SMITHSONIAN CONTRIBUTIONS TO NUMBER 33

A Monograph of the Hale ()

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 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 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. 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 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 found in differ in the production of cilia. Both have fatty P. schindle7.i and P. xantholepis (Figure 4e,f); they acids and cannot be separated easily by a chemical are mostly marginal. test. PvsTuLEs.-These are characteristic of four species, Confusion may also arise with species of Hypot~a- P. Iiaynchinensis, P. leucotylizn (Figure 5g), P. NUMBER 33 3

FIGURE1.-Epicortical pores oti the surface of Parnze[irzn species: a,b, P. ettornzis (X 400 and 1600) (Jeilicoe 150 in US); c,d, P. muelleri (X 400 and 1600) (Hale 42219); e,!, P. indica (X 400 atid 1600) (Hale 43884). (Scatitiing-electron microphotographs.) 4 SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE2.-Internal structure of Parmelina; a, vertical cross section of P. degelii (Degelius in US) (x 800); b, upper cortex of P. degelii (same section as in a) (x 1600); c, vertical cross sec- tion of P. galbina (Hale 23415) (x 800); d, upper cortex of P. galbina (same section as d) (x 1600); e, epicortex (arrow) of P. pastillifera (Schroppel 143) (x 5000); f, crystals of lecanoric acid on hyphae of P. pastillifera (Schroppel 143) (x 4000). NUMBER 33 5

FIGURE3.-Morphological structures of Parmelina: a, white maculae and marginal cilia of P. consors (Malme 1282 in US) (x 10); b, axillary cilia of P. lindmanii (Nee and Mori 4258 in US) (x 10); c, rhizines of P. galbina (Hale 18930) (x 70 with scanning-electron microscope); d, isidia of P. dissecta (Montes 10121 pro parte) (x 90 with scanning-electron microscope): e, peltate isidia of P. pastillifera (Schroppel 143) (x 90 with scanning electron microscope): f, cross-section of a peltate isidium of P. pastillifera (Schroppel 143) (x 400 with scanning- electron microscope). 6 SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE4.-Morphological structures of Parwelina: a, isidia of P. dissecta (Hale 33375) (x 10); b, peltate isidia of P. pastillifera (Schroppel 143) (x 10); c, ciliate isidia of P. horrescens (Imshaug 22174) (X 10); d, procumbetit isidia of P. spathulata (Koefler, in US) (x 10); e, lobules of P. schindleri (Schindler 4569 in US) (x 10); f, 1ol)ules of P. xantholepis (Hale 25993) (x 10). NUMBER 33 7

FIGURE5.-Morphological structures of Parrnelina: a, soralia of P. cc~irulenta(Weber anti McYean L-51280 in US) x 10); b, soralia of P. eryptochlora (Hale 35357) (x 10); c, pustules of P. su0- fatiTcens (isotjpe in US) (x 10); tl, coarse coralia of P. swinsco7~'ii(Swinscow K 31/13 in US) (x 10); e, pustules of P. spiiniosa (Hale 19897) (x 10): f, pustule of P. ~piiniosa (Hale 19897) (x 100 with scanning-electron microscope); g, pustules of P. leitcotyliza (Hale 29.544) (x 10). 8 SMITHSONIAN CONTRIBUTIONS TO BOTANY spumosa (Figure 5e,f), and P. szibfatiscens (Figure pm. The P. aurzilenta-P. subaurulenta group (sec- 5c). Soredia are not produced on these pustules. tion Myelochroa) has spores 5-10 X 7-15 pm. The Parmelina denegans, on the other hand, is clearly largest spores have been measured in P. versiformis pustulate initially but produces coarse soredia at (10-14 X 18-28 pm), but no other species in the maturity. genus have spores greater than 19 pm long. The SOREDIA.-SOralia are produced in seven species of smallest spores (3-6 X 7-8 pm) occur in the two Parmelina: P. auwlenta, P. cryptochlora, P. dene- anamolous norstictic acid-containing species, P. guns, P. metawvoluta, P. muelleri, P. pilosa, and antillensis and P. phlyctina. P. swinscowii (Figures 5a,b,d, 17b, 18b). They are orbicular and lamina1 or subterminal but never Chemistry marginal and linear as in Parmotrema. The remaining 17 species of Parmelina produce The chemistry of Parmelina species was de- no vegetative propagules, although the very termined with microcrystal tests before 1965 (Hale wrinkled and often fragile upper cortex of P. and Kurokawa, 1964) and later with thin-layer entotheiochroa or P. rhytidodes might be mistaken chromatography (Merck F-254 silica gel pre-coated for pustules (Figure 6). plates). Two solvent systems, hexane-ether-formic APoTHEcIA.-The apothecia are typically adnate acid and benzene-dioxane-acetic acid, have been or sessile and less than 5 mm in diameter, though routinely employed using C. Culberson’s methods Pal-nzelina irrugniis has discs up to 10 mm broad. (1972). I have had the benefit of Dr. Culberson’s The disc is occasionally perforate in P. co1zsors. No advice on the identification of many of the lichen species has a coronate rim, and only two, P. qzier- substances, and she has very kindly communicated cina and P. tiliacea, sometimes have retrorse to me the results of her work on the “horrescens” rhizines on the lower part of the amphithecium. unknown. It should be assumed, incidentally, that, No fertile collections are known as yet for P. unless otherwise stated, the components of each ci.yptochlora, P. degelii, P. hayachinensis, P. iradica, species in tlie taxonomic section refer to the liolo- P. jamesii, P. spathulatcl, or P. ~winscowi~. type or lectotype specimens with additional com- Spore siLe is very uniform within a given species ments on tlie results from other specimens. and not greatly different between unrelated species. Before listing the lichen substances discovered For example, the species in the P. dissecta-P. hor- so far in Pal-melinn, I might point out a most re- wscens group (Figure 9) have spores 8-12 X 12-18 markable feature in its chemical profile. Orcinol

FIGURE6,-MorpholoLgy of Parmelina: a, flaking cortex of P. entotheiochroa (Hale 29.534) (x 4); b, wrinkles of P. rhytidodes (Numnriri 156 in US) (x 10). NUMBER 33 9 depsides (except for gyrophoric and lecanoric acids), acid. The spots react with H,SO, the same as meta-depsides, orcinol depsidones (except for rare orcinol depsides and fall close to gyrophoric and lobaric acid), and the p-orcinol depsides related to lecanoric acids in the usual solvent systems. The barbatic acid are completely lacking. Two other best separation is achieved in benzene-dioxane cortical substances, lichexanthone and usnic acid, where a single large spot, the main “horrescens” are also missing. The general impression is that unknown, falls well above gyrophoric acid. This Parmelina has a relatively “primitive” assemblage unknown has now been detected in P. damaziana, of substances and would fall on a lower evolution- P. horrescens, P. schindleri, and P. subfatiscens. I ary scale than Hypotrachyna, Parmotrerna, and have also recently discovered this substance in an Pseudopamelia but significantly higher than Bzil- undescribed species of Hypoti.achyna. bothrix, Painzelia, and Relicina (Hale, 1967). The widespread medullary component lecanoric ALIPHATICL\cIDs.-The identification of fatty acid occurs in P. pastillifem, P. pnrinata, P. qiier- acids presents many problems and no entirely satis- cina, and P. tiliacea. I+. falls Close to gyrophoric factory techniques have been worked out to deal acid on chromatographic plates antl must be care- with them. The most direct method is to spray the fully distinguished from it by running controls. chromatographic plates ivith water and carefully ?’he toluene-acetic acid solvent system should be watch for opaque, white, water-repellant spots as the used in addition to the other t\vo systems. plate dries. These are the fatty acids. Terpenes have DEPsmosiis.-?’liese substances occur in many a similar reaction but differ in turning bluish, pur- lichen genera but are at best s~~oradicallydeveloped ple, or I~roivnwhen sprayed with H,SO, and heated. in Paimel ina. Pliy logene t icall y “primitive” salazinic Dr. AIyles Keogh of the Universitlatl de 10s Andes, acid and its close acetate galbinic acid predominate hlerida, Venefuela, has also informed me (pers. in section Pai.melina. comm.) that the fatty acids turn brownish when Constictic acid is an accessory substance with heated for 12 hours or more at 100°C (in contrast stictic acid ancl salazinic acid (see below). to the 10 minutes usually needed for development Fumarprotocetraric acid is extremely rare in the of phenolic substances). In any event, protoliches- genus, occurring in only one species, the Aus- terinic acid has been identified in P. expallida tralian-Sew Zealantl endemic P. jnmesii. It is ac- (Kurokawa, 1968a), and unidentified fatty acids companied by protocetraric acid. Succinprotoce- are the principle components in P. endolezica and traric acid is lacking. P. !let ei.ochloti. Galbinic acid is confined to a small group of hPsmEs.-These well-known substances are com- very closely related species: P. grilbincr, P. haynchi- mon in section Pamwlina (those species of Par- nensi,~,P. nic/awvolufa,arid P. ob.scsscr, accompanied melina lacking terpenes) and occur in most of the by secalonic acid X and the ”s~~baui~i~le~ita”terpene pantemperate-montane pantropical species. series. It is easily identified as ;I deep H,SO, orange Atranorin is the primary cortical substance found spot below norstictic acid in benzene-tlioxane. in all species of Paymelina. Lobaric acid is the only orcinol tlepsidone in Gyrophoric acid, a tridepside, occurs in Paimelinn Pni.melinci and occurs as an accessory substance cryptochlora, P. dissecta, P. melanochaeta, P. spnthir- with salwinic acid in P. su~inscou~ii. hta, and P. spztrnosu. It is a probable minor com- Korstictic acid is the main component in two ponent in P. tlninnziana, P. horl-escens, P. .schindlei~i, closely related Caribbean species, P. antillensis and ancl P. subfritiscen.s along with the “horrescens” un- P. phlyctina. A lower spot of the, as yet, unidenti- known. Gyrophoric acid is not easily chromato- fied connorstictic acid may be present. graphed because of the tendency for streaking antl Protocetraric acid occurs only in P. jnmesii with variable R, values that are highly influenced by fumarprotocetraric acid (see above). solvent composition. In addition, one or more as- Salazinic acid, a very common substance in the sociated spots, still unidentified, may appear on Parmeliaceae, occurs in P. enotmis, P. .rimplicior, the plates. P. ,\ ZL’ in.sc ow i i, P. 1 I sa ?nhave n.c is, P. ve~sif 07-rn is, ant1 “Horrescens” unknown is actually several sub- P. zuallichinna. It is a rare accessory with leucotylin stances presumed to be depsides related to lecanoric in P. el-assata antl may be accompanied by constictic 10 S\IITHSONIAS CONTRIBUTIONS TO BOTANY acid in P. metarevoluta. All of these species, ex- remains to be seen whether this variation has any cepting P. versiformis, are distributed in the Old taxonomic value. World following a pattern seen in Bulbothrix Zeorin is a well known lichen substance. The (Hale, 1976b). stereochemistry of its hopane skeleton was only re- Stictic acid has been discovered in only one spe- cently studied by Yosioka et al. (1968b). It occurs in cies, P. mnelleri, where it is accompanied by con- all species in section 1\4pelochroa and has been re- stictic acid and two unknown compounds falling ported in Acroscyplirrs, Cladonin spp., Hppotra- between stictic and constictic acids in benezene- chpa majoris (Vainio) Hale, Lecanom, Sephroma, dioxane. Peliigera, and the Physciaceae. It may be identified TERPENEL-A closely related series of triterpenes as the highest blue spot on the chromatographic with a liopane skeleton have been described for plates in Iiotli solvent systems (Figure 7). species in section Myelochroa. Their structures have Prc~iE.\’Ts.-.\Iedullary pigments are especially been determined by Yosioka and his group in Ja- characteristic of species in section hfyelochroa. The pan using nuclear magnetic resonance (NhIR) chromatography of these is extremely difficult and spectra, mass spectrometry, etc. The tliin-layer ~is~iallpunsuccessful. There is considerable streak- chromatography is not yet standardized or well un- ing and individual components cannot be distin- derstood, and I have not tried to identify individual guished. spots. Typical profiles for the terpene-containing Secalonic acid X was first ideniifietl by Yosioka species of Parmelina are illustrated in Figure 7. et al. (1968a:2090), who established its identity with Leucotylic acid, a colorless compound closely entothein. It also occurs in ergot (Shibata et al., related to both zeorin and leucotylin, was first elu- 1964). Yosioka found secalonic acid A in Parmelina cidated by Yosioka (1966). It was isolated from Pa?.- nii~rilcntm, P. enfotheiochroa, P. pel-isiclians, and melina leucotyliza, although my results indicate that P. subauixlenta, and I presume it is the major leucotylin is, in fact, the main component of this pigment in all species of section ilfpelochroa with species. I suspect that their material included a a yellowish orange medulla, as well as in P. immis- mixture of P. crassata, which does contain leuco- cens and P. lintlmanii, both lacking terpenes. Un- tylic acid. The species of Pnrmelina which contain identified reddish pigments accompany secalonic leucotylic acid (or at least have the same terpenic acid 4 in the lower medullary layer of P. amagien- profile as P. airrulentn, as illustrated in Figure 7) ,pis and P. dcnegans. are P. aurzilenta, P. degelii, P. il-rugans, and P. rhytidodes. Phytogeography and Speciation Leucotylin, the progenitor of leucotylic acid, is the main terpene component in P. amagiensis, P. Pawnelinti occurs primarily on trees in secondary crassata, P. denegans, P. entotheiochroa, P. galbina, forests in lemperate zones and at higher elevations P. hayachinensis, P. indica, P. leucotyliza, P. meta- in the tropics. A number of the commoner species revoluta, P. obsessa, P. perisidians, P. subaurulenta, may also occur on rocks. There are, however, only and P. xantholepis. It forms a low major spot in four oliligately saxicolous species, P. enownis, P. indica, P. obsessa, and P. usambaiensi.r. The Pnr- benzene-dioxane, but I have generally identified it melina of various geopolitical units are as a profile of several terpene spots as illustrated in enumerated below. It is altogether obvious that Figure 7. many countries are underrepresented because they Yosioka (Yosioka and Nakanislii, 1963; Yosioka have not been visited by lichen collectors. and Nakanishi, 1966) has isolated at least five other compounds related to leucotylin in “Parmelia ento- XORTHAMERIC.~ theiochroa.” I presume that at least some of these United States: P. antillensis, P. aurulenta, P. dissecta, P. make up the numerous spots resolved in hexane- galbi17n, P. hoi 1 escciis, P. ~ne/ar~~volii/a,P. obsessa, and P. ether (see Figure 7),but no attempt has been made sfiu in osa. to identify any of them on the plates. There seems to be great variation in the intensity of the spots, MEXICOALD C~NTRALAMERIC~ reflecting different concentrations in the thallus. It Mexico: P. antillensis, P. aurulenta, P. dissecta, P. hor- NUMBER 33 11 rescens, P. ivztniscens, P. Zindnlanii, P. niuelieri, and P. WEITISDIE s spumosa. Guatemala: P. horrescens. Cuba: P. antiliensis, P. dissecta, P. horrescens, and P. Panama: P. dissect(/, P. Izorre,ce,ls, and P. subfutiscens. pizlyctinn.

FIGURE7.-Chromatographic profiles of Pnrnielinu species: profiles in the hexane-ether solvent system (top) and benzene-dioxaiie solvent system (bottom). (1 1 P. uicndenla, 2 = P. degelii, 3 = P. irrugans, 4 = P. rhytidot/u.i, 5 = P. ci)nrrgie!i,iis, G = P. ci-nssata, 7 = P. denegnns, 8 = P. entotheiochroa, 9 = P. indlica, 10 = P. iettcotyliza, 11 = P. perisidians, 12 = P. subaurulentn, 13 = P. santholepis, 14 = P. plbi!rcr, 1.5 = P. tnetarevoltrtrr, lG = P. ohsessa, Z = zeorin spot, G = galbinic acid, S = salarinic acid, and C = constictic acid: leucotylic acid is presumed to be the series of spots in 1 to 4 just below rerorin, and leucotylin is the lowest series in 5-10 in benzene-dioxane; atranorin falls just above the point where the photographs were trimmed.) SAIITHSONIAN CONTRIBUTIONS TO BOTANY

Jamaica: P. antillensis, P. dissecta, P. horrescens, P. subaurulenta, P. tiliacea, P. wallichiana, and P. xantholepis. phlyctina, P. spumosa, and P. subfatiscens. Sri Lanka: P. denegans, P. dissecta, P. perisidians, and P. Hispaniola: P. antillensis, P. dissecta, P. horrescens, and subaurulenta. P. phlyctina. Thailand: P. expallida, P. perisidians, P. usambarensis, P. Puerto Rico: P. phlyctina. wallichiana, and P. xantholepis. Lesser Antilles (including Trinidad): P. anlillensis, P. Indochina: P. aurulenta. cryptochlora, P. dissecta, and P. subfatiscens. China (including Hong Kong): P. aurulenta, P. irrugans, P. quercina, P. subaurulenta, and P. wallichiana. SOUTHAMERICA Korea: P. entotheiochoroa. Japan: P. amagiensis, P. aurulenta, P. crassata, P. dissecta, Colombia: P. lindmanii, P. melanochaeta, and P. spumosa. P. entotheiochroa, P. galbiria, P. hayachiizensis, P. horrescens, Venezuela: P. antillensis, P. dissecta, P. horrescens, P. P. irrugans, P. leucotyliza, P. metareuoluta, P. perisidians, P. lindmanii, P. muelleri, and P. spumosa. quercina, P. rhytidodes, P. spumosa, and P. wallichiana. Ecuador: P. pilosa. Taiwan: P. azcrulenta, P. dissecta, P. expallida, P. horres- Peru: P. muelleri. cens, P. spirmosa, P. subaurulenta, and P. wallichiana. Brazil: P. antillensis, P. aurulenta, P. consom, P. dama- Philippines: P. aurulenta, P. dissecta, P. horrescens, P. ziana, P. dissecta, P. heteroloba, P. lindmanii, P. melano- perisidians, P. wallichiana, and P. xantholepis. chaeta, P. muelleri, P. schindleri, P. spumosa, and P. Malaya: P. wallichiana. wersiformis. Indonesia: P. aurulenta, P. dissecta, P. horrescens, P. spu- Uruguay: P. cons or,^, P. horrescens, P. lindmanii, and P. mom, P. wallichiana, and P. xantholepis. pilosa . Sabah: P. denegans, P. leucotyliza, and P. wallichiana. Paraguay: P. consors, P. lindmanii, and P. melanochaeta. New Guinea: P. aurulenta. Argentina: P. consors, P. lindrnanii, P. muelleri, P. pilosa, Australia (including New Zealand): P. degelii, P. endo- and P. uersiformis. leuca, P. horrescens, P. jamesii, P. pruinata, P. qziercina, and Chile: P. horrescens, P. pilosa, P. spumosa, and P. swins- P. spumosa. cowii. Hawaiian Islands: P. aurulenta.

EUROPE To summarize, t.he New World has a total flora of 20 species, 12 of them endemic, 3 (P. pilosa, P. Western Europe: P. dissecta, P. horrescens, P. pastillifera, P. quercina, and P. tiliacea. subfatiscens, and P. swinscowii) shared with Africa, Russia: P. aurulenta, P. qziercina, and P. tiliacea. and the remaining 5 (P. aurzilenta, P. dissecta, P. Israel: P. tiliacea. horrescens, P. quercina, and P. spumosa) essentially pantemperate or montane pantropical. AFRIC4 Europe has a small Paimelina flora. Dahl and Morocco: P. tiliacea. Krog (1973) list only P. tiliacea in Scandinavia. The Tunisia. P. tiliacea. checklist by James (1965) for Great Britain includes Ivory Coast: P. usambarensis and P. wallichiana. P. dissecta, P. horrescens, P. qziercina, and P. Guinea: P. usambarensis and P. wallichiana. tiliacen. For the rest of Europe, Poelt (1969) adds Uganda: P. usambarensis and P. wallichiana. Kenya: P. dissecta, P. pilosa, P. swinscowii, and P. wallichi- only P. caipoirhizans (= P. qziercina) and P. pastilli- ana. fera. Angola: P. wallichiana. Africa has only two endemic species, P. enormis Rhodesia: P. wallichiava. and P. spnthdatn, and nine other species either Zambia: P. enormis. shared with the Americas, Asia, or pantemperate in Tanzania: P. aurulenta, P. dissecta, P. usambarensis, and P. wallichiana. distribution. Union of South Africa (including Swaziland): P. dissecta, Asia is especially rich in Parmelina with 15 spe- P. horrescens, P. pilosa, P. spathulata, P. spumosa, P. sub- cies on the Indian subcontinent and 16 in Japan fatiscens, and P. wallichiana. and eastern Asia. Tropical Southeast Asia has nine Madagascar-RPunion: P. aurulenta, P. spumosa, and P. species, none of them endemic and all occurring wallichiana. in the higher elevation cloud forests. The genus is not represented at all in the lowland dipterocarp ASIAAND PACIFICAREAS rain forests from Sri Lanka to Indonesia. Pakistan: P. aurulenta, P. quercina, and P. tiliacea. The Australia-New Zealand region has seven India (including Nepal): P. aurulenta, P. cryptochlora, P. denegans, P. dissecta, P. expallida, P. horrescens, P. indica, species, of which four, P. degelii, P. endoleuca, P. P. perisidians, P. rhytidodes, P. simplicior, P. spumosa, P. jamesii, and P. pruinata, are endemic. No Parme- NUMBER 33 13

FIGURE8.-Number of species in section Myeloclztm in each major geographical division.

linae have been collected on the Pacific Islands ex- (1972) also regards P. aimilenla as a Tertiary relict. cept for P. cl~irttlenta at higher elevations in Ha- Pai.melina has several complex groups of species waii. The total Old World flora (Africa, Asia, and that have evidently resulted from combined chem- Australia-New Zealantl) comprises 26 species. ical and morphological evolution. The species in One distrihution pattern of considerable phyto- section iMyelochroa, for example, all of which con- geographic interest stands out. The species of sec- tain terpenes and almost always have secalonic tion Myeloch ma, a rather homogeneous group, are acid A, a yellow pigment, have evolved from two concentrated in India and eastern Asia (Figure 8) chemically dissimilar ancestors, one, the smaller and only one of them, P. aurulentn, occurs in all group, with leucotylic acid and resembling P. ir- geographic areas (Figure 10). The species in section g.iigans, and the other, a larger series with leuco- Pasmelina, on the other hand, are not strongly tylin, resembling the present-day P. subazirzilenfa. Concentrated in any region. The numerous morplis derived from these presump- TVithin section Myelochl-oa the species related to tive parents or their ancestors, P. nmagiensis, P. P. galbina form a particularly interesting group. As aiti.itlenta, P. crassata, P. clegelii, P. denegnns, P. pointed out by Kurokawa (1972), P. galbina, the entotheiochroa, P. leucotyliza, P. perisidians, P. fertile progenitor species, behaves as an Arcto- Thytidodes, and P. xnntholcpis, have diversified tertiary plant which migrated southward and has mostly by production of soretlia, isidia, pustules, survived in Japan and eastern Korth America. A and wrinkles, variation in apotliecial diameter, etc. rare sorediate morph, P. metawvoluta, has the AIorph forniation (see Hale, 1975a:13) is not same range, whereas the isidiate morph, P. ob.tessa, strongly delineated for the most part since the par- is restricted to North America, and the pustulate ents cannot be traced directly. \Ve are left to con- morph, P. hayachinensis, to Japan. W. Culberson clude that section Alyelochron is an ancient group 14 SMITHSONIAN CONTRIBUTIONS TO BOTANY evolved over a long period. The P. galbina group, Hale and Kurokawa (1964: 130) grouped the narrow- as mentioned above, forms the only exception to lobed, marginally ciliate Parrneliae in Parmelia this rule. subgenus Parmelia section Imbricaria (Schreber) E. ILIorph formation has also been an important Fries, and it is essentially this group that I have seg- mode of speciation in section Parmelina. The fol- regated as a distinct genus, Parmelina (Hale, 1974: lowing parent morph-vegetative morph series are 482). indisputable: P. consors-P. pilosa (sorediate), P. Parmelina is a rather heterogeneous assemblage of quercina-P. tiliacea (isidiate), and P. imrniscens-P. 47 species differing in lobe width, production of lindmanii (isidiate). Parmelina phlyctina and cilia, pigmentation, and rhizine branching. Two isidiate P. antillensis are extremely close but less major groups can be recognized, one including clear-cut morphs. A large complex of species have those species lacking terpenes and one with terpene- been derived from now extinct parents in the P. containing species. They may be conveniently re- dissecta-P. horrescens group, as illustrated in Figure garded as sections. 9. Like the P. subaurulenta group of section Mye- lochroa, the parent morphs of these numerous vegetative morphs are either extinct or have yet to Section Parmelina be discovered. The remaining species in the genus appear to Parmelia section Hypotrachyna subsection Myelolezica Asa- have no discernible interrelationships or morphs. hina, 1952:74 [type-species: Purrnelina tiliacea (Hoffmann) Hale]. These include P. endoleuca, P. enormis, P. expal- lida, P. heteroloba (part of the P. darnaziana se- The species in this section are characterized by ries?), P. pruinata, P. sirnplicior, P. swinscowii, P. the lack of any triterpenes. Except for two species, usambarensis, P. veisiformis, and P. wallichiana. P. immiscens and P. lindmanii, the medulla is white and unpigmented. Several internally homogeneous Classification of Parmelina groups can be recognized. For example, the P. dissecta-P. horrescens group (Figure 9) includes The taxon Lichen section Imbricaria was first seven species with very similar lobe configuration, proposed by Schreber (1791:767). He cited no spe- abundant marginal cilia, and gyrophoric acid or cies, only Squamaria Hoffmann, which is invalid as the closely related “horrescens” unknown. Heavily the later homonym of Squamah Ludwig (Phane- white-niaculate P. consow, P. pilosa, and P. rnuelleri, rogamae). Acharius (1794:250) adopted the name as having coarse, leathery thalli and thick, furcate a tribe of Lichen and included in it L. olivaceus L. marginal cilia, are obviously related. and L. tiliaceus Hoffmann, as well as several other The predominantly European complex of spe- species now recognized as belonging to Hypogymnta, cies, P. qztcrcina-P. iiliacea-P. pastillifera, contains Parmeliopsis, Physcia, and Xanthoria. XIichaux lecanoric acid and forms an easily recognized but (1803) raised the name to generic rank and cited isolated group. Parmelina antillensis and P. phlyc- one species, a lichen now called Anzia colpodes tina, both unusual in containing norstictic acid, (Acharius) Stizenberger. Fries (1825:242) later trans- have no close relatives in the genus. Two other ferred it to Parmelia as Parmelia section Imbricaria, New World species, P. immiscens and P. lindmanii and Koerber (1855:68) used it at the generic rank, with a yellow medulla, form an isolated branch of even though he realized that the name was a later this section. homonym of Imbricaria Decandolle (Phaneroga- The species with salazinic acid, P. enormis, P. mae). Koerber and contemporary lichenologists simplicior, P. swinscowii, P. usambarensis, P. versi- placed many species that we now consider parme- formis, and P. wallichiana, have little in common lioid in Imbricaria, while at the same time using except their chemistry. Finally, the remaining spe- Parmelia for many species now placed in Physcia. cies, P. endoleuca, P. expallida, P. heteioloba, P. After the delimitation of Parrnelia in a modern jamesii, and P. pririnata, have no obvious common sense by authors such as Sluller and Nylander, ancestry or affinities with other species in the sec- Imbricaria was relegated to synonymy under it. tion. NUMBER 33 15

Progenitor

ProgenitorI- Progenitor (gyrophoric acid) (“horrescens” unkn.)

P. horrescensI P. schindleri P. sirbfutiscens Progenitor (whiteProgenitor rnedullaj (isidiate) (lobulate) (pustulate) (yellow medulla) ,I P. spir ntosu (pustulate) r-l-l P. dissei.tu P. sputhulatu P. cryptochloru (isidiate) (lobulate-isidiate) (sorediate)

FIGURE9.-Hypothetical derivation of species in the Parmelina dissecta-P. horrescens group. hrot shoivn are P. damaziana, a presumptile nonisidiate, nonsorediate derivative of the “hor- rescens” progenitor, and P. melanochaetn, a possible relative of P. dissecta but derived from a different progenitor.

Section Myelochroa (Asahina) Hale, new status tyliza, P. perisidzans, P. subauiulenta, and P. xan- tholepis) with leucotylin. Parmelina indica can Parmelia section Hypotrachyna subsection Myelochroa Asa- probably be classified in the P. szi baurulenta sub- hina, 1952374 [type-species: Parmelina aurdenta (Tucker- group although it has a small saxicolous thallus man) Hale]. and lacks secalonic acid A. The affinities of Parmelina lie primarily with All of the species in this section contain zeorin Parmotrema Alassalongo, a broader lobed, loosely and either leucotylin or leucotylic acid and asso- attached, often ciliate genus (Hale, 1965). Palme- ciated terpenes, and all, excepting P. indica, produce Zina, however, has a far less evolved chemistry, nar- varying amounts of the yellow-orange pigment rower adnate lobes, consistently small spores, and secalonic acid A and possibly other, as yet, unidenti- adnate or sessile, usually imperforate apothecia. fied pigments. This group is highly restricted to Where soredia are produced, they are lamina1 Asia, and the 17 species have very close affinities, in rather than marginal in contrast to most Parmo- sharp contrast to section Parmelina. trema species. All in all, however, the apparent Two apparently unrelated groups are recognira- intergradation between broad-lobed Parmelinae and ble in the section. One, the P. galbina group (P. gal- the smaller lobed species of Parmot?-emu,involving bina, P. hayachinensis, P. metarevoluta, and P. ob- perhaps six or eight species, poses difficult problems sessa), is characterized by galbinic acid in addition in the correct generic identification of individual to leucotylin and secalonic acid A. The other group specimens and for the “lumper,” at least, raises includes two closely related subgroups, the P. auru- questions on the distinctness of the two genera as lenta subgroup (P. aul-ulenta, P. degelii, P. irru- circumscribed here. I do not expect that the prob- guns, and P. rhytidodes) with leucotylic acid, and lem can be resolved to the satisfaction of all lichen- the P. subazirulenta subgroup (P. amagiensis, P. ologists, but the basic differences should become crassata, P. denegans, P. entotheiochroa, P. leuco- clearer as more collections are made in tropical 16 SMITHSONIAN CONTRIBUTIONS TO BOTANY countries and our knowledge of soredial formation, species lacking isidia, soredia, and pustules; (2) significance of chemical variation, etc., broadens. those with isidia, lobulate isidia, or lobules; and The key below is divided into three sections: (1) (3) those with soredia or pustules.

Key to the Species of Parmelina

1. THALLUSLACKING ISIDIA, SOREDIA, AND PUSTULES (Cortex flaking off in Parmelina entothetochroa and P. phlyctina, and heavily rugose in P. crassata and P. rlzytidodes)

1. Medulla more or less completely yellow to yellow-orange (pigmented mostly below the apothecia in P. galbina). 2. Upper surface hea\ily rugose (Figure 6b) 35. P. rhytidodes 2. Upper surface more or less plane. 3. Upper cortex vei) fragile, flaking awa) over extensive areas (Figure 6a) . 4. Medulla deep yellow 13. P. entotheiochroa 4. Medulla white (turning reddish if impropeily curated) 3 1, P. phylctina 3. Upper cortex entire and continuous. 5. Lower half of medulla darker reddish orange 1. P. amagiensis 5. Loner half of medulla not darkly pigmented. 6. Lobes narrow and sublinear, 1-2 mm wide 15. P. galbina 6. Lobes broader, subirregular, 2-4 mm wide. 7. Collected in Mexico 19. P. immiscens 7. Collected in Asia. 8. Apothecia 3 mm or less in diameter 40. P. subaurulenta 8. Apothecia 3-10 mm in diameter. 9. Thallus coarse, about 250 pm thick: leucotylin present 5. P. crassoto 9. Thallus tnembranous, about 150 pm thick: IeuLotylic acid present 21. P. irrugans 1. Medulla entirely white. 10. Lower surface pale brown. 11. Collected on rocks in Africa 12. P. enormis 11. Collected in treer in South America 45. P. versiformis 10. Lone1 surface black. 12. Medulla K+ yellow turning red. 13. Lobes sublinear, 1-2 mm wide 15. P. galbina 13. Lobes broader, 2-5 mm wide. 14. Upper cortex fragile, flaking off 31. P. phlyctina 14. Upper cortex firm, continuous. 15. Collected in India 37. P. simplicior 15. Collected in South America 45. P. versiformis 12. Medulla K negatike. 16. Medulla Cf red. 17. Cortex distinctly white maculate 34. P. quercina 17. Cortex emaculate 33. P. pruinnta 16. Medulla C-. 18. Thallus coriaceous, heavily whlte-maculate; lobes 2-5 mm jvide 4. P. consors 18. Thallus thin, emaculate: lobes 1-3 mm i\ide. 19. Medulla KCf rose; “horrescens” unknor\n present 7. P. damaziana 19. Medulla KC-: fatty acids present. 20. Collected in Australia 11. P. endoleuca 20. Collected in South America 17. P. heteroloba

11. THALLUSISIDIATE, LOBULATE-ISIDIATE, OR LOBULATE

I. Medulla yellow to pale orange-yellow. NUMBER 33 15

2. Thallu\ densely lobulate uithout formation of isidial initials (Figure 4f) 47. P. xantholepis 2. Thallus noimally isidiate or lobulate-isidlate. 3. Isidia large, doisiientral and lobulate 8. P. degelii 3. Isidia normal, cylindrical and erect. 4. Lobes subirregular and apicall) rotund, 3-6 mm wide 24. P. lindmanii 4. Lobes sublinear, 1-2 mni Ttide. 5. Plants saxicolous (rarel) coi ticolous) 111 eastetn Sorth America 28. P. obsessa 5. Plant5 corticolous in Southeast A5ia 30. P. perisidians 1. Medulla M hite. 6. Thallus densel) lobulate I\ ithout isidial initlals (Fignre 4e) 36. P. schindleri 6. Thallus isidiate 01 lobulate-isidiate. 7. lsidia in part ptocumbent and lobulate (Fignre 4c,rl). 8. Medulla C+ rose 38. P. qpathulata 8. Medulla C- 18. P. hori escens 7. lsitlia erect, cylintliical. 9. Medulla Kf jellou turning red. 10. Collected in tiopical Ameiica, norstictic acid piesent 2. P. antillensis 10. Collected in Asia and Aflica; salazinic acid present. 11. Lobes subirregulal, cioitded, apicall) iotund 46. P. urallichiana 1 I. Lobes bublineai, looselp dii aricate 44. P. usambarensis 9. Medulla I(- or faintly )elloit. 12. Medulla Cf rose 01 led. 13. Isidia apically flattened and peltate 29. P. pastillifera 13. Isidia cylindiical, not peltate. 14. Medulla C+ intense red (lecanoric acid) 43. P. tiliacea 14. Medulla C+ lose (g)iophoiic acid). 15. Lobes narrow, 1-2 mni nide; upper surface emaculate: isidia eciliate 10. P. dissecta 15. Lobes broader, 24 mm itide; upper surface maculate; isidia ciliate 25. P. melanochaeta 12 4iedulln C-. 16. Medulla P+ red 22. P. jamesii 16. Medulla P-. 17. Louei surface bronn to hlacLening in palt 14. P. expallida 17. Lower suiface unifoimly black. 18. Isidia apically ciliate; medulla KC+ rose 18, P. hort escens 18. Isidia eciliate; medulla KC- 20. P. indico

111. THALLUSSOREDIATE, SOREDIATE-PUSTULATE, OR PU5TULATE

1. Medulla entirely white. 2. Lobes broad and apicall) rotund; coi tex con

10. Pustules forming in capitate masscs (Figuie 5g) 23. P. Zeucotyliza 10. Pustules irregular, intermixed with wrinkles (Figure 6b) 35. P. rhytidodes 9. Lobes narrow and sublinear, less than 2 mm wide. 11. Medulla K+ reddish, P+ orange 16. P. hayachinensis 11. Medulla K-, P-, C+ rose, pigmented bery faint yellow 39. P. spumosa 8. Thallus sorediate or pustulate-sorediate at maturit). 12. Medulla deeper reddish orange In the lower half 9. P. denegans 12. Medulla uniforiiil) pigmented to nhite and pigmented only under the soralia. 13. Lobes 2-4 mm wide; medulla P- 3. P. aurulenta 13. Lobes less than 2 mm wide; medulla P+ orange 26. P. metarevoluta

Species Treatment the few specimens available, and does not seem to be a parent morph for the latter. The 47 species of Parmelina are arranged below in alphabetical order. Location of specimens listed SPPCIMENSEx \MIsFD.-Japan: Prov. Ettchu, Yanagisawa under “Specimens Examined” are indicated by the 599a (TNS); Prov. Hyuga, Kurokawa 50058 (TNS, US). standard acronyms except for those col- lected by Hale, all of which are preserved in the 2. Parmelina antillensis Smithsonian Institution (US). FIGURE11 b

1. Parmelina amagiensis, new combination Parmelirin antillensis (Nylander) Hale, 1974:482. Parmelia antillensis Nylander, 1868:264 [type collection: FIGUREIla Matouba, Guadeloupe, Husnot 445 (H. Nylander herbar- ium number 35119, lectotype; isolectotypes in G, P)]. Parnzelia arriagiensis Asahina, 1951a:228 [type collection: Mt. Parinelia blastica ’I’ainio, 1896:32 [type collection: Shawford Amagi, Prov. Izu, Japan, Asahina 95 (TNS, lectotype)]. Estate, Dominica, Elliott 899 (TUR, lectotype; isolectotype in BM)]. DEScRIPTIoN.-Thallus loosely adnate on bark or mosses over bark, buff mineral gray, 4-8 cm broad; DEscRIPTIoN.-Thallus adnate on bark, fragile lobes more or less subirregular, imbricate, 2-3 mm and membranous, up to 10 cm across, greenish min- wide, the marginal cilia numerous, simple, up to eral gray, often more or less white-pruinose at the 1 mm long; upper surface plane, white-maculate, lobe tips; lobes irregularly branched, subimbricate becoming heavily pycnidiate, isidia and soredia to convoluted, the marginal cilia sparse, mostly in lacking; medulla deep salmon orange; lower sur- lobe axils, to 0.5 mm long; upper surface shiny, face black, densely rhizinate, the rhizines black, plane, cortex continuous or cracked with age; simple or sparsely squarrosely branched, 0.5-1 .O mm densely isidiate, the isidia fine, to 0.4 mm high, long. Apothecia sessile to subpedicillate, 2-4 mm sparingly branched or simple; lower surface black, in diameter; spores 8, 6-7 X 9-1 1 pm. brown and shiny in a narrow zone along the mar- CHEMISTRY.-cOrteX K + yellow (atranorin); rne- gins, densely rhizinate, the rhizines simple. Apo- dulla more intensively yellow with color tests thecia adnate, 3-5 mm in diameter, the amphi- (zeorin, leucotylin and related “subaurulenta” ter- thecium isidiate; spores 8, 5-6 X 7-8 pm. penes, secalonic acid A, and unidentified pigments). CHEMISTRY.-cOrteX K + yellow (atranorin); rne- DIsTRIBuTIoN.-Japan. dulla K+ yellow turning red, C-, KC-, P+ REMARKS.-ThiS rare Japanese species is very orange (norstictic and connorstictic acids). close to P. denegans, a pustulate or sorediate- DIsTRIsuTIoN.-Caribbean region. pustulate species in tropical Asia. Both have a REMARKs.-Parmelia antillensis and its close Ca- more deeply pigmented medulla, especially in the ribbean relative P. phlyctina are the only two spe- lower half, than other members of section Myelo- cies in the genus containing norstictic acid. This chroa. I have not been able to resolve these pig trait, along with the rather broad, apically rotund ments with the usual chromatographic solvent lobes, might tempt one to place them in Parmo- systems. Parmelina amagiensis appears to be some- trema Massalongo, where, in fact, I had originally what more robust than P. denegans, judging from suggested they might belong (Hale, 1959). The NUMBER 33 19 lower surface, however, has at most a narrow bare Tschangscha, Hunan, China, Hnndel-Mazzetti 11454 (WU, or papillate zone, the lobes are generally adnate at lectotyie)]. Parnielia sihestris Degelius, 1940:47 [type collection: Togue the tips, not ascending, and the cilia are mostly in Pond near Mt. Katahdin, Maine, Degelius (Degelius her- the axils of the lobes. Both P. antillensis and P. barium [not seen]: isolectotype in US)]. phlyctinn share traits intermediate between Parme- Pnrmelia aurulenta var. silrwstris (Degelius) Degelius, 1941 :58. Zina and Parmotrema and have evolved in a very restricted region, the rain forests of the Caribbean DEscRIPrIoN.-Thall~is adnate on hark or rock, basin (excepting the unusual record of P. until- pale greenish mineral gray, 4-10 cni broad; lobes 1en.yi.r.from the southeastern United States discussed sublinear to subirregular, apically subrotund, 2-4 in Hale, 1971b:46). Other parmelioid groups have mni wide, the marginal cilia irregularly dispersed evolved here, eg., Pseudoparmelia martinicnnn but mostly in the lobe axils, to 0.8 mni long; upper (Nylander) Hale and P. raunkiaeri (Vainio) Hale surface shiny, plane to rugulose, soon sorediate to and perhaps also Relicinn eximbricnta (Gyelnik) i)iistLilate-sorecliate, the soralia up to 1 mm in diam- Hale (1975b, 1976a), but these are all lowland spe- eter, coalescing into large subcapitate clumps, the cies in dry, disturbed habitats. Although Parmelina soretlia coarse (Figure 5a); medulla white and be- ontillensis is often collected on citrus trees at 600- coming orange sulfur yellow only beneath soralia 700 m elevation, its primary habitat is the canopy antl near exposed cracks in the upper cortex or branches of trees in virgin rain forest. entirely sulfur yellow; lower surface black and moderately to densely rhizinate, the rhizines black, SrECI\rEss Ex4\rrXED.-L?nited States: , Yoshi- simple or becoming sparsely f urcate or squarrosely viiira et al. 660872 (US). Mexico: Chiapas, Hale 20238.’ Cuba: Oriente, Iinsliairg 250.51 (MSC). Jamaica: Cztlberson branched. Apothecia rare, adnate to substipitate, 13648, 13777, 13787 (DUKE), Znishaug 14088, 14627, 14943, the amphithecium sorediate-pListulate, 2-5 mm in 1.5321 (MSC), 14593, 15295 (MSC, US), Orcult 3551b, 5626b diameter; spores poorly developecl, 7 x 12 pm. (US), Plitt (US). Haiti: Oueit, Inishaug 22642 (MSC). Puerto CmxmmY.-Cortex K + yellow (atranorin); me- Rico: Stinisoti 1440 (DUKE). Guadeloupe: Culberson 14511 dulla negative with color reagents or if pigmented (DUKE), Oitestal 1916 (US). Martinique: Citlberson 14682 areas tested becoming more intensely yellow (zeorin, (DUKE), Uegeliuc (Degelius herbarium). Dominica: Irnshaug 32726 (MSC), Hale in Lichenes Selecti Exsiccati 942 (H, LD, leucotylic acid and related terpenes, antl secalonic US) (for additional records see Hale 1971a:lO). St. Lucia: acid A). Inishaug 29712 (MSC). St. Vincent: Znishaug 30504 (MSC). DIsTRIsvi-Io~.-Panteni~)er~~eantl montane pan- Grenada: Hale 38298, Inishaug 16092, 16335 (MSC). Trini- tropical (Figure 10). dad: Hale 38291, Inisliaug 32042 (MSC). Barbados: Elliott REMARKs.-Parmelina auritlenta is one of the most (TUR). \’enezuela: MC-rida, Hale 42751, 42941. Brazil: Minas Gerais, T’ninio in Lichenes Brasilienses Exsiccati 675 (TUR) widespread species in the genus and, at least in the (di7rributed as Parnielia a~tiazonica). United States, is one of the most commonly col- lected foliose . It is also unusual in being the only species of the subaurulenta complex that oc- 3. Parmelina aurulenta curs in the New World, perhaps suggesting merely that efficient dispersal by soredia is the prime reason for the pantemperate distribution. Parmelina aurulenta (‘Tuckerman) Hale, 1974:482. Soredial formation is extremely variable. The Parmelia auruletzta Tuckerman, 18583424 [type collection: Harpers Ferry, Virginia, Tuckernia?i (FH-Tuck, lectotype)]. lectotype, for example, is densely pustulate with Parmelia tiliacea var. eporescens Miiller Argoviensis, well developed soredia that obscure most of the 1887:316 [type collection: Siberia, Russia, Lahm 5 and 6 pustular initials (Figure 5a). At other times, the (G, lectotype: isolectotype in W)]. pustules become only sparsely and coarsely soredi- Pnrnzelia albido-strarninea Hue, 1899:161 [type collection: ate, this being especially true of saxicolous forms. Sanctum Dionysium, Reunion, Rodriguez (PC, lectotype)]. Parnielia subrevoluta Harmand, 1928:326 [type collection: In no case, however, do the pustules remain intact Cha Pon, Indochina, Demnnge (PC, lectotype)]. as in P. leucotyliza, a Japanese species. In Japan the Pornielia hitnunensis Zahlbruckner, 1930:187 [type collection: two species may also be separated by chemistry since P. leztcotyliza produces the “subaurulenta” ’ Entries without herharium designations of specimens terpenes. There is a very minor amount of variabil- collected by Hale are in US. ity in chromatograms of P. aul-zdenta from different 20 SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE10.-Distribution of Parmefina aurzrlenta hased on all available herharium specimens. geographic areas but it does not seem significant or stands and in heavily shaded maple (Acer saccha- consistent. I did not, incidentally, recheck the l-urn) climax forests. Parmelina aul-ulenta occurs terpene profiles of Pawnelia tiliacea var. efiol-esrens, frequently with Parmelia ruderta Acharius and P. albido-strarninea, P. subrevoluta, or P. hunanen- Pseudopal-rnelia capel-ata (L.) Hale. sis, but assumed that they have the “aurulenta” ter- In northern Wisconsin, an area of hardwood- penes since they are sorediate and fall outside the conifer forests more comparable in climate to range of P. leucotyliza. Should any of them be northern Europe, P. aurulenta had an overall fre- found to contain “subaurulenta” terpenes, the no- quency of only Il.670 at the 1.3 m level and 1.3% menclature of P. aurulenta would not change. at the base. It avoided Pinus spp. and grew almost The parent morph of Pal-melina awulenta might exclusively on Acel- and Querczis (W. Culberson, well be P. irl-ugans, which is identical in chemistry. 1955). This avoidance of Pinus was demonstrated It would be desirable, however, to examine more even more graphically by W. Culberson (1958) in specimens of P. irrugans in relation to P. aurulenta an ecological survey exclusively of Pinus in North in Japan before making a final decision. Carolina. Although otherwise common in this state, The ecology of P. aurulenta has been intensively not a single collection was made on conifers. studied in Wisconsin, an area of typical deciduous SPECI~IESSEx.\xIINED.-United States and Canada: See Hale and conifer forests well within its range. For ex- (1958384) for a map of distribution based on specimens in ample, I found in 1955 that it occurred on 30y0 of various American herbaria. Mexico: Michoacin, Wirth 329 a 2800 tree sample at the 1.3 m level on the trunk (US). Brazil: Minas Gerais, Eiten 6932 (US). Europe: Russia, Vasiljeva 9 (US). Tanzania: Tanga Province, Santesson 23358, and on 21y0 at base level in mesic deciduous forests. 23359 (Santesson herliarium, US). Union of South Africa: It was most common in closed oak-hickory stands Satal, Alniboria 8071, 8079 (LD); Transvaal, Alrnborn 7225 and much rarer in both open, savanna-like oak (LD). Madagascar: des .4 bboyes (TUR), Lernaitre (H). Paki- NUMBER 33 21

FIGURE11.-Species of Parmelina: a, P. nmagiensis (Kurokawa 550058 in US); b, P. antillensis (Hale 38291); c, P. aurulentn (Hale 18941); d, P. consors (Reit- and Klein 15973 in US); e, P. crassutn (Kurofmwu 550446 in US); 1, P. cryfitochlora (Hale 37782). (Scale in mm.) 22 SMITHSOSIAN CONTRIBUTIONS TO BOTANY

Stan: Ahmad 211 (L), Iqbnl 831, 839, 843 (US). Nepal: Noikett DIsTRIBuTroN.-South America. 9368 (BM). India: Almora District, Arvasthi 3970 (Aivasthi REhfARKs.-This robust, coriaceous lichen is char- herbarium, US); Kangra District, Hoeg 1492, 1497 (Awasthi herbarium); Punjab: Schelpe 3260 (BM); Tamil Natlu: Hale acterized by its heavily white-maculate cortex (Fig- 43664, 43812, 43816; Uttar Pradesh, Awasthi 3374 (Awasthi ure 3a), thick, furcate cilia (Figure 3a), and lack of herbarium); !Vest Bengal, Togashi (TNS). Sri Lanka: Degel- medullary chemistry. It is also the only species as- ius As407 (Degelius herbarium), Thwaites 32 (BM). China: signed to Parmelina with perforate apothecia, a Kiangsi Province, Tai (BM). Taiwan: Kurokawa 1359, 1390, trait usually associated with Parmotrema hfassa- 2694 (TNS). Korea: Den (TNS). Hong Kong: without collec- longo. The dense rhizine mat below to the margin tor, 1897 (BM, US); Hale 46393. Japan: Prov. Aki, Yan- agisatca 746-1 (TNS); Prov. Awa, Inobe 23 (TNS); Prov. and the generally close adnation, however, do not Higo, Mayebnra 169 (TNS) Togashi in Lichenes Japoniae conform to Parmotrema. Paymelina consors is evi- Exsiccati 177 (US); Prov. Hizen, Kurokawa 6261.5 (TNS): dently a common lichen in dry, scrubby forests and Prov. Kii, Kurokawa 59129 (TNS, US), Sasnki 481 (TNS); on trees along roads. The presumptive sorediate Prov. Settsu, Hale 29417, 29428; Prov. Shimotsuke, Kiirokaroa morph, P. pilosa, has a broader distribution in 56509 (TNS). Philippines: Benguet, Degeliiis As-888 (Degelius herbarium). Indonesia: Java, Groenhart 5872 (L, US). New South America and also occurs in Africa. Guinea: IMorobe District, Weber and McVean L-51280 SPECIMENS Ex4MIsEo.-Brazil: WawrU 669 (M); Mato Groqso, (COLO, US). Hawaiian Islands: Maui, Faurie 514 (LD), 866 Monte\ 10146 (DUKE, LD, US), Minaq Gerais, hfalme 201B (BM, PC), Hnle 31543; Kauai, Hale 31747, 31755. (S), I’ainio in Lichenes Brasilienses Exsiccati 398 (BM, M, TUR); Rio de Janeiro, Burchell 2303 (BM), Gardner 37 (BM), Glnziou 1821, 1836 (M), 1840 (M, S), Hemmendorff 4. Parmelina consors 5386 (S, UPS), Milne (BM); Rio Grande do Sul, Malme 1282 (LD, S, UC, UPS, US); Santa Catarina, Reitz and Klein FIGUKElld 12901, 15973, 16032a (US): SZo Paulo, Eiteti 5731 (US), Gehlt (US), 5924 (MICH), Puiggari (G). Paraguay: Itapua, Malme Pnrmelinci consors (N)lander) Hale, 1974:482. 1462 (S, UPS, US). Uiuguay: Latellaja, Lamb (H). Argentina: Pnrnielia co?isors Sylander, 1885:613 [type collection: Minas Buenos Aires, Kuhnemann 22 (S), Santesson 76, 77a (S); Gerais, Brazil, Weddell (H, Nylander herbarium number Misiones, Montes 36 (US), 3353 (WIS). 35277, lectotype)]. Parmelia balansae Miiller Argoviensis, 1888a: 1 [type collec- tion: AsunciOn, Paraguay, Balansn 8 (G, lectotype)]. Parmelin sampaiana Hue, 1899: 170 [type collection: S%o 5. Parmelina crassata, new species Paulo, Brazil, Sampaio (PC, lectotype)]. Parmelia continentalis Lynge, 1914:lll [type collection: FIGURElle Corumba, Mato Grosso, Brazil, hfalme 48 (S, lectotype)]. DEscRIPTIoN.-Thallus laxe adnatus, corticola, DEsCRIPTIoN.-Thallus adnate on bark, coria- crassatus coriaceusque, cinereo-albidus vel obscur- ceous greenish mineral gray and turning deep olive- ascens, usque ad 15 cm latus, lobis sublinearibus buff in the herbarium, 5-10 cm in diameter; lobes vel subirregularibus, contiguis vel congestis, 3-6 irregularly branched, sublinear-elongate to subir- mm latis, margine ciliatis, ciliis irregulariter dis- regular, apically rotund, often imbricate, 1-4 mm persis, nigris, 0.3-0.8 mm longis, superne planus vel wide, the margins crenate, moderately to densely aetate rugulosus, nitidus, continuus, isidiis soredi- ciliate, the cilia stout, blackish brown to black, isque destitutus, cortex superior ca. 12 pm crassus, becoming furcate, 0.2-1 .O mm long; upper surface epicorticatus, stratum gonidale 16-18 pm crassum, shiny, becoming heavily maculate, sometimes medulla omnino sulphureo-salmonea, 160-2 10 pm pruinose towards the tips, isidia and soredia crassa, cortex inferior fuscus, 14-16 pm crassus, lacking; medulla white; lower surface black, mod- subtus niger, dense rhizinosus, rhizinis nigris, ni- erately rhizinate, the rhizines black, thick and thin tidis, 1-2 mm longis, simplicibus vel pro parte intermixed, simple or squarrose. Apothecia substi- furcatis vel squarroso-ramosis. Apothecia numerosa, pitate, 2-8 mm in diameter, amphithecium smooth, subpedicillata, ainphithecio rugoso, usque ad 10 the disc cinnamon brown, rarely perforate; spores mm diamentro, sporis octonis, 6-8 X 14-15 pm. 8,S-lZ X 14-19 pm. CHEnmTR\r.-Cortex K + yellow (atranorin); CHEMISTRY.-Cortex K+ yellow (atranorin); medulla more intensively yellow with color tests medulla negative with color tests (no substances (zeorin, leucotylin and associated terpenes, and demonstrated). secalonic acid A). NUMBER 33 23

HoLOTYPE.--;\ft. Akagi, Japan, S. Kzirokazun mens from the type-locality in Dominica. It is 550466, 23 August 1955 (US; isotype in TNS). rather common at one locality (base of Morne DIsTRrBLJTr0N.-Japan. Anglais) on trees in a pasture at 600-800 m eleva- REivARKs.-This is another member of the P. tion, One collection that I made in South India sitbaic~iilenfagroup so abundantly developed in also appears to lie this species. It is so small and Japan. It has the same chemistry as P. subazirulenta inconspicuous that it has undoubtedly been over- but is well separated by the larger, leathery thallus, looked elsewhere or, iC collected, has been set aside averaging 218 pm in thickness (sample of 10 speci- as unidentifiable. It has the same chemistry as mens), as opposed to 150 pm for P. subaurzilenta. P. dissecta, a common species with which it is It also has significantly larger, substipitate apothe- clearly allied as the sorediate morph of the extinct cia. Parmelia in-ugans, which has large apothecia, or still undiscovered parent morpli. It also has the can be distinguished by chemistry (the “aurulenta” same chemistry as sorediate Hypotl-achynn l-evoliita terpenes) and thinner thallus. (Floerke) Hale, which woulcl normally be amply This species previously had been identified distinguished by a larger tliallus, more ascending (along with P. irrugans) as “Parmelia homogenes” lobes, dichotomously branched rhizines, and in Japan, but, as explained below, P. homogenes broader soretliate pustules. A poorly developed was misinterpreted and is in reality the same as specimen of H. iavolicta, however, and a large speci- P. .siibnicrulentn, Puvmelia cmssatn is a common men of P. c~pfochlol-awould have to be separated species in Japan, but since I have not had an with extreme care. opportunity to re-examine tlie chemistry of the rich QI’ECIMFNS ExA\rINl.o.-~ominic;i: Hale 3535i, 37782. India: collections in TNS, I have cited below only those Tamil Nadu, Hale 40203. specimens also represented in US.

SIVCIMFX Ex \~is~o.--Japaii: Prov. Kozuke, Kurokarc~a 7. Parmelina damaziana, new combination .58583 (TNS,US) ; Prov. Sagami, Kurokarca 58039 (TNS, US) : I’rov. Suruga, ‘4saliina 539, 2G27 (US). FIGURE120

Partrielin danzaziatia Zahlbruckner, 1905:.541 [type collection: 6. Parmelina cryptochlora Mt. Ituculumi, Brazil, Daniazio 137j (W,lectotype; iso- lectotype in G)]. FIGUREllf Partnelin brachpconidia Zahlbruckner, 1908:465 [type collec- tion: \‘ellozo, Serra (lo Our0 Preto, Braiil, Daniazio 1741 PorrridiTia crjptochloro (Vainio) Hale, 19743482. (LV, lectot!pe)]. Pnrrnelin crjptoclilorn 1-ainio, 1896:34 [type collection: Lau- Pnririelin brnchyrovidin var. cliloroccirpn Zahlbruckner, dat, Dominica. Elliott 912 (BM, lectot!pe)]. 1908:466 [type collection: Cachoeira do Campo, Brazil, DEscRIPTIos.-Thallus closely adnate on bark, Dnmnzio I740 (W, lectotype)]. Pnrtnelici crystallorum Lynge, 1914:128 [t\pe collection: Cor- whitish mineral gray, 1-3 cm broad; lobes sub- covado, Rio de ,janeiro, Brazil, Mnlme 59* (S, lectotype)]. linear, separate to crowded, 1-2 mm wide, the mar- ginal cilia sparsely developed, 0.1-0.3 mm long; D~sc~~~l~o~.-Thallubclosely adnate on twigs, upper surface plane to convex, sorediate toward the whitish ashy mineral gray, 3-6 cm broad; lobes tips, tlie soralia capitate, up to 1 mm in diameter short, sublinear-elongate to subirregular, 1-3 mm (Figure 5b), the sorediate lobe tips becoming wide, tlie marginal cilia evenly dispersed, about 0.5 revolute: lower surface black, sparsely rhizinate, tlie mm long; upper surface plane, shiny; lower surface rliizines black, simple. Apotliecia not seen. Black, moderately rhizinate, the rhizines simple, CHEhIISTRY.-COrteX K + yellow (atranorin): black. Apothecia common, sessile to substipitate, to medulla K-, C+ rose, KC+ red, P- (gyroplioric 12 mm in diameter, the disc flat, often radially split; acid). spores 8,s-12 X 12-18 pm. DISTRIBUTIOS.-~YeSt Indies and India (?). CHEmsTRY.-Cortex K + yellow (atranorin); REMARKS.--ASI explained in my study of the medulla K-, C-, KCf rose, P- (“horrescens” Pawneline of Dominica (Hale, 1971a: 12), Vainio’s unknown). type collection is so poor that I had not been able DIsrRIBuTIo,v.-So~itli !imerica. to interpret it until I collected additional speci- RE~r,~RIts.-AlthoUgll clearly distinct because of 24 SSIITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE12.4pecies of Parmelina: a, P. damaziana (Dainazio 1375 in W); b, P. degelii (Degelius A-I82 in Degelius herbarium); c, P. de?iegans (Hale 43855); d, P. dissecta (Hale 14999); e, P. endoleuca (Weber in Lichenes Exsiccati 244 in US): f, P. enorinis (Jellicoe in US). (Scale in mm.) NUMBER 33 25 the chemical constitutents, P. damaziana is a rare degelii is named in honor of Dr. Gunnar Degelius, and not well comprehended species. The lectotype who has enriched our knowledge of lichen distribu- was collected on tree branches whereas Lynge’s tions through his careful collecting efforts in the species was apparently saxicolous. Burchell 1105-06 tropics. is tentatively placed here since the chemistry, while not clear, is closest to the “horrescens” type. Parme- 9. Parmelina denegans lina damaziana has rather large spores, just as the other members of the P. horrescens group. It could FILLRF 12C be regarded theoretically as the parent morph for P. howescens, P. schindleria, and P. subfatiscens, but Parmelzna denegans (Nylander) Hale, 1974,482. Parnelta denegans Njlander, 1900 6 [tjpe collection: Ram- all of these vegetative morphs have smaller, more podde, Ceylon, Almquzst (H, Njlander herbarium number fragile thalli. 35129, lectotype; isolectotype in S)]. SPECIXIENS ExAalrNED.-BraLil: Burchell 1105-06 (BM,US). DE~CRIPTI~X.-Thallus adnate to closely adnate on bark, light greenish mineral gray, 5-9 cm broad; 8. Parmelina degelii, new species lobes irregularly branched, sublinear, imbricate, 1-3 mm wide, the marginal cilia irregularly dispersed, FIGURE12h 0.4-0.7 mm long; upper surface shiny, weakly macu- DEscRIPrroN.-Thallus adnatus, corticola, fragilis, late, plane but soon developing small pustulate pallide viridi-albicans, 6-8 cm latus, lobis plus areas, the pustules eventually breaking open minusve subirreqularibus, 2-3 mm latis, margine apically and forming coarse soredia; medulla pale aetate lobulato-dissectis, ciliatis, ciliis usque ad 0.8 orange-salmon, in part K+ purple; lower surface mm longis, superne planus vel aetate rugulosus, black, densely rhizinate, the rhizines black, simple nitidus, sparse vel modice isidiatus, isidiis primum to sparsely branched. Apothecia sessile, 2-3 mm in cylindricis, mox procumbentibus, dorsiventralibus, diameter; spores 8, 5-7 X 7-9 p.m. expansis, ad 0.6 mm longis, cortex superior 12 pm CHEhmTRY.-Cortex K + yellow (atranorin); crassus, epicorticatus, stratum gonidiale 12-14 pm medulla more intensively yellow-orange with color crassum, medulla pallide sulfurea, 60-90 pm tests (zeorin, leucotylin and associated terpenes, crassa, cortex inferior paraplectenchymatus, brun- secalonic acid A, and possibly anthraquinones). neus, 12 p,m crassus, subtus niger, dense rhizinosus, DIsTRIsuTIoN.-India, Sri Lanka, and Sabah. rhizinis nigris, simplicibus, 0.5-09 mm longis. REMARKS.-Except for the darker reddish orange Apothecia ignota. pigment in the lower part of the medulla, this spe- CHEMIsTRY.-Cortex K + yellow (atranorin); cies is the pustulate morph of P. subaurulenta. medulla more intensively yellow with color tests They have similar lobe configuration and habitats, occurring at about 2000 m elevation in open forests. (zeorin, leucotylic acid, apparently secalonic acid The lectotype specimen is fertile and has few A, and traces of other pigments and terpenes). pustules. The specimens that I collected are heavily HoLoTYPE.-Rain forest, Cairns Cove, Queens- pustulate-sorediate by contrast. land, Australia, elevation 600 m, G. Degelius A-I82 (Degelius herbarium; isotype in US). SPECIMENSExAM1xFD.-India: Tamil Nadu, Foreau 4128 DI~TRIRUTI~N.-A~Straha. (Awa3thi herbarium, US), Hale 40239, 43636, 43762, 43855. Sri Lanka: Thruaites (UPS). Sabah: Hale 29020. REMARKS.-This is the only species of section Myelochroa that has been discovered in Australia. It is remarkable that such a conspicuous lichen 10. Parmelina dissecta should be collected so recently in Queensland. The FIGCJRE12d. 13 terpene chemistry is close to P. aurulenta except for the absence of one spot (Figure 7). Regardless Parmelina dissecta (Nylander) Hale, 1974:482. of chemistry, however, it is distinct from all other Parmelia dissecta h’ylander, 18823451 [type collection: Fon- tainebleau, France, A’ylander (H, Nylander herbarium species in the P. subaurulenta complex because of number 35131, lectotype; isolectotype in PC)]. the production of large lobulate isidia. Parmelina Parinelia laeuigata var. gracilis f. furfuracea Miiller Argovien- 26 SMITHSONIAN CONTRIBUTIONS TO BOTANY

sis, 1888c:529 [type collection: Faxina, Brazil, Puiggari 47 amphithecium isidiate: 1-4 mm in diameter: spores (G, lectotype)]. 8,8-10 X 12-17 pm. Parmelia minarum Vainio, 1890:48. CHEMIsmY.-Cortex K+ yellow (atranorin); Parmelia amazonica var. hwnotii Hue, 1899:158. Parmelia puiggari Gyelnik, 1931:288 [based on Parmelia medulla K-, C+ rose, KC+ red, P- gyrophoric lamigata var. gracilis f. furfuracea Muller Argoviensis]. acid with or without associated unknown sub- Parmelia camtschadalis var. epiphylla Cengia Sambo, stances). 1938:379. DIsTRIBuTI0N.-Pantemperate and montane pan- Parmelia hubrichtii Berry, 1941:102. tropical (Figure 13). [For full citation of synonymy, see Hale, 1971a:6.] REMAws.-This is one of the most widespread and commonly collected species in the genus, DEScRIPTIoN.-Tha1Ius adnate on bark or rock, especially in temperate zones, occurring on a wide yellowish glaucous to pale greenish mineral gray, variety of substrates in open, secondary forests. In 3-7 cm broad: lobes sublinear-elongate, contiguous, tropical regions it is strictly montane, usually being 1-3 mm wide, the marginal cilia irregularly dis- most abundant in cloud forests up to 2300 m eleva- persed, mostly simple, to 0.7 mm long: upper sur- tion. Morphological variation is wide, even though face shiny, emaculate, plane to convex, moderately the basic characters, isidia, ciliate lobes, and gyro- to densely isidiate, the isidia cylindrical, erect, phoric acid, are constant. The lobes, for example, often branched, less than 0.5 mm high; medulla are narrow, sublinear, and separate in the lectotype white; lower surface black, moderately rhizinate, specimen. At the other extreme are plants such as the rhizines black, shiny, simple or sparsely those collected in Dominica (Hale, 1971a) with branched. Apothecia adnate, the rim crenate, the broader, contiguous to subimbricate lobes.

FIGURE 13.-Distribution of Parmelina dissecta based on all available herbarium specimens. NUMBER 33 27

Parmelina dissecta has been correctly identified 11. Parmelina endoleuca, new combination by European lichenologists both on the Continent FIGURE12e and in tropical regions, Degelius (1941:60) was the first to identify the species correctly in North Parmelia endole.uca TaFlor, 1847: 167 [type collection: Swan America. He noted two populations differing in C Rixer, Australia, Drurnmond (FH-Tajl, lectotjpe)]. test (C+ rose or C-) and presence or absence of DEscRIP’rIos.-Thallus closely adnate on bark, cilia on the isidia. Hale and Kurokawa (1962) whitish mineral gray, 2-4 cm broad; lobes sublinear determined that the C- ciliate population is P. to subirregular, short and crowded, 1-2 mm wide, horrescens. No parent morph has been discovered the marginal cilia sparse, less than 0.3 mm long; for P. dissecta, but it forms a convenient nucleus upper surface shiny, plane to minutely rugulose, of the “dissecta” group, including isidiate but sig- becoming more strongly rugose and lobulate toward nificantly larger P. melanochaeta, sorediate P. cryp- the center, heavily pycnidiate, soredia and isidia tochlora, and isidiate-lobulate P. subfatiscens (Fig- lacking; medulla white: lower surface dark brown ure 9). at the margins and black toward the center, mod- SPECIMENI EXAMINED(selected).-United States: Pennsyl- erately rhizinate, the rhizines simple, brown but vania, Hale 17140; Marjland, Hale 14367; West Virginia, blackening at maturity. Apothecia very numerous, Hale 11470: Ohio, Hale 13910, 15735; Kentucky, Hale 13770; I’irginia. Hale 38419, Liittrell 3799 (US); Tennessee, Hale sessile, 1-2.5 mm in diameter, the disc dark brown, 31124, 37038; North Carolina, Culberson 7099 (DUKE), Hale plane; spores 8, 6-7 X 11-12 pm. 30645; South Carolina, Culberson 7459 (DUKE), Hale 16601; CHExmrRY.--Cortex K + yellow (atranorin); Alabama, Hale 33937, Skorepa 4547 (US); Georgia, Culber- medulla negative with color tests (an unidentified son 7249 (DUKE), Hale 7509; Florida, Nelson (US) (see fatty acid). Moore, 1968:220 for additional records in Florida); Missis- DIsTRIsuTIoN.-Austra~ia. sippi, Hale 7977, Hubricht 1532 (US): Texas, Hale 5242. Mexico: Chiapas, Hale 20304; \‘era Cruz, Hale 21231. Pan- REalARKs.-In my earlier work on Parmelia I ama: Chiriqui, Hale 38879. Cuba: Oriente, Imshaug 24720 placed this species in synomymy under Parmelina (MSC). Jamaica: Inishaug 14172 (MSC). Guadeloupe: Dim galbina, a Japanese-North American species. Since 1030 (TUR). Dominica: Hale 35569 (see Hale, 1971a:13, for the Australian locality seemed at variance with the additional records). Martinique: Degeliits (Degelius herbar- Arcto-Tertiary distribution pattern of P. galbina ium). St. T’incent: Guilding 46 (BM). Trinidad: Imshaug 32103 (MSC). Colombia: Antioquia, Nee and Mori 4258 (US). and since my first chemical tests had been done l’enezuela: Distrito Federal, Dennis 1568 (BM); Mkrida, Hale with crystal techniques, I have checked the type of 42543; TAchira, Hale 42540, 45112. Brazil: Minas Gerais, Parmelia endoleiica again and found that it con- Eiten 6867 (US); Parana, Montes 10121 (hIVM). Europe: tains neither galbinic acid nor any of the associated Portugal, Persson (UPS), Sampaio in Lichenes de Portugal terpenes that characterize P. galbina. I have con- 252 (LD, US), Tauares 97 (H, LD, WIS), 632 (US); Spain, Degelius (S), Santesson 19428 (UPS), Schauer in Lichenes cluded, therefore, that Parmelia endoleuca is a good Selecti Exsiccati 265 (LD, US); France, des Abbayes in species in spite of the very close external resem- Lichenes Armoricani Spectabiles Exsiccati 88 (LD, W), Has- blance to P. galbina. The fatty acid falls higher on selrot (S); Italy, Gresino (S), Sbarbaro (US). Kenya: Rift the chromatographic plates than either caperatic Valley Province, Maas Geesteraniis 6067 (L). Tanzania: Tanga acid or protolichesterinic acid. The species is Prov., Santesson 23137 (UPS). Union of South Africa, Cape Province, Almborn 301, 670, 3201 (LD); Natal, Almborn endemic to the dry scrub forests of eastern 9192, 9525 (LD), Hoeg (TRH). Swaziland: Almborn 7904 Australia. (LD). India: Tamil Nadu, Degelius As-238 (Degelius herbar- SPECIVENSExA\rIN~.o.-Australia: Australian Capital Terri- ium), Hale 40222, 43715, 43852; West Bengal, Degelius As- tory, Weber in Lichenes Exsiccati 244 (COLO, US). 21 1 (Degelius herbarium). Sri Lanka: Degelius As-466 (Degel- ius herbarium, US). Malay: Pahang, Hale 30137, 30483. Taiwan: Kurokawa 2400 (TNS). Japan: Prov. Aki, Hale 12. Parmelina enormis 29368: Prov. Higo, Mayebara 1.58 (TNS); Prov. Kazusa, Kuro- kawa 64096 (TNS); Prov. Ohmi, Hale 29484; Prov. Owari, FIGURE12f Asahina 146 (TNS); Prov. Settsu, Hale 29411. Philippines: Mountain Province, Degelius As-% (Degelius herbarium), Parmelina enormis (Hale) Hale, 1974:482. Hale 26040. Indonesia: Java, Groenhart 6022 (L), 8483 Parmelia enormis Hale, 1972c:344 [type collection: Nyaka (BOR); Sumatra, Groenhart 967 (L). Hawaii: Hale 31340, Plateau, Zambia, Jellicoe, September 1968 (BM, holotype; 32933. isotype in US)]. 28 SMITHSONIAN CONTRIBUTIONS TO BOTANY

DESCRIPTIoN.-Thahs expanded, loosely attached the amphithecium rugose, 2-4 mm in diameter; on rock, coriaceous, whitish to ivory mineral gray, spores 8,5 X 10-1 1 ,urn. 10-30 cm broad; lobes sublinear, dichotomously CHEMwRY.-Cortex K+ yellow (atranorin); branched, imbricate and crowded toward the cen- medulla more intensively yellow with color tests ter, 5-8 mm wide, the margins entire, ciliate mostly (zeorin, leucotylin and associated terpenes, and in the lobe axils, the cilia to 2.0 mm long; upper secalonic acid A). surface plane to convex, continuous, lacking soredia DIsTRIBuTI0N.-Korea and Japan. and isidia; medulla white; lower surface pale REMARKS.-T~~Sis one of the commonest lichens brown, densely rhizinate, the rhizines pale brown, in Japan, recognizable by the large rather fragile simple. Apothecia numerous, subpedicillate, 3-8 thallus with patches of the yellow medulla visible mm in diameter; spores 8, 6-7 x 8-11 where the cortex breaks away (Figure 6a). It CHEMIsTRY.-Cortex K + yellow (atranorin); grows on both trees and rocks in open forests. The medulla I<+ yellow turning red, C-, KC-, P+ densely pustulate-rugose areas are more or less iden- orange (salazinic acid). tical to those of P. rhytidodes but they differ in DIsTRIBuTI0N.-Zambia. breaking open. In addition, P. rhytidodes has a dif- REMARKS.-This species does not fit well in ferent terpene profile, the “aurulenta” series. Par- Parrnelina if judged from lobe configuration alone. melina entotheiochroa is clearly a member of the It appears to be a very large, typically sublinear P. subaurulenta group, most closely related to P. Hypotrachyna, but the rhizines are simple (not leucotyliza and lacking a parent morph. dichotomously branched), the lower surface brown SPECIhfEKS ExAhr1xEo.-Korea: FUjikaWa (TNS). Japan: (not black), and the marginal cilia are distinct. No Prov. Aki, Hale 29534; Prov. Awa, Kishida (TNS); Prov. vegetative morphs are known. The species is known Bitchu, Yoshino (TNS); Prov. Etchu, Asahina (TNS); Prov. only from the type collection but should be found Higo, Fujikawa (TNS); Prov. Ise, Uemara 102 (US); Prov. IZU, on rock outcrops in countries adjacent to Zambia. Asahina (TNS); Prov. Kii, Kurokawa 64131 (TNS), Numajiri 394 (TNS); Prov. Kozuke, Uematsu 388 (TSS); Prov. Musa- shi, Kurokawa 550535 (TXS, US), Nakaji 50 (TNS); Prov. Ohmi, Hale 29456a; Prov. Rikuzen, Iishiba 43 (US); Prov. 13. Parmelina entotheiochroa Satsuma, Fujikawa (TNS); Prov. Settsu, Ui 2667 (TSS); Prov. Shimotsuke, Asahina (TNS), Culberson 10680 (DUKE, US); FIGURE14a Prov. Shinano, Kurokawa 51620 (TSS, US), Togashi 5100 (TNS); Prov. Suruga, Asahina (TNS). Parmelina entotheiochroa (Hue) Hale, 1974:482. Parrnelia entotheiochroa Hue, 1899: 161 [type collection: Hakodate, Japan, Faurie 104 pro parte (PC, lectotype; iso- 14. Parrnelina expallida lectotype in BM)]. FIGURE14b DEscRIP’rIoN.-Tha11us adnate to loosely attached, Parnaelina expallida (Kurokawa) Hale, 1974:482. fragile and easily breaking apart, saxicolous and Parrnelia expallida Kurokawa, 1968a: 191 [type collection: Mt. corticolous, light greenish mineral gray with small Nanfong, Prov. Kaoshiung, Taiwan, Kurokawa 2930 (TNS, holo type)]. sulfur-yellow patches where the cortex has broken away, 8-10 cm broad; lobes subirregular, congested, DESCRIPTIoN.-Thallus adnate on bark, whitish 2-3 mm wide, marginal cilia produced mostly in mineral gray, 5-7 cm broad; lobes sublinear, more the lobe axils, about 0.5 mm long; upper surface or less imbricate, 1-3 mm wide, the marginal cilia plane and shiny on young lobes but soon becoming sparse, simple, about 0.5 mm long; upper surface strongly rugose-wrinkled to pustulate, the surface plane, emaculate and opaque, densely isidiate, the dull and in part white pruinose, the wrinkles or isidia cylindrical, simple or becoming coralloid- pustules breaking open and large areas of cortex branched, 0.3-1.0 mm high; medulla white; lower (1-3 mm across) breaking away to expose the deep surface pale brown or darkening, moderately sulfur-colored medulla; lower surface black except rhizinate, the rhizines simple or rarely furcate, for a narrow brown zone at the tips, densely rhizin- about 0.5 mm long. Apothecia rare, sessile, the rim ate to the margins, the rhizines black and shiny, crenate and the amphithecium isidiate, 2-5 mm in simple or sparsely furcate. Apothecia rare, sessile, diameter; spores 8, 8-10 X 14-17 ,urn. NUMBER 33 29

FIGURI:14.--Species of Parmeliiza: a, P. entotheiochroa (Hale 29456a); b, P. expollida (Kurokair-a 2930 in TKS); c, P. gnlbina (Hole 2341.5); tl, P. hoyachitzensis (Kurokawa 67081 in TNS): e, P. hetemloba (Schiflner in \\'); f, P. liorrescens (Hnle 36999). 90 SMITHSONIAN CONTRIBUTIONS TO BOTANY CHEMIs-rRY.-Cortex K + yellow (atranorin); simple. Apothecia very common, adnate, 2-5 mm in medulla negative with color tests (protolichesterinic diameter, the disc cinnamon brown; spores 8, 7-9 X acid and an unidentified fatty acid). 10-13 pm. DIsmInuTIoN.-India, Thailand, and Taiwan. CHEMIsrRY.-Cortex K + yellow (atranorin); As pointed out by Kurokawa (1968a:191), an medulla K+ yellow, C- or C+ yellow, KC-, unusual feature of this species is the brown lower P + pale orange (galbinic acid, zeorin, secalonic surface, indicating some relationship with P. versi- acid A, trace of salazinic acid, and leucotylin and forrnis, a South American species. It is also one of associated terpenes). the few Parrnelinae to produce a fatty acid. DIsTRIsvTIo;v.-Japan and eastern North America. REMARKs.-During a visit to the Academy of SrEcIarENs EXAMINF‘.D.-See Kurokawa (1968a:193) for records Natural Sciences in Philadelphia in 1958 I found a from India, Taiwan, and Japan. specimen in the hfuhlenberg herbarium without a name but with a penciled number “15.2.” I later 15. Parmelina galbina found the same plant in the Acharian collection (H) also numbered 15.2 and described by Acharius FIGURES14c, 15 as Parrnelia galbina. This discovery clarified the status of one of the very common corticolous lichens Parmelina galbina (Acharius) Hale, 1974:482. in eastern North America. W. Culberson (1961: 173) Parmelia galbina Acharius, 1814: 195 [type collection: North subsequently studied the problem and lectotypified America (?Pennsylvania), Muhlenberg (H-Ach, lectotype; isolectotype in PH)]. P. subquercifolia Hue as synonymous with P. gal- Parmelia tiliacea var. minor Muller Argoviensis, 1877:X bina. The second syntype of Hue’s species was [type collection: near Dallas, Texas, Boll (G, lectotype; determined by Culberson to be Parmelia livida isolectotypes in FH, M, W)]. (= Hypotrachyna livida (Taylor) Hale), another Parmelia tiliacea var. sulphurosa Tuckerman, 1882:57 [type very common corticolous lichen in eastern North collection: Illinois, Hall (FH-Tuck, lectotype)]. Parmelia deminuta Hue, 1899:156 [type collection: Texas, America distinguished by more or less distinctly Boll (PC, lectotype)]. dichotomously branched rhizines and the lividic Parmelia subquercifolia Hue, 1899: 157 [type collection: Ohio, acid complex (Hale, 1975a:45). As a rule, H. livida Sullivant (PC, lectotype)]. has a whiter cast than P. galbina, is more robust, Parmelia subquercifolia var. rugosa Hue, 1899: 175 [type col- and lacks pigments in the medulla or under the lection: Oyama, Japan, Faurie (not seen)]. Parmelia tiliacea subquercifolia (Hue) Merrill and Burnham apothecia. Still, care must be taken when identi- in Burnham, 1922:75 [rank not designated]. fying these two species. One other distinguishing Parmelia quercina var. sulphurosa (Tuckerman) Zahlbruck- character is the unique moniliform cells in the ner, 1929:192. medulla of P. galbina and its morphs, P. hayachin- Parmelia sulphurosa (Tuckerman) Fink, 1935:328. Parmelia laeuigata ssp. extremi-orientalis f. rugosa (Hue) ensis, P. metarevoluta, and P. obsessa, first described Asahina, 1951c:291. by Asahina (1951~).The cells, however, are not Parmelia subquercifolia f. subradiata Asahina, 1952:98 [type easy to find and their exact significance is unknown collection: base of Mt. Fuji, Japan, Yamamoto (TNS, lecto- at this time. type) (nomen nudum in Asahina, 1951c:291)]. Parmelia galbina var. rugosa (Hue) Asahina, 1963:225. SPECIMENS EXAMINED.-United States: See W. Culberson Parmelia galbina var. subradiata (Asahina) Asahina, 1963:225. (1961:lil) for a discussion of the distribution and map of localities in North America based on specimens in DUKE DEscRIPTIoN.-Thallus closely adnate on bark, and US (Figure 15). Japan: Prov. Aki, Kurokawa 64431 yellowish-glaucous to greenish mineral gray, 3-10 (TXS); Prov. Bungo, Kurokawa 63191 (TNS); Prov. Hizen, cm in diameter; lobes short, sublinear, contiguous, Kurokawa 62588 (TNS); Prov. Hoki, Yasuda (TNS); Prov. 1-2 mm wide, marginal cilia mostly in axils, some- Hyogo, Tagawa L15 (TNS); Prov. Kozuke, Degelius As-1072 times only sparsely developed, less than 0.5 mm (Degelius herbarium); Prov. Mikawa, Takaki 353 (TNS): long; upper surface plane to rugulose, continuous, Prov. Musashi, Kurokawa 64279 (TNS); Prov. Mutsu, Kuro- kawa 550385 (TNS); Prov. Shimotsuke, Kurokawa 64057 pycnidia usually numerous; medulla white or pale (TNS); Prov. Shinano, Kurokawa 51758 (TNS, US), 59179 yellow to orange, especially beneath the apothecia; (TNS); Prov. Totomi, h’akanishi 27 (Kobe University); Prov. lower surface densely rhizinate. the rhizines black, Ugo, Suzuki 272 (TNS). NUMBER 33 31

17. Parmelina heteroloba

FIGURE14e

Parrnelina heteroloba (Zahlbruckner) Hale, 1974:482. Parrnelia heteroloba Zahlhruckner, 1909: 171 [tlpe collection: Mt. Itatiaya, Rio de Janeiro, Brazil, Schiflner (W, lecto- type)l. DEscRIPTIoN.-Thallus adnate on bark, light buff mineral gray in the herbarium, 5-8 cm broad; lobes more or less sublinear, short and imbricate, be- coming lobulate toward the center, 2-3 mm wide, the marginal cilia mostly in the lobe axils, to 0.4 mm long; upper surface shiny, plane to rugulose, becoming somewhat lobulate; lower surface black except for a narrow brown zone at the tips, densely rhizinate, the rhizines black, simple or sparsely furcate. Apothecia numerous, substipitate, the disc splitting, 2-9 mm in diameter; spores 8, 8 X 12 pm. CHEMIsTRY.-Cortex K + yellow (atranorin); me- dulla negative with color tests (an unidentified fatty acid and a faint unknown spot, perhaps related to FIGURE15.-Distribution of Parrnelina galbina in Korth America (taken froin W. Culberson, 1961). one of the “horrescens” unknown). DIsmIsuTIoN.-Brazil. REMARKS.-siXICe this species is represented only by the single type collection, it is difficult to gen- 16. Parmelina hayachinensis eralize on its affinities to other Parmelinae. It would probably be mistaken for P. damaziana, which has FIGURE14d a different chemistry (“horrescens” unknown) and Parrnelina hayaclzinensis (Kurokait-a) Hale, 1971:482. smaller spores. It is only one of three species in Prrrnlelia ha)achinensis Kurokawa, 1968b:350 [type collection: Paimelina with fatty acids, the others being P. hit. Hayachine, Proy. Rikuchu, Japan, Kurokawa 67081 endolezica and P. expallida. Parmelilza heteroloba (TSS, holotype)]. has not been recollected at Alt. Itatiaya, a well DEscRIPnoN.-Thallus adnate on bark, whitish to known site that has been visited by several experi- greenish mineral gray, 5-8 cm broad; lobes sub- enced lichen collectors, and it may be an extinct linear, 1.5-3 mm wide, marginal cilia simple, about species. 0.2 mm long; upper surface shiny, faintly maculate, becoming densely pustulate, the pustules becoming 18. Parmelina horrescens aggregated, erupting but not forming soredia; medulla white; lower surface black, densely rhizin- FIGURE14f ate, the rhizines black, simple. Apothecia not seen. CHE~IsTRY.-Cortex K + yellow (atranorin); Paririelina horresce?is (.laylor) Hale, 1974:482. medulla K+ yellow turning red, C-, KC-, P+ Pnr,,relin korrescens Taylor in Mackay, 1836:144 [t!Iie collec- orange (galbinic acid, leucotylin, and zeorin). tion: Dunkerron Mountains, Kerry, Ireland, Taylor (FH- Tayl, lectotype)]. DxsmIsuTIoN.-Japan. REMARKS.-AS pointed out by Kurokawa (1968b: Dk:scRrp?.xos.-Thallus closely adnate to adnate on 350), this species is a member of the P. galbina bark, rocks, or mosses over rocks, whitish to green- complex, representing the pustulate morph. It is ish mineral gray, 2-5 cm broad; lobes more or less known only from the type collection growing on dichotomously branched, sublinear, often crowded Cryptomeria in Japan. and imbricate, 0.5-2.0 mm wide, the margins cre- SMITHSONIAN CONTRIBUTIONS TO BOTANY

nate, often becoming lobulate, ciliate, the cilia more (US); Kentucky, Hale 13743a; West Virginia, Hale 10612, or less evenly dispersed, black, simple, 0.3-0.8 mm 11772, 11878; Virginia, Hale 18388, 33153, Reed 9091 (Reed herbarium), Roller 400 (US); h‘orth Carolina, Culberson long; upper surface shiny, emaculate, densely isidi- 5114, 5706, 5744, 7134 (DUKE), Hale 18042, 15058, Imshaug ate, the isidia cylindrical, often branched and 22174, 22178, 22353 (MSC, US); Tennessee, Hale 31106, 36921, apically spinulate or short-ciliate, in part becoming 36959, 36970, Moore 284 (US), Phillips 358, 377 (US), Skorepa procumbent; medulla white; lower surface black, 5527 (US); South Carolina, Hale 7723; Georgia, Hale 7405, moderately rhizinate, the rhizines black, simple. 7537, 16761, 16776, 30882, 30885; Alabama, Hale 7186, 7216, Apothecia rare, sessile, 2-4 mm in diameter, the 31164, 33779, 33918, 34121, 34174, McCullough 2195 (US); Florida, Hale 21685 (for additional records see Moore, amphithecium isidiate, the disc splitting at ma- 1968:220). Mexico: Chiapas, Hale 20204, 20223, 20400a, 20414, turity; spores 8, 10-12 X 16-18 pm. 20549, 21086. Guatemala: Baja Vera Paz, Hale 45828. Pan- CHExmTRY.-Cortex K + yellow (atranorin); me- ama: Darien, >Vori and Gentry 4309 (US); Panami, Hale dulla K -, C - or C + faint rose, KC + rose or red, 38451. Cuba: Oriente, Imshaug 24737, 24810, 24932 (MSC). P (trace of gyrophoric acid, “horrescens” un- Jamaica: Zmshnug 14216 (MSC). Dominican Republic: Cordil- - lera Central, Imshaug 23516 (MSC), Wetmore 3739 (MSC); La known falling above gyrophoric acid on chroma- T’ega, Allnrrl 17695a (US). Haiti: Ouest, Fahius 2-4 (US), tographic plates, and one or two other unidentified Ini.rhaug 22768, 22837, 22860 (MSC, US), Wetmore 3222 s POt S) . (MSC); Sud, Imshaug 23233 (MSC, US). Venezuela: Distrito DIsTRIsuTIozr.-Pantemperate and montane pan- Federal, Deiinis 2394 (BM), Santesson 6679 (S); Wrida, Hale tropical. 42038, 42067, 42952, 45121, 45201. Urugna): Trienta y Tres, Osorio 5931 (MT‘M). France: Harmand (DUKE). Spain: Ponte- REMARK\.-,& discussed under P. dissecta, this vedra, Schnurr (XI); Tenerife, Zmshazcg 34476, 35677 (MSC). species only recently was differentiated correctly Union of South Africa: Cape Province, Alniborn 1442 (LD). from P. dissecta (Hale and Kurokawa, 1962:2), an India: Tamil Nadn, Hale 43784. Philippines: Mountain Pror- isidiate lichen with gyrophoric acid. Parmelina ince, Hale 26531. Indonesia: Java, Groenhart 2903 (L, US), horrescens is characterized by dense fine isidia with 6279 (L), Kurokau,a 2071 (TNS). Taiwan: Kurokawa 693 (TSS). Japan: Proy. Buzen, Kiirokawa 62468, 63166 (TNS); greater or lesser development of short apical cilia Prov. Kii, Kurokawa 64127 (TNS);Prov. Ohmi, Hale 29465. (Figure 4c). The isidia sometimes become procum- Australia: Sew South \Vales, Cizeel L1708 (NSW), Craigie bent and lobulate. On the whole, the lobes are (NSW). Sew Zealand: Wade 85 (BM, US). somewhat narrower and more appressed than in P. dissecta. Parinelina horrescens has essentially the 19. same geographic range as P. dissecta and occurs Parmelina immiscens most abundantly in the temperate deciduous FIGURE16a forests of the eastern United States and Japan. It is montane in the tropics, occurring as high as 3000 m Pnrmelina irnmiscens (Nylander) Hale, 1974:482. in the paramos of Venezuela. Parmelia immiscens Nylander, 1885:606 [type collection: The chemistry of P. horrcscens is now being Orizaba, Mexico, Galeotti 6879 (PC, 1ectot)pe; isolectotype in H, Xilander herbarium number 35674)l. studied by several lichen chemists. A number of Pnrmelia michoacanensis Body de Lesdain, 1914:7 [type unidentified spots appear in both hexane and ben- collection: Jeslis del Monte, Morelia, Michoacan, Mexico, zene solvent systems, one of which is probably a Arsene 4456 (US, lectotype; isolectotypes in COLO, DUKE, trace of gyrophoric acid. The other spots fall above G, LE, and UPS)]. and below gyrophoric acid and seem to represent DEscRIPTIoN.-Thallus adnate on bark, pale tur- closely related depsides. tle green, 5-10 cm in diameter; lobes subirregular The parent morph of P. horrescens is probably and apically rotund, 2-6 mm wide, the marginal extinct. Chemically identical P. damaziana, a non- cilia mostly in the axils; upper surface plane, con- isidiate Brazilian species, is larger and more robust tinuous, often pruinose near the tips; medulla although obviously from the same stock as the sulphur yellow; lower surface densely rhizinate, parent of P. horrescens. Among the parallel morphs, short rhizinate and pale brown along the margins, both pustulate-sorediate P. subfatisceiis and lobulate the rhizines simple or squarrosely branched. Apoth- P. schindzeri are very close in lobe configuration, ecia numerous, adnate, the disc pale, 2-7 mm in adnation, and thallus texture. diameter; spores 8, 4-6 X 7-12 pm. SPECIMENSEX&MINED.-United States: Illinois, Skorepa 4627 CmmsmY.-Cortex K + yellow (atranorin); NUMBER 33 33

FIGURE16.4pecies of Parrnelina: a, P. irnrniscens (Andne 4456 in US); b, P. indica (Hale 43884); c, P. irrugans (Kurokauu 55342); ti, P. jarnesii (Du Rieti 3117:3 in US); e, P. leucotyliza (Hale 29402); f, P. lindrnnnii (Maline 450 in S). (Scale in mm.) 34 SMITHSONIAN CONTRIBUTIONS TO BOTANY medulla K+, C+, KC+ more intensely yellow, 21. Parmelia irrugans, new combination P - (unidentified yellow pigments). DIsTRIsuT1oN.-Mexico. FIGURE16c REMARKs.-Parmelina immiscens is the only spe- Parmelia irrugans Nylander, 1890:26 [type collection: Simon- cies in the genus endemic to Mexico. It occurs oseki, Japan, A lmquist (H, Nylander herbarium number rather rarely on trees in open oak-pine forests in 35551, lectotype; isolectotype in S)]. the arid highlands (2000-2400 m elevation). Al- Parmelia insinuata Hue, 1899: 158 [type collection: Ta-Pin- though the medulla is distinctly yellow, no terpenes Tze, Yunnan, China, Delavay 3008 (PC, lectotype); not P. insinuata ”).lander, 1856: 3243. are produced. The pigment seems to be secalonic Parmelia xanthocarpa Hue, 1899:178 [type collection: Ta- acid A but chromatographic tests are inconclusive. Long-Tan near Ta-Pin-Tze, Yunnan, China, Delavay 23 This species and its presumptive isidiate morph (PC, lectotype)]. P. Zindmanii, therefore, are unrelated to P. sub- Parmelia insinuatula Zahlbruckner, 1929: 169 [based on P. aurulenta and similar Asian species with a yellow insinuata Hue]. medulla. DEsCRIPTIoN.-Thallus adnate to loosely adnate,

SPECIMENS EXAMINED.-MeXicO: Durango, Cramer 1998 corticolous, firm, 4-10 cm broad; lobes sublinear (KAN, US); Jalisco, Pringle 10706 (US); Oaxaca, Hale 20825. to subirregular with more or less rotund apices, contiguous, 2-5 mm wide, the marginal cilia dis- tinct, mostly in the axils, to 0.8 mm long; medulla 20. Parmelina indica, new species pale orange or yellow; lower surface black, densely rhizinate, the rhizines simple or becoming squar- FIGURE166 rosely branched. Apothecia very common, adnate to sessile, 3-10 mm in diameter; spores 8, 8-10 X DEscRIPTIoN.-Thallus arcte adnatus, saxicola, 12-15 pm, often poorly developed. obscure albo-cinereus, 2-3 cm latus, lobis sub- CHEMIsmY.--Cortex K + yellow (atranorin); linearibus, contiguis, 0.8-1.2 mm latis, ciliis mar- medulla more intensively yellow with color tests gine irregulariter dispersis, simplicibus, 0.1-0.3 mm (zeorin, leucotylic acid and associated terpenes, and longis, superne planus, continuus vel rimosus, secalonic acid A). nitidus, modice vel dense isidiatus, isidiis cylin- DIsmIsuTIoN.-China and Japan. dricis vel leviter inflatis, praecipue simplicibus, REMARKS.-This species has not been recognized usque ad 0.3 mm altis; cortex superior 12 pm by contemporary lichenologists or included in lists crassus, epicorticatus, stratum gonidiale 15-18 pm of Asian Parmelinae. It is characterized by the crassum, medulla alba, 100-120 pm crassa, cortex presence of the “aurulenta” terpene series, the thin, inferior paraplectenchymatus, 12-14 pm crassus; expanded thallus, and large apothecia (3-10 mm in subtus niger, dense rhizinosus, rhizinis nigris, sim- diameter), invariably larger than those of P. sub- plicibus, 0.4 mm longis. Apothecia ignota. aurulenta (3 mm or less). The spores are slightly CHEMIsmY.-Cortex K + yellow (atranorin); but not significantly larger than those in P. SUB- medulla negative with color tests zeorin, leucotylin aurulenta. As a rule, the size of apothecia will and traces of associated “subaurulenta” terpenes). separate the two species. Two of Nylander’s syn- HOL0TYPE.-on rocks in river below Silver Cas- types of P. subaurulenta (see “Specimens Exam- cade, Kodaikanal, Tamil Nadu, India, elevation ined” below) are actually P. irrugans. about 1800 m, M. E. Hale 43884, 24 January 1975 Parmelina irrugans has a more restricted range (US: isotype in Poona). than P. subaurulenta. It does not occur in India DIsmIsuTIoN.-India. where P. subaurulenta is so common. It is, how- REMARKS.-This small saxicolous lichen was col- ever, more frequent in Japan than one might lected on a large rock outcropping in an undis- expect from the specimens examined because I did turbed stream bed. Externally it is close to the not borrow the specimens at TNS for redetermina- American P. obsessa but lacks both yellow pig- tion. Many annotated by me in 1964 in Tokyo as ments and galbinic acid. It is the only terpene- P. homogenes are probably this species. containing Parmelina without pigments. SPECIMENSEXAMINED.-China: Montiguy (H. Nylander her- NUMBER 93 35 barium number 35668, a syntype of Parmelia subaurulenta [type collection: Inokashira, near Tokyo, Japan, Asahina Kllander); Hunan, Handel-Mazzetti 2469 (US, \V); Yunnan, 23a (W, lectotype)]. Delaruy (H, K)lander herbarium number 35666). Japan: Parmelia leucotyliza f. rugulosa Asahina, 1952:94 [tvpe col- illaries (H, Sylander herbarium number 35667, probable syn- lection: Mt. Higane, Prov. Suruga, Japan, Yamashita 17 type of Pormelia subaurulenta Nylander); Prov. Kii, Kuro- (TNS, lectotype)]. kawn 59064 (TNS, US); Prov. Kikuchu, Kzirokazua 59322 Parmelia leucotyliza f. sziblaevis Asahina, 1952:95 [type collec- (TNS, US); Prov. Kozuhe, Kurokazua 58581 (TNS, US); Prov. tion: Kadoike, Pro\.. Suruga, Japan, Asahina (TSS, lecto- Mutsu, Kurokawa 550341, 55342, 56162, 58582 (TNS, US); type)]. Prov. Ohmi, Hale 29456, 29470; Prov. Rikuchu, Kurokawa 59322 (TNS, US); Prov. Shimane. Omura 23 (US); Prov. DEscRIPTIo;u.-Thallus adnate to loosely attached Suruga, Culberson 10738 (DUKE, US). on bark or rocks, light greenish mineral gray, 8-12 cm broad; lobes sublinear to subirregular, con- tiguous, 2-4 mm wide, the marginal cilia mostly 22. Parmelina jamesii, new combination in lobe axils, about 0.5 mm long; upper surface FIGURE16d shiny or becoming dull white pruinose at the tips, plane but soon becoming pustulate, the pustules Parmelia jamesii Hale, 1972b:179 [type collection: Welling- breaking open and coalescing into large clumps ton, New Zealand, James SZ2118 (US, holotype; isotypes in BM, TNS)]. without formation of soredia; medulla very pale salmon colored to white; lower surface black ex- DEscRIPTros.-Thallus adnate on bark, rather cept for a dark brown zone at the tips, densely fragile, whitish mineral gray, 5-10 cm broad; lobes rhizinate, the rhizines black and shiny, simple or sublinear, contiguous, 1.5-3.0 mm wide, the mar- sparsely squarrosely branched. Apothecia rare, sub- gins sparsely ciliate, the cilia irregularly dispersed, stitpitate, the amphithecium pustulate, 1-3 mm in long; upper surface plane but becoming rugulose diameter; spores 8, 6 X 11-12 pm. with age, shiny or becoming opaque and white- CHEmsTRY.-Cortex K + yellow (atranorin); pruinose at the tips, sometimes faintly white- medulla more intensively yellow with color tests reticulate, moderately isidiate, the isidia cylindrical, (zeorin, leucotylin and related “subaurulenta” ter- thin, erect, up to 0.3 mm high; lower surface black, penes, secalonic acid A, and traces of unidentified moderately rhizinate, the rhizines simple or sparsely substances). squarrosely branched. Apothecia unknown. DIsTRIBuTIoN.-Japan and hlalaysia (Sabah). CHEmsTRY.-Cortex K+ yellow (atranorin); REMARKS.-Originally when studying the Jap medulla K-, C-, KC-, P+ red (fumarproto- anese Paymeliae, I synonymized P. leucotyliza under cetraric acid and a trace of protocetraric acid). P. aurulenta. These two species, however, are differ- DIsTRIBuTI0N.-Eastern Australia and New ent in several important respects. First, they have Zealand, different terpene chemistry. Second, the pustules of REMARKs.-Parmelina jamesii is the only species Paymelina leucotyliza do not become sorediate, producing ,B-orcinol depsidone fumarprotocetraric whereas those in P. aurulenta are often densely acid. It is also one of the few species endemic to sorediate. Lastly, P. leucotyliza is restricted to Japan the Australian region. It might be mistaken super- except for a tentatively identified specimen from ficially for a Hypotrachyna except that the lobe margins are distinctly ciliate and the rhizines Sabah. It seems to be much more common than simple. P. aurulenta in Japan, occurring on rocks and trees in open forests, trees along roads, and rocks in rice fields. 23. Parmelina leucotyliza SPECIXIE~SExAaIisED.-Japan: Prov. Aki, Hale 29535, 29445, 29544; Prov. Hyuga, Hale 29677; Prov. Iwaki, Kurokarua FIGURE16e 58079 (TNS, US); Prov. Izu, Asahina in Lichenes Japoniae Parmelina leucotylizn (Sylander) Hale, 1974:482. Exsiccati 139 (US), Kurokawa 58004, 58600 (TNS, US); Prov. Parmelia leucotyliza Nylander, 1890:27 [type collection: Rock- Kazusa, Kurokawa 59002 (TSS, L’S); Prov. Kii, Kurokawa osan, Japan, Almquist (H, Nylander herbarium number 59087, 59131, 59132 (TNS, US); Prov. Musashi, Kurokawa 35196, lectotype: isolectotype in S)]. 51040 (TNS, US); Prov. Settsu, Hale 29402, 29433; Shizuoka Parmelia fmudans ssp. subfraudans Zahlbruckner, 1927:352 Prefecture, Omura 157 (US). Malaysia: Sabah, Hale 29102. 36 SMITHSONIAN CONTRIBUTIONS TO BOTANY

24. Pamtelina lindmanii Parmelia inelanochaeta Kurokaira in Hale and Kurokawa, 1964:133 [type collection: Santa Anna de Chapada, Mato FIGURE16f Grosso, Brazil, Malme 2243 (S, lectot) pe: isolectotypes in Parmelina lindmanii (Lynge) Hale, 1974:483. UC, US)]. Parrnelia lindrnanii Lynge, 1914:74 [type collection: Porto DEscRIrTIos.-Thallus adnate on bark, turning Alegre, Rio Grande do Sul, Brazil, Maime 450 (S, lecto- olive-buff to cream-buff in the herbarium, 4-7 cm type)l. Parmelia tiliacea var. sulphurosa f. asperata Miiller Argovien- in diameter; lobes irregularly branched, sometimes sis, 1883:46 [type collection: Argentina, Balansa 5 (G, sublinear-elongate, 2-6 mm wide, the margins more lectotype)]. or less crenate, ciliate, the cilia black, mostly simple, DESCRIPTION.-Thallus adnate on bark, pale buff 1-2 mm long; upper surface shiny, maculate, mod- to yellowish mineral gray, 5-8 cm broad; lobes sub- erately to densely isidiate, isidia thin, cylindrical, irregular, apically rotund, 3-5 mm wide, the mar- usually branched, often with black spinules or short ginal cilia mostly in lobe axils, less than 0.5 mm cilia; medulla white; lower surface black, dark to long; upper surface plane to rugose with age, shiny, pale brown in a rather wide zone near the tips, moderately to densely isidiate, the isidia cylindrical, moderately rhizinate, rhizines black to blackish unbranched, to 0.3 mm high; medulla uniformly brown, simple. Apothecia adnate to substipitate, pale sulfur-yellow tinged with orange; lower sur- 1-3 mm in diameter, the amphithecium isidiate, face black except for a narrow marginal brown spinulate, the disc Vandyke-brown; spores 8, 8-10 x zone, moderately rhizinate, the rhizines black, sim- 13-15 pm. ple. Apothecia rare, sessile, 2-5 mm in diameter; CHEiwsTRY.-Cortex K + yellow (atranorin); me- spores 8,6-10 X 9-14 pm. dulla K-, C+, KC+ rose, P- (gyrophoric acid). CHExmTRY.-Cortex K + yellow (atranorin); DIsTRIBuTIoN.-South America. medulla more intensely yellow with K and C, P- REMARKS.-This species is very close to P. dissecfa (probably secalonic acid A and unidentified in general morphology and has the same chemistry. pigments). The lobes, however, are much broader, the isidia DISTRIBUTION.-RleXiCO and South America. ciliate, the marginal cilia long and distinct, and REhlARKS.-This corticolous lichen is character- the upper cortex strongly white-maculate. It occurs ized by the pale yellow-orange medulla and the in a rather restricted area from Brazil into Para- isidia. It is probably the isidiate morph of the guay with one specimen recorded from Colombia. Mexican endemic P. immiscens. When I first began SPECIMF\SExA\IINED.-Colombia: Cundinamarca, Flenniken study of Pal-nzelia, I confused Pal-melina lindmanii 1745 (US). Brazil: Mato Grosso, Malme 2397B (S). Paraguay: and Pal-melia endosulphur-ea ( = Parmotl-ema endo- Asuncih, Schinini 17333 (M\'M): Gran Chaco, hlalme (S). sulphur-eum (Hillmann) Hale) (Hale, 1960:20). Both Additional records from Brazil and Paraguai are listed in species have a yellow-orange medulla and isidia, but Hale and Kurokana, 1964:133. P. lindmanii has more adnate lobes, cilia in the axils, and no gyrophoric acid. Both species lack 26. Parmelina metarevoluta terpenes. FIGURE17b SPECIMENSEx.4JrINEo.-Mexico: Queretaro, Nakanishi 257 (US); Tamaulipas, Nakanishi 106 (US): \'era Cruz, Hale Par??7e[i?7arnetaraloluta (Asahina) Hale, 1974:483. 19389, 19685. Colombia: Antioquia, Nee and Mori 4238 (US). Pnrmelia tnetarevolirtn Asahina, 1960:97 [type collection: Venezuela: Merida, Rodrigues 168 (US). Brazil: Rio Grande AzusaJama, Prov. Shinano, Japan, h'uno and Kurokawa do Sul, Lima 3 (US). Paragua:; Caacupe, Balansa (G, H). 59243 (TSS, lectot) pe; isolectotype in US)]. Uruguay: Durazno, Osorio 2772 (DUKE, UPS): Lavalleja, Osorio 3822 (MVM, US): Paysandu, Lamb (BM, H); Soriano, DEscRIPTIo;u.-Thallus adnate on bark or rock, Rosengurtt (H); Trinidad, Osorio 3603 (LD). Argentina: pale olive-buff, 2-6 cm in diameter; lobes dicho- James (BM): Buenos Aires, Santesson 58, 100, 446 (S). tomously branched, sublinear-elongate, more or less ascending at the tips, 1-4 mm wide, lobules 25. Parmelina melanochaeta sometimes present on older lobes, the margins more FIGURE17a or less crenate, narrowly black rimmed, ciliate, the Parmelina nielanochaeta (Kurokawa) Hale, 1974:483. cilia black, shiny, simple, 0.2-0.5 mm long; upper NUMBER 33 37

FIGURE17.4pecies of Parinelina: a, P. melanochaeta (21ialnie 2243 in S); b, P. inetarevoluta (Kurokawa 59180 in US); c, P. niiielleri (Hale 42277): d, P. obsessa (Hale 19186); e, P. pnsr~llt- fera (specimen in US); f, P. perisidrans (Togashi in US). (Scale in mm.) 38 SMITHSONIAN CONTRIBUTIONS TO BOTANY surface smooth and shiny, not maculate, more or less granular, the soralia round, separate; lower less rugose on older lobes, sorediate, soralia sub- surface uniformly black, moderately rhizinate, the terminal, capitate, pale orange below; lower surface rhizines simple, black. Apothecia rare, sessile, the black, densely rhizinate; the rhizines black, shiny, amphithecium sorediate, disk brick red, 2-5 mm in simple, about 0.5 mm long. Apothecia very rare, diameter; spores 8,7-I0 X 12-15 pm. adnate, 1-5 in diameter, the rim more or less CHEMIsTRY.-Cortex K + yellow (atranorin); me- crenate, sorediate, the disc dark brown; spores 8, dulla K+ persistent yellow, C-, KC-, P+ pale 7 X 10-12pm. orange (stictic acid with or without constictic acid). CmMIsmY.-Cortex K + yellow (atranorin); me- DISTRIRUTI0N.-Mexico and South America. dulla K + reddish, C -, KC -, P + orange (galbinic REMARKS.--When 1 first collected this species on acid, rarely constictic acid, trace of salazinic acid, shade trees in a coffee plantation in Venezuela, I zeorin, leucotylin and associated terpenes, and identified it tentatively as P. pilosa because of the secalonic acid A). laminal soralia and distinct white maculae. Chem- DIsTR1suTIoN.-Eastern United States, Japan, and ical tests, however, showed that it contained stictic China. acid and would have to be identified with P. REMARKL-ASpointed out by Kurokawa (1968b: muelleri. After comparing the two species in the 351) this is the sorediate morph of P. galbina. It herbarium, I concluded that P. pilosa is larger and was first collected and correctly identified in the more robust on the average, but the two are very United States by Dr. Satoshi Nakanishi in 1971. closely related. Vainio (1890) had in fact compared Two additional collections have since been identi- P. muelleii with Parmelia balansa f. sorediata (= fied, one of them (Plitt 236) filed under Physcia Parmelina pilosa) which he distinguished with a orbicularis, which is very superficially similar in negative K test. They have different geographic lobe configuration and capitate soralia. It has the ranges, P. muelleri being more common in northern same Arcto-Tertiary distribution pattern as P. gal- South America and possibly occurring at a higher bina but is extremely rare in North America. elevation than P. pilosa. There is no esorediate, stictic acid-containing parent morph comparable to SPECIMENSExAMmEu.-United States: Ohio, Wetmore 18051 (MI", US); Maryland, Plitt 236 (US); Tennessee, Nakanishi the P. consors-P. pilosa species pair. 238 (US). Japan: Prov. Bungo, Kurokawa 62329 (TR'S); Prov. SPECIMENSExAMINED.-Mexico: Michoackn, Wirth 329 (US); Ise, Asahina (TNS); Prov. Omi, Asahina (TNS); Prov. Vera Cruz, Hale 21124. Venezuela: MCrida, Hale 42219, Shinano, Asahina (TNS). Kurokawa 58326 (TNS); 59180 42230, 42256, 42277, Oberroinkler and Poelt 7529 (M, US); (TNS, US) 59243, 59244 (TNS); Prov. Suruga, Asahina (TR'S); Thchira, Hale 45716. Peru: Cuzco, lltis 3214 (WIS). Argen- Prov. Totomi, Asahina 160 (TNS); Prov. Yamashiro, Asa- tina: TucumPn, Culberson 14909 (DUKE, US). hina (TNS). China: Manchuria, Asahina 29 (TNS).

28. Parmelina obsessa 27. Parmelina muelleri FIGURE17c FIGURE17c Parmelina obsessa (Acharius) Hale, 1974:483. Parmelina muelleri (Vainio) Hale, 1974:483. Parmelia obsessa Acharius, 1814:195. [type collection: North Parmelia muelleri Vainio, 1890:49 [type collection: Sitio, America (?Pennsylvania), Muhlenberg (H-Ach, lectotype; Minas Gerais, Brazil, Vainio in Lichenes Brasilienses isolectotype in PH)]. Exsiccati 948 (TUR, Vainio herbarium number 2677, lecto- Parmelia finkii Zahlbruckner in Hedrick, 1934: 162 [type type; isolectotypes in BM, FH, M, UPS)]. collection: Williamsville, Wayne County, Missouri, Russell (W, lectotype; isolectotype in MICH)]. DEScRIPTIoN.-Tha11us closely adnate on bark, lichen green, turning honey yellow in the her- DEScRIPTIoN.-Tha11us closely adnate on rock barium, 5 cm or more in diameter; lobes irregularly (rarely on bark), pale smoke gray to light mineral branched, rounded at the tips, 2-3 mm wide, the gray, 3-6 cm broad; lobes irregularly branched, margins evenly ciliate, the cilia simple or rarely sublinear, contiguous, 1-2 mm wide, the margins branched, black, shiny, up to 1.5 mm long; upper dissected, sparsely ciliate, especially in axils, the surface strongly white-maculate, irregularly cracked cilia black, simple, to 0.3 mm long; upper surface on older lobes, sorediate, soredia laminal, more or densely isidiate, the isidia cylindrical, sometimes NUMBER 88 39 branched; medulla white to pale dull green-yellow; CHEMISTRY.-cOrteX K + yellow (atranorin); me- lower surface black, sparsely to moderately rhi- dulla K-, C+, KC+ red, P- (lecanoric acid). zinate, dark brown and short rhizinate near the tips, DIsTRrsuTIoN.-Great Britain and Europe. the rhizines simple. Apothecia adnate, 1-3 mm in REMARKS.-while this species has long been diameter, the disc brown, the amphithecium isidi- recognized as a distinct variety (Poelt, 1961:194), it ate; spores 8,4-5 X 6-10 pm. was only recently raised to species level. Dobson CHEmsTRY.-Cortex K + yellow (atranorin); me- and Hawksworth (1976) have compared it in detail dulla more intensively yellow with color tests with the normally isidiate P. tiliacea, finding that (zeorin, galbinic acid, trace of salazinic acid, leuco- P. pastillifera has, on the average, somewhat nar- tylin and associated terpenes, and secalonic acid A). rower lobes (2-6 mm) than P. tiliacea (3-8 mm) and DIsTRIsuTI0N.-Eastern United States. denser rhizines.2 Parmelina pastillifera also has a REMARKs.-Parme~ina obsessa is another Acharian somewhat smaller, slightly more bluish thallus. The species that had been lost from the literature since ecological differences are pronounced, for P. pastil- 1814. Not only were extremely few collections Zifera occurred in England west of the 813 mm available, but also many were incorrectly identified. rainfall isopleth and the 2OC January mean iso- One in US was listed as “Parmelia prolixa,” and as therm. This agrees with Schauer’s (1965:80) defini- recently as 1958 P. obses,ra was identified as Parme- tion of P. pastillifera as an oceanic species in lia finkii (Hale, 1958:82). Recent collecting by central Europe. American lichenologists, however, has shown that SPECIMFNS EXAMINED.-EUrOpe: France, Lambinon in it is extremely common on sandstone outcrops in Lichenes Selecti Exsiccati 363 (LD, US); Germany, Schroppel closed oak forests and can tolerate considerable in Lichenes Alpium 143 (US); Austria, Strasser in Crypto- shade. As a closely appressed saxicolous lichen it is gamae Exsiccatae Vindobonensi 3062b (US); Yugoslavia, not easy to collect. Kurokawa (1968b) correctly Vtzda in Lichenes Selecti Exsiccati 762 (US). recognizes it as the isidiate morph of P. galbina. No comparable isidiate population has evolved in 30. Parmelina perisidians Japan. FIGURE17f SPECIMENSEXAMINED (Selected).-New Hampshire, Willey (US): Pennsylvania, Hale 16288, 17486; Maryland, Hale Parmelia perisidians (Nylander) Hale, 1974:483. 14422; West Virginia, Hale 10311, 10660, 12408, 15003; Ken- Parmelia perisidians (h’jlander, 1900:6 [type collection: Ram- tucky, Hale 13866; Indiana, Hale 14182; Illinois, Hale 13935, podde, Ceylon, Almquist (S, 1ectot)pe; isolectotype in H, 14022; Michigan, Hale 33889; Wisconsin, Hale 23529: Min- Nylander herbarium number 35673)l. nesota, Hale 23602; Virginia, Hale 12018, 18803, Luttrell Parmelia subsulphuuata Asahina, 1951a:228 [t)pe collection; 3864 (US), Tucker 6588 (US); North Carolina, Culberson Higashi-Shirakawa, Prov. Mino, Japan, Yasue (TNS, lecto- 6491, 10418 (DUKE, US), Hale 16358; Tennessee, Hale 37083; type)l. South Carolina, Hale 7698; Georgia, Hale 7404, 30605; Ala- bama, Hale 7098; Louisiana, Hale 33862; Arkansas, Hale DEscRIPTIoN.-Thallus adnate on bark, greenish 2972, 4030; Kansas, Hale 4749; Oklahoma, Hale 4886, 5054, mineral gray, 3-6 cm in diameter; lobes sublinear, Keck 1493 (US); Texas, Hale 5452. 0.5-2 mm wide, the marginal cilia mostly in the axils; upper surface plane, continuous, densely isidiate, the isidia simple or branched; medulla 29. Parmelina pastillifera, new combination sea-foam yellow; lower surface densely rhizinate, FIGURE17e the rhizines black, simple or squarrosely branched. Apothecia adnate, 2-5 mm in diameter, the am- Parmelia scortea var. pastillifera Harmand, 19103558 [type collection: Bussang, Vosges, France, Claude1 and Harmand phithecium isidiate; spores 8,5-7 X 7-1 1 pm. in Lichenes Gallici Exsiccati 491 (H, lectotype; isolectotypes CHEMISTRY.-cOrteX K + yellow (a tranorin); me- in BM, BPI)]. dulla K+, C+ KC+ more intensely yellow, P-, Parmelia pastillifera (Harmand) Schubert and Klement, (zeorin, leucotylin and related terpenes, and 1966:58. secalonic acid A). DEscRIPTIoN.-Thallus largely as in P. tiliacea (p. 48) except isidia dark tipped, peltate, and *I wish to thank Dr. Hawksworth for allowing me to see apically flattened (Figures 3e,f, 4b). his manuscript before publication. 40 SMITHSONIAN CONTRIBUTIONS TO BOTANY

DIsTRIBuTI0N.-Southeast Asia from India to of norstictic acid, and small spores. It is closely re- Japan. lated to isidiate P. antillensis but is not the direct REMARKS.-This densely isidiate species is closely parent morph since P. antillensis has a continuous related to P. amagiensis and may represent its isidi- cortex. It is not, as implied in my original descrip- ate morph, although differences in pigments, so far tion, a member of Amphigymnia (= Parmotrema) unresolved, may exclude this possibility. Both because the thallus is adnate overall, cilia are pro- species have the typical “subaurulenta” terpene duced in the axils, and the bare zone at the series. Parmelina perisidians has a broad distribu- margins below is very narrow. Both P. phlyctina tion in the higher elevation forests of tropical Asia and P. antzllensis, however, constitute an anomolous and the temperate forests of Japan. element in Parmelina, not only because of the somewhat amphigymnioid lobation but also be- SPECIMENS ExAMIKED.-India: Karnataka, Patwardhan 74.3283 (Poona, US): Tamil Nadu, Hale 40101, 40.102, Togashi cause of the production of norstictic acid. (TNS, US). Sri Lanka: see Kurokawa and Mineta (1973:74) SPECIW ?s Exs~i~\k~.-hfexico:Chiapas, Hale 20202 (S, for records in Ceylon. Thailand: Kurokawa 1818 (TNS, US). US), Cuba: Oriente, Imshnug 27174 (MSC, US), 25146, 25148 Philippines: Rlountain Province, Hale 26166a. Japan: Prov. (MSC), ,\lorton 9394 (US). Jamaica: Imshaug 14614, 14714, Kozuke, Asahina (TNS); Proc. Ifusashi, Asahina (TNS); 15211, 15362, 15375, 15458, 15502, 15524, 15536 (MSC), 13179, Kurokawa 550009 (TKS, LS), 64284, 64285 (TNS); Prov. 14727, 15146, 15453, 15513 (MSC, US). Haiti: Sud, Wetmore Settsu, Togashi (TNS, US). 3373 (MSC). Dominican Republic: Cordillera Central, Wet- more 3728 (LD, MSC), 3738 (MSC, US). Puerto Rico: Imshaug 29562 (MSC). 3 1. Parmelina phlyctina

FIGURE18a 32. Parmelina pilosa

Parmelina phlyctina (Hale), Hale, 1974.483. FIGURE18b Parmelia phlyctina Hale, 1959:129 [tjpe collection, Blue Parmelina pilosa (Stizenlxrger) Hale, 1974:483. Mountains, Jamaica, lnzshaicg 14908 (MSC, holotjpe; iso- Parmeha pilosa Stizenberger, 180: 165 [type collection: qpe in US)]. Rhenoster Ricer, Taaibosch Kranz Mountains, Orange Free State, Union of South Africa, Rehmann (ZT, lecto- DEscRIPTIoN.-Thallus adnate to loosely adnate type; isolectotype in H)]. on bark, membranous and fragile, light mineral Parrnelin halansae var. sorediata Muller Argoviensis, 1888a:Z gray but turning pink in the herbarium if improp- [tjpe collection: Montevideo, Uruguaj, Arechavaleta (G, erly dried, 5-15 cm broad; lobes subirregular, lectotype)]. apically rotund, 5-10 mm wide, the margins entire Parnielia subbalansae Gyelnik, 1931:288 (based on Parmelia balansae ~ar.sorediata Miiller Argoviensis. to lobulate, sparsely ciliate in the lobe axils, the cilia up to 0.5 mm long; upper surface plane, DEscRIPTIoN.-Thallus adnate to loosely attached smooth to rugose in older parts, upper cortex on bark, coriaceous, light buff mineral gray, 5-12 fragile and flaking off in small pieces, isidia and cm broad; lobes subirregular, apically subrotund, soredia lacking; lower surface black except for a the marginal cilia coarse, becoming furcate, 0.5-1 .O narrow brown zone at the tips, sparcely rhiAnate, mm long; upper surface shiny and heavily white- the rhizines simple. Apothecia rare, adnate, 3-8 mm maculate, plane to minutely pitted, sorediate, the in diameter; spores 8, 3-5 X 7-8 pm. soralia orbicular, about 0.5 mm in diameter, sepa- CHEMIsmY.-Cortex K + yellow (atranorin); me- rate or coalescing into extensive lamina1 sorediate dulla K+ yellow turning red, C-, KC-, P+ areas; medulla white; lower surface black, densely orange (norstictic and connorstictic acids). rhizinate, coarse and fine rhizines intermixed, DISTRIBUTION.-hkXiCO and the West Indies. simple to sparcely furcate. Apothecia rare, sub- REMARKS.-sinCe my discription of this species in stipitate; spores not seen. 1959, additional collections have been made in CHE;MIsTRY.-Cortex K + yellow (atranorin); me- Mexico, Cuba, and other islands in the West Indies. dulla negative with color reagents (no substances It occurs in wet upland rain forests or secondary demonstrated). mist forests. The main distinguishing features are DISTRIRLTIoN.-~outh America and southern the fragile thallus with a flaking cortex, presence Africa. NUMBER 33 41

FIGURE18.-Species of Parmelina: a, P. phlyctina (Imshaug 13179 in US); b, P. pilosa (McOwan in US); c, P. pruinata (Kurokawa in Lichenes Selecti Exsiccati 81 in US); (1, P. quercina (“car- porrhizans” population) (Schroppel and Poelt in Lichenes Alpiurn 8 in US); e, P. quercina (Tavares 1082 in US); f, P. rhytidodes (Kurokawa 58601 in US). (Scale in mm.) 42 SMITHSONIAN CONTRIBUTIONS TO BOTANY

REMARKs.-Parmelina pilosa is the sorediate The marginal cilia are sparse and sometimes diffi- morph of P. consors. It occurs on a variety of trees cult to determine. It could, in fact, be misidentified (Celtis, Erythz’na, Scutia), even on fence posts, and as a Pseudoparmelia but no corticolous nonisidiate in open or secondary habitats up to 2000 m eleva- species of that genus with lecanoric acid are pres- tion. It must be distinguished carefully from P. ently known. muelleri, which rarely occurs in this range and con- SPECIMENSExAMINED.-Australia: Australian Capital Ter- tains stictic acid. ritory, Kurokawa in Lichenes Rariores Selecti Exsiccati 81 SPECIMENSEXAMINED.-Ecuador: Pichincha, Wiggins 38945 (US), Leuthner (US); New South Wales, Boorman 1387, (COLO). Uruguay: Canelones, Felippone 434 (G); Durazno, L1389 (NSW), Doing (L, US), McVean 6358 (COLO), Weber Osorio 2979 (DUKE); Flores, Osorio 3519 (MVM, US); and McVean L-47913, L-49886 (COLO, US). Wilson 96 Florida, Osorio 1299 (F); Minas, Osorio 2049, 2056 (MVM, (NSW); Queensland, Gwytherl (BM); South Australia, Rogers US); Soriano, Rosengurth (H). Argentina: Buenos Aires, 886 (US); Tasmania, Bratt 1380b (US), James 2117 2125 Grassi 719 (US), Santesson 75 (S), Schnyder (G), Venturi (BM, US), Weymouth 65 (BM, US), 432 (BM); Victoria, Fil- 2853 (SI); Cordoba, Giardelli 1025 (SI), Petersen 23 (S); Entre son 6639, 7004 (US), James 223 (BM, US). New Zealand: Rios, Santesson 126 (S, US); Salta, Fries 21 (S), James (BM), Mason 90 (BM). Lorentz (M). Chile: Valparaiso, Santesson 3084 (S). Kenya: Nyanza Province, Maas Geesteranus 4957 (L, US). Rhodesia: Hoeg (TRH). Union of South Africa: Basutoland, KofEer 34. Parmelina quercina (LD); Cape Province, Hoeg (LD, TRH), Kofler (LD), Mc- FIGURE18d,e Owan (BM, US); Natal, Hoeg (TRH). Parmelina quercina (Willdenow) Hale, 1974:483. Lichen quercinw Willdenow, 1787:353 [type collection: Thier- 33. Parmelina pruinata garten and near Tegel, Germany (not seen; illustrated in pl. 7: fig. l3)]. 18c FIGURE Imbricaria quercina (Willdenow) de Lamarck and de Can- Parmelina pruinata (Muller Argoviensis) Hale, 1974:483. dolle, 1805:389. Parmelia pruinata Muller Argoviensis, 1883:46 [type collec- Parmelia carporrhizans Taylor, 1847:163 [type collection: tion: Yorke Peninsula, Australia, Tepper (G, lectotype)]. Canary Islands, Lemann (FH-Tayl, lectotype)]. Parmelia tiliacea var. afixa Stirton, 1899:485 [type collection: Parmelia sinuosa var. hypothrix Nylander, 1856:301 [type Queensland, Australia, Bailey (?) 376 (BM, lectotype)]. collection: Pyrenees, France, Nylander (H, lectotype)]. Parmelia atricha Nylander, 1873:271 [type collection: La DEsCRIPTIoN.-ThaIIus closely adnate on twigs, Preste, Pyrenees, France, Nylander 16 (H, lectotype (not ashy white, 1-5 cm broad; lobes sublinear-elongate, seen); isolectotypes in FH, BM)]. variable, 1-3 mm wide, marginal cilia sparse, mostly Parmelia tiliacea var. hypothrix (Nylander) Muller Argovien- sis, 1888b:196. in lobe axils, up to 0.5 mm long; upper surface Parmelia revoluta var. grantilata Harmand, 1897:221 [type plane to convex, rugulose, more or less pruinose; collection: Docelles, Vosges, France, Harmand 284 (DUKE, medulla white; lower surface black, sparsely rhi- lecto type)]. zinate, the rhizines black, simple. Apothecia con- Parmelia quercina (Willdenow) Vainio, 1899:279. spicuous, adnate, 24 mm in diameter, the disc Parmelia budapestinensis Gyelnik, 1932b3212 [type collection: Budapest, Hungary, Timkd (BP, holotype)]. often pruinose; spores 8,7-8 X 10-12 pm. Parmelia carporrhizans f. malicola Gyelnik, 1932b:212 [type CHEMIsTRY.-Cortex K+ yellow (atranorin); me- collection: St. George, Hungary (Austria?), Zahlbruckner dulla K -, C +, KC + red, P - (lecanoric acid). (BP. holotype)]. DISTRIBUTION.-Australia and New Zealand. Parmelia yalungana Zahlbruckner, 1934b:206 [type collec- REMARKS.-This species is rather variable in lobe tion: between the Litang and Yalung rivers between Muli Gomba, Baurong, and Wa-Erh-Dje, Szechuan, China, Rock configuration and width but typically grows closely 16720 (W, lectotype)]. adnate on small twigs and branches of trees in arid scrub land. It has no apparent relation with P. DEscRIPTIoN.-Tha1Ius closely adnate on bark, quercina, the only other nonsorediate, lecanoric rarely on rock, rather coriaceous, light olive-gray, acid-containing species in the genus, characterized sometimes turning olive-buff in the herbarium, by much coarser lobes and white maculae in the 5-10 cm broad; lobes dichotomously or irregularly cortex. Muller mentions “subtus alba” in his origi- branched, sublinear, separate or becoming im- nal description, but he was actually describing a bricate, 1.5-4.5 mm wide, the margins smooth to Physcia species growing intermingled with the type. more or less crenate, often narrowly black rimmed, NUMBER 33 43 moderately to densely ciliate, the cilia black, simple, Asahina (TNS); Prov. Tosa, Kurokawa 64091 (TNS), 64094 0.2-1 mm long; upper surface smooth and shiny, (TNS, US). Australia: New South Wales, Du Rietz 577b faintly to strongly white-maculate, irregularly (UPS, US), Flockton 884 (US). Mostly with Retrorse Rhizines on the Apothecia: United rugose and cracked on older lobes, sometimes partly States: See W. Culberson (1961:172) for a list of specimens pruinose, isidia and soredia lacking; lower surface examined and a distribution map. Europe: England, Borrer black, densely rhizinate, the rhizines black, shiny, (US), Crombie (BM, US), Holl 15 (BM),James 75 (BM): simple or more rarely squarrose, 1-3 mm long. France, Crozals (US), Rondon in Lichenes Selecti Exsiccati Apothecia numerous, substipitate, 1.5-5 mm in 268 (US); Spain, Bauza (LD); Portugal, Tavares in Lichenes Lusitaniae Selecti Exsiccati 217 (CS); Germany, Poelt in diameter, the amphithecium smooth or retrorse- Lichenes Alpiurn 8; Switzerland, Barkman 4294a (UPS): Aus- rhizinaee, the disc burnt umber; spores 8, 5-7 x tria, Stezner in Crjptogamae Exsiccatae Vindobonensi 4134 6-10 pm. (F, NY); Italy, Alrnborn (LD), Baunigartner in Cryptogarnae CHEmsmY.--Cortex K + yellow (atranorin); me- Exsiccatae Vindobonensi 3165 (NY, US). Tunisia: Runeinaik dulla K-, C+ KC+ red, P- (lecanoric acid). (LD). DISTRIBUTION.-califOrnia, Europe, Pakistan, Ne- pal, eastern Asia, and Australia. 35. Parmelina rhytidodes, new species REMARKS.-ASa common member of the Euro- pean flora this was one of the first FIGURE18f species of Parmelia to be described. The records from eastern North America previously identified as DEscfuPTIo;u.-Thallus adnatus ve1 laxe adnatus, Parmelia quercina are now recognized as Parmelina corticola et saxicola, pallide olivaceo-cinereus, 8-12 galbina or Hypotrachyna Zivida (Taylor) Hale. The cm latus, lobis subirregularibus, imbricatis con- interesting distribution pattern, centered in western gestisque, 2-4 mm latis, margine irregulariter North America and Europe, is typical of the Medi- ciliatis, ciliis nigris, 0.3-0.6 mm longis; superne terranean type, although the species occurs less nitidus vel apicem versus albo-pruinosus, primum commonly in eastern Asia and Australia. Still this planus sed mox dense et omnino rugosus (Figure is a remarkably broader range than its isidiate 6b), rugis non eruptentibus, cortice integro vel rare morph P. filiacea. rumpente; cortex superior 12-14 pm crassus, epicor- The development of rhizines on the lower side of ticatus, stratum gonidiale 14-16 pm crassum, me- the apothecia has been used to justify a separate dulla pallide salmonea, 110-150 pni crassa, cortex population, called Parmelia carporrhizans. As W. inferior brunneus, paraplectenchymatus, 15-17 pm Culberson (1961: 168) points out, however, virtually crassus; subtus niger, dense rhizinosus, rhizinis nigris all specimens in California have these rhizines, and vel apice brunneis, simplicibus vel sparse squarroso- many European specimens identified as P. quercina ramosis. Apothecia numerosa, sessilia vel substipi- are often found to have them, sometimes only very tata, disco plano, 3-8 mm diametro; sporis 8, sparsely developed (Dobson and Hawksworth, 1976). 9-11 X 12-14,um. This trait does not seem to warrant specific status, CHEMIsmY.-Cortex K + yellow (atranorin); me- although Schauer (1965:71) found this population dulla more intensively yellow with color tests to have a strongly oceanic distribution pattern in (zeorin, leucotylic acid and related terpenes, seca- western Europe. The problem will have to be solved Ionic acid A, and traces of unidentified substances). through careful field studies in Europe. HOLoTYPE.-Amagi Pass, Prov. Izu, Japan, S. Kurokawa 58601, 1 December 1958 (US, holotype; SPECIMFNSEXAMINED (selected).-Without Retrorse Rhizines: isotype in TNS). Europe: Denmark, Rostrup (LD); Germany, Schroppel and DIsTRIBuTIoN.-Japan and Nepal. Poelt in Lichenes Alpiurn 7 (H, LD, US); Switzerland, Mereschkovsky (US), Seifriz (US); Austria, Eggerth in Flora REMARKS.-After I had examined specimens iden- Exsiccata Austro-Hztngarica 1542 (US); Czechoslovakia, Vdzda tified as Pa?melina entotheiochron with thin-layer in Lichenes Bohernoslouakiae Exsiccati 201 (LD, US); Hun- chromatography, there remained a small group with gary, Tirnkd in Lichenotheca 135 (H, LD); Spain, Schindler the “aurulenta” terpene profile. On further study 4303 (KR, US); Portugal, Tavares 1082 (US). Pakistan: Iqbal I found that these are morphologically different 180 (US). Nepal: Stainton 4098 (BM, US). China: Manchuria, Asahina 28 (TNS); Shensi Prov., Hugh (BM). Japan: Prov. from typical P. entotheiochroa in that the ridges Hyuga, Hale 29650, Kurokawa 65065 (TNS); Prov. Tango, which develop very densely do not burst open or 44 SSIITHSONIAX CONTRIBUTIONS TO BOTANY flake off. While P. rhytidodes can be recognized Lire 4e). They do not originate from isidia and for with practice from the external morphology alone, this reason P. schindleri cannot be considered as a chemical test is desirable for confirmation. The a modified form of P. horrescens, which may occa- species is probably not as common in Japan as P. sionally have cylindrical and flattened isidia inter- entotheiochroa, but I have not had the opportunity mixed. The species is named in honor of Dr. H. to re-examine the numerous specimens in TNS with Schindler, the first lichenologist to collect the chromatography. The collections made by Poelt in species since Vainio in 1885, although this region Nepal are tentatively placed here. They are very has been visited by many lichen collectors. Vainio densely rugose and have crowded lobes, modified had misidentified his exsiccate number 1284 as ecologically by the high exposed elevation where Parmelia coronata var. isidiosa Rluller Argoviensis they were collected. Parmelina rhytidodes is appar- (= Bztlbothrix fungicola (Lynge) Hale). ently an offshoot with sorediate P. aurulenta from SPFCIMENS ExAMIxED.-Brazil: Rio de Janeiro, Schindler P. irrugans or a now extinct parent morph similar 4569,4577 (KR, US). to it.

SPECIMENSExAMINED.-Japan: Prov. Awa, Kurokawa 56554 (TSS, US); Prov. Hyuga, Hale 29602; Hiroshima Prefecture: 37. Parmelina simplicior Nakanishi and Oshio 5731 (US); Kanto, Nurnariri 156 (US). FIGURE19b Nepal: Poelt L-747, L-754 (M, US). Parmelina simplicior (Hale) Hale, 1974:483. Parrnelia sirnplicior Hale, 1972a:99 [type collection: Panch- 36. Parmelina schindleri, new species gani, \Vestern Ghats, India, D. D. Awasthi 4056 (US, holo- type; isotype in Awasthi herbarium)]. FIGURE19a DEscRIPTIoN.-Thallus adnate on bark, buff min- DEscRIPrIoN.-ThallUs adnatus vel appressus, eral gray, coriaceous, 8-10 cm broad; lobes elongate, fragilis, corticola, pallide albo-cinereus, 2-4 cm more or less subirregular, contiguous and becoming latus, lobis sublinearibus, brevibus, contiguis, ca. imbricate, 3-5 mm wide, the axils sparsely ciliate, 1-1.5 mm latis, margine ciliatis, ciliis 0.2-0.4 nim the cilia 0.5 mm long; upper surface plane, contin- longis, simplicibus, margine et pro parte superficie LIOLIS, emaculate, isidia and soredia lacking; medulla lobulatis, lobulis congestis, suberectis, ramosis, 0.1- white; lower surface black, sparsely to moderately 0.2 mm latis et usque ad 1 mm longis, margine rhizinate, the rhizines simple, black. Apothecia breve ciliatis; superne nitidus, planus; cortex supe- common, sessile, 3-6 mm in diameter; spores 8, rior 14-15 pm crassus, epicorticatus, stratum gonidi- 4 X 6pm. ale 12 pm crassum, medulla alba, 50-65 pm crassa, cortex inferior paraplectenchymatus, 12 pm crassus; CHEMIsmY.--Cortex K + yellow (atranorin); me- subtus niger, modice rhizinosus, rhizinis nigris, sim- dulla K+ yellow turning red, C-, KC-, P+ plicibus vel sparse furcatis. Apothecia rara, sessilia, orange (salazinic acid). margine crenato, 3-4 mm diametro, sporis 8:nae, DISTRIBUTIoN.-~outh India. 10 X 16-18pm. REMARKs.-Parmelina simphior has a rather CHEMIsrRY.-Cortex K + yellow (atranorin); me- leathery thallus and very sparsely developed axil- dulla K-, C-, KC+ rose, P- (traces of gyro- lary cilia. As with many other Indian lichens it phoric acid and the “horrescens” unknown). does not exhibit clear-cut traits that enable one to HoLoTYPE.-Caraca, hhas Gerais, Brazil, I/ainio place it immediately in a particular genus. I had in Lichenes Brasilienses Exsiccati 1284 (BM; isotypes mentioned a possible relationship to isidiate P. in FH, RI, TUR, and UPS). wallichiana, which has much larger spores and a DIsTRIBuTIoN.-Brazil. more membranous thallus, but these two species are RExmKs.-The chemical constituents place this not at all related. Pal-melina simplicior grows on species in the “horrescens” group, where it is the roadside trees in the Western Ghats region of lobulate morph of a parent species represented by India where an intense monsoon season from June P. damaziana or a now extinct progenitor similar to September alternates with a long period of almost to it. The dense lobules are easily recognized (Fig total drought. Parmelina wallichiana, on the other NUMBER 33 45

FIGURE19.-Species of Purmelina: a, P. schindleri (I’ainio 1284 in BM); 6, P. .rirnplicior (Arcnstlzi 4056 in US); c, P. spathulato (Alnzborn 931B in US); d, P. spurnma (Halt, 42934a); e, P. subaw-ulenta (Hale 43705); f, P. subfatitcens (Hale 35091). (Scale in mm.) 46 SMITHSONIAN COSTRIBUTIONS TO BOTANY hand, occurs at higher elevations with less climatic surface moderately rhizinate, black, the rhizines stress. simple or furcate. Apothecia rare, adnate, 1-3 mm in diameter, the amphithecium pustulate; spores 8, SPECIMENSExAMIxED.-lndia: Maharashtra, Hale 40004, 7-8 X 12-14,um. 40007,40046, 40090,43972. CHmiIsTRY.-Cortex K + yellow (atranorin); me- dulla K-, C+ rose, KC+ red, P-, (gyrophoric 38. Parmelina spathulata acid, an unidentified pigment, and frequently a FIGURE19c white fluorescent spot). DIsTRInuTroN.-Pantropical at higher elevations. Parmelina spathulata (Kurokawa) Hale, 19743483. Parmelia spathulata Kurokawa in Hale and Kurokawa, REMARKS.-T~~Swidespread but rather rare 1964:133 [type collection: Skeleton Gorge, Wynberg, Union species is characterized by the dense, erupting but of South Africa, Alrnbortz 305 (LD, holotype; isotype in nonsoretliate pustules (Figure 5e,f) and the pale Wl. yellowish medulla. The main constituent, gyro- DEsCRIPTIoN.-Thallus adnate on bark, pale phoric acid, places it in the P. dissecta group, but whitish glaucous-green, fragile, 2-5 cm broad; lobes it probably represents a different line of evolution sublinear, crowded, 1-3 mm wide, the marginal from a now extinct parent morph (Figure 9). The cilia evenly dispersed, about 0.5 mm long; upper chemistry is somewhat aberrant from P. dissecta and surface plane, continuous, moderately isidiate, the related species judging by the presence of a white isidia initially cylindrical and erect but soon be- UV-fluorescent spot in more than half of the speci- coming procumbent and flattened; lower surface mens (including the lectotype) from both the New black, moderately rhizinate, the rhizines black, TVorltl and the Old World. The spot is not reactive simple to rarely furcate. Apothecia not seen. with H2S0, and its identity is not known. CmhiIsmY.-Cortex K + yellow (atranorin); me- Parmelina spurnosa has ecological requirements dulla K-, C+, KC+ rose to red, P- (gyrophoric similar to those of P. dissecta and P. horrescens. acid). Much less common at temperate latitudes than the DIsTRIBuTIoN.-South Africa. latter two, it grows throughout the tropics on REMARKs.-This lobulate-isidiate species (Figure exposed trees (hardwoods and conifers) and more 4d) is obviously a member of the P. dissecta group. rarely on rocks in forests up to 2500 m elevation. It occurs only in southern Africa and is much less SPEcIhfENs Ex.AarIxm.-Mexico: Chiapas, Hale 19897, 20110a, common than P. dissrcfn, which has smaller, normal 20150. Cuba: Hioram 6682 (US). Jamaica: Imshaug 14279, is idia . 15646 (MSC, US). Colombia: Antioquia, Nee and Mori 4258a (US). Venezuela: Merida , Hale 42957, 42934a, 43037a. SPrcihfEss Ex.iMlsED.-~nion of South Africa: Cape Pror- Brazil: Sao Paulo, Eiten and Mimura 5738 (US). Chile: ince, Kofler (LD, US). Additional records from South Africa Chiloe, Santesson 2264 (S, US): Valdivia, Santesson 7066 (S). are given in Hale and Kurokaiva (1964:134). St. Helena: Loveridge (AM). Union of South Africa: Cape Province, Almborn 749, 2663 (LD), 3966 (LD, US), Degelius SA-403 (Degelius herbarium, US); Natal, Almborn 10069 39. Parmelina spumosa (LD). Madagascar: Lemaitre (H). Indonesia: Java, Groenhart 2218 (L), Neemoort 3354 (BOR), Oka 4089 (L). India: Tamil FIGURE19d Nadu, Hale 43829. Japan: Prov. Aki, Hale 29529, 29540; Prov. Parmrlina spumosa (Asahina) Hale, 1974:483. Hizen, Kurokawa 63159 (TNS): Prov. Izu, Asahina (TNS): Parinelia spz~mosa Asahina, 1951b:259 [type collection: Prov. Mino, Asahiim (TNS); Prov. Ohmi, Hale 29472, 29482; Higashi-Murayama, Kita-Tatna-gun, Prov. Musashi, Japan, Prov. Sagami, Asahina 40 (TNS), Kurokawa 58038 (TNS, US); Asahina (TNS, lectotype)]. Prov. Shimofusa, Asahina (TNS); Prov. Survo, Asahina 125 (TNS). Taiwan: Kurokawa 494 (TNS). Australia: Victoria, DEsCRIPTIoN.-Thallus closely adnate on bark, Johnson (BM). New Zealand: Knight (UPS), Martin 493, 566 fragile, pale olive gray, 2-6 cm in diameter; lobes (AM). sublinear, 0.5-2 mm wide, the marginal cilia dis- tinct and evenly dispersed, about 0.5 mm long; 40. Parmelina subaurulenta upper surface plane, continuous, emaculate, densely pustulate-isidiate, the pustules bursting but not FIGURE19e becoming sorediate; medulla faintly yellow; lower Parmelina siibaurulenta (Nylander) Hale, I974:483. NUMBER 33 47

Parrnelia subaurulenta Nylander, 1885:606 [type collection: a significant difference in spore size, P. homogenes Narkanda, 5. W’. Himalayas, India, Skoliczka (H, Nylander must be regarded as a synonym of P. siibauixlenta. herbarium number 35672, lectotype)]. The species most often confused with P. sitbaum- Parmelia homogenes Nylander, 38853607 [type collection: lenfn, especially outside of India, is P. if-rugans, India, Hooker and Thotnson 1942 (H, Nylander herbarium number 35664, lectotype)]. which has consistently larger apothecia (up to 10 Parmelia conspicita Hue, 1899:1/15 [type collection: Kiao Che- mm in diameter), somewhat larger spores, and the Tung, above Kiang-l’n, Yunnan, China, Delavay (PC, “aurulenta” terpene series. T’wo of Nylander’s lectotype)]. syntypes for Parmelia sztbaurulenta are, in fact, Parrnelia homalotera Hue, 1899:159 [type collection: above P. imgans, as discussed above under that species. Mo-So-Yn, Yunnan, China, Delavay (PC, lectotype)]. Parrnelia fecrrnrla Hue, 1899:169 [type collection: Ta-Pin-Tze, A11 specimens should be tested with chromatog- Yunnan, China, Delavay 5 (PC, lectotype)]. raphy, although diameter of the apothecia is often Parmelia srcbcrernea Zahlbruckner, 1934b:208 [type collection: adequate to separate the species in this group. All Betiveen Muli Gomba, Baurong, and T\’a-Erh-Dje, Setch- type specimens listed in the synomymy above were Ivan, China, Rock 16720 pro parte (I%’,lectotype)]. tested with thin-layer chromatography except for Parmelia subaurulenta var. myriocarpa Asahina, 19.5la:227 Pnmelia coiispiczta Hue, which is placed here on [type collection: Mt. Koya, Prov. Kii, Japan, Inuinaru 1232 (TSS, lectotype)]. the basis of the small apothecia. Parmelia homogenes f. mitior Asahina, 1952:78 [type collec- tioil: Yamanaka, Prov. Kai, Japan, Asahitla (TNS. SPECIMEKSEx.iMIs.ro.-Sepal: Poelt 156 (hf). India: Sikkim, lect o t ype)]. Hnra et al. (TSS, US); Tamil Sadu, Degelius As352 (Degelius herbarium), Hale 40214, 43705, 43761, 43834, Parmelia rnyriocarpa (Asahina) Chao, 1964:149. Perinftet (H, Nylander herbarium number 35670, syntype of Parnielina hotnoge?res (Sylander) Hale, 1974:482 Parrnelia srcbaurdeiita), Watt 119 (BM): TTest Bengal, DE~CRIPTION.-T~~~~L~Sadnate to closely adnate on .4wnsthi 3919, 3921 (Aivasthi herbarium. US). Sri Lanka: bark, light greenish mineral gray, rather fragile, Almquist (H, Nylander herbarium number 35665). China: Yunnan, Handel-Mazzrtti 89, 2453, 2453 (US, JV), Rock 11747 4-1 0 cm broad; lobes sublinear, subimbricate, 2-4 (US). Taiivan: Kntrokawn 873, 1208 (TSS). Japan: Afoseley mm wide, the marginal cilia quite dense in the (H, Nylander herbarium number 35669, syntype of Parmelia axils, irregularly dispersed at the lobe tips, 0.4-0.7 s7tbaurulen/n); .4lrnquist (H, Nylantler herbarium number mm long; upper surface plane, faintly maculate, 36671). shiny, isidia and soredia lacking: medulla pale orange-yellow; lower surface black, densely rhi- 41. Parmelina subfatiscens zinate, the rhizines simple or becoming squarrosely branched. Apothecia very numerous, adnate, 1-3 FIGURE19f mm in diameter; spores 8, 6-10 X 8-14 pm, usually Parinelina subfatiscens (Kurolawa) Hale, 1974:483. abundantly developed Parmelfa siibfafiscens Kuroka\\a in Hale and Kurokawa, CHExmTRY.-Cortex K + yellow (atranorin); me- 1964:134 [t)pe collection: Louis Trichartlt, Zoutpansberg, dulla more intensively yellow with color tests Transiaal, Union of South Ahica, Al/nborn 6443 (LD, holotype; isotype in US)]. (zeorin, leucotylin and associated terpenes, secalonic acid A, and traces of unidentified substances). DEscRIPTIos.-ThallLis closely adnate on bark, DIsTRIiivTIoN.-Eastern Asia from India to Japan. fragile, whitish mineral gray, 3-5 cm broad; lobes RE:hfARKs.-Parmelina su bazirirlentn is character- sublinear-elongate, separate to contiguous, 03-15 ized by a thin thallus (average thickness for 20 speci- mm wide, the marginal cilia distinct, simple, to 1.0 mens 154 pm), numerous apothecia rarely exceeding mm long: upper surface plane, shiny, continuous, 3 mm in diameter, and well developed spores 6-9 x pustulate laminally and subterminally (Figure 5c), 8-12 pm. Nylander (1885:60i) described Paimelin exposed medulla in center of pustules turning black, homogenes at the same time on the basis of a single pustules remaining entire or producing very coarse specimen, citing as the main difference spore size sorediate or isidiate-sorediate masses, often short (“similis siibazmilenta . . . sed sporis majoribus”). ciliate in African material but bery rarely so in the Nylander measured spores 8-10 X 14-16 pm but my New 1Yorld; medulla white; lower surface black, own measurements on the type specimen gave densely rhizinate, the rhizines simple, black. 7-8 X 12-13 pm, putting it well within the range Apothecia rare, adnate, 1.5-4.0 mm in diameter; of spore size for Pal-rnelina subaurulenta. Without spores 8,8-9 X 12-14 pm. 48 SMITHSONIAN CONTRIBUTIONS TO BOTANY

CHEMISTRY.-cOrteX K + yellow (atranorin); me- sorediate; medulla white; lower surface black, dulla K -, C -, KC + rose, P - (trace of gyrophoric densely rhizinate, the rhizines black, simple. acid and the “horrescens” unknown substances). Apothecia unknown. DISTRIBUTIoN.-~outh Africa and the Caribbean CHEMIsTRY.-Cortex K + yellow (atranorin); me- region. dulla K-, C-, KC+ rose, P- (lobaric acid and REMARKS.-The pustules of P. subfatiscens (Fig- salazinic acid). ure 5c) often erupt, leaving a central area of exposed REMARKS.-sinCe describing P. swinscowii, 1 have medulla that blackens. Other pustules become identified it among collections made by Rolf coarsely sorediate with the soredia sometimes as- Santesson in southernmost Chile. The Chilean and suming the form of tiny isidia-like masses. The African populations are identical in morphology specimens from Africa usually produce short cilia and chemistry. In Kenya P. swinscowii grows in the on these isidioid masses and on the surface of the alpine zone at over 3500 m elevation, and in Chile pustules. The tropical American population very it occupies similar barren habitats on rock and soil rarely has any cilia. The chemistry of the neotrop- near sea level. ical and African populations is, as far as I can As mentioned in the original description, P. determine, within the limits of the solvent systems swinscowii does not fit well into any of the parmeli- available, identical and equivalent to that in P. oid groups. It superficially resembles an “evernii- horrescens. Dey (1975:433) suggests that the neo- form” Parmelia, such as sorediate, salazinic acid- tropical specimens, along with some from the containing Everniastwm sorocheilum (Vainio) Hale, southern Appalachian Mountains in the United but the rhizines are relatively short and the lobes States, may represent a new species, based in part not at all canaliculate. It intergrades to a certain on the absence of cilia on the pustules. It is also extent with abnormally small forms of Parmotrema true that the ecological requirements seem to be cetratum (Acharius) Hale or P. reticulatum (Taylor) different. The neotropical specimens were all col- Choisy, except that the upper surface is not reticu- lected in upland virgin rain forest at or above 500 late and the lobes are sublinear. The species is m elevation, whereas the African localities are in probably best left in Parmelina where it is much drier forests. I prefer to keep a broader spe- anomolous largely because of the presence of lobaric cies concept until more complete morphological and acid. ecological data can establish the case one way or the SPECIVEM EXAhZISED.-chile: Magallanes, Santesson 2159, other. 6381a, 6381b, 6478 (S, US); Tierra del Fuego, Santesson 1244 (S, US). Specimens from Kenya are listed in Hale, 1973b:4. SPECIMFUSEXA\IINID.-Panama. Code, Hale 43564. See Hale (1971a:24) for additional records from Dominica. 43.

42. Parmelina swinscowii FIGURE20b FIGURE20a Parmelina tiliacea (Hoffmann) Hale, 1974:483. Lichen tiliaceus Hoffmann, 1784:96, pl. 16: fig. 2 [type collec- Parmelina swinscowii (Hale) Hale, 1974:483. tion: Europe (not seen but illustrated in Hoffmann)]. Parmelia swinscowii Hale, 1973b:1 [type collection: Mt. Kenja, Lichen scorteus Acharius, 1798:119 [type collection: Sweden Central Province, Kenya, T. D. V. Swinscow K 31/33 (H-Ach, lectotype)]. (BM, holotype; isotype in US)]. Parmelin tiliacea (Hoffmann) Acharius, 1803:215. DEscRIPTIoN.-Thallus growing on soil or over Zmbricaria tiliacea (Hoffmann) Koerber, 1855:70. Parmelia quercifolia var. scortea f. microphylla Massalongo, mosses on soil, loosely attached, whitish mineral 1856:175 [type collection: Campofontana, Verona, Italy, gray or darkening, fragile, 3-5 cm broad; lobes Massalongo in Lichenes Ital. 329 (UPS, lectotype)]. sublinear-elongate, imbricate, 1.5-2.0 mm wide, the Parmelia scortea var. papillosa Gylenik, 1931:284 [type col- margins sparsely ciliate, the cilia black, simple, lection: Ndgrad, Hungary, Foriss 599 (BP, holotype)]. about 0.5 mm long; upper surface plane to convex, Pnrmelia scortea var. ramificn Gyelnik, 1931:285 [type collec- tion: S6sko Mountains, Hungary, Timkd 4521 (BP, shiny, becoming rugose and cracked toward the holotype)]. lobe tips, the tips remaining entire or becoming Parmelia scortea f. visegradensis Gyelnik, 1932a:444 [type minutely dissected, turning coarsely pustulate- collection: Visegrld, Pest, Hungary, Timkd (BP, holotype)]. NUMBER 33 49 50 SMITHSONIAN CONTRIBUTIONS TO BOTANY

DEscRIPTION.-Tha1lus adnate on bark or rock, SxczcIhfEvs EXAMINED(selected) .-Europe: England, Crom- light mineral gray to mineral gray, turning olive- bie (BM), Davies (BM), Larbalestzer 292 (BM); Norway, Hoeg (US), Magnusson 9193 (US); Sweden, Almborn (LD), buff to deep olive-buff in the herbarium, 5-15 cm Asplund (US), Blomberg in Lichenes Sueciae Exsiccati 64 or more in diameter: lobes irregularly branched, (LD), Santesson 12711 (US), Vrnng in Kqptogamae Exsiccatae sublinear-elongate, often imbricate, rounded at the Vindobonensi 3718 (LD, US); Finland, Kari in Lichenes apices, 2-6 mm wide, the margins more or less Fenniae Exsiccati 194, 1103 (H), Linkoln in Lichenes Fenniae crenate and undulate, sometimes narrowly black Ewsiccati 392 (H, US); Poland, Glanc in Lichenotheca Polonica rimmed, ciliate, the cilia black, coarse, shiny, simple, 171 (UPS); Russia, in Lichenotheca Rossica 17 (WIS), Oxner (US): Bulgaria, Szatala (US): France, Crozals (US), Rondon 0.2-0.7 mm long; upper surface more or less shiny, in Lichenes Selecti Exsiccati 440 (US); Germany, Erichsen white maculate, usually partly pruinose, irregularly (US), Hiltmann in Kryptogamae Exsiccatae Vindobonensi cracked on older lobes, densely isidiate, the isidia 3062 (US). Paul and Schroppel in Lichenes Alpium 9 (US); cylindrical, short, rarely branched, usually black- Czechosloi akia, Pisut in Lichenes Slovakiae Exsiccati 73 (US): Spain, Culberson 11954 (DUKE, US); Portugal, Sam- ening at the tips; medulla white; lower surface paio in Lichenes de Portugal 250 (US); Italy, Steiner in black, moderately to densely rhizinate, rhizines Ki yptogamae Exsiccatae Vindobonensi 4327 (US). Israel: black, simple, 1-2 mm long. Apothecia rare, adnate, Pharaii 11 (US). Tunisia: Runemark (LD). Morocco: Ernst sometimes retrosely rhizinate, to 4 mm in diameter; 2206 (US), hrewboula 198 (BM). Bahrain: Ahrnad L162 (L, spores 8, 5-6 X 8-11 pm. US). Pakistan: Iqbal 180 (L), 511,540 (US). India: Kashmir, Kapoor 991 (Awasthi herbarium), Kaitl 4006 (Awasthi her- CHEMISTRY.-cOrteX K + yellow (atranorin); me- barium), Watt 117, 455 (BM). dulla K-, C+, KC+ red, P- (lecanoric acid). DIsTRIBuTIoN.-Europe to western India. REMARKS.-ThiS classical European species was 44. Parmelina usambarensis the first Parmelina to be described. Except for the nomenclatural confusion with Acharius’ Parmelia FIGURE2Oc scortea, it has been correctly identified by all Parmelina usambarensis (Steiner and Zahlbruckner) Hale, lichenologists, excluding, of course, those in Amer- 1964:483. ica, who used the name for Parmelina galbina. Parmelia usambarensis Steiner and Zahlbruckner in Zahl- Hypotrachyna liuida (Taylor) Hale, and other bruckner, 1926:524 [type collection: Lutindi, Tanzania, narrow-lobed lichens. Brunnthaler (W, lectotype)]. Parmelia lneuigatoides des Abbayes, 1951:970 [type collec- Parmelina tiliacea is the isidiate morph of P. tion: Fouta-Djalon, Dalaba, Guinea, des Abbayes (RES, q uercina, which, surprisingly, has a much broader lectotype)]. geographic range, contrary to the situation for most species pairs. A close relative in Britain and Europe DEscRIPTIox.-Thallus loosely attached on rock, is P. pastillifera, which has peltate isidia and differ- whitish mineral gray, to 10 cm broad; lobes broadly ent ecological requirements (Dobson and Hawks- sublinear, divaricate to contiguous, 2-5 mm wide, worth, 1976). Taken together, the three species the marginal cilia irregularly dispersed but mostly form a distinct group in Parmelina characterized in the lobe axils, up to 1 mm long; upper surface shiny, plane, continuous or cracking in older parts, by the white maculae and presence of lecanoric sparsely to densely isidiate, the isidia cylindrical, acid. Only one other species in the genus, P. simple to branched, to 0.4 mm high; medulla pruinata, contains this acid. white; lower surface black and shiny except for a As one of the commonest parmelioid lichens in narrow dark brown zone at the tips, moderately to Europe, P. tiliacea has been mentioned and stud- sparsely rhizinate, the rhizines black, simple, 1 mm ied in detail by many workers. Sernander-Du Rietz or more long. Apothecia rare, substipitate, the (1926), for example, found that it tends to occur amphithecium isidiate, 2-3 mm in diameter; spores in ornithocoprous or dust-laden habitats in Scan- 8,6 X 12pm. dinavia. She later conducted a careful study of REMARKS.-PaYmelina usambarensis is a rather apothecial formation, showing that growth of new large, loosely attached, saxicolous lichen. The isidia apothecia could be correlated with heavy precipita- are often quite sparse. It does not conform perfectly tion in warm periods of the summer (Sernander-Du to the concept of Paymelina as I define the genus, Rietz, 1957). but the simple rhizines and coarse, long marginal NUMBER 33 51 cilia at least remove it from Hypotrachyna (Vainio) in the New World with salazinic acid, excepting Hale. On the other hand, it cannot be accommo- rare P. min.rcowii. It is also unusual in having very dated in Parmotrema LIassalongo because of the large spores with a thick episporium as well as a sublinear subdivaricate lobes. There are no close lower surface that is dark brown to black rather relatives, except, very superficially, the African en- than entirely black. The closest relative appears to demic Parmelina enormis. be the Asian P. expallida, an isidiate species with smaller spores (Kurokawa, 19G8a: 192). Although P. SPECIMFSSEx \vIsm.-Guinea: Baldruin 9849a (BM) . Ivory Coast: Man, tles .4bbnyes (LD, REN), Santesson 10632 (S, veisiformis occurs in a botanically well-known re- US). Uganda: Su’inscow 2U 15/6 (BM, US), Thornas 1678, gion, it has not been collected since 1901. 3029 (BM). Thailand: Kuroknwa 1804, 1813, 1939, 1956 (TXS), 1958 (TSS, US), Kurokawa in 1-icheria Rariores et Criiici Si~ec~si~,ssEx.iwAFD.-Brazil: hhas Gerais, r’C’flTr7lifig 283, Exsiccati 41 (US). 289, 286 (M);Rio de Janeiro, B~lrclzell 2409 (BM), Glaziou 1822, 1823 (M).

45. Parmelina versiformis 46. Parmelina wallichiana FIGURE2Od FIGURES20e, 21 Parmeliiza versiformis (Krempelhnber) Hale, 1964:483. Pnvr!ielin uersiformis Krempelhul)er, 1878:464 [type collec- Parmelinn wallichiniia (Taylor) Hale, 1974:483. tion: Argentina, Lorentz crnd Hieronymuy (M, lectotype: Parmelia wallichiana Taylor, 1847:l 76 [type collection: Sepal, isolectotypes in G, PC)]. Wnlliciz (FH-TayI, lectotype)]. Parmeiia inuta/a Vainio, 1890.3‘) [t)pe collection: Sitio, Parinelia tiliacea rar. fximfa Steiner, 1888:138 [type collection: Minas Gerais, Brazil, Vainio in Lichenes Brasilienses Usambara, Tanzania, Meyer (G, lectotype)]. Exsiccati 539 (TUR, 1ectot)pe: isolectotypes in Bhl, FH, Parmelia juflodii Steiner, 19073640 [type collection: Sana- UPS)]. torium, Cape Proiince, Union of South Africa, Junod 978 Parmelia catliari?ie,!sis Miiller Argoviensis, 1891:239 [type (G, lectot!pe)]. colleclion: Near Santa Catarina, Brazil, Uie 1891 (G, lecto- Parmelia nimandaira,la Zahlbrnckner, 1934a:55 [type collec- type; isolectotjpe in \V)]. tion: Mt. rlrisan, Taiivan, Asahimi 63 (W,lectotype; iso- Parmelia weltsteinii Zahlbrnckner, 1909:173 [type collection: lectotype in TSS)]. Near Taipas, Sao l’aulo, Brazil, Schiffner and Wettsteiii Parrnelia nimandairana f. sirblaeui.~Asahina, 19523131) [type (W,lectotype, isolectotype in G)]. collection: hlt. Buko, Prov. hfusashi, Japan, Asahiiza 35 (TNS, lectotype)]. DE~CRIPTION.-T~~~~USadnate to loosely adnate on bark, turning deep olive-buff in the herbarium, DEscRIPTioN.-Thallus adnate to loosely adnate about 8 cm in diameter; lobes irregularly branched, on bark or rock, yellowish glaucous to pale sublinear-elongate, more or less imbricate, 2-8 mm glaucous-green, 5-20 cm in diameter; lobes irregu- wide, the margins crenate, ciliate mostly in lobe larly branched, apically rotund, 3-10 mm wide, the axils, the cilia often sparse, to 0.5 mm long; upper margins more or less crenate, lobulate with age, surface more or less shiny, not maculate, wrinkled short ciliate, especially in the axils, the cilia about and irregularly cracked on older lobes, soredia and 0.5 mm long; upper surface shiny, not maculate, isidia lacking; medulla white; lower surface dark irregularly cracked on older lobes, sparsely isidiate, brown or blackening, more or less wrinkled, sparsely isidia cylindrical, mostly Simply, less than 1 mm rhizinate, the rhizines pale brown, simple, 0.1-0.3 high; medulla white; lower surface black, mod- mm long. Xpothecia common, substipitate, 2-10 erately to sparsely short rhizinate, dark brown and mm in diameter, the rim crenate and undulate, the naked or papillate in a rather wide Lone near the amphithecium rugose, the disc blackish brown; tips, the rhizines black, simple. Apothecia rare, spores 8, 10-14 X 18-28 pm, the episporium about adnate, 2-7 mm in diameter; spores 8, 8-10 x 3 pm thick. 14-18 Fm. CHEmsmY.-Cortex K + yellow (atranorin); me- CHEnmTRu.-Cortex K + yellow (atranorin); me- dulla K+ yellow turning red, C-, KC-, P+ dulla K+ yellow turning red, C-, KC-, P+ orange (salazinic acid). orange (salazinic acid). DISTRIBUTION.-~OUth America. DISTRIBuTIoN.-Africa and Asia (Figure 2 1). REMARKS.--This is the only species of Parmelina REMARKS.-This is the most widespread and com- 52 SMITHSONIAN CONTRIBUTIONS TO BOTANY

monly collected Paimelina in the Old World. It is tributed as Parmelia isidiza). Nepal: Awasthi 2156, 2157, easily recognized by the adnate, moderately isidiate 2198, 2203 (Arvasthi herbarium), i\Torkett 5533 (BM), Togashi thallus that grows in a variety of habitats in the (TNS, US). Sikkih: Bose 136 (Awasthi herbarium). India: highlands of Africa, India, and eastern Asia, rang- Uttar Pradesh, Awasthi 3175, 3509, 3814, 3980, 3986 (Awasthi herbarium), Hara (TNS, US); Tamil Nadu, Foreau 43 ing from temperate Japan to both higher elevations (Awasthi herbarium), Hale 40220, 40274, 43620, 43740, 43870. in the tropics and to rather low elevations near Thailand: Kerr L4 (BM). Malaya: Pahang: Hale 30094, the equator in Guinea and the Ivory Coast. The 30217, 30218, 30219. Taiwan: Kurokawa 29, 530, 626, 2926 parent morph is no longer extant nor are the (TNS). China: Kwantung Province, Petig 12578 (US); Man- parallel sorediate or pustulate morphs. churia, Asahina 25 (TNS). Hong Kong: Thrower 1586 (US). Japan: Prov. Aki, Hale 29508; Prov. Chikuzen, Kurokawa SPECIMENSEXAMINED.-GUinea: N’Zerekore, Santesson 10568b 62545 (TNS); Prov. Higo Kiirokawn 63076 (TNS); Prov. Hizen, (UPS, US). Ivory Coast: Man, Santesson 10650 (UPS). Uganda: Kurokawa 63157 (TXS); I’rov. Hyuga, Hale 29645; Prov. Iyo, Dummer 5064 (US), Swinscow 2U 24/20 (BM, US). Kenya: Kurokau~a60098 (TNS); Prov. Iru, Asahina 94 (Th’S); Prov. Nyanza Province, Maas Geesteratius 10935 (L, LD, US), 4950, Kii, Asahina (TNS), Kzirokawa 60237 (TNS, US), Kurokawa 11164 (L); Rift \’alley Province, hfaas Geesteranus 4832 (L). Angola: Cuanza Sul, Degelius (Degelius herbarium, US). in Lichenes Rariores et Critici Selecti 42 (US); Prov. Musashi, Rhodesia: Hoeg (TRH). Tanzania: Arusha Province, San- Asahina 11 (TNS), Ktirokawa 59155 (TNS, US); Prov. Ohmi, tesson 21339 (UPS, US); Kiliuiarijaro Province, Santesson Hale 29460; Prov. Satsuma, Kurokawa 63043 (TNS); Tosa, 20961, 20989, 20990 (UPS, US); Moshi District, Burnet T122 Fujikawa (TNS). Yoshimirra 1157 (US). Philippines: Benguet, (BM, US). Swaziland: Almborn 7901 (LD), Kofler (LD). Union Degelius As-787 (Degelius herbarium), Hale 26760, Hale of South Africa: Natal, Almborn 8494 (LD), 8683, 9732, 9871 in Lichenes Selecti Exsiccati 672 (US); Mountain Province, (LD, US), Hoeg (TRH), Kofler (LD); Transvaal, Almborn Haze 26413. Sabah: Hale 28707, 28807, 29063, 29071. Indonesia: 6430 (LD). Madagascar: des Abbayes in Licheiies Madagas- Java, Groenhart 3238 (L, US), 4358, 5881, 5913, 5917 (L), 5948 carienses et Borbonici Selecti Exsiccati 10 (UPS, US, dis- (L, US), Schiflner 3294 (US), Voogd 2079 (L). NUMBER 33 53

47. Parmelina xantholepis Feinando Po, West Africa, G. Thornson (Bhf, lectotype; CLAM, isolectotype)]. FIGURE20f The type material is too fragmentary for ade- Parmeliria sun tholepis (Montagne and van den Bosch) Hale, quate study. It is a nonsorediate, nonisidiate, sub- 1974:438. Parrne/ia xantholepis hIontagne and van den Bosch, irregularly lobed species with a black lower surface, 1855:428 [type collection: Gode, Java, Teydom (L, simple rhizines, and numerous marginal cilia. The lectotype)]. spores are large, 15 X 28 pm. Atranorin and sala- Parrnelia hiformis T’ainio f. dataensis Vainio [= f. biformis], zinic acid are present. It is almost certainly a 1909:660 [type collection: Mt. Data, Luzon, Philippines, Pnrmelina, and its status will only be clarified as Alerrill 4987 (TUR, T’ainio herbarium number 2607, lectotype)]. more collections are made in TYest Africa. Parmelia biformis f. pauniensis T’ainio, 1909:660 [type collec- tion: Pauai, Benguet, Luzon, Philippines, Mearns 4434 (TUR, T’ainio herbarium number 2605, lectotype)]. ParmeIin orchidophlia DEscRIP’rIos.-Thallus adnate on bark or rock, Pnrnielin orcliidoljhila Dodge, 1953:374 [type collection: yellowish mineral gray, very fragile, 4-15 cm broad; Nyinabitsa, TVestern Pro\-ince, Uganda, Omastin 118.1 lobes sublinear to subirregular, 1-3 mm wide, (Bhf, lectot! pe)] crowded, dissected, the marginal cilia coarse, 0.2-0.8 The type collection is too Eragmentary for study. mm long; upper surface plane, maculate, becoming Salazinic acid was proved niicrochemically. It may densely lobulate toward the lobe margins, isidia be Parmelina zunllichiann (Taylor) Hale. and soredia lacking; medulla barium yellow; lower surface black, densely rhizinate, the rhizines black, long, simple or squarrose. Apothecia adnate, 1-3 Parmelia tiliacea var. lezicina mm in diameter; spores 6-8 X 9-13 pm. Parmelin tilicea Tar. leucintr Miillrr Argoviensis, 1880:267 CHEXIISTRY.-cOl.teX K + yellow (atranorin); me- [type collection: Sear Petropolis, Brazil, Deventer (G, dulla more intensively yellow with K and C, P- lec totype)]. (zeorin, leucotylin and related terpenes, and seca- lonic acid A.) This variety is equal to Hypotrachyna dactylifera DISTRImTIoN.-hdia and Nepal to the Philip- (Vainio) Hale (Hale, 1975a:30). pines. REM.mKs.-This species is easily identified by the Pal-melia tilacea var. rimzilosa fragile, lobulate thallus with a pale yellow-orange medulla. It is closely related to the P. amagiensis-P. Pnrrrielin tilicea tar, rriricilosa Muller Aigoiiensis, 1882:458 clenegans series, definitely in chemistry but less so in [tJpe collection: Socotra, h’nlfoirr (C, lectotipe)]. thallus texture. It is usually collected at the base of trees or on rocks, often growing among mosses, at No type specimen could be located at Geneva. One of two specimens so labeled by hluller is Par- higher elevations (1000-2300 m) in evergreen hard- motrema reticiilatum (Taylor) Choisy, and the wood cloud forests. other is Pseudoparmelia carneopruinata (Zahl- SPECIMESSEx \MIsIm-h’epaI: Poelt 101, 111 (hI) , Norkett bruckner) Hale. 9227 (BM). India: Tamil Nadu, Foreau 4118 (ALvasthi her- barium, US), Hale 43862; Uttar Pradesh, Awasthi 3839 (Awasthi herbarium). Thailand: Kziroknwa 1669 (TNS, US). Pal-melina bagziioensis Philippines: Mountain Province, Hale 25993, 26096, 26187, 26344, 26554. Indonesia: Java, Groenhart 2841, 5939 (L), Panneliizu bnguioensis (Hale) Hale, 1974:482 [type collection: 5941, 6004 (L, US). Baguio, Mountain Province, Philippines, Ha/e 26768 (US, holotype)].

Doubtful and Rejected Names Closer examination of this sorediate species con- vinced me that it is a member of the genus Hypo- Parmelia coilocarpa tl-achyna (H. bagtrioensis (Hale) Hale, new combi- Parriwlia coilocnrpn Stirton, 1877-78:202 [t)pe collection: nation). The rhizines are rather sparse but clearly 54 SMITHSONIAN CONTRIBUTIONS TO BOTANY

dichotomously branched at maturity. In any event, I have placed this species in synonymy under the chemistry-fumarprotocetraric acid-was ano- Pa rme 1 ina szi bau ru 1en t a. malous for Parmelina, and this was the first clue to its correct generic status. Parmelina nylanderi

Parmelina carporrhizans Parmelina nylanderi (Lynge) Hale, 1974:483. Parmelia nylanderi Lynge, 1914:82 [type collection: Sear $450 Parmelina carporrhizans (Taylor) Hale, 1974:482. Jeronyno, Serra da Chapada, Mato Grosso, Brazil, Malme 2747 (S, lectotype)]. I now consider this to be a synonym of Parme- lina quercina. I originally considered this loosely attached saxi- colous lichen to be a member of Parmelina, but after examining a second collection (Eiten 3429 Parmelina crystallorum (US) from S5o Paulo, Brazil), I now feel that the chemistry (usnic acid in the cortex and salazinic Parmelina crystallorum (Lynge) Hale, 1974:482 and gyroplioric acid in the medulla) and general This species is synonymized under Parmelina habit, in spite of the narrow ciliate lobes, place it damaziana. more appropriately in Parmotrema (P. nylanderi (Lynge) Hale, new combination). It is related to the much broader lobed P. delicatulzim (Vainio) Parme 1ina h om ogen es Hale group so common on sandstone in South Parmelina homogenes (Nylander) Hale, 1974:482. America. Literature Cited

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55 56 SMITHSONIAN CONTRIBUTIONS TO BOTANY

1958. Studies on the Chemistry and Distribution of North Hedrick, J. American Lichens (10-13). Bqologist, 61:81-85. 1934. New Genera and Species of Lichens from the Her- 1959. New or Interesting Parmelias from North and Trop- barium of Bruce Fink, 11. Mycologia, 26:153-166. ical America. Bryologist, 62:123-132. Hoffmann, G. F. 1960. A Revision of the South American Species of Par- 1784. Enumeratio Lichenum. 102 pages. Erlangae. melia Determined by Lynge. Contributions from the Hue, A. M. United States National Herbarium, 36: 141. 1899. Lichenes extra-europaei. Nouvelles Archives du Mu- 1965. A Monograph of Parmelia Subgenus Amphigymnia. seum Paris, series 3, 1: 1-250. Contributions from the United States National Her- James, P. W. barium, 36: 193-358. 1965. A New Check-List of British Lichens. Lichenologist, 1967. Chemistry and Evolution in Lichens. Israel Journal 3:95-153. of Botany, 15:150-157. Koerber, G. W. 1971a. Morden-Smithsonian Expedition to Dominica: The 1855. Systenza Lichenum Gernianiae. 458 pages. Breslau. Lichens (Parmeliaceae). Smithsonian Contributions Krempelhuber, A. to Botany, 4: 1-25. 1878. Lichenes collecti in republica Argentina a Doctori- 1971b. Two Species of Parmelia New to North America. bus Lorentz et Hieronymus. Flora, 61:461-464. Bryologist, 74:44-47. Kurokawa, S. 1972a. Four New Species of Parmelia (Lichenes) from India 1968a. Parmelia expallida, a Xew Lichen Species from and the Philippines. Bryologist, 75:97-101. Eastern Asia. Bulletin of the National Science Mu- 1972b. Pamielia jamesii, an Unusual Species in Section seum, 11:191-194. Imbricaria (Lichenes) from Australia and New Zea- 1968b. New or Noteworthy Species of Parmelia of Japan. land. Phytologia, 23: 179. Journal of Japanese Botany, 43:349-353. 1972~.Six New Species of Parmelia (Lichenes) from Atrica. 1972. Probable Mode of Differentiation of Lichens in Phytologia, 23:343-349. Japan and Eastern North America. Pages 139-146 1973a. Fine Structure of the Cortex in the Lichen Family in A. Graham, Floristics and Paleofloristics of Asia Parmeliaceae Viewed with the Scanning-Electron and North America. Amsterdam. Microscope. Smithsonian Contributions to Botany, Kurokawa, S., and M. Mineta 10: 1-92. 1973. Enumeration of Parmeliae of Ceylon. Annual Re- 1973b. New Parmeliae (Lichenes) from Africa. Phytologia, port of the Noto Marine Lnboratory, 13:71-76. 27~1-6. de Lamarck, J., and A. P. de Candolle 1974. Bulbothrix, Parmelina, Relicina, and Xanthopar- 1805. Flore Francaise. Volume 2, 600 pages. Paris. melia, Four New Genera in the Parmeliaceae (Lich- Lynge, B. enes). Phytologia, 28:47%490. 1914. Die Flechten der ersten Regnellschen Expedition. 1975a. A Revision of the Lichen Genus Hypotrachyna Die Gattungen Pseudoparmelia gen. nov. und Par- (Parmeliaceae) in Tropical America. Smithsonian melia Ach. Arkiv for Botanik, 13 (13):l-172. Contributions to Botany, 25:l-73. Massalongo, A. B. 1975b. A Monograph of the Lichen Genus Relicina 1856. Schedae Criticae. Part 10, number 328. (Parmeliaceae). Smithsonian Contributions to Bot- hfichaux, A. any, 26: 1-32. 1803. Flora Boreali-Americana. Volume 2, 340 pages. Paris. 1976a. A Monograph of the Lichen Genus Pseudoparmelia Montagne, J. K. C., and R. B. van den Bosch Lynge (Parmeliaceae). Smithsonian Contributions to 1855. Lichenes, Collemaceae. Pages 427-494 in F. Jung- Botany, 31:l-62. huhn, PIantae Junghuhnianae [sive Enumeratio 1976b. A Monograph of the Lichen Genus Bulbothrix Hale Plantarum quas in insulis Java et Sumatra]. (Parmeliaceae). Smithsonian Contributions to Bot- Moore, B. J. 1968. The Macrolichen Flora of Florida. Bryologist, 71: any, 32: 1-29. 161-266. H ale, M E., Jr., and S. Kurokawa Muller Argoviensis, J. 1962. Parmelia Species First Described from the British 1877. Lichenologische Beitrage, V: Lichenen aus Texas. Isles. Lichendogist, 2: 1-5. Flora, 60:77-80. 1964. Studies on Parmelia Subgenus Parmelia. Contribu- 1880. Licheologische Beitrage, XI. Flora, 63:260-268. tions from the United States h’ational Herbarium, 1882. Diagnoses Lichenum Socotrensium novorum. Pro- 36~121-191. ceedings of the Royal Society of Edinburgh, 11:457- H armand, J. 472. 1897. Catalogue descriptif des Lichens observes dans la 1883. Lichenologische Beitrage, XVII. Flora, 66:4548. Lorraine. Bulletin de la Socidtd des Sciences de 1887. Lichenologische Beitrage, XXVI. Flora, 70:316-320. Nancy, series 2, 14:l-510. 1888a. Lichenes Montevidenses quos legit et communicavit 1910. Lichens de France. Volume 4, pages 483-732. Paris. Prof. Arechavaleta. Revue Mycologique, 10: 1-5. 1928. Lichens d’Indo-Chine recueillis par M. V. Demange. 1888b. Lichenologische Beitrage, XXIX. Flora, 71 : 195-208. Annales Cryptogarnie Exotique, 1:319-337. 1888~.Lichenologische Beitrage, XXX. Flora, 71 :528-552. NUMBER 33 57

1891. Lichenologische Beitrage, XXXV. Flora, 74:371-382. 1847. New Lichens, Principally from the Herbarium of Nylander, W. W. J. Hooker. Hooker Journal oj Botany, 6:148- 1856. Prodromus Lichenographiae Galliae et Algeriae. 197. Actes de la Societe Lznndenne de Bordeaux, 21:249- Tuckerman, E. 467. 1858. Supplement to an Enumeration of North American 1868. Enumeration des lichen, recoltes par M. T. Husnot Lichens: Part First, Containing Brief Diagnoses of aux Antilles francaises. Bulletin de la Societe Lin- New Species. American Journal of Science and Arts, ndenne de Normandie, (2)3:259-280. series 2, 25:422-430. 1873. Observata Lichenologica in Pyrenaeis orientalibus. 1882. A Synopsis of the North American Lichens. 261 Bulletin de la Societe Linntenne de Normandie, pages. Boston. (2)6:256-328. Vainio, E. A. 1882. Addenda nova ad Lichenographiam europaeam. 1890. Etude sur la classification naturelle et la morpho- Flora, 653491458. logie des Lichens du Bresil. Acta Societatis pro 1885. Parmeliae exoticae novae. Flora, 68:605-615. Fauna et Flora Fennica, 7(7):1-247. 1890. Lichenes Japoniae. 122 pages. Paris. 1896. Lichenes Antillarum a W. R. Elliott collecti. Jour- 1900. Lichenes Ceylonenses. Acta Societatis Scientiarum nal of Botany British and Foteign, 34:31-36. Fennicae, 26(10):791-821. 1899. Lichenes in Caucaso et in Peninsula Taurica annis Poelt, J. 1884-1885 ab H. Lojka et hi. a Dechy collecti. 1961. Mitteleuropaische Flechten, VII. Mitteilungen der Terineszetrajz i Fuzetek, 22:269-343. Botanischen Staatssammlung Munchen, 4: 171-197. 1909. Lichenes Insularum Philippinarum, I. Philippine 1969. Bestimmungsschliusel europascher Flechten. 757 Journal of Science, 4:651-662. pages. Lehre. Willdenow, C. L. Schauer, ..-1. 1787. Florae Berolinensis Prodromus. 440 pages. Berlin. 1965. Oceanische Flechten im Nordalpenraum. Portu- Yosioka, I. galiae Acta Biologica, (B)8:17-229. 1966. The Structure of Leucotylic Acid, a New Triterpenic Schreber, J. C. D. Acid from a Lichen. Tetrahedron Letters, 6:607- 1791. Linne, Genera Plantarum. Eighth edition, 872 pages. 612. Frankfurt am Main. Yosioka, I., and T. Nakanishi Schubert, R., and 0. Klement 1963. Structure of Leucotylin. Chemical Pharmaceutical 1966. Beitrag zur Flechtenflora von Sord- und LMittel- Bulletin, 11: 1468-1470. indien. Nova Hedwigia, 11 : 1-73. 1966. On the Triterpenic Constituents of a Lichen, Par- Sernander-Du Rietz, G. melia entotheiochroa Hue, Zeorin, Leucotylin, 1926. Parnzelia tiliacea, en kustlav och marin inlandsrelikt Leucotylic Acid, and Five Sew Related Triterpenic i Skandinavien. Svensk Botanisk Tidskrift, 20:352- Acids. Chemical Pharmaceutical Bulletin, 14:804- 365. 807. 1957. Om yttre faktores inverkan pa apotheciebildningen Yosioka, I., T. Nakanishi, S. Izumi, and I. Kitagawa hos Parmelia tiliacea. Svensk Botanisk Tidskrift, 1968a. Structure of a Lichen Pigment Entothein and its 51:454-488. Identity with Secalonic Acid A, a Major Ergot Pig- Shibata, S., S. Natori, and S. Udaga1r.a ment. Chemical Pharmaceutical Bulletin, 16:2090, 1964. List of Fungal Products. 170 pages. Tokjo. 2091. Steiner, J. Yosioka, I., T. Nakanishi, and Kitagawa 1888. Ueber afrikanische Flechten. Jahres-Bericht der I. 3968b. On the Stereostructures of Zeorin and Leucotylin. Schlesischen Gesellschaft jur vaterlundische Cultur, 66: 133-150. Tetrahedron Letters, 12:1485-1490. 1907. Lichenes austro-africani. Bulletin de l’Herbier Bois- Zahlbruckner, A. sier, 2(7) : 637-646. 1905. Lichenes a cl. Damazio in Brasilia lecti. Bulletin de Stirton, J. I’Herbim Boissier, series 2, 5:539-543. 1877-78. On Certain Lichens Belonging to the Genus 1908. Beitriige zur Flechtenflora Brasiliens. Bulletin de Parmelia. Scottish Naturalist, 4:200-203. 1’Herbier Boissier, series 2, 8:459468. 1899. Lichenes. In F. M. Bailey, Contributions to the 1909. Lichenes (Flechten). In R. I\’ettstein and 1’. Schiff- Flora of Queensland. Queensland Agricultural Jour- ner, Ergebnisse der botanischen Expedition der nal, 5:483-488. kaiserlichen Akademie der Wissenschaften nach Stizenberger, E. Sudhrasilien 1901. Denkschrift Akademie der Wis- 1890. Lichenaea africana. Bericht uber die Taigkeit der senschaften in Wien. Mathematisch-A’aturwissen- St. Gallischen Naturruissenschaftlic~ieii Gesellschaft, schajtliche Klasse, 83:87-211. 1888-89: 105-209. 1926. Afrikanische Flechten (Lichenes). Botanische Jahr- Ta)lor, T. bucher jur Systetnatik, 60:468-552. 1836. Lichens. In J. T. hlackay, Flora Hibernica. 279 1927. Additamenta ad Lichenographiatn Japoniae. Bo- pages. Dublin. tanical Magazine (Tokyo), 41:313-364. 58 S\iITHSOXI 4S CONTRIBUTIOSS TO BOTASY

1929. Catalogus iichenum tcniiwsrriic. T.olume 6, 323 1934a. Flechten der Inrel Forniosa. FeddP Reperiorium pages. Leipiig. 33:22-C,F. 1930. Lichenes. 171 H. Handel-hlazretti, Sjmboiae Sinica, 1934. Nachtrage zur Flechtenflora Chinas. Hedwigia, 3:1-234. 74: 193-21 3. Index

(Synonyms in italics) lmbricaria quercina, 42 hubrichtii, 26 subaurulenta, 47 tiliacea, 48 hrinanensis, 19 siibaurulenta var. inyriocarpa, 47 Lichen quercinus, 42 immiscens, 32 subbalansae, 40 tiliaceus, 48 insinuata, 34 subcremea, 47 Par in elia a 1b ido -s tra minea, 19 insin itat ula, 34 sribfatisceiis, 47 amagiensis, 18 irrugans, 34 subquercifolia, 30 ainazonica var. hirsnotii, 26 jarnesii, 35 subqiiercifolia f. Aiibradiata, 30 antillensis, 18 junodii, 51 subreuolitta, 19 atricha, 42 laruignta ssp. estremi-orienialis, 30 subsz~lphitrata,39 aurulenta, 19 iaeuigata var. gracilis f. furfuracea, 25 tilracea. 48 halansae, 22 laevigatoides, 50 tiliacea \ ar. afjixa, 42 balansae \ar. sorediata, 40 lericotylita, 35 tilicicea Tar. efflorescens, 19 biformis f. dataensis, 53 leucotqlix f. rugulosa, 35 tiiiacva var. exirnin, ,51 biforinis f. pnuaiensis, 53 leiico/ylizn f. siiblaevis, 35 tiliacen ~ar.hsmthri\, 42 blastica, 18 lin d ilia nii, 36 tiiiacra \ar. leuciiia, 53 brachpconidia, 23 mrlaiiochneta, 36 tiliacea var. minor, 30 brachsconidia \ ar. chlorocarpa, 23 nietnrevoiiita, 36 ti/iacen \ar. rimiilosn, 53 budapestiiiensis, 42 inichoncnnensis, 32 t iliacea su bq ii r rci folio, 30 ca m t rchadalis i'ar. epiphy lla , 26 ni innJIL m, 26 tilinrea var. siiipliurouz,, 30 carporrhizans, 42 riii/el/eri, 38 tiliaren Iar. siilphurosa f. asperata, 36 carporrhizans f. inalicola, 42 uiiitata, 51 usariibarensis, 50 catharinensis, 51 myriocnrpa, 47 versiforniis, 5 1 coilocarpa, 53 11 ima nda i ra n a, 5 1 wailichiaiia, 51 consors, 22 iiimaiidairnna f. sublaevis, 51 wettsteinii, 51 conspiciia, 47 nylanderi, 54 yalungana, 42 con t in entalis, 22 obsessa, 36 vnnthorarpa, 34 cryptochlora, 23 orchidophila, 53 santholefiis, 53 crystalloruin, 23 pastillifera, 39 Parmelina amagiensis, 18 damaziana, 23 perisidians, 39 antillensis, 18 deminuta, 30 phlyctina, 40 aurulenta, 19 denegans, 25 pilosa, 40 baguioensis, 53 dissecta, 25 pruinata, 42 carporrhizans, 54 endoleuca, 27 piiiggari, 26 consors, 22 enormis, 27 quercifolia var. scrotea f. inicrophylla, crassata. 22 entotheiochroa, 28 48 cr) ptochlora, 23 expallida, 28 quercina, 42 crystallorum, 54 fecunda, 47 querrina var. stclphurosa, 30 tlamaziana, 23 finkii, 38 revolnta i.ar. granulata, 42 degelii, 25 fraudans ssp. subfraudans, 35 sampniana, 22 tlenegans, 25 galbina, 30 .scortea var. papillosa, 48 dissecta, 25 galbina var. rugosa, 30 scortra var. pastillifera, 39 endoleuca, 27 galbina 1-ar. sitbradiata, 30 scortea iar. ramifica, 48 enormis, 27 hayachinensis, 31 scortea var. visegradensis, 48 entotheiochroa, 28 heteroloba, 31 silvestris, 19 expallida, 28 homalotera, 47 simplicior, 44 galbina, 30 homogenes, 47 sinuosa var. hypothrix, 42 hayachinensis, 31 homogenes f. minor, 47 spathulata, 46 heteroloba, 31 horrescens, 31 sp umosa, 46 homogenes, 47, 54

59 SMITHSONIAN CONTRIBUTIONS TO BOTANY horrescens, 31 obsessa, 38 spathulata, 46 immiscens, 32 pastillifera, 39 spumosa, 46 indica, 34 perisidians, 39 suhaurulenta, 46 jamesii, 35 phlyctina, 40 suhfatiscens, 47 leucotyliza, 35 pilosa, 40 swinscowii, 48 lindmanii, 36 pruinata, 42 tiliacea, 48 melanochaeta, 36 quercina, 42 usamharensis, 50 metarevoluta, 36 rhytidodes, 43 versiformis, 51 muelleri, 38 schindleri, 44 wallichiana, 51 nylanderi, 54 simplicior, 44 xantholepis, 53