SMITHSONIAN CONTRIBUTIONS TO BOTANY 0 NUMBER 40 

Morden-Smithsonian Expedition 

to Dominica: 

The Lichens (Graphidaceae) 

Michael Wirth 

and Mason E. Hale, Jr. 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1978 
ABSTRACT 

LVii-th, Rl., and Mason E. Hale, Jr. ~Iortlen-Smitlisonian Expedition to 

Dominica: The Lichens (Graphidaceae). Smithsonian Contribulions to Botany, 

number 40, 64 pages, 23 figures, 1978.-The four main genera of the Graphidaceae 

in Dominica, Gruphis (27 species), Phaeogiaphis (7 species), Graplzina (25 species), 

and Plraeographina (6 species), are treated monographically with full synonymy, 

descriptions, chemistry, and illustrations. The family as a whole is common in all 

primary and secondary forests and at all elevations. Eight new species are 

de scr i I] etl : Graph i i i I 1 \ ha 11 g i i, G . isid i ife~ci , Plici <: og~rr f, Ii is ?ri oitle n i i, G YN p h i II a 

cmrneoviridis, G. sziberythrella, G. triphoroides, Phaeogiaphitza utrovermicularis, 

and P. coriaria. Three new combinations, Gmphina coliinzbina (Tuckerman) 

Wirth and Hale, G. illinata (Eschweiler) Wirth and Hale, and G. plurispora 

(Redinger) TYirth and Hale, are also made. 

OFFICIAL PLL~LICATIOA DATE is handstam ed in a limited number of initial copies and is recorded 

in the Institution’s anilual report, SnzitEsoriian l.ear. SERIFS COLEK DESIGS: Leaf clearing from the 

Katsura tree CerridiphJllum japonicum Siebold and Zuccarini. 

Libraiy of Congress Cataloging in Publication Data 

LVirth, Michael. 

Xlorden-Smithsonian Expedition to Dominica: the Lichens (Graphidaceae). 

(Smithsonian contributions to botany ; no. 40) 

Bibliography: p. 

Includes index. 

Supt. of Docs. no.: SI 1.29:40 

1, Graphidaceae. 2. Lichen-Dominica. 3. Xlorden-Smithsonian Expedition to Dominica I. 

Hale, Mason E. joint author. 11. Title. 111. Series: Smithsonian Institution. Smithsonian 

contributions to botany ; no. 40) 

QWS2747 no. 40 [QK585.GS] 581’.08s [389’.1] 77-608313 
Contents 

Page 

Introduction 1 

Generic Delimitation ............................................... 1 

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

Species Delimitation ............................................... 3 

Spores 3 

Hymenium 3 

Exciple 3 

Thallus 5 

Chemistry 5 

Ecology 6 

6 

Key to the Genera of the Graphidacea nica 8 

Key to the Species of Graphis ....................................... 8 

25 

45 

Incorrect and Omitted Names ....................................... 49 

Index ............................................................ 52 

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

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

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

Evolution in the Graphidaceae ..... 

Key to the Species of Phaeographis ................................... Key to the Species of Graphina .................................. 29 

Key to the Species of Phaeographina ................................. 

Literature Cited ................................................... 50 

................................... ........... 54 

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

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

... 

111 

Morden-Smi thsonian Expedition 

to Dominica: 

The Lichens (Graphidaceae) 

Michael Wirth 

and Mason E. Hale, 3r. 

Introduction 

This study is a continuation of a floristic treatment of the lichen flora of Dominica. The first 

article on the Parmeliaceae (Hale, 1971) and a 

second on the Thelotremataceae (Hale, 1974) should 

be consulted for general information on the geography and ecology of the island. The specimens examined in this study came from four sources: the 

historical collections of Elliott (preserved in BM 

and TUR) and the more recent materials of Hale 

(US), Imshaug (MSC), and Wirth (US). A variety of 

localities and habitats was visited by each collector, 

although Hale worked primarily in the higher elevation rain forest that was being logged 1968-1972, 

and Imshaug and Wirth studied both the high 

mountain tops and lower, disturbed forests. While 

we do not claim that all of the Graphidaceae have 

been discovered, this treatment must include a high 

percentage of the graphidaceous flora of Dominica. 

We owe special thanks to hlrs. William J. Morden, 

who generously supported field and herbarium 

studies by Hale. Dr. Henry Imshaug very kindly 

placed his collections at our disposal. We are also 

indebted to Dr. Minoru Nakanishi for the use of his 

unpublished data and keys for many of the Asian 

Michael Wirth, New England College, Henniker, New Hampshire 03242. Mason E. Hale, Jr., Department of Botany, 

National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. 

species. Finally, we thank curators of the herbaria 

and institutions who so promptly sent specimens on 

loans, in particular Dr. Teuvo Ahti (H), Dr. Reino 

Alava (TUR), and Mr. Peter James (BM). 

Generic Delimitation 

The delimitation of the Graphidaceae is still a 

matter of difficulty. Most lichenologists now reject 

from the family all those groups that have obviously 

branched paraphyses, globular asci with a bitunicate 

nassasceous structure, and cylindrical spore locules. 

Thus, Chiodecton, Opegrapha, Arthonia, and their 

segregate genera are generally excluded from the 

Graphidaceae sensu stricto. Melaspilea also seems 

best excluded; even though the paraphyses frequently appear free and unbranched and the twoloculed spores are shared by a few Graphis species, 

the structure of the asci in Melaspilea seems quite 

dissimilar to that of Graphis. 

Using the criteria above, one is left with a group 

of four closely related genera without stroma-like 

tissue (Graphis, Phaeographis, Graphina, Phaeographina), four parallel genera with such tissue 

(Glyph is, Sarcographa, Medusu lina, Sarcograph ina), 

and a number of minor genera whose relationships 

are unclear (Helminthocarpon, Aulaxina, Gyrostomum, etc.). Although it is customary to refer to 

these major genera as ascohymenial, even that seems 

to be in doubt (cf. Groenhart, 1965:4). We have, in 

any case, continued the ascohymenial terminology 

1 
2 

usually applied to the family, using the terms “hymenium” and “paraphyses” without necessarily 

accepting them a5 ontogenetically correct. 

To a great extent, all genera (in phanerogams 

as well as in lichens) are artificial coiistructs, in that 

no fixed criteria exist for delimiting any taxa above 

the biological species. In groups Ivhere cytogenetics, 

breeding experiments, etc., can reveal something of 

the direction of evolution, genera may in fact be an 

approximation of biological (i.e. evolutionary) relatedness. Obviously, in the crustose lichens, such 

information is mostly lacking. In addition, ontogenetic studies of lichen reproductive structures 

are scarce and frequently open to different interpretations: for example: TVliat exactly is a hymeniuni? 

Is the structure of the ascus of great importance, or 

little?-and so on. 

In the light of such uncertainty, a reasonable 

approach seems to be the recognition of genera on 

practical grounds, so that at least problems of 

identification and curating are more manageable. 

On such grounds, the major genera of the Graphidaceae are reasonably delimited. By using $pore 

septation and color as primary criteria, and the 

development of stroma-like tissue as a secondary 

criterion, one can distinguish the major genera of 

the family with relative ease. 

To be sure, specimens (and species) exist that 

bridge the gap betlveen these form genera and tend 

to blur the distinctions. In Graphina IJeStilOideS, 

for example, only the end locules of otlienvise typical Graphis spores become longitudinally divided. 

Many of the very small-spored Gmpiiina species 

(G. insigni.r, G. colliculosa, etc.) have barely muriform spores which, wlien not quite mature, will 

resemble four-loculed Gmpli is spores. The problems of spore color and septation in such borderline species as Graphinn achasii and Pl7acogixpliina 

chi.ywcarpa have been mentioned elsewhere (TVirth 

and Hale, 1963). 

Intermediate states also exist between the sti-omatoid and nonstromatoid genera. A fine example 

can be seen in many specimens of Phacogmphis 

exaltata, which frequently show, in portions of a 

single thallus, characteristic? of both Phucogi-aphis 

and Sawographa. 

There i3, however, a more serious objection to 

spore genera. One can regard a genus as consisting 

of species that share a large number of characterstates in common; this approach, common among 

SUITHSONIAN COKTRIBCTIOSS TO BOTASY 

“numerical” taxonomists, assumes that “relatedness” in the biological sense frequently can be 

approximated by measuring the over-all similarity 

of taxa. As there are few (if any) good biological 

criteria for genera in crustose lichens, this approach 

may be quite useful. One can find many examples 

of species that are extremely similar but are presently placed in different genera only because of a 

difference in spore color and/or septation. TZ’e will 

call sucli forms “sporomorphs,” defined as follows: 

trvo or inore species that are extremely similar (or 

identical) in external morphology, anatomy, cheniistry, and spore size, as far as we can determine 

with present techniques, but that are presently 

placed in different genera on the basis of spore 

septatioii and /or spore color only. 

Sporomorphs are common in the Graphidaceae; 

good examples occurring at least in part in Dominica are Pliueogmph is art h on ioide.5 Graph i.r h ii m ilis 

and Gvuphina incrzi.rtan.r :I G1-aphi.r dunzastii. 

The exi5tence of iporomorphs casts considerable 

doubt on the ability of spore features to establish 

genera of closely related (i.e. very similar over-all) 

species. Is it possible, for example, that the muriform condition simply represents a later ontogenetic 

stage of the Graphis spore that has become genetically fixed? If such were the case (and indeed all 

muriform spores seem to go through a Gwphis-like 

juvenile stage), then Gmphina incwstans may be 

n recent descendant of Graphis clunrastii, and these 

two species should be regarded as closely related. 

Placing these species in different genera, while 

grouping G. inci‘tistans with otherwise very dissimilar Gmphina species, would then be illogical. 

If this objection to spore-based genera is accepted, 

then one is faced with a considerable problem of 

nomenclature. Ignoring the spore coIor and septation as major generic criteria, then only two choices 

appear to be acceptable. One choice would be the 

erection of scores of segregate genera based on overall similarity of exciple structure and chemistry. It 

is not apparent to us how much such a system 

could be established or used without the help of 

a computer. The second generic approach would 

be to reduce all the spore-based genera back into 

Gmphi.~. Such a system, which may be biologically 

quite acceptable, would raise immense practical 

problems; there are simply too many species of 

Grap1iicl;iceae (over one thousand) to be handled 

conveniently. 
NUMBER 40 3 

In light of these practical problems, we have 

chosen to accept the major spore-based genera for 

the Graphidaceae. Whenever possible, however, we 

have indicated similar species and sporomorphs, regardless of present generic lines. 

Species Delimitation 

With essentially no knowledge of speciation in 

the Graphidaceae, we have had to use the traditional approach of concentrating on discontinuities 

of morphology, anatomy, chemistry, and size. This 

approach works best with large collections of specimens; unfortunately, in the Graphidaceae, as in 

other tropical crustose groups, one is too often 

accomplishing no more than cataloguing types. In 

light of this problem we have elected to maintain 

a very narrow species concept, leaving “lumping” to 

some future monographer blessed with abundant 

collections. 

SpoREs.--Several features associated with spores 

supply useful means of separating species. In a family with such a large range of spore size (6 pm to 

300 pm), it is not surprising to find discontinuities 

of size classes. We have, whenever possible, maintained species on this basis. Within a given individual, however, spore size commonly varies by a 

factor of 1.5 to 2; with additional collections, many 

of the species presently maintained primarily on 

spore size may vanish, while in other cases more 

species will be described. 

A second spore character of considerable interest 

is number per ascus. In many specimens, this number remains quite constant, and can be used as a 

species criterion. A good example is Graphina 

chlorocarpa and G. balbisii; individuals with one 

spore per ascus are assigned to G. chlorocarpa and 

all others to G. balbisii. Unfortunately, the spore 

number is quite variable in other species; Graphina 

virginea commonly will have 2, 4, 6, or 8 spores per 

ascus in the same thallus. Only larger population 

samples, when they become available, can help us 

solve this problem. 

The number of locules per spore, though roughly 

correlated with spore size, may be useful where only 

four locules per spore occur. This condition seems 

constant, and we have accepted species (particularly 

in Phaeogmphis) where the main, and perhaps only, 

distinction consists of four versus more than four 

locules per spore. 

HYMExIuM.-Although hymenial character-states 

supply much useful information, we have seen very 

few cases where species are established primarily on 

hymenial features, One such feature is the presence 

of droplets or inclusions in the hymenium; it is not 

yet clear if these inspersions are a constant feature 

within a species or not. At least one species has 

been based primarily on these inclusions (Graphis 

inspersa Redinger (1935)). It should be noted that 

the droplets or particles disappear in Hoyer’s 

medium, and hence must be looked for in water 

mounts. 

In all the descriptions given here, hymenium 

height is stated. We have found this measurement 

to be a very useful single feature to indicate gross 

apothecium size. In species with very heavily carbonized exciples the hymenium height becomes proportionally small; for most other species, it is an 

excellent indicator of over-all size. Hymenial measurements include the epihymenium (“epithecium” 

of older literature), which is frequently darkened. 

ExcIPLE.-Exciple characters have traditionally 

formed the backbone of species delimitation in 

Graphidaceae. In addition, almost all subgeneric 

taxa are based on excipular characters (see Redinger, 1933:4 and 1935:Z). Many of these characters 

now appear to be quite variable and unreliable. In 

the discussion below, it should be emphasized that 

all observations were made from very thin hand 

sections, mounted in Hoyer’s medium, and briefly 

boiled to drive out air. Without this approach, it 

is very difficult to determine accurately excipular 

structure. Occasionally, three to four days must 

elapse after mounting a section before all minute 

air bubbles disappear; when present, they may 

mimic carbonization very closely, particularly in 

the labial tips of totally uncarbonized exciples. 

Excip2e Curbonmztion: Carboniiation is not an 

all-or-nothing proposition. Many ascocarps will 

show heavy carbonization in portions of a thallus 

and only partial or slight blackening in others. We 

have maintained several species on the basis of incomplete vs. complete carbonization of the lateral 

exciple walls, but more abundant collections may 

force such taxa into synonymy. 

One odd variant of carbonization appears in 

some graphids. The bases of the exciples are connected by a more or less continuous black to brown 

(partially carbonized) band. Such a condition appears commonly in Graphina insignis, and less fre- 
4 SMITHSOXIAN CONTRIBUTIONS TO BOTASY 

quently in G. uirginea and G. confluens. We have 

found no correlation with any other features; in 

fact, the black band may be quite absent in parts 

of a thallus and present elsewhere. 

Many species of Graphidaceae lack carbonization 

completely. In some of these, the exciple itself 

seems to be non-existent. In many of the fissurine 

Graphis species (i.e., those with fissure-like apothecia in surface view, and four-locular spores), the 

term “exciple” may be inapplicable, as the hymenium would seem to be bounded by unmodified 

hyphae; in many of these, what appear to be 

excipular labia are, in fact, masses of bark cells. 

It may well be that presence of large amounts of 

bark laterally bounding the hymenium is significant as an indicator of endophloeodal development 

of the ascocarp. 

Open us. Closed Exciples: In many specimens 

with carbonized exciples, all sections from a thallus 

will be consistently open (or consistently closed) 

below, but many others will vary from completely 

open to completely closed within the same thallus. 

The weight commonly placed on this character-state 

to establish species, subgenera, or sections is unjustified. \\‘hen no carbonization exists, it becomes 

even more difficult to determine if closure is present, even in a single transverse section. It seems 

likely that many species presently separated only 

by excipular closure will prove synonymous with 

more collections. 

In addition, it may well be that degree of basal 

carbonization, and hence the overall height of the 

ascocarp above the thallus, is a function of age. 

This possibility is discussed more fully below. 

Labial Striae: Another character on which much 

emphasis has been placed, probably incorrectly, is 

the absence, presence, and degree of striation in the 

labia. Two basic types of striae occur in the 

Graphids and must be treated separately. 

In the first type, occurring in both carbonized and 

clear exciples, the surface of the lips is obviously 

indented, forming distinct longitudinal grooves. 

This character is usually quite variable within a 

single thallus; maintaining species on degree of 

striation must be regarded with great suspicion. In 

fact, many specimens with no striations on the 

majority of apothecia can be found to have them 

on a few. Many species-pairs can be found in the 

family where the members are distinguished only 

by presence or absence of longitudinal striae (see, 

for example, Graphis anguilliformis and G. flexib ilis). 

It is possible that the striations in both carbonized and clear exciples may be a function of age. 

Specimens that have high ascocarps with thick bases 

and (frequently) disintegrating hymenia (all presumed to be signs of age) also usually have many 

striae. On the same thallus one can occasionally 

find low ascocarps with less basal carbonization and 

vigorous (fertile) hymenia; these structures are frequently associated with fewer (sometimes no) labial 

striae. In addition, in species with both clear and 

carbonized exciples, new hymenia arise below old 

ones: this phenomenon, which may be seasonal, 

apparently ruptures the excipular tissue below the 

old hymenium, and compresses it into new lateral 

striae. This phenomenon has been pointed out 

earlier for carbonized species (Redinger, 1935: 98). 

In uncarbonized forms, an increase in the number 

of internal striae (described below) occurs; we have 

seen it very frequently in sterile Mexican material 

of cf. Gmphina peplophora (Figure 1). If this interpretation is correct, then the number of striae per 

ascocarp would increase seasonally (at least in these 

species), and hence would be useless taxonomically. 
NUMBER 40 5 

The second major type of striae occurs only in 

uncarbonized exciples. What appear to be dentations or striae occur within the labial tissue, without any trace of surface grooves. Examination of 

many ascocarps on large thalli reveals that these 

striations arise with age, apparently by lateral portions of the hymenium compacting, darkening, and 

being incorporated into the lateral tissue surrounding the hymenium. Along with these compactions, 

the portion of the exciple below the hymenium 

thickens and raises the whole ascocarp farther from 

the thallus surface. Thus, a single specimen may 

have low ascocarps with entire lips and an open 

exciple and high ascocarps with massive bases and 

(internally) striate labia. The cross sections of 

Graphina colliculosa exciples (Figures 8f,g; 9a) illustrate these conditions well. 

Labial Convergence: The degree of convergence 

of labial apices, commonly used to separate species 

and subgenera, is also beset with problems. 

One common occurrence is the rapid spreading 

of otherwise convergent labia of carbonized exciples 

as soon as a moistened ascocarp is sectioned. Such 

species as Graphis candidissima Zahlbruckner and 

G. subamylacea Zahlbruckner will appear to belong 

to Section Phanerographa (intact, spreading carbonized lips) when in fact the spreading is an artifact 

caused by the hymenium swelling rapidly. 

A more serious problem arises in many species 

with uncarbonized exciples. In Phaeographis albida 

(Figures 7a,b) and Graphina colliculosa (Figures 

8f,g; 9a), ascocarps with convergent labia (no disc 

visible) and divergent labia (prominent disc visible) 

may occur on the same thallus. If the two forms 

occur on different specimens, the surface appearance will differ strikingly and result in the description of two species. 

THALLuS.-Very few character-states of importance are derived from the thallus. Color, in general, is not a practical feature, with the possible 

exception of those endophloeodal thalli which frequently signal this condition by a distinctive yellowbrown shade. 

The thalli of some species show a distinct prosoplectenchymatous upper layer. The hyphae seem 

to align perpendicularly to the ascocarps, and hence 

cross-sections of the lirellae almost always exhibit 

characteristic parallel filaments. This condition 

seems to be consistent within species, and may 

prove useful in distinguishing members of certain 

species complexes (see, particularly, the Graphis 

triticea group). 

Very few species of Graphidaceae show highly 

pronounced surface features such as soredia or isidia. More subtle aspects of texture may be constant 

enough for systematic purposes, as in thalli with 

pronounced glossy or powdery surfaces. A lower 

cortex is lacking in the family, with the exception 

of the extremely odd Graphis coriacea Vainio from 

Guadeloupe. 

In all cases, we have found the algal constituents 

of no use for species level distinctions. 

CHEMISTRY.-The Iodine Reaction: In most species of Graphidaceae, water squashes of mature 

spores react with almost any concentration of iodine 

solution and rapidly turn a deep blue-violet. The 

hymenium in most species is iodine negative, i.e., 

absorbs the solution and turns yellow-orange, but 

never violet. Within a species the reactions seem to 

be quite constant; see Phaeographis albida for the 

one exception known to date. 

Many of the “fissurine” Graphis species, and 

many species of Phaeographis, have spores that 

are iodine negative, or oddly and slowly I positive 

(turning red-purple). In a few species (Graphina 

virginea, G. diorygmatoides (Vainio) Zahlbruckner, 

G. confluens (Fee) Mueller Argoviensis, Phaeographis albida), the entire hymenium, or at least 

the epihymenium, turns bright violet with iodine. 

The reaction seems constant enough to make it very 

useful in species identification. It is unclear as to 

what compounds are involved in the reaction. 

Lichen Acids and Other Compounds: In general, 

we have tried to erect and maintain species that 

are chemically homogeneous (for exceptions, see 

Phaeographis exaltata and Graphina platyleuca). 

In many cases, this has meant maintaining species 

pairs (or triplets) entirely on chemical grounds. It 

would appear that the Graphidaceae will have 

many cases where a series of morphologically identical species are separable only by the presence or 

absence of stictic and/or norstictic acids. 

Recent chemical testing shows that more than 

half of the species in the Graphidaceae are TLC 

negative, in other words lacking any phenolic secondary products. The most common lichen substance in the TLC positive remainder is unquestionably norstictic acid. Fairly abundant, though 

not common, are stictic acid and the “quintaria” 

unknowns. Very uncommon are psoromic, lecanoric, 
6 S\IITHSONIAN COKTRIBUTIOSS TO BOTASY 

protocetraric, salazinic acids, and lichexanthone. A 

few species (Phacograpliis haetnatiics, P. cinnabarina (Fee) Mueller Argoviensis, Phneograplzina 

rliyysornq!m (Kacldi) Reclinger, among others) ha1.e 

pigments (anthraquinones?). 

All specimens were tested with normal thin-laTer 

chromatographic techniques using silica gel aluminum-backed LIerck precoated plates. Two solvent 

systems were always used: hexane-ether-formic acid 

and benzene-dioxane-acetic acid. l’isualization was 

accomplished with 10 yo sulfuric acid and heating 

at 110” C. 

A list of the chemicals found and species that 

contain them is given belorv. It should be assumed 

that the chemistry for each species represents that 

of the type material, unless otherwise indicated, 

and all other specimens cited. 

Lecanoric acid: G~-nphis nfzelii 

Norstictic acid: Graphis desquninescens, G. dussii, G. elegnns, 

G. isidiifera, G. librntn, G. lumbricina, Phaegraphis suh- 

/i,<l’iflfl, ~~I~tf~~JtJiit itJ/fi//(iJ ItJJt, G. i/;\fX’l~\O, G. fl/OfT/eilCii 

faccescor\). G. ~~scltiloitllnlr/glt, <;. illbelyllltc~lln, and Phneocollospo,.n 

grit,+> 1, i ~i ii n 1 roi ‘YJ iri ic ii lii > i F 

Sorstictic acid with stictic and constictic acitlr: Grophiiin 

I’rotoce t r:ii-ic xitl: G! npltiiiit plnty leitcn 

Psol-omir acitl: (;rnphi,iir roiii?izbi,in 

“Quintaria” unknowns: Phaeogrnphis exnltatn and Grnphina 

t’irginea 

Salazitiic acid: Gi-ofiitiJra rolliriilosct aiid G. mn~rescrns: G. 

plot! lezicn (accessory) 

Stictic acid: Grnphis iiiitJinstioitlrc, G. iiii\ho!(gii. G. lefitocarpa, G. iriticea, G. turgiduln, Grnphina triphora, G. 

iJirgiJirn, and Phneogrnphinn oscitans 

TLC negative (no phenolic substances present): Grnphis 

n il fi VPT sit, G , /I )I f i nc t ii o \(I , C; . n ~i g ii ilii fo i-iii ic , G . /ICY i b ili c, 

G. glnitcescens, G. grnnimitis, G. huinilis, G. insirliosn, G. 

Zonguln, G. olivncen, G. rigiduln, G. rimulosn, G. siibele- 

~OJ~F, G. sitbnitiduia, G. tnchygrnphn, Phneogrnphis nlhirln, 

P. nrthoiiioides, P. hneniatites, P. mordenii, P. rosen, 

Grnphiiia nclinrii. G. iiiltcrib~iic, G. cnrnroi,iritlii, G. chloroenrfin, G. dimidinta, G. frumentnrin, G. illfiiotn, G. incriistn)is, G. inscnlpfn, G. innrclln, G. nzrdo, G. plziriiporn, 

G. rujopzilidn, G. zwstitoides, Phneogrnphiiiu cnesiopniinosn, P. corinrin, P. difformis, P. eiliottii, and (in part) 

P. exnltntn 

Pigments: Phneographis hnernatitey 

Ecology 

The famiIy Graphidaceae is one of the largest 

groups in the lichens with over 1100 published 

names. One should not be surprised, therefore, to 

find that the species have become adapted to a wide 

range of habitats and often, as in the case of 

Graphis afzclii, seem to have a wide ecological amplitude in both secondary and primary forests. This 

is in contrast to the closely related Thelotremataceae that are highly restricted to undisturbed rain 

forest in Dominica (Hale, 1954) and other tropical 

areas. 

TVe are not able to categorize all the species ecologically even for such a small island as Dominica 

since many are represented by only one or two collections. However, the areas of mid elevation virgin 

rain fore5t generally have the greatest number of 

species. Of the commoner species the following, for 

example, appear to be more or less confined to well 

developed rain forests: Graphis oliuacea (also rarely 

found in elfin forest), Graphinn chlorocarpa, G. 

colliczilosa, and G. illinata. These species are common in and probably restricted to elfin or mossy 

forest: Graph i.s ndpl-e.r.~n, G. n ngzi illiforvn i F, G. 

/zim byicina, G. tachygrapha, G. triticen, Phaeoqrnpliic c~xtil/niii, and P. mordr77ii. These occur 

chiefly in low elevation, disturbed habitats: Graphis 

derqzramescen.c, Gmphina ant illarum, and G. uirginfa. Further notes are given under “Habitat” for 

each species but these should be considered tentative until more collections are made. 

Evolution in the Graphidaceae 

In order to speculate on the evolution of nonstromatoid graphids, several assumptions must be 

accepted. First, one must assume that the group is 

relatively natural, i.e., nionophyletic. Second, since 

data froin genetics and cytology are still lacking, 

one inlist assume that at least some phylogeny can 

lie inferred froin ontogeny. 

.\lchough the pitfalls and iallacies of using ontogen? as ;I source for evolutionary speculation are 

!$-ell known. there seems little choice in the Graphitlaceae. Specifically, stages of spore development 

may supply the only concrete bases yet available. 

The follon-ing four observations on spore ontogeny should be considered: 

1. All Graphid spores start as a one-loculed (onecelled) structure, progress to two locules, then to 

four (rarely three) uniseriate chambers, then to 

multiloculx stages. 

2. Longitudinal spore septae occur only after 

most transverse septae have been laid down. 
NUMBER 40 7 

3. Monosporous asci always (?) begin as plurisporous asci, followed by abortion of most spores 

in their early stages. Monosporous asci occur only 

(?) in Gsaphina and Phaeographina. Four-locular 

spores always occur as a group of eight per ascus; 

most spores with only transverse septae also occur 

as a group of eight per ascus. 

4a. Pigmentation of spores occurs last, after 

spores reach essentially full size and septation. 

4b. Pigmentation is accomplished by thickening 

of walls in all (?) Phaeographis, but in very few 

Phaeographina species. Those Phaeographina species which do have thickened walls also have rather 

irregularly distributed spore locules, and the spores 

are rather small and similar in size to the bulk of 

Phaeographis spores. 

From these observations one might draw the following highly speculative conclusions: 

1. Species with two- or four-loculed spores are 

most like primitive (ancestral) stock, unless other 

data indicate that these forms are reduced or neotenous derivatives. 

2. Muriform-spored species are derived from ancestors with Graphis-like spores. 

3. Monosporous forms are derived from ancestors 

with plurisporous asci. 

4a. Pigmentation may occur at any stage of spore 

septation, but usually represents the last event in 

spore development. Therefore sporomorphs which 

differ only in pigmentation are extremely closely 

related (i.e., one “step” apart). 

4b. As most Phaeographina species do not have 

thick-walled spores, they are probably derived from 

Graphina sporomorphs by added pigmentation, 

rather than from Phaeographis sporomorphs by 

added septation. The less common thick-walled 

Phaeogsaphina species may be derived from irregularly-loculed Graphina ancestors (also rather uncommon) or from Phaeographis-like forms. 

Putting these speculations together in graphic 

form, we propose a very hypothetical scheme of 

evolutionary relationships (Figure 2). 

................................................ GRAPHIS ....... ..... ..a** .* .* : Ancestral stock ..’ spores 2- Iocular, 

.: c~earffew extant 

i G. turbulenta, etc.) 

species, G. coriacea, 

.................. *. 

few- loculed 

thick- wa I led 

monosporous .: ...... Phaeographina .... .... PHAEOGRAPH I NA .................. 

FIGURE 2.-Hypothetical origin and relationships of the genera of the Graphidaceae. 
8 

This scheme immediately gives rise to certain 

questions, problems, and objections, to wit: Where 

does carbonization fit in this scheme? Graphis species with few-loculed spores, supposedly primitive in 

the proposed arrangement, very rarely show any 

carbonization. At the same time, lack of carbonization is also the more common condition in the supposedly most advanced groups (i.e., those with 

monosporous asci and muriform spores). In addition, many of the fissurine Graphis species, supposedly primitive, show an apparently specialized 

(prosoplectenchymatous) upper cortex. One way to 

resolve these contradictions might be to consider 

the carbonized Graphis species with multilocular 

spores as equivalent to an ancestral stock, and to 

regard the fissurine and other forms with very 

simple spores and little carbonization as derived 

(reduced) forms, the spore simplicity perhaps 

arising neotenically. 

Another problem arising from this scheme is the 

admitted artificiality of the present four-genus system for the nonstromatoid taxa. Phaeographina, for 

example, could arise from two fairly disparate 

stocks. 

Most important, perhaps, is to ask what selection 

pressures may be involved in generating these 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

spore states? For example: a fissurine Gjaphzs 

would produce a maximum of four germination 

tubes per spore and hence have a low number of 

tubes in each of many landing sites. A monosporous muriform species produces a great many 

tubes in each of fewer sites. Is this relationship 

significant in terms of colonizing or competing? To 

the best of our knowledge, there are no data on correlations of spore-types and habitat. The fact that 

one can find all the spore variants on a single twig 

does not necessarily mean that there is no competitile advantage associated with the variants. In the 

Ryrsonima scrub of western Mexico, for example, 

all the spore forms occur abundantly, shoulder to 

shoulder. HoweLer, this may be a very equable 

habitat, comparable to the tropical rain forest for 

the angiosperms; other more demanding habitats 

and substrates might re\ eal significant preferences 

and correlations, In addition, the abundance of adjacent forms may be misleading in that a succession 

may be occurring too slowly to be apparent immediately. 

The same lack of data exists with respect to distribution of carbonized vs. uncarbonized forms, 

endo- 1s. epiphloeodal thalli, etc. Clearly, much 

field work remains to be done. 

Key to the Genera of Graphidaceae in Dominica 

(Although all genera of the Graphidaceae that are known to occur in Dominica are keyed 

belois, the txvo stromatoid groups, Glyphis and Snrcogrnphn, are not treated in the text. Each 

is represented by only one species (l’ainio, 1896, 1915.) 

1. Ascocarps embedded in prominently raised, stromatoid tissue. 

2. 

1. Ascocarps not embedded in raised stromatoid tissue. 

3. Spores clear and hyaline, the lvalls usually thin. 

4. Spores with transverse septae only ................................................................................ Gru~~~s 

4. Spores with transverse and longitudinal septae ...................................................... Graphina 

3. Spores brown, the !calls occasionally thickened. 

5. Spores with transverse and longitudinal septae, the Isalls rarely thickened .......................... 

5. Spores with transverse and longitudinal septate, the walls rarely thickened .... 

2. Spores clear, with transverse septae only .......... .......................... Glyphis 

Spores brown, with transverse septae only ................................................................ Sarcograph~~ 

................................................................................................................ .Phaeographis 

....................................................................................................................... .Phaeographina 

Key to the Species of Graphis 

1. Spores always four locular. 

2. .4scocarps fissurine, arising as a thalline crack; disc usually concealed or sunken. 

3. Exciple carbonized. 

4. Exciple heavily carhoniied laterally, spores I+ ........................................ I?. G. hurnilis 
NUMBER 40 9 

nized only at labial tips, spores I- (reddish) ...................... ................................................. 25. G* tachYgraPha 

................................... ...... 11. G. gramrnitis 

.... 

3. Exciple totally uncarbonized. 

5. Ascocarps usually exposing a red-brown disc in 

long ............................ 5. Ascocarps fissurine th 

6. Ascocarps embedded in swollen tissue (“puffs”). 

7. Stictic acid present .................................................. .. , ..26. G. triticea 

.............................. 14. G. insidiosa 

6. Ascocarps not embedded in swollen t 

‘ mmonly to 10 mm 

7. No substances present ................ 

8. Spores less than 11 pm long; no substances present .......... 24. G. subnitidula 

8. Spores more than 13 pm long; stictic acid present ............ 6. G. dumastioides 

9. Ascocarps white-powdery; exciple carbonized; lecanoric acid present .......... 2. G. afxelii 

norstictic acid present .................................................................................. 15. G. isidiifera 

present ...................................................................... 11. G. grammitis 

Exciples red-brown to barely carbonized at labial apices ........................ 10. G. glaucescens. 

2. Ascocarps not fissurine; disc broad, obvious, not depressed within a thalline crack. 

9. Ascocarps not powdery; exciple carbonized; lecanoric acid absent. 

10. Thallus isidiate; thalline margin coarsely white-mealy; asocarps nearly round; 

10. Thallus not isidiate. thalline margin not mealy; ascocarps elongate; no substances 

1. Spores six locular o 

11. 

11. Exciples distinctly carbonized. 

12. Labia distinctly striate. 

13. Exciples with distinct red-yellow areas below the carbonized portions. 

14. Thalline margin prominent; spores over 45 pm long ..... 

14. Thalline margin absent or nearly so; spores under 35 gm . 

times dirty-brown below). 

15. Ascocarps usually without a prominent thalline margin. 

16. 

16. Exciples closed below; no substances present 

15. Ascocarps consistently with a thalline margin. 

17. Norstictic acid present ......................... 17. Norstictic acid absent. 

13. Exciples lacking red-yellow tissue below the carbonized portions (but someExciples open below; norstictic acid present ............. 8. G. elegans 

......... 22. G. rimulosa 

19. G. lumbricina 

18. Exciple open below. 

19. Spores usually 70-90 pm long .............................. 19. Spores usually 50-65 pm long ............................ 

20. Exciple massively carbonized below .................... 9. G. flexibilis 

18. Exciple closed below. 

20. Exciple barely closed below ...... 18. G. Iongula 

12. Labia intact or irregular, not distinctly striate. 

21. Exciples distinctly open below. 

22. Stictic acid only present ............................................... ..... 16. G. Zeptocarpa 

22. Norstictic acid only present. 

23. Spores less than 36 pm long ...................................................... 17. G. Zibrata 

23. Spores over 45 pm long ................................. 

21. Exciples distinctly closed below, or the lateral 

proaching each other very closely basally. 

24. Ascocarps raised, without any thalline margin visible, short (1-2 mm long), 

Opegrapha-like ................................................................................ 1. G. adpressa 

24. Ascocarps flush to raised but always with a thalline margin, usually over 

2 mm long, not Opegrapha-like. 

25. Spores under 51 pm long. 

26. Stictic or norstictic acid present. 

27. Norstictic acid present .................................... 3. G. desquamescens 

27. Stictic acid present ................................................ 13. G. imshaugii 

26. No substances present .................................................. 3. G. anfractuosa 

25. Spores usually over 60pm long. 

28. Stictic acid present ..... 

28. No substances present .... 4. G. anguilliformis 
10 S\IITHSOSIAN COSTRIBLTIOSS TO BOTASY 

1. Graphis adpressa 

FKVRE 90; PLATE la 

Grnphis nrlp).essn I'ainio, 1890: 119 [type collection: Carasqa, 

Braril, l'oinio 1289 (TUR, 1ectot)pe)l. 

~)~.S~:KIP-I~IOZ.--~'~I;I~~L~S it,hite, t?;r:i)., or pile 

brown, thin but continuous, shining. Xscocarps 

quit2 black, raised antl protuberant, usually unbranched, 110 thnlline nixgin 1-isible, 0.5-2 mn 

long, slender. Disc not visible in surface view. Ascocarp transverse section: liymeniuin 170-250 ,uin 

high, I-; exciple black, closed, lips intact and convergent, frequently with an extremely thin thalline 

covering. Spores 8 'ascus, 10-18 locular, 13-15 X 

50-70 pm, I+ blue. 

CHESZISTRY.-NO substances present. 

H.AsrTAT.-Rain forest above 2600 feet. 

DISCL-SSIOS.-GVZ~~~S adpwssa externally resembles an Ojiegmplia or Xrlas/iilea antl is unlikely to 

be confused with any other Dominican Gmphis. It 

exhibits comiderably less variation in morphology 

than most Graphids. Extremely similar but for 

spore septation and size is Gmphina niida. 

Srrcr~rss Ex.&>risrD.-Fresh IVater Lake, 2600-2800 ft, 

Hnle 33485, 3.5487 (US), 2600-2877 ft, Iiushntig 32849.4, 

32852A, 32864 (LiSC); Trois Pitons, 2900-3150 ft, Imshnicg 

33090 (MSC); Riorne Diablotin, 33004300 ft, Iinshaiig 32940 

(NSC); Jlicotrin, ca 3000 ft, TVirth 464 (US). 

2. Graphis afzelii 

FIGURE 3b: PLATE lb 

Grojiliir ajxlii .\charius, 1814:86 [tvpe tollcction: Guinea, 

Afzelizu, s.11. (H, lectotype; UPS, isotype)]. 

I)F~C:RIP~-ION.--T~I;I~~II bi~ownisii tiin to gray, continuous, smooth. Ascocarps usually compIeteIy 

covered by ii poTvdery white thalline layer, appearing black only where this layer is rubbed, usually 

unbranched, 1.0-6.0 mm long, to nearly 1 nm wide. 

Disc not visible in surface view. Ascocarp transverse 

section: hymenium 75-1.50 high, I-; exciple 

black laterally, brown and closed to pale and open 

basally, lips conr.ergent, intact, with a proininent 

thalline covering extending to their apices. Spores 

S'ajcus, always 4 locular, 6-9 x 16-23 ,pn, I+ 

blue. 

C€IE~rIsTRY.-I,ecanoric acid (in the thalline 

apothecial covering only). 

HABITAT.-D~~ sea-level scrub to virgin upland 

rain forest; apparently with broad ecological amplitude. 

DISCU~SIOS\'.-GYU~~~S afzelii is a common pantropic species, unique in the family in having 

lecanoric acid. This acid, along with the distinctive 

exciple structure, render it unlikely to be confused 

with any other graphid. 

SPrcl\rrss ExlMIsEo.-Pointe Round, 100 ft, Im~hntcg 

33484.i (JISC): Londerry Agricultural Area, 100 ft, Imshnug 

SL'B9T (\ISC): iicar Ma(Iii~ii, 100 ft, Hnlr 3,5618 (CS); nnrrtl 

of Rinche, 200 ft, Hole 39644 (US); Can-Dom logging area, 

Se~\-foundland, 800 ft, Hale 35239, 35116. 35447 (US); 8 

mile5 north of l'ont Cnrsi., 1400 ft, Tl'irth ,546 (1-S); Cnn-I)oni 

losgiiis area, Dlenu Croiiitnier, 1600-l700 ft, Hde 35183 (US). 

3. Graphis anfractuosa 

FIGURE 3C; PLATE 1C 

Grnphis niifrcictttorn Eschweiler i)i Martius, 1833:86 [type 

collection: Caitete, 13ruil. s.n. (\I, lcctcit\pc; G, isot\pe)]. 

~~sCRIPTros.-~ha~~us dull white to gray, thin, 

continuous. Ascocarps partially emergent, the labial 

tips appearing as slender, flexuose, black, sparingly 

branched lines 1.0-3.0 min long. Disc not visible 

in surface vieIv. Xscocarp transverse section: 

hynienium (SO-) 90-130 p.m high, I-; exciple black 

laterally, frequently dark brown to black below, 

usually closed, lips more or less intact, convergent. 

Spores S'ascus, 8-12 locular, 9-10 X 40-50 pm, 

I+ blue. 

CHEMISTRY.-KO substances present. 

HAnITAT.-I<ain forest lower branches, ca. 1400 

feet (good specimen); coastal scrub (poor specimen). 

DIscc.ssIos.-The lectotype specimen of Graphis 

anfraciuo.~n is large but apparently without spores; 

the isotype at Geneva is quite sinall but fertile. 

\'cry similar is Graphis desqiiameYCen.7, which differs in having smaller spores and norstictic acid. 

S~rc~\ir\-s Ex.t\rrsED.-Scotts Head, sea lei-el, PT'irth 938 

(US); Emeialtl Pool, ca 1400 ft, Tl'irtlz 504a (US). 

4. Graphis anguilliforniis 

FIGURE 3d; PLATE Id 

Grophis n?igziillifonnis Taylor, 1847:152 [t)pe collection: 

St. \'incent, s.n. (FH, lectotype)]. 

DEscRrPTIos.-Thallus tan to light brown, thick, 

glossy, frequently cracked. Ascocarps large, protuberant, unbranched to sparingly branched, straight 
NUMBER 40 11 

FIGURE J.-Cross sections of apothecia of Graphis: a, G. adpirssa (Vainio 1289); 6, G. afzelii 

(lectotype in H); c, G. anfractuosa (lectotype in G); d, G. anguilliformis (lectotvpe in FH); 

c, G. rlesqucrt,icsce?is (Glnzzon 5082); f, G. dtrmastioides (Fink 723 in MICH). 

to flexuose in the larger lirellae, covered at least 

half by a concolorous thalline margin, the exposed 

labial tips black, 0.5-10 mm long, to nearly 0.5 mm 

wide. Disc not visible in surface view. Ascocarp CHEMISTRY.-NO substances present. 

transverse section: hymenium 190-250 pm high (to HABITAT.-Rain forest above 1500 feet, up into 

375 pm in a few extreme cases), I-; exciple black, 

massive, closed, lips entire, convergent, thalline DISCUSSION.-GYU~~~~ anguilliformis (“anguillaemargin prominent. Spores (2-) 4-6 (-8?)/ascus, 12- 

18 locular, 12-15 X (65) 75-110 (-130) pm, I+ 

blue. 

elfin forest. 
12 SMITHSONIAN COSTRIBUTIOSS TO BOTANY 

formis” in the original description) is the commonest 

graphid in Dominica; we have also seen abundant 

material of this very large species from Puerto Rico, 

Mexico, and Colombia. A number of species are 

quite similar, as follows. 

Opegrapha rhizocola Fee (1824) is probably an 

earlier name for G. anguilliformis but the type (G) 

is sterile and very small. In light of the depauperate 

type and the common usage of Taylor’s name, we 

suggest that the Fee name should be permanently 

dropped. 

Graphis cooperta Zenker (1829) is supposedly a 

synonym of 0. rhizocola (fide Mueller Argoviensis, 

1887), but we have not been able to locate the type. 

Graphis seminuda Mueller Argoviensis (1891) is 

likely to prove a synonym of G. anguilliformis; an 

isotype specimen (US) differs only in the more or 

less open exciple. 

Graphis tumidula (Fee) Sprengel (1827) is quite 

close; it differs in size (hymenium frequently 400 

pm high) and in the very large spores (40 locular, 

to 300 pm long). According to Mueller Argoviensis 

(1887) this species is a synonym of Graphis cinerea 

Fee (1824); however, the type of G. cinerea (G) 

is striate. 

G~aphzs tuigzdula llueller Argoviensis differs in 

having stictic acid. 

Graphis Pexibilis differs only in being somewhat 

smaller, and in having striate labia. 

SPECIMEVS ExAv Ixm.-Can-Dom logging area, Dleau Gommier, 1600-1700 ft, Hale 35154 (US); Can-Dom logging area, 

Pont Cassb, 2000 ft, Hale 35084, 35112 (US); Can-Dom log 

ging area, Brantridge Estate, 1700 ft, Hale 35282 (US); Morne 

Anglais, 300-3600 ft Hale 39326a, 35349, 35351 (US), Ellrott 

sn (TUR); Boiling Lake, 2400-3000 ft, Hale 35759, 35760 

(US): Fresh Water Lake, 2600-2877 ft, Imshaug 32848 (MSC); 

Boeii Lake, 2750 ft, Imshaug 33225 (MSC); Central Forest 

Reserie, 1500 ft, Imshaug 335541 (MSC): Morne 

3000-3500 ft, Imshaug 32955 (MSC); 2200-2600 

38174, 38191 (US). 

Diahlotin, 

ft, Hale 

5. Graphis desquamescens 

FIGURE 3e; PLATE le 

Graphis desquamescens (Fee) Zahlbruckner, 1909: 108. 

Opegrapha desquamescens Fee, 1874:24 [t! pe collection: Brazil, 

Glaziou 5082 (M, lectotype)]. 

DEScRIPTIoN.-Thal1us white to off-white, continuous, slightly roughened and matte. Ascocarps 

flush to emergent, slender, flexuose, black, frequently branched, usually with a low thalline margin, (I-) 2-4 (-6) mm long. Disc barely visible in 

surface view. Ascocarp transverse section: hymenium 

60-70 (-120) pm high, I-; exciple black, more or 

less closed, labia convergent, usually entire but 

barely striate in occasional sections. Spores 8/ascus, 

6-9 locular, 6-8 x (20-) 23-35 (-50) pm, I+, blue. 

CHEMISTRY.-NorstiCtiC acid (crystal test only for 

the lectotype). 

HABITAT.-Dry scrub from sea level to 700 feet. 

DIscussIoN.-This species occurs throughout the 

Neotropics; it has been reported (probably correctly) from Japan by Nakanishi (1966). Very similar to Graphis desquamescens is G. intricata Fee 

(1824), separable only by the smaller spores (less 

than 13 pm long) in the latter. Also similar is G. 

anfractuosa, which lacks norstictic acid and has 

somewhat larger spores. 

SPECIMENS ExAsIIsED.-Rodney’s Rock, sea level, Hale 

39581 (US); north of Mero, 50 ft, Hale 35067, 35449, 35727 

(US); Grande SaJ anee, 200-290 ft, Imshaug 33285A, 33291A 

(MSC); hlt. St. Mary’s, 600-700 ft, Imshaug 33448.4 (MSC). 

6. Graphis dumastioides 

FIGURE 3f; PLATE If 

Graphis dumastioides Fink, 1927:213 [type collection: Mameyes, Puerto Rico, Fznk 723 (MICH, lectotype; FH, US, 

isotypes)]. 

DEscRrPTIou.-Thallus brown to greenish tan, 

glossy, continuous, rather thick for a fissurine species (to 150 pm), with a distinct prosoplectenchymatous upper layer. Ascocarps appear as flexuous, 

unbranched to sparingly branched fissures in the 

thallus, the margins usually not concolorous with 

the rest of the thallus, 0.5-4.0 mm long. Disc 

sunken and usually not easily visible in surface 

view. Ascocarp transverse section: 11) menium 60- 

100 pm high, I-; exciple rudimentary at best, the 

lips consisting of barely modified uncarbonized 

thalline tissue with included bark cells. Spores 

Slascus, always 4 locular, (6-) 8-9 X (14-) 18-25 

pm, I- or I+ slow, faint and reddish. 

CHEMISTRY.-stiCtiC acid. 

HABITAT.-Rain forest. 

DIscLssIou.-For a discussion of the fi55Ul ine 

complex of species, see Graphis triticea. 

SPECIVE~ EXA\II\ED -Road to Morne Jean, 2000-2400 ft, 

Hale 37722 (US). 
NUMBER 40 13 

7. Graphis cf. dussii 

FIGURE 4a; PLATE 2a 

Graphis dussii Vainio, 1899:259 [type collection: Guadeloupe, 

Phaeographis dussii (Vainio) Zahlbruckner, 1923:371. 

Duss 515 (TUR, lectotype)] 

DrscRIPTIoN.-Tha~hs off-white, thick, glossy. 

Ascocarps in type oryzaeform only, mostly 0.5 X 1.0 

mm, in Dominican specimen to 3.0 mm long, both 

usually unbranched and more or less straight, with 

a low thalline margin. Disc usually not visible in 

surface view. Ascocarp transverse section: hymenium 

140-150 pm high, I-; exciple black laterally, open 

below, lips convergent, entirely to barely crenate. 

Spores 6-8/ascus, (lo-) 12-16 locular, 6-10 X 60- 

100 pm (46 pm long fide Vainio, but none so short 

found on the type), I+ blue. 

CHEMISTRY.-Norstictic acid. 

HABITAT.-upland rain forest. 

DIscussIoN.-Although Vainio described the 

spores of Graphis dussii as darkened, we can find 

only typical Graphis spores in the type. 

On the type specimen, only short oryzaeform 

Iirellae are present; the Dominican material has 

mostly longer ascocarps, with very few as short as 

in the type. As apothecial length is rather variable 

in the family, we are assuming that additional collections will bridge the gap between these two 

specimens. We are thus using the name to cover the 

largest specimens, with large spores and norstictic 

acid, of the Eugraphis alliance. For a more complete discussion of this complex group, see under 

Graphis leptocarpa. 

35443 (US). 

SPECIMEN EXAMINED.-Morne Diablotin, 32004600 ft, Hale 

8. Graphis elegans 

FIGURE 4b; PLATE 2b 

Graphis elegans (Smith) Acharius, 1814:85. 

Opegrapha elegans Smith in Smith and Sowerby, 1807:16 

[type collection: Britain (BM); only a cross-section of the 

BM specimen seen]. 

DEscRIPTIoN.-Thallus grayish white, continuous, 

glossy to slightly roughened. Ascocarps black, raised, 

flexuous, usually unbranched, thalline margin 

usually absent, striae distinctly visible from above, 

2.0-6.0 mm long, less than 0.5 mm wide. Disc not 

visible in surface view. Ascocarp transverse section: 

hymenium 90-100 pm high, I-; exciple black, more 

or less open below, lips convergent, from barely to 

very striate (on same specimen). Spores 8/ascus, 

9-12 locular, 10-11 X 50-60 pm, I+ blue. 

CHEMISTRY.-NOIS tictic acid (see discussion). 

HABITAT.-secondary and mossy forests above 

1800 feet. 

DIscussIoN.-Without having seen the type of 

Graphis elegans, the identification of the Dominican material is somewhat uncertain, but our material matches the “classic” concept of the European 

species very closely. In addition, at least some of the 

specimens from northern Europe also contain norstictic acid. Specimens are also recorded from New 

Zealand (Hayward, 1978) that match both the 

Dominican and European material very closely. If 

these identifications are all correct, then G. elegans 

must be a cosmopolitan species. 

Closely related to G. elegans are the species of 

the G. rimulosa complex, from which the former is 

distinguished by larger spores and the presence of 

norstictic acid. 

SPECIMENS EXAMINED.-Giraudel, 1800-2000 ft, Hale 35476 

(US); Fresh Water Lake, 2600-2800 ft, Hale 35325 (US). 

9. Graphis flexibilis 

FIGURE 4c; PLATE 2c 

Graphis fiexibilis Krempelhuber, 1876:414 [type collection: 

Brazil, Glariou 5106 (M, lectotype)]. 

DEscRIPTIoN.-Thallus gray to white, usualiy 

smooth and glossy, occasionally somewhat roughened and cracked, thick, cortex prosoplectenchymatous or nearly so. Ascocarps prominently raised, 

mostly flexuose, occasionally branched, black and 

obviously striate above, half to nearly completely 

covered by a thalline margin, 2.0-4.0 mm long 

(commonly longer in non-Dominican material), 

slender. Disc not visible in surface view. Ascocarp 

transverse sections: hymenium (50-) 100-150 pm 

high, I-; proportionately small for the massively 

carbonized exciple (ca. 300 pm high), exciple always 

quite closed below, lips convergent, with 14 striae. 

Spores 6-8/ascus, (15-) 20-30 locular, 13-16 X 60- 

150 (-190) pm (60-110 pm in the type specimen), 

I + blue. 

CHEMISTRY.-TYpe, P-, no TLC made; Dominican material, no substances present. 
14 SXlITHSOSIAS COSTRIBUTIONS TO BOTASY 

b 

FIGURE 4.-Cross sections of apothecia of Graphis: a, G. diissii (D14ss 515); 6, G. eiegam (lectotype 

in BJI); c, G. fiexibilk (Glaziou 5016): d, G. glnucrscetis (lectotype in C); f, G. imshnugii 

(I?nsiiaug 32721B); g, G. hutnilis (,Iferrill 9067); ir, G. hu?tii!is (Hale 35698). 
NUMBER 40 15 

HABITAT.-BelOW the lower edges of the wet rain 

forest. 

DIscussIoN.-Externally Graphis fiexibizis strongly 

resembles G. anguilliformis and Graphina acharii 

and can be considered a sporomorph of the latter. 

Many other species must be treated in any discussion of G. fiexibilis, as follows. 

Graphis angustata Eschweiler (1833) is sup. 

posedly an earlier name for G. fiexibizis (fide Zahlbruckner, 1923); however, the type (M!) has apparently always been found to be sterile (see 

Mueller Argoviensis 1888b3509). We could find no 

spores on the type, and hence this name should be 

discarded permanently. Graphis angustata var. 

denudalum Vainio (1915) has a proportionally 

small hymenium, nearly intact lips, contains an 

unknown substance, and is hence quite dissimilar 

to the “classic” concept of G. angustata; G. angustata var. ingarum Vainio (1915), also unrelated, 

belongs in the Etigraphis alliance. 

Graphis cinerea Fee (1824) is very similar in 

morphology and anatomy; although the type now 

seems to be sterile, Mueller Argoviensis (1887) 

found very large spores (to 250 pm), with only 1-3 

per ascus. For the moment, this species can thus be 

separated from G. fiexibizis by its spores. 

Graphis calcea (Fee) Massalongo (1853) can tentatively be separated by its lower ascocarps (nearly 

flush) and fewer striae. 

Graphis congesta (Fee) Mueller Argoviensis (1887) 

is much like G. lumbricina (below), but differs in 

its shorter, asteroidly-branched ascocarps. 

Graphis longula differs in being more sunken, 

smaller, and in having a nearly open exciple. 

Graphis Zumbricina differs in the presence of 

norstictic acid and in the occasionally open exciple. 

Graphis oZivacea is separable only by the paler 

red-yellow excipular base. 

Graphis Yigidula can tentatively be separated by 

its open exciple and smaller size. 

Graphis tumidzilella Fink (1927) is separable 

only by its smaller spores (under 50 pm). 

SPECIMENS EXAMINED.-Bois Serpe, ca. 1000 ft, Itnshaug 

32772A (MSC); Ridgefield Estate, 1100-1200 ft, Imshaug 

33360 (MSC): Shawford Estate, Elliott 1854 p.p. (TUR). 

10. Graphis glaucescens 

FIGURE 4d; PLATE 2d 

Graphis glaucescens Fee 1824:3G [type collection: South 

America, s.n. (G, lectotype)]. 

DEscRIPTIoN.-Thallus white, continuous, minutely roughened. Ascocarps concolorous with the 

thallus, more or less black only where the thalline 

covering is rubbed, slightly raised, branched and 

flexuose, 1 .O-3.0 mm long, slender. Disc occasionally 

visible as a slightly darker central area. Ascocarp 

transverse section: hymenium 75-100 pm high, I-, 

epihymenium somewhat darkened; exciple more or 

less open, lips more or less convergent, apically 

red-brown to barely carbonized, paler below. Spores 

4-) 6 (-8)/ascus, 6-12 locular, 7-12 x (35-) 42-55 

pm, I+ blue. 

CHEMISTRY.-NO substances present. 

HABITAT.-Lowland cultivated areas. 

DIscussIoN.-Graphis caesioglauca Redinger 

(1 935) is indistinguishable, externally and internally, from G. glaucescens. Transverse sections of the 

type (Malme 1526, S) indicate the same labial structure, and the Redinger illustrations are incorrect. 

We have not yet determined the chemistry of G. 

caesioglauca. 

shaug 33313A (MSC). 

SPECIMENS EXAMIxED.-Brookhill Estate, 100-150 ft, Im1 1. Graphis grammitis 

PLATE 2e 

Graphis grainrnitis Fee 1824:47 [type collection: South America, s.n. (G, lectotype)]. 

DEscRIP’rIoN.-Thallus thick, prosoplectenchymatous, continuous, gray to greenish brown, usually 

glossy. Apothecia nearly fissurine at first, then 

strongly gaping, but with margins not separating 

laterally from the disc or hymenium as in “typical” 

fissurine species, occasionally branched, flexuose, 

commonly 10 mm long, slender. Disc exposed in 

at least a portion of the ascocarps, red-brown. Ascocarp transverse section: hymenium 60 pm high (35- 

40 in the type), epihymenium usually darkened, I-; 

exciple barely closed, yellow-brown to red-brown, 

lips convergent to strongly spreading, apically sometimes darkened. Spores 8/ascus, always 4 locular, 

4-8 X 10-14 pm, I+ blue. 

CHEMISTRY.-NO substances present. 

HABITAT.-Elfin forest and upland rain forest 

(Dominica only). 

DrscussIor\;.-In early stages, the ascocarps of 

Graphis grammitis are quite fissurine, and appear 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 16 

to be a nongaping equivalent of G. dumastii. (The 

type, which is quite poor, consists mostly of this 

fissurine condition), Later stages, however, develop 

rather prominent margins and appear very much 

like Graphina colliculosa externally. 

Quite similar in morphology and anatomy is 

Graphis floridana Tuckerman (1888); the latter species, however, has norstictic acid and probably has 

submuriform spores (and hence is referable to 

Graphina). 

SPECIMENS EXAMINED.-LayOu Road, northwest of Pont 

CassC, 1400 ft, Hale 38005 (US): Elfin woodland, Boeri Lake, 

2750 ft, Imshaug 33208 (MSC). 

12. Graphis humilis 

FIGURE 4h; PLATE 2j 

Graphis hurnilis l’ainio, 1921:256 [type collection: Philippines, Merrill 9067 (TUR, not seen; US, isotype)]. 

DEscRIPTIoN.-Thallus dirty white to pale tan, 

thin (endophloeodal?), continuous, matte. Ascocarps fissurine, barely raised, mostly straight and 

unbranched, the dark exciples showing through a 

thin thalline covering, 0.5-2.0 mm long. Disc not 

visible in a surface view. Ascocarp transverse section: hymenium 60-75 pm high, I-; exciple quite 

open (rudimentary) basally, lips convergent, carbonized apically only (type specimen) to apically 

and laterall). Spores 8,’acus, alwa) s 4 locular, 7-9 

x 14-25 pm, I+ blue. 

CHEMISTRY.-NO substances present. 

HABITAT.-wet lowland and coastal forest. 

DIscussIox.-The Dominican material matches 

the Philippine isotype closely, differing only in having somewhat larger spores and more carbonized 

labia. The carbonization makes this species distinct 

in the fissurine alliance; the closest relative is probably Graphis tachygrapha, which has less carbonization, and I- spores. A more complete discussion of 

the complex alliance of fissurine Graphis species 

may be found under G. triticea. 

A sporomorph of G. humilis (with I- spores, however) is Phaeographis arthonioides. 

SPECIME\ ExA\.rIuED.-I\.Iadjiiii, 100 ft, Hale 35698 (Us) 

13. Graphis imshaugii, new species 

FIGURE 4f; PLATE 3a 

DEscRIPTIoN.-Tha~lus continuus, laevis, prosoplectenchymatus. Xpothecia subiinmersa vel semiemergentia, flexuosa, ramulosa, 24 mm longa, 

margine thallino crassiusculo, cum thallo concolore, 

ab eo fissura tenuissima separato. Excipulum vulgo 

integrum vel subintegrum, fuligineum, labiis integris, erectis; hymenium 120-140 pm altum, I-. Asci 

8 spori; sporae decolores, 8-10 loculares, 6-9 X 

(17-) 23-35 pin, I + coeruleae. 

Thallus continuous, thick, glossy, green-gray, cortex prosoplectenchymatous. Ascocarps usually 

raised (in type) to nearly flush (in the Mexican 

specimen), flexuose, commonly branched, 2-4 mm 

long, slender, the prominent thalline margin commonly separating from the exciple, which is black 

and pruinose. Disc barely visible in surface view. 

Ascocarp transverse section: hymenium 120-140 pm 

high, I-; exciple black, closed to nearly open, lips 

intact to rarely slightly striate, upright to somewhat 

spreading. Spores 8iascus, 8-10 locular, 6-8 X 

(17-) 25-35 pm, I+ blue. 

CHEMISTRY.-stictic acid, constictic acid. 

HoLoTYPE.-Cultivated area, South Chiltern 

Estate Road, 1500-1700 feet, Imshaug 32i24B 

DIscussIo;u.-Graphis imshaugii is closely related 

to G. subamylacea Zahlbruckner (1921), from hlexico. The former species differs in the glossy (not 

farinulose) thallus, more robust ascocarps, and in 

the distinct fissure which separates the exciple and 

the thalline margin. 

Pringle 412 (Tamaulipas, Tampico, Mexico, 

MICH), previously identified by us (Wirth and 

Hale, 1963) as G. subamylacea, is properly referable 

to G. imshaugii. 

(RISC). 

14. Graphis insidiosa 

FIGURE 5,; PLATE 3b 

GwphiA i,z,itiio.tn (Knight aiid Mitten) Hooker El., 1867:586. 

Fissurina insidiosa Knight and Mitten, 1860: 102 [type collection: h’ew Zealand, Knight 259 (BM, lectotype designated 

by Ha)ward, 1978)l. 

Graphis beaurnontii Tuckerman, 1888:124 [type collection: 

Alabama, USA, Beaumont 324 (FH, lectotype)]. 

Graphis inteiversa SJlander, 1889:49 [type collection: Florida, USA, Eckfeldt 339 (H, lectotype)]. 

Graphis lactea var. clazisn Vninio, 1919:163 [type collection: 

Soufrikre, Dominica, Elliott 151 1 (TUR, lectotype)]. 

DEscRrPTIoN.-Thallus greenish to green-tan, 

usually thick and cracked, glossy but with frequent 
NUMBER 40 17 

, 

FIGURE 5.-Cross sections of apothecia of Graphis: a, G. insidiosa (Knight 259); b, G. isidiifera 

(Hale 35179); c, G. lefitocarfia (lectotype in G); d, G. librata (Knight s.n.); e, G. longula 

(Glaziou 5497); f, G. lumbricina (Duss 1036); g, G. olivacea (Malme 2267); h, G. rigidula (Pittiers s.n.). 
18 SlIITHSOSIAS COSTRIBUTIOSS TO BOTASY 

bumps and rugosities, in section showing a poorly 

developed prosoplectenchyinatous cortex. Xscocarps 

starting as fissurine cracks, usually developing 

raised and swollen concolorous thalline margins, 

usually unbranched, 1-4 mm long. Disc usually 

not visible in surface view. Ascocarp transverse section: liymeniuin 90-140 pm high, I-; exciple pale, 

open, lips convergent to slightly gaping, pale to 

barely darkened apically, rudimentary below. 

Spores 8 ascus, always 4 locular, (5-) 8-1 1 x (1 I-) 

13-22 (-24) pm, I- or I+ slow, reddish. 

CHEMISTRY.--NO substances present in all except 

Hale 35313, which has 2-3 unknown spots in TLC. 

HABITAT.-\Vet lowland forest (US), wet upland 

forest (Dominica). 

DIscr-ssIo;u.-Examination of the types of Graphis 

insidiosa, G. beaunzontii and G. interversa indicates 

that these three taxa are identical. The type of C;. 

lactea var. clazisa differs in lacking the cracking of 

the thallus and in having rather shorter, smaller 

ascocarps. The nearly continuous, uncracked thallus 

is also found in the Hale and TVirth collections; 

we feel, however, that these differences ’are too 

minor to warrant naming. 

Grapkis insidiosa is most similar to G. triticea, 

froin which it differs in lacking stictic acid, having 

lower ascocarps with a less elaborate exciple, and in 

a less prominently prosoplectench~inatous cortex. 

The two species are, however, quite similar and are 

most easily separated by their chemistry. 

A more complete discussion of the fissurine complex ma): be found under Graphis friticea. 

SPECIMESS Ex4vlsro.-Can-Dom logging area, Dleau Gommier, 1600-1T00 ft, Hale 33313 (US); hlicotrin, ca 3000 ft, 

T’l’irth 463.1 (US). 

15. Graphis isidiifera, new species 

FIGURE 5b; PLATE 3c 

DesCRtPTIos.-Thallus laevis, isidiatu5; StratUni 

corticale prosoplectenchymatum. Apothecia immersa, rotunda vel sublirellina, ca. 1 inn1 longa, 0.5 

min lata, margine thallino, crasso, pallido, farinuloso, discum bene superante, disco dilatato, salmoneo. Excipuluin integrum, pallidum, labiis 

integris, divergentibus; hymeniuin 70-80 pm altum, 

I-. Asci 8 spori; sporae decolores, 4 loculares, 4-6 

x 12-13 pm, I-. 

Th ;I 1 1 LI i t 11 i c k, cl-;I c ke (1, gl o 5s y , pale greenish 

gray, distinctly isidiate, cortex prosoplectenchymatous. Ascocarps with prominent white mealy margins, nearly round to quite irregular, clumped and 

branched, ca. 1 mm long, 0.5 mm wide. Disc quite 

exposed, pale salmon-pink, lightly pruinose, Ascocarp transverse section: hymeniuin 70-80 pm high, 

I-; exciple closed, pale, lips entire, spreading. 

Spores 8/ascus, always 4-locular, 4-6 X 12-13 

pn, I-. 

CHE\zIsiRY.-Sorstictic acid, unknown substance. 

HoLo?-YPE.-\Tirgin upland rain forest, Can-Dom 

logging area at Dleau Gommier, 1600-1700 feet, 

Hale 35179 (US). 

Drscuss~os.-Grapkis isidiifera is unlikely to be 

confused with any other Graplzis species. The combination of isidia, cracked prosoplectenchymatous 

thallus, white-margined irregular ascocarps and 

flesh colored disc render this species quite distinct. 

It is probably related to the fissurine Gmphis species, but differs from them in the very broad disc 

and prominent mealy margin. 

Isidia are very rare in the Graphidaceae; to the 

best of our knowledge, the only other tuberculateisidiate species occur in Grapkino, (\,iz., Gw,b/tina 

dirt7orplrorles (Sylaiidei-) Zahlbrucknel-, 1923), 

Graphina dea 1 bata (Nylander) Mueller Argoviensis 

(1895a), and its close allies G. albostriata (Vainio) 

Zahlbruckner (1 923), G. heteroplacoides Redinger 

(1933), G. rinizilosa Redinger (1933), and Phaeographina (= Graphina!) includens (Vainio) Zahlbruckner (1923). 

16. Graphis leptocarpa 

FICLRE 5c; PLATE 9d 

Graphis leptocarpa FCe, 1824:36 [type collection: South 

America, Humboldt and Bonpland s.n. (G, lectotype)]. 

DEscRrPTIoN.-Thallus white to off-white, continuous, smooth to slightly roughened. Ascocarps 

raised, black, unbranched to occasionally branched, 

straight to somewhat flexuose, with a slightly raised 

thalline margin, 1-3 mni long, slender. Disc not 

visible in surface view. Ascocarp transverse section: 

hymeniuin 90-100 pin high, I-; exciple black laterally, absent below, lips more or less convergent. 

Spores 8 ascus, 8-9 locular, 5-8 x 16-23 pm (in 

the tjpe) to 42 pin (in the Dominican and other 

material), I + blue. 

CHEhIISTRY.--StiCtiC acid (and usually constictic 

acid). 
NUMBER 40 19 

HABITAT.-FrOm sea level scrub to rain forest. 

Probably pantropical. 

DIscussIoN.-The Eugraphis alliance is one of 

the most frustrating species complexes in the family. The type species of the Graphidaceae, Graphis 

scripta, belongs here; unfortunately this species has 

not yet been typified properly. In addition, as G. 

scripta is probably a north temperate population, 

we have refrained from calling any tropical material by this name. 

Of the numerous names published before 1900, 

the following must be considered in any discussion 

of tropical eugraphids (i.e., those with dimidiate 

carbonized exciples and spores in the 20-40 pm 

range): 

Graphis lineola Acharius (1810) is probably the 

oldest and best name for those tropical eugraphids 

which are P- (and presumably TLC negative, although as far as we know the type has not yet been 

chromatographed). 

Graphis tenella Acharius (1814), another P negative species, may not belong in this complex at all, 

even though the “classic” concept of the species 

places it here. Type material of G. tenella at Helsinki has striate labia; if all typical material is 

striate, then the historical concept of the species 

is incorrect. 

Graphis furcata Fee (1824), which is TLC negative, is probably s)-nonymous with G. lineoln. 

Graphis pavoniana FPe (1824), another TLC 

negative type, may be distinct from G. lineola in 

being larger overall, with only 4-6 larger spores (to 

50 pin) per a5cus. 

Graphis libl-ata is the norstictic equivalent of G. 

lineola (P-) and G. leptocal-pa (stictic acid). 

Graphis caesiella Vainio (1890), with norstictic 

acid, differs from G. Zibrata primarily in having 

pruinose ascocarps and discs. 

Graphis dussii (above), with norstictic acid, differs from G. librata in overall size and in having 

larger spores. 

SPrCI\rr:ss ExA~ilsED.-Pointe Michel, 200 ft, Zrushntrg 

33109.4 (MSC); Morne Conliabon, Elliott 1533 (TUR). 

17. Graphis librata 

FIGURE 5d; PLATE 3e 

Graphis libratn Knight, 1884:404 [type collection: New Zealand, Knight 67:23 (WELT, lectotype designated by Hayward, 1978)l. 

Graphis crebra Vainio, 1899:256 [type collection: Guadeloupe, 

Graphis plzirnierae Vainio, 1915:161 [type collection: GuadeGraphis tenellula 1.ainio 1915:169 [tjpe collection: Santo 

Duss 541 (TUR, lectotype)]. 

loupe, Duss 1189 (TUR, lectotype)]. 

Domingo, Raunkiaer 490 p.p. (TUR, lectotype)]. 

DEscRIPTroN.-Thallus gray-white, continuous to 

matte. Ascocarps quite variable even on a single 

thallus, usually more or less raised, straight to 

flexuose, commonly branched, black, the thalline 

margin prominent but low, 1-4 mm long, slender. 

Disc not visible in surface view. Ascocarp transverse 

section: hymenium 50-100 pm high, I-; exciple 

quite open below, lips carbonized, entire, more or 

less convergent; spores 8/ascus, 6-8 locular, 5-9 X 

18-35 pm, I + blue. 

CHEMISTRY.-NOrStiCtiC acid. 

HABITAT.-coastal scrub to lower rain forest. 

DISCL!SSION.-G~-U~~~~ librata is the norstictic 

equivalent of Graphis leptocarpa; for a more complete discussion of the tropical Eugraphis complex, 

see the latter species. 

SPECIMENS ExAMInED.-Roseau Botanic Garden, ca 100 ft, 

Wirth 468 (US): Pointe Michel, 200 ft, Imshaug 33112A 

(MSC); Coulibistri, 200 ft, Hale 35757 (US); Emerald Pool, 

Wi7th 504B (US). 

18. Graphis longula 

FIGURE 5e; PLATE 3f 

Graphis longtila Krempelhuber, 1876:414 [type collection: 

Graphis flavicnns Mueller Argoviensis, 1893a:457 [type collecPhueographis longtila (Krempelhuber) Zahlbi-uckner, 1923: 

Brazil, G[azioti 5497 (If, lectotype)]. 

tion: Rio, Brazil, Glaziou 5498 p,p. (M, lectotype)]. 

379. 

Z)ESCRIPTI~~..-T~~~~IUS white, grenisli Tvliite, 01’ 

gray, continuous, smooth and glossy to slightly 

roughened and bumpy. Ascocarps more or less 

emergent, black apically, half to two-thirds covered 

by the prominent thalline margin, usually straight 

and unbranched, obviously striate, 1-6 (-10) mm 

long, slender. Disc not visible in surface view. Ascocarp transverse section: hymenium 60-90 pm high, 

I-; exciple black, more or less closed to barely open 

(variable even within a single thallus), lips convergent, striate (appearing only crenate when the 

grooves are abundant and closely packed). Spores 

(?4-) 6-8jascus, (8-) 10-17 locular, 9-13 X (40-) 

70-90 pm, I+ blue. 
20 SMITHSOSIAS COSTRIBCTIOSS TO BOTASY 

CHEMISTRY.-Type of Graphis longula P-, no 

TLC available; type of G. flavicans and Dominican 

specimens, no substances present. 

HABITAT.-LOWer edges of rain forest. 

DIscvssIos.-The types of Graphis longula and 

G. flauicans are extremely similar; note the consecutive collector’s numbers. 

The exciple cross-section illustrated for the type 

of G. longula in IVirth and Hale (1963) is misleading; thinner sections reveal that the barely crenate 

appearance is, in fact, a result of a very closely 

packed deep striae (an excellent example of the 

pitfalls awaiting the graphidologist). 

We have seen material referable to G. longula 

from Mexico and Costa Rica, indicating that it will 

probably be found through most of the Neotropics. 

Very similar (and perhaps synonymous) is Graphis 

rigidula (p. 21), Tvhich is tentatively separated by 

its smaller overall size and somewhat smaller spores. 

Also similar are many of the species that center 

around Graphis flexibilis; a more complete discussion may be found under the latter. Graphis 

fiexibilis itself is separable by its massively closed 

exciple and larger more protuberant ascocarps. 

The Graphis rirnulosa group differs in having 

smaller spores and more emergent ascocarps with 

no thalline margin. 

SPECIJIESS EXAuIsEn.-Morne Bruce, 400 ft, Inishaug 33243 

(MSC); Can-Dom logging area, Sewfoundland, 800 ft, Hale 

33229 (US). 

19. Graphis lumbricina 

FIGLRE 5f; PLATE 4a 

Graphis lurn6ricina l’ainio, 1899:256 [tlpe collection: Cuadeloupe, Duss 1036 (TUR, lectotype; FH, isotjpe)]. 

DEscRrp?Io,u.-Thallus gray, continuous, smooth. 

Ascocarps at first sunken, nearly nonstriate, then 

striate, protuberant and large, apically black, 

mostly covered by a prominent thalline margin, 

occasionally branched, flexuose, to 5 mm long, less 

than 0.4 mm wide. No disc visible in surface view. 

Ascocarp transverse section: hymenium (100-) IiO200 pm high, I-; exciple black, closed to brown 

below and nearly open, lips convergent, distinctly 

striate. Spores 8/ascus, 10-16 locular, 12-20 x (50-) 

75-100 (120) pm, I+ bIue. 

CHEhlISTRY.-h70rStiCtiC acid (isotype). 

HABITAT.-Rain forest, mossy forest. 

DIscussIoh-.-Graphis lum bricina is very similar 

to G. flexibilis; the latter differs in a more massively 

carbonized excipular base and in lacking norstictic 

acid. Graphis congesta (Fee) hlueller Argoviensis 

(1887) differs only in having short, congested, 

asteroidly-branched ascocarps. For a more complete 

discussion of this alliance, see Graphis flexibilis. 

SPECIMENS Ex.~hrI\En.-Can-Dom logging area, xerifoundland, 800 ft, Hale 33210 (US); Fresh lVater Lake, 2600-2800 

ft, Hale 33449 (US); Tmis Pitons, 29-3150 ft, Imskniig 

?;3084A, 33087 (XISC). 

20. Graphis olivacea 

FIGURE 5g; PLATE 4b 

Graphis olivacea Redinger, 1933:53 [type collection: hfatto 

Grosso, Brazil, Naljne 226SB (S, lectotype)]. 

DEsCRIPTIoh-..-Thallus pale olive to whitish, 

smooth, glossy, continuous to rather eroded, and 

restricted to near the ascocarps in the high elevation specimens. Ascocarps straight to flexuose, occasionally branched, apically black and striate (at 

least in the largest ant1 oldest lirellae), frequently 

with pale streaks of thalline material between the 

striae, the thalline margin prominent, 1-6 mm 

long, slender. Disc not visible in surface view. Ascocarp transverse section: hymenium 130-200 pm 

high, I-; exciple black above, yellow-red to brownred below, usually closed, lips convergent, striate. 

Spores 6-8/ascus, 10-18 locular, 8-10 (-13) X (45) 

60-90 (-125) pm, If blue. 

CHEMISTRY.-~O substances present. 

HABITAT.--T\‘et rain forest into elfin forest, where 

it is very coninion on Clusia uenenosa with Phaeographis exnltata and P. mordenii. 

D~scuss~o~.-Graphis olivacea is very similar to 

G. ltcxibilis, from which it differs in having the excipular base constructed of partially uncarbonized, 

yellow- to red-brolvn tissue. It may well be that this 

difference is not significant. Also quite similar is 

For a more complete discussion of this alliance, see 

G. flexibilis. 

Graphis olivacea may prove to be a synonym of 

the Philippine G. glauconigrn L‘ainio (1921), which 

appears to differ only in the smaller ascocarps and 

brown thallus. 

c J. 1 : lgidula, ’ which differs in being smaller overall. 

Sr~c~sr~si Esisiism-Can-Dom logging area, Se\c.foundland, 800 ft, Hale 35062 (LTS); Dleau Gommier Forest Reserve, 
NUMBER 40 21 

1200 ft, Hale 37988 [US); South Chiltern Estate Road, 1500- 

1700 ft, Imshaug 32730 (MSC); Soufriere Ridge, 2200-3100 ft, 

lmskaug 33069A [MSC); Trois Pitons, 2900-3150 ft, Imshaug 

33089 (MSC), 4000-4672 ft, Irnshaug 32874A [MSC), summit, 

Wirth 487, 491 (US); Morne Diablotin, 4300-4550 ft, Imshaug 

32904A, 32909 (MSC). 

21. Graphis rigidula 

FIGURE 5h; PLATE 4c 

Graphis rigidula Mueller Arg0.i iensis, 1891 :58 [type collection: San Jose, Costa Rica, Pittier 5291 (G, lectotype)], 

DEscRIPTIoN.-Thahs white to gray, continuous, 

nearly glossy to minutely roughened. Ascocarps 

black above, flexuose, occasionally branched, covered 

at least halfway by the thalline margin (frequently 

covered nearly to the apex in the type), sometimes 

showing white thalline streaks between the black 

striae, 2-5 mm long, slender. Disc not visible in 

surface view. Ascocarp transverse section: hymenium 

60-90 pm high, I-; exciple black, open below, lips 

shallowly striate, thalline margin prominent in all 

sections. Spores 4-6 (-8?)/ascus, 10-15 locular, 10- 

12 x 50-65 pm, I+ blue. 

CHEMISTRY.--NO substances present (? trace of 

norstictic acid in the type?). 

HABITAT.-secondary forest. 

DIscussIoN.-Graphis rigidula is extremely similar to G. Zongula; it is tentatively maintained here 

because of its smaller overall size and slightly 

smaller spores. For a more complete discussion of 

this species complex, see Graphis flexibilis. 

(MSC). 

SPECIVEN EXAMI.\IED.-BOiS kpe, 1000 ft, ImshUUg 32777A 

22. Graphis rimulosa 

FIGURE 6a; PLATE 4d 

Graphis riniulosa (Montagne) Trevisan, 1853:ll. 

Opegrapha rimulosa Montagne, 1842:271 [type collection: 

Guyana, Lefirieur 200 (P, lectotype)]. 

DEscR:PTIoN.-Thalhs white to gray, continuous, 

slightly roughened. Ascocarps raised, black, straight 

to flexuose, occasionally branched, clearly striate in 

surface view, no thalline margin evident, 1-4 mm 

long, slender. Ascocarp transverse section: hymenium 60-90 pm high, I-; exciple black, usually 

quite closed, lips convergent, striate. Spores 6-8/ 

ascus, (8-) 10-13 locular, 9-10 x (25-) 40-50 (-55) 

pm, I+ blue. 

CHEMISTRY.-NO substances present. 

HABITAT.-LowlandS and cultivated areas. 

DIscussIoN.-Very similar to Graphis rimulosa 

are the following two species. 

Graphis duplicata Acharius (1814): The type 

material at Helsinki is externally identical to the 

lectotype of G. rimulosa. A section from the isotype 

at Uppsala shows an open excipular base, which appears to be the only difference between the two 

species. 

Graphis strialula (Acharius) Sprengel (1827): The 

type material at Helsinki has much shorter ascocarps than G. riinulosa and resembles Opegrapha 

or Melaspilea. In addition, a section from the isotype at Uppsala shows a more or less open excipular base. 

This group of species almost grades into the 

smallest members of the Graphis flexibilis complex, 

from which they differ in consistently lacking any 

thalline margin. 

Hale 35699 (US); Roseau l'allev, Elliott 125 (TUR); Central 

Forest Resene, 1500 ft, lmskaug 33549A (MSC). 

SPECIMESS EXAMIKED.-sOUth Of Portsmouth, sea level, 

23. Graphis subelegans 

FIGURE 6b; PLATE 4e 

Graphis subelegans Kylander, 1891:42 [tjpe collection: San 

Luis Potosi, Mexico, Pringle 162 (H, lectotype)]. 

DEscRIPTIoN.-Thallus usually yellow-brown, offwhite in the Dominican specimen, continuous, 

smooth to rugose, glossy. Ascocarps usually quite 

raised, flexuose, rarely branched, black, obviously 

striate, no thalline margin present, 1-6 mm long, 

0.4-0.7 mm wide. No disc visible in surface view. 

Ascocarp transverse section: hymenium 100-120 pm 

high, I-; exciple black laterally, yellow to yellowred below, open, lips convergent, quite striate. 

Spores 8/ascus, 6-8 locular, 6-8 (-10) X 25-30 pm, 

I + blue. 

CHEXIISTRY.-NO substances present. 

HABITAT.-sea level scrub (probably low elevation exposed scrub throughout its range). 

D~scuss~o~.-Graphis su belegans has been placed 

in synonymy (Zahlbruckner, 1923) of G. endoxantha Xylander (1868); however, the type of the 

latter (Puncher s.n., H) was sterile when described 

(and is still so) and we recommend that this name 

be permanently dropped. 
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

-- 

FIGURE 6.-Cross sections of apothecia of Graphis: a, G. rimulosa (lectotype in P); b, G. subeZegans (Pringle 162); c, G. subnitidula (Wright 155); d, G. tachygrapha (Lindig 869); e, G. 

triticea (Lindig 841); f, G. turgidda (lectotype in BM). 

There is a large group of species related to 

G. su belegans, all characterized by striate black 

labia embedded in basal red-yellow tissue. Most 

similar is probably G. proserpens Vainio (1909), 

which is smaller overall, with more sunken ascocarps. Much additional collecting is necessary to 

unravel this alliance. 

(US). 

SPECIMEN ExAMINED.-Rodney’s Rock, sea level, Hale 35546 

24. Graphis subnitidula 

FIGURE 6c; PLATE 4f 

Graphis subnitidula Nylander in Tuckerman, 1888:123 [type 

collection: Cuba, Wright 155 (US, lectotype; ?FH, isotype, 

not seen)]. 

Graphina subnitidula (Nylander in Tuckerman) Zahlbruckner, 1923:427. 

DESCRIPTION.-Tha~~US brownish to greenish, 
NUMBER 40 23 

smooth, glossy, continuous, distinctly prosoplectenchymatous in section. Ascocarps fissurine, arising 

as a swelling which then cracks and gapes, straight, 

unbranched, 0.3-0.5 (-1.0) mm long. Disc just visible between the gaping lips. Ascocarp transverse 

section: hymenium 80-100 pm high, I-; exciple 

open, lips pale, convergent but exposing some epihymenium. Spores 8/ascus, 4 locular, 5 x 8-10 pm, 

I- or I+ faint, reddish, slow. 

CHEMISTRY.-NO substances present (in type; 

faint unknown near norstictic in the Dominican 

material). 

HABITAT.-upper rain forest. 

DIscussIoN.-The type description of Graphis 

subnitidula cites two specimens: the Wright collection from Cuba (which is a Graphis, and which 

Nylander would have seen) and a later collection 

by Austin from Florida (which is a Graphina and 

probably was never seen by Nylander). Tuckerman’s description clearly recognizes the mixed nature of the spores; we have chosen to follow what 

appears to be Nylander’s original concept and consider this a Graphis. Zahlbruckner’s transfer to 

Graphina then represents a superfluous name. 

Graphis subnitidula is the smallest species yet 

found in the fissurine alliance; for a more complete 

discussion of its relatives, see Giaphis triticea. 

(US). 

SPECIMEN EXAMINED.-Pont Casse, 2200-2600 ft, Hale, 37676 

25. Graphis tachygrapha 

FICXRE 6d; PLATE 5a 

Graphis tachygrapha Xylander, 1863:371 [type collection: 

Colombia, Lindig 869 (H, lectotype; FH, isotype)]. 

Graphis lactea var. dominicana Vainio, 1915:162 [type collection: Trois Pitons, Dominica, Elliott 535 (TUR, lectotype)]. 

Graphis timida Vainio, 1923: 142 [type collection; Trinidad, 

Thaxter 30 (FH, lectotype)]. 

DESCRIPTION.-Thallus very thin, apparently endophloeodal, probably taking most of its color and 

texture from bark and /or epidermis. Ascocarps fissurine, beginning as cracks and finally gaping, unbranched, 0.5-1.0 mm long. Dark disc visible in 

gaping stages. Ascocarp transverse section: hymenium 60-90 pm high, I-, epihymenium slightly to 

much darker than the rest of the hymenium; exciple rudimentary below, lips convergent when 

young, then spreading broadly, partially or incompletely carbonized at least at the apices, sometimes 

to half the height of the hymenium. Spores 8/ascus, 

always 4 locular, 6-9 X 16-22 pm, I- or I+ slow, 

reddish, pale. 

CHEMISTRY.--NO substances present. 

HABITAT.-Elfin forest in Dominica; the type of 

G. tachygrapha is from 2600 meters in Colombia. 

DrscussIoN.-Very similar to Graphis tachygrapha 

is G. humilis, which is tentatively separable by its 

I+ spores and by the much more heavily carbonized lips. For a more complete discussion of the 

fissurine alliance, see Graphis triticea. 

Trois Pitons, summit, 4000-4672 ft, Imshaug 32862 (MSC). 

SPECIMENS EXAMINED.-Mt. Soufrirre, Elliott 1515 (TUR); 

26. Graphis triticea 

FIGURE 6e; PLATE 5b 

Graphis triticea Nylander, 1863:367 [type collection: Villeta, 

Colombia, Lindig 841 (H, lectotype; FH, isotype)]. 

DEsCRiPTION.-ThallUs tan-yellow to grayish, 

thick, glossy, almost always deeply cracked, cortex 

quite prosoplectenchymatous. Ascocarps concolorous with the thallus, arising as fissures in swollen 

thalline margins, sometimes barely revealing the 

tips of the labia and the disc (barely gaping); total 

raised portion 1-4 mm long, to nearly 1 mm wide. 

Ascocarp transverse section: hymenium 120-150 pm 

high, I-; exciple yellow below, open to almost 

closed, laterally yellow, mixed with bark cells, 

apically convergent to somewhat spreading, frequently with peculiar dense whitish tissue at the 

labial apices, the whole exciple embedded in quite 

swollen thalline margin tissue (“puffs”). Spores 

8/ascus, always 4 locular, appearing quite rotund 

and almost halonate because of what appears to be 

a soft, hygroscopic thick outer layer, 10-15 X 15-20 

pm, I- or I + slow, reddish. 

CHEMISTRY.-stiCtiC acid. 

HABITAT.-h or just below elfin forest: probably pantropical in high altitude rain forests. 

DIscvssIoN.-within the fissurine alliance, 

Graphis triticea represents the extreme of ascocarp 

elevation. The most similar species in the group is 

G. insidiosa, which lacks stictic acid. In addition, 

G. insidiosa usually lacks the peculiar white labial 

apices of most New World specimens of G. triticea. 

Most New Zealand specimens of G. triticea (Hayward, 1978) also lack this white tissue. 

The number of names in the fissurine alliance is 
24 SMITHSONIAN CONTRIBUTIONS TO BOTAXY 

large. To distinguish these generally rather inconspicuous species, we have found the following features most useful: (1) presence or absence of stictic 

acid; (2) thickness and degree of specialized organization of the cortex (prosoplectenchymatous versus 

not so); (3) swelling of thalline tissue lateral to the 

exciple, producing emergences in which the fissure 

is embedded (“puffs”); (4) degree of gaping of mature ascocarps, producing a visible disc in extreme 

cases (this may be a variable character and needs 

additional study); (5) degree of labial carbonization 

(rare in the group); (6) iodine reaction of spores; 

(7) overall size of ascocarps and size of spores (this 

character is useful only occasionally, as most fissurines are quite similar in this respect). 

Using these seven criteria, we tentatively propose 

a scheme to separate many of the common fissurines 

(many type specimens of other names remain to be 

seen). 

Graphis bonplandiae (Fee) Mueller Argoviensis 

(1887): An early fissurine name that should be 

permanently dropped; the type was sterile when 

described. 

Graphis dumastii Fee (1824): No stictic acid; 

thallus thick, prosoplectenchymatous; no “puffing”; 

ascocarps gaping; no carbonization; spores I- (or 

reddish); size average for the alliance (nonstictic 

equivalent of G. dumastioidies). Graphis grammitica Nylander (1866) may be synonym. 

Graphis dumastioides Fink: Stictic acid; thallus 

thick, prosoplectenchymatous; no “puffing”; ascocarps gaping; no carbonization; spores I- (or reddish) average size for the alliance (stictic acid equivalent of G. durnastii). 

Graphis humilis Vainio: No stictic acid; thallus 

thin; no “puffing”; ascocarps non-gaping; exciples 

carbonized; spores I + ; slightly smaller ascocarps 

than average. 

Graphis inquinata (Knight and Mitten) Hooker 

f. (1867): Stictic acid; thallus thick, apparently not 

prosoplectenchymatous; some “puffing”; ascocarps 

nongaping; exciple carbonized; spores I- (or reddish); average size for the alliance (rather like a 

carbonized equivalent of G. triticea). 

Graphis insidioja (Knight and Mitten) Hooker 

f.: No stictic acid; thallus thick, apparently prosoplectenchymatous; usually quite “puffed”; ascocarps usually not gaping; no carbonization; spores 

I- (or reddish); ascocarps slightly larger than average (nonstictic equivalent of G. triticea). 

Graphis subnitidula Nylander in Tuckerman: 

No stictic acid; thallus thin; no “puffing”; ascocarps 

sometimes gaping; no carbonization; spores I- (or 

reddish); very small spores and ascocarps. 

Graphis tachygrapha Nylander: No stictic acid; 

thallus thin; no “puffing”; ascocarps sometimes 

gaping; exciple partially carbonized; spores I-; 

ascocarps slightly smaller than average. 

Graphis triticea Nylander: Stictic acid; thallus 

thick, prosoplectenchymatous; ascocarps strongly 

“puffed,” usually not gaping; no carbonization; 

spores I- (or reddish); ascocarps larger than average 

and spores rather rotund (stictic acid equivalent of 

G. insidiosa). 

Another critical early species usually placed in 

this alliance is Graphis lactea (Fee) Sprengel (1827). 

Preliminary examination of the type (G) indicates 

some doubt as to its inclusion here. 

It must also be noted that a fissurine-like series 

exists in Graphina, wherein the spores frequently 

have only three transverse septa (as in this group) 

but also one or two longitudinal septa. Graphina 

incrustans appears to be a sporomorph of Graphis 

dumastii; in all probability a whole series of sporomorphs between these two alliances awaits discovery. 

SPECIMENS ExAMIhED.-Morne Anglais, 3000-3600 ft, Hale 

35324, 35326B, 39377 (US); Morne Diablotin, 3200-4600 ft, 

Hale 39438, 35442 (US), 4300-4500 ft, Imshaug 32912 (MSC), 

Trois Pitons, Elliott 533 (TUR), 3000-1000 ft, lrnshaug 

32886 (MSC), summit, Wzrth 482 (US): Micotrin, ca 3000 ft, 

Wirth 463B (US). 

27. Graphis turgidula 

FIGURE 6f; PLATE 5c 

Graphis turgidda hiueller ArgoJ iensis, 1895a:457 [type collection: Mauritius, s.n. (BM, lectotype)]. 

DEscRIPrros.-Thallus white to off-white, thick, 

continuous, glossy. Ascocarps prominently raised, 

black above, thalline margin quite pronounced, 

straight to more or less flexuose, rarely branched, 

1-4 mm long. Ascocarp transverse section: hymenium (130-) 150-250 pm high, I-; exciple black, 

massively closed below, lips convergent, intact, 

laterally bordered nearly to the apices by a thick 

thalline margin. Spores 6-8/ascus, 11-18 locular, 

10-11 x (40-) 60-90 (-105) pm, I+ blue. 

CHEhlISTRY.-stiCtiC acid. 
KUhlBER 40 25 

HABITAT.-Dominican specimen from lower montane rain forest, on Hibiscus elatus. 

D~sccss~o~.--Almost certainly synonymous with 

Graphis turgidula are G. marginifera Vainio (1921) 

and G. tonglonensis Vainio (1921), both from the 

Philippines. These two species differ only in having 

more exaggerated thalline margins, a feature that 

varies considerably even within a single thallus. 

Graphis turgidula is extremely similar to G. anguilliformis, and is separable only in having stictic 

acid. One fairly poor specimen from Dominica 

(Hale 35069, Can-Dom logging area, Newfoundland) is morphologically identical to G. turgidula 

but has norstictic acid. Whether this represents a 

different species, or a chemically polymorphic population, remains to be seen. 

SPECIMEN EX.AMI&ED.-Sorth of Pont Cask, ca 1400 ft, 

Wirth 544B, (US). 

Key to the Species of Phaeographis 

1. Exciples with carbonized areas. 

2. Ascocarps fissure-like, inconspicuous; exciple open ................. 29. P. arthonioides 

2. Ascocarps robust: exciple heavily carbonized below 

3. Ascocarps pink or cinnabar-red in surface view. 

.......................... 30. P. exaltata 

1. Exciple without carbonization. 

4. Ascocarps cinnabar; exciples closed .......................................... 31. P. haematites 

4. Ascocarps pink; exciples rudimentary ........................... .......... 33. P. rosea 

3. Ascocarps white, grey or blackish in surface view. 

5. ....................... 32. P. mordenii 

5. Spores under 35 pm long. 

6. Spores 4 locular, under 20 pm long: ascocarps usually clumped and radiately 

34. P. cf. subtigrina 

6. Spores 6-8 locular, over 20 pm long, ascocarps not clumped or radiately branched 

Spores over 100 pm long, usually much longer 

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

............................................................. .... ................................................................... 28. P. albida 

28. Phaeographis albida 

FIGURE 7a,b; PLATE 5d 

Phaeographis nlbida (Vainio) Zahlbruckner, 1923:364. 

Graphis albida I’ainio, 1896:251 [t!pe collection: JIorne 

Anglais, Dominica, Elliott 528 (TUR, lectotype)]. 

DEscRIPTIoN.-Thallus white to off-white, continuous, matte. Ascocarps raised, unbranched to 

quite branched, flexuose, covered to the apex by a 

concolorous thalline margin, 0.5-5.0 mm long. 

Disc sometimes completely covered by the convergent labia, sometimes completely exposed, dark 

brown in surface view. Ascocarps transverse section: 

hymenium (70-) 80-150 pm high, I+ blue or I- 

(see discussion), epihymenium very dark; exciple 

yellow-red below and laterally, not well developed, 

lips convergent to quite spreading, tips brown to 

almost carbonized, dark portions probably derived 

(at least in part) by lateral compression of the epihymenium. Spores (4-) 6-8/ascus, 6-8 locular, (6-) 

8-13 x 20-35 pm, frequently halonate after release, 

I+ blue (in the type some cells appearing very 

irregularly muriform). 

CHEMISTRY.--NO substances present. 

HABITAT.-Elfin forest and high altitude rain 

forest. 

DIsccssIoN.-Phaeographis albida is a distinct 

species, apparently endemic to Dominica. The Hale 

and Imshaug collections have ascocarps with spreading labia and much more disc exposed than the 

type; we have found, however, that this character 

is very variable, even within a single thallus, in species with uncarbonized exciples. A very good parallel example is illustrated for Graphina colliculosa. 

This is the only species known to us with a variable iodine reaction of the hymenium. 

Extremely similar to P. albida is Phaeographina 

oscitans; the latter can be distinguished only by the 

spores, the consistently I- hjmenium, and the 

presence of stictic acid. 

SPECI\IE\S Ex \\II~ED -Fresh I\‘ater Lake, 2600-2800 ft, Hale 

334j3 (US); Morne Diablotin, 3300-4300 ft, Inzshat~g 32936 

pp. prsc). 

29. Phaeographis arthonioides 

FIGURE 7c; PLATE 3c 

Phaeogiaphis artlzonzoides (Vainio) Zahlbruckner, 1923:364. 
26 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 7.-Cross sections of apothecia of Phaeographis: a, P. albida (Elliott 528); b, P. albida 

(Hale 35453); c, P. arthonioides (Raunkiaer 548); d, P. exaltata (Elliott 1301); e, P. mordenii 

(Hale 35071); j, P. haematites (Hale 38013); g, P. cf. subtigrina (Wirth 444); h, P. rosea (Elliott 

523). 
NUMBER 40 27 

Graphis arthonioides Vainio, 1915:155 [type collection: St. 

Phaeographis sexloculata Fink, 1927:215 [type collection: 

Croix, Raiinkiaer 548 (FH, isotype)]. 

Puerto Rico, Fink 1436 (MICH, lectotype)]. 

DEscRIPTIoN.-Thallus gray to whitish, continuous, glossy, rather thin. Ascocarps fissurine, barely 

raised, margins blackish to black, mostly unbranched, straight to somewhat flexuose, usually 

1 mm or less long. Disc usually not visible in surface view. Ascocarp transverse section: hymenium 

65-70 ym high, I-, epihymenium dark; exciple 

rudimentary below, lips more or less convergent, 

irregularly carbonired at least apically, frequently 

to the base of the hymenium. Spores 8/ascus, 6-0 

locular, 7-10 x 19-26 pm, I- or I+ slow, reddish. 

CHEmSTRY.-Isotype of P. arthonioides has a 

faint unknown; lectotype of P. sexloculata is P-, 

too small for adequate TLC; Dominican specimen 

contains no substances. 

HABITAT.-LoWlandS in exposed or cultivated 

areas. 

DIscvssIoN.-Extremely similar to Phaeographis 

arthonioides is P. decipiens (Fee) Mueller Argoviensis (1887), also from the Caribbean; the latter differs 

only in having stictic acid. Differing only in spores 

is Graphis hunzilis Vainio. 

shaug 33128 (MSC). 

SPECIML\ EXAMI!ED.-EaSt of Pointe Michel, 200 ft, Im30. Phaeographis exaltata 

FIGURE 7d; PLATES 5f, 6a 

Phaeographis exaltata (Montagne and van den Bosch) MuelLecanactis exaltata Montagne and van den Bosch, 1855:475 

ler Argoviensis, 1882:336. 

[type collection: Java, Junghuhn s.n. (L, lectotype)]. 

DEscRIPTIoru.-Thallus nearly white to gray, 

thick, usually glossy, continuous. Ascocarps very 

prominent, from Sarcographa-like and circular to 

elongate and flexuose (on one thallus), thalline margin very prominent, from l x l mm to l x 6 mm. 

Disc wide, black to gray-pruinose. Ascocarp transverse section: hymenium 120-200 pm high, I-, epihymenium blackened; exciple black, heavily closed 

below, lips usually quite spreading, intact, frequently incompletely carbonized. Spores 8/ascus, 

6-8 locular. 7-11 x 20-30 pm, I- or I+ faint, 

slow, reddish. 

cHEMISTRY.-k “Discussion.” 

HABITAT.-upper montane and elfin forest. One 

of the three major graphids on elfin forest Clusia 

uenenosa (with Graphis olivacea and Phaeographis 

m or d e n i i) . DiscussIox.-Phaeographis exaltata is a widespread, common pantropical species of high altitude 

rain forests. The large size, distinctive appearance, 

and heavily carbonized exciple render it unlikely 

to be confused with any other species of Phaeographis. Individual specimens (and frequently parts 

parts of a single specimen) can mimic Sarcographa 

closely; the only consistent difference is the lack of 

cracking and partitioning of the disc so common 

in Sarcographa. 

Partially because of the distinctive morphology, 

we find it difficult to split P. exaltata into two chemical species. The vast majority of the Paleotropic 

specimens (including the type) have no lichen acids; 

the vast majority of Neotropic specimens have at 

least one of the three “quintaria” unknowns. Even 

within the relatively small area of Dominica both 

chemical forms can be found. 

As P. exaltata is both common and fairly well 

represented in herbaria, it would seem to be an 

excellent subject for a future study of chemical 

variation in a crustose species. 

SPECIMENS ExAMlNED.-can-Dom logging area, Brantridge 

Estate, 1700 ft, Hale 35301, 35321 (US); Can-Dom logging 

area, Ponte Casse, 2000 ft, Hale 35009, 35091, 35100, 35115, 

35116 (US); Boiling Lake, 2400-3000 ft, Hale 35622A (US); 

hforne Anglais, 3000-3600 ft, Hale 35322 (US); Trois Pitons, 

30004000 ft, Imshaug 32883, 32888 (MSC), summit, Wirth 

480, 483 (US); Morne Diablotin, 35004300 ft, Imshaug 32934 

(MSC), 3200-4600 ft, Hale 35323, 35445 (US); Prince Rupert, 

Elliott 1307 (TUR). 

3 1. Phaeographis haematites 

FIGURE 7f; PLATE 6b 

Phaeographis haematites (Fee) Mueller Argoviensis, 1882:384. 

Graphis haematites Fee, 1824:45 [type collection: South 

America, s.n. (G, lectotype)]. 

DEscRIPTIoN.-Thallus tan to pale brown, continuous, smooth to matte. Ascocarps raised, prominent, occasionally branched, usually quite sinuous, 

thalline margin very prominent, usually covering 

the ascocarp to the apex (but frequently separated 

from the disc and labia by a distinct crack), up to 

15 mm long, slender. Disc (and labial tips) deep to 

bright cinnabar-orange, disc quite obvious in sur- 
28 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

face view. Ascocarp transverse section: hymenium 

80-100 pm high, I-; exciple red, uncarbonized, 

more or less closed, the pigment soluble in Hoyer's 

medium, labia more or less spreading, somewhat 

thickened apically. Spores 8/ascus, 6-10 locular, 9- 

11 x 21-35 pm, I-. 

CHEMISTRY.-NO substances present except for 

the red pigment. 

HABITAT.-Rain forest in Dominica; probably 

from lowland scrub to high rain forest elsewhere in 

its range. 

DIscvssIo~.-Phaeographis cinnabarina (Fke) 

hfueller Argoviensis (1887) is the only species likely 

to be confused with P. haematites. The former can 

easily be distinguished by its thinner, duller ascocarps, with a brownish rudimentary exciple, and 

the much smaller hymenium with much darkened 

epihymenium. Somewhat similar is Phaeographina 

chrysocarpa (Raddi) Redinger (1933), which differs 

in the spores and in the heavily carbonized exciple. 

SPECISIES ExAMINED.-Dleau Gommier Forest Reserve, 1200 

ft, Hale 38013 (US). 

32. Phaeographis mordenii, new species 

FIGURE 7e; PLATE 6c 

DEscRIP.rIoN.-Thallus continuus, laevis vel ruguloso-inaequalis. Apothecia sessilia, rotunda vel 

lirellina, simplicia, recta, 2-8 longa, ca. 1 mm lata, 

disco dilatato, primum pallido, dein nigricante, 

pruinoso. Excipulum subintegrum, flavum vel rufoferrugineum, labiis integris, divergentibus; hymenium inspersum, 200-250 pm altum, I-. Asci 6-8 

spori, sporae fuscescentes, 30-40 (-50) loculares, 

12-17 x 120-220 pm, in mentem revocans sporae 

Conotremae, I + rufo-fuscae. 

Thallus light gray and glossy near the apothecia, 

white and matte where more exposed, continuous, 

distinctly rugulose in spots. Ascocarps prominently 

raised, nearly round to irregular to elongate, rarely 

branched, with a prominent tlialline margin, 2-8 

mm long, ca. 1 mm wide. Disc very broad, initially 

concolorous with the thallus, then dark blackishgray and pruinose. Xscocarp trans\.erse section: hjmenium 200-250 pm high, heavily inipersetl and 

beaded, epihymenium darkened, I-; exciple closed 

to nearly open, reddish yellow, labia intact, divergent, covered by the thalline margin. Spores 6-8/ 

ascus, brown, very long and slender, 30-40 (-50) 

locular, 12-17 x 120-220 pm, occasionally with 

irregular biocellations in the end chambers, chamber walls frequently thickened, reminiscent of 

Conotrema spores, I+ slow, eventually dark redbrown. 

CHEMISTRY.-~O substances present. 

HOLOTYPE.-~~OSS~ and elfin forest, trail to summit of Morne Diablotin, Dominica, 3200-4600 feet, 

Hale 35071 (US). 

DIscvssIox.-Phaeogi.aphis mol-denii is one of the 

three characteristic and abundant graphids on 

Clusia venenosa in the elfin forest. It is quite distinct in its large size and enormous, thick-walled 

spores; there would seem to be no closely related 

species. 

Occasionally, some spores will show biocellate 

ends, similar to those of Graphina vestoides; however, these extra locules are irregular and seem to 

represent disintegration of cell contents rather than 

a submuriform condition. 

SPECIMENS EXAMlNED.->forne Diablotin, 35004300 ft, Zmshazrg 32922, 32945A (MSC); Morne Trois Pitons, 30004000 

ft, Zmshaug 32884 (MSC), summit, on Clusia uenenosa, Wirth 

479, 481, 484, 485, 486 (US). 

33. Phaeographis rosea 

FIGURE ih; PLATE Gd 

Phaeographis rosea (I'ainio) Zahlbruckner, 1923:385. 

Graphis roses \.ainio, 1896:259 [type collection: Morne 

Anglais, Dominica, Elliott 523 (TUR, lectotype)]. 

DEScRIPTIo~.-Thallus tan, continuous, smooth 

and shining. Ascocarps barely emergent, rather fissurine, occasionally branched, flexuose, the margins 

distinctly stained rose-pink, 5-15 mm long, slender. 

Disc not visible in surface view. Ascocarp transverse 

section: hymenium 140-160 pm high, I-; exciple 

rudimentary, labia convergent, brownish above. 

Spores 8.'ascus, always 4 locular, 9-12 X (13-) 

18-25 pm, I+ blue. 

CHEI\ZISTRY.-~O substances present. 

HABITAT.-upper montane forest. 

DIscvssIos.-Phaeographis rosea is known only 

from the type collection. The rose-pink pigmenta, 

tion that renders this species so distinct is apparently a normal condition; occasionally, some 

graphids will show discolored, reddish areas on 

thalli of normally unpigmented species (see Gra- 
NUMBER 40 

phina diinidiata for a Dominican example). The 

color in P. yosea, however, is restricted to the ascocarp margins, and appears to be normal. 

34. Phaeographis cf. subtigrina 

FIGURE 7g; PLATE 6e 

Phaeographk subtigrina (T'ainio) Zahlbruckner, 1923:387. 

Graphis subtigriria Vainio, 19OT:17T [type collection: Koh 

Chang, Gulf of Siam, Schmidt XXI (TUR, lectotype)]. 

DEscRIPTIoN.-Thallus thin, off-white, continuous, matte, Ascocarps usually clumped, commonly 

radiately branched, forming clusters to 2 mm in 

diameter, flush, thalline margin frequently separating from the thallus. Disc broad, gray, lightly but 

distinctly pruinose. Ascocarp transverse section: 

hymenium 115 pm high (Dominican specimens; no 

measurement for the type), I-; exciple pale below, 

closed to nearly open, lips darker but not carbonized, intact, divergent. Spores S/ascus, thick-walled, 

4 locular, 4-6 X 10-17 pm, I-. 

CHEMISTRY.-NOrStiCtiC acid only (type); norstictic and stictic acid (Dominican specimens). 

HABITAT.-one Dominican specimen from sealevel, the other from upland rain forest. Apparently 

of great ecological amplitude. 

DIscussIozr.-Phaeog?-aphis subtigrina has been 

reported (Vainio, 1915) from Guadelope, but we 

have not seen these specimens. The Dominican 

material differs from the type primarily in the 

chemistry. 

As in many species of Phaeographis, the ascocarp 

form in P. subtigrina varies greatly within the same 

thallus. In the Dominican specimens these structures vary from very narrow, heavily branched, and 

clumped to quite broad, barely branched, and 

hardly clumped. 

Quite similar to P. subtigrina is P. dimorpha 

(Nylander) Zahlbruckner (1923), which differs in 

having a more prominent margin and more raised 

ascocarps. 

SPECNE~S Ex4~1h~~.-Roseau Botanic Garden, lVirth 444 

(US); Emerald Pool, Wiith 505 (US). 

Key to the Species of Graphina 

1. Exciples distinctly carbonized. 

2. Disc broad and obvious in surface view, labia quite divergent. 

3. 

3. Spores without colloidal layer; ascocarps more or less raised. 

Spores with broad persistent colloidal layer; ascocarps flush with thalline surface .......... 

.................................................................................................................................. 41. G. collospora 

4. Spores I/ascus; protocetraric acid present ........... 52. G. platyleuca 

4. Spores more than I/ascus, some darkening; n sent .................................. 

.............................................. G3. Phaeographina cf. difformis 

.... 

2. Disc narrow, not prominent in surface view, labia more or less convergent. 

5. Ascocarps with powdery tvhite covering, easily rubbed to 

6. Spores under 20 &rn long; salazinic acid present ........... 

6. Spores over 40 pm long; stictic acid present .................. 

5. Ascocarps without a poivdery white coiering. 

7. Labia entire or nearly so. 

7. G triphoroides 

abial apices (upper ?,'* or less of total exciple 

........................................................ 43. G. diniidiata 

o base of hymenium or beyond. 

9. Ascocarps Opregraphn-like, 1-2 mm long, without a thalline margin ................ 

...................................................................................................................... ~1. G. nuda 

9. Ascocarps elongate, flexuose, with a thalline margin. 

10. Spores less than 61 pm long. 

11. Norstictic acid present; mature spores 8/acus ................................ 

................................................................................ . pseudoanaloga 

11. KO substances present; mature spores usually 4-6/ascus ........................ 

................................................................................................ 53. G. plurispora 
30 SMITHSOKIAS COSTRIBUTIONS TO BOTASY 

10. Spores over 85 pm long. 

12. Only the ends of the spores muriform .................... 58. G. uestitoides 

12. Entire spore densely muriform. 

13. Ascocarps under 0.5 mm wide; hymenium under 250 pm high. 

35, G. acharii 

13. Ascocarps massive, usually 1 mm wide; hymenium over 350 pm 

high .... ..46. G. illinata 

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

7. Labia striate. 

14. 

14. Exciple without yellowish areas, closed or nearly so. 

15. Spores under 61 long. 

Exciple yellow to yellow-brown laterally, more or less open .... 37. G. antillarum 

Norstictic acid present; mature spores 8/ascus ...... 54. G. pseudoanaloga 

.............................................................................. ~3. G. plurispora 

16. 

16. No substances present; mature spores usually 4-6/ascus ............................ 

15. Spores over 64 @rn long. 

17. Only ends of spores muriform ........................................ 58. G. vestitoides 

17. Entire spore densely muriform. 

18. Exciple base irregularly carbonized; spores always l/ascus ................ 

............ .............................. 49. G. macella 

18. Exciple base usually evenly carbonized; spores usually 4Iascus ........ 

.................................................................................................. 35. G. acharii 

19. Apothecia fissurine, i.e., originating as a crack, the disc usually concealed by or sunken 

between the narrow.to gaping fissure walls. 

20. Spores l/ascus, over 50 pn long ........................ 48. G. insculpta 

20. Spores 6-8/ascus, under 50 pm long. 

21. Thallus prosoplectenchymatous .................... 47. G. incrustans 

21. Thallus not prosoplectenchymatous. 

22. Psoromic acid present .................................. ..42. G. columbina 

22. No substances present ................................... , G. cf. adscribens 

1. Exciples totally uncarbonized. 

19. Ascocarps not fissurine. 

23. Disc broad, easily seen in surface view. 

24. 

24. Disc whitish, gray to greenish; spores over 17 p long; salazinic acid absent. 

Disc reddish: spores under 17 pm long; salazinic acid present .... 40. G. colli~~lo~a 

25. Spores 17-28 long; norstictic acid present .................... 56. G. suberythrella 

25. Spores 50-150 pm long; “quintaria” unknowns present ............ 59. G. virginea 

23. Disc narrow, not prominent in surface view. 

26. Margins of asocarps distinctly reddish to pinkish brown in surface view, a 

least in upper portions. 

27. Spores l/ascus, over 80 pm long ............... 39. G. chlorocarpa 

27. Spores Z-SIascus, under 75 pm long. 

28. Korstictic acid present; exciples with internal striae only ............................ 

28. So substances present; internal striae absent. 

29. 

29. Mature spores 8/ascus, less long; labia entire .............. 

.......................................................................... 38. G. carneouir 

.............................................................................. 44. G. dispersa 

Mature spores 2-4 ascus, over 35 ,pm long; labia slightly striate ........ 

.................................................. ................... 55. G. rufopallida 

26. Margins of ascocarps tan, white, or greenish. 

30. Spores more than 40 pm long; salazinic acid absent. 

31. Ascocarps always massive and protuberant; hymenium over 200 #m 

31. Ascocarps rarely protuberant, never massive; hymenium less than 200 

&m high, at least the epihymenium I+ blue; “quintaria” unknowns 

present 59. G. uirginea 

30. Spores less than 17 pm long; salazinic acid present ............. 40. G. coll~culosa 

high; I-; no substances present. .................................... 45. G. frumentaria 

................................................ 
NUMBER 40 31 

35. Graphina acharii 

FIGURE 8a; PLATE 6f 

Graphina achaii (Fee) Mueller Argoviensis, 1887:38. 

Graphis acharii Fee, 1824:39 [type collection: South America, 

Graphis inturgescens Krempelhuber, 1876:383 [type collecGraphitia inturgescens (Krempelhuber) Mueller Argoviensis, 

s.n. (G. lectotype)]. 

tion: Brazil, Glaiiou 6286 (BM, lectotype; M, isotype)]. 

1888a:545. 

D~sc~r~~rorv.-Tha~~us gray to off-white, thick, 

continuous to cracked. Ascocarps large, strongly 

raised, black above, straight to flexuose, occasionally branched, covered nearly to the tops by a 

prominent thalline margin, 1-6 mm long. Disc not 

visible in surface view. Ascocarp transverse section: 

liymenium 150-200 pm high, proportionately small, 

I-; exciple black, massively closed, labia convergent, 

nearly entire to crenate to distinctly striate (often 

within a single ascocarp). Spores (in the type of G. 

acharii) usually 41ascus but occasionally one or 

three (and rarely six), densely muriform, 18-25 x 

95-170 pm, I+ blue. 

CHEMISTRY.-NO substances present. 

HABITAT.-secondary forest (Dominica). 

DIscvssIoN.-According to Mueller Argoviensis 

(1887), the spores in the type of G. acharii have 

biocellate ends; however, we have been able to find 

only densely muriform spores in this same material. 

In an earlier treatment of this species (Wirth and 

Hale, 1963), we had not seen the type, and followed 

Mueller in including specimens with biocellate 

spores in G. acharii. Although additional collections may prove these spore variants to be part of 

the same continuum, we are presently restricting 

the name G. acharii to those forms with densely 

muriform spores only. Those specimens with biocellate spores can be referred to G. vestitoides 

The ascocarps in G. acharii vary from nearly 

oryzaeform to quite long, frequently within the 

same specimen. Also very variable is the number of 

spores per ascus, and the degree of striation of the 

labia. The subentire labia characteristic of G. inturgescens can frequently be found on the same 

thallus with the heavily striate labia of typical 

G. acharii. 

This species is extremely common throughout the 

Neotropics; a sporomorph is Graphis flexibilis. 

Somewhat similar among the Dominican Gra- 

(P. 44). 

phina species is G. macella, which differs in being 

smaller overall and in having an irregularly carbonized excipular base. 

(US). 

SPECIMEN ExAM1xED.-Syndicate Estate, 1800 ft, Hale 35555 

36. Graphina cf. adscribens 

FIGURE 8b; PI ATE 7a 

Graphina adscribens (Sylander) Mueller Argoviensis, 1892a: 

Graphis adscribens Nylander, 1868: 177 [type collection: Lifu, 

284. 

New Caledonia, Thiebaut s.n. (H, lectotype.]. 

DEsCRIPTIoN.-Thallus white, thin, continuous to 

cracked. Ascocarps very inconspicuous in the Dominican material and in the type photographs, fissurine, slightly mealy in the Dominican specimen, 

concolorous with the thallus (at least in the Dominican specimen), less than 1 mm long. Disc not visible in surface view. Ascocarp transverse section: 

hymenium 100-120 pm high, (I- Dominican specimen); exciple rudimentary in the Dominican material, apparently slightly reddish in the type, lips 

convergent, intact. Spores 8/ascus, muriform 8-9 x 

21-25 pn (type), 9-10 X 2340 (Dominica), I- in 

type (fide Nylander), I+ reddish, slow in the Dominican material. 

CHEJIISTRY.-NO substances present (both specimens). 

HABITAT.-on cocos at beach (Dominica only). 

DIscussIoN.-This tentative identification is 

based on a black and white photograph of the 

lectotype (which is a very small specimen) and a 

sketch of the excipular cross-section of the type. According to both Nylander and Mueller, the type is 

reminiscent of Gyaphina chlorocarpa and Graphis 

grammitis, both of which have reddish ascocarps in 

surface view. As the Dominican specimen is uniformly pale, the identification must remain tentative until the type can be examined. 

SPECIMEN EXAXINED.-.~~S du Me, Wirth 519 (US). 

37. Graplaina antillarum 

FIGCRE 8c: PLATE 7b 

Graphina antillarum (\'ainio) Zahlbruckner, 1923:398. 

Graphis an tillarum Yainio, 18993255 [tkpe collections, Guadeloupe, Duss 540 (FH, isotype)]. 
32 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

C 

FIGURE 8.-Cross sections of apothecia of Graphinn: a, G. acharii (lectotvpe in G); b, G. cf. 

adscribens (Wirth 519); c, G. antillarum (Duss s.n.); d, G. cnrneouiridis (Haie 3i642); e, G. 

chlorocarpa (lectotype in G); f, G. col[iculosa (lectotype of G. eugeniae, Merrill 3471): g, G. 

colliculosa (lectotype in PC). 

Graphis acuminata Vainio, 1915:147 [type collection: St. Jan, Graphina sulcata Fink, 1927:217 [type collection: Puerto Rico, 

Raunkiaer 437 (TUR, lectotype)]. Fink 659 (MICH, lectotype)]. 

DESCRIPTION.-T~~~~~US gra) to white, continuous, 

matte. Ascocarps nearly flush to partially emergent, 

black, commonly branched and flexuose, thalline 

Graphis platycarpoides Yainio, 1915: 145 [type collection, 

Graphina acuminata (Yainio) Zahlbruckner, 1923:393. 

Graphina platycarpoides (Yainio) Zahlbruckner, 1923:419. 

Guadeloupe, Duss 1198 (TUR, lectotype)]. 
NUMBER 40 33 

margin absent or nearly so, 5-6 mm long, slender. 

Ascocarp transverse section: hymenium 80-100 

(-120) pm high, I-, exciple black laterally, yellow 

to yellow-brown below, more or less open, labia convergent, barely to quite strongly striate. Spores (I-) 

2-4 (-6)/ascus, distinctly muriform, 15-22 X 28-46 

(-60) pm, I+ blue. 

CHEMISTRY.-NOlStiCtiC acid. 

HABITAT.-LoWland scrub forest and higher altitude cultivated areas and road cuts. 

DrscussIoN.-Among the Graphina species with 

carbonized, striate, dimidiate exciples, G. antillarum is fairly distinct by virtue of its 24 small, 

rotund spores per ascus, and the presence of norstictic acid. There are, however, several other species that are extremely similar and may prove to 

be conspecific: 

Gmphina bzpartita XIueller Argoviensis (1 888c) 

differs in having a thin thalline layer over the ascocarps and in having only the very tips of the labia 

carbonized. 

Graphina deserpens (Vainio) Zahlbruckner (1923) 

differs in having 8 spores per ascus and stictic acid. 

It is probably a synonym of G. parilis, below. 

Graphina elongata (Vainio) Zahlbruckner (1923) 

differs in having both stictic acid and norstictic 

acid, more dendritically branched apothecia, and no 

basal yellow area in the exciple. 

Graphina parilis (Krempelhuber) Mueller Argoviensis (1892b) differs in having stictic acid and 8 

spores per ascus. 

Most of the Dominican specimens fit comfortably within the boundaries defined here for G. 

antillarum. One, however (Wirth 544c), has the 

brown exciple base of G. elongata but lacks the 

stictic acid of G. elongata. 

SPECIMENS ExAMINED.-Prince Rupert Bay, Irnshaug 33529A 

(MSC); Brookhill Estate, 100-150 ft, Imshaug 33307.4 (MSC); 

Roseau Botanic Garden, 100 ft Wirth 445 (US); north of 

Bioche, 200 ft, Hale 35745; South Chiltern Estate, 1300 ft, 

Zmshaug 33050, 33036 (MSC); near Pont Cassk, 1400 ft, Wirth 

544c (US). 

38. Graphina carneoviridis, new species 

Frcum 8d; PLATE 7c 

DESCRIPTION.-Tha11uS continuus, laevis, stratum 

corticale prosoplectenchymatum. Apothecia sessilia, 

recta vel flexuosa, 1-2 (-3) mm longa. Excipulum 

dimidiatum, rufo-fuscum, labiis convergentibus, integris vel indistincte incisis; hymenium 75 pm 

altum, I-. Asci 8 spori; sporae decolores, murales, 

loculis horizontalibus 3, loculus transversis 2, 10-12 

X 15-17 pm, I+ coeruleae. 

Thallus greenish, continuous to somewhat 

cracked, glossy, prosoplectenchymatous. Ascocarps 

raised, occasionally branched, straight to Bexuose, 

pink-brown above, laterally bounded by a green 

thalline margin, 1-2 (-3) mm long, slender. Disc 

not visible in surface view. Ascocarp transverse section: hymenium 75 pm high, I-; exciple reddishbrown, open below, labia convergent, intact to 

barely striate. Spores 8/ascus, 3 X 2 locular, 10-12 

X 15-17 pm, I+ blue. 

CHEMISTRY.-NO substances present. 

HoLoTYPE.-hIixed disturbed and primary rain 

forest, trail from Brigantin through XIiddleham 

Estate toward Morne Trois Pitons, Dominica, elevation 2200-2600 feet, Hale 37642 (US). 

DISCLJSSION.-G~U~~~~U carneouiridis is externally 

rather similar to Graphina dispersa, from which it 

differs in having smaller spores, a more open exciple with nearly intact labia, a prosoplectenchymatous cortex, and in a different chemistrv. 

39. Graphina chlorocarba 

FIGURE 8e; PLATE 7d 

Graphina chlorocarpa (Fee) Mueller Argoviensis, 1887:44. 

Graphis chlorocarpa Fee, 1824:47 [tjpe collection: Pe? 14. s.n. 

(G, lectotype)]. 

Graphina balbisii var. monospora Redinger, 1933:61 [type 

collection, Maltne 494 (S, lectotype; FH, isotype)]. 

DEsCRrPTroN.-Thallus tan to grey-green, continuous, glossy, smooth to slightly rugose. Ascocarps 

usually raised (nearly flush when young), straight 

to quite flexuose, usually unbranched, concolorous 

with the thallus except (usually) for the disc, which 

is pale orange-brown, 1-5 mm long, about 0.5 mm 

wide. Ascocarp transverse section: hymenium 120- 

180 pm high, I-; exciple yellow-brown to redbrown, open below, labia convergent, lightly striate, 

the tips of the striae sometimes slightly darkened. 

Spores l/ascus, densely muriform, 24-40 X 80-130 

pm, I+ blue. 

CHEMISTRY.-Both types and most specimens: no 

substances present. A few Dominican specimens 

with unknown substances. 

HABITAT.-A~I fertile specimens from rain forest; 

sterile material (see discussion) from sea level and 
34 SMITHSONIAN CONTRIBUTIOKS TO BOTANY 

mid altitude cultivated areas. 

DIscussIorv.-The only neotropical G~aphina species likely to be confused with G. chlorocarpa is 

G. DrclDi.cii (Fee) Mueller .irgovieiisis. Both these 

species share the distinctive orange-brown disc area 

and the striate reddish exciple; the only difference 

lies in the number of spores per ascus. In G. chlorocarpa the asci are always monosporous; in G. balbisii the asci usually contain 3-4 slightly smaller 

spores, occasionally 6 or 2, but never only one. As 

yet, 1iaI.e seen no ,pecinien~ that bridge the 

rather small gap between these two species. 

l\Tithout spores, it is not possible to be sure which 

of these two species is in hand. All the fertile Dominican material referable to G. chlorocarpa is 

from rain forest; however, four additional specimens from cultivated areas (Zmshaug 33024, 33417, 

33424, Wivth 446) are sterile and may be either of 

the two species. No fertile material of G. balbisii 

has yet been identified from the Island. 

(US); Can-Dom logging area, Pont CassC., 2000 ft, Hale 39097, 

35129 (US); Ridgefield Estate, 1100-1200 ft, In7shaug 33417, 

33424 (JISC, both sterile): South Chiltern Estate, 1300 ft, 

I~nsiiaug 33024 (lISC, sterile); Roseau Botanic Garden, 100 

ft, Wirth 446 (US, sterile). 

SPECI.UESS ExAMINED.--Sear Laudat, 800 ft, Hale 35797 

40. Graphina colliculosa 

FIGURES 8f,g; PLATE 7e 

Graphina colliculosa (hfontagne) Hale, 1976: 156. 

Sclerophyton coliicuZosu~n Montagne, 1851 :61 [type collection: 

Graphis intortnila Stirton, 1881 :I86 [type collection: Assam, 

Graphis eugeniae Yainio, 1921:262 [type collection: PhilipGraphis colliculosa (Montagne) Zahlbruckner, 1923:299. 

Graphina intortula (Stirton) Zahlbruckner, 1923:411. 

Graphina aibonite7isis Fink, 1927:219 [type collection: Puerto 

Guyana, Leprieur 1406 (P, lectotype: Bhf, isotype)]. 

s.n. (BM, lectotype)]. 

pines, Merrill 3971 (TUR, lectotype: FH, isotype)]. 

Rico, Fink 2017 (MICH, lectotype)]. 

~ESCRIPTI~S.-T~~~IILI~ gray to pale tan to faintly 

greenish, smooth, glossy, continuous, thick, the 

upper layer compacted and prosoplectenchymatous 

in section. Ascocarps raised, sinuous, commonly 

branched, to 30 mm long, slender. Disc sometimes 

prominently displayed as a red-brown line in surface view, sometimes completely covered by the concolorous thalline margin. Ascocarp transverse section: hymenium 80-100 pm high, I-, epihymenium 

darkened; exciple rather ill-defined, yellow to reddish-yellow, usually closed below, labia quite convergent to quite divergent, more or less intact or 

having the appearance of internal striae. Spores 

8/ascus, 4 x 1-2 locular, 5-7 x 11-16 pm, I+ blue. 

CIIEMrsTRY.-Salazinic acid. 

HABITA.r.-Rain forest canopy in Dominica; elsewhere found in both rain forest and secondary 

forest. 

DISCLXIOS.--GYU~~~~U colliculosn is an extremely 

vigorous, pantropical species; we have seen specimens from much of the Caribbean (Cuba, Puerto 

Rico, St. Vincent, Grenada, Trinidad, St. Lucia), 

Panama, Guayana, Tahiti, Fiji, hssam, and the 

Philippines. 

As in many species with totally uncarbonized 

exciples, the degree of labial divergence is quite 

variable (see Phaeographis nlbida for a parallel condition). The taxa listed as synonyms differ from 

each other primarily in the extent to which the redbrown disc is exposed; as complete exposure and 

complete concealment can be found within a single 

thallus, this character seems clearly useless to distinguish species. 

A second feature that is quite variable in many 

species with uncarbonized labia is the presence of 

“internal” striae, i.e., dark lines within the lips. 

These darkened masses of cells seem to arise with 

age, by lateral compaction of old hymenium. The 

type of G. aibonitensis (Figure 9a) is a good example of the extreme of this condition. Note also the 

very thick basal closure of the exciple, which seems 

also to be correlated with age. 

Chemically, G. colliculosn is unlikely to be confused with any other species, as salazinic acid is 

quite rare in the family. However, several other 

species are quite 5iinilar in inorpholog), as follows. 

Graphina eiythrella (Montagne) Zahlbruckner 

(1923) is distinguished by having norstictic acid, 

larger spores, and somewhat shorter, more raised 

ascocarps. Some of the sterile, norstictic acid-containing syntypes of G. eugenine (MacGregor 41315, 

Elmer 15077) may be referable to this species. 

G9,aphina riopiedwnsis Fink (1927) differs only 

in the somewhat larger spores and in having stictic 

acid. 

SPECIVENS ExAlrINED.<astle Bruce Road, 1000 ft, Hale 

38177 (US): Can-Dom logging area, Dleau Gommier, 1600- 

1700 ft, Hale 35175 (US); Soufriere Ridge, 2200-3100 ft, 

Imshaug 33068 (MSC). 
NUMBER 40 35 

FIGURE 9.4ross sections of apothecia of Graphina: a, G. colliculosa (lectotype of G. aibonitensis 

Fink); b, G. collospora (Boergesen s.n.); c, G. columbina (Beaumont s.n.); d, G. dimidiata 

(Vainio 332); e, G. dispersa (Malme 2038B); f, G. frumentaria (lectotype in G); g, G. illinata 

(lectotype in M); h, G. incrustans (lectotype in G). 
36 SSIITHSOSIAN COSTRIBUTIOSS TO BOTAKY 

41. Graphina collospora 

FIGURE 9b; PLATE 7f 

Graphina collospora (l‘ainio) Zahlbruckner, 1923:402. 

Graphis collospora T.ainio, 19153153 [type collection: hft. 

Eagles, St. Croix, Boergeseri s.n. (TUR, 1ectot)pe)l. 

DEsCRIPTioS.-l’hallus greenish gray, thick, continuous, matte. Ascocarps flush with the thallus, 

lips completely covered by the thalline margin 

which frequently cracks away from the rest of the 

thallus, ascocarps occasionally branched, flexuose, 

0.5-2.5 mm long. Disc very prominent in surface 

view, brown and lightly pruinose. Ascocarp transverse section: hymenium 90-100 pm high, epihymenium brown, upper portion of hymenium 

and all of epihymenium I+ blue. Exciple brownblack, barely closed to barely open, labia quite 

divergent. Spores 1 jascus, densely muriform, 25-37 

x (SO-) 105-130 pm, with broad pale gelatinous 

outer layer, ca. 15 pm thick, which frequently persists even after ejection from the ascus, I+ blue. 

CHEMISTRY.-Norstictic acid, stictic acid, constictic acid. 

HABITAT.-on cocos at the shore. 

DiscussroN.-Graphina collospora is part of the 

complex centering around G. confluens (Fee) Mueller Argoviensis, which is very similar in morphology 

and anatomy but lacks the odd colloidal spore 

layer. It also differs in chemistry (lichexanthone, 

stictic acid, and constictic acid). 

The Dominican specimen seems to represent 

only the second collection of G. collospora. 

(US). 

SPECIMEYS EXAMINED.-.4ns du Me, sea level, Wirth 520 

42. Graphina columbina, new combination 

FIGURE 9c; PLATE 8a 

Graphis coltimbina Tuckerman, 1888:123 [tvpe collection: 

Alabama, USA, Beaumont s.n. (FH, lectotype)]. 

Fissurina virginalis Tuckerman ex Kylander, 1889:50 [type 

collection: Florida, USA, s.n. (US, lectotype)]. 

Graphis virginalis Tuckerman ex Eckfeldt, 1890:256. 

Graphim virginalis (Tuckerman ex Nylander) Mueller Argoviensis, 1895b:4T. 

Phaeographina columbina (Tuckerman) Zahlbruckner, 1923: 

436. 

DEscRIPTrox.-Thallus greenish to brownish, 

continuous, smooth to matte. Ascocarps fissurine, 

the margins paler than the surrounding thallus, 

straight to flexuose, occasionally branched, 0.5-5 

mm long, slender. Disc not visible in surface view. 

Ascocarp transverse section: hymenium 90-125 pm 

high, I-; exciple yellowish, rudimentary, labia convergent to somewhat gaping, entire, containing 

many bark cells. Spores 6-8/ascus, irregularly muriform, 1-3 x 3-6 locular, clear (!), (8-), 13-19 X 

20-30 (-40) pm, I+ blue. 

CHEMISTRY.-PSOrOmiC acid. 

HABIrAT.-Lowland secondary growth. 

DIscussIoN.-The spores of G. coltim bina are 

clear in all stages, and hence Zahlbruckner’s transfer to Phaeographina is unnecessary. This species is 

part of the complex centered around G. incrustans. 

It is most easily distinguished from its relatives by 

the presence of psoromic acid, a very rare compound in the Graphidaceae. 

SPECIMEXS EXAMIxED.-Roseau Botanic Garden, 100 ft, 

Wirth 446 (US). 

43. Graphina dimidiata 

FIGURE 9d; PLATE 86 

Graphina dimidiata (T’ainio) Zahlbruckner, 1923:404. 

Graphis dimidiata Yainio, 1890:108 [type collection: Brazil, 

Vainio 322 (TUR, lectotype)]. 

DESCRIPTIos.-Thallus grayish to off-white, cow 

tinuous to cracked, glossy. Apothecia black, slightly 

raised, straight to somewhat flexuose, rarely 

branched, occasionally with a low thalline margin, 

0.5-2 (-3) mm long, slender. Disc usually not visible in surface 1-iew, occxionally appearing as a 

lighter line between the labia. Ascocarp transverse 

section: hymenium 90-1 15 pm high, epihymenium 

darkened, I-; exciple open below, lips entire, convergent to spreading, carbonized only on the upper 

half or less. Spores (?4) 8/ascus, 2-4 X 4-6 locular, 

(8-) 10-15 x (13-) 18-26 pm, I+ blue. 

CHEILIISTRY.-KO substances present. 

HABITAT.-Rain forest, mossy forest. 

DIscussIoK.-Graphina dimidiata is distinct 

among the small-spored GI-nphinrt species in its 

dimidiate, entire exciple, which is carbonized only 

in the upper portion of the labia. 

One of the Dominican specimens (Hale 35466) 

has the thallus irregularly spotted with a bright 

pink pigment. This coloration is not restricted to 

the apothecial margins, as in Phaeographis albida, 

and seems to be either an injury or infection. 
NUMBER 40 37 

SPECIMENS ExAMlNED.-Madjini, 100 ft, Hale 35697 (us); 

Freshwater Lake, 2600-2800 ft, Hale 35466 (US). 

44. Graphina dispersa 

FIGURE 9e; PLATE 8c 

Graphina dispersa Redinger, 1933:67 [type collection: Brad, 

Malme 2038B (S, lectotype)]. 

DEscRIPTIoN.-Thahs in the type epilithic and 

scattered, in the Dominican specimen corticolous, 

continuous, pale tan, matte. Ascocarps raised, in the 

type mostly whitish laterally, in the Dominican 

specimen whitish to concolorous with the thallus, 

in the type mostly 1 mm long, in the Dominican specimen 1-3 mm long, both rarely branched, straight 

to slightly curved, both frequently pale to reddish 

tan above. Disc not visible in surface view. Ascocarp transverse section: hymenium 70-100 pm high 

in type, 130-150 pm in the Dominican specimen, 

I-; exciple pale red-brown, closed, labia convergent, 

internally striate. Spores 6-8/ascus, muriform, 9-12 

X 20-30 pm, I+ blue. 

CHEMISTRY.-Norstictic acid. 

HABITAT.-virgin upland rain forest (Dominica 

only). 

DIscussIoN.-The illustration of the exciple that 

accompanies Redinger’s original description is incorrect, in that the exciple is shown as quite dark. 

Thin sections are difficult to prepare from specimens 

on rock; presumably the illustration was taken from 

a rather thick section. The Dominican material differs from the type in being corticolous and more 

vigorous overall. 

Graphina dispersa is similar to G. chlorocarpa 

but differs in spore number and size and in chemistry. Also similar is Graphina carneoviridis, which 

differs in having smaller spores, a paler more poorly 

developed exciple, and a different chemistry. 

SPECIhfEN EXAMINED.->fOrne Diablotin, 2200-2600 ft, Hale 

38090 (US). 

45. Graphina frumentaria 

FIGURE 9f; PLATE 8d 

Graphina frumentarh (Fee) MuelIer Argoviensis, I880:40. 

Graphis frumentaria Fee 1824:45 [type collection: Peru, 

Mutis s.n. (G, lectotype)]. 

DEscRIPTIoN.-Thallus buff-gray, smooth to somewhat rugose, glossy, occasionally cracked. Ascocarps 

very large and emergent, covered to the apices by 

the thalline margin, usually unbranched, straight 

to somewhat curved, 1-4 mm long, to nearly 1 mm 

wide. Ascocarp transverse section: hymenium 250- 

300 pm high, I-; exciple yellow, occasionally redbrown below, open, labia convergent, more or less 

intact, covered completely by pale thalline tissue. 

Spores 6-8/ascus, densely muriform, 18-24 X 42-60 

pm, I+ blue. 

CHEMISTRY.-NO substances present. 

HABITAT.-virgin rain forest. 

DIscussIoN.-Graphina frumentaria is somewhat 

similar to G. chlorocarpa but differs in its very large 

ascocarps, entire labia, and in having 6-8 spores 

per ascus. The description and illustration of this 

species in Redinger (1933:69) are probably incorrect in that the specimen has labia with internal 

striae and the overall size is distinctly less than the 

type. 

SPECIMEN ExAhrINED.-Can-Dom logging area, Newfoundland, 800 ft, Hale 35242 (US). 

46. Graphina illinata, new combination 

FIGURE 9g; PLATE 8e 

Graphis illinata Eschweiler in Martius, 1833:82 [type collection: Brazil, s.n. (M, lectotype)]. 

DEscRrPTIor\..-Thallus slate gray to whitish, 

smooth, glossy, thick. Ascocarps very protuberant, 

straight to occasionally flexuose, usually unbranched, covered completely by the thalline margin, only rarely showing a trace of the black exciple, 

0.5-6 mm long, mostly ca. 1 mm wide. Disc not 

visible in surface view. Ascocarp transverse section: 

hymenium 350-710 (!) pm high, I-; exciple black, 

massively closed, labia more or less convergent, 

more or less entire, completely covered by the thalline margin. Spores 1 iascus, densely muriform, 

30-50 X 110-225 pm, I+ blue. 

CHEMISTRY.-NO substances present (trace norstictic acid on one specimen). 

HABITAT.-virgin upland rain forest. 

DIscussIoN.-The traditional concept of G. illinata, i.e., as an earlier name for Graphis anguilliformis, is quite incorrect. The lectotype has one 

very large muriform spore per ascus and thus must 

be transferred to Graphina. It is interesting to note 

that this very large, very distinct and obvious species was found in Dominica by only one of the four 
SMITHSOXIAS CONTRIBUTIOSS TO BOTANY 38 

collectors. The implication, of course, is that many 

other less conspicuous canopy species remain uncollected on the island. 

SPECIMENS Ex.AwI\ED-Can-Dorn logging area, Biantridge 

Estate, 1700 ft, Hale 35276, 35281, 35284 (US); Can-Dom logging area, Pont Casse, 2000 ft, Hale 35082, 33083, 35124 (US). 

47. Graphina incrustans 

FIGURE 9h; PLATE 81 

Graphina incrustans (Fee) Mueller Argoviensis 1887:47. 

Fissurina iricrustarrs Fee, 1824:60 [type collection: South 

Graphis rubigitiosa Fee, 1824:47 [type collection: South AmerGraphina rubiginosa (Fee) Mueller Argoviensis, 1887:44. 

Graphis glaucoderina Xylander ex Tuckerman, 1888: 124 (type 

Graphis dehiscens l'ainio, 1890: 11 1 [type collection: Brazil, 

Fissurina nitidescens Nylander, 1890: 108 [type collection: 

Graphina glaucoderma (Xylander ex Tuckerman) Mueller 

Graphis nitidescens (Nylander) Vainio, 1915: 151. 

Graphina nitidesceiis (Sylander) Riddle, 1918: 115. 

Graphina dehiscens (Vainio) Redinger, 1933: 13. 

America, s.n. (G, lectotype)]. 

ica, s.n. (G, lectotype)]. 

collection: Cuba, Wright 61 (FH, lectotype)]. 

Vainio 306 (TUR, lectotype)]. 

Florida, USA, Calkins 31 (H, lectotype)]. 

Argoviensis 1895b347. 

DEScRrPTroN.-Thallus yellow-brown to yellowgreen, glossy to somewhat matte, smooth, thick, 

upper layer distinctly prosoplectenchymatous in 

transverse section. Ascocarps fissurine, starting as a 

crack flush with the thallus, finally gaping, occasionally branched, usually concolorous with the 

thallus, nearly round to quite elongate and flexuose, 0.5-6 mm long. Disc exposed but too sunken 

to be easily visible in surface view. Ascocarp transverse section: hymenium (45-) 75-150 pm high, I-, 

epihymenium slightly darkened; exciple rudimentary, yellow to pale red-brown, more or less open, 

labia quite convergent to quite spreading, apices 

of lips sometimes slightly darkened, with much included bark. Spores 8/ascus, 1-2 (-3) x 4-6 locular, 

6-11 x 13-35 pm, I- or I+ slow, reddish. 

CHEMISTRY.-NO substances present. 

HABiTAT.-Rain forest and elfin forest. 

DIscussIoN.-Graphina incrustans is a member of 

a species complex which appears to parallel the 

fissurine Graphis species (see Graphis triticea). It is 

a sporomorph of Graphis dumastii; undoubtedly a 

whole series of sporomorphs will be found between 

these two groups. 

The species listed in synonymy overlap with each 

other completely and cannot be maintained on any 

reasonable grounds. TSe have concentrated on Neotropic species, but many Paleotropic types will undoubtedly be involved here also; we have not yet 

seen TLC analyses for these. 

Most similar to G. incrustans is probably G. 

babingtonii (Montagne) Zahlbruckner; this species 

is tentatively separable by its more strongly gaping 

ascocarps and in having only 4-6 spores per ascus. 

Graphina insculpta is morphologically identical, 

separable only in having one very large spore per 

ascus. 

SPECIMEXS Ex4\fIsED.-hficotrin, 3000 ft, Wirth 465 (us); 

South Chiltern Estate, 1200-1500 ft, Imshaug 32802 (MSC); 

Freshwater Lake, 2600-2877 ft, Imshaug 32857A (MSC). 

48. Graphina insculpta 

FIGURE 10a: PLATE 9a 

Grnphina insculpta (Eschweiler in hfartius) hfueller kgoviensis, 1888b:307. 

Diovgrna insculpta Eschweiler iri ;\fartius, 1828:9 [type collection: Brazil, s.n. (M, lectotype)]. 

Graphis iriscdpta (Eschweiler iii Martius) Sylander, 1863:371. 

DEsCRIPTIos.-Thallus greenish to yellow-brown, 

smooth, gloss), thick, quite prosoplectenchymatous 

in section. Ascocarps fissurine, concolorous with the 

thallus, usually unbranched, straight to slightly 

curved, 0.5-1.5 mm long. Ascocarp transverse section: hymenium 90-120 pm high, I-; exciple rudimentary, yellow to pale red-brown, open, labia 

convergent to spreading, with much included bark. 

Spores l/'ascus, densely muriform, 20-30 X (50-) 

70-75 (-100) pm, I- or I+ slow, reddish. 

CHEMISTRY-NO substances present. 

HABITAT.-Rain forest. 

DISCLJSSI~K.-G?-U~~~~U insculpta is morphologically identical to G. incrustans and can be separated only by having one very large spore per 

ascus. 

(US). 

SPECIMENS EX.4alINED.-Near Pont Case, 1400 ft, Hale 37819 

49. Graphina macella 

FIGURE lob; PLATE 9b 

Graphina inacella (Krempelhuber) Mueller Argoviensis, 1880: 

39. 
NUMBER 40 39 

, 

FIGURE 10.-Cross sections of apothecia of Graphina: a, G. insculpta (lectotype in G); b, G. 

macella (Glaziou 6289B); c, G. marcescens (lectotype in G); d, G. nuda (Hale 35119); e, G. 

platyleuca (Thiebaut s.n.); f, G. plurispora (Malme 998); g, G. pseudoanaloga (Duss 1590); 

h, G. rufopallida (Duss 527). 
40 SMITHSOXIAN COSTRIBUTIOXS TO BOTANY 

Graphis macella Krempelhuber, 1876:380 [type collection: 

Brazil, Glaziou 6289b (RI, 1ectot)pe)l. 

DESCRIPTioN.-Thallus gray to white, continuous, 

smooth to lightly rugose, glossy to somewhat matte. 

Ascocarps raised but not prominently so, straight 

to curved, usually unbranched, apically black or 

black with thin white thalline stripes, laterally 

covered by the thalline margin, 1-5 mm long, less 

than 0.5 nim wide. Disc not visible in surface view. 

Ascocarp transverse section: 1i)menium 140-160 

pm high, I-; exciple black laterally and above, frequently brown or irregularly Carbonized below, 

closed to nearly open, labia convergent, striate. 

Spores l/ascus, densely muriform, (20-) 2540 x 

(-65) 95-130 pm, I+ blue. 

CHEMISTRY.-NO substances present. 

HABITAT.-Rain forest (Dominica). 

DIscussroN.-Graphina macella is similar to G. 

acharii but is smaller overall, more sunken, consistently has monosporous asci, and has an irregularly carbonized excipular base. In addition to the 

Brazilian and Dominican material, we have seen 

speciments from llexico, indicating that this species will probably be found throughout the Neotropics. 

35723 (US). 

SPECIMEN EXAMINED.--\’alley of Desolation, 2800 ft, Hale 

50. Graphina marcesceizs 

FIGURE 1Oc; PLATE 9c 

Graphina marcescens (Fee) Mueller Argoviensis, 1887:42. 

Graphis marcescens Fee 1824:38 [type collection: South America, Humboldt et Bonpland s.n. (G, lectotype)]. 

Graphis intricata Eschweiler in Martius, 183359 (non 

Graphis intricata Fee, 1824:42) [type collection: Brazil, 

Martius s.n. (hf, lectotype)]. 

Graphina intricata (Eschweiler in Martiusj hlueller Argoviensis, 1888b:510. 

Graphilia plittii Zahlbruckner, 1928:60 [type collection: Florida, USA, Plitt s.n. (US, lectotype; L, isotype)]. 

DEscRIPTIoS.-Thallus off-white, continuous, 

matte to nearly powdery. Xscocarps more or less 

emergent, rarely branched, quite flexuose, completely covered by a concolorous thalline margin 

which is easily dislodged, exposing the black exciples below, 1-6 mm long, ca. 0.3 mm wide. Ascocarp transverse section: hymenium (60-) 75-120 

(-140) pm, high, I-; exciple black above, more or 

Iess closed below by paler yellow to red or brown 

tissue, labia convergent, entire to slightly crenate, 

completely thalline covered. Spores 8/ascus, 1-2 X 

4-5 locular, 5-8 x 12-16 pm, I+ blue. 

CHEMrsmY.-Salazinic acid. 

HABIrhT.-upland rain forest (Dominica; see 

remarks). 

DIsccssIoh-.-Graphina marcescens is unlikely to 

be confused with any other Graphina species. Three 

characteristics account for this individuality: The 

red-brown exciple base, the presence of salazinic 

acid, very raie in the family, and the powdery thalline covering, easily rubbed, completely covering 

the ascocarps. Externall), G. marcescens is identical 

to Graphis candidmima Zahlbruckner (1923) but is 

easily separable from that species by spores and 

chemistry. 

It seems likely that this species will be found to 

have a wide range, both geographically and ecologically. The Dominican specimen (and probably the 

type of G. mal-cescens) is from high virgin rain 

forest; the type of G. plittii is from Florida, and 

hence probably lowland forest. In addition, we 

hale seen a specimen from 2200 meters, collected in 

India (Hale 35312X), which differs from the Neotropical material only in being somewhat larger 

and in having the exciple base a bit more carbonized. 

SPECIlIEV Ex~~IhED.-can-Dom logging area, Dleau Gommier, 1600-1700 ft, Hale 34312.4 (US). 

51. Graphina nuda 

FIGURE 10d: PLATE 9d 

Graphina nicda Magnusson, 19553266 [tvpe collection: Mauna 

Kea, Haw‘iii, Faurie 1023b (UPS, holotype)]. 

DEscRIPTIox.-Thallus gray to tan-gray, continuous, smooth to minutely roughened. Ascocarps 

very prominent, black, straight to slightly curved, 

unbranched, without a thalline margin, resembling 

Graphis adpl-essa, 1-2 mm long, slender. Disc not 

visible in surface view. .4scocarp transverse section: 

hymenium 135-150 pm high, I-; exciple black 

usually closed but occasionally irregular at the base, 

lips intact, convergent. Spores 8/ascus, uniseriate, 

1-4 x 4 locular, 16-18 X 20-30 pm, I+ blue. 

CHEJIISTRY.-NO substances present. 

HABITAT.-coastal cocos to virgin upland rain 

DISCLWON.-GYU~~~~U nuda is distinctive among 

forest; apparently of wide ecological amplitude. 
NUMBER 40 41 

Dominican Graphina species in its short, Opegrapha-like ascocarps, externally (and internally) 

indistinguishable from Graphis adpressa. Closely 

related to G. nuda is G. substriatula (Nylander) 

Zahlbruckner (1923), which has larger spores and 

is striate in some portions of the type. Also similar 

is G. sulcatula Mueller Argoviensis (1888d), which 

has longer, sinuous ascocarps, and exciples that are 

nearly open and nearly crenate. Graphina ruiziana 

(FCe) Mueller Argoviensis (1887), which is externally and internally very similar, differs in having 

much smaller spores. 

SPECIMENS EXAMINED.-Prince Rupert Bay, Zmshaug 33536.4 

(MSC); Can-Dom logging area, Pont Cad, 2000 ft, Hale 

35092, 35119 (US). 

52. Graphina platyleuca 

FICURE 10e; PLATE 9e 

Graphina platyleuca (Nylander) Zahlbruckner, 1923:420. 

Graphis platyleuca Nylander, 1868:114 [type collection: New 

Caledonia, Thiebaut s.n. (H, lectotype)]. 

Graphina platycarpa Fink, 1927:219 (non G. platycarpa (Eschweiler in Martius) Zahlbruckner, 1902:389) [type collection: 

Puerto Rico, Fink 1774 (MICH, lectotype)]. 

Graphina platycarpinu Zahlbruckner, 1932:Z 14 (nomen noviim 

for G. platycarpa Fink). 

DEscRIPTIorv.-Thallus greenish white, continuous or occasionally cracked, matte to nearly powdery-mealy. Ascocarps usually strongly raised, 

rotund to somewhat elongate, rarely branched, 

labia completely covered by the thalline margin, 

1-2 (-5) mm long, ca. 1 mm wide. Disc prominent, 

concolorous to brownish black, lightly pruinose. 

Ascocarp transverse section: hymenium 150-250 

pm, high, I + blue, epihymenium dark; exciple 

irregularly carbonized, quite black in parts, redbrown in other patches, closed, lips quite divergent, 

entire. Spores 1 iascus, densely muriform, 3540 X 

120-165 pm, I+ blue. 

CHEMIsmY.-Protocetraric acid only (types of G. 

platyleuca, G. platycarpa), or protocetraric acid 

with salazinic acid and norstictic acid (see discussion). 

HABITAT.-PrimarY rain forest (Dominica). 

DI~CUSSION.-AS presently recognized, G. platyleuca is a chemically heterogeneous species. However, protocetraric acid is quite rare in the Graphidaceae and serves to tie the Dominican specimen 

into the taxon. The only other material we have 

seen that is referable here is the “type” of one of 

Nylander’s nomina nuda, Graphis glaucoleuca, 

from Cuba. This specimen is chemically like the 

Dominican material. Additional collections may reveal that G. platyleuca shows similar chemical variation to that found in Phaeographis exaltata. 

Most similar to G. platyleuca is G. confluens 

(Fee) Mueller Argoviensis (1887); the latter species 

can be distinguished by somewhat less emergent 

ascocarps and a different chemistry (lichexanthone, 

stictic acid, and constictic acid). 

ft, Hale 37955 (US). 

SPECIhIEN ExAMINED.-Dleau Gommier Forest Reserve, 1200 

53. Graphina plurispora, new combination 

FIGURE lo!; PL.4TE 9f 

Graphina pseud osop h ist ica var. plurispora Redinger , 193 3: 36 

[type Collection: Brad, Malme 998 (S, lectotype)]. 

DEscRIPrIorV.-Thallus continuous, gray-white, 

smooth and glossy to matte (on the same thallus). 

Ascocarps barely raised, black above, frequently 

with white stripes, with a well-developed thalline 

margin laterally, quite flexuose, 2-5 mm long, slender. Disc not visible in surface view. Ascocarp 

transverse section; hymenium 60-100 pm high (on 

the same specimen), I-; exciple very variable 

(within one ascocarp), black, distinctly oI:en to distinctly closed, lips convergent, perfectly intact to 

quite striate. Spores begining as Siascus, but only 

(2-) 4-6 maturing, muriform, 1-3 X 8-13 locular, 

10-15 X (27-) 35-50 (-60) pm, I+ blue. 

CHEMISTRY.-NO substances present. 

HABITAT.-cultivated areas above 1500 feet 

(Dominica only). 

DrscussIoN.-Redinger’s variety differs from the 

type of Graphina pseudosophistica in two importain respects. The latter has consistently monosporous asci and has the “quintaria” unknowns. 

Because of these differences, we have raised the 

variety to species rank. Still, Graphina plurispol-a 

is both highly variable and part of a yery confusing 

array of species. Most closely related are the following 

Graphina disseypens (Nylander) Mueller Argoviensis (1880) is tentatively separable because of the 

quite dendritically branched ascocarps and consistently open exciples. 

Graphina elongatoradians Fink (1927) is another 
42 SXIITHSONIAN CONTRIBUTIONS TO BOTANY 

dendritically branched relative with longer, more 

slender ascocarps, with open exciples, and a different chemistr) (two unknown substances). 

Graphzna subvelata (Stirton) Zahlbruckner (1923) 

differs in being non-striate and in having open exciples. See H,i\wdrd (1gi8) foi d discussion of 

G. subuelata and its near relatives, including New 

JVorld species. 

In addition, G. plurispora bears a number of 

similarities to the G. antzllarum complex; for a 

more complete discussion, see the latter species. 

1500 ft, Irnshaug 32i24.A, 32727 (MSC) 

SPECIMEM EXAVIXED -South Chilteiti F\tate Road, 1500- 

54. Graphina pseudoanaloga 

FIGURE log; PLATE 10a 

Graphina pseudoanaloga (Vainio) Zahlbruckner, 1923:421. 

Graphis pseudoanaloga L’ainio, 1915:145 [type collection: 

Guadeloupe, Duss 1590 (TUR, lectotype)]. 

DEScRIPTIox.-Thallus gray-white, thin, continuous, matte. Ascocarps slightly raised, black above, 

straight to flexuose, rarely branched, covered about 

half-way by the thalline margin, 0.5-4 mm long, 

slender. Disc not visibIe in surface view. Ascocarp 

transverse section: hymenium 75-1 15 pm high, inspersed (apparently clear in TVirth 526), epih)menium dark, I-; exciple black, closed to occasionally 

nearly open, labia convergent, entire to barely 

crenate, rarely with an occasional stria (all on one 

thallus). Spores 8/ascus, 1-3 X 7-10 locular, 9-12 

x 22-45 pm, If blue. 

CHEl\zISTRY.-r\TOrStiCtiC acid. 

HABITAT.-Beach to upland virgin rain forest; 

apparently of wide ecological amplitude. 

DIscussIoN.--Graphina pseudonnaloga is very 

similar to G. annloga (Nylander) Zahlbruckner 

(1923), which differs in having smaller spores and 

an open exciple. 

SPECIMENS ExAMISED.-Cabrit, sea level, Wirth 526 (US); 

Can-Dom logging area, Brantridge Estate, 1700 ft, Hale 33283 

(US). 

55. Graphina rufopallida 

FIGURE 10h; PLATE 10b 

Graphina rufopallida (T’ainio) Zahlbruclner, 1923:423 (“rufoGraphis rufopallida T’ainio, 1915:149 [type collection: Guadepallens,” sphalm.). 

loupe, Duss 527 (TUR, lectotype)]. 

ford Estate, Dominica, Elliott 1852 (TUR, lectotype)]. 

Graphis verrnicularis \.ainio, 1915: 148 [type collection: ShawCraphina verrniczilaris (L’ainio) Zahlbruckner, 1923:430. 

DEscRIPTIos.-Thallus off-white, smooth, continuous. Ascocarps somewhat emergent, tan to redbrown, rarely branched, flexuose and intertwined, 

1-7 mm long, slender. Disc not visible in surface 

view. Ascocarp transverse section: hymenium 

70-100 pm high, I-; exciple more or less open, 

labia convergent, more or less striate, dark redbrown or fuscescent apically (but not carbonized), 

paler yellow-brown below. Spores 6-81immature 

ascus, apparently always aborting to 2-4/mature 

ascus, densely muriform, 15-30 X 36-75 pm, I+ 

blue. 

CHEMISTRY.-NO substances present. 

HABITAT.-upland rain forest. 

DISCUSSIOS.-G~U~~~~U rufopallida is most similar to G. bulbisii (see discussion under G. chlorocarpa), from which it differs in the lower, much 

longer, more slender and intricating ascocarps. 

Itre have seen no specimens of this species other 

than the two types cited. 

SPECIMEN ExAxrrsm.-Shawford Estate, Elliott 1852 (TUR). 

56. Graphim suberythrella, new species 

FIGURE lla; PLATE 1Oc 

DEscRIPTIox.-Thallus continuus, laevis, stratum 

corticale bene evolutum. Apothecia subimmersa, 

flesuosa. vulgo iimplicia, 2-4 mm longa. disco lato, 

pruinoso. Excipulum integrum, flavidum vel rufoferrugineum, labiis integris, divergentibus. Hymenium 110-140 pm altum, I+ coeruleum. Asci 8 

spori; sporae decolores, murales, loculis horizontalibus 5-7, loculis transversis 1-3, 10-14 X 17-28 pm, 

I + coeruleae, in juventute halonae indutae. 

Thallus gray, glossy, continuous, rather thick, 

cortical layer compact and distinct. Ascocarps 

barely raised, flexuose, occasionally branched, 2-4 

mm long, slender, concolorous with the thallus. 

Disc broad, dark gray, lightly pruinose. Ascocarp 

transverse section: hymenium 110-140 pm high, at 

least the epihymenium I+ blue; exciple J-ellow to 

red-brown, closed, lips intact and divergent. Spores 

8/ascus, 1-3 X 5-7 locular, halonate in early stages, 

10-14 X 17-28 pm, I+ blue. 

CHEMISTRY.-NOrStiCtic acid. 
NUMBER 40 43 

FIGURE 11.-Cross sections of apthecia of Graphina: a, G. suberythrella (Imshaug 32699); b, 

G. triphoroides (Hale 35161); c, G. vestitoides (Fink 1986); d, G. virginea (Wirth 547); e, G. 

virginea (lectotype of G. triangularis, Pringle 17); j, G. virginea (lectotype of G. obtectula, 

Tonduz). 

HoLorYPE.-Cultivated area, South Chiltern Es- G. erythrella (Montagne) Zahlbruckner (1923), but 

tate Road, Dominica, elevation 1500-1700 feet, differs in its more immersed ascocarps, gray disc, 

Imshaug 32699 (MSC); isotype, US). I- hymenium and larger spores. Also similar is 

DISCUSSION.--G~~~~~~U suberythrella resembles G. insignis (Vainio) Zahlbruckner (1923), which dif- 
44 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

fers in having wider ascocarps and smaller spores, 

and in lacking both norstictic acid and the I+ 

hymenium. 

SPECIMEX ExAMIsED.-can-Dom logging area, Newfoundland, 800 ft, Hale 35224 (US). 

57. Graphina triphoroides, new species 

FIGURE llb: PLATE 10d 

DEscRIPTIoN.-Thallus continuus, farinosus. Apothecia alte sessilia, simplicia, margine thallino 

usque ad verticem vestita, 1-6 mm longa, 1 mm 

lata. Excipulum dimidiatum, fuligineum, labiis integris vel irregulariter crenulatis, erectis vel convergentibus; hymenium 250-700 pm altum, I-; Asci 

6 (-8?) spori; sporae decolores, murales, loculis horizontalibus 7-10, loculis transversis 2-4, 18-30 X 

47-75 pm, I+ coeruleae. 

Thallus white to off-white, continuous, mealy. 

Ascocarps large, very protuberant, unbranched, 

covered to the apices by a mealy concolorous thalline margin, irregularly blackened where the covering is rubbed, 1-6 mm long, I mm wide. Disc 

usually not visible in surface view. Ascocarp transverse section: hymenium 250-700 (!) pm high, I-; 

exciple black, more or less yellow-brown and open 

below, labia erect to convergent, entire to somewhat irregular in outline. Spores 6 (-S?)/ ascus, 

muriform, 24 X 7-10 locular, 18-30 X 47-75 

pm, I+ blue. 

CHEJfISTRY.--StiCtiC acid. 

HoLoTYPE.-can-Dom logging area, Dleau Gommier, 1600-1700 ft, Hale 35161, January 1969 (US). 

DIsccssIoN.-Graphina triphoroides is part of a 

very distinct group of three New r\'orld species, 

all characterized by very large, thalline-covered apothecia and dimidiate, carbonized exciples. Graphina 

triphol-a (Sylander) Alueller Argoviensis (1880) differs in having 34 spores per BSCLIS, no lichen acids, 

and carbonization restricted more to the upper portions of the labia. Graphina cleitops (Fee) RIueller 

Argoviensis (1887), the third species, differs in being 

monosporous. 

SPECIMENS Ex~~~~m.--Can-Dorn logging area, Dleau Gommier, 1600-1700 ft, Hale 35128 (US); Wi-ight 15, Cuba (US) 

("type" of Graphis triphomides Kylander, unpublished 

name). 

58. Graphina vestitoides 

FIGURE llc; PLATE 10e 

Graphina vestitoides Fink, 1927:218 [type collection: Puerto 

Phaeographis cervicuiata Redinger, 1933:76 [type collection: 

Rib, Fink 1986 (MICH, lectotype)]. 

Brazil, Maline 3531 (S, lectotype; UPS, isotype)]. 

DEscRIPrIos.-Thallus gray, continuous, smooth. 

Ascocarps quite protuberant, frequently branched, 

flexuose, varying from black above and partially 

covered by a thalline margin to concolorous (completely covered), to 10 mm long, ca. 0.5 mm wide. 

Ascocarp transverse section: hymenium 150-2 10 pm 

high, I-; exciple black, usually closed, lips convergent, striate to nearly entire. Spores 4-6 (-8)/ascus, 

biocellate only in the terminal few sections, 12-18 

X 85-140 pm, I+ blue. 

CHEMISTRY.-NO substances present. 

HhBITAT.-upland virgin and secondary forest. 

DIscr,ssIos.-C;raphina vestitoides is separable 

from G. acharii only in the peculiarly biocellate 

spores, which additional collections may prove to 

be an unreliable character. 

The type material of Phaeographis ceruiculata 

has mature spores that are quite clear, and the 

specimen is identical to G. vestitoides. 

SPECIMENS ExAMrhm.-Freshwater Lake, 2600-2877 ft, ImsIzaug 32859'4 (MSC); near Pont Cad, 1400 ft, Wirth 544a 

(US). 

59. Graphina virginea 

FIGURE lld,e,f; PLATE lOf 

Graphina virginea (Eschweiler in Martius) Mueller -4rgoviensir, 1880:41. 

Leiog~amma virgineum Eschweiler in Martius, 1833:98 [type 

collection: Brazil, Afartiiis s.n. (XI, lectotype)]. 

Graphinn obtectdn Mueller Argoviensis, 1892b: 133 [type collection: Costa Rica, Pitfier (3166 (G, lectotype); Figure llf)]. 

Graphina t~ielaleucn Miieller Argoviensis, 1895,: 144 [type collection: Roscau, Dominica, Eckfeldt 70 (G, lectotype)]. 

Grnfihina paluiei-i Zahlbruckner, 1921:231 [type collection: 

Mexico, Pringle 9 (MICH, isotype)]. 

Grnpliina triangiiinris Zahlbruckner 1921 :232 [type collection: 

Mexico, Priizgle 17 (MICH, isotype)]. 

Graphis collospot~elin I'ainio, 1923: 141 [type collection: Trinidad, Tliaxter 10 (TUR, lectotype)]. 

Graphifin collosfiorelin (Vainio) Zahlbruckner, 1923:212. 

DESCRIPTIO~.-~~~~I~~LIS greenish white to gray, 

thick, continuous except for a fissure usually paralleling each ascocarp, smooth to mealy, occasion- 
NUMBER 40 45 

ally quite rugose (when ascocarps are densely clustered), Ascocarps nearly flush with the thallus, 

rarely somewhat raised, margins concoloi ous to 

much paler than the thallus, occasionally branched, 

flexuose, 1-5 mm long, slender. Disc in surface view 

varying from concealed to quite broad, concolorous 

to darker than the thallus. Ascocarp transverse section: hymenium 120-200 pm high, epihjmenium 

darkened, at least the epihymenium I+ blue. 

Exciple yellowish, rudimentary, sometimes thickened below and/or with a basal dark (nearly carbonized) band (see discussion), labia more or less 

divergent, occasionally with internal striae (see discussion) but otherwise entire. Spores 2-3, 4-6 or 

8/ascus (!), most numbers occurring within a single thallus, 1-4 (-7) X 11-20 locular, (12-) 15-25 X 

50-80 (-130) pm, I+ blue. 

CHEMISTRY.-At least one of the 3 so-called 

“Quintaria” unknowns; trace of stictic acid and 

constictic acid (type of G. uzrgznea only). 

HABITAT.-secondary growth and cultivated trees 

and shrubs, from sea level to over 1000 meters. 

DIscussrox.--In its usual form Graphina uirginea 

is an extremely vigorous species, forming thalli up 

to 15 cm in diameter. The thallus is normally 

greenish white, matte, and continuous except for 

characteristic fissures bordering the ascocarp margin. The ascocarps themselves are commonl) slightly 

paler than the rest of the thallus, and do not occur 

in close proximity to each other. In all likelihood, 

the species is weedy and of rapid growth, as large 

colonies are commonly found on such fast-growing 

cultivated plants as cacao and blue Mahoe (HibisInternally, this most common form of the species 

has a rather rudimentary exciple and a very variable number of spores per ascus. 

Morphologically and anatomically, Graphina 

collosporella and G. obtectula cannot be separated 

from the normal form of G. virginea. 

The other taxa listed as synonyms all show some 

cus). 

structural variation from the typical pattern. In 

the case of G. palmeri and G. melaleuca the thallus 

in surface view appears quite rugose and the ascocarps somewhat emergent. It seems quite likely 

that this appearance comes with age; dense clustering of more typical ascocarps, with their associated 

fissures, could easily produce such a form. A probable additional synonym for this “rumpled” variant of G. uirginea is G. monophora (Nylander) 

Zahlbruckner (1923). 

The type of G. melnleiica also has a broader, 

browner disc than is normally found in G. uirginea; 

externally this specimen is indistinguishable from G. 

mendax (Nylander) Mueller-Argoviensis (1 888a). 

The different disc is, however, a matter of degree 

rather than of kind; occasionally one can find such 

discs on parts of more typical thalli. 

Graphilia palmrri shows the internal striae that 

seem to arise with age and lateral compaction of 

the old hymenium; we do not find such striae to 

be at all dependable in distinguishing species (see 

Graphina colliculosa for a parallel variation). 

The internal striae, coupled with a more or less 

continuous semicarbonized band below the exciples, 

is also characteristic of the type of G. triangularis, 

which externally is much like typical virginea. 

The band is occasionally found in thalli of quite 

normal G. uirginea, frequently unconnected with 

the ascocarps. The thickened exciple base of the 

type of G. triangularis is another condition that 

we feel may well be simply a factor of age (again, 

see G. colliculosa for parallels). 

All the forms of G. virginea are tied strongly 

together by their chemistry. All have the rather 

rare It blue reaction of (at least) the epihymenium; all share at least two of the three “Quintaria” 

acids. 

SPEci\ftNs Ex4\iINEo.-Biookhi~1 Estate, 100-150 ft, Imshaug 33321.4 (MSC); Cocoa Centre on Layou Rixer, 200 ft, 

Wirth 517 (US); north of Pont Casse., 1400 ft, Wirth 543, 545, 

547, 548 (US). 

Key to the Species of Phaeographina 

1 Exciple with carbonized areas. 

2. Exciples carbonized and closed below ...........................).......)...............,....,..,.. 63. P. cf. diflormis 

2. Exciples carbonized only at labial tips. 

3. Ascocarps massive, 1-3 mm wide: no substances present ........_........... 61. P. caesiopruinosa 

3. Ascocarps slender, less than 0.4 mm wide: stictic acid present .,......,....... 65. P. oscitans 
46 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

1. Exciple uncarbonized. 

4. Labia convergent: spores more than 40 Gm long .................................................... 64. P. eZZiottii 

4. Labia divergent, exposing the disc; spores less than 40 &m long. 

mmonly over 5 mm long; norstictic acid present 

5 mm long; norstictic acid abse 

.......................................................... 60. P. atrmtermicdaris 

6. Spores 20-27 rm long; stictic acid present ................................. 

6. Spores 33-85 gm long; no substances present .............................................. 62. P. coriaria 

60. Phaeographina atrovermicularis, new species 

FIGURE 12a; PLATE lla 

DEscRIPTIoN.-Thal~us continuus, sublaevigatus. 

Apothecia adpresso-sessilia, vulgo 1-amulosa, 3-10 

mm longa, disco dilatato, nigro. Excipulum dimidiatum, fuscum, labiis integris, divergentibus; hymenium 145-220 pm altum, I-. Asci 6-8 spori; 

sporae fuscescentes, murales, loculis horizontalibus 

6-9, loculis transversis 1-4, 15-17 X 25-35 pm, 

I + coeruleae. 

Thallus thin, light tan, continuous to apparently 

eroded except near the ascocarps, smooth to matte. 

Ascocarps very long, raised, frequently branched, 

the pale thalline margin very prominent, 3-10 mm 

long, slender. Disc very prominent, black, occasionally faintly pruinose. Ascocarp transverse section: hymenium 145-220 ,urn high, I-, lightly beadedinspersed; exciple open below, laterally brownishblack, labia divergent, intact, bounded by a thick 

parenchymatous thalline margin. Spores’ 6-8/ascus, 

1-4 X 6-9 loculed, 15-17 x 25-35 pm, I+ blue. 

CHEMISTRY.-NOrStiCtiC acid. 

HoLoTYPE.-can-Dom logging area, Brantridge 

Estate, Dominica, elevation 1700 feet, Hale 35278 

DISCUSSION.-B~ virtue of its long black ascocarps, Phaeographina atrovermicularis is quite distinct and is apparently not closely related to any 

other New World species. Most similar is P. intercedens AIueller Argoviensis (1 888e), which differs 

in having shorter, broader, stockier ascocarps, larger 

spores, an unknown acid in addition to norstictic, 

and no trace of carbonization in the exciple. Also 

somewhat similar is the widespread P. caesiopruinosa, which has much shorter, broader ascocarps, 

strongly pruinose discs, and lacks norstictic acid. 

SPFCIME\S ExAMINrD.-can-Dom logging area, Pont Cassk, 

2000 ft, Hale 35093 (US); rain forest, Jean Estate, 2000 ft, 

Hale 35068 (US). 

(US). 

61. Pltaeographina caesiopruinosa 

FIGURE 12b; PLATE llb 

Phaeographina caesiopruinosa (Fee) Mueller Argoviensis, 

Arthonia caesiopruinosa Fke, 1837:36 [type collection: South 

1887:49. 

America, s.n. (G, lectotype)]. 

DEscRIPTIoN.-Thallus pale brown, smooth to 

slightly roughened, continuous. Ascocarps quite elevated, rarely branched, straight to flexuose, thalline 

margin reaching to disc, 1-10 mm long, 1-3 mm 

wide. Disc very prominent in surface view, lightly 

to very heavily pruinose. Ascocarp transverse section: hymenium 150-200 pm high, I-, epihymenium darkened; exciple black laterally, open below, 

lips divergent. Spores (4-) 6-8/ascus, densely muriform, 12-17 x 45-75 (-100) pm, I+ slow, reddish 

blue. 

CHEMISTRY.-NO substances present. 

HABITAT.-cUltivated areas, uplands. 

DIscussIoN.-Phaeographina caesiopruinosa is a 

very common New World species, unlikely to be 

misidentified. All other New World species with 

very large ascocarps and carbonized exciples have 

monosporous asci. 

Zmshaug 32731 (MSC). 

SPECIMEN EXAMINED.-sOUth Chiltern Estate, 1500-1700 ft, 

62. Phaeographina coriaria, new species 

FIGURE 12c; PLATE llc 

DEscRrmIoN.-Thallus continuus, laevis, coriaceus. Apothecia adpresso-sessilia, recta vel arcuata, 

simplicia vel raro ramulosa, 1-5 mm longa et ca. 

0.3 mm lata, disco dilatato, nigro, leviter pruinoso. 

Excipulum rufo-ferrugineum, dimidiatum, labiis integris, divergentibus; hymenium ca. 150 pm altum, 

I-. Asci 4-6 (-8?) spori; sporae fuscescentes, mu- 
NUMBER 40 47 

FIGURE 12.-Cross sections of apothecia of Phaeographina: a, P. atrovermicularis (Hale 35278); 

b, P. caesiopruinosa (lectotype in G): c, P. coriaria (Hale 35378); d, P. difjormis (Fink 1874); 

e, P. elliottii (Elliott 1338); f, P. oscitans (Mann s.n.). 

rales, loculis horizontalibus 8-10, loculis transversis 1-3, 9-10 X 33-35 pm, I+ rufo-fuscae. 

velum. Ascocarps slightly raised, straight to somewhat curved, rarely branched, 1-5 mm long, ca. 

0.3 mm wide. Disc broad, black, with a light pruThallus continuous, smooth, tan, resembling ina. Ascocarp transverse section: hymenium ca. 150 

pm high, beaded, epihymenium dark, I-; exciple 
48 SMITHSONIAS COXTRIBUTIOSS TO BOTASY 

rudimentary to very pale below, laterally red-brown, 

lips intact, divergent. Spores 4-6 (-8?) lascus, brown, 

8-10 x 1-3 locular, 9-10 x 33-35 pm, I+ red, 

slowly darkening. 

CHEMISTRY.-~O substances present. 

HOLOTYPE.--;\~OSSY forest, Morne Anglais, DOminica, elevation 3000-3600 feet, Hale 35378 (US). 

DIscussIoN.-Phaeographzna coriaria is most similar to the following three New 'CYorld species: 

Phaeographina arechaualetae Mueller Argoviensis 

(1888~) differs in having distinctly carbonized labia 

and rotund spores. 

Phaeographina caracasana Mueller Argoviensis 

(1887) differs in having much broader ascocarps 

with rounder apices and in having smaller spores. 

Phaeographina elliptica IVirth and Hale (1963) 

differs in chemistry and in having monosporous 

asci. 

63. Phaeographina cf. diflormis 

FIGURE 12d; PLATE lld 

Phaeographina diffomis Fink, 1927:220 [type collection: 

Aibonito, Puerto Rico, Fink 1874 (MICH, lectotype)]. 

DEscRIPTIoN.-Thallus thin, continuous, glossy, 

greenish gray. Xscocarps nearly round to elongate 

and irregular, ends round, occasionally shortl) 

branched, partly immersed, 0.8-1 3 mm long, ca. 

0.8 mm wide, with a raised thalline margin. Disc 

broad, brown-black, occasionally faintly pruinose. 

Ascocarp transverse section: hymenium 90-125 pm 

high, epihymenium very dark, I-; exciple brownblack, closed, lips intact, divergent. Spores 8/ascus 

in the type, 4/ascus in the Dominican specimen, 

densely muriform, pale to brown in the type, 

mostly pale in the Dominican specimen, 10-15 X 

(25-) 30-48 pm in the type, 18-22 x 55-75 in the 

Dominican specimen, I+ blue. 

CHEMISTRY.-UnknO\Vn in the type, no substances present in the Dominican specimen. 

HABITAT.-Elfin forest (Dominican specimen 

only). 

DrscvssIos.-;\ltliough the Dominican specimen 

is externally identical to the type, and is indistinguishable in cross-section, the spore size and number make this identification tentative. In addition, 

it should be noted that both specimens have large 

numbers of clear spores, making this species difficult to assign to genus. In fact, this species may 

represent a link to the smaller members of the 

Graphinn confluens alliance, such as G. cymbegraplzn (Nylander) Zahlbruckner (1 923), which is 

strikingly similar. 

Very similar to P. difformis is P. explicans Fink 

ex Hedrick (1933), which differs in its more elongate, rather sunken apothecia. 

SPECI\IEI\ Exi\rIzlED.-summit of Morne Diablotin, 4300- 

4350 ft, Imshaug 32913,4 (MSC). 

64. Phaeographina elliottii 

FIGURE 12e; PLATE lle 

Phaeographina elliottii (Vainio) Zahlbruckner, 1923:438. 

Graphis elliottii T'ainio, 1915:143 [type collection: Dominica, 

Roseau Valley, Elliott 1338 (TUR, lectotype)]. 

DEScRIPTIos.-Thallus thin, whitish, matte, continuous. Ascocarps slightly raised, rarely branched, 

straight to somewhat curved, thalline margin prominent, to 1.5 mm. long, slender. Disc concealed to 

open and dark gray. Ascocarp transverse section: 

hymenium completely (?) disintegrated in the type, 

I+ fide Vainio; exciple yellow-red, not well developed, lips convergent to quite spreading, tips 

brown, dark portions probably derived (at least 

in part) by lateral compression of the dark epihymenium. Spores (fide Vainio) 2-8/ascus, muriform, 

22-28 X 44-54 pm. 

CHEMISTRY.-NO substances present. 

HABITAT.--('On trees in Roseau Valley." 

DIscussIos.-The type of Phaeographina elliottii 

now seems to be completely sterile. Both externally 

and internally, the type specimen closely resembles 

Phaeographis albida; it may be possible that Vainio 

mistook entire disintegrating asci for spores. We 

have seen no other material that might be refera6le 

to this Phaeographina species 

65. Phaeographina oscitans 

FIGURE 12f; PLATE llf 

Phaeographinn oscitarzs (Tuckerman) Zahlbruckner, 1923:442. 

Graphis oscitans Tuckerman, 1868:231 [type collection: Oahu, 

Hawaii, Marin s.n. (FH, lectotype)]. 
NUMBER 40 49 

DEScRIPTIoN.-Thallus continuous, smooth to 

matte, off-white. Ascocarps slightly raised, flexuose, 

rarely branched, pale thalline margins prominent, 

1-2 mm long, slender. Disc prominent, gray-black. 

Ascocarp transverse section: hymenium 125-1 50 

pm high, I-; exciple rudimentary below, yellow laterally, lips slightly convergent to divergent, more 

or less carbonized apically, entire to once-sulcate. 

Spores 8/ascus, brown, 1-3 X 6-8 locular, 6-9 X 

(15-) 20-27 pm, IS blue. 

CHEMISTRY.-stiCtiC acid. 

HABITAT.-virgin upland rain forest (Dominica 

only). 

DIscussIos.-Phaeographjna oscitans is externally 

very similar to Phaeographis albida, but can easily 

be separated by spores and chemistry. Also quite 

similar is Phaeographina pachnodes (Fee) Mueller 

Argoviensis (1887), which differs in having larger 

spores, no trace of striae, and in having norstictic 

acid; and P. turgida (Fee) hfueller Argoviensis 

(1887), which would seem to be the TLC negative 

equivalent of P. pachnodes. 

SPEcnfExs ExAMIsED.-caii-Dom logging area, Pont Casse,

2000 ft, Hale 35127, 35131 (US). 

Incorrect and Omitted Names 

Graphis bonplandiae: See Graphis triticea for Elliott 533. 

Graphis cooperta: See G?-aphis anguilliformis for Elliott s. n. 

Graphis (Phaeographis) diversa: See Phaeographis exaltata for 

Graphis dziplicata: See Graphis rirntclosa for Elliott 125 and 

Graphis duplicata var. subduplicata Vaiiiio: Elliott I509 is 

Graphis lactea: Elliott 535 is the type of G. lactea var, 

Graphis lactea Xar. clausa: See Gyaphis insidiosa for Elliott 

Graphis lineola: See Graphis leptocarpa for Elliott 1535. 

Graphis (Phaeographis) lobata: The only Dominican collection (Elliott 1365 in TUR) is correctly identified to species, 

but it is not included in our list because the generic status 

is unclear. It is probably closer to Phaeotrenza in the 

Thelotremataceae. 

Graphis (Phaeographis) inedicsaeformis: Elliott 529 in BM is 

a fragmentary specimen, which may be an immature 

Phaeogr-aphis esaltata. There is no specimen so identified 

in TUR. 

Graphis (Graphina) sttbnitida: Elliott 171 is indeterminable. 

Graphis tenella \ar. epiphaea I‘ainio: Not found in TUR or 

Graphis (Graphiiia) verrnicularis: See Graphina rufopallida 

Elliott 526. 

Graphis flesibilis for Elliott 3954 pro parte. 

sterile. 

doiiiinicana T’ainio; see Graphis tachygrapha. 

1511. 

BM and presumed to be lost or misfiled. 

for Elliott 1852. 
Literature Cited 

Acharius, E. 

1810. Lichenographia universalis. 696 pages. Gottingae. 

1814. Synopsis methodica lichenum. 392 pages. Lund. 

1890. A Further Enumeration of Some Lichens of the 

United States. Bulletin of the Torrey Botanical Club, 

Eckfeldt, J. W. 

173255-257. 

Eschweiler, F. G. 

1828. 

1833. 

Fke, A. 

1824. 

1837. 

1874. 

Lichens. In C.F.P. Martius, Icones Plantarum Cryptogamicarum. 138 pages. Monachii. 

Lichens. In C.F.P. Martius, Flora Brasiliensis, volume 1, 390 pages. Stuttgart. 

Essai sur les cryptogames des ecorces exotiques officinales. 167 pages. Paris. 

Essai sur les cryptogames des ecorces exotiques, 11: 

Supplement and rekision. 137 pages. Paris. 

Materiaux pour une flore lichenologique du Bresil, 

11: Les Graphidkes. Bulletin de la Societe Botanique 

de France, 21:21-32. 

Fink, B. 

1927. Yew Species of Lichens from Porto Rico. Mycologia, 

19:206221. 

Groenhart, P. 

1965. Studies in Ascostromatic Lichen-Fungi, I: The 

Problem of Ascohymeniales and Ascoloculares. Persoonia, 4: 1-7. 

Morden-Smithsonian Expedition to Dominica: The 

Lichens (Parmeliaceae). Smithsonian Contributions 

to Botany, 431-25. 

Morden-Smithsonian Expedition to Dominica: The 

Lichens (Thelotremataceae). Smithsonian Contributions to Botany, 16:l-46. 

1976. Lichenes Americani Exsiccati, no. 156. Washington, 

D.C. 

Hayward, G. 

The Lichen Families Graphidaceae and Opegraphaceae in New Zealand. New Zealand Journal of 

Botany [In press.] 

Sew Genera and Species of Lichens from the Herbarium of Bruce Fink, I. Mycologia, 25:303-316. 

Hale, M. E., Jr. 

1971. 

1974. 

1978. 

Hedrick, J. 

1933. 

Hooker, J. D. 

1867. Handbook of the New Zealand Flora. 798 pages. 

London. 

Knight, C. 

1884. On the Lichenographia of New Zealand. Transactions and Proceedings of the New Zealand Institute, 

16:400-408. 

Contributions to the Lichenographia of New ZeaKnight, C., and W. Mitten 

1860. 

land. Transactions of the Linnean Society of 

London, 23: 101-106. 

Krempelhuber, A. 

1876. Lichenes Brasilienses, collecti a D. A. Glaziou in 

provincia brasiliensi Rio Janeiro. Flora, 59:378-384, 

412-4 16. 

Magnusson, A. H. 

A Catalogue of the Hawaiian Lichens. Arkiv for 

Botanik, 3( 10):223-402. 

Memorie Lichenographiche. 185 pages. Verona. 

Troisieme Centurie de Plantes cellulaires exotiques 

nouvelles. Annales Sciences Naturelles Botanique, 

series 2, 18:241-282. 

Cryptogamis Guyanensis seu Plantarum Cellularium 

in Guyana Gallica Annis 1835-1849 a C1. Leprieur 

Collectarum Enumeratio Universalis. Annales Sciences Naturelles Botanique, series 3, 16:47-81. 

1855. Lichenes, Collemaceae. In F. Junghuhn, Plantae 

Junghuhnianae sive Enumeratio Plantarum quas in 

insulis Java et Sumatra, pages 427-494. 

1880. Lichenologische Beitrage, X. Flora, 63:40-45. 

1882. Lichenologische Beitrage, XI’. Flora, 65:381-386. 

1887. Graphideae Feeanae. Memoires de la Societede 

Physique et d’Histoire Naturelle de Geneve, 29:3-80. 

1888a. Lichenologische Beitrage, XXX. Flora, 71:528-552. 

1888b. Revision Lichenum Eschweilerianorum. Flora, 71: 

1888c. Lichenes Paraguayenses a cl. Balansa lecti. Revue 

1888d. Lichenes Paraguayenses. Revue Mycologique, 10: 

1888e. Lichenes Paraguayenses. Revue Mycologique, 10: 

1891. Lichenes. In T. Durand and H. Pittier, Lichens 

Primitiae Florae Costaricensis. Bulletin de la SocittC 

Royale Botanique Belgique, 30:49-97. 

1955. 

Massalongo, A. 

Montagne, C. 

1853. 

1842. 

1851. 

Montagne, C., and R. B. van den Bosch 

Mueller Argoviensis, J. 

507-513. 

Mycologique, 38: 1-32. 

115-120. 

177-184. 

1892a. Lichenes Exotici. Hedwigia, 31 :276-284. 

1892b. Lichens, Seconde Enumeration. In T. Durand and 

H. Pittier, Primitiae Florae Costaricensis, 111. Bulletin de la Societe Royale Botanique Belgique, 32: 

1895a. Thelotremeae et Graphideae Novae Quae Praesertim ex Herb. Reg. Kewensi Exponit. Journal of the 

Linnean Society of London, Botany, 30:451-463. 

1895b. Graphideae Eckfeldtianae in Louisiana et Florida 

lecte. Bulletin de l’Herbier Boissier, 3:41-50. 

1895~. Lichenes Exotici, IV. Hedwigia, 34:139-145. 

122-1 73. 
NUMBER 40 51 

- 1827. 

Stirton, 

1881. 

Taylor, 

1847. 

Nakanishi, M. 

1966. Taxonomical Studies on the Family Graphidaceae 

of Japan. Journal of Science of the Hiroshima University, series B (Botany), 11:51-126. 

Nylander, W. 

1863. Lichenes. In J. Triana and J. E. Planchon, Prodromus Florae h’ovo-Granatensis ou Enumeration 

des Plantes de la Nouvelle-Grenade. Annales Sciences Naturelles Botanique, series 4, 20:228-382. 

1866. Collectio Lichenum ex Insula Cuba. Flora, 49:289- 

295. 

1868. Synopsis Lichenum Sovae-Caledoniae. Bulletin de 

la Societe Linne‘enne de Normandie, (2)2:39-140. 

1889. Lichenes Insularurn Guineensium. 54 pages. Paris. 

1890. Lichenes Japoniae. 122 pages. Paris. 

1891. Sertum Lichenaeae Tropicae e Labuan et Singapore. 

45 pages. Paris. 

Die Graphidineen der ersten Regnell’schen Expedition nach Brasilien, 1892-94, 11: Graphina und 

Phaeographina. Arkiv for Botanik, 26A(1): 1-105. 

1935. Die Graphidineen der ersten Regnell’schen Expedition nach Brasilien, 1892-94, 111: Graphis und 

Phaeographis, nebst einem Nachtrage zu Graphina. 

Arkiv for Botanik, 27A(2):1-103. 

Report on the Lichens of St. Thomas and St. Jan. 

In N. L. Britton, The Flora of the American Virgin 

Islands. Brooklyn Botanic Garden Memoirs, 1: 109- 

116. 

Redinger, K. 

1933. 

Riddle, L. W. 

1918. 

Smith, J. E., and J. Sowerby 

Sprengel, K. 

1807. English Botany. \.olume 26, plate 1812. 

Systema Vegetabilium. Volume 4, edition 16, part 2, 

410 pages. Gottingae. 

plants. Proceedings of the American Academy of 

Arts and Science, 7: 143-235. 

Synopsis of North American Lichens, Part 11. 176 

pages. New Bedford. 

Etude sur la classification naturelle et la morphologie des Lichens de Bred. Acta Societatis pro 

Fauna et Flora Fennica, 7(7): 1-256. 

1896. Lichenes Antillarum a W. R. Elliott collecti. Journal 

of Botany British and Foreign, 34:204-266. 

1899. Lichens novi rarioresque, 111. Hedwigia, 38:(253)- 

(259). 

1907. Lichenes novi rarioresque, IV. Hedwigia, 46: 168-181. 

1909. Lichenes. In J. Schmidt, Flora of Koh Chang, Part 

IX. Botanisk Tidsskrift, 29:104-151. 

1915. Additamenta ad lichenographia antillarum illustrandam. Annales Academiae Scientiarurn Fennicae, 

1921. Lichenes insularum Philippinarum, 111. Annales 

Academiae Scientiarum Fennicae, 15A: 1-368. 

1923. Lichenes in insula Trinidad a Professor R. Thaxter 

collecti. Proceedings of the American Academy of 

Arts and Sciences, 58:131-147. 

The Lichen Family Graphidaceae in Mexico. Contributions from the United States National Herbarium, 36:63-119. 

1888. 

Vainio, E. A. 

1890. 

6A( 7) : 1-226. 

Wirth, M., and M. E. Hale, Jr. 

1963. 

Zahlbruckner, A. 

1902. Studien uber brasilianische Flechten. Sitzungsberichte der kaiserlichen Akademie der Wissenschaft, 

Mathematisch-Naturwissenschaftliche Klasse, 111: 

357-432. 

Lichenes (Flechten). In R. Wettstein and V. Schiffner, Ergebnisse der botanischen Expedition der 

Akademie der Wissenschaften nach Sudbrasilien 

1901. Denkschriften der Akademie der Wissenschaften in Wien. Mathematisch-naturwissenschaftliche 

1909. 

J. Klasse, 83:87-211. 

On the Vegetable Parasites on the Tea Plant, More 1921. Neue Flechten IX. Annales Mycologici, 19:224-242. 

Especially That of Assam. Proceedings of the Philo- 1923. Catalogus lichenum universalis. \‘olume 2, 815 pages. 

sophical Society of Glasgow, 13:181-193. Leipzig: Gebruder Borntraeger. 

T. 1928. Lichenes. In C. Keissler, Schedae ad “Kryptogamas 

New Lichens, Principally from the Herbarium of exsiccatas.” Annalen des Naturhistorischen Museums 

W. J. Hooker. Hooker Journal of Botany, 6:148-197. 

1853. Spighe e Paglie. 65 pages. Padova. 

in Wien, 42:47-74. 

pages. Leipzig: Gebruder Borntraeger. 

1932. Catalogus lichenum universalis. Volume 8, 612 Trevisan, S.-L. 

Zenker, J. K. 

Tuckerman, E. 1829. Lichenes. In F. Goebel and G. Kunze, Pharma1866. Lichenes. In H. Mann, Enumeration of Hawaiian ceutische Waarenkunde, 1:109-199. 
Index 

(Synonyms in italics) 

Arthonia caesiopruinosa, 46 

Diorygma insculpta, 38 

Fissurina incrustans, 38 

insidiosa, 16 

nitidescens, 38 

virginalis, 36 

acuminata, 32 

adscribens, 31 

aibonitensis, 34 

albostriata, 18 

analoga, 42 

antillarum, 31 

babingtonii, 38 

balbisii, 34 

balbisii var. monospora, 33 

bipartita, 33 

carneoviridis, 33 

chlorocarpa, 33 

cleitops, 44 

colliculosa, 34 

collospora, 36 

collosporella, 44 

columbina, 36 

confluens, 41 

cymbergrapha, 48 

dealbata, 18 

dehiscens, 38 

deserpens, 33 

dimidiata, 36 

dimorphodes, 18 

dispersa, 37 

disserpens, 41 

elongata, 33 

elongatoradians, 41 

erythrella, 34, 43 

frumentaria, 37 

glaucoderma, 38 

heteroplacoides, 18 

illinata, 37 

incrustans, 38 

insculpta, 38 

insidiosa, 16 

insignis, 43 

intortula, 34 

intricata, 40 

iuturgescens, 31 

macella, 38 

Graphina acharii, 31 

marcescens, 40 

melaleuca, 44 

monophora, 45 

nitidescens, 38 

nuda, 40 

obtectula, 44 

palmeri, 44 

parilis, 33 

platycarpa, 41 

platycarpinu, 41 

platycarpoides, 32 

platyleuca, 41 

plittii, 40 

plurispora, 41 

pseudoanaloga, 42 

pseudosophistica var. plurispora, 41 

riopiedrensis, 34 

rufopallida, 42 

ruiriana, 41 

suberythrella, 42 

subnitidula, 22 

substriatula, 41 

subvelata, 42 

sulcata, 32 

sulcatula, 41 

triangularis, 44 

triphora, 44 

triphoroides, 44 

vermicularis, 42 

vestitoides, 44 

virginalis, 36 

virginea, 44 

Graphis acharii, 31 

acuminata, 32 

adpressa, 10 

adscribens, 31 

afzelii, 10 

albida, 25 

anfractuosa, 10 

anguilliformis, 10 

angustata, 15 

angustata var. denudata, 15 

antillarum, 31 

antillarum var. ingarum, 15 

arthonioides, 27 

beaumontii, 16 

bonplandiae, 24, 49 

caesiella, 19 

caesioglauca, 15 

calcea, 15 

candidissima, 40 

chlorocarpa, 33 

cinerea, 15 

colliculosa, 34 

collospora, 36 

collosporella, 44 

coiumbina, 36 

congesra, 15, 20 

cooperta, 12, 49 

dehiscens, 38 

desquamescens, 12 

dimidiata, 36 

ditersa, 49 

dumastii, 24 

dumastioides, 12 

duplicata, 21, 49 

duplicata var. subduplicata, 49 

dussii, 13 

elegans, 13 

elliottii, 48 

endoxantha, 21 

eugeniae, 34 

flavicarrs, 19 

flexibilis, 13 

frunientaria, 31 

furcata, 19 

glaucescens, 15 

glaucoderma, 38 

glauconigra, 20 

grammitis, 15 

haematites, 27 

humilis, 16, 24 

illinata, 37 

imshaugii, 16 

inquinata, 24 

insculpta, 38 

inszdiosa, 16 

interuersa, 16 

intortula, 34 

intricata, 12, 40 

tnturgescens, 31 

isidiifera, 18 

lactea, 24 

lactea var. clausa, 16, 49 

lactea var. dominicana, 23 

leptocarpa, 18 

52 
NUMBER 40 53 

librata, 19 

lineola, 19,49 

lobata, 49 

longula, 19 

lumbricina, 20 

marcescens, 40 

marginifera, 25 

medusaeformis, 49 

nitidescens, 38 

olivacea, 20 

oscitans, 48 

pavoniana, 19 

platy carpoides, 32 

platyleuca, 41 

proserpens, 22 

pseudoanaloga, 42 

rigidula, 21 

rimulosa, 21 

rosea, 28 

rubiginosa, 38 

rufopallida, 42 

seminuda, 12 

striatula, 21 

subamylacea, 16 

subelegans, 21 

subnitida, 49 

subnitidula, 22,24 

subtigrina, 29 

tachygrapha, 23 

tenella, 19 

tenella var. epiphaea, 49 

timida, 23 

tongloensis, 25 

triticea, 23 

tumidula, 12 

tumidulella, 15 

turgidula, 24 

vermicularis, 42, 49 

virginalis, 36 

Lecanactis exaltata, 27 

Leiogramma virgineum, 44 

Opegrapha desquamescens, 12 

elegans, 13 

rhlizocola, 12 

rimulosa, 21 

atrovermicularis, 46 

caesiopruinosa, 46 

caracasana, 48 

chrysocarpa, 28 

Phaeographina arechavaletae, 48 

columbina, 36 

coriaria, 46 

difformis, 48 

elliottii, 48 

elliptica, 48 

explicans, 48 

includens, 18 

intercedens, 46 

oscitans, 48 

pachnodes, 49 

turgida, 49 

ceruiculata, 44 

cinnabarina, 28 

decipiens, 27 

dimorpha, 29 

dussii, 13 

exaltata, 27 

haematites, 27 

longula, 19 

mordenii, 28 

rosea, 28 

sexloculata, 27 

subtigrina, 29 

Phaeographis albida, 25 

Sclerophyton colliculosum, 34 
54 ShlITHSONIAN COSTRIBUTIOSS TO BOTAhY 

PLATE 1.-Species of Graphis: a, G. adpresstc (Hale 35487); b, G. afieiii (Irnshaug 33489'4); c, 

G. nnfractuosa (Wirth 538); d, G. anguillifornzis (Hale 35282); e, G. desqztamescens (Hale 

33449); f, G. dumastioidrs (Hale 37722). (Plates 1-10 x 10.) 
NUMBER 40 

PLATE 2.-Species of Graphis: a, G. dussii (Hale 35443); b, G. elegans (Hale 35476): c, G. pexibilis (Zmshaug 33360); d, G. glaucescens (Zmshaug 33313A); e, G. grammitis (Hale 38005); f, G. 

humilis (Hale 35698). 

55 
56 SMITHSOSIAN CONTRIBUTIOSS TO BOTANY 

PLATE 3.-Species of Graphis: a, G. irnshaugii (Imshntig 32724 in MSC): b, G. insidiosa (Hale 

35313); c, G. isidiifera (Hale 35179); d, G. leptocarpn (Imshatig 33109.A); e, G. librata (Wirth 

538); f. G. Eongula (Hale 35225). 
NUMBER 40 

PLATE 4.-Species of Graphis: a, G. lumbricina (Hale 35240); b, G. olivacea (Hale 37988); c, 

G. rigidula (Imshaug 32777A); d, G. rimulosa (Hale 35699); e, G. subelegans (Hale 35546); 

j, G. subnitidula (Hale 37676). 

57 
58 SMITHSONIAS COSTRIBUTIONS TO BOTANY 

PLATE 5.--Species of Graphis and Phaeographis: a, G. tachygmpha (Imshaug 32862A); b, G. 

triticea (Hale 35324); c, G. turgidula (Wirth 544b); d, P. aibida (Hale 35453); e, P. arthonioides 

(Imshaug 33128); f, P. exaltata (Hale 35301). 
NUMBER 40 

PLATE B.-Species of Phaeographis and Graphina: a, P. exaltata (Hale 35321); b, P. haematites 

(Hale 38013); c, P. mordenii (Hale 35071); d, P. rosea (Elliott in BM); e, P. subtigrina (Wirth 

444); j, G. acharii (Hale 35555). 

59 
60 SMITHSOSIAS COSTRIBUTIONS TO BOTASY 

PLATE 7.--Species of Graphina: a, G. adscribens (Wirth 519); b, G. antillaruni (Hale 35745); 

c, G. carneoviridis (Hale 37642); d, G. chlorocarpa (Hale 35129); e, G. colliculosa (Hale 35175); 

f, G. collospora (Wirth 520). 
NUMBER 40 

PLATE &-Species of Graphina: a, G. columbina (Wirth 446); b, G. dimidiata (Hale 35697); 

c, G. dispersa (Hale 38090); d, G. frumentaria (Hale 35242); e, G. illinata (Hale 35284); 

f, G. incrustans (Wirth 465). 

61 
62 SMITHSOXIAN COSTRIBUTIONS TO BOTASY 

PLATE 9.--Species of Graphina: a, G. insculpta (Hale 37819); b, G. macella (Hale 35723); c, 

G. marcexens (Hale 35312); d, G. nuda (Hale 35119); e, G. platyleuca (Hale 37955); j, G. 

plurispora (Imshaug 37724.4). 
NUhlBER 40 

PLATE 10.-Species of Graphina: a, G. pseudoanaloga (Hale 35283): b, G. rufopallida (Elliott 

1832); c, G. suberythrella (Zmshaug 32699); d, G. triphoroides (Hale 35161); e, G. vestitoides 

(Zmshaug 32859A); f, G. virginea (Wirth 547). 

63 
64 SMITHSONIAN CONTRIBUTIOSS TO BOTANY 

PLATE 1 1.-Species of Phaeographina: a, P. atrouermictilaris (Hale 35278); b, P. raesiopruinosa 

(Znzshaug 32713); c, P. coriaria (Hale 35378); d, P. dioormis (Zrnshaug 32913.4); e, P. elliottii 

(EIEiott 1338 in TUR); f, P. oscitans (Hale 35131).