrom (US , TNS); Cerro Azul , Morelia, G. Arsene
3 730, 3984 (US); Mal Paris de Curucu, old lava flow badlands 12 km west of Uruapan, M. Wirth 337 (US). Brazil. Minas Gerais: Serra da Mantiqueira, oberhalb vonYi la Monte Yerde, etwa 30 km ostlich von Camanducaia, 1970 m, K. Kalb & G . Plost(distributed asK. Kalb: Lich. Neotropici 144) (US).

Parmotrema permaculatum (Hale) Kurok ., comb. nov.
Parmelia permaculata Hale, Phytologia 21 : 425 . 1971. Holotype : Mexico, Veracruz, Open pasture, 9 km east of Jalapa along highway 140, on deciduous trees, elevation 1240 m, M. E. Hale 19406 & T. R. Soderstrom (US; isotype in TNS).
Thallus adnate, coriaceous, whitish gray, up to 20 cm broad; lobes irregularly divided, with rotund apices, broadly crenate and ciliate at the margin, 8- 15 mm wide ,lacking soredia, isidia and elongated laciniae; upper surface glossy, strongly macu-

Taxonomi c Notes on Parme/ia reparata (Parmeliaceae, Li chenes) and the Related Species

Fig. 4.

9

Ho lotype of Parme/ia permaculata Hale (Bar = I cm ).

late, rugose and reticulately cracked on older lobes; cilia more or less erect, moderate
to dense, sometimes branched, 1- 2 mm long; lower surface black but dark brown andnaked near the apices (for about 5- 7 mm), sometimes white-mottled along the margin; rhizines moderate, uniformly elongate, 1- 2 mm long. Thallus 110- 140 ,um thick;upper cortex variable in thickness, I 0- 20 ,um thick, gonidial layer subcontinuous,
7- 20 )..lm thick, medulla 65- 120 ,um thick, lower cortex pale brown, ea I 0 ,um thick.
Apothecia sessile, 5- 15(20) mm in diameter, amphithecium reticulately maculate,smooth to reticulately rugose, disk pale brown, perforate; hymenium ea 70 ,um high,asci subclavate, ea 23 X 52 )..lm in size, spores 8- 10 X 13- 16 ,um.
Chemistry. Atranorin, salazinic acid and consalazinic acid.
The present species is confused easily with Canomaculina subcaperata because
these species form similar lobes that are strongly maculate on the upper surface andrather densely ciliate along the margin of lobes. Both species produce atranorin,salazinic acid and consalazinic acid. P permaculatum was reduced to a synonym of
C. subcaperata by B. Hale & De Priest ( 1999). However, the dimorphic rhizines,
which are characteristic of Canomaculina , are lacking in the present species, al though some black verrucae are formed usually on the black lower surface. Thus thepresent species is considered to belong to Parmotrema rather than Canomaculina .
This species may be also confused with P eurysacum. However, compared withthe latter species, it has rather coriaceous thalli composed of lobes with strongly maculate upper surfaces that lack subpalmately divided laciniae.
The species is known only from Alabama in the United States and Veracruz in10

Syo Kurokawa

Mexico at present.
Specimens examined. U.S.A. Alabama: Mobile Co., C. Mohr s.n. (two packets)(US). Mexico. Veracruz: Mirador, an Yuccastammen, ea. I 000 m, T. A. Purpus 210(US , TNS); Open pasture, scattered Cactus and Acacia, 9 km east of Ja1apa along
highway 140, on deciduous trees, elevation 1240 m, M. E. Hale 19378 & T. R. Soderstrom (US, TNS); the same, M.E. Hale 19394 & T. R. Soderstrom (US); Scatteredshade trees in coffee plantation, 64 km southwest of junction of highways 140 and
155, northeast of Huatusco, elevation 1310 m, M. E. Hale 19529 & T. R. Soderstrom(US).

Acknowledgements
I express my sincere thanks to Dr. P. T. DePriest of the Department of Botany,
Smithsonian Institution in Washington D.C. and Dr. H. Kashiwadani of the Department of Botany, National Science Museum, Tokyo for the loan of specimens. I amvery grateful to the curators of the Botany Department, The Natural History Museum, London and of the Conservatoire et Jardin Botaniques de la Geneve for the loanof type specimens. I am also grateful to Dr. K. H. Moon of Sookmyung Women 's
University for making TLC tests and Dr. P. T. DePriest of the Smithsonian Institution
for revising the English text.

References
Eli x, J. A. 1997. The lichen genera Canomacu/ina and Rime/iel/a (Ascomycot ina, Parmeliaceae). Mycotaxon, 65 :475-479.
Hale, B. W. & P. T. De Priest 1999. Mason E. Hale's list of epithets in the Parmelioid genera . Th e Bryologist, 102 : 462- 572.
Hale, M. E. 1965. A monograph of Parmelia subgen us Amphigymnia. Contr. US. Nat. Herb. , 36: 193358.
Hale, M. E. 197 1. Parmelia permaculata, a new lichen from Alabama and Mexico. Phyrologia , 21 :45.
Hale, M. E. 1974. New combinations in the lichen genus Parmotrema Massalongo. Phytologia , 28:334- 339.
Kurokawa, S. 199 1. Rimeliella, a new li chen genus re lated to Rimelia of the Parmeliaceae. Ann. Tsukuba
Bot. Gard. , 10: 1- 14.
Stirton, J. 1877- 78. On certa in lichens belonging to the gen us Parme/ia. Scott. Natural., 4: 200- 203.