SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 26

A Monograph of the Lichen Genus Relicina (Parmeliaceae)

Mason E. Hale, Jr.

SMITHSONIAN INSTITUTION PRESS City of Washington 1975 ABSTRACT

Hale, Mason E., Jr. A Monograph of the Lichen Genus Relicina (Parmelia- ceae). Smithsonian Contributions to Botany, number 26, 32 pages, 16 figures, 1975.-A revision on the world level is made for Relicina, a generic segregate of Parmelia characterized by having bulbate cilia on the lobe margins and by 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. amphithrix, R. incongrua, R. precircumnodata, and R. subconnivens, are described and one new combination, R. relicinula (Muller Argau) Hale, is made.

OFFICIALPUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution’s annual report, Smithsonian Year. SI PRESSNUMBER 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, no. 26. QKl.S2747 no. 26 [QK585.P2] 581’.08s [589’.1] 75-6191 19 Contents

Page Introduction ...... 1 Morphological Characters ...... 2 Chemistry ...... 11 Ecology and Habitats ...... 11 Phytogeography ...... 12 Evolution and Speciation ...... 13 Position of Relicina in the Parmeliaceae ...... 15 Key to Species of Relicina ...... 16 Species Treatment ...... 17 Literature Cited ...... 32

iii

A Monograph of the Lichen Genus Relicina (Parmeliaceae)

Mason E. Hale, Jr.

Introduction series of collections from the Bogor herbarium and described seven more species, mainly from Java Relicina is a generic segregate of the collective and Borneo (Kalimantan and Sabah). Most of genus Parmelia. It is characterized by marginal bul- these, as with previous species, were known only bate cilia and the constant occurrence of usnic acid, from type collections. It became obvious during a yellow pigment, in the cortex. The genus includes the course of their work, however, that these mar- 24 species, almost all of them occurring in tropical ginally bulbate species formed a natural group, Southeast Asia. This paper is a monographic treat- quite distinct from section Xanthoparmelia where ment of the genus based on herbarium and field they had previously been classified. They recog- studies. nized the 18 species then known as a series in The first species of Relicina was described by subgenus Paymelia section Imbricaria subsection E. Fries as Parmelia relicina on the basis of a speci- B icorn u tae. men from “Rawak” in 1825. Neither Linnaeus nor M7hile this publication (Hale and Kurokawa, Acharius had seen any material belonging to this 1964) was actually in press, I had an opportunity genus. Laurer described the second species, P. lim- to collect lichens in Malaysia, the Philippines, and bata, an Australian endemic, in 1827. This species Japan. What I discovered explained why this is well known because it was issued in several peculiar assemblage of species had been so poorly widely distributed exsiccati. collected in the past and represented by so few In the remainder of the 19th century only five herbarium specimens. Relicina inhabits canopy more taxa were discovered: Parmelia abstrusa branches in the vast lowland dipterocarp forests Vainio, P. circzrmnodata Nylander, P. relicinella from Burma to the Solomon Islands. I collected at Nylander, P. relicinula Muller Argau, and P. lim- logging heads where these giant trees, 60 m or bata f. isidiosa Muller Argau (=Relicinu sydney- more high, were being felled commercially. Most ensis). Between 1900 and 1964 an additional four earlier collectors had obviously not been able to taxa were described: P. abstrzisa f. laevigata Lynge find (or did not try to find) felled trees. For ex- (= Relicina subabstrusa), P. nigrociliata Hillmann, ample, many lichens were collected by botanists in P. samoensis Zahlbruckner, and P. samoensis var. the Philippines and studied by Vainio for his eximbvicata Gyelnik (=Relicina eximbricata). extensive Philippine lichen flora, yet he did not Hillmann (1940) recognized a natural relationship report a single species of Relicina, in reality the between six species described from the Old World commonest foliose lichens in the Philippines! An- and constructed a key to them. other habitat where Relicinn has proved abundant In I964 Hale and Kurokawa (1964) examined a is the oak forests found at high elevation through-

Mason E. Hale, Jr., Department of Botany, h’ational Museum out southeastern Asia. These areas have not been of Natural History, Smithsonian Institution, Washington, visited by lichenologists either. D.C. 20560. The most reassuring aspect of my field work was 1 2 ShiITHSONIAN CONTRIBUTIONS TO BOTANY

that all of the species described by Hale and extremely uniform in lobation. The lobes are Kurokawa were rediscovered, often in great abun- generally narrow and sublinear, dichotomously dance and comprising the major foliose element in branched, and contiguous. Small marginal lobules lowland rain forest. As one might expect, more often develop. undescribed species were found. I described two VEGETATIVE PROPAGULES.-one Of the mOSt un- species, Parmelia puorescens and P. malesiana usual features of Relicina is that no soredia are (Hale, 1965), and Kurokawa (1965) described other produced by any of the species. Soredia are usually species from Japan (P. echinocarpa and P. sub- the most common diaspore in the Parmeliaceae. turgida). In the final revision of this monograph Isidia, however, are produced by five species, four more new species are being described to bring R. abstrusn, R. amphithrix (Figure la), R. circum- the total for the genus to 24 species. nodata, R. planiuscula, and R. sydneyensis (Figure ACKNOWLEDGMENTS-Iwish to thank the follow- 3i). The isidia are cylindrical and mostly un- ing curators of museums who kindly sent specimens branched but may become procumbent and lobu- on loan: Dr. S. Ahlner, Dr. T. Ahti, Dr. R. Alava, late in R. amphithrix and to a lesser extent in Dr. 0. Almborn, Dr. C. E. Bonner, Mme. Jovet- R. planiuscula. True lobules that do not originate Ast, Dr. R. A. Maas Geesteranus, Dr. H. Merx- from isidia are produced by R. luteoviridis (Figure muller, Dr. H. Riedl, and Dr. R. Santesson. 14e) and R. schizospatha (Figure Ib). Appressed Dr. S. Kurokawa assisted with species descrip- palmately branched lobules (Figure lc) are char- tions, identification, and chemical studies both in acteristic of some populations of R. amphithrix. 1960 in Washington and in 1964-1965 when I EPICORTEXAND CoRTEX.-The internal structure studied at the Tokyo National Science Museum. of Relicina was studied by making longitudinal The field studies were supported by the National sections of the lobes and examining them under a Science Foundation under the Japan-US. Coopera- scanning-electron microscope (SEM). All of the tive Science Program and by the Smithsonian Re- species are illustrated in Figures ld-3i. I had pre- search Foundation. In this connection I wish to viously determined with the SEM that Relicina express sincerest thanks to Dr. G. Velasquez and has a pored epicortex and a basic palisade plecten- Dr. Flora Uyenco of the University of the Philip- chymatous upper cortex (Hale, 1972). More de- pines, who arranged for trips to lumbering areas tailed investigations of internal structure for the in the Philippines. I am also indebted to Mrs. purposes of this monograph have confirmed this Sheila Collonette for arranging a trip to Mt. Kina- structure. Most species have an irregularly arranged balu, to Dr. W. Meijer for trips to eastern Sabah, palisade layer 10-1 5 pm thick, rarely protruding and to Mr. J. Anderson for assistance in Sarawak. downward to 20 pm (Figure 4c-h). The pored epi- Dr. Manuel Lopez Figueiras kindly provided sup- cortex (Figure 4a,b) is about 0.6 pm thick as in port and transportation in Venezuela. The project other epicorticate genera (Hale, 1973). A particu- would never have been completed without the larly well-developed palisade structure, here called generous logistic support of lumber companies and columnar, is characteristic of R. flziorescens and forestry departments in the Philippines, Malaya, R. planiusctila, two related species with a coria- Sarawak, and Sabah. ceous thallus. The upper cortex consists of packed The scanning-electron microphotographs were columnar cells 20-25 pm high overlain by the taken by Mr. Walter Brown of the Smithsonian epicortex (Figure 5a,c). A similarly well-developed Scanning-electron Microscope Laboratory. Herba- columnar structure is known for R. Zziteovirdir rium specimens were photopaplied by the Smith- (Figure 5b) and R. subahstriisa (Figure 54, both sonian Photo

FIGURE1.-Morphology of Relicina: a: isidia of R. arnphithrix (Hale 30066) (about X 10); b, lobules of R. schizospatha (Groenhart 2219) (about X 10); c, lobules of R. arnphithrix (Hale 30287) (about X 10); d, R. abstrusa (Oberruinkler and Poelt); e, R. acrobotrys (A’eervoort 60 pp.); f, R. arnflhithrix (Hale 30051); g, R. circurnnodata (Hale 26580); h, R. connivens (Hoog land 4726); i, R. echinocarpa (Hale 29659). (Scale in lower right corner of d-i is 10 km.) 4 ShlITHSONIAN CONTRIBUTIONS TO BOTANY

FICWRE2.--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

FIGURE3.-Morphology of Relicina: a, R. relicinella (Nee and Mori 3766); b, R. relicinula (Hale 24402); c, R. sarnoensis (Hale 26955); d, R. schizospatha (Groenhart 2219); e, R. subabstrusa (Malrne sm.); f, R. subconnivens (Hale 30054); g, R. sublanea (Groeizhart 8409); h, R. sublim- bata (Kurokawa 1872); i, R. sydneyensis (Kurokawa 58622). (Scale in lower right corner is 10 km.) 6 SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE4.--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

FIGURE5.-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 TO BOTANY

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. It is altogether possible, however, that they act as propagules. On rare occasions one can see lobules or even submacroscopic thalli that seem to have originated from detached bulbae. Since they do not contain algae, they would have to function as giant fungal diaspores and encounter suitable symbiotic algae before continuing growth. LOWERSURFACE 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 FIGURE&-Bulbate cilia of R. abstrusa (about X 20). sparsely branched rhizines often occur in species with a black lower surface, as in R. abstrzisa (Fig- ure Sa), R. amphithrix, R. connivens, R. echino- MEDULLA.-The medulla occupies about two- carpa, R. exinibyicnta, R. puorescens, R. incongrua, thirds of the thallus and varies considerably in R. liitcoviyidk, R. planiusciila, R. relicinella, R. density. In R. amphithrix, for example, a very relicinitla, R. samoensis, R. schizospatha, R. sub- small species, the hyphae are very loosely organized ab.rtursa (Figure 8r), and R. subconnivens. Two (Figure If). Most species, however, have a densely species with a brown lower surface also have sim- packed medulla (see in particular Figures lg and ple rhizines: R. limbata (Figure 8b) and R. syd- 2h). Crystals of lichen substances encrust the neyensis. Both R. malesiana and R. siiblimbata, hyphae, especially in the upper half of the medulla as well as some of the other lowland species, may (Figure 3a,f). have rather densely branched, almost squarrose LOWERCORTFX.-The lower cortex in all species rhizines, I,ut these are always black and shiny consists of dense pacliydermatous paraplectenchy- without agglutination. Other species with a pale ma. It is up to 20 pm thick in larger coriaceous loiver surface, including R. arrobotl-ys (Figure 84, species, such as R. planizisculn (Figure 5f) but cor- R. cimimnodatn, R. pwcirrzimnodata (Figure 8e), respondingly thinner (10-16 pm) in less coriaceous R. i.omo.sis.cinia (Figure Sj), and R. .siiblanea (Fig- species, such as P. luteoviridis (Figure 5e) or R. ure 8g), have densely branched, bushy, and in part subabstmwa (Figure 5g). agglutinated rhizines that form a rvoolly mat about R~JLRATE CIr.IA.-The marginal inflated bulbate 1 mm thick. The rhizines differ from squarrose cilia so characteristic of Relicina were probably types, such as one finds in Pamielia szi2cnta Taylor, first clearly recognized by Nylander when he de- in the pale color, the blunt rather than pointed scril)ed P0l'melifl cii ciimiiodnta in 1883. These tips, and the adherence of the various ramified structures are produced along lobe margins with parts. I have not seen this Iiranching pattern out- great regularity (Figure 6). They begin on new side of Relirino. lobe5 as ordinary cilia but soon become basally A~o~~~,~1~.-,4potlieciaare very commonly pro- inflated (Figure 70). The ciliate tip ma) be lost duced in Rclirinn. Only two species, R. Iiiteoviiidiq or remain and even become densely branched. Ma- and R. schizo.spatha, lack them. Details on fre- ture globose I,ulliae are actually 1101101v (Figure quency are presented under the species descrip- 7b) and filled with a colorless fluid. The heavily tions. The discs are generally almut 1-3 mm in carlioni7ed ~valls are uniformlv thickened and diameter antl imperforate (see, for example, Fig- cornpod of densely conglu tinatecl paraplectenc1i~- ures ISf, 14c, 35b, and 16b). The rim i? smooth to ma. A parallel-oriented procoplectench~niato~~~crenate antl either coronate, that is, provided with structure, an orientation characteristic of cilia, can- black 1xill)ae (Figure 9) or ecoronate. TVhile these NUhlBER 26 9 bulbae appear to be the same as bulbate cilia, they apothecia (Figure 9). These were first described by are in fact erect pycnidia (Figure 7c,d) with an Laurer (1827) for Relicina limbata. apical pore, as most earlier workers seem to have SPORES.-Tlle asci contain the normal comple- concluded. The thallus also has the normal im- ment of eight spores as in other Parmeliaceae. mersed pycnidia that one would expect to find in a Relicilia ~elicinella,however, produces multispored parmelioid genus. asci with 16-32 spores. The spores are elliptical, Bulbate cilia (retrorse rhizines) often develop ovoid, or nearly round, between 2-6 pm wide and around the base of both coronate and ecoronate 3-10 pm long. The one exception here is the un-

FIGURE7.--Structure of bulbate cilia and amphithecial pycnidia: a, bulbate cilia on lobe margins of R. sublaiiea (Hale 30280) sectioned longitudinally (X 100); b, longitudinal section of the base of a bulbate cilium of R. precircumnodata (Hale 25004a) (X 400); c, amphithecial pycnidia of R. circurnnodata (Hale 26433) (X 170); d, vertical section of an amphithecial pyciiidium of R. circumizodata (Hale 26433) (X 500). 10 ShIITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE8.-Rhizine structure of Relicina: a, R. abstrusa (Oberwinkler and Poelt); b, R. limbata (Degelius A-368); c, R. subabstrusa (Malme s.n.); d, R. acrobotrys (A'eervoort 60 pp.); e, R. precircumnodata (Hale 25004a); f, R. ramosissima (Hale 30075); g, R. sublanea (Groenhart 8409). (All about X 80.) NUMBER 26 11

Stictic acid: R. limbata, R. sydneyensis. Succinprotocetraric acid: R. eximbricata, R. ramosissima, R. sublimbata. 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. The identification of these acids with TLC pre- sents no special problems. The protocetraric acid FIGURE9.-Ornamentation of apothecia: coronate rim of R. and related acids are separated most effectively in circumnodata (left) and retrorse rhizines of R. fluorescens hexane, where fumarprotocetraric acid is highest, (both about X 15). (Drawings by N. Halliday.) and succinprotocetraric acid lowest with proto- cetraric acid falling between them. Separation im- usual bicornute spore of R. precircumnodata and proves with an extra 10-20 ml of ether added to its vegetative morph R. circumnodata. These are the solvent. Both barbatic and 4-0-demethylbarbatic 2-3 X 10-13 pm in size, similar to the spores of acids occur in trace amounts; diffractaic acid oc- Bttlbothrix bicor-nuta (Muller Argau) Hale. curs in high concentration. Species containing echinocarpic acid produce several additional spots, none of which have been identified. Chemistry An unusual feature of Relicina is the predomi- The chemistry of Relicina was first summarized nance of P+ depsidones and the unidentified echi- by Hale and Kurokawa (1964), who used micro- nocarpic acid, which appears to be P+ depside. crystal tests and later some thin-layer chromatog- Echinocarpic acid seems to be especially charac- raphy (TLC). All of the specimens have been teristic of Relicina, although it occurs rarely in reexamined, as far as possible, with TLC (Merck Hypotr-achyna, usually in conjunction with bar- silica gel precoated plates) for this revision, using batic acid (Culberson and Hale, 1973), and in two solvent systems, hexane-ether-formic acid and Parmotrema along with protocetraric acid (Hale, benzene-dioxane-acetic acid. The substances identi- 197 1). fied in Relicina are listed below. Species with chem- The substances occur in several patterns. A com- ical strains are listed two or more times. mon one is acid A or acid A + B, as in R. acro- botryc (salazinic acid or salazinic acid with bar- Alectoronic acid: R. fluorescens. Atranorin: Sporadically produced in R. acrobotrys, R. lim- batic acid). An acid-deficient pattern (acid A or bata, K. planiuscula, and R. sydneyensis. no acid) is evident in R. amphithrix (no medullary Barbatic acid: R. abstrusa, R. acrobotrys, R. incongrua. substance or echinocarpic acid present). Both R. Caperatic acid: R. subconnivens. echinocarpa and R. fluorescens show the pattern Consalazinic acid: R. acrobotrys. acid A, acid A + B, and acid B. With R. echino- Diffractaic acid: R. abstrusa, R. incongrua. carpa this involves salazinic and echinocarpic acids Echinocarpic acid: R. amphithrix, R. echinocarpa, R. @ores- cens, R. planiuscula, R. samoensis, R. schizospatha. and with R. flziorescens alectoronic and echino- 4-0-Demethylbarbatic acid: R. abstrusa, R. acrobotrys, R. carpic acid?. Other examples of variation are dis- incongrua. cussed under the various species. I prefer not to Fumarprotocetraric acid: R. exirnbricata, R. malesiana, R. treat these chemical populations as species because ramosissima, R. relicinella, R. sublimbata. of the absence of any strong correlating morpho- Gyrophoric acid: R. luteoviridis. logical or geographical characters. Norstictic acid: R. abstrusa, R. limbata, R. subabstrusa, R. sydneyensis. Protocetraric acid: R. circumnodata, R. precircumnodata, R. sublanea. Ecology and Habitats “Quintaria” unknown: R. abstrusa. Protolichesterinic acid: R. connivens. Relicina occupies two main habitats, the canopy Salazinic acid: R. abstrusa, R. acrobotrys, R. echinocarpa, R. branches of dipterocarps and other trees in the subabstrusa. lowland rain forest and the trunk and branches of 12 S>lITHSOZJIAN CONTRIBUTIOTS TO BOTANY

Qzie)ciis at high elevations. A few species grow in as yet unknown in Europe, India and central Asia, secondary forests on trees such as coconut or even and Africa, except for a single isolated collection on rocks. of R. szibnbstrzisa on the Comoro Islands near The typical lowland dipterocarp forests occur in Madagascar. dense stands from sea level to less than 1000 m The genus obviously evolved in the ancient elevation. The highest locality at which I collected dipterocarp forests of Southeast Asia. The endemic on tlipterocarp was 850 m in Negros Occidental, neotropical species may be remnants of older more Philippines. Those species which occur in this ele- widespread species, although they are unrelated to vational range (and often only up to 400 m) any species now found in the Old World. Alter- include R. circiimnodata, R. conniven y, R. male- natively they may have evolved separately in the tiana, R. precircnmnodntn, R. rarnosissimn, R. re- New 12rorltl where conditions have obviously not licitiiila, R. wbabstrtisa, R. subconnivens, antl R. been favorable for rapid evolution and expansion tiiblanea. In the New T\'orld one finds R. incon- of the genus. grrra in a similar range of elevation. In almost all The number of species known in various geo- case5 I collected these species in areas where trees graphic areas is summarized in Figure 10. The were being felled commercially, and since the dip- actual species by region are tabulated as follows: terocarp forests do not usually regenerate, Relicinn Japan: R. nbstrusn, R. echinocnrpa, R. planiusculn, and R. could become extinct as the forests are timbered sydn eyensis. throughout southeast Asia. Taiwan: R. nbstrirsn, R. nanlesiann, R. plnniuscula, R. sub- Oak and, on Luzon, Philippines, Pinii~make up nbstrirsn, and R. sydneyensis. the typical vegetation above about 1400 m eleva- Philippines: R. nhstrusn, R. ninphithrix, R. circumnodntn, R. tion on mountains in Southeast Asia. The charac- conniz~ens,R. mnlesinnn, R. plnnizisciiln, R. precircumno- dntn, R. roniosissinzn, R. reliciniiln, R. snmoensis, R. schizo- acteristic species here are R. acrobotyrs, R. fliiores- spothn, R. sitbnbstritso, and R. sublnnen. fen$, R. liitroviridis, R. planiuscziln, and R. hfalaya: R. nbstrusn, R. ninphithrix, R. circumnodntn, R. tchizo7t)atho. These oak forests are being cut over plan iitsc~iln,R. rninosissiinn, R. relicin iiln, R. schi:ospathn, for vegetable farms and as with the dipterocarp R. s~cbconnivenr,and R. srihlnnen. forests many of the Relicinn species will eventually Indonesia: R. nbstritsn, R. acrobotrys, R. connivens, R. plani- hecome extinct. usculo, R. minosissinin, R. relicinuln, R. samoensis, R. schizospnthn, R. sciblnnen, and R. sydneyensis. Several temperate species occur on trees and Sarawak: R. nbstrzisn, R. circiimnodntn, and R. subnbstrusn. rocks at relatively low elevation. These include Sabah: R. nrrobotrys, R. nmphithrix, R. fluorexens, R. luteo- R. echinocnrpa, R. limbata, and R. sydneyensis. ?tiridis, R. ~nnlrsinnn,R. plirniirscitln, R. rnmosissimn, and Also occurring at low elevation but in disturbed R. schizospnthn. forest5 in tropical regions are R. samoensis antl New Guinea: R. connivens, R. fluorescens, and R. malesinnn. R. cxim bi icatn. Relicinn wblimbatn seems to occur Australia: R. limbntn, R. subnbstrnsn. and R. sydneyensis. Pacific Islands: R. connizxws and R. snmoensis. in drier subtropical areas of Burma and Thailand Thailand-Burma: R. siiblimbnto. at mid elevations. Comoro Islands: R. sit bn bstrusn. Two species, one restricted to Asia (R. ampki- U.S..A,: R. nbstrtm and R. eximbricntn. thrix) and the other pantropical outside of Africa hlexico: R. nbstrusa. (R. nbrtriiro), have very great elevational ranges, IVest Indie?: R. nbstrusn and R. eximbricntn. Panama: R. nhstrtisn and R. incongrira. from 150 m to over 1600 m. Colombia: R. nbsfrusn, R. relicinella. Venezuela: R. nbstrusn. Phyto geo gr aphy Brazil: R. nbstriisn, R. incongrzln, R. relicinello, and R. sub- nbst rusn. The most distinctive feature of Relicinn is the Paraguay: R. nbstritsn and R. sribnbstrusn. high degree of endemism in Southeast Asia. Out- Argentina: R. nbstrusn. side of R. abstrzim and R. wbabstrzisn, which occur The commonest species of Relicina are as fol- in the Neotropics a5 well as in Asia, and three neo- lows lvith number of collections examined: R. sub- tropical endemics (R. eximbl-icntn, R. incongrua, ob.ttrii,ro (52), R. omphithrix (43), R. circiimnodnta R. relicinella), all remaining 19 species occur only (40), R. plnrii?i.tcziln (37), R. reliciniila (31), R. in the Old TVorld. At the same time the genus is nbstrir.tn (26), and R. schizospnthn (20). Rarities NUMBER 26 13

?

include R. luteoviridis (3), R. precircumnodata the most characteristic acid, echinocarpic acid, is (3), and R. siibconniveizs (1). as yet unknown. The following discussions, there- The localities where especially large numbers of fore, will cover only the more obvious lines of specimens were collected include logging areas in evolution. Negros Occidental, Philippines (44 collections), Selangor, Malaya (40), and Mountain Province, MORPHOLOGICALEVOLUTION Philippines (36). The richest localities had six or seven species each with considerable differences in FomrATIoN OF MORPHS.-A vegetative morph is species composition. defined as an isidiate or sorediate population which is absolutely identical in chemistry and morphology (excepting the diaspores) to a fertile, nonisidiate Evolution and Speciation or nonsorediate parent population. A discussion of the development of the theory of morps in lichens Very little is known or even hypothesized about is presented by Hale (1975). In Hypot?”aclzyna,for evolution and speciation in lichens. Lacking any example, more than half of the speciation can be fossil record, lichenologists are left with few clues attributed to the formation of morphs. A similar that can be used to trace evolution of species and situation is known in Paimotl-emn (Hale, 1965a; genera. Chemistry has proven to be a valuable Cullierson, 1973). independent check in some groups, especially the Relirintl is exceptional in that soredia are not Parmeliaceae. I have already discussed in some formed by any of the species. Isidia do occur, how- detail the evolution of species in Hypotmchyna ever, in five species, and four of these can be (Hale, 1975), where hybrid chemistries and rich paired witli nonisidiate sexual morphs: development of vegetative morphs permit one to R. 0 bsfYlLsn-R. Sli 110 bstru~n draw a fairly complete picture of speciation. By R. rimi inii odatn-R . precircu iniiodnto comparison, Rrlicintl has fewer vegetative morphs R. ~~la?riz~scziln-R.fliiorcsrciis (echinocarpic acid population) and the chemistry is less varied. The structure of R. sydiieyensis-R. limbata 14 SMITHSONIAN CONTRIBUTIONS TO BOTANY

The acid deficient population of R. amphithrix seen in R. connivens (ecoronate) and R. subconni- is closely related to nonisidiate R. relicinula and vens (coronate) where the spores have not changed may represent a vegetative morph, but variation in but the chemistry is altered (protolichesternic acid isidia and lobule development is so great that the versus caperatic acid). In most other species of evidence for a direct derivation is poor. ReZicina, however, parallel ecoronate-coronate RHIZINES.-The type of rhizine, simple or populations are not closely matched, or as in the branched, appears to be an important character. R. eximbricata group in the New World no ecoro- The five species with branched agglutinated, pale nate parent populations are extant. Conversely no rhizines seem to form a close group that originated coronate species have evolved from the R. limbata in and has not migrated out of the dipterocarp group. forests. One, R. acrobotrys, occurs only in high elevation forests and, perhaps significantly, is chem- CHEMICALEVOLUTION ically completely distinct from the other four species in containing salazinic and barbatic acids. Chemical evolution is presumed to involve The other species contain protocetraric acid or genetic processes that lead to differentiation of a fumarprotocetraric acid and occur only at low parent species into two or more chemical popula- elevations. One series, including R. 1-amosissima tions. The patterns may be extremely clear-cut and antl R. rziblanea, is ecoronate and has small ovoid easy to follow, as in Hypotrachyna (Hale, 1975), spores; the other series, including R. precircumno- or too complex to interpret. Relicina has a less data antl its isidiate morpli R. circzimnodata, is varied chemistry but several examples can be coronate and has cornute spores. The main lines given. One is the difference between the two main of evolution here seem to be ecological isolation populations of R. abstnisa and R. subnbstrzisa. All and differentiation in apothecial and spore charac- specimens of these two related species contain ters (Figure 11). approximately equal amounts of norstictic acid CORONATEAPOTHECIA.-I am assuming that ecor- and salazinic acid in the New World. Most speci- onate species are more primitive than coronate mens in the Old IVorld, however, even though species and gave rise by mutation to coronate externally indistinguishable from those in the New species. This seems fairly evident in the R. sub- T.Vorlc1, usually lack salazinic acid or have at most lanea-R. cimimnodnta group discussed above trace amounts. The constancy of the difference where the coronate condition is paralleled by spore suggests that the two major populations have been evolution and where chemistry is not significantly isolated for a long period and that no genetic changed. A somewhat different pattern can be interchange has occurred during this period of isolation. The New World populations have in Progeniror 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 Ecoronate progenitor Coronate progenitor example to R. incongrua. If true, we can expect other chemical evolution in the New World, espe- R. acrobotrys/\ cially since apothecia are common in both the 4 isidiate anti noni5idiate populations, whereas the R. sublanea R. prerirci~mnodata more conservative Old World isidiate population I usually lacks apotliecia and gives no evidence of J continuing chemical evolution. R. raniosissima Another example of possible backcrossing can be cited for the R. fEuoYescens group. The two R. rirriimnodata species here, R. piiol-escens and R. planiurciila, are FIGURE11.-Hypothetical evolution of species with branched, morphologically very close because of the strongly pale rhizines. developed columnar cortex and similar habitats at NUMBER 26 15 higher elevations. The presumptive parent, R. A common situation in Relicina is the differen- Puorescens, has a complex chemical population tiation of species into parallel chemical groups structure, alectoronic acid alone, echinocarpic acid with various combinations of fumarprotocetraric alone, and the two acids occurring jointly (Figure acid and other substances. For example R. sublim- 12). The three populations occur together in sig- bata and R. nzalesiana are essentially identical but nificant numbers in Sabah. The vegetative morph, R. sublimbata produces succinprotocetraric acid R. planiuscula, contains only echinocarpic acid. If along with fumarprotocetraric acid, while R. male- we are correct in assuming that R. planiuscula siana produces only fumarprotocetraric acid. In evolved from R. fluorescens or a sexual morph this instance the species have a different geographic similar to it, as shown in Figure 12, then the range, R. sublimbata occurring in Burma and morph must have developed before R. fiuorescens Thailand, R. malesiana from Taiwan south to underwent chemical evolution. A corallary of this New Guinea apart from the Asian mainland. assumption is that the progenitor population con- tained echinocarpic acid. Perhaps the alectoronic acid-containing populations of R. fiuorescens have Position of Relicina in the Parmeliaceae not had time to develop isidiate morphs, or if they have evolved they cannot, for reasons we cannot Relicina has an isolated position in the Parme- even guess at this time, compete with the now liaceae because of its morphology and phytogeog- widespread echinocarpic acid-containing morph raphy. First, it has nothing to do with Xantho- R. planiuscula. parmelia, with which it shares only the yellow Another example of how species have diverged cortical pigment usnic acid. The only closely slightly in chemistry and apparently become repro- related genus is Bulbothrix (Hale, 1974), which ductively isolated is R. eximbricata and R. relici- also has marginal bulbate cilia and coronate apo- nella in the New World. While these externally thecia but which produces only atranorin in the similar species share the production of fumarproto- cortex. Bulbothrix, however, is rare in the range cetraric acid, R. eximbricata also produces succin- of Relicina. Only B. subdissecta (Nylander) Hale protocetraric acid, while R. relicinella contains an and B. pigmentarea (Hale) Hale occur in the low- unidentified P + compound, also apparently re- land rain forests of Southeast Asia and B. Jubin- lated to protocetraric acid. More significantly, R. flata (Hale) Hale in oak forests where one finds relicinella has multispored asci. Relicina. Bulbothrix is largely a genus of the drier The species containing echinocarpic acid form subtropical zones, being especially common in the niicleiis of the genus in Southeast Asia. We southeastern United States and Africa. It generally cannot speculate on the chemical evolution of this has larger spores, 12-19 pm long, twice the average group until the molecular structure of echinocarpic size of those in Relicina, although a few species do acid is determined. have spores only 2-4 pm long. The chemistry of Bulbothrix is different in many respects and there is little evidence of any significant chromosomal Progenitor exchange with Relicina that might appear in the chemistry. Many species of Bulbothrix, for exam- ple, have gyrophoric or salazinic acid, both of these very rare in Relicina. No barbatic, difiactaic, echi- R. fruoresrens R. frtdoresrens nocarpic, furmaprotocetraric, or stictic acids are [echinocarpic acid] [alectoronic acid] produced. On the other hand, Bulbothrix produces lecanoric, lobaric, and colensoic acids, all unknown in Relicina. I would argue then that Bulbothrix and Relicina as they presently stand are not closely R. planiiisrrda R. fruorescens related biologically. This does not rule out the [echinocarpic & alectoronic acids] possibility that they have a distant common origin FIGURE12.-Hypothetical evolution of chemical populations which may have centered on the usnic acid- in R. fluorexens and R. planiuscula. 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. planiusda 2. Isidia becoming dorsiventral and lobulate or only lobules present. 7. Lobes narrow and appressed, 0.5-1.0 mm wide. 3. R. amphithrix 7. Lobes broader and adnate, 1-3 mm wide. 8. Medulla C+ red; lobules densely developed. 11. R. luteoviridis 8 Medulla C- or dull )ellowish red (and Pf red); lobules not dense. 9. Thallus coriaceous; upper cortex columnar. 13 R. planiuscula 9. Thallus thinner; upper cortex not columnar. 19. R. schizospatha 1. Thallus lacking isidia and lobules (some adventitious marginal lobules may be present). 10. Lower surface tan to pale brown. 11. Rhizines simple. 10. R. lirnbata 11. Rhizines densely branched. 12. Medulla K+ red. 2. R. acrobotrys 12. Medulla K-. 13. Apothecia present, coronate. 14. R. precircumnodata 13. Apothecia ecoronate or lacking. 14. Protocetraric acid present. 22. R. sublanea 14. Fumarprotocetraric acid present. 15. R. ramodssima 10. Lower surface black. 15. Apothecia present, coronate. 16. Medulla K+ red. 20. R. subabstrusa 16. Medulla K-. 17 Medulla P-. 18. Diffractaic acid present; New World species. 9. R. incongrua 18. Diffractaic acid absent; Old World species 19. Lobes narrow and appressed, 0.5-1 0 mm Tside. 17. R. relicinula 19. Lobes broader and adnate, 1-3 mm wide. 2 1. R. subconnivens 17 Medulla P+ orange red. 20. Echinocarpic acid present; Old World species. 18. R. sanioensis 20. Fumarprotocetraric acid present; New World species. 21. Spores 8/ascus. 7. R. eximbricata 21. Spores 16-32/ascus. 16. R. relicinella 15. Apothecia ecoronate or lacking. 22. Medulla K+ red. 23. Salazinic acid only present. 6. R. echinocmpa 23. Norstictic and salazinic acids present. 20. R. subabstrusa 22. Medulla K-. 24. Medulla P -. 25, Lobes narrow and appressed, 0.5-1 mm wide. 17. R. relicinula 25. Lobes broader and adnate, 1-3 mm wide. 26 Upper cortex columnar; alectoronic acid present. 8. R. Puorescens 26. Upper cortex not columnar; protolichesterinic acid present. 5. R. connivens 24. Medulla Pf orange red. 27. Echinocarpic acid present 28. Upper cortex columnar. 8. R. puorescem 28. Upper cortex not columnar. 6. R. echinocarpa NUMBER 26 17

27. Fumarprotocetraric acid present. 29. Succinprotocetraric acid present. 23. R. sublimbata 29. Succinprotocetraric acid lacking. 12. R. rnalesiana

Species Treatment isidiate, the disc carob brown; spores 8, 4-5 X 5-6 pm. Relicina CHEhiIsTRY.-I1/Iedulla Kf yellow turning red, C--, KC-, Pi- orange, norstictic and usnic acids Relicina (Hale and Kurokawa) Hale, 1974:484. with or without salazinic acid and very rarely with Pnrmelia subgenus Purmelia section lmbricaria subsection Bicornutae series Relicinae Hale and Kurokawa, 1964: 135. diffractaic acid and a trace of barbatic acid and 4-0-denietliylbarbatic acid. A “quintaria” unknown, Type-species: Parmelia relicinn Fries, 1825:283. the reddish pink spot just below norstictic acid in My typification of the genus (Hale, 1974) was both solvent systems, occurs in about one-quarter not completely accurate since Parmelia relicinn of the specimens. must be designated as the type-species even though Rmr.mKS.-This is a very common species in it becomes a tautonym when transferred to Reli- both the New 1170rld and the Old World. As with cinn. Furthermore, my typification of Pnrmelia R. szibabstmsn, the presumptive nonisidiate par- eumorphn Hepp, based on an incorrectly deter- ent, the Sew World specimens produce norstictic mined specimen in the Rijksherbarium (L), which and salazinic acids in nearly equal concentration, I considered to be a valid type, is in error. As but salazinic acid is either absent or present in Muller Argau (1882) had pointed out, Hepp’s trace amounts in Old 1170rld specimens. The two species is a Coccocnrpia and the specimen which populations also differ in elevational range. In Muller Argau examined (in G) is indeed a Cocco- Southeast Asia R. nbstix~nocc~irs commonly from cnl-pin, although the same number in L is a 100 m to 1400 m elevation. The same species in Relicinn species. Finally, Hepp’s protologue ob- South America ranges from 700-2150 m. Xpothecia viouslj refers to a nonparmelioid genus. are much more common in the New Tl’orld popu- The species are listed alphabetically in the list- lations (5Sm, frequency) than in the Old Ii’orld ing below. All collections by Hale are deposited (7yo frequency). in the National Museum of Natural History, One specimen from Brazil (Lichenes nirsfro- Smithsonian Institution, and the herbarium acro- nmeiicnni 95) contained diffractaic acid and a trace nym (US) is not given. 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 1. Relicina abstrusa species such as R. incongrirn, which contains only diffractaic and barbatic acids. FIGURE13a Vainio (1 890) considered Parmelin lim bntn f. Relicinn nbstrusu (Vainio) Hale, 1974:484. i,sidio.cn hfuller Argau to be a synonym of P. nb- Parnzelin nbsti ma Vainio, 1890:G4 [ripe-collection: Caraca, .rfriisa, but Muller’s taxon is now recognized as Minas Gerais, Brazil, 1/ni?iio 1347 (not 1346) (TUR, lecto- Relicinn ,xydney~?i,si,r.A\saliina (1951) identified nor- type; UPS, isolectotype)]. stictic and stictic acids in Japanese specimens, Thallus closely adnate on bark or rocks, 6-10 whicli lie identified as P. nb.rt~ir.rnand these too cm in diameter, pale chartreuse yellow; lobes sub- represent Rclirinn ,sydneyei?sis. linear, 1-2 mm wide; upper surface plane to con- SFECISI~~SExh\rIxrD.--Florida: Baker Coniltv. HI!/<’21968. vex, continuous or faintly maculate, moderately Xlexico: Chiapas. Hnlf 20028 (C. ITS), 20120, 20898. 2014,;. isidiate, isidia short, mostly simple: bulbate cilia Panama: Panama, Hnlr 88116. 38447. Jnniaic:i: Ci/lber.ioti moderately inflated; lower surface black and rliizi- 13748 (DKTKE), Plift (1-S). Coloml,ia: r.iiidig 701 (RJI. P). l’enezuela: Federal, Ol~rrri’i~iklrrnnrf Poclt (\I. t.S): Merida, nate, the rhi7ines simple, shiny (Figure 80). Apo- Hale 42070, 48886; 3Ierindn. I~cii.csclii289ic (t’q). R~i~il:Rio thecia adnate, 1-4 mm in diameter, coronate, de Jatieiro. Eitrti 6541, 7175R. 7671 (1‘s). Glntioic 2000 (I’PS): basally retrorsely rhizinate, the amphithecium Minas Gerais, I’nitlio 1589 (TI’R. Ill’s): Rio C.r:inde do Sill, 18 SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE13.--Species of Relicina: a, R. abstrusa (Hale 20393); b, R. acrobotrys (h'eeruoort GO pp.); c, R. amphithrix (Hale 30051); d, R. circumnodata (Maingay in BM); e, R. conniuens (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), 3. Relicina amphithrix, new species 1857 (BM, S), 1865B (S), 1884 (S, W), 2745 (S), 2749B (S), in Lichenes austroamericani 94 (G, H, UPS), 95 (G, H, LD, S, FIGURE13c UPS, US). Paraguay: Paraguari, Malme 1537 (S, US). Argen- Thallus appressus, corticola, viridi-flavicans, 2-6 tina: Misiones, Osorio 4666 (WIS). Japan: Hyuga, Hale cm diametro, lobis sublinearibus, angustis, 0.3- 29617, 29646, 29647, 29667; Kii, Kurokawa 59085 (TNS, US). Philippines: Mountain, Hale 25769, 25818, 25828, 25881, 1.O mm latis, margine lobulascentibus, lobulis raro 25888; Cagayan, Hale 25625, 25645, 25674, 25675, 25734; palmato-dactyloideis, bulbo-ciliatis, bulbis inflatis, Negros Occidental, Hale 26432. Malaya: Pahang, Hale 30090, congestis, aetate apice ramosis, superne dense isi- 30148, 30149; Selangor, Hale 29927, 30058, 30059, 30072, 30090, diatis, isidiis cylindricis, cylindricis vel sparse ra- 20149, 30296, 20397, 30298. Sarawak: Hale 29993. Java: Groen- mosis, pro parte isidiato-lobulascentibus; cortex hart 5868 (US), A'eercioort 4110 (BO). See Kurokawa (1965: 265) for records for Taiwan. 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 2. Relicina acrobotys rhizinosus, rhizinis nigris, nitidis, simplicibus vel sparse ramosis. Apothecia rara, adnata, usque ad FIGURE13b 1 mm diametro, amphithecio coronato, basin retror- Relicina acrobotrys (Kurokawa) Hale, 1974:484. so-rhizinato, disco imperforator, hymenio 30-40 pm Parmelia acrobotrys Kurokawa in Hale and Kurokawa, 1964: alto, sporis octonis, 3 X 4 pm. 142 [type-collection: Tjibodas, Java, Neeruoort 60 pr. p. CHEMISTRY.-hfedulla K - , C -k , KC reddish, (BO, holotype; US, isotype)]. + P+ red or medulla not reacting with color tests, Thallus adnate on bark, rather coriaceous, 6-12 usnic and echinocarpic acids and associated un- cm in diameter; lobes sublinear-elongate, 1-2.5 known substances or only usnic acid present. mm wide, narrowly black rimmed; bulbate cilia HoLoTYPE.-oak forest, elevation about 1600 m, tapered, not strongly inflated; upper surface plane Cameron Highlands, Pahang, Malaya, M. E. Hale or rugulose, continuous; lower surface black, 30203, 2 March 1965 (US). densely rhizinate except for a narrow brown zone REMARKs.-This is the most variable species in near the tips, the rhizines black, irregularly densely Relicina and may well represent more than one branched (Figure 8d). Apothecia numerous (fre- species. The typical material is clearly isidiate, but quency loo%), adnate, 1-2.5 mm in diameter, the isidia may become procumbent or lobulate. In ecoronate, with basal retrorse rhizines; spores 8, the extreme case the lobules are far more numerous 2-3 X 4-6 pm. than isidia over the thallus surface. Intergradation CHEMIsTRY.-Medulla K+ yellow turning red, with R. schizospatha is a distinct possibility al- C--, KC- or KC+ orange, P-, atranorin, usnic, though the two species are fairly easily separated consalazinic, and salacinic acids with or without on thallus size and lobe width. A small group of barbatic acid and 4-0-demethylbarbatic acid. specimens has closely appressed, mostly marginal HABITATS.-on trunk and branches of Quercus palmately branched lobules (Figure Ic) often in open forests at 1400-1600 m elevation. occurring without isidia but seeming to originate DIsTRIBuTIoN.-Java and Sabah. from or to intergrade with isidia when isidia are REXfARKs.-This is the only species with bushy present. agglutinated rhizines that occurs at high elevation. Relicina amphithrix is probably related to R. It is also the only species in Relicina with salazinic relicinicla since both have very narrow lobes and acid as the main chemical component. While the coronate apothecia. Relicina wlicin !/la, hoicever, type contains only salazinic acid, specimens col- does not produce echinocarpic acid and would lected later in Sabah all contain barbatic acid in therefore differ from the bulk of the specimens addition. Barbatic acid is also produced in trace identified as R. amphithi.ix. On the other hand. it amounts by R. abstrusa and R. incongrzta, but seems probable that the acid deficient population these New World populations have no relationship of R. amphithrix is the isidiate morph of R. whatsoever to R. acrobotrys. relicinzila. SPECIMENSEXAMINED.-sabah: Hale 28126, 28540, 29123. SrEcrartxs EX~VINFD(echinocnipic ncid present, isidinte or Indonesia: Java, Kurokawa 2167 (TNS, US). becoming lobalnte-isidinte).--Philippiues: AIouiitnin, Hole 20 ShIITHSONIAN CONTRIBUTIOXS TO BOTANY

25824; Negros Occidental, Hale 26420, 26437, 26445, 26480, Philippines, northern Borneo, and Malaya. It has 26481, 26522, 26569, 26630, 26654; Agusan, Hale 24481, 25064, not yet been collected south of this region in 25191; Surigao del Stir, Hale 24674. Malaya: Pahang, Hale Indonesia but probably occurs there ;IS well. 30204, 30211, 30479, 20493; Selangor, Hale 30066, 30070. Sabah: Hale 28239, 28820, 29076. SPECIX~ENSExAhrl~ED.-Philippines: Mountain, Hale 25787, SPECIMENSEXARIINFD (echinocarpic acid, marginally palmate- 23802, 25810, 25820, 25841, 25873, 25902: Cagayan, Hale lobulate).-Philippines: hfountain, Hale 25770, 25789; Agusan, 24410, 25736; Quezon, Hale 26878, 26879, 26887, 26900, 26920, Hale 24462. Malaya: Selangor, Hale 30060, 30063, 30256, 26923, 26926, 26928, 26934, 26980; Negros Occidental, Hale 30257, 30258, 30259, 30265, 30283, 30287, 31185, 31186. 26421, 26433, 26474, 26500, 26317, 26561, 26572, 26580, 26599, SPECIMENSEXAMINED (echinocarpic acid absent, isidiate or 26631, 26633, 26651, 266.52. XIalaya: Pahang, Hale 30144; becoming lobulate isidiate).-Philippines: Agusan, Hale Selangor, Hole 30056. Sarawak: Hale 29995, 29996, 29997, 24454. Malaya: Selangor, Hale 30051, 30064, 30065, 30069, 299999, 30437. 50071, 30288.

5. Relicina connivens 4. Relicina circumnodata FIGURE13e FIGURE13d Relicirm ro~iuivens (Kurokarva) Hale, 1974:484. Relicinn circumnodata (Sylander) Hale, 1974:484. Partrielin rotinivens Kurokawa irz Hale and Kurokawa, 1964: Parmelia circumnodata Sylander in Crombie, 1883:51 [type- 142 [type collection: Milne nay district, Papua, Sew collection: Government Hill, Penang, Malaya, Maingay 22 Guinea, Hooglond 4i26 (BM, holotype; L, US, isotypes)]. (BM,lectotype; FH, H, isolectotypes)]. Tliallus closely adnate on bark, 3-8 cm in diam- Thallus adnate on bark, coriaceous, 3-10 cm in eter; lobes sublinear-elongate, 0.7-2 mm wide; buI- diameter; lobes sublinear-elongate, 1-3 mm wide; bate cilia dense and conspicuous, strongly inflated upper surface plane to convex, becoming rugulose, and becoming globose and apically densely continuous to faintly maculate; bulbate cilia large, Ixanclied; upper surface plane to convex, continu- becoming globose; lower surface black, densely ous or faintly maculate, sparsely isidiate, the isidia rhirinate, the rhizines black, siniple to irregularly simple, to 0.5 mm high; lower surface pale brown, densely branched. Apothecia common (frequency rhizinate, the rhizines pale, more or less densely My;), atlnate, 1-2.5 mm in diameter, the rim branched and agglutinated. hpothecia rare (fre- ecoronate, the base retrorsely rhizinate; spores 8, quency lo%), adnate, 1-2 mm in diameter, rim 2-3 X 4-6 pm. coronate; spores 8, cornute, 3 X 10-12 pm. CHEMlSTRY.-;\/IedUl~aK-, C-, KC-, P--, pro- CHEX\'rISTRY.-MedU11a K-, C-, KC- or KCS- tolichesteric and usnic acids. rose, Pf red, protocetraric and usnic acids. HAnrTATs.-on trunk and canopy branches of HABITATs.-on trunks and canopy branches of tlipterocarps in rain forest and on other trees in tlipterocarp and other trees in rain forest at sea secondary forest at 200-850 m elevation. level to I300 m elevation. DIsTRIBUTroN.-Phi~ippines, Indonesia, New DlsTRIBuTIoN.-iC~alaya, Philippines, Sarawak. Guinea, Guam, and the Solomon Islands. RnrmKs.-This species was described by Ny- Rnr.mKs.-This species has proved to be wide- lander from sterile material. Apothecia are now spread hut not common in a band from the Philip- known from 4 of 39 specimens that I have been pines through Xew Guinea to the Solomon Islands, able to examine. All are coronate and have un- occurring generally at 300-800 m elevation. No usual bicornute spores, exactly the same as those other species in the genus produces protolichesteri- in the presumptive parent species R. precirciirnno- nic acid. It pro1)ably would be mistaken for R. data. The bulbate cilia are very large, globose, m(~le.~in~in(P+ red) without appropriate chemical and crowded on older parts of the lobes, also as in testing. Closely related R. szibconniuens has coro- the parent species. The lower surface is consistently nate apothecia but if collected sterile it could be pale brown and the rhizines initially simple but differentiated by the presence of caperatic acid. soon branching in the pecular bushy manner of SPECI\IENS ExAzrisrD.-Pliilippines: Segroes Occidental, this group. Hale 26457, 26,709, 26626a: Agusan, Hale, 25025, 25034, 25044. The species is very common throughout the Colomon Islands: Guadalcanal, Hill 9057 (EM,US). hddi- NUMBER 26 21 tional records from the Molucca Islands and Guam are listed 1938:288 [type-collection: Monte Rus, Cuba, Hioram 10506 in Hale and Kurokawa (1964:143). (BP, holotype)]. Parmelia eximbricata (Gyelnik) Hale and Kurokawa, 1964: 143. 6. Relicina echinocarpa Thallus closely adnate on twigs or bark, 3-5 cm FIGURE13f broad; lobes sublinear, 0.7-1.5 mm wide; upper surface plane and continuous; lower surface black, Relicina echinocarpa (Kurokawa) Hale, 1974:484. densely rhizinate, the rhizines simple. Apothecia Parmelia echinocarpa Kurokawa, 1965:265 [type-collection: Mt. Hikosan, Prov. Buzen, Japan, Kurokawa 63163 (TNS, very common (frequency 100yo), adnate, 0.5-1.5 holot ype)]. mm in diameter, the exciple coronate, the base often retrorsely rhizinate; spores 8, 4-5 X 6-9 pm. Thallus adnate, to appressed, corticolous or saxi- CHEMISTRY.-bfedUlla K - , c - , KC - , P + brick colous, 2-6 cm in diameter; lobes sublinear- red, fumarprotocetraric, succinprotocetraric, and elongate, 1-2 mm wide; bulbate cilia gradually in- usnic acids. flated; upper surface plane, faintly maculate; lower HARITATs.-T~u~~sand branches of trees (cash- surface black, densely rhizinate, the rhizines simple ew, Eugenia) in open areas from sea level to 600 m or sparsely branched. Apothecia numerous (fre- elevation. quency 10070), adnate, ecoronate, 1-2.5 mm in D!s~~~~uTro~.-Floridaand the West Indies. diameter, with basal retrorse rhizines; spores 8, REMARKs.-Relicina eximbricata has been col- 3-5 X 6-8 pm. lected for the most part in disturbed secondary CHEhiISTRY.-Medulla K+ yellow, c- or c+ forest. It is rather rare if the few collections avail- pale yellow, KC-, Pf orange, echinocarpic and able reflect its true abundance. I was unable, for usnic acids and a C+ unknown with or without example, to recollect any specimens at the locality salazinic acid. in Florida visited by Swanson. It is extremely close HARITATS.-on trunk and canopy branches, to R. relicinella in external appearance but differs more rarely on rocks, in open deciduous forest at in spore number (8 versus 16-32) and presence of low elevation. succinprotocetraric acid. D I STR I R UTI ON.-J a pa n . SPECIMENS EXAhTINED.-see Hale & Kurokawa (1964: 144) for REMARKs.-Relicina echinocarpa is a small, close- specimens from Florida, Cuba, Grand Cayman, Jamaica, and ly adnate species collected on upper branches of the Dominican Republic. deciduous trees or on rocks in southern Japan, where it appears to be endemic. While it usually contains echinocarpic acid, it is not related to the 8. Relicina fluorescens tropical members of the genus containing this acid. FIGURE14b For example, R. .mmoensis has more globose bul- he and coronate apothecia. Relicina Jluorescens (Hale) Hale, 1974:484. The chemical variation in R. echinocarpa follows Parmelia fluorescens Hale, 1965:202 [type-collection: between a typical addition pattern. Specimens may contain Kamharanga and the second radio tower, Tourist Trail, Kinahalu National Park, Sabah, Hale 28637 (US, holotype; echinocarpic acid alone, echinocarpic and salazinic TNS, isotype)]. acids together, and in one or two specimens ap- parently salazinic acid alone. Thallus loosely attached to adnate, coriaceous, corticolous, 4-7 cm broad; lobes sublinear-elongate, SP€CIMF.NSExAMlNFD.-Japan: Hyuga, Hale 29596, 2961 3, 1-3 mm wide; bulbate cilia tapered, not strongly 29626, 29638, 29648, 29658, 29659: Higo, Mayebara (TNS). inflated; upper surface plane to rugulose and finely pitted, faintly maculate; upper cortex strongly 7. Relicina eximbricata columnar (Figure 5a); surface black, densely rhizi- nate, the rhizines simple. to sparsely branched. FIGURE140 Apothecia common (frequency 50%,), adnate, 2-3 Relicinn eximbricntn (Gyclnik) Hale, 1974:484. mm in diameter, ecoronate, usually with basal Parmelia sanioensis Zahlhruckncr var. eximbricata Gyelnik, retrorse rhizines; spores 8, 4 X 5 pm. 22 SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE14.-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 retrorso-rhizinato, hymenio 45-55 pm alto, sporis P- or P+ usnic and alectoronic acids with or octonis, 4-5 X 5-7 ym. without echinocarpic acid or echinocarpic acid CHEMISTRY.-Meduh K-, c - , KC -, KC +, alone in the medulla. P - , usnic, diffractaic, barbatic, and 4-O-demethyl- HABITATS.-on trunk and branches of @MCUS barbatic acids. and other trees in open forest at 1600 to 2400 m HoLoTYPE.-santa Anna da Chapada, Mato elevation. Grosso, Brazil, Malme 2433C, 27 February 1894 DIsTRiBuTIoN.-sabah and New Guinea. (S; isotype in US). REMARKs.-Relicina fluorescens was first recog- HABITAT.-oII trees in primary forests at low to nized by the presence of alectoronic acid. On later mid elevation. study I decided to lump here a series of chemically DIsTRIBUTI0N.-Panama and Brazil. variable specimens occurring at high elevation, REMARKS.-This rare species is externally identi- mostly in Sabah. All have coriaceous thalli and a cal with R. subabstrzisa, which contains norstictic columnar cortex. Once again the primary substance and salazinic acids. Since this seems to represent a is echinocarpic acid with or without alectoronic replacement type of chemical population, I am acid, but a sizeable part of the population may considering it as a distinct species, very closely lack echinocarpic acid. The unknown substance related to R. subabstrusa and probably derived occurring with alectoronic acid in four specimens with it from a common progenitor. Both of these listed below has an RFof about 0.25 in the hexane species have unfortunately been only rarely col- solvent and about 0.6 in the benzene solvent. It is lected in tropical America. brownish when developed with sulfuric acid. a- SPECIMENSExAMINED.-Panama: Canal Zone, Hale 43448. Collatolic acid does not occur in this species. Brazil: Mato Grosso, Alalme 2445 (S, US). The isidiate morph of R. fluorescens is R. plan- iuscula, which has a broader geographic range. 10. Relicina limbata SPECIMENSEXAMINED (alectoronic acid present).-New FIGURE14d Guinea. Western Highland: Kurokawa 75 in Lichenes Rari- ores et Critici Exsiccati (US). See Hale (1965: 203) for addi- Relicina limbata (Laurer) Hale, 1974:484. tional records from Sabah. Parmelia limbata Laurer, 1827:39 [type-collection: Australia, SPECIMENSEXAMINED (alectoronic and echinocarpic acids Sieber (M, lectotype)]. present).-Sabah: Hale 28510, 28512, 29048, 29248. Parmelia sphaerospora Knight, 1882:49 [type-collection: Syd- SPECIMENSEXAMINED (echinocarpic acid present).-Sabah: ney, Australia, Knight (H, lectotype) (not Parmeh Hale 28703, 28941, 29088. sphaerospora Nylander, 1859:254)]. SPECIMENSEXAMINED (alectoronic acid and an unidentified Parmelia insinuata Nylander, 1886:324 [based 011 Pnrmelia substance).-Sabah: Hale 28322, 28750, 29031, 29053. sphaerospora Knight]. Thallus adnate to loosely attached on bark or rock, coriaceous, 4-9 cm in diameter; lobes sub- 9. Relicina incongrua, new species linear-elongate, 0.8-2.5 mm wide; bulbate cilia FIGURE14c conspicuous, up to 2.5 mm long; upper surface plane to convex, maculate, cracked on older lobes; Thallus adnatus vel appressus, corticola, viridi- lower surface brown to blackish brown, moderately flavicans, circa 6 cm diametro, lobis sublinearibus, rhizinate, the rhizines pale brown, coarse, mostly contiguis, 0.7-1.5 mm latis, margine modice simple (Figure 8b). Apothecia yery common (fre- bulbato-ciliatis, bulbis aetate inflatis globosisque; quency 100~o),adnate, 1-6 inm in diameter, ecoro- cortex superior circa 10 pm crassus, stratum goni- nate, basally retrorsely rhizinate; spores 8, 4-6 X diale 10-12 pm crassum, medulla 70-80 pm crassa, 7-10 pm. cortex inferior 10-12 pm crassis (Figure 2c); sub- CHEMISTRY.-h’~edU11a K+ yellow, c-, KC-, tus niger, modice vel dense rhizinosus, rhizinis P+ pale orange, norstictic, stictic, and usnic acids nigris, nitidis, simplicibus vel sparse ramosis. Apo- with or without atranorin. thecia numerosa (frequencia 100~o),adnata, usque HABIThT.-on rocks (granite, SaIldStOlle), rarel\ ad 2 mm diametro, amphithecio coronato, basin on trees, in open forests at low elevation. 24 SXIITHSONIAN CONTRIBUTIONS TO BOTANY

DIsTRrBuTIoN.-Australia and Tasmania, SPECIMENSExAMINED.-Sabah: Hale 29069, hleijer B548a REMARKS.-ThiS well-known lichen has been col- (BO, US). lected only in eastern Australia and Tasmania, where it appears to be common. It was elegantly 12. Relicina malesinna illustrated in color by Laurer (1827), yet Knight, not apparently having seen Laurer’s publication, FIGURE14f described it as a new species, Parmelia sphaero- Relicina inalesiana (Hale) Hale, 1974:484. spora. Knight then sent a specimen to Nylander, Parmelia malesiana Hale, 1965:203 [type-collection: Florida who immediately published a new name, P. insin- logging area, about 30 km southeast of Butuan City, uata, since Knight’s epithet was a homonym of Agusan Prov., Philippines, Hale 25370 (US, holotype; Nylander’s own P. sphaeospora published in 1859. TNS, UPS, isotypes)]. Vainio (1890) comprehended the species correct- Thallus adnate on bark, 6-8 cm broad; lobes ly, but Asahina’s (1934) reports from Japan are sublinear-elongate, 1.5-2.5 mm wide; bulbate cilia Relicina sydneyensis, the isidiate morph of R. gradually inflated; upper surface plane to rugulose, limbata. faintly maculate; medulla white, very rarely in SPECIMENSExAMINED.-Australia: Queensland, Bailey 165 part orange red; lower surface dark brown or (BM), Hartmann (US); New South Wales, Chase (US), Du blackening, densely rhizinate, the rhizines black, Rietz 60b, 4025 (UPS, US), Home (BM), Moore in Krypto- simple to squarrosely branched. Apothecia numer- gamae Vindobonensis Exsiccntae 570 (BM, H, LD, NY, UPS, US), Weber and hlcVean 47322, 49914 (COLO, US), Wilson ous (frequency 80y0), adnate, ecoronate; spores 8, (TNS), in Lichenes Rariores Exsiccati 34 (BPI, NY); Tas- 2-4 X 3-7 pm, medulla below the hymenium rare- mania, Brown (BM), Degelius A-368 (US). ly orange red. CHEMISTRY.-Medulla K--, c-, KC-, P-k red, usnic and fumarprotocetraric acids and if pig- 11. Relicina luteoviridis mented an unidentified K + purple anthraquinone. FIGURE14e HABITAT.-On trunk and branches of diptero- Relicina luteoviridis (Kurokawa) Hale, 1974:484. carps and other trees in rain forest at 150-600 m Parmelia luteoviridis Kurokawa in Hale and Kurokawa, 1964: elevation. 144 [type-collection: B. Papan, Beu territory, E. Kutai, DISTRIBUTIoN.-TaiWan, Philippines, Sabah, and Borneo, Meijer B1948 (BO, holotype; US, isotype)]. New Guinea. REMARKS.-The lower surface of this species is Thallus adnate on bark, fragile, 5-8 cm in diam- generally black but may be in part dark brown. eter; lobes sublinear-elongate, 1.5-3 mm wide; The rhizines when fully branched appear to be upper surface plane, faintly maculate, lobulate, squarrose, black, rather thin, and shiny. It is some- the lobules large, dorsiventral, marginally bulbate- what related, then, to species like R. ramosissima, ciliate, procumbent, often dichotomously branched which has a consistently pale brown lower surface (Figure 14e); cortex columnar (Figure 5b); bulbate and pale agglutinated rhizines, succinprotocetraric cilia long and tapered; lower surface black, densely acid in addition to fumarprotocetraric acid, and rhizinate, the rhizines black, simple or sparsely smaller spores. It is also chemically related to R. branched. Apothecia not seen. stiblimbata, which contains both fumarprotocetra- CHEMIsTRY.-Medulla K--, C+ rose, KC+ red, ric and succinprotocetraric acids. P-, gyrophoric and usnic acids with or without a tranorin. SPECIMENSExAMINED.-NeW Guinea: Hollandia, Royen and HABITATS.-On upper branches of evergreen trees Sleumer 6232 (L, US). See Hale (1965:203) for additional in rain forest at 600-1400 m elevation. records for Taiwan, Philippines, and Sabah. DIs”RIsuTIoN.-Sabah. REMARKs.-Relicina luteoviridis can be recog- 13. Relicina planiuscula nized by the large lobules and the unique chem- FIGURE15a istry. It is the only species containing gyrophoric acid. Only three collections are known, all from Reliciiza planiuscula (Kurokawa) Hale, 1974:484. Borneo. Parmelia planiuscula Kurokawa in Hale and Kurokawa, 1964: NUMBER 26 25

144 [type-collection: Kandang Badak, Java, Neeruoort 427 pm crassum, medulla 100-120 pm crassa, cortex in- (BO, holotype; US, isotype)]. ferior circa 12 pm crassus (Figure 2h); subtus pal- lide castaneus, dense rhizinosus, rhizinis pallidis, Thallus adnate on bark, coriaceous, 5-9 cm in dense ramosis atque agglutinatis. Apothecia adnata, diameter; lobes sublinear-elongate, 1-3 mm wide; usque ad 2 mm diametro, amphithecio crenato, upper surface plane, faintly maculate, sparsely to moderately isidiate, the isidia simple, less than coronato, basin retrorso-rhizinato, hymenio 70-80 0.7 mm high, frequently procumbent, rarely lobu- pm alto, sporis octonis, bicornutis, 2-3 X 11-13 late; upper cortex strongly columnar (Figure 5c); P. lower surface black, moderately rhizinate, the CHEMIsTRY.-Medulla K-, c -, P-k red, proto- rhizines simple, black. Apothecia rare (frequency cetraric and usnic acids. 11%), adnate, 1-2 mm in diameter, ecoronate, the HOLOTYPE.-virgin dipterocarp forest, Tungao amphithecium isidiate; spores 8, 4 X 5 pm. Logging Area, 40 km SE Butuan City, Agusan CHEMISTRY.-Medulla K -k yellowish, c -k faint Province, Philippines, elevation about 400 m, yellow orange, KC - , P + yellow, echinocarpic and M. E. Hale 25044a, July 1964 (US). usnic acids and a C+ unknown with or without HABITATS.-on canopy branches of dipterocarps atranorin. in rain forest at 100-400 m elevation. HABITATS.-on trunk and branches of Quercus, DrsTRIsuTIoN.-Philippines. Pinirr, and other trees in open forests and mossy REMARKS.-ThiS species represents the presump- forest at 1500-2200 m (at lower elevation in Japan tive nonisidiate morph of R. circumnodata. Both and Taiwan). contain protocetraric acid and have unusual bi- DIsTRlnUTI0N.-Japan, Taiwan, Philippines, Ma- cornute spores. Relicina precircumnodata is by far laya, Indonesia, and Sabah. the rarer of this species pair, occurring only at REMARKs.-This is the commonest species of three widely separate localities in the Philippines. Reliciria at higher elevations in Southeast Asia. SPECIMENSExAvINED.-Philippines: Cagayan, Hale 24418; The presumptive parent of this isidiate morph is Mountain, Hale 25826. R. flziorcsccns, which has a much more restricted distribution. While most specimens have sparse to moderate development of cylindrical isidia, others 15. Relicina ramosissima have procumbent lobulate isidia. The population FIGURE15c on Pin ?is in the northern Philippines is especially strongly isidiate-lobulate, and I had at first con- Relicifla ramosissima (Kurokawa) Hale, 1974:485. sidered it to be a separate species. Parmelia ramosissima Kurokawa in Hale and Kurokawa, 1964: 145 [type-collection: Tenimber Island, Moluccas, SPECIMENSExAhrmm.-Japan. Hyuga: Hale 29665. Philip- BuwaEda 4583 (BO, holotype; US, isotype)]. pines: Mountain, Hale 26088, 26089, 26102, 26113, 26117, 26173, 26195, 26806. Malaya: Pahang, Hale 29929, 29943,' Thallus adnate, corticolous, 3-10 cm in diam- 29944, 29945, 29946, 29949, 30481, 30497. Sabah: Hale 28006, eter; lobes sublinear-elongate, 0.7-2 mm wide; bul- 28040, 28063, 28234, 28250, 28257, 28372, 28400, 28508, 28546, 28580, 28581, 28603, 28700, 28815, 29035, 29168, 29250. In- bate cilia gradually inflated to gobose, becoming donesia: Java, Groenhart 2217 (L); Sumatra, Steenis 3895 apically branched; upper surface plane, maculate; PO). lower surface pale to dark brown, moderately rhizi- nate, the rhizines simple to densely branched and agglutinated, pale (Figure 8f). Apothecia common 14. Relicina precircumnodata, new species (frequency 93%), adnate, 1-2.5 mm in diameter, FIGURE15b ecoronate, often retrorsely rhizinate at the base; spores 8, 2 X 3 pm. Thallus adnatus, corticola, viridi-flavicans, 4-10 CHEMISTRY.-Medulla K - , c -, KC -k reddish, cm diametro, lobis sublinearibus, contiguis vel sep- P+ red, usnic acid, fumarprotocetraric acid, and aratis, 1-2 mm latis, margine dense bulbo-ciliatis, succinprotocetraric acid. bulbis globosis, sorediis isidiisque destitutis; cortex HAnITAT.-on trunk and canopy branches of superior circa 12 pm crassus, stratum gonidiale 10 dipterocarps in rain forest at 150-300 m elevation. 26 S.MITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE15.--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 Parmelia relicina Fries, 1825:283 [type-collection: Gaudichaud Indonesia. 54, Rawak (Pacific area) (UPS, lectotype; P. isolectotype)]. REMARKS.-This species has a wide distribution Thallus closely adnate to appressed on bark, 2-6 in the dipterocarp forests. It superficially resem- cm broad; lobes variable, short, contiguous and bles R. sublanea, which differs in producing pro- sublinear to separate and sublinear-elongate, 0.3- tocetraric acid, in having large spores (4-5 X 7-9 1.5 mm wide; bulbate cilia short but conspicuous, pm), and in occurring generally above 300-400 m strongly inflated, sometimes apically branched; elevation, higher than R. ramosissima. upper surface plane, continuous to faintly macu- SPECIMENS EXAMINED.-Philippines: Agusan, Hale 25245, late, without lobules or becoming marginally lobu- 25360. Malaya: Selangor, Hale 29923, 30062, 30074, 30075, late, isidia lacking; lower surface black, densely 30290, 30291, 30300. Sabah: Hale 30352, 30355, 30356, 30357, rhizinate, the rhizines black, simple to sparsely 30366. Indonesia: Java, Groenhart 8374 (BO). branched. Apothecia very common (frequency 9473, adnate, about 1 mm in diameter, the exciple 16. Relicina relicinella coronate, the base often retrorsely rhizinate; spores X FIGURE15d 8, 4-5 5-7 pm. CHEMISTRY.-Medulla K-, c-, KC-, P-, Relicina relicinella (Nylander) Hale, 1974:485. usnic acid and unidentified colorless substances. Parmelia relicinella Nylander 1885:215 [type-collection: San- HABITAT.-on trunk and canopy branches of tarem, Brazil, Spruce 136 (H, lectotype; BM, G, P, isolecto- dipterocarps and other trees in rain forest at 100- types)]. 400 m elevation. Thallus closely adnate on bark, 3-6 cm in diam- DIsTRIBUTIoN.-Phi~ippines, Malaya, and Indo- eter; lobes sublinear, 0.5-1.5 mm wide; bulbate nesia. cilia distinct, short; upper surface plane, continu- REMARKs.-The characteristic features of this ous; lower surface black, densely rhizinate, the widespread species are the narrow appressed lobes rhizines simple or sparsely branched. Apothecia and lack of lichen substances in the medulla. It is common (frequency loo%), adnate, 0.5-1.5 mm an extremely variable species, however, particularly in diameter, the disc concave to plane, the exciple in regard to production of marginal lobes. Some coronate; spores 16-32, 3-4 X 5-6 pm. specimens have no obvious lobules. Muller’s type- CHEMISTRY.-Medulla K-, c-, KC-, Pf red, specimen is somewhat lobulate. He described Par- usnic acid, fumarprotocetraric acid, and an un- melia relicinztla as being 2-3 times narrower than identified P+ compound. P. relicina Fries, but I doubt that he saw Fries’ HABITAT.-on tree trunks and branches in rain type. The excellent material of P. relicina in Paris forest at 100-200 m elevation. has very narrow lobes. This specimen does lack DIsTRIBuTIoY.-Co~ombia and Brazil. lobules. Its chemistry is still undecided since, al- REMARKs.-The most unusual feature of this though it is negative with color tests, a dull, high species is the multispored asci, recognized by unidentified spot appears on TLC plates. Similar Nylander (1885) in his original description. It is but not identical spots were discovered in several very similar externally to R. eximbricata but seems of the Philippine specimens. to occur only in virgin rain forest. It may be a Asahina (1934) reported Parmelia relicina from common species in the vast lowland rain forests of southern Japan. I have not seen this material but South America, but no lichenologists have yet had suspect that it is Relicina echinocarpa. an opportunity to visit these habitats. SPECIMENS EXAMINED.-Philippines: Mountain, Hale 25849, SPECIMENSEXAhmxED.-Colombia: Santander, Nee and Mori 25871: Cagayan, Hale 24402, 24416, 25604, 25638, 25652, 3766. Brazil: Amazonas, Spruce 58 (BM, US). 25657, 25658, 25737; Cavite, Hale 26836; Sorsogon, Elmer 16031 (F, US): Agusan, Hale 24996, 25011, 25659; Surigao del 17. Relicina relicinula, new combination Sur, Hale 24641, 24698, 24700; Zamboanga del Sur, Hale 24521, 24704, 24709, 24733, 24738, 24748, 24807, 24826, 24827, FIGURE15e 24861, 25159, 25313. Malaya: Selangor, Hale 29925, 30281. Parmelia relicinula Muller Argau, 1882:317 [type-collection: Java: Groenhart 1702 (L). Thursday Island: Hartmann in Java, Junghuhn 3728 (G, lectotype)]. Lichenotheca Universalis 159 (S, US). 28 SMITHSONIAN CONTRIBUTIONS TO BOTANY

18. Relicina samoensis black, moderately rhizinate, the rhizines simple. Apothecia unknown. FIGURE15f CHEMISTRY.-Medulla K- yellow, C- or C+ Relicina samoensis (Zahlbruckner) Hale, 1974:485. faint orange, KC-, P+ orange, usnic and echino- Parmelia samoensis Zahlbruckner, 1908:272 [type-collection: carpic acids and a C+ unknown. Malifa, Upolu, Rechinger 5005 (W, lectotype)]. HABITAT.-on trunk and branches of QUeYCUS and other trees in rain forest at 1300-1600 m Thallus adnate to closely adnate, corticolous, 4-8 elevation. cm in diameter; lobes rather short, sublinear, 0.3-1 DIsTRIBuTIoN.-hfa~aya, Philippines, Sabah, and mm wide; bulbate cilia conspicuous, globose, up Indonesia. to 1 mm long; upper surface plane, continuous; REMARKS-The distinguishing feature of this lower surface black, densely rhizinate, the rhizines species is the distinct marginal and lamina1 10- black, simple or sparsely branched. Apothecia ad- bules (Figure 1b). These lobules are dorsiventral, nate, very common (frequency 100’%), 1-3 mm in simple to forked, and sometimes subascending. Bul- diameter, the exciple coronate, the base retrorsely bate cilia are rarely developed on the margins in rhizinate; spores 8, 5-6 X 6-7 pm. contrast, for example, to the abundant production CHEMISTRY.-hfedd1a K+ yellow, C faint + of bulbils in R. luteoviridis. There is some inter- orange, KC-, P+ yellow orange, usnic and echino- gradation with echinocarpic-acid-containing speci- carpic acids and a C+ unknown. mens of R. ampkitkrix, which have strongly lobu- HABITAT.-on coconut palm and other trees in late isidia. On the average R. ampkitkrix would disturbed areas at 100-700 m elevation. have much narrower lobes than R. sckirospatha DIsTRrBUTION.-~hi~ippines, Java, New Cab and some trace of isidial initials. In addition R. donia, Samoa, and Society Islands. ampkitkrix is rare at the higher elevations where REMARKs.-Relicina samoensis is a lowland spe- R. sckizospatkn normally occurs. In addition, al- cies occurring in secondary forests. The lobes vary though R. sckizospatka is sterile, I suspect that the considerably in size but are usually narrow and apothecia, when discovered, will be ecoronate. closely adnate. This is the only species with echino- carpic acid that has coronate apothecia. SPECIMENSEXAMIh.ED.-Philippines: Bukidnon, Sulit 14806 (US). Malaya: Pahang, Hale 30089, 30142, 30210, 30216, 30221, SPECIMENSExAM1sED.-Philippines: Quezon, Hale 26955. 30222, 30223, 30486, 30494. Sabah: Hale 28156, 28157, 28221, Indonesia: Halmahera, Croenhart 8343 (BO): Java, Bredijn 29026, 29037, 29117, 29285. Indonesia: Java, Groenhart 2219 1099 (BO), Palmer 614 (US). Samoa: Reinecke 22 p. p. (BO, (L, US), Ooststroom 14198, 14300 (L, US). A record from US). New Caledonia: Hill 11590 (BM). Society Islands: Moore L3 (US). Taiwan by Kurokawa (1965:267), which I have not been allowed to examine, is probably R. amphithrix.

19. Relicina schizospatha 20. Relicina subabstrusa FIGURE16a FIGURE16b Relicina schizospatha (Kurokawa) Hale, 1974:485. Parmelia schizospatha Kurokawa in Hale and Kurokawa, Relicina subabstrusa (Gyelnik) Hale, 1974:485. 1964:146 [type-collection: Gegerbentang, Java, Neeruoort Parmelia subabstrusa Gyelnik, 1931:288 [based on P. abstrusa 1062 (BO, holotype; US, isotype)]. f. laevigata Lynge]. Parmelia limbata f. endococcinea Muller Argau, 1887:318 Thallus adnate, corticolous, rather fragile, 3-6 [type-collection: Russell River, Australia, Sayer 19 (G, cm in diameter; lobes short, sublinear, 1-2 mm lectotype)]. wide; hulhate cilia not conspicuous or dense; upper Parmelia abstrusa f. laevigata Lynge, 1914: 147 [type-collection: Bocca da Serra, Serra da Chapada, Mato Grosso, Brazil, surface plane, faintly maculate, becoming more or Malme (S, lectotype)]. less densely lobulate along the margins and sur- Parmelia endococcinea (Muller Argau) Gyelnik, 1931:288. face, the lobules dorsiventral, procumbent to sub- Parmelia decaryana Gyelnik, 1934: 153 [type-collection: Tsant- ascending, sometimes dactyloid branched, rarely sani, Anjouan, Comoro Island, Decary (BP, holotype)]. with marginal hulhils (Figure 16); lower surface Parmelia kilauae f. laevigata (Lynge) Gyelnik, 1935:37. NUMBER 26 29

FIGURE16.-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 (Kim- kawa 1872); f, R. sydneyensis (Weber 49048). (Scale in mm.) 30 SMITHSONJAN CONTRIBUTIONS TO BOTANY

Thallus adnate on bark, 4-8 cm broad; lobes simplicibus vel sparse ramosis. Apothecia adnata, sublinear, 0.5-2 mm wide; bulbate cilia numerous, 0.5-1 .O mm diametro, amphithecio coronato, basin strongly inflated, to 1 mm long; upper surface retrorso-rhizinato, disco carneo, imperforato, sporis plane, continuous; lower surface black, densely octonis, 3 X 4 ym. rhizinate, the rhizines black, simple (Figure 8c). CHEMISTRY.-~edUlla negative with color tests, Apothecia very common (frequency 83y0), adnate, usnic and caperatic acids. 1-3 mm in diameter, the exciple strongly coronate; HOLOTYPE.-virgin dipterocarp forest, Ulu Gom- spores 8, 4 X 7-8 pm. bak Forest Reserve, 15 km NE of Kuala Lumpur, CHEiWIsTRY.-Medulla K-k yellow turning red, elevation about 300 m, M. E. Hale 30054, 5 March C-, KC-, P+ orange, norstictic and usnic acids 1965 (US). with or without salazinic acid. REMARKS.-This species was first identified as HABITAT.-on trunk and branches of diptero- R. connivens. It differs primarily in having coro- carps in Old World rain forest and in secondary nate apothecia and very small spores and in con- forest at sea level to 850 m elevation and on trees taining caperatic acid rather than protolichesteri- at mid elevations in the New World. nic acid. It is known only from the type collection. DIsTRIBUTION.-Taiwan, Philippines, Sarawak, Australia, Comoro Islands, Brazil, and Paraguay. REMARKS.-Relicina subabstrusa has the broad- 22. Relicin,a sublanea est geographic range of any species in the genus but is commonly collected only in the Philippines. FIGURE16d As with R. abstrzisa, its presumptive isidiate morph, Relicina sublanea (Kurokawa) Hale, 1974:485. the Old Miorld population lacks or has only traces Parmelia sublanea Kurokawa in Hale and Kurokawa, 1964: of salazinic acid, whereas salazinic is produced in 146 [type-collection: Halmahera Island, Indonesia, Groen- heavy concentration in the New World. Muller’s kardt 8409 (BO, holotype; US, isotype)]. specimen of f. endococcinea is red because of the decomposition of norstictic acid. Thallus adnate on bark, coriaceous, 6-10 cm in diameter; lobes sublinear-elongate, 1-2 mm wide; SPECIMENS EXAhrINED.-Phi~ippines: Mountain, Hale 25760, bulbate cilia strongly inflated; upper surface more 25762, 25772, 25581, 25797, 25800, 25801, 25812, 25817, 25838, or less convex, continuous; lower surface pale 25858, 25863, 25879, 25898, 25901, 25802a, 25913, 26506; Cagayan, Hale 25608, 25654, 25661, 25751; Cavite, Hale 26811, brown, densely rhizinate, the rhizines densely 26828; Polillo, Leiberg 532 (US); Negros Occidental, Hale branched, and agglutinated (Figure 8g). Apothecia 26427, 26444, 26449, 26473, 26494, 26494a, 26510, 26519, 26564, common (frequency 607L), adnate, 1-4.5 mm in 26581, 26626, 26632, 26693, 26696; Agusan, Hale 24460, 24474, diameter, ecoronate; spores 8, 4-5 X 7-9 ym. 24984, 25003, 25014, 25029, 25033, 25048; Basilan, Hale 24935, CHEMISTRY.-hfedUlki K-, C--, KC+ rose, P+ 24949, 25141, 25333. Sarawak: Hale 29991. Paraguay: Balansa orange red, usnic and protocetraric acids. (H). See Kurokawa (19633267) for records from Taiwan. HABITAT.-on trunk and canopy branches of dipterocarps in rain forest at 300-850 m elevation. 21. Relicina subconniuens, new species DIsTRIBuTIoN.-Southern Philippines, Malaya, and Indonesia. FIGURE16e REMARKS.-This is another typical dipterocarp forest lichen characterized by pale agglutinated Thallus adnatus, corticola, viridi-flavicans, circa rhizines and protocetraric acid. Its relation to 10 cm latus, lobis sublinearibus, contiguis, sparse R. rnmosissimn is discussed under that species. The lobulatis, margine dense bulbociliatis, bulbis aetate only other nonisidiate species with protocetraric inflatis, globosis, isidiis sorediisque destitutis; cor- acid, R. precirczimnodata, has coronate apothecia. tex superior 10-12 ym crassus, stratum gonidiale SPECIMENS ExAMINED.-Philippines: Negros Occidental, circa 12 pm crassum, medulla 60-80 ym crassa, Hale 26459, 26578, 26650, 26657; Surigao del Sur, Hale 24635, cortex inferior 16-18 pm crassus (Figure 3f); sub- 24655; Zarnboanga del Sur, Hale 24793, 23310; Basilan, 24920, tus niger, dense rhizinosus, rhizines nigris, nitidis, 24922. Malaya: Selangor, Hale 30280. NUMBER 26 31

23. Relicina sublimbata Thallus adnate on bark or rocks, 3-8 cm in diameter; lobes sublinear to sublinear-elongate, FIGURE16e contiguous, 0.8-1.5 mm wide; bulbate cilia tapered Relicina sublimbata (Nylander) Hale, 1974:485. to strongly inflated; upper surface plane, faintly Purmeliu sublimbata Nylander, 1885:615 [type-collection: maculate, irregularly cracked with age, isidiate, the Yomah in Pegu, Burma, Brandis 595 (M, lectotype; H, isolectotype)]. isidia simple, to 0.8 mm high; lower surface brown to dark brown, moderately rhizinate, the rhizines Thallus adnate on bark or rock, coriaceous, 4-8 brown or black, mostly simple. Apothecia rare, cm in diameter; lobes sublinear-elongate, 1-2.5 mm adnate, 1.5-3.5 mm in diameter, ecoronate, the base wide; bulbate cilia coarse, inflated, to 1 mm long; retrorsely rhizinate; spores 8, 4-5 X 7-8 pm. upper surface plane, continuous; lower surface CHEMISTRY.-Medulla K + yellow turning red, black, densely rhizinate, the rhizines black, simple C-, KC-, P+ orange, usnic, norstictic, and to sparsely branched. Apothecia adnate, 1-2 mm stictic acids with or without atranorin. in diameter, ecoronate; spores 8, 4-5 X 5-6 pm. HABITAT.-on bark of trees and more rarely on CHEMISTRY.-Medulla K + reddish, c -, KC - , rocks in open forests at low elevation. P + orange red, usnic acid, fumarprotocetraric acid, DIsrrcIsuTIoN.-Japan, Taiwan, Java, and Aus- and succinprotocetrarir acid. tralia. HABITAT.-& tree trunks and rocks at mid ele- KEMARKs.-Muller Argau recognized the close vation (1300-1400 m in Thailand). relation between this and R. limbata; R. sydney- DIsTRIsuTroN.-Burma and Thailand. ensis would appear to be the isidiate morph of REMARKs.-Nylander (1885) separated this spe- R. limbata. It has identical chemistry but seems cies from R. limbata because of the smaller spores to occur more frequently on trees and, of course, (4.5 X 6-8 pm) and whitish color. The two species to have a broader geographic range. Chromato- are, in fact, totally unrelated, Relicina sublimbata graphic profiles of the types of Parmelia limbata f. is actually very close to R. malesiana, which pro- isidiosa and P. subturgida are identical. Hale and duces fumarprotocetraric acid alone, has more Kurokawa (1964) proposed the name Parmelia branched rhizines and smaller spores, and does not tumescens but later discovered that the type of occur on mainland Asia. Relicina sublimbata P. sydneyensis Gyelnik is actually sparsely isidiate, seems to be restricted to the Burma-Thailand re- not nonisidiate. gion but little is known of its full range there. SPECIMENS EXAMINED.-Japan: IZU, Kurokawa 58622 (Us). SPECIMENSExAMINm.-Thailand: Kurokawa 1872 (TNS, Indonesia: Java, Neruoort 5874 (BO). Australia: New South US). Wales, Weber and McVean L-49048 (COLO, US); Queens- land, Du Rietz 4234:l (UPS, US). See Kurokawa (1965:268) 24. Relicina sydneyensb for records from Taiwan.

FIGUREI6f Nomen Inquirendum: Parmelia nigrociliata Relicina sydneyensis (Gyelnik) Hale, 1974:485. Parmelia sydneyensis Gyelnik, 1938:292 [type-collection: Moss- Parmelia nigrociliata Hillmann, 1940:39 [type-collection: man’s Bay, Sydney, Australia, Cheel (BP, holotype)]. Mutautu, Samoa, Vaupel (destroyed at B); (not Parmelia Parmelia limbata Laurer f. isidiosn Muller Argau, 1887:59 nigrociliata Bouly de Lesdain ex anno 1933)l. [type-collection: Richmond River, Australia, Hodgkinson (G, lectotype)]. This species cannot be typified since the type Purmelia tumescens Hale and Kurokawa, 1964:147 [based on (and only known) collection was lost at Berlin. P. limbata f. isidiosa Muller Argau]. The description and remarks by Hillmann, how- Parmelia subturgida Kurokawa, 1965:268 [type-collection: Bandagamori, Prov. Tosa, Japan, Makino (TNS, holotype; ever, leave little doubt that it is synonymous with US, isotype)]. Relicina samoensis (Zahlbruckner) Hale. Literature Cited Asahina, Y. 1975. A Revision of the Lichen Genus Hypotrachyna in 1934. Uber Parmelia abstrusa Vain., P. limbata Laur. und Tropical America. Smithsonian Contributions to P. relicina Fr. aus Japan. The Journal of Japanese Botany, 25:l-73. Botany, 10:299-304. Hale, M. E., and S. Kurokawa 1951. Lichenes Japoniae novae vel minus cognitae (3). 1964. Studies on Parmelia subgenus Parmelia. Contribu- The Journal of Japanese Botany, 26:193-198. tions from the United States National Herbarium, Crombie, J. M. 36:121-191. 1883. On a Collection of Exotic Lichens Made in Eastern Hillman, J. Asia by the Late Dr. A. C. Maingay. Journal of the 1940. Neue oder wenig bekannte Flechten aus aller Welt, Linnean Society, Botany, 20:48-69. 111. Fedde Repertorium, 48:34-40. Knight, C. Culberson, C. F., and M. E. Hale, Jr. 1882. Contribution to the Lichenographia of New South 1973. Chemical and Morphological Evolution in Parmelia IVales. Transactions of the Linnean Society of Lon- sect. Hypotrachyna: Product of Ancient Hybridiza- don, Botany, new series, 2:37-51. tion? Brittonia, 25:162-173. Kurokawa, S. Culberson, W. L. 1965. Revision of Series Relicina of the Genus Parmelia 1973. The Parmelia perforata Group: Niche Character- in Japan and Taiwan. The Japanese Journal of istics of Chemical Races, Speciation by Parallel Botany, 40: 264-269. Evolution, and a New Taxonomy. Bryologist, 76: Laurer, J. F. 20-29. 1827. Sieber’sche Lichenen. Linnaea, 238-46. Fries, E Lynge, B. 1825. Systema Orbis vegetabilis. Part 1, 374 pages. Lund. 1914. Die Flechten der ersten Regnellschen Expedition. Gyelnik, V. Die Gattungen Pseudoparmelia gen. nov. und Par- 1931. Additamenta ad cognitionem Parmeliarum, 11. melia Ach. Arkiv for Botanik, 13 (13):1-172. Fedde Repertorium, 29:273-291. Muller Argau, J. 1934. Additamenta ad cognitionem Parmeliarum, V. 1882. Lichenologische Beitrage, XX. Flora, 65:316322. Fedde Repertorium, 36: 151-156. 1887. Lichenologische Beitrage, XXVI. Flora, 70:316-320. 1935. Revisio Typorum ab auctoribus variis descriptorum Nylander, W. 1. Annales Musei Nationalis Hungarici, 29: 1-54. 1859. Lichenes in regionibus exoticis quibusdam vigentes, 1938. Additamenta ab cognitionem Parmeliarum, 8. An- cum synopticis enumerationibus. Annales Sciences nales Mycologici, 36:267-294. Naturelles Botanique, series 4, 11:205-264. Hale, M. E. Jr. 1885. Parmeliae exoticae novae. Flora, 68:605-615. 1965a. A Monograph of Parmelia Subgenus Amphigymnia. 1886. Lichenes nonnulli Australiensis. Flora, 69:323-328. Contributions from the United States National Her- Rechinger, K. H. barium, 36: 193-358. 1907. Botanische und zoologische Ergebniss einer wissen- 1965b3. Six New Species of Parrnelia from Southeast Asia. schaftlichen Forschungsreise nach der Samoainseln, The Japanese Journal of Botany, 40:199-205. dem Neuguinea-Archipel und den Solomonsinseln, 1971. Parmelia afluens, a New Species of Lichen in Sub- IV: Die Flechten von A. Zahlbruckner. Denkschrift genus Amphigymnia with a Yellow Medulla. Phy- Akademie der Wissenschaften in Wien, 81:250-292. tologin, 22:141,142. Vainio, E. A. 1973. Fine Structure of the Cortex in the Lichen Family 1890. Ptude sur la classification naturelle et la morphol- Parmeliaceae Viewed with the Scanningelectron ogie des Lichens du Bresil. Acta Societas Fauna et Microscope. Smithsonian Contributions to Botany, Flora Fennica, 7 (7):l-247. 10: 1-92. Zollinger, H. 1974. Bulbothrix, Parmelina, Relicina, and Xanthopar- 1854. Systematisches Verzeichniss der im indischen Archi- me&, Four New Genera in the Parmeliaceae pel in den Jahren 1842-1848 gesammetten sowie der (Lichenes). Phytologia, 28:479-490. aus Japan empfangenen Pflanzen. 160 pages. Zurich.

* U.S. GOVERNMENT PRINTING OFFICE: 1975-58S-80/082S 32