A Monograph of the Lichen Genus Relicina (Parmeliaceae)

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

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