A Monograph of the Lichen Genus Parmelina Hale (Parmeliaceae)

A Monograph of the Lichen Genus Parmelina Hale (Parmeliaceae)

SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 33 A Monograph of the Lichen Genus Parmelina Hale (Parmeliaceae) Mason E. Hale, Jr. SMITHSONIAN INSTITUTION PRESS City of Washington 1976 ABSTRACT Hale, Mason E., Jr. A Monograph of the Lichen Genus Parmelina Hale (Par- meliaceae). Smithsonian Contyibutions to Botany, number 33, 60 pages, 21 figures, 1976.-The 47 species of Parmelina are revised on the world level. Two sections are recognized: section Parmelzna with 30 species widely distributed in temperate to tropical montane regions and section Myelochroa with 17 terpene- containing species concentrated in eastern and southern Asia. The genus is most closely related to Parmotrema hlassalongo. Five new species, P. crassata Hale, P. degelii Hale P. indica Hale, P. rhytidodes Hale, and P. schindleri Hale, are described, and six new combinations proposed, P. amagiensis (Asahina) Hale, P. damaziana (Zahlbruckner) Hale, P. endoleuca (Taylor) Hale, P. irrugans (Nylander) Hale, P. jamesii (Hale) Hale, and P. pastillifera (Harmand) Hale. New combinations are also made for Hypotrachyna baguioensis (Hale) Hale and Parmotrema nylanderi (Lynge) Hale. OFFICIALPUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution’s annual report, Srnithsonian Year. SERIES COVER DESIGN: Leaf clearing from the katsura tree Cercidiphyllum juponicum Siebold and Zuccarini. Library of Congress Cataloging in Publication Data Hale, Mason E. A monograph of the lichen genus Parmelina Hale (Parmeliaceae) (Smithsonian contributions to botany ; no. 33) Bibliography: p. Includes index. Supt. of Docs. no.: SI 1.29:33 1. Parmelina. I. Title. 11. Series: Smithsonian Institution. Smithsonian contributions to botany : no. 33. QKl.SZ747 no. 33 [QK585.P2] 581’.08s [589’.1] 76-608065 Contents Page Introduction ................................................. 1 Morphology ............................................... 2 Chemistry ................................................... 8 Phytogeography and Speciation ............................... 10 Classification of Parmelina ...................................... 14 Key to the Species of Parmelina ......................... 16 Species Treatment . , ..... ... ...... 18 Doubtful and Rejected Names .................... .. ... 53 Literature Cited ......................................... 55 Index ....................................... 59 iii A Monograph of the Lichen Genus Parmelina Hale (Parmeliaceae) Mason E. Hale, Jr. Introduction and Dr. R. Santesson were also geneiously placed at my disposal. The material in these various her- The genus Parmelina Hale is a segregate of Par- baria was chemically tested and annotated between melia Acharius comprising 47 species widely dis- 1958 and 1975, but not all of it has been re- tributed in temperate and tropical regions. It is examined for possible name changes in the light of recogni7ed by the narrow, adnate lobes, marginal new synonymies, species concepts, or improved cilia, and absence of usnic acid in the cortex (Hale, chemical techniques. 1974:438). Some of the species included here were Any monograph is bound to be more complete if formerl) classified in the ill-defined (and invalid) the author has had an opportunity to observe and genus Zmbitca?ia (Schreber) Michaux (Hale and collect specimens in the field. I am especially grate- Kurokawa, 1964: 130). This world level monograph ful to the following colleagues for assistance in is based on the study of all available type speci- arranging excursions: blr, J. Anderson (Sarawak), mens, collections preserved in the major herbaria, Alrs. Sheila Collenette (Sabah), Dr. S. Kurokawa and personal field work. The format follows my (Japan), Dr. 51. L6pez-Figueii-as (l’enezuela), Dr. earlier treatments of Bulbothrtx (Hale, 1976b), W. Meijer (Sabah), Dr. hI. Nakanishi and Dr. S. Hypotrachyna (Hale, 1975a), Pseudopawnella Nakanishi (Japan), Dr. P. G. Patwardhan (India), (Hale, 1976a), and Rellcina (Hale, 197513). Dr. Stella Thrower (Hong Kong), and Dr. Flora I wish to thank curators of the following insti- Uyenco (Philippines). I have also conducted field tutions who sent specimens on loan so promptl) and studies in Llexico and Central America, the Lesser often allowed extensions of loan periods: BM, BO, Antilles, and Malaya. BP, COLO, DUKE, F, FH, FI, G, GLAM, H, KAN, The scanning-electron photographs were pre- KR, L, LD, Rl, LlICH, NSC, RlVAf, NSLV, NY, pared by the Smithsonian Scanning-Electron Rficro- PC, PH, REX;, S, SI, TNS, TRH, TUR, UC, UPS, scope Laboratory. Photographs of the specimens W, LVIS, WU, and ZT. Specimens in TNS were were taken by the Smithsonian Photographic Ser- annotated during 1964-65 while I studied in Tokyo. vices. The priyate herbaria of Dr. D. D. Awasthi, Dr. Grants in support of this ret,earch have been re- Gunnar Degelius, Dr S Nakanishi, Dr. Clyde Reed, ceived from the National Science Foundation (Sys- tematic Biology and the U. S-Japan Cooperative Mason E Hale, Jr, Department of Botan), National Mu- Science Program), hforden-Smi thsonian Expedition seum of Aatiital Histor), Smithsonian Institution, Washang- to Dominica, National Geographic Society, and the ton, D. C. 20160. Smithsonian Research Foundat ion. I 2 SSIITHSONIAPi COSTRIBUTIONS TO BOTANY Morphology chyna that lack cilia but may have a very dense rhizinal mat below. The mat may assume a mar- THALISISSTRLCTURE.-Pa?-melina basically has a ginal position and resemble cilia. These “cilia,” closely appressed to adnate foliose thallus with nar- however, will be dichotomously branched. Such a row lobes (1-4 mm wide). Most species have more distinction fails in many specimens of H. rcvoluta or less subirregular lobation and apically rotund (Floerke) Hale having rather sparsely branched lobes (Figures I&, 18b), although narrow lobed rhizines and a few marginally positioned “cilia.” If species are sublinear (Figures 12d, 16c,). There is, incorrectly interpreted as having furcate rhizines however, a wide range of variation in lobe config- and sparse marginal cilia, such a lichen might be uration and width, much more than exists, for ex- identified as Parmelina cryptochlom, which has ample, in Hypotmchynn (Hale, 1975a). more powdery capitate soralia, or as pustulate P. The internal anatomy is similar to that of other spiimosn, which has a pale yellowish medulla. All epicorticate genera (Hale, 1973a). A thin, generally of these species have gvrophoric acid. palisade plectenchymatous cortex is overlain by a One further example of parallelism involving pored epicortex (Figures 1 and 2). The medulla Pame 1in a rl is sec t a and Hypo t ru c h y n a n eod issec t a consists of loosely packed hyphae, often encrusted (Hale) Hale can be mentioned. The latter has with lichen substances (Figure 2f). The lower cor- clearly dichotomously branched rhizines and lacks tex is p”p1ectenchymatous (Figure 20-d). cilia, yet both contain the same chemical, gyropho- The upper surface is strongly white maculate in ric acid, and have virtually identical lobe configu- Pwmelina con.son (Figure 3a), P. miielleii, and P. ration. It is difficult, if not impossible, to decide at pilostr, less conspiciiously so in P. nzelanochaetn, P. this time whether intergeneric hybridization has pa.stillifei.a, P. qiiercina, and P. tiliacea, and faintly occurred in these cases and resulted in “hybrid” or not at all in the remaining species. species. The lower surface is black with only three ex- RHIzIms.-?’he rhiLines of Parmelina are of two ceptions. Pai.melina cnormis has a uniformly pale types: simple to sparsely furcate (Figure 3c) and brown lower surface, and P. expallida and P. versi- squarrose. Squarrose rhizines, that is, those with a forrnis are darker brown tending toward black at main axis and short lateral branches, are confined the center. mostly to some species in section Myelochroa. This CILIA.-The marginal cilia which characterize pattern is also known in a few species of Parmelia Pnrrnelina are usually distinct but may be variably (e.g., P. ,szilcata Taylor) and Pai.motl-ema (the P. dispersed around the lobe margins. In many species 7.cticnlOtzLm group). they occur more or less regularly both on lobe tips ISIDIAAND LosuLEs.-Isidia are normally cylin- and in the axils (Figures 3a, lia, 18b). In others drical and erect as in other parmelioid genera and cilia may be lacking at the lobe tips and confined occur in the following 13 species: P. antillensis, P. to the axils (Figure 3b) or may be so sparse and expallidla, P. di.csecta (Figure 34, P. horrescens, P. inconspicuous in a few species, such as P. .rimplicioy, jamesii, P. indica, P. lindmanii, P. melanochaeta, that they are overlooked. In these cases one might P. obsessn, P. perisidians, P. tiliacea, P. usambaren- tend to classify them in the genus Pseutlopaimelia sis, and P. urnllichinna. Isidia are distinctly lobulate (Hale, 1976a), which is differentiated from Par- in P. degelii ant1 P. spatlziilata (Figure 44 and melina, among other ways, by the total absence of uniquely peltate in P. pastillifem (Figures 3e, 4b), a marginal cilia. Great care must be taken to estab- close relative of P. tiliacea, Apical cilia almost al- lish the presence or absence of cilia in the lobe ways occur in P. howescens and P. melanochaeta axils. Another example of this problem is Pa?-melin(i (Figure 4c) and resemble those in PaYmotTema crin i- endoleum and P.rezidopnrmelin siibtiliacea (Nylan- tiinz (Acliarius) Choisy. der) Hale, two very similar species in Australia that Lobules not originating from isidia are

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