Botany
Falkland Islands (Islas Malvinas) Hepaticae and Anthocerotophyta: A Taxonomic and Phytogeographic Study
John J. Engcl
Sov< nber 30, 1990 Publication 1416
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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 |
|
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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
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ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA
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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
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14
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ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA
15
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ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA
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ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA
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ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA
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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-
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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-
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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-
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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
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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
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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:
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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