SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 25 

A Revision of the Lichen Genus 

Hypotrachyna (Parmeliaceae) 

in Tropical America 

Mason E. Hale, Jr. 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1975 
ABSTRACT 

Hale, Mason E., Jr. A Revision of the Lichen Genus Hypotrachyna (Parmeliaceae) in Tropical America. Smithsonian Contributions to Botany, number 25, 

73 pages, 20 figures, 1975.-A revision is made for 77 species of Hypotrachyna 

occurring in tropical America. Fifty-eight of these are endemic to the New 

World. These species are classified loosely in chemical constellations based on 

the secondary depside-depsidone products. The possible role of hybridization 

of fertile species and the evolution of vegetative morphs are discussed. Five new 

species are described: H. andensis, H. lopezii, H. partita, H. producta, and 

H. protenta. The remaining 72 species are transferred from the genus Parmelia 

to Hypotrachyna. 

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded 

in the Institution’s annual report, Smithsonian Year. SI PRESS NUMBER 5273. SERIES COVER DESIGN: 

Leaf clearing from the katsura tree Cercidiphyllurn japonicum Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data. 

Hale, Mason E. 

A revision of the lichen genus Hypotrachyna (Parmeliaceae) in Tropical America. 

(Smithsonian contributions to botany, no. 25) 

Includes index. 

Supt. of Docs. no.: SI 1.29:25 

1. Hypotrachyna. 2. Lichens-Latin America. I. Title. 11. Series: Smithsonian Institution. 

QK1 .S2747 no. 25 [QK585.P2] 581’.08s [589’.1] 74-32050 

Smithsonian contributions to botany, no. 25. 
Contents 

Introduction ...................................................... 

History of Hypotrachyna Research ............................ 

Collections ........................................... 

Morphological Characters .......................................... 

Chemical Characters ............................................... 

Speciation in Hypotrachyna ........... ........ ....... 

Classification of Hypotrachyna .... .................... .. 

The Hypotrachyna Flora ................................ .... 

Habitats and Ecology .............................................. 

Phytogeographical Relationships .................................... 

Nomenclature ..................................................... 

Keys to the Species ................................................ 

Species Treatment ................................................. 

Literature Cited ..................................... 

Index ....... ..................... 

Page 

1 

2 

3 

3 

8 

11 

15 

16 

17 

18 

19 

19 

23 

69 

72 

... 

111 

A Revison of the Lichen Genus 

Hypotrachyna (Parmeliaceae) 

in Tropical America 

Mason E. Hale, Jr. 

Introduction 

I began study of the Parmeliaceae in 1958 with 

the realization that synonymy and nomenclature of 

the nearly 600 species in the family could be solved 

only through a world-level revision. A monograph 

of the relatively conservative genus Parmotrema 

Massalongo (as Parmelia subgenus Amphigymnia) 

was completed in 1964 (Hale, 1965) along with a 

preliminary study of the other tropical sections of 

the collective genus Parmelia (Hale and Kurokawa, 

1964). 

Although I had examined all the type specimens 

and a considerable body of herbarium material, 

two factors mitigated against a prompt completion 

of a final monograph for all groups: (1) the inadequacy of chemical techniques available prior to 

1965 that were needed to resolve certain chemical 

groups such as the barbatic and lividic acid groups 

and even separation of the norstictic acid group; 

and (2) the lack of sufficient specimens and field 

experience in tropical America, where many new 

species were obviously yet to be found. 

The development and standardization of thinlayer chromatography has answered the problem of 

chemical techniques (see C. Culberson, 1972a). 

We can now routinely achieve resolution heretofore 

impossible and make accurate identifications of 

the lichen substances, A number of previously unMason E. Hale, Jr., Department of Botany, National Museum 

of Natural History, Smithsonian Institution, Washington, D.C. 

20560. 

known substances have been isolated and described. 

The few remaining problems will eventually be 

solved with high-performance liquid chromatography and mass spectrometry (C. Culberson, 

1972b). 

The inadequacy of collections will probably 

never be fully remedied, but we now have access 

to many more collections than were available in 

1965. I have had the opportunity of visiting several 

tropical areas where I could appreciate more fully 

the habitats and ecology of the species, data which 

are only rarely well documented on herbarium 

labels. 

The time has come, therefore, when a "final" 

summary can be made for the parmelioid genera. 

This paper summarizes the genus Hypotrachyna 

(Vainio) Hale for tropical America, including the 

region from the West Indies and Mexico southward to temperate Argentina and Chile. The 19 

species known from North America proper are not 

included, but actually only one (H. vz'rginka) is 

endemic there and the remaining 18 also occur in 

tropical '4merica. 

ACKNOWLEDGMENTS.-~ wish to reiterate thanks 

to the curators-previously acknowledged in Hale 

(1965) and Hale and Kurokawa (1964)-who SO 

generously loaned and in many cases reloaned 

valuable specimens. Special thanks are due Dr. S. 

Kurokawa, whose collaborative help in 1960-1964 

speeded up study of hundreds of type specimens. 

I am extremely grateful to Dr. M. Lopez Figueiras 

of Universidad de 10s Andes for the opportunity 

1 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

to collect extensively with him in the Venezuelan 

Andes. Mr. Eric Lewis, Ms. Marianne See, and Dr. 

Michael Wirth helped at various times with testing of specimens by thin-layer chromatography. 

The scanning-electron microphotographs were 

taken by Mr. Walter Brown of the Smithsonian 

Scanning-Electron Microscope Laboratory. 

Field studies in Mexico (1960) were supported 

by the National Science Foundation, and in Dominica by the Morden-Smithsonian Expedition 

(Hale, 1971a). The work in the past several years 

has been supported by grants from the Smithsonian 

Research Foundation. 

History of Hypotrachyna Research 

Studies on Hypotrachyna (and for all other tropical Parmelias) appeared only as scattered floristic 

lists prior to 1890. Krempelhuber, Tuckerman, 

Muller Argau, Nylander, and others received for 

identification small lots of lichens (usually collected 

by phanerogamic botanists) and published the 

results without illustrations, keys, or really useful 

descriptions. There was little cooperation between 

workers or sharing of information or specimens, 

Vainio (1890) was the first lichenologist to make 

a truly comprehensive study of tropical lichens. He 

collected extensively in Rio de Janeiro and Minas 

Gerais, Brazil, in 1885. The resulting publication, 

“Etude sur la Classification Naturelle et la hlorphologie des Lichens du Bresil,” remains a classic and 

Vainio’s well-curated herbarium in Turku and his 

exsiccate, Lichenes brarilienses-distributed, as far 

as I have been able to determine, to BM, FH, G, K 

(now at BRI), RI, P, and UPS-attest his tremendous ability and industry. When identifying specimens, Vainio used Nylander’s herbarium and also 

made a study trip to the Paris museum. This enabled him to present a more definitive study with 

a low level of synonymy. 

Vainio collected 13 species now classified in the 

genus Hypotrachyna: H. brasiliana (Nylander) 

Hale, H. consirnilis (Vainio) Hale, H. dactylifera 

(Vainio) Hale, H. flavida (Zahlbruckner) Hale (as 

Parmelia flava Krempelhuber), H. gracilescens 

(Vainio) Hale, H. intercalanda (Vainio) Hale, H. 

rnicroblasta (Vainio) Hale, H. novella (Vainio) 

Hale, H. plzrriformis (Nylander) Hale, H. revoluta 

(Floerke) Hale, H. subaflinis (Zahlbruckner) Hale 

(as Parmelia aflinzs Vainio), and H. velloziae 

(Vainio) Hale. Material he identified as Parmelia 

gracilis (Muller Argau) Vainio is misidentified 

H. physcioides. Altogether, Vainio described eight 

of these species as new. In spite of his efforts, however, only a portion of the Hypotrachyna species 

were collected; the general region of southeastern 

Brazil is now known to have nearly 40 species. 

The next major collecting program also was 

undertaken by a lichenologist, G. A. A. Malme, 

under the Regnell expeditions in the 1890s, primarily in Mato Grosso. The magnificently prepared 

specimens, now preserved in Stockholm, were 

studied mostly by Malme himself, but Parmelia 

specimens were turned over to B. Lynge, a Norwegian lichenologist who had no previous experience 

in tropical species, and this at a time when Vainio 

was the most logical worker and acknowledged 

authority. Lynge, however, did a creditable job, 

dealing with 110 species, of which 61 were proposed 

as new (Lynge, 1914, 1917, 1925). Only 18 of these 

species are now classified in Hypotrachyna. Lynge 

relied heavily on Vainio’s exsiccate preserved at 

UPS, saw some Nylander types, and visited Zahlbruckner in Vienna. His publications contain much 

valuable data and some excellent illustrations. 

The only other work that could be considered as 

a flora with keys and descriptions is that by Zahlbruckner (1909), completed just before Lynge’s 

study. It was based on collections made by a nonlichenologist, V. Schiffner, in the Sao Paulo area. 

The specimens are preserved at WU and W with 

duplicates at BM. 

Nore recently, Vareschi (1962) compiled a catalog of Venezuelan Pal-meliac Fvith many illustrations, but the species identifications have not been 

checked b) a monographer. 

My own field experience in tropical America 

began in Mexico, which has a large Hypotrachyna 

flora. I have also collected intensively in the Lesser 

Antilles, Trinidad, Guatemala, Panama, and Venezuela. These excursions have given me some idea 

of the ecological behavior of the genus from sea 

level to 3700 m, and while the area covered is but 

a small fraction of the total land mass of tropical 

America, it seems reasonable to extrapolate from 

this experience in discussing the habitats where the 

species might be expected to occur. 
NUMBER 25 3 

Collections 

Any revision is only as good and complete as the 

specimens available for study. Tropical America is 

a vast area, some parts of which have been relatively well explored, others are completely unknown 

and often inaccessible. Even a country as seemingly 

small as Panama has great tracts of unexplored rain 

and cloud forest that cannot be reached except by 

a major expeditionary effort. On the other hand, 

some areas, while uncollected, would probably have 

few species of Hypotrachyna. Such areas include, 

in particular, the Amazon Basin, which has a generally low elevation and a hot moist climate, hardly 

conducive to a rich foliose lichen flora. Deserts or 

very dry regions as in northern Chile, northern 

Mexico, and Baja California, and even northeastern Brazil, also are poor prospects for 

Hypotrachyna. 

The major areas of virgin forest in the tropics 

are, of course, being destroyed-cut over, burned, 

and colonized-as quickly as they can be reached. 

Such areas include even the higher elevations where 

Hypotrachyna is so abundant but where the soils 

are ideal for vegetable farming. 

The following list shows the countries where 

Hypotrachyna has been collected, the principal collectors, and the herbaria where their collections 

are housed: 

BAHAMAS: Britton (XY). 

MEXICO: Arsene (US), Galeotti (P), Hale (US), Wirth (US). 

WEST INDIES (all islands): Imshaug (ILISC). 

CUBA: Hioram (US), Jt’right (FH, L, P, UPS, US). 

JAMAICA: Morton (US), Orcutt (US), Plitt (US). 

DOMINICAN REPUBLIC: Allard (US). 

VIRGIN ISLANDS: Raunkiaer and Boergesen (C, TUR). 

GUADELOUPE: Culberson (DUKE), Duss (TUR), Husnot (H, 

DOMI~ICA: Elliott (BM, TUR), Hale (US). 

ST. VINCENT: Elliott (BM, TUR). 

TRINIDAD: Hale (US). 

GUATEMALA: Hale (US), Standley (F, US). 

HONDURAS: tandley (F). 

COSTA RICA: Culberson (DUKE), Pittier (G, US). 

PANAMA: Hale (US), Scholander (US). 

COLOMBIA: Cuatrecasas (US), Killip and Smith (US), Lindig 

VESEZUELA: Dennis (BM), Hale (US), Magdefrau (M), Poelt 

FRENCH GUYANA: Leprieur (P), 

PERU: Bryan (WIS), Gomez (US), Herrera (US), Iltis (TYIS). 

ECUADOR: Jameson (BM), Meyer (W). 

BOLIVIA: Mandon (H). 

P). 

(BM [original], FH, H, M, P), Weir (BM). 

(M, US), Santesson (S), Vareschi (VEN). 

CHILE: Gay (P), Imshaug (MSC), Mahu (personal, US), Santesson (S, UPS), Skottsberg (S). 

BRAZIL: Damazio (G), Deventer (G), Eiten (US), Glaziou (G, 

P), Hohnel (W), Irwin (NY, US), Kurokawa (TNS; specimens not available for study), Malme (S), Puiggari (G), 

Reitz and Klein (US), Schiffner (W, WU), Spruce (BM, H, 

NY), Ule (G), Vainio (TUR [original], BM, FH, M, P, 

UPS), Weddel (P, H), Warming (M). 

PARAGUAY: Balansa (G), Malme (S). 

URUGUAY: Felippone (W), Herter (H), Hosseus (H), Osorio 

ARGENTINA: Arechavaleta (G), Balansa (G), Dusen (S), Lamb 

(MVM). 

(CAN), Lorentz and Hieronymus (M). 

Noticeable gaps remain in Bolivia, Ecuador, Peru, 

Venezuela outside the Andes, and areas of Brazil 

bordering these countries. The unevenness of collecting can also be appreciated by examining lists 

of species collected in each country (see p. 18). 

I have not attempted to evaluate literature reports of species in floristic lists (exclusive of type 

species) other than to examine the herbarium 

collections on which these were based and annotate 

them with revised names as needed. The sheer 

number of collections involved discourages one 

from tracing, verifying, and correcting each published name. Because of the extensive synonymy, 

lack of adequate chemical testing, and the generally low level of accuracy in identification, older 

published reports are quite untrustworthy. 

Morphological Characters 

Hypotrachyna is a segregate of the collective 

genus Parmelia characterized by narrow, apically 

truncate lobes, a black lower surface, and dichotomously branched rhizines (Hale, 1974a). Hale 

and Kurokawa (1964) had earlier discovered that 

presence or absence of marginal cilia, type of cilia 

(normal or bulbate), and branching pattern of 

rhizines were the most consistent characters that 

could be used to subdivide the more than 400 

species of Parmelia. The segregate Hypotrachyna is 

one of the more clear-cut groups in the Parmeliaceae 

because of the distinctive rhizine branching. 

INTERNAL SmucTuRE.-Little attention has been 

paid to differences in internal structure of Parmeliae 

and how they might aid in a more natural classification of the genera. Tlihile carefully sectioning 

the Brazilian collections, Lynge (1914) noted that 

the species related to Parmelia microsticta had a 

rather thick paraplectenchymatous cortex in con- 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

trast to a palisade plectenchymatous cortex found 

in Hypotrachyna species. RIy recent work with the 

scanning-electron microscope (SEM) showed that 

all species of Hypotrachyna so far examined (Hale, 

I97Za) have a palisade cortex that is overlain by a 

thin, pored polysaccliaride epicortex (Figure la) 

which cannot be seen with light microscopes. It is 

the cortex, along with dichotomously branched 

rhizines, that I use to distinguish Hypotrachyna. 

Parmelia sensu strict0 has a paraplectenchymatous 

cortex without a pored epicortex, as in the typespecies of the genus, P. saxatilis (L.) Acharius. 

The cross sections of Hypotrachyna are consequently rather uniform from species to species. 

The palisade cortical cells form a layer 12p-181i 

thick. The TreDoztxia-containing algal layer is 

12~-2411 thick, the medulla 50y-150p thick, and the 

paraplectenchymatous lower cortex 12p-18p thick 

(Figure 10). In general, very small species (2-5 cm 

in diameter) are quite thin (medulla 50p-60p), 

FIGURE 1.-Morphology of Hypotrachyna: a, surface of H. croceopustulata (Imshaug 22275) (X 

500 with SEM); b, cross section of H. croceopustulata (Hale 33269) (X 500 with SEM); c, rhizines 

of H. densirhizinata (Scholander) (X 100 with SEM); d, cross section of thallus and a soralium 

of H. densirhizinata (Hale 43194) (X 75 with SEM). 
NUMBER 25 5 

while larger species (5-25 cm in diameter) have 

proportionately thicker layers. The rhizines consist of conglutinated strands of prosoplectenchymatous hyphae and are moderately to richly dichotomously branched (Figure lc). 

LOBE CONFICURATION.-HYPO~YUC~~~U is extremely homogeneous in overall lobe configuration. It is 

possible, however, to recognize two broad types in 

tropical America: an extremely elongate, divaricate, 

and little-branched type as in H. caraccensis (Figure 

8a) or M. lopezii (Figure 13c), and a shorter-lobed, 

more or less closely branched and imbricate type 

exemplified by H. Zivida (Figure 13b). There are, 

of course, numerous intermediates between these 

two extremes, as in H. laeuigata (Figure 120. A 

third type of configuration is found in the very 

rare H. koyaensis, which has the broader, almost 

subirregular lobes (Figure 1%) characteristic of 

many species of Hypotrachyna endemic to Asia. 

SoREDIA.-These vegetative diaspores originate in 

FIGURE 2.-Vegetative structures of Hypotrachyna (all X 10): a, diffuse soralia of H. revoluta 

(Hale 43293a); b, orbicular soralia of H. sinuosa (Hale 43071a); c, pustulate soralia of H. 

croceopustulata (Hale 33269); d, pustules of H. dactylifera (Hale 42075). 
SXIITHSONIAN CONTRIBUTIONS TO BOTANY 

soralia (Figure Id) and erupt as a powder. The 

shape and location of the soralia on lobes are important taxonomic characters used to recognize 

species. Soredia occur in 18 of the 77 species of 

tropical American Hypotrachyna: H. breuirhiza, 

H. croceopzistulata, H. densirhizinata, H. endochlol-a, H. exsplendens, H. pavovirens, H. gondylophora, H. immaculata, H. laeuigata, H. oostingii, 

H. prodzicta, H. pseudosinuosa, H. revoluta (Figure 2a), H. rockii, H. sinuosa (Figure 2b), H. subaffinis, H. thpsanota, and H. velloziae. They are 

generally subterminal to rarely lamina1 and produced in orbicular soralia. Soralia rarely become 

diffuse, as in H. reuoluta. In a few species pustules 

are formed initially but later become sorediate, as 

in H. croceopustulata (Figure 2c). Details of abnormal soredial formation will be found in the 

species discussions. 

PusTuLEs.-These are coarse, inflated, extremely 

fragile, isidialike structures (Figures Zd, 3a), but 

they are related to soralia since they often become 

sorediate. True pustules are hollow and tend to 

FIGURE 3.-Vegetative structures of Hypotrachyna: a, pustules of H. formosana (Eiten 7671a) 

(X 60 with SEM); b, enlargement of pustule in a (X 300); c, cross section of thallus and isidia 

of H. imbricatula (Quiros 1479) (X 100 with SEM); d, enlargement of isidia in c (X 300). 
FIGURE 4.-Vegetative structures of Hypotrachyna: a, cross section of an isidium of H. costaricensis (Hale 42201) (X 1000 with SEICI); b, lohulate isidia of H. andensis (Hale 42658a) (X 100 

with SEM); c, lobulate isidia of H. bogotensis (Oberwinkler and Poelt 7544) (X 10); d, lobules 

of H. ~achista (Hale 43139) (X 10); e, lobules of H. chlorina (Hale 43104) (X 10); f, maculae 

on a lobe of H. subplumbeata (Moore and Wood 4944) (X 10). 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

break open apically without soredial formation. 

Only three species of Hypotrachyna treated here 

have this type of pustule: H. dactylifera, H. formosana, and H. malmei. Under SEM the surface of 

pustules is quite rough with loosely aggregated 

hyphae lacking an outer polysaccharide layer 

(Figure 3b). More work needs to be done on these 

structures. 

IsIDIA.-Fingerlike, cylindrical isidia, densely 

filled with medullary hyphae and about 0.1 mm in 

diameter, are well-known lichen structures (Figures 3c, 3d). They occur in 16 species of Hypotrachyna: H. andensis, H. bogotensis, H. consirnilis, H. costaricensis, H. dentella, H. ensifolia, 

H. imbricatula, H. isidiocera, H. koyaensis, H. 

microblasta, H. neodissecta, H. osorioi, H. partita, 

H. rhabdiformis, H. steyermarkii, and H. subphysodalica. 

Isidia are often presented in textbooks as containing extensions of thalloid tissues. When viewed 

with the scanning-electron microscope, however, we 

see little internal organization (Figure 4a). There 

is no true cortical structure and no pored epicortex, 

only a thin, smooth, continuous polysaccharide 

sheet covering the weakly organized mass of algae 

and hyphae inside. 

While most isidia show little variation except in 

branching, lobulate isidia sometimes are found to 

occur in six of the above-named species: H. andensis, H. bogotensis, H. ensifolia, H. imbricatula, H. 

partita, and H. steyermarkii. In such cases the 

isidia begin normally but soon become procumbent 

and even dorsiventral, as in true lobules (Figure 

4b,c). Cilia will rarely form (cf. H. steyermarkii). 

When viewed with the SEM, the lobulate isidia 

appear to have the same level of organization as 

isidia, lacking, for example, a distinct cortex. 

LoBuLEs.-Almost any species of Hypotrachyna 

may develop small adventitious lobes with age. 

Such lobes can be regarded as regenerative structures with internal structure and a pored epicortex 

comparable to the main lobes when viewed under 

the scanning-electron microscope. As defined here, 

lobules are a consistent species character, abundantly or distinctively produced and not associated 

with isidia. Four species have them: H. chlorina 

(Figure 4e), H. prolongata, H. rachista (Figure 

4d), and H. singularis. I do not mean to imply, 

however, that these lobules are identical in origin 

and formation. 

MAcuLAE.-These are mottled, light to dark 

patches in the cortex visible at X 10 magnification 

(Figure 4f). They are not detectable as surface 

features under the scanning-electron microscope 

but seem to result from darker clumps of algae 

contrasted with the lighter algae-free hyphal matrix. 

Although some species of Hypotrachyna are always 

maculate, this is a difficult character to determine 

in practice and I have avoided using maculae as a 

primary character. 

APOTHECIAL CHARACTERS.-Apothecia have been 

found in 63 species of Hypotrachyna but so far are 

unknown in H. exsplendens, H. gondylophora, H. 

lineariloba, H. monilifera, H. oostingii, H. partita, 

H. physodalica, H. producta, H. prolongata, H. 

rachista, H. steyermarkii, H. subphysodalica, H. 

thysanota, and H. velloziae. Apothecia are present, 

but no spores developed, in H. croceopustulata, H. 

laevigata, H. protoboliviana, and H. subplumbeata. 

Apothecia are uniformly small, 1-5 mm in diameter (rarely to 10 mm), adnate (stipitate only in 

H. peruviana), and imperforate. The hymenium is 

40~-60~ high. The unicellular spores show a very 

small range in size, usually 4p-lop wide and 8y-16p 

long. A few species, notably those with yellow pigments (as H. endochlora) and H. pluriformis have 

significantly larger spores, up to 25p long, but no 

species of Hypotrachyna has spores more than 30p 

long. 

Pycnidia are often produced by apothecia-bearing 

species. The microconidia are about 1p wide and 

5p-7~ long in all species investigated. I have not 

included them in the descriptions since they seem 

to have no taxonomic value. Articles by Nylander, 

Lynge, and Vainio can be consulted for further 

data on pycnidia and microconidia in various 

species. 

Chemical Characters 

Beginning with Nylander in the 1860s and 

continuing with Vainio, Zahlbruckner, and Lynge, 

color tests with KOH and Ca(OC1)2 have played an 

important role in identifying species of Parmelia, 

and information on the tests always has been included in the species descriptions. Only Miiller 

Argau refused to use chemistry, even though it was 

shown to be a useful aid in identification. 

It was Asahina in the few years from 1933-1940 

who perfected the microcrystal tests by which 
NUMBER 25 9 

specific lichen substances could be distinguished. 

This meant, for example, that a KOH+ red species 

could be shown to contain norstictic acid, salazinic 

acid, or other component. 

The next significant advance in chemical techniques was partition chromatography, first with 

paper (Wachtmeister, 1952; Mitsuno, 1953) and 

later on silica gel (thin-layer chromatography or 

TLC) (Stahl and Schorn, 1961). TLC is now the 

accepted standard for routine qualitative identification of lichen substances and there is little 

chance it will be replaced by a more sophisticated 

technique in the near future. High-performance 

liquid chromatography (C. Culberson, 1972b) is 

and will remain far too expensive to be widely 

available to taxonomists. 

My original chemical analyses of Parmelia specimens were accomplished with microcrystal tests 

well before 1965 when TLC was just coming into 

use. Many of these specimens were returned to 

herbaria from which they had been borrowed, and 

it has been necessary in the last few years to borrow 

them again (especially types) and perform TLC 

tests. Unfortunately, some which I annotated between 1958 and 1964 were not rechecked. All 

material examined since 1965 has been studied 

with TLC. 

Culberson’s methods have been used exclusively 

(C. Culberson, 1972a). I have also benefited from 

being able to consult with Dr. Culberson on numerous occasions on the identification of difficult or 

unknown acids. 

Following is an alphabetical list of all identifiable 

substances, the species that contain them, and 

associated substances, that is, those with which they 

may occur. Unknowns are given at the end. Further 

details on joint occurrences of substances, accessory 

substances, etc., are given in discussions of the 

species. 

Alectoronic acid.-H. degelii, H. densirhizinata, H. ensifolia, 

H. exsplendens, H. gigas, H. lineariloba, H. rnalrnei, and 

H. protenta. Associated substances: atranorin, a-collatolic 

acid, gyrophoric acid, lichexanthone, and protocetraric acid. 

An additional qualitative test for alectoronic acid is bright 

white to bluish white fluorescence in longwave ultraviolet. 

Anziaic acid.-H. ducalis, H. eitenii, H. partita, H. producta, 

and H. rachista. Associated substances: atranorin, lichexanthone, and perlatolic acid. Several unknown spots commonly occur in TLC profiles of anziaic acid. 

Atranorin.-In all species except H. brasiliana, H. caraccensis, H. enderythraea, H. erythrodes, H. flauida, H. pauovirens, H. formosana, H. lopezii, H. malmei, H. microblasta, 

H. minima, H. novella, H. osteoleuca, H. physodalica, H. 

protoboliviana, H. reducens, H. siluatica, H. subafinis, H. 

subphysodalica, and H. uelloziae. Associated substances: all 

substances except lichexanthone and usnic acid. Atranorin 

is the main cortical substance in Hypotrachyna; it is absent 

only when replaced by usnic acid or lichexanthone, although in a few cases it may be present in trace amounts. 

Barbatic acid.-H. chlorina, H. dentella, H. endochlora, H. 

imbricatula, H. isidiocera, H. laeuigata, H. monilifera, H. 

peruviana, H. physcioides, H. protoboliviana, H. steyermarkii, and X. subplurnbeata. Associated substances: Atranorin, 4-0-demethylbarbatic acid, obtusatic and norobtusatic 

acids (constant accessories except in H. subplurnbeata), 

echinocarpic acid, entothein, usnic acid, and unidentified 

yellow pigments. A complete discussion of the chemistry 

of these species is provided by Culberson and Hale (1973). 

Chromatography of the barbatic group is very difficult; the 

most promising resolution is obtained with toluene-acetic 

acid. All specimens usually react C+ orange, a reaction 

now known to be caused by 4-0-demethylbarbatic acid, not 

barbatic acid itself. 

Colensoinic acid.-H. dactylifera, H. erythrodes, H. formosana, H. immaculata, H. livida, H. novella, and H. palmarum. Associated substances: atranorin, 4-0-demethylphysodic acid, lichexanthone, lividic acid, norcolensoinic 

acid, physodic acid, rhodophyscin, the “palmarum” unknown, and traces of other unknowns. See the discussion 

under lividic acid. 

a-Collatolic acid.-H. degelii, H. densirhizinata, H.. exsplendens, H. gigas, and H. protenta. Associated substances: alectoronic acid, atranorin, and gyrophoric acid. a-Collatolic 

acid is not a constant accessory component with alectoronic 

acid, as one may see by comparing the lists of species. 

Constictic acid.-H. andensis, H. loperii, and H. sinuosa. Associated substances: norstictic acid, salazinic acid, stictic 

acid, and usnic acid. Constictic acid invariably occurs with 

stictic acid in Hypothrachyna. Its structure is very close to 

stictic acid. 

4-0-Demethylbarbatic acid.-See barbatic acid. 

4-0-Demethylphysodic acid.-See lividic acid. 

Echinocarpic acid.-H. dentella, H. subafinis, and H. thysanota. Associated substances: atranorin, barbatic acid complex, gyrophoric acid, lichexanthone, and microphyllinic 

acid, The structure of this P+ orange-red acid is as yet 

undetermined. It forms a low discrete H,SO,+ orange spot 

in hexane-ether and a low, dull gray streak in benzenedioxane solvent systems. 

Entothein.-H. chlorina, H. endochlora, H. isidiocera, and 

H. peruviana. Associated substances: atranorin, barbatic 

acid complex, obtusatic acid complex, and unidentified 

yellow pigments. 

Evernic acid.-H. bogotensis; H. chicitae, H, pulvinata, and 

H. rockii. Associated substances: atranorin, lecanoric acid, 

norobtusatic acid, obtusatic acid, and rhodophyscin. Lecanoric acid is a constant accessory. Culberson and Hale (1973) 

discuss this group more fully in relation to the species con- 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

taining barbatic acid. They hypothesize that obtusatic acid 

could be the product of hybridization between evernic acid 

and barbatic acid. 

Fumarprotocetraric acid.-H. gondylophora. Associated substances: atranorin, rhodophyscin, and succinprotocetraric 

acid. Fumarprotocetraric acid would seem to be an extremely rare substance in Hypotrachyna. 

Galbinic acid.-H. caraccensis, H. enderythraea, and H. 

microblasta. Associated substances: norstictic acid, salazinic 

acid, and usnic acid. Galbinic acid was first described from 

Parmelina galbina (Acharius) Hale where it is a primary 

constituent. It has also been reported in Usnea. In Hypotrachyna it always occurs with norstictic and salazinic acids. 

The molecular structure has recently been determined by 

Dr. Myles Keogh as an acetate of salazinic acid. 

Gyrophoric acid.-H. gigas, H. neodissecta, H. oostingii, H. 

osorioi, H. pluriformis, H. revoluta, and H. thysanota. Associated substances: alectoronic acid, atranorin, echinocarpic 

acid, microphyllinic acid, and unknowns. Gyrophoric acid 

is an accessory substance only in H. gigas. 

Lecanoric acid.-H. bogotensis, H. chicitae, H. pulvinata, and 

H. rockii. Associated substances: atranorin, evernic acid, 

norobtusatic acid, obtusatic acid, and rhodophyscin. Although common as a primary constituent (along with 

atranorin) in most parmelioid genera, lecanoric acid always 

occurs in Hypotrachyna with and in lesser concentration 

than evernic acid (C. Culberson, 1972b). The color test 

with calcium hypochlorite is therefore weak or indistinct. 

Lichexanthone.-H. brasiliana, H. eitenii, H. erythrodes, H. 

formosana, H. malmei, H. minima, H. novella, H. osteoleuca, H. silvatica, and H. subafinis. Associated substances: 

alectoronic acid, anziaic acid, atranorin, colensoinic acid, 

4-0-demethylphysodic acid, echinocarpic acid, lividic acid, 

norcolensoinic acid, olivetoric acid, physodic acid, protocetraric acid, rhodophyscin, and unknown pigments. Lichexanthone is a cortical substance which fluoresces brilliant 

yellow-orange under longwave ultraviolet. It completely replaces atranorin and usnic acid in the cortex, although 

traces of atranorin may accompany it in a few species. It is 

noteworthy that no Hypotrachyna species endemic to the 

Old World produces lichexanthone. 

Lividic acid.-H. dactylifera, H. erythrodes, H. formosana, 

H. irnmaculata, H. livida, and H. novella. Associated substances: atranorin, colensoinic acid, 4-0-demethylphysodic 

acid, lichexanthone, norcolensoinic acid, physodic acid, rhodophyscin, and unknowns. C. Culberson (1966) isolated lividic 

acid from H. livida but its molecular structure (apparently 

close to physodic acid) is still unknown. It can be identified 

with TLC, preferably using the hexane and toluene solvent 

systems. All of the associated substances fall out in this 

sequence from highest to lowest RF in hexane: atranorin 

or lichexanthone, colensoinic acid, norcolensoinic acid, 4-0- 

demethylphysodic acid, physodic acid, and lividic acid (see 

C. Culberson, 1966). Various unknowns may appear below 

and above lividic acid. In toluene the resolution is not as 

good and only four or five distinct spots may appear in 

this sequence: atranorin or lichexanthone, colensoinic acid, 

4-8-demethylphysodic acid, norcolensoinic acid, and lividic 

acid. Perfect resolution of spots is only rarely achieved with 

these solvent systems, and we must await more satisfactory 

separation that will come from high-performance liquid 

chromatography. 

Microphyllinic acid.-H. thysanota. Associated substances: 

atranorin, echinocarpic acid, and gyrophoric acid. The 

identity of microphyllinic acid was established by Mr. Jon 

Dey while comparing the type of Parmelia nakanishii with 

his P. oostingii. It is the first reported occurrence outside 

of Cetraria, although C. Culberson postulated its existence 

in Hypotrachyna (Culberson and Hale, 1973). 

Norcolensoinic acid.-See discussion under lividic acid. 

Norobtusatic acid.-All species containing obtusatic acid (see 

lists under barbatic acid). Associated substances: see discussions under barbatic acid and evernic acid. 

Norstictic acid.-H. andensis, H. brevirhiza, H. caraccensis, 

H. enderythraea, H. pavovirens, H. lopezii, H. microblasta, 

H. reducens, H. rhabdiformis, H. sinuosa, and H. sublaevigata. Associated substances: atranorin, constictic acid, galbinic acid, protocetraric acid, salazinic acid, stictic acid, and 

usnic acid. One of the first positive results of TLC was the 

resolution of norstictic acid and its closely related associated 

substances. Norstictic acid occurs in trace amounts with 

salazinic acid and in substantial amounts with stictic acid. 

The accessory substance connorstictic acid has been reported in Parmotrema (W. Culberson, 1973). Connorstictic 

acid is very close to salazinic acid and may occur in Hypotrachyna although I have not been able to identify it positively. 

Obtusatic acid.-All species containing barbatic acid, H. 

chicitae. Associated substances: see discussions under barbatic acid. 

Olivetoric acid.-H. intercalanda and H. osteoleuca. Associated substances: atranorin, lichexanthone, rhodophyscin, 

and unknowns. This CS red acid can be easily identified 

by means of TLC. 

Perlatolic acid.-Hi. partita. Associated substances: anziaic 

acid and atranorin. Perlatolic acid, rare for Hypotrachyna, 

was identified in only one specimen of H. partita along 

with closely related anziaic acid. Its co-occurrence with 

anziaic acid would not be unexpected. 

Physodalic acid.-H. physodalica and H. subphysodalica. Associated substances: protocetraric acid and usnic acid. This 

acid is biogenetically closely related to protocetraric acid 

and often occurs with it in Hypogymnia. However, only 

one of the 14 species of Hypotrachyna having protocetraric 

acid contains physodalic acid. 

Physodic acid.-See discussions under lividic acid. 

Protocetraric acid.-H. bahiana, H. brasiliana, H. consirnilis, 

H. contradicta, H. croceopustulata, H. flavida, H. Pavovirens, H. koyaensis, H. malmei, H. minima, H. obscurella, H. 

physodalica, H. pseudosinuosa, H. silvatica, and H. velloziae. Associated substances: alectoronic acid, atranorin, 

lichexanthone, norstictic acid, physodalic acid, rhodophyscin, salazinic acid, and usnic acid. This is the commonest medullary substance in Hypotrachyna, although 

none of the species containing it are especially widespread. 

The best separation (especially from fumarprotocetraric) 

is achieved in the hexane solvent system using 10-20 ml 

more ether than recommended by C. Culberson (1972a). 
NUMBER 25 11 

Rhodophyscin.-H. bahiana, H. consimilis, H. croceopustulata, H. dactylifera, H. erythrodes, H. gondylophora, H. 

immaculata, H. rockii, and H. velloziae. Associated substances: atranorin, colensoinic acid, evernic acid, fumarprotocetraric acid, lecanoric acid, lichexanthone, lividic 

acid complex, protocetraric acid, succinprotocetraric acid, 

and usnic acid. This red pigment is assumed to be an 

anthraquinone first isolated from Physcia. It occurs sporadically in the species listed, either in the lower medulla or 

under soralia. 

Salazinic acid.-H. andensis, H. boquetensis, H. brevirhiza, 

H. caraccensis, H. enderythraea, H. microblasta, H. reducens, H. sinuosa, H. sublaevigata, and H. subsaxatilis. 

Associated substances: atranorin, constictic acid, galbinic 

acid, norstictic acid, stictic acid, and usnic acid. I have not 

conclusively identified consalazinic acid, an unknown substance that may accompany salazinic acid. 

Stictic acid.-H. andensis, H. lopezii, and H. sinuosa. Associated substances: constictic acid, norstictic acid, usnic 

acid, and two unknowns. This acid is quite rare in Hypotrachyna and is accompanied by the same substances that 

are found in other parmelioid genera. 

Succinprotocetraric acid.-H. gondylophora. Associated substances: atranorin and fumarprotocetraric acid. This recently described acid (Baker et al., 1973) often accompanies 

fumarprotocetraric acid in the genus Xanthoparrnelia. 

Usnic acid.-H. andensis, H. caraccensis, H. enderythraea, H. 

flavida, H. flavovirens, H. lopezii, H. microblasta, H. physodalica, H. protoboliviana, H. reducens, H. sinuosa, H. subphysodalica, and H. velloxiae. Associated substances: atranorin, barbatic acid complex, constictic acid, galbinic acid, 

norstictic acid, obtusatic acid complex, physodalic acid, 

protocetraric acid, rhodophyscin, salacinic acid, and stictic 

acid. Usnic acid, a common cortical pigment, ordinarily 

replaces atranorin or lichexanthone, although traces of 

atranorin may be present in some species. It occurs only 

with P+ 8-orcinol depsidones with the sole exception of 

H. protoboliviana, which contains the P - barbatic acid 

complex. 

Fatty acids.-H. costaricensis, H. koyaensis, and H. singularis. 

Associated substances: atranorin, protocetraric acid. Fatty 

acids are represented by caperatic and protolichesterinic 

acids, perhaps others, all difficult to identify with TLC. 

“Gracilescens” unknown.-H. gracilescens. Associated substance: atranorin. 

“Oostingii” unknowns.-H. oostingii. Associated substances: 

atranorin and gyrophoric acid. These unknowns are white 

fluorescent in longwave ultraviolet but differ from alectoronic acid. 

“Palmarum” unknown.-H. palmarum. Associated substances: 

atranorin and colensoinic acid. This unknown acid forms 

a distinct spot below colensoinic acid different from any of 

the other colensoinic-lividic acid complex substances but 

obviously related to them. 

“Prolongata” unknown.-H. prolongata. Associated substance: 

atranorin. This C+ red compound could not be identified 

with any other lichen substances reacting C+. 

“Silvatica” anthraquinones-H. silvntica. Associated substances: atranorin, lichexanthone, and protecetraric acid. 

This rusty red pigment occurs throughout the lower medulla. 

Speciation in Hypotrachyna 

One of the least discussed topics in lichenology 

is species evolution. As Thomson (1961) points out, 

we have little of the kinds of cytological or fossil 

evidence that is so often available in other plant 

groups. An important clue in lichenology, however, 

has proven to be chemistry, and, as stated by 

Culberson and Culberson (1 970), comparative 

phytochemistry is the most useful independent 

check that exists to evaluate the naturalness of 

systems based on morphology. This is especially 

true in lichens, where differentiation of chemical 

races is a far more obvious and frequent means of 

evolutionary differentiation than morphology (W. 

Culberson, 1970). 

I will discuss here primarily the origin and 

direction of speciation in Hypotrachyna, although 

many of the observations and conclusions should 

apply to other foliose genera. Briefly, I will deal 

with three aspects: possible hybridization of parent 

species, formation of vegetative morphs, and habitat 

selection. 

HYBRIDIZATION 

Lichenologists probably would agree that crustose 

lichens reproduce sexually since most species lack 

any other obvious means of propagation. On the 

other hand, it is assumed that foliose and other 

macrolichens generally reproduce vegetatively by 

means of diaspores (soredia, isidia, etc.). Little has 

been said about the reproduction of those foliose 

lichens which lack vegetative diaspores. If two fertile lichens should hybridize, we would expect gene 

exchange and possible mutations, processes that 

could give rise to new variations and species. 

Unfortunately, we are not sure how lichens reproduce sexually. There is no way at this time to 

make experimental crossings and to study cytological details. It is assumed that microconidia (spermatia or pycnidiospores) are capable of migrating 

to the ascogonial coil, forming a dicaryon with the 

female element, and existing in a brief one-celled 

sporophytic stage before meiosis and spore formntion take place. Each of the eight ascospores can be 

germinated and will, at least in theory, reconstitute 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

a new thallus when a suitable lichenizable alga is 

entrapped. 

Several recent studies have indeed shown that 

variation in the chemistry of certain populations 

can best be explained on the assumption of sexual 

reproduction and hybridization. I can cite my own 

work in Cetraria ciliaris (Hale, 1963) and the Culbersons’ recent study of the Parmelia perforata group 

(Culberson and Culberson, 1973). The most convincing evidence for Hypotrachyna has been presented by Culberson and Hale (1973) for those 

species producing evernic acid or barbatic acid. 

Their data are summarized in Figure 5. Culberson 

showed that hybridization of plants in the New 

World with type I Old World chemistry (lM-l = 

evernic acid and 1-1 = lecanoric acid) and type 

11 chemistry (zM-2 = barbatic acid and 2-2 = 

4-0-demethylbarbatic acid) theoretically could give 

rise to a new population containing obtusatic acid 

(lM-2) and norobtusatic acid (1-2). Old World 

populations which had no opportunity to cross 

with progenitors containing evernic acid also now 

lack any species with obtusatic acid. On the other 

hand, these populations free of obtusatic acid are 

absent in the New World except for rare H. subplumbeata, leaving us to conclude that hybridization produced populations containing obtusatic 

acid and that such populations had a very significant advantage over those lacking this acid and 

have displaced them. 

Another example of possible hybridization is 

illustrated by species which previously would have 

been considered to contain “accessory” substances. 

One could reasonably postulate, for example, that 

gyrophoric acid occurring with alectoronic acid in 

some specimens of H. gigas was contributed by a 

now extinct progenitor which contained gyrophoric 

acid and which hybridized with the still extant 

parent containing alectoronic acid. The same reasoning could be applied to H. physodalica, which 

produces protocetraric acid with or without physodalic acid. 

A final example concerns the production of 

cortical substances, principally atranorin, lichexanthone, and usnic acid. While we are totally ignorant 

of the biogenesis of these substances, one might 

assume that atranorin is the normal component and 

that some mutation could block this substance in 

favor of lichexanthone (see Hale, 1972b, for H. 

formosana versus H. pustulifera) or usnic acid. I 

can cite the following species pairs: H. livida 

(atranorine) and H. novella (lichexanthone); H. 

HYPOTHETICAL PROGENITORS / PRIMARY 

SPECIES 

P massartit 

p mplurnbeata I ‘\ MU TAT IONS 

SECONDARY 

S PEC I ES’ 

P addita 

p adjuflcta 

P execta 

P orientalis 

P boaotensis 

tavlorensis 1- P rock’i 

H Y BRI Di 2 AT IONS 

P chicitae 

(2M-2),(2-2), (IM-2), (1-2) 

P dentelia 

p diqitata 

p irnbricatula 

P laevigata 

P virqinica 

FIGURE 5.-Proposed pathway of chemical and morphological evolution leading to species in the 

barbatic-evernic-acid complex (from Culberson and Hale, 1973). 
NUMBER 25 13 

contradicta (atranorin) and H. brasilianu (lichexanthone); and H. physcioides (atranorin) and 

H. protoboliviana (usnic acid). 

The genus Hypotrachyna in tropical America 

has 36 species that have apothecia and lack the 

usual vegetative diaspores. These include 26 species 

endemic to the New World. They represent a gene 

pool which has produced, and in many cases is 

still producing, hybridizations and mutations responsible for the creation of new species. As I will 

show below, many of the extinct sexual morphs in 

Hypotrachyna are survived by their vegetative 

morphs. 

In summary, the evidence now available supports 

a working hypothesis that sexual reproduction has 

occurred in Hypotrachyna and that it has contributed significantly to the morphological and, 

especially, chemical diversity of the genus. 

FORMATION OF VEGETATIVE MORPHS 

The primary vegetative diaspores in Hypotrachyna are soredia and isidia. These never occur 

together and seem to represent two separate lines 

of morphological evolution, as I had previously 

concluded in Parmotrema (Hale, 1965). Most 

lichenologists of the 19th century treated soredia 

and isidia as minor characters unworthy of species 

rank. It was, in fact, not until 1924 that Du Rietz 

presented a comprehensive discussion of these 

structures. He not only carefully described their 

orientation on the thallus but discussed their biological and systematic significance, concluding that 

sympatric sorediate-nonsorediate, isidiate-nonisidiate populations are best treated as varieties. He felt, 

however, that allopatric populations should be 

treated as distinct species. 

In the last decade we have gained a new perspective on the significance of soredia and isidia, 

even if we still do not know why they are produced. 

When studying Parmotrema on a world level, for 

example, I analyzed a number of “counterpart” 

species, that is, pairs of sorediate-nonsorediate and 

isidiate-nonisidiate species which shared otherwise 

identical morphology and chemistry (Hale, 1965). 

Fourteen of the 34 sorediate species had fertile nonsorediate counterparts. The sexual morphs were 

generally endemic to relatively small continental 

areas while the sorediate counterparts, as a rule, 

had much wider distributions. 

Poelt (1970) expanded these and other examples 

into a more comprehensive hypothesis of species 

pairs (“Artenpaare”). Briefly, he postulated that 

the asexual sorediate and isidiate morphs are derived from sexual morphs and that they behave 

apomictically without any means of further variation save highly unlikely somatic mutations. These 

secondary species, to quote Poelt, are blind alleys 

which have conserved the chemistry of the primary 

sexual morph. Later, Poelt (1972) expanded this 

hypothesis to include differentiation of a parent 

species into two or more chemically different, 

sexually reproducing species, each of which could 

produce stable secondary apomictic species. In 

addition, any or all of the primary species theoretically comprising species pairs may have become 

extinct. 

An upshot of this hypothesis is that morphologically identical but chemically variable sorediate 

(or isidiate) species, having probably evolved parallel from different parents, would have to be con 

sidered polyphyletic. Culberson and Culberson 

(1973) first suggested this possibility and used it to 

explain convincingly the origin of speciation in 

the Parmotrema perforatum group. A further consequence is that polyphyletic “chemical species” 

almost certainly are distinct species. 

If the species pair concept is in fact true, it goes 

far toward explaining the chemical diversity among 

vegetative morphs. On the other hand, it is supported by no proof other than that offered by 

comparative biochemistry. As such, it may be an 

oversimplification that might discourage lichenologists from exploring other evolutionary processes. Some sorediate and isidiate species, for example, 

do produce pycnidia and microconidia which could 

act as fertilizing elements of fertile populations. 

These morphs need not remain simply as “blind 

alleys.” Alternative hypotheses may come to light 

as we gain a better understanding of soredial and 

isidial formation, the role of microconidia, and the 

mechanics and frequency of gene recombination 

in lichens. 

I believe it is profitable, nonetheless, to analyze 

species pairs in Hypotrachyna and see how much 

they have contributed to the evolution of the genus. 

The method is very simple: using a list of diasporebearing species, a search is made among nondiaspore-bearing fertile species which might be considered perfect morphological and chemical parents. 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

These parents are defined as strictly as possible 

with the realization that the sexual morphs may 

still be evolving and that the pairs may differ because of environmental modifications induced by 

occupying different habitats. 

SOREDIATE MoRPHs.-There are eighteen sorediate 

species in Hypotrachyna and nine of these can be 

reasonably paired with presumptive parents (given 

in alphabetic order of the sorediate species): 

H. brevirhixa - H. sublaevigata 

H. densirhizinata - H. gigas 

H. exsplendens - H. degelii 

H. immaculata - H. livida 

H. laevigata - H. physcioides 

H. producta - H. ducalis 

H. revoluta - H. pluriformis 

H. rockii - H. pulvinata 

H. sinuosa - H. reducens 

The remaining nine species cannot be so closely 

matched. Their origin can be hypothesized in a few 

cases. Hypotrachyna flavovirens, for example, normally contains only protocetraric acid but one 

specimen contains norstictic acid as well. The 

norstictic acid probably was contributed by a parent 

similar to H. reducens that crossed with a now 

extinct progenitor containing protocetraric acid. 

To cite another example, it is perhaps significant 

that H. densirhizinata is represented by a single 

chemical population whereas the presumptive parent H. gigas has two populations, one with gyrophoric acid in addition to alectoronic acid and one 

with alectoronic acid only. We are left to conclude 

that the sorediate morph H. densirhizinata evolved 

before the parent species H. gigas acquired gyrophoric acid, probably by crossing with a now extinct progenitor related to H. thysanota. 

Let us next take a group of very similar species 

which, for all practical purposes, can be distinguished only by means of chemical tests: 

H. endochlora (barbatic acid and pigments) 

H. densirhizinata (alectoronic acid) 

H. gondylophora (fumarprotocetraric acid) 

H. laevigata (barbatic acid) 

H. oostingii (gyrophoric acid) 

H. producta (anziaic acid) 

H. rockii (evernic and lecanoric acids) 

H. thysanota (gyrophoric, microphyllinic, and echinocarpic 

acids) 

If we accept the species pairs concept to explain 

this degree of chemical speciation, then we can 

appreciate how complicated is the evolutionary 

pattern involved. Can we assume that these eight 

species originated from a single sexually reproducing ancestor because of the very similar lobe configuration and mode of forming soredia? Would 

it follow, then, that the present chemical diversity 

resulted when the ancestor hybridized with other 

sexual morphs to give eight chemical populations, 

each of which produced a vegetative morph? While 

four of the above species seem to have parents, 

four (H. endochlora, H. gondylophora, H. oostingi 

and H. thysanota) have no chemically comparable 

sexual morphs which, we must assume, are either 

extinct, to be represented now by four chemically 

stable vegetative morphs, or yet to be discovered in 

tropical America. If our reasoning is true, then 

the presumptive parents of the remaining sorediate 

species in the genus, H. croceopustulata, H. 

flavovirens, H. pseudosinuosa, H. subafinis, and 

H. velloziae, are probably extinct. 

ISIDIATE Mo~p~s.-Al though lichenologists are 

ignorant of the causes of isidial formation, they 

rarely ever lump isidiate and nonisidiate species in 

the same taxon. In Hypotrachyna we find that 

seven of the sixteen isidiate species have chemically 

identical nonisidiate presumptive parents: 

H. andensis - H. reducens 

H. bogotensis - H. pulvinata 

H. consimilis - H. bahiana 

H. ensifolia - H. gigas 

H. imbricatula - H. physcioides 

H. microblasta - H. enderythraea 

H. partita - H. ducalis 

Two of the remaining nine isidiate species, H. 

dentella and H. isidiocera, are closely related to 

the H. physcioides group but differ somewhat 

chemically. Hypotrachyna neodissecta may have 

evolved from H. pluriformis but has much smaller 

spores. All of these vegetative morphs are rather 

easily distinguished morphologically and no “chemical species” complexes comparable to the H. 

laevigata group discussed above have evolved. We 

might assume, then, that the progenitors of isidiate 

morphs have hybridized or mutated to a lesser 

degree than the parents of sorediate morphs or 

that the isidiate morphs, once created, have had a 

lower survival rate. 

PUSTULATE MORPHS.-TWO pustulate species have 

perfect nonpustulate parents: H. dactylifera-H. 

livida and H. formosana-H. novella. Hypotrachyna 

malmei has no nonpustulate counterpart. 
NUMBER 25 15 

OTHER MoRPHs.-None of the lobulate species, 

except perhaps H. rachista-H. ducalis, has a perfect nonlobulate parent. Hypotrachyna chlorina 

obviously is related to the H. physcioides complex 

because it produces barbatic acid; H. prolongata 

and H. singularis stand isolated. 

SERIES OF MORPHS.-In the following species a 

progenitor has apparently given rise to more than 

one kind of morph: 

H. ducalis-H. producta (sorediate)-H. partita (isidiate)-H. 

rachista (lobulate) 

H. livida-H. immaculata (sorediate)-H. dactylifera (pustulate) 

H. physcioides-H. laevigata (sorediate)-H. imbricatula (isidiH. pulvinata-H. rockii (sorediate)-H. bogotensis (isidiate) 

H. reducens-H. sinuosa (sorediate)-H. andensis (isidiate) 

This seems to imply that most sexual morphs produce either sorediate, isidiate, or pustulate vegetative morphs and only rarely produce two or more 

different kinds. 

In conclusion, we find that Hypotrachyna consists of 36 potential sexually reproducing parent or 

progenitor species. Twelve of these have actually 

produced 18 morphs, giving a total of 54 species 

(out of the 77 recognized in this monograph) whose 

possible evolutionary development can be traced. 

The remaining 23 asexual species can be assigned 

to various chemical constellations, but a few simply 

cannot be explained at this time. 

It is my contention that the high rate of development of asexual morphs-involving 66 percent of 

the sexual morphs-and the coexistence of many of 

these presumptive progenitors are signs of comparatively recent origin for the genus. In other 

words, Hypotrachyna has been speciating rapidly 

since the continents drifted apart. 

ate) 

HABITAT SELECTION 

The concept of habitat selection as first proposed 

by Culberson 2nd Culberson (1967) helped to 

explain the microdistribution of chemical populations of Ramalina siliquosa. Three species, sympatric over their broad geographic range, segregated 

at different exposures on rocks into remarkably pure 

populations, an example of how genetically defined 

physiological races revealed phenotypically by 

chemistry (rather than morphology) were associated with distinct habitats. 

Given sufficient time 2nd resources, I believe 

similar kinds of habitat selection could be demonstrated among Hypotrachyna species. For example, 

species containing barbatic acid, especially H. 

physcioides, seem indiscriminate in selecting habitats, rock, trees, or soil. The tree population seems 

less robust and has more distinct maculae than the 

saxicolous population. Two distinct species may 

eventually evolve because of habitat segregation. 

Other examples could be cited, as the possible distant relationship between H. bahiana (corticolous) 

and H. contradicta (saxicolous) or between H. 

degelii (corticolous) and H. protenta (saxicolous). 

Classification of Hypotrachyna 

Vainio proposed three sections in his Brazilian 

study: Amphigymnia, Hypotrachyna, and Xanthoparmelia. Hypotrachyna was subdivided into three 

groups without formal rank: Irregularis, Cyclocheila, and Linearis. It is this last group by which 

I typified subgenus Hypotrachyna (Vainio) Hale 

and Kurokawa with Parmelia brasiliana Nylander 

as the lectotype species (Hale and Kurokawa, 

1964). I recently elevated this taxon to generic level 

(Hale, 1974a). 

Vainio assigned eleven species to Hypotrachyna, 

and, in fact, only one that he listed, Parmelia 

coronata Fee, does not belong here. Two yellow 

species, P. /lava and P. velloziae, were placed in 

section Xanthoparmelia but they have dichotomously branched rhizines and belong in 

Hypotrachy nu. 

While Hypotrachyna contains about 110 species, 

there is no practical internal subdivision into subgenera or sections. The most useful approach is to 

combine the species into chemical constellations 

proposed by Culberson and Hale (1973), which 

are based on the probably biogenetic relationship 

of the chemicals in Hypotrachyna (Figure 6). 

It is assumed that simple depsides, for example, 

represent the lowest evolutionary level and mixed 

depsidones the highest. Accordingly, in my opinion, 

species in the lividic acid complex (including the 

boxes for colensoinic acid and 4-O-methylphysodic 

acid) should and do represent the highest level of 

evolution in the genus. Without question, the 

barbatic-evernic acid complex has the highest level 

of speciation. Another advanced group, with 

alectoronic acid (H. gigas, H. degelii, etc.), seems 

to have evolved in parallel along different lines 
16 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

SIMPLE I) 

DEPSIDONES alec toroni c norcolenso'nic 

FIGURE 6.-Biogenetic relationships among secondary products in Hypotrachyna. Compounds 

connected by lines have similar chemical structures but are not necessarily derivable from the 

other. 

and has a lower level of speciation. As one might 

expect, the P+ acid series represented by H. 

brasilianu, H. enderythraea, H. reducens, H. subZaevigata, etc., seems to include more conservative 

species, some pantropical. 

The various species of Hypotrachyna are grouped 

provisionally (below) according to the biogenetic 

chart (Figure 6) and a consideration of morphological traits. The groups joined by dashes have a 

generally higher level of internal homogeneity than 

those separated by commas. 

Fatty acids.-H. costaricensis and H. singularis. 

PD+ compounds.-Norstictic acid derivatives: H. andensisH. reducens-H. rhabdiformis-H. sinuosa, H. boquetensisH. brevirhiza-Hi. sublaevigata, H. caraccensis-H. enderythraea-H. microblasta, H. lopezii, H. subsaxatilis. Protocetraric acid derivatives: H. brasiliana-H. bahiana, H. consirnilis-H. contradicta-H. croceopustulata-H. pseudosinuosaH. silvatica, H. flavida, H. flavovirens, H. gondylophora, H. 

koyaensis, H. malmei-H. minima, H. obscurella, H. physodalica, H. subafinis-H. velloziae, H. subphysodalica. 

Evernic-1ecanoric.-H. bogotensis-H. chicitae-H. pulvinataH. rockii. 

Gyrophoric: H. neodissecta-H. osorioi-H. pluriformis-H. revoluta. 

Barbatic: H. subplumbeata. 

Obtusatic: H. chlorina-H. dentella-H. endochlora-H. imbricatula-H. isidiocera-H. laevigata-H. monolifera-Hi. peruviana-H. physcioides-H. steyermarkii, H. protoboliviana. 

Microphyllinic: H. thysanota. 

Alectoronic: H. degelii-H. exsplendens-H. protenta, H. densirhizinata-H. ensifolia-H. gigas-H. lineariloba, (?)H. 

oostingii. 

Perlatolic-anziaic: H. ducalis-H. partita-H. producta-H. prolongata-H. rachista, H. eitenii. 

Olivetoric: H. intercalanda-H. osteoleuca. 

Colensoinic: H. dactylifera-H. erythrodes-H. formosana-H. 

gracilescens-H. immaculata-H. livida-H. novella-H. palmarum. 

More detailed discussions of some of these groups 

will be found in the section on species treatment. 

Habitats and Ecology 

Hypotrachyna is primarily a genus of higher 

elevations throughout its range in the tropics. It 

grows near sea level only at its northern limits in 

southern United States and southern limits in Chile 

and Argentina. It is absent or very rare in xerophytic areas and in lowland (0-500 m elevation) 

rain forest. 

The optimal area for Hypotrachyna is the zone 

between 1300 and 2400 m elevation in the major 

mountain chains of tropical America. The climate 

here is quite uniform throughout the year and precipitation comes in the form of rain and frequent 

clouds or mists. There are about 55 species here. 

Most are corticolous. Where rock outcrops are 
NUMBER 25 

Phytogeographical region 

17 

Endemic species Extraneotropical species 

Total flora No. Percent No. Percent 

abundant, however, a number of obligately saxicolous species may occur: H. brasiliana, H. contradicta, H. erythrodes, H. flavida, H. obscurella, 

H. osteoleuca, H. protenta, H. subsaxatilis, etc. 

Even some typically corticolous species may grow 

on rocks: H. formosana, H. imbricatula, H. microblasta, H. physcioides, H. revoluta, etc. 

The area above 2400 m in the American tropics 

becomes more sparsely forested and intergrades 

with the true treeless paramo. The Hypotrachyna 

flora here consists of 36 species, including many 

from the lower zone. 

The high paramo has much greater climatic 

extremes than other zones. The 20 species occurring here (3600-4500 m elevation) include some of 

the most showy and spectacular ones: H. caraccensis and H. gigas. Hypotrachyna chicitae, H. 

physodalica, and H. subplumbeata are the only 

species apparently restricted to high paramo. These 

paramo lichens grow on rocks, over grass and moss 

tussocks, and at the base of small shrubs, forming 

a conspicuous part of the vegetation. 

The zone below 1000-1200 m in the tropics is 

largely true rain forest and, relatively speaking, is 

poor in terms of number and diversity of foliose 

species. Only 14 species of Hypotrachyna have been 

recorded below 1000 m. These include those from 

temperate and subtropical latitudes in the United 

States and southern South America as well as 

“weedy” species with broad elevational amplitude 

such as H. costaricensis, H. formosana, H. laevigata, 

H. microblasta, H. revoluta, and H. rockii. 

North America .............................. 

West Indies ...................................... 

Andes ................................................ 

Mexico/Central America .............. 

Brazil/Uruguay .............................. 

Chile/Argentina ........................... 

Phytogeographical Relationships 

Hypotrachyna, primarily a tropical genus, is 

concentrated in the New World, There are 58 

19 1 5 6 32 

29 2 I 12 41 

40 4 10 14 35 

46 9 20 11 24 

38 11 29 8 21 

15 2 13 7 47 

species that occur only in the Americas, while Asia 

and Africa together have only 32 endemics. 

The American tropics, however, differ considerably in floristic composition and degree of endemism. Six major regions can be recognized, as shown 

in Table 1, where comparisons are made with 

pantropical elements and Europe and Africa. Europe has only one endemic, H. taylorensis, and 

four other species, H. endochlora, H. laevigata, 

H. revoluta, and H. sinuosa, all pantropical. 

Two regions of high endemism stand out: the 

Andean region (Colombia, Venezuela, Peru, Ecuador, and Bolivia) and the area from Minas Gerais 

and Mato Grosso in Brazil to adjacent Uruguay. 

The farther one goes from these centers the lower 

the degree of endemism and speciation. Actually, 

the Andes and southeastern Brazil have many 

species in common and this number surely will be 

increased as more collecting is done. 

Nineteen of the 77 species of Hypotrachyna 

occur outside the New World: H. bogotensis and 

H. laevigata are known from Europe: H. brevirhiza, H. costaricensis, H. endochlora, H. formosana, 

H. koyaensis, H. microblasta, H. pseudosinuosa, 

H. revoluta, H. rockii, and H. sinuosa are pantropical or occur on at least one Old World land 

mass; and the most interesting group, comprising 

H. dactylifera, H. degelii, H. densirhizinata, H. 

exsplendens, H. immaculata, H. livida, and H. 

subplumbeata, occurs outside of the New World 

only in southern Africa, a classic pattern for plants 

which evolved before the continents drifted apart. 

In the latter group there is a preponderance of 

species with the lividic acid complex. 

TABLE 1 .-Total Hypotrachyna flora, number and percentage of endemic species, 

and number and percentage of species also occurring outside the neotropics, 

according to the major phytogeographical regions. 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

The Hypotrachyna Flora 

The flora as presented at this time comprises 77 

species. The most commonly collected ones (with 

number of collections examined) are H. imbricatula 

(81 +), H. costaricensis (65+), H. bogotensis 

(60+), H. microblasta (52+), H. rockii (50+), 

H. physcioides (35+), H. laevigata (35), H. 

sinuosa (33), H. caraccensis (32), H. gigas (31), 

H. pulvinata (31), H. densirhizinata (25), and 

H. livida (25). It is noteworthy that the four commonest species are isidiate. 

The rarities in the Hypotrachyna flora, found 

only once or twice, include H. contradicta, H. 

ducalis, H. eitenii, H. malmei, H. minima, H. 

monilifera, H. neodissecta, H. prolongata, H. 

protoboliviana, H. singularis, H. subsaxatilis, and 

H. subplumbeata. 

The number of species in each country or 

political entity in tropical America varies from 

highs of 42 species in Venezuela and 37 in Brazil 

to only one species in Honduras, Puerto Rico, and 

Trinidad. These differences obviously reflect the 

intensity of collecting and not necessarily a poor 

flora. The following listing of the Hypotrachyna 

flora of each country shows how much work remains 

to be done, and hopefully it will stimulate further 

collecting efforts: 

ARGENTINA: H. brevirhiza, H. densirhizinata, H. gigas, H. 

intercalanda, H. livida, and H. pluriformis. 

BOLIVIA: H. caraccensis, H. costaricensis, H. densirhizinata, H. 

enderythraea, H. gigas, H. gondylophora, H. physcioides, 

H. pulvinata, H. revoluta, and H. sinuosa. 

BRAZIL: H. bahiana, H. brasiliana, H. caraccensis, H. chlorina, 

H. consimilis, H. contradicta, H. costaricensis, H. dactylifera, H. degelii, H. dentella, H. eitenii, H. enderythraea, 

H. endochlora, H. erythrodes, H. pavida, H. formosana, H. 

gracilescens, H. imbricatula, H. immaculata, H. intercalanda, H. isidiocera, H. laevigata, H. livida, H. malmei, 

H. microblasta, H. minima, H. novella, H. obscurella, H. 

palmarum, H. peruviana, H. pluriformis, H. protenta, H. 

pulvinata, H. rockii, H. silvatica, H. subafinis, and H. 

velloziae. 

CHILE: H. bogotensis, H. brevirhiza, H. dactylifera, H. densirhizinata, H. pavovirens, H. formosana, H. imbricatula, H. 

laevigata, H. oostingii, H. pseudosinuosa, H. revoluta, H. 

rockii, H. sinuosa, H. sublaevigata, and H. subphysodalica. 

COLOMBIA: H. andensis, H. bogotensis, H. caraccensis, H. 

costaricensis, H. degelii, H. densirhizinata, H. endochlora, 

H. ensifolia, H. Pavida, H. gigas, H. imbricatula, H. isidiocera, H. laevigata, H. microblasta, H. osteoleuca, H. physcioides, H. physodalira, H. producta, H. protenta, H. 

pulvinata, H. rachista, H. reducens, H. rockii, and H. 

sinuosa. 

COSTA RICA: H. andensis, H. bogotensis, H. caraccensis, H. 

chicitae, H. costaricensis, H. dactylifera, H. densirhizinata, 

H. endochlora, H. ensifolia, H. exsplendens, H. gondylophora, H. imbricatula, H. laevigata, H. microblasta, H. 

partita, H. physcioides, H. protenta, H. protoboliviana, H. 

pulvinata, H. rockii, and H. sinuosa. 

CUBA: H. costaricensis, H. formosana, H. imbricatula, H. 

microblasta, and H. rockii. 

DOMINICAN REPUBLIC: H. bogotensis, H. consimilis, H. costaricensis, H. densirhizinata, H. endochlora, H. ensifolia, 

H. gondylophora, H. imbricatula, H. isidiocera, H. laevigata, H. lineariloba, H. microblasta, H. physcioides, H. 

pseudosinuosa, H. rachista, H. revoluta, H. rockii, H. sinuosa, H. subafinis, and H. velloziae. 

ECUADOR: H. bogotensis, H. brevirhiza, H. caraccensis, H. 

densirhizinata, H. enderythraea, H. gigas, H. microblasta, 

H. pulvinata, and H. sinuosa. 

GUATEMALA: H. caraccensis, H. costaricensis, H. densirhizinata, 

H. exsplendens, H. formosana, H. imbricatula, H. isidiocera, 

H. microblasta, H. physcioides, H. pulvinata, H. rockii, and 

H. subafinis. 

GUYANA: H. imbricatula and H. laevigata. 

HAITI: H. bahiana, H. bogotensis, H. chlorina, H. costaricensis, H. croceopustulata, H. densirhizinata, H. ensifolia, 

H. formosana, H. imbricatula, H. microblasta, H. physcioides, H. prolongata, H. pseudosinuosa, H. rockii, H. 

sinuosa, H. subafinis, and H. velloziae. 

HONDURAS: H.imbricatula. 

JAMAICA: H. bogotensis, H. costaricensis, H. enderythaea, H. 

ensifolia, H. exsplendens, H. imbricatula, H. isidiocera, H. 

laevigata, H. microblasta, H. pseudosinuosa, H. pulvinata, 

and H. rockii. 

LESSER ANTILLES (Dominica, Grenada, Guadeloupe, Martinique, and St. Vincent): H. costaricensis, H. imbricatula, 

H. laevigata, H. microblasta, H. pseudosinuosa, and H. 

rockii. 

MEXICO: H. bogotensis, H. consimilis, H. costaricensis, H. 

croceopustulata, H. dactylifera, H. densirhizinata, H. dentella, H. endochlora, H. ensifolia, H. exsplendens, H. formosana, H. gigas, H. gondylophora, H. imbricatula, H. 

isidiocera, H. koyaensis, H. laevigata, H. microblasta, H. 

physcioides, H, pluriformis, H. pseudosinuosa, H. pulvinata, 

H. rachista, H. rockii, H. sinuosa, H. subafinis, H. sublaevigata, H. subplumbeata, H. subaxatilis, and H. thysanota. 

PANAMA: H. boquetensis, H. caraccensis, H. chlorina, H. COStaricensis, H. dactylifera, H. densirhizinata, H. endochlora, 

H. ensifolia, H. formosana, H. gigas, H. imbricatula, H. 

isidiocera, H. microblasta, H. partita, H. physcioides, H. 

protenta, H. pulvinata, H. rachista, H. reducens, H. rhabdiformis, H. rockii, and H. sublaevigata. 

PERU: H, andensis, H. bogotensis, H. caraccensis, H. chicitae, 

H. chlorina, H. costaricensis, H. degelii, H. densirhizinata, 

H. ducalis, H. enderythraea, H. erythrodes, H. pavida, H. 

formosana, H. gigas, H. imbricatula, H. microblasta, H. 

peruviana, H. physodalica, H. protenta, H. pulvinata, H. 

rachista, H. reducens, H. rhabdiformis, H. rockii, and H. 

singularis. 

PUERTO RICO: H. consirnilis. 
NUMBER 25 19 

TRINIDAD: H. imbricatula. 

URUGUAY: H. dactylifera, H. endochlora, H. livida, H. osorioi, 

and H. phiformis. 

VENEZUELA: H. andensis, H. bogotensis, H. brevirhiza, H. 

caraccensis, H. chicitae, H. chlorina, H. consimilis, H. costaricensis, H. dactylifera, H. densirhizinata, H. dentella, H. 

ducalis, H. enderythraea, H. endochlora, H. ensifolia, H. 

erythrodes, H. pavida, H. formosana, H. gigas, H. gondylophora, H. imbricatula, H. immaculata, H. isidiocera, H. 

laevigata, H. livida, H. loperii, H. microblasta, H. monilifera, H. neodissecta, H. novella, H. obscurella, H. osteoleuca, H. partita, H. physcioides, H. physodalica, H. protenta, H. pulvinata, H. rachista, H. revoluta, H. rockii, H. 

sinuosa, and H. steyermarkii. 

Nomenclature 

One of the most time-consuming tasks in any 

monographic effort is unraveling problems in 

nomenclature and typification. The literature must 

be searched as completely as possible, the identity 

and location of type specimens must be determined, 

and, where needed, lectotypes must be selected. I 

have decided to use a narrow interpretation of the 

Code of Botanical Nomenclature in defining types. 

A holotype, as I employ the term here, is the 

specimen actually designated as such by an author. 

This includes all species described since 1958 when 

the designation “holotype” became mandatory for 

a single element on which a new taxon is based. 

Before 1958 Gyelnik seems to be the only lichenologist who consistently used this term in a correct, 

modern sense. All other authors either failed to 

cite a holotype collection or left to later workers 

the job of deciding which of two or more specimens 

cited in a species description is the lectotype. 

A lectotype, then, in my definition, includes two 

cases, one in which one of the published syntypes 

is selected as the lectotype and another where the 

author listed only one collection in his description 

but failed to call it a holotype and often did not 

state clearly where the specimen was housed. In 

other words, I am often selecting one specimen out 

of several duplicate collections as the lectotype. 

The point of departure here is that I had formerly 

applied the term “holotype” to a collection which 

represented the only material cited by an author. 

Because of the element of selection involved, even 

as to location of the specimen, I feel the use of the 

term “lectotype” is justified. 

It should be emphasized that selection of lectotypes is not binding on later workers. The Code 

allows us to lectotypify a species again if better 

evidence can be produced to support the change. 

This may happen, for example, when a second 

worker discovers a specimen in another collection 

that is in better condition or is more appropriate 

than the previously designated lectotype. A possibly undesirable amount of nomenclatorial instability may be introduced in these circumstances, 

but this seems a better course than rigidly following a lectotypification which the earlier worker 

himself admits may be faulty. 

Keys to the Species 

The following keys are divided into three 

major groups: sorediate, isidiate, and nonsorediatenonisidiate species. The main division under each 

group is presence or absence of usnic acid (cortex 

greenish yellow or whitish to ashy gray). Following 

this, a combination of pigment and morphological 

characters is used, but ultimately chemical tests are 

called for. The standard color tests with KOH, 

calcium hypochlorite (or other bleach), and pphenylenediamine suffice to identify a number of 

species and species groups, but below this level it 

is highly desirable, if not sometimes mandatory, to 

identify the substances with microchemical tests, 

preferably thin-layer chromatography. Further clues 

as to the species identity can be gained by comparing the specimens with the photographs and by 

consulting the descriptions and geographical distribution. The geographic ranges of many species will 

undoubtedly be extended as more collections are 

made, but geography is still a valuable aid. 

Key to Species with Thallus Sorediate or Sorediate-Pustulate 

1. Thallus greenish yellow (usnic acid present). 

2. Lobes very narrow, less than 1 mm wide; medulla K- ........................... 77. H. velloxiae 

2. Lobes 1-3 mm wide; medulla Kf red or K-. 

3. Medulla KS yellow turning red ..................................................................... 69. H. sinuosa 

3. Medulla K- (protocetraric acid) ................................................................... 26. H. fluvovirens 
20 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

1. Thallus whitish to ashy gray (usnic acid lacking). 

4. Medulla uniformly yellow .................................................................................... 21. H. endochlora 

4. Medulla white. 

5. Soralia distinct, generally terminal or subterminal, soredia powdery. 

6. Medulla K+ red (salazinic acid) ................................................................. 6. H. brmirhixa 

6. Medulla K- or slowly dull yellowish. 

7. Medulla Cf rose, red, or orange-red. 

8. Soralia orbicular, distinct ............................................................ H. lmigata group 

(Barbatic acid: 36. H. Zaevigata; evernic-lecanoric acids: 66. H. rockii; gyrophoric acid: 47. H. oostingii; anziaic acid: 56. H. producta; gyrophoricechinocarpic acids: 76. H. thysanota.) 

8. Soralia more irregular and diffuse (Figure 2a); gyrophoric acid present .......... 

.................................................................................................. 

9. Medulla P+ red. 

10. Lobes narrow, 1-2 mm wide; thallus closely adnate ...................................... 

10. 

11. Thallus adnate; lobes short ...................................... 

11. Thallus loosely attached; lobes longdivaricate .. 

...................... 60. H. pseudosinuosa 

Lobes 2-5 mm wide; thallus loosely attached ............. 29. H. gondylophora 

9. Medulla P-. 

5. Soralia laminal toward the center of the thallus. 

12. Soralia with powdery soredia. 

13. 

13. Lobes 1-5 mm wide; thallus loosely adnate. 

14. 

14. Medulla C-. 

Lobes narrow, about 1 mm wide; thallus closely adnate .......... 71. H. subaffinis 

Medulla C+ red (gyrophoric acid) ....................................... 64. H. revolwta 

15. Medulla P 4- red ........ 13. H. croceopustulata 

.............. 32. H. immaculata 

................................ 

12. Soralia pustulate with sparse coarse soredia or pustules present. 

16. Lichexanthone present (thallus yellow in UV light). 

17. Lobes narrow, 1-2 mm; confined to Brazil ............................ 40. H. malmei 

17. Lobes 1-5 mm wide; pantropical ........................................ 27. H. formosana 

16. Lichexanthone absent (no fluorescense in UV). 

18. Medulla Cf rose (gyrophoric acid); isidiate pustules solid ... 48. H. osorioi 

18. Medulla C-. 

Pustules becoming sorediate (Figure 2c) ............ 13. H. croceopustulata 

Pustules not sorediate (Figure 2d) .............................. 14. H. dactylifera 

19. 

19. 

Key to Species with Thallus Isidiate, Isidiate-Lobulate, or Pustulate 

1. Thallus greenish esent). 

2. Medulla K- ............................................................ 73. H. subphysodalica 

3. Isidia remaining cylindrical .......................................................................... 41, H. microblasta 

3. Isidia distinctly lobulate .................................................................................... 

1. Thallus whitish to ashy gray. 

4. Medulla uniformly yellow ........................................................................................ 34. K. isidiocera 

4. Medulla white. 

5. Isidia mostly laminal on the lobe surface. 

6. Isidia inflated and hollow (pustules). See dichotomy 17 in the sorediate key above. 

6. Isidia cylindrical, solid, not strongly inflated. 

7. Medulla KS yellow turning red ..................... ...................... 65. H. rhabdiformis 

7. Medulla K- or slowly pale yellowish. 

8. Medulla C+ (and KC+) orange (barbatic acid group). 

9. Lobes narrow and appressed, 1-2 mm wide; isidia ciliate. ........................ 

............................................ ..................................................... 70. H. steyermarkii 

9. Lobes 2-5 mm wide, isidia eciliate. 

10. Medulla P- ........................................................................ 31. H. imbricatula 
NUMBER 25 21 

10. Medulla P f red ..................................................................... 17. H. dentella 

8. Medulla CS rose or red or C-. 

11. Medulla C+ rose or red (anziaic or gyrophoric acids). 

12. Isidia not cylindrical. 

12. Isidia cylindrical, erect .................................................. .44. H. neodissecta 

13. Isidia becoming lobulate, procumbent ........................ 51, H. partita 

13. Isidia club-shaped, crumbling apically ....................... 48. H. osorioi 

14. Isidia hollow (pustules) .................................................. 14. H. dactylifera 

Lobes sublinear, 1-3 mm wide ........................ 10. H. consirnilis 

16. Lobes subirregular, 3-7 mm wide .................... 35. H. koyaensis 

17. Isidia cylindrical, erect; fatty acids present ............................ 

11. Medulla C- or faint C+ rose (lecanoric acid). 

14. Isidia solid, not pustulate. 

15. Medulla Pf red (protocetraric acid). 

16. 

15. Medulla P-. 

............................................. .., , , , . . , , .. , .. , ..12. H. costdcen& 

18. Alectoronic acid pres (strongly white fluorescent 

......................... 22. H. ensifolia 

18. Evernic and lecanoric present (weak to negative 

17. Isidia becoming lobulate, procumbent. 

under UV) ..................... 

fluorescence) .................................................... 3. H. bogotensis 

19. Anziaic acid present ........................................................................................ 6 2. H. rachista 

19. Unknown C+ substance present: known only from Haiti ........... ..57. H. POlMgata 

Key to Species with Thallus Lacking Soredia, Isidia, and Pustules 

5. Isidia marginal toward lobe tips (Figure 44. 

1. Thallus greenish yellow (usnic acid present). 

2. Lobes long and little branched, divaricate; thallus loosely attached. 

3. Medulla K- .......................................................................................... ..54. H. physodalica 

3. Medulla K+ yellow turning red. 

4. Stictic acid present .............................................................................................. 39. H. lopezii 

4. Salazinic acid present ...................................................................................... 7. H. cmaccensis 

2. Lobes shorter, moderately branched; thallus generally adnate. 

5. Medulla K+ yellow turning red. 

6. Thallus corticolous ..................... ................................................. 63. H. reducens 

6. Thallus saxicolous ...................... .......................................... 20. H. enderythraea 

5. Medulla K- or KS very pale yellow. 

7. Thallus corticolous; medulla C + orange ............................... 59. H. protobolivima 

7. Thallus saxicolous; medulla C- ............................................................. 25. H. flavida 

1. Thallus whitish to ashy gray. 

8. All or most of the medulla pigmented yellow or red. 

9. Thallus saxicolous. 

10. Medulla C + deep red (olivetoric acid) ................................................ 49. H. osteoleuca 

11. Margins of lobes densely lobulate (Figure 4e) .......................................... 9. H. chlorina 

12. Pigment pale yellow, throughout the medulla ............................ 52. H. peruviana 

12. Pigment red, occurring in the lower half of the medulla .......... 67. H. silvatica 

14. Medulla C+ orange (barbatic acid group) ................................... 43. H. monilifera 

15. Medulla C+ deep red (anziaic acid) ............................................. 18. H. ducalis 

Thallus small, 3-6 cm broad ................................................ 37. H. lineariloba 

10. Medulla C- .......... ............................................................. ......... 23. H. erythrodes 

9. Thallus corticolous. 

11. Margins of lobes sparcely lobulate or lacking lobules. 

8. Medulla white throughout. 

13. Lobes long and little branched; thallus loosely attached. 

14. Medulla Cf rose or red or C-. 

15. Medulla C+ rose or C- (alectoronic acid). 

16. 

16. Thallus larger, more than 8 cm broad ......................................... 28. H. gigas 

13. Lobes shorter and moderately branched; thallus adnate. 
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

17. Thallus collected on rocks. 

18. Medulla K+ yellow turning red (salazinic acid) .................... 75. H. subsaxatilis 

18. Medulla K- or K+ pale yellow. 

19. Medulla C+ orange or red. 

20. 

20. Medulla C+ red or rose. 

21. Medulla all white. 

Medulla C+ orange (barbatic acid) .......................... 53. H. physcioides 

21. Lower part of medulla pigmented orange .......... 49. H. osteoleuca 

22. Medulla C + deep red (anziaic acid) .................. 19. H. eitenii 

22. Medulla C+ rose or C- ........................................ 8. H. chicitae 

19. Medulla C- . 23. Medulla P+ red (protocetraric acid). 

24. Lichexanthone present in the cortex (UV+ orange). 

25. Lobes short, brittle and fragile .......................... 42. H. minima 

26. Thallus dark gray; lobes about 1 mm wide, short .................. 

.......................................................................... 46. H. obscurella 

llus whitish gray; lobes 2-4 mm wide ..11. H. contradicta 

25. Lobes sublinear-elongate, coriaceous ............... .5. H. brasiliana 

24. Lichexanthone absent (UV-). 

23. Medulla P-. 

27. Lichexanthone present in cortex (UV+ orange) ................... 

27. Lichexanthone absent. 

.................... ............................................................ 23. H. eryth 

28. Lividic acid group present 

28. Lividic acid goup absent. 

............................ 38. H. livida 

29. Alectoronic acid present .............................. 58. H. potenta 

29. “Gracilescens” unknown present ......... 30. H. gracilescens 

31. Norstictic acid present ........................................................ 72. H. sublmigata 

31. Norstictic acid absent ............................................................. 4. H. boquetensis 

32. Medulla PS red (protocetraric acid) ......................................... 2. H. bahiuna 

17. Thallus collected on trees. 

30. Medulla K+ yellow turning red (salazinic acid). 

30. Medulla K- or K+ pale yellow. 

32. Medulla P - . 33. Medulla C+ orange, red, or rose. 

34. Medulla C+ orange (barbatic acid group). 

35. Apothecia strongly stalked .............................. 52. H. peruuimta 

35. Apothecia adnate. 

36. Maculae absent to weakly developed 53. H. physcioides 

36. Maculae strongly developed (Figure 40 ............. 

................................................... 74. H. subp 

34. Medulla C+ red or rose. 

37. Lobes subirregular, broadly canaliculate: C 4- rose (gyroph37. Lobes sublinear, not canaliculate. 

oric acid) .......................................................... 55. H. pluriformis 

present .......................................................... 61. H. pulvinatn 

38. Maculae usually conspicuous; evernic and lecanoric acids 

38. Maculae absent: olivetoric acid present ............................. 

................................................................... 33. H. intercalanda 

39. Lobe margins distinctly lobulate ......................... 68. H. singularis 

40. Lichexanthone present in cortex (UV+ orange) .................... 

............................................................................. 45. H. novella 

40. Lichexanthone absent. 

33. Medulla C--. 

39. Lobe margins without lobulae. 

41. Alectoronic acid present ................................ 15. H. degelii 

42. Lividic acid present .................................. 38. H. livida 

42. “Palmarum” unknown present ... 50. H. palmmum 

41. Alectoronic acid absent. 
NUMBER 25 28 

Species Treatment 

The species are arranged in alphabetic order. 

Standard herbarium acronyms are given to show 

the location of specimens cited except for collections by Hale, all of which are in US. 

1. Hypotrachyna andensis, new species 

FIGURE la 

Thallus arcte adnatus vel adnatus, fragilis, 4-7 

cm latus, viridi-flavicans, lobis sublinearibus, separatis, 1-3 mm latis; superne isidiatus, isidiis lobulatis; cortex superior 12p-14p crassus, stratum gonidiale 24p-28p crassum, medulla alba, 120p-130y 

crassa, cortex inferior 12~-14~ crassus; subtus niger, 

dense rhizinosus. Apothecia substipitata, sporis 

simplicibus, lop X 14p. 

Thallus closely adnate on twigs or on soil, fragile, 

4-7 cm broad, yellowish green; lobes sublinear, 

rather elongate, mostly separate, 1-3 mm wide; 

upper surface plane, continuous, becoming isidiate, 

the isidia at length lobulate, the lobules dorsiventral, about 0.5 mm long, often with short cilia; 

lower surface densely rhizinate, the rhizines 

moderately dichotomously branched. Apothecia 

substipitate, 1-3 mm in diameter; spores lop X 14~. 

CHEmsmY.-Cortex K-, medulla K+ yellow 

turning red, C-, P+ orange (usnic acid, norstictic 

acid, and salazinic acid (holotype); or more rarely 

norstictic acid, stictic acid, and constictic acid). 

TYPE CoLLEcTIoN.-San Martin, Peru, Holligan 

L 128 (BM, holotype). 

DIsTRIsuTIoN.-Central America, Colombia, Venezuela, and Peru. 

HABITAT.-On branches of small shrubs in the 

paramo or on road banks at 2800-3700 m elevation. 

REMARKS.-This paramo species usually is 

densely isidiate-lobulate. The presumptive progenitor is H. reducens. The chemical variation 

parallels that in H. sinuosa, the sorediate morph. 

Almeda 2457 (DUKE). Colombia: Cundinamarca, 

Cuatrecasas 2706813 (US). Venezuela: Merida, Hale 

42658a, 43072, 43146. Peru: Bryan 596 (WIS). 

SPECIMENS EXAMINED.-costa Rica: Cartago, 

2. Hypotrachyna bahiana, new combination 

FIGURE 7b 

Parmelia bahiana Sylander, 1885: 612. [Type collection: Bahia, 

Brazil, Blanchet (H, lectotype; M, P, isolectotopes).] 

Parrnelia insinuans Nylander, 1885:612. [Type collection: 

Tropical America, Bonpland (H, lectotype; P, isolectotype).] 

Parmelia pava f. albescens Krempelhuber, 1873:lO. [Type collection: Serra da Mantiqueira, Brazil, Warming (M, lectotype).] 

Thallus adnate, mineral gray, 3-6 cm broad; 

lobes sublinear, short, 1-2 mm wide; upper surface 

plane, continuous, soredia and isidia lacking; lower 

surface moderately to densely rhizinate, the rhizines densely dichotomously branched. Apothecia 

common, adnate, 2-4 mm in diameter; spores 

6p-8p X lop-13p. 

CHEMISTRY.-cOrteX Kf yellow, medulla K--, 

C -, P + orange-red (atranorin, protocetraric acid, 

and, if pigmented in the lower medulla, 

rhodophyscin). 

DISTRIBUTION.-weSt Indies, Brazil. 

HABITAT.-On trees in open forest at 1000-2000 

m elevation. 

REMARKs.-Hypotrachyna bahiana is externally 

similar to H. physcioides but differs in chemistry. 

It is common only in Brazil. The isidiate morph 

is H. consirnilis. 

SPECIMENS EXAMINED.-Haiti: Sud, Zmshaug 

23229, Wetmore 3364 (MSC). Brazil: Minas Gerais, Warming 280 (M); Rio de Janeiro, Eiten 

6433, 7502 (US), Glaziou 1119 (G,H,P). 

3. Hypotrachyna bogotensis, new combination 

FIGURE 7c 

Parmelia bogotensis Vainio, 1899: (122). [Type collection: 

Bogota, Colombia, Weir 53 (TUR, lectotype; BM, isolectoParrnelia culmigena Zahlbruckner, 1905:81. [Type collection: 

type).] 

El Altar, Ecuador, Meyer 381 (\$’, lectotype).] 

Thallus adnate to loosely attached, 6-12 cm 

broad; lobes sublinear, dichotomously branched, 

1-6 mm wide; upper surface plane, shiny, whitemaculate, densely isidiate, the isidia cylindrical 

or at length procumbent and dorsiventral; lower 

surface densely rhizinate, the rhizines finely 

branched and forming a woolly mat. Apothecia 

very rare, adnate, to 5 mm in diameter; spores 

CHEiwsTRY.-Cortex K+ yellow, medulla K-, 

C+, KC+ rose to red, P- (atranorin, evernic 

acid, and lecanoric acid). 

America, West Indies, South America south to 

6y x 121.1. 

DISTRIBUTION AND HABITAT.-Mexico, Central 
24 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 7.-Species of Hypotmchyna: a, H. andensis (Holligan L128); b, H. bahiana (Eiten 7502); 

c, H. bogotensis (Oberwinkler and Poelt 7544); d, H. boquetensis (Hale 38878) e, H. brasilianu 

(Reitz and Klein 10187); f, H. brevirhiia (Hale 42532). (Scale in mm.) 
NUMBER 25 25 

Chile on trees in secondary and primary forests 

(oak, pine, Drimys), shrubs, fence posts, rocks, 

and soil on road banks at 1800-4100 m elevation. 

REMARKS.-This is an extremely variable and 

widespread species. The isidia may be quite sparse 

and merely papillate or even subsorediate but can 

eventually enlarge and grow procumbent (Figure 

4c), almost as narrow lobules, as Vainio (1899) 

has noted. Parmelia culmigena represents this 

extreme form, It is closely related to both H. 

pulvinata, the presumptive progenitor, and H. 

rockii, a sorediate morph. Care must be taken to 

distinguish it from H. ensifolia (alectoronic acid 

present) when, as is often the case, both species 

occur in the same habitat, especially on roadsides. 

ico: Chiapas, Hale 20266, 20541, 20542; Oaxaca, 

Hale 20684, 20788; Sinaloa, Culberson 13384, 

13421 (DUKE). Costa Rica: Alajuela, Culberson 

12369 (DUKE). Jamaica: Zmshaug 13044, 15442, 

15465 (MSC, US), Plitt 129 (US). Haiti: Ouest, 

Zmshazig 22959 (MSC, US), 22958 (MSC); Sud, 

Zmshaug 23232 (MSC), Wetmore 3294, 3369 

(MSC). Dominican Republic: La Vega, Zmshaug 

23371, 23402, 23406 (MSC, US), 23537, 23548 

MSC), Wetmore 3481, 3613, 3620 (MSC, US), 

3522, 3627, 3662 (MSC). Colombia: Boyaca, 

Cuatrecasas 1364a (US); Santander, Killip and 

Smith 18942 (NY), 18594 (US). Venezuela: MCrida, Hale 42387, 42607, 43120, 43200, 43277, 

43350, Poelt and Oberwinkler 13172b, 14427~ 

(M). Peru: Bryan 399 (F). Ecuador: Jameson 

144 (BM). Chile: Aysen, Santesson 4309, 4345, 

4488, 6943, 6945 (S); Chiloe, Santesson 4192 (S); 

fuan Fernandez, Skottsbeyg 87 (NY, S, US), 198 

(BM), Zmshaug 37384 (REX); Valdivia, Santesson 3274, 3521 (S, US), 3272, 3387, 3404, 3475, 

3556, 3710, 7149 (S). 

REPRESENTATIVE s PECIM EN S EXAM INED.-MeX4. Hypotrachyna boquetensis, new combination 

FIGURE 7d 

Parmelia boquetensis Hale, 1974b:265. [Type collection: Boquete, Chiriqui, Panama, Hale 38878 (US).] 

Thallus closely adnate on branches, 8-12 cm 

broad, greenish ashy gray; lobes subirregular, 

3-4 mm wide, the margins toward the center becoming lobulate; upper surface smooth, cracked 

with age and with conspicuous marginal pycnidia; 

lower surface black, sparsely to moderately rhizinate. Apothecia common, adnate, 2-3 mm in diameter; spores 6p X 12p. 

CHEMIsmY.-Cortex Kf yellow, medulla K+ 

red, P+ orange (atranorin and salazinic acid). 

DISTRIBUTION.-Panama. 

HABITAT.-on trunks and branches of hardwood 

trees in open pastures at about 1500 m elevation. 

REMARKS.-ThiS species is still known only from 

Panama, where it is rather common in the 

Chiriqui area. While at first sight similar to H. 

sublaevigata, the lobes are larger and more irregular, pruina does not develop, and the chemistry 

is different. 

SPECIMENS EXAMINED.-Panama: Chiriqui, Hale 

38811, 38831, 38848, 38902. 

5. Hypotrachyna brasiliana, new combination 

FIGURE 7e 

Parmelia brasilianu Nylander, 1883:611. [Type collection: 

Organ Mountains, Brazil, Weddell (H, lectotype; P, isolectotype).] 

Parmelia glaziouii Muller Argau, 1889:354. [Type collection: 

Rio de Janeiro, Brazil, Glaziou 16670 (G, lectotype).] 

Thallus loosely adnate on rocks, coriaceous, 

ashy white, 6-20 cm in diameter; lobe sublinear, 

divaricate, 1-4 mm wide; upper surface plane, 

continuous, irregularly wrinkled and cracked with 

age; lower surface sparsely to moderately rhizinate, 

rhizines sparsely dichotomously branched. Apothecia common, adnate, 2-5 mm in diameter; 

spores 4y-6p X 8p-1Oy. 

red (lichexanthone and protocetraric acid). 

CHEMISTRY.-cOrteX K-, medulla K-, c-, Pf 

DIsTRIsuTIoN.-SoUtheaStern Brazil. 

HABITAT.-On rocks at open rocky slopes at 

1000-2300 m elevation. 

REMARKs.-HyPotrachyna brasiliana is one of 

the more conspicuous and commonly collected 

saxicolous lichens in the Rio area. It grows 

loosely, often on sandstone, forming large whitish 

colonies that are easily detached. Two related 

species in the same habitats, H. contradicta (lichexanthone absent) and H. eitenii (medulla Cf 

red), would have to be distinguished by chemical 

tests. 

SPECIMENS EXAMINED.-Brazil: Mato Grosso, 

iMalme 309, 312, 321B, 2246B (S); Minas Gerais, 
26 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Eiten 7028, 7031 (US), Malme 312A (UPS), 

iMosen 2319bis (UPS), Vainio 1184 (BM, M, 

TUR, UPS), 1189 (M, TUR), 1249 (TUR), 

1280 (BM, TUR, UPS); Rio de Janeiro, Eiten 

6594A (US); Santa Catarina, Reitz and Klein 

10187 (US), Ule 155 (G); SHo Paulo, Schiflner 

(W). 

6. Hypotrachyna brevirhiza, new combination 

FIGURE 7f 

Parmelia breuirhiza Kurokawa in Hale and Kurokawa, 1964: 

166. [Type collection: Isla Riesco, Magallanes, Chile, Suntesson 2066 (S, holotype; US, isotype).] 

Thallus closely adnate, ashy white, 3-6 cm 

broad; lobes subirregular and short, 1-5 mm wide; 

upper surface plane to rugulose, dull, frequently 

white-pruinose, sorediate toward the lobe tips and 

in part laminally, the soredia coarse and rather 

diffuse; lower surface densely rhizinate, the rhizines densely dichotomously branched. Apothecia 

very rare, adnate, to 5 mm in diameter; spores 

poorly developed, 3p-4~ X 6p-8~. 

CHEMIsmY.-Cortex K+ yellow, medulla K+ 

yellow turning red, P+ orange (atranorin and 

salazinic acid). 

DISTRIBUTION.-PantrOpiCal in equatorial regions and in southern Chile and Argentina. 

HABITAT.-on trees in mist forest, secondary 

forest, and on rocks at 2000-3800 m elevation 

(but near sea level in Chile). 

REhfARKs.-Additional collections of this Species 

were made in the Pacific region and Africa after 

it was described from Chile. It is essentially pantropical in montane regions. The rather diffuse 

soralia, very dense rhizine mat, and tendency to 

form coarse pruina are characteristic features. 

Closely related H. majoris (Vainio) Hale, known 

from Madagascar and eastern Asia, consistently 

produces zeorine in addition to salazinic acid. A 

negative KOH test would separate H. pseudosinuosa from H. breuirhiza. 

SPECIMENS ExAMINED.-Venezuela: Merida, Hale 

42377; Tachira, Hale 42532. Ecuador: Pinchincha, Knight 1084B (MiIS). Chile: Santiago, Mahu 

2133 (US). Other records from Chile and Argentina are listed in Hale and Kurokawa (1964:167). 

7. Hypotrachyna caraccensis, new combination 

FIGURE 8a 

Parmelia caraccensis Taylor, 1847: 163. [Type collection: Caracas, Venezuela, Linden 576 (FH-TAYL, lectotype; BM, G, 

isolec totypes).] 

Parmelia sinuosa var. caraccensis (Taylor) Lindsay, 1859:218. 

Parmelia caraccensis var. guatemalensis Steiner, 1903:234. 

[Type collection: Guatemala, Friedrichsthal (W, lectotype).] 

Parmelia caraccensis yar. guatemalensis f. adspersa Zahlbruckner, 1905:82. [Type collection: El Altar, Ecuador, Meyer 

341 (W, lectotype).] 

Parmelia endorubra Gyelnik, 1934: 154. [Type collection: Peru, 

Lillie 1040 (BP, holotype).] 

Thallus loosely attached, massicot yellow, 6-30 

cm in diameter; lobes long-linear, separate, 2-6 

mm wide; upper surface plane, continuous, lacking soredia and isidia; lower surface densely 

rhizinate, the rhizines densely dichotomously 

branched, forming a projecting mat along the 

lobes. Apothecia adnate, 3-9 mm in diameter; 

spores 6p-8~ X 11p-13p. 

orange (usnic acid, galbinic acid, norstictic acid, 

and salazinic acid). 

DIsTRIBuTIoN.-Central America southward in 

the Andes to Bolivia and in southeastern Brazil. 

ses at 1500-4200 m elevation. 

REMARKs.-This must be regarded as the most 

conspicuous and typical of all the Hypotrachynae 

in the paramo region. It grows in large mats on 

exposed humus or among grass tussocks. When 

pressed and dried it tends to fragment. It could 

only be confused with unusually large or loosely 

adnate specimens of H. enderythraea, which has 

imbricate, smaller lobes, or with H. lopezii, which 

contains stictic acid. 

The syntype from Quito, Ecuador (Jameson 

1845), is this species. The spores of var. guatemalensis are given as 14p-20p long, larger than 

in other material examined, but I doubt that this 

difference justifies varietal rank. 

SPECIMENS ExAMINED.-costa Rica: Cartago, Almeda 737e (DUKE), Brenes 34 (US), Dodge (US), 

Standley 35221 (US); San Jose, Crosby 3928 

(DUKE). Panama: Chiriqui, Kozlousky and Saw- 

~PY 178 (lVIS), Scholander (US). Colombia: Boyaca: Grzibb and Guymer L85b (BM); Cundinamarca, Barclny 6107 (US), Lindig 2594 (BM, M, 

CHEMISTRY.-cOrteX K-, medulla K+ red, P+ 

HABITAT.-On hardwood trees, humus, and mOS- 
NUMBER 25 

FIGURE 8.-Species of Hypotrachyna: a, H. caraccensis (Hale 42446); b, H. chicitae (Culberson 

13210); c, H. chlorina (Eiten 7671b); d, H. consirnilis (Eiten 7293); e, H. contradicta (Reitz and 

Klein 15727); f, H. costaricensis (Hale 42139). (Scale in mm.) 

21 
28 ShfITHSONIAN CONTRIBUTIONS TO BOTANY 

P), Soderstrom 1305 (US), Weir 17 (BM); Santander, Ariste (US), Killip and Smith 17562 (US). 

Venezuela: Merida, Dennis 1925 (BM, US), Hale 

42446, 42676, 42862, 43063, Nilson 1523 (UPS), 

Oberwinkler 13171b (M), Vareschi 946, 1472 

(US). Peru: Holligan 60B (BM), Lechler 2728 

(G, H, M). Ecuador: Jameson 137 (BM, US). 

Bolivia: La Paz: Julio 416 (US). Brazil: Minas 

Gerais, Damazio 13 (TUR); Rio de Janeiro, 

Lobb (BM). 

8. Hypotrachyna chicitae, new combination 

FIGURE 8b 

Parmelia chicitae Hale, 1971:30. [Type collection: Asunci6n, 

Costa Rica, Culberson 13210 (US, holotype; DUKE, TNS, 

isotypes).] 

Thallus loosely to closely adnate, fragile, often 

forming a thick, cushion-like mass, ashy white or 

blackening toward the center, 4-10 cm broad; 

lobes sublinear, elongate, often crowded, 1.5-4 

mm wide; upper surface plane, shiny, lacking 

soredia and isidia but with numerous pycnidia; 

lower surface densely rhizinate, the rhizines 

sparsely dichotomously branched. Apothecia rare, 

adnate, 3-6 mm in diameter; spores 5p X 6p-7p. 

C- (or C+ faint rose), KC+ rose, P- (atranorin, evernic acid, lecanoric acid, obtusatic acid, 

and norobtusatic acid). 

DISTRIBUTION.-cOStB Rica, Venezuela, and Peru. 

HABI-rAT.-On rocks, soil, or mosses in paramo 

at 3300-3600 m elevation. 

REMARKS.-The saxicolous habitat at high elevations and unusual chemistry set this species 

apart from other members of the evernic acid constellation. As Culberson and Hale (1973) discovered, it falls midway in chemical evolution in 

the group. Chromatography of the complex mixture of acids is difficult. It could be confused with 

only one other paramo species, H. protenta, 

which contains alectoronic acid (C-). 

SPECIMENS EXAMINED.-Venezuela: Mkrida, Hanselmann 50 (DUKE, US). Other records from 

Costa Rica, Venezuela, and Peru are listed by 

Hale (1971b:31). 

CHEMISTRY.-cOrteX Kf yellow, medulla K-, 

9. Hypotrachyna chlorina, new combination 

FIGURE 8c 

Parmelia chlorina Muller Argau, 1880:267. [Type collection: 

Petropolis, Brazil, Deuenter (G, lectotype).] 

Thallus adnate to loosely adnate, quite fragile, 

greenish or yellowish gray, to 8 cm broad; lobes 

short, crowded, sublinear, 2-5 mm wide; upper 

surface plane, shiny, becoming densely lobulate 

toward and at the margins, the lobules delicate, 

elongate, simple or furcate; lower surface moderately rhizinate. Apothecia rare, substipitate, 

3-7 mm in diameter; spores 7p X 17p, the asci 

turning blue with IKI. 

CHEMIsmY.-Cortex K+ yellow, medulla Kf, 

C+ yellowish, P- (atranorin, barbatic acid, 

obtusatic acid, enthothein, and unidentified 

pigments). 

DIsrRmuTIoN.-West Indies, Panama, Venezuela, Peru, and Brazil. 

HABITAT.-On trees, mossy logs, or rocks in 

cloud forest or secondary forest at 900-2500 m 

elevation. 

REMARKS.-The yellow medulla and production 

of dense lobules (Figure 4e) are characteristic of 

this species. The lobules have the same pored epicortical structure as the main thallus, as revealed 

with the scanning-electron microscope. The chemistry places it in the H. physcioides complex but 

there are no close relatives, although on hasty 

examination it might be misidentified as H. 

isidiocera, a clearly isidiate species, on account of 

the similar yellow medulla. 

SPECIMENS EXAMINED.-Haiti: Ouest, Zmshaug 

22740 (MSC, US). Panama: Chiriqui, Hale 38754, 

38770. Venezuela: Merida, Hale 42735, 43014. 

Peru: Cuzco, Hewera 3466a (US). Brazil: Minas 

Gerais, Eiten 6940, 7052 (US); Rio de Janeiro, 

Eiten 6573, 751 9 (US). 

10. Hypotracltynn consirnilis, new combination 

FIGURE 8d 

Parmelia consirnilis Vainio, 1890:58. [Type collection: Carassa, 

Minas Gerais, Brazil, Vainio 1295 (TUR, lectotype; BM, 

isotype).] 

Thallus adnate on bark, 3-6 cm broad, light 

mineral gray but turning light tan in the her- 
NUMBER 25 29 

barium; lobes sublinear, becoming crowded, 1-3 

mm wide; upper surface plane, continuous, becoming densely isidiate, isidia mostly simple; 

lower surface moderately rhizinate, the rhizines 

densely dichotomously branched. Apothecia rare, 

adnate, 3-5 mm in diameter, the amphithecium 

isidiate; spores 5y-8y X 8y-1111. 

CHEMIsmY.-Cortex K+ yellow; medulla K-, 

C -, P + orange (atranorin, protocetraric acid, 

and, if pigmented in lower medulla, rhodophyscin). 

DIsTRIBUTI0N.-Mexico, West Indies, Venezuela, 

Brazil. 

HABITAT.-On hardwoods and conifers in open 

woods at 800-1600 m. 

REhfARKs.-Hypotrachyna consimilis is as rare 

in the American tropics as its presumptive progenitor H. bahiana. It is smaller and more compact than H. imbricatula (P-). 

SPECIMENS EXAMINED.-MeXiCO: Chiapas, Hale 

20375. Puerto Rico: Home 1162 (US). Dominican Republic: La Vega, AZZard 17469 (US). 

Venezuela: Valencia, Magdefrau 493 (M). Brazil: 

Minas Gerais, Vainio 1133 (BM, K, M, TUR, 

UPS); Rio de Janeiro, Eiten 7293 (US). 

11. Hypotrachyna contradicta, new combination 

FIGURE 8e 

Parmelia contradicta Hale, 1974b:265. [Type collection: 

Teresopolis, Rio de Janeiro, Brazil, Watson 521 (BM, holotype; US, isotype).] 

Thallus loosely attached on rock, rather coriaceous, whitish mineral gray; lobes linear and 

elongate, dichotomously branched, 1.5-2.0 mm 

wide, lacking soredia and isidia; upper surface 

plane and shiny; lower surface black, sparingly 

rhizinate, the rhizines coarse, sparsely dichotomously branched. Apothecia abundant, adnate, 2-4 

mm in diameter; spores 4p X 6y. 

CHEMISTRY.-MedUlh K-, P + orange-red 

(atranorin and protocetraric acid). 

DIsTRInvTIoN.-Southeastern Brazil. 

HABITAT.-on open sandstone outcrops at about 

1000 m elevation. 

REMARKs.-The absence of yellow-fluorescing 

lichexanthone in the cortex separates this species 

from the more widespread H. brasiliana; otherwise, the two species are very close, A possible 

relative occurs in South Africa: H. fissicarpa 

(Kurokawa) Hale, which has large, radially split 

apothecia and more imbricate, sublinear lobes. 

SPECIMENS EXAMINED.-Brazil: Santa Catarina, 

Reitz and Klein 15727 (US). 

12. Hypotrachyna costaricensis, new combination 

FIGURE 8f 

Parmelia costaricensis Nylander in Polakowsky, 1877:225. 

[Type collection: Angostura, Costa Rica, Polakowsky (H, 

lectotype) .] 

Parmelia hypotrachyna Nylander, 1860:405. [Type collection: 

Tolima, Colombia, Goudot (P, lectotype).] 

Parmelia sublaeuigata f. isidiosa Muller Argau, 1880:267. 

Parmelia tropica Vainio, 1896:33. 

Parmelia tropica var. deformis Vainio, 1896:33. 

Parmelia deformis (Vainio) Vainio, 1907: 169. 

Parmelia amoena Zahlbruckner, 1908:464. [For full citations, 

see Hale, 1971a:ll.] 

Thallus adnate to loosely adnate, growing on 

rocks or trees, whitish to greenish mineral gray, 

6-10 cm in diameter; lobes sublinear to irregular, 

often imbricate, 2-6 mm wide; upper surface 

plane, usually strongly maculate, moderately isidiate, the isidia cylindrical, erect, simple to 

branched; lower surface moderately to densely 

rhizinate, forming a thick mat visible along the 

margins from above, the rhizines densely dichotomously branched. Apothecia rare, adnate, 2-7 

mm in diameter; spores 4y-6y X 8y-1011. 

CHEMIsTRY.-Cortex K + yellow, medulla negative with all reagents (atranorin and protolichesterinic and possibly caperatic acids). 

DIsrRIBuTIoN.-Mexico, Central America, West 

Indies, Andean region from Venezuela to Bolivia, 

Brazil, and Southeast Asia. 

HABITAT.-on deciduous trees, conifers, palm, 

rocks, and (rarely) soil, in scrub forests, secondary cloud forests, and coffee plantations at 1000- 

3500 m elevation. 

 REMARK^.-€€^^^ trachyna costaricensis is the 

second most common species in the genus after 

H. imbricatula in tropical America. It can be 

recognized by the uniformly isidiate surface, often 

with strong white maculation, dense rhizine mat, 

and negative chemical reactions. It occurs over a 

wide range of habitats and elevation. The fatty 

acid constituent has not been precisely identified. 

According to Dr. Chicita Culberson, it probably 

is near caperatic acid but has a different R, 

value. 

The oldest name, Parmelia hypotrachyna, is 
SMITHSONIAN CONTRIBUTIOXS TO BOTANY 

invalid in Hypotrachyna as a tautonym. It is represented by a rather unusual specimen containing 

traces of gyrophoric acid along with a fatty acid, 

but no other specimens contain this acid. Zahlbruckner (1929) transferred it to Anzia without 

seeing the type, apparently since Nylander mentioned its affinity with Anzia colpodes, presumably 

because of the dense hypothallus-like rhizine mat. 

Zahlbruckner also described P. amoena in 1908 

without having checked the type of P. tropica, 

another synonym, which he guessed was close. 

SPECIMENS EXAMINED.-Mexico: Chiapas, Hale 

20036, 20046, 20479; Mexico, Cain 27597 (TNT, 

US); Michoacan, Hale 20860, 20966; Veracruz, 

Liebmann 7562d (TUR). Guatemala: Alta Verapaz, Johnson 787 (US), Standley 70064, 70347 

(US); Baja Verapaz, Hale 38358, 38617. Costa 

Ria: Cartago, Standley 34985 (US), Stone 

(DUKE); San Jose, Standley 47329 (US). Panama: Chiriqui, Hale 38728, 38732, 38783, 38814, 

38851, 38928, Scholander (US), Wagner 42 (M). 

Cuba: Las Villas, Imshaug 24619, 24633A (MSC); 

Oriente, Imshaug 2471 1 (MSC, US). Jamaica: 

Imshaug 13926 (MSC, US), 14368, 14650, 14912, 

16225 (MSC), Orcutt 5657 (US). Haiti: Ouest, 

Zmshaug 22596 (MSC). Dominican Republic: 

La Vega, Imshaug 23814 (MSC). Dominica: 

Elliott 276 (lectotype of P. tropica Vainio) (BM, 

TUR), 12 (lectotype of P. tropica var. deformis 

Vainio) (BM, TUR), Hale 35596a. Martinique: 

Degelius (DEGEL). St. Vincent: Elliott (TUR). 

Grenada: Imshaug 16235 (MSC, US), 16260, 

16271 (MSC). Colombia: Santander, Cuatrecasas 

et al. 12470 (US), Killip and Smith 19385 (US), 

Lindig 2531 (UPS); Tolima, Quimby (US). 

Venezuela: Federal, Santesson 6678b (S). Merida, 

Hale 42201, 42242, 42271, 42738, 43301, Oberwinkler and Poelt (US). Peru: Amazonas, Wurdack 483 (US); Cuzco, Thomasson (UPS). Bolivia: La Paz, Eyerdam 25139 (F). Brazil: Minas 

Gerais, Damazio 1453 (lectotype of P. amoena 

Zahlbruckner) (WU, BM, H), Eiten 6954 (US); 

Rio de Janeiro, Ainsworth 557 (BM, US), Deventer 49 (lectotype of P. sublaevigata f. isidiosa 

Miiller Argau) (G), Eiten 7671 (US). 

13. Hypotrachyna croceopustulata, 

new combination 

FIGURE 9a 

Parnielia croceopustulata Kurokawa in Hale and Kurokawa, 

1964: 169. [Type collection: Grandfather Mountain, Avery 

County, North Carolina, Irnshaug 22275 (MSC, holotype; 

US, isotype).] 

Thallus closely adnate to bark, whitish mineral 

gray, 4-8 cm in diameter; lobes sublinear, becoming crowded, 1.5-3.5 mm in width; upper surface 

plane to rugulose, sorediate-pustulate, the soralia 

coarse and coalescing, mostly laminal, the medulla 

below often yellow ochre in color; lower surface 

moderately rhizinate. Apothecia rare, adnate, 1-6 

mm in diameter; mature spores lacking. 

C -, P + orange-red (atranorin, protocetraric 

acid, and rhodophyscin). 

DIsTRIBuTI0N.-Southeastern United States, Mexico, West Indies. 

HABITAT.-on conifers (Abies, Picea, Pinus) in 

open forests at 1400-2000 m ,elevation. 

REMARKS.-when first described, Parmelia croceopustzilata was based in part on Hypotrachyna 

gondylophora, which is also Pi- red but contains 

fumarprotocetraric acid but which, more significantly, has strictly subterminal soralia. This error 

was later corrected (Hale, 1967) and H. croceopustulata was typified as including only laminally 

pustulate-sorediate plants (Figure 2c) with protocetraric acid. It is a relatively rare species, perhaps 

most common on conifers in the southern Appalachian Mountains but absent from South America. 

Nakanishi I79 (US). Haiti: Ouest, Wetmore 3213 

(MSC, US); Sud, Zmshaug 23229 (MSC, US). 

CHEMISTRY.-COrteX K+ yellow, medulla K-, 

SPECIMENS EXAMINED.-MeXiCO: Tamaulipas, 

14. Hypotrachyna dactylifera, new combination 

FIGURE 9b 

Parrnelia dactylifera Vainio, 1890357. [Type collection: Lafayette, Minas Gerais, Brazil, Vainio 363 (TUR, lectotype; BM, 

FH, M, P, UPS, isolectotypes).] 

Parrnelia tilacen var. leucina Muller Argau, 1880:267. [Type 

collection: Petropolis, Brazil, Deventer (G, lectotype).] 

Thallus closely adnate, tannish mineral gray, 

3-8 cm broad; lobes sublinear, often crowded, 

1-4 mm wide; upper surface plane, continuous, 

with age becoming rugulose, becoming densely 

isidiate, the isidia very large, about 1 mm high, 

fragile and usually breaking open and pustulate 

apically without formation of soredia; lower sur- 
NUMBER 25 31 

face moderately rhizinate, the rhizines densely 

dichotomously branched. Apothecia rare, adnate 

to substipitate, 1-4 mm in diameter; spores 

CHEMIsmY.-Cortex K+ yellow, medulla K-, 

C-, KC+ rose, P- (atranorin, colensoinic acid, 

norcolensoinic acid, 4-0-demethylphysodic acid, 

physodic acid, lividic acid, and associated unknowns, and, if pigmented in the lower medulla, 

rhodophyscin). 

DIsTRIBuT1oN.-Mexico, Costa Rica, Panama, 

Venezuela, Chile, Brazil, Uruguay, and Africa. 

HABITAT.-On trees (hardwoods and conifers) 

and rarely on rocks, in open secondary forest or 

cloud forest at 1000-2800 m elevation. 

REMARKs-The isidia of this species are very 

large and inflated and probably are correctly interpreted as pustules. They are exceedingly variable and the range of variation is still unsettled 

in spite of the fair number of collections available. 

The chemistry is identical with that of the presumptive progenitor H. livida. Vainio (1890) 

surmised that Parmelia sublaevigata f. isidiosa 

Muller Argau might belong here, but it is a synonym of H. costaricensis. 

SPECIMENS EXAMINED.-kfeXiCO: Chiapas, Hale 

20288, 20375; Durango, Weber 33708 (COLO); 

Veracruz, Hale 19520. Costa Rica: Cartago, Torres 

156 (F). Panama: Chiriqui, Hale 38722, 38927, 

Scholander (US). Venezuela: Federal, Santesson 

6677a (S); Merida: Hale 42042, 43252, Poelt 7525 

(US). Brazil: Rio de Janeiro, Dusen 11 (S), 

Eiten 6529, 6534, 7126 (US); Sao Paulo, Sampaio 

(P). Uruguay: Osorio 5925 (MVM). Chile: Chiloe, Lobb (BM). 

5p-7~ X 9p-13p. 

15. Hypotrachyna degelii, new combination 

FIGURE 9c 

Parmelia degelii Hale in Hale and Kurokawa, 1964:170. [Type 

collection: Angola, Mexico, Degelius (Degelius herbarium, 

holotype: US, isotype).] 

Thallus rather closely adnate, coriaceous, tannish mineral gray in the herbarium, 4-8 cm 

broad; lobes short, crowded, sublinear, 1-2.5 mm 

wide; upper surface plane to rugulose, shiny, 

faintly white-maculate, lacking soredia and isidia; 

lower surface moderately rhizinate. Apothecia common, adnate to substipitate, 3-8 mm in diameter, 

the amphithecium white-maculate, incised; spores 

CHEiwsmY.-Cortex K+ yellow, medulla K-, 

C--, KC+ rose, P- (atranorin, alectoronic acid, 

and a-collatolic acid). 

DISTRIBUTION.-Colombia, Peru, Brazil, and 

Africa. 

HABITAT.-On trees, rarely rocks, in open forest 

at 1000-3500 m elevation. 

REMARKs.-Hypotrachyna degelii resembles other 

short-lobed species such as H. bahiana (protocetraric acid present) and H. physcioides (barbatic 

acid group present) and must be identified with a 

chemical test. It is biogenetically unrelated to these 

species and probably falls closer to the H. Zivida 

group. The sorediate morph is H. exsplendens. 

SPECIMENS ExAMINED.-co~ombia: Cundinamarca, King 505 (US). Peru: Bryan 703 (S, US, 

WIS). Brazil: Mato Grosso, Malme 2545" (S); 

Rio Grande do Sul, Malme 941 (S). 

4p-5p x 8p-llp. 

16. Hypotrachyna densirhizinata, new combination 

FIGURE 9d 

Parmelia densirhizinata Kurokawa in Hale and Kurokawa, 

1964: 171. [Type collection: Volcan, Chiriqui, Panama, 

Scholander (US, holotype).] 

Thallus loosely attached, whitish mineral gray, 

6-20 cm broad; lobes linear-elongate, often divaricate, separate, 2-6 mm wide; upper surface plane, 

continuous, sorediate-pustulate near the tips, the 

soralia subcapitate; lower surface densely rhizinate, 

the rhizines projecting as a mat along the margins, densely dichotomously branched. Apothecia 

very rare, substipitate, to 10 mm in diameter; 

spores lop X 17 p. 

CHEMIsmY.-Cortex K+ yellow, medulla K-, 

C--, KC+ rose, P- (atranorin, alectoronic acid, 

and a-collatolic acid). 

DIsTRIBuTIoN.-Southeastern United States, Mexico, Central America, West Indies, Colombia, 

Venezuela, Ecuador, Peru, Bolivia, Chile, Argentina, and Africa. 

HABITAT.-& trees (open pine forests, Drimys, 

Baccharis), rocks, and mossy humus in open areas 

and on roadside banks at 2000-3700 m elevation. 

REMARKS.-The lobes of this species are not as 

elongate on the average as those of its presumptive progenitor H. gigas. It has a much wider 
32 ShIITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 9.--Species of Hypotrachyna: a, H. croceopustulata (Imshaug 22383); b, H. dactylifera 

(Hale 42075); c, H. degelii (Bryan 703); d, H. densirhizinata (Hale 43194); e, H. dentella (MeCullough 570); f, H. ducalis (Holligan L59B). (Scale in mm.) 
NUMBER 25 33 

geographic range than H. gigas-as we might expect for a sorediate morph-from the cold temperate mountains of the southern Appalachians to 

southern Chile. The chemistry is uniform, and no 

specimens contain gyrophoric acid, strongly suggesting that H. densirhizinata originated before 

H. gigas interbred with a gyrophoric acidcontaining population. Without a chemical test 

H. densirhizinata could be confused with any of 

the species in the chemical complex including H. 

laevigata, H. oostingii, and H. thysanota. 

SPECIMENS EXAMINED.-MexicO: Oaxaca, Sharp 

et al. 2684 (NICH). Guatemala: Solola, Kellerman (OSU, US). Costa Rica: Cartago, Culberson 

12416 (DUKE). Colombia: Tolima, Quimby (US). 

Venezuela: Merida, Hale 42389, 42695, 42808, 

43194, Hanselmann 62 (DUKE, US). Chile: Juan 

Fernandez, Imshaug 37469 (MSC); Valdivia, Santesson 3604 (S, US). Argentina: Lamb 6003 

(UPS). For additional records from Guatemala, 

Panama, Haiti, Dominican Republic, Colombia, 

Peru, Ecuador, Bolivia, and Chile see Hale and 

Kurokawa (1964: 171-172). 

17. Hypotrachyna dentella, new combination 

FIGURE 9e 

Parrnelia dentella Hale and Kurokawa, 1964: 172. [Type collection: Cheaha State Park, Clay County, Alabama, McCullough 570 (US, holotype; TNS, isotype).] 

Parmelia laevigata var. ceratina Muller Argau, 1880:267. 

[Type collection: Petropolis, Brazil, Deventer (G, lectotype).] 

Thallus loosely adnate on rocks, 3-8 cm broad, 

greenish mineral gray; lobes subirregular to linear, short, 2-6 mm wide; upper surface plane, 

continuous, often quite white-maculate, densely 

isidiate, the isidia simple to branched, up to 0.7 

mm high; lower surface sparsely to moderately 

rhizinate, the rhizines sparsely dichotomously 

branched. Apothecia rare, adnate, 1-2 mm in 

diameter; spores 4p-6~ X 8p-101.1. 

CHEMIsmtY.-Cortex K+ yellow, medulla K+ 

yellow, C- or C+ pale yellow, P+ orange 

(atranorin, barbatic acid, echinocarpic acid, 4-0- 

demethylbarbatic acid, obtusatic acid, and norobtusatic acid). 

DIsmtIsuTIoN.-Southeastern United States, Mexico, Venezuela, and Brazil. 

HABITAT.-On rocks in open woods and pastures at about 1000 m elevation. 

REMARKS.-This species is closely related to H. 

imbricatula but differs in being smaller and in 

producing P + echinocarpic acid. 

SPECIMENS EXAMINED.-Mexico: Tamaulipas, 

Nakanishi 11 1 (US). Venezuela: Merida; Hale 

42727. Brazil: Rio de Janeiro, Eiten 6532 (US). 

18. Hypotrachyna ducalis, new combination 

FIGURE 9f 

Parmelia ducalis Jatta, 1908:407. [Type collection: Mobuku 

Valley, Nabitava verso Kirchuchu, and Duroni TV of Valle 

dei Laghi, Ruwenzori, Aputi (not seen).] 

Thallus loosely adnate, whitish mineral gray, 

about 10 cm broad; lobes linear-elongate, separate 

to divaricate, 2-5 mm wide; upper surface plane, 

shiny, soredia and isidia lacking; lower surface 

densely rhizinate, the rhizines densely dichotomously branched. Apothecia seen only in African material: 3p-4p X 8p-9p. 

CHEMIsmY.-Cortex K+ yellow, medulla KC+ red, PDIsTRIsurIoN.-Venezuela, Peru, and Africa. 

HABITAT.-oVer mosses and humus at the base 

of trees in thickets in open areas and paramo at 

3000-3800 m elevation. 

REMARKS.-The type of H. ducalis has not yet 

been found, but Jatta's description clearly refers 

to a nonsorediate linear-lobed species similar to 

other collections made in the mountains of Kenya 

and Uganda. In any event, in chemistry and 

morphology the population in South .hierica is 

identical to the population in Africa. Externally, 

H. ducalis is close to H. gigas, which contains 

alectoronic acid. Vegetative morphs include H. 

producta (sorediate), H. partita (isidiate-lobdate), and H. rachistn (lobulate). 

SPEcIivENs EXARIINED.-veneZuela: Merida, Hale 

43059. Peru: San hiartin, Hollignn L59B (BII). 

(atranorin and anziaic acid). 

19. Hypotrachyna eitenii, new combination 

FIGURE 10n 

Pnrrnelin eitenii Hale, 1974b:2GG. [Tlpe collection: Seirn dos 

Orgaos, Rio de Janeiro, Brazil, G. n~d I-. Eiteti 7123 (L'S,.] 

Thall~i~ loosely attached to rocks, brittle. min- 
34 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

era1 gray, about 6 cm in diameter; lobes sublinear, 

dichotomously branched, about 3 mm wide; upper 

surface plane, without soredia or isidia; lower 

surface sparsely to moderately rhizinate, the rhizines long. Apothecia abundant, adnate, to 4 mm 

in diameter; spores 5p X 7p-8y. 

CHEMIsmY.-Cortex K+ yellowish, medulla K- , 

Cf red, P- (trace atranorin, lichexanthone, and 

anziaic acid). 

DIsmIsuTI0N.-Southeastern Brazil. 

HABITAT.-Rocks in scrubby forest at about 

1800 m elevation. 

REMARKS.-ThiS saxicolous species is very close 

to H. brasiliana in external appearance and in 

habitat. As a possible recent derivative of the H. 

brasiliana group, it diverges very significantly in 

chemical biogenesis. 

20. Hypotrachyna enderythraea, new combination 

FIGURE 10b 

Parmelia enderythraea Zahlbruckner, 1929: 136. [Type collection: Based on P. erythrocardia Zahlbruckner.] 

Parmelia erythrocardia Zahlbruckner, 1908, p. 466. [Type collection: Camarinhas, Brazil, Damazio 1761 (W, lectotype). 

Not P. erythrocardia (Miiller Argau) Vainio, 1900:7.] 

Thallus adnate to loosely adnate on rock or 

soil, massicot yellow, 4-8 cm in diameter: lobes 

variable, linear-elongate, often crowded, 1-3 mm 

wide; upper surface plane, continuous, sometimes 

pruinose at the tips; lower surface moderately 

rhizinate, the rhizines often projecting beyond the 

margins. Apothecia rare, adnate, 2-15 mm in 

diameter; spores 7p-8p X 131.1-151.1. 

CHE;\IIsmY.-Cortex K-; medulla K+, Pf 

orange to red (galbinic, norstictic, salazinic, and 

usnic acids). 

DIsTRIBuTIoN.-Panama, Jamaica, Andean region from Venezuela to Bolivia, and southeastern 

Brazil. 

HABITAT.-On soil and rocks in open forests and 

in the paramo at 2200-3700 m elevation. 

REMARKs.-Degree of lobe adnation and width 

are rather variable in this species. The lobes may 

he separate and elongate to imbricate and short, 

variation that may result from environmental 

modification, for it grows in exposed paramo 

areas. The chemi5try places this species close to 

H. microhlmta, a closely related isidiate morph. 

Hypotrachyna caraccensis is a good candidate for 

the sexual morph because of the identical chemistry, but it is much larger and divaricate. 

Culbel-son 13730, 13733 (DUKE, US). Venezuela: 

Mkrida, Hale 42464, 42898, 43150, 43218, 43324, 

43340 (US), Hanselmann 33 (DUKE, US), Oberwinkler and Poelt 7542, 7547 (M, US). Peru: 

Chachapoyas, Matthews (US), Wurdack 484 (US); 

Cuzco, Iltis 3313 (WIS), Thomasson (UPS). Bolivia: Julio 414 (US). Ecuador: Fiske (US). 

SPECIMENS EXAMINED.-Jamaica: Culberson and 

21. Hypotrachyna endochlora, new combination 

FIGURE 1Oc 

Parmelia endochlora Leighton, 1871:140. [Type collection: 

Askew Wood, Kerry, Ireland, Taylor (BM, lectotype).] 

Parmelia millaniana Stirton, 1874:79. [Type collection: Loch 

Long, Argyleshire, Scotland, Macmillan (BM, lectotype).] 

Parmelia xanthomyela Nylander, 1874:306. [Type collection: 

Fontainebleau, France, Nylander (H, lectotype).] 

Parmelia pittieri Muller Argau in Durand and Pittier, 1891: 

53. [Type collection: Rancho Rotundo, Costa Rica, Pittier 

5050 (G, lectotype).] 

Parmelia madagascariensis Vainio, 1898: (33). [Type collection: Tanala, Ambohimitombo, Madagascar, Forsyth 562 

(TUR, lectotype; BM, isolectotype).] 

Parmelia xanthomyela f. sorediosa Hue, 1899: 146. [Type collection: Mafate, Bourbon, Rodriguez (P, lectotype).] 

Parmelia endochlora f. sorediosa (Hue) Zahlbruckner, 1929: 

166. 

Parmelia gallicana Gyelnik, 1931:288. [Type collection: Based 

on Parmelia xanthomyela f. sorediosa Hue).] 

Thallus loosely adnate, greenish mineral gray, 

8-13 cm broad; lobes sublinear, margins sometimes dissected with age, 2-6 mm wide; upper surface shiny, plane, continuous, faintly to strongly 

white-maculate, pustulate, the pustules mainly 

suhterminal, soon turning granular-sorediate; medulla pale cendre green; lower surface moderately 

to densely rhizinate, the rhizines densely dichotomously branched. Apothecia rare, adnate, 4-14 

mm in diameter; spores 9y-14p X 18p-22p. 

C+ orange, P- (atranorin, barbatic acid, obtusatic acid, norobtusatic acid, entothein, undetermined pigments, rarely echinocarpic acid). 

DIsmIBuTIoN.-Mexico, Central America, Colombia, Venezuela, Brazil, western Europe, South 

Africa, Madagascar, St. Helena, Hawaii. 

HABITAT.-on trees, rarely rocks, and over mosCHEMISTRY.-cOTteX K+ yellow, medulla K- , 
NUMBER 25 35 

ses in mature forests, primary cloud forest, and 

scrubby areas at 1700-3500 m elevation. 

REMARKs.-This widespread species is very similar to H. laeuigata except for the yellowish medulla. It occurs over a broader range, however, 

including Africa and the Pacific. The pigmentation is apparently the same as that in H. chlorina 

and H. isidiocera but the results with TLC are 

unsatisfactory for resolution. Actually the chemistry is variable, with one or two of the minor 

constituents and echinocarpic acid sometimes 

absent. 

SPECIMENS EXAMINED.-Mexico: Michoacan, Hale 

20973. Costa Rica: San Jose, Culberson 12572 

(DUKE). Panama: Chiriqui, Hale 38931, Scholander (US). Dominican Republic: La Vega, Wetmore 3509 (MSC). Colombia: Tolima, Quimby 

37 (US). Venezuela: Mkrida, Hale 43261. Brazil: 

Rio de Janeiro, Eiten 6577, 7093, 7709 (US). 

Uruguay: Higgins (BM). 

22. Hypotrachyna ensifolia, new combination 

FIGURE 10d 

Parmelia ensifolia Kurokawa in Hale and Kurokawa, 1964: 172. 

[Type collection: Alto de la Bandera, La Vega, Dominican 

Republic, lmshaug 23430 (MSC, holotype; US, isotype).] 

Parmelia lobulifera var. insensitiva Degelius, 1941:63. [Type 

collection: Forney Ridge, Swain County, North Carolina, 

Degelius (Degelius herbarium, lectotype).] 

Thallus adnate, whitish mineral gray, 5-11 cm 

broad; lobes sublinear, dichotomously branched, 

1-5 mm wide; upper surface shiny, continuous, 

becoming densely isidiate-lobulate on the surface 

and along the margins, the lobules up to 1.5 mm 

long, often ciliate; lower surface moderately 

rhizinate, the rhizines densely dichotomously 

branched. Apothecia rare, adnate, 3-4 mm in 

diameter; spores 8p-lop X 14p-18p. 

CHEMIsmY.-Cortex K+ yellow, medulla K--, 

C--, KC+ rose, P- (atranorin, alectoronic acid, 

and a-collatolic acid or with a-collatolic present 

in trace amounts or lacking). 

DIsmIBuTIoN.-Costa Rica, Panama, West Indies, Colombia, Venezuela, and New Zealand. 

HABITAT.-on trees or boulders in cloud forest 

or on soil banks at 2000-3700 m elevation. 

REMARKs.-Hypotrachyna ensifolia is characterized by abundant lobulate isidia, often becoming 

ciliate. It may represent the isidiate morph of H. 

gigas although the lobes are shorter. It often 

occurs with morphologically similar H. bogotensis, 

which can be distinguished by a C+ rose color 

test or absence of white fluorescence in ultraviolet. 

berson 12360 (DUKE, US). Colombia: Santander, 

Killip and Smith 20281 (US). Venezuela: MCrida, 

Hale 42390, 42455, 42897, 43131, Oberwinkler and 

Poelt 7511 (M, US). Additional records from 

Mexico, Panama, Jamaica, Haiti, Dominican Republic, and Venezuela are listed in Hale and 

Kurokawa (1964: 173). 

SPECIMENS EXAMINED.-cOSta Rica: Cartago, CUl23. Hypotrachyna erythrodes, new combination 

FIGURE 10e 

Parmelia erythrodes (Zahlbruckner) Hale and Kurokawa, 

1964:173. 

Parmelia brasiliana var. erythrodes Zahlbruckner, 1909: 169. 

[Type collection: Mt. Itatiaya, Minas Gerais, Brazil, SchiBner (WIJ, lectotype).] 

Thallus adnate to closely adnate on rock, often 

brittle and forming fragile mats, whitish ivory or 

very pale yellowish ashy gray, 4-8 cm broad; lobes 

sublinear, becoming crowded and imbricate, 1-3 

mm wide; upper surface plane, continuous, without soredia or isidia; medulla white or turning 

dull salmon; lower surface moderately rhizinate, 

the rhizines sparsely dichotomously branched. 

Apothecia common, adnate, 2-6 mm in diameter; 

spores 5p6p X 9p-12p. 

P - (lichexanthone, 4-0-demethylphysodic acid, 

and physodic acid, with or without lividic acid 

and colensoinic acid, and rhodophyscin with 

associated unknowns). 

DIsmIBvnoN.-Venezuela, Peru, and Brazil. 

HABITAT.-on rocks and stone walls in sparsely 

forested areas and in the paramo at 1600-3500 m 

elevation. 

REhfARKS.-This is one of the commonest saxicolous lichens in the paramo along with H. osteoleuca, which contains olivetoric acid. Both of these 

species contain lichexanthone in the cortex and 

have a distinctive whitish to pale cream color. 

Hypotyachyna erythrodes is differentiated by the 

presence of the lividic acid complex and frequent 

production of a pale salmon or orange pigment 

in the medulla. Almost half of the specimens 

from Venezuela were found to lack lividic acid 

CHEMISTRY.-cOrteX K--, medulla K-, c-, 
36 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 10.-Species of Hypotrachyna: a, H. eitenii (Eiten 7125); b, H. enderythraea (Hale 

42464); c, H. endochlora (Hale 43261); d, H. ensifolia (Zrnshaug 23430); e, H. erythrodes (Hale 

43321);f, H. exspendens (Standleyj. (Scale in mm.) 
NUMBER 25 57 

and to moduce almost random assortments of the Based on P. frava Krempelhuber not P. paua Rebentisch, I 

remaining components. TLC cannot adequately 

resolve these substances, and a final analysis of the 

chemical variation will have to come with highpressure liquid chromatography. 

SPECIMENS EXAMINED.-VeneZUeki: Merida, Dennis 2370 (BM), Hale 42469, 42472, 42488, 42619, 

42791, 42792, 42798, 42809, 42825, 42863, 43147, 

43186A, 43321, 43329; Tachira, Hale 42556. Peru: 

Lobb (BM). Brazil: Minas Gerais, Eiten 7030 

(US). 

24. Hypotrachyna exsplendens, new combination 

FIGURE lOf 

Parmelia exsplendens Hale in Hale and Kurokawa, 1964: 174. 

[Type collection: Murdock’s Gap, Blue Mountains, Jamaica, Zmshaug 15306 (MSC, holotype; US, isotype).] 

Thallus adnate on bark, tannish mineral gray, 

4-8 cm broad; lobes short, sublinear, 1.5-2.5 mm 

wide; upper surface plane, shiny, strongly whitemaculate, sorediate, the soralia subterminal, capitate; lower surface densely rhizinate, the rhizines 

densely dichotomously branched. Apothecia not 

seen. 

CHEMIsmY.-Cortex KS yellow, medulla K-, 

P-, C-, KC+ rose (atranorin, alectoronic acid, 

and a-collatolic acid). 

DIsmIBuTIoN.-Mexico, West Indies, Guatemala, 

Costa Rica, South Africa. 

HABITAT.-on trees in evergreen forests at 900- 

1200 m elevation. 

REMARKs.-The diagnostic characters are subterminal capitate soralia, distinct white maculae, 

and presence of alectoronic acid. This species is 

much smaller and has more imbricate, adnate 

lobes than H. densirhizinata, another species with 

alectoronic acid. The presumptive progenitor is 

H. degelii, which also is known from Africa. 

(US). Jamaica: Culberson 13509 (DUKE). Other 

records from Mexico, Guatemala, and Jamaica 

are listed in Hale and Kurokawa (1964:175). 

SPECIMENS EXAMINED.-cOSta Rica: BreneS 47 

1804:303.] 

Parmelia frava Krempelhuber, 1873: 10. [Type collection: Serra 

de Piedade, Brazil, Warming 294 (M, lectotype; G, UPS, 

isolectotypes).] 

Parmelia frava var. stellata Lynge, 1914: 150. [Type collection: 

Sao Joao d’el Rey, Minas Gerais, Malme 311 (S, lectotype).] 

Parmelia flava var. subdichotoma Lynge, 1914:149. [Type collection: SLo JoLo d’el Rey, Minas Gerais, Brazil, Malme 310 

(S, lectotype).] 

Thallus adnate to loosely attached, coriaceous, 

sea-foam green, 3-5 cm broad; lobes sublinear, 

separate, 0.5-2 mm wide; upper surface plane to 

convex, continuous or irregularly cracked on older 

lobes; lower surface moderately rhizinate, the 

rhizines long, sparsely dichotomously branched, 

often projecting beyond the lobe margins. Apothecia adnate, 1-3.5 mm in diameter; spores 5p-6~ 

CHEMIsmY.-Cortex K--, medulla K-, C-, or 

KCS rose, P+ orange-red (usnic acid and protocetraric acid). 

DIsTRIsuTIoN.-Colombia, Venezuela, Peru, and 

Brazil. 

HABITAT.-On open sandstone outcrops or 

(rarely) on soil at 1200-4100 m elevation. 

REMARKS.-This saxicolous species has a fairly 

restricted distribution in northern South America. 

Vainio (1890) noted the branched rhizines in his 

descriptions and classified the species in section 

Xanthoparmelia because of the yellow color of the 

thallus. Krempelhuber (1873) reported the type 

on tree bark, but the actual specimens appear to 

be saxicolous. The great variability in lobe width 

led Lynge (1914) to describe two new varieties, 

both narrow-lobed. 

SPECIMENS ExA~fINED.-Co~ombia: Santander, 

Killip and Smith 15186 (NY, US), 17527 (US). 

Venezuela: hlerida, Hale 42121a, Oberwinkley and 

Poelt (hl, US). Peru: Lobb (Bhi). Brazil: Bahia, 

Blanchet (hi); hiinas Gerais, Damnzio 959 (TV), 

1402 (G), Eiten 7003, 5027 (US), Hensclien 

(UPS, US), T’ainio 1163 (Bll, TUR, UPS), 1522 

(Bhl, &I, TUR); Rio de Janeiro, Gaztdichaird 9 

x lop-12p. 

(H). 

25. Hypotrachyna flavida, new combination 26. Hypotrachyna flacocirens, new combination 

FIGURE lla FIGURE Ilb 

Parmelia Pauida Zahlbruckner, 1929: 137. [Type collection: Partnelia pnvovirens Kurokawa in Hale and Kurokawa, 196-1: 
38 S.MITHSONIAN CONTRIBUTIONS TO BOTANY 

176. [Type collection: 50 km SE Natales, Magallanes, Chile, 

Santesson 5681 (S, holotype; US, isotype).] 

Thallus adnate, sea-foam green, 3-6 cm in diameter; lobes sublinear, short and sometimes 

imbricate, 1-2.5 mm wide; upper surface plane, 

shiny, becoming rugulose with age, lobulate toward the center, sorediate, the soralia mainly 

subterminal, capitate; lower surface densely 

rhizinate, the rhizines densely dichotomously 

branched. Apothecia adnate, 24 mm in diameter, 

the amphithecium rugose; spores 71.1-lop X 

CmmsmY.-Cortex K-, medulla K-, C-, 

P + orange-red (usnic acid and protocetraric acid). 

DIsTRIBUTIoN.--Southern Chile. 

HABITAT.-oII trees (Nothofagus) and on rocks 

at low elevation. 

REMARKs.-This species is essentially indistinguishable from the typical form of H. sinuosa, 

a pantemperate-pantropical montane species also 

common in the range of H. flavovirens. The 

chemical pattern (protocetraric or protocetraric 

with norstictic and salazinic acids) suggests a backcrossing between the progenitor of H. flavovirens 

(extinct?) and that of H. sinuosa (H. reducens). 

SPECIMENS EXAMINED.-chile: Magallanes, Suntesson 8191 (S, US); Tierra del Fuego, Santesson 

7460 (S). Other records from Chile are listed in 

Hale and Kurokawa (1964: 176). 

11p-15p. 

27. Hypotrachyna formosana, new combination 

FIGURE llc 

Parmelia formosana Zahlbruckner, 1934:57. [Type collection: 

Mt. Arisan, Nimandaira, Taiwan, Asahina 70 (W, lectotype; 

TNS, isolectotype).] 

Thallus adnate to closely adnate, whitish mineral gray, 3-8 cm in diameter; lobes sublinear, 

crowded, 1-3 mm wide; upper surface plane to 

rugulose, continuous, moderately to densely pustulate over most of the surface, the pustules breaking open apically but not becoming sorediate; 

medulla white or ochre below the pustules; lower 

surface moderately rhizinate. Apothecia rare, adnate, the amphithecium pustulate, 3-5 mm in 

diameter; spores 4y-6p X 7p-llp. 

Cmarrsmy.-Cortex K-, medulla K+ reddish, 

C--, P- or P+ orange (lichexanthone, colensoinic acid, norcolensoinic acid, 4-0-demethylphysodic acid, physodic acid, lividic acid, and 

associated unknowns, rarely protocetraric acid, 

and, if pigmented, rhodophyscin). 

Central America, West Indies, Venezuela, Peru, 

Brazil, Africa, India, southeastern Asia, and 

Japan. 

HABITAT.-on trees (hardwoods and conifers) 

in upland cloud forest, secondary forests, and pine 

forests and on rocks in open areas at 900-3000 m 

in tropical regions and down to sea level in temperate areas. 

REMARKs.-HyPotrachyna formosana is pantropical with wide ecological amplitude, ocurring 

on both trees and rocks. The dense pustules and 

positive orange fluorescence under ultraviolet 

instantly identify it. It belongs to the H. livida 

complex and the sexual morph seems to be rare 

H. novella. The chemistry is very uniform and 

identical with that of H. liuida except for the rare 

occurrence of protocetraric acid. The second syn 

type of Parmelia formosana, (Asahina 60 from 

Taiwan) can be identified with H. exsecta (Taylor) Hale, an Asian species with barbatic acid. 

SPECIMENS EXAMINED.-Mexico: Chiapas, Hale 

19942, 201 10, 20417, 20469, 20806; Michoacan, 

Hale 20852. Guatemala: Alta Verapaz, Standley 

71865 (US); Baja Verapaz, Hale 38370, 38384; 

Huehuetenango, Standley 82325 (US). Panama: 

Chiriqui, Scholander (US). Cuba: Oriente, Imshaug 24757, 24760 (MSC). Haiti: Ouest, Imshaug 

22546 (MSC). Venezuela: Merida, Hale 42069, 

42570. Peru: San Martin, Allard 22578b (US). 

Brazil: Rio de Janeiro, Eiten 6460, 7671a (US). 

Chile: Chiloe, Santesson 4107 (S); Valdivia, Santesson 7261 (S). 

DISTRIBUTION.-EaStern United States, Mexico, 

28. Hypotrachyna gigas, new combination 

FIGURE lld 

Parmelia gigas Kurokawa in Hale and Kurokawa, 1964:177. 

[Type collection: Bogota, Colombia, Joseph B7O (US, holotype: DUKE, LD, MSC, REN, S, TNS, UPS, WIS, isotypes).] 

Parmelia exporrecta Kurokawa in Hale and Kurokawa, 1964: 

174. [Type collection: Cerro San Felipe, Oaxaca, Mexico, 

Hale 20722 (US, holotype).] 

Thallus loosely attached and divaricate, rarely 

pendulous, 8-40 cm broad; lobes elongate, linear, 

separate, 3-10 mm wide; upper surface plane, continuous, without soredia or isidia; lower surface 
NUMBER 25 

FIGURE 1 1.-Species of Hypotrnchyna: n, H. flavida (Eiten 7003); 6, €I. flnriorJirens (Satitussoti 

5681); c, H. formosann (Asahina F70); d, H. gigns (Hole 42373); e, H. gotidylophorn (Hale 

20879); f, H. gracikscens (Vainio 1917 in BM). (Scale in mm.) 

39 
40 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

moderately to densely rhizinate, the rhizines conspicuous as a mat along the lobe margins, densely 

dichotomously branched. Apothecia very rare, 

substipitate, up to 15 mm in diameter, the amphithecium rugose; spores 8p-lop X 16p-l8p, the 

episporium 1.5p thick. 

CHEMIsmY.-Cortex K+ yellow, medulla K-, 

C- or C+ rose, KC+ red, P- (atranorin, alectoronic acid, and a-collatolic acid with gyrophoric 

acid as an accessory substance). 

DIsmIBuTIoN.-Mexico, Panama, Colombia, 

Venezuela, Ecuador, Peru, and Bolivia. 

HABITAT.-on small trees, humus, soil along 

road banks, and rarely rocks in high-elevation, 

humid forest and in the paramo at 2600-3900 m 

elevation. 

REMARKS.-This is one of the lichen spectaculars of the paramo region along with H. caraccensis. It grows loosely on humus and festoons 

small shrubs in the paramo. Individual lobes may 

be up to 20 cm long. At lower elevations it forms 

extensive suberect colonies on exposed tree trunks. 

The two chemical populations are morphologically identical and appear to be sympatric, at least 

in Venezuela. I am regarding these as one species 

for the present. The holotype of Parmelia exporrecta contains gyrophoric acid in addition to alectoronic acid and a-collatolic acid; the holotype of 

P. gigas lacks gyrophoric acid. 

SPECIMENS EXAMINED.-Mexico: Oaxaca, Hale 

20723 (DUKE, US), Skorepa 5559 (US); Veracruz, Liebmann 116 (UPS). Colombia: Boyaca, 

Cuatrecasas 1348 (US); Cundinamarca, Lindig 

2573 (P), Ariate (UPS). Venezuela: Merida, 

Hale 42360, 42367, 42376, 42673, 43158, 43187, 

43231, Linden 1088 (BM), 1089 (P), Nilsson 1508 

(UPS), Oberwinkler and Poelt 7519 (M, US). 

Peru: Cuzco, Gomez 631 (US), Holligan 38 (BM). 

Bolivia: Bang 1853 (H, NY), Spruce 174 (BM). 

Argentina: Patagonia, Dusen 122 (S). Additional 

localitie., for Mexico, Panama, Venezuela, Colombia, and Ecuador are listed in Hale and 

Kurokawa (1964: 177, 178). 

29. Hypotrachyna gondylophora, new combination 

FIGURE lle 

Purrnelia gondylophoru Hale, 1967:420. [Type collection: 

Clingmans Dome, Swain County, North Carolina, Hale 

33570 (US, holotype; DUKE, FH, UPS, isotypes).] 

Thallus loosely attached, whitish mineral gray, 

8-15 cm broad; upper surface plane, smooth, sometimes faintly white-maculate, sorediate toward the 

tips, the soralia capitate; lower surface densely 

rhizinate, the rhizines densely dichotomously 

branched. Apothecia not seen. 

CHEMIsmY.-Cortex K+ yellow, medulla K-, 

C -, P + orange-red (atranorin, fumarprotocetraric acid, and succinprotocetraric acid along 

with rhodophyscin (under soralia) if pigment 

present). 

DISTRIBUTION.-Southeastern United States, Mexico, Costa Rica, Dominican Republic, Venezuela, 

Bolivia, and Africa. 

HABITAT.-on trees (hardwoods and conifers), 

rarely on rocks, in cloud forests at 1000-2200 m 

elevation. 

REMARKS.-The most unusual feature of this 

species is the chemistry; no other Hypotrachyna 

is known to contain fumarprotocetraric acid 

(Hale, 1967). Otherwise it is externally similar to 

H. densirhizinata or robust forms of H. laevigata. 

SPECIMENS EXAMINED.-Mexico: Chiapas, Hale 

19920; Oaxaca, Nakanishi 349 (US); Veracruz, 

Hale 20867, 20879. Costa Rica: San Jose, Stone 

(DUKE). Dominican Republic: La Vega, 

Zmshaug 23470 (MSC, US). Venezuela: Merida, 

Oberuinkler and Poelt 7530 (M, US). Bolivia: 

Inquisive, Overbeck 5455 (US). 

30. Hypotrachyna gracilescens, new combination 

FIGURE llf 

Purrnelia gracilescens Vainio, 1890:53. [Type collection: Carassa, Minas Gerais, Brazil, Vuinio 1517 (TUR, lectotype).] 

Thallus loosely adnate on rock, buff mineral 

gray, 3-5 cm in diameter; lobes sublinear, dichotomously branched, more or less divaricateimbricate, 1-2 mm wide; upper surface plane, 

shiny, without maculae, soredia, or isidia; lower 

surface moderately rhizinate, the rhizines sparsely 

dichotomously branched. Apothecia adnate, 1-2 

mm in diameter; spores 7p X llp. 

(atranorin and one or two unidentified 

substances). 

CHEMISTRY.-cOrteX Kf yellow, K--, c-, PDIsTRIBuTIoN.-Southeastern Brazil. 

HABITAT.-on open rock outcrops at 1400-1500 

m elevation. 
KUMBER 25 41 

REMARKS.-This very rare saxicolous species is 

externally close to H. liuida but has an unusual 

chemistry. Without knowing the molecular structure of the “gracilescens” unknown, however, we 

cannot speculate on its relation to species containing lividic acid. Vainio (1890) differentiated the 

species from Parmelia chilena, P. bahiana, and P. 

subsinuosa (all KC+ red) by the negative KC 

reaction . SPECIMENS EXAMINED.-BraZi1: Minas Gerais, 

Damazio 934 (G), Vainio 1218, 1532 (BM, TUR). 

31. Hypotrachyna imbricatula, new combination 

FIGURE 12a 

Parmelia irnbricatula Zahlbruckner, 1909: 168. [Type collection: Itapecirica, Sao Paulo, Brazil, Schiflner (W, lectotype; MICH, isotype).] 

Parmelia lobulifera var. Zuteoreagens Degelius, 1941:61. [Type 

collection: Myrtle Point, Mt. LeConte, Tennessee, U.S.A., 

Degelius (DEGEL, lectotype).] For additional synonymy 

see Hale, 1971a:16. 

Thallus adnate, 5-10 cm in diameter, whitish 

mineral gray but often turning pale tannish in 

the herbarium; lobes sublinear, often crowded, 

2-4 mm wide; upper surface plane, shiny, usually 

strongly white-maculate, moderately isidiate, the 

isidia thin, cylindrical, simple to branched, sometimes darkening at the tips; lower surface densely 

rhizinate, the rhizines densely dichotomously 

branched. Apothecia infrequently developed, 2- 

10 mm in diameter; spores 7p-lop X llp-16p. 

CHEiwsmY.-Cortex K+ yellow, medulla K-, 

C+ orange, P- (atranorin, barbatic acid, 4-0- 

demethylbarbatic acid, obtusatic acid, and norobtusatic acid). 

DIsTR1BuTIoN.-Southeastern United States southward to Chile. 

HABITAT.-on trees (oaks, pine, juniper, Araucaria, Arbutus, coconut palm, citrus), stumps, 

rocks, and (rarely) soil, in primary and secondary 

forests at 200-2600 m elevation. 

REMARKS.-Hypotrachyna imbricatula is the commonest and most widely distributed species in the 

genus. It also has wide ecological amplitude. The 

isidia usually are well developed and may rarely 

become in part lobulate or procumbent. In the 

herbarium it often turns light brownish with age. 

The presumptive progenitor is H. physcioides and 

the sorediate morph H. laevigata. A very similar 

species, H. bogotensis, is separated by the C+ rose 

(rather than orange) reaction caused by evernic 

and lecanoric acids. The very common H. costaricensis has broader lobes and a uniformly dense 

rhizine mat below, as well as completely negative 

color tests. 

Chiapas, Hale 20021, 20197, 20365, 20455, 21202; 

Michoacan, Arsene 8032, 8245 (US), Darrow 

5155 (Darrow herbarium), Kiener 18456 (DUKE); 

Oaxaca, Hale 20652; Sinaloa, Breedlove 16758d 

(US); Veracruz, Hale 21059, 21 146. Guatemala: 

Baja Verapaz, Hale 38322, 38341; Jalapa, Standley 76825 (F, US); Totonicapan, Standley 84542 

(F, US); Zacapa, Steyermark (US). Honduras: 

Morazan, Standley and Molina 4320 (F). Costa 

Rica: Cartago, Culberson 13219 (DUKE); San 

Jose, Culberson 12403, 12628 (DUKE), Standley 

43033, Standley and Valerio 43174, 43479 (US). 

Panama: Chiriqui, Almeda 1025A (DUKE), Hale 

38720, 38820, 38884, 38911, Maxon 4989 (US), 

Scholander (US). Cuba: Oriente, Imshaug 25030 

(MSC); Pinar del Rio, Imshaug 25221, 25225 

(MSC, US). Jamaica: Imshaug 12950, 13382, 

13417 (MSC, US), 15278, 15531 (MSC), Orcutt 

5626 (US), Plitt (US). Haiti: Ouest, Imshaug 

22958, 22980, 23069 (MSC, US), Wetmore 3001, 

3044, 3121, 3151 (MSC, US); Sud, Imshaug 23235 

(MSC, US). Dominican Republic: La Vega, Allard 17695, 17709, 17715 (US), Imshaug 23628, 

23683 (MSC, US), Wetmore 3428, 3513 (MSC, 

US). Guadeloupe: Hale 38280. Dominica: Hale 

35345. Trinidad: Hale 38212, 38294. Colombia: 

Santander, Cuatrecasas 12470 (US), Charetier 

211 (US), Killip and Smith 18942 (US). Venezuela: Bolivar, Steyermark and Dunsterville 92981 

(US); Federal, Jahn 249 (US); Merida, Hale 

42185, 42761, 43302. Peru: Cuzco, Herrera 3461a 

(US), Worthley (US). Guyana: Lillie 47 (BM). 

Brazil: Bahia, Willey (US); Minas Gerais, Mosen 

2319 (UPS), Vainio 1085 (TUR); Rio de Janeiro, 

Eiten 6512 (US); Santa Catarina, Reitz and Klein 

11991, 15501 (US). Chile: Valdivia, Santesson 

3785 (S, US). 

REPRESENTATIVE SPECIMENS EXAMINED.-MexicO: 

32. Hypotrachyna immaculata, new combination 

FIGURE 12b 

Parmelia immaculata Kurokawa in Hale and Kurokawa, 1964: 
42 SiMITHSONIAN COKTRIBUTIOXS TO BOTANY 

178. [Type collection: Zoutpansberg, Union of South Africa, 

Almborn 6551 (LD, holotype; US, isotype).] 

Thallus loosely adnate to adnate, very fragile, 

whitish mineral gray, 5-7 cm broad; lobes sublinear, contiguous, 1.5-3 mm wide; upper surface 

plane, shiny, sorediate, the soralia mostly subterminal, large and capitate, often pigmented 

ochre below, 1-2 mm in diameter; lower surface 

densely rhizinate, the rhizines densely dichotomously branched. Apothecia rare, adnate, 2-3 mm 

in diameter; spores 6p X 11p. 

CHEMISTRY.-cOrteX K f yellow, medulla Kor K+ reddish, P- or P+ dull reddish, C 

(atranorin, colensoinic acid, 4-0-demethylphysodic acid, lividic acid, and associated unknown 

substances). 

D1sTRIBuTIoN.-Venezuela, Brazil, and South 

Africa. 

HABITAT.-& trees, rarely on soil, in open forest 

and along roadbanks at 2000-3000 m elevation. 

REMARKS.-The South American material of 

this poorly known species is variable in size and 

lobation but compares with the African population in overall morphology and chemistry. Additional collections are needed to settle any doubt. 

In any event, it would be considered as the 

sorediate morph of H. livida. On reexamining the 

type of H. immaculata I found that it consists of 

a mixture of two species, one (apparently new) 

with lichexanthone in the cortex and one (lectotype) with atranorin. 

SPECIMENS EXAMINED.-Venezuela: Merida, Hale 

42900, 42969. Brazil: Minas Gerais, Warming 

281 (M). 

33. Hypotrachyna intercalanda, new combination 

FIGURE 12c 

Parmelia intercalanda Vainio, 1890:53. [Type collection: Sitio, 

Minas Gerais, Brazil, Vainio 899 (TUR, Vainio herbarium 

number 3014, lectotype; BM, M, UPS, isolectotypes).] 

Parmelia fragilis Lynge, 1914, p. 123. [Type collection: Santa 

Anna da Chapada, Mato Grosso, Brazil, Malrne 2365* (S, 

lectotype).] 

Thallus adnate to closely adnate on bark, tannish mineral gray, 3-6 cm broad; lobes sublinear, 

crowded to separate, 0.5-2 mm wide; upper surface plane to rugulose, continuous, faintly maculate, without soredia or isidia; medulla white; 

lower surface moderately rhizinate, the rhizines 

densely dichotomously branched. Apothecia common, adnate to substipitate, the disc light brown, 

2-4 mm in diameter; spores 5p-6p X Sp-lOp. 

CHEMIsmY.-Cortex K+ yellow, medulla K--, 

Cf red (atranorin and olivetoric acid). 

DrsmIBuTIoN.-Southeastern Brazil and Argentina. 

HABITAT.-on trees, rarely on humus over rocks, 

in open woods at about 500-1200 m elevation. 

REMARKS.-The C+ red test caused by olivetoric 

acid was first noted by Vainio (1890) in the 

original description. Hypotrachyna osteoleuca also 

produces this rare acid, along with lichexanthone, 

but the two species have different geographic 

distribution and ecology. According to C. Culberson's scheme (Culberson and Hale, 1973), 

a species such as this one could have originated 

from progenitors containing microphyllinic acid 

or perlatolic acid. Further biochemical transformations of H. osteoleuca itself might have produced 4-0-methylphysodic-physodic acid progenitors of the H. livida complex. 

Malme in Lichenes Austroamericani 88 (G, LD, 

UPS); Minas Gerais, Henschen (UPS, US), Vainio 

936 (TUR, UPS) (syntype of P. intercalanda); 

Parana, Montes 10116 (DUKE, LD, MVM, US); 

S5o Paulo, Puiggari (G); Santa Catarina, Reitz 

and Klein 15985 (US). Argentina: Misiones, 

Montes 10106 (MVM, US), Osorio 4297 (MVM, 

SPECIMENS EXAMINED.-Brazil: Mato Grosso, 

US). 

34. Hypotrachynu isidiocera, new combination 

FIGURE 12d 

Parmelia isidiocera Nylander, 1860:382. [Type collection: 

Cape Krusenstern, American Arctic, Beechey (BM, lectotype: H, isolectotype).] 

Thallus loosely adnate, fragile, greenish mineral gray, 5-15 cm broad; lobes sublinear, short, 

often crowded; upper surface shiny, strongly whitemaculate, becoming densely isidiate toward the 

margins, the isidia tall, erect, easily breaking off 

at the tips to reveal the yellow medulla; medulla 

sulphur yellow; lower surface densely rhizinate, 

the rhizines densely dichotomously branched. Apothecia rare, substipitate, 2-8 mm in diameter, the 

rim isidiate, rugose; spores 7y-8~ X 12p-16p. 
NUMBER 25 

FIGURE 12.--Species of Hypotrachyna: a, H. imbricatula (Hale 21059); b, H. immaculata (Hale 

42900); c, H. intercalanda (Osorio 4297); d, H. isidiocera (Hale 20219); e, H. koyaensis (Nakanishi 

284); f, H. laevigata (Hale 43106). (Scale in mm.) 

43 
44 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

CHEMIsmY.-Cortex K 4- yellow, medulla K+, 

C+ yellowish, P- (atranorin, barbatic acid, 

obtusatic acid, entothein, undetermined pigments, 

and other substances, rarely echinocarpic acid and 

a trace of norobtusatic acid). 

DISTRIBUTION.-Mexico, Central America, West 

Indies, Colombia, Venezuela, Brazil, Hawaii. 

HABITAT.-on trees (oak, pine) in virgin pine 

forests, cloud forests, and on rocks in open woods 

at 1000-3000 m elevation. 

REMARKs.-This robust lichen is easily recognized by the bright yellow medulla exposed as 

the large isidia break away. While obviously a 

member of the H. physcioides group, it is distant 

from any known species with respect to isidial 

formation (largely marginal) and brittleness of 

the thallus. The type is listed as “American 

Arctic” but this is surely a mistake since the 

species is known only from montane tropical 

regions. 

SPECIMENS EXAMINED.-?VfeXiCO: Chiapas, Hale 

20219 (DUKE, US). Guatemala: Alta Verapaz, 

Johnson 171g (US), Panama: Chiriqui, Hale 

38690, Scholander (US). Jamaica: Britton 1110, 

1121 (FH), Culberson 13502, 13894 (DUKE), 

Harris 10010 (BM, FH), Imshaug 12993 (MSC, 

US), 14937 (MSC). Dominican Republic: La 

Vega, Allard 16555G (US). Colombia: Tolima, 

Quimby (US). Venezuela: Federal, Santesson 

6677B (S); Merida, Hale 43289. Brazil: Minas 

Gerais, Damazio in Zahlbruckner, Lichenes Rariores 93 (BM, P, UPS); Rio de Janeiro, Gregory 

and Ainsworth (UPS). 

35. Hypotrachyna koyaensis, new combination 

FIGURE 1% 

Parmelia koyaensis Asahina, 1953:67. [Type collection: Mt. 

Koya, Kishu, Japan, Asahina 52101 (TNS, lectotype).] 

Thallus adnate, membranaceous, ashy white, up 

to 10 cm broad; lobes irregular, subrotund apically, with crenate margins, 4-10 mm wide; upper 

surface plane, continuous, moderately isidiate, the 

isidia simple to rarely branching, cylindrical; 

lower surface black but with a narrow brown 

zone at the margins, moderately rhizinate, the 

rhizines moderately dichotomously branched. Apothecia rare, stipitate, 1-3 mm in diameter; spores 

8p X 16~. 

CHEMIsmY.-Cortex K + yellow, medulla K-, 

C--, P+ orange-red (atranorin, protocetraric 

acid, and an unidentified fatty acid). 

DISTRIBUTION.-Japan, Southeast Asia, Mexico. 

HABITAT.-On trees (Pinus) at 2100 m elevation. 

REMARKS.-ThiS species has the subirregular 

rotund lobation so typical of Asian species. The 

isidia are sparce and the rhizines sparcely to moderately branched. Its occurrence in the New World 

is difficult to explain, 

nishi 284 (US). 

SPECIMENS EXAMINED.-h‘feXiCO: Oaxaca, NUkU36. Hypotrachyna laevigata, new combination 

FIGURE 12f 

Lichen laeuigatus Smith, 1808: 1852. [Type collection: Anglesey, Caernarvonshire, England, Dauies (LINN, lectotype).] 

Parmelia laeviga ta (Smith) Acharius, 18 14: 2 12. 

Parmelia boliviana var. cephalota Zahlbruckner, 1926:16. 

[Type collection: Loma Frei Jorge, Coquimbo, Chile, 

Skottsberg 448 (W, lectotype; S, isolectotype),] 

Thallus adnate to loosely adnate, greenish to 

whitish mineral gray, 3-10 cm broad; lobes linear, 

1.5-6.0 mm wide; upper surface plane, shiny, usually strongly white-maculate, pustulate-sorediate 

toward the lobe tips, soralia capitate; lower surface 

densely rhizinate, rhizines densely branched. Apothecia rarely developed, to 5 mm in diameter, 

adnate; spores not seen. 

CHEiwsmY.-Cortex K+ yellow, medulla K-, 

C+ orange, P- (atranorin, barbatic acid, 4-0- 

demethylbarbatic acid, obtusatic acid, and norobtusatic acid). 

DIsTRIBUTION.-~nited States southward to Chile, 

Europe. 

HABITAT.-oII trees (hardwoods, conifers, 

Drimys, Nothofagus) and rocks in open cloud or 

rain forests at 600-3700 m elevation. 

REMARKS.-This was one of the first Hypotrachyna species to be described on the basis of 

European material, but the name has been 

applied in floristic lists to many different narrowlobed species. The soredia are well developed 

and the maculae often very distinct. The presumptive progenitor is H. physcioides, and the 

parallel isidiate morph is H. imbricatula. According to Culberson and Hale (1973), the obtusatic 

acid originated from a cross between a progenitor 
NUMBER 25 

containing evernic acid (H. pulvinata) and a progenitor containing barbatic acid (extinct or now 

represented by H. massartii (Hue) Hale in Asia). 

The species evolved parallel with H. rockii, the 

sorediate morph of H. pulvinata, and the two are 

very similar externally. 

SPECIMENS EXAMINED.-Mexico: Chiapas, Hale 

19885; Oaxaca, Yoshimura 650533 (NISH). Costa 

Rica: Cartago, Culberson 12419, 13221 (DUKE); 

San Jose, Ream (WIS). Jamaica: Plitt (US). 

Dominican Republic: La Vega, Imshaug 23386, 

23561 (MSC, US). Guadeloupe: Culberson 14503 

(DUKE). Dominica: Elliott 1321, 1556, 1559 

(TUR), Hale (see data in Hale, 1971a:17). COlombia: Cundinamarca, Killip 11900 (US), Nee 

and Mori 4219 (US), Weir 53 (BM); Santander, 

Chawtier 189 (US); Tolima, King 506 (US). 

Venezuela: hlerida, Hale 422 14, 42647, 43 106, 

43206. Guyana: McConnell and Quelch 503 

(BhI). Brazil: Rio de Janeiro, Eden 7093 (US). 

Chile: Aysen, Santesson 6948 (S, US); Coquimbo, 

Munoz 1257 (S), Sparre 2933 (S); Juan Fernandez, Skottsberg 285 (UPS); Valdivia, Santesson 

3273, 3277, 3464, 3786, 3698, 7571 (S). 

37. Hypotrachyna lineariloba, new combination 

FIGURE 13a 

Parmelia lineariloba Kurokawa in Hale and Kurokawa, 1964: 

180. [Type collection: Alto de la Bandera, La Vega, Dominican Republic, Wetmore 3475 (MSC, holotype; US, isotype).] 

Thallus loosely attached to bark, rather fragile, ashy white, 3-6 cm broad; lobes elongate, 

linear, dichotomously branched, subdivaricate, 

0.5-2 mm wide; upper surface plane, shiny, continuous, lacking soredia and isidia; lower surface 

moderately rhizinate, the rhizines about 1 mm 

long, sparsely dichotomously branched. Apothecia 

not seen. 

C-, KC+ rose, Pacid). 

CHEMISTRY.-cOrteX K+ yellow, medulla K--, 

(atranorin and alectoronic 

~IsTRIBUTIox.-Dominican Republic. 

HABITAT.-on trunks of Pinus in open forests 

on ridges a[ 2400-2800 m elevation. 

RERIARKS.-NO new collections of this species 

have been made and it may indeed be endemic 

to the pine forests of the West Indies. It resembles 

H. gigaT, a larger species which always contains 

a-collatolic acid along with alectoronic acid. 

SPECIMENS ExhhZINED.-Additional records for 

the Dominican Republic are listed in Hale and 

Kurokawa (1964: 181). 

38. Hypotrachyna liuida, new combination 

FIGURE 13b 

Parmelia livida Taylor, 1847: 171. [Type collection: New Orleans, Louisiana, Hooker Herbarium (FH-TAYL, lectotype; 

BM, H, isolectotypes).] 

Parmelia osseoalbida Lynge, 1914: 133. [Type collection: Porto 

Alegre, Rio Grande do Sul, Brazil, Malne 595 (S, lectotype; 

UC, UPS, isolectotypes).] 

Thallus closely adnate, rather coriaceous (on 

bark) to fragile (on rock), whitish mineral gray, 

4-9 cm broad; lobes sublinear, dichotomously 

branched, contiguous, 1-2 nim wide; upper surface shiny, plane to rugulose, lacking soredia and 

isidia; lower surface moderately rhizinate, the 

rhizines sparsely dichotomously branched. Apothecia numerous, adnate to substipitate, 1-5 mm 

in diameter; spores 5p X lop-12p. 

CHEMIsmY.-Cortex K + yellow, medulla K-, 

C-, KC+ rose, P- (atranorin, colensoinic acid, 

norcolensoinic acid, 4-0-demethylphysodic acid, 

physodic acid, lividic acid, and associated 

unknowns). 

DISTRIBUTION.-SOUtheaStern United States, Bahamas, Venezuela, southeastern Brazil southward 

to Argentina). 

HABITAT.-On tree trunks and branches or 

(more rarely) on rocks, in open forests and gallery forests at 500-2800 m elevation (to near sea 

level in the United States). 

R~~~~~s.-Hypotrachyna lividais characterized 

by the adnate thallus, lack of diaspores, and complex chemistry. While we still do not know the 

structure of lividic acid, the presence of at least 

colensoinic and physodic acids would place the 

species on a high evolutionary scale in the genus 

(Culberson and Hale, 1973). Some specimens (in 

Brazil) appear to lack lividic acid and contain 

colensoinic acid and related compounds. These 

may represent a newly evolving species but careful studies with high-pressure liquid chromatography are needed to settle the chemistry. 

The species is by far most common in the deciduous forests of southeastern United States 

(W. Culberson, 1961) and then reappears in a 

climatically similar zone in southern Brazil to 
46 ShlITHSONIAN CONTRIBUTIONS TO BOTAKY 

FIGURE 13.--Species of Hypotrachyna: a, H. lineariloba (Wetmore 3425); b, H. livida (Reitz and 

Klein 15150); c, H. lopezii (Hale 42463); d, H. mnltnei (ibfalme 2750); e, H. microblasta (Hale 

42031); f, H. minima (Malrne 2747B in S). (Scale in mm.) 
NUMBER 25 47 

Argentina. The rock form described by Lynge as 

Paymelia osseoalbida is somewhat more fragile 

than corticolous forms. 

Hypotrachyna livida is the progenitor of several 

vegetative morphs: H. immaculata (sorediate), H. 

dactylifera (isidiate-pustulate), and H. pustulata 

(pustulate). This whole complex has evolved 

parallel with H. novella (a species containing 

lichexanthone), as discussed below. 

tesson 6682 (S, US); Merida, Hale 42833; Valencia, 

Magdefrau 389 (M). Brazil: Minas Gerais, Vainio 

1400 (BM, TUR, as P. gracilis); Rio de Janeiro, 

Eiten and Eiten 6598, 7396, 7403, 7468 (US), Glaziou (G); Rio Grande do Sul, iMaZme 1330B (S); 

Sao Paulo, Dhen 11 (S); Santa Catarina, Reitz and 

Klein 13213, 15185, 15809, 16002 (US). Uruguay: 

Lavalleja, Hosseus 45 (H), Osorio 6480 (MVM, 

US); Maldonado, Hosseus 93457a (H), Osorio 5040 

(MVM); Rocha, Hosseus 43, 47 (H, M); Treinta y 

Tres, Osorio 6000 (MVM, US). Argentina: Without collector (M). 

SPECIMENS EXAMINED.-VeneZuela: Federal, Sari39. Hypotrachyna lopezii, new species 

FIGURE 13c 

Thallus laxe adnatus, viridi-flavicans, lobis 

lineari-elongatis, divaricatis, nigromarginatis, 1.5- 

2 mm latis; cortex superior 14p-16p crassus, 

stratum gonidiale 24y-18p crassum, medulla alba, 

130y crassa, cortex inferior 18p-20y crassus; subtus niger, sparse rhizinosus vel papillatus, rhizinis 

longis. Apothecia rara, substipitata, sporis simplicibus, 7p X 14p. 

Thallus loosely attached, greenish yellow, to 12 

cm broad; lobes linear-elongate, sparsely dichotomously branched, divaricate and almost appearing subfruticose, black rimmed, 1.5-2 mm wide; 

upper surface plane to concave, shiny, lacking 

isidia and soredia; lower surface in part bare and 

papillate, otherwise sparsely rhizinate, the rhizines 

long, sparsely dichotomously branched. Apothecia 

rare, substipitate, 8-11 mm in diameter, the 

amphithecium rugose; spores 7p X 14p. 

CHEmsmy.-Cortex K-, medulla K+ yellow 

turning red, C-, P+ orange (usnic acid, norstictic acid, stictic acid, and constictic acid). 

TYPE CoLLECTIoN.-La Aguada, Merida, Venezuela, Hale 43208, 6 February 1974 (US, 

holotype). 

DIsTRiBuTIoN.-Venezuela. 

HAnrTAT.-On base of shrubs and twigs in open 

rocky paramo at 2800-3500 m elevation. 

REMARKs.-This paramo species might be confused initially with H. caraccensis, but the lobes 

are narrower, usually black rimmed, and the 

rhizines very sparse. The chemistry is unusual, 

stictic acid being rare in Hypotrachyna. The 

species is named in honor of Prof. Manuel 

Lopez Figueiras. 

SPECIMENS ExAhmED.-Venezuela: Merida, Hale 

42463, 43 142, 43 195. 

40. Hypotrachyna malmei, new combination 

FIGURE 13d 

Parmelia rnalmei Lynge, 1914:116. [Type collection: Bocca da 

Serra, Mato Grosso, Brazil, Malme 2750 (S, lectotype; W, 

isolectotype).] 

Thallus loosely adnate on rocks, very fragile, 

3-5 cm in diameter, ashy white; lobes sublinear, 

short, crowded, 0.5-1.5 mm wide; upper surface 

plane, dull, pustulate, the pustules entire or bursting open apically, without forming soredia; medulla in part yellow-ochre below; lower surface 

moderately rhizinate, the rhizines sparsely dichotomously branched. Apothecia adnate, the amphithecium pustulate, 0.5-2 mm in diameter; spores 

6p-8~ X 81.1-12~. 

CHEMISTRY.-cOrteX K--, medulla K- or K+ 

reddish, P + orange (lichexanthone, protocetraric 

acid, alectoronic acid, and rhodophyscin). 

DIsTRIBuTIoN.-~oUtheastern Brazil. 

HABITAT.-on open sandstone outcrops. 

RE;CZARKs.-This rare saxicolous species is characterized by the pustules and unusual combination of chemicals. It will undoubtedly be found 

again in southeastern Brazil with more intensive 

collecting. I had originally synonymized it with 

H. minima (Hale, 1960) and compared it with H. 

foymosana. It is, however, morphologically and 

chemically distinct from both of these. 

41. Hypotrachyna microblasta, new combination 

FIGURE 13e 

Parinelia rnicroblasta Vainio, 1890:57. [Type collection: Carassa, Minas Gerais, Brazil, Vainio 1214 (TUR, lectotype; 

BM, M, P, UPS, isolectotypes).] 
48 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Parmelia revoluta f. isidiosa Muller Argau, 1891:378. 

Parmelia propagulifera Vainio, 1899: (123). 

Parmelia jarnaicensis Vainio, 1915:23. 

Parmelia pseudorevoluta Gyelnik, 1931:289. 

Parmelia neopropagulifera Gyelnik, 1938a:39. 

Parmelia endorubra f. imbricatiformis Gyelnik, 1938b3277. 

Parmelia norstictica Hale, 1959:128. 

[For full synonymy, see Hale, 1971a:18.] 

Thallus adnate to loosely attached, 5-15 cm in 

diameter, pale yellowish green, turning reddish 

in the herbarium if improperly curated; lobes sublinear, 1-4 mm wide; upper surface plane, shiny, 

without maculae, moderately to densely isidiate, 

the isidia cylindrical, tall, simple or coralloidbranched; lower surface densely rhizinate, the 

rhizines densely dichotomously branched. Apothecia 2-6 mm in diameter, adnate, the amphithecium isidiate; spores 6p-8p X 10p-13p. 

CHEMIsmY.-Cortex K-, medulla K+ red, P+ 

orange (atranorin (traces), usnic acid, norstictic 

acid, salazinic acid, and galbinic acid). 

DIsmIBuTIoN.-Mexico, Central America, West 

Indies, Andean region from Venezuela to Peru, 

southeastern Brazil, Hawaii, and Southeast Asia. 

HABITAT.-On trees (hardwoods and conifers), 

rocks, and soil in open secondary forests, pastures, 

and road banks at 900-3100 m elevation. 

RE~fARKs.-Hypotrachyna rnicroblasta is common in a wide range of habitats throughout tropical America. The extensive synonymy is difficult 

to understand in view of the general uniformity 

of the species. The thallus frequently discolors 

red when improperly pressed and dried. The 

chemistry relates it to H. enderythraea, the presumptive nonisidiate sexual morph or a population derived from it. 

ico: Chiapas, Hale 20038, 20210, 20221, 20353, 

20385, 20553; Michoacan, Arsene (US). Guatemala: Quetzaltenango, Standley 86085 (US). Costa 

Rica: Alajuela, Standley and Torres 47471 (US); 

Cartago, Standley and Valerio 46807 (US). Panama: Chiriqui, Hale 38799, 38842, 38912, 38925, 

Scholander (US). Cuba: Oriente, Znzshaug 25141, 

25179 (MSC), Morton 9595, 9596 (US). Jamaica: 

Curnrnings (FH), Hart 54 (NY), Zmshaug 12943, 

13057, 13308, 13313 (MSC), 13327 (LD, MSC), 

13857, 13923, 14815, 14898, 15452 (MSC, US), 

Maxon 8634 (US), Plitt (US). Haiti: Ouest, 

Imshaug 23088, Wetrnore 2993 (MSC, US). Dominican Republic: La Vega, Allard 17726A (US), 

REPRESENTATIVE SPECIMENS EXAMINED.-MeXZrnshaug 23389 (MSC), 23573 (MSC, US). Guadeloupe: Duss 676 (NY). Dominica: Hale 35450. 

Colombia: Cundinamarca, Cuatrecasas 231, 27068a 

(US), Weir 78 (BM); Santander, Killip and 

Smith 18942a (US). Venezuela: Bolivar, Steyermark 59817 (US); Federal, Santesson 6683 (S) 

Merida, Hale 42146, 42192, 42773, 43028, Oberzuinkler and Poelt (M, US), Vareschi 883 (US). 

Peru: Cuzco, Marin 21916 (F). Ecuador: Imbabura, Wiggins 38941 (COLO). Brazil: Sowerby 

(lectotype of P. reuoluta f. isidiosa Miiller Argau) 

(G); Minas Gerais, Vainio 1160 (BM, M, TUR, 

UPS). 

42. Hypotrachyna minima, new combination 

FIGURE 13f 

Parmelia minima Lynge, 1914: 139. [Type collection: Bocca da 

Serra, Mato Grosso, Brazil, Malme 2747 bis (S, lectotype).] 

Thallus adnate on rocks, fragile and breaking 

apart, creamy whitish gray and darkening toward 

the center, 6-8 cm broad; lobes sublinear, crowded 

and imbricate, 0.5-1.5 mm wide; upper surface 

plane to rugulose, continuous to cracked with age, 

pycnidia and papillate apothecial initials abundant; medulla orange-red in the lower part; lower 

surface sparsely to moderately rhizinate, the rhizines sparsely dichotomously branched. Apothecia 

common, adnate, 1-2.5 mm in diameter, the 

amphithecium crenate and bullate; spores 7y X 

lop. 

CHEMISTRY.-COrteX K-, medulla K-, C-, 

KC + reddish, P + orange (lichexanthone and 

protocetraric acid). 

DISTRIBUTION.-SOUtheaStern Brazil. 

HABITAT.-on open sandstone rocks at about 

1000 m elevation. 

REMARKS.-ThiS saxicolous species is as rare as 

its close relative H. rnalrnei in Brazil. The pustules, which I originally described (Hale, 1960), 

are entire and confined to the amphithecium. 

This difference, in addition to chemistry (lack of 

alectoronic acid), clearly separates H. minima 

from H. rnalrnei. 

43. Hypotrachyna monilifera, new combination 

FIGURE 14a 

Parmelia monilifera Kurokawa in Hale and Kurokawa, 1964: 
NUhIBER 25 49 

182. [Type collection: La Aguada, Merida, Venezuela, 

Magdefrau 651 (M, holotype; US, isotype).] 

Thallus loosely adnate on bark, 10-15 cm broad, 

whitish mineral gray; lobes linear-divaricate, 2-6 

mm wide; upper surface plane, continuous; lower 

surface densely rhizinate, the rhizines moderately 

branched, conspicuously moniliform-segmented, 

projecting as a mat beyond the lobe margins. 

Apothecia not present. 

CHEMIsmY.-Cortex K+ yellow, medulla K-, 

C+ orange, P- (atranorin, barbatic acid, 4-0- 

demethylbarbatic acid, possibly traces of obtusatic 

and norobtusatic acids). 

D~smIBuTIoN.-Venezuela. 

HABITAT.-On bark of small trees in lower 

paramo region at about 2800 m elevation. 

REMARKS.-while superficially close to H. gigas, 

this species has a stiffer thallus and peculiar moniliform rhizines easily detected with a hand lens. 

I was fortunate in being able to re-collect it at the 

type locality where, apparently, it is very rare in 

comparison with H. gigas and corticolous. The 

chemistry relates it to the H. physcioides group, 

but the elongate lobes and the rhizines put it in 

an isolated position in the genus. 

SPECIMENS EXAMINED.-Venezuela: Merida, Hale 

43085. 

44. Hypotrachyna neodissecta, new combination 

FIGURE 14b 

Parmelia neodissecta Hale, 1971c:94. [Type collection: 6 km 

SE of Zouepo, N’Zerekore, Guinea, Santesson 10597d (UPS, 

holotype; US, isotype).] 

Thallus loosely attached, rather fragile, pale 

greenish ashy gray, 3-7 cm broad; lobes sublinear 

to subirregular, in part subascending, becoming 

subcaniculate, 1-3 mm wide; upper surface shiny, 

continuous, becoming densely isidiate, isidia cylindrical, mostly simple; lower surface densely 

rhizinate, the rhizines moderately dichotomously 

branched. Apothecia adnate, the amphithecium 

isidiate, 2-3 mm in diameter; spores not found 

(in African material, 6p-8~ X 12p-14p). 

CHEMISTRY.-cOrteX K+ yellow, medulla K-, 

P-, C+ red (atranorin and gyrophoric acid). 

DISTRIBUTION.-VeneZUela, Africa, and India. 

HABITAT.-on trees in secondary forest at 2000- 

2200 m elevation. 

REMARKS.-HyPotrachyna neodissecta was first 

discovered in Guinea and subsequently collected 

in other African localities and in India. I would 

not have expected to find it in the New World. 

It is distinguished from Purmelinu dissecta (Nylander) Hale, for which it would almost certainly 

be mistaken, by the branched rather than simple 

rhizines, lack of cilia, and overall canaliculaterevolute pattern of lobation. 

SPECIMENS EXAMlNED.-VeneZuela: Merida, Hale 

42973. 

45. Hypotrachyna nouella, new combination 

FIGURE 14c 

Parmelia novella Vainio, 1890:56. [Type collection: Sitio, 

Minas Gerais, Brazil, Vuinio 1028 (TUR, lectotype; BM, 

FH, M, isolectotypes).] 

Parmelia brasilianu var. novella (Vainio) Lynge, 1914: 115. 

Parfnelia regis Lynge, 1914: 126. [Type collection: S5o Paulo 

d’el Rey, Minas Gerais, Brazil, Malme 178 (S, lectotype; 

UPS, isolectotype).] 

Thallus closely adnate on bark, tannish mineral gray, 2-4 cm broad; lobes sublinear, crowded, 

1-2.5 mm wide; upper surface plane, continuous, 

darkening toward the center; lower surface black, 

densely rhizinate, the rhizines densely dichotomously branched. Apothecia common, adnate, 2-7 

mm in diameter, the amphithecium crenulate; 

spores 6p X 12p. 

rose or C - , P - (lichexanthone, colensoinic acid, 

norcolensoinic acid, 4-O-demethylphysodic acid, 

lividic acid, physodic acid, and associated unknown substances). 

DISTRIBUTION.-veneZUela and southeastern 

Brazil. 

HABITAT.-on trees in open forests at 1000-2000 

m elevation. 

REMARKs.-HypotraChyna novella is essentially 

identical to H. liuida except for the replacement 

of atranorin by lichexanthone. It also has a much 

more restricted distribution. Chemically identical 

H. formosana could be considered as the pustulate 

vegetative morph. 

Magdejrau 398 (M). Brazil: Mato Grosso, Malme 

(S, UPS, US); Rio de Janeiro, Eiten 7724a (US); 

Sao Paulo, Hemmendorf (S). 

CHEMISTRY.-cOrteX K-, medulla K--, Cf 

SPECIMENS EXAMINED.-veneZuela: ValenCia, 
50 SSIITHSONIAN CONTRIBUTIOSS TO BOTANY 

FIGURE 14.-Species of Hypotrachyna: a, H. monilifera (Magdefrau 651); b, H. neodissecta (Santesson 10597d); c, H. novella (Malme I78 in s); d, H. obscurella (Hale 43342); e, H. oostingii 

(Dey 2403); f, H. osorioi (Osorio 6507). (Scale in mm.) 
NUhlBER 25 51 

46. Hypotrachyna obscurella, new combination 

FIGURE 14d 

Parrnelia gracilescens f. obscurella Vainio, 1890:54. [Type 

collection: Carassa, Minas Gerais, Brazil, Vainio 1180 (TUR, 

lectotype; BM, isolectotype).] 

Thallus very closely adnate to rocks, dark mineral gray, 4-6 cm broad; lobes sublinear, short, 

crowded, 1-1.5 mm wide; upper surface plane, 

shiny, cracked transversely with age, pycnidia 

numerous, isidia and soredia lacking; lower surface moderately rhizinate, the rhizines sparsely 

dichotomously branched. Apothecia common, adnate, 1-2.5 mm in diameter; spores 6y X 8y. 

CHEMISTRY.-cOrteX K-k yellow, medulla K-, 

C-, KC- or KC+ reddish, Pf orange (atranorin and protocetraric acid). 

DISTRIBUTION.-Venezuela and Brazil. 

HABITAT.-on exposed rocks in open areas and 

in the paramo at 1500-3500 m elevation. 

REMARKs.-This dark, small, saxicolous lichen 

blends in with the background igneous rock substratum and is difficult to collect because of the 

close adnation. It is very common on boulders in 

the paramo regions of Venezuela and probably 

occurs throughout the Andes. There are no close 

relatives in the genus. 

SPECIMENS EXAMINED.-T'eneZuela: Merida, Hale 

43188A 43322, 43342. Brazil: Minas Gerais, 

Vainio 1431 (BM, TUR). 

47. Hypotrachyna oostingii, new combination 

FIGURE 14e 

Parmelia oostingii Dey, 1974:362. [Type collection: Balsam 

Cone, Yancey County, North Carolina, Dey 1236 (DUKE, 

holotype; US, isotype).] 

Thallus loosely attached, firm, whitish mineral 

gray, 8-12 cm broad; lobes sublinear, dichotomously branched, more or less separate, 2-5 mm 

wide; upper surface plane, shiny, pustulate, the 

pustules subterminal, becoming coarsely sorediate 

or remaining as open pits; lower surface moderately to densely rhizinate, the rhizines moderately 

dichotomously branched. Apothecia not seen. 

C+ red, P- (atranorin, gyrophoric acid, and 

weakly UVS fluorescent unknown substances). 

CHEMISTRY.-cOrteX K+ yellow, medulla K-, 

DIsixIsucrIox.-Southeastern United States and 

southern Chile. 

HABITAT.-on trees (conifers, Nothofagus) in 

open woods at lower elevations in Chile, at 2000 

m in southeastern United States. 

REMARKs.-Externally this species is very Close 

to H. laevigata and H. thysanota. Specimens from 

Chile had been misidentified as H. densirhizinata. 

Chemistry places it closest to H. thysanota, which 

would react both C+ red (gyrophoric acid) and 

P+ red (echinocarpic acid). 

SPECIMENS EXAMINED.-chile: Tierra del Fuego, 

Santesson 6799 (S, US) . 

48. Hypotrachyna osorioi, new combination 

FIGURE 14f 

Parrnelia osorioi Hale, 1974133266. [Type collection: Abra de 

Cotto, Lavalleja, Uruguay, Osorio 6507 (MVM, holotype; 

US, isotype).] 

Thallus loosely adnate on rock, fragile, light 

tannish mineral gray, about 8 cm in diameter; 

lobes sublinear, about 2 mm wide; upper surface 

plane, continuous, coarsely isidiate, isidia simple 

and cylindrical to coralloid-branched, apically 

breaking down and almost pustulate or sorediate, 

dense with age; lower surface moderately 

rhizinate, rhizines moderately dichotomously 

branched. Apothecia rare, adnate, 3-7 mm in 

diameter; spores 6y X 12y. 

C+ red, P- (atranorin and gyrophoric acid). 

CHEMISTRY .-Cortex K f yellow, medulla K- , 

DISTRIBUTION.-UrugUay. 

HABITAT.-on stones in open forest at less than 

1000 m elevation. 

REMARKS.-The isidia of this rather fragile 

species are well developed and tend to enlarge 

apically and crumble, unlike any other in the 

genus. It may be related to H. pluriformis-H. 

revoluta, although the lobes are more regularly 

linear. 

SPECIMENS EXAMINED.-urUgUaY: Rocha, Hosseus 48 (H); San Jose, Osorio 6178 (MVM). 

49. Hypotrachyna osteoleuca, new combination 

FIGURE 15a 

Parrnelia osteoleuca Nylander, 1863:439. [Type collection: 
52 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Bogota, Colombia, Lindig 2528 (H, lectotype; BM, FH, G, 

H, M, P, IV, isolectotypes).] 

Thallus closely adnate or becoming loosely 

attached as thick thallus mats form, ashy white or 

darkening, 5-10 cm broad; lobes sublinear, often 

imbricate and crowded, 1-3 mm wide; upper surface plane, smooth, continuous, lacking soredia 

and isidia but usually heavily pycnidiate; medulla 

white to pale salmon; lower surface sparsely to 

densely rhizinate, the rhizines densely dichotomously branched. Apothecia common, adnate to 

substipitate, the disc dark brown, 1-4 mm in diameter; spores 5p-6p X 7p-9p. 

CHEMISTRY.-cOrteX K--, medulla K-, cf red, 

P - (lichexanthone, olivetoric acid, rhodophyscin 

(in lower medulla), and unidentified substances). 

DIsRTIBuTIoN.-Colombia and Venezuela. 

HABITAT.-on exposed rocks in the paramo at 

2300-3000 m elevation. 

REMARKS.-The positive C test was first noted 

by Vainio (1890), who also noted the superficial 

resemblance to H. brasilianu. In Venezuela this 

species occurs intermixed on boulders in the 

paramo with H. erythrodes, which reacts C-. 

SPECIMENS ExAhfINED.-colombia: Santander, 

Killip and Smith 15186 (US). Venezuela: Merida, 

Hale 42680, 42853, 42864, 43311, 43323, 43336. 

50. Hypotrachyna palmarum, new combination 

FIGURE l5b 

Parmelia palmarun2. Lynge, 1914: 136. [Tjpe collection: Buriti, 

Serra da Chapada, Mato Grosso, Brazil, Malnie 2243Ct (S, 

lectotype).] 

Parmelia zahlbruckneri Lynge, 1914: 125. [Type collection: 

Bocca da Serra, Seria da Chapada, Mato Grosso, Brazil, 

Malme S, lectotype).] 

Thallus adnate, turning cream or cinnamonbuff in the herbarium, rather fragile, 3-8 cm 

broad; lobes sublinear, crowded, more or less 

imbricate, short, 1-3 mm wide; upper surface 

shiny, plane to rugose and cracked on older lobes, 

isidia and soredia lacking; medulla white to very 

pale yellow; lower surface moderately rhizinate, 

the rhizines rather sparsely dichotomously 

branched. Apothecia common, adnate, the amphithecium crenate, the disc often split, 2-6 mm in 

diameter; spores 5y-ip X 9p-11p. 

CHE1\IISTRY.-cOrteX K-k yellow, medulla K-, 

C-, KCf rose, P- (atranorin, colensoinic acid, 

the “palmarum” unknown, and other unidentified substances). 

DIsTRIB1JTIoN.-Southeastern Brazil. 

HABITAT.-on trees or sandstone outcrops in 

open or secondary forests at 600-1500 m elevation. 

REMARKS.-This rather rare species is closely 

related to H. livida since they both contain 

colensoinic acid. Hopefully, the full relationship 

will be revealed when the molecular structure of 

the “palmarum” unknown is determined. The 

rock form, described by Lynge as Parmelia zahlbruckneri, is indistinguishable in chemistry and 

morphology from the corticolous forms. This parallels the situation with Hypotrachyna lividaParmelia osseoalbida. 

SPECIMEKS EXAhiINED.-BraZil: Mat0 Grosso, 

Malme in Lichenes Austroamericani 90 (H, UPS), 

Malme 2393**** (S, US), Richards 6355 (BM); 

Minas Gerais, Warming 279 (M); Santa Catarina, 

Reitr and Klein 16005 (US); SHo Paulo, Eiten 

and Eiten 3255, 3256 (US). 

51. Hypotrachyna partita, new species 

FIGURE 15c 

Thallus laxe adnatus, membraneus, cinereoalbicans, usque ad 15 cm latus, lobis sublinearibus, 

4-7 mm latis, superne isidiatis, mox lobulascentibus, lobulis 0.3 mm latis, 1 mm longis; cortex 

superior 16p-18p crassus, stratum gonidiale 22p24p crassum, medulla alba, 80p-9Oy crassa, cortex 

inferior 16p-18p crassus; subtus niger, dense 

rhizinosus. Apothecia ignota. 

Thallus loosely adnate, rather membranaceous 

and fragile, greenish mineral gray, to 15 cm 

broad; lobes sublinear to subirregular, the tips 

often broadly rotund, 4-7 mm wide; upper surface 

plane, shiny, initially isidiate, the isidia becoming 

strongly lobulate, the lobules dorsiventral, about 

0.3 mm wide and 1 mm long, simple to palmately 

divided, developing both laminally and marginally and becoming very dense in older parts; 

lower surface densely rhizinate, the rhizines 

densely dichotomously branched. Apothecia not 

seen. 

CHEmsmY.-Cortex K+ yellow, medulla K-, 

Cf red, P- (atranorin and anziaic acid rarely 

accompanied by perlatolic acid). 
NUMBER 25 

FIGURE 15.--Species of Hypotrachyna: a, H. osteoleuca (Hale 43332); 6, H. palmarum (Eiten 

3255); c, H. partita (Hale 43143); d, H. peruviana (Lechler 2727 in G); e, H. physcioides (Hale 

43354); f, H. physodalica (Mora 984). (Scale in mm.) 

53 
54 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TYPE CoLLEcTI0Ns.-Between La Aguada and 

La Montana, Merida, Venezuela, Hale 43143, 6 

February 1974 (US, holotype). 

DIsnIBuTIoN.-Central America and Venezuela. 

HABITAT.-on trunks of trees in open or cloud 

forest and on soil at road banks at 2200-3600 m 

elevation. 

REMARKs.-This species is common on stabilized roadsides where superficially morphologically 

similar H. bogotensis and H. ensifolia also occur. 

The lobulate isidia of H. partita are extremely 

variable and often densely developed. Perlatolic 

acid was detected in only one specimen (Culberson 12395) which has very large, almost palmately 

divided lobules and had been subjected to recent 

lava ash fall. 

Nee and Mori 3540 (US). Panama: Chiriqui, 

Hale 38930. Venezuela: Merida, Hale 42118, 

42144, 42852, 43065, 43167, 43191, 43205, 43242. 

SPECIMENS EXAMINED.-cOSta Rica: Cartago, 

52. Hypotrachyna peruviana, new combination 

FIGURE 15d 

Parmelia peruviana Nylander, in Crombie, 1876:228. [Type 

collection: Tatanara, Peru, Lechler 2727 (H, lectotype; 

BM, G, M, P, S, UPS, isolectotypes).] 

Thallus loosely adnate on bark, rather fragile, 

4-7 cm in diameter; lobes short, sublinear, 2-6 

mm wide; upper surface plane to rugulose, faintly 

maculate, lacking isidia and soredia; medulla pale 

chalcedony yellow; lower surface densely rhizinate, the rhizines 1-2 mm long, dichotomously 

branched. Apothecia conspicuously stipitate to 

columnar, 3-6 mm high, 2-5 mm in diameter, the 

amphithecium rugose; spores 7p-12p X 11p-18p. 

yellowish, C + orange (atranorin, barbatic acid, 

obtusatic acid, entothein, and unidentified 

pigments). 

CHEMISTRY.-cOrteX K+ yellow, medulla K+ 

DIsmIBuTIox.-Peru and Brazil. 

HABITAT.-On trees in open forests at 1000-1500 

m elevation. 

REhrARKs.-The coarsely stalked apothecia are 

distinctive for this species. The yellow pigment 

is very pale and probably different from that in 

H. endochlora. The species is related to H. physcioides, as Nylander (1885) suspected, but has 

diverged considerably from any presumptive progenitor in this group because of the unusual 

apothecia and production of pigments. 

SPECIMENS EXAMINED.-BraZil: Rio de Janeiro, 

Gregory and Ainsworth 529, 530 (BM, US); 

Santa Catarina: Reitz and Klein 16091 (US). 

53. Hypotrachyna physcioides, new combination 

FIGURE 15e 

Parmelia Physcioides Nylander, 1860:385. [Type collection: 

Silla de Caracas, Venezuela, Humboldt and Bonpland (P, 

lectotype; BM, H, isolectotypes).] 

Parmelia laevigata var. gracilis Muller Argau, 1879: 169. [Type 

collection: Bogota, Colombia, Andrt 923 bis (G, lectotype; 

BM, US, isolectotypes),] 

Parmelia boliviana Nylander, 1885:612. [Type collection: 

Prov. Larecaja and Caupolican, Bolivia, Weddell (P. lectotype).] 

Parmelia gracilis (Muller Argau) Vainio, 1890:55. 

Parmelia digitata Lynge, 1914:98. [Type collection: Santa 

Anna da Chapada, Mato Grosso, Brazil, Malme 2545 (S, 

lectotype; LD, US, W, isolectotypes).] 

Parmelia confusula Zahlbruckner, 1929: 162. [Type collection: 

Based on P. gracilis (Muller Argau) Vainio, not P. gracilis 

Sprengel, 1827:277, or P. gracilis Muller Argau, 1887:317).] 

Thallus adnate to loosely attached, usually 

rather coriaceous, 6-15 cm broad, ashy white to 

tannish mineral gray; lobes sublinear, separate to 

crowded, 2-6 mm wide; upper surface plane to 

convex, dull and sometimes white pruinose at the 

tips to shiny and distinctly white-maculate, rarely 

becoming sparsely to densely lobulate-laciniate 

toward the center, isidia and soredia lacking; 

lower surface densely rhizinate, the rhizines 

densely dichotomously branched. Apothecia adnate to substipitate, 2-10 mm in diameter; spores 

6p-9 pX 12p-16p. 

CHEMISTRY.-cOrteX K + yellow, medulla K-, 

C- or C+ orange, KC+ orange, P- (atranorin, 

barbatic acid, 4-0-demethylbarbatic acid, obtusatic 

acid, and norobtusatic acid). 

DIsnIBuTIoN.-Mexico, Central America, West 

Indies, Colombia, Venezuela, Peru, Ecuador, Bolivia, Brazil, and Southeast Asia. 

HABITAT.-on trees (conifers and hardwoods) 

and rocks in open or disturbed forests at 1500- 

4100 m elevation. 

REMARKs.-Hypotrachyna physcioides is common in the northern part of its range but apparently lacking in most of Brazil and southward. 

It is extremely variable, Typical soil- or rock- 
NUMBER 25 55 

inhabiting specimens (as in the type) are almost 

coriaceous and have dull linear lobes. Some 

corticolous forms (as in the type of Parmelia 

boliviana) are thinner and have shorter, sublinear lobes and a shiny surface with white 

maculae. Many intermediates can be found. The 

chemistry, however, is uniform in all cases. 

A rather densely lobulate population described 

by Lynge as Parmelia digitata probably is only an 

extreme form which intergrades with the more 

normal and abundant nonlobulate population. 

This species is the presumptive progenitor of 

various vegetative morphs, H. laevigata (sorediate) and H. imbricatula (isidiate), as well as 

being part of the same constellation that produced 

H. chlorina, H. endochlora, H. isidiocera, H. 

peruviana, etc. The possible origin of H. physcioides from hybridization between evernic-acidcontaining H. pulvinata and a barbatic-acidcontaining progenitor similar to H. massartii 

(Hue) Hale is discussed by Culberson and Hale 

(1973). 

SPECIMENS EXAMINED.-Mexico: Puebla, Arsene 4253 (US). Guatemala: Alta Verapaz, Standley 91328 (US). Costa Rica: Alajuela, Standley 

and Torres 47720 (US); Cartago, Culberson 12394 

(DUKE, US), Stork L-38 (US); San Jose, Culberson 13215 (DUKE). Panama: Chiriqui, Martin 

2751 (US). Haiti: Ouest, Imshaug 22990, 22991 

(MSC), Wetmore 3121 (RISC); Sud, Imshaug 

23238 (MSC). Dominican Republic: La Vega, 

Zmshazig 23459, 23628 (MSC). Colombia: Cundinamarca, King 472 (US), Nee and Mori (US), 

Soderstl-om 1305b, 1305c (US), Weir 27 (BM, 

US); Santander, Killip and Smith 15644, 18593 

(US). Venezuela: Merida, Hale 42171, 42433, 

42485, 42804, 43126, 43224, 43326, Hanselmann 

168 (DUKE, US), Magdefrau 698 (h4, US), Nash 

2013, 2014 (US). 

54. Hypotrachyna physodalica, new combination 

FIGURE 15f 

Parmelia physodalica Hale, 1971d:436. [Type collection: 

Guasca, Cundinamarca, Colombia, Perez-Arbelaez 1104 (US, 

holotype).] 

Thallus loosely adnate, often scattered, coriaceous but easily breaking apart, sea-foam yellow, to 

8 cm broad; lobes sublinear, elongate, sparsely 

dichotomously branched, separate, 1-3 mm wide; 

upper surface plane, shiny, maculae, soredia, and 

isidia lacking; lower surface moderately to densely 

rhizinate, the rhizines dichotomously branched. 

Apothecia not seen. 

P+ orange-red (usnic acid, physodalic acid, and 

a trace of protocetraric acid or usnic acid and 

protocetraric acid only). 

DIsTRIsuTIoN.-Colombia, Venezuela, and Peru. 

HABITAT.-on soil and over humus in the paramo at 3700-4100 m elevation. 

REMARKS.-This paramo species is easily recognized by the deep greenish yellow color and sublinear divaricate lobes. Both H. caraccensis, a 

much larger species, and H. Eopezii, which has a 

nearly bare lower surface, react K+ red or yellow. 

The type of H. physodalica was reported to contain only physodalic acid, but subsequent TLC 

tests showed traces of protocetraric acid. Specimens collected later contain only protocetraric 

acid. I am provisionally regarding these chemically variable populations as a single species, at 

least until we know more about the biogenesis and 

co-occurrence of these closely related acids. 

SPECIMENS ExAhfINED.-Colombia: Cundinamarca, Mora 984 (US); Santander, Killip and 

Smith (17527 (US). Venezuela: Merida, Hanselmann 340 (DUKE, US), Oberwinkler and Poelt 

(M, US). Peru: San Martin, Holligan L60A 

CHEMISTRY.-cOrteX K--, medulla K--, c-, 

(BM) * 

55. Hypotrachyna pluriformis, new combination 

FIGURE 16a 

Parmelia pluriformis Nylander, 1860:381. [Type collection: 

Brazil, Weddell (H, lectotype; P, isolectotype).] 

Parmelia pluriformis var. chlorocarpa Muller Argau, 1881:85. 

[Type collection: Apiahy, Brazil, Puiggari 125e (G, lectoParmelia pluriformis var. multifida Muller Argau, 1881:85. 

[Type collection: Apiahy, Brazil, Puiggari 1283 (G, lectoParmelia everniaeformis Zahlbruckner, 1902:416. [Type collection: Petropolis, Brazil, Hohnel 163 (W, lectotype).] 

Parmelia subpluriformis Zahlbruckner, 1909: 172. [Type c01- 

lection: Mt. Jaragui, Sao Paulo, Brazil, Schiflner (W, lectotype).] 

type).] 

type).] 

Thallus adnate or loosely attached to bark, 

greenish mineral gray, 5-8 cm broad; lobes sublinear to subirregular, imbricate, sometimes be- 
56 ShlITHSONIAN CONTRIBUTIONS TO BOTANY 

coming canaliculate, 1-5 mm wide; upper surface 

plane to convex, continuous, lacking soredia and 

isidia; lower surface black in the center but with 

a broad brown zone along the margins, moderately 

rhizinate. Apothecia common, substipitate, 2-10 

mm in diameter, the amphithecium rugulose, the 

disc often radially split; spores Sp-11~ X 14p-25p, 

C+ red, P- (atranorin and gyrophoric acid). 

DIsTRIBUTION.-~~exico, Brazil, Uruguay, and 

Argentina. 

HAnITAT.-On trees in open forest at 1000-2300 

m elevation. 

REhIARKS.-The lobes of this species are unique 

in being subirregular and in part canaliculate and 

revolute. The thallus has a coriaceous texture. 

Spore size is variable but definitely larger than 

in most Hypotrachynae. Zahlbruckner (1909), in 

fact, recognized Parnzelia subpluriformis on the 

basis of large, thick-walled spores, but I would 

consider this as merely the extreme on a broad 

scale of spore size. It has the same chemistry as 

H. reuoluta and H. osoiioi, two vegetative morphs 

that originated from a sexual morph similar to 

H. pluriformis. 

SPECIMENS ExAwNm.-Alexico: Puebla, Hale 

19621. Brazil: hIinas Gerais, Burclzell 1105/68, 

2252 (BRI), Christina (P), Vainio 794 (Rl, TUR), 

1049, 1052 (BRI, TUR); Rio de Janeiro, Glazioir 

1832 (UPS); Sao Paulo, Gehit '(US, W), Lindberg (UPS), Schiflnei (BRI, G). Uruguaj: Canelones, Row-Mato 1031 (S); Florida, Imshaug 

424064 (RISC). Argentina: RIisiones, ilfontes 

10073 (AfVM). 

CHEILIISTRY.-cOrteX K+ yellow, medulla K- , 

56. Hypotrachyna producta, new species 

FIGURE 16b 

Thallus laxe adnatus, 6-7 cm diametro, 

cinereo-albicans, lobis sublinearibus, 2-3 latis, 

superne sorediatis; cortex superior 13y-15p crassus, stratum gonidiale 14u-16~ crassum, medulla 

alba, 65y-9Op crassa, cortex inferior, 12p crassus; 

subtus niger, modice rhizinosus. Apothecia ignota. 

Thallus loosely adnate on bark, whitish mineral gray, 6-10 cm in diameter; lobes sublinear, 

becoming elongate and subdivaticate, 2-4 mm 

wide; upper surface shiny, plane, sorediate, the 

soredia subterminal in orbicular soralia, the sorediate lobes becoming subrevolute; lower surface moderately rhizinate, the rhizines moderately to densely 

dichotomously branched. Apothecia not seen. 

CHEwsmY.-Cortex K+ yellow, medulla K-, 

C+ red, P- (atranorin and anziaic acid). 

TYPE CoLLEcTIoiv.-On Picea rubens, Mt. LeConte, Sevier County, Tennessee, Nakanishi 174, 

8 hlay 1971 (US, holotype; KOBE, isotype). 

lombia, and Africa. 

HABITAT.-On trunks and branches of trees 

(hardwoods and conifers) in open forest at 

2000-3600 m. 

REhrARKs.-Hypotrachyna producta is morphologically similar to other sorediate species such 

as H. laevigata and H. oostingii. It would have to 

be identified with a chemical test. The presumptive progenitor is H. ducalis, which is also rarely 

collected in tropical America. 

SPECIMENS ExAxfINED.-Colombia: Cundinamarca, Killip 341 18 (US); Santander, Killip and 

Smith 17798 (US). 

DISTRIBUTIoN.-SOutheastern United States, Co57. Hypotrachyna prolongata, new combination 

FIGURE 16c 

Pnimelici fiiolongntn Kurokai+a in Hale and Kuiokaira, 1964: 

182. [Tjpe collection: Pic Nacaja, Sud, Haiti, Wetmore 

3233 (RISC, holotjpe; US, isoqpe).] 

Thallus adnate to looselj adnate, mineral gray, 

5-10 cm in diameter; lobes sublinear, 1.5-4 mm 

wide; upper surface plane, shiny, faintly maculate, spaisely lobulate-isidiate along the margins, 

the "isidia" in part lobulate, sometimes branched, 

2-3 mm long; lower surface densely rhizinate. 

.\pothecia not seen. 

C+ red, P- (atranorin and an unidentified C+ 

substance). 

~H~h11SlR~.-~01 tex K+ yellow, medulla K--, 

DISTRIBUTIOA .-Haiti. 

HABITAT.-OII pine trees at about 2000 m 

eleLation. 

RE\IARKS.-ThiS rare species is still known only 

fiorn the tj pe collection in Haiti. Specimens from 

other localities so identified on the basis of a Cf 

1 etl reaction T$ ere found to contain anziaic acid 

and to ha\e a different kind of isidial formation. 

These have been recognized as H. rachista (Hale, 

197ld). The C+ substance in H. p?-olongata 
NUMBER 25 

FIGURE lG.-Species of Hypotrachyna: a, H. plurifortnis (Schiflner in IV); b, H. producta (Sakanishi 174); c. H. prolongata (Wetmore 3233); d, H. proterita (Hale 43365); e, H. protoboliviana 

(Flenniken 1874); f, H. pseudosinuosa (Hale 35450). (Scale in mm.) 

57 
58 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

does not compare with anziaic acid, gyrophoric 

acid, lecanoric acid, or olivetoric acid and probably represents a new compound. 

58. Hypotrachyna protenta, new species 

FIGURE 16d 

Thallus laxe adnatus, saxicola, 6-10 cm diametro, lobis sublinearibus, imbricatis, 2-4 mm 

latis; cortex superior 141.1-16p crassus, stratum 

gonidiale 24p crassum, medulla alba, 8Op-9Op 

crassa, rare pro parte aurantiaca, cortex inferior 

14p-16p crassus; subtus niger, modice rhizinosus. 

Apothecia numerosa, substipitata, sporis simplicibus, 6p X 12p. 

Thallus rather loosely adnate on rocks, the colonies easily breaking apart, whitish ashy, 6-10 cm 

broad; lobes sublinear, more or less crowded or 

imbricate, variable, 2-4 mm wide, sometimes becoming densely appressed lobulate toward older 

parts; upper surface shiny, plane to minutely rugulose, often heavily pycnidiate, soredia and isidia 

lacking; lower surface densely rhizinate, the 

rhizines sparsely to moderately dichotomously 

branched, sometimes projecting as a narrow mat 

along the margins. Apothecia numerous, adnate, 

4-8 mm in diameter; spores 6p X 12p. 

CHEMIsmY.-Cortex K+ yellow, medulla K-, 

C-, KC+ red, P- (atranorin, alectoronic acid, 

a-collatolic acid, and, if pigmented, rhodophyscin). 

TYPE CoLLEcTIoiv.-Open forest along stream, 

on rocks, El Valle, Merida, Venezuela, Hale 

43365, February 1974 (US, holotype; DUKE, TNS, 

UPS, isotypes). 

DIsTRIBuTIoN.-Centra1 America, Colombia, and 

Venezuela southward to Peru, and Brazil. 

HABITAT.-on rocks in open forests and in the 

paramo at 1700-4000 m elevation. 

RExfARKs.-Hypotrachyna protenta grows on 

open boulders or even soil in paramo and subparamo regions. Typically the lobes are densely 

imbricate and black rimmed but under some conditions they are almost separate and divaricate. 

The species resembles H. chicitae but can be distinguished by the negative C reaction and white 

fluorescence under ultraviolet. It can also be separated from small, high-elevation forms of H. 

physcioides (C+ orange) by a color test. 

SPECIMENS ExAMINED.-costa Rica: Stork (DUKE, 

H, MICH, US, W). Panama: Chiriqui, Hale 

38796, Scholander (US). Colombia: Cundinamarca, King 505 (US). Venezuela: Merida, Hale 

42130a, 42193, 42915, 42929, 43030, 43278, 43288, 

43339. Peru: Cuzco, Staflord 1008 (F, US). Brazil: 

Santa Catarina, Reitz and Klein 17545 (US); SZo 

Paulo, Schiflner (BM, W). 

59. Hypotrachyna protoboliviana, new combination 

FIGURE 16e 

Parmelia protoboliviana Hale, 1974b:267. [Type collection: 

Volcan Irazu, Cartago, Costa Rica, Flenniken 1874 (US).] 

Thallus adnate on bark, rather fragile, about 8 

cm in diameter, light yellowish green; lobes more 

or less sublinear, short, 4-5 mm wide; upper surface smooth, without isidia or soredia; lower surface densely rhizinate, the rhizines densely 

dichotomously branched. Apothecia poorly developed, about 1 mm in diameter; spores lacking. 

orange, P- (usnic acid, barbatic acid, 4-0-demethylbarbatic acid, obtusatic acid, and norobtusatic acid). 

CHEMISTRY.-cOrteX K-, medulla K--, c+ 

DISTRIBUTION.-cOSta Rica. 

HABITAT.-oII trees in open forest at about 

3200 m elevation. 

REMARKs.-Identical to H. physcioides except 

for the replacement of atranorin by usnic acid, 

H. protoboliuiana is known only from Costa Rica 

in an active volcanic region. As a very recently 

evolved species it has not given rise to any vegetative morphs comparable to those of H. physcioides. 

Flenniken 2358 (US). 

SPECIXIEXS EXAhZINED.-COSta Rica: Cartago, 

60. Hypotrachyna pseudosinuosa, new combination 

FIGURE 16f 

Parmelia pseudosinuosa Asahina, 1951:329. [Type collection: 

Shimohirano-mura, Shinano, Japan, Takahashi 99 (TNS, 

lectotype).] 

Parmelia anaptychioides Kurokawa in Hale and Kurokawa, 

1964: 165. [Type collection: Pic0 Trujillo, Dominican Republic, Wetmore 3617 (hlSC, holotype; US, isotype).] 

Thallus closely adnate, light mineral gray, 3-9 

cm in diameter; lobes sublinear, short and sometimes crowded, 1-4 mm wide; upper surface 
NUMBER 25 59 

plane, continuous, sorediate, the soralia capitate, 

mainly subterminal; lower surface moderately 

rhizinate. Apothecia rare, adnate, 2-4 mm in 

diameter; spores 6p-8y X 8p-lop. 

CHEiwsmY.-Cortex K+ yellow, medulla K-, 

P + orange red (atranorin, protocetraric acid). 

South Africa, Japan, and Southeast Asia. 

HABnAT.-on tree trunks and small branches 

(hardwoods, Pinus) in secondary forests at 1800- 

2200 m elevation. 

REMARKS.-The thallus of H. pseudosinuosa is 

often small and closely adnate, as in the type of 

Parmelia anaptychioides (Hale, 1971a). It is one 

of the few species of Hypotrachya first described 

from Asia and later found to be pantropical. 

There is no presumptive parent species. 

SPECIMENS EXAMINED.-Mexico: Oaxaca, Nakanishi 285 (US). Dominica: Hale 35358, 35488. 

Chile: Valparaiso, Imshaug 36628, 36642 (MSC). 

Records from Jamaica and Haiti are listed in 

Hale and Kurokawa (1964: 166). 

DISTRIBUTION.-Mexico, West Indies, Peru, 

61. Hypotrachyna puluinnta, new combination 

FIGURE 17a 

Parrnelia pulvinata Fee, 1824: 123. [Type collection: Tropical 

America (G, lectotype).] 

Parmelia subsinuosa Nylander, 1885:613. [Type collection: 

Tropical America, BonPland (H, Nylander herbarium 

number 35169, lectotype; P, isolectotype).] 

P. caraccensis f. pulvinata (Fee) Zahlbruckner, 1929:58. 

Thallus adnate to loosely attached, whitish mineral gray, 8-12 cm broad; lobes short and often 

crowded, sublinear, 2-6 mm wide; upper surface 

plane, shiny, white-maculate; lower surface 

densely rhizinate, the rhizines often projecting as 

a mat beyond the margins, densely dichotomously 

branched. Apothecia common, adnate, 5-17 mm 

in diameter, the amphithecium strongly whitemaculate; spores 6p-7p X lop-13p. 

C+ red, KC+ red, P- (atranorin, evernic acid, 

and lecanoric acid. 

DISTRIBUTION.-Southwestern United States, 

Mexico, Central America, Jamaica, Andean region from Venezuela to Bolivia, and southeastern 

Brazil. 

HABITAT.-on trees (oak, pine), soil, over 

CHEMISTRY.-cOrteX K+ yellow, medulla K--, 

mosses in pine forests, scrubby mist forest, and 

paramo at 1200-4300 m elevation. 

recognized by the white maculae, dense rhizine 

mat, and C+ rose or red color test. It is very common in the dry upland oak-pine forests from 

southern New Mexico-Arizona southward to Guatemala but becomes quite rare in South America. 

The vegetative morphs, H. rockii (sorediate) and 

H. bogotensis (isidiate), are two of the commonest 

species in tropical America. It is H. pulvinata or a 

progenitor similar to it which probably crossed 

with a barbatic-acid-containing progenitor such 

as H. massartii (Hue) Hale to give rise to the 

obtusatic-acid-containing constellation including 

H. physcioides and its vegetative morphs, H. 

peruviana, etc. (Culberson and Hale, 1973). 

SPECIMENS EXAMINED.-Mexico: Chiapas, Hale 

20330, 20571; Federal, Kramer 1817 (US); Mexico, Hale 19290; Oaxaca, Hale 20672, 20688, 20691, 

20700, 20784; Sinaloa, Culberson 13356 (DUKE). 

Guatemala: Quetzaltenango, Steyermark 34898 

(US); Totonicapan, Standley 84507 (US). Costa 

Rica: San Jose, Standley and Valerio 43750 (US). 

Panama: Chiriqui, Scholander (US). Jamaica: 

Imshaug 15495 (MSC), Plitt (US). Colombia: 

Magdalena, Flenniken 2295 (US); Santander, 

Killip and Smith 19560 (US); Tolima, Killip and 

Varela 34678a (US). Venezuela: Merida, Dennis 

1940 (BM), Hale 42365, 42374, 42460. Ecuador: 

Pichincha, Jameson 145 (BM), Ugent (US, WIS). 

Peru: Bryan 579 (US, WIS). Bolivia: Cochabamba, Ugent (US, WIS); Larecaja, Mandon 

1753 (BM, S). Brazil: Minas Gerais, Henschen 

REMARKS.-Hypotrachyna pu~vinata is easily 

(UPS). 

62. Hypotrachyna rachista, new combination 

FIGURE 17b 

Parrnelia rachista Hale, 1971d3436. [Type collection: Clingmans Dome, Swain County, North Carolina, Kurokawa 6755 

(US; isotype in TNS).] 

Parmelia lobulijera var. sanguineoreagens Degelius, 1941:63. 

[Type collection: Mt. Kephert, North Carolina, Degelius 

(US, lectotype).] 

Thallus adnate to loosely adnate, 6-12 cm 

broad, whitish mineral gray; lobes linear, dichotomously branched, variable, 1-5 mm wide; upper 

surface plane, continuous, becoming dactyloidisidiate at the lobe tips, isidia dorsiventral, often 
60 SMITHSONIAX COXTRIBUTIOKS TO BOTANY 

dissected; lower surface black but sometimes with 

a narrow white rim below the lobe tips, densely 

rhizinate, the rhizines densely dichotomously 

branched. Apothecia not seen. 

C+ red, P- (atranorin and anziaic acid). 

DIsTRIsuTIoN.-Southeastern United States, MeXico, Panama, Dominican Republic, Colombia, 

Venezuela, and Peru. 

HABrrAT.-On trees (hardwoods and conifers), 

over mosses, and on soil at road banks at 2000- 

3500 m elevation. 

REMARKS.-The isidia of this species are unusual in branching and position on the thallus 

(Figure 4d) and one might question whether they 

should be called isidia or lobules. The distinctive 

chemistry relates H. rachista to H. ducalis and its 

sorediate morph H. producta. The species is most 

commonly collected in the southern Appalachian 

Mountains in the United States and seems rare in 

tropical America. 

SPECIMENS ExAMINED.-Dominican Republic: 

La Vega, Imshaug 23402 (MSC). Venezuela: 

Merida, Hale 42610, 42648, 42896, 43139, 43239. 

Peru: Pasco, Tovar 397 (BM). For additional records from Mexico, Panama, and Colombia see 

Hale (1971:437). 

CHEMISTRY.-cOrteX K+ yellow, medulla K--, 

63. Hypotrachyna reducens, new combination 

FIGURE 17c 

Parrnelin reducens Nylander, 1963:438. [Type collection: Colombia, Lindig 2743 (H, Nylander herbarium number 

35100, lectotype).] 

Thallus adnate to loosely attached, sea-foam 

yellow, 2-5 cm in diameter; lobes sublinear, separate, 1-2 mm wide; upper surface plane to 

rugulose, faintly white-maculate, lacking soredia 

and isidia; lower surface densely rhizinate, the 

rhizines sometimes projecting out as a mat along 

the margins. Apothecia common, substipitate, the 

disc becoming radially split, 1-5 mm in diameter; 

spores 5p-7p X 9p-llp. 

CHEmsmY.-Cortex K-, medulla K+ yellow 

turning red, C-, P+ orange-red (usnic acid, norstictic acid, salazinic acid, sometimes a trace of 

a tranorin). 

DIsmIBuTIoN.-Panama, Colombia, Peru, Sabah, 

and New Guinea. 

HABITAT.-On trunk and twigs of hardwood 

trees in exposed areas at 2800-3500 m elevation. 

REhIARKs.-while rarely collected, H. reducens 

is easily identified since it is the only nonsorediate, 

nonisidiate yellow species (outside of rare H. protobohiana) on trees. It is always fertile. The typical habitat seems to be exposed shrubs in the 

paramo. The presumed sorediate morph, H. sinuosa, may occur in similar habitats. 

SPECIMENS ExAMINED.-Panama: Chiriqui, Scholander (US). Colombia: Humbry-Tracy 313 

(BM); Lindig 122 (BM, M), 799 (M, P, syntypes 

of P. reducens). Peru: Recuay, Gomez 549 (US). 

64. Hypotrachyna revoluta, new combination 

FIGURE 17d 

Parinelia revoluta Floerke, 1815:ll. [Type collection: Germany, Floerke (FH-Tuck, lectotype).] 

Thallus loosely adnate, rather fragile, greenish mineral gray, 3-8 cm broad; lobes sublinear 

to subirregular, often short and more or less ascending at the tips, subcanaliculate, 1-5 mm wide; 

upper surface plane, shiny, becoming very 

coarsely pustulate toward the lobe tips, the pustules becoming coarsely sorediate and sometimes 

eroding centrally leaving bare blackish areas of 

medulla, heavily sorediate lobes becoming revolute; lower surface black but with a brownish zone 

along the margins, sparsely rhizinate, the rhizines 

sparsely dichotomously branched. Apothecia very 

rare, adnate to substipitate, 2-6 mm in diameter; 

spores 6p-8~ X 10p-12p. 

CHEmsmY.-Cortex K+ yellow, medulla K-, 

C+ red, P- (atranorin and gyrophoric acid). 

DIsTRIBuTIoN.-Eastern United States, West Indies, Venezuela to Chile in the Andes, Europe, 

South Africa, Japan, and Indonesia. 

HABITAT.-On trees (hardwoods and conifers) 

in open or cloud forest and on rocks at 1000-4500 

m elevation (at sea level in Chile). 

REXIARKs.-This classical European species 

(Almborn, 1942) has a montane pantropical distribution. The name, however, has been applied 

incorrectly to many different species in the tropics. 

There is a wide and often perplexing range of 

variation in soredial formation, but in general the 

soredia are lamina1 to subterminal, causing the 

lobes to become broadly revolute, and may even 
SUMBER 25 

FIGURE 17.-Species of Hy$otmchyria: a, H. pulvit7ata (Hale 20784): 0, H. rochisfa (Hole 43139): 

c, H. reducens (Liridig 122 in &I); d, H. reuolnfa (Male 43293a); e, H. r/inbdifor?riis (Il.iirtfack 

483); f, H. rockii (Rock 7). (Scale in mm.) 

61 
62 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

erode away leaving large, bare pustule-like areas 

(Figure 2a). The lobes also vary widely in width 

and degree of erectness. Spore size for European 

material was determined by Hillmann (1936) to 

be up to 16p long, somewhat larger than American material. A presumptive progenitor, H. pluriformis, has significantly larger spores, up to 251.1 

long. Vainio’s determination of his number 1140 

as Parmelia revoluta from Brazil is incorrect; it 

is H. rockii. 

SPECIMENS EXAMINED.-Dominican Republic: 

La Vega, Allard 16530A (US). Venezuela: Merida, 

Hale 42995, 43296, 43306. Bolivia: Brooke 6819 

(US), Overbeek 5455 (US). Chile: Chiloe, Santesson 4029 (S, US), 4031 (S); Valdivia, Rabenhorst (M, US), Santesson 2363 (S). 

65. Hypotrachyna rhahdiformis, new combination 

FIGURE 17e 

Parmelia rhabdijormis Kurokawa in Hale and Kurokawa, 

1964: 183. [Type collection: El Boquete, Chiriqui, Panama, 

Maxon 5128a (US, holotype).] 

Thallus adnate, light tannish mineral gray, 

5-8 cm broad; lobes sublinear, short, imbricate, 

2-6 mm wide; upper surface plane, continuous, 

moderately isidiate, the isidia mostly simple and 

cylindrical, to 0.5 mm high or more, becoming 

club-shaped or lobulate; lower surface densely 

rhizinate, the rhizines densely dichotomously 

branched. Apothecia adnate, 1-5 mm in diameter, 

the amphithecium isidiate; spores 6p-8p X 

1 5p-2 1 p. 

CHEMISTRY.-COrteX K + yellow, medulla K + 

yellow turning red, C-, P+ orange (atranorin, 

norstictic acid, and in Iltis 3005 and Wurdack 

483 a trace of stictic acid). 

Asia. 

HABITAT.-oII trees or on soil in cloud forest at 

1300-2300 m elevation. 

REMARKs.-This isidiate species has a unique 

chemistry. Superficially it resembles H. costaricensis, which has very dense rhizines and reacts 

K-. The collections from Peru are more robust 

than the type but agree in other features. 

SPECIMENS EXAM INED.-Peru: Chachapoyas, 

Wurdack 483 (US); Urubamba, Zltis 3005 (US, 

DISTRIBUTION.-Panama and Peru, Southeast 

WIS). 

66. Hypotrachyna rockii, new combination 

FIGURE 175 

Partnelia rockii Zahlbruckner, 1912:379. [Type collection: 

Kauai, Hawaii, Rock 7 (W, lectotype; US, isotype).] 

Thallus adnate to loosely attached, 5-12 cm in 

diameter, whitish mineral gray; lobes sublinear, 

1-6 mm wide; upper surface plane, shiny, often 

distinctly white-maculate, pustulate initially near 

the lobe tips, usually becoming coarsely sorediate; 

medulla white except for pale orange spots under 

the soralia; lower surface black, densely rhizinate. 

Apothecia rarely developed, adnate, 2-8 mm in 

diameter; spores 5p-7p X 11p-14p. 

CHEMISTRY.-COrteX K + yellow, medulla K - , 

C+ rose, P- (atranorin, evernic acid, and lecanoric acid). 

DISTRIBUTION.-SOutheaStern United States to 

southern Chile, South Africa, Southeast Asia, and 

Hawaii. 

HABITAT.-on tree bark (oaks, pine, Ficus, 

Drimys) and rocks in open forests at 1000-3300 m 

elevation. 

REMARKs.-Hypotrachyna rockii is a very common lichen in the highland oak-pine forests of 

Mexico and Central America as well as in the 

West Indies. It is very similar externally to H. 

laevigata in having distinct maculae and subterminal soredia but can be distinguished from it by 

a color test with C. The presumptive progenitor 

is H. pulvinata (Culberson and Hale, 1973). 

REPRESENTATIVE SPECIMENS EXAMINED.-hfexico: 

Chiapas, Hale 202 14, 20242, 20290, 20534, 2056 1 ; 

Michoaah, Arsene 3746 (US), Hale 20850, 20987; 

Puebla, Hale 21048; Veracruz, Hale 21116, 21176. 

Guatemala: Alta Verapaz, Standley 69331 (US); 

Baja Verapaz, Hale 38333; Chimaltenango, Standley 60097 (US); Huehuetenango, Standley (US); 

Totonicapan, Standley 84522 (F). Costa Rica: 

San Jose, Standley and Valerio 43898 (US). Panama: Chiriqui, Hale 38768, 38819, 38863, Scholander (US). Cuba: Oriente, Zmshaug 25045 

(MSC, US). Jamaica: Zmshaug 12954, 13802, 

15424, 15541 (MSC, US), Orcutt 5626c (US), 

Plitt (US). Haiti: Ouest, Zmshaug 22591 (MSC), 

22538 (US), Wetmore 2973, (MSC); Sud, Wetmore 3365 (MSC). Dominican Republic: Cordillera Central, Zmshaug 23537 (MSC, US), 23639 

(MSC); La Vega, Allard 16529a (US). Dominica: 
NUMBER 25 63 

Hale 35586. Martinique: Culberson 14555 

(DUKE). Colombia: Santander, Charetier 202 

(US). Venezuela: Federal, Vareschi 427c (US); 

Merida, Hale 42553, 42734, 42974, 43287. Peru: 

Cuzco, Zltis 3004 (US, WIS), Thomasson (UPS). 

Chile: Magallanes, Santesson 8010 (S, US); T' ierra 

del Fuego, Santesson 6793 (S, US); Valdivia, 

Mahu 1695 (US). Brazil: Minas Gerais, Vainio 1140 

(BW 

67. Hypotrachyna silvatica, new combination 

FIGURE 18a 

Parmelia silvatica Lynge, 1914: 118. [Type collection: Santa 

Anna da Chapada, Mato Grosso, Brazil, Malme 2393' (S, 

lectotype).] 

Parmelia cruslacea Lynge, 1914: 108. [Type collection: Santa 

Anna da Chapada, Mato Grosso, Brazil, Malme (S, lectoParmelia silvatica var. pinnata Lynge, 1914: 120. [Type collection: Santa Anna da Chapada, Mato Grosso, Brazil, 

Malme 2393*** (S, lectotype).] 

Parmelia silvatica var. radiata Lynge, 1914: 120. [Type collection: Santa Anna da Chapada, Mato Grosso, Brazil, Mahe 

2393** (S, lectotype).] 

Thallus closely adnate on bark, whitish mineral 

gray, 2-6 cm in diameter; lobes sublinear, often 

crowded, 0.5-2 mm wide; upper surface plane, 

shiny, faintly maculate; medulla in the lower half 

antimony yellow; lower surface moderately rhizinate, the rhizines densely dichotomously branched. 

Apothecia common, adnate, 1-2 mm in diameter; 

spores 5p6p X lop-12y. 

P + orange-red (lichexanthone, protocetraric acid, 

and an unidentified anthraquinone). 

type).] 

CHEMISTRY.-cOrteX K--, medulla K--, C-, 

DIsTRIsuTIorv.-Southern Brazil. 

HABITAT.-On trees in open forests. 

REMARKS.-This rare lichen resembles H. livida 

externally but the chemistry is completely different. Some specimens are very closely adnate and 

have narrow lobes. These were described by Lynge 

under the synonyms listed above. 

SPECIMENS ExAMINm.-Brad: h4ato Grosso, 

Malme (S, UPS) in Lichenes Austroamericani 

87 (G, H, LD, UPS), 2271 Ba (S), 2511 Ba (S). 

68. Hypotrachyna singularis, new combination 

FIGURE 18b 

Parnzelia singularis Hale, 1974b:267. [Type collection: Cerros 

Calla Calla, Chachapoyas, Peru, Hutchinson and Wright 

5704 (US, holotype; UC, isotype).] 

Thallus loosely adnate on branches, subcoriaceous, whitish to pale yellowish mineral gray, 

about 8 cm broad; lobes subirregular to sublinear, 

3-5 mm wide, marginally dissected and lobulate, 

the lobules to 1 mm long, narrow; upper surface 

smooth and shiny, continuous; lower surface 

densely rhizinate, the rhizines densely dichotomously branched. Apothecia numerous, adnate, 

to 5 mm in diameter; spores 6p7p X 10p-12p. 

CmMIsmY.-Cortex K+ yellow, medulla K-, 

C-, P- (atranorin and apparently a fatty substance not further identified). 

DISTRIBUTION.-PerU. 

HABITAT.-oII tree in open forest at 3100 m 

elevation. 

RE~~ARKs.-HyPotrachyna singularis is characterized by the rather coriaceous thallus and numerous elongate marginal lobules. It is known 

only from the type collection. 

69. Hypotrachyna sinuosa, new combination 

FIGURE 18c 

Lichen sinuosus Smith, 1809:2050. [Type collection: DumParmelia sinuosa (Smith) Acharius: 1814:207. 

Parmelia despreauxii Delise ex Duby, 1830:602. [Type collection: Vire, France, Despreaux and Briquebec (not seen).] 

Parmelia sinuosa var. virescens Kremplehuber, 1861:131. 

[Type collection: Mittenwald, Germany, Krempelhuber 

(M, lectotype; UPS, isolectotype).] 

Parmelia meyeri Zahlbruckner, 1905:82. [Type collection: 

Chimborazo, Ecuador, Meyer 365 (W, lectotype).] 

friesshire, England, Burgess (LINK, lectotype).] 

Thallus adnate to loosely adnate on bark and 

twigs, rarely on rock, sometimes suberect at the 

periphery, pale greenish yellow, 2-7 cm broad; 

lobes sublinear, short, 0.7-3 mm wide; upper surface plane, continuous, sorediate, less commonly 

pustulate-sorediate, the soralia capitate, orbicular 

to diffuse, subterminal; lower surface densely 

rhizinate, the rhizines sometimes forming a marginal mat. Apothecia very rare, adnate, 1-4 mm 

in diameter; spores 7p-9p X lop-14y. 

CmMIsmY.--Cortex K-, medulla K+ yellow 

turning red, P+ orange (usnic acid and salacinic 

acid, sometimes with norstictic acid and a trace of 

stictic acid). 
64 SMITHSONIAK CONTRIBUTIOSS TO BOTAKY 

FIGURE 18.--Species of Hypotrachyna: a, H. silvatica (Malnze in S); b, H. singularis (Hutchi?iso?z 

and It’right 57041; c, H. sinuosa (Hale 430Tla); d, H. steyermaikii (Steyermark 106123); e, H. 

subafinis (Imshaug 22531); f, H. sublaevigata (Hale 38810). (Scale in mm.) 
NUMBER 25 65 

DIsTRIBuTIoN.-A~aska to Oregon, southeastern 

United States, Mexico, West Indies, Andean region from Venezuela to Chile, Europe, southeastern Asia from southern Japan to Java. 

HABITATS.-On trees (hardwoods and conifers) 

and rarely rocks and on small shrubs in the 

paramo at 2000-3500 m (to near sea level in 

boreal areas). 

REMARKS.-The development of soredia in this 

pantropical-montane, pantemperate species is extremely variable. The type and the bulk of the 

specimens examined have discrete subterminal 

soralia and powdery soredia. At the other extreme, the soralia (as in Parmelia meyeri) may 

originate as laminal to subterminal pustules which 

become coarsely sorediate or rarely remain esorediate. There are various intergradations between 

these types. Most of the problematic specimens 

have been collected in the West Indies or in the 

high Andes. The chemistry also is variable, typically salazinic acid with or without norstictic acid 

but sometimes with stictic acid or different combinations of these. We may be dealing with several intergrading morphological and chemical 

populations but for the present they are most conveniently recognized as a single species. 

Smith cited three syntypes: the Dumfriesshire 

lectotype, a Menzies specimen from Nova Scotia 

(LINN, US) which can be identified as Parmelina galbina (Acharius) Hale, and a Turner 

and Hooker collection from Ballacheulish which 

I have not located. Parmelia despreauxii is represented by collections in BM and G apparently 

identified by Delise himself, but the most appropriate lectotype probably is housed in Strasbourg 

(STR). A syntype of P. meyel-i (Meyer 388) is not 

a Pamelia at all but is a member of the Lecanoraceae. 

anishi 370 (KOBE) 393 (US); Puebla, Van Horn 

59199 (US); Veracruz, Hale 20897. Costa Rica: 

San Jose, Culberson 12580, 13199 (DUKE); 

Haiti: Ouest, Zmshaug 23016 (MSC, US). Dominican Republic: La Vega, Zmshaug 23392, 23518 

(MSC, US), Wetmore 3673 (MSC, US). Colombia: Boyaca, Cuatrecasas 1787a (US); Cundinamarca, Linclig 2548 (P); Santander, Killip and 

Smith 18595 (US). Venezuela: Merida, Hale 

42711, 42794, 42820, 43210, Nash 1983 (US), 

Schwabe (M). Bolivia: Cochachamba, Ugent (US, 

SPECIhlENS ExAhfINED.-Mexico: OaXaCa, NUkWIS). Chile: Aysen: Santesson 4603, 8413 (UPS); 

Juan Fernandez, Imshaug 37440 (MSC); Magallanes, Santesson 1848, 1994, 6441, 7827 (S), 6586 

(UPS); Tierra del Fuego, Santesson 404, 420, 

7864 (S), 5471 (UPS); Valdivia, Santesson 7262 

(9 

70. Hypotrachyna steyermarkii, new combination 

FIGURE 18d 

Parrnelia steyerrnarkii Hale, 1974b:268. [Type collection: 

Sierra Parima, Amazonas, Venezuela, Steyermark 106123 

(US).] 

Thallus closely adnate on mosses on bark, fragile, light mineral gray, about 8 cm broad; lobes 

narrow, dichotomously branched, 1-2 mm broad; 

upper surface plane, shiny, isidiate, the isidia 

initially simple and cylindrical but soon procumbent and in part lobulate, becoming ciliate; 

lower surface moderately rhizinate, the rhizines 

sparsely dichotomously branched. Apothecia not 

seen. 

Cmammu.-Cortex Kf yellow, medulla K-, 

C+ orange, P- (atranorin, barbatic acid, 4-0- 

demethylbarbatic acid, obtusatic acid, and norobtusatic acid). 

DISTRIBUTION.-VeneZUeIa. 

HABITAT.-On a small tree in forest at 1300 m 

elevation. 

REMARKS.-This species is distinguished from 

chemically identical H. imbricatula by the very 

narrow lobes and the consistently procumbent, 

ciliate isidia. 

71. Hypotrachyna subafinis, new combination 

FIGURE 18e 

Parrnelia subafinis Zahlbruckner, 1929:213. [Type collection: 

Based on P. ufjiinis Vainio, not P. afinis (Dickson) Acharius, 

1803:212 (= Pannuria).] 

Parrnelia ufiinis Vainio, 1890:52. [Type collection: Carassa, 

Minas Gerais, Brazil, Vainio 1331 (TUR, Vainio herbarium 

number 3010, lectotype; BM, isolectotype).] 

Thallus closely adnate on bark, sea-foam yellow, 3-5 cm broad; lobes sublinear, dichotomously 

branched, 0.4-2 mm wide; upper surface plane, 

continuous, sorediate, the soralia capitate, mostly 

laminal; lower surface densely rhizinate, the 

rhizines often projecting along the margins. Apo- 
66 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

thecia rare, adnate, 0.7-2.5 mm in diameter; 

spores 5p-7p X 8p-11p. 

CHEMIsmY.-Cortex K+ yellowish, medulla 

K-, C-, P+ pale orange (usnic acid, echinocarpic acid, and associated unknowns). 

DIsTRIBuTIoN.-Mexico, Central America, West 

Indies, Brazil. 

HABITAT.-oII trees (oaks and pines) in open 

forests at 1000-2000 m elevation. 

REMARKs.-This is a very small, closely adnate 

species without any close relatives, except perhaps 

H. velloziae. TVhile Vainio’s type was collected on 

hardwood trees, almost all collections from the 

West Indies, where it is most common, were taken 

from Pinus bark. 

SPECIMENS EXAMINED.-Mexico: Chiapas, Hale 

20352, 20384. Guatemala: Baja Verapaz, Hale 

38308, 38342. Haiti: Ouest, Zmshaug 22531 (MSC, 

US), 22539, 22764, 23081, 23599, 23761 (MSC), 

Wetmore 3203 (MSC, US). Dominican Republic: 

La Vega, Allard 16542, 16544, 16552, 16581, 16589 

(US); Santiago, Wetmore 3882 (MSC, US). Brazil: Minas Gerais, Vainio 771 (BM, M, TUR), 

986, 3010 (TUR), 1391 (TUR, UPS). 

72. Hypotrachyna sublaevigata, new combination 

FIGURE 1Sf 

Parmelia tiliacea var. sublaevigata Nylander, 1860:383. [Type 

collection: South America, Bonpland (P, lectotype; H, isolectotype),] 

Parmelia sublaevigata (Nylander) Piylander, 1867:306. 

Parmelia chilena Nylander, 18853612. [Type collection: Chile, 

M. Gay (P, lectotype).] 

Parmelin canescens Kurokawa in Hale and Kurokawa, 1964: 

167. [Type collection: Colchagua, Chile, Dusen 92 (S, holotype; US, isotype).] 

Thallus closely adnate on bark or twigs, 3-8 

cm in diameter, dull mineral gray but often turning red in the herbarium because of improper 

curating; lobes subirregular and short, crowded, 

2-5 mm wide; upper surface plane to rugulose, 

continuous, often becoming whitish pruinose; 

lower surface densely short rhizinate. Apothecia 

common, substipitate, 2-5 mm in diameter; spores 

C~~~~sm~.--Medulla K+ yellow turning red, 

P + orange (atranorin, norstictic and salazinic 

DIsmInuTIoN.-Mexico, Panama, and Chile. 

acids). 

5p-5~ X 8y-lOp. 

HABITAT.-on oaks, pine, Drimys, and Discaria 

in open or secondary forests at 1500-2100 m 

elevation. 

REMARKS.-The lobes of H. sublaeuigata tend 

to be subrotund, crowded, and closely adnate. 

Apothecia are usually numerous. In Panama this 

species could be confused with H. boquetensis, 

which contains only salazinic acid and has longer 

lobes. Nylander’s syntype of Parmelia tiliacea var. 

sublaevigata is Leprieur 504 (P), an isidiate 

species in the genus Bul bothrix Hale. Parmelia 

canescens was differentiated from H. sublaevigata 

only by the presence of norstictic acid, but later 

tests with thin-layer chromatography showed that 

salazinic acid also is present. 

SPECIMENS EXAMINED.-Mexico: Durango, Bell 

s.n. (DUKE), Fox M87 (US); Michoacan, Arsene 

4046b, 4459b (US); Sinaloa, Culberson 13449 

(DUKE). Panama: Chiriqui, Hale 38810. Chile: 

Curico, Mahu 3492 (US); Santiago, Mahu 28, 

2133a (US); Tierra del Fuego, Santesson 6004 

(S, US); Valdivia, Santesson 3719, 3730 (S, US); 

3722 (S); Valparaiso, Mahu 1140 (US). 

73. Hypotrachyna subphysodalica, 

new combination 

FIGURE 19a 

Parmelia subphysodalica, Hale, 1974b:268. [Type collection: 

Puerto Ballena, Chile, Imshaug 43121A (MSC, holotype; 

US, isotype).] 

Thallus closely adnate on twigs, yellowish 

green, 6-7 cm broad; lobes sublinear, about 2 mm 

wide; upper surface plane, shiny, moderately 

isidiate, the isidia cylindrical, simple, becoming 

short ciliate, about 0.3 mm high; lower surface 

moderately rhizinate. Apothecia not seen. 

CmmsmY.--Cortex K--, medulla K--, P + 

orange-red (usnic acid and physodalic acid). 

DIsmrnuTIoN.-Southern Chile. 

HABITAT.-oII tree in mossy forest near sea level. 

REMARKS.-The diagnostic features of this rare 

species are the yellow color, isidia, and presence 

of physodalic acid. While it might be mistaken 

for H. microblasta, its range is far south of that 

species. It is not related to the only other1 

physodalic-acid-containing species in the genus, H. 

physodalica, a paramo lichen in the northern 

Andes. 
NUMBER 25 67 

74. Hypotrachyna subplumbeata, new combination 

FIGURE 19b 

Parmelia subplumbeata Dodge, 1959: 106. [Type collection: 

Parrnelia kahusiensis Dodge, 1959: 105. [Type collection: Mt. 

Mt. Karisimbi, Congo, Derscheid 2504 (US, lectotype).] 

Kahusi, Congo, Hendrickx 4300 (EAH, holotype).] 

Thallus loosely adnate, rather coriaceous, 5-10 

cm broad; lobes sublinear, crowded, 2-4 mm wide; 

upper surface smooth, heavily white-maculate, 

without soredia or isidia; lower surface densely 

rhizinate, the rhizines densely dichotomously 

branched. Apothecia abundant, substipitate, 2-5 

mm in diameter; spores not developed. 

CHEMISTRY.-cOrteX K+ yellow, medulla K-, 

C+ orange, P- (atranorin, barbatic acid, and 

4-0-demethylbarbatic acid). 

DIsTRIBUTI0N.-Mexico and Africa. 

HABITAT.-oII small trees near the tree line at 

about 3800 m elevation. 

FIGURE 19.-Species of Hypotrachyna: a, H. subphysodalica (Zmshaug 43121.4); b, H. subplumbeata (Moore and Wood 4944); c, H. subsaxntilis (Arsene 3974); d, H. thysanota (Hale 21118). 

(Scale in mm.) 
68 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

REMARKs.-According to Culberson and Hale 

(1973), H. subplumbeata is the remnant of an 

ancient population (now mostly extinct in the 

New World but widespread in the Old World) 

that contains barbatic acid without obtusatic acid. 

It could be the progenitor species which hybridized with H. pziluinata to produce the obtusaticacid-containing species in the H. physcioides constellation. The thallus is rather coriaceous and 

the maculae strongly developed, just as in the 

African population. 

SPECIMENS EXAiCIINED.-kfeXiCO: Mexico, Kiener 

18577 (DUKE), Moore and Wood 4944 (US). 

75. Hypotrachyna subsaxatilis, new combination 

FIGURE 19c 

Parmelia subsaxatilis Bouly de Lesdain, 1914:6. [Type collection: Cerro Azul, Michoacan, Mexico, Arsene 3973 (P, lectotype) .I 

Thallus loosely adnate on rock, dark mineral 

gray, 4-8 cm broad; lobes sublinear, short, 

crowded, 1-3.5 mm wide; upper surface plane to 

rugulose and cracked, lacking isidia and soredia; 

lower surface sparsely to moderately rhizinate, 

the rhizines sparsely dichotomously branched. 

Apothecia adnate, 1.5-4.5 mm in diameter; spores 

CHEiwsmY.-Cortex K+ yellow, medulla K+ 

yellow turning red, P+ orange-red (atranorin and 

salazinic acid). 

7p-91.1 X 11p-15p. 

DISTRIBUTION.-Mexico. 

HABITAT.-on rocks and soil over rocks in open 

areas at about 2200 m elevation. 

REMARKS.-While amply distinct and collected in 

quantity by Arsene, this species remains a rarity 

known only from Mexico. It is unrelated to any 

other species in the genus. 

SPECIMENS EXAMINED.-MeXiCO: Michoacan, 

Arsene 3974, 3974b (US). 

76. Hypotrachyna thysanota, new combination 

FIGURE 19d 

Parmelia thysanota Kurokawa in Hale and Kurokawa, 1964: 

186. [Type collection: 11 km E Las Vigas, Veracruz, Mexico, 

Hale 20943 (US, holotype).] 

Partnelia nakanishii Hale, 1971d3434. [Type collection: Mt. 

Leconte, Sevier County, Tennessee, Nakanishi 205 (US, 

holotype; KOBE, DUKE, TKS, isotypes).] 

Thallus loosely adnate, whitish mineral gray, 

to 15 cm broad; lobes linear, dichotomously 

branched, subdivaricate, 2-5 mm wide; upper 

surface plane, shiny, becoming pruinose near the 

tips, sorediate, the soralia mostly subterminal, 

often protruding, coalescent; lower surface densely 

rhizinate, the rhizines densely dichotomously 

branched, often projecting as a mat beyond the 

margins. Apothecia not seen. 

CHEhmmY.-Cortex K+ yellow, medulla K-, 

C- red, P+ red (atranorin, echinocarpic acid, 

gyrophoric acid, and microphyllinic acid). 

DIsTRIBuTIoN.-Southeastern United States and 

Mexico. 

HABITAT.-oII conifers (Abies, Pinus) in open 

forests at about 2000 m elevation. 

REMARKs.-Gyrophoric acid was the only constituent identified with microcrystal tests for this 

species. Dey (1974) has analyzed the type specimen with thin-layer chromatography and discovered the additional substances. This is the only 

report of microphyllinic acid outside of Cetraria, 

although Culberson and Hale (1973) postulated 

its possible occurrence in Hypotrachyna. Superficially, this species is similar to H. Zaeuigata or 

H. densirhizinata; no nonsorediate progenitor 

has been discovered. 

SPECIMENS ExAMINED.-Additional records from 

the United States and Mexico are listed in Hale 

and Kurokawa (1964:186), and in Hale (1971d: 

434) under Parmelia nakanishii. 

77. Hypotrachyna velloziae, new combination 

FIGURE 20 

Parmelia velloziae Vainio, 1890:64. [Type collection: Carassa, 

Minas Gerais, Brazil, Vainio 1455 (TUR, Vainio herbarium 

number 2892, lectotype; BM, FH, M, UPS, isolectotypes).] 

Thallus closely adnate to bark, fragile, pale 

viridine yellow, 3-7 cm in diameter; lobes sublinear, dichotomously branched, 0.5-2 mm wide; 

upper surface plane, continuous, sorediate, the 

soralia mostly laminal, capitate, often coalescing; 

medulla turning yellow-ochre under the soralia; 
NUMBER 25 69 

lower surface moderately rhizinate, the rhizines 

sparsely dichotomously branched. Apothecia not 

seen. 

CHE;LIIsmY.-Cortex K-, medulla K-, PS 

orange-red (usnic acid, protocetraric acid, and 

rhodophyscin). 

DISTRIBUTION.-west Indies and Brazil. 

HABITAT.-on trunk of trees (Pinus) in open or 

virgin pine forests (West Indies) at 1200-1600 m 

elevation. 

REhlARKs.-H~potraChylza uelloziae parallels H. 

subafinis in distribution (mainly West Indies) 

and habitat (pine trees) but differs completely in 

chemistry. There is no nonsorediate sexual morph. 

SPECIMENS ExAMINED.-Haiti: Ouest, Imshaug 

22535 (MSC, US), Wetmow 3071, 3217 (MSC, US). 

FIGURE 20.-Hypotrachyna uelloziae (Wetmore 307). Dominican Republic: La Vega, Allard 16541a, 

(Scale in mm.) 16562d (US). 

Literature Cited 

Acharius, E. 

1803. Methodus qua omnes detectos Lichenes, 394 pages. 

1814. Synopsis methodica lichenum. 392 pages. Lund. 

1942. Distribution and Ecology of Some South ScandiStockholm. 

Almborn, 0. 

navian Lichens. 252 pages. Lund. 

.4sahina, Y. 

1951. Lichenes Japoniae novae vel minus cognitae (7). 

1953. Lichenes Japoniae novac vel minus cognitae (10). 

1968. Lichenologische Notizen (209). Journal of Japanese 

Baker, C. J., .4. Elix, D. P. H. Murphy, S. Kurokawa, and 

R. H. Grover 

Parmelia reptans, a New Lichen Species Producing 

the Depsidone, Succinprotocetraric .Acid. Australian Journal of Botany, 213137-140. 

Journal of Japanese Botany, 26:329-334. 

Journal of Japanese Botany, 28:65-68. 

Botany, 43:97-101. 

1973. 

Bouly de Lesdain, hl. 

Crombie, J. M. 

1914. Lichens de Mexique. 31 pages. Mexico. 

1876. On the Lichens Collected by Prof. R. 0. Cunningham. Journal of the Linneaiz Sociely of London, 

15~222-230. 

Culherson, C. F. 

1966. The Structure of a New Depsidone from the Lichen 

Parmelia livida. Phytochenistry, 53835-818. 

1972a. Improved Conditions and New Data for the Identification of Lichen Products by a Standardised 

Thin-Layer Chromatographic hfcthod. Journal of 

Chromatography, 72:113-125. 

1972b. High-speed Liquid Chromatography of Lichen Extracts. Bryologist, 55:54-62. 

Culberson, C. F., and M. E. Hale, Jr. 

Chemical and RloIphological Evolution in Parmelia 

sect. Hypotmchynn: Product of Ancient Hybfidization? Brittonia, 253162-173. 

The Parmelia qum~ina Group in North America. 

American ,Journal of Botany, 48: 168-174. 

Chemosystematics and Ecology of Lichen-Forming 

Fungi. Annual Xeuiew of Ecology and Systematics, 

1 : 153-1 70. 

The Parmelia perforata Group: Siche Characteristics of Chemical Races, Speciation by Parallel Evolution, and a Sew Taxonomy. Bryologkt, 76:20-29. 

1967. Hahitat Selection by Chemically Differentiated 

Races of Lichens. Science, 158:1195-1197. 

1970. .A Phylogenetic View of Chemical Evolution in thc 

Lichens. Bryologist, 73:l-31. 

1973. Parallel Evolution in Lichen-Forming Fungi. 

Science, 180:196-198. 

1973. 

Culberson, TV. L. 

1961. 

1970. 

1973. 

Culherson, Tt’. L., and C. F. Culberson 

Degelius, G. 

1941. Contributions to the Lichcn Flora of h’orth America, 11. The Lichen Flora of the Great Smoky 

Mountains. Arkiv for Botanik, 304 (3):l-80. 

Dey, J. 

1074. Nerc- Interesting I’armcliae (Lichens) from the 

Southern .Appalachianr. Castanea, 39:360-369. 

Some Lichens of Tropical Africa, 111. Parmeliaceae. 

Annals of the Missouri Botanical Garden, 46:39-193. 

Dodge, C. TV. 

1959. 
70 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Duby, J, E. 

1830. Aug. Pyrami de Candolle Botanicon gallicum. 1068 

pages. Paris. 

Durand, T., and H. Pittier 

1891. Primitae Florae Costaricensis. Volume 2, pages 49- 

97. Bruxelles. 

Du Rietz, G. 

1924. Die Soredien und Isidien der Flechten. Svensk BotanFee, A. 

1824. 

isk Tidskrift, 18:371-396. 

Essai sur les cryptogames des ecorces exotiques oficinales. 167 pages. Paris. 

Floerke, H. G. 

Gyelnik, V. 

1815. Deutsche Lichenen, I. 18 pages. Rostock. 

1931. Additamenta ad cognitionem Parmeliarum, 11. 

Fedde Repertorium, 29:273-291. 

1934. Additamenta ad cognitionem Parmeliarum, V. 

Fedde Repertorium, 36: 151-156. 

1938a. Revisio typorum ab auctoribus variis descriptorum. 

Annales Musei Nationalis Hungarici, 31:2-57. 

1938b. Additamenta ad cognitionem Parmeliarum. Annales Mycologica, 36:267-294. 

Hale, M. E., Jr. 

1959. New or Interesting Parmelias from North and 

Tropical America. Bryologist, 62: 123-132. 

1960. A Revision of the South American Species of 

Parmelia Determined by Lynge. Contributions 

from the United States National Herbarium, 36: 

1-41. 

1963. Populations of Chemical Strains in the Lichen 

Cetraria ciliaris. Brittonia, 15: 126-133. 

1965. A Monograph of Parmelia subgenus Amphigymnia. 

Contributions from the United States National Herbarium, 36: 193-358. 

1967. New Taxa in Cetraria, Parmelia, and Parmeliopsis. 

Bryologist, 70:414-422. 

1971a. Morden-Smithsonian Expedition to Dominica: The 

Lichens (Parmeliaceae). Smithsonian Contributions 

to Botany, 4:l-25. 

1971b. Five New Parmeliae from Tropical America. Phytologia, 22:30-35. 

1971c. New Parmeliae (Lichens) from Africa. Phytologia, 

2294-96. 

1971d. New Species of Parmelia sect. Hypotrachyna 

(Lichenes). Phytologia, 22:433438. 

1972a. Fine Structure of the Cortex in the Lichen Family 

Parmeliaceae Viewed with the Scanning-Electron 

Microscope. Smithsonian Contributions to Botany, 

1972b. Parmelia pustulifera, a New Lichen from Southeastern United States. Brittonia, 24:22-27. 

1974a. Delimitation of the Lichen Genus Hypotrachyna 

(Vainio) Hale. Phytologia, 28:340-342. 

1974b. New Species of Parmelia (Lichenes) from Tropical 

America, 1. Phytologia, 28:265-269. 

10: 1-92. 

Hale, M. E., Jr., and S. Kurokawa 

1964. Studies on Parmelia subgenus Parmelia. Contributions from the United States National Herbarium, 

36: 121-191. 

Hillmann, J. 

1936. Parmeliaceae. Section 3, pages 1-309, in part 5, 

volume 9, in L. Rabenhorst, Rabenhorst's Kryptogamenpora. Leipzig. 

Hue, A. M. 

1899. Lichenes extra-europaei. Nouvelles Archives du 

Museum Paris, series 3, 1:l-250. 

Species novae in excelsis Ruwenzori in expeditione 

Duck Aprutii lectae, IV: Lichenes. Annuli di 

Botanica, 6:407-409. 

Verzeichnis der Lichenen, welche M. Wagner auf 

seinen Reisen in Centralamerika 1858/59 gesammelt 

hat. Flora, 44:129-131. 

1873. Lichenes brasilienses in Warming Symbolae ad 

floram Brasiliae cognoscendam particula, XIV. 

Videnskabelige Meddelelser fra Dansk Naturhistorisk forening i Kjobenhavn, 25: 1-35. 

The Lichen-Flora of Great Britain, Ireland and the 

Channel Islands. 502 pages. Shrewsbury. 

VIII. Memoir on the Spermagones and Pycnides of 

Filamentous, Fruticulose, and Foliaceous Lichens. 

Transactions of the Royal Society of Edinburgh, 

Jatta, A. 

1908. 

Krempelhuber, A. 

1861. 

Leighton, W. A. 

1871. 

Lindsay, W. L. 

1859. 

22:lOl-303. 

Lynge, B. 

1914. Die Flechten der ersten Regnellschen Expedition. 

Die Gattungen Pseudoparmelia gen. nov. und 

Parmelia Ach. Arkiv for Botanik, 13 (13):l-172. 

1917. Ueber einige Regnellschen Parmelian aus Matto 

Grosso, Brasilien. Arkiv for Botanik, 15 (1):1-4. 

1925. On Some South American Lichens of the Genera 

Parmelia, Candelaria, Teloschistes, and Pyxine. Nyt 

Magazin for Naturvidenskaberne, 62:83-97. 

Mitsuno, M. 

1953. Paper Chromatography of Lichen Substances, I. 

Pharmaceutical Bulletin (Tokyo), 14:659-669. 

Muller Argau, J. 

1879. Lichenes aequinoctiali-americani a cl. E. Andre 

annis 1875-76 praesertim in editroribus Ecuador et 

in Nova Granata lecti. Revue Mycologique, 1:163- 

171. 

1880. Lichenologische Beitrage, XI. Flora, 63:259-268. 

1881. Lichenologische Beitrage, XII. Flora, 64:81-88. 

1887. Lichenologische Beitrage, XXVI. Flora, 70:316-320. 

1889. Lichenes Sebastianopolitani lecti a cl. Dr. Glaziou. 

Nuovo Giornale Botanic0 Italiano, 21:353-364. 

1891. Lichenologische Beitrage, XXXV. Flora, 74:371-382. 

1860. Synopsis methodica lichenum, I. 430 pages. Paris. 

1863. Lichenographiae Novo-Granatensis Prodromus. Acta 

Nylander, W. 

Societas Scientarum Fennica, 7:415-504. 
NUMBER 25 

1867. Lichenes. Additamentum in J. Triana and J. E. 

Planchon, Prodromus Florae Nova-Granatensis ou 

Enumeration des Plantes de la Nouvelle-Grenade. 

Annales Sciences Naturelles Botanique, series 5, I: 

1874. Addenda nova ad Lichenographiam europaeam. 

1885. Parrneliae exoticae novae. Flora, 68:605-615. 

1970. Das Konzept der Artenpaare bei den Flechten. 

. Deutsche Botanische Gesellschaft Neue Folge, 4: 

Die taxonomische Behandlung von Artenpaare bei 

den Flechten. Botaniska Notiser, 125:77-81. 

Polakowsky, H. 

Bryophytas et cormophytas costaricensis anno 1875 

lectas. Journal of Botany British and Foreign, 13: 

225, 226. 

Rebentisch, J. F. 

Prodrornus Florae Neomarchicae. 406 pages. Berlin. 

Smith, J. E. 

301-354. 

Flora, 573305-318. 

Poelt, J. 

187-198. 

1972. 

1877. 

1804. 

1808. English Botany. Volume 26, page 1852. London. 

1809. English Botany. Volume 29, page 2050. London. 

1827. Systerna Vegetabiliurn, 4. Edition 16, part 2, 410 

Sprengel, K. 

pages. Gottingae. 

Stahl, E., and P. J. Schorn 

1961. Dunnschicht-chromatographie hygrophiler Arzneipflanzenauszuge, VIII. Mitteilung: Cumarine, Flavonderivate, Hydroxysauren, Gerbstoffe, Antracenderivate und Flechteninhaltsstoffe, Hoppe-Seyler's 

Zeitschrift fur Physiologische Chemie, 325:263-274. 

1903. Flechten von Kamerun und dem Kamerunberg 

(Fako), gesammelt von Alfred Bornmuller in den 

Jahren 1897 und 1898. Verhandlungen der 

Kaiserlich-koniglichen zoologisch-botanischen Gesellschaft in Wien, 53:227-236. 

Steiner, J. 

Stirton, J. 

1874. New British Lichens. Grevillea, 3:79. 

Taylor, T. 

1847. New Lichens, Principally from the Herbarium of 

W. J. Hooker. Hooker Journal of Botany, G:148- 

197. 

Thomson, J. W. 

1961. Evolution in the Lichen Genus Physcia. Pages 267- 

271 in Recent Advances in Botany. Toronto: University of Toronto Press. 

71 

Vainio, E. A. 

1890. 

1896. 

1898. 

1899. 

1900. 

1907. 

1915. 

Etude sur la classification naturelle et la morphologie des Lichens du Bresil. Acta Societas Fauna et 

Flora Fennica, 7 (7):l-247. 

Lichenes Antillarum a W. R. Elliott collecti. 

Journal of Botany British and Foreign, 34:31-36. 

Lichenes quos in Madagscaria centrali Dr. C. Forsyth Major collegit. 

Lichenes novi rarioresque, I. Hedwigia, 38: (121)- 

(125). 

Reactiones lichenum a J. Mullero Argoviensi descriptorum. Memoire E'Herbier Bossier, 5: 1-17. 

Lichenes novi rarioresque, IV. Hedwigia, 46: 168- 

181. 

Additamenta ad lichenographia antillarum illustrandam. Annales Academiae Scientarurn Fennicae, 

Hedwigia, 37: (33)- (37). 

6~ (7):l-226. 

Vareschi, V. 

1962. Resultados liquenologicos de excursiones effectuadas 

en Venezuela, No. 2: Acerca del genero Parrnelia en 

Venezuela. Acta Biologica Venezuelica, 3:201-232. 

Studies on the Chemistry of Lichens, I: Separation 

of Depside Components by Paper Chromatography. 

Acta Chernica Scandinavica, 6: 81 8-825. 

Wachtmeister, C. A. 

1952. 

Zahlbruckner, A. 

1902. 

1905. 

1908. 

1909. 

1912. 

1926. 

1929. 

1934. 

Studien uber brasilianische Flechten. Sitzungsberichte der kaiserliche Akademie Wissenschift. 

Mathematisch-naturuGssenschajtliche Klasse, .4bteilung 1, 111:357432. 

Flechten, im Hochlande Ecuadors gesammelt von 

Prof. Dr. Hans Meyer im Jahre 1903. Beihefte zur 

Botanischer Centralblatt, 19:75-84. 

Beitrage zur Flechtenflora Brasiliens. Bulletin de 

1'Herbier Bossier, series 2, 8:459-468. 

Lichenes (Flechten). In Ergebnisse der botanischen 

Expedition der kaiserlichen Akademie der TVissenschaften nach Sudbrasilien 1901. Denkschrift 

Akademie der Wissenschajt in Wien. Mathematischnaturwissenschaftliche Klasse, 83:87-211. 

Neue Flechten. Annales Mycologici, 103359-384. 

Chilenische Flechten gesammelt von C. Skottsberg. 

Meddelanden fran Goteborgs Botaniska Tradgard, 

Catalogus lichenum universalis. Volume 6, 323 

pages. Leipzig. 

Flechten der Insel Formosa. Fedde Repertoriitm, 

2: 1-26. 

33:22-68. 
Index 

(Synonyms in italics) 

Hypotrachyna andensis, 23 

bahiana, 23 

bogotensis, 23 

boquetensis, 25 

brasiliana, 25 

brevirhiza, 26 

caraccensis, 26 

chicitae, 28 

chlorina, 28 

consimilis, 28 

cantradicta, 29 

costaricensis, 29 

croceopustulata, 30 

dactylifera, 30 

degelii, 31 

densirhizinata, 31 

dentella, 33 

ducalis, 33 

eitenii, 33 

enderythraea, 34 

endochlora, 34 

ensifolia, 35 

erythrodes, 35 

exsplendens, 37 

flavida, 37 

flavovirens, 37 

formosana, 38 

gigas, 38 

gondylophora, 40 

gracilescens, 40 

imbricatula, 41 

immaculata, 41 

intercalanda, 42 

isidiocera, 42 

koyaensis, 44 

laevigata, 44 

lineariloba, 45 

livida, 45 

lopezii, 47 

malmei, 47 

microblasta, 47 

minima, 48 

monilifera, 48 

neodissecta, 49 

novella, 49 

obscurella, 51 

oostingii, 51 

osorioi, 51 

osteoleuca, 51 

palmarum, 52 

partita, 52 

peruviana, 54 

physcioides, 54 

physodalica, 55 

pluriformis, 55 

producta, 56 

prolongata, 56 

protenta, 58 

protoboliviana, 58 

pseudosinuosa, 58 

pulvinata, 59 

rachista, 59 

reducens, GO 

revoluta, GO 

rhabdiformis, 62 

rockii, 62 

silvatica, 63 

singularis, 63 

sinuosa, 63 

steyermarkii, 65 

subaffinis, 65 

sublaevigata, 66 

subphysodalica, 66 

subplumbeata, 67 

subsaxatilis, 68 

thysanota, 68 

velloziae, 68 

sinuosus, 63 

amoena, 29 

anaptychioides, 58 

bahiana, 23 

bogotensis, 23 

boliviana, 54 

boliviana var. cephalota, 44 

boquetensis, 25 

brasiliana, 25 

brasiliana var. erythrodes, 35 

brasiliana var. novella, 49 

brevirhiza, 26 

canescens, 66 

caraccensis, 26 

caraccensis var. guatemalensis, 26 

caraccensis f. pulvinata, 59 

chicitae, 28 

Lichen laevigatus, 44 

Parinelia afinis, 65 

chilena, 66 

chlorina, 28 

confusula, 54 

consimilis, 28 

contradicta, 29 

costaricensis, 29 

croceopus t ti la ta, 30 

crustacea, 63 

culmigena, 23 

dactylifera, 30 

deformis, 29 

degelii, 31 

densirhizinata, 31 

dentella, 33 

despreauxii, 63 

digitata, 54 

ducalis, 33 

eitenii, 33 

enderythraea, 34 

endochlora, 34 

endochlora f. sorediosa, 34 

endorubra, 26 

endorzrbra f. imbricatiformis, 48 

ensifolia, 35 

eryt h rocard ia, 34 

erythrodes, 33 

everniaeformis, 55 

exporrecta, 38 

exsplendens, 37 

flava, 37 

flava f. albescens, 23 

flava var. stellata, 37 

pava var. subdichotoma, 37 

flavida, 37 

flavovirens, 37 

formosana, 38 

fragilis, 42 

gallicana, 34 

gigas, 38 

glaziovii, 25 

gondylophora, 40 

gracilescens, 40 

gracilescens f. obscurella, 51 

gracilis, 54 

hypotrachyna, 29 

imbricatula, 41 

immaculata, 41 

insinuans, 23 

72 
NUMBER 25 73 

intercalanda, 42 

isidiocera, 42 

jamaicensis, 48 

kahusiensis, 67 

koyaensis, 44 

laevigata, 44 

laevigata var. gracilis, 54 

lineariloba, 45 

livida, 45 

lobulifera var. insensitiva, 35 

lobulifera var. luteoreagens, 41 

lobulifera var. sanguineoreagens, 59 

madagascariensis, 34 

malmei, 47 

meyeri, 63 

microblasta, 47 

millaniana, 34 

minima, 48 

monilifera, 48 

nakanishii, 68 

neodissecta, 49 

neopropagulifera, 48 

norstictica, 48 

novella, 49 

oostingii, 51 

osorioi, 51 

osseoalbida, 45 

osteoleuca, 51 

palmarum, 52 

peruviana, 54 

physcioides, 54 

physodalica, 55 

pittieri, 34 

pluriformis, 55 

pluriformis var. chlorocarpa, 55 

pluriformis var. multifida, 55 

prolongata, 56 

propagulifera, 48 

protoboliviana, 58 

pseudorevoluta, 48 

pseudosinuosa, 58 

pulvinata, 59 

rachista, 59 

reducens, 60 

regis, 49 

revoluta, 60 

revoluta f. isidiosa, 48 

rhabdiformis, 62 

rockii, 62 

silvatica, 63 

silvatica var. pinnata, 63 

silvatica var. radiata, 63 

singularis, 63 

sinuosa, 63 

sinuosa var. caraccensis, 26 

sinuosa var. virescens, 63 

steyermarkii, 65 

subafinis, 65 

sublaevigata, 66 

sublaevigata f. isidiosa, 29 

subphysodalica, 66 

subplumbeata, 67 

subpluriformis, 55 

subsaxatilis, 68 

subsinuosa, 59 

thysanota, 68 

tiliacea var. leucina, 30 

tiliacea var. sublaevigata, 66 

tropica, 29 

tropica var. deformis, 29 

velloziae, 68 

xanthomyela, 34 

xanthomyela f. sorediosa, 34 

zahlbruckneri, 52 

* U.S. GOVERNMENT PRINTING OFFICE: 1975- 586-378/19