SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 25 A Revision of the Lichen Genus Hypotrachyna (Parmeliaceae) in Tropical America Mason E. Hale, Jr. SMITHSONIAN INSTITUTION PRESS City of Washington 1975 ABSTRACT Hale, Mason E., Jr. A Revision of the Lichen Genus Hypotrachyna (Parmelia- ceae) in Tropical America. Smithsonian Contributions to Botany, number 25, 73 pages, 20 figures, 1975.-A revision is made for 77 species of Hypotrachyna occurring in tropical America. Fifty-eight of these are endemic to the New World. These species are classified loosely in chemical constellations based on the secondary depside-depsidone products. The possible role of hybridization of fertile species and the evolution of vegetative morphs are discussed. Five new species are described: H. andensis, H. lopezii, H. partita, H. producta, and H. protenta. The remaining 72 species are transferred from the genus Parmelia to Hypotrachyna. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution?s annual report, Smithsonian Year. SI PRESS NUMBER 5273. SERIES COVER DESIGN: Leaf clearing from the katsura tree Cercidiphyllurn japonicum Siebold and Zuccarini. Library of Congress Cataloging in Publication Data. Hale, Mason E. A revision of the lichen genus Hypotrachyna (Parmeliaceae) in Tropical America. (Smithsonian contributions to botany, no. 25) Includes index. Supt. of Docs. no.: SI 1.29:25 1. Hypotrachyna. 2. Lichens-Latin America. I. Title. 11. Series: Smithsonian Institution. QK1 .S2747 no. 25 [QK585.P2] 581?.08s [589?.1] 74-32050 Smithsonian contributions to botany, no. 25. Contents Introduction ...................................................... History of Hypotrachyna Research ............................ Collections ........................................... Morphological Characters .......................................... Chemical Characters ............................................... Speciation in Hypotrachyna ........... ........ ....... Classification of Hypotrachyna .... .................... .. The Hypotrachyna Flora ................................ .... Habitats and Ecology .............................................. Phytogeographical Relationships .................................... Nomenclature ..................................................... Keys to the Species ................................................ Species Treatment ................................................. Literature Cited ..................................... Index ....... ..................... Page 1 2 3 3 8 11 15 16 17 18 19 19 23 69 72 ... 111 A Revison of the Lichen Genus Hypotrachyna (Parmeliaceae) in Tropical America Mason E. Hale, Jr. Introduction I began study of the Parmeliaceae in 1958 with the realization that synonymy and nomenclature of the nearly 600 species in the family could be solved only through a world-level revision. A monograph of the relatively conservative genus Parmotrema Massalongo (as Parmelia subgenus Amphigymnia) was completed in 1964 (Hale, 1965) along with a preliminary study of the other tropical sections of the collective genus Parmelia (Hale and Kurokawa, 1964). Although I had examined all the type specimens and a considerable body of herbarium material, two factors mitigated against a prompt completion of a final monograph for all groups: (1) the inade- quacy of chemical techniques available prior to 1965 that were needed to resolve certain chemical groups such as the barbatic and lividic acid groups and even separation of the norstictic acid group; and (2) the lack of sufficient specimens and field experience in tropical America, where many new species were obviously yet to be found. The development and standardization of thin- layer chromatography has answered the problem of chemical techniques (see C. Culberson, 1972a). We can now routinely achieve resolution heretofore impossible and make accurate identifications of the lichen substances, A number of previously un- Mason E. Hale, Jr., Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. known substances have been isolated and described. The few remaining problems will eventually be solved with high-performance liquid chromatog- raphy and mass spectrometry (C. Culberson, 1972b). The inadequacy of collections will probably never be fully remedied, but we now have access to many more collections than were available in 1965. I have had the opportunity of visiting several tropical areas where I could appreciate more fully the habitats and ecology of the species, data which are only rarely well documented on herbarium labels. The time has come, therefore, when a "final" summary can be made for the parmelioid genera. This paper summarizes the genus Hypotrachyna (Vainio) Hale for tropical America, including the region from the West Indies and Mexico south- ward to temperate Argentina and Chile. The 19 species known from North America proper are not included, but actually only one (H. vz'rginka) is endemic there and the remaining 18 also occur in tropical '4merica. ACKNOWLEDGMENTS.-~ wish to reiterate thanks to the curators-previously acknowledged in Hale (1965) and Hale and Kurokawa (1964)-who SO generously loaned and in many cases reloaned valuable specimens. Special thanks are due Dr. S. Kurokawa, whose collaborative help in 1960-1964 speeded up study of hundreds of type specimens. I am extremely grateful to Dr. M. Lopez Figueiras of Universidad de 10s Andes for the opportunity 1 2 SMITHSONIAN CONTRIBUTIONS TO BOTANY to collect extensively with him in the Venezuelan Andes. Mr. Eric Lewis, Ms. Marianne See, and Dr. Michael Wirth helped at various times with test- ing of specimens by thin-layer chromatography. The scanning-electron microphotographs were taken by Mr. Walter Brown of the Smithsonian Scanning-Electron Microscope Laboratory. Field studies in Mexico (1960) were supported by the National Science Foundation, and in Dom- inica by the Morden-Smithsonian Expedition (Hale, 1971a). The work in the past several years has been supported by grants from the Smithsonian Research Foundation. History of Hypotrachyna Research Studies on Hypotrachyna (and for all other trop- ical Parmelias) appeared only as scattered floristic lists prior to 1890. Krempelhuber, Tuckerman, Muller Argau, Nylander, and others received for identification small lots of lichens (usually collected by phanerogamic botanists) and published the results without illustrations, keys, or really useful descriptions. There was little cooperation between workers or sharing of information or specimens, Vainio (1890) was the first lichenologist to make a truly comprehensive study of tropical lichens. He collected extensively in Rio de Janeiro and Minas Gerais, Brazil, in 1885. The resulting publication, ?Etude sur la Classification Naturelle et la hlorph- ologie des Lichens du Bresil,? remains a classic and Vainio?s well-curated herbarium in Turku and his exsiccate, Lichenes brarilienses-distributed, as far as I have been able to determine, to BM, FH, G, K (now at BRI), RI, P, and UPS-attest his tremen- dous ability and industry. When identifying speci- mens, Vainio used Nylander?s herbarium and also made a study trip to the Paris museum. This en- abled him to present a more definitive study with a low level of synonymy. Vainio collected 13 species now classified in the genus Hypotrachyna: H. brasiliana (Nylander) Hale, H. consirnilis (Vainio) Hale, H. dactylifera (Vainio) Hale, H. flavida (Zahlbruckner) Hale (as Parmelia flava Krempelhuber), H. gracilescens (Vainio) Hale, H. intercalanda (Vainio) Hale, H. rnicroblasta (Vainio) Hale, H. novella (Vainio) Hale, H. plzrriformis (Nylander) Hale, H. revoluta (Floerke) Hale, H. subaflinis (Zahlbruckner) Hale (as Parmelia aflinzs Vainio), and H. velloziae (Vainio) Hale. Material he identified as Parmelia gracilis (Muller Argau) Vainio is misidentified H. physcioides. Altogether, Vainio described eight of these species as new. In spite of his efforts, how- ever, only a portion of the Hypotrachyna species were collected; the general region of southeastern Brazil is now known to have nearly 40 species. The next major collecting program also was undertaken by a lichenologist, G. A. A. Malme, under the Regnell expeditions in the 1890s, pri- marily in Mato Grosso. The magnificently prepared specimens, now preserved in Stockholm, were studied mostly by Malme himself, but Parmelia specimens were turned over to B. Lynge, a Norwe- gian lichenologist who had no previous experience in tropical species, and this at a time when Vainio was the most logical worker and acknowledged authority. Lynge, however, did a creditable job, dealing with 110 species, of which 61 were proposed as new (Lynge, 1914, 1917, 1925). Only 18 of these species are now classified in Hypotrachyna. Lynge relied heavily on Vainio?s exsiccate preserved at UPS, saw some Nylander types, and visited Zahl- bruckner in Vienna. His publications contain much valuable data and some excellent illustrations. The only other work that could be considered as a flora with keys and descriptions is that by Zahl- bruckner (1909), completed just before Lynge?s study. It was based on collections made by a non- lichenologist, V. Schiffner, in the Sao Paulo area. The specimens are preserved at WU and W with duplicates at BM. Nore recently, Vareschi (1962) compiled a cata- log of Venezuelan Pal-meliac Fvith many illustra- tions, but the species identifications have not been checked b) a monographer. My own field experience in tropical America began in Mexico, which has a large Hypotrachyna flora. I have also collected intensively in the Lesser Antilles, Trinidad, Guatemala, Panama, and Vene- zuela. These excursions have given me some idea of the ecological behavior of the genus from sea level to 3700 m, and while the area covered is but a small fraction of the total land mass of tropical America, it seems reasonable to extrapolate from this experience in discussing the habitats where the species might be expected to occur. NUMBER 25 3 Collections Any revision is only as good and complete as the specimens available for study. Tropical America is a vast area, some parts of which have been rela- tively well explored, others are completely unknown and often inaccessible. Even a country as seemingly small as Panama has great tracts of unexplored rain and cloud forest that cannot be reached except by a major expeditionary effort. On the other hand, some areas, while uncollected, would probably have few species of Hypotrachyna. Such areas include, in particular, the Amazon Basin, which has a gen- erally low elevation and a hot moist climate, hardly conducive to a rich foliose lichen flora. Deserts or very dry regions as in northern Chile, northern Mexico, and Baja California, and even north- eastern Brazil, also are poor prospects for Hypotrachyna. The major areas of virgin forest in the tropics are, of course, being destroyed-cut over, burned, and colonized-as quickly as they can be reached. Such areas include even the higher elevations where Hypotrachyna is so abundant but where the soils are ideal for vegetable farming. The following list shows the countries where Hypotrachyna has been collected, the principal col- lectors, and the herbaria where their collections are housed: BAHAMAS: Britton (XY). MEXICO: Arsene (US), Galeotti (P), Hale (US), Wirth (US). WEST INDIES (all islands): Imshaug (ILISC). CUBA: Hioram (US), Jt?right (FH, L, P, UPS, US). JAMAICA: Morton (US), Orcutt (US), Plitt (US). DOMINICAN REPUBLIC: Allard (US). VIRGIN ISLANDS: Raunkiaer and Boergesen (C, TUR). GUADELOUPE: Culberson (DUKE), Duss (TUR), Husnot (H, DOMI~ICA: Elliott (BM, TUR), Hale (US). ST. VINCENT: Elliott (BM, TUR). TRINIDAD: Hale (US). GUATEMALA: Hale (US), Standley (F, US). HONDURAS: Standley (F). COSTA RICA: Culberson (DUKE), Pittier (G, US). PANAMA: Hale (US), Scholander (US). COLOMBIA: Cuatrecasas (US), Killip and Smith (US), Lindig VESEZUELA: Dennis (BM), Hale (US), Magdefrau (M), Poelt FRENCH GUYANA: Leprieur (P), PERU: Bryan (WIS), Gomez (US), Herrera (US), Iltis (TYIS). ECUADOR: Jameson (BM), Meyer (W). BOLIVIA: Mandon (H). P). (BM [original], FH, H, M, P), Weir (BM). (M, US), Santesson (S), Vareschi (VEN). CHILE: Gay (P), Imshaug (MSC), Mahu (personal, US), San- tesson (S, UPS), Skottsberg (S). BRAZIL: Damazio (G), Deventer (G), Eiten (US), Glaziou (G, P), Hohnel (W), Irwin (NY, US), Kurokawa (TNS; speci- mens not available for study), Malme (S), Puiggari (G), Reitz and Klein (US), Schiffner (W, WU), Spruce (BM, H, NY), Ule (G), Vainio (TUR [original], BM, FH, M, P, UPS), Weddel (P, H), Warming (M). PARAGUAY: Balansa (G), Malme (S). URUGUAY: Felippone (W), Herter (H), Hosseus (H), Osorio ARGENTINA: Arechavaleta (G), Balansa (G), Dusen (S), Lamb (MVM). (CAN), Lorentz and Hieronymus (M). Noticeable gaps remain in Bolivia, Ecuador, Peru, Venezuela outside the Andes, and areas of Brazil bordering these countries. The unevenness of col- lecting can also be appreciated by examining lists of species collected in each country (see p. 18). I have not attempted to evaluate literature re- ports of species in floristic lists (exclusive of type species) other than to examine the herbarium collections on which these were based and annotate them with revised names as needed. The sheer number of collections involved discourages one from tracing, verifying, and correcting each pub- lished name. Because of the extensive synonymy, lack of adequate chemical testing, and the gen- erally low level of accuracy in identification, older published reports are quite untrustworthy. Morphological Characters Hypotrachyna is a segregate of the collective genus Parmelia characterized by narrow, apically truncate lobes, a black lower surface, and dicho- tomously branched rhizines (Hale, 1974a). Hale and Kurokawa (1964) had earlier discovered that presence or absence of marginal cilia, type of cilia (normal or bulbate), and branching pattern of rhizines were the most consistent characters that could be used to subdivide the more than 400 species of Parmelia. The segregate Hypotrachyna is one of the more clear-cut groups in the Parmeliaceae because of the distinctive rhizine branching. INTERNAL SmucTuRE.-Little attention has been paid to differences in internal structure of Parmeliae and how they might aid in a more natural classi- fication of the genera. Tlihile carefully sectioning the Brazilian collections, Lynge (1914) noted that the species related to Parmelia microsticta had a rather thick paraplectenchymatous cortex in con- 4 SMITHSONIAN CONTRIBUTIONS TO BOTANY trast to a palisade plectenchymatous cortex found in Hypotrachyna species. RIy recent work with the scanning-electron microscope (SEM) showed that all species of Hypotrachyna so far examined (Hale, I97Za) have a palisade cortex that is overlain by a thin, pored polysaccliaride epicortex (Figure la) which cannot be seen with light microscopes. It is the cortex, along with dichotomously branched rhizines, that I use to distinguish Hypotrachyna. Parmelia sensu strict0 has a paraplectenchymatous cortex without a pored epicortex, as in the type- species of the genus, P. saxatilis (L.) Acharius. The cross sections of Hypotrachyna are con- sequently rather uniform from species to species. The palisade cortical cells form a layer 12p-181i thick. The TreDoztxia-containing algal layer is 12~-2411 thick, the medulla 50y-150p thick, and the paraplectenchymatous lower cortex 12p-18p thick (Figure 10). In general, very small species (2-5 cm in diameter) are quite thin (medulla 50p-60p), FIGURE 1.-Morphology of Hypotrachyna: a, surface of H. croceopustulata (Imshaug 22275) (X 500 with SEM); b, cross section of H. croceopustulata (Hale 33269) (X 500 with SEM); c, rhizines of H. densirhizinata (Scholander) (X 100 with SEM); d, cross section of thallus and a soralium of H. densirhizinata (Hale 43194) (X 75 with SEM). NUMBER 25 5 while larger species (5-25 cm in diameter) have proportionately thicker layers. The rhizines con- sist of conglutinated strands of prosoplectenchyma- tous hyphae and are moderately to richly dicho- tomously branched (Figure lc). LOBE CONFICURATION.-HYPO~YUC~~~U is extreme- ly homogeneous in overall lobe configuration. It is possible, however, to recognize two broad types in tropical America: an extremely elongate, divaricate, and little-branched type as in H. caraccensis (Figure 8a) or M. lopezii (Figure 13c), and a shorter-lobed, more or less closely branched and imbricate type exemplified by H. Zivida (Figure 13b). There are, of course, numerous intermediates between these two extremes, as in H. laeuigata (Figure 120. A third type of configuration is found in the very rare H. koyaensis, which has the broader, almost subirregular lobes (Figure 1%) characteristic of many species of Hypotrachyna endemic to Asia. SoREDIA.-These vegetative diaspores originate in FIGURE 2.-Vegetative structures of Hypotrachyna (all X 10): a, diffuse soralia of H. revoluta (Hale 43293a); b, orbicular soralia of H. sinuosa (Hale 43071a); c, pustulate soralia of H. croceopustulata (Hale 33269); d, pustules of H. dactylifera (Hale 42075). SXIITHSONIAN CONTRIBUTIONS TO BOTANY soralia (Figure Id) and erupt as a powder. The shape and location of the soralia on lobes are im- portant taxonomic characters used to recognize species. Soredia occur in 18 of the 77 species of tropical American Hypotrachyna: H. breuirhiza, H. croceopzistulata, H. densirhizinata, H. endo- chlol-a, H. exsplendens, H. pavovirens, H. gondylo- phora, H. immaculata, H. laeuigata, H. oostingii, H. prodzicta, H. pseudosinuosa, H. revoluta (Fig- ure 2a), H. rockii, H. sinuosa (Figure 2b), H. sub- affinis, H. thpsanota, and H. velloziae. They are generally subterminal to rarely lamina1 and pro- duced in orbicular soralia. Soralia rarely become diffuse, as in H. reuoluta. In a few species pustules are formed initially but later become sorediate, as in H. croceopustulata (Figure 2c). Details of ab- normal soredial formation will be found in the species discussions. PusTuLEs.-These are coarse, inflated, extremely fragile, isidialike structures (Figures Zd, 3a), but they are related to soralia since they often become sorediate. True pustules are hollow and tend to FIGURE 3.-Vegetative structures of Hypotrachyna: a, pustules of H. formosana (Eiten 7671a) (X 60 with SEM); b, enlargement of pustule in a (X 300); c, cross section of thallus and isidia of H. imbricatula (Quiros 1479) (X 100 with SEM); d, enlargement of isidia in c (X 300). FIGURE 4.-Vegetative structures of Hypotrachyna: a, cross section of an isidium of H. costari- censis (Hale 42201) (X 1000 with SEICI); b, lohulate isidia of H. andensis (Hale 42658a) (X 100 with SEM); c, lobulate isidia of H. bogotensis (Oberwinkler and Poelt 7544) (X 10); d, lobules of H. ~achista (Hale 43139) (X 10); e, lobules of H. chlorina (Hale 43104) (X 10); f, maculae on a lobe of H. subplumbeata (Moore and Wood 4944) (X 10). 8 SMITHSONIAN CONTRIBUTIONS TO BOTANY break open apically without soredial formation. Only three species of Hypotrachyna treated here have this type of pustule: H. dactylifera, H. for- mosana, and H. malmei. Under SEM the surface of pustules is quite rough with loosely aggregated hyphae lacking an outer polysaccharide layer (Figure 3b). More work needs to be done on these structures. IsIDIA.-Fingerlike, cylindrical isidia, densely filled with medullary hyphae and about 0.1 mm in diameter, are well-known lichen structures (Fig- ures 3c, 3d). They occur in 16 species of Hypo- trachyna: H. andensis, H. bogotensis, H. con- sirnilis, H. costaricensis, H. dentella, H. ensifolia, H. imbricatula, H. isidiocera, H. koyaensis, H. microblasta, H. neodissecta, H. osorioi, H. partita, H. rhabdiformis, H. steyermarkii, and H. sub- physodalica. Isidia are often presented in textbooks as con- taining extensions of thalloid tissues. When viewed with the scanning-electron microscope, however, we see little internal organization (Figure 4a). There is no true cortical structure and no pored epicortex, only a thin, smooth, continuous polysaccharide sheet covering the weakly organized mass of algae and hyphae inside. While most isidia show little variation except in branching, lobulate isidia sometimes are found to occur in six of the above-named species: H. anden- sis, H. bogotensis, H. ensifolia, H. imbricatula, H. partita, and H. steyermarkii. In such cases the isidia begin normally but soon become procumbent and even dorsiventral, as in true lobules (Figure 4b,c). Cilia will rarely form (cf. H. steyermarkii). When viewed with the SEM, the lobulate isidia appear to have the same level of organization as isidia, lacking, for example, a distinct cortex. LoBuLEs.-Almost any species of Hypotrachyna may develop small adventitious lobes with age. Such lobes can be regarded as regenerative struc- tures with internal structure and a pored epicortex comparable to the main lobes when viewed under the scanning-electron microscope. As defined here, lobules are a consistent species character, abun- dantly or distinctively produced and not associated with isidia. Four species have them: H. chlorina (Figure 4e), H. prolongata, H. rachista (Figure 4d), and H. singularis. I do not mean to imply, however, that these lobules are identical in origin and formation. MAcuLAE.-These are mottled, light to dark patches in the cortex visible at X 10 magnification (Figure 4f). They are not detectable as surface features under the scanning-electron microscope but seem to result from darker clumps of algae contrasted with the lighter algae-free hyphal matrix. Although some species of Hypotrachyna are always maculate, this is a difficult character to determine in practice and I have avoided using maculae as a primary character. APOTHECIAL CHARACTERS.-Apothecia have been found in 63 species of Hypotrachyna but so far are unknown in H. exsplendens, H. gondylophora, H. lineariloba, H. monilifera, H. oostingii, H. partita, H. physodalica, H. producta, H. prolongata, H. rachista, H. steyermarkii, H. subphysodalica, H. thysanota, and H. velloziae. Apothecia are present, but no spores developed, in H. croceopustulata, H. laevigata, H. protoboliviana, and H. subplumbeata. Apothecia are uniformly small, 1-5 mm in diam- eter (rarely to 10 mm), adnate (stipitate only in H. peruviana), and imperforate. The hymenium is 40~-60~ high. The unicellular spores show a very small range in size, usually 4p-lop wide and 8y-16p long. A few species, notably those with yellow pig- ments (as H. endochlora) and H. pluriformis have significantly larger spores, up to 25p long, but no species of Hypotrachyna has spores more than 30p long. Pycnidia are often produced by apothecia-bearing species. The microconidia are about 1p wide and 5p-7~ long in all species investigated. I have not included them in the descriptions since they seem to have no taxonomic value. Articles by Nylander, Lynge, and Vainio can be consulted for further data on pycnidia and microconidia in various species. Chemical Characters Beginning with Nylander in the 1860s and continuing with Vainio, Zahlbruckner, and Lynge, color tests with KOH and Ca(OC1)2 have played an important role in identifying species of Parmelia, and information on the tests always has been in- cluded in the species descriptions. Only Miiller Argau refused to use chemistry, even though it was shown to be a useful aid in identification. It was Asahina in the few years from 1933-1940 who perfected the microcrystal tests by which NUMBER 25 9 specific lichen substances could be distinguished. This meant, for example, that a KOH+ red species could be shown to contain norstictic acid, salazinic acid, or other component. The next significant advance in chemical tech- niques was partition chromatography, first with paper (Wachtmeister, 1952; Mitsuno, 1953) and later on silica gel (thin-layer chromatography or TLC) (Stahl and Schorn, 1961). TLC is now the accepted standard for routine qualitative identifi- cation of lichen substances and there is little chance it will be replaced by a more sophisticated technique in the near future. High-performance liquid chromatography (C. Culberson, 1972b) is and will remain far too expensive to be widely available to taxonomists. My original chemical analyses of Parmelia speci- mens were accomplished with microcrystal tests well before 1965 when TLC was just coming into use. Many of these specimens were returned to herbaria from which they had been borrowed, and it has been necessary in the last few years to borrow them again (especially types) and perform TLC tests. Unfortunately, some which I annotated be- tween 1958 and 1964 were not rechecked. All material examined since 1965 has been studied with TLC. Culberson?s methods have been used exclusively (C. Culberson, 1972a). I have also benefited from being able to consult with Dr. Culberson on numer- ous occasions on the identification of difficult or unknown acids. Following is an alphabetical list of all identifiable substances, the species that contain them, and associated substances, that is, those with which they may occur. Unknowns are given at the end. Further details on joint occurrences of substances, accessory substances, etc., are given in discussions of the species. Alectoronic acid.-H. degelii, H. densirhizinata, H. ensifolia, H. exsplendens, H. gigas, H. lineariloba, H. rnalrnei, and H. protenta. Associated substances: atranorin, a-collatolic acid, gyrophoric acid, lichexanthone, and protocetraric acid. An additional qualitative test for alectoronic acid is bright white to bluish white fluorescence in longwave ultraviolet. Anziaic acid.-H. ducalis, H. eitenii, H. partita, H. producta, and H. rachista. Associated substances: atranorin, lichexan- thone, and perlatolic acid. Several unknown spots com- monly occur in TLC profiles of anziaic acid. Atranorin.-In all species except H. brasiliana, H. caraccen- sis, H. enderythraea, H. erythrodes, H. flauida, H. pauovir- ens, H. formosana, H. lopezii, H. malmei, H. microblasta, H. minima, H. novella, H. osteoleuca, H. physodalica, H. protoboliviana, H. reducens, H. siluatica, H. subafinis, H. subphysodalica, and H. uelloziae. Associated substances: all substances except lichexanthone and usnic acid. Atranorin is the main cortical substance in Hypotrachyna; it is absent only when replaced by usnic acid or lichexanthone, al- though in a few cases it may be present in trace amounts. Barbatic acid.-H. chlorina, H. dentella, H. endochlora, H. imbricatula, H. isidiocera, H. laeuigata, H. monilifera, H. peruviana, H. physcioides, H. protoboliviana, H. steyer- markii, and X. subplurnbeata. Associated substances: Atran- orin, 4-0-demethylbarbatic acid, obtusatic and norobtusatic acids (constant accessories except in H. subplurnbeata), echinocarpic acid, entothein, usnic acid, and unidentified yellow pigments. A complete discussion of the chemistry of these species is provided by Culberson and Hale (1973). Chromatography of the barbatic group is very difficult; the most promising resolution is obtained with toluene-acetic acid. All specimens usually react C+ orange, a reaction now known to be caused by 4-0-demethylbarbatic acid, not barbatic acid itself. Colensoinic acid.-H. dactylifera, H. erythrodes, H. formo- sana, H. immaculata, H. livida, H. novella, and H. pal- marum. Associated substances: atranorin, 4-0-demethyl- physodic acid, lichexanthone, lividic acid, norcolensoinic acid, physodic acid, rhodophyscin, the ?palmarum? un- known, and traces of other unknowns. See the discussion under lividic acid. a-Collatolic acid.-H. degelii, H. densirhizinata, H.. exsplen- dens, H. gigas, and H. protenta. Associated substances: alec- toronic acid, atranorin, and gyrophoric acid. a-Collatolic acid is not a constant accessory component with alectoronic acid, as one may see by comparing the lists of species. Constictic acid.-H. andensis, H. loperii, and H. sinuosa. As- sociated substances: norstictic acid, salazinic acid, stictic acid, and usnic acid. Constictic acid invariably occurs with stictic acid in Hypothrachyna. Its structure is very close to stictic acid. 4-0-Demethylbarbatic acid.-See barbatic acid. 4-0-Demethylphysodic acid.-See lividic acid. Echinocarpic acid.-H. dentella, H. subafinis, and H. thy- sanota. Associated substances: atranorin, barbatic acid com- plex, gyrophoric acid, lichexanthone, and microphyllinic acid, The structure of this P+ orange-red acid is as yet undetermined. It forms a low discrete H,SO,+ orange spot in hexane-ether and a low, dull gray streak in benzene- dioxane solvent systems. Entothein.-H. chlorina, H. endochlora, H. isidiocera, and H. peruviana. Associated substances: atranorin, barbatic acid complex, obtusatic acid complex, and unidentified yellow pigments. Evernic acid.-H. bogotensis; H. chicitae, H, pulvinata, and H. rockii. Associated substances: atranorin, lecanoric acid, norobtusatic acid, obtusatic acid, and rhodophyscin. Lecan- oric acid is a constant accessory. Culberson and Hale (1973) discuss this group more fully in relation to the species con- SMITHSONIAN CONTRIBUTIONS TO BOTANY taining barbatic acid. They hypothesize that obtusatic acid could be the product of hybridization between evernic acid and barbatic acid. Fumarprotocetraric acid.-H. gondylophora. Associated sub- stances: atranorin, rhodophyscin, and succinprotocetraric acid. Fumarprotocetraric acid would seem to be an ex- tremely rare substance in Hypotrachyna. Galbinic acid.-H. caraccensis, H. enderythraea, and H. microblasta. Associated substances: norstictic acid, salazinic acid, and usnic acid. Galbinic acid was first described from Parmelina galbina (Acharius) Hale where it is a primary constituent. It has also been reported in Usnea. In Hypo- trachyna it always occurs with norstictic and salazinic acids. The molecular structure has recently been determined by Dr. Myles Keogh as an acetate of salazinic acid. Gyrophoric acid.-H. gigas, H. neodissecta, H. oostingii, H. osorioi, H. pluriformis, H. revoluta, and H. thysanota. As- sociated substances: alectoronic acid, atranorin, echinocarpic acid, microphyllinic acid, and unknowns. Gyrophoric acid is an accessory substance only in H. gigas. Lecanoric acid.-H. bogotensis, H. chicitae, H. pulvinata, and H. rockii. Associated substances: atranorin, evernic acid, norobtusatic acid, obtusatic acid, and rhodophyscin. Al- though common as a primary constituent (along with atranorin) in most parmelioid genera, lecanoric acid always occurs in Hypotrachyna with and in lesser concentration than evernic acid (C. Culberson, 1972b). The color test with calcium hypochlorite is therefore weak or indistinct. Lichexanthone.-H. brasiliana, H. eitenii, H. erythrodes, H. formosana, H. malmei, H. minima, H. novella, H. osteo- leuca, H. silvatica, and H. subafinis. Associated substances: alectoronic acid, anziaic acid, atranorin, colensoinic acid, 4-0-demethylphysodic acid, echinocarpic acid, lividic acid, norcolensoinic acid, olivetoric acid, physodic acid, proto- cetraric acid, rhodophyscin, and unknown pigments. Liche- xanthone is a cortical substance which fluoresces brilliant yellow-orange under longwave ultraviolet. It completely re- places atranorin and usnic acid in the cortex, although traces of atranorin may accompany it in a few species. It is noteworthy that no Hypotrachyna species endemic to the Old World produces lichexanthone. Lividic acid.-H. dactylifera, H. erythrodes, H. formosana, H. irnmaculata, H. livida, and H. novella. Associated sub- stances: atranorin, colensoinic acid, 4-0-demethylphysodic acid, lichexanthone, norcolensoinic acid, physodic acid, rho- dophyscin, and unknowns. C. Culberson (1966) isolated lividic acid from H. livida but its molecular structure (apparently close to physodic acid) is still unknown. It can be identified with TLC, preferably using the hexane and toluene solvent systems. All of the associated substances fall out in this sequence from highest to lowest RF in hexane: atranorin or lichexanthone, colensoinic acid, norcolensoinic acid, 4-0- demethylphysodic acid, physodic acid, and lividic acid (see C. Culberson, 1966). Various unknowns may appear below and above lividic acid. In toluene the resolution is not as good and only four or five distinct spots may appear in this sequence: atranorin or lichexanthone, colensoinic acid, 4-8-demethylphysodic acid, norcolensoinic acid, and lividic acid. Perfect resolution of spots is only rarely achieved with these solvent systems, and we must await more satisfactory separation that will come from high-performance liquid chromatography. Microphyllinic acid.-H. thysanota. Associated substances: atranorin, echinocarpic acid, and gyrophoric acid. The identity of microphyllinic acid was established by Mr. Jon Dey while comparing the type of Parmelia nakanishii with his P. oostingii. It is the first reported occurrence outside of Cetraria, although C. Culberson postulated its existence in Hypotrachyna (Culberson and Hale, 1973). Norcolensoinic acid.-See discussion under lividic acid. Norobtusatic acid.-All species containing obtusatic acid (see lists under barbatic acid). Associated substances: see discus- sions under barbatic acid and evernic acid. Norstictic acid.-H. andensis, H. brevirhiza, H. caraccensis, H. enderythraea, H. pavovirens, H. lopezii, H. microblasta, H. reducens, H. rhabdiformis, H. sinuosa, and H. sublaevi- gata. Associated substances: atranorin, constictic acid, gal- binic acid, protocetraric acid, salazinic acid, stictic acid, and usnic acid. One of the first positive results of TLC was the resolution of norstictic acid and its closely related associated substances. Norstictic acid occurs in trace amounts with salazinic acid and in substantial amounts with stictic acid. The accessory substance connorstictic acid has been re- ported in Parmotrema (W. Culberson, 1973). Connorstictic acid is very close to salazinic acid and may occur in Hypo- trachyna although I have not been able to identify it posi- tively. Obtusatic acid.-All species containing barbatic acid, H. chicitae. Associated substances: see discussions under bar- batic acid. Olivetoric acid.-H. intercalanda and H. osteoleuca. Asso- ciated substances: atranorin, lichexanthone, rhodophyscin, and unknowns. This CS red acid can be easily identified by means of TLC. Perlatolic acid.-Hi. partita. Associated substances: anziaic acid and atranorin. Perlatolic acid, rare for Hypotrachyna, was identified in only one specimen of H. partita along with closely related anziaic acid. Its co-occurrence with anziaic acid would not be unexpected. Physodalic acid.-H. physodalica and H. subphysodalica. As- sociated substances: protocetraric acid and usnic acid. This acid is biogenetically closely related to protocetraric acid and often occurs with it in Hypogymnia. However, only one of the 14 species of Hypotrachyna having protocetraric acid contains physodalic acid. Physodic acid.-See discussions under lividic acid. Protocetraric acid.-H. bahiana, H. brasiliana, H. consirnilis, H. contradicta, H. croceopustulata, H. flavida, H. Pavovir- ens, H. koyaensis, H. malmei, H. minima, H. obscurella, H. physodalica, H. pseudosinuosa, H. silvatica, and H. vel- loziae. Associated substances: alectoronic acid, atranorin, lichexanthone, norstictic acid, physodalic acid, rhodo- physcin, salazinic acid, and usnic acid. This is the com- monest medullary substance in Hypotrachyna, although none of the species containing it are especially widespread. The best separation (especially from fumarprotocetraric) is achieved in the hexane solvent system using 10-20 ml more ether than recommended by C. Culberson (1972a). NUMBER 25 11 Rhodophyscin.-H. bahiana, H. consimilis, H. croceopustu- lata, H. dactylifera, H. erythrodes, H. gondylophora, H. immaculata, H. rockii, and H. velloziae. Associated sub- stances: atranorin, colensoinic acid, evernic acid, fumar- protocetraric acid, lecanoric acid, lichexanthone, lividic acid complex, protocetraric acid, succinprotocetraric acid, and usnic acid. This red pigment is assumed to be an anthraquinone first isolated from Physcia. It occurs sporad- ically in the species listed, either in the lower medulla or under soralia. Salazinic acid.-H. andensis, H. boquetensis, H. brevirhiza, H. caraccensis, H. enderythraea, H. microblasta, H. re- ducens, H. sinuosa, H. sublaevigata, and H. subsaxatilis. Associated substances: atranorin, constictic acid, galbinic acid, norstictic acid, stictic acid, and usnic acid. I have not conclusively identified consalazinic acid, an unknown sub- stance that may accompany salazinic acid. Stictic acid.-H. andensis, H. lopezii, and H. sinuosa. As- sociated substances: constictic acid, norstictic acid, usnic acid, and two unknowns. This acid is quite rare in Hypo- trachyna and is accompanied by the same substances that are found in other parmelioid genera. Succinprotocetraric acid.-H. gondylophora. Associated sub- stances: atranorin and fumarprotocetraric acid. This re- cently described acid (Baker et al., 1973) often accompanies fumarprotocetraric acid in the genus Xanthoparrnelia. Usnic acid.-H. andensis, H. caraccensis, H. enderythraea, H. flavida, H. flavovirens, H. lopezii, H. microblasta, H. physo- dalica, H. protoboliviana, H. reducens, H. sinuosa, H. sub- physodalica, and H. velloxiae. Associated substances: atra- norin, barbatic acid complex, constictic acid, galbinic acid, norstictic acid, obtusatic acid complex, physodalic acid, protocetraric acid, rhodophyscin, salacinic acid, and stictic acid. Usnic acid, a common cortical pigment, ordinarily replaces atranorin or lichexanthone, although traces of atranorin may be present in some species. It occurs only with P+ 8-orcinol depsidones with the sole exception of H. protoboliviana, which contains the P - barbatic acid complex. Fatty acids.-H. costaricensis, H. koyaensis, and H. singularis. Associated substances: atranorin, protocetraric acid. Fatty acids are represented by caperatic and protolichesterinic acids, perhaps others, all difficult to identify with TLC. ?Gracilescens? unknown.-H. gracilescens. Associated sub- stance: atranorin. ?Oostingii? unknowns.-H. oostingii. Associated substances: atranorin and gyrophoric acid. These unknowns are white fluorescent in longwave ultraviolet but differ from alec- toronic acid. ?Palmarum? unknown.-H. palmarum. Associated substances: atranorin and colensoinic acid. This unknown acid forms a distinct spot below colensoinic acid different from any of the other colensoinic-lividic acid complex substances but obviously related to them. ?Prolongata? unknown.-H. prolongata. Associated substance: atranorin. This C+ red compound could not be identified with any other lichen substances reacting C+. ?Silvatica? anthraquinones-H. silvntica. Associated sub- stances: atranorin, lichexanthone, and protecetraric acid. This rusty red pigment occurs throughout the lower me- dulla. Speciation in Hypotrachyna One of the least discussed topics in lichenology is species evolution. As Thomson (1961) points out, we have little of the kinds of cytological or fossil evidence that is so often available in other plant groups. An important clue in lichenology, however, has proven to be chemistry, and, as stated by Culberson and Culberson (1 970), comparative phytochemistry is the most useful independent check that exists to evaluate the naturalness of systems based on morphology. This is especially true in lichens, where differentiation of chemical races is a far more obvious and frequent means of evolutionary differentiation than morphology (W. Culberson, 1970). I will discuss here primarily the origin and direction of speciation in Hypotrachyna, although many of the observations and conclusions should apply to other foliose genera. Briefly, I will deal with three aspects: possible hybridization of parent species, formation of vegetative morphs, and habitat selection. HYBRIDIZATION Lichenologists probably would agree that crustose lichens reproduce sexually since most species lack any other obvious means of propagation. On the other hand, it is assumed that foliose and other macrolichens generally reproduce vegetatively by means of diaspores (soredia, isidia, etc.). Little has been said about the reproduction of those foliose lichens which lack vegetative diaspores. If two fer- tile lichens should hybridize, we would expect gene exchange and possible mutations, processes that could give rise to new variations and species. Unfortunately, we are not sure how lichens re- produce sexually. There is no way at this time to make experimental crossings and to study cytolog- ical details. It is assumed that microconidia (sper- matia or pycnidiospores) are capable of migrating to the ascogonial coil, forming a dicaryon with the female element, and existing in a brief one-celled sporophytic stage before meiosis and spore formn- tion take place. Each of the eight ascospores can be germinated and will, at least in theory, reconstitute 12 SMITHSONIAN CONTRIBUTIONS TO BOTANY a new thallus when a suitable lichenizable alga is entrapped. Several recent studies have indeed shown that variation in the chemistry of certain populations can best be explained on the assumption of sexual reproduction and hybridization. I can cite my own work in Cetraria ciliaris (Hale, 1963) and the Culber- sons? recent study of the Parmelia perforata group (Culberson and Culberson, 1973). The most con- vincing evidence for Hypotrachyna has been pre- sented by Culberson and Hale (1973) for those species producing evernic acid or barbatic acid. Their data are summarized in Figure 5. Culberson showed that hybridization of plants in the New World with type I Old World chemistry (lM-l = evernic acid and 1-1 = lecanoric acid) and type 11 chemistry (zM-2 = barbatic acid and 2-2 = 4-0-demethylbarbatic acid) theoretically could give rise to a new population containing obtusatic acid (lM-2) and norobtusatic acid (1-2). Old World populations which had no opportunity to cross with progenitors containing evernic acid also now lack any species with obtusatic acid. On the other hand, these populations free of obtusatic acid are absent in the New World except for rare H. sub- plumbeata, leaving us to conclude that hybridi- zation produced populations containing obtusatic acid and that such populations had a very signifi- cant advantage over those lacking this acid and have displaced them. Another example of possible hybridization is illustrated by species which previously would have been considered to contain ?accessory? substances. One could reasonably postulate, for example, that gyrophoric acid occurring with alectoronic acid in some specimens of H. gigas was contributed by a now extinct progenitor which contained gyrophoric acid and which hybridized with the still extant parent containing alectoronic acid. The same rea- soning could be applied to H. physodalica, which produces protocetraric acid with or without physo- dalic acid. A final example concerns the production of cortical substances, principally atranorin, lichexan- thone, and usnic acid. While we are totally ignorant of the biogenesis of these substances, one might assume that atranorin is the normal component and that some mutation could block this substance in favor of lichexanthone (see Hale, 1972b, for H. formosana versus H. pustulifera) or usnic acid. I can cite the following species pairs: H. livida (atranorine) and H. novella (lichexanthone); H. HYPOTHETICAL PROGENITORS / PRIMARY SPECIES P massartit p mplurnbeata I ?\ MU TAT IONS SECONDARY S PEC I ES? P addita p adjuflcta P execta P orientalis P boaotensis tavlorensis 1- P rock?i H Y BRI Di 2 AT IONS P chicitae (2M-2),(2-2), (IM-2), (1-2) P dentelia p diqitata p irnbricatula P laevigata P virqinica FIGURE 5.-Proposed pathway of chemical and morphological evolution leading to species in the barbatic-evernic-acid complex (from Culberson and Hale, 1973). NUMBER 25 13 contradicta (atranorin) and H. brasilianu (lich- exanthone); and H. physcioides (atranorin) and H. protoboliviana (usnic acid). The genus Hypotrachyna in tropical America has 36 species that have apothecia and lack the usual vegetative diaspores. These include 26 species endemic to the New World. They represent a gene pool which has produced, and in many cases is still producing, hybridizations and mutations re- sponsible for the creation of new species. As I will show below, many of the extinct sexual morphs in Hypotrachyna are survived by their vegetative morphs. In summary, the evidence now available supports a working hypothesis that sexual reproduction has occurred in Hypotrachyna and that it has con- tributed significantly to the morphological and, especially, chemical diversity of the genus. FORMATION OF VEGETATIVE MORPHS The primary vegetative diaspores in Hypo- trachyna are soredia and isidia. These never occur together and seem to represent two separate lines of morphological evolution, as I had previously concluded in Parmotrema (Hale, 1965). Most lichenologists of the 19th century treated soredia and isidia as minor characters unworthy of species rank. It was, in fact, not until 1924 that Du Rietz presented a comprehensive discussion of these structures. He not only carefully described their orientation on the thallus but discussed their bio- logical and systematic significance, concluding that sympatric sorediate-nonsorediate, isidiate-nonisidi- ate populations are best treated as varieties. He felt, however, that allopatric populations should be treated as distinct species. In the last decade we have gained a new per- spective on the significance of soredia and isidia, even if we still do not know why they are produced. When studying Parmotrema on a world level, for example, I analyzed a number of ?counterpart? species, that is, pairs of sorediate-nonsorediate and isidiate-nonisidiate species which shared otherwise identical morphology and chemistry (Hale, 1965). Fourteen of the 34 sorediate species had fertile non- sorediate counterparts. The sexual morphs were generally endemic to relatively small continental areas while the sorediate counterparts, as a rule, had much wider distributions. Poelt (1970) expanded these and other examples into a more comprehensive hypothesis of species pairs (?Artenpaare?). Briefly, he postulated that the asexual sorediate and isidiate morphs are de- rived from sexual morphs and that they behave apomictically without any means of further varia- tion save highly unlikely somatic mutations. These secondary species, to quote Poelt, are blind alleys which have conserved the chemistry of the primary sexual morph. Later, Poelt (1972) expanded this hypothesis to include differentiation of a parent species into two or more chemically different, sexually reproducing species, each of which could produce stable secondary apomictic species. In addition, any or all of the primary species theoret- ically comprising species pairs may have become extinct. An upshot of this hypothesis is that morphologi- cally identical but chemically variable sorediate (or isidiate) species, having probably evolved par- allel from different parents, would have to be con sidered polyphyletic. Culberson and Culberson (1973) first suggested this possibility and used it to explain convincingly the origin of speciation in the Parmotrema perforatum group. A further con- sequence is that polyphyletic ?chemical species? almost certainly are distinct species. If the species pair concept is in fact true, it goes far toward explaining the chemical diversity among vegetative morphs. On the other hand, it is sup- ported by no proof other than that offered by comparative biochemistry. As such, it may be an oversimplification that might discourage lichen- ologists from exploring other evolutionary proces- ses. Some sorediate and isidiate species, for example, do produce pycnidia and microconidia which could act as fertilizing elements of fertile populations. These morphs need not remain simply as ?blind alleys.? Alternative hypotheses may come to light as we gain a better understanding of soredial and isidial formation, the role of microconidia, and the mechanics and frequency of gene recombination in lichens. I believe it is profitable, nonetheless, to analyze species pairs in Hypotrachyna and see how much they have contributed to the evolution of the genus. The method is very simple: using a list of diaspore- bearing species, a search is made among nondia- spore-bearing fertile species which might be con- sidered perfect morphological and chemical parents. 14 SMITHSONIAN CONTRIBUTIONS TO BOTANY These parents are defined as strictly as possible with the realization that the sexual morphs may still be evolving and that the pairs may differ be- cause of environmental modifications induced by occupying different habitats. SOREDIATE MoRPHs.-There are eighteen sorediate species in Hypotrachyna and nine of these can be reasonably paired with presumptive parents (given in alphabetic order of the sorediate species): H. brevirhixa - H. sublaevigata H. densirhizinata - H. gigas H. exsplendens - H. degelii H. immaculata - H. livida H. laevigata - H. physcioides H. producta - H. ducalis H. revoluta - H. pluriformis H. rockii - H. pulvinata H. sinuosa - H. reducens The remaining nine species cannot be so closely matched. Their origin can be hypothesized in a few cases. Hypotrachyna flavovirens, for example, nor- mally contains only protocetraric acid but one specimen contains norstictic acid as well. The norstictic acid probably was contributed by a parent similar to H. reducens that crossed with a now extinct progenitor containing protocetraric acid. To cite another example, it is perhaps significant that H. densirhizinata is represented by a single chemical population whereas the presumptive par- ent H. gigas has two populations, one with gyro- phoric acid in addition to alectoronic acid and one with alectoronic acid only. We are left to conclude that the sorediate morph H. densirhizinata evolved before the parent species H. gigas acquired gyro- phoric acid, probably by crossing with a now ex- tinct progenitor related to H. thysanota. Let us next take a group of very similar species which, for all practical purposes, can be distin- guished only by means of chemical tests: H. endochlora (barbatic acid and pigments) H. densirhizinata (alectoronic acid) H. gondylophora (fumarprotocetraric acid) H. laevigata (barbatic acid) H. oostingii (gyrophoric acid) H. producta (anziaic acid) H. rockii (evernic and lecanoric acids) H. thysanota (gyrophoric, microphyllinic, and echinocarpic acids) If we accept the species pairs concept to explain this degree of chemical speciation, then we can appreciate how complicated is the evolutionary pattern involved. Can we assume that these eight species originated from a single sexually reproduc- ing ancestor because of the very similar lobe con- figuration and mode of forming soredia? Would it follow, then, that the present chemical diversity resulted when the ancestor hybridized with other sexual morphs to give eight chemical populations, each of which produced a vegetative morph? While four of the above species seem to have parents, four (H. endochlora, H. gondylophora, H. oostingi and H. thysanota) have no chemically comparable sexual morphs which, we must assume, are either extinct, to be represented now by four chemically stable vegetative morphs, or yet to be discovered in tropical America. If our reasoning is true, then the presumptive parents of the remaining sorediate species in the genus, H. croceopustulata, H. flavovirens, H. pseudosinuosa, H. subafinis, and H. velloziae, are probably extinct. ISIDIATE Mo~p~s.-Al though lichenologists are ignorant of the causes of isidial formation, they rarely ever lump isidiate and nonisidiate species in the same taxon. In Hypotrachyna we find that seven of the sixteen isidiate species have chemically identical nonisidiate presumptive parents: H. andensis - H. reducens H. bogotensis - H. pulvinata H. consimilis - H. bahiana H. ensifolia - H. gigas H. imbricatula - H. physcioides H. microblasta - H. enderythraea H. partita - H. ducalis Two of the remaining nine isidiate species, H. dentella and H. isidiocera, are closely related to the H. physcioides group but differ somewhat chemically. Hypotrachyna neodissecta may have evolved from H. pluriformis but has much smaller spores. All of these vegetative morphs are rather easily distinguished morphologically and no ?chem- ical species? complexes comparable to the H. laevigata group discussed above have evolved. We might assume, then, that the progenitors of isidiate morphs have hybridized or mutated to a lesser degree than the parents of sorediate morphs or that the isidiate morphs, once created, have had a lower survival rate. PUSTULATE MORPHS.-TWO pustulate species have perfect nonpustulate parents: H. dactylifera-H. livida and H. formosana-H. novella. Hypotrachyna malmei has no nonpustulate counterpart. NUMBER 25 15 OTHER MoRPHs.-None of the lobulate species, except perhaps H. rachista-H. ducalis, has a per- fect nonlobulate parent. Hypotrachyna chlorina obviously is related to the H. physcioides complex because it produces barbatic acid; H. prolongata and H. singularis stand isolated. SERIES OF MORPHS.-In the following species a progenitor has apparently given rise to more than one kind of morph: H. ducalis-H. producta (sorediate)-H. partita (isidiate)-H. rachista (lobulate) H. livida-H. immaculata (sorediate)-H. dactylifera (pustulate) H. physcioides-H. laevigata (sorediate)-H. imbricatula (isidi- H. pulvinata-H. rockii (sorediate)-H. bogotensis (isidiate) H. reducens-H. sinuosa (sorediate)-H. andensis (isidiate) This seems to imply that most sexual morphs pro- duce either sorediate, isidiate, or pustulate vegeta- tive morphs and only rarely produce two or more different kinds. In conclusion, we find that Hypotrachyna con- sists of 36 potential sexually reproducing parent or progenitor species. Twelve of these have actually produced 18 morphs, giving a total of 54 species (out of the 77 recognized in this monograph) whose possible evolutionary development can be traced. The remaining 23 asexual species can be assigned to various chemical constellations, but a few simply cannot be explained at this time. It is my contention that the high rate of develop- ment of asexual morphs-involving 66 percent of the sexual morphs-and the coexistence of many of these presumptive progenitors are signs of com- paratively recent origin for the genus. In other words, Hypotrachyna has been speciating rapidly since the continents drifted apart. ate) HABITAT SELECTION The concept of habitat selection as first proposed by Culberson 2nd Culberson (1967) helped to explain the microdistribution of chemical popu- lations of Ramalina siliquosa. Three species, sym- patric over their broad geographic range, segregated at different exposures on rocks into remarkably pure populations, an example of how genetically defined physiological races revealed phenotypically by chemistry (rather than morphology) were associ- ated with distinct habitats. Given sufficient time 2nd resources, I believe similar kinds of habitat selection could be demon- strated among Hypotrachyna species. For example, species containing barbatic acid, especially H. physcioides, seem indiscriminate in selecting hab- itats, rock, trees, or soil. The tree population seems less robust and has more distinct maculae than the saxicolous population. Two distinct species may eventually evolve because of habitat segregation. Other examples could be cited, as the possible dis- tant relationship between H. bahiana (corticolous) and H. contradicta (saxicolous) or between H. degelii (corticolous) and H. protenta (saxicolous). Classification of Hypotrachyna Vainio proposed three sections in his Brazilian study: Amphigymnia, Hypotrachyna, and Xantho- parmelia. Hypotrachyna was subdivided into three groups without formal rank: Irregularis, Cyclo- cheila, and Linearis. It is this last group by which I typified subgenus Hypotrachyna (Vainio) Hale and Kurokawa with Parmelia brasiliana Nylander as the lectotype species (Hale and Kurokawa, 1964). I recently elevated this taxon to generic level (Hale, 1974a). Vainio assigned eleven species to Hypotrachyna, and, in fact, only one that he listed, Parmelia coronata Fee, does not belong here. Two yellow species, P. /lava and P. velloziae, were placed in section Xanthoparmelia but they have dichoto- mously branched rhizines and belong in Hypotrachy nu. While Hypotrachyna contains about 110 species, there is no practical internal subdivision into sub- genera or sections. The most useful approach is to combine the species into chemical constellations proposed by Culberson and Hale (1973), which are based on the probably biogenetic relationship of the chemicals in Hypotrachyna (Figure 6). It is assumed that simple depsides, for example, represent the lowest evolutionary level and mixed depsidones the highest. Accordingly, in my opinion, species in the lividic acid complex (including the boxes for colensoinic acid and 4-O-methylphysodic acid) should and do represent the highest level of evolution in the genus. Without question, the barbatic-evernic acid complex has the highest level of speciation. Another advanced group, with alectoronic acid (H. gigas, H. degelii, etc.), seems to have evolved in parallel along different lines 16 SMITHSONIAN CONTRIBUTIONS TO BOTANY SIMPLE I) DEPSIDONES alec toroni c norcolenso'nic FIGURE 6.-Biogenetic relationships among secondary products in Hypotrachyna. Compounds connected by lines have similar chemical structures but are not necessarily derivable from the other. and has a lower level of speciation. As one might expect, the P+ acid series represented by H. brasilianu, H. enderythraea, H. reducens, H. sub- Zaevigata, etc., seems to include more conservative species, some pantropical. The various species of Hypotrachyna are grouped provisionally (below) according to the biogenetic chart (Figure 6) and a consideration of morpho- logical traits. The groups joined by dashes have a generally higher level of internal homogeneity than those separated by commas. Fatty acids.-H. costaricensis and H. singularis. PD+ compounds.-Norstictic acid derivatives: H. andensis- H. reducens-H. rhabdiformis-H. sinuosa, H. boquetensis- H. brevirhiza-Hi. sublaevigata, H. caraccensis-H. endery- thraea-H. microblasta, H. lopezii, H. subsaxatilis. Proto- cetraric acid derivatives: H. brasiliana-H. bahiana, H. con- sirnilis-H. contradicta-H. croceopustulata-H. pseudosinuosa- H. silvatica, H. flavida, H. flavovirens, H. gondylophora, H. koyaensis, H. malmei-H. minima, H. obscurella, H. physo- dalica, H. subafinis-H. velloziae, H. subphysodalica. Evernic-1ecanoric.-H. bogotensis-H. chicitae-H. pulvinata- H. rockii. Gyrophoric: H. neodissecta-H. osorioi-H. pluriformis-H. re- voluta. Barbatic: H. subplumbeata. Obtusatic: H. chlorina-H. dentella-H. endochlora-H. imbri- catula-H. isidiocera-H. laevigata-H. monolifera-Hi. peru- viana-H. physcioides-H. steyermarkii, H. protoboliviana. Microphyllinic: H. thysanota. Alectoronic: H. degelii-H. exsplendens-H. protenta, H. den- sirhizinata-H. ensifolia-H. gigas-H. lineariloba, (?)H. oostingii. Perlatolic-anziaic: H. ducalis-H. partita-H. producta-H. pro- longata-H. rachista, H. eitenii. Olivetoric: H. intercalanda-H. osteoleuca. Colensoinic: H. dactylifera-H. erythrodes-H. formosana-H. gracilescens-H. immaculata-H. livida-H. novella-H. pal- marum. More detailed discussions of some of these groups will be found in the section on species treatment. Habitats and Ecology Hypotrachyna is primarily a genus of higher elevations throughout its range in the tropics. It grows near sea level only at its northern limits in southern United States and southern limits in Chile and Argentina. It is absent or very rare in xero- phytic areas and in lowland (0-500 m elevation) rain forest. The optimal area for Hypotrachyna is the zone between 1300 and 2400 m elevation in the major mountain chains of tropical America. The climate here is quite uniform throughout the year and pre- cipitation comes in the form of rain and frequent clouds or mists. There are about 55 species here. Most are corticolous. Where rock outcrops are NUMBER 25 Phytogeographical region 17 Endemic species Extraneotropical species Total flora No. Percent No. Percent abundant, however, a number of obligately saxi- colous species may occur: H. brasiliana, H. contra- dicta, H. erythrodes, H. flavida, H. obscurella, H. osteoleuca, H. protenta, H. subsaxatilis, etc. Even some typically corticolous species may grow on rocks: H. formosana, H. imbricatula, H. micro- blasta, H. physcioides, H. revoluta, etc. The area above 2400 m in the American tropics becomes more sparsely forested and intergrades with the true treeless paramo. The Hypotrachyna flora here consists of 36 species, including many from the lower zone. The high paramo has much greater climatic extremes than other zones. The 20 species occur- ring here (3600-4500 m elevation) include some of the most showy and spectacular ones: H. carac- censis and H. gigas. Hypotrachyna chicitae, H. physodalica, and H. subplumbeata are the only species apparently restricted to high paramo. These paramo lichens grow on rocks, over grass and moss tussocks, and at the base of small shrubs, forming a conspicuous part of the vegetation. The zone below 1000-1200 m in the tropics is largely true rain forest and, relatively speaking, is poor in terms of number and diversity of foliose species. Only 14 species of Hypotrachyna have been recorded below 1000 m. These include those from temperate and subtropical latitudes in the United States and southern South America as well as ?weedy? species with broad elevational amplitude such as H. costaricensis, H. formosana, H. laevigata, H. microblasta, H. revoluta, and H. rockii. North America .............................. West Indies ...................................... Andes ................................................ Mexico/Central America .............. Brazil/Uruguay .............................. Chile/Argentina ........................... Phytogeographical Relationships Hypotrachyna, primarily a tropical genus, is concentrated in the New World, There are 58 19 1 5 6 32 29 2 I 12 41 40 4 10 14 35 46 9 20 11 24 38 11 29 8 21 15 2 13 7 47 species that occur only in the Americas, while Asia and Africa together have only 32 endemics. The American tropics, however, differ consider- ably in floristic composition and degree of endem- ism. Six major regions can be recognized, as shown in Table 1, where comparisons are made with pantropical elements and Europe and Africa. Eu- rope has only one endemic, H. taylorensis, and four other species, H. endochlora, H. laevigata, H. revoluta, and H. sinuosa, all pantropical. Two regions of high endemism stand out: the Andean region (Colombia, Venezuela, Peru, Ecua- dor, and Bolivia) and the area from Minas Gerais and Mato Grosso in Brazil to adjacent Uruguay. The farther one goes from these centers the lower the degree of endemism and speciation. Actually, the Andes and southeastern Brazil have many species in common and this number surely will be increased as more collecting is done. Nineteen of the 77 species of Hypotrachyna occur outside the New World: H. bogotensis and H. laevigata are known from Europe: H. brevir- hiza, H. costaricensis, H. endochlora, H. formosana, H. koyaensis, H. microblasta, H. pseudosinuosa, H. revoluta, H. rockii, and H. sinuosa are pan- tropical or occur on at least one Old World land mass; and the most interesting group, comprising H. dactylifera, H. degelii, H. densirhizinata, H. exsplendens, H. immaculata, H. livida, and H. subplumbeata, occurs outside of the New World only in southern Africa, a classic pattern for plants which evolved before the continents drifted apart. In the latter group there is a preponderance of species with the lividic acid complex. TABLE 1 .-Total Hypotrachyna flora, number and percentage of endemic species, and number and percentage of species also occurring outside the neotropics, according to the major phytogeographical regions. 18 SMITHSONIAN CONTRIBUTIONS TO BOTANY The Hypotrachyna Flora The flora as presented at this time comprises 77 species. The most commonly collected ones (with number of collections examined) are H. imbricatula (81 +), H. costaricensis (65+), H. bogotensis (60+), H. microblasta (52+), H. rockii (50+), H. physcioides (35+), H. laevigata (35), H. sinuosa (33), H. caraccensis (32), H. gigas (31), H. pulvinata (31), H. densirhizinata (25), and H. livida (25). It is noteworthy that the four com- monest species are isidiate. The rarities in the Hypotrachyna flora, found only once or twice, include H. contradicta, H. ducalis, H. eitenii, H. malmei, H. minima, H. monilifera, H. neodissecta, H. prolongata, H. protoboliviana, H. singularis, H. subsaxatilis, and H. subplumbeata. The number of species in each country or political entity in tropical America varies from highs of 42 species in Venezuela and 37 in Brazil to only one species in Honduras, Puerto Rico, and Trinidad. These differences obviously reflect the intensity of collecting and not necessarily a poor flora. The following listing of the Hypotrachyna flora of each country shows how much work remains to be done, and hopefully it will stimulate further collecting efforts: ARGENTINA: H. brevirhiza, H. densirhizinata, H. gigas, H. intercalanda, H. livida, and H. pluriformis. BOLIVIA: H. caraccensis, H. costaricensis, H. densirhizinata, H. enderythraea, H. gigas, H. gondylophora, H. physcioides, H. pulvinata, H. revoluta, and H. sinuosa. BRAZIL: H. bahiana, H. brasiliana, H. caraccensis, H. chlorina, H. consimilis, H. contradicta, H. costaricensis, H. dactyli- fera, H. degelii, H. dentella, H. eitenii, H. enderythraea, H. endochlora, H. erythrodes, H. pavida, H. formosana, H. gracilescens, H. imbricatula, H. immaculata, H. inter- calanda, H. isidiocera, H. laevigata, H. livida, H. malmei, H. microblasta, H. minima, H. novella, H. obscurella, H. palmarum, H. peruviana, H. pluriformis, H. protenta, H. pulvinata, H. rockii, H. silvatica, H. subafinis, and H. velloziae. CHILE: H. bogotensis, H. brevirhiza, H. dactylifera, H. densir- hizinata, H. pavovirens, H. formosana, H. imbricatula, H. laevigata, H. oostingii, H. pseudosinuosa, H. revoluta, H. rockii, H. sinuosa, H. sublaevigata, and H. subphysodalica. COLOMBIA: H. andensis, H. bogotensis, H. caraccensis, H. costaricensis, H. degelii, H. densirhizinata, H. endochlora, H. ensifolia, H. Pavida, H. gigas, H. imbricatula, H. isidio- cera, H. laevigata, H. microblasta, H. osteoleuca, H. phys- cioides, H. physodalira, H. producta, H. protenta, H. pulvinata, H. rachista, H. reducens, H. rockii, and H. sinuosa. COSTA RICA: H. andensis, H. bogotensis, H. caraccensis, H. chicitae, H. costaricensis, H. dactylifera, H. densirhizinata, H. endochlora, H. ensifolia, H. exsplendens, H. gondylo- phora, H. imbricatula, H. laevigata, H. microblasta, H. partita, H. physcioides, H. protenta, H. protoboliviana, H. pulvinata, H. rockii, and H. sinuosa. CUBA: H. costaricensis, H. formosana, H. imbricatula, H. microblasta, and H. rockii. DOMINICAN REPUBLIC: H. bogotensis, H. consimilis, H. cos- taricensis, H. densirhizinata, H. endochlora, H. ensifolia, H. gondylophora, H. imbricatula, H. isidiocera, H. laevi- gata, H. lineariloba, H. microblasta, H. physcioides, H. pseudosinuosa, H. rachista, H. revoluta, H. rockii, H. sinu- osa, H. subafinis, and H. velloziae. ECUADOR: H. bogotensis, H. brevirhiza, H. caraccensis, H. densirhizinata, H. enderythraea, H. gigas, H. microblasta, H. pulvinata, and H. sinuosa. GUATEMALA: H. caraccensis, H. costaricensis, H. densirhizinata, H. exsplendens, H. formosana, H. imbricatula, H. isidiocera, H. microblasta, H. physcioides, H. pulvinata, H. rockii, and H. subafinis. GUYANA: H. imbricatula and H. laevigata. HAITI: H. bahiana, H. bogotensis, H. chlorina, H. costari- censis, H. croceopustulata, H. densirhizinata, H. ensifolia, H. formosana, H. imbricatula, H. microblasta, H. physci- oides, H. prolongata, H. pseudosinuosa, H. rockii, H. sinuosa, H. subafinis, and H. velloziae. HONDURAS: H. imbricatula. JAMAICA: H. bogotensis, H. costaricensis, H. enderythaea, H. ensifolia, H. exsplendens, H. imbricatula, H. isidiocera, H. laevigata, H. microblasta, H. pseudosinuosa, H. pulvinata, and H. rockii. LESSER ANTILLES (Dominica, Grenada, Guadeloupe, Marti- nique, and St. Vincent): H. costaricensis, H. imbricatula, H. laevigata, H. microblasta, H. pseudosinuosa, and H. rockii. MEXICO: H. bogotensis, H. consimilis, H. costaricensis, H. croceopustulata, H. dactylifera, H. densirhizinata, H. den- tella, H. endochlora, H. ensifolia, H. exsplendens, H. for- mosana, H. gigas, H. gondylophora, H. imbricatula, H. isidiocera, H. koyaensis, H. laevigata, H. microblasta, H. physcioides, H, pluriformis, H. pseudosinuosa, H. pulvinata, H. rachista, H. rockii, H. sinuosa, H. subafinis, H. sub- laevigata, H. subplumbeata, H. subaxatilis, and H. thy- sanota. PANAMA: H. boquetensis, H. caraccensis, H. chlorina, H. COS- taricensis, H. dactylifera, H. densirhizinata, H. endochlora, H. ensifolia, H. formosana, H. gigas, H. imbricatula, H. isidiocera, H. microblasta, H. partita, H. physcioides, H. protenta, H. pulvinata, H. rachista, H. reducens, H. rhab- diformis, H. rockii, and H. sublaevigata. PERU: H, andensis, H. bogotensis, H. caraccensis, H. chicitae, H. chlorina, H. costaricensis, H. degelii, H. densirhizinata, H. ducalis, H. enderythraea, H. erythrodes, H. pavida, H. formosana, H. gigas, H. imbricatula, H. microblasta, H. peruviana, H. physodalica, H. protenta, H. pulvinata, H. rachista, H. reducens, H. rhabdiformis, H. rockii, and H. singularis. PUERTO RICO: H. consirnilis. NUMBER 25 19 TRINIDAD: H. imbricatula. URUGUAY: H. dactylifera, H. endochlora, H. livida, H. osorioi, and H. phiformis. VENEZUELA: H. andensis, H. bogotensis, H. brevirhiza, H. caraccensis, H. chicitae, H. chlorina, H. consimilis, H. cos- taricensis, H. dactylifera, H. densirhizinata, H. dentella, H. ducalis, H. enderythraea, H. endochlora, H. ensifolia, H. erythrodes, H. pavida, H. formosana, H. gigas, H. gondylo- phora, H. imbricatula, H. immaculata, H. isidiocera, H. laevigata, H. livida, H. loperii, H. microblasta, H. moni- lifera, H. neodissecta, H. novella, H. obscurella, H. osteo- leuca, H. partita, H. physcioides, H. physodalica, H. pro- tenta, H. pulvinata, H. rachista, H. revoluta, H. rockii, H. sinuosa, and H. steyermarkii. Nomenclature One of the most time-consuming tasks in any monographic effort is unraveling problems in nomenclature and typification. The literature must be searched as completely as possible, the identity and location of type specimens must be determined, and, where needed, lectotypes must be selected. I have decided to use a narrow interpretation of the Code of Botanical Nomenclature in defining types. A holotype, as I employ the term here, is the specimen actually designated as such by an author. This includes all species described since 1958 when the designation ?holotype? became mandatory for a single element on which a new taxon is based. Before 1958 Gyelnik seems to be the only lichen- ologist who consistently used this term in a correct, modern sense. All other authors either failed to cite a holotype collection or left to later workers the job of deciding which of two or more specimens cited in a species description is the lectotype. A lectotype, then, in my definition, includes two cases, one in which one of the published syntypes is selected as the lectotype and another where the author listed only one collection in his description but failed to call it a holotype and often did not state clearly where the specimen was housed. In other words, I am often selecting one specimen out of several duplicate collections as the lectotype. The point of departure here is that I had formerly applied the term ?holotype? to a collection which represented the only material cited by an author. Because of the element of selection involved, even as to location of the specimen, I feel the use of the term ?lectotype? is justified. It should be emphasized that selection of lecto- types is not binding on later workers. The Code allows us to lectotypify a species again if better evidence can be produced to support the change. This may happen, for example, when a second worker discovers a specimen in another collection that is in better condition or is more appropriate than the previously designated lectotype. A pos- sibly undesirable amount of nomenclatorial in- stability may be introduced in these circumstances, but this seems a better course than rigidly follow- ing a lectotypification which the earlier worker himself admits may be faulty. Keys to the Species The following keys are divided into three major groups: sorediate, isidiate, and nonsorediate- nonisidiate species. The main division under each group is presence or absence of usnic acid (cortex greenish yellow or whitish to ashy gray). Following this, a combination of pigment and morphological characters is used, but ultimately chemical tests are called for. The standard color tests with KOH, calcium hypochlorite (or other bleach), and p- phenylenediamine suffice to identify a number of species and species groups, but below this level it is highly desirable, if not sometimes mandatory, to identify the substances with microchemical tests, preferably thin-layer chromatography. Further clues as to the species identity can be gained by compar- ing the specimens with the photographs and by consulting the descriptions and geographical distri- bution. The geographic ranges of many species will undoubtedly be extended as more collections are made, but geography is still a valuable aid. Key to Species with Thallus Sorediate or Sorediate-Pustulate 1. Thallus greenish yellow (usnic acid present). 2. Lobes very narrow, less than 1 mm wide; medulla K- ........................... 77. H. velloxiae 2. Lobes 1-3 mm wide; medulla Kf red or K-. 3. Medulla KS yellow turning red ..................................................................... 69. H. sinuosa 3. Medulla K- (protocetraric acid) ................................................................... 26. H. fluvovirens 20 SMITHSONIAN CONTRIBUTIONS TO BOTANY 1. Thallus whitish to ashy gray (usnic acid lacking). 4. Medulla uniformly yellow .................................................................................... 21. H. endochlora 4. Medulla white. 5. Soralia distinct, generally terminal or subterminal, soredia powdery. 6. Medulla K+ red (salazinic acid) ................................................................. 6. H. brmirhixa 6. Medulla K- or slowly dull yellowish. 7. Medulla Cf rose, red, or orange-red. 8. Soralia orbicular, distinct ............................................................ H. lmigata group (Barbatic acid: 36. H. Zaevigata; evernic-lecanoric acids: 66. H. rockii; gyro- phoric acid: 47. H. oostingii; anziaic acid: 56. H. producta; gyrophoric- echinocarpic acids: 76. H. thysanota.) 8. Soralia more irregular and diffuse (Figure 2a); gyrophoric acid present .......... .................................................................................................. 9. Medulla P+ red. 10. Lobes narrow, 1-2 mm wide; thallus closely adnate ...................................... 10. 11. Thallus adnate; lobes short ...................................... 11. Thallus loosely attached; lobes longdivaricate .. ...................... 60. H. pseudosinuosa Lobes 2-5 mm wide; thallus loosely attached ............. 29. H. gondylophora 9. Medulla P-. 5. Soralia laminal toward the center of the thallus. 12. Soralia with powdery soredia. 13. 13. Lobes 1-5 mm wide; thallus loosely adnate. 14. 14. Medulla C-. Lobes narrow, about 1 mm wide; thallus closely adnate .......... 71. H. subaffinis Medulla C+ red (gyrophoric acid) ....................................... 64. H. revolwta 15. Medulla P 4- red ........ 13. H. croceopustulata .............. 32. H. immaculata ................................ 12. Soralia pustulate with sparse coarse soredia or pustules present. 16. Lichexanthone present (thallus yellow in UV light). 17. Lobes narrow, 1-2 mm; confined to Brazil ............................ 40. H. malmei 17. Lobes 1-5 mm wide; pantropical ........................................ 27. H. formosana 16. Lichexanthone absent (no fluorescense in UV). 18. Medulla Cf rose (gyrophoric acid); isidiate pustules solid ... 48. H. osorioi 18. Medulla C-. Pustules becoming sorediate (Figure 2c) ............ 13. H. croceopustulata Pustules not sorediate (Figure 2d) .............................. 14. H. dactylifera 19. 19. Key to Species with Thallus Isidiate, Isidiate-Lobulate, or Pustulate 1. Thallus greenish esent). 2. Medulla K- ............................................................ 73. H. subphysodalica 3. Isidia remaining cylindrical .......................................................................... 41, H. microblasta 3. Isidia distinctly lobulate .................................................................................... 1. Thallus whitish to ashy gray. 4. Medulla uniformly yellow ........................................................................................ 34. K. isidiocera 4. Medulla white. 5. Isidia mostly laminal on the lobe surface. 6. Isidia inflated and hollow (pustules). See dichotomy 17 in the sorediate key above. 6. Isidia cylindrical, solid, not strongly inflated. 7. Medulla KS yellow turning red ..................... ...................... 65. H. rhabdiformis 7. Medulla K- or slowly pale yellowish. 8. Medulla C+ (and KC+) orange (barbatic acid group). 9. Lobes narrow and appressed, 1-2 mm wide; isidia ciliate. ........................ ............................................ ..................................................... 70. H. steyermarkii 9. Lobes 2-5 mm wide, isidia eciliate. 10. Medulla P- ........................................................................ 31. H. imbricatula NUMBER 25 21 10. Medulla P f red ..................................................................... 17. H. dentella 8. Medulla CS rose or red or C-. 11. Medulla C+ rose or red (anziaic or gyrophoric acids). 12. Isidia not cylindrical. 12. Isidia cylindrical, erect .................................................. .44. H. neodissecta 13. Isidia becoming lobulate, procumbent ........................ 51, H. partita 13. Isidia club-shaped, crumbling apically ....................... 48. H. osorioi 14. Isidia hollow (pustules) .................................................. 14. H. dactylifera Lobes sublinear, 1-3 mm wide ........................ 10. H. consirnilis 16. Lobes subirregular, 3-7 mm wide .................... 35. H. koyaensis 17. Isidia cylindrical, erect; fatty acids present ............................ 11. Medulla C- or faint C+ rose (lecanoric acid). 14. Isidia solid, not pustulate. 15. Medulla Pf red (protocetraric acid). 16. 15. Medulla P-. ............................................. .., , , , . . , , .. , .. , ..12. H. costdcen& 18. Alectoronic acid pres (strongly white fluorescent ......................... 22. H. ensifolia 18. Evernic and lecanoric present (weak to negative 17. Isidia becoming lobulate, procumbent. under UV) ..................... fluorescence) .................................................... 3. H. bogotensis 19. Anziaic acid present ........................................................................................ 6 2. H. rachista 19. Unknown C+ substance present: known only from Haiti ........... ..57. H. POlMgata Key to Species with Thallus Lacking Soredia, Isidia, and Pustules 5. Isidia marginal toward lobe tips (Figure 44. 1. Thallus greenish yellow (usnic acid present). 2. Lobes long and little branched, divaricate; thallus loosely attached. 3. Medulla K- .......................................................................................... ..54. H. physodalica 3. Medulla K+ yellow turning red. 4. Stictic acid present .............................................................................................. 39. H. lopezii 4. Salazinic acid present ...................................................................................... 7. H. cmaccensis 2. Lobes shorter, moderately branched; thallus generally adnate. 5. Medulla K+ yellow turning red. 6. Thallus corticolous ..................... ................................................. 63. H. reducens 6. Thallus saxicolous ...................... .......................................... 20. H. enderythraea 5. Medulla K- or KS very pale yellow. 7. Thallus corticolous; medulla C + orange ............................... 59. H. protobolivima 7. Thallus saxicolous; medulla C- ............................................................. 25. H. flavida 1. Thallus whitish to ashy gray. 8. All or most of the medulla pigmented yellow or red. 9. Thallus saxicolous. 10. Medulla C + deep red (olivetoric acid) ................................................ 49. H. osteoleuca 11. Margins of lobes densely lobulate (Figure 4e) .......................................... 9. H. chlorina 12. Pigment pale yellow, throughout the medulla ............................ 52. H. peruviana 12. Pigment red, occurring in the lower half of the medulla .......... 67. H. silvatica 14. Medulla C+ orange (barbatic acid group) ................................... 43. H. monilifera 15. Medulla C+ deep red (anziaic acid) ............................................. 18. H. ducalis Thallus small, 3-6 cm broad ................................................ 37. H. lineariloba 10. Medulla C- .......... ............................................................. ......... 23. H. erythrodes 9. Thallus corticolous. 11. Margins of lobes sparcely lobulate or lacking lobules. 8. Medulla white throughout. 13. Lobes long and little branched; thallus loosely attached. 14. Medulla Cf rose or red or C-. 15. Medulla C+ rose or C- (alectoronic acid). 16. 16. Thallus larger, more than 8 cm broad ......................................... 28. H. gigas 13. Lobes shorter and moderately branched; thallus adnate. 22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 17. Thallus collected on rocks. 18. Medulla K+ yellow turning red (salazinic acid) .................... 75. H. subsaxatilis 18. Medulla K- or K+ pale yellow. 19. Medulla C+ orange or red. 20. 20. Medulla C+ red or rose. 21. Medulla all white. Medulla C+ orange (barbatic acid) .......................... 53. H. physcioides 21. Lower part of medulla pigmented orange .......... 49. H. osteoleuca 22. Medulla C + deep red (anziaic acid) .................. 19. H. eitenii 22. Medulla C+ rose or C- ........................................ 8. H. chicitae 19. Medulla C- . 23. Medulla P+ red (protocetraric acid). 24. Lichexanthone present in the cortex (UV+ orange). 25. Lobes short, brittle and fragile .......................... 42. H. minima 26. Thallus dark gray; lobes about 1 mm wide, short .................. .......................................................................... 46. H. obscurella llus whitish gray; lobes 2-4 mm wide ..11. H. contradicta 25. Lobes sublinear-elongate, coriaceous ............... .5. H. brasiliana 24. Lichexanthone absent (UV-). 23. Medulla P-. 27. Lichexanthone present in cortex (UV+ orange) ................... 27. Lichexanthone absent. .................... ............................................................ 23. H. eryth 28. Lividic acid group present 28. Lividic acid goup absent. ............................ 38. H. livida 29. Alectoronic acid present .............................. 58. H. potenta 29. ?Gracilescens? unknown present ......... 30. H. gracilescens 31. Norstictic acid present ........................................................ 72. H. sublmigata 31. Norstictic acid absent ............................................................. 4. H. boquetensis 32. Medulla PS red (protocetraric acid) ......................................... 2. H. bahiuna 17. Thallus collected on trees. 30. Medulla K+ yellow turning red (salazinic acid). 30. Medulla K- or K+ pale yellow. 32. Medulla P - . 33. Medulla C+ orange, red, or rose. 34. Medulla C+ orange (barbatic acid group). 35. Apothecia strongly stalked .............................. 52. H. peruuimta 35. Apothecia adnate. 36. Maculae absent to weakly developed 53. H. physcioides 36. Maculae strongly developed (Figure 40 ............. ................................................... 74. H. subp 34. Medulla C+ red or rose. 37. Lobes subirregular, broadly canaliculate: C 4- rose (gyroph- 37. Lobes sublinear, not canaliculate. oric acid) .......................................................... 55. H. pluriformis present .......................................................... 61. H. pulvinatn 38. Maculae usually conspicuous; evernic and lecanoric acids 38. Maculae absent: olivetoric acid present ............................. ................................................................... 33. H. intercalanda 39. Lobe margins distinctly lobulate ......................... 68. H. singularis 40. Lichexanthone present in cortex (UV+ orange) .................... ............................................................................. 45. H. novella 40. Lichexanthone absent. 33. Medulla C--. 39. Lobe margins without lobulae. 41. Alectoronic acid present ................................ 15. H. degelii 42. Lividic acid present .................................. 38. H. livida 42. ?Palmarum? unknown present ... 50. H. palmmum 41. Alectoronic acid absent. NUMBER 25 28 Species Treatment The species are arranged in alphabetic order. Standard herbarium acronyms are given to show the location of specimens cited except for collec- tions by Hale, all of which are in US. 1. Hypotrachyna andensis, new species FIGURE la Thallus arcte adnatus vel adnatus, fragilis, 4-7 cm latus, viridi-flavicans, lobis sublinearibus, sep- aratis, 1-3 mm latis; superne isidiatus, isidiis lobu- latis; cortex superior 12p-14p crassus, stratum gon- idiale 24p-28p crassum, medulla alba, 120p-130y crassa, cortex inferior 12~-14~ crassus; subtus niger, dense rhizinosus. Apothecia substipitata, sporis simplicibus, lop X 14p. Thallus closely adnate on twigs or on soil, fragile, 4-7 cm broad, yellowish green; lobes sublinear, rather elongate, mostly separate, 1-3 mm wide; upper surface plane, continuous, becoming isidiate, the isidia at length lobulate, the lobules dorsiven- tral, about 0.5 mm long, often with short cilia; lower surface densely rhizinate, the rhizines moderately dichotomously branched. Apothecia substipitate, 1-3 mm in diameter; spores lop X 14~. CHEmsmY.-Cortex K-, medulla K+ yellow turning red, C-, P+ orange (usnic acid, norstictic acid, and salazinic acid (holotype); or more rarely norstictic acid, stictic acid, and constictic acid). TYPE CoLLEcTIoN.-San Martin, Peru, Holligan L 128 (BM, holotype). DIsTRIsuTIoN.-Central America, Colombia, Ven- ezuela, and Peru. HABITAT.-On branches of small shrubs in the paramo or on road banks at 2800-3700 m elevation. REMARKS.-This paramo species usually is densely isidiate-lobulate. The presumptive pro- genitor is H. reducens. The chemical variation parallels that in H. sinuosa, the sorediate morph. Almeda 2457 (DUKE). Colombia: Cundinamarca, Cuatrecasas 2706813 (US). Venezuela: Merida, Hale 42658a, 43072, 43146. Peru: Bryan 596 (WIS). SPECIMENS EXAMINED.-costa Rica: Cartago, 2. Hypotrachyna bahiana, new combination FIGURE 7b Parmelia bahiana Sylander, 1885: 612. [Type collection: Bahia, Brazil, Blanchet (H, lectotype; M, P, isolectotopes).] Parrnelia insinuans Nylander, 1885:612. [Type collection: Tropical America, Bonpland (H, lectotype; P, isolectotype).] Parmelia pava f. albescens Krempelhuber, 1873:lO. [Type col- lection: Serra da Mantiqueira, Brazil, Warming (M, lecto- type).] Thallus adnate, mineral gray, 3-6 cm broad; lobes sublinear, short, 1-2 mm wide; upper surface plane, continuous, soredia and isidia lacking; lower surface moderately to densely rhizinate, the rhiz- ines densely dichotomously branched. Apothecia common, adnate, 2-4 mm in diameter; spores 6p-8p X lop-13p. CHEMISTRY.-cOrteX Kf yellow, medulla K--, C -, P + orange-red (atranorin, protocetraric acid, and, if pigmented in the lower medulla, rhodophyscin). DISTRIBUTION.-weSt Indies, Brazil. HABITAT.-On trees in open forest at 1000-2000 m elevation. REMARKs.-Hypotrachyna bahiana is externally similar to H. physcioides but differs in chemistry. It is common only in Brazil. The isidiate morph is H. consirnilis. SPECIMENS EXAMINED.-Haiti: Sud, Zmshaug 23229, Wetmore 3364 (MSC). Brazil: Minas Ger- ais, Warming 280 (M); Rio de Janeiro, Eiten 6433, 7502 (US), Glaziou 1119 (G,H,P). 3. Hypotrachyna bogotensis, new combination FIGURE 7c Parmelia bogotensis Vainio, 1899: (122). [Type collection: Bogota, Colombia, Weir 53 (TUR, lectotype; BM, isolecto- Parrnelia culmigena Zahlbruckner, 1905:81. [Type collection: type).] El Altar, Ecuador, Meyer 381 (\$?, lectotype).] Thallus adnate to loosely attached, 6-12 cm broad; lobes sublinear, dichotomously branched, 1-6 mm wide; upper surface plane, shiny, white- maculate, densely isidiate, the isidia cylindrical or at length procumbent and dorsiventral; lower surface densely rhizinate, the rhizines finely branched and forming a woolly mat. Apothecia very rare, adnate, to 5 mm in diameter; spores CHEiwsTRY.-Cortex K+ yellow, medulla K-, C+, KC+ rose to red, P- (atranorin, evernic acid, and lecanoric acid). America, West Indies, South America south to 6y x 121.1. DISTRIBUTION AND HABITAT.-Mexico, Central 24 SMITHSONIAN CONTRIBUTIONS TO BOTANY FIGURE 7.-Species of Hypotmchyna: a, H. andensis (Holligan L128); b, H. bahiana (Eiten 7502); c, H. bogotensis (Oberwinkler and Poelt 7544); d, H. boquetensis (Hale 38878) e, H. brasilianu (Reitz and Klein 10187); f, H. brevirhiia (Hale 42532). (Scale in mm.) NUMBER 25 25 Chile on trees in secondary and primary forests (oak, pine, Drimys), shrubs, fence posts, rocks, and soil on road banks at 1800-4100 m elevation. REMARKS.-This is an extremely variable and widespread species. The isidia may be quite sparse and merely papillate or even subsorediate but can eventually enlarge and grow procumbent (Figure 4c), almost as narrow lobules, as Vainio (1899) has noted. Parmelia culmigena represents this extreme form, It is closely related to both H. pulvinata, the presumptive progenitor, and H. rockii, a sorediate morph. Care must be taken to distinguish it from H. ensifolia (alectoronic acid present) when, as is often the case, both species occur in the same habitat, especially on roadsides. ico: Chiapas, Hale 20266, 20541, 20542; Oaxaca, Hale 20684, 20788; Sinaloa, Culberson 13384, 13421 (DUKE). Costa Rica: Alajuela, Culberson 12369 (DUKE). Jamaica: Zmshaug 13044, 15442, 15465 (MSC, US), Plitt 129 (US). Haiti: Ouest, Zmshazig 22959 (MSC, US), 22958 (MSC); Sud, Zmshaug 23232 (MSC), Wetmore 3294, 3369 (MSC). Dominican Republic: La Vega, Zmshaug 23371, 23402, 23406 (MSC, US), 23537, 23548 MSC), Wetmore 3481, 3613, 3620 (MSC, US), 3522, 3627, 3662 (MSC). Colombia: Boyaca, Cuatrecasas 1364a (US); Santander, Killip and Smith 18942 (NY), 18594 (US). Venezuela: MCr- ida, Hale 42387, 42607, 43120, 43200, 43277, 43350, Poelt and Oberwinkler 13172b, 14427~ (M). Peru: Bryan 399 (F). Ecuador: Jameson 144 (BM). Chile: Aysen, Santesson 4309, 4345, 4488, 6943, 6945 (S); Chiloe, Santesson 4192 (S); fuan Fernandez, Skottsbeyg 87 (NY, S, US), 198 (BM), Zmshaug 37384 (REX); Valdivia, Santes- son 3274, 3521 (S, US), 3272, 3387, 3404, 3475, 3556, 3710, 7149 (S). REPRESENTATIVE s PECIM EN S EXAM INED.-MeX- 4. Hypotrachyna boquetensis, new combination FIGURE 7d Parmelia boquetensis Hale, 1974b:265. [Type collection: Bo- quete, Chiriqui, Panama, Hale 38878 (US).] Thallus closely adnate on branches, 8-12 cm broad, greenish ashy gray; lobes subirregular, 3-4 mm wide, the margins toward the center be- coming lobulate; upper surface smooth, cracked with age and with conspicuous marginal pycnidia; lower surface black, sparsely to moderately rhizin- ate. Apothecia common, adnate, 2-3 mm in diam- eter; spores 6p X 12p. CHEMIsmY.-Cortex Kf yellow, medulla K+ red, P+ orange (atranorin and salazinic acid). DISTRIBUTION.-Panama. HABITAT.-on trunks and branches of hardwood trees in open pastures at about 1500 m elevation. REMARKS.-ThiS species is still known only from Panama, where it is rather common in the Chiriqui area. While at first sight similar to H. sublaevigata, the lobes are larger and more irreg- ular, pruina does not develop, and the chemistry is different. SPECIMENS EXAMINED.-Panama: Chiriqui, Hale 38811, 38831, 38848, 38902. 5. Hypotrachyna brasiliana, new combination FIGURE 7e Parmelia brasilianu Nylander, 1883:611. [Type collection: Organ Mountains, Brazil, Weddell (H, lectotype; P, iso- lectotype).] Parmelia glaziouii Muller Argau, 1889:354. [Type collection: Rio de Janeiro, Brazil, Glaziou 16670 (G, lectotype).] Thallus loosely adnate on rocks, coriaceous, ashy white, 6-20 cm in diameter; lobe sublinear, divaricate, 1-4 mm wide; upper surface plane, continuous, irregularly wrinkled and cracked with age; lower surface sparsely to moderately rhizinate, rhizines sparsely dichotomously branched. Apo- thecia common, adnate, 2-5 mm in diameter; spores 4y-6p X 8p-1Oy. red (lichexanthone and protocetraric acid). CHEMISTRY.-cOrteX K-, medulla K-, c-, Pf DIsTRIsuTIoN.-SoUtheaStern Brazil. HABITAT.-On rocks at open rocky slopes at 1000-2300 m elevation. REMARKs.-HyPotrachyna brasiliana is one of the more conspicuous and commonly collected saxicolous lichens in the Rio area. It grows loosely, often on sandstone, forming large whitish colonies that are easily detached. Two related species in the same habitats, H. contradicta (lich- exanthone absent) and H. eitenii (medulla Cf red), would have to be distinguished by chemical tests. SPECIMENS EXAMINED.-Brazil: Mato Grosso, iMalme 309, 312, 321B, 2246B (S); Minas Gerais, 26 SMITHSONIAN CONTRIBUTIONS TO BOTANY Eiten 7028, 7031 (US), Malme 312A (UPS), iMosen 2319bis (UPS), Vainio 1184 (BM, M, TUR, UPS), 1189 (M, TUR), 1249 (TUR), 1280 (BM, TUR, UPS); Rio de Janeiro, Eiten 6594A (US); Santa Catarina, Reitz and Klein 10187 (US), Ule 155 (G); SHo Paulo, Schiflner (W). 6. Hypotrachyna brevirhiza, new combination FIGURE 7f Parmelia breuirhiza Kurokawa in Hale and Kurokawa, 1964: 166. [Type collection: Isla Riesco, Magallanes, Chile, Sun- tesson 2066 (S, holotype; US, isotype).] Thallus closely adnate, ashy white, 3-6 cm broad; lobes subirregular and short, 1-5 mm wide; upper surface plane to rugulose, dull, frequently white-pruinose, sorediate toward the lobe tips and in part laminally, the soredia coarse and rather diffuse; lower surface densely rhizinate, the rhiz- ines densely dichotomously branched. Apothecia very rare, adnate, to 5 mm in diameter; spores poorly developed, 3p-4~ X 6p-8~. CHEMIsmY.-Cortex K+ yellow, medulla K+ yellow turning red, P+ orange (atranorin and salazinic acid). DISTRIBUTION.-PantrOpiCal in equatorial re- gions and in southern Chile and Argentina. HABITAT.-on trees in mist forest, secondary forest, and on rocks at 2000-3800 m elevation (but near sea level in Chile). REhfARKs.-Additional collections of this Species were made in the Pacific region and Africa after it was described from Chile. It is essentially pan- tropical in montane regions. The rather diffuse soralia, very dense rhizine mat, and tendency to form coarse pruina are characteristic features. Closely related H. majoris (Vainio) Hale, known from Madagascar and eastern Asia, consistently produces zeorine in addition to salazinic acid. A negative KOH test would separate H. pseudo- sinuosa from H. breuirhiza. SPECIMENS ExAMINED.-Venezuela: Merida, Hale 42377; Tachira, Hale 42532. Ecuador: Pinchin- cha, Knight 1084B (MiIS). Chile: Santiago, Mahu 2133 (US). Other records from Chile and Argen- tina are listed in Hale and Kurokawa (1964:167). 7. Hypotrachyna caraccensis, new combination FIGURE 8a Parmelia caraccensis Taylor, 1847: 163. [Type collection: Ca- racas, Venezuela, Linden 576 (FH-TAYL, lectotype; BM, G, isolec totypes).] Parmelia sinuosa var. caraccensis (Taylor) Lindsay, 1859:218. Parmelia caraccensis var. guatemalensis Steiner, 1903:234. [Type collection: Guatemala, Friedrichsthal (W, lectotype).] Parmelia caraccensis yar. guatemalensis f. adspersa Zahlbruck- ner, 1905:82. [Type collection: El Altar, Ecuador, Meyer 341 (W, lectotype).] Parmelia endorubra Gyelnik, 1934: 154. [Type collection: Peru, Lillie 1040 (BP, holotype).] Thallus loosely attached, massicot yellow, 6-30 cm in diameter; lobes long-linear, separate, 2-6 mm wide; upper surface plane, continuous, lack- ing soredia and isidia; lower surface densely rhizinate, the rhizines densely dichotomously branched, forming a projecting mat along the lobes. Apothecia adnate, 3-9 mm in diameter; spores 6p-8~ X 11p-13p. orange (usnic acid, galbinic acid, norstictic acid, and salazinic acid). DIsTRIBuTIoN.-Central America southward in the Andes to Bolivia and in southeastern Brazil. ses at 1500-4200 m elevation. REMARKs.-This must be regarded as the most conspicuous and typical of all the Hypotrachynae in the paramo region. It grows in large mats on exposed humus or among grass tussocks. When pressed and dried it tends to fragment. It could only be confused with unusually large or loosely adnate specimens of H. enderythraea, which has imbricate, smaller lobes, or with H. lopezii, which contains stictic acid. The syntype from Quito, Ecuador (Jameson 1845), is this species. The spores of var. guate- malensis are given as 14p-20p long, larger than in other material examined, but I doubt that this difference justifies varietal rank. SPECIMENS ExAMINED.-costa Rica: Cartago, Al- meda 737e (DUKE), Brenes 34 (US), Dodge (US), Standley 35221 (US); San Jose, Crosby 3928 (DUKE). Panama: Chiriqui, Kozlousky and Saw- ~PY 178 (lVIS), Scholander (US). Colombia: Boy- aca: Grzibb and Guymer L85b (BM); Cundina- marca, Barclny 6107 (US), Lindig 2594 (BM, M, CHEMISTRY.-cOrteX K-, medulla K+ red, P+ HABITAT.-On hardwood trees, humus, and mOS- NUMBER 25 FIGURE 8.-Species of Hypotrachyna: a, H. caraccensis (Hale 42446); b, H. chicitae (Culberson 13210); c, H. chlorina (Eiten 7671b); d, H. consirnilis (Eiten 7293); e, H. contradicta (Reitz and Klein 15727); f, H. costaricensis (Hale 42139). (Scale in mm.) 21 28 ShfITHSONIAN CONTRIBUTIONS TO BOTANY P), Soderstrom 1305 (US), Weir 17 (BM); San- tander, Ariste (US), Killip and Smith 17562 (US). Venezuela: Merida, Dennis 1925 (BM, US), Hale 42446, 42676, 42862, 43063, Nilson 1523 (UPS), Oberwinkler 13171b (M), Vareschi 946, 1472 (US). Peru: Holligan 60B (BM), Lechler 2728 (G, H, M). Ecuador: Jameson 137 (BM, US). Bolivia: La Paz: Julio 416 (US). Brazil: Minas Gerais, Damazio 13 (TUR); Rio de Janeiro, Lobb (BM). 8. Hypotrachyna chicitae, new combination FIGURE 8b Parmelia chicitae Hale, 1971:30. [Type collection: Asunci6n, Costa Rica, Culberson 13210 (US, holotype; DUKE, TNS, isotypes).] Thallus loosely to closely adnate, fragile, often forming a thick, cushion-like mass, ashy white or blackening toward the center, 4-10 cm broad; lobes sublinear, elongate, often crowded, 1.5-4 mm wide; upper surface plane, shiny, lacking soredia and isidia but with numerous pycnidia; lower surface densely rhizinate, the rhizines sparsely dichotomously branched. Apothecia rare, adnate, 3-6 mm in diameter; spores 5p X 6p-7p. C- (or C+ faint rose), KC+ rose, P- (atra- norin, evernic acid, lecanoric acid, obtusatic acid, and norobtusatic acid). DISTRIBUTION.-cOStB Rica, Venezuela, and Peru. HABI-rAT.-On rocks, soil, or mosses in paramo at 3300-3600 m elevation. REMARKS.-The saxicolous habitat at high ele- vations and unusual chemistry set this species apart from other members of the evernic acid con- stellation. As Culberson and Hale (1973) dis- covered, it falls midway in chemical evolution in the group. Chromatography of the complex mix- ture of acids is difficult. It could be confused with only one other paramo species, H. protenta, which contains alectoronic acid (C-). SPECIMENS EXAMINED.-Venezuela: Mkrida, Han- selmann 50 (DUKE, US). Other records from Costa Rica, Venezuela, and Peru are listed by Hale (1971b:31). CHEMISTRY.-cOrteX Kf yellow, medulla K-, 9. Hypotrachyna chlorina, new combination FIGURE 8c Parmelia chlorina Muller Argau, 1880:267. [Type collection: Petropolis, Brazil, Deuenter (G, lectotype).] Thallus adnate to loosely adnate, quite fragile, greenish or yellowish gray, to 8 cm broad; lobes short, crowded, sublinear, 2-5 mm wide; upper surface plane, shiny, becoming densely lobulate toward and at the margins, the lobules delicate, elongate, simple or furcate; lower surface mod- erately rhizinate. Apothecia rare, substipitate, 3-7 mm in diameter; spores 7p X 17p, the asci turning blue with IKI. CHEMIsmY.-Cortex K+ yellow, medulla Kf, C+ yellowish, P- (atranorin, barbatic acid, obtusatic acid, enthothein, and unidentified pigments). DIsrRmuTIoN.-West Indies, Panama, Venezu- ela, Peru, and Brazil. HABITAT.-On trees, mossy logs, or rocks in cloud forest or secondary forest at 900-2500 m elevation. REMARKS.-The yellow medulla and production of dense lobules (Figure 4e) are characteristic of this species. The lobules have the same pored epi- cortical structure as the main thallus, as revealed with the scanning-electron microscope. The chem- istry places it in the H. physcioides complex but there are no close relatives, although on hasty examination it might be misidentified as H. isidiocera, a clearly isidiate species, on account of the similar yellow medulla. SPECIMENS EXAMINED.-Haiti: Ouest, Zmshaug 22740 (MSC, US). Panama: Chiriqui, Hale 38754, 38770. Venezuela: Merida, Hale 42735, 43014. Peru: Cuzco, Hewera 3466a (US). Brazil: Minas Gerais, Eiten 6940, 7052 (US); Rio de Janeiro, Eiten 6573, 751 9 (US). 10. Hypotracltynn consirnilis, new combination FIGURE 8d Parmelia consirnilis Vainio, 1890:58. [Type collection: Carassa, Minas Gerais, Brazil, Vainio 1295 (TUR, lectotype; BM, isotype).] Thallus adnate on bark, 3-6 cm broad, light mineral gray but turning light tan in the her- NUMBER 25 29 barium; lobes sublinear, becoming crowded, 1-3 mm wide; upper surface plane, continuous, be- coming densely isidiate, isidia mostly simple; lower surface moderately rhizinate, the rhizines densely dichotomously branched. Apothecia rare, adnate, 3-5 mm in diameter, the amphithecium isidiate; spores 5y-8y X 8y-1111. CHEMIsmY.-Cortex K+ yellow; medulla K-, C -, P + orange (atranorin, protocetraric acid, and, if pigmented in lower medulla, rhodophyscin). DIsTRIBUTI0N.-Mexico, West Indies, Venezuela, Brazil. HABITAT.-On hardwoods and conifers in open woods at 800-1600 m. REhfARKs.-Hypotrachyna consimilis is as rare in the American tropics as its presumptive pro- genitor H. bahiana. It is smaller and more com- pact than H. imbricatula (P-). SPECIMENS EXAMINED.-MeXiCO: Chiapas, Hale 20375. Puerto Rico: Home 1162 (US). Domini- can Republic: La Vega, AZZard 17469 (US). Venezuela: Valencia, Magdefrau 493 (M). Brazil: Minas Gerais, Vainio 1133 (BM, K, M, TUR, UPS); Rio de Janeiro, Eiten 7293 (US). 11. Hypotrachyna contradicta, new combination FIGURE 8e Parmelia contradicta Hale, 1974b:265. [Type collection: Teresopolis, Rio de Janeiro, Brazil, Watson 521 (BM, holo- type; US, isotype).] Thallus loosely attached on rock, rather cori- aceous, whitish mineral gray; lobes linear and elongate, dichotomously branched, 1.5-2.0 mm wide, lacking soredia and isidia; upper surface plane and shiny; lower surface black, sparingly rhizinate, the rhizines coarse, sparsely dichotom- ously branched. Apothecia abundant, adnate, 2-4 mm in diameter; spores 4p X 6y. CHEMISTRY.-MedUlh K-, P + orange-red (atranorin and protocetraric acid). DIsTRInvTIoN.-Southeastern Brazil. HABITAT.-on open sandstone outcrops at about 1000 m elevation. REMARKs.-The absence of yellow-fluorescing lichexanthone in the cortex separates this species from the more widespread H. brasiliana; other- wise, the two species are very close, A possible relative occurs in South Africa: H. fissicarpa (Kurokawa) Hale, which has large, radially split apothecia and more imbricate, sublinear lobes. SPECIMENS EXAMINED.-Brazil: Santa Catarina, Reitz and Klein 15727 (US). 12. Hypotrachyna costaricensis, new combination FIGURE 8f Parmelia costaricensis Nylander in Polakowsky, 1877:225. [Type collection: Angostura, Costa Rica, Polakowsky (H, lectotype) .] Parmelia hypotrachyna Nylander, 1860:405. [Type collection: Tolima, Colombia, Goudot (P, lectotype).] Parmelia sublaeuigata f. isidiosa Muller Argau, 1880:267. Parmelia tropica Vainio, 1896:33. Parmelia tropica var. deformis Vainio, 1896:33. Parmelia deformis (Vainio) Vainio, 1907: 169. Parmelia amoena Zahlbruckner, 1908:464. [For full citations, see Hale, 1971a:ll.] Thallus adnate to loosely adnate, growing on rocks or trees, whitish to greenish mineral gray, 6-10 cm in diameter; lobes sublinear to irregular, often imbricate, 2-6 mm wide; upper surface plane, usually strongly maculate, moderately isid- iate, the isidia cylindrical, erect, simple to branched; lower surface moderately to densely rhizinate, forming a thick mat visible along the margins from above, the rhizines densely dichot- omously branched. Apothecia rare, adnate, 2-7 mm in diameter; spores 4y-6y X 8y-1011. CHEMIsTRY.-Cortex K + yellow, medulla neg- ative with all reagents (atranorin and protolich- esterinic and possibly caperatic acids). DIsrRIBuTIoN.-Mexico, Central America, West Indies, Andean region from Venezuela to Bolivia, Brazil, and Southeast Asia. HABITAT.-on deciduous trees, conifers, palm, rocks, and (rarely) soil, in scrub forests, secon- dary cloud forests, and coffee plantations at 1000- 3500 m elevation. REMARK^.-??^^^ trachyna costaricensis is the second most common species in the genus after H. imbricatula in tropical America. It can be recognized by the uniformly isidiate surface, often with strong white maculation, dense rhizine mat, and negative chemical reactions. It occurs over a wide range of habitats and elevation. The fatty acid constituent has not been precisely identified. According to Dr. Chicita Culberson, it probably is near caperatic acid but has a different R, value. The oldest name, Parmelia hypotrachyna, is SMITHSONIAN CONTRIBUTIOXS TO BOTANY invalid in Hypotrachyna as a tautonym. It is rep- resented by a rather unusual specimen containing traces of gyrophoric acid along with a fatty acid, but no other specimens contain this acid. Zahl- bruckner (1929) transferred it to Anzia without seeing the type, apparently since Nylander men- tioned its affinity with Anzia colpodes, presumably because of the dense hypothallus-like rhizine mat. Zahlbruckner also described P. amoena in 1908 without having checked the type of P. tropica, another synonym, which he guessed was close. SPECIMENS EXAMINED.-Mexico: Chiapas, Hale 20036, 20046, 20479; Mexico, Cain 27597 (TNT, US); Michoacan, Hale 20860, 20966; Veracruz, Liebmann 7562d (TUR). Guatemala: Alta Vera- paz, Johnson 787 (US), Standley 70064, 70347 (US); Baja Verapaz, Hale 38358, 38617. Costa Ria: Cartago, Standley 34985 (US), Stone (DUKE); San Jose, Standley 47329 (US). Pan- ama: Chiriqui, Hale 38728, 38732, 38783, 38814, 38851, 38928, Scholander (US), Wagner 42 (M). Cuba: Las Villas, Imshaug 24619, 24633A (MSC); Oriente, Imshaug 2471 1 (MSC, US). Jamaica: Imshaug 13926 (MSC, US), 14368, 14650, 14912, 16225 (MSC), Orcutt 5657 (US). Haiti: Ouest, Zmshaug 22596 (MSC). Dominican Republic: La Vega, Imshaug 23814 (MSC). Dominica: Elliott 276 (lectotype of P. tropica Vainio) (BM, TUR), 12 (lectotype of P. tropica var. deformis Vainio) (BM, TUR), Hale 35596a. Martinique: Degelius (DEGEL). St. Vincent: Elliott (TUR). Grenada: Imshaug 16235 (MSC, US), 16260, 16271 (MSC). Colombia: Santander, Cuatrecasas et al. 12470 (US), Killip and Smith 19385 (US), Lindig 2531 (UPS); Tolima, Quimby (US). Venezuela: Federal, Santesson 6678b (S). Merida, Hale 42201, 42242, 42271, 42738, 43301, Ober- winkler and Poelt (US). Peru: Amazonas, Wur- dack 483 (US); Cuzco, Thomasson (UPS). Bo- livia: La Paz, Eyerdam 25139 (F). Brazil: Minas Gerais, Damazio 1453 (lectotype of P. amoena Zahlbruckner) (WU, BM, H), Eiten 6954 (US); Rio de Janeiro, Ainsworth 557 (BM, US), De- venter 49 (lectotype of P. sublaevigata f. isidiosa Miiller Argau) (G), Eiten 7671 (US). 13. Hypotrachyna croceopustulata, new combination FIGURE 9a Parnielia croceopustulata Kurokawa in Hale and Kurokawa, 1964: 169. [Type collection: Grandfather Mountain, Avery County, North Carolina, Irnshaug 22275 (MSC, holotype; US, isotype).] Thallus closely adnate to bark, whitish mineral gray, 4-8 cm in diameter; lobes sublinear, becom- ing crowded, 1.5-3.5 mm in width; upper surface plane to rugulose, sorediate-pustulate, the soralia coarse and coalescing, mostly laminal, the medulla below often yellow ochre in color; lower surface moderately rhizinate. Apothecia rare, adnate, 1-6 mm in diameter; mature spores lacking. C -, P + orange-red (atranorin, protocetraric acid, and rhodophyscin). DIsTRIBuTI0N.-Southeastern United States, Mex- ico, West Indies. HABITAT.-on conifers (Abies, Picea, Pinus) in open forests at 1400-2000 m ,elevation. REMARKS.-when first described, Parmelia croceo- pustzilata was based in part on Hypotrachyna gondylophora, which is also Pi- red but contains fumarprotocetraric acid but which, more signifi- cantly, has strictly subterminal soralia. This error was later corrected (Hale, 1967) and H. croceo- pustulata was typified as including only laminally pustulate-sorediate plants (Figure 2c) with proto- cetraric acid. It is a relatively rare species, perhaps most common on conifers in the southern Appa- lachian Mountains but absent from South America. Nakanishi I79 (US). Haiti: Ouest, Wetmore 3213 (MSC, US); Sud, Zmshaug 23229 (MSC, US). CHEMISTRY.-COrteX K+ yellow, medulla K-, SPECIMENS EXAMINED.-MeXiCO: Tamaulipas, 14. Hypotrachyna dactylifera, new combination FIGURE 9b Parrnelia dactylifera Vainio, 1890357. [Type collection: Lafay- ette, Minas Gerais, Brazil, Vainio 363 (TUR, lectotype; BM, FH, M, P, UPS, isolectotypes).] Parrnelia tilacen var. leucina Muller Argau, 1880:267. [Type collection: Petropolis, Brazil, Deventer (G, lectotype).] Thallus closely adnate, tannish mineral gray, 3-8 cm broad; lobes sublinear, often crowded, 1-4 mm wide; upper surface plane, continuous, with age becoming rugulose, becoming densely isidiate, the isidia very large, about 1 mm high, fragile and usually breaking open and pustulate apically without formation of soredia; lower sur- NUMBER 25 31 face moderately rhizinate, the rhizines densely dichotomously branched. Apothecia rare, adnate to substipitate, 1-4 mm in diameter; spores CHEMIsmY.-Cortex K+ yellow, medulla K-, C-, KC+ rose, P- (atranorin, colensoinic acid, norcolensoinic acid, 4-0-demethylphysodic acid, physodic acid, lividic acid, and associated un- knowns, and, if pigmented in the lower medulla, rhodophyscin). DIsTRIBuT1oN.-Mexico, Costa Rica, Panama, Venezuela, Chile, Brazil, Uruguay, and Africa. HABITAT.-On trees (hardwoods and conifers) and rarely on rocks, in open secondary forest or cloud forest at 1000-2800 m elevation. REMARKs-The isidia of this species are very large and inflated and probably are correctly in- terpreted as pustules. They are exceedingly var- iable and the range of variation is still unsettled in spite of the fair number of collections available. The chemistry is identical with that of the pre- sumptive progenitor H. livida. Vainio (1890) surmised that Parmelia sublaevigata f. isidiosa Muller Argau might belong here, but it is a syn- onym of H. costaricensis. SPECIMENS EXAMINED.-kfeXiCO: Chiapas, Hale 20288, 20375; Durango, Weber 33708 (COLO); Veracruz, Hale 19520. Costa Rica: Cartago, Torres 156 (F). Panama: Chiriqui, Hale 38722, 38927, Scholander (US). Venezuela: Federal, Santesson 6677a (S); Merida: Hale 42042, 43252, Poelt 7525 (US). Brazil: Rio de Janeiro, Dusen 11 (S), Eiten 6529, 6534, 7126 (US); Sao Paulo, Sampaio (P). Uruguay: Osorio 5925 (MVM). Chile: Chi- loe, Lobb (BM). 5p-7~ X 9p-13p. 15. Hypotrachyna degelii, new combination FIGURE 9c Parmelia degelii Hale in Hale and Kurokawa, 1964:170. [Type collection: Angola, Mexico, Degelius (Degelius herbarium, holotype: US, isotype).] Thallus rather closely adnate, coriaceous, tan- nish mineral gray in the herbarium, 4-8 cm broad; lobes short, crowded, sublinear, 1-2.5 mm wide; upper surface plane to rugulose, shiny, faintly white-maculate, lacking soredia and isidia; lower surface moderately rhizinate. Apothecia com- mon, adnate to substipitate, 3-8 mm in diameter, the amphithecium white-maculate, incised; spores CHEiwsmY.-Cortex K+ yellow, medulla K-, C--, KC+ rose, P- (atranorin, alectoronic acid, and a-collatolic acid). DISTRIBUTION.-Colombia, Peru, Brazil, and Africa. HABITAT.-On trees, rarely rocks, in open forest at 1000-3500 m elevation. REMARKs.-Hypotrachyna degelii resembles other short-lobed species such as H. bahiana (proto- cetraric acid present) and H. physcioides (barbatic acid group present) and must be identified with a chemical test. It is biogenetically unrelated to these species and probably falls closer to the H. Zivida group. The sorediate morph is H. exsplendens. SPECIMENS ExAMINED.-co~ombia: Cundin- amarca, King 505 (US). Peru: Bryan 703 (S, US, WIS). Brazil: Mato Grosso, Malme 2545" (S); Rio Grande do Sul, Malme 941 (S). 4p-5p x 8p-llp. 16. Hypotrachyna densirhizinata, new combination FIGURE 9d Parmelia densirhizinata Kurokawa in Hale and Kurokawa, 1964: 171. [Type collection: Volcan, Chiriqui, Panama, Scholander (US, holotype).] Thallus loosely attached, whitish mineral gray, 6-20 cm broad; lobes linear-elongate, often divar- icate, separate, 2-6 mm wide; upper surface plane, continuous, sorediate-pustulate near the tips, the soralia subcapitate; lower surface densely rhizinate, the rhizines projecting as a mat along the mar- gins, densely dichotomously branched. Apothecia very rare, substipitate, to 10 mm in diameter; spores lop X 17 p. CHEMIsmY.-Cortex K+ yellow, medulla K-, C--, KC+ rose, P- (atranorin, alectoronic acid, and a-collatolic acid). DIsTRIBuTIoN.-Southeastern United States, Mex- ico, Central America, West Indies, Colombia, Venezuela, Ecuador, Peru, Bolivia, Chile, Argen- tina, and Africa. HABITAT.-& trees (open pine forests, Drimys, Baccharis), rocks, and mossy humus in open areas and on roadside banks at 2000-3700 m elevation. REMARKS.-The lobes of this species are not as elongate on the average as those of its presump- tive progenitor H. gigas. It has a much wider 32 ShIITHSONIAN CONTRIBUTIONS TO BOTANY FIGURE 9.--Species of Hypotrachyna: a, H. croceopustulata (Imshaug 22383); b, H. dactylifera (Hale 42075); c, H. degelii (Bryan 703); d, H. densirhizinata (Hale 43194); e, H. dentella (Me- Cullough 570); f, H. ducalis (Holligan L59B). (Scale in mm.) NUMBER 25 33 geographic range than H. gigas-as we might ex- pect for a sorediate morph-from the cold temper- ate mountains of the southern Appalachians to southern Chile. The chemistry is uniform, and no specimens contain gyrophoric acid, strongly sug- gesting that H. densirhizinata originated before H. gigas interbred with a gyrophoric acid- containing population. Without a chemical test H. densirhizinata could be confused with any of the species in the chemical complex including H. laevigata, H. oostingii, and H. thysanota. SPECIMENS EXAMINED.-MexicO: Oaxaca, Sharp et al. 2684 (NICH). Guatemala: Solola, Keller- man (OSU, US). Costa Rica: Cartago, Culberson 12416 (DUKE). Colombia: Tolima, Quimby (US). Venezuela: Merida, Hale 42389, 42695, 42808, 43194, Hanselmann 62 (DUKE, US). Chile: Juan Fernandez, Imshaug 37469 (MSC); Valdivia, San- tesson 3604 (S, US). Argentina: Lamb 6003 (UPS). For additional records from Guatemala, Panama, Haiti, Dominican Republic, Colombia, Peru, Ecuador, Bolivia, and Chile see Hale and Kurokawa (1964: 171-172). 17. Hypotrachyna dentella, new combination FIGURE 9e Parrnelia dentella Hale and Kurokawa, 1964: 172. [Type col- lection: Cheaha State Park, Clay County, Alabama, Mc- Cullough 570 (US, holotype; TNS, isotype).] Parmelia laevigata var. ceratina Muller Argau, 1880:267. [Type collection: Petropolis, Brazil, Deventer (G, lecto- type).] Thallus loosely adnate on rocks, 3-8 cm broad, greenish mineral gray; lobes subirregular to lin- ear, short, 2-6 mm wide; upper surface plane, continuous, often quite white-maculate, densely isidiate, the isidia simple to branched, up to 0.7 mm high; lower surface sparsely to moderately rhizinate, the rhizines sparsely dichotomously branched. Apothecia rare, adnate, 1-2 mm in diameter; spores 4p-6~ X 8p-101.1. CHEMIsmtY.-Cortex K+ yellow, medulla K+ yellow, C- or C+ pale yellow, P+ orange (atranorin, barbatic acid, echinocarpic acid, 4-0- demethylbarbatic acid, obtusatic acid, and norob- tusatic acid). DIsmtIsuTIoN.-Southeastern United States, Mex- ico, Venezuela, and Brazil. HABITAT.-On rocks in open woods and pas- tures at about 1000 m elevation. REMARKS.-This species is closely related to H. imbricatula but differs in being smaller and in producing P + echinocarpic acid. SPECIMENS EXAMINED.-Mexico: Tamaulipas, Nakanishi 11 1 (US). Venezuela: Merida; Hale 42727. Brazil: Rio de Janeiro, Eiten 6532 (US). 18. Hypotrachyna ducalis, new combination FIGURE 9f Parmelia ducalis Jatta, 1908:407. [Type collection: Mobuku Valley, Nabitava verso Kirchuchu, and Duroni TV of Valle dei Laghi, Ruwenzori, Aputi (not seen).] Thallus loosely adnate, whitish mineral gray, about 10 cm broad; lobes linear-elongate, separate to divaricate, 2-5 mm wide; upper surface plane, shiny, soredia and isidia lacking; lower surface densely rhizinate, the rhizines densely dichot- omously branched. Apothecia seen only in Afri- can material: 3p-4p X 8p-9p. CHEMIsmY.-Cortex K+ yellow, medulla K- C+ red, P- DIsTRIsurIoN.-Venezuela, Peru, and Africa. HABITAT.-oVer mosses and humus at the base of trees in thickets in open areas and paramo at 3000-3800 m elevation. REMARKS.-The type of H. ducalis has not yet been found, but Jatta's description clearly refers to a nonsorediate linear-lobed species similar to other collections made in the mountains of Kenya and Uganda. In any event, in chemistry and morphology the population in South .hierica is identical to the population in Africa. Externally, H. ducalis is close to H. gigas, which contains alectoronic acid. Vegetative morphs include H. producta (sorediate), H. partita (isidiate-lob- date), and H. rachistn (lobulate). SPEcIivENs EXARIINED.-veneZuela: Merida, Hale 43059. Peru: San hiartin, Hollignn L59B (BII). (atranorin and anziaic acid). 19. Hypotrachyna eitenii, new combination FIGURE 10n Pnrrnelin eitenii Hale, 1974b:2GG. [Tlpe collection: Seirn dos Orgaos, Rio de Janeiro, Brazil, G. n~d I-. Eiteti 7123 (L'S,.] Thall~i~ loosely attached to rocks, brittle. min- 34 SMITHSONIAN CONTRIBUTIONS TO BOTANY era1 gray, about 6 cm in diameter; lobes sublinear, dichotomously branched, about 3 mm wide; upper surface plane, without soredia or isidia; lower surface sparsely to moderately rhizinate, the rhiz- ines long. Apothecia abundant, adnate, to 4 mm in diameter; spores 5p X 7p-8y. CHEMIsmY.-Cortex K+ yellowish, medulla K- , Cf red, P- (trace atranorin, lichexanthone, and anziaic acid). DIsmIsuTI0N.-Southeastern Brazil. HABITAT.-Rocks in scrubby forest at about 1800 m elevation. REMARKS.-ThiS saxicolous species is very close to H. brasiliana in external appearance and in habitat. As a possible recent derivative of the H. brasiliana group, it diverges very significantly in chemical biogenesis. 20. Hypotrachyna enderythraea, new combination FIGURE 10b Parmelia enderythraea Zahlbruckner, 1929: 136. [Type collec- tion: Based on P. erythrocardia Zahlbruckner.] Parmelia erythrocardia Zahlbruckner, 1908, p. 466. [Type col- lection: Camarinhas, Brazil, Damazio 1761 (W, lectotype). Not P. erythrocardia (Miiller Argau) Vainio, 1900:7.] Thallus adnate to loosely adnate on rock or soil, massicot yellow, 4-8 cm in diameter: lobes variable, linear-elongate, often crowded, 1-3 mm wide; upper surface plane, continuous, sometimes pruinose at the tips; lower surface moderately rhizinate, the rhizines often projecting beyond the margins. Apothecia rare, adnate, 2-15 mm in diameter; spores 7p-8p X 131.1-151.1. CHE;\IIsmY.-Cortex K-; medulla K+, Pf orange to red (galbinic, norstictic, salazinic, and usnic acids). DIsTRIBuTIoN.-Panama, Jamaica, Andean re- gion from Venezuela to Bolivia, and southeastern Brazil. HABITAT.-On soil and rocks in open forests and in the paramo at 2200-3700 m elevation. REMARKs.-Degree of lobe adnation and width are rather variable in this species. The lobes may he separate and elongate to imbricate and short, variation that may result from environmental modification, for it grows in exposed paramo areas. The chemi5try places this species close to H. microhlmta, a closely related isidiate morph. Hypotrachyna caraccensis is a good candidate for the sexual morph because of the identical chem- istry, but it is much larger and divaricate. Culbel-son 13730, 13733 (DUKE, US). Venezuela: Mkrida, Hale 42464, 42898, 43150, 43218, 43324, 43340 (US), Hanselmann 33 (DUKE, US), Ober- winkler and Poelt 7542, 7547 (M, US). Peru: Chachapoyas, Matthews (US), Wurdack 484 (US); Cuzco, Iltis 3313 (WIS), Thomasson (UPS). Bo- livia: Julio 414 (US). Ecuador: Fiske (US). SPECIMENS EXAMINED.-Jamaica: Culberson and 21. Hypotrachyna endochlora, new combination FIGURE 1Oc Parmelia endochlora Leighton, 1871:140. [Type collection: Askew Wood, Kerry, Ireland, Taylor (BM, lectotype).] Parmelia millaniana Stirton, 1874:79. [Type collection: Loch Long, Argyleshire, Scotland, Macmillan (BM, lectotype).] Parmelia xanthomyela Nylander, 1874:306. [Type collection: Fontainebleau, France, Nylander (H, lectotype).] Parmelia pittieri Muller Argau in Durand and Pittier, 1891: 53. [Type collection: Rancho Rotundo, Costa Rica, Pittier 5050 (G, lectotype).] Parmelia madagascariensis Vainio, 1898: (33). [Type collec- tion: Tanala, Ambohimitombo, Madagascar, Forsyth 562 (TUR, lectotype; BM, isolectotype).] Parmelia xanthomyela f. sorediosa Hue, 1899: 146. [Type col- lection: Mafate, Bourbon, Rodriguez (P, lectotype).] Parmelia endochlora f. sorediosa (Hue) Zahlbruckner, 1929: 166. Parmelia gallicana Gyelnik, 1931:288. [Type collection: Based on Parmelia xanthomyela f. sorediosa Hue).] Thallus loosely adnate, greenish mineral gray, 8-13 cm broad; lobes sublinear, margins some- times dissected with age, 2-6 mm wide; upper sur- face shiny, plane, continuous, faintly to strongly white-maculate, pustulate, the pustules mainly suhterminal, soon turning granular-sorediate; me- dulla pale cendre green; lower surface moderately to densely rhizinate, the rhizines densely dichot- omously branched. Apothecia rare, adnate, 4-14 mm in diameter; spores 9y-14p X 18p-22p. C+ orange, P- (atranorin, barbatic acid, ob- tusatic acid, norobtusatic acid, entothein, unde- termined pigments, rarely echinocarpic acid). DIsmIBuTIoN.-Mexico, Central America, Co- lombia, Venezuela, Brazil, western Europe, South Africa, Madagascar, St. Helena, Hawaii. HABITAT.-on trees, rarely rocks, and over mos- CHEMISTRY.-cOTteX K+ yellow, medulla K- , NUMBER 25 35 ses in mature forests, primary cloud forest, and scrubby areas at 1700-3500 m elevation. REMARKs.-This widespread species is very sim- ilar to H. laeuigata except for the yellowish me- dulla. It occurs over a broader range, however, including Africa and the Pacific. The pigmenta- tion is apparently the same as that in H. chlorina and H. isidiocera but the results with TLC are unsatisfactory for resolution. Actually the chem- istry is variable, with one or two of the minor constituents and echinocarpic acid sometimes absent. SPECIMENS EXAMINED.-Mexico: Michoacan, Hale 20973. Costa Rica: San Jose, Culberson 12572 (DUKE). Panama: Chiriqui, Hale 38931, Scho- lander (US). Dominican Republic: La Vega, Wet- more 3509 (MSC). Colombia: Tolima, Quimby 37 (US). Venezuela: Mkrida, Hale 43261. Brazil: Rio de Janeiro, Eiten 6577, 7093, 7709 (US). Uruguay: Higgins (BM). 22. Hypotrachyna ensifolia, new combination FIGURE 10d Parmelia ensifolia Kurokawa in Hale and Kurokawa, 1964: 172. [Type collection: Alto de la Bandera, La Vega, Dominican Republic, lmshaug 23430 (MSC, holotype; US, isotype).] Parmelia lobulifera var. insensitiva Degelius, 1941:63. [Type collection: Forney Ridge, Swain County, North Carolina, Degelius (Degelius herbarium, lectotype).] Thallus adnate, whitish mineral gray, 5-11 cm broad; lobes sublinear, dichotomously branched, 1-5 mm wide; upper surface shiny, continuous, becoming densely isidiate-lobulate on the surface and along the margins, the lobules up to 1.5 mm long, often ciliate; lower surface moderately rhizinate, the rhizines densely dichotomously branched. Apothecia rare, adnate, 3-4 mm in diameter; spores 8p-lop X 14p-18p. CHEMIsmY.-Cortex K+ yellow, medulla K--, C--, KC+ rose, P- (atranorin, alectoronic acid, and a-collatolic acid or with a-collatolic present in trace amounts or lacking). DIsmIBuTIoN.-Costa Rica, Panama, West In- dies, Colombia, Venezuela, and New Zealand. HABITAT.-on trees or boulders in cloud forest or on soil banks at 2000-3700 m elevation. REMARKs.-Hypotrachyna ensifolia is character- ized by abundant lobulate isidia, often becoming ciliate. It may represent the isidiate morph of H. gigas although the lobes are shorter. It often occurs with morphologically similar H. bogotensis, which can be distinguished by a C+ rose color test or absence of white fluorescence in ultraviolet. berson 12360 (DUKE, US). Colombia: Santander, Killip and Smith 20281 (US). Venezuela: MCrida, Hale 42390, 42455, 42897, 43131, Oberwinkler and Poelt 7511 (M, US). Additional records from Mexico, Panama, Jamaica, Haiti, Dominican Re- public, and Venezuela are listed in Hale and Kurokawa (1964: 173). SPECIMENS EXAMINED.-cOSta Rica: Cartago, CUl- 23. Hypotrachyna erythrodes, new combination FIGURE 10e Parmelia erythrodes (Zahlbruckner) Hale and Kurokawa, 1964:173. Parmelia brasiliana var. erythrodes Zahlbruckner, 1909: 169. [Type collection: Mt. Itatiaya, Minas Gerais, Brazil, SchiB- ner (WIJ, lectotype).] Thallus adnate to closely adnate on rock, often brittle and forming fragile mats, whitish ivory or very pale yellowish ashy gray, 4-8 cm broad; lobes sublinear, becoming crowded and imbricate, 1-3 mm wide; upper surface plane, continuous, with- out soredia or isidia; medulla white or turning dull salmon; lower surface moderately rhizinate, the rhizines sparsely dichotomously branched. Apothecia common, adnate, 2-6 mm in diameter; spores 5p6p X 9p-12p. P - (lichexanthone, 4-0-demethylphysodic acid, and physodic acid, with or without lividic acid and colensoinic acid, and rhodophyscin with associated unknowns). DIsmIBvnoN.-Venezuela, Peru, and Brazil. HABITAT.-on rocks and stone walls in sparsely forested areas and in the paramo at 1600-3500 m elevation. REhfARKS.-This is one of the commonest saxi- colous lichens in the paramo along with H. osteo- leuca, which contains olivetoric acid. Both of these species contain lichexanthone in the cortex and have a distinctive whitish to pale cream color. Hypotyachyna erythrodes is differentiated by the presence of the lividic acid complex and frequent production of a pale salmon or orange pigment in the medulla. Almost half of the specimens from Venezuela were found to lack lividic acid CHEMISTRY.-cOrteX K--, medulla K-, c-, 36 SMITHSONIAN CONTRIBUTIONS TO BOTANY FIGURE 10.-Species of Hypotrachyna: a, H. eitenii (Eiten 7125); b, H. enderythraea (Hale 42464); c, H. endochlora (Hale 43261); d, H. ensifolia (Zrnshaug 23430); e, H. erythrodes (Hale 43321);f, H. exspendens (Standleyj. (Scale in mm.) NUMBER 25 57 and to moduce almost random assortments of the Based on P. frava Krempelhuber not P. paua Rebentisch, I remaining components. TLC cannot adequately resolve these substances, and a final analysis of the chemical variation will have to come with high- pressure liquid chromatography. SPECIMENS EXAMINED.-VeneZUeki: Merida, Den- nis 2370 (BM), Hale 42469, 42472, 42488, 42619, 42791, 42792, 42798, 42809, 42825, 42863, 43147, 43186A, 43321, 43329; Tachira, Hale 42556. Peru: Lobb (BM). Brazil: Minas Gerais, Eiten 7030 (US). 24. Hypotrachyna exsplendens, new combination FIGURE lOf Parmelia exsplendens Hale in Hale and Kurokawa, 1964: 174. [Type collection: Murdock?s Gap, Blue Mountains, Ja- maica, Zmshaug 15306 (MSC, holotype; US, isotype).] Thallus adnate on bark, tannish mineral gray, 4-8 cm broad; lobes short, sublinear, 1.5-2.5 mm wide; upper surface plane, shiny, strongly white- maculate, sorediate, the soralia subterminal, capi- tate; lower surface densely rhizinate, the rhizines densely dichotomously branched. Apothecia not seen. CHEMIsmY.-Cortex KS yellow, medulla K-, P-, C-, KC+ rose (atranorin, alectoronic acid, and a-collatolic acid). DIsmIBuTIoN.-Mexico, West Indies, Guatemala, Costa Rica, South Africa. HABITAT.-on trees in evergreen forests at 900- 1200 m elevation. REMARKs.-The diagnostic characters are sub- terminal capitate soralia, distinct white maculae, and presence of alectoronic acid. This species is much smaller and has more imbricate, adnate lobes than H. densirhizinata, another species with alectoronic acid. The presumptive progenitor is H. degelii, which also is known from Africa. (US). Jamaica: Culberson 13509 (DUKE). Other records from Mexico, Guatemala, and Jamaica are listed in Hale and Kurokawa (1964:175). SPECIMENS EXAMINED.-cOSta Rica: BreneS 47 1804:303.] Parmelia frava Krempelhuber, 1873: 10. [Type collection: Serra de Piedade, Brazil, Warming 294 (M, lectotype; G, UPS, isolectotypes).] Parmelia frava var. stellata Lynge, 1914: 150. [Type collection: Sao Joao d?el Rey, Minas Gerais, Malme 311 (S, lectotype).] Parmelia flava var. subdichotoma Lynge, 1914:149. [Type col- lection: SLo JoLo d?el Rey, Minas Gerais, Brazil, Malme 310 (S, lectotype).] Thallus adnate to loosely attached, coriaceous, sea-foam green, 3-5 cm broad; lobes sublinear, separate, 0.5-2 mm wide; upper surface plane to convex, continuous or irregularly cracked on older lobes; lower surface moderately rhizinate, the rhizines long, sparsely dichotomously branched, often projecting beyond the lobe margins. Apoth- ecia adnate, 1-3.5 mm in diameter; spores 5p-6~ CHEMIsmY.-Cortex K--, medulla K-, C-, or KCS rose, P+ orange-red (usnic acid and pro- tocetraric acid). DIsTRIsuTIoN.-Colombia, Venezuela, Peru, and Brazil. HABITAT.-On open sandstone outcrops or (rarely) on soil at 1200-4100 m elevation. REMARKS.-This saxicolous species has a fairly restricted distribution in northern South America. Vainio (1890) noted the branched rhizines in his descriptions and classified the species in section Xanthoparmelia because of the yellow color of the thallus. Krempelhuber (1873) reported the type on tree bark, but the actual specimens appear to be saxicolous. The great variability in lobe width led Lynge (1914) to describe two new varieties, both narrow-lobed. SPECIMENS ExA~fINED.-Co~ombia: Santander, Killip and Smith 15186 (NY, US), 17527 (US). Venezuela: hlerida, Hale 42121a, Oberwinkley and Poelt (hl, US). Peru: Lobb (Bhi). Brazil: Bahia, Blanchet (hi); hiinas Gerais, Damnzio 959 (TV), 1402 (G), Eiten 7003, 5027 (US), Hensclien (UPS, US), T?ainio 1163 (Bll, TUR, UPS), 1522 (Bhl, &I, TUR); Rio de Janeiro, Gaztdichaird 9 x lop-12p. (H). 25. Hypotrachyna flavida, new combination 26. Hypotrachyna flacocirens, new combination FIGURE lla FIGURE Ilb Parmelia Pauida Zahlbruckner, 1929: 137. [Type collection: Partnelia pnvovirens Kurokawa in Hale and Kurokawa, 196-1: 38 S.MITHSONIAN CONTRIBUTIONS TO BOTANY 176. [Type collection: 50 km SE Natales, Magallanes, Chile, Santesson 5681 (S, holotype; US, isotype).] Thallus adnate, sea-foam green, 3-6 cm in di- ameter; lobes sublinear, short and sometimes imbricate, 1-2.5 mm wide; upper surface plane, shiny, becoming rugulose with age, lobulate to- ward the center, sorediate, the soralia mainly subterminal, capitate; lower surface densely rhizinate, the rhizines densely dichotomously branched. Apothecia adnate, 24 mm in diameter, the amphithecium rugose; spores 71.1-lop X CmmsmY.-Cortex K-, medulla K-, C-, P + orange-red (usnic acid and protocetraric acid). DIsTRIBUTIoN.--Southern Chile. HABITAT.-oII trees (Nothofagus) and on rocks at low elevation. REMARKs.-This species is essentially indistin- guishable from the typical form of H. sinuosa, a pantemperate-pantropical montane species also common in the range of H. flavovirens. The chemical pattern (protocetraric or protocetraric with norstictic and salazinic acids) suggests a back- crossing between the progenitor of H. flavovirens (extinct?) and that of H. sinuosa (H. reducens). SPECIMENS EXAMINED.-chile: Magallanes, Sun- tesson 8191 (S, US); Tierra del Fuego, Santesson 7460 (S). Other records from Chile are listed in Hale and Kurokawa (1964: 176). 11p-15p. 27. Hypotrachyna formosana, new combination FIGURE llc Parmelia formosana Zahlbruckner, 1934:57. [Type collection: Mt. Arisan, Nimandaira, Taiwan, Asahina 70 (W, lectotype; TNS, isolectotype).] Thallus adnate to closely adnate, whitish min- eral gray, 3-8 cm in diameter; lobes sublinear, crowded, 1-3 mm wide; upper surface plane to rugulose, continuous, moderately to densely pus- tulate over most of the surface, the pustules break- ing open apically but not becoming sorediate; medulla white or ochre below the pustules; lower surface moderately rhizinate. Apothecia rare, ad- nate, the amphithecium pustulate, 3-5 mm in diameter; spores 4y-6p X 7p-llp. Cmarrsmy.-Cortex K-, medulla K+ reddish, C--, P- or P+ orange (lichexanthone, colen- soinic acid, norcolensoinic acid, 4-0-demethyl- physodic acid, physodic acid, lividic acid, and associated unknowns, rarely protocetraric acid, and, if pigmented, rhodophyscin). Central America, West Indies, Venezuela, Peru, Brazil, Africa, India, southeastern Asia, and Japan. HABITAT.-on trees (hardwoods and conifers) in upland cloud forest, secondary forests, and pine forests and on rocks in open areas at 900-3000 m in tropical regions and down to sea level in tem- perate areas. REMARKs.-HyPotrachyna formosana is pan- tropical with wide ecological amplitude, ocurring on both trees and rocks. The dense pustules and positive orange fluorescence under ultraviolet instantly identify it. It belongs to the H. livida complex and the sexual morph seems to be rare H. novella. The chemistry is very uniform and identical with that of H. liuida except for the rare occurrence of protocetraric acid. The second syn type of Parmelia formosana, (Asahina 60 from Taiwan) can be identified with H. exsecta (Tay- lor) Hale, an Asian species with barbatic acid. SPECIMENS EXAMINED.-Mexico: Chiapas, Hale 19942, 201 10, 20417, 20469, 20806; Michoacan, Hale 20852. Guatemala: Alta Verapaz, Standley 71865 (US); Baja Verapaz, Hale 38370, 38384; Huehuetenango, Standley 82325 (US). Panama: Chiriqui, Scholander (US). Cuba: Oriente, Ims- haug 24757, 24760 (MSC). Haiti: Ouest, Imshaug 22546 (MSC). Venezuela: Merida, Hale 42069, 42570. Peru: San Martin, Allard 22578b (US). Brazil: Rio de Janeiro, Eiten 6460, 7671a (US). Chile: Chiloe, Santesson 4107 (S); Valdivia, San- tesson 7261 (S). DISTRIBUTION.-EaStern United States, Mexico, 28. Hypotrachyna gigas, new combination FIGURE lld Parmelia gigas Kurokawa in Hale and Kurokawa, 1964:177. [Type collection: Bogota, Colombia, Joseph B7O (US, holo- type: DUKE, LD, MSC, REN, S, TNS, UPS, WIS, isotypes).] Parmelia exporrecta Kurokawa in Hale and Kurokawa, 1964: 174. [Type collection: Cerro San Felipe, Oaxaca, Mexico, Hale 20722 (US, holotype).] Thallus loosely attached and divaricate, rarely pendulous, 8-40 cm broad; lobes elongate, linear, separate, 3-10 mm wide; upper surface plane, con- tinuous, without soredia or isidia; lower surface NUMBER 25 FIGURE 1 1.-Species of Hypotrnchyna: n, H. flavida (Eiten 7003); 6, ?I. flnriorJirens (Satitussoti 5681); c, H. formosann (Asahina F70); d, H. gigns (Hole 42373); e, H. gotidylophorn (Hale 20879); f, H. gracikscens (Vainio 1917 in BM). (Scale in mm.) 39 40 SMITHSONIAN CONTRIBUTIONS TO BOTANY moderately to densely rhizinate, the rhizines con- spicuous as a mat along the lobe margins, densely dichotomously branched. Apothecia very rare, substipitate, up to 15 mm in diameter, the amphi- thecium rugose; spores 8p-lop X 16p-l8p, the episporium 1.5p thick. CHEMIsmY.-Cortex K+ yellow, medulla K-, C- or C+ rose, KC+ red, P- (atranorin, alector- onic acid, and a-collatolic acid with gyrophoric acid as an accessory substance). DIsmIBuTIoN.-Mexico, Panama, Colombia, Venezuela, Ecuador, Peru, and Bolivia. HABITAT.-on small trees, humus, soil along road banks, and rarely rocks in high-elevation, humid forest and in the paramo at 2600-3900 m elevation. REMARKS.-This is one of the lichen spectacu- lars of the paramo region along with H. carac- censis. It grows loosely on humus and festoons small shrubs in the paramo. Individual lobes may be up to 20 cm long. At lower elevations it forms extensive suberect colonies on exposed tree trunks. The two chemical populations are morphologi- cally identical and appear to be sympatric, at least in Venezuela. I am regarding these as one species for the present. The holotype of Parmelia expor- recta contains gyrophoric acid in addition to alec- toronic acid and a-collatolic acid; the holotype of P. gigas lacks gyrophoric acid. SPECIMENS EXAMINED.-Mexico: Oaxaca, Hale 20723 (DUKE, US), Skorepa 5559 (US); Vera- cruz, Liebmann 116 (UPS). Colombia: Boyaca, Cuatrecasas 1348 (US); Cundinamarca, Lindig 2573 (P), Ariate (UPS). Venezuela: Merida, Hale 42360, 42367, 42376, 42673, 43158, 43187, 43231, Linden 1088 (BM), 1089 (P), Nilsson 1508 (UPS), Oberwinkler and Poelt 7519 (M, US). Peru: Cuzco, Gomez 631 (US), Holligan 38 (BM). Bolivia: Bang 1853 (H, NY), Spruce 174 (BM). Argentina: Patagonia, Dusen 122 (S). Additional localitie., for Mexico, Panama, Venezuela, Co- lombia, and Ecuador are listed in Hale and Kurokawa (1964: 177, 178). 29. Hypotrachyna gondylophora, new combination FIGURE lle Purrnelia gondylophoru Hale, 1967:420. [Type collection: Clingmans Dome, Swain County, North Carolina, Hale 33570 (US, holotype; DUKE, FH, UPS, isotypes).] Thallus loosely attached, whitish mineral gray, 8-15 cm broad; upper surface plane, smooth, some- times faintly white-maculate, sorediate toward the tips, the soralia capitate; lower surface densely rhizinate, the rhizines densely dichotomously branched. Apothecia not seen. CHEMIsmY.-Cortex K+ yellow, medulla K-, C -, P + orange-red (atranorin, fumarproto- cetraric acid, and succinprotocetraric acid along with rhodophyscin (under soralia) if pigment present). DISTRIBUTION.-Southeastern United States, Mex- ico, Costa Rica, Dominican Republic, Venezuela, Bolivia, and Africa. HABITAT.-on trees (hardwoods and conifers), rarely on rocks, in cloud forests at 1000-2200 m elevation. REMARKS.-The most unusual feature of this species is the chemistry; no other Hypotrachyna is known to contain fumarprotocetraric acid (Hale, 1967). Otherwise it is externally similar to H. densirhizinata or robust forms of H. laevigata. SPECIMENS EXAMINED.-Mexico: Chiapas, Hale 19920; Oaxaca, Nakanishi 349 (US); Veracruz, Hale 20867, 20879. Costa Rica: San Jose, Stone (DUKE). Dominican Republic: La Vega, Zmshaug 23470 (MSC, US). Venezuela: Merida, Oberuinkler and Poelt 7530 (M, US). Bolivia: Inquisive, Overbeck 5455 (US). 30. Hypotrachyna gracilescens, new combination FIGURE llf Purrnelia gracilescens Vainio, 1890:53. [Type collection: Ca- rassa, Minas Gerais, Brazil, Vuinio 1517 (TUR, lectotype).] Thallus loosely adnate on rock, buff mineral gray, 3-5 cm in diameter; lobes sublinear, dichot- omously branched, more or less divaricate- imbricate, 1-2 mm wide; upper surface plane, shiny, without maculae, soredia, or isidia; lower surface moderately rhizinate, the rhizines sparsely dichotomously branched. Apothecia adnate, 1-2 mm in diameter; spores 7p X llp. (atranorin and one or two unidentified substances). CHEMISTRY.-cOrteX Kf yellow, K--, c-, P- DIsTRIBuTIoN.-Southeastern Brazil. HABITAT.-on open rock outcrops at 1400-1500 m elevation. KUMBER 25 41 REMARKS.-This very rare saxicolous species is externally close to H. liuida but has an unusual chemistry. Without knowing the molecular struc- ture of the ?gracilescens? unknown, however, we cannot speculate on its relation to species contain- ing lividic acid. Vainio (1890) differentiated the species from Parmelia chilena, P. bahiana, and P. subsinuosa (all KC+ red) by the negative KC reaction . SPECIMENS EXAMINED.-BraZi1: Minas Gerais, Damazio 934 (G), Vainio 1218, 1532 (BM, TUR). 31. Hypotrachyna imbricatula, new combination FIGURE 12a Parmelia irnbricatula Zahlbruckner, 1909: 168. [Type collec- tion: Itapecirica, Sao Paulo, Brazil, Schiflner (W, lecto- type; MICH, isotype).] Parmelia lobulifera var. Zuteoreagens Degelius, 1941:61. [Type collection: Myrtle Point, Mt. LeConte, Tennessee, U.S.A., Degelius (DEGEL, lectotype).] For additional synonymy see Hale, 1971a:16. Thallus adnate, 5-10 cm in diameter, whitish mineral gray but often turning pale tannish in the herbarium; lobes sublinear, often crowded, 2-4 mm wide; upper surface plane, shiny, usually strongly white-maculate, moderately isidiate, the isidia thin, cylindrical, simple to branched, some- times darkening at the tips; lower surface densely rhizinate, the rhizines densely dichotomously branched. Apothecia infrequently developed, 2- 10 mm in diameter; spores 7p-lop X llp-16p. CHEiwsmY.-Cortex K+ yellow, medulla K-, C+ orange, P- (atranorin, barbatic acid, 4-0- demethylbarbatic acid, obtusatic acid, and norob- tusatic acid). DIsTR1BuTIoN.-Southeastern United States south- ward to Chile. HABITAT.-on trees (oaks, pine, juniper, Arau- caria, Arbutus, coconut palm, citrus), stumps, rocks, and (rarely) soil, in primary and secondary forests at 200-2600 m elevation. REMARKS.-Hypotrachyna imbricatula is the com- monest and most widely distributed species in the genus. It also has wide ecological amplitude. The isidia usually are well developed and may rarely become in part lobulate or procumbent. In the herbarium it often turns light brownish with age. The presumptive progenitor is H. physcioides and the sorediate morph H. laevigata. A very similar species, H. bogotensis, is separated by the C+ rose (rather than orange) reaction caused by evernic and lecanoric acids. The very common H. costa- ricensis has broader lobes and a uniformly dense rhizine mat below, as well as completely negative color tests. Chiapas, Hale 20021, 20197, 20365, 20455, 21202; Michoacan, Arsene 8032, 8245 (US), Darrow 5155 (Darrow herbarium), Kiener 18456 (DUKE); Oaxaca, Hale 20652; Sinaloa, Breedlove 16758d (US); Veracruz, Hale 21059, 21 146. Guatemala: Baja Verapaz, Hale 38322, 38341; Jalapa, Stand- ley 76825 (F, US); Totonicapan, Standley 84542 (F, US); Zacapa, Steyermark (US). Honduras: Morazan, Standley and Molina 4320 (F). Costa Rica: Cartago, Culberson 13219 (DUKE); San Jose, Culberson 12403, 12628 (DUKE), Standley 43033, Standley and Valerio 43174, 43479 (US). Panama: Chiriqui, Almeda 1025A (DUKE), Hale 38720, 38820, 38884, 38911, Maxon 4989 (US), Scholander (US). Cuba: Oriente, Imshaug 25030 (MSC); Pinar del Rio, Imshaug 25221, 25225 (MSC, US). Jamaica: Imshaug 12950, 13382, 13417 (MSC, US), 15278, 15531 (MSC), Orcutt 5626 (US), Plitt (US). Haiti: Ouest, Imshaug 22958, 22980, 23069 (MSC, US), Wetmore 3001, 3044, 3121, 3151 (MSC, US); Sud, Imshaug 23235 (MSC, US). Dominican Republic: La Vega, Al- lard 17695, 17709, 17715 (US), Imshaug 23628, 23683 (MSC, US), Wetmore 3428, 3513 (MSC, US). Guadeloupe: Hale 38280. Dominica: Hale 35345. Trinidad: Hale 38212, 38294. Colombia: Santander, Cuatrecasas 12470 (US), Charetier 211 (US), Killip and Smith 18942 (US). Venezu- ela: Bolivar, Steyermark and Dunsterville 92981 (US); Federal, Jahn 249 (US); Merida, Hale 42185, 42761, 43302. Peru: Cuzco, Herrera 3461a (US), Worthley (US). Guyana: Lillie 47 (BM). Brazil: Bahia, Willey (US); Minas Gerais, Mosen 2319 (UPS), Vainio 1085 (TUR); Rio de Janeiro, Eiten 6512 (US); Santa Catarina, Reitz and Klein 11991, 15501 (US). Chile: Valdivia, Santesson 3785 (S, US). REPRESENTATIVE SPECIMENS EXAMINED.-MexicO: 32. Hypotrachyna immaculata, new combination FIGURE 12b Parmelia immaculata Kurokawa in Hale and Kurokawa, 1964: 42 SiMITHSONIAN COKTRIBUTIOXS TO BOTANY 178. [Type collection: Zoutpansberg, Union of South Africa, Almborn 6551 (LD, holotype; US, isotype).] Thallus loosely adnate to adnate, very fragile, whitish mineral gray, 5-7 cm broad; lobes sub- linear, contiguous, 1.5-3 mm wide; upper surface plane, shiny, sorediate, the soralia mostly sub- terminal, large and capitate, often pigmented ochre below, 1-2 mm in diameter; lower surface densely rhizinate, the rhizines densely dichotom- ously branched. Apothecia rare, adnate, 2-3 mm in diameter; spores 6p X 11p. CHEMISTRY.-cOrteX K f yellow, medulla K- or K+ reddish, P- or P+ dull reddish, C (atranorin, colensoinic acid, 4-0-demethylphyso- dic acid, lividic acid, and associated unknown substances). D1sTRIBuTIoN.-Venezuela, Brazil, and South Africa. HABITAT.-& trees, rarely on soil, in open forest and along roadbanks at 2000-3000 m elevation. REMARKS.-The South American material of this poorly known species is variable in size and lobation but compares with the African popula- tion in overall morphology and chemistry. Ad- ditional collections are needed to settle any doubt. In any event, it would be considered as the sorediate morph of H. livida. On reexamining the type of H. immaculata I found that it consists of a mixture of two species, one (apparently new) with lichexanthone in the cortex and one (lecto- type) with atranorin. SPECIMENS EXAMINED.-Venezuela: Merida, Hale 42900, 42969. Brazil: Minas Gerais, Warming 281 (M). 33. Hypotrachyna intercalanda, new combination FIGURE 12c Parmelia intercalanda Vainio, 1890:53. [Type collection: Sitio, Minas Gerais, Brazil, Vainio 899 (TUR, Vainio herbarium number 3014, lectotype; BM, M, UPS, isolectotypes).] Parmelia fragilis Lynge, 1914, p. 123. [Type collection: Santa Anna da Chapada, Mato Grosso, Brazil, Malrne 2365* (S, lectotype).] Thallus adnate to closely adnate on bark, tan- nish mineral gray, 3-6 cm broad; lobes sublinear, crowded to separate, 0.5-2 mm wide; upper sur- face plane to rugulose, continuous, faintly macu- late, without soredia or isidia; medulla white; lower surface moderately rhizinate, the rhizines densely dichotomously branched. Apothecia com- mon, adnate to substipitate, the disc light brown, 2-4 mm in diameter; spores 5p-6p X Sp-lOp. CHEMIsmY.-Cortex K+ yellow, medulla K--, Cf red (atranorin and olivetoric acid). DrsmIBuTIoN.-Southeastern Brazil and Argen- tina. HABITAT.-on trees, rarely on humus over rocks, in open woods at about 500-1200 m elevation. REMARKS.-The C+ red test caused by olivetoric acid was first noted by Vainio (1890) in the original description. Hypotrachyna osteoleuca also produces this rare acid, along with lichexanthone, but the two species have different geographic distribution and ecology. According to C. Culber- son's scheme (Culberson and Hale, 1973), a species such as this one could have originated from progenitors containing microphyllinic acid or perlatolic acid. Further biochemical transfor- mations of H. osteoleuca itself might have pro- duced 4-0-methylphysodic-physodic acid progeni- tors of the H. livida complex. Malme in Lichenes Austroamericani 88 (G, LD, UPS); Minas Gerais, Henschen (UPS, US), Vainio 936 (TUR, UPS) (syntype of P. intercalanda); Parana, Montes 10116 (DUKE, LD, MVM, US); S5o Paulo, Puiggari (G); Santa Catarina, Reitz and Klein 15985 (US). Argentina: Misiones, Montes 10106 (MVM, US), Osorio 4297 (MVM, SPECIMENS EXAMINED.-Brazil: Mato Grosso, US). 34. Hypotrachynu isidiocera, new combination FIGURE 12d Parmelia isidiocera Nylander, 1860:382. [Type collection: Cape Krusenstern, American Arctic, Beechey (BM, lecto- type: H, isolectotype).] Thallus loosely adnate, fragile, greenish min- eral gray, 5-15 cm broad; lobes sublinear, short, often crowded; upper surface shiny, strongly white- maculate, becoming densely isidiate toward the margins, the isidia tall, erect, easily breaking off at the tips to reveal the yellow medulla; medulla sulphur yellow; lower surface densely rhizinate, the rhizines densely dichotomously branched. Apo- thecia rare, substipitate, 2-8 mm in diameter, the rim isidiate, rugose; spores 7y-8~ X 12p-16p. NUMBER 25 FIGURE 12.--Species of Hypotrachyna: a, H. imbricatula (Hale 21059); b, H. immaculata (Hale 42900); c, H. intercalanda (Osorio 4297); d, H. isidiocera (Hale 20219); e, H. koyaensis (Nakanishi 284); f, H. laevigata (Hale 43106). (Scale in mm.) 43 44 SMITHSONIAN CONTRIBUTIONS TO BOTANY CHEMIsmY.-Cortex K 4- yellow, medulla K+, C+ yellowish, P- (atranorin, barbatic acid, obtusatic acid, entothein, undetermined pigments, and other substances, rarely echinocarpic acid and a trace of norobtusatic acid). DISTRIBUTION.-Mexico, Central America, West Indies, Colombia, Venezuela, Brazil, Hawaii. HABITAT.-on trees (oak, pine) in virgin pine forests, cloud forests, and on rocks in open woods at 1000-3000 m elevation. REMARKs.-This robust lichen is easily recog- nized by the bright yellow medulla exposed as the large isidia break away. While obviously a member of the H. physcioides group, it is distant from any known species with respect to isidial formation (largely marginal) and brittleness of the thallus. The type is listed as ?American Arctic? but this is surely a mistake since the species is known only from montane tropical regions. SPECIMENS EXAMINED.-?VfeXiCO: Chiapas, Hale 20219 (DUKE, US). Guatemala: Alta Verapaz, Johnson 171g (US), Panama: Chiriqui, Hale 38690, Scholander (US). Jamaica: Britton 1110, 1121 (FH), Culberson 13502, 13894 (DUKE), Harris 10010 (BM, FH), Imshaug 12993 (MSC, US), 14937 (MSC). Dominican Republic: La Vega, Allard 16555G (US). Colombia: Tolima, Quimby (US). Venezuela: Federal, Santesson 6677B (S); Merida, Hale 43289. Brazil: Minas Gerais, Damazio in Zahlbruckner, Lichenes Rari- ores 93 (BM, P, UPS); Rio de Janeiro, Gregory and Ainsworth (UPS). 35. Hypotrachyna koyaensis, new combination FIGURE 1% Parmelia koyaensis Asahina, 1953:67. [Type collection: Mt. Koya, Kishu, Japan, Asahina 52101 (TNS, lectotype).] Thallus adnate, membranaceous, ashy white, up to 10 cm broad; lobes irregular, subrotund api- cally, with crenate margins, 4-10 mm wide; upper surface plane, continuous, moderately isidiate, the isidia simple to rarely branching, cylindrical; lower surface black but with a narrow brown zone at the margins, moderately rhizinate, the rhizines moderately dichotomously branched. Apo- thecia rare, stipitate, 1-3 mm in diameter; spores 8p X 16~. CHEMIsmY.-Cortex K + yellow, medulla K-, C--, P+ orange-red (atranorin, protocetraric acid, and an unidentified fatty acid). DISTRIBUTION.-Japan, Southeast Asia, Mexico. HABITAT.-On trees (Pinus) at 2100 m elevation. REMARKS.-ThiS species has the subirregular rotund lobation so typical of Asian species. The isidia are sparce and the rhizines sparcely to mod- erately branched. Its occurrence in the New World is difficult to explain, nishi 284 (US). SPECIMENS EXAMINED.-h?feXiCO: Oaxaca, NUkU- 36. Hypotrachyna laevigata, new combination FIGURE 12f Lichen laeuigatus Smith, 1808: 1852. [Type collection: Angle- sey, Caernarvonshire, England, Dauies (LINN, lectotype).] Parmelia laeviga ta (Smith) Acharius, 18 14: 2 12. Parmelia boliviana var. cephalota Zahlbruckner, 1926:16. [Type collection: Loma Frei Jorge, Coquimbo, Chile, Skottsberg 448 (W, lectotype; S, isolectotype),] Thallus adnate to loosely adnate, greenish to whitish mineral gray, 3-10 cm broad; lobes linear, 1.5-6.0 mm wide; upper surface plane, shiny, usu- ally strongly white-maculate, pustulate-sorediate toward the lobe tips, soralia capitate; lower surface densely rhizinate, rhizines densely branched. Apo- thecia rarely developed, to 5 mm in diameter, adnate; spores not seen. CHEiwsmY.-Cortex K+ yellow, medulla K-, C+ orange, P- (atranorin, barbatic acid, 4-0- demethylbarbatic acid, obtusatic acid, and nor- obtusatic acid). DIsTRIBUTION.-~nited States southward to Chile, Europe. HABITAT.-oII trees (hardwoods, conifers, Drimys, Nothofagus) and rocks in open cloud or rain forests at 600-3700 m elevation. REMARKS.-This was one of the first Hypo- trachyna species to be described on the basis of European material, but the name has been applied in floristic lists to many different narrow- lobed species. The soredia are well developed and the maculae often very distinct. The pre- sumptive progenitor is H. physcioides, and the parallel isidiate morph is H. imbricatula. Accord- ing to Culberson and Hale (1973), the obtusatic acid originated from a cross between a progenitor NUMBER 25 containing evernic acid (H. pulvinata) and a pro- genitor containing barbatic acid (extinct or now represented by H. massartii (Hue) Hale in Asia). The species evolved parallel with H. rockii, the sorediate morph of H. pulvinata, and the two are very similar externally. SPECIMENS EXAMINED.-Mexico: Chiapas, Hale 19885; Oaxaca, Yoshimura 650533 (NISH). Costa Rica: Cartago, Culberson 12419, 13221 (DUKE); San Jose, Ream (WIS). Jamaica: Plitt (US). Dominican Republic: La Vega, Imshaug 23386, 23561 (MSC, US). Guadeloupe: Culberson 14503 (DUKE). Dominica: Elliott 1321, 1556, 1559 (TUR), Hale (see data in Hale, 1971a:17). CO- lombia: Cundinamarca, Killip 11900 (US), Nee and Mori 4219 (US), Weir 53 (BM); Santander, Chawtier 189 (US); Tolima, King 506 (US). Venezuela: hlerida, Hale 422 14, 42647, 43 106, 43206. Guyana: McConnell and Quelch 503 (BhI). Brazil: Rio de Janeiro, Eden 7093 (US). Chile: Aysen, Santesson 6948 (S, US); Coquimbo, Munoz 1257 (S), Sparre 2933 (S); Juan Fernan- dez, Skottsberg 285 (UPS); Valdivia, Santesson 3273, 3277, 3464, 3786, 3698, 7571 (S). 37. Hypotrachyna lineariloba, new combination FIGURE 13a Parmelia lineariloba Kurokawa in Hale and Kurokawa, 1964: 180. [Type collection: Alto de la Bandera, La Vega, Domini- can Republic, Wetmore 3475 (MSC, holotype; US, isotype).] Thallus loosely attached to bark, rather frag- ile, ashy white, 3-6 cm broad; lobes elongate, linear, dichotomously branched, subdivaricate, 0.5-2 mm wide; upper surface plane, shiny, con- tinuous, lacking soredia and isidia; lower surface moderately rhizinate, the rhizines about 1 mm long, sparsely dichotomously branched. Apothecia not seen. C-, KC+ rose, P- acid). CHEMISTRY.-cOrteX K+ yellow, medulla K--, (atranorin and alectoronic ~IsTRIBUTIox.-Dominican Republic. HABITAT.-on trunks of Pinus in open forests on ridges a[ 2400-2800 m elevation. RERIARKS.-NO new collections of this species have been made and it may indeed be endemic to the pine forests of the West Indies. It resembles H. gigaT, a larger species which always contains a-collatolic acid along with alectoronic acid. SPECIMENS ExhhZINED.-Additional records for the Dominican Republic are listed in Hale and Kurokawa (1964: 181). 38. Hypotrachyna liuida, new combination FIGURE 13b Parmelia livida Taylor, 1847: 171. [Type collection: New Or- leans, Louisiana, Hooker Herbarium (FH-TAYL, lectotype; BM, H, isolectotypes).] Parmelia osseoalbida Lynge, 1914: 133. [Type collection: Porto Alegre, Rio Grande do Sul, Brazil, Malne 595 (S, lectotype; UC, UPS, isolectotypes).] Thallus closely adnate, rather coriaceous (on bark) to fragile (on rock), whitish mineral gray, 4-9 cm broad; lobes sublinear, dichotomously branched, contiguous, 1-2 nim wide; upper sur- face shiny, plane to rugulose, lacking soredia and isidia; lower surface moderately rhizinate, the rhizines sparsely dichotomously branched. Apo- thecia numerous, adnate to substipitate, 1-5 mm in diameter; spores 5p X lop-12p. CHEMIsmY.-Cortex K + yellow, medulla K-, C-, KC+ rose, P- (atranorin, colensoinic acid, norcolensoinic acid, 4-0-demethylphysodic acid, physodic acid, lividic acid, and associated unknowns). DISTRIBUTION.-SOUtheaStern United States, Ba- hamas, Venezuela, southeastern Brazil southward to Argentina). HABITAT.-On tree trunks and branches or (more rarely) on rocks, in open forests and gal- lery forests at 500-2800 m elevation (to near sea level in the United States). R~~~~~s.-Hypotrachyna livida is characterized by the adnate thallus, lack of diaspores, and com- plex chemistry. While we still do not know the structure of lividic acid, the presence of at least colensoinic and physodic acids would place the species on a high evolutionary scale in the genus (Culberson and Hale, 1973). Some specimens (in Brazil) appear to lack lividic acid and contain colensoinic acid and related compounds. These may represent a newly evolving species but care- ful studies with high-pressure liquid chroma- tography are needed to settle the chemistry. The species is by far most common in the de- ciduous forests of southeastern United States (W. Culberson, 1961) and then reappears in a climatically similar zone in southern Brazil to 46 ShlITHSONIAN CONTRIBUTIONS TO BOTAKY FIGURE 13.--Species of Hypotrachyna: a, H. lineariloba (Wetmore 3425); b, H. livida (Reitz and Klein 15150); c, H. lopezii (Hale 42463); d, H. mnltnei (ibfalme 2750); e, H. microblasta (Hale 42031); f, H. minima (Malrne 2747B in S). (Scale in mm.) NUMBER 25 47 Argentina. The rock form described by Lynge as Paymelia osseoalbida is somewhat more fragile than corticolous forms. Hypotrachyna livida is the progenitor of several vegetative morphs: H. immaculata (sorediate), H. dactylifera (isidiate-pustulate), and H. pustulata (pustulate). This whole complex has evolved parallel with H. novella (a species containing lichexanthone), as discussed below. tesson 6682 (S, US); Merida, Hale 42833; Valencia, Magdefrau 389 (M). Brazil: Minas Gerais, Vainio 1400 (BM, TUR, as P. gracilis); Rio de Janeiro, Eiten and Eiten 6598, 7396, 7403, 7468 (US), Gla- ziou (G); Rio Grande do Sul, iMaZme 1330B (S); Sao Paulo, Dhen 11 (S); Santa Catarina, Reitz and Klein 13213, 15185, 15809, 16002 (US). Uruguay: Lavalleja, Hosseus 45 (H), Osorio 6480 (MVM, US); Maldonado, Hosseus 93457a (H), Osorio 5040 (MVM); Rocha, Hosseus 43, 47 (H, M); Treinta y Tres, Osorio 6000 (MVM, US). Argentina: With- out collector (M). SPECIMENS EXAMINED.-VeneZuela: Federal, Sari- 39. Hypotrachyna lopezii, new species FIGURE 13c Thallus laxe adnatus, viridi-flavicans, lobis lineari-elongatis, divaricatis, nigromarginatis, 1.5- 2 mm latis; cortex superior 14p-16p crassus, stratum gonidiale 24y-18p crassum, medulla alba, 130y crassa, cortex inferior 18p-20y crassus; sub- tus niger, sparse rhizinosus vel papillatus, rhizinis longis. Apothecia rara, substipitata, sporis simpli- cibus, 7p X 14p. Thallus loosely attached, greenish yellow, to 12 cm broad; lobes linear-elongate, sparsely dichot- omously branched, divaricate and almost appear- ing subfruticose, black rimmed, 1.5-2 mm wide; upper surface plane to concave, shiny, lacking isidia and soredia; lower surface in part bare and papillate, otherwise sparsely rhizinate, the rhizines long, sparsely dichotomously branched. Apothecia rare, substipitate, 8-11 mm in diameter, the amphithecium rugose; spores 7p X 14p. CHEmsmy.-Cortex K-, medulla K+ yellow turning red, C-, P+ orange (usnic acid, nor- stictic acid, stictic acid, and constictic acid). TYPE CoLLECTIoN.-La Aguada, Merida, Ven- ezuela, Hale 43208, 6 February 1974 (US, holotype). DIsTRiBuTIoN.-Venezuela. HAnrTAT.-On base of shrubs and twigs in open rocky paramo at 2800-3500 m elevation. REMARKs.-This paramo species might be con- fused initially with H. caraccensis, but the lobes are narrower, usually black rimmed, and the rhizines very sparse. The chemistry is unusual, stictic acid being rare in Hypotrachyna. The species is named in honor of Prof. Manuel Lopez Figueiras. SPECIMENS ExAhmED.-Venezuela: Merida, Hale 42463, 43 142, 43 195. 40. Hypotrachyna malmei, new combination FIGURE 13d Parmelia rnalmei Lynge, 1914:116. [Type collection: Bocca da Serra, Mato Grosso, Brazil, Malme 2750 (S, lectotype; W, isolectotype).] Thallus loosely adnate on rocks, very fragile, 3-5 cm in diameter, ashy white; lobes sublinear, short, crowded, 0.5-1.5 mm wide; upper surface plane, dull, pustulate, the pustules entire or burst- ing open apically, without forming soredia; me- dulla in part yellow-ochre below; lower surface moderately rhizinate, the rhizines sparsely dichot- omously branched. Apothecia adnate, the amphi- thecium pustulate, 0.5-2 mm in diameter; spores 6p-8~ X 81.1-12~. CHEMISTRY.-cOrteX K--, medulla K- or K+ reddish, P + orange (lichexanthone, protocetraric acid, alectoronic acid, and rhodophyscin). DIsTRIBuTIoN.-~oUtheastern Brazil. HABITAT.-on open sandstone outcrops. RE;CZARKs.-This rare saxicolous species is char- acterized by the pustules and unusual combina- tion of chemicals. It will undoubtedly be found again in southeastern Brazil with more intensive collecting. I had originally synonymized it with H. minima (Hale, 1960) and compared it with H. foymosana. It is, however, morphologically and chemically distinct from both of these. 41. Hypotrachyna microblasta, new combination FIGURE 13e Parinelia rnicroblasta Vainio, 1890:57. [Type collection: Ca- rassa, Minas Gerais, Brazil, Vainio 1214 (TUR, lectotype; BM, M, P, UPS, isolectotypes).] 48 SMITHSONIAN CONTRIBUTIONS TO BOTANY Parmelia revoluta f. isidiosa Muller Argau, 1891:378. Parmelia propagulifera Vainio, 1899: (123). Parmelia jarnaicensis Vainio, 1915:23. Parmelia pseudorevoluta Gyelnik, 1931:289. Parmelia neopropagulifera Gyelnik, 1938a:39. Parmelia endorubra f. imbricatiformis Gyelnik, 1938b3277. Parmelia norstictica Hale, 1959:128. [For full synonymy, see Hale, 1971a:18.] Thallus adnate to loosely attached, 5-15 cm in diameter, pale yellowish green, turning reddish in the herbarium if improperly curated; lobes sub- linear, 1-4 mm wide; upper surface plane, shiny, without maculae, moderately to densely isidiate, the isidia cylindrical, tall, simple or coralloid- branched; lower surface densely rhizinate, the rhizines densely dichotomously branched. Apo- thecia 2-6 mm in diameter, adnate, the amphi- thecium isidiate; spores 6p-8p X 10p-13p. CHEMIsmY.-Cortex K-, medulla K+ red, P+ orange (atranorin (traces), usnic acid, norstictic acid, salazinic acid, and galbinic acid). DIsmIBuTIoN.-Mexico, Central America, West Indies, Andean region from Venezuela to Peru, southeastern Brazil, Hawaii, and Southeast Asia. HABITAT.-On trees (hardwoods and conifers), rocks, and soil in open secondary forests, pastures, and road banks at 900-3100 m elevation. RE~fARKs.-Hypotrachyna rnicroblasta is com- mon in a wide range of habitats throughout trop- ical America. The extensive synonymy is difficult to understand in view of the general uniformity of the species. The thallus frequently discolors red when improperly pressed and dried. The chemistry relates it to H. enderythraea, the pre- sumptive nonisidiate sexual morph or a popula- tion derived from it. ico: Chiapas, Hale 20038, 20210, 20221, 20353, 20385, 20553; Michoacan, Arsene (US). Guate- mala: Quetzaltenango, Standley 86085 (US). Costa Rica: Alajuela, Standley and Torres 47471 (US); Cartago, Standley and Valerio 46807 (US). Pan- ama: Chiriqui, Hale 38799, 38842, 38912, 38925, Scholander (US). Cuba: Oriente, Znzshaug 25141, 25179 (MSC), Morton 9595, 9596 (US). Jamaica: Curnrnings (FH), Hart 54 (NY), Zmshaug 12943, 13057, 13308, 13313 (MSC), 13327 (LD, MSC), 13857, 13923, 14815, 14898, 15452 (MSC, US), Maxon 8634 (US), Plitt (US). Haiti: Ouest, Imshaug 23088, Wetrnore 2993 (MSC, US). Dom- inican Republic: La Vega, Allard 17726A (US), REPRESENTATIVE SPECIMENS EXAMINED.-MeX- Zrnshaug 23389 (MSC), 23573 (MSC, US). Guade- loupe: Duss 676 (NY). Dominica: Hale 35450. Colombia: Cundinamarca, Cuatrecasas 231, 27068a (US), Weir 78 (BM); Santander, Killip and Smith 18942a (US). Venezuela: Bolivar, Steyer- mark 59817 (US); Federal, Santesson 6683 (S) Merida, Hale 42146, 42192, 42773, 43028, Ober- zuinkler and Poelt (M, US), Vareschi 883 (US). Peru: Cuzco, Marin 21916 (F). Ecuador: Imba- bura, Wiggins 38941 (COLO). Brazil: Sowerby (lectotype of P. reuoluta f. isidiosa Miiller Argau) (G); Minas Gerais, Vainio 1160 (BM, M, TUR, UPS). 42. Hypotrachyna minima, new combination FIGURE 13f Parmelia minima Lynge, 1914: 139. [Type collection: Bocca da Serra, Mato Grosso, Brazil, Malme 2747 bis (S, lectotype).] Thallus adnate on rocks, fragile and breaking apart, creamy whitish gray and darkening toward the center, 6-8 cm broad; lobes sublinear, crowded and imbricate, 0.5-1.5 mm wide; upper surface plane to rugulose, continuous to cracked with age, pycnidia and papillate apothecial initials abun- dant; medulla orange-red in the lower part; lower surface sparsely to moderately rhizinate, the rhiz- ines sparsely dichotomously branched. Apothecia common, adnate, 1-2.5 mm in diameter, the amphithecium crenate and bullate; spores 7y X lop. CHEMISTRY.-COrteX K-, medulla K-, C-, KC + reddish, P + orange (lichexanthone and protocetraric acid). DISTRIBUTION.-SOUtheaStern Brazil. HABITAT.-on open sandstone rocks at about 1000 m elevation. REMARKS.-ThiS saxicolous species is as rare as its close relative H. rnalrnei in Brazil. The pus- tules, which I originally described (Hale, 1960), are entire and confined to the amphithecium. This difference, in addition to chemistry (lack of alectoronic acid), clearly separates H. minima from H. rnalrnei. 43. Hypotrachyna monilifera, new combination FIGURE 14a Parmelia monilifera Kurokawa in Hale and Kurokawa, 1964: NUhIBER 25 49 182. [Type collection: La Aguada, Merida, Venezuela, Magdefrau 651 (M, holotype; US, isotype).] Thallus loosely adnate on bark, 10-15 cm broad, whitish mineral gray; lobes linear-divaricate, 2-6 mm wide; upper surface plane, continuous; lower surface densely rhizinate, the rhizines moderately branched, conspicuously moniliform-segmented, projecting as a mat beyond the lobe margins. Apothecia not present. CHEMIsmY.-Cortex K+ yellow, medulla K-, C+ orange, P- (atranorin, barbatic acid, 4-0- demethylbarbatic acid, possibly traces of obtusatic and norobtusatic acids). D~smIBuTIoN.-Venezuela. HABITAT.-On bark of small trees in lower paramo region at about 2800 m elevation. REMARKS.-while superficially close to H. gigas, this species has a stiffer thallus and peculiar moni- liform rhizines easily detected with a hand lens. I was fortunate in being able to re-collect it at the type locality where, apparently, it is very rare in comparison with H. gigas and corticolous. The chemistry relates it to the H. physcioides group, but the elongate lobes and the rhizines put it in an isolated position in the genus. SPECIMENS EXAMINED.-Venezuela: Merida, Hale 43085. 44. Hypotrachyna neodissecta, new combination FIGURE 14b Parmelia neodissecta Hale, 1971c:94. [Type collection: 6 km SE of Zouepo, N?Zerekore, Guinea, Santesson 10597d (UPS, holotype; US, isotype).] Thallus loosely attached, rather fragile, pale greenish ashy gray, 3-7 cm broad; lobes sublinear to subirregular, in part subascending, becoming subcaniculate, 1-3 mm wide; upper surface shiny, continuous, becoming densely isidiate, isidia cy- lindrical, mostly simple; lower surface densely rhizinate, the rhizines moderately dichotomously branched. Apothecia adnate, the amphithecium isidiate, 2-3 mm in diameter; spores not found (in African material, 6p-8~ X 12p-14p). CHEMISTRY.-cOrteX K+ yellow, medulla K-, P-, C+ red (atranorin and gyrophoric acid). DISTRIBUTION.-VeneZUela, Africa, and India. HABITAT.-on trees in secondary forest at 2000- 2200 m elevation. REMARKS.-HyPotrachyna neodissecta was first discovered in Guinea and subsequently collected in other African localities and in India. I would not have expected to find it in the New World. It is distinguished from Purmelinu dissecta (Ny- lander) Hale, for which it would almost certainly be mistaken, by the branched rather than simple rhizines, lack of cilia, and overall canaliculate- revolute pattern of lobation. SPECIMENS EXAMlNED.-VeneZuela: Merida, Hale 42973. 45. Hypotrachyna nouella, new combination FIGURE 14c Parmelia novella Vainio, 1890:56. [Type collection: Sitio, Minas Gerais, Brazil, Vuinio 1028 (TUR, lectotype; BM, FH, M, isolectotypes).] Parmelia brasilianu var. novella (Vainio) Lynge, 1914: 115. Parfnelia regis Lynge, 1914: 126. [Type collection: S5o Paulo d?el Rey, Minas Gerais, Brazil, Malme 178 (S, lectotype; UPS, isolectotype).] Thallus closely adnate on bark, tannish min- eral gray, 2-4 cm broad; lobes sublinear, crowded, 1-2.5 mm wide; upper surface plane, continuous, darkening toward the center; lower surface black, densely rhizinate, the rhizines densely dichotom- ously branched. Apothecia common, adnate, 2-7 mm in diameter, the amphithecium crenulate; spores 6p X 12p. rose or C - , P - (lichexanthone, colensoinic acid, norcolensoinic acid, 4-O-demethylphysodic acid, lividic acid, physodic acid, and associated un- known substances). DISTRIBUTION.-veneZUela and southeastern Brazil. HABITAT.-on trees in open forests at 1000-2000 m elevation. REMARKs.-HypotraChyna novella is essentially identical to H. liuida except for the replacement of atranorin by lichexanthone. It also has a much more restricted distribution. Chemically identical H. formosana could be considered as the pustulate vegetative morph. Magdejrau 398 (M). Brazil: Mato Grosso, Malme (S, UPS, US); Rio de Janeiro, Eiten 7724a (US); Sao Paulo, Hemmendorf (S). CHEMISTRY.-cOrteX K-, medulla K--, Cf SPECIMENS EXAMINED.-veneZuela: ValenCia, 50 SSIITHSONIAN CONTRIBUTIOSS TO BOTANY FIGURE 14.-Species of Hypotrachyna: a, H. monilifera (Magdefrau 651); b, H. neodissecta (San- tesson 10597d); c, H. novella (Malme I78 in s); d, H. obscurella (Hale 43342); e, H. oostingii (Dey 2403); f, H. osorioi (Osorio 6507). (Scale in mm.) NUhlBER 25 51 46. Hypotrachyna obscurella, new combination FIGURE 14d Parrnelia gracilescens f. obscurella Vainio, 1890:54. [Type collection: Carassa, Minas Gerais, Brazil, Vainio 1180 (TUR, lectotype; BM, isolectotype).] Thallus very closely adnate to rocks, dark min- eral gray, 4-6 cm broad; lobes sublinear, short, crowded, 1-1.5 mm wide; upper surface plane, shiny, cracked transversely with age, pycnidia numerous, isidia and soredia lacking; lower sur- face moderately rhizinate, the rhizines sparsely dichotomously branched. Apothecia common, ad- nate, 1-2.5 mm in diameter; spores 6y X 8y. CHEMISTRY.-cOrteX K-k yellow, medulla K-, C-, KC- or KC+ reddish, Pf orange (atran- orin and protocetraric acid). DISTRIBUTION.-Venezuela and Brazil. HABITAT.-on exposed rocks in open areas and in the paramo at 1500-3500 m elevation. REMARKs.-This dark, small, saxicolous lichen blends in with the background igneous rock sub- stratum and is difficult to collect because of the close adnation. It is very common on boulders in the paramo regions of Venezuela and probably occurs throughout the Andes. There are no close relatives in the genus. SPECIMENS EXAMINED.-T'eneZuela: Merida, Hale 43188A 43322, 43342. Brazil: Minas Gerais, Vainio 1431 (BM, TUR). 47. Hypotrachyna oostingii, new combination FIGURE 14e Parmelia oostingii Dey, 1974:362. [Type collection: Balsam Cone, Yancey County, North Carolina, Dey 1236 (DUKE, holotype; US, isotype).] Thallus loosely attached, firm, whitish mineral gray, 8-12 cm broad; lobes sublinear, dichotom- ously branched, more or less separate, 2-5 mm wide; upper surface plane, shiny, pustulate, the pustules subterminal, becoming coarsely sorediate or remaining as open pits; lower surface moder- ately to densely rhizinate, the rhizines moderately dichotomously branched. Apothecia not seen. C+ red, P- (atranorin, gyrophoric acid, and weakly UVS fluorescent unknown substances). CHEMISTRY.-cOrteX K+ yellow, medulla K-, DIsixIsucrIox.-Southeastern United States and southern Chile. HABITAT.-on trees (conifers, Nothofagus) in open woods at lower elevations in Chile, at 2000 m in southeastern United States. REMARKs.-Externally this species is very Close to H. laevigata and H. thysanota. Specimens from Chile had been misidentified as H. densirhizinata. Chemistry places it closest to H. thysanota, which would react both C+ red (gyrophoric acid) and P+ red (echinocarpic acid). SPECIMENS EXAMINED.-chile: Tierra del Fuego, Santesson 6799 (S, US) . 48. Hypotrachyna osorioi, new combination FIGURE 14f Parrnelia osorioi Hale, 1974133266. [Type collection: Abra de Cotto, Lavalleja, Uruguay, Osorio 6507 (MVM, holotype; US, isotype).] Thallus loosely adnate on rock, fragile, light tannish mineral gray, about 8 cm in diameter; lobes sublinear, about 2 mm wide; upper surface plane, continuous, coarsely isidiate, isidia simple and cylindrical to coralloid-branched, apically breaking down and almost pustulate or sorediate, dense with age; lower surface moderately rhizinate, rhizines moderately dichotomously branched. Apothecia rare, adnate, 3-7 mm in diameter; spores 6y X 12y. C+ red, P- (atranorin and gyrophoric acid). CHEMISTRY .-Cortex K f yellow, medulla K- , DISTRIBUTION.-UrugUay. HABITAT.-on stones in open forest at less than 1000 m elevation. REMARKS.-The isidia of this rather fragile species are well developed and tend to enlarge apically and crumble, unlike any other in the genus. It may be related to H. pluriformis-H. revoluta, although the lobes are more regularly linear. SPECIMENS EXAMINED.-urUgUaY: Rocha, Hos- seus 48 (H); San Jose, Osorio 6178 (MVM). 49. Hypotrachyna osteoleuca, new combination FIGURE 15a Parrnelia osteoleuca Nylander, 1863:439. [Type collection: 52 SMITHSONIAN CONTRIBUTIONS TO BOTANY Bogota, Colombia, Lindig 2528 (H, lectotype; BM, FH, G, H, M, P, IV, isolectotypes).] Thallus closely adnate or becoming loosely attached as thick thallus mats form, ashy white or darkening, 5-10 cm broad; lobes sublinear, often imbricate and crowded, 1-3 mm wide; upper sur- face plane, smooth, continuous, lacking soredia and isidia but usually heavily pycnidiate; medulla white to pale salmon; lower surface sparsely to densely rhizinate, the rhizines densely dichotom- ously branched. Apothecia common, adnate to substipitate, the disc dark brown, 1-4 mm in diam- eter; spores 5p-6p X 7p-9p. CHEMISTRY.-cOrteX K--, medulla K-, cf red, P - (lichexanthone, olivetoric acid, rhodophyscin (in lower medulla), and unidentified substances). DIsRTIBuTIoN.-Colombia and Venezuela. HABITAT.-on exposed rocks in the paramo at 2300-3000 m elevation. REMARKS.-The positive C test was first noted by Vainio (1890), who also noted the superficial resemblance to H. brasilianu. In Venezuela this species occurs intermixed on boulders in the paramo with H. erythrodes, which reacts C-. SPECIMENS ExAhfINED.-colombia: Santander, Killip and Smith 15186 (US). Venezuela: Merida, Hale 42680, 42853, 42864, 43311, 43323, 43336. 50. Hypotrachyna palmarum, new combination FIGURE l5b Parmelia palmarun2. Lynge, 1914: 136. [Tjpe collection: Buriti, Serra da Chapada, Mato Grosso, Brazil, Malnie 2243Ct (S, lectotype).] Parmelia zahlbruckneri Lynge, 1914: 125. [Type collection: Bocca da Serra, Seria da Chapada, Mato Grosso, Brazil, Malme S, lectotype).] Thallus adnate, turning cream or cinnamon- buff in the herbarium, rather fragile, 3-8 cm broad; lobes sublinear, crowded, more or less imbricate, short, 1-3 mm wide; upper surface shiny, plane to rugose and cracked on older lobes, isidia and soredia lacking; medulla white to very pale yellow; lower surface moderately rhizinate, the rhizines rather sparsely dichotomously branched. Apothecia common, adnate, the amphi- thecium crenate, the disc often split, 2-6 mm in diameter; spores 5y-ip X 9p-11p. CHE1\IISTRY.-cOrteX K-k yellow, medulla K-, C-, KCf rose, P- (atranorin, colensoinic acid, the ?palmarum? unknown, and other unidenti- fied substances). DIsTRIB1JTIoN.-Southeastern Brazil. HABITAT.-on trees or sandstone outcrops in open or secondary forests at 600-1500 m elevation. REMARKS.-This rather rare species is closely related to H. livida since they both contain colensoinic acid. Hopefully, the full relationship will be revealed when the molecular structure of the ?palmarum? unknown is determined. The rock form, described by Lynge as Parmelia zahl- bruckneri, is indistinguishable in chemistry and morphology from the corticolous forms. This par- allels the situation with Hypotrachyna livida- Parmelia osseoalbida. SPECIMEKS EXAhiINED.-BraZil: Mat0 Grosso, Malme in Lichenes Austroamericani 90 (H, UPS), Malme 2393**** (S, US), Richards 6355 (BM); Minas Gerais, Warming 279 (M); Santa Catarina, Reitr and Klein 16005 (US); SHo Paulo, Eiten and Eiten 3255, 3256 (US). 51. Hypotrachyna partita, new species FIGURE 15c Thallus laxe adnatus, membraneus, cinereo- albicans, usque ad 15 cm latus, lobis sublinearibus, 4-7 mm latis, superne isidiatis, mox lobulascenti- bus, lobulis 0.3 mm latis, 1 mm longis; cortex superior 16p-18p crassus, stratum gonidiale 22p- 24p crassum, medulla alba, 80p-9Oy crassa, cortex inferior 16p-18p crassus; subtus niger, dense rhizinosus. Apothecia ignota. Thallus loosely adnate, rather membranaceous and fragile, greenish mineral gray, to 15 cm broad; lobes sublinear to subirregular, the tips often broadly rotund, 4-7 mm wide; upper surface plane, shiny, initially isidiate, the isidia becoming strongly lobulate, the lobules dorsiventral, about 0.3 mm wide and 1 mm long, simple to palmately divided, developing both laminally and margin- ally and becoming very dense in older parts; lower surface densely rhizinate, the rhizines densely dichotomously branched. Apothecia not seen. CHEmsmY.-Cortex K+ yellow, medulla K-, Cf red, P- (atranorin and anziaic acid rarely accompanied by perlatolic acid). NUMBER 25 FIGURE 15.--Species of Hypotrachyna: a, H. osteoleuca (Hale 43332); 6, H. palmarum (Eiten 3255); c, H. partita (Hale 43143); d, H. peruviana (Lechler 2727 in G); e, H. physcioides (Hale 43354); f, H. physodalica (Mora 984). (Scale in mm.) 53 54 SMITHSONIAN CONTRIBUTIONS TO BOTANY TYPE CoLLEcTI0Ns.-Between La Aguada and La Montana, Merida, Venezuela, Hale 43143, 6 February 1974 (US, holotype). DIsnIBuTIoN.-Central America and Venezuela. HABITAT.-on trunks of trees in open or cloud forest and on soil at road banks at 2200-3600 m elevation. REMARKs.-This species is common on stabil- ized roadsides where superficially morphologically similar H. bogotensis and H. ensifolia also occur. The lobulate isidia of H. partita are extremely variable and often densely developed. Perlatolic acid was detected in only one specimen (Culber- son 12395) which has very large, almost palmately divided lobules and had been subjected to recent lava ash fall. Nee and Mori 3540 (US). Panama: Chiriqui, Hale 38930. Venezuela: Merida, Hale 42118, 42144, 42852, 43065, 43167, 43191, 43205, 43242. SPECIMENS EXAMINED.-cOSta Rica: Cartago, 52. Hypotrachyna peruviana, new combination FIGURE 15d Parmelia peruviana Nylander, in Crombie, 1876:228. [Type collection: Tatanara, Peru, Lechler 2727 (H, lectotype; BM, G, M, P, S, UPS, isolectotypes).] Thallus loosely adnate on bark, rather fragile, 4-7 cm in diameter; lobes short, sublinear, 2-6 mm wide; upper surface plane to rugulose, faintly maculate, lacking isidia and soredia; medulla pale chalcedony yellow; lower surface densely rhizin- ate, the rhizines 1-2 mm long, dichotomously branched. Apothecia conspicuously stipitate to columnar, 3-6 mm high, 2-5 mm in diameter, the amphithecium rugose; spores 7p-12p X 11p-18p. yellowish, C + orange (atranorin, barbatic acid, obtusatic acid, entothein, and unidentified pigments). CHEMISTRY.-cOrteX K+ yellow, medulla K+ DIsmIBuTIox.-Peru and Brazil. HABITAT.-On trees in open forests at 1000-1500 m elevation. REhrARKs.-The coarsely stalked apothecia are distinctive for this species. The yellow pigment is very pale and probably different from that in H. endochlora. The species is related to H. phys- cioides, as Nylander (1885) suspected, but has diverged considerably from any presumptive pro- genitor in this group because of the unusual apothecia and production of pigments. SPECIMENS EXAMINED.-BraZil: Rio de Janeiro, Gregory and Ainsworth 529, 530 (BM, US); Santa Catarina: Reitz and Klein 16091 (US). 53. Hypotrachyna physcioides, new combination FIGURE 15e Parmelia Physcioides Nylander, 1860:385. [Type collection: Silla de Caracas, Venezuela, Humboldt and Bonpland (P, lectotype; BM, H, isolectotypes).] Parmelia laevigata var. gracilis Muller Argau, 1879: 169. [Type collection: Bogota, Colombia, Andrt 923 bis (G, lectotype; BM, US, isolectotypes),] Parmelia boliviana Nylander, 1885:612. [Type collection: Prov. Larecaja and Caupolican, Bolivia, Weddell (P. lecto- type).] Parmelia gracilis (Muller Argau) Vainio, 1890:55. Parmelia digitata Lynge, 1914:98. [Type collection: Santa Anna da Chapada, Mato Grosso, Brazil, Malme 2545 (S, lectotype; LD, US, W, isolectotypes).] Parmelia confusula Zahlbruckner, 1929: 162. [Type collection: Based on P. gracilis (Muller Argau) Vainio, not P. gracilis Sprengel, 1827:277, or P. gracilis Muller Argau, 1887:317).] Thallus adnate to loosely attached, usually rather coriaceous, 6-15 cm broad, ashy white to tannish mineral gray; lobes sublinear, separate to crowded, 2-6 mm wide; upper surface plane to convex, dull and sometimes white pruinose at the tips to shiny and distinctly white-maculate, rarely becoming sparsely to densely lobulate-laciniate toward the center, isidia and soredia lacking; lower surface densely rhizinate, the rhizines densely dichotomously branched. Apothecia ad- nate to substipitate, 2-10 mm in diameter; spores 6p-9 pX 12p-16p. CHEMISTRY.-cOrteX K + yellow, medulla K-, C- or C+ orange, KC+ orange, P- (atranorin, barbatic acid, 4-0-demethylbarbatic acid, obtusatic acid, and norobtusatic acid). DIsnIBuTIoN.-Mexico, Central America, West Indies, Colombia, Venezuela, Peru, Ecuador, Bo- livia, Brazil, and Southeast Asia. HABITAT.-on trees (conifers and hardwoods) and rocks in open or disturbed forests at 1500- 4100 m elevation. REMARKs.-Hypotrachyna physcioides is com- mon in the northern part of its range but ap- parently lacking in most of Brazil and southward. It is extremely variable, Typical soil- or rock- NUMBER 25 55 inhabiting specimens (as in the type) are almost coriaceous and have dull linear lobes. Some corticolous forms (as in the type of Parmelia boliviana) are thinner and have shorter, sub- linear lobes and a shiny surface with white maculae. Many intermediates can be found. The chemistry, however, is uniform in all cases. A rather densely lobulate population described by Lynge as Parmelia digitata probably is only an extreme form which intergrades with the more normal and abundant nonlobulate population. This species is the presumptive progenitor of various vegetative morphs, H. laevigata (soredi- ate) and H. imbricatula (isidiate), as well as being part of the same constellation that produced H. chlorina, H. endochlora, H. isidiocera, H. peruviana, etc. The possible origin of H. phys- cioides from hybridization between evernic-acid- containing H. pulvinata and a barbatic-acid- containing progenitor similar to H. massartii (Hue) Hale is discussed by Culberson and Hale (1973). SPECIMENS EXAMINED.-Mexico: Puebla, Ar- sene 4253 (US). Guatemala: Alta Verapaz, Stand- ley 91328 (US). Costa Rica: Alajuela, Standley and Torres 47720 (US); Cartago, Culberson 12394 (DUKE, US), Stork L-38 (US); San Jose, Culber- son 13215 (DUKE). Panama: Chiriqui, Martin 2751 (US). Haiti: Ouest, Imshaug 22990, 22991 (MSC), Wetmore 3121 (RISC); Sud, Imshaug 23238 (MSC). Dominican Republic: La Vega, Zmshazig 23459, 23628 (MSC). Colombia: Cun- dinamarca, King 472 (US), Nee and Mori (US), Soderstl-om 1305b, 1305c (US), Weir 27 (BM, US); Santander, Killip and Smith 15644, 18593 (US). Venezuela: Merida, Hale 42171, 42433, 42485, 42804, 43126, 43224, 43326, Hanselmann 168 (DUKE, US), Magdefrau 698 (h4, US), Nash 2013, 2014 (US). 54. Hypotrachyna physodalica, new combination FIGURE 15f Parmelia physodalica Hale, 1971d:436. [Type collection: Guasca, Cundinamarca, Colombia, Perez-Arbelaez 1104 (US, holotype).] Thallus loosely adnate, often scattered, coriace- ous but easily breaking apart, sea-foam yellow, to 8 cm broad; lobes sublinear, elongate, sparsely dichotomously branched, separate, 1-3 mm wide; upper surface plane, shiny, maculae, soredia, and isidia lacking; lower surface moderately to densely rhizinate, the rhizines dichotomously branched. Apothecia not seen. P+ orange-red (usnic acid, physodalic acid, and a trace of protocetraric acid or usnic acid and protocetraric acid only). DIsTRIsuTIoN.-Colombia, Venezuela, and Peru. HABITAT.-on soil and over humus in the par- amo at 3700-4100 m elevation. REMARKS.-This paramo species is easily recog- nized by the deep greenish yellow color and sub- linear divaricate lobes. Both H. caraccensis, a much larger species, and H. Eopezii, which has a nearly bare lower surface, react K+ red or yellow. The type of H. physodalica was reported to con- tain only physodalic acid, but subsequent TLC tests showed traces of protocetraric acid. Speci- mens collected later contain only protocetraric acid. I am provisionally regarding these chemi- cally variable populations as a single species, at least until we know more about the biogenesis and co-occurrence of these closely related acids. SPECIMENS ExAhfINED.-Colombia: Cundina- marca, Mora 984 (US); Santander, Killip and Smith (17527 (US). Venezuela: Merida, Hansel- mann 340 (DUKE, US), Oberwinkler and Poelt (M, US). Peru: San Martin, Holligan L60A CHEMISTRY.-cOrteX K--, medulla K--, c-, (BM) * 55. Hypotrachyna pluriformis, new combination FIGURE 16a Parmelia pluriformis Nylander, 1860:381. [Type collection: Brazil, Weddell (H, lectotype; P, isolectotype).] Parmelia pluriformis var. chlorocarpa Muller Argau, 1881:85. [Type collection: Apiahy, Brazil, Puiggari 125e (G, lecto- Parmelia pluriformis var. multifida Muller Argau, 1881:85. [Type collection: Apiahy, Brazil, Puiggari 1283 (G, lecto- Parmelia everniaeformis Zahlbruckner, 1902:416. [Type col- lection: Petropolis, Brazil, Hohnel 163 (W, lectotype).] Parmelia subpluriformis Zahlbruckner, 1909: 172. [Type c01- lection: Mt. Jaragui, Sao Paulo, Brazil, Schiflner (W, lecto- type).] type).] type).] Thallus adnate or loosely attached to bark, greenish mineral gray, 5-8 cm broad; lobes sub- linear to subirregular, imbricate, sometimes be- 56 ShlITHSONIAN CONTRIBUTIONS TO BOTANY coming canaliculate, 1-5 mm wide; upper surface plane to convex, continuous, lacking soredia and isidia; lower surface black in the center but with a broad brown zone along the margins, moderately rhizinate. Apothecia common, substipitate, 2-10 mm in diameter, the amphithecium rugulose, the disc often radially split; spores Sp-11~ X 14p-25p, C+ red, P- (atranorin and gyrophoric acid). DIsTRIBUTION.-~~exico, Brazil, Uruguay, and Argentina. HAnITAT.-On trees in open forest at 1000-2300 m elevation. REhIARKS.-The lobes of this species are unique in being subirregular and in part canaliculate and revolute. The thallus has a coriaceous texture. Spore size is variable but definitely larger than in most Hypotrachynae. Zahlbruckner (1909), in fact, recognized Parnzelia subpluriformis on the basis of large, thick-walled spores, but I would consider this as merely the extreme on a broad scale of spore size. It has the same chemistry as H. reuoluta and H. osoiioi, two vegetative morphs that originated from a sexual morph similar to H. pluriformis. SPECIMENS ExAwNm.-Alexico: Puebla, Hale 19621. Brazil: hIinas Gerais, Burclzell 1105/68, 2252 (BRI), Christina (P), Vainio 794 (Rl, TUR), 1049, 1052 (BRI, TUR); Rio de Janeiro, Glazioir 1832 (UPS); Sao Paulo, Gehit '(US, W), Lind- berg (UPS), Schiflnei (BRI, G). Uruguaj: Cane- lones, Row-Mato 1031 (S); Florida, Imshaug 424064 (RISC). Argentina: RIisiones, ilfontes 10073 (AfVM). CHEILIISTRY.-cOrteX K+ yellow, medulla K- , 56. Hypotrachyna producta, new species FIGURE 16b Thallus laxe adnatus, 6-7 cm diametro, cinereo-albicans, lobis sublinearibus, 2-3 latis, superne sorediatis; cortex superior 13y-15p cras- sus, stratum gonidiale 14u-16~ crassum, medulla alba, 65y-9Op crassa, cortex inferior, 12p crassus; subtus niger, modice rhizinosus. Apothecia ignota. Thallus loosely adnate on bark, whitish min- eral gray, 6-10 cm in diameter; lobes sublinear, becoming elongate and subdivaticate, 2-4 mm wide; upper surface shiny, plane, sorediate, the soredia subterminal in orbicular soralia, the soredi- ate lobes becoming subrevolute; lower surface mod- erately rhizinate, the rhizines moderately to densely dichotomously branched. Apothecia not seen. CHEwsmY.-Cortex K+ yellow, medulla K-, C+ red, P- (atranorin and anziaic acid). TYPE CoLLEcTIoiv.-On Picea rubens, Mt. Le- Conte, Sevier County, Tennessee, Nakanishi 174, 8 hlay 1971 (US, holotype; KOBE, isotype). lombia, and Africa. HABITAT.-On trunks and branches of trees (hardwoods and conifers) in open forest at 2000-3600 m. REhrARKs.-Hypotrachyna producta is morph- ologically similar to other sorediate species such as H. laevigata and H. oostingii. It would have to be identified with a chemical test. The presump- tive progenitor is H. ducalis, which is also rarely collected in tropical America. SPECIMENS ExAxfINED.-Colombia: Cundin- amarca, Killip 341 18 (US); Santander, Killip and Smith 17798 (US). DISTRIBUTIoN.-SOutheastern United States, Co- 57. Hypotrachyna prolongata, new combination FIGURE 16c Pnimelici fiiolongntn Kurokai+a in Hale and Kuiokaira, 1964: 182. [Tjpe collection: Pic Nacaja, Sud, Haiti, Wetmore 3233 (RISC, holotjpe; US, isoqpe).] Thallus adnate to looselj adnate, mineral gray, 5-10 cm in diameter; lobes sublinear, 1.5-4 mm wide; upper surface plane, shiny, faintly macu- late, spaisely lobulate-isidiate along the margins, the "isidia" in part lobulate, sometimes branched, 2-3 mm long; lower surface densely rhizinate. .\pothecia not seen. C+ red, P- (atranorin and an unidentified C+ substance). ~H~h11SlR~.-~01 tex K+ yellow, medulla K--, DISTRIBUTIOA .-Haiti. HABITAT.-OII pine trees at about 2000 m eleLation. RE\IARKS.-ThiS rare species is still known only fiorn the tj pe collection in Haiti. Specimens from other localities so identified on the basis of a Cf 1 etl reaction T$ ere found to contain anziaic acid and to ha\e a different kind of isidial formation. These have been recognized as H. rachista (Hale, 197ld). The C+ substance in H. p?-olongata NUMBER 25 FIGURE lG.-Species of Hypotrachyna: a, H. plurifortnis (Schiflner in IV); b, H. producta (Saka- nishi 174); c. H. prolongata (Wetmore 3233); d, H. proterita (Hale 43365); e, H. protoboliviana (Flenniken 1874); f, H. pseudosinuosa (Hale 35450). (Scale in mm.) 57 58 SMITHSONIAN CONTRIBUTIONS TO BOTANY does not compare with anziaic acid, gyrophoric acid, lecanoric acid, or olivetoric acid and prob- ably represents a new compound. 58. Hypotrachyna protenta, new species FIGURE 16d Thallus laxe adnatus, saxicola, 6-10 cm dia- metro, lobis sublinearibus, imbricatis, 2-4 mm latis; cortex superior 141.1-16p crassus, stratum gonidiale 24p crassum, medulla alba, 8Op-9Op crassa, rare pro parte aurantiaca, cortex inferior 14p-16p crassus; subtus niger, modice rhizinosus. Apothecia numerosa, substipitata, sporis simplici- bus, 6p X 12p. Thallus rather loosely adnate on rocks, the col- onies easily breaking apart, whitish ashy, 6-10 cm broad; lobes sublinear, more or less crowded or imbricate, variable, 2-4 mm wide, sometimes be- coming densely appressed lobulate toward older parts; upper surface shiny, plane to minutely rug- ulose, often heavily pycnidiate, soredia and isidia lacking; lower surface densely rhizinate, the rhizines sparsely to moderately dichotomously branched, sometimes projecting as a narrow mat along the margins. Apothecia numerous, adnate, 4-8 mm in diameter; spores 6p X 12p. CHEMIsmY.-Cortex K+ yellow, medulla K-, C-, KC+ red, P- (atranorin, alectoronic acid, a-collatolic acid, and, if pigmented, rhodophyscin). TYPE CoLLEcTIoiv.-Open forest along stream, on rocks, El Valle, Merida, Venezuela, Hale 43365, February 1974 (US, holotype; DUKE, TNS, UPS, isotypes). DIsTRIBuTIoN.-Centra1 America, Colombia, and Venezuela southward to Peru, and Brazil. HABITAT.-on rocks in open forests and in the paramo at 1700-4000 m elevation. RExfARKs.-Hypotrachyna protenta grows on open boulders or even soil in paramo and sub- paramo regions. Typically the lobes are densely imbricate and black rimmed but under some con- ditions they are almost separate and divaricate. The species resembles H. chicitae but can be dis- tinguished by the negative C reaction and white fluorescence under ultraviolet. It can also be sep- arated from small, high-elevation forms of H. physcioides (C+ orange) by a color test. SPECIMENS ExAMINED.-costa Rica: Stork (DUKE, H, MICH, US, W). Panama: Chiriqui, Hale 38796, Scholander (US). Colombia: Cundina- marca, King 505 (US). Venezuela: Merida, Hale 42130a, 42193, 42915, 42929, 43030, 43278, 43288, 43339. Peru: Cuzco, Staflord 1008 (F, US). Brazil: Santa Catarina, Reitz and Klein 17545 (US); SZo Paulo, Schiflner (BM, W). 59. Hypotrachyna protoboliviana, new combination FIGURE 16e Parmelia protoboliviana Hale, 1974b:267. [Type collection: Volcan Irazu, Cartago, Costa Rica, Flenniken 1874 (US).] Thallus adnate on bark, rather fragile, about 8 cm in diameter, light yellowish green; lobes more or less sublinear, short, 4-5 mm wide; upper sur- face smooth, without isidia or soredia; lower sur- face densely rhizinate, the rhizines densely dichotomously branched. Apothecia poorly de- veloped, about 1 mm in diameter; spores lacking. orange, P- (usnic acid, barbatic acid, 4-0-de- methylbarbatic acid, obtusatic acid, and norob- tusatic acid). CHEMISTRY.-cOrteX K-, medulla K--, c+ DISTRIBUTION.-cOSta Rica. HABITAT.-oII trees in open forest at about 3200 m elevation. REMARKs.-Identical to H. physcioides except for the replacement of atranorin by usnic acid, H. protoboliuiana is known only from Costa Rica in an active volcanic region. As a very recently evolved species it has not given rise to any vegeta- tive morphs comparable to those of H. physcioides. Flenniken 2358 (US). SPECIXIEXS EXAhZINED.-COSta Rica: Cartago, 60. Hypotrachyna pseudosinuosa, new combination FIGURE 16f Parmelia pseudosinuosa Asahina, 1951:329. [Type collection: Shimohirano-mura, Shinano, Japan, Takahashi 99 (TNS, lectotype).] Parmelia anaptychioides Kurokawa in Hale and Kurokawa, 1964: 165. [Type collection: Pic0 Trujillo, Dominican Re- public, Wetmore 3617 (hlSC, holotype; US, isotype).] Thallus closely adnate, light mineral gray, 3-9 cm in diameter; lobes sublinear, short and some- times crowded, 1-4 mm wide; upper surface NUMBER 25 59 plane, continuous, sorediate, the soralia capitate, mainly subterminal; lower surface moderately rhizinate. Apothecia rare, adnate, 2-4 mm in diameter; spores 6p-8y X 8p-lop. CHEiwsmY.-Cortex K+ yellow, medulla K-, P + orange red (atranorin, protocetraric acid). South Africa, Japan, and Southeast Asia. HABnAT.-on tree trunks and small branches (hardwoods, Pinus) in secondary forests at 1800- 2200 m elevation. REMARKS.-The thallus of H. pseudosinuosa is often small and closely adnate, as in the type of Parmelia anaptychioides (Hale, 1971a). It is one of the few species of Hypotrachya first described from Asia and later found to be pantropical. There is no presumptive parent species. SPECIMENS EXAMINED.-Mexico: Oaxaca, Nak- anishi 285 (US). Dominica: Hale 35358, 35488. Chile: Valparaiso, Imshaug 36628, 36642 (MSC). Records from Jamaica and Haiti are listed in Hale and Kurokawa (1964: 166). DISTRIBUTION.-Mexico, West Indies, Peru, 61. Hypotrachyna puluinnta, new combination FIGURE 17a Parrnelia pulvinata Fee, 1824: 123. [Type collection: Tropical America (G, lectotype).] Parmelia subsinuosa Nylander, 1885:613. [Type collection: Tropical America, BonPland (H, Nylander herbarium number 35169, lectotype; P, isolectotype).] P. caraccensis f. pulvinata (Fee) Zahlbruckner, 1929:58. Thallus adnate to loosely attached, whitish min- eral gray, 8-12 cm broad; lobes short and often crowded, sublinear, 2-6 mm wide; upper surface plane, shiny, white-maculate; lower surface densely rhizinate, the rhizines often projecting as a mat beyond the margins, densely dichotomously branched. Apothecia common, adnate, 5-17 mm in diameter, the amphithecium strongly white- maculate; spores 6p-7p X lop-13p. C+ red, KC+ red, P- (atranorin, evernic acid, and lecanoric acid. DISTRIBUTION.-Southwestern United States, Mexico, Central America, Jamaica, Andean re- gion from Venezuela to Bolivia, and southeastern Brazil. HABITAT.-on trees (oak, pine), soil, over CHEMISTRY.-cOrteX K+ yellow, medulla K--, mosses in pine forests, scrubby mist forest, and paramo at 1200-4300 m elevation. recognized by the white maculae, dense rhizine mat, and C+ rose or red color test. It is very com- mon in the dry upland oak-pine forests from southern New Mexico-Arizona southward to Gua- temala but becomes quite rare in South America. The vegetative morphs, H. rockii (sorediate) and H. bogotensis (isidiate), are two of the commonest species in tropical America. It is H. pulvinata or a progenitor similar to it which probably crossed with a barbatic-acid-containing progenitor such as H. massartii (Hue) Hale to give rise to the obtusatic-acid-containing constellation including H. physcioides and its vegetative morphs, H. peruviana, etc. (Culberson and Hale, 1973). SPECIMENS EXAMINED.-Mexico: Chiapas, Hale 20330, 20571; Federal, Kramer 1817 (US); Mex- ico, Hale 19290; Oaxaca, Hale 20672, 20688, 20691, 20700, 20784; Sinaloa, Culberson 13356 (DUKE). Guatemala: Quetzaltenango, Steyermark 34898 (US); Totonicapan, Standley 84507 (US). Costa Rica: San Jose, Standley and Valerio 43750 (US). Panama: Chiriqui, Scholander (US). Jamaica: Imshaug 15495 (MSC), Plitt (US). Colombia: Magdalena, Flenniken 2295 (US); Santander, Killip and Smith 19560 (US); Tolima, Killip and Varela 34678a (US). Venezuela: Merida, Dennis 1940 (BM), Hale 42365, 42374, 42460. Ecuador: Pichincha, Jameson 145 (BM), Ugent (US, WIS). Peru: Bryan 579 (US, WIS). Bolivia: Cocha- bamba, Ugent (US, WIS); Larecaja, Mandon 1753 (BM, S). Brazil: Minas Gerais, Henschen REMARKS.-Hypotrachyna pu~vinata is easily (UPS). 62. Hypotrachyna rachista, new combination FIGURE 17b Parrnelia rachista Hale, 1971d3436. [Type collection: Cling- mans Dome, Swain County, North Carolina, Kurokawa 6755 (US; isotype in TNS).] Parmelia lobulijera var. sanguineoreagens Degelius, 1941:63. [Type collection: Mt. Kephert, North Carolina, Degelius (US, lectotype).] Thallus adnate to loosely adnate, 6-12 cm broad, whitish mineral gray; lobes linear, dichot- omously branched, variable, 1-5 mm wide; upper surface plane, continuous, becoming dactyloid- isidiate at the lobe tips, isidia dorsiventral, often 60 SMITHSONIAX COXTRIBUTIOKS TO BOTANY dissected; lower surface black but sometimes with a narrow white rim below the lobe tips, densely rhizinate, the rhizines densely dichotomously branched. Apothecia not seen. C+ red, P- (atranorin and anziaic acid). DIsTRIsuTIoN.-Southeastern United States, MeX- ico, Panama, Dominican Republic, Colombia, Venezuela, and Peru. HABrrAT.-On trees (hardwoods and conifers), over mosses, and on soil at road banks at 2000- 3500 m elevation. REMARKS.-The isidia of this species are un- usual in branching and position on the thallus (Figure 4d) and one might question whether they should be called isidia or lobules. The distinctive chemistry relates H. rachista to H. ducalis and its sorediate morph H. producta. The species is most commonly collected in the southern Appalachian Mountains in the United States and seems rare in tropical America. SPECIMENS ExAMINED.-Dominican Republic: La Vega, Imshaug 23402 (MSC). Venezuela: Merida, Hale 42610, 42648, 42896, 43139, 43239. Peru: Pasco, Tovar 397 (BM). For additional rec- ords from Mexico, Panama, and Colombia see Hale (1971:437). CHEMISTRY.-cOrteX K+ yellow, medulla K--, 63. Hypotrachyna reducens, new combination FIGURE 17c Parrnelin reducens Nylander, 1963:438. [Type collection: Co- lombia, Lindig 2743 (H, Nylander herbarium number 35100, lectotype).] Thallus adnate to loosely attached, sea-foam yellow, 2-5 cm in diameter; lobes sublinear, sep- arate, 1-2 mm wide; upper surface plane to rugulose, faintly white-maculate, lacking soredia and isidia; lower surface densely rhizinate, the rhizines sometimes projecting out as a mat along the margins. Apothecia common, substipitate, the disc becoming radially split, 1-5 mm in diameter; spores 5p-7p X 9p-llp. CHEmsmY.-Cortex K-, medulla K+ yellow turning red, C-, P+ orange-red (usnic acid, nor- stictic acid, salazinic acid, sometimes a trace of a tranorin). DIsmIBuTIoN.-Panama, Colombia, Peru, Sabah, and New Guinea. HABITAT.-On trunk and twigs of hardwood trees in exposed areas at 2800-3500 m elevation. REhIARKs.-while rarely collected, H. reducens is easily identified since it is the only nonsorediate, nonisidiate yellow species (outside of rare H. pro- tobohiana) on trees. It is always fertile. The typ- ical habitat seems to be exposed shrubs in the paramo. The presumed sorediate morph, H. sin- uosa, may occur in similar habitats. SPECIMENS ExAMINED.-Panama: Chiriqui, Scho- lander (US). Colombia: Humbry-Tracy 313 (BM); Lindig 122 (BM, M), 799 (M, P, syntypes of P. reducens). Peru: Recuay, Gomez 549 (US). 64. Hypotrachyna revoluta, new combination FIGURE 17d Parinelia revoluta Floerke, 1815:ll. [Type collection: Ger- many, Floerke (FH-Tuck, lectotype).] Thallus loosely adnate, rather fragile, green- ish mineral gray, 3-8 cm broad; lobes sublinear to subirregular, often short and more or less as- cending at the tips, subcanaliculate, 1-5 mm wide; upper surface plane, shiny, becoming very coarsely pustulate toward the lobe tips, the pus- tules becoming coarsely sorediate and sometimes eroding centrally leaving bare blackish areas of medulla, heavily sorediate lobes becoming revo- lute; lower surface black but with a brownish zone along the margins, sparsely rhizinate, the rhizines sparsely dichotomously branched. Apothecia very rare, adnate to substipitate, 2-6 mm in diameter; spores 6p-8~ X 10p-12p. CHEmsmY.-Cortex K+ yellow, medulla K-, C+ red, P- (atranorin and gyrophoric acid). DIsTRIBuTIoN.-Eastern United States, West In- dies, Venezuela to Chile in the Andes, Europe, South Africa, Japan, and Indonesia. HABITAT.-On trees (hardwoods and conifers) in open or cloud forest and on rocks at 1000-4500 m elevation (at sea level in Chile). REXIARKs.-This classical European species (Almborn, 1942) has a montane pantropical dis- tribution. The name, however, has been applied incorrectly to many different species in the tropics. There is a wide and often perplexing range of variation in soredial formation, but in general the soredia are lamina1 to subterminal, causing the lobes to become broadly revolute, and may even SUMBER 25 FIGURE 17.-Species of Hy$otmchyria: a, H. pulvit7ata (Hale 20784): 0, H. rochisfa (Hole 43139): c, H. reducens (Liridig 122 in &I); d, H. reuolnfa (Male 43293a); e, H. r/inbdifor?riis (Il.iirtfack 483); f, H. rockii (Rock 7). (Scale in mm.) 61 62 SMITHSONIAN CONTRIBUTIONS TO BOTANY erode away leaving large, bare pustule-like areas (Figure 2a). The lobes also vary widely in width and degree of erectness. Spore size for European material was determined by Hillmann (1936) to be up to 16p long, somewhat larger than Ameri- can material. A presumptive progenitor, H. pluri- formis, has significantly larger spores, up to 251.1 long. Vainio?s determination of his number 1140 as Parmelia revoluta from Brazil is incorrect; it is H. rockii. SPECIMENS EXAMINED.-Dominican Republic: La Vega, Allard 16530A (US). Venezuela: Merida, Hale 42995, 43296, 43306. Bolivia: Brooke 6819 (US), Overbeek 5455 (US). Chile: Chiloe, San- tesson 4029 (S, US), 4031 (S); Valdivia, Raben- horst (M, US), Santesson 2363 (S). 65. Hypotrachyna rhahdiformis, new combination FIGURE 17e Parmelia rhabdijormis Kurokawa in Hale and Kurokawa, 1964: 183. [Type collection: El Boquete, Chiriqui, Panama, Maxon 5128a (US, holotype).] Thallus adnate, light tannish mineral gray, 5-8 cm broad; lobes sublinear, short, imbricate, 2-6 mm wide; upper surface plane, continuous, moderately isidiate, the isidia mostly simple and cylindrical, to 0.5 mm high or more, becoming club-shaped or lobulate; lower surface densely rhizinate, the rhizines densely dichotomously branched. Apothecia adnate, 1-5 mm in diameter, the amphithecium isidiate; spores 6p-8p X 1 5p-2 1 p. CHEMISTRY.-COrteX K + yellow, medulla K + yellow turning red, C-, P+ orange (atranorin, norstictic acid, and in Iltis 3005 and Wurdack 483 a trace of stictic acid). Asia. HABITAT.-oII trees or on soil in cloud forest at 1300-2300 m elevation. REMARKs.-This isidiate species has a unique chemistry. Superficially it resembles H. costari- censis, which has very dense rhizines and reacts K-. The collections from Peru are more robust than the type but agree in other features. SPECIMENS EXAM INED.-Peru: Chachapoyas, Wurdack 483 (US); Urubamba, Zltis 3005 (US, DISTRIBUTION.-Panama and Peru, Southeast WIS). 66. Hypotrachyna rockii, new combination FIGURE 175 Partnelia rockii Zahlbruckner, 1912:379. [Type collection: Kauai, Hawaii, Rock 7 (W, lectotype; US, isotype).] Thallus adnate to loosely attached, 5-12 cm in diameter, whitish mineral gray; lobes sublinear, 1-6 mm wide; upper surface plane, shiny, often distinctly white-maculate, pustulate initially near the lobe tips, usually becoming coarsely sorediate; medulla white except for pale orange spots under the soralia; lower surface black, densely rhizinate. Apothecia rarely developed, adnate, 2-8 mm in diameter; spores 5p-7p X 11p-14p. CHEMISTRY.-COrteX K + yellow, medulla K - , C+ rose, P- (atranorin, evernic acid, and lec- anoric acid). DISTRIBUTION.-SOutheaStern United States to southern Chile, South Africa, Southeast Asia, and Hawaii. HABITAT.-on tree bark (oaks, pine, Ficus, Drimys) and rocks in open forests at 1000-3300 m elevation. REMARKs.-Hypotrachyna rockii is a very com- mon lichen in the highland oak-pine forests of Mexico and Central America as well as in the West Indies. It is very similar externally to H. laevigata in having distinct maculae and subter- minal soredia but can be distinguished from it by a color test with C. The presumptive progenitor is H. pulvinata (Culberson and Hale, 1973). REPRESENTATIVE SPECIMENS EXAMINED.-hfexico: Chiapas, Hale 202 14, 20242, 20290, 20534, 2056 1 ; Michoaah, Arsene 3746 (US), Hale 20850, 20987; Puebla, Hale 21048; Veracruz, Hale 21116, 21176. Guatemala: Alta Verapaz, Standley 69331 (US); Baja Verapaz, Hale 38333; Chimaltenango, Stand- ley 60097 (US); Huehuetenango, Standley (US); Totonicapan, Standley 84522 (F). Costa Rica: San Jose, Standley and Valerio 43898 (US). Pan- ama: Chiriqui, Hale 38768, 38819, 38863, Scho- lander (US). Cuba: Oriente, Zmshaug 25045 (MSC, US). Jamaica: Zmshaug 12954, 13802, 15424, 15541 (MSC, US), Orcutt 5626c (US), Plitt (US). Haiti: Ouest, Zmshaug 22591 (MSC), 22538 (US), Wetmore 2973, (MSC); Sud, Wet- more 3365 (MSC). Dominican Republic: Cordil- lera Central, Zmshaug 23537 (MSC, US), 23639 (MSC); La Vega, Allard 16529a (US). Dominica: NUMBER 25 63 Hale 35586. Martinique: Culberson 14555 (DUKE). Colombia: Santander, Charetier 202 (US). Venezuela: Federal, Vareschi 427c (US); Merida, Hale 42553, 42734, 42974, 43287. Peru: Cuzco, Zltis 3004 (US, WIS), Thomasson (UPS). Chile: Magallanes, Santesson 8010 (S, US); T' ierra del Fuego, Santesson 6793 (S, US); Valdivia, Mahu 1695 (US). Brazil: Minas Gerais, Vainio 1140 (BW 67. Hypotrachyna silvatica, new combination FIGURE 18a Parmelia silvatica Lynge, 1914: 118. [Type collection: Santa Anna da Chapada, Mato Grosso, Brazil, Malme 2393' (S, lectotype).] Parmelia cruslacea Lynge, 1914: 108. [Type collection: Santa Anna da Chapada, Mato Grosso, Brazil, Malme (S, lecto- Parmelia silvatica var. pinnata Lynge, 1914: 120. [Type col- lection: Santa Anna da Chapada, Mato Grosso, Brazil, Malme 2393*** (S, lectotype).] Parmelia silvatica var. radiata Lynge, 1914: 120. [Type collec- tion: Santa Anna da Chapada, Mato Grosso, Brazil, Mahe 2393** (S, lectotype).] Thallus closely adnate on bark, whitish mineral gray, 2-6 cm in diameter; lobes sublinear, often crowded, 0.5-2 mm wide; upper surface plane, shiny, faintly maculate; medulla in the lower half antimony yellow; lower surface moderately rhizi- nate, the rhizines densely dichotomously branched. Apothecia common, adnate, 1-2 mm in diameter; spores 5p6p X lop-12y. P + orange-red (lichexanthone, protocetraric acid, and an unidentified anthraquinone). type).] CHEMISTRY.-cOrteX K--, medulla K--, C-, DIsTRIsuTIorv.-Southern Brazil. HABITAT.-On trees in open forests. REMARKS.-This rare lichen resembles H. livida externally but the chemistry is completely differ- ent. Some specimens are very closely adnate and have narrow lobes. These were described by Lynge under the synonyms listed above. SPECIMENS ExAMINm.-Brad: h4ato Grosso, Malme (S, UPS) in Lichenes Austroamericani 87 (G, H, LD, UPS), 2271 Ba (S), 2511 Ba (S). 68. Hypotrachyna singularis, new combination FIGURE 18b Parnzelia singularis Hale, 1974b:267. [Type collection: Cerros Calla Calla, Chachapoyas, Peru, Hutchinson and Wright 5704 (US, holotype; UC, isotype).] Thallus loosely adnate on branches, subcori- aceous, whitish to pale yellowish mineral gray, about 8 cm broad; lobes subirregular to sublinear, 3-5 mm wide, marginally dissected and lobulate, the lobules to 1 mm long, narrow; upper surface smooth and shiny, continuous; lower surface densely rhizinate, the rhizines densely dichot- omously branched. Apothecia numerous, adnate, to 5 mm in diameter; spores 6p7p X 10p-12p. CmMIsmY.-Cortex K+ yellow, medulla K-, C-, P- (atranorin and apparently a fatty sub- stance not further identified). DISTRIBUTION.-PerU. HABITAT.-oII tree in open forest at 3100 m elevation. RE~~ARKs.-HyPotrachyna singularis is charac- terized by the rather coriaceous thallus and nu- merous elongate marginal lobules. It is known only from the type collection. 69. Hypotrachyna sinuosa, new combination FIGURE 18c Lichen sinuosus Smith, 1809:2050. [Type collection: Dum- Parmelia sinuosa (Smith) Acharius: 1814:207. Parmelia despreauxii Delise ex Duby, 1830:602. [Type collec- tion: Vire, France, Despreaux and Briquebec (not seen).] Parmelia sinuosa var. virescens Kremplehuber, 1861:131. [Type collection: Mittenwald, Germany, Krempelhuber (M, lectotype; UPS, isolectotype).] Parmelia meyeri Zahlbruckner, 1905:82. [Type collection: Chimborazo, Ecuador, Meyer 365 (W, lectotype).] friesshire, England, Burgess (LINK, lectotype).] Thallus adnate to loosely adnate on bark and twigs, rarely on rock, sometimes suberect at the periphery, pale greenish yellow, 2-7 cm broad; lobes sublinear, short, 0.7-3 mm wide; upper sur- face plane, continuous, sorediate, less commonly pustulate-sorediate, the soralia capitate, orbicular to diffuse, subterminal; lower surface densely rhizinate, the rhizines sometimes forming a mar- ginal mat. Apothecia very rare, adnate, 1-4 mm in diameter; spores 7p-9p X lop-14y. CmMIsmY.--Cortex K-, medulla K+ yellow turning red, P+ orange (usnic acid and salacinic acid, sometimes with norstictic acid and a trace of stictic acid). 64 SMITHSONIAK CONTRIBUTIOSS TO BOTAKY FIGURE 18.--Species of Hypotrachyna: a, H. silvatica (Malnze in S); b, H. singularis (Hutchi?iso?z and It?right 57041; c, H. sinuosa (Hale 430Tla); d, H. steyermaikii (Steyermark 106123); e, H. subafinis (Imshaug 22531); f, H. sublaevigata (Hale 38810). (Scale in mm.) NUMBER 25 65 DIsTRIBuTIoN.-A~aska to Oregon, southeastern United States, Mexico, West Indies, Andean re- gion from Venezuela to Chile, Europe, south- eastern Asia from southern Japan to Java. HABITATS.-On trees (hardwoods and conifers) and rarely rocks and on small shrubs in the paramo at 2000-3500 m (to near sea level in boreal areas). REMARKS.-The development of soredia in this pantropical-montane, pantemperate species is ex- tremely variable. The type and the bulk of the specimens examined have discrete subterminal soralia and powdery soredia. At the other ex- treme, the soralia (as in Parmelia meyeri) may originate as laminal to subterminal pustules which become coarsely sorediate or rarely remain esored- iate. There are various intergradations between these types. Most of the problematic specimens have been collected in the West Indies or in the high Andes. The chemistry also is variable, typ- ically salazinic acid with or without norstictic acid but sometimes with stictic acid or different com- binations of these. We may be dealing with sev- eral intergrading morphological and chemical populations but for the present they are most con- veniently recognized as a single species. Smith cited three syntypes: the Dumfriesshire lectotype, a Menzies specimen from Nova Scotia (LINN, US) which can be identified as Par- melina galbina (Acharius) Hale, and a Turner and Hooker collection from Ballacheulish which I have not located. Parmelia despreauxii is rep- resented by collections in BM and G apparently identified by Delise himself, but the most appro- priate lectotype probably is housed in Strasbourg (STR). A syntype of P. meyel-i (Meyer 388) is not a Pamelia at all but is a member of the Lecanor- aceae. anishi 370 (KOBE) 393 (US); Puebla, Van Horn 59199 (US); Veracruz, Hale 20897. Costa Rica: San Jose, Culberson 12580, 13199 (DUKE); Haiti: Ouest, Zmshaug 23016 (MSC, US). Dom- inican Republic: La Vega, Zmshaug 23392, 23518 (MSC, US), Wetmore 3673 (MSC, US). Colom- bia: Boyaca, Cuatrecasas 1787a (US); Cundina- marca, Linclig 2548 (P); Santander, Killip and Smith 18595 (US). Venezuela: Merida, Hale 42711, 42794, 42820, 43210, Nash 1983 (US), Schwabe (M). Bolivia: Cochachamba, Ugent (US, SPECIhlENS ExAhfINED.-Mexico: OaXaCa, NUk- WIS). Chile: Aysen: Santesson 4603, 8413 (UPS); Juan Fernandez, Imshaug 37440 (MSC); Magal- lanes, Santesson 1848, 1994, 6441, 7827 (S), 6586 (UPS); Tierra del Fuego, Santesson 404, 420, 7864 (S), 5471 (UPS); Valdivia, Santesson 7262 (9 70. Hypotrachyna steyermarkii, new combination FIGURE 18d Parrnelia steyerrnarkii Hale, 1974b:268. [Type collection: Sierra Parima, Amazonas, Venezuela, Steyermark 106123 (US).] Thallus closely adnate on mosses on bark, frag- ile, light mineral gray, about 8 cm broad; lobes narrow, dichotomously branched, 1-2 mm broad; upper surface plane, shiny, isidiate, the isidia initially simple and cylindrical but soon pro- cumbent and in part lobulate, becoming ciliate; lower surface moderately rhizinate, the rhizines sparsely dichotomously branched. Apothecia not seen. Cmammu.-Cortex Kf yellow, medulla K-, C+ orange, P- (atranorin, barbatic acid, 4-0- demethylbarbatic acid, obtusatic acid, and nor- obtusatic acid). DISTRIBUTION.-VeneZUeIa. HABITAT.-On a small tree in forest at 1300 m elevation. REMARKS.-This species is distinguished from chemically identical H. imbricatula by the very narrow lobes and the consistently procumbent, ciliate isidia. 71. Hypotrachyna subafinis, new combination FIGURE 18e Parrnelia subafinis Zahlbruckner, 1929:213. [Type collection: Based on P. ufjiinis Vainio, not P. afinis (Dickson) Acharius, 1803:212 (= Pannuria).] Parrnelia ufiinis Vainio, 1890:52. [Type collection: Carassa, Minas Gerais, Brazil, Vainio 1331 (TUR, Vainio herbarium number 3010, lectotype; BM, isolectotype).] Thallus closely adnate on bark, sea-foam yel- low, 3-5 cm broad; lobes sublinear, dichotomously branched, 0.4-2 mm wide; upper surface plane, continuous, sorediate, the soralia capitate, mostly laminal; lower surface densely rhizinate, the rhizines often projecting along the margins. Apo- 66 SMITHSONIAN CONTRIBUTIONS TO BOTANY thecia rare, adnate, 0.7-2.5 mm in diameter; spores 5p-7p X 8p-11p. CHEMIsmY.-Cortex K+ yellowish, medulla K-, C-, P+ pale orange (usnic acid, echi- nocarpic acid, and associated unknowns). DIsTRIBuTIoN.-Mexico, Central America, West Indies, Brazil. HABITAT.-oII trees (oaks and pines) in open forests at 1000-2000 m elevation. REMARKs.-This is a very small, closely adnate species without any close relatives, except perhaps H. velloziae. TVhile Vainio?s type was collected on hardwood trees, almost all collections from the West Indies, where it is most common, were taken from Pinus bark. SPECIMENS EXAMINED.-Mexico: Chiapas, Hale 20352, 20384. Guatemala: Baja Verapaz, Hale 38308, 38342. Haiti: Ouest, Zmshaug 22531 (MSC, US), 22539, 22764, 23081, 23599, 23761 (MSC), Wetmore 3203 (MSC, US). Dominican Republic: La Vega, Allard 16542, 16544, 16552, 16581, 16589 (US); Santiago, Wetmore 3882 (MSC, US). Bra- zil: Minas Gerais, Vainio 771 (BM, M, TUR), 986, 3010 (TUR), 1391 (TUR, UPS). 72. Hypotrachyna sublaevigata, new combination FIGURE 1Sf Parmelia tiliacea var. sublaevigata Nylander, 1860:383. [Type collection: South America, Bonpland (P, lectotype; H, iso- lectotype),] Parmelia sublaevigata (Nylander) Piylander, 1867:306. Parmelia chilena Nylander, 18853612. [Type collection: Chile, M. Gay (P, lectotype).] Parmelin canescens Kurokawa in Hale and Kurokawa, 1964: 167. [Type collection: Colchagua, Chile, Dusen 92 (S, holo- type; US, isotype).] Thallus closely adnate on bark or twigs, 3-8 cm in diameter, dull mineral gray but often turn- ing red in the herbarium because of improper curating; lobes subirregular and short, crowded, 2-5 mm wide; upper surface plane to rugulose, continuous, often becoming whitish pruinose; lower surface densely short rhizinate. Apothecia common, substipitate, 2-5 mm in diameter; spores C~~~~sm~.--Medulla K+ yellow turning red, P + orange (atranorin, norstictic and salazinic DIsmInuTIoN.-Mexico, Panama, and Chile. acids). 5p-5~ X 8y-lOp. HABITAT.-on oaks, pine, Drimys, and Discaria in open or secondary forests at 1500-2100 m elevation. REMARKS.-The lobes of H. sublaeuigata tend to be subrotund, crowded, and closely adnate. Apothecia are usually numerous. In Panama this species could be confused with H. boquetensis, which contains only salazinic acid and has longer lobes. Nylander?s syntype of Parmelia tiliacea var. sublaevigata is Leprieur 504 (P), an isidiate species in the genus Bul bothrix Hale. Parmelia canescens was differentiated from H. sublaevigata only by the presence of norstictic acid, but later tests with thin-layer chromatography showed that salazinic acid also is present. SPECIMENS EXAMINED.-Mexico: Durango, Bell s.n. (DUKE), Fox M87 (US); Michoacan, Arsene 4046b, 4459b (US); Sinaloa, Culberson 13449 (DUKE). Panama: Chiriqui, Hale 38810. Chile: Curico, Mahu 3492 (US); Santiago, Mahu 28, 2133a (US); Tierra del Fuego, Santesson 6004 (S, US); Valdivia, Santesson 3719, 3730 (S, US); 3722 (S); Valparaiso, Mahu 1140 (US). 73. Hypotrachyna subphysodalica, new combination FIGURE 19a Parmelia subphysodalica, Hale, 1974b:268. [Type collection: Puerto Ballena, Chile, Imshaug 43121A (MSC, holotype; US, isotype).] Thallus closely adnate on twigs, yellowish green, 6-7 cm broad; lobes sublinear, about 2 mm wide; upper surface plane, shiny, moderately isidiate, the isidia cylindrical, simple, becoming short ciliate, about 0.3 mm high; lower surface moderately rhizinate. Apothecia not seen. CmmsmY.--Cortex K--, medulla K--, P + orange-red (usnic acid and physodalic acid). DIsmrnuTIoN.-Southern Chile. HABITAT.-oII tree in mossy forest near sea level. REMARKS.-The diagnostic features of this rare species are the yellow color, isidia, and presence of physodalic acid. While it might be mistaken for H. microblasta, its range is far south of that species. It is not related to the only other1 physodalic-acid-containing species in the genus, H. physodalica, a paramo lichen in the northern Andes. NUMBER 25 67 74. Hypotrachyna subplumbeata, new combination FIGURE 19b Parmelia subplumbeata Dodge, 1959: 106. [Type collection: Parrnelia kahusiensis Dodge, 1959: 105. [Type collection: Mt. Mt. Karisimbi, Congo, Derscheid 2504 (US, lectotype).] Kahusi, Congo, Hendrickx 4300 (EAH, holotype).] Thallus loosely adnate, rather coriaceous, 5-10 cm broad; lobes sublinear, crowded, 2-4 mm wide; upper surface smooth, heavily white-maculate, without soredia or isidia; lower surface densely rhizinate, the rhizines densely dichotomously branched. Apothecia abundant, substipitate, 2-5 mm in diameter; spores not developed. CHEMISTRY.-cOrteX K+ yellow, medulla K-, C+ orange, P- (atranorin, barbatic acid, and 4-0-demethylbarbatic acid). DIsTRIBUTI0N.-Mexico and Africa. HABITAT.-oII small trees near the tree line at about 3800 m elevation. FIGURE 19.-Species of Hypotrachyna: a, H. subphysodalica (Zmshaug 43121.4); b, H. subplum- beata (Moore and Wood 4944); c, H. subsaxntilis (Arsene 3974); d, H. thysanota (Hale 21118). (Scale in mm.) 68 SMITHSONIAN CONTRIBUTIONS TO BOTANY REMARKs.-According to Culberson and Hale (1973), H. subplumbeata is the remnant of an ancient population (now mostly extinct in the New World but widespread in the Old World) that contains barbatic acid without obtusatic acid. It could be the progenitor species which hybrid- ized with H. pziluinata to produce the obtusatic- acid-containing species in the H. physcioides con- stellation. The thallus is rather coriaceous and the maculae strongly developed, just as in the African population. SPECIMENS EXAiCIINED.-kfeXiCO: Mexico, Kiener 18577 (DUKE), Moore and Wood 4944 (US). 75. Hypotrachyna subsaxatilis, new combination FIGURE 19c Parmelia subsaxatilis Bouly de Lesdain, 1914:6. [Type collec- tion: Cerro Azul, Michoacan, Mexico, Arsene 3973 (P, lec- totype) .I Thallus loosely adnate on rock, dark mineral gray, 4-8 cm broad; lobes sublinear, short, crowded, 1-3.5 mm wide; upper surface plane to rugulose and cracked, lacking isidia and soredia; lower surface sparsely to moderately rhizinate, the rhizines sparsely dichotomously branched. Apothecia adnate, 1.5-4.5 mm in diameter; spores CHEiwsmY.-Cortex K+ yellow, medulla K+ yellow turning red, P+ orange-red (atranorin and salazinic acid). 7p-91.1 X 11p-15p. DISTRIBUTION.-Mexico. HABITAT.-on rocks and soil over rocks in open areas at about 2200 m elevation. REMARKS.-While amply distinct and collected in quantity by Arsene, this species remains a rarity known only from Mexico. It is unrelated to any other species in the genus. SPECIMENS EXAMINED.-MeXiCO: Michoacan, Arsene 3974, 3974b (US). 76. Hypotrachyna thysanota, new combination FIGURE 19d Parmelia thysanota Kurokawa in Hale and Kurokawa, 1964: 186. [Type collection: 11 km E Las Vigas, Veracruz, Mexico, Hale 20943 (US, holotype).] Partnelia nakanishii Hale, 1971d3434. [Type collection: Mt. Leconte, Sevier County, Tennessee, Nakanishi 205 (US, holotype; KOBE, DUKE, TKS, isotypes).] Thallus loosely adnate, whitish mineral gray, to 15 cm broad; lobes linear, dichotomously branched, subdivaricate, 2-5 mm wide; upper surface plane, shiny, becoming pruinose near the tips, sorediate, the soralia mostly subterminal, often protruding, coalescent; lower surface densely rhizinate, the rhizines densely dichotomously branched, often projecting as a mat beyond the margins. Apothecia not seen. CHEhmmY.-Cortex K+ yellow, medulla K-, C- red, P+ red (atranorin, echinocarpic acid, gyrophoric acid, and microphyllinic acid). DIsTRIBuTIoN.-Southeastern United States and Mexico. HABITAT.-oII conifers (Abies, Pinus) in open forests at about 2000 m elevation. REMARKs.-Gyrophoric acid was the only con- stituent identified with microcrystal tests for this species. Dey (1974) has analyzed the type speci- men with thin-layer chromatography and dis- covered the additional substances. This is the only report of microphyllinic acid outside of Cetraria, although Culberson and Hale (1973) postulated its possible occurrence in Hypotrachyna. Super- ficially, this species is similar to H. Zaeuigata or H. densirhizinata; no nonsorediate progenitor has been discovered. SPECIMENS ExAMINED.-Additional records from the United States and Mexico are listed in Hale and Kurokawa (1964:186), and in Hale (1971d: 434) under Parmelia nakanishii. 77. Hypotrachyna velloziae, new combination FIGURE 20 Parmelia velloziae Vainio, 1890:64. [Type collection: Carassa, Minas Gerais, Brazil, Vainio 1455 (TUR, Vainio herbarium number 2892, lectotype; BM, FH, M, UPS, isolectotypes).] Thallus closely adnate to bark, fragile, pale viridine yellow, 3-7 cm in diameter; lobes sub- linear, dichotomously branched, 0.5-2 mm wide; upper surface plane, continuous, sorediate, the soralia mostly laminal, capitate, often coalescing; medulla turning yellow-ochre under the soralia; NUMBER 25 69 lower surface moderately rhizinate, the rhizines sparsely dichotomously branched. Apothecia not seen. CHE;LIIsmY.-Cortex K-, medulla K-, PS orange-red (usnic acid, protocetraric acid, and rhodophyscin). DISTRIBUTION.-west Indies and Brazil. HABITAT.-on trunk of trees (Pinus) in open or virgin pine forests (West Indies) at 1200-1600 m elevation. REhlARKs.-H~potraChylza uelloziae parallels H. subafinis in distribution (mainly West Indies) and habitat (pine trees) but differs completely in chemistry. There is no nonsorediate sexual morph. SPECIMENS ExAMINED.-Haiti: Ouest, Imshaug 22535 (MSC, US), Wetmow 3071, 3217 (MSC, US). FIGURE 20.-Hypotrachyna uelloziae (Wetmore 307). Dominican Republic: La Vega, Allard 16541a, (Scale in mm.) 16562d (US). Literature Cited Acharius, E. 1803. Methodus qua omnes detectos Lichenes, 394 pages. 1814. Synopsis methodica lichenum. 392 pages. Lund. 1942. Distribution and Ecology of Some South Scandi- Stockholm. Almborn, 0. navian Lichens. 252 pages. Lund. .4sahina, Y. 1951. Lichenes Japoniae novae vel minus cognitae (7). 1953. Lichenes Japoniae novac vel minus cognitae (10). 1968. Lichenologische Notizen (209). Journal of Japanese Baker, C. J., .4. Elix, D. P. H. Murphy, S. Kurokawa, and R. H. Grover Parmelia reptans, a New Lichen Species Producing the Depsidone, Succinprotocetraric .Acid. Austra- lian Journal of Botany, 213137-140. Journal of Japanese Botany, 26:329-334. Journal of Japanese Botany, 28:65-68. Botany, 43:97-101. 1973. Bouly de Lesdain, hl. Crombie, J. M. 1914. Lichens de Mexique. 31 pages. Mexico. 1876. On the Lichens Collected by Prof. R. 0. Cunning- ham. Journal of the Linneaiz Sociely of London, 15~222-230. Culherson, C. F. 1966. The Structure of a New Depsidone from the Lichen Parmelia livida. Phytochenistry, 53835-818. 1972a. Improved Conditions and New Data for the Iden- tification of Lichen Products by a Standardised Thin-Layer Chromatographic hfcthod. Journal of Chromatography, 72:113-125. 1972b. High-speed Liquid Chromatography of Lichen Ex- tracts. Bryologist, 55:54-62. Culberson, C. F., and M. E. Hale, Jr. Chemical and RloIphological Evolution in Parmelia sect. Hypotmchynn: Product of Ancient Hybfidiza- tion? Brittonia, 253162-173. The Parmelia qum~ina Group in North America. American ,Journal of Botany, 48: 168-174. Chemosystematics and Ecology of Lichen-Forming Fungi. Annual Xeuiew of Ecology and Systematics, 1 : 153-1 70. The Parmelia perforata Group: Siche Characteris- tics of Chemical Races, Speciation by Parallel Evo- lution, and a Sew Taxonomy. Bryologkt, 76:20-29. 1967. Hahitat Selection by Chemically Differentiated Races of Lichens. Science, 158:1195-1197. 1970. .A Phylogenetic View of Chemical Evolution in thc Lichens. Bryologist, 73:l-31. 1973. Parallel Evolution in Lichen-Forming Fungi. Science, 180:196-198. 1973. Culberson, TV. L. 1961. 1970. 1973. Culherson, Tt?. L., and C. F. Culberson Degelius, G. 1941. Contributions to the Lichcn Flora of h?orth Amer- ica, 11. The Lichen Flora of the Great Smoky Mountains. Arkiv for Botanik, 304 (3):l-80. Dey, J. 1074. Nerc- Interesting I?armcliae (Lichens) from the Southern .Appalachianr. Castanea, 39:360-369. Some Lichens of Tropical Africa, 111. Parmeliaceae. Annals of the Missouri Botanical Garden, 46:39-193. Dodge, C. TV. 1959. 70 SMITHSONIAN CONTRIBUTIONS TO BOTANY Duby, J, E. 1830. Aug. Pyrami de Candolle Botanicon gallicum. 1068 pages. Paris. Durand, T., and H. Pittier 1891. Primitae Florae Costaricensis. Volume 2, pages 49- 97. Bruxelles. Du Rietz, G. 1924. Die Soredien und Isidien der Flechten. Svensk Botan- Fee, A. 1824. isk Tidskrift, 18:371-396. Essai sur les cryptogames des ecorces exotiques ofi- cinales. 167 pages. Paris. Floerke, H. G. Gyelnik, V. 1815. Deutsche Lichenen, I. 18 pages. Rostock. 1931. Additamenta ad cognitionem Parmeliarum, 11. Fedde Repertorium, 29:273-291. 1934. Additamenta ad cognitionem Parmeliarum, V. Fedde Repertorium, 36: 151-156. 1938a. Revisio typorum ab auctoribus variis descriptorum. Annales Musei Nationalis Hungarici, 31:2-57. 1938b. Additamenta ad cognitionem Parmeliarum. An- nales Mycologica, 36:267-294. Hale, M. E., Jr. 1959. New or Interesting Parmelias from North and Tropical America. Bryologist, 62: 123-132. 1960. A Revision of the South American Species of Parmelia Determined by Lynge. Contributions from the United States National Herbarium, 36: 1-41. 1963. Populations of Chemical Strains in the Lichen Cetraria ciliaris. Brittonia, 15: 126-133. 1965. A Monograph of Parmelia subgenus Amphigymnia. Contributions from the United States National Her- barium, 36: 193-358. 1967. New Taxa in Cetraria, Parmelia, and Parmeliopsis. Bryologist, 70:414-422. 1971a. Morden-Smithsonian Expedition to Dominica: The Lichens (Parmeliaceae). Smithsonian Contributions to Botany, 4:l-25. 1971b. Five New Parmeliae from Tropical America. Phy- tologia, 22:30-35. 1971c. New Parmeliae (Lichens) from Africa. Phytologia, 2294-96. 1971d. New Species of Parmelia sect. Hypotrachyna (Lichenes). Phytologia, 22:433438. 1972a. Fine Structure of the Cortex in the Lichen Family Parmeliaceae Viewed with the Scanning-Electron Microscope. Smithsonian Contributions to Botany, 1972b. Parmelia pustulifera, a New Lichen from South- eastern United States. Brittonia, 24:22-27. 1974a. Delimitation of the Lichen Genus Hypotrachyna (Vainio) Hale. Phytologia, 28:340-342. 1974b. New Species of Parmelia (Lichenes) from Tropical America, 1. Phytologia, 28:265-269. 10: 1-92. Hale, M. E., Jr., and S. Kurokawa 1964. Studies on Parmelia subgenus Parmelia. Contribu- tions from the United States National Herbarium, 36: 121-191. Hillmann, J. 1936. Parmeliaceae. Section 3, pages 1-309, in part 5, volume 9, in L. Rabenhorst, Rabenhorst's Krypto- gamenpora. Leipzig. Hue, A. M. 1899. Lichenes extra-europaei. Nouvelles Archives du Museum Paris, series 3, 1:l-250. Species novae in excelsis Ruwenzori in expeditione Duck Aprutii lectae, IV: Lichenes. Annuli di Botanica, 6:407-409. Verzeichnis der Lichenen, welche M. Wagner auf seinen Reisen in Centralamerika 1858/59 gesammelt hat. Flora, 44:129-131. 1873. Lichenes brasilienses in Warming Symbolae ad floram Brasiliae cognoscendam particula, XIV. Videnskabelige Meddelelser fra Dansk Naturhis- torisk forening i Kjobenhavn, 25: 1-35. The Lichen-Flora of Great Britain, Ireland and the Channel Islands. 502 pages. Shrewsbury. VIII. Memoir on the Spermagones and Pycnides of Filamentous, Fruticulose, and Foliaceous Lichens. Transactions of the Royal Society of Edinburgh, Jatta, A. 1908. Krempelhuber, A. 1861. Leighton, W. A. 1871. Lindsay, W. L. 1859. 22:lOl-303. Lynge, B. 1914. Die Flechten der ersten Regnellschen Expedition. Die Gattungen Pseudoparmelia gen. nov. und Parmelia Ach. Arkiv for Botanik, 13 (13):l-172. 1917. Ueber einige Regnellschen Parmelian aus Matto Grosso, Brasilien. Arkiv for Botanik, 15 (1):1-4. 1925. On Some South American Lichens of the Genera Parmelia, Candelaria, Teloschistes, and Pyxine. Nyt Magazin for Naturvidenskaberne, 62:83-97. Mitsuno, M. 1953. Paper Chromatography of Lichen Substances, I. Pharmaceutical Bulletin (Tokyo), 14:659-669. Muller Argau, J. 1879. Lichenes aequinoctiali-americani a cl. E. Andre annis 1875-76 praesertim in editroribus Ecuador et in Nova Granata lecti. Revue Mycologique, 1:163- 171. 1880. Lichenologische Beitrage, XI. Flora, 63:259-268. 1881. Lichenologische Beitrage, XII. Flora, 64:81-88. 1887. Lichenologische Beitrage, XXVI. Flora, 70:316-320. 1889. Lichenes Sebastianopolitani lecti a cl. Dr. Glaziou. Nuovo Giornale Botanic0 Italiano, 21:353-364. 1891. Lichenologische Beitrage, XXXV. Flora, 74:371-382. 1860. Synopsis methodica lichenum, I. 430 pages. Paris. 1863. Lichenographiae Novo-Granatensis Prodromus. Acta Nylander, W. Societas Scientarum Fennica, 7:415-504. NUMBER 25 1867. Lichenes. Additamentum in J. Triana and J. E. Planchon, Prodromus Florae Nova-Granatensis ou Enumeration des Plantes de la Nouvelle-Grenade. Annales Sciences Naturelles Botanique, series 5, I: 1874. Addenda nova ad Lichenographiam europaeam. 1885. Parrneliae exoticae novae. Flora, 68:605-615. 1970. Das Konzept der Artenpaare bei den Flechten. . Deutsche Botanische Gesellschaft Neue Folge, 4: Die taxonomische Behandlung von Artenpaare bei den Flechten. Botaniska Notiser, 125:77-81. Polakowsky, H. Bryophytas et cormophytas costaricensis anno 1875 lectas. Journal of Botany British and Foreign, 13: 225, 226. Rebentisch, J. F. Prodrornus Florae Neomarchicae. 406 pages. Berlin. Smith, J. E. 301-354. Flora, 573305-318. Poelt, J. 187-198. 1972. 1877. 1804. 1808. English Botany. Volume 26, page 1852. London. 1809. English Botany. Volume 29, page 2050. London. 1827. Systerna Vegetabiliurn, 4. Edition 16, part 2, 410 Sprengel, K. pages. Gottingae. Stahl, E., and P. J. Schorn 1961. Dunnschicht-chromatographie hygrophiler Arzneip- flanzenauszuge, VIII. Mitteilung: Cumarine, Flavon- derivate, Hydroxysauren, Gerbstoffe, Antracenderiv- ate und Flechteninhaltsstoffe, Hoppe-Seyler's Zeitschrift fur Physiologische Chemie, 325:263-274. 1903. Flechten von Kamerun und dem Kamerunberg (Fako), gesammelt von Alfred Bornmuller in den Jahren 1897 und 1898. Verhandlungen der Kaiserlich-koniglichen zoologisch-botanischen Gesell- schaft in Wien, 53:227-236. Steiner, J. Stirton, J. 1874. New British Lichens. Grevillea, 3:79. Taylor, T. 1847. New Lichens, Principally from the Herbarium of W. J. Hooker. Hooker Journal of Botany, G:148- 197. Thomson, J. W. 1961. Evolution in the Lichen Genus Physcia. Pages 267- 271 in Recent Advances in Botany. Toronto: Uni- versity of Toronto Press. 71 Vainio, E. A. 1890. 1896. 1898. 1899. 1900. 1907. 1915. Etude sur la classification naturelle et la morphol- ogie des Lichens du Bresil. Acta Societas Fauna et Flora Fennica, 7 (7):l-247. Lichenes Antillarum a W. R. Elliott collecti. Journal of Botany British and Foreign, 34:31-36. Lichenes quos in Madagscaria centrali Dr. C. For- syth Major collegit. Lichenes novi rarioresque, I. Hedwigia, 38: (121)- (125). Reactiones lichenum a J. Mullero Argoviensi de- scriptorum. Memoire E'Herbier Bossier, 5: 1-17. Lichenes novi rarioresque, IV. Hedwigia, 46: 168- 181. Additamenta ad lichenographia antillarum illus- trandam. Annales Academiae Scientarurn Fennicae, Hedwigia, 37: (33)- (37). 6~ (7):l-226. Vareschi, V. 1962. Resultados liquenologicos de excursiones effectuadas en Venezuela, No. 2: Acerca del genero Parrnelia en Venezuela. Acta Biologica Venezuelica, 3:201-232. Studies on the Chemistry of Lichens, I: Separation of Depside Components by Paper Chromatography. Acta Chernica Scandinavica, 6: 81 8-825. Wachtmeister, C. A. 1952. Zahlbruckner, A. 1902. 1905. 1908. 1909. 1912. 1926. 1929. 1934. Studien uber brasilianische Flechten. Sitzungs- berichte der kaiserliche Akademie Wissenschift. Mathematisch-naturuGssenschajtliche Klasse, .4bteil- ung 1, 111:357432. Flechten, im Hochlande Ecuadors gesammelt von Prof. Dr. Hans Meyer im Jahre 1903. Beihefte zur Botanischer Centralblatt, 19:75-84. Beitrage zur Flechtenflora Brasiliens. Bulletin de 1'Herbier Bossier, series 2, 8:459-468. Lichenes (Flechten). In Ergebnisse der botanischen Expedition der kaiserlichen Akademie der TVissen- schaften nach Sudbrasilien 1901. Denkschrift Akademie der Wissenschajt in Wien. Mathematisch- naturwissenschaftliche Klasse, 83:87-211. Neue Flechten. Annales Mycologici, 103359-384. Chilenische Flechten gesammelt von C. Skottsberg. Meddelanden fran Goteborgs Botaniska Tradgard, Catalogus lichenum universalis. Volume 6, 323 pages. Leipzig. Flechten der Insel Formosa. Fedde Repertoriitm, 2: 1-26. 33:22-68. Index (Synonyms in italics) Hypotrachyna andensis, 23 bahiana, 23 bogotensis, 23 boquetensis, 25 brasiliana, 25 brevirhiza, 26 caraccensis, 26 chicitae, 28 chlorina, 28 consimilis, 28 cantradicta, 29 costaricensis, 29 croceopustulata, 30 dactylifera, 30 degelii, 31 densirhizinata, 31 dentella, 33 ducalis, 33 eitenii, 33 enderythraea, 34 endochlora, 34 ensifolia, 35 erythrodes, 35 exsplendens, 37 flavida, 37 flavovirens, 37 formosana, 38 gigas, 38 gondylophora, 40 gracilescens, 40 imbricatula, 41 immaculata, 41 intercalanda, 42 isidiocera, 42 koyaensis, 44 laevigata, 44 lineariloba, 45 livida, 45 lopezii, 47 malmei, 47 microblasta, 47 minima, 48 monilifera, 48 neodissecta, 49 novella, 49 obscurella, 51 oostingii, 51 osorioi, 51 osteoleuca, 51 palmarum, 52 partita, 52 peruviana, 54 physcioides, 54 physodalica, 55 pluriformis, 55 producta, 56 prolongata, 56 protenta, 58 protoboliviana, 58 pseudosinuosa, 58 pulvinata, 59 rachista, 59 reducens, GO revoluta, GO rhabdiformis, 62 rockii, 62 silvatica, 63 singularis, 63 sinuosa, 63 steyermarkii, 65 subaffinis, 65 sublaevigata, 66 subphysodalica, 66 subplumbeata, 67 subsaxatilis, 68 thysanota, 68 velloziae, 68 sinuosus, 63 amoena, 29 anaptychioides, 58 bahiana, 23 bogotensis, 23 boliviana, 54 boliviana var. cephalota, 44 boquetensis, 25 brasiliana, 25 brasiliana var. erythrodes, 35 brasiliana var. novella, 49 brevirhiza, 26 canescens, 66 caraccensis, 26 caraccensis var. guatemalensis, 26 caraccensis f. pulvinata, 59 chicitae, 28 Lichen laevigatus, 44 Parinelia afinis, 65 chilena, 66 chlorina, 28 confusula, 54 consimilis, 28 contradicta, 29 costaricensis, 29 croceopus t ti la ta, 30 crustacea, 63 culmigena, 23 dactylifera, 30 deformis, 29 degelii, 31 densirhizinata, 31 dentella, 33 despreauxii, 63 digitata, 54 ducalis, 33 eitenii, 33 enderythraea, 34 endochlora, 34 endochlora f. sorediosa, 34 endorubra, 26 endorzrbra f. imbricatiformis, 48 ensifolia, 35 eryt h rocard ia, 34 erythrodes, 33 everniaeformis, 55 exporrecta, 38 exsplendens, 37 flava, 37 flava f. albescens, 23 flava var. stellata, 37 pava var. subdichotoma, 37 flavida, 37 flavovirens, 37 formosana, 38 fragilis, 42 gallicana, 34 gigas, 38 glaziovii, 25 gondylophora, 40 gracilescens, 40 gracilescens f. obscurella, 51 gracilis, 54 hypotrachyna, 29 imbricatula, 41 immaculata, 41 insinuans, 23 72 NUMBER 25 73 intercalanda, 42 isidiocera, 42 jamaicensis, 48 kahusiensis, 67 koyaensis, 44 laevigata, 44 laevigata var. gracilis, 54 lineariloba, 45 livida, 45 lobulifera var. insensitiva, 35 lobulifera var. luteoreagens, 41 lobulifera var. sanguineoreagens, 59 madagascariensis, 34 malmei, 47 meyeri, 63 microblasta, 47 millaniana, 34 minima, 48 monilifera, 48 nakanishii, 68 neodissecta, 49 neopropagulifera, 48 norstictica, 48 novella, 49 oostingii, 51 osorioi, 51 osseoalbida, 45 osteoleuca, 51 palmarum, 52 peruviana, 54 physcioides, 54 physodalica, 55 pittieri, 34 pluriformis, 55 pluriformis var. chlorocarpa, 55 pluriformis var. multifida, 55 prolongata, 56 propagulifera, 48 protoboliviana, 58 pseudorevoluta, 48 pseudosinuosa, 58 pulvinata, 59 rachista, 59 reducens, 60 regis, 49 revoluta, 60 revoluta f. isidiosa, 48 rhabdiformis, 62 rockii, 62 silvatica, 63 silvatica var. pinnata, 63 silvatica var. radiata, 63 singularis, 63 sinuosa, 63 sinuosa var. caraccensis, 26 sinuosa var. virescens, 63 steyermarkii, 65 subafinis, 65 sublaevigata, 66 sublaevigata f. isidiosa, 29 subphysodalica, 66 subplumbeata, 67 subpluriformis, 55 subsaxatilis, 68 subsinuosa, 59 thysanota, 68 tiliacea var. leucina, 30 tiliacea var. sublaevigata, 66 tropica, 29 tropica var. deformis, 29 velloziae, 68 xanthomyela, 34 xanthomyela f. sorediosa, 34 zahlbruckneri, 52 * U.S. GOVERNMENT PRINTING OFFICE: 1975- 586-378/19