S M I T H S O N I A N C O N T R I B U T I O N S T O B O T A N Y 0 N U M B E R 4 0 Morden-Smithsonian Expedition to Dominica: The Lichens (Graphidaceae) Michael Wirth and Mason E. Hale, Jr. SMITHSONIAN INSTITUTION PRESS City of Washington 1978 A B S T R A C T LVii-th, R l . , and Mason E. Hale, Jr. ~Iortlen-Smitlisonian Expedition to Dominica: The Lichens (Graphidaceae). Smithsonian Contribulions to Botany, number 40, 64 pages, 23 figures, 1978.-The four main genera of the Graphidaceae in Dominica, Gruphis (27 species), Phaeogiaphis (7 species), Graplzina (25 species), and Plraeographina (6 species), are treated monographically with full synonymy, descriptions, chemistry, and illustrations. The family as a whole is common in all primary and secondary forests and at all elevations. Eight new species are de scr i I] etl : Graph i i i I 1 \ h a 11 g i i , G . isid i ife~ci , Plici <: og~rr f, Ii is ? r i oitle n i i, G YN p h i I I a cmrneoviridis, G. sziberythrella, G. triphoroides, Phaeogiaphitza utrovermicularis, and P. coriaria. Three new combinations, Gmphina coliinzbina (Tuckerman) Wirth and Hale, G. i l l inata (Eschweiler) Wirth and Hale, and G. plurispora (Redinger) TYirth and Hale, are also made. OFFICIAL P L L ~ L I C A T I O A DATE is handstam ed in a limited number of initial copies and is recorded in the Institution?s anilual report, SnzitEsoriian l.ear. SERIFS COLEK DESIGS: Leaf clearing from the Katsura tree Cerr id iphJ l lum japonicum Siebold and Zuccarini. Libraiy of Congress Cataloging in Publication Data LVirth, Michael. Xlorden-Smithsonian Expedition to Dominica: the Lichens (Graphidaceae). (Smithsonian contributions to botany ; no. 40) Bibliography: p. Includes index. Supt. of Docs. no.: SI 1.29:40 1, Graphidaceae. 2. Lichen-Dominica. 3. Xlorden-Smithsonian Expedition to Dominica I . Hale, Mason E. joint author. 11. Title. 111. Series: Smithsonian Institution. Smithsonian contributions to botany ; no. 40) QWS2747 no. 40 [QK585.GS] 581?.08s [389?.1] 77-608313 Contents Page Introduction 1 Generic Delimitation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Species Delimitation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Spores 3 Hymenium 3 Exciple 3 Thallus 5 Chemistry 5 Ecology 6 6 Key to the Genera of the Graphidacea nica 8 Key to the Species of Graphis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 25 45 Incorrect and Omitted Names . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Evolution in the Graphidaceae . . . . . Key to the Species of Phaeographis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key to the Species of Graphina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 Key to the Species of Phaeographina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Literature Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... 111 Morden-Smi thsonian Expedition to Dominica: The Lichens (Graphidaceae) Michael Wirth and Mason E. Hale, 3 r . 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 hlrs. William J. Morden, who generously supported field and herbarium studies by Hale. Dr. Henry Imshaug very kindly placed his collections at our disposal. We are also indebted to Dr. Minoru Nakanishi for the use of his unpublished data and keys for many of the Asian Michael Wir th , 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, Phaeographis, Graphina, Phaeo- graphina), four parallel genera with such tissue (Glyph is, Sarcographa, Medusu lina, Sarcograph ina), and a number of minor genera whose relationships are unclear (Helminthocarpon, Aulaxina, Gyro- stomum, etc.). Although it is customary to refer to these major genera as ascohymenial, even that seems to be in doubt (cf. Groenhart, 1965:4). We have, in any case, continued the ascohymenial terminology 1 2 usually applied to the family, using the terms ?hy- menium? and ?paraphyses? without necessarily accepting them a5 ontogenetically correct. T o a great extent, all genera ( in phanerogams as well as in lichens) are artificial coiistructs, in that no fixed criteria exist for delimiting any taxa above the biological species. In groups Ivhere cytogenetics, breeding experiments, etc., can reveal something of the direction of evolution, genera may in fact be an approximation of biological (i.e. evolutionary) 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 different interpre- tations: for example: TVliat exactly is a hymeniuni? Is the structure of the ascus of great importance, or little?-and so on. In the light of such uncertainty, a reasonable approach seems to be the recognition of genera on practical grounds, so that at least problems of identification and curating are more manageable. On such grounds, the major genera of the Graphi- daceae are reasonably delimited. By using $pore septation and color as primary criteria, and the development of stroma-like tissue as a secondary criterion, one can distinguish the major genera of the family with relative ease. T o be sure, specimens (and species) exist that bridge the gap betlveen these form genera and tend to blur the distinctions. In Graphina IJeStilOideS, for example, only the end locules of otlienvise typi- cal Graphis spores become longitudinally divided. Many of the very small-spored Gmpiiina species (G. insigni.r, G. colliculosa, etc.) have barely muri- form spores which, wlien not quite mature, will resemble four-loculed Gmpli is spores. The prob- lems of spore color and septation in such border- line species as Graphinn achasii and Pl7acogixpliina chi.ywcarpa have been mentioned elsewhere (TVirth and Hale, 1963). Intermediate states also exist between the sti-om- atoid and nonstromatoid genera. A fine example can be seen in many specimens of Phacogmphis exal tata, which frequently show, in portions of a single thallus, characteristic? of both Phucogi-aphis and Sawographa. There i3, however, a more serious objection to spore genera. One can regard a genus as consisting of species that share a large number of character- states in common; this approach, common among SUITHSONIAN COKTRIBCTIOSS T O BOTASY ?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 quite 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. TZ?e will call sucli forms ?sporomorphs,? defined as follows: trvo or inore species that are extremely similar (or identical) in external morphology, anatomy, cheni- 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 septatioii and /or spore color only. Sporomorphs are common in the Graphidaceae; good examples occurring at least in part in Domi- nica are Pliueogmph is art h on ioide.5 Graph i.r h ii m il i s and Gvuphina incrzi.rtan.r :I G1-aphi.r dunzastii. The exi5tence of iporomorphs casts considerable doubt on the ability of spore features to establish genera of closely related (i.e. very similar over-all) species. Is i t 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 Gwphis-like juvenile stage), then Gmphina incwstans may be n recent descendant of Graphis clunrastii, and these two species should be regarded as closely related. Placing these species in different genera, while grouping G. inci?tistans with otherwise very dis- similar Gmphina species, would then be illogical. If this objection to spore-based genera is accepted, then one is faced with a considerable problem of nomenclature. Ignoring the spore coIor and 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 would be to reduce all the spore-based genera back into Gmphi .~ . Such a system, which may be biologically quite acceptable, would raise immense practical problems; there are simply too many species of Grap1iicl;iceae (over one thousand) to be handled conveniently. NUMBER 40 3 In light of these practical problems, we have chosen to accept the major spore-based genera for the Graphidaceae. Whenever possible, however, we have indicated similar species and sporomorphs, 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 species. In a fam- ily with such a large range of spore size (6 pm to 300 pm), it is not surprising to find discontinuities of size classes. We have, whenever possible, 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 Phaeogmphis) where the main, and perhaps only, distinction consists of four versus more than four locules per spore. HYMExIuM.-Although hymenial character-states supply much useful information, we have seen very few cases where species are established primarily on hymenial features, One such feature is the presence of droplets or inclusions in the hymenium; i t 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:Z). Many of these characters now appear to be quite variable and unreliable. In the discussion below, it should be emphasized that all observations were made from very thin hand sections, mounted in Hoyer?s medium, and briefly boiled to drive out air. Without this approach, it is very difficult to determine accurately excipular structure. Occasionally, three to four days must elapse after mounting a section before all minute air bubbles disappear; when present, they may mimic carbonization very closely, particularly in the labial tips of totally uncarbonized exciples. Excip2e Curbonmztion: Carboniiation i s 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 SMITHSOXIAN CONTRIBUTIONS T O BOTASY quently in G. uirginea and G . confluens. We have found no correlation with any other features; in fact, the black band may be quite absent in parts of a thallus and present elsewhere. Many species of Graphidaceae lack carbonization completely. I n 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; i n many of these, what appear to be excipular labia are, in fact, masses of bark cells. I t may well be that presence of large amounts of bark laterally bounding the hymenium is signifi- cant as an indicator of endophloeodal development of the ascocarp. Open us. Closed Exciples: In many specimens with carbonized exciples, all sections from a thallus will be consistently open (or consistently closed) below, but many others will vary from completely open to completely closed within the same thallus. T h e weight commonly placed on this character-state to establish species, subgenera, or sections is un- justified. \\?hen 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. I n 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 i s 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- b i l is) . 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. O n 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. Gmphina 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 8f,g; 9a) 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 8f ,g; 9a), ascocarps with convergent labia (no disc visible) and divergent labia (prominent disc visible) may occur on the same thallus. If the two forms occur on different specimens, the surface 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 React ion: 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 S\IITHSONIAN COKTRIBUTIOSS T O BOTASY protocetraric, salazinic acids, and lichexanthone. A few species (Phacograpliis haetnatiics, P. cinnaba- rina (Fee) Mueller Argoviensis, Phneograplzina rliyysornq!m (Kacldi) Reclinger, among others) ha1.e pigments (anthraquinones?). All specimens were tested with normal thin-laTer chromatographic techniques using silica gel alum- inum-backed LIerck precoated plates. Two solvent systems were always used: hexane-ether-formic acid and benzene-dioxane-acetic acid. l?isualization was accomplished with 10 yo sulfuric acid and heating at 110? C. A l i s t of the chemicals found and species that contain them is given belorv. It should be assumed that the chemistry for each species represents that of the type material, unless otherwise indicated, and all other specimens cited. Lecanoric acid: G~-nph i s nfzelii Norstictic acid: Graphis desquninescens, G. dussii, G. elegnns, G. isidiifera, G. librntn, G. lumbricina, Phaegraphis s u h - /i, 1, i ~i ii n 1 r o i ? Y J i r i ic i i lii > i F Sorst ic t ic acid with stictic and constictic acitlr: Grophiiin I?rotoce t r:ii-ic x i t l : G ! nplti i i i t plnty l e i t cn Psol-omir acitl: (;rnphi,iir roi i i?izbi, in ?Quintaria? unknowns: Phaeogrnphis exnltatn and Grnphina t?irginea Salazitiic acid: Gi-ofiitiJra rolliriilosct a i id G. m n ~ r e s c r n s : G . plot! lezicn (accessory) Stictic acid: Grnphis iiiitJinstioitlrc, G. iiii\ho!(gii. G. le f i to- carpa, G. iriticea, G. turgiduln, Grnphina triphora, G. iJirgiJirn, and Phneogrnphinn oscitans T L C negative (no phenolic substances present): Grnphis n il f i V P T sit, G , / I ) I f i nc t i i o \ ( I , C; . n ~i g i i ilii fo i-iii ic , G . / I C Y i b ili c, G. glnitcescens, G. grnnimitis, G. huinilis, G. insirliosn, G. Zonguln, G. olivncen, G. rigiduln, G. rimulosn, G. siibele- ~ O J ~ F , G. sitbnitiduia, G. tnchygrnphn, Phneogrnphis nlhirln, P . nrthoiiioides, P . hneniatites, P . mordenii, P. rosen, Grnphi i ia nclinrii . G. iiiltcrib~iic, G . cnrnroi,iritlii, G. chlo- roenrfin, G. dimidinta, G. frumentnrin, G. illfiiotn, G. i n - criistn)is, G. inscnlpfn, G. innrclln, G. nzrdo, G. plziriiporn, G. rujopzilidn, G. zwstitoides, Phneogrnphiiiu cnesiopnii- nosn, P . corinrin, P . difformis, P . eiliottii, and (in part) P . e x n l t n t n Pigments: Phneographis hnernatitey Ecology T h e famiIy Graphidaceae is one of the largest groups in the lichens with over 1100 published names. One should not be surprised, therefore, to find that the species have become adapted to a wide range of habitats and often, as in the case of Graphis afzclii, seem to have a wide ecological 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, 1954) and other tropical areas. TVe 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 fore5t generally have the greatest number of species. Of the commoner species the following, for example, appear to be more or less confined to well developed rain forests: Graphis oliuacea (also rarely found in elfin forest), Graphinn chlorocarpa, G. colliczilosa, and G. illinata. These species are com- mon in and probably restricted to elfin or mossy forest: Graph i.s ndpl-e.r.~n, G. n ngzi illiforvn i F, G. /zim byicina, G. tachygrapha, G . triticen, Phaeo- qrnpliic c ~ x t i l / n i i i , and P . mordr77ii. These occur chiefly in low elevation, disturbed habitats: Graph i s derqzramescen.c, G m p h i n a ant i l larum, and G. uir- g i n f a . 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., nionophyletic. Second, since data froin genetics and cytology are still lacking, one inlist assume that a t least some phylogeny can lie inferred froin ontogeny. .\lchough the pitfalls and iallacies of using on- togen? a s ;I source for evolutionary speculation are !$-ell known. there seems little choice in the Graph- itlaceae. Specifically, stages of spore development m a y supply the only concrete bases yet available. T h e follon-ing four observations on spore ontog- eny should be considered: 1 . All Graphid spores start as a one-loculed (one- celled) structure, progress to two locules, then to four (rarely three) uniseriate chambers, then to multi loculx 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 Gsaphina and Phaeographina. Four-locular spores always occur as a group of eight per ascus; most spores with only transverse septae also occur as a group of eight per ascus. 4a. Pigmentation of spores occurs last, after spores reach essentially full size and septation. 4b. Pigmentation is accomplished by thickening of walls in all (?) Phaeographis, but in very few Phaeographina species. Those Phaeographina 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 Graphis-like spores. 3. Monosporous forms are derived from ancestors with plurisporous asci. 4a. Pigmentation may occur at any stage of spore septation, but usually represents the last event in spore development. Therefore sporomorphs which differ only in pigmentation are extremely closely related (i.e., one ?step? apart). 4b. As most Phaeographina species do not have thick-walled spores, they are probably derived from Graphina sporomorphs by added pigmentation, rather than from Phaeographis sporomorphs by added septation. The less common thick-walled Phaeogsaphina species may be derived from irregu- larly-loculed Graphina ancestors (also rather un- common) or from Phaeographis-like forms. Putting these speculations together in graphic form, we propose a very hypothetical scheme of evolutionary relationships (Figure 2). ................................................ GRAPHIS ....... ..... . . a * * .* . * : A n c e s t r a l s t o c k ..? spores 2 - Iocu la r , .: c ~ e a r f f e w extant i G . tu rbu len ta , e tc . ) species, G. cor iacea, .................. *. few- l o c u l e d t h i c k - wa I led monosporous .: ...... Phaeographina .... .... PHAEOGRAPH I NA .................. FIGURE 2.-Hypothetical origin and relationships of the genera of the Graphidaceae. 8 This scheme immediately gives rise to certain questions, problems, and objections, to wit: Where does carbonization fit in this scheme? Graphis spe- cies with few-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 SMITHSONIAN CONTRIBUTIONS T O BOTANY spore states? For example: a fissurine Gjaphzs would produce a maximum of four germination tubes per spore and hence have a low number of tubes in each of many landing sites. A 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- tile advantage associated with the variants. In the Ryrsonima scrub of western Mexico, for example, all the spore forms occur abundantly, shoulder to shoulder. HoweLer, this may be a very equable habitat, comparable to the tropical rain forest for the angiosperms; other more demanding habitats and substrates might re\ eal significant preferences and correlations, In addition, the abundance of 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- 1s. epiphloeodal thalli, etc. Clearly, much field work remains to be done. Key to the Genera of Graphidaceae in Dominica (Although all genera of the Graphidaceae that are known to occur in Dominica are keyed belois, the txvo stromatoid groups, Glyphis and Snrcogrnphn, are not treated in the text. Each is represented by only one species (l?ainio, 1896, 1915.) 1. Ascocarps embedded in prominently raised, stromatoid tissue. 2. 1. Ascocarps not embedded in raised stromatoid tissue. 3. Spores clear and hyaline, the lvalls usually thin. 4. Spores with transverse septae only ................................................................................ G r u ~ ~ ~ s 4 . Spores with transverse and longitudinal septae ...................................................... Graphina 3. Spores brown, the !calls occasionally thickened. 5 . Spores with transverse and longitudinal septae, the Isalls rarely thickened .......................... 5 . Spores with transverse and longitudinal septate, the walls rarely thickened .... 2. Spores clear, with transverse septae only .......... .......................... Glyphis Spores brown, with transverse septae only ................................................................ Sarcograph~~ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Phaeographis ....................................................................................................................... .Phaeographina Key to the Species of Graphis 1. Spores always four locular. 2. .4scocarps fissurine, arising as a thalline crack; disc usually concealed or sunken. 3. Exciple carbonized. 4. Exciple heavily carhoniied laterally, spores I + ........................................ I? . G. hurnilis NUMBER 40 9 nized only at labial tips, spores I- (reddish) ...................... ................................................. 25. G* tachYgraPha ................................... ...... 11. G. gramrnitis .... 3. Exciple totally uncarbonized. 5. Ascocarps usually exposing a red-brown disc in long ............................ 5. Ascocarps fissurine th 6. Ascocarps embedded in swollen tissue (?puffs?). 7. Stictic acid present .................................................. .. , ..26. G. triticea .............................. 14. G. insidiosa 6. Ascocarps not embedded in swollen t ? mmonly to 10 mm 7. No substances present ................ 8. Spores less than 11 pm long; no substances present .......... 24. G. subnitidula 8. Spores more than 13 pm long; stictic acid present ............ 6. G. dumastioides 9. Ascocarps white-powdery; exciple carbonized; lecanoric acid present .......... 2. G. afxelii norstictic acid present .................................................................................. 15. G. isidiifera present ...................................................................... 11. G. grammitis Exciples red-brown to barely carbonized at labial apices ........................ 10. G . glaucescens. 2. Ascocarps not fissurine; disc broad, obvious, not depressed within a thalline crack. 9. Ascocarps not powdery; exciple carbonized; lecanoric acid absent. 10. Thallus isidiate; thalline margin coarsely white-mealy; asocarps nearly round; 10. Thallus not isidiate. thalline margin not mealy; ascocarps elongate; no substances 1. Spores six locular o 11. 11. Exciples distinctly carbonized. 12. Labia distinctly striate. 13. Exciples with distinct red-yellow areas below the carbonized portions. 14. Thalline margin prominent; spores over 45 pm long ..... 14. Thalline margin absent or nearly so; spores under 35 g m . times dirty-brown below). 15. Ascocarps usually without a prominent thalline margin. 16. 16. Exciples closed below; no substances present 15. Ascocarps consistently with a thalline margin. 17. Norstictic acid present ......................... 17. Norstictic acid absent. 13. Exciples lacking red-yellow tissue below the carbonized portions (but some- Exciples open below; norstictic acid present ............. 8. G . elegans ......... 22. G . rimulosa 19. G. lumbricina 18. Exciple open below. 19. Spores usually 70-90 pm long .............................. 19. Spores usually 50-65 pm long ............................ 20. Exciple massively carbonized below .................... 9. G. flexibilis 18. Exciple closed below. 20. Exciple barely closed below ...... 18. G. Iongula 12. Labia intact or irregular, not distinctly striate. 21. Exciples distinctly open below. 22. Stictic acid only present ............................................... ..... 16. G. Zeptocarpa 22. Norstictic acid only present. 23. Spores less than 36 pm long ...................................................... 17. G. Zibrata 23. Spores over 45 pm long ................................. 21. Exciples distinctly closed below, or the lateral proaching each other very closely basally. 24. Ascocarps raised, without any thalline margin visible, short (1-2 mm long), Opegrapha-like ................................................................................ 1. G . adpressa 24. Ascocarps flush to raised but always with a thalline margin, usually over 2 mm long, not Opegrapha-like. 25. Spores under 51 pm long. 26. Stictic or norstictic acid present. 27. Norstictic acid present .................................... 3. G. desquamescens 27. Stictic acid present ................................................ 13. G. imshaugii 26. No substances present .................................................. 3. G . anfractuosa 25. Spores usually over 60pm long. 28. Stictic acid present ..... 28. No substances present .... 4 . G . anguilliformis 10 S\IITHSOSIAN C O S T R I B L T I O S S T O BOTASY 1. Graphis adpressa FKVRE 90; PLATE l a Grnphis nrlp).essn I'ainio, 1890: 119 [type collection: Carasqa, Braril, l'oinio 1289 (TUR, 1ectot)pe)l. ~ ) ~ . S ~ : K I P - I ~ I O Z . - - ~ ' ~ I ; I ~ ~ L ~ S it,hite, t?;r:i)., or pile brown, thin but continuous, shining. Xscocarps quit2 black, raised antl protuberant, usually un- branched, 110 thnlline nixgin 1-isible, 0.5-2 mn long, slender. Disc not visible in surface view. Asco- carp transverse section: liymeniuin 170-250 ,uin 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 pm, I + blue. CHESZISTRY.-NO substances present. H.AsrTAT.-Rain forest above 2600 feet. DISCL-SSIOS.-GVZ~~~S adpwssa externally resem- bles an Ojiegmplia or Xrlas/ i i lea antl is unlikely to be confused with any other Dominican Gmphis. It exhibits comiderably less variation in morphology than most Graphids. Extremely similar but for spore septation and size i s Gmphina niida. S r r c r ~ r s s Ex.&>risrD.-Fresh IVater Lake, 2600-2800 f t , Hnle 33485, 3.5487 (US), 2600-2877 f t , Iiushntig 32849.4, 32852A, 32864 (LiSC); Trois Pitons, 2900-3150 f t , Imshnicg 33090 (MSC); Riorne Diablotin, 33004300 ft, Iinshaiig 32940 (NSC); Jlicotrin, ca 3000 ft, TVirth 464 (US). 2. Graphis afzelii FIGURE 3b: PLATE l b Grojiliir a j x l i i .\charius, 1814:86 [ t vpe tollcction: Guinea, Afzelizu, s.11. (H, lectotype; UPS, isotype)]. I )F~C:RIP~-ION.- -T~I ; I~~II bi~ownisii tiin to gray, con- tinuous, smooth. Ascocarps usually compIeteIy covered b y ii poTvdery white thalline layer, appear- ing black only where this layer is rubbed, usually unbranched, 1.0-6.0 mm long, to nearly 1 n m wide. Disc not visible in surface view. Ascocarp transverse section: hymenium 75-1.50 high, I-; exciple black laterally, brown and closed to pale and open basally, lips conr.ergent, intact, with a proininent thalline covering extending to their apices. Spores S'ajcus, always 4 locular, 6-9 x 16-23 ,pn, I + blue. C?IE~rIsTRY.-I,ecanoric acid (in the thalline apothecial covering only). HABITAT.-D~~ sea-level scrub to virgin upland rain forest; apparently with broad ecological ampli- tude. DISCU~SIOS\'.-GYU~~~S afzelii is a common pan- tropic species, unique in the family in having lecanoric acid. This acid, along with the distinctive exciple structure, render i t unlikely to be confused with any other graphid. SPrcl\rrss ExlMIsEo.-Pointe Round, 100 f t , I m ~ h n t c g 33484.i (JISC): Londerry Agricultural Area, 100 ft, Imshnug SL'B9T ( \ I S C ) : iicar M a ( I i i ~ i i , 100 f t , Hnlr 3,5618 (CS); nnrrtl of Rinche, 200 ft, Hole 39644 (US); Can-Dom logging area, Se~\ - foundland , 800 f t , Hale 35239, 35116. 35447 (US); 8 mile5 north of l'ont Cnrsi., 1400 ft, T l ' i r t h ,546 (1-S); Cnn-I)oni losgiiis area, Dlenu Croiiitnier, 1600-l700 ft, H d e 35183 (US). 3 . Graphis anfractuosa F I G U R E 3C; PLATE 1 C Grnphis niifrcictttorn Eschweiler i ) i Martius, 1833:86 [type collection: Cai te te , 13rui l . s . n . ( \ I , lcctcit\pc; G , isot\pe)]. ~ ~ s C R I P T r o s . - ~ h a ~ ~ u s dull white to gray, thin, continuous. Ascocarps partially emergent, the labial tips appearing as slender, flexuose, black, sparingly branched lines 1.0-3.0 min long. Disc not visible in surface vieIv. Xscocarp transverse section: hynienium (SO-) 90-130 p.m high, I-; exciple black laterally, frequently dark brown to black below, usually closed, lips more or less intact, convergent. Spores S'ascus, 8-12 locular, 9-10 X 40-50 pm, I + blue. CHEMISTRY.-KO substances present. HAnITAT.-Iforne Diablotin, 35004300 f t , Zm- shazrg 32922, 32945A (MSC); Morne Trois Pitons, 30004000 f t , Zmshaug 32884 (MSC), summit, on Clusia uenenosa, Wirth 479, 481, 484, 485, 486 (US). 33. Phaeographis rosea FIGURE i h ; PLATE Gd Phaeographis rosea (I'ainio) Zahlbruckner, 1923:385. Graphis roses \.ainio, 1896:259 [type collection: Morne Anglais, Dominica, Elliott 523 (TUR, lectotype)]. DEScRIPTIo~.-Thallus tan, continuous, smooth and shining. Ascocarps barely emergent, rather 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 pm high, I-; exciple rudimentary, labia convergent, brownish above. Spores 8.'ascus, always 4 locular, 9-12 X (13-) 18-25 pm, I + blue. CHEI\ZISTRY.-~O substances present. HABITAT.-upper montane forest. DIscvssIos.-Phaeographis rosea is known only from the type collection. The rose-pink pigmenta, tion that renders this species so distinct is appar- ently a normal condition; occasionally, some graphids will show discolored, reddish areas on thalli of normally unpigmented species (see Gra- NUMBER 40 phina diinidiata for a Dominican example). T h e color in P. yosea, however, is restricted to the asco- carp margins, and appears to be normal. 34. Phaeographis cf. subtigrina FIGURE 7g; PLATE 6e Phaeographk subtigrina (T'ainio) Zahlbruckner, 1923:387. Graph i s subtigriria Vainio, 19OT:17T [type collection: Koh Chang, Gulf of Siam, Schmid t 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 pm 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 S/ascus, thick-walled, 4 locular, 4-6 X 10-17 pm, 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. DIscussIozr.-Phaeog?-aphis subtigrina has been reported (Vainio, 1915) from Guadelope, but we have not seen these specimens. The Dominican material differs from the type primarily in the chemistry. As in many species of Phaeographis, the ascocarp form in P. subtigrina varies greatly within the same thallus. In the Dominican specimens these 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. S P E C N E ~ S E x 4 ~ 1 h ~ ~ . - R o s e a u Botanic Garden, lVir th 444 (US); Emerald Pool, Wiith 505 (US). Key to the Species of Graphina 1 . Exciples distinctly carbonized. 2. Disc broad and obvious in surface view, labia quite divergent. 3. 3. Spores without colloidal layer; ascocarps more or less raised. Spores with broad persistent colloidal layer; ascocarps flush with thalline surface .......... .................................................................................................................................. 41. G. collospora 4. Spores I/ascus; protocetraric acid present ........... 52. G. platyleuca 4. Spores more than I/ascus, some darkening; n sent .................................. .............................................. G3. Phaeographina cf. difformis .... 2. Disc narrow, not prominent in surface view, labia more or less convergent. 5 . Ascocarps with powdery tvhite covering, easily rubbed to 6. Spores under 20 &rn long; salazinic acid present ........... 6. Spores over 40 pm long; stictic acid present .................. 5 . Ascocarps without a poivdery white coiering. 7. Labia entire or nearly so. 7. G triphoroides abial apices (upper ?,'* or less of total exciple ........................................................ 43. G . diniidiata o base of hymenium or beyond. 9. Ascocarps Opregraphn-like, 1-2 mm long, without a thalline margin ................ ...................................................................................................................... ~ 1 . G . nuda 9. Ascocarps elongate, flexuose, with a thalline margin. 10. Spores less than 61 pm long. 11. Norstictic acid present; mature spores 8/acus ................................ ................................................................................ . pseudoanaloga 11. KO substances present; mature spores usually 4-6/ascus ........................ ................................................................................................ 53. G . plurispora 30 SMITHSOKIAS C O S T R I B U T I O N S T O BOTASY 10. Spores over 85 pm long. 12. Only the ends of the spores muriform .................... 58. G. uestitoides 12. Entire spore densely muriform. 13. Ascocarps under 0.5 mm wide; hymenium under 250 pm high. 35, G. acharii 13. Ascocarps massive, usually 1 mm wide; hymenium over 350 pm high .... ..46. G. illinata ................................................................................................ 7. Labia striate. 14. 14. Exciple without yellowish areas, closed or nearly so. 15. Spores under 61 long. Exciple yellow to yellow-brown laterally, more or less open .... 37. G. antillarum Norstictic acid present; mature spores 8/ascus ...... 54. G. pseudoanaloga .............................................................................. ~ 3 . G. plurispora 16. 16. No substances present; mature spores usually 4-6/ascus ............................ 15. Spores over 64 @rn long. 17. Only ends of spores muriform ........................................ 58. G. vestitoides 17. Entire spore densely muriform. 18. Exciple base irregularly carbonized; spores always l/ascus ................ ............ .............................. 49. G. macella 18. Exciple base usually evenly carbonized; spores usually 4Iascus ........ .................................................................................................. 35. G. acharii 19. Apothecia fissurine, i.e., originating as a crack, the disc usually concealed by or sunken between the narrow.to gaping fissure walls. 20. Spores l/ascus, over 50 pn long ........................ 48. G. insculpta 20. Spores 6-8/ascus, under 50 pm long. 21. Thallus prosoplectenchymatous .................... 47. G . incrustans 21. Thallus not prosoplectenchymatous. 22. Psoromic acid present .................................. ..42. G . columbina 22. No substances present ................................... , G. cf. adscribens 1. Exciples totally uncarbonized. 19. Ascocarps not fissurine. 23. Disc broad, easily seen in surface view. 24. 24. Disc whitish, gray to greenish; spores over 17 p long; salazinic acid absent. Disc reddish: spores under 17 pm long; salazinic acid present .... 40. G. c o l l i ~ ~ l o ~ a 25. Spores 17-28 long; norstictic acid present .................... 56. G. suberythrella 25. Spores 50-150 pm long; ?quintaria? unknowns present ............ 59. G. virginea 23. Disc narrow, not prominent in surface view. 26. Margins of asocarps distinctly reddish to pinkish brown in surface view, a least in upper portions. 27. Spores l/ascus, over 80 pm long ............... 39. G. chlorocarpa 27. Spores Z-SIascus, under 7 5 pm long. 28. Korstictic acid present; exciples with internal striae only ............................ 28. So substances present; internal striae absent. 29. 29. Mature spores 8/ascus, less long; labia entire .............. .......................................................................... 38. G. carneouir .............................................................................. 44. G. dispersa Mature spores 2-4 ascus, over 35 ,pm long; labia slightly striate ........ ................... 55. G . rufopallida .................................................. 26. Margins of ascocarps tan, white, o r greenish. 30. Spores more than 40 pm long; salazinic acid absent. 31. Ascocarps always massive and protuberant; hymenium over 200 #m 31. Ascocarps rarely protuberant, never massive; hymenium less than 200 &m high, at least the epihymenium I + blue; ?quintaria? unknowns present 59. G. uirginea 30. Spores less than 17 pm long; salazinic acid present ............. 40. G . coll~culosa high; I-; no substances present. .................................... 45. G . frumentaria ................................................ NUMBER 40 31 35. Graphina acharii FIGURE 8a; PLATE 6f Graphina achaii (Fee) Mueller Argoviensis, 1887:38. Graphis acharii Fee, 1824:39 [type collection: South America, Graphis inturgescens Krempelhuber, 1876:383 [type collec- Graphitia inturgescens (Krempelhuber) Mueller Argoviensis, s.n. (G. lectotype)]. tion: Brazil, Glaiiou 6286 (BM, lectotype; M, isotype)]. 1888a:545. D ~ s c ~ r ~ ~ r o r v . - T h a ~ ~ u s gray to off-white, thick, 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: liymenium 150-200 pm high, proportionately small, I-; exciple black, massively closed, labia convergent, nearly entire to crenate to distinctly striate (often within a single ascocarp). Spores (in the type of G. acharii) usually 41ascus but occasionally one or three (and rarely six), densely muriform, 18-25 x 95-170 pm, I + blue. CHEMISTRY.-NO substances present. HABITAT.-secondary forest (Dominica). DIscvssIoN.-According to Mueller Argoviensis (1887), the spores in the type of G. acharii have biocellate ends; however, we have been able to find only densely muriform spores in this same material. In an earlier treatment of this species (Wirth and Hale, 1963), we had not seen the type, and followed Mueller in including specimens with biocellate spores in G. acharii. Although additional 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 T h e 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- (P. 44). phina species is G. macella, which differs in being smaller overall and in having an irregularly car- bonized excipular base. (US). SPECIMEN ExAM1xED.-Syndicate Estate, 1800 ft, Hale 35555 36. Graphina cf. adscribens FIGURE 8b; PI ATE 7a Graphina adscribens (Sylander) Mueller Argoviensis, 1892a: Graphis adscribens Nylander, 1868: 177 [type collection: Lifu, 284. New Caledonia, Thiebaut s.n. (H, lectotype.]. DEsCRIPTIoN.-Thallus white, thin, continuous to cracked. Ascocarps very inconspicuous in the 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 pm 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 21-25 pn (type), 9-10 X 2 3 4 0 (Dominica), I- in type (fide Nylander), I + reddish, slow in the Do- minican material. CHEJIISTRY.-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 Gyaphina 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 EXAXINED.- .~~S du Me, Wirth 519 (US). 37. Graplaina antillarum FIGCRE 8c: PLATE 7b Graphina antillarum (\'ainio) Zahlbruckner, 1923:398. Graphis an tillarum Yainio, 18993255 [tkpe collections, Guade- loupe, Duss 540 (FH, isotype)]. 32 SMITHSONIAN CONTRIBUTIONS T O BOTANY C FIGURE 8.-Cross sections of apothecia of Graphinn: a , G. acharii (lectotvpe in G); b, G. cf. adscribens (Wirth 519); c, G. antillarum (Duss s.n.); d , G. cnrneouiridis (Haie 3i642); e, G. chlorocarpa (lectotype in G); f , G. col[iculosa (lectotype of G. eugeniae, Merril l 3471): g, G. colliculosa (lectotype in PC). Graphis acuminata Vainio, 1915:147 [type collection: St. Jan, Graphina sulcata Fink, 1927:217 [type collection: Puerto Rico, Raunkiaer 437 (TUR, lectotype)]. Fink 659 (MICH, lectotype)]. D E S C R I P T I O N . - T ~ ~ ~ ~ ~ U S gra) to white, continuous, matte. Ascocarps nearly flush to partially emergent, black, commonly branched and flexuose, thalline Graphis platycarpoides Yainio, 1915: 145 [type collection, Graphina acuminata (Yainio) Zahlbruckner, 1923:393. Graphina platycarpoides (Yainio) Zahlbruckner, 1923:419. Guadeloupe, Duss 1198 (TUR, lectotype)]. NUMBER 40 33 margin absent or nearly so, 5-6 mm long, slender. Ascocarp transverse section: hymenium 80-100 (-120) pm high, I-, exciple black laterally, yellow to yellow-brown below, more or less open, labia con- vergent, barely to quite strongly striate. Spores (I-) 2-4 (-6)/ascus, distinctly muriform, 15-22 X 28-46 (-60) pm, I + blue. CHEMISTRY.-NOlStiCtiC acid. HABITAT.-LoWland scrub forest and higher alti- tude cultivated areas and road cuts. DrscussIoN.-Among the Graphina species with carbonized, striate, dimidiate exciples, G. antil- larum 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: Gmphina bzpartita XIueller Argoviensis ( 1 888c) 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. antillarum. One, however (Wir th 544c), has the brown exciple base of G. elongata but lacks the stictic acid of G. elongata. SPECIMENS ExAMINED.-Prince Rupert Bay, Irnshaug 33529A (MSC); Brookhill Estate, 100-150 ft, Imshaug 33307.4 (MSC); Roseau Botanic Garden, 100 f t Wirth 445 (US); north of Bioche, 200 f t , Hale 35745; South Chiltern Estate, 1300 f t , Zmshaug 33050, 33036 (MSC); near Pont Cassk, 1400 ft, Wirth 544c (US). 38. Graphina carneoviridis, new species Frcum 8d; PLATE 7c DESCRIPTION.-Tha11uS continuus, laevis, stratum corticale prosoplectenchymatum. Apothecia sessilia, recta vel flexuosa, 1-2 (-3) mm longa. Excipulum dimidiatum, rufo-fuscum, labiis convergentibus, in- tegris vel indistincte incisis; hymenium 75 pm altum, I-. Asci 8 spori; sporae decolores, murales, loculis horizontalibus 3, loculus transversis 2, 10-12 X 15-17 pm, I+ coeruleae. Thallus greenish, continuous to somewhat cracked, glossy, prosoplectenchymatous. Ascocarps raised, occasionally branched, straight to Bexuose, pink-brown above, laterally bounded by a green thalline margin, 1-2 (-3) mm long, slender. Disc not visible in surface view. Ascocarp transverse sec- tion: hymenium 75 pm high, I-; exciple reddish- brown, open below, labia convergent, intact to barely striate. Spores 8/ascus, 3 X 2 locular, 10-12 X 15-17 pm, I + blue. CHEMISTRY.-NO substances present. HoLoTYPE.-hIixed disturbed and primary rain forest, trail from Brigantin through XIiddleham Estate toward Morne Trois Pitons, Dominica, ele- vation 2200-2600 feet, Hale 37642 (US). D I S C L J S S I O N . - G ~ U ~ ~ ~ ~ U carneouiridis 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 chemistrv. 39. Graphina chlorocarba FIGURE 8e; PLATE 7d Graphina chlorocarpa (Fee) Mueller Argoviensis, 1887:44. Graphis chlorocarpa Fee, 1824:47 [ t j p e collection: Pe? 14. s.n. (G, lectotype)]. Graphina balbisii var. monospora Redinger, 1933:61 [type collection, Maltne 494 (S, lectotype; FH, isotype)]. DEsCRrPTroN.-Thallus tan to grey-green, contin- uous, glossy, smooth to slightly rugose. Ascocarps usually raised (nearly flush when young), straight to quite flexuose, usually unbranched, concolorous with the thallus except (usually) for the disc, which is pale orange-brown, 1-5 mm long, about 0.5 mm wide. Ascocarp transverse section: hymenium 120- 180 pm high, I-; exciple yellow-brown to red- brown, open below, labia convergent, lightly striate, the tips of the striae sometimes slightly darkened. Spores l/ascus, densely muriform, 24-40 X 80-130 pm, I+ blue. CHEMISTRY.-Both types and most specimens: no substances present. A few Dominican specimens with unknown substances. HABITAT.-A~I fertile specimens from rain forest; sterile material (see discussion) from sea level and 34 SMITHSONIAN CONTRIBUTIOKS TO BOTANY mid altitude cultivated areas. DIscussIorv.-The only neotropical G~aphina spe- cies likely to be confused with G. chlorocarpa is G. DrclDi.cii (Fee) Mueller .irgovieiisis. Both these species share the distinctive orange-brown disc area and the striate reddish exciple; the only difference lies in the number of spores per ascus. I n 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, 1iaI.e seen no ,pecinien~ that bridge the rather small gap between these two species. l\Tithout spores, i t 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 (Zmshaug 33024, 33417, 33424, Wivth 446) are sterile and may be either of the two species. No fertile material of G. balbisii has yet been identified from the Island. (US); Can-Dom logging area, Pont CassC., 2000 ft, Hale 39097, 35129 (US); Ridgefield Estate, 1100-1200 ft, In7shaug 33417, 33424 (JISC, both sterile): South Chiltern Estate, 1300 ft, I ~ n s i i a u g 33024 ( l ISC, sterile); Roseau Botanic Garden, 100 ft, Wirth 446 (US, sterile). SPECI.UESS ExAMINED.--Sear Laudat, 800 f t , Hale 35797 40. Graphina colliculosa FIGURES 8f,g; PLATE 7e Graphina colliculosa (hfontagne) Hale, 1976: 156. Sclerophyton coliicuZosu~n Montagne, 1851 :61 [type collection: Graphis intortnila Stirton, 1881 :I86 [type collection: Assam, Graphis eugeniae Yainio, 1921:262 [type collection: Philip- Graphis colliculosa (Montagne) Zahlbruckner, 1923:299. Graphina intortula (Stirton) Zahlbruckner, 1923:411. Graphina aibonite7isis Fink, 1927:219 [type collection: Puerto Guyana, Leprieur 1406 (P, lectotype: Bhf, isotype)]. s.n. (BM, lectotype)]. pines, Merrill 3971 ( T U R , lectotype: FH, isotype)]. Rico, Fink 2017 (MICH, lectotype)]. ~ E S C R I P T I ~ S . - T ~ ~ ~ I I L I ~ 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 pm 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 pm, I + blue. CIIEMrsTRY.-Salazinic acid. HABITA.r.-Rain forest canopy in Dominica; else- where found in both rain forest and secondary forest. DISCLXIOS.--GYU~~~~U colliculosn 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, hssam, and the Philippines. As in many species with totally uncarbonized exciples, the degree of labial divergence is quite variable (see Phaeographis n lb ida 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. T h e 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. colliculosn is unlikely to be con- fused with any other species, as salazinic acid is quite rare in the family. However, several other species are quite 5iinilar in inorpholog), a s follows. Graphina eiythrella (Montagne) Zahlbruckner (1923) is distinguished by having norstictic acid, larger spores, and somewhat shorter, more raised ascocarps. Some of the sterile, norstictic acid-con- taining syntypes of G. eugenine (MacGregor 41315, Elmer 15077) may be referable to this species. G9,aphina riopiedwnsis Fink (1927) differs only in the somewhat larger spores and in having stictic acid. SPECIVENS ExAlrINED.fOrne Diablotin, 2200-2600 ft, Hale 38090 (US). 45. Graphina frumentaria FIGURE 9f; PLATE 8d Graphina frumentarh (Fee) MuelIer Argoviensis, I880:40. Graphis frumentaria Fee 1824:45 [type collection: Peru, Mutis s.n. (G, lectotype)]. DEscRIPTIoN.-Thallus buff-gray, smooth to 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 pm 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 pm, I+ blue. CHEMISTRY.-NO substances present. HABITAT.-virgin rain forest. DIscussIoN.-Graphina frumentaria is somewhat similar to G. chlorocarpa but differs in its very large ascocarps, entire labia, and in having 6-8 spores per ascus. T h e 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 ExAhrINED.-Can-Dom logging area, Newfound- land, 800 ft, Hale 35242 (US). 46. Graphina illinata, new combination FIGURE 9g; PLATE 8e Graphis illinata Eschweiler in Martius, 1833:82 [type collec- tion: Brazil, s.n. (M, lectotype)]. DEscRrPTIor\..-Thallus slate gray to whitish, smooth, glossy, thick. Ascocarps very protuberant, 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 (!) pm high, I-; exciple black, massively closed, labia more or less convergent, more or less entire, completely covered by the thal- line margin. Spores 1 iascus, densely muriform, 30-50 X 110-225 pm, I+ 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 SMITHSOXIAS CONTRIBUTIOSS T O BOTANY 38 collectors. T h e implication, of course, is that many other less conspicuous canopy species remain uncol- lected on the island. SPECIMENS Ex.AwI\ED-Can-Dorn logging area, Biantridge Estate, 1700 ft, Hale 35276, 35281, 35284 (US); Can-Dom log- ging area, Pont Casse, 2000 ft, Hale 35082, 33083, 35124 (US). 47. Graphina incrustans FIGURE 9h; PLATE 81 Graphina incrustans (Fee) Mueller Argoviensis 1887:47. Fissurina iricrustarrs Fee, 1824:60 [type collection: South Graphis rubigitiosa Fee, 1824:47 [type collection: South Amer- Graphina rubiginosa (Fee) Mueller Argoviensis, 1887:44. Graphis glaucoderina Xylander ex Tuckerman, 1888: 124 (type Graphis dehiscens l'ainio, 1890: 11 1 [type collection: Brazil, Fissurina nit idescens Nylander, 1890: 108 [type collection: Graphina glaucoderma (Xylander ex Tuckerman) Mueller Graphis nit idescens (Nylander) Vainio, 1915: 151. Graphina nitidesceiis (Sylander) Riddle, 1918: 115. Graphina dehiscens (Vainio) Redinger, 1933: 13. America, s.n. (G , lectotype)]. ica, s.n. (G, lectotype)]. collection: Cuba, Wright 61 (FH, lectotype)]. Vainio 306 (TUR, lectotype)]. Florida, USA, Calkins 31 (H, lectotype)]. Argoviensis 1895b347. DEScRrPTroN.-Thallus yellow-brown to 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 pm 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 pm, I- or I + slow, reddish. CHEMISTRY.-NO substances present. HABiTAT.-Rain forest and elfin forest. DIscussIoN.-Graphina incrustans is a member of a species complex which appears to parallel the fissurine Graphis species (see Graphis triticea). It is a sporomorph of Graphis dumastii; undoubtedly a whole series of sporomorphs will be found between these two groups. T h e species listed in synonymy overlap with each other completely and cannot be maintained on any reasonable grounds. TSe have concentrated on Neo- tropic species, but many Paleotropic types will un- doubtedly be involved here also; we have not yet seen T L C analyses for these. Most similar to G. incrustans is probably G. babingtonii (Montagne) Zahlbruckner; this species is tentatively separable by its more strongly gaping ascocarps and in having only 4-6 spores per ascus. Graphina insculpta is morphologically identical, separable only in having one very large spore per ascus. SPECIMEXS Ex4\fIsED.-hficotrin, 3000 f t , Wirth 465 (us); South Chiltern Estate, 1200-1500 ft, Imshaug 32802 (MSC); Freshwater Lake, 2600-2877 ft, Imshaug 32857A (MSC). 48. Graphina insculpta FIGURE 10a: PLATE 9a Grnphina insculp ta (Eschweiler in hfartius) hfueller k g o - viensis, 1888b:307. Diovgrna insculpta Eschweiler i r i ;\fartius, 1828:9 [type col- lection: Brazil, s.n. (M, lectotype)]. Graphis ir iscdpta (Eschweiler iii Martius) Sylander , 1863:371. DEsCRIPTIos.-Thallus greenish to yellow-brown, smooth, gloss), thick, quite prosoplectenchymatous in section. Ascocarps fissurine, concolorous with the thallus, usually unbranched, straight to slightly curved, 0.5-1.5 mm long. Ascocarp transverse sec- tion: hymenium 90-120 pm high, I-; exciple rudi- mentary, yellow to pale red-brown, open, labia convergent to spreading, with much included bark. Spores l/'ascus, densely muriform, 20-30 X (50-) 70-75 (-100) pm, I- or I + slow, reddish. CHEMISTRY-NO substances present. HABITAT.-Rain forest. D I S C L J S S I ~ K . - G ? - U ~ ~ ~ ~ U insculpta is morpholog- ically identical to G. incrustans and can be sep- arated only by having one very large spore per ascus. (US). SPECIMENS EX.4alINED.-Near Pont C a s e , 1400 ft, Hale 37819 49. Graphina macella FIGURE lob; PLATE 9b Graphina inacella (Krempelhuber) Mueller Argoviensis, 1880: 39. NUMBER 40 39 , FIGURE 10.-Cross sections of apothecia of Graphina: a, G. insculpta (lectotype in G); b, G. macella (Glaziou 6289B); c, G. marcescens (lectotype in G); d, G. nuda (Hale 35119); e, G. platyleuca (Thiebaut s.n.); f, G. plurispora (Malme 998); g, G. pseudoanaloga (Duss 1590); h, G. rufopallida (Duss 527). 40 SMITHSOXIAN COSTRIBUTIOXS T O BOTANY Graphis macella Krempelhuber, 1876:380 [type collection: Brazil, Glaziou 6289b (RI, 1ectot)pe)l. DESCRIPTioN.-Thallus gray to white, continuous, smooth to lightly rugose, glossy to somewhat matte. Ascocarps raised but not prominently so, straight to curved, usually unbranched, apically black or black with thin white thalline stripes, laterally covered by the thalline margin, 1-5 mm long, less than 0.5 nim wide. Disc not visible in surface view. Ascocarp transverse section: 1i)menium 140-160 pm high, I-; exciple black laterally and above, fre- quently brown or irregularly Carbonized below, closed to nearly open, labia convergent, striate. Spores l/ascus, densely muriform, (20-) 2 5 4 0 x (-65) 95-130 pm, I + blue. CHEMISTRY.-NO substances present. HABITAT.-Rain forest (Dominica). DIscussroN.-Graphina macella is similar to G. acharii but is smaller overall, more sunken, 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 llexico, indicating that this spe- cies will probably be found throughout the Neo- tropics. 35723 (US). SPECIMEN EXAMINED.--\?alley of Desolation, 2800 ft, Hale 50. Graphina marcesceizs FIGURE 1Oc; PLATE 9c Graphina marcescens (Fee) Mueller Argoviensis, 1887:42. Graphis marcescens Fee 1824:38 [type collection: South Amer- ica, Humboldt et Bonpland s.n. (G, lectotype)]. Graphis intricata Eschweiler in Martius, 183359 (non Graphis intricata Fee, 1824:42) [type collection: Brazil, Martius s.n. (hf, lectotype)]. Graphina intricata (Eschweiler in Martiusj hlueller Argo- viensis, 1888b:510. Graphilia p l i t t i i Zahlbruckner, 1928:60 [type collection: Flor- ida, USA, Plitt s.n. (US, lectotype; L, isotype)]. DEscRIPTIoS.-Thallus off-white, continuous, matte to nearly powdery. Xscocarps more or less emergent, rarely branched, quite flexuose, 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) pm, high, I-; exciple black above, more or Iess closed below by paler yellow to red or brown tissue, labia convergent, entire to slightly crenate, completely thalline covered. Spores 8/ascus, 1-2 X 4-5 locular, 5-8 x 12-16 pm, I + blue. CHEMrsmY.-Salazinic acid. HABIrhT.-upland rain forest (Dominica; see remarks). DIsccssIoh-.-Graphina marcescens is unlikely to be confused with any other Graphina species. Three characteristics account for this individuality: The red-brown exciple base, the presence of salazinic acid, very raie in the family, and the powdery thal- line covering, easily rubbed, completely covering the ascocarps. Externall), G. marcescens is identical to Graphis candidmima Zahlbruckner (1923) but i s 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. mal-cescens) is from high virgin rain forest; the type of G. pli t t i i is from Florida, and hence probably lowland forest. In addition, we hale seen a specimen from 2200 meters, collected in India (Hale 35312X), which differs from the Neo- tropical material only in being somewhat larger and in having the exciple base a bit more carbon- ized. SPECIlIEV Ex~~IhED.-can-Dom logging area, Dleau Gom- mier, 1600-1700 ft, Hale 34312.4 (US). 51. Graphina nuda FIGURE 10d: PLATE 9d Graphina nicda Magnusson, 19553266 [tvpe collection: Mauna Kea, Haw?iii, Faurie 1023b (UPS, holotype)]. DEscRIPTIox.-Thallus gray to tan-gray, contin- uous, smooth to minutely roughened. Ascocarps very prominent, black, straight to slightly curved, unbranched, without a thalline margin, resembling Graphis adpl-essa, 1-2 mm long, slender. Disc not visible in surface view. .4scocarp transverse section: hymenium 135-150 p m high, I-; exciple black usually closed but occasionally irregular at the base, lips intact, convergent. Spores 8/ascus, uniseriate, 1-4 x 4 locular, 16-18 X 20-30 pm, I + blue. CHEJIISTRY.-NO substances present. HABITAT.-coastal cocos to virgin upland rain D I S C L W O N . - G Y U ~ ~ ~ ~ U nuda is distinctive among forest; apparently of wide ecological amplitude. 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 (FCe) Mueller Argoviensis (1887), which is exter- nally and internally very similar, differs in having much smaller spores. SPECIMENS EXAMINED.-Prince Rupert Bay, Zmshaug 33536.4 (MSC); Can-Dom logging area, Pont C a d , 2000 ft, Hale 35092, 35119 (US). 52. Graphina platyleuca FICURE 10e; PLATE 9e Graphina platyleuca (Nylander) Zahlbruckner, 1923:420. Graphis platyleuca Nylander, 1868:114 [type collection: New Caledonia, Thiebaut s.n. (H, lectotype)]. Graphina platycarpa Fink, 1927:219 (non G. platycarpa (Esch- weiler in Martius) Zahlbruckner, 1902:389) [type collection: Puerto Rico, Fink 1774 (MICH, lectotype)]. Graphina platycarpinu Zahlbruckner, 1932:Z 14 (nomen noviim for G . platycarpa Fink). DEscRIPTIorv.-Thallus greenish white, 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 pm, high, I + blue, epihymenium dark; exciple irregularly carbonized, quite black in parts, red- brown in other patches, closed, lips quite divergent, entire. Spores 1 iascus, densely muriform, 3 5 4 0 X 120-165 pm, I+ blue. CHEMIsmY.-Protocetraric acid only (types of G. platyleuca, G . platycarpa), or protocetraric acid with salazinic acid and norstictic acid (see discus- sion). HABITAT.-PrimarY rain forest (Dominica). DI~CUSSION.-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). ft, Hale 37955 (US). SPECIhIEN ExAMINED.-Dleau Gommier Forest Reserve, 1200 53. Graphina plurispora, new combination FIGURE lo!; PL.4TE 9f Graphina pseud osop h is t ica var. plurispora Redinger , 193 3: 36 [type Collection: B r a d , Malme 998 (S, lectotype)]. DEscRIPrIorV.-Thallus continuous, gray-white, smooth and glossy to matte (on the same thallus). Ascocarps barely raised, black above, frequently with white stripes, with a well-developed thalline margin laterally, quite flexuose, 2-5 mm long, slen- der. Disc not visible in surface view. Ascocarp transverse section; hymenium 60-100 pm high (on the same specimen), I-; exciple very variable (within one ascocarp), black, distinctly oI:en to dis- tinctly closed, lips convergent, perfectly intact to quite striate. Spores begining as Siascus, but only (2-) 4-6 maturing, muriform, 1-3 X 8-13 locular, 10-15 X (27-) 35-50 (-60) pm, I+ blue. CHEMISTRY.-NO substances present. HABITAT.-cultivated areas above 1500 feet (Dominica only). DrscussIoN.-Redinger?s variety differs from the type of Graphina pseudosophistica in two impor- tain respects. The latter has consistently mono- sporous asci and has the ?quintaria? unknowns. Because of these differences, we have raised the variety to species rank. Still, Graphina plurispol-a is both highly variable and part of a yery confusing array of species. Most closely related are the fol- lowing Graphina disseypens (Nylander) Mueller Argovi- ensis (1880) is tentatively separable because of the quite dendritically branched ascocarps and consist- ently open exciples. Graphina elongatoradians Fink (1927) is another 42 SXIITHSONIAN CONTRIBUTIONS T O BOTANY dendritically branched relative with longer, more slender ascocarps, with open exciples, and a differ- ent chemistr) (two unknown substances). Graphzna subvelata (Stirton) Zahlbruckner (1923) differs in being non-striate and in having open ex- ciples. See H,i\wdrd (1gi8) foi d discussion of G. subuelata and its near relatives, including New JVorld species. In addition, G. plurispora bears a number of similarities to the G. antzllarum complex; for a more complete discussion, see the latter species. 1500 ft, Irnshaug 32i24.A, 32727 (MSC) SPECIMEM E X A V I X E D -South Chilteiti F\ ta te Road, 1500- 54. Graphina pseudoanaloga FIGURE log; PLATE 10a Graphina pseudoanaloga (Vainio) Zahlbruckner, 1923:421. Graphis pseudoanaloga L?ainio, 1915:145 [type collection: Guadeloupe, Duss 1590 ( T U R , lectotype)]. DEScRIPTIox.-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 visibIe in surface view. Ascocarp transverse section: hymenium 75-1 15 pm high, in- spersed (apparently clear i n TVirth 526), epih)me- 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 pm, I f blue. CHEl\zISTRY.-r\TOrStiCtiC acid. HABITAT.-Beach to upland virgin rain forest; apparently of wide ecological amplitude. DIscussIoN.--Graphina pseudonnaloga is very similar to G. annloga (Nylander) Zahlbruckner (1923), which differs in having smaller spores and an open exciple. SPECIMENS ExAMISED.-Cabrit, sea level, Wirth 526 (US); Can-Dom logging area, Brantridge Estate, 1700 ft, Hale 33283 (US). 55. Graphina rufopallida FIGURE 10h; PLATE 10b Graphina rufopallida (T?ainio) Zahlbruclner, 1923:423 (?rufo- Graphis rufopallida T?ainio, 1915:149 [type collection: Guade- pallens,? sphalm.). loupe, Duss 527 (TUR, lectotype)]. ford Estate, Dominica, Elliott 1852 (TUR, lectotype)]. Graphis verrnicularis \.ainio, 1915: 148 [type collection: Shaw- Craphina verrniczilaris (L?ainio) Zahlbruckner, 1923:430. DEscRIPTIos.-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 pm 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 6-81immature ascus, apparently always aborting to 2-4/mature ascus, densely muriform, 15-30 X 36-75 pm, I+ blue. CHEMISTRY.-NO substances present. HABITAT.-upland rain forest. D I S C U S S I O S . - G ~ U ~ ~ ~ ~ U rufopallida is most simi- lar to G. bulbisii (see discussion under G. chloro- carpa), from which it differs in the lower, much longer, more slender and intricating ascocarps. Itre have seen no specimens of this species other than the two types cited. SPECIMEN ExAxrrsm.-Shawford Estate, Elliott 1852 (TUR). 56. Graphim suberythrella, new species FIGURE l l a ; PLATE 1Oc DEscRIPTIox.-Thallus continuus, laevis, stratum corticale bene evolutum. Apothecia subimmersa, flesuosa. vulgo iimplicia, 2-4 mm longa. disco lato, pruinoso. Excipulum integrum, flavidum vel rufo- ferrugineum, labiis integris, divergentibus. Hy- menium 110-140 pm altum, I+ coeruleum. Asci 8 spori; sporae decolores, murales, loculis horizontali- bus 5-7, loculis transversis 1-3, 10-14 X 17-28 pm, I + coeruleae, in juventute halonae indutae. Thallus gray, glossy, continuous, rather thick, cortical layer compact and distinct. Ascocarps barely raised, flexuose, occasionally branched, 2-4 mm long, slender, concolorous with the thallus. Disc broad, dark gray, lightly pruinose. Ascocarp transverse section: hymenium 110-140 pm high, at least the epihymenium I+ blue; exciple J-ellow to red-brown, closed, lips intact and divergent. Spores 8/ascus, 1-3 X 5-7 locular, halonate in early stages, 10-14 X 17-28 pm, I+ blue. CHEMISTRY.-NOrStiCtic acid. NUMBER 40 43 FIGURE 11.-Cross sections of apthecia of Graphina: a, G. suberythrella (Imshaug 32699); b, G. triphoroides (Hale 35161); c , G. vestitoides (Fink 1986); d, G. virginea (Wirth 547); e, G. virginea (lectotype of G. triangularis, Pringle 17); j , G. virginea (lectotype of G. obtectula, Tonduz). HoLorYPE.-Cultivated area, South Chiltern Es- G. erythrella (Montagne) Zahlbruckner (1923), but tate Road, Dominica, elevation 1500-1700 feet, differs in its more immersed ascocarps, gray disc, Imshaug 32699 (MSC); isotype, US). I- hymenium and larger spores. Also similar is D I S C U S S I O N . - - G ~ ~ ~ ~ ~ ~ U suberythrella resembles G. insignis (Vainio) Zahlbruckner (1923), which dif- 44 SMITHSONIAN CONTRIBUTIONS T O BOTANY fers in having wider ascocarps and smaller spores, and in lacking both norstictic acid and the I+ hymenium. SPECIMEX ExAMIsED.-can-Dom logging area, Newfound- land, 800 ft, Hale 35224 (US). 57. Graphina triphoroides, new species FIGURE l l b : PLATE 10d DEscRIPTIoN.-Thallus continuus, farinosus. Apo- thecia alte sessilia, simplicia, margine thallino usque ad verticem vestita, 1-6 mm longa, 1 mm lata. Excipulum dimidiatum, fuligineum, labiis in- tegris vel irregulariter crenulatis, erectis vel con- vergentibus; hymenium 250-700 pm altum, I-; Asci 6 (-8?) spori; sporae decolores, murales, loculis hori- zontalibus 7-10, loculis transversis 2-4, 18-30 X 47-75 pm, I+ coeruleae. Thallus white to off-white, continuous, mealy. Ascocarps large, very protuberant, unbranched, covered to the apices by a mealy concolorous thal- line margin, irregularly blackened where the cov- ering is rubbed, 1-6 mm long, I mm wide. Disc usually not visible in surface view. Ascocarp trans- verse section: hymenium 250-700 (!) pm 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 (-S?)/ ascus, muriform, 2 4 X 7-10 locular, 18-30 X 47-75 pm, I+ blue. CHEJfISTRY.--StiCtiC acid. HoLoTYPE.-can-Dom logging area, Dleau Gom- mier, 1600-1700 ft , Hale 35161, January 1969 (US). DIsccssIoN.-Graphina triphoroides is part of a very distinct group of three New r\'orld species, all characterized by very large, thalline-covered apo- thecia and dimidiate, carbonized exciples. Graphina triphol-a (Sylander) Alueller Argoviensis (1880) dif- fers in having 3 4 spores per BSCLIS, no lichen acids, and carbonization restricted more to the upper por- tions of the labia. Graphina cleitops (Fee) RIueller Argoviensis (1887), the third species, differs in being monosporous. SPECIMENS E x ~ ~ ~ ~ m . - - C a n - D o r n logging area, Dleau Gom- mier, 1600-1700 ft, Hale 35128 (US); Wi-ight 15, Cuba (US) ("type" of Graphis triphomides Kylander, unpublished name). 58. Graphina vestitoides FIGURE l lc ; PLATE 10e Graphina vestitoides Fink, 1927:218 [type collection: Puerto Phaeographis cervicuiata Redinger, 1933:76 [type collection: R i b , Fink 1986 (MICH, lectotype)]. Brazil, Maline 3531 (S, lectotype; UPS, isotype)]. DEscRIPrIos.-Thallus gray, continuous, smooth. Ascocarps quite protuberant, frequently branched, flexuose, varying from black above and partially covered by a thalline margin to concolorous (com- pletely covered), to 10 mm long, ca. 0.5 mm wide. Ascocarp transverse section: hymenium 150-2 10 pm high, I-; exciple black, usually closed, lips conver- gent, striate to nearly entire. Spores 4-6 (-8)/ascus, biocellate only in the terminal few sections, 12-18 X 85-140 pm, I + blue. CHEMISTRY.-NO substances present. HhBITAT.-upland virgin and secondary forest. DIscr,ssIos.-C;raphina vestitoides is separable from G. acharii only in the peculiarly biocellate spores, which additional collections may prove to be an unreliable character. The type material of Phaeographis ceruiculata has mature spores that are quite clear, and the specimen is identical to G. vestitoides. SPECIMENS ExAMrhm.-Freshwater Lake, 2600-2877 f t , Ims- Izaug 32859'4 (MSC); near Pont C a d , 1400 ft, Wirth 544a (US). 59. Graphina virginea FIGURE lld,e,f; PLATE l O f Graphina virginea (Eschweiler in Martius) Mueller -4rgovien- sir, 1880:41. L e i o g ~ a m m a virgineum Eschweiler i n Martius, 1833:98 [type collection: Brazil, Afartiiis s.n. (XI, lectotype)]. Graphinn obtec tdn Mueller Argoviensis, 1892b: 133 [type col- lection: Costa Rica, Pit f ier (3166 (G, lectotype); Figure l lf)] . Graphina t ~ i e l a l e u c n Miieller Argoviensis, 1895,: 144 [type col- lection: Roscau, Dominica, Eckfeldt 70 (G, lectotype)]. Grnfihina paluiei-i Zahlbruckner, 1921:231 [type collection: Mexico, Pringle 9 (MICH, isotype)]. Grnpliina triangiiinris Zahlbruckner 1921 :232 [type collection: Mexico, Priizgle 17 (MICH, isotype)]. Graphis collospot~elin I'ainio, 1923: 141 [type collection: Trini- dad, Tl iax t e r 10 (TUR, lectotype)]. Graphif in collosfiorelin (Vainio) Zahlbruckner, 1923:212. D E S C R I P T I O ~ . - ~ ~ ~ ~ I ~ ~ L I S 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 concoloi ous to much paler than the thallus, occasionally branched, flexuose, 1-5 mm long, slender. Disc in surface view varying from concealed to quite broad, concolorous to darker than the thallus. Ascocarp transverse sec- tion: hymenium 120-200 pm high, epihjmenium darkened, at least the epihymenium I + 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) pm, I+ blue. CHEMISTRY.-At least one of the 3 so-called ?Quintaria? unknowns; trace of stictic acid and constictic acid (type of G. uzrgznea only). HABITAT.-secondary growth and cultivated trees and shrubs, from sea level to over 1000 meters. DIscussrox.--In its usual form Graphina uirginea is an extremely vigorous species, forming thalli up to 15 cm in diameter. The thallus is normally greenish white, matte, and continuous except for characteristic fissures bordering the ascocarp mar- gin. The ascocarps themselves are commonl) slightly paler than the rest of the thallus, and do not occur in close proximity to each other. In all likelihood, the species is weedy and of rapid growth, as large colonies are commonly found on such fast-growing cultivated plants as cacao and blue Mahoe (Hibis- 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. T h e other taxa listed as synonyms all show some cus). structural variation from the typical pattern. In the case of G. palmeri and G. melaleuca the thallus in surface view appears quite rugose and the asco- carps somewhat emergent. I t 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. uirginea is G. monophora (Nylander) Zahlbruckner (1923). The type of G. melnleiica also has a broader, browner disc than is normally found in G. uirginea; externally this specimen is indistinguishable from G. mendax (Nylander) Mueller-Argoviensis (1 888a). The different disc is, however, a matter of degree rather than of kind; occasionally one can find such discs on parts of more typical thalli. Graphilia palmrr i shows the internal striae that seem to arise with age and lateral compaction of the old hymenium; we do not find such striae to be at all dependable in distinguishing species (see Graphina colliculosa for a parallel variation). The internal striae, coupled with a more or less continuous semicarbonized band below the exciples, is also characteristic of the type of G. triangularis, which externally is much like typical virginea. The band is occasionally found in thalli of quite normal G. uirginea, frequently unconnected with the ascocarps. The thickened exciple base of the type of G. triangularis is another condition that we feel may well be simply a factor of age (again, see G. colliculosa for parallels). All the forms of G. virginea are tied strongly together by their chemistry. All have the rather rare It blue reaction of (at least) the epihymen- ium; all share at least two of the three ?Quintaria? acids. SPEci \ f tNs Ex4\iINEo.-Biookhi~1 Estate, 100-150 ft, Ims- haug 33321.4 (MSC); Cocoa Centre on Layou Rixer, 200 ft, Wirth 517 (US); north of Pont Casse., 1400 ft, Wirth 543, 545, 547, 548 (US). Key to the Species of Phaeographina 1 Exciple with carbonized areas. 2. Exciples carbonized and closed below ...........................).......)...............,....,..,.. 63 P. cf. diflormis 2. Exciples carbonized only a t labial tips. 3. Ascocarps massive, 1-3 mm wide: no substances present ........_........... 61. P . caesiopruinosa 3. Ascocarps slender, less than 0.4 mm wide: stictic acid present .,......,....... 65. P . oscitans 46 SMITHSONIAN CONTRIBUTIONS TO BOTANY 1. Exciple uncarbonized. 4. Labia convergent: spores more than 40 Gm long .................................................... 64. P. eZZiottii 4. Labia divergent, exposing the disc; spores less than 40 &m long. mmonly over 5 mm long; norstictic acid present 5 mm long; norstictic acid abse .......................................................... 60. P. atrmtermicdaris 6 . Spores 20-27 rm long; stictic acid present ................................. 6. Spores 33-85 gm long; no substances present .............................................. 62. P. coriaria 60. Phaeographina atrovermicularis, new species FIGURE 12a; PLATE l l a DEscRIPTIoN.-Thal~us continuus, sublaevigatus. Apothecia adpresso-sessilia, vulgo 1-amulosa, 3-10 mm longa, disco dilatato, nigro. Excipulum dimi- diatum, fuscum, labiis integris, divergentibus; hy- menium 145-220 pm altum, I-. Asci 6-8 spori; sporae fuscescentes, murales, loculis horizontalibus 6-9, loculis transversis 1-4, 15-17 X 25-35 pm, I + coeruleae. Thallus thin, light tan, continuous to apparently eroded except near the ascocarps, smooth to matte. Ascocarps very long, raised, frequently branched, the pale thalline margin very prominent, 3-10 mm long, slender. Disc very prominent, black, occa- sionally faintly pruinose. Ascocarp transverse sec- tion: hymenium 145-220 ,urn 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+ blue. CHEMISTRY.-NOrStiCtiC acid. HoLoTYPE.-can-Dom logging area, Brantridge Estate, Dominica, elevation 1700 feet, Hale 35278 DISCUSSION.-B~ 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- cedens AIueller Argoviensis (1 888e), which differs in having shorter, broader, stockier ascocarps, larger spores, an unknown acid in addition to norstictic, and no trace of carbonization in the exciple. Also somewhat similar is the widespread P . caesioprui- nosa, which has much shorter, broader ascocarps, strongly pruinose discs, and lacks norstictic acid. SPFCIME\S ExAMINrD.-can-Dom logging area, Pont Cassk, 2000 f t , Hale 35093 (US); rain forest, Jean Estate, 2000 ft, Hale 35068 (US). (US). 61. Pltaeographina caesiopruinosa FIGURE 12b; PLATE l l b Phaeographina caesiopruinosa (Fee) Mueller Argoviensis, Arthonia caesiopruinosa Fke, 1837:36 [type collection: South 1887:49. America, s.n. (G, lectotype)]. DEscRIPTIoN.-Thallus pale brown, smooth to slightly roughened, continuous. Ascocarps quite ele- vated, rarely branched, straight to flexuose, thalline margin reaching to disc, 1-10 mm long, 1-3 mm wide. Disc very prominent in surface view, lightly to very heavily pruinose. Ascocarp transverse 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, I+ slow, reddish blue. CHEMISTRY.-NO substances present. HABITAT.-cUltivated areas, uplands. DIscussIoN.-Phaeographina caesiopruinosa is a very common New World species, unlikely to be misidentified. All other New World species with very large ascocarps and carbonized exciples have monosporous asci. Zmshaug 32731 (MSC). SPECIMEN EXAMINED.-sOUth Chiltern Estate, 1500-1700 f t , 62. Phaeographina coriaria, new species FIGURE 12c; PLATE l l c DEscRrmIoN.-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. difjormis (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 pm, I+ rufo-fuscae. 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- Thallus continuous, smooth, tan, resembling ina. Ascocarp transverse section: hymenium ca. 150 pm high, beaded, epihymenium dark, I-; exciple 48 SMITHSONIAS COXTRIBUTIOSS T O BOTASY rudimentary to very pale below, laterally red-brown, lips intact, divergent. Spores 4-6 (-8?) lascus, brown, 8-10 x 1-3 locular, 9-10 x 33-35 pm, I + red, slowly darkening. CHEMISTRY.-~O substances present. HOLOTYPE.--;\~OSSY forest, Morne Anglais, DO- minica, elevation 3000-3600 feet, Hale 35378 (US). DIscussIoN.-Phaeographzna coriaria is most sim- ilar to the following three New 'CYorld species: Phaeographina arechaualetae Mueller Argoviensis (1888~) differs in having distinctly carbonized labia and rotund spores. Phaeographina caracasana Mueller Argoviensis (1887) differs in having much broader ascocarps with rounder apices and in having smaller spores. Phaeographina elliptica IVirth and Hale (1963) differs in chemistry and in having monosporous asci. 63. Phaeographina cf. diflormis FIGURE 12d; PLATE l l d Phaeographina d i f fomis Fink, 1927:220 [type collection: Aibonito, Puerto Rico, Fink 1874 (MICH, lectotype)]. DEscRIPTIoN.-Thallus thin, continuous, glossy, greenish gray. Xscocarps nearly round to elongate and irregular, ends round, occasionally shortl) branched, partly immersed, 0.8-1 3 mm long, ca. 0.8 mm wide, with a raised thalline margin. Disc broad, brown-black, occasionally faintly pruinose. Ascocarp transverse section: hymenium 90-125 pm high, epihymenium very dark, I-; exciple 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 pm in the type, 18-22 x 55-75 in the Dominican specimen, I+ blue. CHEMISTRY.-UnknO\Vn in the type, no sub- stances present in the Dominican specimen. HABITAT.-Elfin forest (Dominican specimen only). DrscvssIos.-;\ltliough 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 Graphinn confluens alliance, such as G. cymbe- graplzn (Nylander) Zahlbruckner (1 923), which is strikingly similar. Very similar to P. difformis is P. explicans Fink ex Hedrick (1933), which differs in its more elon- gate, rather sunken apothecia. SPECI\IEI\ Exi\rIzlED.-summit of Morne Diablotin, 4300- 4350 f t , Imshaug 32913,4 (MSC). 64. Phaeographina elliottii FIGURE 12e; PLATE l l e Phaeographina elliottii (Vainio) Zahlbruckner, 1923:438. Graphis elliottii T'ainio, 1915:143 [type collection: Dominica, Roseau Valley, Elliott 1338 (TUR, lectotype)]. DEScRIPTIos.-Thallus thin, whitish, matte, 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, I + 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 pm. CHEMISTRY.-NO substances present. HABITAT.--('On trees in Roseau Valley." DIscussIos.-The type of Phaeographina elliottii now seems to be completely sterile. Both externally and internally, the type specimen closely resembles Phaeographis albida; it may be possible that Vainio mistook entire disintegrating asci for spores. We have seen no other material that might be refera6le to this Phaeographina species 65. Phaeographina oscitans FIGURE 12f; PLATE l l f Phaeographinn oscitarzs (Tuckerman) Zahlbruckner, 1923:442. Graphis oscitans Tuckerman, 1868:231 [type collection: Oahu, Hawaii, M a r i n s.n. (FH, lectotype)]. NUMBER 40 49 DEScRIPTIoN.-Thallus continuous, smooth to matte, off-white. Ascocarps slightly raised, flexuose, rarely branched, pale thalline margins prominent, 1-2 mm long, slender. Disc prominent, gray-black. Ascocarp transverse section: hymenium 125-1 50 pm high, I-; exciple rudimentary below, yellow 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 pm, IS blue. CHEMISTRY.-stiCtiC acid. HABITAT.-virgin upland rain forest (Dominica only). DIscussIos.-Phaeographjna oscitans is externally very similar to Phaeographis albida, but can easily be separated by spores and chemistry. Also quite similar is Phaeographina pachnodes (Fee) Mueller Argoviensis (1887), which differs in having larger spores, no trace of striae, and in having norstictic acid; and P. turgida (Fee) hfueller Argoviensis (1887), which would seem to be the T L C negative equivalent of P. pachnodes. SPEcnfExs ExAMIsED.-caii-Dom logging area, Pont Casse, 2000 ft, Hale 35127, 35131 (US). Incorrect and Omitted Names Graphis bonplandiae: See Graphis triticea for Elliott 533. Graphis cooperta: See G?-aphis anguilliformis for Elliott s. n. Graphis (Phaeographis) diversa: See Phaeographis exaltata for Graphis dziplicata: See Graphis rirntclosa for Elliott 125 and Graphis dupl ica ta var. subduplicata Vaiiiio: Elliott I509 is Graphis lactea: Elliott 535 is the type of G. lactea var, Graphis lactea Xar. clausa: See Gyaphis insidiosa for Elliott Graphis lineola: See Graphis leptocarpa for Elliott 1535. Graphis (Phaeographis) lobata: T h e only Dominican collec- tion (Elliott 1365 in T U R ) is correctly identified to species, but it is not included in our list because the generic status is unclear. I t is probably closer to Phaeotrenza in the Thelotremataceae. Graphis (Phaeographis) inedicsaeformis: Elliott 529 in B M is a fragmentary specimen, which may be an immature Phaeogr-aphis esaltata. There is no specimen so identified in TUR. Graphis (Graphina) sttbnitida: Elliott 171 is indeterminable. Graphis tenella \ a r . epiphaea I?ainio: Not found in T U R or Graphis (Graphiiia) verrnicularis: See Graphina ru fopal l ida Elliott 526. Graphis flesibilis for Elliott 3954 pro parte. sterile. doiiiinicana T?ainio; see Graphis tachygrapha. 1511. BM and presumed to be lost or misfiled. for Elliott 1852. Literature Cited Acharius, E. 1810. Lichenographia universalis. 696 pages. Gottingae. 1814. Synopsis methodica lichenum. 392 pages. Lund. 1890. A Further Enumeration of Some Lichens of the United States. Bulletin of the Torrey Botanical Club, Eckfeldt, J. W. 173255-257. Eschweiler, F. G. 1828. 1833. Fke, A. 1824. 1837. 1874. Lichens. I n C.F.P. Martius, Icones Plantarum Cryp- togamicarum. 138 pages. Monachii. Lichens. In C.F.P. Martius, Flora Brasiliensis, vol- ume 1, 390 pages. Stuttgart. Essai sur les cryptogames des ecorces exotiques of- ficinales. 167 pages. Paris. Essai sur les cryptogames des ecorces exotiques, 11: Supplement and rekision. 137 pages. Paris. Materiaux pour une flore lichenologique du Bresil, 11: Les Graphidkes. Bulletin de la Societe Botanique de France, 21:21-32. Fink, B. 1927. Yew Species of Lichens from Porto Rico. Mycologia, 19:206221. Groenhart, P. 1965. Studies in Ascostromatic Lichen-Fungi, I: The Problem of Ascohymeniales and Ascoloculares. Per- soonia, 4: 1-7. Morden-Smithsonian Expedition to Dominica: The Lichens (Parmeliaceae). Smithsonian Contributions to Botany, 431-25. Morden-Smithsonian Expedition to Dominica: T h e Lichens (Thelotremataceae). Smithsonian Contribu- tions to Botany, 16:l-46. 1976. Lichenes Americani Exsiccati, no. 156. Washington, D.C. Hayward, G. T h e Lichen Families Graphidaceae and Opegrapha- ceae in New Zealand. New Zealand Journal of Botany [In press.] Sew Genera and Species of Lichens from the Her- barium of Bruce Fink, I . Mycologia, 25:303-316. Hale, M. E., Jr . 1971. 1974. 1978. Hedrick, J. 1933. Hooker, J. D. 1867. Handbook of the New Zealand Flora. 798 pages. London. Knight, C. 1884. On the Lichenographia of New Zealand. Transac- tions and Proceedings of the New Zealand Institute, 16:400-408. Contributions to the Lichenographia of New Zea- Knight, C., and W. Mitten 1860. land. Transactions of the Linnean Society of London, 23: 101-106. Krempelhuber, A. 1876. Lichenes Brasilienses, collecti a D. A. Glaziou in provincia brasiliensi Rio Janeiro. Flora, 59:378-384, 412-4 16. Magnusson, A. H. A Catalogue of the Hawaiian Lichens. Arkiv for Botanik, 3( 10):223-402. Memorie Lichenographiche. 185 pages. Verona. Troisieme Centurie de Plantes cellulaires exotiques nouvelles. Annales Sciences Naturelles Botanique, series 2, 18:241-282. Cryptogamis Guyanensis seu Plantarum Cellularium in Guyana Gallica Annis 1835-1849 a C1. Leprieur Collectarum Enumeratio Universalis. Annales Sci- ences Naturelles Botanique, series 3, 16:47-81. 1855. Lichenes, Collemaceae. I n F. Junghuhn, Plantae Junghuhnianae sive Enumeratio Plantarum quas in insulis Java et Sumatra, pages 427-494. 1880. Lichenologische Beitrage, X. Flora, 63:40-45. 1882. Lichenologische Beitrage, XI?. Flora, 65:381-386. 1887. Graphideae Feeanae. Memoires de la Societede Physique et d?Histoire Naturelle de Geneve, 29:3-80. 1888a. Lichenologische Beitrage, XXX. Flora, 71:528-552. 1888b. Revision Lichenum Eschweilerianorum. Flora, 71: 1888c. Lichenes Paraguayenses a cl. Balansa lecti. Revue 1888d. Lichenes Paraguayenses. Revue Mycologique, 10: 1888e. Lichenes Paraguayenses. Revue Mycologique, 10: 1891. Lichenes. I n T. Durand and H. Pittier, Lichens Primitiae Florae Costaricensis. Bulletin de la SocittC Royale Botanique Belgique, 30:49-97. 1955. Massalongo, A. 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Planchon, Pro- dromus Florae h?ovo-Granatensis ou Enumeration des Plantes de la Nouvelle-Grenade. Annales Sci- ences Naturelles Botanique, series 4, 20:228-382. 1866. Collectio Lichenum ex Insula Cuba. Flora, 49:289- 295. 1868. Synopsis Lichenum Sovae-Caledoniae. Bulletin de la Societe Linne?enne de Normandie, (2)2:39-140. 1889. Lichenes Insularurn Guineensium. 54 pages. Paris. 1890. Lichenes Japoniae. 122 pages. Paris. 1891. Sertum Lichenaeae Tropicae e Labuan et Singapore. 45 pages. Paris. Die Graphidineen der ersten Regnell?schen Expedi- tion nach Brasilien, 1892-94, 11: Graphina und Phaeographina. Arkiv for Botanik, 26A(1): 1-105. 1935. Die Graphidineen der ersten Regnell?schen Expe- dition nach Brasilien, 1892-94, 111: Graphis und Phaeographis, nebst einem Nachtrage zu Graphina. Arkiv for Botanik, 27A(2):1-103. Report on the Lichens of St. Thomas and St. Jan. I n N. L. Britton, T h e Flora of the American Virgin Islands. Brooklyn Botanic Garden Memoirs, 1: 109- 116. Redinger, K. 1933. Riddle, L. W. 1918. Smith, J. E., and J. Sowerby Sprengel, K. 1807. English Botany. \.olume 26, plate 1812. Systema Vegetabilium. Volume 4, edition 16, part 2, 410 pages. Gottingae. plants. Proceedings of the American Academy of Arts and Science, 7: 143-235. Synopsis of North American Lichens, Part 11. 176 pages. New Bedford. Etude sur la classification naturelle et la morpho- logie des Lichens de Bred . Acta Societatis pro Fauna et Flora Fennica, 7(7): 1-256. 1896. Lichenes Antillarum a W. R. Elliott collecti. Journal of Botany British and Foreign, 34:204-266. 1899. Lichens novi rarioresque, 111. Hedwigia, 38:(253)- (259). 1907. Lichenes novi rarioresque, IV. Hedwigia, 46: 168-181. 1909. Lichenes. In J. Schmidt, Flora of Koh Chang, Part IX. Botanisk Tidsskrift, 29:104-151. 1915. Additamenta ad lichenographia antillarum illustran- dam. Annales Academiae Scientiarurn Fennicae, 1921. Lichenes insularum Philippinarum, 111. 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Index (Synonyms in italics) Arthonia caesiopruinosa, 46 Diorygma insculpta, 38 Fissurina incrustans, 38 insidiosa, 16 nitidescens, 38 virginalis, 36 acuminata, 32 adscribens, 31 aibonitensis, 34 albostriata, 18 analoga, 42 antillarum, 31 babingtonii, 38 balbisii, 34 balbisii var. monospora, 33 bipartita, 33 carneoviridis, 33 chlorocarpa, 33 cleitops, 44 colliculosa, 34 collospora, 36 collosporella, 44 columbina, 36 confluens, 41 cymbergrapha, 48 dealbata, 18 dehiscens, 38 deserpens, 33 dimidiata, 36 dimorphodes, 18 dispersa, 37 disserpens, 41 elongata, 33 elongatoradians, 41 erythrella, 34, 43 frumentaria, 37 glaucoderma, 38 heteroplacoides, 18 illinata, 37 incrustans, 38 insculpta, 38 insidiosa, 16 insignis, 43 intortula, 34 intricata, 40 iuturgescens, 31 macella, 38 Graphina acharii, 31 marcescens, 40 melaleuca, 44 monophora, 45 nitidescens, 38 nuda, 40 obtectula, 44 palmeri, 44 parilis, 33 platycarpa, 41 platycarpinu, 41 platycarpoides, 32 platyleuca, 41 plittii , 40 plurispora, 41 pseudoanaloga, 42 pseudosophistica var. plurispora, 41 riopiedrensis, 34 rufopallida, 42 ruiriana, 41 suberythrella, 42 subnitidula, 22 substriatula, 41 subvelata, 42 sulcata, 32 sulcatula, 41 triangularis, 44 triphora, 44 triphoroides, 44 vermicularis, 42 vestitoides, 44 virginalis, 36 virginea, 44 Graphis acharii, 31 acuminata, 32 adpressa, 10 adscribens, 31 afzelii, 10 alb ida , 25 anfractuosa, 10 anguilliformis, 10 angustata, 15 angustata var. denudata, 15 antillarum, 31 antillarum var. ingarum, 15 arthonioides, 27 beaumontii, 16 bonplandiae, 24, 49 caesiella, 19 caesioglauca, 15 calcea, 15 candidissima, 40 chlorocarpa, 33 cinerea, 15 colliculosa, 34 collospora, 36 collosporella, 44 coiumbina, 36 congesra, 15, 20 cooperta, 12, 49 dehiscens, 38 desquamescens, 12 dimidiata, 36 ditersa, 49 dumastii, 24 dumastioides, 12 duplicata, 21, 49 duplicata var. subduplicata, 49 dussii, 13 elegans, 13 elliottii, 48 endoxantha, 21 eugeniae, 34 flavicarrs, 19 flexibilis, 13 frunientaria, 31 furcata, 19 glaucescens, 15 glaucoderma, 38 glauconigra, 20 grammitis, 15 haematites, 27 humilis, 16, 24 illinata, 37 imshaugii, 16 inquinata, 24 insculpta, 38 inszdiosa, 16 interuersa, 16 intortula, 34 intricata, 12, 40 tnturgescens, 31 isidiifera, 18 lactea, 24 lactea var. clausa, 16, 49 lactea var. dominicana, 23 leptocarpa, 18 52 NUMBER 40 53 librata, 19 lineola, 19,49 lobata, 49 longula, 19 lumbricina, 20 marcescens, 40 marginifera, 25 medusaeformis, 49 nitidescens, 38 olivacea, 20 oscitans, 48 pavoniana, 19 platy carpoides, 32 platyleuca, 41 proserpens, 22 pseudoanaloga, 42 rigidula, 21 rimulosa, 21 rosea, 28 rubiginosa, 38 rufopallida, 42 seminuda, 12 striatula, 21 subamylacea, 16 subelegans, 21 subnitida, 49 subnitidula, 22,24 subtigrina, 29 tachygrapha, 23 tenella, 19 tenella var. epiphaea, 49 timida, 23 tongloensis, 25 triticea, 23 tumidula, 12 tumidulella, 15 turgidula, 24 vermicularis, 42, 49 virginalis, 36 Lecanactis exaltata, 27 Leiogramma virgineum, 44 Opegrapha desquamescens, 12 elegans, 13 rhlizocola, 12 rimulosa, 21 atrovermicularis, 46 caesiopruinosa, 46 caracasana, 48 chrysocarpa, 28 Phaeographina arechavaletae, 48 columbina, 36 coriaria, 46 difformis, 48 elliottii, 48 elliptica, 48 explicans, 48 includens, 18 intercedens, 46 oscitans, 48 pachnodes, 49 turgida, 49 ceruiculata, 44 cinnabarina, 28 decipiens, 27 dimorpha, 29 dussii, 13 exaltata, 27 haematites, 27 longula, 19 mordenii, 28 rosea, 28 sexloculata, 27 subtigrina, 29 Phaeographis albida, 25 Sclerophyton colliculosum, 34 54 Shl ITHSONIAN C O S T R I B U T I O S S T O B O T A h Y PLATE 1.-Species of Graphis: a, G. adpresstc (Hale 35487); b, G. afieiii (Irnshaug 33489'4); c, G. nnfractuosa ( W i r t h 538); d , G. anguillifornzis (Hale 35282); e, G. desqztamescens (Hale 33449); f, G. dumastioidrs (Hale 37722). (Plates 1-10 x 10.) NUMBER 40 PLATE 2.-Species of Graphis: a, G . dussii (Hale 35443); b, G. elegans (Hale 35476): c, G . pexibi- lis (Zmshaug 33360); d, G. glaucescens (Zmshaug 33313A); e, G . grammitis (Hale 38005); f, G . humilis (Hale 35698). 55 56 SMITHSOSIAN CONTRIBUTIOSS TO BOTANY PLATE 3.-Species of Graphis: a , G. irnshaugii (Imshntig 32724 i n MSC): b, G. insidiosa (Hale 35313); c, G. isidiifera (Hale 35179); d , G. leptocarpn (Imshatig 33109.A); e, G. librata ( W i r t h 538); f . G. Eongula (Hale 35225). NUMBER 40 PLATE 4.-Species of Graphis: a, G . lumbricina (Hale 35240); b, G . olivacea (Hale 37988); c, G . rigidula (Imshaug 32777A); d , G . rimulosa (Hale 35699); e, G . subelegans (Hale 35546); j , G . subnitidula (Hale 37676). 57 58 SMITHSONIAS C O S T R I B U T I O N S T O BOTANY PLATE 5.--Species of Graphis and Phaeographis: a , G. tachygmpha (Imshaug 32862A); b, G. triticea (Hale 35324); c, G. turgidula (Wirth 544b); d, P . aibida (Hale 35453); e, P . arthonioides (Imshaug 33128); f, P. exaltata (Hale 35301). NUMBER 40 PLATE B.-Species of Phaeographis and Graphina: a , P. exaltata (Hale 35321); b, P. haematites (Hale 38013); c, P. mordeni i (Ha le 35071); d , P. rosea (Elliott in BM); e, P . subtigrina (Wir th 444); j , G. acharii (Hale 35555). 59 60 S M I T H S O S I A S C O S T R I B U T I O N S TO BOTASY PLATE 7.--Species of Graphina: a , G . adscribens (Wirth 519); b, G. antillaruni (Hale 35745); c, G. carneoviridis (Hale 37642); d, G. chlorocarpa (Hale 35129); e, G. colliculosa (Hale 35175); f, G . collospora ( W i r t h 520). NUMBER 40 PLATE &-Species of Graphina: a, G. columbina (Wirth 446); b, G . dimidiata (Hale 35697); c, G. dispersa (Hale 38090); d, G. frumentaria (Hale 35242); e, G. illinata (Hale 35284); f , G. incrustans (Wirth 465). 61 62 SMITHSOXIAN COSTRIBUTIONS TO BOTASY PLATE 9.--Species of Graphina: a, G. insculpta (Hale 37819); b, G. macella (Hale 35723); c, G. marcexens (Hale 35312); d , G. nuda (Hale 35119); e, G. platyleuca (Hale 37955); j , G . plurispora (Imshaug 37724.4). NUhlBER 40 PLATE 10.-Species of Graphina: a , G. pseudoanaloga (Hale 35283): b, G. rufopallida (Elliott 1832); c, G. suberythrella (Zmshaug 32699); d, G. triphoroides (Hale 35161); e, G. vestitoides (Zmshaug 32859A); f, G. virginea (Wir th 547). 63 64 SMITHSONIAN CONTRIBUTIOSS T O BOTANY PLATE 1 1.-Species of Phaeographina: a, P . atrouermictilaris (Hale 35278); b, P. raesiopruinosa (Znzshaug 32713); c, P . coriaria (Hale 35378); d, P. dioormis (Zrnshaug 32913.4); e, P . elliottii (EIEiott 1338 in TUR); f, P . oscitans (Hale 35131).