S M I T H S O N I A N  C O N T R I B U T I O N S  T O  B O T A N Y  N U M B E R  2 6  
A Monograph of the Lichen Genus 
Relicina (Parmeliaceae) 
Mason E. Hale, Jr .  
SMITHSONIAN INSTITUTION PRESS 
City of Washington 
1975 
A B S T R A C T  
Hale, Mason E., J r .  A Monograph of the Lichen Genus Relicina (Parmelia- 
ceae). Smithsonian Contribut ions 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 pro- 
ducing 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, fumarproto- 
cetraric 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. amphi thr ix ,  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. I t  is characterized by marginal bul- 
bate 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 treat- 
ment of the genus based on herbarium and field 
studies. 
T h e  first species of Relicina was described by 
E. Fries as Parmelia relicina on the basis of a speci- 
men from ?Rawak? in 1825. Neither Linnaeus nor 
Acharius had seen any material belonging to this 
genus. Laurer described the second species, P. lim- 
bata, an Australian endemic, in 1827. This  species 
is well known because i t  was issued in several 
widely distributed exsiccati. 
I n  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. lim- 
bata f .  isidiosa Muller Argau (=Relicinu sydney- 
ensis). 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 mar- 
ginally bulbate species formed a natural group, 
quite distinct from section Xanthoparmelia where 
they had previously been classified. They recog- 
nized 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 ex- 
ample, 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! An- 
other habitat where Relicinn has proved abundant 
is the oak forests found at high elevation through- 
out southeastern Asia. These areas have not been 
visited by lichenologists either. 
The  most reassuring aspect of my field work was 
1 
2 ShiITHSONIAN CONTRIBUTIONS T O  BOTANY 
that all of the species described by Hale and 
Kurokawa were rediscovered, often in great abun- 
dance 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. sub- 
turgida). 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 follow- 
ing 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. Jovet- 
Ast, Dr. R.  A. Maas Geesteranus, Dr. H. Merx- 
muller, Dr. H. Riedl, and Dr. R. Santesson. 
Dr. S. Kurokawa assisted with species descrip- 
tions, 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. Coopera- 
tive Science Program and by the Smithsonian Re- 
search Foundation. In  this connection I wish to 
express sincerest thanks to Dr. G. Velasquez and 
Dr. Flora Uyenco of the University of the Philip- 
pines, who arranged for trips to lumbering areas 
in the Philippines. I am also indebted to Mrs. 
Sheila Collonette for arranging a trip to Mt. Kina- 
balu, to Dr. W. Meijer for trips to eastern Sabah, 
and to Mr. J .  Anderson for assistance in Sarawak. 
Dr. Manuel Lopez Figueiras kindly provided sup- 
port and transportation in Venezuela. T h e  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. Herba- 
rium specimens were photopaplied by the Smith- 
sonian 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. amphi thr ix  (Figure la), R.  circum- 
nodata, R. planiuscula, and R. sydneyensis (Figure 
3i). The isidia are cylindrical and mostly un- 
branched but may become procumbent and lobu- 
late in R. amphi thr ix  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 l c )  are char- 
acteristic of some populations of R. amphi thr ix .  
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 pre- 
viously determined with the SEM that Relicina 
has a pored epicortex and a basic palisade plecten- 
chymatous upper cortex (Hale, 1972). More de- 
tailed 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 epi- 
cortex (Figure 4a,b) is about 0.6 pm thick as in 
other epicorticate genera (Hale, 1973). A particu- 
larly well-developed palisade structure, here called 
columnar, is characteristic of R.  flziorescens and 
R. planiusctila, two related species with a coria- 
ceous 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 5 b )  and R. subahstriisa (Figure 5 4 ,  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 T O  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  a n d  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. sublim- 
bata (Kurokawa 1872); i, R. sydneyensis (Kurokawa 58622). (Scale in lower right corner is 
10 km.) 
6 SMITHSONIAN CONTRIBUTIONS T O  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. sub- 
abstrusa (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 T O  BOTANY 
FIGURE &-Bulbate cilia of R. abstrusa (about X 20). 
MEDULLA.-The medulla occupies about two- 
thirds of the thallus and varies considerably in 
density. In  R. amphi thr ix ,  for example, a very 
small species, the hyphae are very loosely organized 
(Figure I f ) .  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 paraplectenchy- 
ma. It is up  to 20 pm thick in larger coriaceous 
species, such as R. planizisculn (Figure 5f )  but cor- 
respondingly thinner (10-16 pm) in less coriaceous 
species, such as P. luteoviridis (Figure 5e) or R. 
subabstmwa (Figure 5g). 
R ~ J L  RATE CIr.IA.-The marginal inflated bulbate 
cilia so characteristic of Relicina were probably 
first clearly recognized by Nylander when he de- 
scril)ed P0l'melifl cii c i imi iodnta 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 m a )  be lost 
or remain and even become densely branched. Ma- 
ture globose I,ulliae are actually 1101101v (Figure 
7 b )  and filled with a colorless fluid. The  heavily 
carlioni7ed ~valls are uniformlv thickened and 
c o r n p o d  of densely conglu tinatecl paraplectenc1i~- 
ma. A parallel-oriented procoplectench~niato~~~ 
structure, an orientation characteristic of cilia, can- 
not be distinguished at  the base but occurs far- 
ther 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 un- 
known. I t  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 sur- 
face is either black through carbonization or me- 
dium 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 (Fig- 
ure Sa), R. a m p h i t h r i x ,  R.  connivens, R .  echino- 
carpa,  R.  exinibyicnta, R. puorescens, R.  incongrua, 
R.  liitcoviyidk, R .  planiusci i la ,  R. relicinella, R .  
relicinitla, R. samoensis, R.  schizospatha, R.  sub-  
ab.rtursa (Figure 8r), and R.  subconnivens. Two 
species with a brown lower surface also have sim- 
ple rhizines: R .  l imbata (Figure 8 b )  and R. syd- 
neyensis. 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 8 4 ,  
R .  c i m i m n o d a t n ,  R .  pwcirrzimnodata (Figure 8e), 
R.  i.omo.sis.cinia (Figure S j ) ,  and R. .siiblanea (Fig- 
ure 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 out- 
side of Relirino. 
A ~ o ~ ~ ~ , ~ 1 ~ . - , 4 p o t l i e c i a  ar  very commonly pro- 
duced in Rclir inn.  Only two species, R.  Iiiteoviiidiq 
and R. schizo.spatha, lack them. Details on fre- 
quency are presented under the species descrip- 
tions. T h e  discs are generally almut 1-3 mm in 
diameter antl imperforate (see, for example, Fig- 
ures ISf ,  14c, 35b,  and 16b). The  rim i? smooth to 
crenate antl either coronate, t h a t  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 im- 
mersed 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 comple- 
ment of eight spores as in other Parmeliaceae. 
Relicilia ~ e l i c i n e l l a ,  however, produces multispored 
asci with 16-32 spores. T h e  spores are elliptical, 
ovoid, or nearly round, between 2-6 pm wide and 
3-10 pm long. T h e  one exception here is the un- 
FIGURE 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 T O  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 1 1  
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 
T h e  chemistry of Relicina was first summarized 
by Hale and Kurokawa (1964), who used micro- 
crystal tests and later some thin-layer chromatog- 
raphy (TLC). All of the specimens have been 
reexamined, as far as possible, with T L C  (Merck 
silica gel precoated plates) for this revision, using 
two solvent systems, hexane-ether-formic acid and 
benzene-dioxane-acetic acid. T h e  substances identi- 
fied in Relicina are listed below. Species with chem- 
ical strains are listed two or more times. 
Alectoronic acid: R. fluorescens. 
Atranorin: Sporadically produced in R.  acrobotrys, R. lim- 
Barbatic 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 .  @ores- 
4-0-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 T L C  pre- 
sents 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 proto- 
cetraric acid falling between them. Separation im- 
proves 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 oc- 
curs in high concentration. Species containing 
echinocarpic acid produce several additional spots, 
none of which have been identified. 
An unusual feature of Relicina is the predomi- 
nance of P+ depsidones and the unidentified echi- 
nocarpic acid, which appears to be P+ depside. 
Echinocarpic acid seems to be especially charac- 
teristic of Relicina, although it occurs rarely in 
Hypotr-achyna, usually in conjunction with bar- 
batic acid (Culberson and Hale, 1973), and in 
Parmotrema along with protocetraric acid (Hale, 
197 1). 
The  substances occur in several patterns. A com- 
mon one is acid A or acid A + B, as in R. acro- 
botryc (salazinic acid or salazinic acid with bar- 
batic 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. echino- 
carpa this involves salazinic and echinocarpic acids 
and with R.  flziorescens alectoronic and echino- 
carpic acid?. Other examples of variation are dis- 
cussed under the various species. I prefer not to 
treat these chemical populations as species because 
of the absence of any strong correlating morpho- 
logical 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 T O  BOTANY 
Qzie)ciis at high elevations. A few species grow in 
secondary forests on trees such as coconut or even 
on rocks. 
T h e  typical lowland dipterocarp forests occur in 
dense stands from sea level to less than 1000 m 
elevation. T h e  highest locality at which I collected 
on tlipterocarp was 850 m in Negros Occidental, 
Philippines. Those species which occur in  this ele- 
vational range (and often only up  to 400 m) 
include R. circiimnodata, R.  conniven y, R. male- 
tiana, R.  precircnmnodntn, R. rarnosissimn, R .  re- 
licitiiila, R.  wbabstrtisa, R. subconnivens, antl R. 
tiiblanea. In the New T\'orld one finds R. incon-  
grrra 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 dip- 
terocarp forests do not usually regenerate, Relicinn 
could become extinct as the forests are timbered 
throughout southeast Asia. 
Oak and ,  on Luzon, Philippines, P i n i i ~  make up 
the typical vegetation above about 1400 m eleva- 
tion on mountains in Southeast Asia. T h e  charac- 
acteristic species here are R. acrobotyrs, R. fliiores- 
f e n $ ,  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. l imbata,  and R. sydneyensis. 
Also occurring a t  low elevation but in disturbed 
forest5 in tropical regions are R. samoensis antl 
R. cxim bi icatn. Rel ic inn w b l i m  batn seems to occur 
in drier subtropical areas of Burma and Thailand 
a t  mid elevations. 
Two species, one restricted to Asia (R.  a m p k i -  
thrix) 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 
T h e  most distinctive feature of Relicinn is the 
high degree of endemism in Southeast Asia. Out- 
side of R. abstrzim and R.  wbabstrzisn, which occur 
in the Neotropics a5 well as in Asia, and three neo- 
tropical 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. 
T h e  genus obviously evolved in the ancient 
dipterocarp forests of Southeast Asia. T h e  endemic 
neotropical species may be remnants of older more 
widespread species, although they are unrelated to 
any species now found in the Old World. Alter- 
natively they may have evolved separately in the 
New 12rorltl where conditions have obviously not 
been favorable for rapid evolution and expansion 
of the genus. 
T h e  number of species known in various geo- 
graphic areas is summarized in Figure 10. T h e  
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. sub- 
nbstrirsn, and R .  sydneyensis. 
Philippines: R.  nhstrusn, R.  ninphi thrix ,  R .  circumnodntn, R. 
conniz~ens, R .  mnlesinnn, R.  plnnizisciiln, R. precircumno- 
dntn, R .  roniosissinzn, R .  reliciniiln, R .  snmoensis, R .  schizo- 
spothn, R. sitbnbstritso, and R.  sublnnen. 
hfalaya: R .  nbstrusn, R .  ninphi thrix ,  R .  circumnodntn, R. 
p lan  i i t sc~i ln ,  R .  rninosissiinn, R. relicin i i ln, R.  schi:ospathn, 
R .  s~cbconnivenr, and R. srihlnnen. 
Indonesia: R.  nbstritsn, R .  acrobotrys, R. connivens, R .  plani- 
usculo, 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 .  nmph i thr i x ,  R .  f luorexens,  R. luteo- 
?tiridis, R .  ~nnlrs innn,  R .  plirniirscitln, R .  rnmosissimn, and 
R.  schizospnthn. 
New Guinea: R.  connivens, R .  fluorescens, and R.  malesinnn. 
Australia: R .  l imbntn,  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. n b s t r t m  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. sub- 
Paraguay: R.  nbstritsn and R.  sribnbstrusn. 
Argentina: R.  nbstrusn. 
nbst rusn. 
T h e  commonest species of Relicina are as fol- 
lows lvith number of collections examined: R. sub- 
ob.ttrii,ro (52), R .  omphi thr ix  (43), R. circiimnodnta 
(40), R. plnrii?i.tcziln (37), R. reliciniila (31), R. 
nbstrir.tn (26), and R. schizospnthn (20). Rarities 
N U M B E R  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. T h e  structure of 
the most characteristic acid, echinocarpic acid, is 
as yet unknown. T h e  following discussions, there- 
fore, will cover only the more obvious lines of 
evolution. 
MORPHOLOGICAL EVOLUTION 
F o m r A T I o N  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 b y  any of the species. Isidia do occur, how- 
ever, in five species, and four of these can be 
paired witli nonisidiate sexual morphs: 
R.  0 bsf YlLsn-R. S l i  110 b s t r u ~ n  
R. r i m i  i n  ii odatn-R . precircu i n  i iodnto 
R. ~~la?r iz~scz i ln-R.  fliiorcsrciis (echinocarpic acid population) 
R.  sydiieyensis-R. limbata 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
The  acid deficient population of R. amphi thr ix  
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 chem- 
ically 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. precircumno- 
data 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 charac- 
ters (Figure 11). 
CORONATE APOTHECIA.-I am assuming that ecor- 
onate species are more primitive than coronate 
species and gave rise by mutation to coronate 
species. T h i s  seems fairly evident in the R. sub- 
lanea-R. c i m i m n o d n t a  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.  subconni-  
vens (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 ecoro- 
nate parent populations are extant. Conversely no 
coronate species have evolved from the R. l imbata 
group. 
CHEMICAL EVOLUTION 
Chemical evolution is presumed to involve 
genetic processes that lead to differentiation of a 
parent species into two or more chemical popula- 
tions. 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 speci- 
mens 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 Bra- 
zilian 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, espe- 
cially 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). T h e  three populations occur together in sig- 
nificant numbers in Sabah. T h e  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 con- 
tained 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 repro- 
ductively isolated is R. eximbricata and R. relici- 
nella in the New World. While these externally 
similar species share the production of fumarproto- 
cetraric acid, R.  eximbricata also produces succin- 
protocetraric acid, while R.  relicinella contains an 
unidentified P + compound, also apparently re- 
lated to protocetraric acid. More significantly, R. 
relicinella has multispored asci. 
T h e  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. 
P r o g e n i t o r  
R. fruoresrens R. frtdoresrens 
[echinocarpic acid] [alectoronic acid] 
R. planiiisrrda R. fruorescens 
[echinocarpic & alectoronic acids] 
FIGURE 12.-Hypothetical  e v o l u t i o n  of c h e m i c a l  p o p u l a t i o n s  
in R .  fluorexens and R .  planiuscula. 
A common situation in Relicina is the differen- 
tiation of species into parallel chemical groups 
with various combinations of fumarprotocetraric 
acid and other substances. For example R. subl im-  
bata and R. nzalesiana are essentially identical but 
R. subl imbata produces succinprotocetraric acid 
along with fumarprotocetraric acid, while R.  male- 
siana produces only fumarprotocetraric acid. In  
this instance the species have a different geographic 
range, R.  subl imbata 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 Parme- 
liaceae because of its morphology and phytogeog- 
raphy. First, i t  has nothing to do with X a n t h o -  
parmelia,  with which it shares only the yellow 
cortical pigment usnic acid. The  only closely 
related genus is B u l b o t h r i x  (Hale, 1974), which 
also has marginal bulbate cilia and coronate apo- 
thecia but which produces only atranorin in the 
cortex. Bulbothr ix ,  however, is rare in the range 
of Relicina.  Only B .  subdissecta (Nylander) Hale 
and B .  pigmentarea (Hale) Hale occur in the low- 
land rain forests of Southeast Asia and B .  Jubin- 
flata (Hale) Hale in oak forests where one finds 
Relicina.  Bulbothr ix  is largely a genus of the drier 
subtropical zones, being especially common in 
southeastern United States and Africa. I t  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 
B u l b o t h r i x  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 Bulbothr ix ,  for exam- 
ple, have gyrophoric or salazinic acid, both of these 
very rare in Relicina.  No barbatic, difiactaic, echi- 
nocarpic, furmaprotocetraric, or stictic acids are 
produced. On the other hand, Bulbothr ix  produces 
lecanoric, lobaric, and colensoic acids, all unknown 
in Relicina.  I would argue then that Bulbothr ix  
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 acid- 
atranorin 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. p lan iusda  
3. R. amphithrix 7. Lobes narrow and appressed, 0.5-1.0 mm wide. 
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 P f  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 P f  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 Reli-  
cinn. Furthermore, my typification of Pnrmelia 
eumorphn Hepp, based on an incorrectly deter- 
mined 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 Cocco- 
cnl-pin, although the same number in L is a 
Relicinn species. Finally, Hepp?s protologue ob- 
viouslj refers to a nonparmelioid genus. 
T h e  species are listed alphabetically in the list- 
ing below. All collections by Hale are deposited 
in the National Museum of Natural History, 
Smithsonian Institution, and the herbarium acro- 
nym (US) is not given. 
1. Relicina abstrusa 
FIGURE 13a 
Relicinn nbstrusu (Vainio) Hale, 1974:484. 
Parnzelin nbsti m a  Vainio, 1890:G4 [ripe-collection: Caraca, 
Minas Gerais, Brazil, 1/ni?iio 1347 (not 1346) ( T U R ,  lecto- 
type; UPS, isolectotype)]. 
Thallus closely adnate on bark or rocks, 6-10 
cm in diameter, pale chartreuse yellow; lobes sub- 
linear, 1-2 mm wide; upper surface plane to con- 
vex, continuous or faintly maculate, moderately 
isidiate, isidia short, mostly simple: bulbate cilia 
moderately inflated; lower surface black and rliizi- 
nate, the rhi7ines simple, shiny (Figure 80). Apo- 
thecia 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 K f  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 par- 
ent, the S e w  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.  n b s t i x ~ 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 popu- 
lations (5Sm, frequency) than in the Old Ii?orld 
(7yo frequency). 
One specimen from Brazil (Lichenes nirsfro- 
nmeiicnni  95) contained diffractaic acid and a trace 
of barbatic acid in addition to the main compo- 
nents 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 nor- 
stictic and stictic acids in Japanese specimens, 
whicli lie identified as P. nb.r t~ i r . rn  and these too 
represent Rclirinn ,sydneyei?sis. 
S F E C I S I ~ ~ S  Exh\rIxrD.--Florida: Baker Coniltv. HI!/<? 21968. 
Xlexico: Chiapas. H n l f  20028 ( C .  ITS), 20120, 20898. 2014,;. 
Panama: Panama, Hnlr  88116. 38447. Jnniaic:i: Ci/lber.ioti 
13748 (DKTKE), P l i f t  (1-S). Coloml,ia: r.iiidig 701 ( R J I .  P). 
l?enezuela: Federal, Ol~rrr i ? i~ ik l r r  nnrf Poclt (\I .  t . S ) :  M e r i d a ,
Hale 42070, 48886; 3Ierindn. I~cii.csclii 289ic (t?q). R ~ i ~ i l :  Rio 
de  Jatieiro. Ei t r t i  6541, 7175R.  7671 (1?s). G l n t i o i c  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 B M ) ;  e, R .  conniuens (Hoog- 
land 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). Argen- 
tina: 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: Groen- 
hart 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 (fre- 
quency 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. 
I t  is also the only species in Relicina with salazinic 
acid as the main chemical component. While the 
type contains only salazinic acid, specimens col- 
lected 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 isi- 
diatis, isidiis cylindricis, cylindricis vel sparse ra- 
mosis, 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 retror- 
so-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 un- 
known 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 al- 
though 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 wl ic in  ! / l a ,  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. i t  
seems probable that the acid deficient population 
of R. amphithrix is the isidiate morph of R.  
relicinzila. 
SrEcrartxs E X ~ V I N F D  (echinocnipic ncid present, isidinte or 
becoming lobalnte-isidinte).--Philippiues: AIouiitnin, H o l e  
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 palmate- 
lobulate).-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 o r  
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. 
S P E C I X ~ E N S  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  i n  Crombie, 1883:51 [type- 
collection: Government Hill,  Penang, Malaya, Maingay 22 
(BM, lectotype; FH, H, isolectotypes)]. 
Tliallus closely adnate on bark, 3-8 cm in diam- 
eter; lobes sublinear-elongate, 0.7-2 mm wide; buI- 
bate cilia dense and conspicuous, strongly inflated 
and becoming globose and apically densely 
Ixanclied; upper surface plane to convex, continu- 
ous 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 (fre- 
quency lo%), adnate, 1-2 mm in diameter, rim 
coronate; spores 8, cornute, 3 X 10-12 pm. 
rose, P f  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  Ny- 
lander from sterile material. Apothecia are now 
known from 4 of 39 specimens that I have been 
able to examine. All are coronate and have un- 
usual bicornute spores, exactly the same as those 
in the presumptive parent species R. precirciirnno- 
data. 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 KCS- 
Relicirm ro~iuivens (Kurokarva) Hale, 1974:484. 
Partrielin rotinivens Kurokawa irz Hale a n d  Kurokawa, 1964: 
142 [type collection: Milne nay district, Papua,  S e w  
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 wide- 
spread hut not common in a band from the Philip- 
pines through Xew Guinea to the Solomon Islands, 
occurring generally at 300-800 m elevation. No 
other species in the genus produces protolichesteri- 
nic acid. It pro1)ably would be mistaken for R .  
m ( ~ l e . ~ i n ~ i n  (P+ red) without appropriate chemical 
testing. Closely related R.  szibconniuens has coro- 
nate apothecia but if collected sterile it could be 
differentiated b y  the presence of caperatic acid. 
CHEMlSTRY.-;\/IedUl~a K-, C-, KC-, P--, pro- 
SPECI\IENS ExAzrisrD.-Pliilippines: Segroes Occidental, 
Hale 26457, 26,709, 26626a: Agusan, Hale, 25025, 25034, 25044. 
Colomon Islands: Guadalcanal, Hill 9057 (EM, US). h d d i -  
N U M B E R  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 saxi- 
colous, 2-6 cm in diameter; lobes sublinear- 
elongate, 1-2 mm wide; bulbate cilia gradually in- 
flated; upper surface plane, faintly maculate; lower 
surface black, densely rhizinate, the rhizines simple 
or sparsely branched. Apothecia numerous (fre- 
quency 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-, P f  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, close- 
ly 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, i t  is not related to the 
tropical members of the genus containing this acid. 
For example, R.  .mmoensis has more globose bul- 
h e  and coronate apothecia. 
T h e  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 ap- 
parently 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 (cash- 
ew, Eugenia) in open areas from sea level to 600 m 
elevation. 
D ! s ~ ~ ~ ~ u T r o ~ . - F l o r i d a  and the West Indies. 
REMARKs.-Relicina eximbricata has been col- 
lected for the most part in disturbed secondary 
forest. It is rather rare if the few collections avail- 
able reflect i ts  true abundance. I was unable, for 
example, to recollect any specimens at the locality 
in Florida visited by Swanson. I t  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 rhizi- 
nate, 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 @ M C U S  
and other trees in open forest at 1600 to 2400 m 
elevation. 
DIsTRiBuTIoN.-sabah and New Guinea. 
REMARKs.-Relicina fluorescens was first recog- 
nized 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. I t  is 
brownish when developed with sulfuric acid. a- 
Collatolic acid does not occur in  this species. 
T h e  isidiate morph of R. fluorescens is R. plan- 
iuscula, which has a broader geographic range. 
SPECIMENS EXAMINED (alectoronic acid present).-New 
Guinea. Western Highland: Kurokawa 75 in Lichenes Rari- 
ores et Critici  Exsiccati (US). See Hale (1965: 203) for addi- 
tional 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, viridi- 
flavicans, 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 goni- 
diale 10-12 pm crassum, medulla 70-80 pm crassa, 
cortex inferior 10-12 pm crassis (Figure 2c); sub- 
tus niger, modice vel dense rhizinosus, rhizinis 
nigris, nitidis, simplicibus vel sparse ramosis. Apo- 
thecia 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-demethyl- 
barbatic 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 identi- 
cal 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 col- 
lected 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: Syd- 
ney, Australia, Knight (H, lectotype) (not P a r m e h  
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 sub- 
linear-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 (fre- 
quency 100~o) ,  adnate, 1-6 inm in diameter, ecoro- 
nate, 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 col- 
lected 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 sphaero- 
spora. Knight then sent a specimen to Nylander, 
who immediately published a new name, P. insin- 
uata, since Knight?s epithet was a homonym of 
Nylander?s own P. sphaeospora published in 1859. 
Vainio (1890) comprehended the species correct- 
ly, 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 Krypto- 
gamae 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); Tas- 
mania, 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 diam- 
eter; lobes sublinear-elongate, 1.5-3 mm wide; 
upper surface plane, faintly maculate, lobulate, 
the lobules large, dorsiventral, marginally bulbate- 
ciliate, 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 recog- 
nized by the large lobules and the unique chem- 
istry. I t  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 numer- 
ous (frequency 80y0), adnate, ecoronate; spores 8, 
2-4 X 3-7 pm, medulla below the hymenium rare- 
ly orange red. 
CHEMISTRY.-Medulla K--, c - ,  KC-, P-k red, 
usnic and fumarprotocetraric acids and if pig- 
mented an unidentified K + purple anthraquinone. 
HABITAT.-On trunk and branches of diptero- 
carps 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 some- 
what 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 fumarprotocetra- 
ric 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 lobu- 
late; 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 a t  1500-2200 m (at lower elevation in Japan 
and Taiwan). 
DIsTRlnUTI0N.-Japan, Taiwan, Philippines, Ma- 
laya, Indonesia, and Sabah. 
REMARKs.-This is the commonest species of 
Reliciria at higher elevations in Southeast Asia. 
T h e  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. T h e  population 
on Pin ?is in the northern Philippines is especially 
strongly isidiate-lobulate, and I had at  first con- 
sidered i t  to be a separate species. 
SPECIMENS ExAhrmm.-Japan. Hyuga: Hale 29665. Philip- 
pines: 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. In- 
donesia: Java, Groenhart 2217 (L); Sumatra, Steenis 3895 
P O ) .  
14. Relicina precircumnodata, new species 
FIGURE 15b 
Thallus adnatus, corticola, viridi-flavicans, 4-10 
cm diametro, lobis sublinearibus, contiguis vel sep- 
aratis, 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 in- 
ferior circa 12 pm crassus (Figure 2h) ;  subtus pal- 
lide 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, proto- 
cetraric 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 presump- 
tive nonisidiate morph of R. circumnodata. Both 
contain protocetraric acid and have unusual bi- 
cornute 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 diam- 
eter; lobes sublinear-elongate, 0.7-2 mm wide; bul- 
bate cilia gradually inflated to gobose, becoming 
apically branched; upper surface plane, maculate; 
lower surface pale to dark brown, moderately rhizi- 
nate, 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 resem- 
bles R. sublanea, which differs in producing pro- 
tocetraric 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: San- 
tarem, Brazil, Spruce 136 (H, lectotype; BM, G ,  P, isolecto- 
types)]. 
Thallus closely adnate on bark, 3-6 cm in diam- 
eter; lobes sublinear, 0.5-1.5 mm wide; bulbate 
cilia distinct, short; upper surface plane, continu- 
ous; 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-, P f  red, 
usnic acid, fumarprotocetraric acid, and an un- 
identified 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 macu- 
late, without lobules or becoming marginally lobu- 
late, 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 Indo- 
nesia. 
REMARKs.-The characteristic features of this 
widespread species are the narrow appressed lobes 
and lack of lichen substances in the medulla. I t  is 
an extremely variable species, however, particularly 
in regard to production of marginal lobes. Some 
specimens have no obvious lobules. Muller?s type- 
specimen is somewhat lobulate. He described Par- 
melia 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. I ts  chemistry is still undecided since, al- 
though it is negative with color tests, a dull, high 
unidentified spot appears on T L C  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 ad- 
nate, 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 echino- 
carpic 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 C a b  
donia, Samoa, and Society Islands. 
REMARKs.-Relicina samoensis is a lowland spe- 
cies occurring in secondary forests. T h e  lobes vary 
considerably in size but are usually narrow and 
closely adnate. This is the only species with echino- 
carpic 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 sur- 
face, the lobules dorsiventral, procumbent to sub- 
ascending, 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 echino- 
carpic acids and a C +  unknown. 
HABITAT.-on trunk and branches of Q U e Y C U S  
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. Bul- 
bate cilia are rarely developed on the margins in 
contrast, for example, to the abundant production 
of bulbils in R. luteoviridis. There is some inter- 
gradation with echinocarpic-acid-containing speci- 
mens of R .  ampkitkrix, which have strongly lobu- 
late 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, al- 
though 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: Tsant- 
Parmelia 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 ( K i m -  
kawa 1872); f, R. sydneyensis (Weber 49048). (Scale in mm.) 
29 
30 SMITHSONJAN CONTRIBUTIONS T O  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 diptero- 
carps 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 broad- 
est 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; cor- 
tex superior 10-12 ym crassus, stratum gonidiale 
circa 12 pm crassum, medulla 60-80 ym crassa, 
cortex inferior 16-18 pm crassus (Figure 3f); sub- 
tus 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 Gom- 
bak 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 coro- 
nate apothecia and very small spores and in con- 
taining caperatic acid rather than protolichesteri- 
nic 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, Groen- 
kardt 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 ele- 
vation (1300-1400 m in Thailand). 
DIsTRIsuTroN.-Burma and Thailand. 
REMARKs.-Nylander (1 885) separated this spe- 
cies from R. l imbata because of the smaller spores 
(4.5 X 6-8 pm) and whitish color. T h e  two species 
are, in fact, totally unrelated, Relicina subl imbata 
is actually very close to R. malesiana, which pro- 
duces fumarprotocetraric acid alone, has more 
branched rhizines and smaller spores, and does not 
occur on mainland Asia. Relicina subl imbata 
seems to be restricted to the Burma-Thailand re- 
gion 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: Moss- 
man?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 Aus- 
tralia. 
KEMARKs.-Muller Argau recognized the close 
relation between this and R. limbata; R. sydney- 
ensis 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. Chromato- 
graphic 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, W e b e r  and McVean L-49048 (COLO, US); Queens- 
land, D u  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, how- 
ever, leave little doubt that it is synonymous with 
Relicina samoensis (Zahlbruckner) Hale. 
Literature Cited 
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1934. Uber Parmelia abstrusa Vain., P. limbata Laur. und 
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1883. 
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1973. 
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Fine Structure of the Cortex in the Lichen Family 
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Bulbothrix, Parmelina, Relicina, and Xanthopar- 
me&, 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 
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Studies on Parmelia subgenus Parmelia. Contribu- 
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Hale, M. E., and S. Kurokawa 
1964. 
36:121-191. 
Hillman, J. 
1940. 
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Neue oder wenig bekannte Flechten aus aller Welt, 
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