SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 26 

A Monograph of the Lichen Genus 

Relicina (Parmeliaceae) 

Mason E. Hale, Jr. 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1975 
ABSTRACT 

Hale, Mason E., Jr. A Monograph of the Lichen Genus Relicina (Parmeliaceae). Smithsonian Contributions to Botany, number 26, 32 pages, 16 figures, 

1975.-A revision on the world level is made for Relicina, a generic segregate of 

Parmelia characterized by having bulbate cilia on the lobe margins and by producing usnic acid in the cortex. Of the 24 species presently known, 19 occur in 

the Old World tropics, with the greatest concentration in the lowland dipterocarp 

forests, 3 are endemic to the New World tropics, and 2 occur in both the New 

and Old World outside of Africa. The most important taxonomic characters are 

type of rhizine (simple or branched), presence of coronate apothecia, isidia, and 

chemistry. The major chemical constituents are echinocarpic acid, fumarprotocetraric acid, and protocetraric acid. The genus is considered to be of fairly 

recent origin but rather conservative in terms of morphological and chemical 

evolution. Four new species, R. amphithrix, R. incongrua, R. precircumnodata, 

and R. subconnivens, are described and one new combination, R. relicinula 

(Muller Argau) Hale, is made. 

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded 

in the Institution’s annual report, Smithsonian Year. SI PRESS NUMBER 5356. SERIF^ COVER DESIGN: 

Leaf clearing from the Katsura tree Cercidiphyllum japonicum Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data 

Hale, Mason E. 

A monograph of the lichen genus Relicina (Parmeliaceae). 

(Smithsonian contributions to botany, no. 26) 

Supt. of Docs. no.: SI 1.29.26 

1. Relicina. I. Title. 11. Series: Smithsonian Institution. Smithsonian contributions to botany, 

QKl.S2747 no. 26 [QK585.P2] 581’.08s [589’.1] 75-6191 19 no. 26. 
Contents 

Page 

Introduction 1 

Morphological Characters .......................................... 2 

Ecology and Habitats .............................................. 11 

Phytogeography ....... ............................. ...... 12 

Evolution and Speciation .......................................... 13 

Position of Relicina in the Parmeliaceae ............................. 15 

Key to Species of Relicina .......................................... 

Literature Cited ................................................... 32 

...................................................... 

Chemistry ........................................................ 11 

16 

Species Treatment ................................................. 17 

iii 

A Monograph of the Lichen Genus 

Relicina (Parmeliaceae) 

Mason E. Hale, Jr. 

Introduction 

Relicina is a generic segregate of the collective 

genus Parmelia. It is characterized by marginal bulbate cilia and the constant occurrence of usnic acid, 

a yellow pigment, in the cortex. The genus includes 

24 species, almost all of them occurring in tropical 

Southeast Asia. This paper is a monographic treatment of the genus based on herbarium and field 

studies. 

The first species of Relicina was described by 

E. Fries as Parmelia relicina on the basis of a specimen from “Rawak” in 1825. Neither Linnaeus nor 

Acharius had seen any material belonging to this 

genus. Laurer described the second species, P. limbata, an Australian endemic, in 1827. This species 

is well known because it was issued in several 

widely distributed exsiccati. 

In the remainder of the 19th century only five 

more taxa were discovered: Parmelia abstrusa 

Vainio, P. circzrmnodata Nylander, P. relicinella 

Nylander, P. relicinula Muller Argau, and P. limbata f. isidiosa Muller Argau (=Relicinu sydneyensis). Between 1900 and 1964 an additional four 

taxa were described: P. abstrzisa f. laevigata Lynge 

(= Relicina subabstrusa), P. nigrociliata Hillmann, 

P. samoensis Zahlbruckner, and P. samoensis var. 

eximbvicata Gyelnik (=Relicina eximbricata). 

Hillmann (1940) recognized a natural relationship 

between six species described from the Old World 

and constructed a key to them. 

In I964 Hale and Kurokawa (1964) examined a 

Mason E. Hale, Jr., Department of Botany, h’ational Museum 

of Natural History, Smithsonian Institution, Washington, 

D.C. 20560. 

series of collections from the Bogor herbarium and 

described seven more species, mainly from Java 

and Borneo (Kalimantan and Sabah). Most of 

these, as with previous species, were known only 

from type collections. It became obvious during 

the course of their work, however, that these marginally bulbate species formed a natural group, 

quite distinct from section Xanthoparmelia where 

they had previously been classified. They recognized the 18 species then known as a series in 

subgenus Paymelia section Imbricaria subsection 

B icorn u tae. 

M7hile this publication (Hale and Kurokawa, 

1964) was actually in press, I had an opportunity 

to collect lichens in Malaysia, the Philippines, and 

Japan. What I discovered explained why this 

peculiar assemblage of species had been so poorly 

collected in the past and represented by so few 

herbarium specimens. Relicina inhabits canopy 

branches in the vast lowland dipterocarp forests 

from Burma to the Solomon Islands. I collected at 

logging heads where these giant trees, 60 m or 

more high, were being felled commercially. Most 

earlier collectors had obviously not been able to 

find (or did not try to find) felled trees. For example, many lichens were collected by botanists in 

the Philippines and studied by Vainio for his 

extensive Philippine lichen flora, yet he did not 

report a single species of Relicina, in reality the 

commonest foliose lichens in the Philippines! Another habitat where Relicinn has proved abundant 

is the oak forests found at high elevation throughout southeastern Asia. These areas have not been 

visited by lichenologists either. 

The most reassuring aspect of my field work was 

1 
2 ShiITHSONIAN CONTRIBUTIONS TO BOTANY 

that all of the species described by Hale and 

Kurokawa were rediscovered, often in great abundance and comprising the major foliose element in 

lowland rain forest. As one might expect, more 

undescribed species were found. I described two 

species, Parmelia puorescens and P. malesiana 

(Hale, 1965), and Kurokawa (1965) described other 

species from Japan (P. echinocarpa and P. subturgida). In the final revision of this monograph 

four more new species are being described to bring 

the total for the genus to 24 species. 

ACKNOWLEDGMENTS-I wish to thank the following curators of museums who kindly sent specimens 

on loan: Dr. S. Ahlner, Dr. T. Ahti, Dr. R. Alava, 

Dr. 0. Almborn, Dr. C. E. Bonner, Mme. JovetAst, Dr. R. A. Maas Geesteranus, Dr. H. Merxmuller, Dr. H. Riedl, and Dr. R. Santesson. 

Dr. S. Kurokawa assisted with species descriptions, identification, and chemical studies both in 

1960 in Washington and in 1964-1965 when I 

studied at the Tokyo National Science Museum. 

The field studies were supported by the National 

Science Foundation under the Japan-US. Cooperative Science Program and by the Smithsonian Research Foundation. In this connection I wish to 

express sincerest thanks to Dr. G. Velasquez and 

Dr. Flora Uyenco of the University of the Philippines, who arranged for trips to lumbering areas 

in the Philippines. I am also indebted to Mrs. 

Sheila Collonette for arranging a trip to Mt. Kinabalu, to Dr. W. Meijer for trips to eastern Sabah, 

and to Mr. J. Anderson for assistance in Sarawak. 

Dr. Manuel Lopez Figueiras kindly provided support and transportation in Venezuela. The project 

would never have been completed without the 

generous logistic support of lumber companies and 

forestry departments in the Philippines, Malaya, 

Sarawak, and Sabah. 

The scanning-electron microphotographs were 

taken by Mr. Walter Brown of the Smithsonian 

Scanning-electron Microscope Laboratory. Herbarium specimens were photopaplied by the Smithsonian Photo<graphic Services. 

Morphological Characters 

THAI.I.US ADNATIoN.-Most species are closely 

adnate and usually collected still attached to the 

bark. None have ascending lobes. 

LOBE CoNFrGuRATIoN.-Species of Relicina are 

extremely uniform in lobation. The lobes are 

generally narrow and sublinear, dichotomously 

branched, and contiguous. Small marginal lobules 

often develop. 

usual features of Relicina is that no soredia are 

produced by any of the species. Soredia are usually 

the most common diaspore in the Parmeliaceae. 

Isidia, however, are produced by five species, 

R. abstrusn, R. amphithrix (Figure la), R. circumnodata, R. planiuscula, and R. sydneyensis (Figure 

3i). The isidia are cylindrical and mostly unbranched but may become procumbent and lobulate in R. amphithrix and to a lesser extent in 

R. planiuscula. True lobules that do not originate 

from isidia are produced by R. luteoviridis (Figure 

14e) and R. schizospatha (Figure Ib). Appressed 

palmately branched lobules (Figure lc) are characteristic of some populations of R. amphithrix. 

EPICORTEX AND CoRTEX.-The internal structure 

of Relicina was studied by making longitudinal 

sections of the lobes and examining them under a 

scanning-electron microscope (SEM). All of the 

species are illustrated in Figures ld-3i. I had previously determined with the SEM that Relicina 

has a pored epicortex and a basic palisade plectenchymatous upper cortex (Hale, 1972). More detailed investigations of internal structure for the 

purposes of this monograph have confirmed this 

structure. Most species have an irregularly arranged 

palisade layer 10-1 5 pm thick, rarely protruding 

downward to 20 pm (Figure 4c-h). The pored epicortex (Figure 4a,b) is about 0.6 pm thick as in 

other epicorticate genera (Hale, 1973). A particularly well-developed palisade structure, here called 

columnar, is characteristic of R. flziorescens and 

R. planiusctila, two related species with a coriaceous thallus. The upper cortex consists of packed 

columnar cells 20-25 pm high overlain by the 

epicortex (Figure 5a,c). A similarly well-developed 

columnar structure is known for R. Zziteovirdir 

(Figure 5b) and R. subahstriisa (Figure 54, both 

smaller noncoriaceous species, where the cortex is 

about 20 pm thick. 

ALGAL LAYER.-ThiS layer is difficult to delimit 

in SEM photographs. It is often simply an area of 

loosely organized hyphae 10-20 pm thick, not 

sharplv differentiated from the cortex above or the 

medulla below. Algae are scattered sparsely in this 

7one (Figure ld,f). 

VEGETATIVE PROPAGULES.-one Of the mOSt un- 
NUMBER 26 3 

FIGURE 1.-Morphology of Relicina: a: isidia of R. arnphithrix (Hale 30066) (about X 10); 

b, lobules of R. schizospatha (Groenhart 2219) (about X 10); c, lobules of R. arnphithrix (Hale 

30287) (about X 10); d, R. abstrusa (Oberruinkler and Poelt); e, R. acrobotrys (A’eervoort 60 

pp.); f, R. arnflhithrix (Hale 30051); g, R. circurnnodata (Hale 26580); h, R. connivens (Hoog 

land 4726); i, R. echinocarpa (Hale 29659). (Scale in lower right corner of d-i is 10 km.) 
4 ShlITHSONIAN CONTRIBUTIONS TO BOTANY 

FICWRE 2.--Morphology of Relicina: a, R. eximbricata (Swanson C-745); b, R. puorescens (Hale 

29248); c, R. incorigrua (Malme s.n.); d, R. lirnbata (Degelius '4-368); e, R. luteoviridis (hleijer 

13194d); 1, R. malesiana (Hale 25370); g, R. planiuscula (Hale 26806); h, R. precircumnodata 

(Hale 25004a); i, R. rarnosissima (Hale 30075). (Scale in lower right corner is 10 km.) 
NUMBER 26 5 

FIGURE 3.-Morphology of Relicina: a, R. relicinella (Nee and Mori 3766); b, R. relicinula (Hale 

24402); c, R. sarnoensis (Hale 26955); d, R. schizospatha (Groenhart 2219); e, R. subabstrusa 

(Malrne sm.); f, R. subconnivens (Hale 30054); g, R. sublanea (Groeizhart 8409); h, R. sublimbata (Kurokawa 1872); i, R. sydneyensis (Kurokawa 58622). (Scale in lower right corner is 

10 km.) 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 4.--Surface and upper palisade cortex structure in Relicina: a, b, surface of R. incongrua 

(Malme s.n.) X 500 and X 2000); c, R. acrobotrys (Neervoort 60 pp.); d, R. incongrua (Malme 

s.n.); e, R. malesiana (Hale 25370); f, R. ramosissima (Hale 30075); g, R. sublanea (Groenhart 

8409); h, R. sublimbata (Kurokawa 1872). (c-h all about X 1000.) 
NUMBER 26 7 

FIGURE 5.-Upper columnar cortex (a-d) and lower cortex (e-g) in Relicina: a, R. fluorescens 

(Hale 29248); b, R. luteoviridis (Meijer B 1946); c, R. planiuscula (Hale 26806); d, R. subabstrusa (Hale 26564); e, R. Zuteoviridis (Meijer B 1946); f, R. planiuscula (Hale 26806); g, R. 

subabstrusa (Hale 26564). (Scale of 10 pm for all figures indicated in lower right corner of 5g). 
8 ShIITHSONIAN CONTRIBUTIOSS TO BOTANY 

FIGURE &-Bulbate cilia of R. abstrusa (about X 20). 

MEDULLA.-The medulla occupies about twothirds of the thallus and varies considerably in 

density. In R. amphithrix, for example, a very 

small species, the hyphae are very loosely organized 

(Figure If). Most species, however, have a densely 

packed medulla (see in particular Figures lg and 

2h). Crystals of lichen substances encrust the 

hyphae, especially in the upper half of the medulla 

(Figure 3a,f). 

LOWER CORTFX.-The lower cortex in all species 

consists of dense pacliydermatous paraplectenchyma. It is up to 20 pm thick in larger coriaceous 

species, such as R. planizisculn (Figure 5f) but correspondingly thinner (10-16 pm) in less coriaceous 

species, such as P. luteoviridis (Figure 5e) or R. 

subabstmwa (Figure 5g). 

R~JL RATE CIr.IA.-The marginal inflated bulbate 

cilia so characteristic of Relicina were probably 

first clearly recognized by Nylander when he descril)ed P0l'melifl cii ciimiiodnta in 1883. These 

structures are produced along lobe margins with 

great regularity (Figure 6). They begin on new 

lobe5 as ordinary cilia but soon become basally 

inflated (Figure 70). The ciliate tip ma) be lost 

or remain and even become densely branched. Mature globose I,ulliae are actually 1101101v (Figure 

7b) and filled with a colorless fluid. The heavily 

carlioni7ed ~valls are uniformlv thickened and 

cornpod of densely conglu tinatecl paraplectenc1i~- 

ma. A parallel-oriented procoplectench~niato~~~ 

structure, an orientation characteristic of cilia, cannot be distinguished at the base but occurs farther along where the cilia were tapered. Inflated 

bulbae have undergone more differentiation than 

that involved simply in inflating the base of cilia. 

The function of marginal bulbate cilia is unknown. It is altogether possible, however, that they 

act as propagules. On rare occasions one can see 

lobules or even submacroscopic thalli that seem to 

have originated from detached bulbae. Since they 

do not contain algae, they would have to function 

as giant fungal diaspores and encounter suitable 

symbiotic algae before continuing growth. 

LOWER SURFACE AND RHIZ1NEs.-The lower surface is either black through carbonization or medium to very pale brown. The type of rhizine is 

broadly correlated with this color. Simple or 

sparsely branched rhizines often occur in species 

with a black lower surface, as in R. abstrzisa (Figure Sa), R. amphithrix, R. connivens, R. echinocarpa, R. exinibyicnta, R. puorescens, R. incongrua, 

R. liitcoviyidk, R. planiusciila, R. relicinella, R. 

relicinitla, R. samoensis, R. schizospatha, R. subab.rtursa (Figure 8r), and R. subconnivens. Two 

species with a brown lower surface also have simple rhizines: R. limbata (Figure 8b) and R. sydneyensis. Both R. malesiana and R. siiblimbata, 

as well as some of the other lowland species, may 

have rather densely branched, almost squarrose 

rhizines, I,ut these are always black and shiny 

without agglutination. Other species with a pale 

loiver surface, including R. arrobotl-ys (Figure 84, 

R. cimimnodatn, R. pwcirrzimnodata (Figure 8e), 

R. i.omo.sis.cinia (Figure Sj), and R. .siiblanea (Figure 8g), have densely branched, bushy, and in part 

agglutinated rhizines that form a rvoolly mat about 

1 mm thick. The rhizines differ from squarrose 

types, such as one finds in Pamielia szi2cnta Taylor, 

in the pale color, the blunt rather than pointed 

tips, and the adherence of the various ramified 

parts. I have not seen this Iiranching pattern outside of Relirino. 

A~o~~~,~1~.-,4potliecia ar very commonly produced in Rclirinn. Only two species, R. Iiiteoviiidiq 

and R. schizo.spatha, lack them. Details on frequency are presented under the species descriptions. The discs are generally almut 1-3 mm in 

diameter antl imperforate (see, for example, Figures ISf, 14c, 35b, and 16b). The rim i? smooth to 

crenate antl either coronate, that is, provided with 

black 1xill)ae (Figure 9) or ecoronate. TVhile these 
NUhlBER 26 9 

bulbae appear to be the same as bulbate cilia, they 

are in fact erect pycnidia (Figure 7c,d) with an 

apical pore, as most earlier workers seem to have 

concluded. The thallus also has the normal immersed pycnidia that one would expect to find in a 

parmelioid genus. 

Bulbate cilia (retrorse rhizines) often develop 

around the base of both coronate and ecoronate 

apothecia (Figure 9). These were first described by 

Laurer (1827) for Relicina limbata. 

SPORES.-Tlle asci contain the normal complement of eight spores as in other Parmeliaceae. 

Relicilia ~elicinella, however, produces multispored 

asci with 16-32 spores. The spores are elliptical, 

ovoid, or nearly round, between 2-6 pm wide and 

3-10 pm long. The one exception here is the unFIGURE 7.--Structure of bulbate cilia and amphithecial pycnidia: a, bulbate cilia on lobe margins 

of R. sublaiiea (Hale 30280) sectioned longitudinally (X 100); b, longitudinal section of the 

base of a bulbate cilium of R. precircumnodata (Hale 25004a) (X 400); c, amphithecial 

pycnidia of R. circurnnodata (Hale 26433) (X 170); d, vertical section of an amphithecial 

pyciiidium of R. circumizodata (Hale 26433) (X 500). 
10 ShIITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 8.-Rhizine structure of Relicina: a, R. abstrusa (Oberwinkler and Poelt); b, R. limbata 

(Degelius A-368); c, R. subabstrusa (Malme s.n.); d, R. acrobotrys (A'eervoort 60 pp.); e, R. 

precircumnodata (Hale 25004a); f, R. ramosissima (Hale 30075); g, R. sublanea (Groenhart 

8409). (All about X 80.) 
NUMBER 26 11 

FIGURE 9.-Ornamentation of apothecia: coronate rim of R. 

circumnodata (left) and retrorse rhizines of R. fluorescens 

(both about X 15). (Drawings by N. Halliday.) 

usual bicornute spore of R. precircumnodata and 

its vegetative morph R. circumnodata. These are 

2-3 X 10-13 pm in size, similar to the spores of 

Bttlbothrix bicor-nuta (Muller Argau) Hale. 

Chemistry 

The chemistry of Relicina was first summarized 

by Hale and Kurokawa (1964), who used microcrystal tests and later some thin-layer chromatography (TLC). All of the specimens have been 

reexamined, as far as possible, with TLC (Merck 

silica gel precoated plates) for this revision, using 

two solvent systems, hexane-ether-formic acid and 

benzene-dioxane-acetic acid. The substances identified in Relicina are listed below. Species with chemical strains are listed two or more times. 

Alectoronic acid: R. fluorescens. 

Atranorin: Sporadically produced in R. acrobotrys, R. limBarbatic acid: R. abstrusa, R. acrobotrys, R. incongrua. 

Caperatic acid: R. subconnivens. 

Consalazinic acid: R. acrobotrys. 

Diffractaic acid: R. abstrusa, R. incongrua. 

Echinocarpic acid: R. amphithrix, R. echinocarpa, R. @ores4-0-Demethylbarbatic acid: R. abstrusa, R. acrobotrys, R. 

Fumarprotocetraric acid: R. exirnbricata, R. malesiana, R. 

Gyrophoric acid: R. luteoviridis. 

Norstictic acid: R. abstrusa, R. limbata, R. subabstrusa, R. 

Protocetraric acid: R. circumnodata, R. precircumnodata, R. 

“Quintaria” unknown: R. abstrusa. 

Protolichesterinic acid: R. connivens. 

Salazinic acid: R. abstrusa, R. acrobotrys, R. echinocarpa, R. 

bata, K. planiuscula, and R. sydneyensis. 

cens, R. planiuscula, R. samoensis, R. schizospatha. 

incongrua. 

ramosissima, R. relicinella, R. sublimbata. 

sydneyensis. 

sublanea. 

subabstrusa. 

Stictic acid: R. limbata, R. sydneyensis. 

Succinprotocetraric acid: R. eximbricata, R. ramosissima, R. 

Usnic acid: All species of Relicina. 

P+ unknown: R. relicinella. 

C+ unknown: All species containing echinocarpic acid. 

Other unknowns: R. fluorescens (with alectoronic acid). 

No medullary substances: R. amphithrix, R. relicinula. 

sublimbata. 

The identification of these acids with TLC presents no special problems. The protocetraric acid 

and related acids are separated most effectively in 

hexane, where fumarprotocetraric acid is highest, 

and succinprotocetraric acid lowest with protocetraric acid falling between them. Separation improves with an extra 10-20 ml of ether added to 

the solvent. Both barbatic and 4-0-demethylbarbatic 

acids occur in trace amounts; diffractaic acid occurs in high concentration. Species containing 

echinocarpic acid produce several additional spots, 

none of which have been identified. 

An unusual feature of Relicina is the predominance of P+ depsidones and the unidentified echinocarpic acid, which appears to be P+ depside. 

Echinocarpic acid seems to be especially characteristic of Relicina, although it occurs rarely in 

Hypotr-achyna, usually in conjunction with barbatic acid (Culberson and Hale, 1973), and in 

Parmotrema along with protocetraric acid (Hale, 

197 1). 

The substances occur in several patterns. A common one is acid A or acid A + B, as in R. acrobotryc (salazinic acid or salazinic acid with barbatic acid). An acid-deficient pattern (acid A or 

no acid) is evident in R. amphithrix (no medullary 

substance or echinocarpic acid present). Both R. 

echinocarpa and R. fluorescens show the pattern 

acid A, acid A + B, and acid B. With R. echinocarpa this involves salazinic and echinocarpic acids 

and with R. flziorescens alectoronic and echinocarpic acid?. Other examples of variation are discussed under the various species. I prefer not to 

treat these chemical populations as species because 

of the absence of any strong correlating morphological or geographical characters. 

Ecology and Habitats 

Relicina occupies two main habitats, the canopy 

branches of dipterocarps and other trees in the 

lowland rain forest and the trunk and branches of 
12 S>lITHSOZJIAN CONTRIBUTIOTS TO BOTANY 

Qzie)ciis at high elevations. A few species grow in 

secondary forests on trees such as coconut or even 

on rocks. 

The typical lowland dipterocarp forests occur in 

dense stands from sea level to less than 1000 m 

elevation. The highest locality at which I collected 

on tlipterocarp was 850 m in Negros Occidental, 

Philippines. Those species which occur in this elevational range (and often only up to 400 m) 

include R. circiimnodata, R. conniven y, R. maletiana, R. precircnmnodntn, R. rarnosissimn, R. relicitiiila, R. wbabstrtisa, R. subconnivens, antl R. 

tiiblanea. In the New T\'orld one finds R. incongrrra in a similar range of elevation. In almost all 

case5 I collected these species in areas where trees 

were being felled commercially, and since the dipterocarp forests do not usually regenerate, Relicinn 

could become extinct as the forests are timbered 

throughout southeast Asia. 

Oak and, on Luzon, Philippines, Pinii~ make up 

the typical vegetation above about 1400 m elevation on mountains in Southeast Asia. The characacteristic species here are R. acrobotyrs, R. fliioresfen$, R. liitroviridis, R. planiuscziln, and R. 

tchizo7t)atho. These oak forests are being cut over 

for vegetable farms and as with the dipterocarp 

forests many of the Relicinn species will eventually 

hecome extinct. 

Several temperate species occur on trees and 

rocks at relatively low elevation. These include 

R. echinocnrpa, R. limbata, and R. sydneyensis. 

Also occurring at low elevation but in disturbed 

forest5 in tropical regions are R. samoensis antl 

R. cxim bi icatn. Relicinn wblim batn seems to occur 

in drier subtropical areas of Burma and Thailand 

at mid elevations. 

Two species, one restricted to Asia (R. ampkithrix) and the other pantropical outside of Africa 

(R. nbrtriiro), have very great elevational ranges, 

from 150 m to over 1600 m. 

Phyto geo gr aph y 

The most distinctive feature of Relicinn is the 

high degree of endemism in Southeast Asia. Outside of R. abstrzim and R. wbabstrzisn, which occur 

in the Neotropics a5 well as in Asia, and three neotropical endemics (R. eximbl-icntn, R. incongrua, 

R. relicinella), all remaining 19 species occur only 

in the Old TVorld. At the same time the genus is 

as yet unknown in Europe, India and central Asia, 

and Africa, except for a single isolated collection 

of R. szibnbstrzisa on the Comoro Islands near 

Madagascar. 

The genus obviously evolved in the ancient 

dipterocarp forests of Southeast Asia. The endemic 

neotropical species may be remnants of older more 

widespread species, although they are unrelated to 

any species now found in the Old World. Alternatively they may have evolved separately in the 

New 12rorltl where conditions have obviously not 

been favorable for rapid evolution and expansion 

of the genus. 

The number of species known in various geographic areas is summarized in Figure 10. The 

actual species by region are tabulated as follows: 

Japan: R. nbstrusn, R. echinocnrpa, R. planiusculn, and R. 

sydn eyensis. 

Taiwan: R. nbstrirsn, R. nanlesiann, R. plnniuscula, R. subnbstrirsn, and R. sydneyensis. 

Philippines: R. nhstrusn, R. ninphithrix, R. circumnodntn, R. 

conniz~ens, R. mnlesinnn, R. plnnizisciiln, R. precircumnodntn, R. roniosissinzn, R. reliciniiln, R. snmoensis, R. schizospothn, R. sitbnbstritso, and R. sublnnen. 

hfalaya: R. nbstrusn, R. ninphithrix, R. circumnodntn, R. 

plan iitsc~iln, R. rninosissiinn, R. relicin iiln, R. schi:ospathn, 

R. s~cbconnivenr, and R. srihlnnen. 

Indonesia: R. nbstritsn, R. acrobotrys, R. connivens, R. planiusculo, R. minosissinin, R. relicinuln, R. samoensis, R. 

schizospnthn, R. sciblnnen, and R. sydneyensis. 

Sarawak: R. nbstrzisn, R. circiimnodntn, and R. subnbstrusn. 

Sabah: R. nrrobotrys, R. nmphithrix, R. fluorexens, R. luteo- 

?tiridis, R. ~nnlrsinnn, R. plirniirscitln, R. rnmosissimn, and 

R. schizospnthn. 

New Guinea: R. connivens, R. fluorescens, and R. malesinnn. 

Australia: R. limbntn, R. subnbstrnsn. and R. sydneyensis. 

Pacific Islands: R. connizxws and R. snmoensis. 

Thailand-Burma: R. siiblimbnto. 

Comoro Islands: R. sit bn bstrusn. 

U.S..A,: R. nbstrtm and R. eximbricntn. 

hlexico: R. nbstrusa. 

IVest Indie?: R. nbstrusn and R. eximbricntn. 

Panama: R. nhstrtisn and R. incongrira. 

Colombia: R. nbsfrusn, R. relicinella. 

Venezuela: R. nbstrusn. 

Brazil: R. nbstriisn, R. incongrzln, R. relicinello, and R. subParaguay: R. nbstritsn and R. sribnbstrusn. 

Argentina: R. nbstrusn. 

nbst rusn. 

The commonest species of Relicina are as follows lvith number of collections examined: R. subob.ttrii,ro (52), R. omphithrix (43), R. circiimnodnta 

(40), R. plnrii?i.tcziln (37), R. reliciniila (31), R. 

nbstrir.tn (26), and R. schizospnthn (20). Rarities 
NUMBER 26 13 

? 

include R. luteoviridis (3), R. precircumnodata 

(3), and R. siibconniveizs (1). 

The localities where especially large numbers of 

specimens were collected include logging areas in 

Negros Occidental, Philippines (44 collections), 

Selangor, Malaya (40), and Mountain Province, 

Philippines (36). The richest localities had six or 

seven species each with considerable differences in 

species composition. 

Evolution and Speciation 

Very little is known or even hypothesized about 

evolution and speciation in lichens. Lacking any 

fossil record, lichenologists are left with few clues 

that can be used to trace evolution of species and 

genera. Chemistry has proven to be a valuable 

independent check in some groups, especially the 

Parmeliaceae. I have already discussed in some 

detail the evolution of species in Hypotmchyna 

(Hale, 1975), where hybrid chemistries and rich 

development of vegetative morphs permit one to 

draw a fairly complete picture of speciation. By 

comparison, Rrlicintl has fewer vegetative morphs 

and the chemistry is less varied. The structure of 

the most characteristic acid, echinocarpic acid, is 

as yet unknown. The following discussions, therefore, will cover only the more obvious lines of 

evolution. 

MORPHOLOGICAL EVOLUTION 

FomrATIoN OF MORPHS.-A vegetative morph is 

defined as an isidiate or sorediate population which 

is absolutely identical in chemistry and morphology 

(excepting the diaspores) to a fertile, nonisidiate 

or nonsorediate parent population. A discussion of 

the development of the theory of morps in lichens 

is presented by Hale (1975). In Hypot?”aclzyna, for 

example, more than half of the speciation can be 

attributed to the formation of morphs. A similar 

situation is known in Paimotl-emn (Hale, 1965a; 

Cullierson, 1973). 

Relirintl is exceptional in that soredia are not 

formed by any of the species. Isidia do occur, however, in five species, and four of these can be 

paired witli nonisidiate sexual morphs: 

R. 0 bsf YlLsn-R. Sli 110 bstru~n 

R. rimi in ii odatn-R . precircu in iiodnto 

R. ~~la?riz~scziln-R. fliiorcsrciis (echinocarpic acid population) 

R. sydiieyensis-R. limbata 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

The acid deficient population of R. amphithrix 

is closely related to nonisidiate R. relicinula and 

may represent a vegetative morph, but variation in 

isidia and lobule development is so great that the 

evidence for a direct derivation is poor. 

RHIZINES.-The type of rhizine, simple or 

branched, appears to be an important character. 

The five species with branched agglutinated, pale 

rhizines seem to form a close group that originated 

in and has not migrated out of the dipterocarp 

forests. One, R. acrobotrys, occurs only in high 

elevation forests and, perhaps significantly, is chemically completely distinct from the other four 

species in containing salazinic and barbatic acids. 

The other species contain protocetraric acid or 

fumarprotocetraric acid and occur only at low 

elevations. One series, including R. 1-amosissima 

antl R. rziblanea, is ecoronate and has small ovoid 

spores; the other series, including R. precircumnodata antl its isidiate morpli R. circzimnodata, is 

coronate and has cornute spores. The main lines 

of evolution here seem to be ecological isolation 

and differentiation in apothecial and spore characters (Figure 11). 

CORONATE APOTHECIA.-I am assuming that ecoronate species are more primitive than coronate 

species and gave rise by mutation to coronate 

species. This seems fairly evident in the R. sublanea-R. cimimnodnta group discussed above 

where the coronate condition is paralleled by spore 

evolution and where chemistry is not significantly 

changed. A somewhat different pattern can be 

Progeniror 

Ecoronate progenitor Coronate progenitor 

R. acrobotrys /\ 

4 

I 

R. sublanea R. prerirci~mnodata 

J 

R. raniosissima 

R. rirriimnodata 

FIGURE 11.-Hypothetical evolution of species with branched, 

pale rhizines. 

seen in R. connivens (ecoronate) and R. subconnivens (coronate) where the spores have not changed 

but the chemistry is altered (protolichesternic acid 

versus caperatic acid). In most other species of 

ReZicina, however, parallel ecoronate-coronate 

populations are not closely matched, or as in the 

R. eximbricata group in the New World no ecoronate parent populations are extant. Conversely no 

coronate species have evolved from the R. limbata 

group. 

CHEMICAL EVOLUTION 

Chemical evolution is presumed to involve 

genetic processes that lead to differentiation of a 

parent species into two or more chemical populations. The patterns may be extremely clear-cut and 

easy to follow, as in Hypotrachyna (Hale, 1975), 

or too complex to interpret. Relicina has a less 

varied chemistry but several examples can be 

given. One is the difference between the two main 

populations of R. abstnisa and R. subnbstrzisa. All 

specimens of these two related species contain 

approximately equal amounts of norstictic acid 

and salazinic acid in the New World. Most specimens in the Old IVorld, however, even though 

externally indistinguishable from those in the New 

T.Vorlc1, usually lack salazinic acid or have at most 

trace amounts. The constancy of the difference 

suggests that the two major populations have been 

isolated for a long period and that no genetic 

interchange has occurred during this period of 

isolation. The New World populations have in 

fact begun to evolve in another direction, if we 

correctly interpret the evidence from a single Brazilian collection, which contains diffractaic and 

barbatic acids, indicating a backcross with a parent 

morpli containing diffractaic acid, similar for 

example to R. incongrua. If true, we can expect 

other chemical evolution in the New World, especially since apothecia are common in both the 

isidiate anti noni5idiate populations, whereas the 

more conservative Old World isidiate population 

usually lacks apotliecia and gives no evidence of 

continuing chemical evolution. 

Another example of possible backcrossing can 

be cited for the R. fEuoYescens group. The two 

species here, R. piiol-escens and R. planiurciila, are 

morphologically very close because of the strongly 

developed columnar cortex and similar habitats at 
NUMBER 26 15 

higher elevations. The presumptive parent, R. 

Puorescens, has a complex chemical population 

structure, alectoronic acid alone, echinocarpic acid 

alone, and the two acids occurring jointly (Figure 

12). The three populations occur together in significant numbers in Sabah. The vegetative morph, 

R. planiuscula, contains only echinocarpic acid. If 

we are correct in assuming that R. planiuscula 

evolved from R. fluorescens or a sexual morph 

similar to it, as shown in Figure 12, then the 

morph must have developed before R. fiuorescens 

underwent chemical evolution. A corallary of this 

assumption is that the progenitor population contained echinocarpic acid. Perhaps the alectoronic 

acid-containing populations of R. fiuorescens have 

not had time to develop isidiate morphs, or if they 

have evolved they cannot, for reasons we cannot 

even guess at this time, compete with the now 

widespread echinocarpic acid-containing morph 

R. planiuscula. 

Another example of how species have diverged 

slightly in chemistry and apparently become reproductively isolated is R. eximbricata and R. relicinella in the New World. While these externally 

similar species share the production of fumarprotocetraric acid, R. eximbricata also produces succinprotocetraric acid, while R. relicinella contains an 

unidentified P + compound, also apparently related to protocetraric acid. More significantly, R. 

relicinella has multispored asci. 

The species containing echinocarpic acid form 

the niicleiis of the genus in Southeast Asia. We 

cannot speculate on the chemical evolution of this 

group until the molecular structure of echinocarpic 

acid is determined. 

Progenitor 

R. fruoresrens R. frtdoresrens 

[echinocarpic acid] [alectoronic acid] 

R. planiiisrrda R. fruorescens 

[echinocarpic & alectoronic acids] 

FIGURE 12.-Hypothetical evolution of chemical populations 

in R. fluorexens and R. planiuscula. 

A common situation in Relicina is the differentiation of species into parallel chemical groups 

with various combinations of fumarprotocetraric 

acid and other substances. For example R. sublimbata and R. nzalesiana are essentially identical but 

R. sublimbata produces succinprotocetraric acid 

along with fumarprotocetraric acid, while R. malesiana produces only fumarprotocetraric acid. In 

this instance the species have a different geographic 

range, R. sublimbata occurring in Burma and 

Thailand, R. malesiana from Taiwan south to 

New Guinea apart from the Asian mainland. 

Position of Relicina in the Parmeliaceae 

Relicina has an isolated position in the Parmeliaceae because of its morphology and phytogeography. First, it has nothing to do with Xanthoparmelia, with which it shares only the yellow 

cortical pigment usnic acid. The only closely 

related genus is Bulbothrix (Hale, 1974), which 

also has marginal bulbate cilia and coronate apothecia but which produces only atranorin in the 

cortex. Bulbothrix, however, is rare in the range 

of Relicina. Only B. subdissecta (Nylander) Hale 

and B. pigmentarea (Hale) Hale occur in the lowland rain forests of Southeast Asia and B. Jubinflata (Hale) Hale in oak forests where one finds 

Relicina. Bulbothrix is largely a genus of the drier 

subtropical zones, being especially common in 

southeastern United States and Africa. It generally 

has larger spores, 12-19 pm long, twice the average 

size of those in Relicina, although a few species do 

have spores only 2-4 pm long. The chemistry of 

Bulbothrix is different in many respects and there 

is little evidence of any significant chromosomal 

exchange with Relicina that might appear in the 

chemistry. Many species of Bulbothrix, for example, have gyrophoric or salazinic acid, both of these 

very rare in Relicina. No barbatic, difiactaic, echinocarpic, furmaprotocetraric, or stictic acids are 

produced. On the other hand, Bulbothrix produces 

lecanoric, lobaric, and colensoic acids, all unknown 

in Relicina. I would argue then that Bulbothrix 

and Relicina as they presently stand are not closely 

related biologically. This does not rule out the 

possibility that they have a distant common origin 

which may have centered on the usnic acidatranorin split. 
16 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Key to Species of Relicina 

1. Thallus isidiate, isidiate-lobulate, or lobulate. 

2. Isidia distinct, cylindrical. 

3. Lower surface pale brown. 

4. Rhizines densely branched. 4. R. circumriodata 

4. Rhizines simple. 24. R. sydwyensis 

3. Lower surface black. 

5. Lobes very narrow and appressed, 0.5-1.0 mm wide 3. R. amphithrix 

5. Lobes broader, adnate, 1-3 mm wide. 

6. Medulla K+ red. 1. R. abstrusa 

6 Medulla K-. 13. R. planiusda 

7. Lobes narrow and appressed, 0.5-1.0 mm wide. 3. R. amphithrix 

7. Lobes broader and adnate, 1-3 mm wide. 

8. Medulla C+ red; lobules densely developed. 11. R. luteoviridis 

8 Medulla C- or dull )ellowish red (and Pf red); lobules not dense. 

9. Thallus coriaceous; upper cortex columnar. 13 R. planiuscula 

9. Thallus thinner; upper cortex not columnar. 19. R. schizospatha 

2. Isidia becoming dorsiventral and lobulate or only lobules present. 

1. Thallus lacking isidia and lobules (some adventitious marginal lobules may be present). 

10. Lower surface tan to pale brown. 

11. Rhizines simple. 

11. Rhizines densely branched. 

10. R. lirnbata 

2. R. acrobotrys 

14. R. precircumnodata 

22. R. sublanea 

15. R. ramodssima 

12. Medulla K+ red. 

12. Medulla K-. 

13. Apothecia present, coronate. 

13. Apothecia ecoronate or lacking. 

14. Protocetraric acid present. 

14. Fumarprotocetraric acid present. 

10. Lower surface black. 

15. Apothecia present, coronate. 

16. Medulla K+ red. 

16. Medulla K-. 

20. R. subabstrusa 

17 Medulla P-. 

18. Diffractaic acid present; New World species. 

18. Diffractaic acid absent; Old World species 

9. R. incongrua 

17. R. relicinula 

2 1. R. subconnivens 

18. R. sanioensis 

7. R. eximbricata 

16. R. relicinella 

19. Lobes narrow and appressed, 0.5-1 0 mm Tside. 

19. Lobes broader and adnate, 1-3 mm wide. 

17 Medulla P+ orange red. 

20. Echinocarpic acid present; Old World species. 

20. Fumarprotocetraric acid present; New World species. 

21. Spores 8/ascus. 

21. Spores 16-32/ascus. 

15. Apothecia ecoronate or lacking. 

22. Medulla K+ red. 

23. Salazinic acid only present. 

23. Norstictic and salazinic acids present. 

24. Medulla P -. 

6. R. echinocmpa 

20. R. subabstrusa 

22. Medulla K-. 

25, Lobes narrow and appressed, 0.5-1 mm wide. 

25. Lobes broader and adnate, 1-3 mm wide. 

17. R. relicinula 

8. R. Puorescens 

5. R. connivens 

26 Upper cortex columnar; alectoronic acid present. 

26. Upper cortex not columnar; protolichesterinic acid present. 

24. Medulla Pf orange red. 

27. Echinocarpic acid present 

28. Upper cortex columnar. 

28. Upper cortex not columnar. 

8. R. puorescem 

6. R. echinocarpa 
NUMBER 26 17 

Relicina (Hale and 

Pnrmelia subgenus 

Bicornutae series 

Type-species: 

27. Fumarprotocetraric acid present. 

29. Succinprotocetraric acid present. 

29. Succinprotocetraric acid lacking. 

23. R. sublimbata 

12. R. rnalesiana 

Species Treatment 

Relicina 

Kurokawa) Hale, 1974:484. 

Purmelia section lmbricaria subsection 

Relicinae Hale and Kurokawa, 1964: 135. 

Parmelia relicinn Fries, 1825:283. 

My typification of the genus (Hale, 1974) was 

not completely accurate since Parmelia relicinn 

must be designated as the type-species even though 

it becomes a tautonym when transferred to Relicinn. Furthermore, my typification of Pnrmelia 

eumorphn Hepp, based on an incorrectly determined specimen in the Rijksherbarium (L), which 

I considered to be a valid type, is in error. As 

Muller Argau (1882) had pointed out, Hepp’s 

species is a Coccocnrpia and the specimen which 

Muller Argau examined (in G) is indeed a Coccocnl-pin, although the same number in L is a 

Relicinn species. Finally, Hepp’s protologue obviouslj refers to a nonparmelioid genus. 

The species are listed alphabetically in the listing below. All collections by Hale are deposited 

in the National Museum of Natural History, 

Smithsonian Institution, and the herbarium acronym (US) is not given. 

1. Relicina abstrusa 

FIGURE 13a 

Relicinn nbstrusu (Vainio) Hale, 1974:484. 

Parnzelin nbsti ma Vainio, 1890:G4 [ripe-collection: Caraca, 

Minas Gerais, Brazil, 1/ni?iio 1347 (not 1346) (TUR, lectotype; UPS, isolectotype)]. 

Thallus closely adnate on bark or rocks, 6-10 

cm in diameter, pale chartreuse yellow; lobes sublinear, 1-2 mm wide; upper surface plane to convex, continuous or faintly maculate, moderately 

isidiate, isidia short, mostly simple: bulbate cilia 

moderately inflated; lower surface black and rliizinate, the rhi7ines simple, shiny (Figure 80). Apothecia adnate, 1-4 mm in diameter, coronate, 

basally retrorsely rhizinate, the amphithecium 

isidiate, the disc carob brown; spores 8, 4-5 X 

5-6 pm. 

CHEhiIsTRY.-I1/Iedulla Kf yellow turning red, 

C--, KC-, Pi- orange, norstictic and usnic acids 

with or without salazinic acid and very rarely with 

diffractaic acid and a trace of barbatic acid and 

4-0-denietliylbarbatic acid. A “quintaria” unknown, 

the reddish pink spot just below norstictic acid in 

both solvent systems, occurs in about one-quarter 

of the specimens. 

Rmr.mKS.-This is a very common species in 

both the New 1170rld and the Old World. As with 

R. szibabstmsn, the presumptive nonisidiate parent, the Sew World specimens produce norstictic 

and salazinic acids in nearly equal concentration, 

but salazinic acid is either absent or present in 

trace amounts in Old 1170rld specimens. The two 

populations also differ in elevational range. In 

Southeast Asia R. nbstix~n occ~irs commonly from 

100 m to 1400 m elevation. The same species in 

South America ranges from 700-2150 m. Xpothecia 

are much more common in the New Tl’orld populations (5Sm, frequency) than in the Old Ii’orld 

(7yo frequency). 

One specimen from Brazil (Lichenes nirsfronmeiicnni 95) contained diffractaic acid and a trace 

of barbatic acid in addition to the main components listed above. This unusual combination of 

acids could have originated as a backcross with a 

species such as R. incongrirn, which contains only 

diffractaic and barbatic acids. 

Vainio (1 890) considered Parmelin lim bntn f. 

i,sidio.cn hfuller Argau to be a synonym of P. nb- 

.rfriisa, but Muller’s taxon is now recognized as 

Relicinn ,xydney~?i,si,r. A\saliina (1951) identified norstictic and stictic acids in Japanese specimens, 

whicli lie identified as P. nb.rt~ir.rn and these too 

represent Rclirinn ,sydneyei?sis. 

SFECISI~~S Exh\rIxrD.--Florida: Baker Coniltv. HI!/<’ 21968. 

Xlexico: Chiapas. Hnlf 20028 (C. ITS), 20120, 20898. 2014,;. 

Panama: Panama, Hnlr 88116. 38447. Jnniaic:i: Ci/lber.ioti 

13748 (DKTKE), Plift (1-S). Coloml,ia: r.iiidig 701 (RJI. P). 

l’enezuela: Federal, Ol~rrri’i~iklrr nnrf Poclt (\I. t.S): Merida,

Hale 42070, 48886; 3Ierindn. I~cii.csclii 289ic (t’q). R~i~il: Rio 

de Jatieiro. Eitrti 6541, 7175R. 7671 (1‘s). Glntioic 2000 (I’PS): 

Minas Gerais, I’nitlio 1589 (TI’R. Ill’s): Rio C.r:inde do Sill, 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 13.--Species of Relicina: a, R. abstrusa (Hale 20393); b, R. acrobotrys (h'eeruoort GO pp.); 

c, R. amphithrix (Hale 30051); d, R. circumnodata (Maingay in BM); e, R. conniuens (Hoogland 4526); j, R. echinocarpa (Hale 29626). (Scale in mm.) 
NUMBER 26 19 

Rambo 52, 91 (US); Mato Grosso, Malme 1481B (S), 1537 (S), 

1857 (BM, S), 1865B (S), 1884 (S, W), 2745 (S), 2749B (S), in 

Lichenes austroamericani 94 (G, H, UPS), 95 (G, H, LD, S, 

UPS, US). Paraguay: Paraguari, Malme 1537 (S, US). Argentina: Misiones, Osorio 4666 (WIS). Japan: Hyuga, Hale 

29617, 29646, 29647, 29667; Kii, Kurokawa 59085 (TNS, US). 

Philippines: Mountain, Hale 25769, 25818, 25828, 25881, 

25888; Cagayan, Hale 25625, 25645, 25674, 25675, 25734; 

Negros Occidental, Hale 26432. Malaya: Pahang, Hale 30090, 

30148, 30149; Selangor, Hale 29927, 30058, 30059, 30072, 30090, 

20149, 30296, 20397, 30298. Sarawak: Hale 29993. Java: Groenhart 5868 (US), A'eercioort 4110 (BO). See Kurokawa (1965: 

265) for records for Taiwan. 

2. Relicina acrobot ys 

FIGURE 13b 

Relicina acrobotrys (Kurokawa) Hale, 1974:484. 

Parmelia acrobotrys Kurokawa in Hale and Kurokawa, 1964: 

142 [type-collection: Tjibodas, Java, Neeruoort 60 pr. p. 

(BO, holotype; US, isotype)]. 

Thallus adnate on bark, rather coriaceous, 6-12 

cm in diameter; lobes sublinear-elongate, 1-2.5 

mm wide, narrowly black rimmed; bulbate cilia 

tapered, not strongly inflated; upper surface plane 

or rugulose, continuous; lower surface black, 

densely rhizinate except for a narrow brown zone 

near the tips, the rhizines black, irregularly densely 

branched (Figure 8d). Apothecia numerous (frequency loo%), adnate, 1-2.5 mm in diameter, 

ecoronate, with basal retrorse rhizines; spores 8, 

2-3 X 4-6 pm. 

CHEMIsTRY.-Medulla K+ yellow turning red, 

C--, KC- or KC+ orange, P-, atranorin, usnic, 

consalazinic, and salacinic acids with or without 

barbatic acid and 4-0-demethylbarbatic acid. 

HABITATS.-on trunk and branches of Quercus 

in open forests at 1400-1600 m elevation. 

DIsTRIBuTIoN.-Java and Sabah. 

REXfARKs.-This is the only species with bushy 

agglutinated rhizines that occurs at high elevation. 

It is also the only species in Relicina with salazinic 

acid as the main chemical component. While the 

type contains only salazinic acid, specimens collected later in Sabah all contain barbatic acid in 

addition. Barbatic acid is also produced in trace 

amounts by R. abstrusa and R. incongrzta, but 

these New World populations have no relationship 

whatsoever to R. acrobotrys. 

Indonesia: Java, Kurokawa 2167 (TNS, US). 

SPECIMENS EXAMINED.-sabah: Hale 28126, 28540, 29123. 

3. Relicina amphithrix, new species 

FIGURE 13c 

Thallus appressus, corticola, viridi-flavicans, 2-6 

cm diametro, lobis sublinearibus, angustis, 0.3- 

1 .O mm latis, margine lobulascentibus, lobulis raro 

palmato-dactyloideis, bulbo-ciliatis, bulbis inflatis, 

congestis, aetate apice ramosis, superne dense isidiatis, isidiis cylindricis, cylindricis vel sparse ramosis, pro parte isidiato-lobulascentibus; cortex 

superior 8-10 pm crassus, stratum gonidiale 10 pm 

crassum, medulla 30-40 pm crassa, cortex inferior 

9-10 pm crassus (Figure 10; subtus niger, dense 

rhizinosus, rhizinis nigris, nitidis, simplicibus vel 

sparse ramosis. Apothecia rara, adnata, usque ad 

1 mm diametro, amphithecio coronato, basin retrorso-rhizinato, disco imperforator, hymenio 30-40 pm 

alto, sporis octonis, 3 X 4 pm. 

CHEMISTRY.-hfedulla K - , C -k , KC + reddish, 

P+ red or medulla not reacting with color tests, 

usnic and echinocarpic acids and associated unknown substances or only usnic acid present. 

HoLoTYPE.-oak forest, elevation about 1600 m, 

Cameron Highlands, Pahang, Malaya, M. E. Hale 

30203, 2 March 1965 (US). 

REMARKs.-This is the most variable species in 

Relicina and may well represent more than one 

species. The typical material is clearly isidiate, but 

the isidia may become procumbent or lobulate. In 

the extreme case the lobules are far more numerous 

than isidia over the thallus surface. Intergradation 

with R. schizospatha is a distinct possibility although the two species are fairly easily separated 

on thallus size and lobe width. A small group of 

specimens has closely appressed, mostly marginal 

palmately branched lobules (Figure Ic) often 

occurring without isidia but seeming to originate 

from or to intergrade with isidia when isidia are 

present. 

Relicina amphithrix is probably related to R. 

relicinicla since both have very narrow lobes and 

coronate apothecia. Relicina wlicin !/la, hoicever, 

does not produce echinocarpic acid and would 

therefore differ from the bulk of the specimens 

identified as R. amphithi.ix. On the other hand. it 

seems probable that the acid deficient population 

of R. amphithrix is the isidiate morph of R. 

relicinzila. 

SrEcrartxs EX~VINFD (echinocnipic ncid present, isidinte or 

becoming lobalnte-isidinte).--Philippiues: AIouiitnin, Hole 
20 ShIITHSONIAN CONTRIBUTIOXS TO BOTANY 

25824; Negros Occidental, Hale 26420, 26437, 26445, 26480, 

26481, 26522, 26569, 26630, 26654; Agusan, Hale 24481, 25064, 

25191; Surigao del Stir, Hale 24674. Malaya: Pahang, Hale 

30204, 30211, 30479, 20493; Selangor, Hale 30066, 30070. 

Sabah: Hale 28239, 28820, 29076. 

SPECIMENS EXARIINFD (echinocarpic acid, marginally palmatelobulate).-Philippines: hfountain, Hale 25770, 25789; Agusan, 

Hale 24462. Malaya: Selangor, Hale 30060, 30063, 30256, 

30257, 30258, 30259, 30265, 30283, 30287, 31185, 31186. 

SPECIMENS EXAMINED (echinocarpic acid absent, isidiate or 

becoming lobulate isidiate).-Philippines: Agusan, Hale 

24454. Malaya: Selangor, Hale 30051, 30064, 30065, 30069, 

50071, 30288. 

Philippines, northern Borneo, and Malaya. It has 

not yet been collected south of this region in 

Indonesia but probably occurs there ;IS well. 

SPECIX~ENS ExAhrl~ED.-Philippines: Mountain, Hale 25787, 

23802, 25810, 25820, 25841, 25873, 25902: Cagayan, Hale 

24410, 25736; Quezon, Hale 26878, 26879, 26887, 26900, 26920, 

26923, 26926, 26928, 26934, 26980; Negros Occidental, Hale 

26421, 26433, 26474, 26500, 26317, 26561, 26572, 26580, 26599, 

26631, 26633, 26651, 266.52. XIalaya: Pahang, Hale 30144; 

Selangor, Hole 30056. Sarawak: Hale 29995, 29996, 29997, 

299999, 30437. 

5. Relicina connivens 

4. Relicina circumnodata 

FIGURE 13e 

FIGURE 13d 

Relicinn circumnodata (Sylander) Hale, 1974:484. 

Parmelia circumnodata Sylander in Crombie, 1883:51 [typecollection: Government Hill, Penang, Malaya, Maingay 22 

(BM, lectotype; FH, H, isolectotypes)]. 

Tliallus closely adnate on bark, 3-8 cm in diameter; lobes sublinear-elongate, 0.7-2 mm wide; buIbate cilia dense and conspicuous, strongly inflated 

and becoming globose and apically densely 

Ixanclied; upper surface plane to convex, continuous or faintly maculate, sparsely isidiate, the isidia 

simple, to 0.5 mm high; lower surface pale brown, 

rhizinate, the rhizines pale, more or less densely 

branched and agglutinated. hpothecia rare (frequency lo%), adnate, 1-2 mm in diameter, rim 

coronate; spores 8, cornute, 3 X 10-12 pm. 

rose, Pf red, protocetraric and usnic acids. 

HABITATs.-on trunks and canopy branches of 

tlipterocarp and other trees in rain forest at sea 

level to I300 m elevation. 

DlsTRIBuTIoN.-iC~alaya, Philippines, Sarawak. 

RnrmKs.-This species was described by Nylander from sterile material. Apothecia are now 

known from 4 of 39 specimens that I have been 

able to examine. All are coronate and have unusual bicornute spores, exactly the same as those 

in the presumptive parent species R. precirciirnnodata. The bulbate cilia are very large, globose, 

and crowded on older parts of the lobes, also as in 

the parent species. The lower surface is consistently 

pale brown and the rhizines initially simple but 

soon branching in the pecular bushy manner of 

this group. 

The species is very common throughout the 

CHEX\'rISTRY.-MedU11a K-, C-, KC- or KCSRelicirm ro~iuivens (Kurokarva) Hale, 1974:484. 

Partrielin rotinivens Kurokawa irz Hale and Kurokawa, 1964: 

142 [type collection: Milne nay district, Papua, Sew 

Guinea, Hooglond 4i26 (BM, holotype; L, US, isotypes)]. 

Thallus adnate on bark, coriaceous, 3-10 cm in 

diameter; lobes sublinear-elongate, 1-3 mm wide; 

upper surface plane to convex, becoming rugulose, 

continuous to faintly maculate; bulbate cilia large, 

becoming globose; lower surface black, densely 

rhirinate, the rhizines black, siniple to irregularly 

densely branched. Apothecia common (frequency 

My;), atlnate, 1-2.5 mm in diameter, the rim 

ecoronate, the base retrorsely rhizinate; spores 8, 

2-3 X 4-6 pm. 

tolichesteric and usnic acids. 

HAnrTATs.-on trunk and canopy branches of 

tlipterocarps in rain forest and on other trees in 

secondary forest at 200-850 m elevation. 

DIsTRIBUTroN.-Phi~ippines, Indonesia, New 

Guinea, Guam, and the Solomon Islands. 

Rnr.mKs.-This species has proved to be widespread hut not common in a band from the Philippines through Xew Guinea to the Solomon Islands, 

occurring generally at 300-800 m elevation. No 

other species in the genus produces protolichesterinic acid. It pro1)ably would be mistaken for R. 

m(~le.~in~in (P+ red) without appropriate chemical 

testing. Closely related R. szibconniuens has coronate apothecia but if collected sterile it could be 

differentiated by the presence of caperatic acid. 

CHEMlSTRY.-;\/IedUl~a K-, C-, KC-, P--, proSPECI\IENS ExAzrisrD.-Pliilippines: Segroes Occidental, 

Hale 26457, 26,709, 26626a: Agusan, Hale, 25025, 25034, 25044. 

Colomon Islands: Guadalcanal, Hill 9057 (EM, US). hddi- 
NUMBER 26 21 

tional records from the Molucca Islands and Guam are listed 

in Hale and Kurokawa (1964:143). 

6. Relicina echinocarpa 

FIGURE 13f 

Relicina echinocarpa (Kurokawa) Hale, 1974:484. 

Parmelia echinocarpa Kurokawa, 1965:265 [type-collection: 

Mt. Hikosan, Prov. Buzen, Japan, Kurokawa 63163 (TNS, 

holot ype)]. 

Thallus adnate, to appressed, corticolous or saxicolous, 2-6 cm in diameter; lobes sublinearelongate, 1-2 mm wide; bulbate cilia gradually inflated; upper surface plane, faintly maculate; lower 

surface black, densely rhizinate, the rhizines simple 

or sparsely branched. Apothecia numerous (frequency 10070), adnate, ecoronate, 1-2.5 mm in 

diameter, with basal retrorse rhizines; spores 8, 

3-5 X 6-8 pm. 

CHEhiISTRY.-Medulla K+ yellow, c- or c+ 

pale yellow, KC-, Pf orange, echinocarpic and 

usnic acids and a C+ unknown with or without 

salazinic acid. 

HARITATS.-on trunk and canopy branches, 

more rarely on rocks, in open deciduous forest at 

low elevation. 

D I STR I R UTI ON. -J a pa n . 

REMARKs.-Relicina echinocarpa is a small, closely adnate species collected on upper branches of 

deciduous trees or on rocks in southern Japan, 

where it appears to be endemic. While it usually 

contains echinocarpic acid, it is not related to the 

tropical members of the genus containing this acid. 

For example, R. .mmoensis has more globose bulhe and coronate apothecia. 

The chemical variation in R. echinocarpa follows 

a typical addition pattern. Specimens may contain 

echinocarpic acid alone, echinocarpic and salazinic 

acids together, and in one or two specimens apparently salazinic acid alone. 

SP€CIMF.NS ExAMlNFD.-Japan: Hyuga, Hale 29596, 2961 3, 

29626, 29638, 29648, 29658, 29659: Higo, Mayebara (TNS). 

7. Relicina eximbricata 

FIGURE 140 

Relicinn eximbricntn (Gyclnik) Hale, 1974:484. 

Parmelia sanioensis Zahlhruckncr var. eximbricata Gyelnik, 

1938:288 [type-collection: Monte Rus, Cuba, Hioram 10506 

(BP, holotype)]. 

Parmelia eximbricata (Gyelnik) Hale and Kurokawa, 1964: 

143. 

Thallus closely adnate on twigs or bark, 3-5 cm 

broad; lobes sublinear, 0.7-1.5 mm wide; upper 

surface plane and continuous; lower surface black, 

densely rhizinate, the rhizines simple. Apothecia 

very common (frequency 100yo), adnate, 0.5-1.5 

mm in diameter, the exciple coronate, the base 

often retrorsely rhizinate; spores 8, 4-5 X 6-9 pm. 

CHEMISTRY.-bfedUlla K - , c - , KC - , P + brick 

red, fumarprotocetraric, succinprotocetraric, and 

usnic acids. 

HARITATs.-T~u~~s and branches of trees (cashew, Eugenia) in open areas from sea level to 600 m 

elevation. 

D!s~~~~uTro~.-Florida and the West Indies. 

REMARKs.-Relicina eximbricata has been collected for the most part in disturbed secondary 

forest. It is rather rare if the few collections available reflect its true abundance. I was unable, for 

example, to recollect any specimens at the locality 

in Florida visited by Swanson. It is extremely close 

to R. relicinella in external appearance but differs 

in spore number (8 versus 16-32) and presence of 

succinprotocetraric acid. 

specimens from Florida, Cuba, Grand Cayman, Jamaica, and 

the Dominican Republic. 

SPECIMENS EXAhTINED.-see Hale & Kurokawa (1964: 144) for 

8. Relicina fluorescens 

FIGURE 14b 

Relicina Jluorescens (Hale) Hale, 1974:484. 

Parmelia fluorescens Hale, 1965:202 [type-collection: between 

Kamharanga and the second radio tower, Tourist Trail, 

Kinahalu National Park, Sabah, Hale 28637 (US, holotype; 

TNS, isotype)]. 

Thallus loosely attached to adnate, coriaceous, 

corticolous, 4-7 cm broad; lobes sublinear-elongate, 

1-3 mm wide; bulbate cilia tapered, not strongly 

inflated; upper surface plane to rugulose and finely 

pitted, faintly maculate; upper cortex strongly 

columnar (Figure 5a); surface black, densely rhizinate, the rhizines simple. to sparsely branched. 

Apothecia common (frequency 50%,), adnate, 2-3 

mm in diameter, ecoronate, usually with basal 

retrorse rhizines; spores 8, 4 X 5 pm. 
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 14.-Species of Relicina: a, R. eximbricata (Imshaug 24518); b, R. puorescens (Hale 

29088); c, R. incongrua (Malme 2433C); d, R. limbata (Du Rietz 4025); e, R. luteoviridis (Hale 

29069); f, R. malesiana (Royen 6232). (Scale in mm.) 
NUMBER 26 23 

CHEMISTRY.-Medulla K-, C--, KC+ rose, or 

P- or P+ usnic and alectoronic acids with or 

without echinocarpic acid or echinocarpic acid 

alone in the medulla. 

HABITATS.-on trunk and branches of @MCUS 

and other trees in open forest at 1600 to 2400 m 

elevation. 

DIsTRiBuTIoN.-sabah and New Guinea. 

REMARKs.-Relicina fluorescens was first recognized by the presence of alectoronic acid. On later 

study I decided to lump here a series of chemically 

variable specimens occurring at high elevation, 

mostly in Sabah. All have coriaceous thalli and a 

columnar cortex. Once again the primary substance 

is echinocarpic acid with or without alectoronic 

acid, but a sizeable part of the population may 

lack echinocarpic acid. The unknown substance 

occurring with alectoronic acid in four specimens 

listed below has an RF of about 0.25 in the hexane 

solvent and about 0.6 in the benzene solvent. It is 

brownish when developed with sulfuric acid. aCollatolic acid does not occur in this species. 

The isidiate morph of R. fluorescens is R. planiuscula, which has a broader geographic range. 

SPECIMENS EXAMINED (alectoronic acid present).-New 

Guinea. Western Highland: Kurokawa 75 in Lichenes Rariores et Critici Exsiccati (US). See Hale (1965: 203) for additional records from Sabah. 

SPECIMENS EXAMINED (alectoronic and echinocarpic acids 

present).-Sabah: Hale 28510, 28512, 29048, 29248. 

SPECIMENS EXAMINED (echinocarpic acid present).-Sabah: 

Hale 28703, 28941, 29088. 

SPECIMENS EXAMINED (alectoronic acid and an unidentified 

substance).-Sabah: Hale 28322, 28750, 29031, 29053. 

9. Relicina incongrua, new species 

FIGURE 14c 

Thallus adnatus vel appressus, corticola, viridiflavicans, circa 6 cm diametro, lobis sublinearibus, 

contiguis, 0.7-1.5 mm latis, margine modice 

bulbato-ciliatis, bulbis aetate inflatis globosisque; 

cortex superior circa 10 pm crassus, stratum gonidiale 10-12 pm crassum, medulla 70-80 pm crassa, 

cortex inferior 10-12 pm crassis (Figure 2c); subtus niger, modice vel dense rhizinosus, rhizinis 

nigris, nitidis, simplicibus vel sparse ramosis. Apothecia numerosa (frequencia 100~o), adnata, usque 

ad 2 mm diametro, amphithecio coronato, basin 

retrorso-rhizinato, hymenio 45-55 pm alto, sporis 

octonis, 4-5 X 5-7 ym. 

CHEMISTRY.-Meduh K-, c - , KC -, KC +, 

P - , usnic, diffractaic, barbatic, and 4-O-demethylbarbatic acids. 

HoLoTYPE.-santa Anna da Chapada, Mato 

Grosso, Brazil, Malme 2433C, 27 February 1894 

(S; isotype in US). 

HABITAT.-oII trees in primary forests at low to 

mid elevation. 

DIsTRIBUTI0N.-Panama and Brazil. 

REMARKS.-This rare species is externally identical with R. subabstrzisa, which contains norstictic 

and salazinic acids. Since this seems to represent a 

replacement type of chemical population, I am 

considering it as a distinct species, very closely 

related to R. subabstrusa and probably derived 

with it from a common progenitor. Both of these 

species have unfortunately been only rarely collected in tropical America. 

Brazil: Mato Grosso, Alalme 2445 (S, US). 

SPECIMENS ExAMINED.-Panama: Canal Zone, Hale 43448. 

10. Relicina limbata 

FIGURE 14d 

Relicina limbata (Laurer) Hale, 1974:484. 

Parmelia limbata Laurer, 1827:39 [type-collection: Australia, 

Sieber (M, lectotype)]. 

Parmelia sphaerospora Knight, 1882:49 [type-collection: Sydney, Australia, Knight (H, lectotype) (not Parmeh 

sphaerospora Nylander, 1859:254)]. 

Parmelia insinuata Nylander, 1886:324 [based 011 Pnrmelia 

sphaerospora Knight]. 

Thallus adnate to loosely attached on bark or 

rock, coriaceous, 4-9 cm in diameter; lobes sublinear-elongate, 0.8-2.5 mm wide; bulbate cilia 

conspicuous, up to 2.5 mm long; upper surface 

plane to convex, maculate, cracked on older lobes; 

lower surface brown to blackish brown, moderately 

rhizinate, the rhizines pale brown, coarse, mostly 

simple (Figure 8b). Apothecia yery common (frequency 100~o), adnate, 1-6 inm in diameter, ecoronate, basally retrorsely rhizinate; spores 8, 4-6 X 

7-10 pm. 

P+ pale orange, norstictic, stictic, and usnic acids 

with or without atranorin. 

on trees, in open forests at low elevation. 

CHEMISTRY.-h’~edU11a K+ yellow, c-, KC-, 

HABIThT.-on rocks (granite, SaIldStOlle), rarel\ 
24 SXIITHSONIAN CONTRIBUTIONS TO BOTANY 

DIsTRrBuTIoN.-Australia and Tasmania, 

REMARKS.-ThiS well-known lichen has been collected only in eastern Australia and Tasmania, 

where it appears to be common. It was elegantly 

illustrated in color by Laurer (1827), yet Knight, 

not apparently having seen Laurer’s publication, 

described it as a new species, Parmelia sphaerospora. Knight then sent a specimen to Nylander, 

who immediately published a new name, P. insinuata, since Knight’s epithet was a homonym of 

Nylander’s own P. sphaeospora published in 1859. 

Vainio (1890) comprehended the species correctly, but Asahina’s (1934) reports from Japan are 

Relicina sydneyensis, the isidiate morph of R. 

limbata. 

SPECIMENS ExAMINED.-Australia: Queensland, Bailey 165 

(BM), Hartmann (US); New South Wales, Chase (US), Du 

Rietz 60b, 4025 (UPS, US), Home (BM), Moore in Kryptogamae Vindobonensis Exsiccntae 570 (BM, H, LD, NY, UPS, 

US), Weber and hlcVean 47322, 49914 (COLO, US), Wilson 

(TNS), in Lichenes Rariores Exsiccati 34 (BPI, NY); Tasmania, Brown (BM), Degelius A-368 (US). 

11. Relicina luteoviridis 

FIGURE 14e 

Relicina luteoviridis (Kurokawa) Hale, 1974:484. 

Parmelia luteoviridis Kurokawa in Hale and Kurokawa, 1964: 

144 [type-collection: B. Papan, Beu territory, E. Kutai, 

Borneo, Meijer B1948 (BO, holotype; US, isotype)]. 

Thallus adnate on bark, fragile, 5-8 cm in diameter; lobes sublinear-elongate, 1.5-3 mm wide; 

upper surface plane, faintly maculate, lobulate, 

the lobules large, dorsiventral, marginally bulbateciliate, procumbent, often dichotomously branched 

(Figure 14e); cortex columnar (Figure 5b); bulbate 

cilia long and tapered; lower surface black, densely 

rhizinate, the rhizines black, simple or sparsely 

branched. Apothecia not seen. 

CHEMIsTRY.-Medulla K--, C+ rose, KC+ red, 

P-, gyrophoric and usnic acids with or without 

a tranorin. 

HABITATS.-On upper branches of evergreen trees 

in rain forest at 600-1400 m elevation. 

DIs”RIsuTIoN.-Sabah. 

REMARKs.-Relicina luteoviridis can be recognized by the large lobules and the unique chemistry. It is the only species containing gyrophoric 

acid. Only three collections are known, all from 

Borneo. 

SPECIMENS ExAMINED.-Sabah: Hale 29069, hleijer B548a 

(BO, US). 

12. Relicina malesinna 

FIGURE 14f 

Relicina inalesiana (Hale) Hale, 1974:484. 

Parmelia malesiana Hale, 1965:203 [type-collection: Florida 

logging area, about 30 km southeast of Butuan City, 

Agusan Prov., Philippines, Hale 25370 (US, holotype; 

TNS, UPS, isotypes)]. 

Thallus adnate on bark, 6-8 cm broad; lobes 

sublinear-elongate, 1.5-2.5 mm wide; bulbate cilia 

gradually inflated; upper surface plane to rugulose, 

faintly maculate; medulla white, very rarely in 

part orange red; lower surface dark brown or 

blackening, densely rhizinate, the rhizines black, 

simple to squarrosely branched. Apothecia numerous (frequency 80y0), adnate, ecoronate; spores 8, 

2-4 X 3-7 pm, medulla below the hymenium rarely orange red. 

CHEMISTRY.-Medulla K--, c-, KC-, P-k red, 

usnic and fumarprotocetraric acids and if pigmented an unidentified K + purple anthraquinone. 

HABITAT.-On trunk and branches of dipterocarps and other trees in rain forest at 150-600 m 

elevation. 

DISTRIBUTIoN.-TaiWan, Philippines, Sabah, and 

New Guinea. 

REMARKS.-The lower surface of this species is 

generally black but may be in part dark brown. 

The rhizines when fully branched appear to be 

squarrose, black, rather thin, and shiny. It is somewhat related, then, to species like R. ramosissima, 

which has a consistently pale brown lower surface 

and pale agglutinated rhizines, succinprotocetraric 

acid in addition to fumarprotocetraric acid, and 

smaller spores. It is also chemically related to R. 

stiblimbata, which contains both fumarprotocetraric and succinprotocetraric acids. 

SPECIMENS ExAMINED.-NeW Guinea: Hollandia, Royen and 

Sleumer 6232 (L, US). See Hale (1965:203) for additional 

records for Taiwan, Philippines, and Sabah. 

13. Relicina planiuscula 

FIGURE 15a 

Reliciiza planiuscula (Kurokawa) Hale, 1974:484. 

Parmelia planiuscula Kurokawa in Hale and Kurokawa, 1964: 
NUMBER 26 25 

144 [type-collection: Kandang Badak, Java, Neeruoort 427 

(BO, holotype; US, isotype)]. 

Thallus adnate on bark, coriaceous, 5-9 cm in 

diameter; lobes sublinear-elongate, 1-3 mm wide; 

upper surface plane, faintly maculate, sparsely to 

moderately isidiate, the isidia simple, less than 

0.7 mm high, frequently procumbent, rarely lobulate; upper cortex strongly columnar (Figure 5c); 

lower surface black, moderately rhizinate, the 

rhizines simple, black. Apothecia rare (frequency 

11%), adnate, 1-2 mm in diameter, ecoronate, the 

amphithecium isidiate; spores 8, 4 X 5 pm. 

CHEMISTRY .-Medulla K -k yellowish, c -k faint 

yellow orange, KC - , P + yellow, echinocarpic and 

usnic acids and a C+ unknown with or without 

atranorin. 

HABITATS.-on trunk and branches of Quercus, 

Pinirr, and other trees in open forests and mossy 

forest at 1500-2200 m (at lower elevation in Japan 

and Taiwan). 

DIsTRlnUTI0N.-Japan, Taiwan, Philippines, Malaya, Indonesia, and Sabah. 

REMARKs.-This is the commonest species of 

Reliciria at higher elevations in Southeast Asia. 

The presumptive parent of this isidiate morph is 

R. flziorcsccns, which has a much more restricted 

distribution. While most specimens have sparse to 

moderate development of cylindrical isidia, others 

have procumbent lobulate isidia. The population 

on Pin ?is in the northern Philippines is especially 

strongly isidiate-lobulate, and I had at first considered it to be a separate species. 

SPECIMENS ExAhrmm.-Japan. Hyuga: Hale 29665. Philippines: Mountain, Hale 26088, 26089, 26102, 26113, 26117, 

26173, 26195, 26806. Malaya: Pahang, Hale 29929, 29943,' 

29944, 29945, 29946, 29949, 30481, 30497. Sabah: Hale 28006, 

28040, 28063, 28234, 28250, 28257, 28372, 28400, 28508, 28546, 

28580, 28581, 28603, 28700, 28815, 29035, 29168, 29250. Indonesia: Java, Groenhart 2217 (L); Sumatra, Steenis 3895 

PO). 

14. Relicina precircumnodata, new species 

FIGURE 15b 

Thallus adnatus, corticola, viridi-flavicans, 4-10 

cm diametro, lobis sublinearibus, contiguis vel separatis, 1-2 mm latis, margine dense bulbo-ciliatis, 

bulbis globosis, sorediis isidiisque destitutis; cortex 

superior circa 12 pm crassus, stratum gonidiale 10 

pm crassum, medulla 100-120 pm crassa, cortex inferior circa 12 pm crassus (Figure 2h); subtus pallide castaneus, dense rhizinosus, rhizinis pallidis, 

dense ramosis atque agglutinatis. Apothecia adnata, 

usque ad 2 mm diametro, amphithecio crenato, 

coronato, basin retrorso-rhizinato, hymenio 70-80 

pm alto, sporis octonis, bicornutis, 2-3 X 11-13 

P. 

CHEMIsTRY.-Medulla K-, c -, P-k red, protocetraric and usnic acids. 

HOLOTYPE.-virgin dipterocarp forest, Tungao 

Logging Area, 40 km SE Butuan City, Agusan 

Province, Philippines, elevation about 400 m, 

M. E. Hale 25044a, July 1964 (US). 

HABITATS.-on canopy branches of dipterocarps 

in rain forest at 100-400 m elevation. 

DrsTRIsuTIoN.-Philippines. 

REMARKS.-ThiS species represents the presumptive nonisidiate morph of R. circumnodata. Both 

contain protocetraric acid and have unusual bicornute spores. Relicina precircumnodata is by far 

the rarer of this species pair, occurring only at 

three widely separate localities in the Philippines. 

SPECIMENS ExAvINED.-Philippines: Cagayan, Hale 24418; 

Mountain, Hale 25826. 

15. Relicina ramosissima 

FIGURE 15c 

Relicifla ramosissima (Kurokawa) Hale, 1974:485. 

Parmelia ramosissima Kurokawa in Hale and Kurokawa, 

1964: 145 [type-collection: Tenimber Island, Moluccas, 

BuwaEda 4583 (BO, holotype; US, isotype)]. 

Thallus adnate, corticolous, 3-10 cm in diameter; lobes sublinear-elongate, 0.7-2 mm wide; bulbate cilia gradually inflated to gobose, becoming 

apically branched; upper surface plane, maculate; 

lower surface pale to dark brown, moderately rhizinate, the rhizines simple to densely branched and 

agglutinated, pale (Figure 8f). Apothecia common 

(frequency 93%), adnate, 1-2.5 mm in diameter, 

ecoronate, often retrorsely rhizinate at the base; 

spores 8, 2 X 3 pm. 

CHEMISTRY.-Medulla K - , c -, KC -k reddish, 

P+ red, usnic acid, fumarprotocetraric acid, and 

succinprotocetraric acid. 

HAnITAT.-on trunk and canopy branches of 

dipterocarps in rain forest at 150-300 m elevation. 
26 S.MITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 15.--Species of Relicina: a, R. planiuscula (Hale 26806); b, R. precircumnodata (Hale 

24418); c, R. ramosissima (Hale 29923); d, R. relicinella (Nee 3766); e, R. relicinula (Hale 

24402); 1, R. sarnoensis (Hale 26955). (Scale in mm.) 
NUMBER 26 27 

DIsTRIBLJTIoN.-Malaya, Philippines, Sabah, and 

Indonesia. 

REMARKS.-This species has a wide distribution 

in the dipterocarp forests. It superficially resembles R. sublanea, which differs in producing protocetraric acid, in having large spores (4-5 X 7-9 

pm), and in occurring generally above 300-400 m 

elevation, higher than R. ramosissima. 

SPECIMENS EXAMINED.-Philippines: Agusan, Hale 25245, 

25360. Malaya: Selangor, Hale 29923, 30062, 30074, 30075, 

30290, 30291, 30300. Sabah: Hale 30352, 30355, 30356, 30357, 

30366. Indonesia: Java, Groenhart 8374 (BO). 

16. Relicina relicinella 

FIGURE 15d 

Relicina relicinella (Nylander) Hale, 1974:485. 

Parmelia relicinella Nylander 1885:215 [type-collection: Santarem, Brazil, Spruce 136 (H, lectotype; BM, G, P, isolectotypes)]. 

Thallus closely adnate on bark, 3-6 cm in diameter; lobes sublinear, 0.5-1.5 mm wide; bulbate 

cilia distinct, short; upper surface plane, continuous; lower surface black, densely rhizinate, the 

rhizines simple or sparsely branched. Apothecia 

common (frequency loo%), adnate, 0.5-1.5 mm 

in diameter, the disc concave to plane, the exciple 

coronate; spores 16-32, 3-4 X 5-6 pm. 

CHEMISTRY.-Medulla K-, c-, KC-, Pf red, 

usnic acid, fumarprotocetraric acid, and an unidentified P+ compound. 

HABITAT.-on tree trunks and branches in rain 

forest at 100-200 m elevation. 

DIsTRIBuTIoY.-Co~ombia and Brazil. 

REMARKs.-The most unusual feature of this 

species is the multispored asci, recognized by 

Nylander (1885) in his original description. It is 

very similar externally to R. eximbricata but seems 

to occur only in virgin rain forest. It may be a 

common species in the vast lowland rain forests of 

South America, but no lichenologists have yet had 

an opportunity to visit these habitats. 

3766. Brazil: Amazonas, Spruce 58 (BM, US). 

SPECIMENS EXAhmxED.-Colombia: Santander, Nee and Mori 

17. Relicina relicinula, new combination 

FIGURE 15e 

Parmelia relicinula Muller Argau, 1882:317 [type-collection: 

Java, Junghuhn 3728 (G, lectotype)]. 

Parmelia relicina Fries, 1825:283 [type-collection: Gaudichaud 

54, Rawak (Pacific area) (UPS, lectotype; P. isolectotype)]. 

Thallus closely adnate to appressed on bark, 2-6 

cm broad; lobes variable, short, contiguous and 

sublinear to separate and sublinear-elongate, 0.3- 

1.5 mm wide; bulbate cilia short but conspicuous, 

strongly inflated, sometimes apically branched; 

upper surface plane, continuous to faintly maculate, without lobules or becoming marginally lobulate, isidia lacking; lower surface black, densely 

rhizinate, the rhizines black, simple to sparsely 

branched. Apothecia very common (frequency 

9473, adnate, about 1 mm in diameter, the exciple 

coronate, the base often retrorsely rhizinate; spores 

8, 4-5 X 5-7 pm. 

CHEMISTRY.-Medulla K-, c-, KC-, P-, 

usnic acid and unidentified colorless substances. 

HABITAT.-on trunk and canopy branches of 

dipterocarps and other trees in rain forest at 100- 

400 m elevation. 

DIsTRIBUTIoN.-Phi~ippines, Malaya, and Indonesia. 

REMARKs.-The characteristic features of this 

widespread species are the narrow appressed lobes 

and lack of lichen substances in the medulla. It is 

an extremely variable species, however, particularly 

in regard to production of marginal lobes. Some 

specimens have no obvious lobules. Muller’s typespecimen is somewhat lobulate. He described Parmelia relicinztla as being 2-3 times narrower than 

P. relicina Fries, but I doubt that he saw Fries’ 

type. The excellent material of P. relicina in Paris 

has very narrow lobes. This specimen does lack 

lobules. Its chemistry is still undecided since, although it is negative with color tests, a dull, high 

unidentified spot appears on TLC plates. Similar 

but not identical spots were discovered in several 

of the Philippine specimens. 

Asahina (1934) reported Parmelia relicina from 

southern Japan. I have not seen this material but 

suspect that it is Relicina echinocarpa. 

SPECIMENS EXAMINED.-Philippines: Mountain, Hale 25849, 

25871: Cagayan, Hale 24402, 24416, 25604, 25638, 25652, 

25657, 25658, 25737; Cavite, Hale 26836; Sorsogon, Elmer 

16031 (F, US): Agusan, Hale 24996, 25011, 25659; Surigao del 

Sur, Hale 24641, 24698, 24700; Zamboanga del Sur, Hale 

24521, 24704, 24709, 24733, 24738, 24748, 24807, 24826, 24827, 

24861, 25159, 25313. Malaya: Selangor, Hale 29925, 30281. 

Java: Groenhart 1702 (L). Thursday Island: Hartmann in 

Lichenotheca Universalis 159 (S, US). 
28 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

18. Relicina samoensis 

FIGURE 15f 

Relicina samoensis (Zahlbruckner) Hale, 1974:485. 

Parmelia samoensis Zahlbruckner, 1908:272 [type-collection: 

Malifa, Upolu, Rechinger 5005 (W, lectotype)]. 

Thallus adnate to closely adnate, corticolous, 4-8 

cm in diameter; lobes rather short, sublinear, 0.3-1 

mm wide; bulbate cilia conspicuous, globose, up 

to 1 mm long; upper surface plane, continuous; 

lower surface black, densely rhizinate, the rhizines 

black, simple or sparsely branched. Apothecia adnate, very common (frequency 100’%), 1-3 mm in 

diameter, the exciple coronate, the base retrorsely 

rhizinate; spores 8, 5-6 X 6-7 pm. 

CHEMISTRY.-hfedd1a K+ yellow, C + faint 

orange, KC-, P+ yellow orange, usnic and echinocarpic acids and a C+ unknown. 

HABITAT.-on coconut palm and other trees in 

disturbed areas at 100-700 m elevation. 

DIsTRrBUTION.-~hi~ippines, Java, New Cab 

donia, Samoa, and Society Islands. 

REMARKs.-Relicina samoensis is a lowland species occurring in secondary forests. The lobes vary 

considerably in size but are usually narrow and 

closely adnate. This is the only species with echinocarpic acid that has coronate apothecia. 

SPECIMENS ExAM1sED.-Philippines: Quezon, Hale 26955. 

Indonesia: Halmahera, Croenhart 8343 (BO): Java, Bredijn 

1099 (BO), Palmer 614 (US). Samoa: Reinecke 22 p. p. (BO, 

US). New Caledonia: Hill 11590 (BM). Society Islands: Moore 

L3 (US). 

19. Relicina schizospatha 

FIGURE 16a 

Relicina schizospatha (Kurokawa) Hale, 1974:485. 

Parmelia schizospatha Kurokawa in Hale and Kurokawa, 

1964: 146 [type-collection: Gegerbentang, Java, Neeruoort 

1062 (BO, holotype; US, isotype)]. 

Thallus adnate, corticolous, rather fragile, 3-6 

cm in diameter; lobes short, sublinear, 1-2 mm 

wide; hulhate cilia not conspicuous or dense; upper 

surface plane, faintly maculate, becoming more or 

less densely lobulate along the margins and surface, the lobules dorsiventral, procumbent to subascending, sometimes dactyloid branched, rarely 

with marginal hulhils (Figure 16); lower surface 

black, moderately rhizinate, the rhizines simple. 

Apothecia unknown. 

faint orange, KC-, P+ orange, usnic and echinocarpic acids and a C+ unknown. 

HABITAT.-on trunk and branches of QUeYCUS 

and other trees in rain forest at 1300-1600 m 

elevation. 

DIsTRIBuTIoN.-hfa~aya, Philippines, Sabah, and 

Indonesia. 

REMARKS-The distinguishing feature of this 

species is the distinct marginal and lamina1 10- 

bules (Figure 1 b). These lobules are dorsiventral, 

simple to forked, and sometimes subascending. Bulbate cilia are rarely developed on the margins in 

contrast, for example, to the abundant production 

of bulbils in R. luteoviridis. There is some intergradation with echinocarpic-acid-containing specimens of R. ampkitkrix, which have strongly lobulate isidia. On the average R. ampkitkrix would 

have much narrower lobes than R. sckirospatha 

and some trace of isidial initials. In addition R. 

ampkitkrix is rare at the higher elevations where 

R. sckizospatkn normally occurs. In addition, although R. sckizospatka is sterile, I suspect that the 

apothecia, when discovered, will be ecoronate. 

CHEMISTRY.-Medulla K- yellow, C- or C+ 

SPECIMENS EXAMIh.ED.-Philippines: Bukidnon, Sulit 14806 

(US). Malaya: Pahang, Hale 30089, 30142, 30210, 30216, 30221, 

30222, 30223, 30486, 30494. Sabah: Hale 28156, 28157, 28221, 

29026, 29037, 29117, 29285. Indonesia: Java, Groenhart 2219 

(L, US), Ooststroom 14198, 14300 (L, US). A record from 

Taiwan by Kurokawa (1965:267), which I have not been 

allowed to examine, is probably R. amphithrix. 

20. Relicina subabstrusa 

FIGURE 16b 

Relicina subabstrusa (Gyelnik) Hale, 1974:485. 

Parmelia subabstrusa Gyelnik, 1931:288 [based on P. abstrusa 

f. laevigata Lynge]. 

Parmelia limbata f. endococcinea Muller Argau, 1887:318 

[type-collection: Russell River, Australia, Sayer 19 (G, 

lectotype)]. 

Parmelia abstrusa f. laevigata Lynge, 1914: 147 [type-collection: 

Bocca da Serra, Serra da Chapada, Mato Grosso, Brazil, 

Malme (S, lectotype)]. 

Parmelia endococcinea (Muller Argau) Gyelnik, 1931:288. 

Parmelia decaryana Gyelnik, 1934: 153 [type-collection: TsantParmelia kilauae f. laevigata (Lynge) Gyelnik, 1935:37. 

sani, Anjouan, Comoro Island, Decary (BP, holotype)]. 
NUMBER 26 

FIGURE 16.-Species of Relicina: a, R. schizospatha (Neervoort 1062); b, R. subabstrusa (Hale 

25800); c, R. subconnivens (Hale 30054): d, R. sublanea (Hale 26650); e, R. siLblimbatn (Kimkawa 1872); f, R. sydneyensis (Weber 49048). (Scale in mm.) 

29 
30 SMITHSONJAN CONTRIBUTIONS TO BOTANY 

Thallus adnate on bark, 4-8 cm broad; lobes 

sublinear, 0.5-2 mm wide; bulbate cilia numerous, 

strongly inflated, to 1 mm long; upper surface 

plane, continuous; lower surface black, densely 

rhizinate, the rhizines black, simple (Figure 8c). 

Apothecia very common (frequency 83y0), adnate, 

1-3 mm in diameter, the exciple strongly coronate; 

spores 8, 4 X 7-8 pm. 

CHEiWIsTRY.-Medulla K-k yellow turning red, 

C-, KC-, P+ orange, norstictic and usnic acids 

with or without salazinic acid. 

HABITAT.-on trunk and branches of dipterocarps in Old World rain forest and in secondary 

forest at sea level to 850 m elevation and on trees 

at mid elevations in the New World. 

DIsTRIBUTION.-Taiwan, Philippines, Sarawak, 

Australia, Comoro Islands, Brazil, and Paraguay. 

REMARKS.-Relicina subabstrusa has the broadest geographic range of any species in the genus 

but is commonly collected only in the Philippines. 

As with R. abstrzisa, its presumptive isidiate morph, 

the Old Miorld population lacks or has only traces 

of salazinic acid, whereas salazinic is produced in 

heavy concentration in the New World. Muller’s 

specimen of f. endococcinea is red because of the 

decomposition of norstictic acid. 

SPECIMENS EXAhrINED.-Phi~ippines: Mountain, Hale 25760, 

25762, 25772, 25581, 25797, 25800, 25801, 25812, 25817, 25838, 

25858, 25863, 25879, 25898, 25901, 25802a, 25913, 26506; 

Cagayan, Hale 25608, 25654, 25661, 25751; Cavite, Hale 26811, 

26828; Polillo, Leiberg 532 (US); Negros Occidental, Hale 

26427, 26444, 26449, 26473, 26494, 26494a, 26510, 26519, 26564, 

26581, 26626, 26632, 26693, 26696; Agusan, Hale 24460, 24474, 

24984, 25003, 25014, 25029, 25033, 25048; Basilan, Hale 24935, 

24949, 25141, 25333. Sarawak: Hale 29991. Paraguay: Balansa 

(H). See Kurokawa (19633267) for records from Taiwan. 

21. Relicina subconniuens, new species 

FIGURE 16e 

Thallus adnatus, corticola, viridi-flavicans, circa 

10 cm latus, lobis sublinearibus, contiguis, sparse 

lobulatis, margine dense bulbociliatis, bulbis aetate 

inflatis, globosis, isidiis sorediisque destitutis; cortex superior 10-12 ym crassus, stratum gonidiale 

circa 12 pm crassum, medulla 60-80 ym crassa, 

cortex inferior 16-18 pm crassus (Figure 3f); subtus niger, dense rhizinosus, rhizines nigris, nitidis, 

simplicibus vel sparse ramosis. Apothecia adnata, 

0.5-1 .O mm diametro, amphithecio coronato, basin 

retrorso-rhizinato, disco carneo, imperforato, sporis 

octonis, 3 X 4 ym. 

CHEMISTRY.-~edUlla negative with color tests, 

usnic and caperatic acids. 

HOLOTYPE.-virgin dipterocarp forest, Ulu Gombak Forest Reserve, 15 km NE of Kuala Lumpur, 

elevation about 300 m, M. E. Hale 30054, 5 March 

REMARKS.-This species was first identified as 

R. connivens. It differs primarily in having coronate apothecia and very small spores and in containing caperatic acid rather than protolichesterinic acid. It is known only from the type collection. 

1965 (US). 

22. Relicin,a sublanea 

FIGURE 16d 

Relicina sublanea (Kurokawa) Hale, 1974:485. 

Parmelia sublanea Kurokawa in Hale and Kurokawa, 1964: 

146 [type-collection: Halmahera Island, Indonesia, Groenkardt 8409 (BO, holotype; US, isotype)]. 

Thallus adnate on bark, coriaceous, 6-10 cm in 

diameter; lobes sublinear-elongate, 1-2 mm wide; 

bulbate cilia strongly inflated; upper surface more 

or less convex, continuous; lower surface pale 

brown, densely rhizinate, the rhizines densely 

branched, and agglutinated (Figure 8g). Apothecia 

common (frequency 607L), adnate, 1-4.5 mm in 

diameter, ecoronate; spores 8, 4-5 X 7-9 ym. 

CHEMISTRY.-hfedUlki K-, C--, KC+ rose, P+ 

orange red, usnic and protocetraric acids. 

HABITAT.-on trunk and canopy branches of 

dipterocarps in rain forest at 300-850 m elevation. 

DIsTRIBuTIoN.-Southern Philippines, Malaya, 

and Indonesia. 

REMARKS.-This is another typical dipterocarp 

forest lichen characterized by pale agglutinated 

rhizines and protocetraric acid. Its relation to 

R. rnmosissimn is discussed under that species. The 

only other nonisidiate species with protocetraric 

acid, R. precirczimnodata, has coronate apothecia. 

SPECIMENS ExAMINED.-Philippines: Negros Occidental, 

Hale 26459, 26578, 26650, 26657; Surigao del Sur, Hale 24635, 

24655; Zarnboanga del Sur, Hale 24793, 23310; Basilan, 24920, 

24922. Malaya: Selangor, Hale 30280. 
NUMBER 26 31 

23. Relicina sublimbata 

FIGURE 16e 

Relicina sublimbata (Nylander) Hale, 1974:485. 

Purmeliu sublimbata Nylander, 1885:615 [type-collection: 

Yomah in Pegu, Burma, Brandis 595 (M, lectotype; H, 

isolectotype)]. 

Thallus adnate on bark or rock, coriaceous, 4-8 

cm in diameter; lobes sublinear-elongate, 1-2.5 mm 

wide; bulbate cilia coarse, inflated, to 1 mm long; 

upper surface plane, continuous; lower surface 

black, densely rhizinate, the rhizines black, simple 

to sparsely branched. Apothecia adnate, 1-2 mm 

in diameter, ecoronate; spores 8, 4-5 X 5-6 pm. 

CHEMISTRY.-Medulla K + reddish, c -, KC - , 

P + orange red, usnic acid, fumarprotocetraric acid, 

and succinprotocetrarir acid. 

HABITAT.-& tree trunks and rocks at mid elevation (1300-1400 m in Thailand). 

DIsTRIsuTroN.-Burma and Thailand. 

REMARKs.-Nylander (1 885) separated this species from R. limbata because of the smaller spores 

(4.5 X 6-8 pm) and whitish color. The two species 

are, in fact, totally unrelated, Relicina sublimbata 

is actually very close to R. malesiana, which produces fumarprotocetraric acid alone, has more 

branched rhizines and smaller spores, and does not 

occur on mainland Asia. Relicina sublimbata 

seems to be restricted to the Burma-Thailand region but little is known of its full range there. 

US). 

SPECIMENS ExAMINm.-Thailand: Kurokawa 1872 (TNS, 

24. Relicina sydneyensb 

FIGURE I6f 

Relicina sydneyensis (Gyelnik) Hale, 1974:485. 

Parmelia sydneyensis Gyelnik, 1938:292 [type-collection: Mossman’s Bay, Sydney, Australia, Cheel (BP, holotype)]. 

Parmelia limbata Laurer f. isidiosn Muller Argau, 1887:59 

[type-collection: Richmond River, Australia, Hodgkinson 

(G, lectotype)]. 

Purmelia tumescens Hale and Kurokawa, 1964:147 [based on 

P. limbata f. isidiosa Muller Argau]. 

Parmelia subturgida Kurokawa, 1965:268 [type-collection: 

Bandagamori, Prov. Tosa, Japan, Makino (TNS, holotype; 

US, isotype)]. 

Thallus adnate on bark or rocks, 3-8 cm in 

diameter; lobes sublinear to sublinear-elongate, 

contiguous, 0.8-1.5 mm wide; bulbate cilia tapered 

to strongly inflated; upper surface plane, faintly 

maculate, irregularly cracked with age, isidiate, the 

isidia simple, to 0.8 mm high; lower surface brown 

to dark brown, moderately rhizinate, the rhizines 

brown or black, mostly simple. Apothecia rare, 

adnate, 1.5-3.5 mm in diameter, ecoronate, the base 

retrorsely rhizinate; spores 8, 4-5 X 7-8 pm. 

CHEMISTRY.-Medulla K + yellow turning red, 

C-, KC-, P+ orange, usnic, norstictic, and 

stictic acids with or without atranorin. 

HABITAT.-on bark of trees and more rarely on 

rocks in open forests at low elevation. 

DIsrrcIsuTIoN.-Japan, Taiwan, Java, and Australia. 

KEMARKs.-Muller Argau recognized the close 

relation between this and R. limbata; R. sydneyensis would appear to be the isidiate morph of 

R. limbata. It has identical chemistry but seems 

to occur more frequently on trees and, of course, 

to have a broader geographic range. Chromatographic profiles of the types of Parmelia limbata f. 

isidiosa and P. subturgida are identical. Hale and 

Kurokawa (1 964) proposed the name Parmelia 

tumescens but later discovered that the type of 

P. sydneyensis Gyelnik is actually sparsely isidiate, 

not nonisidiate. 

SPECIMENS EXAMINED.-Japan: IZU, Kurokawa 58622 (Us). 

Indonesia: Java, Neruoort 5874 (BO). Australia: New South 

Wales, Weber and McVean L-49048 (COLO, US); Queensland, Du Rietz 4234:l (UPS, US). See Kurokawa (1965:268) 

for records from Taiwan. 

Nomen Inquirendum: Parmelia nigrociliata 

Parmelia nigrociliata Hillmann, 1940:39 [type-collection: 

Mutautu, Samoa, Vaupel (destroyed at B); (not Parmelia 

nigrociliata Bouly de Lesdain ex anno 1933)l. 

This species cannot be typified since the type 

(and only known) collection was lost at Berlin. 

The description and remarks by Hillmann, however, leave little doubt that it is synonymous with 

Relicina samoensis (Zahlbruckner) Hale. 
Literature Cited 

Asahina, Y. 

1934. Uber Parmelia abstrusa Vain., P. limbata Laur. und 

P. relicina Fr. aus Japan. The Journal of Japanese 

Botany, 10:299-304. 

1951. Lichenes Japoniae novae vel minus cognitae (3). 

The Journal of Japanese Botany, 26:193-198. 

On a Collection of Exotic Lichens Made in Eastern 

Asia by the Late Dr. A. C. Maingay. Journal of the 

Linnean Society, Botany, 20:48-69. 

Chemical and Morphological Evolution in Parmelia 

sect. Hypotrachyna: Product of Ancient Hybridization? Brittonia, 25:162-173. 

Crombie, J. M. 

1883. 

Culberson, C. F., and M. E. Hale, Jr. 

1973. 

Culberson, W. L. 

1973. 

Fries, E 

1825. 

Gyelnik, 

1931. 

1934. 

1935. 

1938. 

The Parmelia perforata Group: Niche Characteristics of Chemical Races, Speciation by Parallel 

Evolution, and a New Taxonomy. Bryologist, 76: 

20-29. 

Systema Orbis vegetabilis. Part 1, 374 pages. Lund. 

V. 

Additamenta ad cognitionem Parmeliarum, 11. 

Fedde Repertorium, 29:273-291. 

Additamenta ad cognitionem Parmeliarum, V. 

Fedde Repertorium, 36: 151-156. 

Revisio Typorum ab auctoribus variis descriptorum 

1. Annales Musei Nationalis Hungarici, 29: 1-54. 

Additamenta ab cognitionem Parmeliarum, 8. Annales Mycologici, 36:267-294. 

Hale, M. E. Jr. 

1965a. 

1965b3. 

1971. 

1973. 

1974. 

A Monograph of Parmelia Subgenus Amphigymnia. 

Contributions from the United States National Herbarium, 36: 193-358. 

Six New Species of Parrnelia from Southeast Asia. 

The Japanese Journal of Botany, 40:199-205. 

Parmelia afluens, a New Species of Lichen in Subgenus Amphigymnia with a Yellow Medulla. Phytologin, 22:141,142. 

Fine Structure of the Cortex in the Lichen Family 

Parmeliaceae Viewed with the Scanningelectron 

Microscope. Smithsonian Contributions to Botany, 

10: 1-92. 

Bulbothrix, Parmelina, Relicina, and Xanthoparme&, Four New Genera in the Parmeliaceae 

(Lichenes). Phytologia, 28:479-490. 

1975. A Revision of the Lichen Genus Hypotrachyna in 

Tropical America. Smithsonian Contributions to 

Botany, 25:l-73. 

Studies on Parmelia subgenus Parmelia. Contributions from the United States National Herbarium, 

Hale, M. E., and S. Kurokawa 

1964. 

36:121-191. 

Hillman, J. 

1940. 

Knight, C. 

1882. 

Neue oder wenig bekannte Flechten aus aller Welt, 

111. Fedde Repertorium, 48:34-40. 

Contribution to the Lichenographia of New South 

IVales. Transactions of the Linnean Society of London, Botany, new series, 2:37-51. 

Kurokawa, S. 

1965. Revision of Series Relicina of the Genus Parmelia 

in Japan and Taiwan. The Japanese Journal of 

Botany, 40: 264-269. 

Laurer, J. F. 

1827. Sieber’sche Lichenen. Linnaea, 238-46. 

Lynge, B. 

1914. Die Flechten der ersten Regnellschen Expedition. 

Die Gattungen Pseudoparmelia gen. nov. und Parmelia Ach. Arkiv for Botanik, 13 (13):1-172. 

Lichenologische Beitrage, XX. Flora, 65:316322. 

Lichenologische Beitrage, XXVI. Flora, 70:316-320. 

Lichenes in regionibus exoticis quibusdam vigentes, 

cum synopticis enumerationibus. Annales Sciences 

Naturelles Botanique, series 4, 11:205-264. 

1885. Parmeliae exoticae novae. Flora, 68:605-615. 

1886. Lichenes nonnulli Australiensis. Flora, 69:323-328. 

1907. Botanische und zoologische Ergebniss einer wissenschaftlichen Forschungsreise nach der Samoainseln, 

dem Neuguinea-Archipel und den Solomonsinseln, 

IV: Die Flechten von A. Zahlbruckner. Denkschrift 

Akademie der Wissenschaften in Wien, 81:250-292. 

Ptude sur la classification naturelle et la morphologie des Lichens du Bresil. Acta Societas Fauna et 

Flora Fennica, 7 (7):l-247. 

Systematisches Verzeichniss der im indischen Archipel in den Jahren 1842-1848 gesammetten sowie der 

aus Japan empfangenen Pflanzen. 160 pages. Zurich. 

Muller Argau, J. 

1882. 

1887. 

1859. 

Nylander, W. 

Rechinger, K. H. 

Vainio, E. A. 

1890. 

Zollinger, H. 

1854. 

* U.S. GOVERNMENT PRINTING OFFICE: 1975-58S-80/082S 

32