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to Dominica:
The Lichens (Graphidaceae
MICHAEL WIRTH and
MASON E. HALE, JR.
SMITHSONIAN CONTRIBUTIONS TO'BOTANY • NUMBER 40
I
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S. Dillon Ripley Secretary
Smithsonian Institution
SMITHSONIAN CONTRIBUTIONS TO BOTANY • NUMBER 40
Morden-Smithsonian Expedition to Dominica:
The Lichens (Graphidaceae)
Michael Wirth and Mason E. Hale, Jr.
ISSUED
OCT
SMITHSONIAN PUBLICATIONS
SMITHSONIAN INSTITUTION PRESS City of Washington 1978
ABSTRACT
Wirth, M., and Mason E. Hale, Jr. Morden-Smithsonian Expedition to Dominica: The Lichens (Graphidaceae). Smithsonian Contributions to Botany, number 40, 64 pages, 23 figures, 1978. — The four main genera of the Graphidaceae in Dom.inica, Graphis (27 species), Phaeographis (7 species), Graphina (25 species), and Phaeographina (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 described; Graphis imshaiigii, G. isidiifera, Phaeographis rnordenii, Graphina carneoviridis, G. suberythrella, G. triphoroides, Phaeographina atroverrnicularis, and P. coriaria. Three new combinations, Graphina colurnbina (Tuckerman) Wirth and Hale, G. illinata (Eschweiler) Wirth and Hale, and G. plurispora (Redinger) Wirth and Hale, are also made.
i T AD;
Official fublication date is handstamped in a limited number of initial copies and is recorded in the Institution’s annual report, Smithsonian Year. Serifs cover design: Leaf clearing from the Katsura tree Cercidiphyllum japonicum Siebold and Zuccarini.
Library of Congress Cataloging in Publication Data Wirth, Michael.
Morden-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
I. Graphidaceae. 2. Lichen — Dominica. 3. Morden-Smithsonian Expedition to Dominica 1. Hale, Mason E. joint author. 11. Title. III. Series: Smithsonian Institution. Smithsonian contributions to botany ; no. 40)
QKLS2747 no. 40 [QK585.GS] 581'.08s [389'.I] 77-608313
Contents
Page
Introduction 1
Generic Delimitation 1
Species Delimitation 3
Spores 3
Hymenium 3
Exciple 3
Thallus 5
Chemistry 5
Ecology 6
Evolution in the Graphidaceae 6
Key to the Genera of the Graphidaceae in Dominica 8
Key to the Species of Gr aphis 8
Key to the Species of Phaeographis 25
Key to the Species of Graphina 29
Key to the Species of Phaeographina 45
Incorrect and Omitted Names 49
Literature Cited 50
Index 52
Plates 54
iii
Morden-Smithsonian Expedition to Dominica:
The Lichens (Graphidaceae)
Michael Wirth and Mason E. Hale^ Jr.
Introduction
This study is a continuation of a floristic treat- ment 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 geogra- phy and ecology of the island. The specimens ex- amined 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 eleva- tion 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 Mrs. 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 Hamp- shire 03242. Mason E. Hale, Jr., Department of Botany, National Museum of Natural History, Smithsonian Institu- tion, 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 fre- quently appear free and unbranched and the two- loculed 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, Phaeogr aphis, Graphina, Phaeo- graphina), four parallel genera with such tissue (Glyphis, Sarcographa, Medusulina, Sarcographina), and a number of minor genera whose relationships are unclear (Helminthocarpon, Aulaxina, Gyro- stomiim, 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
SMITHSONIAN CONTRIBUTIONS TO BOTANY
usually applied to the family, using the terms “hy- menium” and “paraphyses” without necessarily accepting them as ontogenetically correct.
To a great extent, all genera (in phanerogams as well as in lichens) are artificial constructs, in that no fixed criteria exist for delimiting any taxa above the biological species. In groups where cytogenetics, breeding experiments, etc., can reveal something of the direction of evolution, genera may in fact be an approximation of biological (i.e. evolutionary) re- latedness. Obviously, in the crustose lichens, such information is mostly lacking. In addition, onto- genetic studies of lichen reproductive structures are scarce and frequently open to dilferent interpre- tations; for example: What exactly is a hymenium? 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 Graphi- daceae are reasonably delimited. By using spore 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 s|recies) exist that bridge the gap between these form genera and tend to blur the distinctions. In Graphina vestitoides, for example, only the end locules of otherwise typi- cal Graphis spores become longitudinally divided. Many of the very small-spored Graphina species (G. insignis, G. coUicidosa, etc.) have barely muri- form spores which, when not quite mature, will resemble four-loculed Graphis spores. The prob- lems of spore color and septation in such border- line species as Graphina acharii and Phaeographina chrysocarpa have been mentioned elsewhere (Wirth and Hale, 1963).
Intermediate states also exist between the strom- atoid and nonstromatoid genera. A fine example can be seen in many specimens of Phaeographis exallata, which frequently show, in portions of a single thallus, characteristics of both Phaeographis and Sarcographa.
There is, however, a more serious objection to sjiore genera. One can regard a genus as consisting of species that share a large number of character- states in common; this approach, common among
“numerical” taxonomists, assumes that “related- ness” 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 c^uite useful. One can find many examples of species that are extremely similar but are pres- ently placed in different genera only because of a difference in spore color and/or septation. We will call such forms “sporomorphs,” defined as follows: two or more species that are extremely similar (or identical) in external morphology, anatomy, chem- istry, 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 septation and/or spore color only.
Sporomorphs are common in the Graphidaceae; good examples occurring at least in part in Domi- nica are Phaeographis arthonioides j Graphis hiimilis and Graphina incrustans j Graphis dumastii.
The existence of sporomorphs 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 muri- form condition simply represents a later ontogenetic stage of the Graphis spore that has become genet- ically fixed? If such were the case (and indeed all muriform spores seem to go through a Graphis-\ik.e juvenile stage), then Graphina incrustans may be a recent descendant of Graphis dumastii, and these two species should be regarded as closely related. Placing these species in different genera, while grouping G. incrustans with otherwise very dis- similar Graphina 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 color and septa- tion 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 over- all 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 w'ould be to reduce all the spore-based genera back into Graphis. Such a system, which may be biologically quite acceptable, would raise immense practical problems; there are simply too many species of Graphidaceae (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, re- gardless of present generic lines.
Species Delimitation
With essentially no knowledge of speciation in the Graphidaceae, we have had to use the tradi- tional approach of concentrating on discontinuities of morphology, anatomy, chemistry, and size. This approach works best with large collections of speci- mens; 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 sjrecies. In a fam- ily with such a large range of spore size (6 .^um to 300 it is not surprising to find discontinuities of size classes. We have, whenever possible, main- tained species on this basis. Within a given indi- vidual, 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 num- ber 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 Phaeographis) where the main, and perhaps only, distinction consists of four versus more than four locules per spore.
Hymenium. — 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 car- bonized exciples the hymenium height becomes pro- portionally small; for most other species, it is an excellent indicator of over-all size. Hymenial meas- urements 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 Redin- ger, 1933:4 and 1935:2). 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.
Exciple Carbonization: Garbonization 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 in- complete 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 con- nected by a more or less continuous black to brown (partially carbonized) band. Such a condition ap- pears commonly in Graphina insignis, and less fre-
4
SMITHSONIAN CONTRIBUTIONS TO BOTANY
quently in G. virginea 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 apothe- cia in surface view, and four-locular spores), the term “exciple” may be inapplicable, as the hymen- ium 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 tliat presence of large amounts of bark laterally bounding the hymenium is signifi- cant as an indicator of endophloeodal development of the ascocarp.
Open vs. 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 un- justified. When no carbonization exists, it becomes even more difficult to determine if closure is pres- ent, 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. flexi- bilis).
It is possible that the striations in both carbo- nized and clear exciples may be a function of age. Specimens that have high ascocarps with thick bases and (frequently) disintegrating hymenia (all pre- sumed 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 fre- quently 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. Graphina peplophora (Figure 1). If this inter- pretation 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 denta- tions or striae occur within the labial tissue, with- out any trace of surface grooves. Examination of many ascocarps on large thalli reveals that these striations arise with age, apparently by lateral por- tions of the hymenium compacting, darkening, and being incorporated into the lateral tissue surround- ing 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 8/,g; 9rt) illus- trate 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 carbo- nized 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 8/,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 appear- ance will differ strikingly and result in the descrip- tion of two species.
Thallus. — Very few character-states of impor- tance are derived from the thallus. Color, in gen- eral, is not a practical feature, with the possible exception of those endophloeodal thalli which fre- quently signal this condition by a distinctive yellow- brown shade.
The thalli of some species show a distinct proso- plectenchymatous 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 isi- dia. 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 spe- cies 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, Phaeo- graphis 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 iden- tical 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 sec- ondary products. The most common lichen sub- stance in the TLC positive remainder is unques- tionably norstictic acid. Fairly abundant, though not common, are stictic acid and the “quintaria” unknowns. Very uncommon are psoromic, lecanoric.
6
SMITHSONIAN CONTRIBUTIONS TO BOTANY
protocetraric, salazinic acids, and lichexanthone. A few species {Phaeogr aphis haematites, P. cinnaba- rina (Fee) Mueller Argoviensis, Phaeographina chrysocarpa (Raddi) Redinger, among others) have pigments (anthraquinones?).
All specimens were tested with normal thin-layer chromatographic techniques using silica gel alum- inum-backed Merck precoated plates. Two solvent systems were always used: hexane-ether-formic acid and benzene-dioxane-acetic acid. Visualization was accomplished with 10% sulfuric acid and heating at 110° C.
A list of the chemicals found and species that contain them is given below. 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: Graphis afzelii
Norstictic acid: Graphis desquainescens, G. dussii, G. elegans, G. isidiifera, G. librata, G. lumbricma, Phaegraphis sub- tigrina, Graphiua anliltarum, G. dispersa, G. plalyleuca (accessory), G. psetidoatialoga, G. suberythrella, and Phaeo- graphina at rove rinicula ris
Norstictic acid with stictic and constictic acids: Graphina cotlospora
Protocetraric acid: Graphina platyleuca Psoromic acid: Graphina coliimbina
“Quintaria” unknowns: Phaeographis exaltata and Graphina virginea
Salazinic acid: Graphina colliculosa and G. marcescens; G. platyleuca (accessory)
Slictic acid: Graphis dumastioides, G. i)nsha2tgii, G. lepto- carpa, G. triticea, G. turgidula, Graphina triphora, G. virginea, and Phaeographina oscitans TLC negative (no phenolic substances present): Graphis adpressa, G. anfractuosa, G. anguilliforniis, G. flexibilis, G. glaucescens, G. grammitis, G. humilis, G. insidiosa, G. longula, G. olivacea, G. rigidula, G. rimulosa, G. subele- gans, G. subnitidula, G. tachygrapha, Phaeographis albida, P. arthonioides, P. haematites, P. mordenii, P. rosea, Graphina acharii, G. adscribens, G. carneoviridis, G. chlo- rocarpa, G. dimidiata, G. frumentaria, G. illinata, G. in- crustans, G. insculpta, G. macella, G. mida, G. plurispora, G. rufopallida, G. vestitoides, Phaeographina caesioprui- nosa, P. coriaria, P. difformis, P. elliottii, and (in part) P. exaltata
Pigments: Phaeographis haematites
Ecology
The family Graph idaceae 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 afzelii, seem to have a wide ecological am- plitude in both secondary and primary forests. This is in contrast to the closely related Thelotrema- taceae that are highly restricted to undisturbed rain forest in Dominica (Hale, 1974) and other tropical areas.
We are not able to categorize all the species eco- logically even for such a small island as Dominica since many are represented by only one or two col- lections. However, the areas of mid elevation virgin rain forest 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 olivacea (also rarely found in elfin forest), Graphina chlorocarpa, G. colliculosa, and G. illinata. These species are com- mon in and probably restricted to elfin or mossy forest: Crraphis adpressa, G. anguilliformis, G. lumbricina, G. tachygrapha, G. triticea, Phaeo- graphis exaltata, and P. mordenii. These occur chiefly in low elevation, disturbed habitats: Graphis desquamescens, Graphina antillarum, and G. vir- ginea. Further notes are given under “Habitat” for each species but these should be considered tenta- tive until more collections are made.
Evolution in the Graphidaceae
In order to speculate on the evolution of non- stromatoid graphids, several assumptions must be accepted. First, one must assume that the group is relatively natural, i.e., monophyletic. Second, since data from genetics and cytology are still lacking, one must assume that at least some phylogeny can be inferred from ontogeny.
Although the pitfalls and fallacies of using on- togeny as a source for evolutionary speculation are well known, there seems little choice in the Graph- idaceae. Specifically, stages of spore development may supply the only concrete bases yet available.
The following four observations on spore ontog- eny should be considered:
1. All Graphic! spores start as a one-loculed (one- celled) structure, progress to two locules, then to four (rarely three) uniseriate chambers, then to multilocular stages.
2. Longitudinal spore septae occur only after most transverse septae have been laid down.
NUMBER 40
7
3. Monosporous asci always (?) begin as pluri- sporous asci, followed by abortion of most spores in their early stages. Monosporous asci occur only (?) in Graphma 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 spe- cies 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 fol- lowing 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 neo-
tenous derivatives.
2. Muriform-spored species are derived from an- cestors with Grap/aVlike 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 Graphma sporomorphs by added pigmentation, rather than from Phaeographis sporomorphs by added septation. The less common thick-walled Phaeographina species may be derived from irregu- larly-loculed Graphina ancestors (also rather un- common) or from Phaeographis-\i\:.t forms.
Putting these speculations together in graphic form, we propose a very hypothetical scheme of evolutionary relationships (Figure 2).
Ancestral stock; spores 2- locular, c lear ( few extant
species G. corlacea G. turbulenta. etc.)
Figure 2. — Hypothetical origin and relationships of the genera of the Graphidaceae.
8
SMITHSONIAN CONTRIBUTIONS TO BOTANY
This scheme immediately gives rise to certain questions, problems, and objections, to wit: Where does carbonization fit in this scheme? Graphis spe- cies with £ew-loculed spores, supposedly primitive in the proposed arrangement, very rarely show any carbonization. At the same time, lack of carboniza- tion is also the more common condition in the sup- posedly most advanced groups (i.e., those with monosporous asci and muriform spores). In addi- tion, many of the fissurine Graphis species, sup- posedly 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 sys- tem 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
spore states? For example: a fissurine Graphis 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 mono- sporous 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 cor- relations 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 competi- tive advantage associated with the variants. In the Byrsonima scrub of western Mexico, for example, all the spore forms occur abundantly, shoulder to shoulder. However, this may be a very equable habitat, comparable to the tropical rain forest for the angiosperms; other more demanding habitats and substrates might reveal significant preferences and correlations. In addition, the abundance of ad- jacent forms may be misleading in that a succession may be occurring too slowly to be apparent imme- diately.
The same lack of data exists with respect to dis- tribution of carbonized vs. uncarbonized forms, endo- vs. 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 below, the two stromatoid groups, Glyphis and Sarcographa, are not treated in the text. Each is represented by only one species (Y’ainio, 1896, 1915.)
1. Ascocarps embedded in prominently raised, stromatoid tissue.
2. Spores clear, with transverse septae only Glyphis
2. Spores brown, with transverse septae only Sarcographa
1. Ascocarps not embedded in raised stromatoid tissue.
3. Spores clear and hyaline, the walls usually thin.
4. Spores with transverse septae only Graphis
4. Spores with transverse and longitudinal septae Graphina
3. Spores brown, the walls occasionallv thickened.
5. Spores with transverse and longitudinal septae, the walls rarely thickened
Phaeographis
5. Spores with transverse and longitudinal septate, the walls rarely thickened
Phaeograph ina
Key to the Species of Graphis
1. Spores always four locular.
2. .Ascocarps fissurine, arising as a thalline crack; disc usually concealed or sunken.
3. Exciple carbonized.
4. Exciple heavily carbonized laterally, spores 1 +
12. G. humilis
4. Exciple carbonized only at labial tips, spores I— (reddish)
25. G. tachygrapha
3. Exciple totally uncarbonized.
5. Ascocarps usually exposing a red-brown disc in portions, commonly to 10 mm
long 11. G. grammitis
5. Ascocarps fissurine throughout, rarely more than 3 mm long, usually shorter.
6. Ascocarps embedded in swollen tissue (‘‘puffs'’).
7. Stictic acid present 26. G. triticea
7. No substances present 14. G. insidiosa
6. Ascocarps not embedded in swollen tissue.
8. Spores less than 11 /j,m long; no substances present 24. G. subnitidula
8. Spores more than 13 /^m long; stictic acid present 6. G. dumastioides
2. Ascocarps not fissurine; disc broad, obvious, not depressed within a thalline crack.
9. Ascocarps white-powdery; exciple carbonized; lecanoric acid present 2. G. ajzelii
9. Ascocarps not powdery; exciple carbonized; lecanoric acid absent.
10. Thallus isidiate; thalline margin coarsely white-mealy; asocarps nearly round;
norstictic acid present 15. G. isidiifera
10. Thallus not isidiate; thalline margin not mealy; ascocarps elongate; no substances
present 11. G. grammitis
Spores six locular or more.
11. Exciples red-brown to barely carbonized at labial apices 10. G. glaucescens.
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 yum long 20. G. olivacea
14. Thalline margin absent or nearly so; spores under 35 ^m ....23. G. subelegans 13. Exciples lacking red-yellow tissue below the carbonized portions (but some- times dirty-brown below).
15. Ascocarps usually without a prominent thalline margin.
16. Exciples open below; norstictic acid present 8. G. elegans
16. Exciples closed below; no substances present 22. G. rimtdosa
15. Ascocarps consistently with a thalline margin.
17. Norstictic acid present 19. G. lumbricina
17. Norstictic acid absent.
18. Exciple open below.
19. Spores usually 70-90 j^m long 18. G. longula
19. Spores usually 50-65 jj.m long 21. G. rigidula
18. Exciple closed below.
20. Exciple massively carbonized below 9. G. flexibilis
20. Exciple barely closed below 18. G. longula
12. Labia intact or irregular, not distinctly striate.
21. Exciples distinctly open below.
22. Stictic acid only present 16. G. leptocarpa
22. Norstictic acid only present.
23. Spores less than 36 fixa long 17. G. librata
23. Spores over 45 jum long 7. G. dussii
21. Exciples distinctly closed below, or the lateral portions of the exciples ap- 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 fim long.
26. Stictic or norstictic acid present.
27. Norstictic acid present 5. G. desquamescens
27. Stictic acid present 13. G. imshaugii
26. No substances present 3. G. anfractuosa
25. Spores usually over 60yum long.
28. Stictic acid present 27. G. turgidula
28. No substances present 4. G. anguillijormis
10
SMITHSONIAN CONTRIBUTIONS TO BOTANY
1. Gr aphis adpressa Figure 3a; Plate la
Graphis adpressa Vainio, 1890:119 [type collection: Carassa,
Brazil, Vainio 1289 (TUR, lectotype)].
Description. — Thallus white, gray, or pale brown, thin but continuous, shining. Ascocarps quite black, raised and protuberant, usually un- branched, no thalline margin visible, 0.5-2 mm long, slender. Disc not visible in surface view. Asco- carp transverse section: hymenium 170-250 p,m high, I-; exciple black, closed, lips intact and con- vergent, frequently with an extremely thin thalline covering. Spores 8/ascus, 10-18 locular, 13-15 X 50-70 p.m, I-f blue.
Chemistry. — No substances present.
Habitat. — Rain forest above 2600 feet.
Discussion. — Graphis adpressa externally resem- bles an O pegrapha or Melaspilea and is unlikely to be confused with any other Dominican Graphis. It exhibits considerably less variation in morphology than most Graphids. Extremely similar but for spore septation and size is Graphina niida.
Specimens Examined. — Fresh Water Lake, 2600-2800 ft. Hale 35485, 35487 (US), 2600-2877 ft, Imshaug 32849A, 32852A, 32854 (MSC); Trois Pitons, 2900-3150 ft, Imshaug 33090 (MSC); Morne Diablotin, 3500-4300 ft, Imshaug 32940 (MSC); Micotrin, ca 3000 ft, Wirth 464 (US).
2. Graphis afzelii Figure 35; Plate lb
Graphis afzelii .\charius, 1814:85 [type collection: Guinea,
Afzelius, s.n. (H, lectotype; UPS, isotype)].
Description. — Thallus brownish tan to gray, con- tinuous, smooth. Ascocarps usually completely covered by a jrowdery white thalline layer, appear- ing black only where this layer is rubbed, usually unbranched, 1. 0-6.0 mm long, to nearly 1 mm wide. Disc not visible in surface view. Ascocarp transverse section: hymenium 75-150 pm high, I-; exciple black laterally, brown and closed to pale and open basally, lips convergent, intact, with a prominent thalline covering extending to their apices. Spores 8/ascus, always 4 locular, 6-9 X 16-23 pm, I-P blue.
Chemistry. — Lecanoric acid (in the thalline apothecial covering only).
Habitat. — Dry sea-level scrub to virgin upland
rain forest; apparently with broad ecological ampli- tude.
Discussion. — Graphis afzelii is a common pan- tropic 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.
Specimens Examined. — Pointe Round, 100 ft, Imshaug 33484A (MSC); Londerry Agricultural Area, 100 ft, Imshaug 32897 (MSC); near Madjini, 100 ft, Hale 35618 (US); north of Bioche, 200 ft. Hale 35644 (US); Can-Dom logging area, Newfoundland, 800 ft. Hale 35239, 35446, 35447 (US); 8 miles north of Pont Casse, 1400 ft, Wirth 546 (US); Can-Dom logging area, Dleau Gommier, 1600-1700 ft. Hale 35183 (US).
3. Graphis anfractuosa Eigure 3c; Plate Ic
Graphis anfractuosa Eschw'eiler in Martins, 1833:86 [type
collection: Caitete, Brazil, s.n. (M, lectotype; G, isotype)].
Description. — Thallus dull white to gray, thin, continuous. Ascocarps partially emergent, the labial tips appearing as slender, flexuose, black, sparingly branched lines 1. 0-3.0 mm long. Disc not visible in surface view. Ascocarp transverse section: hymenium (80-) 90-130 pm high, I-; exciple black laterally, frequently dark brown to black below, usually closed, lips more or less intact, convergent. Spores 8/ascus, 8-12 locular, 9-10 X 40-50 pm, I-t- blue.
Chemistry.^ — No substances present.
Habitat. — Rain forest lower branches, ca. 1400 feet (good specimen); coastal scrub (poor specimen).
Discussion. — The lectotype specimen of Graphis anfractuosa is large but apparently without spores; the isotype at Geneva is quite small but fertile.
Very similar is Graphis desquarnescens, which dif- fers in having smaller spores and norstictic acid.
■Specimens Examined. — Scotts Ffead, sea level, Wirth 538 (US); Emerald Pool, ca 1400 ft, Wirth 504a (US).
4. Graphis anguilliformis Eigure 3d; Plate Id
Graphis angtiilliformis Taylor, 1847:152 [type collection:
St. \’incent, s.n. (FH, lectotype)].
Description. — Thallus tan to light brown, thick, glossy, frequently cracked. Ascocarps large, protu- berant, unbranched to sparingly branched, straight
NUMBER 40
11
Figure 3. — Cross sections of ajxithecia of Graphis: a, G. adpressa {Vainio 1289); b, G. afzelii (lectotype in H); c, G. anfractuosa (lectotype in G); d, G. anguilliformis (lectotype in FH); e, G. desquamescens {Glaziou 5082); /, G. dumastioides (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 transverse section; hymenium 190-250 p,m high (to 375 ju,m in a few extreme cases), I-; exciple black, massive, closed, lips entire, convergent, thalline
margin prominent. Spores (2-) 4-6 (-8?)/ascus, 12- 18 locular, 12-15 X (65-) 75-110 (-130) fcm, 1 + blue.
Chemistry. — No substances present.
Habitat. — Rain forest above 1500 feet, up into elfin forest.
Discussion. — Graphis anguilliformis (“anguillae-
12
SMITHSONIAN CONTRIBUTIONS TO BOTANY
formis” in the original description) is the commonest graphid in Dominica; we have also seen abundant material of this very large sjrecies 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.
Gr aphis cooper ta Zenker (1829) is supposedly a synonym of O. 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 (Fte) Sprengel (1827) is quite close; it differs in size (hymenium frequently 400 [xm high) and in the very large spores (40 locular, to 300 ^m 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.
Graphis turgidula Mueller Argoviensis differs in having stictic acid.
Graphis flexibilis differs only in being somewhat smaller, and in having striate labia.
Specimens Examined. — Can-Dom logging area, Dleaii Gom- mier, 1600-1700 ft, Hale 35154 (US); Can-Dom logging area, Pont Casse, 2000 ft. Hale 35084, 35112 (US); Can-Dom log- ging area, Brantridge Estate, 1700 ft. Hale 35282 (US); Morne Anglais, 300-3600 ft, Hale 35326a, 35349, 35351 (US), Elliott s.n. (TUR); Boiling Lake, 2400-3000 ft. Hale 35759, 35760 (US); Fresh Water Lake, 2600-2877 ft, Imshaug 32848 (MSC); Boeri Lake, 2750 ft, Irnshaug 33225 (MSC); Central Forest Reserve, 1500 ft, Imshaug 33554.\ (MSC); Morne Diahlotin, 3000-3500 ft, Irnshaug 32955 (MSC); 2200-2600 ft. Hale 38174, 38191 (US).
5. Graphis desquamescens Figure 3e; Plate \e
Graphis desquamescens (Fee) Zahlbruckner, 1909:108. Opegrapha desquamescens Fee, 1874:24 [type collection: Brazil,
Glaziou 5082 (M, lectotype)].
Description. — Thallus white to off-white, con- tinuous, slightly roughened and matte. Ascocarps flush to emergent, slender, flexuose, black, fre-
quently branched, usually with a low thalline mar- gin, (1-) 2-4 (-6) mm long. Disc barely visible in surface view. Ascocarp transverse section: hymenium 60-70 (-120) jim 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) y^m, 1 4-, 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 cor- rectly) from Japan by Nakanishi (1966). Very sim- ilar to Graphis desquamescens is G. intricata Fee (1824), separable only by the smaller spores (less than 15 /xm long) in the latter. Also similar is G. anfractuosa, which lacks norstictic acid and has somewhat larger spores.
Specimens Examined. — Rodney’s Rock, sea level. Hale 35581 (US); north of Mero, 50 ft. Hale 35067, 35449, 35727 (US); Grande Savanee, 200-250 ft, Imshaug 33285A, 33291A (MSC); Mt. St. Mary’s, 600-700 ft, Imshaug 33448A (MSC).
6. Graphis dumastioides Figure 3/; Plate If
Graphis dumastioides Fink, 1927:213 [type collection: Ma-
meyes, Puerto Rico, Fink 723 (MICH, lectotype; FH, US,
iso types)].
Description. — Thallus brown to greenish tan, glossy, continuous, rather thick for a fissurine spe- cies (to 150 /xm), with a distinct prosoplectenchyma- tous 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: hymenium 60- 100 /xin high, I-; exciple rudimentary at best, the lips consisting of barely modified uncarbonized thalline tissue with included bark cells. Spores 8/ascus, always 4 locular, (6-) 8-9 X (14—) 18-25 /xm, I- or I-f- slow, faint and reddish.
Chemistry. — Stictic acid.
Habitat. — Rain forest.
Discu-SSIon. — For a discussion of the fissurine complex of species, see Graphis triticea.
Specimen Examined. — Road to Morne Jean, 2000-2400 ft. Hale 37722 (US).
NUMBER 40
13
7. Graphis cf. dussii Figure 4a; Plate 2a
Graphis dussii Vainio, 1899:255 [type collection: Guadeloupe,
Duss 515 (TUR, lectotype)]
Phaeographis dussii (Vainio) Zahlbruckner, 1923:371.
Discription. — Thallus 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 unbranchecl and more or less straight, with a low thalline margin. Disc usually not visible in surface view. Ascocarp transverse section: hymenium 140-150 /xm high, I-; exciple black laterally, open below, lips convergent, entirely to barely crenate. Spores 6-8/ascus, (10-) 12-16 locular, 6-10 X 60- 100 (lUi (46 fxm long fide Vainio, but none so short found on the type), 1+ 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 lirellae 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 col- lections 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 com- plete discussion of this complex group, see under Graphis leplocarpa.
Specimen Examined. — Morne Diablotin, 3200-4600 ft. Hale 35443 (US).
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 p,m 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 p,m, I-f blue.
Chemistry.- — Norstictic acid (see discussion).
Habitat. — Secondary and mossy forests above 1800 feet.
Discussion. — Without having seen the type of Graphis elegans, the identification of the Domini- can material is somewhat uncertain, but our mate- rial matches the “classic” concept of the European species very closely. In addition, at least some of the specimens from northern Europe also contain nor- stictic 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 flexibilis Krempelhuber, 1876:414 [type collection:
Brazil, Glaziou 5106 (M, lectotype)].
Description. — Thallus gray to white, usually smooth and glossy, occasionally somewhat rough- ened and cracked, thick, cortex prosoplectenchyma- tous 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 pin high), exciple always quite closed below, lips convergent, with 1-4 striae. Spores 6-8/ascus, (15-) 20-30 locular, 13-16 X 60- 150 (-190) pm (60-110 pm in the type specimen), 1+ blue.
Chemistry. — Type, P-, no TLC made; Domini- can material, no substances present.
14
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Figure 4. — Cross sections of apothecia of Graphis: a, G. dussii (Diiss 515); b, G. elegans (lectotype in BM); c, G. flexibilis (Glazioii 5016); d, G. glaucescens (lectotype in G); /, G. imshaugii (Imshaug 32724B); g, G. humilis {Merrill 9067); li, G. humilis {Hale 35698).
NUMBER 40
15
Habitat. — Below the lower edges of the wet rain forest.
Discussion. — Externally Graphis flexibilis strongly resembles G. anguilliformis and Graphina acharii and can be considered a sporomorph of the latter. Many other species must be treated in any discus- sion of G. flexibilis, as follows.
Graphis angustata Eschweiler (1833) is sup- posedly an earlier name for G. flexibilis {fide Zahl- bruckner, 1923); however, the type (M!) has ap- parently always been found to be sterile (see Mueller Argoviensis 1888b:509). 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. an- gustata var. ingarum Vainio (1915), also unrelated, belongs in the Eugraphis alliance.
Graphis cinerea Eee (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 ju,m), with only 1-3 per ascus. Eor the moment, this species can thus be separated from G. flexibilis by its spores.
Graphis calcea (Eee) Massalongo (1853) can tenta- tively be separated by its lower ascocarps (nearly flush) and fewer striae.
Graphis congesta (Eee) 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 lumbricina differs in the presence of norstictic acid and in the occasionally open exciple.
Graphis olivacea is separable only by the paler red-yellow excipular base.
Graphis rigidula can tentatively be separated by its open exciple and smaller size.
Graphis tumidulella Fink (1927) is separable only by its smaller spores (under 50 ju,m).
Specimens Examined. — Bois Serpe, ca. 1000 ft, Imshaug 32772A (MSC): Ridgefield Estate, 1100-1200 ft, Imshatig .33360 (MSC); Shawford Estate, Elliott 1854 p.p. (TUR).
10. Graphis glaucescens Figure 4d; Plate 2d
Graphis glaucescens Fee 1824:36 [type collection: South
America, s.n. (G, lectotype)].
Description. — Thallus white, continuous, mi- nutely roughened. Ascocarps concolorous with the thallus, more or less black only where the thalline covering is rubbed, slightly raised, branched and flexuose, 1. 0-3.0 mm long, slender. Disc occasionally visible as a slightly darker central area. Ascocarp transverse section: hymenium 75-100 ju,m 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 Jim, 1+ blue.
Chemistry. — No substances present.
Habitat. — Lowland cultivated areas.
Discussion. — Graphis caesioglauca Redinger (1935) is indistinguishable, externally and intern- ally, from G. glaucescens. Transverse sections of the type {Malrne 1526, S) indicate the same labial struc- ture, and the Redinger illustrations are incorrect. We have not yet determined the chemistry of G. caesioglauca.
Specimens Examined. — Brookhill Estate, 100-150 ft, Im- shaug 33313A (MSC).
1 1. Graphis grammitis Plate 2e
Graphis grammitis Fee 1824:47 [type collection: South Amer- ica, s.n. (G, lectotype)].
Description. — Thallus thick, prosoplectenchyma- tous, 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 expo.sed in at least a portion of the ascocarps, red-brown. Asco- carp transverse section: hymenium 60 p.m high (35- 40 in the type), epihymenium usually darkened, I-; exciple barely closed, yellow-brown to red-brown, lips convergent to strongly spreading, apically some- times darkened. Spores 8/ascus, always 4 locular, 4-8 X 10-14 p.m, I-b blue.
Chemistry. — No substances present.
Habitat. — Elfin forest and upland rain forest (Dominica only).
Discussion. — In early stages, the ascocarps of Graphis grammitis are quite fissurine, and appear
16
SMITHSONIAN CONTRIBUTIONS TO BOTANY
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 colliciilosa externally.
Quite similar in morphology and anatomy is Graphis floridana Tuckerman (1888); the latter spe- cies, however, has norstictic acid and probably has submuriform spores (and hence is referable to Ciraphina ).
Specimens Examined. — Layou Road, northwest of Pont Casse, 1400 ft, Hale 38005 (US): Elfin woodland, Boeri Lake, 2750 ft, Imshaug 33208 (MSC).
12. Graphis humilis Figure 4/i; Plate 2/
Graphis humilis Vainio, 1921:256 [type collection: Philip- pines, Merrill 9067 (TUR, not seen; US, isotype)].
Description. — Thallus dirty white to pale tan, thin (endophloeodal?), continuous, matte. Asco- carps 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 sec- tion: hymenium 60-75 p,m high, I-; exciple quite open (rudimentary) basally, lips convergent, car- bonized apically only (type specimen) to apically and laterally. Spores 8/ascus, always 4 locular, 7-9 X 14-25 /xm, I-f- blue.
Chemistry. — No substances present.
Habitat.^ — Wet lowland and coastal forest.
Discussion. — The Dominican material matches the Philippine isotype closely, differing only in hav- ing somewhat larger spores and more carbonized labia. The carbonization makes this species distinct in the fissurine alliance; the closest relative is prob- ably Graphis tachygrapha, which has less carboniza- tion, 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, how- ever) is Phaeographis arthonioides.
Specimen Examined. — Madjiiii, 100 ft, Hale 35698 (US).
13. Graphis imshaugii, new species Figure 4/; Plate 3a
Description. — Thallus continuus, laevis, proso-
plectenchymatus. Apothecia subimmersa vel semi- emergentia, flexuosa, ramulosa, 2^ mm longa, margine thallino crassiusculo, cum thallo concolore, ab eo fissura tenuissima separato. Excipulum vulgo integrum vel subintegium, fuligineum, labiis inte- gris, erectis; hymenium 120-140 ju,m ahum, I-. Asci 8 spori; sporae decolores, 8-10 loculares, 6-9 X (17-) 23-35 /xin, 1+ coeruleae.
Thallus continuous, thick, glossy, green-gray, cor- tex 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 com- monly separating from the exciple, which is black and pruinose. Disc barely visible in surface view. Ascocarp transverse section: hymenium 120-140 jxm high, I-; exciple black, closed to nearly open, lips intact to rarely slightly striate, upright to somewhat spreading. Spores 8/ascus, 8-10 locular, 6-8 X (17-) 25-35 jum, 14- blue.
Chemistry. — Stictic acid, constictic acid. Holotype. — Cultivated area. South Chiltern Estate Road, 1500-1700 feet, Imshaug 32724B (MSC).
Discussion. — Graphis imshaugii is closely related to G. subamylacea Zahlbruckner (1921), from Mex- ico. 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.
14. Graphis insidiosa Figure 5a; Plate 3&
Graphis insidiosa (Knight anti Mitten) Hooker lil., 1867:586. Fissurina insidiosa Knight and Mitten, 1860:102 [type collec- tion: New Zealand, Knight 259 (BM, lectotype designated by Hayward, 1978)].
Graphis beaumontii Tuckerman, 1888:124 [type collection:
Alabama, USA, Beaumont 324 (EH, lectotype)].
Graphis interversa Nylander, 1889:49 [type collection: Flor- ida, USA, Eckjeldt 339 (H, lectotype)].
Graphis lactea var. clausa \'ainio, 1915:163 [type collection: Sotifriere, Dominica, Elliott 1511 (TUR, lectotype)].
Description. — 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. leptocarpa (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 {Pit- tiers s.n.).
18
SMITHSONIAN CONTRIBUTIONS TO BOTANY
bumps and rugosities, in section showing a poorly developed prosoplectenchymatous cortex. Ascocarps 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 sec- tion; hymenium 90-140 ^um high, I-; exciple pale, open, lips convergent to slightly gaping, pale to barely darkened apically, rudimentary below. Spores 8/ascus, always 4 locular, (5-) 8-11 X (11-) 13-22 (-24) /xm, I- or 1 4- slow, reddish.
Chemistry. — No substances present in all except Hale 35313, which has 2-3 unknown spots in TLC.
Habitat. — Wet lowland forest (US), wet upland forest (Dominica).
Discussion. — Examination of the types of Graphis insidiosa, G. beaumontii and G. interversa indicates that these three taxa are identical. The type of G. lactea var. clausa 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 Wirth collections; we feel, however, that these differences are too minor to warrant naming.
Graphis insidiosa is most similar to G. triticea, from which it differs in lacking stictic acid, having lower ascocarps with a less elaborate exciple, and in a less prominently prosoplectenchymatous cortex. The two species are, however, quite similar and are most easily separated by their chemistry.
A more complete discussion of the fissurine com- plex may be found under Graphis triticea.
Specimens Examined. — Can-Dom logging area, Dleau Gom- mier, 1600-1700 ft. Hale 35313 (US); Micotrin, ca 3000 ft, Wirth 463A (US).
15. Graphis isidiifera, new species Figure 5b; Plate 3c
Description. — Thallus laevis, isidiatus; stratum corticale prosoplectenchymatum. Apothecia im- mersa, rotunda vel sublirellina, ca. 1 mm longa, 0.5 mm lata, margine thallino, crasso, pallido, farinu- loso, discum bene superante, disco clilatato, sal- moneo. Excipulum integrum, pallidum, labiis integris, divergentibus; hymenium 70-80 ^m ahum, I-. Asci 8 spori; sporae decolores, 4 loculares, 4-6 X 12-13 /xm, I-.
Thallus thick, cracked, glossy, pale greenish gray, distinctly isidiate, cortex prosoplectenchyma-
tous. Ascocarps with prominent white mealy mar- gins, nearly round to quite irregular, clumped and branched, ca. 1 mm long, 0.5 mm wide. Disc quite exposed, pale salmon-pink, lightly pruinose, Asco- carp transverse section: hymenium 70-80 ^am high, I-; exciple closed, pale, lips entire, spreading. Spores 8/ascus, always 4-locular, 4—6 X 12-13 /tm, I-.
Chemistry. — Norstictic acid, unknown substance.
Holotype. — Virgin upland rain forest, Can-Dom logging area at Dleau Gommier, 1600-1700 feet. Hale 35179 (US).
Discussion. — Graphis isidiifera is unlikely to be confused with any other Graphis species. The com- bination 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 Graphis spe- cies, 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 tuberculate- isidiate species occur in Graphina, (viz., Graphina dimorphodes (Nylander) Zahlbruckner, 1923), Graphina dealbata (Nylander) Mueller Argoviensis (1895a), and its close allies G. albostriata (Vainio) Zahlbruckner (1923), G. heteroplacoides Redinger (1933), G. rimiilosa Redinger (1933), and Phaeo- graphina { = Graphina!) includens (Vainio) Zahl- bruckner (1923).
16. Graphis leptocarpa Figure 5c; Plate 3d
Graphis leptocarpa F^e, 1824:36 [type collection: South
America, Humboldt and Bonpland s.n. (G, lectotype)].
Description. — Thallus white to off-white, con- tinuous, smooth to slightly roughened. Ascocarps raised, black, unbranched to occasionally branched, straight to somewhat flexuose, with a slightly raised thalline margin, 1-3 mm long, slender. Disc not visible in surface view. Ascocarp transverse section: hymenium 90-100 pin high, I-; exciple black later- ally, absent below, lips more or less convergent. Spores 8/ascus, 8-9 locular, 5-8 X 16-23 pm (in the type) to 42 pin (in the Dominican and other material), I-l- blue.
Chemistry. — 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 fam- ily. The type species of the Graphidaceae, Gr aphis 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 mate- rial 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 ju,m range):
Graphis lineola Acharius (1810) is probably the oldest and best name for those tropical eugraphids which are P- (and presumably TLC negative, al- though as far as we know the type has not yet been chromatographed).
Graphis tenella Acharius (1814), another P nega- tive 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 Hel- sinki 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 nega- tive, is probably synonymous with G. lineola.
Graphis pavoniana Fee (1824), another TLC negative type, may be distinct from G. lineola in being larger overall, with only 4-6 larger spores (to 50 fjLxn) per ascus.
Graphis librata is the norstictic equivalent of G. lineola (P-) and G. leptocarpa (stictic acid).
Graphis caesiella Vainio (1890), with norstictic acid, differs from G. librata primarily in having pruinose ascocarps and discs.
Graphis dussii (above), with norstictic acid, dif- fers from G. librata in overall size and in having larger spores.
Specimens Examined. — Pointe Michel, 200 ft, Imshang 33I09A (MSC); Morne Conliabon, Elliott 1535 (TUR).
17. Graphis librata Figure 5d; Plate 3e
Graphis librata Knight, 1884:404 [type collection: New Zea- land, Knight 67:23 (WELT, lectotype designated by Hay- ward, 1978)].
Graphis crebra Vainio, 1899:256 [type collection: Guadeloupe, Duss 541 (TUR, lectotype)].
Graphis plumierae Vainio, 1915:161 [type collection: Guade- loupe, Duss 1189 (TUR, lectotype)].
Graphis tenellula Vainio 1915:169 [type collection: Santo Domingo, Raunkiaer 490 p.p. (TUR, lectotype)].
Description. — 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 p,m high, I-; exciple quite open below, lips carbonized, entire, more or less convergent; spores 8/ascus, 6-8 locular, 5-9 X 18-35 pin, 1 4- blue.
Chemistry. — Norstictic acid.
Habitat. — Coastal scrub to lower rain forest. Discussion. — Graphis librata is the norstictic equivalent of Graphis leptocarpa; for a more com- plete 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 331 12A (MSC); Coulibistri, 200 ft. Hale 35757 (US); Emerald Pool, Wirth 504B (US).
18. Graphis longula Figure 5e; Plate 3/
Graphis longula Krempelhtiber, 1876:414 [type collection;
Brazil, Glaziou 5497 (M, lectotype)].
Graphis flavicans Mueller Argoviensis, 1895a:457 [type collec- tion: Rio, Brazil, Glaziou 5498 p.p. (M, lectotype)]. Phaeographis longula (Krempelhuber) Zahlbruckner, 1923: 379.
Description. — Thallus white, grenish white, or 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. Asco- carp transverse section: hymenium 60-90 pm high, I-; exciple black, more or less closed to barely open (variable even within a single thallus), lips conver- gent, striate (appearing only crenate when the grooves are abundant and closely packed). Spores (?4-) 6-8/ascus, (8-) 10-17 locular, 9-13 X (40-) 70-90 pm, 1 4- blue.
20
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Chemistry. — Type o£ Graphis longula P-, no TLC available; type of G. flavicans and Dominican specimens, no substances present.
Habitat. — Lower edges of rain forest.
Discussion. — The tyjres of Graphis longula and G. flavicans are extremely similar; note the consecu- tive collector’s numbers.
The exciple cross-section illustrated for the type of G. longula in Wirth and Hale (1963) is mislead- ing; 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), which 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 dis- cussion may be found under the latter. Graphis flexibilis itself is separable by its massively closed exciple and larger more protuberant ascocarps.
The Graphis rimulosa gioup differs in having smaller spores and more emergent ascocarps with no thalline margin.
Specimens Examined. — Morne Bruce, 400 ft, Imshaug 33243 (MSC); Caii-Doin logging area, Newfoundland, 800 ft. Hale 33225 (US).
19. Graphis lumbricina
Figure 5/; Plate 4a
Graphis lumbricina \'ainio, 1899:256 [type collection: Guade- loupe, Duss 1036 (TUR, lectotype; FH, isotype)].
Description. — 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-) 170- 200 p,m 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) ^m, I-t- blue.
Chemistry. — Norstictic acid (isotype).
Habitat. — Rain forest, mossy forest.
Discussion. — Graphis lumbricina is very similar to G. flexibilis; the latter differs in a more massively carbonized excipular base and in lacking norstictic acid. Graphis congesta (Fee) Mueller Argoviensis (1887) differs only in having short, congested, asteroidly-branched ascocarps. For a more complete discussion of this alliance, see Graphis flexibilis.
Specimens Examined. — Can-Dom logging area, Newfound- land, 800 ft. Hale 35240 (US); Fresh Water Lake, 2600-2800 ft. Hale 35449 (US); Trois Pitons, 29-3150 ft, Imshaug 33084A, 33087 (MSC).
20. Graphis olivacea Figure 5g; Plate Ab
Graphis olivacea Redinger, 1935:55 [type collection: Matto
Grosso, Brazil, Malme 2267B (S, lectotype)].
Description. — Thallus pale olive to whitish, smooth, glossy, continuous to rather eroded, and restricted to near the ascocarps in the high eleva- tion specimens. Ascocarps straight to flexuose, occa- sionally branched, apically black and striate (at least in the largest and 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. Asco- carp transverse section: hymenium 130-200 ju,m high, I-; exciple black above, yellow-red to brown- red below, usually closed, lips convergent, striate. Spores 6-8/ascus, 10-18 locular, 8-10 (-13) X (45-) 60-90 (-125) /cm, 1 4- blue.
Chemistry. — No substances present.
Habitat. — Wet rain forest into elfin forest, where it is very common on Clusia venetiosa with Phaeo- graphis exaltala and P. rnorclenii.
Discussion. — Graphis olivacea is very similar to G. flexibilis, from which it differs in having the ex- cipular base constructed of partially uncarbonized, yellow- to red-brown tissue. It may well be that this difference is not significant. Also quite similar is G. rigidula, which differs in being smaller overall. For a more complete discussion of this alliance, see G. flexibilis.
Graphis olivacea may prove to be a synonym of the Philippine G. glauconigra Vainio (1921), which appears to differ only in the smaller ascocarps and brown thallus.
Si’Eci.MENS Examined. — Caii-Dom logging area, Newfound- land, 800 ft, Hale 35062 (US); Dleau Gommier Forest Reserve,
NUMBER 40
21
1200 ft. Hale 37988 [US); South Chiltern Estate Road, 1500- 1700 ft, Imshaug 32730 (MSC); Soufri^re Ridge, 2200-3100 ft, Imshaiig 33069A [MSC); Trois Pitons, 2900-3150 ft, Imshaug 33089 (MSC), 4000-4672 ft, Imshaug 32874A [MSC), summit, Wirth 487, 491 (US); Morne Diablotin, 4300-4550 ft, Imshaug 32904A, 32909 (MSC).
21. Gr aphis rigidula Figure 5h; Plate 4c
Graphis rigidula Mueller Argoviensis, 1891:78 [type collec- tion; San Jose, Costa Rica, Pittier 5291 (G, lectotype)].
Description. — Thallus 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 jxm 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 /xm, I-f blue.
Chemistry. — No substances present (? trace of norstictic acid in the type?).
Habitat. — Secondary forest.
Discussion. — Graphis rigidula is extremely sim- ilar to G. longula; 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.
Specimen Examined. — Bois Serpe, 1000 ft, Imshaug 32777A (MSC).
22. Graphis rimulosa Figure 6a; Plate 4d
Graphis rimulosa (Montague) Trevisan, 1853:11.
Opegrapha rimulosa Montagne, 1842:271 [type collection:
Guyana, Leprieur 200 (P, lectotype)].
Description. — Thallus 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: hyme- nium 60-90 fj.m 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) firm, 1+ 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 ap- pears to be the only difference between the two species.
Graphis strialula (Acharius) Sprengel (1827): The type material at Helsinki has much shorter asco- carps than G. rimulosa and resembles Opegrapha or Melaspilea. In addition, a section from the iso- type at Uppsala shows a more or less oj^en excipu- lar 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.
Specimens Examined. — South of Portsmouth, sea level. Hale 35699 (US); Roseau V'alley, Elliott 125 (TUR); Central Forest Reserve, 1500 ft, Imshaug 33549.\ (MSC).
23. Graphis subelegans Figure 66; Plate 4e
Graphis subelegans Nylancler, 1891:42 [type collection: San
Luis Potosi, Mexico, Pringle 162 (H, lectotype)].
Description. — Tliallus usually yellow-brown, off- white 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 /xm high, I-; exciple black laterally, yellow to yellow- red below, open, lips convergent, cjuite striate. Spores 8/ascus, 6-8 locular, 6—8 (-10) X 25-30 /xm, 1+ blue.
Chemistry. — No substances present.
Habitat. — Sea level scrub (probably low eleva- tion exposed scrub throughout its range).
Discussion. — Graphis subelegans has been placed in synonymy (Zahlbruckner, 1923) of G. endox- antha Nylancler (1868); however, the type of the latter [Pancher 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. rirnulosa (lectotype in P); b, G. siibe- legans (Pringle 162); c, G. subnitidula (Wright 155); d, G. tachygrapha (Lindig 869); e, G. triticea (Lindig 841); /, G. turgidula (lectotype in BM).
There is a large group of species related to G. suhelegans, 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 asco- carps. Much additional collecting is necessary to unravel this alliance.
Specimen Examined. — Rodney’s Rock, sea level. Hale 35546 (US).
24. Graphis subnitidula
Figure 6c; Plate 4/
Graphis subnitidula Nylander m Tuckerman, 1888:123 [type collection: Cuba, Wright 155 (US, lectotype; ?FH, isotype, not seen)].
Graphma subnitidula (Nylander in Tuckerman) Zahlbruck- ner, 1923:427.
Description. — Thallus brownish to greenish.
NUMBER 40
23
smooth, glossy, continuous, distinctly prosoplecten- chymatous 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 visi- ble between the gaping lips. Ascocarp transverse section: hymenium 80-100 jxm high, I-; exciple open, lips pale, convergent but exposing some epi- hymenium. Spores 8/ascus, 4 locular, 5 X 8-10 fixn, I- or 1+ 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 collec- tion 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). Tucker- man’s description clearly recognizes the mixed na- ture of the spores; we have chosen to follow what appears to be Nylander’s original concept and con- sider 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 Graphis triticea.
Specimen Examined.— Pont Casse, 2200-2600 ft, Hale, 37676 (US).
25. Graphis tachygrapha Fignre 6d; Plate 5a
Graphis tachygrapha Nylander, 1863:371 [type collection:
Colombia, Lindig 869 (H, lectotype; FH, isotype)].
Graphis lactea var. dominicana Vainio, 1915:162 [type collec- tion: Trois Pitons, Dominica, Elliott 535 (TUR, lectotype)]. Graphis timida Vainio, 1923:142 [type collection; Trinidad, Thaxter 30 (FH, lectotype)].
Description. — Thallus very thin, apparently en- dophloeodal, probably taking most of its color and texture from bark and/or epidermis. Ascocarps fis- surine, beginning as cracks and finally gaping, un- branched, 0. 5-1.0 mm long. Dark disc visible in gaping stages. Ascocarp transverse section: hyme- nium 60-90 p,m high, I-, epihymenium slightly to much darker than the rest of the hymenium; ex- ciple rudimentary below, lips convergent when young, then spreading broadly, partially or incom- pletely 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 p,m, I- or 1 4- slow, reddish, pale.
Chemistry. — No substances present.
Habitat. — Elfin forest in Dominica; the type of G. tachygrapha is from 2600 meters in Colombia.
Discussion. — Very similar to Graphis tachygrapha is G. humilis, which is tentatively separable by its I-b spores and by the much more heavily carbon- ized lips. For a more complete discussion of the fissurine alliance, see Graphis triticea.
Specimens Examined. — Mt. Soufriere, Elliott 1515 (TUR); Trois Pitons, summit, 4000-4672 ft, Imshaug 32862 (MSC).
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 gTayish, thick, glossy, almost always deeply cracked, cortex quite prosoplectenchymatous. Ascocarps concolor- ous 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 p,m high, I-; exciple yellow below, open to almost closed, laterally yellow, mixed with bark cells, apically convergent to somewhat spreading, fre- quently 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 /j,m, I- or 1 4- slow, reddish.
Chemistry. — Stictic acid.
Habitat. — In or just below elfin forest; prob- ably pantropical in high altitude rain forests.
Discussion. — 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 (Hay- ward, 1978) also lack this white tissue.
The number of names in the fissurine alliance is
24
SMITHSONIAN CONTRIBUTIONS TO BOTANY
large. To distinguish these generally rather incon- spicuous species, we have found the following fea- tures most useful: (1) presence or absence of stictic acid; (2) thickness and degree of specialized organi- zation 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 ma- ture 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 gioup); (6) iodine reaction of spores; (7) overall size of ascocarps and size of spores (this character is useful only occasionally, as most fis- surines 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 (Fte) Mueller Argoviensis (1887): An early fissurine name that should be permanently dropped; the type was sterile when described.
Graphis durnastii 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 gram- mitica Nylander (1866) may be synonym.
Graphis diimastioides Fink: Stictic acid; thallus thick, prosoplectenchymatous; no “puffing”; asco- carps gaping; no carbonization; spores I- (or red- dish) average size for the alliance (stictic acid equiv- alent of G. durnastii).
Graphis humilis Vainio: No stictic acid; thallus thin; no “puffing”; ascocarps non-gaping; exciples carbonized; spores I-H; slightly smaller ascocarps than average.
Crraphis inquinata (Knight and Mitten) Hooker f. (1867): Stictic acid; thallus thick, apparently not prosoplectenchymatous; some “puffing”; ascocarps nongaping; exciple carbonized; spores I- (or red- dish); average size for the alliance (rather like a carbonized equivalent of G. triticea).
Graphis insidiosa (Knight and Mitten) Hooker f.: No stictic acid; thallus thick, apparently proso- plectenchymatous; usually quite “puffed”; asco- carps usually not gaping; no carbonization; spores I- (or reddish); ascocarps slightly larger than aver- age (nonstictic equivalent of G. triticea).
Graphis suhnitidula 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 durnastii; in all probability a whole series of sporo- morphs between these two alliances awaits dis- covery.
Specimens Examined. — Morne Anglais, 3000-3600 ft. Hale 35324, 35326B, 35377 (US); Morne Diablotin, 3200-4600 ft.
Hale 35438, 35442 (US), 4300-4500 ft, Imshaug 32912 (MSC), Trois Pitons, Elliott 533 (TUR), 3000-4000 ft, Imshaug 32886 (MSC), summit, Wirth 482 (US); Micotrin, ca 3000 ft, Wirth 463B (US).
27. Graphis turgidula Figure 6/; Pe.ate 5c
Graphis turgidula Mueller Argoviensis, 1895a;457 [type col- lection: Mauritius, s.n. (BM, lectotype)].
Description. — ^Thallus white to off-white, thick, continuous, glossy. Ascocarps prominently raised, black above, thalline margin quite pronounced, straight to more or less ffexuose, rarely branched, 1-4 mm long. Ascocarp transverse section: hyme- nium (130-) 150-250 p,m high, I-; exciple black, massively clo.sed 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) /xin, 1+ blue.
Chemistry. — Stictic acid.
NUMBER 40
25
Habitat. — Dominican specimen from lower mon- tane rain forest, on Hibiscus elatus.
Discussion. — 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. an-
g^lilliformis, and is separable only in having stictic acid. One fairly poor specimen from Dominica {Hale 35069, Can-Dom logging area, Newfound- land) is morphologically identical to G. turgidula but has norstictic acid. Whether this represents a different species, or a chemically polymorphic popu- lation, remains to be seen.
Specimen Examined. — North of Pont Cass^, 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 30. P. exaltata
1. Exciple without carbonization.
3. Ascocarps pink or cinnabar-red in surface view.
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. Spores over 100 p.m long, usually much longer 32. P. mordenii
5. Spores under 35 yum long.
6. Spores 4 locular, under 20 yum long; ascocarps usually clumped and radiately
branched 34. P. cf. subtigrina
6. Spores 6-8 locular, over 20 fiva long, ascocarps not clumped or radiately branched 28. P. albidh
28. Phaeographis albida Figure 7a, b; Plate 5d
Phaeographis albida (Vainio) Zahlbruckner, 1923:364. Graphis albida \'ainio, 1896:251 [type collection: Morne
Anglais, Dominica, Elliott 528 (TUR, lectotype)].
Description. — Thallus white to off-white, con- tinuous, 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 conver- gent labia, sometimes completely exposed, dark brown in surface view. Ascocarps transverse section: hymenium (70-) 80-150 yu,m high, I-f- 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 epi- hymenium. Spores (4-) 6-8/ascus, 6-8 locular, (6-) 8-13 X 20-35 yarn, frequently halonate after release, I-f- blue (in the type some cells appearing very irregularly muriform).
Chemistry. — No substances present.
Habitat. — Elfin forest and high altitude rain forest.
Discussion. — Phaeographis albida is a distinct species, apparently endemic to Dominica. The Hale and Imshaug collections have ascocarps with spread- ing 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 spe- cies with uncarbonized exciples. A very good paral- lel example is illustrated for Graphina colliculosa.
This is the only species known to us with a varia- ble 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- hymenium, and the presence of stictic acid.
Specimens Examined. — Fresh Water Lake, 2600-2800 ft. Hale 35453 (US): Morne ]5iablotiu, 3500^300 ft, Imshaug 32936 p.p. (MSC).
29. Phaeographis arthonioides
Figure 7c; Plate 5c
Phaeographis arthonioides (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); /, P. haematites {Hale 38013); g, P. cf. subtigrina {Wirth 444); h, P. rosea {Elliott 523).
NUMBER 40
27
Graphis arthonioides V^ainio, 1915:155 [type collection: St.
Croix, Raimkiaer 548 (FH, isotype)].
Phaeographis sexloculata Fink, 1927:215 [type collection:
Puerto Rico, Fink 1436 (MICH, lectotype)].
Description. — Thallus giay to whitish, contin- uous, glossy, rather thin. Ascocarps fissurine, barely raised, margins blackish to black, mostly un- branched, straight to somewhat flexuose, usually 1 mm or less long. Disc usually not visible in sur- face view. Ascocarp transverse section: hymenium 65-70 /xm high, I-, epihymenium dark; exciple rudimentary below, lips more or less convergent, irregularly carbonized at least apically, frequently to the base of the hymenium. Spores 8/ascus, 6-8 locular, 7-10 X 19-26 pm, I- or I-f slow, reddish.
Chemistry. — 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.
Discussion. — Extremely similar to Phaeographis arthonioides is P. decipiens (Fee) Mueller Argovien- sis (1887), also from the Caribbean; the latter differs only in having stictic acid. Differing only in spores is Graphis humilis Vainio.
Specimen Examined. — East of Pointe Michel, 200 ft. Ini- shaug 33128 (MSC).
30. Phaeographis exaltata Figure 7d; Plates 5/, 6a
Phaeographis exaltata (Montague and van den Bosch) Muel- ler Argoviensis, 1882:336.
Lecanactis exaltata Montague and van den Bosch, 1855:475
[type collection: Java, Junghuhn s.n. {L, lectotype)].
Description. — Thallus nearly white to gray, thick, usually glossy, continuous. Ascocarps very prominent, from Sarcographa-\\k.e and circular to elongate and flexuose (on one thallus), thalline mar- gin very prominent, from 1x1 mm to 1 X 6 mm. Disc wide, black to gray-pruinose. Ascocarp trans- verse section: hymenium 120-200 pm high, I-, epi- hymenium blackened; exciple black, heavily closed below, lips usually quite spreading, intact, fre- quently incompletely carbonized. Spores 8/ascus, 6-8 locular, 7-1 1 X 20-30 pm, I- or I-F faint, slow, reddish.
Chemistry. — See “Discussion.”
Habitat. — Upper montane and elfin forest. One of the three major graphids on elfin forest Clusia venenosa (with Graphis olivacea and Phaeographis mordenii).
Discussion. — Phaeographis exaltata is a wide- spread, 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 Phaeo- graphis. 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 chem- ical 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 Examined. — 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); Morne Anglais, 3000-3600 ft. Hale 35322 (US); Trois Pitons, 3000-4000 ft, Imshaug 32883, 32888 (MSC), summit, Wirth 480, 483 (US); Morne Diablotin, 3500-4300 ft, Imshaug 32934 (MSC), 3200-4600 ft, Hale 35323, 35445 <US); Prince Rupert, Elliott 1307 (TUR).
31. Phaeographis haematites Eigure 7/; 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, con- tinuous, smooth to matte. Ascocarps raised, promi- nent, 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 /xin 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 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.
Discussion. — Phaeographis cinnabarina (Fee) Mueller Argoviensis (1887) is the only species likely to be confused with P. haematites. The former can easily be distinguished by its thinner, duller asco- carps, 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.
Specimen Examined. — Dleau Gommier Forest Reserve, 1200 ft. Hale 38013 (US).
32. Phaeographis mordenii, new species Figure 7e: Plate 6c
Description.— Thallus continuus, laevis vel rugu- loso-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 rufo- ferrugineum, labiis integris, divergentibus; hyme- nium inspersum, 200-250 /xm altum, I-. Asci 6-8 spori, sporae fuscescentes, 30-40 (-50) loculares, 12-17 X 120-220 fjLxn, in mentem revocans sporae Conotremae, 1+ rufo-fuscae.
Thallus light gi ay 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 thalline margin, 2-8 mm long, ca. 1 mm wide. Disc very broad, initially concolorous with the thallus, then dark blackish- gray and prtiinose. Ascocarp transverse section: hy- menium 200-250 /un high, heavily inspersed and beaded, epihymenium darkened, I-; exciple closed to nearly open, reddish yellow, labia intact, diver- gent, covered by the thalline margin. Spores 6-8/ ascus, brown, very long and slender, 30-40 (-50)
locular, 12-17 X 120-220 ^m, occasionally with irregular biocellations in the end chambers, cham- ber walls frequently thickened, reminiscent of Conotrema spores, 14- slow, eventually dark red- brown.
Chemistry. — No substances present.
Holotype. — Mossy and elfin forest, trail to sum- mit of Morne Diablotin, Dominica, 3200-4600 feet. Hale 35071 (US).
Discussion. — Phaeographis mordenii is one of the three characteristic and abundant graphids on Clusia venenosa in the elfin forest. It is quite dis- tinct 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; how- ever, these extra locules are irregular and seem to represent disintegration of cell contents rather than a submuriform condition.
Specimens Examined. — Monie Diablotin, 3500-4300 ft, Im- shaitg 32922, 32945A (MSC); Morne Trois Pitons, 3000-41000 ft, Irnshaiig 32884 (MSC), summit, on Clusia venenosa, Wirth 479, 481, 484, 485, 486 (US).
33. Phaeographis rosea Figure 7/;; Plate 6d
Phaeographis rosea (\’ainio) Zahlbruckner, 1923:385.
Graphis rosea \’ainio, 1896:259 [type collection: Morne
Anglais, Dominica, Elliott 523 (TUR, lectotype)].
Description. — Thallus tan, continuous, smooth and shining. Ascocarps barely emergent, rather fis- surine, 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 p,m high, I-; exciple rudimentary, labia convergent, brownish above. Spores 8/ascus, always 4 locular, 9-12 X (13-) 18-25 p,m, 14- blue.
Chemistry. — No substances present.
Habitat. — Upper montane forest.
Discussion. — Phaeographis rosea is known only from the type collection. The rose-pink pigmenta- tion that renders this species so distinct is appar- ently a normal condition; occasionally, some graphids will show discolored, reddish areas on thalli of normally unpigmented species (see Gra-
NUMBER 40
29
phina dimidiata for a Dominican example). The color in P. rosea, however, is restricted to the asco- carp margins, and appears to be normal.
34. Phaeographis cf. subtigrina
Figure 7g; Pl.\te 6e
Phaeographis subtigrina (\'ainio) Zahlbruckner, 1923:387. Graphis subtigrina Vainio, 1907:177 [type collection: Koh Chang, Gulf of Siam, Schmidt XXI (TUR, lectotype)].
Description. — Thallus thin, off-white, continu- ous, matte, Ascocarps usually clumped, commonly radiately branched, forming clusters to 2 mm in diameter, flush, thalline margin frequently separat- ing from the thallus. Disc broad, gray, lightly but distinctly pruinose. Ascocarp transverse section: hymenium 115 /xin high (Dominican specimens; no measurement for the type), I-; exciple pale below, closed to nearly open, lips darker but not carbon- ized, intact, divergent. Spores 8/ascus, thick-walled, 4 locular, 4-6 X 10-17 /xm, I-.
Chemistry. — Norstictic acid only (type); norstic- tic and stictic acid (Dominican specimens).
Habitat. — One Dominican specimen from sea- level, the other from upland rain forest. Apparently of great ecological amplitude.
Discussion. — Phaeographis 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. subtigrma varies gueatly within the same thallus. In the Dominican specimens these struc- tures 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.
Speci.viens Examined. — Roseau Botanic Garden, iVirth 444 (US); Emerald Pool, Wirth 505 (US).
Key to the Species of Graphina
1. Exdples distinctly carbonized.
2. Disc broad and obvious in surface \iew, labia quite di\ergent.
3. Spores with broad persistent colloidal layer; ascocarps flush with thalline surface
41. G. collospora
3. Spores without colloidal layer; ascocarps more or less raised.
4. Spores 1/ascus; protocetraric acid present 52. G. platyleuca
4. Spores more than I/ascus, some darkening; no substances present
63. Phaeographina cf. di(formis
2. Disc narrow, not prominent in surface view, labia more or less con\crgent.
5. Ascocarps with powdery white covering, easily ridjbeil to expose the black exciple.
6. Spores under 20 jj.m long; salazinic acid present 50. G. viarcescem
6. Spores over 40 ,um long; stictic acid present 57. G triphoroides
5. Ascocarps without a powdery white covering.
7. Labia entire or nearly so.
8. Exciple carbonization limited to labial apices (upper i/o or less of total exciple
height 43. G. dimidiata
8. Exciple carbonization extending to base of hymenium or beyond.
9. Ascocarps Opregrapha-like, 1—2 mm long, without a thalline margin
51. G. Hilda
9. Ascocarps elongate, flexuose, with a thalline margin.
10. Spores less than 61 jj.m long.
11. Norstictic acid present; mature spores 8/actis
54. G. pseitdoaiialoga
11. No substances present; mature spores usually 4-6/ascus
53. G. pliirispora
30
SMITHSONIAN CONTRIBUTIONS TO BOTANY
10. Spores over 85 /tin long.
12- Only the ends of the spores muriform 58. G. vestitoides
12. Entire spore densely muriform.
13. Ascocarps under 0.5 mm wide; hymenium under 250 p.m high.
35. G. acharii
13. Ascocarps massive, usually 1 mm wide; hymenium over 350 ^m high 46. G. illinata
7. Labia striate.
14. Exciple yellow to yellow-brown laterally, more or less oj>en ....37. G. antillarum
14. Exciple without yellowish areas, closed or nearly so.
15. Spores under 61 _um long.
16. Norstictic acid present; mature spores 8/ascus 54. G. pseudoanaloga
16. No substances present; mature spores usually 4-6/ascus
53. G. plurispora
15. Spores over 64 long.
17. Only ends of spores muriform 58. G. vestitoides
17. Entire spore densely muriform.
18. Exciple base irregularly carbonized; spores always 1/ascus
49. G. macella
18. Exciple base usually evenly carbonized; sjxtres usually 4/ascus
35. G. acharii
1. Exciples totally uncarbonized.
19. Apothecia fissurine, i.e., originating as a crack, the disc usually concealed by or sunken
between the narrow, to gaping fissure walls.
20. Sptores 1/ascus, over 50 /an long 48. G. insculpta
20. Spores 6-8/ascus, under 50 ^um long.
21. Thallus prosoplectenchymatous 47. G. incrustans
21. Thallus not prosoplectenchymatous.
22. Psoromic acid present 42. G. columbina
22. No substances present 36. G. cf. adscribens
19. Ascocarps not fissurine.
23. Disc broad, easily seen in surface view.
24. Disc reddish; spores under 17 /iin long; salazinic acid present ....40. G. coUiculosa 24. Disc whitish, gray to greenish; spores over 17 /im long; salazinic acid absent.
25. Spores 17-28 /jin long; norstictic acid present 56. G. suberythrella
25. Spores 50-150 fxm 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 1/ascus, over 80 jum long 39. G. chlorocarpa
27. Spores 2-8/ascus, under 75 //in long.
28. Norstictic acid present; exciples with intenral striae only
44. G. dispersa
28. No substances present; internal striae absent.
29. Mature spores 2-4 ascus, over 35 ^m long; labia slightly striate
55. G. rufopallida
29. Mature spores 8/ascus, less than 18 jim long; labia entire
38. G. carneoviridis
26. Margins of ascocarps tan, white, or greenish.
30. Spores less than 17 /im long; salazinic acid present 40. G. coUiculosa
30. Spores more than 40 fitn long; salazinic acid absent.
31. Ascocarps always massive and protuberant; hymenium over 200 /^m
high; I-; no substances present 45. G. jrumentaria
31. Ascocarps rarely protuberant, never massive; hymenium less than 200 jum high, at least the epihymenium 1+ blue; “quintaria” unknowns present 59. G. virginea
NUMBER 40
31
35. Graphina acharii Figure 8a; Plate 6/
Graphina achaii (Fee) Mueller Argoviensis, 1887:38.
Graphis acharii Fee, 1824:39 [type collection: South America, s.n. (G. lectotype)].
Graphis inturgescens Krempelhuber, 1876:383 [type collec- tion: Brazil, Glaziou 6286 (BM, lectotype; M, isotype)]. Graphina inturgescens (Krempelhuber) Mueller Argoviensis, 1888a: 545.
Description. — I'hallus gray to oli-white, tliick, continuous to cracked. Ascocarps large, strongly raised, black above, straight to flexuose, occasion- ally branched, covered nearly to the tops by a prominent thalline margin, 1-6 mm long. Disc not visible in surface view. Ascocarp transverse section: hymenium 150-200 pin higli, 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 4/ascus but occasionally one or three (and rarely six), densely muriform, 18-25 X 95-170 pm, I-l- blue.
Chemistry. — No substances present.
Habitat. — Secondary forest (Dominica). Discussion. — 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 collec- tions 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 bio- cellate spores can be referred to G. vestitoides (p. 44).
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. in- turgescens 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-
phina species is G. macella, which differs in being smaller overall and in having an irregularly car- bonized excipular base.
Specimen Examined. — Syndicate Estate, 1800 ft. Hale 35555 (US).
36. Graphma cf. adscribens Figure 8b; Plate la
Graphina adscribens (Nylaiider) Mueller Argoviensis, 1892a:
284.
Graphis adscribens Nylander, 1868:177 [type collection: Lifu,
New Caledonia, Thiebaul s.n. (H, lectotype.].
Description. — Thallus white, thin, continuous to cracked. Ascocarps very inconspicuous in the Domi- nican material and in the type photographs, fis- surine, slightly mealy in the Dominican specimen, concolorous with the thallus (at least in the Domin- ican specimen), less than 1 mm long. Disc not visi- ble in surface view. Ascocarp transverse section: hymenium 100-120 pin high, (I- Dominican speci- men); exciple rudimentary in the Dominican mate- rial, apparently slightly reddish in the type, lips convergent, intact. Spores 8/ascus, muriform 8-9 X 21-27 pin (type), 9-10 X 23-40 (Dominica), I- in type (fide Nylander), 1+ reddish, slow in the Do- minican material.
Chemistry.^ — No substances present (both speci- mens).
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. Ac- cording to both Nylander and Mueller, the type is reminiscent of Graphina chlorocarpa and Graphis grammitis, both of which have reddish ascocarps in surface view. As the Dominican specimen is uni- formly pale, the identification must remain tenta- tive until the type can be examined.
Specimen Examined. — Ans du Me, Wirth 519 (US).
37. Graphina antillarum Eigure 8c; Plate lb
Graphma antillarum (Vainio) Zahlbruckner, 1923:398.
Graphis antillarum \'ainio, 1899:255 [type collections, Guade- loupe, Duss 540 (FH, isotype)].
32
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Figure 8. — Cross sections of apothecia of Graphhia: a, G. ncharii (lectotype in G); b, G. cf. adscribens (Wirth 519); c, G. antillarum (Duss s.n.); d, G. carneoviridis {Hale 37642); e, G. chlorocarpa (lectotype in G); f, G. colUculosa (lectotype of G. eugeriiae, Merrill 3471); g, G. colliculosa (lectotype in PC).
Graphis acuminata Vainio, 1915:147 [type collection: St. Jan, Raunhiaer 437 (TUR, lectotype)].
Graphis platycarpoides Vainio, 1915:145 [type collection, Gnacleloupc, Duss 1198 (TUR, lectotype)].
Graphina acuminata (Vainio) Zahlbruckner, 1923:393. Graphina platycarpoides (Vainio) Zahlbruckner, 1923:419.
Graphina sulcata Fink, 1927:217 [type collection: Puerto Rico, Fink 659 (MICH, lectotype)].
Descriptio.n. — I’hallus gray to white, continuous, matte. Ascocarps nearly flush to partially emergent, black, commonly branched and flexuose, thalline
NUMBER 40
33
margin absent or nearly so, 5-6 mm long, slender. Ascocarp transverse section: hymenium 80-100 (-120) fim high, I-, exciple black laterally, yellow to yellow-brown below, more or less open, labia con- vergent, barely to quite strongly striate. Spores (1-) 2-4 (-6)/ascus, distinctly muriform, 15-22 X 28-46 (-60) p.m, 1 + blue.
Chemistry. — Norstictic acid.
Habitat. — Lowland scrub forest and higher alti- tude cultivated areas and road cuts.
Discussion. — Among the Graphina species with carbonized, striate, dimidiate exciples, G. ajitil- larurn is fairly distinct by virtue of its 2-4 small, rotund spores per ascus, and the presence of nor- stictic acid. There are, however, several other spe- cies that are extremely similar and may prove to be conspecific:
Graphina bipartita Mueller Argoviensis (1888c) differs in having a thin thalline layer over the asco- carps 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 Argo- viensis (1892b) differs in having stictic acid and 8 spores per ascus.
Most of the Dominican specimens fit comfort- ably within the boundaries defined here for G. antillarurn. 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, Imshaug 33529A (MSC): Brookhill Estate, 100-150 ft, Imshaug 33307.\ (MSC); Roseau Botanic Garden, 100 ft Wirth 445 (US); north of Bioche, 200 ft, Hale 35745; South Chiltern Estate, 1300 ft, Imshaug 33050, 33036 (MSC); near Pont Casse, 1400 ft, Wirth 544c (US).
38. Graphina carneoviridis, new species Figure 8d; Plate 7c
Description. — Thallus continuus, laevis, stratum corticale prosoplectenchymatum. Apothecia sessilia, recta vel flexuosa, 1-2 (-3) mm longa. Excipulum dimidiatum, rufo-fuscum, labiis convergentibus, in-
tegris vel indistincte incisis; hymenium 75 p,m ahum, I-. Asci 8 spori; sporae decolores, murales, loculis horizontalibus 3, loculus transversis 2, 10-12 X 15-17 p.m, 1+ coeruleae.
Thallus greenish, continuous to somewhat cracked, glossy, prosoplectenchymatous. Ascocarps raised, occasionally branched, straight to fiexuose, pink-brown above, laterally bounded by a green thalline margin, 1-2 (-3) mm long, slender. Disc not visible in surface view. Ascocarp transverse sec- tion: hymenium 75 p,m high, I-; exciple reddish- brown, open below, labia convergent, intact to barely striate. Spores 8 /ascus, 3x2 locular, 10-12 X 15-17 /xm, 1+ blue.
Chemistry. — No substances present.
Holotype. — Mixed disturbed and primary rain forest, trail from Brigantin tlirough Middleham Estate toward Morne Trois Pitons, Dominica, ele- vation 2200-2600 feet. Hale 37642 (US).
Discussion. — Graphina carneoviridis is externally rather similar to Graphina dispersa, from which it differs in having smaller spores, a more open ex- ciple with nearly intact labia, a prosoplectenchyma- tous cortex, and in a different chemistry.
39. Graphina chlorocarpa Figure 8e; Plate Id
Graphina chlorocarpa (Fee) Mueller Argoviensis, 1887:44. Graphis chlorocarpa Fee, 1824:47 [type collection: Peru. s.n.
(G, lectotype)].
Graphina balbisii var. monospora Reclinger, 1933:61 [type
collection, Malme 494 (S, lectotype; FH, isotype)].
Description. — Thallus tan to grey-green, contin- uous, glossy, smooth to slightly rugose. Ascocarps usually raised (nearly flush when young), straight to quite fiexuose, 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 fiva high, I-; exciple yellow-brown to red- brown, open below, labia convergent, lightly striate, the tips of the striae sometimes slightly darkened. Spores 1/ascus, densely muriform, 24-40 X 80-130 fiui, I-f blue.
Chemistry. — Both types and most specimens: no substances present. A few Dominican specimens with unknown substances.
Habitat. — All fertile specimens from rain forest; sterile material (see discussion) from sea level and
34
SMITHSONIAN CONTRIBUTIONS TO BOTANY
mid altitude cultivated areas.
Discussion. — The only neotropical Graphina spe- cies likely to be confused with G. chlorocarpa is G. balbisii (Fee) Mueller Argoviensis. 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. chloro- carpa the asci are always monosporous; in G. bal- bisii the asci usually contain 3-4 slightly smaller spores, occasionally 6 or 2, but never only one. As yet, we have seen no specimens that bridge the rather small gap between these two species.
Without spores, it is not possible to be sure which of these two species is in hand. All the fertile Do- minican material referable to G. chlorocarpa is from rain forest; however, four additional speci- mens from cultivated areas {Imshaug 33024, 33417, 33424, Wirth 446) are sterile and may be either of the two species. No fertile material of G. balbisii has yet been identified from the Island.
Specimens Examined. — Near Laudat, 800 ft, Hale 35797 (US); Can-Dom logging area, Pont Cassc, 2000 ft, Hale 35097, 35129 (US); Ridgefield Estate, 1100-1200 ft, Imshaug 33417, 33424 (MSG, both sterile); South Chiltern Estate, 1300 ft, Imshaug 33024 (MSG, sterile); Roseau Botanic Garden, 100 ft, Wirth 446 (US, sterile).
40. Graphina colliculosa Eigures 8/,g; Plate 7e
Graphina colliculosa (Montague) Hale, 1976:156.
Sclerophyton colliculosum Montague, 1851:61 [type collection:
Guyana, Leprieur 1406 (P, lectotype; BM, isotype)]. Graphis intortula Stirton, 1881:186 [type collection: Assam, s.n. (BM, lectotype)].
Graphis cugeniae Y'ainio, 1921:262 [type collection: Philip- pines, Merrill 3971 (TUR, lectotype; EH, isotype)]. Graphis colliculosa (Montague) Zahlbruckner, 1923:299. Graphina intortula (Stirton) Zahlbruckner, 1923:411. Graphina aibonitensis Fink, 1927:215 [type collection: Ptierto Rico, Fink 2017 (MIGH, lectotype)].
Description. — Tlialhis 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 sur- face view, .sometimes completely covered by the con- colorous thalline margin. Ascocarp transverse sec- tion: hymenium 80-100 p,m high, I-, epihymenium
darkened; exciple rather ill-defined, yellow to red- dish-yellow, usually closed below, labia quite con- vergent 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 p,m, I-F blue.
Chemistry. — Salazinic acid.
Habitat. — Rain forest canopy in Dominica; else- where found in both rain forest and secondary forest.
Discussion. — Graphina colliculosa is an extremely vigorous, pantropical species; we have seen speci- mens from much of the Caribbean (Cuba, Puerto Rico, St. Vincent, Grenada, Trinidad, St. Lucia), Panama, Guayana, Tahiti, Fiji, Assam, and the Philippines.
As in many species with totally uncarbonized exciples, the degree of labial divergence is quite variable (see Phaeographis albida for a parallel con- dition). The taxa listed as synonyms differ from each other primarily in the extent to which the red- brown disc is exposed; as complete exposure and complete concealment can be found within a single thallus, this character seems clearly useless to dis- tinguish 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 exam- ple 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. colliculosa is unlikely to be con- fused with any other species, as salazinic acid is quite rare in the family. However, several other species are (juite similar in morphology, as follows.
Graphina erythrella (Montagne) Zahlbruckner (1923) is distinguished by having norstictic acid, larger spores, and somewhat shorter, more raised ascocarps. Some of the sterile, norstictic acid-con- taining syntypes of G. eugeniae (MacGregor 41315, Elmer 15077) may be referable to this species.
Graphina riopiedrensis Fink (1927) differs only in the somewhat larger spores and in having stictic acid.
Specimens Examined. — Gastle Bruce Road, 1000 ft. Hale 38177 (US); Gan-Dom logging area, Dleau Gommier, 1600- 1700 ft, Hale 35175 (US); Soufriere Ridge, 2200-3100 ft, Imshaug 33068 (MSG).
NUMBER 40
35
Figure 9. — Cross sections of apothecia of Graphina: a, G. colUculosa (lectotype of G. aibonitensis Fink); b, G. collospora (Boergesen s.n.); c, G. colurnbina {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
SMITHSONIAN CONTRIBUTIONS TO BOTANY
41. Graphina collospora Figure 9b; Plate 7/
Graphina collospora (\’ainio) Zahlbruckner, 1923:402.
Graphis collospora Vainio, 1915:153 [type collection: Mt.
Eagles, St. Croix, Boergeseii s.n. (TUR. lectotype)].
Description. — Thallus greenish gray, thick, con- tinuous, 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 trans- verse section; hymenium 90-100 ^m high, epi- hymenium brown, upper portion of hymenium and all of epihymenium 1+ blue. Exciple brown- black, barely closed to barely open, labia quite divergent. Spores 1/ascus, densely muriform, 25-37 X (80-) 105-130 ^m, with broad pale gelatinous outer layer, ca. 15 /xm thick, which frequently per- sists even after ejection from the ascus, l-f blue.
Chemistry. — Norstictic acid, stictic acid, constic- tic acid.
Habitat. — On Cocos at the shore.
Discussion. — Graphina collospora is part of the complex centering around G. confluens (Fee) Muel- ler 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.
Specimens Examined. — Ans du Me, sea level, Wirth 520 (US).
42. Graphina columbina, new combination Figure 9c; Plate 8a
Graphis columbina Tuckermaii, 1888:123 [type collection:
Alabama, USA, Beaumont s.n. (FFI, lectotype)].
Fissurina virginalis Tuckernian ex Nylander, 1889:50 [type
collection: Florida, Li^SA, s.n. (US, lectotype)].
Graphis virginalis Tuckerman ex Eckfeldt, 1890:256. Graphina virgiiialis (Tuckerman ex Nylander) Mueller Argo- viensis, 1895b:47.
Phaeographina columbina (Tuckerman) Zahlbruckner, 1923:
436.
Description. — 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 /xm high, I-; exciple yellowish, rudimentary, labia con- vergent to somewhat gaping, entire, containing many bark cells. Spores 6-8/ascus, irregularly muri- form, 1-3 X 3-6 locular, clear (!), (8-), 13-19 X 20-30 (-40) ixm, I-+- blue.
Chemistry. — Psoromic acid.
Habitat. — Lowland secondary growth.
Discussion. — The spores of G. columbina are clear in all stages, and hence Zahlbruckner’s trans- fer 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 com- pound in the Graphidaceae.
Speclmens Examined. — Roseau Botanic Garden, 100 ft, Wirth 446 (US).
43. Graphina dimidiata Figure 9d; Plate 8fc
Graphina dimidiata (Vainio) Zahlbruckner, 1923:404.
Graphis dimidiata \'ainio, 1890:108 [type collection: Brazil,
Vainio 322 (TUR, lectotype)].
Description. — lliallus grayish to off-white, con- 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 visi- ble in surface view, occasionally appearing as a lighter line between the labia. Ascocarp transverse section: hymenium 90-115 /xm high, epihymenium darkened, I-; exciple open below, lips entire, con- vergent 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 ^m, 1+ blue.
Chemistry — No substances present.
Habitat. — Rain forest, mossy forest.
Discussion. — Graphina dimidiata is distinct among tlie small-spored Crraphina 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 Examined. — 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: Brasil,
Malme 2038B (S, lectotype)].
Description. — Thallus 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 spec- imen 1-3 mm long, both rarely branched, straight to slightly curved, both frequently pale to reddish tan above. Disc not visible in surface view. Asco- carp transverse section: hymenium 70-100 ^um high in type, 130-150 /xm in the Dominican specimen, I-; exciple pale red-brown, closed, labia convergent, internally striate. Spores 6-8/ascus, muriform, 9-12 X 20-30 pxn, 1+ blue.
Chemistry. — Norstictic acid.
Habitat. — ^Virgin upland rain forest (Dominica only).
Discussion. — The illustration of the exciple that accompanies Redinger’s original description is in- correct, 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 dif- fers 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 chem- istry. Also similar is Graphina carneoviridis, which differs in having smaller spores, a paler more poorly developed exciple, and a different chemistry.
Specimen Examined.— Morne Diablotin, 2200-2600 ft. Hale 38090 (US).
45. Graphina frumentaria Figure 9/; Plate 8d
Graphina frumentaria (Fee) Mueller Argovieiisis, 1880:40. Graphis frumentaria Fee 1824:45 [type collection: Peru,
Mutis s.n. (G, lectotype)].
Description. — Thallus buff-gray, smooth to some- what 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 [xva high, I-; exciple yellow, occasionally red- brown 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 /xin, I-f blue.
Chemistry. — No substances present.
Habitat.^ — Virgin rain forest.
Discussion. — Graphma 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 incor- rect in that the specimen has labia with internal striae and the overall size is distinctly less than the type.
Specimen Examined. — Caii-Dom logging area, Newfound- land, 800 ft. Hale 35242 (US).
46. Graphina illinata, new combination Figure 9g; Plate 8e
Graphis illinata Eschweiler in Martins, 1833:82 [type collec- tion: Brazil, s.n. (M, lectotype)].
Df.scription. — Thallus slate gray to whitish, smooth, glossy, thick. A.scocarps very prottiberant, straight to occasionally flexuose, usually un- branched, covered completely by the thalline mar- gin, 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 (!) p,m high, 1-; exciple black, massively closed, labia more or less convergent, more or less entire, completely covered by the thal- line margin. Spores l/asctis, densely muriform, 30-50 X 110-225 pm, 1+ blue.
Chemistry. — No substances present (trace nor- stictic acid on one specimen).
Habitat. — Virgin upland rain forest.
Discussion. — The traditional concept of G. illi- nata, i.e., as an earlier name for Graphis anguilli- formis, 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 spe- cies was found in Dominica by only one of the four
38
SMITHSONIAN CONTRIBUTIONS TO BOTANY
collectors. The implication, of course, is that many other less conspicuous canopy species remain uncol- lected on the island.
Specimens Examined — Can-Dom logging area, Brantridgc Estate, 1700 ft, Hale 35276, 35281, 35284 (US); Can-Dom log- ging area, Pont Casse, 2000 ft. Hale 35082, 35083, 35124 (US).
47. Graphina incrustans Figure 9/;; Plate 8/
Graphina incrustans (Fee) Mueller Argoviensis 1887:47. Fissurina incrustans Fee, 1824:60 [type collection; South America, s.n. (G, lectotype)].
Graphis rubiginosa Fee, 1824:47 [type collection: South Amer- ica, s.n. (G, lectotype)].
Graphina rubiginosa (Fee) Mueller Argoviensis, 1887:44. Graphis glaucoderma Nylander ex Tuckerman, 1888:124 (type collection; Cuba, Wright 61 (FH, lectotype)].
Graphis dehiscens Vainio, 1890:111 [type collection: Brazil, Vainio 306 (TUR, lectotype)].
Fissurina nitidescens Nylander, 1890:108 [type collection:
Florida, USA, Calkins 31 (H, lectotype)].
Graphina glaucoderma (Nylander ex Tuckerman) Mueller Argoviensis 1895b:47.
Graphis nitidescens (Nylander) Vainio, 1915:151.
Graphina nitidescens (Nylander) Riddle, 1918:115.
Graphina dehiscens (Vainio) Redlnger, 1933:13.
Description. — Thallus yellow-brown to yellow- green, 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, occa- sionally branched, usually concolorous with the thallus, nearly round to quite elongate and flex- uose, 0.5-6 mm long. Disc exposed but too sunken to be easily visible in surface view. Ascocarp trans- verse section: hymenium (45-) 75-150 /rin high, I-, epihymenium slightly darkened; exciple rudimen- tary, yellow to pale red-brown, more or less open, labia quite convergent to quite spreading, apices of lips sometimes slightly darkened, with much in- cluded bark. Spores 8/ascus, 1-2 (-3) X 4-6 locular, 6-11 X 13-35 /xm, I- or 1+ 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. We have concentrated on Neo- tropic species, but many Paleotropic types will un- doubtedly 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.
.Specimens Examined. — Micotriii, 3000 ft, Wirth 465 (US); South Chiltern Estate, 1200-1500 ft, Irnshaug 32802 (MSG); Freshwater Lake, 2600-2877 ft, Irnshaug 32857A (MSG).
48. Graphina insculpta Figure 10a; Plate 9a
Graphina insculpta (Eschweiler in Martius) Mueller Argo- viensis, 1888b:507.
Diorygma insculpta Eschweiler in Martius, 1828:9 [type col- lection: Brazil, s.n. (M, lectotype)].
Graphis insculpta (Eschweiler in Martius) Nylander, 1863:371.
Description. — Thallus greenish to yellow-brown, smooth, glossy, 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 sec- tion: hymenium 90-120 /xm high, I-; exciple rudi- mentary, yellow to pale red-brown, open, labia convergent to spreading, with much included bark. Spores 1 / ascus, densely muriform, 20-30 X (50-) 70-75 (-100) /xm, I- or 1+ slow, reddish. Chemistry. — No substances present.
Habitat. — Rain forest.
Discu.ssion. — Graphina insculpta is morpholog- ically identical to G. incrustans and can be sep- arated only by having one very large spore per
ascus.
Specimens Examined. — Near Pont Cass^, 1400 ft. Hale 37819 (US).
49. Graphina macella Figure 10b; Plate 9b
Graphina macella (Krempelhuber) Mueller Argoviensis, 1880: 39.
NUMBER 40
39
Figure 10.— Cross sections of apothecia of Graphina: a, G. insculpta (lectotype in G); b, tnacella {Glaziou 6289B); c, G. marcescens (lectotype in G); d, G. nuda {Hale 35119); e, platyleuca {Thiebaut s.n.); f, G. plurispora (Malme 998); g, G. pseudoanaloga (Duss 1590); h, G. rufopallida (Duss 527).
O O
40
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Graphis macella Krempelhuber, 1876:380 [type collection: Brazil, Glaziou 6289b (M, lectotype)].
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 tlialline stripes, laterally covered by the thalline margin, 1-5 mm long, less than 0.5 mm wide. Disc not visible in surface view. Ascocarp transverse section: hymenium 140-160 p,m high, I-; exciple black laterally and above, fre- quently brown or irregularly carbonized below, closed to nearly open, labia convergent, striate. Spores 1 / ascus, densely muriform, (20-) 25-40 X (-65) 95-130 p,m, I-b blue.
Chemistry. — No substances present.
Habitat. — Rain forest (Dominica).
Discussion. — Graphina macella is similar to G. acharii but is smaller overall, more sunken, con- sistently has monosporous asci, and has an irregu- larly carbonized excipular base. In addition to the Brazilian and Dominican material, we have seen speciments from Mexico, indicating that this spe- cies will probably be found throughout the Neo- tropics.
Specimen Examined. — Valley of Desolation, 2800 ft, Hale 35723 (US).
50. Graphina marcescens Figure 10c; Plate 9c
Graphina marcescens (Fee) Mueller Argoviensis, 1887:42. Graphis marcescens Fee 1824:38 [type collection: South Amer- ica, Humboldt et Boiipland s.n. (G, lectotype)].
Graphis intricata Eschweiler in Martius, 1833:79 (non Graphis intricata Fee, 1824:42) [type collection: Brazil, Martius s.n. (M, lectotype)].
Graphina intricata (Eschweiler in Martius) Mueller Argo- viensis, 1888b:510.
Graphina pUttii Zahlbruckner, 1928:60 [type collection: Flor- ida, USA, Plitt s.n. (US, lectotype; L, isotype)].
Description. — Thallus off-white, continuous, matte to nearly powdery. Ascocarps more or less emergent, rarely branched, quite flexuose, com- pletely covered by a concolorous thalline margin which is easily dislodged, exposing the black ex- ciples below, 1-6 mm long, ca. 0.3 mm wide. Asco- carp transverse section: hymenium (60-) 75-120 (-140) ^m, high, I-; exciple black above, more or less 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 /jja, 1+ blue.
Chemistry. — Salazinic acid.
Habitat. — Upland rain forest (Dominica; see remarks).
Discussion. — 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 rare in the family, and the powdery thal- line covering, easily rubbed, completely covering the ascocarps. Externally, G. marcescens is identical to Graphis candidissima 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 ecolog- ically. The Dominican specimen (and probably the type of G. marcescens) is from high virgin rain forest; the type of G. plittii is from Florida, and hence probably lowland forest. In addition, we have seen a specimen from 2200 meters, collected in India (Hale 353 12A), which differs from the Neo- tropical material only in being somewhat larger and in having the exciple base a bit more carbon- ized.
Specimen Examined. — Can-Dom logging area, Dleau Gom- mier, 1600-1700 ft. Hale S4312A (US).
51. Graphina nuda Figure lOd; Plate 9d
Graphina Jiuda Magnusson, 1955:266 [type collection: Mauna
Kea, Hawaii, Faurie 1025b (UPS, holotype)].
Description. — Thallus gray to tan-gray, contin- uous, smooth to minutely roughened. Ascocarps very prominent, black, straight to slightly curved, unbranclied, without a thalline margin, resembling Graphis adpressa, 1-2 mm long, slender. Disc not visible in surface view. Ascocarp transverse section: hymenium 135-150 p,m high, I-; exciple black usually closed but occasionally irregular at the base, lips intact, convergent. Spores 8/asciis, uniseriate, 1-4 X 4 locular, 16-18 X 20-30 ^am, I -I- blue.
Chemistry. — No substances present.
Habitat. — Coastal Cocos to virgin upland rain forest; apparently of wide ecological amplitude.
Discussion.-— Grap/tma nuda is distinctive among
NUMBER 40
41
Dominican Graphina species in its short, Ope- grapha-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 (Fee) Mueller Argoviensis (1887), which is exter- nally and internally very similar, differs in having much smaller spores.
Specimens Examined. — Prince Rupert Bay, Irnshaug 33536A (MSC); Can-Dom logging area, Pont Casse, 2000 ft, Hale 35092, 35119 (US).
52. Graphina platyleuca Figure lOe; 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 (Esch- weiler in Martins) Zahlbruckner, 1902:385) [type collection: Puerto Rico, Fink 1774 (MICH, lectotype)].
Graphina platycarpina Zahlbruckner, 1932:214 (nomen novum for G. platycarpa Fink).
Description. — Thallus greenish white, continu- ous or occasionally cracked, matte to nearly pow- dery-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 fim, high, 1+ blue, epihymenium dark; exciple irregularly carbonized, quite black in parts, red- brown in other patches, closed, lips quite divergent, entire. Spores 1 / ascus, densely muriform, 35-^10 X 120-165 p,m, I-l- blue.
Chemistry. — Protocetraric acid only (types of G. platyleuca, G. platycarpa), or protocetraric acid with salazinic acid and norstictic acid (see discus- sion).
Habitat.- — ^Primary rain forest (Dominica). Discussion. — As presently recognized, G. platy- leuca is a chemically heterogeneous species. How- ever, protocetraric acid is quite rare in the Graphi- daceae 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 re- veal that G. platyleuca shows similar chemical vari- ation 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).
Specimen Examined. — Dleau Gommier Forest Reserve, 1200 ft. Hale 37955 (US).
53. Graphina plurispora, new combination Figure 10/; Plate 9/
Graphina pseuclosophistica var. plurispora Redinger, 1933:36
[type Collection: Brasil, Malme 998 (S, lectotype)].
Description. — Thallus continuous, gray-white, smooth and glossy to matte (on the same thallus). Ascocarps barely raised, black above, frecjuently with white stripes, with a well-developed thalline margin laterally, quite Ilexuose, 2-5 mm long, slen- der. Disc not visible in surface view. Ascocarp transverse section; hymenium 60-100 p,m high (on the same specimen), I-; exciple very variable (within one ascocarp), black, distinctly open to dis- tinctly closed, lips convergent, perfectly intact to cjuite striate. Spores begining as 8/ascus, but only (2-) 4-6 maturing, muriform, 1-3 X 8-13 locular, 10-15 X (27-) 35-50 (-60) /xin. Id- blue.
Chemistry. — No substances present.
Habitat. — Cultivated areas above 1500 feet (Dominica only).
Discussion. — Redinger’s variety differs from the type of Graphina pseudosophistica in two impor- tain I'espects. The latter has consistently mono- sporous asci and has the “quintaria” unknowns. Became of these differences, we have raised the variety to species rank. Still, Graphina plurispora is l)oth highly variable and part of a very confusing array of species. Most closely related are the fol- lowing
Graphina disserpens (Nylander) Mueller Argovi- ensis (1880) is tentatively separable because of the cjuite dendritically branched ascocarps and consist- ently open exciples.
Graphina elongatoradians Fink (1927) is another
42
SMITHSONIAN CONTRIBUTIONS TO BOTANY
dendritically branched relative with longer, more slender ascocarps, with open exciples, and a differ- ent chemistry (two unknown substances).
Graphiiia subvelata (Stirton) Zahlbruckner (1923) differs in being non-striate and in having open ex- ciples. See Hayward (1978) for a discussion of G. subvelata and its near relatives, including New World species.
In addition, G. plurispora bears a number of similarities to the G. antillarum complex; for a more complete discussion, see the latter species.
Specimens Examined. — South Cliiltcin Estate Road, 1500- 1700 ft, Imshaug 32724A, 32727 (MSC).
54. Graphina pseudoanaloga
Figure lOg; Plate 10a
Graphina pseudoanaloga (\'ainio) Zahlbruckner, 1923:421. Graphis pseudoanaloga Vaiuio, 1915:145 [type collection:
Guadeloupe, Duss 1590 (TUR, lectotype)].
Description, — Thallus gray-white, thin, continu- ous, 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 visible in surface view. Ascocarp transverse section: hymenium 75-115 jim high, in- spcrsed (apparently clear in Wirth 526), epihyme- nium 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 /cm, I-l- blue.
Chemistry. — Norstictic acid.
Habitat. — Beach to upland virgin rain forest; apparently of wide ecological amplitude.
Discussion. — Graphina pseudoanaloga is very- similar to G. analoga (Nylander) Zahlbruckner (1923), which differs in having smaller spores and an open exciple.
•Specimens Examined. — Cabrit, sea level, Wirth 526 (US); Caii-Dom logging area, Brantridge Estate, 1700 ft. Hale ,35283 (US).
55. Graphina rufopallida Figure 10/j; Plate 105
Graphina rufopallida (Vainio) Zahlbruckner, 1923:423 (“rufo-
pallens,” sphalm.).
Graphis rufopallida Vainio, 1915:149 [type collection: Guade-
loupe, Duss 527 (TUR, lectotype)].
Graphis vermicularis Vainio, 1915:148 [type collection: Shaw-
ford Estate, Dominica, Elliott 1852 (TUR, lectotype)]. Graphina vermicularis (Vainio) Zahlbruckner, 1923:430.
Description. — Thallus off-white, smooth, contin- uous. Ascocarps somewhat emergent, tan to red- brown, rarely branched, flexuose and intertwined, 1-7 mm long, slender. Disc not visible in surface view. Ascocarp transverse section: hymenium 70-100 i^m high, I-; exciple more or less open, labia convergent, more or less striate, dark red- brown or fuscescent apically (but not carbonized), paler yellow-brown below. Spores 6-8/immature ascus, apparently always aborting to 2-4/mature ascus, densely muriform, 15-30 X 36—75 /cm, 1 + blue.
Chemistry. — No substances present.
Habitat. — Upland rain forest.
Dkscussion. — Graphina rufopallida is most simi- lar to G. balbisii (see discussion under G. chloro- carpa), from which it differs in the lower, much longer, more slender and intricating ascocarps.
We have seen no specimens of this species other than the two types cited.
.Specimen Examined. — Shawford Estate, Elliott 1852 (TUR).
56. Graphina suberythrella, new species Figure 11a; Plate 10c
Description. — Thallus continuus, laevis, stratum corticale bene evolutum. Apothecia subimmersa, flexuosa, vulgo simplicia, 2-4 mm longa, disco lato, pruinoso. Excipulum integrum, flavidum vel rufo- ferrugineum, labiis integris, divergentibus. Hy- menium 110-140 /^m ahum, 1+ coeruleum. Asci 8 spori; sporae decolores, murales, loculis horizontali- bus 5-7, loculis transversis 1-3, 10-14 X 17-28 fxm, 1+ 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 /xm high, at least the epihymenium 1+ blue; exciple yellow 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 /xm, 1+ blue.
Chemistry. — Norstictic acid.
NUMBER 40
43
Figure 11. — Cross sections of apothecia of Graphina: a, G. suberythreUa {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); /, G. virginea (lectotype of G. obtectula, Tonduz).
Holotype.- — Cultivated area. South Chiltern Es- tate Road, Dominica, elevation 1500-1700 feet, Imshaug 32699 (MSC); isotype, US).
Discussion. — Graphina suberythreUa resembles
G. erythrella (Montagne) Zahlbruckner (1923), but differs in its more immersed ascocarps, gray disc, I- hymenium and larger spores. Also similar is 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 1 + hymenium.
Specimen Examined. — Can-Dom logging area, Newfound- land, 800 ft. Hale 35224 (US).
57. Graphina triphoroides, new species Figure lib; Plate lOd
Description. — Thallus continuus, farinosus. Apo- thecia alte sessilia, simplicia, margine thallino usque ad verticein vestita, 1-6 mm longa, 1 mm lata. Excipulum dimidiatum, fuligineum, labiis in- tegTis vel irregulariter crenulatis, erectis vel con- vergentibus; hymenium 250-700 fxm ahum, I-; Asci 6 (-8?) spoil; sporae decolores, murales, loculis hori- zontalibus 7-10, loculis transversis 2-4, 18-30 X 47-75 juin, 1+ coeruleae.
Thallus white to off-white, continuous, mealy. Ascocarps large, very protuberant, unbranched, covered to the apices by a mealy concolorous thal- line margin, irregularly blackened where the cov- ering is rubbed, 1-6 mm long, 1 mm wide. Disc usually not visible in surface view. Ascocarp trans- verse section: hymenium 250-700 (!) ^m high, I-; exciple black, more or less yellow-brown and open below, labia erect to convergent, entire to some- what irregular in outline. Spores 6 (-8?)/ ascus, muriform, 2-4 X 7-10 locular, 18-30 X 47-75 [xm, 1+ blue.
Chemistry. — Stictic acid.
Holotype. — Can-Dom logging area, Dleau Gom- mier, 1600-1700 ft, Hale 35161, January 1969 (US).
Discussion. — Graphina triphoroides is part of a very distinct group of three New World species, all characterized by very large, thalline-covered apo- thecia and dimidiate, carbonized exciples. Graphina triphora (Nylander) Mueller Argoviensis (1880) dif- fers in having 3-4 spores per ascus, no lichen acids, and carbonization restricted more to the upper por- tions of the labia. Graphina cleitops (Fee) Mueller Argoviensis (1887), the third species, differs in being monosporous.
Specimens Examined.- — Caii-Dom logging area, Dleau Gom- mier, 1600-1700 ft, Hale 35128 (US); Wright 15, Cuba (US) (“type” of Graphis triphoroides Nylander, unpublished name).
58. Graphina vestitoides Figure 11c; Plate lOe
Graphina vestitoides Fink, 1927:218 [type collection: Puerto
Rico, Fink 1986 (MICH, lectotype)].
Phaeographis cerviculata Redinger, 1935:76 [type collection:
Brazil, Malme 3531 (S, lectotype; UPS, isotype)].
Description. — Thallus gray, continuous, smooth. Ascocarps quite protuberant, frequently branched, ffexuose, varying from black above and partially covered by a thalline margin to concolorous (com- pletely covered), to 10 mm long, ca. 0.5 mm wide. Ascocarp transverse section: hymenium 150-210 fxm high, I-; exciple black, usually closed, lips conver- gent, striate to nearly entire. Spores 4-6 (-8)/ascus, biocellate only in the terminal few sections, 12-18 X 85-140 jxm, 1+ blue.
Chemistry. — No substances present.
Habitat. — Upland virgin and secondary forest.
Discu-SSIOn. — Graphina 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 cerviculata has mature spores that are quite clear, and the specimen is identical to G. vestitoides.
Specimens Examined. — Freshwater Lake, 2600-2877 ft, Ims- haug 32859A (MSC); near Pont Casse, 1400 ft, Wirth 544a (US).
59. Graphina virginea Figure lld,e,f; Plate 10/
Graphina virginea (Eschweiler m Martius) Mueller Argovien- sis, 1880:41.
Leiogramma virgineum E.schweiler in Martius, 1833:98 [type collection: Brazil, Martius s.n. (M, lectotyjre)].
Graphina obtectula Mueller Argoviensis, 1892b: 153 [type col- lection: Costa Rica, Pit tier 6166 (G, lectotype); Figure 11/)]. Graphina melaleuca Mueller Argoviensis, 1895c: 144 [type col- lection: Roseau, Dominica, Eckfeldt 70 (G, lectotype)]. Graphina palmeri Zahlbrtickner, 1921:231 [type collection;
Mexico, Pringle 9 (MICH, isotype)].
Graphina triangularis Zahlbruckner 1921:232 [type collection:
Mexico, Pringle 17 (MICH, isotype)].
Graphis collosporella \’alnio, 1923:141 [type collection: Trini- dad, Thaxter 10 (TUR, lectotype)].
Graphina collosporella (Vainio) Zahlbruckner, 1923:212.
De.scription. — Thalltis greenish white to gray, thick, continuous except for a fissure usually par- alleling each ascocarp, smooth to mealy, occasion-
NUMBER 40
45
ally quite rugose (when ascocarps are densely clus- tered). Ascocarps nearly flush with the thallus, rarely somewhat raised, margins concolorous 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 sec- tion: hymenium 120-200 /xm high, epihymenium darkened, at least the epihymenium 1+ blue. Exciple yellowish, rudimentary, sometimes thick- ened below and/or with a basal dark (nearly car- bonized) band (see discussion), labia more or less divergent, occasionally with internal striae (see dis- cussion) but otherwise entire. Spores 2-3, 4-6 or 8/ascus (!), most numbers occurring within a sin- gle thallus, 1-4 (-7) X 11-20 locular, (12-) 15-25 X 50-80 (-130) /xm, 1+ blue.
Chemistry. — At least one of the 3 so-called “Quintaria” unknowns; trace of stictic acid and constictic acid (type of G. virginea only).
Habitat. — Secondary growth and cultivated trees and shrubs, from sea level to over 1000 meters.
Discussion. — In its usual form Graphina virginea 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 mar- gin. The ascocarps themselves are commonly 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 (Hibis- cus).
Internally, this most common form of the species has a rather rudimentary exciple and a very varia- ble 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
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 asco- carps somewhat emergent. It seems quite likely that this appearance comes with age; dense cluster- ing of more typical ascocarps, with their associated fissures, could easily produce such a form. A prob- able additional synonym for this “rumpled” vari- ant of G. virginea is G. monophora (Nylander) Zahlbruckner (1923).
The type of G. melaleuca also has a broader, browner disc than is normally found in G. virginea; externally this specimen is indistinguishable from G. rnendax (Nylander) Mueller- Argoviensis (1888a). 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.
Graphina palmeri 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. virginea, 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 1+ blue reaction of (at least) the epihyraen- ium; all share at least two of the three “Quintaria” acids.
Specimens Examined. — Brookhill Estate, 100-150 ft, Ims- haiig 33321A (MSC); Cocoa Centre on Layon River, 200 ft, Wirth 517 (US); north of Pont Cass^ 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. difformis
2. Exciples carbonized only at labial tips.
3. Ascocarps massive, 1-3 mm wide; no substances present 61. P. caesiopridnosa
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 p.m long 64. P. elliottii
4. Labia divergent, exposing the disc; spores less than 40 p,m long.
5. Ascocarps prominently raised, commonly over 5 mm long; norstictic acid present
60. P. atrovermicvdaris
5. .Ascocarps barely raised, less than 5 mm long; norstictic acid absent.
6. Spores 20-27 ./im long; stictic acid present 65. P. oscitans
6. Spores 33-35 jiva. long; no substances present 62. P. coriaria
60. Phaeographina atrovermicularis, new species Figure 12a; Plate 11a
Description. — Thallus continuus, sublaevigatus. Apothecia adpresso-sessilia, vulgo ramulosa, 3-10 mm longa, disco dilatato, nigro. Excipulum dimi- diatum, fuscum, labiis integris, divergentibus; hy- menium 145-220 p,m altum, I-. Asci 6-8 spori; sporae fuscescentes, murales, loculis horizontalibus 6-9, loculis transversis 1-4, 15-17 X 25-35 pm, 1+ 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, occa- sionally faintly pruinose. Ascocarp transverse sec- tion: hymenium 145-220 pm high, I-, lightly beaded- inspersed; exciple open below, laterally brownish- black, 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-f blue.
Chemistry. — Norstictic acid.
Holotype.^ — Can-Dom logging area, Brantridge Estate, Dominica, elevation 1700 feet. Hale 35278 (US).
Discussion. — By virtue of its long black asco- carps, Phaeographina atrovermicularis is quite dis- tinct and is apparently not closely related to any other New World species. Most similar is P. inter- ceclens Mueller Argoviensis (1888e), 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. caesioprui- nosa, which has much shorter, broader ascocarps, strongly pruinose discs, and lacks norstictic acid.
Specimens Examined. — Can-Dom logging area, Pont Casse, 2000 ft. Hale 35093 (US); rain forest, Jean Estate, 2000 ft. Hale 35068 (US).
61. Phaeographina caesiopruinosa Figure 12&; Plate life
Phaeographina caesiopruinosa (F6e) Mueller Argoviensis,
1887:49.
Arthonia caesiopruinosa Fte, 1837:36 [type collection: South
America, s.n. (G, lectotype)].
Description. — Thallus pale brown, smooth to slightly roughened, continuous. Ascocarps quite ele- vated, rarely branched, straight to ffexuose, 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 sec- tion: hymenium 150-200 pm high, I-, epihymen- ium darkened; exciple black laterally, open below, lips divergent. Spores (4-) 6-8/ascus, densely muri- form, 12-17 X 45-75 (-100) pm, 1+ 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.
Specimen Examined.^ — -South Chiltern Estate, 1500-1700 ft, Irnshaug 32731 (MSC).
62. Phaeographina coriaria^ new species Figure 12c; Plate 11c
Description. — Thallus continuus, laevis, coria- ceus. 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 in- tegris, 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. difformis {Fink 1874); e, P. elliottii {Elliott 1338); f, P. oscitans {Mann s.n.).
rales, loculis horizontalibus 8-10, loculis transver- sis 1-3, 9-10 X 33-35 //.m, 1+ rufo-fuscae.
Thallus continuous, smooth, tan, resembling velum. Ascocarps slightly raised, straight to some-
what curved, rarely branched, 1-5 mm long, ca. 0.3 mm wide. Disc broad, black, with a light pru- ina. Ascocarp transverse section: hymenium ca. 150 lim high, beaded, epihymenium dark, I-; exciple
48
SMITHSONIAN CONTRIBUTIONS TO BOTANY
rudimentary to very pale below, laterally red-brown, lips intact, divergent. Spores 4-6 (-8?)/ascus, brown, 8-10 X 1-3 locular, 9-10 X 33-35 /xm, 1+ red, slowly darkening.
Chemistry. — No substances present.
Holotype. — Mossy forest, Morne Anglais, Do- minica, elevation 3000-3600 feet. Hale 35378 (US).
Discussion. — Phaeographina coriaria is most sim- ilar to the following three New World species:
Phaeographina arechavaletae Mueller Argoviensis (1888c) 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 Wirth and Hale (1963) differs in chemistry and in having monosporous asci.
63. Phaeographina cf. difformis Figure 12d; Plate llrf
Phaeographina difformis Fink, 1927:220 [type collection:
Aibonito, Puerto Rico, Pink 1874 (MICH, lectotype)].
Description. — Thallus thin, continuous, glossy, greenish gray. Ascocarps nearly round to elongate and irregular, ends round, occasionally shortly branched, partly immersed, 0.8-1. 5 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 p,m high, epihymenium very dark, I-; exciple brown- black, 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 /xin in the type, 18-22 X 55-75 in the Dominican specimen, 1+ blue.
Chemistry. — Unknown in the type, no sub- stances present in the Dominican specimen.
Habitat. — Elfin forest (Dominican specimen only).
Discussion. — Although the Dominican specimen is externally identical to the type, and is indistin- guishable in cross-section, the spore size and num- ber make this identification tentative. In addition, it should be noted that both specimens have large
numbers of clear spores, making this species diffi- cult to assign to genus. In fact, this species may represent a link to the smaller members of the Graphina confluens alliance, such as G. cymbe- grapha (Nylander) Zahlbruckner (1923), which is strikingly similar.
Very similar to P. difformis is P. explicans Fink ex Hedrick (1933), which differs in its more elon- gate, rather sunken apothecia.
Specimen Examined. — Summit of Morne Diablotin, 4300- 4550 ft, Imshaug 32913A (MSC).
64. Phaeographina elliottii
Figure 12c; Plate lie
Phaeographina elliottii (Vainio) Zahlbruckner, 1923:438. Graphis elliottii Vainio, 1915:143 [type collection: Dominica,
Roseau Valley, Elliott 1338 (TUR, lectotype)].
Description. — Thallus thin, whitish, matte, con- tinuous. Ascocarps slightly raised, rarely branched, straight to somewhat curved, thalline margin promi- nent, to 1.5 mm. long, slender. Disc concealed to open and dark gray. Ascocarp transverse section: hymenium completely (?) disintegrated in the type, 1+ fide Vainio; exciple yellow-red, not well devel- oped, lips convergent to quite spreading, tips brown, dark portions probably derived (at least in part) by lateral compression of the dark epihy- menium. Spores (fide Vainio) 2-8/ascus, muriform, 22-28 X 44-54 ^am.
Chemistry. — -No substances present.
Habitat. — “On trees in Roseau Valley.”
Discussion. — 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 referable to this Phaeographina species
65. Phaeographina oscitans
Figure 12/; Plate 11/
Phaeographina oscitans (Tuckerman) Zahlbruckner, 1923:442. Graphis oscitans Tuckerman, 1868:231 [type collection: Oahu,
Hawaii, Mann 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 125-150 lj.m high, I-; exciple rudimentary below, yellow lat- erally, 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 jam, 1+ blue.
Chemistry. — Stictic acid.
Habitat. — -Virgin upland rain forest (Dominica only).
Discussion. — Phaeographina 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) Mueller Aigoviensis (1887), which would seem to be the TLC negative equivalent of P. pachnodes.
Specimens Examined. — Can-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 Graphis anguilliformis for Elliott s. n.
Graphis (Phaeographis) diversa: See Phaeographis exaltata for Elliott 526.
Graphis duplicata: See Graphis rimulosa for Elliott 125 and Graphis flexibilis for Elliott 1954 pro parte.
Graphis duplicata var. subduplicata Vainio: Elliott 1509 is sterile.
Graphis lactea: Elliott 535 is the type of G. lactea var. dominicana \'ainio; see Graphis tachygrapha.
Graphis lactea \ar. clausa: See Graphis insidiosa for Elliott
1511.
Graphis lineola: See Graphis leptocarpa for Elliott 1535.
Graphis (Phaeographis) lobata: The only Dominican collec- tion (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 Phaeotrema in the Thelotreinataceae.
Graphis (Phaeographis) inedusaeforinis: Elliott 529 in BM is a fragmentary specimen, which may be an immature Phaeographis exaltata. There is no specimen so identified in TUR.
Graphis (Graphina) subnitida: Elliott 171 is indeterminable.
Graphis tenella var. epiphaea Vainio: Not found in TUR or BM and presumed to be lost or misfiled.
Graphis (Graphina) verrnicularis: See Graphina rufopallida for Elliott 1852.
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NUMBER 40
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Index
(Synonyms in italics)
Arthonia caesiopruinosa, 46 Diorygma insculpta, 38 Fissurina incruslans, 38 insidiosa, 16 nitidescens, 38 virginal is, 36 Graphina acharii, 31 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 inlricata, 40 inturgescens, 31 macella, 38
marcescens, 40 melaleuca, 44 monophora, 45 nitidescens, 38 nuda, 40 obtectula, 44 palmeri, 44 parilis, 33 platycarpa, 41 platycarpina, 41 platycarpoides, 32 platyleuca, 41 plittii, 40 plurispora, 41 pseudoanaloga, 42 pseudosophistica var. plurispora, riopiedrensis, 34 rufopallida, 42 ruiziana, 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
columbina, 36
congesta, 15, 20
cooperta, 12, 49
dehiscens, 38
desquamescens, 12
dimidiata, 36
diversa, 49
dumastii, 24
dumastioides, 12
duplicata, 21, 49
duplicata var. subduplicata, 49
dussii, 13
elegans, 13
elliottii, 48
endoxantha, 21
eugeniae, 34
flavicans, 19
flexibilis, 13
frumentaria, 37
furcata, 19
glaucescens, 15
glaucoderma, 38
glauconigra, 20
grammitis, 15
haematites, 27
hum ills, 16, 24
illinata, 37
imshaugii, 16
inquinata, 24
insculpta, 38
insidiosa, 16
interversa, 6
intortula, 34
intricata, 12, 40
inturgescens, 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 platycarpoides, 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 tirnida, 23 tongloensis, 25 triticea, 23 tuniidula, 12 tumidulella, 15 turgidula, 24 vermicularis, 42, 49 virginalis, 36
Lecanactis exaltata, 27
Leiogramma virgineum, 44
Opegrapha desquamescens, 12 elegans, 13 rhizocola, 12 rimulosa, 21
Phaeographina arechavaletae, 48 atrovermicularis, 46 caesiopruinosa, 46 caracasana, 48 chrysocarpa, 28
columbina, 36 coriaria, 46 difformis, 48 elliottii, 48 elliptica, 48 explicans, 48 includens, 18 intercedens, 46 oscitans, 48 pachnodes, 49 turgida, 49
Phaeographis albida, 25 ceiviculata, 44 cinnabarina, 28 decipiens, 27 dimorpha, 29 dussii, 13 exaltata, 27 haematites, 27 longula, 19 mordenii, 28 rosea, 28 sexloculata, 27 subtigrina, 29
Sclerophyton colliculosum, 34
54
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Plate 1. — Species of Graphis: a, G. adpressa {Hale 35487); b, G. afzelii {Imshaug 33489A): c, G. anfractuosa (Wirth 538); d, G. anguillifonnis {Hale 35282); e, G. desquamescens {Hale 35449); /, G. dumastioides {Hale 37722). (Plates 1-10 X 10.)
NUMBER 40
55
Plate 2. Species of Graphis: a, G. dussii (Hale 35443); b, G. elegans (Hale 35476); c, G. flexibi- Its (Imshaug 33360); d, G. glaucescens (Imshaug 33313A); e, G. gramrnitis (Hale 38005); /, G. humilis (Hale 35698).
56
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Plate 3. — Species of Graphis: a, G. imshnugii (Imshaiig 32724 in MSC); b, G. insidiosa {Hale 35313); c, G. isidiifera {Hale 35179); d, G. leptocarpa {Imshaug 33109A); e, G. librata {Wirth 538); /. G. Ion gill a {Hale 35225).
NUMBER 40
57
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); G. subnitidula {Hale Sldld).
58
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Plate 5— Species of Grapliis and Phaeographis: a, G. tachygrapha (Imshaug 32862A); b, G. triticea (Hale 35324); c, G. turgidula {Wirlh 544b); d, P. albida {Hale 35453); e, P. arthonioides {Imshaug 33128); f, P. exaltata {Hale 35301).
NUMBER 40
59
Plate 6. — 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); /, G. acharii {Hale 35555).
60
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Pi.ATE 7. — Species of Graphiua: a, G. adscribens (Wirth 519); b, G. antillariim {Hale 35745); c, G. carneoviridis {Hale 37642): d, G. chlorocarpa {Hale 35129); e, G. colliculosa {Hale 35175); f, G. collospora {Wirth 520).
NUMBER 40
61
Plate 8. — Species of Graphina; a, G. columbina (Wirth 44G); b, G. dimidiata {Hale 35697): c, G. dispersa {Hale 38090); d, G. jrumentaria {Hale 35242); e, G. ilUnata {Hale 35284); /, G. incrustans {Wirth 465).
62
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Plate 9. — Species of Graphina: a, G. insciilpta {Hale 37819); b, G. macella {Hale 35723); c, G. marcescens {Hale 35312); d, G. nuda {Hale 35119); e, G. platyleuca {Hale 37955); f, G. plurispora {Imshaug 37724A).
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63
Plate 10. — Species of Graphina: a, G. pseiidoanaloga {Hale 35283); b, G. rufopallida {Elliott 1852); c, G. suberythrella {Imshaug 32699); d, G. triphoroides {Hale 35161); e, G. vestitoides {Imshaug 32859A); f, G. virgmea {Wirth 547).
64
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Plate 11. — Species of Phaeographina: a, P. airovermicularis {Hale 35278); b, P. caesiopruinosa {Imshaug 32713); c, P. coriaria {Hale 35S18); d, P. difformis {Imsltaug 32913A); e, P. elUottii {Elliott 1338 in TUR); f, P. oscitans {Hale 35131).
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