BULLETIN OF THE UNITED STATES NATIONAL MUSEUM 


CONTRIBUTIONS FROM THE UNITED STATES NATIONAL HERBARIUM 


VoLuME 36 


LICHENS 
ALGAE 


Mason E. Hate, Jr., Wirt1am RANDOLPH TayYLor, 
Micuae.t WirtH, Syo KuroKAwa 


SMITHSONIAN INSTITUTION PRESS ¢ WASHINGTON, D.C. © 1968 


Publications of the United States National Herbarium 


The United States National Herbarium, which was founded by 
the Smithsonian Institution, was transferred in the year 1868 to the 
Department of Agriculture and continued to be maintained by that 
department until July 1, 1896, when it was returned to the official 
custody of the Smithsonian Institution. The Department of Agri- 
culture, however, continued to publish the series of botanical reports 
entitled “Contributions from the United States National Herbarium,” 
which it had begun in the year 1890, until, on July 1, 1902, the National 
Museum, in pursuance of an act of Congress, assumed responsibility 
for the publication. The first seven volumes of the series were issued 
by the Department of Agriculture. 
Frank A. Tayvor, 
Director, United States National Museum. 


il 


CONTENTS 


Parr 1. A revision of the South American species of Parmelia 
determined by Lynge. By Mason E. Hale, Jr. 
(Published May 18,1960) .......... 

Part 2. Marine algae from the tropical Atlantic Ocean: 
V. Algae from the Lesser Antilles. By William 
Randolph Taylor. (Published January 31, 
1962)... 2 

Parr 38. The lichen family Graphidaceae in Mexico. By 
Michael Wirth and Mason E. Hale, Jr. (Published 
December 6, 1963) ........2.22. 

Part 4. Studies on Parmelia subgenus Parmelia.* By Mason 
EK. Hale, Jr., and Syo Kurokawa. (Published 
August 26,1964)... ......020224282, 

Part 5. A monograph of Parmelia subgenus Amphigymnia. 
By Mason E. Hale, Jr. (Published April 8, 1965) . 


Page 


1-41 


43-62 


63-119 


121-191 


193-358 


*Errata: Line 23, page 164, should read ‘‘24. Rhizines moderately branched; 


lobes subrotund, often whitish .. .” 


iil 


UNITED STATES NATIONAL MUSEUM 


CONTRIBUTIONS FROM THE UNITED StaTEs NATIONAL HERBARIUM 


VotumeE 36, Part 1 


A REVISION OF THE SOUTH AMERICAN 
SPECIES OF PARMELIA DETERMINED 
BY LYNGE 


By Mason E. Hate, Jr. 


BULLETIN OF THE UNITED STATES NATIONAL Museum 


SMITHSONIAN INSTITUTION e¢ WASHINGTON, D.C, e 1960 


A REVISION OF THE SOUTH AMERICAN SPECIES OF 
PARMELIA DETERMINED BY LYNGE 


Mason E. Hate, Jr. 


Introduction 


Although the Norwegian lichenologist Bernt Lynge is remembered 
as the foremost authority on arctic lichens, early in his career he 
published several articles of fundamental importance on tropical 
lichens, especially in the genera Anaptychia-and Parmelia. His 
major work (1914) involved the determination of the rich and well- 
prepared Parmelia collections, now preserved at Stockholm, of the 
various Regnell expeditions from 1892 to 1902 to Brazil and Paraguay. 
A second short paper (1917) merely added the descriptions of two new 
species, and a third paper (1925) summarized the determinations of 
miscellaneous Brazilian Parmelias at Uppsala, Stockholm, and 
Berlin. 

Altogether Lynge identified a total of 110 taxa, 61 of them new to 
science, and gave exhaustive Latin descriptions of nearly all the 
species and excellent photographs of 30 taxa. It has been possible 
to reexamine 99 of these taxa; specimens of the remaining 11 were 
destroyed at Berlin or have otherwise not been located. The purpose 
of this report is to revise the determinations of these taxa with 
particular attention to adequate typification and analysis of chemical 
components. A full understanding of Lynge’s species is essential 
before a monographic treatment of Parmelia in South America can 
be attempted. 

The Regnell collections contained an extraordinarily large number 
of endemic species, many of which have never been collected again. 
Lynge, although on the whole he was a conservative taxonomist, 
wisely described these as new species instead of ascribing them to 
older names. At the same time the thoroughness of his work is at- 
tested by the fact that I have found it necessary to describe only 
one additional new species, Parmelia microdactyla. Most of my 
effort was expended in correcting misidentifications and synonymy 
stemming from Lynge’s failure to typify the species correctly. 

For the loan of valuable type specimens and other material, I am 
indebted to Dr. H. des Abbayes, Université de Rennes, Rennes, France; 
Dr. Sten Ahlner, Naturhistoriska Riksmuseum, Stockholm, Sweden; 
Dr. Charles Baehni, Conservatoire et Jardin Botanique, Geneva, 

1 


2 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Switzerland; Dr. M. Skytte Christiansen, Botanical Museum, Leiden, 
Netherlands; Dr. Ilmari Hiitonen, University of Helsinki, Helsinki, 
Finland; Dr. I. Mackenzie Lamb, Farlow Herbarium, Cambridge, 
Massachusetts; Dr. Joseph Poelt, Botanische Staatssammlung, 
Munich, Germany; Dr. Rolf Santesson, Botanical Museum, Uppsala 
University, Uppsala, Sweden; and Dr. Antero Vaarama, Botanical 
Institute of the University, Turku, Finland. 


Chemistry of the Species 


The chemical components of each specimen were determined by 
means of the standard microchemical crystal tests of Asahina (1954), 
chromatography (Wachtmeister, 1956), and fluorescence analysis 
(Hale, 1956). Chromatography is required to separate the pigmented 
substances; fluorescence analysis at 3,600 A. is the easiest method of 
detecting the presence of alectoronic acid in the medulla (by a bright 
white fluorescence) and lichexanthone in the cortex (by a brilliant 
orange-yellow fluorescence). Color tests supplementing the micro- 
chemical tests were made under a low power binocular directly on the 
thallus and medulla with fresh reagents. Luynge made a number of 
faulty chemical color tests which resulted in some misidentifications 
and in some species being described as new unnecessarily. 

Thirty lichen substances, including 17 positively identified and 13 
unknown, were demonstrated. A list of the 17 positively identified 
acids and the species in which they occurred is given below. Changes 
in the nomenclature of Lynge’s names should be checked in the list of 
synonyms and misidentifications on p. 39. 


1. Alectoronic acid: 2, Atranorine—Continued 
P. bahiana Nyl. P. flava var. subdichotoma Lynge 
P. laongit Lynge P. malmet Lynge 
P. latissima f. microspora Lynge P. minima Lynge 
P. melanothriz (Mont.) Vain. P. portoalegrensis Lynge 
P. melanothriz f. microspora Lynge P. regis Lynge 
P. rigida Lynge P, regnelliit Lynge 
P. subproboscidea Lynge P. regnellii f. arida Lynge 
P. subrugata var. arcuata Lynge P. rutidota Hook. & Tayl. 
P. wainit A. L. Smith P. silvatica Lynge 
2. Atranorine: P. silvatica var. pinnata Lynge 

Present in all taxa except: P. silvatica var. radiata Lynge 
P. abstrusa Vain. | P. viridescens Lynge 
P. abstrusa f. laevigata Lynge 3. Barbatic acid: 
P. brasiliana Nyl. P. digitata Lynge 
P. brasiliana var. novella (Vain.) ° 7 

L P, regnellit Lynge 

ynge a 

P. chapadensis Lynge P. uleana Mill. Arg. 
P. crustacea Lynge 4, Divaricatic acid: 
P. flava var. stellata Lynge P, rupicola Lynge 


HALE—REVISION OF PARMELIA SPECIES 3 


5. Fumarprotocetraric acid: 


P. rutidota Hook. & Tayl. 
P. rutidota f. filizans Lynge 


6. Gyrophoric acid: 


7. 


. acartospora Zahlbr. 
. coronata Fée 
. marginalis Lynge 
. minarum Vain. 
. pluriformis Nyl. 
. riograndensis Lynge 
. rudecta Ach. 
. sancti-angelit Lynge 
. zanthina (Mill. Arg.) Vain. 
Lecanoric acid: 
P. paraguartensis Lynge 
P. rissoensis Lynge 
P. tinctorum Nyl. 


VUVVVVBMTDVNWY 


8. Lichexanthone: 


9. 


10. 


11. 


12. 


P. brasiliana Ny). 
P. brasiliana var. novella (Vain.) 
Lynge 
. crustacea Lynge 
. malmei Lynge 
. minima Lynge 
. regis Lynge 
. stlvatica Lynge 
. silvatica var. pinnata Lynge 
. silvatica var. radiata Lynge 
Norstictic acid: 
P. abstrusa Vain. 
P. abstrusa f. laevigata Lynge 
Olivetoric acid: 
P. fragilis Lynge 
P. revoluta Floerke 
Perlatolic acid: 
P. wainioana Lynge 
Protocetraric acid: 
P, amazonica Nyl. 
P. consimilis Vain. 
P. crustacea Lynge 
P. cyliphora (Ach.) Vain. 
P. fatiscens Lynge 
P. fistulata Tayl. 
P. flava var. stellata Lynge 
P. flava var. subdichotoma Lynge 
P. latissima Fée 
P. latissima var. 
Kremplh. 
P. latissima var. minima Lynge 
P. leucozantha Mill. Arg. 
P. malmei Lynge 
P. minima Lynge 
P. rutidota Hook. & Tayl. 


ih cde: 


corniculata 


12. 


13. 


14. 


15. 


16. 


Protocetraric acid—Continued 
P. saccatiloba Tayl. 

P. silvatica Lynge 

P. silvatica var. pinnata Lynge 
P. silvatica var. radiata Lynge 


Protolichesteric acid: 

P. canaliculata Lynge 

P. microsticta Mill. Arg. 

P. riograndensis Lynge 

P. zanthina (Mill. Arg.) Vain. 


Salacinic acid: 

P. cetrata Ach. 

P. cetrata var. corniculata Mill. Arg. 

. cetrata ssp. radiata Lynge 

. cinerascens Lynge 

. continua Lynge 

. cristifera Lynge 
kamtschadalis var. 

(Mey. & Flot.) Nyl. 

latissima Fée 

magna Lynge 

. microdactyla Hale 

. nylandert Lynge 

. radians Lynge 

. rupta Lynge 

Stictic acid: 

P. conspersa (Ach.) Ach. 

P. eciliata (Nyl.) Nyl. 

. longiconida Lynge 

. portoalegrensis Lynge 

. regnellit Lynge 

. scrobicularis Kremplh. 

. uleana Mill. Arg. 

Usnic acid: 

. abstrusa Vain. 

. absirusa f. laevigata Lynge 

. chapadensis Lynge 

. conspersa (Ach.) Ach. 

. cristifera Tayl. 

cyliphora (Ach.) Vain. 

. fatiscens Lynge 

flava var. stellata Lynge 

flava var. subdichotoma Lynge 

. leucozantha Miill. Arg. 

magna Lynge 

. microdactyla Hale 

nylanderi Lynge 

portoalegrensis Lynge 

radians Lynge 

. regnellit Lynge 

. regnellii f. arida Lynge 

. rutidota Hook. & Tayl. 


americana 


Tuy yy DWV 


vu 


yuvyyuyyyuyyyyuyuyy 


4 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


16. Usnie acid—Continued 17. Vulpinic acid: 
P. rutidota f. filizans Lynge P. cornuta Lynge 
P. wleana Mill. Arg. P. persulphurata Nyl. 
P. zanthina (Mill. Arg.) Vain. P. sulphurata Nees & Flot. 


Unknown colorless substances were found in the following species: 


. osseo-albida Lynge 

. palmarum Lynge 

. regis Lynge 

. regnellit Lynge 

. regnellit f. arida Lynge 
riograndensis Lynge 

. saccatiloba Tayl. 

. saccatiloba f. membranacea Lynge 
. subregressa Lynge 

. zahlbrucknert Lynge 


P. annae Lynge 

P. brasiliana var. novella (Vain.) 
Lynge 

. capitata Lynge 

. ceracea Lynge 

. chapadensis Lynge 

. digitata Lynge 

. gracilis (Mill. Arg.) Vain. 

. melanothriz (Mont.) Vain. 

. mesotropa Miill. Arg. 


Vu V VV 


vu 


Unknown pigments were discovered in the following species: 


. regnellas Lynge 

. regnellii f. arida Lynge 

. stlvatica Lynge 

. silvatica var. pinnata Lynge 
. stlvatica var. radiata Lynge 
. uleana Mill. Arg. 


. chapadensis Lynge 

. cornuta var. crocea Lynge 
. crustacea Lynge 

. lindmaniw Lynge 

. malmet Lynge 

. merrillia Lynge 

. minima Lynge 


wu yyw 


Tey VD 


List of Species 


Lynge originally arranged his lists of 110 taxa in phylogenetic 
order. I have, for convenience, rearranged the species in alphabetical 
order and have numbered them consecutively. No descriptions are 
repeated from Lynge since he described meticulously nearly all the 
species in his 1914 article in Arkiv fér Botanik, which is readily 
available in most larger libraries. I have, however, included sup- 
plemental information on the nomenclatural types, synonymy, 
range, color reactions and chemical components, and pertinent com- 
ments on the status of the species and their relatives. I have not 
attempted to improve or expand Lynge’s excellent, though outdated 
key, for our knowledge of the Parmelias of South America is so 
incomplete that no key can do justice to the genus now. 

The Malme specimens from the Regnell collections, all of which are 
preserved at Stockholm, are cited only by number or date, since 
Lynge gives complete label data for these in his lists. Holotypes and 
lectotypes are cited in full. Specimens collected by Dusén, Henschen, 
or Regnell, mostly preserved at Uppsala, are also cited in full. 

Lynge did not designate holotypes for his new taxa in his publica- 
tions, but he did write “originaleksemplar’”’ on one herbarium packet 


HALE—REVISION OF PARMELIA SPECIES 5 


if two or more specimens were included in the original concept. I 

have selected such packets as lectotypes except in the case of P. 

zahlbruckneri Lynge (see p. 38). A summary of the new taxa reduced 

to synonymy and corrected identifications is given at the end of 

this list (p. 39). 

1. Parmelia abstrusa Vain. Acta Soc. Faun. & Fl. Fenn. 7:64. 1890.—Lynge, 
1914, p. 145, pl. 5, figs. 8, 9; 1925, p. 84. 

Lectotype: Caraga, Minas Gerais, Brazil, Vainio 1347 (TUR). 

Range: Brazil, Colombia, West Indies, Japan. 

Additional specimens examined: Malme 1481B, 1537, 1857, 1865B, 
1884, 2745, 2749B. 

Reactions: Thallus K—, medulla K+ red, C—, KC—, P+ orange- 
red, norstictic and usnic acids present. 

The specimens seen by Lynge fall within the range of variation of 
Vainio’s species, although the thickness of the thallus is quite variable. 
A very close relative with thinner lobes is P. jamaicensis Vain., which 
also contains norstictic acid and usnic acid, and probably intergrades 
with P. abstrusa in the West Indies. Both species appear to be 
common on soil and rocks as well as on tree bark. Parmelia micro- 
blasta Vain., judging from the poor type material, is in this same group 
of isidiate species with norstictic acid but differs in lacking usnic acid. 
2. Parmelia abstrusa f. laevigata Lynge, Ark. Bot. 13, No. 13:147. 1914. 

Parmelia subabstrusa Gyel. Repert. Sp. Nov. 29:288. 1931. Based on P. 
abstrusa f. laevigata Lynge. 

Lectotype: Bocca da Serra, Serra da Chapada, Mato Grosso, 
Brazil, Malme, June 15, 1894 (S). 

Range: Known only from the type locality. 

Reactions: Thallus K—, medulla K+ red, C—~, KC—, P+ orange- 
red, norstictic and usnic acids present. 

The lack of isidia and the conspicuous coronate apothecia set this 
entity quite apart from P. abstrusa f. abstrusa. In fact, we may well 
wonder why Lynge did not recognize it as a new species. The adop- 
tion of Gyelnik’s name as a distinct species is recommended, although 
it does not clear up the confusion surrounding this difficult group. 
Two specimens seen by Lynge (Malme 2445 and March 7, 1894) 
react K— in the medulla and may be classified either as a chemical 
strain of P. subabstrusa or as an undescribed species more closely 
related to P. relicina Fr. 

3. Parmelia acanthifolia Pers. in Gaudich. Voyage Uranie, Bot. 197. 1826.— 
Lynge, 1925, p. 84. PuaTE 1 

Type: Rio de Janeiro, Brazil, Gaudichaud 13 (P). 

The specimen determined by Lynge in the Berlin Museum is pre- 
sumably destroyed. Persoon’s type is apparently an abnormal form 


6 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


of P. cetrata Ach. The upper cortex is less reticulately rimose than 

expected, but the plant contains atranorine and salacinic acid and has 

spores (8-11 12-16 ») comparable to P. cetrata. Vainio’s determina- 

tion of P. acanthifolia from Brazil (Lich. Bras. Fs. 737) is based on 

P. subcaperata Kremplh. 

4, Parmelia acariospora Zahlbr. Denkschr. Akad. Wiss. Math. Naturw. Wien 
83:169. 1909.—Lynge 1914, p. 105. 

Type: Barra Mansa, Itapecirica, Séo Paulo, Brazil, Schaffner & 
Wettstecn (BPI, isotype). 

Range: Brazil. 

Additional specimens examined: Malme 2509B, 2532B*, 2545. 

Reactions: Thallus K-+ yellow, medulla K—, C+ rose, KC+ red, 
P—, atranorine and gyrophoric acid present. 

This uniform species is easily recognized by its thin fragile thallus 
and delicate isidia. Parmelia minarum Vain. is similar in chemistry 
and appearance but has a more ashy color and firmer lobes. Parmelia 
granatensis Nyl. and P. chileana Nyl. are also related to this species, 
but their exact status is unsettled. 


5. Parmelia amazonica Ny]. Flora 68:611. 1885—Lynge, 1914, p. 101. 


Type: Santarem, Amazon River, Bahia, Brazil, Spruce 111 (H). 

Range: Florida, West Indies, Central America, Brazil. 

Additional specimen examined: Malme 2408. 

Reactions: Thallus K+ yellow, medulla K—,C—, KC+ rose, P+ 
orange-red, atranorine and protocetraric acid present (not proved 
microchemically in the type specimen). 

Parmelia amazonica is distinguished by the presence of protoce- 
traric acid. Parmelia consimilis Vain. of similar chemical constitu- 
tion has thicker crowded isidia. 


6. Parmelia annae Lynge, Ark. Bot. 13, No. 13:88, pl. 2, fig. 6. 1914. 


Lectotype: Santa Anna da Chapada, Mato Grosso, Brazil, Malme 
2368B (S). 

Additional specimen examined: Malme, June 27, 1894. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC+ red, P—, 
atranorine and two colorless unknowns, one forming tetragonal 
lamellae, the other needles in the reagent G.A.W. 

The two specimens of P. annae and P. ceracea Lynge (see p. 9) 
agree chemically and morphologically with P. recipienda Nyl. (type: 
Brazil, Nylander Herbarium No. 35212, H), and both species are 
synonyms of P. recipienda. The medulla of P. annae was reported 
by Lynge to be K+ red. Evidently he obtained such a reaction 
from a single unpublished collection (Colonia Risso, Paraguay, 
Malme 1959, 8), labeled P. annae but actually belonging to P. sub- 
caperata Kremplh., which contains salacinic acid (K+ red). 


HALE—REVISION OF PARMELIA SPECIES 7 


7. Parmelia bahiana Ny]. Flora 68:612. 1885.—Lynge 1914, p. 134; 1925, p. 84. 


Lectotype: Rio de Janeiro, Glaziou 1999 (H). 

Range: Brazil. 

Additional specimen examined: Malme 2545**. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC+ red, P—, 
atranorine and alectoronic acid (in Malme 2545**) or an unidentified 
substance (in the lectotype), other specimens not tested. 

The only diagnostic character of P. bahiana is the KC + medullary 
reaction attributable to two different substances, as far as I can 
determine. Since the chemically different specimens are so close 
morphologically, I dare not describe Lynge’s material as a new species, 
for it belongs to the particularly difficult nonisidiate, esorediate group 
of the section Hypotrachyna. The specimens collected by Dusén 
and Warming (UPS) were too fragmentary for certain identification. 


8. Parmelia balansae Mill. Arg. Rev. Mycol. 10:1. 1888,—Lynge, 1914, p. 51. 


Lectotype: Asuncidén, Paraguay, Balansa 8 (G). 

Lynge determined Malme 201B as this species; the specimen has 
not been made available for study, but there is little doubt but that 
Lynge’s determination is correct. Parmelia balansae is closely re- 
lated to P. consors (see p. 11). 


9. Parmelia balansae var. sorediata Miill. Arg. Rev. Mycol. 10:2. 1888.— 
Lynge, 1914, p. 53, pl. 1, fig. 6; 1925, p. 84. 


Parmelia subbalansae Gyel. Repert. Sp. Nov. 33:288. 1931. Based on P- 
balansae var. sorediata Miill. Arg. 

Type: Montevideo, Uruguay, Arechavaleta in 1887 (G). 

Range: Argentina, Brazil, Uruguay. 

Additional specimen examined: Malme, September 7-9, 1894. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC—, P-, 
atranorine present. 

Lynge’s specimen agrees perfectly with the holotype of the variety, 
which, following Gyelnik, should be recognized as a distinct species, 
P. subbalansae Gyel., which differs from P. balansae var. balansae in 
possessing laminal soredia. Parmelia subbalansae appears to be much 
more common than P. balansae. 

10. Parmelia borreri (J. E. Smith) Turn. Trans. Linn. Soc. 9:148. 1808.— 
Lynge, 1925, p. 84. 
Lichen borrert J. E. Smith, Eng. Bot. 25:1780. 1807. 

The material of this species determined by Lynge was destroyed 
at Berlin. The citation as given above is the correct name for the 
currently accepted P. dubia (Wulf. in Jacq.) Schaer. non (Ach.) 
Floerke. 


8 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


11. Parmelia brasiliana Nyl. Flora 68:611. 1885,—Lynge, 1914, p. 113. 


Type: Organ Mountains, Rio de Janeiro, Brazil, Weddell (not 
seen). 

Specimens examined: Malme 309, 312B. Reactions: Thallus K—, 
medulla K—, C—, KC—, P—, lichexanthone present. 

Although the type has not been checked, the identity of this 
distinctive species has not been questioned by previous workers. 
12. Parmelia brasiliana var. glaziovii (Mill. Arg.) Lynge, Nyt Mag. Naturv. 


62:85. 1925. 
Parmelia glaziovid Mill. Arg. Nuov. Giorn. Bot. Ital. 21:353. 1889. 


The Henschen specimen cited by Lynge and preserved at Uppsala 
has not been available for study. 

13. Parmelia brasiliana var. novella (Vain.) Lynge, Ark. Bot. 13, No. 13:115. 
1914.—Lynge, 1925, p. 85. 
Parmelia novella Vain. Acta Soc. Faun. & Fl. Fenn. 7:56. 1890. 

Type: Minas Gerais, Brazil, Vainio 1028 (FH, isotype). 

Range: Brazil. 

Additional specimens examined: Malme 2246B; Hemmensdorf 
in 1898 (UPS). 

Reactions: Thallus K—, medulla K+ dull reddish, C—, KC+ 
reddish, P+ faint orange-red, lichexanthone and unknown sub- 
stances present. 

The status of this entity is not clear at this time. It probably 
falls nearer P. silvatica Lynge (see p. 34) than P. brasiliana because 
of the small, adnate thallus. 

14. Parmelia canaliculata Lynge, Ark. Bot. 13, No. 13:28, pl. 1, fig. 1. 1914.— 
Lynge, 1925, p. 85. 

Holotype: Cachoeira, Rio Grande do Sul, Brazil, Malme 1055, 
February 17, 1893 (S). 

Range: Brazil, Argentina, Uruguay. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC—, P-, 
atranorine and protolichesteric acid present. 

This unique species appears superficially to be in the P. melanothriz 
group, but on closer inspection it is found to have pseudocyphellae 
and lack cilia. Most of the specimens determined by Lynge in 1925 
were destroyed at Berlin. 


15. Parmelia capitata Lynge, Ark. Bot. 13, No. 13:59, pl. 1, figs. 4, §. 1914. 

Holotype: Rio Vermelho, near Bahia, Bahia, Brazil, Malme, 
October 1894 (S). 

Range: Brazil, West Indies. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC—, P-, 
atranorine and an unidentified fatty substance near caperatic acid. 


HALE—REVISION OF PARMELIA SPECIES 9 


The name P. capitata has appeared in the literature several times, 
but such reports are usually based on misidentifications. Parmelia 
capitata is a small plant—so small that one would question its in- 
clusion in the section Amphigymnia. Parmelia sanctae-crucis Vain. 
(cf. Hale, 1959a, p. 22), a common tropical species with similar chem- 
istry, 1s twice the size of P. capitata. One specimen from the West 
Indies (Roseau, Dominica, Evans, US, YU) can probably be referred 
here. 


16. Parmelia ceracea Lynge, Ark. Bot. 13, No. 13:97. 1914,—Lynge, 1925, p. 85. 


Lectotype: Pilcomayo, Gran Chaco, Paraguay, Malme, September 
7, 1893 (S). 

Additional specimens examined: Caldas, Minas Gerais, Brazil, 
Henschen in 1868 (UPS). 

Reactions: As in P. annae Lynge. 

The two specimens agree chemically and morphologically with 
P. recipienda Nyl. (see under P. annae Lynge, p. 6), a rare but 
widespread species in Brazil, Argentina, and Paraguay. 

17. Parmelia cetrata Ach. Syn. Lich. 198. 1814.—Lynge, 1914, p. 90; 1925, 
p. 86. 


Type: North America, Muhlenburg (PH, isotype). 

Range: Cosmopolitan in temperate and tropical regions. 

Additional specimens examined: Malme 463, June 25, 1894, 
October 14-16, 1902. 

Reactions: Thallus K+ yellow, medulla K+ red, C—, KC—, P+ 
orange-red, atranorine and salacinic acid present. 

Du Rietz (1924, p. 330) correctly circumscribed P. cetrata as an 
esorediate species with a reticulately rimose cortex and a uniformly 
rhizinate underside. Although specimens determined by Lynge 
which I have been able to verify all lack soredia, his concept of the 
species included sorediate plants also, which should be referred to 
P. reticulata Tayl. 

18. Parmelia cetrata f. ciliosa Viaud-Grand-Marais, Bull. Soc. Ouest France 
2:156. 1892.—Lynge, 1925, p. 86. 

Material of this form which Lynge examined was not available for 

study. 
19. Parmelia cetrata var. corniculata (Kremplh.) Miill. Arg. Hedwigia 32:228. 
1891.—Lynge, 1914, p. 93; 1925, p. 86. 
Parmelia perforata var. corniculata Kremplh. Naturhist. For. Kjébenhavn 
Vid. Medd. 1873:11. 


Type: Rio de Janeiro, Brazil, Warming (not seen). 
Range: North and South America. 


10 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Specimens examined: Malme 313, September 3, 1893, and January 
25, 1893. Reactions as in P. cetrata var. cetrata. 
The correctness of this identification is undoubted. 


20. Parmelia cetrata subsp. radiata Lynge, Ark. Bot. 13, No. 13:94. 1914. 


Lectotype: Colonia Risso, near Rfo Apa, Paraguay, Malme 1834, 
September 23, 1893 (S). 

Range: Brazil and Paraguay. 

Additional specimen examined: Malme, June 15, 1894. 

Reactions: As in P. cetrata var. cetrata. 

Lynge based this subspecies on the more or less radiating arrange- 
ment of the lobes and noted in addition that the upper cortex was not 
reticulately rimose to the margins, but merely white-maculate. It is 
not uncommon, however, to find a similar condition in specimens of 
both P. cetrata and P. reticulata Tayl. 


21. Parmelia chapadensis Lynge, Ark. Bot. 13, No. 13:153. 1914. 


Holotype: Near Bocca da Serra, Serra da Chapada, Mato Grosso, 
Brazil, Malme 2297B, January 21, 1894 (S). 

Range: Known only from the type locality. 

Reactions: Thallus K+ yellowish, KC+ strong yellow, medulla 
K+ yellow, C—, KC+ orange-yellow, P+ orange-yellow, usnic acid, 
an unidentified pale yellow pigment (in the medulla), and a P+ sub- 
stance, probably near protocetraric acid. 

This saxicolous species at first seems like an oversized specimen of 
the crustose Rinodina oreina (Ach.) Mass. rather than a Parmelia. 
It is a unique species with no near relatives in the genus. 


22. Parmelia cinerascens Lynge, Ark. Bot. 13, No. 13:104. 1914. 


Holotype: Paraguari, Paraguay, Malme 1498, August 2, 1893 (S). 

Range: Known only from the type locality. 

Reactions: Thallus K+ yellow, medulla K+ red, C—, KC—, P+ 
orange-red, atranorine and salacinic acid present. 

In gross appearance this species is near P. minarum Vain. but pro- 
duces salacinie acid. 
23. Parmelia coccinea Lynge, Ark. Bot. 15, No. 1:3. 1915. 

Parmelia ochrococcinea Zahlbr. Cat. Lich. Univ. 8:562. 1932. A new name 
for P. coccinea Lynge non Clem. 

Lynge (1925, p. 97) had already reduced this species to synonymy 
under Pyzxine coccifera (Fée) Nyl., when Zahlbruckner made a new 
name. 


24, Parmelia congruens Ach. Lich. Univ. 491. 1810-—Lynge, 1925, p. 87. 


Type: North America, Swartz. The identity of this species bas 
long been a source of confusion to lichenologists. There is no speci- 
men in the Acharian herbarium according to Mr. Teuvo Ahti (in litt.). 


HALE—REVISION OF PARMELIA SPECIES ll 


A presumed isotype at Uppsala is a pale yellow, nonisidiate plant of 
the section Xanthoparmelia, even though Acharius described it as 
“‘albo-pallescens . . . sordide albo.”’ The chemistry of this fragmen- 
tary specimen is not clear. 

Two specimens identified by Lynge have not been seen. A dupli- 
cate of one of them (Argentina, Lorentz & Hieronymus, M) is the type 
of P. taractica Kremplh., a recognized species in the section Xantho- 
parmelia not to be compared with P. congruens. 

25. Parmelia consimilis Vain. Acta Soc. Faun. & Fl. Fenn. 7:58. 1890.—Lynge, 
1925, p. 87. 

Lectotype: Sitio, Minas Gerais, Brazil, Vainio 1133 (FH). 

Range: Brazil, West Indies. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC+ rose, P+ 
orange-red, atranorine and protocetraric acid present. 

The specimen of Henschen (UPS) examined by Lynge consists of 
an Anaptychia species and a fragment of a sorediate Parmelia, not 
P. consimilis, which is isidiate. 

26. Parmelia consors Nyl. Flora 68:613. 1885.—Lynge, 1914, p. 95, pl. 3, 
fig. 1; 1925, p. 87. 

Lectotype: Brazil, Weddell (H, Nylander Herbarium No. 35276). 

Range: Brazil. 

Additional specimen examined: Malme 1282. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC—, P-, 
atranorine present. 

Malme’s specimen is well developed and typical. This species 
must be very near P. balansae Miill. Arg. Both have a rigid, finely 
white-maculate thallus and produce only atranorine. Parmelia con- 
sors has a dark underside, densely rhizinate to the margins, whereas 
P. balansae becomes lighter brown at the margins with a narrow 
papillate or almost bare zone below. 

27. Parmelia conspersa (Ach.) Ach. Meth. Lich. 205. 1803.—Lynge, 1914, p. 142; 
1925, p. 87. 
Lichen conspersus Ach. Lich. Suec. Prod. 118. 1798. 

Type: Europe (not seen). According to Gyelnik (1936, p. 120), 
the Acharian type is a mixture of nonisidiate P. conspersa and isidiate 
P. isidiata (Anzi) Gyel. I have advocated the acceptance of both 
species (Hale, 1955). 

Range: Cosmopolitan. 

Specimens examined: Malme 563, 1346. Reactions: Thallus K—, 
medulla K+ yellow, C—, KC—, P+ pale orange, stictic and usnic 
acids present. 

The two Malme specimens are isidiate and should be classified . 
as P. isidiata, an entity which Lynge recognized as a variety. A 


12 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


third specimen (Malme 248) lacking isidia is P. flavida Zahlbr. (see 
under P. flava Kremplh., p. 15). 


28. Parmelia conspersa var, hypoclysta Ny]. Syn. Meth. Lich. 1:391. 1860.— 
Lynge, 1925, p. 87. 


The specimen seen by Lynge was destroyed at Berlin. 


29. Parmelia continentalis Lynge, Ark. Bot. 13, No. 13:111. 1914.—Lynge, 
1925, p. 87. 

Holotype: Corumba, Mato Grosso, Brazil, Malme 48, August 1, 
1894 (S). 

Range: Brazil, Paraguay. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC—, P-, 
atranorine present. 

The only distinguishing feature of this Hypotrachyna species is the 
exceptionally rigid thallus. 


30. Parmelia continua Lynge, Ark. Bot. 13, No. 13:109. 1914. 


Holotype: Buriti, Serra da Chapada, Mato Grosso, Brazil, Malme 
June 19, 1894 (S). 

Range: Known only from the type locality. 

Reactions: Thallus K+ yellow, medulla K+ red, C—, KC— 
P+ orange-red, atranorine and salacinic acid present. 

This unusual member of the section Hypotrachyna lacks soredia 
and isidia and produces salacinic acid. It may be a nonisidiate 
variant of P. cinerascens Lynge. 


31. Parmelia cornuta Lynge, Ark. Bot. 13, No. 13:76, pl. 2, fig. 6. 1914. 


d 


Holotype: Santa Anna da Chapada, Mato Grosso, Brazil, Malme 
2477, March 2, 1894 (S). 

Range: Known only from the type locality. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC—, P-, 
atranorine and vulpinic acid present. 

Parmelia cornuta has the same chemical composition as the better 
known P. sulphurata Nees & Flot. but lacks isidia and has longer 
cilia and a more rigid thallus. 

32. Parmelia cornuta var, crocea Lynge, Ark. Bot. 13, No. 13:78. 1914. 
Parmelia crocea (Lynge) Gyel. Repert. Sp. Nov. 29:287. 1931. This is a 
later homonym of Parmelia crocea (Ach.) Sprengl. 

Holotype: Santa Anna da Chapada, Mato Grosso, Brazil, Malme 
2477 bis, March 2, 1894 (S). 

Range: Known only from the type locality. 

Reactions: Thallus K+ yellow, medulla flava K-+- yellowish, C—, 
KC-+ yellowish, P—, medulla crocea K+ purple, atranorine, a pale 
yellow-orange pigment (also known in P. lindmanii Lynge and 


HALE—REVISION OF PARMELIA SPECIES 13 


P. merrill Lynge (= P. lyngeana Zahlbr.)), and an unidentified 
anthraquinone. 

Although Lynge relied on the orange-red lower medulla to differen- 
tiate this variety, it actually contains a yellow pigment entirely 
different from that in var. cornuta (vulpinic acid). On the basis 
of both spore size and chemistry, var. crocea is closer to P. merrillit 
Lynge (see p. 24) than to P. cornuta. We should continue to recognize 
it as a variety rather than coin another new species name, at least 
until additional material throws more light on the range of variation 
in this group. 

33. Parmelia coronata Fée, Essai Crypt. Ecorces, 123, pl. 31, fig. 2. 1824.— 
Lynge 1914, p. 121; 1925, p. 88. 

Type: American tropics (not seen); Fée’s color plate permits 
almost certain identification of the species. 

Range: Tropical America. 

Specimens examined: Malme 2392, 2511Ba, 2522G, 2525a. Re- 
actions: Thallus K+ yellow, medulla K—, C+ rose, KC+ red, 
P—, atranorine and gyrophoric acid present. 

The Malme specimens are well developed and typical. 

34. Parmelia cristifera Tayl. London Journ. Bot. 6:165. 1847.—Lynge, 1914, 
p. 46. 

Lectotype: Calcutta, India, Wallich (FH). 

Range: Tropical regions, especially in the Pacific area. 

Reactions: Thallus K-+ yellow, medulla K+ red, C—, KC—, P+ 
orange-red, atranorine and salacinic acid present. 

The single specimen cited by Lynge (Malme, June 16, 1894) con- 
tains atranorine, protocetraric acid, and usnic acid, and has the 
upper cortex more or less reticulately cracked. It is certainly not 
referable to P. cristifera but may be allied to P. dominicana Vain. 
The material, however, is too poor for satisfactory determination. 


35. Parmelia crustacea Lynge, Ark. Bot. 13, No. 13:108, pl. 3, fig. 4. 1914. 


Holotype: Santa Anna da Chapada, Mato Grosso, Brazil, Malme, 
February 21, 1894 (S). 

Range: Known only from the type locality. 

Reactions: Thallus K—, medulla alba K—, C—, KC-+ faint rose, 
P+ orange-red, medulla crocea K+ purple, lichexanthone, pro- 
tocetraric acid, and an unidentified anthraquinone present. 

This species is very closely related to P. stlvatica Lynge (see p. 34), 
from which it is distinguished by a more adnate growth habit. Lynge’s 
separation of the two species in his key (1914, pp. 20-21) into different 
groups (Cyclocheila and Sublineares) is entirely unsatisfactory. 


14 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


36. Parmelia crystallorum Lynge, Ark. Bot. 18, No. 13:128. 1914. 


Holotype: Corcovado, Rio de Janeiro, Brazil, Malme 59*, August 
14, 1892 (S). 

Range: Known only from the type locality. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC—, P-, 
atranorine present. 

Lynge characterized the species by the large colorless crystals which 
precipitated from the medulla in KOH; the identity of these crystals 
is unknown, and the type specimen is too small and fragile for ade- 
quate chemical analyses. The species resembles P. zahlbruckneri 
Lynge (see under P. gracilis, p. 17) in general appearance, especially 
with regard to the lobation and the ivory or ashy colored shiny 
surface. 

37. Parmelia cyliphora (Ach.) Vain. Acta Soc. Faun. & Fl. Fenn. 13:7. 1896.— 
Lynge, 1914, p. 60. 
Parmelia caperata (L.) Ach. var. cyliphora Ach. Syn. Lich. 196. 1814. 
P. caperata auct. 

Type: North America (UPS, isotype). The Acharian isotype is 
somewhat smaller than but otherwise identical with P. caperata var. 
caperata. The identity of Lichen caperatus L. was first questioned by 
Vainio (1886), who discovered a specimen of Cetraria pinastri (Scop.) 
S. F. Gray in the packet labeled Lichen caperatus in the Linnaean 
Herbarium. He assumed that this was the type specimen and there- 
fore used the epithet caperatus for the Cetraria species, reduced 
pinastri to synonymy, and resurrected the Acharian var. cyliphora as 
the valid name for P. caperata auct. A study of Linnaeus’ original 
diagnosis (Sp. Pl. 1147, 1753), however, indicates that the type of 
Lichen caperatus is probably not to be found in the Linnaean Her- 
barium. Linnaeus based his species on old phrase-names by Royen, 
Guettard, Dillenius, and Morison. The type of one of these pre- 
Linnaean species must be selected as the lectotype of Lichen caperatus, 
an action which has never been taken. Both Dillenius (Hist. Musc. 
pl. 25, fig. 97.A, B, 1741) and Morison (Hist. ox. 633, pl. 7, fig. 1, 1699) 
illustrate a broad lobed Parmelia apparently identical with P. caperata 
auct., not a narrow lobes species such as Cetraria pinastro. 

Specimen examined: Malme 2156. 

. Reactions: Thallus K—, medulla K—, C—, KC+ rose, P+ orange- 
red, usnic and protocetraric acids present. 

The specimen appears to be typical P. caperata except for the lack 
of soredia, a condition not unknown in North American plants. 

38. Parmelia digitata Lynge, Ark. Bot. 13, No. 13:98, pl. 3, fig. 4. 1914. 


Holotype: Santa Anna da Chapada, Mato Grosso, Brazil, Malme 
2545, March 12, 1894 (8). 
Range: Brazil, West Indies. 


HALE—REVISION OF PARMELIA SPECIES 15 


Reactions: Thallus K+ yellow, medulla K—, C+ light orange, 
KC-+ deep orange, P—, atranorine and barbatic acid present. 
Parmelia digitata belongs to the difficult and highly variable P. 
laevigata group. It has distinct marginal laciniae and lobules much 
as in P. lobulifera Degel. from North Carolina. A specimen from 
Jamaica (Imshaug 14910, MSC) is identical with Lynge’s type. 
39, Parmelia eciliata (Nyl.) Nyl. in Fournier, Mex. Pl.I:3. 1872.—Lynge, 1914, 
p. 72. PLATE 1 
Parmelia crinita var. eciliata Nyl. Flora 52:291. 1869. 
Type: Orizaba, Mexico, Bourgeau in 1865 (H). 
Range: Argentina, Brazil, West Indies, Mexico. 
Additional specimen examined: Malme 627. 
Reactions: Thallus K-+ yellow, medulla K+ yellow, C—, KC—, 
P+ pale orange, atranorine and stictic acid present. 
The Malme collection compares well with Nylander’s holotype in 
spore size and lobation, but has somewhat shorter cilia. The species 
is discussed more fully under P. urceolata Eschw. (see p. 37). 


40. Parmelia fatiscens Lynge, Ark. Bot. 15, No. 1:1. 1917. PLATE 2 


Holotype: Santa Anna da Chapada, Mato Grosso, Brazil, Malme, 
October 16, 1902 (S). 

Range: Known only from the type locality. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC-+ rose, 
P+ orange-red, atranorine, protocetraric acid, and traces of usnic 
acid in the cortex. 

This species is characterized by large coralloid outgrowths which 
cannot properly be classified as isidia or soredia, although they 
sometimes become sorediate or pustular-isidiate. These peculiar 
growths also occur in P. fasciculata Vain. from Colombia, which 
contains atranorine and protocetraric acid, and in an undescribed 
species from Liberia, Africa. Certain forms of P. robusta Degel. 
have an atypical formation of tiny sorediate laciniae which coalesce 
into fasciculate coralloid growths resembling those of P. fatiscens. 

41. Parmelia flava Kremplh. var. stellata Lynge, Ark. Bot. 13, No. 13:150, 
pl. 5, figs. 6, 6. 1914. 

Parmelia flavida Zahibr. var. stellata (Lynge) Zahlbr. Cat. Lich. Univ. 
6:137. 1929. Parmelia flavida is a new name for P. flava Kremplh. non 
Rebent. 

Type: Sao Joao d’el Rey, Minas Gerais, Brazil, Malme 311, Sep- 
tember 1, 1894 (S). 

Range: Brazil. 

Reactions: Thallus K—, medulla K—, C—, KC+ rose, P+ 
orange-red, usnic and protocetraric acids present. 


532390—60——2 


16 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


The holotype of P. flava Kremplh. (Serra da Piedade, Minas Gerais, 
Brazil, Warming 294, M), which Lynge did not examine, has much 
broader, obtuse lobes (1.5-2.0 mm. wide) than the new variety (0.8-1.0 
mm. wide), but is otherwise similar in chemistry and gross appearance. 
42. Parmelia flava var. subdichotoma Lynge, Ark. Bot. 13, No. 13:149, pl. 6, 

fig. 7. 1914—Lynge, 1925, p. 88. 


Parmelia flavida Zahlbr. var. subdichotoma (Lynge) Zahlbr. Cat. Lich. 
Univ. 6:137. 1929. 


Type: Sio Jodo d’el Rey, Minas Gerais, Brazil, Malme 310, Sep- 
tember 1, 1894 (S). 

Range: Brazil. 

Additional specimen examined: Caldas, Minas Gerais, Brazil, 
Henschen in 1868 (UPS). 

Reactions: As in P. flava var. stellata Lynge. 

This variety can hardly be distinguished from var. stellata. We 
have no choice but to maintain both varieties, which seem to be 
distinct from var. flavida, until the range of variation of the species 
is more completely known. 


43. Parmelia fragilis Lynge, Ark. Bot. 13, No. 13:123. 1914. 


Holotype: Santa Anna da Chapada, Mato Grosso, Brazil, Malme 
2365*, February 19, 1894 (S). 

Range: Brazil. 

Reactions: Thallus K+ yellow, medulla K—, C+ orange-red, 
KC-+ deep orange-red, P—, atranorine and olivetoric acid present. 

This plant is identical in every respect with P. intercalanda Vain. 
(syntype: Lich. Bras. Hrs. 899, FH). This species is apparently the 
first of the section Hypotrachyna in which olivetoric acid has been 
demonstrated. Lynge suggests a similarity with P. microblasta Vain., 
which does have the same gross appearance but which differs quite 
significantly in having isidia and norstictic acid (K+ red). 
44. Parmelia fungicola Lynge, Ark. Bot. 13, No. 13:129. 1914. 


Holotype: Santa Anna da Chapada, Mato Grosso, Brazil, Malme 
2438B, February 27,-1894 (S). 

Range: Known only from the type locality. 

Reactions: Thallus K+ yellow, medulla K—, C+ rose, KC-+ red, 
P—, atranorine present, gyrophoric acid suspected but not proved 
microchemically because of the scarcity of material. 

At first glance P. fungicola seems no more than a small isidiate form 
of P. coronata Fée. The contaminating fungus on the underside is a 
conspicuous feature. Lynge erroneously gave the medullary reaction 
as K+ yellow and C—, an understandable mistake in view of the 
small size of the thallus. 


HALE—REVISION OF PARMELIA SPECIES 17 


45. Parmelia gracilis (Miill. Arg.) Vain. Ann. Acad. Soc. Faun. & FI. Fenn. 
7:55. 1890.—Lynge, 1914, p. 124. 
Parmelia laevigata var. gracilis Mill. Arg. Rev. Mycol. 1:169. 1879. 
P. confusula Zahlbr. Cat. Lich. Univ. 6:162. 1929. A new name for P. 
gracilis (Mill. Arg.) Vain., non Sprengel nec Mill. Arg. 

Type: Boqueron de Bogoté, Colombia, André 923 bis, December 21, 
1875 (G, holotype; US, isotype). 

Range: Colombia, Brazil. 

Additional specimens examined: Malme 2241, 2749. 

Reactions: Thallus K+ yellow, medulla K+ reddish, C—, KC+ 
red, P+ pale orange, atranorine and unidentified substances present; 
the acetone extract is a thick pale orange crust. 

Parmelia confusula Zahlbr. is antedated by P. zahlbrucknert Lynge, 
which is identical with the type of P. gracilis (Mill. Arg.) Vain. (see 
p. 38). A noteworthy feature of this saxicolous lichen, other than the 
peculiar chemical reactions, is the shiny, whitish gray upper cortex, 
often transversely cracked and infested by a small black fungus. 


46. Parmelia hieronymi Lynge, Nyt Mag. Naturv. 62:88. 1925. 


The specimen on which this species was based was destroyed at 
Berlin. I have seen no herbarium specimens agreeing with the 
original description. 

47, Parmelia isidiophora Zahlbr. Sitzungsb. Akad. Wiss. Math. Naturw. (Wien) 
111:420, pl. 1. 1902.—Lynge, 1925, p. 89. 

Type: Botanical Garden, Rio de Janeiro, Brazil, Héhnel 169 (not 
seen, but appears to be typical P. caroliniana Nyl. from Zahlbruck- 
ner’s photograph). 

Specimen examined: Caldas, Minas Gerais, Brazil, Henschen in 
1868 (UPS). 

The single Henschen collection, verified by Zahlbruckner, is typical 
P. caroliniana Nyl., a widespread corticolous species in North and 
South America (cf. Hale, 1959a, p. 17). The specimen contains 
atranorine and perlatolic acid. 

48. Parmelia kamtschadalis (Ach.) Eschw. var. americana (Mey. & Flot.) Nyl. 
Ann. Sci. Nat. Bot., ser. 4. 11:215. 1859.—Lynge, 1925, p. 89. 
Evernia americana Mey. & Flot. Verh. Kaiser Leopold Carol. Akad. Natur- 
forsch. 19, suppl. 1:211. 1843. 

Type: Tropical America (destroyed at Berlin). 

Specimen examined: Serra da Itatiaia, Brazil, Dusén, May 1902 
(UPS). 

Reactions: Thallus K+ yellow, medulla K+ red, C—, KC—, P+ 
orange-red, atranorine and salacinic acid present. 

The material conforms with the accepted concept of P. amerieana 
(Mey. & Flot.) Mont., which differs from P. cirrhata Fr. in being isi- 
diate. Both species are widespread in mountains of tropical regions. 


18 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Parmelia kamtschadalis (Ach.) Eschw. is a completely ‘unrelated 

species in the section Xanthoparmelia. 

49, Parmelia laceratula Nyl. Mem. Soc. Imp. Sci. Nat. Cherbourg 5:105. 
1857.—Lynge, 1914, p. 100, pl 3, fig. 3. 

Holotype: Burnet and Brisbane River, Australia, Miller (H). 

Range: Australia. 

Reactions: Thallus K+ yellow, medulla K—, C+ red, KC-+ red, 
P—, atranorine and lecanoric acid present. 

Parmelia laceratula is a synonym of P. subflava Tayl. (type: Van 
Diemen’s Land, FH). The types of both species have peculiar coarse 
marginal isidia and no pseudocyphellae except on the receptacle of 
the apothecia. The Malme specimens (533 and 828) seen by Lynge 
have the same chemistry but are densely pseudocyphellate on the 
upper surface. They are closest to P. riograndensis Lynge (=P. cf. 
bolliana Mill. Arg. see p. 31), but the thallus is very fragile. Lynge 
gave an incorrect chemical test (C—) and failed to include the species 
with the other pseudocyphellate Amphigymnias. 


50. Parmelia laongii Lynge, Ark. Bot. 13, No. 13:68, pl. 1, fig. 3. 1914. 


Holotype: Santa Anna da Chapada, Mato Grosso, Brazil, Malme 
2392*, February 21, 1894 (S). 

Range: Known only from the type locality. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC+ red, P—, 
atranorine and alectoronic acid present; cilia K+ violet. 

Parmelia laongit is a member of the P. melanothriz group (see p. —), 
closest to P. argentina Kremplh. but with smaller spores (8-10 X 16-18y 
vs. 10-13 19-23) and cilia K+ violet. 

51. Parmelia latissima Fée, Ess. Crypt. Suppl. 119, pl. 38, fig. 4. 1837.— 
Lynge, 1914, p. 41. 

Type: American tropics and Jamaica (not seen but well illustrated 
in color by Fée). 

Range: Tropical regions. 

Specimen examined: Malme 2742B. Reactions: Thallus K+ yel- 
low, medulla K+ red, C—~, KC—, P+ orange-red, atranorine and 
salacinic acid present. 

Only this one Malme specimen seems to be typical P. latissima. 
Four other specimens examined by Lynge (Malme 2364, February 21, 
1894, June 15 and June 25, 1894) contained atranorine and pro- 
tocetraric acid and should be identified as P. zollingeri Hepp, a wide- 
spread tropical and subtropical species, apparently much more common 
than P. latissima. Aside from chemical differences, P. zollingert seems 
to have slightly smaller spores than P. latissima (18-24 vs. 28-32u, 
both with the episporium 3-4 wide). 


HALE—REVISION OF PARMELIA SPECIES 19 


52. Parmelia latissima var. corniculata Kremplh. Flora 61:463. 1878.—Lynge, 
1914, p. 44. 

Holotype: Argentina, Lorentz & Hieronymus (M). 

Range: Argentina, Brazil. 

Additional specimens examined: Malme 2738, 2743B, 2512Ba. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC+ rose, 
P+ orange-red, atranorine and protocetraric acid present. 

The specimens seen by Lynge correspond perfectly to Krempel- 
huber’s original type. The K— reaction was overlooked by Lynge. 
If we should follow a strict chemical criterion, var. corniculata should 
be transferred to P. zollingeri, which also contains protocetraric acid. 
However, a formal recombination of the varietal name would be 
premature at this time, when the differences between P. latissima 
and P. zollingert are so poorly understood. 


53. Parmelia latissima f. microspora Lynge, Ark. Bot. 13, No. 13:45, 1914. 


Lectotype: Near Bocca da Serra, Serra da Chapada, Mato Grosso, 
Brazil, AMfalme 2244*, January 19, 1894 (S). 

Additional specimen examined: Malme 2244. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC+ red, P—, 
atranorine and alectoronic acid present. 

Both of the specimens identified by Lynge are the same as P. 
wainit A. L. Smith (see p. 29, under P. proboscidea). He described 
the form in part on the basis of a faulty color test, the medulla 
supposedly K+ yellow changing to red but actually K—. 


54. Parmelia latissima var. minima Lynge, Ark. Bot. 13, No. 13:45. 1914. 


Holotype: Buriti, Serra da Chapada, Mato Grosso, Brazil, Malme 
2243C**, January 20, 1894 (S). 

Range: Known only from the type locality. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC+ rose, P+ 
orange-red, atranorine and protocetraric acid present. 

Lynge at first described this plantas a newspecies, ‘‘P.sublatissima,”’ 
in the herbarium. It is a much smaller plant than P, latissima, 
although otherwise comparable in morphology, and contains protoce- 
traric acid, more characteristic of P. zollingeri Hepp. We cannot 
validly judge this new variety on the basis of a single small specimen. 
59. Parmelia leucoxantha Miill. Arg. Flora 64:85. 1881.—Lynge, 1914, p. 82. 

Type: Apiahy, Brazil, Puiggari 1050, March 1880 (G). 

Range: Brazil. 

Additional specimen examined: Malme, April 25, 1894. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC+ rose, 
P+ orange-red, atranorine, protocetraric acid, and usnic acid present. 

Hillmann (1939) considered Lynge’s determination to be incorrect. 


20 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Miiller’s type consists of four or five pieces of lichen pasted on a 
single card. The bulk of the material is identical with Malme’s 
plant in chemistry and morphology, but the largest piece is a mixture 
of P. leucoxantha and another very similar species, apparently P. 
nylandert Lynge (see p. 25), which has cilia and reacts K+ red 
(salacinic acid). It is probably this aberrant thallus on which Vainio 
(1900) based his K-++ red reaction and report of sparse cilia. The 
K— eciliate material, equivalent to Lynge’s concept, should be 
designated as the lectotype of P. leucorantha. 


56. Parmelia lindmanii Lynge, Ark. Bot. 13, No. 13:74. 1914. PLATE 3 


Holotype: Porto Alegre, Rio Grande do Sul, Brazil, Malme 450, 
September 25, 1892 (5). 

Range: Southern United States, Mexico, West Indies, Central and 
South America, Africa. 

Reactions: Thallus K+ yellow, medulla K+ yellowish,C—, KC+ 
yellowish, P—, atranorine and an unidentified pale orange-yellow 
pigment (in the medulla) present. 

This distinct species has a wide distribution in the tropics, yet it 
has almost always been misidentified in herbaria as P. sulphurata 
Nees & Flot. or P. tinctorum Ny]. (see Hale, 1959, p. 20). The pale 
yellowish pigment, also known in P. merrillit Lynge (p. 24) and P. 
cornuta var. crocea Lynge (p. 12), forms a long streak on chroma- 
tographic paper with butanol as a solvent. In contrast, vulpinic 
acid, which has a deeper lemon-yellow color, rises up the paper 
without any streaking. A chromatographic test is often necessary 
to separate the species from P. sulphurata. 

57. Parmelia longiconida Lynge, Ark. Bot. 13, No. 13:130, pl. 3, figs. 7,8. 1914.— 
Lynge, 1925, p. 89. 

Holotype: Near Rio Apa, Colonia Risso, Paraguay, Malme 1949, 
October 21, 1893 (S). 

Range: Paraguay, Brazil. 

Reactions: Thallus K+ yellow, medulla K+ yellow, C—, KC—, 
P-+ orange, atranorine and stictic acid present. 

This species is conspecific with P. scrobicularis Kremplh. from Brazil 
(holotype: Lagoa Santa, Warming, M). Two unusual characters 
are the pruinose apothecia, very rare in Parmelia, and the strongly 
reticulately rugose upper cortex. Parmelia sbarbaronis B. de Lesd. 
(Hale, 1959a, p. 23) differs only in having soredia and shorter conidia, 


58. Parmelia magna Lynge, Ark. Bot. 13, No. 13:83, 1914.—Lynge, 1925, p. 89. 
Holotype: Sao Joao d’el Rey, Minas Gerais, Brazil, Malme 269, 
August 31, 1892 (S). 
Range: Brazil. 


CONTR. NAT. HERB. VOL. 36 HALE—PLATE 1 


Pamelia @Ciliata (Nyl.) Nyl. 
(Holotype ) 


Mat. chim: Atranorine, stictic acid. 
Examined by Mason E. Hale, Jr. U. S. National Museum IITI.1953 


Above: Parmelia acanthifolia Pers. (holotype, Gaudichaud 13, P). Below: Parmelia 


eeiltaia (Nyl) Nyl. (holotwpe, Bourgeau, 11). 


CONTR. NAT. HERB. VOL. 36 HALE—PLATE 2 


Parmelia fatiscens Lynee (holotype, Wale, S$). 


CONTR. NAT. HERB. VOL. 36 HALE—PLATE 3 


Above: Parmelia lindmanit Lynge (holotype, MWalme 450, 8). Below: Parmelia proboscidea 
var. ornatula Zahlbr. (holotype, Damazio 1090, G). 


CONTR. NAT. HERB. VOL. 36 HALE—PLATE 4 


Parmelia mesotropa Mull. Are. Gectotys pe, Balansa, G). 


CONTR. NAT. HERB. VOL. 36 HALE--PLATE 5 


ty fre ar _ 
whee AIA tee AS Aw?’ 


Above: Parmelia saneti-angelit Lynge (holotype, Malme, $). Below: Parmelia bicornuta 
\Till. Arg. (holotype, Leyland, G). 


HALE—REVISION OF PARMELIA SPECIES 21 


Reactions: Thallus K+ yellow, medulla K+ red, C-, KC—, P+ 
orange-red, atranorine, salacinic and usnic acids present. 

Parmelia magna is a large lichen, lacking isidia and soredia, with 
wide rotund, monophyllous lobes that easily break away from the 
main thallus. It is distantly related to P. flavescens (Kremplh.) 
Nyl.; its relation to other species with usnic acid may be illustrated 
in the following key: 


1. Thallus isidiate; medulla K+ red or K—. 


2. Medulla K-+ red (salacinic acid) . . . . P. flavescens (Kremplh.) Nyl. 
2. Medulla K—. 
3. Medulla C—, KC+ rose. . . . P. madagascariacea (Hue) des Abb. 


3. Medulla C+ rose (gyrophoric acid) or C—, KC—. 
P. xanthina (Mill. Arg.) Vain. (see p. 38) 
1. Thallus without isidia; medulla K+ red (salacinic acid). 
4, Thallus fragile; lobes breaking apart. . ....... P. magna Lynge 
4, Thallus not fragile; lobes intact. 
5. Lobes elongate, conspicuously ciliate. 
P. radians Lynge (=P. delicatula Vain., see p. 29) 
5. Lobes short, more or less imbricate, cilia very sparse. 
P. microdactyla Hale (see p. 21) 


A second specimen identified by Lynge as P. magna (Rio de Janeiro, 
Widgren, UPS) is a different plant, here proposed as a new species: 


Parmelia microdactyla sp. nov. 


Thallus laxe adnatus, 8-10 cm. latus, dilute flavescens, irregulariter 
lobatus, lobis 8-12 mm. latis, 10-15 mm. longis, nonnihil imbricatus, 
superne planus, laevigatus, nec isidiatus, neque sorediatus, margine 
integro vel plus minusve dactyloideo-lobulato, lobulis discretis, seriatis, 
0.2-0.4 mm. latis, 0.4-0.6 mm. longis, parce ciliato, ciliis usque ad 
0.5 mm. longis, medulla alba, subtus niger et rhizinosus, ambitum 
versus castaneus, glaber. Apothecia ignota; pycnidia numerosa, 
conidiis 6 » longis. Thallus KOH—, medulla KOH+ rubra, C—, 
KC—, P+ aurantiacus, acidum salacinicum et acidum usnicum 
continens. 

Type in the Botanical Museum of the University of Uppsala, col- 
lected by Widgren, Rio de Janeiro, Brazil; isotype in the Riksmuseum, 
Stockholm. 

This new species is closest to the isidiate P. flavescens, but the 
isidia are replaced by peculiar marginal lobules. Zahlbruckner 
unnecessarily complicated the nomenclature of this group when he 
was revising the South American collections of Schiffner and Wettstein. 
Krempelhuber had described P. glaberrima 8 flavescens, which Ny- 
lander raised to species rank in 1885. Although Nylander actually 
saw an isotype of P. glaberrima B flavescens when he made the new 
combination, Zahlbruckner followed Vainio (1896, p. 33) in the belief 


22 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


that “P. flavescens Nyl.” was based on a gray, not a yellow specimen. 
Of course it is immaterial whether Nylander applied the new combi- 
nation to a correctly identified plant or not. The holotype of P. 
glaberrima B flavescens, while not as yellow as typical P. zanthina 
(Mill. Arg.) Vain., does contain usnic acid. Following his own 
peculiar rules of nomenclature, Zahlbruckner wanted to make the 
combination “P. flavescens Zahlbr.,” but since this name was pre- 
occupied by “P. flavescens Nyl.”, he had to propose a new name, 
P. protoflavescens. For the supposedly gray plant seen by Nylander, 
he proposed the name P. pseudoflavescens, which is no more than a 
nomen nudum. The synonymy is summarized below: 

Parmelia flavescens (Kremplh.) Nyl. Flora 68:607. 1885. 

P. glaberrina B flavescens Kremplh. Flora 52:223. 1869. Type: Rio de 

Janeiro, Brazil, Glaziou (M). 


P. protoflavescens Zahlbr. Denkschr. Akad. Wiss. Math. Naturw. Wien 83:176. 
1909. Illegitimate name. 


P. pseudoflavescens Zahlbr. Denkschr. Akad. Wiss. Math. Naturw. Wien 
83:176. 1909. Nomen nudum. 
59. Parmelia malmei Lynge, Ark. Bot. 13, No. 13:116, pl. 2, figs. 3, 4. 1914. 


Holotype: Near Bocca da Serra, Serra da Chapada, Mato Grosso, 
Brazil, Malme 2750, June 5, 1894 (S). 

Range: Brazil. 

Reactions: Thallus K—, medulla alba K—, C—, KC+ rose, P+ 
orange-red, medulla crocea K+ purple, lichexanthone, protocetraric 
acid, and an unidentified anthraquinone present. 

Differing only in the greater development of pustules, this species 
falls within the morphological and chemical range of P. minima Lynge 
(see p. 25) and should be considered synonymous with it. The 
species has some affinity with P. formosana Zahlbr. (Hale, 1958b, p. 
89), a larger plant which also contains lichexanthone (but not pro- 
tocetraric acid) and has similar pustular outgrowths. 


60. Parmelia marginalis Lynge, Ark. Bot. 13, No. 13:112. 1914. 


Holotype: Santa Anna da Chapada, Mato Grosso, Brazil, Malme 
2393 **** February 21, 1894 (S). 

Range: Known only from the type locality. 

Reactions: Thallus K+ yellow, medulla K—, C+ rose, KC+ red, 
P—, atranorine and gyrophoric acid present. 

Parmelia marginalis adds yet another element of variability to the 
already large and difficult group of isidiate species in the section 
Hypotrachyna. It is characterized by an ashy white thallus (as op- 
posed to the buff hue of the P. minarum group). It resembles P. 


HALE—REVISION OF PARMELIA SPECIES 23 


hookert Tayl. (C+ red, lecanoric acid) rather closely, but it should 
not be classified merely as a chemical strain of that species. 


61. Parmelia melanothrix (Mont.) Vain. Ann. Soc. Faun. & Fl. Fenn. 7:30. 
1890.—Lynge, 1914, p. 54; 1925, p. 90. 
Parmelia urceolata var. melanothriz Mont. Ann. Sci. Nat. Ser. II, Bot. 2:372. 
1834, 


Type: Brazil, Gaudichaud 89 bis (not seen); Vainio apparently saw 
the type in the Paris Museum and found it to react KC—. 

Range: Tropical regions. 

Specimens examined: Malme 201, 314, 967 (S); Glaziow 1835; 
Caldas, Minas Gerais, Brazil, Henschen in 1868; Regnell, s.d.; Brazil, 
Guillemin 127; and Rio de Janeiro, Warming (UPS). Reactions: 
Thallus K+ yellow, medulla K—, C—, KC—, P—, atranorine and 
an unknown fatty substance present. 

Parmelia melanothriz belongs to a small, well-circumscribed group 
of primarily tropical species which have a white-maculate cortex, 
marginal cilia, and a pale zone below at the margin contrasting with 
the black rhizinate center. Its relationship to the other species of 
this group in the Western Hemisphere is shown in the following key: 


1. Margins sorediate; apothecia unknown. ... .. .. . P. hypotropa Nyl. 
1. Soredia lacking; apothecia invariably present. 
2. Apothecia perforate. 
3. Salacinic acid present; spores 5-7 X 8-104. P. uruguensis Kremplh. 
3. Norstictic acid present; spores 7-8 X 12-14 up. 
P. perforata (Jacq.) Ach. 
2. Apothecia imperforate. 
4, Medulla KC-; spores 10-12 X 24-26 uy; episporium 3 uz. 
P. melanothrix (Mont.) Vain. 
4, Thallus KC-+ red, alectoronic acid present; episporium less than 2 yu. 
5. Cilia K—. 
6. Thallus membranaceous; spores 6-7 X 17-19 yu. 
P. argentina Kremplh. 
6. Thallus rigid, covered with large apothecia; spores (after Lynge) 
12-138 X 17-24 4H......4.4.6. P. rigida Lynge (see p. 31) 
5. Cilia K+ violet. 
7. Apothecia eciliate or at most sparsely ciliate, pale buff below at the 
margin; spores 6-8 X 11-14 uz. 
P. subproboscidea Lynge (see p. 35) 
7. Apothecia ciliate, tinged below pale reddish-orange; spores 8-10 X 
16-18 u......66 5622s P. laongii Lynge (see p. 18) 


Two additional specimens identified by Lynge as P. melanothriz 
(Malme 2522F* and February 19, 1894) reacting KC+ red (not 
KC— as reported by Lynge) are typical P. argentina Kremplh. 
(holotype: Argentina, Lorentz & Hieronymus, M). 


24 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


62. Parmelia melanothrix f, microspora Lynge, ad int., Ark. Bot. 13, No. 
13:56, 1914. 


Lectotype: Villa Morra, Asuncién, Paraguay, Malme 1585C, 
August 14, 1893 (S). 

Additional specimens examined: Malme, September 3 and September 
7 (two packets), 1893. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC+ red, P—, 
atranorine and alectoronic acid present. 

As a new form ‘‘ad interim,” this entity has no taxonomic status. 
The spores are smaller than those of typical P. melanothriz, and the 
chemical reaction as given by Lynge (KC—) is incorrect. This 
combination of morphological and chemical characters is found in P. 
subproboscidea Lynge (see p. 35), with which the Malme specimens 
are identical. A fourth specimen (Malme, September 3, 1893) is too 
fragmentary for verification. 

63. Parmelia merrillii Lynge, Ark. Bot. 13, No. 13:79. 1914. 


Parmelia lyngeana Zahlbr. Cat. Lich. Univ. 6:243. 1929. A new name for 
P. merrillit Lynge, non Vainio, Phil. Journ. Sci. 4:658. 1909. 


Holotype: Coxipé Igreja, near Cuyabé, Mato Grosso, Brazil, 
Malme 2198B, December 27, 1893 (S). 

Range: Known only from the type locality. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC+ yellowish, 
P—, atranorine and an unidentified pale orange-yellow pigment (in 
the medulla) present. 

The species is very close to P. cornuta var. crocea Lynge (see p. 12) 
in morphology and chemistry, but the type material (the only collec- 
tion of the species) is in poor condition. 

64. Parmelia microsticta Mill. Arg. Flora 62:164. 1879—Lynge, 1914, p. 24; 
1925, p. 90. 
Parmelia borrert var. allophylla Kremplh. Flora 61:438. 1878. Type: 
Argentina, Lorentz & Hieronymus (M). 

Type: Montevideo, Uruguay (not seen). 

Additional specimens examined: Malme 940, January 25, 1893; 
June 1893; September 11 and 13, 1894. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC—, P-, 
atranorine and protolichesteric acid present. 

This pseudocyphellate species is distinguished by a rigid thallus 
and a negative C reaction. It is otherwise similar to P. bolliana 
Mill. Arg. 
64a. Parmelia microsticta var. riograndensis (Lynge) Lynge, Nyt Mag. Naturv. 

62:90. 1925. 

This combination was based on P. riograndensis Lynge, which is 

discussed on p. 31. 


HALE—REVISION OF PARMELIA SPECIES 25 


65. Parmelia minarum Vain. Ann. Soc. Faun. & FI. Fenn. 7:48. 1890.—Lynge, 
1914, p. 106. 

Type: Sitio, Minas Gerais, Brazil, Vainio 1040 (FH, isotype). 

Range: Central and South America, West Indies, Africa. 

Reactions: Thallus K+ yellow, medulla K—, C+ rose, KC+ red, 
P—, atranorine and gyrophoric acid present. 

The specimen identified by Lynge (Malme 2243) cannot be referred 
to P. minarum. It is a large plant with marginal cilia, twice the size 
of typical P. minarum, although both entities contain atranorine 
and gyrophoric acid. This is in all probability a new species, which I 
would hesitate to describe from the single collection. 


66. Parmelia minima Lynge, Ark. Bot. 13, No. 13: 139, pl. 5, figs. 3,4. 1914. 


Lectotype: Near Bocca da Serra, Mato Grosso, Brazil, Malme 
2747 bis, June 15, 1894 (S). 

Range: Brazil. 

Additional specimens examined: Malme 2747, s.n. 

Reactions: Thallus K—, medulla alba K—, C—, KC+ rose, 
P+ orange-red, medulla crocea K+ purple, lichexanthone, pro- 
tocetraric acid, and an unidentified anthraquinone present. 

Lynge’s “originaleksemplar” is an intimate mixture of an ashy 
white plant with sorediate pustules and a yellowish one with isidia. 
His Latin description includes the morphological and chemical 
characters of the ashy plant and the external color of the yellowish 
plant. There is no doubt that the ashy plant should be the lectotype 
of P. minima, of which P. malmei Lynge is probably a synonym (see 
p. 22). The yellowish plant is too fragmentary for identification. 
67. Parmelia nylanderi Lynge, Ark. Bot. 13, No. 13:82. 1914. 

Holotype: Near Sao Jeronymo, Serra da Chapada, Mato Grosso, 
Brazil, Mdalme 2747, June 3, 1894 (S). 

Range: Brazil. 

Reactions: Thallus K+ yellow, medulla K+ red, C—, KC—, P+ 
orange-red, atranorine, salacinic acid, and usnic acid present. 

This saxicolous species may be related to P. leucorantha Miill. 
Arg. (see p. 19), which contains protocetraric acid and lacks cilia. 
68. Parmelia olivaria Hue, Lich. Extra-Europ. 195. 1899.—Lynge, 1925, p. 91. 


The specimen from Argentina so identified by Lynge was destroyed 
at Berlin. 


69. Parmelia osseo-albida Lynge, ad int., Ark. Bot. 13, No. 13: 133. 1914. 


Lectotype: Porto Alegre, Rio Grande do Sul, Brazil, Malme 595, 
October 15, 1892 (S). - 
Additional specimen examined: Malme 1330B. 


26 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Reactions: Thallus K+ yellow, medulla K+ reddish, C—, KC+ 
red, P+ pale orange, atranorine and unidentified substances present; 
the acetone extract is a pale orange crust. 

This species, which must be rejected since it was proposed pro- 
visionally, differs from P. zahlbrucknert Lynge (see under P. gracilis, 
p. 17) in having more irregular lobing and a looser attachment to the 
rock substratum. Considering that the plants share an unusual 
chemistry, I do not believe that the slight morphological variation 
constitutes a valid specific difference. As a matter of fact, in his key 
Lynge separated P. osseo-albida (“medulla KOH non coloratur’’) 
from both P. gracilis and P. zahlbruckneri (“medulla KOH rubescens’’) 
by a simple error: The medullary reaction of P. osseo-albida is given 
in the diagnosis (p. 134) as “‘intus e flavo rubescens,”’ 

70. Parmelia pachyderma Hue, Lich. Extra-Europ. 1387, pl. 4, figs. 1,2. 1899.— 
Lynge 1914, p. 137; 1925, p. 91. 

Type: Montevideo, Uruguay, Dr. Courbon 536 (not seen). 

The specimen identified by Lynge (Malme 1320) is identical with 
Parmelia fistulata Tayl. (syntypes: Argentina, Tweedie, and Monte- 
video, Uruguay, Darwin, FH). Zahlbruckner (1929, p. 60) inexplic- 
ably reduced it to a variety of P. cirrhata Fr. The two syntypes, 
however, differ significantly from P. cirrhata in having protocetraric 
acid and atranorine, rather than salacinic acid, and a beautifully 
effigurate white-maculate cortex. Judging from illustrations of the 
types, Hue’s long-accepted name P. pachyderma and P. felipponer 
Lindau should be placed in synonymy under P. fistulata. 


71. Parmelia palmarum Lynge, ad. int., Ark. Bot. 13, No. 13:136. 1914. 


Holotype: Buriti, Serra da Chapada, Mato Grosso, Brazil, Malme 
2243C*, January 20, 1894 (S). 

Since Lynge proposed this new species ‘‘ad interim” and did not 
definitely accept it, it must be rejected. It is nevertheless a valid 
species, which may be characterized as follows, the description com- 
piled from Lynge’s original diagnosis of P. palmarum: 

Parmelia palmarum Lynge 
Parmelia palmarum Lynge, ad int., Ark. Bot. 13, No. 13:136, cf. pl. 4, 
figs. 1,2. 1914. 

Thallus laxe adpressus, mollis, cinereus vel vix flavescens, laciniis 
non bene radiantibus, 1-2 mm. latis, contiguis vel vulgo imbricatis, 
transversim rugosis, di- vel trichotomiter lobatis et crebre divergenter 
ramosis, lacinulis truncatis vel rotundatis et crenulatis, centrum 
versus lacinulis secundariis numerosis, parvis, digitatim vel coralli- 
formiter ramosis, sorediis, isidiis et ciliis deficientibus, subtus niger, 
ad apicem anguste castaneus, usque ad ambitum rhiziniis nigris, 
ramosis, sat dense instructus. Cortex superior 20-25 u altus, medulla 


HALE—REVISION OF PARMELIA SPECIES 27 


alba, 80-150 yu alta, cortex inferior fuscus vel fusconiger, 13-15 p altus. 
Apothecia numerosa, diam. 5-6 mm., sessilia, non perforata, disco 
flavofuscente, subopaco, non pruinoso; ‘hymenium superne fuscescens, 
ceterum decolor, 50-55 4; sporae anguste limbatae, ovales, 9.5-11 u 
longae, 5.3-6.6 u crassae; conceptacula pycnoconidiorum subglobosa, 
100-160 u alta, 100-110 uw lata, conidiis 4-5 pu longis. Thallus K+ 
flavescens, medulla K—, C—, KC+ roseus, P—, atranorinum et 
materiam jgnotam continens. 

Type in the Naturhistoriska Riksmuseum, Stockholm, collected at 
Buriti, Serra da Chapada, Mato Grosso, Brazil, by G. A. Malme, No. 
2243C*, January 20, 1894. 

This species is a close relative of P. livida Tayl. (type: New Orleans, 
Hook, Herb., FH), a common corticolous lichen in the southeastern 
United States with identical external appearance, although the thallus 
is more ashy white, and a similar KC+ reaction; P. livida differs 
principally i in having smaller spores, 7-8 u long, uniseriately arranged 
in the asci. Parmelia bahiana Nyl. could easily be confused with P. 
palmarum, except that it has a dense mat of rhizines projecting beyond 
the margins of the lobes. Lynge’s “originaleksemplar”’ of P. zahl- 
brukneri (see p. 38) and one collection of P. regis Lynge (see p. 30) 
are both referable to P. palmarum. 


72. Parmelia paraguariensis Lynge, Ark. Bot. 13, No. 13:71, pl. 1, fig. 7. 1914. 


Holotype: Cierro Negro, Paraguari, Paraguay, Malme 1539, 
August 8, 1893 (S). 

Reactions: Thallus K+ yellow, medulla K—, C+ red, KC+ red, 
P—, atranorine and lecanoric acid present. 

This species is conspecific with P. africana Mill. Arg. (lectotype: 
Ghattas, Djur, Africa, Schweinfurth in 1877, G), a rather widespread 
species in Africa. A diagnostic character, aside from the unexpected 
occurrence of lecanoric acid, is the white-maculate upper cortex. 
Parmelia rissoensis Lynge (see p. 31) is merely a poorly developed 
specimen of this same species. 

73. Parmelia perforata (Jacq.) Ach. Meth. Lich. 217. 1803.—Lynge, 1925, p. 91. 
Lichen perforatus Jacq. Coll. Bot. 1:116, pl. JI. 1786. 

Type: Pennsylvania; not seen, but the original specimen is well 
illustrated in the color plate by Jacquin. 

Range: Southeastern United States. 

Lynge determined two packets as P. perforata (Caldas, Minas 
Gerais, Brazil, Henschen in 1868, UPS), but both are misidentified. 
One specimen is P. subcaperata Kremplh., and the other is P. leu- 
cosemotheta Hue, both of which contain atranorine and salacinic acid. 
Parmelia perforata, a species endemic to the United States, contains 
norstictic acid. 

532390—60——_3 


28 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


74. Parmelia persulphurata Nyl. in Cromb. Journ. Linn. Soc, London 16:219. 
1877.—Lynge, 1914, p. 80. 


Type: Not seen; the holotype was collected by Crombie in Bahia, 
Brazil. Nylander (Flora 68:606, 1885) later published the same 
diagnosis that he sent to Crombie in litt. and cited two specimens, 
from Cuba and Louisiana, which are the same as the holotype of P. 
sulphurata Nees & Flot. according to Hillmann (1939). 

The Malme specimen (June 26, 1894) identified by Lynge lacks 
isidia and cannot therefore be P. persulphurata (if we are correct in 
assuming the latter species is synonymous with P. sulphurata). 
Actually it is closer to P. cornuta Lynge (see p. 12), but the material 
is far too poor for sound judgment. 


75. Parmelia pluriformis Ny]. Synops. Lich, 381. 1858-60.—Lynge, 1925, p. 91. 


Type: Minas Gerais, Brazil, Weddell, Nyl. Herb. No. 35585, (H). 

Range: Brazil. 

Additional specimen examined: Sao Paulo, Brazil, Lindberg, April 
1854 (UPS). 

Reactions: Thallus K+ yellow, medulla K—, C+ rose, KC+ red, 
P—, atranorine and gyrophoric acid present. 

Lynge reported his determination with doubt, but the material is 
fully typical. 

76. Parmelia portoalegrensis Lynge, Ark. Bot. 13, No. 13:147, pl. 4, fig. 13. 
1914, 

Lectotype: Porto Alegre, Rio Grande do Sul, Brazil, Malme 571, 
October 15, 1892 (S). 

Additional specimen examined: Malme 560. 

Reactions: Thallus K—, medulla K+ yellow, C—, KC—, P+ 
light orange, usnic and stictic acids present. 

The tropics abound in minute isidiate species of the section Xantho- 
parmelia Vain. One of the first to be described, P. adpressa Kremplh., 
must be rejected as an orthographic variant and later homonym of 
P. appressa Mey. & Flot. The holotype of P. adpressa (Rio de 
Janeiro, Glaziou 3842, M) contains usnic acid and stictic acid and 
appears in other respects to be typical P. isidiata (Anzi) Gyel. Miiller 
described a variety of this species, P. adpressa var. stenophylloides 
Mill. Arg., which Vainio raised to the rank of species as P. steno- 
phylloides (Mill. Arg.) Vain. This entity also contains stictic acid 
but seems to be amply distinct from P. isidiata by reason of the very 
narrow lobes (to 0.5 mm wide). It has been suggested (cf. des 
Abbayes, 1958, p. 5) that P. congensis Stein., published in 1889, is 
identical with P. stenophylloides and should be the correct name for 
this entity. I have not checked the type of P. congensis, an African 
species, but the description seems to conform to P. stenophylloides. 


HALE——REVISION OF PARMELIA SPECIES 29 


Parmelia portoalegrensis is hardly distinguishable from this narrow 

lobed entity, whatever its correct name, except for the more crowded, 

contiguous lobes, and is perhaps merely a growth form on very smooth 

rock faces. It should be considered provisionally as a synonym of 

P. congensis. 

77. Parmelia proboscidea Tayl. in Mack. Fl. Hibern. 2:143. 1836.—Lynge, 
1914, p. 38; 1925, p. 91. 

Type: Dunkerron Mountains, Ireland (FH); conspecific with the 
earlier P. crinita Ach. (cf. Hale, 1958a, p. 179). 

Reactions: Thallus K+ yellow, medulla K+ yellow, C—, KC—, 
P+ pale orange, atranorine and stictic acid present. 

Lynge followed Miller and Vainio in an incorrect interpretation 
of Taylor’s species. A. L. Smith, who was familiar with P. probosci- 
dea from Great Britain and knew that it was simply P. crinita, real- 
ized that the tropical plants so named were a different species. She 
gave them a new name, P. wainii A. L. Smith (Journ. Linn. Soc. 
London, Bot. 46:85, 1922), and cited as a basionym P. proboscidea 
(sensu Vainio, Acta Soc. Faun. & Fl. Fenn. 7:29, 1890). Vainio’s Lich. 
Bras, Exs. Nos. 400 (FH), 582B, 973, and 1000 are therefore syntypes 
of P. wainit. Two packets labeled P. proboscidea by Lynge (Malme 
314B and June 15, 1894) are identical with Lich. Bras. Exs. 400. 
Zahlbruckner (1904, p. 135) had actually described this entity earlier 
as P. proboscidea var. ornatula Zahlbr. (holotype: Serra do Ouro Preto, 
Damazio 1090, G; plate 3). Parmelia wainii is characterized by an 
ample thallus, marginally ciliate and without soredia or isidia; it 
produces atranorine and alectoronic acid (KC+ red). At the present 
time it is known from Brazil and Africa. 


78. Parmelia prolixa var, rosea Lynge, Nyt Mag. Naturv. 62:91. 1925. 


The specimen on which Lynge based this new variety was destroyed 
at Berlin. 


79. Parmelia radians Lynge, ad int., Ark. Bot. 13, No. 13:85. 1914. 


Holotype: Sao Jodo d’el Rey, Minas Gerais, Brazil, Malme 203, 
August 30, 1892 (S). 

Reactions: Thallus K+ yellow, medulla K+ red, C—, KC—, P+ 
orange-red, atranorine, salacinic acid, and usnic acid present. 

Parmelia radians should be rejected as a provisional new species 
“ad interim.’’ The type specimen is identical with P. delicatula 
Vain. (type: Lich. Bras. Exs. 1256, FH), although Lynge thought 
that Vainio’s plant had a deeper yellow color than his own. The rela- 
tion of P. delicatula to other tropical species with usnic acid in the 
section Amphigymnia is given in the key on p. 21. 


30 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


80. Parmelia regis Lynge, Ark. Bot. 13, No, 13:126. 1914. 


Lectotype: Sao Jodo d’el Rey, Minas Gerais, Brazil, Malme 178, 
August 30, 1892 (S). 

Range: Known only from the type locality. 

Reactions: Thallus K—, medulla K+ wine-red, C—, KC+ pale 
red, P-+ pale orange, lichexanthone and unidentified substances 
present. 

The type of P. regis cannot be told externally from P. silvatica 
Lynge (see p. 34); it differs chiefly in lacking an anthraquinone 
pigment in the lower medulla. The two species are separated in 
Lynge’s key (p. 21) by: “1. Thallus KOH superne non coloratur”’ 
(P. silvatica), and ‘2. Thallus superne flavescens” (P. regis). Lynge 
apparently established a yellow K test for P. regis from the two mis- 
identified syntypes discussed below, since the ‘“‘originaleksemplar”’ 
is K—. While P. regis thus delimited differs only in chemical charac- 
ters from P. silvatica, it should be retained as a separate species until 
we are more familiar with the range of chemical variation in this 
group. 

The other two specimens cited by Lynge (Malme 941 and 2393****) 
differ from the lectotype in lacking lichexanthone and reacting K+- 
yellow on the cortex and K— in the medulla. One specimen (2393 ****) 
is identical with P. palmarum Lynge (see p. 26 under P. palmarum 
Lynge), while the other, which Lynge had first called a new species 
in manuscript, P. cachoeirae, is too fragmentary for proper study. 


81. Parmelia regnellii Lynge, Ark. Bot. 13, No. 13:140, pl. 5, figs. 1, 2. 1914. 


Lectotype: Sao Joao d’el Rey, Minas Gerais, Brazil, Malme 308, 
September 1, 1892 (S). 

Range: Brazil. 

Additional specimens examined: Malme 179, 2748B, 2745B, s.n. 

Reactions: Thallus K—, medulla alba K+ yellowish, C+ yellowish, 
KC+ orange-yellow, P— (P+ in 2745B), medulla crocea K+ 
purple, usnic and barbatic acids (stictic in 2745B), an unidentified 
anthraquinone and other substances present. 

Parmelia regnellii is a most remarkable lichen with a deep yellow 
thallus and in part a deep saffron medulla. Although Malme col- 
lected the species at three widely separated localities, it has not 
appeared in the literature or been seen in herbaria since Lynge’s 
description. 

82. Parmelia regnellii f. arida Lynge, Ark. Bot. 13, No. 13:141. 1914. 


Holotype: Near Bocca da Serra, Serra da Chapada, Mato Grosso, 
Brazil Malme 2240, January 21, 1894 (S). 

Range: Known only from the type locality. 

Reactions: As in P. regnellia f. regnellar. 


HALE—REVISION OF PARMELIA SPECIES 31 


This form is much smaller than the typical form but has a similar 
chemistry. 
83. Parmelia revoluta Floerke, Deutsch. Lich. 1:11. 1815—Lynge, 1925, p. 92. 


Lynge based his record on two small fragments (Henschen in 1868, 
UPS), which in my opinion are not identifiable. Although P. revoluta 
has appeared in several lists of tropical American Parmelias, I doubt 
that the identifications are correct. 


$4. Parmelia rigida Lynge, Ark. Bot. 13, No. 13:50, pl. 2, fig. 2. 1914. 


Holotype: Piratiny, Rio Grande do Sul, Brazil, Malme 827B, 
December 17, 1892 (5). 

Range: Known only from the type locality. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC+ red, 
P—, atranorine, and alectoronic acid present. 

The exceedingly rigid thallus without soredia or isidia and the 
numerous large apothecia set this species apart from all other parme- 
lias. It may be placed in the P. melanothriz group (see. p. 23) because 
of the white-maculate cortex and the presence of alectoronic acid. 
Lynge mistakenly gave the medullary reaction as KC—. 

85. Parmelia riograndensis Lynge, Ark. Bot. 13, No.13:26, pl. 1, fig. 2. 1914. 
Parmelia microsticta Mill. Arg. var. riograndensis (Lynge) Lynge, Nyt 
Mag. Naturv. 62:90, 1925. 

Lectotype: Porto Alegre, Rio Grande do Sul, Brazil, Malme 461, 
September 25, 1892 (S). 

Reactions: Thallus K+ yellow, medulla K—, C—, KC—, P-, 
atranorine and protolichesteric acid present. 

Additional specimen examined: Malme 1282B. Reactions: Thallus 
K-+ yellow, medulla K—, C+ rose, KC+ red, P—, atranorine and 
gyrophoric acid present. . 

Lynge based his description on two packets, one labeled “original- 
eksemplar,”’ reacting C+ rose, which was erroneously designated by 
me (1958a, p. 180) as the holotype, and another reacting C—, which is 
the legitimate lectotype since it agrees with Lynge’s C test. The 
two chemical strains are morphologically indistinguishable and near, 
if not equal to, P. bolliana Mill. Arg., which has the same chemical 
strains in the southwestern United States (Culberson & Culberson, 
1956). A fuller understanding of P. riograndensis will depend on 
further study of P. mierosticta Mill. Arg. and P. squamuligera Sant., 
but provisionally it can be regarded as a synonym of P. bolliana. 


86. Parmelia rissoensis Lynge, Ark. Bot. 13, No. 13:69. 1914. 


Holotype: Near Rio Apa, Colonia Risso, Paraguay, Malme 1895B, 
October 7, 1893 (S). 


32 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Reactions: Thallus K+ yellow, medulla K—, C+ red, KC+ red, 
P—, atranorine and lecanoric acid present. 

This entity is conspecific with P. africana Miill. Arg. and a poorly 
developed state of its synonymous species, P. paraguariensis Lynge 
(see p. 27). 

87. Parmelia rudecta Ach. Syn. Lich. 197. 1814.—Lynge, 1914, p. 29; 1925, 
p. 92. 

Type: North America, Muhlenberg (PH, isotype). 

Reactions: Thallus K+ yellow, medulla K—, C+ red, KC+ red, 
P—, atranorine and lecanoric acid present. 

Additional specimens examined: Malme, July 9, 1894; Glaziou 
1834 (UPS). Both of these specimens contain gyrophoric acid 
(C+ rose). 

The specimens are identical in morphology with the typical North 
American P. rudecta, which produces lecanoric acid constantly 
(Culberson & Culberson, 1956). The significance of the chemical 
difference in South American specimens has not yet been evaluated. 


88. Parmelia rupicola Lynge, Ark. Bot. 13, No. 13:132, pl. 3, figs. 5, 6. 1914. 


Holotype: Porto Alegre, Rio Grande do Sul, Brazil, Malme1339, 
June 2, 1893 (S). 

Range: Known only from the type locality. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC—, P—, 
atranorine and divaricatic acid present. 

Parmelia rupicola mimics the crustose Lecanora muralis (Schreb.) 
Rabh. very closely in size and habit. Divaricatic acid is known in 
only one other Parmelia species, P. texana Tuck, a widely distributed 
corticolous species in temperate and tropical regions. 


89. Parmelia rupta Lynge, Ark. Bot. 18, No. 13:40. 1914. 


Holotype: Rio Negro, Gran Chaco, Paraguay, Malme, September 
14, 1893 (S). 

Range: Known only from the type locality. 

Reactions: Thallus K+ yellow, medulla K+ red, C—, KC—, P+ 
orange-red, atranorine and salacinic acid present. 

The single holotype collection has the appearance of P. cetrata Ach. 
with a reticulately rimose cortex. However, it has a rather distinct 
naked zone below at the margins. The species cannot be evaluated 
from this single rather poor specimen. 

90. Parmelia rutidota Hook. & Tayl. Lond. Journ. Bot. 3:645. 1844-—Lynge, 
1914, p. 151; 1925, p. 92. 
Type: Van Dieman’s Land (FH). 
Range: Australia, South America. 


HALE—REVISION OF PARMELIA SPECIES 33 


Reactions: Thallus K—, medulla K—, C—, KC+ rose, P+ 
orange-red, usnic and protocetraric acids present. 

Additional specimen examined: A4alme 715. 

The Malme specimen has the same morphology, habit, and spore 
size (13-16 X 7-10 ») as Taylor’s type, but it contains fumarproto- 
cetraric acid. It is probably no more than a chemical strain of little 
significance, and does not warrant species rank at this time. Par- 
melia rutidota is a plant of the southern hemisphere. Asahina’s 
report from Japan (1952, p. 142) is a misidentified species of the 
P. texana group. 


91. Parmelia rutidota f. filizans Lynge, Ark. Bot. 13, No. 13:153. 1914. 


Holotype: Quinta, near Rio Grande, Rio Grande do Sul, Brazil, 
Malme 727, December 3, 1892 (S). 

Range: Known only from the type locality. 

Reactions: Thallus K—, medulla K—, C—, KC— or brownish, 
P+ red, fumarprotocetraric acid and usnic acid present. 

This is a well characterized form with conspicuous filiform marginal 
laciniae. 

92. Parmelia saccatiloba Tayl. Lond. Journ. Bot. 6:174. 1847.—Lynge, 1914, 
p. 65. 

Lectotype: Pitcairn’s Island, Beechey (FH, cf. Hale, 1958, p. 180). 

Reactions: Thallus K+ yellow, medulla K—, C—, KC+ rose, P+ 
orange-red, atranorine and protocetraric acid present. 

The five specimens thus identified by Lynge (Malme 1678**, 2156B, 
2392**, s.n.) all contain atranorine and a fatty substance near caper- 
atic acid, and lack the isidia characteristic of P. saccatiloboa. They 
may be identified as P. mesotropa Mill. Arg. (type: Asuncién, 
Paraguay, Balansa in 1878, G, plate 4), a common species which Lynge 
failed to study. 

93. Parmelia saccatiloba f, membranacea Lynge, ad int., Ark. Bot. 18, No. 
13:67. 1914. 

Holotype: Pileomayo, Gran Chaco, Paraguay, Malme, September 
2, 1893 (S). 

Reactions: Thallus K++ yellow, medulla K—, C—, KC—, P-, 
atranorine and an unknown fatty substance near caperatic acid 
present. 

This form, which is not validly published, is also identical with P. 
mesotropa Mill. Arg. The smaller spores as reported by Lynge 
probably have no significance. 

94. Parmelia sancti-angelii Lynge, Ark. Bot. 13, No. 13:35, 1914.—Lynge, 1925, 
p. 92. PLATE 5 

Holotype: Colonia Santo Angelo, near Cachoeira, Rio Grande do 

Sul, Brazil, Malme, January 25, 1893 (S). 


34 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Range: Mexico, West Indies, South America, Africa, China. 

Additional specimens examined: Caldas, Minas Gerais, Brazil, 
Henschen in 1868 (UPS). 

Reactions: Thallus K+- yellow, medulla K—, C+ rose, KC+ red, 
P—, atranorine and gyrophoric acid present. 

Parmelia sancti-angelii is a distinctive species now known to be 
widespread in tropical regions. In size and general morphology, it 
resembles P. maxima Hue (P. claudelii (Harm.) Vain.), which con- 
tains salacinic acid (K+ red). There is considerable variance in 
spore size, which is difficult to assess in a species so often collected 
sterile. Liynge reported spores 26-33 yu long with a wide episporium; 
other tropical collections which I have measured have much smaller 
spores, only up to 16 » long, with a thin episporium. Differences of 
this magnitude are generally not expected. 


95. Parmelia semilunata Lynge, Ark. Bot. 13, No. 13:23, pl. 5, figs. 10-12. 1914. 


Holotype: Buriti, Serra da Chapada, Mato Grosso, Brazil, Malme, 
June 19, 1894 (S). 

Range: Known only from the type locality. 

Reactions: Specimens too fragmentary for testing; Lynge reported 
the thallus K+ yellow, medulla C—, K—. 

Externally P. semilunata could be mistaken for a species in the P. 
coronata group because of the small steel-gray thallus with coronate 
apothecia. The spores, however, are distinctly two-horned, a peculiar 
character known only in P. schiffneri Zahlbr. and P. bicornuta Mill. 
Arg., the latter species a much larger plant (plate 5) with lecanoric 
acid (type: Rio de Janeiro, Leyland, G). In view of the great mor- 
phological diversity of the three species it seems doubtful whether 
they should be joined together in a separate section of the genus 
(Section Bicornutae Lynge, Ark. Bot. 13, No. 13:23, 1914) solely on 
the basis of the unique spores. 


96. Parmelia silvatica Lynge, Ark. Bot. 13, No. 13:118, pl. 4, fig. 14. 1914. 


Lectotype: Santa Anna da Chapada, Mato Grosso, Brazil, Malme 
2393*, February 21, 1894 (S). 

Range: Brazil. 

Additional specimens examined: Malme, March 3, 1894, June 25, 
1894. 

Reactions: Thallus K—, medulla alba K—, C—, KC+ rose, P+ 
orange-red, medulla crocea K+ purple, lichexanthone, protocetraric 
acid, and an unidentified anthraquinone present. 

This well-circumscribed species is characterized by an unusual 
combination of chemical components. It is very near P. regis Lynge 


HALE—REVISION OF PARMELIA SPECIES 35 


(see p. 30), which lacks the pigment, and not far removed from species 
in the P. bahiana-P. palmarum group (see p. 26 under P. palmarum 
Lynge). 

97. Parmelia silvatica var, pinnata Lynge, Ark. Bot. 13, No. 13:120. 1914. 


Holotype: Santa Anna da Chapada, Mato Grosso, Brazil, Malme 
2393***, February 21, 1894 (S). 

Range: Known only from the type locality. 

Reactions: As in P. silvatica var. silvatica. 

This variety is a broader-lobed plant than var. silvatica. I doubt 
that it could be told from the typical variety in a larger series of 
specimens, and I therefore do not believe it deserves any taxonomic 
rank. 


98. Parmelia silvatica var. radiata Lynge, Ark. Bot. 13, No. 13:120. 1914. 


Holotype: Santa Anna da Chapada, Mato Grosso, Brazil, Malme 
2393**, February 21, 1894 (S). 

Range: Known only from the type locality. 

Reactions: As in P. silvatica var. silvatica. 

This variety differs from var. silvatica only in being more closely 
adnate to the bark, and in this respect it approaches P. crustacea 
Lynge (see p. 13) closely. I do not believe it deserves taxonomic 
status. 


99. Parmelia soredica Nyl. Flora 68:608. 1885.—Lynge, 1925, p. 92. 


A single specimen collected by Fries in Argentina (S) and cited 
by Lynge has not been available for study. 


100. Parmelia subproboscidea Lynge, Ark. Bot. 13, No. 13:36. 1914. 


Holotype: Assuncion, Paraguay, Malme 1678 in 1893 (S). 

Range: Paraguay. 

Reactions: Thallus K-+ yellow, medulla K—, C—, KC+ red, P—, 
atranorine and alectoronic acid present; cilia K+ violet. 

Parmelia subproboscidea has no relationship at all with P. pro- 
boscidea Tayl. (= P. crinita Ach.) but is a member of the P. mela- 
nothriz group (see under P. melanothriz, p. 23). It differs from 
P. laongii Lynge in having slightly smaller spores and a paler under- 
side. Parmelia melanothriz f. microspora Lynge (see p. 24) is referable 
to this species. The recognition of P. subproboscidea makes the 
typification of P. urceolata Eschw. (Icon. Pl. Crypt. 23, 1827) essen- 
tial. Though Eschweiler’s color plate is not as clear as we would 
like, there is a strong possibility that it is the same as P. subpro- 
boscidea. In any event, when the type of P. urceolata is located 
and studied, that name will doubtless replace one of those of the 
presently recognized members of the P. melanothriz group. 


36 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


101. Parmelia subregressa Lynge, ad. int., Ark. Bot. 13, No. 13:58. 1914. 
Parmelia subregressa, Lynge, Nyt. Mag. Naturv. 62:93. 1925. 

Holotype: Paraguari, Paraguay, Malme 1525B, August 7, 1893 (5). 

Reactions: Thallus K+ yellow, medulla K—, C—, KC—, P-, 
atranorine and a fatty substance present. 

Parmelia subregressa Lynge, ‘ad interim,” dating from 1914, must be 
rejected as a provisional name. The holotype is identical morpho- 
logically and chemically with P. mesotropa Mill. Arg. (see p. 38, 
under P. saccatiloba Tayl.). In the second publication dating from 
1925, Lynge listed the species without provisional status, so that this 
is the valid date of publication of P. subregressa. The specimen on 
which the second determination was based (Pampa Blanca, Argen- 
tina, Fries, S) has not been seen, but the type of the species would 
still be the Malme specimen. 


102. Parmelia subrugata Kremplh. var. arcuata Lynge, Ark. Bot. 13, No. 13:48. 
1914. 


Holotype: Porto Alegre, Rio Grande do Sul, Malme 440, September 
25, 1892 (S). 

Reactions: Thallus K-+ yellow, medulla K—, C—, KC+ red, 
P—, atranorine and alectoronic acid present. 

The holotype of P. subrugata Kremplh. (Serra dos Orgéos, Minas 
Gerais, Brazil, Helmreichen, M) has more or less arcuate conidia such 
as Lynge described for his new variety. Since the two entities are 
completely similar in morphology and chemistry, var. arcuata may 
be regarded simply as a synonym of var. subrugata. 

103. Parmelia sulphurata Nees & Flot. Linnaea 9:501. 1834,—Lynge, 1914, p. 
75; 1925, p. 94. 

Type: Destroyed at Berlin, but once examined by Hillmann (1939), 
who compared it with P. persulphurata Nyl. (see p. 28). 

Range: Tropical regions. 

Specimen examined: Malme 146. Reactions: Thallus K+ yellow, 
medulla K—, C—, KC—, P—, atranorine and vulpinic acid present. 

Lynge’s determination agrees with the present concept of the 
species. The density of isidia and marginal cilia varies widely. 
104. Parmelia tinctorum Nyl. Flora 55:547. 1872.—Lynge, 1914, p. 32; 1925, 

p. 94. 

Type: Nylander apparently based the species on a specimen from 
the Canary Islands collected by Despréaux (not seen). 

Range: Cosmopolitan in tropical and subtropical regions. 

Specimens examined: Malme 1889, 1895 Ac, 1480, 2369, 2243C, 196, 
and September 7, 1893. Reactions: Thallus K+ yellow, medulla K—, 
C+ red, KC+ red, P—, atranorine and lecanoric acid present. 


HALE—REVISION OF PARMELIA SPECIES 37 


Nylander should be cited as the author of this weedy tropical 
species, since Despréaux merely provided a specimen with an her- 
barium name. 


105. Parmelia uleana Mill. Arg. Flora 72:506. 1889.—Lynge, 1914, p. 155, pl. 
4, fig. 6. 

Type: Nova Cintra, Rio de Janeiro, Brazil, Ule 10 (G). 

Range: Southern United States, West Indies, Central and South 
America, and Africa. 

Additional specimens examined: Malme 61 and August 16, 1894; 
Malme 91, 2366, 2418, 2435B, 2519, 2642, 2742, and June 16, 1894. 

Reactions: Thallus K—, medulla K+ yellowish, C—, KC+ pale 
orange, P—, usnic and barbatic acids, and an unidentified yellowish 
pigment in the medulla present. The first two specimens cited con- 
tain stictic acid (P+ pale orange) instead of barbatic acid. 

Parmelia uleana is a synonym of P. sphaerospora Nyl. (Hale, 
1959a, p. 129), a very common tropical species distinguished by nearly 
spherical spores, a thick minutely rugulose cortex, and lack of soredia 
or isidia. The chemical composition is not perfectly clear at present. 
106. Parmelia urceolata Eschw. Icon. Pl. Crypt. 23. 1827.—Lynge, 1914, p. 

64; 1925, p. 95. 

Type: Not seen, but illustrated by Eschweiler (see under P. sub- 
proboscidea Lynge, p. 35). 

The two specimens determined by Lynge (Malme 827, and Zarati, 
Argentina, September 1894) contain atranorine and stictic acid and 
should be referred to P. eciliata (Nyl.) Nyl. (see p. 15). 

107. Parmelia viridescens Lynge, Ark. Bot. 13, No. 13:117, pl. 3, figs. 9, 10. 
1914. 

Holotype: Santa Anna da Chapada, Mato Grosso, Brazil, Malme 
2453, February 28, 1894 (S). 

Range: Known only from the type locality. 

Reactions: Thallus K—, medulla K—, C—, KC—, P—, no lichen 
substances proved. 

The type is a pale greenish plant without soredia or isidia. It 
resembles no other species known to me. 


108. Parmelia wainioana Lynge, Ark. Bot. 13, No. 13:87. 1914. 


Holotype: Santa Anna da Chapada, Mato Grosso, Brazil, Malme 
2435C, February 27, 1894 (S). 

Reactions: Thallus K+ yellow, medulla K—~, C—, KC—, P-, 
atranorine and perlatolic acid present. 

This species is synonymous with P. caroliniana Nyl., a common 
corticolous species from the southern United States southward to 
Brazil (Hale, 1959a, p. 17). 


38 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


109. Parmelia xanthina (Miill. Arg.) Vain. Ann. Soc. Faun. & Fl. Fenn. 7:37. 
1890. —Lynge, 1914, p. 85. 
Parmelia proboscidea var. zanthina Mill. Arg. Flora 67:616. 1884. 
P. perlata var. xanthina (Mill. Arg.) Stizb. Ber, St. Gall. Naturw. Gesell. 
1888-1889:156. 


Type: Central Madagascar, Hildebrandt (G). 

Range: Brazil, Madagascar. 

Reactions: Thallus K+ yellow, medulla K—, C—, KC—, P-, 
atranorine, and usnic and protolichesteric acids present. 

Des Abbayes (1958, p. 21) gave considerable attention to the 
status of this species. Although he was unable to typify Miiller’s 
var. zanthina satisfactorily, he referrred plants reacting C— in the 
medulla to P. aberrans (Vain.) des Abb., and the plants reacting C+ 
rose to P. zanthina Vain. nec Mill. Arg. The latter citation is 
incorrect inasmuch as any new combination is based on the type of 
the basionym, regardless of whether the combining author saw it 
or not. A specimen labeled P. perlata var. xanthina Mill. Arg. which 
I received from Geneva and which I believe is the holotype of P. 
proboscidea var. zanthina reacts C—, as indicated above. Therefore, 
P. aberrans is only a synonym of P. zanthina. Plants reacting KC+ 
rose have been identified with P. madagascariacea (Hue) des Abb. 
(des Abbayes, 1958, p. 22), Plants reacting C+ rose, such as 
Vainio’s Lich. Bras. Exs. 1181 and the two Malme specimens seen by 
Lynge (Malme 2748 and June 22, 1894) are indistinguishable from 
both P. madagascariacea and P. zanthina (C—) except for the C+ 
rose test caused by gyrophoric acid. JI would prefer to rank the 
C+ plants as a chemical strain of P. zanthina for the present. It 
may be desirable to recognize them as a distinct species in the future 
when the whole P. ranthina group (see under P. magna Lynge, p. 20) 
has been more precisely delimited. 

110. Parmelia zahlbruckneri Lynge, Ark. Bot. 13, No. 13:325, pl. 4, figs. 1, 2. 
1914. 

Lectotype: Near Bocca da Serra, Serra da Chapada, Mato Grosso, 
Brazil, Malme, June 15, 1894 (S). 

Reactions: Thallus K+ yellow, medulla K+ red, C—, KC+ 
reddish, P+ pale orange, atranorine, and unknown substances 
present. The acetone extract is a pale orange crust. 

Parmelia zahlbruckneri was based on two specimens. The one 
labeled ‘“Originaleksemplar”’ and photographed (pl. 4, figs. 1, #) 
by Lynge (Malme, same data as the lectotype above) reacts K-, 
C—, KC-+ rose in the medulla and is best referred to P. palmarum 
Lynge (see under P. palmarum Lynge, p. 26). The other syntype, 


HALE—REVISION OF PARMELIA SPECIES 39 


here designated the lectotype since it conforms better with the 
original diagnosis, is identical in every respect with P. gracilis (Mill. 
Arg.) Vain. Since P. gracilis is a later homonym and P. confusula 
Zahlbr., a new name, was published in 1929, P. zahlbrucknert becomes 
the correct name for this entity (see under P. gracilis, p. 17). 


Summary of Synonymy, Misidentifications, and Changes in 
Nomenclature 


Lynge identified a total of 110 taxa, 99 of which I have been able 
to verify. Of the 61 new taxa proposed by Lynge, two (P. hieronymi 
and P. proliza var. rosea) were destroyed at Berlin, and one (P. 
coccinea) was transferred by Lynge himself to Pyzxine coccifera (Fée) 
Nyl. Lynge also made two new combinations, P. brasilana var. 
glaziovit (Mill. Arg.) Lynge and var. novella (Vain.) Lynge. Two 
infraspecific taxa (P. abstrusa f. laevigata Lynge and P. cornuta var. 
crocea Lynge) have been raised to species rank by Gyelnik, and a 
later homonym (P. merrillii Lynge non Vain.) has received a new 
name. I have reduced 24 species, varieties, or forms to synonymy 
in the above list of species. A summary of these changes follows: 


. abstrusa f. laevigata Lynge=P. subabstrusa Gyel. 
. annae Lynge=P. recipienda Nyl. 
. ceracea Lynge=P. recipienda Nyl. 
. coccinea Lynge= Pyzine coccifera (Fée) Nyl. 
. cornuta var. crocea Lynge= P. crocea (Lynge) Gyel. non Sprengl. 
. fragilis Lynge= P. intercalanda Vain. 
latissima f. microspora Lynge=P. wainit A. L, Smith 
longiconida Lynge= P. scrobicularis Kremplh, 
malmet Lynge=P. minima Lynge, probably 
melanothrix {. microspora Lynge (invalid name)=P. subproboscidea 
Lynge 
merrillii Lynge, non Vain.=P. lyngeana Zahlbr. 
. osseo-albida Lynge (invalid name) = P. zahlbrucknert Lynge 
palmarum Lynge (invalid name) =P. palmarum ex Hale 
paraguariensis Lynge=P. africana Mill. Arg. 
. portoalegrensis Lynge= P, congensis Stein. provisionally 
radians Lynge (invalid name) =P. delicatula Vain. 
riograndensis Lynge= P. bolliana Mill. Arg., probably 
. rissoensis Lynge=P. africana Mill. Arg. 
. saccatiloba f. membranacea Lynge (invalid name)=P. mesotropa Mill. 
Arg. 
. silvatica var. pinnata Lynge= P. silvatica var. silvatica 
. silvatica var, radiata Lynge= P. silvatica var. silvatica 
. subregressa Lynge=P. mesotropa Miill. Arg. 
. subrugata Kremplh. var. arcuata Lynge=P. subrugata var, subrugata 
. wainioana Lynge= P. caroliniana Nyl. 


wu My uVUDUYD Ty vy yyy 


40 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Misidentifications and changes in nomenclature include the follow- 
ing 20 taxa: 


P, abstrusa f. laevigata Lynge pr. p.=P. ef. relicina Fr. 

P. balansae var. sorediata Mill. Agr.=P. subbalansae Gyel. 

P. conspersa (Ach.) Ach.=P. isidiata (Anzi) Gyel.+ P. flavida Zahlbr. 

P, cyliphora (Ach.) Vain.=P. caperata (L.) Ach. 

P. flava Kremplh. var. stellata Lynge=P. flavida Zahlbr var. stellata (Lynge) 

Zahlbr. 

P. flava var. subdichotoma Lynge=P. flavida Zahlbr. var. subdichotoma 
(Lynge) Zahlbr. 

. gracilis (Mill. Arg.) Vain. =P. zahlbruckneri Lynge 

. isidiophora Zahlbr.= P. caroliniana Ny). 

. kamtschadalis var. americana (Mey. & Flot.) Nyl.=P. americana (Mey. 

Flot.) Mont. 

. laceratula Nyl.=Parmelia sp. near P. riograndensis Lynge and P. 

squamuligera Sant. 

. latissima Fée pr. p.=P. zollingeri Hepp 

magna Lynge pr. p.=P. microdactyla Hale 

melanothriz (Mont.) Vain. pr. p.=P. argentina Kremplh. 

. pachyderma Hue=P. fistulata Tayl. 

. perforata (Jacq.) Ach.= P. leucosemotheta Hue+ P. subcaperata Kremplh. 

. persulphurata Nyl.=P. cornuta Lynge, probably. 

. proboscidea Tayl.=P. wainit A, L. Smith 

. saccatiloba Tayl.=P. mesotropa Mill. Arg. 

. uleana Mill. Arg.=P. sphaerospora Ny). 

. urceolata Eschw.= P. eciliata (Nyl.) Nyl. 


VeVyVUVV TY TY VU 


Four other species, P. consimilis Vain., P. eristfera Tayl., P. mi- 
narum Vain., and P. revoluta Floerke, were also misidentified, but it 
was impossible to correct the names because of the poor condition of 
the specimens. 


Bibliography 


AsAHINA, Y. 1952. Lichens of Japan. II. Genus Parmelia. Res. Inst. Nat. 
Resources. 1-172. Tokyo. 
————. 1954. Chemistry of lichen substances. 1-240. Tokyo. 
CuLperson, W. L., anp C. F. Cuntserson. 1956. The systematics of the 
Parmelia dubia group in North America. Amer, Journ. Bot. 43:678-687. 
Des Apsayes, H. 1958. Lichens récoltés en Guinée Francaise et en Cdéte 
d’Ivoire. IX. Supplement aux Parmeliacées. Bull. Inst. Fr. Afr. Noire 
20: 1-27. 

Du Rietz, G. EB. 1924. Flechtensystematische Studien. IV. Bot. Not. 1924: 
329-342. 

GyeLnik, V. 1936. Revisio typorum ab auctoribus variis descriptorum. 
II. Ann. Mus. Nat. Hungar. Bot. 30:119-135. 


HALE—REVISION OF PARMELIA SPECIES 41 


Haug, M.E., Jr. 1955. Xanthoparmelia in North America. I. The Parmelia 
conspersa-stenophylla group. Bull. Torrey Bot. Club 82:9-21. 

—. 1956. Fluorescence of lichen depsides and depsidones as a taxonomic 

criterion. Castanea 21:30—32. 

1958a. Chemical components of type specimens in Parmelia. I. 

Brittonia 10:177-180. 

1958b. The occurrence of Parmelia formosana in North America. 

Castanea 23:89, 80. 

1959a. New or interesting species of Parmelia in North America. 

Bryol. 62:16—-24. 

—. 1959b. New or interesting Parmelias from North and tropical 
America. Bryol. 62:123-132. 

HittMaNnn, J. 1939. Bemerkungen iiber einige Arten der Flechtengattung 
Parmelia. I. Hedwigia 78:249-267. 

Lynaz, B. 1914. Die Flechten der ersten Regnellschen Expedition. Die 
Gattungen Pseudoparmelia gen. nov. und Parmelia Ach. Ark. Bot. 13, 
No. 13:1-172. 

————. 1917. Ueber einige Regnellschen Parmelian aus Matto Grosso, 
Brasilien. Ark. Bot. 15, No. 1:1-4. 

—. 1925. On some South American lichens of the genera Parmelia, 
Candelaria, Teloschistes, and Pyxine. Nyt Mag. Naturv. 62:83-97. 

Maenusson, A. H. anp A. ZAHLBRUCKNER. 1944. Hawaiian lichens. II. The 
families Lecideaceae to Parmeliaceae. Ark. Bot. 31A, No. 6:1-109. 

Vainio, E. A. 1886. Revisio lichenum in herbario Linnaei asservatorum. 
Medd. Soe. Faun. & Fl. Fenn. 16:1-10. 

1896. Lichenes Antillarum a W. R. Elliott collecti. Journ. Bot. 
Brit. & For. 34:31-36. 

1900. Reactiones lichenum J. Miillero Argoviensi descriptorum. 
Mem. Herb. Boissier. No. 5:1-17. 

Wacutmerster, C. A. 1956. Identification of lichen acids by paper chroma- 
tography. Bot. Not. 109:313-324. 

ZAHLBRUCKNER, A. 1904. Lichenes a cl. Damazio in montibus Serra do 
Ouro Preto Brasiliae lecti. Bull. Herb. Boissier, ser. 2. 4:134-136. 

————. 1929. Catalogus lichenum universalis. 6:1-618. Leipzig. 


U.S. GOVERNMENT PRINTING OFFICE: 1960 


UNITED STATES NATIONAL MUSEUM 


CONTRIBUTIONS FROM THE UNITED STATES NATIONAL HERBARIUM 


VoLumeE 35, Part 2 


MARINE ALGAE FROM THE TROPICAL 
ATLANTIC OCEAN: 
V. ALGAE FROM THE LESSER ANTILLES 


By Wittiam RANDOLPH TAYLOR 


BULLETIN OF THE UNITED StTaTES NATIONAL Museum 


SMITHSONIAN INSTITUTION e WASHINGTON, D.C. e 1962 


MARINE ALGAE 
FROM THE TROPICAL ATLANTIC OCEAN: 
V. ALGAE FROM THE LESSER ANTILLES 


WILu1AM Ranpo.tpn TAYLOR 


Introduction 


In preparing a general account of the marine algae of the West 
Indies and neighboring coasts the writer has had many occasions to 
note apparent discontinuities of range, or ranges of conspicuous algae 
much less extensive that it seems reasonable to expect. One who 
knows the West Indian algal flora well discounts these deceptive 
ranges, but is nevertheless restricted in the range he can cite for species. 
The algae of very few places in the West Indies or nearby have been 
thoroughly studied; even those places that have been visited and the 
algae superficially studied are widely scattered. 

The marine algae of the Virgin Islands (while still the Danish West 
Indies) were thoroughly studied and described by Bgrgesen (1913-20), 
and his papers are the most informative of any dealing with West 
Indian algae. Famous among the algal works on the Lesser Antilles 
are the books based on the collections of A. Schramm and H. Mazé 
in Guadeloupe (Schramm and Mazé 1865, 1866; Mazé and Schramm 
1870-77), the identifications having been made by the brothers 
Crouan, well-known phycologists of Brest. These are rare books in 
the original issues. The collections were made with great discern- 
ment, but the identifications took little account of the great varia- 
bility of many of these plants, and the publications while proposing 
scores of new species seldom provided descriptions of the plants. It 
appears that no definitive set of specimens was set aside; several 
institutions possess considerable series of them but the specimens 
under a given collection number are not always identical as to species. 
Hamel and Hamel-Joukov supplemented these works later (1929, 
1931), as did Questel (1951). 

Grieve gave a brief list of Dominican algae (1909). The algae of 
Martinique have often been collected but reported only in scattered 
notes. A study of the algae of Barbados (Vickers 1905, 1908) yielded 
a rather limited list and a handsome atlas of selected Chlorophyceae 
and Phaeophyceae. The writer has reported on small collections 
from the Lesser Antilles in a few papers (1929, 1940, 1942). Lists 

43 


44 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


from several sources appear in the reports of the famous H.M.S. 
“Challenger” expedition, but they are in part simply recapitulations 
of earlier-published accounts. 

The only comprehensive catalog of West Indian (including Lesser 
Antilles) algae is that of Murray (1888, 1889) which brings together 
all reports up to that date in one list, but unfortunately it is quite 
uncritical, including records almost without question or verification, 
in particular those of Schramm and Mazé without remark, even when 
their names are nomina nuda. The writer in his recent (1960) general 
work has attempted to select the most reliable reports from the Lesser 
Antilles as well as other West Indian and mainland countries and to 
segregate those which are doubtful. 

It is very satisfying to be able to report now on useful collections 
from the British Virgin Islands, Barbuda, Nevis, and Antigua, and 
even more substantial ones from St. Kitts, St. Lucia, and Grenada, 
which in the very great majority of cases provide the first records of 
these plants from those islands, and in several cases extend the known 
ranges of the plants along the chain. These collections did not come 
to hand in time to be studied and the records entirely incorporated in 
the general account mentioned above (Taylor 1960), though it was 
possible to introduce a very considerable number of the Grenada 
records and a few others of special interest. No general conclusions 
can be derived from these collections—they fill in gaps in our knowl- 
edge but do not change our general picture of the West Indian algal 
flora. 

The material from the Smithsonian-Bredin Caribbean Expeditions 
of 1958 and 1959 was submitted through the kindness of Dr. Fenner 
A. Chace, Jr., Curator of Marine Invertebrates, U.S. National 
Museum, and Dr. Waldo L. Schmitt, Emeritus Head-Curator of 
Zoology and one of the collectors on the expeditions. That from the 
Institute of Jamaica-Arnold Arboretum (Harvard University) Ex- 
pedition was submitted by C. Bernard Lewis, Director of the Institute, 
on behalf of the collector, Mr. George R. Proctor and of Mrs. Lena 
Green, the curatorial assistant who prepared the material for study. 
To all of these the writer is exceedingly grateful for the opportunity of 
studying the specimens. The first set of the Smithsonian-Bredin 
collection is in the U.S. National Museum, and that of the plants 
collected by Mr. Proctor in the Science Museum of the Institute of 
Jamaica, 

Localities 


As a large number of species are involved and as it is impossible in 
any case to designate the precise environment of individual speci- 
mens, since they were grouped together in large jars, a general charac- 
terization of each major station will be given below. The individual 


CONTR. U.S. NAT. HERB. VOL. 36 TAYLOR—PLATE 1 


Figures 1-2.—-Sarcodtotheca caribaea; 1, spermatangial plant, 1.35; 2, type, cysto- 


carpic plant, 1.35. 


CONTR. U.S. NAT. HERB. VOL. 36 TAYLOR —PLATE 2 


Fictres 1-2.—-Meristotheca tobagensis: 1, cystocarpie plant, 1.7; 2, type, sperma- 


rangial plant, < 1.9. 


TAYLOR—PLATE 3 


U.S. NAT. HERB. VOL. 36 


CONTR. 


Vl 


0 yuryd duiodseqay rene! ‘THT urd sidivsoisia ‘T Spunyjizup viopyy ff 


C 


] SAMI] 


CONTR. U.S. NAT. HERB, VOL. 26 TAYLOR—PLATE 4 


Figures 1-2-1, Sarcodiotheca carthaea, tetrasporic plant, X13. 2. Waldoia antillana, 


detail of part of a tetrasporic plant showing stichidia, * 2.3. 


TAYLOR—MARINE ALGAE FROM LESSER ANTILLES 45 


numbers for the Smithsonian specimens were supplied by the writer 
when mounting the plants for study; those with the Proctor specimens 
were assigned by the Science Museum. 


SMITHSONIAN-BREDIN CARIBBEAN EXPEDITION oF 1958 


The algae of the Smithsonian-Bredin Expeditions were collected 
incidental to other tasks by various persons, particularly by Dr. Waldo 
L. Schmitt, to whose continuing interest in all living things of the 
shore, and generosity in contributing the plants for study, the writer 
has been deeply indebted for many years. Dr. Schmitt collected at 
most of the localities; Dr. Richard S. Cowan collected at Pigeon 
Island, St. Lucia, at some of the Barbuda localities, and Man-of-War 
Bay, Tobago, and Dr. Paul Denckla collected the algae at Antigua. 

Station 9-58. British Virgin Islands, B.W.I., Guano Island, White 
Bay. Reefs in a cove, from honeycombed rocks and Porites clumps, 
just off shore and at a depth of about 2 meters, Mar. 28, 1958. 
Collector: W. L. Schmitt. 

Stations 18 & 21-58. British Virgin Islands, B.W.I., Peter Island, 
Little Bay. Dredged from 4.5-15.0 meters, Mar. 29, 30, 1958. 
Collector: W. L. Schmitt. 

Station 42-58. British Virgin Islands, B.W.I., Anegada Island, 
Pomato Point. Collected over the edge of a reef and on eel-grass 
flats at depths to one meter, Apr. 8, 1958. Collector: W. L. Schmitt. 

Station 67-58. Nevis, B.W.J., Charlestown. Dredged about the 
anchorage over a weedy bottom at 3.6-5.5 meters, Apr. 16, 1958. 
Collector: W. L. Schmitt. 

Station 113a-58. Barbuda, B.W.I., Gravenor Landing. Col- 
lected along shore in honeycombed coral rock and limestone, and in 
turtle grass beds, Apr. 28, 1958. Collector: W. L. Schmitt. 


SMITHSONIAN-BREDIN CARIBBEAN EXPEDITION oF 1959 


Tobago, B.W.I., Man-of-War Bay, Apr. 7, 1959. Collector: R. 8. 
Cowan. 

St. Lucia, B.W.I., Marigot Bay, Apr. 14, 1959. Collectors: W. L. 
Schmitt et al. 

St. Lucia, B.W.I., Pigeon Island, Apr. 16, 1959. Collector: R. S. 
Cowan. 

Barbuda, B.W.I., Barbuda lagoon, Apr. 25, 1959. Collector: R.S. 
Cowan. 

Barbuda, B.W.I., Cocoa Point, Apr. 26, 1959. Collector: R. 8S. 
Cowan. 

Station 119-59. Antigua, B.W.I., Reeds Beach, a private beach 
in the Mill Reef Club area north of English Harbor, May 4, 1959. 
Collector: Paul Denckla. 


46 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


INSTITUTE OF JAMAICA-ARNOLD ARBORETUM EXPEDITIONS OF 1958, 1959 


Collector for all stations, George L. Proctor. No station numbers 
were supplied. 

St. Kitts, B.W.I., Black Rocks. Tidepools in rough lava rock, 
subject to violent wave action, Mar. 14, 1959. 

St. Kitts, B.W.I., Conaree Beach. Attached to rocks lightly 
covered by shifting sand at depths of 1.5-3.0 dm., Mar. 28, 1959. 

St. Kitts, B.W.I., Frigate Bay. Attached to boulders at depths of 
1.5-3.0 dm., Mar. 29, 1959. 

St. Kitts, B.W.I., North Friars Bay. On ledges and in tidepools 
washed by violent surf, Mar. 29, 1959. 

St. Kitts, B.W.I., South Friars Bay. On partly submerged rocks 
accompanied by some sand, Mar. 29, 1959. 

Nevis, B.W.I., near Red Cliff. Collected along a rocky shore at 
depths of 1.5-3.0 dm., Mar. 20, 1959. 

St. Lucia, B.W.I., Marquis Bay. Collected along rocky shores, 
May 16, 1958. 

St. Lucia, B.W.I., Vigie Point. From rocky ledges in heavy surf, 
June 1, 1958. 


LIST OF SPECIES 
Myxophyceae 
Oscillatoriaceae 
Lyngbya majuscula Harv. ex Gom. Ann. Sci. Nat., Bot. VII, 16: 131. 1892. 


Antigua: Reeds Beach, coll. Denckla, Smiths.-Bredin no. 119-59a. A very 
widely distributed species.! 


Chlorophyceae 


Ulvaceae 


Enteromorpha flexuosa (Wulf.) J. Ag. Lunds Univ. Arsskr., Afd. Math. Natur- 
vetensk. 19(2): 126. 1883. 
Grenapva: True Blue, coll. Proctor no. A-2699. Range from Bermuda and 
North Carolina to Uruguay. 


Ulva fasciata Delile. Flore d’Egypte 153. 1813. 


Nevis: Charlestown, coll. Schmitt, Smiths.-Bredin no. 67-58a. Range from 
Bermuda and North Carolina to Uruguay. 


Cladophoraceae 


Chaetomorpha brachygona Harv. Smiths. Contr. Knowl. 10: 87. 1858. 
Nevis: Charlestown, coll. Schmitt, Smiths.-Bredin no. 67-58e. Range from 
Bermuda and North Carolina to Brazil. 


Chaetomorpha crassa (C. Ag.) Kitz. Phycol. Germ. 204. 1845. 
St. Kirts: Frigate Bay, coll. Proctor no. A-3166. The easternmost record to 
date. Range hitherto Bermuda, Virgin Islands, and St. Barthélemy. 


1 The range as given applies only to the distribution of these plants in the warmer latitudes of the western 
Atlantic Ocean. Some of them occur in other seas as well. 


TAYLOR—MARINE ALGAE FROM LESSER ANTILLES 47 


Chaetomorpha media (C. Ag.) Kiitz. Spec. Alg. 380. 1849. 

British Virain Isuanps: Guano Island, coll. Schmitt, Smiths.-Bredin no. 2. 
Sr. Kitts: Black Rocks, coll. Proctor no. A-3206; Conaree Beach, coll. Proctor 
no. A-3136. Str. Lucia: Marquis Bay, coll. Proctor no. A-2782; Vigie Point, 
Proctor no. A-2790. Range from Bermuda to Brazil. 


Cladophora fascicularis (Mert.) Kiitz. Linnaea 17: 91. 1843. 

BarBupa: Cocoa Point, coll. Smiths.-Bredin no. 84. Sr. Kirrs: Black Rocks, 
coll. Proctor no. A-3198; S. Friars Bay, coll. Proctor no. A-3173. Sr. Lucta: 
Marguis Bay, Proctor no. A-2775. Range from Bermuda and North Carolina 
to Uruguay. 


Cladophora fuliginosa Kiitz. Spec. Alg. 415. 1849. 
Sr. Kirrs: Conaree Beach, coll. Proctor no. A-3121. The easternmost record 
to date. Range hitherto from Bermuda to the southern Netherlands Antilles. 


Dasycladaceae 


Batophora oerstedii J. Agardh, Ofvers. K. Vetensk.-Akad. Férhandl., 11: 108. 
1854. 
BarRBupa: Barbuda Lagoon, coll. Cowan, Smiths.-Bredin no. 75. Range from 
Bermuda to British Honduras. 


Acetabularia crenulata Lamour. Hist. Polyp. Corall. Flex. 249. 1816. 
BarBupa: Barbuda Lagoon, coll. Cowan, Smiths.-Bredin no. 70. Range from 
Bermuda to Venezuela. 
Valoniaceae 


Halicystis osterhoutii L. R. & A. H. Blinks, Bull. Torrey Bot. Club, 57: 389. 
1930. 
Antigua: Reeds Beach, coll. Denckla, Smiths.-Bredin no. 119-59m. Range 
from Bermuda to the Grenadines, but at few stations. 


Valonia macrophysa Kiitz. Phycol. Gen. 307. 1863. 

British Virgin Is.tanps: Anegada Island, Pomato Point, coll. Schmitt, 
Smiths.-Bredin no. 42-58b. A slight northeastward range extension. Range 
hitherto from Bermuda to the American Virgin Islands. 


Valonia ventricosa J. Ag. Lunds Univ. Arsskr., Afd. Math. Naturvetensk., 23(2). 
96. 1887. 
British Virain Isuanps: Anegada Island, Pomato Point, coll. Schmitt, 
Smiths.-Bredin no. 42-58a. Range from Bermuda to Brazil. 


Dictyosphaeria cavernosa (Forssk.) Bgrg. Dansk Bot. Ark. 8(2): 2. 1932. 

British Viren Istanps: Anegada Island, Pomato Point, coll. Schmitt, 
Smiths.-Bredin no. 42-58c. Sr. Kirrs: Black Rocks, coll. Proctor no. A-3190; 
Conaree Beach, coll. Proctor no. A-3137. ANm@ua: Reeds Beach, coll. Denckla, 
Smiths.-Bredin no. 119-59g. Range from Bermuda to Brazil. 


Cladophoropsis membranacea (C. Ag.) Bgrg. Overs. K. Danske Vidensk. Selsk: 
Forhandl. 1905(3): 288. 1905. 

Sr. Kirrs: Black Rocks, coll. Proctor no. A-3199. AnNntTiaua: Reeds Beach. 
coll. Denckla, Smiths.-Bredin no. 119-59f. Sr. Lucia: Pigeon Island, coll. 
Cowan, Smiths.-Bredin no. 48. Grenapa: Grenada Bay, coll. Proctor no. 
A-2741. Range from Bermuda to Brazil. 


Anadyomene stellata (Wulf.) C. Ag. Spec. Alg. Rite Cogn. 1: 400. 1822. 
Barsupa: Barbuda Lagoon, coll. Cowan, Smiths.-Bredin no. 74. Range from 
Bermuda to Brazil. 


48 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Bryopsidaceae 


Bryopsis pennata Lamour. Nouv. Bull. Sci. Soc. Philom. 1: 333, 1809. 
Sr. Kirts: Black Rocks, coll. Proctor no. A-3191. Range from Bermuda to 
Brazil. 
Caulerpaceae 


Caulerpa cupressoides (West) C. Ag. Spec. Alg. Rite Cogn. 1: 441. 1822. 
Nevis: Red Cliff (v. lycopodium (J. Ag.) Weber-van Bosse f. alternifolia 
Weber-van Bosse), coll. Proctor no. A-3209; (v. Lycopodium f. disticha (Weber-van 
Bosse) Collins), coll. Proctor no. A~3220; (v. Lycopodium toward f. elegans (Crouan) 
Weber-van Bosse), coll. Proctor no. A-3223. Sr. Lucra: Marigot Bay (v. 
cupressoides), coll. Schmitt, Smiths.-Bredin no. 42: Pigeon Island (v. lycopo- 
dium), coll. Cowan, Smiths.-Bredin no. 51. Range from Bermuda to Brazil. 


Caulerpa microphysa (Weber-van Bosse) J. Feldm. Rev. Gen. Bot. 62: 424. 
1955. 
Sr. Lucra: Pigeon Island, coll. Cowan, Smiths.-Bredin no. 46. Range from 
Bermuda to Venezuela. 


Caulerpa prolifera (Forssk.) Lamour. Nouv. Bull. Sci. Soc. Philom. 1: 332. 
1809. 

GrenapDa: Grenada Bay, Proctor A-2718. Range from Bermuda and Florida 
to Brazil. 


Caulerpa racemosa (Forssk.) J. Ag. Lunds Univ. Arsskr., Afd. Math. Natur- 
vetensk. 9(8): 35. 1873. 

British VirGin Isuanps: Guano Island, coll. Schmitt, Smiths.-Bredin no. 1. 
Barsupa: Cocoa Point, coll. Cowan, Smiths.-Bredin no. 85. Sr. Lucra: 
Marquis Bay, coll. Proctor no. A-2778; Pigeon Island, coll. Cowan, Smiths.-Bredin 
no. 52; Vigie Point, coll. Proctor no. A-2796. Range from Bermuda and Florida 
to Brazil. 


Caulerpa sertularioides (Gmel.) Howe, Bull. Torrey Bot. Club 32: 576. 1905. 

BarRBupDa: Cocoa Point, partly approaching v. farlowit (Weber-van Bosse) 
Berg., coll. Cowan, Smiths.-Bredin no. 86. Sr. Kirrs: Conaree Beach (partly 
f. brevipes (J. Ag.) Sved.), coll. Proctor no. A-3140; South Friars Bay, coll. 
Proctor no. A-3176. Nevis: Charlestown, coll. Schmitt, Smiths.-Bredin no. 
67-58d. Sr. Lucia: Pigeon Island (f. brevipes (J. Ag.) Sved.), coll. Cowan, 
Smiths.-Bredin no. 47; Vigie Point, coll. Proctor A-2792. Range from Bermuda 
and Florida to Brazil. 


Caulerpa verticillata J. Ag. Ofvers. K. Vetensk.-Akad. Férhandl. 4: 6. 1847. 
GrenaDa: True Blue, coll. Proctor A-2701. Range from Bermuda and 
Florida to Brazil. 
Codiaceae 


Avrainvillea nigricans Dec. Ann. Sci. Nat., Bot., II, 18: 108. 1842. 
Sr, Lucta: Vigie Point, coll. Proctor no. A-2813 (in part). Range from 
Bermuda and Florida to Brazil. 


Avrainvillea rawsonii (Dickie) Howe, Bull. Torrey Bot. Club 34: 510. 1907. 
Sr. Kirrs: Conaree Beach, coll. Proctor no. A-3115. Sr. Lucia: Vigie Point, 
coll. Proctor no. A-2806. Range from Bermuda to Barbados. 
Udotea flabellum (Ell. & Sol.) Lamour. Nouv. Bull. Sci. Soc. Philom. 3: 180. 
1812. 
BarBupva: Cocoa Point, coll. Cowan, Smiths.-Bredin no. 83; Barbuda Lagoon 
coll. Cowan, Smiths.-Bredin no. 72. ANntTiaua: Reeds Beach, coll. Denckla, 


TAYLOR~~MARINE ALGAE FROM LESSER ANTILLES 49 


Smiths.-Bredin no. 119-59e. Sr. Lucia: Pigeon Island, coll. Cowan, Smiths.- 
Bredin no. 49. Range from Bermuda and Florida to Brazil. 


Penicillus capitatus Lamarck, Ann. Mus. Hist. Nat. 20: 299. 1813. 

BarBuDA: Barbuda Lagoon, coll. Cowan, Smiths.-Bredin no. 73. Nevis: 
Charlestown, coll. Schmitt, Smiths.-Bredin no. 67-58c. ANTIauUA: Reeds Beach, 
coll. Denckla, Smiths.-Bredin no. 119-59i. Sr. Lucia: Vigie Point, coll. Proc- 
tor no. A-2809; Pigeon Island, coll. Cowan, Smiths.-Bredin no. 50. Range from 
Bermuda and Florida to Las Aves Island, and at Trinidade Island, off Brazil. 


Penicillus dumetosus (Lamour.) Blainv. Man. Actin. Zoophyt. 553. 1834. 

Sr. Kirrs: Conaree Beach, coll. Proctor no. A-3124. Sr. Lucra: Pigeon 
Island, coll. Cowan, Smiths.-Bredin no. 45. Grenapa: Grenada Bay, coll. 
Proctor no. A-2714. Range from Bermuda and Florida to Colombia. 


Halimeda discoidea Dec. Ann. Sci. Nat., Bot. 18: 102. 1842. 

Sr, Lucia: Vigie Point, coll. Proctor A-2805; Pigeon Island, coll. Cowan, 
Smiths.-Bredin no. 44. A slight eastward extension of range in the West Indies. 
Range from Florida to Brazil. 


Halimeda incrassata (Ellis) Lamour. Hist. Polyp. Corall. Flex. 307. 1816. 
Barsupa: Cocoa Point, coll. Cowan, Smiths.-Bredin no. 87; Barbuda Lagoon 
coll. Cowan, Smiths.-Bredin no. 71. Nxvis: Charlestown, coll. Schmitt, Smiths.- 
Bredin no. 67-58b. St. Lucia: Marigot Bay, coll. Schmitt, Smiths.-Bredin no. 
43. Range from Bermuda and Florida to Grenada and the Netherlands Antilles. 


Halimeda opuntia (L.) Lamour. Hist. Polyp. Corall. Flex. 308. 1816. 
Barsupa: Cocoa Point, coll. Cowan, Smiths.-Bredin no. 82. Sr. Kirvs: 
Conaree Beach, coll. Proctor no. A-3144. Nevis: Red Cliff, coll. Proctor no. 
A-3221. Grenapba: True Blue, coll. Proctor no. A-2703. Range from Florida 
to Brazil. 
Codium intertextum Coll. & Herv. Proc. Amer. Acad. Arts Sci. 53: 54. 1917. 
St. BARTHELEMY: Anse de Cul-de-Sac, coll. Le Gallo no. 62, Sept. 15,1955. Sr. 
Kirts: Conaree Beach, coll. Proctor no. A~3141. Sr. Lucia: Pigeon Island, 
coll. Cowan, Smiths.-Bredin no. 66. Range from Bermuda and Florida to Brazil. 


Codium isthmocladum Vickers, Ann. Sci. Nat., Bot., IX, 1: 57. 1905. 
St. Lucia: Pigeon Island, coll. Cowan, Smiths.-Bredin no. 65. Range from 
Bermuda and North Carolina to Brazil. 


Xanthophyceae 


Vaucheriaceae 


Vaucheria dichotoma (L.) C. Ag. Synop. Alg. Scand. 47. 1817. 
Grenapva: True Blue, coll. Proctor no. A-2090. Range from Bermuda to 
Grenada, but reported from only a few stations. 


Phaeophyceae 


Ectocarpaceae 


Ectocarpus breviarticulatus J. Ag. Ofvers. K. Vetensk.-Akad. Férhandl. 4(1): 7. 
1847. 
Sr. Kirrs: Conaree Beach, coll. Proctor no. A-3131. Sr. Lucta: Marquis 
Bay, coll. Proctor no. A-2785; Vigie Point, coll. Proctor no. A-2822. GRENADA: 
Grenada Bay, coll. Proctor no. A-2729. Range from Jamaica to Brazil. 


597062—62——__2 


50 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Giffordia mitchellae (Harv.) Hamel, Bot. Notiser, 1939: 66. 1939. 

Sr. Lucia: Vigie Point, coll. Proctor no. A-2820. The easternmost record of 
this species in the West Indies to date. Ranging from Bermuda and North 
Carolina to Guadeloupe, and reported from Brazil. 


Dictyotaceae 


Dilophus guineensis (Kitz.) J. Ag. Lunds Univ. Arsskr., Afd. Math. Natur- 
vetensk. 17(4): 106. 1882. 

St. Krrrs: Conaree Beach, coll. Proctor nos. A-3127, A-3129; North Friars 
Bay, coll. Proctor no. A-3186. Nuzvis: Near Red Cliff, coll. Proctor nos. A-3212, 
A-3214. Sr. Lucia: Pigeon Island, coll. Cowan, Smiths.-Bredin no. 54; Vigie 
Point, coll. Proctor no. A-2816. Range from Bermuda and Florida to Venezuela. 


Dictyota bartayresii Lamour. Nouv. Bull. Sci. Soc. Philom. 1: 331. 1809. 
Antigua: Reeds Beach, coll. Denckla, Smiths.-Bredin no. 119-59c. Range 
from Bermuda and Florida to Brazil. 


Dictyota cervicornis Kiitz. Tab. Phycol. 9: 11. 1859. 
Antigua: Reeds Beach, coll. Denckla, Smiths.-Bredin no. 119-59d. Range 
from Bermuda and North Carolina to Brazil. 


Dictyota ciliolata Kitz. Tab. Phycol. 9: 12. 1859. 
Sr. Krrrs: Conaree Beach, coll. Proctor nos. A-3122, A-3123, A-3125. Range 
from Bermuda and Florida to Brazil. 


Dictyota dentata Lamour. Nouv. Bull. Sci. Soc. Philom. 1: 331. 1809. 

Sr. Kirrs: Conaree Beach, coll. Proctor A-3133. St. Lucta: Pigeon Island, 
coll. Cowan, Smiths.-Bredin no. 57. Grenapa: Grenada Bay, coll. Proctor no. 
A-2716. Range from Bermuda and Florida to Brazil. 


Dictyota dichotoma (Huds.) Lamour. Nouv. Bull. Sci. Soc. Philom. 1: 331. 
1809. 
Nevis: Charlestown, coll. Schmitt, Smiths.-Bredin no. 67-58h. Range from 
Bermuda and North Carolina to Brazil. 


Dictyota jamaicensis Taylor, Mar. Alg. East. Trop. Subtrop. Coasts Amer. 630. 

1960. 

St. Kirrs: Frigate Bay, coll. Proctor no. A-3164. Sr. Lucra: Vigie Point, 

coll. Proctor no. A-2798. Hitherto known from Jamaica, Grenada, and 
Venezuela. 


Dictyopteris delicatula Lamour. Nouv. Bull. Sci. Soc. Philom. 1: 332. 1809. 

Sr. Kirrs: Conaree Beach, coll. Proctor no. A-3130 (in part). Sr. Lucia: 
Pigeon Island, coll. Cowan, Smiths.-Bredin nos. 44 (mixed with Halimeda dis- 
coidea), 55 (in part). Range from Bermuda and Florida to Brazil. 


Dictyopteris jamaicensis Taylor, Mar. Alg. East. Trop. Subtrop. Coasts Amer. 
631. 1960. 
GRENADA: Grenada Bay, coll. Proctor no. A-2739. Previously only known 
from Jamaica. 


Dictyopteris justii Lamour. Nouv. Bull. Sci. Soc. Philom. 1: 332. 1809. 
St. Lucra: Pigeon Island, coll. Cowan, Smiths.-Bredin no. 56. Range from 
Bermuda and Florida to Brazil. 


Pocockiella variegata (Lamour.) Papenf. Amer. Journ. Bot. 30: 467. 1943. 

Nevis: Charlestown, coll. Schmitt, Smiths.-Bredin no. 67-58g. Sr. Lucia, 
Pigeon Island, coll. Cowan. Smiths.-Bredin no. 53. Range from Bermuda and 
North Carolina to Brazil. 


TAYLOR—MARINE ALGAE FROM LESSER ANTILLES ol 


Stypopodium zonale (Lamour.) Papenf. Bot. Notiser, 1940: 205. 1940. 
Sr. Lucia: Pigeon Island, coll. Cowan, Smiths.-Bredin no. 64. Range from 
Bermuda and Florida to Brazil. 


Padina gymnospora (Kiitz.) Vickers, Ann. Sci. Nat., Bot., IX, 1: 58. 1905. 

Sr. Kirrs: Conaree Beach, coll. Proctor no. A-3147. Nevis: Near Red Cliff, 
coll. Proctor no. A-3216. Sr. Lucia: Vigie Point, coll. Proctor no. A-2812. 
GRENADA: Grenada Bay, coll. Proctor no. A-2711. Range from Bermuda and 
Florida to Brazil. 


Padina sanctae-crucis Bgrg. Dansk Bot. Ark, 2(2): 45. 1914. 
GrenapDa: Grenada Bay, coll. Proctor no. A-2712. Range from Bermuda 
and Florida to Brazil. 


Padina vickersiae Hoyt in Britt. & Millsp. Bahama Flora 595. 1920. 
Sr. Kirrs: Frigate Bay, coll. Proctor no. A-3156. Grenapa: True Blue, coll. 
Proctor no. A-2696. Range from Bermuda and North Carolina to Brazil. 


Punctariaceae 


Colpomenia sinuosa (Roth) Derb. & Sol. Comptes Rend. Séanc. Acad. Sci. 
Suppl. 1: 11. 1856. 
Nevis: Red Cliff, coll. Proctor no. A-3225. Range from Bermuda and North 
Carolina to Brazil. 


Chnoospora minima (Hering) Papenf. Journ. So. Afr. Bot. 22(2): 69. 1956. 
Sr. Lucta: Vigie Point, coll. Proctor no. A-2781. Tosaago: Man-of-War 
Bay, coll. Cowan, Smiths.-Bredin no. 27. Range from Redonda Island to Brazil. 


Sargassaceae 


Sargassum fluitans Bgrg. Dansk Bot. Ark. 2(2): 66. 1914. 

Nevis: Red Cliff, coll. Proctor no. A-3126. AntTiaua: Reeds Beach, coll. 
Denckla, Smiths.-Bredin no. 119-59j. Range from Bermuda and Florida to 
Panama. 


Sargassum natans (L.) J. Meyen, Arch. fir Naturgesch. [Wiegm.], 4(2): 185. 
1838. 

BarBuva: Cocoa Point, coll. Cowan, Smiths.-Bredin nos. 88, 93. NeEvis: 
Red Cliff, coll. Proctor no. A-3125. AntTiaua: Reeds Beach, coll. Denckla, 
Smiths.-Bredin no. 119-59k. Range from Bermuda and North Carolina to 
British Honduras. 


Sargassum platycarpum Mont. Ann. Sci. Nat., Bot. IT, 18: 248. 1842. 

Sr. Kirts: Conaree Beach, coll. Proctor no. A-3151. Range from Bermuda to 
Brazil. 
Sargassum polyceratium Mont. Ann. Sci. Nat., Bot., II, 8: 356. 1837. 

Sr. Kirts: Conaree Beach, coll. Proctor no. A-3149. Range from Bermuda 
and Florida to Brazil. 


Sargassum rigidulum Kiitz. Spec. Alg. 615. 1849. 
Sr. Kirrs: Black Rocks, coll. Proctor no. A-3200. Range from Bermuda to 


Brazil. 


Sargassum vulgare C. Ag. Spec. Alg. Rite Cogn. 1: 3. 1820. 

Sr. Lucra: Pigeon Island, coll. Cowan, Smiths.-Bredin no. 59; Vigie Point, 
coll. Proctor no. A-2797. Grenapa: Grenada Bay, coll. Proctor no. A-2735. 
Var. foliosissium (Lamour.) J. Ag.: British Virain Isuanps: Guano Island, 
coll. Schmitt, Smiths.-Bredin no. 3. Range from Bermuda and Florida to Brazil. 


52 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Turbinaria tricostata Barton, Trans. Linn. Soc., Bot. II, 3: 218. 1891. 
Barsupa: Cocoa Point, coll. Cowan, Smiths.-Bredin no. 94. Range from 
Bermuda to the Netherlands Antilles. 


Turbinaria turbinata (L.) Kuntze, Rev. Gen. Plant. 3: 434. 1898. 
Sr. Krrrs: Conaree Beach, coll. Proctor no. A-3142. Range from Florida to 
Brazil. 


Rhodophyceae 


Helminthocladiaceae 


Liagora ceranoides Lamour. Hist. Polyp. Corall. Flex. 239. 1816. 
Sr. Kitts: North Friars Bay, coll. Proctor no. A-3185. Range from Bermuda 
and Florida to Brazil. 


Liagora farinosa Lamour. Hist. Polyp. Corall. Flex. 240. 1816. 
Sr. Kirts: Frigate Bay, coll. Proctor no. A-3159. Range from Bermuda and 
Florida to Brazil. 


Liagora valida Harv. Smithson. Contr. Knowl. 5(5): 138. 1853. 
Sr. Kirts: South Friars Bay, coll. Proctor no. A-3195 (young and soft). 
Range from Bermuda and Florida to Panama. 


Chaetangiaceae 


Galaxaura cylindrica (Ell. & Sol.) Lamour. Expos. Méthod. Genres Ordre 
Polypiers 22. 1821. 
Sr. Kitts: Conaree Beach, coll. Proctor no. A-3139. Range from Bermuda 
and Florida to Brazil. 


Galaxaura oblongata (Ell. & Sol.) Lamour. Hist. Polyp. Corall. Flex. 262. 1816. 
Sr. Lucra: Vigie Point, coll. Proctor no. A-2799. Range from Bermuda and 
Florida to Brazil. 


Galaxaura obtusata (Ell. & Sol.) Lamour. Hist. Polyp. Corall. Flex. 262. 1816. 
Nevis: Near Red Cliff, coll. Proctor no. A-3224. Range from Bermuda and 
Florida to Brazil. 


Galaxaura rugosa (Ell. & Sol.) Lamour. Hist. Polyp. Corall. Flex. 263. 1816. 

St. Kitts: Conaree Beach, coll. Proctor no. A-3138; Frigate Bay, coll. Proctor 
no. A-3161. St. Lucia: Vigie Point, coll. Proctor no. A-2795. Range from 
Bermuda and Florida to Brazil. 


Galaxaura squalida Kjellm. K. Svenska Vetensk.-Akad. Handl. 33(1): 55. 
1900. 

Str. Kirrs: Frigate Bay, coll. Proctor no. A-3170. Sr. Lucra: Pigeon Island, 

coll. Cowan, Smiths.-Bredin no. 61. Range from Bermuda and Florida to 
Venezuela, 


Galaxaura subverticillata Kjellm. K. Svenska Vetensk.-Akad. Handl. 33(1): 48. 
1900. 
Sr. Lucta: Pigeon Island, coll. Cowan, Smiths.-Bredin no. 62. Range from 
Bermuda and Florida to British Honduras. 


Gelidiaceae 


Gelidiella acerosa (Forssk.) Feldm. & Hamel, Rev. Gén. Bot. 46: 533. 1934. 

St. Lucra, Pigeon Island, coll. Cowan, Smiths.-Bredin no. 69. Sr. Lucia: 
Marquis Bay: coll. Proctor no. A-2774; Vigie Point, coll. Proctor no. A-2793. 
GrenapDa: Westerhall Point, coll. Proctor no. A-2683. Range from Bermuda 
and Florida to Brazil. 


TAYLOR—MARINE ALGAE FROM LESSER ANTILLES 53 


Rhizophyllidaceae 


Ochtodes secundiramea (Mont.) Howe in Britt. & Millsp. Bahama Fl. 583. 
1920. 
Sr. Kirrs: Black Rocks, coll. Proctor no. A-3189. Sr. Lucra: Vigie Point, 
coll. Proctor no. A-2825. Range from the Bahamas to Trinidad. 


Corallinaceae 


Fosliella farinosa (Lamour.) Howe zn Britt. & Millsp. Bahama Fl. 587. 1920. 

Sr. Kirrs: Conaree Beach, coll. Proctor no. A-3127. Antigua: Reeds Beach, 
coll. Denckla, Smiths.-Bredin no. 119-59p. Range from Bermuda and North 
Carolina to Brazil. 


Fosliella lejolisii (Rosan.) Howe in Britt. & Millsp. Bahama Fl. 588. 1920. 
Nevis: Charlestown, coll. Schmitt, Smiths.-Bredin no. 67-58i. Range from 
Prince Edward Island to Brazil. 


Ampbhiroa fragilissima (L.) Lamour. Hist. Polyp. Corall. Flex. 298. 1816. 

ANEGADA IsLAND: Pomato Point, coll. Schmitt, Smiths.-Bredin no. 42-58d 
(lesser part). Sr. Lucrta: Pigeon Island, coll. Smiths.-Bredin no. 68 (lesser part). 
Antiaua: Reeds Beach, coll. Denckla, Smiths.-Bredin no. 119-591. Range from 
Bermuda and North Carolina to Brazil. 


Amphiroa hancockii Taylor, Allan Hancock Atlantic Exped. 2: 95. 1942, 
Sr. Lucia: Pigeon Island, coll. Cowan, Smiths.-Bredin no. 63. Hitherto only 
known from Hispaniola and Panama, 


Amphiroa rigida Lamour. var. antillana Bgrg. Dansk Bot. Ark. 3(1c): 183. 
1917. 
Sr. Lucta: Pigeon Island, coll. Cowan, Smiths.-Bredin nos. 67, 68. Range 
from Florida to Panama. 


Corallina officinalis Linn., Fauna Suecica, edit. alt. 539. 1761. 

Sr. Lucta: Marquis Bay, coll. Proctor A-2770. A very interesting find, 
because, although this plant occurs further south as well as north, reports of it 
from the West Indies are extremely untrustworthy. The material is quite typical. 
Range south to North Carolina, and again from Brazil and Uruguay southward, 
with this as a new intermediate station. 


Jania adhaerens Lamour. Hist. Polyp. Corall. Flex. 270. 1816. 

ANEGADA IsLAND: Pomato Point, coll. Schmitt, Smiths.-Bredin no. 42-58d 
(lesser part). Barsupa: Barbuda Lagoon, coll. Cowan, Smiths.-Bredin no. 81 
(lesser part). Sr. Lucia: Pigeon Island, coll. Cowan, Smiths.-Bredin no. 68. 
The last is probably a new eastern record for the West Indies. 


Jania capillacea Harv. Smithson. Contr. Knowl. 5(5): 84. 1853. 
8t. Kirrs: Black Rocks, coll. Proctor no. A-3203. Sr. Lucia: Vigie Point, 
coll. Proctor nos. A-2804, A—2810 (in small part). Range from Florida to Brazil. 


Jania rubens (L.) Lamour. Nouv. Bull. Sci. Soc. Philom. 3: 186. 1812. 
Angea@apDA Istanp Pomato Point, coll. Schmitt, Smiths.-Bredin no. 42—58e (in 
part). Barsupa: Barbuda Lagoon, coll. Cowan, Smiths.-Bredin no. 79. 
GreNaApDA: Grenada Bay, coll. Proctor no. A-2730. Range from Bermuda and 
North Carolina to Uruguay. 
Grateloupiaceae 


Halymenia floresia (Clem.) C. Ag. Spec. Alg. Rite Cogn. 1: 209. 1822. 
Sr. Lucia: Vigie Point, coll. Proctor no. A-2801. Range from Bermuda and 
North Carolina to Brazil. 


54 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Grateloupia cuneifolia J. Ag. Ofvers. K. Vetensk.-Akad, Férhandl. 6: 85. 1849. 

Sr. Kirrs: Black Rocks, coll. Proctor nos. A-3145, A-3208. Sr. Lucia: 
Marquis Bay, coll. Proctor no. A-2771. A slight eastward extension of range in 
the West Indies. Range from Jamaica to Uruguay. 


Grateloupia dichotoma J. Ag. Algae Maris Medit. Adriat. 103. 1842. 
Sr, Kirrs: Black Rocks, coll. Proctor no. A-3192. Range from Jamaica to 
Trinidad. 


Grateloupia filicina (Wulf.) C. Ag. Spec. Alg. Rite Cogn. 1: 223. 1822. 

Barsupa: Barbuda Lagoon, coll. Cowan, Smiths.-Bredin no. 92. Sr. Krrrs: 
South Friars Bay, coll. Proctor no. A~3172; Frigate Bay, coll. Proctor no. A-3155, 
A-3157. Range from North Carolina to Brazil. 


Cryptonemia luxurians (Mert.) J. Ag. Spec. Gen. Ord. Alg. 2: 228. 1851. 
Grenapa: Grenada Bay, coll. Proctor no. A-2721. Range from Bermuda 
and Florida to Brazil. 


Gracilariaceae 


Gracilaria curtissiae J. Ag. Lunds Univ. Arsskr. Afd. Math. Naturvetensk. 
21(8): 61. 1886. 
Grenapa: Grenada Bay, coll. Proctor no. A-2722. 


Gracilaria debilis (Forssk.) Bgrg. Dansk Bot. Ark. 8(2): 7. 1982. 

Sr, Krrrs: Conaree Beach, coll. Proctor nos. A-3116, A-3118a, b, A-3143; 
North Friars Bay, coll. Proctor no. A-3181. Nevis: Red Cliff, coll. Proctor no. 
A-3222. Grenapa: Grenada Bay, coll. Proctor no. A-2237. Range from 
Bermuda and Florida to Brazil. 


Gracilaria domingensis Sonder, herb., Sphaerococcus domingensis Kiitz. Tab- 
Phycol. 19: 8. 1869; G. domingensis Sonder ex Collins, Proc. Amer. Acad. 
Arts Sci. 37: 254. 1901. 

Sr. Krrrs: Conaree Beach, coll. Proctor no. A-3132. Grenapa: Grenada 
Bay, coll. Proctor no. A-2723. Range from Jamaica to Brazil. 

Gracilaria ferox J. Ag. Spec. Gen. Ord. Alg. 2(2): 592. 1852. 

Sr, Krrrs: Conaree Beach, coll. Proctor no. A-3150; North Friars Bay, coll. 
Proctor no. A-3179; Frigate Bay, coll. Proctor no. A-3171. Sr. Lucia: Marquis 
Bay, coll. Proctor no. A-2786. Range from Bermuda and Florida to Brazil. 
Gracilaria foliifera (Forssk.) Bgrg. Dansk Bot. Ark. 8(2): 7. 1932. 

Grenapba: True Blue, coll. Proctor nos. A-2698, A-2707. ToxBaco: Man-of- 
War Bay, coll. Cowan, Smiths.-Bredin no. 31. Range from Bermuda and North 
Carolina to Uruguay. 

Gracilaria verrucosa (Huds.) Papenf. Hydrobiol. 2: 195. 1950. 

St. Kirrs: Conaree Beach, coll. Proctor no. A-3117. Nevis: Charlestown, 
coll. Schmitt, Smiths.-Bredin no. 67-58k. Topaco: Man-of-War Bay, coll. 
Cowan, Smiths.-Bredin no. 28, Range from Bermuda and North Carolina to 
Brazil. 

Solieriaceae 


Sarcodiotheca caribaea nov. sp. Puate 1, Figures 1, 2; Puatre 4, Figure 1 


Plantae omnes infra breviter teretes supra ad laminan 4-8 irregu- 
lariter dichotome furcatam, ramis ligulatis 4-10 mm. lat. super 
furcationes inferiores, 1-2 mm. in divisionibus supremis, marginibus 
integris, expansae; tetrasporangia super thallam dispersa, 30-38 4 


TAYLOR—MARINE ALGAE FROM LESSER ANTILLES 55 


diam., 45-55 yw longa; cystocarpi prominentes, marginales, intus 
parenchymatici; pericarpi ostiolati, intus filamentosi, nonnullis filis 
inter gonimoblastos ad centrum sterile extensis. 

Plants arising from very small, lobed, cushion-shaped holdfasts, 
briefly terete below and often with 1-3 proliferations from the hold- 
fast or the stalk, each expanding above to a 4-8 times irregularly 
dichotomous blade with a tapering base, the strap-shaped branches 
4-10 mm. broad above the lower forkings, 1-2 mm. in the uppermost 
divisions, the tips rather acute, the margins entire. Blade thickness 
to 400-500 u, the cortex of one layer of small cells and the outer 
medulla of two layers, the inner ones rather large; medullary cavity 
small, loosely crossed by oblique thicker-walled filaments about 
18-25 » in diameter, and thinner-walled ones about 4.5-14 ». Tetra- 
sporangia scattered over the thallus, extending inward from the 
cortex, zonate, 30-38 uw in diameter, 45-55 » long. Spermatangia 
widely distributed in patches over the surface. Cystocarps promi- 
nent, marginal, with a parenchymatous core, the pericarps ostiolate, 
filamentous within, with some strands extending between the gonimo- 
blasts to the sterile center; carposporangia produced in short centri- 
petal series. 

Type in the U.S. National Herbarium, collected in Man-of-War Bay, Tobago, 
British West Indies, April 7, 1959, by Richard 8. Cowan (No. 41). 

These plants agree in most respects with the description of Sarco- 
diotheca, the only particular discrepancy lying in the restriction of 
the cystocarps to the margin, where they are strongly elevated. 
Individuals of west-coast species of the genus may, on occasion, have 
some of them along the margin. Since this is the first record of the 
genus in the western Atlantic in a family not represented there by 
many species, it is of particular interest. 

Agardhiella tenera (J. Ag.) Schmitz, Flora 72: 441. 1889. 

Tospaco: Man-of-War Bay, coll. Cowan, Smiths.-Bredin no. 38. Range from 
North Carolina to Brazil. 

Eucheuma isiforme (C. Ag.) J. Ag. Ofvers. K. Vetensk.-Akad. Forhandl. 4: 16. 


1847. 
Barsupa: Cocoa Point, coll. Cowan, Smiths.-Bredin no. 95; Gravenor Land- 


ing, coll. Cowan, Smiths.-Bredin no. 26. This records a slight eastward extension 
of range. Range hitherto Bermuda and Florida to St. Barthélemy. 


Meristotheca tobagensis nov. sp. PLATE 2, Fiaures 1, 2 

Plantae foliosac, in margine irregulariter furcatae; plantae sperma- 
tangiales identidem irregulariter ad subdichotome furcatae, in partibus 
inferioribus ad 1.0 cm. lat., modice ad valde eroso-dentatae; plantae 
cystocarpicae lobos relative latiores, simpliciores magis irregulares, 
sursum valde aculeato-dentatae; cystocarpi 0.5-2.5 mm. diam., super- 
ficiales aut clevati super superficiem dispersi. 


56 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Plants foliose, to at least 6 cm. in height, the fleshy blades reaching 
a thickness of 530 yw, marginally irregularly forked, but probably 
sexually dimorphic. Structurally showing an outer cortex of one 
layer of oval cells about 5 yw in diameter, 15 » tall, below a firm gelat- 
inous cuticle, and resting on a layer of small rounded ones. Outer 
medulla of about 3 layers of thick-walled cells, the inner ones being 
very large, enclosing a filamentous central medulla, this being rather 
loose in the younger parts, dense near the base. Spermatangial plants 
several times irregularly to subdichotomously forked, to 1.0 cm. broad 
in the lower parts, or 1.5 cm. where subcuneate below a fork, the 
ultimate divisions more band-shaped and tapering, moderately to 
strongly erose-dentate. Cystocarpic plants with relatively broader, 
simpler and more irregular lobes rather than forking blades, these 
very strongly aculeate-dentate above and, scattered over the surface, 
bearing conspicuous superficial or elevated cystocarps 0.5-2.5 mm. in 
diameter, each with a thick ostiolate pericarp, this parenchymatous 
without but with a filamentous lining. Gonimoblasts surrounding a 
parenchymatous ‘core, this externally interrupted by groups of fila- 
ments spreading over the surface and bearing the small carpospor- 
angia. Tetrasporangial plants not seen. 

Type in the U.S. National Herbarium, collected in Man-of-War Bay, Tobago, 
British West Indies, April 7, 1959, by Richard 8. Cowan (no. 40, spermatangial 
and cystocarpic plants). 

In the absence of tetrasporangial material and with too little 
cystocarpic material to permit a developmental study, the generic 
assignment of these plants must be considered tentative. They are 
far smaller and less broadly foliar than M. florida Kylin, an ill-known 
plant although reported from Florida to Panama, and more regularly 
branched than the type species, M. papulosa (Mont.) J. Ag., which is 
not known to occur in the West Indies. 


Hypneaceae 


Hypnea musciformis (Wulf.) Lamour. Mém. Muséum [Paris], 20: 43. 1813. 

St. Kirrs: Black Rocks, coll. Proctor no. A-3194; Conaree Beach, coll. 
Proctor no. A-3120; South Friars Bay, coll. Proctor no. A-3174; Frigate Bay, 
coll. Proctor nos. A-3158, A-3162. Sr. Lucia: Vigie Point, coll. Proctor no. 
A-2807. Range from Bermuda and North Carolina to Brazil. 


Hypnea spinella (C. Ag.) Kitz. Tab. Phycol. 18: 9. 1868. 
Sr. Kirrs: Frigate Bay, coll. Proctor no. A-3168. Sr. Lucra: Vigie Point, 
coll. Proctor no, A~2821. Range from Bermuda to Brazil. 


Phyllophoraceae 


Gymnogongrus tenuis (J. Ag.) J. Ag. Ofvers. K. Vetensk.-Akad. Férhandl. 6: 88. 
1849. 
Sr. Kirts: Frigate Bay, coll. Proctor no. A-3160. Reported from North 
Carolina, Guadeloupe, Tobago, Brazil, and Uruguay. 


TAYLOR—-MARINE ALGAE FROM LESSER ANTILLES 57 


Champiaceae 


Coelothrix irregularis (Harv.) Bgrg. Dansk Bot. Ark. 3: 389. 1920. 
Sr. Lucia: Vigie Point, coll. Proctor no. A-2824, Range from Bermuda and 
Florida to British Honduras. 
Ceramiaceae 


Ceramium nitens (C. Ag.) J. Ag. Spec. Gen. Ord, Alg. 2: 130. 1851. 

Sr. Lucta: Vigie Point, coll. Proctor no, A~2823. Within the West Indies this 
is a slight eastward extension of range. Hitherto known from Bermuda and 
Florida to Martinique and along the mainland to Colombia. 


Ceramium subtile J. Ag. Spec. Gen. Ord. Alg. 2: 120. 1851. 
GRENADA: True Blue, coll. Proctor no, A-2695. Range from Bermuda and 
Florida to Barbados and the Netherlands Antilles. 


Centroceras clavulatum (C. Ag.) Mont. in Durieu, Flore Algérie 1: 140. 1846. 

St. Kirrs: Black Rocks, coll. Proctor no. A~8207. Sr. Lucta: Marquis Bay, 
coll. Proctor no. A-2787. Grenapa: True Blue, coll. Proctor no. A—2693. 
Range from Bermuda and Florida to Brazil. 


Spyridia aculeata (Schimp.) Kitz. Phye. Gen. 377. 1843. 

GRENADA: Grenada Bay, coll. Proctor no. A-2733. Tospaco: Man-of-War 
Bay, coll. Cowan, Smiths.-Bredin no. 33. Range from Bermuda and North 
Carolina to Brazil. 


Spyridia filamentosa (Wulf.) Harv. in Smith, English Fl. 5(1): 337. 1833. 

Sr. Lucta: Marquis Bay, coll. Proctor A-2767, Grenapa: Westerhall Point, 
coll. Proctor nos, A-2686, A-2688. Range from Bermuda and North Carolina to 
Brazil. 

Delesseriaceae 


Grinnellia americana (C. Ag.) Harvey var. caribaea Taylor, Hancock Atlantic 
Exped. 2: 127. 1942. 
TosaGco: Man-of-War Bay, coll. Cowan, Smiths.-Bredin no. 30. The species 
itself chiefly northern in range, but with the variety known from North and South 
Carolina, the Netherlands Antilles and Venezuela as well. 


Nitophyllum punctatum (Stack.) Grev. Alg. Britt. 79. 1830. 

Topaco: Man-of-War Bay, coll. Cowan, Smiths.-Bredin no. 29. Conspicu- 
ously large and handsome cystocarpic plants. This is the first well-substantiated 
record for the West Indies. 

Dasyaceae 


Heterosiphonia gibbesii (Harv.) Falk. Fauna et Flora Golf. Neapel 26: 653. 
1901, 

Sr. Kirrs: North Friars Bay, coll. Proctor no. A-3183. Grenapa: Grenada 

Bay, coll. Proctor no, A~2725. Range from Bermuda and Florida to Venezuela. 


Rhodomelaceae 


Falkenbergia hillebrandii (Bornet) Falk. Fauna et Flora Golf. Neapel 26: 689. 
1901. 
Antiava: Reeds Beach, coll. Denckla, Smiths.-Bredin no. 119-590. Range 
from Bermuda and Florida to Brazil. 


Polysiphonia ferulacea Suhr in J. Ag. Spec. Gen. Ord. Alg. 2: 980. 1863. 

St. Kirrs: Conaree Beach, coll. Proctor no. A-3119; South Friars Bay, coll. 
Proctor no, A-3177.. Grenapa: Grenada Bay, coll. Proctor no, A-2719. Range 
from Bermuda and Florida to Venezuela. 


597062— 62——__3 


58 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Polysiphonia subtilissima Mont. Ann. Sci. Nat, Bot. II, 13: 199. 1840. 

Sr. Krrrs: Conaree Beach, coll. Proctor no. A-3152. Tospaco: Man-of-War 
Bay, coll. Cowan, Smiths.-Bredin no. 37. Range from Bermuda and North 
Carolina to Brazil. 


Bryothamnion triquetrum (Gmel.) Howe, Journ. N.Y. Bot. Gard. 16: 222. 
1915. 

ANEGADA IsLAND: Pomato Point, coll, Schmitt, Smiths.-Bredin no, 42-58f. 
Sr. Kirrs: Conaree Beach, coll. Proctor no. A-3130. Nevis: Red Cliff, coll. 
Proctor no. A-3211. Grenapa: Grenada Bay, coll. Proctor no. A-2727. Range 
from Florida to Brazil. 


Digenia simplex (Wulf.) C. Ag. Spec. Alg. Rite Cogn. 1: 388. 1822. 

Barsupa: Barbuda Lagoon, coll. Cowan, Smiths.-Bredin no. 81. Sr. Kirrs: 
Black Rocks, coll. Proctor no, A-8202. Antigua: Reeds Beach, coll. Denckla, 
Smiths.-Bredin no. 119-59h. Sr. Lucra: Vigie Point, coll. Proctor no. A-2810. 
Grenapa: Grenada Bay, coll. Proctor no. A-2726. Range from Bermuda and 
Florida to Brazil. 


Murrayella periclados (C. Ag.) Schmitz, Ber. Deutsche Bot. Ges. 11: 227. 1893. 
Grenapa: True Blue, coll. Proctor no. A-2692; Westerhall Point, coll. Proctor 
no. A-2687. Range from Bermuda and Florida to French Guiana. 


Bostrychia tenella (Vahl) J. Ag. Spec. Gen. Ord. Alg. 2: 869. 1863. 
Grenapa: Westerhall Point, coll, Proctor no, A-2684, Range from Bermuda 
and Florida to Brazil. 


Dipterosiphonia dendritica (C. Ag.) Schmitz in Engler & Prantl, Nat. Pflanzenf, 
1(2): 463. 1897. 

Sr. Lucra: Marquis Bay, coll. Proctor no, A~-2769. On Corallina officinalis, 
so crowded and mosslike that the typical branch arrangement was seldom seen. 
This is the easternmost and only the second record in the West Indies, it having 
been reported from Beata Island by Bgrgesen. 


Amansia multifida Lamour. Nouv. Bull. Sci. Soc. Philom. 1: 332. 1809, 
Nevis: Near Red Cliff, coll. Proctor no. A-3218. Range from Florida to 
Brazil. 


Vidalia obtusiloba (Mert.) J. Ag. Spec. Gen. Ord. Alg., 2(3): 1128. 1863. 
Grenapva: Grenada Bay, coll. Proctor no. A-2724. Range from Florida to 
Brazil. 


Chondria atropurpurea Harv. Smiths. Contr, Knowl. 5(5): 22. 1853. 


Grenapa: Grenada Bay, coll. Proctor no. A-2732. The easternmost record 
within the West Indies. Range from Bermuda and North Carolina to Brazil. 


Chondria littoralis Harv. Smiths. Contr. Knowl. 5(5): 23. 1853. 
Sr. Krrrs: Conaree Beach, coll. Proctor no. A-3154, an exceptionally attenuate 
form. Range from Bermuda and North Carolina to Colombia. 


Chondria tenuissima (G. & W.) C. Ag. Synop. Alg. Scand. xviii. 1817. 

Topaco: Man-of-War Bay, coll. Cowan, Smiths.-Bredin no. 32. The eastern- 
most record within the West Indies. Range hitherto from North Carolina to 
Guadeloupe and Venezuela. 


Waldoia nov. gen. 


Laminae planae alterne marginaliter conformiter ramosae, ecostatae, 
cellulis axialibus non perspicuis per nonnulla fila rhizoidea circumdatis, 


TAYLOR—-MARINE ALGAE FROM LESSER ANTILLES 59 


cellulis pericentralibus non perspicuis; tetrasporangia in stichidiis 
simplicibus cylindricis super superficilem axium ramorumque ordi- 
nariorum dispositis; pericarpi sessiles aut breviter stipitati, in super- 
ficie axium ramorumque ordinarium dispositi. Species typicum: 
Waldoia antillana. 

Plants with flat blades, alternately marginally conformably branched, 
ecostate, the axial cells obscure. Tetrasporangia in simple cylindrical 
stichidia borne over the surfaces of ordinary axes and branches. Peri- 
carps sessile or briefly stalked, borne on the surfaces of ordinary axes 
and branches. 


Waldoia antillana nov. sp. PLATE 3, Ficures 1, 2; Prats 4, Figure 2 


Plantae ex haptero crasse fibroso parce ramoso, in laminis primariis 
lineari-lanceolatis ecostatis gradatim expansae, laminis per alti- 
tudinem plantae totam plerumque extensis, usque ad 4-7 mm. lat.; 
plantae bis vel ter abunde ramosae; tetrasporangia in stichidiis ad 
4-5 mm. longis, 250-400 u diam., subcylindrica; pericarp super ramos 
laterales atque versus extremitates axis primarii dispersi, rotundati, 
ca. 0.6 mm. diam., ostiolo paululum producto. 

Plants to 13 cm. high (perhaps more), rose-red to dull purplish, 
soft, closely adherent to paper except at the base, arising from a 
sparingly branched coarsely fibrous holdfast, this giving rise to a few 
erect, foliar axes; axes very briefly terete below, the stalklike portion 
2-15 mm. long, sometimes with a few terete or flattened proliferations, 
gradually expanded to the primary linear-lanceolate ecostate blades, 
these generally extending the full height of the plant and becoming 
4-7 mm. broad, the margins slightly undulate-crenate, the apex 
somewhat narrowed but the tip obtuse to retuse; growth from an 
evident apical cell somewhat projecting or sunken in the apical notch 
(if present), and surrounded by a few obsolescent trichoblasts. Fronds 
abundantly 2- or 3-times branched, the branches, marginal, alternate, 
the branch attachments subterete to flat but contracted, the bases 
obtusely tapered, the branches otherwise like the primary axis. 
Tetrasporangia in notable stichidial branchlets scattered over the 
faces of the blades, often in abundance, to 4-5 mm. long, 250-400 u 
diam., at first subconical, becoming subcylindrical with a slightly 
contracted base and briefly tapered tip, the sporangia formed pro- 
gressively near the growing point and soon discharged. Pericarps 
scattered over the lateral branches and toward the ends of the primary 
axes, superficial, sessile, or contracted below and briefly stalked, 
rounded, about 0.5 mm. in diameter, the ostiole slightly produced. 
Spermatangial plants not seen. 

Type in the U.S. National Herbarium, collected in Man-of-War Bay, Tobago, 
British West Indies, April 7, 1959, by Richard S. Cowan (no. 39, tetrasporangial 
plants). 


60 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


These plants represent a particularly strange and unexpected 
addition to the Caribbean flora, for the small and specialized genus 
Cladhymenia to which they at first seemed to belong is primarily a 
Pacific one. Once before the writer hesitantly assigned some sterile 
Colombian material to Cladhymenia, and naturally one must consider 
the possibility that the same species is represented in both collections. 
There is little about the histological characters to distinguish these 
plants: The cortex cells in surface view seem about one-third broader 
in the older parts of these Tobagan specimens. It is even more 
difficult to recognize the primary axial cell and pericentral cells than 
in C. lanceifolia. What probably is the primary axis seems to be 
surrounded by a few irregular small elements rhizoidal in nature, and 
the large cells immediately outside these are not regular in number or 
position. The Tobagan plants are also far more elegantly branched, 
with longer and more symmetrical axes than the plants from Colombia, 
which do not adhere to paper except in the youngest proliferations. 
The possibility remains that the Colombian specimens are fragmentary 
and proliferous basal portions of old plants. However, the aspects of 
the two are so very different, as may be seen by comparing the figures 
of C. lanceifolia (Taylor 1942, pl. 19) with those given here of Waldova 
antillana, that it seems safer to describe the new material as a new 
species rather than to risk confusing the two. 

The generic assignment becomes critical now that fertile material 
is at hand. Until fertile material directly connected with the Colom- 
bian plants comes to hand nothing can be done about the status of 
C. lanceifolia, which, on the basis of sterile material, cannot be 
separated from Cladhymenia. Contrariwise, the Tobagan material 
with the tetrasporangia in simple stichidia all over the surfaces of 
the blades and not as marginal structures, and with the pericarps 
sessile or briefly stalked and similarly scattered over the surfaces 
rather than in marginal projections, is in its reproductive organs 
very different indeed, although belonging to the same section of the 
family. It seems appropriate, therefore, to make Tobagan material 
the type of a new genus Wa/doia, named in appreciation of the help 
given by Dr. Waldo Schmitt, and to leave the transfer of the Co- 
lombian species name until evidence to justify it appears.? 


Acanthophora muscoides (L.) Bory, Voy. de la Coquille, Bot. Crypt. 156, 
1828. 
Sr. Kirts: Frigate Bay, coll. Proctor no. A-3163. Range from Florida to 
Brazil, a less common species than the next. 


2 The writer wishes to express appreciation of the kindness of Professor G. F. Papenfuss in examining 
representative Tobagan specimens and offering valuable suggestions which in general confirmed the writer’s 
own observations. 


TAYLOR—MARINE ALGAE FROM LESSER ANTILLES 61 


Acanthophora spicifera (Vahl) Bgrg. Bot. Tidsskr. 30: 201. 1910. 

Barsupa: Cocoa Point, coll. Cowan, Smiths.-Bredin, no. 89. Sr. Krrts: 
Black Rocks, coll. Proctor no. A~3197, St. Lucia: Pigeon Island, coll. Cowan, 
Smiths.-Bredin no. 60; Vigie Point, coll. Proctor no. A-2811. GRENADA: 
Westerhall Point, coll. Proctor no. A-2685. Range from Bermuda and Florida 
to Brazil. 


Laurencia gemmifera Harv. Smiths. Contr. Knowl. 5(5): 73. 1853. 

Bargupa: Barbuda Lagoon, coll. Cowan, Smiths.-Bredin no. 77. NeEvis: 
Charlestown, coll. Schmitt, Smiths.-Bredin no. 67-58j. Range from Bermuda 
and North Carolina to Brazil. 


Laurencia obtusa (Huds.) Lamour. Mém. du Museum [Paris] 20: 42. 1813. 

VIRGIN IsLANDs: Guano Island, coll. Schmitt, Smiths.-Bredin no. 4. BAaRrBUDA: 
Barbuda Lagoon, coll. Cowan, Smiths.-Bredin no. 91. Range from Bermuda 
and Florida to Brazil. 


Laurencia papillosa (Forssk.) Grey. Alg. Britt. lii. 1830. 

ANEGADA IsLaNnD: Pomato Point, coll. Schmitt, Smiths.-Bredin no. 42-58d 
(major part). Sr. Kitrs: Black Rocks, coll. Proctor no. A-3196; Conaree Beach, 
coll, Proctor no. A-3128; South Friars Bay, coll. Proctor no. A-3178; Frigate 
Bay, coll. Proctor no. A-8167. Sr. Lucrta: Marquis Bay, coll. Proctor no. 
A-2773. Range from Bermuda and Florida to Brazil. 


Laurencia scoparia J. Ag. Spec. Gen. Ord. Alg. 2: 746. 1863. 

St. Kirrs: Black Rocks, coll. Proctor no. A-3187; Conaree Beach, coll. Proctor 
no, A-3128; North Friars Bay, coll. Proctor no. A-3182. Sr. Lucia: Marquis 
Bay, coll. Proctor no. A-2772; Vigie Point, coll. Proctor no. A-2803. Range 
from Jamaica to Brazil. 


Bibliography 
Bércusen, F. 
1913-1920. The marine algae of the Danish West Indies. I. Chlorophyceae 
Dansk Bot. Arkiv, 1(4) ; 1-158 + 2, 126 figs., map, 1913; II. Phaeophyceae’ 
2(2): 1-66 + 2, figs. 1-44, 1914; III. Rhodophyceae 3: 1-80, figs. 1-86; 
1915; 81-144, figs. 87-148, 1916; 145-240, figs. 149-230, 1917; 241-304, 
figs. 231-307, 1918; 305-368, figs. 308-360, 1919; 369-504, figs. 361-435, 
1920. 
Dicxiz, G. 
1874. On the marine algae of Barbados. Journ. Linn. Soc. (Bot.) 14: 
146-152, pl. 11. 1875. 
GRIEVE, 8. 
1909. Note on some sea-weeds from the Island of Dominica, British West 
Indies. Trans. Bot. Soc. Edinburgh, 24(1); 7-12. 
HAMEL, G. 
1929. Contribution 4 la flore algologique des Antilles. Ann. Crypt. 
Exotique, 2: 53-58, 9 figs. 
HameE., G. and Hamet-Joukov, A. 
1931. Algues des Antilles frangaises (Exsiccata). Fasc. 1-3. Paris. 
Howe, M. A. 
1928. Notes on some marine algae from Brazil and Barbados. Journ. 
Washington Acad. Sci. 18: 186-194, 2 figs. 


62 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Mazk, H., and ScurammM, A. 

1870-77. Essai de classification des algues de la Guadeloupe. (Designated 
the second edition, but actually the third edition of this work, originally 
issued with A. Schramm as the senior author.) xix + 283 + iii pp. 
Basse-Terre, Guadeloupe. 

Murray, G. 

1888, 1889. Catalogue of the marine algae of the West Indian region. 
Journ. Bot. Brit. and For. 26: 193-196, 1888; 27: 237-242, 257-262, 
298-305, 1889. London. 

QuESTEL, A. 

1951, Algae, pp. 193-208, in La Flore de la Guadeloupe. Géographie 
Générale de la Guadeloupe et Dépendences, 1m et 111. Géographie 
Biologique, 1.—La Flore. 327 pp., 8 pls., 117 figs., 2 maps. Basse- 
Terre, Guadeloupe. 

Scuramm, A., and Mazf, H. 

1865. Essai de classification des algues dela Guadeloupe. (1st ed., printed). 
ii + 52 pp. Basse-Terre, Guadeloupe. 

1866. Essai de classification des algues de la Guadeloupe. (2nd ed., litho- 
graphed), iii + 144 pp. Cayenne, French Guiana. 

Tayitor, WILLIAM RANDOLPH. 

1929. Notes on algae from the tropical Atlantic Ocean [I]. Amer. Journ. 
Bot. 16: 621-630, 13 figs., pl. 62. 

1940. Marine algae of the Smithsonian-Hartford Expedition to the West 
Indies, 1937. Contr. U.S. Nat. Herb. 28: 549-562, pl. 20. 

1942. Caribbean marine algae of the Allan Hancock Expedition, 1939. 
Rep. Allan Hancock Atlantic Exped. 2: 1-198, 20 pls. Los Angeles. 

1960. Marine Algae of the Eastern Tropical and Subtropical Coasts of the 
Americas. ix + 870 pp., 14 figs., 80 pl. University of Michigan Press, 
Ann Arbor. 

Vann, M. 

1802. Endeel kryptogamiske planter fra St. Croix. Skrifter af Natur- 

historie-Selskabet [Copenhagen], 5(2); 29-47. 
Vickers, A. 

1905. Liste des algues marines de la Barbade. Ann. Sci. Nat. (Bot.) IX, 
1: 45-66. 

+1908. Phycologia Barbadensis. Iconographie des algues marines ré- 
coltées 4 l’Ile Barbade (Antilles): (Chlorophycées et Phéophycées). 
Text by M. H. Shaw, i-ix. Part I, Chlorophyceae, pp. 1-30, 53 pls. + 
14h, '43b, 44b, 44c; II, Phaeophyceae, pp. 31-44, 34 pls. + 6b, 24b. 
Paris. 


U.S. GOVERNMENT PRINTING OFFICE: 1962 


UNITED STATES NATIONAL MUSEUM 


CoNTRIBUTIONS FROM THE UNITED STATES NATIONAL HERBARIUM 


Vo.LumE 36, Part 3 


THE LICHEN FAMILY GRAPHIDACEAE 
IN MEXICO 


By Micuae.t Wirts and 
Mason E. Hate, Jr. 


BULLETIN OF THE UNITED STATES NaTIONAL Museum 
SMITHSONIAN INSTITUTION «+ WASHINGTON, D.C. « 1963 


THE LICHEN FAMILY GRAPHIDACEAE 
IN MEXICO 


By Micwaet Wirt and 
Mason E. Hate, Jr. ! 


Introduction 


The Graphidaceae form a large but very poorly known family of 
corticolous, crustose lichens widely distributed in tropical regions. 
The senior author has long had an interest in studying this group, and 
during his tenure at the Smithsonian Institution as a research assist- 
ant, in 1960-61, was afforded the opportunity to carry these plans 
forward. It was decided that a revision of the Graphidaceae of 
Mexico would be a realistically delimited area of study. Mexico is 
known to have a rich tropical flora, and large representative collections 
are available for study, including some collections made by the junior 
author and T. R. Soderstrom in southern Mexico and the Pringle 
material from central Mexico. 

In the initial stages of the study it became obvious that type speci- 
mens of extra-Mexican species would have to be examined, especially 
in view of the extensive synonymy that was to be anticipated in a 
family which has never been treated monographically. To this end, 
types of many species described by Krempelhuber, Miiller-Argau, 
Nylander, and Vainio were examined. Altogether types of more 
than 100 taxa were critically investigated. Unfortunately few of the 
species described by Fée in 1824 were available, and in most cases 
the current concepts of these species had to be followed. Redinger’s 
(1933, 1935) excellent revisions of Brazilian Graphidaceae have been 
especially helpful in this respect. 

The Fée types are reportedly preserved at Rio de Janeiro, but in- 
quiries there failed to uncover more than a few species. It has been 
learned recently that a nearly complete series of isotypes was prepared 
by Miiller-Argau when he revised the Fée species in 1887. These 
specimens are still in the process of being curated and incorporated 


1 Based on research partially supported by a grant from the National Science Foundation, 
63 


64 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


into the general cryptogamic herbarium at Geneva. Because of their 
fundamental importance in the taxonomy of the Graphidaceae and 
other tropical crustose families, Fée’s many species must eventually 
be carefully typified, and it is hoped that this can be accomplished in 
the near future. 

The circumscription of the Graphidaceae is far from settled (cf. 
Santesson, 1952, pp. 44, 45), because this family once included a 
number of ascolocular genera (¢.g., Opegrapha). As here considered, 
the family is taken to include all those ascohymenial lichens which 
have lirelliform ascocarps and lack columellae. By this definition, 
all the ascohymenial representatives of the Chiodectonaceae should 
probably be referred to the Graphidaceae. ‘This transfer would actu- 
ally be in accord with natural relationships, as the so-called stromata 
of the Chiodectonaceae are not significantly different from the asco- 
carps of the Graphidaceae. It is not uncommon to find individuals 
with polymorphic ascocarps such that half of an individual can be 
referred to one family and half to another. Sarcographa (Chiodecto- 
naceae) and Phaeographis (Graphidaceae) are so closely allied as to 
make identification at the species level nearly impossible. The lack of 
consistency in the formation of stromatoid tissue renders the taxonomy 
of this group very unclear. 

Another major difficulty in delimiting the Graphidaceae becomes 
evident in considering the related family Thelotremataceae, which is 
usually characterized by having round ascocarps. Some species in 
the Thelotremataceae have lirelline ascocarps and show variations 
paralleling those of the Graphidaceae. In addition, some species in 
the Graphidaceae (e.g., Graphina confluens (Fée) Mill. Arg., G. hemi- 
sphaerica (Vain.) Zahlbr., and @. quassiaecola (Fée) Mill. Arg.) ex- 
hibit a definite trend towards the formation of round apothecia, 
structurally identical with those of many thelotremataceous species. 
A further link can be found in Graphina macrospora (Kremplh.) Mill. 
Arg., in the ascocarps of which are found columellae (cf. fig. 4), usually 
considered as being restricted to the Thelotremataceae. It is obvious 
that the shape of the ascocarps is not a valid criterion for separating 
families and perhaps not even useful for generic delimitation. Much 
more work on developmental anatomy is necessary before workable 
families can be erected in these lichens. 

The genera of the Graphidaceae are traditionally separated on the 
bases of algal host, spore color, and spore septation. As Santesson 
(1952, p. 46) has already pointed out, the genus of algal host is a poor 
criterion in lichen taxonomy. Both spore color and septation are 
characters of sometimes limited usefulness in generic delimitation. 
There are groups of species which have constantly colorless, thin- 
walled, four-celled spores. There is also a natural group of species 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 65 


with brown, thick-walled, five- to eight-celled spores. However, in 
other cases spore color and septation seem to be too variable to serve 
as major taxonomic features; they are often so unstable as to vary 
within a single specimen. See the discussions under Graphina acharii 
(p. 72) and Phaeographina chrysocarpa (p. 102) for a more detailed 
account of this variability. 

The genus Melaspilea, usually separated from Graphis by the cubic 
shape of the spore locules, has a perfect counterpart in the Graphis 
species allied to G@. turbulenta Tuck. The spores in both of these 
groups are almost always 2-locular and identical, and separating them 
into two genera is quite unrealistic. The spores of Melaspilea often 
show variation in color from hyaline to dark brown. 

Within the four major genera, Graphis, Phaeographis, Graphina, 
and Phaeographina, parallel subgeneric divisions into sections have 
been established. ‘These have been summarized, for the most part, 
by Redinger (1933, 1935), who recognized 50 of these taxa. The 
characters used in sectional analysis are discussed below. They have 
proved in our studies to be highly inconstant and intergrading, and a 
single species or occasionally a single individual may be placed in 
either of two sections with equal facility. 

1. Degree of excipular carbonization. This character is often useful 
on the specific level, but it varies considerably. It is difficult to divide 
the species objectively into major groups on the criteria of ‘‘exciple 
entirely black,” “exciple black above and colored below,” or “exciple 
completely brown or colorless.” Too often random carbonization 
will occur in portions of an exciple (see figs. 8, 10-13). In addition, 
it is not easy to delimit brown exciples from black exciples, since the 
pigmentation often intergrades. 

With regard to carbonization, it might be mentioned that it is neces- 
sary to cut sections of the ascocarps as thin as possible, since un- 
carbonized tissues will often appear black in thick sections. Included 
air will also cause the same appearance. We have found that the best 
results are obtained when Hoyer’s solution (Anderson 1954) is used 
as the mounting and clearing agent and the slides are allowed to stand 
on a warming plate for at least a half hour. This medium clears tissues 
quickly and completely, yet it does not change their relative densities. 
Moreover it is an excellent solution in which to demonstrate colored 
spores, as these do not decolorize at all, while the hymenium and asci 
become transparent and clear. 

2. Open or closed exciples. Exciples in the Graphidaceae cannot 
always be easily separated into two major types on the basis of whether 
they are closed basally or not (see figs. 31-35). Even in species with 
highly carbonized exciples it is often possible to find the basal portion 
turning brown and uncarbonized or becoming so thin as to be closed 


66 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


in one part of an ascocarp and open in another. It is difficult to 
determine where apothecial tissue ends and where thallus or sub- 
stratum begins in species with only partial carbonization or no dark- 
ening of the exciple. In fact, we have been unable to state for many 
type specimens whether the exciple is open or closed, because one 
section of an ascocarp would differ from the next. 

3. Striation of the exciple. The degree to which excipular labia 
become striate is not a constant feature. Many individuals exhibit 
labia which are entire in some portions, crenate in others, and striate 
elsewhere (see figs. 14-16, 24-27). Furthermore the number of 
lamellae or dentations in prominently striate labia seems to be 
correlated with age. 

4. Convergent or divergent labia. This character is only partially 
useful as a taxonomic criterion. Some species have constantly open 
disks; others vary from rimiform at one stage to gaping at others. 
(See figs. 31-35, 66, 69, and 70.) 

All four of the above characters are useful in separating individual 
species but intergrade far too much to be maintained as sectional 
criteria. We have therefore decided to dispense with sectional 
names in this study. It is impractical to use the myriad sections 
now recognized until the genera of the Graphidaceae are monographed 
and new sectional criteria established. 

In investigating the Graphidaceae of Mexico, we have attempted 
to assay specimens for lichen acids whenever possible. Unfortunately 
type material was sometimes too fragmentary to extract, and color 
reactions alone had to be used. A KOH-+ red reaction is often 
caused by an unknown substance that does not recrystallize in 
G.A.o-T. Stictic acid may give a KOH— reaction even though a 
microchemical test with G.A.o-T. is positive. This color reaction 
is complicated when the thallus is discolored or the acid is present in 
seemingly very low concentration. 

Salacinic acid and atranorine, so common in foliose lichens, appear 
to be very rare in the Graphidaceae. On the other hand, norstictic 
acid (KOH-+ red) seems to be one of the most common acids. In 
addition to atranorine, stictic acid, norstictic acid, and salacinic acid, 
we have demonstrated zeorine, lecanoric acid, protocetraric acid, 
lichexanthone, fatty substances, and several unknown acids. 

There seems to be some correlation between chemistry and ana- 
tomical features in a few species. Graphina parilis, with 8-spored 
asci and stictic acid, and G. sulcata, with 2- to 6-spored asci and 
norstictic acid, are examples of this phenomenon. In other species 
there are two or more chemical strains but with no apparent cor- 
relation with anatomy. It remains to be seen exactly what taxonomic 
importance chemical components will have in this family. With the 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 67 


exception of Rhizocarpon (Runemark, 1956), no crustose groups 
have been studied chemically. To aid other investigators we have 
appended on page 112 a list of type specimens which we have examined 
for color reactions and lichen acids. 

In this study we have described five new species, Graphina peplo- 
phora, Graphis glaucopis, G. stygioarachnoidea, Phaeographina strigops, 
and P. elliptica, and made two new combinations, Graphina mexicana 
(Zahlbr.) Wirth & Hale and Phaeog~aphina leiogrammodes (Kremplh.) 
Wirth & Hale. 

For the prompt loan of valuable type specimens we are very much 
indebted to Dr. Sten Ahlner (Naturhistoriska Riksmuseum, Stock- 
holm), Dr. R. Alava (Botanical Institute, Turku), Dr. Charles Baehni 
(Conservatoire et Jardin Botaniques, Geneva), Dr. I. Mackenzie 
Lamb (Farlow Herbarium, Cambridge), Dr. Josef Poelt (Botanische 
Staatssammlung, Munich), Dr. K. Rechinger (Naturhistorisches 
Museum, Vienna), Dr. C. T. Rizzini (Jardim Botdnico, Rio de 
Janeiro), Dr. H. Roivainen (Botanical Museum, Helsinki), Dr. 
Robert Shaffer (Herbarium, University of Michigan, Ann Arbor), and 
Dr. H. W. Vogelmann (University of Vermont, Burlington). 


Key to Species 


1. Ascocarps cinnabarred. ........ 39. Phaeographina chrysocarpa 
1. Ascocarps brown, black, gray, or white. 
2. Spores always 2 locular. ......4.4.. 36. Melaspilea polymorpha 


2. Spores 3 to many locular. 
3. Spores transversely 3 septate (4 locular) only, no other septae present. 
4. Spores brown, thick walled. 
5. Ascocarps dendritieally branched; exciple entirely black; disc black or 


gray . 2. 1. ee ee ee ~ ees 45. Phacographis sericea 
5. Ascocarps more or less unbranched; exciple red-brown, black only in 
spots; disc brown. ...... . . . 57. Phaeographis inusta 


4. Spores hyaline (except when senile or obviously shrunken), thin walled. 
6. Ascocarps sessile; exciple black, covered by a white C+ red veil. 
22. Graphis afzelii 
6. Ascocarps more or less immersed, C—; exciple uncarbonized. 
7. Exciple well developed, much thickened below; ascocarps lirelline, 


immersed... . 1... ee eee 29. Graphis grammatica 
7. Exciple poorly developed, bark-inspersed; mature ascocarps ellip- 
tical, with flaring margins. . . . . 32. Graphis platycarpella 


3. Spores transversely 4 or more septate or muriform. 
8. Spores transversely septate only, never muriform or apically biocellate. 
9. Spores brown, thick walled. 

10. Ascocarps sessile, prominent, often stromatoid; exciple entirely 
black, closed, much thickened below; hymenium with oil 
droplets. .... rr: 2h Phaeographis exaltata 

10. Ascocarps somewhat “emergent, never sessile, thalline border 
never prominent or stromatoid; exciple lacking or not greatly 
thickened below; hymenium not inspersed. 


68 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


11. Ascocarps rarely branched, slender, elongate; exciple black, 
dimidiate. . ..... . . 46. Phaeographis sexloculata 
11. Ascocarps dendritically branched, to 0.4 mm. wide; exciple 
closed, brown and black. . . 43. Phaeographis dendritica 
9. Spores colorless (brownish when senile or shriveled), thin walled. 
12. Exciple totally uncarbonized, with no black or very dark brown 
tissues. 
13. Labia lightly sulcate, sulcae externally concealed by a smooth 
thalline veil. 
14. Ascocarps slender, lirelline. . . . . 30. Graphis implicata 
14. Ascocarps rotund to elliptical . . 7. Graphina hololeucoides 
13. Labia entire. 
15. Ascocarps dendritic to asteroid, immersed; spores muriform 
at maturity. ....... .. 21. Graphina virginea 
15. Ascocarps more or less unbranched, elliptical, arranged in 
rows, subsessile; spores never muriform. 
28. Graphis glaucopis 
12. Exciple with black or dark tissue (often limited to the labial 
apices). 
16. Exciple carbonized laterally only (often the labial apices only), 
pale or light brown below. 
17. Labia consisting of partially free carbonized hyphae; exciple 
otherwise totally decolorized. 
34. Graphis stygioarachnoidea 
17. Labia entire or striate but never composed of free individually 
carbonized hyphae. 
18. Labia distinctly striate. 
19. Spores up to 30u long. 
20. Ascocarps quite sessile, prominent; exciple heavily 


inspersed with crystals . . 26. Graphis endoxantha 
20. Ascocarps emergent to subsessile; exciple without 
crystals . . . .. . . . 33. Graphis proserpens 


19. Spores more than 30u long, usually much longer. 
21. Exciple and labia black; sulcae few. 
49. Graphis striatula 
21. Exciple partially brown; sulcae many, deep, black; 
spores finally muriform. . . 13. Graphina parilis 
18. Labia never striate. 
22. Labia barely fuscescent, never highly carbonized; exciple 
pale, more or less closed; ascocarps elliptical. 
28. Graphis glaucopis 
22. Labia distinctly carbonized; exciple more or less dimidi- 
ate; ascocarps lirelline. 
23. Ascocarpa distinctly pruinose . 24. Graphis caesiella 
23. Ascocarps black, epruinose. 
48. Graphis scripta group 
16. Exciple carbonized laterally and below or dark brown below. 
24. Ascocarps caesiopruinose; labia more or less divergent. 
35. Graphis subamylacea 
24. Ascocarps black or covered by the thallus, epruinose; labia 
more or less convergent. 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 69 


25. Spores very long and slender, vermiform; ascocarps con- 
colorous with the thallus; labia denticulate. 
50. Graphis vermiformis 
25. Spores not vermiform; ascocarps showing some traces of 
black. 
26. Labia more or less entire. 
27. Thallus KOH—, P—; labia always lightly crenate or 
denticulate . ...... . 31. Graphis longula 
27. Thallus KOH+ reddish; labia entire. 

28. Ascocarps robust, covered nearly to the apex by a 
heavy thalline veil; spores 2-6 per ascus, over 
75u long; no acids demonstrated. 

23. Graphis anguilliformis 

28. Ascocarps slender, Opegrapha-like, black; spores 8 
per ascus, under 30y long; norstictic acid present. 

25. Graphis desquamescens 
26. Labia denticulate to striate. 
29. Exciple distinctly carbonized throughout. 

30. End locules of some spores biocellate. 

1. Graphina acharii 

30. End locules of all spores entire. 

31. Ascocarps robust, about 0.6 mm. wide; labia 
deeply striate . .. . 27. Graphis flexibilis 
31. Ascocarps slender, usually less than 0.4 mm. wide; 

labia crenate to denticulate. 
31. Graphis longula 

29. Exciple open or brown below. 
32. Spores eventually muriform; stictic acid present. 

13. Graphina parilis 

32. Spores never muriform. 

33. Exciple dark brown; labia dissected into free, 
disconnected black lamellae. 
33. Graphis proserpens 
33. Exciple completely black; labia confluent with the 
rest of the exciple. 

34. Spores 2-6 per ascus, more than 40y long; 
exciple usually closed below, at least by a 
thin carbonized strip; ascocarps flexuose, 

occasionally branched. 
31. Graphis longula 
34. Spores 8 per ascus, less than 45y long; exciple 
open below, rarely almost closed; ascocarps 
Opegrapha-like, usually unbranched, straight. 
49. Graphis striatula 
8. Spores transversely and longitudinally septate (sometimes only in the 

apical portions). 
35. Exciples with no carbonized tissue. 
36. Spores one per ascus. 
37. Ascocarps and thallus concolorous. 
38. Labia striate; exciple pale below the hymenium. 

3. Graphina bipartita 


70 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


38. Labia entire; exciple black or very dark below the hymenium. 
4. Graphina confluens 
37. Ascocarps (at least the disc) darker than the thallus. 

39. Hymenium with prominent crystalline inspersions. 

42, Phaeographina elliptica 

39. Hymenium without crystals. 

56. Phaeographina scalpturata 
36. Spores 2-8 per ascus. 
40. Labia distinctly striate, the striae often externally concealed 
by a thalline covering. 
41. Ascocarps subsessile, lighter than the thallus. 
42. Ascocarps slender, quite flexuose; not mealy. 
2. Graphina balbisii 
42. Ascocarps robust, round to straight, mealy. 
43. Ascocarps round to elliptical; spores 4—6 per ascus. 
7. Graphina hololeucoides 
43. Ascocarps elongate; spores 2—4 per ascus. 
14. Graphina peplophora 

41. Ascocarps more or less immersed, concolorous with the thallus. 

44, Exciple basally continuous with the thallus by a carbonized 
band; spores 4-8 per ascus. . 19. Graphina triangularis 
44. Exciple not basally continuous, colorless below; spores 8 
perascus... . . .. . « 12. Graphina palmeri 

40. Labia entire, not striate. 

45. Exciple basally continuous with thallus by a band of more or 
less carbonized tissue; ascocarps dendritic, fused; disc wide, 
pruinose. ....... ee ee 8. Graphina insignis 

45. Exciples discrete, never : basally continuous. 

46. Spores distinctly brown; exciples red-brown, distinctly 
closed; labia more or less divergent. 
47. Ascocarps emergent, with a prominent thalline margin; 
discs pruinose; spores more than 25y long. 
37. Phaeographina asteroides 
47. Ascocarps immersed, without a thalline margin; discs 
black; spores less than 22y long. 
41. Phaeographina strigops 
46. Spores colorless (except brownish when senile); exciples 
pale to fuscescent, barely closed or open; labia more or 
less convergent. 
48. Spores more than 50y long, usually much more; ascocarps 
dendritically branched, never fissurine. 
21. Graphina virginea 
48. Spores less than 40u long; ascocarps rarely dendritically 
branched, often fissurine. 
49. Ascocarps intricately intertwined and grouped, gaping. 
11. Graphina mexicana 
49. Ascocarps never intricately intertwined. 
50. Ascocarps KOH— or pale, short, more or less un- 
branched, gaping; labia dark brown or fuscescent. 
20. Graphina virginalis 
50. Ascocarps KOH-+ purple to black, long slender, 
branched, usually not gaping; labia pale or 
reddish. . . . .. . . 16, Graphina scolecitis 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 71 


35. Exciple with carbonized tissue. 
51. Spores one per ascus. 
52. Labia striate. 


53. Exciple closed, entirely black . . . 10. Graphina macella 
53. Exciple more or less open, pale or brownish, only the labia 
carbonized . ....... . . 38. Graphina bipartita 


52. Labia entire. 
54. Exciples closed, often basally continuous; ascocarps round to 


lirelline. . . . . . . . . 4 Graphina confluens 

54, Exciples more or less open, never basally continuous; ascocarps 
lirelline. 

55. Disc very wide, pruinose. . . . 40. Phaeographina sp. 

55. Disc narrow, epruinose.... . 6. Graphina hiascens 


51. Spores 2-8 per ascus. 
56. Exciples basally continuous by a more or less carbonized layer. 
57. Labia striate, pale; spores over 45y long. 
19. Graphina triangularis 
57. Labia entire, brown; spores up to 15 long. 
8. Graphina insignis 
56. Exciples not basally continuous. 
58. Labia divergent, disc very wide. 
38. Phaeographina caesiopruinosa 
58. Labia more or less convergent, disc rimiform. 
59. Exciples mostly brown or pale, only the labia fuscescent 
or darker brown. 
60. Labia striate; ascocarps not fissurine; spores more than 


45u long... we ee 1D Graphina palmeri 
60. Labia entire; ascocarps fissurine at maturity; spores less 
than 40u long. . . . . . . 20. Graphina virginalis 


59. Labia and/or exciples distinctly carbonized. 
61. Labia entire, sinuous or barely striate. 
62. Ascocarps intricately intertwined in discrete groups, 
naked, sessile, black. 
18. Graphina sulcatula var. conglomerata 
62. Ascocarps not intertwined in discrete groups. 
63. Ascocarps asteroidly branched; labia sulcate in some 
portions; exciple dimidiate. 
5. Graphina elongata 
63. Ascocarps not asteroidly branched; labia substriate 
to sulcate. 
64. Exciple more or less open, brown or pale below. 
13. Graphina parilis 
64. Exciple closed, entirely black. 
9. Graphina inturgescens 
61. Labia distinctly dentate or sulcate. 
65. Ascocarps asteroidly branched. 
5. Graphina elongata 
65. Ascocarps never asteroidly branched. 
66. Exciple thick and more or less completely carbonized 
below the hymenium. 
67. Spores 65-150u% long, commonly muriform only 
in the end cells; labia regularly striate. 
1. Graphina acharii 


72 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


67. Spores 30-45u long; labia irregularly dissected. 
15. Graphina rimulosa 
66. Exciple brown or lacking below the hymenium. 

68. Exciple completely black laterally, dimidiate and 
brownish below the hymenium; labia lightly 
few suleate . . . . 47. Graphina sophistica 

68. Exciple partially brown laterally; labia more or 
less free, black, embedded in the brownish 
exciple. 

69. Labia very irregularly dissected and lacerate. 
15. Graphina rimulosa 
69. Labia lamellar, regular. 
70. Spores 2-6 per ascus, with 5-8 transverse 
septae, 20-35 long; norstictic acid present. 
17. Graphina sulcata 
70. Spores 8 per ascus, with 12-20 transverse 
septae, 28-80y long; stictic acid. 
13. Graphina parilis 


1. Graphina 


1. Graphina acharii (Fée) Mill. Arg. Mem. Soc. Phys. Hist. Nat. Genéve 29, 

no. 8:38. 1887. FIaurReE 1 

Graphis acharii Fée, Essai Crypt. Ecorces, 39, pl. 10, fig. 4, 1824. Type: 
South America (not seen). 


Thallus smooth to rough, continuous. Ascocarps sessile, large, very 
variable, covered up to or nearly to the apex by a thalline margin; 
exciple black, closed; labia convergent, striate. ‘Spores 2-8 per ascus, 
18-30 1-6 locular, colorless or brownish, 15-25% 65-150 u. 

Reactions: KOH+ red, P—, o-T—, no acids demonstrated.’ 

Specimens examined: Chiapas: El Suspiro, Hale 20230, 20237, 
north of Tuxtla Gutiérrez, Hale 20067, Lagos de Monte Bello, Hale 
20437 (US). Morelos: Cuernavaca, Pringle 429, 15373 (FH, VT). 
Tamaulipas: Tamasopo, Pringle 34a (FH), 166 p.p. (FH, MICH). 
Vera Cruz: South of Catemaco, Hale 19821 (S, US), Orizaba, Pringle 
3.n. (FH, MICH), Jalapa, Pringle 15398 (FH, MICH, VT). 

Graphina acharii shows an enormous amount of variation in ascocarp 
size and shape and spore septation and color. The ascocarps range 
from elongate and black above to oryzaeform and totally covered 
with thalline tissue. The spores may be muriform throughout or 
only geminate at the ends. The completely muriform spores are 
quite often pale brownish. Considering these characters, we find it 
difficult to maintain the entity originally described by Krempelhuber 
as Graphis inturgescens, which differs from Graphina acharwi only in 
the subentire labia. 


2 In these lists of the species, all specimens examined contained the substances indicated under the heading 
‘Reactions’ unless otherwise indicated. 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 73 


Ficures 1—3.—1. Graphina acharii (Fée) Mill. Arg. (Pringle 166 p.p.). 2. G. balbisti (Fée) 
Mill. Arg. (Pringle 242). 3. G. peralbida (Nyl.) Zahlbr. (holotype, Pringle 222). 


Graphina acharvi apparently exhibits the same type of variation 
that is found in Phaeographina chrysocarpa. A broad treatment of 
the species will be necessary, based on careful examination of the 
holotype and the types of several related species, Graphis ilinata 
Eschw. in Mart., Phaeographis cerviculata Redgr., Graphina vestitoides 
Fink, G. dealbata (Nyl.) Mill. Arg., and G. heteroplaca Mill. Arg. 

2. Graphina balbisii (Fée) Mill. Arg. Flora 65:397. 1882. FIGURE 2 
Graphis balbisii Fée, Essai Crypt. Ecorces 48, pl. 10, fig. 5, 1824. Type: 
South America (not seen). 

Thallus continuous, smooth to irregular. Ascocarps lighter in color 
than the thallus, usually unbranched, quite slender and flexuose, 1-4 
mm. long; disc rimiform; exciple red-yellow, closed, about 250 u high; 
labia 1-3 sulcate, convergent. Spores 4-5 per ascus, 13-18X4-6 
locular, 18-26 50-110 yu, occasionally somewhat brownish. 

Reactions: KOH+ red, P—, o-T—, no acids demonstrated. 

Specimens examined: Morelos: Cuernavaca, Pringle 242 (VT), 426 
(MICH). 

Redinger (1933, p. 61) placed Graphina balbisii and his variety 
monospora in section Chlorographina. However, the type of var. 
monospora, and what may be a fragment of the type of Graphis 
balbisit in MICH, both show distinctly striate labia, as Miller (1887, 
p. 45) had earlier observed for the species. 


74 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Pringle 119, reported by Eckfeldt (1892, p. 252) as “‘Graphis balbisii 

Nyl.,” is referable to Graphis implicata Fée. 
3. Graphina bipartita Mill. Arg. Rev. Mycol. 10:19. 1888. FIGcurRE 3 
Graphis peralbida Nyl. Lich. Trop. Singapore et Labuan 42. 1891. Holo- 
type: Las Palmas, San Luis Potosi, Mexico, Pringle 222 (H, isotypes in 
FH, VT). 
Graphina peralbida (Nyl.) Zahlbr. Cat. Lich. Univ. 2:418. 1923. 

Holotype: Assuncion, Paraguay, Balansa 1876 (G). 

Thallus nitid to minutely roughened, continuous or somewhat 
fissured. Ascocarps immersed to emergent, unbranched or nearly 
so, usually concolorous with the thallus, 1-3 mm. long, about 0.4 mm. 
wide, flexuose and somewhat intricate; disc very narrow, occasionally 
blackish; exciple open or partially closed; labia convergent, with 
1-3 dark or carbonized striae; hymenium 60-100 » high. Spores one 
per ascus, densely muriform, hyaline or brownish, 23-40 60-100 u. 

Reactions: KOH-- reddish, norstictic acid. 

In the type description, @. peralbida is cited from a Mexican col- 
lection by Eckfeldt. One year later, Eckfeldt (1892, p. 252) listed 
Graphis peralbida giving Pringle as the collector. As the holotype 
specimen (H) is marked as coming from San Luis Potosi and is identical 
with Pringle 202 in FH and VT, it is certain that this collection by 
Pringle is the type and that Nylander made a mistaken reference in 
his publication. This type is identical with Graphina bipartita, which 
is probably pantropical, as we have seen several specimens from Hawaii. 
4. Graphina confluens (Fée) Mill. Arg. Mem. Soc. Phys. Hist. Nat. Genéve 

29, no. 8:45. 1887. Fieures 4-13 

Arthonia confluens Fée, Essai Crypt. Ecorces 55, pl. 14, fig. 6, 1824. Type: 
South America (see below). 

?Graphis delicatula Fée, Bull. Soc. Bot. France 21:28. 1874. Syntypes: Brazil, 
Glaziou 3226, 3396 (not seen; see following species). 

Graphis macrospora Kremplh. Flora 59:380. 1876. Syntypes: Brazil, Glaziou 
3379, 3380 (M), 3226 (not seen; this is also a syntype of G. delicatula). 

Graphina macrospora (Kremplh.) Mill. Arg. Flora 71:508. 1888. 

?Graphina platygrapta Mill. Arg. Flora 71:495. 1888. Type: Cayey, Puerto 
Rico, Sintents 25 p.p. (not seen). 

Graphina epiglauca Mill. Arg. Bull. Soc. Bot. Belg. 32:152. 1893. Type: 
Costa Rica, Pittier (US, isotype). 

Graphina collatinensis Redgr. Hedw. 73:59, 1933. Type: Collatina, Espiritu 
Santo, Brazil, Bornmiiller 13 (not seen). 

Graphina collatinensis var. ocellartiformis Redgr. Ark. Bot. 26A, no. 1:24. 
1933. Syntypes: Brazil, Malme 2351, 2416, 3565, 3576 (8). 

Graphina collatinensis var. lirelliformis Redgr. Ark. Bot. 26A, no. 1:25. 
1933. Holotype: Corcovado, Rio de Janeiro, Brazil, Malme 84 (8). 


Thallus light colored, continuous, thin to quite thick, often des- 
quamescent. Ascocarps very variable in size and shape, immersed 
to emergent or sessile, round to lirelline, always covered by a thalline 
margin; disc narrow to wide, lightly to heavily pruinose; exciple 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 15 


closed, brown or carbonaceous, often basally continuous in a sub- 
thalline layer; labia more or less entire, distant, and divergent; hyme- 
nium intact or rarely with an incipient columella. Spores one per 
ascus, densely muriform, hyaline, yellowish, or pale brownish, 25-50 
65-175 p. 

Reactions: See below. 

Specimens examined: Chiapas: El Suspiro, Hale 20155, 20169 (US). 
Tamaulipas: Tampico, Pringle 8 (W). Vera Cruz: South of Catemaco, 
Hale 19803 (S, US). 

Arthonia confluens Fée was based on two specimens, one from South 
America on Cinchona and one from Guadeloupe on Jacquinia. Miiller 


Ficures 4-9.—4. Graphina macrospora (Kremplh.) Mill. Arg. (syntype, Glaziou 3379). 
5. G. confluens (Fée) Mill. Arg. Variation in spore shape. 6. G. epiglauca Mill. Arg. 
(isotype, Pittier s.n.). 7. G. collatinensis var. ocellartiformis Redgr. (syntype, Malme 
3565). 8. G. collatinensis var. lirelliformis Redgr. (holotype, Malme 84). 9. G. confluens 
(Fée) Mull. Arg. (Pringle 8). 


76 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


discovered that the syntypes were heterogeneous and referred the 
Guadeloupe specimen to Opegrapha. We have examined the same 
specimen (from Rio de Janeiro) and can confirm that it is an Opegrapha. 
The South American material, which becomes the lectotype, has not 
been seen by us, but Miiller’s description leaves little doubt in our 
minds as to its identity. 

Graphina confluens seems to be one of the most variable, complex, 
and widespread Graphina species yet encountered. ‘The entity cuts 
across generic and family boundaries with a continuous, interrelated 
series of forms which are difficult to separate. In addition, quite a 
number of species have been described which are probably additional 
variants. The difficulties involved in this group may best be sum- 
marized by examining the following salient features. 

1. The extremes of thallus form in this group are quite distinct. 
At one end of the spectrum of variants are thin, barely continuous, 
closely adherent thalli; at the opposite end are those forms with thick 
desquamescent thalli, often with a basal carbonaceous layer. All 
intermediate stages can be found. In the related Graphina platy- 
carpina Zahblbr. the thallus is areolate. 

2. The ascocarps vary from completely lirelline to completely 
round, commonly within the same thallus. Specimens with immersed 
apothecia-like ascocarps approach the Thelotremataceae very closely. 
The degree of immersion in the species is highly variable. The ex- 
tremes are distinct, but the presence of numerous intermediate speci- 
mens renders separation nearly impossible. 

3. The exciples vary in structure from more or less completely car- 
bonized forms (Graphina epiglauca, G. collatinensis var. ocellaritformis, 
and G. macrospora, figs. 4, 6, 7) to entirely brown extremes showing 
carbonization only in the subthalline layer (G@. collatinensis var. 
lirelliformis, fig. 8). Between these two extremes one can find exciples 
which are carbonized only in spots or blackened above and brown be- 
low (figs. 10-13). 

One of the syntypes of G. macrospora (Glaziou 3379) has exciples 
which show distinct columellae forming. In all other details of anat- 
omy it is identical with the syntypes of G. collatinensis var. ocellari- 
iformis. The presence of columellae is not altogether unexpected and 
provides further evidence of the unity of the Graphidaceae and much 
of the Thelotremataceae. 

4. Spores that are large and densely muriform, with but one per 
ascus, are characteristic of all individuals of this polymorphic species. 
However, the spores vary from completely hyaline to yellowish or 
brownish. It is thus sometimes difficult to decide whether certain 
specimens should be placed in Graphina or Phaeographina. 

5. The distribution of Graphina confluens as here delimited extends 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 77 


Ficures 10-13.—10. Graphina confluens (Fée) Mill. Arg. (Wright 23). 11. G. confluens 
(Hioram 5300). 12. G. confluens (Williams 7102). 13. Graphis glaucoleuca Nyl. nom. 
nud. (Wright 4). 


from Mexico, Costa Rica, and the West Indies, to Brazil and Africa. 
There seems to be no correlation between anatomy and distribution. 
6. Lichen acids are present in a bewildering array, apparently unre- 
lated to either anatomy or geographic distribution. Acids demon- 
strated are as follows (the numbers in parentheses indicate number of 
specimens) : 
Stictic acid only: Philippines (1). 
Stictic acid, lichexanthone, and a fatty substance: Costa Rica (1), Mexico (3), 
Cuba (3), Jamaica (1), Puerto Rico (2), and Brazil (2). 
Stictic acid and an unknown: Puerto Rico (2). 
Lichexanthone only: Cuba (1) and Brazil (3). 
Lichexanthone, zeorine, atranorine: Brazil (1). 
Norstictic acid: Cuba (1) and Brazil (1). 
5. Graphina elongata (Vain.) Zahlbr. Cat. Lich. Univ. 2:405. 1923. 
Ficaures 14-16 
Graphis elongata Vain. Acta Soc. Faun. Fl. Fenn. 7, no. 7:107. 1890. 
Holotype: Sitio, Minas Gerais, Brazil, Vainio 782 (TUR). 
Graphina elongatoradians Fink, Mycologia 19:218. 1927. Holotype: Manatf, 
Puerto Rico, Fink 2049 (MICH). Reactions: KOH+ red, P— (?), 
o-T—, no acids demonstrated. 


Thallus smooth, continuous. Ascocarps slender, semiemergent to 
subsessile, occurring in large, radiately branched asteroid groups up to 


15 mm. in diameter; exciple black, open; labia 1-3 striate, convergent; 
686-879—63——2 


78 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


hymenium 50 u high. Spores 2-4 (-6) per ascus, 7-9 1-4 locular, 
13-19 K 25-50 u. 

Reactions: KOH-+ red, norstictic acid. 

Specimen examined: Chiapas: Lagos de Monte Bello, Hale 21069 
(US). 

The excipular labia of this species are often only faintly striate, 
and it is necessary to examine sections of several ascocarps to ensure 
proper identification. The holotype of G. elongatoradians differs from 
that of G. elongata only in having somewhat smaller, less emergent 
ascocarps and in apparently lacking norstictic acid, although the 
color reaction indicates its presence. 

6. Graphina hiascens (Fée) Mill. Arg. Mem. Soc. Phys. Hist. Nat. Genéve 29 

no. 8:42. 1887. Figures 17-20 

Opegrapha hiascens Fée, Suppl. Essai Crypt. Ecorces, 25, pl. 36, fig. 1, 1837. 
Type: South America (not seen, isotype at MICH?). 

?Opegrapha endochroma Fée, Essai Crypt. Ecorces, 31. 1824. Type: 
Tropical America (not seen). 

Graphis subserpentina Nyl. Acta Soc. Sci. Fenn. 7:465. 1863. Type: 
Ceylon, Gardner (US, isotype). 

Graphina subserpentina (Nyl.) Miill. Arg. Bull. Soc. Bot. Belg. 32:152. 
1893. 

Graphis macgregortt Vain. Ann. Acad. Sci. Fenn. 15A, no. 6:207. 1921. 
Type: Guam, MacGregor 607 (US, isotype). 

Graphis orientalis Vain. Ann. Acad. Sci. Fenn. 15A, no. 6:206. 1921. 
Type: Philippines, Merrill 6716 (isosyntype, US). 

Graphina orientalis (Vain.) Zahlbr. Cat. Lich. Univ. 2:416. 1923. 

Graphina macgregorit (Vain.) Zahlbr. Cat. Lich. Univ. 2:413. 1923. 


Thallus smooth, granular, or warty, continuous. Ascocarps un- 
branched, lightly flexuose, semiemergent or subsessile, 0.5-2 mm. 
long, about 0.5 mm. wide, black above, with a more or less prominent 
lateral thalline margin; exciple black, open; labia more or less con- 
vergent, entire; hymenium 100-200 » high. Spores one per ascus, 
occasionally faint brown, 13-161-8 locular, 18-32 70-115 uy. 

Reactions: KOH-+ red, nostictic acid. 

Specimens examined: Tamaulipas: Tampico, Pringle 5454, 5457 
(MICH). 

Fée published the name Opegrapha endochroma in 1824. Later, in 
his Supplement, he renamed the same entity O. hiascens, supposedly 
on the basis of better material. Miller, in transferring it to Graphina, 
chose to use the later epithet Aiascens and listed O. endochroma as a 
synonym, Although endochroma has nomenclatural priority, the new 
combination should not be made until the materials in Fée’s collec- 
tions have been thoroughly examined. 

An isosyntype of Graphis orientalis (US) differs from typical Gra- 
phina hiascens only in having totally immersed ascocarps. In our 
opinion this is insufficient grounds for maintaining this entity as a 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 79 


<  @ 


Ficures 14-16.—14. Graphina elongata (Vain.) Zahlbr. (holotype, Vainio 782). 15-16. G. 
elongatoradians Fink (holotype, Fink 2049). 


Ficures 17-20.—17. Graphina subserpentina (Nyl.) Mill. Arg. (isotype, Gardner s.n.). 
18. G. macgregorii (Vain.) Zahlb. (isotype, MacGregor 607). 19. G. ortentalis (Vain.) 
Zahlbr. (isosyntype, Merrill 6716). 20. G. hiascens var. clausior (Vain.) Zahlbr. (isotype, 
Ramos &§ Edano 29548). 


distinct species, since the degree of immersion is highly variable in this 
species. Isotypes of G. macgregorit and G. subserpentina differ from 
G. hiascens only in minor intergrading characters. An isotype of 
G. hiascens var. clausior (Vain.) Zahlbr. (US) from the Philippines is 
distinct in having elongate intricate ascocarps (fig. 20) and in pro- 
ducing stictic acid only. 


80 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


7. Graphina hololeucoides (Nyl.) Mill. Arg. Flora 65:397. 1882. 
Figure 21 

Graphis hololeucotdes Nyl. Ann. Sci. Nat. Bot. ser. 4, 20:266. 1863. Type: 
Mexico, Ghiesbreght (H, isotype). 

Thallus uneven, often discontinuous. Ascocarps sessile or sub- 
sessile, lighter than the thallus, slightly mealy, unbranched, round to 
elliptical, 0.5-0.8 mm. wide, 1-1.5 mm. long; exciple barely closed, 
colorless below; labia internally irregularly striate, slightly darkened; 
hymenium about 100 » high. Spores 4-6 per ascus, 20-3014 
locular, 15-18%60-100 u. 

Reactions: Thallus KOH—, P—; ascocarps KOH+- faint, 
P+ red. No microchemical tests made. 

Although the isotype is too small to extract, the color reactions 
suggest the presence of protocetraric acid. 

8. Graphina insignis (Vain.) Zahlbr. Cat. Lich. Univ. 2:410. 1923. 
Figures 22, 23 

Graphis insignis Vain. Acta Soc. Faun. Fl. Fenn. 7, no. 7:112. 1890. 
Lectotype: Carassa, Minas Gerais, Brazil, Vainio 1209 (TUR). 

Graphina insignis var. primaria Redgr. Ark. Bot. 26A, no. 1:21. 1933. 
Syntypes: Brazil, Malme 2299, 3509 p.p. (8). 

Graphina insignis var. imperfecta Redgr. Ark. Bot. 26A, no. 1:21. 1933. 
Holotype: Brazil, Malme 3514 (8). Reactions: KOH+ red, P—, gummy 
residue, atranorine only. 

Graphina insignis var. tartarea Redgr. Ark. Bot. 26A, no. 1:22. 1933. 
Syntypes: Brazil, Malme 2549, 3509 p.p. (S). 

Thallus thick, nitid, continuous or fissured, usually carbonized near 
the substrate, often with soredia-like eruptions. Ascocarps im- 
mersed, usually bordered by a lighter area of thallus, irregular, flexuose, 
simple to asteroidly branched, often anastomosing into large contorted 
fertile areas, 1-5 mm. long, up to 1 mm. wide; disc plane or depressed, 
caesiopruinose; exciple closed, pale to dark brown, sometimes car- 
bonized at the very base; labia divergent, entire; hymenium 100-175 yu 
high, epithecium prominent. Spores 8 per ascus, ovoid, 4-5 1-3 
locular, 5-8 10-15 u. 

Reactions: KOH+ red, P—, o-T—, gummy residue, no acids 
demonstrated. 

Specimen examined: Chiapas: Lagos de Monte Bello, Hale 20449 
(DUKE, REN, S, TNS, US). 

Graphina insignis was placed by Redinger in the section Platy- 
graphopsis. However, examination of the Vainio syntypes and 
Redinger’s types showed that the exciples are usually uncarbonized 
or only rarely carbonized at the base. The thick band of black tissue 
which separates the thallus from the substrate is quite variable, and 
it seems superfluous to name each variation. 

This species is apparently restricted to substrata of decaying 
organic matter, such as rotten wood, old twigs, grass, and detritus on 


WIRTH & HALE—MEXICAN GRAPHIDACEAE Sl 


Ficures 22-23.—22. Graphina insignis (Vain.) Zahlbr. (lectotype, Vainio 1209). 23. G. 
insignis (Hale 20449). 


rocks. It forms a heavy thallus with characteristic soredia-like 
outgrowths and frequently fused ascocarps. It must have a very 
rapid growth rate to be able to colonize such unstable habitats. 

All of the specimens react KOH-+ red and yield from acetone a 
clear gummy residue which is unrecrystallizable. Only one specimen 
(the holotype of var. ¢mperfecta) yielded atranorine. 


9. Graphina inturgescens (Kremplh.) Mill. Arg. Flora 71:545. 1888. 


Fiaures 24-27 

Graphis inturgescens Kremplh. Flora 59:383. 1876. Holotype: Brazil, 
Glaztou 6286 (M). 

Graphis phaeospora Vain. Acta Soc. Faun. Fl. Fenn. 7, no. 7:97. 1890. 
Holotype: Sitio, Minas Gerais, Brazil, Vainio 682 (TUR). 

Graphina acharii var. subintegra Zahlbr. Ann. Mycol. 19:230. 1921. 
Types: Tampico, Tamaulipas, Mexico, Pringle 5 p.p., 25 p.p. (MICH, 
isosyntypes). 

Phacographina phaeospora (Vain.) Zahlbr. Cat. Lich. Univ. 2:443. 1923. 

Thallus continuous, thick, somewhat roughened, saxicolous or 
corticolous. Ascocarps sessile, flexuose, elongate, rarely branched, to 
6 mm. long, covered up to or nearly to the apex by a thalline margin; 
exciple black, closed; labia convergent, entire or crenate; hymenium 
150-300 » high. Spores 2-4 (-6?) per ascus, colorless or nearly so, 
17-30 1-5 locular, 15-30 80-160 u. 


82 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Reactions: KOH+ reddish, P—, no acids demonstrated. 

We have been unable to find any differences between the various 
species listed above other than the fact that the holotype of G. 
inturgescens 1s saxicolous while the others are corticolous. The spores 
in all three types are colorless or rarely pale brown, Vainio’s name 
phaeospora notwithstanding! The labia vary from entire to crenate 
and are covered by a thin to thick thalline veil. 

It is altogether likely that @. inturgescens is referable to the poly- 
morphic G. achariz. Graphina inturgescens can be provisionally 
separated by the subentire rather than distinctly striate labia. 

10. Graphina macella (Kremplh.) Mill. Arg. Flora 63:23. 1880. Fiaure 28 
Graphis macella Kremplh. Flora 59:380. 1876. Holotype: Brazil, Glaziou 
6289b (M). 

Thallus continuous, smooth. Ascocarps semiemergent to sub- 
sessile, stout, sparmgly branched, straight to flexuose, black above, 
white striped, 1-5 mm. long, about 0.5 mm. wide; exciple black, 
closed; labia convergent, 1-4 sulcate; hymenium 150-200 » high. 
Spores one per ascus, 25~35 3-12 locular, 20-40 x 65-130 uy. 

Reactions: KOH+ yellow, P—, 0-T—, G.E.—, no acids demon- 
strated. 

Specimen examined: Chiapas: Southeast of San Cristébal, Hale 
20275 (US). 

Graphina macella is closely related to G. acharii, from which it differs 
in having only one rather rotund spore per ascus. The Mexican ma- 
terial differs from the holotype of G. macella in having fewer striae, 
and the exciple is peculiarly infuscated below. ‘There is little doubt 
however as to the identity of the two specimens. 

11. Graphina mexicana (Zahlbr.) comb. nov. Figure 29 
Enterodictyon mexicanum Zahlbr. Ann. Mycol. 19:233. 1921. Holotype: 
Cuernavaca, Mexico, Pringle 20 (W). 

Thallus smooth to minutely roughened, continuous. Ascocarps in 
densely intricated clusters, appearing as gaping, very light-colored 
fissures; exciple colorless or rudimentary, barely closed to open; labia 
convergent at first, then somewhat spreading, composed at least 
partially of bark cells; hymenium about 90 u high. Spores 6-8 per 
ascus, occasionally halonate, 4-5 1-3 locular, 8-13 23-40 up. 

Reactions: KOH-+ red, P—, o-T—, no acids demonstrated. 

Specimen examined: San Luis Potosi: Las Palmas, Pringle 225 
(FH, VT). 

The holotype of Enterodictyon mexicanum is obviously referable to 
Graphina. Pringle 225 was determined by Miiller (1894, p. 92) as 
Medusulina nitida (Eschw. in Mart.) Miill. Arg., but we could find no 
trace of anything resembling stromatoid tissue in the specimen. 
Since the type of M. nitida has not been available for comparison and 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 83 


Ficures 24-27,—24-25. Graphina inturgescens (Kremplh.) Mull. Arg. (holotype, Glaziou 
6286). 26-27. Phaeographina phaeospora (Vain.) Zahlbr. (holotype, Vainio 682). 


Figure 29.—Graphina mexicana (Zahlbr.) Wirth & Hale (holotype, Pringle 20). 


is probably lost, it is best to transfer Enterodictyon mexicanum to 
Graphina, even though it may eventually be proved to be a synonym 
of Graphina nitida (Eschw. in Mart.) Mill. Arg. 


84 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


12. Graphina palmeri Zahlbr. Ann. Mycol. 19:231. 1921. Figure 30 


Type: Cuernavaca, Morelos, Mexico, Pringle 9 (MICH, isotype). 

Thallus much fissured and irregular. Ascocarps small, more or less 
immersed to semiemergent, 0.5-1.5 mm. long, slender, concolorous 
with the thallus; exciple closed, brown laterally, colorless below the 
hymenium; labia sulcate, with brown inclusions, covered by a thick 
thalline margin. Spores 8 per ascus, 12-153-6 locular, 15-18 45- 
75 p. 

Reactions: KOH-+ red, unknown acid in G.A. o-T. 

The unknown acid in the type is identical with that found in 
Graphina triangularis, @. virginea, and Phaeographis exaltata. In 
G.A. o-T. the crystals are somewhat similar to those of barbatic acid. 


13. Graphina parilis (Kremplh.) Miill. Arg. Bull. Soc. Bot. Belg. 32:152. 1892. 
Fieures 31-35 
Graphis parilis Kremplh. Flora 59:422. 1876. Holotype: Brazil, Glaziou 
3394 (M). Reactions: KOH+ yellowish, no chemical tests made. 
Graphis annulata Kremplh. Flora 59:445. 1876. Holotype: Brazil, Gla- 
ziou 3388 (M). Reactions: KOH+ yellowish, no chemical tests made. 
Graphina acromelaena Mill. Arg. Bull. Soc. Bot. Belg. 32:152. 1893. 
Holotype: San José, Costa Riea, Tonduz (G). 
Graphina achrophaea Mill. Arg. Bull. Herb. Boiss. 3:46. 1895. Type: 
Louisiana, Langlois 751 (US, isotype). 
Graphis subducta Vain. Ann. Acad. Sci. Fenn. 15A, no. 6:203. 1921. Type: 
Luzon, Philippines, Merrill 8576 (US, isotype). 
Graphina pringlei Zahlbr. Ann. Mycol. 19:230. 1921. Type: Tampico, 
Tamaulipas, Mexico, Pringle 11 (not seen). 
Graphina subducta (Vain.) Zahlbr. Cat. Lich. Univ. 2:426. 1923. 
Graphina acrophaea f. multilamellosa Redgr. Ark. Bot. 26A,no.1:52, 1933, 
Holotype: Colonia Risso, Paraguay, Malme 3545 (8). 
Graphina puiggarit f. corumbensis Redgr. Ark. Bot. 26A, no, 1:56. 19383. 
Syntypes: Matto Grosso, Brazil, Malme 3631, 3638 (8). 


Thallus continuous, smooth to minutely roughened. Ascocarps 
dispersed to clustered, semiemergent to subsessile, unbranched to 
3-4 branched, lightly flexuose to curved and intricate, black above, 
often with a raised thalline margin, 1-5 mm. long, 0.3-0.6 mm. wide; 
exciple black above, yellow or pale brown below, closed or open; 
labia convergent, 4-10 striate, often barely so in some portions of 
the ascocarps. Spores 8 per ascus, occasionally halonate, 12-20 1-7 
locular, 10-20 28-80 u. 

Reactions: KOH-+ yellow, stictic acid (accompanied by atranorine 
only in the holotype of Graphina acrophaea f. multilamellosa Redgr. 
and in Hale 21145 and Pringle 160 p.p.). 

Specimens examined: San Luis Potosi: Tamasopo, Pringle 160 p.p. 
(VT). Vera Cruz: South of Xico near Teocello canyon, Hale 21145, 
south of Catemaco, Hale 21085, northeast of Huatusco, Hale 19491 
(US). 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 85 


Ficures 30-35.—30. Graphina palmeri Zahlbr. (holotype, Pringle 9). 31. G. parilis (Krem- 
pelh.) Mill. Arg. (holotype, Glaziou 3394). 32. Graphis annulata Kremplh. (holotype, 
Glaziou 3388). 33. Graphina acrophaea Mill. Arg. (isotype, Langlois 751). 34. G. 
acrophaea {. multilamellosa Redgr. (holotype, Malme 3545). 35. G. putggarti £. corum- 
bensis Redgr. (syntype, Malme 3636). 


Examination of the seven types listed above in synonymy disclosed 
that these individuals are parts of a single variable species. Separat- 
ing them on the basis of open or closed exciples is difficult, since this 
character is not constant and intergrading forms are common. Spore 
size and septation are very similar, considering that these characters 
normally vary greatly even within a single ascocarp. The number of 
striae in the labia is apparently related to the age of the specimen, 
and therefore a name such as Graphina acrophaea f. multilamellosa is 
unnecessary. 


86 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


The type of Graphina pringlei Zahlbr. is not at Vienna and may 
have been lost in World War II. It seems most likely that it is a 
synonym of G. parilis, at least on the basis of the original description 
and on that of Redinger (1933, p. 50) who last borrowed the specimen. 
Zahlbruckner cited KOH— as a reaction, but this is not unusual 
because stictic acid is sometimes so scarce that color tests (though not 
microchemical tests) are negative. 

Although Graphina puiggarii var. corumbensis Redgr. is referable 
to G. parilis, the holotype of @. pucggarit var. puiggarii Mill. Arg. 
(Puiggart 506, G) isnot. G. puiggarit has a completely black, almost 
closed exciple of different construction from that found in G@. parilis. 

The type of @. acromelaena was given by Miiller as Pittier 6167, but 
the holotype is labeled Tonduz. According to Dr. C. E. B. Bonner 
(in litt.), “Types which were published with Pittier numbers appear 
to have been collected by Tonduz and numbered after Miller had 
studied the material.”’ 


14. Graphina peplophora sp. nov. FIGuRE 36 


Thallus corticola, continuus, laevis, 70-80, crassus, strato 
corticale plus minusve cellulare, 20-25 u crasso. Apothecia dispersa, 
emergentia vel subsessilia, simplicia, raro furcata, recta vel undulata, 
1-5 mm. longa, 0.5-1 mm. lata, superne albofarinosa, quam thallo 
pallidiore, excipulo dimidiato, rufescente, labiis convergentibus, in 
5-6 striis clavatis divisis; hymenium ellipticum vel rotundatum, 
125-250 » altum. Sporae decolores vel leviter fuscescentes, 35-78 pu 
longae, 18-36 yu latae, loculis horizontalibus 10-16, loculis transversis 
5-8. 

Reactions: KOH-+ red, P—, o-T—, no acids demonstrated. 

Type in the U.S. National Herbarium, collected at Tampico, 
Mexico, July 1896, by C. G. Pringle, no. 408 (isotype in MICH). 

Specimens examined: Mexico. Tamaulipas: Tampico, Pringle 
15460 (FH, MICH). Cuba. Wright 5, 425, 426 (US). Bonin 
Islands, Wright (US). 

Nylander published this species as a nomen nudum under Graphis 
leucocarpa in Flora 69:103, 1886. On the basis of specimens examined 
so far, it is difficult to decide whether this entity should be placed in 
Graphina or Phaeographina. The spores vary from completely 
hyaline to pale brown. Its affinities lie structurally with Graphina, 
particularly with G. corcovadensis Redgr. which differs in having one 
spored asci, ascocarps and thallus concolorous, and exciples closed. 


15. Graphina rimulosa Redgr. Ark. Bot. 26A, no. 1:39. 1933. FIGuRB 37 
Holotype: Rio Grande do Sul, Brazil, Malme 1089 (S). 


Thallus smooth, continuous, rather irregular. Ascocarps emergent, 
flexuose and irregular, rarely branched, 2-4 mm. long, about 0.4 mm. 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 87 


Ficures 36-37.—36, Graphina peplophora Wirth & Hale (holotype, Pringle 408). 37. G. 
rimulosa Redgr. (holotype, Malme 1089), 


wide, generally somewhat clumped in elevated groups, each ascocarp 
surrounded and partially covered by a whitish thalline border; exciple 
closed, very irregular, partially carbonized, partially dark brown; 
labia irregularly striate or disintegrated, partially covered with a 
loose thalline veil; hymenium 120-140 uw high. Spores 6-8 per ascus, 
10-13 * 1-4 locular, 10-15*30-45 u. 

Reactions: KOH-+ reddish, no microchemical tests made. 

Specimen examined. Hidalgo: Honey Station, Pringle 10863 p.p. 
(VT). 

The Pringle collection is nearly identical with the Brazilian holotype. 
Both show the peculiar whitish lateral covering over the ascocarps 
and the highly irregular exciple, features which characterize this 
species. 

16. Graphina scolecitis (Tuck.) Fink, Lich. Flora U.S. 115. 1935. Fiacure 38 
Graphis scolecitis Tuck. Gen. Lich. 210. 1872. Type: Alabama, U.S.A., 
Beaumont (US, isotype). 
Graphina cinerea Fink, Mycologia 19:216. 1927. Holotype: Mayagiiez, 
Puerto Rico, Fink 1163 (MICH). 


Graphina olivobrunnea Fink, Mycologia 19:216. 1927. Holotype: Mayagiiez, 
Puerto Rico, Fink 1224 (MICH). 


Thallus smooth to minutely roughened, continuous. Ascocarps 
emergent to subsessile, usually darker than the thallus but occasionally 
concolorous, slender, unbranched to 2-3 branched, 1-5 mm. long; 
exciple open to nearly closed, red-brown; labia entire, often loosely 
constructed apically; hymenium 50-120 uw high. Spores 8 per ascus, 
4-8 1-3 locular, 5-10 13-29 up. 

Reaction: Thallus KOH—, P—; ascocarps KOH-+ dark purple, 
no microchemical tests made. 

Specimen examined. San Luis Potosi: Las Palmas, Pringle 233 
(FH, H, US, VT). 


88 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Pringle 233 was identified as the New Caledonian Graphina 
adscribens (Nyl.) Mill. Arg. by Eckfeldt (1892, p. 252). However, 
we have not checked this type, and there is little reason to suppose 
that Eckfeldt made a correct determination. The specimens above 
are all referable to G. scolecitis, which was published four years later 
than G. adscribens. The two Fink species differ from typical G. 
scolecitis only in being somewhat smaller. 


17. Graphina sulcata Fink, Mycologia 19:217, 1927. Ficures 39, 40 


Holotype: Rfo Piedras, Puerto Rico, Fink 659 (MICH). 

Thallus smooth to minutely roughened, continuous. Ascocarps 
subsessile to semi-immersed, straight to flexuose, entire or sparingly 
branched, 1-5 mm. long, 0.2-0.5 mm. wide; exciple black above, 
yellow or brownish laterally, open or nearly closed below; labia 3-4 
sulcate, convergent. Spores (2-)4-6 per ascus, 5-82-5 locular, 
10-20 * 20-34 yp. 

Reactions: KOH-+ red, norstictic acid. 

Specimens examined: Chiapas: El Suspiro, Hale 20233, 20068 (US). 

Graphina sulcata differs from the very closely allied G@. parilis in 
having 2-6, not 8, spores per ascus, constantly fewer locules per spore, 
and norstictic rather than stictic acid. 


18. Graphina sulcatula Mill. Arg. var. conglomerata Mill. Arg. Rev. Mycol. 
10:119. 1888. Fiaure 41 


Syntypes: Paraguay, Balansa 4185a (G), 485 (not seen). 

Thallus smooth, very thin. Ascocarps sessile, black, clustered in 
round or irregular clumps, 1-2 mm. in diameter; disc dark brown; 
exciple black, closed; labia convergent, entire to lightly crenate; 
hymenium 75-90 » high. Spores 6-8 per ascus, 6-8X1-4 locular, 
10-13 X30-35 uy. 

Reactions: KOH—, P—, o-T—, no acids demonstrated. 

Specimens examined: Oaxaca: Northwest of Tehuantepec, Hale 
20629 (US). 

The ascocarps of this taxon form distinct isolated clumps which 
are elevated well above the thallus. They are heavily carbonized, 
rather brittle, and difficult to section properly. 

19. Graphina triangularis Zahlbr. Ann. Mycol. 19:232. 1921. Ficure 42 
Type: Tampico, Tamaulipas, Mexico, Pringle 17 (MICH, isotype). 

Thallus smooth, very thick, continuous or lightly fissured. Asco- 
carps more or less immersed, 1-2 branched, flexuose, slender, to 2 mm. 
long; exciple brown, closed, basally continuous in a carbonized band; 
labia lightly striate, slightly convergent; hymenium triangular, 
280-300 » high; epithecium brown, thick. Spores 4-8 per ascus, 
12-15 5-7 locular, 15-25 45-75 uy. 

Reactions: KOH-+ red, unknown acid in G.A. o-T. 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 89 


Ficures 39-40.—39. Graphina sulcata Fink (holotype, Fink 659). 40. G. sulcata (Hale 
20068). 


Ficures 41-42 —41. Graphina sulcatula var. conglomerata Mull. Arg. (Hale 20629). 42. G. 
triangularis Zahlbr. (isotype, Pringle 17). 


Graphina triangularis is a distinct species, unlikely to be confused 
with any other. It is somewhat similar internally to G aibonitensis 
Fink, in that the labia consist of elongate striae arising from a thick- 
ened base. The unknown acid is also found in Graphina palmeri, 
G. virginea, and Phaeographis exaltata. 


90 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


20. Graphina virginalis (Ny].) Mill. Arg. Bull. Herb. Boiss. 3:47. 1895. 
FIGURE 43 

Fissurina virginalis Nyl. Lich. Insul. Guin. 50. 1889. Type: Florida, U.S.A. 
(US, isotype). 

Graphis virginalis Tuck. in Eckf. Bull. Torr. Bot. Club 17:256. 1890. 
Based on Fissurina virginalis Nyl. 

Graphina virginalis (Tuck.) Redgr. Ark. Bot. 26A, no. 1:58. 1933. Super- 
fluous combination. 

Thallus smooth, continuous. Ascocarps dispersed to somewhat 
clustered, usually lighter than the thallus, immersed to emergent, 
sparingly branched, straight to rather flexuose, to 3 mm. long; disc 
appearing first as a light colored line, then often becoming elevated 
and fissurine; exciple open or nearly so, colorless laterally; labia 
convergent, light to dark brown; hymenium about 100 » high. Spores 
8 per ascus, elliptic, often halonate, irregularly few celled, 14-18 
26-40 uy. 

Reaction: KOH-+ red, P—, no microchemical test made. 

Specimen examined: Chiapas: El Suspiro, Hale 20225 (US). 

Graphina virginalis is part of the nitida-leuconephela complex, a 
group in need of careful revision. 

21. Graphina virginea (Eschw. in Mart.) Mill. Arg. Flora 63:41. 1890. 
Figure 44 
Leiogramma virgineum Eschw. in Mart. Fl. Bras. 1:98. 1833. Type: Near 
Par&, Brazil, s.c. (not seen and possibly destroyed). 

Thallus thick, smooth to minutely granular, continuous or lightly 
fissured. Ascocarps whitish, immersed, usually asteroidly branched 
and intricate, 1-4 mm. long; exciple pale, rudimentary, open; labia 
upright, more or less separate. Spores 2-8 per ascus, elliptical or 
elongate, 15-25 3-7 locular, 13-21 50-130 u. 

Reaction: KOH-+ red, unknown acid in G.A. o-T. 

Specimens examined: Vera Cruz: Northeast of San Andrés Tuxtla, 
Hale 19792, northwest of Alvarado, Hale 19756, Orizaba, Fr. Miiller 
(US). 


TH 
ad Sti iT} ee: 
CREED 6 


Ficures 43-44.—43. Graphina virginalis (Nyl.) Mull. Arg. (isotype). 44. G. virginea 
(Eschw. in Mart.) Mull. Arg. (Fr. Muller s.n.). 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 9] 


The Miiller collection from Orizaba was identified as Graphis cometia 
Fée by Nylander (1858, p. 381), a species considered to be a synonym 
of G. virginea. 

Graphina virginea seems to vary in the number of spores per ascus. 
Some specimens have two to three per ascus, others four to six, others 
six to eight, with no apparent correlations with anatomy or chemistry. 
The unknown acid is identical with that found in Phaeographis ezaltata, 
Graphina palmer, and G@. triangularis. 


2. Graphis 


22. Graphis afzelii Ach. Syn. Lich. 85. 1814. Fiaures 45, 46 
Holotype: Guinea, Afzelius (H). Isotype in UPS. 
Graphis atroalba Vain. Acta Soc. Faun. Fl. Fenn. 7, no. 7:123. 1890, non 
Kremplh. 1875. Holotype: Rio de Janeiro, Brazil, Vainio 189 (TUR). 
Graphis atroleuca Zahlbr. Cat. Lich. Univ. 2:294. 1923. Based on 
G. atroalba Vain. 


Thallus smooth, continuous. Ascocarps sessile, unbranched, usually 
completely covered by a white thalloid veil, 1-5 mm. long to 1 mm. 
wide; exciple closed, black and thick above, brown or pale below, often 
quite thin; labia usually entire, convergent; hymenium 75-125 u high. 
Spores 8 per ascus, four locular, 6-9 16-23 u. 

Reactions: Ascocarps C+ red, lecanoric acid present. 

Specimen examined: Chiapas: Road to El Suspiro, Hale 20099 
(S, TNS, US). 

The holotype of Graphis atroalba Vain. is only a depauperate speci- 
men of @. afzelvi, lacking most of the white veil which normally covers 
the ascocarps. However, a portion which remains reacts C+ red, 
and in all other details, the taxa are identical. 

23. Graphis anguilliformis Tayl. Lond. Journ. Bot. 6:152. 1847. Ficure 47 

Holotype: St. Vincent’s, West Indies (FH-Tay]). 


Ficures 45-46.—45. Graphis afzelii Ach. (isotype). 46. G. afzelii (Hale 20099). 


92 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Thallus smooth to nodular, continuous. Ascocarps sessile, simple 
to 3-4 branched, 1-5 mm. long, 0.6-0.8 mm. wide, concolorous with 
the thallus except at the very apex which is usually black; exciple 
black, entire, covered almost to the apex by a thalline veil; labia 
entire, convergent; hymenium 175-200 » high. Spores 2-6 per ascus, 
12-18 locular, 10-13 75-100 u long. 

Reactions: KOH-+ red, P—, 0-T—, no acids demonstrated. 

Specimen examined: Hidalgo: Honey Station, Pringle 10864 (MICH, 
VT). 

Graphis anguilliformis is usually considered as a synonym of the 
Brazilian G. illinata Eschw., the type of which we have not been able 
to find. The Pringle material from Mexico is identical in all ways 
with Taylor’s type, and it is perhaps best to continue using this name, 
which has been typified. 

24. Graphis caesiella Vain, Acta Soc. Faun. Fl. Fenn. 7, no. 7:122. 1890. 
Fiaures 48-50 
Holotype: Rio de Janeiro, Brazil, Vainto 45 (TUR). 
Graphis yaucoensis Fink, Mycologia 19:213. 1927. Holotype: Yauco, 
Puerto Rico, Fink 1691 (MICH). 

Thallus continuous, smooth to minutely roughened. Ascocarps 
distinctly pruinose, immersed, slender, simple to branched and 
subintricate, often bordered by an irregular slightly raised thalline 
margin, 0.5-4 mm. long; exciple black, open; labia entire, divergent or 
slightly convergent; hymenium 75-125 » high. Spores 8 per ascus, 
6-10 locular, 6-10 20-40 uy. 

Reactions: KOH-+ red, norstictic acid. 

Specimen examined: Tamaulipas: Tampico, Pringle 62 (FH, 
MICH). 

Graphis caesiella can be separated from the other species in the 
G. scripta complex by the distinctly pruinose ascocarps. It is quite 
closely related to G. subamylacea Zahlbr., which differs only in having 
a closed exciple. Graphis yaucoensis is identical in all respects with 
the holotype of @. caesiella. 

Chemically G. caesiella is somewhat variable outside of Mexico. 
The majority of 48 specimens tested contained norstictic acid, but a 
significant number yielded stictic or protocetraric acids. Thirty-seven 
specimens with norstictic acid were collected in Louisiana, Mexico, 
Puerto Rico, Hispaniola, Brazil, and Samoa. Seven specimens with 
protocetraric acid were from Florida, Cuba, Hispaniola, and Puerto 
Rico. Four with stictic acid were restricted to the Dominican Re- 
public. There seems to be no correlation between the chemistry and 
distribution on the basis of this small sample. 


25. Graphis desquamescens Fée, Bull. Soc. Bot. France 21:24. 1876. 
Figure 51 
Syntypes: Brazil, Glaziow 5082 (M), 5016 (see below). 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 93 


Graphis compulsa Kremplh. Flora 59:419, 1876. Based on G. desquames- 
cens Fée. 


Thallus continuous, minutely roughened to smooth. Ascocarps 
subsessile, black, Opegrapha-like, slender, often branched, flexuose, 
0.5-3 mm. long, often with a low thalline margin; exciple closed, black; 
labia convergent, entire or very slightly crenate; hymenium 75 yu high. 
Spores 8 per ascus, 6-8 locular, 5-8 20-28 yp. 

Reactions: KOH-+ red, norstictic acid. 

Specimen examined: Vera Cruz: East of Cérdoba, Hale 19731 (US). 

Graphis compulsa Kremplh. is a superfluous new name for G@. des- 
quamescens Fée and was based on Glaziou 5082. This specimen (M) 
has norstictic acid and is identical with material collected from many 
localities throughout the American tropics. Unfortunately we have 
yet to examine the other syntype of this species. 


nono 


4 
OW 


LDP LL 
EG GG 
“ 50 


Ficures 47-50.—47. Graphis anguilliformis Tayl. (holotype). 48. G. caesiella Vain. (holo- 
type, Vainio 45). 49. G. yaucoensis Fink (holotype, Fink 1691). 50. G. caesiella Vain. 
(Pringle 62). 


Ficure 51.—Graphis compulsa Kremplh. (holotype, Glaziou 5082). 
686—879—63-—_3 


94 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


26. Graphis endoxantha Nyl. Bull. Soc. Linn. Norm. ser. 2, 2:110. 1868. 
Figures 52, 53 
Graphis subelegans Nyl. Lich. Trop. Labuan Singapore 42. 1891. Holotype: 
San Luis Potos{f, Mexico, Pringle 162 (H). 

Holotype: New Caledonia, Pancher (H). 

Thallus smooth, nitid, continuous. Ascocarps sessile, prominent, 
flexuose, rarely branched, 1-6 mm. long, 0.5-0.7 mm. wide; exciple 
nearly open to closed, usually much thickened below, yellow, with 
many prominent crystalline inclusions; labia with many striae, black; 
hymenium about 100 » high. Spores 8 per ascus, 6-7 locular, 6-8X 
25-30 uy. 

Reactions: KOH+ reddish, P—, no microchemical tests made. 

Specimens examined: San Luis Potosi: Las Palmas, Pringle 217 
(VT), 166 p.p. (FH). 

The holotype of G. endoxantha is sterile but it is unquestionably 
identical with @. subelegans. The species is closely allied to G. proser- 
pens Vain., differing in the nitid thallus, the very prominent ascocarps, 
and the abundant excipular inclusions, 


27. Graphis flexibilis Kremplh. Flora 59:414. 1876. Fiaure 54 


Holotype: Brazil, Glaziou 5106 (M). 

Thallus smooth to warty or fissured. Ascocarps sessile, black 
above, commonly branched, flexuose, acute, 5-13 mm. long, about 
0.6 mm. wide; exciple black, closed, often thickened below; labia 
convergent, 1-6 sulcate, often with included crystals; hymenium 
50-150 whigh. Spores 6-8 per ascus, 15-21 locular, 12-13 60-105 uy. 

Reaction: KOH-+ reddish, P—, o-T—, G.E.— no acids demon- 
strated. 

Specimen examined: Chiapas: West of Tuxtla Gutiérrez, Hale 
19894 (US). 

Zahlbruckner (1923, p. 293) placed Graphis flexibilis as a synonym 
of G. angustata Eschw. in Mart. As a rule, Eschweiler’s types are 
not available for study to our knowledge, and since in this case the 
species cannot be typified, it seems appropriate to use Krempelhuber’s 
name. 


28. Graphis glaucopis sp. nov. Figure 55 


Thallus corticola, continuus, laevis vel nitidulus, 200-250 » crassus, 
strato corticale arachnoideo, 25-30 u crasso. Apothecia subsessilia, 
elliptica vel oblongo-elliptica, plus minusve simplicia, 0.5-2 mm. 
longa, 0.4 mm. lata, albofarinosa, superne fuscescentia, excipulo plus 
minusve integro, pallido, labiis convergentibus, integris, fuscofuli- 
gineis; hymenium 150-200 4 altum, epithecio indictincto. Sporae 
6nae, loculis 25-27, 16-26 < 75-260 u longae. 

Reactions: Thallus KOH-+ orange, P—; ascocarps KOH-+ orange, 
P+ red, o-T—, G.E.—, no acids demonstrated. 


WIRTH & HALE—-MEXICAN GRAPHIDACEAE 95 


Ficures 52-53.—52. Graphis subelegans Nyl. (holotype, Pringle 162). 53. G. endoxantha 
Nyl. (Pringle 166 p.p.). 


200 00DHNDH00020000% 


WU 
equ 


Ficures 54, 55.—54. Graphis flexibilis Kremplh. (holotype, Glaziou 5106). 55. G. glaucopss 
Wirth & Hale (holotype, Hale 20219). 


Type in the U.S. National Herbarium, collected in Mexico, El 
Suspiro, Chiapas, March 22, 1960, by M. E. Hale (no. 20219) (iso- 
types in S, TNS). 

Graphis glaucopis is somewhat difficult to place as to genus. The 
extremely thick thallus and the heavy walled spores indicate affini- 
ties with Ocellularia. However, the distinctly elliptical ascocarps, 
the rimiform disc, and the lack of a columella make it necessary to 
put the species in Graphis. 

This species is unusual in the extreme variation in spore size to be 
found within a single ascocarp. Spore length is a very variable charac- 
ter in the majority of the species of the Graphidaceae, and @. glau- 
copis would seem to represent the extreme of this tendency. 


29. Graphis grammatica Ny]. Flora 49:292, 1866. Fieure 56 
Type: Cuba (see below). 


96 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Thallus very thick, often fissured and uneven. Ascocarps densely 
clustered, immersed to depressed, curved and intricate, to 1 mm. 
long, very slender; disc light brown; exciple pale brown, closed, 150- 
200 uw high, heavily thickened below the hymenium. Spores 8 per 
ascus, 4 locular, 4-6 10-18 uy. 

Reactions: KOH-+ yellow, stictic acid. 

Specimen examined: Chiapas: El Sumidero, Tuxtla Gutiérrez, Hale 
20042 (US). 

Graphis grammatica was originally described without citation of 
collector or number; “Cuba” is the only indication of provenance. 
Among the Wright Cuban collections, no. 36 is labeled G. grammatica 
Nyl., and these specimens (FH, US) are probably isotypes. They 
compare very well with the Mexican material (which is sterile), except 
that Wright 36 yielded no stictic acid. 


30. Graphis implicata Fée, Bull. Soc. Bot. France 21:27. 1874. Figures 57-59 
Graphis chlorocarpella Ny]. in Kremplh. Flora 59:413. 1876. Based on G. 


implicata Fée. 


Graphis balbisina Nyl. Lich. Trop. Singapore et Labuan 42. 1891. Type: 
Mexico (see below). 


Type: Brazil, Glaztow 5036 (MICH, isotype). Reactions: too frag- 
mentary to test. 

Thallus continuous or fissured, often uneven. Ascocarps semi- 
emergent to subsessile, lighter than the thallus, occasionally branched, 
flexuose and somewhat intricate; exciple closed or nearly so, reddish or 
yellowish brown; labia lightly sulcate, convergent, often with dark- 
ened apices. Spores 8 per ascus, 11-21 locular, 11-13 52-107 p. 

Reactions: KOH-+ red, norstictic acid. 

Specimens examined: Nuevo Leén: Monterrey, Pringle 118, 119 
(FH, US, VT). 

Both Graphis ‘mplicata and @. chlorocarpella are based on Glaziou 
5036, and are therefore synonymous, as has already been pointed out 
by Zahlbruckner (1909, p. 108). The isotype in Fink’s herbarium is 
too fragmentary to test; Pringle 119 yielded an extremely small 
amount of norstictic acid. 

Graphis balbisina Nyl. was described from Mexico with Eckfeldt 
as the collector. One year later Eckfeldt (1892, p. 253) listed this 
species from Mexico, under Pringle’s collection 118. It may therefore 
be assumed that this was the specimen on which Nylander based his 
description and should be considered as the type (isotypes in FH, 
US, VT). Nylander’s dealings with Eckfeldt are frequently marked 
by confusion. 


31. Graphis longula Kremplh. Flora 59:414. 1876. Figure 60 


Phaeographis longula (Kremplh.) Zahlbr. Denkschr. Akad. Wiss. Wien 
83:109. 1909. 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 97 


Ficures 56-59.—56, Graphis grammatica Ny]. (Wright 36). 57. G. implicata Fée (isotype, 
Glaziou 5036). 58. G. balbisina Nyl. (isotype, Pringle 118). 59. G. implicata Fée 
(Pringle 119). 


Holotype: Brazil, Glaziou 5497 (M). 

Thallus smooth to minutely roughened, continuous. Ascocarps 
black above, slender, subsessile, elongate, quite flexuose, occasionally 
branched, 2-10 mm. long, often with a low thalline margin; exciple 
black, closed; labia convergent, crenate to dentate; hymenium 80-90 u 
high. Spores 2-6 per ascus, colorless, 10-17 locular, 9-13 X (40)75-90 u. 

Reactions: KOH— or KOH+ yellow, P—, no acids demonstrated. 


Z0GG00000000003 


Ficures 60, 61.—60. Graphis longula Kremplh. (holotype, Glaziou 5497). 61. G. platycar- 
pella Mill. Arg. (Pringle 210). 


98 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Specimen examined: Chiapas: North of Berriozibal, Hale 20225 
(US). 

The spores of Graphis longula are quite colorless at maturity, and 
Zahlbruckner’s transfer to Phaeographis is unnecessary. It is possible, 
however, that this species may be referable to Graphis angustata 
Eschw. in Mart. 


32. Graphis platycarpella Mill. Arg. Bull. Soc. Bot. Belg. 30:81. 1891. 
Ficure 61 


Type: Baie de Salinas, Costa Rica, Pittier 5236 (not seen). 

Thallus smooth, continuous, fairly thick. Ascocarp immersed, 
round to lirelline, usually unbranched and aggregated, 1-3 mm. long, 
0.5-1 mm. wide; thalline margin prominent, white, mealy, erect and 
flaring, exposing a flat irregular “disc” (actually the excipular labia) ; 
exciple open or partially closed, pale and thin laterally; labia thick, 
light to dark brown, entire, convergent; hymenium 100-150 u high. 
Spores usually 4-8 per ascus, 4 locular, 6-12 12-26 up. 

Reactions: KOH—, P—, o-T—, no acids demonstrated. 

Specimen examined: San Luis Potosi: Las Palmas, Pringle 210 
(FH, VT). 

The Pringle material was reported as “Graphis lactea Ny.” 
[= Graphis lactea (Fée) Sprengl.] by Eckfeldt (1892, p. 252). However, 
a fragmentary isotype of Fissurina lactea Fée (H) differs internally 
from the Mexican material. The description of Graphis platycarpella 
is more in accord with Pringle 210 and this specimen is tentatively 
identified as such. This entity is part of the complex series of Graphis 
species which are characterized by 4 locular spores, pale exciples, and 
more or less fissurine ascocarps. Such species as Graphis durandi 
Mill. Arg., G. grossula Mill. Arg., @. humilis Vain., @. lecanorina 
Mill. Arg., G. schizogramma Vain., and @. stromatoides Magn. should 
be carefully compared in an effort to determine reliable species criteria 
in this alliance. This group should also be examined with regard 
to its relationship with the Graphina nitida-leuconephela-mexicana 
complex. 

33. Graphis proserpens Vain. Bot. Tidsskr. 29:132. 1909. Figures 62, 63 

Graphis disserpens Vain. Acta Soc. Faun. FI. Fenn. 7, no. 7:123. 1890, non 

G. disserpens Nyl. Holotype: Sitio, Minas Gerais, Brazil, Vainio 1091 
(TUR). 

Thallus smooth, continuous. Ascocarps unbranched, subsessile, 
1-4 mm. long, 0.2-0.5 mm. wide, black, occasionally with a small 
lateral thalline margin; exciple black above, red-brown laterally, 
suffused below, open or nearly closed; labia convergent, 4-7 striate; 
hymenium 50-60 » high. Spores 6-8 per ascus, 7-9 locular, 7-8X 
18-28 uw. 

Reactions: KOH+ red, P—, o-T—, no acids demonstrated. 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 99 


Ficures 62, 63.—62. Graphis proserpens Vain. (holotype, Vainio 1091). 63. G. proserpens 
(Pringle 15423). 


Specimens examined: Morelos: Near Cuernavaca, Pringle 15380 
(FH), 15423 (FH, VT). 

The name Graphis disserpens Vain. was apparently a lapsus calamt, 
as Vainio mentioned G. disserpens Nyl. in his original description. 
Vainio himself proposed a new name in 1909. 

Graphis proserpens is part of the rimulosa-elegans-duplicata-striatula 
complex. It is possible that further study may prove these entities 
to be synonymous. Separating them on excipular characters seems 
rather difficult at the present time. 


34. Graphis stygioarachnoideay sp. nov. Fiaure 64 


Thallus corticola, continuus, 38-42, crassus, strato corticale 
arachnoideo, 13-15 wcrasso. Apothecia dispersa, subsessilia, flexuosa, 
lirellina, raro furcata, 2-5 mm. longa, 0.3-0.4 mm. lata, quam thallo 
pallidiore, excipulo plus minusve integro, pallido, labiis convergentibus, 
fuscescentibus; hymenium 100-120 » altum. Sporae 8nae, 5.5-8X 
22-42 uw, loculis 11-16. 


Ficure 64.—Graphis stygioarachnoidea Wirth & Hale (holotype, Hale 20399). 


Reactions: KOH-+ reddish, P+, o-T—, no acids demonstrated. 
Type in the U.S. National Herbarium, collected in Mexico, Lagos 
de Monte Bello, Chiapas, March 25, 1960, by M. E. Hale (no. 20399). 
This new species is quite closely related to G. mosquitensis Tuck., 
from which it differs in having the hyphae of the labia free and 


100 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


distinctly carbonized, and in the more robust, nearly sessile ascocarps. 
Although the labia in the holotype of @. mosquitensis occasionally 
show an apical looseness of construction, these hyphae never become 
free or distinctly black and carbonized. 


35. Graphis subamylacea Zahlbr. Ann. Mycol. 19:229. 1921. Figure 65 


Type: Cuernavaca, Morelos, Mexico, Pringle 24 (MICH, isotype). 

Thallus continuous, to 100 « thick, smooth or minutely roughened. 
Ascocarps immersed, very variable, slender, much branched, often 
radiate or anastomosing, 1-5 mm. long, about 0.3 mm. wide, level 
with the thallus or sometimes bordered by very low thalline margins, 
lightly pruinose or black; exciple black, usually closed; labia entire, 
divergent; hymenium 100-120 4 high, opaque. Spores 8 per ascus, 
6-8 locular, 5-8 17-30 uy. 


I, tray | 
Su td] ) 


dé 


1 


———— 


Ficures 65, 66.—65. Graphis subamylacea Zahlbr. (isotype). 66. Melaspilea polymorpha 
Mull. Arg. (holotype, Pringle 98). 


Reactions: KOH+ yellow, stictic acid. 

Specimen examined: Tamaulipas: Tampico, Pringle 412 (MICH). 

This species, though fairly distinct, is extremely variable in the 
form, size, and pruinosity of the ascocarps, the isotype has ascocarps 
ranging from quite small and highly pruinose to large and almost 
black. The second collection (Pringle 412) is more robust, and 
externally much like Graphis caesiella Vain. 


3. Melaspilea 


36. Melaspilea polymorpha Miill. Arg. Bull. Herb. Boiss. 2:92. 1894. 
FIGurE 66 

Holotype: Near Monterrey, Mexico, Pringle 98 (G). 

Thallus thin, smooth. Ascocarps sessile, unbranched, slightly 
flexuose, to 1 mm. long; exciple black, closed; labia entire, convergent 
but exposing a fairly wide disc; hymenium about 60 u high. Spores 
2 locular, brown, 4-8 per ascus, 5-7 14-17 yp. 

Reactions: KOH—, P—, no acids demonstrated. 


WIRTH & HALE—MEXICAN GRAPHIDACEAR 101 


4. Phaeographina 


37. Phaeographina asteroides Fink, Mycologia 19:218. 1927. FiaurEs 67, 68 
Phaeographina caesiopruinosella Fink, Mycologia 19:219. 1927. Holotype: 
Naranjito, Puerto Rico, Fink 140 (MICH). 

Holotype: Mayagiiez, Puerto Rico, Fink 981 (MICH). 

Thallus smooth, continuous. Ascocarps usually much branched, 
1-3 mm. long, 0.3-0.7 mm. wide, emergent to subsessile; thalline 
margin more or less prominent, often somewhat lighter than the 
thallus; disc (and often the whole ascocarp) pruinose; exciple brownish, 
closed; labia entire, more or less divergent. Spores 8 per ascus, 
brown, 5-10 2-3 locular, 8-18 25-50 p. 

Reactions: KOH-+ red, P—, o-T—, no acids demonstrated. 

Specimen examined: San Luis Potosi; Las Palmas, Pringle 224 


(FH, VT). 


Q ) 
B05 8000 


Ficures 67, 68.—67. Phaeographina asteroides Fink (holotype, Fink 981). 68. P. caesio- 
pruinosella Fink (holotype, Fink 140). 


Pringle 224 was reported by Miiller (1894, p. 92) as Graphina 
caestoradians (Leight.) Mill. Arg. [=Phaeographina caesioradians 
(Leight.) Redgr.]. Redinger (1933, p. 99) placed both of Fink’s 
species in synonymy of P. caesioradians, but since the type of the 
Leighton species has not been examined and since Redinger has incor- 
rectly handled other Fink species, this synonymy cannot be verified. 
In any event, the two Puerto Rican species are identical (except for a 
slight difference in spore size), and the Pringle material is within the 
range of variation shown in the Puerto Rican types. 

38. Phaeographina caesiopruinosa (Fée) Miill. Arg. Mem. Soc. Phys. Hist. Nat. 

Genéve 29, no. 8:49. 1887. FIGureE 70 

Arthonia caesiopruinosa Fée, Suppl. Essai Crypt. Ecorces 36, pl. 40, fig. 4, 
1837. Type: Tropical America (not seen). 

Thallus smooth to slightly roughened, continuous. Ascocarps 
large, emergent to subsessile, elongate, to nearly round, occasionally 
branched, 1-10 mm. long, 1-3 mm. wide, with a wide, usually promi- 
nent, elevated thalline margin; disc flat, lightly to heavily pruinose; 


102 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


exciple black, open to nearly closed, usually black laterally and absent 
below; hymenium 100-250 yu high, clear or inspersed. Spores 8 per 
ascus, gray to brown, 10-20X2-6 locular, 15-20 45-120 u. 

Reactions: KOH-+ reddish or yellow. 

Specimens examined: Chiapas: El Suspiro, Hale 20133, 20215; west 
of Tuxtla Gutiérrez, Hale 19903 (US). Nuevo Leén: Monterrey, 
Pringle 113 (FH, VT, US). Vera Cruz: North of Fortin de las Flores, 
Hale 19695; north of Huatusco, Hale 19482 (US). 

Examination of a large number of specimens from the southern 
United States and tropical America referable to this common species 
indicated an intergrading series of excipular forms, varying from com- 
pletely dimidiate to distinctly carbonized below. There is also inter- 
gradation between heavily inspersed hymenia and clear hymenia, 
elongate nearly unbranched ascocarps and stout lobulate ascocarps. 
None of these characters seem to show any mutual interrelation. For 
the moment, it seems best to treat the aggregate as a single widespread 
variable species. 

39. Phacographina chrysocarpa (Raddi) Redgr. Ark. Bot. 26A, no. 1:83. 1933. 
FIGURE 69 
Opegrapha chrysocarpa Raddi, Atti Soe. Ital. Sci. 344. 1820. Type: Mandioca, 
Brazil (not seen). 
Graphis chrysocarpa (Raddi) Sprengl. Syst. Veg. 4:253. 1827. 
Graphina chrysocarpa (Raddi) Miill. Arg. Flora 63:41. 1880. 
Graphis miniata Redgr. Ark. Bot. 27A, no. 3:26. 1935. Syntypes: Brazil, 
Mosén 3145, 3313, 3577 (S). 

Thallus smooth, continuous. Ascocarps subsessile to sessile, un- 
branched to asteroidly branched, straight to quite curved, 1-7 mm. 
long, more or less covered by a cinnabar-red thalline veil, black 
beneath; exciple black, closed; labia convergent, striate. Spores 4-6 
per ascus, colorless, brown, or gray, transversely septate or bi-ocellate 
at the ends to completely muriform, 10-15 65-150 » long. 

Reactions: Thallus KOH—, P—, no acids demonstrated ; ascocarps 
KOH- purple. 

Specimen examined: Chiapas: El Suspiro, Hale 20173 (S, TNS, 
US). 

Phaeographina chrysocarpa is distinguished by its brilliantly colored 
cinnabar ascocarps. The large series of specimens examined from 
tropical America showed great variation in spore septation and color. 
In fact, it was possible to find specimens with cinnabar ascocarps and 
black closed exciples which could be referred to Graphis, Graphina 
Phaeographis, or Phaeographina. In some cases a single specimen 
could be assigned to all four genera. As shown in the list of syno- 
nyms, this entity has already been placed in three of these genera. 

The variation in spore characters throws considerable doubt on the 
validity of the four traditional genera. Specimens are usually identi- 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 103 


Ficures 69, 70.—69. Phaeographina chrysocarpa (Raddi) Redgr. (Hale 20173). 70. P. 
caesiopruninosa (Fée) Mill. Arg. (Pringle 113). 


fied as P. chrysocarpa on the basis of the cinnabarine ascocarps and 
entire exciples, regardless of variation in the spores. Is it not possible 
that other species, especially those lacking such an obvious unifying 
feature as a pigmented ascocarp, may be assigned to several different 
genera, when in fact they are simply a single species with polymorphic 
spores? One may suggest as an example the mass of species which 
cluster about Graphina acharit. 

It should also be noted that the cinnabarine ascocarp covering in 
P. chrysocarpa sometimes becomes quite reduced, occurring only at 
the ends of the lirellae. If this pigmented covering disappeared 
completely, the resulting species would be very close to Graphina 
acharii, ® species which shows similar spore variation. The im- 
plication of this is obvious. 

One of the syntypes of Graphis miniata Redgr. has brown, muriform 
spores, and it is therefore included in synonymy here. Actually the 
structure of the ascocarp in this species as illustrated by Redinger is 
inaccurate; internally G. miniata is exactly like typical Phaeographina 
chrysocarpa. 

One other species in the Graphidaceae occurs with bright cinnabar 
ascocarps, Graphina malmei Redgr. The holotype (Brazil, Malme 
3508, S) is indeed distinct, as the exciple is pallid below, and the labia 
constructed of free lamellae. A second collection from Brazil (Weir 
6223, BPI) confirms the validity of this species. 

40. Phaeographina sp. FIGuRE 71 
Phaeographina scalpturata auct. non (Ach.) Mill. Arg. 

Thallus continuous, smooth to roughened. Ascocarps sessile, very 
large, sparingly branched, acute, straight to nearly intricate, to 15 
mm. long and nearly 2 mm. wide, with a prominent elevated, light- 
colored thalline margin; disc flat, caesiopruinose; exciple carbonized, 
dimidiate or sometimes barely closed below. Spores 1 per ascus, 
brownish, 20-35 150-200 yu, densely muriform. 


104 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Reactions: KOH-+ red, norstictic acid. 

Specimens examined: Chiapas: West of Tuxtla Gutiérrez, Hale 
19898 (US). San Luis Potosi: Canoas, Pringle 305 (VT). Vera 
Cruz: Northeast of Huatusco, Hale 19453; north of Fortin de las 
Flores, Hale 19696, 19704 (US). 

An examination of an isotype of P. scalpturata (Ach.) Mill. Arg. 
(see p.110) disclosed that the traditional concept of the species is in- 
correct. The entity described above, often identified as P. scalpturata 
by such workers as Nylander, Miiller, and Redinger, differs consid- 
erably from authentic material of that species. The present entity 
has enormous ascocarps, carbonized exciples, prominent thalline 
margins, and very large spores; it is extremely widespread and com- 
mon and must certainly have a valid name, although we have not 
yet been able to determine what it might be. 


41. Phaeographina strigops sp. nov. FIaurRE 72 


Thallus corticola, continuus, nitidus, 100-170» crassus, strato 
corticale arachnoideo, 13-15 » crasso. Apothecia elongata, flexuosa 
et intricata, 4-5 ramulosa, 1-5 mm. longa, 0.1-0.3 mm. lata, disco 
dilatato, concavo, excipulo integro, fuscorufescente, tenue, labiis in- 
teeris, divergentibus; hymenium semicirculare, 90-100» altum. 
Sporae Snae, fuscescentes, 9-13 X 18-21 yu, loculis horizontalibus 5-6, 
loculis transversis 1-2. 

Reactions: KOH-+ yellow, P—, 0o-T unknown acid. 

Type in the U.S. National Herbarium, collected in Mexico, Lagos 
de Monte Bello, Chiapas, March 25, 1960, by M. E. Hale (no. 20383). 

This new species is allied to the Brazilian P. oxalifera Redgr., from 
which it differs in having a black disc, elongate, intricate ascocarps, 
and an unknown acid instead of norstictic acid. 


42. Phaeographina elliptica, sp, nov. FIGuRE 73 


Thallus corticola, continuus vel fissus, laevis, 100-150 yw crassus, 
strato corticale subcellulare, 60-75 yw crasso. Apothecia immersa, 
simplicia, vulgo recta, elliptica vel sublirellina, apicibus rotundatis, 
0.75-2 mm. longa, 0.3-0.5 mm. lata, disco planato vel concavo, nigro 
vel leviter pruinoso, labiis integris, divergentibus, tenuibus; hymenium 
inspersum, crystallis magnis decoloratis includentibus, 120-150 u 
altum, epithecio fusco. Sporae fuscescentes, Inae, 23-26 90-110 4g, 
loculis horizontalibus 22-25, loculis transversis 2-6. 

Reactions: KOH+ orange, P—, unknown crystals in o-T. 

Type in the U.S. National Herbarium, collected in Mexico, north 
of Berrioz4bal, Chiapas, March 22, 1960, by M. E. Hale (no. 20088). 

Phacographina elliptica is closely related to P. scalpturata (Ach.) 
Mill. Arg., differing primarily in the peculiar crystalline inclusions in 
the hymenium. In addition, the thallus shows considerable fissuring, 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 105 


Ficures 72, 73.—72. Phacographina strigops Wirth & Hale (holotype, Hale 20383). 73. 
P. elliptica Wirth & Hale (holotype, Hale 20088). 


and the ascocarps are more crowded and elliptical than in typical 
P. scalpturata. 


5. Phaeographis 


43. Phaeographis dendritica (Ach.) Mill. Arg. Flora 65:382. 1882. Ficure 74 
Opegrapha dendritica Ach. Meth. Lich. 31. 1803. Type: Southern Spain 
(UPS, isotype). 

Thallus smooth, continuous. Ascocarps semiemergent, often 
desquamescent, simple to asteroidly branched, acute, surrounded by a 
low, narrow thalline border, 1-3 mm. long, about 0.4 mm. wide; 
disc black, flat; exciple dark brown to black, closed; labia entire, 
spreading; hymenium 80-120 » high. Spores 8 per ascus, brown, 
6-8 locular, 8-11 26-42 up. 


Ficure 74.—Phaeographis dendritica (Ach.) Mill. Arg. (isotype). 


Reactions: KOH-+ red, norstictic acid. 

Specimens examined: Chiapas: West of Tuxtla Gutiérrez, Hale 
19895 (US). Vera Cruz: Orizaba, Fr. Miller (US). 

The Mexican specimens are identical in all respects with the isotype. 
It is altogether possible that the great variability attributed to 


106 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


P. dendritica may be incorrect and that the numerous subspecific taxa 
described represent other species. 
44. Phaeographis exaltata (Mont. et v.d. Bosch) Mill. Arg. Flora 65:336. 1882. 
Figure 75 
Lecanactis exaltata Mont. et v. d. Bosch, Pl. Junghuhn, 4:475. 1855. Type: 
Java, Junghuhn (not seen). 

Thallus continuous, smooth to warty. Ascocarps sessile, sparingly 
branched or unbranched, rotund to lirelline, stout, 0.5-4 mm. long, 
0.5-1 mm. wide, often apically rounded, laterally with a very promi- 
nent thalline margin much as in Sarcographa; disc black, wide; exciple 
black, closed, thickened below; labia entire, divergent; hymenium 
inspersed with brown oil droplets, 120-150 wu high. Spores 8 per 
ascus, 6-10 locular, 8-11 23-32 u. 

Reactions: KOH+ yellow, unknown acid in o-T. 

Specimen examined: Chiapas: Lagos de Monte Bello, Hale 
20394 (US). 

This species is one of the numerous transition species between 
Sarcographa and Phaeographis and could easily be placed in either 
genus. The unknown lichen acid is identical with that found in 
Graphina virginea, G. triangularis, etc. 

45. Phaeographis sericea (Eschw. in Mart.) Mill. Arg. Flora 71:523. 1888. 

Figure 77 

Leiogramma sericeum Eschw. in Mart. Icon. Pl. Crypt. 2:34. 1828. Type: 
Brazil (not seen). 

Thallus thin, more or less continuous, smooth. Ascocarps black, 
often very variable, some lirellae asteroidly branched, immersed, 
narrow, apically acute, with a mealy, prominent thalline margin, disc 
black, other lirellae subsessile, lacking a mealy margin, irregularly 
branched and intricate, apically rounded, disc lightly pruinose, to 
0.5 mm. wide; exciple black, closed, thickened below; labia at first 
convergent, then divergent, entire; hymenium 80-90 » high. Spores 
8 per ascus, 4 locular, 7-8 X 18-21 u. 

Reactions: KOH+ yellow, P—, o-T—, no acids demonstrated. 

Specimens examined: Chiapas: North of Berriozabal, Hale 20107 
(S, US). Vera Cruz: East of Cérdoba, Hale 19727 (US). 

The variation in ascocarp shape in the cited specimens is extremely 
instructive. Most clusters appear to originate as immersed, radiately 
branched, acute ascocarps with prominent mealy margins and black 
discs. Sometimes these ‘juvenile’ forms are simple and unbranched 
with rotund apices. However, it is not possible to consider these 
stages as immature, since spores are commonly present, and the 
“adult” stage is often found interconnected. This later stage lacks 
the mealy margin, is always subsessile, has lightly pruinose discs and 
rotund apices, and has an irregular, intricated mode of branching. 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 107 


Ficures 75, 76.—75. Phacographis exaltata (Mont. & v. d. Bosch) Mill. Arg. (Hale 20394). 
76. P. inusta (Ach.) Mill. Arg. (isotype, Kalm). 


Figures 77, 78.—77. Phaeographis sericea (Eschw. in Mart.) Mill. Arg. (Hale 19727). 
78. P. sexloculata Fink (holotype, Fink 1436). 


This great variability of form seems to be rather common in those 
Graphidaceae which have thick walled brown spores with carbonized 
exciples. In the light of this variation, we should re-examine carefully 
the species criteria within this group. 

There are at least two other species of Phaeographis which may well 
prove to be synonyms of P. sericea: P. aggregata Redgr. and P. sub- 
stellata Zahlbr., but we have not yet examined the type specimens. 


46. Phaeographis sexloculata Fink, Mycologia 19:215. 1927. FiauRE 78 


Holotype: Yauco, Puerto Rico, Fink 1436 (MICH). 

Thallus smooth, continuous. Ascocarps black, slender, flexuose, 
semiemergent, unbranched to occasionally branched, 1-3 mm. long; 
exciple black, open; labia entire, erect or somewhat convergent, but 
exposing the hymenium; hymenium 65-80 » high. Spores 8 per ascus, 
commonly 6 locular, 8-10 18-26 un. 

Reactions: KOH-+ yellowish, P—, no microchemical tests made. 

Specimen examined: Carmen Island, Rabenhorst 25 p.p. (M). 

The Mexican material of this species was reported by Kremplehuber 
(1876, p. 148) as Graphis (Phaeographis) leiogrammodes. However, 
the holotype of @. letogrammodes has muriform spores. 


108 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Excluded or Doubtful Species 


47. Graphina sophistica (Nyl.) Mill. Arg. Flora 63:40. 1880. 
Graphis sophistica Nyl. Ann. Sci. Nat. Bot. ser. 4, 19:359. 1868. Syntypes: 
Villeta, Colombia, Lindig 876, 902, 2726, 2737 (not seen). 

Thallus continuous, smooth. Ascocarps subsessile, sparingly 
branched or unbranched, black; disc narrow, epruinose; exciple black, 
open or closed; labia convergent, entire or lightly crenate. Spores 
2-4 per ascus, 12-20 35-65 yp. 

This species was reported from Monterrey, Mexico, by Miiller 
(1894, p. 92), but we have seen neither this specimen, the type, or 
any Mexican material referable here. 


48. Graphis scripta (L.) Ach. Vet. Akad. Nya Handl. (Stockholm) 145. 1809. 
Lichen scriptus L. Sp. Pl. 1140. 1753. 


Graphis scripta and a number of closely allied species must be 
critically examined before any sound taxonomic judgments can be 
made about this widespread group. Redinger (1935, p. 8) separated 
G. tenella Ach., G. leptocarpa Fée, G. scripta, G. lineola Ach., and 
G. pavoniana Fée on the basis of spore size, degree of ascocarp immer- 
sion, and length and branching of the ascocarps. The first three of 
these four criteria are nearly useless in this group. The isotypes of 
G. tenella Ach. (UPS) have ascocarps with convergent labia which 
become striate with age. The isotype of G. lineola Ach. (UPS) con- 
tains a Melaspilea, an Opegrapha (which conceivably could be a 
lectotype), and a sterile unidentifiable entity. 

Graphis scripta itself can probably be typified through the Dillenian 
syntype cited by Linnaeus. This would still leave approximately 
190 named varieties, forms, etc., listed in Zahlbruckner’s Catalog. 
A critical study of these taxa in addition to numerous related species 
in Graphis and Phaeographis will pose a formidable problem for future 
lichenologists. 


49, Graphis striatula (Ach.) Sprengl. Syst. Veg. 4:250. 1827. Figure 79 
Opegrapha striatula Ach. Syn. Lich. 74. 1814. Type: Guinea (UPS, 
isotype). Reactions: KOH-+ reddish, P—, no microchemical tests made. 
Thallus thin, discontinuous. Ascocarps sessile, black, Opegrapha- 
like, straight to slightly flexuose, unbranched, 0.5-2 mm. long, about 
0.3 mm. wide; exciple black, usually open, 100 4 high; labia con- 
vergent, striate. Spores 8 per ascus, 11-13 locular, 8-10 X38-438 ug. 
This species was reported from Orizaba, Mexico, by Nylander 
(1858, p. 381), along with two new varieties, var. swblaevis Nyl. and 
var. pulverulenta Nyl. The former variety was described as having 
branched ascocarps, the latter as having barely striate labia. Miller 
(1887, p. 35) made the transfer Graphis duplicata var. sublaevis (Nyl.) 
Mill. Arg. He distinguished G. duplicata from G. striatula only by 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 109 


000000000000000000000000055 


Ficures 79, 80.—79. Graphis striatula (Ach.) Sprengl. (isotype). 80. Graphis vermiformis 
(Eschw. in Mart.) Nyl. (s. c., FH-Tayl). 


the more sessile unbordered ascocarps of the latter. We have not 
seen any material of these taxa from Mexico. 

The isotypes of G@. striatula and G. duplicata actually cannot be 
separated on the basis of sessile vs. nonsessile ascocarps, although 
they may be maintained by the gross form of the lirellae. In G. 
striatula, they are Opegrapha-like, unbranched, nearly straight, and 
short. In G. duplicata, they are elongate, branched, curved and 
flexuose. It would seem therefore that G. striatula var. sublaevis is 
referable to G. duplicata, as Miiller indicated. 


50. Graphis vermiformis (Eschw. in Mart.) Nyl. Flora 41:381. 1858. 
Fiaure 80 
Graphis illinata var. vermiformis Eschw. in Mart. FI. Bras. 1:83. 1833. 
Type: Near Pard, Brazil (not seen). 

Thallus thin, continuous, slightly roughened. Ascocarps sessile, 
unbranched, 1-3 mm. long, 0.5 mm. wide, concolorous with the 
thallus; exciple black, closed; labia convergent, more or less denticu- 
late or dentate, completely covered by a thalline veil. Spores 8 per 
ascus, vermiform, 20-25 locular, 7-9 65-75 yu. 

This species was reported from Orizaba, Mexico, by Nylander 
(1858, p. 381), but we have not verified this report. The Guianan 
specimen on which the above description was based is in the Taylor 
Herbarium (FH), labeled ‘“Graphis illinita Eschw.” and reacts 
KOH-+ brown, P+ brown. 

Graphis vermiformis is listed by Zahlbruckner (1923, p. 297) as a 
synonym of G. candidata Nyl. However, Nylander’s elevation of var. 
vermiformis to species level antedates the publication of G. candidata 
by 16 years. Hence if the two taxa are identical, G. vermiformis has 
priority. This species is closely related to Graphis anguilliformis 
Tayl., differing in the more slender vermiform spores and completely 
covered ascocarps. 

686-879 —63—— 4 


110 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


51. Melaspilea lentiginosa (Lyell in Leight.) Mill. Arg. Mem. Soc. Phys. Hist. 
Nat. Genéve 29, No. 8:19. 1887. 

Opegrapha lentiginosa Lyell in Leight. Ann. Mag. Nat. Hist. 2, No. 13:211. 
1854. Types: New Forest, Hants., Lyell (US, isosyntype); St. Leonard’s 
Forest, Sussex, England, Borrer (not seen). 

This species was reported from Orizaba, Mexico, by Nylander 
(1858, p. 381) but we have not checked this record. The isosyntype 
of Opegrapha lentiginosa, and all specimens we have seen that were 
referred here by Redinger (1938, p. 228), are lichen parasites on a 
Phaeographis species. 

52. Melaspilea leucinoides Mill. Arg. Bull. Herb. Boiss. 2:92. 1894. 


Holotype: Jalisco, Mexico, Pringle 216 (G). 
This entity is an Opegrapha. 
53. Melaspilea mesophlebia (Nyl.) Mill. Arg. Hedwigia 34:143. 1895. 
Opegrapha mesophlebia Nyl. Sert. Lich. Trop. Singapore et Labuan 42. 
1891. 
Opegrapha mesophlabia Ny]. Bull. Torrey Bot. Club 19:250. 1892. A 
sphalm. Holotype: San Luis Potosf, Mexico, Pringle 230 (II). 


This species also belongs in the genus Opegrapha. 


54. Melaspilea microphlebia (Nyl.) Zahlbr. Cat. Lich. Univ. 2:279. 1923. 
Opegrapha microphlebia Nyl. Bull. Soc. Linn. Norm. 2, no. 3:272. 1869. 
Type: Guadeloupe, Husnot (not seen). 


Eckfeldt (1892, p. 250) identified Pringle s.n., from Monterrey, 
Mexico, as Opegrapha microphlebia Nyl. We have checked the Pringle 
collection (VT) and found that it is indeed an Opegrapha. 

55. Phaeographina leiogrammodes (Kremplh.), comb. nov. Figure 81 
Graphis leiogrammodes Kremplh. Natur. For. Kjében. Vid. Medd. 5:25. 
1873. Holotype: Lagda Santa, Brazil, Warming 22 (M). Reactions: 
KOH+ reddish, P—, o-T-—, no acids demonstrated. 
Phacographis leiogrammodes (Kremplh.) Miill. Arg. Nuov. Giorn. Bot. Ital. 
23:397. 1891. 

The holotype specimen is undoubtedly a Phaeographina, externally 
quite similar to P. exilior (Vain.) Zahlbr. The juvenile spores are 
transversely 4 locular, as described, but at maturity one or all of the 
4 locules become biocellate. 

Krempelhuber (1876, p. 148) reported this species from Mexico, 
but these collections are all referable to Phaeographis. 

56. Phacographina scalpturata (Ach.) Miill. Arg. Flora 65:399. 1882. 
Figure 82 
Graphis scalpturata Ach. Syn. Lich. 86. 1814. Type: South America 
(UPS, isotype). Reactions: KOH+ reddish, P—, no microchemical 
tests made. 

Thallus smooth, continuous. Ascocarps barely emergent, flexuose, 
2-6 mm. long, about 0.4 mm. wide, apically rounded, often surrounded 
by a lighter thalline area; disc flat, wide, black or lightly pruinose; 


WIRTH & HALE—-MEXICAN GRAPHIDACEAE 111 


Figure 81.—Phaeographina leiogrammodes (Kremplh.) Wirth & Hale (holotype, 
Warming 22). 


Figure 82.—Phaeographina scalpturata (Ach.) Mill. Arg. (isotype). 


exciple pale brown, open; labia entire, divergent; hymenium clear, 
150 » high. Spores 1 per ascus, densely muriform, 23-26 80-85 uy. 

A record from Mexico (Pringle 335) reported by Davis (1936) has 
not been seen. Phaeographina scalpturata is evidently a completely 
misinterpreted species. Many previous workers have mentioned a 
carbonized exciple and placed the species in section Hleutheroloma. 
Much of the material we have seen under this name, identified by 
Nylander, Miiller, Redinger, etc., has black exciples, huge spores, 
sessile ascocarps, and prominent thalline margins. This entity, for 
which we have not been able to determine a specific name, was listed 
on p. 103 as Phaeographinasp. We have examined very few collections, 
none from Mexico, that can be identified with P. scalpturata as here 
delimited. 

57. Phaeographis inusta (Ach.) Miill. Arg. Flora 65:383. 1882. Figure 76 
Graphis inusta Ach. Syn. Lich. 85. 1814. Type: Canada, Kalm (UPS, 
isotype). Reactions: KOH+ reddish, P—, o-T—, no acids demonstrated. 

Thallus continuous, minutely roughened to nitid. Ascocarps semi- 
emergent, unbranched to sparingly branched, elliptical to lirelline, 
apically rounded, 0.5-1.2 mm. long, about 0.2 mm. wide, usually 
without a thalline margin; dise dark brown, slightly concave, wide; 
exciple red-brown, closed, thickened below; labia entire, divergent, 
often slightly carbonized; hymenium about 60 » high. Spores (?6-)8 
per ascus, 4 locular, 6-10 16-21 u. 

Nylander’s report from Mexico (1858, p. 381) has not been verified. 
An examination of the isotype of P. inusta disclosed that the tradi- 
tional concept of the species is probably incorrect. P. inusta is usually 
regarded as being much like P. dendritica, differing primarily in having 
an open exciple. However, the authentic material has constantly 
4 locular spores and brown closed exciples. 


112 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Phaeographis inustoides Fink (holotype: Puerto Rico, Heller 4430, 


MICH) is apparently distinct. 


It has 5-7 locules per spore, norstictic 


acid, pale labial apices, and a prominently cellular cortex (chondroid 
in the sense of Redinger, a fact which throws doubt on his placing this 
entity as a synonym of P. punctiformis (Eschw.) Mill. Arg.). 


Type Specimens Examined and Their 
Chemical Reactions 


GRAPHINA 

Graphina acharii var. subintegra 
Zahlbr. [=G. inturgescens (Kremplh. 
Mill. Arg.]. Pringle 5 p.p. (MICH, 
isosyntype) KOH—, P—. 

Graphina acromelaena Mill. Arg. 
[=G. parilis (Kremplh.) Miill. Arg.]. 
Tonduz s.n. (G, holotype): KOH+ 
yellow, stictic acid. 

Graphina acrophaea Mill. Arg. [=G. 
parilis (Kremplh.) Mill. Arg.]. 
Langlois 751 (US, isosyntype): 
KOH + yellow, stictic acid. 

Graphina aibonitensis Fink. Fink 
2017 (MICH, holotype): KOH+ 
red, salacinic acid. 

Graphina balbisii var. monospora 
Redgr. Malme 494 (S, holotype; 
FH, isotype): KOH—, P—. 

Graphina bipartita Mill. Arg. Balansa 
1876 (G, holotype): KOH+ red, 
norstictic acid. 

Graphina bothynocarpa Redgr. Malme 
2464 (5, holotype): KOH—, KC—, 
P-. 

Graphina cinerea Fink. [=G. scoleci- 
tis (Tuck.) Fink]. Fink 1163 
(MICH, holotype): Thallus KOH—, 
P—; ascocarps KOH+ blackish, 
P-. 

Graphina collatinensis var. lirelliformis 
Redgr. [=G. confluens (Fée) Mill. 
Arg.]. Malme 84 (S, holotype): 
norstictic acid. 

Graphina corcovadensis Redgr. Malme 
41 (8, holotype): KOH-+ yellow, 
stictic acid. 

Graphina epiglauca Mill. Arg. [=G. 
confluens (Fée) Mill. Arg.]. Pitter 
s.n. (US, isotype): KOH+ yellow, 
stictic acid, lichexanthone. 


Graphina  heteroplacoides Redgr. 
Malme 1227 (S. holotype): KOH+ 
red, P—, o-T unknown acid. 

Graphina incerta Redgr. Malme 3644 
(S, syntype): KOH—, P—; Malme 
1890f (8S, syntype) [=Graphis 
(Graphina) leucopepla Tuck.]: P+ 
red, protocetraric acid. 

Graphina interstes Mill. Arg. Tonduz 
s.n. (G. holotype): KOH+ red, o-T 
unknown acid. 

Graphina luridoolivacea Fink. Fink 
657 (MICH, holotype): KOH+ 
red, P—, o-T—. 

Graphina malmei Redgr. Malme 3508 
(S, holotype): Thallus KOH— 
P—; ascocarps cinnabar, KOH+ 
purple. 

Graphina obtectula Mill. Arg. Ton- 
duz s.n. (G, holotype): KOH+ yel- 
low, o-T unknown acid. 

Graphina olivobrunnea Fink. [=G. 
scolecitis (Tuck.) Fink]. Fink 1224 
(MICH, holotype): Thallus KOH—, 
P—; ascocarps KOH-+ purple, P—, 
o-T—. 

Graphina palmeri Zahlbr. Pringle 9 
(MICH, isotype): KOH+ yellow, 
o-T unknown acid. 

Graphina platycarpa Fink [non G. 
platyearpa (Eschw. in Mart.) Mill. 
Arg.] [=G. platycarpina Zahlbr.]. 
Fink 1774 (MICH, holotype): P+ 
red, protocetraric acid. 

Graphina plittii Zahlbr. Plitt s.n. 
(US, BPI, isotypes): KOH+ red, 
salacinic acid. 

Graphina pseudophlyetis var. mono- 
spora Redgr. Malme 2023, 2177, 
3678, 3679 (S, syntypes): KOH—, 
Pp—. 


WIRTH & HALE—-MEXICAN GRAPHIDACEAE 113 


Graphina puiggarii Mill. Arg. Puzg- 
gart 506 (G, holotype): KOH—, 


D 


Graphina puiggarii var. corumbensis 
Redgr. [=G. parilis (Kremplh.) 
Mill. Arg.]. Malme 3639, 3636, 


3637 (5S, syntypes): KOH+ yellow, 
stictic acid. 
Graphina reniformis var. subastroidea 


Redgr. Malme 3520 (S, holotype): 
P+ red, protocetraric acid and 
lichexanthone. 


Graphina rimulosa Redgr. Malme 
1089 (S, holotype): KOH+ reddish, 
P-. 

Graphina riopiedrensis Fink. Fink 
2167 (MICH, holotype): KOH+ 
yellowish, stictic acid. 

Graphina sulcata Fink. Fink 659 
(MICH, holotype): KOH + red, nor- 
stictic acid. 

Graphina sulcatula var. conglomerata 
Mill. Arg. Balansa 4185a (G, 
syntype): KOH—, P-—. 

Graphina triangularis ZahIbr. Pringle 
17 (MICH, isotype): KOH+ yellow- 
red, o-T unknown acids. 

Graphina vestitoides Fink. Fink 1986 
(MICH, holotype): KOH—, P—. 


GRAPHIS 

Graphis abaphoides Nyl. (Graphina 
abaphoides (Nyl.) Mill. Arg.). 
Calkins 134 (US,  isosyntype): 
KOH+ yellowish, P+ red, proto- 
cetraric acid and an unknown acid. 

Graphis adpressa Vain. Vainio 1289 
(TUR, holotype): KOH—, P-. 

Graphis afzelii Ach. (UPS, isotype): 
Ascocarps KOH—, P—, C+ red, 
lecanoric acid. 

Graphis albescens Vain. Vatnio s. n. 
(TUR, holotype): KOH+ yellow, 
stictic acid. 

Graphis albida Fink. Fink 1777 
(MICH, holotype): P+ red, pro- 
tocetraric acid. 


Graphis albostriata Vain. [= 
Graphina albostriata (Vain.) 
Zahlbr.]. Vainio 1538 (TUR, 


holotype): KOH+ yellow, P-, 
o-T unknown acid. 


Graphis anguinaeformis Vain. [= 
Graphina anguinaeformis (Vain.) 
Zahlbr.]. Vainio 274 (TUR 


holotype): KOH—, P-. 

Graphis arecae Vain. Merrill 6724 
US, isotype): KOH+ yellow, 
stictic acid. 

Graphis atroalba Vain. non Kremplh. 
(=G. afzelii Ach.). Vainio 
189 (TUR, holotype): Ascocarps 
KOH—, C+ red. 

Graphis brachycarpa Vain. 
1092 (TUR, holotype): 
P-. 

Graphis caesiella Vain. Vainio 45 
(TUR, holotype): KOH-+ red, nor- 
stictic acid. 

Graphis caesioglauca Redgr. Malme 
1526 (S, holotype): KOH— P-—. 


Vainio 
KOH-, 


Graphis _carassensis Vain. [= 
Graphina carassensis (Vain.) 
Zahlbr.]. Vainio 1467 (TUR, 


holotype): KOH+ reddish, P-—, 
no acids demonstrated. 

Graphis cladophora Vain. [=Gra- 
phina cladophora (Vain.) Zahlbr.]. 
Merrill 7990 (US, isotype): KOH+ 
red, norstictic acid. 

Graphis collosporella Vain. [=Gra- 
phina collosporella (Vain.) Zahlbr.]. 
Thaxter 9, 10 (FH, syntypes): 
KOH-+ yellowish, unknown acid. 

Graphis compulsa Kremplh. [=G. 
desquamescens Fée]. Glaziou 5082 
(M, holotype): KOH+ red, nor- 
stictic acid. 

Graphis dehiscens Vain. [=Gra- 
phina!]. Vainio 306 (TUR, holo- 
type): KOH+?, P—. 

Graphis diaphoroides Miill. Arg. 
Lahm 91 (US, isosyntype): KOH+ 
red, norstictic acid. 

Graphis dimidiata Vain. [=Graphina 
dimidiata (Vain.) Zahlbr.]. Vainio 
332 (TUR, holotype): KOH—, P—. 

Graphis diorygmatoides Vain. 
[=Graphina diorygmatoides 
(Vain.) Zahlbr.]. Merrill 8515 (US, 
isotype): KOH-+ red, norstictic acid. 

Graphis elongata Vain. [=Graphina 
elongata (Vain.) Zahlbr.]. Vainio 
782 (TUR, holotype): KOH+ red, 
norstictie acid. 


114 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM, 


Graphis endoxantha Nyl. Pancher 
s.n. (H, holotype): KOH-+ reddish, 
P—. 

Graphis eugeniae Vain. Whitford 1087 
(US, isosyntype): KOH-+ red, sala- 
cinic acid. 

Graphis floridana Tuck. (US, isotype): 
KOH-+ red, norstictie acid. 

Graphis granulocarpa Vain. Vainio 
3680 (TUR, holotype): KOH -4- red- 
dish, P—. 

Graphis hiascens var. clausior Vain. 
[=Graphina hiascens var. clausior 
(Vain.) Zahlbr.]. Ramos & Edano 
29548 (US, isotype): KOH+ yellow, 
stictic acid. 

Graphis hololeucoides Nyl. [=Gra- 
phina hololeucoides (Nyl.) Mill. 
Arg.]. Ghiesbreght s.n. (H, isotype): 
Ascocarps KOH-+ faint yellowish, 
P+. 

Graphis humilis Vain. Merrill 9067 
(US, isotype): KOH+ red, o-T 
unknown acid. 

Graphis illota Mull. Arg. 
holotype): KOH—, P—. 

Graphis immersa Fipk. Fink 1613 
(MICH, holotype): KOH—, P+. 

Graphis includens Vain. [=Gra- 
phina!]. Vainio 765 (TUR, holo- 
type): KOH+ reddish, P—, 0-T—. 

Graphis inidita var. pularensis Vain. 
[=Graphina inidita var. pularensis 
(Vain.) Zahlbr.}]. Ramos 19446 (US, 


Ule 273 (G, 


isotype): KOH+ yellow, stictic 
acid. 
Graphis isidiosa Vain. [=Phaeo- 


graphina isidiosa (Vain.) Zahlbr.]. 
Merrill 6647 (US,  isosyntype): 
KOH-4 red, norstictic acid. 

Graphis leucopepla Tuck. [=Gra- 
phinal!]. Wilson 56 (FH, holotype): 
Thallus KOH—, P-—; ascocarps 
KOH-—, P+. 

Graphis lutescens Kremplh. non Fée 
[=Phaeographina lutescens 
(Kremplh.) Zahlbr.]. Glaztou 3374 
(M, holotype): KOH-+ red, norstic- 
tic acid. 


Graphis macgregorii Vain. [=Gra- 
phina hiascens (Fée) Mill. Arg.]. 
MacGregor (607 US, isotype): 


KOH-4 red, norstictie acid. 


Graphis macrospora Kremplh. 
{[=Graphina confluens (Fée) Mill. 
Arg.]. Glaziouw 3379 (M, syntype): 
KOH-+ yellow, stictic acid, lichex- 
anthone, a fatty substance. 


Graphis miniata Redgr. [=Phaeo- 
graphina  chrysocarpa (Raddi) 
Redgr.]. Mosén 3145, 3318, 3577 


(S, syntypes): Thallus KOH—, P—; 
ascocarps KOI-+ purple. 

Graphis olivacea Redgr. Malme 2267 
(S, holotype): IKOH-+ reddish, P—. 

Graphis orientalis Vain. [=Graphina 
hiascens (Fée) Mill. Arg.]. Merrill 
6716 (US, isotype): KOH+ red, 
norstictic acid. 

Graphis oscitans Tuck. [=Phaeo- 
graphina oscitans (Tuck.) Zahlbr.]. 
Mann, sn. (US, isotype): KOH-+ 
yellow, stictice acid. 

Graphis peralbida Nyl. [=Graphina 
bipartita Mill. Arg.]. Pringle 222 
(H, holotype; FH, VT, isotypes): 
KOH-+ red, norstictie acid. 

Graphis phaeospora Vain. [=Gra- 
phina inturgescens (IXremplh.) Mill. 
Arg.]. Vainio 682 (TUR, holo- 
type): KOH—, P—. 

Graphis pseudosophistica Vain. 
[=Graphina pseudosophistica 
(Vain.) Zahlbr.]. Vainio 757 
(TUR, syntype): KOH-4- reddish, 
P—, o-T unknown acid; Vainio 
1003 (TUR, syntype): KOH-+ red, 
P—; Vainio 1404 (TUR, syntype): 
KOH—, P-. 

Graphis rimulosa var. lignicola Fink. 


Fink 1873 (MICH, holotype): 
KOH—, P-—. 
Graphis sitiana Vain. Vainio 533 


(TUR, holotype): KOH—, P—. 

Graphis subamylacea Zahlbr. Pringle 
24 (MICH, isotype): KOH+ yel- 
low, stictic acid. 

Graphis subeabbalistica Vain. [=Gra- 
phina!]. Vainio 1246 (TUR, holo- 
type): KOH-+ reddish, P—. 

Graphis subducta Vain. [=Graphina 
parilis (Kremplh.) Mill. Arg.]. Mer- 
rill 8576 (US, isotype): KOH+- 
yellow, stictic acid. 


WIRTH & HALE—MEXICAN GRAPHIDACEAE 115 


Graphis subelegans Nyl. [=G. endox- 
antha Nyl.J]. Pringle 162 (H, 
holotype): KOH-+ red, P—. 

Graphis subparilis Nyl. Calkins s.n. 
(US, isotype): Thallus KOH—, 
P—; ascocarps KOH-+ purple. 

Graphis subserpentina Nyl. [=Gra- 
phina hiascens (Fée) Mill. Arg.]. 
Gardner s.n. (US, isotype): KOH+ 
red, norstictic acid. 

Graphis tongloensis Vain. Merrill 
7985 (US, isotype): KOH—, P—. 

Graphis tumidella Fink. Fink 1737 
(MICH, holotype): KOH—, P-—. 

Graphis turbulenta Nyl. Calkins s.n. 
(US, isotype): KOH-+ red, nor- 
stictic acid. 

Graphis virens Mill. Arg. Ule 275 
(G, holotype): KOH+ red, P—, 
o-T-—. 

Graphis yaucoensis Fink [=G. caesi- 
ella Vain.]. Fink 1691 (MICH, 
holotype): KOH-+ red, norstictic 
acid. 

MEDUSULINA 

Medusulina texana Mill. Arg. Lck- 
feldt 56A (G, holotype): KOH+ 
yellow, stictic acid. 

OPEGRAPHA 

Opegrapha dendritica Ach. [= Phaeo- 
graphis dendritica (Ach.) Miill. 
Arg.]. (UPS, isotype): KOH+ red, 
norstictic acid. 

PHAEOGRAPHINA 

Phaeographina caesiopruinosa var. ab- 
breviata Redgr. Malme 1888, 1492, 
1562 (5S, syntypes): KOH-+ red, 
P—, o-T-. 


Phaeographina difformis Fink. Fink 
1874 (MICH, holotype): KOH—, 
P-. 

Phaeographina epruinosa Redgr. [= P. 
explicans Fink in Hedrick]. Malme 
3662 (8S, holotype): KOH+ yellow- 
red, o-T—. 

Phaeographina myriogloena Mill. Arg. 
Leprieur 196 (G, holotype): KOH+ 
red, P—, o-T—. 

Phaeographina oxalifera Redgr. 
Malme 3664 (8, syntype): KOH+ 
red, norstictic acid. 

Phaeographina rhodoplaca Mill. Arg. 
Tonduz s.n. (G, holotype): KOH—, 
P-. 

PHAEOGRAPHIS 

Phaeographis astroidea Mill. Arg. 
[=Graphis!]. YZonduz s.n. (G, 
holotype): KOH+ reddish, P—. 

Phaeographis  cerviculata  Redgr. 
[=Graphina!]. Malme 3531 (8, holo- 
type): KOH-+ reddish, P—. 

Phaeographis inustioides Fink. Heller 
4430 (MICH, holotype): KOH+ 
red, norstictic acid and an unknown 
acid. 

Phaeographis pezizoidea var. pruinosa 
Redgr. Malme 2586C, 2030 (8, 
syntypes): KOH-+ red, norstictic 
acid, 

Phaeographis praestans Mill. Arg. 
{[=Graphis!]. Tonduzs.n. (G, holo- 
type): KOH—, P—. 

Phaeographis radiatoramosa Redgr. 
Mosén 3578 (S, holotype): KOH+ 
red, norstictic acid. 


Bibliography 


Anpmrson, L. E. 1954. Hoyer’s solution as a rapid permanent mounting 
medium for bryophytes. Bryologist 57:242-244, 

Davis, H. B. 1936. Life and works of Cyrus Guernsey Pringle. Univ. of 
Vermont, Burlington. 765 pp. 

Ecxrevpt, J. W. 1892. An enumeration of some rare North American 
lichens. Bull. Torrey Bot. Club 19:249-253. 

KREMPELHUBER, A. 1876. Lichenes Mexicani quos legit 1875 R. Rabenhorst. 
Hedwigia 15:148, 149. 

Miitter-Araau, J. 1894. Lichenes Eckfeldtiana. Bull. Herb. Boiss. 2, no. 
2:89-93. 

NyrtanpER, W. 1858. Lichenes collecti in Mexico a Fr. Miller. Flora 
41:377-381. 

Repincer, K. 1933. Die Graphidineen der ersten Regnell’schen Expedition 
nach Brasilien 1892-94. Ark. Bot. 26A, no. 1:1-105. 

1935. Die Graphidineen der ersten Regnell’schen Expedition nach 

Brasilien 1892-94. II. Ark. Bot. 27A, no. 3:1-103. 

1938. Graphidaceae. In Rabenhorst, Kryptogamen Flora, 9, part 2, 
no. 1:181-404. 

RuNnEMARKE, H. 1956. Studies in Rhizocarpon. 1. Taxonomy of the yellow 
species in Europe. Opera Bot. 2, no. 1:1-152. 

Santesson, R. 1952. Foliicolous lichens 1. Symb. Bot. Ups. 12, no. 1:1-590. 


116 


Index 


(Synonyms in italics. 


Arthonia 


caesiopruinosa, 101 
confluens, 74, 75 


Enterodictyon 


mexicanum, 82 


Fissurina 


virginalis, 90 


Graphina 


acharii, 65, 72, 82, 103 

achari var. subintegra, 81, 112 

acromelaena, 84, 86, 112 

acrophaea, 84, 112 

acrophaea f{. multilamellosa, 84, 85 

adscribens, 88 

aibonitensis, 89, 112 

balbisii, 73 

balbisii var. monospora, 112 

bipartita, 74, 112 

bothynocarpa, 112 

caesioradians, 101 

chrysocarpa, 102 

cinerea, 87, 112 

collatinensis, 74 

collatinensis var. lirelliformis, 74, 
76, 112 

collatinensis var. ocellariiformis, 74, 
76 

confluens, 64, 74 

corcovadensis, 86, 112 

dealbata, 73 

elongata, 77 

elongatoradians, 77, 78 

epiglauca, 74, 76, 112 

hemisphaerica, 64 

heteroplaca, 73 

heteroplacoides, 112 

hiascens, 78 

hiascens var. clausior, 79 

hololeucoides, 80 

incerta, 112 

insignis, 80 

insignis var. imperfecta, 80, 81 

insignis var. primaria, 80 

insignis var. tartarea, 80 


Page numbers of principal entries in boldface.) 


Graphina—Continued 


interstes, 112 

inturgescens, 81 

luridoolivacea, 112 

macella, 82 

macgregorit, 78, 79 

macrospora, 64, 74, 76 

malmei, 103, 112 

mexicana, 67, 82 

nitida, 83 

obtectula, 112 

olivobrunnea, 87, 112 

orientalis, 78 

palmeri, 84, 89, 91, 112 

parilis, 66, 84, 88 

peplophora, 67, 86 

peralbida, 74 

platycarpa, 112 

platygrapta, 74, 76 

plittii, 112 

pringlei, 84, 86 

pseudophlyctis var. monospora, 112 
puiggarit f. corumbensis, 84, 86, 113 
puiggarii f. puiggarii, 86, 113 
quassiaecola, 64 

reniformis var. subastroidea, 113 
rimulosa, 86, 113 

riopiedrensis, 113 

scolecitis, 87 

sophistica, 108 

subducta, 84 

subserpentina, 78, 79 

sulcata, 66, 88, 113 

sulcatula var. conglomerata, 88, 113 
triangularis, 84, 88, 91, 106, 113 
vestitoides, 73, 113 

virginalis, 90 

virginea, 84, 89, 90, 106 


Graphis 


abaphoides, 113 
acharii, 72 
adpressa, 113 
afzelii, 91, 113 
albescens, 113 


117 


118 


Graphis—Continued 
albida, 113 
albostriata, 113 
anguilliformis, 91, 109 
anguinaeformis, 113 
angustata, 94, 98 
annulata, 84 
arecae, 113 
atroalba, 91, 113 
atroleuca, 91 
balbisiz, 73 
balbisii var. monospora, 73 
balbisina, 96 
brachycarpa, 113 
caesiella, 92, 100, 113 
caesioglauca, 113 
candidata, 109 
carassensis, 113 
chlorocarpella, 96 
chrysocarpa, 102 
cladophora, 113 
collosporella, 113 
cometia, 91 
compulsa, 938, 113 
dehiscens, 113 
delicatula, 74 
desquamescens, 92 
diaphoroides, 113 
dimidiata, 113 
diorygmatoides, 113 
disserpens, 98 
duplicata var. sublaevis, 108 
durandi, 98 
elongata, 77, 113 
endoxantha, 94, 114 
eugeniae, 114 
flexibilis, 94 
floridana, 114 
glaucopis, 67, 94 
grammatica, 95 
granulocarpa, 114 
grossula, 95 
hiascens var. clausior, 114 
hololeucoides, 80, 114 
humilis, 98, 114 
illinata, 73 
illinata var. vermiformis, 109 
illota, 114 
immersa, 114 
implicata, 74, 81, 96 
includens, 114 
inidita var. pularensis, 114 
insignis, 80 


INDEX 


Graphis— Continued 
inturgescens, 81 
inusta, 111 
isidiosa, 114 
lactea, 98 
lecanorina, 98 
leiogrammodes, 107, 110 
leptocarpa, 108 
leucocarpa, 86 
leucopepla, 114 
lineola, 108 
longula, 96 
lutescens, 114 
macella, 82 
macgregortt, 78, 114 
macrospora, 74, 114 
mintata, 102, 103, 114 
mosquitensis, 99 
olivacea, 114 
ortentalis, 78, 114 
oscitans, 114 
parilis, 84 
peralbida, 74, 114 
phaeospora, $1, 114 
platycarpella, 98 
proserpens, 94, 98 
pseudosophistica, 114 
rimulosa var. lignicola, 114 
scalpturata, 110 
schizogramma, 98 
scolecitis, 87 
scripta, 92, 108 
sitiana, 114 
sophistica, 108 
striatula, 108 
striatula var. pulverulenta, 108 
striatula var. sublaevis, 108 
stromatoides, 98 
stygioarachnoidea, 67, 99 
subamylacea, 92, 100, 114 
subcabbalistica, 114 
subducta, 84, 114 
subelegans, 94, 115 
subparilis, 115 
subserpentina, 78, 115 
tenella, 108 
tongloensis, 115 
tumidella, 115 
turbulenta, 65, 115 
vermiformis, 109 
virens, 115 
virginalis, 90 
yaucoensis, 92 


INDEX 


Lecanactis 
exaltata, 106 
Leiogramma 
sericeum, 106 
virgineum, 90 
Medusulina 
nitida, 82 
texana, 115 
Melaspilea 
lentiginosa, 110 
leucinoides, 110 
mesophlebia, 110 
microphlebia, 110 
polymorpha, 100 
Opegrapha 
chrysocarpa, 102 
dendritica, 105, 115 
endochroma, 78 
hiascens, 78 
lentiginosa, 110 
mesophlabia, 110 
mesophlebia, 110 
microphlebia, 110 
striatula, 108 
Phaeographina 
asteroides, 101 
caesiopruinosa, 101 
caesiopruinosa var. abbreviata, 115 
caestopruinosella, 101 
caesioradians, 101 
chrysocarpa, 65, 73, 102 
difformis, 115 


Phaeographina—Continued 
elliptica, 67, 104 
epruinosa, 115 
exilior, 110 
leiogrammodes, 110 
myriogloena, 115 
oxalifera, 104, 115 
phaeospora, 81 
rhodoplaca, 115 
scalpturata, 104, 110 
sericea, 106 
sp., 103 
strigops, 67, 104 

Phaeographis 
ageregata, 107 
astroidea, 115 
cerviculata, 73, 115 
dendritica, 105, 111 
exaltata, 84, 89, 91, 106 
inusta, 111 
inustoides, 112, 115 
leiogrammodes, 110 
longula, 96 
pezizoidea var. pruinosa, 115 
praestans, 115 
punctiformis, 112 
radiatoramosa, 115 
sericea, 106 
sexloculata, 107 
substellata, 107 

Sarcographa, 106 


U.S. GOVERNMENT PRINTING OFFICE: 1963 


119 


UNITED STATES NATTONAL MUSEUM 


CONTRIBUTIONS FROM THE UNITED StTaTEs NATIONAL HERBARIUM 


Vo.LuME 36, Parr 4 


STUDIES ON PARMELIA 
SUBGENUS PARMELIA 


By Mason FE, Hate, Jr., and Syo Kurokawa 


BULLETIN OF THE UNITED StTaTES NaTIonaAL MusEuM 


SMITHSONIAN INSTITUTION e WASHINGTON, D.C. e 1964 


STUDIES ON PARMELIA SUBGENUS PARMELIA 


By Mason E. Hate, Jr., and Syo Kurokawa 


Introduction 


At least 250 distinct species, most of which occur in the tropics, 
are included in Parmelia subgenus Parmelia. They are generally 
characterized by lobes that are narrow (0.5-4.0 mm. wide), sublinear 
to subirregular, and often apically truncate. The apotheciaare adnate 
and usually imperforate, and there are rhizines over all or most of 
the lower surface. This broadly delimited group has usually been 
called section Hypotrachyna Vain., but we propose to recognize it as 
a subgenus, subgenus Parmelia, typified by P. saxatilis (L.) Ach. 

Subgenus Parmelia is coordinate with subgenus Amphigymnia 
(Vain.) Dodge, the broad-lobed Parmelias, and subgenus Xantho- 
parmelia (Vain.) Hale, comb. nov., based on section Xanthoparmelia 
Vain. (1890, p. 60), with Parn elia conspersa (Ach.) Ach. as the type 
species. The Xanthoparmelias are similar in configuration to many 
species in subgenus Parmelia, but they differ in being saxicolous, 
always containing usnic acid, and having simple rhizines. The brown 
Parmelias (sections Melanoparmelia Zahlbr., Vainioellae Gyel., and 
Olivascentes (Hue) Hillm.) are excluded from subgenus Parmelia, 
although we have not yet decided on their exact position in the genus. 

In the course of preparing a world monograph of subgenus Parmelia, 
we have examined the types of most of the described species and 
reevaluated the subgeneric classification. This preliminary study 
will present the outlines of a new sectional classification, descriptions 
of 52 new species, 2 new combinations, and 4 new names, and pre- 
liminary keys to the species in the major sections. A final mono- 
graph must await more extensive study of general herbarium material 
as well as field work, since many species are still known only from their 
type localities. 

This study has been made possible by the prompt and generous 
cooperation of the curators and directors of museums and university 
herbaria in lending type specimens and other valuable collections and 
in rendering assistance and providing facilities for study during visits. 
They include Dr. Sten Ahlner (Naturhistoriska Riksmuseet, Stock- 

121 


122 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


holm), Dr. Reino Alava (Botanical Institute, Turku), Dr. Ove 
Almborn (Botanical Museum, Lund), Dr. Y. Asahina (Research 
Institute for Natural Resources, Tokyo), Dr. Charles Baehni (Con- 
servatoire et Jardin Botaniques, Geneva), M. P. Bourrelly (Muséum 
National d’Histoire Naturelle, Paris), M. Skytte Christiansen (Botan- 
ical Museum, Copenhagen), Dr. G. Cufodontis (University of Vienna, 
Vienna), Dr. W. L. Culberson (Duke University, Durham), Dr. H. 
des Abbayes (Université de Rennes, Rennes), Dr. Henry Imshaug 
(Michigan State University, East Lansing), Mr. Peter James (British 
Museum, London), Dr. A. Kostermanns (Herbarium Bogoriense, 
Bogor), Dr. I. M. Lamb (Farlow Herbarium, Cambridge), Dr. R. A. 
Maas Geesteranus (Rijksherbarium, Leiden), Dr. J. Millar (Chicago 
Natural History Museum, Chicago), Dr. G. Moggi (Istituto Botanico, 
Firenze), Dr. E. Miller (Institut fiir Spezielle Botanik, Ziirich), C. E. 
Palmar (Glasgow Art Gallery and Museums, Glasgow), Dr. J. Poelt 
(Botanische Staatssammlung, Munich), Dr. K. Rechinger (Natur- 
historisches Museum, Vienna), Dr. C. Rogerson (New York Botanical 
Garden, New York), Dr. H. Roivainen (Botanical Museum, Helsinki), 
Dr. Rolf Santesson (Institute for Systematic Botany, Uppsala), Sir 
George Taylor (Royal Botanic Gardens, Kew), Dr. J. W. Thomson 
(University of Wisconsin, Madison), Dr. K. Verseghy (Museum of 
Natural History, Budapest), Dr. R. Vincenzo (Istituto Botanico, 
Rome), and Dr. R. Woodson (Missouri Botanical Garden, St. Louis). 

Drs. G. Degelius and D. D. Awasthi (abbreviated DEGEL and 
AWAS respectively in the list of species) have kindly sent us their 
valuable private collections. Dr. Z. Chernohorsky arranged for loans 
of Gyelnik’s collections and kindly provided facilities for study in 
Prague. Mr. M. Wirth assisted in the early work of chemical testing. 
Drs. Rolf Santesson and Ove Almborn have kindly read various parts 
of the manuscript and given invaluable aid in problems of sectional 
classification. 

This study has been supported in part by a grant from the National 
Science Foundation. 


Morphological Characters 


The following are the more important morphological characters 
that we have found in subgenus Parmelia. Some of them have been 
overlooked or poorly understood by previous workers and will be 
discussed in detail. 

1. Rhizines: Are produced more or less uniformly over the entire 
lower surface in the majority of species. Certain species with sub- 
rotund subirregular lobes, however, may have a narrow though distinct 
bare or papillate zone around the margins. Three major types of 
rhizines may be distinguished: simple and unbranched or rarely 


HALE & KUROKAWA—SUBGENUS PARMELIA 123 


sparsely furcate; dichotomously and usually richly branched; and 
squarrosely branched (fig. 1). Branching of the rhizines has proved 


Ficure 1.—Schematic drawings of the three types of rhizines: Simple, squarrosely branched, 
and dichotomously branched (left to right). 


to be one of the most important characters on which our proposed 
sectional classification is based. 

2. Cilia: Two types of ciliate outgrowths occur on the margins of 
lobes, simple cilia and bulbate cilia. Simple cilia, comparable to those 
known in Anaptychia or Cetraria, occur in at least 34 species of sub- 
genus Parmelia (pl. 1). These same species usually have a black lower 
surface and simple rhizines (e.g., P. dissecta Nyl. and P. tiliacea 
(Hoffm.) Ach.). Bulbate cilia differ from simple cilia in having a 
conspicuously inflated base. This structure (pl. 1) is known in some 
45 species, all of them tropical. It has not been consistently recog- 
nized by lichenologists, although commemorated in names such as 
P. circumnodata Nyl. Bulbate species may have branched or un- 
branched rhizines and a pale or black lower surface. Bulbate cilia 
may merely be modifications of simple cilia, but they are distributed 
around the lobes with much greater regularity. Chemical data 
(table 1) indicate that simple-ciliate and bulbate-ciliate species are 
far more closely related to each other than to any other groups in 
the subgenus. Both types of cilia are probably related to rhizines 
and, as in the case of rhizines, are of very great importance in sub- 
generic classification. 

3. Lobules: These structures have usually been confused with 
isidia. They apparently originate as isidia but soon begin to flatten 
and grow horizontally (cf. Parmelia ensifolia, pl. 7). At maturity 
they are more or less dorsiventral and often ciliate. Lobules are 
rather rare and best seen in P. culmigena Zahlbr., P. digitata Lynge, 
P. horrescens Tayl., or P. ensifolia Kurokawa. 

4. Pustules: Are coarse papillar outgrowths from the upper cortex. 
They are extremely fragile and seem to lack the thalloid structure of 
isidia. They may remain intact, as in P. dactylifera Vain., or at 
length erupt or burst open apically and form dense pustular masses, 
with little or no tendency to become sorediate (pl. 1). If coarse 


124 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


soredia form, they are not so dense as to obscure the basic pustular 
structure. Typically pustulate species include P. aurulenta Tuck., 
P. endochlora Leight., P. formosana Zahlbr., and P. spumosa Asah. 

5. Maculae: Are submacroscopic white dots uniformly scattered in 
the upper cortex. They are apparently caused by irregular clumping 
of algal colonies in the gonidial layer. They are at times difficult to 
recognize, and most lichenologists have in fact overlooked them. 
Maculae are nevertheless important, especially at the species level. 
They occur in such well-known species as P. laevigata (Sm.) Ach. and 
P. tiliacea (Hoffm.) Ach. 

6. Pseudocyphellae: Here we refer to actual pores in the upper 
cortex. There are apparently two different kinds. One is found in 
species related to P. borreri (Sm.) Turn., where the pores are distinctly 
orbicular or elliptical and scattered irregularly in the upper cortex 
(pl. 1). They are correlated with the presence of simple rhizines 
and P— acids (gyrophorie acid, lecanoric acid, and fatty acids). 
A second type of pore is found in species related to P. sazatilis (L.) 
Ach., where the pores are angular or effigurate and are arranged in 
more or less definite patterns on the cortex and margins of lobes 
(pl. 1). These species usually contain a P+ acid (protocetraric or 
salacinic acid) and have a black lower surface with squarrosely 
branched rhizines. 

7. Reticulation: The upper cortex of a small group of species 
related to P. reticulata Tayl. is finely reticulately maculate and 
fissured (pl. 1). Initially the cortex is maculate to the margins of 
the lobes and at maturity actual fissuring usually occurs. Many 
species of Parmelia are irregularly or more or less reticulately cracked 
on older lobes, but young lobes lack any reticulation. Reticulate 
species always have a black lower surface and simple to squarrosely 
branched rhizines. 

8. Isidia and soredia: These two characters are well known to 
lichenologists through the work of Du Rietz (1924) and require no 
further discussion here. They are valuable species characters in 
Parmelia but have no value at the sectional level. 

9. Apothecia: These characters offer little basis for classification 
in subgenus Parmelia. Spores are uniformly small, generally only 
6-18 » in length, and attain a length of 30 » in only a few species such 
as P. leucoms Kremplh., P. mutata Vain., and P. versiformis Kremplh. 
Except for P. cetrata Ach., P. homotoma N yl., and P. reticulata Tayl., 
the apothecial disc is never perforate. There is little significant varia- 
tion in degree of adnation on the thallus, except in such unusual 
pedicellate species as P. peruviana Nyl. Coronate exciples and basal 
retrorse rhizines are often found on the apothecia of species with 


HALE & KUROKAWA—SUBGENUS PARMELIA 125 


bulbate cilia. Parmelia carporrhizans Tayl., a ciliate species, is 
densely rhizinate at the base of the apothecia. 

10. Anatomy: The internal anatomy is for the most part uniform. 
The total thickness of the thallus varies from about 60 4 to 250 uy; 
the cortex is often thinner and more fragile than in subgenus 
Amphigymnia. Peculiar moniliform cells are found in the medulla 
of P. galbina Ach., P. obsessa Ach., and P. metarevoluta Asah., three 
closely related species (cf. Asahina, 1952, p. 98). Future workers 
may place greater reliance on internal characters, but for the time 
being this approach seems unrewarding. 


Chemical Characters 


The taxonomy of subgenus Parmelia cannot be studied fully with- 
out the use of chemical tests. This does not imply that we are 
accepting or deliberately describing so-called “chemical species.” 
Anyone who works with a genus such as Parmelia, which has a wide 


TaBLE 1.—Number of species in the various sections and subsections containing 
19 important chemical substances. 


Section Section 
Parmelia Imbricaria 
- ese | S| cg | Se | ae 

gs g & lares : 83 § 3 cheila chyna 

84 | 33 38 | SE 

ga | Be £2 | 2a 

g |e" a |a 
Number of species 30 | 25 10 46 | 33 46 83 
Gyrophorie acid —| 6] — 7) 4) — 6 
Lecanoric acid —|10} — 3 4 2 = [5] 
Salacinic acid 28 | — 7 10 4 3 8 
Stictic acid —|— 2 2 5 1 
Norstictic acid —|— — 4 2 — 5 
Protocetraric acid 2) — — 3 1 12 15 
Usnic acid —|— — 18 1 9 12 
Rhodophyscin —|-|] — — 3 9 
K— pigments —|—| — {|—/ 11 5 5 
Olivetoric acid — |}; — —_— —|— — 4 
Cryptoclorophaeic acid —}— 1 —|—|] — 1 
Evernic acid —|— — —|— 1 5 
Divaricatic acid —|— — —|— 8 — 
Perlatolic acid —|— — —|— 1 — 
Alectoronic acid —|— — —|— — 7 
Barbatie acid —|— — —}|}— 1 11 
Lichexanthone —|— — —|— — 7 
KC+ unknown (“livida”) | —}|—| — | —|— 1 5 


« Lecanoric acid in section Hypotrachyna is an accessory component with evernic acid. 


126 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


variety of color tests and chemical constituents, soon realizes that 
the lichen acids are absolutley indispensable aids in species identi- 
fication. We have tested all specimens with the usual microchemical 
methods of Asahina and Shibata (1954). Detailed discussions of 
problematic or unidentified substances encountered will be delayed 
until a final monograph is completed. In the meantime, however, 
we have prepared a preliminary table showing the number of species 
within each section or subsection containing each of 19 identifiable 
lichen substances (table 1). Certain noteworthy features of this 
table will be discussed below under the various sectional headings. 

The color of the thallus and medulla is described for many of the 
species according to Ridgway’s Color Standards and Color Nomen- 
clature (1912); such colors are indicated by the notation R in paren- 


theses. 
Subgeneric Classification 


A classification for subgenus Parmelia is based on results from our 
own morphological and chemical studies and on reviews of previously 
proposed classifications, especially those of Vainio (1890, 1923), 
Gyelnik (1932), and Asahina (1952). 

1. Section Parmelia 
Subsection Parmelia 
Subsection Simplices Hale & Kurokawa 
2. Section Irregulares (Vain.) Vain. 
3. Section Imbricaria (Schreb.) Fr. 
Subsection Imbricaria 
Subsection Bicornutae (Lynge) Hale & Kurokawa 
Series Bicornutae 
Series Relicinae Hale & Kurokawa 
4. Section Cyclochetla (Vain.) Ras. 
5. Section Hypotrachyna Vain. 

Typification, descriptions, and discussions of these taxa are 
presented below under appropriate sectional headings. Keys to 
species in the major sections are included whenever possible in order 
to show the relationships of the new species to those already known. 
The keys do not include a small number of rare problematic species 
that have not yet been characterized. It is anticipated of course 
that more new species and revisions of names will be made in the final 
monograph. It should be emphasized that all species names in the 
keys are based on the actual study of holotype or lectotype specimens. 


Key to Sections 


1. Cortex pseudocyphellate, with or without effigurate maculae; rhizines simple 
Or squarrose .... 1.6. eee eee eee eee 1. Section Parmelia (p. 127) 


HALE & KUROKAWA—SUBGENUS PARMELIA 127 


1. Thallus lacking pseudocyphellae; rhizines simple, dichotomously or squarrosely 


branched. 
2. Lobes more or less rotund; upper cortex uniformly reticulately maculate 
or fissured to the margin . . . 2. Section Irregulares (Vain.) Vain. (p. 129) 


2. Lobes sublinear and truncate to subrotund; upper cortex continuous, 
fissured only on older lobes. 
3. Cilia or bulbate cilia present on margins of lobes. 
3. Section Imbricaria (Schreb.) Fr. (p. 130) 
3. Cilia absent. 
4, Rhizinessimple....... 4, Section Cyclocheila (Vain.) Ris. (p. 147) 
4. Rhizines dichotomously branched. 
5. Section Hypotrachyna Vain. (p. 159) 


Subgenus Parmelia 


Type species: Parmelia sazxatilis (L.) Ach. 

Thallus appressed to adnate, rarely subascending; lobes sublinear 
to subirregular, rarely broad and rotund, the margins entire and 
smooth, ciliate, or bulbate-ciliate; undersurface black or brown, 
uniformly rhizinate to the margins, or, in species with rotund lobes, 
with a narrow bare or papillate zone, the rhizines simple and un- 
branched, dichotomously branched, or squarrosely branched. 
Apothecia more or less adnate, rarely pedicellate, the disc imperforate 
(except in section Irregulares); spores simple, suborbicular to ellip- 
soid, usually less than 20 p» long. 

Parmelia has been proposed as a nomen genericum conservandum 
with P. sazatilis (L.) Ach. as the lectotype species (cf. International 
Code of Botanical Nomenclature, p. 220, 1961). This has been the 
unanimous choice as a lectotype for the genus (cf. Dodge, 1959), 
and we see no objections to it. Parmelia sazxatilis is therefore the 
type of subgenus and section Parmelia. 


1. Section Parmelia 


Thallus adnate; lobes sublinear, rarely subirregular and subrotund ; 
upper cortex pseudocyphellate, with or without effigurate maculae; 
lower surface black or brown, the rhizines simple or squarrosely 
branched. 

Imshaug (1957) placed section Hypotrachyna Vain. in synonymy 
under section Parmelia. As will be shown below, Vainio’s section 
is based on P. brasiliana Nyl., a tropical species with dichotomously 
branched rhizines and no pseudocyphellae. 

As we previously indicated, there are two quite distinct types of 
pseudocyphellae in this section, one typically elliptical and correlated 


128 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


with simple rhizines and the other effigurate-maculate and correlated 
with squarrosely branched rhizines. They are best treated as two 
subsections, Parmelia and Simplices, as follows: 


Subsection Parmelia 


Section Hypotrachyna subsection Myeloleuca Asah. (1952, p. 24). Lecto- 
type: P. saxatilis (L.) Ach. 

Upper cortex with effigurate maculae; lower surface black, the 
rhizines squarrosely branched or simple. 

Asahina’s subsection Myeloleuca included all species of subgenus 
Parmelia lacking medullary pigments. We feel that the most suitable 
choice of a lectotype for this extremely broad subsection would be 
P. sazxatilis. According to the International Code of Botanical 
Nomenclature (Art. 22), the rule of tautonomy applies only to the 
subgeneric and sectional ranks. Notwithstanding, we prefer to retain 
the spirit of the Code and use Parmelia as a subsectional epithet as 
well, even though Asahina’s subsection Myeloleuca is validly published 
and synonymous. 

As we have delimited this subsection, it contains about 30 species. 
The upper cortex is often ridged with the effigurate pseudocyphellae 
forming patterns along the ridges (cf. pl. 1, fig. 4). As far as is known, 
all species contain salacinic acid, except P. insensitiva (Magn.) Anders. 
and P. pseudosulcata Gyel., which contain protocetraric acid. Most 
of the species are confined to montane or subboreal areas in the 
Northern and Southern Hemispheres, with the greatest abundance in 
eastern Asia, where at least 15 species have been reported. Most of 
the Asian species have been summarized by Asahina (1952). There 
is a small group of Australian species, including P. tenuirima Tayl. 
and P. signata Nyl., which have not yet been fully investigated. The 
common American and European species P. fraudans Nyl., P. 
omphalodes (L.) Ach., P. saxatilis (L.) Ach., and P. sulcata Tayl. are 
well known. 


Subsection Simplices Hale & Kurokawa, subsect. nov. 


Thallus superne pseudocyphellatus, pseudocyphellis suborbiculari- 
bus vel ovalibus, subtus niger vel pallide castaneus, rhizinis 
simplicibus. 

Type species: Parmelia borreri (Sm.) Turn. 

This subsection contains at least 25 specics, but their taxonomy has 
not yet been completely worked out. They are found commonly in 
savannas or dry temperate forests. The pseudocyphellae are similar 
to those of P. cetrarioides (Duby) Nyl., an Amphigymnia species. 
The rhizines are always simple and quite long. The chemistry is 
extremely simple, the medulla reacting P—,C-+ rose or red (gyrophor- 


HALE & KUROKAWA—SUBGENUS PARMELIA 129 


ic or lecanoric acids), or C— (fatty acids). There is very strong 
correlation between the color of the lower surface and chemical 
constitution. All species with lecanoric acid, including the well- 
known P. subrudecta Nyl. (P. dubia (Wulf.) Schaer.) and P. rudecta 
Ach., have a pale brown lower surface; and all species with gyrophoric 
acid, such as P. borreri (Sm.) Turn. (P. pseudoborrert Asah.) and P. 
borrerina Nyl., have a jet-black lower surface. Those with fatty acids 
have either a pale (e.g., P. bolliana Mill. Arg. and P. canaliculata 
Lynge) or a black lower surface (e.g., P. appalachensis Culb. and 
P. microsticta Mill. Arg.) (cf. Culberson, 1962). 


2. Section Irregulares (Vain.) Vain. (1923, p. 34) 


Section Hypotrachyna * Irregularis Vain. (1890, p. 38). 
Section Reticulatae Du Rietz (1924b, p. 331). 
Lectotype species: Parmelia reticulata Tayl. 
Section Hypotrachyna subsection Irregulares (Vain.) Gyel. (1932, p. 224). 
Section Hypotrachyna subsection Irregulares (Vain.) Hillm. (1934, p. 188). 
Superfluous combination. 


Type species: Parmelia cetrata Ach. 

Thallus loosely adnate ; lobes subirregular, usually broad and rotund; 
upper cortex reticulately maculate and at maturity reticulately fissured 
to the margins; lower surface black, the rhizines simple or at maturity 
squarrosely branched. Apothecia more or less substipitate, the disc 
perforate or imperforate, 

Vainio first proposed Jrregulares in 1890 as a group name under 
section Hypotrachyna without a designated rank. The obvious lecto- 
type is Parmelia cetrata Ach. He proposed two other groups of 
equivalent rank, Cyclocheila (lectotype P. amazonica Nyl.) and 
Sublinearis (lectotype P. brasiliana Nyl.). These three groups were 
separated by subtle and overlapping differences in lobe configuration 
and presence of rhizines, papillae, or a narrow bare zone below along 
the margins. No lichenologist has successfully employed this classi- 
fication and most recent workers in fact have rejected it (cf. Asahina, 
1952; Maas Geesteranus, 1947). 

Almost all lichenologists except Rasinen (1943), however, have 
overlooked the fact that Vainio radically revised his own classification 
in a study of Philippine lichens (1923). He eliminated section 
Hypotrachyna and proposed in its place two sections, section Irregu- 
lares (including the former Cyclocheila and Trregularis) and section 
Sublineares. This realignment more or less successfully circumvented 
the difficulties of his former classification by recognizing only two 
general types of lobe configuration, subirregular and apically rotund 
and sublinear and truncate. 

Du Rietz’ section Reticulatae was sect up parallel with section 
Hypotrachyna Vain. and section Amphigymnia Vain. Du Rietz based 


130 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


the section not only on P. cetrata and P. reticulata but also on P. 
perforata (Jacq.) Ach., which is heavily maculate but lacks reticulate 
fissures. If P. perforata is excluded and placed in subgenus Am- 
phigymnia, section Reticulatae corresponds exactly with section 
Irregulares. 

This small section is close to Amphigymnia in lobation, but the 
lower surface is usually rhizinate or papillate to the margins. Fur- 
thermore, the rhizines are squarrosely branched at maturity, whereas 
the rhizines of Amphigymnia species are always simple and often 
sparse. There are about ten species in section Jrregulares, some very 
widely distributed in temperate and subtropical areas. The com- 
monly collected species are P. balansae Mill. Arg., P. cetrata Ach., 
P. homotoma Nyl., P. macrocarpoides Vain., P. pilosa Stein., P. 
reticulata Tayl., and P. subisidiosa (Mill. Arg.) Dodge. 


3. Section Imbricaria (Schreb.) Fr. (1825, p. 242) 
Lichen sect. Imbricarta Schreb. (1791, p. 767). 


Lectotype species: Parmelia tiliacea (Hoffm.) Ach. 

Thallus adnate; lobes sublinear to subirregular, the margins ciliate 
or bulbate-ciliate; lower surface black or brown, the rhizines simple 
or branched. Apothecia adnate, the exciple frequently coronate. 

Section Imbricaria was first proposed by Schreber under the genus 
Lichen and characterized as follows: “Syn. Squamaria Hoftm. Frondes 
subfoliaceae, membranaceac, imbricatae, depressae, flexiles. Fr. 
scutellae.” Squamaria is a genus set up by Hoffmann to accommodate 
the species now recognized as Cetraria pinastri (Scop.) Réhl. and C. 
juniperina (L.) Ach. Although Schreber cites only the Hoffmann 
name, we are not obliged to use it in typifying Imbricaria. It seems 
more appropriate to retain Imbricaria in the sense that it was so 
commonly used in the 19th century, that is, for the narrow lobed 
species of Parmelia. Acharius (1794) was the first to adopt Schreber’s 
name and included under it species now classified in the genera 
Physcia, Parmeliopsis, Xanthoria, and Parmelia. Of these, Parmelia 
tiliacea is the most suitable choice as a lectotype for section Imbricaria. 

Section Imbricaria may be divided into two subsections, Imbricaria 
and Bicornutae (p. 135), on the basis of whether marginal cilia or 
marginal bulbate cilia are present. 


Subsection Imbricaria 


Subsection Myelochroa Asah. (1952, p. 74). 
Lectotype species: Parmelia aurulenta Tuck. 


Thallus with simple marginal cilia, bulbate cilia lacking; lower 


surface usually black, the rhizines simple or sparsely branched or 
rarely squarrosely branched. Apothecia adnate, ecoronate. 


HALE & KUROKAWA—SUBGENUS PARMELIA 131 


In his study of the Japanese Parmelias, Asahina (1952) divided 
section Hypotrachyna Vain. into two subsections, Myelochroa (pig- 
ments present in the medulla) and AMyeloleuca (medulla white). We 
have already reduced subsection Myeloleuca to synonymy under sub- 
section Parmelia. The six pigmented species that Asahina included 
in subsection Myelochroa are all related to P. aurulenta Tuck. They 
all also have marginal cilia, a character that we consider to be of 
basic importance in classification. As far as we have been able to 
determine, medullary pigments have no value above the species level. 
Actually there are a number of pigmented species outside of Japan, 
unknown to Asahina at the time (e.g., P. isidiocera Nyl. and P. sil- 
vatica Lynge), which have branched rhizines and lack cilia and are 
classified below in section Hypotrachyna. 

Subsection Jmbricaria is a rather heterogeneous group of 33 species 
widely distributed in temperate and subtropical regions. The well- 
known species P. dissecta Nyl., P. quercina (Willd.) Vain., and P. 
tiliacea (Hoffm.) Ach. belong here. The distribution of chemical sub- 
stances is very similar to subsection Bicornutae (cf. table 1) and the 
almost complete absence of sorediate species is similar. ‘There seems 
to be no further natural subdivision of the subsection. 


Key to Species in Subsection Imbricaria 


1, Thallus isidiate or isidiate-lobulate. 

2. Isidia lobulate at maturity, dorsiventral. 
3. Medulla C+ rose (gyrophoric acid). . . 2. P. spathulata Kurokawa 
3. Medulla C—, KC+ rose (unknown)... . . . . P. horrescens Tayl. 

2. Thallus isidiate, the isidia cylindrical. 

4, Medulla pigmented yellow orange, K—. 
5. Upper surface distinctly maculate....... P. perisidians Ny]l. 
5. Upper surface not maculate. 
6. Thallus saxicolous; lobes sublinear, 1.0-1.5 mm. wide. 
P. obsessa Ach. 
6. Thallus corticolous; lobes subrotund, 2-5 mm. wide. 
P. lindmannii Lynge 
4. Medulla white. 
7. Medulla K+ yellow or red, C—. 
8. Medulla K+ persistent yellow; stictic acid present. 
P. internexa Nyl. 
8. Medulla K+ yellow turning red. 
9. Norstictic acid present; tropical America. . . P.antillensis Nyl. 
9. Salacinic acid present; Old World. 
10. Lobes subrotund, the marginal cilia short. 
P. wallichiana Tayl. 
10. Lobes sublinear, the marginal cilia often 1-3 mm. long. 
P. usambarensis Stein. & Zahlbr. 
7. Medulla K—, C+ rose or red or C—. 
11. MedullaC— . Le 


te ee ee . « « « « P. ikomae Asah. 
11. Medulla C+ rose or red. 


132 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


12. Cortex strongly maculate; medulla C+ red (lecanoric acid). 
P. tiliacea (Hoffm.) Ach. 
12, Cortex weakly maculate or emaculate; medulla C+ rose (gyro- 
phoric acid). 
13. Lobes subrotund, 2-5 mm. wide. 
1. P. melanochaeta Kurokawa 
13. Lobes sublinear, 1-2mm.wide. ... .. . P.dissecta Nyl. 
1. Thallus lacking isidia or lobules. 
14. Thallus pustulate; soredia scarcely if at all developed. 
15. Medulla white .... . . . . 3.P.subfatiscens Kurokawa 
15. Medulla pale yellow or orange yellow. 
16. Lobes 1-2 mm. wide; pustules fine, not sorediate; medulla C+ rose 
(gyrophoric acid) ..... rr P. spumosa Asah., 
16. Lobes 2-4 mm. wide; pustules coarsely sorediate; medulla C—. 
P. aurulenta Tuck. 
14. Thallus lacking pustules. 
17. Thallus sorediate. 


18. Medulla yellow orange. . ......... . P. aurulenta Tuck, 
18. Medulla white. 
19. Thallus yellowish green (usnic acid). . . . . P. nylanderi Lynge 


19. Thallus mineral gray (usniec acid lacking). 
20. Soredia subterminal; moniliform cells in the medulla. 
P. metarevoluta Asah. 
20. Soredia laminal; moniliform cells lacking . . . P. muelleri Vain. 
17. Thallus lacking soredia. 
21. Thallus pale brown below... . . . . P. versiformis Kremplh. 
21. Thallus jet black below. 
22, Medulla white (even under the apothecia). 
23. Medulla C+ blood red (lecanoriec acid). 
24, Lobes about 1 mm. wide... . . . P. pruinata Miill. Arg. 
24. Lobes 2-5 mm. wide. 
25. Thallus strongly maculate; apothecia with retrorse rhi- 
zines... . . . . . . P. carporrhizans Tayl. 
25. Thallus weakly maculate: retrorse rhizines Jacking. 
P. quercina (Willd.) Vain. 
23. Medulla C—, P+ orange red. 
26. Medulla K—, P+ (protocetraric acid). 
P. michauxiana Zahlbr. 
26. Medulla K+ red. 
27. Thallus fragile, the cortex flaking off; spores 7-8 » long; 
norstictic acid present . . . . .. . . P.phlyctinaHale 
27. Thallus coriaceous, the cortex firm; spores 20-28 » long; 
salacinic acid present. .... . . . P. mutata Vain, 
22. Medulla yellow or yellow orange (especially under the apothecia), 
28. Pigment K+ purple... . . .. . . P.denegans Nyl. 
28. Pigment K— or K+ more deeply ‘yellow. 
29. Lobes sublinear, 0.5-1.5 mm. wide. 
30. Medulla P-+ reddish (unknown); thallus firm. 
P. galbina Ach. 
30. Medulla P—; thallus fragile. 
P. xantholepis Mont. & v.d. Bosch. 
29. Lobes subrotund, 3-6 mm. wide. 


HALE & KUROKAWA—SUBGENUS PARMELIA 133 


31. Upper cortex without maculae; apothecia numerous; Mexi- 


CO... eee ee ee . . .. . P.immiscens Nyl. 
31. Upper cortex more or less distinctly maculate; Asia. 
32. Cortex fragile, flaking off . . . P. entotheiochroa Hue 
32. Cortex continuous, firm. 
33. Spores 7-124 long. ..... P. subaurulenta Nyl. 
33. Spores 13-17 long. ...... P. homogenes Nyl. 
1. Parmelia melanochaeta Kurokawa, sp. nov. PLATE 3 


Thallus laxe adnatus, olivaceco-albicans, 4-7 cm. latus, lobis ir- 
regularibus vel sublinearibus, subrotundatis, 2-6 mm. latis, margine 
crenatis, ciliatis, superne albomaculatus, isidiatus, subtus niger, 
rhizinosus, rhizinis nigris, simplicibus. Apothecia adnata vel sub- 
stipitata, 1-3 mm. diametro; hymenium 80-90 uw altum; sporae 
8-10 13-15 up. 

Thallus loosely attached to bark, turning olive buff or cream buff 
(R) in the herbarium, 4—7 cm. in diameter; lobes irregularly branched, 
irregular to sublinear, 2-6 mm. wide, 110-150 u thick, the margins 
crenate, ciliate, the cilia mostly simple, coarse, 1-2 mm. long (cf. pl. 1); 
upper surface shiny, rather distinctly white-maculate, moderately to 
densely isidiate, the isidia thin, cylindrical, often branched and with 
black spinules or short cilia; medulla white; undersurface black, dark 
brown in a rather wide zone near the tips, moderately rhizinate, the 
rhizines black to blackish brown, simple. Apothecia adnate to sub- 
stipitate, 1-3 mm. in diameter, amphithecium isidiate, spinulate, disc 
vandyke brown (R); hymenium 80-90 u high; spores 8-10 13-15 yp. 

Reactions: Thallus K+ yellow; medulla K—, C-++ rose, KC-+ red, 
P—, atranorine and gyrophoric acid present. 

Type in the Naturhistoriska Riksmuseet, Stockholm, collected at 
Santa Anna da Chapada, Mato Grosso, Brazil, Jan. 18, 1894, by 
G. A. Malme (no. 2243; isotype in US). 

This is an unusual species that could be mistaken for P. dissecta 
Nyl. In fact, Lynge identified the same material as P. minarum 
Vain., a synonym of P. dissecta (cf. Hale, 1960). Although the 
chemistry of the two species is identical, P. melanochaeta is much 
larger and loosely attached and has broad rotund lobes. It seems 
to be common in southern Brazil and adjacent Paraguay. 

Additional specimens examined: Brazil: Buriti, Cerra da Chapada, 
Malme 2243, 2397B (S); Corumb4, Mato Grosso, Malme, August 
1894 (S). Paraguay: Gran Chaco, Malme, Sept. 14, 1893 (S); 
Colonia Risso, Malme 1862 Aa*, 1862 Ba*, 1911 Ba (S). 


2. Parmelia spathulata Kurokawa, sp. nov. 

Thallus adnatus, corticola, olivaceo-albicans, 2-5 cm. diametro, 
lobis sublinearibus, plus minusve imbricatis, 1-3 mm. latis, margine 
dissectis, superne sparse ciliatis, nitidus, emaculatus, isidiato-lobulatus, 


134 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


lobulis dorsiventralibus, apice dilatatis, subtus niger, rhizinosus, 
rhizinis nigris, sparse ramosis. Apothecia ignota. 

Thallus adnate, growing over mosses on bark, pale glaucous green 
(R), 2-5 cm. in diameter; lobes irregularly branched, subimbricate, 
1-3 mm. wide, 90-130 uw thick, the margins sparsely ciliate, dissected; 
upper surface shiny, without maculae, covered with numerous small 
isidioid lobules, the lobules dorsiventral, subascending, often branched 
and dilatated towards the apices; medulla white; undersurface black, 
rhizinate, the rhizines black, shiny, simple to sparsely branched. 
Apothecia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C-++ rose, KC+ red, 
P—, atranorine and gyrophoric acid present. 

Type in the Botanical Museum, Lund University, collected north- 
west of Tea-room, Skeleton Gorge, Wynberg, Cape Province, Union 
of South Africa, July 18, 1953, by Ove Almborn (no. 305; isotype in 
US). 

This species is characterized by having numerous spathulate lobules 
and simple to sparsely branched rhizines. It is closely related to 
P. dissecta Nyl., which has similar chemistry but cylindrical isidia 
rather than lobules. Parmelia spathulata is apparently restricted to 
South Africa. 

Additional specimens examined: Union of South Africa: Polela 
Forest, Polela, Natal, Almborn 9516 (LD, US); Kirstenbosch, Cape 
Province, Arnell 437 (LD). 

3. Parmelia subfatiscens Kurokawa, sp. nov. 

Thallus adnatus, albido-cinerascens, 4-7 cm. latus, lobis subline- 
aribus, 0.5-1.5 mm. latis, margine ciliatis, superne planus, emaculatus, 
pustulatus, subtus niger, dense rhizinosus, rhizinis nigris, simplicibus. 
Apothecia adnata, 1.5-4.0 mm. diametro; hymenium 60-75 y altum; 
sporae 8-9 12-14 uy. 

Thallus adnate on bark, pale olive gray to light mineral gray (R), 
4—7 cm. in diameter; lobes dichotomously or irregularly branched, 
sublinear-elongate, 0.5-1.5 mm. wide, 80-100 u thick, the margins 
crenate, ciliate, the cilia black, simple, 0.5-1.0 mm. long; upper 
surface plane and smooth, shiny, not maculate, pustulate, the pustules 
not becoming sorediate, often ciliate; medulla white; undersurface 
black, densely rhizinate, the rhizines black, simple, about 1.0 mm. 
long. Apothecia adnate, 1.5-4.0 mm. in diameter, margins smooth, 
disc clove brown (R), radially split; hymenium 60-75 u high; spores 
8-9 X 12-14 up. 

Reactions: Thallus K-+- yellow; medulla K—, C—, KC+ rose, 
P—, atranorine and an unknown KC-+ substance present. 

Type in the Botanical Museum, Lund University, collected at 
Punch Bow! Inn, north of Louis Trichardt, Zoutpansberg, Transvaal, 


HALE & KUROKAWA—SUBGENUS PARMELIA 135 


Union of South Africa, Oct. 10, 1953, by Ove Almborn (no. 6443; 
isotype in US). 

This species is very close to P. dissecta Nyl. but it is distinctly 
pustulate rather than isidiate and the medulla is C—, whereas P. 
dissecta is C+ rose (gyrophoric acid). The pustules are similar to 
those of P. spuwmosa Asah., a smaller species with a pale yellow 
medulla and gyrophoric acid. Parmelia subfatiscens has the same 
West Indies-Africa distribution characteristic of P. exsplendens Hale, 
P. suffixa Stirt., and P. ventricosa Hale & Kurokawa. 

Additional specimens examined: Jamaica: Blue Mountains: North 
slope below Corn Puss, Jmshaug 14550 (MSC); New Haven Gap, 
Imshaug 15137 (MSC, US). Union of South Africa: Natal: Bosch- 
fontein Forest, Lions River Distr., Almborn 8717 (LD); Transvaal: 
Forest Drive from Houtbosch to Tzaneen, Pietersburg, Almborn 
6797 (LD); Cape Province: Deepwalls, Knysna, Almborn 3433 (LD). 

Subsection Bicornutae (Lynge) Hale & Kurokawa, comb. noy. 
Section Bicornuta Lynge (1913, p. 17). Type species: P. semilunata Lynge. 
Subgenus Bicornuta (Lynge) Gyel. (1932, p. 219). 

Thallus with marginal bulbate cilia; lower surface black or brown, 
the rhizines simple or branched. Apothecia frequently coronate. 

Lynge first proposed section Bicornuta on the basis of two species, 
P. semilunata and P. schiffnert Zahlbr., both of which have peculiar 
bicornute spores. As a matter of fact, there is apparently only one 
other species in Parmelia with such spores, P. bicornuta Mill. Arg. 
However, these species also have marginal bulbate cilia, one of the 
most fundamental characters that we have found in subgenus Parmelia. 
Normally ciliate species (subsection Imbricaria) appear to be closely 
related in chemistry (see table 1). 

This subsection of 46 primarily tropical species may be further 
subdivided into two series, based on the presence or absence of usnic 
acid, as follows: 


Series Bicornutae 

Thallus mineral gray; usnic acid lacking. 

Series Relicinae Hale & Kurokawa, ser. nov. 

Thallus stramineus vel viridiflavicans, acidum usnicum continens. 

Type species: Parmelia relicina Fr. 

When the bulbate-ciliate species are divided on the presence or 
absence of usnic acid, there is significant geographical correlation. 
In the series Bicornutae seven of the nine species with salacinic acid 
occur only in Africa. Of the nine C+ species, six are restricted to 
tropical America. Among the remaining species, five occur only in 
South America, two in Africa, one in America-Africa, and one 

722-891—64——_2 


136 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


in Asia. In other words, the species in this series are most common 
in Africa and South America. 

In the series Pelicinae there are no species at allin Africa. Thirteen 
occur only in southeastern Asia and Australia, two in tropical America 
and the Pacific, two in tropical America only, and one in America- 
Asia. This series then shows very strong representation in south- 
eastern Asia. 


Key to Species in Subsection Bicornutae Series Bicornutae 


1. Thallus isidiate or lobulate-isidiate. 
2. Lower surface pale brown. 
3. Medulla K+ red (salacinic acid). 
4, Thallus large, the lobes 2-5 mm. wide; upper surface not distinctly 


maculate.... . .. .. P. isidiza Nyl. 

4. Thallus smaller, the lobes 1. 0-2. 5 mm, “wide; upper surface distinctly 
maculate .... . . . P. subglandulifera Hue 

3. Medulla K—, C+ rose (gyrophorie acid). ... . . P. scortella Nyl. 


2. Lower surface jet black. 
5. Isidia becoming dorsiventral, lobulate; medulla C+ rose (gyrophoric 


acid), 
6. Apothecia ecoronate. .......... +... . =. P. suffixa Stirt. 
6. Apothecia coronate............ . .P. fungicola Lynge 


5. Isidia normal, cylindrical. 
7. Medulla K-+ red. 


8. Norstictic acid present. . . . .6. P. ventricosa Hale & Kurokawa 
8. Salacinic acid present. 
9. Plants saxicolous. . . .... . . 4 P. decurtata Kurokawa 
9. Plants corticolous ... . . .P. tabacina Mont. & v. d. Bosch. 


7. Medulla K—, 
10. Medulla C—, KC+ red (unknown present). 
2. P. apophysata Hale & Kurokawa 
10. Medulla C+ rose or red. 
11. Medulla C+ deep red (lecanorie acid) . . . P. laevigatula Nyl. 
11. Medulla C+ rose (gyrophoric acid). 
12. Thallus tightly adnate, the lobes 0.5-1.0 mm. wide; Asia. 
P. subdissecta Ny]. 
12. Thallus loosely adnate, the lobes 1.0-3.0 mm. wide; tropical 


America. . ..... +... . «ss. . . P. papyrina Fée 
1. Thallus without isidia. 


13. Lower surface pale brown. 
14. Apothecia coronate; norstictic acid present . P. subcoronata Miill. Arg. 
14. Apothecia ecoronate; salacinic acid present. 
15. Upper surface distinctly maculate . ... . . .P. hypocraea Vain. 
15, Upper surface dull, not maculate. 
16. Thallus small, the lobes 0.7-2.0 mm. wide; South America, 
P. continua Lynge 
16. Thallus medium sized, the lobes 2.0-5.0 mm. wide; Asia. 
P. setschwanensis Zahlbr. 
13. Lower surface jet black. 
17. Medulla K+ red (salacinic acid). 


HALE & KUROKAWA—SUBGENUS PARMELIA 137 


18. Upper surface coarsely pustulate, pustules breaking open, esorediate. 
5. P. pustulata Hale 
18. Upper surface smooth, pustules lacking. 
19. Spores 7-9 uw long; Africa. . . P. sensibilis Stein. & Zahlbr. 
19. Spores 14-19 uw long; eastern Asia . . .P. meizospora (Nyl.) Nyl. 
17. Medulla K—. 
20. Medulla C—. 
21. Thallus adnate, the lobes 1-2 mm. wide. 
22. Thallus coriaceous; bulbate cilia conspicuous; India. 
3. P. bulbochaeta Hale 
22. Thallus fragile; bulbate cilia small; South America. 
P. viridescens Lynge 
21. Thallus appressed, the lobes less than 1 mm. wide. 
23. Lobes 0.3-1.0 mm. wide; spores elliptical. 
1. P. affixa Hale & Kurokawa 
23. Lobes 0.1-0.2 mm. wide; spores bicornute. 
24. Spores 4-5ywlong........ . . .P.schiffmeri Zahlbr. 
24. Spores 15-20 w long. . . . .. =... .P.semilunata Lynge 
20. Medulla C+ rose or red. 
25. Medulla C+ deep red (lecanoric acid). 
26. Spores bicornute. . .... .. . . .P. bicornuta Mill. Arg. 
26. Spores elliptical . .. . . . P. confoederata Culb. 
25. Medulla C+ rose (eyrophoric acid). 
27. Apothecia coronate; upper cortex without maculae P. coronata Fée 
27. Apothecia ecoronate; upper cortex distinctly maculate. 
P. atrichella Nyl. 
1. Parmelia affixa Hale & Kurokawa, stat. et nom. nov. 
Parmelia coronata Fée var. denudata Vain. in Welw. Cat. Afr. Pl. 2, no. 2: 401. 
1901. Type collection: Morro de Lopollo, Huilla, Angola, Welwitsch 
33 (TUR, holotype; BM, isotype). 

Thallus very closely adnate on bark, turning olive buff to tea green 
in the herbarium, about 3 cm. in diameter; lobes dichotomously 
branched, sublinear-elongate, 0.3-1.0 mm. wide, 70-904 thick, the 
margins more or less crenate, with dense bulbate cilia; upper surface 
plane, shiny, emaculate, cracked on older lobes, isidia and soredia 
lacking; medulla white; undersurface black, densely rhizinate, rhi- 
zines black, shiny, simple. Apothecia numerous, adnate, 0.5-1.5 
mm. in diameter, exciple smooth to crenate, coronate, the base 
retrorsely rhizinate; hymenium 40-50 y high; spores 4-58-10 yn, 
episporium 1 p» thick, 

Reactions: Thallus K+ yellow; medulla K—,C—, KC+ rose, P—, 
atranorine and a KC-+ rose unknown substance (not further identified 
because of paucity of material). 

The name Parmelia denudata has already been used by Hampe 
(Linnaea 17:121. 1843) for a species now recognized as Everniopsis 
trulla (Ach.) Nyl. Parmelia affirais a small easily overlooked species 
that is probably more common than suspected in southwest Africa. 


It is one of the few coronate species in Africa. 


138 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Additional specimen examined: Africa: 10 km, north of S& da 
Bandeira, Huila, Angola, Degelius, Mar. 2, 1960 (DEGEL, US). 


2. Parmelia apophysata Hale & Kurokawa, sp. nov. 

Thallus arcte adnatus, corticola, albido-olivascens, 3-6 cm. dia- 
metro, lobis sublinearibus, 0.7-1.5 mm. latis 90-130 u crassis, margine 
bulbato-ciliatis, superne nitidus, isidiatus, isidiis cylindricis, subtus 
niger, rhizinosus, rhizinis dichotome ramosis. Apothecia ignota. 

Thallus closely adnate on bark, turning light olive buff in the her- 
barium, 3-6 em. in diameter; lobes sublinear-elongate, more or 
less dichotomously branched, 0.7-1.5 mm. wide, 90-130 u thick, 
the margins densely bulbate-ciliate; upper surface plane, shiny, 
sometimes faintly maculate, often irregularly cracked on older lobes, 
sparsely to moderately isidiate, isidia simple, cylindrical, less than 
0.3 mm. high; undersurface black, with a broad pale brown zone 
near the tips of lobes, short rhizinate, rhizines! pale to black, shiny, 
densely branched. Apothecia unknown. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ reddish, 
P—, atranorine and an unknown KC-+ substance (near alectoronic 
acid?) present. 

Type in the U.S. National Herbarium, collected on mangrove tree 
in pine barrens, vicinity of Piedra Blanca, La Vega, Dominican 
Republic, Oct. 14, 1947, by H. A. Allard (no. 16078). 

Parmelia apophysata is characterized by having a medium-sized 
thallus, simple cylindrical isidia, bulbate cilia and branched rhizines, 
and faintly maculate lobes. The unknown KC+ substance is notice- 
ably fluorescent in ultraviolet light. This species resembles P. sub- 
dissecta Nyl., which has more adnate smaller lobes and a C+ pale-red 
reaction caused by gyrophoric acid. Parmelia apophysata is known 
only from the type collection but it is certain to be found in other 
collections from tropical America. 

3. Parmelia bulbochaeta Hale, sp. nov. PuaTE 1 

Thallus adnatus vel laxe adnatus, albido-cinerascens, coriaceus, 
4-8 cm. diametro, lobis sublinearibus, 1.5-2.5 mm. latis, margine 
bulbato-ciliatis, superne planus, nitidus, emaculatus, isidiis sorediisque 
destitutus, subtus nigricans, dense rhizinosus, rhizinis nigricantibus, 
dichotome ramosis. Apothecia adnata, coronata, 2-3 mm. diametro; 
hymenium 40-50 p altum; sporae 45 u. 

Thallus adnate to loosely adnate, dark mineral gray, coriaceous, 
4—8 cm. in diameter; lobes sublinear, 1.5-2.5 mm. wide, 160-210 u 
thick, the margins entire, conspicuously bulbate-ciliate, bulbules with 
simple or branched apical cilia; upper surface plane, shiny, without 
maculae, usually heavily pycnidiate, soredia and isidia lacking; 
medulla white, dense; undersurface black, becoming brown at the 


HALE & KUROKAWA—SUBGENUS PARMELIA 139 


tips, densely rhizinate, rhizines black, brownish black near the 
margins, dichotomously branched. Apothecia adnate, 2-3 mm. in 
diameter, coronate; hymenium 40-50 uv high; spores 45 up. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC—, P-, 
only atranorine present. 

Type in the herbarium of D. D. Awasthi, collected at Perumal 
coffee plantation, Shembaganur, Madurai district, India, elev. 5300- 
5700 ft., Dec. 23, 1959, by D. D. Awasthi (no. 4347; isotype in US). 

Parmelia bulbochaeta resembles the South American species P. 
coronata Fée and P. wiridescens Lynge in the presence of coronate 
apothecia, black undersurface, and branched rhizines, and in the 
absence of maculae. It differs in having a more robust coriaceous 
thallus and more conspicuous bulbate cilia and no chemicals other 
than atranorine. It is known only from two collections made in 
South India. 

Additional specimen examined: College compound, Shembaganur, 
Madurai, India, Awasthi 4275 (AWAS). 

4. Parmelia decurtata Kurokawa, sp. nov. 

Thallus adnatus, saxicola, olivaceo-cinerascens, 3-7 cm. latus, lobis 
subirregularibus, subimbricatis, 1-3 mm. latis, margine crenatis, 
bulbato-ciliatis, superne planus, nitidus, emaculatus, isidiatus, subtus 
nigricans, rhizinosus, rhizinis nigris, simplicibus. Apothecia ignota. 

Thallus closely adnate on rock, pale olive buff to pale olive gray 
(R), 3-7 cm. in diameter; lobes irregularly branched, subirregular to 
sublinear, more or less imbricate, 1-3 mm. wide, 220-300 yu thick, the 
margins crenate, moderately bulbate-ciliate; uppersurface plane, 
shiny, often cracked on older lobes, isidiate, the isidia cylindrical, 
often more or less granular, rarely branched; medulla white; under- 
surface dark brown to black, moderately to densely rhizinate, the 
rhizines black, simple, about 1 mm. long. Apothecia not seen. 

Reactions: Thallus K+ yellow; medulla K+ yellow turning red, 
C—, KC—, P+ pale orange red, atranorine and salacinic acid present. 

Type in the Botanical Museum, Lund University, collected 10 
miles southeast of Lydenburg, Transvaal, Union of South Africa, 
Oct. 20, 1953, by Ove Almborn (no. 7388; isotype in US). 

Parmelia decurtata bears a very close resemblance to P. tabacina 
Mont. & v.d. Bosch., a widespread corticolous species. They both 
have isidia, bulbate cilia, and salacinic acid. Parmelia decurtata 
differs in being strictly saxicolous, lacking any trace of maculae, and 
having smaller, often papillate, black-tipped isidia. 

Additional specimens examined: Union of South Africa: Natal: 
Cathedral Peak Area, Bergville Distr., elev. 6000 ft., Almborn 9328 


140 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


(LD); Cape Province: Bains Kloof, Wellington Distr., Almborn 5053 
(LD). 
5. Parmelia pustulata Hale, sp. nov. 

Thallus adnatus, corticola, membranaceus, 6-10 cm. diametro, 
olivaceo-albicans, lobis subirregularibus, 4-6 mm. latis, apice subro- 
tundatis, margine bulbato-ciliatis, superne planus vel rugosus, nitidus, 
isidiato-pustulatus, pustulis inflatis, subtus niger, modice rhizinosus, 
rhizinis simplicibus, ambitu castaneus, nudus vel papillatus. Apo- 
thecia ignota. 

Thallus adnate on bark, membranaceous, 6-10 cm. in diameter, 
olivaceous to light mineral gray; lobes more or less subirregular, 4—6 
mm. wide, the apices subrotund, the margins broadly crenate, in- 
conspicuously bulbate-ciliate; upper surface plane to rugose, shiny, 
becoming moderately isidiate-pustulate, pustules inflated, irregular, 
breaking open apically, soredia absent; medulla white; lower surface 
black and moderately rhizinate at the center, the rhizines black, 
simple, the marginal zone bare or papillate, brown. Apothecia not 
seen. 

Reactions: Thallus K+ yellow; medulla K+ yellow turning red, 
C—, KC—, P+ pale orange red, atranorine and salacinic acid present. 

Type in the U.S. National Herbarium, collected in a bamboo forest, 
Urundi, Africa, elev. 2200 m., by Deuse (communicated by Dr. J. L. 
Ramaut, Liége, Belgium). 

This species is unique in subsection Bicornutae in having large 
distinct pustules. Isidiate P. tabacina Mont. & v.d. Bosch. has similar 
lobe configuration and chemistry and is probably related to it. Par- 
melia pustulata is known only from the type collection. 

6. Parmelia ventricosa Hale & Kurokawa, stat. et nom. nov. 

Parmelia isidiza var. domingensis Vain. Ann. Acad. Sci. Fenn. 6, no. 7:17. 
1915. Type collection: La Cumbra, Santo Domingo (Dominican Re- 
public), Raunkiaer 492 (TUR, holotype; C, isotype). 

Thallus adnate on bark, pale glaucous green, 3-5 cm. in diameter; 
lobes irregularly branched, 1-5 mm. wide, 65-80» thick; margins 
crenate, with bulbate cilia; upper surface plane and smooth, shiny, 
weakly maculate, more or less wrinkled on older lobes, isidiate, the 
isidia mostly simple, fine and short; medulla white; undersurface 
uniformly black, short-rhizinate, the rhizines black, simple. Apo- 
thecia adnate, coronate, more than 2.5 mm. in diameter, exciple 
crenate, amphithecium smooth, weakly maculate, disc russet, imper- 
forate; hymenium 50-60 uw high; spores 7-8><13-17 yu, the episporium 
1.0-1.5 pw thick. 

Reactions: Thallus K-++ yellow; medulla K+ yellow turning red, 
C—, KC—, P+ pale orange, atranorine and norstictic acid present. 


HALE & KUROKAWA—SUBGENUS PARMELIA 141 


The name Parmelia domingensis is preoccupied by P. domingensis 
Ach., which is now recognized as a species of Anaptychia. Parmelia 
ventricosa and P. rsidiza Nyl. share isidia and bulbate cilia, but P. 
ventricosa has coronate apothecia and norstictic acid. Parmelia 
isidiza, a common African lichen, has ecoronate apothecia and salacinic 
acid. Parmelia ventricosa is a rather rare species in the Caribbean 
region with a disjunct locality in South Africa. 

Additional specimens examined: Mexico: Chiapas: Near ranch, El 
Suspiro, Hale 20166 (US). Union of South Africa: Transvaal: Drive 
from Houtbosch to Tzaneen, Pietersburg, Almborn 6813 (LD). 


Key to Species in Subsection Bicornutae Series Relicinae 


1. Thallus isidiate or isidiate-lobulate. 
2. Isidia dorsiventral, lobulate. 
3. Medulla P+ red (unknown)... .. . 7. P. schizospatha Kurokawa 
3. Medulla P—, C+ rose (gyrophoric acid) . 4. P. luteoviridis Kurokawa 
2. Isidia normal, cylindrical. 
4, Lower surface pale to dark brown. 
5. Upper surface maculate; rhizines simple. 
9. P. tumescens Hale & Kurokawa 
5. Upper surface emaculate; rhizines branched . P. cireumnodata Nyl. 
4, Lower surface jet black. 
6. Lobes more or less subirregular, to 3 mm. wide; medulla P+, K— 
(unknown). ........... . 5. P. planiuscula Kurokawa 
6. Lobes sublinear, 1-2 mm. wide; medulla P+, K+ red (norstictie acid). 
P. abstrusa Vain. 
1. Isidia and lobules lacking. 
7. Lower surface pale to dark brown. 
8. Base of apothecia retrorsely rhizinate; plants saxicolous; stictic acid 


present. .... . . . . . P.limbata Laur. 
8. Base of apothecia without retrorse rhizines: ; plants corticolous; stictic acid 
lacking. 


9. Rhizines black; protocetraric acid present. 
6. P. ramosissima Kurokawa 
9. Rhizines uniformly pale brown. 
10. Medulla K+ red (salacinic acid). . .1. P. acrobotrys Kurokawa 
10. Medulla K— (protocetraric acid) . . . 8 P. sublanea Kurokawa 
7. Lower surface jet black. 
11. Medulla P—, K—. 
12. Lobes appressed, 0.2-0.8 mm. wide. . . P.relicinula Miill. Arg. 
12. Lobes adnate, 0.7-2.0 mm. wide. 
13. Rhizines mostly branched; protolichesteric acid present. 
2. P. connivens Kurokawa 


13. Rhizines mostly simple; fatty acids absent. . . .P. relicina Fr. 
11. Medulla P+, K+ red or K—. 
14. Medulla K+ red (norstictic acid)... . P. subabstrusa Gyel. 


14. Medulla K—. 
15, Apothecia ecoronate; P+ unknown present . P. sublimbata Ny]. 
15. Apothecia coronate; fumarprotocetraric acid present. 


142 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


16. Asci 16-32-spored. . .... 2... 0... P. relicinella Nyl. 
16. Asci 8-spored. 
17. Marginal bulbules ciliate, 0.3-1.0 mm. long; Pacific area. 
P. samoensis Zahlbr. 
17. Marginal bulbules not ciliate, 0.1-0.2 mm. long; tropical 


America. . . 3. P. eximbricata (Gyel.) Hale & Kurokawa 


1. Parmelia acrobotrys Kurokawa, sp. nov. PLATE 2 


Thallus adnatus, viridi-flavicans, 6-12 cm. diametro, lobis subline- 
aribus, 1.0-2.5 mm. latis, margine bulbato-ciliatis, superne plerumque 
convexus, albomaculatus, isidiis sorediisque destitutus, subtus pallide 
castaneus, modice rhizinosus, rhizinis dense ramosis, pallidis. Apo- 
thecia substipitata, 1.0-3.5 mm. diametro; hymenium 50-60 y» altum; 
sporae 4-5 7-8 uy. 

Thallus adnate on bark, olive buff (R), 6-12 cm. in diameter; lobes 
dichotomously branched, sublinear-elongate, 1.0-2.5 mm. wide, 240- 
320 uw thick, the margins smooth, with bulbate cilia; upper surface 
more or less convex, shiny, maculate, isidia and soredia lacking; me- 
dulla white; undersurface pinkish buff (R), moderately rhizinate, 
rhizines densely branched towards the tips, pale. Apothecia sub- 
stipitate, without retrorse rhizines, 1.0-3.5 mm. in diameter, exciple 
crenate, amphithecium smooth, disc burnt amber (R); hymenium 50- 
60 » high; spores 4-5 X7-8 p. 

Reactions: Thallus K+ yellowish; medulla K+ yellow turning red, 
C—, KC—, P+ pale orange red, atranorine, usnic acid, and salacinic 
acid. 

Type in Herbarium Bogoriense, collected at Tjibodas, Java, May 
1949, by Neervoort (no. 60 pr. p.; isotype in US). 

Parmelia acrobotrys is closely related to P. sublanea Kurokawa. 
Both species have a pale undersurface, pale brown rhizines, ecoronate 
apothecia, and similar spores. Parmelia acrobotrys has coarser rhi- 
zines and contains salacinic acid, whereas P. sublanea contains proto- 
cetraric acid. The saxicolous species P. limbata Laur. is superficially 
similar but contains stictic acid. Parmelia acrobotrys is known only 
from the type collection. 


2. Parmelia connivens Kurokawa, sp. nov. PLATE 2 


Thallus adnatus, viridiflavicans, 3-5 cm. diametro, lobis subline- 
aribus, 1.0-2.5 mm. latis, margine bulbato-ciliatis, superne planus vel 
convexus, sorediis et isidiis destitutus, subtus niger, dense rhizinosus, 
rhizinis nigris, ramosis. Apothecia adnata, 1.0-2.5 mm. diametro, 
retrorso-rhizinata; hymenium 45-60 u altum; sporae 2-3 «4-6 up. 

Thallus adnate, deep sea-foam green (R), turning cinnamon buff 
(R) in the herbarium, 3-5 cm. in diameter; lobes dichotomously 
branched, sublinear-elongate, 1.0-2.5 mm. wide, 120-160 u thick, the 


HALE & KUROKAWA—SUBGENUS PARMELIA 143 


older lobes becoming lobulate, the margins smooth, narrowly black- 
rimmed near the tips, with bulbate cilia; upper surface plane to 
slightly convex, smooth but becoming rugulose and cracked on older 
lobes, soredia and isidia lacking; undersurface almost uniformly black, 
densely rhizinate, the rhizines black, irregularly branched, sometimes 
projecting beyond the margins. Apothecia adnate, 1.0-2.5 mm. in 
diameter, retrorsely rhizinate, exciple crenate, disc seal brown (R); 
hymenium 45-60 » high; spores 2-3 X4-6 u. 

Reactions: Thallus K+ yellowish; medulla K—, C—, KC—, P-, 
atranorine, usnic acid, and protolichesteric acid present. 

Type in the Kew Herbarium, collected in the Milne Bay district, 
Papua, New Guinea, Sept. 11, 1954, by R. D. Hoogland (no. 4726; 
isotypes in L, US). 

At first glance this species resembles P. acrobotrys Kurokawa and 
P. ramosissima Kurokawa but it is easily distinguished by the black 
undersurface and the negative reaction with P. No other species in 
series felicinae produces a fatty acid. It is a typical Indonesian 
species. 

Additional specimens examined: Molucca Islands: Tanimbar Island, 
P. Jamdena, Buwalda 4246 (BO). Guam: McGregor s.n. (H). 


3. Parmelia eximbricata (Gyel.) Hale & Kurokawa, comb. nov. 


Parmelia samoensis Zahlbr. var. eximbricata Gyel. Ann. Mycol. 36:288. 
1938. Type collection: Monte Rus, Cuba, Hioram 10506 (BP, holotype). 


Thallus closely adnate on bark, massicot yellow (R), 3-5 cm. in 
diameter; lobes more or less dichotomously branched, sublinear, 
0.7-1.5 mm. wide, 160-200 thick; margins crenate, with small 
bulbate cilia, 0.1-0.2 mm. long; upper surface plane, smooth and 
shiny emaculate, isidia and soredia lacking; medulla white; under- 
surface black, densely rhizinate, the rhizines black, simple. Apothecia 
common, adnate, 0.5-1.5 mm. in diameter, exciple coronate, the base 
of the amphithecium often retrorsely rhizinate; hymenium 45-55 yu 
high; spores 4-5 7-9 yu, the episporium 1 yw thick. 

Reactions: Thallus K—; medulla K—, C—, KC—, P+ brick red, 
fumarprotocetraric acid and usnic acid present. 

Parmelia eximbricata resembles the Pacific species P. samoensis 
Zahlbr. in having coronate apothecia and fumarprotocetraric acid. 
The marginal bulbate cilia of P. samoensis however are longer and 
more conspicuous, and tipped with distinct cilia. This species is 
widely distributed in the Caribbean as far north as southern Florida, 
usually occurring at low elevations. 

Additional specimens examined: Florida: Big Pine Key, Florida 
Keys, Swanson & Dowling C-795 (US). Jamaica: Wareika House, 
summit of Long Mountain, St. Andrew Imshaug 13492 (MSC); north 


144 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


slope of Montpelier, St. Catherine, Jmshaug 14261 (MSC, US). 
Grand Cayman: Near Georgetown, Jmshaug 24385 (MSC); center of 
Island, Jmshaug 24518 (MSC, US). Dominican Republic: Vicinity 
of Ciudad Trujillo, Santo Domingo, Allard 16186, 16192 (US); near 
Rio Chavén, east of La Romana, La Altagracia, Hassler, July 31, 
1935 (WISC). Cuba: Without locality, Wright, Lich. Cub. ser. II, 
10 (US). 


4. Parmelia luteoviridis Kurokawa, sp. nov. PLATE 2 


Thallus adnatus, corticola, viridiflavicans, ca. 7 cm. diametro, 
lobis sublinearibus, 1.5-3.0 mm. latis, margine dense lobulatis, 
bulbato-ciliatis, superne nitidus, lobulatus, lobulis vulgo ramosis, 
subtus niger, rhizinosus, rhizinis nigris, sparse ramosis. Apothecia 
ignota. 

Thallus adnate on bark, deep sea-foam green to light dull green 
yellow (R), about 7 cm. in diameter; lobes dichotomously branched, 
sublinear-elongate, 1.5-3.0 mm. wide, 160-200 thick, the margins 
lobulate, with shiny bulbate cilia; upper surface plane and smooth, 
shiny, faintly maculate, lobulate, the lobules dorsiventral, not as- 
cending, often dichotomously branched, rhizinate below; undersurface 
black, rhizinate, the rhizines black, shiny, sparsely branched. Apoth- 
ecia not seen. 

Reactions: Thallus K+ yellowish; medulla K—, C+ rose, KC+ 
red, P—, atranorine, usnic acid, and gyrophoric acid. 

Type in the Herbarium Bogoriense, collected on peak of B. Papan, 
Beu territory, E. Kutai, Borneo, July 8, 1952, by W. Meijer (no. 
B1948; isotype in US). 

This species is characterized by the conspicuous yellow richly lobu- 
late thallus and by the production of gyrophoric acid. It is the only 
species in series Felicinae with a C++ substance. It resembles P. schizo- 
spatha Kurokawa in having lobules and a black lower surface but 
has wider lobes and greater development of lobules. Parmelia schizo- 
spatha is C—,P+ red. Parmelia luteoviridis is still known only from 
the type locality. 

Additional specimen examined: Borneo: Same data as the holotype, 
Meijer B1548a (BO). 


5. Parmelia planiusecula Kurokawa, sp. nov. 


Thallus adnatus, corticola, viridiflavicans, usque ad 9 cm. latus, 
lobis sublinearibus, 1-3 mm. latis, margine subcrenatis, bulbato- 
ciliatis, superne planus, nitidus, sparse isidiatus, subtus niger, modice 
rhizinosus, rhizinis nigris, simplicibus. Apothecia ignota. 

Thallus adnate to loosely attached over mosses on bark, rather 
rigid, turning deep olive buff (R) in the herbarium, about 9 cm. in 
diameter; lobes irregularly branched, sublinear-elongate, 1-3 mm. 


HALE & KUROKAWA—SUBGENUS PARMELIA 145 


wide, 150-190 u thick, the margins more or less crenate, with bulbate 
cilia; upper surface plane and smooth, shiny, faintly maculate, sparsely 
isidiate, isidia simple, cylindrical, less than 0.7 mm. high; medulla 
white; undersurface black, moderately rhizinate, the rhizines black, 
shiny, simple. Apothecia not seen. 

Reactions: Thallus K+ yellowish; medulla K—, C—, KC—, P+ 
orange red, atranorine, usnic acid, and an unknown P+ substance. 

Type in the Herbarium Bogoriense, collected at Kandang Badak, 
Java, May 27, 1949, by Neervoort (no. 427; isotype in US). 

Externally this species resembles P. tumescens Hale & Kurokawa. 
It differs in having a black undersurface and a P+ unknown substance 
instead of stictic acid. Parmelia cireumnodata Nyl., a much smaller 
lichen, has short sparse isidia, a pale undersurface with branched 
rhizines, and protocetraric acid. Parmelia abstrusa Vain., the fourth 
isidiate species in series Relicinae, has a thick rigid thallus and pro- 
duces norstictic acid. Parmelia planiuscula is still known only from 
the type locality. 


6. Parmelia ramosissima Kurokawa, sp. nov. PLATE 2 


Thallus adnatus, corticola, obscure viridiflavicans, ca. 3 cm. dia- 
metro, lobis sublinearibus, 0.7—2.0 mm. latis, margine integris, bulbato- 
ciliatis, superne nitidus, albomaculatus, isidiis sorediisque destitutus, 
subtus pallide castaneus, rhizinosus, rhizinis nigricans, dense ramosis. 
Apothecia adnata, ecoronata, retrorso-rhizinata; hymenium 40-50 u 
altum; sporae 23 uy. 

Thallus adnate on bark, turning dark olive buff (R) in the herbar- 
ium, more than 3 cm. in diameter; lobes dichotomously branched, 
sublinear-elongate, 0.7-2.0 mm. wide, 200-240 » thick, the margins 
smooth, with bulbate cilia; upper surface plane and smooth, rather 
shiny, maculate, isidia and soredia lacking; medulla white; under- 
surface pale brown to brown, moderately rhizinate, the rhizines black- 
ening, finely and densely branched. Apothecia adnate, ecoronate, 
1.0-2.5 mm. in diameter, exciple crenate, often basally retrorsely 
rhizinate, disc warm blackish brown (R); hymenium 40-50 p high; 
spores about 23 uz. 

Reactions: Thallus K+ yellowish; medulla K—, C—, KC-+ red- 
dish, P+ orange red, usnic acid and protocetraric acid present. 

Type in Herbarium Bogoriense, collected on Tanimbar Island, 
Moluccas, Apr. 2, 1938, by P. Buwalda (no. 4583; isotype in US). 

This species resembles P. acrobotrys Kurokawa in habit and in the 
pale undersurface. However, P. ramosissima is distinctly maculate 
and contains protocetraric rather than salacinic acid. It has the 
smallest spores of any species in section Relicinae. It is presently 
known only from the type locality. 


146 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


7. Parmelia schizospatha Kurokawa, sp. nov. PLATE 2 

Thallus adnatus, corticola, obscure viridiflavicans, ca. 5 em. di- 
ametro, lobis sublinearibus, 1-2 mm. latis, margine subcrenatis, 
bulbato-ciliatis, lobulatis, lobulis ramosis, superne planus, nitidulus, 
lobulatus, subtus niger, modice rhizinosus, rhizinis nigris, simplicibus. 
Apothecia ignota. 

Thallus adnate on bark, turning deep olive buff (R) in the herbar- 
ium, about 5 cm. in diameter; lobes irregularly branched, sublinear- 
elongate, 1-2 mm. wide, 100-120 » thick, the margins more or less 
crenate, with black bulbate cilia, lobulate, the lobules dorsiventral, 
often dichotomously branched, not ascending; upper surface plane 
and smooth, faintly maculate, somewhat shiny, lobulate, the lobules 
the same as the marginal ones; undersurface black, moderately rhi- 
zinate, the rhizines black, shiny, simple. Apothecia not seen. 

Reactions: Thallus K+ yellowish; medulla K—, C—, KC—, P+ 
orange red, usnic acid and an unknown P+ substance present. 

Type in Herbarium Bogoriense, collected at Gegerbentang, Java, 
June 27, 1949, by Neervoort (no. 1062; isotype in US). 

This new species is identical with P. sublimbata Nyl. in chemistry 
but it is distinctly lobulate. The lobules are quite fragile and fre- 
quently branched. At present the species is known only from the 
type locality. 

8. Parmelia sublanea Kurokawa, sp. nov. 

Thallus adnatus, rigidulus, obscure viridiflavicans, ca. 10 cm. 
diametro, lobis sublinearibus, subimbricatis, 1-2 mm. latis, margine 
integris, bulbato-ciliatis, ciliis apice ramosis, superne planus vel 
convexus, nitidus, emaculatus, isidiis sorediisque destitutus, subtus 
pallide castaneus, dense rhizinosus, rhizinis pallide castaneis, dense 
ramosis. Apothecia substipitata, 1.0-4.5 mm. diametro; hymenium 
40-50 » altum; sporae 4-5X7-9 un. 

Thallus rather rigid, olive buff to deep olive buff (R), corticolous, 
about 10 cm. in diameter; lobes dichotomously branched, sublinear- 
elongate, subimbricate, 1-2 mm. wide, 220-3004 thick, the margins 
entire, often brown rimmed, with bulbate cilia; upper surface plane 
to more or less convex, smooth and shiny, without maculae, isidia and 
soredia lacking; medulla white; undersurface pale brown, densely 
rhizinate, the rhizines pale brown, densely branched, often forming a 
woolly mat. Apothecia substipitate, 1.0-4.5 mm. in diameter, exciple 
crenate, more or less undulate, amphithecium smooth, disc seal brown 
(R); hymenium 40-504 high; spores 4-5 X7-9 u 

Reactions: Thallus K-++ yellowish; medulla K—, C—, KC+ 
reddish, P-++ orange red, usnic acid and protocetraric acid present. 

Type in Herbarium Bogoriense, collected on route between Ake 


HALE & KUROKAWA—SUBGENUS PARMELIA 147 


Biaur and Tolewang, Halmahera Island, Oct. 12, 1951, by P. Groen- 
hardt (no. 8409; isotype in US). 

As previously discussed, P. sublanea is very similar to P. acrobotrys 
except in chemistry. Parmelia ramosissima has similar chemistry 
but the rhizines are darker. The bulbate cilia of P. sublanea are 
unusual in being almost spherical and quite large. The species is 
known only from the type locality. 


9. Parmelia tumescens Hale & Kurokawa, stat. et nom. nov. 


Parmelia limbata Laur. f. isidiosa Mill. Arg. Flora 70:59. 1887. Type 
collection: Richmond River, Australia, Hodgkinson s.n. (G, holotype). 


Thallus adnate on bark, turning deep olive buff in the herbarium 
about 4 cm. in diameter; lobes dichotomously branched, sublinear, 
1.0-1.5 mm. wide, 190-230, thick, 'the margins subcrenate, with 
bulbate cilia; upper surface more or less convex, irregularly cracked 
on older lobes, white-maculate, isidiate, the isidia cylindrical, simple or 
branched, up to 0.5 mm. high; medulla white; undersurface brown, 
moderately rhizinate, the rhizines simple, rather thick, dark brown to 
black. Apothecia not seen. 

Reactions: Thallus K—; medulla K+ yellow, C—, KC—, P+ pale 
orange red, usnic acid and stictic acid present. 

The specific epithet isidiosa is preempted by Parmelia isidiosa 
(Mill. Arg.) Hale, a species in subgenus Xanthoparmelia. Except for 
the presence of isidia, P. tumescens is very similar to P. limbata Laur., 
another Australian species that occurs commonly on rocks rather 
than bark. 

Additional specimens examined: Australia: Eastern Creek, New 
South Wales, Wilson (H); Brisbane, Queensland, Bailey (BM). 


4, Section Cyclocheila (Vain.) Ras. (1943, p. 10) 


Section Hypotrachyna *Cyclocheila Vain. (1890, p. 47). 

Pseudoparmelia Lynge (1913, p. 15). Type species: P. cyphellata (Lynge) 
Sant. 

Section Subflavescentes (Vain.) Gyel. subsection Eciliatae Gyel. (1932, p. 
225). Lectotype: P. caperata (L.) Ach. 

Section Hypotrachyna subsection Cyclocheila (Vain.) Gyel. (1932, p. 224). 

Section Hypotrachyna subsection Cyclocheila (Vain.) Hillm. (1934, p. 187). 
Superfluous combination. 

Section Pseudoparmelia (Lynge) Ris. (1943, p. 10). 


Lectotype species: Parmelia amazonica Ny]. 

Thallus adnate to loosely attached; lobes sublinear and truncate 
to subirregular and subrotund; lower surface black or brown, the 
rhizines simple, often coarse. 

As we previously mentioned (p. 129), Vainio first recognized Cyclo- 
cheila as a group under section Hypotrachyna, characterized by 


148 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


subrotund lobes and including three species, P. amazonica Nyl., P. 
minarum Vain. (=P. dissecta Nyl.), and P. muelleri Vain. The latter 
two species are ciliate and should be classified in our section Imbricaria 
subsection Imbricaria. Vainio (1923) later combined this group and 
Trregulares into a single section Irregulares (Vain.) Vain. We have 
amended section Irregulares to include only species with a reticulate 
upper cortex, marginal cilia, and usually squarrosely branched rhizines. 

The type species of section Cyclocheila, Parmelia amazonica, has 
simple unbranched rhizines and lacks any marginal cilia. There are 
45 additional eciliate species with simple rhizines that can be classified 
here. These include typical sublinear-lobed species (e. g., P. sphaero- 
spora Nyl., P. texana Tuck.) and a number of subirregular, subrotund- 
lobed species, such as P. amazonica Nyl., P. caperata (L.) Ach., P. 
caroliniana Nyl., and P. soredians Nyl. Some of these species have 
a narrow bare or papillate zone below at the margins and could be 
confused with species in subgenus Amphigymnia. Section Cyclocheila 
is best represented in Africa where 18 of the species are endemic; they 
are often saxicolous. Divaricatic acid is especially common along 
with a number of other K—, P— acids also found in section Hypo- 
trachyna (see below, also cf. table 1). 

Lynge first proposed the genus Pseudoparmelia on the supposed 
discovery of ‘“pseudocyphellae” on the lower surface of Parmelia 
cyphellata, a mistake followed by Riisiinen (1943). Santesson (1944) 
has conclusively shown that these structures are merely breaks or 
gashes in the lower cortex where rhizines have been torn away. 
Gyelnik (1933) used this genus name to accommodate the pseudo- 
cyphellate Amphigymnia species related to Parmelia cetrarioides (Duby) 
Nyl., but this application of the name seems to be incorrect. 

In his original publication, Ras&inen (1943) did not actually indicate 
himself as the author of section Cyclocheila but listed Vainio alone, 
apparently on the mistaken assumption that Vainio had recognized 
this taxon at the sectional level previously. Rdasinen clearly cites 
Cyclocheila as a section, however, and we believe he should be cited as 
the author of the new combination. 


Key to Species in Section Cyclocheila 


1. Thallus isidiate or pustulate. 
2. Thallus distinctly pustulate; soredia absent or only sparsely developed. 
3. Thallus corticolous; lobes rather broad, 2-5 mm. wide. 


4. Medulla P— (divaricatic acid). . .... 4. P. eruptens Kurokawa 

4, Medulla P+ red (protocetraric acid). . ... . P. raunkiaeri Vain. 
3. Thallus saxicolous; lobes narrow, to 1 mm. wide. 

5. Medulla P+ red (unknown). ..... 6. P. imperfecta Kurokawa 


5. Medulla P—. 


HALE & KUROKAWA—SUBGENUS PARMELIA 149 


6. Caperatic acid present .... . . 8. P. pustulescens Kurokawa 
6. Divaricatic acid present . ..... =... . P. owariensis Asah. 
2. Thallus with cylindrical isidia. 
7. Lower surface pale brown. 


8. Medulla pale yellow .. ..... . . P.cyphellata (Lynge) Sant. 
8. Medulla white. 
9. Lobes broad, 2-5 mm. wide; subrotund . . 9. P. salacinifera Hale 


9. Lobes narrow, 0.5-2.0 mm. wide, sublinear. 
10. Thallus mineral gray, usnic acid lacking. 2. P. areana Kurokawa 
10. Thallus yellowish green (usnic acid). 
11. Lobes 1.5-2.0 mm. wide; rhizines black. 
P. abstrusoides des Abb. 
11. Lobes 0.5-1.5 mm. wide; rhizines mostly pale. 
12. Medulla P—, KC-+ orange (barbatic acid). 
P. rahengensis Vain. 
12, Medulla P+ red (protocetraric acid) . . P. malaccensis Nyl. 
7. Lower surface jet black. 
13. Thallus yellowish green (usnic acid) . . . . P. ecaperata Mill. Arg. 
13. Thallus mineral gray (usnie acid lacking). 
14. Thallus saxicolous. 
15. Medulla C++ deep red (lecanoric acid) . 1. P. annexa Kurokawa 
15. Medulla C—, P+ pale orange (stictic acid). 
7. P. ischnoides Kurokawa 
14. Thallus corticolous. 
16. Medulla P+ orange red. 
17. Medulla K+ red (salacinicacid). .. . P. cinerascens Lynge 
17. Medulla K— or brownish (protocetrariec acid). 
18. Isidia rather irregular, often becoming granular. 
P. martinicana Nyl. 
18. Isidiacylindrical,normal. . . . . . . P.amazonica Ny]. 
16. Medulla P—. 
19. Upper cortex finely cracked; perlatolic acid present. 
P. caroliniana Nyl. 
19. Upper cortex continuous or irregularly cracked with age. 
20. Lobes subrotund, 2-5 mm. wide; KC-+-red unknown present. 
P. meiosperma (Hue) Dodge 
20. Lobes sublinear, 1.5-2.0 mm. wide; divaricatic acid present. 
P. concrescens Vain. 
1. Thallus lacking pustules or isidia. 
21. Thallus sorediate. 
22. Thallus yellowish green (usnic acid). 
23. Medulla K-+red (salacinic acid); soredia farinose. P. soredians Ny). 
23. Medulla K— (protocetraric acid); soredia coarse, originating from 
pustules... ... 2... 2.04 P. caperata (L.) Ach. 
22. Thallus mineral gray (usnic acid lacking). 
24. Upper cortex reticulately ridged and wrinkled; stictic acid present. 
25. Lobes subrotund, 2-5 mm. wide .. . P. crozalsiana B. de Lesd. 
25. Lobes sublinear, 1.0-2.5 mm. wide . . P. carneopruinata Zablbr. 
24. Upper cortex plane, not ridged. 
26. Soredia strongly capitate, submarginal. 
P. cryptochlorophaea Hale 
26. Soredia laminal, not strongly capitate. 


CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


27. Yellowish pigment present under soralia; KC+ unknown 

present . .. . . . P. aptata Kremplh. 
27. Pigments absent; divar . P. texana Tuck. 
21. Thallus lacking soredia. 


28. Lower surface pale to dark brown. 
29. Medulla pigmented, the pigment K+ purple black. 


30. Pigment present throughout the medulla. 
11. P. violacea Kurokawa 


150 


icatic acid present . . 


30. Pigment present only in lower half of the medulla. 
P. hypomilta Fée 


29. Medulla white or pale yellow, the pigment K—. 
31. Medulla pale yellow. 
32. Thallus saxicolous; lobes 1 mm. wide or less. 
P. chapadensis Lynge 
32. Thallus corticolous; lobes 2-5 mm. wide . P. sphaerospora Nyl. 


31. Medulla white, without any pigments. 
33. Thallus corticolous; medulla P-+ orange red (protocetraric acid). 
. P. intertexta Mont. & v.d. Bosch, 


34. Usnic acid present . . 
34. Usnic acid absent. 
35. Spores 5-78-11 y; Africa . . . P.somaliensis Mill. Arg, 
35. Spores 14-18 26-28 w; South America. 
P. leucopis Kremplh. 


33. Thallus saxicolous; medulla P—. 
36. Medulla C+ deep red (lecanoric acid) . . P. molybdiza Nyl.- 
36. Medulla C— ... .. . 10. P. tortula Kurokawa 
28. Lower surface jet black. 


37. Medulla pigmented, the pigment K+ purple black. 
38. Thallus yellowish green (usnic acid). . P.rutidota Hook. & Tayl. 


38. Thallus mineral gray (usnic acid lacking). 


39. Thallus saxicolous, the lobes subareolate, appressed. 
P. lecanoracea Mill. Arg. 


39. Thallus corticolous, the lobes loosely adnate. 
5. P. heterochroa Hale & Kurokawa 


37. Medulla white; pigments absent. - 
40. Thallus saxicolous. 
41. Lobes appressed to the substratum. 
42. Medulla C+ red (lecanoric acid) . . . . .P.molybdiza Nyl. 
42, Medulla C—. 
43. Medulla K+ yellow (stictic acid). 
P. xanthomelaena Mill. Arg. 
43. Medulla K— (divaricatic acid) . . P. rupicola Lynge 
41. Lobes loosely adnate. 
44, Medulla P+ red (protocetraric acid). . .3. P. caribaea Hale 
44. Medulla P— (divaricatie acid) . . .P. rodriguesiana Hue 


40. Thallus corticolous. 
45. Thallus yellowish green (usnic acid) . P. rutidota Hook. & Tay). 


45. Thallus mineral gray (usnic acid lacking). 


46. Medulla K+ yellow (stictic acid); upper cortex reticulately 
ridged ..... . . P. scrobicularis Kremplh. 
P. subtiliacea Nyl. 


46. Medulla K—; upper cortex plane. . .. . 


HALE & KUROKAWA—SUBGENUS PARMELIA 151 


1. Parmelia annexa Kurokawa, sp. nov. PLATE 4 


Thallus arcte adnatus, saxicola, albido-cinerascens, 3-7 cm. latus, 
lobis sublinearibus, centrum versus subimbricatis, 0.7-3.5 mm. latis, 
margine crenatis, superne nitidus, emaculatus, isidiatus, subtus niger 
sparse rhizinosus, rhizinis nigris, simplicibus. Apothecia adnata, 
1.5-5.0 mm. diametro; hymenium 70-80 yp altum; sporae 5X7-8 uy. 

Thallus tightly adnate on rock, pale olive gray to light mineral gray 
(R), 3-7 cm. in diameter; lobes irregularly branched, sublinear- 
elongate, more or less imbricate towards the center, 0.7-3.5 mm. 
wide, 180-240 » thick, the margins crenate; upper surface plane, 
shiny, not maculate, irregularly cracked with age, isidiate, the isidia 
cylindrical, mostly simple, darkening at the tips, about 0.2 mm. high; 
medulla white; undersurface black, sparsely rhizinate, the rhizines 
black, simple. Apothecia adnate, 1.5-5.0 mm. in diameter, exciple 
undulate, amphithecium isidiate, disc snuff brown (R); hymenium 
70-80 » high; spores 5X7-8 u. 

Reactions: Thallus K+ yellow; medulla K—, C+ deep red, KC+ 
red, P—, atranorine and lecanoric acid present. 

Type in the Botanical Museum, Lund University, collected at 
Mossel River, 3 miles east of Hermanus, Caledon, Cape Province, 
Union of South Africa, Sept. 29, 1953, by Ove Almborn (no. 5683; 
isotype in US). 

This is a typical South African saxicolous lichen clearly related to 
P. molybdiza Nyl., the only other species of the section Cyclocheila 
with lecanoric acid. Parmelia molybdiza differs in lacking isidia and 
in having a pale brown, rarely black, undersurface. Both species 
grow tightly adnate on exposed acidic rocks over a wide area in 
southern Africa. 

Additional specimens examined: Angola: Malanje: Between Ben- 
guela. and Coporolo, Degelius, Feb. 2, 1960 (DEGEL), Duque de 
Braganca, Degelius, Mar. 5, 1960 (DEGEL, US). Union of South 
Africa: Natal: 8 miles south of Lomkile, Nongoma Distr., Héeg, 
August 1929 (TRH) ; Cape Province: Cape Town, Héeg, May 31, 1929 
(TRH). Mossel River, Caledon Distr., Almborn 5701 (LD), Camps 
Bay, Cape Distr., Almborn 4402 (LD). 

2. Parmelia arcana Kurokawa, sp. nov. PuaTE 4 

Thallus arcte adnatus, saxicola, albido-cinerascens, 3-7 cm. latus, 
lobis sublinearibus, 0.3-1.0 mm. latis, margine subcrenatis, superne 
nitidus, emaculatus, isidiatus, medulla pro parte pallide flavescens, 
subtus castaneus, sparse rhizinosus, rhizinis castaneis, simplicibus. 
Apothecia ignota. 

Thallus tightly adnate on rock, pale to dark olive gray (R), 3-7 cm. 
in diameter; lobes dichotomously or irregularly branched, sublinear- 

722-891—64_8 


152 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


elongate, areolate toward the center, 0.3-1.0 mm. wide, 100-150 u 
thick, the margins more or less crenate; upper surface plane to convex 
rather shiny, not maculate, tangentially cracked on older lobes 
isidiate, the isidia cylindrical, mostly simple, about 0.2 mm. high; 
medulla white, partly pale yellow in the upper half; undersurface 
brown to pale brown, sparsely rhizinate, the rhizines brown, simple. 
Apothecia not seen. 

Reactions: Thallus K-++ yellow; medulla K—, C—, KC—, P-, 
pigment if present K—, atranorine, an unidentified fatty acid, and an 
unidentified pale yellow pigment present. 

Type in the herbarium of G. Degelius, collected in desert northeast 
of Baba, Mocdmedes, Angola, Feb. 5, 1960, by G. Degelius (isotype in 
US). 

Parmelia arcana is a tightly adnate saxicolous species with the 
same habit and lobe configuration as P. annera Kurokawa and P. 
molybdiza Nyl. These latter two species contain lecanoric acid. 
The pale undersurface of P. molybdiza is reminiscent of P. arcana, 
and the isidia of P. anneza are similar to those of P. arcana except 
that they are generally more papillate. 

Additional specimen examined: Union of South Africa: Konings- 
kroon, Elliott Distr., Cape Prov., Héeg, Dec. 1, 1929 (TRH). 

3. Parmelia caribaea Hale, sp. nov. PLATE 3 

Thallus adnatus, eborinus vel cinereo-albicans, saxicola, 6-10 em. 
diametro, lobis parum lobatis, irregulariter sublinearibus, subimbri- 
catis, margine integris, superne planus, demum rugosus, sorediis 
isidiisque destitutus, subtus niger, modice rhizinosus, rhizinis crassis, 
nigris, simplicibus, ambitu fusco-castaneus, anguste nudus vel papil- 
latus. Apothecia subpedicellata, urceolata, usque ad 5 mm. diametro; 
hymenium 50-60 u altum; sporae 4-5 10-13 p. 

Thallus adnate, saxicolous, ivory to whitish gray, 6-10 cm. wide, 
rather coriaceous; lobes irregularly sublinear, infrequently branched, 
more or less imbricate, 3-6 mm. wide, 200-250 yp, the margins entire 
to undulate, more or less black-rimmed, soredia and isidia lacking; 
upper surface plane, rugulose with age, cracked on older lobes; medulla 
white; undersurface black, moderately rhizinate, the rhizines thick, 
simple, the marginal zone brown, narrowly naked or papillate. Apo- 
thecia subpedicellate, at first globose, urceolate at maturity, up to 
5 mm. in diameter, amphithecium rugose, the disc imperforate; 
hymenium 50-60 » high; spores 4-5 10-13 yp. 

Reactions: Thallus K+. yellow; medulla K—, KC+ rose, C—, 
P+ orange red, atranorine and protocetraric acid present. 

Type in the Missouri Botanical Garden, collected at Flanc Nord, 
Morne de Villet, St. Barthelemy, West Indies, Oct. 23, 1952, by 
C. Le Gallo (no. 494; isotype in US). 


HALE & KUROKAWA—SUBGENUS PARMELIA 153 


This remarkable species has been collected only on the island of 
St. Barthelemy. It is a saxicolous lichen without any obvious 
affinities. The mode of branching and the undersurface resemble 
P. nairobiensis Stein. & Zahlbr. and related South African species 
that contain divaricatic acid. Parmelia caribaea also has the black- 
rimmed margins typical of these species but the upper surface lacks 
any distinct white-netted reticulation. The apothecia are peculiar 
in that the disc may remain closed until quite late in the development 
of apothecia, so that young apothecia appear like spherical buds. 

Additional specimens examined: St. Barthelemy: Saline, Le Gallo 
491 (MO, US); Morne de Villet, Le Gallo 473 (MO, US); without 
specific locality, Le Gallo 2604 (MSC). 


4. Parmelia eruptens Kurokawa, sp. nov. PLATE 3 
Thallus adnatus, corticola, olivaceo-albicans, 5-8 cm. diametro, 
lobis subirregularibus, 2-8 mm. latis, margine crenatis, superne 
emaculatus, isidiatus, isidiis crassis, inflatis, apice demum pustules- 
centibus, subtus niger, sparse rhizinosus, ambitu castaneus, sub- 
nudus, rhizinis nigricans, simplicibus. Apothecia adnata, 1-3 mm. 
diametro; hymenium 90-100 » altum; sporae 5-7%10-12 u. 

Thallus adnate on bark, marguerite yellow to olive buff (R), 5-8 
cm. in diameter; lobes irregularly branched, subrotund, 2-8 mm. 
wide, 190-240, thick, the margins crenate, dark brown-rimmed near 
the tips; upper surface smooth and shiny, without maculae, moderately 
to densely isidiate, the isidia irregularly inflated, narrower at the base, 
at length breaking open at the tips; medulla white; undersurface 
black, brown and naked in a narrow zone near the tips, sparsely 
rhizinate, the rhizines pale near the margins, blackening at the center, 
simple. Apothecia adnate, 1-3 mm. in diameter, exciple more or less 
crenate, amphithecium isidiate, disc vandyke brown (R); hymenium 
90-100 u high; spores 5-7 10-12 u. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC—or KC + 
fleeting purple violet, P—, atranorine and divaricatic acid present. 

Type in the Botanical Museum, Lund University, collected 5 miles 
east of Mokobulaan, Lydenburg, Transvaal, Union of South Africa, 
Oct. 21, 1953, by Ove Almborn (no. 7498; isotype in US). 

Parmelia eruptens is characterized by rather large subirregular 
lobes and unusually large pustules which break open apically without 
soredial formation. It has much broader and more loosely attached 
lobes than other corticolous species containing divaricatic acid, such as 
P. nairobiensis Stein. & Zahlbr. and P. texana Tuck., though it is 
obviously related to these species. Parmelia eruptens is apparently 
endemic to South Africa. 

Additional specimens examined: Union of South Africa: Transvaal: 
Mount Anderson, Sabie-Lydenburg, Maas Geesteranus 6453 (L, US), 


154 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


6455 (L); Mogambique: Near Rock Pueshang, west of Martins Falls, 
Manica and Sofala distrs., Mitchell 332 (US). 


5. Parmelia heterochroa Hale & Kurokawa, nom. nov. 


Parmelia hypoxantha Stirt. Trans. New Zealand Inst. 32:76. 1899. Type 
collection: Warwick, Queensland, Australia, Gwyther s.n. (BM, holotype). 
Non P. hypoxantha Mill. Arg. Flora 64:85. 1881 (= Chondropsis semi- 
viridis Nyl.). 

Parmelia tiliacea (Hoffm.) Ach. var. stenophylla Mill. Arg. Flora 66:46. 
1883. Type collection: Toowoomba, Queensland, Australia, Hartmann 
s.n. (G, holotype). 

P. tiliacea var. rugulata Mill. Arg. Nuov. Giorn. Bot. Ital. 23:388. 1891. 
Type collection: Brisbane, Australia, Batley s.n. (G, lectotype). 

P. tiliacea var. converula Mill. Arg. Bull. Herb. Boiss. 4:90. 1896. Type 
collection: Brisbane, Australia, Shirley 1665 (G, holotype) [collector on 
label given as Bailey]. 

Type collection: Warwick, Queensland, Australia, Gwyther s.n. 
(BM). 

Thallus adnate, mineral gray, turning olive buff in the herbarium, 
4-12 cm. in diameter; lobes more or less dichotomously branched, 
sublinear-elongate, 14 mm. wide, 160-210, thick, the margins more 
or less crenate, often black-rimmed near the tips; upper surface plane 
to convex, smooth, shiny, irregularly rugose and cracked on older 
lobes, isidia and soredia lacking; medulla white above, capucine 
buff to zinc orange in the lower half; undersurface black, moderately 
to densely rhizinate, the rhizines black, simple, irregularly to squar- 
rosely branched, 0.5-1.7 mm. long. Apothecia adnate, 1-53 mm. in 
diameter, exciple smooth to crenate or undulate, disc imperforate; 
hymenium 70-80 » high; spores 7-8 10-12 u, episporium 1 yu thick. 

Reactions: Thallus K+- yellow; medulla K—, C—, KC-+ rose, 
P-+ orange red, pigment K-+ purple, atranorine, protocetraric acid, 
and an unidentified anthraquinone pigment present. 

This remarkable species is known only from Queensland, Australia, 
where it appears to be common. Superficially, it resembles P. 
liwida Tayl. or even P. brasiliana Nyl. but the lower medulla is con- 
spicuously pigmented. The rhizines are mostly simple, though 
occasionally squarrosely branched. It is the only species outside of 
section Parmelia and section Irregulares that has squarrose rhizines, 
but it is clearly unrelated to these two sections. On a gross morpho- 
logical basis we would prefer to put P. heterochroa in section Hypo- 
trachyna but the peculiar rhizines indicate a closer affinity to section 
Cyclocheila. 

Additional specimens examined: Australia: Toowoomba, Queens- 
land, Hartmann 45 (G) (syntype of P. tiliacea var. rugulata) ; Queens- 
land, Batley 334 (BM). 


HALE & KUROKAWA-—SUBGENUS PARMELIA 155 


6. Parmelia imperfecta Kurokawa, sp. nov. 


Thallus arcte adnatus, saxicola, albido-cinerascens, 2-4 cm. dia- 
metro, lobis sublinearibus, 0.5-2.0 mm. latis, margine crenatis, 
superne nitidus, isidiato-pustulatus, subtus niger, sparse rhizinosus, 
rhizinis nigris, simplicibus. Apothecia ignota. 

Thallus closely or loosely adnate on rock, pale gull gray to light 
mineral gray (R), 2-4 cm. in diameter; lobes irregularly branched, 
sublinear-elongate, 0.5-2.0 mm. wide, 120-170» thick, the margins 
crenate; upper surface plane to more or less concave, shiny, irregularly 
cracked on older lobes, isidiate-pustulate, the pustules short, erupting ; 
medulla white; undersurface black, sparsely rhizinate, the rhizines 
simple, about 0.2 mm. long. Apothecia not seen. 

Reactions: Thallus K+ yellow; medulla K— or K+ yellowish, 
C—, KC—, P+ orange red, atranorine and an unknown P+ sub- 
stance (forming small balls in G. A. o—T.) present. 

Type in the Botanical Museum, Lund University, collected at the 
Cathedral Peak Area, Bergville, Natal, Union of South Africa, 
Nov. 2, 1953, by Ove Almborn (no. 8815; isotype in US). 

Parmelia imperfecta is another saxicolous lichen endemic to South 
Africa. It is apparently related to P. annexa Kurokawa, P. arcana 
Kurokawa, and P. molybdiza Nyl. The pustules are rather sparse, 
thick and short and burst open apically without formation of soredia. 
Almost identical pustules are found in the saxicolous Japanese species 
P. owariensis Asah., which differs principally in containing divaricatic 
acid. 

Additional specimens examined: Union of South Africa: Transvaal: 
Punch Bowl Inn, north of Louis Trichardt, Almborn 6491 (LD) ; 
Louis Trichardt, near Punch Bowl, Zoutpansberg, Almborn 6108 
(LD, US). 


7. Parmelia ischnoides Kurokawa, sp. nov. PLATE 4 


Thallus arcte adnatus, saxicola, cinereo-albicans, 4-10 cm. latus, 
lobis sublinearibus, 0.5-2.0 mm. latis, superne nitidus, emaculatus, 
isidiatus, subtus niger, sparse rhizinosus, rhizinis nigris, simplicibus. 
Apothecia adnata, 1-2 mm. diametro; hymenium 70-80 u altum; 
sporae 5-6 X7-8 un. 

Thallus closely adnate on rock, pearl gray to mineral gray (R), 
4-10 cm. in diameter; lobes irregularly branched, sublinear-elongate, 
more or less imbricate with age, 0.5-2.0 mm. wide, 200-280 » thick, 
the margins entire to subcrenate; upper surface plane to convex, 
shiny, not maculate, isidiate, the isidia cylindrical, short and simple, 
darkening apically; medulla white; undersurface black, sparsely 
rhizinate, the rhizines black, simple. Apothecia adnate, 1-2 mm. 


156 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


in diameter, exciple more or less crenate, amphithecium isidiate, disc 
light seal brown (R); hymenium 70-80 » high; spores 5-6 7-8 uy. 

Reactions: Thallus K+- yellow; medulla K+ yellow, C—, KC—, 
P+ pale orange, atranorine and stictic acid present. 

Type in the Botanical Museum, Lund University, collected on 
rocks near Window Stream, Kirstenbosch, Wynberg, Cape Province, 
Union of South Africa, Aug. 3, 1953, by Ove Almborn (no. 1698; iso- 
type in US). 

Parmela ischnoides adds still another member to that unique group 
of saxicolous species so common in the Cape region (see discussions 
under P. annexa and P. imperfecta). It is characterized by adnate, 
linear and more or less separate lobes as illustrated in plate 4 to broad 
and more crowded lobes up to 2 mm. wide. Isidia are often quite 
dense but always rather small or even at times papillate. The 
chemical component, stictic acid, is quite rare in the section Cyclocheila. 

Additional specimens examined: Union of South Africa: Cape 
Province: Kirstenbosch, Wynberg Distr., Almborn 124, 1443 (LD); 
Window Gorge, Wynberg Distr., Almborn 788 (LD); near Kloof N eck, 
Camps Bay Road, Cape Distr., Almborn 864 (LD). 


§. Parmelia pustulescens Kurokawa, sp. nov. Puatr 4 


Thallus arcte adnatus, saxicola, albido-cinerascens, 3-5 cm. latus, 
lobis sublinearibus, 0.3-1.2 mm. latis, margine crenatis, superne 
nitidus, emaculatus, isidiatus, isidiis irregulariter inflatis, demum 
apice pustulescentibus, subtus niger, sparse rhizinosus, rhizinis nigris, 
simplicibus. Apothecia adnata, 0.7-2.0 mm. diametro; hymenium 
45-55 pw altum; sporae 4-5 7-9 u. 

Thallus tightly adnate on rock, pearl gray to light mineral gray (R), 
3-5 cm, in diameter; lobes dichotomously to irregularly branched, 
sublinear-elongate, 0.3-1.2 mm, wide, 100-130 p thick, the margins 
crenate, narrowly black-rimmed near the tips; upper surface plane 
and shiny, without maculae, irregularly cracked on older lobes, isid- 
iate, the isidia cylindrical, irregularly inflated, at length bursting 
open apically; medulla white; undersurface black, sparsely rhizinate, 
the rhizines simple, black, 0.1-0.2 mm. long. Apothecia adnate, 
0.7-2.0 mm. in diameter, exciple isidiate-pustulate, disc clove brown 
(R); hymenium 45-55 uw high; spores 4-5 X7-9 up. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC—, P-, 
atranorine and caperatic acid present. 

Type in the herbarium of G. Degelius, collected near Fazenda 
Canjangue, Vila Flor, Humbo, Angola, Feb. 18, 1960, by G. Degelius 
(isotype in US). 

Parmelia pustulescens is an adnate saxicolous lichen virtually in- 
distinguishable from P. imperfecta Kurokawa and P. owariensis Asah. 


HALE & KUROKAWA—SUBGENUS PARMELIA 157 


in the development of pustules. (See discussion under P. imperfecta.) 
Parmelia pustulescens differs chiefly in containing caperatic acid. It 
is known only from the type specimen. 

9. Parmelia salacinifera Hale, sp. nov. PLATE 9 

Thallus adnatus, corticola, olivaceo-albicans, 6-12 cm. diametro, 
lobis subirregularibus, subrotundatis, 3-6 mm. latis, 170-190 4 crassis, 
superne planus, nitidus, modice isidiatus, subtus pallide castaneus, 
modice rhizinosus, rhizinis concoloribus, simplicibus, ambitu anguste 
nudus vel papillatus. Apothecia adnata, 2-4 mm. diametro; hyme- 
nium 30-38 u altum; sporae non evolutae. 

Thallus adnate, corticolous, olivaceous to mineral gray, 6-12 cm, 
in diameter; lobes subirregular, 3-6 mm. wide, 170-190, thick, 
apically subrotund, the margins entire to subcrenate; upper surface 
plane, rugose and fissured with age, shiny, moderately isidiate, the 
isidia cylindrical, usually simple, 0.04-0.08 X0.16-0.30 mm.; under- 
surface pale brown, moderately rhizinate, the rhizines pale brown, 
simple, the marginal zone bare or papillate. Apothecia adnate, 2-4 
mm. in diameter, amphithecium isidiate, disc imperforate; hymenium 
30-38 u high; spores not developed. 

Reactions: Thallus K+ yellow; medulla K+ yellow turning red, 
C—, KC—, P-+ pale orange red, atranorine and salacinic acid present. 

Type in the U.S. National Herbarium, collected at Sanford, Semi- 
nole County, Florida, March 1925, by S. Rapp (isotype in FLAS). 

Parmelia salacinifera is a typical member of section Cyclocheila 
with irregular subrotund lobes. The thallus is usually quite closely 
adnate on bark. At first glance it seems indistinguishable from 
P. amazonica Ny]l., which occurs from Florida to Brazil; P. amazonica 
has a jet black undersurface and protocetraric acid. Parmelia 
salacinifera undoubtedly occurs generally in the Caribbean region. 
A typical habitat is burned palm trunks in pastures. 

Additional specimens examined: Georgia: 5 mi. SW. of Waycross, 
Ware Co., Hale 16843 (US). Florida: Tomoka State Forest, Volusia 
Co., Hale 17063 (US); near Pensacola, Escambia Co., Hale 7994 (US); 
Dog Lake Recreation Area, Hale, 16714 (US) ;6 mi. SW. of Tallahassee, 
Leon Co., Hale 16984 (US); Orange Park, Clay Co., Hale 17735 (US) ; 
vicinity of Fort Myers, Lee Co., Standley 13084 (US); Sanford, 
Seminole Co., Rapp 472, May 1919, February 1924 (FLAS). Mexico: 
Km. 956 on Hwy. 190, west of Ocozocoautla, Chiapas, [Zale 20607 
(US). Brazil: Mato Grosso: Bocca da Serra, Malme s.n. (UPS). 

10. Parmelia tortula Kurokawa, sp. nov. PLATE 4 

Thallus adpressus, saxicola, albido-cinerascens, 5-10 cm. latus, 
lobis subirregularibus vel sublinearibus, apice subrotundatus, 1.5-4.0 
mm. latis, margine crenatis, superne nitidus, emaculatus, isidiis 


158 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


sorediisque destitutus, subtus pallide castaneus, sparse rhizinosus, 
rhizinis castaneis, simplicibus. Apothecia adnata vel substipitata, 
1-5 mm. diametro; hymenium 40-50 uz altum; sporae 5-6 X7-9 p. 

Thallus closely adnate on rock, yellowish glaucous to light olive 
gray (R), 5-10 cm. in diameter; lobes irregularly branched, sub- 
irregular to sublinear-elongate, more or less subrotund apically, often 
twisted and contorted, 1.54.0 mm. wide, 180-220» thick, the margins 
entire to crenate, black-rimmed at the tips; upper surface shiny, 
often faintly pruinose, without maculae, rugulose and distinctly 
cracked on older lobes, isidia and soredia lacking; medulla white; 
undersurface pale brown, sparsely rhizinate, the rhizines simple, 
pale brown, about 0.3 mm. long. Apothecia adnate to substipitate, 
1-5 mm. in diameter, exciple and amphithecium smooth, disc bone 
brown (R); hymenium 40-50 » high; spores 5-6 X7-9 p. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC—, or 
KC-- faint rose, P—, atranorine and unknown colorless substances 
present. 

Type in the Botanical Museum, Lund University, collected on 
rocks between O’okiep and Springbok, Namaqualand, Cape Province, 
Union of South Africa, Sep. 15, 1953, by Ove Almborn (no. 4805; 
isotype in US). 

Parmelia tortula is a saxicolous species common in the Cape region 
and obviously related to P. molybdiza Nyl. Generally it is larger 
and the colonies more extensive than in P. molybdiza. The lobes are 
often more or less twisted and, at least near the tips, partially free of 
the substratum; P. molybdiza is less robust and contains lecanoric 
acid. The chemistry of P. tortula is not entirely clear at this time. 
The KC+ test is variable, often weak or absent. Microchemical 
tests have not revealed the cause of this reaction. 

Additional specimens examined: Union of South Africa: Orange 
Free State: Thabanchis, Héeg, Aug. 12, 1929 (TRH); Cape Province: 
4 miles south-southwest of Oudtshoorn, Oudtshoorn Distr., Almborn 
4199 (LD); Saldanha, Malmesbury Distr., Almborn 4988, 4996 (LD); 
Sea Point, Cape Town, Wynberg Distr., Héeg, May 31, 1929 (TRH); 
Dordricht, Héeg, Nov. 21, 1929 (TRH). 


11. Parmelia violacea Kurokawa, sp. nov. 


Thallus adpressus vel adnatus, saxicola, olivaceo vel viridi-cineras- 
cens, 3-6 cm. diametro, lobis sublinearibus, 1-3 mm. latis, margine 
crenatis, superne nitidus, emaculatus, isidiis sorediisque destitutus, 
medulla purpureus, subtus pallide castaneus, modice rhizinosus, 
rhizinis castaneis, simplicibus. Apothecia adnata, 1.0-1.5 mm. 
diametro; sporis non vVisis. 


HALE & KUROKAWA—SUBGENUS PARMELIA 159 


Thallus closely adnate on rock, smoke gray to dark olive gray 
(R), 3-6 cm. in diameter; lobes irregularly branched, sublinear- 
elongate, 1-3 mm. wide, 160-200 » thick, the margins crenate; upper 
surface shiny, without maculae, distinctly rugulose on older lobes, 
isidia and soredia lacking; medulla vinaceous purple to indian purple 
(R); undersurface pale brown, moderately rhizinate, the rhizines 
pale brown, simple, 0.5-1.0 mm. long. Apothecia adnate, 1.0-1.5 
mm. in diameter, exciple and amphithecium smooth, disc cameo 
brown to vandyke brown (R); mature spores not seen. 

Reactions: Thallus K+ yellow; medulla K+ deep purple, P—, 
atranorine and an unidentified hydroxanthraquinone present. 

Type in the Botanical Museum, Lund University, collected at 
Blinkwater Ravine, slopes of Table Mountain, Cape Province, Union 
of South Africa, Aug. 5, 1953, by Ove Almborn (no. 1771; isotype 
in US). 

Parmelia violacea has a brilliant deep rusty-red medulla, frequently 
exposed as the upper cortex breaks away. In this respect it is very 
similar to P. endomiltoides Nyl., a Xanthoparmelia species with usnic 
acid and two unidentified pigments, one of them (of lower R,) identical 
with that in P. violacea. Both of these species are found on Table 
Mountain in Cape Province. 


5. Section Hypotrachyna Vain. (1890, p. 38) 


Section Hypotrachyna* Sublinearts Vain. (1890, p. 50). 

Section Sublineares (Vain.) Vain. (19238, p. 34). 

Section Hypotrachyna subsection Sublineares (Vain.) Gyel. (1932, p. 223). 

Section Hypotrachyna subsection Sublineares (Vain.) Hillm. (1934, p. 187). 
Superfluous combination. 


Type species: Parmelia brasiliana Nyl. 

Thallus adnate to loosely attached and subdivaricate; lobes sub- 
linear to linear-elongate; lower surface jet black, the rhizines more or 
less richly dichotomously branched. 

The type species of this section should be selected from the Brazilian 
species cited in Vainio’s original study. We believe that he considered 
his group Sublinearis to represent the typical concept of section Hypo- 
trachyna and P. brasiliana Nyl. to be a typical species in this group. 
Two other groups described at the same time, Cyclocheila (= section 
Cyclocheila (Vain.) Ris.) and Irregularis (= section Irregulares (Vain.) 
Vain.), were described as having subirregular rotund lobes with a 
narrow bare zone below. In his study of Philippine lichens, Vainio 
(1923) put the subirregular species into a new section (/rregulares) 
and the sublinear species into section Sublineares (Vain.) Vain. He 
included Parmelia americana (Mey. & Flot.) Mont. and P. sorocheila 


160 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Vain. in section Sublineares, but these are everniiform species that 
should be transferred to the genus Psewdevernia Zopf. 

As we have delimited section Hypotrachyna, it does not include 
P. sazatilis (L.) Ach., the type of section Parmelia, which has pseudo- 
cyphellae and squarrosely branched rhizines. For this reason, sec- 
tion Parmelia, which includes P. sazatilis, and section Hypotrachyna 
cannot be considered as synonymous. 

Section Hypotrachyna is the largest yet surprisingly the most homo- 
geneous group of species in subgenus Parmelia. All species have a 
black lower surface with more or less distinctly dichotomously branched 
rhizines, sometimes so dense as to form a marginal mat projecting 
beyond the margins. Cilia are absent, although projecting branched 
rhizines might be mistaken for cilia. The greatest concentration of 
species is in tropical America, where 48 of the 83 species are endemic. 
Fourteen species are endemic to Asia and only six to Africa. 

Morphological and chemical diversity reach a peak in this section. 
There is an abundance of sorediate species (14). There are a number 
of acids not found in any of the other sections of subgenus Parmelia 
except section Cyclocheila (cf. table 1). These include alectoronic 
acid, barbatic acid, divaricatic acid, evernic acid, lichexanthone, olive- 
toric acid, and perlatolic acid. All of these substances are nega- 
tive with KOH, paraphenylenediamine, calcium hypochlorite (except 
for olivetoric acid), and most are KC-+ red and more or less fluores- 
cent in ultraviolet light. In addition, if depsides, they have a basic 
2-methoxy substitution on the acid ring (cf. Asahina and Shibata, 
1954, p. 54), whereas the other depsides usually have a 2-hydroxy 
substitution. The full significance of this chemical variation has not 
yet been assessed, but the overall correlation between the chemical 
constituents and our sectional classification appears to be sound. 


Key to Species in Section Hypotrachyna Vain. 


1. Thallusisidiate 2... 1. 1 we ee ee ee ee ee I. 
1. Thallus lacking isidia. 
2. Thallus sorediate or pustulate ... 1... 2.0.22, . . ID. (p. 161). 
2. Thallus lacking isidia, soredia, or pustules. ....... III. (p. 163). 


I. THALLUS ISIDIATE 


1. Medulla yellow, the pigment K—. 
2. Medulla C—; isidia dense ......... .. P.isidiocera Nyl. 
2. Medulla C+ red (unknown); isidia variable, sparse, simple to coralloid. 
23. P. prolongata Kurokawa 
1. Medulla white, or if pigmented, the pigment K+ purple. 
3. Upper cortex distinctly maculate. 
4, Medulla K—, C—, KC—, P— (fatty acids) ... P. costaricensis Nyl. 


HALE & KUROKAWA—SUBGENUS PARMELIA 161 


4. Medulla P+ yellow to red or KC+ red or orange. 
5. Medulla K+ yellow, P+ yellow (unknown). 
9. P. dentella Hale & Kurokawa 
5. Medulla K—, KC-+ red or orange. 
6. Medulla C_, KC-+ orange (barbatic acid). 


7. Isidia cylindrical . ......... . P. imbricatula Zahlbr. 

7. Isidia lobulate, dorsiventral ..... .. « P. digitata Lynge 
6. Medulla C—, KC+ rose (evernic acid). 

8. Isidia cylindrical . . ..... =... .. . P. bogotensis Vain. 

8. Isidialobulate, dorsiventral . ... .. . P. culmigena Zahlbr. 


3. Upper cortex not distinctly maculate. 
9. Medulla P+ yellow to orange red. 
10. Medulla K— (protocetraric acid). 
11. Medulla ochraceous yellow in the lower half . P. consimilis Vain. 
11. Medulla entirely white ....... .. . P. koyaensis Asah. 
10. Medulla K+ yellow or red. 
12. Thallus yellowish green (usnic acid); norstictic acid present. 
P. microblasta Vain. 
12. Thallus grayish green (usnic acid lacking). 
13. Rhizines finely branched; isidia mostly simple; norstictic acid 


present. ...... . . . 24. P. rhabdiformis Kurokawa 
13. Rhizines sparsely to moderately branched; isidia often branched; 
stictic acid present ....... 5. P. crenata Kurokawa 


9. Medulla P—. 
14. Medulla C+ red or yellow or KC-+ red or orange. 
15. Medulla C+ deep red, KC+ red (unknown). 
23. P. prolongata Kurokawa 
15. Medulla C— or C+ yellow, KC+ orange (barbatie acid). 
16. Isidia cylindrical .. . . . . « P. imbricatula Zahlbr. 
16. Isidia lobulate, dorsiventral ..... . . P. digitata Lynge 
14. Medulla C—, KC—, or KC-+ red. 
17. Medulla KC+ red or rosy. 
18. Isidia inflated or pustular; medulla usually ochraceous yellow 
under the isidia. ...... . . . . P. dactylifera Vain. 
18. Isidia simple or lobulate; pigments absent. 
19. Isidia lobulate, dorsiventral; alectoronic acid present. 
10. P. ensifolia Kurokawa 
19. Isidia cylindrical; KC-+ unknown present. 
P. nodakensis Asah. 
17. Medulla C—, KC—; fatty acids present. 
20. Upper cortex weakly maculate; rhizines densely branched; 
America. ... . . . . . P. costaricensis Nyl. 
20. Upper cortex without maculae; ‘thizines moderately branched; Asia. 
18. P. infirma Kurokawa 


II. THALLUS SOREDIATE OR PUSTULATE 


1. Pustules distinct, without soredia or only very sparsely sorediate. 


2. Thallus yellowish green (usnic acid). . ... . . . . P. meyeri Zahlbr. 
2. Thallus mineral gray (usnic acid lacking). 
3. Medulla pale yellow orange, pigment K— . . . P. endochlora Leight. 


3. Medulla white or in part pigmented, pigment K+ purple. 
4. Pigments present, K+ purple. 


162 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


5. Thallus 2-4 em. broad, the lobes up to 1 mm. wide . P. malmei Lynge 
5. Thallus 3-10 cm. broad, the lobes 1.0-2.5 mm. wide. 
P. formosana Zahlbr. 
4, Pigments absent. 
6. Medulla C+ rose (gyrophoric acid) . . .. . P. revoluta Floerke 
6. Medulla C—, KC+ orange (barbatic acid). 
7. Pustules distinct, becoming sorediate; Asia . . . P. exsecta Tayl. 
7. Pustules irregular, esorediate; cortex fragile; North America. 
29. P. virginica Hale 
1. Pustules indistinct or absent; soredia usually abundant, farinose. 
8. Upper cortex distinctly maculate. 
9. Medulla pale yellow orange throughout . . . . P. endochlora Leight. 
9. Medulla white. 
10. Medulla KC+ orange (barbatic acid) . . P. laevigata (Sm.) Ach. 
10. Medulla KC-+ red or rose. 
11. Evernic acid present .......... . P.rockii Zahlbr. 
11. Alectoronic acid present . ..... . 13. P. exsplendens Hale 
8. Upper cortex without maculae or at most in part faintly maculate. 
12. Thallus yellowish green (usnic acid). 
13. Medulla K+ yellow turning red (salacinic acid). 
P. sinuosa (Sm.) Ach. 
13. Medulla K—, P+ orange red (protocetraric acid). 
14. Soralia mainly terminal; medulla ochraceous (rhodophyscin) under 
the soralia ........ +e. es . P. velloziae Vain. 
14. Soralia mainly subterminal; pigments lacking. 
15. P. flavovirens Kurokawa 
12. Thallus mineral gray (usnic acid lacking). 
15. Medulla P+ yellow to orange red. 
16. Medulla K+ yellow turning to red (salacinic acid). 
17. Rhizines moderately branched; zeorine present. 
P. majoris Vain. 
17. Rhizines finely branched; zeorine absent. 
2. P. brevirhiza Kurokawa 
16. Medulla K—. 
18. Lichexanthone present; P-+ unknown present. 
P. subaffinis Zahlbr. 
18. Lichexanthone absent; protocetraric acid present. 
19. Orange-red pigment (rhodophyscin) under the soralia. 
6. P. croceopustulata Kurokawa 
19. Pigments absent under the soralia. 
20. Lobes 1.5-2.0 mm. wide; soralia mostly subterminal. 
P. pseudosinuosa Asah. 
20. Lobes 0.5-1.0 mm. wide; soralia mostly laminal. 
1. P. anaptychioides Kurokawa 
15. Medulla P—. 
21. Orange-yellow pigment (rhodophyscin) under the soralia. 
19. P. leiophylla Kurokawa 
21. Pigments absent. 
22. Medulla C+ rose, red, or orange red. 
23. Medulla C+ red or orange red (olivetoric acid). 
21. P. lividescens Kurokawa 
23. Medulla C+ rose (gyrophoric acid). 


HALE & KUROKAWA—SUBGENUS PARMELIA 163 


24. Lobes linear-elongate, subdivaricate; Mexico. 
28. P. thysanota Kurokawa 
24. Lobes loosely adnate, sublinear; Africa .. P. revoluta Flk. 
22. Medulla C—, KC-+ red or orange red. 
25. Medulla KC+ orange (barbatic acid) ... P. rockii Zahlbr. 
25. Medulla KC-+ rose or red. 
26. KC+ unknown present; Africa. 
17. P. immaculata Kurokawa 
26. Alectoronic acid present; tropical America. 
27. Lobes linear-elongate, subdivaricate. 
8. P. densirhizinata Kurokawa 


27. Lobes sublinear, loosely adnate to subimbricate. 
13. P. exsplendens Hale 


Ill. THALLUS LACKING ISIDIA, SOREDIA, AND PUSTULES 


1. Thallus yellowish green (usnic acid). 
. P. flavida Zahlbr. 


2. Thallus saxicolous. ....... re 
2. Thallus corticolous. 
3. Medulla K—, P+ orangered. ........+--s P. flavida Zahlbr. 
3. Medulla K+ red, P+ orange red. 
4. Salacinic acid present ........ . 4. P. citrella Kurokawa 


4. Norstictic acid present. 
5. Thallus closely adnate; lobes sublinear; rhizines densely branched. 
P. reducens Nyl. 
5. Thallus loosely adnate to subdivaricate; lobes linear-elongate; 
rhizines moderately branched. 
6. Lobes 1-2 mm. wide, sparsely rhizinate, subimbricate. 
P. enderythraea Zahlbr. 
6. Lobes 2-4 mm. wide, densely rhizinate, subdivaricate. 
P. caraccensis Tayl. 
1. Thallus mineral gray (usnic acid lacking). 
7. Medulla pale yellow or in part ochraceous. 


8. Pigment K—. 
9. Apothecia stalked; rhizines richly branched .. P. peruviana Nyl. 


9. Apothecia adnate; rhizines sparsely to moderately branched. 
25. P. rigidula Kurokawa 
8. Pigment K+ purple to purple black. 
10. Thallus orange fluorescent under UV light (lichexanthone present). 
11. Upper cortex maculate; thallus corticolous. . P. silvatica Lynge 
11. Upper cortex not maculate; thallus saxicolous. 
12. Medulla C+ red (olivetoric acid). . . P.osteoleuca Nyl, 
12. Medulla C—, P+ red (unknown). 
ll. P. erythrodes (Zahlbr.) Hale & Kurokawa 
10. Thallus not fluorescent (lichexanthone absent). 
13. Medulla P+ red (protocetraric acid)... . P.bahiana Nyl. 
13. Medulla P— (fatty acids)... .. . . P.lythgoeana Dodge 
7. Medulla white, no pigments present. 
14. Medulla P+ yellow to orange red. 
15. Medulla K+ yellow turning red. 
16. Norstictic acid present . ... . . 3. P. canescens Kurokawa 


16. Salacinic acid present. 


164 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


17. Thallus corticolous; rhizines densely branched. 
P. sublaevigata (Nyl.) Nyl. 
17. Thallus saxicolous; rhizines moderately branched 
P. subsaxatilis B. de Lesd. 
15. Medulla K— or K+ faint brownish. 
18. Thallus saxicolous. 
19. Cortex orange fluorescent in UV light (lichexanthone present). 
P. brasiliana Nyl. 
19. Cortex not fluorescent (lichexanthone absent). 
P. osseoalbida Lynge 
18. Thallus corticolous. 
20, Medulla KC+ orange (barbatic acid present). 
26. P. scytodes Kurokawa 
20. Medulla KC+ rose (protocetraric acid present), 
21. Spores 15-25 yp long. 
22. Spores 20-25 u long; fatty acid present . P. keitauensis Asabh, 
22, Spores 15-20 yu long; fatty acids lacking. 
P. manilensis Vain. 
21. Spores 8-12 » long. 
23. Apothecia large, the dise split radially; Africa. 
14. P. fissicarpa Kurokawa 
23. Apothecia small, the disc entire; tropical America. 
24, Rhizines moderately branched lobes subrotund often, whitish 
pruinose... . . . . P.insinuans Nyl. 
24 Rhizines fine, densley branched; lobes sublinear, epruinose. 


P. bahiana Ny]l. 
14. Medulla P—. 


25. Medulla C—, KC+ orange (barbatie acid). 
26. Thallus subdivaricate, the lobes linear-elongate; rhizines mostly 


moniliform ......... . 22. P. monilifera Kurokawa 
26. Thallus adnate, the lobes sublinear; rhizines normal, not moniliform. 
27. Upper cortex distinctly maculate. . . . . . . P. boliviana Nyl. 


27. Upper cortex not distinctly maculate, 
28. Amphithecium strongly rugose; Java. 
P. bostrychodes Zahlbr, 
28. Amphithecium smooth; tropical America. 
29. Lobes sublinear to linear-elongate; rhizines densely branched. 
P. physcioides Ny]. 
29. Lobes shorter, subimbricate; rhizines sparsely to moderately 
branched. ... . .. . . P. zahlbruckneri Lynge 
25. Medulla C+ rose or red or C_, “KC+ red, 
30. Medulla C+ rose (ayrophoric acid). 
31. Lobes linear-elongate, subdivaricate. 
12. P. exporrecta Kurokawa 
31. Lobes sublinear, adnate. 
32. Lobes plane to more or less canaliculate, pale brown in a rather 
wide zone at the margins below; spores 14-21 yu long. 
P. pluriformis Nyl. 
32. Lobes plane, pale brown only in a very narrow zone below; 
spores to 12 4 long. 
33. Thallus large, 4-8 em. broad, the lobes 2-4 mm. wide. 
27. P. seytophylla Kurokawa 


HALE & KUROKAWA—SUBGENUS PARMELIA 165 


33. Thallus smaller, 2-5 cm. broad, the lobes 1-2 mm. wide. 
P. revolutella Ny]. 
30. Medulla C+ red or C— (gyrophoric acid lacking). 
34. Medulla C+ red to orange red (olivetoric acid). 
P. intercalanda Vain. 
34. Medulla C—. 
35. Medulla KC-+ red (alectoronic acid). 
36. Thallus adnate, the lobes sublinear. . . . 7. P. degelii Hale 
36. Thallus loosely attached, subdivaricate. 
37. Lobes narrow, 1-2 mm. wide. 
20. P. lineariloba Kurokawa 
37. Lobes wider, 2-4 mm. wide. 
38. Rhizines densely branched, about 1 mm. long; tropical 


America. . . 1... 2 eee 16. P. gigas Kurokawa 
38. Rhizines moderately branched, 2 mm. or more long; 
Hawaii. .......+-.-. P. cervicornis Tuck. 

35. Medulla KC+ rose (alectoronic acid absent). 
39. Upper cortex distinctly maculate. . .. . P. pulvinata Fée 


39. Upper cortex not distinctly maculate. 
40. Upper cortex flaking away, fragile; evernic acid present. 
P. taylorensis Mitch. 
40. Upper cortex continuous, not flaking; KC-+ unknown sub- 
stances present. 
41. Spores uniseriate, 6-8 » long; North America. 
P. livida Tay]l. 
41. Spores biseriate, 14-18 » long; South America. 
42. Thallus corticolous; apothecial disc flesh-colored. 
P. damaziana Zahlbr. 
42. Thallus saxicolous; disc dark brown. 
P. gracilescens Vain. 


1. Parmelia anaptychioides Kurokawa, sp. nov. PuatTE 5 

Thallus adnatus, cinereo-albicans, 2-5 cm. diametro, lobis sub- 
linearibus, separatis, 0.5-2.0 mm. latis, superne plus minusve convexus, 
sorediatus, subtus niger, rhizinosus, rhizinis nigris, dichotome ramosis. 
Apothecia ignota. 

Thallus adnate on bark, pale olive gray to gray (R), 2-5 cm. in 
diameter; lobes dichotomously branched, sublinear-elongate, separate 
at the circumference, 0.5-2.0 mm. wide, 160-220 u thick, the margins 
smooth, narrowly black-rimmed; upper surface more or less convex, 
shiny on younger lobes, dull and irregularly cracked on older lobes, 
sorediate, the soredia mainly laminal, 1-2 mm. in diameter, often 
fusing; medulla white; undersurface black, dark brown in a narrow 
zone at the tips, the rhizines black, branched. Apothecia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ faint red, 
P+ orange red, atranorine and protocetraric acid present. 

Type in Michigan State University, collected on low ridge connect- 
ing La Rucilla and Pico Trujillo, Maciso Central, Cordillera Central, 


166 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


elev. 8000 ft., Dominican Republic, August 8, 1958, by C. Wetmore 
(no. 3617; isotype in US). 

This species is related to P. pseudosinuosa Asah. but the soralia are 
mainly laminal. The lobes are sublinear to linear-elongate and 
narrower than in P. pseudosinuosa. It is also close to P. subaffinis 
Zahlbr., especially in size and position of soralia, but it is differentiated 
by the absence of lichexanthone. Parmelia anaptychioides is known 
only from the West Indies, where it occurs on pine trees at higher 
elevations. 

Additional specimens examined: Jamaica: Blue Mountain Peak, 
7000 ft., Imshaug 15533 (MSC). Haiti: South of Forét des Pins, 
5500 ft., Dept. ’ Ouest, Imshaug 22779 (MSC). 

2. Parmelia brevirhiza Kurokawa, sp. nov. PLATE 3 

Thallus adnatus, ca. 5 cm. diametro, olivaceo-cinerascens, lobis 
subirregularibus, 1-4 mm, latis, subrotundatis, margine crenatis, 
superne opacus vel nitidulus, submaculatus, partim leviter albo- 
pruinosus, sorediatus, sorediis capitatis, subtus niger, dense brevi- 
rhizinosus, rhizinis nigris, dichotome ramosis. Apothecia rara, 
adnata, usque ad 5 mm. diametro; hymenium 50-55 yp altum; sporae 
male evolutae, 3-4 6-8 u. 

Thallus adnate on bark, cream buff (R), about 5 cm. in diameter; 
lobes subirregular to sublinear, more or less subrotund and sub- 
imbricate, 1-4 mm. wide, 140-170 u thick, the margins subcrenate; 
upper surface plane to rugulose, shiny and more or less white-maculate 
to dull and lightly white-pruinose, sorediate, the soredia mostly 
subterminal, strongly capitate, diffuse, medulla white; undersurface 
black, densely short rhizinate, dark brown and papillate in a very 
narrow zone at the tips, the rhizines black, finely branched, about 0.1 
mm. long, projecting as a narrow mat beyond the margins. Apothecia 
very rare, adnate, to 5 mm. in diameter, amphithecium sorediate, 
dise plane; hymenium 50-55 uw high; spores poorly developed, 3-4 
6-8 uy. 

Reactions: Thallus K-+ yellow; medulla K+ yellow turning red, 
C—,KC— P- pale orange red, atranorine and salacinic acid present. 

Type in the Naturhistoriska Riksmuseet, Stockholm, collected in 
Nothofagus rain forest, Isla Riesco, Mina Elena, Terr. Magallanes, 
Chile, Apr. 29, 1940, by R. Santesson (no. 2066; isotype in US). 

Parmelia brevirhiza has a distinct whitish cast, caused by faint 
pruinosity, and large subterminal and projecting soralia 2-4 mm. in 
diameter. The sorediate lobes are strongly revolute or contorted. 
This species resembles P. majoris Vain., which is known from Asia 
and Madagascar, in the presence of soredia and salacinic acid. Par- 
melia brevirhiza has more densely branched rhizines, which form a 
mat on the undersurface. Parmelia majoris contains zeorine in 


HALE & KUROKAWA—SUBGENUS PARMELIA 167 


addition to salacinic acid. Parmelia brevirhiza is very common in 
southern Argentina and Chile and there is one quite unexpected record 
from Java. 

Additional specimens examined: Argentina: Lago Roca, Santesson 
1085 (S); Rfo Piper, Ushuaia, Tierra del Fuego, Santesson 417 (S); 
Lago Verde, near Futalaufquen, Prov. Chubut, Lamb 5864 (CAN, 
US). Chile: Punta Guapacho, Peninsula Lacui, Isla Chiloé, Prov. 
Chiloé, Santesson 4135 (S); Cerros Divisaderos, Coyhaique, Terr. 
Aysén, Santesson 4456 (S); Mina Elena, Isla Riesco, Santesson 7849 
(S); Estancia Maria, Seno Skyring, Terr. Magallanes, Santesson 1974, 
1997 (S); Puerto Yartou, Canal Whiteside, Santesson 5791, 6759 (S); 
Rio Condor, south of Puerto Yartou, Canal Whiteside, Santesson 
5905 (S); Puerto Navarino, Isla Navarino, Santesson 7404 (S, US); 
Porvenir, Tierra del Fuego, Santesson 5372 (5). Java: Near Madang- 
Aer Batumbuk, Groenhart 9268 (BO). 

3. Parmelia canescens Kurokawa, sp. nov. 

Thallus adnatus, corticola, olivaceo-albicans, 3-7 cm. latus, lobis 
irregularibus, subrotundatis, 1-5 mm. latis, margine subcrenatis, su- 
perne opacus, parte albo-pruinosus, isidiis sorediisque destitutus, 
subtus niger, rhizinosus, rhizinis nigris, sparse vel modice ramosis, 
Apothecia adnata, 1.5-4.5 mm. diametro; hymenium 40-50 yu altum; 
sporae 5-68-10 yp. 

Thallus adnate on twigs, pale olive buff (R), 3-7 cm. in diameter; 
lobes irregularly branched, subrotund apically, 1-5 mm. wide, the 
margins more or less crenate, rarely lobulate; upper surface plane, 
dull, sometimes faintly white-pruinose, not maculate, irregularly 
rugose or cracked on older lobes, isidia and soredia lacking; medulla 
white; undersurface uniformly black, sparsely to moderately branched ; 
rhizines black, moderately branched, about 0.5 mm. long. Apothecia 
adnate, amphithecium rugulose, pruinose, disc vandyke brown (R); 
hymenium 40-50 yu high; spores 5-6 X8-10 yu. 

Reactions: Thallus K+ yellow; medulla K-+ yellow turning red C—, 
KC—, P+ pale orange, atranorine and norstictic acid present. 

Type in the Naturhistoriska Riksmusect, Stockholm, collected at 
Hacienda de Cayquenes, Colchagua, Chile, Aug. 22, 1896, by P. Dusén 
(no. 92; isotype in US). 

Parmelia canescens externally resembles P. sublaevigata (Nyl.) Nyl. 
in having a similar adnate thallus and irregularly branched lobes. 
Parmelia canescens contains norstictic acid, whereas P. sublaevigata 
contains salacinic acid. Moreover, the rhizines of P. canescens are 
sparsely to moderately branched while those of P. sublaevigata are 
more densely and finely branched. It is hoped that further collec- 
tions of these rare species will further clarify their relationships. 

722-891—64_4 


168 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


4. Parmelia citrella Kurokawa, sp. nov. 


Thallus adnatus, corticola, viridiflavicans, 3-8 cm. latus, lobis sub- 
linearibus, subimbricatis, 1.0-2.5 mm. latis, superne nitidulus, isidiis 
sorediisque destitutus, subtus niger, rhizinosus, rhizinis nigris, furcatis. 
Apothecia adnata, 2-10 mm. diametro; hymenium 25-35 yw altum; 
sporae 7-9 12-14 up. 

Thallus adnate on bark, sea-foam yellow (R), 3-8 cm. in diameter; 
lobes dichotomously branched, sublinear to linear-elongate, more or 
less imbricate, 1.0-2.5 mm. wide, 90-140 thick, the margins smooth, 
often narrowly black-rimmed; uppersurface plane to convex, rather 
shiny, isidia and soredia lacking, pycnidia forming blackish papillae 
on older lobes; medulla white; undersurface black, rhizinate, the 
rhizines black, rather long, moderately branched, often projecting 
beyond the margins. Apothecia adnate, 2-10 mm. in diameter, 
exciple crenate to undulate, disc Hay’s brown to light seal brown 
(R), radially split, amphithecium smooth to rugose; hymenium 25- 
35 high; spores 7-9 12-14 pu. 

Reactions: Thallus K+ yellowish; medulla K+ yellow turning 
red, C—, KC—, P+ pale orange red, usnic acid and salacinic acid 
present. 

Type in the Herbarium Bogoriense, collected at G. Gedeh, Java, 
elev. 2900 m., March 7, 1938, by H. van Woerden (isotype in US). 

Parmelia citrella is very close to P. reducens Nyl., but the thallus 
is more loosely attached and the rhizines are consistently longer. 
Parmelia reducens also differs chemically in producing norstictic 
acid, and on the basis of a few measurements the spores seem to be 
slightly smaller (9-11 long). Parmelia citrella appears primarily 
to be an Indonesian species, while P. reducens is known only from 
South America. 

Additional specimens examined: Panama: Craters edge, Volc4n 
Chiriqui, Prov. Chiriquf, Scholander, Dec. 5-12, 1948 (MO, US). 
Colombia: Sierra de Santa Maria, elev. S000 ft., Humbry-Troy 
313 (K). Java: G. Gedeh, van Woerden 2023 (BO); G. Pangerango, 
Reinvaan 150 (BO); no locality, Horsfield (BM). 

5. Parmelia crenata Kurokawa, sp. nov. 

Thallus laxe adnatus, muscicola, albido-cinerascens, 3-6 cm. dia- 
metro, lobis subirregularibus, subimbricatis, apice subrotundatis 
vel subtruncatis, 1.5-5.0 mm. latis, margine crenatis, superne 
laevigatus, emaculatus, isidiatus, subtus niger, ambitu castaneus, 
sparse rhizinosus, rhizinis nigris, modice furcatis. Apothecia ignota. 

Thallus loosely attached, growing over mosses on rocks, yellowish 
glaucous to light mineral gray (R), 3-6 cm. in diameter; lobes ir- 
regularly branched, more or less imbricate, rounded at the tips, 


HALE & KUROKAWA—SUBGENUS PARMELIA 169 


1.5-5.0 mm. wide, 90-1104 thick, the margins crenate; upper surface 
smooth, without maculae, moderately isidiate, the isidia short, less 
than 0.5 mm. high, often branched; medulla white; undersurface 
black, pale brown in a narrow zone at the tips, sparsely rhizinate, 
the rhizines black, sparsely to moderately branched. Apothecia not 
seen. 

Reactions: Thallus K++ yellow; medulla K+ yellow, C—, KC—, 
P+ pale orange, atranorine and stictic acid present. 

Type in the National Science Museum, Tokyo, collected on Mount 
Kuishi, Tosa-gun, Tosa-ken, Shikoku, Japan, Dec. 14, 1960, by 58. 
Kurokawa (no. 60154; isotype in US). 

Parmelia crenata might at first be mistaken for P. koyaensis Asah., 
but the lobes are consistently smaller and more or less sublinear. 
It is the only species in section Hypotrachyna that contains stictic 
acid. Parmelia koyaensis contains protocetraric acid. Thus far 
P. crenata is known only from the type locality in Japan. 


6. Parmelia croceopustulata Kurokawa, sp. nov. PLATE 9 


Thallus adnatus, albido-cinerascens, ca. 5 cm. diametro, lobis 
sublinearibus, 1.5-3.5 mm. latis, margine integris, superne nitidus, 
emaculatus, aetate rugulosus, pustulato-sorediatus, soraliis partim 
subcoloratis, subtus niger, rhizinosus, rhizinis nigris, ramosis. 
Apothecia adnata, 1-6 mm. diametro; hymenium 50-60» altum; 
sporae non visae. 

Thallus adnate on bark, yellowish glaucous (R), 4-9 em. in diameter; 
lobes dichotomously branched, sublinear, 1.5-3.5 mm. wide, 150- 
220 » thick, the margins entire to more or less dissected; upper surface 
shiny, rugulose with age, without maculae, sorediate, the soralia 
capitate and subterminal, 1-5 mm. in diameter, originating from 
pustules; medulla below the soralia yellow ochre (R), otherwise 
white; undersurface black, rhizinate, the rhizines black, shiny, 
branched. Apothecia rare, adnate, 1-6 mm. in diameter, exciple 
sorediate, disc chestnut brown (R), radially split; hymenium 50-60 u 
high; mature spores not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC— or KC+ 
reddish, P-++ orange red or in part P—, soralia P+ orange red, pig- 
ment under the soralia K-+ purple, atranorine, protocetraric acid, 
and rhodophyscin present. 

Type in Michigan State University, collected on fir around summit 
of Grandfather Mountain, Avery Co., North Carolina, elev. 5964 ft., 
June 23, 1958, by H. Imshaug (no. 22275; isotype in US). 

Parmelia croceopustulata is related to species in the P. livida group, 
especially P. formosana Zahlbr. and P. lecophylla Kurokawa, both fo 
which also produce rhodophyscin. Parmelia formosana differs in 
having nonsorediate pustules and lichexanthone. Parmelia leiophylla 


170 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


has similar sorediate pustules but produces olivetoric acid. This 
species seems to be restricted to higher elevations in Hispaniola and 
Jamaica, with several interesting disjunct localities in the Appalachian 
Mountains. 

Additional specimens examined: United States: Virginia: White 
Top Mountain, Washington Co., elev. 5200 ft., Hale 18662 (US); 
North Carolina: Roan Mountain, Avery Co., elev. 6200 ft., Hale 18071 
(US); Mt. Mitchell, Yancey Co., elev. 6684 ft., ZImshaug 22383, 
22390 (MSC, US); Newfound Gap, Swain Co., Imshaug 22435 (MSC, 
US); Mt. Pisgah, Haywood Co., Green in 1959 (US). Dominican 
Republic: Trail to Alto de la Bandera, Cordillera Central, La Vega, 
Imshaug 23470 (MSC, US). Haiti: Pic La Selle, Morne La Selle, 
elev. 8844 ft., Wetmore 3120 pr. p. (MSC); above Le Refuge, Montagne 
Noire, Kenscoff, Imshaug 22538 (MSC). Jamaica: Mossmans Peak, 
Imshaug 14726 (MSC); High Peak, Blue Mountains, elev. 6800 ft., 
Imshaug 15254 (MSC). 


7. Parmelia degelii Hale, sp. nov. Puate 5 

Thallus arcte adnatus, corticola, cinereo-albicans, 4—8 cm. diametro, 
lobis sublinearibus, 1-2 mm. latis, contiguis, superne rugulosus, 
isidiis sorediisque destitutus, subtus modice rhizinosus, rhizinis sparse 
vel modice dichotome ramosis. Apothecia adnata, 3-5 mm. diametro; 
hymenium 50-60 uv altum; sporae 4-5 X8-11 un. 

Thallus closely adnate on bark, 4-8 cm. in diameter, mineral gray; 
lobes sublinear, 1-2 mm. wide, 140-170 y thick, contiguous, the 
margins entire; upper surface dull to more or less shiny, rugulose, 
transversely rugose with age, isidia and soredia lacking; medulla 
white; undersurface black, moderately rhizinate, the rhizines sparsely 
to moderately branched, dichotomous. Apothecia adnate, 3-5 mm. 
in diameter, the exciple crenate, disc plane; hymenium 50-60 yu high; 
spores 4-5 8-11 up. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC-+ red, P—, 
atranorine, alectoronic acid, and @-collatolic acid present. 

Type in the herbarium of G. Degelius, collected between Luso and 
Cachipoque, Moxico, Angola, about 1300 m. elevation, Feb. 16, 1960, 
by G. Degelius (isotype in US). 

Parmelia degelii is very close to P. livida Tayl. in general habit and 
color, but P. livida, a temperate American lichen with several localities 
in South Africa, has smaller spores (4-5 6-8 y), somewhat denser 
and more richly branched rhizines, and a different chemistry (KC-+ 
red unknown). Parmelia degelii is unusual in being the second known 
species of section Hypotrachyna with alectoronic acid in Africa. The 
other species is P. ersplendens Hale, which also occurs in the Caribbean 
area. The five remaining species in section Hypotrachyna with alec- 
toronic acid occur exclusively in tropical America (P. densirhizinata 


HALE & KUROKAWA—SUBGENUS PARMELIA 171 


Kurokawa, P. ensifolia Kurokawa, P. gigas Kurokawa, and P. 
lineariloba Kurokawa) or in Hawaii (P. cervicornis Tuck.). 

Additional specimens examined: Angola: Same locality and data 
as the holotype (DEGEL); Rio Kuiriri, near Kassuango, Bie, Goss- 
weiler 3256d (BM). 

8. Parmelia densirhizinata Kurokawa, sp. nov. PLATE 8 

Thallus laxe adnatus, corticola, albido-cinerascens, 6-13 cm. latus, 
lobis lineari-elongatis, subdivaricatis, 2-6 mm. latis, margine integris, 
superne planus, emaculatus, pustulato-sorediatus, soraliis subtermi- 
nalibus, subtus niger, rhizinosus, rhizinis densis, dichotome ramosis. 
Apothecia ignota. 

Thallus loosely adnate on bark, pale glaucous green (R), 6-13 
mm. in diameter; lobes dichotomously branched, linear-elongate, 
subdivaricate, 2-6 mm. wide, 110-150 thick, the margins smooth, 
narrowly black-rimmed; upper surface plane, continuous, without 
maculae, sorediate, the soralia mainly subterminal, 1-3 mm. in diam- 
eter, originating from pustules; medulla white; undersurface black, 
densely rhizinate, the rhizines black, shiny, dichotomously branched, 
1-2 mm. long, projecting in a mat beyond the margins. Apothecia 
not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ red, P-, 
atranorine and alectoronic acid present. 

Type in the U.S. National Herbarium, collected on the east side of 
Volean Chiriqui, Prov. Chiriqui, Panama, elev. 1500-3000 ft., Dec. 
5-12, 1948, by P. F. Scholander. 

Parmelia densirhizinata is a typical subdivaricate species related 
to P. gigas Kurokawa, a tropical American species, and P. cervicornis 
Tuck., a Hawaiian species. They all produce alectoronic acid, but 
P. densirhizinata is easily distinguished by the subterminal sorediate 
pustules. It is widespread in tropical America, occurring In moun- 
tains between 2000 and 3750 m. elevation. 

Additional specimens examined: Guatemala: Pacaj4, region of 
Desconsuelo, Totonicapan, Standley 84522 (MO). Panama: Craters 
edge, Volc&n Chiriqui, Prov. Chiriqui, Scholander, Dec. 11, 1948 (US). 
Haiti: Ridge east of Pic La Selle, Morne La Selle, elev. 8300 ft., 
Imshaug 23026 (MSC). Dominican Republic: Trail to Alto de la 
Bandera Cordillera Central, La Vega, elev. 7200 ft., Imshaug 23486 
(MSC). Colombia: Péramo de Guasca, Dept. Cundinamarca, Killip 
34118 (US); Chorreén San Paulina, Nevada del Cocuy, Dept. Boyaca, 
elev. 3750 m., Cuatrecasas 1348a (US). Ecuador: Mount Tunguragua, 
Spruce 175 (KK). Peru: Pampalea, Dept. Ayacucho, elev. 3200 m., 
Killip & Smith 22237 (US). Bolivia: Unduavi Valley, Julio 413 (US). 
Chile: Isla de Chiloe, Joseph 2245 (US); Cerro Tralcan, Lago Rinihue, 


172 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Prov. Valdivia, Santesson 3451 (S); Canal Whiteside, Puerto Yartou, 
Tierra del Fuego, Santesson 6799 (S). 
9. Parmelia dentella Hale & Kurokawa, sp. nov. Puats 5 

Thallus laxe adnatus, saxicola, viridi-cinerascens, 3-8 cm. diametro, 
lobis sublinearibus vel subirregularibus, subimbricatis, 2-6 mm. 
latis, margine dentato-crenatis, superne planus, albomaculatus, dense 
isidiatus, subtus nigricans, sparse vel modice rhizinosus, rhizinis 
nigris, ramosis. Apothecia ignota. 

Thallus loosely adnate on shaded rocks, yellowish glaucous to 
glaucous green (R), 3-8 cm. in diameter; lobes irregularly branched, 
sublinear to subirregular, more or less imbricate, 2-6 mm. wide, 
150-230 » thick, the margins in part dentate-crenate; upper surface 
plane, distinctly white-maculate, densely isidiate, the isidia often 
branched, cylindrical, less than 0.7 mm. high; medulla white; under- 
surface black, pale brown in a rather broad zone at the tips, sparsely 
to moderately rhizinate, the rhizines black, shiny, moderately 
branched, about 1 mm. long. Apothecia not seen. 

Reactions: Thallus K+ yellow; medulla K+ yellowish, C—, 
KC—, P+ pale orange red, atranorine and an unknown P-++ substance 
present. 

Type in the U.S. National Herbarium, collected on granite at 
Cheaha State Park, Clay County, Alabama, Mar. 16, 1962, by H. 
McCullough (no. 570; isotypes in TNS and the herbarium of Howard 
College, Birmingham, Alabama). 

Parmelia dentella bears a striking resemblance to the endemic 
Japanese species P. crenata Kurokawa. It differs from P. crenata in 
having the upper cortex distinctly white-maculate and in producing 
a P+ unknown substance instead of stictic acid. It also resembles 
P. costaricensis Nyl., especially in the maculae, isidia, and size 
of the thallus, but the rhizines of P. costaricensis are much more 
densely branched and its main component is caperatic acid (P—). 
Parmelia dentella is still known only from the type locality in Alabama, 
a rather dry oak forest rich in many kinds of lichens. It will probably 
be found again in the foothills of the Appalachian Mountains. 

10. Parmelia ensifolia Kurokawa, sp. nov. Pate 7 

Thallus adnatus, albido-cinerascens, 5-11 cm. latus, lobis subline- 
aribus, 1-4 mm. latis, margine lobulatis, superne nitidus, emaculatus, 
isidiato-lobulatus, subtus niger, rhizinosus, rhizinis nigris, dichotome 
ramosis. Apothecia adnata, 3-4 mm. diametro; hymenium 50-65 y 
altum; sporae 8-10 14-18 u. 

Thallus loosely adnate, corticolous, pearl eray (R), 5-11 cm. in 
diameter ; lobes dichotomously branched, sublinear-elongate, 1-4 mm. 
wide, 160-200 u thick, the margins with numerous lobules, the lobules 


HALE & KUROKAWA—SUBGENUS PARMELIA 173 


ascending, dorsiventral, short-rhizinate below, often branched, up 
to 1.5 mm. long; upper surface shiny, without maculae, short-isidiate- 
lobulate to lobulate on the margins; medulla white; undersurface 
black, densely rhizinate, the rhizines black, shiny, dichotomously 
branched. Apothecia adnate, 3-4 mm. in diameter, disc tawny 
olive (R), amphithecium rugose; hymenium 50-65 yu high; spores 
8-10 14-18 pz. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC-+ rose or 
red, P—, atranorine and alectoronic acid present. 

Type in Michigan State University, collected in pine forest below 
summit of Alto de la Bandera, Cordillera Central, La Vega, Dominican 
Republic, elev. 8300 ft., Aug. 5, 1958, by H. Imshaug (no. 23430; 
isotype in US). 

Parmelia ensifolia has sublinear lobes and laminal and marginal 
lobules similar to those of P. culmigena Zahlbr. and P. digitata Lynge. 
The latter two species, however, are distinctly maculate and differ 
chemically, P. culmigena having evernic acid and P. digitata having 
barbatic acid. Parmelia ensifolia is distinguished by the presence 
of alectoronic acid and lack of maculae. It seems to be rather common 
in tropical America. 

Additional specimens examined: Mexico: No data on locality, 
1890 (S). Panama: Vole4n Chiriqui, Prov. Chiriqui, Scholander, 
Dec. 5-12, 1948 (MO, US). Jamaica: Summit of Blue Mountain 
Peak, Mazon 9865 (US). Haiti: Pic La Selle, Morne La Selle, Wetmore 
3086 (MSC, US). Dominican Republic: Below summit of Alto de 
la Bandera, Cordillera Central, La Vega, elev. 8300 ft., Wetmore 3511 
(MSC, US). Venezuela: Pan de Azticar, Mérida, Jahn 241 (US); 
Laguna de Canoa, Sierra de Santo Domingo, Mérida, Dennis 1924, 
1950 (K). 


11. Parmelia erythrodes (Zahlbr.) Hale & Kurokawa, comb. nov. 


Parmelia brasiliana Nyl. var. erythrodes Zahlbr. Denkschr. Akad. Wiss. 
Wien Math. Naturw. 83: 169. 1927. Type collection: Mt. Itatiaya, 
Minas Gerais, Brazil, Schiffner, September 1901 (WU, holotype). 

Thallus adnate on rock, ivory yellow, deep mouse gray towards 
the center, about 5 cm. in diameter; lobes dichotomously branched, 
sublinear-elongate, 1-2 mm. wide, 140-190, thick, the margins 
entire to subcrenate; upper surface plane, shiny, emaculate, tan- 
gentially cracked on older lobes, isidia and soredia lacking; medulla 
pale pinkish buff in the lower half, white above; undersurface uni- 
formly black, moderately rhizinate, the rhizines black, moderately 
branched, 0.5-1.0 mm. long. Apothecia adnate, 2-6 mm. in diameter, 
exciple undulate, disc imperforate, blackish brown; hymenium 50- 
60 » high; spores 5-69-10 yw, episporium 1 yw thick. 


174 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Reactions: Thallus K-++ yellow; medulla K—, C—, KC+ rose, 
P—, pigment K-+ reddish purple, atranorine, a KC+ unknown 
(apparently the same as the one in P. livida Tayl.), rhodophyscin, 
and lichexanthone. 

Parmelia erythrodes is very close to P. brasiliana Ny]l., differing 
chiefly in chemistry. Parmelia brasiliana lacks rhodophyscin and 
the KC-+ substance but contains lichexanthone and in addition 
protocetraric acid. Both species are typically saxicolous and occur 
rather rarely in South America. 

Additional specimen examined: Peru: Without locality, Lobb s.n. 
(BM). 


12. Parmelia exporrecta Kurokawa, sp. nov. PLATE 6 


Thallus laxe adnatus, albido-cinerascens, 7-10 cm. latus, lobis 
lineari-elongatis, subdivaricatis, 2-6 mm. latis, superne planus, 
nitidus, emaculatus, sorediis isidiisque destitutus, subtus niger, 
rhizinosus, rhizinis nigris, dense ramosis. Apothecia ignota. 

Thallus loosely adnate on bark, pale glaucous green (R), 7-10 cm. 
in diameter; lobes dichotomously branched, linear-elongate, subdi- 
varicate, 2-6 mm. wide, 150-220 u thick, the margins smooth, rarely 
lobulate; upper surface plane and smooth, shiny, often pruinose near 
the tips, without maculae, isidia or soredia; medulla white; under- 
surface black, rhizinate, the rhizines black, shiny, densely branched, 
about 1 mm. long. Apothecia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C+ rose, KC+ 
red, P—, atranorine and gyrophoric acid present. 

Type in the United States National Herbarium, collected in mature 
pine-oak forest, on trail leading to Cerro San Felipe, Oaxaca, Mexico, 
elev. about 3000 m., Apr. 1, 1960, by M. E. Hale (no. 20722). 

Parmelia exporrecta seems to be the nonsorediate ally of P. thysanota 
Kurokawa (see below). It has a large subdivaricate thallus and linear- 
elongate lobes, much as in P. gigas Kurokawa, which differs in con- 
taining alectoronic acid. 

Additional specimen examined: Mexico: Vera Cruz: Orizaba, Lieb- 
mann 116 (UPS). 


13. Parmelia exsplendens Hale, sp. nov. 

Thallus adnatus, viridi- vel cinereo-albicans, 4-8 cm. diametro, 
lobis sublinearibus, 1.5-2.5 mm. latis, superne nitidus, valde albomac- 
ulatus, sorediatus, soralis subterminalibus, subtus niger, dense 
rhizinosus, rhizinis dichotome ramosis. Apothecia ignota. 

Thallus adnate, greenish to whitish gray, 4-8 cm. across, corti- 
colous; lobes sublinear, dichotomously branched, 1.5-2.5 mm. wide, 
160-200 » thick, the margins entire to subcrenate; upper surface 
plane, shiny, strongly white-maculate, sorediate, soralia capitate, 


HALE & KUROKAWA—-SUBGENUS PARMELIA 175 


subterminal; medulla white; undersurface black, densely rhizinate, 
the rhizines black, dichotomously branched. Apothecia unknown. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ red, 
P—, atranorine and alectoronic acid present. 

Type in Michigan State University, collected on ridge northwest of 
Murdock’s Gap, Blue Mountains, Jamaica, elev. 3600 ft., Mar. 19, 
1958, by H. Imshaug (no. 15306; isotype in US). 

This species resembles P. metarevoluta Asah. from Asia in having 
a medium-sized thallus and subterminal capitate soralia. However, 
it is distinctly white-maculate and the rhizines are densely branched. 
Parmelia exsplendens is fairly common in the Caribbean with one 
disjunct record from South Africa. 

Additional specimens examined: Mexico: Virgin upland jungle, 
El Suspiro, Chiapas, Hale 20229 (US). Guatemala: Near Tactic, 
Alta Verapaz, Standley s.n. (MO, no. 185561). Jamaica: Gap south- 
east of Catherine’s Peak, Imshaug 13894 (MSC); gap on northeast 
spur of High Peak, Imshaug 15287 (MSC); Woodcutters Gap, 
Imshaug 13423 (MSC, US); Summit, Blue Mountains, Imshaug 
13855 (MSC); west slope of John Crow Mountains, above Corn Puss 
Gap, Imshaug 14577 (MSC). Union of South Africa: 5 mi. east of 
Mokobulaan, Lydenberg, Transvaal, Almborn 7496 (LD). 

14. Parmelia fissicarpa Kurokawa, sp. nov. PLATE 7 

Thallus adnatus vel laxe adnatus, albido-cinerascens, ca. 14 cm. 
diametro, lobis sublinearibus vel subirregularibus, 2-4 mm. latis, 
margine subcrenatis superne nitidulus, emaculatus, isidiis sorediisque 
destitutus, subtus niger, dense rhizinosus, rhizinis nigris, ramosis. 
Apothecia substipitata, ca. 10 mm. diametro; hymenium 60-70 pz 
altum; sporae 6-7 X8-10u. 

Thallus adnate to loosely attached on rock, pale olive buff (R), 
about 14 cm. in diameter; lobes irregularly branched, sublinear to 
subirregular, 2-4 mm. wide, 210-250 » thick, the margins subcrenate; 
upper surface rather shiny, without maculae, the older lobes rugulose, 
isidia and soredia lacking; medulla white; undersurface black, 
densely rhizinate, the rhizines black, shiny, moderately branched, 
often more than 1.5 mm. long. Apothecia substipitate, 10 mm. or 
more in diameter, exciple undulate, amphitheclum rugose, disc 
carob brown (R), radially split; hymenium 60-70 4 high; spores 
6-7 X8-10 un. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ faint 
red, P+ orange red, atranorine and protocetraric acid present. 

Type in the Botanical Museum, Lund University, collected 2 miles 
south of forester’s office, Cathedral Peak Area, Bergville, Natal, Union 
of South Africa, Nov. 7, 1953, by Ove Almborn (no. 9374; isotype in 
US). 


176 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


This peculiar species has an exceptionally large thallus and 
apothecia. It resembles P. insinuans Nyl., a tropical American 
species, and P. keitauensis Asah. and P. manilensis Vain., both Asian 
species, in having rather irregularly branched lobes and protocetraric 
acid. Although P. fissicarpa and P. insinuans have similar spore size, 
the thallus of P. fissicarpa is much larger and the rhizines are longer. 
In addition, P. insinuans often produces the pigment rhodophyscin 
along with protocetraric acid. Parmelia fissicarpa is clearly dis- 
tinguished from P. keitauensis by the absence of fatty acids and by 
smaller spores. Parmelia manilensis differs in having adnate and 
quite small apothecia (1-3 mm. in diameter) and large spores. 


15. Parmelia flavovirens Kurokawa, sp. nov. PLATE 8 


Thallus adnatus, corticola, viridiflavicans, 3-6 cm. diametro, lobis 
sublinearibus, 1.0-2.5 mm. latis, margine subcrenatis, superne nitidus, 
aetate rugulosus, centrum versus lobulatus, sorediatus, soraliis sub- 
terminalibus, subtus niger, dense rhizinosus, rhizinis nigris, ramosis. 
Apothecia adnata, 2-4 mm. diametro; hymenium 60-75 » altum; 
sporae 7-10 X11-15 gz. 

Thallus adnate on bark, sea-foam green (R), 3-6 cm. in diameter; 
lobes more or less irregularly branched, sublinear, 1.0-2.5 mm. wide, 
130-150 » thick, the margins more or less crenate, often narrowly 
black-rimmed ; upper surface shiny near the tips, more or less rugulose 
and irregularly cracked on older lobes, sometimes lobulate toward the 
center, sorediate, soralia mainly subterminal; medulla white; under- 
surface black, densely rhizinate, the rhizines black, branched. Apo- 
thecia adnate, 2-4 mm. in diameter, exciple crenate, amphithecilum 
rugose, disc light seal brown (R); hymenium 60-75 » high; spores 
7-10 X11-15 p. 

Reactions: Thallus K+ yellowish; medulla K—, C—, KC-+ faint 
red, P+ deep orange red, usnic acid, and protocetraric acid present. 

Type in the Naturhistoriska Riksmuseet, Stockholm, collected 
near Hotel Rio Rubens, 50 km. southeast of Natales, Magallanes, 
Chile, Jan. 14, 1941, by R. Santesson (no. 5681; isotype in US). 

Parmelia flavovirens is very close in external appearance to P. 
sinuosa (Sm.) Ach. and P. velloziae Vain. The last named species 
has similar chemistry but contains in addition the orange-red pigment 
rhodophyscin under the soralia. Parmelia sinuosa produces salacinic 
acid. Parmelia flavovirens is apparently endemic to southernmost 
Chile where it occurs intermingled with P. sinuosa. 

Additional specimens examined: Chile: Puerto Navarino, Isla 
Navarino, Tierra del Fuego, Santesson 1261 (S); Ays¢én: Coyhafgue, 
Santesson 8406 (S). 


HALE & KUROKAWA—-SUBGENUS PARMELIA 177 


16. Parmelia gigas Kurokawa, sp. nov. PuaTE 8 


Thallus laxe adnatus, corticola, albido-cinerascens, 15-40 cm. 
diametro, lobis lineari-elongatis, subdivaricatis, 3-10 mm. latis, 
margine integris, superne planus, emaculatus, isidiis sorediisque desti- 
tutus, subtus niger, breve rhizinosus, rhizinis nigris, dense ramosis. 
Apothecia usque ad 15 cm. diametro, substipitata; hymenium 50-65 u 
altum; sporae 8-10 16-18 y. 

Thallus loosely attached on bark, mosses, or humus, turning from 
ivory yellow to cream buff (R) in the herbarium, 15-40 ecm. in di- 
ameter; lobes dichotomously branched, linear-elongate, subdivaricate 
and separate, 3-10 mm. wide, 220-330 » thick, the margins smooth, 
rarely short lobulate; upper surface plane and smooth, shiny, without 
maculae, isidia and soredia lacking; medulla white; undersurface 
black, short rhizinate, partly naked, the rhizines black, shiny, densely 
branched, about 1 mm. long, forming a mat projecting narrowly 
beyond the margins. Apothecia rare, to 15 mm. in diameter, sub- 
stipitate, amphithecium strongly rugose, faintly maculate; hymenium 
50-65 » high; spores 8-10 X 16-18 u, the episporium 1.5 y» thick. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ red, 
P—, atranorine and alectoronic acid present. 

Type in the United States National Herbarium, collected at Laguna 
Verjou, above Bogot4, Colombia, 1922, by Ariste Joseph (no. B70; 
isotypes in DUKE, REN, 8, TNS). 

This species has a large divaricate thallus, long linear-elongate lobes, 
and densely branched, short rhizines. Externally it is very similar 
to and has been misidentified as P. caraccensis Tayl., which differs 
significantly in producing usnic and norstictic acids. It is also 
related to the Hawaiian species P. cervicornis Tuck. in that it produces 
the same chemical substances. The thallus of P. gigas, however, is much 
larger and the lobes ordinarily wider than in P. cervicornis. The 
rhizines of P. gigas, furthermore, are short, usually about 1 mm. long, 
and form a dense mat on the lower side, whereas the rhizines of P. 
cervicornis are often more than 2 mm. long and relatively sparse. 
Parmelia gigas is widely distributed in tropical America, being 
especially common at higher elevations (up to 3900 m.) in cloud 
forests. 

Additional specimens examined: Mexico: Oaxaca: Trail leading to 
Cerro san Felipe, elev. 2920-3000 m., Hale 20707, 20723, 20783, 21100 
(US). Panama: Craters edge, Vole4n Chiriquf, Prov. Chiriqui, Schol- 
ander, Dec. 12, 1948 (MO). Venezuela: Pan de Aztcar, Cordillera de 
Mérida, elev. 4000 m., Jahn 240 (US). Colombia: Carrizales, north 
of Las Brisas, Cordillera Occidental, Dept. Valle, Cuatrecasas 20545 
(US); eastern slope of Paramo de las Coloradas, Dept. Santander, 
elev. 3300-3900 m., Killip & Smith 18394, 18481 (US); Los Gaques, 


178 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


western slope of Paramo de Guasca, Dept. Cundinamarca, elev. 3250 
m., Killip 34064B (US); without locality: Zindig 2573 (UPS). Ecua- 
dor: Paramo de Minza Chica, Tunguragua, elev. 3800 m., Penland 
404 (MO, US); Paramo de Volcén Cayambe, Dept. Pichincha, elev. 
13,000 ft., Prescott (MSC, US). 

17. Parmelia immaculata Kurokawa, sp. nov. 

Thallus adnatus, corticola, albido-cinerascens, 4-9 cm. latus, lobis 
sublinearibus, contiguis, 1-3 mm. latis, superne nitidulus, aetate 
rugosulus, sorediatus, soraliis laminalibus, subtus sparse rhizinosus, 
rhizinis nigris, brevibus, dichotome ramosis. Apothecia rara, sub- 
stipitata, 1.5-4.5 mm. diametro; hymenium 50-60 uw altum; sporae 
5-7 X9-10 p. 

Thallus adnate on bark, yellowish glaucous (R), 4-9 cm. across; 
lobes dichotomously branched, sublinear-elongate, contiguous, 1-3 mm, 
wide, 110-200 u thick, the margins smooth or sparsely lobulate with 
age; upper surface rather shiny, plane, becoming rugulose and irregu- 
larly cracked on older lobes, sorediate, the soralia orbicular, laminal, 
often fusing; medulla white; undersurface sparsely rhizinate, minutely 
rugulose, black, the rhizines black, short, dichotomously branched. 
Apothecia more or less stipitate, 1.5-4.5 mm. in diameter, exciple 
crenate, amphithecium sorediate, disc vandyke brown (R), radially 
split; hymenium 50-60 yp high; spores 5-79-10 uy. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ rose, P—, 
pigment if present K+ purple, atranorine, KC-+ unknown, and 
rhodophyscin present. 

Type in the Botanical Museum, Lund University, collected at 
Punch Bow] Inn, north of Louis Trichardt, Zoutpansberg, Transvaal, 
Union of South Africa, Oct. 11, 1953, by Ove Almborn (no. 6551; 
isotype in US). 

Parmelia immaculata belongs to that group of species near P. dactyl- 
ifera Vain. and P. livida Tayl., all of which contain the same unknown 
KC+ substance. Parmelia dactylifera is pustulate-isidiate, whereas 
P. livida lacks pustules and soredia. Parmelia immaculata is appar- 
ently a very common corticolous lichen in South Africa, occurring 
frequently with P. leiophylla Kurokawa. <A single specimen from 
Java also seems to belong to this species. 

Additional specimens examined: Africa: South of Felixburg, South- 
ern Rhodesia, Héeg, April 2, 1930 (TRH); Punch Bowl Inn, north of 
Louis Trichardt, Zoutpansberg, Transvaal, Almborn 6440, 6550 (LD), 
6247 (LD, US); Polela Forest, Polela, Natal, Almborn 9511 (LD, US); 
Stormsrivier, Humansdorp, Almborn 4132 (LD); Grootvadersbosch, 
Swellendam, Cape Province, Almborn 2194, 2195, 2196 (LD) ; Grahams- 
town, Albany, Cape Province, Héeg, December 1929 (TRH). Java: 
Tjibodas, Neervoort 3140 (BO). 


HALE & KUROKAWA—SUBGENUS PARMELIA 179 


18. Parmelia infirma Kurokawa, sp. nov. PLATE 7 


Thallus laxe adnatus, corticola, olivaceo-albicans, 3-8 cm. diametro, 
lobis sublinearibus, subimbricatis, 1.0-3.5 mm. latis, margine sub- 
crenatis, superne nitidus, emaculatus, isidiatus, subtus niger, breviter 
rhizinosus, rhizinis nigris, ramosis. Apothecia adnata, 1.0-2.5 mm. 
diametro; hymenium 60-85 » altum; sporae 9-13%15-19 p . 

Thallus loosely adnate on bark, yellowish glaucous (R), 3-8 cm. 
across; lobes irregularly branched, sublinear, sometimes imbricate, 
1.0-3.5 mm. wide, 130-170 » thick, the margins more or less crenate, 
sometimes blackrimmed; upper surface shiny, without maculae, 
isidiate, the isidia cylindrical, moderate, simple to coralloid branched ; 
medulla white; undersurface black, short-rhizinate, the rhizines 
black, shiny, branched. Apothecia adnate, 1.0-2.5 mm. in diameter, 
exciple entire, amphithecium usually isidiate, disc seal brown (R); 
hymenium 60-85 y» high; spores 9-13 15-19 un. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC—, P-, 
atranorine, caperatic and protolichesteric acids present. 

Type in the National Science Museum, Tokyo, collected at Dar- 
jeeling, Kurseong, India, Apr. 12, 1960, by H. Hara et al. (isotype in 
US). 

This new species resembles P. koyaensis Asah. in external appearance, 
spore size, and presence of fatty acids. The isidia of P. infirma are 
shorter and sometimes become more or less granular. Parmelia 
koyaensis is P-++ red, containing protocetraric acid, which has not 
been demonstrated in P. infirma. The related isidiate species P. 
nodakensis Asah. is KC+ rose and P. erenata Kurokawa contains 
stictic acid (K+ yellow). Parmelia infirma also resembles the tropical 
American species P. costaricensis Nyl., which is distinctly maculate. 
Parmelia infirma is known only from the type locality. 


19. Parmelia leiophylla Kurokawa, sp. nov. PLaTE 9 


Thallus adnatus, corticola, cinereo-albicans, 3-7 cm. diametro, 
lobis sublinearibus, contiguis, 1-3 mm. latis, superne planus, nitidulus, 
aetate rugosus, pustulatus, medulla pro parte ochracea, subtus niger, 
rhizinosus, rhizinis nigris, dichotome ramosis. Apothecia rara, 
adnata, usque ad 1 cm. diametro; hymenium 50-55 » altum; sporae 
9-6 X 10-13 pu. 

Thallus adnate, corticolous, yellowish glaucous (R), 3-7 cm. across; 
lobes dichotomously branched, sublinear, contiguous, 1-3 mm. wide, 
130-180 u thick, the margins smooth; upper surface plane, rather shiny, 
rugulose on older lobes, pustulate, not distinctly sorediate, without 
maculae; medulla white, turning yellow ochre (R) under the pustules; 
undersurface black, rhizinate, the rhizines black, shiny, dichotomously 
branched, rarely projecting beyond the margins. Apothecia very 


180 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


rare, adnate, to 1 cm. in diameter, amphithecium rugose, disc split 
with age, rarely abnormally perforate; hymenium 50-55 yw high; 
spores 5-6 10-13 u. 

Reactions: Thallus K+ yellow; medulla K—, C+ deep red, KC+ 
red, P—, pigment K+ purple, atranorine, olivetoric acid, and rhodo- 
physcin present. 

Type in the Botanical Museum, Lund University, collected on 
Forest Drive from Houtbosch to Tzaneen, Pietersburg, Transvaal, 
Union of South Africa, Oct. 12, 1953, by Ove Almborn (no. 6796; 
isotype in US). 

This species resembles P. livida Tayl. in general habitus but is 
separated by the sorediate pustules and chemistry. Parmelia wm- 
maculata Kurokawa, which has similar distribution in South Africa, 
is distinctly sorediate and C—, KC+ rose. Parmelia formosana 
Zahlbr. is strictly pustulate and contains lichexanthone. The Ameri- 
can species P. croceopustulata Kurokawa has similar coarsely sorediate 
pustules and rhodophyscin, but the main component is protocetraric 
acid (C—, P-+ red). 

Additional specimens examined: Union of South Africa: Transvaal: 
Punch Bowl Inn, north of Louis Trichardt, Zoutpansberg, Almborn 
6435 (LD); Natal: South of Nkandhla, Nkandhla, Almborn 8074 (LD), 
Impetyne Forest near Weza, Alfred, Almborn 9969 (LD), Indumeni 
Forest, Cathedral Peak Area, Bergville, Almborn 9191 (LD); Cape 
Province: Deepwall Forest, Knysna, Arnell 1632a, 1645 (LD), Table 
Mountain, Arnell 861 (LD), Maas Geesteranus 12167 (L), along 
Skeleton Stream, east slopes of Table Mountain, Wynberg, Almborn 
11153 (LD), Fern Kloof, Albany, Almborn 10735 (LD). 


20. Parmelia lineariloba Kurokawa, sp. nov. PLATE 6 


Thallus adnatus, corticola, cinereo-albicans, 3-5 cm. diametro, lobis 
sublinearibus, elongatis, subdivaricatis, separatis, 0.5-2.0 mm. latis, 
superne planus, nitidus, emaculatus, isidiis sorediisque destitutus, 
subtus niger, rhizinosus, rhizinislongis, nigris, sparse ramosis. Apothe- 
cia ignota. 

Thallus adnate on bark, pale gull grey (R), 3-5 cm. in diameter; 
lobes dichotomously branched, linear-elongate, subdivaricate, sepa- 
rate, 0.5-2.0 mm. wide, 100-130 u thick, the margins smooth, narrowly 
black-rimmed; upper surface plane and smooth, shiny, without 
maculae, isidia and soredia lacking; medulla white; undersurface 
black, rhizinate, the rhizines black, long and sparsely branched, 
about 1 mm. long. Apothecia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ red, P—, 
atranorine and alectoronic acid present. 


HALE & KUROKAWA—SUBGENUS PARMELIA 181 


Type in Michigan State University, collected in pine forest at 
summit of Alto de la Bandera, Cordillera Central, La Vega, Dominican 
Republic, elev. 2800 m., Aug. 5, 1958, by C. Wetmore (no. 3475; 
isotype in US). 

Parmelia lineariloba has sublinear lobes that are quite narrow, long, 
and usually subdivaricate. The rhizines are rather long and only 
moderately branched. The chemically related subdivaricate species 
P. gigas Kurokawa and P. cervicornis Tuck. are much larger in size. 
Parmelia lineariloba is at present known only from the Dominican 
Republic, where it is found at high elevations in pine forests. We 
might expect it to be common in similar habitats elsewhere in the 
West Indies. 

Additional specimens examined: Dominican Republic: Ridge con- 
necting La Rucilla and Pico Trujillo, Macisco Central, elev. 8000 ft., 
Wetmore 3599, 3628 (MSC, US). 


21. Parmelia lividescens Kurokawa, sp. nov. 


Thallus adnatus, corticola, cinereo-albicans, 5-10 cm. diametro, 
lobis sublinearibus, 1.5-3.0 mm. latis, margine subcrenatis, superne 
nitidulus, sorediatus, soraliis subterminalibus, subtus sparse rhizi- 
nosus, niger, rhizinis nigris, dichotome ramosis. Apothecia adnata, 
2-3 mm. diametro; hymenium 40-50 » altum; sporae 5-79-11 p. 

Thallus adnate on bark, yellowish glaucous (R), 5-10 em. across; 
lobes dichotomously branched, sublinear, separate at the circumfer- 
ence, 1.5-3.0 mm. wide, 140-180 4 thick, the margins subcrenate; 
upper surface shiny on younger lobes, dull and irregularly cracked on 
older lobes, sorediate, soralia orbicular, laminal and subterminal; 
medulla white; undersurface sparsely rhizinate, black, the rhizines 
black, dichotomously branched. Apothecia adnate, 2-3 mm. in 
diameter, disc saccardo’s brown (R), radially split; hymenium 40-50 
high; spores 5-79-11 up. 

Reactions: Thallus K+ yellow; medulla K—, C+ red, KC+ red, 
P—, atranorine and olivetoric acid present. 

Type in the Botanical Museum, Lund University, collected on bark, 
Polela Forest, Polela, Natal, Union of South Africa, Nov. 10, 1953, by 
Ove Almborn (no. 9490; isotype in US). 

This species resembles P. revoluta Flk. in the presence of subterminal 
capitate soralia and in the absence of maculae. Parmelia revoluta, 
however, produces gyrophoric acid and has more sparsely branched 
rhizines. Parmelia leiophylla Kurokawa has the same chemical con- 
tent as P. lindescens but differs in having laminal sorediate pustules. 

Additional specimen examined: Union of South Africa: 6 mi. 
north of Knysna, Knysna, Cape Province, Almborn 2665 (LD). 


182 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


22. Parmelia monilifera Kurokawa, sp. nov. PLATE 9 

Thallus adnatus, corticola, cinereo-albicans, ca. 12 cm. diametro, 
lobis sublineari-elongatis, subdivaricatis, 2-6 mm. latis, superne 
planus, nitidus, emaculatus, isidiis sorediisque destitutus, subtus 
niger, dense rhizinosus, rhizinis nigris, ramosis, saepe moniliformibus. 
Apothecia ignota. 

Thallus loosely adnate on bark, grayish white, about 12 cm. in 
diameter; lobes dichotomously branched, sublinear-elongate, sub- 
divaricate, 2-6 mm. wide, 200-250 » thick, the margins smooth; 
upper surface smooth, plane to concave, shiny, without maculae, 
isidia and soredia lacking; medulla white; undersurface black, densely 
rhizinate, the rhizines black, shiny, branched, often moniliform or 
opuntioid, projecting as a mat beyond the margins. Apothecia not 
seen. 

Reactions: Thallus K+ yellow; medulla K—, C— or C+ pale 
orange, KC+ deep orange red, P—, atranorine, barbatic acid, and 
an unknown KC-+ substance present. 

Type in the Botanische Staatssammlung, Munich, collected at 
La Aguada, Mérida, Venezuela, Apr. 2, 1958, by K. Magdefrau 
(no. 651; isotype in US). 

This species resembles P. gigas Kurokawa, P. cervicornis Tuck., 
and P. exporrecta Kurokawa in having a large divaricate thallus and 
sublinear-elongate lobes. It is clearly separated by the coarse, 
branched, moniliform or opuntioid rhizines, a type that we have 
not seen in any other species of Parmelia. It is especially distinct 
chemically, since related species have alectoronic acid, or less com- 
monly gyrophoric acid. The species is known only from the type 
locality in Venezuela. 

23. Parmelia prolongata Kurokawa, sp. nov. PLATE 6 

Thallus laxe adnatus, corticola, olivaceo-albicans, 5-10 cm. dia- 
metro, lobis sublinearibus, 1.5-4.0 mm. latis, margine partim isidiato- 
dissectis, superne planus, nitidus, exigue albomaculatus, sparse 
isidiatus, medulla alba vel pallide flavescens, subtus niger, dense 
rhizinosus, rhizinis nigris, dense ramosis. Apothecia ignota. 

Thallus loosely adnate on bark, sea-foam yellow (R) turning olive 
buff in the herbarium, 5-10 cm. across; lobes dichotomously branched, 
sublinear, elongate, sometimes subimbricate, 1.5-4.0 mm. wide, 100- 
200 » thick, the margins smooth or in part with isidial branchlets 
or isidia, often elongate, ascending or suberect, dorsiventral or 
becoming cylindrical and coralloid branched; upper surface plane 
and smooth, weakly maculate, rather shiny, often sparsely isidiate, 
the isidia cylindrical, variable, sometimes branched; medulla white 
to cartridge buff (R); undersurface black, densely rhizinate, the 


HALE & KUROKAWA—SUBGENUS PARMELIA 183 


rhizines black, shiny, densely branched, often forming a mat. Apo- 
thecia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C+ deep red, KC-++ 
red, P—, atranorine, a C+ unknown substance, and frequently a 
pale-yellowish pigment present. 

Type in Michigan State University, collected on ridge leading to 
Pic Macaya, Morne Macaya, Dept. du Sud, Haiti, elev. 6400-7000 ft., 
July 27, 1958, by C. Wetmore (no. 3233; isotype in US). 

Parmelia prolongata is characterized by peculiar laminal and mar- 
ginal isidia which are cylindrical at first but which at length become 
dorsiventral, suberect, and more or less branched, and as much as 
5 mm. long. However, the production of isidia and lobules is ex- 
tremely variable and they may be reduced to small marginal isidia. 
All specimens have in common a deep C+ reaction caused by an 
unknown substance apparently related to olivetoric or anziaic acid. 
The pale-yellow pigment may or may not be clearly present and its 
identity is unknown. Parmelia prolongata is found at higher eleva- 
tions, often on conifer bark, in the Caribbean and Andean regions. 

Additional specimens examined: Mexico: Chiapas: 18 km. south- 
east of San Crist6ébal, Hale 20286 (US). Panama: Prov. Chiriqui: 
East side of Volcin Chiriqui, Scholander, Dec. 5-12, 1948 (US). 
Haiti: Ridge leading to Pic Macaya, Morne Macaya, Dept. du Sud, 
elev. 6400-7000 ft., Wetmore 3233 (MSC, US), 3286 (MSC). Colom- 
bia: Western slopes of Paéramo de Cruz Verde, Dept. Cundinamarca, 
3150 m., Cuatrecasas 450Z (US). 


24. Parmelia rhabdiformis Kurokawa, sp. nov. 


Thallus adnatus, corticola, olivaceo-albicans in herbario, ca. 8 cm. 
diametro, lobis sublinearibus, 2-6 mm. latis, subimbricatis, margine 
plus minusve dissectis, superne planus, nitidus, emaculatus, isidiatus, 
subtus niger, dense rhizinosus, rhizinis nigris, dense ramosis. Apo- 
thecia adnata, 1-5 mm. diametro; hymenium 60-80 » altum; sporae 
6-8 X 15-21 yp. 

Thallus adnate on bark, turning cream buff (R) in the herbarium, 
about 8 cm. in diameter; lobes irregularly branched, sublinear, 2-6 
mm. wide, subimbricate, 150-200 » thick, the margins more or less 
dissected; upper surface plane, shiny, without maculae, isidiate, the 
isidia mostly simple, club-shaped, about 0.5 mm. high; medulla white; 
undersurface black, densely rhizinate, the rhizines black, densely and 
finely branched, forming a woolly mat. Apothecia adnate, 1-5 mm. 
in diameter, exciple crenate, amphithecium isidiate, disc sayal brown 
(R), sometimes radially split; hymenium 60-80 » high; spores 6-8 X 15- 
21 py. 

722-891 —64——-5 


184 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Reactions: Thallus K+ yellow; medulla K+ yellow turning red, 
C—, KC—, P+ pale orange red, atranorine and norstictic acid 
present. 

Type in the U.S. National Herbarium, collected in the vicinity of 
El] Boquete, Prov. Chiriqui, Panama, Mar. 2, 1911, by W. R. Maxon 
(no. 5128a). 

Except for the production of norstictic acid, this species resembles 
P. infrma Kurokawa and P. crenata Kurokawa, both Asian species. 
The isidia of P. rhabdiformis are mostly simple and clavate and rather 
elongate, to 0.5 mm. high. It is known only from the type locality in 
Panama. 

25. Parmelia rigidula Kurokawa, sp. nov. 

Thallus adnatus, corticola, olivaceo-cinerascens, 7-12 cm. diametro, 
lobis sublinearibus vel subirregularibus, subimbricatis, usque ad 7 
mm. latis, margine subcrenatis, superne nitidulus, exigue albomacu- 
latus vel emaculatus, sorediis isidiisque destitutus, medulla pallide 
flavicans, subtus niger, rhizinosus, rhizinis nigris, modice ramosis. 
Apothecia adnata, 2-7 mm. diametro; hymenium 70-85 y altum; 
sporae 10-12%15-18 up. 

Thallus adnate on bark, pale olive buff (R), 7-12 cm. in diameter; 
lobes irregularly branched, sublinear to subirregular, more or less 
imbricate, up to 7 mm. wide, 210-250 » thick, the margins crenate, 
often narrowly black-rimmed; upper surface plane to concave, more 
or less shiny, weakly maculate or without maculae; medulla sea-foam 
yellow to pale-pinkish buff (R), K—; under surface black, moderately 
rhizinate, the rhizines black, matted, sparsely to moderately branched. 
Apothecia adnate, 2-7 mm. diameter, exciple smooth to undulate, 
dise dark livid-brown (R), sometimes radially split; hymenium 70-85 yu 
high; spores 10-12 15-18 up. 

Reactions: Thallus K-+ yellow; medulla K—, C—, KC—, P-, 
atranorine and a pale-yellow pigment present. 

Type in the National Science Museum, Tokyo, collected at Phalut, 
Darjeeling, India, June 6, 1960, by H. Hara (isotype in US). 

This species has a peculiar coriaceous thallus as in P. scytodes 
Kurokawa and P. scytophylla Kurokawa. However, it is distinguished 
by producing a pale-yellow-orange pigment in the medulla (apparently 
not entothein). Superficially it resembles P. subaurulenta Nyl., but 
the rhizines are sparsely to moderately branched, not simple. Only 
two specimens have been collected, both in northern India, where it 
is endemic. 

Additional specimen examined: India: Tongloo, Himalayas, Watt 
7035 (BM). 


HALE & KUROKAWA—SUBGENUS PARMELIA 185 


26. Parmelia scytodes Kurokawa, sp. nov. PLATE 6 

Thallus adnatus vel laxe adnatus, corticola, cinereo-albicans, ca. 
20 cm. diametro, lobis sublinearibus, 2-5 mm. latis, superne planus, 
emaculatus, isidiis sorediisque destitutus, subtus niger, dense rhizino- 
sus, rhizinis nigris, modice ramosis. Apothecia ignota. 

Thallus adnate to loosely adnate on bark, rather coriaceous, pale 
smoke gray (R), about 20 cm. across; lobes irregularly branched, 
sublinear, 2-5 mm. wide, 150-200 uw thick, the margins subcrenate, 
narrowly black-rimmed; upper surface plane and smooth, becoming 
rugose on older lobes, without maculae, isidia and soredia lacking; 
medulla white; undersurface black, moderately to densely rhizinate, 
the rhizines black, shiny, moderately branched. Apothecia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ orange, 
P+ orange red, atranorine, barbatic acid, and an unidentified P+- 
acid present. 

Type in the National Science Museum, Tokyo, collected at Dar- 
jeeling, Batasi, India, May 6, 1960, by Togashi (isotype in US). 

This new species is characterized by the large rather coriaceous 
thallus. Externally it resembles P. fissicarpa Kurokawa, an African 
species with a P+ reaction but no barbatic acid. Parmelia bostry- 
chodes Zahlbr. and P. physcioides Nyl., both of which produce barbatic 
acid without a P-+ substance, have smaller sublinear lobes. Parmelia 
scytodes is known only from the type locality in India. 


27. Parmelia scytophylla Kurokawa, sp. nov. Puate 7 


Thallus adnatus, corticola, cinereo-albicans, 5-14 cm. diametro, 
lobis sublinearibus vel subirregularibus, margine subcrenatis, superne 
planus, nitidus, exigue albomaculatus, isidiis sorediisque destitutus, 
subtus niger, modice rhizinosus, rhizinis nigris, sparse ramosis. Apo- 
thecia adnata, ca. 10 mm. diametro; hymenium 35-404 altum; 
sporae 5-6 X89 u. 

Thallus adnate to loosely adnate on bark, yellowish glaucous to 
pale olive gray (R), 5-14 cm. in diameter; lobes irregularly branched, 
sublinear to subirregular, 2-6 mm. wide, 250-350 u thick, the margins 
more or less crenate, narrowly black-rimmed; upper surface plane 
and smooth, shiny, faintly maculate, isidia and soredia lacking; 
medulla white; undersurface black, moderately rhizinate, blackish 
brown and naked or papillate in a narrow zone around the tips, the 
rhizines black, shiny, sparsely branched. Apothecia adnate to sub- 
stipitate, often more than 10 cm. in diameter, exciple undulate, 
amphithecium smooth, faintly maculate, disc warm blackish brown 
to light seal brown (R); hymenium 35—40 uw high; spores 5-6 X8-9 u. 

Reactions: Thallus K+ yellow; medulla K—, C+ rose, KC+- 
red, P—, atranorine and gyrophoric acid present. 


186 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Type in the National Science Museum, Tokyo, collected at 
Migotang, Sikkim, at 3900 m. elevation, May 30, 1960, by H. Hara 
(isotype in US). 

Parmelia scytophylla is very similar to two other species known 
only in the Himalayan region, P. rigidula Kurokawa and P. scytodes 
Kurokawa. All have rather large coriaceous. thalli. Parmelia 
scytodes contains barbatic acid and a P+ unknown, and P. rigidula 
contains a pale-yellow pigment and has larger spores, 15-18 yu long. 

Additional specimens examined: Nepal: Langtang Village, Polunin 
M32 (BM). India: Nilgherries, Watt s.n. (BM); Darjeeling, Awasthi 
12 (H). 


28. Parmelia thysanota Kurokawa, sp. nov. 


Thallus laxe adnatus, corticola, cinereo-albicans, 5-13 cm. latus, 
lobis sublinearibus, subdivaricatis, separatis, 2-5 mm. latis, superne 
planus, nitidulus, apice leviter albopruinosus, sorediatus, soraliis 
subterminalibus, subtus niger, dense rhizinosus, rhizinis nigris, 
dense ramosis. Apothecia ignota. 

Thallus loosely adnate on bark, pale-glaucous green (R), 5-13 cm. 
in diameter; lobes dichotomously branched, linear or sublinear- 
elongate, subdivaricate, separate at the circumference, 2-5 mm. wide, 
140-180 uw thick, the margins entire, narrowly black-rimmed; upper 
surface plane, rather shiny, often pruinose near the tips, sorediate, 
soralia laminal and subterminal, often protruding, coalescing; medulla 
white; undersurface densely rhizinate, black, the rhizines black, 
shiny, densely branched, usually projecting in a narrow dense mat 
beyond the margins. Apothecia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C+ rose, KC+- 
red, P—, atranorine and gyrophoric acid present. 

Type in the United States National Herbarium, collected in pine-oak 
forest on lava, 11 km. east of Las Vigas on Highway 140, Veracruz, 
Mexico, Mar. 11, 1960, by M. E. Hale (no. 20943). 

Parmelia thysanota is very closely related to P. exporrecta Kuro- 
kawa in having a subdivaricate thallus and in producing gyrophoric 
acid. It differs in having subterminal soralia; P. exporrecta lacks 
any soralia. It is probable that these two species represent a sore- 
diate-nonsorediate phase, but the material at hand is not sufficient 
for a final decision. 

Additional specimen examined: Mexico: Same locality data as 
the holotype, Hale 21118 (US). 


29. Parmelia virginica Hale, sp. nov. PLATE 5 

Thallus adnatus, albido-cinerascens, nonnihil fragilis, corticola, 
5-7 cm. diametro, lobis sublinearibus, 1.5-2.5 mm. latis, margine 
centrum versus paulo lobulato-dissectis, superne nitidus, minute 


HALE & KUROKAWA—SUBGENUS PARMELIA 187 


rugulosus, aetate valde irregulariter rugosus, cortice fragili, 
pustulescente, sorediis destitutus, subtus niger, modice rhizinosus, 
rhizinis modice dichotome ramosis. Apothecia ignota. 

Thallus adnate, mineral gray, rather fragile, corticolous; lobes sub- 
linear, 1.5-2.5 mm. wide, 110-130 u thick, more or less subimbricate, 
towards the center lobulate-dissected; upper surface shiny, minutely 
rugulose, strongly rugose with age, the cortex fragile and breaking 
away, forming coarse pustules, soredia lacking; medulla white; under- 
surface black, brown near the tips, moderately rhizinate, the rhizines 
black, moderately branched. Apothecia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ orange, 
P—, atranorine, barbatic acid, and an unknown KC-+ substance 
present. 

Type in the U.S. National Herbarium, collected at Hawksbill, 
Shennandoah National Park, Page Co., Virginia, elev. 1200 m., 
June 26, 1962, by M. Hale (no. 21592; isotypes in DUKE, LD, M, 
MSC, REN, 5, TNS, WISC). 

Parmelia virginica is virtually identical with P. taylorensis Mitch. 
(P. rugosa Tayl.non Fr.) in regard to the fragile cortex and the develop- 
ment of pustules. The pustules originate along the ridges of wrinkles 
and are therefore usually elongate. Parmelia taylorensis (cf. Mitchell, 
1962; Hale, 1962) is typically a European species which differs in 
having evernic and lecanoric acids and more richly branched rhizines. 
The type collection of P. virginica was made on exposed dwarf spruce 
trunks. It is probably common in similar habitats in the Appa- 
lachians from Virginia through North Carolina. 

Additional specimen examined: North Carolina: Lich-Log ridge, 
Blue Ridge Parkway, McDowell Co., elev. 4660 ft., Becking 5908031 
(US). 


Bibliography 


Acuarius, E. 1794. Férsék til en férbattrad lafvarnes indelning. K. Vet. Acad 
Nya Handl. 237-259. 

ASAHINA, Y. 1952. Lichens of Japan II. Genus Parmelia. Res. Inst. Nat. 
Resources. 162 pp. Tokyo. 

and S. Sarpata, 1954. Chemistry of lichen substances. Japan Soc. for 
Promotion of Science. 240 pp. Tokyo. 

CuLBEerson, W. L. 1962. Some pseudocyphellate Parmelias. Nova Hedw. 
4:564-577. 

Du Ruietz, G. E. 1924a. Die soredien und isidien der Flechten. Svensk Bot. 
Tidskr. 18: 371-396. 

1924b. Flechtensystematische Studien. IV. Bot. Not. 1924:329-342. 

Fries, E. 1825. Systema orbis vegetabilis. Lund. 


188 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Gyeunik, V. 1932. Additamenta ad cognitionem parmeliarum III. Repert. 
Sp. Nov. Fedde 30:209-226. 

1933. Lichenes varii novi criticique. Acta Faun. Fl. Univ. I, 
1, no. 5,6:3-10. 

Haus, M. E., Jr. 1960. A revision of the South American species of Parmelia 
determined by Lynge. Contr. U.S. Nat. Herb. 36, pt. 1:1-41. 

and 8S. Kurokawa. 1962. Parmelia species first described from the 
British Isles. Lichenologist 2:1-5. 

Hiutmann, J. In Rabenhorst, Kryptogamenflora. 1934. Familie Parmeliaceac. 
9(5), part 3:1-309. 

ImsuaucG, H. A. 1957. Alpine lichens of the western United States and adjacent 
Canada I. The macrolichens. Bryologist 60:177-272. 

Lynaz, B. 1914. Die Flechten der ersten Regnellschen Expedition. Die 
Gattungen Pseudoparmelia gen. nov. und Parmelia Ach. Ark. Bot. 13, no, 
13:1-172. 

Maas GEEsTERANUS, R. A. 1947. Revision of the lichens of the Netherlands. 
I. Parmeliaceae. Blumea 6:1-199. 

Mircuetyt, M. E. 1961. L’element eu-oceanique dans la flore lichenique du 
sud-ouest de V’Irlande. Revista Biol. 2, no. 3-4:177-256. 

RASANEN, V. 1943. Das System der Flechten. Act. Bot. Fenn. 33:1-82. 

SANTESSON, R. 1944. Pseudoparmelia Lynge, a lichen genus to be rejected. 
Svensk Bot. Tidskr. 36:471-474. 

ScurEBER, J. 1791. Linné, Genera plantarum, ed. 8. 

Varnio, E. A. 1890. Etude sur la classification naturelle et la morphologie des 
lichens du Brésil. Acta Soc. Faun. Fl. Fenn. 7, no, 7:1-240. 

1923. Lichenes Insularum Philippinarum, IV. Ann. Acad. Sci. Fenn. 

Ser. A, 19, no. 15:1-84. 


Index 


(To species and varieties of subgenus Parmelia. 


Synonyms in 


italics; page numbers of principal entries in boldface.) 


abstrusa, 141, 145 
abstrusoides, 149 

acrobotrys, 141, 142, 143, 145, 147 
affixa, 137 

amazonica, 129, 147, 148, 149, 157 
americana, 159 
anaptychioides, 162, 165 
annexa, 149, 151, 152, 155, 156 
antillensis, 131 

apophysata, 136, 138 
appalachensis, 129 

aptata, 150 

arcana, 149, 151, 155 
atrichella, 137, 

aurulenta, 124, 130, 131, 132 
bahiana, 163, 164 

balansae, 130 

bicornuta, 135, 137 

bogotensis, 161 

boliviana, 164 

bolliana, 129 

borreri, 124, 128, 129 
borrerina, 129 

bostrychodes, 164, 185 
brasiliana, 127, 129, 154, 159, 164, 174 
brasiliana var. erythrodes, 173 
brevirhiza, 162, 166 
bulbochaeta, 137, 138 
canaliculata, 129 

canescens, 163, 167 

caperata, 148, 149 

caraccensis, 163, 177 

caribaea, 150, 152, 153 
carneopruinata, 149 
caroliniana, 148, 149 
carporrhizans, 125, 132 
cervicornis, 165, 171, 177, 181, 182 
cetrarioides, 128, 148 

cetrata, 124, 129, 130 
chapadensis, 150 

cinerascens, 149 


circumnodata, 123, 141, 145 

citrella, 163, 168 

concrescens, 149 

confoederata, 137 

connivens, 141, 142 

consimilis, 161 

continua, 136 

coronata, 137, 139 

coronata var. denudata, 137 

costaricensis, 160, 161, 172, 179 

crenata, 161, 168, 172, 179, 184 

croceopustulata, 162, 169, 180 

crozalsiana, 149 

cryptochlorophaea, 149 

culmigena, 123, 161, 173 

cyphellata, 147, 148, 149 

dactylifera, 123, 161, 178 

damaziana, 165 

decurtata, 136, 139 

degelii, 165, 170 

denegans, 132 

densirhizinata, 163, 170, 171 

dentella, 161, 172 

denudata, 137 

digitata, 123, 161, 173 

dissecta, 123, 131, 132, 133, 134, 135, 
148 

domingensis, 141 

dubia, 129 

ecaperata, 149 

enderythraea, 163 

endochlora, 124, 161, 162 

endomiltoides, 159 

ensifolia, 123, 161, 171, 172, 173 

entotheiochroa, 133 

eruptens, 148, 153 

erythrodes, 163, 173 

eximbricata, 142, 143 

exporrecta, 164, 174, 182, 186 

exsecta, 162 

exsplendens, 135, 162, 163, 170, 174 


189 


190 


fissicarpa, 164, 175, 185 
flavida, 163 

flavovirens, 162, 176 
formosana, 124, 162, 169, 180 
fraudans, 128 

fungicola, 136 

galbina, 125, 132 

gigas, 165, 171, 174, 177, 181, 182 
gracilescens, 165 
heterochroa, 150, 154 
homogenes, 133 
homotoma, 124, 130 
horrescens, 123, 131 
hypocraea, 136 
hypomilta, 150 
hypoxantha, 154 

ikomae, 131 

imbricatula, 161 
immaculata, 163, 178, 180 
immiscens, 133 
imperfecta, 148, 155, 156 
infirma, 161, 179, 184 
insensitiva, 128 
insinuans, 164, 176 
intercalanda, 165 
internexa, 131 

intertexta, 150 

ischnoides, 149, 155 
isidiocera, 131, 160 
isidiosa, 147 

isidiza, 136, 141 

isidiza var. domingensis, 140 
keitauensis, 164, 176 
koyaensis, 161, 169, 179 
laevigata, 124, 162 
laevigatula, 136 
lecanoracea, 150 
leiophylla, 162, 169, 178, 179, 181 
leucopis, 124, 150 
limbata, 141, 142, 147 
limbata f. isidiosa, 147 
lindmannii, 131 
lineariloba, 165, 171, 180 
livida, 154, 165, 170, 174, 180 
lividescens, 162, 181 
luteoviridis, 141, 144 
lythgoeana, 163 
macrocarpoides, 130 
majoris, 162, 166 
malaccensis, 149 

malmei, 162 

manilensis, 164, 176 
martinicana, 149 


INDEX 


meiosperma, 149 
meizospora, 137 
melanochaeta, 132, 133 
metarevoluta, 125, 132, 175 
meyeri, 161 
michauxiana, 132 
microblasta, 161 
microsticta, 129 
minarum, 133, 148 
molybdiza, 150, 151, 152, 155, 158 
monilifera, 164, 182 
muelleri, 132, 148 
mutata, 124, 132 
nairobiensis, 153 
nodakensis, 161, 179 
nylanderi, 132 

obsessa, 125, 131 
omphalodes, 128 
osseoalbida, 164 
osteoleuca, 163 
owariensis, 149, 155, 156 
papyrina, 136 

perforata, 130 
perisidians, 131 
peruviana, 124, 163 
phlyctina, 132 
physcioides, 164, 185 
pilosa, 130 

planiuscula, 141, 144, 145 
pluriformis, 164 
prolongata, 160, 161, 182 
pruinata, 132 
pseudoborreri, 129 
pseudosinuosa, 162, 166 
pseudosulcata, 128 


.pulvinata, 165 


pustulata, 137, 140 
pustulescens, 149, 156 
quercina, 131, 132 
rahengensis, 149 
ramosissima, 141, 143, 145, 147 
raunkiaeri, 148 
reducens, 163, 168 
relicina, 135, 141 
relicinella, 142 
relicinula, 141 
reticulata, 124, 129, 130 
revoluta, 162, 163, 181 
revolutella, 165 
rhabdiformis, 161, 183 
rigidula, 163, 184, 186 
rockii, 162, 163 
rodriguesiana, 150 


INDEX 191 


rudecta, 129 subglandulifera, 136 
rugosa, 187 subisidiosa, 130 

rupicola, 150 sublaevigata, 164, 167 
rutidota, 150 sublanea, 141, 142, 146. 147 
salacinifera, 149, 157 sublimbata, 141, 146 
samoensis, 142, 143 subrudecta, 129 

samoensis var. eximbricata, 143 subsaxatilis, 164 

saxatilis, 124, 127, 128, 160 subtiliacea, 150 

schiffneri, 135, 137 suffixa, 135, 136 
schizospatha, 141, 144, 146 suleata, 128 

scortella, 136 tabacina, 156, 159, 140 
scrobicularis, 150 taylorensis, 165, 187 
seytodes, 164, 184, 185, 186 tenuirima, 128 

scytophylla, 164, 184, 185 texana, 148, 150, 153 
semilunata, 135, 137 thysanota, 165, 174, 186 
sensibilis, 1387 tiliacca, 123, 124, 130, 131, 132 
setschwanensis, 136 tiliaeea var. convexula, 154 
signata, 125 tiliacea var. rugulata, 7/54 
silvatica, 151, 165 tiliacea var. stenophylla, 154 
sinuosa, 162, 176 tortula, 150, 157 
somalicnsis, 150 tumescens, 141, 145, 147 
soredians, 148, 149 usambarensis, 131 
sorocheila, 159 velloziae, 162, 176 
spathulata, 131, 133, 134 ventricosa, 135, 136, 140, 141 
sphaerospora, 148, 150 versiformis, 124, 132 
spumosa, 124, 132, 155 violacca, 150, 158 
subabstrusa, 141 virginica, 162, 186 
subaffinis, 162, 166 viridescens, 137, 139 
subaurulenta, 135, 184 wallichiana, 131 
subcoronata, 136 xantholepis, 132 
subdissecta, 136, 138 xanthomelaena, 150 
subfatiseens, 132, 134, 135 zahlbruckneri, 164 


CONTR. NAT. HERB. VOL. 36 HALE & KUROKAWA PLATE 1 


1. Pustules of Parmelia aurulenta Vuck. (10). 2. Reticulate upper cortex of Parnrelta 
reticulata "Vavl. (10). 3. Marginal cilia of Parmelia melanochaeta Kurokawa (0<5). 
4. Effigurate pseudccy phellae of Parmelia saxatilty (1..) Ach. (05). 5. Pseudoeyphellae of 
Parmelia subrudecta Nvl. (<5). 6. Bulbate cilia of Parmelta tabacina Mont. & v.d. Bosch. 
(5). 7. Parmelia buthochaeta ale Gsowpe, Aceastht 4347, US, >< 2). 


CONTR. NAT. HERB. VOL. 36 HALE & KUROKAWA PLATE 2 


I. Parmelia luteoviridis Kurokawa Csoty pe, Meljer BIYES, US, 022). 2. Parmelta : 
wns Kurokawa (holotype, Moorland 4726, hk, 2). 3. Parmelia acrobeirys Kurokawa (se 
type, Neervoort 60. pop, US, 2). 4. Parmelia ramosioima Kurokawa (isotype, 
Buccalda 4583, US, 02). 3. Parinelia schs patha Kurokawa Gsotrpe, Neersvert 1062, 


US, >. 2). 


CONTR. NAT. HERB. VOL. 36 HALE & KUROKAWA—PLATE 3 


a 


lL. Parmelia caribaea Wale Gsotype, Le Gallo 494, US, 2<2). 2. Parmelia brevirhisa 
Kurokawa (holotwpe, Saatesson 2066, S, 1). 3. Parmelia erwptens Kurokawa (isotype, 
Almborn 7498, LD, 2). 4. Parmelia melanochaeta Kurokawa (holotype, Aalme 2243, 


5, 2). 


CONTR. NAT. HERB. VOL. 36 HALE & KUROKAWA—PLATE 4 


1. Parmelia annexa Warckawa (holotype, dlmvorn 56083, LD. 2). 2. Parmelia pustu- 
lescens Kurokawa CGsoty pe, Develius, Feb. 1s, 1960, DEGEL 5). 3. Parmelta tschnoides 
Kurokawa (holotype, Miborn 169s, LD, 02). 4. Parmelia arcana Kurokawa (isotype, 
Develrus, Web. 3, 1960, DEG (82). 5. Parmelia tortula Waurokawa (holotype, Almborn 


#805, LD, <2). 


CONTR. NAT HERB. VOL. 36 HALE & KUROKAWA —~PLATE 5 


1. Parmelia degelii Wale Cholotwpe, Degelius, Feb. 16, 1960, DIGIL, <2). 2. Parmelta 
anaptychivides Kurokawa (holotype, /etmore 3617, NISC, 1'4). 3. Parmelia virginica 
ILlale (holotype, Male 21502, Us, 2). 4. Parmelta dentella Vale & Kurokawa (holotype, 


MeCullough 370, Us, <2). 


CONTR. NAT. HERB. VOL. 36 HALE & KUROKAWA-— PLATE 6 


I. Parmelia scytodes Kurokawa Gsotype, Togasht, May 6, 1960, US, 1). 2. Parmelia 
lineartloba Kurokawa (holotype, Metnore 3475, MSC, <2). 3. Parmelia ex porrecta Kuro- 
kawa (holotype, Hale 20722, US, 1). 4. Parmelia prolongata Kurokawa (holotype, Met- 


more 3233, NISC, <4), 


CONTR. NAT. HERB. VOL. 36 HALE & KUROKAWA PLATE 7 


lL. Parmelia enstfolia Kurokawa (detail showing lobules) (holotype, Jmshaug 23430, 
MSC, 02). 20 Parmelia scytophylla Kurokawa (isotype, fara, May 30, 1960, US, >< 1). 
3. Parmelia fissicarpa Kurokawa (holotype, d/mborn 9374, LD, <1). 4. Parmelia infirma 

) 


Kurokawa (isotype, Hara, Apr. 12, 1960, US, > 2). 


CONTR. NAT. HERB. VOL. 36 HALE & KUROKAWA ~PLATE 8 


lL. Parmelia flavovirens Kurokawa (holotype, Santesson 5681, S, %1!4). 2. Parmelia 
denstrhisinata Kurokawa (holotype, Scholander, Dec. 5-12, 1948, US, > 3/4). 3. Parmelia 


gigas Kurokawa (holotype, A¢riste B70, US, >< 1). 


CONTR. NAT. HERB. VOL. 36 HALE & KUROKAWA ~~ PLATE 9 


lL. Parmelia monilitera” Nurokawa (holotype, Magdefraw O51, \I 1). 2.) Parmelta 


eroceo pustidata Kurokawa (holowpe, fmshaug 22275, MSC, 1). 3. Parmelta leto phylla 
Kurokawa (holotype, fmborn 6796, LD. 3/4). 4. Parmelta salacinifera Wale (holotype. 


Rapp larch 1925, US. <1) 


UNITED STATES NATIONAL MUSEUM 


CONTRIBUTIONS FROM THE UNITED States NATIONAL HERBARIUM 


VoLuME 36, Part 5 


A MONOGRAPH OF PARMELIA 
SUBGENUS AMPHIGYMNIA 


By Mason E. Hate, Jr. 


BULLETIN OF THE UNITED States NationaL MusEUM 
SMITHSONIAN INSTITUTION e WASHINGTON, D.C. e 1965 


A MONOGRAPH OF PARMELIA SUBGENUS AMPHIGYMNIA 


By Mason E. Hate, Jr. 


Introduction 


Parmelia has strangely been overlooked in the recent burst of 
monographic activity in lichens. It is a common, easily recognized 
foliose genus, well represented in most herbaria, with over 1,000 
names in the literature. Traditionally the European lichenologists 
have passed it over as an “easy” genus and concentrated on the more 
difficult crustose groups. It must be understood, of course, that 
Europe has a very poor Parmelia flora, especially in Amphigymnias, 
certainly not one to attract the attention of a serious monographer. 

This monograph deals only with subgenus Amphigymnia (Vain.) 
Dodge, a remarkably homogeneous group of 106 species. Studies 
are concurrently under progress on subgenus Parmelia (Hale & 
Kurokawa, 1964) and subgenus Xanthoparmelia (Vain.) Hale, two 
distinct groups with approximately 275 and 80 species, respectively. 
As here delimited, Amphigymnia is characterized by broad, more or 
less rotund lobes, 6 to 20 mm. wide, bearing unbranched, often sparse 
and coarse rhizines on the lower side, with a broad distinct bare zone 
around the margins. The apothecia are typically pedicellate and 
rather often have perforate discs, characters virtually unknown in 
other groups of the genus. The species are not difficult to separate; 
characters of an absolute or qualitative nature predominate. There 
is little of the troublesome intergradation between species that makes 
genera such as Usnea or Collema so difficult. 

Some overlap and intergradation occurs between a few Amphigymnia 
species and those of subgenus Parmelia, especially sections Irregulares 
(Vain.) Vain. and Cyclocheila (Vain.) Ris., which may have rotund 
(not linear) lobes and a distinct though narrow, bare zone below. 
An excellent example is P. caperata (L.) Ach., a well-known species 
formerly placed in subgenus Amphigymnia but apparently better 
put into subgenus Parmelia (cf. Hale & Kurokawa, 1964). As a 
result, we have relieved ourselves of these problems here and con- 
centrated in subgenus Amphigymnia & very homogeneous group of 
species. 

193 


194 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


The methods of study in this monograph were first to survey the 
literature as completely as possible and second to examine as many 
holotypes as possible. -Most of the original descriptions were repro- 
duced by photostat and assembled in notebooks for rapid reference. 
All references were taken from the original publications. Proper 
citations of the specimens and the location of the types were deter- 
mined and the available type specimens examined. 

Personal fieldwork has included studies of the flora in much of the 
United States, especially from Virginia to Texas, eastern Canada, 
and southern Mexico. About half of the 106 species were actually 
observed in the field. Fortunately, contemporary lichenologists have 
made extensive and excellent collections in the West Indies (Plitt, 
Imshaug, Wetmore), Chile (Santesson), Africa (Almborn, Degelius, 
des Abbayes, Maas Geesteranus, Héeg), Java (Groenhart), and Japan 
(Asahina, Kurokawa). The most serious gaps are in tropical South 
America, Australia, the Philippines, Borneo, New Guinea, and China. 
It is estimated that exhaustive fieldwork would turn up another 10 
or 20 new species, but it is doubtful that the subgenus will ever be 
found to contain more than 130 species, at least as delimited by the 
presently used taxonomic criteria. 

The following curators and directors have kindly loaned type 
specimens and other valuable collections: Dr. Sten Ahlner (Natur- 
historiska Riksmuseet, Stockholm), Dr. Reino Alava (Botanical In- 
stitute, Turku), Dr. Ove Almborn (Botanical Museum, Lund), Dr. Y. 
Asahina (Research Institute for Natural Resources, Tokyo), Dr. 
Charles Baehni (Conservatoire et Jardin Botaniques, Geneva), M. P. 
Bourrelly (Muséum National d’Histoire Naturelle, Paris), E. H. 
Bryan (Bernice P. Bishop Museum, Honolulu), Dr. A. Burkart 
(Instituto de BotAnico Darwinion, San Isidro), M. Choisy (Lyon), 
M. Skytte Christiansen (Botanical Museum, Copenhagen), Dr. G. 
Cufodontis (University of Vienna, Vienna), Dr. W. L. Culberson 
(Duke University, Durham), Dr. H. des Abbayes (Université de 
Rennes, Rennes), Dr. Henry Imshaug (Michigan State University, 
East Lansing), Mr. Peter James (British Museum, London), Dr. A. 
Kostermanns (Herbarium Bogoriense, Bogor), Dr. I. M. Lamb 
(Farlow Herbarium, Cambridge), Dr. R. A. Maas Geesteranus 
(Rijksherbarium, Leiden), Dr. P. J. Greenway (The East African 
Herbarium, Nairobi), Dr. J. Millar (Chicago Natural History Mu- 
seum, Chicago), Dr. G. Moggi (Istituto Botanico, Firenze), Dr. E. 
Miller (Institut fiir Spezielle Botanik, Ziirich), C. E. Palmar (Glas- 
gow Art Gallery and Museums, Glasgow), Dr. J. Poelt (Botanische 
Staatssammlung, Munich), Dr. K. Rechinger (Naturhistorisches Mu- 
seum, Vienna), Dr. C. Rizzini (Jardim Botanico, Rio de Janeiro), 
Dr. W. Robyns (Jardin Botanique, Bruxelles), Dr. C. Rogerson (New 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 195 


York Botanical Garden, New York), Dr. H. Roivainen (Botanical 
Museum, Helsinki), Dr. Rolf Santesson (Institute for Systematic 
Botany, Uppsala), Dr. R. Shaffer (University of Michigan, Ann 
Arbor), Sir George Taylor (Royal Botanic Gardens, Kew), Dr. J. W. 
Thomson (University of Wisconsin, Madison), Dr. K. Verseghy 
(Museum of Natural History, Budapest), Dr. R. Vincenzo (Istituto 
Botanico, Rome), Dr. E. F. Warburg (Oxford University, Oxford), 
and Dr. R. Woodson (Missouri Botanical Garden, St. Louis). Dr. 
Ove Almborn kindly forwarded to me for study loans from the National 
Herbarium, Pretoria, and from the Botanical Department, Trond- 
heim. 

In addition Drs. G. Degelius, T. Seshadri, and D. D. Awasthi 
(abbreviated DEGEL, SESH, and AWAS, respectively, in the list of 
species) have loaned us their valuable private collections. 

I have had profitable exchanges of information and unpublished 
data with Drs. H. des Abbayes and Y. Asahina. Dr. Syo Kurokawa 
has aided in the chemical testing of many collections. Dr. J. L. 
Ramaut has also carried out chemical investigations of some of the 
species and kindly communicated his results to me. 

This work has been supported in part by a grant from the National 
Science Foundation. Fieldwork in southern United States was sup- 
ported in part by grants previously received from the American 
Philosophical Society and the Society of Sigma Xi. 


Historical Survey 


Taxa properly assigned to subgenus Amphigymnia have been de- 
scribed by at least 54 different lichenologists. This effort, starting 
with Hudson in 1762, has resulted in some 350 names of various 
taxonomic rank, of which 83 are accepted in this monograph. Because 
of the scattering of efforts by so many lichenologists, detailed sum- 
maries of the contributions of each worker will not be attempted here, 
but the following overall survey may be helpful in understanding the 
development of the subgenus. 

Before Acharius proposed the genus Parmelia in 1803, only two 
species now recognized as Amphigymnias had been described, Lichen 
perforatus Jacq. and L. perlatus Huds. Between 1803 and the be- 
ginning of Nylander’s work in 1860, the most important workers were 
Sprengel, E. Fries, Eschweiler, Fée, Taylor, Hepp, and Montagne. 
During this period of early explorations about 10 new species now 
recognized as valid were described. 

From about 1860 to 1900 there was a period of intense activity by 
Nylander, Miller-Argau, Krempelhuber, Stirton, Tuckerman, Laurer, 
and others, during which more than 120 new names were created. 
Of these, about 30 are still accepted as valid. Miller and Nylander, 


196 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


the giants of this period, were complete opposites in methods and 
species concepts. Nylander perfected color tests and had a narrow 
refined species concept. Of the 30 names in Amphigymnia that he 
proposed, 15 are still valid, a very low percentage of synonymy 
indeed. Miiller, on the other hand, did not use chemistry and had a 
distressingly broad species concept. He described many varieties 
and forms, none of which are presently recognized. Of the 45 taxa 
he proposed, in fact, only six are accepted in this monograph. 
Krempelhuber made some very valuable investigations of the South 
American Parmelias which were largely overlooked by later workers. 
Stirton described a number of exotic species. Contributions by 
Tuckerman, Laurer, and others, were much smaller but nevertheless 
significant. Nearly all of the publications in this period were in the 
form of short floristic notes. 

From about 1900 to 1930 another group of lichenologists carried on 
the impetus of the previous period. The most active workers were 
Vainio, Zahlbruckner, Hue, Lynge, Du Rietz, Steiner, Harmand, 
Bouly de Lesdain, Choisy, and Cengio-Samba. Vainio on the whole 
was a careful worker, proposing 26 names, of which 12 are valid 
species. His specimens, however, are notoriously scrappy, although 
well preserved. Zahlbruckner proposed 27 names in Amphigymnia, 
of which only 6 are accepted here. Lynge wrote the most valuable 
tropical flora, based on the rich Regnell collections from Brazil (cf. 
Hale, 1960). He proposed 22 names, of which 3 are acceptable. Hue 
was a prolific worker, describing some 23 taxa from tropical areas, 6 
of them good species. Altogether, from the late 1890’s to about 
1930, somewhat over 100 new names were proposed and about 30 
of these are accepted in this monograph. The floristic lists by these 
workers tend to be more comprehensive than those published before 
1900, but fragmentary floristic notes still predominated. 

The period since 1930 has marked the beginning of a modern 
synthesis of Parmelia on a worldwide scale and full acceptance of the 
International Code of Botanical Nomenclature. Major floristic 
works have been published by des Abbayes (1958, 1961) for Mada- 
gascar and western Africa and by Dodge (1953, 1959) for eastern and 
South Africa. There have been comprehensive floristic treatments 
of the European species (Hillmann, 1934; Maas Geesteranus, 1947; 
Tavares, 1945) and of the Japanese species (Asahina, 1952). The 
North American species were studied by Berry (1941). In the ab- 
sence of a world monograph, however, many of the new species 
described since 1930 have only increased the already overburdened 
synonymy. Of nearly 100 taxa proposed in this period, only some 15 
are accepted in this monograph, 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 197 


Nomenclature 


Lichenologists have been slow to appreciate the importance of 
nomenclature. Even now that the International Code has been uni- 
versally accepted, some lichenologists are still publishing new species 
without designating holotypes and making new combinations without 
citing the basionyms. The concept of absolute priority regardless of 
rank was strong among older workers and has not yet completely died. 
This general lack of precision has resulted in endless confusion in 
floristic lists and the faithful perpetuation of serious errors in 
identification. 

All too many of the older lichenologists failed to designate holo- 
types but listed two or more syntype specimens, leaving to later 
workers the important task of selecting lectotypes. Fortunately 
this practice is no longer permissible under the Code. The type 
method is based strongly on priority, and admittedly it becomes 
difficult. to apply when holotypes are lost, unavailable for loan, or too 
fragmentary for recognition. The use of descriptions alone to charac- 
terize lichens has proved to be far less reliable than in other groups 
of cryptogams. When a type is unavailable, we have two choices, 
to ignore the name and list it as a nomen inquirendum or to select a 
neotype. 

Neotypes are definitely problematic in lichenology because original 
descriptions are often wholly inadequate and illustrations are rarely 
made. Fortunately, we have as a sound precedent the careful work 
of Santesson (1952), who proposed a number of neotypes for the 
foliicolous lichens. Neotypes at their best are indispensable sources 
for the stabilization of names. In Parmelia and many other genera 
they are particularly valuable in establishing the chemical com- 
ponents of the type of the species. At their worst, neotypes 
are merely new species in disguise. The Code does not require a 
taxonomist to create neotypes merely because untypifiable names 
exist. If creation of a neotype deliberately displaces a later typi- 
fiable name, I believe it should be made with great hesitation. It has 
been necessary in subgenus Amphigymnia to select three neotypes, 
those for P. subcrinita Nyl., P. sulphurata Nees & Flot., and P. 
perforata (Jacq.) Ach. All other species or varieties without type 
specimens are ignored for the present (see list of nomina inquirenda 
on p. 343). 

Mention should be made here on the recognition of synonyms. The 
mere listing of a name as a synonym has too often been taken as prima 
facie evidence of identity. In reality, judgment of synonymy is often 
a personal decision that parallels a lichenologist’s species concept. For 


198 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


example, Miiller-Argau had an extremely broad species concept, and 
when he made studies of the important tropical species described by 
Fée, Eschweiler, and Taylor, he reduced many names to synonymy. 
These were listed as synonyms by Zahlbruckner in his Catalogus 
Lichenum Universalis and have been so considered without further 
study by succeeding lichenologists. Actually, although many of these 
names represent valid species, they have been effectively lost in the 
literature, and progress in tropical lichen study has been held back 
because of this. 

A listing of synonymy in this monograph indicates as nearly absolute 
identity in morphology and chemistry as is possible to determine with 
the methods at hand. We have tended therefore to follow a relatively 
narrow species concept. It is a simple matter for other lichenologists 
to combine names they feel are synonyms, but at least until more 
adequate field collections are available, many of them must stand as 
distinct species. It seems probable that a much finer species concept 
will prevail in 50 years, with greater emphasis on now unrecognized or 
unappreciated morphological and chemical characters. 


Morphological Characters 


Parmelia subgenus Amphigymnia is fortunately rich in distinct and 
easily recognized taxonomic characters. However, there has been no 
general agreement as to the value and importance of these characters 
nor have lichenologists consistently reported them with equal clarity 
when describing species. Descriptions in the literature run the gamut 
from Zahlbruckner’s exhaustive, though not always pertinent, full- 
page descriptions to Gyelnik’s or Nylander’s one sentence statements. 
Asahina (1952) is one of the few lichenologists who has categorically 
listed the fundamental species characters for Parmelia. These are, in 
his opinion, cilia, soredia, isidia, spore size, pseudocyphellae and 
maculae, and chemical components. 

Neither Asahina nor other lichenologists have tried to justify their 
selection of various morphological characters. It is assumed that 
most are based on taxonomic intuition, a notoriously variable factor. 
Obviously the most easily recognized traits, cilia, soredia, and isidia, 
are first selected as species characters, and the more subtle, less 
easily recognized traits are either overlooked or given some rank 
according to each taxonomist’s judgment. It would be advantageous 
to be able to evaluate each taxonomic character quantitatively so 
that some objective measure of its importance can be arrived at. 
For example, are soredia and isidia of comparable value as species 
characters? Do cilia on the margin of the amphithecium have any 
taxonomic value? 


CONTR. NAT. HERB. VOL. 36 HALEPLATE 1 


1. Isidia of Parmelia crintta Ach. (Fink V4, US). 2. Sorediate isidia of Pe mellissti Dodve 
(Imvhaug 14235, US). 3. Isidia of P. preudotinctorwm des Abb. (des Ahbayes, US). 4. 
Coralloid isidiate outgrowths of P. ramuscuda Wale (holotype, BAL). 5. \larginal soredia of 
P. cristifera Vayl. (fale 20073, US). 6. Pustules of Po rimulora Dodge (dlinborn 1897, 
LD). All photographs enlarged © 4. 


TPO OAS He 2 


HALE PLATE 2 


CONTR. NAT. HERB. VOL 36 


7. Apotheeta of Parmelia hanntnetoniana Nall Are. (Develius. Feb. 16, 1960, LS) 1). 
S. Lower side of P. perforata (Jacq.) Nch. (Hale 21737, US) (2). 9. Lower side ot 2. 
tinclorum NV. (flale 17068, US) (* 2). 10. Maculae on the upper cortex of 2. perforata 
(Jacg.) Neh. (* 3). LL. Po vearn th ALT. Sm. showing marginal cilia Gsotype., US) ¢ 1) 
12. Lower side of Po subtinetoria Zahlbr. fale 16649. US) (2) 


CONTR. NAT. HERB. VOL. 36 HALE -PLATE 3 


13. Parmelia ebortna Wale Gsotyvpe, US). L4. PP. domintcana Vain. (Allard 15958, US). 
15. P. myelochroa Hale (holotype, US). All photographs «1, 


CONTR. NAT. HERB. VOL. 36 HALE--PLATE 4 


16. Parmelia cristifera Vayl. (Ogata s.n., US). 17. PL rubifaciens Wale (holotwpe, US). 


Both photographs 1. 


CONTR. NAT. HERB. VOL. 36 HALE-—PLATE 5 


Mat. ci. © Lo oo <a 


Examined « ; f ; Z if , Q , 
Sat, Jan-acea, /, Mall OF RV ieb 


¢ 
ane Le o. Ct. et gg oe : 
Je “ , Ba Key LAT?” 


18. Parmelia peralhida \lale (holotype, FH). 19. P. praesorediosa Nyl. (holotype, H). 


3oth photographs >< 1. 


CONTIY. NAT. HERB. VOL. 36 HALE PLATE 6 


20. Parmelta sollingert Vepp Cholotype, Ly (xsd). 21. Pe favotineta Hale Gsotype, US) 
(2), 


CONTR. NAT. HERB. VOL. 36 HALE PLATE 7 


22. Parmelia abnuens Nyl. Qectotype, HI). 23. P. maclayana Mull. Arg. Gsotype, G). 


24. P. amboimensis Dodge (Gossweetler 10008, US). All photographs l. 


CONTR. NAT. HERB. VOL 


36 


25. Parmelia merrillit Vain. 


Both photographs <1. 


(Masuda 456, UNS). 


Jf 


HALE PLATE 8 


» Po eurysaca tlue (holotype, P). 


CONTR. NAT. HERB. VOL. 36 


HALE 


PLATE 9 


30th photographs 1. 


27. Parmelia margaritata Wue (holotype, P). 


28. P, siuppea Lavi. 


(Maury 


3318, P). 


CONTR NAT. HERB. VOL 36 HALE PLATE 10 


9 Parmelta procera Stren & Za Ibr. (Greencray LOOT, IN). sO. PP r} da Theale | 


CONTR. NAT. HERB. VOL. 36 HALE PLATE 11 


31. Parmelia miranda Wale (holotype, US) (2). 32. P. flavescens (Xremplh.) Nyl. 
(holotype, M) (1). 


CONTR. NAT. HERB. VOL. 36 


HALE PLATE 12 


3. Parmelia subtinctoria Zablbr. (holotype, W). 34. P. subswanpia Nyl. (holotype, HH). 
5. P. subeaperata Kremplh.  (Glasiow 1839, TW). All photographs >< 1. 


CONTR. NAT. HERB. VOL. 36 HALE—PLATE 13 


36. Parmelia preudonilgherrensts Asah. (holotype, UNS). 37. P. diacidula Hale (holotype 
LD). 38. P. coopert Stein. & Zahlbr. (holotype, W). All photographs 1. 


CONTR. NAT. HERB. VOL. 36 HALE PLATE 14 


be t" 

% %, ¢ 
20287 
39. Parmelia reparata Stirt. (fale 19406, US). 40. P. coralliformis Hale (holotype, US). 
Both photographs ™ 1, 


CONTR. NAT. HERB. VOL. 36 HALE—PLATE 15 


41. Parmelia chiapensis Wale Gsotywpe, US). 42. P. hababtana Gyel. (lectotwpe, G). 
43. Po argentina WKremplh. Gsotype, US). All photographs 1. 


CONTR. NAT. HERB. VOL. 36 HALE PLATE 16 


44. Parmelia leucoseniotheta ue (holotype, P). 45. Pe. subrugata Kremplh. (holotype, M1). 


Both photographs <1. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 199 


PARMELIA (Amphigymnia) 00... .00.ccccccccccee cece ec eee ce esate eee 

@® 1 Publication : ec... occ cee ceeeceecseecccsee cosseseeeses cope os ceese ceeepaessoenameesesecevevee veveeees cesta. 31 ©@ 
Symomy My £2... ccc cce cee cc eee eeeceeeececee eee ecte cee etee 

@ 2 30 @ 
Thallus sa... eeececcsceee ccs ccsnsneneececeeeseercees cereeuse cnseteeeceeee 

@ 3 29 @ 
Ss) 

@ 4 28 e 
Chemistry to.....-20. cece ce cee ceeeeee eee eeeee 

CS an 27 @ 

@ 6 1—isidiate 9-—alb-ambit 17—protocetrar 25—P-C-K. % © 
2—sored-isid 10—cum fret 18—salacinic 26 —fatty 
3—sored-marg 11—gyrophoric 19—norstictic 27—-cilia K+ 

e 7 4—-sored-laml 12—lecanoric 20—stictic 28—saxicol 235 ©@ 
5—pseudocyph 13—olivetoric 21-—P+ unkn 29—- 
6—maculate 14—C+ unknown 22—usnic 30— 

@ 8 7—reticulate 15—alectoron 23—vulpin 31 4 © 
§—ciliate 16—KC+t unkn 24—sulph 

@e 9 10 1] 12 13 14 15 16 17 18 19 20 21 22 23 @ 

e °e e @ e 6 @  ] 6 e i ® e 


Ficure 1.—Marginal punch card (58 inches) used to record data for each species. 


Because we have had available data on all known species of 
Amphigymnia, we have taken this opportunity to present below not 
only descriptions of the major characters but also their frequency and 
degree of association with each other, in order to evaluate their rela- 
tive taxonomic importance. The data for each species were repro- 
duced on marginal punch cards (fig. 1) to facilitate rapid calculation 
of frequencies and association. All data were derived first from the 
type specimen and supplemented by studies of herbarium specimens 
compared with types. 


ISIDIA AND SOREDIA 


Isidia are one of the major characters useful at the species level. 
They occur in 25 of the 106 species of Amphigymnia. There is con- 
siderable variation in the structure and development of isidia. Sixteen 
species have normal cylindrical isidia, ranging from simple to more or 
less branched with a diameter of 0.04-0.10 mm. (pl. 1). These isidia 
may be short-ciliate in robust specimens of P. aberrans, P. crinita, 
P. pseudocrinita, P. subcrinita, and P. xzanthina. The isidia of P. 
pseudotinctorum are exceptionally large and thickened, up to 0.3 mm. 
in diameter (pl. 1). In P. mellissti, P. paulensis, and P. subcorallina, 
the isidia are in part granular or sorediate, but always recognizable 
as isidia (pl. 1). In P. lophogena and P. setchellit the isidia are ir- 
regularly thickened and at length pustular, without developing 
soredia. These latter structures are classified as isidia here only 
as a matter of convenience; they may represent a structure quite 
distinct from isidia. The most peculiar isidia, if indeed they are 

729-018—65——3 


200 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


isidia, are the coralloid outgrowths of P. flavotincta, P. fasciculata, 
P. coralliformis, and P. ramuscula (pl. 1). These are very large 
coralloid structures, produced laminally and marginally from small 
laciniae, which apically become irregularly thickened and at times 
pustulate or sorediate. 

Soredia occur in 34 species. Soredia are much less variable than 
isidia and usually differ only in position on the thallus. The most 
typical position in Amphigymnia is strictly marginal in linear soralia, 
characteristic of 25 of the 34 species. Well-known examples are P. 
austrosinensis, P. cristifera, and P. stuppea (pl. 1; fig. 2). In older 
thalli marginal soralia may eventually cause the lobe margins to 
become involute. Two species, P. hypomiltoides and P. natalensis, 
have marginal as well as extensive laminal soralia, with laminal 
soralia predominating. Four species have soredia which originate 
submarginally; that is, the soredia originate just back from the 
thallus margin. The soralia may be linear and, advancing on a broad 
front, soon cause the lobes to become strongly revolute (fig. 2), as 
in P. perlata and P. ochroglauca. In P. arnoldii and P. margaritata, 
the soralia are mostly orbicular and originate on short marginal 
laciniae which also become revolute (fig. 2). Parmelia bangu, P. 
cerypotozantha, P. fracta, and P. rimulosa (pl. 1) have strictly laminal, 


Figure 2a-c.—Soredial formation in Amphigymnia: a, Parmelia stuppea Tayl.; b, P. 
perlata (Huds.) Ach.; c, P. margaritata Hue. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 201 


irregular soralia which originate from cracks or pustules. These 
intergrade to a certain extent with sorediate or pustulate isidia but 
are probably best considered as a modification of soredia. 

Plants which lack soredia or isidia comprise the remaining 47 
species. These plants may have lobules or a rugose upper cortex, 
but they are always distinguishable from sorediate or isidiate plants. 

The degree of association between isidia, soredia, and lack of 
soredia and isidia (abbreviated NIS) is shown in table 1. This 
table, as an example, shows that of the 25 isidiate species, 68% have 
cilia; that of the 34 sorediate species, 77% have cilia; and that of 
the 47 species lacking soredia or isidia, 72% have cilia. Such close 
percentages may be interpreted to mean that there is little significant 
association between soredia, isidia, and lack of soredia and isidia. 


TaBLeE 1.—Percentage of isidiate, sorediate, and nonsorediate-nonisidiate (NIS) 
species with each of 16 different traits. 


Trait Isidiate Sorediate NIS 

(Number of species) (25) (34) (47) 

gyrophoric acid 12 12 4 
lecanoric acid 8 9 6 
alectoronic acid 8 21 26 
eryptochlorophaeic acid 4 3 4 
protocetraric acid 28 15 13 
salacinic acid 16 18 13 
norstictic acid 0 6 6 
stictic acid 8 3 2 
usnic acid 20 15 2 
maculae 12 27 38 
cilia 68 77 72 
white rim below 0 9 15 
perforate disc 6 30 43 
small spores 53 55 47 
intermediate spores 24 30 26 
large spores 24 15 26 


It is noteworthy that in the case of alectoronic acid, protocetraric 
acid, maculae, white rim below, and perforate apothecia, the sorediate— 
NIS species have similar percentages of association, in each case more 
or less obviously different from the isidiate species. ‘There is only one 
example, usnic acid, where isidiate and sorediate plants share a similar 
percentage of association measurably different from that of NIS 
plants and no outstanding example for isidiate-NIS. These obser- 
vations suggest that sorediate-NIS species are closely, perhaps geneti- 
cally, related and that isidiate plants belong to an entirely different 
lineage. There is good grounds then for believing that isidia are good 
species characters and that two otherwise identical plants differing 
chiefly in the presence or absence of isidia are separate species. 


202 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Further confirmation of the distinctness of isidia can be gained 
by taking the isidiate species one by one and searching among the 
nonisidiate species for “counterparts,” species which seem to be iden- 
tical in every character except for the absence of isidia. Obviously 
some degree of personal judgment is called for in this selection, but on 
the whole we have used the narrowest possible interpretation of iden- 
tity. For example, there are 2 isidiate species which have both 
sorediate and NIS counterparts. 


Isidiate Sorediate NIS 
P. endosulphurea P. araucartarum P. myelochroa 
P. subtinctoria P. subsumpta P. subcaperata 


Two species appear to have sorediate counterparts only: P. erinita- 
P. perlata, and P. erasmia—P. hypomiltoides. There is only one isidiate 
species with an apparent NIS counterpart: P. paulensis—P. melano- 
thriz. The remaining 20 isidiate species have no obvious counter- 
parts. Most species pairs that look promising are found to differ in 
spore size, perforation of apothecia, or other characters. 

If we proceed now to a similar analysis of sorediate species, we 
find many species with counterpart nonsorediate species, as follows: 


Sorediate Nonsorediate 
P. cristifera P. latissima 
P. dilatata P. zollingert 
P. austrosinensis P. andina 
P. coopert P. hololoba 
P. hypotropa P. perforata 
P. hababiana P. abessinica 
P. reparata P. leucosemotheta 
P. pseudonilgherrensis P. nilgherrensis 
P. praesorediosa P. mesotropa 
P. defecta P, soyauxit 


Combining these with previously mentioned cases, about 14 of the 
total 34 sorediate species are counterparts of nonsorediate species. 
This comparison, along with the frequency data (table 1) already 
mentioned, suggests that presence of soredia is a significantly weaker 
species character than isidia. We may be dealing simply with species 
that may or may not produce soredia or produce them only rarely 
or under special conditions. 

On the whole, however, counterpart sorediate species have a more 
northerly range (or in the Southern Hemisphere more southerly) 
than the nonsorediate species. For example, P. latissima (fig. 3) 
is strictly tropical in distribution. It is common in Cuba but does 
not occur in southern Florida, only 140 km. distant. On the other 
hand, the sorediate counterpart, P. cristifera, has a much broader 
distribution; it occurs not only in Cuba but also as far north as South 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 2038 


Ficures 3-4.—Distribution: 3, Parmelia latissima Fée; 4, P. cristifera Tayl. 


Carolina and Georgia in the United States (fig. 4). In South Africa 
P. andina is almost completely lacking south of the Tropic of Capri- 
corn (fig. 5), whereas sorediate P. austrosinensis is extremely common 
on both sides of this line (fig. 6). Furthermore, P. andina is com- 
pletely absent in North America and Mexico. Parmelia hypotropa 
(fig. 8) has a more northerly center of distribution in the United 
States than the nonsorediate counterpart P. perforata (fig. 7). 

The existence of counterpart species may have some significance 
in interpreting speciation in Amphigymnia. The ancestral species 
were probably isidiate or nonsorediate. At the present time almost 
all nonsorediate species are confined to tropical regions and many 
of them are restricted to one continent. By contrast half of the 
pantropical species are sorediate. In some of the species, e.g., P. 
latissima, perfect sorediate counterparts (P. cristifera) have de- 
veloped, and most of them are much more widely distributed in 
temperate areas than the nonsorediate counterpart. However, in 
some cases, such as P. diacidula, P. direagens, P. grayana, P. rubi- 
faciens, P. sancti-angelti, and P. subarnoldii, no nonsorediate ancestral 
forms have survived, if indeed they ever existed. Finally, these 


204 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Ficures 5-6.—Distribution: 5, Parmelia andina Mill. Arg.; 6, P. austrosinensis Zahlbr. 


various counterpart species may be slowly evolving and diverging. 
For example, P. tunctorum would be a perfect isidiate counterpart to 
P. andina and P. austrosinensis if it were maculate and had perforate 
apothecia. Parmelia eurysaca would be a logical counterpart of 
sorediate P. stuppea, except that it has distinct laciniae. Many 
other examples could be cited. 


CILIA 


Cilia are very common in Amphigymnia, occurring in 77 of the 
106 species, sporadically produced in the axils of the lobes in P. 
zollingert, P. dilatata, and P. conformata, and entirely lacking in the 
remaining 26 species. The longest cilia, such as those of P. cooperi, 
P. rampoddensis, or P. wainii, are up to 6 mm. long and very con- 
spicuous (pl. 2). At the other extreme, the cilia of P. perlata may be 
quite sparse and not more than 1 mm. long. The cilia of P. argentina 
and P. mellissii sometimes react K-+ purple, but we do not believe that 
this reaction has any taxonomic value. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 205 


Ficures 7-8.—Distribution in the United States: 7, Parmelia perforata (Jacq.) Ach.; 
8, P. hypotropa Nyl. 


The degree of association of cilia with other morphological and 
chemical characters is shown in table 2. This table shows, for ex- 
ample, that of the 77 ciliate species, 3% have lecanoric acid and of 
the 29 nonciliate species, 21% have lecanoric acid. Cilia have a 
positive or negative association with more than half of the characters 
listed. We believe that this high degree of correlation indicates that 
cilia are important taxonomic characters. This hypothesis is 
strengthened by the fact that there are almost no counterpart ciliate- 


206 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


nonciliate species. The only clear-cut examples are P. hololoba and 
P. andina. It is impossible to find exact counterparts for the other 
76 ciliate species among the 29 nonciliate species. 


TABLE 2.—Percentage of species with or without maculae, cilia, or perforate disc 
having various tratts. 


Percentage of species having 


Maculae Cilia Perforate disc 
Trait Present Absent Present Absent Present Absent 

(Number of species) (30) (76) (77) (29) (27) (56) 
gyrophoric acid 10 8 12 0 4 5 
lecanoric acid 10 7 3 21 18 4 
alectoronic acid 22 18 26 3 30 16 
cryptochlorophaeic acid 10 0 4 0 7 3 
protocetraric acid 3 21 10 31 0 23 
salacinic acid 22 13 18 10 22 14 
norstictic acid 10 3 4 14 11 2 
stictic acid 0 5 5 0 0 7 
usnic acid 0 12 12 7 0 14 
maculae — — 35 10 59 20 
cilia 90 66 — — 82 68 
white rim below 37 0 12 7 26 7 
perforate disc 59 21 37 24 —_— — 
small spores 63 45 57 35 74 39 
intermediate spores 26 27 18 43 22 28 
large spores 11 29 25 22 4 33 

MACULAE 


Maculae in Amphigymnia are more or less regular, submacroscopic 
white spots in the upper cortex (pl. 2). They are related to but should 
not be confused with the irregular effigurate pseudocyphellae in 
P. saxatilis (L.) Ach. or P. sulcata Tayl. or with typical pseudocy- 
phellae. Maculae are not always easy to distinguish in Amphigymnia 
and have usually been overlooked by lichenologists. Some 27 species 
have distinct maculae, 3 have variable or questionable maculae, and 
76 have a normal opaque cortex without maculae. Maculae occur, 
however, on the amphithecium and pedicels of nearly all species. In 
some of these the maculae may be distinct at the base of the pedicels 
but do not clearly go beyond this point on the thallus, as in P. eciliata. 

Maculae are strongly positively or negatively associated with seven 
other characters (table 2). Of these characters, cryptochlorophaeic 
acid, protocetraric acid, white rim below, and perforate apothecia 
share a similar degree of association with cilia. Maculae would then 
seem to be good species characters, especially since there are no 
recognizable maculate-nonmaculate counterpart species. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 207 


LOWER SIDE 


The lower side in Amphigymnia is more variable than usually 
thought. Typically most species are jet black, with rather long, 
unbranched, moderate to sparse rhizines. The outer rim is naked, 
shiny, dark brown or less commonly mottled tan or brown and ivory 
(pl. 2). About 91 of the species in this monograph can be classified 
in this group. 

A second group also has a black rhizinate center, but the marginal 
area is entirely pale, ivory to white, and conspicuously set off from 
the black center (pl. 2). There are at least 11 species with this so- 
called white rim: 


P. abessinica P. paulensis 
P. argentina P. perforata 
P. euneta P. rigida 

P. hababiana P,. subrugata 
P. hypotropa P. uruguensis 
P. melanothriz 


It will be noted from table 2 that a white rim is highly associated 
with maculae, perforate apothecia, and cilia, and seems to be a sound 
character. 

A third type of lower side found in three species is characterized 
by very short, rather dense rhizines produced to or nearly to the 
margin (pl. 2). The lower cortex is entirely pale brown or brown or 
blackened toward the center with age. These species (P. subcaperata, 
P. subsumpta, and P. subtinctoria) also have a strongly maculate 
cortex, cilia, and salacinic or cryptochlorophaeic acid. There are no 
counterpart species with a black lower side. The species could be 
placed in subgenus Parmelia, but because of their large broad lobes 
and lack of reticulation they are more properly treated here. 


APOTHECIAL CHARACTERS 


The typical apothecium is more or less distinctly pedicellate and 
has a rugose, maculate amphithecium. In P. hanningtoniana the 
amphithecium is very coarsely rugose to lobulate (pl. 2). The exciple 
is most frequently entire, but in P. abessinica, P. abnuens, P. argentina, 
P. corniculans, P. euneta, P. hanningtoniana, P. melanothriz, P. 
merrill, P. ornatula, P. perforata, P. rigida, P. subrugata, P. uruguen- 
sis, and P. wainii, the exciple is more or less dentate and frequently 
but not always ciliate. Cilia are not constant in any one species and 
appear to have no consistent value as a species character. The 
exciple is conspicuously laciniate in P. appendiculata, P. cornuta, and 
P. disparilis. Apothecia are not known for some 23 species, of which 
14 are sorediate, 8 are isidiate, and only 1 is without soredia or isidia. 


208 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Spores are uniformly simple and unicellular in Amphigymnia and 
the only significant variation is in size and in the thickness of the 
spore wall. There are three major size classes: (1) small thin-walled 
spores, 12-18 » long, with the episporium 1.0-1.5 » thick; (2) inter- 
mediate spores, 18-25 » long, with the episporium 1.5-2.0 yu thick; 
and (3) large spores, 25-40 » long, with the episporium 2.0-4.0 u 
thick. ‘These three size classes represent at best three clusters on a 
scale of continuous variation. However, it is not too difficult to 
assign any given species to one of these classes. The most significant 
area of intergradation is in species with intermediate or large spores, 
where P. abessinica, P. austrosinensis, P. praesorediosa, and some 
others could well be placed in either group. 

Examination of tables 1 and 2 will show that spore size is not 
consistently correlated with other morphological characters. Spore 
size is, however, strongly correlated with some of the rare acids 
(table 3). We are inclined to feel that spore size alone is not a valid 
species character, but it is undeniably a valuable key character in 
identification work, especially for species with alectoronic acid. 
There are no species in this monograph based solely on spore size, 
although there are a few problematic species where there are rather 
significant differences in spore size (see P. abnuens and P. crimta). 
As a rule, wide divergence in spore size is an indication that we are 
dealing with two species and careful study will often reveal unnoticed 
differences in other morphological characters that confirm this. 


Taste 3.—Percentage of species with large, intermediate, or small spores that 
contain each of 8 different lichen acids. 


Large Intermediate Small 


(Number of species) (19) (22) (42) 
gyrophoric acid 0 4 10 
lecanoric acid 0 0 17 
alectoronic acid 10 32 19 
protocetraric acid 21 27 5 
salacinic acid 10 0 26 
norstictic acid 0 5 7 
stictic acid 21 0 0 
usnie acid 5 14 10 


The most important apothecial character seems to be the perforate 
disc, found in at least 27 species. The discs of both small and large 
apothecia are clearly perforate, leaving a gaping hole which opens 
directly into the hollow stipe. Nonperforate species have a smooth 
continuous disc which may split radially with age but only rarely 
develops a true perforation. This character correlates rather strongly 
with a number of other characters (table 2), including protocetraric 
acid, maculae, and white rim. Perforation is apparently a valid 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 209 


species character, especially in the absence of perforate-nonperforate 
counterpart species. 

Other apothecial characters have been noted but appear to have 
little immediate value in the taxonomy of Amphigymnia. The height 
of the hymenium, for example, seems to vary in proportion to spore 
size. Small-spored species have a hymenium 35 to 65 u high, large- 
spored species 90 to 130 w high. In actual practice the height of the 
hymenium may vary 20 to 30 yu in different specimens of the same 
species. The size of the thecium is also directly proportional to spore 
size and has no particular value in descriptions. Pycnidia have not 
been consistently investigated because they seem to have no impor- 
tance in the taxonomy of Amphigymnia; they are often absent or 
extremely difficult to find. 


ANATOMICAL CHARACTERS 


Anatomical characters play no role in the taxonomy of Amphigymnia 
at this time. Dodge (1959) gives a good summary of the internal 
structure of Parmelia. The thickness of the upper cortex and lower 
cortex depends partly on the age of the thallus so that a number of 
quite different measurements can be made on the same thallus. The 
importance of variation in the structure and arrangement of the 
medullary hyphae or the organization of the algal colonies is as yet 
unknown. Such variation has been useful in interpreting Usnea 
(Asahina, 1954a), but it has not been used in foliose genera. 


Chemistry 


The use of color tests in Parmelia taxonomy started with Nylander 
in 1866, and virtually every major lichenologist to date, excepting 
Miiller-Argau and Fink, has used color tests as key characters in the 
genus. Refinements in identification of the chemicals were brought 
about by Asahina’s microchemical tests (cf. Asahina & Shibata, 1954), 
which finally placed the nonspecific color reactions on a firm founda- 
tion. No one can study Parmelia microchemically without being 
profoundly impressed by the practical importance of lichen acids in 
species identification. In addition, the presence of certain acids is 
highly correlated with morphological characters, as previously shown, 
and geography, indicating that the acids have a more basic value than 
simply their use in identification work. 

Although the chemical color tests are widely used, it is both surpris- 
ing and disconcerting to find how inaccurate or incomplete are some of 
the tests reported in the literature by many recent lichenologists. If 
color tests are used at all, it is imperative that the results be trust- 
worthy. Reports of colors with KOH and P are usually accurate, but 


210 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


tests with C or KC are often erroneous. The Ca (O Cl), reagent is all 
too often old or decomposed and fails to give any color reaction. This 
reagent should be prepared freshly each day and tested first on 
Parmelia tinctorum or P. rudecta, both of which contain lecanoric acid. 
In this connection it should be mentioned that microchemical tests 
pose even greater possibilities for error. It is hoped that a suitable 
manual for identifying lichen acids will appear in the near future and 
forestall this possibility. 


Taxonomy of Chemical Strains 


The use of chemistry in lichen taxonomy is a subject of active 
discussion by present-day lichenologists. The easy application of 
Asahina’s microchemical methods has encouraged more and more 
workers to use chemical tests. The evidence in favor of using chem- 
istry in identifying and characterizing lichens is now so overwhelming 
that few modern workers deny its use. The moot point, however, is 
whether chemical characters alone deserve taxonomic rank, and if so, 
what this rank should be. 

There are now a number of well-documented examples of lichen 
species that are morphologically inseparable but chemically distinct. 
Parmelia is particularly rich in chemical strains. Many of these 
chemical variants show more or less clear phytogeographic differences 
(cf. Culberson & Culberson, 1956; Hale, 1956a), but none has yet been 
correlated with habitat, substratum, or microclimate. In fact, all 
evidence so far suggests that chemical strains result from genetic 
control. 

The arguments in favor of species rank for chemical strains are in 
part pragmatic. Such species are given immediate recognition at 
the highest possible rank, where they will appear in keys and indexes 
without danger of being lost. As a corollary, of course, these taxa 
may be separated in taxonomic lists far from the parent species and 
the names alone give no inkling of otherwise very close affinities 
among the species. However, by using species rank, lichenologists 
are compelled to make more careful and complete identifications and 
at the same time add to the growing body of data on chemical varia- 
tion. Even though we cannot yet fully evaluate these data, the fact 
that most present-day lichenologists ascribe little taxonomic value 
to chemistry does not preclude the possibility that chemical variation 
will be of very great importance in future lichen taxonomy. Certainly 
no one can deny that the taxonomy of Cladonia has not been im- 
measurably clarified and advanced by employing chemistry to the 
fullest degree. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 211 


Another choice is an infraspecific rank. Any infraspecific taxon 
shares the disadvantages (for bibliographers in particular) of a trino- 
mial system, but at least all chemical variants would appear together 
in lists or indexes of species. Unfortunately few lichenologists have 
ever agreed on criteria for recognizing varieties or forms, and in the 
past these two taxa have been hopelessly abused. Moreover, most 
taxonomists do not routinely identify species to varietal or other sub- 
specific rank. In other words, if chemical taxa were described as 
varieties, we would in effect discourage investigation of chemical 
variation. Literature records would usually refer only to group 
names, such as Cladonia chlorophaea (Flk.) Spreng., and interested 
taxonomists would have to reexamine the specimens to determine if 
the chemical variants C. grayi Merr., C. cryptochlorophaea Asahb.., 
etc., were included. The category of subspecies has appeared sporadi- 
cally in lichen literature and was used to some extent by Runemark 
(1956) for describing chemical variation in the yellow Rhizocarpons. 
As far as I am concerned, this taxon has the same disadvantages as 
any other infraspecific rank. 

A last means of recognizing chemical variation is the extra-nomen- 
clatorial device of numbered ‘‘chemical strains” first proposed by 
Lamb (1951) and followed by several lichenologists (cf. Hale 1952). 
This category fails to provide a stable system of naming strains since 
it is not subject to rules of nomenclature. However, the use of 
chemical strain notations has a most acceptable temporary advantage 
if in a preliminary study we wish to indicate the existence of chemical 
variation without being forced to propose formal names. 

Evidence in subgenus Amphigymnia either favoring or opposing 
taxonomic recognition of chemical variation is at best inconclusive. 
The species which by any reasonable definition can be considered as 
chemical strains are listed in table 4. These include 8 parent species 
(left column) and a total of 10 chemical variants. It is surprising 
that of the 10 strains, 7 have already been described in the literature, 
in each case without any cognizance of the older parent name. 

Closer study of table 4 shows a preponderance of species with 
fatty acids or fatty acids and a color reacting acid, most often cryp- 
tochlorophaeic or salacinic acids. Of these species, P. hababiana 
(figs. 26, 27), P. subsumpta, P. abessinica, P. subcaperata, P. ranthina 
(but not including P. aberrans) (figs. 10-12), P. subtinctoria, and their 
respective chemical strains each have essentially identical distribution 
patterns and, as far as we can tell, similar habitat requirements. 
I do not believe that these variants can be considered as distinct 
species at this time. Both chemical and geographic data suggest 
variable populations of single species. 


212 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


TaBLeE 4.—Chemical strains in Amphigymnia. 


Strain 1 Strain 2 Strain 3 
P, hababiana Gyel. unnamed species (fatty) — 
(eryptochlorophaeic 
+ fatty) 
P. abessinica Kremplh. P. glaucocarpoides Zahlbr. — 
(cryptochlorophaeic (fatty) 
+fatty) 
P. subcaperata Kremplh. —P. recipienda Ny). _— 
(salacinic) (eryptochlorophaeic 
+fatty) 
P. subsumpta Nyl. unnamed species — 
(salacinic) (cryptochlorophaeic 
+ fatty) 
P. subtinctoria Zahlbr. P. haitiensis Hale unnamed species (salacinic, 
(salacinic) (cryptochlorophaeic cryptochlorophaeic, 
+ fatty) + fatty) 
P. zanthina (Mill. Arg.) P. madagascariacea (Hue) P. aberrans (Vain.) des 
Vain. des Abb. Abb. 
(fatty) (KC+unknown + fatty) (gyrophoric) 
P. perforata (Jacq.) Ach. P. rigida Lynge — 
(norstictic) (alectoronic) 
P. margaritata Hue P. arnoldit Du Rietz — 
(salacinic) (alectoronic) 


The three remaining examples show well-defined differences in 
distribution patterns. Parmelia margaritata (fig. 9) is confined to 
east-central United States, whereas P. arnoldii (figs. 9, 16) is common 
in the mountains of western and eastern North America, Mexico, 
West Indies, South America, and Europe. Parmelia zanthina (in- 


het Tee 


Ms. H : 
vs, We weeeieowel 
tase Pe 
~ 
a 


Ficure 9.—Distribution in the United States of Parmelia arnoldii Du Rietz (@) and P. 
margaritata Hue (©). 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 213 


Ficures 10-12.—Distribution: 10, Parmelia xanthina (Mill. Arg.) Vain. (protolichesteric 
acid present); 11, P. madagascariacea (Hue) des Abb. (KC+ red unknown strain of 
P. xanthina); 12, P. aberrans (Vain.) des Abb. (gyrophoric acid present). 


cluding P. madagascariacea) occurs widely in India, Africa, and 
North America (figs. 10, 11), but P. aberrans (fig. 12) is restricted 
to tropical America. Parmelia rigida occurs in southern South 
America and, with P. perforata, in the southern United States (fig. 
29). Parmelia perforata extends much farther north in the United 
States (fig. 7). On the basis of the distribution patterns of these 


214 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


species and the distinctiveness of the chemicals involved, I propose 
to recognize P. arnoldii, P. aberrans, and P. rigida as distinct species 
in addition to the earlier described chemical variants P. margaritata, 
P. zanthina, and P. perforata. 

A definitive solution to the problem of recognizing chemical varia- 
tion in lichens, if indeed such can ever be attained, must await two 
developments, a complete manual bringing up to date the identifica- 
tion of lichen acids and far more exhaustive field and herbarium 
studies of chemical strains. If chemical strains are not worthy of 
such complete investigation, then they are certainly not worthy of 
specific epithets and every effort should be made not to encumber 
the literature with hastily proposed chemical species. 


Identification of Lichen Substances 


A total of 23 different lichen substances, 15 of them definitely 
identified, has been demonstrated in the 106 species of Amphigymnia 
treated in this monograph. Almost every specimen examined was 
chemically tested. The substances were identified with routine mi- 
crochemical procedures generally used by lichenologists, including color 
tests (concentrated aqueous solution of KOH, concentrated freshly 
prepared aqueous solution of Ca (O Cl), and a 5% alcoholic solution 
of paraphenylenediamine (C,H,(NH,).); Asahina’s microchemical 
tests prepared according to Evans (1943); and chromatography with 
Whatman No. 1 paper. It must be emphasized that exhaustive 
analyses were not carried out because of limitations of equipment 
and space. Our attention was directed primarily at the substances 
of immediate use in taxonomy. 

1. Alectoronic acid. This is the commonest substance, outside of 
atranorine, occurring in subgenus Amphigymnia. It was demon- 
strated in the following 21 species: 


P. argentina P. natalensis 
P. arnoldii* P. nilgherrensis* 
P. breviciliata P. ornatula 
P. chiapensis* > : er 
. . procera 
5 Chenoa P. pseudonilgherrensis 
P. rampoddensis* 
P, erasmia P. rigida 
P. hypomiltoides P. rimulosa* 
P. maclayana P, subrugata* 
P, mellissii* P. wainit 


Alectoronic acid is always deposited in the medulla, It usually 
occurs with small quantities of e-collatolic acid, but I have usually 
not attempted to make absolute identification of a-collatolic acid. 
Species in the above list marked with asterisks may also contain the 
red-orange pigment rhodophyscin, which is scattered in moribund 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 215 


parts of the lower medulla. Alectoronic acid is identified by a bright 
fluorescence test, by the gummy residue from acetone, and by recrys- 
tallization in G.A.W. as small fan-shaped lamellae (Asahina, 1938, 
fig. 73). 

2. Atranorine. This lichen substance occurs in varying concen- 
tration in all species of subgenus Amphigymnia. It is typically pro- 
duced in the upper cortex where its presence is indicated by a distinct 
K+ pale-yellow reaction. It also probably occurs in the carbonized 
lower cortex as well, although here color tests are obscured. Only 
one species, P. flavotincta, contains atranorine in the medulla, a very 
rare occurrence in Parmelia. Atranorine is very easily identified in 
G.A.o-T. orinG.E. Because of its universal occurrence in Amphigym- 
nia species, it has little diagnostic value, and it is merely observed as 
a constant component when tests are made for other lichen acids, 
especially with G.A.o-T. 

3. Caperatic acid. This fatty acid has been positively identified 
in three species: P. diacidula (as an accessory component), P. meso- 
tropa, and P. praesorediosa. Unfortunately the identification of lichen 
fatty acids has not been completely worked out. We have recognized 
caperatic acid by the formation of dendroid or warty globules in G.E. 
(cf. Asahina 1959, fig. 2). Caperatic acid is closely related to and 
probably occurs with protolichesteric acid in other species. 

4. Cryptochlorophaeic acid. This acid was first discovered in 
Cladonia but later found in Parmelia (Hale, 1959a). Its chemical 
structure is unknown but it seems to be a depside. It has been found 
so far in six species of Amphigymnia: 


P. abessinica P. subcaperata 
P. hababiana P. subsumpta 
P. schimperi P. subtinctoria 


In all of these species it is accompanied by protolichesteric acid, 
and in P. subtinctoria salacinic acid may be present as well. Each 
of these species, excepting P. schimperi, is represented by a chemical 
strain containing protolichesteric acid alone (P. abessinica, and P. 
hababiana) or salacinic acid (P. subcaperata, P. subsumpta, and P. 
subtinctoria). The medulla of species containing cryptochlorophaeic 
acid is KC-+ deep red. A gummy residue from acetone recrystallizes 
as long curving needles in G.A.W. It should be noted that no crystals 
form in G.E. 

5. Gyrophoric acid. This acid, widely distributed in lichen genera, 
occurs in nine species of subgenus Amphigymnia: 


P. aberrans P. lophogena 
P. diacidula P. permutata 
P. euneta P. pseudocrinita*® 
P. hanningtoniana P. sancti-angelii* 


P. lobulascens 
729—018—65——_4 


216 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


In the two species marked with asterisks rhodophyscin is an acces- 
sory component. Parmelia permutata contains in addition a K— 
orange-yellow pigment, P. diacidula contains caperatic acid, and 
P. aberrans contains usnic acid. The medulla of all of these species 
reacts C+ rose and the whitish residue from acetone recrystallizes 
as small warts or granules in G.E. or G.A.W. In problems where it 
must be absolutely differentiated from olivetoric or lecanoric acids, a 
chromatographic test is necessary; gyrophoric has a very low R, value 
(Hale, 1956b). 

6. Lecanoric acid. This acid, structurally related to gyrophoric 
acid, occurs in eight species, most of them common in Africa: 


P. andina P. hololoba 

P. austrosinensis P. pseudotinctorum 
P. coopert P. soyauxit 

P. defecta P. tinctorum 


The medulla is C+ deep blood red. The thick white acetone residue 
crystallizes as needle clusters in both G.E. and G.A.W. Chromato- 
eraphic identification is very easy, lecanoric acid having an R; range 
midway between gyrophoric acid and olivetoric acid. 

7. Norstictic Acid. This is a rather rare acid in subgenus 
Amphigymnia, occurring in P. aldabrensis, P. crassescens, P. hypotropa, 
P. perforata, and P. rubifaciens. It may be intermingled with stictic 
acid in P. aldabrensis and P. hypotropa. The medullary color test 
is K+ yellow turning to red and P+ pale orange. Microchemical 
identification with G.A.o-T. is extremely simple, with the immediate 
formation of 4-angled yellow lamellae or small elliptical clusters. 

8. Olivetoric acid. The only Amphigymnia species containing 
olivetoric acid are P. abnuens and P. amboimensis. The medulla is 
C+ deep red orange. The acetone residue is gummy and recrys- 
tallizes with some difficulty in G.A.W. A chromatographic test is 
usually needed for positive identification. 

9. Protocetraric acid. This acid, with salacinic acid, is the second 
commonest acid in Amphigymnia, occurring in these 17 species: 


P. blanchetii P. merrillit 

P. conformata* P. pachyspora 

P. dilatata* P. peralbida 

P. disparilis P, saccatiloba 

P. dominicana* P. setchellit 

P. eborina P. subarnoldit 

P. fasciculata* P. subcorallina 

P. fracta* P. viridiflava* 
P. zollingers 


Species with asterisks usually also contain usnic acid, and P. 
subcorallina contains protolichesteric acid. The medullary reaction is 
K-— or faint brownish, C—, KC+ transient rose, and P+ brick red. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 217 


The acetone residue is usually small and recrystallizes as yellow balls 
from G.A.o-T. They can be mistaken for aggregates of salacinic acid 
(K-+ red) but have a deeper color and less soluble residue. 

10. Protolichesteric acid. As with caperatic acid, this fatty acid 
can only be identified with rather unsatisfactory microchemical tests 
with G.E. In this reagent thin angular or feathery plates are formed 
(cf. Asahina, 1959, fig. 4), sometimes aggregated and intergrading with 
caperatic acid. It occurs as the chief component in P. abessinica 
(ch. str.), P. grayana, P. hababiana (ch. str.), P. melanothriz, and in 
P. ochroglauca and P. xanthina accompanied with usnic acid. It also 
occurs as an accessory component in species containing cryptochloro- 
phaeic acid. In P. subcorallina it occurs with protocetraric acid, in P. 
paulensis and P. zanthina (ch. str.) with unknown KC-+ substances, 
and in P. crocoides and P. cryptoxantha with unidentified K— pigments. 
Because of the rather variable crystal shapes in G.E., we are not 
certain that the only fatty acid in these 14 species is protolichesteric 
acid or even that true protolichesteric acid is always present. It is 
hoped that future work in identification of these acids will help solve 
these problems. 

11. Rhodophyscin. This red-orange hydroxyanthraquinone pig- 
ment was first discovered in Physcia endococcinea. It occurs in ten 
species of Amphigymnia as an accessory component with alectoronic or 
gyrophoric acids (see lists above under these acids). It reacts K+ 
deep purple. It is most easily identified with chromatography, form- 
ing a distinct spot at R, .55-.65 (n-butanol-NH,OH). Rhodophyscin 
occurs near the lower cortex in thalli that seem moribund or old. It 
must be carefully distinguished from the unknown K-+ pigment No. 1 
in P. erasmia, P. hypomiltoides, and P. mesogenes (see below). 

12. Salacinic acid. This common K+ acid occurs in 17 species: 


P. coralliformis P. ramuscula 
P. cristifera P, reparata 

P. delicatula* P. stuppea 

P. eurysaca P. subcaperata 
P. flavescens* P. subcrinita 
P. latissima P. subsumpta* 
P. leucosemotheta P. subtinctoria 
P. margaritata P, uruguensis 
P. miranda* 


Species with asterisks also usually contain usnic acid; P. subcaperata, 
P. subsumpta, and P. subtinctoria have chemical strains with crypto- 
chlorophaeic acid. Salacinic acid is first recognized by a deep red 
color test with K in the medulla. A usually abundant residue from 
acetone recrystallizes as deep yellow, boat-shaped plates in G.A.o—T. 
If aggregated, they may be confused at first with norstictic acid, 
which is also K+ red. 


218 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


13. Stictic acid. Stictic acid occurs as the main component in only 
four species, P. bangii, P. erinita, P. eciliata, and P. perlata. It is an 
accessory component with norstictic acid in P. aldabrensis and P. 
hypotropa. The medulla reacts persistent K-+ yellow and unmis- 
takable colorless hexagons recrystallize in G.A.o—T. 

14. Usnic acid. This yellow cortical pigment has been demon- 
strated in 12 species as an accessory component with gyrophoric acid 
(P. aberrans), protocetraric acid (P. conformata, P. dominicana, P. 
fasciculata, P. fracta, P. viridiflava), protolichesteric acid (P. ochro- 
glauca, P. zanthina), salacinic acid (P. delicatula, P. flavescens, P. 
miranda), or a KC+ unknown (P. zanthina). It is found only rarely 
and in low concentration in P. subsumpta. Usnic acid can be tenta- 
tively identified by the yellow color of the thallus and is positively 
identified by a microchemical test with G.E. or with chromatography. 

15. Vulpinic acid. This brilliant lemon-yellow pigment has been 
found in two species of subgenus Amphigymnia: P. cornuta and P. sul- 
phurata. Microchemical identification can be made with G.E. (cf. 
Asahina, 1954b). Care should be taken not to confuse this pigment 
with other unidentified K— pigments (see below) which are pale yel- 
low orange or yellow. 

16. Unidentified hydroxyanthraquinone pigment No. 1. This con- 
spicuous red-orange pigment occurs in the medulla of P. erasmia and 
P. hypomiltoides, both of which also contain alectoronic acid, and in 
P. mesogenes, which contains unknown substances. This pigment 
forms a streak with a lower R, than rhodophyscin. It is more or less 
uniformly deposited, generally near the lower cortex, even at lobe 
tips, and is not associated with moribund parts of the medulla. 

17. Unidentified K— pigment No. 1. The following six species 
contain the same pale yellow-orange pigment, heavily concentrated in 
the medulla of apothecia and in the lower half or all of the medulla: 
P. appendiculata, P. araucariarum, P. endosulphurea, P. myelochroa, P. 
permutata, and P. subcolorata. This pigment is accompanied by gy- 
rophoric acid in P. permutata. In the remaining species the accom- 
panying components have not been identified. Barbatic acid is sus- 
pected but not yet positively identified. The K, C, and P reactions 
are vague and microchemical tests have been fruitless. 

18. Unidentified K— pigment No. 2. Two species, P. cryptoxantha 
and P. ebulliens, contain traces of a very faint unidentified yellow 
pigment. 

19. P+ Unknown. Parmelia direagens, in addition to alectoronic 
acid, contains an unidentified P+ substance in the soredia, which 
forms deep-yellow warts in G.A.o-T. 

20. K+ Unknown. Parmelia inexspectata contains an unknown 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 219 


K+, P— substance which has not been identified with any crystal 
tests. 

21. Unknown KC-+ substance No. 1. One chemical strain of P. 
rvanthina (P. madagascariacea strain) reacts KC+ red. Abundant 
colorless needles and aggregate clusters form in G.E., but they have 
not been identified. 

22. Unknown KC-+ substance No. 2. This unidentified compound 
is found in P. pawlensis. It reacts KC+ faint red; no identification 
has been made. 

23. Unknown substances. Parmelia mesogenes Nyl. contains a 
colorless substance that is unreactive with color tests. It forms large 
needle clusters in G.E. 


Geographical Distribution 


Phytogeography of lichens is actually no more than a presentation 
of known distribution patterns interpreted with reference to patterns 
already established for phanerogams and ferns. Higher plants have 
a fossil record, and there is a more or less coherent link between 
modern and fossil forms. Lichens have no fossil records and we 
know nothing of their evolution in geological ages. We will therefore 
confine our discussions here to a summary of the distribution patterns. 
These discussions must be tempered with the fact that more thorough 
collecting will undoubtedly extend the distributions of many of the 
species now considered to be endemic. 

The following 26 species are pantropical. By our definition this 
means that they have been found on at least three of the major 
continental areas, North or South America, Europe, Africa, Asia, 
and Australia, although they may be very rare in some parts of their 
ranges. 


P. andina (fig. 5) P. praesorediosa (fig. 15) 
P. austrosinensis (fig. 6) P. rampoddensis 

P. breviciliata (fig. 23) P. sanctt-angelit 

P. crinita (fig. 17) P. stuppea (fig. 21) 

P. cristifera (fig. 4) P. subarnoldit 

P. dilatata P. subcrinita 

P. endosulphurea (fig. 14) P. subrugata 

P. euneta P, subsumpta 

P. hababiana (figs. 26, 27) P. subtinctoria (fig. 24) 
P. hypotropa P. sulphurata (fig. 22) 
P, mellissii P. tinctorum 

P. perlata (fig. 20) P. xzanthina (figs. 10, 11) 
P. permutata P. zollingert 


The most interesting observation that can be made on the cosmo- 
politan species is that half of them (13) are sorediate. Only five 
species (P. andina, P. breviciliata, P. euneta, P. subrugata, and P. 
zollingeri) lack soredia and isidia. 


220 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


The following 30 species are endemic to the Americas. Many of 
these are restricted to Mexico, the West Indies, or South America, 
as may be seen in the lists of localities below. In contrast to the 
pantropical species, half (15) of the species here lack soredia and 
isidia. Only eight are sorediate. 


P. aberrans (fig. 12) P. eurysaca 
P. abnuens P. flavescens 
P. araucariarum P, flavotincta 
P. argentina P. fracta 

P. bangit P. hypomuiltoides 
P. blanchetit P. margaritata 
P. chiapensis P. mesogenes 
P. coralliformis P. mesotropa 
P. cornuta P. miranda 

P. crassescens P. myelochroa 
P. crocoides P. peralbida 
P. delicatula P, rigida 

P, eborina P. rubifaciens 
P. ebulliens P. uruguensis 
P, erasmia P, viridiflava 


Twenty-two species have been found only in Africa, mostly central 
and southern Africa. As in the Americas, about half of them lack 
soredia and isidia. 


P. amboimensis P. maclayana 

P. cooperi (fig. 25) P, natalensis 

P. cryptoxantha P. ochroglauca 

P. defecta P. ornatula 

P. diacidula P. pachyspora 

P. direagens P. pseudocrinita 
P. hanningtoniana P. pseudotinctorum 
P. hololoba P. rimulosa 

P. inexpectata P. schimperi 

P. lobulascens P. soyauxit 

P, lophogena P. subcolorata 
Seven species occur only in tropical America and Africa: 
P, abessinica P. fasciculata 

P. conformata P. leucosemotheta 
P. disparilis P, wainit 


P. eciliata (fig. 18) 


Three additional species, P. appendiculata, P. melanothriz, and 
P. paulensis, are typically American but have disjunct distributions 
in Madagascar or nearby islands. 

Only four species, P. corniculans, P. merrillii, P. nilgherrensis 
(fig. 28), and P. subcorallina, are restricted to Asia. Two species, 
P. grayana (fig. 19) and P. pseudonilgherrensis, occur in Asia and in 
Africa. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 931 


Ficures 13-15.—Distribution: 13, Parmelia dominicana Vain.; 14, Parmelia endosulphurea 
(Hillm.) Hale; 15, Parmelia praesorediosa Nyl. 


Three species, P. ramuscula, P. saccatiloba, and P. setchellu, are 
endemic to the Pacific region. On the whole, this region has a very 
poor Amphigymnia flora. 

Two species, P. reparata and P. subcaperata, have an unusual 
distribution in America and Australia-New Zealand. These may 
eventually prove to be pantropical, although their absence in Africa 
seems reasonably well established. It should be noted in passing 
that there are no Amphigymnia species endemic to Australia or New 


Zealand. 


222 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


oS, 
geil 
© 
oo 


Ficures 16-18.—Distribution: 16, Parmelia arnoldii Du Rietz; 17, Parmelia crinita Ach.; 
18, Parmelia eciliata (Nyl.) Nyl. 


The remaining species are not easily classified. These include 
P. aldabrensis (Africa, India Ocean), P. arnoldii (temperate North 
and South America, Europe), P. dominicana (Hawaii, Tropical 
America, Africa), P. latissima (Tropical America, India), P. pancheri 
(New Caledonia, Asia), P. perforata (North America, Europe, Mada- 
gascar), and P. procera (Africa, Asia, New Caledonia). 

A further breakdown of the species according to the total number 
in various geographic areas is instructive and shows parallelism with 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 223 


an? eg sg - 
ES Ps Se 


A “Ss << 9 % 
ay e e os : na > 


‘ 


Ficures 19-21.—Distribution: 19, Parmelia grayana Hue; 20, Parmelia perlata (Huds.) 
Ach.; 21, Parmelia stuppea Tayl. 


the degree of endemicity. North America north of Mexico has an 
Amphigymnia flora of 25 species, Mexico and Central America 47, 
the West Indies 27, and South America 51. Europe has only 8 species 
(P. arnoldui, P. austrosinensis, P. crinita, P. dilatata, P. hypotropa, 
P. perforata, P. perlata, and P. stuppea). Africa has a grand total of 
64 species, making it the richest area. Asia has 36 species, the Pacific 
region 10, and Australia and New Zealand 17. 

The ecology of the Amphigymnia species seems to be comparatively 


224 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Ficures 22-24.—Distribution: 22, Parmelia sulphurata Nees & Flot.; 23, Parmelia 
brevictliata Hale.; 24, Parmelia subtinctoria Zahlbr. (specimens with salacinic acid). 


simple. Almost all of them flourish best in open rather dry deciduous 
forests or savannas at low to medium elevation. In dense deciduous 
forests the typical habitat is the canopy branches. In open savannas 
or pastures or on shade trees in coffee or banana plantations the lower 
trunks are thickly covered with Amphigymnias. At sea level the 
most commonly collected species are the pantropical weeds such as 
P. praesorediosa and P. tinctorum. The richest development of species, 
at least in the tropics, appears to be at elevations between 500 and 
2000 m. Few species are found above 3000 m. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 225 


‘co 


Ficures 25-27.—Distribution: 25, Parmelia cooperi Stein. & Zahlbr.; 26-27, Parmelia 
hababiana Gyel.: 26, protolichesteric acid strain (=P. glaucocarpoides Zahlbr.); 27, 
cryptochlorophaeic acid strain). 


Subgeneric Classification 


The number of previously described subgeneric taxa in Amphigym- 
nia is small and most of them, in fact, have been proposed by only 
two lichenologists, Vainio and Gyelnik. The typification of these 
few names, however, has been extremely difficult, mainly because 
Vainio’s classification, used by almost all lichenologists, is based on 


296 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


color of the thallus (presence or absence of usnic acid), a character 
which we have found to be of secondary importance. Because of this, 
it might be well to review first the important morphological characters 
that we have determined in subgenus Amphigymnia, especially with 
respect to their possible use in setting up a subgeneric classification. 
These data may be summarized as follows: 

1. Isidia, soredia, and the lack of isidia and soredia in general show 
very weak association with other morphological and chemical 
characters. ‘There was rather significant correlation between soredi- 
ate-NIS species but not enough to justify splitting them from the 
isidiate species. 

2. Maculae, cilia, and perforate apothecia all show a high degree of 
association with each other and with white rim below. 

3. Spore size is not consistently correlated with any characters 
outside of some of the rarer acids. 

4. The presence of usnic acid is a rather rare trait negatively 
associated with maculae and perforate apothecia. 

After considering these data, we must conclude that there is no one 
dominating character that will immediately divide Amphigymnia 
into natural sections. The primary characters are cilia, maculae, 
perforate apothecia, and white rim below, all of which are mutually 
associated and show strong association with some of the lichen acids. 
Of these four characters, perforate apothecia are least usable because 
nearly 25% of the species are sterile. Maculae are useful but are 
difficult to recognize without considerable experience. White rim 
below can also be troublesome to distinguish and is a relatively rare 
character. This leaves cilia as the most likely and certainly the most 
easily recognized character for a major sectional division. 

If we proceed now to a study of subgeneric taxa proposed by past 
workers, we find only one basic system, which was first proposed by 
Vainio (1890) as the result of his experience with the lichen flora of 
Rio de Janeiro. He divided Parmelia into three sections: Amphi- 
gymnia, Hypotrachyna (=Parmelia), and Xanthoparmelia. These 
sections, now recognized as subgenera, were carefully delimited for 
the most part and have been correctly interpreted by most lichenolo- 
gists. In this monograph our attention will be directed only at 
subgenus Amphigymnia. 

Within section Amphigymnia, Vainio erected two groups without 
formal rank, based entirely on color of the thallus: Subglaucescentes, 
gray species (lacking usnic acid) and Subflavescentes, the yellow-green 
species (usnic acid present). As far as we know, no one has ever 
attempted to typify Vainio’s subgeneric taxa. It seems reasonable 
to assume that he considered Subglaucescentes to be the typical group 
in Amphigymnia since it is the largest and is presented first. The 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 227 


Subflavescentes by contrast is a smaller group of only three species. 
The selection of a lectotype for Subflavescentes is not difficult since 
the species listed here by Vainio all contain usnic acid and are ciliate. 
Parmelia delicatula seems to be the most suitable type species and 
has also been mentioned as such by Gyelnik (1932, p. 225). 

The eight species assigned by Vainio to Subglaucescentes form a 
motley group, some ciliate, some nonciliate. Selection of a type must 
unfortunately be arbitrary. Since the lectotype of Subflavescentes is 
ciliate, it would seem desirable to select a nonciliate species for Sub- 
glaucescentes, especially in view of our reliance on cilia as a primary 
character. The best known nonciliate species listed is P. tinctorum 
Nyl. (identified by Vainio as P. coralloidea (Mey. & Flot.) Vain.), a 
common pantropical species. As will be seen, Gyelnik also defined 
Subglaucescentes as including typically nonciliate species as well as 
ciliate ones. 

The second major subgeneric classification for the Amphigymnias 
was proposed by Gyelnik in 1932. It is essentially a modification of 
Vainio’s. Gyelnik did not recognize section Amphigymnia but 
adopted in its place two sections, Subflavescentes (Vain.) Gyel. and 
Subglaucescentes (Vain.) Gyel., and created within each section 
parallel nonciliate—ciliate subsections as follows: 

Section Subflavescentes (Vain.) Gyel. 

Subsection Eciliatae Gyel. (lectotype P. caperata) 
Subsection Ciliatae Gyel. (type P. delicatula) 

Section Subglaucescentes (Vain.) Gyel. 

Subsection Eciliolae Gyel. (lectotype P. perlata) 
Subsection Ornaticolae Gyel. (lectotype P. perforata) 

Gyelnik made several errors which make typification rather 
arbitrary. Of three species listed for subsection Eciliatae, one (P. 
kernstockii Lynge & Zahlbr.=P. flaventior Stirt.) is pseudocyphellate 
and two (P. soredians Nyl. and P. caperata (L.) Ach.) belong in 
subgenus Parmelia. Parmelia perlata, the type of subsection Ecilio- 
lae, is actually ciliate and it is possible that Gyelnik did not interpret 
this species correctly. 

Our studies correlating morphology and chemistry in Amphigymnia 
have convinced us that thallus color is useless as a sectional character. 
If our assessment of the importance of cilia is correct, Gyelnik has 
come closest to a natural subdivision of subgenus Amphigymma, 
although it is complicated by his acceptance of Vainio’s two color 
based groups. By recombining taxa described by Vainio and Gyelnik, 
we are proposing the following classification. Typification and dis- 
cussions of each taxon will be presented below in the list of species. 


Subgenus Amphigymnia (Vain.) Dodge 
1. Section Amphigymnia (Section Subglaucescentes (Vain.) Gyel.) 


228 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


2. Section Subflavescentes (Vain.) Gyel. 
Subsection Subflavescentes 
Series Subflavescentes 
Series Emaculatae Hale 
Subsection Ornaticolae Gyel. 
Series Subpallidae Hale 
Series Ornaticolae 


Key to the Subgeneric Taxa 


1. Margins and apices of lobes without cilia or axils of lobes rarely sparsely 


ciliate. . 2... 1 ee ee ee ee ee 6C1 Section Amphigymnia 

1. Margins and apices of lobes distinctly ciliate . . 2. Section Subflavescentes 
2. Upper cortex without distinct maculae. . . . Subsection Subflavescentes 
3. Thallus yellow (usnic acid). . ..... . . . Series Subflavescentes 

3. Thallus mineral gray (usnic acid lacking). . . . . . Series Emaculatae 

2. Upper cortex distinctly maculate . . . . . . . . Subsection Ornaticolae 


4. Lower side uniformly pale brown, short rhizinate to the margins. 
Series Subpallidae 
4. Lower side black and rhizinate in the center, brown to white or mottled 
and naked along the margins. . ..... . . . Series Ornaticolae 


Artificial Key to Species 


1. Thallus isidiate, sorediate-isidiate, or pustulate ............ C0 
1. Thallus lacking isidia. 
2. Thallus sorediate or pustulate . ......... =... . II (p. 230). 
2. Thallus lacking soredia or pustules. . . ....... . . . ITI (p. 232). 


I. THALLUS ISIDIATE, SOREDIATE-ISIDIATE, OR PUSTULATE 


1. Margins of lobes smooth, without cilia. 
2. Medulla pale yellow orange. ........ ... 13. P. endosulphurea 
2. Medulla white. 
3. Thallus yellowish green, usnic acid present. 
4. Isidia normal, to 1 mm. high, laminal. . . . . . 31. P. conformata 
4. Isidia coralloid, to 5 mm. high, mostly marginal. . 14. P. fasciculata 
3. Thallus mineral gray to buff, usnic acid lacking. 
5. Isidia coralloid, to 5 mm. high, mostly marginal. 
6. Medulla K+ red (salacinic acid). ..... . .23. P. ramuscula 
6. Medulla K—, P+ red (protocetraric acid) . . . 14. P. fasciculata 
5. Isidia laminal, to 1 mm. high, not conspicuously coralloid. 
7. Isidia very thick, 0.2-0.3 mm. in diameter. 
8. Medulla C+ red (lecanoric acid) ; Africa. 22. P. pseudotinctorum 
8. Medulla C—, P+ red (protocetraric acid) ; Pacific region. 
26. P. setchellii 
7. Isidia thin, usually less than 0.1 mm. thick. 
9. Medulla C+ red (lecanoric acid). . . . .. . 28. P. tinctorum 
9. Medulla C—, P+ red (protocetraric acid). 
10. Thallus coriaceous; spores large; Pacific region. 
25. P. saccatiloba 
10. Thallus membranous; spores small; tropical America. 
20. P. peralbida 
1. Margins of lobes distinctly ciliate. 
11. Medulla deep lemon yellow, K—. ....... .. . 78 P. sulphurata 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 229 


11. Medulla white or pigmented, pigment K+ purple. 
12, Thallus yellowish green, usnic acid present. 
13. Lower side pale below, short rhizinate or papillate to the margin. 
82. P. subtinctoria 
13. Lower side jet black, the margins naked. 
14. Medulla K+ red or brown. 


15. Plants saxicolous; salacinic acid present. . . 33. P. flavescens 
15. Plants corticolous; protocetraric and fumarprotocetraric acid 
present .........2.4..... 81. P. conformata 


14. Medulla K—. 
16. Medulla P+ red (protocetraric and fumarprotocetraric acid). 
31. P. conformata 
16. Medulla P—. 
17. Medulla C+ rose (gyrophoric acid). . . . . 30. P. aberrans 
17. Medulla C—, KC— or KC+ rose. . ... 37. P. xanthina 
12. Thallus mineral or ashy gray to buff, usnic acid lacking. 
18. Isidia coralloid, marginal, 1-5 mm. high. 


19. Cortex heavily white maculate. . . .. . . 88. P. coralliformis 
19. Cortex emaculate. 
20. Medulla K+ yellow (atranorine). .... . 54. P. flavotincta 
20. Medulla K— (alectoronic acid). . . ... . . 63. P. mellissii 


18. Isidia simple to moderately branched, to 1 mm. high. 
21. Isidia thick, pustular; medulla C+ rose (gyrophoric acid). 
60. P. lophogena 
21. Isidia thin, not pustulate (but sometimes sorediate) ; medulla C—. 
22. Isidia in part granular or sorediate. 
23. Medulla P+ red (protocetraric acid) . . 76. P. subcorallina 


23. Medulla P—. 
24, Alectoronic acid present . .... - . . . 63. P. mellissii 
24. Protolichesteric acid present ...... 98. P. paulensis 


22. Isidia normal, cylindrical. 
25. Medulla in part orange red, pigment K+ purple. 
26. Medulla C+ rose (gyrophoric acid) ; Africa. 
70. P. pseudocrinita 
26. Medulla C— (alectoronic acid) ; Central America and Mexico. 
52. P. erasmia 
25. Medulla entirely white. 
27. Lower side uniformly brown, short rhizinate to the margin. 
82. P. subtinctoria 
27. Lower side jet black, the margins more or less naked. 
28. Medulla C+ rose (gyrophoric acid); Africa. 
70. P. pseudocrinita 
28. Medulla C— ; cosmopolitan species. 
29. Medulla K+ persistent yellow (stictic acid). 
47. P. crinita 
29. Medulla K+ red (salacinic acid). 
30. Cortex strongly maculate; lower side rhizinate or papil- 
late to margin, pale brown in a broad zone. 
82. P. subtinctoria 
30. Cortex emaculate or faintly maculate; lower side naked 
and shiny dark brown along the margins. 
77. P. subcrinita 


230 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


II, THALLUS SOREDIATE OR PUSTULATE 


1. Margins of lobes smooth, without cilia. 
2. Medulla pale yellow orange... ..- s+ ees . 2. P. araucariarum 
2. Medulla white. 
3. Medulla K+ red. 
4, Soredia farinose, soralia elongate; thallus loosely attached; salacinic acid 


present 26 6 6 ee ee ee ee ee te 5. P. cristifera 
4. Soredia more or less conglutinated, soralia irregular; thallus adnate; 
norstictic acid present . . ........ =. . 2& P. rubifaciens 


3. Medulla K— or K+ brownish. 
5. Medulla P+ red (protocetraric acid). 
6. Soredia farinose, yellowish, usnic acid abundant . 10. P. dominicana 
6. Soredia granular, white, usnic acid usually absent . . 8. P. dilatata 
5. Medulla P—. 
7. Medulla C—, KC— (caperatic acid). . . . . 21. P. praesorediosa 
7. Medulla C+, KC-+ red (lecanoric acid). 
8. Plants saxicolous, dark below at the margin. . . . 7. P. defecta 
8. Plants corticolous, often pale or white rimmed below. 
3. P. austrosinensis 
1. Margins of lobes ciliate. 
9, Thallus yellowish green, usnic acid present. 
10. Lower side pale, short rhizinate to the margin . . . 81. P. subsumpta 
10. Lower side black, naked at the margin. 
11. Soredia laminal in extensive pustules ...... . . 55. P. fracta 
11. Soredia marginal or submarginal in linear soralia. 
12. Medulla K+ red (salacinic acid); Mexico. . . . 34. P. miranda 
12. Medulla K— or K+ brown. 
13. Medulla P— (protolichesteric acid); Africa . 35. P. ochroglauca 
13. Medulla P+ red (protocetraric acid); tropical America. 
36. P. viridiflava 
9. Thallus mineral or whitish gray to buff, usnic acid absent. 
14. Soredia chiefly laminal, often originating from extensive pustular 
eruptions. 
15. Soralia orbicular; thallus coriaceous, upper cortex maculate; Africa. 
96. P. natalensis 
15. Soralia originating from irregular cracks and pustules; thallus mem- 
branous, cortex emaculate. 
16. Medulla K—, P— (alectoronic acid); South Africa . 72. P. rimulosa 
16. Medulla K+ yellow or red, P+ yellowish. 
17. Stictic acid present; South America . ... . . 42. P. bangii 
17. Stictic acid absent; Africa... .. . . 48. P. cryptoxantha 
14. Soredia chiefly marginal, pustules lacking. 
18. Medulla in part with red or yellow-orange pigments. 
19. Pigments K—; medulla C+ rose (gyrophoric acid). 
68. P. permutata 
19. Pigments K+ purple. 
20. Medulla C+ rose (gyrophoric acid) . . . 73. P. sancti-angelii 
20. Medulla C—, KC-+ red (alectoronic acid). 
21. Pigment conspicuous throughout the medulla; soredia sub- 
marginal and subisidiate . . . . . . 58. P. hypomiltoides 
21. Pigment scattered near the lower cortex; soredia farinose. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 231 


22. Soredia originating submarginally; sorediate lobes revolute. 
41. P. arnoldii 
22. Soredia terminal; sorediate lobes involute. 
71. P. rampoddensis 
18. Medulla white, pigments absent. 
23. Medulla K+ yellow or yellow turning red. 
24. Cortex distinctly maculate. 
25. Lower side usually pale, rhizinate or papillate to the margin. 
81. P. subsumpta 
25. Lower side black, brown or white and naked along the margins. 
26. Lower side with a white marginal rim; norstictic or stictic 
acid present. . ... » + + « « 92. P. hypotropa 
26. Lower side brown along the | margins; salacinic acid present. 
93. P. leucosemotheta 
24. Cortex dull, emaculate. 


27. Soralia linear, terminal. . . ...... . . 74. P. stuppea 

27, Soralia orbicular to sublinear, submarginal, the sorediate lobes 
revolute. 

28. Soralia chiefly on lobe margins, mostly linear; stictie acid 

present. .... ... . 67. P. perlata 

28. Soralia chiefly on marginal lobules or laciniae, mostly orbicu- 

lar; salacinic acid present. . . . . 62. P. margaritata 


23. Medulla K—. 
29. Medulla P+ red. 
30. Only soredia reacting P+ orange red; South Africa. 
50. P. direagens 
30. Soredia and medulla both reacting P+ orange red; pantropical. 
31. Cilia produced densely around lobe tips; soralia linear. 
75. P. subarnoldii 
31. Cilia sparse or absent at lobe tips; soralia irregular. 
8. P. dilatata 
29. Medulla P—. 
32. Medulla C+ rose or red. 
33. Cortex distinctly maculate. 
34. Medulla C+ rose (gyrophoric acid). . 94. P. lobulascens 


34. Medulla C+ red (lecanoric acid) . . . . 87. P. cooperi 
33. Cortex emaculate or in part faintly maculate. 
35. Medulla C+ red (lecanorie acid). . . . 87. P. cooperi 


35. Medulla C+ rose (gyrophorie acid). 
36. Soralia farinose, linear; pantropical. 
73. P. sancti-angelii 
36. Soralia coarse, irregular; caperatic acid present; South 
Africa»... . 2 1... 49 P. diacidula 
32. Medulla C—. 
37. Lower side rhizinate or papillate to the margin, pale in a broad 
zone .... . . .. . . 81, P. subsumpta 
37. Lower side with a . broad naked zone, jet black at the center, 
pale or dark brown at the margin. 
38. Cortex maculate. 
39. Lower side with a broad ivory zone. . 90. P. hababiana 
39. Lower side dark brown at the margins. 
100. P. pseudonilgherrensis 


729-018—65——5 


232 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


38. Cortex emaculate; lower side dark at the margin. 

40. Medulla KC— (protolichesteric acid); Africa and Asia. 
56. P. grayana 

40. Medulla KC+ red (alectoronic acid); pantropical. 
41. Soralia in part submarginal; sorediate lobes revolute. 
41. P. arnoldii 

41. Soralia terminal; sorediate lobes involute. 

71. P. rampoddensis 


Ill. THALLUS LACKING SOREDIA OR PUSTULES 


1. Margins of lobes lacking cilia. 
2. Medulla pigmented orange red or yellow. 
3. Pigment K+ purple. ........4..... 16. P. mesogenes 
3. Pigment K—. 
4. Pigment pale yellow; acetone residue bubbling in G.E. 
12. P. ebulliens 
4. Pigment yellow orange; acetone residue not bubbling. 
5, Protolichesteric acid present . ....... . . .6. P. crocoides 
5. Fatty acids lacking. . .......... . 18. P. myelochroa 
2. Medulla white. 
6. Medulla K+ red. 
7. Spores 28-32 y; salacinic acid present... . . . . 15. P. latissima 
7. Spores 20-25 yu; norstictic acid present . . . . . . .4. P. crassescens 
6. Medulla K— or K+ brownish. 
8. Medulla P+ orange red. 


9. Plants saxicolous; spores 11-16 ulong. ..... . Il. P. eborina 
9. Plants corticolous; spores 16-22 y long. 
10. Cortex maculate; margins laciniate . . . . . . 9. P. disparilis 


10. Cortex emaculate; margins smooth, rarely laciniate. 
29. P. zollingeri 


8. Medulla P—. 
11. Medulla C+ red (lecanoric acid). 
12. Plants saxicolous; Africa. . ........ . 27. P. soyauxii 
12. Plants corticolous; pantropical... ..... .. IL. P.andina 
11. Medulla C—. 
13. Medulla KC+ red (alectoronic acid). . . . . 19. P. pancheri 


13. Medulla KC—. 
14. Spores 26-28 »; unknown substance effervescent in G.E. 
12. P. ebulliens 
14. Spores 16-21 yu; caperatic acid present . . . 17. P. mesotropa 
1. Margins of lobes ciliate. 
15. Thallus yellowish green, usnic acid present; South America. 
32. P. delicatula 
15. Thallus whitish or mineral gray to buff, usnic acid lacking. 
16. Medulla, at least in part, pigmented yellow or orange yellow. 
17. Pigment K—, present in most of the medulla. 
18. Exciple of apothecia entire, disc perforate; Africa. 
104. P. subcolorata 
18. Exciple dentate or laciniate disc imperforate. 
19. Vulpinic acid present. . . .. . . . 46. P. cornuta 
19. Unknown orange-yellow pigments present . 40. P. appendiculata 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 233 


17. Pigment K+ purple, scattered near the lower cortex. 
20. Spores 18-24 uw; cortex more or less distinctly maculate. 
21. Thallus coriaceous, strongly maculate; lobes adnate; Asia. 
97. P. nilgherrensis 
21. Thallus membranous, weakly maculate; lobes suberect; tropical 


America»... 2... 2... . 86. P. chiapensis 

20. Spores 28-34 yw; cortex not distinctly maculate. 
22. Apothecia imperforate;pantropical. . . . . .105. P. subrugata 
22. Apothecia perforate; Asia... ... . . 45. P. corniculans 


16. Medulla white, pigments lacking. 
23. Apothecia present. 
24. Apothecial disc imperforate. 
25. Medulla K+ yellow or red, P+ yellow. 
26. Stictic acid present; tropical America and South Africa. 
51. P. eciliata 
26. Unknown acid present; Africa ..... 59. P. inexspectata 
25. Medulla K—. 
27. Medulla P+ red (protocetraric acid). 
28. Spores large, 26-34 u, the episporium 2-4 yz thick. 
29. Lobe margins and frequently apothecia laciniate; Asia. 
64. P. merrillii 
29. Lobes and apothecia entire; Africa. . 66. P. pachyspora 
28. Spores small or intermediate, to 22 » long, the episporium 
less than 2 p thick. 
30. Spores small, 10-15 » long; thallus saxicolous. 
43. P. blanchetii 
30. Spores intermediate, 18-22 » long; thallus corticolous. 
29. P. zollingeri 
27. Medulla P—. 
31. Medulla C+ rose or red. 
32. Spores large, 22-34 » long; South America. 38. P. abnuens 
32. Spores small, 10-14 » long; Africa . 39. P. amboimensis 
31. Medulla C—. 
33. Spores large, 24-34 u long. 
34. Medulla KC— (protolichesteric acid). 
95. P. melanothrix 
34. Medulla KC+ (alectoronic acid). 
35. Lower side pale at the margin; exciple dentate; pan- 
tropical. . ......2..- 105. P. subrugata 
35. Lower side dark brown, exciple entire; Asia. 
97. P. nilgherrensis 
33. Spores small or intermediate, to 22 » long. 
36. Cortex maculate. 
37. Exciple ciliate; white rim below; alectoronic acid 
present; tropical America. . . . 85. P. argentina 
37. Exciple entire; margin dark brown below; crypto- 
chlorophaeic acid present; Africa. 103. P. schimperi 
36. Cortex emaculate. 
38. Plants saxicolous. ....... 44. P. breviciliata 
38. Plants corticolous. 


234 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


39. Cilia dense; spores 18-22 plong .. 79. P. wainii 
39. Cilia sparse; spores 12-16 plong . . 69. P. procera 
24. Apothecia perforate. 
40. Spores large, 28-34 w long. ..... . 45. P. corniculans 
40. Spores small or intermediate, to 22 » long. 
41. Medulla K+ red. 
42. Cortex dull, emaculate. . ..... . . 53. P. eurysaca 
42. Cortex shiny, maculate. 
43. Lower side with black center and white rim around the 
margin . wee ee ee 99. P. perforata 
43. Lower side with black or pale brown center and brown 
margin. 
44. Lower side uniformly brown or pale brown. 
45. Lower side rhizinate to the margin. 
80. P. subcaperata 
45. Lower side naked at the margin . 84. P. aldabrensis 
44. Lower side black at the center. 
46. Lower side rugose, short rhizinate to the margin; 


southern South America . . . 106. P. uruguensis 
46, Lower side smooth, naked at the margin; tropical 
America and Australia. . .. . 101. P. reparata 


41. Medulla K—. 
47, Medulla C+ rose or red. 
48. Cortex maculate; medulla C+ rose (gyrophoric acid). 
89. P. euneta 
48. Cortex dull, emaculate; medulla C+ red (lecanoric acid). 
57. P. hololoba 
47. Medulla C—. 
49. Lower side uniformly pale brown, rhizinate to the margin. 
80. P. subcaperata 
49. Lower side black at the center, paler at the margins. 
50. Cortex dull, emaculate; lower side brown at the margins. 
51. Plants saxicolous; exciple ciliate . . 65. P. ornatula 
51. Plants corticolous; exciple eciliate. 
61. P. maclayana 
50. Cortex shiny, maculate; lower side usually pale ivory to 
white, brown, or mottled below at the margins. 
52. Amphithecium coarsely rugose-lobulate. 
91. P. hanningtoniana 
52. Amphithecium smooth or rugose. 
53. MedullaKC— ...... . . 83. P. abessinica 
53. Medulla KC+ red. 
54. Cryptochlorophaeic acid present. 
83. P. abessinica 
54. Alectoronic acid present. 
55. Lower side with a broad white margin. 
102. P. rigida 
55° Lower side mottled or brown at the margin. 
56. Thallus coriaceous; cilia coarse; Asia. 
97. P. nilgherrensis 
56. Thallus membranous; cilia fine; tropical 
America. .... . . 86. P. chiapensis 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 235 


23. Apothecia not present. 
57. Medulla K—. P-. 
58. Medulla C+ red or rose. 


59. Lobes sublinear, laciniate, emaculate . . 39. P. amboimensis 
59. Lobes rotund, maculate ......... . 89. P. euneta 
58. Medulla C—. 
60. Cortex strongly white-maculate . . . . . 80. P. subcaperata 
60. Cortex dull, lacking white maculae. 
61. Thallus corticolous, laciniate . ... . 45. P. corniculans 
61. Thallus saxicolous, without laciniae . . 44. P. breviciliata 


57. Medulla P+ and/or K+ yellow to red. 
62. Cortex strongly white-maculate. 
63. White rim below or brown and naked. . . 99. P. perforata 
63. Brown below with fine rhizines to the margin. 
80. P. subcaperata 
62. Cortex dull, emaculate. 
64. Medulla K+ distinct yellow or red. 
65. Lobes with distinct marginal laciniae; salacinic acid present. 
53. P. eurysaca 
65. Lobes with few laciniae; salacinic acid lacking. 
66. Stictic acid present; tropical America and South Africa. 
51. P. eciliata 
66. Unknown substance present; Africa . 59. P. inexspectata 
64. Medulla K— or K+ faint brownish yellow (protocetraric acid 


present). 
67. Thallus saxicolous; lobes distinctly ciliate . 43. P. blanchetii 
67. Thallus corticolous; cilia rare. ..... 29. P. zollingeri 


Subgenus Amphigymnia 


Section Amphigymnia Vain. (1890, p. 28). 
Parmotrema Mass. Atti I. R. Ist. Veneto, III, 5: 248. 1860. Lectotype: P. 
perforata (Jacq.) Ach. 

Type species: Parmelia tinctorum Nyl. 

Thallus adnate to loosely attached, 5-30 cm. in diameter; lobes 
broad, rotund, 5-20 mm. wide; lower side black or brown, rhizinate 
in the center, the rhizines simple, often coarse, the margins usually with 
a broad naked zone; apothecia more or less pedicellate, the disc often 
perforate. 


1. Section Amphigymnia 


Section Amphigymnia * Subglaucescentes Vain. (1890, p. 28, as “Subglauce- 
scens’’). 

Section Subglaucescentes (Vain.) Gyel. (1932, p. 225). 

Section Subglaucescentes subsection Eciliolae Gyel. (1932, p. 225). 

Section Amphigymnia subsection Subglaucescentes (Vain.) Hillm. (1934, p. 
243). Superfluous combination. 


Thallus adnate to loosely attached; lobes broad, rotund, 5-20 mm. 
wide, the margins eciliate or rarely with sparse cilia in the axils; 
lower side black and rhizinate, the marginal zone brown, ivory to 
white, or mottled and naked. 


236 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


The 29 species in section Amphigymnia seem to form a natural 
grouping. Most have a brown rather than white marginal rim below 
and all except P. andina, P. austrosinensis, and P. disparilis lack 
maculae. Five of the eight species in the subgenus with lecanoric 
acid are found in this section; only one species (P. pancheri) contains 
alectoronic acid. Eleven of the species are endemic to the Americas, 
five to Asia, and only three to Africa. These facts seem to indicate 
that section Amphigymnia is the most primitive group in the subgenus. 


1. Parmelia andina Mill. Arg. Rev. Mycol. 1: 169. 1879. 

Parmelia abessinica var. nuda Mill. Arg. Flora 62:289. 1879. Type collec- 
tion: Seriba Ghattas, Djur, Africa, Schweinfurth (G, lectotype; M, US, W, 
isotypes). 

P. africana Mill. Arg. Flora 63:265. 1880. Based on P. abessinica var. 
nuda Mill. Arg. 

P. hildebrandtii Kremplh. f. nuda Mill. Arg. Flora 74:376. 1891. Type 
collection: South Africa, Hannington (G, lectotype). 

P. zambesica Mill. Arg. Verhl. Zool. Bot. Gesell. Wien 53:296. 1893. Type 
collection: Boroma, Africa, Menyhart s.n. (G, lectotype). 

P. pedicellata Stein. Sitzungsber. Akad. Wiss. Wien 106:214. 1897. Type 
collection: Matchakos, Kenya, Liechtenstein, 1896 (WU, holotype). 

P. olivetorum var. esorediata Vain. in Welw. Cat. Afr. Pl. 2:399. 1901. 
Type collection: Serra de Mongollo, Welwitsch 110 (TUR, lectotype; BM, 
isotype). 

P. olivetorum var. hyporysalea Vain. in Welw. Cat. Afr. Pl. 2:399. 1901. 
Type collection: Morro de Lopolla, Angola, Welwitsch 27 (TUR, lectotype; 
BM, isotype). 

P. menyhartit Stein. Verh. Zool. Bot. Gesell. Wien 53:235. 1903. Type 
collection: Zambesi, Africa, Menyhart(?) 475 pr. p. (WU, holotype). 

P. paraguariensis Lynge, Ark. Bot. 13, no. 13:71, pl. 1, fig. 7. 1914. Type 
collection: Cierro Negro, Paraguari, Paraguay, Malme 1539 (S, holotype). 

P. rissoensis Lynge, Ark. Bot. 13, no. 13:69. 1914. Type collection: Near 
Rfo Apa, Colonia Risso, Paraguay, Malme 1895B (8, holotype). 

P. hyporysalea (Vain.) Vain., Bot. Mag. Tokyo 35:47. 1921. 

P. hyporysalea (Vainio) Stein. & Zahlbr. Bot. Jahrb. Engler 60:533. 1926. 
Superfluous combination. 

P. hyporysalea var. menyhartit (Stein.) Stein. & Zahlbr. Bot. Jahrb. Engler 
60:535. 1926. 

P. pedicellata var. subbullata Stein. & Zahlbr. Bot. Jahrb. Engler 60:536. 
1926. Type collection: Bukoba, Lake Victoria, Africa, Schréder 323 (W, 
holotype). 

P. dalei Dodge, Ann. Mo. Bot. Gard. 46:131. 1959. Type collection: 
Toro, Fort Portal, Uganda, Dale L44 (K, holotype). 

P. subbullata (Stein. & Zahlbr.) Dodge, Ann. Mo. Bot. Gard. 46:162. 1959. 

P. thomasit Dodge, Ann. Mo. Bot. Gard. 46:168. 1959. Type collection: 
Mt. Otse, West Nile, Uganda, Thomas 1962 (K, holotype). 


Type collection: Cisne, Ecuador, André 4324 bis (G, holotype; K, 
isotype). 

Thallus often very large, to 30 cm. broad, loosely attached to bark, 
light mineral gray; lobes large and rotund, 10-20 mm. wide, margins 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 237 


smooth to broadly crenate, cilia lacking; upper surface plane to 
broadly undulous, continuous or reticulately cracked with age, 
faintly to distinctly white-maculate, isidia and soredia lacking; 
lower side black and sparsely rhizinate, brown and naked in a broad 
zone at the margins. Apothecia numerous, large, to 20 mm. in diame- 
ter, stalked, amphithecium rugose, white-maculate, disc perforate; 
hymenium 50-70 » high; spores 7-10%13-18 yu, the episporium 
1.0-1.5 » thick; pycnidia present, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C+ blood red, KC+ 
blood red, P—, atranorine and lecanoric acid present. 

The large synonymy of P. andina seems unaccountable. It is a 
conspicuous, extremely uniform lichen with a combination of mor- 
phological and chemical characters quite unique in the subgenus 
Amphigymnia. Large perforate apothecia are almost always present 
in great abundance. The maculae are usually distinct but may be 
rather faint on older lobes. It is most common in savannas of central 
and south-central Africa as far south as the Tropic of Capricorn but 
completely absent from the Cape area. It is much rarer in other 
tropical areas in South America and Asia (fig. 5). The sorediate 
counterpart is P. austrosinensis Zahlbr. 

Additional specimens examined: 


COLOMBIA: No locality, Blaghorne, s.d. (K). BOLIVIA: Cuaco: Fataranda, 
Fries L156 (S). PARAGUAY: Paraguari, Malme 1529 (MO). ARGENTINA: 
Satta: General Ballividn, Ordn, Digilio-Grassi 676 (MO). 

SUDAN: Mongalla, Dandy 523 (BM). IVORY COAST: Crercite DomBoxro: 
Mt. Orombo-Boka, Santesson 10724a (UPS). LIBERIA: Crenrrat Province: 
Piatah, Gbarnga, Baldwin 12383 (US). GUINEA: Cerrcure or N’Z&REKORS: 
Nimba Mtns., Santesson 10516b (UPS). SIERRA LEONE: Nijala, Deighton 
M5976 (K). CAMEROONS: Between Jaunde and Dengdeng, Mildbraed 8403 
(K). ANGOLA: Brnzurta: Between Benzuela and Coporolo, Degelius, Feb. 2, 
1960 (DEGEL); Bré: Chinguar, Degelius, Feb. 9, 1960 (DEGEL); between Vila 
General Machado and Coemba, Degelius, Feb. 10, 1960 (DEGEL); Cuanza Sut: 
Near Santa Comba Dao, Degelius, Mar. 3, 1960 (DEGEL); Capir, Amboim, 
Gossweiler 9907 (BM); Huta: Bimba, Humpala Plateau, Degelius, Feb. 6, 1960 
(DEGEL); Matance: Northeast of Lunda, Dundo, Gossweiler 13928 (US); 
Moxico: Luso, Degelius, Feb. 11, 1960 (DEGEL). REPUBLIC OF CONGO: 
Brazzaville, Degelius, Mar. 12, 1960 (DEGEL). CONGO: N. Kivu: Nyaru- 
sambo, Goma, Degelius, Mar. 16, 1960 (DEGEL). RUANDA-URUNDI: 
Usumbara, Degelius, Mar. 17, 1960 (DEGEL); west of Albertville, Héeg, Feb. 25, 
1930 (TRH, US); Patambalu, Tazlfer 43 (BR); Kiyaka-Kwango, Devred 2607 
(BR); near Lukafu, Schmitz 1839 (BR); Eala, Louis 2123 (BR). NORTHERN 
RHODESIA: Abercorn Distr., Bullock 2103 pars (K). SOUTHERN RHO- 
DESIA: Zimbabwe, Héeg, Mar. 4, 1930 (TRH); Makoni, Hyles 215 (PRE); 
Salisbury, Montagu 1809 (PRE); Dora River, Umtali, Schwetckerdt 2219 (PRE). 
KENYA: CeEntrat Prov.: Nairobi, Maas Geesteranus 10273 (L, LD); 4 mi. W. of 
Nairobi, Maas Geesteranus 4393 (L, LD). TANGANYIKA: Dodoma, Héeg, 
Mar. 16, 1930 (TRH); Kigonsero, at Peramiho, Detrich, 1958 (KLEM). UGAN- 
DA: Budonga Forest, Bungoro, Taylor 3319 (BM); Kilpayo, Dimmer 848 (BM); 


938 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


West Nile, below Metu Rest Camp, Chancellor, Sept. 14, 1953 (BM). MOCAM- 
BIQUE: Catembe, Lorengo Marques, Borle 35 (PRE). UNION OF SOUTH 
AFRICA: Nata: Weenin, Héeg, Oct. 20, 1929 (TRH); Pinetown, Doidge 1546 
(PRE). 

MADAGASCAR: Céte de Mahafaly, Perrier de la Bathie, June 1910 (LD). 
INDIA: Madras, Shuter, s.d. (K); Mapurar Distr.: Shembaganur, Foreau 4176 
(AWAS). THAILAND: Doi Sutep, Tsuyama 10, 11 (TNS). TAHITI: Wragge, 
December 1905 (BM). . 

2. Parmelia araucariarum Zahlbr. Denkschr. Akad. Wiss. Math. Naturw. 
Wien 83:179. 1909. 


Type collection: Near SAo Amaro, Séo Paulo, Brazil, Schiffner 
(W, holotype; BPI, MICH, isotypes). 

Thallus often large, 10-15 cm. in diameter, loosely attached to bark, 
mineral gray with a yellowish tinge at times; lobes rotund, 7-14 mm. 
wide, becoming crowded and suberect toward the center, margins 
entire, sorediate, soralia linear to irregular, sorediate lobes sinuate and 
dissected, soredia granular; upper surface shiny, continuous to reticu- 
lately cracked with age; medulla pale yellow orange; lower side black 
and sparsely rhizinate, dark brown and naked in a broad zone along 
the margins. Apothecia and pycnidia unknown. 

Reactions: Thallus K+ yellow; medulla K+, C+, KC+ more 
intensely yellow, P—, atranorine, an unidentified pigment, and 
unknown substances present. 

Externally P. araucariarum closely resembles P. dilatata Vain., but 
the medulla is pigmented orange yellow and the lobe margins are not as 
dissected. It is apparently the sorediate phase of P. myelochroa Hale, 
although the chemicals accompanying the unknown pigment have not 
yet been fully determined for either species. Parmelia araucariarum 
is endemic and widespread in South America. 

Additional specimens examined: 

BRITISH GUIANA: Essequibo River, Moraballi Creek, Richards 239 (K). 
PERU: San Martin: Tingo Marfa, elev. 625-1100 m., Allard 20700, 21893, 
22524 (US); HuAnuco: Hacienda Exito, Churubamba, Mexia 8249a (F, US); 
AyacucHo: Aina, between Huanta and Rfo Apurimac, Killip & Smith 23142 (US). 
BRAZIL: No locality, Gardner (M). PARAGUAY: Villa Morro, Asuncion, 


Malme 1585 (S). ARGENTINA: Satta: Rio Pescado, Digilio-Grassi 384, 469 
(MO). 


3. Parmelia austrosinensis Zahlbr. Symb. Sin. 3:192. 1930. 

Parmelia olivetorum var. hypomelaena Kremplh. Verh. Zool. Bot. Ver. 
Wien 26:443. 1876. Type collection: Lima, Peru, Barranca (M, holo- 
type; UPS, isotype). 

P. olivetorum var. sorediosa Vainio in Welw. Cat. Afr. Pl. 2:399. 1901. Type 
collection: Serra de Muxatila, Cazengo, Angola, Welwitsch 112 (TUR, 
holotype; BM, isotype). 

P. hyporysalea (Vainio) Vainio var. cinerascens Vainio, Bot. Mag. Tokyo 
35:47. 1921. Type collection: Prov. Awaji, Japan, Mikuma 140 pr. p. 
(TUR, Vain. herb. no. 2564, holotype). 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 239 


P,. mesotropa var. compactior Zahlbr. Symb. Sin. 3:192. 1930. Type 
collection: Sanyingpan, north of Yunnanfu, Yunnan, China, Handel- 
Mazzettt 581 (WU, holotype; W, isotype). 

P. meridionalis Tavares, Port. Acta Biol., ser. B, 1:159, pl. 10, fig. 2. 1945. 
Type collection: Arredores de Setubal, Estremadura, Portugal, Tavares 
(LISU, holotype; not seen); Portela de Sacavém, Estremadura, Portugal, 
Tavares (M, paratype). 

P. cazengensis Dodge, Ann. Mo. Bot. Gard. 46:152. 1959. Based on 
P. olivetorum var. sorediosa Vainio. 

P. olivetoroides Dodge, Ann. Mo. Bot. Gard. 46:154. 1959. Type collec- 
tion: Cape of Good Hope, s.c. (FH-Tuck, holotype). 


Type collection: Gwanyinschan, near Kweiyang, Kweitschou, 
China, Handel-Mazzetts 10580 (WU, lectotype; BPI, W, isotypes; cf. 
Hale (1959a) for details of typification). 

Thallus 6-10 cm. in diameter, loosely attached to bark, very light 
mineral gray; lobes broad, 10-15 mm. wide, often ascending, rotund, 
margins sorediate, sinuate, soralia linear; upper surface plane, dull to 
shiny and more or less clearly maculate; lower side black and rhizinate 
at the center, naked and light brown, mottled to ivory, or white in 
a broad zone along the margins. Apothecia very rare, substipitate, 
to 10 mm. in diameter, amphithecium maculate, sparsely sorediate, 
disc widely perforate; hymenium 75 yu high; spores 6-10X10-16 uy, 
episporium 1.5 u; pyenidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C+ blood red, KC+ 
blood red, P—, atranorine and lecanoric acid present. 

Parmelia austrosinensis is the sorediate counterpart of P. andina 
Mill. Arg. It is a pantropical species most common in savannas 
of Africa and southern South America (fig. 6). When typically 
developed, the thallus is medium sized, suberect, with strictly mar- 
ginal soralia and a distinct white zone below. Maculae range from 
quite distinct to virtually absent. Apothecia are extremely rare. 


Additional specimens examined: 


U.S.: Missourt: Kirbyville, Taney Co., Hale 4400 (US); Lovutstana: New 
Orleans, Orleans Co., Tainturier s.n. (US); Texas: East of Fort Worth, Tarrant 
Co., Stretilzer s.n. (WISC); 18 m. east of San Antonio, Bexar Co., Hubricht B1926 
(MO); Pearsall, Frio Co., Darrow 4864 (DUKE); Duffau, Erath Co., Hale 5494 
(US); 10 mi. west of Leakey, Real Co., Darrow 4901 (US). MEXICO: San 
Luis Porosf: Del Maiz, Cain 27608 (TRTC, US); Guanasuato: South of San 
Luis de la Paz, Catn 27562a (TRTC, US); Pursua: 69 km. east of Puebla, Hale 
19336 (US); Paseo Viejo, Arstne 4197 (US); Veracruz: 9 km. east of Jalapa, 
Hale 19412 (US); Oaxaca: 53 km. northwest of Oaxaca, Hale 20801 (US); 
Cutapas: South of Teopisca, Hale 20508 (US). 

HAITI: Below Furcy, Dept. de Ouest, Wetmore 2678 (MSC, US). DO- 
MINICAN REPUBLIC: Gauma, Santiago, Wetmore 3911 (MSC, US); Los 
Amaceyes, Cordillera Setentrional, Wetmore 3392 (MSC). 

VENEZUELA: Lara: Barquisimeta, Saer 2, 135 (US). COLOMBIA: Chipa- 
que, André 1226 (K). URUGUAY: Parque Rivera, Montevideo, Herter 99689 


240 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


(MO, US). ARGENTINA: Pampa del Infierno, Meyer 2106 (MO); no locality, 
Lorentz & Hieronymus (M). 

ETHIOPIA: Addis Ababa, Negri s.n. (F). CONGO: Yangambi, Lueveo 
Plateau, Louis 8155 (BR); Kiyaka-Kwango, Devred 2607 (BR). KENYA: 
NarvasHa Prov.: Kinangop, Aberdare Mountains, Taylor 1616 (BM); CENTRAL 
Prov.: 5 km. west of Nairobi, Maas Geesteranus 10271 (L, LD); Rirr Vatiey 
Prov.: Menengai, Nakuru Distr., Maas Geesteranus 4610, 10281 (L, LD, US), 
4556, 4621, 4622 (L). MOCAMBIQUE: Lourenco Marqugs: 2 km. east of Nama- 
acha, Almborn 7106 (LD); 3 km. north of Vilahuiza, Almborn 6884 (LD); 
Lourengo Marques, Junoa 474 (PRE). SOUTHERN RHODESIA: Zimbabwe, 
Héeg, Feb. 2, 1930 (TRH); Gwelo, Héeg, Jan. 2, 19830 (TRH). UNION OF 
SOUTH AFRICA: Ornance FREE State: Clarens, Plank s.n. (PRE); TRANSVAAL: 
Pretoria, Smith 6274 (PRE), Almborn 5878 (LD); Sibasa, Reusburg 1573 (PRE); 
Bushbuckridge, Wager 1386 (PRE); Basurotanp: Near Masite Mountains, 
Hewitt sn. (TRH); Narau; Boschfontein Forest, Lions River Distr., Almborn 
8724 (LD); 4 mi. south of Pietermaritzburg, Almborn 8541 (LD); Weinen, Héeg, 
Oct. 20, 1929 (TRH); northeast of Eschowe, Eschowe Distr., Héeg, Sept. 8, 1929 
(TRH); Upper Umkomaas, Impendhle, Héeg, Oct. 3, 1929 (TRH); Innersdale, 
Estcourt, Héeg, Nov. 3, 1929 (TRH); north of Hluklurve, Nongoma, Zululand, 
Héeg, Aug. 17, 1929 (TRH); Cars Province: Koningskroon Farm, Elliot, Héeg, 
Dec. 1, 1929 (TRH); Port Alfred, Héeg, Dec. 16, 1926 (TRH); Woodbury, 
Alexandria, Héeg, Aug. 20, 1929 (TRH); Dordrecht, Héeg, Nov. 21, 1929 (TRH); 
Port Elizabeth, Héeg, August 1929 (TRH); Round House, Lions Head Distr., 
Arnell 1146¢ (LD); Oudtohoorn, Congo Caves, Arnell 1450a (LD); Wilderness, 
George, Arnell 1344b (LD); Kentani, Pegler 1231 (PRE); Kingwilliamstown, 
Victoria 33 (PRE); Erste River, Stellenbosch, Garside 100, 148 (BM); 13 mi. 
east of East London, Almborn 10666 (LD); east of Kirstenbosch, Wynberg 
Distr., Almborn 23 (LD); Grahamstown, Albany Distr., Britton s.n. (TRH), 
Almborn 10861 (LD); near Coega, Uitenhage, Almborn 4065, 4107 (LD). 

INDIA: Above village Naret, Askote, Almora, Northwest Himalayas, Awasthi 
3297 (AWAS). JAPAN: Prov. Toromi: Kawasaki, Kurokawa 51074 (LD, 
TNS, US). SUMATRA: Sitarang, Toba, Ouwehand 175 (BO); Karshoogvlakte, 
Lérzing 7052 (BO). JAVA: Tjiumbuleuit, near Bandung, Groenhart 8855, 8867 
(BO); Pasir Larongé, Lugard 10 (BO); Tjibodas, Neervoort 65 p.p. (BO); Tjipanas, 
Lugard 18 (BO). AUSTRALIA: QuEENSLAND: Brisbane, Bailey s.n. (BM). 


4, Parmelia crassescens Stirton, Royal Phil. Soc. Glasgow Proc. 10:161. 1877. 


Type collection: Amazonas, South America, Trail 17 (BM, holotype; 
GLAM, isotype). 

Thallus loosely adnate, 10-15 cm. wide, light mineral gray, turning 
buff in the herbarium; lobes rotund, 7-12 mm. wide, more or less 
convoluted in age, margins crenate to irregularly lobulate, soredia, 
isidia, and cilia lacking; upper surface dull, continuous but becoming 
rugose and reticulately cracked with age; lower side black and sparsely 
rhizinate at the center, brown and naked in a broad zone at the 
margins. Apothecia more or less adnate, 3-5 mm. in diameter, disc 
imperforate; hymenium 80-90 » high; spores 10-1220-25 uy, epi- 
sporium 1.5-2.0 » thick; pycnidia rare, conidia not seen. 

Reactions: Thallus K-++ yellow; medulla K+ yellow turning red, 
C—, KC—, P+ yellow orange, atranorine and norstictic acid present. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 9A 


The type of P. crassescens is a fragment which at first seemed to be 
identical with P. latissima Fée. It is chemically distinct from 
P. latissima, however, in producing norstictic acid, a rare acid in 
subgenus Amphigymnia. Two fertile collections from South America 
have consistently smaller spores than P. latissima. 

Additional specimens examined: 


BRITISH GUIANA: Basin of Rupununi River, Yupukari, Smith 2256 (K, 
MO, US). BRAZIL: Serra do Itatiaia, Hemmendorff 12 (8). 


5. Parmelia cristifera Tayl. London Journ. Bot. 6:165. 1847. Puares 1, 4 

Parmelia perforata var. ulophylla Mey. & Flot. Nov. Act. Acad. Caes.-Leop.- 

Carol. Naturf. 19, suppl. 1: 218. 1843. Type collection: Oahu, Hawaiian 
Islands (G, lectotype). 

P. hildebrandtii Kremplh. Linnaea 41:139. 1877. Type collection: Jo- 
hanna Island, Comoro Islands, Hildebrandt 1866c (BM, K, isotypes). 

P. hildebrandtii f. sorediosa Mill. Arg. Flora 74:376. 1891. Type collection: 
Comoro Island, Schweinfurth s.n. (G, lectotype; K, isotype). 

P. mesotropa Mill. Arg. f. sorediosa Mill. Arg. Flora 74:377. 1891. Type 
collection: Cachar, India, Keenan (K, holotype; G, isotype). 

P. latissima Fée f. cristifera (Tayl.) Hue, Nouv. Arch. Mus. Paris, ser. 4, 
1:205. 1899. 

P. claudelit var. clemensiae Vain., Phil. Journ. Sci. 4:659. 1909. Type col- 
lection: Lake Lanao, Mindanao, Philippines, Clemens 1319 (TUR, Vain. 
herb. no. 2542, lectotype). 

P. submesotropa Gyel. Repert. Sp. Nov. Fedde 29:288. 1931. Based on 
P. mesotropa f. sorediosa Mill. Arg. 

P. cristifera f. cinerata Zahlbr. Repert. Sp. Nov. Fedde 33:58. 1934. Type 
collection: Kuwarus, Formosa, Asahina 51 (W, holotype; BPI, isotype). 

P. gardneri Dodge, Ann. Mo. Bot. Gard. 46:179. 1959. Type collection: 
Brazil, G. Gardner (FH-Tayl, holotype). 

P. gossweileri Dodge, Ann. Mo. Bot. Gard. 46:153. 1959. Type collec- 
tion: Chiloango, Angola, Gossweiler 8091 (K, holotype). 

P. imerinensis Dodge, Ann. Mo. Bot. Gard. 46:143. 1959. Type collection: 
East Imerina, Madagascar, Hildebrandt (FH, holotype). 


Type collection: Calcutta, India, Wallich (FH-Tayl, lectotype). 

Thallus expanded, 10-25 cm. in diameter, loosely attached to bark, 
light mineral gray; lobes broad and rotund, 12-20 mm. wide, margins 
sorediate, soralia linear, sorediate lobes sinuous, more or less involute, 
cilia lacking; upper surface dull, continuous, cracked with age; lower 
side black and sparsely rhizinate, naked and brown in a broad zone 
along the margins. Apothecia rare, usually poorly developed, adnate, 
1-5 mm. in diameter, disc imperforate; hymenium 100-120 yu high; 
spores 13-18 26-35 yp; episporium 3-4 y thick; pycnidia rare, conidia 
not seen. 

Reactions: Thallus K+ yellow; medulla K+ yellow turning red, 
C—, KC—, P+ orange red, atranorine and salacinic acid present. 

Parmelia cristifera is one of the commonest pantropical Parmelias, 
collected frequently by all botanists. It is characterized by abundant 


242 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


marginal soralia and salacinic acid. Apothecia are very rare; spores 
are large, indicating a close affinity with P. latissima Fée, which is 
strictly tropical and primarily American. Parmelia cristifera is espe- 
cially common in tropical America and the Pacific region (cf. fig. 4), 
but it is surprisingly rare in Africa and unknown from all of eastern 
Africa. The only species with which P. cristifera might be easily 
confused is P. dilatata Vain., which differs in containing protocetraric 
acid (medulla K—), in having sparse development of cilia, and in 
more laciniate margins with coarse irregular soralia. 

Miller based P. hildebrandtii f. sorediosa on five syntypes from 
Africa, but only the one listed above was found at Geneva. The type 
collections of P. perforata var. ulophylla Mey. & Flot. were destroyed 
at Berlin, but fortunately Hillmann (1940) had examined the material. 
It consisted of a mixture of P. cetrata Ach. and P. cristifera Tayl. 
The isotype at Geneva is a mixture of the same two species, of which 
P. cristifera is selected as the lectotype. 


Additional specimens examined: 


U.S.: Frormpa: Suwanee River State Park, Suwanee Co., Hale 17573 (US); 
5 mi. north of Okeechobee, Okeechobee Co., Hale 17723 (US); 5 mi. north of 
Silver Glen Springs, Marion Co., Hale 17591 (US); Sanford, Seminole Co., Rapp, 
Merrill, Lich. Exs. 54 (FH, US); Highlands Hammock State Park, Highlands 
Co., Hale 16923 (US); 12 mi. west of Daytona Beach, Volusia Co., Hale 17083 
(US); Lakeland, Polk Co., McFarlin 230 (FLAS); Hillsborough State Park, 
Hillsborough Co., Schallert sn. (FLAS). MEXICO: Jautsco: Santa Cruz de 
Vallarta, Mexia 1299 (US); Hipateo: Cuyamaloya, Pringle 10753 (US); Vera 
Cruz: 15 mi. south of Catemaco, Hale 19846, (US); 46 km. southwest of junction 
of highways 140 and 155, northeast of Huatusco, Hale 19463 (REN, US), 19456 
(MSC, US); Pursia: Cholula, Arséne 4297 (US); Curapas: El. Suspiro, 10 km. 
north of Berriozdébal, Hale 20220 (US); El Sumidero, near Tuxtla Gutiérrez, 
Hale 20073 (8, TNS, US); Hacienda, Matuda s.n. (MICH). BRITISH HON- 
DURAS: Bermuda Bank, Lundell 1981 (MICH). NICARAGUA: Mosquito 
Coast, Schramm s.n. (US); Sangsangta Distr., Schramm 50 (US). COSTA 
RICA: Hamburg Finca, below Cairo, Prov. Limén, Standley 48670 (US); San 
Isidro Coronado, Alfaro 32398 (US). PANAMA: Cana, Zone: Juan Mina, 
Chagres River, Bartlett 16822 (MICH); Las Cascadas Plantation, near Summit, 
Standley 29500 (US). 

BAHAMAS: Great Bahama, Brace 3608 (NY); New Providence, Britton 3311 
(NY); Maidenhead Coppice, Britton 3233, 3238 (NY). CUBA: No locality, 
Wright, Lich. Cubae 68 (K, UPS, US); Pinar del Rio, Cuesta 395 (NY); Baracoa, 
Johnston s.n. (FH); near Santa Barbara, Isle of Pines, Ross s.n. (F); La Prenda, 
Manuel 36a, Hroram 5226 (BPI); Loma del Gato, Sierra Maestra, Leén 10614 
(BPI, NY); Puerto Boniato Ridge, Santiago de Cuba, Oriente, Imshaug 24652 
(MSC, US). HAITI: St. Michel de l’Atalaye, Leonard 7857 (NY); near Kala- 
croix, Dept. Artibonite, Leonard 7837 (US) ; northwest of Jacmel, Thomas 75 (MO), 
77 (NY, US); below Citadelle, south of Milot, Wetmore 2838 (MSC). DOMIN- 
ICAN REPUBLIC: Sabana de la Rosa, near Km. 28, on Durante highway, 
Allard 15982b, 15994 (US); Piedra Blanea, La Vega, Allard 18025 (US); Hato 
Major, Siboey, Thomas 53, (FH, MO, NY); Consuelo, Macoris, Taylor 126 (FH, 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 243 


NY); without locality, Raunkiaer 497 (C). JAMAICA: Without locality, 
Hansen (C); Mandeville, Cushman 11, 147 (FH); Westmoreland Hills, New 
Market, Britton 524 (FH, NY); Farm Hill, St. Thomas, Orcutt 3551 (US); near 
Castleton, Mazon 737 (US); Strawberry Hill, Cinchona, Plitt 46 (BPI); Fern 
Gully, St. Ann, Imshaug 15796 (MSC, US); Lime Hall to Green Park, St. Ann, 
Imshaug 15849 (MSC, US); Mt. Diablo, St. Catherine, Imshaug 13766, 13767 
(MSC); Wareika House, summit of Long Mountain, Zmshaug 13499 (MSC). 
PUERTO RICO: Barranquitas, Hioram 220, 221 (NY, FH), Sintenis 43 (G); 
near Cayey, Sintenis 27, 29 (G); near Adjuntas, Sintenis 77, 79, 110a (G), 96 
(G, K); Mayaguez, Fink 1282 (FH, M, MICH); Naranjito, Fink 273 (FH, 
MICH); Aibonito, Fink 1837 (FH, MICH); 7 miles south of Caguas, Heller 295, 
309 (FH, NY); Monte Llano, Cayey, Goll 460 (US); Caguitas, Goll 361 (US); 
Arecibo to Utuado, Howe 385 (NY); Mt. Morales, near Utuado, Howe 440 (FH, 
NY); Rio de Maricao, Britton 4057 (FH, NY); Vega Baja, Britton 1435 (FH, 
NY); San Narciso, Britton 7332 (NY); Monte Montoso, Britton 4152, 4173 
(FH, NY); Luquillo Mountains, Wilson 55 (FH, NY), 295 (NY); Campo Alegre, 
Stevenson 2477 (FH, US). VIRGIN ISLANDS: Virgin Gorda, Virgin Peak, 
Smith 10575 (US); Sr. Crorx: Mt. Eagle, Boergesen s.n. (C), Thompson 462 
(NY); Sr. Tuomas: Crown, Britton 1441 (FH, NY); without locality, Eggers 
sn. (C). GUADELOUPE: Without locality, Questel 3810 (MO), L’ Herminier 
s.n. (UPS); Prise d’Eau, LeGallo 2724 (MSC); near Lamentin, LeGallo 583 
(MO); Vernon, Basse-Terre, Degelius, June 23, 1958 (DEGEL). DOMINICA: 
Roseau, Evans 64 (FH, NY, YU). MARTINIQUE: Le Morne Rouge, De- 
gelius, May 20, 1958 (DEGEL); Ajoupa-Bouillon, Degelius, May 25, 1958 
(DEGEL). GRENADA: Epping Forest, St. David, Imshaug 16146 (MSC, 
US). 

SURINAM: Without locality, s.c. (K). BRITISH GUIANA: PI. Vryheid, 
Linder 949 (MICH); near Kaieteur Falls, Cowan & Soderstrom 2041 (US); along 
Mure-mure Creek, Kaieteur Plateau, Cowan & Soderstrom 2203 (US). VENE- 
ZUELA: Between Cotiza and Las Venados, near Caracas, Allert 78 (US) ; BoLfvar: 
Guaiquinima, R{fo Paragua, Killip 37845 (US). COLOMBIA: Dept. SANTANDER: 
Northern slope of Mesa de los Santos, Killip & Smith 14996 (US). PERU: 
Dept. San Martin: Tingo Marfa, Allard 20562 (US); Derr. Ayacucno: Aina, 
between Huanta and Rio Apurimac, Killip & Smith 22572: (US). BRAZIL: 
Near Santos, Sao Paulo, Schiffner s.n. (M). 

GUINEA: Issia, cercle of Dalva, Santesson 10400a (UPS). IVORY COAST: 
20 km. west of Séguéla, Santesson 10673b (UPS). LIBERIA: 10 miles east of 
Zwedru, Tchien Distr., Baldwin 12412 (US). NIGERIA: Nkami Road, Oban, 
Talbots.n. (BM). CONGO: Eala, Guesquiére 2081 (BR, US); Yangambi, Luweo 
Plateau, Louis 7150 (BR). ANGOLA: Buco Zau, Cabinda, Gossweiler 7240 
(BM); Sera Subluali, Maiambe, Gosswezler 8033 (BM). MAURITIUS: Robillard 
s.n. (M, US). BOURBON: Rodriguez 8, 21 (P). 

BURMA: Without locality, McMillan 279 (US). THAILAND: Udaun, 
Kerr 215 (K); Chieng Dao, Tsuyama 4 (TNS, US); Neeckey, Kostermans 532 
(BO). FORMOSA: Taroko, Ogata s.n. (TNS, US). PHILIPPINES: Bohol, 
Ramos 11585 (F); Tayasas: Lucban, Luzon, Elmer 7227 (US); Rizau: Luzon, 
Ramos 22477 (US). SUMATRA: Near Lubuk Sikaping, Tandjung Alai, Groen- 
hart 8991 (BO); Boekit Koelampi, Theunissen s.n. (BO); Fort de Kock, Jacobson 10 
(BO). SARAWAK: Kuching, Brooke 462(BM). BORNEO: Peak of B. Papan, 
Kutai, Meijer B2045 (BO). CELEBES: Rante Lemo, Kjeliberg 33 (BO). 
AMBON ISLAND: Laha, Ruinen s.n. (BO). NEW GUINEA: Nondugl, 
Western Highlands Distr., Hoogland 3194 (BM). 


244 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


FIJI: Virt Levu: Mba, vicinity of Nandarivatu, Smith 5968 (US). MAR- 
QUESAS ISLANDS: Nukuhiva, near Hakaui, Brown 472 (BISH). RARO- 
TONGA: Parks 22396 (F). PITCAIRN ISLAND: Parlver Valley Ridge, Sé. 
John 14963 (BISH). HAWAIIAN ISLANDS: Matt: Honokahau Drainage 
Basin, Forbes 587 (BISH); Kavat: Faurie 74, 75, 76, 77, 84 (BM); Hawalt: 
Milo, Faurie 853, 854 (BM); Oanu: Honolulu, Faurie 448 (BM); Pohakea Pass, 
Waianae Mountains, Fosberg 9510 (US); Kalimoa Valley, Rock 86 (US); Tantalus, 
Heller 2160 (US) ; without locality, Heller, Merrill Lich. Exs. 200 (M, US). NEW 
ZEALAND: Without locality, Banks & Solander, August 1769 (BM). 

6. Parmelia crocoides Hale, sp. nov. 

Thallus laxe adnatus, 6-12 cm. diametro, cinereo-glaucescens, lobis 
rotundatis, 8-14 mm. latis, margine integris vel aetate incisis, superne 
planus, nitidus, levissime albomaculatus, aetate rugosus, rimosus, 
sorediis, isidiis, ciliis destitutis, strato corticeo superiore 10-13 yu 
crasso, strato gonidiali 16-20 » crasso, medulla croceo-lutea, 110-130 yu 
crassa, strato corticeo inferiore 12-14 uw crasso, subtus niger, sparse 
rhizinosus, ambitu fusco-castaneus, late nudus. Apothecia ignota; 
pycnidia numerosa, conidiis non visis. Thallus K-+  flavescens; 
medulla K+, C+, KC+ intensius croceo-lutea, P—, atranorinum, 
acidum protolichestericum, et pigmentum K— ignotum continens. 

Type in the U.S. National Herbarium, collected between San Pedro 
de Montes de Oca and Curridabat, San José, Costa Rica, Feb. 2, 1924, 
by P. C. Standley (no. 32772). 

This peculiar species appeared at first to be a relative of P. latissima 
Fée but the upper surface is shiny and more or less maculate and the 
medulla is pigmented pale orange red. Protolichesteric acid is present 
in low concentration. This species is known only from the type 
locality. 


7. Parmelia defecta Hale, sp. nov. 

Thallus adnatus vel laxe adnatus, rigidulus, plerumque saxicola, 
6-12 cm. diametro, albicans, parum lobatus, lobis rotundatis, sinuatis, 
plus minusve congestis ad centrum, superficie et partim margine 
sorediatis vel pustulato-sorediatis, sorediis diffusis, granulosis, soraliis 
marginalibus irregulariter linearibus, superne opacus, laevigatus, 
strato corticeo superiore 14-18 y» crasso, strato gonidiali 16-20 p crasso, 
medulla alba, 70-90 u crassa, strato corticeo inferiore 12—15 u crasso, 
inferne niger, sparse rhizinosus, ambitu castaneus vel albo-variegatus, 
late nudus. Apothecia atque pycnidia ignota. Thallus K+ flave- 
scens; medulla K—, C+ rubescens, KC-+ rubescens, P—, atranorinum 
et acidum lecanoricum continens. 

Type in Lund University, collected in Indumeni Forest, Cathedral 
Park area, Bergville, Natal, Africa, Nov. 3, 1953, by Ove Almborn 
(no. 8934). 

This species appears to be the sorediate counterpart of P. soyawxi 
Mill. Arg. Both species are saxicolous and endemic to Africa. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 245 


Parmelia austrosinensis Zahlbr., normally a corticolous lichen, differs 
in broader, more or less suberect lobes with a pale zone below. These 
two sorediate species may intergrade to some extent but careful field 
studies would be needed to clarify their relationship. 

Additional specimens examined: 

UGANDA: Entebbe, Proctor, Mar. 9, 1959 (K, US). SWAZILAND: 15 mi. 
southwest of Piggs Peak, Mbabane, Almborn 7927 (LD). UNION OF SOUTH 
AFRICA: OrancGE FREE Stare: 5 mi. west of Ladybrand, Maas Geesteranus 
6521 (L); Natau: Indumeni Forest, Bergville, Almborn 9319 (LD); Carg Prov.: 
Dordrecht, Héeg, Nov. 21, 1929 (TRH); near the Punch Bowl, Louis Trichardt, 
Zoutpansberg, Almborn 6095 (LD). MADAGASCAR: Montasoa, Rdsdnen (H). 


8. Parmelia dilatata Vain. Acta Soc. Faun. Fl. Fenn. 7, no. 7:33. 1890. 

Parmelia cetrarioides f. integra Harm. Ann. Crypt. Exot. 1:327. 1928. Type 
collection: Botanical Garden, Singapore, Demange s.n. (P, holotype). 

P. robusta Degel. Goteb. Kungl. Vet. Vitterh. Samh. Handl., ser. B, 1, no. 
7:33. 1941. Type collection: Forét de Crannou, Finistére, France, 
Picquenard in 1898 (P, holotype). 

P. kauaiensis Zahlbr. in Magn. & Zahlbr. Ark. Bot. 31A, no. 6:99. 1944. 
Type collection: Halemanu to Kaholuamano, Kauai, Hawaii, Kusche 
5506 (BPI, holotype; W, isotype). 

P. lobulascens var. isidiosissima Dodge, Ann. Mo. Bot. Gard. 40:375. 1953. 
Type collection: Gbense, Sefadu, Sierra Leone, Adames (K, holotype). 
P. sieberi Dodge, Ann. Mo. Bot. Gard. 46:148. 1959. Type collection: 

Mauritius, Sieber 44 (FH, holotype). 


Type collection: Sitio, Minas Gerais, Brazil, Vainio, Inch. Bras. 
Exs. 397 (TUR, holotype, Vain. herb. no. 2548; BM, FH, M, UPS, 
isotypes). 

Thallus loosely attached to bark, 10-20 cm. broad, mineral gray 
or faintly yellowish gray; lobes rotund, 10-15 mm. wide, margins 
entire at the tips, laterally becoming lacerate, sorediate, soredia 
coarse, mostly in linear soralia or developing on short irregular 
laciniae, cilia lacking or abnormally developed in the axils, 0.5-1.5 
mm. long; upper surface smooth, rather shiny, faintly white-maculate 
or dull, reticulately cracked with age; lower side black and sparsely 
rhizinate, brown to mottled ivory and naked in a broad zone at the 
margins. Apothecia rare, 3-5 mm. in diameter, adnate, disc im- 
perforate; hymenium 70-80 » high; spores 8-10 18-22 y, episporium 
about 2 » thick; pycnidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ reddish, 
P+ brick red, atranorine, protocetraric acid, and rarely traces of 
usnic acid present. 

Parmelia dilatata is one of the commonest pantropical lichens. 
It is unfortunate that the oldest epithet, P. dilatata, is represented 
only by a single apparently abnormal collection from Rio de Janeiro, 
which contains a trace of usnic acid and is eciliate. The population 
lacking usnic acid and having at least some sparse cilia is represented 


246 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


perfectly by the holotype of P. robusta and is by far the commoner 
phase of this species. Development of cilia is especially variable. 
Specimens from the West Indies in particular may have conspicuous 
cilia in the lobe axils and very rarely at lobe tips. Cilia may be so 
abundant that confusion with P. subarnoldit des Abb. is not unlikely. 
However, P. subarnoldii has distinct cilia around the lobe tips as 
well as finer soredia and more regular, linear soralia. The soralia 
of P. dilatata are typically irregular and produced on small marginal 
laciniae. Possible points of confusion with P. eristifera Tayl. and 
P. fasciculata Vain. are discussed under these species. 


Additional specimens examined: 


U.S.: TENNESSEE: Cross Roads, Overton Co., Phillips 334 (US); Sourn Caro- 
LINA: Santee State Park, Orangeburg Co., Hale 16485 (US); 10 mi. southwest 
of Walterboro, Colleton Co., Hale 16574 (US); Grorera: 5 mi. south of Quitman, 
Brooks Co., Hale 16765 (US); 10 mi. northeast of Midway, Bryan Co., Hale 
16806 (US); Fiorina: 1 mi. south of Pittman, Lake Co., Hale 17750 (US); 26 
mi. west of Tallahassee, Leon Co., Pursell 100LF30 (MSC); Sebring, Highlands 
Co., McFarlin 566 (FLAS); Bartow, Polk Co., McFarlin 96 (FLAS); Tomoka 
State Park, Volusia Co., Hale 17054 (US); Myakka River State Park, Sarasota 
Co., Hale 16911 (US); Fort Myers, Lee Co., Standley 326 (US) ; Sanford, Seminole 
Co., Rapp s.n. (FLAS); 3 mi. west of Bunnell, Flagler Co., Hale 17039 (US) ; 
Oleno State Park, Alachua Co., Hale 16431 (US). MEXICO: Veracruz: 11 
km. cast of Las Vigas, Hale 20951 (DUKE, MSC, REN, §, TNS, US); Cutapas: 
Just south of Teopisca, Hale 20522 (COLO, DUKE, LISU, MSC, REN, §, 
TNS, US); Km. 1145 on highway 190, west of San Cristébal, Hale 20202 (COLO, 
US); 18 km. southeast of San Cristébal, Hale 20274 (US), 20250 (LISU, US); 
Lagos de Monte Bello, Hale 20413 (COLO, US); Mt. Ovando, Matuda 28 (TNS). 
GUATEMALA: Aura Verapaz: Cobdn, Standley 92061 (MO); Finca Mocea, 
Johnson 170i (US). HONDURAS: Comayagua: Vicinity of Siguatepeque, 
Standley & Chacén P6315 (F); Morazdn: La Montafita, Standley 12366 (F); 
Ex Parafso: Region of Quebrada de Dantos, Standley et al. 1273 (F). BRITISH 
HONDURAS: EI Cayo Distr., Mains 4098 (FH, MICH). NICARAGUA: 
JINoTEGA: Vicinity of Jinotega, Standley 9704 (F). COSTA RICA: San Josh: 
Vicinity of Santa Maria de Dota, Standley 42481 (US); Carraco: Cerro de La 
Carpintera, Standley 34209 (US). PANAMA: Currigvf: Chiriquf Volcano, 
Scholander s.n. (US). 

BERMUDA: Payutes, s.c. (K). CUBA: Pinar peu Rfo: Between Vifales 
and Pinar del Rio, Imshaug 25320 (MSC, US); Sierra de las Yequas, Leén 5264 
(FH); Istx or Pines: Rocky Point, Britton 15415 (FH, NY); Ortenre: Summit 
of Pico Turquino, Sierra Maestra, Imshaug 25128 (MSC, US); slopes of El Gato, 
Loma del Gato, Imshaug 24773 (MSC). HAITI: East end of Montagne Noire, 
near Kenscoff, Imshaug 22533 (MSC, US); ridge leading to Pic Macaya, Morne 
Macaya, Wetmore 3238 (MSC); summit of Téte Etang, Jmshaug 22608 (MSC). 
DOMINICAN REPUBLIC: Casabito, La Vega, Wetmore 3430 (MSC); be- 
tween Pico del Yaque and Chinguela, Wetmore 3723 (MSC, US); Constanza, 
near Valle Nuevo, Allard 17692, 17696 (US); Sabana de la Rosa near Km. 28, 
Allard 15971 (US). JAMAICA: John Crow Peak, Imshaug 15365 (MSC); 
slope of Catherine’s Peak, Blue Mountains, Imshaug 13399 (MSC, US); without 
locality, Hart 122 p.p. (NY); Albion, St. Ann., Imshaug 15914 (MSC); near 
Guys Hill, St. Ann., Imshaug 13647 (MSC). PUERTO RICO: Lares to 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 247 


San Sebastian, Britton 2789 (FH). DOMINICA: Roseau, Evans 57 (NY, YU). 

VENEZUELA: Cerro Yapacana, Rio Orinoco, Maguire 30818 (MO); Avila, 
Vogl s.n. (M); Caracas, Linden 380 (K); Botfvar: Ptari Tepui, Steyermark 
59952 (MO); Salto de Auraima, Rio Paragua, Killip 37371a (US). COLOMBIA: 
Norte DE SANTANDER: Regién del Sarara, Cuatrecasas 12525 (US); MEpELLfn: 
Vicinity of Medellin, Charetier 192 (US); Cunpinamarca: Bogotd, Lesdain s.n. 
(MO), Apollinaire sn. (MO, US). ECUADOR: Recreo, Eggers (F); without 
locality, André 826 (K). PERU: San Martin: Tingo Maria, Allard 21644, 
22424 (US), Morrow 9605 (US). BOLIVIA: Mapiri, Bang 1750 (US); Trinidad, 
Beni, Werdermann 2305b (S). BRAZIL: Minas Gerais: Sitio, Vainio Lich. 
Bras. Exs. 397 (BM, M), 538b (K); Goras: east of Rio Verde on Goiania Road, 
Cutler 8029 (F); Maro Grosso: Santa Anna da Chapada, Robert 650 (BM); 
Buriti, Serra da Chapada, Malme s.n. (8S); Coxipé, near Cuyaba, Malme 2198 (S). 

FRANCE: Coatloch, Finistére, des Abbayes, Lich. Armor. Exs. Sel. 31 (LD); 
Forét du Cranou, just northwest of St. Conval, Bretagne, Santesson 10278a 
(UPS, US). PORTUGAL: Serra de Cintra, Monserrate, Estremadura, T'avares, 
Lich. Lus. Sel. Exs. 45 (H, US, WIS). 

LIBERIA: Belleyella, Boporo Distr., Baldwin 12311 (US). GUINEA: 
Nimba Mountains, cercle of Zérékoré, Santesson 10562a (UPS). CONGO: 
Isalowe Reserve, Louis 8648 (BR); Kabalo, Katanga, Héeg, sn. (TRH). AN- 
GOLA: Moxico: Lucusse, Degelius, Feb. 13, 1960 (DEGEL); between Luso and 
Cachipoque, Degelius, Feb. 16, 1960 (DEGEL); Huma: Caconda, Degelius, Feb. 
8, 1960 (DEGEL); Cuanza-Sut: Faz. Ceres, Mt. Chitandalua, Degelius, Feb. 21, 
1960 (DEGEL); Bri: Munhango, Degelius, Feb. 10, 1960 (DEGEL); Caio, 
Hambe Region, Gossweiler 8013 (BM). MOCAMBIQUE: Tinga-Tinga, Inham- 
bane, Schelpe 4469, 4469a (LD); near Maxixe, Inhambane Mongue, Schelpe 4463a, 
b (LD); Inhaca Island, Leighton 3324, 3325 (LD). SOUTHERN RHODESIA: 
South of Felixburg Station, Héeg s.n. (TRH); M’Ulnaluska River, Richard s.n. 
(K). UNION OF SOUTH AFRICA: Transvaau: Near the Punch Bowl, Louis 
Trichardt, Zoutpansberg, Almborn 6244 (LD). 

SINGAPORE: Johnson A-109 (US). PHILIPPINES: Camp Keithley, 
Mindanao, Clemens 1300 (US). BORNEO: Sarawak, Brooke 10204 (BM). 
AUSTRALIA: New Sours Waturs: Tilba-Tilba, Render s.n. (BM); QuEENs- 
LAND: Cairns, Fitzalan s.n. (BM). 


9, Parmelia disparilis Nyl. Syn. Lich. 1:381. 1860. 
Parmelia odontata Hue, Nouv. Arch. Mus. Paris, ser. 4, 1:185. 1899. Type 
collection: Bourbon, Rodriguez in 1889 (P, holotype). 
P, tephrina Hue, Nouv. Arch. Mus. Paris, ser. 4, 1:183. 1899. Type collec- 
tion: Nilgherries Mountains, Coonoor, India, Gray in 1893 (P, holotype). 
P. appendiculata Fée f. disparilis (Nyl.) des Abb. Mem. Inst. Sci. Madagascar, 
ser. B, 7:11. 1956. 

Type collection: Nossi-Bé, Madagascar, Pervillé 1847-52 (P, 
lectotype). 

Thallus loosely adnate to bark, up to 15 cm. in diameter or more, 
mineral gray; lobes rotund, 10-14 mm. wide, often suberect, margins 
conspicuously laciniate, laciniae to 10 mm. long, becoming caniculate, 
cilia absent or very sparsely developed in axils, 0.5 mm. long; upper 
surface smooth, shiny, more or less distinctly white-maculate, reticu- 
lately cracked with age; lower side black and sparsely rhizinate at the 
center, brown to ivory or mottled and naked in a broad zone along 

729-018—65——6 


248 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


the margins. Apothecia numerous, to 15 mm. in diameter, pedicellate, 
amphithecium rugose, white-maculate, exciple variable, from entire 
to dentate or even laciniate, disc imperforate; hymenium 90-100 yu 
high; spores 8-10X16-21 u, episporium 1.5-2.0 » thick; pycnidia 
common on laciniae, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC-+ reddish, 
P+ brick red, atranorine and protocetraric acid present. 

Parmelia disparilis is very near P. zollingert Hepp except for the 
distinct maculae and laciniae. The spores of P. disparilis tend to be 
somewhat smaller, although perhaps not significantly so. It is a 
rare species occurring in Asia, Africa, and Mexico. Its relation to 
P. appendiculata is discussed under that species. 

Additional specimens examined: 

MEXICO: Curapas: El Sumidero, near Tuxtla Gutiérrez, Hale 20078 
(DUKE, MSC, REN, 8S, TNS, US); road to El Suspiro, north of Berriozdbal, 


Hale 20081, 20115 (US). 
CONGO: Isalowe Flower Reserve, Yangambi, Louis 6596 (BR, US). 


10. Parmelia dominicana Vain. Journ. Bot. Brit. & For. 34:32. 1896.  PLaTre 3 
Parmelia perlata var. flavogranulosa Vain. Ann. Acad. Sci. Fenn. 6, no. 7:13. 
1915. Type collection: Mt. Stewart, St. Croix, Raunkiaer 433 (TUR, 
Vain. herb. no. 2362, holotype; C, isotype). 
P. capitulifera Zahlbr. in Magn. Ark. Bot. 30B, no. 3:9. 1941. Type 
collection: Punaluu, Oahu, Rock 176 (W, lectotype; BPI, isotype). 

Type collection: Crater, Souffriére, Dominica, Hlliott 114 (TUR, 
holotype). 

Thallus adnate to loosely adnate, saxicolous or corticolous, up to 
15 cm. in diameter, mineral gray with a pale yellowish tinge to dis- 
tinctly yellowish green; lobes rotund, 8-12 mm. wide, margins soredi- 
ate laterally, soralia irregular to subcapitate, confluent, in part sub- 
laminal, tinged distinctly yellowish, cilia lacking; upper surface 
opaque, reticulately cracked with age; lower side black and sparsely 
rhizinate, brown and naked in a rather broad zone along the mar- 
gins. Apothecia very rare, 3-5 mm. in diameter, adnate, amphithe- 
cium sorediate, disc imperforate; hymenium 65-70 u high; spores 5-7 < 
16-18 yu, episporium 1.5 » thick; pycnidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ red, 
fading, P+ orange red, atranorine, usnic acid, and protocetraric acid 
present. 

Parmelia dominicana is distinguished by the yellowish farinose 
soralia caused by the large concentration of usnic acid in the soredia. 
Most yellow Parmelias produce usnic acid only in the cortex and the 
soredia remain white. The concentration of usnic acid in the cortex 
of P. dominicana varies considerably, specimens from tropical Amer- 
ica being more or less mineral gray with traces of usnic acid and 
specimens from Hawaii and Africa being distinctly yellowish green. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 249 


It is a typical weedy species in disturbed areas of tropical America, 
occurring more rarely in Hawaii and Africa (fig. 13). 
Additional specimens examined: 


U.S.: Nortu Carouina: Smith Island, Brunswick Co., Culberson 7996 (DUKE, 
US); Fuoripa: Gold Head Branch St. Park, Clay Co., Hale 17695 (US); San- 
ford, Seminole Co., Rapp, Apr. 4, 1924 (FLAS, US); Snake Key, Dade Co., 
Baker, s.n. (FLAS). MEXICO: Veracruz: 24 km. northwest of San Andrés 
Tuxtla, Hale 19775 (US); Oaxaca: Cerro San Felipe, Hale 20747 (US); Km. 686 
on highway 190, northwest of Tehuantepec, Hale 20645 (TNS, US); Cutapas: 
2 km. east of Oaxaca-Chiapas State line, on highway 190, Hale 19888 (S, US); 
Km. 956 on highway 190, west of Ocozocoautla, Hale 20596 (US). NICARA- 
GUA: Jinoreaa: Vicinity of Jinotega, Standley 9573 (F). 

CUBA: Pinar pet Rio: Alturas de Pizarras, Imshaug 25220, 25231 (MSC); 
above Matahambre, Imshaug 25314 (MSC); Ortentr: Slope of El Gato, Loma 
del Gato, Sierra Maestra, Imshaug 24755 (MSC). HAITI: Northwest of Jacmel, 
Thomas 75 (NY). DOMINICAN REPUBLIC: Sabana de la Rosa, near Km. 
28, Duarte Highway, Allard 15958 (US). JAMAICA: Bloxburgh Ridge, St. 
Andrew, Imshaug 15100 (MSC, US). PUERTO RICO: San Juan, Britton 292 
(FH, NY); hills between Cabo Rojo and San German, Britton 4318 (NY). 
LESSER ANTILLES: Virain Isuanps: Virgin Gorda, Fishlock 346 (FH, NY); 
Sr. Crorx: No locality, Raunkiaer 433 (C); Dominica: Roseau, Evans 59 (FH), 
Evans 62 (US, Y), Crater, Souffriere, Elliott sn. (TUR); no locality, Campbell, 
November 1958 (K); St. BarrHetemy: Vité, Le Gallo 426a (MO). 

COLOMBIA: Isla de Providencia, Proctor, Apr. 29, 1948 (MO, PH). 

ANGOLA: Bif: Between Coemba and Munhango, Degelius, Feb. 10, 1960 
(DEGEL); Huma: 84 da Bandeira, 10 km. north of S& da Bandeira, Degelius, 
Feb. 3, 1960 (DEGEL); Moxico: Lucusse, River Lungué Bungo, Degelius, Feb. 
13, 1960 (DEGEL, US); between Luso and Cachipoque, Degelius, Feb. 16, 1960 
(DEGEL). CONGO: Southwest of Albertville, Héeg, Mar. 5, 1930 (TRH). 
SOUTHERN RHODESIA: South of Felixburg, Héeg, Feb. 4, 1930 (TRH). 

HAWAII: Kavatr: Kilauea, Faurie 79 (BM); Manamaulu, Faurie 81 (BM, 
LD); Oanu: Pumaluu Mtns., Rock 176 (BPI, FH, W). 


11. Parmelia eborina Hale, sp. nov. PLATE 3 


Thallus late expansus, rigidulus, laxe umbrosa ad saxa adnatus, 
cinereo-albicans vel eborinus, 10-15 cm. latus, lobis rotundatis, 
pro parte imbricatis, 8-15 mm. latis, ciliis, sorediis, isidiis destitutis, 
superne opacus, continuus, strato corticeo superiore 18-20 u crasso, 
strato gonidiali 20-22 u crasso, medulla alba, 80-100 u crassa, strato 
corticeo inferiore 20-22 yw crasso, subtus ater, sparse rhizinosus, 
ambitu castaneus, late nudus. Apothecia rara, 3-5 mm. diametro, 
nonnihil pedicellata, disco imperforato; hymenium 70-80 y» altum; 
sporae 5-7X11-14 wu, episporio 1 yu; pycnidia numerosa; conidia 
15-7 wp. Thallus K+ flavescens; medulla K—, C—, KC-+ fugiter 
rosea, P+ aurantiaca, atranorinum et acidum_protocetraricum 
continens. 

Type in the Chicago National History Museum, collected in the 
vicinity of El] Zamorano, Morazin, Honduras, Nov. 26, 1946-Jan. 9, 
1947, by P. C. Standley (no. 236). 


250 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Parmelia eborina is probably rather common on shaded rocks in 
Central America and more rarely in the West Indies. The dis- 
tinguishing features are the lack of soredia and isidia, the presence 
of protocetraric acid, and the ashy white, often monophyllous thallus 
which may break apart easily in the herbarium. One other species 
with protocetraric acid and without isidia, P. zollingert Hepp, has 
larger spores, a rigid thallus, and corticolous habit. 

Additional specimens examined: 

HONDURAS: Comayaaua: Comayagua, Standley 5762 (F); Morazn: 
Las Mesas, elev. 900 m., Standley & Williams 46, 49 (F); vicinity of El Zamorano, 
Standley 137 (F, US), 220 (F), 308 (F), 1692 (F, MO), 2267 (F), 11661 (F, US); 
Ex Paraiso: Las Casitas, Standley et al. 596 (F). 

DOMINICAN REPUBLIC: Moncién, Monte Cristo, Valeur 203 (US). 

COLOMBIA: Isla de Providencia, Proctor, Apr. 29, 1948 (MO). 


12. Parmelia ebulliens Hale, sp. nov. 

Thallus laxe adnatus, 8-12 cm. diametro, rigidus, cinereo-albicans 
vel eborinus, lobis 6-10 mm. latis, contiguis et partim imbricatis, 
marginibus integris vel crenatis, plus minusve convolutis irregulariter 
laciniato-lobatis, isidiis, sorediis, ciliis destitutis, superne planus, 
opacus, minute rimosus in aetate, strato corticeo superiore 20-25 yp 
crasso, strato gonidiali 22-26 » crasso, medulla inconspicue flavida, 
100-130 uw crassa, strato corticeo inferiore 14-16 yu, inferne niger, 
sparse rhizinosus, ambitu castaneus, anguste nudus. Apothecia 
numerosa, breve pedicellata, 5-8 mm. diametro, amphithecio rugoso, 
albomaculato, excipulo rimoso, radiato-fissurino, disco imperforato; 
hymenium 100 yu altum; sporae 10-15 25-28 y, episporio 2 » crasso; 
pycnidia numerosa, conidiis 14-5 yu. Thallus K+  flavescens; 
medulla K—, C—, KC—, P—, atranorinum, acidum incoloratum 
ignotum et pigmentum flavidum ignotum continens. 

Type in the University of Michigan, collected at Mt. Ovando, 
Chiapas, Mexico, April 1936, by E. Matuda. 

Parmelia ebulliens externally resembles a small fertile specimen of 
P. zollingert Hepp, or if sterile, P. eborina Hale. The pale-yellow 
medulla and peculiar chemistry set it apart from both of these species. 
The residue from acetone is abundant and resembles the residue of 
evernic acid. However, no crystals are formed in either G.A.W. or 
G.E., and addition of these reagents or KOH causes bubbling and 
dissolution of the residue. Lamellar plates are formed in G.A.Q, 
but these have not been identified further. Parmelia ebulliens is 
apparently corticolous in Mexico, but the Jamaican specimens are 
saxicolous. 

Additional specimens examined: 

MEXICO: Curapas: Mt. Ovando, Matuda, December 1937 (MICH, US). 


JAMAICA: Trail from Content Gap to Flora River, elev. 2850 ft., St. Andrew, 
Imshaug 14365 (MSC). 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 251 


13. Parmelia endosulphurea (Hillm.) Hale, comb. nov. 

Parmelia tinctorum var. endosulphurea Hillm. Repert. Sp. Nov. Fedde 48:8. 
1940. 

P. praetervisa Mill. Arg. var flavicans Mill. Arg. Flora 71:494. 1888. 
Type collection: Cayey, Puerto Rico, Sintenis 21 (G, holotype). 

P. tinctorum var. chrysophora Zahlbr. ex Hillm. Repert. Sp. Nov. Fedde 
48:10. 1940. Type collection: Near Batavia, Java, Schiffner 3196 
(W, lectotype; WU, isotype). 

Type collection: Mexico, Orcutt 4728 (MO, holotype). 

Thallus loosely adnate, 10-15 cm. broad, light mineral gray; lobes 
rotund, 10-15 mm. wide, margins entire, cilia lacking; upper surface 
plane, opaque, densely isidiate, isidia 0.06-0.08 mm. in diameter, to 
3 mm. high, simple to sparingly branched; medulla pale orange yellow, 
especially near the upper cortex; lower side black and sparsely 
rhizinate, brown and naked in a broad zone at the margins. Apothecia 
rare, short-stalked, 5-10 mm. in diameter, disc imperforate; hymenium 
65-80 uw high; spores 6-9 X 19-23 yp, episporium 1.5-2 y» thick; pycnidia 
not seen. 

Reactions: Thallus K+ yellow; medulla K+, C+, KC+, more 
intensely yellow, P—, atranorine, an unidentified pigment, and un- 
known substances present. 

I published a note identifying a large number of tropical specimens 
previously identified as P. sulphurata Nees & Flot. with P. lindmanii 
Lynge (Hale, 1959a). More critical study has shown that P. lindmanii 
is actually a Hypotrachyna species, although quite broad lobed and 
chemically identical with the Amphigymnia species. Parmelia lind- 
mani has axial cilia and small spores (5-8 10-13 x); it is a rather 
rare species known only from southern Brazil, Uruguay, Paraguay, 
and Argentina. All of the remaining isidiate specimens with a pale 
sulphurous medulla are true Amphigymnias, lacking any trace of 
axial cilia, having broader lobes with a distinct bare zone below, 
and, when fertile, intermediate-sized spores (6-10 X 18-23 y). The 
correct name for this species is P. endosulphurea. It is extremely 
common in the Caribbean region but much rarer elsewhere (fig. 14). 
The chemistry is not clear except that the pigment is the same one 
found in P. appendiculata Fée, P. araucariarum Zahlbr., and others. 

Additional specimens examined: 

U.S.: ALABAMA: Pocosin, 4 mi. southeast of Troy, Pike Co., McCullough 453 
(US); Fioripa: Gainesville, Alachua Co., Nelson s.n. (FLAS); Suwanee River 
State Park, Suwance Co., Hale 17636 (US); Orange Park, Clay Co., Hale 17743 
(US); Everglades National Park, Dade Co., Degelius, Nov. 22, 1939 (DEGEL), 
Imshaug 23955 (MSC); Texas: 3 mi. east of Edna, Victoria Co., Darrow (TAES). 
MEXICO: Veracruz: 24 km. northwest of San Andrés Tuxtla, Hale 19763 (US); 
15 km. south of Catemaco, Hale 19797 (US); Curapas: 50 km. west of Tuxtla 


Gutiérrez, Hale 19941 (DUKE, MSC, REN, S, TNS, US). COSTA RICA: 
Guanacaste: Upper slopes of Cerro San José de Libano, Dodge 7962 (MICH); 


202 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Limén: Portete, Quiros 1478 (MO); vicinity of Gudpiles, Standley 37398 (US). 
PANAMA: Juan Mina, Chagres River, Canal Zone, Bartlett 16822 (MICH); 
Barro Colorado Island, Aviles 945 (F). 

BAHAMAS: New Providence, Britton 6583 p.p. (FH, NY). CUBA: Pinar 
peL Rio: Sumidero, Shafer 13926 (FH); Las Vuuas: Trinidad Mountains, 
Imshaug 24598 (MSC). HAITI: Below Citadelle, south of Milot, Wetmore 2826 
(MSC, US), 2859 (MSC); near Port Margot, Nash 99 (FH); Between Petionville 
and Ft. Jacques, Thomas 36a (NY). DOMINICAN REPUBLIC: Los Amaceyes, 
Cordillera Setentrional, Wetmore 3400, 3406 (MSC); Cerrazo, on ridge from 
La Cumbre to Santiago, Wetmore 3844 (MSC, US), 3871 (MSC) ; Guama, Santiago, 
Wetmore 3906 (MSC, US); west of Ciudad Trujillo, Allard 16794a, 16778 (US); 
Repressa Dam, near Mané, Allard 17285, 17287 (US); Hato Major, along Rio 
Magua, Thomas 32 (NY). JAMAICA: 1 mi. northwest of Stewart’s Town, 
Pierce 510 (PH); Farm Hill, St. Thomas, Orcutt 3551a (US); Birch’s Hill, Han- 
over, Imshaug 15724 (MSC); Mt. Diablo, St. Catherine, Imshaug 13782, 14241 
(MSC); Mandeville, Cushman 8 (FH); Santa Cruz Mountains, Britton 483 (FH); 
below August Town, St. Andrew, Imshaug 13637 (MSC); Hopewell, St. Ann, 
Imshaug 15805 (MSC, US). PUERTO RICO: Sardinera, Mona Island, Britton 
1801 (FH, NY); near Maricao, Sintenis 2 (G); near Aibonito, Sintenis 55 p.p. (G); 
Adjuntas, Sintenis 79 (G); Monte Montoso, Britton 4180 (FH, NY); Mayaguez, 
Fink 1326 (FH, G); Naranjito, Fink 279 (FH). U.S. VIRGIN ISLANDS: 
Sr. Tuomas: Bonne Resolution, Britton 444 (FH, NY); Sr. Jonn: Makumli, 
Raunkiaer 442 (C); Sussanaberg, Boergesen s.n. (C). BRITISH VIRGIN 
ISLANDS: Tortoua: Eggers 3192 (C), Fishlock 502 (FH, NY); Virain Gorpa: 
Fishlock 345 (FH, NY). GUADELOUPE: Trois Riviéres, Le Gallo 421, 558 
(MO). MARTINIQUE: Without locality, Husnot 442 (G); Fonds-St.- Denis, 
Degelius, May 26, 1958 (DEGEL); La Pagerie, Degelius, May 17, 1958 (DEGEL). 
ST. LUCIA: Murray (G). BARBADOS: Welchman’s Hall Gully, Evans s.n. 
(YU). TRINIDAD: San Fernando Palmiste, Broadway 8987 (K). 

VENEZUELA: Méripa: Rio Chama Valley, Mérida, Dennis 1673 (K). 
COLOMBIA: Botfvar: Torrecilla, near Turbaco, Killip & Smith 14675 (US). 
CHILE: Without locality, Barclay, 1836 (BM). BRAZIL: Rio pp JANEIRO: 
Rio de Janeiro, Glaziou 1842 (M), Milne, s.n. (K), Vainio, Lich. Bras. Exs. 480 
(BM); Banta: Sao Beulo, Leutzelburg 296a (M); Rio GRANDE DO Sut: Encrusil- 
hada do Sul, Cuezzo 13371 (F); SA0 Pauto: Near Conceicéo de Itanhaen, Schaffner 
(BM). 

IVORY COAST: Forét d’Akadiedon, des Abbayes s.n. (8S). MADAGASCAR: 
Fort Carnot, Decary s.n. (LD). 

JAVA: Buitenzorg Garden, Batavia, Schiffner 3196 (FH). FIJI: Suva, 
Edmondson s.n. (BISH, US). 

Localities already published by Hale (1959a) for Florida, Louisiana, Grenada, 
Fl Salvador, British Honduras, Nicaragua, and Uruguay (excluding the type of 
P. lindmanti Lynge) are not repeated in this list. 


14. Parmelia fasciculata Vain. Hedwigia 38:122. 1899. 
Parmelia fatiscens Lynge, Ark. Bot. 15, no. 1:1. 1917. Type collection: 
Santa Anna da Chapada, Mato Grosso, Brazil, Malme (8, holotype). 
Type collection: Bogot&é, Colombia, Weir 66 (K, holotype; TUR, 
isotype). 
Thallus loosely adnate on bark, 5-15 cm. in diameter, mineral gray, 
often with a yellowish tinge; lobes rotund, 6-12 mm. wide, margins of 
lobe tips entire, lateral margins laciniate and dissected, laciniae 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 253 


developing into large coralloid clusters to 4 mm. high, sometimes 
becoming pustular or sorediate-pustulate, cilia very rarely developed; 
upper surface plane, dull, reticulately cracked with age; lower side 
black and sparsely rhizinate in the center, brown and naked in a broad 
zone along the margins. Apothecia rare, 3-6 mm. in diameter, 
substipitate, amphithecium irregularly pustulate, disc imperforate; 
hymenium 50-65 » high; spores 8-10X18-20 yu, episporium 1.5 yu 
thick; pycnidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ reddish 
orange, fading, P+ brick red, atranorine, protocetraric acid and traces 
of usnic acid present. 

The most unusual feature of P. fasciculata is the large coralloid out- 
growths which superficially resemble tiny thalli of Stereocaulon. They 
originate from marginal laciniae or directly from the upper cortex. 
The tips of the branches are warty and sometimes become pustulate 
but never develop true soredia. The species is quite rare. It seems 
to be related to P. dilatata Vain., which may develop similar, although 
far less conspicuous, coralloid outgrowths along with coarse soredia. 
Both species are faintly maculate. Parmelia ramuscula Hale and P. 
flavotincta Hale have similar coralloid structures but differ in chemistry. 

Additional specimens examined: 

JAMAICA: East Peak, Blue Mountains, Imshaug 14833 (MSC). 

LIBERIA: Firestone Plantations, Harbel, R. Hale, Oct. 1958 (US). 

15. Parmelia latissima Fée, Ess. Crypt., suppl. 119, pl. 38, fig. 4. 1837. 

Type collection: Jamaica, s.c. (G, lectotype). 

Thallus large, expanded, loosely attached to bark, 10-25 cm. in 
diameter ; lobes rotund, 8-20 mm. wide, margins entire; upper surface 
plane, dull, continuous or becoming reticulately cracked with age, 
soredia and isidia lacking; lower side black and rhizinate, brown and 
naked in a broad zone at the margin. Apothecia common, adnate 
to substipitate, disc imperforate; hymenium 100-120 uw high; spores 
14-16 X 28-32 yw, episporium 3-4 yu thick; pycnidia common, conidia 
not seen. 

Reactions: Thallus K+ yellow; medulla K+ yellow turning red, 
C—, KC—, P+ pale orange red, atranorine and salacinic acid present. 

Fée described P. latissima from two collections in tropical America. 
Fortunately Miiller-Argau kept fragments of both of these specimens. 
The lectotype follows the accepted concept of P. latissima, a plant 
with the medulla reacting K+ red; the other specimen is P. zollingeri 
Hepp (medulla K—, protocetraric acid present). Parmelia latissima 
represents the simplest kind of Amphigymnia, having more or less 
adnate imperforate apothecia and lacking isidia, soredia, cilia, and 
maculae. Parmelia zollingeri is very close, as is P. crassescens Stirt., 
but both of these species may be separated by different chemistry and 


254 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


smaller spores. The sorediate counterpart is P. cristifera Tayl. 
Parmelia latissima is virtually restricted to tropical America except 
for one locality in India (cf. fig. 3). The name has been used in- 
correctly in the literature many times and has often been cited as 
the parent species for numerous varietal taxa. 

Additional specimens examined: 


MEXICO: Veracruz: 15 km. south of Catemaco, Hale 19843a (US); Curapas: 
El Sumidero, near Tuxtla Gutiérrez, Hale 20187 (MSC, US); 50 km. west of 
Tuxtla Gutiérrez, Hale 21050 (DUKE, US); El Suspiro, 9 km. north of Ber- 
riozdbal, Hale 21232 (US); Campecne: Tuxpefia, Lundell 1303 (F). HON- 
DURAS: ArvAntipa: Lancetilla, near Tela, Standley 54210, 54606, 55415 (F). 
NICARAGUA: Vicinity of Casa Colorado, near El Crucero, Standley 8603 (F). 
COSTA RICA: Guanacaste: Vicinity of Tilardn, Standley 44428 (US); 
ALAJUELA: San Ramén, Brenes 14462a (MO); Carraao: Potrero at Santiago, 
Dodge 4557 (MICH); Turrialba, Mazon 206 (MO); El Mufieco, south of Navarro, 
Standley 33676 (US). 

BAHAMAS: New Providence, Britton 3239 (FH, NY); Marsh Harbor, Abaco, 
Brace 1658 (FH, NY). CUBA: Without locality, Wright, Lich. Cubae 67 (BM, 
FH, K, M, UPS, US); Pinar pet Rio: Source of Rio Taco-Taco, Sierra de los 
Organos, Morton 4280 (MO, US); San Diego de los Bafios, Harle & Murrill 258 
(FH), Palmer & Riley 607 (US); Hasana: Cayo Largo, Proctor s.n. (PH); 
OrIENTE: Gran Piedra, Shafer 9118 (FH); La Prenda, Hioram 2590 (NY), 6102 
(BPI) ; Loma del Gato, Ledén 9849, 9886, 9998 (BPI, NY), Imshaug 24784, 24805, 
24949 (MSC, US), Hioram 6694 (US). GRAND CAYMAN: East End Island, 
Lewis 11 (PH). HAITI: Northwest of Jacmel, Thomas 77 (FH, MO, US); Port 
Margot to Corrut, Nash 224 (FH); summit of ridge north of Forét des Pins, Dept. 
VOuest, Wetmore 3155 (MSC), Imshaug 22697 (MSC, US). DOMINICAN 
REPUBLIC: Vicinity of Constanza, La Vega, Allard 17694b (US). JAMAICA: 
Moseley Hall Cave, Imshaug 13669 (MSC, US), 18676 (MSC); Coopers Hill, Red 
Hills, St. Andrew, Imshaug 14145 (MSC, US), 13732, 13712, 138715, 18716 (MSC) ; 
Birches Hill, Hanover, Imshaug 15675 (MSC); Mandeville, Cushman 1 (FH, NY); 
Claremont, Orcutt 3979 (US); Castleton, Underwood 1938 (FH), Plitt s.n. (BPI); 
Hollymount, Mt. Diablo, St. Catherine, Imshaug 14218 (MSC, US); Silver Hill 
Gap, Blue Mountains, Imshaug 14083 (MSC); Beaufort, Wullschlaegel 1256 (M); 
Coldspring Gap, Purdie s.n. (IKK); near Lydford P.O., Proctor 8654 (PH); without 
locality, Eggers 3721, Hansen s.n. (C). PUERTO RICO: Mayaguez, Fink 1022 
(NY). GUADELOUPE: Pointe A Pitre, Questal 9 (MO). CURACAO: La 
Vela, Curran & Haman 513 (US). 

VENEZUELA: Botfvar: Rio Paragua, Salto de Auraima, Killip 37371 (MO, 
US). COLOMBIA: Cuoco: Port Utria, Taylor 879 (H, K). PERU: Ayacu- 
cHo: Estrella, Killip & Smith 23068 (US); Mito, Bryan 222 (F); Lamas, Spruce 
112/2 (K). BRAZIL: Maro Grosso: Santa Anna da Chapada, Robert 649 (BM, 
Kk), 650 (BM, K, US), 651 (K). 

INDIA: Calcutta, Wallich (BM). MARQUESAS: Nukuhiva, near Hakaui, 
Brown 472 (BISH, US). 


16. Parmelia mesogenes Nyl. Flora 68:609. 1885. 
Type collection: Orizaba Peak, Mexico, Galeotti 6958 (P, holotype). 
Thallus saxicolous or corticolous, expanded, rather adnate, to 15 
cm. broad, mineral gray; lobes rotund, 8-12 mm. wide, margins entire; 
upper surface plane, opaque, isidia, soredia, and cilia lacking; medulla 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 255 


white or turning orange red near the lower cortex; lower side black 
and moderately rhizinate, brown, shiny, and naked in a broad zone 
at the margins. Apothecia common, stalked to substipitate, 6-8 mm. 
in diameter, disc imperforate; hymenium 115-130 y high; spores 
12-14 X21-80 u, episporium 2.0-2.5 uw; pycnidia common, conidia not 
seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC—, P-, 
pigmented medulla K+ purple, atranorine, an unknown anthraqui- 
none, and unknown colorless substances crystallizing in G.E. 

Parmelia mesogenes has few distinguishing features since it lacks 
soredia, isidia, and maculae and is negative with color tests. The 
unknown acid crystallizes in G.E. as very large needle clusters. The 
pigment, which is found even in the lobe tips, is identical with the 
one in P. erasmia Hale and P. hypomiltoides Vain. Parmelia meso- 
genes is both saxicolous and corticolous in wet cloud forests where 
banana plantations are found. It is endemic to Mexico but will 
probably eventually be found in other parts of Central America. 

Additional specimens examined: 

MEXICO: Veracruz: Mirador, Liebmann 7562b (C); Teocello Canyon, just 
south of Xico, Hale 21150 (8, TNS, US), 21156 (DUKE, REN, US), 21175 (US); 
northeast of Huatusco, Hale 19521 (US); 7 km. north of Fortin de las Flores, 
Hale 19694 (US) ; 46 km. southwest of junction of highways 140 and 155, northeast 
of Huatusco, Hale 19407a (US); Curapas: El Sumidero, Tuxtla Gutiérrez, Hale 
20026 (US). 

17. Parmelia mesotropa Miill Arg. Rev. Mycol. 10:55. 1888. 
Parmelia saccatiloba Tayl. f. membranacea Lynge, Ark. Bot. 13, no. 13:67. 
1914. Type collection: Pilecomayo, Gran Chaco, Paraguay, Malme, 1893 
(S, holotype). 
P. subregressa Lynge, Ark. Bot. 13, no. 13:58. 1914. Type collection: 
Paraguari, Paraguay, Malme 1525B (S, holotype). 

Type collection: Asuncién, Paraguay, Balansa s.n., 1878 (G, 
lectotype). 

Thallus adnate to bark, 5-10 cm. in diameter, light mineral gray; 
lobes rotund, 5-8 mm. wide, margins entire to broadly crenate; 
upper surface plane, opaque, reticulately cracked with age, isidia, 
soredia, and cilia lacking; lower side black and sparsely rhizinate, 
brown and naked in a rather narrow zone along the margins. 
Apothecia numerous, 5-8 mm. in diameter, amphithecium maculate, 
disc imperforate; hymenium 60-90 u» high; spores 7-1115-21 4, 
episporium 1.5-2.0 uw thick; pycnidia common, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC—, P-, 
atranorine and caperatic (mixed with protolichesteric acid?) present. 

Parmelia mesotropa resembles P. latissima Fée and P. zollingeri 
Hepp in external appearance and the lack of soredia, isidia, and cilia, 
but it is too small to be confused with either of these species. It 


256 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


differs furthermore in the P— reaction. The sorediate counterpart 
of P. mesotropa appears to be P. praesorediosa Nyl., a much more 
widely distributed species. 

Additional specimens examined: 

MEXICO: Oaxaca: 77 km. northwest of Tehuantepec, Hale 20631 (US). 
GUATEMALA: Jurtapa: Jutiapa, Standley 75250, 75949 (MO); El Barrial, 
east of Jutiapa, Standley 75816 (F, US); between Jutiapa and La Calera, Standley 
76106 (F). 

BOLIVIA: El Beni, Trinidad, Werdermann 2389 (S). BRAZIL: Mato Grosso: 
Santo Antonio, near Cuyabd, Malme 215CB (5S); Santa Anna da Chapada, 
Malme 2392** (S). URUGUAY: Rio Verde, Chaco, Herter 84954 (MO) 
PARAGUAY: Pilcomayo, Gran Chaco, Malme, Sept. 7, 1893 (8S); Asuncién, 
Malme 1678** (S). 


18. Parmelia myelochroa Hale, sp. nov. PLATE 3 


Thallus laxe adnatus, usque ad 15 cm. latus, rigidus, viridi- vel 
albido-glaucescens, lobis primo rotundatis, demum late revolutis, 
paulo congestis, 8-12 mm. latis, margine parce dentato-laciniatulis 
vel integris, superne laevigatus, nitidulus, centrum versus rugosus, 
cortice continuo, sorediis isidiisque destitutis, strato corticeo superiore 
11-15 p crasso, strato gonidiali 11-15 p» crasso, medulla luteo-flavens, 
100-125 yu crassa, strato corticeo inferiore 11-15 yp crasso, inferne niger, 
sparse rhizinosus, ambitu castaneus, late nudus. Apothecia numer- 
osa, pedicellata, 7-15 mm. lata, disco imperforato, amphithecio 
rugoso, albo-maculato; hymenium 75-90 y» altum; sporae 9-13 K 21-26 
uw, episporio 1.5-2.0 n; pycnidia numerosa, conidiis non visis. Thallus 
K+ flavescens; medulla K+ intensius flavescens, C-++ flavescens, 
KC-+ flavescens, P—, atranorinum, acidum barbaticum (?) et pigmenta 
ignota continens. 

Type in the U.S. National Herbarium, collected in deciduous mist 
forest, elev. 1040 m., 50 km. west of Tuxtla Gutiérrez, Chiapas, 
Mexico, Mar. 18, 1960, by M. E. Hale (no. 21049; isotype in TNS). 

Parmelia myelochroa is a tropical American species rarely seen 
among older herbarium collections. It was recognized by Zahl- 
bruckner from Schiffner’s Brazilian collections as P. sulphurata f. 
nuda, an herbarium name only. Externally it resembles P. latissima 
Fée, with which it is often found. However, P. myelochroa has a 
conspicuous yellow-orange medulla and intermediate spores. The 
chemical components are still not fully known. P. araucariarum 
Zahlbr. and P. endosulphurea (Hillm.) Hale are respectively the 
sorediate and isidiate counterparts of P. myelochroa. 

Additional specimens examined: 

MEXICO: Cutapas: 50 km. west of Tuxtla Gutiérrez, Hale 20214 (US), 19909 
i US). HONDURAS: Comayraavua: Near Siguatepeque, Yuncker et al. 6499 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 257 


PERU: Selva Real, Tingo Marfa, Morrow 9654 (US, WJC). BRAZIL: 
Minas GeErRaAlts: Road to Sio Miguel, Mexia 5239 (MO, 8, US); Sao Paulo: Mt. 
Jaragud, near Taipas, Schiffner, June 1901 (BM, M, WU). 


19. Parmelia pancheri Hue, Nouv. Arch. Mus. Paris, ser. 4, 1:202. 1899. 


Type collection: New Caledonia, Pancher (P, holotype). 

Thallus large, 10-20 cm. in diameter, loosely attached to bark, 
mineral gray; lobes rotund, to 10 mm. wide, more or less imbricate 
and crowded toward the center, margins smooth, sometimes narrowly 
black rimmed, cilia lacking, upper surface smooth to rugose with age, 
soredia and isidia lacking, opaque; lower side black and sparsely 
rhizinate, brown to tan and naked in a broad zone along the margins. 
Apothecia large, to 30 mm. in diameter, stalked, amphithecium 
rugose-maculate, disc perforate; hymenium 50-60 y» high; spores 
5-9 X 12-18 u, episporium 1.0-1.5 » thick; pycnidia abundant. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ red, 
P—, atranorine and alectoronic acid present. 

When I first examined the holotype of P. pancheri, I considered the 
lack of cilia to be an abnormal condition since all other species with 
alectoronic acid have cilia. After examining several other collec- 
tions, I realized that the lack of cilia is normal for this species. 
Maculae are not noticeably developed. Except for the collection 
from Thailand, P. pancheri is known only from New Caledonia. 

Additional specimens examined: 

THAILAND: Doi Sutep, Tsuyama 7 (TNS,US). NEW CALEDONIA: No 
locality, Pionniero, 1907 (DUKE)* 

20. Parmelia peralbida Hale, sp. nov. PLATE 5 


Thallus laxe adnatus vel adnatus, membranaceus vel rigidulus, 
cinereo-candidus vel albicans, 5-10 cm. diametro, lobis rotundatis, 
vulgo monophyllosis, usque ad 15 mm. latis, ciliis destitutis, modice 
isidiatis, isidiis tenuibus, simplicibus, 0.03-0.080.3-1.0 mm., 
superne planus, nitidus, aetate leviter rimosus, strato corticeo superiore 
30-40 y» crasso, strato gonidiali 20 » crasso, medulla alba, 140-150 xu 
crassa, strato corticeo inferiore 20 » crasso, inferne nigricans, sparse 
rhizinosus, versus ambitum castaneus, nitidus, late glaber. Apothecia 
rara, adnata, ad 4 mm. diametro, disco imperforato, amphithecio 
isidiato; hymenium 35-40 » altum; sporae 5-78-10 uw (maturae?), 
episporio 1 u; pycnidia dispersa; conidia 16-7 yu. Thallus K+ 
flavescens; medulla K—, C—, KC+ fugiter rosea, P+ aurantiaca, 
atranorinum et acidum protocetraricum continens. 

Type in the Farlow Herbarium (Tuck), collected in Jamaica, 1884, 
by J. Hart (no. 124; isotype in K). 

This species is characterized by the large whitish thallus, the very 
thin unbranched isidia, and the presence of protocetraric acid. It is 


258 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


usually collected sterile so that the range of variation in spore size is 
uncertain. The low hymenium and the unusually small spores are 
noteworthy, if they prove to be consistent. Parmelia peralbida is 
known only from the Caribbean area. Two other species resemble 
it in having isidia and protocetraric acid. Parmelia saccatiloba Tayl. 
has large saccate lobes, coarser and shorter isidia, and large spores 
(24-26 uw long). It is known from four localities in the South Pacific. 
If it occurs among the sterile specimens from the Caribbean area, 
it would be difficult to tell it from P. peralbida. Parmelia koyaensis 
Asahina, an Asian species in subgenus Parmelia with slightly branched 
rhizines, is smaller but otherwise very similar in external appearance. 
Additional specimens examined: 


MEXICO: Cutapas: Lagos de Monte Bello, Hale 20397 (US). HONDURAS: 
Moraz&n: Vicinity of El Zamorano, Standley 239 (F, US); La Montafita, 
Standley 12373 (F). PANAMA: Cuiriquf: Monte Lirio, Seibert 1011 (MO). 

HAITI: Ridge between Forét des Pins and Petit Source, Wetmore 3076 (MSC). 
JAMAICA: No locality, Hart 102 pr.p. (NY); Main Ridge Gap, Blue Mountains, 
Imshaug 14757 (MSC); east of East Peak, Blue Mountains, I[mshaug 14889 
(MSC); New Haven Gap, Blue Mountains, Imshaug 15153 (MSC). 


21. Parmelia praesorediosa Nyl. Sert. Lich. Trop. Labuan Singapore 18. 1891. 
PuaTE 5 

Parmelia capitata Lynge, Ark. Bot. 18, no. 13:59, pl. 1, figs. 4, 5. 1914. 
Type collection: Rio Vermelho, near Bahia, Bahia, Brazil, Malme (S, 
holotype). 

P. sanctae-crucis Vain. Ann. Acad. Sci. Fenn. 6, no. 7:14. 1915. Type 
collection: Near Fair Plain, St. Croix, Boergesen (C, holotype; TUR, 
isotype). 

P. neglecta Asahina, Journ. Jap. Bot. 17:71, fig. 77. 1941. Type collection: 
Raisha, Formosa, Asahina F, 51 (TNS, holotype). 

P. luzonensis Ris. Ann. Bot. Soc. Zool. Bot. Vanamo 3:78. 1948. Type 
collection: Panai-Kabayan, Benguet, Luzon, Philippines, McGregor 8806 
(H, holotype). 

P. subcetrarioides des Abbayes, Bull. Inst. Fr. Afr. Noire 18:974. 1951. 
Type collection: Kankan, Guinea, Africa, des Abbayes (REN, lectotype; 
US, isotype). 


Type collection: Singapore, Almquist (H, Nyl. herb. no. 35547, 
holotype; 5, isotype). 

Thallus adnate, 5-10 cm. in diameter, mineral gray to buff; lobes 
5-8 mm. wide, rotund, margins suberect and sorediate, soralia often 
crescent-shaped and in part submarginal, cilia lacking; upper surface 
smooth, dull; lower side black and rhizinate in the center, brown or 
mottled ivory at the margins. Apothecia rare, 4-10 mm. in diameter, 
short-stalked, amphithecium rugose, sorediate, disc imperforate; 
hymenium 70-80 u high; spores 7-10 15-21 yu, episporium 1.2-2.0 u 
thick; pycnidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC—, P-, 
atranorine and caperatic acid present. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 259 


Parmelia praesorediosa is a pantropical species (fig. 15). It is 
both corticolous and saxicolous and occurs at low elevations, often on 
cultivated trees, apparently as a weedy species in areas disturbed by 
man. Margins of sorediate lobes are often sinuous and the soralia 
crescent-shaped. Externally it might be mistaken tor small specimens 
of P. cristifera Tayl. or P. dilatata Vain., but the medulla is K—, P—. 
The nonsorediate counterpart is P. mesotropa Mill. Arg., a rarer 
tropical American species. 

Additional specimens examined: 


U.S.: Sourn Carouina: Bulls Island, Charleston Co., Culberson 10031 (DUKE); 
near Creston, Calhoun Co., Culberson 7734, 7742 (DUKE); 3 mi. west of Sumter, 
Sumter Co., Culberson 7773 (DUKE); ALaBamMa: Dupree, Houston Co., McLennan 
6001001 (US); Froripa: 5 mi. east of Greenville, Madison Co., Hale 17607 (US); 
Nittaw, Osceola Co., Hale 17711 (US); 5 mi. south of Tallahassee, Leon Co., 
Hale 16725 (US); Seffner, Hillsboro Co., Cowan s.n. (US); Upsala, Seminole Co., 
Rapp 660 (FLAS); 4 mi. northeast of La Belle, Glades Co., Hale 16867 (US); 
Lovis1aNa: Grand Isles, Jefferson Parish, Drowet 9520 (F); Baton Rouge, East 
Baton Rouge Parish, McFarland 6 (WIS); 22 mi. south of New Orleans, Plaque- 
mines Parish, Smith 12394 (US); Texas: Houston, Harris Co., Fisher 5199a 
(FH). MEXICO: Veracruz: 9 km. east of Jalapa, Hale 19421 (TNS, US); 
Teocello Canyon, south of Xico, Hale 21142 (MSC, US); Oaxaca: Northwest of 
Tehuantepec at Km. 719 on highway 190, Hale 20620 (DUKE, US); Cutapas: 
El Zapotal, Tuxtla Gutiérrez, Hale 199938 (S, US); YucarAn: Tekax, Gaumer 
1210 (F). HONDURAS: ArtAntipa: Near Tela, Standley 53862 (F). 

BAHAMAS: New Providence, Britton 3311 (FH); Soldiers Road, Brace 9790, 
9907 (NY). CUBA: Without locality, herb Montagne (FH, K); Ise or PINEs: 
La Cunagua, Britton 14586 (FH, NY); Prnar peu Rfo: Valle de Silencio, Alturas 
de Pizarras, Imshaug 25293 (MSC, US); Hasana: Santiago de las Vegas, Hermann 
785 (F), Rotg 1 (MO); Cojimar, Arséne 10859 (US); CamaauEy: Camino Antén, 
MacBride & Dahlgren 74 (F); Orrenre: La Prenda, Manuel 5692 (BPI); El 
Gato, Loma del Gato, Imshaug 24743 (MSC); Puerto Boniato Ridge, Santiago 
de Cuba, Imshaug 24653 (MSC). GRAND CAYMAN: Near Georgetown, 
Imshaug 24381 (MSC); East End Island, Lewis 12 (PH). HAITI: Behind 
Hotel Randan, east edge of Les Cayes, Imshaug 23127b (MSC, US); Bayeux, 
between Cap Haitien and Le Borgne, Bartlett 17783 (WIS). DOMINICAN 
REPUBLIC: Hayto Mayor, Thomas 33 (MO, NY); Santiago, Imshaug 23801 
(MSC, US); Ciudad Trujillo, Santo Domingo, Allard 16182a (US); Valdesia 
Valley, Prov. Trujillo, Allard 17351, 17353 (US); along Rfo Inoa at Inoa, Santiago, 
Imshaug 23885 (MSC). JAMAICA: New Castle, Cushman 167 (FH); Clarks 
Town, Trelawny, Imshaug 16048 (MSC); Birch’s Hill, Hanover, Imshaug 15672 
(MSC); Stewart Town, St. Ann, Jmshaug 16012 (MSC, US); Hope River below 
August Town, Imshaug 13629 (MSC, US); Hillshire Hummock, St. Catherine, 
Imshaug 13687 (MSC). PUERTO RICO: Aibonité, Fink 1931 (FH); Manati, 
Fink 2120 (FH); Laguna Tortuguero, Britton 3869 (FH, NY); Rfo Piedras, 
Johnston 947a (FH, NY); San Juan, Hiorams.n. (FH, NY); Campo Alegre, Steven- 
son 2477 (US); near Cayey, Sintenis 38 (G). VIRGIN ISLANDS: Sr. THomas: 
Mandal, Britton 1311 (FH, NY); Bonne Resolution, Britton 446 (FH, NY). 
GUADELOUPE: Basse Terre, Le Gallo 525 (MO); Rivitres, Le Gallo 2602 
(MSC). ST. BARTHELEMY: Vité, Le Gallo 409 (MO); Saline, Le Gallo 560 
(MO). DOMINICA: Roseau, Evans 51, 60 (US, YU). MARTINIQUE: 


260 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Madiana Beach, Degelius, May 13, 1958 (DEGEL). TRINIDAD: Centeno, 
Earle s.n. (US); without locality, Lassen (C). 

VENEZUELA: Lara: Near Barquisimeta, Saer 49 (US). FRENCH GUI- 
ANA: Cayenne, Babington s.n. (UPS). PERU: San Martin: Tingo Marfa, Allard 
21453 (US). BRAZIL: Rio de Janeiro, Milne (K), Widgren (S, UPS); Mrnas 
Gerais: Rio Branco, Mareco, Mexia 5452a (US); Rio GranpE po Sut: Pareci 
Novo, Lima 82 (MO). PARAGUAY: Pileomayo, Gran Chaco, Malme, Sept. 6, 
1893 (S). ARGENTINA: Tucum4n: Pueblo Viejo, Killip 39515 (US). 

GUINEA: 3-4 km. east of Divo, cercle of Grand Lahou, Santesson 10382a 
(UPS); 50 km. west of Guiglo, cercle of Man, Santesson 10487 (UPS). REPUB- 
LIC OF CONGO: Brazzaville, Degelius, Mar. 12, 1960 (DEGEL, US). 
CONGO: Kabalo, Katanga, Héeg, Feb. 22, 1930 (TRH). UNION OF SOUTH 
AFRICA: Transvaat: Wyllie’s Poort, Zoutpansberg, Almborn 6410 (LD); 
Natau: Somkili, Nogoma, Héeg s.n. (TRH). 

INDIA: 4-5 mi. from Tanakpur, Uttar Pradesh, Awastht 3370 (AWAS). 
SUMATRA: Tandjung Alai, Groenhart 8990 (BO); Sawahpari, Groenhart 9210 
(BO); north bank of Lake Singdarak, Groenhart 9287 (BO). JAVA: Bandung, 
Groenhart 8856 (BO). NEW CALEDONIA: Compton s.n. (BM). 

Additional localities cited by Hale (1959a) (as P. sanctae-crucis Vain.) from 
Georgia, Florida, Alabama, Louisiana, the West Indies, Honduras, Nicaragua, 
Guinea, and Japan are not repeated here. 


22. Parmelia pseudotinctorum des Abbayes, Bull. Inst. Fr. Afr. Noire 13:973, 
1951. PuLaTE 1 
Parmelia nitens f. isidiosa Mill. Arg. Bot. Jahrb. Engler 20:255. 1894. 
Type collection: Ririre, Karapo, Seen region, Tanganyika, Africa, Stuhlmann 

3301 (G, holotype; K, isotype?, as 3201). 

P. soredica var. neghelliensis Cengia-Sambo, R. Accad. Ital. Miss. Biol. 
Paese Borana 380. 1939. Type collection: Borana, Ethiopia, Cufodontis 
193a (FI, holotype). 

P. neghelliensis (Cengia-Sambo) Dodge, Ann. Mo. Bot. 46:120. 1959. 

P. pseudotinctorum f. perrugosa des Abbayes, Bull. Inst. Fr. Afr. Noire 13:974. 
1951. Type collection: Dalaba, Fouta-Djalon, cercle de Mamon, Guinea. 
des Abbayes (REN, holotype). 

P. stuhlmannii Dodge, Ann. Mo. Bot. Gard. 46:137. 1959. Based on P. 
nitens f. isidiosa Mill. Arg. 


Type collection: Mt. Tonkoui, cercle de Man, Ivory Coast, des 
Abbayes, Aug. 14, 1948 (REN, lectotype; US, isotype). 

Thallus saxicolous, loosely adnate, 8-15 cm. in diameter, light 
mineral gray; lobes rotund, 8-12 mm. wide, margins entire to crenate; 
upper surface dull, isidiate, isidia inflated and irregular, 0.2-0.3 mm. 
thick, coarsely branched with age, to 0.5 mm. high; lower side black 
and sparsely rhizinate, light brown, shiny, and naked in a broad zone 
along the margins. Apothecia rare, 2-4 mm. in diameter, amphithe- 
cium coarsely isidiate, disc perforate or imperforate; hymenium about 
40 » high; spores poorly developed, 5-88-12 uy, episporium 1 4; 
pycnidia not seen. 

Reactions: Thallus K-++ yellow; medulla K—, C+ blood red, KC-+ 
red, P—, atranorine and lecanoric acid present. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 261 


Parmelia pseudotinetorum is identical with P. tinctorum Nyl. in 
habit and chemistry. It differs chiefly in having large inflated isidia, 
whereas P. tunctorum has uniformly thin cylindrical or even granular 
isidia. 

Additional specimens examined: 

IVORY COAST: Mont Tonkoui, cercle of Man, Santesson 10633 (Almb. Lich. 
Africani 30) (LD, UPS, US, WIS); Mankono, cercle of Seguela, Santesson 10711 
(UPS). 


23. Parmelia ramuscula Hale, sp. nov. Puate 1 


Thallus laxe adnatus, 8-10 cm. diametro, albicans, lobis rotundatis, 
10-15 mm. latis, margine integris, centrum versus laciniato-dissectis, 
laciniis isidiiformibus, coralloideo-ramosis, usque ad 1 cm. altis, 
isidiis fatiscentibus, demum sorediatis, margine ciliatis, ciliis sparsis, 
1 mm. longis, superne opacus, reticulatim rimosus in aetate, strato 
corticeo superiore 24-28 y crasso, strato gonidiali 22-26 yu crasso, 
medulla alba, 110-140 » crassa, strato corticeo inferiore 13-15 4» 
crasso, subtus niger, sparse rhizinosus, ambitu fuscus, late nudus. 
Apothecia atque pycnidia ignota. Thallus K-+ flavescens; medulla 
K-+ rubra, C—, KC—, P+ aurantiaco-flava, atranorinum et acidum 
salacinicum continens. 

Type in the British Museum, collected at Lawas, Sarawak, May 31, 
1955, by W. M. A. Brooke (no. 10031; isotypes in L, US). 

This species has the same coralloid fatiscent outgrowths (pl. 1) 
that characterize P. flavotincta Hale and P. fasciculata Vain. Parmelia 
flavotincta has a more membranous thallus and well-developed cilia. 
Parmelia fasciculata is very close except for chemistry. Parmelia 
ramuscula may have some relation to P. cristifera Tayl., a marginally 
sorediate species, in the same way that P. fasciculata is related to 
P. dilatata Vain. 

Additional specimen examined: 

PHILIPPINES: Rizal, Luzon, Ramos s.n. (H). 

24 Parmelia rubifaciens Hale, sp. nov. PLATE 4 


Thallus adnatus, 8-12 cm. diametro, viridi-glaucescens (in herbario 
olivaceo-glaucescens), lobis rotundatis, 5-8 mm. latis, margine 
sorediatis, soraliis plus minusve conglutinatis, partim submarginalibus, 
ciliis destitutis, superne opacus, rarius rugulosus et demum irregulariter 
rimosus, strato corticeo superiore 20 u crasso, molli, strato gonidiali 
20-25 u crasso, medulla alba, 100-110 yp crassa, strato corticeo inferiore 
12-18 w crasso, inferne niger, sparse rhizinosus, versus ambitum 
castaneus, late nudus. Apothecia et pycnidia ignota. Thallus K+ 
flavescens; medulla K-+ rubescens, C—, KC—, P+ aurantiaca 
vel lutea, atranorinum et acidum norsticticum continens. 


262 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Type in the Chicago Natural History Museum, collected in the 
vicinity of Casa Colorado, near El Crucero, summit of Sierra Managua, 
elev. 800-900 m., Managua, Nicaragua, May 14-25, 1947, by P. C. 
Standley (no. 8409; isotype in US). 

Parmelia rubifaciens externally resembles P. praesorediosa Nyl. 
The soredia of P. rubifaciens however are somewhat more extensive, 
in part submarginal, and often become conglutinated in dense clumps. 
This species may be related to P. crassescens Stirt., a nonsorediate 
species which also contains norstictic acid, but the collections seen 
so far are smaller and more adnate than those of P. crassescens. 

Additional specimens examined: 

MEXICO: Veracruz: 24 km. northwest of San Andres Tuxtla, Hale 19777 
(US); Yucarsn: Tekax, Gawmer 1210d (Ff). GUATEMALA: Prrtn: La Liber- 


tad, Lundell 2237 (MICH); Zacara: Gualdn, Kellerman, Jan. 3, 1906 (OS, US). 
BRAZIL: Minas Gerais: No locality, Burchell 1105 (K). 


25. Parmelia saccatiloba Tayl. London Journ. Bot. 6:174. 1847. 

Parmelia tinctorum var. inactiva Zahlbr. Denkschr. Akad. Wiss. Naturw. 
Wien 81:271. 1908. Type collection: Malifa, Upolu, Western Samoa, Rech- 
inger 4999 (W, lectotype). 

P. inactiva (Zahlbr.) Vain. Bot. Mag. Tokyo 35:47. 1921. 

Type collection: Pitcairn’s Island, Beechey (FH-Tayl, lectotype; 
H, isotype). 

Thallus large, loosely attached on bark, 8-15 cm. broad, mineral 
gray to buff; lobes broad and rotund, 10-20 mm. wide, older lobes 
broadly convoluted and saccate, margins smooth, cilia lacking; upper 
surface dull, reticulately cracked with age, moderately isidiate, isidia 
cylindrical, simple to branched, 0.03-0.08 mm. in diameter, 0.1-0.3 
mm. high; lower side black and sparsely rhizinate, brown and naked 
in a broad zone at the margins. Apothecia adnate, 5-8 mm. in 
diameter, disc imperforate; hymenium 110-130 y» high; spores 12- 
16 X 22-26 yp, episporium 2.5-3.0 u; pycnidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC-+ reddish, 
P+ brick red, atranorine and protocetraric acid present. 

The other syntypes listed by Taylor (Mauritius, Wright, FH- 
Tayl; Brazil, Hook. Herb., BM) are P. tinctorum Nyl., a common 
pantropical species with lecanoric acid. 

The name P. saccatiloba has only rarely been correctly used in the 
literature. Nylander (1885) reported that it was the same as P. 
zollingert Hepp, a nonisidiate species. Vainio (1890) conceived of it 
as a nonsorediate form of P. dilatata Vain. Lynge (1913) was misled 
by these erroneous concepts and identified P. mesotropa Mill. Arg. 
as P. saccatiloba. The lectotype of P. saccatiloba is actually an 
isidiate plant, very similar externally to P. tinetorum Nyl. but with 
tall thin cylindrical isidia and different chemistry. Parmelia 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 263 


peralbida Hale from tropical America has similar chemistry but a 
thinner thallus and smaller spores. Parmelia saccatiloba is apparently 
restricted to the South Pacific. 

Additional specimens examined: 

HENDERSON ISLAND: North end, St. John & Fosberg 15103 (BISH). 
FIJI: Ovalau Island, Herre 5719 (LD). MARSHALL ISLANDS: Arno Atoll, 
Horwitz 9102C (US). 

26. Parmelia setchellii Vain. Univ. Calif. Publ. Bot. 12:5. 1924. 

Type collection: Puadruu, Tahiti, Setchell « Parks (TUR, Vainio 
herb. no. 2456, holotype). 

Thallus loosely adnate on bark, 4-8 cm. in diameter, mineral eray; 
lobes rotund, often becoming crowded, 5-9 mm. wide, margins 
crenate, cilia lacking; upper surface plane, often faintly maculate, 
faintly white reticulate at the tips, finely reticulately cracked with 
age, coarsely pustulate or isidiate-pustulate in a broad marginal 
zone, pustules to 0.3 mm. thick, breaking open but not becoming 
sorediate; lower side black and rhizinate, light tan, shiny, and naked 
in a distinct but narrow zone along the margins. Apothecia and 
pycnidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ reddish, 
P+ brick red, atranorine and protocetraric acid present. 

This unique species is known only from the type collection. It is 
characterized by peculiar inflated isidia which often break open 
and erupt as pustules but do not become sorediate. The lower side 
has a relatively narrow bare zone. Further collections are needed 
before we can ascertain the range of variation and its relation to 
other species. 

27. Parmelia soyauxii Mill. Arg. Linnaea 9:32. 1880. 

Parmelia nitens Mill. Arg. Bot. Jahrb. Engler 20:255. 1894. Type collec- 
tion: Bukoba, Seen region, Tanganyika, Africa, Stuhlmann 4[095] (G, 
holotype; BM, isotype). 

Type collection: Pungo Andongo, Angola, Soyauz 246 (G, holotype). 

Thallus saxicolous, adnate, 6-12 cm. across, whitish to ashy mineral 
gray; lobes rotund, 6-9 mm. wide, soon crowded and more or less 
imbricate toward the center, margins entire to crenate, cilia lacking; 
upper surface shiny at the tips, continuous or becoming cracked with 
age and darkening from superficial fungal infestations; lower side 
black and moderately rhizinate, dark brown and naked in a broad 
zone along the margins. Apothecia adnate to substipitate, amphi- 
thecium dull, disc usually perforate; hymenium 65-75 p high; spores 
5-7 X 12-13 uw, episporium 1 p» thick; pycnidia present, conidia not 
seen. 

Reactions: Thallus K+ yellow; medulla K—, C+ deep red, KC+ 
deep red, P—, atranorine and lecanoric acid present. 


729-018—65——_-7 


264 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Parmelia soyauzii is a saxicolous species apparently most common in 
the dry savannas of western Africa. The thallus is ashy white, 
externally similar to P. breviciliata Hale, another saxicolous species 
with alectoronic acid and cilia. The sorediate counterpart of P. 
soyauaxti is P. defecta Hale. 

Additional specimens examined: 

ANGOLA: Morro de Lopollo, Welwitsch 25 (BM); above S84 da Bandeira, 
Degelius, Feb. 6, 1960 (DEGEL); between Si da Bandeira and Vila Arriaga, 


Huila, Degelius, Feb. 3, 1960 (DEGEL). SOUTHERN RHODESIA: Zim- 
babwe, Héeg, Feb. 2, 1930 (TRH). MADAGASCAR: Mandruka, Lemaitre (H). 


28. Parmelia tinctorum Nyl. Flora 55: 547. 1872. 

Parmelia latissima Fée f. isidiosa Mull. Arg. Linnaea 43:32. 1880. Type 
collection: Near Mayumba, mouth of Banya River, Gabon, Africa, Pechuel- 
Loesche (G, lectotype). 

P. praetervisa Mill. Arg. Flora 63: 276. 1880. Type collection: Java, Zollinger 
449b (G, lectotype). 

P. perlata var. platyloba Mill. Arg. Flora 65:316. 1882. Type collection: 
Japan, Brauns 5 (G, lectotype). 

P. meiosperma var. ecklonii Dodge, Ann. Mo. Bot. Gard. 46:141. 1959. 
Type collection: Cape of Good Hope, Africa, Ecklon (FH-Tuck, holotype). 

Type collection: Canary Islands, Déspréauz (?) (H, holotype). 

Thallus large, 8-20 cm. broad, loosely adnate on rock or bark, 
whitish mineral gray; lobes rotund, 10-15 mm. wide, margins entire, 
cilia lacking; upper surface plane, dull, densely isidiate, isidia papil- 
late-granular to cylindrical, coarse, 0.06—-0.10 mm. in diameter, to 2 
mm. high, simple to rarely coralloid-branched; lower side black and 
sparsely rhizinate at the center, brown, shiny, and naked in a broad 
zone at the margins. Apothecia very rare, to 20 mm. in diameter, 
exciple isidiate-dentate, amphithecium rugose, white-maculate, isi- 
diate, disc imperforate; hymenium 55-65 uw high; spores 7-10 13-15 
u, episporium 1.5 u thick; pycnidia rare, conidia 12-15 yu long. 

Reactions: Thallus K+ yellow; medulla K—, C+ blood red, 
KC-+ red, P—, atranorine and lecanoric acid present. 

This common weedy pantropical species is well known to all lichen- 
ologists. It is immediately identified by the brilliant C+ red reaction 
and by the simple often granular or papillate isidia. Vainio (1890) 
recognized this species as P. coralloidea (Mey. & Flot.) Vain. in prefer- 
ence to P. tinctorum, apparently because he used the concept of ab- 
solute priority. The name is illegitimate and, furthermore, a type 
fragment of P. perlata var. coralloidea Mey. & Flot. at Geneva seems 
to be a Hypotrachyna species, though too small for study. As Hillmann 
reported (1939), this specimen is C—. 

Additional representative collections examined: 


U.S.: Kentucky: 8 mi. northwest of Brownsville, Edmonson Co., Hale 13861 
(US); Virainra: Amburg, Middlesex Co., Luttrell 1709 (MO); Norru Caro.ina: 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 265 


Smith Island, Morton 2219 (US); between Atlantic Beach and Salterpath, Carteret 
Co., Culberson 6846 (DUKE, US); Sourn Caro.rna: Beaufort, Beaufort Co., 
Hale 7622 (US); 12 mi. southwest of Andrews, Georgetown Co., Hale 16548 (US) ; 
Isle of Palms, Charleston Co., Culberson 9041 (DUKE, US); TennesszE: Syca- 
more, Cheatham Co., Phillips s.n. (US); GeorG1a: 3 mi. southwest of Hinesville, 
Liberty Co., Hale 16757 (US); 5 mi. southwest of Waycross, Ware Co., Hale 
16826 (US); FLoripa: 5 mi. east of Greenville, Madison Co., Hale 17616 (US); 
8 mi. east of Ocala, Marion Co., Hale 17032 (US); northwest of Hilliard, Nassau 
Co., Hale 17623 (US); Tomoka State Park, Volusia Co., Hale 17068 (US); San- 
ford, Seminole Co., Rapp, Merr. Lich. Exs. 141 (BM, M, US); Indian River City, 
Brevard Co., Degelius sn. (DEGEL, US); Crewsville, Hardee Co., Hale 16875 
(US); near Pensacola, Escambia Co., Hale 7996 (US); ALABAMA: Oak Mountain 
Park, Shelby Co., Hale 7123 (US); Dothan, Houston Co., McLennan 6002002 
(US); Pocosin, Pike Co., McCullough 467 (US); near Epps, Sumter Co., McCul- 
lough 627 (US); Wilmer, Mobile Co., Hale 7174 (US); Misstssiprt: Lucedale, 
George Co., Hale 7782 (US); Collins, Covington Co., Hale 7971 (US); Tupelo, 
Lee Co., Hale 7808 (US); Lovurtstana: 22 mi. south of New Orleans, Plaquemines 
Parish, Smith 12392 (US) ; Jefferson Island, Iberia Parish, Drowet 9059 (F); Lake 
Charles, Calcasien Parish, Drouet 8743 (F); St. Martinsville, Martin Parish, 
Langlois, Lich. Bor.-Amer. 122 (H, M, MO, US), Decades N. A. Lich. 190 (BM, 
DUKE, US); Arkansas: Near Dallas, Polk Co., Hale 3941 (US); Texas: 7 mi. 
south of Silsbee, Hardin Co., Whitehouse 25955 (MO); near Avinger, Cass Co., 
Hale 5264 (US); near Woodville, Tyler Co., Hale 5226 (US); Karnack, Harrison 
Co., Hale 5324 (US); Nacogdoches, Nacogdoches Co., Hale 5435 (US). MEXICO: 
TAMAULIPAS: Between Mante and Antiguo Morelos, Moore 3636 (US); Jauisco: 
San Sebastian, Mexia 1541b (F); Veracruz: 96 km. east of Cérdoba, Hale 19722 
(COLO, LISU, MSC, US); Teocello Canyon, south of Xico, Hale 21163 (US); 
PurEBLA: St. Barbara, Arséne 808 (US); Oaxaca: Km. 686 on highway 190, 
Hale 20647 (MSC, REN, US); Curapas: south of Teopisca, Hale 20520 (8, TNS, 
US); El Sumidero, Tuxtla Gutiérrez, Hale 21210 (US); YucaTAn: Chichankanab, 
Gaumer 2255 (F); Tekax, Gawmer 1210 (F). GUATEMALA: SACATEPEQUEZ: 
Near Antigua, Standley 58822 (F). HONDURAS: Comayaaua: Siguatepeque, 
Yuncker et al. 6460 (F, MO), Standley & Chacon 6773 (F); Morazhn: El Za- 
morano, Standley 241, 12275 (F). BRITISH HONDURAS: Et Cayo: Cohune 
Ridge, Mains 3828 (MICH). EL SALVADOR: San Vicente: Vicinity of San 
Vicente, Standley 3662 (F). NICARAGUA: Jinorgaa: Sierra west of Jinotega, 
Standley 10267 (F); Cuonrates: La Libertad, Standley 8789 (F). COSTA 
RICA: Guanacaste: Tilar4n, Standley 44476 (US); ALasuELA: Cerros de Pata 
de Gallo, Brenes 222 (MICH); Carraao: North of Cartago, Standley 49590, 
49631 (US). PANAMA: Chiriqui, Wagner 334 (M). 

BERMUDA: Devonshire Marsh, Britton 167 (FH, NY). CUBA: Prnar DEL 
Rio: Northwest of Hotel San Vicente, Imshaug 25232 (MSC, US); Sierra de los 
Organos, Imshaug 25375 (MSC, US); Orrenre: El Gato, Loma del Gato, Sierra 
Maestra, Imshaug 24735 (MSC); Guanténamo, Hioram 18090 (M); Puerto 
Boniato Ridge, Santiago de Cuba, Imshaug 24661 (MSC). HAITI: Northwest 
of Jacmel, Thomas 66 (NY, US); west of Cap Haitien, Imshaug 22679 (MSC) ; 
below Citadelle, south of Milot, Wetmore 2849 (MSC); St. Michel de l Atalaye, 
Leonard 8015a (FH, NY, US). DOMINICAN REPUBLIC: East of La Romana, 
Hassler 1330 (WIS) ; Ciudad Trujillo, Santo Domingo, Allard 15709, 16182, 17012 
(US); Santiago, Imshaug 23800 (MSC, US); Gauma, Wetmore 3905 (MSC); 
Los Amaceyes, Cordillera Setentrional, Wetmore 3378 (MSC, US); Sousa Bay, 
Puerto Plata, Wetmore 3952 (MSC, US); without locality, Raunkiaer 495 (C). 


266 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


JAMAICA: Flora River, St. Andrews, Imshaug 14401 (MSC, US); Albion, St. 
Ann, Imshaug 15896 (MSC), 15920 (MSC, US); Birches Hill, Hanover, Imshaug 
15673 (MSC); summit of Montpelier, St. Catherine, Imshaug 14298 (MSC); 
Bulls Head, Clarendon, Bengry 334 (PH): New Market, Britton 534 (FH). 
PUERTO RICO: 7 mi. south of Caguas, Heller 308 (FH, NY); Lares to San 
Sebastian, Britton 2789 (NY); El Rfo, Goll 307 (US); Naranjito, Pink 295 (NY); 
Adjuntas, Sintenis 79 (M, NY), 87 (G); near Aibonito, Sintenis 55 p.p. (G); 
Mt. Morales, Britton 841 (FH, NY). U.S. VIRGIN ISLANDS: Sr. Tuomas: 
Crown, Raunkiaer 415 (C); Signalhill, Eggers sn. (FH); St. Joun: Bordeux, 
Britton 567 (FH, NY), Raunkiaer 402 (C). BRITISH VIRGIN ISLANDS: 
Virgin Gorda, Fishlock 324 (FH, NY); Tortola, Eggers 3192 (C). TRINIDAD: 
Port of Spain, Degelius, June 17, 1958 (DEGEL). 

VENEZUELA: Guarico: Between Uberito and Los Placeres, Prttier 12395 
(US); Feprrat Distr.: Caracas, Vogl s.n. (M), Dennis 1516, 1519 (K); Maracay, 
Médgdefrau 401 (M). COLOMBIA: Santranper: Between El Roble and Tona, 
Killip & Smith 19445 (US); Mesa de los Santos, Killip & Smith 15148 (US). 
GALAPAGOS ISLANDS: Santa Maria Isl., Taylor 864 (MO). PERU: San 
Martin: Tingo Marfa, Allard 21600, 22577 (US); Cuzco: Valle de Lacco, 
Herrera 2070 (US). BRAZIL: Minas Gerats: Vigosa Agricultural College, 
Mexia 5145a (US); Sitio, Vainio, Lich. Bras. Exs. 1082 (M), 614 (UPS); Rio 
pE JANEIRO: Rio de Janeiro, Copeland 2 (G), Darwin 463 (K); Maro Grosso: 
Near Santa Cruz, Moore 719 (BM); Santa Anna da Chapada, Malme. Lich. 
Austroamer. 81 (H, LD, UPS); Sio Pauto: Near Santos, Schiffner s.n. (M); 
Cruzeiro, Robert sn. (BM). PARAGUAY: Cerro Negro, Paraguari, Malme 
1480 (LD). ARGENTINA: Without locality: Lorentz & Hieronymus (M); 
Jusuy: Laguna de la Brea, Chaco, Fries 48a (S); Satta: Urundel, Ordn, Grassi 
5642 (MO); northwest of Urundel, Killip 39624 (US); Misiones: San Ignacio 
Gisela, Montes 36 (LD); without locality, Borman s.n. (MQ). 

CANARY ISLANDS: La Palma, Bornmiiller 3283 (LD); Puerto de la Cruz, 
Tenerife, Deglius, Jan. 21, 1960 (DEGEL). IVORY COAST: Danane, cercle of 
Man, Santesson 10491a (UPS); Mt. Orombo-Boka, cercle of Dimbokro, Santesson 
10724c (UPS). GUINEA: Mont Nimba, N’Zérékoré, Santesson 10515 (UPS). 
REPUBLIC OF CONGO: Brazzaville, Degelius sn. (DEGEL). CONGO: 
South of Albertville, Héeg s.n. (TRH); Kabalo, Katanga, Héeg s.n. (TRH); 
Victoria Falls, Héeg s.n. (TRH); southwest side of Lake Kivu, Degelius s.n. 
(DEGEL). ANGOLA: Moxico: Between Luso and Cachipoque, Degelzus s.n. 
(DEGEL, US); Bré: Chinguar, Degelius sn. (DEGEL); Matanse: Duque de 
Braganca, Degelius s.n. (DEGEL); Cuanza-Sux: 10 km. north of Cassonque, 
Degelius s.n. (DEGEL); Hurta: North of Quilengues, Degelius s.n. (DEGEL). 
MOCAMBIQUE: 3 km. north of Vila Luiza, Lourengo Marques, Almborn 6886 
(LD). SOUTHERN RHODESIA: Matopo Hills, Eyles 1178 (BM); south of 
Felixburg Station, Héeg s.n. (TRH); between Cashel and Melsetler, Schelpe 4046 
(BM). UNION OF SOUTH AFRICA: Transvaa.: 2 mi. north of Warmbad, 
Waterberg, Almborn 5895 (LD); near Punch Bowl, Zoutpansberg, Almborn 6332, 
6487 (LD); Nara: Boschfontein forest, Lions River Distr., Almborn 8677 (LD); 
Eschowe, Almborn 8492 (LD, US); Pietermaritzburg, Almborn 9629 (LD, US); 
Nottingham, Clouston 1564 (PRE); Carn Province: Near Port Elizabeth, Héeg 
sn. (TRH); Grahamstown, Albany, Héeg s.n. (TRH). MADAGASCAR: 
Mantasoa, Decary s.n. (P); Fort Dauphin, Elliott 2862 (BM). 

NEPAL: Langtang Khola, Polunin M161 (BM). INDIA: Darjeeling, Hima- 
layas, Awasthi 3874 (AWAS); Shembagar, Palni Hills, Awastht 4089 (AWAS). 
THAILAND: Doi Chieng Dao, Tswyama 14 (TNS); Pha Nok Kao, Loei, P.S. 


HALE—-PARMELIA SUBGENUS AMPHIGYMNIA 267 


(K); LAOS: Hase Plantations, Tsuyama 15, 22 (TNS). CHINA: Tien Tai Shan, 
Chekiang Prov., Chiao 14065 (US); Chihli Prov., Clemens 4014 (US). HONG 
KONG: Wright s.n. (US). JAPAN: Otani, Prov. Inaba, [koma 6228 (LD); near 
Mito, Ibaraki, Sato, Lich. Jap. 10 (LD); Ono, Koga, Omi Prov., Kurokawa 520026 
(LD, TNS, US); Shakudai, Shimamoto, Prov. Settu, Kurokawa 57253 (TNS, 
US). 

SUMATRA: Kebondjahe, Depari 4 (BO); Brastagi, Breedveld 21 (BO); Mt. 
Marapi, Groenhart 9152 (BO). JAVA: Kampong Koebang, Neervoort 2181 (BO); 
Tjibodas, Neervoort 41 p.p. (BO). BALI: Batoer, De Voogd s.n. (BO). NEW 
GUINEA: Nondugl, Western Highlands Distr., Hoogland 3194 p.p. (BM). 
GUAM: Agana, Necker 301 (US). MARSHALL ISLANDS: Ine Island, Arno 
Atoll, Stone 1120 (US). NEW HEBRIDES: Hog Harbor, Baker s.n. (BM). 
AUSTRAL ISLANDS: Naa Rurutu, Fosberg 12003 (BISH, MO). MANGA- 
REVA ISLAND: Near Atituiti Point, Fosberg 11112 (BISH, MO, US); Mt. 
Duff, St. John 14489 (MO). SAMOA: Upolu, Reinecke 13 (BO, US). TAHITI: 
Papeari, Herre s.n. (LD, MICH). HAWAIIAN ISLANDS: Kavat: Waiakale, 
Faurte 75 bis (BM). AUSTRALIA: Sydney, New South Wales, Home s.n. (BM). 


29. Parmelia zollingeri Hepp in Zoll. Syst. Verz. 9. 1854. PLATE 6 

Parmelia latissima var. corniculata Kremplh. Flora 61:463. 1878. Type 
collection: Cionega, Argentina, Lorentz & Hieronymus (M, holotype). 

P, neocaledonica Nyl. Flora 68:609. 1885. Type collection: New Caledonia, 
Vieillard (H, Nyl. herb. no. 35304, holotype). 

P. platyphyllina Vain. Hedwigia 46:168. 1907. Koh Chang, Thailand, 
Schmidt XXX (TUR, holotype; C, isotype). 

P, latissima var. minima Lynge, Ark. Bot. 13, no. 13:45. 1914. Type 
collection: Buriti, Serra da Chapada, Mato Grosso, Brazil, Malme 2243C** 
(8, holotype). 

P. bogoriensis Zahlbr. Ann. Crypt. Exot. 1:206. 1928. Type collection: 
Buitenzorg Garden Java, Overeem 4 (W, holotype; BO, MICH, isotypes). 

P. overeemii Zahlbr. Ann. Crypt. Exot. 1:204. 1928. Type collection: Mt. 
Tjibodas, Java, Overeem 94 (W, holotype; BO, isotype). 

P. litoralis Dodge, Ann. Mo. Bot. Gard. 46:169. 1959. Type collection: 
Kilifi, Kenya, Africa, Matt Cass, September 1939 (K, holotype). 

P. robertyt Dodge, Ann. Mo. Bot. Gard. 46:174. 1959. Type collection: 
Bouallé, Boka de Titiékro, Ivory Coast, G. Roberty 13941 (G, holotype). 


Type collection: Bantam, Java, Zollinger 1241 (L, holotype; G, H, 
isotypes). 

Thallus loosely adnate to bark, 10-20 cm. broad, olivaceous mineral 
gray; lobes broad and rotund, 15-18 mm. wide, margins entire or 
lobulate or sublobulate, frequently short ciliate in the axils of lobes, 
cilia 0.5-0.8 mm. long, sometimes branched; upper surface smooth, 
irregularly cracked with age, dull, soredia and isidia lacking; lower 
side black and sparsely rhizinate at the center, brown and naked 
in a broad zone at the margins. Apothecia common, 3-10 mm. in 
diameter, substipitate, amphithecium smooth, disc imperforate; 
hymenium 70-80 u high; spores 7-1018-22 y, episporium 1.5-2.0 pu 
thick; pycnidia common, conidia 8-10 yu long. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC-+ reddish, 
P-+ brick red, atranorine and protocetraric acid present. 


268 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Parmelia zollingeri is a variable widespread pantropical species. 
Externally it is extremely close to P. latissima Fée, which differs 
principally in chemistry (K-- red, salacinic acid) and larger spores 
(28-32 uw long). Most specimens of P. zollingert from tropical America 
lack cilia, but specimens from Asia often have more or less distinct 
axial cilia, up to 0.8 mm. long. We are, however, classifying P. 
zollingeri among the nonciliate species. Another area of variability 
is the development of marginal laciniae which may be richly developed, 
as in P. latissima var. corniculata Kremplh., to completely absent. 
Variability in the development of both cilia and laciniae is largely 
continuous and extremely difficult to classify. The only other species 
that can be compared with P. zollingeri is P. disparilis Nyl., which has 
laciniate lobes and apothecia, maculae, and somewhat smaller spores. 


Additional specimens examined: 


U.S.: FLroripa: Watson Hammock, Big Pine Key, Monroe Co., Killip 40869 
(US); Royal Palm Hammock, Dade Co., Small 7581 (US); Snapper Hammock, 
subtropical Florida, Britton 408 (NY, US). MEXICO: Veracruz: 7 km. north 
of Fortin de las Flores, Hale 19671 (US); Mirador, Ross 625 (M); Curapas: 2 km. 
north of highway 190 on road to Puebla Nueva, Hale 20186 (COLO, DUKE, 
LISU, MSC, REN, S, TNS, US); Km. 916 on highway 190, west of Ocozocoautla, 
Hale 20600 (TNS, US); Cameprecue: Calahmul, Lundell 1172 (F). GUATE- 
MALA: Perén: La Libertad, Lundell 2237 (Ff); Aura Verapaz: Between San 
Crist6ébal and Verapaz, Steyermark 43881 (MO). HONDURAS: Comayagua: 
Near Siguatepeque, Yuncker 6499 (US); ArLANtTIpa: Near Tela, Standley 55431 
(F). BRITISH HONDURAS: Belize River, Lundell 1824 (MICH). COSTA 
RICA: Guanacaste: Los Ayotes, near Tilarén, Standley 45522 (US); Mt. 
Irazi, Bruner 9504 (MO). 

BAHAMAS: New Providence, Britton 646 bis (NY); Watlings Island, Britton 
6205, 6217 (FH, NY); Abaco, Brace 1983 (FH, NY). GRAND CAYMAN: 
Ironshore Area, Spotts Beach, Imshaug 24585 (MSC, US); near Georgetown, 
Imshaug 24398 (MSC, US). CUBA: Without locality, Sagra in 1886 (UPS); 
CamaGuEy: La Gloria, Shafer 767 (FH); Pinar pew Rio: San Diego de los Bafios, 
Leon 5260 (FH); northwest of Hotel San Vicente, Imshaug 25245 (MSC, US); 
Oriente: Antilla, Britton 12523 (FH). HAITI: Furey, Leonard (FH). DO- 
MINICAN REPUBLIC: Sousa Bay, Puerto Plata, Wetmore 3926 (MSC). 
JAMAICA: Without locality, Wright s.n. (K), Newcastle, Cushman 168 (FH); 
Morces Gap, Blue Mountains, Jmshaug 13295 (MSC), Dunns River, St. Ann, 
Imshaug 15753 (MSC, US). 

VENEZUELA: Botfvar: Salto de Auraima, Rio Paragua, Killip 37371b (US). 
COLOMBIA: Norte pe Santanper: Alto de Santa Inés, Cuatrecasas 12451 
(US). PERU: Cuzco: Valle de Lacco, Herrera 2071 (US); Machupicchu, Herrera 
3263 (US); Loreto: Mishuyacu, Klug 1108 (US). BRAZIL: Santarém, Spruce 
112 (K). ARGENTINA: Without locality, Lorente & Hieronymus (M). 

MAURITIUS: McGregor s.n. (BM). LAOS: Hase Plantations, Tsuyama 6, 
16 (TNS). FORMOSA: Kuraru, Asahina s.n. (TNS, US). JAVA: Without 
locality, Horsfield s.n. (BM); vicinity of Goenoeng Boender, Palmer 611 (US). 
PHILIPPINES: Mt. Malinao, Luzon, Edafio 37206 (K), TONGA: Eva: Fuai 
plantation, Yuncker 15302 (US). 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 269 


2. Section Subflavescentes (Vain.) Gyel. (1932, p. 225) 


Section Amphigymnia* Subflavescens Vain. (1890, p. 35) 
Section Amphigymnia subsection Subflavescentes (Vain.) Hillm. (1934, p. 234). 


Type species: Parmelia delicatula Vain. 
Thallus as in section Amphigymnia except margins and apices of 
lobes distinctly ciliate. 


Subsection Subflavescentes 


This subsection may be divided into two series: Subflavescentes 
with usnic acid and Emaculatae (p. 277) without usnic acid. 


Series Subflavescentes 


Thallus yellowish green, usnic acid present; lower side dark brown 
along the margins; apothecia imperforate. 

This small series corresponds exactly to Vainio’s original group 
Subflavescentes. All of the species contain usnic acid in abundance. 
Except for the pantropical P. zanthina and the African endemic 
P. ochroglauca, the species occur chiefly in tropical and subtropical 
America. 


30. Parmelia aberrans (Vain.) des Abbayes, Bull. Inst. Fr. Afr. Noire, ser. 
A, 20:22. 1959. 

Parmelia xanthina f. aberrans Vain. Ann. Acad. Soc. Faun. Fl. Fenn. 7, 
no. 7:37. 1890. 

P. xanthina var. ciliata Zahlbr. Denkschr. Akad. Wiss. Math. Naturw. 
Wien 83:175. 1909. Type collection: Mt. Jaragud, near Taipas, Sao 
Paulo, Brazil, Schiffner, June 1901 (WU, holotype). 

Type collection: Sitio, Minas Gerais, Brazil, Vainio, Lich. Bras: 
Exs. 664 (TUR, Vain. herb. no. 2758, holotype). 

Thallus large, loosely attached on bark or humus, yellowish green, 
10-15 cm. in diameter; lobes rotund, 10-15 mm. wide, margins broadly 
crenate to isidiate-dissected on older lobes, ciliate, cilia 1.0-3.0 mm. 
long; upper surface dull to shiny, smooth to minutely rugulose and 
finely reticulately cracked on older lobes, isidiate, especially in a zone 
near the margins, isidia simple to coralloid-branched and sometimes 
apically ciliate, isidia 0.04-0.06 mm. in diameter, to 0.6 mm. high; 
lower side black, moderately rhizinate, dark brown and naked in a 
broad zone at the margins. Apothecia very short-stalked, 2-4 mm. in 
diameter, amphithecium rugose, isidiate, disc imperforate; hymenium 
65-75 uw high; spores 7-1010-16 yp, episporium 1 yu thick; pycnidia 
present, conidia not seen. 

Reactions: Thallus K-++ more intensely yellow; medulla K—, C+ 
rose, KC-++ deep red, P—, atranorine, usnic acid, and gyrophoric acid 
present. 


270 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


In his original description Vainio gives the diagnostic color test as 
“Medulla CaCl,O, non reagens.” The holotype specimen however is 
clearly C+ rose and contains gyrophoric acid. Since the holotype 
takes precedence over the description, P. aberrans must be regarded as 
C+. The C— population is P. zanthina (Vain.) Mill. Arg. Des 
Abbayes (1958) followed Vainio’s description when elevating f. aberrans 
to species rank and considered it to be C— and P. zanthina to be C+, 
a concept thus the opposite of ours. Parmelia aberrans is morphologi- 
cally indistinguishable from P. zanthina (including P. madagascariacea) 
but has a more restricted distribution, known so far only in tropical 
America (see fig. 12). 

Additional specimens examined: 

MEXICO: Curapas: Hacienda, Matuda s.n. (MICH); 8 km. east of Teopisca, 
Hale 20355 (DUKE, S, TNS, US). GUATEMALA: Zacapa: Between Santa 
Rosalia and San Lorenzo, Steyermark 43179 (MO, US). COSTA RICA: Guana- 
caste: Hacienda Santa Maria, Dodge 6708 (S). PANAMA: Curriquf: Chiriqu{, 
Wagner s.n. (M). 

PERU: No locality, Mathews s.n. (BM). BRAZIL: Minas Gerais: Caraga, 
Vainio, Lich. Bras. Exs. 1181 (BM, M, TUR, UPS); Rio pE Janurmo: Serra dos 
Orgiios, Burchell 2305 (K); Maro Grosso: Bocea da Serra, Malme 2748 (S, UPS) ; 
Santa Anna da Chapada, Malme s.n. (S). PARAGUAY: No locality, Grosse 
s.n. (MO). 


31. Parmelia conformata Vain. Acta Soc. Faun. Fl. Fenn. 7, no. 7:36. 1890, 
Parmelia conformata f. ciliolifera Vain. Acta Soc. Faun. Fl. Fenn. 7, no. 
7:36. 1890. Type collection: Sitio, Minas Gerais, Brazil, Vainio, Lich. 
Bras. Exs. 615 (TUR, Vain. herb. no. 2489, holotype; BM, isotype). 
P. xanthina var. subeciliata Mill. Arg. Hedwigia 30:228. 1892. Based on 
P. conformata Vain. 


Type collection: Sitio, Minas Gerais, Brazil, Vainio, Lich. Bras. 
Exs. 650 (TUR, Vain. herb. no. 2485, lectotype; FH, M, isotypes). 

Thallus adnate, 6-10 cm. in diameter, yellowish green; lobes 
rotund, 7-10 mm. wide, margins entire to dentate or incised toward 
the center, sparsely ciliate, cilia 0.5-1.0 mm. long; upper surface 
plane, continuous or reticulately cracked with age, faintly white- 
reticulate at the tips, densely isidiate, isidia simple to branched, 
0.04-0.07 mm. in diameter, 0.3-0.6 mm. high; lower side black and 
sparsely rhizinate in the center, brown and naked in a broad zone at 
the margins. Apothecia very rare, more or less pedicellate, am- 
phithecium isidiate, disc imperforate; hymenium 50-60 y high; 
spores 9-11 18-25 u, episporium 1.5-2.0 » thick; pycnidia not seen. 

Reactions: Thallus K+ yellow; medulla P+ red, K+ brown, 
C— atranorine, usnic, protocetraric, and fumarprotocetraric acids 
present. 

Parmelia conformata superficially resembles the other yellow isidiate 
species P. zanthina (Mill. Arg.) Vain. and P. flavescens (Kremplh.) 
Nyl. It differs in chemical constituents, larger spores, and sparse 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 271 


development of cilia. Vainio recognized a specimen with more 
conspicuous cilia as f. ciliolifera but this form has no taxonomic 
significance. Except for one locality in Africa, P. conformata is a 
tropical American species most common in pine forests at higher 
elevations in the West Indies. 

Vainio listed as a synonym P. caperata f. isidiosa Mill. Arg. The 
type of this taxon (G) is a fragment that I tentatively identify as 
P. cinerascens Lynge, a Hypotrachyna species. Vainio supplied spore 
data for P. conformata from this specimen, but since Miiller’s plant is 
a different species, these spore data should be ignored. 

Additional specimens examined: 


MEXICO: Curapas: 18 km. S.E. of San Cristébal, Hale 20296 (US); 50 km. 
west of Tuxtla Gutiérrez, Hale 19930 (US); 40 km. southeast of Comitdn, Hale 
20477 (US). PANAMA: Cuiriquf: Chiriqui, Scholander, February 1941 (MO, 
US). 

HAITI: Mission, Fonds Varettes, Leonard 3887 (FH); summit of Montagne 
Noire, near Kenscoff, Wetmore 2767 (MSC); west of Forét des Pins, Wetmore 
3202 (MCS) ; summit ridge on Téte Etang, Wetmore 2659 (MSC). DOMINICAN 
REPUBLIC: Valle Nuevo, Constanza, La Vega, Allard 16540a, 16562b, 16582b, 
17678 (US); Cerrazo, ridge from La Cumbre to Santiago, Wetmore 3842 (MSC); 
north side of Constanza, Cordillera Central, Imshaug 23722 (MSC). JAMAICA: 
Albion, St. Ann, Imshaug 15906 (MCS); Dolphin Head, Hanover, Imshaug 
15652 (MSC); Cooper’s Hill, Red Hills, St. Andrew, Imshaug 14156 (MSC). 

BRAZIL: Minas Gerats: Sitio, Vainio, Lich. Bras. Exs. 538b (K, TUR), 
650b (UPS), 981 (TUR). 

ANGOLA: Huma: Above 84 da Bandeira, Humpala Plateau, Degelius, Feb. 6, 
1960 (DEGEL, US). 


32. Parmelia delicatula Vain. Acta Soc. Faun. Fl. Fenn. 7, no. 7:35. 1890. 
Parmelia subcaperata f. ciliata Zahlbr. Bull. Herb. Boiss., II, 4:135. 1904. 
Type collection: Morro de Sao Sebastiao, Brazil, Damazio 940 (W, holo- 

type; G, isotype). 

P. magna Lynge, Ark. Bot. 18, no. 13:83. 1914. Type collection: SA0 Joao 
d’el Rey, Minas Gerais, Brazil, Malme 269 (S, holotype; LD, MO, UPS, 
isotypes). 

P. radians Lynge, Ark. Bot. 13, no. 13:85. 1914. Type collection: Sao 
Jo&io d’el Rey, Minas Gerais, Brazil, Malme 203 (S, holotype; MO, isotype). 

Parmelia microdactyla Hale, Contr. U.S. Nat. Herb. 36, part 1:21. 1960. 
Type collection: Rio de Janeiro, Brazil, Widgren (UPS, holotype; 8, 
isotype). 

Type collection: Caraga, Minas Gerais, Brazil, Vainio, Lich. 
Bras. Exs. 1256 (TUR, Vain. herb. no. 2754, holotype; BM, FH, 
M, UPS, isotypes). 

Thallus saxicolous, 5-15 cm. in diameter, yellowish green; lobes 
very variable, from rotund and monophyllous to rather narrow and 
subimbricate, 5-15 mm. wide, margins adnate to suberect, sparsely 
ciliate, cilia 1-2 mm. long; upper surface dull, often faintly white- 
pruinose, reticulately cracked with age, isidia and soredia lacking; 


272 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


lower side black and moderately rhizinate, dark brown and naked in 
a zone along the margins. Apothecia adnate to substipitate, 4-10 
mm. in diameter, disc imperforate; hymenium 55-75 yw high; pores 
5-810-14 yu, episporium 1 » thick; pycnidia abundant, conidia 
not seen. 

Reactions: Thallus K+ more intensely yellow; medulla K-+ 
yellow turning red, C—, KC—, P+ pale orange red, atranorine, 
usnic acid, and salacinic acid present. 

Parmelia delicatula is a saxicolous species common in the area 
around Rio de Janeiro. The variability of the species is very great, 
as one can surmise from the number of synonyms. The basic char- 
acters are marginal cilia of varying development, lack of soredia and 
isidia, and presence of salacinic and usnic acids. The lobes may be 
broad and flattened, as in the type of P. magna, or rather narrow with 
ascending margins, as in the types of P. delicatula and P. radians. 
Parmelia microdactyla, first thought to be distinct because of the small 
marginal lobules, is apparently only a rare morphological variant. 

Additional specimens examined: 

BRAZIL: Parani: Serrinha, Dusén, Lich. Austroamer. 85 (H, K, LD, MSC, 
S, UPS); Minas Gerais: Serra do Ouro Preto, Damazio, Zahlbr. Lich. Rar. 


Exs. 80 (BPI, FH, UPS); Serra de Caldas, Mosén 2318a, b (8S). URUGUAY: 
Rocua: Estancia Cerros, Hosseus 111 (H). 


33. Parmelia flavescens (Kremplh.) Nyl. Flora 68:607. 1885. PuaTE 11 

Parmelia glaberrima B flavescens Kremplh. Flora 52:223. 1869. 

P. glaberrima f. flavescens (Kremplh.) Kremplh. Flora 59:73. 1876. 

P. latissima f. flavescens (Kremplh.) Miill. Arg. Linnaea 43:37. 1880. 

P. mauriensis Hue, Nouv. Arch. Mus. Paris, ser. 4, 1:201. 1899. Type 
collection: Abrededores, San Luis de Potosf, Mexico, Maury 7651 (P, 
holotype). 

P. protoflavescens Zahlbr. Denkschr. Akad. Wiss. Math. Naturw. Wien 
83:176. 1909. Illegitimate name based on P. glaberrima B flavescens 
Kremplh. 

P. pseudoflavescens Zahlbr. Denkschr. Akad. Wiss. Math. Naturw. Wien 
83:176. 1909. Nomen nudum. 


Type collection: Brazil, Glaziow 1833 (M, holotype; H, isotype). 

Thallus saxicolous, expanded, up to 25 cm. broad, coriaceous, yel- 
lowish green; lobes rotund, 9-12 mm. wide, margins entire to lacerate 
or dissected, often isidiate, sparsely ciliate, cilia 1.0-1.5 mm. long; 
upper surface opaque, reticulately cracked with age, moderately 
isidiate, isidia simple or sparingly branched, 0.06—0.1 mm. in diameter, 
to 0.6 mm. high, infrequently ciliate; lower side black and sparsely 
rhizinate, dark brown and naked in a broad zone along the margins. 
Apothecia common, to 10 mm. in diameter, amphithecium rugose, 
maculate, sparcely isidiate, disc imperforate or rarely perforate; 
hymenium 40-55 uw high; spores 6-8 10-12 y, episporium 1 yu thick; 
pycnidia common, conidia not seen. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 273 


Reactions: Thallus K+ yellow or more intensely yellow; medulla 
K-+ yellow turning red, C—, KC—, P+ pale orange red, atranorine, 
usnic acid, and salacinic acid present. 

Parmelia flavescens is typically a saxicolous species characterized 
by salacinic acid and isidia. All other isidiate yellow species contain 
K— substances and are usually corticolous. The apothecia of P. 
jflavescens are adnate and most frequently imperforate, although 
rarely perforate. 

The nomenclatorial confusion created by Zahlbruckner has already 
been discussed (Hale, 1960). The identity of P. mauriensis Hue and 
P. flavescens is interesting in that P. mauriensis was formerly thought 
to be asynonym of P. subcrinita Nyl., an isidiate species lacking usnic 
acid. 

Additional specimens examined: 

MEXICO: Isla Socorro, Herrera 18 (MSC). GUATEMALA: Jauapa: Hills 
northeast of Jalapa, Standley 76788 (MO, US); Jalapa, Standley 76530 (M). 
HONDURAS: Morazin: El Zamorano, Standley & Williams 465 (F, US), 
Standley 1691 (MO). 

VENEZUELA: Between Caracas and La Guaira, west of Silla de Caracas, 
Santesson 6670 (S). COLOMBIA: Norte pe SantTanpDER: Vicinity of Toledo, 
elev. 1700-1900 m., Killip & Smith 20514 (US); Cunpinamarca: Bogotd, Weir 
58 (BM, K); Antioquia: Medellin, Charetier 191 (US). BRAZIL: Rio ps 
JANEIRO: Serra dos Orgios, Burchell 2385 (K). 


34. Parmelia miranda Hale, sp. nov. PLATE 11 


Thallus late expansus, membranaceus vel rigidulus, usque ad 10 
cm. latus, viridi-flavicans vel stramineo-albicans, lobis rotundatis, 
vage ramosis, margine sinuato-incisis, sorediatis, soraliis globosis 
vel elongatis, lobis sorediatis plus minusve involutis, ciliatis, cillis 
inconspicuis, 0.5-2.0 mm. longis, superne laevigatus, opacus, 
demum leviter albo-pruinosus, continuus vel paulo rimosus in aetate, 
strato corticeo superiore 12-14 uw crasso, strato gonidiale 26-35 u 
crasso, medulla alba 65-70 u crassa, strato corticeo inferiori 13-15 u 
crasso, inferne niger dense rhizinosus, rhizinis atris, nitidis, ambitu 
fusco-castaneus, late nudus. Apotheciaignota. Thallus K—; medulla 
K+ rubra, C—, KC—, P+ aurantiaca, acidum salacinicum et 
acidum usnicum continens. 

Type in the U.S. National Herbarium, collected on pine-scrub 
oak mountainside, Km. 686 on Hwy. 190, northwest of Tehuantepec, 
Oaxaca, Mexico, Mar. 31, 1960, by M. E. Hale (no. 20653). (Isotypes 
in COLO, DUKE, LISU, MSC, REN, 8, TNS). 

Parmelia miranda is characterized by the large yellowish green 
thallus and marginal soralia and cilia. There are only two other 
comparable species in the subgenus Amphigymnia, P. viridiflava 


274. CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Hale and P. ochroglauca Hale. Parmelia viridiflava is very close to 
P. miranda but differs in chemistry (protocetraric and fumarproto- 
cetraric acids). Parmelia ochroglauca, an African species, has 
distinctly revolute sorediate lobes and contains protolichesteric 
acid. Parmelia miranda is endemic to Mexico where it occurs 
in open oak-pine forests. 

Additional specimens examined: 

MEXICO: Oaxaca: 77 km. northwest of Tehuantepec at km. 719 on highway 
190, Hale 20615 (US); Cutapas: 8 km. east of Teopisca, Hale 20346 (US); 2 km. 
north of highway 190 on road to Puebla Nueva, Hale 20165 (8, US). 

35. Parmelia ochroglauca Hale, sp. nov. 

Thallus laxe adnatus, 8-15 cm. diametro, obscure viridiflavus, lobis 
rotundatis, 8-12 mm. latis, margine subcrenatis, ciliatis, ciliis 0.5- 
1.5 mm. longis, sorediatis, sorediis granulosis, partim submarginalibus, 
lobis sorediatis plus minusve revolutis, superne planus vel rugulosus, 
minute rimosus in aetate, strato corticeo superiore 13-16 yu crasso, 
strato gonidiali 20-25 yu crasso, medulla alba, 80-90 yu crassa, strato 
corticeo inferiore 13-15 » crasso, subtus niger, sparse rhizinosus, 
ambitu castaneus, late nudus. Apothecia atque pycnidia ignota. 
Thallus K-+ flavescens; medulla K—, C—, KC—, P—, atranorinum, 
acidum protolichestericum et acidum usnicum continens. 

Type in the Museum of the Royal Norwegian Society (TRH), 
collected near Barretts Berlin Mine, Barberton, Transvaal, Africa, 
Aug. 8, 1929, by O. A. Héeg (isotypes in LD, US). 

Although P. ochroglauca is known from only one collection, it ap- 
pears to be amply distinct because of a unique combination of charac- 
ters. At first glance it seems to be P. zanthina (Mill. Arg.) Vain., an 
isidiate species with identical chemistry. However, P. ochroglauca is 
distinctly though not conspicuously sorediate, the soredia originating 
for the most part submarginally and eventually causing the lobe mar- 
gins to become more or less revolute. The revolution is similar to but 
not so pronounced as that in P. perlata (Huds.) Ach. The soredia 
here are rather sparse and granular. This species is probably endemic 
to South Africa. 


36. Parmelia viridiflava Hale, sp. nov. 

Thallus laxe adnatus, membranaceus, mollescens, usque ad 10 cm. 
diametro, viridiflavicans, lobis rotundatis, parum lobatis, 9-12 mm. 
latis, integris vel centrum versus dentato-incisis, margine sorediatis, 
soraliis irregulariter sinuato-elongatis, centrum versus praecipue in 
axillis ciliatis, ciliis 1.0-1.5 mm. longis, superne laevigatus, nitidulus, 
aetate rimosus, ad apicem levissime albo-reticulatus, strato corticeo 
superiore 13-15 u crasso, strato gonidiali 18-20 u crasso, medulla alba, 
85-90 pu crassa, strato corticeo inferiore 16-18 yu, inferne niger, modice 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 275 


rhizinosus, ambitu castaneus atque late nudus, nonnunquam albo- 
variegatus. Apothecia pycnidiaque ignota. Thallus K—; medulla 
K—, C—, KC-+ fugiter rosea vel fulvescens, P+ aurantiaca, acidum 
usnicum, acidum protocetraricum et acidum fumarprotocetraricum, 
rarius atranorinum continens. 

Type in the U.S. National Herbarium, collected in mature pine 
forest, elev. 1220 m., at Lagos de Monte Bello, 50 km. southeast of 
Comitfin, Chiapas, Mexico, Mar. 25, 1960, by M. E. Hale (no. 20415; 
isotypes in DUKE, REN, 8S, TNS). 

This species reminds one at once of P. miranda Hale or P. domini- 
cana Vain. The former species differs in having salacinic acid and 
more conspicuous soralia, the latter in lacking cilia. Parmelia 
viridiflava and P. conformata Vain, are the only Amphigymnia species 
with fumarprotocetraric acid. The cilia are often sparse and ir- 
regular but definitely occur at the tips as well as in the axils of the 
lobes. It seems to be an endemic Caribbean species related to P. 
conformata. 

Additional specimens examined: 

HAITI: Montagne Noire, near Kenscoff, elev. 5500 ft., Wetmore 2761 (MSC); 
west of Forét des Pins, Wetmore 2923, 3219 (MSC), Imshaug 22846, 22869 (MSC); 
summit of Téte Etang, between Kenscoff and Furcy, Imshaug 22559 (MSC). 
DOMINICAN REPUBLIC: Gap on main ridge between Pico del Yaque and L. 
Chinguela, elev. 7000 ft., Wetmore 3701, 3706 (MSC). 

37. Parmelia xanthina (Mill. Arg.) Vain. Acta Soc. Faun. Fl. Fenn. 7, no. 7:37. 
1890. 

Parmelia chrysantha Tuck. Syn. N. A. Lich. 55. 1882. Type collection: Blue 
Ridge, Virginia, Tuckerman (FH-Tuck, holotype). Invalid name in syn. 

P. proboscidea Tayl. var. zanthina Mill. Arg. Flora 67: 616. 1884. 

P. perlata var. zanthina (Mill. Arg.) Stizenb. Bericht. St. Gall. Naturw. 
Gesell. 1888-1889:156. 1889. 

P. zanthina f. isidiosa Mill. Arg. Hedwigia 30:229. 1891. Based on P. 
proboscidea var. xanthina Miill. Arg. 

P. caperata var. madagascariacea Hue, Nouv. Arch. Mus. Paris, ser. 4, 1:181. 
1899. Type collection: Ambositra, Madagascar, Rodriguez, 1889. (P, 
holotype). 

P, madagascariacea (Hue) des Abbayes, Bull. Inst. Fr. Afr. Noire, ser. A, 
20:22. 1958. 

P. nyasensis Dodge, Ann. Mo. Bot. Gard. 46:126. 1959. Type collection: 
Mt. Nehisi, Nyasaland, Africa, Brass 16922 (NY, holotype). 


Type collection: Central Madagascar, Hildebrandt (G, holotype). 

Thallus loosely attached, 8-20 cm. in diameter, saxicolous or 
corticolous, yellowish green; lobes rotund 8-12 mm. wide, margins 
isidiate-dissected on older lobes, ciliate, cilia 1.0-2.5 mm. long; upper 
surface plane, dull, reticulately cracked with age, densely isidiate, 
especially near the margins of lobes, isidia simple to coralloid branched, 
0.05-0.08 mm. thick, 1 mm. high or more, often apically ciliate; 


276 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


lower side black and rhizinate at the center, shiny dark brown and 
dark brown in a broad zone along the margins. Apothecia and pyeni- 
dia lacking. 

Reactions: Thallus K+ more intensely yellow; medulla K—, 
O—, KC+ red, P—, atranorine, protolichesteric acid, usnic acid, 
and a KC+ unknown substance recrystallizing as clustered needles 
in G.E.; or K—, C—, KC—, P—, atranorine, protolichesteric acid, 
and usnic acid present. 

Des Abbayes (1958) has studied and attempted to typify the various 
yellow isidiate Amphigymnias. He considered P. zanthina to be C-++ 
rose (gyrophoric acid). Two other morphologically identical strains 
were also recognized: P. aberrans (Vain.) des Abb. (C—) and P. mada- 
gascariacea (Hue) des Abb. (KC-+ red). Although des Abbayes was 
not able to find a satisfactory type for P. proboscidea var. xanthina, | 
received from Geneva a well-developed specimen from Madagascar 
which is apparently Miiller’s holotype. It is C— and contains proto- 
lichesteric acid. Furthermore, the holotype of P. xanthina f. aberrans 
Vain. actually contains gyrophoric acid, contrary to Vainio’s report of 
a C— reaction (see discussion of P. aberrans). Hence the name 
P. aberrans should be applied to the tropical America population that 
reacts C+ rose. Typical P. zanthina with protolichesteric acid alone 
and P. madagascariacea (including P. chrysantha and P. nyasensis) 
with a KC+ unknown have identical ranges in North America, 
Africa, and Asia (figs. 10, 11). By our definition, they should be 
considered as chemical strains of one species, P. zanthina. In North 
America, P. xanthina is common on rock outcrops and on oaks in 
open swamps in the southeastern United States. 

Additional specimens examined: 

Protolichesteric acid present: 

U.S.: AbvaBamMa: Oak Mountain State Park, Shelby Co., Hale 7143 (US); 
Georaia: Near Trenton, Dade Co., Hale 7387 (US); Vogel State Park, Union 
Co., Hale 7355 (US); Cloudland Canyon, Walker Co., Anderson 12969 (DUKK, 
US). MEXICO: Cutapas: 2 km. north of highway 190 on road to Puebla Nueva, 
Hale 20163 (US); Km. 956 on highway 190, west of Ocozocoautla, Hale 20594 
(US). NICARAGUA: JrnotreGa: southwest of Jinotega, Standley 10137 (F). 

SOUTHERN RHODESIA: Zimbabwe Hill, Héeg s.n. (TRH). MADA- 
GASCAR: Road between Ivato and Ambatofinandrahana, des Abbayes, Lich. 
Madagasc. Borb. Sel. Exs. 19 (M, REN, UPS, US). 

Unknown KC-+ substance and protolichesteric acid present: 


U.S.: Viramnta: Natural Bridge, Rockbridge Co., Hermann 15087 (US); Norri 
CaRoLINA: 8 mi. south of Raeford, Hoke Co., Culberson 7600 (DUKE); Dardin, 
Martin Co., Culberson 6684 (DUKE); 1 mi. east of Wise Fork, Jones Co., Culberson 
4926 (DUKE); Soutn Caroiina: 12 mi. southwest of Andrews, Georgetown Co., 
Hale 16543 (US); King Mountain State Park, Cherokee Co., Hale 7707 (US); 
TENNESSEE: Cliff Springs, Overton Co., Phillips 307 (US); Grorata: Near 
Claxton, Evans Co., Hale 7451 (US); Neels Gap, Lumpkin Co., Hale 7538 (US); 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 277 


10 mi. northeast of Midway, Bryan Co., Hale 16799 (US); Unicoi State Park, 
White Co., Culberson 7291 (DUKE); AtaBama; Cheaha State Park, Clay Co., 
McCullough 571 (US); 4 mi. south of Alexander City, Tallapoosa Co., McCullough 
1042 (US); Fuoripa: 5 mi. east of Greenville, Madison Co., Hale 17613 (US); 
near Pensacola, Escambia Co., Hale 7991 (US); Dog Lake Recreation Area, 
Leon Co., Hale 16713 (US). MEXICO: Oaxaca: 53 km. northwest of Oaxaca, 
Hale 20799 (US); Km. 686 on highway 190, northwest of Tehuantepec, Hale 
20646 (DUKE, REN, TNS, US); Cutapas: El Zapotal, Tuxtla Gutiérrez, Hale 
19975 (US). 

MADAGASCAR: Road from Antsirabe to Fianarantsoa, Km. 202, des Abbayes, 
Lich. Madagasc. Borb. Sel. Exs. 20 (M, REN, UPS, US). 

INDIA: Coonoor, Nilgheries Mountains, Gray s.n. (P). 

Additional localities cited by Hale (1959b) from Kentucky, Virginia, North 
Carolina, South Carolina, Tennessee, Georgia, Alabama, Florida, Arkansas, 
Costa Rica, and Madagascar are not repeated here. 


Series Emaculatae, ser. nov. 


Thallus cinereo-albicans, acido usnico destitutus, lobis ciliatis, 
superne emaculatus, subtus margine nigricans, castaneus, rarius 
albo-variegatus vel albicans. 

Type species: Parmelia crinita Ach. 

This large series of 41 species includes 10 cosmopolitan species, 12 
endemic to Africa, and 10 endemic to the Americas. It includes some 
of the more familiar temperate species such as P. arnoldii, P. crinita, 
P. perlata, and P. stuppea. Chemical and morphological diversity is 
very great, but no further subdivision of the series seems possible at 
this time. 


38. Parmelia abnuens Nyl. Flora 68:610. 1885. PLATE 7 


Parmelia callitricha Zahlbr. Denkschr. Akad. Wiss. Math. Naturw. Wien 
83:178. 1909. Type collection: Between SAo Amaro and Barra Mansa, 
and near Palmeiras de Sdio Lourenzo, Sio Paulo, Brazil [one collection] 
Schiffner, June 1901 (W, holotype). 


Type collection: Brazil, Glaziou 1835 (H, Nyl, herb. no. 35290, 
lectotype). 

Thallus loosely attached on bark, 4-8 cm. across, mineral gray; 
lobes rotund, 8-15 mm. wide, becoming quite laciniate, laciniae 
12-4 mm., often appearing canaliculate below, margins densely 
ciliate, cilia 3-5 mm. long; upper surface shiny, distinctly to faintly 
maculate, pitted-rugose with age; lower side black and _ sparsely 
rhizinate at the center, dark brown to mottled tan and naked at the 
margins. Apothecia common, stalked, 5-10 mm. in diameter, am- 
phithecium rugose, strongly maculate, disc imperforate; hymenium 
90-110 w high; spores 12-18%24-35 yu, episporium 2.5-3.0 yw thick; 
pycnidia present, conidia 6-8 yw long (after Nylander). 

Reactions: Thallus K+ yellow; medulla K—, C+ deep orange red, 
KC+ deep orange red, P—, atranorine and olivetoric acid present 
(proved with chromatography). 


278 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Parmelia abnuens is superficially near P. melanothriz (Mont.) Vain. 
and related nonsorediate, nonisidiate species. It is separated by the 
deep C+ red reaction. The only close relative is P. amboimensis 
Dodge, an African species with small spores and an even more laciniate 
thallus. The differences in spore size between the types of P. abnuens 
(20-24 u) and P. callitricha Zablbr. (30-36 uw) are much larger than we 
expect to find in Parmelias, but there seems to be no justification for 
recognizing two species on spore size alone. 

The second syntype on which Nylander based the name P. abnuens 
is a Lorentz collection from Uruguay. This specimen in Nylander’s 
herbarium is P. argentina Kremplh. 

39. Parmelia amboimensis Dodge, Ann. Mo. Bot. Gard. 46:158. 1959. 
PuatEe 7 

Type collection: Amboim, Capir, Cuanza Sul, Angola, Gossweiler 
9993 (K, holotype; US, isotype). 

Thallus large, loosely attached, more or less suberect, 10-15 cm. 
across, mineral gray; lobes 10-12 mm. wide, irregularly branched and 
richly laciniate, laciniae 1-210 mm., appearing canaliculate below, 
margins sparsely ciliate, cilia 1-2 mm. long; upper surface smooth, dull 
to shiny; lower side black and sparsely rhizinate, dark brown to mot- 
tled tan and naked at the margins. Apothecia stalked, 8-15 mm. in 
diameter, amphithecium rugose, faintly maculate, disc imperforate 
hymenium 70-75 u high; spores 7-8 10-13 u, episporium 1 yg thick; 
pycnidia present, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C+ deep orange red, 
KC-+ deep orange red, P—, atranorine and olivetoric acid present 
(proved with chromatography). 

This species has a decidedly everniiform appearance with broadly 
caniculate secondary lobes. The main lobes, however, are wide and 
the apothecia are stalked and rugose, unlike any species in Pseude- 
verma Zopf. The closest relative is the Brazilian species P. abnuens 
Nyl., which differs in having less laciniate lobes and larger spores. 

Additional specimens examined: 

ANGOLA: Cuanza Suu: Carloaongo-Cuvo, Capir, Amboim, Gossweiler 10008, 
10038 (US). 


40. Parmelia appendiculata Fée, Ess. Crypt., Suppl. 118, pl. 38, fig. 3. 1837. 

Parmelia cristata Nyl. Flora 52:291. 1869. Type collection: Caripe, 
Venezuela, Moritz (H, holotype, Nyl. herb. no. 35293). 

P. merrillit Lynge, Ark. Bot. 13, no. 13:79. 1914. Type collection: Coxipé, 
Igreja, near Cuyabé, Mato Grosso, Brazil, Malme 2198B (8S, holotype). 
Not P. merrillit Vain. (1909). 

P. cornuta var. crocea Lynge, Ark. Bot. 13, no. 13:78. 1914. Type collection: 
Santa Anna da Chapada, Mato Grosso, Brazil, Malme 2477 bis (S, holotype; 
W, isotype). (Unidentified anthraquinone pigment present in addition 
to K— yellow pigments.) 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 279 


P. lyngeana Zahlbr. Cat. Lich. Univ. 6:243. 1929. Based on P. merrillit 
Lynge. 

P. raced (Lynge) Gyel. Repert Sp. Nov. Fedde 29:287. 1931. [Non P. 
crocea (Ach.) Spreng. =Solorina.] 

Type collection: Bourbon, s.c. (G, isotype). 

Thallus loosely adnate to bark, 3-6 cm. in diameter, buff mineral 
gray in herbarium, lobes rotund, 5-8 mm. wide, laciniate, laciniae 1 
mm. wide, 3-8 mm. long, margins of lobes sparsely ciliate, cilia 1 mm. 
long; upper surface dull, reticulately cracked with age, soredia and 
isidia lacking; medulla white or in part pale to deep yellow-orange; 
lower side black and sparsely rhizinate, dark brown, naked and rugu- 
lose in a broad zone along the margins. Apothecia short-stalked, 
exciple dentate to dentate-laciniate, sparsely ciliate, amphithecium 
maculate, disc imperforate; hymenium 100-130 yw high; spores 
14-16 X 26-32 yw, the episporium 2.5-3.0 » thick; pycnidia present, 
conidia not seen. 

Reactions: Thallus K+ yellow; medulla K— or K+ yellowish, 
C+ yellowish, KC+ yellowish, P—, atranorine, an unidentified 
pigment, and unknown substances present. 

Although P. appendiculata was described some 125 years ago, the 
name has only rarely appeared in the literature. The type was care- 
fully illustrated by Fée but the most critical character, the chemistry, 
was not known. Fortunately an isotype found at Geneva has enabled 
us to characterize the species accurately for the first time. The 
medulla is white to pale yellow and distinctly yellow under the 
apothecia, with a K— pigment identical with that in P. araucarvarum 
Zablbr. and others. The marginal laciniae are well developed but 
cilia are sparse and short. This species is extremely rare and known 
only from the type collections cited above. 

Des Abbayes (1956) compared P. appendiculata and disparilis Nyl. 
on the basis of Fée’s illustration and on his own collections from 
Madagascar. These two species are indeed similar externally, but 
P. disparilis contains protocetraric acid, lacks any pigment, and has 
smaller spores. 

41. Parmelia arnoldii Du Rietz, Nyt Mag. Naturv. 62:80. 1924. 

Type collection: Munich, Germany, Arnold Exs. 136b (UPS, 
lectotype; DUKE, FH, H, LD, isotypes). 

Thallus loosely attached to bark, 8-20 cm. in diameter, light mineral 
gray; lobes 8-15 mm. wide, with dentate marginal laciniae, 2-4 mm. 
wide, 5-10 mm. long, crowded toward the center; upper surface 
shiny, faintly maculate, sorediate, soredia developing mostly on the 
laciniae, submarginal, eventually coalescing into extensive soralia, 
tips of sorediate lobes usually revolute, margins ciliate, cilia rather 
sparse, 1.0-2.5 mm. long; lower side black and rhizinate at the center, 


%29—-018—_65——_8 


280 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


brown and naked in a broad zone at the margin. Apothecia very rare, 
hymenium 80 yw high; spores 9.5-1215-22 yw; conidia 10.5-11.5 p 
long (after Hillmann, 1934). 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ reddish 
orange, P—, atranorine and alectoronic acid present; rhodophyscin 
also present in the lower medulla in some specimens, reacting K+ 
purple. 

Parmelia arnoldii is a temperate species of both Northern and 
Southern Hemispheres (fig. 16) (cf. Degelius, 1935). It is distinguished 
by the sublaminal soredia and the presence of alectoronic acid, often 
accompanied by rhodophyscin. Parmelia margaritata Hue, which 
differs only in having salacinic acid and lacking rhodophyscin, has a 
restricted distribution in North America (fig. 9). 

The typification of P. arnoldii is somewhat involved. Du Rietz 
proposed it as a new name for “P. nilgherrensis Nyl.” (1874) non P. 
nilgherrensis Nyl. (1869) and a number of other misidentified litera- 
ture reports by Hue and Lynge. Parmelia nilgherrensis Nyl. (1874) isa 
specimen (Arnold Lich. Exs. 136b) from Germany which Nylander 
had misidentified. The original P. nilgherrensis (1869) is an Asian 
species. The most appropriate lectotype for P. arnoldii is Arnold 
136b, a typically developed specimen widely distributed in the larger 
herbaria. 

Additional specimens examined: 

CANADA: Brrrisn Conumsra: Cedar Hill, Vancouver Island, Melburn 90 (US); 
Spring Island, Vancouver Island, Szczawinski 418 (US). U.S.: Maine: Mt. 
Katahdin, Degelius sn. (US); West Virainra: 5 mi. northeast of Thornwood, 
Pocahontas Co., Hale 12362 (US) ; Spruce Knob, Pendleton Co., Hale 15460 (US); 
MicuiGaan: Isle Royale, Keweenaw Co., Wetmore 5044 (LD); Minnesora: 2 mi. 
northwest of Illgen City, Lake Co., Spross s.n. (US); Virainta: Hawksbill Moun- 
tain, Madison Co., Hale 18937 (US); Whitetop Mountain, Washington Co., 
Hale 18739 (US); TENNESSEE: Roan Mountain, Carter Co., Phillips 374 (US); 
Kentucky: Rockcastle River at Route 192, Pulaski Co., Reed 56753 (Reed 
Herbarium); Wasuineron: Hoh River, Jefferson Co., Brown 160 (US); Long- 
beach, Pacific Co., Howard 4851 (WIS); OrEGon: Road between Bridal Veil Falls 
and Larch Mountain, Multnomah Co., Hale 21587 (US); Canirornra: Pilarcitos 
Creek Canyon, San Mateo Co., Thomson 4761 (WIS), Cain 26388 (TRT, US), 
Herre 825 (MO); Near Saratoga, Santa Clara Co., Cain 26394a (TRT, US); 
Mt. Tamalpais, Marin Co., Morton & Howell 11905 (US). MEXICO: Mexico: 
Popocatepetl, Cain 27595 (TRT, US); 10 km. southwest of Cahuacin, Madrigal 
1426 (US); Pursia: Near Llano Grande, Hale 19302 (DUKE, MSC, REN, §, 
TNS, US); Oaxaca: Cerro San Felipe, Hale 20708 (US). GUATEMALA: 
CHIMALTENANGO: Region of Santa Elena, Standley 58718 (MO), 

HAITI: Near summit of Morne Macaya, Wetmore 3352 (MSC), 3358 (MSC, 
US); summit of Téte Etang, Imshaug 22614 (MSC, US). DOMINICAN RE- 
PUBLIC: Summit of Alto de la Bandera, Wetmore 3483 (MSC, US); southwest 
slope of La Rucilla, Maciso de los Yaques, Imshaug 23614 (MSC, US); La Nevera, 
Cordillera Central, Wetmore 3450 (MSC, US), Imshaug 23342 (MSC, US). 
GUADELOUPE: Palmer s.n. (FH). 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 281 


CHILE: Vauraraiso: Alto del Puerto, Valparaiso, Santesson 2761, 2798, 
2878 (S); Concrrci6n: Coronel, Santesson 2379 (S); Vaupivia: Malihue, Rio 
San Perdo, Santesson 3348 (S); Corral, Santesson 2329 (S); Enco, Lago Riiiihue, 
Santesson 7595 (S); Aysin: Puerto Aysén, Santesson 6934 (8). 

FRANCE: Lorges, des Abbayes, Lich. Armor. Spect. Exs. 32 (LD). GER- 
MANY: Munich, Arnold, Lich. Monac. 492 (MICH); Bleckenau, Schwaben, 
Férster, Lich. Alp. 194 (LD, M, US). POLAND: Puszeza Bukowa, Bieszezady 
Zachodnie, Glance in Lich. Polonica 173 (UPS). NORWAY: Rogaland, Havaas, 
Lich. Norv. Occ. Exs. 84 (LD). 


42. Parmelia bangii Vain. Bot. Tidsskr. 29:104. 1909. 

Type collection: La Paz, Bolivia, Bang 13 (H, holotype, Nyl. 
herb. no. 35500). 

Thallus loosely attached to bark, 8-15 cm. in diameter, turning 
buff in the herbarium; lobes rotund, irregular, 5-8 mm. wide, margins 
entire, very sparsely ciliate, cilia 1.0-1.5 mm. long, mostly along the 
lateral edges; upper surface dull, continuous or reticulately cracked 
with age, cortex distintegrating in large areas near the margins and 
developing into sorediate pustules; lower side black and moder- 
ately rhizinate, light brown, shiny, and naked along the margins. 
Apothecia adnate, amphithecium pustulate, disc imperforate; hyme- 
nium 70-80 » high; spores 11-16 26-36 y, episporium 3-4 yu thick; 
pycnidia not seen. 

Reactions: Thallus K+ yellow; medulla K+ yellow, C—, KC—, 
P-+ pale orange red, atranorine and stictic acid present. 

This is a rare species apparently endemic to tropical South America. 
The spore size and chemistry place it close to P. perlata (Huds.) 
Ach., but the development of extensive laminal sorediate pustules is 
unique. 

Additional specimen examined: 

COLOMBIA: Chapinero, Ariste-Joseph A355a (US). 

43. Parmelia blanchetii Hue, Nouv. Arch. Mus. Paris, ser. 4, 1:199. 1899. 
Type collection: Bahia, Brazil, Blanchet (P, holotype; BM, isotype). 
Thallus loosely adnate on rock, 8-10 cm. in diameter, light mineral 

eray; lobes rotund to irregular, 5-8 mm. wide, margins crenate to 
dentate-lobulate, especially toward the center, more or less black- 
rimmed, ciliate, cilia 1.5-2.5 mm. long; upper surface plane, shiny, 
reticulately cracked with age, isidia and soredia lacking; lower side 
black and sparsely rhizinate, dark brown and naked in a broad zone 
along the margins. Apothecia stalked, 4-6 mm. in diameter, amphi- 
thecium slightly rugose, disc imperforate; hymenium 50-60 y high; 
spores 5-6<12-15 yu, episporium 1 y; pycnidia present, conidia not 
seen. 

Reactions: Thallus K++ yellow; medulla K—, KC-+ reddish orange, 
fading rapidly, P+ orange red, atranorine and protocetraric acid 
present. 


282 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


This unique species is known from only two collections. It is 
apparently saxicolous. Externally it is rather close to P. eciliata 
(Nyl.) Nyl., a corticolous species which has stictic acid and large 
spores. 

Additional specimen examined: 


COLOMBIA: Vaurrs: Mesa de Yambi, Rfo Karur, Schultes 19125 (FH, 
US). 

44, Parmelia breviciliata Hale, sp. nov. 

Parmelia latissima f. microspora Lynge, Ark. Bot. 13, no. 13:45. 1914. 
Type collection: Near Bocca da Serra, Serra da Chapada, Mato Grosso, 
Brazil, Malme 2244* (S, lectotype). 

Thallus laxe adnatus, saxicola, rigidulus, cinereo-albicans, 5-15 cm. 
latus, lobis rotundatis, 8-12 mm. latis, margine integris vel acetate 
irregulariter lobulatis, modice ciliatis, ciliis brevibus, 1-2 mm. longis, 
superne planus, opacus, continuus vel aetate plus minusve rimosus, 
medulla alba, subtus niger, sparse rhizinosus, ambitu castaneus, 
nudus. Apothecia 4-10 mm. diametro, primo urceolata, substipitata, 
disco imperforato, amphithecio opaco, excipulo integro; hymenium 
45-55 uw altum; sporae 6-10X12-17 wu, episporio 1.0-1.5 y lato. 
Thallus K-+ flavescens; medulla K—, C—, KC+ rosea, P—, atra- 
norinum et acidum alectoronicum continens. 

Type in Lund University, collected 4 miles southeast of Pilgrims 
Rest, Transvaal, Union of South Africa, Oct. 10, 1953, by Ove Alm- 
born (no. 7800; isotype in US). 

Parmelia breviciliata is most common on rocks in South Africa, 
although there are disjunct localities in South America and Thailand 
(fig. 23). It resembles most closely P. wainii A. L. Sm., which differs 
chiefly in having intermediate spores, strongly stalked maculate 
apothecia, a ciliate exciple, and corticolous habit. Parmelia soyauzii 
Mill. Arg., another saxicolous African lichen, is superficially very 
similar but lacks cilia and contains lecanoric acid. A second Malme 
collection (2244) from Mato Grosso was tentively identified by Lynge 
as P. proboscidea f. incrassata Lynge; this specimen (S) is P. breviciliata. 

Additional specimens examined: 


BRAZIL: Maro Grosso: Near Bocca de Serra, Serra da Chapada, Malme 
s.n., 2244 (S). 

IVORY COAST: Mont Tonkoui, cercle of Man, Santesson 10634 (UPS). 
CAMEROONS: Bafut-Ngem Forest Reserve, Brunt 773 (K, US). UGANDA: 
Entebbe, Proctors.n. (K). ANGOLA: Huma: North of 84-da-Bandeira, Degelius 
s.n. (DEGEL, US). SOUTHERN RHODESIA: Zimbabwe, Héeg s.n. (TRH, 
US). UNION OF SOUTH AFRICA: Transvaau: 6 mi. north of Houtbosch, 
Almborn 6720 (US); Barberton, Héeg s.n. (TRH); Fairy Glen, Pretoria, Bottomly 
719 (PRE); Nara: Table Mountain, Pietermaritzburg, Almborn 8591, 8592 
(LD). MADAGASCAR: Ambositra, Forsyth 629 (BM); road from Tananarive 
to Ankazobe, Km. 48, des Abbayes, Lich. Madagasc. Borb. Sel. Exs. 15 (M, REN, 
UPS, US). THAILAND: Kao Krading, Kerr L23 (K). 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 283 


45. Parmelia corniculans Nyl. Flora 68:607. 1885. 

Type collection: Java, Horsfield (H, lectotype, Nyl. herb. no. 
35308). 

Thallus loosely attached, 7-12 cm. broad, light mineral gray; lobes 
rotund, 10-13 mm. wide, margins smooth to more or less crenate, more 
or less dentate-laciniate with age, ciliate, cilia to 2 mm. long; upper 
surface dull, plane, soredia and isidia lacking; lower side black and 
sparsely rhizinate, dark brown to brown and naked in a broad zone 
along the margins. Apothecia 4-8 mm. in diameter, stalked, exciple 
more or less dentate, short-ciliate, amphithecium rugulose, maculate, 
disc perforate; hymenium 80-85 yu; spores 16-18 30-35 yu, episporium 
3 p thick; pycnidia numerous, conidia 5-6 u long. 

Reactions: Thallus K+ yellow; medulla K—, KC-+ reddish orange, 
C—, P—, atranorine and alectoronic acid (mixed with a-collatolic 
acid) present; rhodophyscin often present in the lower medulla, K+ 
purple. 

This species seems to be restricted to Malaysia and is therefore 
quite separated geographically from comparable species with alec- 
tornic acid in Africa and America. The only really closely related 
species is P. maclayana Mill. Arg., which lacks such conspicuous 
laciniae and has small spores. Parmelia merrillii Vain., an Asian 
species, has similar laciniae and large spores, but differs in containing 
protocetraric acid and having imperforate apothecia. 

Additional specimens examined: 

PHILIPPINES: Luzon: Mt. Tonglon, Ramos 5493 (US); Baguio, Benguet, 
Elmer 8563 (US). JAVA: Mt. Gedeh, Groenhart 8812 (BO). LAOS: Hase 
Plantation, Tsuyama 13 (TNS). 

46. Parmelia cornuta Lynge, Ark. Bot. 13, no. 13:76, pl. 2, fig. 5. 1914. 

Type collection: Santa Anna da Chapada, Mato Grosso, Brazil, 
G. A. Malme 2477 (S, holotype). 

Thallus loosely attached, 10-15 cm. across, buff mineral gray in 
herbarium; lobes rotund, 7-10 mm. wide, margins entire, becoming 
dentate-laciniate towards the center, ciliate, cilia sparse, 2-3 mm. 
long; upper surface dull, more or less rugulose, cracked with age, 
isidia and soredia lacking; medulla lemon yellow; lower side black, 
sparsely rhizinate in the center, brown to tan and naked in a broad 
zone at the margins. Apothecia substipitate, exciple dentate and 
ciliate, amphithecium maculate, disc imperforate; hymenium 110-130 
u» high; spores 13-15 26-32 yu, episporium 3 » thick; pycnidia present, 
conidia not seen. 

Reactions: Thallus K+ yellow; medulla lemon yellow, K—, C—, 
KC—, P-—, atranorine and vulpinic acid present. 

Parmelia cornuta is superficially very near to P. appendiculata 
Fée except for the presence of vulpinic acid. Both species have 


284 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


dentate-laciniate apothecia, sparse cilia, and large spores. Parmelia 
sulphurata Nees & Flot., an isidiate species, is the only other Parmelia 
with vulpinic acid. Parmelia cornuta is still known only from the 
Mato Grosso region. 

Additional specimen examined: 


BRAZIL: Mato Grosso: Buriti, Malme s.n., 2243B (8). 


47, Parmelia crinita Ach. Syn. Lich. 196. 1814. PLATE 1 

Parmelia proboscidea Tayl. in Mack. Fl. Hibern. 143. 1836. Type collec- 
tion: Near Dunkerron, Ireland (FH-Tayl, holotype; isotype in BM). 

P. perlata var. ciliata f. dissectula Nyl. in Leight. Lich. Fl. Gr. Brit. 120. 
1879. Type collection: Killery Bay, Connemara, Ireland, Larbalestier s.n. 

(BM, holotype). 

P. chlorocarpa Mill. Arg. Flora 63:265. 1880. Type collection: Caracas, 
Venezuela, Ernst in 1879 (G, holotype). 

P. schweinfurthii Mill. Arg. Proc. Roy. Soc. Edinburgh 2:459. 1882. 
Type collection: Socotra, Africa, Schweinfurth (G, holotype). 

P. proboscidea var. dissectula (Nyl.) Miill. Arg. Flora 67:616. 1884. 

P. catharinensis Mill. Arg. f. isidiosa Mill. Arg. Flora 74:378. 1891. 
Type collection: Southern Brazil, Leyland (K, holotype; G, isotype). 

P. perlata var. dissectula (Nyl.) Mill. Arg. Bull. Soc. Bot. Belg. 32:127. 1893. 

P. pilosella Hue, Journ. de Bot. 12:247. 1898. Type collection: Coatloch 
Forest, Finistére, France, Picquenard in 1898 (P, lectotype; BM, isotype). 

P. pilosella f. excrescens Hue, Journ. de Bot. 12:249. 1898. Type collection: 
Munich, Germany, Arnold Lich. Exs. 655a (P, lectotype; BM, isotype). 

P. crinita f. pilosella (Hue) Merrill, Bryol. 11:95. 1908. Based on P. 
pilosella Hue. 

(?)P. tuckermanii Du Rietz, Nyt Mag. Naturvid. 62:70. 1924. Type col- 
lection: Wright, Lich. Cubae 69 (UPS, holotype; FH, M, US, YU, iso- 
types). 

P. pseudocatharinensis Gyel. Repert. Sp. Nov. Fedde 29:289. 1931. Based 
on P. catharinensis Miill. Arg. f. isidiosa Mill. Arg. 


Type collection: North America, Muhlenberg (H, holotype). 

Thallus adnate to loosely attached, 8-25 cm. in diameter, mineral 
gray; lobes rotund to irregularly dissected, 6-12 mm. wide, margins 
crenate, often isidiate and dissected on older lobes, ciliate, cilia 1-2 
mm. long; upper surface dull, plane, moderately to densely isidiate, 
isidia simple to coralloid-branched, 0.05-0.1 mm. thick, to 2 mm. 
high, often apically ciliate; lower side black and sparsely rhizinate, 
dark brown to tan, more or less rugose, shiny, and naked in a broad 
to rather narrow zone at the margins. Apothecia rare, 3-8 mm. in 
diameter, substipitate, amphithecium isidiate, disc imperforate; 
hymenium 65-80 y» high; spores 12-16%25-30 yu, episporium 3-4 u 
thick; pycnidia rare, conidia 3-4 u long (after Asahina). 

Reactions: Thallus K+ yellow; medulla K+ yellow, C—, KC—, 
P+ pale orange yellow, atranorine and stictic acid present. 

Parmelia crinita is @ very common cosmopolitan species known to 
all lichenologists. It has a remarkable range of climatic adaptability, 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 285 


from purely tropical areas to temperate and even subboreal forests 
(fig. 17) (cf. Degelius, 1935). The isidia vary widely from simple and 
sparse to densely coralloid-branched and ciliate. The lobes are rather 
small and have a relatively narrow, shiny, often rugose bare zone 
below. It is the only isidiate species in the subgenus with stictic 
acid. Parmelia subcrinita Nyl., which contains salacinic acid, is in 
general a more robust plant with a broad bare zone below. Parmelia 
mellissit Dodge in a few cases has coralloid isidia that could be confused 
with those of P. crinita but it is K— (alectoronic acid) and has smaller 
spores. 

Parmelia internexa Nyl., which is classified in the subgenus Parmelia, 
differs chiefly in size. It has adnate lobes at most 4 mm. wide and a 
narrow papillate zone below. 

Parmelia tuckermanii Du Rietz was described from a Wright 
Cuban collection that contains spores significantly smaller (about 18 u 
long) than in any specimens of P. crinita that I have examined. Since 
spore size is the only difference between it and P. crinita, sterile speci- 
mens must be referred to P. erinita. I believe the smaller spores are 
juvenile or otherwise abnormal. 

Additional representative specimens examined: 


CANADA: Nova Scotra: Hunt’s Point, Denison (US). Ontario: Hibbard 
Bay, Algoma Distr., Cain 26882, 26884 (TRT, US); Brighton, Macoun 67, 175 
(US). U.S.: Maine: Mt. Desert Island, Hancock Co., Davis 68 (US); Rockland, 
Knox Co., Merrill, Lich. Exs. 124 (FLAS, US); Connecricur: Aton Forest, 
Litchfield Co., Hale 71 (US, YU); New Yorr: Jamesville, Onondaga Co., Cook 
s.n. (US); MaryLanp: 6 mi. west of Town Hill, Alleghany Co., Hale 14503 (US). 
WEst VIRGINIA: 4 mi. east of Baker, Hardy Co., Hale 15024 (US); Bluestone 
Dam, Summers Co., Hale 11707 (US); Jaeger, McDowell Co., Hale 10832 (US); 
Ouro: Junction of routes 374 and 56, Hocking Co., Hale 13299 (US); Kenruckyr: 
Near Forest Cottage, Cumberland Co., Hale 13714 (US); Wisconsin: Upper 
Brule River, Douglas Co., Thomson s.n. (WIS); Micniaan: Lower Tahquamenon 
Falls, Chippewa Co., Imshaug 19906 (MSC, US); Iron Bridge, Cheboygan Co., 
Robinson s.n. (US); Minnesota: Grand Portage Island, Cook Co., Fink 114 
(US); Duluth, St. Louis Co., Kimball s.n. (US); Iowa: Fayette Co., Fink s.n. 
(US); Missourr: Near Ava, Douglas Co., Hale 4327 (US); Laquey, Pulaski Co., 
Hale 4079 (US). Virernta: Mountain Lake Biological Station, Giles Co., Hale 
18414 (US); 2 mi. east of Buena Vista, Rockbridge Co., Hale 18345 (US); Brown- 
town Valley Overlook, Skyline Drive, Rappahannock Co., Hale 14732 (US); 
Doughat State Park, Bath Co., Hale 12401 (US); Norru Carouina: 2 mi. south 
of West Jefferson, Ashe Co., Culberson 6306 (DUKE); Cleveland, Rowan Co., 
Culberson 6132 (DUKE); Gastonia, Gastonia Co., Hale 8144 (US); Forney Ridge, 
Great Smoky Mountains, Degelius s.n. (US); Sourn Caroiina: Near Greenville, 
Greenville Co., Hale 7762 (US); 5 mi. east of Walterboro, Colleton Co., Hale 
16502 (US); TENNESSEE: 2 mi. north of Carters Gap, Carter Co., Hale 18003 
(US); Cliff Springs, Overton Co., Phillips 301 (US); Grorera: Chatsworth, 
Murry Co., Hale 7416 (US); Neels Gap, Lumpkin Co., Hale 7559 (US); ALABAMA: 
Oak Mountain State Park, Shelby Co., Hale 7156 (US); 4 mi. southeast of Troy, 
Pocosin area, McCullough 472, 476 (US); Fuorrpa: 8 mi. east of Ocala, Marion 


286 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Co., Hale 17033 (US); Oleno State Park, Alachua Co., Hale 16739 (US); High- 
lands Hammock State Park, Highlands Co., Hale 16937 (US); Arkansas: Nogo, 
Pope Co., Hale 2975 (US); Dardanelle, Yell Co., Hale, Lich. Amer. Exs. 16 (BM, 
C, CAN, FH, FU, ID, ILL, L, LD, LISU, M, MO, NY, 8, TENN, TRT, UC, 
US, W, WIS); Oxtanoma: South of Albion, Pushmataha Co., Hale 4976 (US); 
Texas: Near Woodville, Tyler Co., Hale 5218 (US); Jasper, Jasper Co., Hale 
5411 (US); Arizona: Santa Rita Mountains, Santa Cruz Co., Darrow 4394 
(Darrow Herbarium); MEXICO: Veracruz: 33 km. northeast of Perote, Hale 
20871 (US), 20892 (COLO, US); Oaxaca: Cerro San Felipe, Hale 20787 (LISU, 
US); Curapas: 50 km. west of Tuxtla Gutiérrez on highway 190, Hale 19907 
(DUKE, 8, TNS, US); 14 km. west of San Cristébal, Hale 20566 (US). GUA- 
TEMALA: QuerzaLreNANGo: Between Zunil and Cantel, Standley 83905 (MO). 
HONDURAS: El Paraiso, Giiinope, Valerio 1816 (MO). NICARAGUA: 
JInoTEGA: West of Jinotega, Standley 9766 (F); Masaya: Region of Las Nubes, 
Standley 8749 (F). COSTA RICA: Guanacaste: Aguilares, Standley 46355 (US); 
Cartaao: Along the Rio Reventado, Standley 49601 (US); 1 mi. south of Cartago, 
Danilson 17 (MICH); AuasuELA: San Pedro de San Ramén, Brenes 457 (UPS). 

CUBA: Las Vituas: Trinidad Mountains, Imshaug 24599 (MSC); OrrEnTE: 
1] Gato, Loma del Gato, Sierra Maestra, Imshaug 24819 (MSC, US), 24926 
(MSC, US), 24768 (MSC), Hioram 8467 (FH), Leon 10613 (NY); Loma San 
Juan, Guanténamo, Hioram 10268 p.p. (FH); summit of Pico Turquino, Imshaug 
25105 (MSC). HAITI: Near summit of Morne Macaya, Imshaug 23252 (MSC, 
US); summit of Téte Etang, Imshaug 22587 (MSC). DOMINICAN REPUBLIC: 
Along road at Los Amaceyes, Wetmore 3399 (MSC); near Cerrazos, on ridge from 
La Cumbre to Santiago, Wetmore 3856 (MSC, US), 3862 (MSC); summit of 
Pico de Isabel de Torres, near Puerto Plata, Imshaug 23939 (MSC, US). JA- 
MAICA: Coopers Hill in the Red Hills, Imshaug 13721, 138729, 13744, 14125, 
14141 (MSC); Dolphin Head, Hanover, Imshaug 15647 (MSC); Blue Mountain 
Peak Trail, St. Thomas, Jmshaug 12951 (MSC, US); Hollymount, Mt. Diablo, 
Imshaug 14190 (MSC, US), 14192, 14212 (MSC); Newcastle, Cushman 40 (FH); 
without locality, Eggers 3779 (C). PUERTO RICO: Mt. Torrecella, Sintenis 32 
(G). WIRGIN ISLANDS: Sr. Tuomas: Signal Hill, Eggers s.n. (S); Crown, 
Britton 1441 p.p. (NY). 

VENEZUELA: Méripa: Rio Chama Valley, Mérida, Dennis 2335 (K). 
COLOMBIA: Cunpinamarca: Near Zipaquird, Schultes 11441 (FH, US); 
Touma: Between Cajamarca and summit of Divide, Killip 34678b (US). 
CHILE: Cavtin: Llainia, Gunekel 19453 (MO); Vaupivra: Lago Riifiihue, 
Santesson 3510, 3732, 7146 (S); Como: Petrohué, Hosseus 187 (M); Isla Chiloé, 
Peninsula Lacui, Santesson 4092 (S), 4116 (S, US); Juan FERNANDEZ: Reed s.n. 
(BM), Hassler s.n. (K). 

SCOTLAND: Dungallan Parks, Argyll, Lindahl 176 (UPS). ENGLAND: 
Sussex, Borrer s.n. (US). NORWAY: Skaras, Regefjord, Rogaland, Hasselrot 
(8). FRANCE: Forét du Cranou, Finistére, Almborn, Aug. 20, 1954 (LD, US); 
St. Thurial, Ille-et-Vilaine, Santesson 10084a (UPS, US). HUNGARY: Near 
Ronafired, Szatala, Fl. Hung. Exs. 416 (LD, US). ITALY: Mt. Mulatto, near 
Predazzo, Arnold, Lich. Exs. 1151 (US); Deserto di Santa Anna, Cogoleto, Gresino 
11445 (F). SPAIN: 2 km. northwest of Irurita, Navarra, Santesson 13291j 
(UPS). PORTUGAL: Near Pena, Serra de Sintra, Estremadura, Tavares, Lich. 
Lusit. Sel. Exs. 117 (LD, US, WIS). 

KENYA: Tinderet Forest Reserve, Kisumu-Londiani, Nyanza Prov., Maas 
Geesteranus 11162 (L). UNION OF SOUTH AFRICA: TransvaaL: 5 mi. east 
of Mokobulaan, Lydenberg, Almborn 7495 (LD); north of Louis Trichardt, Zout- 
pansberg, Almborn 6542 (LD); Natau: Eschowe, Héeg s.n. (TRH); Polela 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 287 


Forest, Polela, Almborn 9504 (LD); near Pietermaritzburg, Umgeni, Héeg s.n. 
(TRH); Cathedral Peak Area, Bergville, Almborn 8872 (LD); Cape PRovINcE: 
Tzitzikama Mountains, Maas Geesteranus 6624 (L); Fern Kloof, Albany, Almborn 
10729 (LD); Paarl Rock, Paarl, Almborn 5422 (LD); east slope of Table Mountain, 
Wynberg, Almborn 11311 (LD); Blauwkrantz, Humansdorp, Héeg s.n. (TRH). 

CHINA: Fukien Prov., Sowerby s.n. (US). JAPAN: Ohito, Prov. Idu, 
Kurokawa 58025 (TNS, US). 

JAVA: Mt. Gedeh, Tjibodas, Groenhart 8550, 8764 (BO). FIJI: Mba, Viti 
Levu, Smith 4912 (US). AUSTRALIA: Ravenshoe, Duma Creek, Sherrin s.n. 
(BM). 

48. Parmelia cryptoxantha des Abbayes, Mem. Inst. Sci. Madagascar, ser. B, 
10:115. 1961. 


Type collection: 30 km. northeast of Ankazobe, Forét d’Ambohi- 
tantely, Centre Moyen, Madagascar, des Abbayes (REN, lectotype; 
US, isotype). 

Thallus 8-15 cm. in diameter, membranous, loosely attached, light 
mineral gray; lobes rotund, 8-15 mm. wide, margins crenate to short 
lobulate-dentate, conspicuously ciliate, cilia dense, 3-5 mm. long; 
upper surface plane to rugulose-pitted, cortex fragile and easily 
breaking away, becoming coarsely sorediate-pustulate in a broad 
zone along the margins; medulla very pale yellow to white; lower side 
black and sparsely rhizinate, black to brown, shiny, and naked in a 
broad zone along the margin. Apothecia substipitate, to 5 mm. in 
diameter, exciple short lobulate, sparcely ciliate, amphithecium 
rugose, maculate, disc imperforate; hymenium 75-90 y; spores 
10-12 24-28 yw, episporium 1.5-2.0 uw thick; pycnidia present, 
conidia not seen. 

Reactions: Thallus K+ yellow; medula K+ yellow, C—, P+ pale 
orange yellow, protolichesteric acid, atranorine, unknown P+ acids, 
and a pale yellow pigment present. 

The morphological characters of P. eryptoxantha are rather peculiar: 
Laminal sorediate pustules, similar to those of P. rimulosa Dodge but 
more sparsely developed, long cilia, and a membranous, fragile thallus. 
The chemical components are unknown except for atranorine and 
protolichesteric acid. The pigment is pale and easily overlooked. 

Additional specimen examined: 


NATAL: Forest south of Nkandhla, Nkandhla Distr., Almborn 8065 (LD, US). 


49. Parmelia diacidula Hale, sp. nov. PuatTE 13 


Thallus expansus, membranaceus, laxe adnatus, 10-20 cm. dia- 
metro, viridi-glaucescens, lobis rotundatis, 10-15 mm. latis, margine 
integris vel late crenatis, centrum versus plus minusve breve dis- 
sectis, ciliatis, cillis sparsis, 0.5-2.5 mm. longis, sorediatis, soraliis 
linearibus, irregularibus, sorediis granulosis, superne planus, nitidus, 
levissime albomaculatus, cortice fragili, strato corticeo superiore 


288 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


8-10 u crasso, strato gonidiali 24-26 u crasso, medulla alba, 75-90 u 
crassa, strato corticeo inferiore 13-15 u crasso, inferne niger, sparse 
rhizinosus, ambitu castaneus, late nudus. Apothecia pycnidiaque 
ignota. Thallus K-+ flavescens; medulla K—, C+ rosea, KC+ 
rubescens, P—, atranorinum, acidum gyrophoricum et acidum pro- 
tolichestericum (et caperaticum?) continens. 

Type in Lund University, collected in Boschfontein Forest, Lions 
River, Natal, Africa, Oct. 31, 1953, by Ove Almborn (no. 8679; isotype 
in Us). 

This species superficially resembles P. dilatata Vain, especially 
in the formation of soredia on irregularly dissected lobe margins. 
Cilia, however, are consistently produced even at the lobe tips and the 
thallus is more membranous. No other species of Amphigymnia 
produces gyrophoric and protolichesteric acids together, although 
this combination is known in the P. cirrhata group of the Evernii- 
formes. Parmelia diacidula is rather well collected and seems to be 
endemic to Natal. 

Additional specimens examined: 

UNION OF SOUTH AFRICA: Narau: Indumeni Forest, Cathedral Peak 
Area, Bergville, Almborn 9189 (LD); Ingeli Forest, Alfred, Almborn 10250 (LD); 


Impetyne Forest, near Weza, Almborn 9961 (LD); forest south of Nkandhla, 
Nkandhla, Almborn 8068 (LD). 


50. Parmelia direagens Hale, sp. nov. 


Thallus laxe adnatus, 5-15 cm. diametro, cinereo-albicans, lobis 
rotundatis, 6-12 mm. latis, suberectis, margine integris, sinuatis 
vel breve laciniato-lobatis, sorediatis, soraliis linearibus,  ciliatis, 
ciliis 1.5-2.5 mm. longis, superne nitidus, levissime albomaculatus, 
strato corticeo superiore 15-18 y, strato gonidiali 15-20 », medulla 
alba, 80-100 u crassa, strato corticeo inferiore 12-14 » crasso, subtus 
niger, sparse rhizinosus, ambitu castaneus, late nudus. Apothecia 
atque pycnidia ignota. Thallus K-+ flavescens; medulla K—, 
C—, KC+ rosea, P—, soralia P+ citrina, atranorinum, acidum 
alectoronicum et acidum P+ ignotum continens. 

Type in the British Museum, collected near Kimberly, Cape 
Province, South Africa, by John Shaw (isotype in US). 

This remarkable species could easily be misidentified as P. sancti- 
angelit Lynge, but it differs significantly in chemistry. When P is 
applies to the medulla, no reaction is observed, but the soredia are P+ 
deep yellow. Alectoronic acid is produced throughout the medulla 
and the soredia, but the soredia also contain the P+ yellow unknown. 
There is no other Parmelia species with this reaction. The upper 
surface tends to be faintly white-maculate, whereas P. sancti-angelit is 
not clearly maculate. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 289 


Additional specimen examined: 


UNION OF SOUTH AFRICA: Transvaau: 5 mi. east of Mokobulaan, 
Lydenburg, Almborn 7499 (LD). 


51. Parmelia eciliata (Nyl.) Nyl. in Fournier, Mex. Pl. 3. 1872. 
Parmelia crinita var. eciliata Nyl. Flora 52:291. 1869. 
P. resupina Stirton, Scot. Nat. 4: 298. 1877-78. Type of collection: Near 
Knysna, Cape Prov., Africa, Knobel (BM, holotype). 
(?)P. euryearpa Stein. & Zahlbr. Bot. Jahrb. Engler 60:530. 1926. Type 
collection: Amani, East Usambara, Africa, Brunnthaler s.n. (W, holotype). 


Type collection: Orizaba, Mexico, Bourgeaw (H, Nyl. herb. no. 
35295, holotype; P, isotype). 

Thallus loosely adnate, often found on twigs, membranous, up to 10 
cm. wide, mineral gray; lobes rotund, soon more or less convoluted and 
crowded, margins crenate to lobulate-dissected, lobules to 3 mm. long, 
more or less canaliculate, ciliate, cilia 1.0-1.5 mm. long; upper surface 
plane, shiny, maculate near the base of apothecia, isidia and soredia 
lacking; lower side black and sparsely rhizinate, brown to tan or white 
variegated and naked in a broad zone along the margins. Apothecia 
common, stalked, up to 18 mm. in diameter, amphithecium rugose, 
strongly white-maculate, disc imperforate; hymenium 120-150 u high; 
spores 12-18 X 23-30 y, episporium 3 p» thick; pycnidia present, conidia 
not seen. 

Reactions: Thallus K++ yellow; medulla K+ yellow, C—, KC—, 
P+ pale orange yellow, atranorine and stictic acid present. 

Parmelia eciliata bas no close relatives among species with stictic 
acid. It is morphologically rather similar to P. blanchetii Hue and 
P. abnuens Nyl., both of which differ in chemistry and in various 
minor morphological characters. This species occurs mainly in the 
Caribbean area where it is common on branches of deciduous trees, 
but it is also rather common in southern South America and in South 
Africa (fig. 18). The type of P. eurycarpa is placed in synonymy 
tentatively because the holotype is in poor condition; the presence of 
stictic acid and lack of soredia make it highly probable that it is iden- 
tical with P. eciliata. 

Additional specimens examined: 

MEXICO: Hipateo: Chapulhuacan, Lundell 7214e (MICH); Veracruz: 36 
km. southwest of Orizaba, Hale 19582 (S, US); 64 km. southwest of junction of 
highways 140 and 155, northeast of Huatusco, Hale 19549 (US). NICARAGUA: 
JinorreGa: Cerro de la Cruz, west of Jinotega, Standley 11091 (F). 

CUBA: Oriente: Loma del Gato, Hioram 6693 (BPI); Gran Piedra, Sierra 
Maestra, Imshaug 25035 (MSC). HAITI: Summit of ridge on Téte Etang, elev. 
5850 ft., Wetmore 2661 (MSC); summit of Montagne Noire, elev. 5500 ft., Wet- 


more 2798 (MSC) ; summit of ridge north of Forét des Pins, elev. 5800 ft., Imshaug 
22705 (MSC). JAMAICA: No locality, Wolle, 1875 (US). 


290 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


BRAZIL: Rio pr Janerro: Serra dos Orgiios, Burchell 2251 (K). ARGEN- 
TINA: Near Salta, Smith 4676 (US); Delta of Rio Parand, Buenos Aires, Sentes- 
son 109, 110, 1186 (8S); no locality, Lorentz & Hieronymus, 1872-76 (M). 

UNION OF SOUTH AFRICA: Transvaau: Kaaimansgat Forest, Woodbush, 
Mogg 1990 (PRE); Carr Prov.: Dassies Krantz, Grahamstown, Almborn 4017 
(LD); 6 mi. N. of Knysna, Knysna, Almborn 2619 (LD, US). 


52. Parmelia erasmia Hale, sp. nov. 


Thallus laxe adnatus, saxicola, 5-10 cm. diametro, cinereo-glau- 
cescens, lobis rotundatis, 8-10 mm. latis, margine ciliatis, ciliis 2.0- 
2.5 mm. longis, superne opacus, aetate rimosus, isidiatus, frequenter 
superficie neque margine, isidiis tenuibus ramosisque, strato corticeo 
superiore 20-25 yw crasso, strato gonidiali 18-20 yw crasso, medulla 
60-70 pu alta, parte superiore alba, inferiore pro parte crocea, strato 
corticeo inferiore fusco, 15-20 u crasso, inferne niger, sparse rhizinosus, 
ambitu castaneus, late nudus. Apothecia et pycnidia ignota. 
Thallus K+ flavescens; medulla alba K—, C—, KC+ rosea, P—, 
medulla crocea K+ atropurpurea, atranorinum, acidum alectoroni- 
cum, et pigmentum croceum ignotum continens. 

Type in the Chicago Natural History Museum, collected in the 
vicinity of El Zamorano, Morazin, Honduras, Noy. 26, 1946—Jan. 9, 
1947, by P. C. Standley (no. 210; isotype in US). 

Parmelia erasmia has the gross appearance of the widespread P. 
crinita Ach. The isidia are normal with no obvious tendency to 
become pustulate or sorediate. It might accidentally be placed with 
specimens of P. mellissii Dodge that have more normal isidia and 
well-developed pigments. However it is easy to prove with chroma- 
tography that the pigment in P. mellissii is rhodophyscin and that in 
P. erasmia is the same unknown anthraquinone found in P. hypomil- 
toides Vain. Parmelia hypomuiltoides has coarsely sorediate margins. 
All three species must be carefully studied morphologically and 
chemically before identifications are made. In any event P. erasmia 
is a rare species apparently endemic to pine-oak forests in Mexico and 
Central America, where it is probably more common than the few 
records indicate. 

Additional specimens examined: 

MEXICO: Veracruz: Teocelo Canyon, south of Xico, Hale 21139 (US). 
HONDURAS: Comayraaua: Siguatepeque, Standley & Chacén 6257 (F. US); 


Morazan: Quebrada el Gallo, northeastern slopes of Cerro de Uyuca, Morton 
7044 (REN, US). 


53. Parmelia eurysaca Hue, Nouv. Arch. Mus. Paris, ser 4, 1:194. 1899. PuLare 8 


Type collection: Mexico, Bourgeaw 1865-66 (P, holotype). 

Thallus loosely adnate to bark, coriaceous, ashy white to mineral 
gray, 10-20 cm. broad; lobes rotund, 10-14 mm. wide, conspicuously 
dentate-laciniate toward the center, laciniae 1-2><5-10 mm., margins 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 291 


ciliate, cilia 1-4 mm. long; upper surface plane, opaque to rather 
shiny, reticulately cracked with age, isidia and soredia lacking; lower 
side black and rhizinate in the center, brown to white-variegated in a 
broad zone at the margins. Apothecia 5-15 mm. in diameter, sub- 
stipitate, amphithecium rugose, white-maculate, disc perforate; 
hymenium 35-45 uw high; spores 6-89-12 yu, episporium 1 y» thick; 
pycnidia common, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K+ yellow turning red, 
C—, KC—, P-+ pale orange red, atranorine and salacinic acid present. 

Parmelia eurysaca is characterized by heavily pycnidiate marginal 
laciniae, which are always present in some degree. ‘This is the only 
ciliate species with salacinic acid that lacks soredia or isidia. A very 
close sorediate relative is P. stuppea Tayl., which lacks distinct 
laciniae. Parmelia eurysaca is endemic to central and northern 
Mexico and the adjacent United States, occurring very commonly 
on oak trees on dry open hillsides along with P. stuppea and P. 
praesignis Nyl. 

Additional specimens examined. 

U.S.: Texas: Head of Camp Verde Creek, Bandero Co., Parks 1068 (US, 
WIS). Arizona: Cave Creek, Chiricahua Mountains, Cochise Co., Weber et al. 
24741 (US). MEXICO: San Luis Porosf: Without locality, Parry 1022 (K); 
Alvarez, Palmer 465, 466 (US); Jatisco: San Sebastian, Sierra Madre Occidental, 
Mexia 1381, 1545c (F), 1904a (US); Hrpateo: Cuyamaloya, Pringle 10753 (US); 
El Hiloche, Martinez s.n. (US); Sierra de Pachuca, Madrigal 1415 (US); Vera- 
cruz: 64 km. southwest of junction of highways 140 and 155, northeast of 
Huatusco, Hale 19535 (US); 33 km. northeast of Perote, Hale 21220 (US); 
Mexico: North of Acambay, Cain 27597a (TRT, US); Sierra de las Cruces, 
Pringle 168 (COLO, US); Micuoacdn: 65 km. east of Morelia, Hale 20988 (US); 
Cerro Azul, Morelia, Arséne 3737 (US); 61 km. northwest of Zitdcuaro, Hale 
20861 (COLO, US); Oaxaca: Cerro San Felipe, Hale 20692 (DUKE, REN, §, 
TNS, US), 20709 (MSC, US). 


54. Parmelia flavotincta Hale, sp. nov. PLaTE 6 


Thallus expansus, laxe adnatus, membranaceus, 10-15 cm. latus, 
lobis rotundatis, 8-15 mm. latis, margine integris, centrum versus 
demum laciniatis, laciniis isidiiformibus, coralloideo-ramosis, apice 
granulato-fatiscentibus (ut in P. fasciculata Vain.), margine ciliatis 
ciliis 1-2 mm. longis, superne nitidus, planus, strato corticeo superiore 
20-26 u crasso, strato gonidiali 15-18 u crasso, medulla alba, 90-125 yu 
crassa, strato corticeo inferiore 14-18 yu crasso, inferne niger, sparse 
rhizinosus, ambitu fusco-castaneus, late nudus. Apothecia atque 
pycnidia ignota. Thallus K+ flavescens; medulla K-+ flavescens, 
C—, KC—, P+ flavotincta, atranorinum solum continens. 

Type in Kew Herbarium, collected at Cross Line, 8 mi. east of 
Berbice, British Guiana, July 17, 1919, by A. A. Abrahamson (no. 
229; isotype in US). 


292 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Parmelia flavotincta is the only Amphigymnia species that contains 
atranorine in the medulla. Morphologically it is close to P. fasci- 
culata Vain. and P. ramuscula Hale in the formation of coralloid 
isidial structures but differs in having a more membranous thallus 
and distinct cilia. 

Additional specimen examined: 

COLOMBIA: Vaupts: Cachivera de Jirijirimo, Schultes & Cabrera 11986 
(FH, US). 

55. Parmelia fracta Hale, sp. nov. 


Thallus expansus, laxe adnatus, 10-15 cm. latus, stramineo- 
flavicans, lobis rotundatis, usque ad 20 mm. latis, margine integris, 
centrum versus pro parte dentato-laciniatis, ciliatis, ciliis 1-2 mm. 
longis, pro maxima parte in lobis lateralibus, superne opacus, rugu- 
losus, demum pustulato-sorediatus, cortice partim comminuente, 
strato corticeo superiore 15-18 w crasso, strato gonidiali 14-16 yu 
crasso, medulla alba, 75-95 uw crassa, strato corticeo inferiore 13-15 pu 
crasso, subtus niger, sparse rhizinosus, ambitu castaneus, late nudus. 
Apothecia magna, pedicellata, usque ad 15 mm. diametro, amphi- 
thecio valde rugoso, albomaculato, disco imperforato; hymenium 
120-140 yw» altum; sporae 16-2030-38 yu, episporio 3-4 pw crasso; 
pycnidia rara, conidiis non visis. Thallus K+ flavescens; medulla 
K—, C—, KC-+ rosea, P+ aurantiaca, atranorinum, acidum usnicum, 
et acidum protocetraricum continens. 

Type in the Farlow Herbarium, Harvard University, collected in 
Colombia, South America, by Lindig (no. 740; isotypes in H, M, US). 

The upper cortex of P. fracta is quite fragile, flaking off and develop- 
ing broad pustulate-sorediate areas, reminiscent of P. bangit Vain. 
and P. rimulosa Dodge. It is known only from the type collection. 
Nylander had given it the herbarium name of “‘P. latissima f. flavida.” 


56. Parmelia grayana Hue, Nouv. Arch. Mus. Paris, ser. 4, 1:184. 1899. 
Parmelia cinereoplumbea Harm. in Pitt. & Harm. Bull. Soc. Bot. Fr. 58, 
suppl. 22:32. 1911. Type collection: Riscos de Casitas, Canary Islands, 
Pitard 2001 (DUKE, holotype). 
P. simodensis Asahina, Journ. Jap. Bot. 17:73, fig. 79. 1941. Type collec- 
tion: Simoda, Prov. Izu, Hondo, Japan, Asahina (TNS, holotype). 

Type collection: Coonoor, Nilgherries, India, Gray, 1893 (P, 
holotype). 

Thallus saxicolous, adnate to loosely attached, 3-6 cm. in diameter, 
coriaceous, whitish gray; lobes rotund, rather narrow, 4-8 mm. wide, 
crowded, margins crenate, often ascending, sorediate, soralia linear to 
subglobose, ciliate, cilia sparse to conspicuous, 0.5-1.5 mm. long, 
rather coarse; upper surface opaque, often pruinose, irregularly 
cracked with age, often capitate-sorediate near the margins, coalescing 
with the marginal soredia; lower side black and rhizinate at the 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 293 


center, dark brown and naked or in part papillate along the margins. 
Apothecia rare, adnate, to 4 mm. in diameter, disc perforate; hy- 
menium 50-60 u high; spores 5-712-14 yu, episporium 1.5 yu thick; 
pycnidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC—, P-, 
atranorine and protolichesteric acid present. 

Parmelia grayana is a saxicolous species with wide distribution in 
Asia and Africa (fig. 19). It is superficially like P. praesorediosa Nyl., 
which is sometimes saxicolous but lacks cilia. The thallus of P. 
grayana is usually coriaceous, the upper cortex white pruinose, and 
the cilia thick and coarse. 

Additional specimens examined: 

UNION OF SOUTH AFRICA: Transvaau: Pretoria, Smith 6276 (PRE); 
Punch Bowl Inn, Almborn 6483 (LD, US); Louis Trichardt, Zoutpansberg, 
Almborn 6098 (LD); Narau: Indumeni Forest, Cathedral Peak Area, Almborn 
8929 (LD, US); 1 mi. west of Cathedral Peak Hotel, Bergville Distr., Almborn 
9130 (LD); Carr Province: Mataliele, Héeg s.n. (TRH). 

JAPAN: Mt. Buzan, Shimoda, Prov. Izu, Kurokawa 58630 (TNS, US). 


57. Parmelia hololoba Hale, sp. nov. 


Thallus laxe adnatus vel suberectus, rigidiusculus, usque ad 10 cm. 
diametro, albicans, parum lobatus, lobis magnis, rotundatis, 10-20 
mm. latis, margine sinuatis, ciliatis, ciliis 1.0 mm. longis, superne 
opacus, laevigatus, leviter rimosus in aetate, cortice continuo, strato 
corticeo superiore 12-16 yu crasso, strato gonidiali 10-15 yu crasso, 
medulla alba, 70-85 yu crassa, strato corticeo inferiore 8-10 u crasso, 
inferne niger, sparse rhizinosus, margine castaneus, late nudus. 
Apothecia 5-10 mm. diametro, pedicellata, disco perforato, amphi- 
thecio albo-maculato in aetate; hymenium 60-70 yu altum; sporae 
7-8 10-12 uw; pyenidia numerosa, conidiis non visis. Thallus K+ 
flavescens; medulla K—, C+ rubra, KC+ rubra, P—, atranorinum 
et acidum lecanoricum continens. 

Type in the Kew Herbarium, collected at Entebbe, Uganda, Africa, 
Mar. 3, 1959, by Proctor (packet A; isotype in US). 

Although P. hololoba is known only from the type locality, the 
material is well developed and exhibits a unique combination of charac- 
ters. The presence of cilia in a species with lecanoric is particularly 
unusual. It is probably near P. cooperi Stein. & Zahlbr., which is 
sorediate, and may eventually prove to be the nonsorediate phase of 
this species. Except for the presence of cilia, it is very close to P. 
andina Mull. Arg. 


58. Parmelia hypomiltoides Vain. Acta Soc. Faun. Fl. Fenn. 7, no. 7:35. 1890. 


Type collection: Sitio, Minas Gerais, Brazil, Vainio, Lich. Bras. 
Exs. sn. (TUR, Vain. herb. no. 2543, holotype). 


294 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Thallus loosely adnate, membranaceous, 5-8 cm. in diameter, 
mineral gray; lobes rotund, margins entire to incised, ciliate, cilia 
1.5-2.5 mm. long, sorediate, soredia granular; upper surface shiny, 
faintly maculate, sorediate near the margins, soralia at first puncti- 
form, granular, coalescing irregularly, developing short subisidial 
growths; lower side black and sparsely rhizinate, brown and rugulose, 
naked, in a zone along the margin. Apothecia and pycnidia unknown. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ reddish 
orange, P—, in part orange red, reacting K+ purple, atranorine, 
alectoronic acid, and an unknown anthraquinone present. 

The type of P. hypomiltoides is a very poor fragment. The chief 
diagnostic characters are the orange-red medulla, quite conspicuous 
even near lobe tips, and the coarsely sorediate or even isidiate- 
sorediate laminal and submarginal soralia. By chromatography it 
is possible to show that the red pigment is not rhodophyscin, which 
is otherwise so common in Amphigymnia species with alectoronic 
acid. It is an unnamed pigment (No. 1) also found in P. erasmia 
Hale and P. mesogenes Nyl. Parmelia hypomiltoides could be con- 
fused with P. rampoddensis Nyl. which has rhodophyscin and marginal 
farinose soredia or with P. mellissii Dodge which also has rhodophyscin 
and distinct sorediate-isidia. In both of the latter species the pig- 
ment is associated with older or decomposed parts of the medulla, 
but it is still desirable to make chromatographic tests. Parmelia 
hypomiltoides is extremely rare and confined to southern Brazil, 
although it will probably be found eventually in Central America. 

Additional specimen examined: 


BRAZIL: Sao Pauto: Cruzeiro, Robert, November 1900 (BM). 
59. Parmelia inexspectata des Abbayes, Bull. Inst. Fr. Afr. Noire 20:16. 1958. 


Type collection: Mt Tonkoui, cercle de Man, Ivory Coast, des 
Abbayes (REN, holotype; US, isotype). 

Thallus to 10 cm. broad, loosely attached, mineral gray; lobes 
rotund, 10-15 mm. wide, margins subcrenate, ciliate, cilia 3-6 mm. 
long, upper surface plane, dull, soredia and isidia lacking, lower side 
black and sparsely rhizinate, brown and naked in a broad zone at 
the margins. Apothecia (from original description) to 15 mm. in 
diameter, pedicellate, exciple entire to dentate, amphithecium rugose, 
disc imperforate; spores 11-1423-28 u. 

Reactions: Thallus K+ yellow; medulla K+ orange red, C—, 
KC—, P—, atranorine and an unidentified K+ substance present. 

This species is characterized by a P—, K+ medullary reaction. As 
des Abbayes (1958) has pointed out, P. eurysaca Hue is somewhat 
similar in general appearance but contains salacinic acid, and, we 
might add, has perforate apothecia and small spores. Actually 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 295 


P. inexpectata seems to have no close relatives and more collections 
must be seen before we can assess the range of variation. 
Additional specimens examined: 


IVORY COAST: Near Cinchona station, Mont Tonkoui cercle de Man, 
Santesson 10618b, 10619a (UPS). 


60. Parmelia lophogena des Abbayes, Bull. Inst. Fr. Afr. Noire 20:19. 1958. 


Type collection: Dalaba, Fouta-Djalon, Guinea, Africa, des 
Abbayes (REN, holotype; US, isotype). 

Thallus loosely adnate on bark, 5-8 em. broad, mineral gray; 
lobes rotund, 7-10 mm. wide, margins entire, ciliate, cilia 1.0-1.5 
mm. long; upper surface shiny, plane, continuous, densely lobulate- 
isidiate, isidia inflated, pustulate, irregular, 0.2-0.3 mm. thick, to 
0.5 mm. high, fragile; lower side black and moderately rhizinate, 
pale brown to mottled ivory in a broad zone at the margins. 
Apothecia poorly developed, spores not seen. 

Reactions: Thallus K+ yellow; medulla K—, C+ rose, KC+ 
blood red, P—, atranorine and gyrophoric acid present. 

The more or less lobulate pustulate isidia set P. lophogena apart 
from all other species of Amphigymnia. It is still known only from 
the type collection. 


61. Parmelia maclayana Miill. Arg. Flora 74:376. 1891. PLATE 7 


Type collection: South of Tropic of Capricorn, Africa, Maclay 
(K, holotype; G, isotype). 

Thallus loosely attached, more or less coriaceous, 6-10 cm. in 
diameter, mineral gray; lobes rotund, 10-15 mm. wide, margins 
entire, become short, coarsely digitate-lobulate toward the center, 
ciliate, cilia rather sparse, to 2 mm. long, more common on lateral 
margins; upper surface plane, dull or rather shiny, very faintly 
maculate in part, continuous to more or less cracked with age; isidia 
and soredia lacking; lower side black, sparsely rhizinate, brown 
to tan and naked in a broad zone along the margins. Apothecia 
substipitate, 8-12 mm. in diameter, amphithecium maculate, rugulose, 
disc perforate; hymenium 50-70 yu high; spores 6-10X12-15 p, 
episporium 1.0-1.5 4; pycnidia present, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ orange 
red, P—, atranorine and alectoronic acid present. 

Parmelia maclayana is differentiated from three other nonsorediate 
African Amphigymnia species containing alectoronic acid (P. brevicil- 
vata Hale, P. procera Stein. & Zahlbr., and P. wainti A. L. Sm.) 
by perforate apothecia. Parmelia ornatula Hale is saxicolous and 
has dense cilia. Parmelia maclayana is known only from three widely 
separated localities in Africa. 


729-018—65——_9 


296 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Additional specimens examined: 


SIERRA LEONE: Kaballa, Thomas 2192 (K). TANGANYIKA: Peramiho, 
Dietrich s.n. (M). 

62. Parmelia margaritata Hue, Nouv. Arch. Mus. Paris, ser. 4, 1:193. 1899. 
PuateE 9 

Type collection: Ohio, Sulliwant (P, lectotype). 

Thallus loosely attached, 5-15 cm. in diameter, corticolous, mineral 
gray; main lobes rotund, 7-10 mm. wide, soon branching irregularly 
and developing numerous marginal laciniae, 1.5-2.0 mm. wide, 
3-5 mm. long, margins sparsely ciliate, cilia 1.0-1.5 mm. long; upper 
surface dull to shiny, faintly maculate, reticulately cracked with 
age, sorediate, soralia originating near the tips of laciniae in orbicular 
groups, coalescing into extensive irregular soralia, sorediate lobes 
usually becoming revolute; lower side black and sparsely rhizinate, 
dark brown and naked in a broad or rather narrow zone at the margins. 
Apothecia and pycnidia unknown. 

Reactions: Thallus K+ yellow; medulla K+ yellow turning red, 
C—, KC—, P+ pale orange red, atranorine and salacinic acid present. 

The soralia of P. margaritata originate on the surface of small 
marginal laciniae, eventually extending to the margins and causing 
the laciniae to become revolute. It is virtually identical therefore 
with P. arnoldit Du Rietz except for the more restricted distribution 
(fig. 9) and different chemistry. These two species have quite dif- 
ferent distribution patterns in the deciduous forests of eastern United 
States, P. margaritata being more common in mesic forests and P. 
arnoldit being more common at higher elevations. 

Additional specimens examined: 

U.S.: MassacuuseEtTts: Seavern’s Woods, Faron s.n. (DUKE); PENNSYLVANIA: 
Conewago, Lancaster Co., Heller s.n. (US); Disrrict or Cotumsia: Rock Creek, 
Miller 116 (US); West Vireinia: Pineville, Wyoming Co., Hale 11861 (US); 
Onto: Near Cincinnati, Lea 14 (PH); Marytanp: Red Run, Baltimore Co., 
Plitt 233 (BPI); Kentucky: Stanton, Powell Co., Prescott 66 (WIS) ; 6 mi. south- 
west of London, Laurel Co., Reed 58147 (Reed Herbarium); Wisconsin: Coon 
Valley, Vernon Co., Thomson 1907 (WIS); Iowa: Fayette Co., Fink s.n. (US); 
Vireinta: Browntown Valley Overlook, Skyline Drive, Rappahannock Co., 
Hale 14734 (US); 2 mi. east of Buena Vista, Rockbridge Co., Hale 18263 (US); 
Hone Quarry, Rockingham Co., Hale 18212 (US); near Evington, Campbell Co., 
Hale 15993 (US); Mountain Lake, Giles Co., Hale 12671 (US); 6 mi. east of Burnt 
Chimney, Franklin Co., Hale 15940 (US); Norru Carona: 12 mi. northeast of 
Pinnacle, Stokes Co., Culberson 6336 (DUKE); 8 mi. east of Weaverville, Bun- 
combe Co., Culberson 7161 (DUKE); 3 mi. east of Lenoir, Caldwell Co., Culber- 
son 7224 (DUKE); between Sands and Meat Camp, Watauga Co., Culberson 5982 
(DUKE); TENNEsSEE: Cliff Springs, Overton Co., Phillips s.n. (US); ARKANSAS: 
Eureka Springs, Carroll Co., Hale 2755 (US); 8 mi. south of Hollis, Perry Co., 
Hale 3097 (US); Mt. Ida, Montgomery Co., Hale 3035 (US); near Dallas, Polk 
Co., Hale 3957 (US); Ponea, Newton Co., Hale 2820 (US); Gateway, Benton Co., 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 297 


Hale 4010 (US); Mt. Magazine Park, Logan Co., Hale 3419, 3442 (US); Moun- 
tainburg, Crawford Co., Hale 2669 (US). 


63. Parmelia mellissii Dodge, Ann. Mo. Bot. Gard. 46:134. 1959. PLATE 1 
Parmelia crinita var. inactiva Magn. in Magn. & Zahlbr. Ark. Bot. 31A, no. 
6:104. 1944. Type collection: Between Haelaau and Nakalalua, Maui, 
Hawaii, Selling 5801 (5, holotype). 
P. allardii Hale, Bryol. 62:123. 1959. Type collection: Constanza, La 
Vega, Dominican Republic, Allard 16529 (US, holotype). 


Type collection: St. Helena, Melliss 23 (K, holotype). 

Thallus loosely adnate, corticolous, light mineral gray, 5-10 cm. 
broad; lobes rotund, 6-12 mm. wide, margins crenate to isidiate and 
dissected, ciliate, cilia 1-3 mm. long, KOH+ blue or K—; upper 
surface plane, dull, isidiate, especially near the margins, or sorediate- 
isidiate, isidia eventually becoming granular and sorediate, at ma- 
turity densely coralloid branched and ciliate, up to 3 mm. high; 
medulla white or in part reddish orange near the lower cortex; lower 
side black and rhizinate, naked and brown to tan or mottled white in 
a broad zone near the margins. Apothecia very rare, to 5 mm. wide, 
disc imperforate; hymenium 80-90 y high; spores 10-14 16-22 yu; 
episporium 1.5-2.0 »; pycnidia and conidia lacking. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ red or 
orange red, P—, pigmented medulla K+ purple, atranorine, alec- 
toronic acid, and frequently rhodophyscin present. 

Dodge’s type specimen, although poorly developed, has typical 
coralloid sorediate isidia. The medullary reaction that he gives, 
KC-, is erroneous since alectoronic acid is present. His specimen 
lacks the pigment rhodophyscin which is present in about half of the 
specimens examined, especially those that show signs of decay. 
Rhodophyscin appears to be produced whenever the thallus ages or 
begins to disintegrate and seemingly has no taxonomic value. 

Parmelia mellissii is pantropical with northward extensions into 
temperate United States and Japan. It had been identified as P. 
arnoldit Du Rietz in Japan because most specimens, as in the United 
States, are predominantly coarsely sorediate-isidiate. The range of 
variation from distinct coralloid isidia with scant development of 
soredia to the sorediate phase is very great and cannot be easily 
classified. Confusion with P. arnoldit and even P. rampoddensis 
Nyl. is possible, but the origin and type of soredia separate these 
species. 

Additional specimens examined: 

U.S.: TENNESSEE: Fall Creek Falls, Van Buren Co., Phillips 348 (US); Sout 
Carotina: Honey Hill, Berkeley Co., Culberson 10300 (DUKE); Ge&orara: 


Near Chatsworth, Murray Co., Hale 7439 (US); ALaBaMA: Bankhead National 
Forest, Winston Co., McCullough 574 (US); Fioripa: 6 mi. southwest of Talla- 


298 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


hassee, Leon Co., Hale 16986 (US); Alum Bluff, Apalachicola River, Liberty Co., 
Thomson 5144 (WIS); Texas: Pineland, Sabine Co., Hale 5161 (US). MEXICO: 
Curapas: Lagos de Monte Bello, Hale 20961 (DUKE, MSC, REN, TNS, US); 
El Suspiro, 10 km. north of Berriozibal, Hale 20235 (5S, US). GUATEMALA: 
San Marcos: Barranco Eminencia, Standley 86498 (MO). HONDURAS: 
MorazAn: Lower slopes of Cerro de Uyuca, Standley 12034 (F). COSTA RICA: 
Cuesta de Terraza, Tonduz s.n. (G). 

CUBA: Orrente: Gran Piedra, Sierra Maestra, Imshaug 25044 (MSC, US). 
HAITI: Eastern end of Montagne Noire, near Kenscoff, Imshaug 22541 (MSC, 
US); vicinity of Forét des Pins, SHADA Station, Wetmore 3212 (MSC, US), 
2925, 3074 (MSC), Imshaug 22756 (MSC, US); summit of Pic La Selle, Imshaug 
22982 (MSC, US), Wetmore 3130 (MSC). DOMINICAN REPUBLIC: Summit 
of Isabel de Torres, near Puerto Plata, Imshaug 23927 (MSC); southwest slope 
of La Rucilla, Maciso Central, Imshaug 23595 (LD, MSC, US); between Pico 
del Yaque and Chinguela, Cordillera Central, Wetmore 3698, 3735 (MSC, US). 
JAMAICA: Without locality, Cummings 43 (FH, NY); Abbey Green Woodland, 
Harris 10028 (BM); Hollymount, Mt. Diablo, Jmshaug 14235 (MSC, US); Sir 
Johns Peak, Blue Mountains, Imshaug 15175 (MSC). PUERTO RICO: Adjuntas, 
Sintenis 99, 110a (G). MARTINIQUE: Piton Dumauzé, Degeliuss.n. (DEGEL). 

COLOMBIA: Vicinity of Medellin, Charetier 188 (US); San Isabel, s.c. (G). 
BRAZIL: Minas Gerais, Warming 272 (M). 

CANARY ISLANDS: Teneriffe, Cook 1130 (US). 

INDIA: Above Kurseong, Darjeeling, Awasthi 3923 (AWAS). LAOS: 
Hase Plantations, Tsuyama 1 (TNS). CHINA: Near Yunnanfu, Yunnan, 
Handel-Mazzetti 66 (WU). JAVA: Gegerbentang, Neervoort 3048 (BO); Tjinji- 
roea, Van Slooten s.n. (BO). JAPAN: Tanso-st: Kominato, Prov. Awa, Kuro- 
kawa 56572 (TNS, US); Matsukawa-ura, Prov. Iwaki, Kurokawa 58088 (TNS, 
US). 

Additional records from the United States, Cuba, Jamaica, Venezuela, and 
Formosa listed by Hale (1959b) as P. allardii are not repeated here. 


64. Parmelia merrillii Vainio, Phil. Journ. Sci. 4:658. 1909 PLATE 8 


Type collection: Mt. Halcon, Mindoro, Philippine Islands, Merrill 
6163 (TUR, lectotype). 

Thallus large, loosely attached, 8-10 cm. in diameter, mineral gray; 
lobes 10-15 mm. wide, rotund, margins laciniate toward the center 
of the thallus, laciniae 0.5-1.0 2-5 mm., sparsely ciliate, cilia 1.0-1.5 
mm. long; upper surface dull, reticulately cracked with age, isidia 
and soredia lacking; lower side black, sparsely rhizinate, brown and 
naked in a broad zone along the margins. Apothecia abundant, 
4-10 mm. in diameter, exciple dentate-laciniate, ciliate, amphithe- 
cium rugose, white-maculate, disc imperforate; hymenium 90-110 yu 
high; spores 12-16X26-34 yu, episporium 2.5-3.0 uw thick; pycnidia 
common, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ orange 
red, P+ brick red, atranorine and protocetraric acid present. 

Parmelia merrillii has long narrow laciniae on the lobes and fre- 
quently on the exciple of the apothecia. It could be confused super- 
ficially with P. corniculans Nyl., another Asian species, which differs 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 299 


chiefly in chemistry (alectoronic acid present) and in having perforate 
apothecia. Parmelia pachyspora Hale, an African species, has 
similar chemistry and large spores, but differs in having sparse cilia 
and in lacking distinct marginal laciniae. 

Additional specimens examined: 

BOLIVIA: CocuaBamBa: Schuenca-Carrasco, Cardenas 5284 (US). 

FORMOSA: Chokakurai, Masuda F.546 (TNS, US). BORNEO: Marai 
Parai, Mt. Kinabalu, Clemens 32425 (BO). CELEBES: B. Watoewila, Kjellberg 
41L (BO). SUMATRA: Batang Paloepoeh, Jacobson s.n. (BO). JAVA: 8.1., 
Horsfield (BM). 


65. Parmelia ornatula Hale, sp. nov. PLATE 10 


Thallus adnatus, saxicola, 5-10 cm. diametro, albicans vel cinereo- 
albicans, lobis rotundatis, contiguis, elongatis, 4-7 mm. latis, margine 
suberectis, crenatis, dense ciliatis, ciliis suberectis, 1-2 mm. longis, 
sorediis isidiisque destitutis, superne nitidus, continuus, planus vel 
late concavus, irregulariter rimosus in aetate, strato corticeo superiore 
13-17 p» crasso, strato gonidiali 15-26 » crasso, medulla alba, 50-65 u 
crassa, strato corticeo inferiore 13-15 uw crasso, subtus niger, modice 
rhizinosus, ambitu castaneus, anguste nudus. Apothecia pedicellata, 
4—7 mm. diametro, excipulo ciliato, amphithecio rugoso, albomaculato, 
disco perforato; hymenium 40-50 yw altum; sporae 5-7X10-13 u, 
episporio 1.5 w; pycnidia numerosa, conidiis non visis. Thallus K+ 
flavescens; medulla K—, C—, KC+ rubescens, P—, atranorinum 
et acidum alectoronicum continens. 

Type in herbarium of Gunnar Degelius, collected near Faz. Can- 
jangue, Vila Flor, Huambo, Angola, Africa, Feb. 18, 1960, by G. 
Degelius (isotype in US). 

This is a saxicolous species with rather narrow crowded lobes and 
dense suberect marginal cilia, as well as a few laminal cilia. It would 
appear to be a distant relative of another saxicolous species with 
alectoronic acid, P. breviciliata Hale, which is a larger species with 
imperforate apothecia. Parmelia ornatula is known only from the 
type collection and is probably endemic to Angola. 


66. Parmelia pachyspora Hale, sp. nov. 


Thallus laxe adnatus, expansus, 12-20 cm. diametro, cinereo- 
albicans, lobis rotundatis, 10-15 mm. latis, margine plus minusve 
crenatis, saepe dentato-laciniatis, suberectis, sparse ciliatis, ciliis 
1.0-2.5 mm. longis, sorediis isidiisque destitutis, superne opacus, 
planus, continuus vel irregulariter rimosus, strato corticeo superiore 
13-17 yu crasso, strato gonidiali 20-26 u crasso, medulla alba, 75-100 u 
crassa, strato corticeo inferiore 15-18 yu crasso, inferne niger, sparse 
rhizinosus, ambitu castaneus vel eborinus, late nudus. Apothecia 
pedicellata, 5-8 mm. diametro, excipulo plus minusve ciliato-dentato, 


300 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


amphithecio valde rugoso atque albomaculato, disco imperforato; 
hymenium 80-90 u altum; sporae 13-15 30-34 y, episporio 3 u crasso; 
pycnidia numerosa, conidiis 15-6 yw. Thallus K+ flavescens; 
medulla K—, C—, KC+ rosea, P+ aurantiaca, atranorinum et 
acidum protocetraricum continens. 

Type in the herbarium of Gunnar Degelius, collected 10 km. north 
of S4 da Bandeira, Huila, Angola, Africa, Feb. 3, 1960, by G. Degelius 
(isotype in US). 

The apothecia of P. pachyspora are stalked and conspicuous. The 
closest related species is P. merrillii Vain., an Asian species with 
long marginal laciniae. Parmelia pachyspora is known from several 
localities in southern Africa. 

Additional specimens examined: 

ANGOLA: Moxtco: Between Luso and Cochipoque, Degelius, Feb. 16, 1960 
(DEGEL); Cuanza Sut: Amboim, Gossweiler 10006 (BM). NYASALAND: 
Kyimbila, Stolz 2760 (C); Mt. Massangulo, Gomez 8 (K). SOUTHERN RHO- 


DESIA: Salisbury, Moss 2217 (PRE). MOGAMBIQUE: Nampula Distr., 
Gomes 2134 (PRE). 


67. Parmelia perlata (Huds.) Ach. Meth. Lich. 216. 1803. 

Lichen perlatus Huds. Fl. Angl. 448. 1762. 

Parmelia coniocarpa Laurer, Linnaea 2:39. 1827. Type collection: Aus- 
tralia, Sieber 50 (M, holotype; S, isotype). 

P. coriacea var. perlata (Huds.) Eschw. in Mart. Fl. Bras. 1:206. 1833. 

P. schweinfurthii f. sorediata Mill. Arg. Flora 70:59. 1887. Type collec- 
tion: East Gippsland, Australia, Walter (M, holotype; G, isotype). 

P. trichotera Hue, Journ. de Bot. 12:245. 1898. Type collection: France, 
Malbranche, Lich. Norm. 65 (P, lectotype; DUKE, isotype). 

P. piloselloides Zahlbr. Svensk. Vet. Akad. Handl. 57, no. 6:48. 1917. 
Type collection: Masatierra, Cumberland Bay, Juan Fernandez, Skottsberg 
(W, holotype; 8, isotype). 

P. protosorediata Gyel. Repert. Sp. Nov. Fedde 29:288. 1931. Based on 
P. schweinfurthw f. sorediata Mill. Arg. 

P. cristifera f. pallida Ris. Ann. Bot. Soc. Zool. Bot. Fenn. Vanamo 3:3. 
1944. Type collection: Carrasco, Montevideo, Uruguay, Herter, 1929 
(H, holotype). 

Type collection: Specimen and pl. 20, fig. 39B, Dillenius Hist. 
Muse. 147. 1741 (OXF, lectotype). 

Thallus adnate to loosely attached, 5-15 cm. in diameter, mineral 
gray; lobes 8-15 mm. wide, margins entire, ciliate, cilia sparse, 0.3- 
2.0 mm. long, submarginally sorediate, soredia originating in linear 
soralia, soon eroding a large area and causing the margins to become 
strongly revolute; upper surface smooth, opaque; lower side black 
and moderately rhizinate, dark brown, shiny, and naked in a broad to 
narrow zone along the margins. Apothecia very rare, 3-5 mm. in 
diameter, disc imperforate; hymenium 70-80 yu high; spores 13-16 
22-30 pw, episporium 3 y thick; pycnidia not seen. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 301 


Reactions: Thallus K+ yellow; medulla K+ persistent yellow, 
C—, KC—, P+ pale orange yellow, atranorine and stictic acid present. 

I recently made a study (1961) of the typification of this common 
species in order to stabilize its name. The diagnostic characters are 
the conspicuous submarginal soralia, revolute lobes, and the presence 
of stictic acid. Specimens from Juan Fernandez named as P. pilosel- 
loides Zahlbr. have more extensive sublaminal soredia than normal 
but still fall within the range of P. perlata. The narrow, shiny and 
often rugose naked zone below suggests a relationship between P. 
perlata and isidiate P. crinita Ach., which also produces stictic acid. 
Parmelia perlata is widespread in temperate regions of all continents 
(fig. 20). In Scandinavia it is an oceanic species (Almborn, 1948) 
with a distribution pattern similar to that of P. arnoldii Du Rietz 
and P. erinita Ach. 


Additional specimens examined: 


CANADA: Nova Scotra: Hunt’s Point, Denison (US); Uclucet, Vancouver 
Island, Macoun, Canad. Lich. 44 (DUKE). U.S.: PENNSYLVANIA: 2 mi. east of 
Worlds End State Park, Sullivan Co., Hale 17226 (US); West Virernra: Near 
West Union, Doddridge Co., Hale 10385 (US); Wellford, Kanawha Co., Hale 
10862 (US); Kentucky: Near Baldrock, Laurel Co., Reed 58118 (Reed Her- 
barium); Vrroinra: 1 mi. north of Gillespie, Bedford Co., Hale 15762 (US); 
Whitetop Mountain, Washington Co., Hale 18490 (US); TsNnNEssEE: Near 
Elkmont, Sevier Co., Cain 85 (US); Cliff Springs, Overton Co., Phillips 313 
(US); Norra Caroiina: Near Yadkin Valley, Caldwell Co., Culberson 6507 
(DUKE, US); near Candler, Buncombe Co., Green s.n. (US); summit of Mt. 
Mitchell, Yancey Co., Imshaug 22358 (MSC); Grorata: 5 mi. east of Hiawassee, 
Towns Co., Culberson 7289, 7264 (DUKE); Orecon: Cape Blanco, Port Oxford, 
Imshaug 17655 (MSC); Catirornia: Freshwater, Humboldt Co., Becking 
61060012 (US); Woodside, San Mateo Co., Cain 26378 (TRT, US); Point Reyes, 
Marin Co., Imshaug 17686 (MSC). MEXICO: San Luis Porosf: Alvarez, 
Palmer 467 (US); Hipauao: 20 mi. southwest of Jacala, Cain 27619 (TRT, 
US); Pursia: 3 km. west of Puebla-Veracruz state line on highway 150, Hale 
19645 (US); Mexico: La Cima, Pringle 10725 (US); Veracruz: 33 km. north- 
east of Perote, Hale 19354 (US). PANAMA: Curriquf: Chiriqui volcano, 
Scholander s.n. (MO). 

HAITI: Summit of Téte Etang, Imshaug 22614 (MSC), 22620 (MSC, US). 

CHILE: Juan Fernandez, Skottsberg s.n. (8S), s.c. (K); Coquimso: Fray Jorge, 
Sparre 3020 (8S); Vavpararso: Lago Pefiuelas, Santesson 7002 (S); VALDIVIA: 
Corral, Santesson 7064 (S); Lago Rifftihue, Santesson 3511; Cavutin: Temuco, 
Gunckel 18204 (MO); Cutto&: Ancud, Isla Chiloé, Santesson 4234 (S); Laraquete, 
Hosseus 75 (M). 

IRELAND: Galway, Degelius sn. (UPS). SCOTLAND: Between Salen 
and Drumlang, Argyll, Lindahl 95d (UPS). ENGLAND: Bilsdale, Yorkshire, 
Baker s.n. (US). NORWAY: Seljuasen, Sokndal, Santesson s.n. (US). 

NETHERLANDS: Goes, Zeeland, Maas Geesteranuss.n. (US). POLAND: 
Puszeza Bukowa, Glance, Lich. Polon. 173 (LD). FRANCE: St. Thurial, Ille- 
et-Vilaine, Santesson 10080 (US); Rennes Woods, des Abbayes, Lich. Armor. 
Spect. Exs. 107 (LD); Le Trayas, Estrel, Suza, Crypt. Exs. Vind. 3657 (US); 


302 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Troyes, Culberson 1026 (DUKE, US). SPAIN: South of the Santuario, 
Covadonga, Santesson 13085b (UPS). PORTUGAL: Near Azoia, Estremadura, 
Tavares, Lich. Lus. sel. Exs, 141 (LD, US, WIS); Pinhal, Douro Litoral, Tavares 
4199 (US); Peso, Minho, Tavares sn. (WIS). ITALY: Torriglia, Liguria, 
Almborn s.n. (LD); Arenzano, Sbarbaro s.n. (WIS); Veneto, Mass. Lich. Ital. 
325 (UPS). 

AZORES: Santa Maria, Baldwin 14196 (US); St. Michel, Persson s.n. (UPS). 
TUNISIA: Ain Draham, Runemark s.n. (LD). CANARY ISLANDS: Cumbu 
Nueva, Pitard 2008 (DUKE). MADEIRA: Funchal, Paton s.n. (BM). 
TRISTAN DA CUNHA: Gong Island, van der Merwe 51a (LD). KENYA: 
Tinderet Forest Reserve, Kisumu-Londiani, Nyanza Prov., Maas Geesteranus 
11167 (L). UNION OF SOUTH AFRICA: Transvaat: Houtbosch, Pietersburg, 
Almborn 6768 (LD); Nata: Polela Forest, Polela, Almborn 9514 (LD, US); 
Upper Umkomaas, Impendhle Distr., Héeg sn. (TRH); Care Province: 
3 mi. west of Heidelberg, Swellendam, Héeg s.n. (TRH); Kirstenbosch, Wynberg, 
Almborn 1889 (LD). 

JAPAN: Mt. Oodake, Musashi, Kurokawa 59147 (TNS, US); Shikkari, Prov. 
Mutsu, Kurokawa 550310 (TNS, US). HAWAIIAN ISLANDS: Kauai, Kraus 
1128 (Darrow Herbarium). AUSTRALIA: Illawarra, New South Wales, Kirton 
sn. (BM). NEW ZEALAND: Plymouth, Thomson s.n. (BM); Wellington, 
Buchanan s.n. (BM). 


68. Parmelia permutata Stirton, Scot. Nat. 4:252. 1877-78. 


Type collection: Near Brisbane, Australia, Bailey (BM, holotype; 
GLAM, isotype). 

Thallus 5-15 cm. broad, loosely attached, mineral gray; lobes 
rotund, 8-15 mm. wide, margins often suberect, entire, ciliate, cilia 
2.0-4.5 mm. long, sorediate, soralia elongate, sinuous, rarely sub- 
marginal; upper surface smooth, opaque, reticulately cracked with 
age; medulla white in upper half, becoming orange yellow in lower half; 
lower side black and sparsely rhizinate, brown and naked in a broad 
zone along the margins. Apothecia and pycnidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C+ rose, KC+ red, 
P—, pigment K—, atranorine, gyrophoric acid, and an unidentified 
yellow pigment present. 

Parmelia permutata has a pale orange-yellow medulla with the same 
pigments that are found in P. araucariarum Zahlbr., P. myelochroa 
Hale, and others. In P. permutata the pigments are definitely local- 
ized in the lower half of the medulla and the upper half is white. 
When pigments are heavily concentrated in P. sancti-angelii Lynge, 
which is also C+ rose, it is easy to confuse these two species, but the 
pigment in P. sancti-angelii is rhodophyscin, which reacts K+ 
purple. Parmelia rampoddensis Nyl., which is C—, KC+ red 
(alectoronic acid), is also very similar externally and may produce 
rhodophyscin. In doubtful cases chromatography should be used to 
identify the pigments. Parmelia permutata is a rather rare species 
known only from Australia, Africa, and Asia and a single locality 
in the West Indies. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 303 


Additional specimens examined: 


HAITI: Near Furey, Bro. Fabius, July 1962 (US). 

IVORY COAST: Danané, cercle de Man, des Abbayes, Oct. 19, 1951 (TUR). 
KENYA: Nyanza: Tinderet Forest Reserve, Kisumu-Londiani, Maas Geesteranus 
4954, 4956 pr. p. (L). UNION OF SOUTH AFRICA: Narat: Pietermaritzburg, 
Almborn 9633 (LD); Transvaaut: Near “Punch Bowl,’ Louis Trichardt, Zout- 
pansberg, Almborn 6245 (LD). 

INDIA: Darjeeling, Thomson, s.d. (GLAM). THAILAND: Hase, Tsuyama 
20 (TNS). SUMATRA: Between Lubuk Selasih and Aer Batumbuk, Groen- 
hart 9355 (BO). 

69. Parmelia procera Stein. & Zahlbr. Bot. Jahrb. Engler 60:537. 1926. 
PLATE 10 


Type collection: Near Amani, Tanganyika, Brunnthaler, August 
1890 (W, lectotype). 

Thallus large, 10-15 cm. in diameter, coriaceous, loosely attached, 
buff, mineral gray; lobes rotund, 8-15 mm. wide, often becoming 
convoluted and crowded toward the center, margins smooth or be- 
coming short lobulate-dentate or laciniate, laciniae suberect, sparsely 
to moderately ciliate, cilia 2-4 mm. long; upper surface smooth, at 
length finely reticulately cracked, faintly maculate at the base of 
apothecia; lower side black and sparsely rhizinate, brown and naked 
in a broad zone along the margins. Apothecia very large, to 30 mm. 
in diameter, stalked, amphithecium maculate, disc imperforate; 
hymenium 60-70 » high; spores 5-612-16 yu, episporium 1 4g; 
pycnidia present, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ red, 
P—, atranorine and alectoronic acid present. 

Parmelia procera belongs to that variable difficult group of non- 
sorediate, nonisidiate species with alectoronic acid. It differs from 
P. maclayana Mill. Arg. and P. ornatula Hale in having imperforate 
apothecia. Parmelia brevciliata Hale is saxicolous, and P. wainir 
A. L. Sm. has long cilia and intermediate spores. 

Additional specimens examined: 

TANGANYIKA: Amani, Greenway 1001 (K). MOCAMBIQUE: Massangulo 
Sandrone (F). NYASALAND: Mt. Massangulo, Sousa 8 (K). ANGOLA: 
Capir, Cuanza Sul, Gossweiler 9907 (US). 


THAILAND: Doi Sutep, Tsuwyama 7 (TNS, US). 
NEW CALEDONIA: Compton 1471 (BM). 


70. Parmelia pseudocrinita des Abbayes, Bull. Inst. Fr. Afr. Noire 20:19. 1958. 
Parmelia brauniit Dodge, Ann. Mo. Bot. Gard. 46:135. 1959. Type collec- 
tion: Tanganyika, Braun, Inst. Amani 8603 (EA, holotype). 
P. mannii Dodge, Ann. Mo. Bot. Gard. 46:136. 1959. Type collection: 
Ilha Principe, Africa, Mann, 1861 (K, holotype). 


Type collection: Dalaba, Fouta-Djalon Mountains, Guinea, des 
Abbayes, Feb. 5, 1951 (REN, lectotype; US, isotype). 


304 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Thallus large, 10-15 cm. in diameter, rather coriaceous, loosely 
attached, mineral gray; lobes rotund, 8-12 mm. wide, margins crenaie, 
ciliate, cilia coarse, 1-2 mm. long, upper surface smooth, dull, be- 
coming rugulose or fissured on older lobes, isidiate, isidia simple to 
rarely coralloid branched, 0.05-0.070.2-0.5 mm., rarely ciliate 
apically; medulla white, rarely orange red near the lower cortex; 
lower side black and sparsely rhizinate, brown, shiny, and naked in 
a broad zone at the margins. Apothecia rare, short stalked, 2-3 mm. 
in diameter, amphithecium isidiate, disc imperforate; hymenium 50-65 
u high; spores 7-8 11-13 yu, episporium 1 yp thick. 

Reactions: Thallus K+ yellow; medulla K—, C+ rose, KC+ red, 
P—, pigmented medulla K+ purple, atranorine, gyrophoric acid, and 
(if pigmented) rhodophyscin present. 

Parmelia pseudocrinita is a rather common lichen in southern Africa. 
The overall external appearance reminds one at once of P. crinita 
Ach., but the cilia are coarse, the spores small, and the medulla K—, 
C+ rose. 

Additional specimens examined: 

ANGOLA: Huita: Between Coporolo and Chingorvi, Degelius, Feb. 2, 1960 
(DEGEL); Bri: Coemba, Degelius, Feb. 10, 1960 (DEGEL). CONGO: 15 km. 
west of Goma, Goma, North Kivu, Degelius, Mar. 16, 1960 (DEGEL) ; Yangambi, 
Luweo Plateau, Louis 8151, 8155 pr. p. (BR); north of Elisabethville, Héeg, Apr. 
16, 1980 (TRH). SOUTHERN RHODESIA: Matopos, Borle 54a (PRE). 
UNION OF SOUTH AFRICA: Near Solheim, Eschowe (Zululand), Héeg, Sept. 
5, 1929 (TRH); Narau: 1 mi. north of Umzumbi, Umzinto, Almborn 9870 (LD). 
71. Parmelia rampoddensis Nyl. Acta Soc. Sci. Fenn. 26, no. 10:7. 1900. 

Parmelia proboscidea var. sorediifera Mill. Arg. Flora 67:615. 1884. Type 
specimen: Central Madagascar, Hildebrandt (G, lectotype). 

P. proboscidea f. sorediifera (Mill. Arg.) Mill. Arg. Bull. Soc. Bot. Belg. 
30:53. 1891. 

P. proboscidea f. bulbifera Hue, Nouv. Arch. Mus. Paris, ser. 4, 1:197. 1899. 
Type collection: Coonoor, Nilgherries, India, Gray, 1883 (P, holotype). 

P. poolit Dodge, Ann. Mo. Bot. Gard. 46:146. 1959. Based on P. pro- 
boscidea var. sorediifera Mill. Arg. 

P. subinvoluta Hale, Bryol. 62:130, fig. 4. 1959. Type collection: 4 mi. 
west of Midway, Liberty Co., Georgia, Hale 16807 (US, holotype). 

Type collection: Ramboda, Ceylon, Almquist (H, Nyl. herb. 
no. 35555, holotype; S, isotype). 

Thallus large, expanded, 10-30 cm. wide, loosely attached, whitish 
to mineral gray; lobes broad and rotund, 12-20 mm. wide; margins 
entire, sorediate, soralia linear, 0.2-0.5 mm. wide, sorediate margins 
more or less involute, ciliate, cilia conspicuous, 3-6 mm. long; upper 
surface plane, rugulose with age; medulla white, often in part orange 
red near the lower cortex; lower side black and sparsely rhizinate, 
brown and naked in a broad zone along the margins. Apothecia 
rare, 3-10 mm. in diameter, amphithecium sorediate, disc imperforate; 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 305 


hymenium 65-75 y» high; spores 6-7 10-12 yu, episporium 1 y; pycni- 
dia present, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC-+ red, 
P—, pigmented medulla K+ purple, atranorine, alectoronic acid 
and (if pigmented) rhodophyscin present. 

After describing P. subinvoluta, I had an opportunity to examine 
the type of P. rampoddensis Nyl. from Ceylon, which proved to be 
identical. Further collections of the species were soon found. It 
is in reality a pantropical lichen, although most common in southern 
United States and the Caribbean area. Rhodophyscin is found in a 
small percentage of the specimens. When the pigment is abundant, 
one might identify a specimen as P. hypomiltoides Vain., which 
differs in having more or less submarginal isidiate soredia and a 
different anthraquinone. 

Additional specimens examined: 


U.S.: Nortu Carouina: 5 mi. east of Calypso, Duplin Co., Culberson 6675 
(DUKE); Sovurn Caro.uina: 3 mi. northeast of Darlington, Darlington Co., 
Culberson 7648 (DUKE); ALaBamA: Pocosin area 4 mi. southeast of Troy, Pike 
Co., McCullough 492 (US); Froripa: Tomoka State Park, Volusia Co., Hale 
17062 (US); 5 mi. east of Greenville, Madison Co., Hale 17615 (US); Crewsville, 
Hardee Co., Hale 16876 (US); Lourstana: Near Alexandria, Rapides Parish, 
Logan 881b (WIS), 881c (MO); Texas: 7 mi. south of Silsbee, Hardin Co., 
Whitehouse 25955 (MO); Jasper, Jasper Co., Hale 5416 (US). MEXICO: 
Oaxaca: 47 mi. south of Oaxaca, Kramer 2333 (KANU, US); Veracruz: Mira- 
dor, Purpus 86 (US); Cutapas: 2 km. north of highway 190 on road to Puebla 
Nueva, Hale 20171, 20185 (US); El Zapotal, Tuxtla Gutiérrez, Hale 20004 (US). 

HAITI: Eastern end of Montagne Noire, Imshaug 22543 (MSC). DOMINI- 
CAN REPUBLIC: North of San José de las Matas, Santiago, Wetmore 3897 
(MSC). JAMAICA: Mandeville, Cushman 12 (FH); trail above Mavis Bank 
Road, St. Andrew, Imshaug 14429 (MSC). 

COLOMBIA: Antroaufa: Medellin, Daniel 925 (US). BOLIVIA: 20 km. 
west of San José de Chiquitos, Santa Cruz, Cutler 7071 (F). BRAZIL: Minas 
Gerais: S&0 Jo&o d’el Rey, Malme 270 (8). 

CAMEROONS: Staudt 692 (BM). MOCAMBIQUE: 2 km. east of Namaa- 
cha, Lourengo Marques, Almborn 7107 (LD). UNION OF SOUTH AFRICA: 
TRANSVAAL: Blaauwberg, Pietersburg, Leemann 1533 (PRE). MADAGASCAR: 
Ambatolaona, Benoist 326 (LD). 

INDIA: Senchal, Darjeeling, Togashi s.n. (TNS). FORMOSA: Renegechi, 
Asahina F.55 (TNS, US). NEW GUINEA: Mt. Arfak, Kostermanns 3017 
(BO). AUSTRALIA: Parramatta, New South Wales, s.c. (BM). 

Additional records from the United States, Jamaica, Dominican Republic, and 
Honduras listed by Hale (1959b) as P. subinvoluta are not 1epeated here. 


72. Parmelia rimulosa Dodge, Ann. Mo. Bot. Gard. 46:133. 1959. PuLate 1 
Type collection: Table Mountain, Cape of Good Hope, Union 
of South Africa, MacGillivray (K, holotype). 


Thallus 5-15 cm. in diameter, loosely attached, light mineral gray 
membranous; lobes rotund 7-12 mm. wide, becoming crowded, 


306 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


imbricate and convoluted with age, margins entire to dentate- 
laciniate, ciliate, cilia rather sparse, 2-3 mm. long; upper surface 
plane, becoming quite rugose, reticulately cracked with age, cortex 
cracking apart and flaking away in a broad zone along the margins, 
becoming sorediate or isidiate-pustulate, clearly maculate only at the 
base of apothecia; medulla white, sometimes orange red near the lower 
cortex; lower side black and rhizinate, brown to mottled tan and 
naked in a broad zone along the margins. Apothecia rare, 8-15 mm. 
in diameter, stalked, amphithecium rugose, maculate, disc perforate; 
hymenium 70-80 » high; spores 10-13 X 20-22 yu, episporium 1.5-2.0 
thick; pycnidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC-+ red, P—, 
sometimes pigmented orange red near the lower cortex, K-+ purple, 
atranorine, alectoronic acid, and if pigmented rhodophyscin present. 

Parmelia rimulosa is a typical South African species extremely 
common and confined largely to Natal and the Cape Province. The 
outstanding character is the cracking and flaking of the upper 
cortex, usually giving rise to extensive laminal pustulate-sorediate 
areas. The plants are quite thin and membranous. There seem to 
be no close relatives, although heavily sorediate specimens might be 
confused with P. natalensis Stein. & Zahlbr., a maculate species with 
distinct laminal soredia. 

Additional specimens examined: 

KENYA: Nyanza: Tinderet Forest Reserve, Kisumu-Londiani, Maas Geestera- 
nus 11160 (L). UNION OF SOUTH AFRICA: Nara: South of Nkandhla, 
Nkandhla, Almborn 8069 (LD); along ‘‘Mountain Road,’’ Cathedral Peak Area, 
Bergville, Almborn 9375 (LD); Mgnalsheni, Pohla, Héeg, Oct. 9, 1929 (TRH); 
Pietermaritzburg, Umgeni, Héeg, Sept. 29, 1929 (TRH); Impendhla, Héeg, Oct. 8, 
1929 (TRH); Care Province: Deepwells, Knysna, Almborn 10916 (LD), Maas 
Geesteranus 12163-12165, 12176 (L); Knysna, Knobel, Apr. 12, 1878 (BM); 6 mi. 
N. of Knysna, Almborn 2697 (LD); Grootvadersbosch, Swellendam, Almborn 
2174 (LD); Tzitzikama Mtns., Maas Geesteranus 6624 pr. p. (L); Grahamstown, 
Albany, Héeg, December 1929 (TRH); Fern Kloof, Albany, Almborn 10734, 
10739 (LD); Skeleton Stream, east slopes of Table Mountain, Wynberg, Almborn 
11158, 11161 (LD); near upper Cableway Station, Table Mountain, Almborn 1731, 


1733, 1892, 1897 (LD); Table Mountain, Arnell 1009 (LD); Jonkershoek, Stel- 
lenbosch, Almborn 1968, 11397, 11401 (LD). 


73. Parmelia sancti-angelii Lynge, Ark. Bot. 13, no. 13:35. 1914. 
Parmelia pseudohyporysalea Asahina in Kihara, Faun. Fl. Nepal Himalaya 
54, fig. 19. 1955[?]. Type collection: Downside of Katunje Bazar, Nepal, 
Kihara (TNS, lectotype). 
Type collection: Colonia Santo Angelo, near Cachoeira, Rio 
Grande do Sul, Brazil, Malme s.n. (S, holotype). 
Thallus large, 10-20 cm. in diameter, loosely attached, light mineral 
eray, rather coriaceous; lobes rotund, 8-15 mm. wide, margins entire 
to broadly crenate, often ascending, ciliate, cilia 2-4 mm. long, 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 307 


sorediate, soralia linear, sorediate lobes often becoming involuted; 
upper surface dull, continuous to finely cracked with age, rarely 
sorediate submarginally; medulla white, frequently orange red near 
the lower cortex; lower side black and sparsely rhizinate, light brown 
to mottled ivory and naked in a broad zone along the margins. 
Apothecia rare, more or less adnate, disc imperforate; hymenium 65 yu 
high; spores 7-10 13-18 u, episporium 1 y» thick; pycnidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C+ rose, KC+ red, 
P—, pigmented medulla K+ purple, atranorine, gyrophoric acid, and 
(if pigmented) rhodophyscin present. 

Parmelia sancti-angelti is a very common pantropical, corticolous 
or less commonly saxicolous species which, surprisingly, has only one 
synonym. Rhodophyscin occurs in about half of the specimens, 
including the type of P. pseudohyporysalea but not the type of P. 
sancti-angelii. Parmelia rampoddensis Nyl. is externally very similar 
but contains alectoronic acid (C—, KC+). Parmelia permutata Stirt. 
has a different K— pigment. These three species are closely related 
and chemical tests are sometimes necessary for certain identification. 

Additional specimens examined: 


MEXICO: Hripata@o: Huasco, near Pachuca, Wood s.n. (F); VERACRUZ: 
Mirador, Sartorius s.n. (M), Purpus 215 (US); 9 km. east of Jalapa, Hale 19433 
(S, TNS, US), 21109 (US); northeast of Huatusco, Hale 19505 (DUKE, MSC, 
REN, US); Pugssuia: Xuchitl, Arséne 8236 (US); MicuoacAn: Morelia, Arséne 
4460 (US). GUATEMALA: Hotel Tzanjuyu, Panajachel, Gays.n. (F). COSTA 
RICA: Rio Torres, Pittier 5051 (M); vicinity of Cartago, Standley 35468 (US). 

HAITI: Citadel la Ferriore, Thomas 47 (MO, NY, US); between Petionville 
and Ft. Jacques, Thomas 36c (NY, US). DOMINICAN REPUBLIC: Guama, 
Santiago, Wetmore 3904 (MSC, US); vicinity of Constanza, La Vega, Imshaug 
23318 (MSC, US), Allard 16543 (US); north side of Constanza, Cordillera Central, 
Imshaug 23721 (MSC, US). JAMAICA: Without locality, Cummings 38 (NY). 

VENEZUELA: Miranpa: Corrada del Guayado, Dennis 1565 (K); Los 
Chorros, Dennis 1524A (K); Méripa: El Valle, Magdefrau 584 (M). COLOM- 
BIA: Cauca: La Capilla, Killip 38483 (US). CHILE: Cavurin: Pucon, 
Gunckel 18119 (MO). BRAZIL: Mato Grosso: Santa Anna da Chapada, 
Sladen 321 (BM); SXo Pavto: Piquete, Robert sn. (BM); Santa Albertina, 
Hemmendorf s.n.(S). ARGENTINA: Misiones: San Ignacio, Oniroga K54 bis 
(8). 

CONGO: Yangambi, Luweo Plateau, Louis 8155 p.p. (BR). KENYA: 
Rirt VaLutey Prov.: Eastern Mau Forest Reserve, Maas Geesteranus 11313 p.p. 
(L); Nyanza Prov.: Tinderet Forest Reserve, Kisumu-Londiani Distr., Maas 
Geesteranus 4956 p.p., 5532, 11160 (L). SOUTHERN RHODESIA: Zimbabwe, 
Hoéeg s.n. (TRH). UNION OF SOUTH AFRICA: Narat: Boschfontein Forest, 
Lions River Distr., Almborn 8701 (LD). MADAGASCAR: Tananarive, des 
Abbayes s.n. (REN, US); 30 km. north of Ankazoba, des Abbayes s.n. (REN, 
US). 

SIKKIM: Yokusam, Hara et al. s.n. (TNS, US). INDIA: Darjeeling, 
Hara et al. s.n. (TNS, US). THAILAND: Doi Chieng Dao, Tsuyama 12 (TNS, 
US). LAOS: Ban Phu Phao, Tsuyama 8 (TNS,US ); Hase Plantations, Tsuyama 9 


308 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


(TNS, US). SUMATRA: Raja Karohoogvlakte, Lérzing 7051 p.p. (BO); 
near Aer Batumbuk, Groenhart 8363 (BO); Pagar-alam, Steup 1024 (BO). 
CHINA: Kushan, near Foochow, Fukien, Chung F217 (FH, MICH). CELEBES: 
Rante Lemo, Kjellberg 33 (BO). JAVA: Mt. Gedeh, Tjibodas, Groenhart 8551 
(BO). 


74. Parmelia stuppea Tayl. London Journ. Bot. 6:175. 1847. PLATE 9 

Parmelia perforata var. claudelit Harm. Bull. Soc. Sci. Nancy, n.s. 14: 223. 
1896. Type collection: Docilles, France, Claudel, August 1890 (DUKE, 
holotype). 

P. maxima Hue, Nouv. Arch. Mus. Paris, ser. 4, 1:193. 1899. Type collec- 
tion: Chimaleapan, near Lerma, Mexico, Maury 3318 (P, holotype). 

P. claudelit (Harm.) Vain. Bot, Tidskr. 29:105. 1909. 

P. trichotera Hue var. claudelit (Harm.) Du Rietz, Nyt Mag. Naturv. 62:77. 
1924. 

P. claudelit (Harm.) Tavares, Port. Acta Biol., ser. B, 1:152. 1945. Super- 
fluous combination. 


Type collection: Monterey, California, Beechey (FH-Tayl, holotype; 
BM, K, isotypes). 

Thallus loosely attached to bark, 10-20 cm. in diameter, mineral 
gray; lobes rotund, 10-15 mm. wide, margins often ascending and 
suberect, sorediate, soralia terminal, linear, ciliate, cilia 2-3 mm. long; 
upper surface dull, continuous to cracked with age; lower side black 
and sparsely rhizinate, dark brown to brown and naked in a broad 
zone along the margins. Apothecia (known mostly from collections 
in western United States) very large, 10-30 mm. in diameter, stalked, 
disc perforate or rarely imperforate; hymenium 40-55 yu high; spores 
6-9 12-17 pu, episporium 1.0-1.5 » thick; pycnidia present, conidia 
4-6 pu long. 

Reactions: Thallus K+ yellow; medulla K+ yellow turning red, 
C—, KC—, P+ pale orange red, atranorine and salacinic acid present. 

Parmelia stuppea is represented by a very fragmentary type specimen 
from California. Although badly decomposed, it has marginal 
soredia, a few cilia, and salacinic acid. It is thus similar to modern 
collections of a species that has been called P. claudelii in Europe or 
P. trichotera in North America. It is common in the mountains of 
North America, central America, and Europe, and more rarely in 
Africa and Asia (fig. 21). It is especially common on oak trees in dry 
forests. It intergrades with P. cristifera Tayl. in that a sterile speci- 
men with sparse axial cilia could be identified as P. stuppea or as an 
abnormally ciliate form of P. cristifera. These two species in their 
typical forms are of course quite different in cilia and spore size. 
Parmelia stuppea may also be related to the Mexican species P. 
eurysaca Hue, which lacks soredia and has marginal laciniae. 

Additional specimens examined: 


U.S.: VerRMont: Scanlon Swamp, Leicester, Dutton 1620 (DUKE); Massa- 
CHUSETTs: 5 mi. west of Blanchard, Culberson 4564 (DUKE); Connecticut: 1 mi. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 309 


south of Colebrook, Litchfield Co., Hale 15406 (US); Detaware: Faulkland, 
Commons s.n. (PH); PENNSYLVANIA: 1 mi. east of Shunk, Sullivan Co., Hale 17442 
(US); 2 mi. north of Red Rock, Luzerne Co., Hale 16160 (US); Maryuanp: 1 mi. 
east of Oldtown, Alleghany Co., Hale 14492 (US); Wesr Virainia: Clear Creek, 
Raleigh Co., Hale 11773 (US); Panther Knob, Pendleton Co., Hale 14302 (US); 
near Mace, Pocahontas Co., Hale 12482 (US); Wisconsin: Park Falls, Price Co., 
Culberson 1818 (WIS); Viretnta: Mountain Lake Biological Station, Giles Co., 
Hale 18422 (US); Hawksbill Mountain, Madison Co., Hale 18884 (US); Hungry 
Mother State Park, Smyth Co., Hale 11931 (US); NortuH Carona: 3 mi. north 
of Buckcreek Gap, McDowell Co., Culberson 4864 (DUKE); southeast of Rainbow 
Springs, Macon Co., Anderson 12911 (DUKE); near Waynesville, Haywood Co., 
Standley 5784 (US); TENNESSEE: Cliff Springs, Overton Co., Phillips 314 (US); 
CauirorNIA: Pilarcitos Canyon, San Mateo Co., Cain 26381, 26384 (TRT, US); 
2 mi. east of Pt. Reyes Station, Marin Co., Koch 393 (US); between Gilvey and 
Salinas, San Benito Co., Herre s.n. (US). MEXICO: Veracruz: 11 km. east of 
Las Vigas, Hale 21117 (US); Puresia: 69 km. east of Puebla, Hale 19327 (US); 3 
km. west of Puebla-Veracruz state line on highway 150, Hale 19611 (S, TNS, US); 
Mexico: west of Rio Frio, Hale 19281 (US); MicnHoacdn: 61 km. northwest of 
Zitdcuaro, Hale 20838 (DUKE, MSC, REN, US); Cerro Azul, Morelia, Arséne 
3996, 4016 (US); Oaxaca: 53 km. northwest of Oaxaca, Hale 20814 (COLO, US); 
Cerro San Felipe, Hale 20694 (LISU, 8, TNS, US); Curapas: 14 km. west of San 
Cristébal, Hale 20543 (MSC, REN, US). GUATEMALA: HvuEHUETENANGO: 
Cumbre de la Sierra de los Cuchumatanes, Standley 81167 (MO). COSTA 
RICA: San José: Santa Maria de Dota, Standley 43176 (US). 

GERMANY: Munich, Arnold, Lich. Exs. 145 (MICH); Stuttgart, Wurttem- 
berg, Putzler sn. (LD). PORTUGAL: Serra de Sintra, Estremadura, Tavares 
4783 (US); Serra do Bucaco, Beira Litoral, Tavares 23 (H). 

UNION OF SOUTH AFRICA: Nara: Eschowe, south of Solheim, Héeg s.n. 
(TRH). SOUTHERN RHODESIA: Zimbabwe, Héeg s.n. (TRH). 

INDIA: Shembaganur, Madurai Distr., Awastht 4345 (AWAS). 


75. Parmelia subarnoldii des Abbayes, Mem. Inst. Sci. Madagascar, ser. B, 
10:113. 1961. 

Type collection: Forét de Manjakatompo, Ankaratra, Centre Nord, 
Madagascar, des Abbayes (REN, lectotype; US, isotype). 

Thallus loosely adnate, 8-12 cm. across, mineral gray; lobes rotund, 
7-10 mm. wide, margins often ascending, rather conspicuously ciliate, 
cilia 3-6 mm. long, sorediate, soralia, terminal, linear, sorediate lobes 
sinuous; upper surface plane, shiny, continuous; lower side black, 
sparsely rhizinate, brown, shiny, and naked in a broad zone along 
the margins. Apothecia and pycnidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC-+- reddish, 
P+ brick red, atranorine and protocetraric acid present. 

Parmelia subarnoldii is a rare pantropical species. Externally it is 
very close to P. sancti-angelii Lynge but differs significantly in 
chemistry. It has no relation to P. arnoldii Du Rietz, a temperate 
species with distinct sublaminal soredia. Extreme ciliate forms of 
P. dilatata Vainio, such as occur in the West Indies, are virtually 
identical with P. subarnoldit except that the soredia are coarse and 
irregular and the cilia erratically produced. 


310 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Additional specimens examined: 

MEXICO: Cutapas: 50 km. west of Tuxtla Gutiérrez, Hale 19904 (US), 19935 
(DUKE, REN, §, US). 

BRAZIL: Sao Pauto: Santa Albertina, Hemmendorff, July 3, 1898 (8). 

MADAGASCAR: Manjakatompo, Centre Moyen, Benoist 1355 (LD). 

NEW GUINEA: Mt. Arfak, Angi Gita Lake, Kostermanns 3012 (BO). JAVA: 
Tjinnivrang, Ogata 1001 (TNS). 


76. Parmelia subcorallina Hale, Journ. Jap. Bot. 37:345, fig. 1. 1962. 
P. proboscidea var. corallina Mill. Arg. Flora 67:616. 1884 (p.p.). 


Type collection: Keitau, Taichu, Formosa, Asahina 3312 (TNS, 
holotype; US, isotype). 

Thallus loosely adnate, 10-15 cm. in diameter, mineral gray; 
lobes rotund, 8-15 mm. wide, margins entire or subcrenate, in part 
isidiate-dissected, ciliate, cilia 1.5-3.0 mm. long; upper surface plane, 
continuous or reticulately cracked with age, faintly white-maculate, 
toward the margins moderately granulate-isidiate or isidiate, isidia 
cylindrical to inflated, simple or irregularly branched, 0.08-0.1 mm. 
in diameter, to 0.2 mm. high, often apically ciliate; lower side black 
and sparsely rhizinate, brown and naked in a broad zone along the 
margins. Apothecia not seen; pycnidia present, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC-+ reddish, 
P+ brick red, atranorine, protocetraric acid, and protolichesteric 
acid present. 

Parmelia subcorallina is distinguished by the chemistry and by the 
usually granular isidia, in part close to those of P. mellissit Dodge. 
Since the publication of this species, two additional records from 
Java have been found. It is apparently endemic to Asia and the 
Madagascar region. The photograph in my original publication is 
enlarged two times. 

Additional specimens examined: 


JAVA: Tjibodas, Mezjer 8928 (BO), van Woerden 2192 (BO). 
MAURITIUS: Robillard s.n. (G). SAO TOME: Moller 70 pp. (H). 


77. Parmelia subcrinita Nyl. Lich. Japon. 26. 1890. 


Type collection: Hirosaki, Japan, Almquist (H, Nyl. herb. no. 
35479, neotype). 

Thallus loosely adnate, 10-20 cm. broad, mineral gray; lobes ro- 
tund, 6-12 mm. wide, margins crenate, ciliate, cilia 1-2 mm. long; 
upper surface plane dull to shiny, at times faintly white-maculate, 
becoming white-pruinose towards the tips, reticulately cracked in 
older parts, moderately isidiate, isidia cylindrical, simple to coralloid- 
branched, to 0.08 mm. in diameter and 0.3 mm. high, often ciliate 
apically; lower side jet black and rhizinate, dark brown, shiny, and 
naked in a broad zone at the margins. Apothecia rather rare, 3-6 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA all 


mm. in diameter, disc imperforate; hymenium 65-75 yw high; spores 
8-10 12-14 y, episporium 1 » thick; pycnidia rare, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K+ yellow turning red, 
C—, KC—, P+ pale orange red, atranorine and salacinic acid 
present. 

The typification of P. subcrinita has presented some problems. Ny- 
lander cited a collection by Almquist from Shimonoseki, Japan, in his 
original description. This collection could not be found at Helsinki 
or Stockholm. We have therefore selected as a neotype another 
specimen from Japan identified in Nylander’s handwriting. Par- 
melia subcrinita is easily separated from P. crinita Ach. by chemical 
tests as well as by the smaller spore size, coarser cilia, and the more 
robust thallus. The upper cortex may be faintly maculate, as in the 
neotype, dull and opaque, or even pruinose. Reticulate fissuring with 
age may be pronounced. Another isidiate species with similar range 
and chemistry is P. subtinctoria Zahlbr., a species typically with dis- 
tinct maculae and a uniformly pale lower side, or if darkened, short 
rhizines or papillae to the margin below. 

Additional specimens examined: 


U.S.: West Viretnia: 3 mi. southwest of Jordan Run, Grant Co., Hale 14997 
(US); Wisconsin: Flambeau River State Forest, Sawyer Co., Hale 2053 (WIS); 
VirGINIA: 6 mi. east of Wallaceton, Norfolk Co., Ireland 4177 (US); Little Stony 
Man Cliffs, Page Co., Hale 15177 (US); Nortru Carona: 10 mi. east of Fayette- 
ville, Cumberland Co., Hale 17507 (US); 3 mi. north of Silver Valley, Davidson 
Co., Culberson 6446 (DUKE, US); 4 mi. east of Hayesville, Clay Co., Culberson 
5299 (DUKE); Sourn Carotina: Near Hodges, Greenwood Co., Culberson 7457 
(DUKE, US); Santee State Park, Orangeburg Co., Hale 16494 (US); 12 mi, 
southwest of Andrews, Georgetown Co., Hale 16555 (US); TENNESSEE: Cliff 
Springs, Overton Co., Phillips sn. (US); Grorata: Near McRae, Telfair Co., 
Hale 7573 (US); Toccoa, Stephens Co., Culberson 7346 (DUKE); AtaBama: 15 
mi. southeast of Birmingham, Shelby Co., McCullough 346 (US); Cheaha State 
Park, Cleburne Co., Hale 7086 (US); 2 mi. west of Cedar Cove, Tuscaloosa Co., 
McCullough 404 (US); Frioripa: 1 mi. east of Monticello, Jefferson Co., Hale 
16461 (US); Sanford, Seminole Co., Rapp 495 (FLAS); Suwanee River State 
Park, Suwanee Co., Hale 17650 (US); Mrssissipp1: Near Morton, Scott Co., 
Hale 7923 (US); near Durant, Holmes Co., Hale 7892 (US); near Tupelo, Lee Co., 
Hale 7830 (US); Arxansas: Near Hollis, Perry Co., Hale 3129 (US); Buffalo River 
Park, Marion Co., Hale 4483 (US); Oxtauoma: Near Broken Bow, Hale 4925 
(US); near Ludlow, Le Flore Co., Hale 5044 (US). MEXICO: Veracruz: 15 
km. south of Catemaco, Hale 19850 (US); Mirador, Liebmann 117 (UPS); 
Micuoackn: Rincén, Morelia, Arséne 4407 (US); Cotrma: Isla Socorro, Herrera 
17 (MSC); Oaxaca: Cuyamecalco, Conzatti 3496 (US); Cutaras: 8 km. east of 
Teopisca, Hale 20335 (S, US); Hacienda, Matuda s.n. (MICH). HONDURAS: 
MorazAn: Above El Zamorano, Standley 336 (MO). 

DOMINICAN REPUBLIC: Along Rfo Inoa, at Inoa, Imshaug 23861 (MSC, 
Us). JAMAICA: Clydesdale, Plittsn.. (BPI, US). MARTINIQUE: Caravelle, 
Degelius s.n. (DEGEL). 

VENEZUELA: Northeast of Puerto Ayacucho, Orinoco, Mdgdefrau 115 
(M, US). COLOMBIA: Norte pre Santanpver: La Cabuya, Cuatrecasas 


729—-018—65——_10 


312 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


12057 (US). ECUADOR: Gatapacos IsLtanps: Santa Maria Island, Taylor 
868 (MICH). BRAZIL: Baura: Serra Chugue, Luetzelburg 408 (M); Minas 
Gerais: Caldas, Regnell s.n. (S). 

AZORES: St. Jorge, Persson s.n. (UPS). 

JAPAN: Nirayama, Prov. Idzu, Asahina 56105 (TNS, US), Kurokawa 58009 
(TNS, US). JAVA: West Bantam, Groenhart 9740 (BO); Gegerbentang, 
Neervoort 1187 (BO), NEW CALEDONIA: Nepouis Valley, Williams 14 
(BISH, US). 

78. Parmelia sulphurata Nees & Flot. Linnaea 9:501. 1835. 

Parmelia persulphurata Nyl. in Cromb. Journ. Linn. Soc. Lond. 16:219. 
1877. Type collection: Bahia, Brazil, Crombie (BM, holotype). 

P. brisbanensis Stiiton, Trans. Proc. Roy. Soc. Victoiia 17:69. 1881. Type 
collection: Brisbane, Queensland, Australia, Bailey 228 (BM, holotype; 
GLAM, isotype.) 

Type collection: Cuba, Wright 72 (UPS, neotype; BM, FH, K, 
M, US, isotypes). 

Thallus adnate to loosely attached to bark, 6-10 cm. in diameter, 
yellowish to turtle green; lobes rotund, 5-8 mm. wide, soon irregularly 
branched, margins entire to dissected, ciliate, cilia variable, 1-4 mm. 
long; upper surface plane, dull, reticulately cracked in older parts 
(with yellow medulla showing through), sparsely to densely isidiate, 
isidia cylindrical, simple to branched, 0.03-0.05 mm. in diameter, 
to 3 mm. high; medulla deep lemon yellow; lower side black and 
rhizinate at the center, naked and brown in a broad zone along the 
margins. Apothecia and pycnidia not seen. 

Reactions: Thallus K-++ more intensely yellow; medulla K—, C—, 
KC—, P—, atranorine and vulpinic acid present. 

This species has been correctly identified by most lichenologists 
because of the brilliant lemon-yellow medulla. Nylander, however, 
consistently called it P. persulphurata Nyl. and identified specimens 
of P. endosulphurea (Hillm.) Hale as P. sulphurata. Parmelia 
endosulphurea has different yellow pigments and lacks cilia. The 
Péppig type of P. sulphurata, destroyed at Berlin, had been seen 
earlier by Hillmann (1939), who was able to confirm that the current 
concept of the species is correct. I have therefore selected as a 
neotype a widely distributed exsiccate from the type locality. The 
species is very common in the Caribbean, though not nearly as 
common as P. endosulphurea, and occurs widely in other tropical 
areas (fig. 22). 

Additional specimens examined: 

U.S.: Frortpa: 5 mi. south of Silver Glenn Springs, Marion Co., Hale 17579 
(US); Eldorado, Lake Co., Underwood 2313 (FH); Sanford, Seminole Co., Rapp, 
Merr. Lich Exs. 76 (US); Lourstana: Grand Bayou, Langlois 42 (US). MEXICO: 
CaMPECHE: Tuxpefia, Lundell 1303 (F); Yucarsn: Tizimin, Swallen 2538 


(MICH). HONDURAS: Comayagua: Comayagua, Standley 5829 (F). 
PANAMA: Canau Zone: Barro Colorado Island, Scholander s.n. (US). 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 313 


CUBA: Pinar pvEL Rfo: Northwest of Hotel San Vicente, Imshaug 25235 
(MSC); San Diego de los Bafios, Earle 275 (FH). DOMINICAN REPUBLIC: 
Consuelo, Taylor 152 (FH); north of San José de los Matas, Wetmore 3878 
(MSC); Piedra Blanca, La Vega, Allard 16856 (US). 

FRENCH GUIANA: Belizon, River Combé, Degelius s.n. (DEGEL). 
SURINAM: Without locality, s.c. (G, M). VENEZUELA: Cerro Pavén, 
Rio Atabapo, Mdgdefrau 285 (M). PERU: Loreto: Santa Maria, Allard 22470 
bis (US). 

IVORY COAST: 5 km. north of Abidjan, Santesson 10325 (UPS); 6-7 km. 
north-northeast of Tai, cercle de Man, Santesson 10463b (UPS). CONGO: 
Luweo Plateau, Yangambi, Louis 8172 (BR). REPUBLIC OF CONGO: 
Subluali, Maiumbe, Gossweiler 8136 (BM). 

SUMATRA: Steup s.n. (BO). 


79, Parmelia wainii A. L. Smith, Journ. Linn. Soc. London, Bot. 46:85. 1922. 
Parmelia proboscidea var. ornatula Zahlbr. Bull. Herb. Boiss., Ser. 2, 4:135. 
1904. Type collection: Serra do Ouro Preto, Brazil, Damazio 1090 

(W, holotype; G, isotype). 

Type collection: Caraga, Minas Gerais, Brazil, Vainio, Lich. 
Bras. Exs. 400 (TUR, Vain. herb. no. 2410, lectotype; BM, FH, K, 
M, UPS, isotypes). 

Thallus adnate to loosely attached to bark, 5-15 cm. in diameter, 
mineral gray; lobes rotund, 8-12 mm. wide, margins smooth or 
irregularly digitate-lobulate and crowded in older parts, ciliate, 
cilia 2-5 mm. long; upper surface plane, dull, continuous or becoming 
reticulately cracked with age, isidia and soredia lacking; medulla 
white, rarely turning orange red near the lower cortex; lower side 
black, sparsely rhizinate, dark brown and naked or mottled tan in a 
broad zone along the margins. Apothecia common, stipitate, to 
10 mm. in diameter, exciple entire to short-dentate, ciliate, amphi- 
thecium rugose, white-maculate, disc imperforate; hymenium 75-— 
90 » high; spores 6-1017-22 yu, the episporium 1.5-2.0 » thick; 
pycnidia common, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ red, 
P—, pigmented medulla K+ purple, atranorine, alectoronic acid, 
and (if pigmented) rhodophyscin present. 

A. L. Smith was familiar with P. proboscidea Tayl., a synonym 
of P. crinita Ach., because of her work in British lichens. When 
she saw Vainio’s erroneous determinations of nonisidiate South 
American plants as P. proboscidea, she proposed a new name, P. 
wainit, for his misidentified material. Parmelia wainii is character- 
ized by large rugose ciliate apothecia and conspicuously ciliate lobes. 
Parmelia subrugata Kremplh. differs in having large spores, shorter 
cilia, and more richly branched, laciniate lobes. Parmelia argentina 
Kremplh. differs in having a white rim below and more or less distinct 
maculae. Parmelia wainii is not common but has a characteristic 
South American-South African distribution pattern. 


314 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Additional specimens examined: 


BRAZIL: Minas GeErats: Sitio, Vainio, Lich. Bras. Exs. 582b (TUR). 
NIGERIA: Basse, Dusén, Almb. Lich. Afr. 31 (H, LD, UPS, US). CONGO: 
Crete de Mvakala, Devred 1368 (BR). MOCGAMBIQUE: Lourengo Marques, 
Junoa 472 (PRE). UNION OF SOUTH AFRICA: Narat: Ngoma, Héeg s.n. 
(TRH). 
Subsection Ornaticolae Gyel. (1932, p. 225) 


Thallus loosely adnate to suberect, 5-30 cm. in diameter; lobes 
broad, 5-20 mm. wide, the margins ciliate; upper cortex distinctly 
maculate; lower side brown or black with a brown or white marginal 
zone. 

Type species: Parmelia perforata (Jacq.) Ach. 

This subsection may be divided into two series: Subpallidae, the 
thallus with a pale entirely rhizinate lower side, and Ornaticolae, with 
a black center. 

Series Subpallidae, ser. nov. 


Thallus subtus ex toto pallidus, breve rhizinosus usque ad marginem, 
ceterum ut in subsectione Ornaticolae. 

Type species: P. subsumpta Nyl. 

This series of only three species seems to be unrelated to any other 
group in subgenus Amphigymnia. The species are closely related 
and appear to present a sorediate-isidiate-NIS series with a parallel 
development of chemical strains. 

80. Parmelia subcaperata Kremplh. Nat. For. Kjoeb. Vid. Medd. 1873: 10. 

1873. PLaTE 12 

Parmelia erubescens Stirton, Scot. Nat. 4:201. 1877-78. Type collection: 
Near Brisbane, Australia, Bailey (BM, holotype). 

Parmelia recipienda Nyl. Flora 68:609. 1885. Type collection: Brazil, 
s.c. (H, Nyl. herb. no. 35212, holotype). 

Parmelia imperforata Nyl. Acta Soc. Sci. Fenn. 26, no. 10:7. 1899. Type 
collection: Brazil, Glaziou 1839 (H, Nyl. herb. no. 35425, holotype). 

Parmelia annae Lynge, Ark. Bot. 13, no. 13:88, pl. 2, fig. 6. 1914. Type 
collection: Santa Anna da Chapada, Mato Grosso, Brazil, Malme 2368B 
(S, lectotype). 

P. ceracea Lynge, Ark. Bot. 13, no. 13:97. 1914. Type collection: Pil- 
comayo, Gran Chaco, Paraguay, Malme, Sept. 7, 1893 (S, lectotype). 


Type collection: Lagoa Santa, Serra da Piedade, Brazil, Warming 
297 (M, holotype). 

Thallus loosely attached, 5-12 cm. in diameter, turning buff in 
the herbarium; lobes rotund, 7-15 mm. wide, margins crenate, ciliate, 
cilia 1.0-3.0 mm. long, often branched; upper surface plane, shiny, 
strongly white-maculate, becoming cracked with age; lower side 
uniformly pale tan or darkening toward the center, densely short 
rhizinate to the margins. Apothecia abundant, to 30 mm. in diameter, 
stalked, disc perforate; hymenium 55-65 yw high; spores 6-10 12-19 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 315 


BM, episporium 1.0-1.5 yw thick; pycnidia abundant, conidia 10-13 
u long. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ red, 
P—, atranorine, cryptochlorophaeic acid, and protolichesteric acid 
present; or K-+ yellow turning red, C—, KC—, P+ pale orange 
red, atranorine and salacinic acid present. 

Parmelia subcaperata is characterized by the strongly maculate 
upper cortex, short rhizinate pale lower side, and the lack of soredia 
or isidia. It has the same chemical variation as P. subsumpta Nyl., 
a sorediate counterpart, but the distribution is more restricted. The 
types of P. recipienda, P. annae, and P. ceracea contain cryptochloro- 
phaeic and protolichesteric acids, while the remaining types listed 
above all contain salacinic acid. Except for a single collection in 
Australia, the species is known only from South America. 

Additional specimens examined: 

Salacinic acid present: 

BRAZIL: Minas Gerats: Sitio, Vainio, Lich. Bras. Exs. 737 (TUR, UPS); 
Retiro, Itatiaia, Dusén, Lich. Austroamer. 82, 86 (H, LD, MSC, S, US); Carangola, 
Mexia 4322 (MO, US); Lagoa Santa, Warming 313 (M); Serra dos Orginos, 
Ainsworth & Gregory 553 (BM); Rio pE JANERIO: Rio de Janeiro, Glaztou 1843, 


2001 (M), Malme 372 (8), Fry s.n. (BM); SAo0 Pauto: Piquete, Robert s.n. (BM); 
Rio GRANDE Do Sut: Pareci Nova, Lima 48 (MO). 


Cryptochlorophacic acid present: 


BRAZIL: Minas Gerais: Caldas, Henschen s.n. (UPS); S&0 Paulo: Piquete, 
Robert s.n. (BM). PARAGUAY: Asuncién, Malme 1678 * * * (8); Pilcomayo, 
Gran Chaco, Malme s.n. (MO). ARGENTINA: Jusuy: Quinta, Fries 27 p.p. 
(S); Satta: Urundel, Ordn, Grassi 594 (MO); Formosa: Las Lomitas, Pierotti 
2896 (MO). 


81. Parmelia subsumpta Nyl. Flora 52:117. 1869. PLATE 12 

Parmelia urceolata Eschw. var. nuda Mill. Arg. Flora 63:266. 1880. Type 
collection: Petropolis, Brazil, Deventer 33 (G, holotype). 

P. hypotropa var. imperialis Hue, Nouv. Arch. Mus. Paris, ser. 4, 1:189. 
1899. Type collection: Brazil, S.M. Theresa Christina, 1887 (P, holotype). 

P. petropoliensis Zahlbr. Sitzungsber. Akad. Wiss. Wien Math. Naturw. 
111:426. 1902. Type collection: Petropolis, Brazil, Héhnel 177 (W, 
holotype). 

P. corrugis (Fr.) Mill. Arg. var. imperialis (Hue) Zahlbr. Cat. Lich. 
Univ. 6:236. 1929. 

P. leucozantha Mill. Arg. f. firma Ceng.-Samb. Annali Bot. 22, no. 1:18. 
1939. Type collection: Serra dos Orgéos, Rio de Janeiro, Brazil, Casaretti 
2444 (RO, holotype). 


Type collection: Minas Gerais, Brazil, Glaziou sn. (H, Nyl. herb. 
no. 35451, holotype). 

Thallus loosely adnate to bark, up to 15 cm. broad, rather coriaceous, 
mineral gray turning buff in the herbarium; lobes rotund, 10-12 mm. 
wide, margins often ascending, subcrenate, sorediate, soralia elongate, 


316 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


in older specimens becoming crescent-shaped, sorediate lobes involute, 
ciliate, cilia 1.0-1.5 mm. long; upper surface plane, shiny, strongly 
white-maculate, reticulately cracked with age; lower side uniformly 
pale to dark brown, rarely blackening, densely short rhizinate to the 
margin or with a narrow papillate zone along the margins. Apothecia 
rare, 10-13 mm. in diameter, stalked, amphithecium rugose, white- 
maculate, sorediate, disc imperforate; hymenium 65-75 yw high; 
spores 7-10 12-15 yu, episporium 1.0-1.5 »; pycnidia not seen. 

Reactions: Thallus K+ yellow; medulla K+ yellow turning red, 
C—, KC—, P+ pale orange red, atranorine and salacinic acid pres- 
ent; or medulla K—, C—, KC+ red, P—, atranorine, crypto- 
chlorophaeic acid, and protolichesteric acid present. 

Parmelia subsumpta is a widespread corticolous species in tropical 
America and Africa. The distinguishing features are the strongly 
maculate upper cortex and short rhizinate, pale lower side. The 
soralia are linear and terminal but often enlarge and may become 
involute or even crescent-shaped, as in the holotype of P. subsumpta. 
The lower side may blacken somewhat so as to resemble P. lewcose- 
motheta Hue, which has a broad naked zone below. There are at 
least two chemical strains in P. subsumpta. The most common by 
far (including all the types listed) contains salacinic acid, rarely with 
traces of usnic acid. The second strain with protolichesteric and 
eryptochlorophaeic acids occurs in Nebraska, Brazil, and Peru. 
Since there is similar chemical variation in other species in this series, 
we hesitate to recognize two separate species. 

Additional specimens examined: 

Salacinic acid present: 

U.S.: Georara: Vogel State Park, Union Co., Hale 7325 (US). MEXICO: 
Pursua: 3 km. west of Puebla-Veracruz state line on highway 150, Hale 19653 
(US); Oaxaca: 32 mi. northwest of Oaxaca, Cain 27566d (TRT, US); CHIAPAS: 
14 km. west of San Cristébal, Hale 20568 (TNS, US); south of Teopisca, Hale 
20509 (DUKE, US); El Sumidero Canyon, Tuxtla Gutiérrez, Hale 21044 (US); 
50 km. west of Tuxtla Gutiérrez, Hale 19957 (US). GUATEMALA: Hve- 
HUETENANGO: Rio Pucal, Standley 82404 (MO); Basa Verapaz: Above Santa 
Rosa, Standley 91074 (MO). PANAMA: Curriqui: Llano del Volcano, Scho- 
lander s.n. (MO, US). 

VENEZUELA: Avila, Vogl s.n. (M). BRAZIL: Minas Gerats: Lagoa 
Santa, Warming 307 (M); Sio Pauto: Sao Paulo, Hoehne s.n. (M); Rio GRANDE 
po Suu: Near Cachocira, Santo Angelo, Malme 908 (S). ARGENTINA: With- 
out locality, Lorentz & Hieronymus (M). 

CONGO: Biano, Smitz 3884 (BR). KENYA: Tinderet Forest Reserve, 
Kisumu-Londiani Distr., Maas Geesteranus 11120 (LD, L). UNION OF SOUTH 
AFRICA: Transvaau: Northeast of Haenertsburg, Kapp 1773 (PRE); Natvau: 
Polela Forest, Polela, Almborn 9506 (LD); Carg Province: 6 mi. from Knysna, 
Almborn 2618 (LD). 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 317 


Cryptochlorophaeic and protolichesteric acids present: 


U.S.: Nepraska: Sandhills, Nebraska National Forest, Henzlik s.n. (MSC). 
PERU: Cuzco: Urubamba, Vargas 7274 (MICH, US). BRAZIL: Sanra 
CaTARINA: Rio dos Bugres, Cagador, Reitz 13215 (US). 


82. Parmelia subtinctoria Zahlbr. Symb. Sin. 3:193. 1930. PLATEs 2, 12 

Parmelia proboscidea var. aspera Mill. Arg. Flora 69:258. 1886. Type 
collection: Caracas, Venezuela, Ernst 71 (G, lectotype). 

P. virens f. isidiosa Mill. Arg. Ann. Nat. Hofmus. Wien 7:303. 1892. Type 
collection: Toowoomba, Queensland, Hartmann 58 (G, holotype; W 
isotype). 

P. leucosemotheta Hue f. isidiata Hue, Nouv. Arch. Mus. Paris, ser. 4, 1:192. 
1899. Type collection: Ohio, U.S.A., Lesquereux 262 (P, holotype). 

P. velutina Zahlbr. Ann. Crypt. Exot. 1:206. 1928. Type collection: Mt. 
Tjibodas, Java, van Overeem 89 (W, holotype) [non P. velutina (Ach.) 
Wallr. Fl. Crypt. Germ. 3:552. 1831]. 

P. protovirens Gyel. Repert. Sp. Nov. Fedde 29:289. 1931. Based on P. 
virens f. isidiosa Mill. Arg. 

P. haitiensis Hale, Bryol. 62:20, fig. 2. 1959. Type collection: Blue Moun- 
tains, Jamaica, Orcutt 2987 (US, holotype). 


? 


Type collection: Sanyingpan, north of Yunnanfu, Yunnan, China, 
Handel-Mazzetti 5645 (WU, holotype). 

Thallus adnate to loosely attached, 5-10 cm. in diameter, mineral 
gray but soon turning buff in the herbarium; lobes rotund, 10-15 mm. 
wide, margins crenate, sparsely ciliate, cilia 1.0-1.5 mm. long; upper 
surface plane, shiny, distinctly white-maculate, irregularly cracked 
with age, moderately to densely isidiate, isidia cylindrical, simple to 
branched, 0.04-0.07 mm. in diameter, to 1.0 mm. high, rarely ciliate 
apically; lower side uniformly pale brown, rarely darkening at the 
center, densely short rhizinate to the margin or with a narrow naked 
or papillate zone along the margins. Apothecia rare, 5-8 mm. in 
diameter, amphithecium isidiate, disc imperforate; hymenium 40-60 
» high; spores 5-8><8-11 yw, episporium 1 yu thick; pycnidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C— KC+ red, 
P—, atranorine, cryptochlorophaeic acid, and protolichesteric acid 
present; or medulla K+ yellow turning red, P+ pale orange red, 
atranorine and salacinic acid sometimes mixed with protolichesteric 
acid. 

Parmelia subtinctoria is a widespread pantropical species (fig. 24) 
usually found in herbaria under the names P. crinita Ach. or P. 
suberinita Nyl. Parmelia crinita differs in lack of maculae, large 
spores, black lower side, and the presence of stictic acid. Parmelia 
suberinita lacks distinct maculae and has a black lower side. The 
chemical variation presents an interesting pattern. All specimens 
from the Americas that have been tested (about 50) contain only 


318 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


salacinic acid or cryptochlorophaeic acid and protolichesteric acid. 
Those from Asia, including the holotypes of P. subtinctoria and P. 
velutina, contain a mixture of salacinic, cryptochlorophaeic, and 
protolichesteric acids. Parmelia haitiensis (and P. proboscidea var. 
aspera) contains cryptochlorophaeic and protolichesteric acids and 
represents the type of the K—-, KC+ strain. ‘These variants can be 
considered as no more than chemical strains at this time. 

Additional specimens examined: 

Salacinic acid present (with or without protolichesteric) : 


U.S.: West Virainia: 2 mi. north of Brandywine, Pendleton Co., Hale 18543 
(US); 3 mi. northeast of Alvon, Greenbrier Co., Hale 16649 (US); near Huttons- 
ville, Randolph Co., Hale 12493 (US); Kenrucxy: Near Oil Valley, Wayne Co., 
Hale 13360 (US); near Forest Cottage, Cumberland Co., Hale 13723 (US); 
Pennyrile State Forest, Christian Co., Hale 13217 (US); Intinors: Cadiz, Hardin 
Co., Hale 13947 (US); Iowa: Fayette Co., Fink s.n. (US); Missourr: Near 
Radical, Stone Co., Hale 2549 (US); Aurora, Lawrence Co., Hale 2516 (US); 
Hartville, Wright Co., Hale 4290 (US); Virernta: Mile 102, Blue Ridge Parkway, 
Bedford Co., Hale 19176 (US); Signal Knob Overlook, Warren Co., Hale 14844 
(US); near Bane, Giles Co., Hale 12754 (US); Norru Carotina: Near Bynum, 
Chatham Co., Culberson, Vézda Lich. Sel. Exs. 70 (H, LD, US); near Oxford, 
Granville Co., Culberson 6520 (DUKE, US); 11 mi. north of Burlington, Alamance 
Co., Hale 16364 (US); TENNESSEE: Cliff Springs, Overton Co., Phillips s.n. (US); 
Backbone Mountain Recreation Area, Johnson Co., Hale 18391a (US); GEorata: 
Near Cleveland, White Co., Hale 7457 (US); AtaBama: Pell City, St. Clair Co., 
Hale 7235 (US); 15 mi. southwest of Rockford, Coosa Co., McCullough 1025 
(US); Fiortpa: Sanford, Seminole Co., Rapp s.n. (FLAS, US); Mississippi: 
Holly Springs, Marshall Co., Hale 7841 (US); Louistana: Near Shreveport, 
Caddo Parish, Hale 5620 (US); ArKANsas: 8 mi. south of Hollis, Perry Co., 
Hale 3095 (US); Crystal Springs, Garland Co., Hale 3050 (US); Danville, Yell Co., 
Hale 3734 (US); Black Springs, Montgomery Co., Hale 3896 (US); OkLAHOMA: 
Beaver Bend State Park, McCurtain Co., Hale 4929 (US); north of Sallisaw, 
Sequoyah Co., Hale 5030 (US); Texas: Near Milano, Milam Co., Hale 5471 
(US); near Avinger, Cass Co., Hale 5265 (US); Kansas: Batesville, Woodson Co., 
Hale 4676 (US); near Coyville, Wilson Co., Hale 4728 (US); Artzona: Cave 
Creek, Chiricahua Mountains, Cochise Co., Weber 8787 (COLO, US). MEXICO: 
Hipaua@o: Jacala, Chase 7431 (F, US); Veracruz: 15 km. south of Catemaco, 
Hale 19859 (US); Purnia: 3 km. west of Puebla-Veracruz state line, on highway 
150, Hale 21043 (US); Mexico: North of Acambay, Cain 27593 (TRT, US); 
MicHoacan: 61 km. northwest of Zitdcuaro, Hale 20862 (MSC, US); Oaxaca: 
132 km. northwest of Oaxaca on highway 190, Hale 20826 (DUKE, MSC, REN, 
S, TNS, US); Curapas: El Sumidero, Tuxtla Gutiérrez, Hale 20013 (US), south 
of Teopisea, Hale 20503 (COLO, US). GUATEMALA: Ciudad Vieja, Tejada 
s.n. (FH, US). HONDURAS: Comayaeua: Siguatepeque, Standley 6355 (F). 

CUBA: OrrentTE: Slope of El Gato, Loma del Gato, Imshaug 24746, 24798 
(MSC). HAITI: Eastern end of Montagne Noire, Imshaug 22529 (MSC). 

PARAGUAY: Rio Apa, Colonia Risso, Malme s.n. (S). 

KENYA: Rirr Vatitey Prov.: Cherangani, Maas Geesteranus 4711 (L); 
Nyanza Prov.: Londiani, Kisumu-Londiani, Maas Geesteranus 10935 (L); 
Tinderet Forest Reserve, Maas Geesteranus 11091 (LD, L). SOUTHERN 
RHODESIA: Zimbabwe, Héeg s.n. (TRH), Sim s.n. (BM). SWAZILAND: 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 319 


Piggs Peak, Almborn 7902 (LD). UNION OF SOUTH AFRICA: Transvaat: 
Punch Bowl Inn, north of Louis Trichardt, Almborn 6431 (LD, US); Houtbosch, 
Pietersburg, Almborn 6767 (LD); Nara: Boschfontein Forest, Lions River 
Distr., Almborn 8712 (LD); Indumeni Forest, Cathedral Peak Area, Almborn 
8874 (LD); Natal Table Mountain, Almborn 8577 (LD); Emmersdale, Estcourt, 
Héeg s.n. (TRH); Cape Province: Matatiele, Héeg s.n. (TRH); northern part 
of Swellendam, Almborn 2155 (LD); Stormsrivier, Humansdorp, Almbern 4127 
(LD, US); Keurboomsrivier, Almborn 2945 (LD); Parks Station, Knysna, Arnell 
1500a (LD); 3 mi. east of Hermanus, Caledon, Almborn 5727 (LD). 

INDIA: On way to Kasardeir, Almora Distr., Awasthi 3449 (AWAS). 
CHINA: Near Foochow, Kushan, Fukien, Chung F455 (FH). JAPAN: Honryu 
Temple, Nirayama, Prov. Idu, Kurokawa 58008 (TNS, US); Mt. Yatsugadake, 
Prov. Shinano, Asahinas.n. (TNS, US). 

SUMATRA: Panprama Tabat Patah, Groenhart 9214 (BO). JAVA: Mt. 
Gedeh, van Overeem s.n. (MICH); Tjibodas, van Woerden 5493 (BO); Treanger, 
Bakhutzen 1518 (BO). 


Cryptochlorophaeic and protolichesteric acids present (without 


salacinic acid): 

U.S.: Vireinia: Brokenburg, Spotsylvania Co., Hale 15709 (US); Nortu 
Carouina: Darden, Martin Co., Culberson 6561 (DUKE); near Red Springs, 
Robeson Co., Culberson 7627 (DUKE); Misstssipr1: Near Aderman, Choctaw 
Co., Hale 7802a (US); Arkansas: Cove Lake, Logan Co., Hale 3452 (US); 
MEXICO: Veracruz: Northeast of Huatusco, Hale 19478 (US); Pursia: 3 km. 
west of Puebla-Veracruz state line on highway 150, Hale 19616 (US); CuHrapas: 
50 km. west of Tuxtla Gutiérrez, Hale 19921 (US). 

HAITI: Summit of Montagne Noire, Wetmore 2722, (MSC), 2735 (MSC, US); 
near Boutillier, Morne Hospital, Imshaug 22514 (MSC). JAMAICA: Blue 
Mountains, Orcutt 2987 (US), Imshaug 14789 (MSC, US). 

Additional records from the United States and the West Indies listed by Hale 
(1959a) as P. haitiensis are not repeated here. 


Series Ornaticolae 


Thallus black below in the center, the marginal rim commonly 
white or mottled, naked. 

This is the most highly evolved group in subgenus Amphigymnia. 
Of the 23 species, only three, P. hababiana, P. hypotropa, and P. 
subrugata, may be considered pantropical. Sixteen of the species 
occur only in America or Africa or in both areas. There are two 
subgroups that may be recognized. One is characterized by a dis- 
tinct white rim below along the margins and suberect lobes. It in- 
cludes P. abessinica, P. argentina, P. chiapensis, P. euneta, P. hababi- 
ana, P. hypotropa, P. melanothriz, P. paulensis, P. perforata, P. rigida, 
P. subrugata, and P. uruguensis. Most of these species are centered 
in tropical America. 

A second group has a dark marginal rim and more or less adnate 
lobes. It includes P. cooperi, P. hanningtoniana, P. lobulascens, P. 
natalensis, P. nilgherrensis, P. pseudonilgherrensis, P. schimperi, and 
P. subcolorata, all of which occur only in Africa or Asia. 


320 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


The remaining species, P. coralliformis, P. leucosemotheta, and P. 
reparata, have very strong, almost effigurate maculae and occur chiefly 
in Mexico. 

These three groups seem quite distinct, but relegating them to 
subseries would unnecessarily fragment the proposed subgeneric 
classification at this time. 


83. Parmelia abessinica Kremplh. Linnaea 41: 140. 1877. 

Parmelia glaucocarpa Mill. Arg. Flora 67: 615. 1884. Later homonym of 
P. glacucocarpa Ach. (=Acarospora). Type collection: Nossi-Bé, Mada- 
gascar, Hildebrandt 2150 (G, holotype; FH, FLAS, M, WU, isotypes). 

P. abyssinica Nyl. Flora 68: 608. 1885. Superfluous name based on P. 
abesstnica Kremplh. 

P. hildebrandtii var. ciliata Mill. Arg. Bot. Jahrb. Engler 20: 255. 1894. 
Type collection: Marangu Station, Kilimanjaro, Tanganyika, Volkens 275 
(G, lectotype; BM, isotype). 

P, abyssenica [sic] var. glabrior Stein. & Zahlbr. Bot. Jahrb. Engler 60: 526. 
1926. Type collection: Between Mazumbai and Mzinga, West Usambara, 
Brunnthaler s.n. (WU, holotype). 

P. glaucocarpoides Zahlbr. Cat. Lich. Univ. 6:167. 1929. Based on P. 
glaucocarpa Mill. Arg., non Ach. 

P. nigeriensis Dodge, Ann. Mo. Bot. Gard. 46: 156. 1959. Type collection: 
Mongu Forest Reserve, Panshin, Plateau Province, Nigeria, Keay & King 
37096 (K, holotype). 

Type collection: Ethiopia, Hildebrandt s.n. (M, holotype). 

Thallus adnate to loosely attached, often ascending, to 10 cm. in 
diameter, mineral gray, turning olivaceous in the herbarium; lobes 
rotund, 6-10 mm. wide, margins subcrenate to more or less lobulate 
with age, sparsely to moderately ciliate, cilia 0.5-2.0 mm. long; upper 
surface smooth, more or less distinctly white-maculate, rugulose and 
cracked with age; lower side black and sparsely rhizinate, brown to 
ivory or mottled and naked in a broad zone along the margins. 
Apothecia common, stipitate, 5-13 mm. in diameter, amphithecium 
rugose, white-maculate, exciple variable, entire or dentate and ciliate, 
dise perforate; hymenium 70-80 » high; spores variable, 9-12 X 18-28 un, 
episporium 1.5-2.0 » thick; pyenidia common, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC—, P-, 
atranorine and protolichesteric acid present; or medulla KC-+ red, 
atranorine, protolichesteric acid, and cryptochlorophaeic acid present. 

Parmelia abessinica is the nonsorediate counterpart of P. hababiana 
Gyel. and has a similar geographical distribution and chemical vari- 
ation, with two strains. The cryptochlorophaeic acid strain (includ- 
ing all types listed above except P. glaucocarpa) is less common but 
has a distribution similar to the protolichesteric acid strain. Spore 
size is intermediate and rather variable. The exciple is ciliate in less 
than half the specimens, and there is no apparent correlation of the 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 321 


cilia with chemistry or distribution. Parmelia abessinica occurs on 
twigs and small branches of deciduous trees in open dry woods. 
Additional specimens examined: 
Cryptochlorophaeic and protolichesteric acids present: 
MEXICO: Curapas: El Zapotal, Tuxtla Gutiérrez, Hale 19990 (US). 
ETHIOPIA: North of Debra Marcos, Chokke Mountains, Lythgoe L46, L47 


(K). CONGO: Southwest side of Lake Kivu, Mulungu, South Kivu, Degelius 
s.n. (DEGEL, US). ANGOLA: Bré: Chinguar, Degelius s.n. (DEGEL, US). 


Protolichesteric acid alone present: 


ETHIOPIA: Uoché, Baldrati s.n. (US). CONGO: Bangu, road to Thysville, 
Compére 1149 (BR, US); Mvakala Crater, Devred 1368 (BR); Lobito, Quesyiere 
904 (BR); Lukuga Valley west of Albertville, Héeg s.n. (TRH). KENYA: Ny- 
ANZA Prov.: Tinderet Forest Reserve, Kisumu-Londiani, Maas Geesteranus 4956 
(L, LD). UGANDA: Mt. Eglon, Small J19 p.p. (K). ANGOLA: Huta: Qui- 
lengues, Degelius sn. (DEGEL, US). SOUTHERN RHODESIA: Zimbabwe, 
Héeg s.n. (TRH). UNION OF SOUTH AFRICA: Transvaat: Mambelane, 
Mogg 2110 (PRE). 


84. Parmelia aldabrensis Dodge, Ann. Mo. Bot. Gard. 46:160. 1959. 


Type collection: Aldabra Islands, Fox 220 (K, holotype). 

Thallus adnate to bark, 4-8 cm. in diameter, buff mineral gray 
in herbarium; lobes rotund to irregular, 5-10 mm. wide, margins 
smooth to crenate and short dentate-laciniate, ciliate, cilia 0.5-2.5 
mm. long; upper surface smooth, shiny, more or less coarsely 
white-maculate; lower side brown to dark brown, rarely blackening, 
coarsely rhizinate, pale brown and naked along the margins. 
Apothecia 2-10 mm. in diameter, short-stalked, amphithecium 
maculate, disc perforate; hymenium 50-70 uw high; spores 5-7 13-18 
u, episporium 1 » thick; pycnidia present, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K+ yellow turning to 
red, C—, KC—, P+ orange yellow, atranorine and norstictic acid 
(sometimes mixed with stictic acid) present. 

This is a peculiar species of somewhat doubtful status, although 
the three collections seen are rather uniform. It is very near P. 
perforata (Jacq.) Ach. in several respects: The strong maculation, 
perforate apothecia, small spores, and the presence of norstictic 
acid. It differs from P. perforata in size, being much smaller and 
more adnate, and in having a uniformly brown or darkening lower 
side, in contrast to the jet black center and white rim so characteristic 
of P. perforata. I believe these apparently consistent differences 
justify the recognition of P. aldabrensis. 


Additional specimens examined: 


TANGANYIKA: Wadiboma, Fischer 703 (BM, G). MADAGASCAR: Fort 
Dauphin, Vallée du Moyen Mandrare, Decary, Aug. 1, 1926 (LD). 


322 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


85. Parmeli argentina Kremplh. Flora 61:476. 1878. Puate 15 

Parmelia urceolata var. cladonioides Mill. Arg. Flora 63:266. 1880. Type 
collection: Petropolis, Brazil, Deventer 35 (G, holotype). 

P. melanothriz (Mont.) Vain. var. argentina (Kremplh.) Mill. Arg. Hedwigia 
30:228. 1891. 

P. macrocarpoides Vain. f. phyllophora Hue, Nouv. Arch. Mus. Paris, ser. 4, 
1:177. 1899. Type collection: Brazil, S. M. Theresa Christina, 1889 (P, 
holotype). 

P. laongii Lynge, Ark. Bot. 13, no. 13:68, pl. 1, fig. 3. 1914. Type collection: 
Santa Anna da Chapada, Mato Grosso, Brazil, Malme 2392* (S, holotype). 

P. melanothrixz f. microspora Lynge, Ark. Bot. 13, no. 13:56. 1914. Type 
collection: Villa Morra, Asuncién, Paraguay, Malme 1585C (S, lectotype). 

P. subproboscidea Lynge, Ark. Bot. 13, no. 13:36. 1914. Type collection: 
Asuncié6n, Paraguay, Malme 1678 (S, holotype). 

P. subrugata var. mexicana Vain. Dansk Bot. Ark. 4, no. 11:5. 1926. Type 
collection: Consoquitla, Mexico, Liebmann 7562e (TUR, holotype). 


Type collection: Argentina, Lorentz & Hieronymus (M, holotype; 
BM, H, 8, W, isotypes). 

Thallus to 10 cm. in diameter, loosely attached to suberect on bark, 
mineral gray, turning buff in the herbarium; lobes rotund, broad, 7-12 
mm. wide, margins entire to crenate and dissected, ciliate, cilia 1.5-3.0 
mm. long, K—, or K+ violet; upper surface plane, shiny, more or less 
distinctly white-maculate, soredia and isidia lacking; lower side black 
and sparsely rhizinate at the center, sharply delimited from a buff to 
ivory, naked, rugose zone along the margins. Apothecia common, 
stalked, 5-10 mm. in diameter, exciple short-dentate and ciliate, 
amphithecium maculate, disc imperforate; hymenium 50-70 yp high; 
spores 6-12 11-22 y, episporium 1.5-2.0 » thick; pycnidia common, 
conidia 5-7 yp long. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ orange 
red, P—, atranorine and alectoronic acid present. 

Parmelia argentina is characterized by distinct maculae, imperforate, 
often ciliate apothecia, and a distinct pale or white zone below at the 
margin. The cilia may react K+ violet, as in the type specimens of 
P. laongiit and P. subproboscidea, but this reaction is variable and 
apparently of no taxonomic significance. My preliminary key to the 
group (Hale, 1960) illustrates the futility of separating species on cilia 
reaction or minor variation in spore size. Parmelia argentina could be 
confused with P. rigida Lynge and P. chiapensis Hale, both of which 
have perforate apothecia, with P. subrugata Kremplh., which has 
large spores, or even with P. melanothrix (Mont.) Vain., which has 
protolichesteric acid and large spores. 

Additional specimens examined: 

MEXICO: Cutapas: El Zapotal, near Tuxtla Gutiérrez, Hale 19991, 20005 
(US); El Sumidero, near Tuxtla Gutiérrez, Hale 20061 (US). HONDURAS: 


Comayaaua: Vicinity of Comayagua, Standley & Chacén 5925 (F, US); Ex 
Parafso: Vicinity of Galeras, Standley et al. 1969 (F, US). 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 323 


VENEZUELA: Lara: Barquisimeto, Saer 44 (US). BRAZIL: Minas Gerais: 
Sitio, Vainio, Lich. Bras. Exs. 973 (TUR); Mato Grosso: Corumb4, Malme s.n. 
(S$); no locality, Theresa Christina, 1899 (P). PARAGUAY: Asuncién, 
Malme, Aug. 27, 1893 (UPS); Pileomayo, Gran Chaco, Malme, Sept. 3 and 7, 
1893 (8S). ARGENTINA: Sara: Rio Pescado, Ordn, Digilio-Grassi 382 (MO). 


86. Parmelia chiapensis Hale, sp. nov. Puate 15 


Thallus laxe adnatus, 8-15 cm. latus, albido-glaucescens, lobis 
rotundatis, 7-12 mm. latis, apicem versus margine integris, centrum 
versus denticulato-laciniatis, ciliatis, ciliis 1-2 mm. longis, superne 
nitidus, levissime albomaculatus, aetate irregulariter rimosus, strato 
corticeo superiore 16-20 u crasso, strato gonidiali 22-28 yu crasso, 
medulla superiore alba, medulla inferiore pro parte crocea, toto 
110-130 uw crassa, strato corticeo inferiore 12-14 yw crasso, subtus 
niger sparse rhizinosus, ambitu castaneus, albovariegatus vel albicans, 
late nudus. Apothecia numerosa, pedicellata, ad 12 mm. diametro, 
amphithecio revolutente, rugoso, valde albomaculato, disco late per- 
forato; hymenium 70-80 yu altum; sporae 9-12>18-22 yu, episporio 
1.5-2.0 w crasso; pycnidia numerosa, conidiis non visis. Thallus K+ 
flavescens; medulla alba K—, C—, KC+ rubescens, P—, medulla 
crocea K+ atropurpurea, atranorinum, acidum alectoronicum et 
rhodophyscin continens. 

Type in the National Science Museum, Tokyo, collected at Monte 
Ovando, Chiapas, Mexico, Jan. 1, 1925, by Eizi Matuda (no. 105; 
isotype in US). 

Parmelia chiapensis is superficially very close to P. argentina 
Kremplh., especially in lobation and habit, but has perforate apothecia 
and lacks a distinct white zone below. Parmelia subrugata Kremplh. 
has a larger thallus, imperforate apothecia, and large spores. This 
species is known from the Chiapas highlands in Mexico and adjacent 
Guatemala. 

Additional specimens examined: 

MEXICO: Curapas: Mt. Ovando, Matuda, December 1937 (MICH) and 45 
(TNS). GUATEMALA: QuEzaLTENANGO: South of San Martin, Standley 
83659 (MO). 

87. Parmelia cooperi Stein. & Zahlbr. Bot. Jahrb. Engler 60:528. 1926. 
PLaTE 13 
Parmelia segreganda des Abbayes, Mem. Inst. Sci. Madagascar, ser. B, 
10:112. 1961. Type collection: Ambohitantely forest, near Ankazobe, 
Madagascar, H. des Abbayes, Lich. Mad. Bor. Sel. Exs. 17 (REN, lectotype; 
H, M, UPS, US, isotypes). 


Type collection: Kapprana, Kapland, Africa, Cooper 1813 (W, 
holotype). 

Thallus large, to 15 cm. in diameter, loosely attached, light mineral 
gray; lobes rotund, broad, 10-15 mm. wide, margins sorediate, 
soralia linear, rarely becoming submarginal, sorediate lobes sinuous, 


324 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


margins ciliate, cilia 1.5-3 mm. long; upper surface plane, more or 
less distinctly white-maculate, reticulately cracked with age; lower 
side black, sparsely rhizinate, pale brown and naked in a broad zone 
at the margins. Apothecia poorly developed, adnate, 1-2 mm. in 
diameter, disc imperforate; hymenium 90-95 yu high; spores 
7-10 18-20 pu, episporium 1-1.5 » thick; pycnidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C+ blood red, 
KC+ blood red; P—, atranorine and lecanoric acid present. 

Parmelia cooperi is a distinctive east African lichen (fig. 25) with 
an unusual combination of characters, lecanoric acid and cilia. Ex- 
cept for the presence of cilia and imperforate apothecia, it is very 
close to P. austrosinensis and occupies similar habitats in the dry 
forests of Africa. Dr. des Abbayes kindly showed me an isotype of 
his P. segreganda, which at the time seemed unique to me. Later 
examination of the type of P. cooperi, however, led me to regard them 
as the same species, although the cilia of P. segreganda are exceptionally 
long. 

Additional specimens examined: 

CONGO: Ibanda, Kabare, Spirlet, s.n. (BR); Kalongi, Albert National Park, 
de Witte 10307 (BR, US). KENYA: Rirr Vatiey Prov.: Eastern Mau Forest 
Reserve, Nakuru distr., Maas Geesteranus 11313 (L); Nyanza: Londiani, Kisumu- 
Londiani, Maas Geesternaus 10932 (L, LD). TANGANYIKA: Livingstone 
Mountains, Milo, Rae, 1935 (BM). UGANDA: Kizezi, Naiziru, Dale L48 (K). 
SOUTHERN RHODESIA: Zimbabwe, Héeg, Feb. 2, 1930 (TRH). UNION 
OF SOUTH AFRICA: Transvaau: Houtboschberg, Nelson 1056 (TRH); 
Care Prov.: 6 mi. west of Heidelberg, Héeg, June 12, 1929 (TRH). MADA- 


GASCAR: La Mondraka, Benoist, 1879 (LD); between Tananarive and Ma- 
junga, km. 133, des Abbayes, Aug. 20, 1956 (REN, US). 


88. Parmelia coralliformis Hale, sp. nov. PuLaTE 14 


Thallus laxe adnatus, expansus, rigidulus, usque ad 10 em. latus, 
viridi-glaucescens, lobis rotundatis, parum ramosis, margine integris 
vel sinuato-incisis, centrum versus isidiatis, isidiis dense coralliformi- 
bus ramosis (ut in P. fasciculata Vain.), usque ad 5 mm. altis, apicibus 
nonnullis sparse sorediatis fatiscentibus, margine ciliatis, ciliis nu- 
merosis, 1.0-1.5 mm. longis, superne laevigatus, nitidus, valde 
albomaculatus, cortice continuo vel demum fisso, strato corticeo 
superiore 17-19 y» crasso, strato gonidiali 20-22 » crasso, medulla 
alba, 90-95 crassa, strato corticeo inferiore 17-19 yu crasso, inferne 
niger, rhizinosus, ambitu castaneus, anguste nudus. Apothecia 
ignota. Thallus K+ flavida; medulla K+ rubra, C—, KC—, P+ 
aurantiaca, atranorinum et acidum salacinicum continens. 

Type in the U.S. National Herbarium, collected in mature pine 
forests, elev. 2340 m., 18 km. southeast of San Cristébal, Chiapas, 
Mexico, Mar. 24, 1960, by M. E. Hale (mo. 20287; isotypes in 
REN, 9). 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 325 


Parmelia coralliformis has the large fatiscent coralloid isidioid 
growths typical of P. fasciculata Vain. and P. ramuscula Hale. It 
differs significantly from these two species in having a shiny, dis- 
tinctly maculate upper surface. The rhizines are often produced to 
the margin below in a manner reminiscent of P. subcaperata Kremplh. 
but the center is jet black. It is known only from the type locality 
in Mexico, and may be related to P. leucosemotheta Hue, a common 
Mexican species with similar maculae and chemistry but differing in 
having soredia. 


89. Parmelia euneta Stirton, Scot. Nat. 4:298. 1877-78. 


Type collection: Victoria, Africa, Thomson (BM, holotype; GLAM, 
isotype). 

Thallus loosely adnate, corticolous, often expanded, 10-25 cm. in 
diameter, coriaceous, mineral gray; lobes rotund, margins entire to 
crenate, ciliate, cilia 2-4 mm. long; upper surface plane, shiny, faintly 
to distinctly white-maculate, reticulately cracked with age, isidia and 
soredia lacking; lower side black, sparsely rhizinate, brown to white 
or mottled in a broad zone at the margins. Apothecia stalked, 5-15 
mm. in diameter, amphithecium strongly maculate, exciple usually 
ciliate, disc perforate; hymenium 60-80 u high; spores 7-12 18-26 y, 
episporium 1.5-2.5 » thick; pycnidia numerous, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C+ rose, KC+ 
blood red, P—, atranorine and gyrophoric acid present. 

Parmelia euneta appears to be a pantropical species, although it is 
very rare outside of Africa. It is unique in the presence of maculae, 
gyrophoric acid, and stalked perforate apothecia. The thallus is 
usually quite large and rather coriaceous. The maculae vary con- 
siderably in distinctness but will always be distinct near the base of 
apothecia. The spores are uniformly intermediate in size. It has 
no close relatives with gyrophoric acid, but it does resemble externally 
P. abessinica Kremplh., a smaller more membranous species with 
a similar variation in maculae and with intermediate-sized spores. 
Additional specimens examined: 

HAITI: Kenscoff, Thomas 101 (MO, NY, US). 

SIERRA LEONE: Falaba, Small 450 (K). ANGOLA: Mayumbe, Goss- 
weiler 7181 (BM); Vila Salazar, Cuanza-Norte, Degelius, Mar. 6, 1960 (DEGEL, 
US). CONGO: Isalowe Reserve, Yangambi, Louis 8675 (BR, US). TANGAN- 


YIKA: Milo, Livingstone Mtns., Rae, 1935 (BM). 
CEYLON: No locality, Tiwary, s.d. (BM). 


90. Parmelia hababiana Gyel. Repert. Sp. Nov. Fedde 29:288. 1931. PuarE 15 
Parmelia abessinica var. sorediosa Mill. Arg. Flora 68:501. 1885. Nomen- 
clatural basis of P. hababiana. 
P, nilgherrensis f. subciliaris Vainio, Hedwigia 37:40. 1898. Type collection: 
Ruwenzori, Uganda, Africa, Scott Elliott 218 (TUR, holotype). 
P. subciliaris (Vain.) Dodge, Ann. Mo. Bot. Gard. 40:377. 1953. 


326 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


P. alleniti Dodge, Ann. Mo. Bot. Gard. 46:150. 1959. Type collection: 
Lake Ngunga, Kenya, Allen 1783 (FH-Howe, holotype) (not 1837). 

(2) P. hendrickzii Dodge, Ann. Mo. Bot. Gard. 46:124. 1959. Type collec- 
tion: Kahusi, Congo, Hendrickr 4307 p.p. (EA, holotype). 

Type collection: Near Habab, Ethiopia, Hildebrandt 310 p.p. (G, 
lectotype). 

Thallus loosely adnate to bark, 6-10 cm. in diameter, mineral gray; 
lobes rotund, 7-12 mm. wide, margins often subascending, crenate, 
sorediate, soralia linear or in part capitate and submarginal, sorediate 
lobes becoming revolute with age, ciliate, cilia rather sparse, 0.5-2 
mm. long; upper surface dull to quite shiny, faintly to strongly white- 
maculate, especially near the base of apothecia, rarely becoming white- 
pruinose; lower side black and sparsely rhizinate at the center, brown 
to ivory or mottled and naked in a broad zone at the margins. Apo- 
thecia rare, 2-3 mm. in diameter, amphithecium sorediate, disc per- 
forate; hymenium 60-70 » high; spores 8-1015-81 yu, episporium 
1.5 » thick; pycnidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ red, P—, 
atranorine, cryptochlorophaeic acid, and protolichesteric acid present; 
or medulla K—, C—, KC—, P—, atranorine and protolichesteric acid 
present. 

The name P. hababiana was probably proposed by Gyelnik as a 
nomenclatorial exercise, since he did not see the the type of the variety 
forming the basis of the species or, as far as we know, identify any 
material belonging to this species. It now appears that P. hababiana 
is much more common than had been thought. It is common not 
only in the savannas of Africa but also in the dry upland oak forests 
of Central America and in Asia. The outstanding morphological 
characters are the marginal soralia, cilia, faint to distinct maculae, and 
white or mottled zone below near the margin. Superficially it appears 
very similar to the nonciliate species P. praesorediosa Nyl. or P. 
austrosinensis Zablbr. More than half of the specimens, including all 
the type specimens listed above, contain protolichesteric and crypto- 
chlorophaeic acids (KC+ red) and the remainder contain only proto- 
lichesteric acid (KC—). These two chemical variants have nearly 
identical distributions (figs. 26, 27), although the typical strain with 
cryptochlorophaeic acid is much more common throughout the geo- 
graphic range. Parmelia abessinica Kremplh., which has similar 
chemical variation, is the nonsorediate counterpart of P. hababiana. 

Additional specimens examined: 

Cryptochlorophaeic and protolichesteric acids present: 

U.S.: Arizona: Stewart Creek Campground, Chiricahua Mountains, Cochise 
Co., Weber 828021 (COLO, US). MEXICO: Oaxaca: 53 km. northwest of 


Oaxaca, Hale 20796 (US); 132 km. northwest of Oaxaca, Hale 20835 (US); 
Curapas: Just south of Teopisca, Hale 20499 (S, US); 2 km. north of highway 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 327 


190 on road to Puebla Nueva, Hale 20164 (US). GUATEMALA: Jourtapa: 
Vicinity of Jutiapa, Standley 75949 (F, MO). HONDURAS: Comayagua: 
Siguatepeque, Yuncker 6505 (MO). NICARAGUA: Jinorega: Cerro Sialci, 
Standley 10591 (F). COSTA RICA: La Palma, Valerio 13 (S). 
ARGENTINA: Santiago del Estero, Dept. Bando, Vega 20960 (SI). 
ANGOLA: Bré: Chinguar, Degelius s.n. (DEGEL). KENYA: Rurt VALLEY 
Prov.: Lake Nakuru, Maas Geesteranus 4631 (L), Eastern Mau Forest Reserve, 
Nakuru Distr., Maas Geesteranus 11378 (L); Cherangani, Maas Geesteranus 
4708 (L); Cenrrau Prov.: Nairobi, Maas Geesteranus 4438 (L); Nyanza Prov.: 
Tinderet Forest Camp, Kisumu-Londiani Distr., Maas Geesteranus 4955 (L, 
LD, US). SOUTHERN RHODESIA: Zimbabwe, Héeg s.n. (TRH). UNION 
OF SOUTH AFRICA: Transvaat: Louis Trichardt, Hanglip Forest, Alm- 
born 12011 (LD); Narau: Emmersdale, Estcourt, Héeg s.n. (TRH); Creighton 
Station, Héeg sn. (TRH). MADAGASCAR: Tananarive, Bathie s.n. (LD). 
INDIA: Almora Distr.: Above village Naret, Askote, Awasthi 3297 p.p. 
(AWAS); Ranikhet, Awasthi 3548 (AWAS). CHINA: Schilungba, Yunnan, 
Handel-Mazzetti 255 (MICH). 
Protolichesteric acid only present: 
MEXICO: Oaxaca: Cerro Zempoaltepetl, Hallberg 690 (MICH). 
LIBERIA: Western Prov.: Wohmen, Vonjama, Baldwin 12452 (US). 
ANGOLA: Cuanza-Norre: Salazar, Degelius s.n. (DEGEL); BENnevueEta: 
Between Benguela and Coporolo, Degelius sn. (DEGEL). KENYA: Rurt 
VaLLEY Proy.: Menengai, Nakuru Distr., Maas Geesteranus 10280 (L); Nyanza 
Prov.: Tinderet Forest Reserve, Kisumu-Londiani Distr., Maas Geesteranus 
4949 (LD); Centra Prov.: 4 mi. west of Nairobi, Maas Geesteranus 4394c 
(LD). SOUTHERN RHODESIA: Near Bulawayo, Héeg, s.n. (TRH). UNION 
OF SOUTH AFRICA: Transvaat: 2 mi. north of Warmbad, Waterberg, Almborn 
5889 p.p., 5896 (LD); Natau: Boschfontein Forest, Lions River Distr., Almborn 
8721, 8728 (LD); Natal Table Mountain, Almborn 8555 (LD); Weenen, Héeg 
s.n. (TRH); Emmersdale, Estcourt, Héeg s.n. (TRH); Care Prov.: Near Port 
Elizabeth, Héeg s.n. (TRH). 
91. Parmelia hanningtoniana Mill. Arg. Flora 73:339. 1890. PLATE 2 
Parmelia sinensis Hue f. isidiata Ceng. Samb. Nuov. Giorn. Bot. Ital. 
40:283. 1933. Type collection: Limbolo, Angola, Fenaroli II (FI, holotype) . 


Type collection: South Africa, Hannington (G, holotype; K, 
isotype). 

Thallus adnate to loosely attached, 5-8 cm. in diameter coriaceous, 
greenish mineral gray; lobes rotund, 15-20 mm. wide, margins crenate, 
ciliate, cilia 1.5-2.0 mm. long; upper surface distinctly white- 
maculate, plane to rugulose, isidia and soredia lacking; lower side 
black and sparcely rhizinate in the center, pale tan, rugose, and 
naked in a broad zone at the margins. Apothecia large, 4-10 mm. 
broad, stalked, amphithecium extremely wrinkled and lobulate, 
white-maculate, disc perforate; hymenium 50-60 y high; spores 
7-11X13-18 yw, episporium 1.5 y» thick, pycnidia present, conidia 
not seen. 

Reactions: Thallus K+ yellow; medulla K—, C+ rose, KC+ 
red, P—, atranorine and gyrophoric acid present. 

729-018—65——11 


328 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Parmelia hanningtoniana is apparently restricted to the savannas 
of west Africa. The unusual apothecia (cf. pl. 2) set it apart from 
all other Parmelias. They are large, stalked, and the amphithecium 
is coarsely sublobulate. In other respects it is close to P. euneta 
Stirt. 

Additional specimens examined: 


ANGOLA: Moxico: Between Luso and Cochipoque, Degelius, Feb. 16, 1960 
(DEGEL, US); Koiembe, Quingenge-Benguela, Damann, Tav. Lich. Lusit. Sel. 
Exs. 168 (H, LD, US). CONGO: Biano, 26°3' E., 10°16’ S., Schmitz 3884 (BR). 
92. Parmelia hypotropa Nyl. Syn. Lich. 379. 1860. 

Parmelia hypotropa var. sorediata Mill. Arg. Flora 60:77. 1877. Type col- 
lection: Near Dallas, Texas, Boll (G, holotype; US, isotype). 

P. cetrata var. hypotropoides Nyl. ex Willey, Bot. Gaz. 21:203. 1896. Type 
collection: New Bedford, Massachusetts, Willey (US, holotype). 

P. lividotessellata f. ablephara Hue, Nouv. Arch. Paris, ser. 4, 1:191. 1899. 
Type collection: Near Tonza, Algeria, Africa, Maisonneuve, November 
1840 (P, holotype). 

P. hypoleucina Stein. Oecster. Bot. Zeitschr. 67:282. 1918. Type collection: 
Barro, Portugal, Menyhart 113 (WU, holotype). 

P. trichotera var. subincana Mah. & Gil. Bull. Soe. Bot. Fr. 68:286. 1921 
[1925]. Type collection: Forét de Mamora, near Kenitra, Morocco, 
Gillet 105 bis (P, holotype). 

P. subincana (Mah. & Gil.) Mah. & Gil. Bull. Soc. Bot. Fr. 72:862. 1925. 

P. corrugis var. sorediata (Mill. Arg.) Zahlbr. Cat. Lich. Univ. 6:236. 1929. 

Parmotrema gattefossei Choisy, Bull. Soc. Bot. Fr. 78:458. 1931. Type col- 
lection: Forét Mamora, Morocco, Gattefossé (LY, herb. no. 1698, holo- 
type). 

P. werneri Choisy, Bull. Soc. Bot. Fr. 78:457. 1931. Type collection: Oued 
Cherrat, Morocco, Gattefossé (LY, herb. no. 1753, lectotype). 

Parmelia mueller-argoviensis Gyel. Repert. Sp. Nov. Fedde 29:288. 1931. 
Based on P. hypotropa var. sorediata Mill. Arg. 

P. gattefossei (Choisy) Zahlbr. Cat. Lich. Univ. 8:559. 1932. 

P. werneri (Choisy) Zahlbr. Cat. Lich. Univ. 8:569. 1932. 

P. hypoleucina var. subincana (Mah. & Gil.) Werner, Bull. Soc. Sci. Nat. 
Maroc. 35:57. 1955. 


Type collection: Comancharies, Texas [Mexico?], Berlandier & 
Trécul 2093 (P, holotype; H, isotype). 

Thallus loosely adnate, 8-12 cm. in diameter, light mineral gray, 
sometimes turning pink below in the herbarium; lobes broad, ascend- 
ing, 10-15 mm. wide, margins broadly crenate, ciliate, cilia 1-3 mm. 
long, sorediate, soralia linear, sorediate lobes sinuate; upper surface 
plane, shiny, strongly white-maculate, becoming rugose with age; 
‘ower side black and sparsely rhizinate in the center, outer margin 
white or rarely tan, rugulose, naked. Apothecia extremely rare, to 
4 mm. in diameter, disc perforate; hymenium 40-50 u high; spores 
5-68-10 pu, episporium 1 » thick; pycnidia not seen. 

Reactions: Thallus K+ yellow; medulla K+ yellow or yellow 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 329 


turning red, C—, KC—, P+ pale orange red, atranorine, norstictic 
and/or stictic acid present. 

In his original description Nylander cites Parmelia hypotropa rather 
ambiguously as “*P. hypotropa” under P. perforata. It is possible to 
interpret P. hypotropa as having subspecific rank, but Nylander’s 
brief notes also imply that it could be considered as a variety of P. 
perforata. I prefer to consider it as validly published at the species 
rank. In any event the first use of Parmelia hypotropa at a species 
rank (apparently Nylander in Flora 52: 291. 1869) still antedates 
any of the synonyms, 

Parmelia hypotropa is easily recognized by the distinct maculae, 
marginal cilia and soredia, and the broad white marginal zone below. 
Apothecia are known only from a single collection. It is especially 
common in the southeastern United States (fig. 8) on the lower trunks 
of deciduous trees, and occurs also in southern California, in Mexico, 
in the Mediterranean region, and in one locality in Formosa. It is the 
sorediate counterpart of P. perforata (Jacq.) Ach., which has a similar 
but more southerly range in eastern United States (fig. 7) and is col- 
lected more frequently on the upper branches of deciduous trees. 

All specimens from eastern North America and Mexico contain 
norstictic acid in abundance. Those from California and Europe 
(including the type specimens from Europe and Africa listed above) 
usually contain stictic acid, and norstictic acid cannot always be 
demonstrated with a G.A.o-T test, although a K,CO, test is often 
successful. It is probable that the two acids always occur together 
but that our present chemical tests are not yet refined enough to 
detect them in mixtures. 

Additional representative specimens examined: 

U.S.: Massacuusetrs: New Bedford, Bristol Co., Willey s.n. (US); Connec- 
TIcuT: 1 mi. south of Colebrook, Litchfield Co., Hale 15412 (US); New York: 
Near Sound, Suffolk Co., Imshaug 25770A (H); New Jersey: Greenbank State 
Forest, Burlington Co., Hale 15331 (US); PENNsyLvanra: 4 mi. northwest of 
Dingmans Ferry, Pike Co., Hale 16249 (US); DeLtaware: Laurel, Sussex Co., 
Commons s.n. (PH); Marynanp: 2 mi. northwest of Bristol, Anne Arundel Co., 
Hale 14333 (US); West Virernta: Near Pullman, Ritchie Co., Hale 10791 (US); 
3 mi. northeast of Alvon, Greenbrier Co., Hale 18957; Glen Rogers, Wyoming Co., 
Hale 11643 (US); Onto: Miami Whitewater Forest, Hamilton Co., Taylor 31 (US); 
Newstonville, Clermont Co., Hale 13225 (US); Kentucky: 4 mi. northeast of 
Whitly City, McCreary Co., Hale 13748 (US) ; 2 mi. east of Paducah, McCracken 
Co., Hale 13692 (US); Inprana: Harrison County State Forest, Harrison Co., 
Hale 14038 (US); Intrnors: Grantsburg, Johnson Co., Hale 14013 (US); Wiscon- 
sin: Near Dunnville, Dunn Co., Culberson 3725 (WIS); Wisconsin Dells, Juneau 
Co., Thomson 1825 (WIS); Missourr: Near Houston, Texas Co., Hale 4222 (U8) ; 
Vireinia: Vint Hill Farms, Fauquier Co., Cooke (LD); Lee’s Millpond, Isle of 
Wight Co., Culberson 7027 (DUKE); Prince William State Forest, Prince William 
Co., Hale 15223 (US); Millsboro Springs, Bath Co., Hale 12933 (US); Norra 


330 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


CAROLINA: 3 mi. southeast of Bertha, Currituck Co., Culberson 6968 (DUKE, US); 
3 mi. north of Halifax, Halifax Co., Hale 16332 (US); 10 mi. south of Asheville, 
Buncombe Co., Culberson 7125 (DUKE); TenNessEE: Sycamore, Cheatham Co., 
Phillips s.n. (US); Nashville, Davidson Co., Gattinger s.n. (fertile) (M); GrorGiA: 
Near Blue Ridge, Fannin Co., Hale 7321 (US); ALasama: De Soto State Park, 
De Kalb Co., Hale 7051 (US); Fioripa: 3 mi. south of Sanford, Seminole Co., 
Hale 17679 (US); Eustis, Lake Co., Nash 1879 (US); Mississippi: 4 mi. south of 
Nettleton, Monroe Co., Hubricht B1533 (US); Holly Springs, Marshall Co., 
Hale 7850 (US); Lourstana: Plaquemines Parish, Langlois s.n. (PH, US); 
Arxansas: Near Briggsville, Yell Co., Hale 3747 (US); 0.5 mi. north of Francis, 
Boone Co., Dodge et al. s.n. (US); near Jasper, Newton Co., Hale 3081 (US); 
OxiaHoma: Near Ardmore, Carter Co., Hale 4859 (US); Texas: Meridian, 
Bosque Co., Hale 5201 (US); Wisdom Ranch, Wilson Co., Parks 1098 (WIS); 
20 mi. northwest of Fort Davis, Jeff Davis Co., Parks 1042 (WIS); Duffau, Erath 
Co., Hale 5496 (US); Catrrornia: San Diego, San Diego Co., Palmer 279 (US), 
Alderson 779 (US); Santa Catalina Island, McGregor 26 (US). MEXICO: 
TAMAULIPAS: South of Antigua Morelos, Reeder 1889 (US). 

PORTUGAL: Near Magoito, Estremadura, Tavares, Lich, Lusit. Sel. Exs. 98 
(H, US, WIS); near Azoia, Estremadura, Tavares, Vezda Lich. Sel. Exs. 68 (H, LD, 
US). ITALY: Pisa, Etruria, Vivante, Gyel. Lichenoth. 139 (H, LD); between 
Spezia and Pisa, Hasselrot s.n. (F). 

MOROCCO: Near Rabat, Gattefossé s.n. (P); Oued Bouznika, Chaouia, 
Gattefossé s.n. (P). 

FORMOSA: Keitau, Asahina 3411 (TNS, US). 


93. Parmelia leucosemotheta Hue, Nouv. Arch. Mus. Paris, ser. 4, 1:192. 1899. 
PLaTE 16 

Type collection: Abrededores, San Luis Potosi, Mexico, Maury 
7650 (P, holotype). 

Thallus large, 10-20 em. broad, loosely attached to bark, mineral 
eray; lobes rotund, 10-15 mm. wide, margins entire to dentate- 
laciniate with age, ciliate, cilia 1.5-2.5 mm. long, sorediate, soralia 
mostly on lateral margins, linear; upper surface shiny, strongly 
white-maculate, usually becoming conspicuously reticulately cracked 
with age; lower side black and sparsely rhizinate, dark brown to 
brown or mottled white in a broad zone along the margins. Apoth- 
ecia rare, more or less adnate, to 6 mm. in diameter, amphithecium 
sorediate, disc imperforate or rarely perforate; hymenium 65-75 4 
high; spores 7-10 12-16 y, episporium 1.0-1.5 » thick; pycnidia not 
seen. 

Reactions: Thallus K+ yellow; medulla K+ yellow turning red, 
C—, KC—, P+ pale orange red, atranorine and salacinic acid present. 

Parmelia leucosemotheta is characterized by strong maculation, 
marginal soredia, and a black lower side. It is very similar in many 
respects to P. reparata Stirt., which differs in being nonsorediate and 
in having perforate apothecia. It also closely resembles P. subsumpta 
Nyl., which differs principally in having a pale lower side with short 
dense rhizines and a narrow rhizinate or papillate zone at the margins. 
If specimens of P. subswmpta have an abnormally blackened lower 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 331 


side, one must rely on the extent of papillae in the marginal zone to 
separate them from P. leucosemotheta, which typically has a broad, 
dark and shiny bare zone. These two species have similar distribution 
patterns, although P. lewcosemotheta is much more common in Mexico. 

Additional specimens examined: 

MEXICO: Nurvo Lron: Monterrey, Lacas 521 (F); Hipatco: La Placita, 
Moore 3900 (US); Veracruz: Northeast of Huatusco, Hale 19551 (COLO, 
DUKE, MSC, REN, S, TNS, US); 7 km. north of Fortin de las Flores, Hale 
19709 (S, US), Hale 1690 (DUKE, US); 9 km. east of Jalapa, Hale 21233 (US); 
PurBua: 3 km. west of Puebla-Veracruz state line on highway 150, Hale 19614 
(US); Micnoackn: Morelia, Arséne 8238 (US); Oaxaca: Cerro del Machete, 
Pochutla Distr., Reko 6307a (F); Curapas: 40 km. southeast of Comitan, Hale 
20484 (US), 20483 (COLO, US); 14 km. west of San Cristébal, Hale 20573 (LISU, 
US). GUATEMALA: Jatapa: Jalapa, Standley 76552 (MO). COSTA RICA: 
San Jos&: Santa Maria de Dota, Standley 41727 (US); Carraco: North of 
Cartago, Standley 49616, 49635 (US). 

HAITI: Summit of Montagne Noire, Wetmore 2730 (MSC). 

BRAZIL: Minas Gerais: Caldas, Mosén 2313 (UPS), Henschen s.n. (UPS); 
Sao Pavto: Piquete, Robert sn. (BM). ARGENTINA: Without locality, 
Loreniz & Hieronymus (M). 

UNION OF SOUTH AFRICA: Boschfontein Forest, Lions River Distr., 
Almborn 8674 (LD). 


94. Parmelia lobulascens Steiner, Verh. Zool. Bot. Gesell. Wien 53:234. 1903. 
Type collection: Cameroons, Bornmiiller (WU, holotype). 


Thallus loosely attached, coriaceous, 5-10 cm. in diameter, mineral 
gray; lobes rotund, 8-14 mm. wide, margins entire, lobulate toward 
the center, central lobes often crowded, margins sorediate, soralia 
irregular to sublinear, ciliate, cilia rather sparse, 0.5-2.0 mm. long; 
upper surface shiny, moderately to strongly white-maculate, rugulose 
and cracked with age; lower side black and coarsely rhizinate, margins 
dark brown, naked, rugose. Apothecia and pycnidia unknown. 

Reactions: Thallus K+ yellow; medulla K—, C+ rose, KO+ 
blood red, P—, atranorine and gyrophoric acid present. 

The type of P. lobulascens is a very poor specimen. Laciniae or 
lobules are quite abundant and have irregularly sorediate margins, 
a fact overlooked by Steiner. Fortunately additional collections 
from Africa have clarified the range of variation in the species. The 
lobules may not be conspicuously developed and the soredia may be in 
elongate, more normal soralia. Maculae are always very distinct. 
It is extremely close to P. pseudonilgherrensis Asah., which contains 
alectoronic and a@-collatolic acids and lacks lobules. 

Additional specimens examined: 

ETHIOPIA: North of Debra Marcos, Chokke Mtns., Hiller L85 (K); no 


locality, Schimper, s.d. (BM). KENYA: Nyanza: Tinderet Forest Reserve, 
Kisumu-Londiani, Maas Geesteranus 5532 (L). 


332 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


95. Parmelia melanothrix (Mont.) Vain. Acta Soc. Faun. Fl. Fenn. 7, no. 


7:30. 1890. 
Parmelia urceolata var. melanothriz Mont. Ann. Sci. Nat. Bot., ser. 2, 2:372. 
1834. 


Type collection: Brazil, Gaudichaud 89 bis (P, holotype). 

Thallus 5-15 cm. in diameter, loosely attached to suberect on bark, 
mineral gray; lobes rotund, 6-12 mm. wide, margins crenate, short 
dentate-laciniate, especially toward the center, densely ciliate, cilia 
2-3 mm. long; upper surface plane, shiny, white-maculate, isidia and 
soredia lacking; lower side black and rhizinate in the center, pale 
tan to white or mottled in a broad zone at the margins. Apothecia 
common, stalked, to 10 mm. in diameter, exciple minutely dentate or 
laciniate, ciliate, amphithecium rugose, maculate, disc imperforate; 
hymenium 80-100 » high; spores 10-16 20-26 y, episporium 2-3 yu 
thick; pycnidia common, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC—, P-, 
atranorine and protolichesteric acid present. 

Parmelia melanothriz is one of the few tropical Parmelias that is 
usually correctly identified in herbaria. The salient taxonomic 
features are the large spores, K—, P— reaction, maculae, and cilia. 
The only species with which it might be confused is P. argentina 
Kremplh., which is KC+ red (alectoronic acid) and has intermediate- 
sized spores. Parmelia melanothriz has been commonly collected in 
southeastern Brazil; there is a disjunct locality in Réunion, in the 
Indian Ocean. 


Additional specimens examined: 


BRAZIL: Marannao: Sao Bernardo, Serra de Cubatao, Burchell 3942 (K); 
Minas Gerais: Caldas, Mosén 2314, Lich. Austro-Amer. Herb. Regnell. 358 
(BM, H, MSC, 8, UPS, W); Sitio, Vainio, Lich. Bras. Exs. 950 (TUR, M, UPS); 
no locality, Warming 273 (M); R1o pg Janeiro: Serra dos Orgfos, Helmreichen, 
s.d. (M), Burchell 2403 (K); Rio de Janeiro, Glaziow 1835 (M); Sao Pauto: 
Apiaf, Puiggart, s.d. (M); near Lapa, July 1901, Schiffner (M); Paran&: Fiarehy, 
September 1908, Dusén (M); Ypiranga, Liiderwaldt, Zahlbr. Lich. Rar. Exs. 
222 (BM, BPI, MICH); Rio Granpe vo Sut: Elisenau, Wirttemberg, Born- 
miller 796 (M). 

REUNION ISLAND: Salazie, 1893, Chauret (P), 1889, Rodriguez (P). 


96. Parmelia natalensis Stein. & Zahlbr. Bot. Jahrb. Engler 60:515. 1926. 


Type collection: Van Reenens Pass, Drackensberge, Black Moun- 
tains, Natal, Africa, Brunnthaler (W, holotype). 

Thallus expanded, coriaceous, 8-15 cm. in diameter, mineral gray; 
lobes rotund, 9-15 mm. wide, margins entire, ciliate, cilia 1-3 mm. long, 
sometimes sorediate; upper surface plane, shiny, strongly white- 
maculate, becoming sorediate in a broad zone along the margins, 
soralia orbicular, soon coalescing into extensive sorediate masses; 
lower side black and sparsely rhizinate, brown and naked in a broad 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 333 


zone along the margins. Apothecia rare, 3-8 mm. in diameter, very 
short-stalked, amphithecium sorediate, disc imperforate, often radially 
split; hymenium about 50 » high; spores 10-13 x 18-22 yu, the episporium 
to 2 uw thick; pycnidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ red or 
orange red, P—, atranorine and alectoronic acid present. 

Parmelia natalensis, an African endemic, has a large coriaceous 
thallus with a broad submarginal zone of soralia. The maculae are 
usually very distinct. Parmelia pseudonilgherrensis Nyl., another 
maculate species with alectoronic acid, has strictly marginal soralia. 
Parmelia rimulosa Dodge, which occurs in South Africa, has a more 
membranous fragile thallus and extensive submarginal sorediate 
pustules. 


Additional specimens examined: 


CONGO: Volcan Karisimbi, Plaine de Lukumi, Louis 5411 (BR). KENYA: 
Nyanza: Tinderet Forest Reserve, Kisumu-Londiani, Maas Geesteranus 4956 
(L, LD), 5017 (L, US), 5532 (L), 11100 (L); Londiani, Kisumu-Londiani, Maas 
Geesteranus 10933 (L, LD); Rirr Vattey: Cherangani Mountains, Trans Nzoia 
Distr., Maas Geesteranus 4715 (L), 4829 (L, LD); Eastern Mau Forest Reserve, 
Nakuru Distr., Maas Geesteranus 11313 pr. p. (L). UNION OF SOUTH 
AFRICA: Nata: Polela Forest, Polela, Almborn 9500 (LD). 

97. Parmelia nilgherrensis Nyl. Flora 52:291. 1869. 

Parmelia yunnana Hue, Nouv. Arch. Mus. Paris, ser. 4, 1:186. 1899. Type 
collection: Lopin-chan, Yunnan, China, Delavay, July 31, 1888 (P, 
holotype). 

P. siamensis Vain. Ann Soc. Zool. Bot. Fenn. 1, no. 3:37. 1921. Type collec- 
tion: Doi Sutep, Thailand, Hosseus (TUR, holotype). 


Type collection: Nilgherries Mountains, India, Perrottet (H, Nyl. 
herb. no. 35337, holotype). 

Thallus loosely attached to bark, 8-15 cm. wide, mineral gray, 
turning buff in the herbarium; lobes rotund, 10-15 mm. wide, margins 
entire to crenate or dentate toward the center, ciliate, cilia 1-3 mm. 
long, rather coarse, often furcate; upper surface plane, shiny, strongly 
white-maculate, isidia and soredia lacking; lower side black and 
coarsely rhizinate, dark brown and naked in a broad zone at the 
margins. Apothecia large, numerous, 8-20 mm. in diameter, stalked, 
amphithecium strongly rugose and maculate, disc perforate; hy- 
menium 100-130 yu high; spores 10-18%17-26 u, episporium 2.0- 
2.5 »; pycnidia abundant, conidia 14-16 yu long. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC-+ red or 
orange red, P—, atranorine, a-collatolic acid (predominantly), and 
alectoronic acid present; medulla, if pigmented, K+ purple, rhodo- 
physcin present. 

Parmelia nilgherrensis is a typical Asian species especially common 
in the foothills of the Himalayan Mountains (fig. 28). It is charac- 


334 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Ficure 28.—Distribution of Parmelia nilgherrensis Nyl. 


terized by strong maculae, perforate apothecia, and a dark lower side 
even at the margins. The predominant chemical is a-collatolic acid 
and the medulla does not fluoresce at all. The sorediate counter- 
part appears to be P. pseudonilgherrensis Asah. 

\* Additional specimens examined: 

SIKKIM: Lachen, Hooker 1994 (K). NEPAL: Lantang, Khola, Polunin 
M161 (BM); no locality, Buchanan, Feb. 3, 1803 (BM). INDIA: Wan, Gharwal 
Distr., Pranavananda A-119 (SESH, US); Ihageswar forests, en route to 
Ihagersen, Sarma A-114 (SESH, US); Ranikhet, Senthna 2 (US); Mussoorie, 
Ali 3 (BM), Awasthi 3815 (AWAS), Mehra 18 (MO); Mahasa, Simla, Thomson 
1990 (K); Madras, Gamble 18048 (K); Jogeswar Ridge, Almora Distr., Awasthi 
3505 (AWAS); Shola, near mile 9, Kodaikanal-Berijani Road, Madurai, Foreau 
4238 (AWAS). CEYLON: Naini Jal, Duvay, May 1930 (BM). CHINA: 
Yunnan: Eastern slopes of Likiang Snow Range, Likiang, Rock 11753 (US). 
THAILAND: Doi Sutep, Hosseus (TUR). 


98. Parmelia paulensis Zahlbr. Denkschr. Akad. Wiss. Naturw. Wien 83:175, 
pl. 1, fig. 6. 1909. 

Parmelia dolosa des Abbayes, Mem. Inst. Sci. Madagascar, ser. B, 10:115. 
1961. Type collection: 30 km. northeast of Ankazobe, }orét d’Ambo- 
hitantely, Centre Moyen, Madagascar, des Abbayes (REN, lectotype; 
US, isotype). 

Type collection: Near Taipas, Mount Jaragué, Sao Paulo, 
Brazil, Schiffner (W, holotype). 

Thallus loosely attached, to 10 cm. broad, membranous, mineral 
gray; lobes rotund, to 15 mm. wide, subimbricate and dissected 
toward the center, margins and submarginal areas isidiate or sorediate- 
isidiate, densely ciliate, cilia 2-3 mm. long; upper surface smooth, 
dull to more or less distinctly white-maculate, becoming reticulately 
cracked with age; lower side black and sparsely rhizinate, dark to 
pale brown, ivory, or mottled and naked in a broad zone along the 
margins. Apothecia rare, 3-5 mm. in diameter, exciple short- 
dentate, ciliate, amphithecium rugose and maculate, disc imperforate; 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 335 


hymenium 50-60 » high; spores 13-1526-28 y, episporium 3 2 
thick; pycnidia abundant, conidia to 10 y long. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC-+ faintly 
reddish or KC—, P—, atranorine, protolichesteric acid, and apparently 
an unknown substance present. 

Parmelia pawlensis has sorediate isidia very similar to those of P. 
mellissvi Dodge, a pantropical species with alectoronic acid and inter- 
mediate spores. Unfortunately few specimens of P. paulensis have 
been available for study and the chemical constituents are uncertain. 
Protolichesteric acid is usually demonstrated but one syntype of P. 
dolosa (Forét de Manjakatompo) appears to contain a fatty acid near 
caperatic acid. I was unable to prove gyrophoric acid reported by 
des Abbayes. The KC reaction is either negative or faintly reddish 
but no identifiable lichen substances have been demonstrated. Par- 
melia paulensis has an unusual Madagascar-Brazil distribution shared 
by P. melanothriz (Mont.) Vain., a related species. 

Additional specimens examined: 


MADAGASCAR: Forét de Manjakatompo, Ankaratra, Centre Moyen, des 
Abbayes, July 30, 1956 (REN). REUNION: No locality, Rodriguez, 1890 (P). 


99. Parmelia perforata (Jacq.) Ach. Meth. Lich. 217. 1803. PLATE 2 

Lichen perforatus Jacq. Coll. Bot. III, 1:116, pl. 3. 1786. 

Cetraria corrugis Fr. Syst. Orb. Veg. 1: 283. 1825. Type collection: North 
America, Schweinitz (UPS, holotype). 

Parmelia coriacea var. perforata (Jacq.) Eschw. in Mart. Fl. Bras. 1: 206. 
1833. 

P. perforata var. ciliata Nyl. Syn. Lich. 377. 1860. Based on Lichen 
perforatus Jacq. 

P. hypotropa f. parmata Nyl. in Cromb. Journ. Linn. Soc. London 17:568. 
1880. Type collection: pl. 20, fig. 44 in Dillenius, Hist. Musc. (OXF, 
holotype) (= Parmelia perforata (Jacq.) Ach.). 

P. corrugis (Fr.) Miill. Arg. Flora 70:59. 1887. Based on Cetraria corrugis 
Fr. 

P. hypotropoides Nyl. ex Willey, Bot. Gaz. 21:204. 1896. Type collection: 
New Bedford, Massachusetts, Willey (US, holotype). 

P. corrugis f. parmata (Nyl. in Cromb.) Zahlbr. Cat. Lich. Univ. 6:236. 
1929. 

P. ciliata (Nyl.) Gyel. Report. Sp. Nov. Fedde 30:225. 1932. Based on 
P. perforata var. ciliata Nyl. 

P. erecta Berry, Ann. Mo. Bot. Gard. 28:103. 1941. Type collection: 
Marble Falls, Newton Co., Arkansas, Dodge, Berry, and Johnson (MO, 
holotype). 


Type collection: New England, Tuckerman, Lich. Amer. Septentr. 
Exs. 69 (UPS, neotype; isotypes in FH, L, M, MO). 

Thallus loosely attached to suberect, corticolous, 8-12 cm. in di- 
ameter, light mineral gray or turning pinkish in the herbarium; lobes 
large, often monophyllous, 10-20 mm. wide, margins entire to lacerate 
or even laciniate, ciliate, cilia 2-4 mm. long; upper surface plane to 


336 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


rugulose, shiny, strongly white-maculate, isidia and soredia lacking; 
lower side jet black and sparsely rhizinate at the center, pure white 
to ivory or rarely tan in a distinct broad zone along the margins. 
Apothecia numerous, stalked, 10-20 mm. in diameter, exciple rarely 
ciliate, amphithecium strongly rugose, maculate, disc perforate; 
hymenium 50-65 p high; spores 7-9 X 12-16 u, episporium 1 yp; pycnidia 
numerous, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K+ yellow turning red, 
C—, KC—, P+ pale yellow orange, atranorine and norstictic acid 
present. 

A neotype is required for the typification of P. perforata since the 
holotype is probably destroyed, although represented by an excellent 
color plate. A logical choice of a neotype is the specimen in Tucker- 
man’s exsiccate which is widely distributed in larger herbaria. 

Parmelia perforata is a name which has in the past been applied to 
any large Amphigymnia species with perforate apothecia. Du Rietz 
(1924) did much to clear up the confusion and delimit the species 
more carefully. I had recognized the species as P. hypotropoides Nyl. 
(Hale, 1957) but later examination of Jacquin’s plate left no doubt 
that P. hypotropoides is identical with P. perforata. The norstictic 
acid easily decomposes in improperly dried herbarium specimens and 
turns the white lower side pink or reddish. Other diagnostic charac- 
ters are the distinct maculae, erect perforate apothecia, and the dis- 
tinct white zone below (pl. 2). All of the specimens seen in the early 
part of this study were from eastern North America (sce fig. 7), but 
several collections from Jamaica, Madagascar, and Ireland have ex- 
tended the range considerably. In North America the species is 
most commonly found in the canopy of oak trees, whereas the soredi- 
ate counterpart, P. hypotropa Nyl., is more common on trunks near 
the base. 

It is interesting to note that Lichen cylindricus L. (Sp. Pl. 1150. 
1753), which antedates Lichen perforatus, is based primarily on “Ti 
chenoides foliorum laciniis crinitis’”’ Dill. Hist. Muse. 149, pl. 20, fig. 42. 
1742. Dillenius’ specimen (OXF) is typical P. perforata collected in 
Pennsylvania. Linnaeus cited a second syntype from Lapland; this 
specimen (LINN) is a small Umbilicaria. Linnaeus himself later 
realized that the American specimen did not fit his concept of Lichen 
cylindricus and put this name in the umbilicate series as a synonym 
of L. proboscideus L. without further explanation. All later authors 
have followed Linnaeus’ concept without, however, realizing that L. 
cylindricus was based in large part on Parmelia perforata. 

Additional specimens examined: 


U.S.: Massacnuserts: 1 mi. west of Tolland, Hampden Co., Hale 19092 
(US); Connecticut: 1 mi. south of Colebrook, Litchfield Co., Hale s.n. (US); 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 337 


New York: Chilson Lake, Essex Co., Harris s.n. (US); Roope Isuann: Bristol, 
Howe, Lich. Nov. Angl. 39 (M); Devaware: Laurel, Sussex Co., Commons s.n. 
(PH); Onto: Near Cincinnati, Hamilton Co., Leas.n. (PH); Wisconsin: Fayette, 
Lafayette Co., Cheney 9220 (WIS); Virainia: Skinquarter, Chesterfield Co., 
Hale 14574 (US); 5 mi. northeast of Wyllesburg, Charlotte Co., Hale 15814 (US); 
3 mi. south of Laconia, Mecklenburg Co., Hale 15874 (US); Nort CaRo.ina: 
Shakelford Banks, Carteret Co., Culberson 6789 (DUKE, US); near Candler, 
Buncombe Co., Green s.n. (US); Florence, Pamlico Co., Culberson 5191 (DUKE); 
TENNESSEE: Between Del Rio and Newport, Cocke Co., Sharp TL5517 (WIS); 
Cross Roads, Overton Co., Phillips 336 (US); Sourn Caroiina: Northeast of 
Edgefield, Edgefield Co., Standley 92532 (F); Fuorrpa: 5 mi. west of Clarksville, 
Calhoun Co., Hale 21865 (US); Gainesville, Alachua Co., Nelson s.n. (FLAS); 
10 mi. east of Newport, Jefferson Co., Hale 21737 (US); 2 mi. west of Hosford, 
Liberty Co., Hale 21907 (US); Way Key, Levy Co., Drowet 11124 (F); Lovrsrana: 
Abita Springs, St. Tammany Parish, Drouet 9592a (F); southeast of New Iberia, 
Iberia Parish, Drouet 8937 (F); near Tunica, Feliciana Parish, McFarland 509 
(WIS); Texas: Carrizo Sand Hills, Wilson Co., Parks 8 (US, WIS). 

JAMAICA: Greenwood s.n. (K). 

IRELAND: Without locality, Taylor s.n. (UPS). 

Mapaaascar: Near Ft. Dauphine, Decary s.n. (LD); near Berari, Hildebrandt 
s.n. (FH). 

Additional records from the United States listed as ‘‘Parmelia hypotropoides” 
by Hale (1957) are not repeated here. 


100. Parmelia pseudonilgherrensis Asahina, Journ. Jap. Bot. 29:370, fig. 1. 
1954. PLaTE 13 
Parmelia nigrireagens Dodge, Ann. Mo. Bot. Gard. 46:145. 1959. Type 
collection: Kikandara, Toro Distr., Western Prov., Uganda, Omaston 3766c 

(K, holotype). 

Type collection: Northern Korea, Tsutani, 1936 (TNS, holotype). 

Thallus 5-10 cm. in diameter, loosely attached, mineral gray; lobes 
rotund, 8-15 mm. wide, margins sorediate, soralia linear, sorediate 
lobes sinuous; upper surface smooth, shiny, strongly white-maculate, 
submarginally sparsely sorediate; lower side black and moderately 
rhizinate, dark brown and naked in a broad zone along the margins. 
Apothecia unknown. 

Reactions. Thallus K+ yellow; medulla K—,C—, KC+ red, P—, 
atranorine, alectoronic and a-collatolic acids present. 

Parmelia pseudonilgherrensis is apparently the sorediate phase of 
P. nilgherrensis Nyl. Parmelia lobulascens Stein. & Zahlbr. has similar 
maculae and soredia but contains gyrophoric acid. These species may 
prove to be chemical strains. Although only recently described from 
Korea, P. pseudonilgherrensis seems to be common on high mountains 
in Africa. 

Additional specimens examined: 

KENYA: Kinango, Turner 6388 p. p. (K); western slopes of Mount Kenya, elev. 
3630 m., Mearns 1529 (US). UGANDA: Namwamba Valley, Ruwenzori, 


Taylor 3088 (BM); Mount Elgon, elev. 14,200 ft., Dummer 3397 (US); Kiliman- 
jaro, Héhnel, 1890 (G). 


338 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


NEPAL: Langtang Village, Polunin M179 (BM). KOREA: Kanchi-in, 
Fujikawa, Aug. 2, 1934 (TNS, US). 
101. Parmelia reparata Stirton, Scot. Nat. 4: 201. 1877-78. PLATE 14 
Parmelia virens Mill. Arg. Flora 69: 255. 1886. Type collection: Too- 
woomba, Queensland, Australia, Hartmann in 1882 (G, holotype). 


Type collection: Cave Mountain, Brisbane, Australia, Bailey (BM, 
lectotype). 

Thallus expanded, loosely adnate, up to 20 cm., coriaceous, mineral 
eray; lobes rotund, margins broadly crenate, ciliate, cilia abundant, 
1.5-2.5 mm. long; upper surface plane, shiny, strongly effigurate white- 
maculate, becoming cracked with age, soredia and isidia lacking; lower 
side black and rhizinate, brown to mottled ivory and naked in a broad 
zone along the margins. Apothecia common, stalked, up to 20 mm. 
in diameter, amphithecium rugose, strongly white-maculate, disc 
perforate; hymenium 65-70 » high; spores 6-7 13-16 y, episporium 
1 » thick; pycnidia common, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K+ yellow turning red, 
C—, KC—, P+ orange red, atranorine and salacinic acid present. 

The type of P. reparata is unfortunately not in good condition. 
The lower side is dark and apparently sparsely rhizinate almost to the 
margins, exactly as in the type of P. virens Mill. Arg. The one col- 
lection from New Zealand and all collections from America cited 
below, however, have a broad distinct bare zone below. It is with 
some hesitation therefore that we identify the latter specimens with 
P. reparata, but the other morphological similarities, the strongly 
white-maculate upper cortex and perforate apothecia, are so great as 
to leave no other choice. The problem cannot be solved without 
field work or access to good recent collections from Australia. In 
Mexico, it seems that P. reparata is very closely related to and prob- 
ably the nonsorediate counterpart of P. leucosemotheta Hue. 

Additional specimens examined: 

U.S.: AtaBama: Mobile, Mobile Co., Mohr s.n. (US). MEXICO: Veracruz: 
9 km. east of Jalapa, Hale 19406, 19394 (US); northeast of Huatusco, Hale 19529 
(US); Oaxaca: 32 mi. northwest of Oaxaca, Cain 27566a (TRT, US); Curapas: 
El Sumidero, Tuxtla Gutiérrez, Hale 20078a (US). 

AUSTRALIA: QuEEnsLAND: Brisbane, Bailey s.n. (BM). NEW ZEALAND: 
North Island, Gobby s.n. (BM); Ashburton, Allan (BM). 

102. Parmelia rigida Lynge, Ark. Bot. 13, no. 13:50, pl. 2, fig. 2. 1914. 

Type collection: Piratiny, Rio Grande do Sul, Brazil, Malme 827B 
(S, holotype). 

Thallus loosely attached, 6-12 cm. in diameter, mineral gray; 
lobes often suberect, rotund, 6-12 mm. wide, margins crenate to 
laciniate, ciliate, cilia 1-3 mm. long; upper surface shiny, minutely 
pitted, strongly white-maculate, rugose with age, soredia and isidia 
lacking; lower side black and sparsely rhizinate at the center, white 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 339 


to ivory and naked in a broad zone along the margins. Apothecia 
common, pedicellate, to 20 mm. in diameter, exciple rarely ciliate, 
amphithecium rugose, white-maculate, disc perforate; hymenium 
60-80 » high; spores variable, 7-1214-19 y, episporium 1.0-1.5 yu 
thick; pycnidia common, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ red, 
P—, atranorine and alectoronic acid present. 

Parmelia rigida is virtually indistinguishable from P. perforata 
without chemical tests. There are two main differences which may be 
cited to justify recognizing two distinct species. First, P. rigida 
occurs in extreme southern United States (fig. 29), most commonly 


Ficure 29.—Distribution of Parmelia rigida Lynge in the United States. 


near shorelines, as well as in South America, whereas P. perforata has 
not been collected in South America and has a much broader distri- 
bution in North America (fig. 7). Second, the spores of P. rigida are 
more variable and on the whole slightly larger than those of P. 
perforata. Both species occupy similar habitats in the canopy of 
deciduous trees. Field studies in Florida have shown that the two 
species rarely grow mixed at the same locality. 

Additional specimens examined: 

U.S.: NortH Caroutna: Shakleford Banks, Carteret Co., Culberson 4919 
(DUKE); Smith Island, Brunswick Co., Culberson 8005 (DUKE, US); Sours 
Carotina: Myrtle Beach, Horry Co., Culberson 9085 (DUKE); Isle of Palms, 
Charleston Co., Culberson 9049 (DUKE); Grorara: 10 mi. south of Greensboro, 


Greene Co., Culberson 6599 (DUKE, US); AvaBama: Gulf Beach, Baldwin Co., 
Hale 7263 (US); near Wilmer, Mobile Co., Hale 7167 (US); Fuoripa: 5 mi. 


340 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


south of Lokosee, Osceola Co., Hale 16975 (US); Tomoka State Forest, Volusia 
Co., Hale 17059 (US); Fort Myers, Lee Co., Standley 12898 (US); 5 mi. south of 
Clermont, Lake Co., Hale 16585 (US); Orange Park, Clay Co., Hale 17740 (US); 
5 mi. south of Panacea, Wakulla Co., Hale 21918 (US); 10 mi. east of Newport, 
Jefferson Co., Hale 21710 (US); Eastpoint, Franklin Co., Hale 21692 (US); 
St. Joe Beach, Gulf Co., Hale 21847 (US); east of Pensacola, Escambia Co., 
Hale 7997 (US); 2 mi. north of Jonathan Dickinson State Park, Martin Co., 
Imshaug 23975 (MSC); Sanford, Seminole Co., Rapp s.n. (FLAS); Mississippt: 
Near Hattiesburg, Forrest Co., Hale 7989 (US); 9 mi. south of Montrose, Jasper 
'0., McDaniel L-40 (US); Texas: Saratoga, Hardin Co., Fisher 50007 (DUKE) ; 
Barreda, Cameron Co., Runyon 1587 (MSC); Yturria Ranch, Willacy Co., 
Runyon 3821 (MSC); west of San Augustine, San Augustine Co., Hale 5234 
(US); Pineland, Sabine Co., Hale 5159 (US); Corpus Christi, Nueces Co., Palmer 
11212 (US); east of Franklin, Robertson Co., Walker 45 (US). 

BRAZIL: Rio GranpE po Sut: Piratiny, Malme 827* (8S). URUGUAY: 
Yazuarg, Dept. Tacuarembé, Rosengurtt 2861 (US). ARGENTINA: Jusvy: 
Near Quinta, Fries 27 (S). 


103. Parmelia schimperi Mill. Arg. Hedwigia 31:276. 1892. 


Type collection: Near Debra, Ethiopia, Schimper 13 (G, lecto- 
type; H, UPS, isotypes). 

Thallus 10-15 cm. in diameter, loosely adnate to bark, mineral 
gray; lobes rotund, 10-12 mm. wide, margins entire, sparsely ciliate, 
cilia 0.5-1.0 mm. long; upper surface plane, strongly white-maculate; 
lower side black and moderately rhizinate, brown and naked in a 
broad zone at the margins. Apothecia to 10 mm. in diameter, 
amphithecium rugose, maculate, disc usually perforate; hymenium 
65-70 uw high; spores 10-1217-22 w, episporium 1.0-1.5 yw thick. 

Reactions: Thallus K+ yellow; medulla K—,C—, KC-+ red, P—, 
atranorine, cryptochlorophaeic acid, and protolichesteric acid present. 

This distinctive species is still known only from several collections 
of the lectotype by Schimper. Outside of the unusual chemistry, it 
is distinguished by the strongly maculate cortex and the black lower 
side. In the absence of more material, it is difficult to decide what 
relation it might have to other species with cryptochlorophaeic acid 
such as P. abessinica Kremplh. Additional syntypes cited by Miiller 
(Geraz, Schimper 1396 and Mount Kubbi near Adoa) were not found 
at Geneva and their identity is unknown. 


104. Parmelia subcolorata Hale, sp. nov. 


Thallus late expansus, laxe adnatus, rigidus, 10-15 cm. latus, lobis 
rotundatis, usque ad 15 mm. latis, margine integris, ciliatis, ciliis 
1.0—2.5 mm. longis, superne nitidus, plus minusve valde albomaculatus, 
strato corticeo superiore 14-18 yw crasso, strato gonidiali 13-16 yp 
crasso, medulla aurantiaco-flavida pro maxima parte, partim alba, 
100-135 w crassa, strato corticeo inferiore 12-14 uw crasso, subtus 
niger, sparse rhizinosus, ambitu castaneus vel albovariegatus, late 
nudus. Apothecia usque ad 15 mm. lata, pedicellata, amphithecio 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 341 


rugoso, albomaculato, disco imperforato; hymenium 50-60 » altum; 
sporae 8-10 13-16 y, episporio 1.0-1.5 yw crasso; pycnidia numerosa, 
conidiis non visis. Thallus K+ flavescens; medulla K+, C+, 
KC-+ intensius lutea, P—, atranorinum, pigmentum K— ignotum 
continens. 

Type in the Rijksherbarium, Leiden, collected in Tinderet Forest 
Reserve, Kisumu-Londiani, Nyanza, Kenya, Aug. 1, 1949, by R. A. 
Maas Geesteranus (no. 5359; isotype in US). 

Parmelia subcolorata is a large conspicuous lichen known from only 
two localities in Africa. It differs significantly from similarly pig- 
mented species (P. myelochroa Hale, P. araucariarum Zablbr., etc.) in 
having distinct maculae, perforate apothecia, and long coarse cilia. 

CONGO: Southwest side of Lake Kivu, Nulungu, Degelius, Mar. 19, 1960 
(DEGEL). 

105. Parmelia subrugata Kremplh. Verh. Zool. Bot. Gesell. Wien 18:320. 1868. 
PLATE 16 
Parmelia latissima f. subrugata Nyl. in Fourn. Mex. Pl. 1:3. 1872. Type col- 
lection: Brazil (H, holotype). 
Parmelia cyathina Stirton, Scot. Nat. 4:252. 1877-78. Type collection: 
Near Brisbane, Australia, Bailey 3 (GLAM, holotype). 
Parmelia sinensis Hue, Nouv. Arch. Mus. Paris, ser. 4, 1:187. 1899. Type 
collection: Near Yent-ze-Hay, Yunnan, Delavay, Aug. 8, 1888 (P, lectotype). 
P. subrugata f. integra Hue, Nouv. Arch. Mus. Paris, ser. 4, 1:204, 1899. 
Type collection: Brazil, S. M. Theresa Christina (P, holotype). 
P. subrugata f. arcuata Lynge, Ark. Bot. 13, no. 13:48. 1914. Type collec- 
tion: Porto Alegre, Rio Grande do Sul, Brazil, Malme 440 (S, holotype). 
P. amaniensis Stein. & Zahlbr. Bot. Jahrb. Engler 60:526. 1926. Type 
collection: Amani, East Usambara, Brunnthaler s.n. (W, holotype). 

Type collection: Organ Mountains, Minas Gerais, Brazil, Helmret- 
chen s.n. (M, holotype). 

Thallus loosely attached, corticolous, 7-15 cm. broad, mineral gray, 
turning buff in the herbarium; lobes rotund, 7-15 mm. wide, often 
soon crowded, convoluted, and obscured by lobules, margins crenate 
to dissected-laciniate, laciniae usually becoming canaliculate, suberect, 
moderately ciliate, cilia 1-3 mm. long; upper surface plane to rugulose, 
shiny, opaque or very faintly white-maculate, distinctly maculate at 
the base of apothecia; medulla white, turning orange red frequently 
near the lower cortex; lower side black and sparsely rhizinate at the 
center, brown to ivory or mottled in a broad zone at the margins. 
Apothecia common, stalked, to 12 mm. in diameter, exciple often short 
dentate-laciniate, ciliate, amphithecium heavily rugose and white- 
maculate, disc imperforate; hymenium 100-140 uy high; spores 12-18 
26-34 u, episporium 3-4 yw thick; pycnidia abundant, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K—, C—, KC+ red, P—, 
atranorine and alectoronic acid present; medulla often pigmented, K+ 
purple, rhodophyscin present. 


342 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Des Abbayes (1962) was the first to place P. sinensis in synonymy 
under P. subrugata. These two species at first seem only distantly 
related. Typical P. subrugata has short laciniate, suberect lobes, 
rather distinct maculae, and a distinct pale zone below. Parmelia 
sinensis, known only from type collections, has more coriaceous lobes, 
less evident maculae, and darker zone below. However, both have 
alectoronic acid, large spores, and imperforate apothecia and must be 
considered as members of a single variable population. By applying 
these same criteria, we must also put P. cyathina Stirt., a coriaceous, 
strongly maculate species, and P. amaniensis Stein. & Zahlbr., a 
faintly maculate species, into synonymy. ‘The amount of variation in 
P. subugrata is distressing, but separation of the plants seems to be 
artificial and unworkable. 

Additional specimens examined: 


MEXICO: Pursuia: 5 km. west of Puebla-Veracruz State line on highway 
150, Hale 19640 (DUKE, 8, TNS, US); Curapas: El Suspiro, 10 km. N. of 
Berriozdbal, Hale 20207 (US). GUATEMALA: GuatrEmata: Volcan de Pacaya, 
Standley 58511 (MO). 

HAITI: Summit of Montagne Noire, near Kenscoff, Wetmore 2747 (MSC). 

BRAZIL: Minas Gerais: Caldas, Regnell, s.d. (S); Sitio, Vain. Lich. Bras. 
Exs. 994 (M, UPS); no locality, Burchell 1105:28 (K); Rro pE JANErRo: Serra 
dos Orgiios, Ainsworth & Gregory 523 (BM). ARGENTINA: No locality, 
Lorentz & Hieronymus, 1872-74 (M). 

MOCAMBIQUE: Massangulo, Sandrone s.n. (F). UNION OF SOUTH 
AFRICA: Capge Prov.: Woodbury, Alexandria, Héeg, July 20, 1929 (TRH); 
East London, Héeg, Dec. 20, 1929 (TRH). 

CHINA: Yunnan: Sanyingpan, Yunnan-fu, Handel-Mazzetti 640 (W); Mekong- 
Salween, Gebauer s.n. (W). 

AUSTRALIA: Queensland, Batley, 1899 (BM). 


106. Parmelia uruguensis Kremplh. Flora 61:461. 1878. 

Parmelia dusenit Zahlbr. Ann. Mycol. 6:133. 1908. Type collection: 
Carmen de Patagones, Patagonia, Argentina, Dusén 158 (W, holotype; §, 
isotype). 

P. hieronymi Lynge, Nyt Mag. Naturv. 62:88. 1925. Type collection: 
Abrededores, Pan de Azucar, Argentina, Hieronymus 34 (W, holotype). 

P. perforata var. ciliata Ceng.-Samb. Contr. Sci. Miss. Sales. Don Bosco No. 
6:45. 1930. Type collection: General Acha, Pampa, Patagonia, Argen- 
tina, Macchi, Apr. 25, 1910 (RO, holotype) [non P. perforata var. ciliata 
Nyl.]. 

Type collection: Cérdova and Conceptién, Argentina, Lorentz 
& Meronymus (M, holotype; isotypes in G, TUR). 

Thallus loosely attached to twigs, 5-8 cm. broad, olivaceous in the 
herbarium; lobes rotund, often suberect, 8-15 mm. wide, margins 
crenate, short ciliate, cilia about 1 mm. long; upper surface plane, 
shiny, heavily white-maculate, reticulately cracked with age; lower 
side brown or blackening and short rhizinate at the center, white to 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 343 


pale brown, rugose, short woolly rhizinate to the margins. Apothecia 
common, substipitate, to 10 mm. in diameter, exciple ciliate, disc 
perforate; hymenium 40-55 y high; spores 5-79-13 p, episporium 
1 » thick; pycnidia present, conidia not seen. 

Reactions: Thallus K+ yellow; medulla K+ yellow turning red, 
C—, KC—, P+ pale orange red, atranorine and salacinic acid present. 

Parmelia uruguensis differs rather significantly from other members 
of series Ornaticolae. It is, for example, the only species with salacinic 
acid. It has peculiar fine, short rhizines to the margin, much as in P. 
subcaperata Kremplh. The stalked perforate apothecia, pale margin 
below, suberect lobes, and maculae, however, place it nearer to species 
in series Ornaticolae than to any in series Subpallidae. It is appar- 
ently endemic to Argentina. 


Additional specimen examined: 
ARGENTINA: Buenos ArrEs: Tandil, Yussen 20948 (SI). 


Nomina Inquirenda 


The following names are untypifiable because the type specimens 
are presumed lost, unavailable, or too fragmentary for study. 
Omphalodium mazoense Dodge, Ann. Mo. Bot. Gard. 46:192. 1959. Type 

collection: Mazoe, Southern Rhodesia, Eyles 420 (K, holotype). 

The type collection is too fragmentary for study. It is probably a 
Parmelia species in subsection Ornaticolae, series Ornaticolae, and may 
contain alectoronic acid. 

Parmelia adspersa Vain. Hedwigia 46:168. 1907. Type collection: Lem Ngob, 
Thailand, Schmidt XV, XXXIV (TUR, syntypes; isosyntypes in C). 

All of the collections seen are fragments. The chemical reaction 
in the medulla is K+ yellowish, C—, P+ orange red; the P+ reaction 
is caused by an unknown substance. Des Abbayes (1958) reported 
this species from Africa, but I have not seen the specimen he cited. 
Parmelia arechavaletae Mill. Arg. Rev. Mycol. 10:1. 1888. Type collection: 

Montevideo, Uruguay, Arechavaleta (G, holotype). 

The type collection is fragmentary and has been damaged by 
insects. Atranorine and stictic acid were proved, indicating that the 
species, if isidiate, is probably synonymous with P. crinita Ach. 
Parmelia coriacea contexta Eschw. in Mart. Fl. Brasil. 1:209. 1833. Type 

collection: Amazon River, Martius. 

The type has not been located and may be lost. Eschweiler’s 
notation that it is near P. conspersa might mean that it is not an 
Amphigymnia species. 


729-018—65—_12 


344 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Parmelia coriacea rufa Eschw. in Mart. Fl. Brasil. 1:208. 1833; Icon. Sel. 
Crypt. p. 23, pl. 13, fig. 2. 1827. Type collection: Amazon River, 
Martius. 

The type has not been located and the illustration does not help 
in further identifying this taxon. 

Parmelia coriacea var. urceolata (Eschw.) Eschw. in Mart. Fl. Brasil. 1:207. 
1833. 

See entry under Parmelia urceolata. 

Parmelia crinita Meyer, Entw. Metamor. Fortpfl. Fl. 23. 1825 (non Ach. 
1814). Type collection: Rio de Janeiro, Brazil, Beyrich. 

The type collection has not been located; the original description 
is extremely brief. 

Parmelia crinita f. varians Merrill, Bryol. 11:95. 1908. Type collection: 
Carleton Place, Ontario, Macoun 3828. 

The type collection was not found at Farlow Herbarium. It is 
probably synonymous with P. erinita Ach. 

Parmelia flaveola Hillm. Repert. Sp. Nov. Fedde 49:197. 1940. Type collec- 


lection: Na Pali-Kona Forest Reserve, Kauai, Hawaiian Islands, Krajina 
425. 


The type specimen was destroyed in World War IT. 
Parmelia hildebrandtii var. subcetraria Jatta, Malpighia 19:171. 1905. Type 
collection: Mt. Singalang, Sumatra, Beccari 285, 286. 
The type collections have not been located. 
Parmelia melanoleuca (Willd.) Zenk. Pharm. Waarenk. 1:156. 1827. 
Lichen melanoleucus Willd. in Rom. & Usteri, Mag. fiir Bot. 4:9, fig. 2. 1788. 
Type collection: America meridionalis, s.c. 

In Zahlbruckner’s Catalogus this species is listed as a synonym 
of P. perforata (Jacq.) Ach., which does not occur in tropical America. 
A possible isotype (M) labeled ‘‘Auf Cort. Chin.” contains stictic acid 
and apparently equals P. crinita Ach. The original collection is 
probably at Berlin, but I have not had an opportunity to examine it. 
Parmelia melanothrix var. lacinulata Mill. Arg. Flora 74:376. 1891. Type 

collection: Andes, Bolivia, Pearce (K, holotype; G, isotype). 

The type is a sterile species of the series Ornaticolae with alectoronic 
acid and without soredia or isidia. It is near P. argentina Kremplh. 
but cannot be identified without spores. 

Parmelia modesta Hue, Bull. Soc. Bot. Fr. Mém. 63 (28): 6. 1916. Type col- 
lection: British Africa, Poncins. 

This species is missing from the collections at Paris. 

Parmelia munda (Harm.) Gyel. Repert. Sp. Nov. Fedde, 30:225. 1932. 

See the entry under P. trichotera f. munda Harm. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 345 


Parmelia noyoguinensis Hillm. Repert. Sp. Nov. Fedde 50:334. 1941. Type 
collection: Station Felsspitze, New Guinea, Ledermann 12852. 


The holotype specimen was destroyed in World War II and it is 
not known whether there were any isotypes. 


Parmelia paxinoides Dodge, Ann. Mo. Bot. Gard. 46:119. 1959. Type collec- 
tion: Andrangolaoka, Imerina, Madagascar, Hildebrandt (FH). 

The holotype could not be found in the Farlow Herbarium. Dodge 
often failed to annotate his type specimens with the result that they 
are filed under the original determinations and effectively lost. 
Parmelia pedicellata var. isidiosa Dodge, Ann. Mo. Bot. Gard. 46:144. 1959. 

Type collection: Bugishu, Bulambuli, Uganda, Thomas 549 p.p. (K) 

The type collection was not found at Kew. 


Parmelia perforata var. denticulata Lindsay, Trans. Roy. Soc. Edinburgh 
22:212. 1859. Type collection: Singalelah, Sikkim, Hooker (K). 
The type collection could not be found at Kew or the British 
Museum. 


Parmelia perforata var. digitata Lindsay, Trans. Roy. Soc. Edinburgh 22: 212. 
1859. Type collection: Pedra Bonita, Brazil, Gardner in 1836 (K). 


The type collection could not be found at Kew. 


Parmelia perforata var. incrassata Wedd. Mém. Soc. Nat. Sci. Nat. Cherbourg 
19:264. 1875. Type collection: fle d’Yeu. 
The type specimen has not been located. The description suggests 
Parmelia reticulata Tay). 


Parmelia perlata var. ceratophylla Mont. & v.d. Bosch. in Jungh. Pl. Jungh. 
4:442. 1855. Type collection: Mount. Ungarang, Java. 


The type specimen has not been located. 
Parmelia perlata var. cinchonarum Fée, Ess. Crypt., suppl. 119. 1837. Type 
collection: Peru. 
The type has not been found at either Paris or Geneva. 
Parmelia perlata f. innocua Wallr. Fl. Crypt. Germ. 3:521. 1831. Type collec- 
tion: Germany? 
The type collection has not been found. Wallroth lists P. perlata 
(Huds.) Ach. as a synonym. 


Parmelia perlata var. munda Harm. Bull. Soc. Sci. Nancy, ser. 2, 14:221, 
1897. Type collection: Epinal, Vosges, France, Berher. 


The type collection could not be found in the Harmand herbarium 
at Duke University. 


Parmelia perlata var. munda subvar. innocua Harm. Bull. Soc. Sci. Nancy, 
ser. 2, 14:221. 1897. 


This taxon appears to be based on the same type as P. perlata var. 
munda cited above. 


346 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Parmelia perlata var. opaca Miill. Arg. Mém. Soc. Phys. Nat. Hist. Genave 
16:372. 1862. Type collection: Geneva, Muiller (G). 


Miiller’s original specimen could not be found at Geneva. 


Parmelia perlata f. pilosa Grognot, Pl. Crypt. Cell. 56. 1863. Type collec- 
tion: Brisecou, near Autun. 
The type collection has not been found. 
Parmelia perlata subvar. punctulata Mont. in Sagra, Hist. Ile Cuba. Bot. 
42. 1838. Type collection: Near Havana, Cuba, Sagra. 
The location of the type specimen is unknown. 
Parmelia perlata f. soralifera Woron & Pachun in Pakhunova, Trud. Tiflissk 


Bot. Inst. 1:312. 1934. Type collection: Crater of Mukhera, Pak- 
hunova, 1925. 


The type specimen has not been examined. 


Parmelia perlata f. sorediifera Oxner, Bull. Jard. Bot. Kiev 1:33. 1924. Type 
collection: Minsk. 

The type specimen has not been examined. 

Parmelia platycarpa Stirt. Scot. Nat. 4:252. 1877-78. Type collection: Near 
Brisbane, Australia, Bailey (BM, holotype). 

The type collection is in such poor condition that it could not be 
used as the basis for a description. The chemical constituents are 
atranorine and stictic acid, and the spores, as Stirton found, are 
large (30-37 u long). This species differs from others with stictic 
acid in lacking cilia. 

Parmelia proboscidea var. sorediifera f. saxicola Ceng.-Samb. Nuov. 


Giorn. Bot. Ital. 45:381. 1938. Type collection: Mohulo, Kipengere, 
Tanganyika, Eusebio 13 ter. 


The type specimen has not been examined. 
Parmelia pseudovirens Gyel. Repert. Sp. Nov. Fedde 29: 288, 1931. 
See entry under P. virens 8 sorediata Mill. Arg. 


Parmelia trichotera f. dissecta Oliv. Mem. Acad. Cienc. Art. Barcelona, ser. 
3, 16:485. 1921 [not seen]. 


The type specimen has not been examined. 


Parmelia trichotera f. microphylla B. de Lesd. ex Harm. Lich. France 4:582. 
1909. Type collections: Petite Synthe, Dunkerque, Lesdain; La Massane, 
Pyrenees, Weddell. 


The specimens in Bouly de Lesdain’s herbarium were lost; no dupli- 
cates have been found in the Harmand herbarium at Duke University. 
Parmelia trichotera f. munda Harm. Lich. France 4: 582. 1909. 

The type collection could not be found in Harmand’s herbarium 
at Duke University. 


Parmelia trichotera var. plombii B. de Lesd. Rev. Bryol. Lichen. 19:235. 1950. 
Type collection: Cestas, Gironde, France, Plomb. 


The type collection has not been available for study. 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 347 


Parmelia uberrima Hue, Bull. Soc. Bot. Fr. Mém. 63:9. 1916. Type collection: 
British Africa, Poncins. 


The type has not been found in Hue’s herbarium at Paris. 


Parmelia urceolata Eschw. Icon. Pl. Crypt. 23. 1827. 

This species is illustrated by a color plate which unfortunately 
has so little detail that identification is impossible. It is obviously 
near P. argentina Kremplh. or P. subrugata Kremplh., and if the type 
is found (and typification is made), P. urceolata will be the earliest 
name for one of the species in this group. 

Parmelia urceolata var. sorediifera Miill. Arg. Flora 63:266. 1880. Type 
collection: Petropolis, Brazil, Deventer (G, holotype). 

The type specimen is an indeterminable scrap. 

Parmelia virens 8 sorediata Miill. Arg. Flora 69: 256. 1886. Type collection: 


Toowoomba, Queensland, Australia, Hartmann 49 (G, holotype). 
Parmelia pseudovirens Gyel. 


The type specimen is in poor condition. It is obviously close to 
P. leucosemotheta Hue since it is heavily maculate and contains 
salacinic acid. It could also be near P. subsumpta Nyl. No sound 
judgment can be made until additional material from Australia is 
seen. 

Nomina Rejicienda 
Parmelia glaberrima Kremplh. Flora 52: 223. 1869. Type collections: Suri- 
nam, Wuhlschlegel (not seen); Assam, India, Simons 397 (M). 

This name is a later homonym of P. glaberrima Fr. (Syst. Orb. Veg. 
1:283. 1825), a Lobaria species. The Assam specimen at M is too 
poor for adequate study. 


Parmelia latissima f. sorediata Kremplh. Journ. Mus. Godeffroy 41: 100. 1873. 
Type collection: Ovalau, Fiji, Grdffe 26(?). 


This name is a nomen nudum. 
Parmelia neohollandica Nyl. Flora 69: 324. 1886. 


Although Nylander wrote a short Latin description, there is no 
indication of the locality, except as indicated by the specific name 
neohollandica, i.e. Australia. The only specimen so labeled at Helsinki 
is an unidentifiable scrap collected by Verreaux. 


Parmelia perforata var. isidioidea Kremplh. Denkschr. Kgl. Bayer. Bot. Gesell. 
4:130. 1861. 


This is a nomen nudum. 
Parmelia perforata f. sorediifera (Mill. Arg.) Merrill, Bryol. 30:87. 1927. 
The basionym of this taxon is unknown. 


Parmelia perlata var. laciniata Nyl. Mem. Soc. Imp. Sci. Nat. Cherbourg 5: 301. 
1857. 

This is a nomen nudum; there is no specimen at Helsinki under 
this name. 


348 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Parmelia perlata var. olivetorum f. isidiophora Kremplh. Verh. Zool. Bot. Gesell. 
Wien 18:321. 1868. Type collection: Guatemala, 1841, Friedrichsthal. 


This is a nomen nudum. The type specimen was not found at 
Munich. 


Parmelia perlata var. platysmoides Nyl. ex Stizenb. Ber. St. Gall. Naturw. Gesell. 
1888-89: 157. 1890. Cited collections: Teneriffe, Perrandiére; Madeira, Mandon. 


This is a nomen nudum. The first collection is at Helsinki; it is P. 
crinita Ach. The second could not be found. 


Excluded Names 


Many species placed in the subgenus (or section) Amphigymnia in 
catalogs and floristic lists actually belong to other subgenera in Par- 
melita. Names in the following lists, based on studies of type speci- 
mens, are drawn principally from Asahina (1952), Dodge (1959), 
Hillmann (1934), Lynge (1914), Magnusson (1955), Maas Geesteranus 
(1947), Tavares (1945), Vainio (1890), and Zahlbruckner (1909, 1929). 
These species are currently being investigated in the course of mono- 
graphing other subgenera in Parmelia. They are listed here merely 
to justify their absence in lists of synonyms in this monograph. 


The following species belong in subgenus Parmelia. 
Parmelia albaniensts Dodge, Ann. Mo. Bot. Gard. 46:121. 1959. 
. anitllensis Nyl. Bull. Soc. Linn. Norm., ser 2, 3:264. 1868. 
. aptata Kremplh. in Nyl. Flora 52:291. 1869. 
badia Pers. in Gaud. Voy. Uranie 198. 1827. 
. balansae Mill. Arg. Rev. Mycol. 10:1. 1888. 
. balansae var. sorediata Miill. Arg. Rev. Mycol. 10:2. 1888. 
. btformis Vain. Phil. Journ. Sci. 4:660. 1909. 
. blastica Vain. Journ. Bot. Brit. For. 34:32. 1896. 
. canaliculata Lynge, Ark. Bot. 13, no. 13:28. 1914. 
. caperata (L.) Ach. Meth. Lich. 216. 1803. 
. caperatula (Nyl.) Nyl. Flora 68:606. 1885. 
. coralloidea (Mey. & Flot.) Vain. Acta Soc. Faun. Fl. Fenn. 7, no. 7:33. 
1890. 
. crozalstana B. de Lesd. in Harm. Lich. Fr. 4:368. 1909. 
. cyliphora (Ach.) Vain. Acta Soc. Faun. Fl. Fenn. 13, no. 6:7. 1896. 
. ecaperata Mill. Arg. Flora 74:378. 1891. 
. euplecta Stirt. Scot. Nat. 4:299. 1877-78. 
. ghatiensis Hue, Nouv. Arch. Mus. Paris, ser. 4, 1:198. 1899. 
. hansfordit Dodge, Ann. Mo. Bot. Gard. 46:127. 1959. 
hawatiensis Magn. in Magn. & Zahlbr. Ark. Bot. 31A, no. 6:106. 1944. 
. herreana Zahlbr. Cat. Lich. Univ. 6:239. 1929. 
. hypomilta Fée, Ess. Crypt., suppl. 123. 1837. 
tmmiscens Nyl. Flora 68:606. 1885. 
inhaminensis Dodge, Ann. Mo. Bot. Gard. 46:130. 1959. 
. leucozantha Mill. Arg. Flora 64:85. 1881. 
. livido-tessellata Hue, Nouv. Arch. Mus. Paris, ser. 4, 1:191. 1899. 
. martinicana Ny]. Flora 68:609. 1885. 
. metosperma (Hue) Dodge, Ann. Mo. Bot. Gard. 46:139. 1959. 


Swe VBTVNVVDVVVV VNVvVyVV VINNY 


HALE—PARMELIA SUBGENUS AMPHIGYMNIA 349 


. michauziana Zahlbr. Cat. Lich. Univ. 6:244. 1929. 
. microsticta Mill. Arg. Flora 62:164. 1879. 
natrobiensis Stein. & Zahlbr. Bot. Jahrb. Engler 60:517. 1926 
negata Nyl. Bull. Soc. Linn. Norm., ser. 2, 6:270. 1872. 
nimandairana Zahlbr. Repert. Sp. Nov. Fedde 33:55. 1934. 
nipponica Zahlbr. Bot. Mag. Tokyo 41:353. 1927. 
nylanderi Lynge, Ark. Bot. 13, no. 13:82. 1914. 
olivaria (Ach.) Fr. Lich. Scand. 1:112. 1871. 
owaniana Stirt. Trans. Glas. Soc. Field Nat. 5:213. 1877. 
perlata var. subrevoluta Mill. Arg. Flora 63:267. 1880. 
. praeperlata Nyl. Flora 69:319. 1886. 
pseudoreticulata Tav. Acta Port. Biol. 1B:138. 1945. 
. ramulicola Dodge, Ann. Mo. Bot. Gard. 46:172. 1959. 
raunkiaert Vain. Ann. Acad. Sci. Fenn. 6, no. 7:19. 1915. 
riograndensis Lynge, Ark. Bot. 13, no. 13:26. 1914. 
rudecta Ach. Syn. Lich. 197. 1814. 
ruminata Zablbr. in Magn. & Zahlbr. Ark. Bot. 31A, no. 6:107, 1944. 
somaliensis Mill. Arg. Flora 68:501. 1885. 
soredians Nyl. Flora 55:421. 1872. 
steinert Dodge, Ann. Mo. Bot. Gard. 46:125. 1959 [non P. steineri 
Gyel.]. 
subcaperatula Nyl. Flora 68:606. 1885. 
subglauca Nyl. in Gasil. Journ. de Bot. 8:126. 1894. 
subpluriformis Zahlbr. Denkschr. Akad. Wiss. Math. Naturw. Wien 83:172. 
1909. 
. subpraesignis Nyl. Lich. Environ. Paris 36. 1896. 
 tabacina Mont. & v.d. Bosch. in Mont. Syll. Gen. Sp. Crypt. 327. 1856. 
. wainioana Lynge, Ark. Bot. 13, no. 13:87. 1914. 
. wallichiana Tayl. London Journ. Bot. 6:176. 1847. 


The following species should be classified in the subgenus Xantho- 
parmelia (Vain.) Hale. 


Parmelia amphixantha Mill. Arg. Flora 71: 139. 1888. 

P. dichotoma Mill. Arg. Flora 69: 257. 1886. 

P. hypopsila Mill. Arg. Flora 70:317. 1887. 

P. impleza Stirt. Ann. Rep. Trans. Glasgow Soc. Field Nat. 1872-73: 20. 
1873. 

P. stramineonitens Zahlbr. Ann. Hist. Hofm. 11:195. 1896. 

P. subflabellata Stein. Bull. Herb. Boiss., ser. 2, 7: 639. 1907. 

P. wrightii Dodge, Ann. Mo. Bot. Gard. 46: 128. 1959. 

P. zeyheri Dodge, Ann. Mo. Bot. Gard. 46: 1382. 1959. 


VUDT DVD DVD VVVVDVUVNDVDVDD VD 


The following species have pseudocyphellae and are related to 
Parmelia cetrarioides (Del.) Nyl. The exact position of this group in 
the genus Parmelia has not yet been determined. 


Parmelia andreana Mill. Arg. Rev. Mycol. 1:169. 1879. 

P. cetrarioides (Del. ex Duby) Nyl. Flora 52: 289. 1869. 

P. ethiopica Dodge, Ann. Mo. Bot. Gard. 46:123. 1959. 

P. kernstockii Lynge & Zahlbr. in Zahlbr. Ann. Nat. Hofm. Wien 27: 271. 
1913. 

P. lobarina Zahlbr. Ann. Mycol. 10:381. 1912. 

P. manshurica Asah. Journ. Jap. Bot. 17:75. 1941. 


350 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


P. nitescens Stirt. Scot. Nat. 4:299. 1877-78. 

P. praesignis Nyl. Bull. Soc. Linn. Norm., ser. 2, 6:270. 1873. 
P. pseudolivetorum Asah. Journ. Jap. Bot. 26:16. 1951. 

P. soredica Nyl. Flora 68: 605. 1885. 


The following two species should be classified in the genus Céetraria. 


Parmelia arisanit Zahlbr. Repert. Sp. Nov. Fedde 33:57. 1934. 
P. braunsiana Mill. Arg. Flora 64: 506. 1881. 


Bibliography 


ALMBORN, O. 1948. Distribution and ecology of some south Scandinavian 
lichens. Bot. Not., suppl. 1, no. 2: 1-252. 

AsaHIna, Y. 1938. Mikrochemischer Nachweis der Flechtenstoffe. VII. Journ. 
Jap. Bot. 14:318-323. 

1952. Lichens of Japan. II. Genus Parmelia. Res. Inst. Nat. Re- 

sources 1-172. Tokyo. 

1954a. A new method in describing the relation between cortex, 

medulla, and axis of Usneae. Journ. Jap. Bot. 29:11-17. 

1954b. On the yellow pigment contained in Letharia togashit Asahina. 

Journ. Jap. Bot. 29:33, 34. 

1959. Lichenologische Notizen (150-158). Journ. Jap. Bot. 34:225- 


230. 

, and Surpata, S. 1954. Chemistry of lichen substances. Jap. Soc. 
Prom. Sci. 1-240. Tokyo. 

Berry, E. C. 1941. A monograph of the genus Parmelia in North America, 
north of Mexico. Ann. Mo. Bot. Gard. 28:31-146. 

CuLBerson, W. L., and Cunserson, C. F. 1956. The systematics of the 
Parmelia dubia group in North America. Amer. Journ. Bot. 43:678-687. 

Dece.ius, G. 1935. Das ozeanische Element der Strauch und Laubflechten- 
flora von Skandinavien. Acta Phyto. Suec. 7:1-411. 

Des ApBarEs, H. 1956. Lichens de la region Malgache. I. Mém. Inst. Sci. 
Madagascar, ser. B, 8: 1-26. 

1958. Lichens récoltés en Guinée Frangaise et en Céte d’Ivoire. Bull. 

Inst. Fr. Afr. Noire, ser. A, 20: 1-27. 

1961. Lichens récoltés a Madagascar et a la Réunion (Mission H. des 
Abbayes, 1956). Mém. Inst. Sci. Madagascar, ser. B, 10:81-121. 

Dopaz, C. W. 1953. Some lichens of tropical Africa. Ann. Mo. Bot. Gard. 
40: 271-412. 

1959. Some lichens of tropical Africa, III. Parmeliaceae Ann. Mo. 
Bot. Gard. 46:39-193. 

Du Rie, G. E. 1924. Flechtensystematische Studien. IV. Bot. Not. 
1924:329-342. 

Evans, A. W. 1943. Asahina’s microchemical studies on the Cladoniae. Bull. 
Torrey Bot. Club 70:139-151. 

GYELNIK, V. 1932. Additamenta ad cognitionem Parmeliarum III. Repert. 
Sp. Nov. Fedde 30:209-226. 

Haute, M.E., Jr. 1952. Studies on the lichen Rinodina oreina in North America. 
Bull. Torrey Bot. Club 79:251-259. 

1956a. Chemical strains of the lichen Parmelia furfuracea. Amer. 

Jour. Bot. 43:456-459. 

1956b. 2,4-Dihydroxy depsides in North American lichens. Trans. 

Kansas Acad. Sci. 59:229-232. 

1957. The identity of Parmelia hypotropoides. Bryol. 60:344-347. 

1959a. New or interesting species of Parmelia in North America. 

Bryol. 62:16—24, 


351 


352 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 


Have, M.E., Jr. 1959b. New or interesting Parmelias from North and tropical 
America. Bryol. 62:123-132. 

1960. A revision of the South American species of Parmelia deter- 

mined by Lynge. Contr. U.S. Nat. Herb. 36:1-41. 

1961. The typification of Parmelia perlata (Huds.) Ach. Brittonia 

13:361-367. 

, and Kurokawa, 8. 1964. Studies on Parmelia subgenus Parmelia. 
Contr. U.S. Nat. Herb. 36:121-191. 

HuMann, J. 1934. In Rabenhorst, Kryptogamenflora. 9(5), part 3:1-309. 

1939. Bemerkungen iiber einige Arten der Flechtengattung Parmelia. 

I. Hedwigia 78:249-267. 

1940. In Mattick, F. XIX. Die Flechten der Hawaii-Inseln. Repert. 
Sp. Nov. Fedde 49:187-206. 

Lams, I. M. 1951. On the morphology, phylogeny, and taxonomy of the lichen 
genus Stereocaulon. Canad. Journ. Bot. 29:522-584. 

LyneE, B. 1914. Die Flechten der ersten Regnellschen Expedition. Die 
Gattungen Pseudoparmelia gen. nov. und Parmelia Ach. Ark. Bot. 13, 
no. 13:1-172. 

Maas Grrsreranus, R. A. 1947. Revision of the lichens of the Netherlands. 
I. Parmeliaceae. Blumea 6:1-199. 

Maenusson, A. H. 1955. <A catalogue of the Hawaiian lichens. Ark. Bot., 
ser. 2, 3, no. 10:223-402. 

NYLANDER, W. 1869. Circa reactiones Parmeliarum adnotationes. Flora 52: 
289-293. 

1874. Addenda nova ad lichenographiam europaeam. Flora 57:305- 


318, 


1885. Parmeliae exoticae novae. Flora 68:605-615. 

RuNnEMARK, H. 1956. Studies in Rhizocarpon. I. Taxonomy of the yellow 
species in Europe. Opera Bot. 2, no. 1: 1-152. 

SANTESSON, R. 1952. Foliicolous lichens I. Symb. Bot. Ups. 12, no. 1:1-590. 

Tavares, C. N. 1945. Contribuicéo para o estudo das Parmelidceas Portu- 
guesas. Port. Acta. Biol. 1B:1-210. 

Vainio, E. A. 1890. Etude sur la classification naturelle et la morphologie des 
lichens du Brésil. Acta Soc. Faun. Fl. Fenn. 7, no. 7:27-66. 

ZAHLBRUCKNER, A. 1909. Ergebnisse der botanische Expedition der Akademie 
der Wissenschaften nach Siidbrasilian 1901. Lichenes. Denkschr. Akad. 
Wiss. Naturw. Wien 83:87-211. 

———. 1929. Catalogus lichenum universalis. 6:1-618. 


INDEX 


(To species and infraspecific taxa. 


Synonyms in italics; page numbers of principal 


entries in boldface) 


Cetraria corrugis, 335 
Cladonia chlorophaea, 211 
cryptochlorophaea, 211 
grayi, 211 
Lichen cylindricus, 336 
melanoleucus, 344 
perforatus, 195, 335, 336 
perlatus, 195, 300 
proboscideus, 336 
Omphalodium mazoense, 343 
Parmelia aberrans, 199, 211, 212, 213, 214, 
215, 216, 218, 220, 229, 269, 276 
abessinica, 202, 207, 208, 211, 212, 
215, 217, 220, 234, 319, 320, 325, 
326, 340 
abessinica var. nuda, 236 
abessinica var. sorediosa, 325 
abyssenica var. glabrior, 320 
abyssinica, 320 
abnuens, 207, 208, 216, 220, 233, 
277, 278, 289 
adspersa, 343 
africana, 236 
albaniensis, 348 
aldabrensis, 216, 218, 222, 234, 321 
allardii, 297, 298 
allenit, 326 
amaniensis, 341, 342 
amboimensis, 216, 220, 233, 235, 
278 
amphixantha, 349 
andina, 202, 203, 204, 206, 216, 219, 
232, 236, 239, 293 
andreana, 349 
annae, 314, 315 
antillensis, 348 
appendiculata, 207, 218, 220, 232, 
248, 251, 278, 283 
appendiculata f. disparilis, 247 
aptata, 348 
araucariarum, 202, 218, 220, 230, 
238, 251, 256, 279, 302, 341 
arechavaletae, 343 
argentina, 204, 207, 214, 220, 233, 
278, 318, 319, 322, 323, 332, 344, 347 
arisanii, 350 


Parmelia— Continued 

arnoldii, 200, 212, 214, 222, 223, 231, 
232, 277, 279, 296, 297, 301, 309 

austrosinensis, 200, 202, 203, 204, 
208, 216, 219, 223, 230, 236, 237, 
238, 239, 245, 324, 326 

badia, 348 

balansae, 348 

balansae var. sorediata, 348 

bangii, 200, 218, 220, 230, 281, 292 

biformis, 348 

blanchetii, 216, 220, 233, 235, 281, 
289 

blastica, 348 

bogoriensis, 267 

brauniz, 303 

braunsiana, 350 

breviciliata, 214, 219, 224, 233, 235, 
264, 282, 295, 299, 303 

brisbanensis, 312 

callitricha, 277, 278 

canaliculata, 348 

caperata, 193, 227, 348 

caperata f. isidiosa, 271 

caperata var. madagascariacea, 275 

caperatula, 348 

capitata, 258 

capitulifera, 248 

catharinensis f. isidiosa, 284 

cazengensis, 239 

ceracea, 314, 315 

cetrarioides, 349 

cetrarioides f. integra, 245 

cetrata, 242 

cetrata var. hypotropoides, 328 

chiapensis, 214, 220, 233, 234, 319, 
322, 323 

chlorocarpa, 284 

chrysantha, 275, 276 

ciliata, 335 

cinerascens, 271 

cinereoplumbea, 292 

cirrhata, 288 

claudelit, 308 

claudelit var. clemensiae, 241 


353 


304 


Parmelia— Continued 


conformata, 204, 216, 218, 220, 228, 
229, 270, 275 

conformata f. ciliolifera, 270 

contocarpa, 300 

conspersa, 343 

cooperi, 202, 204, 216, 220, 225, 231, 
293, 319, 323 

coralliformis, 200, 217, 220, 229, 
320, 324 

coralloidea, 227, 264, 348 

coriacea contexta, 343 

coriacea var. perforata, 335 

coriacea var. perlata, 300 

coriacea rufa, 344 

coriacea var. urceolata, 344 

corniculans, 207, 214, 220, 233, 234, 
235, 283, 298 

cornuta, 207, 218, 220, 232, 283 

cornuta var. crocea, 278 

corrugis, 335 

corrugts var. imperialis, 315 

corrugis f. parmata, 335 

corrugis var. sorediata, 328 

crassescens, 216, 220, 232, 240, 253, 
262 

crinita, 199, 202, 208, 218, 219, 222, 
223, 229, 277, 284, 290, 301, 304, 
311, 313, 317, 3438, 344, 348 

crinita var. eciliata, 289 

crinita var. inactiva, 297 

crinita f. pilosella, 284 

crinita f. varians, 344 

cristata, 278 

cristifera, 200, 202, 203, 217, 219, 
230, 241, 246, 254, 259, 261, 308 

cristifera f. cinerata, 241 

cristifera f. pallida, 300 

crocea, 279 

crocoides, 217, 220, 232, 244 

crozalsiana, 348 

cryptoxantha, 200, 217, 218, 220, 
230, 287 

cyathina, 341, 342 

cyliphora, 348 

dalei, 236 

defecta, 202, 216, 220, 230, 244, 264 

delicatula, 217, 218, 220, 227, 232, 
269, 271, 272 

diacidula, 203, 215, 216, 220, 231, 
287 

dichotoma, 349 


INDEX 


Parmelia— Continued 


dilatata, 202, 204, 216, 219, 223, 
230, 231, 238, 242, 245, 253, 259, 
261, 262, 288, 309 

direagens, 203, 214, 218, 220, 231, 
288 

disparilis, 207, 216, 220, 232, 236, 
247, 268, 279 

dolosa, 334, 335 

dominicana, 216, 218, 221, 222, 
230, 248, 275 

dusenti, 342 

eborina, 216, 220, 232, 249, 250 

ebulliens, 218, 220, 232, 250 

ecaperata, 348 

eciliata, 206, 218, 220, 222, 233, 235, 
282, 289 

endosulphurea, 202, 218, 219, 221, 
228, 251, 256, 312 

erasmia, 202, 214, 217, 218, 220, 
229, 255, 290, 294 

erecta, 335 

erubescens, 314 

ethiopica, 349 

euneta, 207, 215, 219, 234, 235, 319, 
325, 328 

euplecta, 348 

eurycarpa, 289 

eurysaca, 204, 217, 220, 234, 235, 
290, 291, 294, 308 

fasciculata, 200, 216, 218, 220, 228, 
246, 252, 261, 291, 292, 324, 325 

fatiscens, 252 

flaventior, 227 

flaveola, 344 

flavescens, 217, 218, 220, 229, 270, 
272 

flavotincta, 200, 215, 220, 229, 253, 
261, 291 

fracta, 200, 216, 218, 220, 230, 292 

gardneri, 241 

gattefosset, 328 

ghattensis, 348 

glaberrima, 347 

glaberrima B flavescens, 272 

glaberrima f. flavescens, 272 

glaucocar pa, 320 

glaucocar poides, 212, 225, 320 

gosswetlert, 241 

grayana, 203, 217, 220, 223, 232, 
292 

hababiana, 202, 207, 211, 212, 215, 
217, 219, 225, 231, 319, 320, 325 


INDEX 355 


Parmelia—Continued 


haitiensis, 212, 317, 318, 319 

hanningtoniana, 207, 215, 220, 234, 
319, 327 

hansfordii, 348 

hawaiiensis, 348 

hendrickxii, 326 

herreana, 348 

hieronymi, 342 

hildebrandtii, 241 

hildebrandtii var. ciliata, 320 

hildebrandtii f. nuda, 236 

hildebrandtii f. sorediosa, 241, 242 

hildebrandtii var. subcetraria, 344 

hololoba, 202, 206, 216, 220, 234, 
293 

hypoleucina, 328 

hypoleucina var. subincana, 328 

hypomilta, 348 

hypomiltoides, 200, 202, 214, 217, 
218, 220, 230, 255, 290, 293, 305 

hypopsila, 349 

hyporysalea, 236 

hyporysalea var. cinerascens, 238 

hyporysalea var. menyhartii, 236 

hypotropa, 202, 203, 205, 207, 216, 
218, 219, 223, 231, 319, 328, 336 

hypotropa var. imperialis, 315 

hypotropa f. parmata, 335 

hypotropa var. sorediata, 328 

hypotropoides, 335, 336, 337 

imerinensis, 241 

immiscens, 348 

amperforata, 314 

implexa, 349 

inactiva, 262 

inexspectata, 218, 220, 233, 235, 294 

inhaminensis, 348 

internexa, 285 

kauatensis, 245 

kernstockii, 227, 349 

koyaensis, 258 

laongit, 322 

latissima, 202, 203, 217, 222, 232, 
241, 242, 244, 253, 255, 256, 268 

latissima var. corniculata, 267, 268 

latissima f. cristifera, 241 

latissima f. flavescens, 272 

latisstma f. flavida, 292 

latissima f. tstdiosa, 264 

latissima f. microspora, 282 

latissima var. minima, 267 

latissima f. sorediata, 347 


Parmelia—Continued 

latissima f. subrugata, 341 

leucosemotheta, 202, 217, 220, 231, 
316, 320, 325, 330, 331, 338, 347 

leucosemotheta f. isidiata, 317 

leucoxantha, 348 

leucorantha f. firma, 315 

lindmanii, 251, 252 

litoralis, 267 

livido-tessellata, 348 

livido-tessellata f. ablephara, 328 

lobarina, 349 

lobulascens, 215, 220, 231, 319, 
331, 337 

lobulascens f. isidiostssima, 245 

lophogena, 199, 215, 220, 229, 295 

luzonensis, 258 

lyngeana, 279 

maclayana, 214, 220, 234, 283, 295, 
303 

macrocarpoides f. phyllophora, 322 

madagascariacea, 212, 213, 219, 270, 
275, 276 

magna, 271, 272 

mannit, 303 

manshurica, 349 

margaritata, 200, 212, 214, 217, 
220, 231, 280, 296 

martinicana, 348 

mauriensis, 272, 273 

mazima, 308 

meiosperma, 348 

meiosperma var. ecklonii, 264 

melanoleuca, 344 

melanothrix, 202, 207, 217, 220, 
233, 278, 319, 322, 332, 335 

melanothriz var. argentina, 322 

melanothrix var. lacinulata, 344 

melanothriz f. microspora, 322 

mellissii, 199, 204, 214, 219, 229, 
285, 290, 294, 297, 310, 335 

menyhartii, 236 

meridionalis, 239 

merrillii, 207, 216, 220, 233, 278, 
283, 298, 300 

mesogenes, 217, 218, 219, 220, 232, 
254, 294 

mesotropa, 202, 215, 220, 232, 
255, 259, 262 

mesotropa var. compactior, 239 

mesotropa f. sorediosa, 241 

michauxiana, 349 

microdactyla, 271, 272 


356 


INDEX 


Parmelia— Continued 


microsticta, 349 

miranda, 217, 218, 220, 230, 273, 
274, 275 

modesta, 344 

mueller-argoviensis, 328 

munda, 344 

myelochroa, 202, 218, 220, 232, 
238, 256, 302, 341 

nairobiensis, 349 

natalensis, 200, 214, 220, 230, 306, 
319, 332 

negata, 349 

neghelliensis, 260 

neglecta, 258 

neocaledonica, 267 

neohollandica, 347 

nigeriensis, 320 

nigrireagens, 337 

nilgherrensis, 202, 214, 220, 233, 
234, 280, 319, 333, 337 

nilgherrensis f. subciliarts, 325 

nimandairana, 349 

nipponica, 349 

nitens, 263 

nitens f. isidiosa, 260 

nitescens, 350 

novoguinensis, 345 

nyasensis, 275, 276 

nylanderi, 349 

ochroglauca, 200, 217, 218, 220, 
230, 269, 274 

odontata, 247 

olivaria, 349 

olivetoroides, 239 

olivetorum var. esorediata, 236 

olivetorum var. hypomelaena, 238 

olivetorum var. hyporysalea, 236 

olivetorum var. sorediosa, 238 

ornatula, 207, 214, 220, 234, 295, 
299, 303 

overeemit, 267 

owaniana, 349 

pachyspora, 216, 220, 233, 299, 300 

pancheri, 214, 222, 232, 236, 257 

paraguariensis, 236 

paulensis, 199, 202, 207, 217, 219, 
220, 229, 319, 334 

paxinoides, 345 

pedicellata, 236 

pedicellata var. isidiosa, 345 

pedicellata var. subbullata, 236 

peralbida, 216, 220, 228, 257, 263 


Parmelia— Continued 

perforata, 197, 202, 203, 205, 207, 
212, 213, 214, 216, 222, 223, 227, 
234, 235, 314, 319, 321, 329, 335, 
339, 344 

perforata var. ciliata, 335, 342 

perforata var. claudelit, 308 

perforata var. denticulata, 345 

perforata var. digitata, 345 

perforata var. incrassata, 345 

perforata var. isidioidea, 347 

perforata f. sorediifera, 347 

perforata var. ulophylla, 241, 242 

perlata, 200, 202, 204, 218, 219, 223, 
227, 231, 274, 277, 281, 300, 301 

perlata var. ceratophylla, 345 

perlata var. ciliata f. dissectula, 284 

perlata var. cinchonarum, 345 

perlata var. coralloidea, 264 

perlata var. dissectula, 284 

perlata var. flavogranulosa, 248 

perlata f. innocua, 345 

perlata var. laciniata, 347 

perlata var. munda, 345 

perlata var. munda subvar. in- 
nocua, 345 

perlata var. olivetorum f. isidio- 
phora, 348 

perlata var. opaca, 346 

perlata f. pilosa, 346 

perlata var. platyloba, 264 

perlata var. platysmoides, 348 

perlata subvar. punctulata, 346 

perlata f. soralifera, 346 

perlata f. sorediifera, 346 

perlata var. subrevoluta, 349 

perlata var. zanthina, 275 

permutata, 215, 216, 218, 219, 230, 
302, 307 

persulphurata, 312 

petropoliensis, 315 

pilosella, 284 

pilosella f. excrescens, 284 

piloselloides, 300, 301 

platycarpa, 346 

platyphyllina, 267 

pooliz, 304 

praeperlata, 349 

praesignis, 291, 350 

praesorediosa, 202, 208, 215, 219, 
221, 224, 230, 256, 258, 262, 293, 
326 

praetervisa, 264 


INDEX 357 


Parmelia— Continued 


praetervisa var. flavicans, 251 

proboscidea, 284, 313 

proboscidea var. aspera, 317, 318 

proboscidea f. bulbtfera, 304 

proboscidea var. corallina, 310 

proboscidea var. dissectula, 284 

proboscidea f. incrassata, 282 

proboscidea var. ornatula, 313 

proboscidea f. sorediifera, 304 

proboscidea var. sorediifera, 304 

proboscidea var. sorediifera f, saxi- 
cola, 346 

proboscidea var. xanthina, 275, 276 

procera, 214, 222, 234, 295, 303 

protoflavescens, 272 

protosorediata, 300 

protovirens, 317 

pseudocatharinensis, 284 

pseudocrinita, 199, 215, 220, 229, 
303 

pseudo flavescens, 272 

pseudohyporysalea, 306, 307 

pseudolivetorum, 350 

pseudonilgherrensis, 202, 214, 220, 
231, 319, 331, 333, 334, 337 

pseudoreticulata, 349 

pseudotinctorum, 199, 216, 220, 
228, 260 

pseudotinctorum f. perrugosa, 260 

pseudovirens, 346, 347 

radians, 271, 272 

rampoddensis, 204, 214, 219, 231, 
232, 294, 297, 302, 304, 307 

ramulicola, 349 

ramuscula, 200, 217, 221, 228, 253, 
261, 292, 325 

raunkiaeri, 349 

recipienda, 212, 314, 315 

reparata, 202, 217, 221, 234, 320, 
330, 338 

resupina, 289 

reticulata, 345 

rigida, 207, 212, 213, 214, 220, 234, 
319, 322, 338 

rimulosa, 200, 214, 220, 230, 287, 
292, 305, 333 

riograndensis, 349 

rissoensis, 236 

robertyi, 267 

robusta, 245, 246 

rubifaciens, 203, 216, 220, 230, 261 


Parmelia—Continued 

rudecta, 210, 349 

ruminata, 349 

saccatiloba, 216, 221, 228, 258, 262 

saccattloba f. membranacea, 255 

sanctae-crucis, 258, 260 

sancti-angelii, 203, 215, 219, 230, 
231, 288, 302, 306, 309 

saxatilis, 206 

schimperi, 215, 220, 233, 319, 340 

schweinfurthii, 284 

schweinfurthit f. sorediata, 300 

segreganda, 323 

setchellii, 199, 216, 221, 228, 263 

siamensis, 333 

stebert, 245 

simodensis, 292 

sinensis, 341, 342 

sinensis f. istdiata, 327 

somaliensis, 349 

soredians, 227, 349 

soredica, 350 

soredica var. neghelliensis, 260 

soyauxii, 202, 216, 220, 232, 244, 
263, 282 

steineri, 349 

stramineonitens, 349 

stuhlmannii, 260 

stuppea, 200, 204, 217, 219, 223, 
231, 277, 291, 308 

subarnoldii, 203, 216, 219, 231, 
246, 309 

subbullata, 236 

subcaperata, 202, 207, 211, 212, 215, 
217, 221, 234, 235, 314, 325, 343 

subcaperata f. ciliata, 271 

subcaperatula, 349 

subcetrarioides, 258 

subciliaris, 325 

subcolorata, 218, 220, 232, 319, 
340 : 

surcorallina, 199, 216, 217, 220, 
229, 310 

subcrinita, 197, 199, 217, 219, 229, 
273, 285, 310, 317 

subflabellata, 349 

subglauca, 349 

subincana, 328 

subinvoluta, 304, 305 

submesotropa, 241 

subpluriformis, 349 

subpraesignis, 349 

subproboscidea, 322 


358 


Parmelia—Continued 


subregressa, 255 

subrugata, 207, 214, 219, 233, 313, 
319, 322, 323, 341, 347 

subrugata f, arcuata, 341 

subrugata f. integra, 341 

subrugata var. mexicana, 322 

subsumpta, 202, 207, 211, 212, 215, 
217, 218, 219, 230, 231, 314, 
315, 330, 347 

subtinctoria, 202, 207, 211, 212, 
215, 217, 219, 224, 229, 311, 
317, 318 

sulcata, 206 

sulphurata, 197, 218, 219, 224, 228, 
251, 284, 312 

sulphurata f. nuda, 256 

tabacina, 349 

tephrina, 247 

thomastt, 236 

tinctorum, 204, 210, 216, 219, 224, 
227, 228, 235, 261, 262, 264 

tinctorum var. chrysophora, 251 

tinctorum var. endosulphurea, 251 

tinctorum var. inactiva, 262 

trichotera, 300, 308 

trichotera var. claudelii, 308 

trichotera f. dissecta, 346 

trichotera f. microphylla, 346 

trichotera f. munda, 344, 346 

trichotera var. plombii, 346 

trichotera var. subincana, 328 

tuckermanit, 284, 285 

uberrima, 347 

urceolata, 344, 347 


INDEX 


Parmelia— Continued 


urceolata var. cladonioides, 322 

urceolata var. melanothriz, 332 

urceolata var. nuda, 315 

urceolata var. sorediifera, 347 

uruguensis, 207, 217, 220, 234, 319, 
342 

velutina, 317, 318 

virens, 338 

virens f. isidiosa, 317 

virens 8 sorediata, 347 

viridiflava, 216, 218, 220, 230, 273, 
274, 275 

wainii, 204, 207, 214, 220, 234, 282, 
295, 303, 313 

wainioana, 349 

wallichiana, 349 

wernert, 328 

wrightii, 349 

xanthina, 199, 211, 212, 213, 214, 
217, 218, 219, 229, 269, 270, 274, 
275 

zanthina f. aberrans, 269, 276 

zanthina var. ciliata, 269 

xanthina f. isidiosa, 275 

zanthina var. subeciliata, 270 

yunnana, 333 

zambesica, 236 

zeyheri, 349 

zollingeri, 202, 204, 216, 219, 232, 
233, 235, 248, 250, 253, 255, 262, 
267 


Parmotrema gattefossei, 328 


wernert, 328 


Physica endococcinea, 217 


U.S. GOVERNMENT PRINTING OFFICE: 1968