lll MllllHHl THE LICHEN GENUS RAMALITNA ACH9 , m THE WEST INDIES WITH NOTES : ‘ 0N ITS ROLE IN THE VEGETATION . . . “ , OFPUERTO RICO ThesnéfoTtheDegrée OTP'h. D MICHIGAN STATE UNIVERSITY - ; » ISMAEL LANDRON COTNCEPCION . ' 1972 A * W J- LIBRARY ”7’ Michigan Sue: r. University This is to certify that the 0%,. I:- thesis entitled THE LICHEN GENUS RAMALINA ACH. IN THE WEST INDIES WITH NOTES ON ITS ROLE IN THE presented by I smael L andrén’ Concepci 6n has been accepted towards fulfillment of the requirements for Ph . D. degree in Bot any % fl. 6 62: Major professor Date M 017639 . L _. BINDING av ‘-‘ HUM} & SflNS‘ \fiEflGK BENCH! 15-1“, , , — ETL‘QRASXHD'N-T : I .: LICHEN GENUS I WITH N07 VEGET Ismae ABSTRACT THE LICHEN GENUS RAMALINA ACH. IN THE WEST INDIES WITH NOTES ON ITS ROLE IN THE VEGETATION 0F PUERTO RICO By Ismael Landrén Concepcion The purpose of this study was to investigate the hchmigenus Ramalina Ach. in the West Indies and assess itsrple in the vegetation of Puerto Rico. The study is basaion collections made during the summer of 1967 and Um winter of 1968—1969 throughout the island, supplemented by herbarium material. The vegetation of the West Indies, Central America, hmatén, Mexico, Venezuela and Florida has been reviewed. Hm physiography, geology, climate and vegetation of Puerto Mco is treated in more detail and maps have been prepared. Thelichen vegetation in Puerto Rico is described within sevmlvegetation types: littoral, lowland rainforest, seasonal-evergreen forest, semi-deciduous forest, lower mon— tmw rainforest, montane forest and montane scrub. The distribution patterns of the species of Ramalina Wiflfln Puerto Rico are discussed and categorized as well as LA 2:12:75 :5 distril :2 "Es: Indies. The taxonomic 1125 a key to twen :uized several Chen sensed on analyses gin tests, and ti 2:29: {1'} lichen 5:2 species includ 5.3.5, chemical r ;:;s::ibution. I smael Landrén the patterns of distribution of species of Ramalina through- out the West Indies. The taxonomic treatment of the West Indian material includes a key to twenty—three (23) species, some of which contained several chemical variants. The chemical variants were based on analyses by spot tests, microchemical crystal— lization tests, and thin-layer chromatography. A total of seventeen (17) lichen substances was demonstrated. Discussion of the species includes notes on nomenclature, taxonomy, diagnosis, chemical reactions, lichen substances, ecology and distribution . EUflfiGHUSl WITH NO] VEGETJ lsmae THE LICHEN GENUS RAMALINA ACH. IN THE WEST INDIES WITH NOTES ON ITS ROLE IN THE VEGETATION OF PUERTO RICO By Ismael Landr6n~Concepci6n A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Botany and Plant Pathology College of Natural Science 1972 .n iedicat it" 50115 c In dedication to my wife Myriam and my sons Ismael Angel and Jesus Manuel. ii A1 Iuish to 8X stony Ml“ PrOf .2ng to the beaut than and tolerai ifriendship and C01 :fnthe use of hi in collections. 1 am deeply lztee, Dr. Rolli 33. iantlon and 1 {32511915 on and c ACKNOWLEDGMENTS I wish to express my deep appreciation and grat- itude to my major professor Dr. Henry A. Imshaug for intro— ducing me to the beautiful world of lichens, for guiding me with humor and tolerance toward the completion of this study, for friendship and concern demonstrated to me and my family and for the use of his personal library and West Indian lichen collections. I am deeply indebted to the members of my graduate committee, Dr. Rollin H. Baker, Dr. Everett S. Beneke, Dr. John E. Cantlon and Dr. William B. Drew for their many suggestions on and criticisms of the manuscript. Thanks are extended to the individuals and institu- tions listed below for the loan of specimens: Dr. I. Mackenzie Lamb (FH), Dr. Howard A. Crum (MICH), Dr. William Louis Culberson (DUKE) and Dr. Mason E. Hale (US). Acknowledgment is made to my fellow graduate students for providing an atmosphere of friendliness and cooperation toward me. Special thanks are due Mr. Ronald M. Taylor for aiding in the preparation of plates. I would like to extend special thanks to the University of Puerto Rico for financial assistance provided during my stay and Michigan State University. Finally, I express my sincere appreciation to my wife and sons for the many sacrifices they have made to enable me to complete my studies. It is to them that this dissertation is dedicated. iii i IEIIEDQENTS . - - SUITABLES . - - ISIIIGURES . - . :.?E?IAIES 3'. TE DIEXDICES F‘Lv mm '.. INTRODUCTION 1- CHER-ll DESC DESI INDIES Florida . The Baham: The Greatl Cuba TABLE OF CONTENTS Page ACKNOWLEDGMENTS . iii LIST OF TABLES . viii LIST OF FIGURES X LIST OF PLATES xii LIST OF APPENDICES . . . xiv Chapter I. INTRODUCTION 1 II. GENERAL DESCRIPTION AND VEGETATION OF THE WEST INDIES AND ADJACENT AREAS . . . . . . . . . 3 Florida . . 3 The Bahamas . . 4 The Greater Antilles 5 Cuba . S Hispaniola 7 Jamaica . . 3 The Lesser Antilles . . . . 9 The Continental Islands of Trinidad and Tobago . . . 10 Central America . 12 Panama . 12 Costa Rica 13 Honduras 13 Yucatan 14 Mexico . 15 Venezuela . 16 III. PUERTO RICO 18 General Description 18 Physiography l8 Geology 20 Climate 22 iv 6 i an Vegetation ‘ Hi5t0r1ca Schemes Dansereau System The Lit The Low Raini swam} Lake The Se: Semi-D The L0 Nontan Palm Colo N055 Mont The MC Distribi Summary . BIOGEOGRTPH Florigtic IntTOCIu West In Endemic COnt‘lne Distribu< Distribu Sum”). Chapter IV. Vegetation . . Historical Review of Classification Schemes . . . Dansereau' s Vegetation Classification System The Littoral Zone . . . The Lowland Rainforest Zone Rainforest Swamp forest Lake and River Ecosystems . . . . The Seasonal Evergreen Forest Zone Semi- Deciduous Forest Zone The Lower Montane Rainforest Zone . Montane Forest Zone Palm Type Forest . . Colorado Type Forest Moss Type Forest Montane Maquis . . . The Montane Scrub Zone Distribution of Ramalina Summary BIOGEOGRAPHICAL AFFINITIES OF THE WEST INDIAN FLORA Floristic Elements of Vascular Plants Introduction West Indian Element Endemic Element . . . Continental Element . Cosmopolitan Element Distribution of Animals in the West Indies Distribution of the Lichen Genus Ramalina . Summary THE GENUS RAMALINA ACH. History of the Genus Morphological and Anatomical Studies. Methods Thallus Soralia . Rhizines Cilia . Tissues Cortex Medulla . . Algal Layer Page 23 23 25 25 28 28 35 37 38 41 44 48 52 55 57 61 65 67 71 76 76 76 77 79 82 87 87 88 91 93 93 101 101 102 105 107 107 107 108 109 110 'mler Apotheci Hymen: Spore Subhy Pycnidi Chemical St Introduct Methods a Color I Microcl Thin L: lichen ST Atrano Capera Crypto Figu DiTari Xorsti Protoc Psoron Rmnh Salazf Sekik; Usnic Subst Unide Chapter Page Apothecia . . . . . . . . . . . . . . . . . 110 Hymenium . . . . . . . . . . . . . . . . 111 Spores . . . . . . . . . . . . . . . . . 112 Subhymenial Tissues . . . . . . . . . . . 113 Pycnidia . . . . . . . . . . . . . . . . . 113 Chemical Studies‘ . . . . . . . . . . . . . . . 114 IntroduCtion . . . . . . . . . . . . . . . . 114 Methods and Techniques . . . . . . . . . . . 117 Color Reactions . . . . . . . . . . . . . . 117 Microchemical Tests . . . . . . . . . . . . 119 Thin Layer Chromatography . . . . . . . . . 121 Lichen Substances in Ramalina . . . . . . . . 124 Atranorin (Plate 1, Figure 1) . . . . . . . 125 Caperatic Acid (Plate I, Figure 2) . . . . 130 Cryptochlorophaeic Acid (Plate 1, Figure 3) . . . . . . . . . . . . . . . . 131 Divaricatic Acid (Plate 11, Figure 4) . . . 134 Norstictic Acid (Plate 11, Figure 5) . . . 135 Protocetraric Acid (Plate 11, Figure 6) . . 136 Psoromic Acid (Plate III, Figure 7) . . . . 136 Ramalinolic Acid (Plate 111, Figure 8) . . 139 Salazinic Acid (Plate III, Figure 9) . . . 140 Sekikaic Acid (Plate IV, Figure 10) . . . . 143 Usnic Acid (Plate IV, Figure 11) . . . . . 144 Substance H (Plate IV, Figure 12) . . . . . 145 Unidentified Substance A (Plate V, Figure 13) . . . . . . . . . . . . . . . 148 Unidentified Substanc B (Plate V, Figure 14) . . . . . . . . . . . . . . . 149 Unidentified Substanc C (Plate V, Figure 15) . . . . . . . . . . . . . . . 150 Unidentified Substance D . . . . . ~ - . 150 Role of Chemical Substances in the Taxonomy of Ramalina 153 Chemogenesis . . . . . . . . . . 153 I Distribution of Lichen Substances ' in Ramalina . . . . . . . . . . . . . . . 155 i Summary . . . . . . . . . . . . . . . 157 Taxonomic Treatment . . . . . . . . . . . . . . 128 i Sub—generic Categories . . ‘.' . . . ... ' Artificial Key to Ramalina Spec1es Occurring 161 in the West Indies and Florida . . . . . . 163 Section Ramalina . . . . . . . . . . . . . . 163 Subsection Myelopoea (Va1n.) Zahlbr. . . . 163 Series Teretiusculae (Va1n.) Zahlbr. . . 1. Ramalina attenuata (Pers.) 163 R. H. Howe . . . . . . . . . . . . 166 2. Ramalina dendrisc01des Nyl. vi SubseCt Smmary - 75.1.de CITED Chapter LITERATURE APPENDICES 3. Ramalina furcellata (Mont. ) Zahlbr. 4. Ramalina EETEHTEEE—(B. de Lesd. ) comB. nov. . . 5. Ramalina monta nei De Not. . . 6. REHEITHE coc learis Zahlbr. . . 7. RamaIina rac1Iis (Pers.) Nyl. . 8. 'RamaIina su5as erata Nyl. . . .. Series ComprelssTEEEfiIEE—Tvain. ) Zahbr. 9. Ramalina com lanata (Sw.) Ach. 10. 'W ama 1na sore 1ant a Nyl. 11. RamaI1na 1e tos erma Nyl. . 12. RamaI1na straminea Pers. ) Ach. 13. RamaIlna cam tos ora Nyl. . 14. Ramalina eruV1ana Ach. . 15.RamaI1na Tarinacea (L. ) Ach. 16. RamaIina Bistorta Nyl. . . l7. Rama[1na cumanensis Fée . . . . . 18. Rama11nasu5peI1uc1da Mull. Arg. 19. Ramallna anceps Nyl. . . . 20. Ramalina perance s Nyl. 21. Ramal1na en r01 es Nyl. . 22. Ramalina usnea (L .i R. H. Howe Subsection F15tu1ar1a (Vain. ) Zahlbr. . . 23.Rama1ina inflata (Hook. f. G Tayl. ) Summary . . . . . . . . . . . . . . . CITED vii Page 171 175 180 186 189 195 203 203 220 223 226 228 230 235 240 243 246 251 256 261 266 275 275 306 311 327 1. Distribution 0: oootane forest 2. Physical chara chemontane f0 3. iercent of end 5. Percent of end in the Lesser E. Aromatic aldel reaction with ?- Ieosides not P‘Phenylenedi color reactio and calcium h Table 1. 2. 10. 11. 1% 1i LIST OF TABLES Distribution of some lichen genera within the montane forest zone at three localities . . . Physical characteristics of three regions within the montane forest zone Percent of endemics in the Greater Antilles Percent of endemics in each of four sub-floras in the Lesser Antilles (Beard, 1947) Aromatic aldehydes giving a positive color reaction with p-phenylenediamine . . . . . . Depsides not giving a color reaction with p-phenylenediamine but giving positive color reaction with potassium hydroxide and calcium hypochlorite . . . . . . . . . . . Depsides not giving a color reaction with PD, K, C or KC Various substances reacting PD- and belonging to different chemical groups Unidentified substances reacting PD—, K-, C-, KC- or KC+ Distribution of the chemical variants of R. subasBerata in the Greater Antilles . . . Percentage of 214 specimens of R. comEIanata from Puerto Rico with each of seven c emica constituents as determined by microchemical tests and thin-layer chromatography Distribution of West Indian species of Ramalina and their chemical variants Species names associated to chemical variants of R. usnea viii Page 64 66 80 81 126 127 128 128 129 201 207 209 270 ‘1, Geographical di variants of Rio :3. Clerical contei Florida Ramal ir Li. list of lichen species of Ram of .3: ' I Page Tag ‘nts of Ramalina usnea. . . . . . . . . . . . . 272 , ical contents.of West Indies and erida Ramalina . . . . . . . . . . . . . . . . . 309 9% List of lichen substances reported in 114 species of Ramalina . . . . . . . . . . . . . . . . 338 ix m5 L. Physiograph1c [Adapted from 1. Geologic map rap PRWRA: b)’ I Associates ’ 3. Map based 0“ rean annual A Caution Shou: this map, Pa: {Adapted froo Sm Juan. P11 1. Climatic map toe Thorn“?li (Adapted fro :. Vegetation ' .. from Danserf E. lose map C0I in Puerto R3 300 and 600 . Distributio‘ 3. Distributio 3. Distributic I Distributic .. Iistributit Iistributir 3 subas e :3IallnlC 3. n {ELIEEUII 3 so asp orot e OCCII" C LIST OF FIGURES Figure Page 1. Physiographic regions of Puerto Rico (Adapted from Mitchell, 1954). . . . . . . . . . . 279 2. Geologic map of Puerto Rico (Adapted from map PRWRA, by Black and Veatch; Domenech 8 Associates, 1971). . . . . . . . . . . . . . . . 280 3. Map based on the period 1931-1960 showing mean annual precipitation in inches. Caution should be used in interpolating this map, particularly in mountainous areas. (Adapted from map by U.S. Weather Bureau, San Juan, Puerto Rico). . . . . . . . . . . . . . 281 4. Climatic map of Puerto Rico according to the Thornwaite system of classification. (Adapted from Thorpe, 1941). . . . . . . . . . . . 282 R Vegetation zones of Puerto Rico (adapted } from Dansereau, 1966). . . . . . . . . . . 283 1 6. Base map containing 75 collection localities in Puerto Rico. Contour lines at approximately 300 and 600 meters. . . . . . . . . . . . . . . . 284 L Distribution map of Ramalina dendriscoides . . . . 285 8 Distribution map of Ramalina furcellata . . . . . 286 9. Distribution map of Ramalina sorediosa . . . . . . 287 10. Distribution map of Ramalina gracilis . . . . . . 288 11. Distribution map of Ramalina subasperata . . . . , 289 12. Distribution map of chemical variant of R. subas erata containing sekikaic and sala21n1c ac1ds. . . . . . . . . . . . . . . . . . 290 13 Distribution map of chemical variant of R. subasperata containing sekikaic and protocetraric acids. . . . 291 “strnuticn m 15. Distribution m l. complanata if. Distribution n l. complanata 1'. Distribution I I. complanata Distribution ‘ i. complanata 13. Distribution '1, Distribution '1. Distribution ‘1. Distribution Distribution ‘1. Distribution Distribution Distribution 7 r. . ' ilWlbUtior Figure Page 14. Distribution map of Ramalina complanata. . . . . . 292 15. Distribution map of chemical variant of R. complanata containing divaricatic acid . . . . 293 16. Distribution map of chemical variant of R. complanata containing protocetraric acid . . . 294 17. Distribution map of chemical variant of R. complanata containing salazinic acid . . . . . 295 18. Distribution map of chemical variant of R. complanata containing unknown substance A . . . 296 19. Distribution map of Ramalina leptospora . . . . . 297 20. Distribution map of Ramalina peruviana . . . . . . 298 21. Distribution map of Ramalina farinacea . . . . . . 299 22. Distribution map of Ramalina bistorta . . . . . . 300 23. Distribution map of Ramalina subpellucida . . . . 301 24. Distribution map of Ramalina anceps . . . . 302 25. Distribution map of Ramalina peranceps . . . . . . 303 § 26. Distribution map of Ramalina dendroides . . . . . 304 I 27. Distribution map of Ramalina usnea . . . . . . . . 305 xi Eigure 3. :igore 3. LIST OF PLATES Plate Page I. Figure l. Crystals formed by atranorin in G.A.o—T solution ( x 65). Extracted from Ramalina complanata . . 133 Figure 2. Crystals formed by caperatic acid in G.E. solution ( x 65). Extracted from Ramalina bistorta . . . . . . . . 133 Figure 3. Crystals formed by cryptochloro- phaeic acid in G.A.W. solution ( x 80). Extracted from Ramalina complanata . . . . . . . . . . . . . . 133 II. Figure 4. Crystals formed by divaricatic acid in G.E. solution ( x 160). Extracted from Ramalina subpellucida . . . . . . 138 Figure 5. Crystals formed by norstictic acid in G.A.o—T. solution ( x 180). Ex— tracted from Ramalina anceps . . . . . 138 Figure 6. Crystals formed by protocetraric acid in G.A.o—T. solution ( x 70). Ex— tracted from Ramalina complanata . . . 138 III. Figure 7. Crystals formed by psoromic acid in G.E. solution ( x 100). Extracted from Ramalina gracilis . . . . . . . . 142 Figure 8. Crystals formed by ramalinolic acid in G.A.W. solution ( x 170). Ex— tracted from Ramalina subasperata . . 142 Figure 9. Crystals formed by salazinic acid in G.A.o-T. solution ( x 70). Extracted from Ramalina peranceps . . 142 xii . Figure 13. 10. Cr‘ Eigure 11. Cr Figure 12. Cr Influx m Figure 14. ( Plate IV. Figure 10. Figure 11. Figure 12. V. Figure 13. Figure 14. Figure 15. Crystals formed by sekikaic acid in G.E. solution ( x 170). Ex— tracted from Ramalina peruviana . Crystals formed by usnic acid in G.E. solution ( x 150). Extracted from Ramalina complanata Crystals formed by substance H in G.E. solution ( x 45). Extracted from Ramalina usnea . Crystals formed by unidentified substance A in G.E. solution ( x 130). Extraced from Ramalina complanata Crystals formed by unidentified substance B in G.E. solution ( x 150). Extracted from Ramalina cochlearis Crystals formed by unidentified substance C in G.E. solution ( x 600). Extracted from PD+ variant of Ramalina sorediosa . xiii Page 147 147 147 152 152 152 i 9..) guilt T. List of Col 3. The Natural the Lichen I. list of Lil in 114 Spe LIST OF APPENDICES Appendix Page A. List of Collection Localities . . . . . . . . . 327 B. The Natural Products Reported from the Lichen Genus Ramalina . . . . . . . . . . . 336 C. List of Lichen Substances Reported in 114 Species of Ramalina . . . . . . . . . . 338 xiv Although th :Ezie best known i its been piecemeal ::~:: the object of ' ~~~~~~ CHAPTER I INTRODUCTION Although the flora of Puerto Rico is probably one ofthe best known in the West Indies,vegetational studies vaebeen piecemeal and incomplete. The lichens have not bemithe object of a general floristic treatment and rqmrts are limited to descriptions of small collections (Wfller, 1888; Riddle, 1915; Fink, 1927; Vainio, 1929; Zdflbruckner, 1930a and Hedrick, 1930). Hedrick compiled alist of lichens reported from Puerto Rico and The Virgin Islands to be published as a portion of Volume VII of the Scientific Survey of Puerto Rico and The Virgin Islands. Hm list contains 710 species and subspecies representing MO genera in 41 families. Imshang (1957) prepared a catalogue of West Indian lichens. This study originated as an investigation of the rolelichens played as components of the vegetation of theisland. Due to the little information available and Um impossibility of studying the entire lichen flora in Um short time available, it was felt that a critical study of the taxonomy and ecology of one genus might flunish insight into the general role played by lichens "is vegetation of (I) ::.:e etudie . The lichen g self to such stud: in difficult gen‘ at either pendent rasaod shrubs, 1: ii: and size. Ft zucal variants l :gnaoce was the tangoized in the Early in t 312i the role of .:\i .‘l in the vegetation of Puerto Rico and provide the basis for future studies. The lichen genus Ramalina was believed to lend itself to such study for several reasons. This taxonomi- cally difficult genus consists of fruticose lichens that grow either pendent or erect on trunks and branches of trees and shrubs, presenting a great variability in growth habit and size. Furthermore, the genus contains numerous chemical variants which required analysis. Of no less importance was the consideration that Ramalina is easily recognized in the field and was known to be locally abun— dant. Early in the study it was realized that to under- stand the role of the genus in the vegetation it was necessary to study the taxonomy of the West Indian species and prepare a key to separate them. Thus, the collections obtained from field studies in the summer of 1967 and in the winter of 1968 were supplemented with herbarium material from adjacent areas.1 A review of the major plant communities in the West Indies and adjacent areas is included as a point of reference for this and future works. It is hoped that this attempt provides an initial step toward future studies in which the floristic aspect of the lichen flora is related to the overall vegetation. E.— Specimens cited in the text as "Landrén" are dEposited in Michigan State University Herbarium (=MSC). GENERAL 1 OF THE HE This is the 'lztistic Kingdom N aergreeo \'egetati( Zzzi, 1961.; Cleasr CHAPTER I I GENERAL DESCRIPTION AND VEGETATION OF THE WEST INDIES AND ADJACENT AREAS This is the northeastern part of the Neotropical Floristic Kingdom which is characterized by tropical evergreen vegetation on loamy soils and high rainfall (Good, 1964; Gleason and Cronquist, 1964). Florida is a long peninsula occupying the southernmost tip of the United States. The region is a crossroad where the northern flora meets the West Indian and mixes with indigenous flora. The simple topography of the peninsula only supports a lowland vegetation and the characterisitic montane formations of the American tropics are absent. Pine communities are found throughout Florida and may be divided into scrub, sand hills and tropical pine associations (Carr, 1949; Laessle, 1942 and 1958). Reindeer-moss lichens (Cladina spp.) cover the ground in Open areas within the scrub association. Hammock com- munities, often found as islands within the pine forest :ssseg‘, Eli), are TEES if temperate ‘ :tharacterized b zurophytic forest tidy distributed E13). South of pt Liars called the indistinct types EgilaIlOD and man :Lzzar development tech. messey, 1911), are composed of broad—leaved evergreen trees of temperate and tropical affinities. These hammocks we characterized by a humid environment in the middle of axerophytic forest. Everglades and swamp communities are TMdely distributed in southern Florida (Davis, 1940 and 1943). South of peninsular Florida is a long fringe of islets called the Florida Keys. Davis (1942) recognized two distinct types of vegetation on these islets: strand vegetation and mangrove swamp. Either type may reach its climax development in an upland forest known locally as hammock. The Bahamas The Bahamas form an archipelago east of Florida mm.north of Cuba and Hispaniola. The islands stand on shallow sandy banks with deep waters between them. The topography is low but hilly and rocky. Once there were extensive pine forests in the major islands enclosing hardwood vegetation similar to the hammocks of Florida Written and Millspaugh, 1920). The present vegetation of the islands consists of second growth and strand com- munities (Howard, 1950)~ IT The Greater pm; in the West dherto Rico. T :‘p'mby Anthony I more), analyzi :zthey were of t The island: serene iluctuat ior :2 the northeast Ezztieast-northwes Iiitryvinters a result in great lc ifislittle as 2( ithe rainforest: 515 iSlands combi rate 4 - °~ evaporati The Greater Antilles The Greater Antilles include the four largest islands in the West Indies, Cuba, Hispaniola, Jamaica andPuerto Rico. They were thought to be of continental ofigin by Anthony (1925) and Schuchert (1935) but Darling— tm1(1938), analyzing distributional evidence, thought thm:they were of oceanic origin. The islands have a tropical oceanic climate without extreme fluctuation in temperature. The prevailing winds are the northeast trade winds while hurricanes follow a smnheast-northwest path. The regional rainfall is seasonal wiflidry winters and wet summers. Local conditions, however, remflt in great local variation with an annual precipitation of as little as 20 inches in dry areas and over 150 inches hithe rainforests. The relatively high temperatures of theislands combined with the trade winds, produce a high rate of evaporation. Cuba Cuba, the largest of the West Indian islands, lies atthe mouth of the Gulf of Mexico and except for the Bahamas is the nearest to the North American continent. A cmmral plain extends throughout the greater part of the island without interruption other than the Trinidad Moun— tahw. The highest mountains, the Sierra Maestra, are fmmd in Provincia de Oriente at the eastern end of the aid ”mine“ even end 0f ‘ numerous Para” 3905. 5011 eroS int the forlotltion g'rogotes . u The coast“ testraod communi‘ TITS which are ‘ 125231 lowlands. 51': types: hard :i ephemeral herb :e inland mogoteE Erelcped (Seifrii The singll tesaraona (Bear {Lei either by th .i‘ilaoi three typ 15;, bushy and s Shoas are prir firstly on se' iiirihed tuo sa Euro and the The fore 5-1 the hroad~le (demon ShOW , island and culminate in Pico de Turquino (6180 ft.) . At the western end of Cuba, in Provincia de Pinar del Rio, are numerous parallel ridges called Cordillera de los Organos. Soil erosion in this province has also brought about the formation of low conical limestone hills known as "mogotes." The coastal vegetation in the island is formed by the strand communities, rock communities and mangrove swamps, which are the most extensive formations of the coastal lowlands. The Cuban deserts have three ecological plant types: hardwood thorny shrubs, perennial succulents and ephemeral herbs. On the coastal limestone hills and the inland mogotes a xerophytic type of vegetation has developed (Seifriz, 1943). The single formation that covers most of Cuba is the savanna (Beard, 1953). Cuban savannas can be classi— fied either by the vegetation or by the soil. In the lowland three types of savanna have been reported: the palm, bushy and sandy. As regard to soil the western savannas are primarily siliceous, while the eastern ones are mostly on serpentine soil. Carabia (1945a and 1945b) described two savanna types of the highlands: the pine savanna and the savanna proper. The forests of Cuba can be divided into pine forests and the broad-leaved forests (Smith, 1954)., The montane vegetation shows four belts: the coastal xerophytic forest, L‘ :2::itlE zone bel tithe smnmit belt soot as well deve Soariola .a— The island :5 Greater Antill :3 the Dominican .siaod is characte 2::stal strip. Ar 7 the middle zone belt or "monte fresco," the upper belt ' formed by a mixed forest (pines, tree ferns and palms), and the summit belt. The cloud forest in the summit belt is not as ’well developed as in the other West Indian islands. Hispaniola The island of Hispaniola is the second largest of the Greater Antilles comprising Haiti in the western end and the Dominican Republic in the eastern two—thirds. The island is characteristically mountainous with a narrow coastal strip. An extensive highland area occupies the central part of the islands. The backbone of the island is a high mountain range extending in an east-west direction. This is the Cordillera Central containing Mount Duarte T (former Mount Trujillo) (10500 ft.), the highest peak in the West Indies. To the north of this range lies the Cordillera Septentrional and between the two ranges is Cibao Valley, which extends into Haiti as Plain du Nord. The most recent treatment of the flora of His— paniola is that of Moscoso (1943). The vegetation was the object of a classical study by Ciferri (1936). Holdridge (1945) recognized four major vegetation zones in the island: the mangrove, dry, moist and pine forest. The mangrove vegetation does not differ from that elsewhere in the West Indies. Most of the vegetation falls within the dry forest type. This includes the sand—beach L‘ “ea-I— :;e:a'.ion and the 5 :23 into open : rind savanna in l Though pine trees ihidge [1942) ha fisarannas. The moist T trim; the low moj 'ii:oss forest, {Lied-by a single :: acquire a 5 avan \wgetation and the savanna. Beard (1953) classified the smmnnas into open savanna, usually devoid of trees and ordmrd savanna in which grasses are ecologically dominant, ahmough pine trees are present. Chardon (1941) and Hohhidge (1942) have studied the ecological aspects of the savannas. The moist forest exhibits three facies in the vege- tatimu the low moist forest, the high mountain forest and thenwss forest. The pine vegetation in the island is fmmwd by a single species, Pinus occidentalis, and tends to acquire a savanna aspect. Jamaica The island of Jamaica lies 18° north of the equator andis third in size in the West Indies. The eastern end mmports the Blue Mountains which culminate in Blue Mountain Peak (7400 ft.). The central part has low mountains, the norfllside has gentle slopes and the south side harbors the dry plains. The vegetation of Jamaica was discussed by Swabey U949) under climatic and edaphic types. Asprey and mmbins (1953) considered that the vegetation falls natur— aly under coastal, lowland and montane formations. The coastal vegetation was described as a desert by Shreve (1910), meless and Asprey (1957) and Asprey and Loveless (1958) MQgested that the dry evergreen formation could be arranged grist of increa! 157115 thiCket uuhcoast; while "richer, bushland Junta in wet ant min the same 2 31:]. No true I Santa, but Aspr take evidence of aggruach to the n li'ztains [Shrew sclerophyll type 1311‘} nor Beard 21113 in their c 13:10.1. hiorder of increasing habitat adversity as follows: bush— 1and,palm thicket, hedge and pavement communities on the norflicoast; while on the south coast the sequence would be flflcket, bushland and scrub. Lowland formations occur in Jamaica in wet and dry limestone. The swamps and marshes cmuain the same associations as in other islands (Chapman, 1944). No true rainforest can be distinguished today in Jamaica, but Asprey and Robbins (1953) find in the tree fern brake evidence of their former existence. The closest approach to the montane rainforest is found in the Blue Mmumains (Shreve, 1914). At high elevations there is a sclerophyll type of vegetation for which neither Barbour U942) nor Beard (1942a, 1944a, 1946, 1955) have counter- parts in their classification system of the montane vege— tation. The Lesser Antilles The Lesser Antilles are a chain of small islands dispersed in an arc between the North and South American continents. These islands are of different geological origin and belong to different political units. According to Davis (1926) the islands are of volcanic origin and never have been connected to the continents. The vegetation of the entire group has been treated by Hodge (1941), Beard (1949) and Vélez (1957). Stehlé g:;,1915a, 1945b 5;; 1913) treate egetation of the 3 her treated the 6 rated Antigua. l iluuland and mom Esraup formatiou zeciluuus forest. Ezrest, the lower Einhytic forest host of th Feet and includu fizetions (Beard 115i$1ands is th km in Domin :16 and Who] iTStehlé [1945bf ssszeu of Beard 53331311 {Orma t .1 10 0941,1945a, 1945b and 1946) treated the French Islands, lbdge (1943) treated Dominica, Howard (1950) studied the vegetation of the Bimini Island Group in The Bahamas and lamr treated the Grenadines (1952). Loveless (1960) treated Antigua. The vegetation falls under the headings oflcwland and montane. The lowland vegetation comprises Um swamp formations, the xerophytic forest, and the dmfiduous forest. The montane formations include the rain— forest, the lower montane forest, the montane thicket, the nmsophytic forest and the elfinwoodland. Most of the lowland vegetation presents a xerophytic aqmct and includes both the dry evergreen and the seasonal formations (Beard, 1946). A characteristic association in theislands is the Pterocarpus swamp. The best rainforest is found in Dominica (Hodge, 1943). This forest develops hired and podzolic soils. The mesophytic forest described by Stehlé (1945b) corresponds in the general classification Swtem of Beard (1949) to the semi—evergreen and evergreen seasonal formations. The Continental Islands of Trinidad and Tobago The island of Trinidad is generally regarded as the Smuhernmost tip of the Lesser Antilles. Geologically the island is part of Venezuela. A mountain range in the northern part Of the island contains the highest peaks, Pico Tucuche andIHCO Aripo (ca. 3000 ft.). Across the center of the :1: runds a rang ages alluvial teu hectutral valley: relative humidity. hang from north ieuet season. R anal range to 4 Beard (192 Ezrthe size of ti E31946) has reel '11 climatic and iihe referred t The lowla 3““ and belonf .... 11 island runds a range of low hills. Between these two ranges alluvial terraces dissected by steep rivers form the central valleys. The climate is tropical with high relative humidity. The prevailing winds are easterly varying from northeast in the dry season to southeast in the wet season. Rainfall varies from 108 inches in the central range to 40 inches in the dry areas. Beard (1945) pointed out that the flora is rich for the size of the island. The same author (Beard, 1942a and 1946) has recognized fourteen vegetation types divided into climatic and edaphic climaxes. The vegetation types can be referred to as lowland or montane. The lowland vegetation types are the most wide- spread and belong to the seasonal formation series or to the semi-evergreen seasonal formation series. The montane forests include the lower montane evergreen rainforest, the montane rainforest and the elfinwoodland. The island of Tobago presents essentially the same characteristics as Trinidad. Both represent a disconnected portion of the South American continent. While the flora of Tobago is poorer than that of Trinidad, the former presents more oceanic elements. The vegetation types are the same in both islands (Beard, 1944b) . Central Ame luragua, Honduras flunto south of tasuothorder the steuala touches :Elaualina have b ~ Ezluras and Yucat 12 Central America Central American includes Panama, Costa Rica, Nicaragua, Honduras, El Salvador, Guatemala, and the part of Mexico south of the isthmus of Tehuantepec. El Salvador does not border the Caribbean and only a minute part of Guatemala touches it. Of the remaining countries specimens of Ramalina have been seen primarily from Panama, Costa Rica, Honduras and Yucat an . Panama The Republic of Panama is the southernmost country of Central America, bordering Colombia on the east, Costa Rica on the west and having the Caribbean and the Pacific Ocean on the north and south respectively. The mountains in the country run from east to west forming a continuous range interrupted only at the Canal Zone. This range reaches its highest peaks at volcan Chiriqui and Bard (above 12000 ft.). The country lies within the tropical climatic zone with temperatures being higher on the Pacific side than along the Caribbean. Schery (1945) and Holdridge (1956) distinguished tropical, sub—tropical, lower montane, and montane vegetation belts. The tropical belt has been well studied at Barro Colorado Island (Standley, 1933) and at Darien (Lamb., 1953). 1.2 1:32 W I’- Costa Rica like Caribbean 31 sultry is uountail :thetoasts. A] (nurses the coun irihhean side rai tileou the Pacif The flora :leutral America :rided the veget: Esltlantic “tie ind forests of :slieute" compris Fizifit side. "1 :e highland veg< 13 Costa Rica Costa Rica is a small country bordered on the east by the Caribbean and on the west by the Pacific Ocean. The country is mountainous and the narrow valleys are restricted to the coasts. A mountain range with volcanic peaks traverses the country rising to about 11700 feet. On the Caribbean side rainfall is abundant throughout the year, while on the Pacific side it is much less and seasonal. The flora of Costa Rica is one of the best known in Central American (Standley, 1938). Standley (1945) divided the vegetation according to geographical regions. The Atlantic "tierra caliente" comprises the lowland and upland forests of the Caribbean side; the Pacific "tierra caliente" comprises the dry savannas and grasslands of the Pacific side. "Tierra templada and tierra fria" include the highland vegetation . Honduras The republic of Honduras is located between Guate— mala and Nicaragua and is triangular in outline with its base forming the northern coast along the Caribbean. The country is rough and mountainous, with the principal ranges extending in an east-west direction in the south central Part of the country. Temperatures range from decidedly tropical on the coast to moderately temperate at high wins. The no mughout the year The flora : 12h knoum. Sta! in and luncker :1tal region. I 2:1:leared for b tithe country the :‘erior areas the saunas [l’ogel, i Silur to that o .... Yucatan f of llexico. ":91 The nor .ltrum a brac iith l4 devations. The northern coast receives ample rainfall fluoughout the year. The flora and vegetation of Honduras are imper— fmxly known. Standley (1931) studied the northern valley flora and Yuncker (1938, 1940, 1945) studied the northern cmmtal region. The vegetation of the northern plains has bemrcleared for banana plantations. Toward the interior ofthe country the mountains are densely forested. In some huerior areas the pine and oak forest forms park-like savannas (Vogel, 1954). In general the vegetation is very shfllar to that of other areas of Central America. Yucatan Yucatan forms a broad peninsula extending into the one of Mexico. For the most part it is flat country with alow sierra (900 ft.) running across the southwestern corner. The northern coast is characterized by sand dunes bordering a brackish lagoon. The temperature is tropical mm the climate shows a sharp contrast between high summer andlow winter rainfall. The natural vegetation is best described as a dry forest that contains features of tropical humid forests and xerophytic woodlands but northern Yucatan tends to support mm-xerophytic vegetation (Bequaert, 1933; Swallen, 1934). The Republi lull herica. Th4 which lies be islierra Madre 0 ssuleuel to 18885 invariable fron trend the lowlant :the northern d: Attempts ‘ 15 Mexico The Republic of Mexico lies between Central and NorfluAmerica. The country is rugged with a central pla- temrwhich lies between the Sierra Madre Occidental and Um Sierra Madre Oriental. Elevations in Mexico range from sealevel to 18885 feet on Monte Orizaba. The climate is venrvariable from north to south and from the high peaks 1mward the lowlands. Rainfall may be as little as 4 inches hithe northern deserts to 200 inches in the mountain zone. Attempts to map the vegetation of Mexico have been mamaby Standley (1930), Ochoterena (1945), and Starker U950). These have been summarized by Pesman (1962). In general the vegetation types have been divided into tem- perate and tropical types as follows: A. Temperate 1. Desert. This includes cactus scrub, creosote bush and alkali flat associations. 2. Mezquite and grassland. Within this zone four communities have been recognized: mezquite scrub 1 short grass prairie, grassland and Agave—cactus. 3. Pine—oak forest. 4. Boreal forest. This zone occupies the peaks where alpine meadows are also present. B. Tropical 5. Arid scrub. This is a mixed thorn forest. 6. Thorn forest. This is dominated by thorny legumes, \u—‘. *m Gentry sane soils 9-1111 10. Rainfc and ti from 1 11. Cloud fog a “M 16 7. Deciduous forest. This forest was described by Gentry (1942) from the canyons. 8. Savanna. This type is characteristic of poor soils along the coasts. 9. Evergreen forest. 10. Rainforest. Occurs on the inaccessible slopes and there is no sharp boundary separating it from the previous type. 11. Cloud forest. This is characterized by dense fog and high humidity. Venezuela The country of Venezuela occupies the most northern part of South America and is bordered by the Caribbean Sea hlthe north. The Paria Peninsula forms the eastern extrem— ity. This peninsula, together with the islands of Trinidad, Tobago, Margarita and Tortugas, is part of a former chain of nwuntains. The highest peak in the country attains 15000 feet in the Andes. The temperature varies from hot in the lowland to freezing in the Andes. Rainfall is usually seasonal with wet winter seasons and dry summer seasons. Pittier and Williams (1945) divided the country in fmu'geographical areas: (1) The coastal range on the north— emipart of the country with xerophytic and thorny types of vegetation; (2) the Andean region with the paramo vegetation; (Q the llanos, which have a variable relief and support mamas and woodl: itle Orinoco, wh Puerto Ri of the Great iripital zone, be p.- ihi°35‘ and 6', texugular in or tension along ‘ 2:111 miles a CHAPTER III PUERTO RICO General Description Puerto Rico is the smallest island and farthest east of the Greater Antilles. It lies well within the tropical zone, between 17°55' and 18°3l' north latitude and 65°35' and 67°15' west longitude. The island is rectangular in outline as shown in Figure 1. The greatest fimmnsion along its main axis is 113 miles and it is about 41 miles across. The largest of the off shore islets are Vieques and Culebra off the eastern coast and Mona off the western coast. The island is surrounded on the north and east by the Atlantic Ocean, on the south by the Caribbean and on the west by El Canal de la Mona. Physiography The first physiographic study of the entire island was done by Lobeck (1922) as part of the Scientific Survey Of Puerto Rico and the Virgin Islands. Lobeck rec0gnized three physiographic regions on the island: the upper pene— Plane, the lower peneplane and the coastal plains. The upper peneplane represents the results of the first period 18 Earth after “1 anion and liftin send lover plate argeute of land eulnd during t this. A detaile: air may be four Lfihlmmg(195 iuhll(l954l h Einsvhich are The most ismnhern coas findngcharactu Eentuest de Almost t iifléedeposit rr‘fera‘u lakes an 19 of erosion after which the island was raised. Subsequent erosion and lifting of the submerged land brought about a second lower plateau called the lower peneplane. Further emergence of land upon which thick deposits of limestone were laid during the Tertiary period formed the coastal plains. A detailed geographical description of the topo- graphy may be found in the works of Pico (1954, 1962 and 1963), Young (1955) and Pico, Buitrago and Berri’os (1969). Mitchell (1954) has distinguished seven physiographic regions which are delineated in Figure 1. The most extensive flat lowland occurs on the northern and southern coasts. According to Roberts (1942) the out— standing characteristic of the northern coastal lowland is the east-west deposition of limestone. Almost the entire shoreline of the island is covered by large deposits of white or yellow sand. The island has several lakes and numerous small rivers. The largest and only fresh water lake is Tortuguero, west of Vega Baja; Laguna San José on the northeast and Laguna Guanica on the south contain brackish water. The largest rivers are Rio Grande de Arecibo and Rio La Plata in the north, Rio Grande de Loi'za in the northeast and Rio Culebrinas in the west coast. On the south the rivers are short and shallow and most of them dry out during the drought season. ". ”'1‘ T On the mos :slizestone belt lese deposits are due thel’ assume :dled'uogotes" < 1:: top known 85 :2ghest relief 0 The main ‘ the eastern en 1i restern parts 21:1 than to the fat to vest. Tl trier of the rar The less 212:: of a rocky fiittoast, cove Tviritton (941 :2: ul, El re 20 On the most southerly part of the northern lowland thelimestone belt is formed by almost pure chalk deposits. flwse deposits are more characteristic of the western end where they assume a conical shape and form steep, low hills cflled ”mogotes” or long loaf—shaped mounts with a narrow flattop known as "pepinos." These low hills may have the nnghest relief of the island due to rainfall erosion. The main mountain range is known as Sierra de Cayey hithe eastern end and Cordillera Central in the central mm.western parts. The range is a little closer to the souflrthan to the north coast and traverses the island from east to west. The highest peaks are attained near the center of the range where Cerro de Punta rises to 1338 meters. The less extensive range of Sierra de Luquillo con— flsts of a rocky, steep and wet chain of mountains along the east coast, covered by dense forest. The highest peaks are Mr.Britton (941 m.), East Peak (1051 m.), The Pinnacles (1050 m.), El Yunque (1065 m.) and 131 Toro (1074 m.). Geology Berkley (1915) carried out the initial geological reconnaissance of the island, which formed the background forsubsequent reports. The series of publications by the New York Academy of Sciences (1919—1931) represents the greatest effort to elucidate the geology of Puerto Rico. Hm results of these surveys were published by Semmes (1919), L;— ~:i;s T1920], Mitcl giTsjierhoff (193‘ arlieruorks and ill a generalized hens [1942) con Eanus: and Thom :f'he Tertiary a1 shell [1954) tl tiierrios (1969' 21 Hodge (1920), Mitchell (1922), Hubbard (1923), Fettke (1924) and Meyerhoff (1931). Meyerhoff (1933) summarized these earlier works and to date is the only publication dealing with a generalized review of the geology of the island. Roberts (1942) conducted a soil survey of the island. Zapp, Bergquist and Thomas (1948) studied and mapped the geology of the Tertiary areas of the north and south coasts and Mitchell (1954) that of the other periods. Pico, Buitrago and Berrios (1969) provided maps of the geology of Puerto Rico. Many papers and maps about different aspects of the geology of the island were compiled in the Transactions of the Second Caribbean Geological Conference held at the University of Puerto Rico at Mayagiiez in 1960 (Kaye and Dunlap, 1960; Hildebrand, 1960; Mitchell, 1960; Weaver, 1960). Meyerhoff (1933) modified Lobeck's concepts about the physiography of Puerto Rico and proposed his own hypo— thesis on the origin and evolution of the island. The origin of the island probably goes back to the Cretaceous. During this period the island seems to have gone through several periods of emergences and submergences. By the end of the Cretaceous soft rocks emerged forming hills and valleys while the hard rocks built up the mountain ranges. By the early part of the Tertiary erosion had worked enough to form hills and mountains as they are seen today. Puerto Rico was connected to Hispaniola probably until the Pleis- tocene . 0n the non tiesiread rocks a run from the CIT imdaut. Figure ilaud. Puerto Ri :iified by land tale winds blow Sierra de luquil‘ is northern slo 2::thi'estern cor id“ of rainfa 22 On the mountains of the island the most common and widespread rocks are volcanic, granitic and diorite. Lime- stone from the Cretaceous and Teriary periods are also dnmdant. Figure 2 shows a generalized geologic map of the island. Climate Puerto Rico has a tropical, marine climate slightly nwdified by land breezes. The moisture laden northeast trade winds blow almost continuously and first strike Sierra de Luquillo. The highest precipitation falls on Hm northern slopes of the mountain ranges. The south and northwestern corners and adjacent islands receive the least mwunt of rainfall because of the absence of mountains, or because the clouds lose their moisture on their way south— ward. The U.S. Weather Bureau records (1967) for the period from 1931 to 1960 show that the average precipita— tion varies from less than 35 inches per year on the south- west to over 200 inches on Sierra Luquillo. The data finther indicate that there are no distinct seasons, but mahfly a rainy period from May to November and a dry period with sporadic showers the rest of the year. The mean annual precipitation in inches is shown in Figure 3. The temperature is ameliorated during the day by Um trade winds and during the night by the mountain breezes. mm to 100°] age from 45°F '9 an 50°F or ris :ehigh peaks an Early part of the The clina 1:21 occurrences A simple lisiders not on 7:1qu (1941). .slaud yields fi EsIi-arid, and a illetion accordi 23 The local climatological data published by the U.S. Weather Bureau (1967) indicate that the mean monthly temperatures along the dry coast range from 75°F in winter to about 80°F during summer. The mean highest annual temperatures range from 91°F to 100°F and the mean lowest annual temperatures range from 45°F to 61°F. Rarely the temperature falls below 50°F or rises over 90°F. Frost is unknown, but on the high peaks and valleys a dense fog is formed during the early part of the day and during winter months. The climate of the island is influenced by period~ ical occurrences of hurricanes and storms. A simple system for climate classification that considers not only rainfall but vegetation as well was used by Thorp (1941). The application of this system to the island yields five climate zones: wet, humid, sub-humid, semi—arid, and arid. Figure 4 shows the climatic classi— fication according to the Thornwaite system (Thorp, 1941). W Historical Review of Classflication Schemes The first attempt to map the vegetation of the island was made by Murphy (1916) who recognized four major types: littoral woodland, moist deciduous forest, tropical rainforest and dry deciduous forest. Although Gleason and Cook (1927) made an exhaustive at“ the plant aflT 0‘ Classm maithin vhicI mthem plain, '61 htheir discuss1 an association accessional sequ Tito (19E Stiuguished for :1:: forest, 5111 :11 forest he . The appl :ieuegetation o licf little val iatiou data is r izrided the vege in. The trog 45‘: forests, 24 study of the plant ecology of the island they did not attempt to map or classify vegetation types. The three geographic areas within which they described the vegetation are the northern plain, the mountain region, and the southern plain. In their discussion of the vegetation they recognized many plant associations and related them through a series of successional sequences to the climax communities. Pico’ (1954) adopted Murphy's classification and distinguished four types of natural vegetation: littoral, moist forest, sub-moist forest and thorn forest. In the moist forest he described tropical and subtropical types. The application of Holdridge‘s system (1947) to the vegetation of Puerto Rico by Kumme and Briscoe (1963) is of little value as it is limited to areas where precipi— tation data is readily available. Kurnme and Briscoe divided the vegetation into tropical and subtropical forma- tions. The tropical formations comprised the dry and moist forests, the limestone vegetation and the tropical wet forest. The subtropical formations include the savanna, subtropical dry forest, moist forest, wet forest and rainforest . After many years of field work Little and Wads- worth (1964) published a classification system that included the vegetation of Puerto Rico and the Virgin Islands. The authors recognized eight climatic forest types and an edaphic littoral type. The coastal vegetation wax.» 11:3. dry cow Trusts. The “1°“ agar Cordillera inereau‘s Ve Ct The foreg isrefore providl tier vegetation leueffort to Etsrtu Rico on a is aud analyze studied within \ Siuhic and clir tie iucorporati: 'Ziuthe vege E:33T'Stems and 55-5 context th '53: described. '35 littoral It The 11 moving the ring in CO“t iascnal eyerg €555, tidal i 25 hmluded moist coastal forests, moist deciduous limestone forests, dry coastal forests, and dry deciduous limestone :mrests. The montane vegetation included the lower and umper Cordillera and the lower and upper Luquillo. Dmmereau's Vegetation Cla551f1cation System The foregoing vegetation schemes are sketchy and Uwrefore provide very little help for the integration of oflwr vegetation aspects within their limited framework. In an effort to describe and interpret the vegetation of hwrto Rico on an experimental basis, Dansereau (1966) drew and analyzed numerous vegetation types. These were studied within vegetation zones extrapolated from physio- graphic and climatic data. Such study provides room for Hm incorporation of the lichen flora in a meaningful way widfin the vegetation zones, the vegetation belts, the ecosystems and finally in the community. It is within nus context that the whole vegetation of the island is best described. The Littoral Zone The littoral zone lies near the ocean shore, Occupying the outer belt of the lowland rainforest and coming in contact with the semi—deciduous forest and seasonal evergreen forest. The topography includes lagoons, reefs, tidal belts, dunes and rivers. The vegetz gained by salt oil, all contribi out conspicuous ngrove forest. round the island Esau paid to the soon. Hodridge tigroue from a s 1339] studied t1 26 The vegetation of the littoral zone is primarily (mtermined by salt water spray, sea winds and calcereous soil,a11 contributing to give a xerophytic aspect. The most conspicuous and important element of this zone is the mangrove forest. In Puerto Rico it forms a broken ring annmd the island and adjacent islets. Much attention has been paid to the mangrove by foresters and other investi— gators. Hodridge (1940) discussed the zonation of the mmgrove from a sylvicultural point of view and Wadsworth U959) studied the rate of growth of Laguncularia and Avicennia. The metabolic and community turn-over aspects of the red mangrove (Rhizophora mangle) were detailed by Gdfley, Odum, and Wilson (1962). The mangrove forest is typical in exhibiting species zonation. Occupying the outer zone from open sea to the high tidal area is found the red mangrove. In this community the trees are supported by stilt roots fixing them to the unstable substratum. Toward the shore the next zone is dominated by the black mangrove (Avicennia nitida) , characterized by sending numerous neumatophores out of the mud. The drier zone is invariably dominated by the white mangrove (Laguncularia racemosa). The mangrove Conocarpus erecta is not always part of the mangrove forest. The mangrove complex varies from 8 meters to 20 nmters in height. There are no other tree species associated with the mangrove, and there is no undergrowth vegetation. i‘IC' '12 spnhvtic com ofcrustose liche olocully known :écrustose liche gut of this zone :11 fern Acrosti user is allowed M and TE Much of :51 drained and Igore uegetat iLILiug and lumb is expanding pg 27 The epiphytic community is represented by colored patches of crustose lichens. In certain areas the black mangrove is locally known as yellow mangrove because yellow colonies of crustose lichens cover the tree trunks. In the inland part of this zone, or where the forests have been cut, the tall fern Acrostichum aureum is dominant. If the salt water is allowed to dry, a salt flat is developed, where Portulaca and Batis dominate. Much of the land formerly occupied by mangroves has been drained and planted to sugar cane or coconut. This mangrove vegetation has also been depleted through charcoal burning and lumbering. Large areas have been claimed by the expanding population for house construction and public facilities such as airports, marinas and roads. On the southern rocky limestones the severe climate of the region coupled with scant soil and high porosity of the substrate have induced a thin thicket of microphyllous, deciduous, woody shrubs and trees. The xerophytic aspect is accentuated by the presence of many thorny and succulent species such as arborescent and other cacti. Epiphytes include many bromeliads but lichens are rare. The most conspicuous lichen is a small, black, Pyrenopsidaceous crust covering most of the exposed rock surface. The sand and dune ecosystem does not differ in com- position and structure from those already described for the rest of the West Indies. The vegetation is dominated by is Eeach grasses :i :cconut gm“ The liche rah trees. COW hrscia and flXil Eelouland Rain Rainfo re :‘duforest remai to topography 2 aiovland mesopl Ziis forest prol Stratification 28 the beach grasses, the hedge-forming Cocculoba uvifera and coconut groves. The lichen vegetation is almost restricted to the palm trees. Common are the genera Cladonia, Parmelia, Physcia and Pyxine. Thelbwland Rainforest ZOne' RainfOrest:--Although no mature stand of lowland rainforest remains on the island, the nature of the soil, the topography and climate point to an early existence of alowland mesophytic forest. The typical structure of this forest probably followed the classical pattern of stratification described by Richards (1952) from elsewhere in the tropics. The geographical distribution of the lowland rain— forest almost coincides with Zones I—V and part of VII in Pic6 (1954, pp. 15—33). The first zone comprises the most littoral belt of the humid northeastern coastal plain. Rainfall is abundant, with about 62 inches a year on the drier parts. The second geographical unit consists of the humid plains of the eastern coast from Fajardo to Maunabo. hlthe third region is included the Caguas valley, which is an interior alluvial plain flooded by the Rio Grande de Loiza. Precipitation on the valley reaches an average of 60inches a year. The coastal valleys of the west coast constitute the fourth geographical zone. It begins north j to TulebrinaS ::‘.:ipalities of rivers Anasco, GI aiofall similar hung the summe do the most ea Eetillas was for :sgiou is formed ziiacent to the In Puert :11 occurs abo 13 Species is t .I'a‘laud rainfor 29 of RfiuCulebrinas and extends southward, comprising the mmflcipalities of Aguada, Aguadilla and the valleys of the rhmrs Afiasco, Guanajibo and Yagfiez. The area receives a rainfall similar to the eastern plains but with a peak mning the summer months. Of the fifth geographical zone mfly the most eastern part, which includes the plains of Patillas was formerly occupied by the rainforest. The last region is formed by the most interior limestone hills adjacent to the Cordillera Central. In Puerto Rico there is a complex of species which never occurs above an elevation of 350 meters. This cohort ofspecies is taken to represent the upper limits of the lowland rainforest zone. Conspicuous in this species com- plex are Manilkara bidentata, Lucuma multiflora, Diospyros ebenaster, Stenostomum obtusifolium, Ixora ferrea, Hernandia sonora and Petitia domingensis (Dansereau, 1966). Thus, the floristic delimitation of the lowland rainforest coin- cides with the upper limits of the above mentioned species and the lower limits of another species complex, especially the Dacryodes excelsa association (Tabonuco type forest). On the coastal belt of the northern plain between SmuJuan and Manati there are extensive narrow areas of white sand where remnants of the lowland rainforest remain. Hm best stand, probably in a process of healing, is at Dorado, occupying the upper flat land back of the Ptero— carpus swamp, a few hundred meters from the beach. 2::qu stumps in trees. The under ciages. Shrubs rant. lianas, 30 The soil is a shallow white calcereous sand covered by a thin layer of leaves and humus. The surface of the soil is moist and cool due to the continuous shade of the canopy. The height of the larger trees is about 20 meters, but old stumps indicate the previous existence of larger trees. The understory is rich in small trees of all sizes and ages. Shrubs are rare and herbaceous vegetation is absent. Lianas, ferns and epiphytes are not particularly abundant. Lichens are confined to crustose growths of white powdery masses. The most abundant tree species are Mammea americana, Calophyllum brasilense, and Eugenia jambos. Other tree genera common in the forest include Dendropanax, Clusia, Manilkara, Hymenaea and the palms Roystonea and Acrocomia. The lower strata is formed by seedlings of the canopy trees and a few shrubs such as Pi_p_e_1;, Psychotria and Chioccoca. Climbers are represented by species of Marcgravia, Vainilla and Smilax. Where the forest has been cleared a secondary growth thicket develops and lichens are prominent. The thicket is best developed on the south shore of Lake Tortuguero, west 0f Vega Baja, where the topography is flat with small, shallow depressions forming temporary pools during excessiVe rains. White sand reaches a depth of 2-3 feet and occasionally up to 10 feet. Underneath the white sand there is a thin layer of black soil and below this, a layer “Sta .1. , irsi-‘arovn C1 3)’ grieisites f '1: Tcrocomia. grtirie has evol‘ Batis of ferns fr Title area is th risen clear c iatsloped, where some type of flu occic ‘_ The icac ‘ “191T separ; ifhushes are ' 1:5 sand and th up. 1““ tl'pical o fibrous 31 ofred-brown clay. The scrub occupies the least recently Mstrubed sites fringed by a screen of tall trees of Chsuarina equisetifolia, Terminalia cattapa and the thorny pahnAcrocomia. Where the lake floods the sand a marshy praiie has evolved covered by high grasses and sedges. Banks of ferns form part of the swampy vegetation. The Mufle area is thus a mosaic; where the natural vegetation has been clear cut a Chrysobalanus icaco thicket has developed, where grassing has been the main disturbance a savanna type of vegetation dominated by coconut palms and Anacardium occidentale has evolved. The icaco scrub is almost exclusively constituted by widely separated colonies of Chrysobalanus icaco shrubs. The bushes are very low with many branches lying flat on Um sand and the higher ones showing signs of the stag head effect typical of the mossy forest. The woody shrubs have a very fibrous wood which is so flexible that it was locally owed in weaving baskets. The leaves are small, shiny and thick. Fruit are abundant during summer time. Byrsonima Spicata, Myrcia Splendens, Calophyllum brasilense, Miconia, Randia, Eugenia,Cecropia and Didymopanax are other trees scattered throughout the thicket. All these are typical Species of the lowland rainforest elsewhere in the West Indies. Epiphytes are abundantly represented by Tillandsia Eflflgygga, small orchids and lichens. Herbaceous vegetation is not very conspicuous except where the thicket has been {tenently tut f0 :stcits of graSS he surface is 6 The epipl do" are a majo: 111(1927] repo hriua (=C1adina any shrub 0f t :i the floor to fruticose epiphl’ 25565 hanging 1 5:: dead twigs . 3735111 associ: 32 frequently cut for burning charcoal. The soil is covered by tufts of grasses and lichens, but in many places the bare surface is exposed. The epiphytic lichens locally known as ”barba de pahx‘are a major element of the community. Gleason and Cod<(l927) reported from here the lichen Cladonia gaggi— fefina (=Cladina sandstedei). I have seen here almost evemrshrub of the thicket covered with masses of lichens muithe floor covered by a layer of soil lichens. The fnfiicose epiphytic lichens of the genus Ramalina occur in nwsses hanging from the bases of the main branches and from dead twigs. Rarely small plants of a red Usnea are found in association with Ramalina. The most abundant Ramalina represented in the flucket is R. sorediosa, reaching up to 10 cm. long. It has a fragile thallus, with rounded branches covered by mmwrous mounds of soredia. Rarely the thallus is in fruit. TWO chemical variants of this species are usually found growing together; the most frequent reacting PD— and the other PD+. Another species of the genus present is R. SEE— planata, a smaller species with canaliculate, papillate branches. The species includes both salazinic acid and protocetraric acid variants. These are indistinguishable hithe field and very often are epiphytes on the trunks of Myrcia splendens. Other aerial lichens are represented by foliose arcies of the genus Parme] Err’estoons twisted arour into shrubs. Abundant spo “19.-W» 8‘ aiuhytic lichens in the tl ruin the genera Physcia, dlfl. These are minor osociation. Underneath and on tiesaud surface is covere liheus locally called " f3 oohe association formed 313T: ' ctedeu (referred to il 1' .‘od ' ‘ & assocuation form 1: r ' . free Or white sand r‘LTtlr distinct shrubb\‘ Edged . - Too d B rown~fr ‘Iher Tarts where there "EOTTTTTlo 33 species of the genus Parmelia. On shaded areas these species form festoons twisted around the'lower branches of the icaco shrubs. Abundant species are 1:. cristifera, P. tinc— torum, P. praesorediosa, and P. sulfurea. Other foliose epiphytic lichens in the thicket associated with Parmelia are in the genera Physcia, Coccocarpia, Leptogium, and Collema. These are minor elements of the epiphytic foliose association. Underneath and on the clear spaces between shrubs the sand surface is covered by extensive patches of shrubby lichens locally called "flor de tierra." The most conspicuous is the association formed by the spongy masses of Cladina sandstedei (referred to in Gleason and Cook, 1927, as Cladonia rangiferina) and Cladonia peltasta. This Cladina- Cladonia association forms a prairie limited only by the presence of white sand. In the field they are recognized by their distinct shrubby growth form and distinguished by the whitish color of the Cladina in contrast to the yellowish- green color of the Cladonia. Many other Cladonia Species of smaller sizes and different forms comprise the floor layer. Red—fruited species are especially abundant on de~ composed wood. Brown—fruited species form sheaths on the wetter parts where there is more soil and less sand. Small Cup-forming species grow at the base of the icaco shrubs in association with sterile podetia forming species. The crustose lichens are not very common in the icaco thicket, gen for the yellow Temn fete poles. The second growth chant palms, Anacardium $1613 splendens forming 1 wetland has a thin carpel The lichen commun titrees except for tufts @subpellucida gro that trees and the smal Wig inefl. The c tried in the forest. T} W94 by circular grow“ W Md many specie: 0n the west coas 34 except for the yellow Temnospora on old stumps and dried fence poles. The second growth savanna is dominated by scattered coconut palms, Anacardium occidentale, Andira inermis, and Myrcia splendens forming the arboreal vegetation. The open woodland has a thin carpet of grasses and low shrubs. The lichen community is rich but not very abundant on trees except for tufts of the small fruticose species Ramalina subpellucida growing on dead branches of the cashew trees and the small flattened foliose Physcia albicans on Andira inermis. The crustose lichen layer is the most varied in the forest. The smooth trunks of Hymenaea are covered by circular growths of Haematomma, Pyxine, Buell'ia, Lecanora and many species of the family Graphidaceae. On the west coast a similar scrub of icaco covers the slopes of the Las Mesas hill near Mayagiiez. The soil here is deep red clay supporting a scrubby vegetation of Chrysobalanus and Miconia. A few large trees of Mammea, Bambusa, and many seedlings of Tabebuia heterophylla constitute the arborescent element. This scrub is dense, low and simple. The main epiphyte is the fruticose lichen Ramalina subpellucida. On the ground, particularly under- neath the bamboo screen typically grows a shallow cup— forming species of Cladonia. The podetia characteristi— cally produce proliferating cups from the center or the edge of the lower cups forming three or four tiers of cups raring branches. 0n the 'l 4.:- fl-a‘...“ . ' shes the ground is dotte rustese lichen Diplochist The eastern limit hmstly occupied by sug: This area, along the It high present good habit rosette-like patches of E rothecia are the most co y 0n loose coarse hhapothecia on top of man feel 21H oped within the 10“ her cut and only large :eseen, . This forest wa ‘H n3?) as the only of it arising that they overlc “ht, which lies betwt interest that they The Dorado fore b . . ehmd the SUbClim 35 bearing branches. On the bare soil toward the steeper slopes the ground is dotted by the black apothecia of the crustose lichen Diplochistes. The eastern limit of the lowland rainforest zone is mostly occupied by sugar cane plantations and settlements. In this area, along the roads, large boulders of Cretaceous origin present good habitats for lichen communities. Large rosette—like patches of Physcia albicans with prominent apothecia are the most common followed by species of Hetero— dermia. On loose coarse soil a species of Cladonia with pale apothecia on top of simple podetia is abundant. Swamp forest.-—A few hundred feet from the shore at Humacao playa and at Dorado Beach a swamp forest has developed within the lowland zone. The one at Humacao has been cut and only large stumps with many new shoots are to be seen. This forest was described by Gleason and Cook (1927) as the only of its type on the island. It is sur- prising that they overlooked the Dorado Pterocarpus swamp forest, which lies between the beach and the white sand rainforest that they so well described. The Dorado forest occupies a shallow depression just behind the subclimax rainforest. The area is a brack— ish muddy swamp formed by the accumulation of fresh water during rainfall, and periodical floods from the nearby sea. The swamp has not been invaded by mangrove or any other '.'...'.-.I:2_,_3'__r=a.. gong—:3 ; . :gherplant association. iznedhy a pure stand of Tunas ”palo de pollo" t resemblance of the base b hrgeplank-like roots ri ink and then extend out iehase of the tree. The forest is ope reaching a height of abor nest is continuous but little undergrowth has d paring in the canopy, a 3T» abundant but those p Snore tree to anOther trees. Epiphytes are re Etc ' W and the strar eh of the buttresses 1 36 lfigher plant association. The center of the swamp is fomwd by a pure stand of Pterocarpus officinalis, locally hmwn as ”palo de pollo” or "chicken tree” because of the resemblance of the base buttresses to a chicken leg. large plank—like roots rise 4—5 meters high along the tmmk and then extend outwards to about 4—5 meters from the base of the tree. The forest is open with large clean bole trees readfing a height of about 100 meters. The crown of the forest is continuous but permits ample light to penetrate. Little undergrowth has developed, but where there is an opening in the canopy, a few shrubs are seen. Lianas are rmt abundant but those present are woody, thick and hang frmnone tree to another connecting the crowns of the trees. Epiphytes are represented by ferns, bromeliads, Lycopodium and the strangler fig Clusia. On the rough bark of the buttresses much deposition of dust and humus harbors extensive growth of large foliaceous Parmelias. These foliose lichens are the only noticeable ones and are represented by two yellow medulla species, Parmelia endo- aflphurea and E. sulfurata. Fringing the forest on the dry sites are many palms and large Bucida buceras trees. The role of animals in the community is worthwhile noting here. The presence of Chmia rosea growing as epiphytes high on the tree trunks cmronly be explained by a constant bird population that iepsits the heavy seeds : and hundreds of burrow hnintaining the deep 5 seasons. Unlike the swamp ’iyyey and the mar he forest described ab m Lake and River El R h’oLaPlata and Rio Gra: 2i ' thhornia cover the sur role community. On the an colonis of Castali ‘ “Tetation on the shallc Ethan w, are rapid a the marshy area arour 37 dqmsits the heavy seeds in the top of the trees. On the pmund hundreds of burrows made by the blue land crab help himaintaining the deep soil wet even during the rainless seasons. Unlike the swamp forest at Humacao playa the Acros- tidumlaureum and the mangrove associations are absent from flw forest described above. Lake and River Ecosystems.~-On the broad delta of Rio La Plata and Rio Grande de Arecibo floating mats of Eichhornia cover the surface of the water and dominate the whole community. On the fresh water lake of Tortuguero dame colonis of Castalia and Nymphoides compose the dominant vegetation on the shallow parts. Along the shore, Iypha and Mariscus, are rapid land builders and form large colonies. hithe marshy area around the lake, Phragmites communis and the cattail Typha_constitute the dominant species. The land is under cultivation and its principal crops have been discussed by Pic6 (1954, 1962). Dansereau 0966) has indicated the principal secondary vegetational transformations on the sites formerly occupied by the rain— forest. Since pre—Columbian days the forests of the island have gone through continuous selective cutting for the most Valuable trees. The forests have been continuously clear mm for fuel, cultivation, grassing, and building. Roads and settlements occupy a large portion of the low areas of 5:2 island and thus many 25th roads in Puerto Ri W, Delonix r_e_g_i udeih_a pentandra, C32 ones on the eastern part frequently found along re Terminalia cattapa, and often loaded with bromel gens Parmelia. \ ~on .. Seasonal Evergreen I The seasonal eve o", - .uestone hills that eat HE: A on the north coast 1Oh‘land rainforest 38 Um island and thus many exotics have been planted. Most ofthe roads in Puerto Rico are hedged by naturalized trees sudias Mangifera indica and bamboos, on the west; Spathodea cumanulata, Delonix regia, Hernanadia sonora on the north andtbiba pentandra, Cassia nodosa and other leguminose trees on the eastern part of the island. Other trees fimquently found along roads are Tabebuia heterophylla, Temfinalia cattapa, and Albizzia lebbek. These trees are often loaded with bromeliads, and foliose lichens of the genus Parmelia. The Seasonal Evergreen Forest Zone The seasonal evergreen forest occupies a range of limestone hills that extends from the east to the extreme west on the north coast. The hills are scattered within the lowland rainforest zone or else they form a chain that lies near the foothills of the Cordillera Central. These lfills are deeply cut by tortuous rivers and highly eroded by dissolution of the limestone material. The hills adopt two types of formations: the sugar loaf or ”mogote" has few spurs and exposed surfaces and has Um appearance of an inverted cone on the coastal plain; the haystack or ”pepino” is more rough and has a flat top wifiisteep cliffs. The vegetation on the hills is deter— IMned by a combination of edaphic features and relief and not by rainfall. .- .-.'—.-.-M=3 II“? "“- o The forest is cha aeoegetation to the ext mist deciduous forest- osign the island vegetat ones, deciduous or ever hodhurry and Wadsworth ( heiduousness on the isl: hard [1942b] considered iheering period. Many leaves in reSponse to sh font to length of day Evidence of the fine ”10995 and moist i 39 The forest is characterized by the periodicity of Um vegetation to the extent that Murphy (1916) called it anmist deciduous forest. However, it is difficult to assign the island vegetation to any of the classical cate- gories, deciduous or evergreen. Dansereau (1966) cited Wommurry and Wadsworth of being of the opinion that deciduousness on the island is related to photperiodicity. Beard (1942b) considered that deciduousness is related to flowering period. Many species in Puerto Rico shed their leaves in response to short days period, others are indif— fermu to length of day or to rainfall. Evidence of the lowland rainforest remains on the lower slopes and moist intervals between hills. Actually these sites are planted to sugar cane, pineapple and minor crops. Sometimes the lowland rainforest includes a complex ofspecies dominated by Cecropia peltata, Acrocomia aculeata ’ anezuma speciosissima and Ochroma pyramidalis. Other species present in the complex are Bucida buceras (southern plains), Plumeria alba, Anthacanthus spinosus, and Gaussia attenuata (endemic to the hills). The lichen vegetation in the seasonal evergreen forest is best developed in those hills immersed within the lowland rainforest sites, where lichens (Ramalina sorediosa and 3. peruviana) are abundant in abandoned fields. Other Ramalina Species are rare, but the foliose community is Very well represented by species of Parmelia, Physcia, Eoeroderuia, Collema and ’._——- moot particularly consp Near the sea, in ‘ on river the hills decr orlevel where a low scr his area the calcereous arrange crust of Calopl hrerouycetous lichens. The seasonal eve] inn interferences in t1 odgrassing. The origir Every steep localities at source of cabinet w Many of the lime inderedw' 1th a large nu Tze ' tree specres are mar .nuost frequent. 0n 1 40 Tkterodermia, Collema and Coccocarpia. Crustaceous lichens arenot particularly conspicuous. Near the sea, in the north coast, along the Guaja— taca river the hills decrease in height and gradually reach sealevel where a low scrub is the only vegetation. In His area the calcereous rocks are exposed and covered by miorange crust of Caloplaca and a black crust of various Pyrenomycetous lichens. The seasonal evergreen forest has suffered many hummrinterferences in the form of lumbering, cultivation, and grassing. The original forest is rarely seen except hivery steep localities. This was in early times an impor- tmn source of cabinet wood, charcoal and fence poles. Many of the limestone hills are traversed by roads bordered with a large number of exotic and native trees. Hm tree species are many but Delonix regia is probably the most frequent. On the moist calcereous hills near the central range the lichen flora of road trees is charac— terized by species that grow toward the road. On the middle slopes of the hills a mesophytic type of forest develops. This is formed by tall trees with straight trunks and sclerophyllous leaves occupying the eroded slopes. Locally the forest is known as the "gateado" forest. The conspicuous species are Coccoloba laurifolia, §.diversifolia and Bucida buceras. Tree ferns are absent “— but epiphytic orchids and ferns are abundant. Lianas and nos are not particularl‘ mtlyas colored spots 0 The steep cliffs hestrangler trees (£133 sending out long roots tl lens and AM fill hthe rocks . 0n the most expo: more and a xerophytic ‘ hrs area represent vari homhy the similarity the dense shade as well the roads help in mai Jud environment. Inlz geoeraR ' amalina and Usnr in y and Physcias are 41 vhms are not particularly conspicuous. Lichens occur rmstly as colored spots on the smooth tree trunks. The steep cliffs of the hills are dominated by thestrangler trees (Ficus laevigata and Clusia rosea) smufing out long roots that hang from the rocks. Many femm and Anthurium fill the numerous pockets and crevices in the rocks. On the most exposed spurs, the drought is more severe and a xerophytic vegetation dominates. Species of Hus area represent various degrees of xeromorphism as flmwn by the similarity with the rainforest. Undoubtedly the dense shade as well as the waterways along the shoulders ofthe roads help in maintaining a cool temperature and a mmmd environment. Inland, fruticose lichens of the genera Ramalina and Usnea, and foliose Parmelias, Hetero— denflas and Physcias are very abundant along roads. Semi—Deciduous Forest Zone The geographical area occupied by the semi-deciduous forest zone on the island comprises the lowland of the southern coast, except the littoral zone, the low limestone hulls of Cretaceous origin on the south and the foothills ofthe Cordillera Central. The topography is flat with steep hills toward the central mountain range. Moisture is very scarce throughout this region, wiflirainfall varying from about 60 inches in the extreme southeast to less than I“ hate is very warm and The vegetation i: hoover completely leaf shedding are discovered heir leaves prior to £1 honght season and still ’11. The general opinic hatdeciduousness is re honght and reproductive The climax commo h'hncaro forest" (Bur 1rh‘floristic affiniti PMS 0f the northern l hthe forest are Bucid x duo - . ww» thetm ad in & lnfest the 42 southeast to less than 30 in the southwest corner. The dimate is very warm and dry for the most part of the year. The vegetation is deciduous, however, the forest is never completely leafless. Different patterns of leaf flwdding are discovered in the forest; some species shed their leaves prior to flowering, others only during the drought season and still others never lose their leaves at all. The general opinion among foresters in the island is that deciduousness is related to short daylight photoperiods, drought and reproductive activities. The climax community of the deciduous forest is the'bucaro forest" (Bucida buceras). This forest shows many floristic affinities with vegetation of the drier parts of the northern limestone hills. The common species hothe forest are Bucida buceras, Savia sessiliflora, Coccoloba laurifolia, and Bursera simaruba. On the steep hills is found an open savanna dominated by Bursera simaruba. The trees are rather tall and widely scattered. Under the trees covering the soil is a layer of tall grasses and herbs. Other species associated to the savanna are the cacti (Cephalocereus, Cactus and Opuntia) and woody shrubs (such as, Plumeria alba, Croton lucidus, and Pictetia aculeata). Epiphytic bromeliads of the genus fillandsia infest the branches of the trees and woody shrubs, The thorn—cactus association occupies the hyper— xerophytic habitat of limestone pavements and is similar T Lu :othe xeromorphic flora In the southern : possland vegetation tha‘ noes depending on the d represent a prairie, 5 trees invade the grassla hseen. The mezquite s Salinas and Laj as. The PTyyjuliflora, P_ar_l W; the grasses a: only. Bromeliads : or the savanna. The lichen ’f10r instose forms. Very d any, Phys\cia and Pa hoard the Cordillera—C ioo teases and becoming lohedP ‘ armelias, small it ' ththen vegetation ‘ 43 tothe xeromorphic flora of the littoral zone. In the southern flats, the fine soil supports a grassland vegetation that takes on many structural appear- ances depending on the dominant species. If only grasses arepresent a prairie, steppe or meadow might develop. If trees invade the grassland, a savanna type of vegetation is seen. The mezquite savanna occupies the flat land near Salinas and Lajas. The dominant arborescent species are Prosopis juliflora, Parkinsonia aculeata and Capparis flexuosa; the grasses are Chloris inflata and Andropogon mmulatus. Bromeliads are the conspicuous aerial community of the savanna. The lichen flora is almost entirely restricted to crustose forms. Very depauparate thalli of Ramalina 99m- planeta, Physcia and Parmelia occur on branches of mezquite. Toward the Cordillera Central the saxicolous vegetation increases and becoming especially abundant are the narrow lobed Parmelias, small Heterodermias, and crustose lichens. The lichen vegetation is richer near the upper slopes with broad lobed Parmelias, fructicose Ramalinas, Usneas and Cladonias being abundant. Thus, it appears that a zonation oflichen species emerges in correlation with increased humidity. The forest was depleted of valuable timbers during the period when ox carts were the principal means of trans— portation. But the major transformation of the original To forest has been done by mores. This has crea oodprairies, which will their secondary growth s on under irrigation has orgar cane fields. The humorous large and SI onplexes which have ino remained of the origina to hover Mont ane Rainf This rainforest the v ' lonland rainforest. aagronp of Species u must and their replar 11' Among tl ”Est are An ' &o \"\e 44 forest has been done by the utilization of the land for pastures. This has created savannas, woodlands, thickets, andgnairies, which will probably never recover from their secondary growth state. Much of the mezquite savanna now under irrigation has been converted into productive sugar cane fields. The southern lowlands have been invaded by numerous large and small settlements and industrial complexes which have induced profound changes in what remained of the original deciduous vegetation. The Lower Montane Rainforest Zone This rainforest presents some characteristics of the lowland rainforest. It is distinguished by the absence of a group of species which are dominant in the lowland forest and their replacement by another complex of species. Hm typical association of the forest is the ”tabonuco type” of Wadsworth (1952a, b) in which Dacryodes excelsa is the dmfinant species. Associated tree species are Magnolia splendens, Sloanea berteriana, Cordia sulcata, Buchenavia capitata and the tree fern Cyathea arborea. The palm Euterpe globosa is scattered throughout the forest. Although a large portion of the montane forest was clear cut for coffee plantations during the past century, a few undisturbed trees give us an idea of the original Composition. Among the privileged trees within the coffee forest are Andira, Nectandra, Inga, Byrsonima, and the reduced BEE vera and The least distur host is that on the $1 Ilnqnillo Mountains . trees forming a continu slide on the understory 'oysnall trees on the f Tyrongh rocks clothed me very rare in this f The lichen flor hthis forest. The f0 hltigera and Sticta a faces. Along the nume provide many habitats concentric growth habi ialt except where the r on through. In these the soil harbors an ex Tnder this condition IT Crbrown fruits grow i By far the 1110E tomonly known as corn trees without regard htecially abundan'C ‘ itoellularia, 93513 11d Bombvlios OT?" 45 hmroduced Inga vera and Erythrina poeppigiana. The least disturbed stand of lower montane rain- forest is that on the slopes of El Verde experimental area on Luquillo Mountains. It is a dense forest with tall trees forming a continuous canopy that casts a permanent shade on the understory. This is primarily represented by small trees on the floor layer. The terrain is covered by rough rocks clothed in cryptogams. Lianas and shrubs are very rare in this forest. The lichen flora on the rocks is very characteristic in this forest. The foliaceous species of Leptogium, Peltigera and Sticta are the most common on the rock sur- faces. Along the numerous creeks large igneous boulders provide many habitats for crustose lichens that present a concentric growth habit. The soil flora is not very abun- dant except where the canopy is thin enough to let light get through. In these areas a thicket develops in which the soil harbors an extensive development of Baeomyces. Under this condition many species of Cladonia bearing red or brown fruits grow in association with Baeomyces. By far the most varied lichens belong under a group commonly known as crusts. These cover the holes of the trees without regard to any substrate specificity. Especially abundant are the genera Porina, Pyrenula, Occellularia, Chiodecton, Graphis, Pertusaria, Lecidia and Bombyliospora. Squamulose lichens belong to the fin” Phyllopsora and Q herepresented by Coccoo pmforest along the fit species of Ramalina and hme_lia and Heterodermi A different type slopes of Carite forest Susy. On these slopes AT and 80°F and the hunches. The rocks deep clays have been de hracterized by slopes toterophylla. Thu5, un original forest is diff of the original forest ylaurina, Roystonea p . . . atropholis garcmlaefc Amarked zonatf Tithe lichen flora, a? it process of species éfadient. High on the lthe lower slops 0f 333. peranceps, some the this group 0f 5 it tree trunks, two grow in 3550‘ innate 46 gmwra Phyllopsora and Crocynia. Some foliaceous species arerepresented by Coccocarpia and Pannaria. Only in the oparforest along the rivers are a few poorly developed species of Ramalina and Usnea. But in this same area Parmelia and Heterodermia are frequent. A different type of vegetation occurs on the lower Hopes of Carite forest at the eastern end of Sierra de Cayey. On these slopes the temperature fluctuates between osfidand 80°F and the annual rainfall varies from 65 to 100 inches. The rocks are of volcanic origin from which deep clays have been developed. Most of the terrain is characterized by slopes planted to Swietenia and Tabebuia heterophylla. Thus, under this managed condition the original forest is difficult to identify. Some species ofthe original forest are Ceiba pentandra, Eugenia jambos, Inga laurina, Roystonea borinquefia, Ficus leavigata and Micropholis garciniaefolia. A marked zonation is evidenced by the distribution of the lichen flora, along the tree trunks as well as in the process of species replacement following the elevation gradient. High on the tree trunks of Tabebuia heterophylla ———————_._____ mlthe lower slops of the mountains hang Ramalina dendroides ——————._._._~ and B. peranceps, sometimes reaching 20 inches in length. Below this group of species, covering the middle third of the tree trunks, two chemical variants of Ramalina com- —_.._ lanata grow in association with two chemical variants of no.3. At the 'l outerhranches, are scat horse of the trees th Andante and are repla ad the yellow fruticos hoses are covered by Pa bier lichen species in hepalm bases which ha Mspp. Higher on occupied by planted tre my boringuefia and th lichen flora is the sa exception that g ance i . dialina peranceps . St holoner mountain rair heir optimum habitats Size and high frequenc lichen vegetation at t Most of the 66 h tobacco belt. Th5 .iotnsion of second g Etriect habitats for second growth Vegetat 3tub, in associatlm 47 3.5ubasperata. At the same level, but occupying the mner branches, are scattered tufts of B. gracilis. Toward the base of the trees the Ramalina species decrease in dumdance and are replaced by a small red species of Usnea and the yellow fruticose Teloschistes flavicans. The tree bases are covered by Parmelia and small Heterodermias. In between the macro lichens many crustose species occur with— mm showing any preference for a particular substrate. Other lichen species in the Tabebuia forest are found on flw palm bases which harbor Pseudocyphellaria aurata and Collema spp. Higher on the mountains the slopes are occupied by planted trees of Swietenia, the endemic Roys- tonea boringuefia and the buttressed Ceiba pentandra. The lichen flora is the same as in the lower forest with the exception that E. anceps now appears in association with Ramalina peranceps. Still higher, at the upper limits of Um lower mountain rainforest, all these species occur in their Optimum habitats as shown by their large thallus file and high frequency. The principal trees harboring lichen vegetation at this limit are MicrOpholis and Ficus. Most of the east central part of the island comprises Um tobacco belt. This area is devoid of forest but a profusion of second growth and fence vegetation offer perfect habitats for lichen communities. Frequently the second growth vegetation consists of a low Psidium guajava scrub, in association with Rapanea, Tabebuia and Erythrina. Velichen community is hiriscoides and g. Eel hing fluids of soredia mediate species R_. su infrequent. Growing ' species we find Telosch' [5g and Heterodermia. characteristic on the d rgetation includes the cflaralina complanata lighways traversing the Spicuous for the dense myfruticose and £011 Bars are mostly Ramali “W, Teloschi W Spp‘ Where the scrul 1°33 hanging mats of I: 3911- are conspicuous 0 PL‘Iltahe Forest Zone Within this 1' Forth (1964] called 1 :Irdillera forest . '1 tire humid conditio11$ Cited three ve get at 1 48 Um lichen community is abundantly represented by Ramalina dendriscoides and g. peruviana, both characterized by hmdng mounds of soredia on the thallus. The small non- sorediate species 3. subasperata and 3. complanata also are frequent. Growing in association with the Ramalina species we find Teloschistes flavicans, many species of IBnea and Heterodermia. The lichen community is more characteristic on the dead twigs of the scrub. The fence vegetation includes the divaricatic acid containing variant of Ramalina complanata and Ramalina gracilis. Along the lfighways traversing the area the road vegetation is con— spicuous for the dense epiphytic community that harbors mmw'fruticose and foliose lichens. The lichens encountered here are mostly Ramalina peruviana, B. usnea, 3. peranceps, B.dendroides, Teloschistes flavicans, Parmelia spp. and Heterodermia spp. Where the scrub approaches the original vegetation long hanging mats of Ramalina anceps, B. usnea and Usnea Spp. are conspicuous on Buchenavia capitata. Montane Forest Zone Within this zone is included what Little and Wads- worth (1964) called the Upper Luquillo forest and the Upper Cordillera forest. The former is differentiated by its nmre humid conditions. Wadsworth (1952a) previously rec0g~ qued three vegetation types within the upper slopes of j high mocmtains: the hype" and the elfin type alias the "tabonuco ty Belong to the lower Ion cussed by Gleason and C ninfoIest. Much of th hcluded in the "montan hisereau (1966) . The montane for herto Rico, all above heLuquillo Mountains thetarite mountains in hthe central part an Sierra de Luqui easterly trade winds, the Sierra. The higheS Elioro [1074m.), El Y1 iflhSm.), The Pinnacle! hmt Britten (941W) - This mountain "her Cretaceous. In in layer of dark g1”? Rid clay, containing 335 surface. This 13 mas the soil is 1’96 Gites it is very 5“ 49 the high mountains: the palm type, the "palo colorado type" and the elfin type. The “palo colorado type" as well as the "tabonuco type" of Wadsworth (1952b), which belong to the lower montane rainforest zone, were dis- cussed by Gleason and Cook (1927) under the heading of rainforest. Much of the mossy forest of these authors is included in the “montane forest zone" delimited by Dansereau (1966). The montane forest zone covers four areas in Puerto Rico, all above 750 meters in elevation. These are the Luquillo Mountains in the eastern part of the island, the Carite mountains in the southeast, Toro Negro Reserve in the central part and, in the west, Maricao Insular Forest. Sierra de Luquillo lies in the path of the north- easterly trade winds, provoking a heavy rainfall throughout the Sierra. The highest peaks in the mountain range are El Toro (1074m.), El Yunque Rock (1065m.), El Yunque Peak (1065m.), The Pinnacles (1050m.), East Peak (1051 m.), and Mount Britton (94lm.). This mountain range is of volcanic origin from the upper Cretaceous. In most areas the soil consists of a thin layer of dark grayish-brown, or grayish—brown plastic, acid clay, containing sharp angular fragments of rocks on the surface. This layer rests on hard rock. In some areas the soil is red, permeable and deep, while in other Places it is very stoney. for this region anthose collected at L level. The mean tempera his station is 70°F, th MIT. The difference aans is 6°F. The annual preci all at least 10 inches ilril. Cloud and fog h h“MIT. which, above 505 and at night. The Carite moun total acreage of 6300 3 Slightly over 1000 m. Elll ft. is 72°F and th can and 80°F. Rainfa‘ ml 100 inches. The parental rc ilee ' l. and clay has 50 For this region the most reliable climatic records arethose collected at La Mina at about 780 m. above sea level. The mean temperature for nine years of record at fins station is 70°F, the maximum is 90°F and the minimum is 52°F. The difference between winter and summer montly means is 6°F. The annual precipitation for the area is 183 inches with at least 10 inches in every month except in March and April. Cloud and fog help in maintaining a high atmospheric humidity, which, above 650 m., reaches 90—100% in cloudy days and at night. The Carite mountains in the Sierra de Cayey have a total acreage of 6300 acres. The highest peak reaches slightly over 1000 m. The mean annual temperature above 2000 ft. is 72°F and the monthly means fluctuate between 65°F and 80°F. Rainfall varies during the year between 65 and 100 inches. The parental rock is of volcanic origin, from which a deep, acid clay has developed. The Maricao Mountain range has peaks above 850 m. The mean annual temperature is 72°F. The annual rainfall fluctuates from 74 inches on the lower slopes to 122 inches above 500 meters. The soil is serpentine in this mountain range. In the center of the island the Toro Negro Sierra Mmports the highest mountain peak on the island, Cerro de E‘nta[1338m.). The max: Iitheninimum is 65 in inches at about 750 m. tull.l°F. This zone is cha nlnfuteppe globosa. I {and at lower elevation W pubescens, Podoc nihClusia minor and Il ‘ Dansereau (1966: rainforest which accord lit the zone. These ar mad 1eaf~p31m’ the pa mum is prolonged at “this and the montane lladsn'orth (19 52 51 Pmfia (1338m.). The maximum precipitation is 125 inches multhe minimum is 65 inches, with an annual mean of 107 inches at about 750 m. The temperature varies from 59.3°F to 74.8°F. This zone is characterized by the typical sierra pahnEuterpe globosa. It contains a number of species not found at lower elevations, such as, Cyathea portoricensis, Cyathea pubescens, Podocarpus coriaceus, Didymopanax gleas- mfii,Clusia minor and Icacorea luquillensis. Dansereau (1966) named four varieties of the montane rainforest which according to him form a continuum through— mn the zone. These are the sierra broad leaf, the sierra broad leaf-palm, the palm and the moss forest. The con- tinuum is prolonged at the summit through the montane mamfls and the montane broad leaf forest. Wadsworth (1952a) distinguished the ”colorado type” wiflfln the moss forest as a separate vegetation type. Owing to the transitional character of the sierra broad leaf—palm it should not be considered a type at the same level as the others, but merely a transition zone exhibiting gradual merger of two types at their point of contact. The quuis or sclerOphyll forest seems to be a constant com~ Ponent of the upper slopes vegetation (Asprey and Robbins, 1953). Experience shows that there is no place in any of Um four areas above 750 m. where all these types occur in a series. W best type is given by sierra palm [Eu—tgpfi 3E la'ms above 600 m. Belc lovernontane rainforesI tall trees. Above this pure stands over wide a‘ he forest occurs is f0 Central it does not app amtains it is poorly Above 700 meter “dually merges into t limits it is Sharply di Tailiorestl The forest is ( lgadense canopy at 1 52 Palm Type Forest.-—An excellent description of this forest type is given by Gleason and Cook (1927). The flerra palm (Euterpe globosa) occurs in the Luquillo Moun- tahm above 600 m. Below this altitude it appears in the lower montane rainforest as scattered individuals among the tall trees. Above this elevation the palms form almost pure stands over wide areas. The lowest elevation at which the forest occurs is found in Luquillo. In the Cordillera Central it does not appear below 900 m. and in the western mountains it is poorly developed. Above 700 meters the palms form a broken belt that gradually merges into the mossy forest while at its lower Hmuts it is sharply differentiated from the lower montane rainforest. The forest is one layered with the palm trees form— ing a dense canopy at about 50—60 feet high. An understory layer is absent; no doubt as a result of the poor light available under the large crowns. The trees are widely spaced when growing in pure stand, but much less when grow- ing in a mixed condition. Trunks are straight, and 6~8 inches in diameter. The most common species in the palm forest are Euterpe globosa, Cecropia peltata, Croton poecilanthus, Alchornea latifolia and Matayba domingensis. Tree climbers are represented by Marcgravia rectiflora and M.‘sintenisii, b0th of which often cover the trees and retard their growth. Epiphytes from the lowe: at the palm forest and hebromeliads Vriesia neaostly absent. Aro seen at the edge of the Among the shrub m the rocky and wet sc ntilaginosa, P. bertei The lichen comr simpler structure than primarily represented l tmtaining blue-green ncnpy the field and s 53 Epiphytes from the lower montane rainforest extend through- mu the palm forest and are well developed here, especially Um bromeliads Vriesia and Guzmania. Vines and large lianas are mostly absent. Aroids and bamboos are occasionally seen at the edge of the forest or in old landslides. Among the shrubs, which are very poorly represented on the rocky and wet soil, the most common are Psychotria thilaginosa, P. berterina, P. maladens and Begonia decandra. The lichen community within the forest is of a much simpler structure than thepmanerogamic communities. It is primarily represented by associations of foliose lichens containing blue—green algae. Different lichen associations occupy the field and soil layer of the forest. These are prominent on the bases of trees and on the surface of rocks lying along the many brooks and waterfalls. Lichens are typically absent from the canopy layer in this forest as well as from the boles of the palm trees. The palm habit of shedding a leaf every month and thus not maintaining a permanent substrate might explain the exis— tence of such a poor lichen flora on the canopy. Some importance might also be attributed to the smooth surface 0f the leaf petiole, rendering it difficult for lichen colonization. As for the bole the reason is quite different. In the palm forest the upright position of the dorsally Channeled leaves causes a steady water flow from the crown in the hole which wash nlclecreases the chance Conspicuous in t ltictaceae represented b lseudocyphellaria aurata nsettes, often a foot i rocks and on branches 0: WW forms dark insenargins on large ' MEETS? Both 1i surfaces adapted for ga 54 dmwithe bole which washes away any incipient lichen colony and decreases the chances of any getting established. Conspicuous in the lichen community is the family Stictaceae represented by the species Sticta weigelii and Pseudocyphellaria aurata. The former forms large grayish rosettes, often a foot broad on the bases of the trees, on rocks and on branches of Cecropia peltata. Pseudocyphel— laria aurata forms dark green colonies with yellow sore- fiose margins on large branches of Tabebuia heterophylla andInga vera. Both lichen species have pitted lower mufaces adapted for gas interchange. The gelatinous lichen Leptogium is often seen growing on palms and on rocks where it appears to grow under optimum environmental conditions. Less frequent, but still conspicuous are the genera Collema, Pannaria and Coccocarpia growing on layers of nmsses and bryophytes on the bases of small shrubs along sunny trails. In open areas of the forest, where the soil is unstable, colonies of Stereocaulon in association with Um inconSpicuous crustose Baeomyces, form a podetioid layer, The following list, arranged in descending order of abundance, includes macrolichens common in the palm forest: Sticta Erioderma Pseudocyphellaria Baeomyces Leptogium Stereocaulon Collema Physcia Coccocarpia Cladonia Colorado T e F‘ the dry slopes where nmcevas not describe heluquillo Mountains : slopes where palms are luest" is seen in the bins (Dansereau, 1966) viabroad leaf forest, tree forest or a montar The dominant s; W); the spanz' Iheiorest contains twc lei'med. The canopy l fee”T0111 the ground, ‘0 ft” high. All the 55 Colorado Type Forest.—-This typical association of the dry slopes where the palm forest never gains dom- inance was not described by Gleason and Cook (1927). In the Luquillo Mountains it occurs on the summit of the slopes where palms are absent. A facies of the "colorado forest" is seen in the drier slopes of the Maricao Moun— tains (Dansereau, 1966). In this region it takes the form of a broad leaf forest, where Clusia minor either forms a tree forest or a montane scrub. The dominant species is ”palo colorado" (Cyrilla racemiflora); the spanish name refers to its red flaky bark. The forest contains two strata which not always are clearly defined. The canopy layer is uniform and never exceeds 50 feet from the ground, and the second layer is between 6 and 20 feet high. All the species in this forest are evergreen, with thick, coriaceous leaves. The trunks are short and crooked with numerous branches starting very low on the trunk. The wind effect is seen in the many trees leaning downhill. Buttresses are not common, but in the broad leaf forest prop roots are typical of Clusia. Many trees produce roots that extend for long distances on the sur— face of the soil or hang for 50 or more feet over the cliffs. Prominent species in the Colorado type are Cyrilla racemiflora, Micropholis garciniaefolia, Calycogonium Squamulosum, Ocotea spathulata, Micropholis chrysophylloides ——-—-————-——_—________ and Clusia minor. The more complex athelower limits of t lnntains supports a non nlbryophytes constitut lichens, but the majori‘ ammd places where org packets on tree trunks , snil. Some lichen gen uc conspicuous for the branches of trees. Th6 trees lying at the edgf Mill) 0 ‘ mum aDDenrs tc HZ . ed by havnr 37" the 31ga1 Componenn honed w I SPEC-16$ frp 56 The more complex structure of the'tolorado forest” atthe lower limits of the moss forest in the Luquillo annains supports a more complex lichen flora. Mosses andbryophytes constitute the habitat for many ground hchens, but the majority of the flora appear to be centered mmund places where organic matter accumulates, such as pockets on tree trunks, crevices in rocks, old stumps, and soil. Some lichen genera of the fruticose association are conspicuous for their large thalli hanging from branches of trees. These are more often seen on large trees lying at the edge of the forest or on isolated sites. The soil layer on the exposed road banks and old land slides supports typical associations of large colonies of the shrubby lichen Stereocaulon, Baeomyces and red fnflted Cladonias. Parmelia and Physcia are occasional nwmbers on loose rocks and rocky soil. On dead twigs of Cyrilla recemiflora the lichen Coenogonium appears to be most abundant. This genus is characterized by having a growth form mostly determined by the algal component (Trentepohlia). Uyenco (1963) reported 7 species from this general area. Where the big trees give way to a more sparse forest, Parmelia and Heterodermia are the dominant genera, hithe wet and shady parts of the forest, scattered colonies of podetioid lichens grow on mosses, bryophytes can the wet soil. The :::Iets and decomposed o nlCladonia wrightii ar Chile is readily recogn hereas Cladonia has a c The lichen assoc npe forest are the foil 1. Cladia-Cladonia \— organic matter Cyrilla racemif . Stereocaulon-Ba \ 2 5011 and freque Species of C1,“ 57 oron the wet soil. The association is typical of tree pockets and decomposed organic matter. Cladia aggregata and Cladonia wrightii are the only members of the association. Media is readily recognized by its perforated branches, whereas Cladonia has a continuous surface. The lichen associations recognizable in the colorado type forest are the following: 1. Cladia—Cladonia wrightii: growing on mosses and organic matter in pockets of the dominant trees Cyrilla racemiflora, Clusia rosea, and Micropholis garciniaefolia. 2. Stereocaulon-Baeomyces: abundant on the exposed soil and frequently associated with red fruited species of Cladonia. 3. Coenogonium-blue-green algae: confined to the trunk and twigs of Cyrilla racemiflora. 4. Usnea—Ramalina: hanging from trees on exposed sites in the forest. The lichen genera common to the colorado type are (a) Fruticose: Usnea, Ramalina, Coenogonium (b) Podetioid: Cladia, Cladonia, Stereocaulon, Baeomyces (c) Foliose: Parmelia, Physcia, Heterodermia Moss Forest Type.——Wadsworth and Bonnet (1951) studied the soil composition of the upper slopes of the forest in 1400111110 in SE factor in the mountain i that the dwarf Vegetati‘ of the many hurricanes l irgly named it the hurri In the Luquillo flanks of the higher 51 other types with which Cordillera it never corn in the eastern mountair and certainly the highc 50g and rain, even dur: hole 700 m. the relat: illdl days and during lithe thicket is from {m in the lower mont The annual rai muhht is rarely of a infl one “C9 of the wind 58 forest in Luquillo in search for a critical distribution factor in the mountain forest type. Murphy (1916) thought fiwt the dwarf vegetation of the moss forest is the result ofthe many hurricanes the island had suffered and accord— ingly named it the hurricane forest. In the Luquillo mountains it descends along the flanks of the higher slops and gradually merges into the other types with which it comes in contact. In the Central Cordillera it never comes down lower than 1000 meters, and hithe eastern mountains it receives the highest rainfall and certainly the highest humidity with frequent clouds, fog and rain, even during the driest part of the year. Above 700 m. the relative humidity is usually 90-100% on cloudy days and during the nights. The mean temperature hlthe thicket is from 69°F to 73°F, about 3-4 degrees lower thmrin the lower montane rainforest. The annual rainfall ranges from 100-200 inches and drought is rarely of any significance. The most important influence of the wind on the vegetation is not in creating physiological drought, but in the mechanical effect on the size and shape of the trees. The soil at higher elevations is usually saturated With water even on exposed situations. Extreme wetness gives rise to bog conditions, including growth of Sphagnum. The moss forest is a two storied community and up to 60 ft. in height. In this dwarf forest many tree I ll 4; sgecies of the lower m0! fond. However, these : haeloped than in their hits the forest trees result of their extende trade the dwarf habit is appear. The trees forrr trenches difficult to t tripping with water frr The shrub laye'. troll amount of light ' Althtupper limits of illlletely covered wit huegreen algae. \lurn fiti 1 ‘ “am 15 a conti 59 species of the lower montane rainforest are frequently found. However, these are usually shorter and less developed than in their original habitat. At the lower lhflts the forest trees are widely spaced, probably as a remflt of their extended branches. With increasing alti- tude the dwarf habit is acquired and crowded conditions appear. The trees form a network of twisted horizontal branches difficult to traverse and the trunks are always dripping with water from rainfall or condensation. The shrub layer is absent, possibly due to the small amount of light that penetrates the dense foliage. At the upper limits of the forest the tree trunks are completely covered with Selaginella krugii and a layer of blue-green algae. Numerous water holding epiphytes, orchids, aroids and ferns are well developed here. The field layer is a continuous mat of Sphagnum and other mosses. Where there is a clearing in the forest, sedges, grasses and bamboo grass cover the ground. The vines Marcgravia rectiflora and M. sintenisii grow as epiphytes on trees when young, but as they mature their roots reach the ground, from where they extend in all directions forming a network of impenetrable lianas. The most important tree species in the moss forest are Tabebuia rigida, Micropholis garciniaefolia, Clusia and Podocarpus coriaceus. Podocarpus coriaceus is an endemic restricted to the summit of Luquillo and Maricao :onntains and represents island. At the upper lil lasheen described in t lichen associations are Stereocaulon prob ably r ——_ podetia approximately 1 lodia. Foliose lichens J . . .arnelaa-Heterodernua : \ trlahebuia rigida and lartof this associati :tnposition of the a55 ““d on occasion Hetero \ Eettrod ' . w Spec1es a tilt ate type, a charac 60 nwuntains and represents the only native conifer on the island. At the upper limits of the montane zone, in what has been described in this paper as moss forest, other lichen associations are more abundant. On the ground level Stereocaulon probably reaches its maximum development with podetia approximately 10 cm. long and covered with cepha— lodia. Foliose lichens in the moss forest belong to the Parmelia—Heterodermia association, covering the mossy trunks of Tabebuia rigida and Tabebuia haemantha. Sticta is often part of this association but never gains dominance. The composition of the association varies with local factors and on occasion Heterodermia replaces Sticta. All the Heterodermia species at the limits of the forest are of the dliate type, a character that seems to be associated with heavy rainfall and low temperature. The most frequent species have a yellow tinge on the ventral surface and large Spurred fruiting bodies. The highest degree of development is reached by Um fruticose association of epiphytic lichens, Usnea— Ramalina, with thalli sometimes reaching over two feet and hanging from the high branches of Micropholis garciniae- folia and Cyrilla racemiflora. These lichens occupy the stt exposed habitats, where wind is strong and rain abun- dant. Usually Usnea is found at higher elevations than 1 r aging. The componen Eillli heavy weights and dvesllirflspecies eno ndwater weight. The inlanalina serve a sin P_annelia is prc association with M Finally, a grot nthenoss forest. I: 31%, Dictyonerna an regarded as a characte lien . are rarely Collec 61 Ramalina. The component species are well adapted to with- stand heavy weights and stresses. An elastic central cord gives Usnea species enough strength to resist wind force mm.water weight. The longitudinal cartilaginous strands in Ramalina serve a similar purpose. Barmelia is probably the most diversified genus in association with Physcia and Heterodermia. Finally, a group of rare lichens can be recognized hithe moss forest. In this group are Cora pavonia, Corella, Dictyonema and Coenogonium. This cannot be regarded as a characteristic association of lichens since they are rarely collected together. The lichen Cora is said to have a Basydiomycete as the mycobiont. The most common macro lichen genera in the moss forest are the following: Stereocaulon Physcia Baeomyces Cora Parmelia Corella Heterodermia Dictyonema Cladonia Sticta Montane Maquis.—-The most physiologically dry and exposed areas within the montane forest zone occur at the Upper limits of the zone where a scrub vegetation deveIOps. hithe western mountains where the rainfall is reduced and 1'22 soil is either serp cegetation is more evid that this type is comPc aortane naquis proper z lhenontane maquis is 2 contains spiny woody sl hpical of the scrub i. The broad leaf tion of the sierra bro habitat. The dominant m E. Egg, the Eli ' 9% some The ePiphytic n fruticose lichens ( nioschistes. Other ‘ 62 the soil is either serpentine or lateritic, this type of vegetation is more evident. Dansereau (1966) suggested that this type is composed of two different facies: the nwntane maquis proper and a montane broad leaf scrub facies. The montane maquis is a sclerophyllous community which contains spiny woody shrubs, but not succulent species. Typical of the scrub is Didymopanax gleasonii. The broad leaf scrub could be considered an expres- sion of the sierra broad leaf forest occupying an unsuitable habitat. The dominant species are woody shrubs of Clusia nunor, 9. krugii, the tree fern Cyathea pubescens, and the tree Calycogonium squamulosum. The epiphytic vegetation in the forest is abundant in fruticose lichens of the general Ramalina, Usnea and Teloschistes. Other epiphytes are bromeliads and orchids. Although Usnea and Ramalina appear to be equally common, Um genus Usnea seems to be more diversified than either Ramalina or Teloschistes. The bright yellow Teloschistes flavicans is more often collected in association with Ramalina species than With those of Usnea. It is frequent on spiny shrubs and its color acquires a whitish tone when growing under shady conditions. The genus Usnea has green, red or yellowish thalli reaching two or more feet as it hangs from trees such as Eflgig, Micropholis and Ficus. The diversity of nrphological and cherni correlated with any pa‘ Ramalina speCi occurs. The common sp dendriscoides, R. grac Thistorta. The last The soil flora alladina-Cladonia ass on areas over rocky colonies. The Cladin: the white sands of tha theCladonia species W11 apothecia. The crustose inse ' Times. On expos orange growths of Ca] a ' renery abundant in Slora he I\’ %, Buei harp . . ”1) Graphidaceae an be foun ldie 750 Ill 1‘ 1. Melt distr 63 nwrphological and Chemical characters appear not to be correlated with any particular habitat requirements. Ramalina species are very common where Usnea occurs. The common species are Ramalina complanata, 3. dendriscoides, B. gracilis, B. anceps, 3. peranceps and 3.bistorta. The last species is restricted to this area. The soil flora characteristically is composed of a Cladina-Cladonia association. This is conspicuous in open areas over rocky terrain, where it forms dense colonies. The Cladina species is the same that occurs on the white sands of the northern coastal plains. Many of the Cladonia species are of the cup forming type bearing brown apothecia. The crustose association appears to be very rich hispeCies. On exposed rocks the surface is covered by orange growths of Caloplaca. Other crustose genera that are very abundant in the forest are Haematomma, Bombylio— £2333, Lecanora, Buellia, Pyxine and many genera of the family Graphidaceae. An analysis of distribution of several lichen genera in Table 1 reveals that at least three distribution patterns can be found within the montane vegetation zone above 750 m. l. Widely distributed within the zone: Parmelia, Physcia, Cladonia, Usnea, Ramalina. 2. Restricted to a particular peak: Cladia, Cora, trailer and Coenogonium :1 the eastern end of t contains of Tom NegIC eastern range of MariCE 3. Common to two 1 this distribution is 5] ihlch are frequently fa of the island but neve m is abunda ntahsent from the ea Tah ' ‘ .le l. Distribution the montane \ Ru: Luqui] (eas1 :Etne ‘ Q “ dado ' w + no .\ + 1331' .w + "“0 onium .w + .‘ + .;fe1 g + .fidm'ces ‘ 64 Corella and Coenogonium confined to the Luquillo mountains hithe eastern end of the island; Lobaria to the central nmuntains of Toro Negro and Cladina restricted to the dry western range of Maricao. 3. Common to two regions and absent from the other. fins distribution is shown by Baeomyces and Stereocaulon which are frequently found in the eastern and central part of the island but never encountered in the western mountains. Teloschistes is abundant in the central and western parts but absent from the eastern side. Table 1. Distribution of some lichen genera within the montane forest zone at three localities. Luquillo Toro Negro Maricao Genus (east) (central) (west) Parmelia + + + Physcia + + + Cladonia + + + Usnea + + + Ramalina + + + Cladia + — — Coenogonium + — - Cora + — ~ Corella + — - Teloschistes — + + EEEEXEEE + + _ Stereocaulon + + - The distribution 0 relationship to climati empires the local envi iota indicate that as t coast,rainfall decrease lishing a trend toward hconjnnction with soi istribution shown by 1 he llontane Scrub Zone The peaks above 1' o the Cordillera Cen itthese areas the rai :tol and the soil is v in vegetation at thi fsent from the drier not the Cordillera 65 The distribution of these genera suggests a close relationship to climatic and edaphic factors. Table 2 compares the local environment in the three regions. The data indicate that as the distance increases from the east coast,rainfall decreases and temperature increases, estab— lishing a trend toward xeric conditions. These factors, hiconjunction with soil features, are responsible for the Mstribution shown by lichens. The Montane Scrub Zone The peaks above 1000 meters in the Luquillo mountains and the Cordillera Central are dominated by a scrub forest. hlthese areas the rainfall is ample, the temperature is cool and the soil is very moist. Wind has a great influence l1Pon vegetation at this elevation. The scrub forest is absent from the drier and lower mountains of the western end of the Cordillera Central at Maricao highlands. This forest has been called in part the hurricane forest by Murphy (1916) and is in part the mossy forest of Gleason and Cook (1927). It represents the elfin woodland described by Beard (1944a) for the Lesser Antilles and the mist or cloud forest of Asprey and Robbins (1953) for Jamaica. Dan- Sereau (1966) referred to it as a montane scrub because the trees are mostly below 8 meters high. Only a few semi—xerophytic species grow on the summltS Where the rocky surface is exposed. Small shrubs, OTChidS n_1_,. _,_ .Ii', _ OCNHCOE ocu Gwzuwz mCOMMOh Dogs» LO moflumflhouumhmcu pntwctic 66 om mcflwcomhom Nu OOH mww owoflymz o0 HONwom we Boa wmma Ohwoz CHOP OH HoNvom we me «Boa OHHHSUSA Taco 7:3 TE ammou pmwo mo .9509 Hamwaflma coaum>oao Eopw mommpmfim Hfiom Hmsacm can: assume new: umogwflm .odoN wmowow onmpcoa map :ngfiz macawoa moanu wo mofiumflhopomhmno HmUflmzam .N manme . the lichen 5133392 tion. Below the prom! topped and show a line of the perforated shrul 0n the high peaks of less temperate affi abundant as in Luquill reduced and the soil 1 forest is similar to t legetation is more var Itpresented by yellow “Em, and small group. Some pendent l lab ‘ ebuia are abundant an . dh. dendrordes. I‘ he Heterode rmias are hit The soil flora i 67 and the lichen Stereocaulon Virgatum constitute the vegeta~ timL Below the promontory, the trees are higher, flat- topped and show a line of wet mosses with scattered nests of the perforated shrubby lichen Cladia aggregata. On the high peaks of Cerro de Punta the vegetation is of less temperate affinities, and the mosses are not so abundant as in Luquillo. Rainfall and wind are much reduced and the soil is less rocky. The structure of the forest is similar to the Luquillo scrub but the lichen vegetation is more varied. The fruticose lichens are best represented by yellow masses of Teloschistes flavicans, Usnea 322., and small thalli of the Ramalina complanata- group. Some pendent Ramalinas on the higher trees of Tabebuia are abundant, such as B. anceps, 3. peranceps, and R. dendroides. In the branches of the Tabebuia rigida _. the Heterodermias are frequently found in association with Sticta. The soil flora is abundant and the lichen species of the sub-fruticose Baeomyces and Stereocaulon cover the exposed surfaces of road shoulders. Many species of Cladonia also grow in the wetter parts of the forest. Distribution of Ramalina To discover the distribution patterns shown by Rama- lina in the island each species was plotted on a map and Compared to a base map containing the 75 collection localities (See Figure Each map contai thin. and 600 m., and collected are indicatec In this manner and their chemical var trap was prepared for in the literature and collection examined fr species lg. montagnei h Specimen seen gave no the genus exhibits set Puerto Rico. 1. llaricao Highl shown by the single 5 68 localities (See Figure 6 and Appendix A). Each map contains two contour lines at approximately 300 m. and 600 m., and the localities where the species were collected are indicated by black dots. In this manner a map for each one of the 15 species and their chemical variants was prepared. (See Figures 7-27). A map was prepared for 3. leptosperma with one spot reported in the literature and another one corresponding to a single collection examined from the herbarium material. The species 3. montagnei has not been mapped because the only specimen seen gave no locality. Including 5. leptosperma, the genus exhibits seven major areas of concentration in Puerto Rico. 1. Maricao Highland Distribution. This pattern is shown by the single species 3. bistorta. The pattern con- tains 7 localities and in each one the species was collected at least once. Within the area the species occurs above 500 m. of elevation. The confinement of 3. bistorta to this area is not correlated with tree substrate specificity, as it occurs on trees of many different species. It is more likely that climatic factors are responsible for the restricted distribution of 3. bistorta. 2. Sierra de Cayey-Maricao Distribution. This distri— bution type is typical of g. gracilis and 3. subasperata, although their frequency decreases toward the west. R. sub» _.‘_ aSperata was collected once in Maricao and once in the l..— :orthern sandy plains. gigsubasperata is end of the Sierra de Ca acid containing popul a1 3. Central Highlar load distribution is C] “w and the . nfR_-conplanata. The film Negro Mountain lower montane rainfo re ‘ontane forest zone. r—M. . 69 northern sandy plains. The protocetraric acid variant of Ramalina subasperata is restricted to the extreme eastern end of the Sierra de Cayey at Guavate, while the salazinic acid containing population is more scattered. 3. Central Highland Distribution. The central high- land distribution is characteristic of 3. usnea, 3. farinace, R. furcellata and the divaricatic acid containing population of R. complanata. The species occurs in the general area of Toro Negro Mountains at elevations coinciding with the lower montane rainforest zone and the lower limits of the nwntane forest zone. The species are not very abundant in the region; 3. furcellata was collected only once, R. fari- nacea 4 times and R. usnea 7 times. 4. Southwestern Distribution Pattern. The southwestern type of distribution is shown by B. leptosperma. The species was reported in 1888 by Muller from near Guanica and later was collected by Britton in Bahia Sucia in the same general area. This area is the driest and represents the most xeric conditions in the island. 5. East—West Highland Distribution. This area com— prises the four highland areas containing the Luquillo Mountains, Sierra de Cayey, Toro Negro Mountains and Mari— cao Mountains. The species with this type of distribution are 3. anceps, B. dendroides, 3. dendriscoides and 3. pg:- anceps. The four species were collected in the same local— ity most of the times. The restriction to such areas suggests that the spec mdtenperature factor the eastern part of th frequency and abundanc h. North-Central ligand 3. sorediosa pattern involves the 1 lands, coinciding witl louernontane rainfor ligis abundant in mfmillency and abun "title the reverse is that the central pois climatic and topogral he occurrence of the . Scattered Di: inhution is shown b‘ the distribution of 70 suggests that the species respond to altitudinal, moisture and temperature factors. However, excess of moisture in the eastern part of the island appears to limit both the frequency and abundance of the species. 6. North-Central Distribution Pattern. Only R. 2333— viana and R. sorediosa have a north-central range. This pattern involves the northern lowland and the central high— lands, coinciding with the former lowland rainforest and lower montane rainforest vegetation ranges. Ramalina peru— viana is abundant in the central highlands and decreases in frequency and abundance toward the northern lowlands, were the reverse is true for E. sorediosa. It appears that the central poisition of the range is determined by climatic and topographic factors with elevation influencing the occurrence of the species within their range. 7. Scattered Distribution Pattern. This type of dis- tribution is shown by g. complanata and 3. subpellucida. The distribution of 5. complanata is better demonstrated by the salazinic acid containing population of the species. Although the species exhibit a general distribution, it is noted that they have been rarely collected in the southern part of the island. It is probable that the aridity of the environment constitutes a barrier to the SPreading of the lichens. General trends set prim-fly by rain ters in the island. correlation between 1: there are only 10 to 1 between the warmest a! hnost cases, howeve‘ liliting factor for t? of plants. This rela that exists between p The vegetatic determined by climate Exception to this is the mangrove and the olnangrove and the hrackish water and t character stay uand climates. Lichen veget tented by crustaceo wag spread ov ‘h the swamp. The seasona Wit of rainfall, Alters-ed by areas 1 71 Summary General trends in vegetation follow the pattern set primarily by rainfall and secondarily by soil charac— ters in the island. Roberts (1942) found very little correlation between temperature and plant distribution, as there are only 10 to 15 degrees of maximum difference between the warmest and the coolest spot on the island. In most cases, however, the amount of rainfall is the limiting factor for the distribution of many assemblages of plants. This relation is in reality the correlation that exists between plants and soil water holding ability. The vegetation types of the island are predominantly determined by climate rather than by edaphic conditions. Exception to this is the littoral vegetation that includes the mangrove and the swamp vegetation. The four species of mangrove and the Pterocarpus swamp are associated with brackish water and their physiognomic and floristic character stay unmodified even through belts of different climates. Lichen vegetation in the mangrove is poorly repre— sented by crustaceous colonies, while large foliaceous Parmelias spread over the roots of the Pterocarpus trees in the swamp. The seasonal evergreen forest receives a substantial amount of rainfall, in many cases equivalent to that received by areas lying within the lowland rainforest zone, inlbecause of this so dim status. The ‘50 soil that promotes phy shallow soil deficien Epiphytic veg the genera Parmelia, seated. The semi-decic rich flat limestone wl htreromorphic nature forest is due primari the year. Thus, the erergreen forest can his is especially tr the top of the limest Sui-deciduous forest It follows t ecergreen, and decid 511 or environmental he also related flo ilident as the xero itpe is characteriz reStricted to the t) Situation is well i 72 and because of this some doubts remain about its climatic climax status. The forest is associated with a type of soil that promotes physiological drought as a result of shallow soil deficient in root room and water retention. Epiphytic vegetation is abundant and lichens of Hm genera Parmelia, Physcia and Ramalina are well repre- sented. The semi-deciduous forest is partially associated with flat limestone where the drought is more effective. But xeromorphic nature of the vegetation in the southern forest is due primarily to poor precipitation throughout Um year. Thus, the semi-deciduous forest and the seasonal evergreen forest can be distinguished by edaphic factors. Nfis is especially true when considering the vegetation at the top of the limestone hills. The common lichen in the semi-deciduous forest belongs to the Pyrenopsidaceae. It follows then, that the littoral, seasonal— evergreen, and deciduous zones are the result of physilogi— cal or environmental drought. The three vegetation types are also related floristically although this is not as evident as the xeromorphism of the species. Each vegetation type is characterized by a core of species which is not restricted to the type but it is dominant in it. This situation is well illustrated by a xeromorphic complex of Species which supplies the dominant species for the three types (Dansereau, 1966). Again the Cecropia forest on the rhinestone hills carrie hinforest and the non principle restated by gates upon contact wi into species cores fo As we approac coast the drought effe the central mountain r the on the north coa: sountains of the islac henontane vegetatic: hgaltitudinal seque brake, moss forest an climate is well drain thenoss forest is re lichaands. That the Vegetation is evi den “Posed slopes where zen territory to col Present evidence th iledaphic factor w firect influence of The montane aigetation because , 73 limestone hills carries representatives of the lowland rainforest and the montane rainforest. This confirms the principle restated by Dansereau (1962) that species aggre- gates upon contact with a different climatic zone break huo species cores for different plant types. As we approach inland from the north or the south coast the drought effects are less efective as one reaches flm central mountain range. These effects are less notice- able on the north coast than on the south coast. On the nwuntains of the island the hygrophytic communities develop. The montane vegetation generally is arranged in the follow— ing altitudinal sequence: lower montane rainforest, palm brake, moss forest and elfin woodland. If the montane climate is well drained and the soil is serpentine, then the moss forest is replaced by broad leaf montane scrub or a montane scrub. The first sequence is typical of the Luquillo Mountains and the second is typical of the Maricao highlands. That the palm brake is a subclimax type of Vegetation is evidenced by the tendency to invade the exposed slopes where hurricanes and landslips have opened new territory to colonization. Wadsworth and Bonnet (1951) present evidence that the montane thicket is the result of an edaphic factor which is more preponderant than the direct influence of rainfall. The montane scrub zone supports a climatic climax Vegetation because, although Beard (1949) thinks it is not .—-:corre1ated with soil, denuntains is a res operate. The vegetation tinuum of succession series under more favo faster rate of replace latter represents a 5] environmental conditic the center of the isl: lowland succession se forest vegetation. T {1927) approach to th Not much is g ieneral characteristi island since there is it recover. Further tion demands more an hauls. This suggest 5iscussed represent deflation than of t growths, which appea 5:11P Changes, is 1e Silt forests that o The lichen 335” Vegetation i 74 correlated with soil, the poor and scarce soil on top of the mountains is a result of heavy rainfall and wind exposure. The vegetation of the island constitutes a con- tinuum of succession at two different levels: the northern series under more favorable conditions appear to work at a faster rate of replacement than the southern series. The latter represents a slow rate of succession due to adverse environmental conditions. The optimum conditions toward the center of the island represent the ideal place for the lowland succession series to meet and culminate in the rain— forest vegetation. This is in synthesis Gleason and Cook's 0927) approach to the vegetation of the island. Not much is gained by trying to discover the general characteristics of the original vegetation of the island since there is little opportunity for the vegetation to recover. Furthermore, the constantly increasing populae tion demands more and more of the remaining semi-forested lands. This suggests that the vegetation zones already discussed represent a truer picture of the high montane vegetation than of the lowland vegetation. Only secondary growths, which appear and disappear as fast as land owner- ship changes, is left to indicate the nature of the exten- sive forests that once covered the island. The lichen vegetation is no different from the higher vegetation in its floristic relationships. Groups Toflichem genera show .inmtions. AS a gene attains mainly the so less are mainly gela [anon among these ar (fl, Pseudocyphell the dry mountain zones sented by well develon @1113 attaining up land, soil lichens, e: (Cladonia, Cladina) a “ growths of the lowlan W are foun The genus RE patterns throughout t l 1152M; (2) Si TW); (3) Callata, B. farinace 75 oflichen genera show certain affinities to definite plant formations. As a general rule the rainforest of Luquillo contains mainly the so called less evolved lichen genera. These are mainly gelatinous and partly lichenized algae. Cmmwn among these are Peltigera, Sticta, Leptogium, Cocco- carpia, Pseudocyphellaria, Coenogonium and Dyctionema. On the dry mountain zones the lichen flora is typically repre- sented by well developed fruticose species of Usnea and Ramalina attaining up to 4 feet in length. On the other hand, soil lichens, especially the podetiod growth form (Cladonia, Cladina) are well developed in the secondary growths of the lowland forest sites, while Cladia and Stereocaulon are found in the wet mossy forest. The genus Ramalina exhibits seven distribution patterns throughout the island: (1) Maricao-highland (3.bistorta); (2) Sierra de Cayey-Maricao (3. gracilis, 3.5ubasperata); (3) central-highland (B. usnea, 3. £33— cellata, g. farinacea); (4) southwestern-lowland (3. lepto~ Sperma); (5) east-west—highland (E. anceps, B. dendroides, R. dendriscoides, R. peranceps); (6) north—central (3. peruviana, R. sorediosa); and (7) scattered (B. complanata, 3.5ubpellucida). BIOGEOGI OF THE Floristic “— 1‘ . .nlnoduction Many biogeogr tions between Cuba ar andfnom Cuba to Honc nnly the land bridge CHAPTER IV BIOGEOGRAPHICAL AFFINITIES OF THE WEST INDIAN FLORA Floristic Elements of Vascular Plants Introduction Many biogeographers have postulated land connec— tions between Cuba and Central America, from Cuba to Yucatan, and from Cuba to Honduras. Schuchert (1935) believed that only the land bridge to Honduras actually existed. These land connections would certainly help explain the strong affinity between the flora of the Great Antilles and that of Central America. The land bridge hypothesis of— fers an explanation to the high degree of endemism in Cuba and the awkward position of Jamaica with respect to many genera that either bypass the island or else have a disjunct distribution with continental land.. The great affinity be— tween the floras of Cuba, Hispaniola and Puerto Rico and their poor relation to that of Jamaica is probably due to the fact that while Jamaica was submerged, these islands harbored refugia which later constituted new centers of distribution. 76 to land bridge llonida or between Sot lend (1949), however extended into the Vic: Land connectil role in the plant dis nselsewhere, but oth neatarole. Wind a buds and land rafts Man as from one cc The problem ( recognized by Anthonl Anette Rico. lest Indian Element This is rega n all the Caribbean :nnst, those species inneater Antillean 77 No land bridge has been postulated between Cuba and Florida or between South America and the Lesser Antilles. Beard (1949), however, suggests that Puerto Rico probably extended into the vicinity of Antigua. Land connections undoubtedly have played a major role in the plant distribution of the West Indies as well as elsewhere, but other means also have probably had as great a role. Wind and sea currents, hurricanes, migratory birds and land rafts can account for distribution of plants as far as from one continent to another. The problem of distribution in the West Indies was recognized by Anthony (1925) while studying the mammals of Puerto Rico. West Indian Element This is regarded as the floristic component shared by all the Caribbean Islands. It presents two aspects; first, those species common to all the islands, and second a Greater Antillean element more or less distinct from a Lesser Antillean element. It is agreed among students of the West Indian flora that except for the littoral zone the greatest simi— larity occurs in the seasonal and montane formations. With regards to the distribution of palms, Ciferri (1936) re— ported that the greatest affinities are between Cuba and lfispaniola, with much less affinity between Hispaniola and L...) Euento Rico, and almos In general the endemil latedto those of Cub: least to those of Jam In looking at htilles, we find the tuba extends into the whole flora is more I honest of Cuba. It the eastern part of t lithe flora of Pueri nes between the ext‘ 78 Puerto Rico, and almost none between Cuba and Puerto Rico. hngeneral the endemic plants of Hispaniola are more re- lated to those of Cuba, less to those of Puerto Rico and least to those of Jamaica. In looking at the geographical areas of the Greater Antilles, we find that the vegetation of extreme eastern Cuba extends into the western part of Hispaniola and the whole flora is more related to that of Hispaniola than to the rest of Cuba. The flora of the Samana Peninsula in the eastern part of Hispaniola appears as a prolongation of the flora of Puerto Rico. Less marked are the affini— ties between the extreme eastern parts of Hispaniola and the island of Jamaica. Fawcett and Rendle (1910), analyz- ing the orchids of Jamaica, found more affinities with Cuba and less with Puerto Rico. The flora of the Lesser Antilles is very homogenous and shows some ramifications northward into the Greater Antilles and southward to Venezuela. Beard (1949) found that the species of the dry zone flora and the montane flora of Puerto Rico are the most apt to appear in the Lesser Antilles islands. Both Beard (1949) in regard to the tree species and Vélez (1957) in regard to the herba- Ceous species of angiosperms found that the islands of the Lesser Antilles are very homogeneous in their plant species. bionic Element ——’—'——— The endemic e exaggerated in early of the islands became denies, although subs communities of Cuba a high degree of endemi endemic genera from E and Sierra de NiPe~ shown by such familil Elphorbiaceae, Urtic homatotal of 757 found 319 endemic SP Urban (1923- ‘he flowering plants endemic ‘ s,1.e., 33.Q v Asprey and F 3P fi'n't ~CleS in Jamaica c it and Rendle (' ilotal of 194 spec n3~ ‘ ,eld 90 species of «5 equivalent to 14 fen ' ~ demnc flowerin filliast 79 Endemic Element The endemic element in the West Indies has been exaggerated in early studies (see Table 3). As the flora of the islands became better known the percentage of en— demics, although substantial, has been reduced. The plant communities of Cuba are well recognized as harboring a high degree of endemism. Carabia (1942; 1945b) listed 16 endemic genera from Sierra de los Organos, Sierra Maestra and Sierra de Nipe. Urban (1923) noted the high endemism shown by such families as Leguminosae, Compositae, Euphorbiaceae, Urticaceae, Piperaceae and Aristolochiaceae. From a total of 757 species included in these families, he found 319 endemic species, i.e., 42.14 percent. Urban (1923—1928, V. 9, p. 39) estimated in 3088 the flowering plants of Hispaniola, of which 1048 were endemics, i.e., 33.93 percent. Asprey and Robbins (1953) recognized 440 endemic species in Jamaica out of a total 969 indigenous plants. Fawett and Rendle (1910) listed 73 endemic orchids out of a total of 194 species examined. The ferns of Jamaica Yield 90 species of endemics (Proctor, 1953); this figure is equivalent to 14% of the Pteridophytes. The percent 0f endemic flowering plants in Jamaica approaches 20% in contrast to Beard's estimation of 7% of endemics in Trinidad, with a total flora of 2200 species, and 13% in both Bahamas and Puerto Rico. Table 3. Percent of l l\) I Table 3. Percent Island CUBA (Urban, 1923- 1928) HISPANIOLA (Urban, 1923- 1928) JAMAICA (Fawcett and Rendle, 1910) (Asprey and Robbins, 1953) PUERTO RICO (Little et a1., 1964) (Dansereau, 1966) 80 of endemics in the Greater Antilles. No. Species 757 Angiosperms 3088 Angiosperms 194 Orchids 969 Indigenous plants 250 Tree species 743 Vascular species - Percent No. Endemlcs Endemics 319 42.14 1048 33.93 73 37.63 440 45.40 29 11.60 107 14.40 The endemic Sl from Puerto Rico 1370 out of a total of aPP percent. The flora of non species of HOWE and Quentin (1937) es plants were endemic, French Antilles, i.e endemics Martinique l are common to both i inthe floras of the 81 The endemic species of vascular plants reported from Puerto Rico by Dansereau (1966) comprise 107 species out of a total of approximately 743, i.e., about 14.40 percent. The flora of the Lesser Antilles comprises about 2000 species of flowering plants (Beard, 1949). Stehle and Quentin (1937) estimated that 165 species of flowering plants were endemic, from a total of 1700 species in the French Antilles, i.e., 9.8 percent. From this number of endemics Martinique has 58 species, Guadalupe 68 and 21 are common to both islands. Endemism is especially high hithe floras of the rainforest and montane forest in Um Lesser Antilles. Beard (1949) concluded from a study of 243 tree species that the swamp flora harbors no en- demics, the dry zone flora 12%, the rainforest flora 42% and the montane forest flora 52% of endemic trees, to make a grand total of 28% endemics out of 243 identified species. In Table 4 the trees have been listed under four sub—floras. Table 4. Percent of endemics in each of four sub—floras in the Lesser Antilles (Beard, 1947). W. Wm“ No. of No. of Percent of species endemics endemics __.__________~____________________.__________________ Swamp Flora 6 0 0 Dry Zone Flora 112 13 12.0 Rainforest Flora 102 43 42.0 Montane Flora 23 12 52-0 Total 243 68 28.0 '(e'le: (1957} in a 5t“ lesser Antilles found complex process that of relationships betw ciation and the time tontimental Element Asprey and Re ofthe Jamaican flor. is strongly Central . had been expressed b study of the Cuban f iffuba showed predo herica, less with C henica. Carabia (1 MeafllC 1101‘ \r h olcanic 9. south American l and a strong flo rufflCd as well as 82 Velez (1957) in a study of the herbaceous species of the Lesser Antilles found that endemism in the islands is a complex process that has to be understood in the light of relationships between rate of migration, rate of spe- ciation and the time factor. Continental Element Asprey and Robbins (1953) concluded from a study of the Jamaican flora that the flora of the Greater Antilles is strongly Central American in origin. A different view had been expressed by Seifriz (1943) as a result of his study of the Cuban flora. Seifriz thought that the flora of Cuba showed predominant affinities to the flora of South America, less with Central America and much less with North America. Carabia (1945b) suggested that Cuba is neither oceanic nor volcanic but a portion of the northern part of the South American continent and for this reason its flora showed a strong floristic resemblance to northern South America as well as to the rest of the Greater Antilles. The Cuban situation is especially unique because it lies near both Florida and Yucatan and still its major relation is with South America. In Pinar del Rio Province (Cuba) the vegetation takes a temperate aspect, as shown by the presence of guercus and Pinus. These two genera are not uncommon in the tropics, both occur in Hispaniola and Central America. n. Cuban oak is 119 hsoutheastem Unit species, including 0 my cannon to south Hispaniola. Still a Florida is the royal not only true for Cu lntilles as well. The relation sole is less underst ecer, points to strO two areas. A numbe lenezuelan ancestry finds in the flora c distributed in the l Honduras as another strong West Indian l The florist ismanifested by th‘ fanny. Swallen (1 li grasses common t all that a total 0‘5 l'lcatan. Taking '5 clude that the £10 fcxito and llucata'ln iii Yucatan. 83 The Cuban oak is Quercus virginiana, the same that occurs hisoutheastern United States. The pines are of several species, including one endemic species; two species are very common to southern Florida and one is represented in lfispaniola. Still another plant abundant in Cuba and Florida is the royal palm, Roystonea. This affinity is not only true for Cuba but for the rest of the Greater Antilles as well. The relationship between Cuba and Yucatan penin- sula is less understood. The proximity of the lands, how- ever, points to strong ties between the vegetation of the two areas. A number of species also point to a Colombian— Venezuelan ancestry of the Cuban flora. Standley (1930) finds in the flora of Yucatan many species which are widely distributed in the West Indies. He also points to British Honduras as another country in Central America with a strong West Indian element. The floristic relationship between Yucatan and Cuba is manifested by the distribution pattern of the grass family. Swallen (1934) showed that only 6 of the 87 species of grasses common to the West Indies are not found in Cuba and that a total of 89 species are common to Mexico and Yucatan. Taking the two figures together we have to con- clude that the floral relationship in the grasses between Mexico and Yucatan is not greater than that between Cuba and Yucatan. Asprey and R continental element several genera. The com. Jeanna. P llpllostylon, Salix the authors clearly Cuba, the first two reach Jamaica or th species of pines in Hispaniola. Salix , but the last two ar eral the northern e proceeds eastward i The Central rlpresented by l_)_i_dy his group of gener lesser Antilles. The South A llassess. The ans 33“.” Species which and Jamaican or to l d‘atlaica. The palm Southern species V. Plento Rico and Ce if the swamp tree 533mm in the Less 84 Asprey and Robbins (1953) studied the northern continental element in the West Indies in relation to several genera. The common northern elements are: Magnolia, Juglans, Pinus caribaea, Pinus occidentale, Phyllostylon, Salix and Quercus. The data presented by the authors clearly show that these genera are found in Cuba, the first two genera reach Puerto Rico and none reach Jamaica or the Lesser Antilles. There are four species of pines in Cuba but only one is present in Hispaniola. Salix, Phyllostylon and Quercus reach Cuba, but the last two are represented in Hispaniola. In gen- eral the northern element decreases in influence as one proceeds eastward in the West Indies. The Central American element in this analysis is represented by Didymopanax, Bombax, Lysiloma, and Brosimum. This group 0f genera is common in both the Greater and Lesser Antilles. The South American element is the most difficult to assess. The analysis of the flora shows that there are many species which are either common to the Lesser Antilles and Jamaica or to the entire Antillean region but not to Jamaica. The palm Euterpe globosa is an example of a southern species very prominent in the Lesser Antilles, Puerto Rico and Central America. The same trend is shown by the swamp tree Pterocarpus officinalis which is very COmmon in the Lesser Antilles, reaches Puerto Rico, 'dispaniola and Jamai epic of a strictly inTrinidad but not Puerto Rico. Howev Finland, and from Tr‘ laIaica. Other gen are of southern ori route, completing t The tertiar Hollick (1928) and Hispaniola (Hollick that the tertiary f ent day flora and t show primarily an a continental affinit Urban (1923 occurring in Hispar Antilles, indicati! the South American continent. The di: 1? species in comm Cannon with contin Common with Centra South America. The expla not easy and an a men some type 0 85 Hispaniola and Jamaica, but never gets into Cuba. An SX‘ ample of a strictly South American genus is Clethra, found in Trinidad but not recorded from the Lesser Antilles or Puerto Rico. However, it extends north to Mexico, via Panama, and from Trinidad extends westward to Cuba and Jamaica. Other genera, such as Podocarpus and Dacryodes, are of southern origin and follow the Lesser Antillean route, completing the arc in Yucatan or Puerto Rico. The tertiary flora of Puerto Rico was studied by Hollick (1928) and the same author analyzed that of ffispaniola (Hollick, 1924). In both cases he concluded that the tertiary flora is basically similar to the pres- ent day flora and that the majority of the tertiary genera show primarily an antillean affinity and secondarily a continental affinity. Urban (1923-1928) reported a number of species occurring in Hispaniola which are absent from the other Antilles, indicating that the affinities are stronger with the South American flora than with the Central or northern continent. The distribution of species is as follows: 17 species in common with North America, 23 species in Common with continental Tropical America, 12 species in common with Central America and 32 species in common with South America. The explanation of this distribution pattern is not easy and an analysis of the species probably will un- cover some type of relationship to the early geological he been associated tron and Steere (195 tion of the mosses clian region, the M of South America. flora is absent fro cially from Puerto The flora 0 but with some conti ported 70 species w item and Floridan a northern South Ame he listed. It has did not reach Puert Pterocarpus and §_11_1 It has been flora of Trinidad : would be anticipatl flora of the Lesse herican element w influence of the G The closest affini flora. 86 history of the island. The moss flora of the West Indies has been associated with that of Venezuela and Brazil. Crum and Steere (1958), however, noted a strong affilia- tion of the mosses to continental floras of the Appala— chian region, the Mexican plateau and the Andean region of South America. The vascular plant element of this flora is absent from most parts of the Antilles, espe- cially from Puerto Rico and the Lesser Antilles. The flora of Puerto Rico is predominantly Antillean but with some continental elements. Dansereau (1966) re— ported 70 species with Antillean, continental Central Amer— ican and Floridan affinities and 78 species with a Caribbean- northern South American distribution out of 743 species which he listed. It has been noted before, that the pines and oaks did not reach Puerto Rico from the north, but that Dacryodes, Pterocarpus and Euterpe reach the island from the south. It has been stated by Beard (1946, 1949) that the flora of Trinidad and Tobago is continental in origin, as would be anticipated from their continental nature. The flora of the Lesser Antilles shows a predominant South American element which decreases toward the west, while the influence of the Greater Antilles decreases toward the east, The closest affinities appear to be with the Puerto Rican flora. ”lone olitan Element The cosmPO1 introduced and “at“ luring the Peri0d of plants such as £13111 from Asia. Others ‘hacendados." Gras andsome shade tree The cosmopo road side and garde Dis 1 The living is characterized b)’ teal America (excep distributed, (3) 3‘ evolution (Darling1 The orderll fauna is illustrat‘ in the islands, of islands; ELIE, is not in Puerto Rico into but not in J Puerto Rico but It The mama and insectivol‘es 87 Cosmopolitan Element The cosmopolitan element is represented by species introduced and naturalized in the West Indies, especially during the period of colonization. Many of these are crop plants such as fruit trees brought from the East Indies or from Asia. Others are ornamentals introduced by the early ”hacendados." Grasses have been introduced from Europe and some shade trees have come from South America. The cosmopolitan element is typical of the shore, road side and garden communities. Distribution of Animals in the West Indies The living and extinct fauna of the Greater Antilles is characterized by being (1) principally derived from Gen- tral America (except for birds), (2) homogeneous and orderly distributed, (3) strikingly poor and (4) result of radiative evolution (Darlington, 1938). The orderly distribution of the Greater Antilles fauna is illustrated by the presence of five genera of frogs in the islands, of which Eleutherodactylus occurs in all the islands; EXlE» is found in Cuba, Hispaniola and Jamaica but not in Puerto Rico; Bufg, in Cuba, Hispaniola and Puerto Rico but not in Jamaica; Leptodactylus in Hispaniola and Puerto Rico but not in Cuba and Jamaica. The mammals are also orderly distributed: sloths and insectivores are known from Cuba, Hispaniola, and Puerto Rico but not from J8! occur in all the lat agreement with Darli oceanic nature of th in which mammals hav gestive of an accumu idue of a continenta sectivores came orig and rodents from Soc Absent from artiodactyls, antea‘ insectivores. 0f m. suP1318, anteaters, through a long evol' 88 Rico but not from Jamaica. Rodents are more numerous and occur in all the larger islands. Carlquist (1965), in agreement with Darlington (1938), found evidence of the oceanic nature of the Greater Antilles in the random way in which mammals have been introduced. The fauna is sug- gestive of an accumulation of immigrants and not the res- idue of a continental fauna. It is probable that the in- sectivores came originally from North America and the sloths and rodents from South America. Absent from the Greater Antilles are perissodactyls, artiodactyls, anteaters, ungulates, brobocidians and modern insectivores. Of more significance is the absence of mar~ supials, anteaters, armadillos and monkeys, which have gone through a long evolution. It is unlikely that land bridges either with North or South America would have allowed such a meager fauna. Bats, in contrast to terrestial mammals are rela— tiVely numerous and diversified (Anthony, 1925) as are also birds (Wetmore, 1927; Biaggi, 1970). Distribution of the Lichen Genus Ramalina It is apparent that the distribution of Ramalina in the West Indies is influenced by the nature and position 0f the islands, climate and topography. Whether the island is calcareous, volcanic or oceanic in origin has much to do With the type of vegetation that it can support. On the other hand the 011m"1 are often limiting 1 plant communities. important distributi The position trade winds and hurr dispersal of lichen imaNE-SW directior island at the extrer ihe SE-NW direction hand provides a mea: lands of the archip the Lesser Antilles In looking 33mm, bats, bird Cannot be ignored. 89 other hand the outcome of climatic and edaphic conditions are often limiting in providing suitable habitats for plant communities. Topography and land relief are also important distributional factors. The position of the island in relation to the trade winds and hurricanes path is closely related to the dispersal of lichen diaspores. The trade winds blowing in a NE-SW direction favor the distribution from island to island at the extreme ends of the Lesser Antillean arc. The SE-NW direction of the hurricane winds on the other hand provides a means of dispersal among the central is— lands of the archipelago and between the lower third of the Lesser Antilles. In looking for dispersal factors the influence of mammals, bats, birds, man, ocean currents, and life rafts cannot be ignored. The distribution of Ramalina presented in Table 14 has been examined in the light of these facts in search of some generalities. Unlike flowering plants and animals Ramalina does not show a Greater Antilles element more or less distinct from the Lesser Antilles. Out of the 23 species present in the West Indies 9 species are common to both the large and small island groups. These include Ramalina anceps, g. dendriscoides, 3. furcellata, 3. gracilis, R. dendroides, 3. peranceps, 3.5ubpellucida and 3. usnea. Of this group only 3. anceps has a continuous dis ohile R_. dendroides , t. W are common South American conti treme of the chain. Dominica, and Guadai other in the path 0: only the species wi Six species FEW} R0 SL1 erecommon to the i 90 has a continuous distribution throughout the West Indies, while R. dendroides, R. dendriscoides, R. peranceps and R. usnea are common to both the outlying islands of the South American continent and Puerto Rico in the other ex- treme of the chain. The larger islands of Martinique, Dominica, and Guadaloupe, which lie relatively near each other in the path of the hurricane winds, have in common only the species with a general distribution. Six species (Ramalina complanata, R. leptosperma, R. montagnei, R. subpellucida, R, peruviana and R. usnea) are common to the islands and Florida. Ramalina peruviana probably has the northern limit of its range in the ever- glades of Florida. Four species from Florida (R. willeyi, R. stenospora, R. paludosa and R. fastigiata were never present in the West Indies. Common to the West Indies and Central America, especially Yucatan, are Ramalina complanata, IFU dendriscoides, R. usnea, R. sorediantha, R. peruviana, 3. sorediosa, R. subpellucida and R. dendroides. The West Indian species represented in South America are: Ramalina complanata, R. scrobiculata, R. sorediantha, R. peruviana, R. farinacea, R. cumanensis, R. sorediosa and R. usnea. A total of 41 chemical variants were identified from the West Indies and Florida material, of which only the divaricatic acid containing variants of Ramalina dendroides and R. furcellata are restricted to the Lesser Antilles. Three substances found in Florida species of inalina were never ocid, 4-0-demethylbz The distribo shows the following 1) A tende: the extremes of so many species the Lesser Anti of which size a tent. 2) h deCI‘E the eastern par 3) A greao 4) A tendc sified both ch. than in Other 91 Ramalina were never present in the West Indies (perlatolic acid, 4-O—demethylbarbatic acid and stenosporic acid). The distribution of Ramalina in the West Indies shows the following characteristics: 1) A tendency for the species to concentrate at the extremes of the chain of islands. The absence of so many species in the islands toward the center of the Lesser Antilles is a function of diverse factors of which size and poor vegetation are the most impor- tant. 2) A decrease in the number of species toward the eastern part of West Indies islands. 3) A greater affinity between Cuba, Hispaniola and Jamaica than between these and Puerto Rico. 4) A tendency for the genus to be more diver- sified both chemically and morphologically in Jamaica than in other islands. 5) A greater floristic affinity with Central and South America than with southern United States. Summary The phanerogamic flora of the West Indies contains a West Indian element shared by all the Caribbean islands, an endemic element which is characteristics of each indi— vidual island, a continental element that shows more affin- ities with Central and South America than with North America, coda subcosrnopolita distribution, eSp¢Cj regions of the worlc The fauna of uted, highly homoge: high degree of ende: continental or ocea evidences. The poo explained by over-w ‘ohetical land conne The same fa Nation of higher the dispersion of I “WY transport 1 ““15 distances. Tl sented in the Grea srhred, both chemi 92 and a subcosmopolitan element of more or less world wide distribution, especially in the tropical and temperate regions of the world. The fauna of the West Indies is orderly distrib- uted, highly homogeneous, and depauperate but showing a high degree of endemism. The islands have been considered continental or oceanic by various authors on distributional evidences. The poor fauna of the Greater Antilles could be explained by over—water dispersal without assuming hypo— thetical land connections. The same factors responsible for over water trans— portation of higher plants and animals are responsible for the dispersion of lichens. Hurricane force winds can pre- sumably transport lichen thalli and diaspores over enor- mous distances. The lichen genus Ramalina is best repre— sented in the Greater Antilles where it is greatly diver- sified, both chemically and morphologically. The genus R (1810, p. 122 and 1) previously placed b of the genus Firing tinguished the new cartilaginous thall llontagne (l Ramalina under the Parmeliaceas and e: include those Rama chanical support '1 Massalongo @1135 inanis at and hyphae running Nylander the tribe Ramalin CHAPTER v THE GENUS RAMALINA ACH. History of the Genus The genus Ramalina was established by Acharius (1810, p. 122 and pp. 598-609) to include ten species previously placed by him in the sub—division Polymeria of the genus Parmelia (Acharius, 1803). Acharius dis— tinguished the new genus Ramalina from Parmelia by its cartilaginous thallus and apothecial characters. Montagne (1852) placed the genera Usnea and Eiflgliga under the sub—tribe Usneas in the tribe Parmeliaceas and established the genus Desmazieria to include those Ramalinas (e.g., R. homalea) without me— Chanical support in the cortex and with a cottony medulla. Massalongo (1854) based the genus Cenozosia on BEEELLEQ inanis and characterized with a fistulose thallus and hthae running perpendicular to the surface. NYlander (1858—60) placed the genus Ramalina in the tribe Ramalinei, which belonged in the series Ramalodei. 93 The anatomio Trevisan (1861) to : (=llsmeaceae Eschw.) halineae Ag. Thi: nov., Evernieae Ha nphistema type of coins algae in all hesmazieria Mont. 1 ~—____. the tribe Evernieae acortical layer f0 Ramalimeae had a to compact filamentous to sub-tribe Cornio Trev. in which the Stizenbergo marieria to includo Euramalina (= sect ties with a double The first Xylander in 1870. utilized the colot groups of species and represented b liornia; Group B, \K_ 1 _: The autl "“15 recommender 94 The anatomical structure of the thallus led Trevisan (1861) to subdivide the family Usneinae Trev. (=Usneaceae Eschw.) into six tribes, one of which was Ramalineae Ag. This tribe shared with Neuropogoneae Trev., Evernieae Mass., and Roccelleae Mass. the amphistema type of thallus in which the medulla con— tains algae in all sides. Trevisan included the genus Desmazieria Mont. in the sub—tribe Euevernieae Trev. of the tribe Evernieae, lacking a central axis layer and a cortical layer formed by filamentous hyphae. The tribe Ramalineae had a cortical layer formed by more or less compact filamentous hyphae. The tribe was subdivided in— to sub-tribe Cornicularieae Hook. and sub—tribe Euramalinese Trev. in which the genus Ramalina was placed. Stizenberger (1862) proposed Ramalina sect. Des— mazieria to include species with simple cortex and sect. Euramalina (= sect. Bitectae Stein.) to include those spe~ cies with a double cortex. The first extensive study on the genus was that of Nylander in 1870. For primary divisions of the genus he utilized the color of the pycnidia and recognized three groups of species: Group A, with totally black pycnidia and represented by 7 species, mainly from Chile and Cal— ifornia; Group B, with partially black pycnidia represented M _ 1The authorities for all taxa follow the abbrevia— tions recommended by Sayre, Bonner and Culberson (1964). dbl- WEE “I pith pale or colorl prised 57 species a were founded partly branches, color real Vainio (189 asabasis for arra stirps Fistularia V branches; stirps lly timuous, arachnoid the stirps Myelopoea rith terete or angt balm, with flatter Reinke (189 habit as a taxonom: pseudocyphellae in be believed Ramal it apothecia structur habit. The sys ten together with Dufc iadiatae of the so would correspond r itmere not for t Llanjouw, 1966) . A study c --=derra and (lanai 95 by R. carpatica Korb. of European distribution; Group C, with pale or colorless pycnidia. The latter group com- prised 57 species arranged into 5 stirps. These stirps were founded partly on thallus morphology, structure of branches, color reaction with KOH, and spore characters. Vainio (1890) utilized the anatomy of branches as a basis for arrangement into stirps. He proposed the stirps Fistularia Vain. for species with inflated, hollow branches; stirps Myelopoea Vain. for species with a con- tinuous, arachnoid or cottony medulla. Vainio subdivided the stirps Myelopoea into the series Teretiusculae Vain., with terete or angular branches, and series Compressiusculae Vain., with flattened branches. Reinke (1895) gave preeminent importance to growth habit as a taxonomic character. He was the first to notice pseudocyphellae in Ramalina while examining R. ecklonii. ow believed Ramalina to be related to Heterodea through apothecia structure, but recognized a difference in growth habit. The system of Hue (1901) placed the genus Ramalina together with Dufourea in the tribe Ramalinee in the family Radiatae of the series Cyclocarpae. The family Radiatae would correspond with the modern concept of the family, if it were not for the fact that it was not named after a genus (Lanjouw, 1966). A study of the cortex of Ramalina species from Madeira and Canary Islands formed the basis for Steiner’s division into secti bad a simple cortex longitudinal course dam orientation to msprotected by a strands, the specie [= sect. Euramalina Stein. was characte the inner layer of Steiner reg to the genus R1393 spore form. This s [1926) in his class Steiner (ll Corticatae Stein. 1 hunt.) Stizenb. r (1852) to designat bosmazieria llont. but was included i Howe (1912 ters which he then tion. Section El ‘o'ith small (9-12 itre included the F Corticatae Ste H- Howe (inch “CK-l and serie: 96 division into sections (1904). Section Corticatae Stein. had a simple cortex formed by hyphae diverging from their longitudinal course in the medulla and taking a perpendic- ular orientation toward the surface. If the algal layer was protected by a cortex as well as by inner longitudinal strands, the species belonged to sect. Bitectae Stein. (= sect. Euramalina Stizenb., 1862). Sect. Ecorticatae Stein. was characterized by having a cortex formed only by the inner layer of longitudinally oriented hyphae. Steiner regarded sect. Ecorticatae Stein. as a link to the genus Alectoria, from which it is separated by its spore form. This section was not incorporated by Zahlbruckner (1926) in his classification system. Steiner (1904) overlooked the fact that sect. Corticatae Stein. had been previously described as Desmazieria (Mont.) Stizenb. This name had first been used by Montagne U852) to designate a genus based on Ramalina homalea Mont. Desmazieria Mont. is not recognized in Zahlbruckner (1926) but was included in his catalogue (Zahlbruckner, 1930b). Howe (1912) provided a system based on spore charac- ters which he thought represented a more natural classifica— tion. Section Ellipsosporae R. H. Howe contained species with small (9—12 u long), ellipsoid spores. In this section were included the series Desmazieria (Mont.) R. H. Howe (= Corticatae Stein.), series Myelopoea Vain., series Cilitae R- H. Howe (including the cilitae species Ramalina crinita Tuck.) and series Fistularia Vain. Section Fus iodide all species lone instituted the basis of sigmoid SP iron California. Du Rietz (1 (Mont.) and sect. E be added under the (l. ll. Howe) Du Rio 1. evernioides and resented the fistul Zahlbruckne [1862) in arrangin: subdivisions . The genus ' in the family Usne {1929, p. 23) on a ima family of its lanaiinaceae allie dwarf fruticose ge While stuc (”agnusson and 2a? danalinopsis base harmelioid thallu SUbstrate . 97 Section Fusisporae R. H. Howe was established to include all species with long (l6—35u) spores. Finally, ibwe instituted the section Bistortae R. H. Howe on the basis of sigmoid spores to include R. bistorta, described from California. Du Rietz (1926) transferred the genus Desmazieria (Mont.) and sect. Euramalina Stizenb. to sub-generic rank. He added under the sub-genus Euramalina sect. Tenuicorticatae (R. H. Howe) Du Rietz. sub-section Solidae Du Rietz included R. evernioides and the sub-section Tubulosae Du Rietz rep— resented the fistulous R. inanis Mont. Zahlbruckner (1926; 1930b) followed Stizenberger 0862) in arranging the sections and Wainio (1890) in their subdivisions. The genus Ramalina, usually placed by taxonomists in the family Usneaceae, was separated from it by watson U929, p. 23) on account of its septate spores, and placed in a family of its own. Watson (1929) regarded the family Ramalinaceae allied to the family Lecaniaceae through the dwarf fruticose genus Thamnolecania. While studying the lichens of Hawaii, Zahlbruckner (Magnusson and Zahlbruckner, 1945) created the section Ramalinopsis based on Ramalina mannii Zahlbr. with a parmelioid thallus and a black lower surface fixed to the substrate. 9,.” too L1? I . le" Ramalina we: by Choisy (1954)' W revealed '9 logy similar to tha an alectorioid morp In consider [1957) concluded ti genus Ramalina and “— he agreed with hat: omtthallus structo he accepted, howeVI sterigmata are com m in which Choisy (l9 Win55. in liaison [1929) who 98 Ramalina was considered a close ally of Alectoria by Choisy (1954). The same author held the opinion that Ramalina revealed through Desmazieria a thalline morpho- logy similar to that of Letharia and through Euramalina an alectorioid morphology. In considering the systematics of Ramalina, Choisy (1957) concluded that the greatest similarity between the genus Ramalina and Usnea was in the pale apothecial disk. He agreed with Watson (1929) that Ramalina, with a differ— ent thallus structure, should constitute a distinct family. He accepted, however, Zahlbruckner's view that exobasidial sterigmata are common to both genera in opposition to Alectoria in which they are endobasidial in origin. Choisy (1957) included both Desmazieria Mont. and Cenozosia Mass. in the family Usneaceae in contrast to Watson (1929) who had characterized the Usneaceae as hav— ing simple spores. The section Protoramalina in the genus Ramalina was established by Choisy (1957) for those species with a cortex composed by hyphae running parallel to the sur- face. He selected R. arabum (Ach.) Meyer and Flot. (= sect. Ecorticatae Stein.) as the type species. In the section Bitectae Stein. he included the subsection Euramalina (Stizenb.) Choisy with R. calicaris (L.) Rohl. as type spe— cies and subsection Fistularia (Vain.) Choisy with R. pusilla Le Prev. as type. His new section Divernia Choisy -- . .. (= sect. Corticatae poeuoc.) Jatta. C however, not validl companied by latin Follmann ar W liont., in the family Rama? h‘ sect. Corticataw 99 (= sect. Corticatae Stein.) was based on R. duriaei (De Not.) Jatta. Choisy's sections and subsections were, however, not validly published because they were not ac— companied by latin diagnoses. Follmann and Huneck (1969) recognized the genera Desmazieria Mont., Ramalina Ach. and Ramalinopsis Zahlb, in the family Ramalinaceae. They found that Desmazieria = sect. Corticatae Stein.) in the sense of Stizenberger (1862) with thin cartilaginous cortex is clearly separated from Ramalina (sens. str.) by the occurrence of ceruchinol and tumidulin. They considered this group to be morpholog— ically, ecologically and chemically homogeneous and agreed with Choisy (1957) in restoring it to a generic rank but within the family Ramalinaceae. The genus Desmazieria Mont. was sub—divided into section Desmazieria with compact medullary hyphae and section Cenozosia with hollow thallus. In the genus Ramalina Ach. (sens. str.) they recognized the section Ramalina and section Ecorticatae Stein. The subsections of section Ramalina followed Vainio (1890) conception of the stirps. Follmann and Huneck (1969) found some evidence of evolutionary parallelism between section Desmazieria and section Ramalina. To show this relationship they estab~ lished the series Cylindricae Follm. and Hun. and the series Complanatae Follm. and Hun. in section Desmazieria to correspond with Zahlbr. and Compres section Myelopoea ( Follmann an lamalinopsis Zahlbr its parmelioid thal iron Ramalina by he ‘_.__ substrate. Recently H: the family name US name Ramalinaceae Usneaceae Esch. (E family name Ramali genus ' ‘ Raw 15 100 to correspond with the series Teretiusculae (Vain.) Zahlbr. and Compressiusculae (Vain.) Zahlbr. in sub— section Myelopoea (Va1n.) Follm. and Hun. Follmann and Huneck (1969) also raised section Ramalinopsis Zahlbr. to a generic level on the basis of its parmelioid thallus. This genus was distinguished from Ramalina by having a lower surface adpressed to the substrate. Recently Hawksworth (1970; 1971) proposed that the family name Usneaceae be retained when the family name Ramalinaceae Ag. (Agardh, 1821) and the family name Usneaceae Esch. (Eschweiler, 1824) are combined. The family name Ramalinaceae should be retained when the genus Ramalina is treated as belonging to a family other than that typified by the genus Usnea. He believed that as the family name Ramalinaceae Ag. was published earlier the name Usneaceae Ag. must be regarded as superfluous if not conserved. Usneaceae Eschw. would therefore be con- sidered illegitimate because it included the type of a previously validly published name in the same rank. W. L. Culberson (1971) contended that regarding these two fami— lies as one has been discredited, and that the use of Usneaceae sensu latissimo has been discarded in the current reevaluation of the families of lichen fungi. C. F. Culberson and W. L. Culberson (1970) noted that chemical evidence strongly supports the segregation : L 35W from th byitself. They co level the fact that commonly present in of Usneaceae. Like series are frequent llorphologi I-lethods After rout all the collection 101 of Ramalina from the family Usneaceae to form a family by itself. They considered of taxonomic value at family level the fact that meta-depsides of orcinol derivatives commonly present in Ramalina are not produced by species of Usneaceae. Likewise, meta-depsides of the B-orcinol series are frequent in Usnea but unknown in Ramalinaceae. Morphological and Anatomical Studies Methods After routine procedures of pressing and drying, all the collections were sorted and annotation cards pre— pared for each specimen. Each packet was examined for mixed collections and separated whenever necessary into different subpackets distinguished by adding letters to the original collection numbers. Chemical variants were included in this segregation. Each specimen was examined under the dissecting microscope for presence of soredia, pseudocyphellae, papillae and other features. A micrometer disk adapted to the dissecting microscope was used to measure apothecia and branches. For the study of apothecia, temporary preparations were made by the use of the squash method. The apothecia were soaked in distilled water or KOH in preparations for further studies. Squash mounts and free hand sections were made to study spores and apothecial tissues. KOH was ezployed as a mount: mess as a clearing : the gelatinous mate reasurements were rm immersion. Routine mic but abandoned as th brittle. A freein 30'30 ll thick. Sec bl Curling of the 1 Morphologir studies, examinati: and literature deSr la ' . w SpeCies. 102 employed as a mounting medium because of its effective- ness as a clearing agent, and its ability to dissolve the gelatinous material binding the paraphyses. All measurements were made in distilled water under oil immersion. Routine microtechnical embedding was attempted but abandoned as the apothecia became too hard and brittle. A freezing microtome was used to cut sections 20-30 u thick. Sections were, however, frequently ruined by curling of the tissue. Morphological data in combination with chemical studies, examination of type specimens, exsiccati material, and literature descriptions were employed to identify the Ramalina species. Thallus The lichen thallus in the genus Ramalina is of the fruticose type reaching 45 cm. in pendant species. The largest species examined were Ramalina anceps and Ramalina usnea while the smallest were 3. subasperata and g. paludosa. The thalli were attached to the sub- strate by basal disks. Zahlbruckner (Magnusson and Zahlbruckner, 1945) described 3. menzii from Hawaii with a parmelioid thallus adPressed by the lower side to the substrate. Most of the and rigid, as recog Surface Features In many spe responsible for lon in species with a r lbw, longi outer layer. In R2 marked by shallow < considered of taxo: moo). it is in t 103 Most of the species in the genus are cartilaginous and rigid, as recognized by Acharius in 1803 and 1810. Surface Features In many species the inner mechanical tissue is responsible for longitudinal striations of the thallus. In species with a nitid surface, such as 3. sorediosa and B. straminea, longitudinal strands are seen beneath the outer layer. In Ramalina gracilis striae are strongly marked by shallow depression on the thallus. These are considered of taxonomic importance, as noted by Vainio (1890). It is in these weak spots that the cortex fre- quently cracks open. The West Indian species are char- acterized by having numerous slits through which the medullary hyphae protrude. These fissures were first reported by Reinke (1895) from the laminae of 3. ecklonii (= g. cumanensis). Darbishire (1901) found such interrup— tions in 3. fraxinea and called them atemporen. Today the atemporen are known as pseudocyphellae. Maas Geesteranus (1947, p. 54) defined them as breaks in the cortex through which the medulla communicates with open air. Du Rietz (1926) classified the pseudocyphellae in the genus Ramalina as follows: papilliform as in E. complanata, maculiform as in g. fraxinea and lineariform as in 3. usnea. Another cha tbepapilla~ This breaks at the 511111111i of mag. These ar hit. complanata th the channels and 211 face of the lobes. iichotomously divir W subaspera‘ uidely scattered o: iiuided. Other Sp 104 Another characteristic feature of the surface is the papilla. This is a low corticated mound that often breaks at the summit to expose the internal hyphae devoid of algae. These are considered of taxonomic significance. In B. complanata they are clustered along the margins of the channels and are also sparsely distributed on the sur- face of the lobes. Frequently, in this species, they are dichotomously divided into two short knob-like branches. Ramalina subasperate is distinctly papillate with papillae widely scattered on the trete branches and not dichotomously divided. Other species such as Ramalina willeyi, 3. montagnei and g. paludosa, which are abundantly collected in Florida, are typically papillate. Tuckerman (1882) described in g. stenospora from southern United States papillose growths which he appro- priately called warts. I have seen the same type of warts in 3. montagnei from Florida and the divaricatic acid var- iant of g. furcellata from Curacao. The wart is a large mound with a flattened summit and a rough surface formed by several small channels. In B. stenospora, 3. montagnei and 3. furcellata these warts are not a constant feature of the species. In the divaricatic acid variant of g. furcellata the warts usually appear on raised areas of the branches. These structures may be regarded as deviations from the typical papillae and are not considered to have taxonomic value. A character W furcellata the raised area bet the surface can be grooved as in Ii. 5 plaiata. The allied W ti'piC: face, that is, witj inflfl the thallu inner wall of the Soralia SOTalla ar 105 A characteristic feature of the surface in Ramalina furcellata is the nodule. This is formed by the raised area between constrictions on terete branches. The surface can be smooth and nitid as R. cochlearis, or grooved as in R. sorediantha, R. cumanensis and R. com— planata. The allied species Ramalina sorediantha and R. scrobiculata typically have a densely scrobiculate sur— face, that is, with many pits or depressions. In Ramalina inflata the thallus is profusely perforated exposing the inner wall of the hollow thallus. Soralia Soralia are frequent in the genus Ramalina. Their presence or absence, position and form provide reliable taxonomic criteria in numerous species. According to Du Rietz (1924, p. 376), soralia are those exposed parts of the lichen thallus which are profusely invaded by algae cells. In this process the algae are surrounded by short hyphae resulting in granular or farinose bodies called soredia. Du Rietz suggested that these soredia were the reproductive structures of many lichens and constituted a dispersal mechanism. He also called attention to the difficulties encountered sometimes in distinguishing soralia from pseudocyphellae. Out of 23 s mine have soralia. soralia by Du Riet: llaas Geesteranus ( under the general well defined spots i.Soralia superf on branches of l. llaculiforn \ round or c 106 Out of 23 species examined from the west Indies nine have soralia. Following the classification of soralia by Du Rietz (1924) and the interpretation by Maas Geesteranus (1947, pp. 54-55) these species fall under the general category of soralia confined to small well defined spots. A.Sora1ia superficial, marginal and lateral, occurring on branches of fruticose lichens. 1. Maculiform soralia. This type originates as a round or oblong spot in Ramalina farinacea, R. peruviana, R. camptospora and R. dendroides. 2, Fissure shaped soralia. The soralia originate as long narrow cracks in the cortex. This type is frequent in R, dendroides, especially in the divaricatic acid containing variant. 3. Marginal soralia. Du Rietz (1926) cited R. peranceps as an example of a species with this type of soralia. After examining the type spec- imen it was found that R. peranceps is not a sorediose species. However, marginal soredia are found in R. sorediantha, R. farinacea and R. dendroides. B.Terminal or subterminal soralia occurring at the tips of branches. 4. Capitiform soralia. These are globose soralia at the tips of main and lateral branches. This type is found in R. dendriscoides, and R. sorediantha —— —-————-———————— — W. 5, Helmet shat __——_———- as globose which spli‘ forming a I are produc cochlearis Rhizines These stru sleties studied. W b)’ Bra Cilia 107 5. Helmet shaped soralia. These soralia originate as globose structures at the tips of branches, which split with vaulting of the upper lip, forming a helmet at the inside of which soredia are produced. In the West Indies only Ramalina cochlearis has this type of soralia. Rhizines These structures are not present in any of the species studied. The rhizines were described for R. landroensis by Brandt (1906). Cilia Cilia are not present in the West Indies species. They were described by Tuckerman (1883) from R. crinita and Howe (1914) established the series Ciliatae R. H. Howe to include this species. Tissues The thallus is heteromerous and consists of layers or rings of tissues. These comprise cortex, algal layer, and medulla. The reproductive structure is the apothecium, in which several specialized tissues can be recognized. A vertical section through the apothecium shows, starting from the top, the epithecium, hymenium with paraphysis and asci, hypothecium and exciple. cortex. - 'T} of Ramalina is f0” hyphae so closely 1 form a continuous 3 second layer 0f sur port to the thallu alongitudinal, C0 W and ii. trete bundles of l twins» 3- a % among ot appears as a conti uatous tissue form Difference lontagne (1852), i did Steiner (1904' in the genus. I The first “311115 were made %Fr. Hi 108 Cortex.—-The cortex of the West Indian species of Ramalina is formed by an outer layer of transverse hyphae so closely packed on the outer surface that they form a continuous layer. At the inside of this layer a second layer of supporting tissue gives mechanical sup- port to the thallus. This inner tissue is organized in a longitudinal, continuous ring as in R. complanata, R. dendroides and R. dendriscoides or separated into dis- crete bundles of longitudinally oriented fibers as in R. sorediosa, R. gracilis, R. usnea, R. anceps and R. straminea among others. In cross section this inner wall appears as a continuous or broken ring of pseudoparenchy— matous tissue formed by thick walled cells. Differences in cortex morphology were used by Montagne (1852), Massalongo (1854), Stizenberger (1862) and Steiner (1904) to distinguish groups of species with~ in the genus. The first studies on the fine structure of the thallus were made by Speerschneider (1855) on Ramalina calicaris Fr. His observations were considered primitive by Brandt (1906) as he did not distinguish between me— chanical tissue and outer cortex tissue. Speerchneider mistook the lumen of the cells for the whole cell and the gelatinous cell membranes for a cementing substance between them. Schwendener (1860) in studying species of Egggliga noted that the inner cortex is intimately attached to the outer corte: ture was not reco gr {1894). Both authr lemon and Steiner (l the algal layer a of this tegument h the 0f tegument 1 ward the Periphery he found that in ( 109 to the outer cortex. The complexity of the cortical struc— ture was not recognized by Nylander (1870) and Crombie (1894). Both authors regarded the cortex in R. evernioides, R. scopulorum and R. pusilla as amorphous, i.e., not cellular. Steiner (1904) called the hyphal complex that covers the algal layer a tegument. On the basis of the arrangements of this tegument he erected three sections. He observed a type of tegument in which the medullary hyphae diverged to- ward the periphery in a transversal orientation. Secondly, he found that in other species an additional inner layer of longitudinal hyphae was present. Thirdly, Steiner noted that in a few species only the inner longitudinally oriented layer was present. The pseudoparenchymatous nature of the cortex was reported by Brandt (1906) after studying the anatomy of European Ramalinas. He observed that with the exception of R. evernioides all species had a supporting tissue. This tissue appeared in cross section as a continuous cartilag- inous ring. Medulla.——The medulla is a white mass of hyphae protected by the cortex. It is a porous tissue of a cottony consistency with numerous lacunae between the hyphae. Spe- cies belonging to the stirps Fistularia Vaini, such as R. inflata, constitute the extreme case in which the medulla seems to be absent or present only as a scant layer around rcentral cavitY- Stizenb. as the ot sue is lacking b“ to provide support Algal Laye iuthe outer zone ingamore or less cells. Nylander ( i-ll p in diameter cells were envelop to be of the Protr reporting the alg; lrebo ' &Puym. a: lpothecia 110 a central cavity. Keissler (1959) pointed to Desmazieria Stizenb. as the other extreme in which the supporting tis- sue is lacking but the medulla is very thick, presumably to provide support. Algal Layer.—-The algal layer in all species lies in the outer zone of the medulla, against the cortex form— ing a more or less continuous ring of clusters of green cells. Nylander (1870) described the algae (gonidia) as 7-18 n in diameter. Wainio (1890) observed that the algal cells were enveloped by a thin membrane and believed them to be of the Protoccoid type. Ahmadjian (1958; 1967) in reporting the algae occurring as lichen symbionts cited 'Trebouxia Puym. as the phycobiont in Ramalina. Apothecia Lecanorine apothecia are commonly found in the non-sorediate species. They are usually terminal in R. inflata, lateral as in R, dendroides or laminal as in R. cumanensis. Species such as R. subasperata, R. sub- pellucida and R. bistorta bear apothecia inserted at the point where the branches_bend in an acute angle taking a geniculate position. In such cases the tips of the branches seem to be growing out from the lower side of the apothecia, The apothecia vary in shape from species to species, being cup-shaped, flattened or convex. They are usually subpedicellate, but in R. usnea and R. anceps may be better described as sessi largest (6-8 mm. b smallest (less that The form a vith respect to th to distinguish RE The thalli ous with the thall tan to brown. C011 less often R. m lettions originati W; walla develop apO' that no fertile t Smith was obs erv ibis 586mg to be excretmns. The 111 described as sessile. Their size is variable, with the largest (6—8 mm. broad) found on R. complanata, and the smallest (less than 1.5 mm.) in R. anceps. The form and relative position of the apothecia with respect to the thallus were used by Acharius (1810) to distinguish Ramalina from Parmelia. The thalline margin of the apothecia is concolor- ous with the thallus, but the disk varies in color from tan to brown. Commonly R. subasterata and R. willeyi and less often R. complanata, exhibit short finger—like pro- jections originating on the thalline margin of the apothecia. Ramalina sorediosa and its allied species R. peru- viana develop apothecia which abort before maturation so that no fertile tissue is differentiated. A type of warty growth was observed growing on the disk of many species. This seems to be a response to insect bites and poisonous excretions. The possibility remains that they are some sort of parasitic growth since no further investigation was done on this structure. Hymenium.—-The hymenium is thickly gelatinous (1 + blue) extending to a height of 30—80 mm. Paraphyses are slender, branched or simple, with capitate apices. The ePithecium is formed by the swollen tips of the parPhYses among which a granulose material (pruina) is deposited. This layer is slightly tan colored, except in A. sorediantha formed by a double spores variously a spores, these are series or in two i organized in two 5 0vmapping those and a straight se 112 in R. sorediantha in which it is brown. Asci are clavate, formed by a double membrane sac containing eight colorless spores variously arranged. In species with ellipsoid spores, these are arranged either in a single oblique series or in two irregular series. Fusiform spores are organized in two sets of four, the upper ones partially overlapping those below. Spores.--Spores are two celled, with thick walls and a straight septum. Ramalina bistorta and often R. camptospora depart from the pattern by producing sigmoid spores with an oblique septum. Large spores such as those produced by the R. usnea complex occasionally are tri-septate. Within the spore cells a large nucleus occupies the center of the lumen leaving a narrow ring of vacuolated cytoplasm. In rare cases I have observed germinating spores with hyphal tubes at the tips. The size of the spores is more or less constant between groups of species. Howe (1914) employed spore form and size to establish subdivisions of genus at the section level. He proposed sections Fusisporae R. H. Howe for species containing fusiform spores, Ellipsosporae R. H. Howe for species producing ellipsoid or oblong Spores and Bistortae R. H. Howe for sigmoid spores. I do not be- lieve that the species fall so neatly into these three categories. Ramalina p have the largest a tbellest Indies S; W (8-10 (8-12 x 3-6 H) in bad the largest 5] A number . fusiform, straigh b M and 113 Ramalina usnea, R. montagnei and R. subpellucida have the largest and narrowest spores (16-30 x 3-5 p) of the West Indies species. The smallest are those of R. leptosperma (8-10 x 2-3 u) in the Antilles and R. paludosa (8-12 x 3—6 p) in Florida. A Mexican species, R. alludens, had the largest spores in tropical America (30-40 x 3—6 p). A number of species produce intermediate size, sub- fusiform, straight spores. This group includes R. peruviana, R. dendroides and R. sorediosa. Although, the size and form of the spores are useful taxonomic characters at the species level every species I have examined contained both straight and bent or curved spores in different proportions. Subhymenial Tissues.——Beneath the hymenium is a layer of colorless, compact hyphae, among which the asci and paraphyses originate. This tissue is called the hy— pothecium. The outer rim of the hypothecium forms the proper margin or exciple. This is the only rim present in lichens with a lecideine or biatorine apothecium. In lichens with a lecanorine apothecium the proper margin is enclosed by a thalloid margin containing an algal layer. In Ramalina the hypothecium is 10—15 u thick in most spe- cies. Pycnidia Pycnidia were found to be abundant in R. sub— asperata, R. peruviana and R. subpellucida. They consist ; :' ms black bottle-ne H the cortex. The c side the conceptac lbese conidia are and exobasidial ir conidia is not kni reproductive funC‘ it Nylander (18 Pycnidia to elabo l 114 of black bottle-necked conceptacles slightly embedded in the cortex. The conidiophores are branched hyphae in— side the conceptacle that cut off conidia from the tips. These conidia are minute (2-6 u), one-celled, cylindrical, and exobasidial in origin. The true function of the conidia is not known. One school of thought advocates the reproductive function of the bodies while another denies it. Nylander (1858—60 and 1870) used the color of the pycnidia to elaborate the classification of the genus. Chemical Studies Introduction The utilization of chemical tests in plant tax— onomy was first made in the lichens by Nylander (1866a, 1866b). He claimed that different species responded dif— ferently by producing characteristic colors when potassium hydroxide or calcium hypochlorite were applies to the me— dulla or cortex. His methods were strongly opposed by Tuckerman (1868, 1872) and Willey (1896) on the basis that well known species were chemically inconsistent. The range of lichenological reagents increased when Asahina (1934) introduced the spot test using p— phenylenediamine (PD). Application of this substance to the lichen thallus produced a yellow or red color in cer- tain lichens but no reaction in others. Asahina (1 crystallization te stances. The use acids has been de: lachtmeister (1951 Sthorn (1961) were gel plates for th type of plate, Ra arated aldehydic substances were 5 [1956]. A quicker used by Santesson lithen substances Fluoresce drenda (1951) to rented both micrr mph). by Studyin: sidones under 111' investigated the 115 Asahina (1936) developed simple microchemical crystallization tests for the recognition of lichen sub- stances. The use of paper chromatography for lichen acids has been described by Asahina and Shibata (1954), Wachtmeister (1956, 1959) and Hesse (1958). Stahl and Schorn (1961) were the first to use hand coated silica gel plates for thin—layer chromatography. Using this type of plate, Ramaut (1963) and Santesson (1965) sep- arated aldehydic aromatic compounds. Aliphatic lichen substances were studied by Bendz, Santesson and Tibell (1966). A quicker method employing precoated plates was used by Santesson (1967) to distinguish more than eighty lichen substances. Fluorescence of lichens acids was employed by Ozenda (1951) to delimit species. Hale (1956a) supple— mented both microchemical tests and partition chromatog— raphy by studying the fluorescence of despides and dep— sidones under ultraviolet light. The same author (1956b) investigated the absorption spectra for such substances. Other methods employed in elucidating the struc- ture of lichen substances have been summarized by Huneck (1968). Some of these are gas chromatography, infra-red, nuclear magnetic, mass spectroscopy and fragmentation techniques. Brandt (1906) was one of the first to report on the chemistry of Ramalina species. He noted that the thallus and rhizin products which inc ated those species lonot. All the s rained usnic acid survey of the gem be compared the S] and usnic, ramalir Later wor C0uponents in the complex of relate also to the quest h oendr, Bantesson 116 thallus and rhizines of certain species produced metabolic products which included calcium oxalate. Thus, he separ- ated those species producing the oxalate from those that do not. All the species of Ramalina that he examined con- tained usnic acid in the cortex. The first chemotaxonomic survey of the genus was done by Zopf (1907) in 14 species. He compared the species as to their production of atranorin and usnic, ramalinolic, cuspidatic, evernic, salazinic and obtusatic acids. Later works comprised investigation of chemical components in the lichen thalli of single species or in a complex of related species. Some attention has been given also to the question of distribution of chemical species. Bendz, Santesson and Wachtmeister (1965) studied the Ramalina ceruchis group, W. L. Culberson (1966) the Ramalina farinacea complex, and later (1967) the Ramalina siliquosa group in Europe. Asahina (1934, 1936, 1937), C. Culberson (1965a), and Huneck and Follmann (1965) have studied the chemistry of single species. Follmann and Huneck (1969) reported on the chemistry of 32 species and compiled the chemistry of more than 40 other species. C. Culberson (1969; 1970) listed the chemical constituents 0f 96 taxa reported prior to 1969 (see Table 16). l l Methods and Technf Color Reactions As a prel tests were carrie cagents used wer noon. The pota any Color was obs the calcium hypoc reaction gave any allllication of K( recorded as KC+ j aFiltered, The Cc ing an alCOholic 117 Methods and Techniques Color Reactions As a preliminary approach to identification, color tests were carried out on all lichen thalli. The color reagents used were those introduced by Nylander (1866a and 1866b). The potassium hydroxide (K) is recorded as K+ if any color was observed and K- is no color was produced; the calcium hypoclorite (C) test is recorded as C+ if the reaction gave any color, C— if the reaction was negative; application of KOH followed by calcium hypochlorite is recorded as KC+ if any color was seen and KC- if no color appeared. The color test developed by Asahina (1934) us- ing an alcoholic solution of p—phenylenediamine is denoted as PD+ if a color was developed and PD— if no color at all was observed when applied to the lichen thallus. The so— lutions of K and C were kept in reagent bottles since both are highly corrosive. In applying the color tests the PD test was the first one to be used since its results deter- mined the reagent to be used in the microchemical test and the solution used to spray the chromatograms. The K, C and KC tests followed the PD test. A fresh PD solution was prepared every time it was used by adding a few drops of alcohol to a pinch of p- Phenylenediamine. It was found that the amount of solute and solvent are not critical in this test. The K solution i used was a 15 pert and commercial ble Imshaug to replace Early in ‘ the color reactior or the lichen tha razor blade under of the color reag micropipette and made under the di 118 used was a 15 percent solution of potassium hydroxide and commercial bleaching solution was suggested by Imshaug to replace calcium hypochlorite. Early in the study the method used to test for the color reaction was to apply the chemicals directly on the lichen thallus after exposing the medulla with a razor blade under the dissecting microscope. Application of the color reagent was done by means of a disposable micropipette and determination of any color change was made under the dissecting microscope. The treated por- tion of the thallus was removed so that the rest of the lichen and/or the study packets were not ruined by the lasting corrosive action of the chemicals employed. How— ever, it was noticed that even with the greatest care ac- cidental impregnation of other parts of the thallus often occurred, that not always the color reaction was fully appreciated especially in old herbarium specimens and finally the well known fact that some species either do not have a well defined medulla, in which case the results were indefinite, or the branches were so thin and brittle that involuntary damage was done to the specimen. For these reasons the filter paper method used by Santesson (1967) was adopted. In implementing this method a regular size filter Paper (Whatman No. l) was cut into four pieces and a frag- ment of a thallus was placed on the center of a filter gaper piece and th plate. Drops of a after the previous substances in a ri ments were discarc the appropriate re the extract ring. tected far more a causing damage to llicrochemical Tes A small h SMUtl Cm-Z was set on a warming me dmu at a tin evaporated bEforc flaw instead Of end time was savr T0 insur. Lt seemed better mulls than to la“ cent Shere diSCa Sta -7. .hs \ Slide w 119 paper piece and then extracted with acetone on the warming plate. Drops of acetone were added (5—10) one at a time, after the previous one had evaporated leaving the extracted substances in a ring around the lichen fragments. The frag- ments were discarded and under the dissecting microscope the appropriate reagent was applied with a micropipette on the extract ring. In this way the color reactions were de— tected far more accurately than on the thallus and without causing damage to the specimens. Microchemical Tests A small heap of thallus fragments in an area of about 1 cm.2 was placed on a chemically clean glass slide set on a warming plate at about 60° C. Acetone was added one drop at a time, waiting until the previous drop had evaporated before adding the next. The use of a warming plate instead of an open flame was found to be convenient and time was saved by doing several samples together. To insure sufficient extract from PD- specimens it seemed better to add more acetone or more pieces of thallus than to try to scrape the scant residue with a razor blade since these are usually gummy substances. After extraction was completed the thallus frag- ments were discarded and a drop of one of several cry- stallizing agents was applied followed by a cover slip. The slide was left on the warming plate until the residue dissolved and ther tion occurred in 2 cases. If no cry: was left overnigh‘ crystallization 01 At this 5 litre magnificatio The crYstals were photographs and t the suspected sup; Care was tions by examinix scope and using ( of test. If Chm \‘isable to use ti chemical test 120 dissolved and then was set aside to cool. Recrystalliza— tion occurred in a few minutes after cooling in most cases. If no crystals were immediately formed the slide was left overnight and usually after a 24 hour period crystallization occurred. At this stage the preparations were examined under 100x magnification for form and color of the crystals. The crystals were identified by comparing them to published photographs and to crystals of material known to contain the suspected substance. Care was taken to avoid extraction of mixed collec- tions by examining each thallus under the dissecting micro- scope and using only one labelled thallus for each series of test. If chromatograms are to be developed it is ad— visable to use the same residue employed for the micro— chemical test. One way of doing this is to extract the thallus in a vial and use part of the extraction for micro- chemical test and part for chromatography. The microchemical test posed some problems when more than one substance was present in the lichen thallus. Sometimes the presence of one substance inhibited the for- mation of crystals by another. Some acids assumed atypical forms when other acids were present. In those plants con— taining PD+ and PD— substances in the medulla, the PD— sub- stance may go unnoticed if chromatograms are not prepared. When chromatographic analysis showed this type of chemistry the sample has to G.A.o-T. for PD+ 5 Wu The abbrer liring solutions 1 by Hale (1961). T iuG.A.o-T., excej crystals in G.E. hood in G.E., Q, 1135 were used: G.A.O-T. (gly G'E- (glyceri G'A'W- (glyce Ba[0gg)2 (a 52 These sol 35 they are fair- alresh SUppr er ChI‘OmatO 121 the sample has to be tested twice for crystals, with G.A.o-T. for PD+ substances and with G.E. or G.A.W. for PD-. The abbreviation and composition of the crystal- lizing solutions used in this study follows those given by Hale (1961). Extracts of PD+ specimens were treated in G.A.o-T., except psoromic acid which forms typical crystals in G.E. Specimens reacting PD- were recrystal- lized in G.E., G.A.W. or Ba (OH)2. The following form- ulas were used: G.A.O-T. (glycerin - alcohol—o-toluidine, 212:1 v/v/v). G.E. (glycerin—acetic acid, 1:1 v/v/v). G.A.W. (glycerin-alcohol—water, 1:121 v/v/V). Ba(OH)Z (a saturated solution of barium hydroxide). These solvents were kept in small reagent bottles as they are fairly stable and there is no need to prepare a fresh supply every day. Thin Layer Chromatography Chromatograms of all collections were done as a routine procedure in an attempt to verify the results of the color and microchemical tests. A second objective was to get information on possible substances not detected by other methods. Fragments of lichen thallus were selected under the dissecting microscope to avoid mixed extraction. The fragments were In.) and extractei duct was concentr. on the warming pl ried out on preco 6060) containing beafast and rel stances. Non-flu as the fluorescer possible to cut t spots 1 gm. aparv with a micropiper the lower edge 03 never spreaded b. cedure was Carri 1. step the eXtTaCt the sheets WEre their Use . This IOTlll TESultS 0V8 122 The fragments were placed in.a flat bottomed vial (6 x 50 mm.) and extracted in warm acetone. The extraction pro- duct was concentrated by evaporating part of the acetone on the warming plate. Thin layer chromatography was car- ried out on precoated sheets (Eastman chromatogram sheets 6060) containing a fluorescent indicator. This proved to be a fast and reliable way for identifying lichen sub— stances. Non-florescent compounds were spotted as easily as the fluorescent ones. After some experience it was possible to cut the sheets in halves and accommodate 19 spots 1 cm. apart in each half. The samples were spotted with a micropipette (drawn out capillary tube) 2 cm. from the lower edge of the sheet, taking care that the spots never spreaded beyond 2 mm. in diameter. The entire pro- cedure was carried out on the warming plate in order to keep the extract confined to as small a spot as possible. The sheets were kept in a dessicating chamber prior to their use. This proved to be of help in providing uni- form results over a period of time. The sheets were developed in two different solvent systems: (A)benzene—dioxane-acetic acid, 90:25:4 v/v/v (Santesson, 1965), and (B)toluene-acetic acid, 9:1 v/v (Santesson, 1967). In the developing chamber the solvent was allowed to ascend to 1 cm. below the upper edge of the plate. After the chromatograms were completely dried, in spots were rer right. Tracing 0i pencil and the C0 The spots tograms with a 5“ iromthalli reaCt tion of p-phenl’le Steiner (I955) is sulfite and 1 gm- rater. a; r For PD- 5 oenzidine as des 11 Identif 3\' ~ ‘ ‘O‘Chl‘omatcig idllllv' 123 the spots were revealed by observation under ultraviolet light. Tracing of the spot was done with a soft lead pencil and the color of fluorescence was noted. The spots were visualized by spraying the chroma- tograms with a suitable reagent. Those lichens substances from thalli reacting PD+ were sprayed with a stable solu- tion of p—phenylenediamine. The formula introduced by Steiner (1955) is prepared by dissolving 10 gm. of sodium sulfite and 1 gm. of p-phenylenediamine in 100 ml. of tap water. For PD- substances a solution of bis—diazotized benzidine as described by Lindstedt (1950) was used. Solu- tion I was prepared by dissolving 5 gm. of benzidine de- hydrochloride in 14 m1. of concentrated HCl, to which water is added to make 1 liter of solution. Solution II was a 10 percent solution of sodium nitrite. The spraying was formed by slowly adding an equal amount of solution 11 to solution I. The wet sheets were dried on the warming plate in order to evaporate the excess spray and accelerate the reactions. Chromatograms were numbered and filed for fu— ture reference. Care was taken to keep apart those sprayed with PD from those with benzidine. Identification of lichen substances were checked by co-chromatogramming with a sample from a thallus with known substances. The method used was a modification of :be simple method These were run on diameter and 12 cr The unknown and t left of a mixture is that Rf values ing much material lichen Substances \_.._ In the ir substances found structure or to . liih each of ser The lich. 1) Arom reaction “it 7 ~i Deps b“ giving a hydroxide ar red, KC+ rec 3) pepE but produci] (Table 7). 124 the simple method employed by Rockland and Dunn (1949). These were run on a strip 2 cm. wide, using vials (2.5 cm. diameter and 12 cm. high) as chromatographic chambers. The unknown and the standard were spotted at the right and left of a mixture of both. The advantage of this technique is that Rf values can be directly compared without expand- ing much material. Lichen Substances in Ramalina _______________________________ In the investigated species of Ramalina the lichen substances found can be grouped according to their chemical structure or to the color reactions produced when treated with each of several widely used reagents. The lichen substances discussed here are grouped as follows: 1) Aromatic aldehydes giving a positive color reaction with p—phenylenediamine (Table 5). 2) Depsides not reacting with p-phenylenediamine but giving a positive color reaction with potassium hYdroxide and calcium hypochlorite, PD-, K+ red, C+ red, KC+ red (Table 6). 3) Depsides not reacting with PD, K, C and KC, but producing coloration with bis—diazotized ben21d1ne (Table 7). fi—hh “z‘amfilvfi 4) Vario vious groups, geneous subst and an unknow 5) Unide depside natur red (Table 9) Atranorin (Plate Atranorii We elaborated : 125 4) Various substances not belonging to the pre— vious groups, but reacting PD—. In this group hetero- geneous substances such as a fatty acid, a dibenzofuran, and an unknown were put together (Table 8). 5) Unidentified lichen substances suggesting a depside nature which react PD—, K—, C-, KC— or KC+ red (Table 9). Atranorin (Plate I, Figure l) Atranorin is a para-depside of the B.-orcinol type elaborated in the cortex of many lichens. In Ramalina it is always associated with PD— substances in species such as R. complanata, 3. dendriscoides, R. anceps and E. QEEQEQLQEE. Its color reactions are K+ pale yellow, PD+ yellow (not seen on the thallus) and C—. The best micro- chemical test is to treat the dry acetone extract with G.A.o-T. The treated residue precipitates as very long slender crystals typically curved, which immediately form feathery masses, usually in radiating clusters. If the residue is treated with G.E. solution the resulting cry— stals are short, simple, colorless prisms with pointed ends. This substance exhibits a dark color under UV and a pale yellow color after reaction with PD. It reacts With benzidine to give a red color. Atranorin has the highest Rf value of all the substances except its derivative Table 5. Aroma? reactio‘ Substance \ Atranorin Chloroatranorin lorstictic acid Protocetraric aci Psoronic acid Salazinic acid \ lSolvent l/Y/YI 2 Solvent 126 Table 5. Aromatic aldehydes giving a positive color reaction with p—phenylenediamine. W Rf x 100 Color Substance Solvent Solvent in UV Color with PD A1 132 Atranorin 70-72 70—72 dark yellow Chloroatranorin 76—78 76-78 dark yellow Norstictic acid 62-64 22-24 dark dull yellow Protocetraric acid 10—12 04-06 dark yellow—orange Psoromic acid 60-62 38-40 bright yellow—green yellow Salazinic acid 15—17 04-06 dark dull yellow W lSolvent A: Benzene-dioxane—acetic acid, 90:25:4 V/V/v, 2Solvent B: Toluene-acetic acid, 9:1 v/v. inn 6. Depsidc p-phen) reactic hypochl Substance kWtOChlorOphae acid \\ lSolvent c/ / 2 SOlVent 127 Table 6. Depsides not giving a color reaction with p-phenylenediamine but giving positive color reaction with potassium hydroxide and calcium hypochlorite. Rf x 100 Substance --—-—-—-—————- Color Color with Solvent Solvent in UV bis—diazotized A1 B2 benzidine Cryptochlorophaeic acid 53-55 30—32 blue yellow-orange Ramalinolic acid 30-36 06—08 blue pink—red W lSolvent A: Benzene—dioxane—acetic acid, 90:25:4 V/v/v. 2Solvent B: Toluene-acetic acid, 9:1 v/v. Table 7. Depside K, C or Substance M Divaricatic acid Sekikaic acid 128 Table 7. Depsides not giving a color reaction with PD, K, C or KC. Rf x 100 Color Color with bis-diazotized Substance Solvent Solvgnt in UV benzidine A B Divaricatic acid 64—66 50~52 blue red-brown Sekikaic acid 63-65 49-51 blue red-brown 1Solvent A: Benzene-dioxane-acetic acid, 90:25:4 v/v/v. 2Solvent B: Toluene-acetic acid, 9:1 v/v. Table 8. Various substances reacting PD— and belonging to different chemical groups. W M Rf x 100 Color Color with Substance Solvent Solvent . bis-diazotized 1n UV b . . 1 2 enZidine A B ________________________________________________________________ Caperatic acid 30-32 0—0 blue faint yellow (fatty acid) Usnic acid 68—70 68-70 dark no color (dibenzofuran) Substance H 0—0 0-0 0-0 no color (unknown) W lSolvent A: Benzene-dioxane-acetic acid, 90:25:4 V/v/v. 2Solvent B: Toluene acetic acid, 9:1 v/v. ,— Table 9. Unident KC- or ‘— ‘— Substance2 __ Soln \— Substance A 6( ( Substance B 61 Substance c 5: Substance D 1‘ 129 Table 9. Unidentifie substances reacting PD—, K—, C-, KC- or KC+. Substancez Rf X 100 Color 5010: with d 3 4 . is- iazotize Solvent A Solvent B in UV benzidine Substance A 60—62 45—47 blue brown 0-0 3Z~34 blue brown Substance B 63—65 51—53 blue brown-yellow 23-25 21-25 blue brown Substance C 58-60 50-52 blue orange Substance D 18-20 08-10 blue orange lSubstance A and B reacted KC+. 2Substance A produced 1 spot in solvent A and 2 spots in solvent B. Substance B produced 2 spots in both solvents A and B. 3Solvent A: Benzene-dioxane-acetic acid, 90: 25:4 v/v/v. 4Solvent B: Toluene—acetic acid, 9:1 v/v. "n :hloroatranorin. Spot just above u both developing s varies with conce more or less tria Caperatic Acid (I Caperatic been shown to oc< 130 Chloroatranorin. Its relative position is marked by a spot just above usnic acid, exhibiting Rf .70— .72 in both developing systems used. The shape of the spot varies with concentration from a narrow streak to a more or less triangular form. Caperatic Acid (Plate I, Figure 2) Caperatic acid is the only fatty acid that has been shown to occur in the species studied. It is the only medullary substance in Ramalina bistorta and 3. camptospora. This fatty acid shows a negative reaction with spot tests on the lichen medulla. Extraction of caperatic acid with acetone is slow but enough material is produced to perform the microchemical test. The residue obtained is a shiny, clear, gummy substance. When G.E. is added to a sample containing Caperatic acid circular cloud—shaped clumps resembling drops of oil are precipitated. These crystals are certainly the most distinctive in the genus being easily recognized when typically formed. However, the crystals do not always adopt the cloud-shaped form and occasionally the pattern is altered by the introduction of lateral branches with a festooned edge. Chromatography of caperatic acid in benzene-dioxane- acetic acid gives a round spot fluorescing blue in UV at UIDJJ if .30- .32. Th: in toluene-acetic When chrc acid are sprayed yellow color is I Chloroatranorin This sub: Atranorin (Hunecl new tha‘ This cortical sul Its Chromatogram with an Rf value 131 Rf .30- .32. This fatty acid produces no chromatogram in toluene—acetic acid. When chromatograms containing spots of caperatic acid are sprayed with benzidine a transient, very pale yellow color is barely distinguished. Chloroatranorin This substance is a dichloride derivative of atranorin (Huneck, 1968). It is an infrequent substance in Ramalina, that whenever is present accompanies atranorin. This cortical substance is not detected by crystal tests. Its chromatogram produces a yellow streak above atranorin with an Rf value of .78 in both running solvents. Cryptochlorophaeic Acid (Plate 1, Figure 3) This depside was reported (Moore, 1968) from the minute species 3. paludosa from Florida. I found this acid in a collection of 3. farinacea from Chile and in a variant of 3. complanata from the West Indies. The color reactions of this acid are K+ turning wine red, KC+ bril— liant red, C+ red and PD—. These reactions are better observed on filter paper extractions of the thallus. The crystals of cryptochlorophaeic acid precipi— tate from G.E. as fine penicillate needles and from G.A.W. as fine, slightly curved needles radiating from a central point. Single needles unite and then branch and rebranch forming feathery aggregates. 132 PLATE I Figure l. Crystals formed by atranorin in G.A.o-T solu- tion ( x 65). Extracted from Ramalina complanata. Figure 3. Crystals formed by cryptochlorophaeic acid in . G.A.W. solution ( x 80). Extracted from BEEEJEE complanata. Chromatoé ultraviolet lighl fluorescence. 1] acid (9:1 v/V) C7 .32 while in ben are.53- .55- T abright orange Cryp to ch W Divaricatic Acid This mec iron orsellic ac , a. u ‘ snea and 3. it, and c-. n 134 Chromatograms of this acid when exposed under ultraviolet light exhibit ellipsoid spots with a blue fluorescence. In a developing system of toluene-acetic acid (9:1 v/v) cryptochlorophaeic acid has an Rf .30— .32 while in benzene—dioxane—acetic acid the Rf values are .53- .55. The acid reacts with benzidine producing a bright orange color. Cryptochlorophaeic acid also occurs in Cladonia cryptochlorophaea and Parmelia cryptochlorophaea. Divaricatic Acid (Plate II, Figure 4) This medullary substance is a depside derived from orsellic acid. It occurs in Ramalina subpellucida, E. usnea and R. complanata. The acid reacts PD-, K—, KC-, and C~. The acetone extract yields abundant residue WhiCh readily recrystallize upon addition of one of sev— eral solvents. When G.E. or G.A.W. is added to the dry eXtract long colorless plates arrange in a criss-cross Pattern, with the plates intersecting each other at right angles, In Ba(OH)2 solution radiating plates in discrete Clusters are formed. Isolated plates somewhat resemble Usnic acid. Divaricatic acid produces a blue fluorescence under ultraviolet light. The acid exhibits a round spot at Rf .64— .66 in benzene—dioxane acetic acid and at Rf -50' .52 in toluene—acetic acid. Chromatograms of all specimens contai! low divaricatic i drolysis. Whenl ated-brown colo Norstictic Acid Norstict of the depsidone is the pr inc ipa] W and R. 16 acid it often cc 011 chromatogramj 135 specimens containing this acid show a blue spot just be- low divaricatic acid which is probably a result of hy- drolysis. When benzidine is applied to the chromatogram a red-brown color is produced. Norstictic Acid (Plate II, Figure 5) Norstictic acid is an aromatic aldehydic acid of the depsidone type derived from B— orsellic acid. It is the principal substance in the medulla of Ramalina anceps and 3. leptosperma. In species containing salazinic acid it often occurs as a secondary product recognized only on chromatograms. A small piece of thallus gives a large amount of a white powder when extracted with acetone. The acid gives a strong red color when treated with KOH and a yellow to orange color with PD. The dried acetone extract containing this acid will yield typical norstictic acid crystals very readily when G.A.o-T. is added. The crystals are flat, rectangular plates with a yellow color which preCiPitfite Singly or arranged in clusters. Under UV the substance has a dark color. Norstictic acid Produces a yellow color in daylight after reacting with P‘Phenylenediamine and fluoresces dull yellow under UV. The chromatogram exhibits a round spot at Rf .62- .64 when de— veloped in benzene—dioxane—acetic acid and at Rf .22' -24 when in toluene-acetic acid. Protocetraric Aci This deps present in Ramal: It reacts giving does not react w upon addition of minute yellow ne Power in the mic 136 Protocetraric Acid (Plate II, Figure 6) This depsidone of the B—orsellic acid group is present in Ramalina complanata and in g. subasperata. It reacts giving a brick-red color with PD and KC, but does not react with K or C. The dry acetone extract upon addition of G.A.o—T. precipitates as bundles of minute yellow needles so closely packed that under low power in the microscope they resemble yellow granules. This reaction needs a fairly large amount of residue for the crystals to precipitate. Excessive heat causes dis— solution of all the extracted material decreasing the chances of recrystallizing the acid. Protocetraric acid is easily chromatographed in both solvent systems used. In benzene-dioxane—acetic acid it develops as an ellipsoid spot with Rf .10- .12 bUt at Rf .40- .60 when toluene—acetic acid is used. The acid reacts yellow-orange with PD in daylight. Under UV the spot has a dark color that fluoresce red- brown after being sprayed with PD. Psoromic Acid (Plate III, Figure 7) This is an aromatic depsidone of the B—orcinol type, It is the major component in Ramalina gracilis. Its color reactions are PD+ deep yellow, K—, KC- and C-. When G.E. is added to the white acetone extract, color— . - re less crystals are formed in a few minutes. These a Crystals formed by divaricatic acid in G.E. Extracted from Ramalina solution ( x 160). 1111222221513. Figure 4. 138 ' PLATE II T i I; very fine curve yeniciilate bun tho-17., the t of thick yello to occur, usual some crystals. Psorom' tence under UV. color which app light. The aci benzene-dioxane acid is very si fluorescence is substance chron Ramalinolic Acf This (14 have examined ' is present in 3 sorediosa and 3 red, C+ red an few seconds to detected on th Hans and on fi necessary to p and apply the 139 very fine curved needles forming radiating clusters or penicillate bundles. If the extract is treated with G.A.o—T., the toluidine salt precipitates as bundles of thick yellow crystals. This reaction is very slow to occur, usually needing an overnight period to form some crystals. Psoromic acid has a strong yellow—green fluores- cence under UV. It reacts with PD producing a yellow color which appears yellow-orange under ultraviolet light. The acid shows a round spot with Rf .60- .62 in benzene-dioxane—acetic acid. The Rf value of psoromic acid is very similar to that of norstictic acid but the fluorescence is different. In toluene-acetic acid the substance chromatograms at Rf .38- .40. Ramalinolic Acid (Plate III, Figure 8) This depside is present in all the species I have examined that contain sekikaic acid. The substance is present in Ramalina peruviana, R. complanata, R. sorediosa and R inflata. It reacts K+ faint red, KC+ red, C+ red and PD-. These positive reactions take a few seconds to develop the red color which often is not detected on the lichen thallus, especially on old speci— mens and on fistulous species. In these cases it is necessary to prepare an acetone extract on filter paper and apply the reagent directly on the extraction. In a. subasperata analysis and by involves the ad which brings ab leaf-like cryst in scattered r stals seen sing Ramalin violet light. forms a round 5 solvent system The spot takes zidine . Salazinic Acid This '1. R'orcinol grou' ulla of R. de__n A. sorediantha Cessory in tho Its Color reac orange. The a the thallus iI microchemical G.A.o-T. to t] 140 R. subasperata the acid is only revealed by microcrystal analysis and by chromatograms. The microchemical test involves the addition of G.E. to the thallus residue which brings about the formation of small ellipsoid leaf-like crystals organized in radiating clusters form— ing scattered rosettes. Rarely are these colorless cry- stals seen singly. Ramalinolic acid fluoresces blue under ultra- violet light. In thin-layer chromatography the acid forms a round spot at Rf .30— .36 when developed in solvent system A and .06— .08 when in solvent system B. The spot takes a pink—red color when sprayed with ben— zidine. Salazinic Acid (Plate III, Figure 9) This is an aromatic aldehydic depsidone of the B-orcinol group. It is the major component in the med— ulla of R. dendroides, R. dendriscoides, R. complanata, R. sorediantha and R. straminea. The substance is ac— cessory in those species containing norstictic acid. Its color reactions are K+ red, PD+ yellow turning orange. The acid is easily extracted with acetone from the thallus in large amounts as a white residue. The microchemical test for this acid is the addition of G.A.o—T. to the dry extract. 141 PLATE III Crystals formed by psoromic acid in G.E. solu— Figure 7. tion ( x 100). Extracted from Ramalina gracilis. amalinolic acid in G.A.W. Figure 8. Crystals formed by r Extracted from Ramalina solution ( x 170). subasperata. Figure 9. Crystals formed by salazinic acid in G.A.orT- \ SOIUtlon ( X 70). Extracted from Ramalina ‘ peranceps. 74.; if I‘ 142 Crystal preparation and precipitates as mhr,either s nhtance is pr 0n chro donated spot andthat react TOtproduces z PDbUtgives a1 hsoccurred. 59bikaic Add This d 143 Crystals are formed after warming slightly the preparation and setting aside to cool. Salazinic acid precipitates as small boat—shaped crystals of a yellow color, either single or in small rosettes. If another substance is present, the crystals appear as blunt boats. On chromatograms this substance appears as an elongated spot at Rf .15— .17 in benzene-dioxane—acetic acid that reacts yellow with p—phenylenediamine. The spot produces a dark color under UV before spraying with PD but gives an intense orange color after the reaction has occurred. Sekikaic Acid (Plate IV, Figure 10) This depside of orsellic acid derivative occurs in association with ramalinolic acid. It is found in the medulla of R. inflata, R, peruviana, R, subasperata and R: sorediosa. Extraction of the acid requires more ace- tone than for any of the others. This acid reacts K-, PD-, C-, KC—. The extract yields a shiny, gummy residue after evaporation. When G.E. or G.A.W. is applied to the residue it recrystallizes as coarse colorless lamellae with parallel sides and square or angular corners. The Crystals occur scattered or associated in clusters of Plates radiating from a central point. Frequently in small groups of crystals pairs of lamellae show over- lapping of their tips. ‘W”"“‘ ' ' Sekikaii der ultraviolet round spot deve acid and at Rf acquires a red- when benzidine distinguishable Sekikai substances amor Usnic Acid (Pl: Usnic ; 1968i« it is 144 Sekikaic acid exhibits a blue color when seen un— der ultraviolet light. In a thin-layer chromatograph a round spot develops at .63- .65 in benzene—dioxane—acetic acid and at Rf .59- .51 in toluene-acetic acid. The spot acquires a red—brown color similar to divaricatic acid when benzidine is applied. In practice this spot is in— distinguishable from divaricatic and perlatolic acids. Sekikaic acid is one of the commonest medullary substances among the species in the genus. Usnic Acid (Plate IV, Figure 11) Usnic acid is a dibenzofuran derivative that gives the yellowish color common in many lichens (Huneck, 1968). It is the most common lichen acid, occurring in all of the Ramalina species studied. Together with atranorin and Chloroatranorin it is exclusively produced in the cortex. In R. cumanensis and in a population of R. usnea and R. camptospora, R. gracilis, it is the only chemical substance of taxonomic value produced by the thalli. Usnic acid promptly precipitates from most micro— Chemical reagents. The crystals adopt various forms that range from yellow angular lamellae to thin short needles, singly or in clusters. The acid is dark when exposed to ultraviolet light, appearing at Rf .68- .70 when chromatographed in benzene- dioxane-acetic acid or in toluene-acetic acid. Characteristica dehighest Rf atranorin. The reddish color w concentrated su Substance H (P1 This sc from @113 . °H1Y medullary W, a r sekikaic aCid : he C0101‘ reac The ac eraPbrat'lon th 145 Characteristically usnic acid occupies the position above the highest Rf for a medullary substance and just below atranorin. The acid does not react with PD, but gives a reddish color with benzidine, and a yellow color with concentrated sulfuric acid. Substance H (Plate IV, Figure 12) This substance was first reported by Evans (1943) from Cladonia. The substance has been shown to be the only medullary substance present in a variant of Ramalina complanata, a variant of R. usnea and in association with sekikaic acid in another variant of R, usnea (= R. subanceps). The color reactions are K—, KC—, PD—, C-. The acetone extract produces a shiny residue upon evaporation that forms identifiable crystals without using a recrystallizing agent. The crystals appear as long par— allel needles connected at different levels by short trans- versal plates. The parallel series of H—forming crystals bear here and there small clusters of crystals. If G.E. is applied to the extract it will recrystallize as long needles bent at the tips. Substance H does not chromatogram in developing systems recommended for fatty acids nor in systems for triterpenoids. 146 PLATE IV Figure 10. Crystals formed by sekikaic acid in G.E. solu— tion ( x 170). Extracted from Ramalina ' peruviana. Figure ll. Crystals formed by usnic acid in G.E. solution ( x 150). Extracted from Ramalina complanata. Figure 12. Crystals formed by substance H in G.E. Solu- tion ( x 45). Extracted from Ramali§§_232§3 _ rggggi CTTTW” , ,___--IIIII| I47 Unidentified Sr (Plate V, Figu This 5‘ of Ramalina g live with PD, sient red wine respect it beh acetone extrac benzene. Wher White gummy Sb 148 Unidentified Substance A (Plate V, Figure 13)’ This substance is present in a chemical variant of Ramalina complanata. Its color reactions are nega- tive with PD, K, and C but reacts KC+ producing a tran- sient red wine color turning orange. In this latter respect it behaves like cryptochlorophaeic acid. The acetone extract is difficult to obtain with acetone or benzene. When finally a residue is produced it is a white gummy substance that sticks firmly to the glass slide. Addition of G.E. or G.A.W. results in condensa- tion of oily drops that overnight precipitate clusters of colorless needles with a common central origin. The Crystals in the microchemical test seem to be due to a single medullary substance. Frequently the clusters originate slightly bent branches resembling perlatolic or imbricaric acids. It was shown that the angle of extinction although similar is not identical to that of perlatolic acid. This difference was confirmed by comv Paring the three acids in different solvent systems. When the acids were led to develop in benzene— dioxane-acetic acid the unknown showed a round spot at Rf .60- .62 which fluoresces blue and reacts with ben— Zidine exhibiting a brown color. The same substance in toluene-acetic acid shows two spots with a lower Rf value bUt with the same color reactions. The uPPer SPOt occurs at Rf .45- .47 and the lower one at Rf .32— .34. The che adepside type gram could repr distinguishable spots may reprr Unidentified 81 [Man V, Figu This u some specimens tain sekikaic not react with reaction. Acetor residue which in Clusters 5( Probable that SidnCe 3.5 ShOl When 149 The chemical behavior of this substance suggests a depside type structure. The two spots in the chromato— gram could represent either two different substances in— distinguishable by microchemical analysis or one of the spots may represent the hydrolysis product of the other. Unidentified Substance B (Plate V, Figure 14) This unidentified substance(s) only occurs in some specimens of Ramalina cochlearis, while others con— tain sekikaic and ramalinolic acids. Substance B does not react with PD, K or C but produces a KC+ red color reaction. Acetone extract of the thallus yields a gummy residue which precipitates colorless plates singly and in clusters somewhat resembling sekikaic acid. It is probable that the extract contains more than one sub— stance as shown by chromatography. When a sample of a thallus containing this sub- stance is chromatogrammed two yellow—orange spots are revealed by benzidine. If the chromatogram is developed in benzene-dioxane—acetic acid an upper round spot with blue fluorescence under UV appears at Rf .63- .65. A trailing spot with the same characteristics shows Rf .23- .25. Similarily the upper and the lower spot has Rf .51- .53 and Rf .21— .23 respectively in toluene-acetiC acid. When Ch] spots produce : acid. Unidentified 8 (Plate V, Figu Substa salazinic acid [PD+ variant) . chromatograms The cc C', and KC-, Produces nume: appear as hex. and do not sh is eaSilY Chr at Rf -58- .6 iiilefl Sprayed CUlor. 150 acid. When chromatograms are sprayed with benzidine the spots produce a yellow color similar to hypoprotocetraric acid. Unidentified Substance C (Plate V, Figure 15) Substance C is a PD— substance that accompanies salazinic acid in some specimens of Ramalina sorediosa (PD+ variant). It is detected by its blue color when chromatograms are exposed to UV light. The color reactions of substance C are PD-, K—, C-, and KC—. Crystallization of the extract in G.E. produces numerous minute crystals, forming prisms that appear as hexagonal lamellae. The prisms are colorless and do not show any type of aggregation. The substance is easily chromatographed in benzene-dioxane-acetic acid at Rf .58- .60 and in toluene-acetic acid at Rf .50— .52. When Sprayed with benzidine the spot assumes an orange color. Unidentified Substance D This unknown substance was found to be present in several specimens of Ramalina dendroides in associa— tion with salazinic acid. It chromatograms as a round SPOt above salazinic acid at an Rf of .18- .20 in . e benene—dioxane-acetic aCid. The fluorescence of substanc D is blue and it reacts orange with benzidine. Figure 13. Figure 14. Figure 15. 151 PLATE V unidentified substance A Crystals formed by Extracted from in G.E. solution ( x 130). Ramalina complanata. Crystals formed by unidentified substance B in G.E. solution ( x 150). Extracted from Ramalina cochlearis. Crystals formed by unidentified substance C in G.E. solution ( x 600). Extracted from D+ variant of Ramalina sorediosa. Crystz UE.solution appear scatte: D reacts PD- , Role of Chemi the Taxonomy Chenogenes is Curre Plain the ocC 153 Crystals of this substance were obtained in the G.E. solution. These are small colorless lenticels that appear scattered without forming aggregates. Substance D reacts PD—, K, C- and KC-. Role of Chemical Substances in the Taxonomy of Ramalina. Chemogenesis Current chemogenetic hypotheses attempt to ex- plain the occurrence of depsides and depsidones in the medulla of the lichen thallus. W. L. Culberson (1967) analyzing the European population of Ramalina siliquosa, suggested that the salazinic acid producing population gave rise to a hypoprotocetraric acid containing strain. By a single step reaction this evolved into a population with protocetraric acid. Through another chain of re- actions the original salazinic acid containing stock pro- duced a population in which salazinic acid was replaced by norstictic acid. Reduction in medullary synthesis reached its extreme in a derived population in which no chemical synthesis occurred. As for stictic acid con— taining plants, they could have originated from the salazinic acid original population or from the norstictic acid variant. Follmann and Huneck (1969) proposed a more gen~ eralized scheme for possible pathways in the synthesis of depsides an of divaricatic the results of divaric acid, carbonic acid ramalinolic ac ways can be so divaricatinia 154 of depsides and depsidones. They visualized the synthesis of divaricatic acid, sekikaic acid and ramalinolic acid as the results of esterification of divaricatinic acid with divaric acid, 3-hydroxydivaricatinic acid, and 3-hydroxy~ carbonic acid respectively. Through methylation of ramalinolic acid boninic acid is obtained. These path- ways can be summarized as follows: + Divaric acid .___——————9divaricatic acid divaricatinia acid + 3—hydroxydivaricatinic-9Sekikaic acid + 3-hydroxycarbonia acid—)Ramalinolic acid Boninic acid Follmann and Huneck considered lecanoric acid as the substrate from which different pathways may lead to lichen acid syntheses. Through methylation of the lecanoric aCid molecule evernic acid and obtusatic acid are obtained. Chlorination of lecanoric acid changes it into tumidulin. A methylation in each of the rings of lecanoric acid transforms it into 4-0—demethylbarbatic acid. This acid is in turn precursor of the cortical depside atranorin and the medullary depsidone hypoprotocetraric acid. Partial oxida- tion of hypoprotocetraric acid yields the intermediate virensic acid. One important pathway in the synthesis of PD+ substances originates by successive partial oxidation reactions star‘ tion produces j to form salazi‘ of the interme methylation of foregoing sche lecanoric acid 1 4‘0-demethylba l Hl'thlprotocetrz d lirensic acid l XorSt'lCtiC ac 155 reactions starting with virensic acid. The first oxida— tion produces protocetraric acid, which in turn oxidates to form salazinic acid. In another synthesis methylation of the intermediate produces norstictic acid. Further methylation of norstictic acid gives stictic acid. The foregoing scheme could be diagrammed as follows: Lecanoric acid ________) Tumidulin 4-0-demethylbarba;;:::;:id~““““9 Evernic acid U l Hypoprotocetraric acid Obtusatic acid Virensic acid Atranorin Norstictic aci;““““‘r~.“~.‘..~9 Protocetraric acid U d Stictic acid Salazinic acid C. F. Culberson (1969) proposed an acetate poly— malonate pathway for lichen substances syntheses which can produce phenolic derivatives through an orsellinic acid-type cyclization. The commonest phenolic acid units derived through this pathway are the orcinol-type units and the B-orcinol—type units. Accordingly, this scheme provides a mechanism through which the three chemical variation patterns present in Ramalina could be explained. Distribution of Lichen Substances 1n Ramalina In an exhaustive review of lichen chemistry C. F, Culberson (1969, 1970) reported a total of 28 primary and secondary subs (1969) reporte atotal of 76 sents a more c Florida specie The list conta stances. (See The 51 in the genus l M and l recognizing tl “Uplanatae p l'elopnent wit and ComPressi The p aPleared ackw 156 secondary substances from 96 taxa. Follmann.and Huneck (1969) reported on the chemistry of 32 species and listed a total of 76 species with known chemistry. Table 16 pre- sents a more complete list to which the West Indies and Florida species and their chemical content has been added. The list contains 114 taxa containing a total of 35 sub- stances. (See Appendix C.) The species placed by Follmann and Huneck (1969) in the genus Desmazieria are here restored to the genus Ramalina and Desmazieria is treated as a section. I am recognizing the series Cylindricae Follm. and Hun. and Complanatae Follm. and Hun. to indicate the parallel de- velopment with the series Tretiusculae (Vain.) Zahlbr. and Compressiusculae (Vain.) Zahlbr. The placement of R. peruviana in Desmazieria appeared ackward and it has been removed and placed in Ramalina. Several species cited by Follmann and Huneck (1969) and C. F. Culberson (1969) have been reevaluated in accordance with the results of the present study. Ramalina dasvpoga has been considered synonomous with R. furcellata, R. ecklonii with R. cumanensis, R. geniculata with R. inflata, R. calicaris var. fastigiata with R. fastigiata. The trio of species formed by R. reagens, R. hypoprotocetrarica, and R. subfarinacea are considered chemical variants of R. farinacea. The sp as having a fa l. Ihave exa and demons trat the data giver substance H cc mentioned by I stances in tl .ests, CTYSt The t 157 The species Ramalina evernioides is referred to as having a fatty acid complex and an unknown substance X. I have examined several specimens of this species and demonstrated substance H (see Evans, 1943). From the data given it is not possible to determine if this substance H corresponds to any of the unknown substances mentioned by Follmann and Huneck (1969). Summary Although chemical reactions have been used for a long time as taxonomic characters, taxonomists have not reached agreement as to the rank that should be assigned to morphologically similar individuals with different chemical composition. In this study the chem— ical data has been used to complement morphological in- formation to characterize the species. Otherwise, iden— tical populations but chemically distinct are referred t0 as variants. Morphological, ecological, or geograph- ical differences would warrant species recognition. The results of this investigation reveal that in the West Indies the genus Ramalina is represented by 23 sPecies which produce a total of 17 different lichen sub- stances in the thallus. These were identified by color tests, crystallography and chromatography. The medullary substances produced by the West Indian species fall in the chemical categories Of dePSIdes (divaricatic a cryptochloroph protocetraric fatty acid (ca cortical subst and Chloroatrz products repm C. Culberson 1% morPhological Plosive eXpre tionary trend than to singl and Huneck (1 158 (divaricatic acid, sekikaic acid, ramalinolic acid and cryptochlorophaeic acid); depsidones (norstictic acid, protocetraric acid, psoromic acid and salazinic acid); fatty acid (caperatic acid) and unknown substances. The cortical substances are usnic acid and the depsides atranorin and Chloroatranorin. Appendix B contains a list of natural products reported from Ramalina compiled from data given by C. Culberson (1970). Ramalina represents a genus in which diversity in morphological development has been accompanied by an ex- plosive expression of biochemical syntheses. This evolu- tionary trend, however, relates to groups of species rather than to single species. In section Desmazieria, Follmann and Huneck (1969) found a positive correlation between the presence of ceruchinol and tumidulin and the cortical mor— Ph010gy. Similarily there is a trend for sekikaic acid to be concentrated in the section Fistularia. The alliance of species with papillae C3: EQEELEEEEE, 3. subasperata, R. attenuata, R. furcellata, R. sgggfliggg) typically Contains salazinic acid. The species R. sub— EEllEEléE and R. montagnei, which have some morphological traits in common also have divaricatic acid in the medulla. Three patterns of chemical variation are discovered in Béflflliflg. The commonest pattern is the replacement of a medUllary substance by a different depside or depsidone. The second chemical pattern is the presence of a dep51de- depsidone pair Culberson, 196 acid and hypop in Ramalina, b same species. in Ramalina g of orcinol dei the same thall (either with s 159 depsidone pair with exactly corresponding structures (C. Culberson, 1969). In this group is 4-0-demethylbarbatic acid and hypoprotocetraric acid. Both substances occur in Ramalina, but have not been found together in the same species. The third type of chemical pattern occurs in Ramalina subasperata. In this species a meta-depside of orcinol derivative (sekikaic acid) occurs together in the same thallus with a depsidone of the B-orcinol type (either with salazinic acid or protocetraric acid). Diagno green, frutico branching dich latter either one or two lay be constituted oriented hyphg inner is longj formed by sepz to long axis < by an arachno: 160 Taxonomic Treatment Diagnosis of genus: Thallus stramineous, rarely green, fruticose, erect to pendent, variable in size; branching dichotomous, branches terete or compressed, the latter either flattened or channeled. Cortex formed by one or two layers; if only one layer is present, it could be constituted either by longitudinally or transversally oriented hyphae; if the cortex is formed by two layers, the inner is longitudinally arranged and may be continuous or formed by separate bundles, the outer layer is perpendicular to long axis of the branches. The medulla is constituted by an arachnoid tissue of loosely interwoven hyphae, without a central chondrioid core. The algae are Trebouxioid and confined to a ring around the medulla, giving the thallus its heteromerous character. Apothecia common, margins concohnous, disks tan to brown; asci clavate, containing eight hyaline, one-septate spores. Pycnidia inconspicuous, dark; conidia cylindrical, exobasidial. Sub-generic Categories Section Desmazieria (Mont.) Stiz. Cortex formed by one layer of thick hyfiuw perpendicular to the long axis of the branches. Subsection Solidae Du Rietz. Medulla present. Series Complanatae Follm. & Hun. Branches flattened. Series Cylindricae Follm. E Hun. Branches terete. Subsection Cenozosia (Mass.). Medulla absent. Seaion Ecortii orientr Rdion Ramalii Subsection M' Series Ter Series Com presse finsection F branch luificial Ke MW IIThallus fig 1 Thaluus not 1 Sorediar L Esoredi; 3~Soredia in: SI SOTEdia no. 4' Bundle: 4- Branche 3- Soredia in ofbranche alsoredia no Ut disper 6' Lacinia h Lacinia soredia .‘ NIEdulla 13+ .‘ Medulla P- 8.Tha11uE Spores S. ThallUS Sterile 161 Section Ecorticatae Stein. Cortex formed by longitudinally oriented hyphae. Section Ramalina. Cortex formed by two hyphal layers. Subsection Myelopoea (Vain.) Zahlbr. Medulla present. Series Teretiusculae (Vain.) Zahlbr. Branches terete. Series Compressiusculae (Vain.) Zahlbr. Branches com- pressed. Subsection Fistularia (Vain.) Zahlbr. Medulla absent, branches swollen. Artificial Ke to Ramalina S ecies Occurring 1n the West Indies and Florida 1. Thallus fistulous, foraminous..... ........ ..... R. inflata l. Thaluus not fistulous, not foraminous........... ..... ... 2 2. Sorediate ...... . ................. ..... .......... ..... 3 2. Esorediate ..... ... ........ . .......... .. ...... . . 10 3. Soredia inside swollen, helmet-like soralia..R. cochlearis 3. Soredia not inside swollen, helmet-like soralia ...... ... 4 4. Branches terete or subterete .............. . ....... ... 5 4. Branches never terete or subterete ............. ...... 6 5. Soredia in globose heads at the tips of branches.. ............................ R. dendriscoides 5. Soredia not in globose heads at the tips of branches, but dispersed throughout the branches ...... .. R. sorediosa 6. Laciniae canaliculate, soredia apical... R. sorediantha _-——_—~__ 6. Laciniae not canaliculate, flattened, soredia marginal ....................... . ...... ....... 7 7. Medulla P+; salazinic acid present ............... . ..... . 8 7. Medulla P—; salazinic acid absent ................. . ..... 9 8. Thallus pendulous, to 30 cm. long, usually fertile, spores large (ll—19 x 4—6u) ..... . ....... ..R. dendroides 8. Thallus not pendulous, to 5 cm. long, usuall sterile, spores small (lO—lZ x 3-6u) ...... R. farinacea N fig=zw - . h 9. Medulla K+ present. . . 9. Medulla 1(- absent. . .. 10. Surfac 10. Surfac 11. Branches c canaliculz 11. Branches r 12. Meduli presei 162 9. Medulla K+; sekikaic and ramalinolic acids present.................................... R. peruviana 9. Medulla K-; sekikaic and ramalinolic acids absent ................................... R. camptospora 10. Surface papillate ................................ ll 10. Surface not papillate... .......... . .............. 17 ll. Branches canaliculate or becoming canaliculate .............................. R. complanata ll. Branches not canaliculate, not becoming canaliculate. 12 12. Medulla C+; cryptochlorophaeic acid present ................................. R. paludosa 12. Medulla C-; cryptochlorophaeic acid absent ....... 13 13. Upper part of branches nodulose ........... R. furcellata 13. Upper part of branches not nodulose..... ........... .. l4 l4. Thallus fruticulose, branches tapering terminally ....................................... 15 14. Thallus not fruticulose, branches not tapering terminally, semi—pendulous ....................... 16 15. Thallus dark brown, main branches with small lateral branchlets. Florida .................. R. willeyi lS. Thallus pale, branches without lateral branchlets. West Indies .............................. R. subasperata 16. Medulla P+; salazinic acid present ..... R. attenuata l6. Medulla P-; salazinic acid absent ...... R. montagnei l7. Laciniae canaliculate or becoming canaliculate ....... 18 17. Laciniae not canaliculate ............................ 20 18. Medulla P-; usnic acid only ........... R. cumanensis 18. Medulla P+; norstictic or salazinic acid present. 19 19. Norstictic acid present ...... . ........... R. 13232222323 19. Salazinic acid present ..................... R. straminea N m. Norstir 20. Norstir n.8alazinic . H.Norstictic 23.Tha11u lacini Zl Thallu lacini 24-Spore sept 34' Spore sept ZR Spores 2i Spores 26-Divaricatf 26'DiVaricat: 163 20. Norstictic or salazinic acid present ............. 22 20. Norstictic and salazinic acids both absent ....... 23 22. Salazinic acid present... .................. R. peranceps 22. Norstictic acid present ....................... R. anceps 23. Thallus pendulous, to 35 cm. long, laciniae contorted ......................... R. usnea 23. Thallus not pendulous, to 10 cm. long, laciniae flattened ............................... 24 24. Spore septum oblique ........................ R. bistorta 24. Spore septum straight ................ . ............... 25 25. Spores ellipsoid, 9-13 x 4-5u ......... R. fastigiata 25. Spores fusiform, 16-22 x 3—5u .................... 26 26. Divaricatic acid present ................ R. subpellucida 26. Divaricatic acid absent ................... R. stenospora Section Ramalina Subsection Myelopoea (Vain.) Zahlbr. Series Teretiusculae (Vain.) Zahlbr. 1. Ramalina attenuata (Pers.) R. H. Howe Physcia attenuata Pers. Ann. Wetterau. Ges. Nat. 2: 18. 1811. Ramalina attenuata (Pers.) R. H. Howe Bryologist 17: 35—36. 1914. Original material: Dominican Republic (non vidi). Lichen rigidus Pers. in Ach. Syn. Meth. Lich. 294, 1814. Ramalina rigida Ach. Syn. Meth. Lich. 294. 1814. Original material: Dominican Republic (UPSE, Si), _;,kq7:fig.;h Des cri‘ _——-—4 high, much bra papillate, sor across, subped hymenium 40-80 taining 8 Spor one-septate, s Medull red, Kc.) (3-. Lichen \ ”5111C acid. Discus \ based On One 5 and cited Phys aQuestion Ina] Nylanr of 13 M I in the POSim the SPecies R W and ] Ll lbUtioH g1“ 164 Description: Thallus stramineous, small, to 5 cm. high, much branched, branches subterete or terete; surface papillate, soredia lacking, Apothecia abundant, small, 2-3 mm. across, subpedicellate; disk dark; epithecium 6—10u thick, hymenium 40—80u thick; hypothecium 10-20u thick. Asci con— taining 8 spores arranged in two series; spores hyaline, one-septate, subfusiform, 11-14 x 2.8-4.7u. Medullary reactions: PD+ orange-yellow, K+ yellow- red, KC—, C-. Lichen substances: Atranorin, salazinic acid and usnic acid. Discussion: Acharius (1814) described R. rigida based on one specimen typified by Person as Lichen rigidus and cited Physcia attenuata as a possible synonym but added a question mark to the name. Nylander (1870) considered R. gracilis Nyl. synonym 0f R. rigida Ach. Tuckerman (1882) adopted a different View in the position of the North American species. He considered the species R. gracilenta, R. rigida var. montagnei, R. gracilis and R. tenuis synonymous with R. rigida. The dis- tribution given by Tuckerman for R. rigida, together with the fact that R. montagnei was described from Cuba, points to the possibility that his confusion was produced by not recognizing the species later described as R. willeyi by R. H. Howe. Ramalina attenuata was elucidated by R. H. Howe in 1914. During fertile specim teristics of R than those giv marked as isot another from S fragment of a the two collec of 1}. M n W. Br considering 3 name g, M Current rules The 51 Ori5111211 mate 165 During the course of this study I have examined a fertile specimen from Jamaica with the morphological charac- teristics of R. attenuata but with spores broader (2.8-4.7u) than those given by R. H. Howe (2.5-3.5u). Two packets marked as isotypes of R. rigida (one from herb. UPS and another from S) were studied. Both contained a single fragment of a sterile, poorly preserved specimen. However, the two collections examined matched very well the concept of R. rigida which in turn fitted the description of R. attenuata. Both species appear to be the same. Thus, I am considering R. rigida synonymous with R. attenuata. The name R. attenuata is the legitimate name according to current rules in nomenclature. The small number of collections and the lack of original material for comparison leave some doubts about the validity of this species. It may remain that better collections from the type locality (Dominican Republic) show that R. attenuata is synonymous with R. furcellata. Ramalina attenuata can be separated from R. furcellata by the presence of nodules at tips of branches. Ecology: Occurs at elevations between 1000 ft. and 2300 ft., which is the region of high humidity in the West Indies. Distribution: Ramalina attenuata was described from Dominican Republic and R. H. Howe (1914) reported it from Jamaica and Puerto Rico. Materi area on hillsi 2300 ft., Dept JAMAICA: Long St. Andrew, R 2. Ramalina 5 w red. to C Nllc acid . M hen Close t lished by t M be Confu 166 Material seen: West Indies. HAITI: Cultivated area on hillside below Citadelle, south of Millot, ca. 2300 ft., Dept. du Nord, Wetmore RREQ, 1958 (MSC). JAMAICA: Long Mountain, Mona, ca. 1000—1300 ft., Parish ‘St. Andrew, Imshaug 13199, 1952 (MSC). 2. Ramalina dendriscoides Nyl. Ramalina dendriscoides Nyl. Flora 59: 412. 1876. Lectotype (nov.): Cuba, Wright ZERR (PHI). Description: Thallus shrubby, small, to 5 cm. high, branching irregularly dichotomous, branches terete or sub- terete at the axils, dendroid, to 2 mm. broad at the base; surface smooth; soredia granulose, soralia small, semi- globose, on tips of terminal or short lateral branchlets; cortex thick, cartilaginous. Apothecia rare, lateral, 0.4-4.0 mm. across; disk pale; asci containing 8 spores in a single row; spores uniseptate, hyaline, ellipsoid, straight or slightly bent 6-16 x 3—3.Su. Medullary reactions: PD+ orange—yellow, K+ orange- red, KC—, c—, Lichen substances: Atranorin, salazinic acid and usnic acid. Discussion of the species: This Cuban species is very close to Ramalina attenuata, from which it is distin- guished by the presence of soredia. Ramalina dendriscoides can be confused with the sorediate species R. peruviana and b sardine by its PD+ ree vi_an_a, and fra globose soral: R_. peruviana. species is thy contrast to t' seen numerous thich C two species a R‘ . Wides. \ Two p ““9 Studied. bright 738" n 167 R. sorediosa. However, it can be separated from the former by its PD+ reaction instead of the PD— reaction of R. peru- viana, and from the latter by its characteristic semi- globose soralia rather than the maculiform type found in R. peruviana. An additional criterion to segregate these species is the shrubby growth form of R. dendriscoides in contrast to the pendulous habit of R. sorediosa. I have seen numerous herbarium specimens identified as R. soredi— antha which clearly belong into R, dendriscoides. These two species are separated by the canaliculate branches of R. sorediantha against the terete branches of R. dendris— coides. Two packets marked as types from herb. FH and US were studied. Both packets were labelled ”on palm in Cuba, Wright 738" with what appeared to be authentic handwriting by Nylander. Each packet contained several specimens with terete branches and semi—globose soralia on terminal branches. Since the author did not give indications as to the selection of a holotype, I am designating the specimen marked 738a (FH) lectotype, mainly because it is abundantly sorediate. The original description of the species did not include the chemical reactions of the type collection. I was fortunate to examine the type material and demonstrated it to be PD+ yellow—orange and K+ yellow red. Thin—layer Cromatograms developed in benzene—dioxane-acetic acid (90:25:4) of of salazinic be demonstrat infrequent in Speci A. dendriscoi tained sekika ll. peruviana. the flora of Ecol elevations r the pine for dominant epir montane rainl associated wi Teloschistes Species belor hollow thall ition with R 168 (90:25:4) of the original material demonstrated the presence of salazinic and usnic acids. Although atranorin could not be demonstrated in the type material, this substance is not infrequent in robust thalli of Puerto Rican material. Specimens reported from Florida by Moore (1968) as R. dendriscoides were confirmed to be PD-, K+ pink and con- tained sekikaic and ramalinolic acids and were identified as R. peruviana. Therefore, the species should be removed from the flora of Florida. Ecology: The species occurs in the West Indies at elevations ranging from 1200 ft. in Jamaica to 5300 ft. in the pine forests of Haiti. In Puerto Rico the species is the dominant epiphyte in the Psidium guajava scrub in the lower montane rainforest. Ramalina dendriscoides is frequently associated with R. subasperata, R. complanata, R. anceps, Teloschistes flavicans and species of Usnea. These Usnea species belong to the red cortex species and those with a hollow thallus. Other macrolichens found growing in associ- ation with R. dendriscoides are species of Heterodermia and Parmelia. The species also grows abundantly on the rough trunks of Erxthrina, Inga and other coffee shade trees. Along roads it is commonly seen on trunks of Spathodea cam— panulata and on fence poles. In rare occasions small Specimens were collected on calacereous rocks along with Cladonia and Heterodermia. lndian-Centra gave the dist subtropics . " In th paniola, Puer and reappears Costa Rica in The follows the central part toward the e abundant near less common i tribution of as east-west [Rig rainforest, L691, 1967 c 16125, 1967 1111A, 1722 169 Distribution: Ramalina dendriscoides has a West Indian-Central American distribution. Zahlbruckner (1930b) gave the distribution as "meridional Europe, tropics and subtropics.” In the West Indies the species occurs in Cuba, His- paniola, Puerto Rico and Jamaica, skips the Lesser Antilles and reappears in Trinidad. It has also been recorded from Costa Rica in Central America. The distribution of the species in Puerto Rico follows the central mountain range, being abundant at the central part of the island and decreasing in frequency toward the eastern and western ends. It is especially abundant near Aibonito, Barranquitas and Toro Negro Forest, less common in Luquillo and Maricao Mountains. The dis- tribution of R. dendriscoides shown on Figure 7 is described as east~west highland. Material seen: West Indies. PUERTO RICO: Luquillo rainforest, Landron 1176C, 1967 (MSC); Guavate mountains, Landron 1331, 1332, 1358D, 1967 (MSC); Maricao, Landron 1697, 1967 (MSC); Da. Juana, Toro Negro, Landrén 1686C, 1690A, ~— 1692A, 1967 (MSC); Barrio Caonillas, Villalba, Landrén 1712, 1716A, 1722A, 1723B, 1967 (MSC); Da. Juana, near road, Toro Negro, Landron 2310D, 2316, 2324, 2349C, 1968 (MSC); Divisoria ) Toro Negro, Landron 1738A, 1739, 1741B, 1743D, 1745B, 1949 .__—__—_———____________a 1967 (MSC); Toro Negro, pasture with scattered trees, Landrén 1811B, 1818A, 1819A, 1926A, 1833A, 1837A, 1838, 1839, 1841 ‘ _—_. —_ _...—- ._.__._. _____ _.___. —_—_ ———., an, M: 1848A, 1 lake, Landrén 3561, 2 landron 23733, 23826, luchillas, Landrén l7 (MSC); Carite Lake, L Cuba, Barranquitas , L 1891A, 1892A, 1894A, -__——_—- 1911A, 1912A, 19131;, ~——————— RSR, 1928B, 1936C, 3551967 (MSC); Ce pug, 1971A, 1979A, 11995222.» 19933, 1967 (use); grim; 182 on (an); unlit. iii 1932 (FH); upper slop Maestra, Oriente, 1111‘! Armenia, 'Loma del Gal 241951959 (MSC); la Sierra Maestra, Oriel facing the sea, near Sierra Maestra, Orie DOMINICAN REPUBLIC: tiago and La Cumbre» 1958 (MSC); valley [ordillera Septentr‘ Délélrtment de l ‘Oue 170 1842A, 1847, 1848A, 1852, 1854C, 1967 (MSC); Utuado, near ——————.-———. lake, Landrén 2262, 2272A, 1968 (MSC); Adjuntas, Cerro Punta, Landrén 2373B, 2382C, 2395B, 1968 (MSC); Orocovis, Barrio Cuchillas, Landr6n 1757A, 1758B, 1762B, 1764A, 1798C, 1967 (MSC); Carite Lake, Landr6n 1865B, 1881B, 1967 (MSC); Monte Cuba, Barranquitas, Landrén 1884B,1885A,1886A,1887A,1888D, __—.—””———__—_——_~_———_ 1891A,1892A,1894A,1896A,1903B,1905B,1907,1909A,1910A, ———_W___——.——_——.——_———_——____ 1911A,1912A, 1913B, 1915B,1916A,1917C,1923A,1925A, 1926A, —_.’W——_———_.—___—_—.—_._—____— 1931C,1928B, 1936C,1940A,1941A,1944A,l946,1948B, 1950A, —_W—___——__—_—__—_—_-_——. 1951, 1967 (MSC); Cerro Pulguillas, Aibonito, Landrén 1965A, 1968A,1971A,1979A,1980A,1982A,1983B,1985A,1986A,1987A, ~W____—_—_———.—__——_—_ 1990A,1992, 1993B, 1995B,1996A,1997C,1999C, 2001, 2006D, WW.——_—__——_—__—-—___——— 1967 (MSC); Fink 1823, 1937, 1916 (MICH). CUBA: Wright 738B 152 (FH); Wright 738 (US); Guantanamo, Bro. Hioram 10026, 1932 (PH); upper slopes of San Juan, Loma del Gato, Sierra Maestra, Oriente, Imshaug 24721, 24731, 1959 (MSC); near Armenia, Loma del Gato, Sierra Maestra, Oriente, Imshaug 24793, 1959 (MSC); lower slopes of San Juan, Loma del Gato, Sierra Maestra, Oriente, Imshaug RRRRR, 1959 (MSC); slope facing the sea, near Colegio de la Salle, Loma del Gato, Sierra Maestra, Oriente, Imshaug EERRR, 1959 (MSC). DOMINICAN REPUBLIC: Cerrazo, 2450 ft., ridge between San- tiago and La Cumbre, Cordillera Septentrional, Wetmore RRRQ, 1958 (MSC); valley around El Chorro, near Constanza, 3700 ft., Cordillera Septentrional Wetmore 3814, 1958 (MSC). HAITI: Department de l'Ouest, summit of Montague Noir, ca.5500 ft., near Kenscoff, Wetmor ridge above Kenscoff, summit of Téte Etang 22 6000 ft., M dos Pins (SHADA stat' JAMAICA: Parish of M14154, 1953 2600 ft., Imshaug 13 Mt. Diablo, Imshaug Hall Cave, near Guys (MSC); along Mount D EM, 1952 (MSC); W Imshaug 15829, 1953 Blanchisseuse Road," Imshaug 31743, 31752 W and Cartago, ca. 124 [MlCH] . 3. Ramalina furcell Everni a fur c 9: 236. 1845. Rama] Univ. 6: 489. 1930. Ramalina £133 28: 203. 1859. Leo Cuba,Wright 49C, A] 171 near Kenscoff, Wetmore Rzgg, 1958 (MSC); north slope of ridge above Kenscoff, 5000 ft., Imshaug 22131, 1958 (MSC); summit of Téte Etang on ridge between Kenscoff and Furcy, 6000 ft., Imshaug gfigfi, 1958 (MSC); low hills west of Forét des Pins (SHADA station), Imshaug E9919, 1958 (MSC). JAMAICA: Parish of St. Andrew, Cooper's Hill, 2450 ft., Imshaug 11111, 1953 (MSC); Parish St. Catherine, Mount Diablo, 2600 ft., Imshaug REZQR, 1952 (MSC); between Hollymount and Mt. Diablo, Imshaug 13326, 1953 (MSC); Parish St. Ann, Mosley Hall Cave, near Guys Hill, 2000 ft., Imshaug 13678, 1952 (MSC); along Mount Diablo—Hollymount Road, 2600 ft., Imshaug 13756, 1952 (MSC); Walkers Wood to Moneague, 1200 ft., Imshaug 15829, 1953 (MSC). TRINIDAD: ridge west of ”Arima- Blanchisseuse Road,” along Las Lapas Road, Northern Range, Imshaug 31743, 31752, 1963 (MSC). Central America. COSTA RICA: Between Aguas Caliente and Cartago, ca. 1240-1400 m., Dodge and Thomas 7094, 1930 (MICH). 3. Ramalina furcellata (Mont.) Zahlbr. Evernia furcellata Mont. in Sagra, Hist. Cuba. Bot. 9: 236. 1845. Ramalina furcellata (Mont.) Zahlbr. Cat. Lich. Univ. 6: 489. 1930. Original material: Cuba (non vidi). Ramalina dasypoga Tuck. Am. Journ. Arts Sci. 11. 28: 203. 1859. Lectotype (nov.): Farallones, Monte Verde, CUba, Wright 49C, April 2 (no year given) (FHl). Description: no cm. long, much b tolmm. in cross se surface nitid, tuber frequent, 1-3 mm. ac pruinose; hymenium ZI containing 8 spores tate, hyaline , ellip- Medullar re 1+ yellow~red or PD~ Lichen subst traces of norstictic divaricatic acid and 42, FH, US) of the 0 Each packet containe annotation K+ yellow the author's intenti 1am designating the packet the lectotype in this packet this 0“} dasypoga. H01 studied matches verj mentioned species . related to R. usneo Alectoria . A care f 172 Description: Thallus pendent, filamentous, fragile, to 6 cm. long, much branched; filaments terete throughout, to 1 mm. in cross section, tips filiform and nodulose; surface nitid, tuberculate, cortex rigid. Apothecia in- frequent, 1-3 mm. across, marginal, concave; disk pale brown, pruinose; hymenium 20—30u thick, paraphyses branched. Asci containing 8 spores arranged in one series; spores unisep- tate, hyaline, ellipsoid, substraight 10-15 x 3-4.5u. Medullary reactions: KC—, C-, PD+ yellow-orange, K+ yellow-red or PD—, K-. Lichen substances: Atranorin, salazinic acid, traces of norstictic acid in thin—layer chromatograms, divaricatic acid and usnic acid. Discussion: I have examined three packets (Wright 42, FH, US) of the original material of Ramalina dasypoga. Each packet contained several thalli with one containing the annotation K+ yellow. However, there is no indication of the author's intention of selecting a type. For this reason I am designating the larger specimen (Wright 42C) in the PH packet the lectotype. Out of the four specimens included in this packet this one represents best the original idea 0f R. dasypoga. However, all the Cuban material I have studied matches very closely the type description of both mentioned species. Tuckerman believed R. dasypoga to be related to R. usneoides Fr. and to the genus Usnea and Alectoria. A careful study showed that R. usneoides .‘ (=R. usnea R. H. How with rather pendulou l‘halline morphology the species belong Ramalina fur but it can be separa The type mat diflgB. de Lesd. ha tinct species, mainl and wider distributi tuted by two chemic which contained sal to the island of Cu This PD— variant is because except for t two variants cannot grounds. ECOlOgX: T1 trees and rocks in I described R. 1233232 hl’grophillous, theI‘ Test Indies R. £135 occurring at its 1C its highest range 5 is included the An‘ 173 (=R. usnea R. H. Howe) is from a different complex of species with rather pendulous thallus and long fusiform spores. Thalline morphology and spore characters, likewise, indicated the species belong under the genus Ramalina. Ramalina furcellata may be confused with R. attenuata but it can be separated by its torulose branches. The type material of Ramalina dasypoga var. sore- diosa B. de Lesd. has been included in this paper as a dis— tinct species, mainly on the basis of its sorediate thallus and wider distribution. Ramalina furcellata is here interpreted as consti— tuted by two chemically diverse populations. A PD+ variant which contained salazinic acid and a PD— variant restricted to the island of Curacao which produced divaricatic acid. This PD~ variant is included within the species R. furcellata because except for the difference in chemical contents, the two variants cannot be distinguished on morphological grounds. Ecology: The original material was collected on trees and rocks in Cuba. Follman and Huneck (1969, p. 189) described R. dasypoga as being ”suboceanic, acidophillous, hygrophillous, thermophillous and photophillous." In the West Indies R. furcellata is typical of high elevations, occurring at its lowest point in Haiti (2400 ft.) and at its highest range in Jamaica (6000 ft.). Within this range is included the Antillean montane forest zone, where rainfall is very ab usually covers the m Distribution tally collected in t Antilles, and in Mar Antilles. Curacao 1 acid containing vari A. furcellata was c toward the central highland distributi is shown in Figure Material se dePunta, 200 m. fr forest, Landrén 239 Hioram R8513, 1923 ( Mayra, 1918 (FH); : R9. Hioram, s.r_1_., . shrubs in coastal t Farallones, Monte V rocks in mountains , on slope of Lorna de 1959 (MSC). HAITI: Kenscoff, Départmea (MSC); in elfinwoo SavaneZombie, DéP (MSC); summit of T 174 rainfall is very abundant throughout the year and mist usually covers the mountain peaks. Distribution: This Cuban species is characteristi- cally collected in the four islands comprising the Greater Antilles, and in Martinique and Curacao in the Lesser Antilles. Curacao is the only place where the divaricatic acid containing variant has been collected. In Puerto Rico, R. furcellata was collected in the montane rainforest zone toward the central part of the island. Thus the central highland distribution found for the species in the island is shown in Figure 8. Material seen: West Indies. PUERTO RICO: Cerro de Punta, 200 m. from the summit, in palm and tree ferns forest, Landron 2391B, 1968 (MSC). CUBA: Las Pailas, Bro. Hioram 6869, 1923 (US); Novaliche, Oriente, Bro. Hioram, 3.3., May 16, 1918 (PH); Finca Santa Rosa, Novaliche, Oriente, R33. Hioram, 5.3., Jan. 15, 1920 (EH); Santiago de Cuba, shrubs in coastal thicket, Bro. Le6n 3.3., 1924 (MICH); Farallones, Monte Verde, Wright RR, 39 (FH); on trees and rocks in mountains, Monte Verde, Wright RR (US); Armenia 0n slope of Loma del Gato, Sierra Maestra, Imshaug 24833, 1959 (MSC). HAITI: summit of Montagne Noir, 5500 ft. near Kenscoff, Department de l'Ouest, Wetmore 2787, 2792, 1958 (MSC); in elfinwoodland, along road from Forét des Pins to Savane-Zombie, Department de l'Ouest, Wetmore 3059, 1958 (MSC); summit of Téte Etang, 6000 ft., between Kenscoff and Furs» 115228 ridge, summit north 5000 ft. , Im‘sh’aug‘ Cushman 4_8_, 1912 (PH 22 1028 (US); along Mo 5413,1952 (MSC); Pa and Mount Diablo, ca (MSC); Parish of St. l_5_9_2l, 15922, 1952 ( ca. 5250 ft. , ridge Imshaug 23705, 1958 from Deux Choux to 1963 (MSC). CURACA AM, 5.3., 1917 ( Central Ame tazén, 900-1000 111. , 0- Ramalina soredit Ramalina _c_1_§ Rev. Bryol. Lich. 7 south Guantanamo Na 5.13.,Jan. 1933 (FE Descriptiol prudent, to 10 cm. out, to 1 mm. in c Surface nit id , gra 175 and Furcy, Imshaug 22595, 1958 (MSC); in mist forest on ridge, summit north of Forét des Pins (SHDA station), ca. 5800 ft., Imshaug 22704, 1958 (MSC). JAMAICA: Mandeville, Cushman fig, 1912 (PH), Cushman, 3.2. (MICH); Orcutt 5041, 1928 (US); along Mount Diablo, Parish of St. Ann, Imshaug, 3.3., 1952 (MSC); Parish of St. Catherine between Hollymount and Mount Diablo, ca. 2750 ft., Imshaug 14229, 14251, 1953 (MSC); Parish of St. Ann, Albion, ca. 2400 ft., Imshaug 15921, 15922, 1952 (MSC). DOMINICAN REPUBLIC: Mist forest ca. 5250 ft., ridge on road, ca. 50 km. from Constanza, Imshaug 23195, 1958 (MSC); MARTINIQUE: Route de la Trace from Deux Choux to Belvéder de Piton Gelé, Imshaug 32645A, 1963 (MSC). CURACAO: St. Christoffelberg, Curran and Haman, 3.3., 1917 (MICH). Central America. COSTA RICA: Cafion of Rio Reven- tazén, 900—1000 m., Dodge and Thomas 4616, 1929 (PH). 4. Ramalina sorediosa (B. de Lesd.) comb. nov. Ramalina daszpoga Tuck var. sorediosa B. de Lesd. Rev. Bryol. Lich. 7: 59. 1934. Lectotype (nov.): Boquerén, south Guantanamo Naval Station, Oriente, Cuba, 332. Hioram, §.n., Jan. 1933 (PH!). DescriEtion: Thallus sordid gray or stramineous, pendent, to 10 cm. long; branches terete, dichotomous through— OUt, to 1 mm. in cross section but thinner toward the apices; Surface nitid, granular soredia in small pockets in major branches; cortex thi fertile, usually abo brown, convex, pruin 40-6011, crowded wit apices; epithecium t cium hyaline, 10-15u taining 8 spores arr hyaline, uniseptate, 176 branches; cortex thick, cartilaginous. Apothecia rarely fertile, usually aborted, small, to 2 mm. across; disk brown, convex, pruinose; hymenium mucilaginous, colorless, 40—60u, crowded with slender paraphyses with capitate apices; epithecium thin, 4-6u thick, light brown; hypothe- cium hyaline, lO—lSu thick. Asci clavate, numerous, con— taining 8 spores arranged in one oblique series; spores hyaline, uniseptate, subfusiform, straight, rarely bent, 10-15 x 3—5u. Medullary reactions: Either PD+ yellow turning orange, K+ yellow-red, KC-, C-, or PD~, K+, KC+, C+ slowly becoming pink. Lichen substances: Atranorin, salazinic acid, seki- kaic acid, ramalinolic acid, usnic acid and substance C. Discussion: This species was originally described as a sorediate variety of Ramalina dasypoga, which has been considered here as synonymous with R. furcellata (Mont.) Zahlb. The type selected by the author to represent the variety name is a poor specimen scarcely sorediate. I have examined several other collections made by Hioram, Wright and Bro. Leon from the type locality which represent the more sorediate extreme of the population. The variety was raised to the rank of species because, unlike R. furcellata, it has soredia, lacks papillae and is rarely fertile. Also P‘, R. sorediosa is different chemically from P— R. furcel- lata. The PD— specimens are included within this species because except for t impossible to separa Ramalina ELI cal populations. On yellow and KOH+ red. salazinic acid in as norstictic acid in 1 norstictic acid is 1 [substance C) which of all the specimen: buried by atranorin The sample . in the medulla seki 177 because except for the chemical constituents they are impossible to separate from the PD- reacting ones. Ramalina sorediosa consists of two different chemi- cal populations. One portion of the population reacted PD+ yellow and KOH+ red. This group contained in the medulla salazinic acid in association with detectable traces of norstictic acid in thin-layer cromatography. Occasionally norstictic acid is replaced by an unidentified PD— substance (substance C) which fluoresces blue under UV. The thalli of all the specimens contained usnic acid frequently accom— panied by atranorin. The sample of the population reacting PD- contained in the medulla sekikaic acid and ramalinolic acid, while usnic acid in the cortex is rarely found in association with atranorin. Ecology: In Puerto Rico, Ramalina sorediosa is a common lichen in the lowland rainforest zone and in the valleys between the ”mogotes" within the seasonal evergreen forest zone. In few instances specimens were collected in the central mountains in the lower montane rainforest zone. The species is abundant in the second growth scrub of Chrysobalanus icaco not very far from the marine influence. On these shrubs the lichen forms dense masses of pendulous thalli that invade any available habitat. The PD- variant is the most abundant and constituted the bulk of the collections from this area. The PD+ type is less abundant and the other type that thalli without causi lichens commonly ass planata (protocetrar [salazinic acid vari The role of to that of R. anceps for bird nests. Distributior the PD— population i region, while the pi b91113 found in the l‘ 178 is less abundant and grows in such a close association with the other type that it is very difficult to separate the thalli without causing some damage to the specimens. Other lichens commonly associated with R. sorediosa are R. E22" planata (protocetraric acid variant) and Parmelia cristifera (salazinic acid variant). The role of the species in the ecosystem is similar to that of R. anceps in that it provides building material for bird nests. Distribution: According to my concept of the Species the PD- population is endemic to the Caribbean floristic region, while the PD+ population is more widely represented, being found in the West Indies, Central America and the Galapagos Islands. Thus, it could be said that the species has a neo—tropical distribution. In the West Indies both chemical variants of the species are present in Cuba and Puerto Rico, but only the PD+ variant has been collected in Jamaica. The species skips the rest of the West Indies islands and reappears again in Yucatan. The presence of the species in Yucatan is not surprising, since it is separated from Cuba by a narrow strait only 90 mi. wide. Likewise, for the same reason the species should be present in the rest of the islands. The species in Puerto Rico is restricted to the northern coastal plain, with occasional intrusions in the central mountain reg is shown on Figure 9 Material see Chrpsohalanus icaco west of Vega Baja, I 615,513,510, 69_6. 73’ 7M: w, 775 ”$715,716, 797, 3038111» 812, 818, 331%, 843A, 8441 M, M, 105113 1%, 1064 1065 1092 179 central mountain region. The distribution of R. sorediosa is shown on Figure 9. Material seen: West Indies. PUERTO RICO: In Chrysobalanus icaco scrub, south shore of Laguna Tortuguero, west of Vega Baja, Landr6n 618B, 659, 672, 676B, 678C, 684, “W“__——_— ggg, 688, 690, 696, 6983, 704, 705, 709, 713, 714, 722, 723, 7253, 728, 729, 7303, 731, 732, 734, 740, 742, 746, 7473, 749A, 751A, 753A, 755A, 758, 762, 763B, 764, 765, 767A, 768, ‘WW—__—_——_—_———_———— 771, 772A, 773A, 775, 776, 778, 779A, 790, 791, 792, 793, _—.___.,,,——.—_.___ 794, 795, 796, 797, 799, 800, 801, 803, 804, 805, 806, 8073, 809, 811, 812, 818, 820, 821, 822, 25, 831, 833A, 834, 836, 839, 840, 843A, 844A, 847, 855, 856, 857, 861, 869, 1049, —_—_W_________——_ 1050A, 1050B, 1051B, 1052, 1053, 1054, 1055, 1057, 1060, 1062B,1064,1065,1066,1068B,1069,1072,1073B,1075, ——W_—.—_—————_————_, 1076, 1079A,1081B,1082,1084,1086,1087A,1090C,1091B, h.—.__””””—___—————_———._ 1092, 1093C,1095,1096,1099,1967 (MSC); Vega Baja, 'shrubby on limbs,” Fink 2155, 1916 (MICH); Sardinera beach, back of Pterocarpus forest 5 km. west of Dorado, Landrén 918, — 1967 (MSC); Orocovis, Cerro Dona Juana, Bosque Toro Negro, in pasture with scattered Buchenavia capitata, Landrén 1807, 1967 (MSC); Barrio Padin, Corozal, route P.R. 568, Landrén 1806, 1967 (MSC); Lago Carite, in coffee plantation, Guayama Landron 1870A, 1967 (MSC); Monte Cuba, in guava scrub, Barranquitas, Landron 1945, 1967 (MSC); abandoned acerola plantation in Barrio Los Hoyos, 3 km. west of Vega Alta along route P.R. 690, Landrén 2181A, 2182, 2190, 2191, 2196 1968 (MSC); beach sc M m, 1968 (I field, west of Vega 22.351» 22g. 22_38. z 223 22m. 22h z Tortuguero Military Vega Baja, Landrén 2 Santiago de Cuba, B1 Slopes of San Juan, 1864 T ' rlglnal mat n w 5‘11 180 1968 (MSC); beach scrub, south of Hilton Hotel, Dorado, Landr6n 2208, 1968 (MSC); Laguna Tortuguero, old pasture field, west of Vega Baja, Landrén 2208, 2215, 2218, 2219, ———.——_—_———_———. _—__——_—_—______—————————__—_ —_———.——.——_.—————— Tortuguero Military Station, sandy forest at sea level; Vega Baja, Landrén 2300C, 2304, 1968 (MSC). CUBA: Oriente, Santiago de Cuba, Bro. Leon 12412, 1926 (PH); in cultivated slopes of San Juan, Loma del Gato, Imshaug 24900, 1959, Boca de Jaibo, Bro. Hioram 5487A, 1921 (US); Wright ER (EH); JAMAICA: Lumsden, Orcutt 3962B, 1927 (US); Parish St. Ann, Hopewell, ca. 1400 ft., Imshaug 15821, 1953 (MSC). Additional material seen: South America. ECUADOR: Near Wedeman Place, inland, Santa Maria I. (Charles or Floreana Id., Galapagos Ids.), Taylor 864, 865, 1934 (MICH). 5. Ramalina montagnei De Not. Ramalina montagnei De Not. Giorn. Bot. Ital. 2: 218. 1864. Original material: Cuba (non vidi). Ramalina subpellucida var. tuberculata Mull. Arg. Flora 71: 493. 1888. Original material: Coamo, Puerto Rico, Sintenis 68 2.3. (non vidi). Description: Thallus stramineous, rarely green, caespitose but sometimes pendulous, usually 3—5 cm. high, rarely to 10 cm. long, much branched, subdichotomous; branches 1 to 2 mm. always terete and at coarsely verrucose. Apothecia nu tan, convex or flat; hymenium 40-6011 thic thick. Asci clavate lapping sets of four W17 Straight but “Mm pink or K-, KC-, C‘ Lichen subs olic and usnic acid D' ' u m- late species With 1 by Montagne from Cu 181 branches 1 to 2 mm. broad, terete or subterete but apices always terete and attenuate; surface white papillate, or coarsely verrucose. Apothecia numerous, marginal, to 3 mm. broad; disk tan, convex or flat; hypothecium colorless, 10 to 15u thick; hymenium 40-60u thick; epithecium dense, colorless, 4-8u thick. Asci clavate, containing eight spores in two over— lapping sets of four; spores uniseptate, hyaline, fusiform, mostly straight but often some appearing bent, 16—22 x 3-5u. Medullary reactions: PD-, either K+, KC+, C+ turning pink or K—, KC-, C-. Lichen substances: Divaricatic, sekikaic, ramalin- olic and usnic acids. Discussion: Ramalina montagnei is a terete, papil— late species with long fusiform spores originally described by Montagne from Cuba. Plants are generally small but occasionally they can attain a large size for the species of about 10 cm. long without being pendulous. In the West Indies the species is represented by small shrubby specimens that are distinguished from the similar species R. subpellu- cida by their terete and papillate branches. The latter Species is not papillate and is rather compressed. Both sPecies have fusiform spores. Ramalina montagnei is chemically variable and represents two different populations. The specimens tested reacted homogeneously PD—, but one portion of them gave a pink color reaction did not produce any shown to contain sek medulla; the K- samp laryconsituent. F1 substance that could Miiller (1888 var. tuberculata as margins (lacineae ar albido-tuberculoso-e oritinal material, l Stevens from the sar t . 30061. However, ti rather than the fus Rundel (197 182 pink color reaction when treated with KOH, while the rest did not produce any color. The K+ population has been shown to contain sekikaic and ramalinolic acids in the medulla; the K— sample contained divaricatic acid as medul- lary consituent. Florida material contained an unidentified substance that could be stenosporic acid. Muller (1888) characterized Ramalina subpellucida var. tuberculata as being densely tuberculate along the margins (lacineae and lacinulae secus margines sat denso albido-tuberculoso—asperae”). Although I have not seen the original material, I have examined a collection by F. S. Stevens from the same locality and the material is sugges— tive of Ramalina montagnei. In Puerto Rico the species is not very common but small thalli of R. subasperata may be confused with R. 222' tagnei. However, the former species has ellipsoid spores rather than the fusiform spores typical of the latter. Rundel (1972) reported collecting R. montagnei from two hills in St. Thomas and identified four chemical strains 0f the species with the following chemical constituents: stenosporic, perlatolic, divaricatic acids and another with no medullary substance. Inasmuch as I have been able to study numerous specimens from the West Indies, I believe the fourth chemical strain reported by Rundel is the variant of 3. gracilis without medullary substance. This latter Species is characterized by two different variants, one smith psoromic acid both variants are h Thomas, Virgin Isla Britton and Marble mixture of both. T 45011. a1t., St. Th four variants of R. tolic and stenospor strate their occurr Florida mat its morphological a the species is ofte and North American fused with R. steno the latter species chemically heteroge either stenosporic four chemical strai stenOSporic, perlat stance. Chemical c from Florida demons were mixed collect: SDecimens of R. 3’3 Stenosporic acid, aPPearance , specim She called cfr. R. 183 with psoromic acid and another lacking medullary substances. Both variants are here reported from Puerto Rico and St. Thomas, Virgin Islands. In fact, one collection made by Britton and Marble in St. Thomas (n. 1356; MICH 1065) is a mixture of both. The locality for this collection is Crown, 450 m. alt., St. Thomas, the same given by Rundel for the four variants of R. montagnei. As to the presence of perla— tolic and stenosporic acids, I have been unable to demon- strate their occurrence in the West Indies. Florida material of the species is more variable in its morphological and chemical characters. In this region the species is often confused with a complex of European and North American species. Commonly R. montagnei is con- fused with R. stenospora Mull. Arg.; however, specimens of the latter species with branches that are in part terete are chemically heterogeneous and without exception produce either stenosporic or perlatolic acid. Moore (1968) reported four chemical strains containing one of four substances: stenosporic, perlatolic, sekikaic or an undetermined sub- stance. Chemical determination of some of her material from Florida demonstrated that packets marked R. montagnei were mixed collections. Under this species were included specimens of R. stenospora containing either perlatolic or stenOSporic acid, and thalli of a somewhat semi—fistulous appearance, specimens containing divaricatic acid of what She called cfr. R. fastigiata. This latter species is distinguished from said spores and lar studied from Flori: acids as medullary Florida indicated 1 which could be take compressed branches terete and papillat Howe (1914) Ramalina mont agnei Species by its late reaction. Moore (1 stated that probabl complex of chemical could be either K+ h. stenospora is al In my opini either sekikaic or W, while t1 Papillate or not ar aCid constitute R. Florida specimens, instead and produc< hilt the Puerto Ric Ecology: VOuntainous region 184 distinguished from R. montagnei by its rather large ellip- soid spores and lack of papillae. Additional material studied from Florida demonstrated sekikaic and ramalinolic acids as medullary substances. Other collections from Florida indicated the presence of R. subpellucida, a species which could be taken for R. montagnei were it not for its compressed branches and lack of papillae rather than being terete and papillate. Howe (1914) thought that R. stenospora merged into Ramalina montagnei and that it differed from this latter species by its lateral apothecia and questionable K+ reaction. Moore (1968) disagreed with Howe's concept and stated that probably the Florida material represented a complex of chemical species. In fact Ramalina montagnei could be either K+ or K- in its medullary reaction, while R. stenospora is always K-. In my opinion the terete papillate plants containing either sekikaic or divaricatic acid belong to Ramalina montagnei, while those plants that are compressed whether papillate or not and containing stenosporic or perlatolic acid constitute R. stenospora. I consider that some of the Florida specimens, which are not papillate but striate instead and produce divaricatic acid are not R. stenospora but the Puerto Rican species R. subpellucida Mull Arg. Ecology: Ramalina montagnei is present in the mountainous regions of the West Indies and in the everglades ",v 5"." '5 and hammocks of F14 from dry areas in 1 United States. Rer late the chemical 5 selection on Signal in St. Thomas, Virg Species represented succession on branc the second stage we However, he could r lation between chem in the West Indies. from material C0116 Additional material shows a southeasten 181 (MSC); Merr. n< West Indie: HAITI: Vicinity o Slope, coast east 39.0.31: 1929 (US). [be. DOMINICAN R tos Amaceyes, Cord 1958 (MSC). North Ame: 185 and hammocks of Florida. The species has been recorded from dry areas in Puerto Rico, St. Thomas and southern United States. Recently Rundel (1972) attempted to corre- late the chemical strains of the species with habitat selection on Signal Hill (455 m.) and Crown Mountain (475 m.) in St. Thomas, Virgin Islands, U.S.A. He believed the species represented the earliest stage of macrolichen succession on branches of trees. According to his study the second stage was dominated by Ramalina denticulata. However, he could not find any significant degree of corre— lation between chemical strains and habitat. Distribution: This species was described from Cuba in the West Indies. It has been identified in this study from material collected in Puerto Rico, Cuba and Hispaniola. Additional material examined from Florida and Louisiana shows a southeastern United States distribution as well. Material seen: Exsiccati examined. Cum. I. no. 181 (MSC); Merr. no. 268 (MICH). West Indies. PUERTO RICO: Stevens 3.3. (MICH). HAITI: Vicinity of Jean Rabel, on shrubs, steep rocky slope, coast east of Bord de la Mer, Leonard and Leonard 12903B, 1929 (US)., CUBA: On twigs of tree, Wright 3.3. (PH). DOMINICAN REPUBLIC: Cloud forest, 3000-3200 ft., Los Amaceyes, Cordillera Septentrional, Imshaug 23297, 1958 (MSC). North America. FLORIDA: Lee C0,, Road to Sanibel .' fn'f A?" vr-t— s: - I, u _.-_—- ... Is., trunks of royr 00., trunk of royal 1966 (DUKE); St. L1 M 33%, 1966 (1 State Park , Harris River State Park, c RR, £0, 1967 (h 131, m, 7-28-31 ( Eckfeldt, 5.11. (MIC Daytona, Thaxter l_f hp, 1898 (PH) . 8991512221. 19 (MICH). LOUIS I; data in three more 6. Ramalina cochle Ramalina g 5: 542. 1905. Oriy Ramalina 13 ll: 52. 1908. Ori 1904 (ent). Ramalina l_1_ 70,2): 59. 1934. ‘32- Hioram, s .n. Descriptir high, sparsely brz 186 Is., trunks of royal palm, Moore 3690, 1966 (DUKE); Dade Co., trunk of royal palm in Yard in Goulds, Moore 3691, 1966 (DUKE); St. Lucie Co., high hammock near Indian River, Moore 3694, 1966 (DUKE); Highlands Co., Highland Hammock State Park, Harris 2550A, 1967 (MSC); Sarasota Co., Myakka River State Park, oak woods scrub along Myakka River, Harris £231» RRRQ, 1967 (MSC); Lakeland, on red maple, MacFarlin ERR, ERR, 7-28-31 (MICH); Turtle Mt., Rglly Rig, ARR; (MICH), Eckfeldt, 3.2. (MICH); McIntosh, Thaxter Rig, 1897 (EH); Daytona, Thaxter l 7, 1898 (PH); Palm Beach, Thaxter 261, 1 8, 1898 (FH). __ Additional material seen: TEXAS: Dallas, Davis 13, l9 (MICH). LOUISIANA: St. Martinsville, Langlois 801; no data in three more collections by Langlois, 1888 (MICH). 6. Ramalina cochlearis Zahlbr. Ramalina cochlearis Zahlbr. Bull. Herb. Boissier II. 5: 542. 1905. Original material: Brazil (non vidi). Ramalina inflata var. soredians Merr. Bry010gist ll: 52. 1908. Original material: Jamaica, Cummings E‘E-: 1904 (PH!). Ramalina hiorami B. de Lesd. Rev. Bryol. Lich. 7(1,2): 59. 1934. Original material: Loma San Juan, Cuba, E33. Hioram, 3.2. (non vidi). Description: Thallus caeSpitose, rigid, to 4 cm. high, sparsely branched; branches linear, dichotomous, terete or subterete tips of branches in soralia; surface ni visible, minutely p tol.5 mm. at the t pminose, concave; physes slender, sim clavate, containing series, spores uni 12-16 x 5-7u. Medullar red or pink, or KC- Lichen subs linolic acid, an u G.E. (substance B) Discussion: characterized by th of the main branche any other occurring based his Ramal ina __.._——————- blerr. on a mixed cc which included R 1er'=1\ris.lahlbr. Hi the sorediose thal All specim ““216 Specimen wh 187 terete or subterete, rarely compressed, to 2 mm. broad, tips of branches inflated and opening into helmet-like soralia; surface nitid; cortex thin, cartilaginous strands visible, minutely papillose. Apothecia rare, very small, to 1.5 mm. at the tip of short thin pedicels; disk tan, pruinose, concave; hymenium colorless, 30-59u thick, para- physes slender, simple or branched, non-septate. Asci clavate, containing 8 spores arranged in two irregular series, spores uniseptate, hyaline, ellipsoid, mostly bent, 12-16 x 5-7u. Medullary reactions: PD-, either K+ red or K-, KC+ red or pink, or KC—, C+ pink or C-. Lichen substances: Atranorin, sekikaic acid, rama— linolic acid, an undetermined substance crystallizing in G.E. (substance B) and usnic acid. Discussion: Ramalina cochlearis is very well characterized by the typically swollen soralia at the tips of the main branches. The species cannot be confused with any other occurring in the West Indies. Merrill (1908) based his Ramalina inflata Hook. f. and Tayl. var. soredians Merr. on a mixed collection made by Cummings in Jamaica, which included R. inflata Hook. f. and Tayl. and R. coch- learis Zahlbr. His description of the variety referred to the sorediose thallus of R. cochlearis. All specimens examined were sterile except for a Single specimen which had a few small apothecia. I have not seen. any other soralia except the sometimes globose turned inside out laciniae in R. coc are not typically According to my in Lesdain (1934) des Cuba under the new Although R PD, it is composed of these gave a wi KC, and contained occasionally atran reacted K-, C- and unidentified chemi B). When a sam two unidentified s tained. These spo With benzidine. T thallus with G .E. that aggregated i Ecology: huntislands of t1Imee’cween 350 188 not seen any other species which show these helmet—like soralia except the European R. baltica Lett. Soralia sometimes globose with opening on the side, sometimes turned inside out with coarse granules exposed. When the laciniae in R. cochlearis are compressed and the soralia are not typically helmet-like, it resembles R. farinacea. According to my interpretation of the species, Bouly de Lesdain (1934) described the species R. cochlearis from Cuba under the new name R. hiorami. Although Ramalina cochlearis reacted negative to PD, it is composed of two different chemical variants. One of these gave a wine red color when treated with K, C, or KC, and contained sekikaic, ramalinolic and usnic acids and occasionally atranorin. The other part of the population reacted K-, C- and KC+ red. This variant contained an unidentified chemical constituent in the medulla (substance B). When a sample of the thallus is chromatOgraphed, two unidentified spots that fluoresce blue under UV are ob- tained. These spots take a yellowish color when sprayed With benzidine. Treatment of the acetone extract of the thallus with G.E. precipitated short attenuate crystals that aggregated in rosettes resembling ramalinolic acid. Ecology: Ramalina cochlearis occurs in the West Indian islands of Jamaica, Hispaniola, and Cuba at eleva— tions between 3500 and 6000 feet. It is found in the montane sclerophyll the Blue Mountains of Haiti. Distributio Indian distribution has not been report Material se ca. 4000 ft. , Blue (MSC); Morces Gap, 1952 (MSC); Silver (MSC); forest regi near Hardwar Gap, montane sclerophyl Parish of St. Andr Plitt s.n., July ] x...— July 9, 1919, P_1_i_t_3 Pine forest with St of Forét des Pins, 23086, 1958 (MSC); slope of ridge abo* Imshaug 22494, 195 7~ Ramalina graci Physcia 209. 1826. Ramali 1360- Original or 189 montane sclerophyll and montane rainforest in the region of the Blue Mountains in Jamaica and in the dry pine forests of Haiti. Distribution: This species has a restricted West Indian distribution, typical of endemic plants. The species has not been reported from Puerto Rico or the Lesser Antilles. Material seen: West Indies. JAMAICA: Farm Hill, ca. 4000 ft., Blue Mountains, Imshaug 14776, 14777, 1953 (MSC); Morces Gap, 4800 ft., Blue Mountains, Imshaug 13274, 1952 (MSC); Silver Gap, Blue Mountains, Imshaug 14089, 1953 (MSC); forest region,ca. 3800 ft. along Moodie‘s Gap Trail near Hardwar Gap, Blue Mountains, Imshaug 13076, 1952 (MSC); montane sclerophyll region near Clifton Mount, 4000 ft., Parish of St. Andrew, Imshaug 13599, 1952 (MSC); Cinchona, Plitt 3.3., July 10, 1919 (US); Bellevue, Plitt 3.3., July 9, 1919, Plitt 3.3., July 21, 1926 (US). HAITI: Pine forest with scattered agave, 5300 ft., 10w hills west of Forét des Pins, Department de l'Ouest, Imshaug 23085, 23086, 1958 (MSC); cultivated and pasture area on north slope of ridge above Kenscoff, 5000 ft., Départment de l'Ouest, Imshaug 22494, 1958 (MSC). CUBA: Wright 3.3. (FH). 7. Ramalina gracilis (Pers.) Nyl. Physcia gracilis Pers. in Gaud. Voy. Uranie Bot. 209. 1826. Ramalina gracilis (Pers.) Nyl. Syn. Lich. 1: 296. 1860. Original material: Brazil, Madagascar and China (non vidi). imam—a an Journ. Bot. 34: 31. St. Vincent, Elliot‘ Description much branched, to 5 0.41.5 mm. broad, striate and often b ally to the tips of to the branches ; h)’ spores uniseptate } W914. 190 Ramalina gracilis subsp. antillarum (Nyl.) Vain. Journ. Bot. 34: 31. 1896. Lectotype (nov): Bow—wood Estate, St. Vincent, Elliott 333 1891—1892 (BMl). Description: Thallus stramineous, pale, caespitose, much branched, to 5 cm. high, branching divaricate; branches 0.4-1.5 mm. broad, terete, attenuate, tips filiform, bases striate and often black, the dark color extending occasion— ally to the tips of minor branches. Apothecia common, small to 2 mm. broad, well distributed throughout the thallus, those subterminally located assume a subgeniculate position; disk tan, pruinose, convex and closely adpressed to the branches; hypothecium colorless, 20-30u thick; hymenium gelatinous, 60—80u thick, paraphyses simple; epithecium thin, never over lOu, pale brown; asci clavate containing 8 spores arranged in two irregular series; spores uniseptate hyaline, broad ellipsoid, straight, 12—15 X 6-9u. Medullary reactions: PD+ yellow-green or PD—, K—, KC—, C—. Lichen substances: Psoromic and usnic acids and occasionally atranorin. Discussion: No original material of Ramalina gracilis has been examined and the species has been inter— preted on the basis of published literature. The material studied fitted the original description of both R. gracilis and subs. antillarum, and the asci matched very closely material. the diagram illustrz (Nylander, 1858-1861 The species the West Indian spew preted the species Species has a diffe w has broad whereas R. attenuat \ zinic acid. The origina indication as to th Vainio ( 191 the diagram illustrated by Nylander from the type specimen. (Nylander, 1858—1860, Plate 8, Figures 25a 8 b). The species cannot be confused with any other of the West Indian species in the genus. Howe (1914) inter— preted the species as synonymous with R. attenuata but the species has a different spore size and chemistry. Ramalina gracilis has broad spores and contains psoromic acid, whereas R. attenuata has narrow spores and contains sala— zinic acid. The original description of R. gracilis gives no indication as to the chemical reactions of the original material. Vainio (1890) identified R. gracilis from Rio de Janeiro as being K—, and later (Vainio, 1896) described R. gracilis subsp. antillarum from St. Vincent also as being K—. He distinguished the subspecies from the species by its more pruinose apothecia and from R. gracilenta by its terete branches instead of flat. I have examined a fragment of the type material of R. gracilis subsp. antil- larum (Elliott 16b in BM) and confirmed it is K— and con- tained no medullary substance. I found the West Indies material to be K-, KC—, C- and either PD— or PD+. The great majority of the specimens produced a bright yellow—green color when PD is applied to the medulla. These specimens contained the substance psoro- mic acid and occasionally atranorin. Usnic acid is a Constant substance in the whole population and it is the only chemical const In the light of the conclude that R. gr arated from R. grac M: '1 lower montane rainl epiphytic on brand m. It is 1% (Sw.) Ach. the Sierra de Cayej i' W and l Parmelia, Heterode microflora. Specimens A. t 01 trees. The spe the thallus and ti is the result of 1 instances the plar field observations numerous branches gluing the plant 1 ( the localitieS Br; bllamder (18 70) a‘ .t from St, Vince: 192 only chemical constituent present in the PD- population. In the light of the foregoing considerations I can only conclude that R. gracilis subsp. antillarum cannot be sep- arated from R. gracilis. Ecology: This Species is commonly found in the lower montane rainforest zone of the island where it is epiphytic on branches of Tabebuia heterophylla and Rapanea ferruginea. It is usually associated with Ramalina 333— planata (Sw.) Ach. On the wet slopes of the eastern end of the Sierra de Cayey it grows together with R. peranceps, R. dendroides and R. subasperata among the fruticose lichens. Parmelia, Heterodermia and crusts compose the rest of the microflora. Specimens without medullary substance were collected on the warm and moist northern coastal plain, on the bark of trees. The species often appears black at the base of the thallus and tips of branches, something which I think is the result of injuries from insect bites. In other instances the plant is more bushy than normal and again field observations showed that around the eaten pieces numerous branches were in the process of regeneration, giving the plant a flimsy appearance. Distribution: The original description mentioned the localities Brazil, Madagascar and China, and later Nylander (1870) added Australia. Vainio (1896) described it from St. Vincent in the Lesser Antilles under the name Ramalina raciliS 5 apan-tropical (1151 In the Wes Rico and has been I Lucia and J amaica. Rico and the Virgi‘ in St. Lucia and 0 The similarity bet populations is in and floristic affi The chemic suggest that in th sified in Puerto F islands or the spe les diversified ar nearer Greater Ant 193 Ramalina gracilis subsp. antillarum. These reports indicate a pan—tropical distribution. In the West Indies the species is common in Puerto Rico and has been collected in the Virgin Islands, St. Lucia and Jamaica. Both chemical variants occur in Puerto Rico and the Virgin Islands, but only the PD+ variant occurs in St. Lucia and only the PD- variant occurs in Jamaica. The similarity between the Puerto Rico and Virgin Islands populations is in accordance with their geographical proximity and floristic affinities. The chemical and distributional data available suggest that in the West Indies either the species has diver- sified in Puerto Rico and from here migrated to the other islands or the species once introduced into the Lesser Antil- les diversified and has been swept by air currents into the nearer Greater Antilles. Although the Caribbean trade winds Would be sufficient to account for the latter alternative, it remains to explain the absence of the species from most of the Lesser Antilles, Cuba and Hispaniola. In the island of Puerto Rico the PD+ population of the species is confined to the central region and western end of the Central Cordillera so that it has a central— highland distribution. The PD- population is very sparse and the few collections I have studied are mainly from the northern coastal zone (Figure 10). Material se mear river 550 m-, M» 1.273. 1% Villalba, Landrén l ‘— _ near Orocovis and ( 1833, 1846A, 18481 Landrén 1858A, 1861 barranquit as , Land: ——_ 1m, 1967 (MSC); 19%. 1986C, 1987 19% 1997A, 1999 194 Material seen: PUERTO RICO: Bosque de Guavate, near river 550 m., Cayey, Landr6n 1229, 1230B, 1231A, 1236A, 1237A, 12403, 1244, 1245, 1257, 1260A, 12683, 1271, 1273, ——-—————————————_.—__.————.——u— 1274A, 12753, 1290C, 1292C, 1293A, 1300A, 1302, 13033, 13043, 1308, 13193, 1329C, 1341C, 1348, 13493, 1350, 1354, 1358A, 1360B, 1373, 1378, 1379, 1380A, 1388C, 1389C, 1391, 1392, 1393A, 1397, 1443B; Barrio Caonillas, Toro Negro, Villalba, Landrén 1715, 1716C, 1967 (MSC); Barrio Padin, near Orocovis and Corozal, Landr6n 1799, 1805A, 1814, 1830B, 1839D, 1846A, 1848D, 1859C, 1967 (MSC); Carite Lake, Guayama, Landrén 1858A, 1863A, 1866B, 1802, 1967 (MSC); Monte Cuba, _.___.____——.__-——————__ 1938C, 1967 (MSC); Aibonito, Landrén 1976A, 1977C, 1978A, 1982C, 1986C, 1987D, 1988, 1989A, 1993E, 1994B, 1995B, 1996B, 1997A, 1999B, 2006A, 1967 (MSC); north of Barran— quitas, ca. 500 m., Landrén 2386, 1968 (MSC); Tabonuco, Monte del Estado, Maricao, Landrén 2426A, 2433B, 2436A, 2437A, 1968 (MSC); Quebradillas, Stevens 5012, 1913 (MICH); Naranjito, Fink, 236A (MICH); Aibonito, Fink 1936, 1915 (MICH); Punta Cangrejos, Johnston and Stevenson 1972A, 1914 (US); Santurce, Stevenson 2949, 1915 (US). VIRGIN ISLANDS: St. Thomas, Britton and Marble 3333, 1913 (MICH). ST. LUCIA: cultivated area, Fond Doux, 1200-1663 ft., north— east of Gros Piton, Quartier of Soufriere, Imshaug 3333, 1963 (MSC); lower montane rainforest, Mt. La Combe, 1000— 1442 ft., Barre de l'Isle Ridge, Quartier of Castries and 4“ bennery, Imshaug fl 1809 ft., Hanover, Plitt, s.n., 1932 l b. Ramalina subasp Ramalina 3p Lectotype (nov.): 1 Descriptior “11990115, profusely apices always tere‘ Papillae small, pu diSpersed: cortex 195 Dennery, Imshaug 30056, 1963 (MSC). JAMAICA: Birchs Hill, 1809 ft., Hanover, Imshaug 15718, 1953 (MSC); Chestervale, Plitt, lUl .3., 1932 (US). 8. Ramalina subasperata Nyl. Ramalina subasperata Nyl. Flora 59: 411. 1876. Lectotype (nov.): Cuba, Wright 239 (FH!). Description: Thallus small, to 4 cm. high, stra— mineous, profusely branched, branches subterete to terete, apices always terete and attenuate; surface papillate, papillae small, punctiform, with white summits, and widely dispersed: cortex thick. Apothecia numerous, to 6 mm. across, subterminal, subgeniculate, concave; disk tan, pruinose; hypothecium 12-18u thick, hyaline; hymenium colorless, 30—50u thick, epithecium tan colored 5—10 thick. Asci clavate containing 8 spores arranged in a single row; spores uniseptate, hyaline, mostly bent, rarely straight, 9-10 x 3.5-6u. Pycnidia black, mostly at the tips of branches. Medullary reactions: KC-, C—, PD+ yellow—orange, K+ yellow-red or PD—, K-. Lichen substances: Atranorin, salazinic acid, protocetraric acid, divaricatic acid, sekikaic acid, rama— linolic acid and usnic acid. Discussion: This species could be mistaken in the field for R. denticulata Nyl., from which it is distinguished by its chemistry an It resembles R. ”l1 These latter two 51: em Florida, but 116 Indies. Ramalinarp cetraric or salazir teristic habit of 1 separates it from I guished by its C+ ; teristic of CI‘yptOr are the small size in contrast to the 4-811), It is evid the cortex, gTOWth chemical diversity complanata stock. 03 and 13. willey \ The origin 196 by its chemistry and small, terete or subterete thallus. It resembles R. willeyi R. H. Howe and R. paludosa Moore. These latter two species are common to the swamps of south- ern Florida, but neither has been reported from the West Indies. Ramalina willeyi is known to contain either proto- cetraric or salazinic acid (Moore, 1968) but its charac— teristic habit of forming spherical, rosette—like tufts separates it from R. subasperata. R. paludosa is distin- guished by its C+ red wine color medullary reaction, charac— teristic of cryptochlorophaeic acid. Additional differences are the small size spores (8-12 x 3-6u) of the R. paludosa in contrast to the larger ones of R. subasperata (14-18 x 4—8u). It is evident that the spore size, thickness of the cortex, growth habit, presence of papillae and its chemical diversity relate R. subasperata to the Ramalina complanata stock. The same observation is true for R. 3333— dosa and R. willeyi. The original description of the species indicates that the species reacted K-; however, since there are two pieces of thalli in the original packet and one is K- (containing divaricatic acid) and the other is K+ (contain— ing salazinic acid) it is obvious that Nylander's descrip— tion refers only to one of them and that probably he did not test the second specimen. Therefore, the numbers 25C and 25D correspond to two different chemical variants of the species which i thallus labelled 2! lectotype. I agree wi' represent R. s_ub_a3 more have been add that the species 5 chemical variants. Individual 197 the species which is based on the description of the K— thallus labelled 25C. Thus, I am designating 25C as the lectotype. I agree with Nylander in that both chemical variants represent R. subasperata. To these two chemical types two more have been added during the course of this study so that the species should be interpreted as having four chemical variants. Individuals of the Puerto Rican population of Rama- lina subasperata contained up to four major constituents: salazinic, protocetraric, sekikaic and ramalinolic acids. The island population falls under the sekikaic acid contain— ing variant. In 61 out of the 71 collections I made in Puerto Rico, every single specimen collected contained seki— kaic, ramalinolic, salazinic and usnic acid. Only two out of the total indicated detectable concentration of the lichen substance atranorin. In the rest of the specimens (10 collections) protocetraric acid is associated with seki- kaic, ramalinolic and usnic acid. In this group, atranorin was also recognized in two specimens. Of 71 specimens collected in the island of Puerto Rico none exhibited the same chemistry as the type material. Some of the thalli reacted K+ and some K—, but all gave a red or yellow color reaction with PD. Subsequent crystallo- graphic analysis confirmed that the K- sample contained protocetraric acid and the K+ portion of the sample had 1“! salazinic acid. F that two other PD- and protocetraric unknown spots was G.E. solvent, in w ramalinolic acids combinations of PE or protocetraric a [sekikaic and ram yet been reported substances were ec DTeCipitants obtai ch onatography [\n analys of R- ub 198 salazinic acid. Further chromotagraphic studies revealed that two other PD- spots were associated with both salazinic and protocetraric acids. Identification of these two unknown spots was done by treating the acetone extract with G.E. solvent, in which typical crystals of sekikaic and ramalinolic acids precipitated. The appearance of these combinations of PD+ B- orcinol type depsidones (salazinic or protocetraric acids) with PD- orcinol type meta—depsides (sekikaic and ramalinolic acids) in the same thallus has not yet been reported in Ramalina. Both the PD+ and the PD- substances were equally abundant judging by the amount of precipitants obtained by micro-chemical tests. Specimens tested from Haiti contained either sala- zinic acid or protocetraric acid alone, but material from Cuba was found to produce either salazinic or protocetraric acids in combination with sekikaic and ramalinolic acids, accompanied by usnic acid and often the accessory substance atranorin. In a high percentage of specimens containing salazinic acid, traces of norstictic acid were detected. This accessory substance was only demonstrated by thin-layer chromatography. An analysis of the total West Indian collections of B. subasperata indicated that salazinic and protocetraric acids can either occur alone or in combination with sekikaic and ramalinolic acids, accompanied by sporadic secondary Substances. Divaricatic acid always occurs alone and it is only known from the zinic, protocetrari together in any cor In accordai the species 3. s_ub_z constituted by foui Chemical variai Chemical varia: Chemical varia Chemical varia It may be be included partly the sekikaic acid 199 only known from the type material. Three substances (sala- zinic, protocetraric and divaricatic acids) never occurred together in any combination of two or more. In accordance with frequent practice in lichenology the species 3. subasperata has been interpreted as being constituted by four chemical variants: Chemical variant I (divaricatic acid or typical) Chemical variant II (salazinic acid) Chemical variant III (protocetraric acid) Chemical variant IV (sekikaic acid) It may be argued that the chemical variant IV should be included partly in II and partly in III. In considering the sekikaic acid containing population a variant by itself, much weight was given to the presence of the common substance. Ecology: This inconspicuous species is typical of the lower montane rainforest in Puerto Rico, especially the wet and cool habitats of the central eastern mountains. In the Carite mountains the species shows the habit of growing mixed with Ramalina complanata (salazinic and protocetraric acid strains), 3. gracilis, B. dendroides and 3. peranceps. Typically, it occupies the exposed sites at the base of the trunks of Tabebuia heterophylla. Toward the central part of the island, near Barran- QUito and Aibonito, at elevations between 400 and 660 meters, abandoned fields are frequently invaded by the tenacious scrub Psidium guajava and Rapanea ferruginea. This scrub vegetation constit M, especial acid, which could together with 3. S and I} gracilis. the hymenial layer PTeViously known 0 from Cuba, Jamaica Species is absent Perhaps this small collectors and £01 from the Lesser A: In Puerto species parallels 200 vegetation constitutes a frequent habitat of Ramalina sub— asperata, especially those specimens containing salazinic acid, which could be collected in a crowded condition together with 3. complanata, 3. dendriscoides, 3. peruviana and 3. gracilis. Numerous specimens were observed to have the hymenial layer eaten away, presumably by insects. Distribution: Ramalina subasperata has been previously known only from Cuba but it is here reported from Cuba, Jamaica, Haiti and Puerto Rico (Table 10). The species is absent from the Lesser Antilles and Florida. Perhaps this small species may have been overlooked by many collectors and for this reason it has not been recorded from the Lesser Antilles. In Puerto Rico the chemical distribution of this species parallels that of 3. complanata. Within the island the species population containing salazinic acid is much more abundant than the protocetraric acid producing type. The first mentioned variant is frequent in the eastern and central part of the island, while the latter is mostly con- fined to the eastern part. Only one specimen was collected in the northern coastal plain of the island, and two in the western mountains. Most early collections of the species were done in the Cordillera Central toward the central part of its range. Figures ll—13 show the Sierra de Cayey— Maricao distribution pattern of the Species. mn1e10. Distrib B. suba __—.————- Chemical Variant luDivaricatic acid IInSalazinic acid HIuProtocetraric acid IVnSekikaic acid \— Table 10 : h nhasperata ant \ hemer Antilles. Cmalmrbored chm main variants Rico variant 1v, 201 Table 10. Distribution of the chemical variants of B. subasperata in the Greater Antilles. Chemical Hispaniola Puerto Variant Cuba Jamaica (Haiti) Rico I-~Divaricatic acid + II--Salazinic acid + + + III--Protocetraric acid + IV—-Sekikaic acid + + Table 10 shows that the distribution of the species 3. subasperata and its chemical variants is limited to the Greater Antilles. The specimens examined indicated that Cuba harbored chemical variants I, II and IV, Hispaniola (Haiti) variants II and III, Jamaica variant I and Puerto Rico variant IV. The range of the species appears to have its center of dispersion in Cuba, which contained the highest number of variants. Toward the eastern part of its range the species showed a decreasing dispersion gradient. At the extreme eastern end lies Puerto Rico with only one variant present while none has been reported from the Lesser Antilles. Material seen: West Indies. PUERTO RICO: Vega Baja, Laguna Tortuguero, Landrén 933’ 1967 (MSC); Bosque Estatal de Guavate, near river, Cayey, Landrén 12313, 1242A, 1265]). £62 13290, 1337, 1360C _— _— 1396, 1967 (MSC); t g, 1967 (MSC); : vation Tower, Land Maricao west of Ob Bosque Toro Negro , Villalba, Landrdn Landron 1730B, 174 Orocovis, Landrén \ Laudron 1781C, 196 1%, 1804C, 1805 team, Landrén l8l \ 202 1242A, 126SD, 1269A, 1275A, 1278A, 1278B, 1292A, 130013, __~—_—__———_—_—__________~______——_.—_ 1329D, 1337, 1360C, 1361B, 1366C, 1366A, 1368B, 1375, 13938, .___________—___————____—-—_~___———_ 1396, 1967 (MSC); Guavate near Park, Cayey, Landr6n 1408B, 1429, 1967 (MSC); Monte del Estado, Maricao, east of Obser- vation Tower, Landrén 1571C, 1967 (MSC); Monte del Estado, Maricao west of Observation Tower, Landrén 1589C, 1967 (MSC); Bosque Toro Negro, Landr6n 1684, 1692F, 1967 (MSC); Caonillas, Villalba, Landrén l7l7E, 1967 (MSC); Divisoria, Toro Negro, Landrén 1730B, 1740A, 1743B, 1967 (MSC); Barrio Cuchillas, Orocovis, Landrén 1776, 1967 (MSC); Barrio Negro, Orocovis, Landron 1781C, 1967 (MSC); Barrio Padin, Orocovis, Landron 1799D, 1804C, 1805C, 1967 (MSC); Cerro Dona Juana, Toro Negro, Landrén 1817B, 1822D, 1833B, 1849B, 1967 (MSC); Lago Carite, Guayama, Landron 1858B, 1865A, 1881E, 1878C, 1882D, 1889C, 1891F,1893C,1894E,1897A,1911B,l917B,1923D, “hm—_——_———_———— 1937D, 1940B, 1942B, 1967 (MSC); Monte Llano, Cidra, Lan-w dron 1956D,1967 (MSC); Aibonito, Landrén 1977A, l977E, 1958B, 1986B, 1993A, 1995C, 1996E, 1997D, 1999B, 2000A, 1967 (MSC); “QM—fi—fi— Cerro Punta, Adjuntas, Landr6n 2390, 1968 (MSC); Monte del Estado, Maricao, Landrén Z433C, 1968 (MSC); Guajataca lake, Landrén 2462B, 1968 (MSC). CUBA: Wright 25D (PH); Loma del Gato, Bro. Hioram 6695, 1923 (US); rock in cultivated area at Armenia on slope of El Gato, Loma del Gato, Sierra Maestra, Oriente, Imshaug 24782, 1959 (MSC). HAITI: Dépt. l'Ouest, summit of Téte Btang, on ridge between Kenscoff and Furcy, ca. 6000 ft., Imshaug 22562, 1958 (MSC); Dépt. du Nord, below the Wetmore L851, 195 8 (FH). Series Cc 9. Ramalina comp] \ Lichen con \— u: 290. 1797. P5 259. 1803. Ramali 1810. Original 1112 Pa ' w 5 221. 1883. Ramalf \ midi), 203 du Nord, below the Citadelle, south of Milot, ca. 2300 ft., Wetmore 2851, 1958 (MSC). JAMAICA: Cummings s.n., 1904 (PH). Series Compressiusculae (Vain.) Zahlbr. 9. Ramalina complanata (Sw.) Ach. Lichen complanatus Sw. in Ach. Kgl. Vet. Nya. Handl. 18: 290. 1797. Parmelia complanata (Sw.) Ach. Meth. Lich. 259. 1803. Ramalina complanata (Sw.) Ach. Lich. Univ. 599. 1810. Original material: Indiae Occidentalis, Swartz (8!). Parmelia denticulata Eschw. in Mart. Fl. Bras. 1: 221. 1883. Ramalina denticulata (Eschw.) Nyl. Act. Soc. Sci. Penn. 7: 434. 1863. Original material: Amazonas (non vidi). Description: Thallus caespitose, stramineous, occasionally green, to 7 cm. long, but more often smaller; branches dichotomous, compressed becoming canaliculate, channels shallow, disappearing toward the tips; surface nitid papillate, papillae in some cases appear double lobu- lated or punctiform; cortex thick, cartilaginous to soft in old specimens, thin in young lichens. Apothecia common, marginal, rarely laminal, adnate, variable in size in accor— dance with the size of the lichens; disk concave, rarely convex, tan colored, pruinose; margin rugose. Asci con- taining 8 spores; spores uniseptate, ellipsoid or ellipsoid— elongate, curved, or bent, variable, 9-16 x 4-9n. Medulla yellow-red or PD- ’ wine or KC-, eithe Lichen sub zinic acid, protoc acid, ramalinolic and unidentified s Discussior \ distinguished by P species Ii. complaI reactions; R. com; produced a K+ real part of the origir tained divaricatit Specimens studied cal character the two species. Th thallus ( e Moore, 1 correlate well wi 204 Medullary reactions: Either PD+ yellow—orange or yellow-red or PD—, either K+ yellow or K—, either KC+ red wine or KC—, either C+ transient red or C-. Lichen substances: Atranorin, substance H, sala- zinic acid, protocetraric acid, divaricatic acid, sekikaic acid, ramalinolic acid, cryptochlorophaeic acid, usnic acid, and unidentified substance A. Discussion: The species Ramalina denticulata was distinguished by Nylander from morphologically similar species 3. complanata (Sw.) Ach. on the basis of chemical reactions; 3. complanata reacted K— while 3. denticulata produced a K+ reaction. I have examined what appears to be part of the original material of B. complanata and it con- tained divaricatic acid and usnic acid. In the numerous specimens studied, I could not find any constant morphologi— cal character that could be safely used to recognize the two species. The smaller spores and the more branched thallus (Moore, 1968) attributed to 3. denticulata do not correlate well with the chemical variation. The validity of such chemical differentiation is even more doubtful when it has been demonstrated that a K+ 0r K- reaction can represent a whole series of medullary substances. Young plants of 3. complanata resemble adult plants 0f 3. subasperata. They can be distinguished in the labor— atory by the combined use of thin—layer chromatography and crystallograPhY- and 3. W Thalli of their chemical rea Application of PD red coloration or Kof the PD+ react aK- reacting subg Crystallog confirmed that thc contained the prot and usnic acid in in this group. Ti those specimens re Salazinic acid acr This group always norin. 205 crystallography. For the distinction between 5. complanata and B. subasperata, see discussion under the latter species. Thalli of 3. complanata are highly variable in their chemical reactions to different testing compounds. Application of PD to the medulla either produces a yellow— red coloration or no color at all. Further treatment with K of the PD+ reacting group separates it into a K+ red and a K— reacting subgroups. Crystallographic tests and thin-layer chromatography confirmed that those specimens reacting PD+, K- invariably contained the protocetraric acid as a medullary substance and usnic acid in the cortex. Rarely atranorin was present in this group. The data obtained by the same methods from those specimens reacting PD+ and K+ showed the presence of salazinic acid accompanied by traces of norstictic acid. This group always yielded usnic acid and frequently atra— norin. The sample with a PD— reaction was of a more hetero- geneous nature than the PD+ one. The application of K to the medulla of these specimens separated them into two categories; one reacted K+, variously red or pink and those with a K— reaction. The first group further reacted C+ and KC+ red, and thus was easy to circumscribe. By the use of micro-chemical tests and chromatography it was shown that these color reactions could be due either to cryptochloro— phaeic or ramalinolic acid. The seconc reacting PD-, K' 3 different chemicai other KC-. I C011: however, it former tolic acid. Chror benzene-dioxene-al Spots which may rt or a single subst determined substa: Specimens contain 0f a single speci the SO-called sub The perce various substance based exclusively Puerto Rico. Four acid 206 The second group of specimens was formed by those reacting PD-, K-, C—. Treatment with KC identified two different chemical variants, one reacted KC+ red and the other KC-. I could not identify the KC+ reacting substance; however, it formed crystals very similar to those of perla- tolic acid. Chromatograms of this unknown substance in benzene-dioxene-acetic acid (90:25:4) showed two different spots which may represent either two different substances or a single substance and its hydrolysis product. This un- determined substance was not further investigated. The KC- specimens contained either divaricatic acid or in the case of a single specimen from Jamaica a substance similar to the so-called substance ”H” in Cladonia. The percentage of specimens containing each of various substances is shown in Table 11. The analysis is based exclusively on a total of 214 collections I made in Puerto Rico. Four acids (salazinic, protocetraric, divaricatic and unknown A) were never found together in pairs or in any other combination. Norstictic acid occurred in every thallus containing salazinic acid but always in trace amounts. Crystallization of norstictic acid in G.A.o~T. Was never accomplished but traces were always demonstrated on chromatograms. Atranorin is an accessory substance, having a frequency of 6.5% in specimens containing either salazinic or protocetraric acid. Table 11. Percen from P const 1 tests Lichen Substances ——_— Usnic acid Salazinic acid Norstictic acid ( Divaricatic acid Protocetraric aci Atranorin Unknown subs t anc e \ Specimens 0f the sample and a frequency of 4. acid. 207 Table 11. Percentage of 214 specimens of 3. com lanata from Puerto Rico with each of seven-chemical constituents as determined by microchemical tests and thin—layer chromatography. Lichen Total Substances Specimens Percentage Usnic acid 214 100 Salazinic acid 106 49.0 Norstictic acid (traces) 106 49.0 Divaricatic acid 53 24.8 Protocetraric acid 46 21.4 Atranorin 14 6.5 Unknown substance A 9 4.2 Specimens with divaricatic acid constituted 24.8% of the sample and those having the unknown substance A with a frequency of 4.2% exhibited association only with usnic acid. This latter acid was present in every thallus I examined and is responsible for the stramineous color of the species. The chemical data of the Puerto Rican population of Ramalina complanata, thus, falls into four chemical variants, each characterized by a particular acid: salazinic acid, 49.0% frequency; divaricatic acid, 24.8% protocetraric acid, 21.4%; and the unknown substance A, 4.2%. A similar analysis for the specimens of the rest of the West Indies showed three other substances that never occurred in combii with any of the £1 Rico. The materi: in addition eithei or substance H (s: Combinati‘ from the West Ind cal categories. divaricatic, prot acids, substance W poses Problem. The Sit 208 occurred in combination with each other or in association with any of the four principal substances found in Puerto Rico. The material examined from other islands contained in addition either cryptochlorophaeic acid, sekikaic acid or substance H (see Table 12). Combination of the data from all specimens examined from the West Indies including Puerto Rico yields 7 chemi— cal categories. These are specimens containing salazinic, divaricatic, protocetraric, sekikaic, cryptochlorophaeic acids, substance H, or unidentified substance A. The high degree of chemical variation within 3. complanata poses a nomenclatural as well as a taxonomic problem. The situation in the species is not different from that elucidated in Usnea strigosa (Hale, 1962), Par— melia tinctoria (Hale, 1965), Ramalina siliquosa (W. L. Culberson, 1967) and Ramalina subdecipiens (Imshaug and Harris, 1971). These authors followed the suggestion forwarded by Lamb (1951) and retained the chemical variants of morphologically indistinguishable specimens as chemical strains within the species. Ramalina complanata is considered here to include 3. denticulata Nyl. and the rest of the chemical vari- ants within the species. In the writer's opinion the Species B. complanata consists of the following chemical variants, each one characterized by typically containing one of seven different principal medullary substances. w we —§;;_g ;: 4' ”‘ 209 + + + + Deon Uflcmb + + + wflom UHEOMOW WHHHONH + m + + + + + UHUN UflHNUflrHN>HHQ + UNUQ UHCHNQHWM. GHNHHOUHJLH .M + MMMMMMMMW .N + 1.. :7 ... 1., i va.rtv1\. UN-NNU-NVPM i ~v~ U... wu~ ~17.HJ1~1.~ 7.: «PN . \. ..,.~.~..-.:,v 1.4 + + + + mowaoomawpfioo .m + + mflmdofimESU .m + + < museumnSm + z ooampmnsm + wflow oaexflxom + + wflum oacflNmHem + . + + wave aflwewpooopoam + wave oflueofigo>flm wflom oaomQQOHomooumzau I mumamfimfioo .m + + + + +-++ + m ooceumnsm wave uaemamom l mflhmofinooo .m + + -+ .IIIIIIIIIII l + + + egommoumawo .m + + + mpHODmfln . 1|l|ll||| l + + dwmdfiowww ..m mmoocm um mpcmfiae> HeoflEono d 8 O .1 I .1 no 8 Q0 8 W G D D .A d f H O I w. n o i w. i m 1 1 m m m w n. m m m. m. use 0 l l I. 9 . . 1 a e 1 u e u G. 1 t. D. u on i a .d e mofloomm ecHHeEem r. m 3 n r.oo a e A T r. u a D. t. 1 r. e P a a on D. o D. D. r. n u r. I u o 3 u a s o a a a o u o r. o o D e I p S D I. b B n B I H O I e B n d A S I. I S u 9 9 m I D e I 1 e e o e u u . u P D; S S ) H S ( 210 {\i I ouqemeSm efiom chums wflow uwmxflxom 1 wave Ufipmoflhm>am I + + + + .III‘ILII. NOQWD .M l‘l‘l‘l O mpcmflpoaom Illl-I‘II‘II'I‘I. m eocHEmHum .m e©H03HHm QSm .m + + eflom ofimxflxom efloe ofiqflweaem wave oahwapooopoam + eflum uHDmoHDm>HQ mumHoMmensm vflom 0HdflNwHem wave uaemHmom emOflonom ecefi>soom + + + + + + + + + + mmoucwhmg + + + nit + + + + + Mich ml o wfiow owmxfixom wfioe oflumoagm>am flocweucoe + + 32%: + + + + NEHOMmowMOH + + + UHUW UHCwD + + + + UHUN UHEOHOW WHHHUQH + + + + + m1 oét ml oil + wfluw Odumofihm>flm + + + + + + wwum UHGHNNHNM wwwHHoOHsm . mlcfil + NOUQCfiHQ% . + + + + + Uflum UMfiflNMHmm + + @flum UHHMUHHM>HQ mODMOHwQOD .m + on /J 47 r1 0 U Wm 147 or «.7. fwrrltfa 1 ,fl/ \(1 no! Vi tflorr U1 11.70 g“ fFfDU I ”I + + < 00¢09099w + E OUfidUWwa + + OHUM Ufidxdxww + + + + + + + @HUd UMCHNdem .r iairlLEf Chemical variz Chemical vari: Chemical vari: Chemical vari Chemical vari Chemical vari Chemical vari The first basis of nomencla in descending 0rd invariably cont ai 211 Chemical variant I (divaricatic acid or typical) Chemical variant II (salazinic acid) Chemical variant III (protocetraric acid) Chemical variant IV (sekikaic acid) Chemical variant V (cryptochlorophaeic acid) Chemical variant VI (unknown substance A) Chemical variant VII (substance H) The first two variants have been ranked on the basis of nomenclatural priority; the other five are ranked in descending order of frequency. Chemical variant IV invariably contained ramalinolic acid in association with sekikaic acid; however, I chose to refer to the strain as a sekikaic acid variant because, unlike ramalinolic, sekikaic acid is easily recognized even when it is present in minute amounts by a simple micro-chemical test in G.E. Follmann and Huneck (1969) reported obtusatic acid from a south African specimen of 3. complanata, which might constitute an eighth chemical variant. This chemical type I have not seen and therefore it is not included in this report. Ecology: It is apparent that these substances are not randomly associated but that they tend to replace each other along a geographical gradient. Huxley (1938) pro- posed the term topocline for any morphological character variation on a regional basis in flowering plants. Although locally the chemical variants are sympatric when the whole spectrum of medul the range of the variation emerges species. Follman a W as being phillous, hygroph and of pan tropic in Puerto Rico of and in salt Spray On the nc the PTO’Cocetraric on the second grc Population is as: rarely With 3 cr Illeiia Cl . ado . \7 fi 1 collected in the 212 spectrum of medullary substances is evaluated throughout the range of the species, some indication of chemo-clinal variation emerges in relation to chemical character of the species. Follman and Huneck (1969, p. 188) described 3. 39m- planata as being "corticolous, acidophillous, subnitro— phillous, hygrophillous, thermophillous and photophillous and of pan tropical affinities." I made several collections in Puerto Rico of specimens growing on calcareous rocks, and in salt sprayed habitats. On the northern lowland coastal plain of Puerto Rico the protocetraric acid containing population is well developed on the second growth scrub of Chrysobalanus icaco. This population is associated with Ramalina sorediosa and more rarely with 3. complanata containing salazinic acid. Par— melia, Cladonia and Cladina are other genera which were collected in the same general area. The divaricatic acid strain was most frequently found on the trunks of Tabebuia heterophylla and Erythrina poeppigiana. Specimens containing the unidentified sub— stance A were collected in association with this variant. Specimens with salazinic acid were also found growing mixed With the PD— variant as well as with the small species 3. subasperata and 3. gracilis. Usually the lichens growing in mixed condition were smaller and rarely fertile. Typ— ically the mixture of the three variants (divaricatic, salazinic. and pr Ram m lower montane rai Population W35 0f not to be confine tions of this Val however, it was i and in the southe In the 1c the species 15 ft cafi and species tion has develOP‘ Generall) ferred cool habil year while the P1 climatic factors . 213 salazinic, and protocetraric acids) occurred on branches of Rapanea ferruginea, one of the common tree species of the lower montane rainforest. The salazinic acid containing population was of a more general occurrence and it appeared not to be confined to a particular vegetation type. Collec— tions of this variant were made in all vegetation zones; however, it was found to be rare on the limestone hills and in the southern deciduous forest. In the lower montane forest and the montane forest the species is found in association with Teloschistes flavi- cans and species of Usnea, especially where a scrub vegeta— tion has developed. Generally the PD- population of the species pre— ferred cool habitats where rainfall is over 100 inches a year while the PD+ population seemed to be independent of climatic factors. It should be noted that at elevations above 500 m. Ramalina complanata is abundant on the trunks of roadside trees and fence poles. Distribution: This species has a tropical and sub- tropical distribution with the center of dispersion appar— ently in the West Indies. Nylander (1870) cited collections from Australia, Africa, Brazil and Peru and Magnusson (1955) reported a PD— population from Hawaii. In North America the species is well represented in the southern United States (Howe, 1914; Moore, 1968). Within the island of Puerto Rico the distribution of B M tion containing P common on the low and unknown A Sub of the island, Wh are widely distri In the We the Greater Antil Six out of the se represented in J3 both islands 3 £014 The Cuban Variant acid types, both zinic acid varian Antilles and, 111‘ only one represer \1 ' raterial [MICH]; W Web. r West Ind] dunes, 7 km. west at sea level, Lar S \ g, fl 5663 6. , Landron 6\l s \, 66\2, 663C, r 3221 \, 22243, @ 214 of R. complanata is tied to ecological factors. The popula- tion containing protocetraric acid is found to be more common on the lowland coastal plains, the divaricatic acid and unknown A substance are typical of the cool central part of the island, whereas the salazinic acid containing plants are widely distributed throughout the island (Figures 14-18). In the West Indies the species is more common in the Greater Antilles than in the Lesser Antilles (Table 12). Six out of the seven chemical variants recognized were represented in Jamaica and Hispaniola, five were common to both islands, four occurred in Puerto Rico and two in Cuba. The Cuban variants are the divaricatic acid and salazinic acid types, both of which reached Florida, while the sala— zinic acid variant is the only one represented in the Lesser Antilles and, likewise, the divaricatic acid variant is the only one represented in Trinidad. Material seen: Exsiccati examined. Merr. no. 97 (MICH); W. Web. no. 147 (MICH). West Indies. PUERTO RICO: Laguna Tortuguero, sand dunes, 7 km. west of Vega Baja, on coconut and cashew forest at sea level, Landrén 519A, 528A, 528B, 529A, 535, 537B, 545, 552A, 555A, 566B, 587A, 602, 1967 (MSC); old pasture field Covered with guava scrub, at sea level, 7 km. west of Vega Baja, Landrén 613, 622C, 623, 632, 633. 638, 650. 652. 654, 1969 (MSC); pastu‘ m, 19;, 717A, 1 77213, 77913, 802, ‘ 843, 844, 858A, 1 Vega Alta, Landr6 tree along countr Barrio Maricao, c old cherry planta 1969 (MSC); Lagun (445C); Bosque Est “93> Cavev, Land 449$. 130713, 132 Disc); Laguna Car 215 1969 (MSC); pasture field, with Chrysobalanus icaco, Landron 700, 703, 717A, 737, 7493, 7513, 7533, 7553, 760A, 766A, 761, ————————_——_————.~——_—_——_—__._— _—._—_——-—_————— 843, 844, 858A, 1967 (MSC); Punta Cerro Gordo, coconut field, Vega Alta, Landron 859, 888, 896, 904B, 1967 (MSC); on cork ——_—_—_ tree along country road P.R. 677, 7 km. south of Vega Alta, Barrio Maricao, ca. 100 m., Landrdn 2286A, 2286C, 1968 (MSC); old cherry plantation, Los Hoyos, Vega Alta, Landr6n 2195, 1969 (MSC); Laguna Tortuguero, southeast of Lake Tortuguero, Landrén 1068, 1981A, 1074, 1078, 1087, 1090B, 1093B, 1967 —————_——_———..———_— —_—_—__—____—_—_— 1293B, 1307B, 1329A, 1341A, 1361C, 1366B, 1369A, 1389C, 1967 (MSC); Laguna Cartagena, pasture invaded by Bucida buceras, Landrdn 1506D, 1969 (MSC); Observation Tower, Monte del Estado, 840 m., Maricao, Landrén 1585B, 1630B, 1967 (MSC); Cerro Farrallén in a Tabebuia rigida forest, Landr6n 1665C, 1967 (MSC); Cerro Dona Juana, in Eucal tus-Euterpe forest, 850 m., Landrén 1686B, 1967 (MSC); Bo. Caonillas, Villalba, in guava Scrub, 800 m., Landrén 1713A, 1713B, 1714A, 1714B, Cordillera Central, Divisoria, 800 m., Villalba, Landron 17293, 17333, 1740, 17423, 1743C, 1967 (MSC); Barrio Cuchillas Montafias de Corozal, Landron 1751, 1763, 1764B, 1778A, 1967 (MSC); E1 Negro, Montafias de Corozal, ca. 550 m., Landrén 1780A, 1780B, 1781B, 1783A, 1783B, 1784A, 1786A, AILU papa, 1967 (use) Landron 1792A, 1_7 1798B, 1800A, 1_8( Juana, Bosque T01 1811, mm. L81 E 1m, 1851C, tation, 600 m., ] guava scrub, 600 1M, 18$, 189: 191$, 191211 19‘ 216 1786B, 1967 (MSC); Barrio Padilla, Montafias de Corozal, Landr6n 1792A, 1792B, 1793B, 1794, 1795A, 1796A, 1796B, _—_———_—__—.——_—.————— 1798B, 1800A, 1800B, 1804A, 1805B, 1967 (MSC); Cerro Dofia ___—_————.—-_————_—_.__ Juana, Bosque Toro Negro, Landr6n 1807B, 1910B, 1911A, 1811C,1816A, 1816B, 1826B, 1827A, 1827B,1830C, 1831A, __W——___—.__-__—_——————.———— 1832C,1836B,1836C,1839B,1839C,1846B,1847B,1848B, —_W——._———__—___—_.__—_————_ 1850A, 1851C, 1855D, 1967 (MSC); Lago Carite in coffee plan— tation, 600 m., Landrén 1870, 1882B, 1967 (MSC); Monte Cuba, guava scrub, 600 m. near Barranquitas Hotel, Landr6n 1884A, 1884D,1885,1893A,1893B,1893D,l894C,1904A,1904B,1905A, ——W_—_——_—_-—_———_——_— 1910B, 1912D, 1913A, l917D, 1925B, 1933B,1937A, 1937B, “W“—_————___—_ 1937C,1938A,1941B,194ZD,1948A,1967 (MSC); Hotel Barran- ~W——_________ quitas, Landrdn 2386A, 2388, 1969 (MSC); Monte Llano, Cidra, Landron 1957B, 1957C, 1967 (MSC); Cerro Pulguillas, mahogany along the road, Aibonito, Landrdn 1970, 1971B, 1972A, 1967 (MSC); Cuy6n, Aibonito, on a guava acrub, 500 m., Landrdn 1976B, 1978B,1978C,1980C,1981B,1987B,l994A,1995A, ——-—————._W—_——_————————-——____ 1997B, 1998B, 1999A, 2000A, 2006C, 1967 (MSC); Bafios de ‘—_——W_———_ Coamo on Delonix regia, Landron 2007, 2008, 1967 (MSC); Barrio Los Hoyos, abandoned field, Vega Alta, Landron 2195, 1968 (MSC); east of Laguna Tortuguero, Vega Baja, Landrén 2221, 2224B, 2229A, 2235A, 2235B, 2242A, 2247, 2256, 1968 ‘_ _.___ —___’ _—_—, ._—_’ ________’ ——, —__, (MSC); Lago dos Bocas, near dam, Utuado, ca. 250 m., Lan— —— men 2262, 2266, 1968 (MSC); Guajataca beach, Landrdn 280 ’ 1968 (MSC); Barrio Maricao, Vega Alta, along road, on Cork tree, Landrdn 2286, 1968 (MSC); Bosque Toro Negro, near,, Divisoria, Landrr‘ Sebastian , Landrc‘ Cafiaboncito, Cagl Espinosa, on trur Aibonito, open r4 llanati, on lemon trees, Cook and 4 m 9588, 19. “fl. 1913 (MICH 217 Divisoria, Landrén 2310C, 1968 (MSC); Guajataca lake, San Sebastian, Landrén 2460, 2461, 2462A, 1969 (MSC); Barrio Cafiaboncito, Caguas, Landrén 2456, 1969 (MSC); Vega Alta, Espinosa, on trunk of grapefruit, Stevenson 2351, 1914; Aibonito, open road posts, Fink 1828, 1878, 1916 (MICH); —— Manati, on lemon twigs, Fink 2119, 1916 (MICH); Cayey, on trees, Cook and Collins 5.3. (MICH); Asomante, Britton and Britton 9588, 1930 (MICH); Lares, Britton, Britton and Hess 2764, 1913 (MICH); Mayagfiez, on Eucalyptus, Fink 1362, 1916 (MICH); Rio Piedras, open road side on posts, Fink 86, 1915 (MICH); Santurce, Stevenson, 3.2. (MICH); Vega Baja, Fink 2154, 1916 (MICH); Coamo Springs, on Pictetia, Britton and Britton 9685 (MICH); Caguas, Heller 3.3. (MICH); Naranjito at 1500—2000 ft. on rocks, Pink 228, 1915—16 (MICH). JAMAICA: Parish of Hanover, Birchs Hill, 1809 ft., Imshaug 15684, 15705, 15708, 1953 (MSC); Parish of Hanover, Dolphin Heads, 1789 ft., Imshaug 15641, 15659, 1953 (MSC); Parish of St. Catherine, on cactus, sea level, Port Henderson, Imshaug 13842, 1952 (MSC); between Long Pond and Hellshire Bay, sea level, Imshaug 13616, 1952 (MSC); summit of Montpellier, 2300 ft., Imshaug 14309, 14263, 14284, 1953 (MSC); Fort Clearance, sea level, Imshaug 13430, 13435, 1952 (MSC); between Hollymount and Mount Diablo, 2750 ft., Imshaug 2,2,, 1953 (MSC); Parish of St. Ann, Dunn's River, 150 ft., imghgpg 15747, 15755, 1953 (MSC); Mosely Hall Cave, 2000 ft., Guvs Hill, Imshaug 13648, 13678, 1952 (MSC); Albion, 2400 ft., {15.41.3145 fl, 1: fl, 1953 (M30) White River, 300 Ridge, 3100 ft-2 willow trunk, 52 1912 (PH); Catad‘ phi 2,1,, 7-1I (413); Kingston, : locality given, 1 ity given, m 1909 (FH); 110 5P' specific localitj area at Armenia, llaestra, Oriente cultivated area 218 Imshaug 15895, 1953 (MSC); Browns Town, 1800 ft., Imshaug 15934, 1953 (MSC); Parish of St. Mary, Prospect Estate, White River, 300 ft., Imshaug 15773, 1953 (MSC); Blouxburgh Ridge, 3100 ft., Imshaug 15068, 1953 (MSC); Rose Hill, on willow trunk, Earle J-58, 1952 (MSC); Arntually, Orcutt 2695, 3073, 3081, 3090, 1927 (US); Mandeville, Cushman 181, 1912 (PH); Catadura, Cushman 23, 1912 (PH); Whitefield Hill, Plitt i'E'a 7—11-26 (US); Old Harbor, Orcutt 2522, 1927 (US); Kingston, Thaxter E'E': Feb. 15, 1891 (MICH); no locality given, Cummings 179, 1904 (PH); no specific local- ity given, Walle §.£. (FH); no locality given, Maxwell 998, 1909 (PH); no specific locality, Hart 103, 1884 (PH); no specific locality, Wright ll, 1909 (PH). CUBA: Cultivated area at Armenia,on slope of El Gato, Loma del Gato, Sierra Maestra, Oriente, Imshaug 24748A, 24748B, 23910, 1959 (MSC); cultivated area facing the sea, below Colegio de la Salle, Loma del Gato, Imshaug 24924, 1959 (MSC); Las Pailas, Bro. —_ Hioram §§5§, 1923 (US): Boca de Jaibo, Bro. Hioram 5495, 1921 (US); Mata Abajo Estate, Guantanamo, Bro Hioram 3.2., Feb. 27, 1920 (PH—180091); Novaliche, Bro. Hioram E‘E" Jan. 15, 1920 (PH—13035); locality not given, Wright 51 (FH). DOMINICAN REPUBLIC: On ridge between Santiago and La Cumbre, Cordillera Septentrional, Wetmore 3836, 3852, 1958 (MSC); in pasture area hill, ca. 500 ft., overlooking Sosfia Bay, Prov. Pto. Plata, Wetmore 3930, 1959 (MSC); along road _ at 10$ Amaceyes, Wetmore 3380, 3388, 1958 (MSC); along Summit 0f ridge 8 4450; near Peace [msc); 0n Slope 0 Plata, M E 250 ft., Imshaug [111CH). HAITI: Kenscoff, ca. 504 Furcy, M E Perchoir, near Bc 425i, 1958 (MSC) Moi, 1958 (MSC: 1958 (MSC); belov 1958 (MSC); vi cir 219 summit of ridge at Guama, Prov. Santiago, Wetmore 3913, 1958 (MSC); near Peace Monument, Santiago, Wetmore 3827, 1958 (MSC); on slope of La Cumbre, between Santiago and Pto. Plata, Imshaug 23850, 1958 (MSC); on palms above Santiago 250 ft., Imshaug 23787, 1958 (MSC); Wright §.g. (Flora Domingensis no. 1), 1871 (PH); Rose 3.3., Feb. 20, 1913 (MICH). HAITI: Department de l'Ouest, north slope above Kenscoff, ca. 5000 ft., Imshaug 22501, 22489, 1958 (MSC); Furcy, Imshaug 22709, 1958 (MSC); on hillside below Le Perchoir, near Boutillier, Morne H6pital, Imshaug 22525, 22501, 1958 (MSC); hillside west of Cap Haitien, Imshaug 22656, 1958 (MSC); Cocos behind Hotel Randan, Imshaug 23151, 1958 (MSC); below Citadelle south of Millot, Wetmore 2840, 1958 (MSC); vicinity of Jean Rabel, Leonard and Leonard 18903A, 1929 (US); vicinity of Cabaret, Baie des Mustiques, Leonard and Leonard 11893, 1929 (US). GRAND CAYMAN: Hell Hole, west of Bay Town, Imshaug 24463, 24475, 1959 (MSC); near Red Bay, Imshaug 24431, 24514, 24527, 24537, 1959 (MSC); near Coral Castle, Imshaug 24540, 24548, $959 (MSC). GUADE— LOUPE: Ste. Anne, Duss 688, 1093 (MICH). TRINIDAD: Coast at Point Fortin, Guapo Bay, Gulf of Paria, Imshaug 32220, 1963 (MSC); trunk of Sabal palm, Broadway 50, 1907 (PH). BARBADOS, Imshaug 16339, 16432, 1953 (MSC). BERMUDA: Trees in wood, Britton, Brown and Searn, five collections without data, 1912 (MICH); Davonshire, Hervey s.n.,l915 (PH). BAHAMAS: San Salvador, beach along French Bay, Gillis 5271B, 19 Central 1143.: 1897-98 (1 1932 (MICH); Sar 2978A (MICH); TL North Am 1912 (MICH); Lor H‘ 4443. 1874 Domingo (non V14 1 Desc\rip 011g) Hot Pendu late, ending ln 0% 220 Gillis 5271B, 1963 (MSC). Central America. YUCATAN: Chichankanab, Gaumer 1343, 1897-98 (MICH); Champoton, Campeche, Steere 1810A, 1932 (MICH); San Miguel, Cozumel Is., Quintana Roo, Steere 2978A (MICH); Tuxpena Campeche, Lundell 1111, 1931 (MICH). North America. FLORIDA: Turtle Mt., Kelly 111, 1912 (MICH); Long Boat Key, Kellz 189, 193 (MICH); Lee Co., Standlez 3.3. (MICH); Dade Co., near Homestead, on roadside palms, Culberson and Culberson 10958, 10854, 1962 (DUKE); Charlotte Co., Fort Meyers, Culberson and Culberson 10877, 1962 (DUKE). South America. COLOMBIA: Nova Granada, Lindig 31 (FH). VENEZUELA: Western end of Tortuga Is., Tazlor 730, 735, 1939 (MICH); Rancho Grande, north of Maracay, Test 3 (MICH). BRITISH GUIANA: Georgetown Bot. Garden, Linder 2339, 1923 (MICH). BRAZIL: Sta. Catherina, Pabst, 3.3. (FH); Kunze 3.3. (FH). PERU: Winterfield 3.3. (FH). 10. Ramalina sorediantha Nyl. Ramalina sorediantha Nyl. Bull. Soc. Linn. Norm. II. 4:143. 1870. Original material: Jamaica and Sto. Domingo (non vidi). Description: Thallus pale stramineous, to 6 cm. long, not pendulous; branches linear, compressed, canalicu- late, ending in dichotomous tips; surface scrobiculate, often minutely punctiform, sorediate; soralia globose, large, to 4 mm. a to a maximum of 5 usually both type convex, almost re thiCk; hymenium 1 Sivole; epithecir containing eight sePtate, hyaline 34.511. red. 221 large, to 4 mm. across, marginal and terminal. Apothecia to a maximum of 5 mm. lateral or dorsal to the channels, usually both types on the same thallus; disk concave or convex, almost red, pruinose; hypothecium colorless, 10-16u thick; hymenium tinged with brown, 50—80u thick; paraphyses simple; epithecium dark brown, 6-10u thick. Asci clavate, containing eight spores in one oblique row; spores uni- septate, hyaline, subfusiform, straight or bent, 12-16 x 3-4.5p. MedullarZ reactions: PD+ yellow-orange, K+ yellow- red. Lichen substances: Atranorin, salazinic acid and usnic acid. Discussion: Ramalina sorediantha is a canaliculate species with terminal or lateral soredia. This species can be confused with 3. leptosperma if the soredia are not well developed, but then it can be distinguished by the presence of salazinic acid. The apothecia are not restricted to a zone, but rather are dispersed on the surface of the laciniae, although most are marginal. The species is not always clearly canaliculate or scrobiculate; however, it is always of a homogeneous chemical constitution and we can rely on this character to separate it from B. leptosperma, Which contains norstictic acid. Of all the West Indian species examined, 3. sore- diantha was the only one to have some red coloring material 4‘ 1 in the hymenial 2 from the epithec decreases toward that this colori: mounting medium ‘ Species that occ forest between 7 Distribu \ Species is chara America. In the Cuba, Jamaica, H Puerto Rico. It 222 in the hymenial layer. This appeared to be moving down from the epithecium since the concentration of the dye decreases toward the base of the hymenium. It is unlikely that this coloring material was due to dissolution by the mounting medium which was distilled water. Ecology: Ramalina sorediantha is a corticolous species that occurs in the zone of the lower montane rain- forest between 750 and 2750 ft. of elevation. Distribution: Zahlbruckner (1930b) noted that the species is characteristic of meridional Europe and Tropical America. In the West Indies it has been recorded from Cuba, Jamaica, Haiti and Dominican Republic but not from Puerto Rico. It is absent from the Lesser Antilles, but the species reappears in Venezuela and Brazil in northern South America and in the Galapagos Islands. The species has also been recorded from Yucatan Peninsula in Central America a short distance from Cuba. This species appears to be sub-cosmopolitan of the neo—tropics in its distribu- tion. Material seen: Exsiccati examined. W. Web. no. 147 (MICH). West Indies. CUBA: La Prenda, Bro. Hioram 6795B, 1923 (US); Wright 3.3. (FH); JAMAICA: Parish of St. Ann, White River, 500 ft., Imshaug 15792, 1953 (MSC); Hopewell, 1400 ft., Imshaug 15916, 1953 (MSC); Lime Hall to Green Park, 1500 ft., Imshaug 15844, 1953 (MSC); Browns Town, 1800 ft. , Imshau Imshaug 15980, _1_ Imshaug 15891, _1_ Goshen, 1000 ft. 800 ft., Imshaug lawny, Clarks To Parish of St. An Hills, 2250 ft., “154% 1L7, 163 (M 39g, 192 area along Summi WM, 1958 Also); along roe trional, Wetmore \ 1°44, Hillside t. Cultimted are a 1411“, ca. 2300 nest of JaCmel .1 223 1800 ft., Imshaug 15932, 1953 (MSC); Goshen, 1000 ft., Imshaug 15980, 15982, 1953 (MSC); Claremont, 1400 ft., Imshaug 15891, 15893, 1953 (MSC); between Hopewell and Goshen, 1000 ft., Imshaug 15989A, 1953 (MSC); Stewart Town, 800 ft., Imshaug 16018, 16022, 1953 (MSC); Parish of Tre- lawny, Clarks Town, 750 ft., Imshaug 16035, 1953 (MSC); Parish of St. Andrew, south slope of Cooper's Hill in Red Hills, 2250 ft., Imshaug 13726, 1952 (MSC); Mandeville, Cushman 117, 163, 1912 (PH); Cushman 3.3. (MICH); Lumsden, Orcutt 3962, 1927 (US). DOMINICAN REPUBLIC: Cultivated area along summit of ridge at Guama, Prov. Santiago, Wet— more 3899, 1958 (MSC); on low shrub over coral rock on coastline at Sosfia, Prov. Puerto Plata, Imshaug 23910, 1958 (MSC); along road Los Amaceyes, 1850 ft., Cordillera Septen- trional, Wetmore 3395, 1958 (MSC). HAITI: Department du Nord, Hillside west of Cap Haitien, Imshaug 22674, 1958 (MSC); cultivated area on hillside below the Citadelle, south of Milot, ca. 2300 ft., Wetmore 2833, 2844, 1958 (MSC); north- west of Jacmel, ca. 3000 ft., Thomas 13, 13, 1935 (FH). Central America. YUCATAN: On trees in forest, Uxmal, Steere 2045, 1932 (MICH). ll. Ramalina 32233323333 Nyl. Ramalina leptosperma Nyl. Flora 59: 412. 1876. Lectotype (nov.): Las Lomas near Canto, Cuba, Wright 27D (PHI). Ramalina Lectotype (nov.) Descript too cm. high, b linear, canalicu face densely scr Apothecia freque adpressed agains cave and pruinos tomously branche SPores arranged hyaline, ellipsc 224 Ramalina scrobiculata Mfill Arg. Flora 68: 500. 1885. Lectotype (nov.): Dominican Republic, Prenleloup 111 (G1). Description: Thallus stramineous, caespitose, rigid, to 6 cm. high, branching dichotomous; branches 2-3 mm. broad, linear, canaliculate, tips usually bifid and compressed; sur— face densely scrobiculate, esorediose; cortex cortilaginous. Apothecia frequent, to 4 mm. across, marginal and closely adpressed against the thallus; disk reddish to dark, con— cave and pruinose; hymenium 60—80p thick, paraphyses dicho- tomously branched at the tips. Asci clavate, containing 8 spores arranged in two regular series, spores uniseptate, hyaline, ellipsoid, straight or curved, small 9-13 x 3—4u. Medullarz reactions: PD+ yellow, K+ yellow-red, KC-, C—. Lichen substances: Atranorin, Chloroatranorin, norstictic acid and usnic acid. Discussion: The types of Ramalina leptosperma and R. scrobiculata were identical as regards spore size, channeled branches and scrobiculate thallus surface with the medulla containing norstictic acid. The species can be confused with non-sorediate specimens of 3. sorediantha. Normally the species are distinguished by the esorediose thallus of E. leptosperma against the sorediate thallus of 5. sorediantha. Addi— tional difference is provided by the production of norstic— tic acid by the former and salazinic acid by the latter. Likewise, the es the production c The medt 1+ red and PD+ ) norstictic acid of salazinic aci layer chromatogr are very variabl Chloroatranorin major substance. it is expected 1 atranorin is 315 1968, p. 282). 225 Likewise, the esorediose species 3. straminea differs by the production of salazinic acid. The medullary color reactions of this species are K+ red and PD+ yellow turning orange. The thalli contain norstictic acid as the major medullary constituent. Traces of salazinic acids were, however, demonstrated by thin- layer chromatograms. Other lichen substances in the species are very variable and inconstant, usnic acid, atranorin and Chloroatranorin appeared in different combinations with the major substance. Because of the origin of Chloroatranorin it is expected that whenever this substance is present atranorin is also present (C. F. Culberson, 1964; Huneck, 1968, p. 282). Ecologz: The species in Florida occurred in the southernmost tip of the Everglades swamps. The Puerto Rican specimens were collected on the wouthwest corner of the island where rainfall is very low and temperature reaches the highest extreme in the island. The vegetation is a xerophytic scrub of leguminous and cacti plants, bor- dered by mangrove swamps. The general area is usually under the influence of sea winds and salt spray. Distribution: The species Ramalina leptOSperma has been identified from southern Florida, Cuba, Dominican Republic, Haiti, and Puerto Rico. It is not known to occur in Jamaica or in the Lesser Antilles, but it occurs in Brazil. From this distribution the species can be regarded as neo- The spec (1888) from 3 1c specimen collect are from localit exhibited a sout Material W 2185 —— 1923 (US); La P1 1' . lChe, M Bro. H' w 3.3. W, Cap Haitie Bahia P1873 Suci Worth Am 2M. 1967 (MS, 4% @515 (FH). 12. Ramalina 1 ) Aw 226 regarded as neo—tropical. The species was reported from Puerto Rico by Mfiller (1888) from a locality near Guanica and I have seen another specimen collected in Bahia Playa Sucia. Both collections are from localities in the south west littoral zone and exhibited a southwestern distribution (Figure 19). Material seen: West Indies. CUBA: Novaliche, Bro. Hioram 2189, 1919 (US); Las Pailas, Bro. Hioram 6868, 1923 (Us); La Prenda, Bro. Hioram 6795B, 1923 (US); Nova- liche, Bro. Hioram s.n., 1920 (lSOSS-FH); Novaliche, Oriente, Bro. Hioram 3.2., 1918 (12371—FH). HAITI: Départment du Nord, Cap Haitian, Imshaug 22663, 1958 (MSC). PUERTO RICO: Bahia Playa Sucia, Britton, Cobwell and Brown 4769, 1915 (US). North America. FLORIDA: Everglades, Harris 2853A, 2856B, 1967 (MSC). Additional material seen: South America. BRAZIL: Pabst 3.3. (FH). 12. Ramalina straminea (Pers.) Ach. Phxscia straminea Pers. Ann. Wetter. Gessellsch. 2: 18. 1811. Ramalina straminea (Pers) Ach. Syn. Meth. Lich. 294. 1814. Original material: Dominican Republic (non vidi). Description: Thallus stramineous, erect, to 4 cm. high; branching subdichotomous; filaments rigid, linear, canaliculate, surface smooth, epapillate and esorediate; cortex cartilag: across; disk ta] spores; spores i ellipsoid, lZ-l Medulla red, Kc-, c-_ Lichen \ Discuss SPecies that is the absence 0 f “13- ama; tiated by the a by Characterist resembltis small 3-6h), from Whi The Species her 227 cortex cartilaginous. Apothecia common, lateral, to 4 mm. across; disk tan, concave or flat. Asci containing 8 spores; spores uniseptate, hyaline, straight or substraight, ellipsoid, 12—17 x 3-4u. Medullary reactions: PD+ yellow-orange, K+ yellow— red, KC-, C—. Lichen substances: Salazinic and usnic acids. Discussion: Ramalina straminea is a canaliculate species that is easily distinguished from R. complanata by the absence of papillae on its surface and their presence in R. complanata. From Ramalina sorediantha it is differen— tiated by the absence of soredia and from R. scrobiculata by characteristically producing salazinic acid, while the latter species produces norstictic acid. The species resembles small specimens of R. alludens (spores 30-40 x 3-6u), from which it differs by its much smaller spores. The species here described is represented in the material studied by a single collection from Santo Domingo; however, it is a good collection that represents Acharius‘ concept of the species. Ecology: There is no record of the habitat of this Species. Distribution: The species has been recorded only from Santo Domingo. Material seen: West Indies. DOMINICAN REPUBLIC: Wright, Parry, Brummel s.n. (Flora Domingensis no. 3), 1871 (PH). 13. Ramalina g Ramalina 4: 120. 1870. C Descript main branches di lets; branches ] ate and filiforn POCket-like sore small, to 2 mm. then subgenicula Colorless, 12-“ Went. R ca \ 1 difficult to di. 228 13. Ramalina camptospora Nyl. Ramalina camptOSpora Nyl. Soc. Linn. Norm. II. 4: 120. 1870. Original material: Cuba, Wright (non vidi). Description: Thallus stramineous, erect, compressed, main branches dichotomous with numerous short lateral branch- lets; branches linear, about l.5 mm. at the base, but attenu— ate and filiform at the apices; soredia in chains of small pocket-like soralia along the margins. Apothecia rare, small, to 2 mm. across, lateral, sometimes subterminal and then subgeniculate; disk tan, pruinose, concave; hypothecium colorless, 12-16n thick; hymenium gelatinous, with simple paraphyses, 3—50u thick; epithecium dark colored, 6-10u thick. Asci clavate, containing eight spores in a single series; spores hyaline, uniseptate, septum straight, often oblique, ellipsoid or curved, 12-16 X 6—8u. Medullary reactions: PD-, K-, KC-, C-. Lichen substances: Caperatic and usnic acids. Discussion: This corticolous species is sorediate along the margins and slightly resembles R. farinacea, but it differs by its bifacial thallus and characteristic soredia disposed in a chain along the margins. This species is allied to R. bistorta by the oblique septum present in many spores and by being the only two Species in the genus Ramalina in which caperatic acid is Present. R. camptospora gives no color reactions and it is difficult to distinguish the fatty acid containing specimens from those that . Nylander his description Cuba (Wright), e is placed by Nyl Species. I have c mens and all of in many of these but the cortex ; 3' W i reactions were j the medulla, whj CaPeretic acid. M ......A_ ,..._...__«'v= i1»;— ’%:€~‘é‘;:12j:_i§ij§:éijé:Tél . 7'7 *—_I 229 from those that do not contain any lichen substances. Nylander (1870) did not cite a type specimen in his description of the species and referred to it as ”In Cuba (Wright), ex. hb. Tuckerman." Ramalina camptospora is placed by Nylander under the K- reacting group of species. I have chemically tested numerous West Indian speci- mens and all of them reacted PD-, K—, C-, KC-. The medulla in many of these specimens was devoid of chemical substances but the cortex produced usnic acid. Other specimens of R. camptospora in the West Indies with the same spot reactions were found to contain a fatty acid substance in the medulla, which was proved by crystallography to be caperatic acid. Ecology: Ramalina camptospora is frequent in the high mountains of the West Indies, between 3000 feet and 6000 feet elevation. Distribution: This West Indian species is found only in Cuba, Jamaica and Haiti and Nicaragua in Central America. It is absent from Puerto Rico and the rest of the Caribbean Islands. Material seen: West Indies. CUBA: Loma del Gato, Sierra Maestra, Oriente, Wright 3.3., 1857 (not the type) (PH); Imshaug 24945, 1959 (MSC); summit of Gran Piedra, ca. 1200 m., Sierra Maestra, Oriente,ImshaQa 25007, 25053’ 1959 (MSC). JAMAICA: Summit of Maccasucker Bump, St. Thomas, 825-102 Trail, near Har 1952 (MSC). DO ridge above Los 23297, 1958 (MS 5500 ft. near K 1958 (MSC); nea l‘Ouest, M 8800 ft., Dept. Of Téte Etang, WOUft” £3532 West of Forét d ridge aPPYOachi Central (FH). 14' Ramal ina R Ramalin DeSC ‘ ...‘ggp cm long, bra Pics Se 3 margin 230 Thomas, 825-1025 m., Maxon 9581, 1926 (PH); Moodie's Gap Trail, near Hardwar Gap, Blue Mountains, Imshaug 13077, 1952 (MSC). DOMINICAN REPUBLIC: Cloud forest 3000-3200 ft., ridge above Los Amaceyes, Cordillera Septentrional, Imshaug 23297, 1958 (MSC). HAITI: Summit of Montagne Noire, ca. 5500 ft. near Kenscoff, Départment de l'Ouest, Wetmore 2749, 1958 (MSC); near Forét des Pins, 5500 ft., Départment de l'Ouest, Imshaug 22852, 23089, 1958 (MSC); Morne La Selle, 8800 ft., Dépt. l'Ouest, Imshaug 22994, 1958 (MSC); summit of Téte Etang, on ridge between Kenscoff and Furcy, ca. 6000 ft., Imshaug 22613, 1958 (MSC); on shrubs in low hills west of Forét des Pins, Wetmore 3224, 1958 (MSC); along ridge approaching Morne Macaya, Wetmore 3304, 1958 (MSC). Central America. NICARAGUA: Wright §.p., 1853—56 (FH). 14. Ramalina peruviana Ach. Ramalina peruviana Ach. Lich. Univ. 599. 1810. Original material: Peru, Lagasca (non vidi). Ramalina finkii Zahlbr. Mycologia 22: 78. 1930. Lectotype (nov): Aibonito, Puerto Rico, Fink 1878A, 1916 (MICH!). Description: Thallus sordid gray, caespitose, to 7 cm. long, branching dichotomous, branches linear, com— Pressed, margins undulate to 2 mm. broad, apices multlfld; surface with pseudocyphella, striate, granulose, sorediate, With distinct soralia along the margins; cortex cartilaginous, strands seen thi small, 1-2 mm. 2 containing eight form or ellipsoi Medulla: Lichen 5 acids . Discuss: recognize by it: bordered by sor; are thin, it re: ChemiStrY, but : whereas R. SOTEc \ The plar 231 strands seen through the thin outer cortex. Apothecia rare, small, 1-2 mm. across; disk concave, brown. Asci clavate, containing eight spores; spores uniseptate, hyaline, subfusi— form or ellipsoid, straight or substraight, 10-17.6 x 3-4.5u. Medullarz reactions: PD—, K+, KC+, C+ pink. Lichen substances: Sekikaic, ramalinolic and usnic acids. Discussion: The species R. peruviana is easy to recognize by its compressed branches with irregular margins bordered by soralia with granulose soredia. If the branches are thin, it resembles R. sorediosa in its morphology and chemistry, but it always will be compressed at the base, whereas R. sorediosa is terete throughout. The plant identified by Moore (1968) as R. dendri- scoides in Florida is in reality R. peruviana. The species R. finkii was collected precisely where I found R. peruviana abundant, emphasizing the identity between the two species. The original collections of Zahlbrucker's species are Fink 1799 and 1878 (MSC, MICH, US). Both numbers are formed by mixed collections that contained R. subasperata, R. dendri— scoides, R. subpellucida, R. complanata and R. finkii. This last species is represented by small pieces of thalli which do not differ from the Acharian concept of R. peruviana. This species gives a faint red color reaction with K, C or KC and a negative reaction with PD. It contains sekikaic, ramalinolic and usnic acids. Zahlbruckner (1930a) reported R. E portions of the red (ramalinoli choosing Fink ] chemical reacti seen by Acharii from Paraguay g contained the s Huneck and F01] 3' perm/\iana v2 strong C+ red c in the medulla. Variant . Ma on the trunk 02 glades (M 1 Core, evel and at e 232 reported R. finkii as being K-; however, I have tested portions of the original material and it reacted K+ faint red (ramalinolic acid). Of these original materials I am choosing Fink 1878A (MICH) as lectotype. Although the chemical reactions of the original material of R. peruviana seen by Acharius is not known, I have found that specimens from Paraguay and Ecuador had the same color reactions and contained the same substances as those from the West Indies. Huneck and Follmann (1965) described a Chilean specimen of R. peruviana var. pollinariaeformes Vain. that produced a strong C+ red color. They identified the depside tumidulin in the medulla. I have not seen material of this chemical variant. Ecologz: In Florida the species has been collected on the trunk of the royal palm Roystonea and in the Ever- glades (Moore, 1968). In the West Indies it occurs at sea level and at elevations between 2000 and 3000 feet. The Puerto Rican population is typical of the lower montane rainforest, toward the center of the island. It is frequent between 1500—2500 ft., especially on the trunks of Swietenia, Tabebuia, Rapanea, Psidium and Erythrina. In the scrub vegetation of Psidium guajava it is usually associated with R. dendriscoides, R. complanata, Usnea, and Teloschistes. It forms large colonies at the base of old tree trunks. The species is absent from the wetter eastern mountains and from the drier western hills. Distrib isin Peru, and haul and Ride Mre reported i In the Gmater Antille Laser Antilles Him. In Puer 0fthe island, of the Cordillg isa130 found 5 233 Distribution: The original locality of this species is in Peru, and it has been reported by Vainio (1890) from Brazil and Riddle (1920) from The Bahamas. The species is here reported from Florida. In the West Indies, the species is primarily a Greater Antillean lichen, since it is absent from the Lesser Antilles except from Trinidad and Curacao (Riddle 1920). In Puerto Rico it is abundant in the central part of the island, less frequent in the central western part of the Cordillera and absent from the southern plains. It is also found in the northern part of the island near the sea shore, thus exhibiting a north-central distribution pattern (Figure 20). The general distribution of the species corresponds with the American tropics and temperate zones. Material seen: Exsiccati examined. Malme Austr. amer. I. no. 202 (MSC). West Indies. PUERTO RICO: Laguna Tortuguero, Vega Baja, Landron QRRR, 131, ZER, 129R, §lia 85R, 1051A, RRRQ, 1061, 1085, 1087C, 1098, 1967 (MSC); Bosque Estatal de Guavate, near river, Cayey, Landrén 1386, 1967 (MSC); Cerro Dofia Juana, 800 ft., Orocovis, Landron 1692C, 1693C, 1967 (MSC); Divisoria, 800 m., Villalba, Landron 1745C, 1967 (MSC); Barrio Cuchillas, Orocovis, Landron 1752, 1754, 1755A, 1756A, 1755B, 1760, 1761, 1783B, 1967 (MSC); Barrio E1 Negro, Orocovis near Morovis, R Bosque Toro Neg trees, Orocovis 14E» 1851B, 1 fl, 1876, 19 —I ., M L88 in, M, l w an, i 9&1967 (Ms M, I956C, 1 ‘Sland, near la 234 Negro, Orocovis, Landrén 1795A, 1967 (MSC); Barrio Padin, near Morovis, Landrdn 1795B, 1798A, 1801, 1803, 1967 (MSC); Bosque Toro Negro, Cerro Dofia Juana, pasture with scattered trees, Orocovis, Landrdn 1825C, 1831F, 1832A, 1836A, 1837B, .——-——-————_—_————_ 1847C, 1851B, 1967 (MSC); Carite lake, near Guayama, Landr6n 1873B, 1876, 1967 (MSC); Monte Cuba, near Barranquitas, 600 m., Landrdn 1884E, 1885, 1887B, 1888B, 1891B, 1892B, 1893D, _———___._—____————_——————. 1894B, 1895B, 1896B, 1905C, 1907B, 1909B, 1910C, 1912C, 1913D, 1915A, 1925A, 1928, 1935A, 1940C, 1942A, 1944B, 1947A, 1948C, 1967 (MSC); Monte Llano, Cidra, Landron 1952A, 1953, 1954A, 1956C, 1957A, 1958A, 1959A, 1967 (MSC); Treasure Island, near lake, Cidra, Landr6n 1964, 1967 (MSC); Cerro Pulguillas, Aibonito, Landrdn 1965B, 1967, 1969B, 1971c, 1973, 1977D, 1978D, 1980B, 1984A, 1987C, 1991, 1993C, 1999F, 2002, 2003, 1967 (MSC); in open spaces, posts and bark of trees, Fink 1799, 1916 (FH, MSC, MICH); Mayagfiez, Fink 1316, _ 1915 (MICH); Vega Baja, Fink 2138, 2154, 1916 (MICH); Maricao, Bro. Hioram §.p., 1911 (MICH); Dorado, icaco scrub, Landron 2199, 1968 (MSC). CUBA: On cultivated slope facing sea below, house of Colegio de la Salle, Loma del Gato, Sierra Maestra, Oriente, Imshaug 24922, 1959 (MSC); 'Boca de Jaibo, Bro. Hioram sggl, 1921 (FH). DOMINICAN REPUB- LIC: Cerrazo, Cordillera Septentrional, Wetmore gggg, 1958 (MSC). JAMAICA: Mandeville, Cushman 41, 1912 (PH); Catadura, Cushman Li, 1912 (FH, MICH); St. Mary Parish, Orcutt 4623, 1928 (US); Prospect Estate, White River, 300 ft., Mtg 15768, Cave, Guys Hill Wood to Moneagt Claremont, 140( 1000 ft. , Imshe Hanover, Lucea, (MSC); Parish c 400 ft., Imshac ‘— Red Hills, 250( Ridge west of ' west of Morne 1 North 1 1' . 5' My “Cw (tide HawkstN [hem Linn c m w '/ 258. 18 235 Imshaug 15768, 1953 (MSC); Parish of St. Ann, Moseley Hall Cave, Guys Hill, 200 ft., Imshaug 13656, 1952 (MSC); Walkers Wood to Moneague, 1200 ft., Imshaug 15839, 1953 (MSC); Claremont, 1400 ft., Imshaug 15872, 1953 (MSC); Goshen, 1000 ft., Imshaug 15872, 15989, 1953 (MSC); Parish of Hanover, Lucea, East River, 50 ft., Imshaug 15728, 1953 (MSC); Parish of St. Andrew, Hope River below August Town, 400 ft., Imshaug 1363;, 1952 (MSC); west of Coopers Hill in Red Hills, 2500 ft., Imshaug 14153, 1953 (MSC). TRINIDAD: Ridge west of ”Arima—Blanchisseuse Road," Las Lapas Road, west of Morne Blue, Imshaug 31717, 1963 (MSC). North America. FLORIDA: Everglades National Park, Dade Co., Moore 3805, 1967 (DUKE); Harris 2856B, 1967 (MSC). Central America. COSTA RICA: Irazu, Bethel §.p., 1913 (MICH). Additional material seen: South America. ECUADOR: Galapagos Is., Weber L-40286, L-44035, 1964 (MICH); BRAZIL: Rio de Janeiro, Linderwaldt §.p. (MICH). 15. Ramalina farinacea (L.) Ach. Lichen farinaceus L. Sp. P1. 1146. 1753. Lectotype (fide Hawksworth in Bry010gist 72: 254-255. 1969): Europe (herb. Linn.: Sheet 1273.110) (non vidi). Ramalina farina— Les CL.) Ach. Lich. Univ. 606. 1810. Ramalina subfarinacea Nyl. Bull. Soc. Linn. Norm. 11. 7: 258. 1872. Original material: Europe (non vidi). Ramal: Lich. 34: 847. Cher), sur Er: son 12258 (DUI Ramali Bryol. Lich. 3 (non vidi). Descri to 5 cm. high; ate, subcompr iculate; sora farinose; cor rare, lateral, cave; hypothec thick, paraphy clavate, conta narrow, subfus Medull orange or red; pink; either I< Lichen Salazinic acic Cryptochloroph Discus Variable. In acid and Hess 236 Ramalina hypoprotocetrarica W. L. Culb. Rev. Bryol. Lich. 34: 847. 1966. Holotype: Montou-sur Bievre (Loir et Cher), sur Erables a c6té de la route, Culberson and Culber- son 12258 (DUKE) (non vidi). Ramalina reagens (B. de Lesd.) W. L. Culb. Rev. Bryol. Lich. 34: 847. 1966. Original material: Europe (non vidi). Description: Thallus caespitose, stramineous pale, to S cm. high; branching abundant, branches linear, attenu- ate, subcompressed to compressed, rarely appearing subcanal— iculate; soralia ellipsoid, marginal and terminal, soredia farinose; cortex thick, artilaginous, and rigid. Apothecia rare, lateral, minute, to 1.5 mm. across; disk pale, con— cave; hypothecium 15—20u thick, opagque; hymenium 30-50u thick, paraphyses simple; epithecium 4—6u thick. Asci clavate, containing 8 spores, spores uniseptate, hyaline, narrow, subfusiform, 12-13 x 3—6u. Medullary reactions: Either PD- or PD+ yellow, orange or red; either K- or K+ brick red, yellow red or pink; either KC+ pink or KC—; either C+ pink or C-. Lichen substances: Atranorin, protocetraric acid, salazinic acid, norstictic acid, hypoprotocetraric acid, Cryptochlorophaeic acid and usnic acid. Discussion: The chemistry of the species is very Variable. In Europe Zopf (1907) identified ramalinolic acid and Hess (1958) reported protocetraric acid. French specimens were a new substanc acid (PD-) . A European mater taining either stictic acid a in association recognized fou cetrarica, R. Saxico tain salaziniu mann and Hunec a specimen col also demonstra from Cape Cod, setts, salazin Canary Islands salazinic acid Materi from a mixed c C- or C+ red. protocetraric Cryptochloroph If the the basis of c Same ground ti 237 specimens were shown by C. F. Culberson (1965b) to contain a new substance identified by her as hypoprotocetraric acid (PD—). A mass study by W. L. Culberson (1966) of European material yielded four chemical types: plants con- taining either protocetraric, hypoprotocetraric or nor- stictic acid alone and those containing norstictic acid in association with salazinic acid. Accordingly, he recognized four species: Ramalina farinacea, R. hypoproto— cetrarica, R. subfarinacea and R. reagens. Saxicolous specimens from Israel were found to con- tain salazinic acid alone (Galun and Lavee, 1966), and F011— mann and Huneck (1969) identified protocetraric acid from a specimen collected in Eldorado Co., California. I have also demonstrated hypoprotocetraric acid in a collection from Cape Cod, norstictic acid in specimens from Massachu- setts, salazinic and norstictic acid in material from the Canary Islands. The specimens from Puerto Rico contained salazinic acid. Material from the sourthern hemisphere was studied from a mixed collection from Chile and found to react PD-, C— or C+ red. Those reacting negative with C contain hypo— protocetraric acid and those giving a positive reaction had Cryptochlorophaeic acid in the medulla. If the four species proposed by Culberson (1966) on the basis of chemical constituents are to be valid, on the Same ground the cryptochlorophaeic acid variant reported here could be reason the two stictic acid 3 Species (R. su tograms of the occurred toget that the demon only limited, and by the ori It is that the world five chemical acid, R. farin acid, R. gags farinacea); va cetrarica); an Unlike of the B- orci one is a depsi Althor quently used 2 Flex, it was I tYpified the i aprotocetrarf I hav: the species , : 238 here could be raised to a species by itself. For the same reason the two strains containing either salazinic or nor— stictic acid alone should be revised and reduced to one species (R. subfarinacea=R. reagens). Thin-layer chroma- tograms of these two species indicated that both substances occurred together in each species. Thus it can be concluded that the demonstration of one or the other substance is only limited, at least in this case, by the method employed and by the origin of the material tested. It is evident, according to the foregoing discussion, that the world population of Ramalina farinacea falls under five chemical variants: variant I (containing protocetraric acid, R. farinacea sensus stricto); variant II (salazinic acid, R. reagens); variant III (norstictic acid, R. sub- .—__.. farinacea); variant IV (hypoprotocetraric acid, R. hypoproto- cetrarica); and variant V (cryptochlorophaeic acid, unnamed). Unlike the first four substances which are depsidones of the B- orcinol type (W. L. Culberson, 1966), the last one is a depside of the orcinol type (C. F. Culberson, 1967). Although the chemical substances have been fre— quently used as taxonomic characters in this species com— Plex, it was not until recently that Hawksworth (1969) typified the Linnean material and designated as lectotype a protocetraric acid containing specimen. I have studied all the reported chemical strains of the species, including those from Europe, and found it impossible to. other on that differences a1 they influence has gone thror medullary cons changes. I dc splitting of I species on the tional charact sity and dist] species is muc addition the 6 very clear. In Puc duals of Rama: its compressec margins. All Ecolo; and Huneck (l! phillous, hyg' the bark of t‘ the marine in Oceanic envir In Pu 0f Erythrina 239 impossible to differentiate one chemical variant from the other on the basis of morphological differences. Minute differences are the result of heredity and environment as they influence each individual. Apparently the Species has gone through an evolutionary diversification in the medullary constituents, not paralled by morphological changes. I do not agree with Culberson (1966) in the splitting of R. farinacea (L.) Ach. into four different species on the basis of chemical, ecological and distribu- tional characters. It has been shown here that the diver- sity and distribution of the chemical substances in the species is much wider than shown by W. L. Culberson and in addition the ecological requirements are not altogether very clear. In Puerto Rico the species resembles some indivi- duals of Ramalina dendriscoides but it is differentiated by its compressed branches with a row of soredia along both margins. All Specimens from Puerto Rico were sterile. Ecology: The species has been described by Follman and Huneck (1969, p. 191) as being ”acidophillous, subnitro— phillous, hygrophillous, and photoneutral.” It grows on the bark of trees and on rocks along the coast receiving the marine influence. The species has been reported from oceanic environments. In Puerto Rico R. farinacea occurred on the trunks of Erythrina poeppigiana, a coffee shade tree introduced to the island frc America. Rama peruvi ana and Distri wnes of North inthe tempera dwmical varia Thespecies ap In distributi thesame patte The sp byMfiller (188 Barranquitas 1 Species showed Materi 182 (MSC); [row West I mfltas Hotel, [MCL 16. Ramalina Ramali M2.1870. Descri mllChbranched, Cmllpressed, 5t 240 the island from the lower slopes of the Andes in South America. Ramalina farinacea is associated with Ramalina peruviana and Ramalina dendriscoides. Distribution: This species is common in the boreal zones of North America, from Alaska to Mexico. It occurs in the temperate lowland of Chile in South America. Many chemical variants are frequent in Europe, Asia and Africa. The species appears to be sub-cosmopolitan in distribution. Its distribution, including California and Chile, follows the same pattern as the Ramalina ceruchis complex. The Species was reported from Aibonito, Puerto Rico, by Mfiller (1888) and I collected several specimens in nearby Barranquitas in the central mountains. In the island the species showed a central highland distribution (Figure 21). Material seen: Exsicatti examined, Cum. I. no. 113, 182 (MSC); Howe no. 57 (MSC). West Indies. PUERTO RICO: Monte Cuba near Barran— quitas Hotel, 600 m., Landrén 1926E, 1934, 1942C, 1944, 1967 (use) . 16. Ramalina bistorta Nyl- Ramalina bistorta Nyl. Bull. Soc. Linn. Norm. II. 4: 142. 1870. Description: Thallus stramineous, pale caespitose, much branched, small to 5 cm. high; branches 0.4-2 -m. broad, compressed, subcanaliculate, tips attenuate or terete; cortex thin, ca veins. Apothec subterminal on disk tan, pruir old specimens; colorless, SO-c large numbers ( branched at thc colored to 1411 an 0bliQue uni: appearing $1ng an Oblique p05 1% thalli of R. c which it is di and the produc 241 cortex thin, cartilaginous strands visible as longitudinal veins. Apothecia abundant, to 4.mm. across, lateral and subterminal on short stipes, often appearing subgeniculate; disk tan, pruinose, convex or-concave; margin with spurs in old specimens; hypothecium hyaline, 20-30u thick; hymenium colorless, 50-69u thick, with a gelatinous matrix supporting large numbers of asci and paraphyses, paraphyses much branched at the tips with globose apices; epithecium, brown colored to l4u thick. Asci clavate containing 8 spores in an oblique uniseriate arrangement; spores uniseptate, hyaline, appearing sigmoid but fusiform-oblongae with the septum in an oblique position, 9-16 x 4-6u. Medullary reactions: PD-, K—, KC—, C—. Lichen substances: Caperatic and usnic acids. Discussion: This species can be mistaken for small thalli of R. cumanensis or R. subpellucida from both of which it is distinguished by its typically sigmoid spores, and the production of caperatic acid. Ramalina bistorta appears to be related to R. camptospora by the spore and chemical characters; however, R. camptospora is a sorediate Species which is absent from Puerto Rico. Ecology: The species occurred at elevations between 3000 and 5000 feet in the West Indies, a zone characterized by a montane vegetation which varies from lower montane rainforest to the more dry montane pine forest. In Puerto Rico the species is usually a small corticolous epiphyte on Randia acul teristic of th The species is and Teloschist dry mountain 5 Distri occurs in Cuba skipping Hispe The di iSrestricted the Cordillerg The di 242 on Randia aculeata and Guettarda scabra, two trees charac- teristic of the montane scrub vegetation above 2500 feet. The species is commonly associated with species of Usnea and Teloschistes flavicans, occupying open habitats on the dry mountain summits of the west end of the central range. Distribution: In the West Indies Ramalina bistorta occurs in Cuba, Jamaica, Puerto Rico, and St. Thomas, skipping Hispaniola and the Lesser Antilles. The distribution of R. bistorta within Puerto Rico is restricted to the highland of the western mountains in the Cordillera Central near Maricao and Utuado (Figure 22). The disjunct Antillean Californian distribution of this species is difficult to explain, since there are no floristic affinities between these two regions. The species has not been reported from outside the original collection site and it is not lested in the North American lichen checklist (Hale and Culberson, 1970). It is expected that if it were a California species it would follow the typical California—Chile distribution of the Desmazieria group of species. However, instead it is common in the Greater Antilles. This points to the West Indies as the probably center of dispersion with some accidental intrusion into California. The possibility remains that perhaps the type was mislabelled. Material seen: West Indies. PUERTO RICO: Upper Slopes of Mt. Morales near Utuado, Howe 1146, 1906 (MICH); montane scrub 1 w, 1967 am Cahterine's Per 1952 (MSC); gag Blue Mountains Mount, on nort] Imshaug 1359911 3600 ft., Blue Slope of Mos sm: Parish of St. 1 243 montane scrub zone on Monte del Estado at Maricao, Landrén 1536B, 1967 and 2421, 1968 (MSC). JAMAICA: Summit of Cahterine's Peak, 5000 ft., Blue Mountains, Imshaug 13323, 1952 (MSC); gap southeast of Catherine's Peak, 4500 ft., Blue Mountains, Imshaug 13880, 1952 (MSC); near Clifton Mount, on northeast slope of Catherin's Peak, 4400 ft., Imshaug 13599A, 1952 (MSC); ridge northwest of Murdocks Gap, 3600 ft., Blue Mountains, Imshaug 15340, 1953 (MSC); south slope of Mossman's Peak, 5000 ft., Imshaug 11959, 1953 (MSC); Parish of St. Andrew, Haberstadt to Bloxburgh, 2800 ft., Imshaug RRRRE, 1953 (MSC); St. Thomas, Farm Hill, Orcutt 3366, Rééi, 5392, 1927 and 1928 (US); Chestervale, Plitt Cummings §.p., 1905 (PH); Hart §.p. (FH). CUBA: Loma del l7. Ramalina cumanensis Fée Ramalina cumanensis Fée, Essai Cryptog. Ecorc. Officin. 135. 1824. Original material: Peru, Hfib. and Eggpl. (non vidi). Ramalina ecklonii Spreng. in L. Syst. Veget. ed. 16. 4(2): 328. 1827. Original material: ”In sylvis ditionis Vitenhagen Afric. austr. Ecklon.” Ramalina linearis f. spinulosa Merr. Bry010gist ll: 50. 1808. Lectotype (nov.): Jamaica, Cummings §.p. (MICH). Descri ladniae linea madnng 20 cm flattened in l hodwcia nume wncave, flat tuning 8 spor hynine, ellip Medull ‘ Lichen 244 Description:. Thallus caespitose to pendent; laciniae linear, usually about 5 cm. but occasionally reaching 20 cm. long, dichotomous, canaliculate, becoming flattened in large specimens, 2-4 mm. wide; esorediate. Apothecia numerous, minute, marginal and laminal; disk concave, flat or convex, tan, pruinose. Asci clavate, con- taining 8 spores, arranged in two series, spores uniseptate, hyaline, ellipsoid, 11—18 x 4.5-8.5u. Medullary reactions: PD-, K-, KC—, C-. Lichen substances: Usnic acid. Discussion: The variability of the species is in accordance with its wide range and diverse habitats. The thallus is either shrubby and canaliculate or pendulous and broad. The dispoisition of the apothecia is either dorsal or ventral on the broad laciniae, but more or less lateral along the canals in the canaliculate ones. The name R. Ecklonii is almost always associated in the liter— ature with short, flattened specimens. While studying this species I was able to examine speciems from Jamaica composed of two types of laciniae, thin, canaliculate as in R. cumanensis and broad and flat as in R. ecklonii. In several pendent specimens it is possible to find that the long linear laciniae are thin and canaliculate toward the base, while the same filament flattens and acquires a lanceolate form toward the apex. The fact that l- M61151; and that in J: which was int] reinforces the to R. cumanens Ecolog habitats and t corticolous s; gTOWing on roc rainforest 201: trees. Distri \ aPpears to be northern and S SPeties occurs 245 R. cumanensis was described from the Cinchona tree in Peru and that in Jamaica the species is abundant on this tree, which was introduced to the island from Soutthmerica, reinforces the idea that the West Indian material belongs to R. cumanensis. Ecology: The species has been reported from various habitats and the most diverse environments. Is is mainly a corticolous species;in a few cases it has been reported growing on rocks. In the West Indies it is frequent in the rainforest zone, and especially on the trunks of Cinchona trees. Distribution: The distribution of this species appears to be tropical and temperate throughout both the northern and southern hemispheres. In the West Indies the species occurs in Jamaica, Haiti and Dominican Republic. Material seen: West Indies. JAMAICA: Cinchona, ca. 1500 m., Maxon 842, 1820 (EH); 4500 ft., Plitt g6_, 1919 (US); Plitt 5.3., July 10, 1919 (US); Plitt 3.2., July 21, 1926 (US); Chestervale, ca. 3290 ft., Plitt l, 12, 1932 (EH); Flamstead and Vicinity, Port Royal, mountains ca. 1000 ft. Maxon 8664, 1926 (PH); NeWcastle, Cushman 125, 1912 (PH); Clydesdale, Plitt 3.2., July 19, 1926 (US); St. Andrew, Bellevue, on tree trunk ca. 3750 ft., Amy van der Porter §.p,, 1950 (MICH); Cummings 181, 1905 (MICH); Cinchona, 5000 ft., Earle 379, 1902 (MSC); St. Andrew, Guava ridge, 2900 ft., Imshaug 14346, 1953 (MSC); Bellevue, 3800 ft., Imshaug 14460, 1953 (MSC). H} Leonard 4428, f fields at side l'Ouest, summit Imshaug 23082, Allaceyes’ Cord: 3E, 1958 (MS( 18. ~ ; m: Ramalir \ 1888' Originaj [9pm. (11011 v; 246 1953 (MSC). HAITI: Vicinity of Furcy on rocks on cliff, Leonard 4428, 4444, 1920 (PH); in wood ravines and open fields at side of paths, Thomas 107 (PH); Department de l‘Ouest, summit of Téte Etang, between Kenscoff and Furcy, Imshaug 22570, 22582, 1958 (MSC); summit of Montagne Noir, Wetmore 2743, 2680, 1958 (MSC); west of Forét des Pins, Imshaug 23082, 22506, 1958 (MSC). DOMINICAN REPUBLIC: Los Amaceyes, Cordillera Septentrional, Wetmore 3402, 3407, 3413, 1958 (MSC); Cerrazos, between La Cumbre to Santiago, Wetmore 3829, 3863, 1958 (MSC). 18. Ramalina subpellucida Mall. Arg. Ramalina subpellucida Mfill. Arg. Flora 71: 492. 1888. Original material: Coamo, Puerto Rico; Sintenis QR p.p. (non vidi). Description: Thallus pale stramineous, 1-3 cm. high, shrubby; laciniae compressed, sometimes subcanalicu- late, attenuate with tips subterete; surface longitudinally striate; cortex thin. Apothecia numerous, 3-4 mm. broad, terminal or sub-terminal, if subterminal then geniculate, but appearing terminal if the branch tips are broken; hy- menium very gelatinous, 40—60u thick with profusion of branched paraphyses with capitate apices. Asci clavate with 8 spores arranged in two groups of four, the upper four overlapping the lower ones; spores uniseptate, hyaline, fusiform, Straight or substraight, 18—22 x 3-5u. Medul Liche mnic acid. Discu wnflued with However, it c pmillate and Du variety R ther (1888) lucida. 1 ha htermined by itis only a variety may V wasterete or 247 Medullary reactions: PD-, K—, C-, KC-. Lichen substances: Atranorin, divaricatic acid and usnic acid. Discussion: In the West Indies this species can be confused with small specimens of R. montagnei or R. bistorta. However, it can be separated from the former by not being papillate and from the latter by not having bistorted spores. The variety R. subpellucida var. tuberculata described by Mfiller (1888) was part of the same collection as R. subpel- lucida. I have seen one specimen from the type locality determined by Riddle as v. tuberculata, but in my opinion it is only a small (1 cm.) R. montagnei. The tuberculate variety may very well be R. montagnei De Not. if indeed it was terete or subterete and tuberculate. Since this last species was described from Cuba and the Florida population may have divaricatic acid, it is expected that the species be also present in Puerto Rico. However, I did not collect the plants in mixed condition with the papillate divaricatic acid producing variant of Ramalina complanata, which could pass for R. subpellucida var. tuberculata. The Species R. subpellucida is easily distinguished from R. bistorta on account of its fusiform spores, straight septum and different chemistry; it produces divaricatic acid, while R. bistorta produces caperatic acid. R. subpellucida reacted PD-, K-, C-, KC—. It con- tained divaricatic acid alone as medullary constituent and usnic acid in the . 13523) was found tr substance. Rarely The specie: Rmaflrp stenospor. group, judging by ‘ the species appear: with the above fir: entiated from R IE not being tubercul. divaricatic instear Ecology: land and lower mon is between 70—100 was described from 248 usnic acid in the cortex. One specimen from Jamaica (Imshaug 13523) was found to contain only usnic acid with no medullary substance. Rarely atranorin is present. The species appears to be related to the stock of Ramalina stenospora, R. montagnei and the Ramalina usnea group, judging by the spore characters. In Florida, where the species appears to be frequent, it may be confused with the above first two mentioned species, but it is differ- entiated from R. montagnei by its flattened branches and by not being tuberculate, and from R. stenospora by producing divaricatic instead of perlatolic or stenosporic acid. Ecology: Ramalina subpellucida is found in the low— land and lower montane rainforest sites, where precipitation is between 70-100 inches a year. The fact that the species was described from the drier southern plain could be explained by the presence of a nearby lagoon, which has now disappeared. This corticolous species was found to be abundant in spots within the palm savanna facie of the white sand second growth vegetation. Almost all the collections were made on dead branches of the orchard tree Anacardium occi~ dentale. The only other lichen growing in association with this species in this area was Ramalina complanata (proto- cetraric acid variant). In the western part of the island the species was collected on a low scrub of Chrysobalanus icaco, exposed to strong sea wind. The largest plants of R. subpellucida were those collec elevations betwee: trunk of Tabebuia association with ‘ i- laria and 3. a Distribut Greater Antilles Lesser Antilles. Islands, thus coV floristic provinc Puerto Rico, but species was Origi: but i have Seen 5 [1915] TePOIted f r. . _ ‘ W Whic L0 13. montagnei 249 were those collected in the central part of the island at elevations between 500 and 800 meters, epiphytic on the trunk of Tabebuia heterophylla. Here the species grew in association with R. complanata (divaricatic acid variant), R. usnea and R. gracilis. Distribution: This species is typical of the Greater Antilles and Guadaloupe and Grand Cayman in the Lesser Antilles. It occurs in Florida and in The Bahama Islands, thus covering the Antillean—southern North American floristic province. Ramalina subpellucida has a wide distribution in Puerto Rico, but locally it is not very abundant. The species was originally reported from the southern coast, but I have seen specimens from all over the island. Riddle (1915) reported from Mona Island a specimen identified as R. montagnei which is much closer to R. subpellucida than to R. montagnei. In general, the species exhibited a scattered distribution pattern (Figure 23). Material seen: West Indies. PUERTO RICO: Vega Baja, Laguna Tortuguero, sand dunes with scattered palms ___—__._____.__ 530, 534, 536, 544, 555B, 556, 558, 566C, 587, 1967 (MSC); old pasture field, south of lake, Landron 613B, 614, 618A, ERR, 620B, 636, 646, 652B, 663A, 664, 669, 697, 1967 (MSC); Barrio Cuchillas, 600 m., Orocovis, Landrdn 1758A, 1765, 1967 (MSC); El Negro, Montafias de Corozal, Landron 1779C, ‘ 1781A, 1783C, 196 Corozal, Landron Lago Carite, Guay @2, 1967 (MSC); 1967 (MSC); Barri Lingerie 2&4 196 1914 (US); Mona I “6313, 1914 ( covered ridge jug Alturas de P1 2 arr 250 1781A, 1783C, 1967 (MSC); Barrio Padin near Orocovis and Corozal, Landrén 1793C, 1796C, 1799C, 1800C, 1967 (MSC); Lago Carite, Guayama, lower montane rainforest, Landrdn 1882, 1967 (MSC); Monte Llano, near Cidra, Landron 1955B, 1967 (MSC); Barrio Canaboncito, 200 m., Caguas, Landron 2457, 2458, 2459, 1968 (MSC); Vega Alta, Barrio Maricao, Landrdn 2286, 1968 (MSC); Barrio Espinosa, Stevenson 2352, 1914 (US); Mona Island, Sardinera, Britton, Cobwell and Hess 1724, 1914 (MICH). CUBA: Prov. Pinar del Rio, oak covered ridge just southwest of Hotel San Vicente, 650 ft., Alturas de Pizarras, Imshaug 25229, 25262, 1959 (MSC); San José de Las Sejas, Hubbard p.p., 1906 (MICH); Wright RE (FH). GRAND CAYMAN: West Bay, between Galleon Beach and West Bay Town, Imshaug 24565, 1959 (MSC); ruinate area with scattered thatch palm and maiden plum near Georgetown, Imshaug 24390B, 1969 (MSC); logwood trees in woods near Hell Hole, West Town, Imshaug 24475, 1959 (MSC). JAMAICA: Parish of St. Andrew, south slope of Cooper's Hill in Red Hills, 2250 ft., Imshaug 13725, 1952 (MSC); between airport and Port Royal, Imshaug 13523, 1952 (MSC); Old Harbor, Orcutt 2522, 1927 (US). BAHAMAS: Gillis 5271A, 1963 (MSC). GUADELOUPE: Petit Canal at Pot Louis, Duss 549, 1902 (MICH). Central America. YUCATAN: Chichankanab, Gaumer 2288B (MICH). North America. FLORIDA: Everglades, Harris 2385B, 2948A, 1967 (MSC). " 19. W ance Ramalina Lectotype (nov.): Descripti 45 cm. long branc compressed, angul and terete; surfa minute, to 1.5 mm disk tan, concave To”; SPores unise SUbstraight, 12-2 251 19. Ramalina anceps Nyl. Ramalina anceps Nyl. Syn. Lich. 1: 291. 1860. Lectotype (nov.): Guadeloupe, Wright §.p. (FHT). Description: Thallus pendulous, stramineous, to 45 cm. long branching irregularly dichotomous; branches sub- compressed, angular and two edged, terminal branches thin and terete; surface smooth, striate. Apothecia rare, minute, to 1.5 mm. broad, lateral and adnate to the thallus; disk tan, concave. Asci containing 8 spores in a single row; spores uniseptate, hyaline, ellipsoid, straight or substraight, 12-22 x 6-8u. Medullary reactions: PD+ yellow, K+ red, KC—, C-. Lichen substances: Atranorin, norstictic acid, traces of salazinic acid and usnic acid. Discussion: Nylander (1858-1860) noted the alec— torian aspect of the species. Howe (1914) interpreted R. anceps as a chemical species, probably a strain of R. usnea, from which it differed by having a K+ reaction, while R. usnea was K—. It is my opinion that R. anceps is a distinct species which not only shows chemical differences but also spore differences. Ramalina anceps has ellipsoid spores 12—22 x 6—8u in size, while R. anceps has fusiform spores 16—22 x 3-5u in size. Chemically, R. anceps is very homogeneous, containing norstictic acid, while R. usnea is chemically variable, with at least four chemical variants recognized in the West Indies. of Luquillo Mount racemiflora and h in the east side is above 150 inch along the edge of mountains R. ance ‘ M, which are central elastic c In Sierra between 500 and 7 the palm forest. on branches of Da M 512- and T ated With R. ance In the ce on BuChe ~ na Q can drier western mou SpeC1es Q 252 Ecology: The species occurred on the exposed slopes of Luquillo Mountains, hanging from branches of Cyrilla racemiflora and Microphollis garciniaefolia. The area lies in the east side of the mountains where rainfall sometimes is above 150 inches a year. It is more abundant on trees along the edge of the forest or isolated Spots. In these mountains R. anceps is always associated with species of Usnea, which are easily distinguished by the presence of a central elastic core which is absent in Ramalina. In Sierra de Carite the Species is most frequent between 500 and 700 meters of elevation, without invading the palm forest. In this part of the island it is epiphytic on branches of Dacryodes excelsa, Ficus laevigata, Micro— pholis Spp. and Tabebuia heterophylla. The lichens associ— ated with R. anceps are R. peranceps, R. dendroides, R. usnea, and Teloschistes flavicans, all of which are pendant and corticolous. In the central mountains the species is collected on Buchenavia capitata and Calophxllum brasilense. On the drier western mountains it is epiphytic on many tree species of Clusia. The associated lichen flora is consti- tuted by the same species throughout the range, although the dominant Species vary. Ramalina anceps was so abundant in former times that it was locally used to stuff mattresses. Distribution: The species was described by A Nylander from Gua West Indies (Imsh is probably the 11 Caribbean Islands Greater Antilles. islands of the Le not reach Florida Within ti widely distribute and montane zones LUHUillo. followi 253 Nylander from Guadeloupe and had been reported from the West Indies (Imshaug, 1957) and Brazil (Vainio, 1890). It is probably the most widely distributed species in the Caribbean Islands. I have seen material only from Haiti in the Greater Antilles. Although it is very common in most islands of the Lesser Antilles and Cuba, the Species does not reach Florida as many other species do. Within the island of Puerto Rico the species is widely distributed throughout the lower montane rainforest and montane zones, extending from the eastern mountains at Luquillo, following a central course to the western end of the island. The species has not been collected in the drier northern and southern coasts. Ramalina anceps has an east—west distribution (Figure 24). Material seen: West Indies. PUERTO RICO: Luquillo Mountains route to Naguabo, 800 m., Landron 1174A, 1176B, 1177, 1182, 1183A, 1184, 1185B, 1186, 1187, 1191, 1967 (MSC); south of Mt. Britton on route P.R. 191, Imshaug 29564, 29567, 29569, 1963 (MSC); Cayey, Bosque Estatal de Guavate, near Park, Landron 1417A, 1426, 1435, 1436A, 1967 (MSC); Cerro Farallon, Guavate, route to Carite, 600 m., Landron _.__.__——.__—-————_—— Carite lake, 600 m., Landrdn 1488, 1489A, 1490A, 1491A, 1495A, 1967 (MSC); coffee plantation, north of Carite lake, Cerro Farallon, trail to Carite lake, Landron 1649A, 1654, 1gp, 1699B, 16_7 833 m., route to lSlIR, 1518, 1527 of observation to ‘ 154i, 1551A, EE west of observati 510136 east of rad 1% 1967 (MSC); @1614. E Landron 2415A, 24 Cerro Doha Juana, 1%: 1679. 1681 1M. 1967 (MSC) “Mn 1819B 5 \s 1967 (MSC); Open 23% M, 233 5 254 1655C, 1699B, 1671A, 1967 (MSC); Maricao, Monte del Estado, 833 m., route to Maricao township, Landr6n 1510, 1512, 1513, _-____—__——_._.._—__ ——_—.———.——.—-— 1549A, 1551A, 1554A, 1559B, 1570A, 1571A, 1967 (MSC); slope west of observation tower, Landron 1586A, 1594, 1967 (MSC); slope east of radio antennas, Landron 1602C, 1607, 1624B, 1626, 1967 (MSC); Slope west of radio antennas, Landr6n 1632B, 1634, 1635, 1636, 1967 (MSC); near Stone House, Landron 2415A, 2424, 1968 (MSC); Bosque Estatal, Toro Negro, Cerro Dona Juana, rainforest, Landr6n 1674B, 1675A, 1676, 1678A, 1679, 1681, 1682B, 1683, 1685A, 1689C, 1694, 1695A, 1696A, 1967 (MSC); pasture field with scattered tress, —.—._———._____-—._——__—._—— 1967 (MSC); open field, 800 m., Landron 2310A, 2314, 2322 2323A, 2330A, 2335A, 2346, 2347A, 1968 (MSC); Divisoria, between Villalba and Orocovis, Landrén 1733C, 1967 (MSC); Villalba, Barrio Caonillas, 800 m., Landron 1704A, 1707A, 1717C, 1726, 1727B, 1967 (MSC); Barranquitas, Monte Cuba, near Hotel Barranquitas, Landron 1890B, 1912B, 1917D, 1922, 1933A, 1967 (MSC); Bro.Hioram 3.3., 1911 (MICH); Lago el Guineo, route to Cerro de Punta, Landr6n 2352B, 2354D, 2355, 2360, 1968 (MSC); Aibonito, Fink 1970, 1916 (MICH). ‘ CUBA: Monte Verde, Wright 735 (FH). DOMINICAN REPUBLIC: Cerrazo, Cordillera Septentrional, Wetmore 3832, 1958 (MSC); JAMAICA: Parish of St. Ann, Bamboo, 2000 ft., Imshaug 1_59_76, 1953 (MSC) Rosanna, 3800 ft. Mountains, ridge w, 1952 (MSC) Dick's Pond Trail (MSC); Chesterval Emailing i-Ew 19 Blanchisseuse Roe ca. 2000 ft., E Naranja (above g2 of St. Joseph, Nc 3T. LUCIA: Quart m 297i: 18 ridge, 2000-2400 montane thicket , 255 15976, 1953 (MSC); Parish of St. Andrew, Bellevue to Mt. Rosanna, 3800 ft., Imshaug 14457, 14497, 1953 (MSC); Blue Mountains, ridge of Mount Horeb, 4400—4600 ft., Imshaug 13159, 1952 (MSC); open reforestation region, 3700 ft., Dick's Pond Trail near Hardwar Gap, Imshaug 13118, 1952 (MSC); Chestervale, Plitt p.p. (US); no locality given, Cummings 5.3., 1904 (FH). TRINIDAD: Ridge west of ”Arima- Blanchisseuse Road“ along Las Lapas Road, west of Morne Blue, ca. 2000 ft., Imshaug 31723, 1963 (MSC); north slope of Naranja (above gap between Naranja and El Tucuche) north of St. Joseph, Northern Range, Imshaug RRRRQ, 1963 (MSC). ST. LUCIA: Quartier Sufriere, Mt. Casteau, ca. 2000 ft., Imshaug RRZRZ, 1963 (MSC); montane thicket, Mt. Tabac ridge, 2000-2400 ft., Imshaug 30270, 30298, 1963 (MSC); montane thicket, 1800—2000 ft., south of Piton Canarie (above Migny), Imshaug 29939, 1963 (MSC); secondary rain— forest, Des Bottes, ca. 2000 ft., north of Soufriere, 30082, 30086, 30103, 1963 (MSC); elfinwoodland type modi- fication of semi—evergreen seasonal forest, summit area of Gros Piton, 2300—2619 ft., Imshaug 30221, 1963 (MSC). MARTINIQUE: Route de la Trace from Deux Choux to Belvéder du Piton Gelé, Imshaug 32645B, 1963 (MSC). GRENADA: Grand Etang, Broadway p.p. (MSC). DOMINICA: Mossy forest, trail to summit of Morne Anglais, ca. 3600 ft., Hale 35431, 1969 (US). 20- WEE Ramalina Lectotype (nov.) : (FHl). Ramalina 71:492. 1888. 19’ p p r 2_0 p p Ramalina 34 32 1896 c 256 20. Ramalina peranceps Nyl. Ramalina peranceps Nyl. Flora 59: 411. 1876. Lectotype (nov.): Cuba, Wright RRR, Nov. 14, no year given (FHl). Ramalina sintenisii var. polyclada Mfill. Arg. Flora 71: 492. 1888. Original material: Puerto Rico, Sintenis RR, p.p., Rg_p.p., RR p.p. (non vidi). Ramalina peranceps f. leptoptelos Vain. Journ. Bot. 34: 32. 1896. Original material: St. Vincent, Elliot 5.2. (non vidi). Description: Thallus stramineous, pendulous, to 12 cm. long, branching dichotomous; laciniae frequently with short straight lateral branches at right angles, flattened, to 3 mm. broad, attenuate with tips often subterete; sur— face striate; esorediate; cortex cartilaginous. Apothecia rare, small, to 2 mm. across, adnate, convex. Asci con— taining 8 spores arranged in a Single row, spores uniseptate, hyaline, ellipsoid to subfusiform, straight or slightly bent, 15-21 x 4-6u. Medullary reactions: PD+ yellow-orange, K+ yellow— red, KC—, C-. Lichen substances: Atranorin, salazinic acid, traces of norstictic acid and usnic acid. Discussion: Nylander (1876) described the species with a morphology similar to Ramalina anceps and with a similar K+ yellow—red reaction. Although the color reaction .‘7 appears to be the that causes the C acid in the medul The speci gical characters. bifacial thallus lateral branchlet Ramalina —‘ from which it dif treated as a sync to the same stock having broader, 1 Salazinic acid. ”1“ng from sea al’ear, the Spec T . W and the 257 appears to be the same, it differs in the chemical substance that causes the color. While R. anceps contains norstictic acid in the medulla, R. peranceps contains salazinic acid. The species also differ in their general morpholo- gical characters. Unlike R. anceps, R. peranceps has a bifacial thallus which often is characterized by short lateral branchlets at right angles to the main branches. Ramalina peranceps may be confused with R. dendroides, from which it differs by the absence of lateral soralia. The lichen R. sintenisii var. polyclada Mfill. Arg. seems to be the same as R. peranceps Nyl., and it is here treated as a synonym of that species. R. peranceps belongs to the same stock of R. usnea, being distinguished by having broader, rigid branches, broader spores and typically salazinic acid. Ecology: Ramalina peranceps occurred at elevations ranging from sea level to 3000 feet in the West Indies. In Puerto Rico the species is found in the wet eastern slopes of the Luquillo Mountains and at higher ele- vations it is replaced by Usnea spp. In other places where rainfall is not so abundant, but still near a hundred inches a year, the Species grows abundantly on the trunks of Tabebuia and the palm Roystonea. In the western mountains where rainfall is much less and markedly seasonal, the species grows epiphytic on low branches of Ficus, often hanging from rocks. The rock habitat, however, species. Distribut West Indian distr paniola in the Gr in almost all the reaches as far so The speci dillera Central 3 the east, but it coaStal Plains. east-west highlan slopes of El Yunq NagUabo, Ca. 500 Bosque EStatal dc Cayey, ca, 500 m. M) 1313, 133 258 habitat, however, is not considered characteristic of the species. Distribution: Ramalina peranceps has a widespread West Indian distribution. The species is absent from His- paniola in the Greater Antilles, but it is well represented in almost all the islands of the Lesser Antilles and reaches as far south as Trinidad and Tobago. The species in Puerto Rico is frequent in the Cor- dillera Central and extends to the Luquillo Mountains in the east, but it is absent from the sourthern and northern coastal plains. Thus, the species was found to exhibit an east-west highland distribution [Figure 25). Material seen: West Indies. PUERTO RICO: East slopes of E1 Yunquein.a caimitillo thicket, route to Naguabo, ca. 500 m., Landron 1182B, 1185A, 1967 (MSC); Bosque Estatal de Guavate, Tabebuia forest near Park Area, ——_—_—_—————————————__—_ to Carite Lake, ca. 750 m., Landron 1455, 1967 (MSC); caimitillo palm forest in midway between Guavate-Carite, route to Carite Lake, ca. 800 m., Landron 1495B, 1497, 1967 (MSC); Monte del Estado, Cordillera Central, Maricao, 833 m., Landron 1534A, 1530, 1539, 1967 (MSC); observation tower, Monte del Estado, Maricao, Landron 1544, 1570B, 1580B, 1583A, 1587, 1588, 1591, 1593B, 1967 (MSC); antenas de radio, Monte C scrub facing 8351 M, w, 196'. House, route to l Cerro Dofia Juana Bosque Toro Negn (MSC); Cordiller; Negro in pasture Carite in coffee (MSC); Monte Lla “Mi M. 1 ca. 500 111., Land ,. 1\Qfl7flandh Vleux Fort, mont 259 de radio, Monte del Estado, steep slope covered by spring scrub facing east, ca. 833 m., Landr6n 1628A, 1633B, 1638, 1639A, 1640, 1967 (MSC); Monte del Estado, 5 km. from Stone House, route to Maricao, Landrén 2416, 2420, 1968 (MSC); Cerro Dofia Juana, betweenlkrestry Building and Rec. Area, Bosque Toro Negro, 850 m., Orocovis, Landron 1707B, 1967 (MSC); Cordillera Central, Divisoria, Villalba, 800 m., Landrén 1747, 1967 (MSC); Cerro Dofia Juana, Bosque Toro Negro in pasture with scattered Buchenavia, ca. 850 m., Landrén 1830A, 1843A, 1967 and 2349B, 1968 (MSC); Lago Carite in coffee plantation, ca. 600 m., Landrén 1880, 1967 (MSC); Monte Llano, 7 km. north of Cidra, ca. 500 m., Landrén 1930C, 1967 (MSC); Cuyon, Aibonito, on guava scrub, ca. 500 m., Landr6n 2005B, 1967 (MSC); Lago e1 Guineo, S.E. shore of lake, Landron 2357, 1968 (MSC); Cerro de Punta, about 200 m. below summit in palm-fern forest, Landron 2400A, 1968 (MSC); Luquillo Mountains, south of Mt. Britton, Route 191, Imshaug 29579, 1963 (MSC). CUBA: Monte Libano, Wright 736 and Wright §°E' (FH). ST. LUCIA: Quartier of Vieux Fort, montane thicket, Mount Grand Magazin, 1800—2000 ft., Imshaug 30021, 30029, 1963 (MSC); montane thicket, 2000 ft., ridge at head of Ravine Marche Gaye, Imshaug 30263, 1963 (MSC); Quartier Soufriére, road south of Piton Canarie (above Migny), 1800—2000 ft., Imshaug 29925, 29749, 29768, 1963 (MSC); montane thicket, Morne Bonin, 1700-2000 ft., Imshaug 29768, 29910, 29917, 1963 (MSC); Mt. Casteau, ca. 2000 ft. , ImShaug between Mount Bel Imshaug M’ 19 near shore of GTE (MSC). sT. VINCE Charlotte, MontrE 304i, 1963 (MSC) summit of Main Ri LL; m. 19 1500 ft., north 5 "Roxborough-Bloo< MARTINIQUE: Rout du Piton Gelé, R from summit of M( Range, between A: 018C) ; south $10] Lalaja-Paria Tra: 260 2000 ft., Imshaug 29813, 1963 (MSC); Quartier Laborie, between Mount Belvéder and Mound Grand Magazin, 1500 ft., Imshaug 30049, 1963 (MSC). GRANADA: Parish of St. George, near shore of Grand fitang, 1500 ft., Imshaug 16093, 1953 (MSC). ST. VINCENT: Cultivated area, 1500 ft., Parish Charlotte, Montreal near Mesopotamia, Imshaug 30816, 30831, 30434, 1963 (MSC). TOBAGO: Lower montane rainforest, summit of Main Ridge, near Parlatuvier, 1600-1700 ft., Imshaug 3&162, 1963 (MSC); lower montane rainforest, ca. 1500 ft., north slope of Main Ridge at about 8 miles on old ”Roxborough-Bloody Mule Road,” Imshaug 3R663, 1963 (MSC); MARTINIQUE: Route to la Trace from Deux Choux to Belvéder du Piton Gelé, Imshaug 3266, 1963 (MSC). TRINIDAD: Ridge from summit of Morne Blue to ”Lalaja-Paria Trail” Northern Range, between Arima and Blanchisseuse, Imshaug 37865, 1963 (MSC); south slope of northern range, north of Arima, along Lalaja—Paria Trail, Imshaug RRRRR, 1963 (MSC); north slope of Naranja (above gap between Naranja and Tucuche), north of St. Joseph, Northern Range, Imshaug glRRZ, 1963 (MSC); Cruger s.n., (FH). JAMAICA: Parish of Hanover, Birchs Hill, 1809 ft., Imshaug 15714, 1953 (MSC); ridge southwest of Dolphin Head, 1750 ft., Imshaug 15624, 1953 (MSC); Blue Mountains, forest hut, above Curn Puss Gap, 3550 ft., Im- shaug 15624, 1952 (MSC). DOMINICA: Parish of St. Luke, cultivated area, 1500—1700 ft., south Chiltern Estate Road, Imshaug 32680, 32701, 1963 (MSC); Parishes of St. Luke and St. Mark, foreste Nome Plat Pays, 21. Ramal ina E Ramalina Norm. II. 4: 112, 26:412. 1876. [non vidi). LeCtOthe (nov .) : Descriptj \ 30 cm. long, brar 261 St. Mark, forested area 2200—3100 ft., Soufriére Ridge, Morne Plat Pays, Imshaug 33059, 1963 (MSC). 21. Ramalina dendroides Nyl. Ramalina rigida f. dendroides Nyl. Bull. Soc. Linn. Norm. II. 4: 112, 1870. Ramalina dendroides (Nyl.) Nyl. Flora 26: 412. 1876. Original material: Martinique, Husnot 46R (non vidi). Ramalina sintenisii Mfill. Arg. Flora 71: 491. 1888. Lectotype (nov.): Adjuntas, Puerto Rico, Sintenis 14 (G!). Description: Thallus stramineous, pendulous, to 30 cm. long, branching dichotomous toward the base, and dendroid or with bushy small branchlets at the apices; laciniae linear, compressed bifacial, varying from 0.2 to 2 mm. wide; surface striate, often split along the margins with numerous soredioid granules protruding, with white tubercules along the margins or on short lateral branches bursting into soredia; soralia not characterically present; cortex thick, rigid. Apothecia frequent, marginal, small, to 3 mm. across; disk concave or convex. Asci containing 8 spores arranged in a single row; spores uniseptate, hyaline, fusiform—ellipsoid, either straight or sub-straight, 12-19 x 4-6u. Medullary reactions: PD+ yellow or PD—, K+ yellow- orange or K-, KC—, C-. Lichen 51 catic acid and u: dendroides from i on a specimen mi The formal diagn in 1876, while (1 to R. dendroides \ sorediate. In 1888 w from Pue marginally tube] material for thj In the same gene Nas C01lected a1 held fTOm Rama the OTmer 1t 1 262 Lichen substances: Atranorin, salazinic acid, divari— catic acid and usnic acid. Discussion: Nylander described Ramalina rigida f. dendroides from Martinique in 1870. This taxon was based on a specimen misidentified as R. complanata (Husnot 469). The formal diagnosis was "sorediform tuberculate.” Later, in 1876, while describing Ramalina dendriscoides, he referred to R. dendroides as being compressed and not being terminally sorediate. In 1888 Muller described the species Ramalina sig— tenisii from Puerto Rico, which was described as being marginally tuberculate. Several numbers were cited as type material for this species but none was selected as the type. In the same general area Ramalina sintenisii var. polyclada was collected and described. Again several numbers were cited but not type was selected. I have been able to study Sintenis 14, a syntype of Ramalina sintenisii and have selected it as lectotype. Additional material studied from Puerto Rico more nearly approaches Nylander‘s interpretation Of R. dendroides than Muller's concept of R. sintenisii. It is my belief that both species represent differ- ent aspects of a variable population and the name R. den— __ droides has priority over R. sintenisii. This species may be difficult to separate in the field from Ramalina peranceps and Ramalina anceps. From the former it is distinguished by the presence of marginal soralia and from acid, which is al I inclut ical populations other PD-. The zinic acid in th gave an unidenti which fluoresce reacting specime belong to a diff “5111C acid in th m: West Indies in t island of Puerto RElma ' . & assoc ation Wit} USHEa Dist ‘ 263 soralia and from the latter by the presence of salazinic acid, which is absent in R. anceps. I include within this species two different chem— ical p0pulations, one of which reacts PD+ yellow and the other PD-. The positive reacting population contains sala— zinic acid in the medulla but a few specimens in addition gave an unidentified PD- spot on thin-layer chromatograms which fluoresce blue under UV (Substance D). The PD- reacting specimens containing divaricatic acid might well belong to a different species. All the specimens contained usnic acid in the cortex but only rarely atranorin. Ecology: This corticolous species was found in the West Indies in the zone between 800 and 3000 feet. In the island of Puerto Rico the species occurred in the lower mon- tane and montane rainforest zones, between 1000 and 2800 feet. Characteristically it hung from the trunks and branches of Tabebuia heterophylla, I. rigida, Roystonea boringuefia and from the branches of Ficus laevigata, Microphollis gar- ciniaefolia and Cyrilla racemiflora. It was not uncommon to collect specimens growing on calcareous rocks, especially in the drier western mountain range. Ramalina dendroides was frequently found growing in association with R. peranceps, R. anceps, Teloschistes and Usnea. Distribution: Ramalina dendroides is an endemic species to the West Indies with a wide distribution within the islands of t the Lesser Anti] Trinidad and T01: ulation of the s divaricatic acid Tobago and Jamai variants of the SPecies showed a 264 the islands of the Greater Antilles except Hispaniola. In the Lesser Antilles the species occurred in St. Lucia, Trinidad and Tobago. While the salazinic containing pop- ulation of the species is spread throughout the islands, the divaricatic acid producing one is confined to Trinidad, Tobago and Jamaica, so that in these three islands both variants of the species are present. In Puerto Rico the species showed an east—west highland distribution (Figure 26). Material seen: West Indies. PUERTO RICO: Luquillo Mountains, E1 Verde, on rocks in Rio Espiritu Santo, Landrén 337, 1967 (MSC); Bosque Estatal de Guavate, in Tabebuia ————__.—————.———.—_ ——.__..______.__—__—._.————_-_—___ lézi, RERE, RERZ, RERR, RRRRR, lgig, 1967 (MSC); Cerro Faralldn, Guavate, route to Carite Lake, ca. 750 m., Landron 1.443. 1w. use. 2122. 13631. Lager. an. 1967 (MSC); caimitillo, palm forest, midway between Guavate and Carite, en route to Carite Lake, Landrén 1489B, 1490B, 1491B, 1967 (MSC); Monte del Estado, Cordillera Central, ca. 833 m., Maricao, Landr6n 1508A, 1509, 1511A, 1514B, 1516, 1517, 1520, 1521A, 1522, 1534B, 1967 (MSC); observation tower, Monte del Estado Maricao, ca. 840 m., Landrén 1543B, 1546, 1549B, 1552A, 1553, 1554B, 1555, 1558, 1565, 1562, 1585A, __..—._———————_—_—_ 1586B, 1595A, 1969 (MSC); Cerro Farallon in Tabebuia rigida forest, trail to Carite Lake, Landron 1604, 1611B, 1622C, an. 1612. 1643 Eucalyptus, 5 ie 1 1686A, 1687, 169 ——____.— Villalba, in gua M, 1743A, y Divisoria, 800 m M, 1815, E 1967 (MSC); Lago M 1915c, 1 265 1641, 1642, 1643A, 1659, 1668, 1967 (MSC); Cerro Dofia Juana, Eucalyptus, sierra palm forest, ca. 850 m., Landron 1674A, 1686A, 1687, 1690B, 1693A, 1697B, 1967 (MSC); Barrio Caonillas Villalba, in guava scrub, ca. 800 m., Landrén 1718A, 1738B, 1742B, 1743A, 1745A, 1748, 1967 (MSC); Cordillera Central, Divisoria, 800 m., Villalba, Landr6n 1809, 1810A, 1811D, 1812A, 1815, 1817A, 1822A, 1824B, 1834A, 1842B, 1845B, 1854B, 1967 (MSC); Lago Carite in coffee plantation, ca. 500 m., Landrén 1915C, 1967 (MSC); Treasure Island, 5 mi. from Cidra, 400 m., Landr6n 1960, 1961, 1962, 1967 (MSC); Barrio Pul- guillas, mahogany trees, Aibonito, Landrén 1972, 1967 (MSC); Bosque Toro Negro,pasture with scattered Buchenavia capitata, Landrén 2310B, 2319A, 2322A, 2343A, 1968 (MSC); Lago E1 Guineo, route to Cerro de Punta, Landrén 2354, 1968 (MSC); landslide along route P.R. 143, near Divisoria, Toro Negro, Landron 2367B, 2371B, 1968 (MSC); Monte del Estado, Barrio Tabonuco, Landrén 2413A, 2431B, 2436A, 1968 (MSC); seven miles south of Caguas, Mr. and Mrs. Heller 296, 1899 (MICH); Arroyo de los Conchos, between Adjuntas and Jayuya, ca. 800-900 m., C. G. Britton 5301A, 1915 (MICH); Aibonito, open field at 200 ft. on rock, Fink 1895, 1932, 1935, 1916 (MICH); Coamo, G. E. Britton (MICH-12413). TOBAGO: Moriah, Weiss RE, 1923 (PH); Easterfield Road, east of Mason Hall, cultivated area ca. 800 ft., Imshaug 31172, 31193, 1963 (MSC); Flagstaff, at northeast end of Charlotteville, ca. 1179 ft., Imshaug 31344, 1963 (MSC); Pigeon Hill Road, 700-1000 ft., ea 1963 (MSC); coas San Fernando, E 800ft., Hillsbc M, 1963 (MSC Charlotteville e TRINIDAD: La V2 319g, 1963 (MS( Négre Maron Traj NeSt 0f La Vache “01116 Bleu Pass 1963 (MSC). JN ILshaLg 15894 2000 ft. Imshai E Law 19 \ daestra, M 22 R . v ama w 11 Lic w SneOideg 266 700-1000 ft., east end of Main Ridge, Imshaug 31763, 31734, 1963 (MSC); coast of Point Fortin, Guapo Bay, southwest of San Fernando, Imshaug 32235, 1963 (MSC); cultivated area, 800 ft., Hillsborough Dam on Mt. St. George, Imshaug 31571, 31582, 1963 (MSC); Pigeon Hill road, 700—900 ft. between Charlotteville and Speyside, Imshaug 31499, 1963, (MSC). TRINIDAD: La Vache Point (east of La Vache Bay) Imshaug 31998, 1963 (MSC); cultivated area in forest clearing, Negre Maron Trail, northern slope of Northern Range, south- west of La Vache, Imshaug 32025, 1963 (MSC); south side of Morne Bleu Pass on Arima-Blanchisseuse Road, Imshaug 32332, 1963 (MSC). JAMAICA: St. Andrew, Claremont, ca. 1400 ft., Imshaug 15894, 1953 (MSC); St. Ann, Brown Town to Bamboo, 2000 ft., Imshaug 15960, 1953 (MSC); Goshen 1000 ft., Im- shaug 15982, 1953 (MSC). CUBA: on slope of El Gato, Sierra Maestra, Imshaug 24800, 24814, 1959 (MSC). 22. Ramalina usnea (L.) R. H. Howe Lichen usnea L. Mantissa 1: 131—132. 1767. Parmelia usneoides Ach. Meth. Lich. 270. 1803. Ramalina usneoides Fr. Lich. Europ. reform. Add. 468. 1831. Ramalina usnea (L.) R. H. Howe, Bry010gist 17: 81. 1914. Original material: ”Indiae Or. Ins. Helenae, Madagascar, Martinicae, Jacqu (Linn.!). Ramalina usneoides var. usneoidella (Nyl.) Bull. Soc. Linn. Norm. II. 4: 122. Ramalina usnea var. usneoidella (Nyl.) R. H. How material: Mexic Ramalina Lectotype (nov.) Descript dulous, to 35 cm branches varying compressed, flat with attenuate c quently lateral] strands visibly APOthecia laterg Pale brown; hch gflatinous, C16, tips glObose 0r 267 (Nyl.) R. H. Howe, Bry010gist 17: 81. 1914. Original material: Mexico, Ghiesbreght (non vidi). Ramalina subanceps Nyl. Flora 59: 411. 1876. Lectotype (nov.): Cuba, Wright RR (FHI). Description: Thallus green or stramineous, pen- dulous, to 35 cm. long; laciniae linear, much branched, branches varying from very thin (0.5 mm.) to broad 5 mm.), compressed, flattened and frequently contorted, thin branches with attenuate often terete apices, while wide branches fre- quently laterally digitate; cortex thick, cartilaginous strands visibly separated by longitudinally running striae. Apothecia lateral, pedicellate, 0.2-6 mm. wide, disk convex, pale brown; hypothecium colorless, 12-16u thick; hymenium gelatinous, clear, 30-40u thick; paraphyses branched with tips globose or capitate. Asci numerous, fertile, contain- ing 8 spores in two sets of four; spores uniseptate, hyaline, fusiform, narrow, 16—35 x 3-50. Medullary reactions: PD-, K— or K+ pink, KC- or KC+ pink, C— or C+ pink. Lichen substances: Divaricatic, sekikaic, ramalin- olic and usnic acids, and substance H. Discussion: Linneaus (1767) emphasized in his original description of Lichen usnea the filamentous, com- pressed character of the thallus. Acharius (1803) stressed the thalli and apothecia characters of the species. Nylander (1870) used the spore size to establish R. usneoides var. M Le subanceps from C andits variety in R.subanceps ofwhich I have chections iden andesignating W mrmmlogical, s enmgh magnitude foreach to dese The spec n a chemical 5p maction_ This broader Spores a asci. 268 usneoidella. Later, in 1876, he described the species R. subanceps from Cuba and stated its similarity to R. usneoides and its variety usneoidella. Nylander did not select a type for R. subanceps; however, he cited Wright RR and RR, both of which I have examined and found indistinguishable from collections identified by the same author as R. usnea. I am designating Wright 21 (FH) as lectotype. There are no morphological, spore, chemical or ecological characters of enough magnitude between plants of different chemical races for each to deserve a name as a different species. The species R. anceps was considered by R. H. Howe as a chemical species differing from R. usnea by its K+ red reaction. This species (R. anceps) also differs by its broader spores always arranged in a single row within the asci. It is on the basis of the combined occurrence of both spore and chemical characters that I recognize R. anceps as a different species. Occasionally R. usnea may be mis- taken in the field for R. peranceps, but they can easily be distinguished by the production of salazinic acid in R. per— anceps and its absence in R. usnea. Ramalina usnea is a PD— species variously reacting K+ or K—. The species has been reported containing sekikaic and ramalinolic acids (Asahina, 1938; Asahina and Shibata, 1954; Santesson, 1967; and Follman and Huneck, 1969). The type specimen of R. 22222 (L.) R. H. Howe was described by him as reacting KOH—. Thin-layer chromatogram of a sample of the type mate shown to contain The West strated that spe icatic acid or t Others may lack that reacted K+ acids. This che as K-. The reas first place, the place, in old 51: be noticed. One content was four which did not cl acetic acid (90: Curved needles 5 Similarly, many substance. 269 of the type material (Linn. 879; Linn. Hb. 1273—278) was shown to contain divaricatic and usnic acids. The West Indian and other material studied demon— strated that specimens reacting K— may either contain divar- icatic acid or the so-called substance H found in Cladonia. Others may lack both substances. The population sample that reacted K+ pink contained sekikaic and ramalinolic acids. This chemical variant has consistently been reported as K—. The reason for this is probably two-fold. In the first place, the reaction occurs slowly and in the second place, in old specimens the color produced is too faint to be noticed. One specimen from Cuba with this K+ medullary content was found also to produce an accessory substance which did not chromatogram in benzene-dioxane-glacial acetic acid (90:25:4) and which recrystallized in G.E. long curved needles similar to those described for substance H. Similarly, many collections proved to contain no medullary substance. Accordingly, the world population of R. usnea can be grouped into four chemical variants: chemical variant I (divaricatic acid), variant II (sekikaic acid), variant III (substance H), and variant IV (no medullary substance). My concept of Ramalina usnea includes four chemical variants, each one of which was previously associated to a Species name. Table 13 summarizes the relationships between Chemical variant and species names. Table 13. Spec of _R Chemical Variant Su 1 Di II Se III Su IV No \ Ecology Species sometim range it occurs C001 to warm, f from continenta In Puer‘ and trunks of T. a u M gflhg 22233 you in associ c raciIIS, R Usnea Spp 270 Table 13. Species names associated to chemical variants of R. usnea. Chemical Lichen Previous Variant Substances Species Epithet I Divaricatic acid Ramalina usnea II Sekikaic acid R. subanceps III Substance H R. pallida Magn. ined. IV None i?“ usneoides auct. Ecology: Ramalina usnea is a corticolous, pendulous species sometimes reaching 2—3 feet long. Throughout its range it occurs in habitats ranging from dry to wet, from cool to warm, from low elevations to high elevations, and from continental to maritime climates. In Puerto Rico the species is epiphytic on branches and trunks of Tabebuia heterophylla, Buchenavia capitata and Ceiba pentandra, in the lower montane rainforest. It grows in association with Ramalina anceps, R. subpellucida, R. gracilis, R. complanata, Teloschistes flavicans and Usnea spp. The numerous forms and varieties of R. usnea that have been described reflect the high degree of variability within the species. This is probably the result of heredi- tary as well as environmental influences. For instance, in the dry islands of Tortuga and Margarita the specimens have broad laciniae mixed in the same thallus with the more typical thin or contorted char: seemed to be 1"! Distril distribution. some correlatit graphical dist: to Panama the t unidentified St medullary 11gb, eSpecially Venn excluding Trin; either divaric; Paragllay haVe ; 271 typical thin ones found on moist mountainous areas. The contorted character of the branches throughout its range seemed to be related to dry land or maritime habitats. Distribution: The species has a sub-cosmopolitan distribution. It is apparent from Table 14 that there is some correlation between chemical variants and the geo- graphical distribution. In Central America from Mexico to Panama the examined collections either produced the unidentified substance H found in Cladonia or have no medullary lichen substances. In northern South America, especially Venezuela and its Caribbean outlying islands, excluding Trinidad and Tobago, the population contained either divaricatic or sekikaic acid. The specimens from Paraguay have sekikaic and ramlinolic acids, while those from Galapagos Islands have only usnic acid. The West Indies and southern United States harbor both the divari— catic and sekikaic types, while Grand Cayman has the divaricatic, the substance H producing variants and that without medullary substances. Specimens from Africa are of the usnic acid producing population. The absence of European records in the table is due to the fact that the material studied in herbaria under the name R. usnea was found to be R. siliguosa. Table 14 is not intended to show the world distri- bution of the species but localities of specimens examined arranged in geographical categories. It is evident that Tmle l4. Geog of E S.United State Nest Indies Caural America N.South Americ S.South Americ Galépagos Is. Africa X— the Specties ten smnhern part 0 vafiants are re In the 272 Table 14. Geographical distribution of chemical variants of Ramalina usnea. Sekikaic Divaricatic Substance No Medullary acid acid H Substance S. United States x x x West Indies x x x x Central America x x N. South America x x S. South America x Galapagos Is. X Africa x the species tends to lack medullary substances toward the southern part of the tropical zone, while all the chemical variants are represented in the American tropics. In the West Indies the divaricatic acid variant is common in the Greater Antilles, Grand Cayman and Curacao, while the sekikaic acid strain is only present in Cuba and Jamaica. While both strains are present in Florida, they are absent from the main arch of the Lesser Antilles. Table 12 shows the distribution of chemical strains in the West Indies. The Puerto Rican population of the species is con— fined to the divaricatic acid containing variant. This has been collected only in the central part of the island. Figure 27 is a map of the species distribution in Puerto Rico, Materiz fl (MICH); L West It Negro, Cerro Dc 168i, 1967 am 800 m., Landrdr Nontafias de C01 Toro Negro, pas 182i» 18% 1 between Casa M: 20$, 1967 (Ms( 23$, 1968 (Ms( no locality g1, Ville, Cushman \ Parish of Hanm Parish of St. 7 273 Material seen: Exsiccati examined. Merr. I. 04, ...—- 236 (MICH); W. Web. RR, 115 (MICH). West Indies. PUERTO RICO: Bosque Estatal Toro Negro, Cerro Dona Juana, 800 m., Orocovis, Landron 1682A, 1685B, 1967 and 2338, 1968 (MSC); Barrio Caonillas, Villalba, 800 m., Landron 1724B, 1725A, 1967 (MSC); Barrio E1 Negro, Montafias de Corozal, 500 m., Landrén 1782, 1786C, 1967 (MSC); Toro Negro, pasture with scattered trees, 850 m., Landron 1822C, 1845A, 1855B, 1967 (MSC); Aibonito, near Cuydn, between Casa Manresa and National Guard Armory, Landrdn 2005, 1967 (MSC); Adjuntas, Cerro de Punta, 1150 m., Landrén 2392, 1968 (MSC); no locality given, Fink 1971, 1916 (MICH); no locality given, Bertero 3.2., 1827 (H). JAMAICA: Mande— ville, Cushman RZR (FH, MICH); Orcutt REEL, 1928 (US); Parish of Hanover, 1809 ft., Imshaug RRQZR, 1953 (MSC); Parish of St. Andrew, seasonal forest, 2450—2550 ft., sum- mit of Cooper's Hill, Imshaug RRZRR, 1952 (MSC); Cooper's Hill, 2450 ft., Imshaug RRRRR, 1953 (MSC); rocky bank on southwest slope near summit of Mt. Rossana, 4000 ft., Rm- shaug 13991, 1953 (MSC); south slope of Cooper's Hill in Red Hills, 2250 ft., Imshaug 13724, 13725A, 1952 (MSC); Parish of St. Ann, Hopewell, 1400 ft., Imshaug RRRRR, 1953 (MSC); Lime Hall to Green Park, 1500 ft., Imshaug REERR, 1953 (MSC); Browns Town to Bamboo, 2000 ft., 15976B, 1953 (MSC); Goshen, 1000 ft., Imshaug RRRRZ, 1953 (MSC); Parish of St. Catherine, Guys Hill, Imshaug 13465, 1952 (MSC); Mount Diablo, 2 Prov. Pinar del Alturas de Piza pines and oaks of Vifiales, 500 Oriente, betwee Sierra Maestra, Jaibo, Bro. Hio Prov. Puerto P1 from Santiago t Cerrazo, 2450 f Cordillera S€pt Cristi: 36 km. “£17925. 19c Pilate, 325 Ill. Bombardopolis, Jacmel, Thomas \ 274 Mount Diablo, 2600 ft., Imshaug 13798, 1952 (MSC). CUBA: Prov. Pinar del Rio, northwest of St. Vincente, 650 ft., Alturas de Pizarras. Imshaug 25218, 1959 (MSC); scattered pines and oaks in low wet area in Valle del Silencio, east of Vifiales, 500 ft., Imshaug RREQR, 1959 (MSC); Prov. Oriente, between El Gato and San Juan, Loma del Gato, Sierra Maestra, Imshaug RRRER, R3233, 1959 (MSC); Boca de Jaibo, Bro. Hioram 5482, 1921 (US). DOMINICAN REPUBLIC: Prov. Puerto Plata, north slope of La Cumbre, along road from Santiago to Puerto Plata, Imshaug 23848, 1958 (MSC); Cerrazo, 2450 ft., on ridge between Santiago and La Cumbre, Cordillera Septentrional, Wetmore 3851, 1958 (MSC); Monte Cristi, 36 km. from Monte Cristi on way to Santiago, Wet- more 17925, 1968 (MSC). HAITI: Dépt. du Nord, vicinity of Pilate, 325 m., E. C. Leonard 9602, 1926 (US); vicinity of Bombardopolis, 210 m., Leonard and Leonard 13469, 1929 (US); Jacmel, Thomas ZR, 1935 (PH); vicinity of La Vallée, Tortue Is., Leonard and Leonard 11531, 1928—1929 (US). GRAND CAY— MAN: On 1ow shrubs on beach at West Bay (between Galleon Beach Hotel and West Bay Town), Imshaug 24380, 1959 (MSC). BAHAMAS: San Salvador Is., northeast corner of island, near Bonefish Bay, Gillis 8867, 1970 (MSC). CURACAO: Cristoffel— berg, Harman and Curran 128, 159, 1917 (FH). MARTINIQUE: _— no data given, Plée 3.3. (G—1522). Central America. MEXICO: Las Palmas, San Luis Potosi, Pringle 206, 1896 (MICH); Oaxaca, Conzatti 176, 1918 (MICH). P. South A] 3.13., 1903 (MSC end of Tortuga Punta Arenas, w (MICH). North A Q, 1931 (MICH (MICH); Sanford 531-3., 1909 1967 (MSC); Sar 275 1918 (MICH). PANAMA: R;1_1_s_.r_i. (FH). South America. VENEZUELA: Coche Is., Blakeslee 3.2., 1903 (MSC, MICH); Robertson E'i" 1900 (MSC); western end of Tortuga Is., Taylor ZRR, Zéi, Zég, 1939 (MICH); Punta Arenas, western end of Tortuga Is., Taylor E'E'» 1939 (MICH). North America. FLORIDA: Long Boat Key, MacFarlin RRR, 1931 (MICH); Ana Maria Key, MacFarlin 121, RRR, 1931 (MICH); Sanford, Kelly 105, 1921 (MICH); Ponce Park Steven- —_.__. son s.n., 1909 (MICH); Volusia Co., Harris 2293, 2963B, 1967 (MSC); Sarasota Co., Moore 1140, 2958, 1968 (DUKE). Additional material seen: COLOMBIA: Santa Marta, Baker R, RR, 1898 (MICH). PARAGUAY: no locality given, Malme 5.2., 1859? (MICH). ECUADOR: Galapagos Is., Academy Bay, Indefatigable Is., Herre 3.3., 1929 (MICH). Subsection Fistularia (Vain). Zahlbr. 23. Ramalina inflata (Hook. f. G Tayl.) Cetraria inflata Hook f. G Tayl. Lond. Journ. Bot. 3: 646. p1. 79./fig. 1. 1844. Ramalina inflata (Hook. f. G Tayl.) Hook f. & Tayl. in Hooker f. Fl. Antarct. I: 194. 1845. Original material: Lord Auckland Is. (PH-Taylor 1284!, BMl). Ramalina geniculata Hook. f. Tayl. Lond. Journ. Bot. 3: 655. 1844. Original material: New Zealand, (PH—Taylor 1282!, EMS). 11:52. 1908. GHQ. Descrip Mgh,sparsely bmad,subteret fomu surface n tmnwus with la florediose; cor mmtly terminal dfik invaginate C0“cal/e; hymeni branched, non-s “ranged in twc mid,curVed 1( Li m divaricatic ac aCid. 276 Ramalina inflata var. soredians Merr. Bry010gist 11: 52. 1908. Original material: Jamaica, Cummings 2.3. (PHI). Description: Thallus caespitose, short, 204 cm. high, sparsely branched, dichotomous; branches 0.5 to 2 mm. broad, subterete, fistulous, rarely compressed, tips fili- form; surface nitid, cartilaginous strands seen, discon- tinuous with large perforations or many small punctures; esorediose; cortex thin; medulla scant. Apothecia common, mostly terminal, or appearing geniculate, conical pedicel; disk invaginate, flattened, tan, 1-3 mm. across, rarely concave; hymenium + epithecium 30-40u thick, paraphyses branched, non-septate. Asci clavate, containing 8 spores arranged in two series; spores uniseptate, hyaline, ellip— soid, curved 10-13 x 4-6u. Medullary substances: Pd—, K— or K+ faint pink, KC— or KC+ faint pink, C- or C+ faint pink. Lichen substances: Atranorin, Chloroatranorin, divaricatic acid, sekikaic acid, ramalinolic acid and usnic acid. Discussion: Hooker and Taylor separated R. inflata from the similar species R. geniculata by being smaller and differently branched. Howe (1914) considered these two species as synonyms and selected R. inflata as the correct name. My comparison of the types also shows that the two Species are identical. Nylande Madeira Is. as Nandos no. 24 a in 1964 from Te 1_at_a differs fr and having apot terminal. This The sor is based on a m Which contained m- Merrll wer,is diffic ofthe scant me M g 333153133 Thewest Indies oussPecies R. [Mw ‘ Zealand) , a SEkikaiC an 277 Nylander (1870) described R. subgeniculata from the Madeira Is. as having a foraminous thallus. I have studied Mandos no. 24 and also several collections made by Imshaug in 1964 from Tenerife (Canary Islands). Ramalina subgenicu— lata differs from R. inflata by being profusely branched and having apothecia lateral rather than terminal or sub- terminal. This Madeira species contains divaricatic acid. The sorediate variety described by Merril (1908) is based on a mixed collection made by Cummings in Jamaica which contained several thalli of R. inflata and R. coch- learis. Merril mistook the swollen tips of the sorediose R. cochlearis for the inflated thallus of R. inflata. Ramalina inflata and its variety soredians have been reported to react negative with K. This reaction, how- ever, is difficult to observe in fistulous specimens because of the scant medulla. Both the type of R. inflata and that of R. geniculata were found to contain divaricatic acid. The West Indies specimens depart markedly from the fistul- ous species R. inflata (Lord Auckland 15.), R. geniculata (New Zealand), and R. subgeniculata (Madeira Is.) by produc- ing sekikaic and ramalinolic acids. Ecology: Ramalina inflata is a corticolous species occurring in the West Indies at elevations that vary from 2500 to 6000 ft. In Lord Auckland Is. it grows on rocks near the sea. Distribi Auckland 15. am (Zahlbruckner, I West Indies (Me‘ species has bee: and Haiti. 1t Islands. Materia Scattered pines side of Constan (MSC); on shrub Constanza, 379C (NSC). HAITI: on ridge betwee M, 1958 (M5 Slope ridge ab< (NSC) ; summit , 278 Distribution: This species was described from Auckland Is. and has been reported from the Pacific region (Zahlbruckner, 1930b), North America (Howe, 1914) and the West Indies (Merrill, 1908). In the West Indies the species has been studied from Jamaica, Dominican Republic and Haiti. It is absent from the rest of the Caribbean Islands. Material seen: West Indies. DOMINICAN REPUBLIC: Scattered pines, ca. 3750 ft., summit of low ridge on north side of Constanza, Cordillera Central, Imshaug 23719, 1958 (MSC); on shrub in stream valley around E1 Chorro, near Constanza, 3790 ft., Cordillera Central, Wetmore 3802, 1958 (MSC). HAITI: Department de l'Ouest, summit of Téte Etang on ridge between Kenscoff and Furcy, ca. 6000 ft., Imshaug 22573, 1958 (MSC); cultivated and pasture area on north slope ridge above Kenscoff, 5000 ft., Imshaug 22504, 1958 (MSC); summit of Montagne Noire, ca. 5500 ft., near Ken- scoff, Wetmore 2752, 2756, 1958 (MSC); on Eucalyptus in cultivated area with a few scattered pines, hillside below Furcy, Wetmore 2671, 1958 (MSC). 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The floristic elements of vascular plants in the West Indies are the West Indian, endemic, continental, and cosmopolitan. In Puerto Rico the lichen flora is described within seven vegetation zones (Dansereau, 1966). These are the littoral, lowland rainforest, seasonal—evergreen forest, semi-deciduous forest, lower montane rainforest, montane forest and montane scrub. The genus Ramalina in Puerto Rico showed seven dis- tribution patterns primarily determined by precipitation, elevation and edaphic factors. The distribution patterns were Maricao-highland, central highland, south—western, east—west, north—central and scattered. The distribution of the genus in the West Indies is influenced by the position of the islands, climate and top- Ography. Ramalina does not show a Greater Antillean element more or less distinct from a Lesser Antillean. Of twenty- three (23) species present in the islands, nine (9) were found to be common to both the Greater and the Lesser Antil- les, four (4) were common to the West Indies and Florida, eight (8) were common to Central America and eight (8) were common to S The Indies is c a a t ext b a d C a g lar d a t Jam e a g Ame The to Include 307 common to South America. The distribution of the genus Ramalina in the West Indies is characterized by a. a tendency of the species to concentrate at the extremes of the chain of islands, b. a decrease in the number of species eastwardly, c. a greater floristic affinity between the three larger islands than between these and Puerto Rico, d. a tendency for the genus to be more diversified in Jamaica than in the other islands, and e. a greater floristic affinity with Central and South America than with North America. The genus Ramalina was described by Acharius in 1810 to include several species previously placed in the genus Parmelia (Acharius, 1803). The genus was monographed for the first time by Nylander (1870) and later R. H. Howe (1914) treated the North American species. The morphology of the genus is very variable in the external as well as in the internal features. Differences in cortical morphology have been the bases for subgeneric divisions. The presence or absence of soredia, papillae, pseudocryphellae, channels, pits and the form of the branches are of taxonomic importance at the species level. Chemical studies using spot tests, microchemical crystallization tests and thin—layer chromatography provided additional criteria for species identification. SeVI in material lichen subS‘ absent from ical varian substance) ' Ramalina. their chemi orcinol der orcinol der acids. The yielded twe 1i&-Oft MYGIOpoea a 308 Seventeen (17) lichen substances were demonstrated in material from the West Indies (Table 15). Four (4) lichen substances were present in species from Florida but absent from the West Indies species. Forty-one (41) chem- ical variants (including those species with but a single substance) were recognized in the West Indian species of Ramalina. The lichen substances are grouped according to their chemical features into para— and meta—depsides of orcinol derivatives, para—depsides and depsidones of B- orcinol derivatives, dibenzofuran derivatives and fatty acids. The taxonomic treatment of the genus Ramalina yielded twenty-three (23) species within the section Rama~ lina. Of these, twenty-two (22) belong to subsection Myelopoea and one (1) to subsection Fistularia. Eight (8) species belong to series Teretiusculae and fourteen (14) to series Compressiusculae in subsection Myelopoea. The species with several chemical variants were Ramalina subasperata (4), Ramalina complanata (7), Ramalina usnea (4) and Ramalina farinacea (5). In this last species material from other areas was included. “QLWCTLu-ulv “CWF—“EWQ + + II. mmv + + + + mmmmwmmmmw meeHnEmm + + mODNOHUCQU NCHHNENK annHu + + + + + WHWEOENESU mflflHmEmm + + + + + NHUCNHQEOU NEHNHEMNN 1T whommopfiflhAmU EEWNIWWH~5HQV§ + + dPHODmHfi dfimaofidd + + + + mMoodm ofifladfiom owHSOmfiammohmfiou moflhom + + + + T83? enigma + + + + + + «Dahommwcom wdflHmEdm mew + + + + + meDpoHOm wCHHdEmm + + + + wmo©5me mcHHdEmm + + + + Hocmwucoe mafiamewm + + + mflafiomgm mcHHmEmm + + + + + npwHHoossw mcflHwEmm IIIIIIIIIIIIII + , + + mopfloomflpwcop mcflfimamm mmU 4 , + + + + mflgmoficooo mafiamEmm + + + mumsoopem mcfifimamm 309 OGHSUmSHUOHOB moflhom woomoao>z COHDoomnom Qsfl fl Q5 Q5 S H d e.d d d N 01v G and D D V wcflHwEmm defipuom n u s 1 e e e s o 1 a a o e _ I. o 1 u. e 1 04% u e N. I m o I.o 1 T. 1 110 A I;A T. d 1 s u I. u I. e e 1 D14 I n s I.. a Did 0 e e 110 o o N. z I o o e D. 1 ofiu 1 1 1 1 u a M s e I. I. m ,9 1 o I. e I. o e e o u u e d I. u u I. e 0 s 3 9mm 3 .9 1 1 1 o o o o I. o o 1 T. I. 1 nae e H 1 I. I e I. 1 o I 1 I. o I. I;+ 1 I a o u s P I. a I. a e o o 05% I. o u o o e o o 1 e ,A o 1 o e I. 3 T: T: B D B T._ O .1 3 B D._ I. B D.. 3 D I. D O. B d T. I. o D. o I. D. I. e 3 q u P I. I. P P 1 T1 e P P _ P 9 TL. 3 T .mmcfiadamm mpfiHOHm pom moflch pmoz mo mucoDQOU anoflaosu .mH manna 310 111 Amy 1NVAHV a: + + + + + .m commDmQSm mmv + + + + + + + + + + + + + + + ... ... + + + + + ... + .m ouqmmeSm Amy .< oucmumnsm nNV .u commDmQSm may .3 anHDmQSm may ”WOUENQmQSW ondxfib eoeficee NEHHNEMM mflHmHSDmflm soapoomnzm MQwaHU®HOm mamw>osom 1 mmoocwao mEMonoumoa douwfiwhmw I wumfl flummm mopflopccow mflmcocmezu muwcmeEou whommODmfiwu ..rAHu 1: wcfiHmem mafiHwEmm mcfifiwamm mcflfimsmm wcfiHwEmm mcHHwEmm wcHHmEmm mcHHmEmm moflfimEmm mcflamfiwm mcflHmEmm mcflHmEmm . a a r...$..~/ UiéjUhiJNIUHLEOU IUerW dwxflflvfld m. 7) 40er Fauarrfldr MHMHOHHde/rszm duced. Haw/56M]. dwOMGOHOm dCHHmEm¢ dWOGSHQN dfiflfldgflfi dDGUflHSOE JNAnifli—idmiadu/H LITERATURE CITED Peri Scientific P Acharius, E. Stoc Literature Cited Periodical abbreviations follow the World List of Scientific Periodicals. Acharius, E. 1803. Methodus qua omnes detectos lichenes. Stockholmiae,394 + 52 pp. . 1810. Lichenographia universalis. Gottingae (Danckwerts), viii + 696 pp. . 1814. Synopsis methodica lichenum. Lundae, xiii + 392 pp. Agardh, K. A. 1821. Aphorism botanici. D. I. - VII. Lundae, (Typ. Berling), 246 pp. Ahmadjian, V. 1958. A guide for the identification of algae occurring as lichen symbionts. Bot. Not. 8(4): 632-644. . 1967. The lichen symbiosis. Blaisdell Pub. Co., Mass., xiii + 152 pp. Anthony, H. E. 1925. The mammals of Porto Rico, living and extinct. N.Y. Acad. Sci., Scient. Surv. P. Rico, 9(1-2): 1—406. Asahina, Y. 1934. Uber die Reaktion von Flechten-Tallus. Acta Phytochim. 8: 47—64. . 1936. Mikrochemischer Nachweis der Flechten— Stoffe. J. Jap. 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Jena xi + 499 pp. APPENDIX A LIST OF COLLECTION LOCALITIES Th Rico visit 1968-1969 poinding d Figure 6. included i collection 1-97 98-229 230-268 2692532 283296 List of Collection Localities _____________________________ The list below contains 75 localities in Puerto Rico visited during the summer of 1967 and winter of 1968-1969 accompanied by collection numbers and the corres- poinding dot number for reference to collection map in Figure 6. Collection numbers 2011 to 2180 are not included in this appendix because they belong to Michigan collections. 1-97 (1) 98-229 (2) 230-268 (3) 269-282 (4) 283—296 (5) HUMACAO DISTRICT: Palo colorado association (Cyrilla racemiflora), El Yunque, 630 m., Luquillo Mountains. (P.R. 191 km. 11.0). 27 June 1967. HUMACAO DISTRICT: Palo colorado association (Cyrilla racemiflora) El Yunque, 680-700 m., Luquillo Mountains (P.R. 930 km. 1.5). 28 June 1967. HUMACAO DISTRICT: Mossy forest, El Yunque, 700 m., Luquillo Mountains (P.R. 191 km. 13.7) 29 June 1967. HUMACAO DISTRICT: Palm Brake, La Mina, 550 m., Luquillo Mountains (P.R. 191). 29 June 1967. HUMACAO DISTRICT: Lower montane forest, palo colorado association (Cyrilla recemiflora) 327 297-338 339-370 371-423 424-517 518-611 ! 612~698 328 Espiritu Santo River, 380 m., El Verde, Luquillo Mountains (P.R. 186). 30 June 1967. 297-338 (6) HUMACAO DISTRICT: El Verde, rocks in Es- piritu Santo River. 380 m., Luquillo Moun- tains (P.R. 186). 23 June 1967. 339-370 (7) HUMACAO DISTRICT: Palo colorado-tabonuco (Cyrilla-Dacryodes) association, E1 Verde, 390 m., (P.R. 186 km. 7.7) l4.June 1967. 371-423 (8) HUMACAO DISTRICT: Tabonuco forest (Dacryodes excelsa), El Verde, 390 m., (P.R. 186 km. 7.7) 21 June 1967. 424-517 (9) HUMACAO DISTRICT: Palo colorado association (Cyrilla racmiflora), along creek, 390 m., (P.R. 186 km. 8). 22 June 1967. 518—611 (10) ARECIBO DISTRICT: Laguna Tortuguero, sand dunes 7 km. west of Vega Baja, at sea level P.R. 2). 3 July 1967. 612—698 (11) ARECIBO DISTRICT: Laguna Tortuguero, old pasture field at sea level, 7 km. west of Vega Baja (P.R. 2). 5 July 1967. 699—791 (12) ARECIBO DISTRICT: Laguna Tortuguero, south of lake, at sea level, 7 km. west of Vega Baja (P.R. 2). 6 July 1967. 792-886 (13) ARECIBO DISTRICT: Laguna Tortuguero, white sand flat, sea level, 7 km. west of Vega Baja (P.R. 2). 7 July 1967. 887-909 910-921 922-951 952-982 983-998 999-1048 1049-1100 329 887—909 (14) ARECIBO DISTRICT: Punta Cerro Gordo, coco- nut palm field at sea level, 25 km. north of Vega Alta (P.R. 690). 8 July 1967. 910—921 (15) ARECIBO DISTRICT. Pterocarpus swamp, Sar- dinera Beach, 5 km. west of Dorado (P.R. 693). 8 July 1967. 922-951 (16) HUMACAO DISTRICT: Mossy forest, East Peak, 1051 m., Luquillo Mountains (P.R. 630). 10 July 1967. 952-982 (17) HUMACAO DISTRICT: Mossy forest, 700 m., Luquillo Mountains (P.R. 630). 11 July 1967. 983-998 (18) HUMACAO DISTRICT: Palm forest, La Mina, 500-550 m., El Yunque, Luquillo Mountains (P.R. 191). 11 July 1967. 999-1048 (19) HUMACAO DISTRICT: Elfin woodland, Mt. Brit- ton, 1065 m., El Yunque, Luquillo Mountains, 11 July 1967. 1049—1100 (20) ARECIBO DISTRICT: Laguna Tortuguero, south- east of lake, 7 km. west of Vega Baja P.R. 2). 12 July 1967. 1101-1109 (21) HUMACAO DISTRICT: Palm brake, route to El Toro Peak, 710 m., El Yunque, Luquillo Moun— tains (P.R. 191 km. 14). 12 July 1967. 1110—1119 (22) HUMACAO DISTRICT: Elfin forest, El Yunque Rock, 1065 m., Luquillo Mountains. 14 July 1967. 1131-1143 1144-1171 1 1172-1192 1193-1225 1226—1402 1403-1443 1444~1487 l488-149: 330 1112-1130 (23) HUMACAO DISTRICT: Elfin woodland, route from Mt. Britton to The Pinnacles, 1000 m., Luquillo Mountains. 14 July 1967. 1131-1143 (24) HUMACAO DISTRICT: Mossy forest, East Peak, 1051 m., Luquillo Mountains. 14 July 1967. 1144—1171 (25) HUMACAO DISTRICT: Semi—sabana, East Peak, 800 m., Luquillo Mountains. 15 July 1967. 1172-1192 (26) HUMACAO DISTRICT: Route to Naguabo, 800 m., Luquillo Mountains (P.R. 191 km. 15). 16 July 1967. 1193—1225 (27) HUMACAO DISTRICT: Route to Naguabo, Rio Sabana, 500 m., Luquillo Mountains (P.R. 191 km. 20) 16 July 1967. 1226-1402 (28) GUAYAMA DISTRICT: Bosque Estatal de Guavate, 550 m., near river, Cayey (P.R. 184 km. 6) l7-18 July 1967. 1403-1443 (29) GUAYAMA DISTRICT: Bosque Estatal de Guavate, near Park, 550 m., Cayey (P.R. 184 km. 8). 19 July 1967. 1444-1487 (30) GUAYAMA DISTRICT: Cerro Farallén, Guavate, 750 m., route to Carite Lake, 600 m., (P.R. 184). 28 July 1967. 1488—1499 (31) GUAYAMA DISTRICT: Carite-Guavate, route to Carite Lake, 600 m., (P.R. 179). 28 July 1967. 1500-1504 1505 1506-1507 1508-1541 1542-1579 1580~1595 1632‘1642 164S~l671 331 1500—1504 (32) MAYAGUEZ DISTRICT: Cerro Las Mesas, 350 m., near Mayagfiez Army Base. 31 July 1967. 1505 (33) MAYAGUEZ DISTRICT: XerOphytic forest, Bos- que de Guénica, Cafia Gorda Beach. 1 August 1967. 1506-1507 (34) MAYAGUEZ DISTRICT: Laguna Cartagena, Barrio Magfiayo (Santana), Lajas. 1 August 1967 1508-1541 (35) MAYAGUEZ DISTRICT: Monte del Estado, Cor- dillera Central, 833 m., Maricao (P.R. 120 km. 12). 1 August 1967. 1542-1579 (36) MAYAGUEZ DISTRICT: Monte del Estado, slope east of observation tower, 840 m., Maricao (P.R. 120 km. 12). 2 August 1967. 1580-1598 (37) MAYAGUEZ DISTRICT: Monte del Estado, slope west of observation tower, 840 m., Maricao (P.R. 120 km. 15). 2 August 1967. 1599—1631 (38) MAYAGUEZ DISTRICT: Monte del Estado, slope east of radio antennas, 833 m., Maricao (P.R. 120 km. 17-20). 2 August 1967. 1632-1644 (39) MAYAGUEZ DISTRICT: Monte del Estado, slope west of radio antennas, 833 m., Maricao (P.R. 120, km. 15—17). 2 August 1967. 1645—1672 (40) GUAYAMA DISTRICT: Guavate, Cerro Farallén, 900 m., route to Carite Lake (P.R. 184). 3 August 1967. 1673-1698 1699-1710 1711-1728 1729-1750 1751-1778 1779-1787 1788—1806 1807~1857 1673—1698 (41) 1699-1710 (42) 1711-1728 (43) 1729-1750 (44) 1751-1778 (45) 1779-1787 (46) 1788-1806 (47) 1807—1857 (48) 1858-1882 (49) 332 PONCE DISTRICT: Bosque Estatal Toro Negro, Cerro Dofia Juana, 800 m., Orocovis (P.R. 143). 3 August 1967. PONCE DISTRICT: Bosque Estatal Toro Negro, near forestry building, 850 m., Orocovis (P.R. 143). 3 August 1967. PONCE DISTRICT: Barrio Caonillas, Villalba, 800 m., (Intersection of routes P.R. 143 and P.R. 151). 3 August 1967. PONCE DISTRICT: Cordillera Central, Divi— soria, 800 m., Villalba (P.R. 143 km. 11). 3 August 1967. PONCE DISTRICT: Barrio Cuchillas, 600 m., 10 km. south of Corozal, Orocovis (P.R. 568). 7 August 1967. PONCE DISTRICT: El Negro, Montafias de Coro— zal, 550 m., south of Corozal, Orocovis (P.R. 805). 7 August 1967. ARECIBO DISTRICT: Barrio Padin, Montafias de Corozal, 500 m., (P.R. 568). 7 August 1967. PONCE DISTRICT: Bosque Estatal Toro Negro, Cerro Dofia Juana, pasture with scattered trees, 850 m., Orocovis (P.R. 143). 8 August 1967. GUAYAMA DISTRICT: Lago Carite, coffee plan- tation, 600 m., Guayama (P.R. 179). 10 August 1967. 1883-195 1952-195$ 1960-1964 1965—1974 i 333 1883—1951 (50) PONCE DISTRICT: Monte Cuba, quitas Hotel, 600 m. near Barran- , Barranquitas (P.R. 156). 11 August 1967. 1952-1959 (51) GUAYAMA DISTRICT: Monte Llano, 500 m. east of Cidra (P.R. 173). 1960-1964 (52) GUAYAMA DISTRICT: , 7 km. 12 August 1967. Treasure Island, 400 m., 1 km. east of Cidra (P.R. 173). 12 August 1967. 1965-1974 (53) PONCE DISTRICT: south of Aibonito, 724 km. 4). 12 August 1967. 1975-2006 (54) PONCE DISTRICT: Cuy6n, 500 m., Aibonito (P.R. 162). 13 August 1967. 2007-2010 (55) PONCE DISTRICT: Bafios de Coamo, 100 m., south of Coamo (P.R. l4). 13 August 1967. 2181-2197 (56) ARECIBO DISTRICT: Barrio Los Hoyos, 50 m ' S 3 km. west of Vega Alta (P.R. 690). 8 December 1968. 2198-2212 (57) ARECIBO DISTRICT: South of Dorado Hilton Hotel, 2 km. west of Dorado at sea level (P.R. 693). 9 December 1968. 2213—2261 (58) ARECIBO DISTRICT: Laguna Tortuguero, old field, east of lake, at sea level, 7 km. west of Vega Baja (P.R. 2). 10 December 1968. 2262-227 2273-228 2281-228 2286 2287-229' 334 2262-2272 (59) ARECIBO DISTRICT: Lago Dos Bocas, 250 m., 10 km. north of Utuado (P.R. 10). 12 Decem— ber 1968. 2273-2280 (60) AGUADILLA DISTRICT: Guajataca Beach, 50 m ' 9 (P.R. 2). 12 December 1968. 2281-2285 (61) AGUADILLA DISTRICT: Along road, 10 km. west of Guajataca Beach (P.R. 2). 12 Decem— ber 1968. 2286 (62) ARECIBO DISTRICT: Barrio Maricao, 100 m., 7 km. south of Vega Alta (P.R. 677). 13 December 1968. 2287—2297 (63) ARECIBO DISTRICT: Laguna Tortuguero, south— east corner of lake, 7 km. west of Vega Baja (P.R. 2). 14 December 1968. 2298—2309 (64) ARECIBO DISTRICT: Tortuguero Army Base, at sea level, 5 km. west of Vega Baja (P.R. 2). 14 December 1968. 2310—2349 (65) PONCE DISTRICT: Bosque Estatal Toro Negro, Cerro Dofia Juana, 850 m., (Intersection P.R. 143 and P.R. 564). 16 December 1968. 2350—2360 (66) PONCE DISTRICT: Lago El Guineo, 1000 m., east of lake, Villalba (P.R. 143). 17 Decem- ber 1968. 2361—2385 (67) PONCE DISTRICT: Barrio Divisoria, 850 m., route from Orocovis to Villalba (P.R. 143). 17 December 1967. 2386-238i 2389-2401 2408-242( 2421-243! 2456~2455 2460~2461 2386-2388 (68) 2389-2407 (69) 2408-2420 (70) 2421-2439 (71) 2440-2449 (72) 2450-2455 (73) 2456-2459 (74) 2460—2462 (75) 335 PONCE DISTRICT: About 10 km. north of Barranquitas, 500 m., (P.R. 152). 17 Decem- ber 1968. PONCE DISTRICT: Cerro de Punta, 1150 m., Adjuntas (P.R. 143). 17 December 1968. MAYAGUEZ DISTRICT: Monte del Estado, near Stone House, 830 m., Cordillera Central, Maricao (P.R. 120). 19 December 1968. MAYAGUEZ DISTRICT: Barrio Tabonuco, 500 m., Monte del Estado, Maricao (P.R. 120). 19 December 1968. GUAYAMA DISTRICT: Cerro Pandura, 500 m., Yabucoa (P.R. 3). 4 January 1969. ARECIBO DISTRICT: Punta Cerro Gordo, north of Vega Alta (P.R. 690). 5 January 1969. GUAYAMA DISTRICT: Barrio Cafiaboncito, 200 m., 5 km. west of Caguas. 8 January 1969. AGUADILLA DISTRICT: Guajataca lake, near Boy Scout Summer Camp, San Sebastian, 7 January 1969. APPENDIX B THE NATURAL PRODUCTS REPORTED FROM THE LICHEN GENUS RAMALINA 2. Evernic acid Methyl 3,5—dichlorolecanorate 4. Perlatolic acid 5. Stenosporic acid Meta—depsides (Orcinol derivatives) 1. Boninic acid 2. Cryptochlorophaeic acid 3. Paludosic acid 4. Ramalinolic acid 5. Sekikaic acid Para—depsides (B-orcinol derivatives) 1. Atranorin 2. Chloroatranorin 3. 4—0—demethylbarbatic acid 4. Obtusatic acid Depsidones (B—orcinol derivatives) 1. Hypoprotocetraric acid 2. Norstictic acid 3. Protocetraric acid 4. Psoromic acid 336 Dibe Fatt Mave II o Prim} 337 5. Salazinic acid 6. Stictic acid Dibenzofuran derivatives 1. Usnic acid Fatty acids 1. Aspicilin 2. Caperatic acid 3. Tetrahydroxy fatty acids Mavelonicacid-derived products 1. (—) - l6ahydroxykaurene II. Primary products (Saccharides) l. D-arabitol 2. Isolichenin 3. Lichenin 4. Mannitol APPENDIX C LIST OF LICHEN SUBSTANCES REPORTED IN 114 SPECIES OF RAMALINA List of lichen substances reported in ll4 species of Ramalina. Table I6. 338 UlAeHqu IIIIIIIIIIIIIIIIIIIII-nn E================Efl==“" I uweuon + , III-I _IIIIIIIIIIIIIIIIIIII-Inln 1041qu9—0 IIIIIIIIIIIIIIIIIIII-In... us I so 1 -sv IIIIIIIIIIIIIIIIIIIII-II EIIIIIIIIIIIIIIIIIIII “iii _III-IIIIIIIIIIIIIIII III-l _III-IIIIIIIIIIIIIIII IIIII a eoumsqns IIIIIIIIIIIIIIIIIIII I...- V eouelsqns IIIIIIIIIIIIIIIIIIII . I...- H eouelsqns .IIIIIIIIIIIIIIIIIII I... p o wen In I. I IIIIIIIII III III ‘ ““ AXOJpAqJewl III I ...-III IIIII F3199 Owns IIIIIIIIIII ‘t‘T—MIIIIIIIIIIIIIIIIIIII -::I _III-IIIIIIIIIIIIIIII ...-II _IIIIIIIIIIIIIIIIIIII - I. ll 4 p109 3! 10" 1 [ewes IIIIIIIIIIIIIIIIIIII -; “II _IIIIIIIIIIIIIIIIIIII "II"- p... museum IIIIIIIIIIIIIIIIIIIH ., I III (>109 31|O+9|J9d IIIIIIIIIIIIIIIIIIII . I...- —IIIIIIIIIIIIIIIIIIII .3 I IIII=IIIIIIIIIIII , II + IIIIIIIIIIIIIIIII .: III III II IIIIIIIIIIIIIIIII —g‘§ IAq4ew I II I more 01494490040Jd04H II IIIIIIIIIIII II . :— IIIIIIIIIIIIIIIIIIIE . fig _III-IIIIIIIIIIIIIII . .- _IIIIIIIIIIIIIIIIIIII I p we ‘I‘ . IIIIIIIIIIIIIIIIII x IIIIIIIIIIIIIIIII . _III _IIIIIIIIIIIIIIIIIIII _III-IIIIIIIIIIIIIII= I: _III-IIIIIIIIIIIIIII _IIEEIEIIIIIIIIIIIIIIIIIIII \ EI- . t) I \\ \ \ U‘) u >K (I) .9 (g m E (U +‘ \\ U >)< m (U — 3 ‘3 co ‘ CD to (o (o * .- ._ (o :5 s. m 0 3 m 0 . Q. U’) (o c: U 4— (o m +— C 0 (U -- 0 3,. _ m c - U (D -— (u (D 4— c: 3 (o m (D L U” ,,_ (g. (D U) E -— c 4— 3 o L Q) (0 co :3 0 <0 0 ___\, (q (o *0 o. 3 L — c c — o C _(1 ~ ‘0 0 *‘ (0.- c :3 G.) _C) O _C to o) + -\— .— -— L > + ~~ E c L -— — U (U E (U — +— ~— :1) c :1 3 (D U " m (u o — — C L L m 4—»\+— (U -— O O O L <0 <9 ‘8 0 0 o g (0 (U (U (o (o (o (U _Q _Q .Q .Q .o .0 U 0 (o . . . . . . . . . . . ' .\OL\ 01 Oil OiICCI0:!mlmloqmtododocmdodcmode/ANN“ IIIII 1‘ b .x \ \Ig :1 \ x. \ or on'\ non-J is ‘\ \j \» .\\|\ '\ .\\\.\ \\\\\ y + k R. K. \{. \{. 339 IIIIIIIIIIIIIIIIIIIIIIIIIIIIII IIIIIIIIIIIIIIIIIIIIIIIIIIIIII IIIIIIIIIIIIIIIIIIIIIIIIIIII + IIIIIIIIIIIIIIIIIIIIIIIIIIIIII _III-IIIIIIIIIIIIIIIIIIIIIIIIII IIIIIIIIIIIIIIIIIIIIIIIIIIII IIIIIIIIIIIIII IIIIIII IIIIIII IIIIIIIIIIIIIIIIIII + IIIIIIIIIII traxinea . tlabelliformis . evernioides . tarinacea* R R. tlaccescens R. R. turceIlata' R commixta R. tastigiata . geniculata . continentalis . exilis . cephalota . cumanensis* R. dendriscoides* . denticuiata* . ceruchis . ceruchoides . cochlearis* . complanata' . dilacerata . combeoides R E 3 3 _R_ Q E E. E E E E E 3 E . chilensis . chondrina . crassa . crinalis . curnowii . dasypoga* R. dendroides* R. druidarum . ecklonii 3 E E B. 3 3 l I (Cont.) Table l6. 340 UlAel}OQ!8 — - i .7. IIIII . III-IIIIIIIIIIIIIIII -W" —. . . I. IIIIII IIIIIIIIIIIIIIIIIIII —-___4 2:22:22: ============------- —a eouemns IIIIIIIIIIII====III ' —v eoumsqns IIIIIIIIIIIIIII I" ‘- H eouesqns IIIIIIIIIIIIIII==== I ' spioe 44494 IIIIIIIIIIIIIIIIII IMIIII ll IIIIIIIIIIIIIIIIIIIIHIIIIHIIIIHIIIIL ) IIII g ... I. . IIIIllllfilfifilllfllfififiiflfiiflilIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII + =-—======"===="'===I . u a _ ‘ _ + Y IIllllfilfifillifllfifllflfiflfiflfllIIHIIIIIIIHIIIIIIHHIIIHIIIHIIIIJIIEEHIEIIEEHIEEI ' IIIIlllllllfilifiliflfllilfifilfllIEIIIIIIIIIIIIIIIHIIIIIIIIHIIIIIIINIIHIIflIIIIflIMIIII IIIIlfllfifilfifllfifllfiifilfilalflllIIflIIIIIIIIIIHIIIIIIlflllflllIIHIIHIIIIHIIIIIIII IIIIIIIIliflliilillifliillfiiflllllIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIHIIIIHIIIIHIHII IIIIIIIIIllfilliilillifiiilflifllflllIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIHIIIIHIIII IIIIIlllllflilifiliiflflfififillflfllIIIIEIIIIIEIIIIIIIIIIIIIHIIHIIIIIIEIIINIIIIIIIflII p109 Div-OHSJON IIIIIIIIIIIII II + II eleaoweoeIOJol qolp I III IIIIIIIIIIII IIIIII I IIII III ‘C U m (U L m L +— (D _ U .11“? b L Q 0 Q 4H =--- + IIIIIIIIII IIIIIIIIIIIIIII . . a IIIIIIIIIIIIIIIIIII IIIIIIIIIII + II“ I I p IIIII III IIIIIIIIIIIII p ,3. . (ammo. ”some IIIIIIIIIIIIIIII II =IIIIIIIIIIIIIIII II __=================== _iummlIIIIIIIIIIIIIIIIIII hyPOProToceiraric hoehneliana inflatal“1 infermedla infermediella inanlS ' _...———""‘ Ramalina species R. gracilis* g. R. homalea _— R. humilis l oil oil oil ama 0a cil (steam ml R. R 341 \nndroon‘sl *3 0“ o. \ R. IIIIII ll. IIIIIIIIIIIIIIIIIIIIIIIIIIIIII ' . IIIIIIIIIIIIIIIIIIIIIIIIIIIIII I... ‘ IIIIIIIIIIIIIIIIIIIIIIIIIIIIII IIII l, ,_ _, IIIIIIIIIIIIIIIIIIIIIIIIIIIIII IIIIIII ‘ IIIIIIIIIIIIIIIIIIIIIIIIIIIIII IIII ;. IIIIIIIIIIIIIIIIIIIIIIIIIIIIII ug-III-IIIIIIIIIIIIIIIIIIIIIIIII .. IIIIIIIIIIIIIIIIIIIIIIIIIIIIII IIIII-IIIIIIIIIIIIIIIIIIIIIIII IIIIIIIIIIIIIIIIIIIIIIIIII |:‘."...u-—EEEEEEEEEEEEEEEEEEEEEEEEEEEEEE --'—============================== ”III-IIIII I I+ I IIIII fiEEEEEEEEEEEEEEn"EEEEEEEEE'""' IIIIF:::IIIIIIIIIII=I IIIIIIIII II £94 ================.=:==:======== M 3?? ...-....---------------------- \Sg‘ « \ nervosa coparia iliquosa . subbreviuscuia . pseudosekika . sorediantha* . nervulosa . polymorpha‘ . subamplicata . subasperata' R. obtusata R. pollinaria E R R E. reagens R. reticulata E. roesleri R_ R: scopulorum .E R, sideriza R. R. sinensis R_ R. sorediosa* R. stenoclada E. stenospora R. straminea* R3 strepsilis E B. B . pusilla . sekika Q a c; R. peranceps* R. peruviana* R. pacifica R. pilulitera \\n\ \ \ \ \\\\ ‘ (ALIA if? R. paludosa \'\q\\\.\\~\ \ T 5. E \ \l. V III-....IIIIII lll (Cont.) Table l6. 342 UlAQIIOQla *West lndian spectes. 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