Botany Falkland Islands (Islas Malvinas) Hepaticae and Anthocerotophyta: A Taxonomic and Phytogeographic Study John J. Engcl Sov< nber 30, 1990 Publication 1416 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY Information for Contributors to Fieldiana nc full • luding title lay be •;ls are Museum of Natural I ! . mdard-weight. v nch paper with v, >kette i. Wang PC, Samna, ' or, or --•requested to submit ! lustrations," page. In m the text should I .1." .stem (pen: format ma, " Manual of Style (13th ed.), publishc rsity of liana, Reft ,k and journal titles should be given in full. Where abbrevuc sirable •sistently should follow Botanico-l ihintianum and -nomic i botanical papers) or Serial Soun j 1983) if botanical authors should I • Draft Index of Author the folloui' H. Pennin; A. comparison of montane and lowland rain f< /uador. iructure, ph . and Pioristics. Journ; < gy, 51: 567- '.he Siona: Cultural patterns in visions, pp. 63-80. In Browman, D. L.,and R. A. ;. Mouton Publishers, The Hague, ;iJs. pp. 785-821. jn Steward, J. II., ed., II., >h American itions. Bulletin 143, Bureau of American i hsonian Mcmala. Part II. Polypodiaceae. F;ieldi;i Aith author's name, figure num!1 •v 'Iliis nd ink drav rrcd) .1 copy ^cl of page if the -TV HF THIS PUBLICATION i .iAOID-FREE PAPER. ;PA1GN FIELDIANA Botany NEW SERIES, NO. 25 Falkland Islands (Islas Malvinas) Hepaticae and Anthocerotophyta: A Taxonomic and Phytogeographic Study John J. Engel Donald Richards Curator of Bryology Department of Botany Field Museum of Natural History Chicago. Illinois 60605-2496 Accepted July 27, 1988 Published November 30, 1990 Publication 1416 - X © 1990 Field Museum of Natural History Library of Congress Catalog Card Number: 90-81279 ISSN 00 15-0746 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents PREFACE vii ACKNOWLEDGMENTS vii ABSTRACT 1 THE FALKLAND ISLANDS Location 1 Topographical Features 1 Geological Features 2 Position of the Falkland Islands in Gon- dwanaland Reconstructions 4 Climate 8 COLLECTORS OF HEPATICAE AND ANTHOCEROTOPHYTA 13 PHYTOSOCIOLOGY Introduction 13 Distribution of Bryophytes in Plant Com- munities of the Falkland Islands Maritime Tussock Formation 28 Oceanic Heath Formation 28 Feldmark Formation 34 Fen and Bog Formation 35 Bush Formation 36 Littoral Vegetation 36 Freshwater Vegetation 36 Sheltered High-Altitude Cliff Vegeta- tion 36 Man-Influenced Communities 37 PHYTOGEOGRAPHY Austral Regions 38 The Phytogeographic Categories 41 Distribution Within the Falklands 45 Comparative Taxonomic Composition of the Hepatic-Anthocerote Flora 47 Origin of the Flora 48 SYSTEMATIC ACCOUNT Notes on Format 49 Introduction to Keys 50 Key to Classes and Orders of Hepaticae and Anthocerotophyta 50 Order Calobryales 51 Order Jungermanniales Key to Genera of the Falkland Islands and Mai:dlaman Zone of South America 51 Family Vetaformaceae 63 Family Lepicoleaceae 63 Lepicolea 63 Family Herbertaceae 65 Triandrophyllutn 65 Family Pseudolepicoleaceae 66 Archeochaete 67 Temnoma 67 Family Trichocoleaceae 68 Family Lepidoziaceae 69 Hyalolepidozia 69 Hygrolembidium 69 Kurzia 71 Lepidozia 72 Paracromastigum 76 Psendocephalozia 78 Telaranea 78 Lepidoziaceae Species Excluded from Falklands 82 Family Calypogeiaceae 82 Family Adelanthaceae 82 Adelanthus 82 Adelanthus Species Excluded from Falklands 88 Wettsteinia 88 Family Cephaloziaceae 89 Cephalozia 89 Metahygrobiella 89 Family Cephaloziellaceae 90 Cephaloziella 90 Cephaloziella Species Excluded from Falklands 91 Family Antheliaceae 91 Family Lophoziaceae 92 Anastrophyllum 92 Anastrophyllum Species Excluded from Falklands 92 Andrewsianthus 92 Cephalolobus 95 Lophozia 96 Nothostrepta 96 Roivainenia 97 Family Jungermanniaceae 98 Cryptochila 98 Jamesoniella 99 Jungermannia 102 Family Gymnomitriaceae 103 Acrolophozia 103 Herzogobryum 104 Family Scapaniaceae 106 Diplophyllum 107 Family Blepharidophyllaceae 108 Blepharidophyllum 108 Family Geocalycaceae 109 Chiloscyphus Ill Chiloscyphus Complex Species Ex- cluded from Falklands 122 Clasmatocolea 123 Clasmatocolea Species Excluded from Falklands . .129 111 Evansianthus 129 Heteroscyphus 1 32 Leptophyllopsis 132 Leptoscyphus 133 Leptoscyphus Species Excluded from Falklands 137 Pachyglossa 137 Pedinophyllopsis 141 Pigafettoa 141 Saccogynidium 142 Family Plagiochilaceae 143 Plagiochila 143 Plagiochila Species Excluded from Falklands 148 Family Acrobolbaceae 148 Acrobolbus 148 Austrolophozia 149 Lethocolea 149 Tylimanthus 151 Acrobolbaceae Species Excluded from Falklands 152 Family Schistochilaceae 152 Schistochila 152 Pachyschistochila 153 Family Balantiopsaceae 1 54 Balantiopsis 1 54 Isotachis 157 Family Pleuroziaceae 158 Family Radulaceae 158 Radula 158 Family Ptilidiaceae 158 Family Lepidolaenaceae 158 Gackstroemia 160 Lepidolaenaceae Species Excluded from Falklands 161 Family Porellaceae 161 Family Jubulaceae 161 Frullania 162 Frullania Species Excluded from Falk- lands 163 Family Lejeuneaceae 1 64 Cheilolejeunea 1 64 Harpalejeunea 166 Lejeunea 166 Lejeuneaceae Species Excluded from Falklands 167 Order Treubiales Family Treubiaceae 167 Order Metzgeriales Key to Genera of the Falkland Islands and Magcllanian Zone of South America 167 Family Phyllothalliaceae 168 Family Pelliaceae 168 Noteroclada . .169 Noteroclada Species Excluded from Falklands 1 69 Family Pallaviciniaceae 169 Jensenia 169 Pallavicinia 1 70 Symphyogyna 172 Family Aneuraceae 1 74 Aneura 174 Riccardia 174 Riccardia Species Excluded from Falk- lands 179 Family Metzgeriaceae 179 Metzgeria 179 Order Marchantiales Key to Genera of the Falkland Islands and Magellanian Zone of South America 1 84 Family Aytoniaceae 1 84 Family Marchantiaceae 1 84 Marchantia 1 84 Marchantia Species Excluded from Falklands 186 Division Anthocerotophyta, Order Antho- cerotales Key to Genera of the Falkland Islands and Magellanian Zone of South America 187 Family Anthocerotaceae 187 Megaceros 187 Anthocerotaceae Species Excluded from Falklands 188 LITERATURE CITED 188 APPENDIX: PERSONAL COLLECTION LOCALI- TIES AND NUMBERS 199 INDEX OF TAXA . . 202 List of Illustrations 1. South polar projection showing position of Falklands relative to other austral areas 2 2. Falkland Islands, with principal eleva- tions indicated 3 3. Reconstruction of Gondwanaland show- ing position of Falkland Islands 5 4. Early Late Jurassic reconstructions, one at modern earth dimensions, the other at a reduced dimension 7 5. Gondwanaland reconstruction showing Falklands in Lower Jurassic and Lower Cretaceous positions 8 IV 6. Mean annual rainfall for 10 Falkland stations 9 7. Map of personal collection localities . . 13 8. Photo of Poa flabellata association .... 29 9. Photo of interlacing blades of Poa fla- bellata 29 1 0. Photo of Cortaderia association 30 1 1 . Photo of stream through Cortaderia as- sociation 30 1 2. Photo of wet depression in Cortaderia association 30 13. Photo of stone run as seen from air over East Falklands 31 14. Photo of Bolax gummifera in stone run 3 1 15. Photo of dwarf shrub heath association 32 16. Photo of stream in dwarf shrub heath association 32 17. Photo of Hebe elliptica component of dwarf shrub heath association 32 18. Photo of dry feldmark 35 19. Photo of sheltered high altitude cliffs (arrow) of Mt. Adam summit area .... 36 20. Photo of grove of planted trees (mainly Populus, Nothofagus, and conifers) at Hill Cove settlement 37 2 1 . Photo of interior of planted tree grove (mainly Nothofagus) at Hill Cove settle- ment 38 22. South polar projection showing delimi- tation of subantarctic region 40 23. Map of Lepicolea ochroleuca (Spreng.) Spruce 64 24. Map of Lepicolea rigida (De Not.) Scott 64 25. Map of Temnoma quadripartitum (Hook.) Mitt 68 26. Map of Hyalolepidozia bicuspidata (Mass.) S. Arnell 70 27. Map of Hygrolembidium isophyllum Schust 70 28. Map of Lepidozia chordulifera Tayl. . . 74 29. Map of Pseudocephalozia quadriloba (Steph.) Schust 77 Map of Telaranea blepharostoma (Steph.) Fulf. 79 Map of Telaranea oligophylla (Lehm. & Lindenb.) Engel 79 Map of Telaranea plumulosa (Lehm. & Lindenb.) Fulf. 81 Map of Telaranea pseudozoopsis (Herz.) Fulf. 81 Map of Adelanthus lindenbergianus (Lehm.) Mitt 84 Adelanthus tenuis Engel & Grolle 85 Adelanthus tenuis Engel & Grolle 87 37. Map of Wettsteinia densiretis (Herz.) Grolle 88 38. Andrewsianthus australis Engel 93 39. Andrewsianthus australis Engel 94 40. Andrewsianthus planifolius Engel 95 4 1 . Map of Nothostrepta bifida (Steph.) Schust 97 42. Map of Roivainenia jacquinotii (Mont.) Grolle 98 43. Map of Cryptochila grandiflora (Lin- denb. & Gott.) Grolle 100 44. Map of Cryptochila paludosa (Steph.) Grolle 101 45. Map of Jamesoniella colorata (Lehm.) Schiffn 101 46. Map of Herzogobryum erosum (Carring. & Pears.) Grolle 105 47. Map of Herzogobryum teres (Carring. & Pears.) Grolle 105 48. Map of Herzogobryum vermiculare (Schiffn.) Grolle 107 49. Map of Blepharidophyllum clandesti- num (Mont.) Lac 107 50. Map of Blepharidophyllum densifolium (Hook.) Angstr. ex Mass 110 5 1 . Map of Blepharidophyllum gottschean- um Grolle 110 52. Map of Chiloscyphus austrigenus (Hook. f. & Tayl.) Engel & Schust 113 53. Map of Chiloscyphus elatus (Gott.) En- gel & Schust 113 54. Map of Chiloscyphus leptanthus (Hook. f. & Tayl.) Engel & Schust 118 55. Map of Chiloscyphus semiteres (Lehm.) Lehm. & Lindenb 120 56. Map of Chiloscyphus sylvaticus Mitt. ..121 57. Map of Chiloscyphus textilis (Hook. f. & Tayl.) Engel & Schust 121 58. Map of Clasmatocolea fulvella (Hook. f. & Tayl.) Grolle 125 59. Map of Clasmatocolea humilis (Hook. f. & Tayl.) Grolle 125 60. Map of Clasmatocolea obvoluta var. cookiana (Mass.) Engel 1 28 6 1 . Map of Clasmatocolea rigens (Hook. f. & Tayl.) Engel 128 62. Map of Clasmatocolea vermicularis (Lehm.) Grolle 130 63. Map of Evansianthus georgiensis (Gott.) Schust. & Engel 131 64. Map of Leptoscyphus aequatus (Hook. f. & Tayl.) Mitt 134 65. Map of Leptoscyphus expansus (Lehm.) Grolle . .136 66. Map of Leptoscyphus patagonicus (Steph.) Grolle 137 67. Map of Pachyglossa dissitifolia Herz. & Grolle 139 68. Map of Pachyglossa fissa (Mitt.) Herz. & Grolle 139 69. Map of Pachyglossa spegazziniana (Mass.) Herz. & Grolle 140 70. Map of Pedinophyllopsis abditus (Sull.) Schust. & H. Inoue 140 7 1 . Map of Saccogynidiwn australe (Mitt.) Grolle 145 72. Map of Plagiochila acanthocaulis Sull. 145 73. Map ofAcroboIbus ochrophyllus (Hook. f. & Tayl.) Schust 150 74. Map of Austrolophozia fuegiensis (Steph.) Schust 150 75. Map of Tylimanthns urvillianus (Mont.) Hassel & Solan 151 76. Map of Balantiopsis cancellata (Nees) Steph 156 77. Map of Balantiopsis erinacea (Hook. f. & Tayl.) Mitt 156 78. Map of Isotachis humectata (Hook. f. & Tayl.) Steph 159 79. Map of Radula helix (Hook. f. & Tayl.) Grolle 159 80. Map of Gackstroemia magellanica (Lam.)Trev 161 81. Frullania microcaulis Gola 165 82. Map of Jensenia pisicolor (Hook. f. & Tayl.) Grolle 171 83. Map of Pallavacinia xiphoides (Hook. f. & Tayl.) Trev 171 84. Map of Symphyogyna hymenophyllum (Hook.) Mont. & Nees 173 85. Map of Metzgeria decipiens (Mass.) Schiffh 182 86. Map of Metzgeria leptoneura Spruce .. 183 87. Photo of archegoniophore-bearing plants of Marchantia berteroana Lehm. & Lindenb 185 88. Close-up photo of archegoniophore- bearing plants of Marchantia berteroana 185 List of Tables ? 1 . Temperature data for Port Stanley, Falk- land Islands 9 2. Chronological list of collectors of Hepati- cae 10 3. Distribution of bryophytes in plant com- munities of the Falkland Islands 14 4. Conspectus of phytogeographical cate- gories 42 VI Preface Acknowledgments The study of Falkland Islands hepatics and an- thocerotes which served as my doctoral disserta- tion was carried out while I was a research assistant to Dr. Henry A. Imshaug at Michigan State Uni- versity. Since the completion of my dissertation in 1972 a significant period of time has elapsed, during which we have seen advances in botanical systematics, community ecology, and geology (see sections on "Geological Features" and "Position of the Falkland Islands in Gondwanaland Recon- structions") of austral regions. In hepaticology, there has been a series of noteworthy publications as the result of work on austral groups by R. M. Schuster, R. Grolle, G. Hassel de Menendez, and others. Further, I have continued to focus my re- search on hepatics of the south temperate and sub- antarctic regions, and I have found that my taxo- nomic concepts have changed with regard to several of the groups covered in this study. For example, much of my research has been directed toward two families that are species-rich in austral areas, namely Schistochilaceae and Geocalycaceae. These studies, in large part conducted jointly with R. M. Schuster, have revealed new genera and species that involve Falkland Islands representatives. I have, therefore, reworked my original manuscript to clarify and update the taxonomy, attempting to incorporate relevant literature published during the 1972-1 987 period. Other parts of the manuscript have been sig- nificantly altered. The section on geology has been rewritten, owing to the vast increase in literature published in the last 1 5 years on the timing and relative positions of land masses relevant to Gon- dwanalandic reconstructions. I have attempted to capsulize that part of this mass of literature rele- vant to the Falklands and to potential migratory pathways available in the far south. The section on phytogeography has been emended to incor- porate some of the studies on community ecology. Most of these are relevant to the "Austral Re- gions" section, which contains an expanded dis- cussion of the classification of austral botanical zones. This treatise leaves a residue of numerous un- solved problems. Resolution of such problems, in a number of cases, is dependent upon future re- visionary and monographic studies. In several in- stances I have merely focused attention on a prob- lem, with the hope that the information herein will be useful to future researchers. I should like to express special heartfelt thanks to Dr. Henry Imshaug for his interest, good hu- mor, invaluable guidance, and stimulating discus- sions throughout my years at Michigan State, and for providing me with the opportunity of collecting in the Falkland Islands. I should like to thank Mr. and Mrs. Les Gleadel, Falkland Islands, in whose home I resided while in Port Stanley. Their hospitality and kindness will not be forgotten. Thanks also to the many other individuals of the Falklands for their generosity, hospitality, and friendship. I would like to express special thanks to my wife, Karen, for assisting me at many and various stages in the preparation of this manuscript, and mostly for her continued encouragement. I am grateful to the following hepaticologists for identifying the Falkland Islands groups in which they specialize: Dr. R. Grolle, Jena, Germany (Cryptochila, Jamesonielld); Dr. Gabriella Hassel de Menendez, Buenos Aires, Argentina (Aneura, Riccardia); Dr. H. Inoue, Tokyo, Japan (Plagi- ochila); and Dr. Jifi Vana, Praha, Czechoslovakia (Jungermannia). I am indebted to the late Dr. H. Roivainen of Helsinki, Finland, who determined my Falkland Islands moss collections. His determinations are incorporated in this treatment, and are especially relevant to the ecology section. My continued research on Falkland Islands he- patics at Field Museum was carried out in part through assistance from the Donald Richards Bryological Fund. This support is acknowledged with gratitude. Thanks also to Mary Lou Grein, who has assisted me with most of the research at Field Museum; her care and attention given to this project is appreciated. I also acknowledge the help of Stephen Dercole of Field Museum, who has assisted at various stages, and of Dr. Bertram Woodland for helpful comments and criticisms of the Geology section. I wish to acknowledge the assistance of the fol- lowing institutions and individuals for the loan of specimens: AAS (with special thanks to Dr. P. Lightowlers for assembling British Antarctic Sur- vey materials for my study), BM, CINC (private herbarium of Dr. Margaret Fulford), FH, G, K.RAM (with thanks to R. Ochyra for making available his collections from the Falklands), M, MANCH, NY, PC, S, UPS, and VER. Many of the maps were prepared from tracings Vll by the author; examples of the different kinds are Figures 1, 2, 6, 7, 23, and 51. The south polar projection was adapted from a National Geo- graphic Society map ( 1 943). The non-detailed map of southern South America and the world projec- tion map were adapted from the Goode base map series, University of Chicago. The detailed map of southern South America (fig. 51) was adapted from several U.S. Naval Oceanographic Office charts. The map used in Figure 85 was kindly sent to me by Dr. Hugo Sjors, University of Uppsala. Figures 2, 6, and 7 were adapted from a Direc- torate of Overseas Surveys map. The photographs were taken by the author, ex- cept where otherwise stated in specific legends. vm Falkland Islands (Islas Malvinas) Hepaticae and Anthocerotophyta: A Taxonomic and Phytogeographic Study Abstract On a visit to the Falkland Islands (Islas Mal- vinas) in 1968, ca. 1 ,000 collections of Hepaticae and Anthocerotophyta were made. These collec- tions, which represent the first made by an he- paticologist in the Falklands, as well as those of earlier visitors such as Sir J. D. Hooker and Dr. Carl Skottsberg, were studied in detail and iden- tified insofar as possible by comparison with per- tinent type material. The results of this study are presented in the form of keys to all taxa and a treatment of each of the 1 3 1 species. These treat- ments include taxonomic and nomenclatural re- marks where pertinent, statements of ecological tolerances and requirements, phytogeographic re- marks, lists of literature records, and lists of ma- terial studied. Particular attention is given to the typification of Falkland Islands and related taxa. The ecology of the Falkland Islands Hepaticae and Anthocerotophyta is assessed by arrangement of the taxa into the communities recognized by previous workers. An assessment of phytogeographic affinities of the Falkland Islands Hepaticae and Anthocero- tophyta is made, with special reference to the char- acteristics and differentiation of south temperate and subantarctic distribution patterns. These pat- terns are historically based on the classification of austral regions which primarily utilize vegetation- al criteria. The relative merits of the several zo- nation schemes is discussed and summarized. The distribution of taxa within the Falklands is dis- cussed, especially with reference to available rain- fall data. The Falkland Islands Location The Falklands Islands lie between 51°00' and 52°30'S, and 57°40' and 61°30'W in the South At- lantic Ocean, 520 km east of the Strait of Magellan; their juxtaposition with South America and with other land masses is shown in Figure 1 . The Falk- lands are an archipelago of more than 230 islands, comprised of two principal islands: East Falkland, with adjacent small islands making up ca. 5,000 sq km; and West Falkland, with adjacent small islands making up ca. 3,500 sq km. The Falkland Islands were previously settled by the French, who referred to them as lies Ma- louines. The islands are currently administered as a British crown colony, but claimed by Argentina and referred to as Islas Malvinas. On various Ar- gentine maps, East Falkland Island is known as Isla Soledad and West Falkland Island as Isla Gran Malvina. While recognizing that the sovereignty of these islands is currently in dispute, for the sake of brevity I refer to them as the Falklands or the Falkland Islands. Topographical Features In general, the Falklands are made up of hilly terrain; they lack steep impressive montane sce- nery. The southern portion (Lafonia) of East Falk- land is low and flat or undulating. The principal mountainous areas extend east-west in the north of the islands (fig. 2), with Mt. Kent (458 m) and Mt. Usborne (705 m) on East Falkland, and Mt. Adam (700 m), Mt. Fegan (360 m), and Byron Heights (520 m) on West Falkland. The Hornby Mountains (including Mt. Maria, 658 m, Muffler Jack Mt., 547 m, and Mt. Moody, 554 m) lie in West Falkland and roughly parallel Falkland Sound. The small islands occasionally reach quite high altitudes, e.g., Keppel Island (Mt. Keppel, 342 m), Saunders Island (Mt. Harston, 433 m, Rookery Mt., 422 m, and Mt. Rees, 372 m) and Weddell Island (Mt. Weddell, 380 m). A large number of valleys, small rivers, streams, and bog- gy areas are present on the islands. ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA r" Mas Afuera Juan Fernandez <^ L Mas a Tierra South Shetland Is. » — South Georgia North Island Chatham Is. \ Bounty Is. Antipodes Is. O — Tristan da Cunha Group Gough Is. New Zealand South Island \ Stewart Is. Macquahe Is FIG. 1. South polar projection showing position of Falklands relative to other austral areas. Geological Features The brief and general account of the geological features provided here merely summarizes the major rock types occurring in the Falklands. Greenway (1972) published a comprehensive ac- count of the geology of the islands; earlier, detailed studies may be found in Adie (1952a,b, 1953, 1958), Andersson (1907), Baker (1922), Cawkell et al. (1960), Halle (1912), and Riggi (1934). The Falklands are composed of both sedimen- tary and crystalline rocks, but in vastly different proportions. The crystalline rocks— schists, gneisses, and intrusive granites— are of very lim- ited occurrence. They make up the Cape Meredith Complex (at the southern tip of West Falkland), which comprises the oldest (Precambrian) rocks in the Falkland Islands. Rex and Tanner (1982) recently used radiometric techniques on speci- mens of Cape Meredith gneiss; the measured ages of 953 M.Y. and 977 M.Y. confirm the Precambrian age of this complex. These ages are similar to those from basement gneisses, in part, in South Africa and Antarctica, both of which in Gondwanaland were closely juxtaposed to the Falkland Plateau (Rex & Tanner, 1982) (see below). The remainder FIELDIANA: BOTANY -MT. HARSTON - 433 m. -MT. REES - 372 m. .ROOKERY MT. -422 m. MUFFLER JACK MT. - 547 m. MT. MARIA - 658 m. . MOODY - 554 m. ^HORNBY MTS. BYRON HEIGHTS - 520 m. MT. FEGAN - 360 m. MT. WEDDELL - 380 m. MT. ADAM - 750 m. J MT. KENT - 458 m. MT. USBORNE - 705 m. FIG. 2. Falkland Islands, with principal elevations indicated. of the Falklands are composed of Palaeozoic and Mesozoic sedimentary rocks, with the exception of intrusive dolerites (see below). Two main geological regions may be recognized within the Falklands. DEVONIAN SANDSTONES AND QUARTZITES— These occupy West Falkland (with the exception of the Cape Meredith region) and the northern half of East Falkland. The quartzite is particularly ev- ident as conspicuous parallel ridges that run east- west, parallel to the trend of folds. The sandstone occurs principally in valley bottoms (Skottsberg, 1913). This "Devono-Carboniferous Group" was classified into three main lithostratigraphical di- visions by Greenway (1972), which, in ascending stratigraphical order, follow. PORT STEPHENS BEDS— These are made up of coarse-grained sandstones, quartzites, and quartz conglomerates. In West Falkland they occur ex- tensively in the Port Stephens area; they are absent in East Falkland. Fox BAY BEDS— These are made up of interbed- ded sandstones and slaty shales, occupying "most of the low-lying ground in West Falklands and that to the north of the quartzite ranges in East Falk- land" (Greenway, 1972, p. 13). PORT STANLEY BEDS— These are made up of quartzites and quartzitic sandstones with some in- tercalated shales. In West Falkland these make up all of the high or mountainous areas north of Fox Bay other than the Hornby Mountains and the Fox Bay Range, while in East Falkland they oc- cupy most of the northern half of the island (Greenway, 1972). UPPER CARBONIFEROUS-PERMO-TRIASSIC SEDI- MENTS—Of tillite, sandstone, and clayey rocks, these sediments make up the southern region (La- fonia) of East Falkland. They comprise the young- est rocks in the archipelago and, in addition to Lafonia, "crop out sparsely on the east and north coasts of West Falkland" (Greenway, 1972, p. 15). The only igneous rocks, other than those in the Cape Meredith complex, are intrusive dolerites. Greenway (1972) used aerial photographs to de- lineate between 300 and 400 dikes, nearly all in West Falkland. Only one dike was mapped for East Falkland. These dolerites have been correlated with similar rocks in the Karoo (South Africa) of Lower Jurassic age, but also with similar rocks in Brazil of Lower Cretaceous age (Greenway, 1972). "It is implied that the similarity between South Amer- ican and South African (and also possibly Falkland ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA Islands) rocks may be due to a former close as- sociation of these continents during the Mesozoic" (Greenway, 1972, p. 19). Greenway ( 1 972, p. 9) states that "it is generally agreed . . . that the Falkland Islands did not sup- port a continuous ice cover in the past." However, glacier and periglacier features are present in the form of cirques, nivation hollows, and stone runs, and two of these features will be briefly considered. CIRQUES— "There are approximately 35 cirques and numerous nivation hollows in the Falkland Islands. They are associated with the three highest mountain areas: Mt. Usborne (705 m) in East Falkland, Mt. Adam (700 m, [fig. 19]) and the adjacent peaks, and the central Hornby Mountains (625 m) in West Falkland. They stand out in con- trast to the smooth mountain slopes on which they occur and they are so similar to glacial cirques occurring in formerly glaciated areas of the United Kingdom and of other countries that their glacial origin cannot be disputed" (Greenway, 1972, p. 9). The restriction of cirques to the highest moun- tains indicates that "only above a particular alti- tude were climatic conditions severe enough for cirque glaciers to have formed" (Greenway, 1972, P. 9). STONE RUNS— The extent of the well-known stone runs or "rivers" of large angular blocks of quartzite is unique in the world. Falkland stone runs are a conspicuous feature in the archipelago (fig. 13), where they are known only from areas underlain by quartzite. Falkland stone runs are assumed to be relicts and to have formed during the Pleistocene "under periglacial conditions, when both freeze-thaw and solifluction were much more prevalent than they are today . . ." (Greenway, 1972, p. 11). Stone run boulders "were derived by weathering and erosion of the resistant quartzites exposed in the ridges above the valleys in which they occur" (Greenway, 1972, p. 11). Position of the Falkland Islands in Gondwanaland Reconstructions The Falklands have geological relationships not only with the South American mainland but with South Africa, and this is in part the basis for the different positions of the Falklands in various Gondwanaland reconstructions. Du Toil (1927, 1937) emphasized these joint similarities, and placed the Falklands between northern Argentina and Cape Province, South Africa. He did so on the basis of similarity in structure and stratigraphy of South Africa and southern Brazil, Uruguay, and northern Argentina in South America and the fact that the Falkland Island rock succession is similar and in part identical to both. Adie (1952b), on the basis of structural, tectonic, and stratigraphical evidence, placed the Falkland Islands 250 km due east of Eastern Province of South Africa and placed them in an inverted position (fig. 3). He pointed out that the fold system in the East and West Falk- land Devonian quartzites may be interpreted as an eastward continuation of the Cape Foldings of South Africa. He further stated that the Falklands represented the missing portion of the Karoo Basin in South Africa. Adie then held that the Falklands traveled from this position to their present one and pointed out that this distance would be no greater than that traveled by the Falklands in the Du Toit model. Recent authors for the most part have published Gondwanaland reconstructions with the Falk- lands positioned nearer to South America. In this respect, Bullard et al. (1965) published a recon- struction that included the continents around the Atlantic Ocean. The Bullard model is quite similar to the Du Toit model, but differs in placing the Falklands as part of the South American block and in a position (relative to South America) very close to that of today. Greenway (1972) also regarded the Falklands as having had a Gondwanaland po- sition on the South American block. She cited re- cent geophysical work, including seismic studies, on the continental margin of Argentina which has confirmed that the Falklands are situated on a basement platform, the Falkland Platform. This platform is an extension of the Argentine conti- nental shelf (Ewing et al., 1971), and is "apparently a continuation of the Deseado massif in (Argen- tine) southern Patagonia" (Greenway, 1972, p. 27; cf. also fig. 1 ). Ewing et al. ( 1 97 1 , p. 7 1 30) describe the platform as "a southward-tilted continental block, apparently a submerged portion of South America." Urien and Zambrano (1973, fig. 7) show the position of a basement ridge that forms the margin of the continent south of ca. 41°S and is "supposed to" continue beneath the Falkland Pla- teau. Greenway (1972) maintains that the Falk- land basement has remained attached to the South American continent during the whole of Phaero- zoic time. The Gondwanaland reconstruction of Bullard et al. ( 1 965) is basically similar to a number of models that have followed. These models sometimes differ in the timing of the Gondwanaland reconstruction and its subsequent breakup, e.g., Schopf (1970), FIELDIANA: BOTANY SOUTH AFRICA FIG. 3. Gondwanaland reconstruction showing position of Falkland Islands (= FI) in relation to South Africa and South America (after Adie, 1952b). Light dashed line = limit of Continental Shelf; dotted line = limit of Karroo Basin; heavy dashed line = Gondwanide foldings. Scotese et al. (1979), and Ronov et al. (1984) il- lustrate a Permian reconstruction; Cooper (1980) has a Permo-Triassic model; Dietz and Holden ( 1 970) and Dietz et al. ( 1 972) both have late Trias- sic models; Barron et al. (1978), Norton and Scla- ter (1979) and Smith et al. (1981) have Jurassic models; and Craddock (1982) has an early Me- sozoic reconstruction. Smith and Hallam (1970) also published reconstructions and state (p. 142) "several independent fields of research suggest that much of the initial rifting probably occurred in Jurassic and Lower Cretaceous times, but that much of the dispersal occurred in Upper Creta- ceous and Tertiary times." The maps in Smith et al. ( 1 98 1 ) illustrate this in successive stages. Among these models, the only ones to include graphically the Falklands are those of Smith et al. and that of Craddock (1982). A number of these models show overlap between the Falkland Plateau and the Ant- arctic Peninsula. In more recent literature this overlap has been considered unacceptable, and various alternative reconstructions have been pro- posed (cf. below). A few models differ from the Bullard et al. ( 1 965) reconstruction in position of land masses, and some of these models involve the Falklands. Barker and Griffiths (1977) recognize the continental nature of the Falkland Plateau and close the gap in the southernmost Atlantic in the Bullard et al. model by rotating the Plateau "eastward with South America ... to lie tightly along the southeast coast of Africa as far northeast as Durban" (p. 1 50). Because of the continental nature of the Falkland Plateau and the overlap of the Antarctic Peninsula with South Africa in the Bullard et al. (and other) models, Barker and Griffiths devised reconstruc- tions that involve placement of West Antarctica (portion including the Peninsula) and East Ant- arctica on independent plates. Barker and Griffiths present two Gondwanaland models, differing mainly in the position of West Antarctica. In both, however, the distance between Cape Horn and Elephant Island is small (500 km or less, fide Bark- er & Griffiths) prior to an opening of Drake Pas- sage and the initiation of the South Atlantic with separation of a continental connection. According to this model, then, it appears that a continuous or near-continuous continental connection existed ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA between South America and the Antarctic Pen- insula, the Cape Horn-Elephant Island zone "act- ing as a hinge for rotation of the Antarctic Pen- insula with respect to South America following Gondwanaland fragmentation" (Barker & Grif- fiths, 1977, p. 153). Owen (1983), a proponent of earth expansion, presents a series of maps depicting land masses through geological time. He includes pairs of maps for various periods, one to show reconstruction at modern earth dimensions, the other to allow for size differences of the earth, i.e., progressive ex- pansion. For example, Owen (1983, map 73) shows an Oxfordian (= early Late Jurassic) reconstruc- tion in which the Falklands are part of the South American block and the Antarctic Peninsula is on a second plate (cf. below) oriented northwest- southeast, with its tip near South Africa and its western boundary touching the Falkland Plateau (fig. 4, top). Map 74 (Owen, 1933), however, ad- justs for an earth ca. 84% of present dimensions, and presents an interesting reconstruction. The Falklands are placed further north than in the oth- er reconstruction and are closely juxtaposed to South America, South Africa, and the Antarctic Peninsula (fig. 4, bottom); a hatched line separates the Falkland Plateau from South America. West Antarctica is on its own plate and fits tightly against South America, the Falkland Plateau, South Af- rica, and East Antarctica. The latter reconstruction of Owen has a much closer fit of land masses than nearly all other models. Controversial questions remain as to whether or not Antarctica was a single unit, whether the peninsula was one autonomous microplate or was subdivided into several microplates, and the his- tory of development of the Weddell Sea. Such ar- guments are relevant to problems of overlap of the Falkland Plateau with continental plates. Watts and Bramall (1981), based upon palaeomagnetic evidence, place West Antarctica on a microplate, while Dalziel (1982a,b) and Longshaw and Grif- fiths (1983) believe that Antarctica should be di- vided into several plates. Scotese et al. (1979) di- vide Antarctica into several microplates on their maps. Dalziel (1982a, p. 11) argues that the solution to the Antarctic Peninsula-Falk- land Plateau overlap problem and hence the ultimate solution to the reconstruction of a "perfect" Gondwanaland, lies in recognizing that West Antarctica consists of a number of discrete or semidiscrete microcontinental blocks that probably moved relative to one another and to the East Antarctic craton dur- ing the breakup of the Gondwana supercon- tinent and the development of the southern oceans. Barren et al. (1978), Harrison et al. (1979), and Powell et al. (1980), however, treat Antarctica as a single unit and propose similar models to resolve the problem of the hypothesis of overlap of Ant- arctica with the Falkland Plateau of earlier recon- structions. The reconstructions of these authors place the Antarctic Peninsula adjacent to the west- ern margin of South America, place the Falkland Islands in a position (in relation to South America) similar to that of today, and show an extension of the Falkland Plateau that is closely juxtaposed to southwest Africa. There is considerable evidence to substantiate that the Falkland Plateau was part of the South American plate after the opening of the South Atlantic (cf., e.g., Rabinowitz & La- Brecque, 1979 with Martin et al., 1982). However, work within the last few years has suggested the existence of more microplates, with one in particular involving a Falkland microplate. Some evidence suggests that in the early Mesozoic the Falkland Plateau was not juxtaposed to South America but instead occupied a different position. Recent studies, for example Martin et al. (1981), discuss a Natal Valley-Falkland Plateau fit in the early Mesozoic. Palaeomagnetic evidence suggests that in the Lower Jurassic, well before opening of the South Atlantic in the Lower Cretaceous, the Falkland Islands were in an inverted position ad- jacent to the Transkei coast of South Africa (fig. 5) and formed the southeast corner of the Karoo basin (Mitchell et al., 1986). As above noted, Adie (1952b) suggested such a position to provide for the alignment of Lafonian diamictite of the Falk- lands and the Dwyka tillite of South Africa, which form part of the Cape Fold Belt. According to the model of Mitchell et al. (1986), between 190 and 1 25 M.Y.A., the Falklands rotated 1 80° and moved as a microplate to its Lower Cretaceous palaeo- position nearer to South America. During initial phases of Gondwanaland breakup involving sep- aration of Antarctica from southern Africa in the Jurassic the Falklands moved as a microplate, ro- tating 1 20° to a position ca. 500 km southeast of Cape Town. The Falkland Plateau and islands sub- sequently drifted to their present position with a further rotation of 60° by the opening of the South Atlantic beginning in the Lower Cretaceous. The Jurassic position of the Falklands postulated by Mitchell et al. ( 1 986) may also help solve problems FIELDIANA: BOTANY tft FIG. 4. Early Late Jurassic reconstructions (after Owen, 1983). In the top reconstruction the Falklands are part of the South American Block. The bottom reconstruction adjusts for an earth ca. 84% of present dimensions, and results in a closer fit of land masses involving southern South America, South Africa, and the Antarctic Peninsula; note that the Falklands are placed farther north than in the upper reconstruction. with the fit of the Antarctic Peninsula during the of the Falkland Islands to the eastern margin of Jurassic. In many other models the Lower Cre- South Africa removes the bulge in the craton, taceous palaeoposition of the Falklands caused a opening up a space in the Jurassic reconstruction southwestern bulge of the Gondwanide craton (fig. that may have been occupied by West Antarctic 5). As Martin (1986, p. 101) states, "The rotation microplates." ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA Tierra del Fuego West Antarctic peninsula FIG. 5. Reconstruction of Gondwanaland showing position of the Falklands in the Lower Jurassic (= FI 1) and in the Lower Cretaceous (= FI 2). The dashed line "represents the southwestern bulge of the Gondwanide craton caused by the Falkland Islands in their Lower Cretaceous (FI 2) palaeoposition, which is removed if the rotated (FI 1) position is adopted" (Martin, 1986, p. 101) (map after Martin, 1986). The new evidence raises many questions re- garding the mechanism of movement of the Falk- land microplate to its Lower Cretaceous palaeo- position, and of the identification and position of microplates neighboring the Falkland microplate in the Lower Cretaceous. Mitchell et al. (1986, p. 1 34) state that it now seems unlikely that the Falkland Pla- teau is a simple extension of the South Amer- ican and South African continental crust, and we suggest that it may be a collage of micro- plates. If, as might be expected, all the mi- croplates recognized in this area experienced complex motions during the break-up and dispersal of southern Gondwanaland, this may explain why reconstructions of southern Gon- dwanaland are so problematical. Climate The Falkland Islands lie on the northern edge of the principal depression belt through Drake Passage (Cawkell et al., 1960). These islands are consequently dominated by the westerly winds, fronts, and air masses moving across this region. Although the Falklands are influenced by their po- sition on the lee side of the Andes, the 520 km of open sea greatly tempers this effect. PRECIPITATION— Moore (1968) has assembled the mean annual precipitation figures for those localities with records available for six or more years (fig. 6). It may be observed that the drier areas of the islands are in the south and the wetter areas in the north. The stations with the highest available figures lie on the immediate lee sides of mountain ranges, and a general gradient may be observed from west to east on either island, with Port Howard the wettest station in West Falkland and Port Stanley one of the wettest in East Falk- land. The 609 mm figure for Port Stanley is second only to the 636 mm of Port San Carlos. The Port Stanley data in Moore (1968) may be conserva- tive, as Davies (1939, p. 2) records "in the order of 635 mm and Cawkell et al. (1960, p. 188) record "about" 686 mm for the capital. Moore (1968) points out that the low rainfall of Pebble FIELDIANA: BOTANY HILL COVE 569 mm PEBBLE IS. 473 mm PORT HOWARD 646 mm PORT SAN CARLOS 636 mm SAN CARLOS 520 mm PORT STANLEY 609 mm FOX BAY H? 407 mm <&r J, v^ DARWIN 469 mm FITZROY 468 mm NORTH ARM 399 mm FIG. 6. Mean annual rainfall at those Falkland stations for which records exist for six or more years (data extracted from Moore, 1968). The stations with the highest available figures lie on the immediate lee side of mountain ranges. Hill Cove is located on the lee side of Mt. Adam, Port Howard on the lee of the Hornby Mountains, and San Carlos on the lee side of the Sussex Mountains. The Verde Mountains lie between San Carlos and Port San Carlos. Island could result from its lying in the "rain shad- ow" of the Mount Adam complex. The difference between San Carlos and Port San Carlos may be due to the Verde Mountains which lie between the two stations (or to some other local factor). Cawkell et al. (1960) attempt to explain the slight summer maximum in rainfall figures for Port Stanley through the greater tendency for convec- tion and showers in the northeast part of East Falkland. The air has become moist and unstable after it has crossed the Drake Passage, and it is pointed out that an unstable air mass approaching the Falkland Islands from the southwest will pro- duce only cumulus cloud cover at Fox Bay and Goose Green. However, after this air mass has been sufficiently heated from the land and perhaps been uplifted by high ground west of Port Stanley, it is able to produce showers in the northeast re- gion of the islands. It is likely that this phenom- enon of heating and uplifting is occurring on both island groups and accounts for the data obtained. The rainfall is distributed fairly even throughout the year, with a slight maximum in summer (De- cember-January). The number of days with rain- fall is high (ca. 250 days) with few days of excessive rainfall. The rainfall intensity is thus low, and daily falls exceeding 13 mm are rare (Cawkell et al., 1960). The Falklands have no permanent snow cover. TEMPERATURE— The temperature data for Port Stanley are shown in Table 1 . While temperature data are available from only Port Stanley, Fox Bay, and Westpoint Island, Moore (1968, p. 7) states there is a "suggestion of increasing temperatures towards the west and south of the islands." HUMIDITY, CLOUD COVER, AND SUNSHINE— Cawkell et al. (1960) indicate that the relative hu- midity average values are high; the mean annual value is 82% at 0900 zr, and varies from a mean of 75% in November to 90% in July. The Falkland Islands witness a large number of overcast days. TABLE 1. Temperature data for Port Stanley, Falk- land Islands (after Moore, 1968). Mean Mean Mini- Maxi- mini- maxi- Month Mean mum mum mum mum January 8.8 0 23.9 5.4 12.9 July 2.2 -7.8 8.9 0 4.3 ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA e — 111 5*1 •£ •= ± E V) a o * £ £&s E Sd 0 s" 0 s s 0 0 I S §i OO A A 3 3 I f ^ S 3 3 O h 3 0 ^ oo o u 'C w 'C p c ° la J3 > ^ OO /—v O £• * .s /3' ^-v 2 ^ •^ ">> ~ (O (A - ,- M O — .2 •o oo H 5 S ^ "o. cS ^ ftOv 0^ - 3 ^^ s> ^ r« Cu e QJ 5 ^^ ^. I ea u •3 s u u H II C T\ ^ O *^ C rt ^ o — — — 3 o 11 — t/5 *O "^ e 3--S E « u O o DC u u 55 55 « \ O S~ •s W e Z *» •* oo oo m *o & i •- o i_ rt O 1 * o ! | 1 1 § ist of co on i| i i"l S s| | .. s 1 J •| rt « u ^d I c/5 || || |5 u "o U I PJ (X) cS jl U Q C Cu u O O C S &< S3 ^ D w ^.1 l| ^f u;-0 > UJ U oo 1. 0 — a 2 p *l-ll e ^^ ^ fN 00 O xi f^' o y ^ . 1 O e U U O BJ eg •o c i— » £ u ^. E* g?h< JS (j c u ! « O lu a" X — 1 2 '5 X) C X> 3 < U 1 1 = C/5 C/5 Bf* O « Z * ' lllfie "tt-ilJM I s I5J w « " HH ^_ 5?^ £3 - -^ 'Z^ o .. w g" &&•§ 05-33 [111* i <^ 2 £ a ute • «•§ b-g ^ u s x« er o for r 911) 03 u ? s-^ 2i g I g III '55 rj w «i v-/ — 10 FIELDIANA: BOTANY g - Herbariui (original collection Sill *,_ 3^1 a"-- !« ? L V} „ r • 1 1? I i 1 3 i*-* 2 -C 3 WJ ^^ 3 3 3 • X 3 T^ a a a ; •gl § § c < f a£ t f 3 .2 — .S o o .0 i II I 1 1 i j j i i •s 5 oo 0 z 10 " «* — oo * PA uT TABLE 2. Continued. * H 3 O t> "— ' "alas- rial's £••0 U 8 o^ 2 P a- -2 |J1^ tf dc§- a illll Si«l illll l£lf fljst tlt-1 o »S >,-o ff « > a — •£'•> "5 S c j= o « -b^o.;^ .-OUU55 ^j J2 |1 11 ^^ 8lfi III j-j s = iij en >;,j ISs Port Stanley area: Mt. Wil- liam Port Stanley area § «s i o J o 5 — 2 2 o o •5 a ^ i >' • c>\ 1 ^ I I 5 2 2 O «N * o . c , « JT »r "2 .i ' « s •S3^o = «^«e=*'a' •- * c 5 £ - § .0 2 o J= .S .S 'o -S '§ | "a -5 if! S |8 1 11 I !j;»tf*'".cll5! •O r-*O.S"O J^OSOO-O c- O "O w— c15 « x^^o C'S'o c-= . I^lal522|i3ll JtilpI fii •8as--S> anal 5l|II > > u, u ad P u * • * • e w -^ PC — B M — ' 03 C/3 • <5 I !- 1 G .2 •£ | 1 1 co r X 03 Ou H o 1 ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 11 Herbarium (original collection) (specimen in LIL, fide Hassel de N (U 4> L C/3 ^. UH ^ 08 by Gaudichaud v •o g J I fi'S 1 1 o •g .2 -S .§ • 3 11 3 3 'B 3 -.3 3 3 3 a £> a a a *w B 3 C c B .2 3 a 3 3 — «£ f If 3 3 g .0 —-.2 0 O a) ^ CD ^ ^ ^ B £ u £ £ 1 u i en CO to ^ 2 ^^ ^ • - 4> « »2 o y _^: _*^ u •M4 ¥ U u d ^H n a a a •s oo 00 a Tjr ^\ s-^. ^^, 00 1 ON CN * 2 * T a 1 1 "S ^^ ^-^ 00 S 3 " >v >» ^ 0 C 1 '^"c u d o en O a M S £ ^e o 1 fN •a c3 — » (4 S P | 1 £ o «- 0 [T | m — oo w * -»-- 12 FIELDIANA: BOTANY FIG. 7. Map of personal collection localities. The numbers indicate regions or islands, which are as follows: 1) Kidney Is., 2) Port William (includes Mt. Low), 3) Port Stanley (includes Mt. Kent), 4) Mt. Usborne, 5) Darwin Settlement, 6) Port Howard, 7) Fox Bay, 8) Mt. Adam, 9) Hill Cove, 10) Westpoint Is., 1 1) Weddell Is., 12) New Is. The average annual duration of sunshine is less than 35% of the total possible for the latitude of Port Stanley (Cawkell et al., 1960). Collectors of Hepaticae and Anthocerotophyta The most significant collections of Hepaticae and Anthocerotophyta are those of Hooker, Skottsberg, and Halle, while those of Lechler and Cunningham are important but fewer in number. Table 2 shows a chronological list of collectors, together with pertinent data. A list of my collection numbers together with collection data may be found in the Appendix (p. 1 99). A statement of the ecology is made for nearly every locality, since ecological data have been omitted under the specimen citation portion of each species. The localities are indicated on Figure 7. All localities are cited with reference to Uni- versal Transverse Mercater Grid (Zone 2 1 ) coor- dinates as shown on maps of Falkland Islands 1:50,000 (Directorate of Overseas Surveys 453, Series H791, 1961). In the Systematic Account, the specimens I have personally collected are indicated by number only, without citation of collector. A complete set of my collections has been deposited in the Cryptogamic Herbarium, Michigan State University (MSC). All other specimens are cited with the collector's name provided. Phytosociology Introduction Skottsberg (1913) made a vegetational analysis of the vascular plants of the Falkland Islands, and Moore (1968) based his account of the Falkland vegetation primarily on that of Skottsberg. Brief accounts of the natural vegetation are also pro- vided in Davies (1939) and Cawkell et al. (1960). In summarizing the bryophyte sociology in the Falkland Islands, I have utilized the community groupings and terminology of Moore ( 1 968). I have, however, added one community grouping, i.e., the sheltered high altitude cliff vegetation, which is primarily cryptogamic. As emphasized by Moore the predominant Falkland plant associations are closely related to one another. The associations and formations, especially those of lowland areas, frequently are not arranged as discrete units, but ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 13 ill X X XXXXXX XX^X § O H .. "G . •. ; •g j» S *•• i ^ s X X _z h 1* 8 u!8 t f X 5 |i w XX XXX XXXX ^tX _- * ft t2 I .*'. ''" r — C — (. A < •5 £ o. * 'S X S O fa •o E g vt o 2l X XXX X XX •o g C C CO 2 2 — "3 ^ ^ ^ g* £ — 01 y B •S S • 11 XX^ XXX X X S 1 A 'S 1 •a e g o .2 ||| o I XX X c I ft ^ - CO •^j ** "a e •S V J? 0 o s 'o a (J ^5 Cb o CM O ^ e. 0 '3 5 S"S « X O fl S £ II 'i XXX XX S Uta f> w e = m s- Jo .S M 'C 3 "** 88 gfl H •S( - 0 * X XXX - " 5 5 111 •S .2 a a c, •= t? •> ^ s 3 w *^ "^ C S ! hlli J | .1 giiiji iiii|iiiij|i|ijliii|| i o 51 I'll J I lljl 1 1 11. 1.1. 311 Sllllll 1 d> ^j O "^X "Xs *^J StCCS.2»QQCS *Si "Si "Si ^ ^* ^ "C "^ "51 "C *5J "51 "51 *5J tU ^s ^J ^C ^J ^J ^J ^^ "^3 ^C ^C ^Q ^5 ^3 ^Q ^Q ^5 ^^ ^^ ^^ ^^ ^^ ^J ^^ ^J ^^ ^J ^^ X 14 FIELDIANA: BOTANY i Is 5 :_ — C i — — fc« XX i? |j a. SI XX XXXX XXX XX X X e o -a js 5 2 o o 3 $ * 'c o S -2 ||c 1 II i5 I & & ft Jl 8 C 51 *^ %*••• — O • v IE • •« R 9 *K 1 j- 1 ^ | '^ < ° 3 O -, _ _ tsS w •5 "tj C 013 - r C m 43 * * > ™ ^ V § , ,.2 p | IIJjJ 1 0 ^ |J cu cu cu o, a. a. •w E uu ul, uu u- u, uu u. !*» o S SS S S S S fa fa 1 o c II D o s z 3:i III ^^ i LANT COMMUNITIES S i i ||l! iiuliiliiii liHliiil {fill ililli ililili! § IIIIJI1 3'§333^§§«1*'S' S. S.-§'-l'-l'"§ "~ -S g-^-^-c-c-c-*:-*: = Mesic feldmark 0. « ^ "^-"i "i"i ^"^ ^ k-2'§'§'3aqQ-S'^-S'^^v^^'^^>^v&' M i- •Q'^^^^a fc! ^!^§ S § S"S'S"S-«'«'«:§-S-2.S.S.S.S^ cu Cu bb"^"^"QCCct3"a"5"3-Si-Si-Si§'§' S1-*: •*: :«: •*: pC -c -s: -s: s U ^!'^J'^^'J'^!'^I^I^^^j05^Q^9^Q^Q^Q>J>^>J^J>J>J>J>J>J>J>J X ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 15 !|-. Xir !i^ ^r ^* ^^ ^^ »5: *^* *^ ^^ *^ *^ ^^ ^v1 ^ ^* ^ ^* i^» ^* ^> t^ t^ »^» »^ p*S ^^ rN S j£5 •s .2 ll X X S -|8 B ll'i XXX XXX Xs X X i o • s *• 4» B * •5 $ I'l'S X i Q ta S g e 2*PM B i* ^ X X X X XX a ^0 *^ o £ _ 5 si X Oi « fc X XX j 5 1 H 1 a •o V o •s 8 -g 3 .2 C O 's X»X X C Si - - U •< g .A, .2 04 »*i h« 3 1 «* c s < S 2 (5 '" XX X X H u S ta - S e S ^ a .2 ™ XX X E « 8'S te A X X X X X X gta 2 1 a § | ill I X Genus, species •a « *» (a ^ i I !! Iiil Li c §^ a-d .S?t ^"S3o2^?««SaStt^ "«o>< o ^ "5.^ § e ,. « I o ^ '5 e •C ^eo Sj|* 4-. o = fa i B-2 2 1 8111 — E I a Si 2 .2 § 2 a ^i "3 § & '* g25g fcC tjouC)C;GCJC;ocjQ^tt.k.t*,^^i;a;a;a;a:a:^^^ ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 17 XXX^ X XX X XX^X X^X XX XX X X XX XXX X X X X X X X X X X X X X X XX X I fc-2 II llll mi *, 111 » i -itila J P" 3 S g 11!!! mil 18 FIELDIANA: BOTANY 2 X Is 5 M 8 1 i u X X 21 H r o s I* x a 4 £ £0 e XXX X XXX 3 $ -5 cc u! •£ 4» >*" >• 23 e If* 1 3 1 s * * .2 ^j •d 0 c ta 'si — .g 5 .2 e M a 2 «Si 8 •* C — (j h $ ** 1 • i wn ' - ^ -. e a M A> -s - ~^ * g g .^ % § ^ •g 5 ^ o £ 'S ^p M >3 ^ c *> 3 S 5*5 < O b -2 .2 £ M "5 £ § 2 s !|* 1 t> u C i •i ° •- H s 1 1 § 1 fa a«^ 8 - II -^ o X S -5 CU 0, 1 i 1 U, LL U, U- bu Lu It, SXS S S SS 2 •*" w 0 'a) V E O LLJ u. Z 1 ! III 1 1 • COMMUNITIE M I sic feldmar I S c^l^S^"5 ^"^"tllli ^^"S a"^ 1 a| ^-11 1 CJ S Q. fills!! Illflliilllllfilili] II Q. U, 5 ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 19 "2 8 -a c t •? XX XXX X X X XX X X X | SH .2 S 1 3 • x xxx 1 S ill X XXXXXX XXX XX | oto 8J ~ (& B Q..S *3 -x s Q ta Ii § £ w X X X X X X XX •s C«ta E •2 1 |fc~ s j|| x x x xx OH '= •6 e - a mtinue *|l X 0 .2 'C ' •o a ! rti QA I i<| X 3 u t^ fe H i w ^|«'| x xxx g », S'l XX XX XX 4» ^ s S E- < 2 S » « '5 ^ ' 08 1 b. < SI 1 1 1 «-ct *enus, species (^, ^ K V) ^ *£* •* O ^J "*• g^^'s;aa -^ g .5 ^ :a "i -S c B a illliliiiill !ltlfil]]fttl!!l|U i « ^f I1 8 1^1 i e 3 ^31 i 1 IS |l 1 1 1 ^ i 1 *^ | S w 111 Illlliilii § H § 1 1 1 1 1 illllllll a, ex, ex, a, ^ 5: 5; 5; 5 5; 5 ^ ^ 5 5 5 5 5 5 £ 5 £ 5 5 £ £ £ £ £ ££ FIELDIANA: BOTANY 1 5 5 - 3 '5 8 C 3 eg c = S H 5 ° Ou " -F - 2f •— XX XX XX X XX XX X X go e o •5 - r 3 S « Jf *£ |s B 3 > *" e ... i c .» '5 c •s * 5 8 | 41 8 « o ta i B.2 •S 'c a k S • •— (j w w ea S 'S ^ ** I ijlli 1 "^i. s, *> * ™ — -^ 2 •o * • <3 .2 c g *s g § U, a; .§ % ™ 1| = « 1 j - - - - i 1 «S H 5 PLANT COMMUNITIES, c< f 1 II I3I 1 8 :=il III ^ ^is ^-SS acjfc1 -asaS.o-^Q^s ^ -SSs •§-«c MF = mesic feldmark. Hiilllllttlalliiliilslf Ilill-l-! Ill iJllllllJJ 8 8.8.8.8.5.8.8.8.5.81 1 1|| || ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA *b X XX^ X X X^X X X a #H | *l XX X •< ft; £ ja «* — ~ s _t = C K ^ H <3 *G OS = r* XX X XX X XX X 1 ° 03 IP SG >!*! x | li XX XX to ! 111 XXX X 1 c i .2 a 3 I If 1 X XX c J" O ^ a •s i I a 8 • JB C ~ '-/ OD ^^ bb X H £ £ l|l| XX X 1 8 eu XX X 1* § § 111 « * *C ^ S 3 X 1*1 1 5 D. 3 «» s | 15 i -s i|a il Pi I j fill fill III ^< ^< ^< ^< ^^ ^* ^< »S \^ ^ij ^ "^ '^ 'v ^^ TP ^^ QQ QQ (to QQ Qs QD QQ QQ 02 fto (to ^i S 22 FIELDIANA: BOTANY | XX X 1 :! C : -j •S 1 H Jo " 4 11 X I" — (c X XX X X £0 e ••8 - t a s« a> fc. <* 00 - ^ • ll B o ^ •- "-C ^> Q O 1 ^ I ~ o , S3 8 • J •* '£* $ '~ OB U £ ^2 •» *» »1 « c a y 4» -5 e B • 3 x 8 — •• &i •>« O O a •* -2 * ~ b .2 .2 ? "5 ll » -T ^ * § .2 -if c *5 *? i! ^ CO «» O O * 'S E < 2 o -t B S .2 § 2 | X X II U. (x. u, X •o c o J2 UJ c | 5 ex 3 «« 1 § 5 ° ? s PLANT COMMUNITIES, c ! ill WG = The Woolly Gut. W 5 ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 23 ** S 5 xo x x x x x x><0x xoxo e (5 H o •S .3 M i > C8 X •< 33 - £ Q, —L ® £ EC III xxxxx xxxx x 2 , 15 Ifl'S X 1 " i i | S 'S X XX i o JS A w 's u< J> "3 4> 4> &• A c C -r 5 o 3 « e * - 3 S * x xx Sf c 5 Ss J? a a a H 1 t I if, f ill i &iifni|tliti$lii i Sii t tn-| « ~S3Ra.'^'c:5'Ca<3a|35jQ-~=5=5 ^CX-^-Q'KX S S"SSis: 1 lil|iliillllll!ii till! t fill I ?l:|«lllllRellli'|-| -||lls 1 l-li ^3 (S^Scolalllliliill Islll 1 1 SI FIELDIANA: BOTANY Is " S3 . a c 8 C 3 « 1 H M O J 5 ** <5 L- _ 11 X X I J £ - - sc ._ X X 330 o 'S -c s a/" _ "-* A > — i C~j •« ^ ^ E ^2 2 •*» 5 e 8 1« o "• i 5.2 js ^ 5 5 e S'2 v» g .2 •*• "s* S2 *- BO U * ^ C3 i2 J> *> i •I SlUi <= e "S r** *• J? U c c^ ^, •c e . i E u, .9 .2 K Ct ^ £ ; II £ S «'s U. ^ J? X « ^«.2e •o g 1 1 1 J J2 b °S-S| S — 1| I Z "S o 111 g 1 o J UJ D ^ Z M r™ i s si ^* 0 3 Q K COMMUNITIES, O £ 1 i « 1 11 ll'sl 3 '|'| a, 1 1,, iiiiiiiiiiHi «i if i- I 111*11 ill ill 11 814 IIjiKil 2 all l| | i|tlltl 1 '111 J jiff llf feldmark; WG | J "§ .3 §••- § •§ •§ 5 Jf-S ^ ^ ^ -2 .2 1 "§."§."1. 5"5"5^"s't8p« u 1 C Ilii^lllfllllll ^llli lllii -ll E u "5 Q Q Q fe -5 -C O 5v55v55vvvvS Sj Sj .•? Ss S\ > §• .5 U. OUOUUUOUQQQQQQQQQQQQQQ^aja: ^^ z ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 25 e ?ll $H xS x^xxxxx x ^ > o •a -C ll x * 1 ufji O i— Srt XX XXXX X X X) * ft * i « I a J •g > o. ° "^ XXX £ *£*& g || X XX •a 0)ta e - -= ~, 2 *- h 5> ^3 / •=. 1 il $H X X XX I u O e o • •- °« , "8 * -t- - L 3 c c 1 80* O -A 2 "<5* X o ^^ 0 at *C •e a o a» CA 1 g J O 3 o < U 55 t2 H « § a JE ^ ft .2 *2 5*1 E "2 3 e 2 1 8« 0 3> 0 , X 1 ll§ II fL X ^5 •a o c fe i 6.2 •s lliJ 5 a S *9 • 9 — U »5 4? "^ 1 c a M 8 -2 _ to 0 o B la 8 I ^ •£ K ^ '5 g .* **^ *r ^ l|§ >. 8 " ^ ^ ^ a i o ^ .0 .2 p S S *• o o £ * 3 I « XX i o 11 cu i i] Uu U. _ - _ E M S S X5 x ,"o _ ___ U S — , E o 1 UJ U. I S PLANT COMMUNITIES, cc Genus, species 2 II l| H |j i lli ill fill sit !t|l! iff! ilillflillliiiiiiiili * = Mesic feldmark pool i-« u- S ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 21 a kind of mosaic of community groupings is rather the pattern. Moore (1968) has recognized five formations. In addition, he has recognized four associations (Senecio candicans, Ammophila-Elymus, Eleo- charis, and Myriophylluni) which do not merit for- mation grouping. These are arranged according to types of vegetation. Distribution of Bryophytes in Plant Communities of the Falkland Islands Distribution of bryophytes in Falkland Islands plant communities is shown in Table 3 (pp. 14- 26). MARITIME TUSSOCK FORMATION— Poa flabel- lata Association— This primarily coastal associa- tion was previously far more prevalent than that observed today (Hooker, 1 847; Moore, 1 968). The unrestricted, extensive grazing of sheep has been primarily responsible for the depletion of this for- mation. Pigs, which destroy plants by digging up the roots, are of lesser significance. Tussock grass today is confined to smaller offshore islets (e.g., Kidney Island) which have not been stocked with sheep, and a few fenced off areas on the larger islands. On Kidney Island, where there are exten- sive areas of almost pure stands of Poa flabellata, this association has reached its maximum devel- opment (fig. 8). As discussed in Moore ( 1 968), Poa flabellata tussocks can attain a height of 2-3 m with each tussock composed of a fibrous stock (1.5-2 m high, 1-1.5 m in diameter) surmounted by a dense crown of leaves. In areas where the community is uninterrupted, the dense canopy of interlaced leaves excludes all associated vascular plant species except Carex trifida (fig. 9). Telara- nea pseudozoopsis and Chiloscyphus lentus were particularly common on the tussock bases while C. leptanthus, C. sabuletorum, and Hygroambly- stegiumfuegianum var. skottsbergii were found on soil between the tussock bases. Poa flabellata is well-developed and forms a monospecific grass- land covering two-thirds of Beauchene Island, the most isolated island of the archipelago (lying 80 km south of East Falkland Island) (Smith & Prince, 1985). Chiloscyphus lentus was found to be the most frequently occurring hepatic species on the island; it "grew around the moist shaded bases of occasional tussock pedestals and on inter-tussock mounds . . ." (Smith & Prince, 1985, p. 242). Oc- casional associates were Chiloscyphus divaricatus, Clasmatocolea vermicularis, and Telaranea pseu- dozoopsis. OCEANIC HEATH FORMATION— This formation has been subdivided into two groups. Cortaderia pilosa is dominant or codominant in one, and dwarl shrubs are prominent in the other. Most of the Falkland Islands are covered by one facies or another of these associations. Cortaderia Association— This association, as Moore (1968, p. 15) stated, "can be developed on most nonswampy ground having indifferent drain- age. It is widespread on level or undulating country below 100 m ... but it is also very common on gentler slopes up to an altitude of ca. 180-200 m" (fig. 10). I have found this association, which is dominated by Cortaderia pilosa, occurring at var- ious altitudes up to ca. 610 m. The grass blades form an excellent protective cover against the sun and especially the winds, while the coarse fibrous peaty humus which develops under the tussocks retains considerable moisture. These factors con- tribute to a protected environment in which a sur- prisingly large number of bryophyte species may be found. Bryophytes directly associated with Cor- taderia, in depressions, pockets, or holes, and de- riving protection and moisture from the associa- tion are listed in Table 3 under the General Terrestrial column heading. This association in- cludes several facies, each differing in the means of providing protection and moisture conserva- tion. These facies will be discussed separately. The Stream Facies— Streams are frequently present in the Cortaderia association (fig. 1 1); often the bank vegetation (which frequently is Gunnera magellanicd) or the stream itself offers protective niches for bryophytes. Chiloscyphus austrigenus is very common here and, together with C. elatus, is the only hepatic that is nearly always submerged in these streams. The Wet Depression Facies— Wet depressions, where the peat is saturated and small pools may occur, provide another means of protection for bryophytes (fig. 1 2). The Stone Run Facies— Stone runs are of fre- quent occurrence in the Cortaderia association (figs. 13-14). Often, bryophytes occur in cool, damp, shaded crevices. The Rock Outcrop Facies— Rock outcrops (par- ticularly those having exposures protected from prevailing winds) are the sites for 12 species of bryophytes. These plants were found associated with overhangs, crevices, and small niches, and on the soil immediately below the outcrops. The Cortaderia association, while not possess- ing the bryophyte species diversity of the closely related dwarf shrub heath, possesses the greatest 28 FIELDIANA: BOTANY FIG. 8. Poa flabe llata associa- tion at Kidney Island. FIG. 9. Interlacing blades of Poa flabellata at Kidney Island; note head of standing individual for scale. Photo by H. A. Imshaug. ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 29 30 FIELDIANA: BOTANY Above: FIG. 13. Stone run as seen from air over East I . ilk lands FIG. 14. Bolax gummifera in stone run. Opposite: Fio. 10. Cortaderia association near Mt. Usborne caravan. Photo by R. C. Harris. FIG. 1 1. Stream through Cortaderia association near Mt. Usborne caravan. FIG. 1 2. Wet depression in Cortaderia association in gap between French Peaks. ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 31 17 32 FIELDIANA: BOTANY assemblage of rare Falkland species, several of which are also rare in southern South America. Dwarf Shrub Heath Associations— Moore ( 1 968, p. 1 5) stated that "The communities included here are usually found on comparatively dry ground, being best developed on rocky ridges, stony areas or places where the immediate subsoil is of rela- tively coarse material so that drainage is good. A hard, dry peat underlies well-developed heath." The principal vascular plant species in the dwarf shrub heath are Empetrum rubrum, Pernettya pumila, Blechnum magellanicum, B. penna- ma- rina. Baccaris magellanica, and Bolax gummifera. Various combinations of these may become co- dominant, and Moore (1968) has listed the fol- lowing associations: Empetrum rubrum-Blech- num penna- marina, Empetrum rubrum- Blechnum magellanicum, Empetrum rubrum- Pernettya pumila, Bolax gummifera- Empetrum rubrum, and Blechnum magellanicum- B. penna- marina. Moore (1968, p. 15) stated "The Empetrum rubrum as- sociation is much the most common" and the dis- cussion below pertains to this association. As in the Cortaderia association, there are several means of protection against the sun and winds as well as the conservation of adequate moisture so that this association has a fairly rich bryophyte flora. Each local facies will be discussed separately. Bryophytes commonly grow on the bases of Blechnum magellanicum, the fronds of which pro- vide an excellent protective cover, particularly when interlaced with fronds of other Blechnum plants or branches of Empetrum (fig. 1 5). The soil below such situations often has a considerable bry- ophyte cover. Frequently, Bolax cushions, partic- ularly old, decayed portions, have a bryophyte (mostly hepatic) cover. The bryophytes in these situations are listed in Table 3 in the dwarf shrub heath column labeled General Terrestrial. The Stream Facies— Streams often occur in the dwarf shrub heath association. Vascular plants such as Blechnum magellanicum, B. penna-marina, Empetrum rubrum, and Gunnera magellanica fre- quently offer a protective cover, which may be very dense, and at the same time provide cool, moist niches for bryophytes (fig. 1 6). Several species grow partially or wholly submerged in the streams, Chiloscyphus austrigenus and C. elatus nearly ex- clusively so. This facies, when compared to the stream facies of the Cortaderia association, while having the same Chiloscyphus species, has quite a different species complement, as well as greater species diversity. The Wet Depression Facies— Like the Corta- deria association, the Empetrum rubrum associ- ation has shallow depressions where the peat is saturated and small pools may occur. The habitat has a characteristic bryophyte flora. Sphagnum falcatum was present in depressions where free- standing water occurred. The Rock Outcrop Facies— Rock outcrops of varying sizes are frequently present in this asso- ciation, with protection provided in a manner sim- ilar to that in the rock outcrop facies of the Cor- taderia association. Bryophytes occur in the various niches associated with the outcrops, as well as on soil at the base of the outcrops. The Hebe Facies— Hebe elliptica, when occur- ring as scattered bushes, may be regarded as a relatively minor component of a dwarf shrub heath (Moore, 1968). A heath of this type was visited in Waterfall Valley, Westpoint Island. The Hebe shrubs occurred on steep slopes on the valley sides (fig. 17), while through the valley bottom flowed a rather fast-moving stream, with Gunnera ma- gellanica forming a dense carpet along the water- course. The sides of the valley were often clifflike, with numerous damp, shaded, sheltered ledges and overhangs. Bryophytes occurring under the wa- terfall or in the spray zone were Megaceros fue- giensis, Riccardia georgiensis, Telaranea pseudo- zoopsis, and Webera albicans. The bryophytes associated with the stream bed were typical of this habitat and consisted of Chiloscyphus austrigenus, C. elatus, Riccardia spectabilis, Telaranea plu- mulosa, T. pseudozoopsis, and Sciaromium con- spissatum. Hebe elliptica also occurred on cliffs facing The Woolly Gut on Westpoint Island. The cliffs were rather moist where numerous crevices, ledges, and overhangs were present. Three hepatics— Chilo- scyphus semiteres, Megaceros fuegiensis, and Metzgeria violacea— occurred exclusively in the Hebe facies. Summary of Dwarf Shrub Heath Hepaticae— The dwarf shrub heath associations have the high- est number of hepatic species of any of the Falk- land associations (75), and of these 17 are unique to it. Of those that are not restricted to this as- sociation in the Falkland Islands, 49% also occur Opposite: FIG. 15. Dwarf shrub heath association near Mt. Adam. FIG. 16. Stream in dwarf shrub heath association near Weddell Island settlement. FIG. 1 7. Hebe elliptica component of dwarf shrub heath association in Waterfall Valley, Westpoint Island. ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 33 in the Cortaderia heaths, which is indicative of the rather close relationship between the associations. However, I regard the bryofloras of the Cortaderia association and dwarf shrub heath associations as sufficiently distinct, and the differences aid in dis- tinguishing the two associations. The dwarf shrub heath associations and Cor- taderia associations are vegetation types which have been highly altered both through grazing and conversion to pastureland, a process which fre- quently involves burning and usage of introduced grasses (commonly Holcus lanatus; Davies, 1 939). Even in the most severely altered state of the oceanic heaths, I have found it possible to deter- mine the natural vegetation type through exami- nation of the bryoflora of stream banks. The stream banks are somewhat less disturbed and here the natural vegetation is least altered. See further com- ments under Man-Influenced Communities, at the end of Phytosociology. FELDMARK FORMATION— The feldmark (= fell- field) formation has been characterized by the fol- lowing four features: 1. Considerable areas of bare, exposed, usually mineral soils with scattered small rocks or boulders. According to Greene (1964), feld- marks have a vegetation cover of one half or less the total area. 2. The tendency for the cushion habit to predom- inate. The feldmark cushions do not form a continuous layer of peat (Wace, 1965), and do not have among them the cushion bog1 dom- inants of other associations (in the Falklands the Astelia association). Likewise Azorella, the principal feldmark cushion species, does not occur in cushion bogs (Wace, 1960). Wace (1960, p. 486) listed the principal species for feldmarks of the following islands: Falkland Islands— Azorella selago, Bolax gummifera Kerguelen— Azorella selago, Colobanthus kerguelensis, Lyallia kerguelensis Macquarie— Azorella selago, Racomitrium crispulum. Dicranoweissia antarctica Tristan da Cunha— Empetrum rubrum, Ra- comitrium crispulum, R. lanuginosum Gough Island — Racomitrium crispulum, Thysanomitrium richardii, Jamesoniella spp. 1 The term cushion bog is used to indicate cushion plant communities on deep peat; see Wace (1965) for a discussion. The cushion habit is predominant in all of the above flowering plants. Bryophytes commonly as- sume either a cushion or compact habit. 3. An often conspicuous cryptogamic component of the vegetation. On the Falkland Islands the lichen Neuropogon is a conspicuous element of the feldmark vegetatiori. Mosses are conspic- uous in Macquarie Island feldmarks, and Ash- ton and Gill (1965) propose that the feldmark community be termed the Azorella- Ditrichum alliance. Ashton and Gill (1965) and Selkirk and Seppelt ( 1 984) regard the Macquarie Island feldmarks as a single variable vegetation unit. Selkirk and Seppelt recorded 63 plant species, 71% of them cryptogams (29 bryophytes, 16 lichens, 1 8 vascular plants). 4. A preference for higher altitudes. Wace (1960) lists feldmark altitudinal ranges for the follow- ing islands: Falkland Islands (600-700 m), Macquarie Island (200-450 m), Kerguelen Is- lands (100-300 m), and Tristan da Cunha and Gough islands (850-1 500 m). Wace ( 1 960) and Wace and Dickson (1965) indicate the distri- bution of feldmark is in large part determined by exposure. Due to exposure factors in the Falkland Islands, feldmarks occasionally occur at lower altitudes in the western part of the archipelago. Moore (1968, p. 16) stated that "Apparently the formation is not solely a re- sponse to high altitude, because a comparable community type occurs among rock detritus along exposed cliff tops on the west coast just north of Cape Meredith . . ." (southernmost point of West Falkland Island). On Westpoint Island it is presumably exposure which results in a feldmark at the summit of Mt. Misery (335 m). Feldmarks may also occur at rather low altitudes on Macquarie Island, where they oc- cur above 175 m (Selkirk & Seppelt, 1984). The general aspect of Falkland feldmarks differs considerably. While there may be a continuum present, there appear to be two general types of feldmarks. Mesic Feldmark— This type is characterized by a rather conspicuous fruticose lichen flora (prin- cipally Neuropogon). The lichen flora composition is apparently a result of the presence of mists and frequent enshrouding clouds. The feldmark com- munity on the summit of Mt. Usborne 1 (ca. 700 m) had a rather lush bryophyte vegetation due presumably to the numerous rather small scattered pools which are present. Certain bryophytes were associated solely with the pools, either submerged, at the edge of, or on rock in the pools. The feld- 34 FIELDIANA: BOTANY FIG. 18. Dry feldmark on ridge of north slope of Mt. Fegen, ca. 275 m. mark community on the summit of Mt. Maria (658 m) is rather xeric and has a conspicuous li- chen cover, but only a single bryophyte (Riccardia opuntiiformis). While Adelanthus integerrimus is the only he- patic found exclusively in this formation, it is of interest to note that the following hepatics are found exclusively in the Mt. Usborne feldmark and shel- tered high altitude cliff vegetation: Acrobolbus och- rophyllus, Anastrophyllum ciliatum, Archeochaete kuehnemannii, Herzogobryum vermiculare, Pachyglossa dissitifolia, and P. fissa. Xeric Feldmark— This type is characterized by a comparatively poor lichen cover (Neuropogon is absent). The moisture available to this feldmark type is derived solely from rain and not from en- shrouding clouds, and exposure is likely the factor which governs the comparative lack of moisture. Two feldmarks of this type were visited: the north- west tip of New Island opposite Landsend Bluff (90 m) and the north slope of Mt. Fegen (ca. 275 m)(fig. 18). Very characteristic of the bryophytes of this for- mation are the following three mosses, all of which grew on exposed rock surfaces: Andreaea pseu- domutabilis, Racomitrium lanuginosum and R. rupestre. On soil in crevices under rock overhangs are Lepidozia chordulifera and Ditrichum sp.; Bryum argenteum may be found on Bolax cush- ions. This feldmark type supports only a very small bryophyte vegetation. Even the rock crevices, which often are relatively moist in the Falkland Islands, were dry and only Lepidozia chordulifera, which is able to grow in a wide range of habitats, was apparently able to tolerate the dryness. FEN AND BOG FORMATION— This formation oc- curs where the water table is just below, at, or above the ground surface. Rostkovia Association— Moore (1968, p. 17) stated, "This [association] is normally confined to the wettest depressions, where drainage is imped- ed, and standing water is often present for long periods." In this association Sphagnum falcatum grew submerged or nearly so, with S. fimbriatum also present. No hepatics were observed. Astelia Association — Moore (1968, p. 17) stat- ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 35 19 FIG. 19. Sheltered high altitude cliffs (arrow) of Mt. Adam summit area. Photo by R. C. Harris. ed, "This [association] is usually found over deep peat and consists of a series of low flat cushions or a dense carpet composed of a few species which all are able to retain water in the dense covering of old leaves and branches surrounding their stems." These cushion bogs grade into the Rost- kovia association and the wetter facies of the Cor- taderia heath. The observed Astelia associations which did not grade into the above associations were dense, compact, and very hard, and no bryo- phytes were observed growing here. June us scheuzerioides Association— No visits were made to this association. BUSH FORMATION— This formation is of re- stricted occurrence in the Falkland Islands, and only two native shrubs contribute to associations of this formation. Chiliotrichum Association— This association was not visited in the Falkland Islands. According to Moore (1968) and Skottsberg (1913), it is best developed along the fringes of streams and rivers. Skottsberg (1913) listed two localities for this as- sociation, both in western West Falkland. Hebe Association— Hebe elliptica, which is re- stricted to the West Falkland Islands, was ob- served at Westpoint Island, but the plants were present only as scattered individuals or small thickets in a dwarf shrub heath (q.v.). When a dense stand of Hebe elliptica occurs, a true Hebe association is said to be present. Such an associ- ation is very rare in the Falkland Islands, and was not visited. A photograph of an example on Fox Island is in Skottsberg (1913, pi. 13). LITTORAL VEGETATION— The Senecio candicans andAmmophila-Elymus associations belong here, and both occur in sandy areas. The former asso- ciation was visited on the south shore of Cape Pembroke Peninsula, near Surf Bay (East Falk- lands), and the latter at Gypsy Cove, Port William region (East Falklands). No bryophytes were found in either of these associations. It is likely that the dune and sandy areas are too unstable for bryo- phyte survival. FRESHWATER VEGETATION— Moore (1968) in- cludes the Eleocharis and Myriophyllum associa- tions in this category. No bryophytes were ob- served in either of these associations. The bryophyte collections which were frequently gath- ered on the banks of shallow lagoons and ponds are included in the association of the surrounding area. SHELTERED HIGH-ALTITUDE CLIFF VEGETA- 36 FIELDIANA: BOTANY FIG. 20. Grove of planted trees (mainly Populus, Nothofagus, and conifers) at Hill Cove settlement. TION— These communities consist principally of bryophytes, and occur above 610 m in the Falk- lands (fig. 1 9). They possess a highly characteristic assemblage of hepatics, and because of their re- striction to high altitudes I believe they deserve separate recognition. Cliffs on ridgetops have nu- merous protective niches such as ledges and rock crevices, the latter frequently beneath overhangs of varying sizes. The crevices, especially when be- neath ledges, are commonly moist, cool, and well shaded, a situation highly favorable to bryophyte growth. When this is accompanied by water seep- age, the bryophytes are particularly luxuriant. The variation in exposures, moisture, light, and the consistently rather cool temperatures probably ac- counts for the considerable diversity of these com- munities. The following hepatics were encountered very frequently and often were intermixed: Austrolo- phozia fuegiensis, Balantiopsis bisbifida, Crypto- chila spp., Herzogobryum teres, and Pachyglossa spp. MAN-INFLUENCED COMMUNITIES — Pasture- land— In the Falkland Islands, there are wide ex- panses of pastureland (see Davies, 1939) that are regularly grazed by sheep. It is of interest to note that all three of the mosses occurring here are quite widespread and weedy, and it is not surprising that all are rather plastic species that are able to tolerate highly disturbed situations. Near the mouth of Cheeks' Creek, Fox Bay region, a collection was made in a stream cut through a pasture. It is prob- able that before conversion to pastureland the vegetation of the area was that of a dwarf shrub heath association, as the bryoflora was similar to the stream in that association, with the exception of the presence here of Leptophyllopsis irregularis. Groves of Planted Trees— A mature, well-de- veloped grove occurs at the Hill Cove settlement. According to Dallimore (1919), the trees were in- troduced in the period 1889-1894. The hardwood grove, which is made up of Nothofagus sp. and Populus sp., has a rather dense canopy with little woody undergrowth (figs. 20-21). The hepatics found here were Chiloscyphus lentus, C. leptan- thus, Leptoscyphus expansus, and Marchantia ber- teroana var. polylepida; all occur in a variety of natural communities. Adjacent to the hardwood grove was a stand of conifers. No bryophytes were observed in this stand or in the coniferous stand on Weddel Island. Gorse Thickets— Gorse (Ulex europaea), a na- ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 37 FIG. 2 1 . Interior of planted tree grove (mainly Nothofagus) at Hill Cove settlement. live of western Europe, is widely utilized in the Falkland Islands as sheep fencing. A gorse thicket was visited at Boca House near Brenton Lock, near Darwin Settlement, East Falklands. On soil be- neath the gorse occurred Leptoscyphus expansus, Brachythecium austrosalebrosum, Bryum argen- teum, B. (sect. Rosulata) sp., and Tortula robusta. Tortula monoica was the only bryophyte actually growing on the gorse and it occurred on the ex- posed roots. Phytogeography Austral Regions Several attempts have been made since 1960 to subdivide and classify austral zones. These clas- sification schemes are based upon climatic and biotic features, and also take into account the dominant influence of oceanic circulation, namely the position of the Antarctic and Subtropical Con- vergences. Wace (1960) devised a system based primarily on lowland vegetation criteria. He combined all those islands bearing oceanic heath and Sphag- num or flowering plant cushion bogs into a tem- perate category, and further divided this group into warm and cool temperate, respectively, ac- cording to the presence or absence of fern-bush communities. He limited usage of the term "sub- antarctic" to include only those islands with a closed herbfield vegetation and in which Sphag- num and cushion bogs are absent. The islands in which there are no flowering plants (or only scat- tered individuals), he considered as "antarctic."2 Wace (1960) compiled the available tempera- ture data and noted that the groupings based on lowland vegetation types correlated well with groupings based on mean annual temperature. The following is his classification scheme: SUBTROPICAL Kermadec Islands Juan Fernandez Islands TEMPERATE Chatham Island Tristan da Cunha, Inaccessi- ble, Nightingale, and Gough islands 2 It must be understood that islands, like Bounty Island off New Zealand, may be without vascular plants as a result of physiographic and biotic factors. 38 FIELDIANA: BOTANY New Amsterdam and St. Paul islands New Zealand shelf islands Falkland Islands Marion and Prince Edward is- lands SUBANTARCTIC Macquarie Island Crozet Islands Kerguelen and off-lying is- lands South Georgia Heard and Macdonald islands ANTARCTIC Bouvet Island South Sandwich, South Ork- ney, and South Shetland is- lands Palmer Archipelago (Graham Land) Greene and Greene (1963) and Greene (1964) adopted Wace's ( 1 960) concept of the term sub- antarctic, but included Marion and Prince Edward islands, and used the term antarctic region to em- brace the antarctic and subantarctic zones (fig. 22). This emended classification has been followed by van Zinderen Bakker (1967, 1971) and Gremmen (1982) for Marion and Prince Edward islands, by Lindsay (1971) for the South Shetland Islands, by Smith (1972) for the South Orkney Islands, and by others. Wace ( 1 965, p. 239) extended his earlier scheme by stating, "In order to arrive at a vegetational classification, the major features of the structure of the plant community should be used exclusively to delimit the different zones." He then proposed to use the limit of tree or woody shrub growth to separate the temperate from the subantarctic and the limit of closed phanerogamic communities to separate the subantarctic from the antarctic vege- tation. He characterized the subantarctic vegeta- tion zone as having terrestrial plant communities with the following four features: 1. Soligenous mires in which the important peat- forming plants are Bryales and Juncaceae (not Sphagnum). 2. Feldmark communities composed of flowering plants with very close-growing and compact mat and cushion forms. 3. Closed herbfield communities in which large- leaved perennial herbs are conspicuous. 4. Communities of large trunk-forming tussock grasses around the coasts. Wace excluded all continental areas and restricted the usage of the term subantarctic to remote is- lands. This zone included South Georgia, Marion, Prince Edward (which in 1960 he listed as tem- perate), Crozet, Kerguelen, Heard, and Macquarie islands. Holdgate's (1970) useful summary of the clas- sification systems of Wace and Greene separates subantarctic and south temperate as follows: SOUTHERN COLD TEMPERATE ZONE— Botanical Definition: From the Subtropical Convergence southward to the southern limit of dwarf shrub vegetation. Region: Western Magellanic Moorland of Chile, Falkland Islands, Tristan-Gough groups, I. St. Paul, Nouvelle Amsterdam, New Zealand shelf islands (Auckland, Campbell, Snares, etc.). SUBANTARCTIC ZONE — Botanical Definition: From the southern limit of dwarf shrub vegetation to the southern limit of extensive closed phanero- gamic vegetation. Region: South Georgia, Marion and Prince Edward, Crozet, Kerguelen, Heard, Macquarie (all are insular and isolated). Holdgate (1977) presented a more detailed clas- sification based upon those of the earlier investi- gators, but with the same circumscription of the subantarctic. Smith (1984) proposed an improved modification of the Holdgate (1977) classification, and made several alterations, among them the subdivision of the subantarctic into "three prov- inces according to their oceanic affinities" (Smith, 1984, p. 64), namely a South Atlantic province for South Georgia, a South Pacific province for Macquarie Island, and a South Indian Ocean prov- ince for the remaining islands. The subantarctic botanical zone sensu Holdgate supports a vascular plant flora of 72 species (cf. Greene & Walton, 1975; Smith, 1984), whereas the southern cold temperate zone has much greater floristic diver- sity. It is evident from the criteria outlined in Wace (I960, 1965) and Greene (1964) that the Falkland Islands are south temperate rather than subant- arctic. The characters which place them in the south temperate region are a widespread oceanic heath formation; the presence of flowering plant cushion bogs; Sphagnum bogs; the presence of woody shrub growth; and the absence of a closed herbfield vegetation. The presence of a feldmark community has been used as a characteristic of the subantarctic region. In the Falkland Islands, this community is present, but certainly does not reach the significant proportions and role that it does in the Kerguelen Islands. Skottsberg (1960) based his austral phytogeo- ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 39 Wen of Greenwich 0° Eut of Greenwich 110° I3S8 FIG. 22. South polar projection showing delimitation of subantarctic region (after Greene, 1964). graphic analysis on the following floristic zones (five further provinces are included by Skottsberg, some including "outposts" for "antarctic types"): 1 . Antarctic zone, south of 60°S, but including the South Sandwich Islands and Bouvet Island 2. Subantarctic zone, between approximately 48°S and 60°S, but including Marion, Prince Ed- ward, and Crozet islands a. Magellanic province i. West Patagonian-Fuegian district ii. Andean Patagonian-Fuegian district iii. Falkland Islands and South Georgia dis- trict b. Kerguelen province. Marion, Prince Ed- ward, Crozet, Kerguelen and Heard islands c. Province of the subantarctic islands of New Zealand 3. The austral zone, between approximately 40°S and 48°S. A conspicuous antarctic floristic ele- ment is present, often forming distinct com- munities in a flora and vegetation of a different origin. a. Valdivian province b. Province of the South Island of New Zea- land, Chatham Islands, and the Tasmanian tableland Godley ( 1 960) adopted the subantarctic zone of 40 FIELDIANA: BOTANY Skottsberg, but remarked (p. 472), "The presence or absence of Sphagnum bogs appears to me to be one of the significant features to be taken into consideration when delimiting a homogeneous subantarctic region." He regarded peat formation as a common feature of the region, and the bogs formed by cushion plants rather than by Sphagna. Smith (1984, p. 64) rejects the subantarctic zone as defined by Godley ( 1 960) and Skottsberg ( 1 960) since "it includes several cool temperate islands with arborescent plants and cushion bogs." Bliss (1979) presents an alternative view and develops a general classification system that em- braces comparable Northern and Southern Hemi- sphere vegetation. He combines alpine, arctic, and antarctic vegetation into one classification system and defines the southern lands as follows: The Subantarctic as used here includes closed and open forests and shrublands of Tierra del Fuego, Falkland Islands, and New Zealand shelf islands (Auckland and Campbell), Tris- tan da Cunha-Gough Islands, and the tall tus- sock grasslands of numerous islands. Antarc- tic as used here includes closed vegetation of grass-forb, herbfields, dwarf shrub, mires, and the open cushion plant fellfields found at higher elevations in some of the above islands [italics mine], but vegetation more common to the more southern islands (Macquarie, South Shetlands, South Orkney, South Geor- gia, Marion Island, Prince Edward Island, lies de Kerguelen, lies Crozet and Herd) (Bliss, 1979, p. 2168). and lichens. Two flowering plants, Deschampsia anlarctica and Colobanthus crassifolius, occur in Antarctica. The antarctic botanical zone was subdivided by Holdgate ( 1 964) into the maritime antarctic, which consists of the west coast of the Antarctic Penin- sula and the adjacent islands in the south Atlantic, where the marine influence allows growth of a wide range of bryophytes and lichens in addition to the two flowering plants; and the more extensive con- tinental antarctic, where bryophyte and lichen communities are only sparsely developed and flowering plants are unknown. Several workers have presented vegetation clas- sification systems for all or portions of the ant- arctic zone. Longton (1967) classifies the vegeta- tion of the maritime antarctic and uses an "Antarctic cryptogam formation" and later (Long- ton, 1973, 1979, 1982) provided classifications of vegetation within the antarctic botanical zone in which we find an "Antarctic nonvascular crypto- gam tundra formation." Gimingham and Smith (1970) and Smith and Gimingham (1976) present classifications of the cryptogamic communities of the maritime antarctic, and Smith (1972) does likewise for the South Orkney Islands; all of these authors use an "Antarctic nonvascular cryptogam tundra formation." Presence versus absence of permafrost is also an important factor in austral regions. Permafrost is lacking in the subantarctic zone, even at higher elevations, but is a feature throughout the antarctic zone. Thus, the terms subantarctic and antarctic as used by Bliss cannot be identified by zones. I concur with Smith (1984, p. 65) that the comparison by Bliss of "Arctic, alpine, Antarctic and sub-Ant- arctic vegetation categories is unsatisfactory be- cause the impoverished southern flora has a dif- ferent balance of life forms." The antarctic region includes regions south of the subantarctic and is characterized by a vege- tation which ". . . is at best a poorly developed tundra" (Rudolph, 1971, p. 192). Longton (1973, p. 2339) points out that "The vegetation through- out the Antarctic zone is less well developed than that at most sites in even the high Arctic, at least in terms of flowering plant representation. . . ." Soligenous mires, herbfields, tussock forming grasses, and cushion plants are absent, and the vegetation is a tundra type dominated by mosses The Phytogeographic Categories The chief difficulty in the assessment of the phy- togeographical relationships of the Falkland Is- lands Hepaticae and Anthocerotophyta is the de- limitation of the south temperate and subantarctic patterns of distribution. I have delimited these patterns after the concept of the zones or regions developed by Skottsberg ( 1 9 1 0, 1 9 1 6, 1 960), God- ley ( 1 960), Wace ( 1 960, 1 965), and Greene ( 1 964). I recognize 1 2 categories of distribution patterns of Falkland Islands Hepaticae (table 4). The An- thocerotophyta fall within only one of these cat- egories. The data for these distributions were gath- ered from specimens examined and reports extracted from the literature. If a report is regarded as questionable, it is not included here. I recognize 1 30 species of Hepaticae and one ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 41 species of Anthocerotophyta belonging to the Falkland Islands flora. SOUTH TEMPERATE— Species occurring within the south temperate regions of the world. American Temperate— Species occurring within the south temperate regions of the American sec- tor. There are eight species which occur in south- ern South America as well as South Georgia (see p. 43). Their distribution is regarded as temperate rather than subantarctic, as northward extension from the subantarctic is not confined to higher altitudes. It has frequently been remarked that the Falk- land vascular plant flora is an extension of the southern South American flora (Hooker, 1847; Skottsberg, 1909, 19 13; Moore, 1 968), a statement to be expected with regard to a continental shelf island. Moore stated that, of the 1 63 native species of vascular plants of the Falkland Islands, 89% occur on the mainland south of 40°S. All except three of the remainder are endemic to the Falkland Islands and are in large part related to mainland taxa. Like the vascular flora, the mainland hepatic representatives extend northward to varying de- grees. Commencing with the endemic category, the remainder of groups are arranged principally ac- cording to the degree of northward extension. Endemic to Falkland Islands (Group 1) Andrewsianthus planifolius Metzgeria engelii Riccardia regularis Schistochila subantarctica Fuegian- Falkland (Group 2)— Taxa occurring in the Falkland Islands and Tierra del Fuego. Aus- trolophoziafuegiensis is the only species belonging here. Several other species occur in the Falklands and in southern South America only within Tierra del Fuego, but such taxa also may be found on South Georgia (and sometimes other subantarctic islands) plus sometimes in the antarctic zone. Thus such species are either subantarctic in distribution (e.g., Acrolophozia fuegiana, Adelanthus integer- rimus. Riccardia granulata, R. saxicola), or ant- arctic in distribution (e.g., Cephalozia badia, Her- zogobryum teres). Magellanian- Falkland (Group 3)— Taxa occur- ring in the Falklands and southern Patagonia north to 48°S or Fuegia and southern Patagonia north to 48°S. The northern boundary was affixed by Skottsberg (1916) to delimit the Magellanian and Valdivian regions and has been widely followed by various authors. TABLE 4. Conspectus of phytogeographical cate- gories. Category Percent- age of No. of total species flora SOUTH TEMPERATE American Temperate Endemic to Falkland Islands 4 3.1 Fuegian-Falkland 1 0.7 Magellanian-Falkland 19 14.5 Valdivian-Falkland 3 2.3 Magellanian + Valdivian + Falkland 52 39.7 Andean-Falkland 2 1.5 Extra-American Temperate Amphipacific temperate 1 3 9.9 Amphiatlantic temperate 1 1 8.4 Pan-south temperate 6 4.6 NONTEMPERATE Subantarctic 12 9.2 Antarctic 6 4.6 Widespread 2 1.5 Total 131 100.0 Of the 1 9 taxa in this group, 74% have been found to occur in the Magellanian moorland. Their occurrence in the moorland is indicated by an as- terisk, and those taxa restricted to the moorland are indicated by a double asterisk. This region occurs south of 48°S to Cabo de Hornos, and is limited by the Magellanian rain forest to the east and the Pacific Ocean to the west (see discussion and map in Godley, 1960; see also Pisano, 1983, who provides a detailed treatment, preferring the term "Magellanic Tundra Complex"). The moor- land is extremely wet, exposed to violent winds, has a permanently saturated peaty soil, and is par- tially to distinctly open. Of the remaining four taxa, Chiloscyphus elatus is known from deciduous Nothofagus forests, Adelanthus tenuis is known from evergreen Nothofagus forests, and Anastro- phyllum ciliatum and Archeochaete kuehnemannii are known from both deciduous and evergreen Nothofagus forests. Adelanthus tenuis Anastrophyllum ciliatum * Andrewsianthus australis Archeochaete kuehnemannii * Balantiopsis bisbifida * Cephalolobus scabrellus Chiloscyphus elatus * Chiloscyphus hookeri 42 FIELDIANA: BOTANY Frullania microcaulis Gackstroemia patagonica Kurzia mollis Kurzia setiformis Leptoscyphus aequatus Pachyschistochila splachnophylla (^Plagiochila dura Plagiochila obovata Riccardia fuscobrunnea Saccogynidium vasculosum Telaranea oligophylla Valdivian- Falkland (Group 4,) —Species in the Falklands and Chile and/or Andean Patagonia (see group 5 for definition) north of 48°S and south of 36°S. See Engel (1978, pp. 21-23) for comments on this zone. All species also occur on Juan Fer- nandez. Jungermannia crassula Metzgeria multiformis Wettsteinia densiretis Valdivian + Magellanian + Falklands (Group 5)— Species that are essentially widespread in the American temperate zone. Of the 52 taxa in this group, 7 1% are known to occur in the Magellanian moorland. Their occurrence in the moorland is indicated by an asterisk. It should be repeated here that the moorland occurs south of 48°S and thus is restricted to the Magellanian floristic region. With regard to the Magellanian region, of the re- maining 29%, Cheilolejeunea savatieriana, Chi- loscyphus sylvaticus, Clasmatocolea rigens, Diplo- phyllum acutilobum, Leptophyllopsis irregularis, Megaceros fuegiensis, Paracromastigum subsim- plex. Riccardia papillosa, and Symphyogyna hochstetteri occur in the deciduous Nothofagus zone, Lejeunea corralensis, Plagiochila acantho- caulis, P. ansata, and P. gayana in the evergreen Nothofagus region, and Chiloscyphus sabuletorum, Riccardia opuntiiformis, and Telaranea pseudo- zoopsis in the deciduous and evergreen Nothofagus regions. The taxa, with the exception of groups 5e-f, are arranged according to their latitudinal range. 5a. South from 45°S: Cheilolejeunea savatieriana * Chiloscyphus divaricatus * Pigafettoa crenulata Riccardia opuntiiformis * Riccardia spectabilis 5b. South from 43°30'S (line from north side of Isla Guafo): * Chiloscyphus leptanthus * Chiloscyphus textilis * Kurzia saddlensis * Leptoscyphus patagonicus Plagiochila acanthocaulis Plagiochila ansata * Riccardia pallidevirens * Telaranea plumulosa 5c. South from 40°S: * Blepharidophyllum gottscheanum * Clasmatocolea obvoluta Diplophyllum acutilobum Megaceros fuegiensis * Plagiochila fagicola * Riccardia alcicornis * Riccardia tenax * Schistochila laminigera (40°07'S) Telaranea pseudozoopsis 5d. South from 36°S; from latitude indicated: * Balantiopsis cancellata (39°48'S) * Balantiopsis erinacea (36°50'S) * Chiloscyphus austrigenus (39°52'S) * Clasmatocolea fulvella (36°50'S) * Cryptochila paludosa (39°52'S) * Frullania boveana (39°53'S) * Frullania magellanica (39°52'S) * Harpalejeunea parasitica (39°48'S) * Isotachis humectata (36°43'S) * Nothostrepta bifida (39°52'S) * Plagiochila elata (39°36'S) * Radula helix (39°38'S) * Telaranea blepharostoma (39°48'S) * Tylimanthus urvilleanus (39°38'S) 5e. Valdivian + Magellanian + Falklands + South Georgia + sometimes also South Sandwich Islands. The species listed here are considered as south temperate rather than subantarctic, as northward extension from the subantarctic is not confined to higher altitudes. * Cephaloziella dusenii Clasmatocolea rigens * Gackstroemia magellanica * Lepicolea rigida * Lepidozia chordulifera * Lepidozia fuegiensis * Riccardia papillosa * Roivainenia jacquinotii 5f. Species about which insufficient knowledge is known concerning range: ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 43 Chiloscyphus sabuletorum Chiloscyphus sylvaticus Lejeunea corralensis Leptophyllopsis irregularis * Metahygrobiella tubulata Paracromastigutn subsimplex Plagiochila gayana Symphyogyna hochstetteri Skottsberg ( 1 9 1 0, 1 9 1 6) and Moore ( 1 968) rec- ognized three vegetation zones in the region south of40°S: 1 . West Patagonia: The region including the west slope of the Andes to the Pacific Ocean, char- acterized by high precipitation and supportive of lush rain forests and Magellanian moorland. 2. Andean Patagonia: The eastern slope of the Andes from base to snow line. This region ex- periences moderate rainfall and supports a de- ciduous, comparatively dry forest. 3. East Patagonia: The steppe region, ranging east from the Andean foothills, characterized by grassland and xerophytic shrubs. As is to be expected, the vast majority of He- paticae in groups 3-5 occur predominantly in the West Patagonian zone. The following Valdivian- Magellanian Hepaticae and Anthocerotophyta oc- cur within the Andean Patagonia zone, and nearly all occur in the Lago Nahuel Huapi region; none are restricted to Andean Patagonia: Chiloscyphus text His Clasmatocolea fulvella Clasmatocolea rigens Frullania magellanica Isotachis humectata Lepidozia chordulifera Megaceros fuegiensis Plagiochila elata Riccardia alcicornis Riccardia papillosa Riccardia tenax Roivainenia jacquinotii Schistochila laminigera Telaranea pseudozoopsis Tylimanthus urvilleanus Megaceros fuegiensis is the only species of Val- divian-Magellanian Hepaticae or Anthoceroto- phyta to occur in East Patagonia. Of the remaining Falkland Islands taxa, Leptoscyphus expansus, Marchantia berteroana, and Symphyogyna hy- menophyllum also occur there. Andean- Falkland (Group 6)— Species extending north of 36°S in the Andes. Of the American tem- perate species, Noteroclada confluens and Pseu- docephalozia quadriloba are the only taxa that have utilized the Andes as a route for northward mi- gration to lower latitudes. Extra-American Temperate— Amp hipacific Temperate— Distribution mainly in temperate parts of the South Pacific in the Southern Hemi- sphere, i.e., temperate South America (occasion- ally Juan Fernandez), New Zealand, Tasmania, and southeast Australia. There are varying degrees of penetration northward into the New Zealand sector, and the taxa have been arranged accord- ingly. One asterisk indicates occurrence on New Zealand shelf islands, two indicate occurrence in New Zealand, and three indicate occurrence on New Zealand shelf islands and New Zealand. Only two taxa also occur in the Northern Hemisphere: Metzgeria decipiens and Triandrophyllum subtri- fidum. a. South Island: ** Metzgeria violacea b. To North Island: ** Heteroscyphus triacanthus ** Pallavicinia xiphoides ** Saccogynidium australe ** Symphogyna hymenophyllum *** Telaranea tetradactyla *** Temnoma quadripartitum c. To Tasmania: *** Chiloscyphus bispinosus *** Lepidozia laevifolia * Triandrophyllum subtrifidum d. To Australia: *** Chiloscyphus lentus ** Clasmatocolea humilis e. To Japan: ** Metzgeria decipiens Amphiatlantic Temperate— Distribution main- ly in temperate parts of the south Atlantic, i.e., temperate South America, South Africa (plus oc- casionally Tristan da Cunha). Species whose sole occurrence in the Indian Ocean sector is on sub- antarctic islands are included here and indicated by an asterisk. They are not considered as sub- antarctic species, as northward extensions from the subantarctic are not restricted to higher alti- tudes. Gradstein et al. ( 1 983) discuss several species that belong to this element. Adelanthus lindenbergianus * Blepharidophyllum clandestinum * Blepharidophyllum densifolium 44 FIELDIANA: BOTANY Hyalolepidozia bicuspidata * Jensenia pisicolor Lepicolea ochroleuca Leptoscyphus expansus * Lethocolea radicosa Paracromastigum tristanianum Riccardia prehensilis Schistochila alata All of these taxa, with the exception of Hyalo- lepidozia bicuspidata and Paracromastigum tris- tanianum, penetrate north via the Andes to lower latitudes and are known from relatively high al- titudes north of 36°S. Pan- South Temperate— Species occurring in temperate regions of South America, New Zea- land-Australia, and South Africa: Acrobolbus ochrophyllus Chiloscyphus semiteres Clasmatocolea vermicularis Cryptochila grandijlora Jamesoniella colorata Marchantia berteroana NONTEMPERATE Subantarctic— Species occurring on one or more subantarctic islands (as denned by Greene, 1964) of at least one sector (e.g., American [A], Indian Ocean [I], or New Zealand [NZ], with northward extensions only at higher altitudes; see comments inEngel(1978, pp. 35-36): Acrolophozia fuegiana (A) Adelanthus integerrimus (A, I) Chiloscyphus humifusus (A, I) Herzogobryum erosum (A, NZ) Herzogobryum vermiculare (A, I) Pachyglossa fissa (A, I) Pachyglossa spegazziniana (A) Pachyschistochila leucophylla (A) Pedinophyllopsis abditus (A, I) Riccardia georgiensis (A, I) Riccardia granulata (A) Riccardia saxicola (A) Antarctic— Species occurring in the antarctic zone (as denned by Greene, 1 964), with northward extensions into the subantarctic or temperate zones only at higher altitudes. Species in this category are able to tolerate considerably colder tempera- tures than those in the subantarctic category. Oc- currence on subantarctic islands is indicated with the same symbols used for subantarctic species. (It should be mentioned that the worldwide dis- tribution of L. hatcheri is bipolar.) Cephalozia badia (A) Evansianthus georgiensis (A) Herzogobryum teres (A, NZ, I) Hygrolembidium isophyllum (A) Lophozia hatcheri (A) Pachyglossa dissitifolia (A) Widespread— Species not in any of the above categories and not pantropical. These species, which have a broad distribution with a range in- volving to a considerable extent the Northern Hemisphere, are Aneura pinguis and Metzgeria leptoneura. Distribution Within the Falklands The taxa below are arranged according to their occurrence in either the East Falklands or the West Falklands. TAXA KNOWN ONLY FROM EAST FALKLANDS— Of these, 48% are known only from the Mt. Us- borne region (indicated by an asterisk): * Acrolophozia fuegiana Andrewsianthus planifolius * Blepharidophyllum gottscheanum Cephalolobus scabrellus Chiloscyphus divaricatus Chiloscyphus text His Clasmatocolea obvoluta * Gackstroemia patagonica * Kurzia mollis * Kurzia saddlensis * Kurzia setiformis * Lejeunea corralensis Lepicolea ochroleuca Lepicolea rigida Leptoscyphus aequatus * Nothostrepta bifida Pachyglossa fissa * Paracromastigum tristanianum Plagiochila fagicola * Riccardia granulata Riccardia prehensilis * Riccardia saxicola * Saccogynidium australe Schistochila alata Telaranea blepharostoma * Telaranea oligophylla Telaranea tetradactyla ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 45 TAXA KNOWN ONLY FROM WEST FALKLANDS Chiloscyphus bispinosus Chiloscyphus humifusus Chiloscyphus semiteres Diplophyllum acutilobum Herzogobryum erosum Leptophyllopsis irregularis Lethocolea radicosa Megaceros fuegiensis Metzgeria violacea Pachyschistochila splachnophylla Paracromastigum subsimplex Pigafettoa crenulata Plagiochila dura Plagiochila obovata Radula helix Riccardia regularis Schistochila subantarctica While I regard the highly local occurrence of many Falkland Islands taxa to be a reflection of the availability of microhabitats and narrow eco- logical requirements, the comparison of hepatics restricted to the East Falklands (27) compared to the West Falklands (17) is impressive, and may well reflect differing climatic conditions within each of the respective island groups. Alternatively, the difference in species diversity between East and West Falklands may be due to critical local levels of temperature and rainfall which are present in various parts of the Falkland Islands (see fig. 6 with rainfall indicated; compare especially San Carlos with Port San Carlos). Port Stanley is one of the wettest areas in the entire archipelago, and this feature coupled with the cool temperatures known for the region may account for the eight taxa which were found to occur only in the Port Stanley-Port William re- gions. These species fall within a variety of phy- togeographic elements. Andrewsianthus planifolius Cephalolobus scabrellus Chiloscyphus divaricatus Lepicolea ochroleuca Riccardia prehensilis Schistochila alata Telaranea blepharostoma Telaranea tetradactyla The northwesternmost portion of the Falkland Islands has higher temperatures, and within this region are confined, or nearly so, three vascular plants (Blechnum chilense, Gavilea macroptera, and Ranunculus acaulis) of the Valdivian element. Chiloscyphus semiteres is known in the Falkland Islands only from Westpoint Island. The species seems to be restricted to rather warm temperature regions, especially in the American sector (i.e., the species is unknown for the Magellanian region), where it may be considered Valdivian. Of the taxa occurring on both East and West Falklands, two were confined to the wettest known stations. Frullania microcaulis and Metzgeria en- gelii were collected at Port Stanley and Port How- ard. Rainfall figures for the Mt. Usbome region would be of great interest. A large number of Hepaticae occur strictly in the Mt. Usborne region as well as at the wettest known stations on record, as the lists below illustrate. It is likely that the area receives as much rainfall as that of Port Stanley, Port How- ard, or Mt. Adam. Following are Hepaticae known only from the regions of Mt. Usborne, Port How- ard, and Mt. Adam: Anastrophyllum ciliatum Cryptochila grandiflora Cryptochila paludosa Herzogobryum teres Herzogobryum vermiculare Hygrolembidium isophyllum Pachyglossa dissitifolia In addition to the above rather sizeable suite of hepatics occurring only in the Mt. Usborne, Port Howard, and Mt. Adam regions, a rather large group of species is also known from combinations of these stations plus the Stanley region. Taxa oc- curring only in two or more of the Port Stanley (S), Mt. Usborne (U), Port Howard (H), and Mt. Adam (A) regions are: Acrobolbus ochrophyllus (U, A) Adelanthus integerrimus (U, H) Archeochaete kuehnemannii (U, A) Austrolophozia fuegiensis (S, U, H, A) Blepharidophyllum densifolium (S, U, A) Cephalozia badia (U, H) Clasmatocolea obvoluta (S, U) Evansianthus georgiensis (U, H) Frullania microcaulis (S, H) Jensenia pisicolor (S, U, H, A) Lepidozia fuegiensis (S, U, H, A) Leptoscyphus aequatus (S, U) Metzgeria engelii (S, H) Pachyglossa spegazziniana (S, U, H, A) Pachyschistochila leucophylla (S, U, H, A) Riccardia fuscobrunnea (U, H) Riccardia pallidevirens (U, A) 46 FIELDIANA: BOTANY Comparative Taxonomic Composition of the Hepatic- Anthocerote Flora It is revealing to compare the Falkland flora to that of southern South America, and especially to the Magellanian region, the area on the mainland most closely related phytogeographically to the Falklands. The Brunswick Peninsula is ideal for this comparison, since it is the southernmost land mass on mainland South America; its flora is strongly Magellanian in composition; and there is a complete floristic account of the Hepaticae of this region— the only one for all of southern South America (Engel, 1978). The following chart con- trasts the floras: Brunswick Falkland Peninsula Islands No. of families No. of genera No. of species No. of species per genus 29 743 193 1.8 26 64 131 2 In making such a comparison, the absence of a taxonomic unit is at times just as interesting or revealing as its presence. The families present in the Brunswick Peninsula but not in the Falklands are Trichocoleaceae, Porellaceae, and Calypogei- aceae. On the other hand, all families occurring in the Falklands also are present on the Brunswick Peninsula. It is also revealing to contrast the ge- neric composition of the Falklands. Genera pres- ent on the Brunswick Peninsula but absent in the Falklands are: Acromastigum Allisoniella Anthoceros Aphanolejeunea Apometzgeria Archilejeunea Austrolejeunea Bazzania Calypogeia Colura Hepatostolonophora 3 In order to increase the utility of the comparison of genera, I have made taxonomic adjustments among Brunswick Peninsula names so that they can be com- pared with Falkland Islands generic names. I have done so in three cases: Heteroscyphus occurs in the Brunswick Peninsula, but was included under Chiloscyphus in Engel (1978); likewise Pachyschistochila taxa were included under Schistochila and Pedinophyllopsis was included under Leptoscyphus. Thus the number of Brunswick Peninsula genera actually cited in Engel (1978) was 71. Herbertus Krunodiplophyllum Lepidolaena Paraschistochila Pleurocladopsis Porella Pseudolepicolea Reboulia Trichocolea Nearly all of the genera in the above list have ecological preferences in the Brunswick Peninsula that are in sharp contrast to any niche available on the Falklands (Engel, 1978). The vast majority of genera absent from the Falklands but present in the Brunswick Peninsula are restricted to the Nothofagus zone, and many are known only from evergreen Nothofagus forests.4 Several genera are absent because their constituent species occur only in niches offered by forest vegetation, e.g., on bark of living trees or on rotted logs. Examples of such taxa are Trichocolea, Bazzania, and Porella, all of which are common in wet forests of southern South America. Other genera commonly utilize filmy fern fronds as a niche, a feature characteristic of wet forests in southern South America, but lacking in the Falklands. Examples are Archilejeunea and Aphanolejeunea. Forests are completely lacking in the Falklands, except for the introduced com- munity at Hill Cove (figs. 20-21). Thus, because of the absence of forest communities in the Falk- lands and consequently the absence of large woody substrates, the lower humidity, and the lack of protection from the sun, a large number of mi- croniches available to bryophytes in a forested area such as on the Brunswick Peninsula are completely lacking in the Falklands. A second important factor relevant to distri- bution patterns of hepatics in the Brunswick Pen- insula is that of differences in rainfall. Nearly all genera of the Brunswick Peninsula that are lacking in the Falklands are restricted in the Brunswick to regions receiving in excess of 1 ,000 mm of rainfall annually. The highest annual rainfall recorded in the Falklands is 646 mm (fig. 6). The 1 ,000-mm rainfall figure seems to be critical in the Brunswick Peninsula, and there are dramatic differences in the numbers of taxa restricted to regions receiving more than 1 ,000 mm annual rainfall and those to 4 Engel (1973b, 1978) was the first bryologist to cor- relate southern South American vascular plant vegeta- tion types and zonation to distribution of hepatics. See these references for details. ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 47 regions receiving less than this amount (see Engel, 1978, table 3, pp. 40-41). The absence of a forest community and the com- paratively low annual rainfall undoubtedly are major factors in accounting for the absence from the Falklands of taxa from a range of different taxonomic levels. The following 1 0 genera occur on the Falklands but are unknown for the Brunswick Peninsula: Acrolophozia Aneura Austrolophozia Jensenia Jungermannia Pachyglossa Pallavicinia Paracromastigum Pigafettoa Wettsteinia Based upon known distribution patterns, some of the taxa (e.g., Acrolophozia, Austrolophozia, and Pigafettoa) are rather rare species throughout their range and perhaps truly are absent from the Bruns- wick Peninsula. Furthermore, two of the genera, Jungermannia and Wettsteinia, have a Valdivian- Falkland distribution and thus are unknown for the entire Magellanian zone. With further collect- ing, however, undoubtedly at least several of these genera will eventually be found in the Brunswick Peninsula. Origin of the Flora The phytogeographic affinities of the Falkland Islands hepatics and single anthocerote (see con- spectus on p. 42) indicate not only a strong re- lationship to South America but also noteworthy relationships to other austral land masses. Long- range dispersal may play a role in explaining some geographical discontinuities of bryophytes (van Zanten & Pocs, 1981; van Zanten & Gradstein, 1988), but the concept of continental drift is cru- cial to understanding floristic similarities, espe- cially at the species level, of austral bryophytes. The various Gondwanaland reconstructions de- picted in the literature vary somewhat in detail, but do not deviate to a major extent from the reconstructions shown in Figure 4. As discussed in Geological Features, the age of Gondwanaland and its subsequent breakup is also subject to dif- fering interpretation; such differences, however, are not of major importance to our discussion. Two points are paramount for our purposes: 1 . With the continuity of the land masses making up Gondwanaland, a migratory route was available that involved South America, South Africa, the fringes of Antarctica, Australia, Tas- mania, and New Zealand. The general impor- tance of this pathway is discussed in papers by, for example, Schuster (1969c, 1972b, 1976, 1979b, 198 la, 1982, 1 98 3) and Raven and Ax- elrod ( 1 974). Discussions relevant to origin and distribution of specific Gondwanalandic he- patic groups may be found in monographs, for example, of Pseudocephalozia (Schuster & En- gel, 1974), of Schistochilaceae (Schuster & En- gel, 1977, 1985), of Clasmatocolea (Engel, 1980) and of Lepidoziaceae subfam. Lembidioideae (Schuster & Engel, 1987). 2. The migratory route resulting from the geo- graphic continuity of ancient Gondwanaland was available for a very long period of time. As I have previously emphasized (Engel, 1980, p. 33), "This migratory pathway is of funda- mental importance in understanding the phy- togeography of the far south." The association and juxtaposition of the Falk- lands, together with the South American block, and other land masses involved with Gon- dwanaland are the foundation of our discussion. Available migratory pathways involving Gon- dwanaland are crucial to understanding the phy- togeographic relationships of the Falkland hepatic flora. Evidence indicates that the Falklands were associated with the South American block at least since the Lower Cretaceous (see fig. 4 and discus- sion on p. 6). The proximity of the Falklands to southern South America over a long period of time is at least partially the reason for the very strong phytogeographic affinity of the islands to southern South America. The conspectus on p. 42 reveals that the majority of the Falkland hepatic flora (61.8%) may be categorized as South American temperate. Falkland-South American temperate distribution patterns differ in the degree of range extension on the mainland, but all are similar in involving the Nothofagus zone of southern South America. The range may be narrow and rather restricted, as in Austrolophozia fuegiensis (fig. 74), to widespread in southern South America, as in Roivainenia jacquinotii (fig. 42), to the Andean- Falkland pattern illustrated by Noteroclada con- fluens and Pseudocephalozia quadriloba (fig. 29). Migratory pathways involving Gondwanaland are also relevant to discussion of the amphipacific (fig. 71, p. 145), amphiatlantic (fig. 23, p. 64) and 48 FIELDIANA: BOTANY pan-south-temperate (fig. 45, p. 101) distribution patterns, together making up 22.9% of the flora, as well as to subantarctic (fig. 46, p. 105) and ant- arctic (fig. 47, p. 105) patterns, together totaling 1 3.8% of the flora. In this connection, the sheltered high altitude vegetation in the Falklands is worthy of special mention (see p. 36). In this community, there is a strikingly high percentage of hepatics that either have a subantarctic or antarctic distri- bution, or also occur at higher elevations in New Zealand and Tasmania or have very close relatives there. The hepatic flora of this association appears to be in large part an old, relict one. A number of wide-ranging taxa (especially the genera Acrolo- phozia, Austrolophozia, Herzogobryum, and Pachyglossd) may well represent remnants of an early, widespread flora involving the Antarctic continent. Schuster (1969c) cited all of these gen- era and more in connection with discussions of antipodal distribution patterns, and stated (p. 56) that "it seems reasonable to regard these as relict groups at least in most cases," and that "presum- ably some of these genera already existed prior to the wide separation between the Tasmania-New Zealand area and Fuegia from Antarctica." Schus- ter (p. 56) cited Acrolophozia (but could have also cited Austrolophozia, Herzogobryum, or Pachy- glossd) as ". . . genera which are alpine today, often ex- isting in areas with summer snows, [and that such genera] may well have survived in what is today Antarctica even after its climate was already fairly inclement. Indeed, ecologically, the striking common denominator of most of the genera cited above (Phyllothallia, Ble- phahdophyllum, Acrolophozia, Austrolopho- zia, Herzogobryum, Pachyglossa) is that they are today essentially confined to treeless al- pine zones or extend to the lowlands only on the subantarctic islands. These taxa may then represent remnants of an early cold-antipodal flora that characterized perhaps the colder up- lands of an Antarctic continent whose low- lying areas harbored Nothofagus or Podocar- pus forests." Close juxtaposition with Africa was postulated over 35 years ago by Adie (1952b, cf. fig. 3). That concept was not widely followed until it was re- cently reexamined, with new evidence, by Martin et al. (1981), Mitchell et al. (1986), and Martin (1986) (see p. 6 and fig. 5). While hepatics as a group are old enough to reflect an Early Jurassic African-Falkland association, there are no exclu- sively African-Falkland taxa; all species that occur in the Falklands and in Africa also occur in South America. This is to be expected if one recalls that movement of a Falkland microplate occurred dur- ing the Jurassic, i.e., at a time that Africa and South America were juxtaposed and part of Gon- dwanaland (fig. 4). The continuity of Africa and South America was the most important event, and of secondary relevance is any possibility of intro- duction of African taxa to South America via a Falkland microplate. Systematic Account Notes on Format The sequence of orders and families follows the classification outlined in Schuster (1979a). The genera are arranged in alphabetical order within families, and species in alphabetical order within genera. Author citations follow the abbreviations in Sayre et al. (1964). Journal citations follow the abbreviations in Botanico-Periodicum-Huntian- um (B-P-H). I have attempted to indicate the status of typ- ification of all taxa; I have followed the concepts outlined in Engel (1978, cf. pp. 51-52), and have provided ecological notes for each species I have collected in the Falklands. The terminology and concepts used in these discussions have been de- fined in Phytosociology (see pp. 1 3-38). I have listed the Falkland Islands literature rec- ords for species that have been previously reported from these islands. The information is presented as it is given in the original source, and no attempt has been made to edit the extractions. For ex- ample, there are several instances where a collector is reported for a locality he did not visit. NOTE: References in which Falkland Islands taxa were originally described are given only in the taxo- nomic citations and are not repeated in the liter- ature record section. The localities in the synonymy sections are cited in the manner provided in gazetteers of the various regions, and it is only to this extent that I have edited the extractions. The following gazetteers were utilized: Chile (U.S. Office of Geography, 1967), Argentina (American Geographical Society of New York, 1944), and New Zealand (New Zea- land Dept. of Lands and Survey, 1968). Abbre- viations of localities follow the American Geo- ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 49 graphical Society indices to maps of Hispanic America (see American Geographical Society of New York . . . Argentina . . . 1944) and are as follows; add 5 to form plural: Arch. —Archipelago A.— Arroyo B.-Bahia Br.— Brazo Cta.— Caleta C.-Cerro Cord.— Cordillera Ens.— Ensenada Esto.— Estuario F.— Fiordo I.-Isla L.— Lago La. — Laguna M.— Monte Mt.— Mount, Moun- tain Pen. — Peninsula Pta.— Punta Port. — Portezuela Pto.— Puerto Q.— Quebrada Ran.— Rancho R.— Rio S.— Seno Sa.— Serra, Sierra Vo. — Ventisquero V.— Volcan I have grouped all Falkland collections together, followed by additional collections seen. The spec- imens I have personally collected are indicated by number only, without citation of collector. Distribution maps are provided for a number of taxa to illustrate the patterns of variation within and between phytogeographic categories as well as taxonomic ranks. Maps are only provided for those taxa for which there are sufficient reliable data. Introduction to Keys The keys include all taxa known to be present on the Falkland Islands as well as all genera that occur in the Magellanian zone of southern South America (cf. p. 42 for definition of this zone). The addition of non-Falkland genera will increase the utility of this treatment and allow the user to key, to generic level, all hepatics and anthocerotes presently known from the Magellanian sector; the keys thus include the vast majority of genera from the Nothofagus zone of South America. Genera (or higher taxa) not known for the Falklands are bracketed in the keys. Key to Classes and Orders of Hepaticae and Anthocerotophyta5 1 . Gametophytic chlorophyllose cells with numerous chloroplasts; chlorophyllose cells usually all (or many) with oil-bodies; gametophyte thalloid or leafy, without stomata, at times with air chambers; sporophyte a nonlinear capsule, usually 4-valved, without a columella, without stomata; sex organs exogenous Division Bryophyta, Class Hepaticae ... 2 2. Rhizoids, if present, all smooth, isolated thickenings excepted; plants leafy or thalloid, if thalloid ± delicate and translucent, without air chambers or pores; cells not sharply dimorphic, oil-bodies usually present in all chlorophyllose cells of gametophyte; capsule wall 2-10 stratose; seta usually long and extruding the sporophyte 3 3. Plants clearly and uniformly leafy; archegonia usually terminal on shoots resulting in a cessation of plant growth; capsule wall 2-10 stratose; rhizoids present or lacking 4 4. Rhizoids usually present; plants usually prostrate to procumbent, usually bilateral and anisophyllous or distichous-leaved; leaves, if unlobed, obliquely inserted. Sporophyte with wall 2-10-stratose, the cells variously thickened or lacking bandlike thickenings, never with annular longitudinal solitary bands. Perianths (or else a coelocaule, marsupium, or other protective device) usually present; if only with a shoot calyptra, the shoot calyptra on a short, postical branch Order Jungermanniales (p. 51) 4. Rhizoids never developed; plants erect, isophyllous, fundamentally radial (without distinct division into antical and postical faces); leaves unlobed, flat, transversely inserted. Sporophyte wall unistratose, with annular, longitudinal, solitary bands; perianth not known to occur, the green, massive, fleshy shoot calyptra replacing it Order Calobryales (p. 5 1) 3. Plants usually thallose (leafy only in Noteroclada and Treubia); archegonia and antheridia scattered on dorsal surface of thallus (occasionally on short thallus branches), or in dorsal groups, not causing cessation in growth of plant; capsule wall 2-5 stratose; rhizoids always present 5 * Key basically adapted from Schuster (1963a). 50 FIELDIANA: BOTANY 5. Plants thalloid, or if leafy, then not with two rows of dorsal leaflike appendages; gametangia variable in position, never in axils of foliar appendages. . . . Order Metzgeriales (p. 167) 5. Thallus leafy, the dorsal surface of thallus with two rows of leaflike appendages; gametangia in axils of dorsal "leaves." Cells strongly dimorphic, some or most with solitary, large oil- bodies that nearly fill the cell lumen, others bearing only chloroplasts [Order Treubiales, Treubia, cf. p. 1 67] 2. Rhizoids typically dimorphic, some tuberculate, others smooth; plants clearly thalloid, without leaflike lobes, the thallus firm, fleshy, opaque, with ventral scales, with air chambers or air canals opening dorsally by pores; cells typically dimorphic, a small minority of generally smaller cells each with a single, large oil-body, but no chloroplasts, the large majority of cells with chloroplasts only; capsule wall always unistratose; seta short, not extruding the sporophyte to any extent. Archegonia on specialized thallus branches (archegoniophores) Order Marchantiales Gametophytic cells (at least superficial cells) each normally with 1 (2-several) chloroplasts; cells lacking oil-bodies; gametophyte prostrate, thalloid, never with leaves, never with air chambers, with stomata on ventral surface; sporophyte linear, 2-valved, with a columella; archegonia and antheridia endogenous Division Anthocerotophyta, Order Anthocerotales Order CALOBRYALES An order with two families, Takakiaceae and Haplomitriaceae, each with one genus. Haplomitrium chilensis Schust. occurs in southern South America (see Schuster, 1971; Engel, 1984) but is absent from the Falklands. Order JUNGERMANNIALES Key to Genera of the Falkland Islands and Magellanian Zone of South America6 1. Leaf with dorsal lobe modified to form a water sac. Ventral merophytes absent (apical cell with 2 cutting faces); androecia and gynoecia on highly reduced, lateral-intercalary branches. Rhizoids scat- tered [Pleuroziaceae, Eopleurozia, cf. p. 1 58] 1 . Leaf with the ventral lobe modified to form a water sac, or with a sac absent altogether; ventral merophytes present, at least 1 cell row in width (apical cell with 3 cutting faces); androecia and gynoecia various in position, but not on highly reduced lateral-intercalary branches (except in some Heteroscyphus spp.) 2 2. Leaves of main stems with orientation (and usually insertion) incubous, at least at dorsal end of insertion (the leaves at times transversely inserted and oriented in Lepicolea), the leaves usually incubously shingled (but ventral portions sometimes modified to form a lobule or water sac); rhizoids always restricted in origin, never scattered on stem, often lacking or rare 3 2. Leaves of main stems varying from transverse in insertion to succubously inserted and oriented, to almost longitudinally (horizontally) oriented (never with ventral margin of leaf bearing, or transformed into, a lobule or sac); rhizoids often copious, often scattered on ventral face of stem 9 3. Leaves divided into a larger dorsal lobe and a solitary, smaller ventral lobe, which is usually saclike or pouchlike; branching exclusively lateral, the branches terminal or infra-axillary, neither vegetative nor sexual branches ever ventral or axillary; underleaves 0-2 lobed or absent 4 6 This key is adapted with considerable modification from Schuster (1963a), with portions from Schuster (1965a, 1966a, 1980a) and Engel (1978). ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 5 1 3. Leaves various, but never with ventral base or lobe modified to form a flap or inflated sac; branching various; underleaves always very large, varying from 2- to 4-lobed 7 4. Underleaves lacking; plants light to yellow green, rarely bronzed, never deep brown or reddish Group I (p. 53) 4. Underleaves present and conspicuous; plants at times brownish or reddish 5 5. Underleaves unlobed and edentate; plants relatively vigorous, often bronzed or brownish to blackish [Group II, p. 54] 5. Underleaves bilobed or at least emarginate 6 6. Lobuli, at least in large part, galeate, almost free from the dorsal lobe; plants often reddish or brownish or dark green Group III (p. 54) 6. Lobuli not saccate or galeate, united for most of their length with dorsal lobe along an elongate keel (the water sac formed partly by lobule, partly by the opposed portion of the lobe); plants green to yellow green. Plants often small and delicate; perianth usually pentagonal, with a beak Group IV (p. 54) 7. Plants with only lateral, terminal branching, without ventral branches and never with ventral flagella or stolons; leaves divided for 0.6 or more their length; leaf and underleaf margins and segments often ciliate or laciniate. Underleaves large and conspicuous Group V (p. 55) 7. Plants with branching not exclusively lateral and terminal; sometimes vegetative and/or sexual branches partly or wholly ventral in origin and abbreviated; plants often with ventral stolons or flagella (at least from older parts of plant); leaves rarely very deeply lobed, never with longly ciliate margins 8 8. Plants frequently isophyllous (both on main stems and branches), the underleaves similar in size and usually in form to lateral leaves (except some species of Isotachis); branches usually few and irregular, subfloral innovations excepted, all or in large part ventral and axillary, slender flagella usually lacking; sexual branches never abbreviated and ventral. Lateral leaves 2-4-fid; leaf insertion only weakly incubous, the orientation subtransverse Group VI (p. 55) 8. Plants normally somewhat to strongly anisophyllous, the underleaves differing in size (and usually in form) from lateral leaves, at least on the branches; underleaves often (if lobed) with fewer lobes, or lobes of different form than those of lateral leaves, or else more shallowly lobed; branches (usually) in large part lateral, usually terminal, the plants often regularly 1-2-pinnately branched; stolons, flagella, or rhizomes bearing vestigial leaves usually present; sexual branches nearly always short, ventral Group VII (p. 55) 9. Stem with underleaves conspicuous throughout (not merely with a minute underleaf here and there), even on sterile stems 10 9. Stem with underleaves either lacking on sterile stems, or minute, or merely small, scattered cilia or laciniae, or groups of slime papillae, never conspicuous (exc. Lophozia hatcheri). Rhizoids almost always scattered over ventral stem surface if present at all 15 10. Leaves sharply complicate bilobed, with a ± smaller (rarely subequal) dorsal lobe lying over a larger ventral lobe; plants with a coelocaule or a fleshy marsupium Group VIII (p. 56) 10. Leaves not complicate bilobed if lobed at all, the lobes lying in nearly the same plane, or leaf merely concave or ± naviculariform, never folded; plants with sporophyte protection various, but mostly with a perianth 11 1 1 . Mature leaves (2)4-6(9)-lobed, often very deeply so, the lobe tips usually ciliiform or acuminate, often the lobe margins with cilia or teeth; stem without a distinct, pellucid hyaloderm; underleaves very large, ca. 0.5-0.9 the area of leaves, similarly lobed and/or ciliate; rhizoids usually sharply restricted to underleaf bases Group IX (p. 57) 1 1 . Mature leaves undivided to 2(3-4)-lobed (occasionally with accessory small lobes or teeth, occasional leaves deeply trifid), if leaves lobed, then the lobe margins entire or dentate, but not with long cilia; stem sometimes with a hyaloderm; underleaves various, but for the most part markedly smaller than leaves and differing from them in shape and form 12 12. Plants nearly or quite isophyllous: the underleaves similar in size and form to lateral leaves, or nearly so; leaves erect to erect-appressed. Plants water-repellant. Leaves unistratose Group X (p. 57) 1 2. Plants quite anisophyllous, or if ± isophyllous (Hygrolembidium, Pachyglossd) then with leaves 52 FIELDIANA: BOTANY polystratose at least at base (except Chiloscyphus subg. Notholophocolea); leaves never erect- appressed 13 13. Rhizoids usually common, scattered over ventral stem surface; underleaves usually ovate to tri- angular, unlobed or obscurely bilobed, or large, bifid and/or ciliate; cells each with dense, conspicuous papillae (except Cephaloziella dusenii where small or absent and A mphicephalozia, where sometimes weak) Group XI (p. 57) 13. Rhizoids always restricted in origin, only at underleaf (sometimes also at leaf) bases, often largely confined to reduced leaves and underleaves of stolons, flagella, or rhizomes, never scattered; cells smooth or with scattered, small fine papillae (if, rarely, with coarser papillae, see Saccogynidium) 14 14. Stem often very soft, pellucid, with a cortex of large hyaline cells (hyaloderm) surrounding a ± small-celled medulla which is usually visible through the cortex by transmitted light (a hyaloderm lacking in Kurzia and Hygrolembidium)', cells not collenchymatous, often hyaline, large; perianth on ± abbreviated, intercalary usually ventral branches, long, triquetrous, often tapered distally; androecia usually on short ventral-intercalary branches Group XIV (p. 60) 14. Stem without a transparent, colorless hyaloderm: the cortical cells never conspicuously larger than medullary cells, the medullary cells not visible through the cortex; cells various, often with distinct trigones; perianth, if developed, usually on a ± elongated stem or branch, or on short lateral (rarely on short ventral) branches, seldom tapered distally; androecia on lateral branches, or if ventral (Saccogynidium) then plants with marsupia Group XII (p. 58) 15. Leaves ± sharply complicate-bilobed (except Blepharidophyllum densifolium, which has a broadly rounded keel), transversely to succubously inserted and oriented; perianth dorsiventrally com- pressed. Leaf margins (and/or apices) often ± denticulate to ciliate Group XIII (p. 59) 15. Leaves lobed or not, but if lobed, then never with lobes sharply bent over each other; perianth (if present) not sharply dorsiventrally compressed 16 16. Stem soft textured, consisting of a conspicuous, hyaline cortex of enlarged cells (surrounding a medulla of much smaller, ± thick-walled cells usually distinct by transmitted light, or medullary cells also soft but large); leaves bilobed, with cells thin-walled and ± hyaline Group XIV (p. 60) 16. Stem anatomy very various, but never with a hyaline cortex of large cells surrounding a medulla of smaller, ± thick-walled cells; leaves various, not bilobed and with leptodermous, hyaline cells 17 17. Leaves inserted ± transversely (at least in part of the dorsal half of the insertion) and transversely oriented; leaves generally appearing ± pectinate when not densely appressed-imbricate. Leaves bilobed (except some species of Herzogobryum). Plants with perianths, never with a marsupium Group XV (p. 61) 17. Leaves obliquely inserted and ± succubously oriented, at least half of the insertion quite oblique on stem to nearly horizontal. Leaves unlobed to short bifid, strongly bilobed only in Lophozia. Plants with sporophyte protection various, at times with a marsupium 18 1 8. Plants with perianths terminal on leafy, main, or lateral stems; androecia intercalary or terminal on leading stems; terminal branching common, absent in a few taxa . . . Group XVI (p. 61) 18. Plants without perianths, with pendent perigynia or with an erect fleshy, rigid shoot calyptra; terminal branching absent. Leaves various, often entire or irregularly dentate; capsule wall 5- 9-stratose . Group XVII (p. 62) GROUP I: Radulaceae, Lejeuneaceae (p.p., cf. also groups II, IV) 1 . Leaves with a wide J- or U-shaped insertion; rhizoids (where present) in fascicles from lobuli; ventral merophytes 3-5 or more cells broad; perianth dorsiventrally compressed, the mouth wide, truncate Radula (p. 1 58) 1. Leaves with a narrow transverse insertion; rhizoids in fascicles from stem; ventral merophytes 1 cell broad; perianth (4)5-keeled, the mouth beaked 2 ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 53 2. Branches of Lejeunea type, with a collar at the base; leaves never sharply dimorphic; inflated lobules with hyaline papilla usually ± proximal to apical tooth, if ± entally displaced (often), then rarely inserted on midline of tooth [Cololejeunea] 2. Branches of Aphanolejeunea type, without a collar; leaves sharply dimorphic: some with inflated lobules, others with lobule vestigial and 1-2-celled; inflated lobes with hyaline papilla entally displaced, at base of the 1-2-celled tooth along its ental midline, or displaced to distal side .... .... [Aphanolejeunea} " GROUP II: Porellaceae, Lejeuneaceae (p.p., cf. also groups I, IV) 1. Lobule similar in shape to the underleaves, lying parallel to them, free from lobe, never inflated, with margins often ventrally reflexed; elaters free, tapered at both ends; plants green (in ours) .... [Porellaceae, Porella, cf. p. 161] 1 . Lobule differing in form and size from underleaves, lying normally parallel with posterior lobe margin and broadly united with it, usually inflated; elaters attached to valves, the free end trumpetlike; plants greenish brown to brownish to blackish 2 2. Stem with cortical cells thick-walled, not forming a hyalodermis; perianth 3-5(6)-keeled; oil- bodies segmented [Archilejeunea] 2. Stem with cortical cells relatively thin-walled, forming a hyalodermis; perianth usually pluriplicate; oil-bodies homogenous [Brachiolejeunea] GROUP HI: Lepidolaenaceae, Jubulaceae 1. Underleaves not becoming galeate; plants with perianths, but without a coelocaule; elaters 1 -spiral; leaf lobes entire Frullania (p. 162) 1 . Underleaves, at least of the ultimate branches, becoming galeate; plants with a large, variously paraphyllose coelocaule and without a perianth, or with a coelocaule subtending a perianth; elaters 2-3-spiral; leaf lobes sometimes armed ; 2 2. Underleaves of main axis bifid, emarginate or (in Falkland species) undivided; main shoot com- monly black brown; stems without paraphyllia Gackstroemia (p. 1 60) 2. Underleaves of main axis quadrifid; main shoot green to ± red; stems with paraphyllia [Lepidogyna] GROUP IV: Lejeuneaceae (p.p., cf. also groups I, II) 1 . Plants with segmentation pendulum-like, i.e., with 1 underleaf per lateral leaf; leaves highly specialized with conversion of the lobule and lobe (or portion of it) to a complex water sac, the lobule closed by a specialized movable valve. Plants with leaves erect, oriented away from the substratum [Colura] 1. Plants with segmentation helical, i.e., with 1 underleaf per pair of lateral leaves; leaves without a complex water sac and valve mechanism 2 2. Leaves sharply constricted at base, with a strongly abbreviated, ± transverse attachment to the stem. Lobes narrow, obovate, elongated; lobule with a very narrow insertion formed by 1-2 cells, obovate, tridentate distally along free margin; underleaves narrow, with filiform lobes which are uniseriate for most of their length [Austrolejeunea] 2. Leaves with a wide, distinctly J- or U-shaped insertion, the dorsal lobe distinctly incubously inserted 3 3. Hyaline papilla of lobule distal to apical tooth; leaves convex, often strongly so; plants firm, often dull and opaque Cheilolejeunea (p. 164) 3. Hyaline papilla of lobule proximal to apical tooth; leaves flat to somewhat convex, rarely strongly so; plants delicate, generally somewhat pellucid 4 54 FIELDIANA: BOTANY 4. Lobes blunt, acute to acuminate, narrowly ovate to lanceolate, 1.5-3 x as long as wide; underleaves obdeltoid, with ± divergent lobes only 1-5 cells wide; plants small, remote- leaved 5 5. Underleaf lobes setaceous to sharply acute, 1-2 cells broad, strongly divaricate [Drepanolejeunea] 5. Underleaf lobes blunt to rounded, 4-5 cells broad usually, with blunt to angulate sinus Harpalejeunea (p. 1 66) 4. Lobes blunt to rounded, orbicular-ovate or ovate-falcate usually, less than 1.5 x as long as wide. Leaf trigones absent or small Lejeunea (p. 1 66) GROUP V: Lepicoleaceae, Ptilidiaceae 1. Leaves usually ± symmetrically (3)4-parted, the lobes entire or dentate to ciliate near base only; underleaves lobed as in leaves; plants without perianths but with a fleshy coelocaule densely covered with scales and paraphyllia; plants with subfloral innovations Lepicolea (p. 63) 1. Leaves very asymmetrically lobed and 2-4(5)-parted, with ciliate to dentate lobes; underleaves bi- lobed; plants with a perianth; plants without subfloral innovations [Ptilidiaceae, Ptilidium, cf. p. 1 58] GROUP VI: Vetaformaceae, Herbertaceae, Balantiopsaceae (p.p., cf. also group VIII) 1. Leaves distinctly vitiate, a "vein" of longer cells running into each lobe; leaves deeply (0.5-0.75) bifid; cells with coarse trigones. Leaf lobes slender, often falcate. Perianths present; bracteolar an- theridia present [Herbertus] 1. Leaves non vitiate, the lobe cells not conspicuously elongated, leaves 0.2-0.5 bifid-trifid; cells wilh irigones absenl lo small. Perianlhs absenl or presenl; bracleolar antheridia absent or present . . 2 2. Leaf and underleaf margins enlire or dentale in basal portion, sparsely armed above; planls usually wilhout conspicuous pigmentalion, al mosl pale brown; male bracleoles wilh anlheridia. Perianlhs presenl or absenl 3 3. Leaves and underleaves 2-3-lobed for 0.4-0.6 Iheir lenglh, ihe lobes broad al base, iriangular; perianlh dislincl Triandrophyllum (p. 65) 3. Leaves and underleaves (2)3-4-lobed for 0.6-0.75 Iheir lenglh, ihe lobes linear-lanceolale, only (3)4-5 cells broad al base; perianlh absenl, a clavale coelocaule presenl [Velaformaceae, Vetaforma, cf. p. 63] 2. Leaf and underleaf margins wilh leelh, when presenl, often in apical portion, nol predominanlly in basal portion; planls usually wilh reddish and/or cheslnul brown lo fuscus pigmenlalion; male bracleoles wilhoul anlheridia. Perigynium presenl, firm, fleshy, with a vestigial perianth al ils lip Isotachis (p. 1 57) GROUP VII: Lepidoziaceae (p.p., cf. also group XIV), Calypogeiaceae 1. Leaves deeply (2)4-6-lobed for usually 0.5 or more Ihe leaf lenglh; planls regularly ± pinnalely branched 2 2. Leaves wilh lamina (in ours) 0.5-4 cells high; leaf lobes uniseriale Ihroughoul, filiform and lerele, al mosl 2 cells wide in basal cell lier; slem wilh a conspicuous hyaloderm layer, formed of large cells wilh walls lillle or not thickened conlrasled lo Ihe smaller medullary cells often wilh ihick walls; leaves usually symmelrical; leaf cells usually large, hyaline, Ihin- or slighlly ihick-walled, Irigones lacking Telaranea (p. 78) 2. Leaves wilh a lamina many cells high; leaf lobes iriangular, never filiform; slem wilhoul a hy- aloderm layer (cortical cells ± thick walled, not conspicuously larger than medullary cells); leaves ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 55 usually conspicuously asymmetric (dorsal margin longer and/or arched); leaf cells usually rigid, often with thick walls and guttulate lumina Lepidozia (p. 72) Leaves with apices edentate or 2-3-dentate-lobate, never deeply lobed, the sinuses rarely descending more than 0.25 the leaf length (and always less than 0.5 the leaf length); branching various, but not regularly pinnate 3 3. Leaves firm, the cells thick-walled or with distinct trigones; leaf apices usually (0)2-3 -dentate; lateral terminal branches frequent to common; ventral intercalary stolons freely developed; stem firm-textured; plants with perianths _ 4 4. Stolons intercalary in origin, from axils of underleaves; stem without a hyaloderm, the cortical cells in many rows; leaf apices typically 2-3 -dentate; pseudodichotomous branching present: Frullania-lype branches only from one side of axis and often subequal to main shoot in vigor [Bazzania] 4. Stolons terminal in origin, replacing one-half of an underleaf; stem with a hyaloderm, at least the branches with only 7 rows of cortical cells; leaf apices 0-2 dentate-lobate; pseudodichot- omous branching lacking, Frullania-type branches arising from both left- and right-hand sides of axis [Acromastigum] 3. Leaves delicate, subhyaline, leptodermous (or with small trigones); leaves entire or finely bidentate at apex; lateral terminal branches few or absent; ventral intercalary stolons lacking; stem soft- textured; plants with marsupia, without perianths. Branching normally all ventral intercalary . . [Calypogeiaceae, Calypogeia, cf. p. 82] GROUP VIII: Balantiopsaceae (p.p., cf. also group VI), Schistochilaceae 1. Apex of dorsal lobe usually suberect, lying at an angle to axis of ventral lobe, the lobes ± divergent; keel between lobes wingless; cells of leaf lobe oriented in ± distinct tiers; sporophyte (unknown for Anisotachis) in a marsupium 2 2. Leaf lobes bifid, rarely subequal, the apices without a border Balantiopsis (p. 154) 2. Leaf lobes undivided, truncate or emarginate, subequal, with apices bordered by 1-several trans- verse rows of cells [Anisotachis] 1 . Apex of dorsal lobe oriented towards apex of ventral lobe; keel between lobes usually with a broad wing; cells of leaf lobes irregularly arranged, not in tiers; sporophyte in a terminal coelocaule lying in the axis of the stem. Leaf lobes undivided, never bifid 3 3. Rhizoids colorless, usually fasciculate, the apices often digitiform and copiously septate, the branches becoming polyseriate; plants never with wall pigments of any part of gametophyte; cells of leaves and underleaves frequently showing secondary septation (asexual reproduction via re- generation); leaves partially to largely polystratose (at least basally and in keelar region), without specialized dentition, the ventral lobes never lamellate or with cellular processes; branches nor- mally of lateral-intercalary or Radula type (in P. reflexistipula with both Frullania and Microle- pidozia-lype branches) Pachyschistochila (p. 1 53) 3. Rhizoids distally at least, magenta to claret red, always scattered, the apices, if ramified, irregularly branched, the branches often juxtaposed and/or anastomosing, usually sparingly if at all septate; plants with pigmentation of remainder of gametophyte present in some taxa; cells of foliar ap- pendages never secondarily septate; leaves (aside from the keelar area in Subg. Eoschistochild) uniformly 1 -stratose, often with specialized dentition (setae, cilia, etc.), often with lamellae and/ or processes of ventral lobes; many taxa retaining Frullania-lype branches (but none with Micro- lepidozia-iype branches) 4 4. Radial symmetry in vegetative, gynoecial, and androecial regions; elaters slender and tortuous, 1 -spiral; leaves unwinged, noncomplanate [Pleurocladopsis] 4. Radial symmetry lacking (in Subg. Austroschistochila subsymmetric in gynoecia alone); elaters usually stout and rigid, always 2-spiral; leaves winged, sharply complanate 5 5. Underleaves and $ and 9 bracteoles distinct Schistochila (p. 1 52) 5. Underleaves and <5 and 9 bracteoles completely lacking [Paraschistochila, p. 152] 56 FIELDIANA: BOTANY GROUP IX: Trichocoleaceae 1. Leaves appearing as a mass of interwoven cilia such that the lamina is obscured; stems paraphyllose; plants with a fleshy, hispid coelocaule, a perianth low or lacking; branching copious, exclusively of Frullania type; plants green, without brown pigmentation [Trichocolea] 1. Leaves without interwoven cilia and/or segments (or if so, e.g., in Temnoma pilosum, then plants with brown pigmentation), with at least part of lamina conspicuous; stems not paraphyllose; plants with a conspicuous perianth, at most with a coelocaule precursor (in Temnoma spp.); branching sparing or sporadic, not exclusively of Frullania type, ventral-intercalary branches at least occasionally present; plants often with brownish pigmentation 2 2. Leaves and underleaves bifid or bisbifid. Leaf lobes always entire and several cells broad (even of subfloral leaves and bracts); plants perfectly isophyllpus v 3 3. Leaves and underleaves bifid. Leaves unistratose [Isophyllaria] 3. Leaves and underleaves (except on juvenile shoot sectors) normally bisbifid 4 4. Leaves polystratose, the lobes rigid, polystratose, ± terete; leaves with knotlike swellings at the base of the median and often also the lateral sinuses; branching exclusively intercalary; perianth mouth short lobate-dentate [Herzogiaria] 4. Leaves unistratose throughout, the lobes thin, flat; leaves devoid of knotlike swellings; branching all or predominantly terminal, of Frullania type; perianth mouth crenulate . . . [Pseudolepicolea] 2. Leaves of mature shoots equally quadrifid. Leaf lobes armed or entire; plants anisophyllous to subisophyllous 5 5. Cells of leaf disc with elongated, thin- to equally thick-walled cells 6 6. Lobes of mature leaves (usually of vegetative shoot sectors, at least in and below gynoecia) with opposed, sharp teeth or cilia usually on both disc margins and lobes, rarely on one only; perichaetial bracts freely spinose-dentate to copiously ciliate; perianth wide at open mouth Temnoma (p. 67) 6. Lobes of mature leaves without cilia or teeth (or, rarely, with an isolated tooth on one or both margins of the disc); perichaetial bracts without trace of teeth or cilia; perianth closely contracted to the narrow mouth. Leaves with segment apices setaceous Archeochaete (p. 67) 5. Cells of leaf disc with well-developed trigones. Leaves and underleaves edentate; leaves 3—4- lobed for 0.6-0.75 their length, the lobes acuminate, uniseriate for much of their length; subfloral bracts usually larger than innermost bracts and with disc margins ciliate or dentate [Archeophylla] GROUP X: Antheliaceae 1 . Plants greenish but with age often distinctly whitish; branching normally terminal lateral, of Frullania type, rarely ventral intercalary; stem cortical cells forming a weak hyaloderm, the cells somewhat swollen and bulging, at least their free walls thin. Oil-bodies lacking [Anthelia. cf. p. 91] 1 . Plants red brown; branching strictly intercalary, both ventral and lateral; stem without a differentiated cortex, the cortical (and medullary) cells very thick walled. Oil-bodies unknown (Grollea, cf. p. 91] GROUP XI: Lophoziaceae (p.p. min., cf. also groups XV, XVI), Acrobolbaceae (p.p. min., cf. also group XVII), and Cephaloziellaceae (cf. also group XV) 1 . Underleaves large, ciliate and/or bifid; without stolons or flagella; plants vigorous (shoots to 3-5 mm wide), usually ± brownish. Cells with small or moderately coarse trigones; cuticle with high, coarse, hemispherical papillae; perianth apex twisted; bracts of innermost series very deeply lacerated, smaller than other bracts or leaves; capsule wall 7-stratose Roivainenia (p. 97) ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 57 Underleaves small or minute, eciliate, usually unlobed; often with flagella or stolons; plants small (shoots to 0.7-2 mm wide) 2 2. Leaves succubously inserted throughout, large (leaf width much greater than stem width); perianth lacking, plants with marsupia Acrobolbus (p. 148) 2. Leaves with at least dorsal half transversely inserted (occasionally dorsal half subsuccubous in C. scabrellus), small, distant; perianth present. Plants with Cephaloziella-\ike facies 3 3. Stolons and flagella present; leaf cell trigones distinct; plants often with brownish to reddish secondary pigments; perianth mouth lobulate-laciniate Cephalolobus (p. 95) 3. Stolons and flagella lacking; leaf cell trigones minute; plants green, without secondary pigments; perianth mouth obscurely shallowly lobulate v [Amphicephalozia] GROUP XII: Geocalycaceae 1 . Plants producing marsupia, perianths absent; sexual branches always on abbreviated, small, ventral- intercalary branches. Leaves ovate, undivided or bidentate (occasionally short bifid in S. vasculoswri), the cuticle very densely papillose; underleaves 0.5-0.75 bifid (in our species), free from and much smaller than leaves; plants dull, opaque, gray or light green plants developing (often) some brownish pigmentation Saccogynidium (p. 142) 1 . Plants producing perianths; sexual branches on apices of unmodified stems or short lateral branches, rarely on short ventral branches 2 2. Perianths ± strongly laterally compressed, basically 2-lipped (the ventral lobe strongly reduced or vestigial); rhizoids often tending to spread away from underleaf bases or (Pedinophyllopsis) ± scattered; plants often ± brownish, often intensely so 3 3. Dorsal-intercalary (Andrewsianthus-type) branching present; leaves (marginal sectors aside) 2- 3-stratose; flagelliform branches present. Underleaves very small, narrower than stem, bifid Evansianthus (p. 1 29) 3. Dorsal-intercalary branching lacking; leaves unistratose throughout; flagelliform branches ab- sent. Underleaves variable, at times small and inconspicuous, at times equal to the leaves, undivided or bifid 4 4. Plants lacking secondary pigments, strikingly laterally compressed and complanate, the leaves strongly erect, often appressed to one another dorsally; underleaves minute, free from leaves, with (1)2 juxtaposed long ciliiform segments; rhizoids in large part scattered; perianths barely exserted beyond bracts; male bracts with paraphyllia. Leaves unlobed and entire Pedinophyllopsis (p. 141) 4. Plants often with red brown pigments, basically somewhat to strongly dorsiventrally flat- tened (if leaves concave and dorsally assurgent, then dorsiventral complanation ill-defined, but plants not laterally compressed), the leaves never laterally appressed; underleaves usually large, typically ± decurrent and often connate with 1 or both leaves, the segments usually at most laciniiform, never long ciliiform; rhizoids confined to underleaf bases (from there sometimes spreading somewhat away from underleaf base); perianths usually distinctly exserted beyond bracts; male bracts lacking paraphyllia. Leaves 2-3- or more dentate to lobed, or unlobed and entire Leptoscyphus (p. 1 33) 2. Perianth trigonous to trigonous-inflated, the mouth equally or subequally trilobate; rhizoids re- stricted to underleaf bases; plants normally lacking brown pigments 5 5. Androecia and gynoecia all or predominantly on unspecialized leafy shoots; androecial position generalized: terminal or intercalary on main axes or branches indeterminate in length, the branches normally bearing at least some sterile leaves (either at branch base or distally). Leaf cells generally leptodermous, without trigones or with trigones small, medium, or absent; underleaves (in ours) not strongly connate on both sides 6 6. Plants stoloniferous; leaves transverse to subsuccubously oriented 7 7. Plants nearly isophyllous, the underleaves conspicuous and rigidly patent; leaves and underleaves polystratose at least at the base; female bracteoles ± similar in size to bracts Pachyglossa (p. 1 37) 58 FIELDIANA: BOTANY 7. Plants clearly anisophyllous, the underleaves highly reduced, inconspicuous, appressed to moderately spreading; leaves unistratose throughout; female bracteoles reduced, in- conspicuous, hardly modified from underleaves [Hepatostolonophora] 6. Plants usually lacking stolons; leaves mostly strongly succubously oriented 8 8. Leaves moderately to deeply adaxially concave, with apices incurved 9 9. Leaves strongly succubous, the insertion a long, strongly oblique line 10 10. Plants clearly anisophyllous; underleaves plane, convex, or sometimes cucullate (ventral view); leaves usually conchiform concave, occasionally moderately adaxially concave. Leaf apices mostly undivided, in a few species 2- or 2-3- lobed Clasmatocolea (p. 1 23) 10. Plants subisophyllous to isophyllous; underleaves occasionally strongly concave (recurved) (ventral view), particularly toward shoot apices; leaves sporadically weakly to rather strongly convex. Leaf apices undivided, entire or 1 -dentate . . . [Concave-leaved phases of Chiloscyphus subg. Notholophocolea] 9. Leaves transverse, subsuccubous or subincubous, the insertion narrow 11 11. Leaf margins (and abaxial leaf faces) crenulate to mamillate with domelike cell wall protuberances; leaves Cephaloziella-\ike, at most slightly concave, without an incurved ventral margin, normally lacking any trace of teeth; leaf insertion transverse to weakly succubous Pigafettoa (p. 141) 1 1 . Leaf margins (and abaxial leaf faces) not crenulate to mamillate with domelike cell wall protuberances; leaves ± Nowellia-like, strongly concave, the ventral margin incurved and accentuating and defining a deep pocket-like concavity in ventral portion of leaf, the leaves with accessory teeth; leaf insertion transverse to weakly incubous. Valdivian [Xenocephalozia] 8. Leaves convex, rarely plane, with apices decurved or deflexed 12 1 2. Leaves with lobes and marginal teeth caducous, often giving leaf apices a ragged appearance; leaf apices often with accessory teeth and laciniae Leptophyllopsis (p. 1 32) 1 2. Leaves entire or if lobed, then with lobes persistent; leaves with marginal teeth, if present, persistent; leaf apices never with a ragged appearance and not with accessory laciniae Chiloscyphus (p. 1 1 1) 5. Androecia and gynoecia on specialized, abbreviated, intercalary branches; androecia (abnormal exceptions aside) on determinate, narrow, compactly spicate, lateral-intercalary branches that never bear normal leaves, and never innovate distally. Leaf cells often with coarse to strongly nodose or triradiate trigones; underleaves often strongly connate on both sides Heteroscyphus (p. 1 32) GROUP XIII: Blepharidophyllaceae, Scapaniaceae 1 . Leaf lobes each short bifid; branching predominantly terminal, of Frullania type; rhizoids in small fascicles (often present only on small plants) from the stem at the ventral bases of leaves Blepharidophyllum (p. 1 08) 1. Leaf lobes undivided, never bifid; branching predominantly lateral-intercalary; rhizoids scattered over ventral surface of stem, never in fascicles 2 2. Leaves unequally complicate bilobed, the dorsal lobe smaller than the ventral (except in ours, where subequal); antheridial stalks uniseriate 3 3. Gemmae angulate to polyhedral to stellate; dorsal and ventral lobes oriented in differing directions; median lobe cells with at most small trigones; perianth weakly flattened, distinctly plicate, contracted toward the mouth Diplophyllum (p. 1 06) 3. Gemmae smooth, ovoid to ellipsoid; dorsal and ventral lobes similarly oriented; median lobe cells with trigones; perianth normally strongly flattened, eplicate, truncate at mouth [Scapania] ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 59 2. Leaves equally complicate-bilobed or nearly so; antheridial stalks biseriate. Dorsal lobe dorsally recurved . [Krunodiplophyllum] GROUP XIV: Cephaloziaceae, Lepidoziaceae (p.p., cf. also group VII) 1 . Rhizoids scattered on ventral surface of axis; vegetative shoots without distinct underleaves; leaves bifid, never 3-4-lobed; cells lacking oil-bodies; seta with 8 epidermal + 4 inner cell rows; plants sometimes with purple or red pigmentation. (Cephaloziaceae) 2 2. Leaf insertion transverse at least dorsally, extending to stem midline; pigmentation, when present, purplish or brownish; leaves ± canaliculate, piano-distichous; branching plastic, Frullania-lype branching always freely developed in addition to lateral-intercalary and ventral-intercalary branch- es; gynoecia terminal on leading axes Metahygrobiella (p. 89) 2. Leaf insertion oblique throughout, the dorsal end oblique to almost longitudinal; pigmentation never purplish, usually absent; leaves never piano-distichous, flat to concave but not canaliculate; branching normally of ventral intercalary type, Frullania-type branches rare (but present in C. badid)\ gynoecia normally on short, ventral branches which occasionally are elongated Cephalozia (p. 89) 1 . Rhizoids (often sparingly present) restricted to bases of underleaves; vegetative shoots with under- leaves distinct throughout; leaves often 3-4(or more)-lobed; seta with 8 or 16 large epidermal cell rows and many inner cell rows that are of much inferior diameter; plants without purple or red pigmentation. (Lepidoziaceae, p.p.) 3 3. Branching predominantly or nearly exclusively terminal, plastic, ofFrullania plus Microlepidozia and/or Acromastigum types 4 4. Terminal branches plastic, i.e., ofFrullania-, Microlepidozia (and in K. mollis, Acromastigum-) type, usually of Frullania type on one side of axis and Microlepidozia type on opposite side of axis; underleaves frequently with 1 segment shorter than others; stem with hyaloderm absent or at best weakly developed. Plants usually firm, the leaves transversely to succubously inserted, usually distant, usually 3-4(6)-lobed Kurzia (p. 71) 4. Terminal branches predominantly ofFrullania type, Acromastigum type sporadic to common, rarely of Microlepidozia type; underleaves not regularly possessing one segment shorter than others; stem with a well-developed hyaloderm. Plants commonly soft and flexuous, the leaves (in ours) transversely inserted, usually 2(-3)-dentate-lobate Paracromastigum7 (p. 76) 3. Branching nearly exclusively intercalary, terminal branches absent, at most rare or sporadic . . 5 5. Leaves bifid, unistratose throughout 6 6. Leaves bifid to 0.5 or more; secondary pigments lacking, plants green; cortical cells in 6 rows; terminal branching, when present, of Frullania and Microlepidozia types Hyalolepidozia 6. Leaves 2(3)-dentate or if lobate, then at most divided to 0.3; secondary pigments present, plants brown; cortical cells in 7-8 rows; terminal branching lacking Paracromastigum tristanianum 5. Leaves unlobed or 3-4(6) dentate to lobate, never bifid, polystratose toward the base in Amer- ican sector species 7 7. Leaves 3—4(6) dentate to lobate; leaf cuticle smooth or at most minutely roughened; stem hyaloderm present; androecia on leading shoots; seta (where known) with 8 rows of epi- dermal cells; capsule wall (where known) 2-stratose Pseudocephalozia (p. 78) 7. Leaves unlobed; leaf cuticle often papillose; stem hyaloderm absent; androecia on short, spicate, basal, intercalary branches; seta with 10-20 rows of epidermal cells; capsule wall 3-5-stratose Hygrolembidium (p. 69) 7 Paracromastigum at times may lack terminal branching, and for that reason the genus is also keyed under the second half of couplet 3. 60 FIELDIANA: BOTANY GROUP XV: Cephaloziellaceae (cf. also group XI), Gymnomitriaceae, Lophoziaceae (p.p., cf. also groups XI, XVI) 1. Leaves usually bilobed to 0.35-0.5 their length; leaf margins not bordered by 1-several rows of hyaline cells 2 2. Perianth not situated at the apex of a fleshy tube; a perigynium lacking; plants without stolons or flagella. Cells with trigones variable: at times lacking, at times coarse; cuticle often coarsely papillose 3 3. Plants minute or small, often wiry; leaves remote, bilobed to 0.5 or more their length. Cells very small, equally thick walled, lacking trigones; perianth mouth usually crenulate 4 4. Leaf lobes plane, 2-10 cells wide at base; leaves bifid to nearly 0.75 their length; seta normally with 4 epidermal cell rows and 4 internal cell rows; plants minute, the shoots usually less than 350 nm wide Cephaloziella (p. 90) 4. Leaf lobes sulcate, 9-28 cells wide at base; leaves bifid to nearly the base; seta with 8(9) epidermal cell rows and (3)4-14 internal cell rows; plants small, the shoots greater than 400 nm wide [Allisoniella] 3. Plants larger, not wiry; leaves bilobed to ca. 0.3-0.45(0.5) their length. Leaf lobes 8 or more cells broad at base 5 5. Leaves strongly dorsally secund, the dorsal end of the line of insertion arcuate and often ± decurrent along dorsal midline of stem. Cells (in Falkland species) with coarse, conspic- uous trigones and thin to sinuous intervening walls; branching variable, terminal, Fmllania type in some taxa, ventral-intercalary in some taxa, but if lateral intercalary then never arising from near the dorsal end of leaf or dorsal side of axis (Andrewsianthus-lype) Anastrophyllum (p. 92) 5. Leaves laterally patent and pectinate-distichous, not dorsally secund, the insertion dorsally transverse, not decurrent. Branching strictly intercalary 6 6. Leaf cuticle with conspicuous, very coarse, hyaline papillae; branching apparently strictly lateral intercalary, Andrewsianthus-lype branches lacking 7 7. Underleaves lacking; leaves large, stiffly, pectinately patent, with gibbous sinuses and reflexed margins; gynoecium without bracteole; perianth weakly plicate distally, the mouth crenulate; autoecious Acrolophozia (p. 103) 7. Underleaves small and ciliiform to large and ovate-lanceolate; leaves small, at least basally suberect, ± concave but not folded, the lobe margins not reflexed; gynoecium with large bracteole; perianth ± strongly plicate and lobulate-laciniate at mouth; dioecious Cephalolobus (p. 95) 6. Leaf cuticle (in Falkland species) smooth, at most rather coarsely striate; branches of Andrewsianthus type, arising from near the dorsal base of the leaf, occasionally appearing to arise from dorsal side of axis. Branches and main axis frequently becoming positively geotropic Andrewsianthus (p. 92) 2. Perianth included within the connivent or suberect bracts, associated with a distinct perigynium that subtends the joint insertion of bracts and perianth; plants with stolons or flagella. Cells with coarse trigones; cuticle smooth [Marsupella] 1. Leaves unlobed or bifid to 0.2; leaf margins often with 1-several rows of hyaline cells forming a border. Leaves erect or erect-appressed, cuticle smooth or finely papillose Herzogobryum (p. 104) GROUP XVI: Plagiochilaceae, Lophoziaceae (p.p., cf. also groups XI, XV), Jungermanniaceae 1. Perianth laterally sharply compressed, wide and truncate at mouth. Vegetative branches lateral in- tercalary and/or terminal and ofFrullania type, never ventral intercalary. Leaves with dorsal margin deflexed to form a convex fold or cnemis; rhizoids scattered; anthocyanin pigments lacking Plagiochila (p. 143) 1 . Perianths terete below, pluriplicate and contracted to the mouth 2 ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 61 2. Rhizoids fasciculate, at ventral end of leaf; leaves opposite, connate at least ventrally. Bracts and bracteole normally united; plants often with reddish to purplish anthocyanin pigments [Syzygiella] 2. Rhizoids scattered, often in dense mats on ventral side of stem; leaves alternate, free ventrally. Bracts and bracteole united or free 3 3. Leaves consistently 2^4-lobed. Underleaves laciniate to ciliate or reduced or absent .... 4 3. Leaves unlobed, usually rounded at apex. Underleaves reduced and inconspicuous 5 4. Ventral leaf margin frequently slightly to strongly reflexed, leaves short bifid; underleaves present, toward the stem apices and only of a few cells (in Falkland species) Nothostrepta (p. 96) 4. Ventral leaf margin plane, not reflexed, leaves 2-4-lobed, often deeply so; underleaves, when present, may be large and conspicuous (ciliate in the Falkland species, L. hatcheri) Lophozia (p. 96) 5. Involucral bracts unlobed and edentate; bracteole lacking; plants soft textured, usually prostrate growing, small in size; antheridial bracts (l)2-3-androus Jungermannia (p. 1 02) 5. Involucral bracts ± lobed and/or dentate, ± smaller than leaves; bracteole usually present; plants ± firm, usually erect growing, often robust; antheridial bracts monandrous 6 6. Stolons or flagellae regularly produced; terminal branching completely absent. Brac- teoles nearly as long as bracts; perianth mouth entire to weakly crenulate; plants often greenish to blackish brown; (455)610-700 m in Falklands Cryptochila (p. 98) 6. Stolons or flagella only rarely and sporadically produced, mostly absent; terminal branching present, but sometimes only occasional. Gynoecium as in Cryptochila; plants often red or magenta or red brown; 75-455(685) m in Falklands Jamesoniella (p. 99) GROUP XVII: Adelanthaceae, Acrobolbaceae 1 . Gynoecia and androecia on reduced, abbreviated, basal, intercalary branches; gynoecia producing an erect, massive shoot calyptra without or (in some taxa of Adelanthus) with a perianth; leaf cuticle smooth. Leaves undivided; plants erect, rigid 2 2. Stem 20-45 cells in diameter, with a collapsing hyaloderm (thus scabrous with age); dorsal leaf margin 4-5 cells thick (in Falkland species); plants green, without secondary pigmentation. Sub- hyalodermal cells in 2-4 layers of thick- walled cells Wettsteinia (p. 88) 2. Stem 8-15 cells in diameter, without a collapsing hyaloderm, smooth, never roughened; leaves unistratose throughout; plants green to red brown Adelanthus (p. 82) 1 . Gynoecia and androecia on terminal or leading leafy shoots (except some populations of Tylimanthus, where on short, basal, intercalary branches); gynoecia producing a marsupium; leaf cuticle often conspicuously roughened. Leaves undivided or 2-4-lobed; plants erect or appressed; bracts of in- nermost series smaller than lower bracts and often reduced to laciniiform projections; capsule tip beaked 3 3. Leaves unlobed, leaf margins entire; marsupium slenderly cylindrical, deeply penetrating sub- stratum; unfertilized archegonia carried downward with marsupial canal, inserted on surface and base of free calyptra. Plants closely prostrate Lethocolea (p. 149) 3. Leaves (in Falkland species) 2-4 lobed (or if undivided, then with leaf margins dentate to lobate), leaf margins entire-den tate-lobate; marsupium ± conoidal to wide cylindrical in shape, not deeply penetrating substratum; unfertilized archegonia remaining external to and at summit of marsupium 4 4. Leafy shoots ± ascending, not adhering to substratum by rhizoids. Gynoecia and androecia variable in position, acrotonic or basitonic; plants a clear, translucent to opaque green, often whitish when dead Tylimanthus (p. 151) 62 FIELDIANA: BOTANY 4. Leafy shoots ± creeping, often entire axis, including shoot tips, adhering to the substratum by rhizoids 5 5. Leaves (in ours) bilobed; cuticle coarsely papillose; rhizoids few, arising from ventral leaf bases, never in dense, decurrent bundles on stem; plants without gemmae; antheridia sol- itary; stolons and flagellae common Acrobolbus (p. 148) 5. Leaves 3-5 lobed; cuticle smooth; rhizoids dense, decurrent along stem in fascicles and forming a ventral mat; plants gemmiparous; antheridia 1-10 per bract; stolons and flagellae lacking Austrolophozia (p. 149) Family VETAFORMACEAE Vetaformaceae Fulf. & J. Tayl., Nova Hedwigia 1:405. 1960. A monotypic family represented by Vetaforma dusenii (Steph.) Fulf. & J. Tayl. of southern South America, but not the Falklands. For details see Fulford (1963b) and Hassel de Menendez and So- lari(1975). Family LEPICOLEACEAE Lepicoleaceae Schust., Nova Hedwigia 5: 27. 31 Jan. 1963. Schust. ex Fulf. in Fulford, Mem. New York Hot. Card. 1 1 : 30. 1 5 March 1 963. Schust., Rev. Bryol. Lichenol. 26: 126. 1957, nom. inval. A family of one genus with ca. 1 0 species. Lep- icolea occurs in temperate areas of South America, Africa, and Australasia and extends to the North- ern Hemisphere in both Asia and Latin America. Lepicolea Lepicolea Dum., Recueil Observ. Jungerm. 20. 1835. Key to Falkland Islands Species of Lepicolea 1. Leaf segments ending in a uniseriate tip of a few to several elongate, noncaducous cells, the 2nd, 3rd, and 4th cells from tip elongate, the tip cell not a slime papilla; oil-bodies hyaline, 4-7 per cell (fide Inoue, 1 984) L. ochroleuca 1 . Leaf segments ending in a uniseriate tip of a few-celled row of short (subquadrate to short-rectangular) cells which are often caducous; the 3rd and 4th cells from the tip short, the tip cell a slime papilla; oil-bodies grayish, 1-3(4) per cell L. rigida Lepicolea ochroleuca (Spreng.) Spruce. Figure 23. Jungermannia ochroleuca Spreng., Syst. Veg. 4(2): 325. 1827. Sendtnera ochroleuca (Spreng.) Nees in G. L. & N., Syn. Hep. 240. 1845. Leperoma och- roleuca (Spreng.) Mitt, in Hook, f., Handb. New Zealand Fl. 2: 754. 1867. Herbertia ochroleuca (Spreng.) Trev., Mem. Reale 1st. Lomb. Sci. Lett. Ill, 4: 397. 1877. Lepicolea ochroleuca (Spreng.) Spruce, Trans. & Proc. Hot. Soc. Edinburgh 15: 345. 1885. Original material: South Africa, Cape of Good Hope, Ecklon s.n. (NY!, STR, cited in Scott, 1960). Jungermannia hirsuta Nees ex Hook. f. & Tayl., Lon- don J. Bot. 3: 289 (in errore pro 389), 475. 1844, nom. nud. Sendtnera ochroleuca 0 mexicana Gott., Kongel. Danske Vidensk. Selsk. Naturvidensk. Math. Afh. II, 6: 140. 1863, syn. fide Scott (1960). REMARKS— This is the first authentic report of Lepicolea ochroleuca from the Falkland Islands, as all previous reports (Gottsche et al., 1845 as Sendtnera; Hooker & Taylor, 1844, Taylor & Hooker, 1 847 as Jungermannia hirsuta) were based upon a Hooker collection which is actually L. rig- ida. ECOLOGY— Rare in the Falklands, where it is known from only three stations. The Mt. Kent plant occurred at the summit, 455 m, on wet rock walls where only a few colonies were observed. The species also was gathered deep in a cool, cave- like hollow of a rock outcrop in a dwarf shrub heath (180 m). Here, as in the above locale, the plants (again, only a few) grew on an extensive, flattish area of the outcrop. Longton and Smith ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 63 23 FIG. 23. Distribution of Lepi- colea ochroleuca (Spreng.) Spruce, plus Honduras north to Mexico. FIG. 24. Distribution of Lepi- colea rigida (De Not.) Scott, plus South Georgia. Open circle indi- cates precise locality unknown. 64 FIELDIANA: BOTANY encountered the species at 1 85 m on damp ground among boulders on the south face of Mt. William. Lepicolea ochroleuca is common in the wet Ma- gellanian moorland, where it may form extensive mounds over bark. It is also quite common on Nothofagus bark in the Valdivian region. The species appears to require a high rainfall and cool temperatures, the absence of the former (as well as the bark habitat) presumably accounts for the rarity of the taxon in the Falklands. The Port Stan- ley region has cooler temperatures and a higher rainfall than other Falkland regions, a point which almost undoubtedly accounts for the presence of L. ochroleuca in this part of the Falklands. PHYTOGEOGRAPHY — Amphiatlantic — South Af- rica; Tristan da Cunha; Falkland Is.; Tierra del Fuego; Patagonian Channels; Valdivian region (West Patagonia north to 39°16'S, Andean Pata- gonia at P. N. Nahuel Huapi); Juan Fernandez Is.; also known from Brazil (Stephani, 1909; Lor- scheitter, 1973). Reported from Bolivia (Spruce, 1890), Hon- duras and Guatemala (Fulford, 1963b; Hassel de Menendez & Solari, 1976), and Mexico (Scott, 1960; Fulford, 1963b; Hassel de Menendez & So- lari, 1 975). The report from India in Hooker (1 867) is regarded as erroneous. The New Zealand reports are either L. attenuata or L. scolopendra, and that from Tasmania is likely L. scolopendra (fide ci- tations in Scott, 1960) (Fig. 23). Hassel de Me- nendez and Solari (1975, p. 37) include New Zea- land, Auckland Island, and Campbell Island in the range of this species. FALKLAND SPECIMENS SEEN— EAST FALKLANDS. Stanley Region: S slopes of Mt. William, 1 85 m, Longton & Smith. Longton 992. 193 (AAS, F); Goat Ridge, ca. 1 80 m (3195); summit of Mt. Kent, 455 m (2740. 2742). Lepicolea rigida (De Not.) Scott. Figure 24. Sendtnera rigida De Not., Mem. Reale Accad. Sci. Torino II, 16: 229. pi. XV, 1-9. 1855. Herbertia rigida (De Not.) Trev., Mem. Reale 1st. Lomb. Sci. Lett. Ill, 4: 397. 1877. Leperoma rigida (De Not.) Mass., Nuovo Giom. Bot. Ital. 17: 252. 1885. Lepicolea rigida (De Not.) Scott, Nova Hedwigia 2: 148. 1960. Original material: Chile, "Prov. Valparaiso, Valparaiso," Puccio (NY!). Lepicolea seriata Herz., Hedwigia 66: 9 1 . / 8. 1 926, syn. fide Scott ( 1 960). Lepicolea ochroleuca var. seriata (Herz.) Herz. in Skottsberg, Nat. Hist. Juan Fernandez Is., Easter Is. 2: 728. 1942. Original material: Chile, Prov. Aisen, Pta. Leopardos, Reichert & Hicken s.n. (JE), cited in Scott (1960) (non vidi). ECOLOGY— See comments for the ecology of L. ochroleuca, which also apply here. Hooker visited only East Falkland and it is likely that his collec- tion was made in a niche similar to those of L. ochroleuca. PHYTOGEOGRAPHY— Valdivian + Magellanian + Falklands— South Georgia; Falkland Is.; Tierra del Fuego; Patagonian Channels; Valdivian region north to 46°32'S, 73°52'W (Punta Leopardos) (fig. 24). FALKLAND SPECIMEN SEEN— EAST FALKLANDS. 1 843, Hookers. n. as Jungermannia hirsuta "with a Frul- lania allied to Jung, lobulata Hook." (FH). Family HERBERTACEAE Herbertaceae K. Mull. (Freib.) ex Fulf. & Hatch., Bryologist 61: 284. 23 Feb. 1959. K. Mull. (Freib.) in Rabenh., Krypt.-Fl. Deutschl. ed. 3, 6(1): 557. 1954, nom. nud. Two genera belong here. Triandrophyllum has ca. four species; it is south temperate but extends north to Guatemala in the Neotropics and to Java in the Paleotropics. Herbertus has ca. 75 species, and occurs rather commonly in wet mossy forests of temperate, but more predominantly, tropical regions. Triandrophyllum Triandrophyllum Fulf. & Hatch., Bryologist 64: 349. 1961 [1962]. Fulf. & Hatch., Bryologist 61: 277. 1958 [1959], nom. inval. sin. gen. typ* Grolle, Bryologist 64: 25. 1961, nom. inval. Triandrophyllum subtrifidum (Hook. f. & Tayl.) Fulf. & Hatch. PHYTOGEOGRAPHY — Amphipacific Temper- ate—Macquarie Is., New Zealand, Tasmania, and the Philippines (Kitagawa, 1981). In the American zone in Tristan da Cunha, South Sandwich Is., South Georgia, Falkland Is., southern South America, and north in the Andes to Guatemala. • See notes in Engel ( 1978). ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 65 Variety trifidum (Gott.) Solan is Andean Ameri- can in distribution, and var.fuscum (Steph.) Solari occurs only in southern South America. REMARKS— Triandrophyllum subtrifidum and T. trifidum had been regarded as closely related species until Solari (1973) rightfully treated them as two varieties of a single entity, T. subtrifidum. Solari also included a third variety, var.fuscum, of south- ern South America. The typical variety differs from var. trifidum by underleaf margins and ventral margins of the leaf entire or occasionally with a few small teeth, leaf cell walls thin to very slightly thickened, and leaves generally bifid. Variety tri- fidum, on the other hand, has underleaf margins and ventral margins of the leaf with several sharp teeth, especially toward the base; leaf cell walls uniformly thickened; and leaves generally trifid. Only the typical variety occurs in the Falklands. Triandrophyllum durum, which was reported for the Falklands by Grolle (1962), has had a check- ered taxonomic position. Fulford and Hatcher (1958) transferred the species to Triandrophyllum, but in Fulford and Hatcher (1961) they placed it as a synonym of their new combination, T. trifi- dum (Gott.) Fulf. & Hatch., which is an older name. Fulford (1963b) included T. durum as a synonym of T. trifidum (even though seeing the correct type), and, based upon these sources, I did the same in Engel (1978). However, Solari (1973) moved the plant to the synonymy of T. subtrifidum var. sub- trifidum, which appears to be correct. The species was similarly placed in Hassel de Menendez and Solari (1975). Fulford and Hatcher (1958, p. 283, sub plate legend) erroneously stated that the type of Herberta dura is a Dusen collection from Fue- gia, and included figures based upon that plant. These figures show the marginal teeth typical of T. trifidum, and it naturally follows that they would place the plant there. This erroneous designation of type specimen is probably the source of error in taxonomic placement. Triandroph) Hum subtrifidum (Hook. f. & Tayl.) Fulf. & Hatch, var. subtrifidum Jungermannia subtrifida Hook. f. & Tayl. London J. Bot. 3: 579. 1844. Isotachis subtrifida (Hook. f. & Tayl.) Mitt, in Hook, f., Bot. Antarc. Voyage 2(2): 149. 1854. Mastigophora subtrifida (Hook, f. & Tayl.) Hodgs., Rev. Bryol. Lichenol. 18: 25. 1949. Triandrophyllum subtrifidum (Hook. f. & Tayl.) Fulf. & Hatch., Bryologist 64: 350. 1961 [1962]. Original material: Tasmania, Hooker (s!). Herberta dura Steph., Hedwigia 34: 44. 1 895, syn.fide Solari (1973). Schisma dura (Steph.) Steph., Spec. Hep. 4: 2 1 . 1 909. Triandrophyllum durum (Steph.) Fujf. & Hatch., Bryologist 61: 279. 1958 [1959]. Original material: "Fretum magellanicum. leg. Hooker fil. Herb Kew, sub nomine Jung, tena- cifolia" (G), cited in Hassel de Menendez & Solari (1975) and Solari (1973); (K), cited in Fulford (\963b) (non vidi). Leioscyphus schismoides Steph., Kongl. Svenska Ve- tenskapsakad. Handl. 46(9): 38. f. 13f-g. 1911, syn. fide Grolle (1964c). Leptoscyphus schis- moides (Steph.) Kiihnem., Revista Centre Estud. Doct. Cien. Nat. 1: 177. 1937. Mylia schismoides (Steph.) Kiihnem., Lilloa 19: 341. 1949. Original material: Falkland Is., Mt. Adam, 700 m, 13 De- cember 1907, Halle & Skottsberg 347 (s!). Isotachis lanciloba Steph., Kongl. Svenska Veten- skapsakad. Handl. 46(9): 70. / 26f-g. 1911, syn. fide Fulford & Hatcher (1958). Original material: Falkland Is., Mt. Usborne, Skottsberg s.n. (G), cited in Fulford (1963b) (non vidi). For a complete synonymy see Solari (1973) and Hassel de Menendez and Solari (1975); only syn- onyms of Falkland species are included here. Triandrophyllum durum is also included; see Re- marks under genus. ECOLOGY— In the Falklands, of sporadic occur- rence on soil or rock under rock ledges of sheltered high altitude cliffs. Rarely in stands of Hebe in dwarf shrub heaths. LITERATURE RECORDS (FALKLANDS)— Anony- mous (Fulford & Hatcher, 1958, 1961; Kuhne- mann, 1 937, as Isotachis lanciloba & Leptoscyphus schismoides; 1949, as /. lanciloba & Mylia schis- moides^ Skottsberg (Fulford, 1963; Hassel de Me- nendez & Solari, 1975, and Solari, 1973, as var. subtrifidum; Stephani, 1922, as 7. lanciloba); Mt. Usborne (Bonner, 1966, as I. lanciloba); Mt. Adam (Hassel de Menendez & Solari, 1975, as var. sub- trifidum; Skottsberg, 1913, as Leioscyphus schis- moides; Solari, 1973, as var. subtrifidum; Ste- phani, 1922, as L. schismoides); Skottsberg & Halle, Mt. Adam (Grolle, 1962, as T. durum). FALKLAND SPECIMENS SEEN -EAST FALKLANDS. Mt. Usborne Region: ridge between Mts. Usborne 1 & 2, 685 m (25/6. 2531B, 2552). WEST FALKLANDS. Mt. Adam: 700 m, 13 Dec. 1907, Halle & Skottsberg 347, syntype of Leioscyphus schismoides (s); ridge S of northern lake, 610m (3026, 3036A, 3040, 3042B). West- point Island: near The Waterfall, 30-90 m (2869— c. per.). Family PSEUDOLEPICOLEACEAE Pseudolepicoleaceae Fulf. & J. Tayl., Nova Hed- wigia 1: 411. 1960. 66 FIELDIANA: BOTANY The nine genera that comprise this family are all small; the largest, Temnoma, has only ca. 10 species. The family is chiefly south temperate-sub- antarctic in range. Archeochaete a Sphagnum bog in Tierra del Fuego, and I have found it in the same habitat in the Brunswick Pen- insula. This Magellanian-Falkland species is oth- erwise known only from several stations in Tierra del Fuego (Hassel de Menendez & Solari, 1975), who state the species occurs at wet edges of peat bogs. The ecology of the Falkland plants thus de- viates considerably from that of South American collections. Archeochaete Schust., J. Hattori Bot. Lab. 26: 262. 1963. Archeochaete kuehnemannii Schust. FALKLAND SPECIMENS SEEN- EAST FALKLANDS. Mt. Usborne Region: summit of Mt. Usborne 1, ca. 700 m (2496D). WEST FALKLANDS. Mt. Adam: ridge S of northern lake, 610 m (3046). Archeochaete kuehnemannii Schust., J. Hattori Bot. Lab. 26: 262. 1963. Pseudolepicolea kuehneman- nii (Schust.) Hassel, Comun. Mus. Argent. Cien. Nat. "Bernardino Rivadavia," Bot. 2: 48. 1974, nom. inval. P. kuehnemannii (Schust.) Hassel in Hassel de Menendez & Solari, Fl. Cryptog. Tierra del Fuego 15: 125. 1975 [1976]. Holotype: Ar- gentina, Terr. Tierra del Fuego, ca. 16-17 km W of Ushuaia, on road to Lapataia, C. Bandera, Feb- ruary 1 96 1 , Schuster & Gamundi de Amos 58852 (hb. Schuster, non vidi). REMARKS— Schuster (1965b), in his description of Archeochaete kuehnemannii, states the leaf lobe apices are uniseriate for a length of 4-6(7) cells. The Falkland plants, however, have lobe apices uniseriate for a length of (1)2-4 cells. Schuster (1965b, p. 816), in distinguishing Archeochaete, states: The more attenuate leaf apices, with the uniserate tips longer than in any species of Lophochaete, are (also) distinctive. In Lopho- chaete the tips are formed by, at most, 2-3 superimposed cells. While the genera Pseudolepicolea (= Lophochaete) and Archeochaete are distinct and well denned, the leaf lobe apices should not be used as a character to differentiate them. Pseudolepicolea has leaves and underleaves subequal in size and form and bisbifid, and perianths 4-5-6 plicate distally. Ar- cheochaete has underleaves at most 0.5 the leaf size and leaves, when quadrifid, with sinuses sub- equal, and perianths 3-plicate distally (see also Schuster, 1966c). The only other member of the genus, A. tern- nomoides Schust., is endemic to Tristan da Cunha. ECOLOGY-PHYTOGEOGRAPHY— Rare; found in a feldmark pool and on a cushion plant on a shel- tered high altitude cliff. The type was collected in Temnoma Temnoma Mitt, in Hooker f., Handb. New Zea- land Fl. 753. 1867. Teinnoma Mitt., Philos. Trans. 168(extra vol.): 32, 33. 1879, err. ty- pogr.9 (et Mitten, 1884). Temnoma quadripartitum (Hook.) Mitt. Figure 25. Jungermannia quadripartita Hook., Musci Exot. 2: pi. 117, f. 1-3. 1820. Temnoma Quadripartitum (Hook.) Mitt., J. Linn. Soc. Bot. 15: 68. 1876. Blepharostoma quadripartitum (Hook.) Trev., Mem. Reale 1st. Lomb. Sci. Lett. Ill, 4: 4 1 7. 1 877. Original material: Argentina, Terr. Tierra del Fuego, I. de los Estados, Menzies (K, v), cited in Schuster (1966d) (non vidi). Jungermannia podophylla Angstr., Ofvers. Forh. Kongl. Svenska Vetenskapsakad. 29(4): 11. 1872, syn. fide Pearson (1887); non Jungermannia po- dophylla Thunb., Prodr. Fl. Cap. 2: 174. 1823 (= Symphyogyna). Original material: Chile, Prov. Magallanes, Pto. del Hambre, Andersson s.n. (s), cited in Schuster ( 1 966d) (non vidi). Lepidozia randii S. Arnell, Svensk Bot. Tidskr. 47(3): 417./ 6. 1953, syn. fide Schuster ( 1 966d). Orig- inal material: Marion Is., Rand 3276 (s), cited in Schuster (1966d) (non vidi). Temnoma subintegrum Steph. ex Fulf, Mem. New York Bot. Gard. 1 1: 57. 1963, syn. fide Schuster ( 1 966d). Blepharostoma quadripartitum var. sub- integrum Steph., Icon. Hep. Blepharostoma no. lOb, nom. nud. Original material: Chile, Prov. Magallanes, Pto. Bueno, Dusen 46 (o, H), cited in Fulford (1963b) and Schuster (1966d), respec- tively (non vidi). REMARKS— The Falkland material is referable to var. quadripartitum and not var. pseudopungens 9 See Engel ( 1 978, p. 69) for notes on the typographical error "Teinnoma." ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 67 25 FIG. 25. Distribution of Temnoma quadripartita (Hook.) Mitt. Schust. or var. randii (S. Arnell) Schust. (See Schuster, 1 966d, for a key to varieties.) ECOLOGY— This taxon is rather rare in the Falk- lands, where only a few gatherings were made in Cortaderia (wet depressions) and dwarf shrub heaths (under rock overhangs of outcrops). PHYTOGEOGRAPH Y — Amphipacific Temper- ate—northward extensions in the American sector (West Patagonia north to 39°16'S; Andean Pata- gonia at P. N. Nahuel Huapi and in the New Zea- land sector into the mountains of North Is. Also in the subantarctic, on Kerguelen Is., Crozet Is., Marion Is., and Prince Edward Is. The report from Tristan da Cunha in Arnell (1958) is based on a specimen of Archeochaete temnomoides (Schuster, 1966c). The report from Inaccessible Is. in Arnell (1958) is questionable (fig. 25). FALKLAND SPECIMENS SEEN -EAST FALKLANDS. Mt. Usborne Region: The Gap, 275-290 m (2470-c., S + young 9, 2472). WEST FALKLANDS. Fox Bay Region: summit of East Head, 180 m (3428— c. per.); summit of Fox Bay Mt., 307 m (3446— c. $). Family TRICHOCOLEACEAE Trichocoleaceae Nakai in Ogura, Ordines, fami- liae . . . Prof. Nakai-Takenosin ut novis edita. 200. 1943. 68 FIELDIANA: BOTANY Trichocoleaceae have 3 genera. Trichocolea, with about 30 species, occurs primarily in Neotropical and south temperate mossy rain forests; one species, T. elegans, occurs in southern South America but is absent from the Falkland's. Leio- mitra has ca. 4 species, all Neotropical. Eotricho- colea has 1 species, found in New Zealand and Tasmania. Family LEPIDOZIACEAE Lepidoziaceae Limpr. in Cohn, Krypt.-Fl. Schles. 1: 310. 1875 ("Lepidozieae"). The Lepidoziaceae are a large, exceedingly poly- typic family of seven subfamilies and 28 genera. The family is represented well in south temperate and Neotropical regions, less so in the Paleotrop- ics, and poorly in the Northern Hemisphere. Hyalolepidozia Hyalolepidozia S. Arnell ex Grolle, Rev. Bryol. Lichenol. 32: 1 79. 1 964 [ 1 963]. S. Arnell, Hot. Not. 115: 203. 1962, nom. inval. syn. descr. lat. Hyalolepidozia bicuspidata (Mass.) S. Arnell ex Grolle. Figure 26. Lepidozia bicuspidata Mass., Nuovo Giorn. Bot. Ital. 17: 239. pi 22, f. 25. 1885. Hyalolepidozia bi- cuspidata (Mass.) S. Arnell, Bot. Not. 115: 213. 1962, nom. illeg. Paracromastigum bicuspidatum (Mass.) Schust., J. Hattori Bot. Lab. 26: 276. 1963. Hyalolepidozia bicuspidata (Mass.) S. Arnell ex Grolle, Rev. Bryol. Lichenol. 32: 1 79. 1963. Orig- inal material: Argentina, Terr. Tierra del Fuego, Is. de los Estados, Pto. Cook, Spegazzini 45b (G), cited in Fulford (1968) (non vidi). REMARKS— After a thorough survey of branch types in the Falkland material of H. bicuspidata, I found both lateral- and ventral-intercalary branches in about equal frequency, and only a sin- gle occurrence of a Fr«//ama-type branch. I was unable to find Microlepidozia-lype branching. Terminal branching is only rarely developed in the genus; in H. bicuspidata, Grolle (1963b) found two Frullania-lype branches (Chilean material) and one Microlepidozia-type branch (South African material). The branching pattern in Hyalolepido- zia readily distinguishes the genus from the closely related genera Bonneria and Paracromastigum. In Bonneria the branching is plastic with Acromas- tigum-, Frullania-, Microlepidozia- and ventral- intercalary branches (with lateral-intercalary branches absent). Paracromastigum has Acromas- tigum- and Frullania-lype branches and rarely ventral-intercalary branches (with Microlepido- zia- and lateral-intercalary branches absent). (See Schuster, 1965a for a discussion of the evolution and relationships within this complex of genera.) Fulford (1968, p. 277), in her generic key, uti- lizes as her primary key character for Hyalolepi- dozia, "underleaves and leaves similar in size and form." This statement presumably is based upon examination of the type, although the ventral view of the stem (her plate 101, fig. Ib) is from nontype material. Massalongo (1885), in his original de- scription of "Lepidozia" bicuspidata, gives "Am- phigastria fol. 2-4-plo minora . . . ," and his fig- ures clearly show the underleaves as considerably smaller than the leaves. The underleaves of the Falkland material of H. bicuspidata are ca. 0.33- 0.50 the leaf size and are never equal to the size of the leaves. ECOLOGY— Collected in dwarf shrub heaths, where it was found on soil, particularly on soil over rock. PHYTOGEOGRAPHY — Amphiatlantic Temper- ate—South Africa, Table Mt.; southern Patagonia, Brunswick Peninsula and Rio Rubens; Patagonian Channels; in Valdivian region north to 41°46'S in West Patagonia; Juan Fernandez Is.; and "Frai Jorge" (no. Chile) (see fig. 26). FALKLAND SPECIMENS SEEN— EAST FALKLANDS. Port William Region: summit ridge of Mt. Low, ca. 245 m (3233). Stanley Region: Sapper Hill, 135 m (3402). Mt. Usborne Region: SE slope of Mt. Usborne 2, ca. 455 m (2612E). WEST FALKLANDS. Westpoint Island: steep slope and cliffs facing The Woolly Gut (2974. 2924, 2931, 2932-c. per.). Hygrolembidium Hygrolembidium Schust., J. Hattori Bot. Lab. 26: 277. 1963. Hygrolembidium isophyllum Schust. Figure 27. Hygrolembidium isophyllum Schust., Nova Hedwigia 15: 467. pi. 56. 1968. Holotype: Argentina, Terr. Tierra del Fuego, C. Garibaldi, near L. Escon- dido, Schuster 583 19e (hb. Schuster!). REMARKS— See Engel (1974) and Schuster and Engel(1987). ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 69 26 70 FIELDIANA: BOTANY ECOLOGY— In the Falklands this species seems to be restricted to sheltered high altitude cliffs, where it grows on soil under rock overhangs or on Azorella cushions. In the Brunswick Peninsula I found the species only in the relatively dry eastern end (8 km west of Punta Arenas, 305-610 m), where it grew on soil among cushion plants. Ray- mond E. Hatcher made several collections of the taxon between Punta Arenas and Puerto Natales (near Rio Rubens), ca. 100 km north of the Strait of Magellan, a locality which is on the eastern side of the Andes and thus relatively dry. On the main- land this taxon seems to be restricted to deciduous Nothofagus forests of Magellanian South America. PHYTOGEOGRAPHY— Antarctic Peninsula— Cape Tuxen, Powell and Coronation Is. (Ochyra and Vana, 1989); South Orkney Is.; Signy Is. (Smith, 1972, p. 24); South Georgia; Falkland Is.; moun- tainous region of Isla Grande de Tierra del Fuego; southern Patagonian Channels in Brunswick Pen- insula and Rio Rubens area. This species is treated as having an Antarctic distribution. Northward extension from the Antarctic zone is at high or relatively high elevations, with the curious excep- tion of the Rio Rubens station (see Engel, 1973b) (fig. 27). FALKLAND SPECIMENS SEEN— EAST FALKLANDS. Mt. Usborne Region: ridge between Mts. Usborne 1 & 2, 685 m (2522, 2556). WEST FALKLANDS. Port Howard: pass SW of Mt. Maria summit, ca. 6 10 m (3102). Mt. Adam: E side of summit ridge, 670-700 m (3008); summit of southernmost peak, 685 m (2991). Kurzia Kurzia v. Mart., Flora 53: 417. 1870. Microlepidozia (Spruce) J0rg., Bergens Mus. Skr. 16: 303. 1934. Lepidozia subg. Microlepidozia Spruce, J. Bot. 14: 165. 1876. Micrisophylla Fulf., Brittonia 14: 124. 1962, syn.fide Schuster, 1969b, p. 41. REMARKS— See Engel (1978) for discussion of placement of the genus Micrisophylla. See also the discussion in Schuster (1980c). Through an oversight, I (Engel, 1978) neglected to make the formal transfer of Micrisophylla to subgeneric rank in Kurzia; this was done in Schuster ( 1 980c, p. 347). Key to Falkland Islands Species of Kurzia 1 . Leaves and underleaves stiff, erect, strongly spreading, bristle-like; leaf lobes ending in a uniseriate row of 2-5 cells; plants, at least in a considerable apical portion, green; Acromastigum-type branches frequently produced K. mollis 1. Leaves and underleaves suberect, leaf lobes usually curved towards the stem, leaves lax, not bristle- like; leaf lobes ending in a uniseriate row of 1-3 cells; plants deep brown throughout, or occasionally golden brown at apices; Acromastigum-lype branches not present 2 2. Leaf lobes (4)5-1 2 cells wide at base; Microlepidozia-lype branching rare and sporadic, branching irregularly pinnate, lacking regularly pinnate alternation of Microlepidozia and Frullania-lype branches; plants ± large, robust K. setiformis 2. Leaf lobes (2)3^4(7) cells wide at base; Microlepidozia-lype branching common, branching fre- quently with ± regularly pinnate alternation of Microlepidozia and Frullania types; plants small, 220-400 n wide, delicate K. saddlensis Opposite: FIG. 26. Distribution of Hyalolepidozia bicuspidata (Mass.) S. Arnell. FIG. 27. Distribution of Hygrolembidium isophyllum Schust. ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 71 kur/ia mollis (Steph.) Engel & Schust. Lepidozia mollis Steph., Spec. Hep. 3: 601. 1909. Mi- crisophylla mollis (Steph.) Fulf., Brittonia 14: 131. 1962. Kurzia mollis (Steph.) Engel & Schust. in Engel, Bryologist 79: 514. 1976. Original mate- rial: Chile, Prov. Magallanes, Pto. Bueno, Dusen s.n. (G), cited in Fulford (1962) (non vidi). ECOLOGY— All three Kurzia taxa are rare in the Falklands: K. mollis was gathered in a small depression at 275-290 m in a Cortaderia heath. PHYTOGEOGRAPHY— Magellanian-Falkland — Falkland Is.; Tierra del Fuego; Patagonian Chan- nels north to 50°59'S. FALKLAND SPECIMEN SEEN— EAST FALKLANDS. Mt. Usborne Region: The Gap, 275-290 m (2465— c. per. + Kurzia saddlensis (Besch. & Mass.) Grolle Lepidozia Senlensis Besch. & Mass., Bull. Mens. Soc. Linn. Paris 1(80): 637. 1886. Micrisophylla sad- dlensis (Besch. & Mass.) Fulf., Brittonia 14: 128. 1962. Kurzia saddlensis (Besch. & Mass.) Grolle, Rev. Bryol. Lichenol. 32: 174. 1963.'° Lectotype (cf. Fulford, 1966): Chile, Prov. Magallanes, I. Saddle, Hariot (PC, non vidi). Lepidozia diver sifolia Steph., Kongl. Svenska Veten- skapsakad. Handl. 46(9): 62. / 23c-f. 191 1, syn. fide Fulford (1962). Original material: Chile, Prov. Chiloe, I. Guaitecas, Melinca, Skottsberg s.n. (G), cited in Fulford (1962) (non vidi). REMARKS— The 1902 Skottsberg collections of Kurzia saddlensis (Lepidozia saddlensis) from the Falkland Islands in Stephani (1905a) are misde- terminations of Lepidozia chordulifera. ECOLOGY— Very rare in the Falklands, where it was mixed with other hepatics (Clasmatocoleaful- vella, Gackstroernia magellanica and Riccardia papillosa) in wet depressions of a Cortaderia heath (455 m). PHYTOGEOGRAPHY— Valdivian + Magellanian- Falkland— Falkland Is.; Tierra del Fuego; Valdi- vian West Patagonia at 43°57'S. Also Tristan da Cunha, Nightingale, and Inaccessible Is. I regard the report from Tasmania in Rodway (1916) as doubtful. LITERATURE RECORDS (FALKLANDS)— Anony- mous (Kuhnemann, 1937, 1949); Skottsberg, Port Stanley & Port Louis (Skottsberg, 1913). FALKLAND SPECIMENS SEEN— EAST FALKLANDS. Mt. Usborne Region: SE slope of Mt. Usborne 2, ca. 455 m (2594B-C. 6, 2597B). Kurzia setiformis (De Not.) Engel & Schust. Lepidozia setiformis De Not., Mem. Reale Accad. Sci. Torino II, 16: 255. / XIII, 1-6. 1855. Mastigo- phora setiformis (De Not.) Trev., Mem. Reale 1st. Lomb. Sci. Lett. Ill, 4: 416. 1877. Micrisophylla setiformis (De Not.) Fulf., Brittonia 14: 127. 1962. Kurzia setiformis (De Not.) Engel & Schust. in Engel, Bryologist 79: 514. 1976. Neotype" (fide Fulford, 1 966): Argentina, Terr. Tierra del Fuego, Is. de los Estados, Speggazzini s.n. (G 344! ex hb. Massalongo M 52). Original material: Chile, "Prov. Valparaiso, Valparaiso," Puccio (non vidi). Lepidozia obscura Angstr. in Steph., Spec. Hep. 3: 602. 1909, syn. fide Fulford (1962). Original ma- terial: Chile, Prov. Magallanes, Strait of Magel- lan, Pto. del Hambre, Andersson s.n. (FH!, G!). Lepidozia fusca Steph., Kongl. Svenska Vetenskap- sakad. Handl. 46(9): 64. / 24h-i. 191 1, syn. fide Fulford (1962). Original material: Chile, Prov. Magallanes, S. Skyring, B. Pinto, Halle & Skotts- berg 170 (G, s), cited in Fulford ( 1 962) (non vidi). Lepidozia cunninghamii Steph., Spec. Hep. 6: 322. 1922, syn. fide Fulford (1962). Original material: Chile, Prov. Magallanes, Pto. Grappler, Cun- ningham inter no. 179 (G), cited in Fulford ( 1 962) (non vidi). ECOLOGY— Very rare in the Falklands, where it was gathered in a wet depression of a Cortaderia heath (455 m) mixed with other hepatics, nearly all of which are rare or relatively so, e.g., Telaranea oligophylla, Clasmatocolea obvoluta var. cook- iana, Hyalolepidozia bicuspidata, and Leptoscy- phus aequatus. PHYTOGEOGRAPHY— Magellanian-Falkland — Falkland Is.; Tierra del Fuego; Patagonian Chan- nels north to 48°55'S. Reportedly disjunct in Tas- mania (fide Fulford, 1962, 1966). FALKLAND SPECIMEN SEEN- EAST FALKLANDS. Mt Usborne Region: SE slope of Mt. Usborne 2, ca. 455 m (2612B). Lepidozia Lepidozia (Dum.) Dum., Recueil Observ. Jun- germ. 19. 1835, nom. cons. Pleuroschismasecl. Lepidozia Dum., Syll. Jungerm. Eur. Indig. 69. 1831. 10 Grolle incorrectly cited the basionym reference as Besch. & Mass. Mission Sci. du Cap Horn 5: 233. 1889. " See Engel (1978) for notes on the neotype locality. 72 FIELDIANA: BOTANY Key to Falkland Islands Species of Lepidozia 1 . At least some leaves and/or underleaves on well-developed stems with marginal teeth-lacininae . . L. chordulifera 1 . Margins of leaves and underleaves entire, very rarely with an isolated tooth 2 2. Leaf segments in conspicuous pairs, the dorsal and ventral segments often reduced to a tooth of a few cells; leaves often distant, often scalelike; (only above 245 m in Falklands, common on sheltered high altitude cliffs) L. fuegiensis 2. Leaf segments not in conspicuous pairs, dorsal and ventral segments larger, never reduced to a few cells; leaves imbricate, leafy, never scalelike; (rather common below 245 m in Falklands, absent on sheltered high altitude cliffs) L. laevifolia Lepidozia chordulifera Tayl. Figure 28. Lepidozia chordulifera Tayl., London J. Bot. 5: 371. 1846. Jungermannia chordulifera (Tayl.) Hook, f. & Tayl. in Hook, f., Bot. An tare. Voyage 1(2): 442. 1847. Mastigophora chordulifera (Tayl.) Trev., Mem. Reale 1st. Lomb. Sci. Lett. Ill, 4: 397. 1877. Original material: Chile, Prov. Aisen & Chiloe, Arch, de los Chonos, 1 834, Darwin 461 (FH— cited in Fulford, 1966, NY!). Lepidozia hastata Steph., Spec. Hep. 3: 605. 1909, syn.fide Fulford (1966). Original material: Chile, Prov. Aisen, R. Aisen, Dusen 83a (G), cited in Fulford (1 966) (non vidi). Lepidozia cuspidata Steph., Kongl. Svenska Veten- skapsakad. Handl. 46(9): 61. f. 23a-b. 191 1, syn. nov. Original material: Western Patagonia, Skottsberg s.n. (G), cited in Fulford (1966) (non vidi). Lepidozia ejffusa Steph., Kongl. Svenska Vetenskap- sakad. Handl. 46(9): 62. / 23g-h. 1911, syn.fide Fulford (1966). Original material: Chile, Prov. Magallanes, Pto. Ramirez and I. Atalaya, Halle & Skottsberg (G), cited in Fulford (1966) (non vidi). Lepidozia fernandeziensis Steph., Kongl. Svenska Ve- tenskapsakad. Handl. 46(9): 63. / 24e. 191 1,12 syn.fide Fulford (1966). Original material: Juan Fernandez Is., Mas a Tierra, near El Yunque, Skottsberg s.n. (G), cited in Fulford (1966) (non vidi); Prov. Chiloe, I. Chiloe, Fundo San Antonio, R. Pudito, Skottsberg (non vidi); Prov. Magal- lanes, R. Azopardo, Skottsberg (non vidi). Lepidozia hariotii Steph., Spec. Hep. 6: 329. 1922, syn.fide Fulford (1966). Original material: Chile, Prov. Magallanes, I. Hermite, Hariot (G, ex hb. Bescherelle), cited in Fulford (1966) (non vidi). Lepidozia microscopica Steph., Spec. Hep. 6: 334. 1 922, syn.fide Fulford (1966). Original material: Chile, Prov. Aisen, Cta. Hale, Dusen s.n. (G), cited in Fulford (1966) (won vidi). Lepidozia angulata Steph. ex Fulf., Mem. New York Bot. Gard. 1 1: 206. 1966, nom. nud., pro syn. 12 Fulford (1966) stated the description and figures of L. fernandeziensis have been interchanged with those of L. disticha in Stephani (1911). REMARKS— See Engel (1978). ECOLOGY— Very common in the Falklands, where it is better able to tolerate quite xeric, ex- posed conditions than any other Falkland hepatic, often occurring in crevices and under overhangs of rock outcrops in dwarf shrub heaths. In this association it is also frequently encountered under Blechnum magellanicum cover as well as on Bolax cushions. Rare in sheltered high altitude cliffs and moist feldmarks. PHYTOGEOGRAPHY— Valdivian + Magellanian + Falklands— South Sandwich Is.; South Georgia; Falkland Is.; Tierra del Fuego; Patagonian Chan- nels; Valdivian region; Juan Fernandez Is. (400- 795 m on Mas a Tierra). The report from Tas- mania in Rodway (1916) requires confirmation (see fig. 28). LITERATURE RECORDS (FALKLANDS)— A nony- mous (Kuhnemann, 1937, 1 949); Skottsberg, Port Stanley (Fulford, 1966; Skottsberg, 1913), Port Harriet (Fulford, 1966); Skottsberg & Halle, Port Harriet (Stephani, 1911). FALKLAND SPECIMENS SEEN -EAST FALKLANDS. 1843, Hooker s.n. as Jungermannia laevifolia var. in- crassata (FH). Port William Region: summit ridge of Mt. Low, ca. 245 m (3230, 3238); N side of Gypsy Cove, sea level (3242-3243, 3252, 3258); Engineer Point head- land, The Narrows, ca. 15m, Imshaug 40618 (MSC); ridge on Engineer Point Peninsula, 18-25 m (2817). Stanley Region: Rookery Bay, ca. 15m, Smith 3026 (AAS, F); Port Stanley, 8 Apr. 1902, Skottsberg s.n. as Lepidozia saddlensis (s); Port Stanley, 1953, Pettingill 32 (CMC, F); near Sapper Hill, 29 Oct. 1907, Skottsberg as L. pallida (s, UPS); Sapper Hill, 135 m (2.792. 2394. 239? 'A, 2399. 2407 -c. per., 2410, 2413-c. sporo.-2414, 2419); near summit of Tumbledown Mt., 215m, Long- ton & Smith, Longton 977 (AAS, F); Tumbledown Mt., 155-230 m (237 '5 A. 2380, 2384, 2391); Goat Ridge, ca. 180 m (3118. 3205, 3208, 3216); summit ridge of N peak of Two Sisters, 245-290 m (2703 A. 2727, 27525, 2735-2736); N peak of Two Sisters, 250 m, Dollman s.n. (Greene 3560a) (AAS); summit of Mt. Kent, 455 m (2766). Mt. Usborne Region: gap between Mt. Usborne 2 and Table Rock, ca. 440 m (2640B); summit of Mt. Usborne 1, ca. 700 m (2495). WEST FALKLANDS. ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 73 FIG. 28. Distribution of Lepidozia chordulifera Tayl., plus South Georgia. Open circle indicates precise locality unknown. Port Howard: Freezer Rocks, E slope of Mt. Maria, 320 m (3141. 3147). Fox Bay Region: summit of East Head, !80 m (3421. 3439); valley E of Sulivan House, ca. 75 m (3531). Mt. Adam: E side of summit ridge, 670-700 m (3017). Hill Cove Region: summit of East French Peak, 305 m (2977); summit of West French Peak, 290 m (296 2-296 3); summit of Mt. Fegen, 335-360 m (3065. 3073. 3076); ridge of N slope of Mt. Fegen, ca. 275 m (3057. 3062). Lepidozia fuegiensis Steph. Lepidozia fuegiensis Steph., Kongl. Svenska Veten- skapsakad. Handl. 46(9): 63. / 24f-g. 191 1. Orig- inal material: Chile, Prov. Magallanes, Pto. Cut- ter, 13 April 1908, Halle & Skottsberg (o!, sub "Patagonia austra.," s!, UPS!); Prov. Magallanes, Cta. Gome/, 26 Feb. 1908, Halle & Skottsberg (s!, UPS!). Lepidozia minuta Steph., Spec. Hep. 3: 603. 1909, non Lepidozia minuta Col., Trans. & Proc. New Zealand Inst. 18: 245. 1886, syn. fide Fulford (1966). Original material: Chile, Prov. Magal- lanes, R. Azopardo, Dusen 93 (o), cited in Fulford (1966) (non vidi). Lepidozia magellanica Steph., Kongl. Svenska Veten- skapsakad. Handl. 46(9): 64./ 24m-n. 191 1, syn. fide Fulford (1966). Original material: Chile, Prov. Magallanes, I. Felix, Skottsberg 594 (s), cited in Fulford (1 966) (non vidi). Lepidozia parva Steph., Kongl. Svenska Vetenskap- sakad. Handl. 46(9): 65. / 24o-p. 1911, syn. fide Fulford (1966). Original material: Chile, Prov. Aisen, Cta. Hale, and Cta. Connor, Hale & Skottsberg (o), cited in Fulford (1966) (non vidi). REMARKS— See Engel (1978) for a comparison of Brunswick Peninsula and Falkland Islands pop- ulations of this species, and for enumeration of features which will identify the taxon. 74 FIELDIANA: BOTANY Stephani (1911) based Lepidozia fuegiensis on specimens from four localities— Pto. Cutter, Cta. Gomez, R. Fontaine, and R. Azopardo. Collec- tions at S and UPS from the first two localities rather closely match the protologue of L. fuegien- sis. The R. Fontaine plants, however, are L. lae- vifolia (1 March 1909, Halle & Scottsberg, UPS!), and I have not been able to locate the R. Azopardo collection. The specimens from G are labeled either "Patagonia austr." or "Fuegia." Since Pto. Cutter is the sole locality in Patagonia (the remaining three being in Tierra del Fuego), I am assuming those labeled "Patagonia austr." are indeed from Pto. Cutter. The single G collection from Fuegia is L. laevifolia and is probably the R. Fontaine plant. ECOLOGY— Found above 245 m in the Falk- lands, where it is of frequent occurrence in crevices of sheltered high altitude cliffs. Also known from feldmark rocks, particularly those in shallow pools, but only rarely in Cortaderia and dwarf shrub heaths. It is of interest to note that L. chordulifera and L. laevifolia are very common below 245 m and only rarely (a single collection of L. chordu- lifera) encountered in sheltered high altitude cliffs. PHYTOGEOGRAPHY— Valdivian + Magellanian + Falklands— South Georgia; Falkland Is.; Tierra del Fuego; southern Patagonian Channels; Bruns- wick Peninsula; Valdivian portion of Patagonian Channels. The "Frai Jorge" report in Herzog (1954) requires confirmation. FALKLAND SPECIMENS SEEN— EAST FALKLANDS. Stanley Region: summit ridge of N peak of Two Sisters, 245-290 m (2724). Mt. Usborne Region: gap between Mt. Usbome 2 and Table Rock, ca. 440 m (2638); SE slope of Mt. Usborne 2, ca. 455 m (2604, 2616— c. per.); ridge between Mts. Usborne 1 & 2, 685 m (2534 '-A, 2539B); summit of Mt. Usborne 1 , ca. 700 m (2491B-2492, 2496); The Gap, 275-290 m (2466). WEST FALKLANDS. Port Howard: pass SW of Mt. Maria summit, ca. 610 m (3083A, 3096, 3107, 3109, 3126B). Mt. Adam: ridge S of northern lake, 610 m (3033, 3035). Lepidozia laevifolia (Hook. f. & Tayl.) G. L. & N. Jungermannia laevifolia Hook. f. & Tayl., London J. Bot. 3: 385. 1844 (3: 285. [sic.] in errorepro 385). Lepidozia laevifolia (Hook. f. & Tayl.) G. L. & N., Syn. Hep. 208. 1845. Mastigophora laevifolia (Hook. f. & Tayl.) Trev., Mem. Reale 1st. Lomb. Sci. Lett. Ill, 4: 416. 1877. Original material: Campbell Is., Hooker (non vidi). Lepidozia Jacquinotii" Steph., Spec. Hep. 3: 604. 1 909, syn. fide. Fulford (1966), non Lepidozia Jacquinotii" Steph., Spec. Hep. 6: 330. 1922. Original material: Chile, Prov. Magallanes, Pto. Gallant, Jacquinot (G) (non vidi), cited in Fulford (1966); Patagonia (without specific location), Du- sen (non vidi); Prov. Chiloe, I. Chiloe, sin. coll. (non vidi). Lepidozia viridissimaSleph., Spec. Hep. 3: 604. 1909, syn. fide Fulford (1966). Original material: Chile, Prov. Magallanes, Strait of Magellan, Dusen (a), cited in Fulford (1966) (non vidi). Lepidozia falklandica Steph., Kongl. Svenska Veten- skapsakad. Handl. 46(9): 63. / 24c-d. 191 1, syn. fide Fulford (1966). Original material: Chile, Prov. Aisen, L. O'Higgins, Hailed Skottsberg; Falkland Is., Mt. Adam, and Hearnden Water, Halle & Skottsberg (G), cited in Fulford (1966) (non vidi). Lepidozia halleana Steph., Kongl. Svenska Veten- skapsakad. Handl. 46(9): 64. / 24k-l. 191 1, syn. nov. Original material: Falkland Is., near Port Stanley, Hearnden Water, 1 Nov. 1907, Halle (G!, s!, UPS!). ECOLOGY— Rather common in dwarf shrub heaths, where it is found on soil under Empetrum, or more rarely on stream banks. It also may be found in rock crevices of outcrops in this associ- ation. PHYTOGEOGRAPHY — Amphipacific Temper- ate—reported from South Georgia; Falkland Is.; the Magellanian and Valdivian regions; Juan Fer- nandez Is.; Macquarie Is.; Auckland and Campbell Is.; New Zealand; Tasmania; Kerguelen Is.; Mar- ion and Prince Edward Is. LITERATURE RECORDS (FALKLANDS)— A nony- mous (Kiihnemann, 1937, 1949 as L. halleana, L. jacquemontii [sic], and L. falklandica); Halle (Ful- ford, 1966, Stephani, 1922 as L. halleana, and Taylor, 1960 as L. halleana); Hooker (Taylor & Hooker, 1847 as Jungermannia); Skottsberg (Ful- ford, 1966), Port Stanley (Skottsberg, 1913 as L. jacquemontii [sic]); Port Stanley and Mt. Adam (Skottsberg, \9l3asL.falklandica, Stephani, 1922 as L. falklandica); Skottsberg & Halle (Fulford, 1966); Port Stanley (Stephani, 1911 as L. jacque- montii [sic]). FALKLAND SPECIMENS SEEN— EAST FALKLANDS. Port William Region: N side of Gypsy Cove, sea level (3259); ridge on Engineer Point Peninsula, 20-25 m (2834). Stanley Region: Sapper Hill, 1 35 m (2398, 24 17 A, 2426); Goat Ridge, ca. 180 m (3199); Sapper Hill, 2 km W of Port Stanley, ca. 50 m, Ochyra 2830/80 (F). Darwin Settlement: Boca House on Brenton Lock, sea level (2696). 13 Stephani, in describing his new species, used the spelling "Jacquemontii," which was the spelling used by subsequent authors. Since the species was named after Honore Jacquinot, I have altered the spelling of the ep- ithet accordingly (see Engel, 1978, p. 99). 14 1 believe there is a distinct possibility that Lepidozia Jacquemontii of Stephani, 1922, leg. Skottsberg at "Fre- tum magellanicum," is based upon the same type as L. jacquemontii of Stephani, 1909. See Engel (1978, p. 99) for discussion. ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 75 Port Louis: 2 Aug. 1902, Skottsberg s.n. as L. saddlensis (s). WEST FALKLANDS. Fox Bay Region: near peat cuttings in Ram Paddocks, 45-60 m (3509); valley E of Sulivan House, ca. 75 m (3536). Notes on Lepidozia Species Until the type of Lepidozia pallida Steph. is lo- cated, the identity of this species must remain open to question. As I have pointed out (Engel, 1978, p. 102), perhaps it is of some significance that all the reports (from five localities) of Lepidozia pal- lida in Stephani (1911) were found to be of Lep- idozia chordulifera. This includes the Falkland specimen from Port Stanley, Sapper Hill. Paracromastigum Paracromastigum Fulf. & J. Tayl., Brittonia 13: 336. 1961. Key to Falkland Islands Species of Paracromastigum * 1. Terminal branching common, of Frullania and Acromastigum types; cortical cells in ca. 12 rows . . . P. subsimplex 1 . Terminal branching absent (in Falkland specimens) or, if present, then rare and only of Frullania type; cortical cells in 7-8 rows P. tristanianum Paracromastigum subsimplex (Steph.) Fulf. & J. Tayl. Lepidozia subsimplex Steph., Kongl. Svensk Veten- skapsakad. Handl. 46(9): 66. / 24r-s. 1911. Par- acromastigum subsimplex (Steph.) Fulf. & J. Tayl., Brittonia 13: 336. 1961. Original material: (?) Chile, Prov. Magallanes, (?) Argentina, Terr. Tierra del Fuego, L. Fagnano, 1908, Halle s. n. (G), cited in Fulford & Taylor (1961) (non vidi). ECOLOGY— This rare species, represented by a single Falkland collection, was gathered on the bank of a wet depression in a Cortaderia heath, 580-595 m. PHYTOGEOGRAPHY— Valdivian + Magellanian + Falklands— Falkland Is.; Isla Grande de Tierra del Fuego; Valdivian region. FALKLAND SPECIMEN SEEN -WEST FALKLANDS. Mt. Adam: basin E of summit, 580-595 m (3002). Paracromastigum tristanianum (Schust.) Engel & Schust. Pseudocephalozia tristaniana Schust., Nova Hedwigia 10: 23. 1965. Paracromastigum tristanianum (Schust.) Engel & Schust. in Schuster & Engel, J. Hattori Bot. Lab. 38: 700. 1974. Holotype: Tris- tan da Cunha, Red Hill, Christopher sen 1229C (G, non vidi). REMARKS— The Falkland Islands plants fit the original description of the species, except for two features. Median leaf cells measure 21-36 ^m wide x 39-53 nm long in Falkland material and "ca. 20-25 x 25-38 Mm" in type material. The type has predominantly intercalary branching, with ter- minal, Frullania-type branches rare and Acro- mastigum-type branches lacking. Falkland mate- rial has abundant intercalary branching from all three merophytes, but no terminal branches were observed. Stolons are abundant, freely branched, and prostrate, while the leafy shoots are erect and issue from the stolon system. Androecia are sim- ilar to the type in being on leading shoots and lacking bracteolar antheridia. Leafy shoots on Falkland plants have branches originating only from basal sectors and never from the zone of mature leaves. If that condition occurs in the type, it is then of diagnostic value, and will differentiate the species from P. subsimplex. In that species, branching freely develops from mature-leaved sectors of shoots. ECOLOGY- PHYTOGEOGRAPHY— Apparently quite rare in the Falklands, where it was collected but once— on a stream bank in a Cortaderia heath. This amphiatlantic temperate species is restricted to the Falkland Is. and Tristan da Cunha. FALKLAND SPECIMEN SEEN -EAST FALKLANDS. Mt. Usborne Region: valley SW of Mt. Usborne, ca. 60 m (2670-c. S). 76 FIELDIANA: BOTANY FIG. 29. Distribution of Pseudocephalozia quadriloba (Steph.) Schust. ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 77 Pseudocephalozia Pseudocephalozia Schust., Nova Hedwigia 10: 2 1 . 1965. Pseudocephalozia quadriloba (Steph.) Schust. Fig- ure 29. Isotachis quadriloba Steph., Bih. Kongl. Svenska Ve- tenskapsakad. Handl. 26(111, 6): 54. 1900. Hy- grolembidium quadrilobum (Steph.) Schust., Nova Hedwigia 10: 23. 1965. Lembidium quadrilobum (Steph.) Fulf., Mem. New York Hot. Gard. 11: 247. 1966. Pseudocephalozia quadriloba (Steph.) Schust., J. Hattori Hot. Lab. 36: 37 1 . 1 973 [ 1972]. Original material: Chile, Prov. Chiloe, I. Guai- tecas, Dusen 182 (o 1 1 102! as Fuegia, without specific locality). Cephalozia quadriloba Steph., Kongl. Svenska Veten- skapsakad. Handl. 46(9): 58. / 22a-b. 1911 = Lophozia crassicaulis Steph., Spec. Hep. 6: 111. 1917, syn. fide Schuster & Engel ( 1 974). Original material: Chile, Prov. Magallanes, W. end of L. Fagnano, Skottsberg 555 (G! as Fuegia, without specific locality). Isotachis granditexta Steph., Kongl. Svenska Veten- skapsakad. Handl. 46(9): 68. / 26c, e. 1911, syn. fide Fulford (1966). Original material: Chile, Prov. Magallanes, L. Fagnano, R. Azopardo region, 10- 1 1 March 1908, Halle 127 (UPS!). REMARKS— See Schuster and Engel (1974). ECOLOGY— See Schuster and Engel (1974, pp. 693-694 and fig. 17). PHYTOGEOGRAPHY— Andean American — Inac- cessible I. (475 m); Falkland Is. (10-300 m); Isla Grande de Tierra del Fuego (85-650 m); southern Patagonian Channels (Brunswick Peninsula and Hotel Rubens-Puerto Natales region); Valdivian region (at 43°57'S, Islas Guaitecas); Colombia (3200-3600 m); Venezuela (3850 m); Costa Rica (2600 m) (see fig. 29). FALKLAND SPECIMENS SEEN— EAST FALKLANDS. Port William Region: summit ridge of Mt. Low, ca. 245 m (3224, 3226B). Stanley Region: headwaters of Mullet Creek Stream, ca. 60 m (3171); summit ridge of N peak of Two Sisters, 245-290 m (2702, 2776). Mt. Usborne Region: The Gap, 275-290 m (2459, 2473). WEST FALKLANDS. Fox Bay Region: near mouth of Cheek's Creek, ca. 12 m (3478). Telaranea Telaranea Spruce ex Schiffn. in Engler & Prantl, DieNaturl. Pflanzenfam. 1(3, 1): 103. 1893. Telaranea Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 365. 1885, nom. inval. in syn. Neolepidozia Fulf. & J. Tayl., Brittonia 11: 81. 1959. Key to Falkland Islands Species of Telaranea 1. Leaf apices 2-3 dentate-lobate, and if lobate, then lobes less than 0.5 the leaf length; leaf disc more than 4 rows of cells high. Leaf segments 2 or 3(4) cells long and 2 cells wide at the base; leaf cells in 8 longitudinal rows Subg. Neolepidozia . . . T. oligophylla 1. Leaf apices 4-6 lobed for 0.5 or more the leaf length; leaf disc 1-4 rows of cells high Subg. Telaranea ... 2 2. Leaves of main axis with lamina of 1 row of cells high, the leaves 2-3 segmented; underleaves conspicuously smaller than the leaves; plants small, filamentous T. pseudozoopsis 2. Leaves of main axis with lamina of 1 .5 to 4 rows of cells high, the leaves 4-6 segmented; underleaves similar to leaves in size; plants large, leafy 3 3. Leaves and underleaves stiff, ascending, bristle-like, the lamina 1.5 cell rows high T. blepharostoma 3. Leaves and underleaves lax, curved, not ascending and bristle-like, the lamina 2-4 cell rows high . . . 4 4. Leaf margins 3-4 cell rows high; shoot apices often recurved; perianth mouth crenulate T. plumulosa 4. Leaf margins 1-2 cell rows high; shoot apices plane, not recurved; perianth mouth long-ciliate . . T. tetradactyla 78 FIELDIANA: BOTANY Telaranea blepharostoma (Steph.) Fulf. Figure 30. Lepidozia Blepharostoma Steph., Bih. Kongl. Svenska Vetenskapsakad. Handl. 26(111, 17): 22. 1901. Telaranea blepharostoma (Steph.) Herz., Rev. Bryol. Lichenol. 20: 189. 1960, nom. illeg. Te- laranea blepharostoma (Steph.) Fulf., Brittonia 15: 73. 1963. Original material: Chile, Prov. Ma- ua llanos. Pto. Angosto, "ex Bot. Mus. Upsal." Dusen 142 (o), cited in Fulford ( 1 963a) (non vidi). Lepidozia trichophylla Angstr. ex Fulf., Mem. New York Bot. Card. 11: 240. 1966, nom. nud., pro syn. REMARKS— As all previous reports of T. ble- pharostoma from the Falklands are based upon misdeterminations of T. plumulosa, this is the first authentic report of Telaranea blepharostoma in these islands. ECOLOG Y-PH YTOGEOGR APH Y — Va Id i via n + Magellanian-Falkland— Falkland Is.; Tierra del Fuego; Patagonian Channels; Valdivian region (see fig. 30). Within the Falklands, it was found to occur only in the Port Stanley region under Blechnum in a dwarf shrub heath, and on moist, sandy, shel- tered, vertical, north-facing rocks along the south shore of the Murrell River. The cooler tempera- tures and higher rainfall in this region may account for its presence there. LITERATURE RECORDS (FALKLANDS)— A nony- mous (Kiihnemann, 1937, 1949 as Lepidozia); Skottsberg—Port Philomel, Port Stanley (Skotts- berg, 1913 as Lepidozia); Skottsberg & Halle— Port Philomel, Port Stanley (Stephani, 1911 as Lepidozia). FALKLAND SPECIMENS SEEN— EAST FALKLANDS. Stanley Region: S shore of Murrell River, opposite Islet Point, 30 m, Longton & Smith, Longton 1027 (AAS, F); Goat Ridge, ca. 180 m (3201). Telaranea oligophylla (Lehm. & Lindenb.) Engel. Figure 31. Jungermannia oligophylla Lehm. & Lindenb. in Leh- mann, Nov. Minus Cogn. Stir. Pug. 6: 26. 1 834. Lepidozia oligophylla (Lehm. & Lindenb.) G. L. & N., Syn. Hep. 201. 1845. Mastigophora oli- gophylla (Lehm. & Lindenb.) Trev., Mem. Reale 1st. Lomb. Sci. Lett. Ill, 4: 415. 1877. Neolepi- dozia oligophylla (Lehm. & Lindenb.) Fulf. & J. Tayl., Brittonia 11: 84. 1959. Telaranea oligo- phylla (Lehm & Lindenb.) Engel, Bryologist 79: 514. 1976. Original material: Argentina, Terr. Tierra del Fuego, I. de los Estados, Menzies s.n. (NY!). REMARKS— See Engel (1978). 30 Fio. 30. Distribution of Telaranea blepharostoma (Steph.) Fulf. FIG. 3 1 . Distribution of Telaranea oligophylla (Lehm. & Lindenb.) Engel. Open circle indicates precise locality unknown. ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 79 ECOLOGY— Very rare in the Falklands, where it was gathered in a wet depression of a Cortaderia heath (455 m). It occurred mixed with other hepat- ics, nearly all of which are rare or relatively so, e.g., Clasmatocolea obvoluta var. cookiana, Hy- alolepidozia bicuspidata, Leptoscyphus aequatus, Kunia setiformis, and Riccardia spectabilis. PH YTOGEOGRAPH Y — Magellanian-Fa 1 k 1 a 1 1 c I — Falkland Is.; Tierra del Fuego; Patagonian Chan- nels north to 49°09'S (see fig. 31). FALKLAND SPECIMEN SEEN— EAST FALKLANDS. Mt. Usborne Region: SE slope of Mt. Usborne 2, ca. 455 m (26 12 A). Telaranea plumulosa (Lehm. & Lindenb.) Fulf. Figure 32. Jungermannia plumulosa Lehm. & Lindenb. in Leh- mann, Nov. Minus Cogn. Stir. Pug. 6: 26. 1834. Lepidozia plumulosa (Lehm. & Lindenb.) G. L. & N., Syn. Hep. 211. 1845. Mastigophora plu- mulosa (Lehm. & Lindenb.) Trev., Mem. Reale 1st. Lomb. Sci. Lett. Ill, 4: 416. 1877. Telaranea plumulosa (Lehm. & Lindenb.) Herz., Rev. Bryol. Lichenol. 29: 189. 1960, nom. illeg. Telaranea plumulosa (Lehm. & Lindenb.) Fulf., Brittonia 15: 77. 1963. Original material: Argentina, Terr. Tierra del Fuego, I. de los Estados, Menzies s.n. (G) "ex hb. Kew," cited in Fulford (1963a) (non vidi). Lepidozia magellanica Gott. ex Fulf, Mem. New York Bot. Gard. 11: 243. 1966, nom. nud., pro syn., non Lepidozia magellanica Steph., Kongl. Sven- ska Vetenskapsakad. Handl. 46(9): 64. 1911 (= L. fuegiensis). REMARKS— This taxon reaches its optimal de- velopment in the evergreen forested region of southern Chile. The plants of the Falkland pop- ulation, when compared with the Chilean plants, are smaller in size, less luxuriant and less robust, often without recurved plant tips, and with few exceptions have leaves which are consistently 4-segmented. Fulford ( 1 963a, 1 966) stated that the leaves of T. plumulosa are 5- or 6-segmented. The leaf lamina and margins of the Falkland plants are 3-4 cells high, data which agrees with the Fulford descriptions. The Falkland Islands plants, which show little variation, are thus somewhat inter- mediate between T. plumulosa and T. tetradac- tyla; however, I believe the concept of the former should be expanded to include plants with 4-seg- mented leaves. ECOLOGY— Very common, characteristic of stream banks, particularly under Gunnera, Blech- num magellanicum, or Empetrum cover and in crevices of outcrops in dwarf shrub heaths. Also common on soil, frequently of stream banks in Cortaderia heaths. It occasionally grows in rock crevices or under overhangs of sheltered high al- titude cliffs. PHYTOGEOGRAPHY— Valdivian + Magellanian + Falkland -Falkland Is.; Tierra del Fuego; Pa- tagonian Channels; Valdivian region (West Pata- gonia only at 43°57'S, on an offshore island); and Mas a Tierra, Juan Fernandez Is. (400-500 m) (fig. 32). LITERATURE RECORDS (FALKLANDS)— A nony- mous (Fulford, 1963a, 1966; Kuhnemann, 1937 as Lepidozia); Lyall (Fulford, 1963a, 1966); Skottsberg, North West Bay (Fulford, 1 963a, 1 966), Port Stanley (Fulford, 1966). FALKLAND SPECIMENS SEEN— EAST FALKLANDS. Stanley Region: near Port Stanley, 1 1 Nov. 1907, Skotts- berg 165 as Lepidozia blepharostoma (s, UPS), headwaters of Mullet Creek Stream, ca. 60 m (3162); between Mt. William and Tumbledown Mt., 1 70 m, Longton & Smith, Longton 979 (AAS, F); summit ridge of N peak of Two Sisters, 245-290 m (2700/1, 2779, 2738); summit of Mt. Kent, 455 m (2746). Mt. Usborne Region: SE slope of Mt. Usborne 2, ca. 455 m (2596); The Gap, 275-290 m (2467, 2477); below The Gap, ca. 90 m (2564, 2567, 2577); S side of The Gap, 260-275 m (2452A); valley SW of Mt. Usborne, ca. 60 m (2655, 2664). WEST FALKLANDS. Port Howard: SW of Mt. Maria summit, ca. 610 m (3131). Fox Bay Region: near peat cuttings in Ram Paddock, 45-60 m (3514— c. 9); N base of Fox Bay Mt., 75 m (5459, 5460, 3464A). Mt. Adam: ridge S of northern lake, 610m (5025, 303 IB). Hill Cove Region: gap between French Peaks, 200-2 1 5 m (2978); summit of West French Peak, 290 m (2949); Hill Cove, Vallentin Sp. 55 (MANCH). Port Philomel: Northwest Bay, 2 Nov. 1907, Halle s. n. as Lepidozia blepharostoma (UPS). West- point Island: near The Waterfall, 30-90 m (2574, 2575). Weddell Island: Waterfall Valley, W of settlement, ca. 125 m (3305 A, 3312, 3321, 3327B, 3328); near head- waters of House Creek, ca. 15 m (5404, 5406, 5407, 5470); summit of peak NE of Mt. Weddell, 335 m (5565); summit of Mt. Weddell, 380 m (5556). Telaranea pseudozoopsis (Herz.) Fulf. Figure 33. Lepidozia pseudozoopsis Herz. in Skottsberg, Nat. Hist. Juan Fernandez Is., Easter I. 2(5): 723./ 5. 1942. Telaranea pseudozoopsis (Herz.) Fulf., Brittonia 15: 71. 1963. Original material: Juan Fernandez Is., Mas a Tierra, Centinela, 530 m, Skottsberg 227 (jE)-cited in Fulford (1966) (non vidi). ECOLOGY— Rather common in the Falklands, typically in moist, shaded situations (moist rock crevices, stream banks, etc.) in dwarf shrub heaths, Hebe associations and (more rarely) in Cortaderia heaths. It is rather common on the bases of tussock grasses in tussock associations. PHYTOGEOGRAPHY— Valdivian + Magellanian + Falkland — Falkland Is.; Tierra del Fuego; 80 FIELDIANA: BOTANY southern Patagonian Channels, Brunswick Pen- insula; Valdivian region; Mas a Tierra, Juan Fer- nandez Is. (400-530 m, where recorded) (fig. 33). LITERATURE RECORD (FALKLANDS): Smith, Beauchene Is. (Smith & Prince, 1985). FALKLAND SPECIMENS SEEN— EAST FALKLANDS. Kidney Island: N of shanty (2756, 2793, 2794, 2800). Stanley Region: Sapper Hill, 135 m (2395); Tumble- down Mt., 155-230 m (2377 B); headwaters of Mullet Creek Stream, ca. 60 m (3165, 3181C); Goat Ridge, ca. 180 m (3212, 3217). Mt. Usborne Region: The Gap, 275-290 m (2461-c. per. + 3); valley SW of Mt. Us- borne, ca. 60 m (2648Q. WEST FALKLANDS. Fox Bay Region: summit of East Head, 1 80 m (3423); sum- mit of Fox Bay Mt., 307 m (3448, 3454). Hill Cove Region: summit of West French Peak, 290 m (2951). Westpoint Island: steep slope and cliffs facing The Wool- ly Gut (29/9, 2922. 2927, 2925, 2939); near The Wa- terfall, 30-90 m (2875A, 2876); W base of Black Hog Hill, adjacent to Devils Nose, ca. 20 m (2597, 2893). Weddell Island: Waterfall Valley, W of settlement, ca. 125m (3307). Telaranea tetradactyla (Hook. f. & Tayl.) Hodgs. Jungermannia tetradactyla Hook. f. & Tayl., London J. Bot. 3: 386. 1844 (3: 286 [sic] in errore pro 386). Lepidozia tetradactyla (Hook. f. & Tayl.) G. L. & N., Syn. Hep. 213. 1845. Mastigophora tetradactyla (Hook. f. & Tayl.) Trev., Mem. Reale 1st. Lomb. Sci. Lett. Ill, 4: 397. 1877. Neolepi- dozia tetradactyla (Hook. f. & Tayl.) Fulf. & J. Tayl., Brittonia 11: 84. 1959. Telaranea tetra- dactyla (Hook. f. & Tayl.) Hodgs., Rec. Domin. Mus. 4: 106. 1962. Original material: Auckland Is., Hookers. n. (NY), cited in Fulford ( 1 963a) (non vidi). Lepidozia lindenbergii Gott. in G. L. & N., Syn. Hep. 213. 1845. syn. fide Fulford (1963a). Mastigo- phora lindenbergii (Gott.) Trev., Mem. Reale 1st. Lomb. Sci. Lett. Ill, 4: 397. 1877. Original ma- terial: New A\i hi ml. without specific locality, "ex hb. Taylor," Hooker s.n. (G, NY), cited in Fulford (1963a)(m>w vidi). Lepidozia disticha Steph., Kongl. Svenska Vetenskap- sakad. Handl. 46(9): 62. / 24a-b. 1911, syn. fide Fulford (1963a). Neolepidozia disticha (Steph.) Fulf. & J. Tayl., Brittonia 1 1: 84. 1959. Original material: Juan Fernandez Is., Mas a Tierra, Skottsberg 117 (G), cited in Fulford ( 1 963a) (non vidi). Lepidozia effusiseta Steph., Spec. Hep. 6: 325. 1922, syn. fide Fulford ( 1 963a). Original material: Juan Fernandez Is.; Chile, Prov. Magallanes, I. Des- olation, Skottsberg (G), cited in Fulford (1963a) (non vidi). REMARKS— Fulford (1963a, p. 277; 1966, pp. 206, 241) stated that the original description and figures of Lepidozia fernandeziensis and L. disticha (syn. of Telaranea tetradactyla) were interchanged . 32 FIG. 32. Distribution of Telaranea plumulosa (Lehm. & Lindenb.) Fulf. FIG. 33. Distribution of Telaranea pseudozoopsis (Herz.) Fulf. ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 81 in Stephani (1911). Herzog (1954) was apparently not aware of Stephani's error when citing (p. 51) "Lepidozia (Telaranea) fernandeziensis St. (= L. effusiseta St.)-" Fulford (1966) has cited the Her- zog error as a misidentification ("Lepidozia [Te- laranea] fernandeziensis Herzog, Rev. Bryol. Li- chenol. 23: 51. 1954 (in error)") in the synonymy of Lepidozia chordulifera. PH YTOGEOGRAPHY— Amphipacific Temper- ate—(?) Falkland Is.; Tierra del Fuego; Patagonian Channels; Valdivian Region (West Patagonia north to 39°48'S); Juan Fernandez Is.; Auckland Is.; and New Zealand. The species is common throughout New Zealand and neighboring islands (Hodgson, 1956). LITERATURE RECORDS (FALKLANDS)— Skotts- berg, Port Stanley (Fulford, 1966). Lepidoziaceae Species Excluded from Falklands Bazzania peruviana (Nees) Trev. This species was reported from the Falklands by Stephani (Spec. Hep. 6: 480. 1924, as Masti- gobryum skottsbergii Steph.) by Kiihnemann ( 1 949) and by Bonner (1963). I have failed after several attempts to locate this specimen. However, a label of a specimen from Geneva of a 1 908 collection, Skottsberg 50, from Mas a Tierra, Juan Fernandez Islands, originally bore the locality "Falkland Ins.," which had been crossed out. It is likely that Ste- phani discovered his error when curating the Skottsberg collections, but did not delete the rec- ord from his Species Hepaticarum. The species occurs in the Magellanian and Valdivian regions, Juan Fernandez Islands, and Tristan da Cunha. It appears not to grow in the Falklands. Family CALYPOGEIACEAE Calypogeiaceae (K. Mull.) H. Arnell in Holmberg, Skand. Fl. 2(a), 189. 1928. A cosmopolitan family with two genera, Caly- pogeia, with ca. 90 species, and Metacalypogeia with four species. One species of Calypogeia, C. sphagnicola (H. Arnell & J. Perss.) Warnst. & Loeske is bipolar in distribution; in the Southern Hemisphere it is known from southern South America, New Zealand, and Tasmania (Schuster, 1969b; Engel, 1978). It has not been recorded for the Falklands. Family ADELANTHACEAE Adelanthaceae (J0rg.) Grolle, J. Hattori Bot. Lab. 35: 327. 1972. Tribe Adelantheae J0rg., Ber- gens Mus. Skr. 16: 257. 1934. The Adelanthaceae have two genera. Adelan- thus, with 1 2 species, is well represented in south temperate and Neotropical regions. There are two species which reach Europe, and one is known only from Borneo. Wettsteinia has four species, and is known from southern South America, Juan Fer- nandez, the Falkland Islands, New Zealand, and the Indomalayan region. Adelanthus Adelanthus Mitt., J. Proc. Linn. Soc., Bot. 7: 243. 1864, nom. cons. Key to Falkland Islands Species of Adelanthus 1 . Dorsal margin of leaf only weakly incurved; leaves without distinct differentiation of a vitta of elongated cells; subapical cells of leaf with walls thin to slightly thickened. Leaf apex occasionally retuse to short bifid; leaf base never auriculate A. tenuis 1 . Dorsal margin of leaf distinctly incurved; leaves on well-developed plants with ± distinctly developed vittae of elongated cells; subapical cells of leaf with walls thick. Leaf apex rounded to angular, not retuse or short bifid; leaves often ± auriculate at the ventral base 2 2. Leaf margins dentate A. lindenbergianus 2. Leaf margins entire A. integerrimus 82 FIELDIANA: BOTANY Adelanthus integerrimus Grolle Adelanthus integerrimus Grolle, J. Hattori Bot. Lab. 35: 340. / 4. 1972. Holotype: Chile, Prov. Ma- gallanes, Punta Arenas, C. Mina Rica, 550 m, Santesson M 746 (s, non vidi). ECOLOGY— This species is sporadic in the Falk- lands, occurring in moist feldmarks, high altitude cliffs, and Cortaderia heaths; it was not found be- low 455 m. PHYTOGEOGRAPHY— Subantarctic in Distribu- tion— A. integerrimus is known from Crozet Is.; South Georgia; Falkland Is.; Tierra del Fuego; southern Patagonian Channels (Brunswick Pen- insula). LITERATURE RECORDS (FALKLANDS)— //a/fe, Mt. Adam (Grolle, 1972). FALKLAND SPECIMENS SEEN— EAST FALKLANDS. Mt. Usborne Region: SE slope of Mt. Usborne 2, ca. 455 m (26J 4); summit of Mt. Usborne 1, ca. 700 m (2496A, 2499-c. 8, 2501C, 2505B). WEST FALKLANDS. Port Howard: pass SW of Mt. Maria summit, ca. 6 1 0 m (3128, 3135). Syn. Hep. 653. 1847. Adelanthus sphalerus (Hook, f. & Tayl.) Steph., Bull. Herb. Boissier II, 8 (8): 599. 1908 (= Spec. Hep. 3: 383). Original ma- terial: Chile, Prov. Magallanes, I. Hermite, Hook- er (w), cited in Grolle (1972) (non vidi). Jungermannia ? haliotiphylla De Not., Mem. Reale Accad. Sci. Torino II, 16: 217. pi. V. 1-7. 1855, syn. cf. Schiffner(1890). Original material: Chile, "Prov. Valparaiso, Valparaiso," Puccio s.n. (L, MANCH, s), cited in Grolle (1972) (non vidi). Plagiochila subviminea Steph. in Herzog, Biblioth. Bot. 21: 212. / 146 d. 1916, syn. fide Grolle (1972). Original material: Bolivia, Herzog 285 3 'la (G), cit- ed in Grolle (1972) (non vidi). Adelanthus trollii Herz., Hedwigia 74: 9 1 ./ 6a~c. 1 934, syn. fide Grolle (1972). Original material: Bolivia, Challana, Mina Fabulosa, Troll 50/a (JE), cited in Grolle (1972) (non vidi). Adelanthus parvus Herz., Rev. Bryol. Lichen. 11: 18. 1938, syn. fide Grolle (1972). Original material: Costa Rica, Prov. San Jose, C. de Las Vueltas, 2700-3000 m, Standley 43723/c (JE), cited in Grolle (1972) (non vidi). Plagiochila subviminea f. paramicola Herz., Rev. Bryol. Lichen. 11: 12. 1 938, syn. fide Grolle (1 972). Lec- totype (cf. Grolle, 1972): Costa Rica, Prov. San Jose, C. de Las Vueltas, 2700-3000 m, Standley & Valeria 43862 (JE, non vidi). Adelanthus lindenbergianus (Lehm.) Mitt. Figure 34. Only American synonyms are included here: Jungermannia lindenbergiana Lehm., Linnaea 4: 367. 1829. Plagiochila lindenbergiana (Lehm.) G. L. & N., Syn. Hep. 59. 1844. Adelanthus lindenberg- ianus (Lehm.) Mitt., J. Proc. Linn. Soc., Bot. 7: 244. 1 864. Adelanthus magellanicus var. linden- bergianus (Lehm.) Schiffn., Nova Acta Acad. Caesar. Leop.-Carol. German. Nat. Cur. 60(2): 261. 1893. Lectotype (cf. Grolle, 1972): South Africa, near Cape Town, Ecklon (s, non vidi). Plagiochila magellanica Lindenb., Spec. Hep. 1(5): 163.1 843, syn. cf. Mitten (1859). Adelanthus ma- gellanicus (Lindenb.) Mitt., J. Proc. Linn. Soc. Bot. 7: 244. 1864. Calyptrocolea magellanica (Lindenb.) Schust., J. Hattori Bot. Lab. 26: 287. 1963, nom. illeg. Lectotype (fide Grolle, 1972): Chile, Prov. Magallanes, B. San Nicolas, Dumont d'Urville (PC, non vidi). Jungermannia unciformis Hook. f. & Tayl., London J. Bot. 3: 457. 1844, syn. cf. Mitten (1859). Pla- giochila unciformis (Hook. f. & Tayl.) G. L. & N., Syn. Hep. 653. 1847. Adelanthus unciformis (Hook. f. & Tayl.) Spruce, J. Bot. 14: 200. 1876. Adelocolea unciformis (Hook. f. & Tayl.) Evans, Bull. Torrey Bot. Club 25: 409. 1898. Original material: Chile, Prov. Magallanes, I. Hermite, Hooker (BM, MANCH, w), cited in Grolle (1972) (non vidi). Jungermannia sphalera Hook. f. & Tayl., London J. Bot. 3: 458. 1844, syn. cf. Mitten (1859). Plagi- ochila sphalera (Hook. f. & Tayl.) G. L. & N., REMARKS— I have been unable to locate the specimen on which the record of Adelanthus ma- gellanicus from the Falklands was based. How- ever, the type of Adelanthus magellanicus is con- specific with A. lindenbergianus, but according to Grolle (in ////.), nearly all other collections sub- sequently determined as A. magellanicus belong to other taxa. Presumably the Falkland report is included among them. Plagiochila coriacea Steph., nom. herb., belongs here; it was collected at Port Harriet by Skottsberg (Hassel de Menendez, 1 983a, p. 123). See Engel ( 1978). ECOLOGY— This common species has a wide ecological amplitude, growing in rock crevices or on a soil layer either over rock or under rock over- hangs of outcrops in the dwarf shrub heath asso- ciation as well as the sheltered high altitude cliffs. The taxon also grows on soil in the Cortaderia (commonly under grass cover) and dwarf shrub heaths. Further, it was gathered on soil at 700 m in or at the margins of feldmark pools. PHYTOGEOGRAPHY— Amphiatlantic Temper- ate—Mascarene Is.; Reunion; Madagascar; South Africa (1000-2000 m); Uganda (3100-3500 m); Congo (3100 m); Tristan da Cunha (200-800 m); Inaccessible I. (300 m); South Georgia; Falkland Is.; Tierra del Fuego; Patagonian Channels; Val- divian region; Juan Fernandez Is.; Andes north to Mexico; and western Ireland (450-700 m) (fig. 34). According to Grolle (1972a), the Australasian ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 83 34 FIG. 34. Distribution of Adelanthus lindenbergianus (Lehm.) Mitt. North to Venezuela and Costa Rica, plus Ireland (after Grolle, 1969). records of A. magellanicus are actually A. occlusus and that from lies de Kerguelen is Jamesoniella sp. Among the Falkland taxa, it is one of the two (with Chiloscyphus semiteres) strictly austral taxa that is disjunct in the British Isles (see fig. 55). LITERATURE RECORDS (FALKLANDS)— Anony- mous (Kuhnemann, 1937, 1949 as A. unciformis); Lechler (Mitten, 1864 as /I. magellanicus); Skotts- berg. Port Stanley, Port Louis (Stephani. 1905a as A. unciformis). Port Louis (Grolle, 1972a as A. lindenbergianus), Port Stanley (Skottsberg, 1913 as A unciformis); Skottsberg & Halle, Weddell Is., Port Harriet (Stephani, 1911 as A. unciformis). FALKLAND SPECIMENS SEEN -EAST FALKLANDS. Port William Region: N side of Gypsy Cove, sea level (3241). Stanley Region: near Sapper Hill, 29 Oct. 1907, Halle & Skottsberg 1 13 as Frullania boveana (s); S slopes of Mt. William, 185 m, Longton & Smith, Longton 983 (AAS, F); Tumbledown Mt., 155-230 m (2373, 2382A); Goat Ridge, ca. 180 m (3190); N peak of Two Sisters, 250-275 m, Dollman s.n. (Greene 356 la, 3563b) (AAS, F); summit ridge of N peak of Two Sisters, 245-290 m (2733— c. per., 2734); at summit of Mt. Kent, 455 m 84 FIELDIANA: BOTANY JJE FIG. 35. Adelanthus tenuis Engel & Grolle. Plant; note ventral-intercalary branching and leaf apex variability, x 10 (from holotype). ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 85 (2763). Mt. Usborne Region: gap between Ml. Usborne 2 and Table Rock, ca. 440 m (2625C. 2626B. 2628. 2632, 2634A, 2640D); SE slope of Mt. Usborne 2, ca. 455 m (2600); ridge between Mts. Usborne 1 & 2, 685 m (2532-c. per., 2554A); The Gap, 275-290 m (2462B, 2463. 2469. 2475); below The Gap, ca. 90 m (2562A). WEST FALKLANDS. Port Howard: Freezer Rocks, on E slope of Mt. Maria, 320 m (3146. 3153); pass SW of Mt. Maria summit, ca. 610 m (3080A, 3133). Fox Bay Region: summit of East Head, 180 m (3417). Mt. Adam: E side of summit ridge, 670-700 m (3019A); E slope of main peak, ca. 6 1 0 m ( 3003 A). Hill Cove Region: summit of East French Peak, 305 m (2964 A— c. per.); summit of Mt. Fegen, 335-360 m (3079). Weddell Island: sum- mit of peak NE of Mt. Weddell, 335 m (3369B); summit of Mt. Weddell, 380 m (3335B, 3345. 3350, 3353). Ad elan thus tenuis Engel & Grolle. Figures 35-36. Adelanthus tenuis Engel & Grolle in Grolle, J. Hattori Bot. Lab. 35: 333. / 1-2. 1972. Holotype: Chile, Prov. Magallanes, Brunswick Peninsula, La. El Parrillar region, ca. 365 m, 21 Dec. 1967, Engel 2093 (MSC!). REMARKS— Because the species plays a conspic- uous role in the Falkland hepatic flora, yet may be confused with other hepatics in these islands, a description is provided below. Plants caespitose, rarely as scattered individuals among other Hepaticae, erect, to 2.3 cm tall, light green to golden brown or brown, often light green toward apex of plant and brown toward base, nitid. Branches frequent, exclusively of ventral-inter- calary type, leafy branches stiff, abruptly erect or prostrate and becoming erect, rarely becoming fla- gelliform toward the apex; stolons often arising from near base of ventral-intercalary branch, pros- trate to occasionally positively geotropic, with small scalelike leaves and scattered rhizoids, the stolons occasionally becoming upright and producing en- larged leaves. Stems 165-235 Mm wide, 135-200 Mm thick, in surface view the cortex cells short to long rect- angular, 1 3-23 Mm wide, 20-85(105) Mm long, with walls slightly to distinctly thickened, the end walls transverse to oblique. Stems in section 10-16 cells in diameter, cortex in 1-2 layers, the cortex poorly defined or distinct and then with very thick- walled cells, the cells 9-18 Mm thick, the cuticle smooth; medulla cells (13)16-30(33) Mm wide, thin walled but with slightly thickened corners. Leaves (0.5)0.55-1.1(1.2) mm long and wide, transverse to succubously oriented (succubous in robust plants), the insertion slightly to distinctly succubous, occasionally slightly recurved at ven- tral end; leaves erect to spreading, occasionally recurved, distant to imbricate, concave (often strongly so near dorsal base) to convex, orbicular to obliquely wide-ovate or obliquely wide-obovate in shape, the margins entire; apex acute to broadly rounded to truncate, occasionally slightly bifid and then divided to 0.15, the lobes acute to broadly rounded, the sinus triangular to rounded, the angle obtuse to narrowly rounded to lunate; dorsal mar- gin slightly to occasionally strongly curved, often repand, slightly to strongly inflexed, especially to- ward the base, not to very short decurrent; ventral margin strongly curved, usually more strongly curved than dorsal margin, the ventral margin often repand, plane to recurved, not to very short de- current. Leaf cells subquadrate to subrectangular, thin to slightly thickened, trigones absent to medium, occasionally with intermediate thickenings toward base; subapical cells 14—23 Mm wide and long, me- dian cells 14—24 Mm wide, 17-28 Mm long, the basal cells longer, 28—47 Mm long; marginal cells with nodular thickenings on outer radial walls, the basal marginal cells in 1-3 rows of shorter cells, 10-18 nm long, bordering the elongated basal-me- dian cells, but the leaves without a distinct differ- entiation of a vitta; cuticle smooth; oil-bodies not seen. Underleaves rudimentary, inconspicuous, con- sisting of only a small cluster of cells in close prox- imity to the ventral base of leaves. Gemmae rather rare, green, thin walled, ovate to elliptic, mostly 1 -celled. Opposite: FIG. 36. Adelanthus tenuis Engel & Grolle. 1-44, Leaves (those with same letter were obtained from same population), x 23. 45, Median leaf cells, x 545. 46, Basal-median leaf cells, x 515. 47, Stem, cross section, x 193. 48, Portion of upper, ventral leaf margin, x515. 49, Cortical cells, surface view, x515. (Figs. 1, 8, from Engel 3232; 2-4, from Engel 2533A; 3, 5-7, from Engel 3103; 9, 1 1, from Engel 2725; 10, 17, from Engel 2457; 12, 13, from Engel2773; 14, 20, from Engel 3221; 15, 1 6, from Engel 3343B; 18, 38, from Engel 3452; 19, 22, from Engel 2966; 21,31, from Engel 3003B; 23, 35, from Engel 3114; 24, 33, 39, 41, 45-49, from Engel 2093 (holotype); 25, 34, from Engel 2108C; 26, 44, from Engel 3453; 27, 40, 43, from Engel 3324A; 28, 29, 32, from Engel 3444; 30, 42, from Engel 2732; 36, 37, from Engel 2640C. Figs. 24-25, 33-34, 39, 41, and 45-49 from southern Chilean material, remainder from Falkland Is. material.) 86 FIELDIANA: BOTANY ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 87 37 FIG. 37. Wettsteinia densiretis (Herz.) Grolle. Plants dioecious. Androecia not seen. Gynoecia on reduced ventral-intercalary branches which arise from lower portion of plant, mature gynoecia not seen. VARIATION— Leaves, while usually oriented subsuccubous, may be transverse or, in robust plants, strongly succubous. The leaves vary con- siderably in size and configuration (figs. 36, 1-44). While the leaf apices exhibit a wide range of vari- ability, usually at least some leaves with slightly bifid apices are present on a given plant; bifid api- ces are an important taxonomic feature, and are useful in separating A. tenuis from "look-alikes." The ventral-intercalary branches exhibit consid- erable variation. Normal leafy branches may be- come flagelliform toward the apex, or conversely a stoloniform branch may become erect and pro- duce leaves typical of normal leafy shoots. In the latter, the stoloniform branches may be quite lengthy (to 1 1 mm) prior to becoming negatively geotropic and producing normal leaves. ECOLOGY— This species appears to be restricted to relatively high altitudes in the southern cold temperate region. In the Brunswick Peninsula it was collected at ca. 365 m in an open Sphagnum bog. In the Falklands this species is rather com- mon above 245 m, and occurs on soil under rock cover, in rock crevices, under ledges, etc., of out- crops in Cortaderia as well as dwarf shrub heath associations. Occasionally it may be found on soil in nonoutcrop situations in these associations, growing with Gackstroemia magellanica and Ade- lanthus lindenbergianus, or on soil over rock where it may be associated with Balantiopsis bisbifida and a filmy fern, Serpyllopsis caespitosa (Gaud.) v. d. Bosch. PH YTOGEOGRAPHY — M ayi'l 1 a n i a n- 1 a I k I a n d — Falkland Is.; Tierra del Fuego; and Brunswick Pen- insula (La. Parrillar region). It is of phytogeograph- ical interest to note that the New Zealand A. gem- miparus (Schust.) Hodgs. is a very close ally of this species. FALKLAND SPECIMENS SEEN -EAST FALKLANDS. Port William Region: summit ridge of Mt. Low, ca. 245 m (3220, 3221, 3224 A, 3232). Stanley Region: Sapper Hill, 2 km W of Port Stanley, 60 m, Ochyra 2852/80 (F); near summit of Tumbledown Mt., 230 m, Longton & Smith, Longton 972 (AAS, F); N peak of Two Sisters, 275 m, Dollman s.n. (Greene 3564a) (AAS, F); summit ridge of N peak of Two Sisters, 245-290 m (2725, 2752/1); summit of Mt. Kent, 455 m (2761 A —c. S, 2773— c. young per.). Mt. Usborne Region: gap between Mt. Us- borne 2 and Table Rock, ca. 440 m (2640Q; ridge be- tween Mts. Usborne 1 & 2, 685 m (2533A, 2554B); The Gap, 275-290 m (2457). WEST FALKLANDS. Port Howard: pass SW of Mt. Maria summit, ca. 610 m (3103— c. $, 3114). Fox Bay Region: summit of Fox Bay Mt., 307 m (3444-c. young per., 3452, 3453A). Mt. Adam: E slope of main peak, ca. 610 m (3003 B). Hill Cove Region: summit of East French Peak, 305 m (2966— c. young per., 2969); summit of Mt. Fegen, 335-360 m (3077). Wedded Island: Waterfall Valley, W of settle- ment, ca. 125 m (3324 A— c. young per.); summit of Mt. Weddell, 380 m (3343B). Adelanthus Species Excluded from Falklands Adelanthus falcatus (Hook.) Mitt. The report of Adelanthus falcatus from the Falk- land Islands in Stephani (1905a) is a misdeter- mined specimen of A. lindenbergianus, fide Grolle (in lift. ; specimens from S examined). Adelanthus (Calyptrocolea) falcatus is a species of New Zea- land (type from Dusky Bay, New Zealand, leg. Menzies), Tasmania and (presumably), southeast Australia (Grolle, 1972a). Wettsteinia Wettsteinia Schiffn., Ann. Jard. Bot. Buitenzorg. Suppl. 2: 44. 1898. 88 FIELDIANA: BOTANY Wettsteinia densiretis (Herz.) Grolle. Figure 37. Tylunanthus densiretis Herz. in Skottsberg, Nat. Hist. Juan Fernandez Is., Easter I. 2(5): 712. / 3a-d. 1942. Wettsteinia densiretis (Herz.) Grolle, J. Hattori Bot. Lab. 28: 99. 1965. Lectotype (fide Grolle 1965c): Juan Fernandez Is., Mas Afuera, Camp Correspondencia, 1140 m, 1917, Skotts- berg 92 (s, non vidi). ECOLOGY— This species is of rather frequent oc- currence in crevices of outcrops, as well as on soil in nonoutcrop situations in dwarf shrub heaths. It more rarely occurs on soil in Cortaderia heaths. PHYTOGEOGRAPH Y — Valdivian-Falkland — Falkland Is.; Valdivian region (ca. 40°S, 73°30'W); and Juan Fernandez Is., Mas Afuera (1 140 m) (see fig. 37). FALKLAND SPECIMENS SEEN— EAST FALKLANDS. Port William Region: summit ridge of Mt. Low, ca. 245 m (3225); ridge on Engineer Point Peninsula, 18-25 m (2823). Stanley Region: Tumbledown Mt., 155-230 m (2377 A. 2381); Goat Ridge, ca. 180 m (3196, 3204); summit ridge of N peak of Two Sisters, 245-290 m (2701 A). Mt. Usborne Region: below The Gap, ca. 90 m (2559, 2563). WEST FALKLANDS. Weddell Island: Waterfall Valley, W of settlement, ca. 125 m (3316); summit of peak NE of Mt. Weddell, 335 m (3369C, 3395). Family CEPHALOZIACEAE Cephaloziaceae Mig., Krypt.-Fl. Deutschl. ... 1: 465. 1904. The family has 14 genera and is worldwide in distribution. Only two genera— Cephalozia and Metahygrobiella— occur in south temperate-sub- antarctic areas, and both are in the Falklands. Cephalozia is worldwide and poorly represented in austral regions, while Metahygrobiella is pre- dominantly south temperate. Cephalozia Cephalozia (Dum.) Dum., Recueil Observ. Jun- germ. 18. 1 835. Jungermannia sect. Cephalo- zia Dum., Syll. Jungerm. Eur. Indig. 60. 1831. Cephalozia badia (Gott.) Steph. Jungermannia badia Gott., Ergebn. Deutsch. Polar- Exped. 2(16): 452. pi l.f. 1-5. 1890. Cephalozia badia (Gott.) Steph., Bull. Herb. Boissier II, 8(7): 483. 1908 (= Spec. Hep. 3: 313). Original ma- terial: South Georgia, Koppenberg, 10 Feb. 1883, Will 37 (o), cited in Fulford (1968) (non vidi). ECOLOGY— In the Falklands, this species was collected on a rock in a feldmark pool (ca. 700 m) and on soil under rock cover of a sheltered high altitude cliff. PHYTOGEOGRAPH Y — Antarctic in Distribu- tion—Antarctic Peninsula; Nelson and King George Islands (cf. Ochyra & Vana, 1 989); South Shetland Is. (Allison & Smith, 1973; Smith, 1979); South Sandwich Is. (Longton & Holdgate, 1979); South Georgia; Falkland Is.; and Tierra del Fuego (Lago Fagnano). Stephani (1911) stated that the Lago Fagnano collection is from a "Berge am Wes- tende von Lago Fagnano." LITERATURE RECORD (FALKLANDS)— Anony- mous (Futford, 1968). FALKLAND SPECIMENS SEEN— EAST FALKLANDS. Mt. Usborne Region: summit of Mt. Usborne 1, ca. 700 m (24968-c. per. + S). WEST FALKLANDS. Port Howard Region: pass SW of Mt. Maria summit, ca. 610 m (3104B). Notes on Cephalozia Species 1. Cephalozia bicuspidata (L.) Dum. There are three reports of this species from the Falklands, all based upon a Hooker collection(s). The taxon was reported (as Jungermannia) in Hooker and Taylor (1844), Taylor and Hooker (1847), and Gottsche et al. (1847). The species is widespread in the Northern Hemisphere, but Schuster ( 1 974) extended the range to include New Zealand with C. bicuspidata subsp. austrigena Schust. Vana (1988) added several more austral localities, namely South Africa, Marion Is., Crozet Is., Tristan da Cunha, and southern South Amer- ica (Brunswick Peninsula). With this emended dis- tribution, it is quite reasonable to expect that C. bicuspidata will eventually be recorded for the Falkland Is., but I have not as yet seen material. Metahygrobiella Metahygrobiella Schust., Bryologist 64: 205. 1961. Metahygrobiella tubulata (Hook. f. & Tayl.) Schust. ex Engel ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 89 Jungermannia tubulata Hook. f. & Tayl., London J. Bot. 3: 463. 1844. Cephalozia tubulata (Hook. f. & Tayl.) Trev., Mem. Reale 1st. Lomb. Sci. Lett. Ill, 4: 4 1 7. 1 877. Metahygrobiella tubulata Schust. ex Fulf., Mem. New York Bot. Card. 1 1(3): 322. 1968 nom. nud., pro syn. Metahygrobiella tubu- lata (Hook. f. & Tayl.) Schust. ex Engel, Bryol- ogist 79: 514. 1976. Original material: Falkland Is., Hooker s.n. (BM!). REMARKS— See comments in Engel (1978), which were based upon Falkland Islands plants (includ- ing type material). ECOLOGY— Rather rare in the Falklands. I en- countered the species in a creek in a Cortaderia heath and in wet depressions of dwarf shrub heaths, while Longton and Smith found the plant in a Sphagnum bog. PHYTOGEOGRAPHY— Valdivian + Magellanian + Falklands— Falkland Is.; Tierra del Fuego; north in Patagonian Channels to 44°1 9'S (Cerro Tesoro); and Valdivian West Patagonia. LITERATURE RECORDS (FALKLANDS)— Anony- mous (Kuhnemann, 1937, 1949;Bonner, 1963, as Cephalozia), Port Louis (Fulford, 1 968 as Cepha- lozia). Hooker (Fulford, 1968 as Cephalozia; G. L. & N., 1847, as Jungermannia; Taylor & Hook- er, 1847 as Jungermannia; Stephani, 1908 as Cephalozia). Skottsberg, Port Stanley (Stephani, 1905a, 1908; Skottsberg, 1913; Fulford, 1968 as Cephalozia). FALKLAND SPECIMENS SEEN— EAST FALKLANDS. Stanley Region: headwaters of Mullet Creek stream, ca. 60 m (3/55); between Mt. William and Tumbledown Mt., 170 m, Longton & Smith, Longton 980 (AAS, F); Goat Ridge, ca. 180 m (3207— c. per., 3214). Mt. Us- borne Region: gap between Mt. Usborne 2 and Table Rock. ca. 440 m (2627-c. 6). WEST FALKLANDS. Hill Cove Region: gap between French Peaks, 200-215 m (29798-c. per.). Family CEPHALOZIELLACEAE Cephaloziellaceae Douin, Bull. Soc. Bot. France 29: 1, 5, 13. 1920. Of the seven genera in this family, Allisoniella (five species) and the monoiypicAmphicephalozia occur only in south temperate regions; both are present in southern South America but are lacking from the Falklands. Cephaloziella (ca. 125 species in the broad sense) is worldwide, including both the antarctic and arctic, and occurs on the Falk- lands. Cephaloziella Cephaloziella (Spruce) SchifFn. in Engler & K. Prantl, Die Natiir. Pflanzenfam. 1(3,1): 98. 1893, nom. cons. Cephalozia subg. Cephalo- ziella Spruce, On Cephalozia 62. 1882. Cephaloziella dusenii Steph. Cephaloziella dusenii Steph., Bih. Kongl. Svenska Ve- tenskapsakad. Handl. 26 (III, 6): 49. 1900. Ceph- alozia dusenii (Steph.) Steph., Bull. Herb. Boissier II, 8(7): 504. 1908 (= Spec. Hep. 3: 334). Alobiella dusenii (Steph.) Steph., Bull. Herb. Boissier II, 8(8): 571. 1908 (= Spec. Hep. 3: 355), nom. illeg., non Alobiella dusenii Steph., Bih. Kongl. Svenska Vetenskapsakad. Handl. 26 (III, 6): 48. 1900 (= Alobiella stephanii Bonn., Candottea 14:98. 1953). Lectotype (fide Bonner, 1953): Chile, Prov. Val- divia, Corral, 5 June 1896, Dusen 94 (G!; isolec- totypes: s!— c. per., UPS!— c. per.). Cephalozia setistipa Steph., Spec. Hep. 6: 440. 1924, syn. nov. Original material: Falkland Is., Rabbit I., Halle s.n. (G 15158!— c. young per. + phoziaceae (J0rg.) Vand. Berg, in Robyns, Fl. Gen. Belgique 1: 221. 1956, nom. cons. The Lophoziaceae, containing 1 7 genera, are best developed in the cool and cold portions of the Northern Hemisphere. The largest genus, Lopho- zia, is widespread in boreal-subarctic-arctic re- gions, and only sporadically represented in the tropics and south temperate zone. Also notable is the rather large number of very small south tem- perate-subantarctic and tropical genera. Notho- strepta (2 species) and Roivainenia (1 species) are examples. Anastrophyllum Anastrophyllum (Spruce) Steph., Hedwigia 32: 1 39. 1893. Jungermannia subg. Anastrophyllum Spruce, J. Hot. 14: 235. 1876. Anastrophyllum cilia turn Steph. Anastrophyllum ciliatum Steph., Hedwigia 32: 139. 1893 ("Anastrophylium"). Sphenolobus ciliatus (Steph.) Steph., Bull. Herb. Boissier 2(2): 175. 1902 (= Spec. Hep. 2: 167). Original material: Argentina, Terr. Tierra del Fuego, I. de los Es- tados, Spegazzini s.n. (G!). Anastrophyllum pampaninii Gola, Nuovo Giorn. Bot. Ital. II, 29: 165. pi. 2,f. 1-2. 1923, syn.fideEngel (1978). Original material: Chile, Prov. Magal- lanes, B. Angelito, 300-400 m, 10 Feb. 1913, De Gasperi (FI!). REMARKS— This species is closely related to An- astrophyllum crebrifolium, a species of southern South America and northern Andes (see Engel, 1978). ECOLOGY— Usually mixed with other Hepaticae above 440 m in the Falklands; fairly common in sheltered high altitude cliffs under overhangs or on cushions of Azorella selago. Also collected in a feldmark (700 m), where it was associated with pools. PHYTOGEOGRAPHY — M agellanian-Falkland — South Georgia; Falkland Is.; southern Magellanian region (Isla de los Estados, Brunswick Peninsula, and Volcan Inga, Canal Gajardo). FALKLAND SPECIMENS SEEN -EAST FALKLANDS. Mt. Usborne Region: gap between Mt. Usborne 2 and Table Rock, ca. 440 m (2630— c. <3); ridge between Mts. Usborne 1 & 2, 685 m (2543 A, 2554E); summit of Mt. Usborne 1, ca. 700 m (2489A, 2491 A, 2496Q. WEST FALKLANDS. Port Howard: pass SW of Mt. Maria summit, ca. 610 m (3080B). Mt. Adam: side of summit ridge, 670-700 m (3014B). Anastrophyllum Species Excluded from Falklands Anastrophyllum subcomplicatum (Lehm. & Lin- denb.) Steph. This species was reported from the Falklands by Herzog (1926), who confused the type locality "Insulis Marianis" (Mariana Islands) with the Falklands (= lies Malouinae, Islas Malvinas). Andrewsianthus Andrewsianthus Schust., Rev. Bryol. Lichenol. 30: 66. 1961. Key to Falkland Islands Species of Andrewsianthus 1. Leaves often canaliculate or conduplicate, 0.25-0.3(0.5) bifid; leaf lobes acutely triangular, the angle of lobe outline 15°-65°(800), the apices occasionally acuminate or apiculate; sinus usually sharply triangular, occasionally narrowly rounded, rarely broadly rounded, never lunate; leaf insertion trans- verse, at most occasionally weakly succubous in dorsal half; ventral-intercalary branches very rare A. australis 1. Leaves usually plane to slightly concave, exceptionally canaliculate, never conduplicate, 0.1-0.25 bifid; leaf lobes acutely to obtusely triangular, the angle of lobe outline 45°-l 1 5°, the apices occasionally somewhat apiculate, never acuminate; sinus broadly rounded to lunate, occasionally triangular; leaf insertion often distinctly succubous; ventral-intercalary branches occasional A. planifolius 92 FIELDIANA: BOTANY 14 15^ 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 FIG. 38. Andrewsianthus australis Engel. 1, Stem, cross section, x 490. 2, Portion of plant with main axis becoming flagelliform, and with lateral-intercalary branch, dorsal view, xlO. 3, Median leaf cells with cuticular striations indicated in upper left, x5l5. 4, Apical portion of dorsal lobe, x5l5. 5, Leaf of main axis, x3i. 6, Ventral view of stem showing sessile, rudimentary underleaf, x5l5. 7, Ventral cortical cells, surface view, x5l5. 8, Slime papillae of basal region of dorsal margin of leaf, x515. 9-23, Leaves of main axis, x20. (Figs. 1-5, 7, 15, 18, 20, 25 from holotype; 6, 8, 14, 21, 22 from Engel 2219B. Chile, Prov. Magallanes, Pto. Cutter, N of copper mine; 10-12 from Engel 216 1G, Chile, Prov. Magallanes, Pto. Cutter, between copper mine and river S of mine; 13, 19, 24, from Engel 3075, Falkland Is., Mt. Fegen; 16-17, 26, from Engel 3020B, Falkland Is., Mt. Adam; 23, 37, from Engel 2301 B. Chile, Prov. Magallanes, Pto. Cutter, N side of copper mine; 28, 32-33, from Engel 3356. Falkland Is., Mt. Weddell; 29-31, from Engel 2743, Falkland Is., Mt. Kent.) Andrewsianthus australis Engel. Figures 38-39. Andrewsianthus australis Engel, Bryologist 75: 328. / 1-42. 1972. Holotype: Chile, Prov. Magallanes, Cta. Amalia, 1 Oct. 1969, Engel 5411 A (MSC!- c. per. + sporo. + <5). REMARKS— See Engel (1972). ECOLOGY— In the Falklands, gathered on soil over rock, or in rock crevices of cliffs, or large rock outcrops in dwarf shrub heaths above 335 m. On a single occasion I found it on soil under rock cover of sheltered high altitude cliffs. PHYTOGEOGRAPHY — Magellanian-Falkland — Falkland Is. and Patagonian Channels (north to 50°56'S, 73°52'W). ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 93 FIG. 39. Andrewsianthus australis Engel. 1, Portion of main axis with perianth, x 3 1 . 2, Seta, cross section, x 1 80. 3, Portion of perianth mouth with cellular detail indicated, x 180. 4, Spore, x515. 5, Capsule and portion of seta, x 3 1 . 6, Capsule wall, cross section, x 5 1 5. 7, Capsule wall, inner layer, x 5 1 5. 8, Capsule wall, outer layer, x 5 1 5. 9, Portion of axis bearing androecia, x31. (Figs. 1-8 from holotype; 9 from Engel 2743, Falkland Is., Mt. Kent.) FALKLAND SPECIMENS SEEN— EAST FALKLANDS. Stanley Region: summit of Mt. Kent, 455 m (2743— c. m p e r at i- in Distribution— The species has not penetrated north beyond temperate regions in any of the sectors. It has, however, penetrated into the subantarctic re- gion, as it occurs on Marion and Prince Edward Islands, Crozet Is., Kerguelen Is., and Macquarie I. Grolle (197 la) stated the records from Neo- tropical regions are erroneous and are mostly spec- imens of J. rubricaulis. Grolle also states the report from South Georgia by Stephani (1911) is based upon a specimen which is actually Cryptochila grandiflora (see fig. 45). LITERATURE RECORDS (FALKLANDS)— Anony- mous (Bonner, 1 966, as /. maluina; Kiihnemann, 1937, 1949, as J. colorata & J. maluina; Spruce, 1 876, as Jungermannia); Hooker (G. L. & N. 1 847, as Jungermannia; Hooker & Taylor, 1844, and Taylor & Hooker, 1847, as Jungermannia; Spruce, 1876, as Jungermannia); Lesson (Stephani, 1901, as J. maluina); Skottsberg, Port Stanley (Grolle, 197 la; Skottsberg, 1913; Stephani, 1905a, as /. colorata & J. maluina), Port Louis (Stephani, 1905a), Port Louis, Mt. Adam, Port William (Skottsberg, 1913), Hornby Mts. and between Ar- row Harbour House and Victoria Creek-Lafonia (Skottsberg, 1913, as J. maluina); Skottsberg & Halle, Port Stanley, Mt. Adam, Roy Cove, Wed- dell Is. (Stephani, 1911), Hornby Mts. (Stephani, 1 9 1 1 , as J. maluina). FALKLAND SPECIMENS SEEN -EAST FALKLANDS. Port William Region: Charles Point, 1954, Pettingill 3 (CINC, F); Sparrow Cove, 1954, Pettingill 45 (cmc, F). "Gottsche (1857) states the original collection was made in 1825. However, Moore (1968) stated Lesson, who was physician and assistant to Dumont d'Urville, was in the Falklands from 20 November- 18 December 1822. Stanley Region: Sapper Hill, 2 km W of Port Stanley, ca. 150 m, Ochyra 2872/80 (F); Sapper Hill, 2.5 km W of Port Stanley, ca. 100 m, Ochyra 2907/80 (F); Sapper Hill, 135 m (2422); Mt. William, rock pinnacles, 185 m, Taylor 224 (AAS, F); between Mt. William and Tumble- down Mt., 170 m, Longton & Smith. Longton 978 (AAS, F); Tumbledown Mt., 155-230 m (2374, 23828-c. per.); Wireless Ridge, near Stanley, 60 m, Longton & Smith, Longton 1016 (AAS, F); N peak of Two Sisters, 275 m, Dollman s.n. (Greene 3558a) (AAS); summit of Mt. Kent, 455m (2760/1). Mt. Usborne Region: ridge between Mts. Usborne 1 & 2, 685 m (2553C-C. per.); The Gap, 275- 290 m (2464); below The Gap, ca. 90 m (2555, 2555). WEST FALKLANDS. Fox Bay Region: valley E of Su- livan House, ca. 75 m (3530, 3533). Hill Cove Region: summit of West French Peak, 290 m (2959); valley near Hill Cove House, Valient in s.n. (MANCH). Roy Cove Re- gion: Roy Cove, Vallentin s.n. (MANCH). Weddell Island: summit of Mt. Weddell, 380 m (3341). Jungermannia Jungermannia L. Spec. PI. ed. 1. 1131. 1753. Jungermannia crassula Nees & Mont. Jungermannia crassula Nees & Mont., Ann. Sci. Nat. Bot. II, 5: 54. 1836. Solenostoma crassula (Nees & Mont.) Steph., Bull. Herb. Boissier 1(5): 497. 1 90 1 (= Spec. Hep. 2: 59). Original material: Juan Fernandez Is., Bertero (G, PC, s, w), cited in Vana ( 1974) (non vidi). Jungermannia domeikoana Mont., Ann. Sci. Nat. Bot. Ill, 4: 349. 1845, syn. fide Stephani (1901). Orig- inal material: southern Chile, without specific lo- cality, Gay (BM, FH, G, PC, s, w), cited in Vana (1974) (non vidi). Solenostoma obtusiflorum Steph., Kongl. Svenska Ve- tenskapsakad. Handl. 46(9): I6.f.5a. 1911, syn. fide Vana (1974). Original material: Juan Fer- nandez Is., Mas Afuera, Skottsberg (G, L, s), cited in Vana (1974) (non vidi). Solenostoma rostratum Steph., Kongl. Svenska Ve- tenskapsakad. Handl. 46(9): 16./ 5b. 1911, syn. fide Vana (1974). Original material: Juan Fer- nandez Is., Mas Afuera, Skottsberg (G, L, s), cited in Vana (1974) (non vidi). Jungermannia simplicissima Steph., Kongl. Svenska Vetenskapsakad. Handl. 46(9): 17./ 5c,d. 1911, syn. fide Vana (1974). Original material: Chile, Prov. Chiloe, I. Chiloe, Pto. Quellon, Halle & Skottsberg (G, s), cited in Vana (1974) (non vidi). Jamesoniella nana Steph., Spec. Hep. 6: 101. 1917, syn. fide Grolle (197 la). Original material: Falk- land Is., Halfway Cove, 1907, Halle (G, hb. Grolle), cited in Vana ( 1 974), (UPS), cited in Grolle (1971) and Vana (1974) (non vidi). REMARKS— Stephani (1911) used the name S. obtusiflorum, but altered the spelling to "obtusi- 102 F1ELDIANA: BOTANY flos" in Species Hepaticarum (Stephani, 1917, p. 82). ECOLOGY— Collected at a single locality in the Falklands, on a stream bank of a dwarf shrub heath. PHYTOGEOGRAPHY — Valdivian-Falkland — Tristan da Cunha, Inaccessible I., Gough I.; Falk- land Is.; Valdivian region (West Patagonia north to 36°50'S); Juan Fernandez Is. (Mas Afuera, Mas a Tierra). (Within the Falklands it was found only in the Port Stanley region, and the cooler temper- atures and higher rainfall may account for its pres- ence there.) LITERATURE RECORDS (FALKLANDS)— Dusen11, Halfway Cove (Bonner, 1966, as Jamesoniella nana\ Vana, 1974). FALKLAND SPECIMENS SEEN— EAST FALKLANDS. Stanley Region: headwaters of Mullet Creek Stream, ca. 60 m (3161A, 3172). Notes on Jungermannia Species Jungermannia stereocauli Bory in Gaudichaud, Botanique, Pt. 4, p. 1 30. In Freycinet, Voyage autour du monde . . . sur les corvettes de S. M. rUranieel la Physicienne. 1827 (1826). This species was described in Gaudichaud (1827) for a Falkland Islands specimen Gaudichaud col- lected. I have not been successful in numerous attempts, over a period of many years, to locate this specimen. It could not be found at PC, or at BM, BR, DS, E, FI, G, GL, H, L, Or W. It is quite impossible to ascertain the identity of the species from the original one line diagnosis: "Foliis ovato- concavis, integerrimis, imbricatis." The species name refers to the lichen Stereocaulon, and the only member of that genus mentioned in the Gau- dichaud Falkland collection is S. turfosum Bory. It was thought that perhaps the lichen might still have the J. stereocauli on the thallus, and that a separate collection of the hepatic had not been made. However, Lamb (1977, p. 326) stated that neither Bory's type, nor any other authentic Ste- reocaulon specimen from the Falklands, is known. The following comments by Stafleu and Cowan 17 Stephani cited Dusen as the collector of the speci- men; this, however, is incorrect, as Dusen did not visit the Falklands. Bonner ( 1 966) stated for the type of this species, "West Falkland, Halfway Cove, s.d., Dus£n s.n., in hb. G." Halfway Cove was visited by Halle and Skotts- berg, and Stephani determined their hepatic collections. In this case Halle made the collection. (1976, p. 921) are relevant: "Gaudichaud gave all his collection to P [and PC]. The collections made on the voyage in 1' Vranie and la Physicienne (1817- 1820) went directly to P. Part of the collections were lost during the voyage when the Vranie stranded in the Falkland Islands (lies Malouines)." Therefore, the possibility exists that a Gaudichaud description of J. stereocauli was saved but the specimen was lost. See also Sayre (1975, p. 326). Family GYMNOMITRIACEAE Gymnomitriaceae Klinggr., Die hoheren Crypto- gamen Preussens 16. 1858. There are nine genera in the Gymnomitriaceae; all are concentrated in arctic or arctic-alpine re- gions. Herzogobryum, however, is strictly austral, and members of that genus are typical of the alpine flora. Acrolophozia is also strictly austral: A. fue- giana occurs in the American sector, A. sulcata Hassel is endemic to South Georgia, and A. pec- tinata Schust. is found in New Zealand. Acrolophozia Acrolophozia Schust., Rev. Bryol. Lichenol. 34: 259. 1966; J. Hattori Bot. Lab. 26: 216, 279. 1963, nom. inval. sin. descr. lat. Acrolophozia fuegiana Schust. Acrolophozia fuegiana Schust., Nova Hedwigia 15: 494, pi 63. 1968. Holotype: Argentina, Terr. Tierra del Fuego, C. Garibaldi, ca. 900 m, Schus- ter 58317 (non vidi). ECOLOGY- PHYTOGEOGRAPHY— This subantarc- tic rare species was found at a single station in the sheltered high altitude cliff vegetation growing with Clasmatocoleafulvella and Herzogobryum teres on rock under a rock overhang. The Falkland ecology of A. fuegiana appears to be quite similar to that of the type, which was gathered on Cerro Gari- baldi, Tierra del Fuego, "ca. 900 m, on damp ledges in alpine tundra . . . associated with Pachyglossa dissitifolia, Clasmatocolea sp. (a species with poly- stratose leaves) [= Evansianthus georgiensis], and Herzogobryum teres" (Schuster 1 968b). All the as- sociates listed above are found in the sheltered high altitude cliff vegetation in the Falklands. The species, which is also known from South Georgia ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 103 (Hassel de Menendez, 1 980a), is very closely allied to A. pectinata Schust. of South Island, New Zea- land. It is significant to note that the New Zealand species is known from ca. 1370-1460 m, on soil over shaded ledges in the snow tussock (alpine) zone. FALKLAND SPECIMEN SEEN- EAST FALKLANDS. Mt. Usborne Region: ridge between Mts. Usborne 1 & 2, 685 m (2521C-C. per.). Herzogobryum Herzogobryum Grolle, Rev. Bryol. Lichenol. 32: 160. 1964 (1963). Chondrophyllum Herz., Rev. Bryol. Lichenol. 21: 46. 1952, non Chondrophyllum Kylin, Acta Univ. Lund. n. s. 20(6): 442. 1924 (Rhodophyta). Key to Falkland Islands Species of Herzogobryum 1 . Cells of leaf margins with considerably thicker walls than median lamina cells; plants monoecious; antheridia paired; perianth not exserted beyond bracts H. erosum 1 . Cells of leaf margins with thinner walls than median lamina cells; plants dioecious; antheridia single; perianth exserted far beyond bracts 2 2. Leaves cucullate; leaf margins plane to weakly crenulate; leaf apex not fissured H. vermiculare 2. Leaves concave; leaf margins strongly crenulate; leaf apex often fissured H. teres Herzogobryum erosum (Caning. & Pears.) Grolle. Fig. 46. Cesia erosa Carring. & Pears., Pap. & Proc. Roy. Soc. Tasmania 1887: 8. pi. 6, f. 1-19. 1888. Gymno- mitrium erosum (Carring. & Pears.) Bast., Pap. & Proc. Roy. Soc. Tasmania 1887: 244. 1888 ("ero- sa"). Herzogobryum erosum (Carring. & Pears.) Grolle, Oesterr. Bot. Z. 1 13: 231. 1966. Original material: Tasmania, Bastow s.n. (BM), cited in Grolle (1966a)(m?H vidi). Cesia stygia var. denticulata Berggr., New Zealand Hep. 4. 1898, syn. fide Grolle (1966a). Acolea denticulata (Berggr.) Steph., Bull. Herb. Boissier 1(2): 143. 1901 (= Spec. Hep. 2: 4). Gymnomit- rium denticulatum (Berggr.) K. Mull. (Freib.) Rev. Bryol. Lichenol. 20: 176-177. 1951. Lectotype (/KteGrolle, 1 966a): New Zealand, South I., West- land, Kelly's Hill, 1200 m, 1874, Berggren 2870 (LD, non vidi). ECOLOGY— Collected only once in the Falk- lands, on sandy soil under a rock ledge of sheltered high altitude cliffs. PHYTOGEOGRAPHY— Subantarctic in Distribu- tion—South Georgia; Falkland Is.; Tierra del Fue- go, Isla de los Estados; southern Patagonian Chan- nels, Brunswick Peninsula; Tristan da Cunha; 500- 2000 m in the New Zealand sector, occurring on Campbell I., Stewart I., and north to Tasmania (see fig. 46). FALKLAND SPECIMEN SEEN -WEST FALKLANDS. Mt. Adam: summit of southernmost peak, 685 m (2986A—C. per. + $). Herzogobryum teres (Carring. & Pears.) Grolle. Figure 47. Jungermannia teres Carring. & Pears., Pap. & Proc. Roy. Soc. Tasmania 1887: 9. pi. 7,f. 1-16. 1888 ("Jungermania"). Jamesoniella teres (Carring. & Pears.) Steph., Bull. Herb. Boissier 1(10): 1037. 1901 (= Spec. Hep. 2: 100). Herzogobryum teres (Carring. & Pears.) Grolle, Oesterr. Bot. Z. 113: 223. 1966. Lectotype (cf. Grolle, 1966a): Tas- mania, Mt. Wellington, Bastow s.n. (MANCH, non vidi). Acolea magellanica Mass. & Steph. in Massalongo, Atti Reale Inst. Veneto Sci. 87: 237. pi. 3, f. 5- 7. 1927, syn. fide Grolle (1966a). Original ma- terial: Argentina, Terr. Tierra del Fuego, I. de los Estados, Penguin Rookery, Spegazzini s.n. (non vidi). Jamesoniella inflexo-limbata Herz., Trans. & Proc. 104 FIELDIANA: BOTANY 46 FIG. 46. Distribution of Herzogobryum erosum (Carring. & Pears.) Grolle. FIG. 47. Distribution of Herzogobryum teres (Carring. & Pears.) Grolle. ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 105 Roy. Soc. New Zealand 68: 41. 1938, syn. fide Grolle (1966a). Lectotype (cf. Grolle, 1966a): New Zealand, North I., Ml. Tongariro, 12 Jan. 1933, Moore 130 (JE, non vidi). ECOLOGY— In the Falklands, this taxon seems restricted to sheltered high altitude cliffs, where it occurs very commonly with Austrolophozia fue- giensis, Balantiopsis bisbifida, Cryptochila spp., Pachyglossa spp., and Triandrophyllum subtrifi- dum. It grows in moist, protected rock crevices, frequently under rock overhangs. Schuster ( 1 968b, p. 499) included this species as an associate of Acrolophoziafuegiana, which was collected at Cer- ro Garibaldi, Tierra del Fuego at "ca. 900 m, on damp ledges in alpine tundra" (p. 499), an eco- logical situation which apparently is quite similar to the sheltered high altitude cliffs in the Falklands. PHYTOGEOGRAPHY— Antarctic in Distribu- tion—South Shetland Is. (Ochyra and Vana, 1989); South Orkney Is. (Smith, 1972); Falkland Is.; Ti- erra del Fuego; Tristan da Cunha; Gough I.; Campbell I.; 1200-2300 m in New Zealand and Tasmania; Marion I. (montane) (fig. 47). FALKLAND SPECIMENS SEEN- EAST FALKLANDS. Mt. Usborne Region: ridge between Mts. Usborne 1 & 2, 685 m (2575-c. $, 2517 A, 2521A-C. 6, 2527, 2528- c. per. + 3, 2530, 2554G). WEST FALKLANDS. Port Howard: pass SW of Mt. Maria summit, ca. 610 m (3087-c. per., 3098-c. $, 3112B, 3315B-C. <5, 3127 'A- c. $, 3129C). Mt. Adam: ridge S of northern lake, 610 m (30 30 A -c. per. + <3, 3036B). Herzogobryum vermiculare (Schiffn.) Grolle. Fig- ure 48. Gymnomitrium vermiculare Schiffn. in Naumann, Forschungsr. Gazelle 4(4): 2. pi. l,f. 9,10. 1890. Herzogobryum vermiculare (Schiffn.) Grolle, J. Hattori Bot. Lab. 28: 103. 1965. Original mate- rial: Kerguelen Is., Successful Harbor, 1 874, Nau- mann s.n. (w), cited in Grolle (1965d) (non vidi). Chondrophyllum cucullatum Herz., Rev. Bryol. Li- chenol. 21: 46. / 1-2. 1952, syn. fide Grolle ( 1 965d). Herzogobryum cucullatum (Herz.) Grolle, Rev. Bryol. Lichenol. 32: 160. 1963. Original ma- terial: Chile, Prov. Aisen, C. Tesoro, ca. 1000 m, 1 3 Feb. 1 940, Schwabe 39cp.p. (JE), cited in Grolle (I965d)(non vidi). ECOLOGY— Found in a moist feldmark (700 m), on Azorella cushions, soil over rock, and in rock crevices of sheltered high altitude cliffs. It is very frequently mixed with other hepatics, particularly with Cryptochila spp., Pachyglossa spp., and Her- zogobryum teres. PHYTOGEOGRAPHY— Subantarctic in Distribu- tion—South Georgia; Falkland Is.; southern South America only at ca. 1 000 m in the Valdi vian region (44°19'S); Kerguelen Is.; Marion I. (see fig. 48). LITERATURE RECORD (FALKLANDS)— Anony- mous (Grolle, 1 969). FALKLAND SPECIMENS SEEN— EAST FALKLANDS. Mt. Usborne Region: ridge between Mts. Usborne 1 & 2, 685 m (25555); summit of Mt. Usborne 1, ca. 700 m (2505D). WEST FALKLANDS. Port Howard: pass SW of Mt. Maria summit, ca. 6 10 m (3097 B-c. per., 3110A, 3111.3129A). Mt. Adam: summit of southernmost peak, 685 m (2986B). Family SCAPANIACEAE Scapaniaceae Migula, Krypt.-Fl. Deutschl. 1: 479. 1904. Two genera of this basically Laurasian family occur regionally— Diplophyllum and Scapania. Scapania is the largest genus, with ca. 58 species. Scapania gamundiae Schust., of southern South America, is the only austral member of the genus and also stands alone in having subequally bilobed leaves. Diplophyllum Diplophyllum (Dum.) Dum., Recueil Observ. Jun- germ. 15.1835, nom. cons. Jungermannia sect. Diplophyllum Dum., Syll. Jungerm. Eur. In- dig. 44. 1831. Diplophyllum acutilobum Steph. Diplophyllum acutilobum Steph., Kongl. Svenska Ve- tenskapsakad. Handl. 46(9): 83. / 32f. 1911. Original material: Chile, Prov. Maga llanos. W end of L. Fagnano, Halle & Skottsberg s.n. (s), cited in Grolle (1965b) (non vidi). REMARKS— See comments in Schuster (1974, p. 193). ECOLOGY— Rare; collected at a single locality, on soil under a rock overhang of sheltered high altitude cliffs. PHYTOGEOGRAPHY— Valdi vian + Magellanian + Falklands— Falkland Is.; Tierra del Fuego, Lago Fagnano; Patagonian Channels; the Valdivian re- gion north to 40°42'S. 106 FIELDIANA: BOTANY 48 FIG. 48. Distribution of Herzogobryum vermiculare (Schiffn.) Grolle. FIG. 49. Distribution Blepharidophyllum clandestinum (Mont.) Lac. ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 107 FALKLAND SPECIMEN SEEN -WEST FALKLANDS. Mt. Adam: ridge S of northern lake, 610 m (3027— c. per. + i). Family BLEPHARIDOPHYLLACEAE Blepharidophyllaceae (Schust.) Schust. ex Engel, comb. nov. Scapaniaceae subf. Blepharido- phylloideae Schust., Hep. Anthoc. N. Amer. 3: 175. 1974. Blepharidophyllaceae (Schust.) Schust. in Clarke & Duckett, Syst. Assoc. Spe- cial Vol. 14: 74. 1980, nom. inval. This family of two genera is strictly austral in distribution. Blepharidophyllum has five species, three centered in the American sector and two Australasian. The southern South American Kru- nodiplophyllum is monotypic. Blepharidophyllum Blepharidophyllum Angstr., Ofvers. Forh. Kongl. Svenska Vetenskapsakad. 30: 151. 1873. Key to Falkland Islands Species of Klepharidophyllum'' 1 . Leaves not keeled, not conduplicately bilobed, bifid to the middle, the insertion curved; leaf cells with large knotlike trigones; perianth mouth with long cilia which consist of long cylindrical cells B. densifolium 1. Leaves keeled, conduplicately bilobed, divided 0.75-0.9 into 2 lobes, the insertion nearly transverse; leaf cells ± equally thick walled, or with thin walls and small trigones; perianth mouth with teeth and short spines, the latter of 1-2 massive, extended cells 2 2. Leaf margins irregular and with numerous small teeth, at least on antical margin of dorsal lobe; leaf segments apiculate; cell walls thick, trigones absent B. clandestinum 2. Leaf margins entire; leaf segments triangular, not apiculate; cell walls thin, with small trigones B. gottscheanum Blepharidophyllum clandestinum (Mont.) Lac. Figure 49. Plagiochila (Scapania) dandestina Mont., Ann. Sci. Nat. Bot. II, 19: 247. 1843. ? Scapania dandes- tina (Mont.) G. L. & N., Syn. Hep. 73. 1844. Jungermannia dandestina (Mont.) Hook. f. & Tayl. i/i Hooker f., Bot. Antarc. Voyage 1(2): 434. 1 847. Martinellia dandestina (Mont.) Trev., Mem. Reale 1st. Lomb. Sci. Lett. Ill, 4: 41 1. 1877. Bal- antiopsis dandestina (Mont.) Schiffn. in Engler & Prantl, Naturl. Pflanzenfam. 1(3, 1): 112. 1893. Diplophyllum dandestinum (Mont.) Steph., Bih. Kongl. Svenska Vetenskapsakad. Handl. 26 (III, 6): 61. 1900. Blepharidophyllum dandestinum (Mont.) Lac., Genera Hep. 27. 1910. Clandarium clandestinum (Mont.) Schust., New Manual Bryology 1: 541. 1983. Original material: (fide Grolle, 1965b, exd. B. gottscheanum): Chile, Prov. Magallanes, Pto. Gallant, "Inter caespites Hypni fluitantis," Hombron (PC, non vidi). Balantiopsis incrassata Mitt., J. Linn. Soc., Bot. 15: 197. 1876, syn.fide Mitten (1879). Original ma- terial: Kerguelen Is., Hill NW of Mt. Crozier, Ea- ton (NY!). 18 Key adapted and modified from Grolle (1965b). ECOLOGY— Rather local in occurrence in the Falklands, where I found it on soil in a Cortaderia heath, on the bases ofBlechnum magellanicum in a dwarf shrub heath, and on a sheltered high al- titude cliff. PHYTOGEOGRAPHY — Amphiatlantic Temper- ate—Falkland Is.; Tierra del Fuego; Patagonian Channels; Valdivian region (to 42°30'S in West Patagonia); also on Kerguelen (see fig. 49). FALKLAND SPECIMENS SEEN -EAST FALKLANDS. Mt. Usborne Region: ridge between Mts. Usborne 1 & 2, 685 m (2525A)- below The Gap, ca. 90 m (2566). WEST FALKLANDS. Weddell Island: Waterfall Val- ley, west of settlement, ca. 125 m (3313A—C. $, 3318). Blepharidophyllum densifolium (Hook.) Angstr. ex Mass. Figure 50. Jungermannia densifolia Hook., Musci Exot. 1 : pi. 36, f. 1-4. 1818. ? Scapania densifolia (Hook.) Nees in G. L. & N., Syn. Hep 72. 1844. Diplophyllum densifolium (Hook.) Mitt., J. Linn. Soc., Bot. 1 5: 69. 1876. Martinellia densifolia (Hook.) Trev., 108 FIELDIANA: BOTANY Mem. Reale 1st. Lomb. Sci. Lett. Ill, 4: 4 1 1 . 1 877. Blepharidophyllum densifolium (Hook.) Angstr. ex Mass., Nuovo Giorn. Bot. Ital. 17: 208. 1885. Original material: Argentina, Terr. Tierra del Fuego, I. de los Estados, Menzies (NY!, s, w), cited inGrolle(1965b). Jungermannia chloroleuca Hook. f. & Tayl., London J. Bot. 3: 467. 1844, syn.fide Bescherelle & Mas- salongo ( 1 889). Scapania chloroleuca (Hook. f. & Tayl.) G. L. & N., Syn. Hep. 662. 1847. Schis- tocalyx chloroleuca (Hook. f. & Tayl.) Lindb., J. Linn. Soc., Bot. 13: 185. 1873, nom. illeg. Mar- tinellia chloroleuca (Hook. f. & Tayl.) Trev., Mem. Reale 1st. Lomb. Sci. Lett. Ill, 4: 41 1. 1877. Ble- pharidophyllum vertebrale var. /3 chloroleucum (Hook. f. & Tayl.) Mass., Nuovo Giorn. Bot. Ital. 1 7: 208. 1885. Blepharidophyllum densifolium var. 7 chloroleucum (Hook. f. & Tayl.) Schiffn. in Nau- mann, Forschungsr. Gazelle 4(4): 9. 1 890. Diplo- phyllum vertebrale var. /3 chloroleucum (Hook. f. & Tayl.) Mass., Atti Reale 1st. Veneto Sci. 87: 221. 1927. Original material: Chile, Prov. Ma- gallanes, I. Hermite, Hooker (NY!). Scapania pycnophylla De Not., Mem. Reale Accad. Sci. Torino II, 16: 215. pi. 3, f. 1-6. 1855, syn. fide Bescherelle & Massalongo (1889). Martinellia pycnophylla (De Not.) Trev., Mem. Reale 1st. Lomb. Sci. Lett. Ill, 4: 411. 1877. Wlepharido- phyllum pycnophyllum (De Not.) Angstr. ex Mass., Nuovo Giorn. Bot. Ital. 17: 208. 1885. Blephar- idophyllum densifolium var. < pycnophyllum (De Not.) Schiffn. in Naumann, Forschungsr. Gazelle 4(4): 9. 1890. Diplophyllum pycnophyllum (De Not.) Steph., Result. Voyage S. Y. Belgica 6(5): 6. 1901. Original material: Chile, "Prov. Valpa- raiso, Valparaiso," Puccio (NY!). Blepharidophyllum fuscum Besch. in Bescherelle & Massalongo, Mission Sci. Cap Horn 5: 209. 1889, nom. nud. pro syn. Blepharidophyllum densifol- ium var. 6 fuscum (Besch.) Schiffn. in Naumann, Forschungsr. Gazelle 4(4): 9. 1 890. ECOLOGY— In southern South America, the species is quite common and exhibits a wide eco- logical amplitude. It is common in mounds of bryophytes in the very wet Magellanian moorland region, as well as in the mesic, deciduous forested zones. The species is surprisingly local in the Falk- lands, where it was found in pools, rock crevices, and between Bolax as well as Azorella cushions on sheltered high altitude cliffs. It rarely occurs on soil of dwarf shrub heaths. On Marion and Prince Edward Islands B. densifolium is the dominant plant species in a component of the "Swamp Com- plex" termed a "Blepharidophyllum densifolium bog" by Huntley (1971). This "nodum" is char- acterized by nearly stagnant anaerobic conditions and by a deep, primarily bryophyte carpet with usually less than 20% vascular plant cover. (See also Gremmen, 1982, who has a "Juncus scheuchzerioides-Blepharidophyllum densifolium complex" for the same islands.) PHYTOGEOGRAPHY— Amphiatlantic Temper- ate—South Georgia; Falkland Is.; Tierra del Fue- go; Patagonian Channels and north in the Valdi- vian region (West Patagonia) to 36°50'S. Quite well represented in the subantarctic, on Kerguelen Is., Crozet Is., Marion and Prince Edward Is., and Gough I. (see fig. 50). FALKLAND SPECIMENS SEEN -EAST FALKLANDS. Stanley Region: Sapper Hill, 2 km W of Port Stanley, 100 m, Ochyra 2876/80 (F); S slopes of Mt. William, 1 85 m, Longton & Smith, Longton 989 (AAS, F); summit ridge of N peak of Two Sisters, 245-290 m (2727). Mt. Usborne Region: ridge between Mts. Usborne 1 & 2, 685 m (2526); summit of Mt. Usborne 1, ca. 700 m (2490, 2493A, 2494, 2504A). WEST FALKLANDS. Mt. Adam: ridge S of northern lake, 610 m (3032). Blepharidophyllum gottscheanum (nolle. Figure 51. Blepharidophyllum gottscheanum Grolle, J. Hattori Bot. Lab. 28: 69. / 4. 1965. Clandarium gotts- cheanum (Mont.) Schust., New Manual Bryology 1: 541. 1983. Holotype: Chile, Prov. Magallanes, I. Desolation, Dusen (JE, non vidi) cited in Grolle (1965b). ECOLOGY— I found this species at but a single locality in the Falklands, where it grew in a Cor- taderia heath at 90 m. PHYTOGEOGRAPHY— Valdivian + Magellanian + Falklands— Falkland Is.; western Tierra del Fuego; Patagonian Channels and north to 40°45'S in the Valdivian region (West Patagonia) (see fig. 51). FALKLAND SPECIMENS SEEN— EAST FALKLANDS. Mt. Usborne Region: below The Gap, ca. 90 m (2562B, 2578B). Family GEOCALYCACEAE Geocalycaceae Klinggr., Die hoheren Cryptoga- men Preussens 34. 1858 ("Geocalyceae"). Geocalycaceae, composed of 20 genera, are worldwide in distribution but much better repre- sented in the Southern Hemisphere. Several small genera are restricted to south temperate-subant- arctic regions (e.g., Evansianthus and Pigafettoa [each 1 sp.], Leptophyllopsis [2 spp.], Pachyglossa [4 spp.], and the larger genera are best represented there [e.g., Clasmatocolea, Heteroscyphus]). The largest genus is Chiloscyphus (s. lat. to include Lophocoled). ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 109 50 , FIG. 50. Distribution of Blepharidophyllum densifolium (Hook.) Angstr. ex Mass. FIG. 51. Distribution of Blepharidophyllum gottscheanum Grolle. Open circle indicates pre- cise locality unknown. 110 FTELDIANA: BOTANY Chiloscyphus Chiloscyphus Corda in Opiz (ed.), Beitrage zur Naturg. ( 1 2): 65 1 . 1 829 ("Cheilocyphos"); coir. Dumortier, Sylloge Jung. Europ. Indig. 67. 1831, nom. et orth. cons. Lophocolea (Dum.) Dum., Recueil Observ. Jungerm. 17. 1835. Jungermannia sect. Lophocolea Dum., Syll. Jungerm. Eur. Indig. 59. 1831. Key to Falkland Islands Species of Chiloscyphus 1. Leaves consistently or at least predominantly bifid 2 2. Leaf lobes and marginal teeth caducous, often giving leaf apices a ragged appearance; leaves often with accessory teeth and laciniae; leaf margins entire to highly dentate. Underleaves deeply bifid, with margins 1-laciniate, and lobes spreading; perianths twice as long as broad; either margin of innermost bract to 8-dentate-laciniate; plants dioecious [Leptophyllopsis irregularis] 2. Leaf lobes and marginal teeth, if present, persistent, leaf apices never with a ragged appearance; leaves not with accessory teeth and laciniae; leaf margins entire to 1-dentate 3 3. Leaves bifid to (0.3)0.4-0.5; leaf segments frequently ventrally sulcate 4 4. Leaf segments widely divergent; leaves basically weakly to strongly cuneate, the lamina widest in distal third; underleaf margins mostly entire, sporadically with a small tooth . . C. divaricatus 4. Leaf segments ± parallel with lamina margins at most only sporadically divergent; leaves basically ovate to subrectangular, the lamina widest in basal third; underleaf margins usually with 1 lacinium or tooth C. bispinosus 3. Leaves bidentate or bifid to at most 0.25; leaf segments plane, never ventrally sulcate .. 5 5. Underleaves connate with the leaves on both sides. Leaves symmetrically ovate or nearly so to subrectangular in shape, the sinus truncate or lunate C. sylvaticus 5. Underleaves connate on only one side or completely free 6 6. Median leaf cells (46)52-78 /*m x 33-65 jmi; perianth terete or obscurely triangular below. Dorsal leaf margin straight or nearly so, entire; ventral margin rounded, especially towards the apex, entire to 1-dentate; underleaves elongate-ovate in shape, as wide as or slightly wider than stem, connate on one side C. textilis 6. Median leaf cells 17-48(52) x 20-43 Aim; perianth triangular below 7 7. Leaf lobes variable; 1 or both commonly rather broadly rounded (often with 1 lobe rounded and the other triangular), or both narrowly triangular. Lobes asymmetric, with one lobe often larger than the other; trigones distinct C. humifusus 1. Leaf lobes consistent in shape, narrow to broadly triangular or piliferous. Lobes ± symmetric 8 8. Leaf segments ending in a uniseriate row of 1-6 cells, the segments narrow to broadly triangular. Underleaf margins entire to 1-dentate or occasionally ciliate- small laciniate; plants monoecious C. lentus 8. Leaf segments ending in a uniseriate row of 4-15 cells, the segments piliform. Underleaves deeply bifid (nearly always almost to base), the lamina margins usually with a single long lacinium-lobe; perianths (3.1)3.6-5.3 x longer than broad, the lobes narrowly triangular, dentate-laciniate; plants monoecious .... C. leptanthus 1 . Leaves (well developed) undivided, emarginate or 1-dentate 9 9. Living plant whitish to light to yellow-green or tawny, without wall pigments. Gynoecia isophyllous or subisophyllous: bracteole 0.75-0.95 the area of the bracts 10 10. Underleaf apices plane to retuse to bidentate; underleaves/re<7w^rt//y convex to ± canaliculate, the margins entire. Underleaves connate on one side; leaf apices entire .... C. austrigenus ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 1 1 1 10. Underleaves bifid to ca. 0.5 or more; underleaves plane and not convex, the margins entire to 4-dentate-laciniate 11 1 1 . Leaves expanded near ventral base and here often reflexed; underleaves 0.5 x stem width to as wide as stem. Leaf apices broadly rounded, never retuse; underleaf segments often differing in size and configuration; stems thick, fleshy C. elatus \ 1 . Leaves not expanded near ventral base; underleaves never less than stem width ... 12 1 2. Leaf apices variable, broadly rounded to truncate to retuse to emarginate to 1 - or bidentate; underleaves as wide as to 1.5x stem width; underleaf margins entire to 2-dentate-small laciniate C. sabuletorum 12. Leaf apices broadly rounded to truncate, never retuse; underleaves (l-)3x stem width; underleaf margins to 4-dentate-large laciniate C. semiteres 9. Living plant warm to castaneous brown, with wall pigmentation. Gynoecia strongly anisophyllous: bracteole ca. 0.25-0.3 as large as bracts, 0.5-0.75 bifid, underleaf-like and often with 1 lateral lacinium on 1 or both sides. Perianths most often on abbreviated lateral-intercalary branches without vegetative leaves C. hookeri Chiloscyphus austrigenus (Hook. f. & Tayl.) Engel & Schust. Figure 52. Jungermannia austrigena Hook. f. & Tayl., London J. Bot. 3: 466. 1 844. Lophocolea austrigena (Hook, f. & Tayl.) G. L. & N., Syn. Hep. 702. 1847. Chiloscyphus austrigenus (Hook. f. & Tayl.) Engel & Schust., Nova Hedwigia 39: 411. 1984. Orig- inal material: Chile, Prov. Magallanes, I. Hermite and/or Cabo de Hornos, Hooker s.n. (BM!— c. per.; NY!— c. per.; s!; w, cited in Grolle, 1962). Jungermannia cavispina Hook. f. & Tayl., London J. Bot. 3: 463. 1844, syn. fide Bescherelle & Mas- salongo (1 889). Lophocolea cavispina (Hook. f. & Tayl.) Fulf., Mem. New York Bot. Gard. 1 1 : 483. 1976, nom. illeg. (ICBN, Art. 57). Lectotype (fide Engel, 1978): Falkland Is., 1843, Hooker s.n. (FH!). Lophocolea triseriata Steph., Bih. Kongl. Svenska Ve- tenskapsakad. Handl. 26 (III, 6): 45. 1900, syn. fide Fulford (1976). Original material: Chile, Prov. Aisen, R. Aisen Valley, 13 Jan. 1897, Dusen s.n. (NY!, s!— c. per., UPS!). Leioscyphus grandistipus Steph., Kongl. Svenska Ve- tenskapsakad. Handl. 46(9): 37. / 13c,d. 1911, syn. fide Grolle ( 1 962). Leptoscyphus grandistipus (Steph.) Kiihnem., Revista Centre Estud. Doct. Ci. Nat. Buenos Aires 1: 176. 1937. Mylia gran- distipa (Steph.) Kuhnem., Lilloa 19: 341. 1949. Lectotype (fide Engel, 1 978): Falkland Is., Hornby Mts., 19 Dec. 1907, Skottsberg 346 (s!). Lophocolea falklandica Steph., Kongl. Svenska Ve- tenskapsakad. Handl. 46(9): 44./ 16e. 191 1, syn. fide Grolle (1962). Lectotype (fide Engel, 1978): Falkland Is., Westpoint I., 8 Dec. 1907, Skotts- berg 351 (s!— c. S). REMARKS— See Engel (1978). Chiloscyphus austrigenus is the correct name for this species, and not Lophocolea cavispina, as Ful- ford (1976) would have it; see Engel (1979) and Grolle (1979). Further, Fulford erroneously in- cluded several synonyms of this species. Chiloscy- phus otiphyllus is a well-differentiated species, with Chiloscyphus notophylloides Mass, and Leioscy- phus oppositifolius Steph. as synonyms (see Engel, 1978). Lophocolea magellanica Schiffn. is a syn- onym of Clasmatocolea humilis var. humilis (fide Engel, 1980). Several syntype specimens of Lophocolea falk- landica have been misdetermined. These speci- mens are referable to Balantiopsis erinacea and Chiloscyphus elatus. The description (especially of the underleaf) and figures, however, agree with the syntypes referable to Chiloscyphus austrigenus. I have designated a Stockholm (S) specimen from Westpoint Island, Falkland Islands, as the lecto- type. Chiloscyphus elatus is mixed with C. austri- genus in the lectotype packet. ECOLOGY— This taxon is rather common in and typical of the bryoflora of streams in Cortaderia and dwarf shrub heaths. It is especially luxuriant when submerged, and then frequently forms thick, pure, extensive masses. It occasionally may be found on stream banks. PHYTOGEOGRAPHY— Valdivian + Magellanian Falklands— Falkland Is.; Tierra del Fuego; Pata- gonian Channels; Valdivian region (north to 39°52'S); Tristan da Cunha; and Inaccessible I. (see fig. 52). There have been no authentic reports from Andean Patagonia; the report in Stephani (1911) from Estrecho Miguens is based upon a specimen of Clasmatocolea vermicularis. The re- ports from the New Zealand sector require con- firmation. LITERATURE RECORDS (FALKLANDS)— Anony- mous (Kiihnemann, 1937, as Leptoscyphus gran- distipus, Lophocolea austrigena & L. falklandica; 112 FIELDIANA: BOTANY 52 FIG. 52. Distribution of Chi- loscyphus auslrigenus (Hook. f. & Tayl.) Engel & Schust., plus Tris- tan da Cunha. Open circle indi- cates precise locality unknown. FIG. 53. Distribution of Chi- loscyphus elatus (Gott.) Engel & Schust. 53 ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 113 Kiihnemann, 1949 as L. austrigena, L. falklandica & Mylia grandistipus; Rodway, 1916); Abbott (Fulford, 1976, as Lophocolea cavispina); Hooker (G. L. & N. 1 847 as J. cavispina; Taylor & Hooker, 1847, as J. cavispina; Stephani, 1906); Lechler, Mt. William, Port Stanley (Fulford, 1976, as L. cavispina); Skottsberg (Stephani, 1 922, as Leioscy- phus grandistipus & Lophocolea falklandica), be- tween Arrow Harbour House and Victoria Creek (Skottsberg, 1913, as Leioscyphus grandistipus); Hornby Mts. (Grolle, 1 962); Port Stanley (Fulford, 1976, as Lophocolea cavispina); Port Stanley, Port Louis (Skottsberg, 1913; Stephani, 1905a); West- point Is. (Stephani, 1913, as L. falklandica); Skottsberg & Halle, Murrell River (Stephani, 1911; Fulford, 1976, as Lophocolea cavispina). FALKLAND SPECIMENS SEEN— without specific locality, Capt. Abbott (BM, NY-C. per.). EAST FALKLANDS. 1 843, Hooker, syntypes of Jungermannia cavispina (BM, s). Stanley Region: Port Stanley, 28 Mar. 1902, Skotts- berg s.n. (s); Port Stanley, Lechler 99 (PC); headwaters of Mullet Creek Stream, ca. 60 m (3160, 3164, 3168-3169); Murrell River, 6 Nov. 1907, Halle (s, UPS); Mt. William, Lechler 99 (NY); Moody Brook, near Stanley, 8 m, Long- ton & Smith, Longton 961 (AAS, F). Mt. Usborne Region: valley SW of Mt. Usborne, ca. 60 m (2647, 2651, 2652B, 2656). WEST FALKLANDS. Mt. Adam: basin E of summit, 580-595 m (2999). Westpoint Island: 8 Oct. 1907, Skottsberg 351, syntype of Lophocolea falklandica (UPS); near The Waterfall, 30-90 m (2864, 2870-2873, 2877). Weddell Island: Waterfall Valley, W of settle- ment, ca. 125 m (3311, 3314, 3317, 3331-3332). Chiloscyphus bispinosus (Hook. f. & Tayl.) Engel & Schust. Jungermannia bispinosa Hook. f. & Tayl., London J. Hot. 3: 378. 1844. Lophocolea bispinosa (Hook, f. & Tayl.) G. L. & N., Syn. Hep. 162. 1845. Chiloscyphus bispinosus (Hook. f. & Tayl.) Engel & Schust., Nova Hedwigia 39: 41 1. 1984. Orig- inal material: Campbell I., Hooker (non vidi). ECOLOGY -Collected only once in the Falk- lands, on soil over rock in a dwarf shrub heath. PH YTOGEOGRAPH Y — Amphi pacific Temper- ate -Macquarie I.; Campbell I.; New Zealand; Tasmania; Australia (Victoria); Falkland Is.; Isles of Scilly (Paton, 1974); Isle of Colonsay, Scotland (Wallace, 1979). Reported from southern South America by Massalongo (1885, 1927); these, how- ever, are erroneous, for they are misdetermina- tions of Chiloscyphus leptanthus (fide NY!, VER!). FALKLAND SPECIMEN SEEN— WEST FALKLANDS. Weddell Island: summit of Mt. Weddell, 385 m (3347). Chiloscyphus divaricatus (Hook. f. & Tayl.) Engel & Schust. Lophocolea divaricata Hook. f. & Tayl., London J. Bot. 5: 367. 1846, non L. divaricata Herz., Arch. Esc. Farm. Fac. Ci. Med. Cordoba 7: 19. 1938. Jungermannia divaricata (Hook. f. & Tayl.) Hook, f. & Tayl. in Hooker f., Bot. Antarc. Voyage 1(2): 437. 1847, non J. divaricata Sm. in Sowerby, En- glish Bot. 10: 719. 1800 (= Cephaloziella), necJ. divaricata Nees, Enum. Plant Crypt. Javae 60. 1830 (cf. Acromastigum). Chiloscyphus divari- catus (Hook. f. & Tayl.) Engel & Schust., Nova Hedwigia 39: 414. 1984. Original material: Chile, Prov. Magallanes, I. Hermite, Hooker (NY!). ECOLOGY— This species is rare not only in the Falklands, but also in southern South America. Falkland plants are restricted to tussock associa- tions, where the species occurs on moist litter on the sides or bases of tussock (Poa flabellata) ped- estals, or between the pedestals. It usually occurs in admixtures with other hepatics, such as Chi- loscyphus lentus or Chiloscyphus sabuletorum. PHYTOGEOGRAPHY— Valdivian + Magellanian + Falklands— Falkland Is.; Tierra del Fuego; Pat- agonian Channels (Brunswick Peninsula); and Valdivian region (45°17'S, 73°43'W). Within the Falklands, it was found only in the Port Stanley region, and Beauchene Is. (52°54'S, 59°09'W). LITERATURE RECORD (FALKLANDS) — Smith, Beauchene Is. (Smith & Prince, 1985). FALKLAND SPECIMENS SEEN— Beauchene Is.: "The Hole," 25 m, Smith 3018, 3019d(AAS, F); above "Blinn's Cove," SE side of island, 10-20 m, Smith 3016 (AAS, F). EAST FALKLANDS. Kidney Island: north of shanty (2788B). Chiloscyphus elatus (Gott.) Engel & Schust. Figure 53. Jungermannia elata Gott., Ergebn. Deutsch. Polar- Exped. 2(16): 450.pl. 7,f. 3-6. 1890. Lophocolea elata (Gott.) Steph., Wiss. Ergebn. Schwed. Siid- polar-Exped. 4(1): 7. 1905. Chiloscyphus elatus (Gott.) Engel & Schust., Nova Hedwigia 39: 414. 1 984. Original material: South Georgia, 1 882, Will (M), cited in Grolle (1962) (non vidi). Leioscyphus set istipus Steph., Bih. Kongl. Svenska Ve- tenskapsakad. Handl. 26 (III, 17): 16. 1901, syn. fide Grolle (1962). Leptoscyphus setistipus (Steph.) Kiihnem., Revista Centra Estud. Doct. Ci. Nat., Buenos Aires 1: 177. 1937. Mylia setistipa (Steph.) Kuhnem., Lilloa 19: 341. 1949. Original mate- rial: Chile, Prov. Magallanes, R. Azopardo, 1 896, Dusen 56 (G, s), cited in Grolle (1962) (non vidi). ECOLOGY— This species seems to be restricted to stream habitats in dwarf shrub heath and Cor- 114 FIELDIANA: BOTANY taderia associations, where it was, without excep- tion, submerged or nearly so. This species, with C. austrigenus, is characteristic of the submerged stream bryoflora. PHYTOGEOGRAPHY— Magellanian-Falkland — South Georgia; Falkland Is.; Fuegia (Isla Grande de Tierra del Fuego); southern Patagonian Chan- nels (Brunswick Peninsula). The report from Gru- po Evangelistas in Stephani (1911) requires in- vestigation (see fig. 53). LITERATURE RECORD (FALKLANDS)— Skottsberg, (?)Eastpoint Is. (Grolle, 1962). FALKLAND SPECIMENS SEEN -EAST FALKLANDS. Lyall, as Jungermannia vasculosa (FH). Mt. Usborne Re- gion: valley SW of Mt. Usborne, ca. 60 m (2667). WEST FALKLANDS. Fox Bay Region: near mouth of Cheek's Creek, ca. 12 m (3496A); near peat cuttings in Ram Paddock, 45-60 m (3518); valley E of Sulivan House, ca. 75 m (3524, 3528). Westpoint Island: Skottsberg as L. falklandica (o, UPS); near The Waterfall, 39-90 m (2559). Chiloscyphus hookeri Engel PHYTOGEOGRAPHY— Magellanian-Falkland — Falkland Is., Tierra del Fuego, and Patagonian Channels north to 49°02'S. REMARKS— Fulford (1976) placed Chiloscyphus hookeri in the synonymy of Leptoscyphus palli- dovirens, regarding the two taxa as "two extremes of another variable species of Leptoscyphus" where she transferred the species. Over the last decade I have studied many collections of both species, and regard the two taxa as quite distinct entities which do not even approach one another. See Engel ( 1 972b, 1978) for key characters which I have used to differentiate the species. I have outlined reasons for classifying C. hookeri in Chiloscyphus (see En- gel, 1972b,p. 154; see also Engel & Schuster, 1984, p. 409). This species has two varieties (see Engel, 1 972b); only the typical variety occurs in the Falklands. It differs from var. constant ifolius Engel in the vari- able leaf apices— broadly rounded to truncate to retuse to single lobed to bifid, even on a single axis. Chiloscyphus hookeri Engel var. hookeri Chiloscyphus hookeri Engel var. hookeri. J. Hattori Bot. Lab. 36: I50.pl. 12. 1973. Holotype— Chile, Prov. Magallanes, I. Hermite, Hooker 12, mixed with syntype plants of Junger- mannia pallido-virens (NY!). ECOLOGY— Rather common in dwarf shrub heaths, especially over 155 m, where it grows un- der rock overhang or Blechnum magellanicum cover in deep rock crevices, etc., more rarely on stream banks in this association, and less common under grass cover in Cortaderia heaths. FALKLAND SPECIMENS SEEN -EAST FALKLANDS. Port William Region: summit ridge of Mt. Low, ca. 245 m (3227-3228); ridge on Engineer Point Peninsula, 1 8- 25 m (2820— c. per.). Stanley Region: S slopes of Mt. William, 185 m, Longton & Smith, Longton 984 (AAS, F); Tumbledown Mt., 155-230 m (2372B-C. 9, 2386); Goat Ridge, ca. 180 m (3197— c. per., 3209-c. per.); summit ridge of N peak of Two Sisters, 245-290 m (2704. 2710C-C. per.). Mt. Usborne Region: The Gap, 275-290 m (2462A-C. S); S side of The Gap, 260-275 m (2454 -c. per.). WEST FALKLANDS. Port Howard: Freezer Rocks, on E slope of Mt. Maria, 320 m (3137). Hornby Mts: 19 Dec. 1907, Halle & Skottsberg 223 as Lophocolea pallido-virens (UPS). Fox Bay Region, sum- mit of East Head, 180 m (3437); near peat cutting in Ram Poddock, 45-60 m (3511-c. S). Mt. Adam, 13 Dec. 1907, Halle & Skottsberg 223 as Lophocolea pal- lido-virens (UPS). Weddell Island: summit of peak NE of Mt. Weddell, 335 m (3369E); summit of Mt. Weddell, 380 m (3334. 3337-c. a, 3343A). Chiloscyphus humifusus (Hook. f. & Tayl.) Engel & Schust. Jungermannia humifusa Hook. f. & Tayl., London J. Bot. 3: 472. 1 844. Lophocolea humifusa (Hook, f. & Tayl.) G. L. & N., Syn. Hep. 695. 1847. Chiloscyphus humifusus (Hook. f. & Tayl.) Engel & Schust., Nova Hedwigia 39: 416. 1984. Orig- inal material: Kerguelen Is., Hooker (NY!; s!, ex "Herbarium Lehmannianum"). REMARKS— This is a distinctive species which may be readily identified by the following features: 1) variable leaf lobes: one or both are commonly rather broadly rounded (often with one lobe rounded and the other triangular), or both are nar- rowly triangular; 2) leaf lobe asymmetry: the ven- tral lobe is usually larger and the dorsal is often reduced to a low, rounded projection or is absent altogether (see Taylor & Hooker, 1847, pi. 159, f. 5); in some populations, however, the ventral lobe is often smaller than the dorsal; 3) underleaves bifid nearly to the base, with the lamina between sinus base and underleaf base only a few cells high; 4) underleaf lobes setaceous, sometimes quite markedly so; 5) trigones distinct, medium and straight-sided to large and slightly bulging; and 6) underleaf lamina with a pronounced lacinium or ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 115 cilium. The last is a curious appendage that varies from erect to spreading. At times its attachment to the underleaf is quite subtle, and it superficially appears to insert on the stem. I have not seen "stipules sometimes with a setaceous process at each side running back from their insertion along the stems" as described by Hooker and Taylor (1844) in the protologue. However, I have ob- served (in the type) long-setaceous underleaf seg- ments that extend distally to the basal-lateral por- tion of the next underleaf above, and superficially appear to originate there. It might possibly be this condition that was the basis for Taylor and Hook- er's figures and mention of underleaf processes. It is doubtful that they are rhizoids, for Taylor and Hooker ( 1 847) illustrate them as multicellular. Intercalary branches in Engel 3382 are plastic in origin and may be ventral, ventral-lateral, or lateral. Lateral branches may issue from above or below the leaf insertion and near the median por- tion of the leaf or near its ventral base. They may often be found between the underleaf base and the ventral base of the leaf immediately basal to it. Most intercalary branches remain dormant and "budlike"; some elongate slightly, and are very narrow compared to the main axis. I know of no other southern South American Chiloscyphus with intercalary branches of this kind. Frullania-type branches are also common. Fulford (1976) placed the species as a synonym of Lophocolea leptantha. ECOLOGY— Rather rare in Falklands; my collec- tions are from rocky outcrops above 335m where over rock or on soil under partial rock cover. The Longton and Smith plant was collected at a much lower elevation (30 m) on moist sandy sheltered vertical north-facing rocks. PHYTOGEOGRAPHY — Subantarctic — Kerguelen Is.; Falkland Is. This is the first authentic record of this species from the Falkland Islands. Stephani (1911) reported Chiloscyphus humifusus from the Falklands, and later Kuhnemann (1937, 1949) in- cluded this reference in his catalogs. The reports are in error, as the specimen is a misdetermination of Leptoscyphus patagonicus. The first of the two reports of the taxon in Stephani (191 1) is a Valle Frias collection, which is a misdetermination of Clasmatocolea humilis. Further, in the Uppsala Herbarium, the following specimens are misde- termined by Stephani as L. humifusa: 1896, Dusen 31 from Punta Arenas (a specimen of Clasmato- colea rigens mixed with Chiloscyphus leptanthus) and 1896, Dusen 304 from Ushuaia (a specimen of Chiloscyphus textilis). All remaining records of the species from southern South America require confirmation. FALKLAND SPECIMENS SEEN— EAST FALKLANDS. Stanley Region: S. shore of Murrell River, opposite Islet Point, 30 m, Longton & Smith, Longton 1026 (AAS, F). WEST FALKLANDS. Port Howard: pass SW of Mt. Maria summit, ca. 610 m (3081, 3105, 3125). Weddell Island: summit of peak NE of Mt. Weddell, 335m (3382, 3398). Chiloscyphus lentus (Hook. f. & Tayl.) Engel & Schust. Jungermannia lenta Hook. f. & Tayl., London J. Bot. 3: 379. 1 844. Lophocolea lenta (Hook. f. & Tayl.) G. L. & N., Syn. Hep. 162. 1845. Chiloscyphus lentus (Hook. f. & Tayl.) Engel & Schust., Nova Hedwigia 39: 418. 1984. Original material: Auck- land Is., Hooker (non vidi). Jungermannia secundifolia Hook. f. & Tayl., London J. Bot. 3:471. lS44,syn.fideMillen(\&55). Loph- ocolea secundifolia (Hook. f. & Tayl.) G. L. & N., Syn. Hep. 693. 1 847. Leclolype (fide Engel, 1978): Falkland Is., Hooker (FH!). REMARKS— See Engel (1978). ECOLOGY— This species is common on bases of tussock in the tussock association. It is also present on outcrops in dwarf shrub heaths where it may be found on soil under rock overhangs and in crev- ices. It was also found on bark of Hebe, Notho- fagus, and fir (the latter two in the grove of planted trees at Hill Cove settlement). PHYTOGEOGRAPHY — Amphipacific — Reported from South Sandwich Is. and South Georgia (Grolle, 1972b; Longton & Holdgate, 1979, both as L. secundifolia); Falkland Is.; Tierra del Fuego; southern Patagonian Channels (Brunswick Pen- insula); the Valdivian region in the American sec- tor; Macquarie, Auckland, Campbell, and Snares Is., New Zealand, Tasmania, and Australia in the New Zealand sector. LITERATURE RECORDS (FALKLANDS)— Anony- mous (Kuhnemann, 1937, 1949, as L. secundifo- lia); Hooker (G. L. & N., 1847, as L. secundifolia; Taylor & Hooker, 1847, as /. secundifolia; Ste- phani, 1906, as L. secundifolia); Skottsberg, Port Stanley, Port Louis (Stephani, 1905a, as L. secun- difolia), Port Stanley (Skottsberg, 1913, as L. se- cundifolia); Skottsberg & Halle, Port Louis, Half- way Cove (Stephani, 1911, as L. secundifolia); Smith, Beauchene Is. (Smith & Prince, 1985). FALKLAND SPECIMENS SEEN— Beauchene Is.: Above Blinn's Cove, SE side of island, 10-20 m., Smith 3015 (AAS, F). EAST FALKLANDS. 1843, Hooker, syntypes 116 FIELDIANA: BOTANY of Jungermannia secundifolia(BM, NY— c. per. + <5); Lyall as L. secundifolia (FH). KIDNEY ISLAND, 1954, Pet- tingill 16 (CMC, F), 77 (CINC); N of shanty (27 88 A, 2792. 2795. 2796. 2797, 2802-c. per. + a). Port William Region: S shore of Cape Pembroke Peninsula, near Surf Bay, sea level (2432. 2434— c. per. + S); ridge on En- gineer Point Peninsula, 18-25 m (2837— c. 3 + young 9). Darwin Settlement: Boca House on Brenton Lock, sea level (2689-c. sporo. + S). WEST FALKLANDS. Fox Bay Region: summit of East Head, 1 80 m (3438A — c. sporo., 3442— c. per. + 77../Kfc Fulford ( 1976). Loph- ocolea alternifolia (Hook. f. & Tayl.) G. L. & N., Syn. Hep. 695. 1847. Lectotype (/kfc Engel, 1978): Falkland Is. (non New Zealand), Lyall s.n. (FH! — c. sporo.). Lophocolea gibbosa Mont., Ann. Sci. Nat. Bot. Ill, 4: 351. 1845, syn.fide Fulford (1976). Original ma- terial: Southern Chile, without specific locality, Gay (PC!— c. per.). Lophocolea cunninghamiiSleph., Bull. Herb. Boissier 6(8): 652. 1906 (= Spec. Hep. 3: 68), syn. fide Fulford (1976). Original material: Chile, Prov. Magallanes, Pto. Eden, ex hb. Kew, Apr. 1868, Cunningham 56 (G!). Lophocolea monoica Steph., Kongl. Svenska Vetensk- apsakad. Handl. 46(9): 48. / 19k-p. 1911. syn. fide Fulford (1976). Lectotype (fide Engel, 1978): Falkland Is., Port Stanley, Sapper Hill, 29 Oct. 1907, Skottsberg s.n. (UPS!— c. per.). REMARKS— See remarks in Engel (1978). The type locality of Lophocolea alternifolia has been confused. The original description cites New Zealand as the type locality. A single sheet in the Taylor herbarium (FH) has the following three sep- arate specimens: a) a packet labeled "Falkland Is- lands" but crossed out with "New Zealand" writ- ten below, collected by Lyall; b) a small packet along with a glued specimen labeled "number 1"; and c) a portion of b which has both been placed in a small packet and glued to the sheet and labeled "number 2." Taylor has noted below the speci- mens that number 1 is from the Falkland Islands and collected by Lyall and was communicated by J. D. Hooker, 1844 (the following note is also in- cluded here "with Chilosc. retusatus Tayl."), and number 2 is from New Zealand, also collected by W. Lyall and communicated by W. Wilson. Taylor and Hooker ( 1 847, p. 436) stated of Jun- germannia alternifolia: "Hab. Falkland Islands" and "This species was erroneously described (Lon- don J. Bot. I.e.) as a native of New Zealand; from whence we have never seen specimens." I believe that specimens labeled numbers 1 and 2 are merely segregates of a single Falkland Islands-Lyall col- lection, as both contain the following identical complex of plants: Chiloscyphus leptanthus, a rath- er flaccid form of Leptoscyphus expansus with truncate leaf apices which are frequently emargin- ate, a moss, and bits of grass. In addition, I have studied a syntype collection from the Hooker Herbarium of the British Museum, and this Lyall- collected specimen bears the Falkland Islands locality only. This specimen also contains, in ad- dition to Chiloscyphus leptanthus, the Leptoscy- phus expansus, a moss, and a grass. Specimen a above, which is only a small segregate, has the moss but not Leptoscyphus expansus or the grass. The confusion surrounding this species likely orig- inated during curating of the specimen after it left Hooker and before it was studied by Taylor. I consider the three Farlow plants to be one collection that has been subdivided, and since two of the specimens have perianths, one with a cap- sule, I have designated these specimens as the lec- totype. The British Museum specimen (syntype) appears sterile. The description and figures of Lophocolea mo- noica in Stephani (1911) are somewhat confusing. Part of this confusion arises out of the heteroge- neous assembly of syntype material. The syntypes from Territory Chubut (Arroyo Carbon), and Azo- pardo-Tal are misdeterminations of Clasmatoco- lea rigens. The syntypes from Isla Dawson, Falk- land Islands (Rabbit Island) and Falkland Islands (Sapper Hill) are plants of Lophocolea monoica. The figures and descriptions of vegetative material are of L. monoica, while those of the perianth clearly do not correspond to the long, narrow peri- anths normally produced by this taxon. The peri- anth drawn and described may be that of a very immature one, or one of C. rigens. Stephani (Spec. Hep., Vol. 6, p. 283) stated for L. monoica that plants are ""dilute brunnea" and revised the description of the perianth for the species, stating ""Perianthia anguste oblonga (7 mm ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 117 54 FIG. 54. Distribution of Chiloscyphus leptanthus (Hook. f. & Tayl.) Engel & Schust. Open circle indicates precise locality unknown. longa, 2.5 mm laid) apice ad V3 triloba, lobis late ligulatis, irregulariter multispinosis" The descrip- tion of the perianth here fits that of L. monoica, i.e., it corresponds to the vegetative material de- scribed and drawn in 1911. Further, in 1922 Ste- phani stated the species was collected in the Falk- land Islands by Skottsberg. Thus, Stephani in the latter reference has excluded South American col- lections. Of the syntypes from the Falklands, that from Sapper Hill has several plants with leaves concave and decurved, and thus best agrees with the description. The UPS collection by Skottsberg s.n. (29 Oct. 1927) from the Falkland Islands (Sap- per Hill) was designated the lectotype of Lopho- colea monoica in Engel (1978). The material of L. monoica agrees well with L. leptantha and is here treated as a synonym. Fulford (1976) considerably extended the dis- tribution of this species by including, as synonyms, C. humifusus from Kerguelen and three taxa from Brazil. I accept C. humifusus as a distinct species, and question the Brazil names as synonyms. For the present I am restricting C. leptanthus to those regions indicated below. ECOLOGY— This taxon is rather rare in the Falk- lands, occurring in wet depressions, on wet stones partially submerged in streams, and on soil of dwarf shrub heaths. The species was also found on the bases of and in between tussock plants in a tussock association. On Beauchene Is. the species occurs on moist tussock litter between Poajlabellata ped- estals, and label data accompanying the Smith col- lection notes that it is "widespread in small amounts— the only 'abundant' bryophyte on the island". It also occurs in the grove of planted trees (mainly hardwood) at Hill Cove settlement and on the floor of the coniferous plantation at Gov- ernment House in Stanley. See comments in Engel (1978) for the ecology of the species in the Bruns- wick Peninsula, where it is rather common. PHYTOGEOGRAPHY— Valdivian + Magellanian + Falklands— Falkland Is.; Tierra del Fuego; Pat- agonian Channels; the Valdivian region (West Pat- agonia to 43°54'S). I have not seen the specimen on which the Juan Fernandez report is based, and its presence there is to be regarded as questionable. There are no authentic reports of this species from Andean Patagonia; the report of L. monoica from Arroyo Carbon in Stephani (191 1) is a misdeter- mination of Clasmatocolea rigens (see fig. 54). LITERATURE RECORDS (FALKLANDS)— Anony- mous (Kuhnemann, 1937, 1949 as L. cunning- hamii & L. monoica); Hooker (Taylor & Hooker, 1847, as Jungermannia alternifolid); Pettingill— Charles Point (Fulford, 1976); Skottsberg (Ste- phani, 1 922 as L. monoica), Port Stanley, Rabbit Is. (Skottsberg, 1913, as L. monoica), Port Louis (Skottsberg, 1913; Stephani, 1905a, both as L. cunninghamii); Smith, Beauchene Is. (Smith & Prince, 1985). FALKLAND SPECIMENS SEEN— Beauchene Is.: "The Hole," 25 m, Smith J079c(AAS); ca. 25 m to N of fresh- water pool at SW end of island, Smith 3014 (AAS, F). EAST FALKLANDS. Lyall, syntype of Jungermannia alternifolia (BM). Kidney Island: N of shanty (2797, 2801 — c. per. -I- 3). Port William Region: Charles Point, 1954, Pettingill 4 (CINC, F); N side of Gypsy Cove, sea level (3253); ridge on Engineer Point Peninsula, 1 8-25 m (2814, 2816, 2824— c. per. + <3, 2835 -c. per. + $). Stanley Region: Port Stanley, Government House, 1 5 m, Long- ton & Smith, Longton 1032 (AAS); Port Stanley, 8 Nov. 1902, Skottsberg s.n. as L. cunninghamii (UPS— c. per.); Moody Brook, 8 m, Longton & Smith, Longton 963 (AAS). Darwin Settlement: S side of Carcass Bay, Darwin Harbour (2682-c. per. + $). WEST FALKLANDS. Hill Cove Region: settlement, 15 m (3052— c. per. +