A Monograph of the Lichen Genus Buzbothrix Hale (Parmeliaceae)
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SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 32 A Monograph of the Lichen Genus BuZbothrix Hale (Parmeliaceae) Mason E. Hale, Jr. SMITHSONIAN INSTITUTION PRESS City of Washington 1976 ABSTRACT Hale, Mason E., Jr. A Monograph of the Lichen Genus Bulbothrix Hale (Parmeliaceae). Smithsoniun Contributions to Botany, number 32, 29 pages, 7 figures, 1976.-A world-level revision is given for the 29 species of Bulbothrix, a generic segregate of Parmelia characterized by marginal bulbate cilia and pro- duction of atranorin in the upper cortex. The genus is primarily tropical in distribution and best developed in secondary forests. The main center of specia- tion is Brazil with 14 species, and the genus is also well represented in Africa. The New World species have a high frequency of depsides and orcinol dep- sidones, whereas the Old World species usually contain p-orcinol depsidones. One new species, B. klementii Hale, and one new combination, B. goebelii (Zenker) Hale, are proposed. OFFICIALI~~'I~C,IC,~TIOS D.yrI: is handstamped in a limited number of initial copics and is rccordetl in thc Institution's annual rcport, Suiithsoninn Yew. SI:RII:SCOVER DMICS: Leaf clcaring from the katsura trcc Cercidiphyllirm jnponicum Siebold and Zuccaritii. Libra1y of Congrcss Cataloging in Publication Data Hale, Mason E. A monograph of the lichcn gcnus Bulbothrix Halc (Parmcliaccac). (Smithsonian contributions to botany : no. 32) Bibliography: p. Includes index. Supt. of Docs. no.: SI 1.29:32 1. Bulbothrix. I. Titlc. 11. Series: Smithsonian Institution. Smithsonian contributions to botany ; no. 32. QKlS2747 110. 32 [QK589.P2] 581'.08~ [589'.1] 75-619423 Contents Pnge Introduction 1 Morphology 1 Chemistr) 3 hforph Formation 1 Phj togeography 5 Classification of Bulbothrix 6 Key to the Species 7 Taxonomic Treatment 9 Literature Cited 27 Index 29 ... 111 A Monograph of the Lichen Genus Bulbothrix Hale (Parmeliaceae) Mason E. Hale, Jr. Introduction Smithsonian Expedition to Dominica, Sational Geographic Society, and the Smithsonian lie- This world monograph of the 29 species of search Foundation. Bztlbothl-ix is part of my continuing effort to revise the parnielioid genera, utili~ingthe most up-to-date analytical tools available, in particular Morphology tlie scanning-electron microscope and thin-layer THE THALr.us.-species of b’iilboth rix have chromatography. Genera already completed in- small, adnate to closely adnate tlialli. The lobes clude Nypotl-nchyna (Vainio) Hale (Hale, 1975a) are suhlinear to more rarely sul,irregular, clepentl- (the neotropical species), Relicinn (Hale and ing on the width. In these respects Rlrlbothrix is Kurokawa) Hale (Hale, 1975b), and Pseudopa?.- closest to Relicinn and Pai.nielinn. The most cliar- melia Lynge (Hale, 1976). Much of the intro- acteristic feature, however, is the development of’ ductory material in these revisions applies equall) marginal bulbate cilia, which are essentially well to Bulbothl-ix and will not be repeated here. identical with those of Relirinn antl regularly dis- I am especiallv thankful to curators of the various persed around the lobe tips antl margins. Tl’lien museums and herbaria, as listed Ly tlie standard rhizines are dichotomously Ixinchetl, the tips of herbarium acronyms under specimens examined, these bulbae may also be branched. who so generously and promptly lent speci- LOWERSURFACE AND RHmNr;.s.-The lower sur- mens for study. Special thanks for assistance in face is black in some of tlie species, dark or pale carrying out field studies are due Dr. M. Lopez- brown in others. Rhizines are either simple to Figueiras of the Universidad de 10s Andes, hferida, sparsely furcate, as in B. nfixn, A. chorc~oct7.ti.s.B. Venezuela; Dr. P. G. Patwardhan, Maharashtra deciwtatn, B. fungicola, B. hypori~nea,R. imshcizrgii Association for tlie Cultivation of Science, Poona, (Figure 1 b), R. isidiza, B. nieizo.rpom, B. pir,~tii- India; and Dr. Flora Uyenco, University of the la t a, B . .ren si b i 1 i.r, B . .re t sc h 7~ 1 n n c n .r i,s, R . .s i i b c om- Philippines, Quezon City. Dr. S. Kurokawa a\- nata, B. siibinflatn, B. tnbncinn, and R. 7~entl-ico,sn, shed with descriptions and preliminary identifi- or more or less richly dichotomously In-anched, as cations of many of the species in 1961. in B. apophysata, B. ntrichelln, R. bicoi~~rtn,B. The following grants received since 1960 are biilbochaeta, B. confoedercltn, R. coi-niitn, R. acknowledged with thanks: Sational Science goebelii, B. klenientii, R. lncuignfiiln (Figure lc), Foundation (Systematic Biology and tlie U.S.- B. schiffnel-i, I?. semilzrnatn, B. ,sztfixn, antl R. Japan Cooperative Science Program), Morden- viiidescens. Branching is a useful and consistent Mason E. Hnlr, Jr., Departnient of Botany, Xutionnl Zflrcrttni taxonomic character. of .Vatuinl History, Smithsonian Institution, Washinqton, There is a close correlation bettveen chemistry D.C.20560. and btanching pattern of the rhizines. All P+ 1 2 SMITHSONIAN CONTRIBUTIONS TO BOTANY FIGURE1.-Morphology of Rnlhothrix: n, cross section of 13. lnerjigntuln showing epicortex (arrow) (Allard 15992a in US) (X 500); b, simple rhizines of B. irnshaugii (Mnhu 3545 in US) (X 80); c, dichotomously branched rhizines of n. lnevigatiiln (Allard 15992a in US) (X 130); d, soralia of B. imsharigii (Rtmdel 7308 in US) (X 10); e, pustules of B. piistdata (holotype) (X 10); f, lobules of B. sufiixa (Hale 42753) (X 10); g, pycnidiate apothecia of B. uiridescens (Imshaug 42530 in US) (X 10). (Photographs Q-c taken with scanning-electron microscope.) NUMBER 32 3 species, that is, those containing norstictic acid, species they are almost substipitate because of a protocetraric acid, or salazinic acid, have simple constricted base. Pycnidia (coronate apothecia) unbranched rhizines. Most of these are Old World are always produced around the rim in B. afixa, species. On the other hand, the remaining spe- B. bulbochaeta, B. CoTonnta, B. fungicola, B. schif- cies, all of which are P- and contain gyrophoric fneri, B. .semilzinnta, B. subcoronato, B. ventricosa, acid, lecanoric acid, similar orcinol depsides and and B. viridescens. The thalli of these species (and depsidones, or only atranorin, have richly branched in the case of B. biilbochnetn, B. coronata, and B. rhizines and occur predominantly in the Nelv viridescens (Figure 1 g) the amphithecium) are World. The only exceptions are B. afixa and B. sometimes heavily pycnidiate with normal im- fungicola, which have mostly simple rhizines mersed pycnidia. These apothecial pycnidia are although they contain gyrophoric acid. As far as apparently the same as those I described for Reli- I know, no other parmelioid genus shows such a rina (Hale, 19751,:8), the only other parmelioitl clear-cut correlation between chemistry and mor- genus with this trait. phology except the Pnrrnelza borrerz group (Hale, Ecoronate apothecia are characteristic of the 1965a), which shows a direct correlation between remaining 15 fertile species. Bzilbothrix npopky- a pale lower surface and the presence of lecanoric sata, B. decirrtntci, B. klementii, and B. piginen- acid and a black lower surface with gyrophoric tncea have never been collected with apothecia. acid. There is no obvious correlation between chem- VFGETATIIF PRoP.uuLEs.--Isidia are regularly istry and presence of coronate apothecia, but the produced in B apophysata, B. decititata, B. fltngt- geographical correlation is strong. All coronate cola, B. goebelii, B. isidiza, B. klementii, B. lnevi- species, excepting B. afixa from Angola and A. qntiiln, B. pigmentacen, B. szibinpatn, B. tnbacina, bitlbochaeta from India, are endemic to or more and B. uentrirosa. They are cylindrical and usuall\ common in the New TZrorltl. Most species endemic unbranched but become procumbent and nearlv to the Old T\'orlcl are ecoronate. dorsiventral in B. fungicola. Dense lamina1 lobules The spores of Ritlbothi.ix are colorless, one- without isidia occur in B. sitfixa (Figure If). celled, and either ovoid or bicornute. Ovoid These are ti ue lobules since they are corticate and spores are rather small, 5-8 ym wide and 6-12 pm may produce marginal bulbate cilia. long on the average. Only B. bulbochnftn, R. Pustules are found in only one species, B. pust- ch07~~oen.sis,B. confoederntn, and 23. Iat4gatitla itlata. These large, inflated structures (Figuie have spores no more than 5-7 ym long, the typi- le) tend to break open apically and seem identical cal range for species of Relicinn (Hale, 197511). with those described in the genus Hypot)achyna As a rule, species with salazinic acid, these being (Hale, 1975a:6). Thev do not form soredia. largely endemic to the Old \l'orld, have the largest Soredia occur only in B. irnshairgii, a Chilean spores, 8-20 pm long. Bicornute spores are known endemic. The soralia are rather diffuse over the for three apparently closely related Brazilian spe- upper surface toward the lobe tips, producing. cies, B. bicornirtn, B. .tchifj'neri, and B. .rrmilirnnta. large patches of farinose soredia (Figure 1d). These spores are quite variable in shape, from A rather large percentage of the species in the nearly ovoid ivitli two short but distinct horns to gent15 produce no special vegetative propagules long, narrow, and semi-lunate. This type of spore other than small adventitious lobules so common has recently heen discovered in trvo species of in all paimelioid genera. This group includes B Relicinn (Hale, 1975b), but other than these five afixn, B at?ichella, B bicorniita, B. biilbochnetn, examples no other species in any parmelioid B. cho7ooenrir, B. confoederata, B. coronntn, B. genera have bicornute spores. \Ye cannot, how- hypocram, B. mezzospora, B. sztbcoronata, B ever, determine the evolutionary significance of schiffneri, B. yemilunata, B. yensibiliy, B. setsch7cw- unique spores until more is known of their cytol- nensis, and B. viridescen y. ,411 have abundant ogy and ontogen?. a po theci a. APOTHECIAICHARACTER$ -The apothecia of Bitl- Chemistry bothiix are uniformlJ small (rarely more than 4 mm in diameter) and sessile or adnate.