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).

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

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FIELDIANA: BOTANY

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

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FIELDIANA: BOTANY

FIG. 7. Map of personal collection localities. The numbers indicate regions or islands, which are as follows: 1)

Kidney Is., 2) Port William (includes Mt. Low), 3) Port Stanley (includes Mt. Kent), 4) Mt. Usborne, 5) Darwin

Settlement, 6) Port Howard, 7) Fox Bay, 8) Mt. Adam, 9) Hill Cove, 10) Westpoint Is., 1 1) Weddell Is., 12) New Is.

The average annual duration of sunshine is less

than 35% of the total possible for the latitude of

Port Stanley (Cawkell et al., 1960).

Collectors of Hepaticae and

Anthocerotophyta

The most significant collections of Hepaticae

and Anthocerotophyta are those of Hooker,

Skottsberg, and Halle, while those of Lechler and

Cunningham are important but fewer in number.

Table 2 shows a chronological list of collectors,

together with pertinent data.

A list of my collection numbers together with

collection data may be found in the Appendix (p.

1 99). A statement of the ecology is made for nearly

every locality, since ecological data have been

omitted under the specimen citation portion of

each species. The localities are indicated on Figure

7. All localities are cited with reference to Uni-

versal Transverse Mercater Grid (Zone 2 1 ) coor-

dinates as shown on maps of Falkland Islands

1:50,000 (Directorate of Overseas Surveys 453,

Series H791, 1961).

In the Systematic Account, the specimens I have

personally collected are indicated by number only,

without citation of collector. A complete set of my

collections has been deposited in the Cryptogamic

Herbarium, Michigan State University (MSC). All

other specimens are cited with the collector's name

provided.

Phytosociology

Introduction

Skottsberg (1913) made a vegetational analysis

of the vascular plants of the Falkland Islands, and

Moore (1968) based his account of the Falkland

vegetation primarily on that of Skottsberg. Brief

accounts of the natural vegetation are also pro-

vided in Davies (1939) and Cawkell et al. (1960).

In summarizing the bryophyte sociology in the

Falkland Islands, I have utilized the community

groupings and terminology of Moore ( 1 968). I have,

however, added one community grouping, i.e., the

sheltered high altitude cliff vegetation, which is

primarily cryptogamic. As emphasized by Moore

the predominant Falkland plant associations are

closely related to one another. The associations

and formations, especially those of lowland areas,

frequently are not arranged as discrete units, but

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

13

14

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16

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ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

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FIELDIANA: BOTANY

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

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FIELDIANA: BOTANY

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

22

FIELDIANA: BOTANY

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23

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25

26

FIELDIANA: BOTANY

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

21

a kind of mosaic of community groupings is rather

the pattern.

Moore (1968) has recognized five formations.

In addition, he has recognized four associations

(Senecio candicans, Ammophila-Elymus, Eleo-

charis, and Myriophylluni) which do not merit for-

mation grouping. These are arranged according to

types of vegetation.

Distribution of Bryophytes in Plant

Communities of the Falkland Islands

Distribution of bryophytes in Falkland Islands

plant communities is shown in Table 3 (pp. 14-

26).

MARITIME TUSSOCK FORMATION— Poa flabel-

lata Association— This primarily coastal associa-

tion was previously far more prevalent than that

observed today (Hooker, 1 847; Moore, 1 968). The

unrestricted, extensive grazing of sheep has been

primarily responsible for the depletion of this for-

mation. Pigs, which destroy plants by digging up

the roots, are of lesser significance. Tussock grass

today is confined to smaller offshore islets (e.g.,

Kidney Island) which have not been stocked with

sheep, and a few fenced off areas on the larger

islands. On Kidney Island, where there are exten-

sive areas of almost pure stands of Poa flabellata,

this association has reached its maximum devel-

opment (fig. 8). As discussed in Moore ( 1 968), Poa

flabellata tussocks can attain a height of 2-3 m

with each tussock composed of a fibrous stock

(1.5-2 m high, 1-1.5 m in diameter) surmounted

by a dense crown of leaves. In areas where the

community is uninterrupted, the dense canopy of

interlaced leaves excludes all associated vascular

plant species except Carex trifida (fig. 9). Telara-

nea pseudozoopsis and Chiloscyphus lentus were

particularly common on the tussock bases while

C. leptanthus, C. sabuletorum, and Hygroambly-

stegiumfuegianum var. skottsbergii were found on

soil between the tussock bases. Poa flabellata is

well-developed and forms a monospecific grass-

land covering two-thirds of Beauchene Island, the

most isolated island of the archipelago (lying 80

km south of East Falkland Island) (Smith & Prince,

1985). Chiloscyphus lentus was found to be the

most frequently occurring hepatic species on the

island; it "grew around the moist shaded bases of

occasional tussock pedestals and on inter-tussock

mounds . . ." (Smith & Prince, 1985, p. 242). Oc-

casional associates were Chiloscyphus divaricatus,

Clasmatocolea vermicularis, and Telaranea pseu-

dozoopsis.

OCEANIC HEATH FORMATION— This formation

has been subdivided into two groups. Cortaderia

pilosa is dominant or codominant in one, and dwarl

shrubs are prominent in the other. Most of the

Falkland Islands are covered by one facies or

another of these associations.

Cortaderia Association— This association, as

Moore (1968, p. 15) stated, "can be developed on

most nonswampy ground having indifferent drain-

age. It is widespread on level or undulating country

below 100 m ... but it is also very common on

gentler slopes up to an altitude of ca. 180-200 m"

(fig. 10). I have found this association, which is

dominated by Cortaderia pilosa, occurring at var-

ious altitudes up to ca. 610 m. The grass blades

form an excellent protective cover against the sun

and especially the winds, while the coarse fibrous

peaty humus which develops under the tussocks

retains considerable moisture. These factors con-

tribute to a protected environment in which a sur-

prisingly large number of bryophyte species may

be found. Bryophytes directly associated with Cor-

taderia, in depressions, pockets, or holes, and de-

riving protection and moisture from the associa-

tion are listed in Table 3 under the General

Terrestrial column heading. This association in-

cludes several facies, each differing in the means

of providing protection and moisture conserva-

tion. These facies will be discussed separately.

The Stream Facies— Streams are frequently

present in the Cortaderia association (fig. 1 1); often

the bank vegetation (which frequently is Gunnera

magellanicd) or the stream itself offers protective

niches for bryophytes. Chiloscyphus austrigenus is

very common here and, together with C. elatus,

is the only hepatic that is nearly always submerged

in these streams.

The Wet Depression Facies— Wet depressions,

where the peat is saturated and small pools may

occur, provide another means of protection for

bryophytes (fig. 1 2).

The Stone Run Facies— Stone runs are of fre-

quent occurrence in the Cortaderia association (figs.

13-14). Often, bryophytes occur in cool, damp,

shaded crevices.

The Rock Outcrop Facies— Rock outcrops (par-

ticularly those having exposures protected from

prevailing winds) are the sites for 12 species of

bryophytes. These plants were found associated

with overhangs, crevices, and small niches, and

on the soil immediately below the outcrops.

The Cortaderia association, while not possess-

ing the bryophyte species diversity of the closely

related dwarf shrub heath, possesses the greatest

28

FIELDIANA: BOTANY

FIG. 8. Poa flabe llata associa-

tion at Kidney Island.

FIG. 9. Interlacing blades of

Poa flabellata at Kidney Island;

note head of standing individual

for scale. Photo by H. A. Imshaug.

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

29

30

FIELDIANA: BOTANY

Above: FIG. 13. Stone run as seen from air over East I . ilk lands

FIG. 14. Bolax gummifera in stone run.

Opposite: Fio. 10. Cortaderia association near Mt. Usborne caravan. Photo by R. C. Harris.

FIG. 1 1. Stream through Cortaderia association near Mt. Usborne caravan.

FIG. 1 2. Wet depression in Cortaderia association in gap between French Peaks.

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

31

17

32

FIELDIANA: BOTANY

assemblage of rare Falkland species, several of

which are also rare in southern South America.

Dwarf Shrub Heath Associations— Moore ( 1 968,

p. 1 5) stated that "The communities included here

are usually found on comparatively dry ground,

being best developed on rocky ridges, stony areas

or places where the immediate subsoil is of rela-

tively coarse material so that drainage is good. A

hard, dry peat underlies well-developed heath."

The principal vascular plant species in the dwarf

shrub heath are Empetrum rubrum, Pernettya

pumila, Blechnum magellanicum, B. penna- ma-

rina. Baccaris magellanica, and Bolax gummifera.

Various combinations of these may become co-

dominant, and Moore (1968) has listed the fol-

lowing associations: Empetrum rubrum-Blech-

num penna- marina, Empetrum rubrum- Blechnum

magellanicum, Empetrum rubrum- Pernettya

pumila, Bolax gummifera- Empetrum rubrum, and

Blechnum magellanicum- B. penna- marina. Moore

(1968, p. 15) stated "The Empetrum rubrum as-

sociation is much the most common" and the dis-

cussion below pertains to this association. As in

the Cortaderia association, there are several means

of protection against the sun and winds as well as

the conservation of adequate moisture so that this

association has a fairly rich bryophyte flora. Each

local facies will be discussed separately.

Bryophytes commonly grow on the bases of

Blechnum magellanicum, the fronds of which pro-

vide an excellent protective cover, particularly

when interlaced with fronds of other Blechnum

plants or branches of Empetrum (fig. 1 5). The soil

below such situations often has a considerable bry-

ophyte cover. Frequently, Bolax cushions, partic-

ularly old, decayed portions, have a bryophyte

(mostly hepatic) cover. The bryophytes in these

situations are listed in Table 3 in the dwarf shrub

heath column labeled General Terrestrial.

The Stream Facies— Streams often occur in the

dwarf shrub heath association. Vascular plants such

as Blechnum magellanicum, B. penna-marina,

Empetrum rubrum, and Gunnera magellanica fre-

quently offer a protective cover, which may be

very dense, and at the same time provide cool,

moist niches for bryophytes (fig. 1 6). Several species

grow partially or wholly submerged in the streams,

Chiloscyphus austrigenus and C. elatus nearly ex-

clusively so. This facies, when compared to the

stream facies of the Cortaderia association, while

having the same Chiloscyphus species, has quite a

different species complement, as well as greater

species diversity.

The Wet Depression Facies— Like the Corta-

deria association, the Empetrum rubrum associ-

ation has shallow depressions where the peat is

saturated and small pools may occur. The habitat

has a characteristic bryophyte flora. Sphagnum

falcatum was present in depressions where free-

standing water occurred.

The Rock Outcrop Facies— Rock outcrops of

varying sizes are frequently present in this asso-

ciation, with protection provided in a manner sim-

ilar to that in the rock outcrop facies of the Cor-

taderia association. Bryophytes occur in the various

niches associated with the outcrops, as well as on

soil at the base of the outcrops.

The Hebe Facies— Hebe elliptica, when occur-

ring as scattered bushes, may be regarded as a

relatively minor component of a dwarf shrub heath

(Moore, 1968). A heath of this type was visited in

Waterfall Valley, Westpoint Island. The Hebe

shrubs occurred on steep slopes on the valley sides

(fig. 17), while through the valley bottom flowed

a rather fast-moving stream, with Gunnera ma-

gellanica forming a dense carpet along the water-

course. The sides of the valley were often clifflike,

with numerous damp, shaded, sheltered ledges and

overhangs. Bryophytes occurring under the wa-

terfall or in the spray zone were Megaceros fue-

giensis, Riccardia georgiensis, Telaranea pseudo-

zoopsis, and Webera albicans. The bryophytes

associated with the stream bed were typical of this

habitat and consisted of Chiloscyphus austrigenus,

C. elatus, Riccardia spectabilis, Telaranea plu-

mulosa, T. pseudozoopsis, and Sciaromium con-

spissatum.

Hebe elliptica also occurred on cliffs facing The

Woolly Gut on Westpoint Island. The cliffs were

rather moist where numerous crevices, ledges, and

overhangs were present. Three hepatics— Chilo-

scyphus semiteres, Megaceros fuegiensis, and

Metzgeria violacea— occurred exclusively in the

Hebe facies.

Summary of Dwarf Shrub Heath Hepaticae—

The dwarf shrub heath associations have the high-

est number of hepatic species of any of the Falk-

land associations (75), and of these 17 are unique

to it. Of those that are not restricted to this as-

sociation in the Falkland Islands, 49% also occur

Opposite: FIG. 15. Dwarf shrub heath association near Mt. Adam.

FIG. 16. Stream in dwarf shrub heath association near Weddell Island settlement.

FIG. 1 7. Hebe elliptica component of dwarf shrub heath association in Waterfall Valley, Westpoint

Island.

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

33

in the Cortaderia heaths, which is indicative of the

rather close relationship between the associations.

However, I regard the bryofloras of the Cortaderia

association and dwarf shrub heath associations as

sufficiently distinct, and the differences aid in dis-

tinguishing the two associations.

The dwarf shrub heath associations and Cor-

taderia associations are vegetation types which

have been highly altered both through grazing and

conversion to pastureland, a process which fre-

quently involves burning and usage of introduced

grasses (commonly Holcus lanatus; Davies, 1 939).

Even in the most severely altered state of the

oceanic heaths, I have found it possible to deter-

mine the natural vegetation type through exami-

nation of the bryoflora of stream banks. The stream

banks are somewhat less disturbed and here the

natural vegetation is least altered. See further com-

ments under Man-Influenced Communities, at the

end of Phytosociology.

FELDMARK FORMATION— The feldmark (= fell-

field) formation has been characterized by the fol-

lowing four features:

1. Considerable areas of bare, exposed, usually

mineral soils with scattered small rocks or

boulders. According to Greene (1964), feld-

marks have a vegetation cover of one half or

less the total area.

2. The tendency for the cushion habit to predom-

inate. The feldmark cushions do not form a

continuous layer of peat (Wace, 1965), and do

not have among them the cushion bog1 dom-

inants of other associations (in the Falklands

the Astelia association). Likewise Azorella, the

principal feldmark cushion species, does not

occur in cushion bogs (Wace, 1960). Wace

(1960, p. 486) listed the principal species for

feldmarks of the following islands:

Falkland Islands— Azorella selago, Bolax

gummifera

Kerguelen— Azorella selago, Colobanthus

kerguelensis, Lyallia kerguelensis

Macquarie— Azorella selago, Racomitrium

crispulum. Dicranoweissia antarctica

Tristan da Cunha— Empetrum rubrum, Ra-

comitrium crispulum, R. lanuginosum

Gough Island — Racomitrium crispulum,

Thysanomitrium richardii, Jamesoniella

spp.

1 The term cushion bog is used to indicate cushion

plant communities on deep peat; see Wace (1965) for a

discussion.

The cushion habit is predominant in all of the

above flowering plants. Bryophytes commonly as-

sume either a cushion or compact habit.

3. An often conspicuous cryptogamic component

of the vegetation. On the Falkland Islands the

lichen Neuropogon is a conspicuous element of

the feldmark vegetatiori. Mosses are conspic-

uous in Macquarie Island feldmarks, and Ash-

ton and Gill (1965) propose that the feldmark

community be termed the Azorella- Ditrichum

alliance. Ashton and Gill (1965) and Selkirk

and Seppelt ( 1 984) regard the Macquarie Island

feldmarks as a single variable vegetation unit.

Selkirk and Seppelt recorded 63 plant species,

71% of them cryptogams (29 bryophytes, 16

lichens, 1 8 vascular plants).

4. A preference for higher altitudes. Wace (1960)

lists feldmark altitudinal ranges for the follow-

ing islands: Falkland Islands (600-700 m),

Macquarie Island (200-450 m), Kerguelen Is-

lands (100-300 m), and Tristan da Cunha and

Gough islands (850-1 500 m). Wace ( 1 960) and

Wace and Dickson (1965) indicate the distri-

bution of feldmark is in large part determined

by exposure. Due to exposure factors in the

Falkland Islands, feldmarks occasionally occur

at lower altitudes in the western part of the

archipelago. Moore (1968, p. 16) stated that

"Apparently the formation is not solely a re-

sponse to high altitude, because a comparable

community type occurs among rock detritus

along exposed cliff tops on the west coast just

north of Cape Meredith . . ." (southernmost

point of West Falkland Island). On Westpoint

Island it is presumably exposure which results

in a feldmark at the summit of Mt. Misery (335

m). Feldmarks may also occur at rather low

altitudes on Macquarie Island, where they oc-

cur above 175 m (Selkirk & Seppelt, 1984).

The general aspect of Falkland feldmarks differs

considerably. While there may be a continuum

present, there appear to be two general types of

feldmarks.

Mesic Feldmark— This type is characterized by

a rather conspicuous fruticose lichen flora (prin-

cipally Neuropogon). The lichen flora composition

is apparently a result of the presence of mists and

frequent enshrouding clouds. The feldmark com-

munity on the summit of Mt. Usborne 1 (ca. 700

m) had a rather lush bryophyte vegetation due

presumably to the numerous rather small scattered

pools which are present. Certain bryophytes were

associated solely with the pools, either submerged,

at the edge of, or on rock in the pools. The feld-

34

FIELDIANA: BOTANY

FIG. 18. Dry feldmark on ridge of north slope of Mt. Fegen, ca. 275 m.

mark community on the summit of Mt. Maria

(658 m) is rather xeric and has a conspicuous li-

chen cover, but only a single bryophyte (Riccardia

opuntiiformis).

While Adelanthus integerrimus is the only he-

patic found exclusively in this formation, it is of

interest to note that the following hepatics are found

exclusively in the Mt. Usborne feldmark and shel-

tered high altitude cliff vegetation: Acrobolbus och-

rophyllus, Anastrophyllum ciliatum, Archeochaete

kuehnemannii, Herzogobryum vermiculare,

Pachyglossa dissitifolia, and P. fissa.

Xeric Feldmark— This type is characterized by

a comparatively poor lichen cover (Neuropogon is

absent). The moisture available to this feldmark

type is derived solely from rain and not from en-

shrouding clouds, and exposure is likely the factor

which governs the comparative lack of moisture.

Two feldmarks of this type were visited: the north-

west tip of New Island opposite Landsend Bluff

(90 m) and the north slope of Mt. Fegen (ca. 275

m)(fig. 18).

Very characteristic of the bryophytes of this for-

mation are the following three mosses, all of which

grew on exposed rock surfaces: Andreaea pseu-

domutabilis, Racomitrium lanuginosum and R.

rupestre. On soil in crevices under rock overhangs

are Lepidozia chordulifera and Ditrichum sp.;

Bryum argenteum may be found on Bolax cush-

ions. This feldmark type supports only a very small

bryophyte vegetation. Even the rock crevices,

which often are relatively moist in the Falkland

Islands, were dry and only Lepidozia chordulifera,

which is able to grow in a wide range of habitats,

was apparently able to tolerate the dryness.

FEN AND BOG FORMATION— This formation oc-

curs where the water table is just below, at, or

above the ground surface.

Rostkovia Association— Moore (1968, p. 17)

stated, "This [association] is normally confined to

the wettest depressions, where drainage is imped-

ed, and standing water is often present for long

periods." In this association Sphagnum falcatum

grew submerged or nearly so, with S. fimbriatum

also present. No hepatics were observed.

Astelia Association — Moore (1968, p. 17) stat-

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

35

19

FIG. 19. Sheltered high altitude cliffs (arrow) of Mt. Adam summit area. Photo by R. C. Harris.

ed, "This [association] is usually found over deep

peat and consists of a series of low flat cushions

or a dense carpet composed of a few species which

all are able to retain water in the dense covering

of old leaves and branches surrounding their

stems." These cushion bogs grade into the Rost-

kovia association and the wetter facies of the Cor-

taderia heath. The observed Astelia associations

which did not grade into the above associations

were dense, compact, and very hard, and no bryo-

phytes were observed growing here.

June us scheuzerioides Association— No visits

were made to this association.

BUSH FORMATION— This formation is of re-

stricted occurrence in the Falkland Islands, and

only two native shrubs contribute to associations

of this formation.

Chiliotrichum Association— This association

was not visited in the Falkland Islands. According

to Moore (1968) and Skottsberg (1913), it is best

developed along the fringes of streams and rivers.

Skottsberg (1913) listed two localities for this as-

sociation, both in western West Falkland.

Hebe Association— Hebe elliptica, which is re-

stricted to the West Falkland Islands, was ob-

served at Westpoint Island, but the plants were

present only as scattered individuals or small

thickets in a dwarf shrub heath (q.v.). When a

dense stand of Hebe elliptica occurs, a true Hebe

association is said to be present. Such an associ-

ation is very rare in the Falkland Islands, and was

not visited. A photograph of an example on Fox

Island is in Skottsberg (1913, pi. 13).

LITTORAL VEGETATION— The Senecio candicans

andAmmophila-Elymus associations belong here,

and both occur in sandy areas. The former asso-

ciation was visited on the south shore of Cape

Pembroke Peninsula, near Surf Bay (East Falk-

lands), and the latter at Gypsy Cove, Port William

region (East Falklands). No bryophytes were found

in either of these associations. It is likely that the

dune and sandy areas are too unstable for bryo-

phyte survival.

FRESHWATER VEGETATION— Moore (1968) in-

cludes the Eleocharis and Myriophyllum associa-

tions in this category. No bryophytes were ob-

served in either of these associations. The

bryophyte collections which were frequently gath-

ered on the banks of shallow lagoons and ponds

are included in the association of the surrounding

area.

SHELTERED HIGH-ALTITUDE CLIFF VEGETA-

36

FIELDIANA: BOTANY

FIG. 20. Grove of planted trees (mainly Populus, Nothofagus, and conifers) at Hill Cove settlement.

TION— These communities consist principally of

bryophytes, and occur above 610 m in the Falk-

lands (fig. 1 9). They possess a highly characteristic

assemblage of hepatics, and because of their re-

striction to high altitudes I believe they deserve

separate recognition. Cliffs on ridgetops have nu-

merous protective niches such as ledges and rock

crevices, the latter frequently beneath overhangs

of varying sizes. The crevices, especially when be-

neath ledges, are commonly moist, cool, and well

shaded, a situation highly favorable to bryophyte

growth. When this is accompanied by water seep-

age, the bryophytes are particularly luxuriant. The

variation in exposures, moisture, light, and the

consistently rather cool temperatures probably ac-

counts for the considerable diversity of these com-

munities.

The following hepatics were encountered very

frequently and often were intermixed: Austrolo-

phozia fuegiensis, Balantiopsis bisbifida, Crypto-

chila spp., Herzogobryum teres, and Pachyglossa

spp.

MAN-INFLUENCED COMMUNITIES — Pasture-

land— In the Falkland Islands, there are wide ex-

panses of pastureland (see Davies, 1939) that are

regularly grazed by sheep. It is of interest to note

that all three of the mosses occurring here are quite

widespread and weedy, and it is not surprising that

all are rather plastic species that are able to tolerate

highly disturbed situations. Near the mouth of

Cheeks' Creek, Fox Bay region, a collection was

made in a stream cut through a pasture. It is prob-

able that before conversion to pastureland the

vegetation of the area was that of a dwarf shrub

heath association, as the bryoflora was similar to

the stream in that association, with the exception

of the presence here of Leptophyllopsis irregularis.

Groves of Planted Trees— A mature, well-de-

veloped grove occurs at the Hill Cove settlement.

According to Dallimore (1919), the trees were in-

troduced in the period 1889-1894. The hardwood

grove, which is made up of Nothofagus sp. and

Populus sp., has a rather dense canopy with little

woody undergrowth (figs. 20-21). The hepatics

found here were Chiloscyphus lentus, C. leptan-

thus, Leptoscyphus expansus, and Marchantia ber-

teroana var. polylepida; all occur in a variety of

natural communities. Adjacent to the hardwood

grove was a stand of conifers. No bryophytes were

observed in this stand or in the coniferous stand

on Weddel Island.

Gorse Thickets— Gorse (Ulex europaea), a na-

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

37

FIG. 2 1 . Interior of planted tree grove (mainly Nothofagus) at Hill Cove settlement.

live of western Europe, is widely utilized in the

Falkland Islands as sheep fencing. A gorse thicket

was visited at Boca House near Brenton Lock, near

Darwin Settlement, East Falklands. On soil be-

neath the gorse occurred Leptoscyphus expansus,

Brachythecium austrosalebrosum, Bryum argen-

teum, B. (sect. Rosulata) sp., and Tortula robusta.

Tortula monoica was the only bryophyte actually

growing on the gorse and it occurred on the ex-

posed roots.

Phytogeography

Austral Regions

Several attempts have been made since 1960 to

subdivide and classify austral zones. These clas-

sification schemes are based upon climatic and

biotic features, and also take into account the

dominant influence of oceanic circulation, namely

the position of the Antarctic and Subtropical Con-

vergences.

Wace (1960) devised a system based primarily

on lowland vegetation criteria. He combined all

those islands bearing oceanic heath and Sphag-

num or flowering plant cushion bogs into a tem-

perate category, and further divided this group

into warm and cool temperate, respectively, ac-

cording to the presence or absence of fern-bush

communities. He limited usage of the term "sub-

antarctic" to include only those islands with a

closed herbfield vegetation and in which Sphag-

num and cushion bogs are absent. The islands in

which there are no flowering plants (or only scat-

tered individuals), he considered as "antarctic."2

Wace (1960) compiled the available tempera-

ture data and noted that the groupings based on

lowland vegetation types correlated well with

groupings based on mean annual temperature. The

following is his classification scheme:

SUBTROPICAL Kermadec Islands

Juan Fernandez Islands

TEMPERATE Chatham Island

Tristan da Cunha, Inaccessi-

ble, Nightingale, and Gough

islands

2 It must be understood that islands, like Bounty Island

off New Zealand, may be without vascular plants as a

result of physiographic and biotic factors.

38

FIELDIANA: BOTANY

New Amsterdam and St. Paul

islands

New Zealand shelf islands

Falkland Islands

Marion and Prince Edward is-

lands

SUBANTARCTIC Macquarie Island

Crozet Islands

Kerguelen and off-lying is-

lands

South Georgia

Heard and Macdonald islands

ANTARCTIC Bouvet Island

South Sandwich, South Ork-

ney, and South Shetland is-

lands

Palmer Archipelago (Graham

Land)

Greene and Greene (1963) and Greene (1964)

adopted Wace's ( 1 960) concept of the term sub-

antarctic, but included Marion and Prince Edward

islands, and used the term antarctic region to em-

brace the antarctic and subantarctic zones (fig. 22).

This emended classification has been followed by

van Zinderen Bakker (1967, 1971) and Gremmen

(1982) for Marion and Prince Edward islands, by

Lindsay (1971) for the South Shetland Islands, by

Smith (1972) for the South Orkney Islands, and

by others.

Wace ( 1 965, p. 239) extended his earlier scheme

by stating, "In order to arrive at a vegetational

classification, the major features of the structure

of the plant community should be used exclusively

to delimit the different zones." He then proposed

to use the limit of tree or woody shrub growth to

separate the temperate from the subantarctic and

the limit of closed phanerogamic communities to

separate the subantarctic from the antarctic vege-

tation. He characterized the subantarctic vegeta-

tion zone as having terrestrial plant communities

with the following four features:

1. Soligenous mires in which the important peat-

forming plants are Bryales and Juncaceae (not

Sphagnum).

2. Feldmark communities composed of flowering

plants with very close-growing and compact

mat and cushion forms.

3. Closed herbfield communities in which large-

leaved perennial herbs are conspicuous.

4. Communities of large trunk-forming tussock

grasses around the coasts.

Wace excluded all continental areas and restricted

the usage of the term subantarctic to remote is-

lands. This zone included South Georgia, Marion,

Prince Edward (which in 1960 he listed as tem-

perate), Crozet, Kerguelen, Heard, and Macquarie

islands.

Holdgate's (1970) useful summary of the clas-

sification systems of Wace and Greene separates

subantarctic and south temperate as follows:

SOUTHERN COLD TEMPERATE ZONE— Botanical

Definition: From the Subtropical Convergence

southward to the southern limit of dwarf shrub

vegetation. Region: Western Magellanic Moorland

of Chile, Falkland Islands, Tristan-Gough groups,

I. St. Paul, Nouvelle Amsterdam, New Zealand

shelf islands (Auckland, Campbell, Snares, etc.).

SUBANTARCTIC ZONE — Botanical Definition:

From the southern limit of dwarf shrub vegetation

to the southern limit of extensive closed phanero-

gamic vegetation. Region: South Georgia, Marion

and Prince Edward, Crozet, Kerguelen, Heard,

Macquarie (all are insular and isolated).

Holdgate (1977) presented a more detailed clas-

sification based upon those of the earlier investi-

gators, but with the same circumscription of the

subantarctic. Smith (1984) proposed an improved

modification of the Holdgate (1977) classification,

and made several alterations, among them the

subdivision of the subantarctic into "three prov-

inces according to their oceanic affinities" (Smith,

1984, p. 64), namely a South Atlantic province

for South Georgia, a South Pacific province for

Macquarie Island, and a South Indian Ocean prov-

ince for the remaining islands. The subantarctic

botanical zone sensu Holdgate supports a vascular

plant flora of 72 species (cf. Greene & Walton,

1975; Smith, 1984), whereas the southern cold

temperate zone has much greater floristic diver-

sity.

It is evident from the criteria outlined in Wace

(I960, 1965) and Greene (1964) that the Falkland

Islands are south temperate rather than subant-

arctic. The characters which place them in the

south temperate region are a widespread oceanic

heath formation; the presence of flowering plant

cushion bogs; Sphagnum bogs; the presence of

woody shrub growth; and the absence of a closed

herbfield vegetation. The presence of a feldmark

community has been used as a characteristic of

the subantarctic region. In the Falkland Islands,

this community is present, but certainly does not

reach the significant proportions and role that it

does in the Kerguelen Islands.

Skottsberg (1960) based his austral phytogeo-

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

39

Wen of Greenwich 0° Eut of Greenwich

110° I3S8

FIG. 22. South polar projection showing delimitation of subantarctic region (after Greene, 1964).

graphic analysis on the following floristic zones

(five further provinces are included by Skottsberg,

some including "outposts" for "antarctic types"):

1 . Antarctic zone, south of 60°S, but including the

South Sandwich Islands and Bouvet Island

2. Subantarctic zone, between approximately 48°S

and 60°S, but including Marion, Prince Ed-

ward, and Crozet islands

a. Magellanic province

i. West Patagonian-Fuegian district

ii. Andean Patagonian-Fuegian district

iii. Falkland Islands and South Georgia dis-

trict

b. Kerguelen province. Marion, Prince Ed-

ward, Crozet, Kerguelen and Heard islands

c. Province of the subantarctic islands of New

Zealand

3. The austral zone, between approximately 40°S

and 48°S. A conspicuous antarctic floristic ele-

ment is present, often forming distinct com-

munities in a flora and vegetation of a different

origin.

a. Valdivian province

b. Province of the South Island of New Zea-

land, Chatham Islands, and the Tasmanian

tableland

Godley ( 1 960) adopted the subantarctic zone of

40

FIELDIANA: BOTANY

Skottsberg, but remarked (p. 472), "The presence

or absence of Sphagnum bogs appears to me to be

one of the significant features to be taken into

consideration when delimiting a homogeneous

subantarctic region." He regarded peat formation

as a common feature of the region, and the bogs

formed by cushion plants rather than by Sphagna.

Smith (1984, p. 64) rejects the subantarctic zone

as defined by Godley ( 1 960) and Skottsberg ( 1 960)

since "it includes several cool temperate islands

with arborescent plants and cushion bogs."

Bliss (1979) presents an alternative view and

develops a general classification system that em-

braces comparable Northern and Southern Hemi-

sphere vegetation. He combines alpine, arctic, and

antarctic vegetation into one classification system

and defines the southern lands as follows:

The Subantarctic as used here includes closed

and open forests and shrublands of Tierra del

Fuego, Falkland Islands, and New Zealand

shelf islands (Auckland and Campbell), Tris-

tan da Cunha-Gough Islands, and the tall tus-

sock grasslands of numerous islands. Antarc-

tic as used here includes closed vegetation of

grass-forb, herbfields, dwarf shrub, mires, and

the open cushion plant fellfields found at

higher elevations in some of the above islands

[italics mine], but vegetation more common

to the more southern islands (Macquarie,

South Shetlands, South Orkney, South Geor-

gia, Marion Island, Prince Edward Island, lies

de Kerguelen, lies Crozet and Herd) (Bliss,

1979, p. 2168).

and lichens. Two flowering plants, Deschampsia

anlarctica and Colobanthus crassifolius, occur in

Antarctica.

The antarctic botanical zone was subdivided by

Holdgate ( 1 964) into the maritime antarctic, which

consists of the west coast of the Antarctic Penin-

sula and the adjacent islands in the south Atlantic,

where the marine influence allows growth of a wide

range of bryophytes and lichens in addition to the

two flowering plants; and the more extensive con-

tinental antarctic, where bryophyte and lichen

communities are only sparsely developed and

flowering plants are unknown.

Several workers have presented vegetation clas-

sification systems for all or portions of the ant-

arctic zone. Longton (1967) classifies the vegeta-

tion of the maritime antarctic and uses an

"Antarctic cryptogam formation" and later (Long-

ton, 1973, 1979, 1982) provided classifications of

vegetation within the antarctic botanical zone in

which we find an "Antarctic nonvascular crypto-

gam tundra formation." Gimingham and Smith

(1970) and Smith and Gimingham (1976) present

classifications of the cryptogamic communities of

the maritime antarctic, and Smith (1972) does

likewise for the South Orkney Islands; all of these

authors use an "Antarctic nonvascular cryptogam

tundra formation."

Presence versus absence of permafrost is also

an important factor in austral regions. Permafrost

is lacking in the subantarctic zone, even at higher

elevations, but is a feature throughout the antarctic

zone.

Thus, the terms subantarctic and antarctic as used

by Bliss cannot be identified by zones. I concur

with Smith (1984, p. 65) that the comparison by

Bliss of "Arctic, alpine, Antarctic and sub-Ant-

arctic vegetation categories is unsatisfactory be-

cause the impoverished southern flora has a dif-

ferent balance of life forms."

The antarctic region includes regions south of

the subantarctic and is characterized by a vege-

tation which ". . . is at best a poorly developed

tundra" (Rudolph, 1971, p. 192). Longton (1973,

p. 2339) points out that "The vegetation through-

out the Antarctic zone is less well developed than

that at most sites in even the high Arctic, at least

in terms of flowering plant representation. . . ."

Soligenous mires, herbfields, tussock forming

grasses, and cushion plants are absent, and the

vegetation is a tundra type dominated by mosses

The Phytogeographic Categories

The chief difficulty in the assessment of the phy-

togeographical relationships of the Falkland Is-

lands Hepaticae and Anthocerotophyta is the de-

limitation of the south temperate and subantarctic

patterns of distribution. I have delimited these

patterns after the concept of the zones or regions

developed by Skottsberg ( 1 9 1 0, 1 9 1 6, 1 960), God-

ley ( 1 960), Wace ( 1 960, 1 965), and Greene ( 1 964).

I recognize 1 2 categories of distribution patterns

of Falkland Islands Hepaticae (table 4). The An-

thocerotophyta fall within only one of these cat-

egories. The data for these distributions were gath-

ered from specimens examined and reports

extracted from the literature. If a report is regarded

as questionable, it is not included here.

I recognize 1 30 species of Hepaticae and one

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

41

species of Anthocerotophyta belonging to the

Falkland Islands flora.

SOUTH TEMPERATE— Species occurring within the

south temperate regions of the world.

American Temperate— Species occurring within

the south temperate regions of the American sec-

tor. There are eight species which occur in south-

ern South America as well as South Georgia (see

p. 43). Their distribution is regarded as temperate

rather than subantarctic, as northward extension

from the subantarctic is not confined to higher

altitudes.

It has frequently been remarked that the Falk-

land vascular plant flora is an extension of the

southern South American flora (Hooker, 1847;

Skottsberg, 1909, 19 13; Moore, 1 968), a statement

to be expected with regard to a continental shelf

island. Moore stated that, of the 1 63 native species

of vascular plants of the Falkland Islands, 89%

occur on the mainland south of 40°S. All except

three of the remainder are endemic to the Falkland

Islands and are in large part related to mainland

taxa. Like the vascular flora, the mainland hepatic

representatives extend northward to varying de-

grees. Commencing with the endemic category, the

remainder of groups are arranged principally ac-

cording to the degree of northward extension.

Endemic to Falkland Islands (Group 1)

Andrewsianthus planifolius

Metzgeria engelii

Riccardia regularis

Schistochila subantarctica

Fuegian- Falkland (Group 2)— Taxa occurring in

the Falkland Islands and Tierra del Fuego. Aus-

trolophoziafuegiensis is the only species belonging

here. Several other species occur in the Falklands

and in southern South America only within Tierra

del Fuego, but such taxa also may be found on

South Georgia (and sometimes other subantarctic

islands) plus sometimes in the antarctic zone. Thus

such species are either subantarctic in distribution

(e.g., Acrolophozia fuegiana, Adelanthus integer-

rimus. Riccardia granulata, R. saxicola), or ant-

arctic in distribution (e.g., Cephalozia badia, Her-

zogobryum teres).

Magellanian- Falkland (Group 3)— Taxa occur-

ring in the Falklands and southern Patagonia north

to 48°S or Fuegia and southern Patagonia north

to 48°S. The northern boundary was affixed by

Skottsberg (1916) to delimit the Magellanian and

Valdivian regions and has been widely followed

by various authors.

SOUTH TEMPERATE

American Temperate

Endemic to Falkland

Islands 4 3.1

Fuegian-Falkland 1 0.7

Magellanian-Falkland 19 14.5

Valdivian-Falkland 3 2.3

Magellanian + Valdivian

+ Falkland 52 39.7

Andean-Falkland 2 1.5

Extra-American Temperate

Amphipacific temperate 1 3 9.9

Amphiatlantic temperate 1 1 8.4

Pan-south temperate 6 4.6

NONTEMPERATE

Subantarctic 12 9.2

Antarctic 6 4.6

Widespread 2 1.5

Total 131 100.0

Of the 1 9 taxa in this group, 74% have been

found to occur in the Magellanian moorland. Their

occurrence in the moorland is indicated by an as-

terisk, and those taxa restricted to the moorland

are indicated by a double asterisk. This region

occurs south of 48°S to Cabo de Hornos, and is

limited by the Magellanian rain forest to the east

and the Pacific Ocean to the west (see discussion

and map in Godley, 1960; see also Pisano, 1983,

who provides a detailed treatment, preferring the

term "Magellanic Tundra Complex"). The moor-

land is extremely wet, exposed to violent winds,

has a permanently saturated peaty soil, and is par-

tially to distinctly open. Of the remaining four

taxa, Chiloscyphus elatus is known from deciduous

Nothofagus forests, Adelanthus tenuis is known

from evergreen Nothofagus forests, and Anastro-

phyllum ciliatum and Archeochaete kuehnemannii

are known from both deciduous and evergreen

Nothofagus forests.

Adelanthus tenuis

Anastrophyllum ciliatum

* Andrewsianthus australis

Archeochaete kuehnemannii

* Balantiopsis bisbifida

* Cephalolobus scabrellus

Chiloscyphus elatus

* Chiloscyphus hookeri

42

FIELDIANA: BOTANY

Frullania microcaulis

Gackstroemia patagonica

Kurzia mollis

Kurzia setiformis

Leptoscyphus aequatus

Pachyschistochila splachnophylla

(^Plagiochila dura

Plagiochila obovata

Riccardia fuscobrunnea

Saccogynidium vasculosum

Telaranea oligophylla

Valdivian- Falkland (Group 4,) —Species in the

Falklands and Chile and/or Andean Patagonia (see

group 5 for definition) north of 48°S and south of

36°S. See Engel (1978, pp. 21-23) for comments

on this zone. All species also occur on Juan Fer-

nandez.

Jungermannia crassula

Metzgeria multiformis

Wettsteinia densiretis

Valdivian + Magellanian + Falklands (Group

5)— Species that are essentially widespread in the

American temperate zone. Of the 52 taxa in this

group, 7 1% are known to occur in the Magellanian

moorland. Their occurrence in the moorland is

indicated by an asterisk. It should be repeated here

that the moorland occurs south of 48°S and thus

is restricted to the Magellanian floristic region.

With regard to the Magellanian region, of the re-

maining 29%, Cheilolejeunea savatieriana, Chi-

loscyphus sylvaticus, Clasmatocolea rigens, Diplo-

phyllum acutilobum, Leptophyllopsis irregularis,

Megaceros fuegiensis, Paracromastigum subsim-

plex. Riccardia papillosa, and Symphyogyna

hochstetteri occur in the deciduous Nothofagus

zone, Lejeunea corralensis, Plagiochila acantho-

caulis, P. ansata, and P. gayana in the evergreen

Nothofagus region, and Chiloscyphus sabuletorum,

Riccardia opuntiiformis, and Telaranea pseudo-

zoopsis in the deciduous and evergreen Nothofagus

regions.

The taxa, with the exception of groups 5e-f, are

arranged according to their latitudinal range.

5a. South from 45°S:

Cheilolejeunea savatieriana

* Chiloscyphus divaricatus

* Pigafettoa crenulata

Riccardia opuntiiformis

* Riccardia spectabilis

5b. South from 43°30'S (line from north side of

Isla Guafo):

* Chiloscyphus leptanthus

* Chiloscyphus textilis

* Kurzia saddlensis

* Leptoscyphus patagonicus

Plagiochila acanthocaulis

Plagiochila ansata

* Riccardia pallidevirens

* Telaranea plumulosa

5c. South from 40°S:

* Blepharidophyllum gottscheanum

* Clasmatocolea obvoluta

Diplophyllum acutilobum

Megaceros fuegiensis

* Plagiochila fagicola

* Riccardia alcicornis

* Riccardia tenax

* Schistochila laminigera (40°07'S)

Telaranea pseudozoopsis

5d. South from 36°S; from latitude indicated:

* Balantiopsis cancellata (39°48'S)

* Balantiopsis erinacea (36°50'S)

* Chiloscyphus austrigenus (39°52'S)

* Clasmatocolea fulvella (36°50'S)

* Cryptochila paludosa (39°52'S)

* Frullania boveana (39°53'S)

* Frullania magellanica (39°52'S)

* Harpalejeunea parasitica (39°48'S)

* Isotachis humectata (36°43'S)

* Nothostrepta bifida (39°52'S)

* Plagiochila elata (39°36'S)

* Radula helix (39°38'S)

* Telaranea blepharostoma (39°48'S)

* Tylimanthus urvilleanus (39°38'S)

5e. Valdivian + Magellanian + Falklands + South

Georgia + sometimes also South Sandwich

Islands.

The species listed here are considered as south

temperate rather than subantarctic, as northward

extension from the subantarctic is not confined to

higher altitudes.

* Cephaloziella dusenii

Clasmatocolea rigens

* Gackstroemia magellanica

* Lepicolea rigida

* Lepidozia chordulifera

* Lepidozia fuegiensis

* Riccardia papillosa

* Roivainenia jacquinotii

5f. Species about which insufficient knowledge is

known concerning range:

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

43

Chiloscyphus sabuletorum

Chiloscyphus sylvaticus

Lejeunea corralensis

Leptophyllopsis irregularis

* Metahygrobiella tubulata

Paracromastigutn subsimplex

Plagiochila gayana

Symphyogyna hochstetteri

Skottsberg ( 1 9 1 0, 1 9 1 6) and Moore ( 1 968) rec-

ognized three vegetation zones in the region south

of40°S:

1 . West Patagonia: The region including the west

slope of the Andes to the Pacific Ocean, char-

acterized by high precipitation and supportive

of lush rain forests and Magellanian moorland.

2. Andean Patagonia: The eastern slope of the

Andes from base to snow line. This region ex-

periences moderate rainfall and supports a de-

ciduous, comparatively dry forest.

3. East Patagonia: The steppe region, ranging east

from the Andean foothills, characterized by

grassland and xerophytic shrubs.

As is to be expected, the vast majority of He-

paticae in groups 3-5 occur predominantly in the

West Patagonian zone. The following Valdivian-

Magellanian Hepaticae and Anthocerotophyta oc-

cur within the Andean Patagonia zone, and nearly

all occur in the Lago Nahuel Huapi region; none

are restricted to Andean Patagonia:

Chiloscyphus text His

Clasmatocolea fulvella

Clasmatocolea rigens

Frullania magellanica

Isotachis humectata

Lepidozia chordulifera

Megaceros fuegiensis

Plagiochila elata

Riccardia alcicornis

Riccardia papillosa

Riccardia tenax

Roivainenia jacquinotii

Schistochila laminigera

Telaranea pseudozoopsis

Tylimanthus urvilleanus

Megaceros fuegiensis is the only species of Val-

divian-Magellanian Hepaticae or Anthoceroto-

phyta to occur in East Patagonia. Of the remaining

Falkland Islands taxa, Leptoscyphus expansus,

Marchantia berteroana, and Symphyogyna hy-

menophyllum also occur there.

Andean- Falkland (Group 6)— Species extending

north of 36°S in the Andes. Of the American tem-

perate species, Noteroclada confluens and Pseu-

docephalozia quadriloba are the only taxa that have

utilized the Andes as a route for northward mi-

gration to lower latitudes.

Extra-American Temperate— Amp hipacific

Temperate— Distribution mainly in temperate

parts of the South Pacific in the Southern Hemi-

sphere, i.e., temperate South America (occasion-

ally Juan Fernandez), New Zealand, Tasmania,

and southeast Australia. There are varying degrees

of penetration northward into the New Zealand

sector, and the taxa have been arranged accord-

ingly. One asterisk indicates occurrence on New

Zealand shelf islands, two indicate occurrence in

New Zealand, and three indicate occurrence on

New Zealand shelf islands and New Zealand. Only

two taxa also occur in the Northern Hemisphere:

Metzgeria decipiens and Triandrophyllum subtri-

fidum.

a. South Island:

** Metzgeria violacea

b. To North Island:

** Heteroscyphus triacanthus

** Pallavicinia xiphoides

** Saccogynidium australe

** Symphogyna hymenophyllum

*** Telaranea tetradactyla

*** Temnoma quadripartitum

c. To Tasmania:

*** Chiloscyphus bispinosus

*** Lepidozia laevifolia

* Triandrophyllum subtrifidum

d. To Australia:

*** Chiloscyphus lentus

** Clasmatocolea humilis

e. To Japan:

** Metzgeria decipiens

Amphiatlantic Temperate— Distribution main-

ly in temperate parts of the south Atlantic, i.e.,

temperate South America, South Africa (plus oc-

casionally Tristan da Cunha). Species whose sole

occurrence in the Indian Ocean sector is on sub-

antarctic islands are included here and indicated

by an asterisk. They are not considered as sub-

antarctic species, as northward extensions from

the subantarctic are not restricted to higher alti-

tudes. Gradstein et al. ( 1 983) discuss several species

that belong to this element.

Adelanthus lindenbergianus

* Blepharidophyllum clandestinum

* Blepharidophyllum densifolium

44

FIELDIANA: BOTANY

Hyalolepidozia bicuspidata

* Jensenia pisicolor

Lepicolea ochroleuca

Leptoscyphus expansus

* Lethocolea radicosa

Paracromastigum tristanianum

Riccardia prehensilis

Schistochila alata

All of these taxa, with the exception of Hyalo-

lepidozia bicuspidata and Paracromastigum tris-

tanianum, penetrate north via the Andes to lower

latitudes and are known from relatively high al-

titudes north of 36°S.

Pan- South Temperate— Species occurring in

temperate regions of South America, New Zea-

land-Australia, and South Africa:

Acrobolbus ochrophyllus

Chiloscyphus semiteres

Clasmatocolea vermicularis

Cryptochila grandijlora

Jamesoniella colorata

Marchantia berteroana

NONTEMPERATE

Subantarctic— Species occurring on one or more

subantarctic islands (as denned by Greene, 1964)

of at least one sector (e.g., American [A], Indian

Ocean [I], or New Zealand [NZ], with northward

extensions only at higher altitudes; see comments

inEngel(1978, pp. 35-36):

Acrolophozia fuegiana (A)

Adelanthus integerrimus (A, I)

Chiloscyphus humifusus (A, I)

Herzogobryum erosum (A, NZ)

Herzogobryum vermiculare (A, I)

Pachyglossa fissa (A, I)

Pachyglossa spegazziniana (A)

Pachyschistochila leucophylla (A)

Pedinophyllopsis abditus (A, I)

Riccardia georgiensis (A, I)

Riccardia granulata (A)

Riccardia saxicola (A)

Antarctic— Species occurring in the antarctic

zone (as denned by Greene, 1 964), with northward

extensions into the subantarctic or temperate zones

only at higher altitudes. Species in this category

are able to tolerate considerably colder tempera-

tures than those in the subantarctic category. Oc-

currence on subantarctic islands is indicated with

the same symbols used for subantarctic species.

(It should be mentioned that the worldwide dis-

tribution of L. hatcheri is bipolar.)

Cephalozia badia (A)

Evansianthus georgiensis (A)

Herzogobryum teres (A, NZ, I)

Hygrolembidium isophyllum (A)

Lophozia hatcheri (A)

Pachyglossa dissitifolia (A)

Widespread— Species not in any of the above

categories and not pantropical. These species,

which have a broad distribution with a range in-

volving to a considerable extent the Northern

Hemisphere, are Aneura pinguis and Metzgeria

leptoneura.

Distribution Within the Falklands

The taxa below are arranged according to their

occurrence in either the East Falklands or the West

Falklands.

TAXA KNOWN ONLY FROM EAST FALKLANDS—

Of these, 48% are known only from the Mt. Us-

borne region (indicated by an asterisk):

* Acrolophozia fuegiana

Andrewsianthus planifolius

* Blepharidophyllum gottscheanum

Cephalolobus scabrellus

Chiloscyphus divaricatus

Chiloscyphus text His

Clasmatocolea obvoluta

* Gackstroemia patagonica

* Kurzia mollis

* Kurzia saddlensis

* Kurzia setiformis

* Lejeunea corralensis

Lepicolea ochroleuca

Lepicolea rigida

Leptoscyphus aequatus

* Nothostrepta bifida

Pachyglossa fissa

* Paracromastigum tristanianum

Plagiochila fagicola

* Riccardia granulata

Riccardia prehensilis

* Riccardia saxicola

* Saccogynidium australe

Schistochila alata

Telaranea blepharostoma

* Telaranea oligophylla

Telaranea tetradactyla

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

45

TAXA KNOWN ONLY FROM WEST FALKLANDS

Chiloscyphus bispinosus

Chiloscyphus humifusus

Chiloscyphus semiteres

Diplophyllum acutilobum

Herzogobryum erosum

Leptophyllopsis irregularis

Lethocolea radicosa

Megaceros fuegiensis

Metzgeria violacea

Pachyschistochila splachnophylla

Paracromastigum subsimplex

Pigafettoa crenulata

Plagiochila dura

Plagiochila obovata

Radula helix

Riccardia regularis

Schistochila subantarctica

While I regard the highly local occurrence of

many Falkland Islands taxa to be a reflection of

the availability of microhabitats and narrow eco-

logical requirements, the comparison of hepatics

restricted to the East Falklands (27) compared to

the West Falklands (17) is impressive, and may

well reflect differing climatic conditions within each

of the respective island groups. Alternatively, the

difference in species diversity between East and

West Falklands may be due to critical local levels

of temperature and rainfall which are present in

various parts of the Falkland Islands (see fig. 6

with rainfall indicated; compare especially San

Carlos with Port San Carlos).

Port Stanley is one of the wettest areas in the

entire archipelago, and this feature coupled with

the cool temperatures known for the region may

account for the eight taxa which were found to

occur only in the Port Stanley-Port William re-

gions. These species fall within a variety of phy-

togeographic elements.

Andrewsianthus planifolius

Cephalolobus scabrellus

Chiloscyphus divaricatus

Lepicolea ochroleuca

Riccardia prehensilis

Schistochila alata

Telaranea blepharostoma

Telaranea tetradactyla

The northwesternmost portion of the Falkland

Islands has higher temperatures, and within this

region are confined, or nearly so, three vascular

plants (Blechnum chilense, Gavilea macroptera, and

Ranunculus acaulis) of the Valdivian element.

Chiloscyphus semiteres is known in the Falkland

Islands only from Westpoint Island. The species

seems to be restricted to rather warm temperature

regions, especially in the American sector (i.e., the

species is unknown for the Magellanian region),

where it may be considered Valdivian.

Of the taxa occurring on both East and West

Falklands, two were confined to the wettest known

stations. Frullania microcaulis and Metzgeria en-

gelii were collected at Port Stanley and Port How-

ard.

Rainfall figures for the Mt. Usbome region would

be of great interest. A large number of Hepaticae

occur strictly in the Mt. Usborne region as well as

at the wettest known stations on record, as the lists

below illustrate. It is likely that the area receives

as much rainfall as that of Port Stanley, Port How-

ard, or Mt. Adam. Following are Hepaticae known

only from the regions of Mt. Usborne, Port How-

ard, and Mt. Adam:

Anastrophyllum ciliatum

Cryptochila grandiflora

Cryptochila paludosa

Herzogobryum teres

Herzogobryum vermiculare

Hygrolembidium isophyllum

Pachyglossa dissitifolia

In addition to the above rather sizeable suite of

hepatics occurring only in the Mt. Usborne, Port

Howard, and Mt. Adam regions, a rather large

group of species is also known from combinations

of these stations plus the Stanley region. Taxa oc-

curring only in two or more of the Port Stanley

(S), Mt. Usborne (U), Port Howard (H), and Mt.

Adam (A) regions are:

Acrobolbus ochrophyllus (U, A)

Adelanthus integerrimus (U, H)

Archeochaete kuehnemannii (U, A)

Austrolophozia fuegiensis (S, U, H, A)

Blepharidophyllum densifolium (S, U, A)

Cephalozia badia (U, H)

Clasmatocolea obvoluta (S, U)

Evansianthus georgiensis (U, H)

Frullania microcaulis (S, H)

Jensenia pisicolor (S, U, H, A)

Lepidozia fuegiensis (S, U, H, A)

Leptoscyphus aequatus (S, U)

Metzgeria engelii (S, H)

Pachyglossa spegazziniana (S, U, H, A)

Pachyschistochila leucophylla (S, U, H, A)

Riccardia fuscobrunnea (U, H)

Riccardia pallidevirens (U, A)

46

FIELDIANA: BOTANY

Comparative Taxonomic Composition

of the Hepatic- Anthocerote Flora

It is revealing to compare the Falkland flora to

that of southern South America, and especially to

the Magellanian region, the area on the mainland

most closely related phytogeographically to the

Falklands. The Brunswick Peninsula is ideal for

this comparison, since it is the southernmost land

mass on mainland South America; its flora is

strongly Magellanian in composition; and there is

a complete floristic account of the Hepaticae of

this region— the only one for all of southern South

America (Engel, 1978). The following chart con-

trasts the floras:

Brunswick Falkland

Peninsula Islands

No. of families

No. of genera

No. of species

No. of species per genus

29

743

193

1.8

26

64

131

2

In making such a comparison, the absence of a

taxonomic unit is at times just as interesting or

revealing as its presence. The families present in

the Brunswick Peninsula but not in the Falklands

are Trichocoleaceae, Porellaceae, and Calypogei-

aceae. On the other hand, all families occurring in

the Falklands also are present on the Brunswick

Peninsula. It is also revealing to contrast the ge-

neric composition of the Falklands. Genera pres-

ent on the Brunswick Peninsula but absent in the

Falklands are:

Acromastigum

Allisoniella

Anthoceros

Aphanolejeunea

Apometzgeria

Archilejeunea

Austrolejeunea

Bazzania

Calypogeia

Colura

Hepatostolonophora

3 In order to increase the utility of the comparison of

genera, I have made taxonomic adjustments among

Brunswick Peninsula names so that they can be com-

pared with Falkland Islands generic names. I have done

so in three cases: Heteroscyphus occurs in the Brunswick

Peninsula, but was included under Chiloscyphus in Engel

(1978); likewise Pachyschistochila taxa were included

under Schistochila and Pedinophyllopsis was included

under Leptoscyphus. Thus the number of Brunswick

Peninsula genera actually cited in Engel (1978) was 71.

Herbertus

Krunodiplophyllum

Lepidolaena

Paraschistochila

Pleurocladopsis

Porella

Pseudolepicolea

Reboulia

Trichocolea

Nearly all of the genera in the above list have

ecological preferences in the Brunswick Peninsula

that are in sharp contrast to any niche available

on the Falklands (Engel, 1978). The vast majority

of genera absent from the Falklands but present

in the Brunswick Peninsula are restricted to the

Nothofagus zone, and many are known only from

evergreen Nothofagus forests.4 Several genera are

absent because their constituent species occur only

in niches offered by forest vegetation, e.g., on bark

of living trees or on rotted logs. Examples of such

taxa are Trichocolea, Bazzania, and Porella, all of

which are common in wet forests of southern South

America. Other genera commonly utilize filmy fern

fronds as a niche, a feature characteristic of wet

forests in southern South America, but lacking in

the Falklands. Examples are Archilejeunea and

Aphanolejeunea. Forests are completely lacking in

the Falklands, except for the introduced com-

munity at Hill Cove (figs. 20-21). Thus, because

of the absence of forest communities in the Falk-

lands and consequently the absence of large woody

substrates, the lower humidity, and the lack of

protection from the sun, a large number of mi-

croniches available to bryophytes in a forested area

such as on the Brunswick Peninsula are completely

lacking in the Falklands.

A second important factor relevant to distri-

bution patterns of hepatics in the Brunswick Pen-

insula is that of differences in rainfall. Nearly all

genera of the Brunswick Peninsula that are lacking

in the Falklands are restricted in the Brunswick to

regions receiving in excess of 1 ,000 mm of rainfall

annually. The highest annual rainfall recorded in

the Falklands is 646 mm (fig. 6). The 1 ,000-mm

rainfall figure seems to be critical in the Brunswick

Peninsula, and there are dramatic differences in

the numbers of taxa restricted to regions receiving

more than 1 ,000 mm annual rainfall and those to

4 Engel (1973b, 1978) was the first bryologist to cor-

relate southern South American vascular plant vegeta-

tion types and zonation to distribution of hepatics. See

these references for details.

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

47

regions receiving less than this amount (see Engel,

1978, table 3, pp. 40-41).

The absence of a forest community and the com-

paratively low annual rainfall undoubtedly are

major factors in accounting for the absence from

the Falklands of taxa from a range of different

taxonomic levels.

The following 1 0 genera occur on the Falklands

but are unknown for the Brunswick Peninsula:

Acrolophozia

Aneura

Austrolophozia

Jensenia

Jungermannia

Pachyglossa

Pallavicinia

Paracromastigum

Pigafettoa

Wettsteinia

Based upon known distribution patterns, some of

the taxa (e.g., Acrolophozia, Austrolophozia, and

Pigafettoa) are rather rare species throughout their

range and perhaps truly are absent from the Bruns-

wick Peninsula. Furthermore, two of the genera,

Jungermannia and Wettsteinia, have a Valdivian-

Falkland distribution and thus are unknown for

the entire Magellanian zone. With further collect-

ing, however, undoubtedly at least several of these

genera will eventually be found in the Brunswick

Peninsula.

Origin of the Flora

The phytogeographic affinities of the Falkland

Islands hepatics and single anthocerote (see con-

spectus on p. 42) indicate not only a strong re-

lationship to South America but also noteworthy

relationships to other austral land masses. Long-

range dispersal may play a role in explaining some

geographical discontinuities of bryophytes (van

Zanten & Pocs, 1981; van Zanten & Gradstein,

1988), but the concept of continental drift is cru-

cial to understanding floristic similarities, espe-

cially at the species level, of austral bryophytes.

The various Gondwanaland reconstructions de-

picted in the literature vary somewhat in detail,

but do not deviate to a major extent from the

reconstructions shown in Figure 4. As discussed

in Geological Features, the age of Gondwanaland

and its subsequent breakup is also subject to dif-

fering interpretation; such differences, however,

are not of major importance to our discussion.

Two points are paramount for our purposes:

1 . With the continuity of the land masses making

up Gondwanaland, a migratory route was

available that involved South America, South

Africa, the fringes of Antarctica, Australia, Tas-

mania, and New Zealand. The general impor-

tance of this pathway is discussed in papers by,

for example, Schuster (1969c, 1972b, 1976,

1979b, 198 la, 1982, 1 98 3) and Raven and Ax-

elrod ( 1 974). Discussions relevant to origin and

distribution of specific Gondwanalandic he-

patic groups may be found in monographs, for

example, of Pseudocephalozia (Schuster & En-

gel, 1974), of Schistochilaceae (Schuster & En-

gel, 1977, 1985), of Clasmatocolea (Engel, 1980)

and of Lepidoziaceae subfam. Lembidioideae

(Schuster & Engel, 1987).

2. The migratory route resulting from the geo-

graphic continuity of ancient Gondwanaland

was available for a very long period of time.

As I have previously emphasized (Engel, 1980,

p. 33), "This migratory pathway is of funda-

mental importance in understanding the phy-

togeography of the far south."

The association and juxtaposition of the Falk-

lands, together with the South American block,

and other land masses involved with Gon-

dwanaland are the foundation of our discussion.

Available migratory pathways involving Gon-

dwanaland are crucial to understanding the phy-

togeographic relationships of the Falkland hepatic

flora. Evidence indicates that the Falklands were

associated with the South American block at least

since the Lower Cretaceous (see fig. 4 and discus-

sion on p. 6). The proximity of the Falklands to

southern South America over a long period of time

is at least partially the reason for the very strong

phytogeographic affinity of the islands to southern

South America. The conspectus on p. 42 reveals

that the majority of the Falkland hepatic flora

(61.8%) may be categorized as South American

temperate. Falkland-South American temperate

distribution patterns differ in the degree of range

extension on the mainland, but all are similar in

involving the Nothofagus zone of southern South

America. The range may be narrow and rather

restricted, as in Austrolophozia fuegiensis (fig. 74),

to widespread in southern South America, as in

Roivainenia jacquinotii (fig. 42), to the Andean-

Falkland pattern illustrated by Noteroclada con-

fluens and Pseudocephalozia quadriloba (fig. 29).

Migratory pathways involving Gondwanaland

are also relevant to discussion of the amphipacific

(fig. 71, p. 145), amphiatlantic (fig. 23, p. 64) and

48

FIELDIANA: BOTANY

pan-south-temperate (fig. 45, p. 101) distribution

patterns, together making up 22.9% of the flora,

as well as to subantarctic (fig. 46, p. 105) and ant-

arctic (fig. 47, p. 105) patterns, together totaling

1 3.8% of the flora. In this connection, the sheltered

high altitude vegetation in the Falklands is worthy

of special mention (see p. 36). In this community,

there is a strikingly high percentage of hepatics

that either have a subantarctic or antarctic distri-

bution, or also occur at higher elevations in New

Zealand and Tasmania or have very close relatives

there. The hepatic flora of this association appears

to be in large part an old, relict one. A number of

wide-ranging taxa (especially the genera Acrolo-

phozia, Austrolophozia, Herzogobryum, and

Pachyglossd) may well represent remnants of an

early, widespread flora involving the Antarctic

continent. Schuster (1969c) cited all of these gen-

era and more in connection with discussions of

antipodal distribution patterns, and stated (p. 56)

that "it seems reasonable to regard these as relict

groups at least in most cases," and that "presum-

ably some of these genera already existed prior to

the wide separation between the Tasmania-New

Zealand area and Fuegia from Antarctica." Schus-

ter (p. 56) cited Acrolophozia (but could have also

cited Austrolophozia, Herzogobryum, or Pachy-

glossd) as

". . . genera which are alpine today, often ex-

isting in areas with summer snows, [and that

such genera] may well have survived in what

is today Antarctica even after its climate was

already fairly inclement. Indeed, ecologically,

the striking common denominator of most of

the genera cited above (Phyllothallia, Ble-

phahdophyllum, Acrolophozia, Austrolopho-

zia, Herzogobryum, Pachyglossa) is that they

are today essentially confined to treeless al-

pine zones or extend to the lowlands only on

the subantarctic islands. These taxa may then

represent remnants of an early cold-antipodal

flora that characterized perhaps the colder up-

lands of an Antarctic continent whose low-

lying areas harbored Nothofagus or Podocar-

pus forests."

Close juxtaposition with Africa was postulated

over 35 years ago by Adie (1952b, cf. fig. 3). That

concept was not widely followed until it was re-

cently reexamined, with new evidence, by Martin

et al. (1981), Mitchell et al. (1986), and Martin

(1986) (see p. 6 and fig. 5). While hepatics as a

group are old enough to reflect an Early Jurassic

African-Falkland association, there are no exclu-

sively African-Falkland taxa; all species that occur

in the Falklands and in Africa also occur in South

America. This is to be expected if one recalls that

movement of a Falkland microplate occurred dur-

ing the Jurassic, i.e., at a time that Africa and

South America were juxtaposed and part of Gon-

dwanaland (fig. 4). The continuity of Africa and

South America was the most important event, and

of secondary relevance is any possibility of intro-

duction of African taxa to South America via a

Falkland microplate.

Systematic Account

Notes on Format

The sequence of orders and families follows the

classification outlined in Schuster (1979a). The

genera are arranged in alphabetical order within

families, and species in alphabetical order within

genera. Author citations follow the abbreviations

in Sayre et al. (1964). Journal citations follow the

abbreviations in Botanico-Periodicum-Huntian-

um (B-P-H).

I have attempted to indicate the status of typ-

ification of all taxa; I have followed the concepts

outlined in Engel (1978, cf. pp. 51-52), and have

provided ecological notes for each species I have

collected in the Falklands. The terminology and

concepts used in these discussions have been de-

fined in Phytosociology (see pp. 1 3-38).

I have listed the Falkland Islands literature rec-

ords for species that have been previously reported

from these islands. The information is presented

as it is given in the original source, and no attempt

has been made to edit the extractions. For ex-

ample, there are several instances where a collector

is reported for a locality he did not visit. NOTE:

References in which Falkland Islands taxa were

originally described are given only in the taxo-

nomic citations and are not repeated in the liter-

ature record section.

The localities in the synonymy sections are cited

in the manner provided in gazetteers of the various

regions, and it is only to this extent that I have

edited the extractions. The following gazetteers

were utilized: Chile (U.S. Office of Geography,

1967), Argentina (American Geographical Society

of New York, 1944), and New Zealand (New Zea-

land Dept. of Lands and Survey, 1968). Abbre-

viations of localities follow the American Geo-

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

49

graphical Society indices to maps of Hispanic

America (see American Geographical Society of

New York . . . Argentina . . . 1944) and are as

follows; add 5 to form plural:

Arch. —Archipelago

A.— Arroyo

B.-Bahia

Br.— Brazo

Cta.— Caleta

C.-Cerro

Cord.— Cordillera

Ens.— Ensenada

Esto.— Estuario

F.— Fiordo

I.-Isla

L.— Lago

La. — Laguna

M.— Monte

Mt.— Mount, Moun-

tain

Pen. — Peninsula

Pta.— Punta

Port. — Portezuela

Pto.— Puerto

Q.— Quebrada

Ran.— Rancho

R.— Rio

S.— Seno

Sa.— Serra, Sierra

Vo. — Ventisquero

V.— Volcan

I have grouped all Falkland collections together,

followed by additional collections seen. The spec-

imens I have personally collected are indicated by

number only, without citation of collector.

Distribution maps are provided for a number

of taxa to illustrate the patterns of variation within

and between phytogeographic categories as well as

taxonomic ranks. Maps are only provided for those

taxa for which there are sufficient reliable data.

Introduction to Keys

The keys include all taxa known to be present on the Falkland Islands as well as all genera that occur

in the Magellanian zone of southern South America (cf. p. 42 for definition of this zone). The addition

of non-Falkland genera will increase the utility of this treatment and allow the user to key, to generic

level, all hepatics and anthocerotes presently known from the Magellanian sector; the keys thus include

the vast majority of genera from the Nothofagus zone of South America. Genera (or higher taxa) not

known for the Falklands are bracketed in the keys.

Key to Classes and Orders of Hepaticae and Anthocerotophyta5

1 . Gametophytic chlorophyllose cells with numerous chloroplasts; chlorophyllose cells usually all (or

many) with oil-bodies; gametophyte thalloid or leafy, without stomata, at times with air chambers;

sporophyte a nonlinear capsule, usually 4-valved, without a columella, without stomata; sex organs

exogenous Division Bryophyta, Class Hepaticae ... 2

2. Rhizoids, if present, all smooth, isolated thickenings excepted; plants leafy or thalloid, if thalloid

± delicate and translucent, without air chambers or pores; cells not sharply dimorphic, oil-bodies

usually present in all chlorophyllose cells of gametophyte; capsule wall 2-10 stratose; seta usually

long and extruding the sporophyte 3

3. Plants clearly and uniformly leafy; archegonia usually terminal on shoots resulting in a cessation

of plant growth; capsule wall 2-10 stratose; rhizoids present or lacking 4

4. Rhizoids usually present; plants usually prostrate to procumbent, usually bilateral and

anisophyllous or distichous-leaved; leaves, if unlobed, obliquely inserted. Sporophyte with

wall 2-10-stratose, the cells variously thickened or lacking bandlike thickenings, never with

annular longitudinal solitary bands. Perianths (or else a coelocaule, marsupium, or other

protective device) usually present; if only with a shoot calyptra, the shoot calyptra on a

short, postical branch Order Jungermanniales (p. 51)

4. Rhizoids never developed; plants erect, isophyllous, fundamentally radial (without distinct

division into antical and postical faces); leaves unlobed, flat, transversely inserted. Sporophyte

wall unistratose, with annular, longitudinal, solitary bands; perianth not known to occur, the

green, massive, fleshy shoot calyptra replacing it Order Calobryales (p. 5 1)

3. Plants usually thallose (leafy only in Noteroclada and Treubia); archegonia and antheridia

scattered on dorsal surface of thallus (occasionally on short thallus branches), or in dorsal

groups, not causing cessation in growth of plant; capsule wall 2-5 stratose; rhizoids always

present 5

* Key basically adapted from Schuster (1963a).

50

FIELDIANA: BOTANY

5. Plants thalloid, or if leafy, then not with two rows of dorsal leaflike appendages; gametangia

variable in position, never in axils of foliar appendages. . . . Order Metzgeriales (p. 167)

5. Thallus leafy, the dorsal surface of thallus with two rows of leaflike appendages; gametangia

in axils of dorsal "leaves." Cells strongly dimorphic, some or most with solitary, large oil-

bodies that nearly fill the cell lumen, others bearing only chloroplasts

[Order Treubiales, Treubia, cf. p. 1 67]

2. Rhizoids typically dimorphic, some tuberculate, others smooth; plants clearly thalloid, without

leaflike lobes, the thallus firm, fleshy, opaque, with ventral scales, with air chambers or air canals

opening dorsally by pores; cells typically dimorphic, a small minority of generally smaller cells

each with a single, large oil-body, but no chloroplasts, the large majority of cells with chloroplasts

only; capsule wall always unistratose; seta short, not extruding the sporophyte to any extent.

Archegonia on specialized thallus branches (archegoniophores) Order Marchantiales

Gametophytic cells (at least superficial cells) each normally with 1 (2-several) chloroplasts; cells

lacking oil-bodies; gametophyte prostrate, thalloid, never with leaves, never with air chambers, with

stomata on ventral surface; sporophyte linear, 2-valved, with a columella; archegonia and antheridia

endogenous Division Anthocerotophyta, Order Anthocerotales

Order CALOBRYALES

An order with two families, Takakiaceae and Haplomitriaceae, each with one genus. Haplomitrium

chilensis Schust. occurs in southern South America (see Schuster, 1971; Engel, 1984) but is absent from

the Falklands.

Order JUNGERMANNIALES

Key to Genera of the Falkland Islands and Magellanian Zone of South America6

1. Leaf with dorsal lobe modified to form a water sac. Ventral merophytes absent (apical cell with 2

cutting faces); androecia and gynoecia on highly reduced, lateral-intercalary branches. Rhizoids scat-

tered [Pleuroziaceae, Eopleurozia, cf. p. 1 58]

1 . Leaf with the ventral lobe modified to form a water sac, or with a sac absent altogether; ventral

merophytes present, at least 1 cell row in width (apical cell with 3 cutting faces); androecia and

gynoecia various in position, but not on highly reduced lateral-intercalary branches (except in some

Heteroscyphus spp.) 2

2. Leaves of main stems with orientation (and usually insertion) incubous, at least at dorsal end of

insertion (the leaves at times transversely inserted and oriented in Lepicolea), the leaves usually

incubously shingled (but ventral portions sometimes modified to form a lobule or water sac);

rhizoids always restricted in origin, never scattered on stem, often lacking or rare 3

2. Leaves of main stems varying from transverse in insertion to succubously inserted and oriented,

to almost longitudinally (horizontally) oriented (never with ventral margin of leaf bearing, or

transformed into, a lobule or sac); rhizoids often copious, often scattered on ventral face of stem

9

3. Leaves divided into a larger dorsal lobe and a solitary, smaller ventral lobe, which is usually

saclike or pouchlike; branching exclusively lateral, the branches terminal or infra-axillary,

neither vegetative nor sexual branches ever ventral or axillary; underleaves 0-2 lobed or absent

4

6 This key is adapted with considerable modification from Schuster (1963a), with portions from Schuster (1965a,

1966a, 1980a) and Engel (1978).

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 5 1

3. Leaves various, but never with ventral base or lobe modified to form a flap or inflated sac;

branching various; underleaves always very large, varying from 2- to 4-lobed 7

4. Underleaves lacking; plants light to yellow green, rarely bronzed, never deep brown or reddish

Group I (p. 53)

4. Underleaves present and conspicuous; plants at times brownish or reddish 5

5. Underleaves unlobed and edentate; plants relatively vigorous, often bronzed or brownish to blackish

[Group II, p. 54]

5. Underleaves bilobed or at least emarginate 6

6. Lobuli, at least in large part, galeate, almost free from the dorsal lobe; plants often reddish or

brownish or dark green Group III (p. 54)

6. Lobuli not saccate or galeate, united for most of their length with dorsal lobe along an elongate

keel (the water sac formed partly by lobule, partly by the opposed portion of the lobe); plants

green to yellow green. Plants often small and delicate; perianth usually pentagonal, with a beak

Group IV (p. 54)

7. Plants with only lateral, terminal branching, without ventral branches and never with ventral flagella

or stolons; leaves divided for 0.6 or more their length; leaf and underleaf margins and segments often

ciliate or laciniate. Underleaves large and conspicuous Group V (p. 55)

7. Plants with branching not exclusively lateral and terminal; sometimes vegetative and/or sexual branches

partly or wholly ventral in origin and abbreviated; plants often with ventral stolons or flagella (at

least from older parts of plant); leaves rarely very deeply lobed, never with longly ciliate margins

8

8. Plants frequently isophyllous (both on main stems and branches), the underleaves similar in size

and usually in form to lateral leaves (except some species of Isotachis); branches usually few and

irregular, subfloral innovations excepted, all or in large part ventral and axillary, slender flagella

usually lacking; sexual branches never abbreviated and ventral. Lateral leaves 2-4-fid; leaf insertion

only weakly incubous, the orientation subtransverse Group VI (p. 55)

8. Plants normally somewhat to strongly anisophyllous, the underleaves differing in size (and usually

in form) from lateral leaves, at least on the branches; underleaves often (if lobed) with fewer lobes,

or lobes of different form than those of lateral leaves, or else more shallowly lobed; branches

(usually) in large part lateral, usually terminal, the plants often regularly 1-2-pinnately branched;

stolons, flagella, or rhizomes bearing vestigial leaves usually present; sexual branches nearly always

short, ventral Group VII (p. 55)

9. Stem with underleaves conspicuous throughout (not merely with a minute underleaf here and there),

even on sterile stems 10

9. Stem with underleaves either lacking on sterile stems, or minute, or merely small, scattered cilia or

laciniae, or groups of slime papillae, never conspicuous (exc. Lophozia hatcheri). Rhizoids almost

always scattered over ventral stem surface if present at all 15

10. Leaves sharply complicate bilobed, with a ± smaller (rarely subequal) dorsal lobe lying over a

larger ventral lobe; plants with a coelocaule or a fleshy marsupium Group VIII (p. 56)

10. Leaves not complicate bilobed if lobed at all, the lobes lying in nearly the same plane, or leaf

merely concave or ± naviculariform, never folded; plants with sporophyte protection various,

but mostly with a perianth 11

1 1 . Mature leaves (2)4-6(9)-lobed, often very deeply so, the lobe tips usually ciliiform or acuminate,

often the lobe margins with cilia or teeth; stem without a distinct, pellucid hyaloderm; underleaves

very large, ca. 0.5-0.9 the area of leaves, similarly lobed and/or ciliate; rhizoids usually sharply

restricted to underleaf bases Group IX (p. 57)

1 1 . Mature leaves undivided to 2(3-4)-lobed (occasionally with accessory small lobes or teeth, occasional

leaves deeply trifid), if leaves lobed, then the lobe margins entire or dentate, but not with long cilia;

stem sometimes with a hyaloderm; underleaves various, but for the most part markedly smaller

than leaves and differing from them in shape and form 12

12. Plants nearly or quite isophyllous: the underleaves similar in size and form to lateral leaves,

or nearly so; leaves erect to erect-appressed. Plants water-repellant. Leaves unistratose

Group X (p. 57)

1 2. Plants quite anisophyllous, or if ± isophyllous (Hygrolembidium, Pachyglossd) then with leaves

52 FIELDIANA: BOTANY

polystratose at least at base (except Chiloscyphus subg. Notholophocolea); leaves never erect-

appressed 13

13. Rhizoids usually common, scattered over ventral stem surface; underleaves usually ovate to tri-

angular, unlobed or obscurely bilobed, or large, bifid and/or ciliate; cells each with dense, conspicuous

papillae (except Cephaloziella dusenii where small or absent and A mphicephalozia, where sometimes

weak) Group XI (p. 57)

13. Rhizoids always restricted in origin, only at underleaf (sometimes also at leaf) bases, often largely

confined to reduced leaves and underleaves of stolons, flagella, or rhizomes, never scattered; cells

smooth or with scattered, small fine papillae (if, rarely, with coarser papillae, see Saccogynidium)

14

14. Stem often very soft, pellucid, with a cortex of large hyaline cells (hyaloderm) surrounding a

± small-celled medulla which is usually visible through the cortex by transmitted light (a

hyaloderm lacking in Kurzia and Hygrolembidium)', cells not collenchymatous, often hyaline,

large; perianth on ± abbreviated, intercalary usually ventral branches, long, triquetrous, often

tapered distally; androecia usually on short ventral-intercalary branches

Group XIV (p. 60)

14. Stem without a transparent, colorless hyaloderm: the cortical cells never conspicuously larger

than medullary cells, the medullary cells not visible through the cortex; cells various, often with

distinct trigones; perianth, if developed, usually on a ± elongated stem or branch, or on short

lateral (rarely on short ventral) branches, seldom tapered distally; androecia on lateral branches,

or if ventral (Saccogynidium) then plants with marsupia Group XII (p. 58)

15. Leaves ± sharply complicate-bilobed (except Blepharidophyllum densifolium, which has a broadly

rounded keel), transversely to succubously inserted and oriented; perianth dorsiventrally com-

pressed. Leaf margins (and/or apices) often ± denticulate to ciliate Group XIII (p. 59)

15. Leaves lobed or not, but if lobed, then never with lobes sharply bent over each other; perianth (if

present) not sharply dorsiventrally compressed 16

16. Stem soft textured, consisting of a conspicuous, hyaline cortex of enlarged cells (surrounding a

medulla of much smaller, ± thick-walled cells usually distinct by transmitted light, or medullary

cells also soft but large); leaves bilobed, with cells thin-walled and ± hyaline

Group XIV (p. 60)

16. Stem anatomy very various, but never with a hyaline cortex of large cells surrounding a medulla

of smaller, ± thick-walled cells; leaves various, not bilobed and with leptodermous, hyaline

cells 17

17. Leaves inserted ± transversely (at least in part of the dorsal half of the insertion) and transversely

oriented; leaves generally appearing ± pectinate when not densely appressed-imbricate. Leaves

bilobed (except some species of Herzogobryum). Plants with perianths, never with a marsupium

Group XV (p. 61)

17. Leaves obliquely inserted and ± succubously oriented, at least half of the insertion quite oblique

on stem to nearly horizontal. Leaves unlobed to short bifid, strongly bilobed only in Lophozia.

Plants with sporophyte protection various, at times with a marsupium 18

1 8. Plants with perianths terminal on leafy, main, or lateral stems; androecia intercalary or terminal

on leading stems; terminal branching common, absent in a few taxa . . . Group XVI (p. 61)

18. Plants without perianths, with pendent perigynia or with an erect fleshy, rigid shoot calyptra;

terminal branching absent. Leaves various, often entire or irregularly dentate; capsule wall 5-

9-stratose . Group XVII (p. 62)

GROUP I: Radulaceae, Lejeuneaceae (p.p., cf. also groups II, IV)

1 . Leaves with a wide J- or U-shaped insertion; rhizoids (where present) in fascicles from lobuli; ventral

merophytes 3-5 or more cells broad; perianth dorsiventrally compressed, the mouth wide, truncate

Radula (p. 1 58)

1. Leaves with a narrow transverse insertion; rhizoids in fascicles from stem; ventral merophytes 1 cell

broad; perianth (4)5-keeled, the mouth beaked 2

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 53

2. Branches of Lejeunea type, with a collar at the base; leaves never sharply dimorphic; inflated

lobules with hyaline papilla usually ± proximal to apical tooth, if ± entally displaced (often),

then rarely inserted on midline of tooth [Cololejeunea]

2. Branches of Aphanolejeunea type, without a collar; leaves sharply dimorphic: some with inflated

lobules, others with lobule vestigial and 1-2-celled; inflated lobes with hyaline papilla entally

displaced, at base of the 1-2-celled tooth along its ental midline, or displaced to distal side ....

.... [Aphanolejeunea}

"

GROUP II: Porellaceae, Lejeuneaceae (p.p., cf. also groups I, IV)

1. Lobule similar in shape to the underleaves, lying parallel to them, free from lobe, never inflated,

with margins often ventrally reflexed; elaters free, tapered at both ends; plants green (in ours) ....

[Porellaceae, Porella, cf. p. 161]

1 . Lobule differing in form and size from underleaves, lying normally parallel with posterior lobe margin

and broadly united with it, usually inflated; elaters attached to valves, the free end trumpetlike; plants

greenish brown to brownish to blackish 2

2. Stem with cortical cells thick-walled, not forming a hyalodermis; perianth 3-5(6)-keeled; oil-

bodies segmented [Archilejeunea]

2. Stem with cortical cells relatively thin-walled, forming a hyalodermis; perianth usually pluriplicate;

oil-bodies homogenous [Brachiolejeunea]

GROUP HI: Lepidolaenaceae, Jubulaceae

1. Underleaves not becoming galeate; plants with perianths, but without a coelocaule; elaters 1 -spiral;

leaf lobes entire Frullania (p. 162)

1 . Underleaves, at least of the ultimate branches, becoming galeate; plants with a large, variously

paraphyllose coelocaule and without a perianth, or with a coelocaule subtending a perianth; elaters

2-3-spiral; leaf lobes sometimes armed ; 2

2. Underleaves of main axis bifid, emarginate or (in Falkland species) undivided; main shoot com-

monly black brown; stems without paraphyllia Gackstroemia (p. 1 60)

2. Underleaves of main axis quadrifid; main shoot green to ± red; stems with paraphyllia

[Lepidogyna]

GROUP IV: Lejeuneaceae (p.p., cf. also groups I, II)

1 . Plants with segmentation pendulum-like, i.e., with 1 underleaf per lateral leaf; leaves highly specialized

with conversion of the lobule and lobe (or portion of it) to a complex water sac, the lobule closed

by a specialized movable valve. Plants with leaves erect, oriented away from the substratum

[Colura]

1. Plants with segmentation helical, i.e., with 1 underleaf per pair of lateral leaves; leaves without a

complex water sac and valve mechanism 2

2. Leaves sharply constricted at base, with a strongly abbreviated, ± transverse attachment to the

stem. Lobes narrow, obovate, elongated; lobule with a very narrow insertion formed by 1-2 cells,

obovate, tridentate distally along free margin; underleaves narrow, with filiform lobes which are

uniseriate for most of their length [Austrolejeunea]

2. Leaves with a wide, distinctly J- or U-shaped insertion, the dorsal lobe distinctly incubously

inserted 3

3. Hyaline papilla of lobule distal to apical tooth; leaves convex, often strongly so; plants firm,

often dull and opaque Cheilolejeunea (p. 164)

3. Hyaline papilla of lobule proximal to apical tooth; leaves flat to somewhat convex, rarely

strongly so; plants delicate, generally somewhat pellucid 4

54 FIELDIANA: BOTANY

4. Lobes blunt, acute to acuminate, narrowly ovate to lanceolate, 1.5-3 x as long as wide;

underleaves obdeltoid, with ± divergent lobes only 1-5 cells wide; plants small, remote-

leaved 5

5. Underleaf lobes setaceous to sharply acute, 1-2 cells broad, strongly divaricate

[Drepanolejeunea]

5. Underleaf lobes blunt to rounded, 4-5 cells broad usually, with blunt to angulate sinus

Harpalejeunea (p. 1 66)

4. Lobes blunt to rounded, orbicular-ovate or ovate-falcate usually, less than 1.5 x as long

as wide. Leaf trigones absent or small Lejeunea (p. 1 66)

GROUP V: Lepicoleaceae, Ptilidiaceae

1. Leaves usually ± symmetrically (3)4-parted, the lobes entire or dentate to ciliate near base only;

underleaves lobed as in leaves; plants without perianths but with a fleshy coelocaule densely covered

with scales and paraphyllia; plants with subfloral innovations Lepicolea (p. 63)

1. Leaves very asymmetrically lobed and 2-4(5)-parted, with ciliate to dentate lobes; underleaves bi-

lobed; plants with a perianth; plants without subfloral innovations

[Ptilidiaceae, Ptilidium, cf. p. 1 58]

GROUP VI: Vetaformaceae, Herbertaceae, Balantiopsaceae (p.p., cf. also group VIII)

1. Leaves distinctly vitiate, a "vein" of longer cells running into each lobe; leaves deeply (0.5-0.75)

bifid; cells with coarse trigones. Leaf lobes slender, often falcate. Perianths present; bracteolar an-

theridia present [Herbertus]

1. Leaves non vitiate, the lobe cells not conspicuously elongated, leaves 0.2-0.5 bifid-trifid; cells wilh

irigones absenl lo small. Perianlhs absenl or presenl; bracleolar antheridia absent or present . . 2

2. Leaf and underleaf margins enlire or dentale in basal portion, sparsely armed above; planls usually

wilhout conspicuous pigmentalion, al mosl pale brown; male bracleoles wilh anlheridia. Perianlhs

presenl or absenl 3

3. Leaves and underleaves 2-3-lobed for 0.4-0.6 Iheir lenglh, ihe lobes broad al base, iriangular;

perianlh dislincl Triandrophyllum (p. 65)

3. Leaves and underleaves (2)3-4-lobed for 0.6-0.75 Iheir lenglh, ihe lobes linear-lanceolale, only

(3)4-5 cells broad al base; perianlh absenl, a clavale coelocaule presenl

[Velaformaceae, Vetaforma, cf. p. 63]

2. Leaf and underleaf margins wilh leelh, when presenl, often in apical portion, nol predominanlly

in basal portion; planls usually wilh reddish and/or cheslnul brown lo fuscus pigmenlalion; male

bracleoles wilhoul anlheridia. Perigynium presenl, firm, fleshy, with a vestigial perianth al ils lip

Isotachis (p. 1 57)

GROUP VII: Lepidoziaceae (p.p., cf. also group XIV), Calypogeiaceae

1. Leaves deeply (2)4-6-lobed for usually 0.5 or more Ihe leaf lenglh; planls regularly ± pinnalely

branched 2

2. Leaves wilh lamina (in ours) 0.5-4 cells high; leaf lobes uniseriale Ihroughoul, filiform and lerele,

al mosl 2 cells wide in basal cell lier; slem wilh a conspicuous hyaloderm layer, formed of large

cells wilh walls lillle or not thickened conlrasled lo Ihe smaller medullary cells often wilh ihick

walls; leaves usually symmelrical; leaf cells usually large, hyaline, Ihin- or slighlly ihick-walled,

Irigones lacking Telaranea (p. 78)

2. Leaves wilh a lamina many cells high; leaf lobes iriangular, never filiform; slem wilhoul a hy-

aloderm layer (cortical cells ± thick walled, not conspicuously larger than medullary cells); leaves

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 55

usually conspicuously asymmetric (dorsal margin longer and/or arched); leaf cells usually rigid,

often with thick walls and guttulate lumina Lepidozia (p. 72)

Leaves with apices edentate or 2-3-dentate-lobate, never deeply lobed, the sinuses rarely descending

more than 0.25 the leaf length (and always less than 0.5 the leaf length); branching various, but not

regularly pinnate 3

3. Leaves firm, the cells thick-walled or with distinct trigones; leaf apices usually (0)2-3 -dentate;

lateral terminal branches frequent to common; ventral intercalary stolons freely developed; stem

firm-textured; plants with perianths _ 4

4. Stolons intercalary in origin, from axils of underleaves; stem without a hyaloderm, the cortical

cells in many rows; leaf apices typically 2-3 -dentate; pseudodichotomous branching present:

Frullania-lype branches only from one side of axis and often subequal to main shoot in vigor

[Bazzania]

4. Stolons terminal in origin, replacing one-half of an underleaf; stem with a hyaloderm, at least

the branches with only 7 rows of cortical cells; leaf apices 0-2 dentate-lobate; pseudodichot-

omous branching lacking, Frullania-type branches arising from both left- and right-hand sides

of axis [Acromastigum]

3. Leaves delicate, subhyaline, leptodermous (or with small trigones); leaves entire or finely bidentate

at apex; lateral terminal branches few or absent; ventral intercalary stolons lacking; stem soft-

textured; plants with marsupia, without perianths. Branching normally all ventral intercalary . .

[Calypogeiaceae, Calypogeia, cf. p. 82]

GROUP VIII: Balantiopsaceae (p.p., cf. also group VI), Schistochilaceae

1. Apex of dorsal lobe usually suberect, lying at an angle to axis of ventral lobe, the lobes ± divergent;

keel between lobes wingless; cells of leaf lobe oriented in ± distinct tiers; sporophyte (unknown for

Anisotachis) in a marsupium 2

2. Leaf lobes bifid, rarely subequal, the apices without a border Balantiopsis (p. 154)

2. Leaf lobes undivided, truncate or emarginate, subequal, with apices bordered by 1-several trans-

verse rows of cells [Anisotachis]

1 . Apex of dorsal lobe oriented towards apex of ventral lobe; keel between lobes usually with a broad

wing; cells of leaf lobes irregularly arranged, not in tiers; sporophyte in a terminal coelocaule lying

in the axis of the stem. Leaf lobes undivided, never bifid 3

3. Rhizoids colorless, usually fasciculate, the apices often digitiform and copiously septate, the

branches becoming polyseriate; plants never with wall pigments of any part of gametophyte; cells

of leaves and underleaves frequently showing secondary septation (asexual reproduction via re-

generation); leaves partially to largely polystratose (at least basally and in keelar region), without

specialized dentition, the ventral lobes never lamellate or with cellular processes; branches nor-

mally of lateral-intercalary or Radula type (in P. reflexistipula with both Frullania and Microle-

pidozia-lype branches) Pachyschistochila (p. 1 53)

3. Rhizoids distally at least, magenta to claret red, always scattered, the apices, if ramified, irregularly

branched, the branches often juxtaposed and/or anastomosing, usually sparingly if at all septate;

plants with pigmentation of remainder of gametophyte present in some taxa; cells of foliar ap-

pendages never secondarily septate; leaves (aside from the keelar area in Subg. Eoschistochild)

uniformly 1 -stratose, often with specialized dentition (setae, cilia, etc.), often with lamellae and/

or processes of ventral lobes; many taxa retaining Frullania-lype branches (but none with Micro-

lepidozia-iype branches) 4

4. Radial symmetry in vegetative, gynoecial, and androecial regions; elaters slender and tortuous,

1 -spiral; leaves unwinged, noncomplanate [Pleurocladopsis]

4. Radial symmetry lacking (in Subg. Austroschistochila subsymmetric in gynoecia alone); elaters

usually stout and rigid, always 2-spiral; leaves winged, sharply complanate 5

5. Underleaves and $ and 9 bracteoles distinct Schistochila (p. 1 52)

5. Underleaves and <5 and 9 bracteoles completely lacking [Paraschistochila, p. 152]

56 FIELDIANA: BOTANY

GROUP IX: Trichocoleaceae

1. Leaves appearing as a mass of interwoven cilia such that the lamina is obscured; stems paraphyllose;

plants with a fleshy, hispid coelocaule, a perianth low or lacking; branching copious, exclusively of

Frullania type; plants green, without brown pigmentation [Trichocolea]

1. Leaves without interwoven cilia and/or segments (or if so, e.g., in Temnoma pilosum, then plants

with brown pigmentation), with at least part of lamina conspicuous; stems not paraphyllose; plants

with a conspicuous perianth, at most with a coelocaule precursor (in Temnoma spp.); branching

sparing or sporadic, not exclusively of Frullania type, ventral-intercalary branches at least occasionally

present; plants often with brownish pigmentation 2

2. Leaves and underleaves bifid or bisbifid. Leaf lobes always entire and several cells broad (even

of subfloral leaves and bracts); plants perfectly isophyllpus v 3

3. Leaves and underleaves bifid. Leaves unistratose [Isophyllaria]

3. Leaves and underleaves (except on juvenile shoot sectors) normally bisbifid 4

4. Leaves polystratose, the lobes rigid, polystratose, ± terete; leaves with knotlike swellings

at the base of the median and often also the lateral sinuses; branching exclusively intercalary;

perianth mouth short lobate-dentate [Herzogiaria]

4. Leaves unistratose throughout, the lobes thin, flat; leaves devoid of knotlike swellings;

branching all or predominantly terminal, of Frullania type; perianth mouth crenulate . . .

[Pseudolepicolea]

2. Leaves of mature shoots equally quadrifid. Leaf lobes armed or entire; plants anisophyllous to

subisophyllous 5

5. Cells of leaf disc with elongated, thin- to equally thick-walled cells 6

6. Lobes of mature leaves (usually of vegetative shoot sectors, at least in and below gynoecia)

with opposed, sharp teeth or cilia usually on both disc margins and lobes, rarely on one

only; perichaetial bracts freely spinose-dentate to copiously ciliate; perianth wide at open

mouth Temnoma (p. 67)

6. Lobes of mature leaves without cilia or teeth (or, rarely, with an isolated tooth on one or

both margins of the disc); perichaetial bracts without trace of teeth or cilia; perianth closely

contracted to the narrow mouth. Leaves with segment apices setaceous

Archeochaete (p. 67)

5. Cells of leaf disc with well-developed trigones. Leaves and underleaves edentate; leaves 3—4-

lobed for 0.6-0.75 their length, the lobes acuminate, uniseriate for much of their length; subfloral

bracts usually larger than innermost bracts and with disc margins ciliate or dentate

[Archeophylla]

GROUP X: Antheliaceae

1 . Plants greenish but with age often distinctly whitish; branching normally terminal lateral, of Frullania

type, rarely ventral intercalary; stem cortical cells forming a weak hyaloderm, the cells somewhat

swollen and bulging, at least their free walls thin. Oil-bodies lacking [Anthelia. cf. p. 91]

1 . Plants red brown; branching strictly intercalary, both ventral and lateral; stem without a differentiated

cortex, the cortical (and medullary) cells very thick walled. Oil-bodies unknown

(Grollea, cf. p. 91]

GROUP XI: Lophoziaceae (p.p. min., cf. also groups XV, XVI), Acrobolbaceae (p.p. min., cf. also group

XVII), and Cephaloziellaceae (cf. also group XV)

1 . Underleaves large, ciliate and/or bifid; without stolons or flagella; plants vigorous (shoots to 3-5 mm

wide), usually ± brownish. Cells with small or moderately coarse trigones; cuticle with high, coarse,

hemispherical papillae; perianth apex twisted; bracts of innermost series very deeply lacerated, smaller

than other bracts or leaves; capsule wall 7-stratose Roivainenia (p. 97)

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 57

Underleaves small or minute, eciliate, usually unlobed; often with flagella or stolons; plants small

(shoots to 0.7-2 mm wide) 2

2. Leaves succubously inserted throughout, large (leaf width much greater than stem width); perianth

lacking, plants with marsupia Acrobolbus (p. 148)

2. Leaves with at least dorsal half transversely inserted (occasionally dorsal half subsuccubous in C.

scabrellus), small, distant; perianth present. Plants with Cephaloziella-\ike facies 3

3. Stolons and flagella present; leaf cell trigones distinct; plants often with brownish to reddish

secondary pigments; perianth mouth lobulate-laciniate Cephalolobus (p. 95)

3. Stolons and flagella lacking; leaf cell trigones minute; plants green, without secondary pigments;

perianth mouth obscurely shallowly lobulate v [Amphicephalozia]

GROUP XII: Geocalycaceae

1 . Plants producing marsupia, perianths absent; sexual branches always on abbreviated, small, ventral-

intercalary branches. Leaves ovate, undivided or bidentate (occasionally short bifid in S. vasculoswri),

the cuticle very densely papillose; underleaves 0.5-0.75 bifid (in our species), free from and much

smaller than leaves; plants dull, opaque, gray or light green plants developing (often) some brownish

pigmentation Saccogynidium (p. 142)

1 . Plants producing perianths; sexual branches on apices of unmodified stems or short lateral branches,

rarely on short ventral branches 2

2. Perianths ± strongly laterally compressed, basically 2-lipped (the ventral lobe strongly reduced

or vestigial); rhizoids often tending to spread away from underleaf bases or (Pedinophyllopsis) ±

scattered; plants often ± brownish, often intensely so 3

3. Dorsal-intercalary (Andrewsianthus-type) branching present; leaves (marginal sectors aside) 2-

3-stratose; flagelliform branches present. Underleaves very small, narrower than stem, bifid

Evansianthus (p. 1 29)

3. Dorsal-intercalary branching lacking; leaves unistratose throughout; flagelliform branches ab-

sent. Underleaves variable, at times small and inconspicuous, at times equal to the leaves,

undivided or bifid 4

4. Plants lacking secondary pigments, strikingly laterally compressed and complanate, the

leaves strongly erect, often appressed to one another dorsally; underleaves minute, free

from leaves, with (1)2 juxtaposed long ciliiform segments; rhizoids in large part scattered;

perianths barely exserted beyond bracts; male bracts with paraphyllia. Leaves unlobed and

entire Pedinophyllopsis (p. 141)

4. Plants often with red brown pigments, basically somewhat to strongly dorsiventrally flat-

tened (if leaves concave and dorsally assurgent, then dorsiventral complanation ill-defined,

but plants not laterally compressed), the leaves never laterally appressed; underleaves usually

large, typically ± decurrent and often connate with 1 or both leaves, the segments usually

at most laciniiform, never long ciliiform; rhizoids confined to underleaf bases (from there

sometimes spreading somewhat away from underleaf base); perianths usually distinctly

exserted beyond bracts; male bracts lacking paraphyllia. Leaves 2-3- or more dentate to

lobed, or unlobed and entire Leptoscyphus (p. 1 33)

2. Perianth trigonous to trigonous-inflated, the mouth equally or subequally trilobate; rhizoids re-

stricted to underleaf bases; plants normally lacking brown pigments 5

5. Androecia and gynoecia all or predominantly on unspecialized leafy shoots; androecial position

generalized: terminal or intercalary on main axes or branches indeterminate in length, the

branches normally bearing at least some sterile leaves (either at branch base or distally). Leaf

cells generally leptodermous, without trigones or with trigones small, medium, or absent;

underleaves (in ours) not strongly connate on both sides 6

6. Plants stoloniferous; leaves transverse to subsuccubously oriented 7

7. Plants nearly isophyllous, the underleaves conspicuous and rigidly patent; leaves and

underleaves polystratose at least at the base; female bracteoles ± similar in size to bracts

Pachyglossa (p. 1 37)

58 FIELDIANA: BOTANY

7. Plants clearly anisophyllous, the underleaves highly reduced, inconspicuous, appressed

to moderately spreading; leaves unistratose throughout; female bracteoles reduced, in-

conspicuous, hardly modified from underleaves [Hepatostolonophora]

6. Plants usually lacking stolons; leaves mostly strongly succubously oriented 8

8. Leaves moderately to deeply adaxially concave, with apices incurved 9

9. Leaves strongly succubous, the insertion a long, strongly oblique line 10

10. Plants clearly anisophyllous; underleaves plane, convex, or sometimes cucullate

(ventral view); leaves usually conchiform concave, occasionally moderately

adaxially concave. Leaf apices mostly undivided, in a few species 2- or 2-3-

lobed Clasmatocolea (p. 1 23)

10. Plants subisophyllous to isophyllous; underleaves occasionally strongly concave

(recurved) (ventral view), particularly toward shoot apices; leaves sporadically

weakly to rather strongly convex. Leaf apices undivided, entire or 1 -dentate . . .

[Concave-leaved phases of Chiloscyphus subg. Notholophocolea]

9. Leaves transverse, subsuccubous or subincubous, the insertion narrow 11

11. Leaf margins (and abaxial leaf faces) crenulate to mamillate with domelike cell

wall protuberances; leaves Cephaloziella-\ike, at most slightly concave, without

an incurved ventral margin, normally lacking any trace of teeth; leaf insertion

transverse to weakly succubous Pigafettoa (p. 141)

1 1 . Leaf margins (and abaxial leaf faces) not crenulate to mamillate with domelike

cell wall protuberances; leaves ± Nowellia-like, strongly concave, the ventral

margin incurved and accentuating and defining a deep pocket-like concavity in

ventral portion of leaf, the leaves with accessory teeth; leaf insertion transverse

to weakly incubous. Valdivian [Xenocephalozia]

8. Leaves convex, rarely plane, with apices decurved or deflexed 12

1 2. Leaves with lobes and marginal teeth caducous, often giving leaf apices a ragged

appearance; leaf apices often with accessory teeth and laciniae

Leptophyllopsis (p. 1 32)

1 2. Leaves entire or if lobed, then with lobes persistent; leaves with marginal teeth, if

present, persistent; leaf apices never with a ragged appearance and not with accessory

laciniae Chiloscyphus (p. 1 1 1)

5. Androecia and gynoecia on specialized, abbreviated, intercalary branches; androecia (abnormal

exceptions aside) on determinate, narrow, compactly spicate, lateral-intercalary branches that

never bear normal leaves, and never innovate distally. Leaf cells often with coarse to strongly

nodose or triradiate trigones; underleaves often strongly connate on both sides

Heteroscyphus (p. 1 32)

GROUP XIII: Blepharidophyllaceae, Scapaniaceae

1 . Leaf lobes each short bifid; branching predominantly terminal, of Frullania type; rhizoids in small

fascicles (often present only on small plants) from the stem at the ventral bases of leaves

Blepharidophyllum (p. 1 08)

1. Leaf lobes undivided, never bifid; branching predominantly lateral-intercalary; rhizoids scattered

over ventral surface of stem, never in fascicles 2

2. Leaves unequally complicate bilobed, the dorsal lobe smaller than the ventral (except in ours,

where subequal); antheridial stalks uniseriate 3

3. Gemmae angulate to polyhedral to stellate; dorsal and ventral lobes oriented in differing

directions; median lobe cells with at most small trigones; perianth weakly flattened, distinctly

plicate, contracted toward the mouth Diplophyllum (p. 1 06)

3. Gemmae smooth, ovoid to ellipsoid; dorsal and ventral lobes similarly oriented; median lobe

cells with trigones; perianth normally strongly flattened, eplicate, truncate at mouth

[Scapania]

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 59

2. Leaves equally complicate-bilobed or nearly so; antheridial stalks biseriate. Dorsal lobe dorsally

recurved . [Krunodiplophyllum]

GROUP XIV: Cephaloziaceae, Lepidoziaceae (p.p., cf. also group VII)

1 . Rhizoids scattered on ventral surface of axis; vegetative shoots without distinct underleaves; leaves

bifid, never 3-4-lobed; cells lacking oil-bodies; seta with 8 epidermal + 4 inner cell rows; plants

sometimes with purple or red pigmentation. (Cephaloziaceae) 2

2. Leaf insertion transverse at least dorsally, extending to stem midline; pigmentation, when present,

purplish or brownish; leaves ± canaliculate, piano-distichous; branching plastic, Frullania-lype

branching always freely developed in addition to lateral-intercalary and ventral-intercalary branch-

es; gynoecia terminal on leading axes Metahygrobiella (p. 89)

2. Leaf insertion oblique throughout, the dorsal end oblique to almost longitudinal; pigmentation

never purplish, usually absent; leaves never piano-distichous, flat to concave but not canaliculate;

branching normally of ventral intercalary type, Frullania-type branches rare (but present in C.

badid)\ gynoecia normally on short, ventral branches which occasionally are elongated

Cephalozia (p. 89)

1 . Rhizoids (often sparingly present) restricted to bases of underleaves; vegetative shoots with under-

leaves distinct throughout; leaves often 3-4(or more)-lobed; seta with 8 or 16 large epidermal cell

rows and many inner cell rows that are of much inferior diameter; plants without purple or red

pigmentation. (Lepidoziaceae, p.p.) 3

3. Branching predominantly or nearly exclusively terminal, plastic, ofFrullania plus Microlepidozia

and/or Acromastigum types 4

4. Terminal branches plastic, i.e., ofFrullania-, Microlepidozia (and in K. mollis, Acromastigum-)

type, usually of Frullania type on one side of axis and Microlepidozia type on opposite side

of axis; underleaves frequently with 1 segment shorter than others; stem with hyaloderm absent

or at best weakly developed. Plants usually firm, the leaves transversely to succubously inserted,

usually distant, usually 3-4(6)-lobed Kurzia (p. 71)

4. Terminal branches predominantly ofFrullania type, Acromastigum type sporadic to common,

rarely of Microlepidozia type; underleaves not regularly possessing one segment shorter than

others; stem with a well-developed hyaloderm. Plants commonly soft and flexuous, the leaves

(in ours) transversely inserted, usually 2(-3)-dentate-lobate Paracromastigum7 (p. 76)

3. Branching nearly exclusively intercalary, terminal branches absent, at most rare or sporadic . . 5

5. Leaves bifid, unistratose throughout 6

6. Leaves bifid to 0.5 or more; secondary pigments lacking, plants green; cortical cells in 6

rows; terminal branching, when present, of Frullania and Microlepidozia types

Hyalolepidozia

6. Leaves 2(3)-dentate or if lobate, then at most divided to 0.3; secondary pigments present,

plants brown; cortical cells in 7-8 rows; terminal branching lacking

Paracromastigum tristanianum

5. Leaves unlobed or 3-4(6) dentate to lobate, never bifid, polystratose toward the base in Amer-

ican sector species 7

7. Leaves 3—4(6) dentate to lobate; leaf cuticle smooth or at most minutely roughened; stem

hyaloderm present; androecia on leading shoots; seta (where known) with 8 rows of epi-

dermal cells; capsule wall (where known) 2-stratose Pseudocephalozia (p. 78)

7. Leaves unlobed; leaf cuticle often papillose; stem hyaloderm absent; androecia on short,

spicate, basal, intercalary branches; seta with 10-20 rows of epidermal cells; capsule wall

3-5-stratose Hygrolembidium (p. 69)

7 Paracromastigum at times may lack terminal branching, and for that reason the genus is also keyed under the

second half of couplet 3.

60 FIELDIANA: BOTANY

GROUP XV: Cephaloziellaceae (cf. also group XI), Gymnomitriaceae, Lophoziaceae (p.p., cf. also groups

XI, XVI)

1. Leaves usually bilobed to 0.35-0.5 their length; leaf margins not bordered by 1-several rows of

hyaline cells 2

2. Perianth not situated at the apex of a fleshy tube; a perigynium lacking; plants without stolons or

flagella. Cells with trigones variable: at times lacking, at times coarse; cuticle often coarsely

papillose 3

3. Plants minute or small, often wiry; leaves remote, bilobed to 0.5 or more their length. Cells

very small, equally thick walled, lacking trigones; perianth mouth usually crenulate 4

4. Leaf lobes plane, 2-10 cells wide at base; leaves bifid to nearly 0.75 their length; seta

normally with 4 epidermal cell rows and 4 internal cell rows; plants minute, the shoots

usually less than 350 nm wide Cephaloziella (p. 90)

4. Leaf lobes sulcate, 9-28 cells wide at base; leaves bifid to nearly the base; seta with 8(9)

epidermal cell rows and (3)4-14 internal cell rows; plants small, the shoots greater than

400 nm wide [Allisoniella]

3. Plants larger, not wiry; leaves bilobed to ca. 0.3-0.45(0.5) their length. Leaf lobes 8 or more

cells broad at base 5

5. Leaves strongly dorsally secund, the dorsal end of the line of insertion arcuate and often

± decurrent along dorsal midline of stem. Cells (in Falkland species) with coarse, conspic-

uous trigones and thin to sinuous intervening walls; branching variable, terminal, Fmllania

type in some taxa, ventral-intercalary in some taxa, but if lateral intercalary then never

arising from near the dorsal end of leaf or dorsal side of axis (Andrewsianthus-lype)

Anastrophyllum (p. 92)

5. Leaves laterally patent and pectinate-distichous, not dorsally secund, the insertion dorsally

transverse, not decurrent. Branching strictly intercalary 6

6. Leaf cuticle with conspicuous, very coarse, hyaline papillae; branching apparently strictly

lateral intercalary, Andrewsianthus-lype branches lacking 7

7. Underleaves lacking; leaves large, stiffly, pectinately patent, with gibbous sinuses and

reflexed margins; gynoecium without bracteole; perianth weakly plicate distally, the

mouth crenulate; autoecious Acrolophozia (p. 103)

7. Underleaves small and ciliiform to large and ovate-lanceolate; leaves small, at least

basally suberect, ± concave but not folded, the lobe margins not reflexed; gynoecium

with large bracteole; perianth ± strongly plicate and lobulate-laciniate at mouth;

dioecious Cephalolobus (p. 95)

6. Leaf cuticle (in Falkland species) smooth, at most rather coarsely striate; branches of

Andrewsianthus type, arising from near the dorsal base of the leaf, occasionally appearing

to arise from dorsal side of axis. Branches and main axis frequently becoming positively

geotropic Andrewsianthus (p. 92)

2. Perianth included within the connivent or suberect bracts, associated with a distinct perigynium

that subtends the joint insertion of bracts and perianth; plants with stolons or flagella. Cells with

coarse trigones; cuticle smooth [Marsupella]

1. Leaves unlobed or bifid to 0.2; leaf margins often with 1-several rows of hyaline cells forming a

border. Leaves erect or erect-appressed, cuticle smooth or finely papillose

Herzogobryum (p. 104)

GROUP XVI: Plagiochilaceae, Lophoziaceae (p.p., cf. also groups XI, XV), Jungermanniaceae

1. Perianth laterally sharply compressed, wide and truncate at mouth. Vegetative branches lateral in-

tercalary and/or terminal and ofFrullania type, never ventral intercalary. Leaves with dorsal margin

deflexed to form a convex fold or cnemis; rhizoids scattered; anthocyanin pigments lacking

Plagiochila (p. 143)

1 . Perianths terete below, pluriplicate and contracted to the mouth 2

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 61

2. Rhizoids fasciculate, at ventral end of leaf; leaves opposite, connate at least ventrally. Bracts and

bracteole normally united; plants often with reddish to purplish anthocyanin pigments

[Syzygiella]

2. Rhizoids scattered, often in dense mats on ventral side of stem; leaves alternate, free ventrally.

Bracts and bracteole united or free 3

3. Leaves consistently 2^4-lobed. Underleaves laciniate to ciliate or reduced or absent .... 4

3. Leaves unlobed, usually rounded at apex. Underleaves reduced and inconspicuous 5

4. Ventral leaf margin frequently slightly to strongly reflexed, leaves short bifid; underleaves

present, toward the stem apices and only of a few cells (in Falkland species)

Nothostrepta (p. 96)

4. Ventral leaf margin plane, not reflexed, leaves 2-4-lobed, often deeply so; underleaves,

when present, may be large and conspicuous (ciliate in the Falkland species, L. hatcheri)

Lophozia (p. 96)

5. Involucral bracts unlobed and edentate; bracteole lacking; plants soft textured, usually

prostrate growing, small in size; antheridial bracts (l)2-3-androus

Jungermannia (p. 1 02)

5. Involucral bracts ± lobed and/or dentate, ± smaller than leaves; bracteole usually

present; plants ± firm, usually erect growing, often robust; antheridial bracts monandrous

6

6. Stolons or flagellae regularly produced; terminal branching completely absent. Brac-

teoles nearly as long as bracts; perianth mouth entire to weakly crenulate; plants

often greenish to blackish brown; (455)610-700 m in Falklands

Cryptochila (p. 98)

6. Stolons or flagella only rarely and sporadically produced, mostly absent; terminal

branching present, but sometimes only occasional. Gynoecium as in Cryptochila;

plants often red or magenta or red brown; 75-455(685) m in Falklands

Jamesoniella (p. 99)

GROUP XVII: Adelanthaceae, Acrobolbaceae

1 . Gynoecia and androecia on reduced, abbreviated, basal, intercalary branches; gynoecia producing an

erect, massive shoot calyptra without or (in some taxa of Adelanthus) with a perianth; leaf cuticle

smooth. Leaves undivided; plants erect, rigid 2

2. Stem 20-45 cells in diameter, with a collapsing hyaloderm (thus scabrous with age); dorsal leaf

margin 4-5 cells thick (in Falkland species); plants green, without secondary pigmentation. Sub-

hyalodermal cells in 2-4 layers of thick- walled cells Wettsteinia (p. 88)

2. Stem 8-15 cells in diameter, without a collapsing hyaloderm, smooth, never roughened; leaves

unistratose throughout; plants green to red brown Adelanthus (p. 82)

1 . Gynoecia and androecia on terminal or leading leafy shoots (except some populations of Tylimanthus,

where on short, basal, intercalary branches); gynoecia producing a marsupium; leaf cuticle often

conspicuously roughened. Leaves undivided or 2-4-lobed; plants erect or appressed; bracts of in-

nermost series smaller than lower bracts and often reduced to laciniiform projections; capsule tip

beaked 3

3. Leaves unlobed, leaf margins entire; marsupium slenderly cylindrical, deeply penetrating sub-

stratum; unfertilized archegonia carried downward with marsupial canal, inserted on surface and

base of free calyptra. Plants closely prostrate Lethocolea (p. 149)

3. Leaves (in Falkland species) 2-4 lobed (or if undivided, then with leaf margins dentate to lobate),

leaf margins entire-den tate-lobate; marsupium ± conoidal to wide cylindrical in shape, not deeply

penetrating substratum; unfertilized archegonia remaining external to and at summit of marsupium

4

4. Leafy shoots ± ascending, not adhering to substratum by rhizoids. Gynoecia and androecia

variable in position, acrotonic or basitonic; plants a clear, translucent to opaque green, often

whitish when dead Tylimanthus (p. 151)

62 FIELDIANA: BOTANY

4. Leafy shoots ± creeping, often entire axis, including shoot tips, adhering to the substratum by

rhizoids 5

5. Leaves (in ours) bilobed; cuticle coarsely papillose; rhizoids few, arising from ventral leaf

bases, never in dense, decurrent bundles on stem; plants without gemmae; antheridia sol-

itary; stolons and flagellae common Acrobolbus (p. 148)

5. Leaves 3-5 lobed; cuticle smooth; rhizoids dense, decurrent along stem in fascicles and

forming a ventral mat; plants gemmiparous; antheridia 1-10 per bract; stolons and flagellae

lacking Austrolophozia (p. 149)

Family VETAFORMACEAE

Vetaformaceae Fulf. & J. Tayl., Nova Hedwigia

1:405. 1960.

A monotypic family represented by Vetaforma

dusenii (Steph.) Fulf. & J. Tayl. of southern South

America, but not the Falklands. For details see

Fulford (1963b) and Hassel de Menendez and So-

lari(1975).

Family LEPICOLEACEAE

Lepicoleaceae Schust., Nova Hedwigia 5: 27. 31

Jan. 1963. Schust. ex Fulf. in Fulford, Mem.

New York Hot. Card. 1 1 : 30. 1 5 March 1 963.

Schust., Rev. Bryol. Lichenol. 26: 126. 1957,

nom. inval.

A family of one genus with ca. 1 0 species. Lep-

icolea occurs in temperate areas of South America,

Africa, and Australasia and extends to the North-

ern Hemisphere in both Asia and Latin America.

Lepicolea

Lepicolea Dum., Recueil Observ. Jungerm. 20.

1835.

Key to Falkland Islands Species of Lepicolea

1. Leaf segments ending in a uniseriate tip of a few to several elongate, noncaducous cells, the 2nd, 3rd,

and 4th cells from tip elongate, the tip cell not a slime papilla; oil-bodies hyaline, 4-7 per cell (fide

Inoue, 1 984) L. ochroleuca

1 . Leaf segments ending in a uniseriate tip of a few-celled row of short (subquadrate to short-rectangular)

cells which are often caducous; the 3rd and 4th cells from the tip short, the tip cell a slime papilla;

oil-bodies grayish, 1-3(4) per cell L. rigida

Lepicolea ochroleuca (Spreng.) Spruce. Figure 23.

Jungermannia ochroleuca Spreng., Syst. Veg. 4(2): 325.

1827. Sendtnera ochroleuca (Spreng.) Nees in G.

L. & N., Syn. Hep. 240. 1845. Leperoma och-

roleuca (Spreng.) Mitt, in Hook, f., Handb. New

Zealand Fl. 2: 754. 1867. Herbertia ochroleuca

(Spreng.) Trev., Mem. Reale 1st. Lomb. Sci. Lett.

Ill, 4: 397. 1877. Lepicolea ochroleuca (Spreng.)

Spruce, Trans. & Proc. Hot. Soc. Edinburgh 15:

345. 1885. Original material: South Africa, Cape

of Good Hope, Ecklon s.n. (NY!, STR, cited in

Scott, 1960).

Jungermannia hirsuta Nees ex Hook. f. & Tayl., Lon-

don J. Bot. 3: 289 (in errore pro 389), 475. 1844,

nom. nud.

Sendtnera ochroleuca 0 mexicana Gott., Kongel.

Danske Vidensk. Selsk. Naturvidensk. Math. Afh.

II, 6: 140. 1863, syn. fide Scott (1960).

REMARKS— This is the first authentic report of

Lepicolea ochroleuca from the Falkland Islands,

as all previous reports (Gottsche et al., 1845 as

Sendtnera; Hooker & Taylor, 1844, Taylor &

Hooker, 1 847 as Jungermannia hirsuta) were based

upon a Hooker collection which is actually L. rig-

ida.

ECOLOGY— Rare in the Falklands, where it is

known from only three stations. The Mt. Kent

plant occurred at the summit, 455 m, on wet rock

walls where only a few colonies were observed.

The species also was gathered deep in a cool, cave-

like hollow of a rock outcrop in a dwarf shrub

heath (180 m). Here, as in the above locale, the

plants (again, only a few) grew on an extensive,

flattish area of the outcrop. Longton and Smith

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

63

23

FIG. 23. Distribution of Lepi-

colea ochroleuca (Spreng.) Spruce,

plus Honduras north to Mexico.

FIG. 24. Distribution of Lepi-

colea rigida (De Not.) Scott, plus

South Georgia. Open circle indi-

cates precise locality unknown.

64

FIELDIANA: BOTANY

encountered the species at 1 85 m on damp ground

among boulders on the south face of Mt. William.

Lepicolea ochroleuca is common in the wet Ma-

gellanian moorland, where it may form extensive

mounds over bark. It is also quite common on

Nothofagus bark in the Valdivian region. The

species appears to require a high rainfall and cool

temperatures, the absence of the former (as well

as the bark habitat) presumably accounts for the

rarity of the taxon in the Falklands. The Port Stan-

ley region has cooler temperatures and a higher

rainfall than other Falkland regions, a point which

almost undoubtedly accounts for the presence of

L. ochroleuca in this part of the Falklands.

PHYTOGEOGRAPHY — Amphiatlantic — South Af-

rica; Tristan da Cunha; Falkland Is.; Tierra del

Fuego; Patagonian Channels; Valdivian region

(West Patagonia north to 39°16'S, Andean Pata-

gonia at P. N. Nahuel Huapi); Juan Fernandez Is.;

also known from Brazil (Stephani, 1909; Lor-

scheitter, 1973).

Reported from Bolivia (Spruce, 1890), Hon-

duras and Guatemala (Fulford, 1963b; Hassel de

Menendez & Solari, 1976), and Mexico (Scott,

1960; Fulford, 1963b; Hassel de Menendez & So-

lari, 1 975). The report from India in Hooker (1 867)

is regarded as erroneous. The New Zealand reports

are either L. attenuata or L. scolopendra, and that

from Tasmania is likely L. scolopendra (fide ci-

tations in Scott, 1960) (Fig. 23). Hassel de Me-

nendez and Solari (1975, p. 37) include New Zea-

land, Auckland Island, and Campbell Island in the

range of this species.

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Stanley Region: S slopes of Mt. William, 1 85 m, Longton

& Smith. Longton 992. 193 (AAS, F); Goat Ridge, ca. 1 80

m (3195); summit of Mt. Kent, 455 m (2740. 2742).

Lepicolea rigida (De Not.) Scott. Figure 24.

Sendtnera rigida De Not., Mem. Reale Accad. Sci.

Torino II, 16: 229. pi. XV, 1-9. 1855. Herbertia

rigida (De Not.) Trev., Mem. Reale 1st. Lomb.

Sci. Lett. Ill, 4: 397. 1877. Leperoma rigida (De

Not.) Mass., Nuovo Giom. Bot. Ital. 17: 252.

1885. Lepicolea rigida (De Not.) Scott, Nova

Hedwigia 2: 148. 1960. Original material: Chile,

"Prov. Valparaiso, Valparaiso," Puccio (NY!).

Lepicolea seriata Herz., Hedwigia 66: 9 1 . / 8. 1 926,

syn. fide Scott ( 1 960). Lepicolea ochroleuca var.

seriata (Herz.) Herz. in Skottsberg, Nat. Hist. Juan

Fernandez Is., Easter Is. 2: 728. 1942. Original

material: Chile, Prov. Aisen, Pta. Leopardos,

Reichert & Hicken s.n. (JE), cited in Scott (1960)

(non vidi).

ECOLOGY— See comments for the ecology of L.

ochroleuca, which also apply here. Hooker visited

only East Falkland and it is likely that his collec-

tion was made in a niche similar to those of L.

ochroleuca.

PHYTOGEOGRAPHY— Valdivian + Magellanian

+ Falklands— South Georgia; Falkland Is.; Tierra

del Fuego; Patagonian Channels; Valdivian region

north to 46°32'S, 73°52'W (Punta Leopardos) (fig.

24).

FALKLAND SPECIMEN SEEN— EAST FALKLANDS.

1 843, Hookers. n. as Jungermannia hirsuta "with a Frul-

lania allied to Jung, lobulata Hook." (FH).

Family HERBERTACEAE

Herbertaceae K. Mull. (Freib.) ex Fulf. & Hatch.,

Bryologist 61: 284. 23 Feb. 1959. K. Mull.

(Freib.) in Rabenh., Krypt.-Fl. Deutschl. ed.

3, 6(1): 557. 1954, nom. nud.

Two genera belong here. Triandrophyllum has

ca. four species; it is south temperate but extends

north to Guatemala in the Neotropics and to Java

in the Paleotropics. Herbertus has ca. 75 species,

and occurs rather commonly in wet mossy forests

of temperate, but more predominantly, tropical

regions.

Triandrophyllum

Triandrophyllum Fulf. & Hatch., Bryologist 64:

349. 1961 [1962]. Fulf. & Hatch., Bryologist

61: 277. 1958 [1959], nom. inval. sin. gen.

typ* Grolle, Bryologist 64: 25. 1961, nom.

inval.

Triandrophyllum subtrifidum (Hook. f. & Tayl.)

Fulf. & Hatch.

PHYTOGEOGRAPHY — Amphipacific Temper-

ate—Macquarie Is., New Zealand, Tasmania, and

the Philippines (Kitagawa, 1981). In the American

zone in Tristan da Cunha, South Sandwich Is.,

South Georgia, Falkland Is., southern South

America, and north in the Andes to Guatemala.

• See notes in Engel ( 1978).

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

65

Variety trifidum (Gott.) Solan is Andean Ameri-

can in distribution, and var.fuscum (Steph.) Solari

occurs only in southern South America.

REMARKS— Triandrophyllum subtrifidum and T.

trifidum had been regarded as closely related species

until Solari (1973) rightfully treated them as two

varieties of a single entity, T. subtrifidum. Solari

also included a third variety, var.fuscum, of south-

ern South America. The typical variety differs from

var. trifidum by underleaf margins and ventral

margins of the leaf entire or occasionally with a

few small teeth, leaf cell walls thin to very slightly

thickened, and leaves generally bifid. Variety tri-

fidum, on the other hand, has underleaf margins

and ventral margins of the leaf with several sharp

teeth, especially toward the base; leaf cell walls

uniformly thickened; and leaves generally trifid.

Only the typical variety occurs in the Falklands.

Triandrophyllum durum, which was reported for

the Falklands by Grolle (1962), has had a check-

ered taxonomic position. Fulford and Hatcher

(1958) transferred the species to Triandrophyllum,

but in Fulford and Hatcher (1961) they placed it

as a synonym of their new combination, T. trifi-

dum (Gott.) Fulf. & Hatch., which is an older name.

Fulford (1963b) included T. durum as a synonym

of T. trifidum (even though seeing the correct type),

and, based upon these sources, I did the same in

Engel (1978). However, Solari (1973) moved the

plant to the synonymy of T. subtrifidum var. sub-

trifidum, which appears to be correct. The species

was similarly placed in Hassel de Menendez and

Solari (1975). Fulford and Hatcher (1958, p. 283,

sub plate legend) erroneously stated that the type

of Herberta dura is a Dusen collection from Fue-

gia, and included figures based upon that plant.

These figures show the marginal teeth typical of

T. trifidum, and it naturally follows that they would

place the plant there. This erroneous designation

of type specimen is probably the source of error

in taxonomic placement.

Triandroph) Hum subtrifidum (Hook. f. & Tayl.)

Fulf. & Hatch, var. subtrifidum

Jungermannia subtrifida Hook. f. & Tayl. London J.

Bot. 3: 579. 1844. Isotachis subtrifida (Hook. f.

& Tayl.) Mitt, in Hook, f., Bot. Antarc. Voyage

2(2): 149. 1854. Mastigophora subtrifida (Hook,

f. & Tayl.) Hodgs., Rev. Bryol. Lichenol. 18: 25.

1949. Triandrophyllum subtrifidum (Hook. f. &

Tayl.) Fulf. & Hatch., Bryologist 64: 350. 1961

[1962]. Original material: Tasmania, Hooker (s!).

Herberta dura Steph., Hedwigia 34: 44. 1 895, syn.fide

Solari (1973). Schisma dura (Steph.) Steph., Spec.

Hep. 4: 2 1 . 1 909. Triandrophyllum durum (Steph.)

Fujf. & Hatch., Bryologist 61: 279. 1958 [1959].

Original material: "Fretum magellanicum. leg.

Hooker fil. Herb Kew, sub nomine Jung, tena-

cifolia" (G), cited in Hassel de Menendez & Solari

(1975) and Solari (1973); (K), cited in Fulford

(\963b) (non vidi).

Leioscyphus schismoides Steph., Kongl. Svenska Ve-

tenskapsakad. Handl. 46(9): 38. f. 13f-g. 1911,

syn. fide Grolle (1964c). Leptoscyphus schis-

moides (Steph.) Kiihnem., Revista Centre Estud.

Doct. Cien. Nat. 1: 177. 1937. Mylia schismoides

(Steph.) Kiihnem., Lilloa 19: 341. 1949. Original

material: Falkland Is., Mt. Adam, 700 m, 13 De-

cember 1907, Halle & Skottsberg 347 (s!).

Isotachis lanciloba Steph., Kongl. Svenska Veten-

skapsakad. Handl. 46(9): 70. / 26f-g. 1911, syn.

fide Fulford & Hatcher (1958). Original material:

Falkland Is., Mt. Usborne, Skottsberg s.n. (G),

cited in Fulford (1963b) (non vidi).

For a complete synonymy see Solari (1973) and

Hassel de Menendez and Solari (1975); only syn-

onyms of Falkland species are included here.

Triandrophyllum durum is also included; see Re-

marks under genus.

ECOLOGY— In the Falklands, of sporadic occur-

rence on soil or rock under rock ledges of sheltered

high altitude cliffs. Rarely in stands of Hebe in

dwarf shrub heaths.

LITERATURE RECORDS (FALKLANDS)— Anony-

mous (Fulford & Hatcher, 1958, 1961; Kuhne-

mann, 1 937, as Isotachis lanciloba & Leptoscyphus

schismoides; 1949, as /. lanciloba & Mylia schis-

moides^ Skottsberg (Fulford, 1963; Hassel de Me-

nendez & Solari, 1975, and Solari, 1973, as var.

subtrifidum; Stephani, 1922, as 7. lanciloba); Mt.

Usborne (Bonner, 1966, as I. lanciloba); Mt. Adam

(Hassel de Menendez & Solari, 1975, as var. sub-

trifidum; Skottsberg, 1913, as Leioscyphus schis-

moides; Solari, 1973, as var. subtrifidum; Ste-

phani, 1922, as L. schismoides); Skottsberg & Halle,

Mt. Adam (Grolle, 1962, as T. durum).

FALKLAND SPECIMENS SEEN -EAST FALKLANDS.

Mt. Usborne Region: ridge between Mts. Usborne 1 &

2, 685 m (25/6. 2531B, 2552). WEST FALKLANDS.

Mt. Adam: 700 m, 13 Dec. 1907, Halle & Skottsberg

347, syntype of Leioscyphus schismoides (s); ridge S of

northern lake, 610m (3026, 3036A, 3040, 3042B). West-

point Island: near The Waterfall, 30-90 m (2869— c.

per.).

Family PSEUDOLEPICOLEACEAE

Pseudolepicoleaceae Fulf. & J. Tayl., Nova Hed-

wigia 1: 411. 1960.

66

FIELDIANA: BOTANY

The nine genera that comprise this family are

all small; the largest, Temnoma, has only ca. 10

species. The family is chiefly south temperate-sub-

antarctic in range.

Archeochaete

a Sphagnum bog in Tierra del Fuego, and I have

found it in the same habitat in the Brunswick Pen-

insula. This Magellanian-Falkland species is oth-

erwise known only from several stations in Tierra

del Fuego (Hassel de Menendez & Solari, 1975),

who state the species occurs at wet edges of peat

bogs. The ecology of the Falkland plants thus de-

viates considerably from that of South American

collections.

Archeochaete Schust., J. Hattori Bot. Lab. 26: 262.

1963.

Archeochaete kuehnemannii Schust.

FALKLAND SPECIMENS SEEN- EAST FALKLANDS.

Mt. Usborne Region: summit of Mt. Usborne 1, ca. 700

m (2496D). WEST FALKLANDS. Mt. Adam: ridge S

of northern lake, 610 m (3046).

Archeochaete kuehnemannii Schust., J. Hattori Bot.

Lab. 26: 262. 1963. Pseudolepicolea kuehneman-

nii (Schust.) Hassel, Comun. Mus. Argent. Cien.

Nat. "Bernardino Rivadavia," Bot. 2: 48. 1974,

nom. inval. P. kuehnemannii (Schust.) Hassel in

Hassel de Menendez & Solari, Fl. Cryptog. Tierra

del Fuego 15: 125. 1975 [1976]. Holotype: Ar-

gentina, Terr. Tierra del Fuego, ca. 16-17 km W

of Ushuaia, on road to Lapataia, C. Bandera, Feb-

ruary 1 96 1 , Schuster & Gamundi de Amos 58852

(hb. Schuster, non vidi).

REMARKS— Schuster (1965b), in his description

of Archeochaete kuehnemannii, states the leaf lobe

apices are uniseriate for a length of 4-6(7) cells.

The Falkland plants, however, have lobe apices

uniseriate for a length of (1)2-4 cells. Schuster

(1965b, p. 816), in distinguishing Archeochaete,

states:

The more attenuate leaf apices, with the

uniserate tips longer than in any species of

Lophochaete, are (also) distinctive. In Lopho-

chaete the tips are formed by, at most, 2-3

superimposed cells.

While the genera Pseudolepicolea (= Lophochaete)

and Archeochaete are distinct and well denned, the

leaf lobe apices should not be used as a character

to differentiate them. Pseudolepicolea has leaves

and underleaves subequal in size and form and

bisbifid, and perianths 4-5-6 plicate distally. Ar-

cheochaete has underleaves at most 0.5 the leaf

size and leaves, when quadrifid, with sinuses sub-

equal, and perianths 3-plicate distally (see also

Schuster, 1966c).

The only other member of the genus, A. tern-

nomoides Schust., is endemic to Tristan da Cunha.

ECOLOGY-PHYTOGEOGRAPHY— Rare; found in a

feldmark pool and on a cushion plant on a shel-

tered high altitude cliff. The type was collected in

Temnoma

Temnoma Mitt, in Hooker f., Handb. New Zea-

land Fl. 753. 1867. Teinnoma Mitt., Philos.

Trans. 168(extra vol.): 32, 33. 1879, err. ty-

pogr.9 (et Mitten, 1884).

Temnoma quadripartitum (Hook.) Mitt. Figure 25.

Jungermannia quadripartita Hook., Musci Exot. 2: pi.

117, f. 1-3. 1820. Temnoma Quadripartitum

(Hook.) Mitt., J. Linn. Soc. Bot. 15: 68. 1876.

Blepharostoma quadripartitum (Hook.) Trev.,

Mem. Reale 1st. Lomb. Sci. Lett. Ill, 4: 4 1 7. 1 877.

Original material: Argentina, Terr. Tierra del

Fuego, I. de los Estados, Menzies (K, v), cited in

Schuster (1966d) (non vidi).

Jungermannia podophylla Angstr., Ofvers. Forh.

Kongl. Svenska Vetenskapsakad. 29(4): 11. 1872,

syn. fide Pearson (1887); non Jungermannia po-

dophylla Thunb., Prodr. Fl. Cap. 2: 174. 1823 (=

Symphyogyna). Original material: Chile, Prov.

Magallanes, Pto. del Hambre, Andersson s.n. (s),

cited in Schuster ( 1 966d) (non vidi).

Lepidozia randii S. Arnell, Svensk Bot. Tidskr. 47(3):

417./ 6. 1953, syn. fide Schuster ( 1 966d). Orig-

inal material: Marion Is., Rand 3276 (s), cited in

Schuster (1966d) (non vidi).

Temnoma subintegrum Steph. ex Fulf, Mem. New

York Bot. Gard. 1 1: 57. 1963, syn. fide Schuster

( 1 966d). Blepharostoma quadripartitum var. sub-

integrum Steph., Icon. Hep. Blepharostoma no.

lOb, nom. nud. Original material: Chile, Prov.

Magallanes, Pto. Bueno, Dusen 46 (o, H), cited in

Fulford (1963b) and Schuster (1966d), respec-

tively (non vidi).

REMARKS— The Falkland material is referable

to var. quadripartitum and not var. pseudopungens

9 See Engel ( 1 978, p. 69) for notes on the typographical

error "Teinnoma."

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

67

25

FIG. 25. Distribution of Temnoma quadripartita (Hook.) Mitt.

Schust. or var. randii (S. Arnell) Schust. (See

Schuster, 1 966d, for a key to varieties.)

ECOLOGY— This taxon is rather rare in the Falk-

lands, where only a few gatherings were made in

Cortaderia (wet depressions) and dwarf shrub

heaths (under rock overhangs of outcrops).

PHYTOGEOGRAPH Y — Amphipacific Temper-

ate—northward extensions in the American sector

(West Patagonia north to 39°16'S; Andean Pata-

gonia at P. N. Nahuel Huapi and in the New Zea-

land sector into the mountains of North Is. Also

in the subantarctic, on Kerguelen Is., Crozet Is.,

Marion Is., and Prince Edward Is. The report from

Tristan da Cunha in Arnell (1958) is based on a

specimen of Archeochaete temnomoides (Schuster,

1966c). The report from Inaccessible Is. in Arnell

(1958) is questionable (fig. 25).

FALKLAND SPECIMENS SEEN -EAST FALKLANDS.

Mt. Usborne Region: The Gap, 275-290 m (2470-c.,

S + young 9, 2472). WEST FALKLANDS. Fox Bay

Region: summit of East Head, 180 m (3428— c. per.);

summit of Fox Bay Mt., 307 m (3446— c. $).

Family TRICHOCOLEACEAE

Trichocoleaceae Nakai in Ogura, Ordines, fami-

liae . . . Prof. Nakai-Takenosin ut novis edita.

200. 1943.

68

FIELDIANA: BOTANY

Trichocoleaceae have 3 genera. Trichocolea, with

about 30 species, occurs primarily in Neotropical

and south temperate mossy rain forests; one

species, T. elegans, occurs in southern South

America but is absent from the Falkland's. Leio-

mitra has ca. 4 species, all Neotropical. Eotricho-

colea has 1 species, found in New Zealand and

Tasmania.

Family LEPIDOZIACEAE

Lepidoziaceae Limpr. in Cohn, Krypt.-Fl. Schles.

1: 310. 1875 ("Lepidozieae").

The Lepidoziaceae are a large, exceedingly poly-

typic family of seven subfamilies and 28 genera.

The family is represented well in south temperate

and Neotropical regions, less so in the Paleotrop-

ics, and poorly in the Northern Hemisphere.

Hyalolepidozia

Hyalolepidozia S. Arnell ex Grolle, Rev. Bryol.

Lichenol. 32: 1 79. 1 964 [ 1 963]. S. Arnell, Hot.

Not. 115: 203. 1962, nom. inval. syn. descr.

lat.

Hyalolepidozia bicuspidata (Mass.) S. Arnell ex

Grolle. Figure 26.

Lepidozia bicuspidata Mass., Nuovo Giorn. Bot. Ital.

17: 239. pi 22, f. 25. 1885. Hyalolepidozia bi-

cuspidata (Mass.) S. Arnell, Bot. Not. 115: 213.

1962, nom. illeg. Paracromastigum bicuspidatum

(Mass.) Schust., J. Hattori Bot. Lab. 26: 276. 1963.

Hyalolepidozia bicuspidata (Mass.) S. Arnell ex

Grolle, Rev. Bryol. Lichenol. 32: 1 79. 1963. Orig-

inal material: Argentina, Terr. Tierra del Fuego,

Is. de los Estados, Pto. Cook, Spegazzini 45b (G),

cited in Fulford (1968) (non vidi).

REMARKS— After a thorough survey of branch

types in the Falkland material of H. bicuspidata,

I found both lateral- and ventral-intercalary

branches in about equal frequency, and only a sin-

gle occurrence of a Fr«//ama-type branch. I was

unable to find Microlepidozia-lype branching.

Terminal branching is only rarely developed in

the genus; in H. bicuspidata, Grolle (1963b) found

two Frullania-lype branches (Chilean material) and

one Microlepidozia-type branch (South African

material). The branching pattern in Hyalolepido-

zia readily distinguishes the genus from the closely

related genera Bonneria and Paracromastigum. In

Bonneria the branching is plastic with Acromas-

tigum-, Frullania-, Microlepidozia- and ventral-

intercalary branches (with lateral-intercalary

branches absent). Paracromastigum has Acromas-

tigum- and Frullania-lype branches and rarely

ventral-intercalary branches (with Microlepido-

zia- and lateral-intercalary branches absent). (See

Schuster, 1965a for a discussion of the evolution

and relationships within this complex of genera.)

Fulford (1968, p. 277), in her generic key, uti-

lizes as her primary key character for Hyalolepi-

dozia, "underleaves and leaves similar in size and

form." This statement presumably is based upon

examination of the type, although the ventral view

of the stem (her plate 101, fig. Ib) is from nontype

material. Massalongo (1885), in his original de-

scription of "Lepidozia" bicuspidata, gives "Am-

phigastria fol. 2-4-plo minora . . . ," and his fig-

ures clearly show the underleaves as considerably

smaller than the leaves. The underleaves of the

Falkland material of H. bicuspidata are ca. 0.33-

0.50 the leaf size and are never equal to the size

of the leaves.

ECOLOGY— Collected in dwarf shrub heaths,

where it was found on soil, particularly on soil

over rock.

PHYTOGEOGRAPHY — Amphiatlantic Temper-

ate—South Africa, Table Mt.; southern Patagonia,

Brunswick Peninsula and Rio Rubens; Patagonian

Channels; in Valdivian region north to 41°46'S in

West Patagonia; Juan Fernandez Is.; and "Frai

Jorge" (no. Chile) (see fig. 26).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Port William Region: summit ridge of Mt. Low, ca. 245

m (3233). Stanley Region: Sapper Hill, 135 m (3402).

Mt. Usborne Region: SE slope of Mt. Usborne 2, ca. 455

m (2612E). WEST FALKLANDS. Westpoint Island:

steep slope and cliffs facing The Woolly Gut (2974. 2924,

2931, 2932-c. per.).

Hygrolembidium

Hygrolembidium Schust., J. Hattori Bot. Lab. 26:

277. 1963.

Hygrolembidium isophyllum Schust. Figure 27.

Hygrolembidium isophyllum Schust., Nova Hedwigia

15: 467. pi. 56. 1968. Holotype: Argentina, Terr.

Tierra del Fuego, C. Garibaldi, near L. Escon-

dido, Schuster 583 19e (hb. Schuster!).

REMARKS— See Engel (1974) and Schuster and

Engel(1987).

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

69

26

70

FIELDIANA: BOTANY

ECOLOGY— In the Falklands this species seems

to be restricted to sheltered high altitude cliffs,

where it grows on soil under rock overhangs or on

Azorella cushions. In the Brunswick Peninsula I

found the species only in the relatively dry eastern

end (8 km west of Punta Arenas, 305-610 m),

where it grew on soil among cushion plants. Ray-

mond E. Hatcher made several collections of the

taxon between Punta Arenas and Puerto Natales

(near Rio Rubens), ca. 100 km north of the Strait

of Magellan, a locality which is on the eastern side

of the Andes and thus relatively dry. On the main-

land this taxon seems to be restricted to deciduous

Nothofagus forests of Magellanian South America.

PHYTOGEOGRAPHY— Antarctic Peninsula— Cape

Tuxen, Powell and Coronation Is. (Ochyra and

Vana, 1989); South Orkney Is.; Signy Is. (Smith,

1972, p. 24); South Georgia; Falkland Is.; moun-

tainous region of Isla Grande de Tierra del Fuego;

southern Patagonian Channels in Brunswick Pen-

insula and Rio Rubens area. This species is treated

as having an Antarctic distribution. Northward

extension from the Antarctic zone is at high or

relatively high elevations, with the curious excep-

tion of the Rio Rubens station (see Engel, 1973b)

(fig. 27).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Mt. Usborne Region: ridge between Mts. Usborne 1 &

2, 685 m (2522, 2556). WEST FALKLANDS. Port

Howard: pass SW of Mt. Maria summit, ca. 6 10 m (3102).

Mt. Adam: E side of summit ridge, 670-700 m (3008);

summit of southernmost peak, 685 m (2991).

Kurzia

Kurzia v. Mart., Flora 53: 417. 1870.

Microlepidozia (Spruce) J0rg., Bergens Mus. Skr. 16:

303. 1934. Lepidozia subg. Microlepidozia Spruce,

J. Bot. 14: 165. 1876.

Micrisophylla Fulf., Brittonia 14: 124. 1962, syn.fide

Schuster, 1969b, p. 41.

REMARKS— See Engel (1978) for discussion of placement of the genus Micrisophylla. See also the

discussion in Schuster (1980c). Through an oversight, I (Engel, 1978) neglected to make the formal

transfer of Micrisophylla to subgeneric rank in Kurzia; this was done in Schuster ( 1 980c, p. 347).

Key to Falkland Islands Species of Kurzia

1 . Leaves and underleaves stiff, erect, strongly spreading, bristle-like; leaf lobes ending in a uniseriate

row of 2-5 cells; plants, at least in a considerable apical portion, green; Acromastigum-type branches

frequently produced K. mollis

1. Leaves and underleaves suberect, leaf lobes usually curved towards the stem, leaves lax, not bristle-

like; leaf lobes ending in a uniseriate row of 1-3 cells; plants deep brown throughout, or occasionally

golden brown at apices; Acromastigum-lype branches not present 2

2. Leaf lobes (4)5-1 2 cells wide at base; Microlepidozia-lype branching rare and sporadic, branching

irregularly pinnate, lacking regularly pinnate alternation of Microlepidozia and Frullania-lype

branches; plants ± large, robust K. setiformis

2. Leaf lobes (2)3^4(7) cells wide at base; Microlepidozia-lype branching common, branching fre-

quently with ± regularly pinnate alternation of Microlepidozia and Frullania types; plants small,

220-400 n wide, delicate K. saddlensis

Opposite:

FIG. 26. Distribution of Hyalolepidozia bicuspidata (Mass.) S. Arnell.

FIG. 27. Distribution of Hygrolembidium isophyllum Schust.

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

71

kur/ia mollis (Steph.) Engel & Schust.

Lepidozia mollis Steph., Spec. Hep. 3: 601. 1909. Mi-

crisophylla mollis (Steph.) Fulf., Brittonia 14: 131.

1962. Kurzia mollis (Steph.) Engel & Schust. in

Engel, Bryologist 79: 514. 1976. Original mate-

rial: Chile, Prov. Magallanes, Pto. Bueno, Dusen

s.n. (G), cited in Fulford (1962) (non vidi).

ECOLOGY— All three Kurzia taxa are rare in the

Falklands: K. mollis was gathered in a small

depression at 275-290 m in a Cortaderia heath.

PHYTOGEOGRAPHY— Magellanian-Falkland —

Falkland Is.; Tierra del Fuego; Patagonian Chan-

nels north to 50°59'S.

FALKLAND SPECIMEN SEEN— EAST FALKLANDS. Mt.

Usborne Region: The Gap, 275-290 m (2465— c. per. +

Kurzia saddlensis (Besch. & Mass.) Grolle

Lepidozia Senlensis Besch. & Mass., Bull. Mens. Soc.

Linn. Paris 1(80): 637. 1886. Micrisophylla sad-

dlensis (Besch. & Mass.) Fulf., Brittonia 14: 128.

1962. Kurzia saddlensis (Besch. & Mass.) Grolle,

Rev. Bryol. Lichenol. 32: 174. 1963.'° Lectotype

(cf. Fulford, 1966): Chile, Prov. Magallanes, I.

Saddle, Hariot (PC, non vidi).

Lepidozia diver sifolia Steph., Kongl. Svenska Veten-

skapsakad. Handl. 46(9): 62. / 23c-f. 191 1, syn.

fide Fulford (1962). Original material: Chile, Prov.

Chiloe, I. Guaitecas, Melinca, Skottsberg s.n. (G),

cited in Fulford (1962) (non vidi).

REMARKS— The 1902 Skottsberg collections of

Kurzia saddlensis (Lepidozia saddlensis) from the

Falkland Islands in Stephani (1905a) are misde-

terminations of Lepidozia chordulifera.

ECOLOGY— Very rare in the Falklands, where it

was mixed with other hepatics (Clasmatocoleaful-

vella, Gackstroernia magellanica and Riccardia

papillosa) in wet depressions of a Cortaderia heath

(455 m).

PHYTOGEOGRAPHY— Valdivian + Magellanian-

Falkland— Falkland Is.; Tierra del Fuego; Valdi-

vian West Patagonia at 43°57'S. Also Tristan da

Cunha, Nightingale, and Inaccessible Is. I regard

the report from Tasmania in Rodway (1916) as

doubtful.

LITERATURE RECORDS (FALKLANDS)— Anony-

mous (Kuhnemann, 1937, 1949); Skottsberg, Port

Stanley & Port Louis (Skottsberg, 1913).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Mt. Usborne Region: SE slope of Mt. Usborne 2, ca. 455

m (2594B-C. 6, 2597B).

Kurzia setiformis (De Not.) Engel & Schust.

Lepidozia setiformis De Not., Mem. Reale Accad. Sci.

Torino II, 16: 255. / XIII, 1-6. 1855. Mastigo-

phora setiformis (De Not.) Trev., Mem. Reale 1st.

Lomb. Sci. Lett. Ill, 4: 416. 1877. Micrisophylla

setiformis (De Not.) Fulf., Brittonia 14: 127. 1962.

Kurzia setiformis (De Not.) Engel & Schust. in

Engel, Bryologist 79: 514. 1976. Neotype" (fide

Fulford, 1 966): Argentina, Terr. Tierra del Fuego,

Is. de los Estados, Speggazzini s.n. (G 344! ex hb.

Massalongo M 52). Original material: Chile, "Prov.

Valparaiso, Valparaiso," Puccio (non vidi).

Lepidozia obscura Angstr. in Steph., Spec. Hep. 3:

602. 1909, syn. fide Fulford (1962). Original ma-

terial: Chile, Prov. Magallanes, Strait of Magel-

lan, Pto. del Hambre, Andersson s.n. (FH!, G!).

Lepidozia fusca Steph., Kongl. Svenska Vetenskap-

sakad. Handl. 46(9): 64. / 24h-i. 191 1, syn. fide

Fulford (1962). Original material: Chile, Prov.

Magallanes, S. Skyring, B. Pinto, Halle & Skotts-

berg 170 (G, s), cited in Fulford ( 1 962) (non vidi).

Lepidozia cunninghamii Steph., Spec. Hep. 6: 322.

1922, syn. fide Fulford (1962). Original material:

Chile, Prov. Magallanes, Pto. Grappler, Cun-

ningham inter no. 179 (G), cited in Fulford ( 1 962)

(non vidi).

ECOLOGY— Very rare in the Falklands, where it

was gathered in a wet depression of a Cortaderia

heath (455 m) mixed with other hepatics, nearly

all of which are rare or relatively so, e.g., Telaranea

oligophylla, Clasmatocolea obvoluta var. cook-

iana, Hyalolepidozia bicuspidata, and Leptoscy-

phus aequatus.

PHYTOGEOGRAPHY— Magellanian-Falkland —

Falkland Is.; Tierra del Fuego; Patagonian Chan-

nels north to 48°55'S. Reportedly disjunct in Tas-

mania (fide Fulford, 1962, 1966).

FALKLAND SPECIMEN SEEN- EAST FALKLANDS. Mt

Usborne Region: SE slope of Mt. Usborne 2, ca. 455 m

(2612B).

Lepidozia

Lepidozia (Dum.) Dum., Recueil Observ. Jun-

germ. 19. 1835, nom. cons. Pleuroschismasecl.

Lepidozia Dum., Syll. Jungerm. Eur. Indig.

69. 1831.

10 Grolle incorrectly cited the basionym reference as

Besch. & Mass. Mission Sci. du Cap Horn 5: 233. 1889.

" See Engel (1978) for notes on the neotype locality.

72

FIELDIANA: BOTANY

Key to Falkland Islands Species of Lepidozia

1 . At least some leaves and/or underleaves on well-developed stems with marginal teeth-lacininae . .

L. chordulifera

1 . Margins of leaves and underleaves entire, very rarely with an isolated tooth 2

2. Leaf segments in conspicuous pairs, the dorsal and ventral segments often reduced to a tooth of

a few cells; leaves often distant, often scalelike; (only above 245 m in Falklands, common on

sheltered high altitude cliffs) L. fuegiensis

2. Leaf segments not in conspicuous pairs, dorsal and ventral segments larger, never reduced to a

few cells; leaves imbricate, leafy, never scalelike; (rather common below 245 m in Falklands,

absent on sheltered high altitude cliffs) L. laevifolia

Lepidozia chordulifera Tayl. Figure 28.

Lepidozia chordulifera Tayl., London J. Bot. 5: 371.

1846. Jungermannia chordulifera (Tayl.) Hook,

f. & Tayl. in Hook, f., Bot. An tare. Voyage 1(2):

442. 1847. Mastigophora chordulifera (Tayl.)

Trev., Mem. Reale 1st. Lomb. Sci. Lett. Ill, 4:

397. 1877. Original material: Chile, Prov. Aisen

& Chiloe, Arch, de los Chonos, 1 834, Darwin 461

(FH— cited in Fulford, 1966, NY!).

Lepidozia hastata Steph., Spec. Hep. 3: 605. 1909,

syn.fide Fulford (1966). Original material: Chile,

Prov. Aisen, R. Aisen, Dusen 83a (G), cited in

Fulford (1 966) (non vidi).

Lepidozia cuspidata Steph., Kongl. Svenska Veten-

skapsakad. Handl. 46(9): 61. f. 23a-b. 191 1, syn.

nov. Original material: Western Patagonia,

Skottsberg s.n. (G), cited in Fulford (1966) (non

vidi).

Lepidozia ejffusa Steph., Kongl. Svenska Vetenskap-

sakad. Handl. 46(9): 62. / 23g-h. 1911, syn.fide

Fulford (1966). Original material: Chile, Prov.

Magallanes, Pto. Ramirez and I. Atalaya, Halle

& Skottsberg (G), cited in Fulford (1966) (non

vidi).

Lepidozia fernandeziensis Steph., Kongl. Svenska Ve-

tenskapsakad. Handl. 46(9): 63. / 24e. 191 1,12

syn.fide Fulford (1966). Original material: Juan

Fernandez Is., Mas a Tierra, near El Yunque,

Skottsberg s.n. (G), cited in Fulford (1966) (non

vidi); Prov. Chiloe, I. Chiloe, Fundo San Antonio,

R. Pudito, Skottsberg (non vidi); Prov. Magal-

lanes, R. Azopardo, Skottsberg (non vidi).

Lepidozia hariotii Steph., Spec. Hep. 6: 329. 1922,

syn.fide Fulford (1966). Original material: Chile,

Prov. Magallanes, I. Hermite, Hariot (G, ex hb.

Bescherelle), cited in Fulford (1966) (non vidi).

Lepidozia microscopica Steph., Spec. Hep. 6: 334. 1 922,

syn.fide Fulford (1966). Original material: Chile,

Prov. Aisen, Cta. Hale, Dusen s.n. (G), cited in

Fulford (1966) (won vidi).

Lepidozia angulata Steph. ex Fulf., Mem. New York

Bot. Gard. 1 1: 206. 1966, nom. nud., pro syn.

12 Fulford (1966) stated the description and figures of

L. fernandeziensis have been interchanged with those of

L. disticha in Stephani (1911).

REMARKS— See Engel (1978).

ECOLOGY— Very common in the Falklands,

where it is better able to tolerate quite xeric, ex-

posed conditions than any other Falkland hepatic,

often occurring in crevices and under overhangs

of rock outcrops in dwarf shrub heaths. In this

association it is also frequently encountered under

Blechnum magellanicum cover as well as on Bolax

cushions. Rare in sheltered high altitude cliffs and

moist feldmarks.

PHYTOGEOGRAPHY— Valdivian + Magellanian

+ Falklands— South Sandwich Is.; South Georgia;

Falkland Is.; Tierra del Fuego; Patagonian Chan-

nels; Valdivian region; Juan Fernandez Is. (400-

795 m on Mas a Tierra). The report from Tas-

mania in Rodway (1916) requires confirmation

(see fig. 28).

LITERATURE RECORDS (FALKLANDS)— A nony-

mous (Kuhnemann, 1937, 1 949); Skottsberg, Port

Stanley (Fulford, 1966; Skottsberg, 1913), Port

Harriet (Fulford, 1966); Skottsberg & Halle, Port

Harriet (Stephani, 1911).

FALKLAND SPECIMENS SEEN -EAST FALKLANDS.

1843, Hooker s.n. as Jungermannia laevifolia var. in-

crassata (FH). Port William Region: summit ridge of Mt.

Low, ca. 245 m (3230, 3238); N side of Gypsy Cove,

sea level (3242-3243, 3252, 3258); Engineer Point head-

land, The Narrows, ca. 15m, Imshaug 40618 (MSC);

ridge on Engineer Point Peninsula, 18-25 m (2817).

Stanley Region: Rookery Bay, ca. 15m, Smith 3026

(AAS, F); Port Stanley, 8 Apr. 1902, Skottsberg s.n. as

Lepidozia saddlensis (s); Port Stanley, 1953, Pettingill

32 (CMC, F); near Sapper Hill, 29 Oct. 1907, Skottsberg

as L. pallida (s, UPS); Sapper Hill, 135 m (2.792. 2394.

239? 'A, 2399. 2407 -c. per., 2410, 2413-c. sporo.-2414,

2419); near summit of Tumbledown Mt., 215m, Long-

ton & Smith, Longton 977 (AAS, F); Tumbledown Mt.,

155-230 m (237 '5 A. 2380, 2384, 2391); Goat Ridge, ca.

180 m (3118. 3205, 3208, 3216); summit ridge of N

peak of Two Sisters, 245-290 m (2703 A. 2727, 27525,

2735-2736); N peak of Two Sisters, 250 m, Dollman

s.n. (Greene 3560a) (AAS); summit of Mt. Kent, 455 m

(2766). Mt. Usborne Region: gap between Mt. Usborne

2 and Table Rock, ca. 440 m (2640B); summit of Mt.

Usborne 1, ca. 700 m (2495). WEST FALKLANDS.

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

73

FIG. 28. Distribution of Lepidozia chordulifera Tayl., plus South Georgia. Open circle indicates precise locality

unknown.

Port Howard: Freezer Rocks, E slope of Mt. Maria, 320

m (3141. 3147). Fox Bay Region: summit of East Head,

!80 m (3421. 3439); valley E of Sulivan House, ca. 75

m (3531). Mt. Adam: E side of summit ridge, 670-700

m (3017). Hill Cove Region: summit of East French

Peak, 305 m (2977); summit of West French Peak, 290

m (296 2-296 3); summit of Mt. Fegen, 335-360 m (3065.

3073. 3076); ridge of N slope of Mt. Fegen, ca. 275 m

(3057. 3062).

Lepidozia fuegiensis Steph.

Lepidozia fuegiensis Steph., Kongl. Svenska Veten-

skapsakad. Handl. 46(9): 63. / 24f-g. 191 1. Orig-

inal material: Chile, Prov. Magallanes, Pto. Cut-

ter, 13 April 1908, Halle & Skottsberg (o!, sub

"Patagonia austra.," s!, UPS!); Prov. Magallanes,

Cta. Gome/, 26 Feb. 1908, Halle & Skottsberg

(s!, UPS!).

Lepidozia minuta Steph., Spec. Hep. 3: 603. 1909,

non Lepidozia minuta Col., Trans. & Proc. New

Zealand Inst. 18: 245. 1886, syn. fide Fulford

(1966). Original material: Chile, Prov. Magal-

lanes, R. Azopardo, Dusen 93 (o), cited in Fulford

(1966) (non vidi).

Lepidozia magellanica Steph., Kongl. Svenska Veten-

skapsakad. Handl. 46(9): 64./ 24m-n. 191 1, syn.

fide Fulford (1966). Original material: Chile, Prov.

Magallanes, I. Felix, Skottsberg 594 (s), cited in

Fulford (1 966) (non vidi).

Lepidozia parva Steph., Kongl. Svenska Vetenskap-

sakad. Handl. 46(9): 65. / 24o-p. 1911, syn. fide

Fulford (1966). Original material: Chile, Prov.

Aisen, Cta. Hale, and Cta. Connor, Hale &

Skottsberg (o), cited in Fulford (1966) (non vidi).

REMARKS— See Engel (1978) for a comparison

of Brunswick Peninsula and Falkland Islands pop-

ulations of this species, and for enumeration of

features which will identify the taxon.

74

FIELDIANA: BOTANY

Stephani (1911) based Lepidozia fuegiensis on

specimens from four localities— Pto. Cutter, Cta.

Gomez, R. Fontaine, and R. Azopardo. Collec-

tions at S and UPS from the first two localities

rather closely match the protologue of L. fuegien-

sis. The R. Fontaine plants, however, are L. lae-

vifolia (1 March 1909, Halle & Scottsberg, UPS!),

and I have not been able to locate the R. Azopardo

collection. The specimens from G are labeled either

"Patagonia austr." or "Fuegia." Since Pto. Cutter

is the sole locality in Patagonia (the remaining

three being in Tierra del Fuego), I am assuming

those labeled "Patagonia austr." are indeed from

Pto. Cutter. The single G collection from Fuegia

is L. laevifolia and is probably the R. Fontaine

plant.

ECOLOGY— Found above 245 m in the Falk-

lands, where it is of frequent occurrence in crevices

of sheltered high altitude cliffs. Also known from

feldmark rocks, particularly those in shallow pools,

but only rarely in Cortaderia and dwarf shrub

heaths. It is of interest to note that L. chordulifera

and L. laevifolia are very common below 245 m

and only rarely (a single collection of L. chordu-

lifera) encountered in sheltered high altitude cliffs.

PHYTOGEOGRAPHY— Valdivian + Magellanian

+ Falklands— South Georgia; Falkland Is.; Tierra

del Fuego; southern Patagonian Channels; Bruns-

wick Peninsula; Valdivian portion of Patagonian

Channels. The "Frai Jorge" report in Herzog (1954)

requires confirmation.

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Stanley Region: summit ridge of N peak of Two Sisters,

245-290 m (2724). Mt. Usborne Region: gap between

Mt. Usbome 2 and Table Rock, ca. 440 m (2638); SE

slope of Mt. Usborne 2, ca. 455 m (2604, 2616— c. per.);

ridge between Mts. Usborne 1 & 2, 685 m (2534 '-A, 2539B);

summit of Mt. Usborne 1 , ca. 700 m (2491B-2492, 2496);

The Gap, 275-290 m (2466). WEST FALKLANDS.

Port Howard: pass SW of Mt. Maria summit, ca. 610 m

(3083A, 3096, 3107, 3109, 3126B). Mt. Adam: ridge S

of northern lake, 610 m (3033, 3035).

Lepidozia laevifolia (Hook. f. & Tayl.) G. L. & N.

Jungermannia laevifolia Hook. f. & Tayl., London J.

Bot. 3: 385. 1844 (3: 285. [sic.] in errorepro 385).

Lepidozia laevifolia (Hook. f. & Tayl.) G. L. &

N., Syn. Hep. 208. 1845. Mastigophora laevifolia

(Hook. f. & Tayl.) Trev., Mem. Reale 1st. Lomb.

Sci. Lett. Ill, 4: 416. 1877. Original material:

Campbell Is., Hooker (non vidi).

Lepidozia Jacquinotii" Steph., Spec. Hep. 3: 604. 1 909,

syn. fide. Fulford (1966), non Lepidozia

Jacquinotii" Steph., Spec. Hep. 6: 330. 1922.

Original material: Chile, Prov. Magallanes, Pto.

Gallant, Jacquinot (G) (non vidi), cited in Fulford

(1966); Patagonia (without specific location), Du-

sen (non vidi); Prov. Chiloe, I. Chiloe, sin. coll.

(non vidi).

Lepidozia viridissimaSleph., Spec. Hep. 3: 604. 1909,

syn. fide Fulford (1966). Original material: Chile,

Prov. Magallanes, Strait of Magellan, Dusen (a),

cited in Fulford (1966) (non vidi).

Lepidozia falklandica Steph., Kongl. Svenska Veten-

skapsakad. Handl. 46(9): 63. / 24c-d. 191 1, syn.

fide Fulford (1966). Original material: Chile, Prov.

Aisen, L. O'Higgins, Hailed Skottsberg; Falkland

Is., Mt. Adam, and Hearnden Water, Halle &

Skottsberg (G), cited in Fulford (1966) (non vidi).

Lepidozia halleana Steph., Kongl. Svenska Veten-

skapsakad. Handl. 46(9): 64. / 24k-l. 191 1, syn.

nov. Original material: Falkland Is., near Port

Stanley, Hearnden Water, 1 Nov. 1907, Halle (G!,

s!, UPS!).

ECOLOGY— Rather common in dwarf shrub

heaths, where it is found on soil under Empetrum,

or more rarely on stream banks. It also may be

found in rock crevices of outcrops in this associ-

ation.

PHYTOGEOGRAPHY — Amphipacific Temper-

ate—reported from South Georgia; Falkland Is.;

the Magellanian and Valdivian regions; Juan Fer-

nandez Is.; Macquarie Is.; Auckland and Campbell

Is.; New Zealand; Tasmania; Kerguelen Is.; Mar-

ion and Prince Edward Is.

LITERATURE RECORDS (FALKLANDS)— A nony-

mous (Kiihnemann, 1937, 1949 as L. halleana, L.

jacquemontii [sic], and L. falklandica); Halle (Ful-

ford, 1966, Stephani, 1922 as L. halleana, and

Taylor, 1960 as L. halleana); Hooker (Taylor &

Hooker, 1847 as Jungermannia); Skottsberg (Ful-

ford, 1966), Port Stanley (Skottsberg, 1913 as L.

jacquemontii [sic]); Port Stanley and Mt. Adam

(Skottsberg, \9l3asL.falklandica, Stephani, 1922

as L. falklandica); Skottsberg & Halle (Fulford,

1966); Port Stanley (Stephani, 1911 as L. jacque-

montii [sic]).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Port William Region: N side of Gypsy Cove, sea level

(3259); ridge on Engineer Point Peninsula, 20-25 m

(2834). Stanley Region: Sapper Hill, 1 35 m (2398, 24 17 A,

2426); Goat Ridge, ca. 180 m (3199); Sapper Hill, 2 km

W of Port Stanley, ca. 50 m, Ochyra 2830/80 (F). Darwin

Settlement: Boca House on Brenton Lock, sea level (2696).

13 Stephani, in describing his new species, used the

spelling "Jacquemontii," which was the spelling used by

subsequent authors. Since the species was named after

Honore Jacquinot, I have altered the spelling of the ep-

ithet accordingly (see Engel, 1978, p. 99).

14 1 believe there is a distinct possibility that Lepidozia

Jacquemontii of Stephani, 1922, leg. Skottsberg at "Fre-

tum magellanicum," is based upon the same type as L.

jacquemontii of Stephani, 1909. See Engel (1978, p. 99)

for discussion.

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

75

Port Louis: 2 Aug. 1902, Skottsberg s.n. as L. saddlensis

(s). WEST FALKLANDS. Fox Bay Region: near peat

cuttings in Ram Paddocks, 45-60 m (3509); valley E of

Sulivan House, ca. 75 m (3536).

Notes on Lepidozia Species

Until the type of Lepidozia pallida Steph. is lo-

cated, the identity of this species must remain open

to question. As I have pointed out (Engel, 1978,

p. 102), perhaps it is of some significance that all

the reports (from five localities) of Lepidozia pal-

lida in Stephani (1911) were found to be of Lep-

idozia chordulifera. This includes the Falkland

specimen from Port Stanley, Sapper Hill.

Paracromastigum

Paracromastigum Fulf. & J. Tayl., Brittonia 13:

336. 1961.

Key to Falkland Islands Species of Paracromastigum

*

1. Terminal branching common, of Frullania and Acromastigum types; cortical cells in ca. 12 rows . . .

P. subsimplex

1 . Terminal branching absent (in Falkland specimens) or, if present, then rare and only of Frullania

type; cortical cells in 7-8 rows P. tristanianum

Paracromastigum subsimplex (Steph.) Fulf. & J.

Tayl.

Lepidozia subsimplex Steph., Kongl. Svensk Veten-

skapsakad. Handl. 46(9): 66. / 24r-s. 1911. Par-

acromastigum subsimplex (Steph.) Fulf. & J. Tayl.,

Brittonia 13: 336. 1961. Original material: (?)

Chile, Prov. Magallanes, (?) Argentina, Terr. Tierra

del Fuego, L. Fagnano, 1908, Halle s. n. (G), cited

in Fulford & Taylor (1961) (non vidi).

ECOLOGY— This rare species, represented by a

single Falkland collection, was gathered on the

bank of a wet depression in a Cortaderia heath,

580-595 m.

PHYTOGEOGRAPHY— Valdivian + Magellanian

+ Falklands— Falkland Is.; Isla Grande de Tierra

del Fuego; Valdivian region.

FALKLAND SPECIMEN SEEN -WEST FALKLANDS.

Mt. Adam: basin E of summit, 580-595 m (3002).

Paracromastigum tristanianum (Schust.) Engel &

Schust.

Pseudocephalozia tristaniana Schust., Nova Hedwigia

10: 23. 1965. Paracromastigum tristanianum

(Schust.) Engel & Schust. in Schuster & Engel, J.

Hattori Bot. Lab. 38: 700. 1974. Holotype: Tris-

tan da Cunha, Red Hill, Christopher sen 1229C

(G, non vidi).

REMARKS— The Falkland Islands plants fit the

original description of the species, except for two

features. Median leaf cells measure 21-36 ^m wide

x 39-53 nm long in Falkland material and "ca.

20-25 x 25-38 Mm" in type material. The type

has predominantly intercalary branching, with ter-

minal, Frullania-type branches rare and Acro-

mastigum-type branches lacking. Falkland mate-

rial has abundant intercalary branching from all

three merophytes, but no terminal branches were

observed. Stolons are abundant, freely branched,

and prostrate, while the leafy shoots are erect and

issue from the stolon system. Androecia are sim-

ilar to the type in being on leading shoots and

lacking bracteolar antheridia. Leafy shoots on

Falkland plants have branches originating only

from basal sectors and never from the zone of

mature leaves. If that condition occurs in the type,

it is then of diagnostic value, and will differentiate

the species from P. subsimplex. In that species,

branching freely develops from mature-leaved

sectors of shoots.

ECOLOGY- PHYTOGEOGRAPHY— Apparently quite

rare in the Falklands, where it was collected but

once— on a stream bank in a Cortaderia heath.

This amphiatlantic temperate species is restricted

to the Falkland Is. and Tristan da Cunha.

FALKLAND SPECIMEN SEEN -EAST FALKLANDS. Mt.

Usborne Region: valley SW of Mt. Usborne, ca. 60 m

(2670-c. S).

76

FIELDIANA: BOTANY

FIG. 29. Distribution of Pseudocephalozia quadriloba (Steph.) Schust.

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

77

Pseudocephalozia

Pseudocephalozia Schust., Nova Hedwigia 10: 2 1 .

1965.

Pseudocephalozia quadriloba (Steph.) Schust. Fig-

ure 29.

Isotachis quadriloba Steph., Bih. Kongl. Svenska Ve-

tenskapsakad. Handl. 26(111, 6): 54. 1900. Hy-

grolembidium quadrilobum (Steph.) Schust., Nova

Hedwigia 10: 23. 1965. Lembidium quadrilobum

(Steph.) Fulf., Mem. New York Hot. Gard. 11:

247. 1966. Pseudocephalozia quadriloba (Steph.)

Schust., J. Hattori Hot. Lab. 36: 37 1 . 1 973 [ 1972].

Original material: Chile, Prov. Chiloe, I. Guai-

tecas, Dusen 182 (o 1 1 102! as Fuegia, without

specific locality).

Cephalozia quadriloba Steph., Kongl. Svenska Veten-

skapsakad. Handl. 46(9): 58. / 22a-b. 1911 =

Lophozia crassicaulis Steph., Spec. Hep. 6: 111.

1917, syn. fide Schuster & Engel ( 1 974). Original

material: Chile, Prov. Magallanes, W. end of L.

Fagnano, Skottsberg 555 (G! as Fuegia, without

specific locality).

Isotachis granditexta Steph., Kongl. Svenska Veten-

skapsakad. Handl. 46(9): 68. / 26c, e. 1911, syn.

fide Fulford (1966). Original material: Chile, Prov.

Magallanes, L. Fagnano, R. Azopardo region, 10-

1 1 March 1908, Halle 127 (UPS!).

REMARKS— See Schuster and Engel (1974).

ECOLOGY— See Schuster and Engel (1974, pp.

693-694 and fig. 17).

PHYTOGEOGRAPHY— Andean American — Inac-

cessible I. (475 m); Falkland Is. (10-300 m); Isla

Grande de Tierra del Fuego (85-650 m); southern

Patagonian Channels (Brunswick Peninsula and

Hotel Rubens-Puerto Natales region); Valdivian

region (at 43°57'S, Islas Guaitecas); Colombia

(3200-3600 m); Venezuela (3850 m); Costa Rica

(2600 m) (see fig. 29).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Port William Region: summit ridge of Mt. Low, ca. 245

m (3224, 3226B). Stanley Region: headwaters of Mullet

Creek Stream, ca. 60 m (3171); summit ridge of N peak

of Two Sisters, 245-290 m (2702, 2776). Mt. Usborne

Region: The Gap, 275-290 m (2459, 2473). WEST

FALKLANDS. Fox Bay Region: near mouth of Cheek's

Creek, ca. 12 m (3478).

Telaranea

Telaranea Spruce ex Schiffn. in Engler & Prantl,

DieNaturl. Pflanzenfam. 1(3, 1): 103. 1893.

Telaranea Spruce, Trans. & Proc. Bot. Soc.

Edinburgh 15: 365. 1885, nom. inval. in syn.

Neolepidozia Fulf. & J. Tayl., Brittonia 11:

81. 1959.

Key to Falkland Islands Species of Telaranea

1. Leaf apices 2-3 dentate-lobate, and if lobate, then lobes less than 0.5 the leaf length; leaf disc more

than 4 rows of cells high. Leaf segments 2 or 3(4) cells long and 2 cells wide at the base; leaf cells in

8 longitudinal rows Subg. Neolepidozia . . . T. oligophylla

1. Leaf apices 4-6 lobed for 0.5 or more the leaf length; leaf disc 1-4 rows of cells high

Subg. Telaranea ... 2

2. Leaves of main axis with lamina of 1 row of cells high, the leaves 2-3 segmented; underleaves

conspicuously smaller than the leaves; plants small, filamentous T. pseudozoopsis

2. Leaves of main axis with lamina of 1 .5 to 4 rows of cells high, the leaves 4-6 segmented; underleaves

similar to leaves in size; plants large, leafy 3

3. Leaves and underleaves stiff, ascending, bristle-like, the lamina 1.5 cell rows high

T. blepharostoma

3. Leaves and underleaves lax, curved, not ascending and bristle-like, the lamina 2-4 cell rows high . . .

4

4. Leaf margins 3-4 cell rows high; shoot apices often recurved; perianth mouth crenulate

T. plumulosa

4. Leaf margins 1-2 cell rows high; shoot apices plane, not recurved; perianth mouth long-ciliate . .

T. tetradactyla

78

FIELDIANA: BOTANY

Telaranea blepharostoma (Steph.) Fulf. Figure 30.

Lepidozia Blepharostoma Steph., Bih. Kongl. Svenska

Vetenskapsakad. Handl. 26(111, 17): 22. 1901.

Telaranea blepharostoma (Steph.) Herz., Rev.

Bryol. Lichenol. 20: 189. 1960, nom. illeg. Te-

laranea blepharostoma (Steph.) Fulf., Brittonia

15: 73. 1963. Original material: Chile, Prov. Ma-

ua llanos. Pto. Angosto, "ex Bot. Mus. Upsal."

Dusen 142 (o), cited in Fulford ( 1 963a) (non vidi).

Lepidozia trichophylla Angstr. ex Fulf., Mem. New

York Bot. Card. 11: 240. 1966, nom. nud., pro

syn.

REMARKS— As all previous reports of T. ble-

pharostoma from the Falklands are based upon

misdeterminations of T. plumulosa, this is the first

authentic report of Telaranea blepharostoma in

these islands.

ECOLOG Y-PH YTOGEOGR APH Y — Va Id i via n +

Magellanian-Falkland— Falkland Is.; Tierra del

Fuego; Patagonian Channels; Valdivian region (see

fig. 30). Within the Falklands, it was found to occur

only in the Port Stanley region under Blechnum

in a dwarf shrub heath, and on moist, sandy, shel-

tered, vertical, north-facing rocks along the south

shore of the Murrell River. The cooler tempera-

tures and higher rainfall in this region may account

for its presence there.

LITERATURE RECORDS (FALKLANDS)— A nony-

mous (Kiihnemann, 1937, 1949 as Lepidozia);

Skottsberg—Port Philomel, Port Stanley (Skotts-

berg, 1913 as Lepidozia); Skottsberg & Halle—

Port Philomel, Port Stanley (Stephani, 1911 as

Lepidozia).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Stanley Region: S shore of Murrell River, opposite Islet

Point, 30 m, Longton & Smith, Longton 1027 (AAS, F);

Goat Ridge, ca. 180 m (3201).

Telaranea oligophylla (Lehm. & Lindenb.) Engel.

Figure 31.

Jungermannia oligophylla Lehm. & Lindenb. in Leh-

mann, Nov. Minus Cogn. Stir. Pug. 6: 26. 1 834.

Lepidozia oligophylla (Lehm. & Lindenb.) G. L.

& N., Syn. Hep. 201. 1845. Mastigophora oli-

gophylla (Lehm. & Lindenb.) Trev., Mem. Reale

1st. Lomb. Sci. Lett. Ill, 4: 415. 1877. Neolepi-

dozia oligophylla (Lehm. & Lindenb.) Fulf. & J.

Tayl., Brittonia 11: 84. 1959. Telaranea oligo-

phylla (Lehm & Lindenb.) Engel, Bryologist 79:

514. 1976. Original material: Argentina, Terr.

Tierra del Fuego, I. de los Estados, Menzies s.n.

(NY!).

REMARKS— See Engel (1978).

30

Fio. 30. Distribution of Telaranea blepharostoma

(Steph.) Fulf.

FIG. 3 1 . Distribution of Telaranea oligophylla (Lehm.

& Lindenb.) Engel. Open circle indicates precise locality

unknown.

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

79

ECOLOGY— Very rare in the Falklands, where it

was gathered in a wet depression of a Cortaderia

heath (455 m). It occurred mixed with other hepat-

ics, nearly all of which are rare or relatively so,

e.g., Clasmatocolea obvoluta var. cookiana, Hy-

alolepidozia bicuspidata, Leptoscyphus aequatus,

Kunia setiformis, and Riccardia spectabilis.

PH YTOGEOGRAPH Y — Magellanian-Fa 1 k 1 a 1 1 c I —

Falkland Is.; Tierra del Fuego; Patagonian Chan-

nels north to 49°09'S (see fig. 31).

FALKLAND SPECIMEN SEEN— EAST FALKLANDS. Mt.

Usborne Region: SE slope of Mt. Usborne 2, ca. 455 m

(26 12 A).

Telaranea plumulosa (Lehm. & Lindenb.) Fulf.

Figure 32.

Jungermannia plumulosa Lehm. & Lindenb. in Leh-

mann, Nov. Minus Cogn. Stir. Pug. 6: 26. 1834.

Lepidozia plumulosa (Lehm. & Lindenb.) G. L.

& N., Syn. Hep. 211. 1845. Mastigophora plu-

mulosa (Lehm. & Lindenb.) Trev., Mem. Reale

1st. Lomb. Sci. Lett. Ill, 4: 416. 1877. Telaranea

plumulosa (Lehm. & Lindenb.) Herz., Rev. Bryol.

Lichenol. 29: 189. 1960, nom. illeg. Telaranea

plumulosa (Lehm. & Lindenb.) Fulf., Brittonia

15: 77. 1963. Original material: Argentina, Terr.

Tierra del Fuego, I. de los Estados, Menzies s.n.

(G) "ex hb. Kew," cited in Fulford (1963a) (non

vidi).

Lepidozia magellanica Gott. ex Fulf, Mem. New York

Bot. Gard. 11: 243. 1966, nom. nud., pro syn.,

non Lepidozia magellanica Steph., Kongl. Sven-

ska Vetenskapsakad. Handl. 46(9): 64. 1911 (=

L. fuegiensis).

REMARKS— This taxon reaches its optimal de-

velopment in the evergreen forested region of

southern Chile. The plants of the Falkland pop-

ulation, when compared with the Chilean plants,

are smaller in size, less luxuriant and less robust,

often without recurved plant tips, and with few

exceptions have leaves which are consistently

4-segmented. Fulford ( 1 963a, 1 966) stated that the

leaves of T. plumulosa are 5- or 6-segmented. The

leaf lamina and margins of the Falkland plants are

3-4 cells high, data which agrees with the Fulford

descriptions. The Falkland Islands plants, which

show little variation, are thus somewhat inter-

mediate between T. plumulosa and T. tetradac-

tyla; however, I believe the concept of the former

should be expanded to include plants with 4-seg-

mented leaves.

ECOLOGY— Very common, characteristic of

stream banks, particularly under Gunnera, Blech-

num magellanicum, or Empetrum cover and in

crevices of outcrops in dwarf shrub heaths. Also

common on soil, frequently of stream banks in

Cortaderia heaths. It occasionally grows in rock

crevices or under overhangs of sheltered high al-

titude cliffs.

PHYTOGEOGRAPHY— Valdivian + Magellanian

+ Falkland -Falkland Is.; Tierra del Fuego; Pa-

tagonian Channels; Valdivian region (West Pata-

gonia only at 43°57'S, on an offshore island); and

Mas a Tierra, Juan Fernandez Is. (400-500 m) (fig.

32).

LITERATURE RECORDS (FALKLANDS)— A nony-

mous (Fulford, 1963a, 1966; Kuhnemann, 1937

as Lepidozia); Lyall (Fulford, 1963a, 1966);

Skottsberg, North West Bay (Fulford, 1 963a, 1 966),

Port Stanley (Fulford, 1966).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Stanley Region: near Port Stanley, 1 1 Nov. 1907, Skotts-

berg 165 as Lepidozia blepharostoma (s, UPS), headwaters

of Mullet Creek Stream, ca. 60 m (3162); between Mt.

William and Tumbledown Mt., 1 70 m, Longton & Smith,

Longton 979 (AAS, F); summit ridge of N peak of Two

Sisters, 245-290 m (2700/1, 2779, 2738); summit of Mt.

Kent, 455 m (2746). Mt. Usborne Region: SE slope of

Mt. Usborne 2, ca. 455 m (2596); The Gap, 275-290 m

(2467, 2477); below The Gap, ca. 90 m (2564, 2567,

2577); S side of The Gap, 260-275 m (2452A); valley

SW of Mt. Usborne, ca. 60 m (2655, 2664). WEST

FALKLANDS. Port Howard: SW of Mt. Maria summit,

ca. 610 m (3131). Fox Bay Region: near peat cuttings in

Ram Paddock, 45-60 m (3514— c. 9); N base of Fox Bay

Mt., 75 m (5459, 5460, 3464A). Mt. Adam: ridge S of

northern lake, 610m (5025, 303 IB). Hill Cove Region:

gap between French Peaks, 200-2 1 5 m (2978); summit

of West French Peak, 290 m (2949); Hill Cove, Vallentin

Sp. 55 (MANCH). Port Philomel: Northwest Bay, 2 Nov.

1907, Halle s. n. as Lepidozia blepharostoma (UPS). West-

point Island: near The Waterfall, 30-90 m (2574, 2575).

Weddell Island: Waterfall Valley, W of settlement, ca.

125 m (3305 A, 3312, 3321, 3327B, 3328); near head-

waters of House Creek, ca. 15 m (5404, 5406, 5407,

5470); summit of peak NE of Mt. Weddell, 335 m (5565);

summit of Mt. Weddell, 380 m (5556).

Telaranea pseudozoopsis (Herz.) Fulf. Figure 33.

Lepidozia pseudozoopsis Herz. in Skottsberg, Nat. Hist.

Juan Fernandez Is., Easter I. 2(5): 723./ 5. 1942.

Telaranea pseudozoopsis (Herz.) Fulf., Brittonia

15: 71. 1963. Original material: Juan Fernandez

Is., Mas a Tierra, Centinela, 530 m, Skottsberg

227 (jE)-cited in Fulford (1966) (non vidi).

ECOLOGY— Rather common in the Falklands,

typically in moist, shaded situations (moist rock

crevices, stream banks, etc.) in dwarf shrub heaths,

Hebe associations and (more rarely) in Cortaderia

heaths. It is rather common on the bases of tussock

grasses in tussock associations.

PHYTOGEOGRAPHY— Valdivian + Magellanian

+ Falkland — Falkland Is.; Tierra del Fuego;

80

FIELDIANA: BOTANY

southern Patagonian Channels, Brunswick Pen-

insula; Valdivian region; Mas a Tierra, Juan Fer-

nandez Is. (400-530 m, where recorded) (fig. 33).

LITERATURE RECORD (FALKLANDS): Smith,

Beauchene Is. (Smith & Prince, 1985).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Kidney Island: N of shanty (2756, 2793, 2794, 2800).

Stanley Region: Sapper Hill, 135 m (2395); Tumble-

down Mt., 155-230 m (2377 B); headwaters of Mullet

Creek Stream, ca. 60 m (3165, 3181C); Goat Ridge, ca.

180 m (3212, 3217). Mt. Usborne Region: The Gap,

275-290 m (2461-c. per. + 3); valley SW of Mt. Us-

borne, ca. 60 m (2648Q. WEST FALKLANDS. Fox

Bay Region: summit of East Head, 1 80 m (3423); sum-

mit of Fox Bay Mt., 307 m (3448, 3454). Hill Cove

Region: summit of West French Peak, 290 m (2951).

Westpoint Island: steep slope and cliffs facing The Wool-

ly Gut (29/9, 2922. 2927, 2925, 2939); near The Wa-

terfall, 30-90 m (2875A, 2876); W base of Black Hog

Hill, adjacent to Devils Nose, ca. 20 m (2597, 2893).

Weddell Island: Waterfall Valley, W of settlement, ca.

125m (3307).

Telaranea tetradactyla (Hook. f. & Tayl.) Hodgs.

Jungermannia tetradactyla Hook. f. & Tayl., London

J. Bot. 3: 386. 1844 (3: 286 [sic] in errore pro

386). Lepidozia tetradactyla (Hook. f. & Tayl.)

G. L. & N., Syn. Hep. 213. 1845. Mastigophora

tetradactyla (Hook. f. & Tayl.) Trev., Mem. Reale

1st. Lomb. Sci. Lett. Ill, 4: 397. 1877. Neolepi-

dozia tetradactyla (Hook. f. & Tayl.) Fulf. & J.

Tayl., Brittonia 11: 84. 1959. Telaranea tetra-

dactyla (Hook. f. & Tayl.) Hodgs., Rec. Domin.

Mus. 4: 106. 1962. Original material: Auckland

Is., Hookers. n. (NY), cited in Fulford ( 1 963a) (non

vidi).

Lepidozia lindenbergii Gott. in G. L. & N., Syn. Hep.

213. 1845. syn. fide Fulford (1963a). Mastigo-

phora lindenbergii (Gott.) Trev., Mem. Reale 1st.

Lomb. Sci. Lett. Ill, 4: 397. 1877. Original ma-

terial: New A\i hi ml. without specific locality, "ex

hb. Taylor," Hooker s.n. (G, NY), cited in Fulford

(1963a)(m>w vidi).

Lepidozia disticha Steph., Kongl. Svenska Vetenskap-

sakad. Handl. 46(9): 62. / 24a-b. 1911, syn. fide

Fulford (1963a). Neolepidozia disticha (Steph.)

Fulf. & J. Tayl., Brittonia 1 1: 84. 1959. Original

material: Juan Fernandez Is., Mas a Tierra,

Skottsberg 117 (G), cited in Fulford ( 1 963a) (non

vidi).

Lepidozia effusiseta Steph., Spec. Hep. 6: 325. 1922,

syn. fide Fulford ( 1 963a). Original material: Juan

Fernandez Is.; Chile, Prov. Magallanes, I. Des-

olation, Skottsberg (G), cited in Fulford (1963a)

(non vidi).

REMARKS— Fulford (1963a, p. 277; 1966, pp.

206, 241) stated that the original description and

figures of Lepidozia fernandeziensis and L. disticha

(syn. of Telaranea tetradactyla) were interchanged

.

32

FIG. 32. Distribution of Telaranea plumulosa (Lehm.

& Lindenb.) Fulf.

FIG. 33. Distribution of Telaranea pseudozoopsis

(Herz.) Fulf.

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

81

in Stephani (1911). Herzog (1954) was apparently

not aware of Stephani's error when citing (p. 51)

"Lepidozia (Telaranea) fernandeziensis St. (= L.

effusiseta St.)-" Fulford (1966) has cited the Her-

zog error as a misidentification ("Lepidozia [Te-

laranea] fernandeziensis Herzog, Rev. Bryol. Li-

chenol. 23: 51. 1954 (in error)") in the synonymy

of Lepidozia chordulifera.

PH YTOGEOGRAPHY— Amphipacific Temper-

ate—(?) Falkland Is.; Tierra del Fuego; Patagonian

Channels; Valdivian Region (West Patagonia north

to 39°48'S); Juan Fernandez Is.; Auckland Is.; and

New Zealand. The species is common throughout

New Zealand and neighboring islands (Hodgson,

1956).

LITERATURE RECORDS (FALKLANDS)— Skotts-

berg, Port Stanley (Fulford, 1966).

Lepidoziaceae Species Excluded from Falklands

Bazzania peruviana (Nees) Trev.

This species was reported from the Falklands

by Stephani (Spec. Hep. 6: 480. 1924, as Masti-

gobryum skottsbergii Steph.) by Kiihnemann ( 1 949)

and by Bonner (1963). I have failed after several

attempts to locate this specimen. However, a label

of a specimen from Geneva of a 1 908 collection,

Skottsberg 50, from Mas a Tierra, Juan Fernandez

Islands, originally bore the locality "Falkland Ins.,"

which had been crossed out. It is likely that Ste-

phani discovered his error when curating the

Skottsberg collections, but did not delete the rec-

ord from his Species Hepaticarum. The species

occurs in the Magellanian and Valdivian regions,

Juan Fernandez Islands, and Tristan da Cunha. It

appears not to grow in the Falklands.

Family CALYPOGEIACEAE

Calypogeiaceae (K. Mull.) H. Arnell in Holmberg,

Skand. Fl. 2(a), 189. 1928.

A cosmopolitan family with two genera, Caly-

pogeia, with ca. 90 species, and Metacalypogeia

with four species. One species of Calypogeia, C.

sphagnicola (H. Arnell & J. Perss.) Warnst. &

Loeske is bipolar in distribution; in the Southern

Hemisphere it is known from southern South

America, New Zealand, and Tasmania (Schuster,

1969b; Engel, 1978). It has not been recorded for

the Falklands.

Family ADELANTHACEAE

Adelanthaceae (J0rg.) Grolle, J. Hattori Bot. Lab.

35: 327. 1972. Tribe Adelantheae J0rg., Ber-

gens Mus. Skr. 16: 257. 1934.

The Adelanthaceae have two genera. Adelan-

thus, with 1 2 species, is well represented in south

temperate and Neotropical regions. There are two

species which reach Europe, and one is known only

from Borneo. Wettsteinia has four species, and is

known from southern South America, Juan Fer-

nandez, the Falkland Islands, New Zealand, and

the Indomalayan region.

Adelanthus

Adelanthus Mitt., J. Proc. Linn. Soc., Bot. 7: 243.

1864, nom. cons.

Key to Falkland Islands Species of Adelanthus

1 . Dorsal margin of leaf only weakly incurved; leaves without distinct differentiation of a vitta of

elongated cells; subapical cells of leaf with walls thin to slightly thickened. Leaf apex occasionally

retuse to short bifid; leaf base never auriculate A. tenuis

1 . Dorsal margin of leaf distinctly incurved; leaves on well-developed plants with ± distinctly developed

vittae of elongated cells; subapical cells of leaf with walls thick. Leaf apex rounded to angular, not

retuse or short bifid; leaves often ± auriculate at the ventral base 2

2. Leaf margins dentate A. lindenbergianus

2. Leaf margins entire A. integerrimus

82

FIELDIANA: BOTANY

Adelanthus integerrimus Grolle

Adelanthus integerrimus Grolle, J. Hattori Bot. Lab.

35: 340. / 4. 1972. Holotype: Chile, Prov. Ma-

gallanes, Punta Arenas, C. Mina Rica, 550 m,

Santesson M 746 (s, non vidi).

ECOLOGY— This species is sporadic in the Falk-

lands, occurring in moist feldmarks, high altitude

cliffs, and Cortaderia heaths; it was not found be-

low 455 m.

PHYTOGEOGRAPHY— Subantarctic in Distribu-

tion— A. integerrimus is known from Crozet Is.;

South Georgia; Falkland Is.; Tierra del Fuego;

southern Patagonian Channels (Brunswick Pen-

insula).

LITERATURE RECORDS (FALKLANDS)— //a/fe, Mt.

Adam (Grolle, 1972).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Mt. Usborne Region: SE slope of Mt. Usborne 2, ca. 455

m (26J 4); summit of Mt. Usborne 1, ca. 700 m (2496A,

2499-c. 8, 2501C, 2505B). WEST FALKLANDS. Port

Howard: pass SW of Mt. Maria summit, ca. 6 1 0 m (3128,

3135).

Syn. Hep. 653. 1847. Adelanthus sphalerus (Hook,

f. & Tayl.) Steph., Bull. Herb. Boissier II, 8 (8):

599. 1908 (= Spec. Hep. 3: 383). Original ma-

terial: Chile, Prov. Magallanes, I. Hermite, Hook-

er (w), cited in Grolle (1972) (non vidi).

Jungermannia ? haliotiphylla De Not., Mem. Reale

Accad. Sci. Torino II, 16: 217. pi. V. 1-7. 1855,

syn. cf. Schiffner(1890). Original material: Chile,

"Prov. Valparaiso, Valparaiso," Puccio s.n. (L,

MANCH, s), cited in Grolle (1972) (non vidi).

Plagiochila subviminea Steph. in Herzog, Biblioth. Bot.

21: 212. / 146 d. 1916, syn. fide Grolle (1972).

Original material: Bolivia, Herzog 285 3 'la (G), cit-

ed in Grolle (1972) (non vidi).

Adelanthus trollii Herz., Hedwigia 74: 9 1 ./ 6a~c. 1 934,

syn. fide Grolle (1972). Original material: Bolivia,

Challana, Mina Fabulosa, Troll 50/a (JE), cited

in Grolle (1972) (non vidi).

Adelanthus parvus Herz., Rev. Bryol. Lichen. 11: 18.

1938, syn. fide Grolle (1972). Original material:

Costa Rica, Prov. San Jose, C. de Las Vueltas,

2700-3000 m, Standley 43723/c (JE), cited in

Grolle (1972) (non vidi).

Plagiochila subviminea f. paramicola Herz., Rev. Bryol.

Lichen. 11: 12. 1 938, syn. fide Grolle (1 972). Lec-

totype (cf. Grolle, 1972): Costa Rica, Prov. San

Jose, C. de Las Vueltas, 2700-3000 m, Standley

& Valeria 43862 (JE, non vidi).

Adelanthus lindenbergianus (Lehm.) Mitt. Figure

34.

Only American synonyms are included here:

Jungermannia lindenbergiana Lehm., Linnaea 4: 367.

1829. Plagiochila lindenbergiana (Lehm.) G. L.

& N., Syn. Hep. 59. 1844. Adelanthus lindenberg-

ianus (Lehm.) Mitt., J. Proc. Linn. Soc., Bot. 7:

244. 1 864. Adelanthus magellanicus var. linden-

bergianus (Lehm.) Schiffn., Nova Acta Acad.

Caesar. Leop.-Carol. German. Nat. Cur. 60(2):

261. 1893. Lectotype (cf. Grolle, 1972): South

Africa, near Cape Town, Ecklon (s, non vidi).

Plagiochila magellanica Lindenb., Spec. Hep. 1(5):

163.1 843, syn. cf. Mitten (1859). Adelanthus ma-

gellanicus (Lindenb.) Mitt., J. Proc. Linn. Soc.

Bot. 7: 244. 1864. Calyptrocolea magellanica

(Lindenb.) Schust., J. Hattori Bot. Lab. 26: 287.

1963, nom. illeg. Lectotype (fide Grolle, 1972):

Chile, Prov. Magallanes, B. San Nicolas, Dumont

d'Urville (PC, non vidi).

Jungermannia unciformis Hook. f. & Tayl., London

J. Bot. 3: 457. 1844, syn. cf. Mitten (1859). Pla-

giochila unciformis (Hook. f. & Tayl.) G. L. & N.,

Syn. Hep. 653. 1847. Adelanthus unciformis

(Hook. f. & Tayl.) Spruce, J. Bot. 14: 200. 1876.

Adelocolea unciformis (Hook. f. & Tayl.) Evans,

Bull. Torrey Bot. Club 25: 409. 1898. Original

material: Chile, Prov. Magallanes, I. Hermite,

Hooker (BM, MANCH, w), cited in Grolle (1972)

(non vidi).

Jungermannia sphalera Hook. f. & Tayl., London J.

Bot. 3: 458. 1844, syn. cf. Mitten (1859). Plagi-

ochila sphalera (Hook. f. & Tayl.) G. L. & N.,

REMARKS— I have been unable to locate the

specimen on which the record of Adelanthus ma-

gellanicus from the Falklands was based. How-

ever, the type of Adelanthus magellanicus is con-

specific with A. lindenbergianus, but according to

Grolle (in ////.), nearly all other collections sub-

sequently determined as A. magellanicus belong

to other taxa. Presumably the Falkland report is

included among them. Plagiochila coriacea Steph.,

nom. herb., belongs here; it was collected at Port

Harriet by Skottsberg (Hassel de Menendez, 1 983a,

p. 123). See Engel ( 1978).

ECOLOGY— This common species has a wide

ecological amplitude, growing in rock crevices or

on a soil layer either over rock or under rock over-

hangs of outcrops in the dwarf shrub heath asso-

ciation as well as the sheltered high altitude cliffs.

The taxon also grows on soil in the Cortaderia

(commonly under grass cover) and dwarf shrub

heaths. Further, it was gathered on soil at 700 m

in or at the margins of feldmark pools.

PHYTOGEOGRAPHY— Amphiatlantic Temper-

ate—Mascarene Is.; Reunion; Madagascar; South

Africa (1000-2000 m); Uganda (3100-3500 m);

Congo (3100 m); Tristan da Cunha (200-800 m);

Inaccessible I. (300 m); South Georgia; Falkland

Is.; Tierra del Fuego; Patagonian Channels; Val-

divian region; Juan Fernandez Is.; Andes north to

Mexico; and western Ireland (450-700 m) (fig. 34).

According to Grolle (1972a), the Australasian

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

83

34

FIG. 34. Distribution of Adelanthus lindenbergianus (Lehm.) Mitt. North to Venezuela and Costa Rica, plus

Ireland (after Grolle, 1969).

records of A. magellanicus are actually A. occlusus

and that from lies de Kerguelen is Jamesoniella

sp.

Among the Falkland taxa, it is one of the two

(with Chiloscyphus semiteres) strictly austral taxa

that is disjunct in the British Isles (see fig. 55).

LITERATURE RECORDS (FALKLANDS)— Anony-

mous (Kuhnemann, 1937, 1949 as A. unciformis);

Lechler (Mitten, 1864 as /I. magellanicus); Skotts-

berg. Port Stanley, Port Louis (Stephani. 1905a as

A. unciformis). Port Louis (Grolle, 1972a as A.

lindenbergianus), Port Stanley (Skottsberg, 1913

as A unciformis); Skottsberg & Halle, Weddell Is.,

Port Harriet (Stephani, 1911 as A. unciformis).

FALKLAND SPECIMENS SEEN -EAST FALKLANDS.

Port William Region: N side of Gypsy Cove, sea level

(3241). Stanley Region: near Sapper Hill, 29 Oct. 1907,

Halle & Skottsberg 1 13 as Frullania boveana (s); S slopes

of Mt. William, 185 m, Longton & Smith, Longton 983

(AAS, F); Tumbledown Mt., 155-230 m (2373, 2382A);

Goat Ridge, ca. 180 m (3190); N peak of Two Sisters,

250-275 m, Dollman s.n. (Greene 356 la, 3563b) (AAS,

F); summit ridge of N peak of Two Sisters, 245-290 m

(2733— c. per., 2734); at summit of Mt. Kent, 455 m

84

FIELDIANA: BOTANY

JJE

FIG. 35. Adelanthus tenuis Engel & Grolle. Plant; note ventral-intercalary branching and leaf apex variability,

x 10 (from holotype).

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

85

(2763). Mt. Usborne Region: gap between Ml. Usborne

2 and Table Rock, ca. 440 m (2625C. 2626B. 2628.

2632, 2634A, 2640D); SE slope of Mt. Usborne 2, ca.

455 m (2600); ridge between Mts. Usborne 1 & 2, 685

m (2532-c. per., 2554A); The Gap, 275-290 m (2462B,

2463. 2469. 2475); below The Gap, ca. 90 m (2562A).

WEST FALKLANDS. Port Howard: Freezer Rocks, on

E slope of Mt. Maria, 320 m (3146. 3153); pass SW of

Mt. Maria summit, ca. 610 m (3080A, 3133). Fox Bay

Region: summit of East Head, 180 m (3417). Mt. Adam:

E side of summit ridge, 670-700 m (3019A); E slope of

main peak, ca. 6 1 0 m ( 3003 A). Hill Cove Region: summit

of East French Peak, 305 m (2964 A— c. per.); summit

of Mt. Fegen, 335-360 m (3079). Weddell Island: sum-

mit of peak NE of Mt. Weddell, 335 m (3369B); summit

of Mt. Weddell, 380 m (3335B, 3345. 3350, 3353).

Ad elan thus tenuis Engel & Grolle. Figures 35-36.

Adelanthus tenuis Engel & Grolle in Grolle, J. Hattori

Bot. Lab. 35: 333. / 1-2. 1972. Holotype: Chile,

Prov. Magallanes, Brunswick Peninsula, La. El

Parrillar region, ca. 365 m, 21 Dec. 1967, Engel

2093 (MSC!).

REMARKS— Because the species plays a conspic-

uous role in the Falkland hepatic flora, yet may

be confused with other hepatics in these islands,

a description is provided below.

Plants caespitose, rarely as scattered individuals

among other Hepaticae, erect, to 2.3 cm tall, light

green to golden brown or brown, often light green

toward apex of plant and brown toward base, nitid.

Branches frequent, exclusively of ventral-inter-

calary type, leafy branches stiff, abruptly erect or

prostrate and becoming erect, rarely becoming fla-

gelliform toward the apex; stolons often arising

from near base of ventral-intercalary branch, pros-

trate to occasionally positively geotropic, with small

scalelike leaves and scattered rhizoids, the stolons

occasionally becoming upright and producing en-

larged leaves.

Stems 165-235 Mm wide, 135-200 Mm thick, in

surface view the cortex cells short to long rect-

angular, 1 3-23 Mm wide, 20-85(105) Mm long, with

walls slightly to distinctly thickened, the end walls

transverse to oblique. Stems in section 10-16 cells

in diameter, cortex in 1-2 layers, the cortex poorly

defined or distinct and then with very thick- walled

cells, the cells 9-18 Mm thick, the cuticle smooth;

medulla cells (13)16-30(33) Mm wide, thin walled

but with slightly thickened corners.

Leaves (0.5)0.55-1.1(1.2) mm long and wide,

transverse to succubously oriented (succubous in

robust plants), the insertion slightly to distinctly

succubous, occasionally slightly recurved at ven-

tral end; leaves erect to spreading, occasionally

recurved, distant to imbricate, concave (often

strongly so near dorsal base) to convex, orbicular

to obliquely wide-ovate or obliquely wide-obovate

in shape, the margins entire; apex acute to broadly

rounded to truncate, occasionally slightly bifid and

then divided to 0.15, the lobes acute to broadly

rounded, the sinus triangular to rounded, the angle

obtuse to narrowly rounded to lunate; dorsal mar-

gin slightly to occasionally strongly curved, often

repand, slightly to strongly inflexed, especially to-

ward the base, not to very short decurrent; ventral

margin strongly curved, usually more strongly

curved than dorsal margin, the ventral margin often

repand, plane to recurved, not to very short de-

current.

Leaf cells subquadrate to subrectangular, thin

to slightly thickened, trigones absent to medium,

occasionally with intermediate thickenings toward

base; subapical cells 14—23 Mm wide and long, me-

dian cells 14—24 Mm wide, 17-28 Mm long, the

basal cells longer, 28—47 Mm long; marginal cells

with nodular thickenings on outer radial walls, the

basal marginal cells in 1-3 rows of shorter cells,

10-18 nm long, bordering the elongated basal-me-

dian cells, but the leaves without a distinct differ-

entiation of a vitta; cuticle smooth; oil-bodies not

seen.

Underleaves rudimentary, inconspicuous, con-

sisting of only a small cluster of cells in close prox-

imity to the ventral base of leaves. Gemmae rather

rare, green, thin walled, ovate to elliptic, mostly

1 -celled.

Opposite:

FIG. 36. Adelanthus tenuis Engel & Grolle. 1-44, Leaves (those with same letter were obtained from same

population), x 23. 45, Median leaf cells, x 545. 46, Basal-median leaf cells, x 515. 47, Stem, cross section, x 193. 48,

Portion of upper, ventral leaf margin, x515. 49, Cortical cells, surface view, x515. (Figs. 1, 8, from Engel 3232;

2-4, from Engel 2533A; 3, 5-7, from Engel 3103; 9, 1 1, from Engel 2725; 10, 17, from Engel 2457; 12, 13, from

Engel2773; 14, 20, from Engel 3221; 15, 1 6, from Engel 3343B; 18, 38, from Engel 3452; 19, 22, from Engel 2966;

21,31, from Engel 3003B; 23, 35, from Engel 3114; 24, 33, 39, 41, 45-49, from Engel 2093 (holotype); 25, 34,

from Engel 2108C; 26, 44, from Engel 3453; 27, 40, 43, from Engel 3324A; 28, 29, 32, from Engel 3444; 30, 42,

from Engel 2732; 36, 37, from Engel 2640C. Figs. 24-25, 33-34, 39, 41, and 45-49 from southern Chilean material,

remainder from Falkland Is. material.)

86

FIELDIANA: BOTANY

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

87

37

FIG. 37. Wettsteinia densiretis (Herz.) Grolle.

Plants dioecious. Androecia not seen. Gynoecia

on reduced ventral-intercalary branches which arise

from lower portion of plant, mature gynoecia not

seen.

VARIATION— Leaves, while usually oriented

subsuccubous, may be transverse or, in robust

plants, strongly succubous. The leaves vary con-

siderably in size and configuration (figs. 36, 1-44).

While the leaf apices exhibit a wide range of vari-

ability, usually at least some leaves with slightly

bifid apices are present on a given plant; bifid api-

ces are an important taxonomic feature, and are

useful in separating A. tenuis from "look-alikes."

The ventral-intercalary branches exhibit consid-

erable variation. Normal leafy branches may be-

come flagelliform toward the apex, or conversely

a stoloniform branch may become erect and pro-

duce leaves typical of normal leafy shoots. In the

latter, the stoloniform branches may be quite

lengthy (to 1 1 mm) prior to becoming negatively

geotropic and producing normal leaves.

ECOLOGY— This species appears to be restricted

to relatively high altitudes in the southern cold

temperate region. In the Brunswick Peninsula it

was collected at ca. 365 m in an open Sphagnum

bog. In the Falklands this species is rather com-

mon above 245 m, and occurs on soil under rock

cover, in rock crevices, under ledges, etc., of out-

crops in Cortaderia as well as dwarf shrub heath

associations. Occasionally it may be found on soil

in nonoutcrop situations in these associations,

growing with Gackstroemia magellanica and Ade-

lanthus lindenbergianus, or on soil over rock where

it may be associated with Balantiopsis bisbifida

and a filmy fern, Serpyllopsis caespitosa (Gaud.)

v. d. Bosch.

PH YTOGEOGRAPHY — M ayi'l 1 a n i a n- 1 a I k I a n d —

Falkland Is.; Tierra del Fuego; and Brunswick Pen-

insula (La. Parrillar region). It is of phytogeograph-

ical interest to note that the New Zealand A. gem-

miparus (Schust.) Hodgs. is a very close ally of this

species.

FALKLAND SPECIMENS SEEN -EAST FALKLANDS.

Port William Region: summit ridge of Mt. Low, ca. 245

m (3220, 3221, 3224 A, 3232). Stanley Region: Sapper

Hill, 2 km W of Port Stanley, 60 m, Ochyra 2852/80

(F); near summit of Tumbledown Mt., 230 m, Longton

& Smith, Longton 972 (AAS, F); N peak of Two Sisters,

275 m, Dollman s.n. (Greene 3564a) (AAS, F); summit

ridge of N peak of Two Sisters, 245-290 m (2725, 2752/1);

summit of Mt. Kent, 455 m (2761 A —c. S, 2773— c.

young per.). Mt. Usborne Region: gap between Mt. Us-

borne 2 and Table Rock, ca. 440 m (2640Q; ridge be-

tween Mts. Usborne 1 & 2, 685 m (2533A, 2554B); The

Gap, 275-290 m (2457). WEST FALKLANDS. Port

Howard: pass SW of Mt. Maria summit, ca. 610 m

(3103— c. $, 3114). Fox Bay Region: summit of Fox Bay

Mt., 307 m (3444-c. young per., 3452, 3453A). Mt.

Adam: E slope of main peak, ca. 610 m (3003 B). Hill

Cove Region: summit of East French Peak, 305 m (2966—

c. young per., 2969); summit of Mt. Fegen, 335-360 m

(3077). Wedded Island: Waterfall Valley, W of settle-

ment, ca. 125 m (3324 A— c. young per.); summit of Mt.

Weddell, 380 m (3343B).

Adelanthus Species Excluded from Falklands

Adelanthus falcatus (Hook.) Mitt.

The report of Adelanthus falcatus from the Falk-

land Islands in Stephani (1905a) is a misdeter-

mined specimen of A. lindenbergianus, fide Grolle

(in lift. ; specimens from S examined). Adelanthus

(Calyptrocolea) falcatus is a species of New Zea-

land (type from Dusky Bay, New Zealand, leg.

Menzies), Tasmania and (presumably), southeast

Australia (Grolle, 1972a).

Wettsteinia

Wettsteinia Schiffn., Ann. Jard. Bot. Buitenzorg.

Suppl. 2: 44. 1898.

88

FIELDIANA: BOTANY

Wettsteinia densiretis (Herz.) Grolle. Figure 37.

Tylunanthus densiretis Herz. in Skottsberg, Nat. Hist.

Juan Fernandez Is., Easter I. 2(5): 712. / 3a-d.

1942. Wettsteinia densiretis (Herz.) Grolle, J.

Hattori Bot. Lab. 28: 99. 1965. Lectotype (fide

Grolle 1965c): Juan Fernandez Is., Mas Afuera,

Camp Correspondencia, 1140 m, 1917, Skotts-

berg 92 (s, non vidi).

ECOLOGY— This species is of rather frequent oc-

currence in crevices of outcrops, as well as on soil

in nonoutcrop situations in dwarf shrub heaths. It

more rarely occurs on soil in Cortaderia heaths.

PHYTOGEOGRAPH Y — Valdivian-Falkland —

Falkland Is.; Valdivian region (ca. 40°S, 73°30'W);

and Juan Fernandez Is., Mas Afuera (1 140 m) (see

fig. 37).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Port William Region: summit ridge of Mt. Low, ca. 245

m (3225); ridge on Engineer Point Peninsula, 18-25 m

(2823). Stanley Region: Tumbledown Mt., 155-230 m

(2377 A. 2381); Goat Ridge, ca. 180 m (3196, 3204);

summit ridge of N peak of Two Sisters, 245-290 m

(2701 A). Mt. Usborne Region: below The Gap, ca. 90

m (2559, 2563). WEST FALKLANDS. Weddell Island:

Waterfall Valley, W of settlement, ca. 125 m (3316);

summit of peak NE of Mt. Weddell, 335 m (3369C,

3395).

Family CEPHALOZIACEAE

Cephaloziaceae Mig., Krypt.-Fl. Deutschl. ... 1:

465. 1904.

The family has 14 genera and is worldwide in

distribution. Only two genera— Cephalozia and

Metahygrobiella— occur in south temperate-sub-

antarctic areas, and both are in the Falklands.

Cephalozia is worldwide and poorly represented

in austral regions, while Metahygrobiella is pre-

dominantly south temperate.

Cephalozia

Cephalozia (Dum.) Dum., Recueil Observ. Jun-

germ. 18. 1 835. Jungermannia sect. Cephalo-

zia Dum., Syll. Jungerm. Eur. Indig. 60. 1831.

Cephalozia badia (Gott.) Steph.

Jungermannia badia Gott., Ergebn. Deutsch. Polar-

Exped. 2(16): 452. pi l.f. 1-5. 1890. Cephalozia

badia (Gott.) Steph., Bull. Herb. Boissier II, 8(7):

483. 1908 (= Spec. Hep. 3: 313). Original ma-

terial: South Georgia, Koppenberg, 10 Feb. 1883,

Will 37 (o), cited in Fulford (1968) (non vidi).

ECOLOGY— In the Falklands, this species was

collected on a rock in a feldmark pool (ca. 700 m)

and on soil under rock cover of a sheltered high

altitude cliff.

PHYTOGEOGRAPH Y — Antarctic in Distribu-

tion—Antarctic Peninsula; Nelson and King

George Islands (cf. Ochyra & Vana, 1 989); South

Shetland Is. (Allison & Smith, 1973; Smith, 1979);

South Sandwich Is. (Longton & Holdgate, 1979);

South Georgia; Falkland Is.; and Tierra del Fuego

(Lago Fagnano). Stephani (1911) stated that the

Lago Fagnano collection is from a "Berge am Wes-

tende von Lago Fagnano."

LITERATURE RECORD (FALKLANDS)— Anony-

mous (Futford, 1968).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Mt. Usborne Region: summit of Mt. Usborne 1, ca. 700

m (24968-c. per. + S). WEST FALKLANDS. Port

Howard Region: pass SW of Mt. Maria summit, ca. 610

m (3104B).

Notes on Cephalozia Species

1. Cephalozia bicuspidata (L.) Dum.

There are three reports of this species from the

Falklands, all based upon a Hooker collection(s).

The taxon was reported (as Jungermannia) in

Hooker and Taylor (1844), Taylor and Hooker

(1847), and Gottsche et al. (1847). The species is

widespread in the Northern Hemisphere, but

Schuster ( 1 974) extended the range to include New

Zealand with C. bicuspidata subsp. austrigena

Schust. Vana (1988) added several more austral

localities, namely South Africa, Marion Is., Crozet

Is., Tristan da Cunha, and southern South Amer-

ica (Brunswick Peninsula). With this emended dis-

tribution, it is quite reasonable to expect that C.

bicuspidata will eventually be recorded for the

Falkland Is., but I have not as yet seen material.

Metahygrobiella

Metahygrobiella Schust., Bryologist 64: 205. 1961.

Metahygrobiella tubulata (Hook. f. & Tayl.) Schust.

ex Engel

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

89

Jungermannia tubulata Hook. f. & Tayl., London J.

Bot. 3: 463. 1844. Cephalozia tubulata (Hook. f.

& Tayl.) Trev., Mem. Reale 1st. Lomb. Sci. Lett.

Ill, 4: 4 1 7. 1 877. Metahygrobiella tubulata Schust.

ex Fulf., Mem. New York Bot. Card. 1 1(3): 322.

1968 nom. nud., pro syn. Metahygrobiella tubu-

lata (Hook. f. & Tayl.) Schust. ex Engel, Bryol-

ogist 79: 514. 1976. Original material: Falkland

Is., Hooker s.n. (BM!).

REMARKS— See comments in Engel (1978), which

were based upon Falkland Islands plants (includ-

ing type material).

ECOLOGY— Rather rare in the Falklands. I en-

countered the species in a creek in a Cortaderia

heath and in wet depressions of dwarf shrub heaths,

while Longton and Smith found the plant in a

Sphagnum bog.

PHYTOGEOGRAPHY— Valdivian + Magellanian

+ Falklands— Falkland Is.; Tierra del Fuego; north

in Patagonian Channels to 44°1 9'S (Cerro Tesoro);

and Valdivian West Patagonia.

LITERATURE RECORDS (FALKLANDS)— Anony-

mous (Kuhnemann, 1937, 1949;Bonner, 1963, as

Cephalozia), Port Louis (Fulford, 1 968 as Cepha-

lozia). Hooker (Fulford, 1968 as Cephalozia; G.

L. & N., 1847, as Jungermannia; Taylor & Hook-

er, 1847 as Jungermannia; Stephani, 1908 as

Cephalozia). Skottsberg, Port Stanley (Stephani,

1905a, 1908; Skottsberg, 1913; Fulford, 1968 as

Cephalozia).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Stanley Region: headwaters of Mullet Creek stream, ca.

60 m (3/55); between Mt. William and Tumbledown

Mt., 170 m, Longton & Smith, Longton 980 (AAS, F);

Goat Ridge, ca. 180 m (3207— c. per., 3214). Mt. Us-

borne Region: gap between Mt. Usborne 2 and Table

Rock. ca. 440 m (2627-c. 6). WEST FALKLANDS.

Hill Cove Region: gap between French Peaks, 200-215

m (29798-c. per.).

Family CEPHALOZIELLACEAE

Cephaloziellaceae Douin, Bull. Soc. Bot. France

29: 1, 5, 13. 1920.

Of the seven genera in this family, Allisoniella

(five species) and the monoiypicAmphicephalozia

occur only in south temperate regions; both are

present in southern South America but are lacking

from the Falklands. Cephaloziella (ca. 125 species

in the broad sense) is worldwide, including both

the antarctic and arctic, and occurs on the Falk-

lands.

Cephaloziella

Cephaloziella (Spruce) SchifFn. in Engler & K.

Prantl, Die Natiir. Pflanzenfam. 1(3,1): 98.

1893, nom. cons. Cephalozia subg. Cephalo-

ziella Spruce, On Cephalozia 62. 1882.

Cephaloziella dusenii Steph.

Cephaloziella dusenii Steph., Bih. Kongl. Svenska Ve-

tenskapsakad. Handl. 26 (III, 6): 49. 1900. Ceph-

alozia dusenii (Steph.) Steph., Bull. Herb. Boissier

II, 8(7): 504. 1908 (= Spec. Hep. 3: 334). Alobiella

dusenii (Steph.) Steph., Bull. Herb. Boissier II,

8(8): 571. 1908 (= Spec. Hep. 3: 355), nom. illeg.,

non Alobiella dusenii Steph., Bih. Kongl. Svenska

Vetenskapsakad. Handl. 26 (III, 6): 48. 1900 (=

Alobiella stephanii Bonn., Candottea 14:98. 1953).

Lectotype (fide Bonner, 1953): Chile, Prov. Val-

divia, Corral, 5 June 1896, Dusen 94 (G!; isolec-

totypes: s!— c. per., UPS!— c. per.).

Cephalozia setistipa Steph., Spec. Hep. 6: 440. 1924,

syn. nov. Original material: Falkland Is., Rabbit

I., Halle s.n. (G 15158!— c. young per. + <J; ap-

parently dioecious).

REMARKS— Fulford (1976) placed this name in

synonymy with Cephaloziella exiliflora (Tayl.)

Douin while Schuster (1971b, 1980a) included C.

dusenii in the synonymy of C. byssacea (A. Roth.)

Warnst. The materials I have seen, including the

lectotype of C. dusenii, are dioecious and therefore

cannot be placed with C. exiliflora, an autoecious

or paroecious plant. Materials from the Brunswick

Peninsula (Engel, 1 978) and Falkland Islands agree

with the lectotype of C. dusenii (sensu Bonner,

1953), but whether or not the lectotype belongs in

the synonymy of C. byssacea I believe still is open

to question, for reasons outlined in Engel (1978).

Future studies indeed may reveal the two taxa as

conspecific, but for the present I prefer to recognize

C. dusenii as a distinct species that occurs in South

Georgia, the Falklands, and southern South Amer-

ica. For additional comments see Engel (1978).

The portion of Cephalozia setistipa available for

study consisted of several sterile shoots, several

shoots bearing young gynoecia, and a single an-

droecial shoot. I found no suggestion that the an-

droecial shoot was attached to a gynoecial axis;

the material thus appears to be dioecious, which

agrees with the protologue of Cephalozia setistipa.

ECOLOGY— This species is of rather common oc-

currence in dwarf shrub heaths, where it occurs on

soil over rock, in rock crevices, and under rock

ledges or Empetrum cover. It also was found on

the basal portion of tussock (Poa flabellata asso-

90

FIELDIANA: BOTANY

elation), and then often in association with Chi-

loscyphus lentus.

PHYTOGEOGRAPHY— Valdivian + Magellanian-

Falkland— South Georgia; Falkland Is.; Patago-

nian Channels; Valdivian region (West Patagonia

north to 39°52'S).

LITERATURE RECORDS (FALKLANDS)— Anony-

mous (Kuhnemann, 1937, 1949 as Cephalozia);

Skottsberg, Port Stanley, Warrah River (Bonner,

1953 as Alobiella stephanii), Port Stanley (Skotts-

berg, 1913 as Cephalozia); Skottsberg & Halle,

Port Stanley, Warrah River (Stephani, 1911 as

Cephalozid).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

1 843, Hooker s.n.. as Jungermannia byssacea (FH). Kid-

ney Island: Pettingill 13. 25 (CINC, F); N of shanty (2790).

Port William Region: S shore of Cape Pembroke Pen-

insula, near Surf Bay, sea level (2438); N side of Gypsy

Cove, sea level (3249). Stanley Region: Port Stanley,

1 907, Skottsberg 22 (o); Sapper Hill, 135m (241 1, 2418-

c. per.); Tumbledown Mt., 155-230 m (23788—c. per.).

WEST FALKLANDS. Warrah River Region: 17 Dec.

1907, Halle & Skottsberg 100 (s, UPS); 1907, Skottsberg

39 (o). Fox Bay Region: summit of East Head, 180 m

(3438B—C. per.); valley E of Sulivan House, ca. 75 m

(3538). Hill Cove Region: summit of West French Peak,

290 m (2946- c. per.). Weddell Island: summit of Mt.

Weddell, 380 m (3352-c. per.).

Notes on Cephaloziella Species

1 . Cephaloziella varians (Gott.) Steph.

This species was originally described by Gottsche

(1890) as a Jungermannia, based on a Will col-

lection from South Georgia. It was transferred to

Cephalozia by Stephani (Result. Voyage S. Y. Bel-

gica 6[5]: 5. 1901 [1902]), and later to Cephalo-

ziella by Stephani (Wiss. Ergebn. Schwed. Siid-

polar Exped. 1901-03. 4[1]: 4. 1905). Note that

even though Stephani placed the species in Cepha-

loziella in 1905, he chose to treat the name as a

Cephalozia in Species Hepaticarum ( 1 908), as well

as in Stephani (1911), where he reported the plant

for the Falkland Islands. Kuhnemann (1937, 1949)

recorded the species from the Falklands in his cat-

alogs. I have not seen material of the species. It

was placed as a synonym of Cephaloziella exili-

flora (Tayl.) Douin by Fulford (1976, p. 413).

Ochyra and Vana (1989), however, recently dis-

cussed Cephaloziella varians and included in its

synonymy C. arctica Bryhn & Douin. With this

revised circumscription, C. varians becomes bi-

polar in distribution. Ochyra and Vana stated that

"Cephaloziella varians is widespread throughout

the maritime Antarctic and it is the only species

of hepatic known to occur on the Continent ex-

cluding the Antarctic Peninsula." Interestingly,

Schuster and Damsholt (1974, p. 326) stated, "C.

arctica s.l. is perhaps the commonest and most

widespread Cephaloziella in Greenland and is one

of the most abundant species of Hepaticae found

there." Cephaloziella varians is an autoecious plant.

Cephaloziella Species Excluded from Falklands

1. Cephaloziella byssacea (A. Roth.) Warnst.

This species was reported from the Falkland

Islands by Hooker and Taylor (1844) and Taylor

and Hooker (1847), both as Jungermannia bys-

sacea, and Kuhnemann ( 1 949). These records are

all based upon a Hooker collection which is ac-

tually Cephaloziella dusenii. The species may in-

deed occur in the Falklands, if, after further in-

vestigation, C. dusenii is found to be a synonym

(see comments under C. dusenii).

2. Cephaloziella starkei (Funck ex Nees) Schiffn.

This taxon was reported from the Falklands by

Kuhnemann (1937), who did so on the basis of

the Taylor and Hooker (1847) reference of Jun-

germannia byssacea from the Falkland Islands (see

above). Kuhnemann (1937) included J. byssacea

as a synonym of C. starkei. In his later catalog,

Kuhnemann (1949) used the name Cephaloziella

byssacea for a Falkland report, without mention

of C. starkei. The Hooker collection of Cephalo-

ziella byssacea from the Falkland Islands is ac-

tually a specimen of Cephaloziella dusenii. Grolle

( 1 970) has shown Jungermannia starkei Funck ex

Nees and Cephaloziella starkei (Funck ex Nees)

Schiffn. are illegitimate names, and includes them

in the synonymy of Cephaloziella divaricata (Sm.)

Schiffn.

Family ANTHELIACEAE

Antheliaceae Schust., J. Hattori Bot. Lab. 26: 236.

1963.

A family with two genera, Anthelia and Grollea,

neither of which occurs on the Falklands. Anthelia

has two arctic-alpine species, one (A. juratzkana

(Limpr.) Trev.) bipolar in distribution and occur-

ring in Tierra del Fuego (see Schuster, 1 974). Grol-

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

91

lea, with only G. antheliopsis Schust., occurs in

southern South America (Hassel de Menendez,

1980b; Schuster, 1972a).

Family LOPHOZIACEAE

Lx>phoziaceae (J0rg.) Vand. Berg, in Robyns, Fl.

Gen. Belgique 1: 221. 1956, nom. cons.

The Lophoziaceae, containing 1 7 genera, are best

developed in the cool and cold portions of the

Northern Hemisphere. The largest genus, Lopho-

zia, is widespread in boreal-subarctic-arctic re-

gions, and only sporadically represented in the

tropics and south temperate zone. Also notable is

the rather large number of very small south tem-

perate-subantarctic and tropical genera. Notho-

strepta (2 species) and Roivainenia (1 species) are

examples.

Anastrophyllum

Anastrophyllum (Spruce) Steph., Hedwigia 32: 1 39.

1893. Jungermannia subg. Anastrophyllum

Spruce, J. Hot. 14: 235. 1876.

Anastrophyllum cilia turn Steph.

Anastrophyllum ciliatum Steph., Hedwigia 32: 139.

1893 ("Anastrophylium"). Sphenolobus ciliatus

(Steph.) Steph., Bull. Herb. Boissier 2(2): 175.

1902 (= Spec. Hep. 2: 167). Original material:

Argentina, Terr. Tierra del Fuego, I. de los Es-

tados, Spegazzini s.n. (G!).

Anastrophyllum pampaninii Gola, Nuovo Giorn. Bot.

Ital. II, 29: 165. pi. 2,f. 1-2. 1923, syn.fideEngel

(1978). Original material: Chile, Prov. Magal-

lanes, B. Angelito, 300-400 m, 10 Feb. 1913, De

Gasperi (FI!).

REMARKS— This species is closely related to An-

astrophyllum crebrifolium, a species of southern

South America and northern Andes (see Engel,

1978).

ECOLOGY— Usually mixed with other Hepaticae

above 440 m in the Falklands; fairly common in

sheltered high altitude cliffs under overhangs or

on cushions of Azorella selago. Also collected in

a feldmark (700 m), where it was associated with

pools.

PHYTOGEOGRAPHY — M agellanian-Falkland —

South Georgia; Falkland Is.; southern Magellanian

region (Isla de los Estados, Brunswick Peninsula,

and Volcan Inga, Canal Gajardo).

FALKLAND SPECIMENS SEEN -EAST FALKLANDS.

Mt. Usborne Region: gap between Mt. Usborne 2 and

Table Rock, ca. 440 m (2630— c. <3); ridge between Mts.

Usborne 1 & 2, 685 m (2543 A, 2554E); summit of Mt.

Usborne 1, ca. 700 m (2489A, 2491 A, 2496Q. WEST

FALKLANDS. Port Howard: pass SW of Mt. Maria

summit, ca. 610 m (3080B). Mt. Adam: side of summit

ridge, 670-700 m (3014B).

Anastrophyllum Species Excluded from Falklands

Anastrophyllum subcomplicatum (Lehm. & Lin-

denb.) Steph.

This species was reported from the Falklands

by Herzog (1926), who confused the type locality

"Insulis Marianis" (Mariana Islands) with the

Falklands (= lies Malouinae, Islas Malvinas).

Andrewsianthus

Andrewsianthus Schust., Rev. Bryol. Lichenol. 30:

66. 1961.

Key to Falkland Islands Species of Andrewsianthus

1. Leaves often canaliculate or conduplicate, 0.25-0.3(0.5) bifid; leaf lobes acutely triangular, the angle

of lobe outline 15°-65°(800), the apices occasionally acuminate or apiculate; sinus usually sharply

triangular, occasionally narrowly rounded, rarely broadly rounded, never lunate; leaf insertion trans-

verse, at most occasionally weakly succubous in dorsal half; ventral-intercalary branches very rare

A. australis

1. Leaves usually plane to slightly concave, exceptionally canaliculate, never conduplicate, 0.1-0.25

bifid; leaf lobes acutely to obtusely triangular, the angle of lobe outline 45°-l 1 5°, the apices occasionally

somewhat apiculate, never acuminate; sinus broadly rounded to lunate, occasionally triangular; leaf

insertion often distinctly succubous; ventral-intercalary branches occasional A. planifolius

92

FIELDIANA: BOTANY

14 15^ 16 17 18 19 20

21 22 23 24 25 26 27 28 29 30 31 32 33

FIG. 38. Andrewsianthus australis Engel. 1, Stem, cross section, x 490. 2, Portion of plant with main axis becoming

flagelliform, and with lateral-intercalary branch, dorsal view, xlO. 3, Median leaf cells with cuticular striations

indicated in upper left, x5l5. 4, Apical portion of dorsal lobe, x5l5. 5, Leaf of main axis, x3i. 6, Ventral view of

stem showing sessile, rudimentary underleaf, x5l5. 7, Ventral cortical cells, surface view, x5l5. 8, Slime papillae

of basal region of dorsal margin of leaf, x515. 9-23, Leaves of main axis, x20. (Figs. 1-5, 7, 15, 18, 20, 25 from

holotype; 6, 8, 14, 21, 22 from Engel 2219B. Chile, Prov. Magallanes, Pto. Cutter, N of copper mine; 10-12 from

Engel 216 1G, Chile, Prov. Magallanes, Pto. Cutter, between copper mine and river S of mine; 13, 19, 24, from Engel

3075, Falkland Is., Mt. Fegen; 16-17, 26, from Engel 3020B, Falkland Is., Mt. Adam; 23, 37, from Engel 2301 B.

Chile, Prov. Magallanes, Pto. Cutter, N side of copper mine; 28, 32-33, from Engel 3356. Falkland Is., Mt. Weddell;

29-31, from Engel 2743, Falkland Is., Mt. Kent.)

Andrewsianthus australis Engel. Figures 38-39.

Andrewsianthus australis Engel, Bryologist 75: 328. /

1-42. 1972. Holotype: Chile, Prov. Magallanes,

Cta. Amalia, 1 Oct. 1969, Engel 5411 A (MSC!-

c. per. + sporo. + <5).

REMARKS— See Engel (1972).

ECOLOGY— In the Falklands, gathered on soil

over rock, or in rock crevices of cliffs, or large rock

outcrops in dwarf shrub heaths above 335 m. On

a single occasion I found it on soil under rock cover

of sheltered high altitude cliffs.

PHYTOGEOGRAPHY — Magellanian-Falkland —

Falkland Is. and Patagonian Channels (north to

50°56'S, 73°52'W).

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

93

FIG. 39. Andrewsianthus australis Engel. 1, Portion of main axis with perianth, x 3 1 . 2, Seta, cross section, x 1 80.

3, Portion of perianth mouth with cellular detail indicated, x 180. 4, Spore, x515. 5, Capsule and portion of seta,

x 3 1 . 6, Capsule wall, cross section, x 5 1 5. 7, Capsule wall, inner layer, x 5 1 5. 8, Capsule wall, outer layer, x 5 1 5. 9,

Portion of axis bearing androecia, x31. (Figs. 1-8 from holotype; 9 from Engel 2743, Falkland Is., Mt. Kent.)

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Stanley Region: summit of Mt. Kent, 455 m (2743— c.

<J). WEST FALKLANDS. Mt. Adam: E side of summit

ridge, 670-700 m (3020B). Hill Cove Region: summit

of Mt. Fegen, 335-360 m (3075-c. per.). Weddell Is-

land: summit of peak NE of Mt. Weddell, 335 m (3356).

Andrewsianthus planifolius Engel. Figure 40.

Andrewsianthus planifolius Engel, Bryologist 75: 332.

/ 43-64. 1972. Holotype: Falkland Is., Goat Ridge,

ca. 180 m, 30 Jan. 1968, Engel 3187B (MSC!-C.

young gynoecia).

REMARKS— See Engel (1972a).

ECOLOGY-PHYTOGEOGRAPHY— Endemic to the

Falkland Islands, where it occurs mixed with Frul-

lania microcaulis.

FALKLAND SPECIMENS SEEN -EAST FALKLANDS.

Stanley Region: Goat Ridge, ca. 180 m (31878—c. young

gynoecia, 3192A).

94

FIELDIANA: BOTANY

16

FIG. 40. Andrewsianthus planifolius Engel. 1, Portion of main axis, x 3 1 . 2, Median leaf cells, x 5 1 5. 3-4, Stalked

and sessile slime papillae of basal region of dorsal margin of leaf, x 5 1 5. 5, Ventral view of stem showing rudimentary

underleaf, x5l5. 6, Cortical cells, surface view, with cuticular st nations indicated, in part, x5l5. 7, Stem, cross

section, x 5 1 5. 8, Apical portion of ventral lobe, x 5 1 5. 9-22, Leaves of main axis, x 3 1 . (Figs. 1-14, 20 from holotype;

15-19, 21-22 from Engel 3 192 A, Falkland Is., Goat Ridge.)

Cephalolobus

Cephalolobus Schust., Rev. Bryol. Lichenol. 34:

244. 1966. Cephalolobus Schust., J. Hattori

Bot. Lab. 26: 21 1, 266. 1963, nom. nud.

Cephalolobus scabrellus (Mass.) Schust.

Cephalozia scabrella Mass., Nuovo Giorn. Bot. Ital.

17: 233.p/. 20, f. 19. 1885. Sphenolobus scabrellus

(Mass-)Steph., Bull. Herb. Boissier 2(2): 170. 1902

(= Spec. Hep. 2: 162). Cephalolobus scabrellus

(Mass.) Schust., Rev. Bryol. Lichenol. 34: 250.

1966. Lectotype (fide Schuster, 1966a): Argen-

tina, Terr. Tierra del Fuego, I. de los Estados,

"Port Cook and Port St. John," 1 882, Spegazzini

(o, non vidi).

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

95

ECOLOGY— Quite rare in the Falklands, where I

found it mixed with Harpalejeunea parasitica in

rock crevices in dwarf shrub heaths. The Ochyra

plant was found on dry quartzite rocks associated

with Hymenophyllum species. Taylor collected the

species at a "joint plane in north-dipping quartz-

ose sandstones, on south facing outcrop" (Taylor

label).

PH YTOGEOGRAPHY — Magellanian-Falkland —

Falkland Is.; Tierra del Fuego; Patagonian Chan-

nels (51°19'S). Within the Falklands, it was found

only in the Port Stanley region, and the cooler

temperatures and higher rainfall in this region may

account for its presence there.

FALKLAND SPECIMENS SEEN -EAST FALKLANDS.

Stanley Region: Sapper Hill, 2 km W of Port Stanley,

ca. 100 m, Ochyra 2883/80 (F); Mt. William, rock pin-

nacles, 185 m, Taylor 226a (AAS, F); summit ridge of N

peak of Two Sisters, 245-290 m (2726B); summit of Mt.

Kent, 455 m (2769Q.

Lophozia

Lophozia (Dum.) Dum., Recueil Observ. Jun-

germ. 17. 1835. Jungermannia sect. Lophozia

Dum., Syll. Jungerm. Eur. Indig. 53. 1831.

Lophozia hatched (Evans) Steph.15

Jungermannia hatcheri Evans, Bull. Torrey Bot. Club

25(8): 417. pi 346, f. 1-7. 1898. Lophozia hatch-

eri (Evans) Steph., Result. Voyage S. Y. Belgica

6(5): 4. 1901. Barbilophozia hatcheri (Evans)

Loeske, Verb. Bot. Vereins Prov. Brandenburg

49: 37. 1907. Original material: Argentina, Terr.

Tierra del Fuego, Lapataia, J. B. Hatcher 35b

(NY!).

ECOLOGY— Rather rare in the Falklands, where

I found it in or at the margin of feldmark pools

(700 m) as well as in a dwarf shrub heath on rock

in a cool, protected niche under rock cover (180

m).

PHYTOGEOGRAPHY— Bipolar in Distribution— In

the Northern Hemisphere, holoarctic and high

subarctic-subalpine to arctic-alpine. In the South-

ern Hemisphere, antarctic in distribution occur-

ring in the Antarctic Peninsula, South Shetland

Is., South Orkney Is. (Signey I.) (for a list of sta-

15 See Schuster 1969b for a full synonymy of L. hatch-

lions on Antarctica and offshore islands, where the

species seems to be frequent, see Ochyra & Vana,

1989), South Georgia, Falkland Is. (above 245 m),

the mountainous region of Isla Grande de Tierra

del Fuego, and the Andes of southern Patagonia.

In the latter two regions I suspect it is either sub-

alpine or alpine.

LITERATURE RECORDS (FALKLANDS)— Anony-

mous (Fulford, 1963c as Barbilophozia; Kiihne-

mann, 1937); Skottsberg & Halle, Mt. Adam

(Grolle, 1960a as Barbilophozia; Stephani, 1911).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Mt. Usborne Region: summit of Mt. Usborne 1, ca. 700

m (2489C, 2501D). WEST FALKLANDS. Fox Bay Re-

gion: summit of East Head, 180 m (3431).

Nothostrepta

Nothostrepa Schust., Phytologia 45: 420. 1980.

Nothostrepta hifida (Steph.) Schust. Figure 4 1 .

Plagiochila bifida Steph., Annuario Reale 1st. Bot.

Roma 2(2): 86. pi. 6,f. 1-6. \886.Anastrophyllum

bifidum (Steph.) Steph., Bih. Kongl. Svenska Ve-

tenskapsakad. Handl. 26 (III, 6): 25. 1900. An-

astrepta bifida (Steph.) Steph., Bull. Herb. Bois-

sier 2(5): 474. 1902 (= Spec. Hep. 2: 193).

Nothostrepta bifida (Steph.) Schust., Phytologia

45: 420. 1980. Original material: Chile, Prov.

Magallanes, Pto. Caracciolo, De Amezaga (non

vidi).

Tylimanthus bilobatus Steph., Kongl. Svenska Ve-

tenskapsakad. Handl. 46(9): 24.f.9b. 1911, syn.

fide Grolle (1961). Lectotype (cf. Grolle, 1961):

Juan Fernandez Is., Mas a Tierra, Valle Colonial,

1908, Skottsberg (VPS, non vidi).

Leioscyphus fernandeziensis Steph., Kongl. Svenska

Vetenskapsakad. Handl. 46(9): 36. / 14a. 1911,

syn. fide Grolle (1961). Lectotype (fide Grolle,

1961): Juan Fernandez Is., Mas Afuera, Camp

Correspondencia, 1908, Skottsberg (UPS, non vidi).

REMARKS— See Schuster (1981b).

ECOLOGY— Collected at but a single station in

the Falklands, on soil under a thick protective grass

cover in a Cortaderia heath.

PHYTOGEOGRAPHY— Valdivian + Magellanian-

Falkland— Falkland Is.; Patagonian Channels;

Valdivian region (north to 39°52'S); and Juan Fer-

nandez Is. Not reported for Tierra del Fuego or

Andean Patagonia (see fig. 41).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Mt. Usborne Region: below The Gap, ca. 90 m (2561-

c.S,2572B, 2573A-C.S).

96

FIELDIANA: BOTANY

Roivainenia

Roivainenia Perss. in Persson & Grolle, Nova

Hedwigia 3: 43. 1961.

Roivainenia jacquinotii (Mont.) Grolle. Figure 42.

Jungermannia jacquinotii Mont., Ann. Sci. Nat. Bot.

II, 19: 250. 1843. Chiloscyphus jacquinotii (Mont.)

Nees in G. L. & N., Syn. Hep. 185. 1845. Roi-

vainenia jacquinotii (Mont.) Grolle in Perss. &

Grolle, Nova Hedwigia 3: 44. 1961. Lectotype

(cf. Persson & Grolle, 1961): Chile, Prov. Ma-

gallanes, Strait of Magellan, Jacquinot (PC!).

Jungermannia antarctica Angstr., Ofvers. Forh. Kongl.

Svenska Vetenskapsakad. 29(4): 10. 1872, syn.

fide Persson & Grolle (1961). Lophozia antarctica

(Angstr.) Evans, Bull. Torrey Bot. Club 25(8): 416.

1898. Roivainenia antarctica (Angstr.) Perss. in

S. Arnell, Svensk Bot. Tidskr. 49: 239. 1955, nom.

illeg. Original material: Chile, Prov. Magallanes,

Pto. del Hambre, 1852, Andersson (non vidi).

Jungermannia pigafettoana Mass., Nuovo Giorn. Bot.

Ital. 17: 217. pi 14, f. 7. 1885, syn. cf. Stephani

(1901). Lophozia pigafettoana (Mass.) Kiihnem.,

Lilloa 19: 344. 1949. Original material: Argen-

tina, Terr. Tierra del Fuego, Ushuaia, Spegazzini

(non vidi).

Jungermannia verrucosa Steph., Hedwigia 34: 51.

1895, syn. cf. Stephani (1901). Original material:

Chile, Prov. Magallanes, Pto. Eden, Cunningham

(non vidi).

Leioscyphus skottsbergii Steph., Wiss. Ergebn. Schwed.

Siidpolarexped.4(l): S.f.6,7. 1 905, syn. fide Pers-

son & Grolle (1961). Mylia skottsbergii (Steph.)

Schust., Amer. Midi. Naturalist 62: 34. 1959.

Lectotype (cf. Persson & Grolle, 1961): South

Georgia, Skottsberg (G, non vidi).

Anastrophyllum verrucosum Steph., Kongl. Svenska

Vetenskapsakad. Handl. 46(9): 21. f. 7c. 1911,

syn. fide Persson & Grolle (1961). Lectotype (cf.

Persson & Grolle, 1961): Southern Patagonia,

Skottsberg (G, non vidi).

Acrobolbus patagonicus Steph., Kongl. Svenska Ve-

tenskapsakad. Handl. 46(9): 23.f.7d. 1911, syn.

fide Persson & Grolle (1961). Lectotype (cf. Grolle

& Persson, 1961): Chile, Prov. Aisen, Coihaique,

Halle (G, non vidi).

Leioscyphus bilobatus Steph., Kongl. Svenska Ve-

tenskapsakad. Handl. 46(9): 35. / 13a,b. 1911,

syn. fide Persson & Grolle (1961). Leptoscyphus

bilobatus (Steph.) Kiihnem., Revista Centro Es-

tud. Doct. Ci. Nat. 1: 175. 1937. Mylia bilobata

(Steph.) Kiihnem., Lilloa 19: 340. 1949. Original

material: Falkland Is., near Port Stanley, Skotts-

berg (UPS), cited in Persson & Grolle (1961) (non

vidi).

ECOLOGY— In the Falklands, in dwarf shrub

heaths under protective cover of Empetrum,

Blechnum magellanicum, and B. penna- marina.

Occasionally on soil or rock of sheltered high al-

titude cliffs. The species is very common in the

41

FIG. 4 1 . Distribution of Nothostrepta bifida (Steph.)

Schust.

deciduous Nothofagus forests of southern South

America.

PHYTOGEOGRAPHY— Valdivian + Magellanian

+ Falklands— South Shetland Is. (Allison and

Smith, 1973; Smith, 1979); South Georgia; Falk-

land Is.; Tierra del Fuego; the Patagonian Chan-

nels; Valdivian region (West Patagonia north to

39°52'S and in Andean Patagonia at P. N. Nahuel

Huapi) (see fig. 42).

LITERATURE RECORDS (FALKLANDS): Anony-

mous (Kiihnemann, 1937 as Leptoscyphus bilo-

batus, 1 949 as Mylia bilobata); Skottsberg (Grolle,

1962; Persson & Grolle, 1961; Skottsberg, 1913;

Stephani, 1922, as Leioscyphus bilobatus).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Port William Region: summit ridge of Mt. Low, ca. 245

m (3229— c. $); N side of Gypsy Cove, sea level (3248,

3256, 3261). Stanley Region: near Radio Station, Stan-

ley, 30 m, Longton & Smith, Longton 1049 (AAS, F);

Sapper Hill, 2 km W of Port Stanley, ca. 40 m, Ochyra

2829/80 (F); Sapper Hill, 2.5 km W of Port Stanley, 60

m, Ochyra 2850/80 (F); Sapper Hill, 135m (2405, 2416);

Tumbledown Mt., 155-230 m (2389); near summit of

Tumbledown Mt., 230 m, Longton & Smith, Longton

971 (AAS, F); summit ridge of N peak of Two Sisters,

245-290 m (2707). Mt. Usborne Region: ridge between

Mts. Usborne 1 and 2, 685 m (2524, 2554Q. WEST

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

97

FIG. 42. Distribution of Roivainenia jacquinotii (Mont.) Grolle, plus South Georgia.

FALKLANDS. Port Howard: pass SW of Mt. Maria

summit, ca. 610 m (3084).

Family JUNGERMANNIACEAE

Jungermanniaceae Reichenb., Botanik fur Da-

men, Kiinstler und Freunde. . . 256. 1828.

The Jungermanniaceae, with 18 genera, are

worldwide in distribution. Genera occurring in the

Falklands are among the largest in the family—

Jungermannia has ca. 1 20 species and Jameson-

iella has 1 3 species. The only predominantly south

temperate genus in the family is Cryptochila (6

species; 5 are south temperate-subantarctic, 1 is

in New Guinea).

Cryptochila

Cryptochila Schust., J. Hattori Bot. Lab. 26: 284.

1963.

Key to Falkland Islands Species of Cryptochila

1 . Stem tips and leaves (of mostly robust plants) curved ventrally; dorsal stem surface usually with

paraphyllia between the leaves (paraphyllia occasionally absent); perianth plicae strongly twisted;

leaves orbicular or reniform. Dorsal margin of leaf (of most robust plants) long decurrent

C. grandiflora

98

FIELDIANA: BOTANY

1. Stem tips and leaves plane or ± curved dorsally; stems without paraphyllia; perianth plicae not

twisted; leaves 1.3-1.5(1.6) times as long as wide, spathulate or ligulate. Base of dorsal margin of

leaf (mostly of robust plants) conspicuously rigid, with a small bistratose swelling ... C. paludosa

Cryptochila grandiflora (Lindenb. & Gott.) Grolle.

Figure 43.

Only American synonyms are included here:

Jungermannia grandiflora Lindenb. & Gott. in G. L.

& N., Syn. Hep. 673. 1847. Jamesoniella gran-

diflora (Lindenb. & Gott.) Jack & Steph., Hed-

wigia 31: 13. 1892. Cryptochila grandiflora (Lin-

denb. & Gott.) Grolle, Feddes Repert. 82(1): 19.

1971. Lectotype (cf. Grolle, 197 la): Chile, Prov.

Valdivia, Valdivia, "mis. Montagne 1845 als J.

colorata; W (Lindenb. Hep. no. 1 807)," sin. coll.

(Gay) (non vidi).

Jungermannia penicillata Loitl. in Szyszylowicz,

Diagn. pi. nov. a C. Jelski in Peruvia lect., P. 1 .

Acad. Litt. Cracov 238. 1894, syn. fide Grolle

(197 la). Original material: Peru, without specific

locality, Jelski 549 (w), cited in Grolle (197 la)

(non vidi).

Jamesoniella allionii Steph. in Herzog, Biblioth. Bot.

87: 182. 1916, syn. fide Grolle ( 197 la). Lectotype

(fide Grolle, 197 la): Bolivia, above Tablas, 3400

m, Herzog 2847 (JE, non vidi).

Jamesoniella pyrogea Mass., Atti Reale 1st. Veneto

Sci. 87: 235. pi. 3, f. 1-4. 1927, syn. fide Grolle

(197 la). Original material: Chile, Prov. Magal-

lanes, I. Basket and B. Sarmiento, Spegazzini (non

vidi).

Jamesoniella pellucida Herz., Hedwigia 74: 85. 1934,

syn. fide Grolle (197 la). Lectotype (fide Grolle,

197 la): Bolivia, "Cejagiirtel von Stillutincara,

Jungas von La Paz," Troll 132 (JE, non vidi).

ECOLOGY— Rather characteristic of the shel-

tered high altitude cliffs, where it occurs on Azo-

rella cushions, under ledges, or occasionally in a

rather exposed rock crevice. It is part of a com-

bination of hepatics frequently associated in this

habitat, i.e., Austrolophoziafuegiensis, Balantiop-

sis bisbifida, Cryptochila paludosa, Herzogobryum

teres, Pachyglossa spp., and Triandrophyllum sub-

trifidum. Commonly mixed with other Hepaticae.

PHYTOGEOGRAPH Y — Pan-south-temperate —

South Sandwich Is.; South Georgia; Falkland Is.

(455-700 m); Tierra del Fuego; Andes north to

Guatemala; Africa from Cape of Good Hope to

Natal (3153 m); Marion I.; Prince Edward Is.;

Crozet Is.; Macquarie I.; northeast New Guinea

(4400-4600 m) (see fig. 43). This taxon is yet

another example of a strictly south temperate ele-

ment occurring on Serra do Itatiaia, Brazil (see

Engel, 1968, p. 119).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Mt. Usborne Region: SE slope of Mt. Usborne 2, ca. 455

m (2605); ridge between Mts. Usborne 1 & 2, 685 m

(2548, see Grolle, 197 la; 2555/4). WEST FALK-

LANDS. Port Howard: pass SW of Mt. Maria summit,

ca. 610 m (3093). Mt. Adam: ridge S of northern lake,

610 m (3042C); E side of summit ridge, 670-700 m

(30I4E); summit of southernmost peak, 685 m (2988).

Cryptochila paludosa (Steph.) Grolle. Figure 44.

Jamesoniella paludosa Steph., Bih. Kongl. Svenska

Vetenskapsakad. Handl. 26 (III, 17): 11. 1901.

Bull. Herb. Boissier 1(10): 1034. 30 Sept. 1901

(= Spec. Hep. 2: 97). Cryptochila paludosa (Steph.)

Grolle, Feddes Repert. 82( 1 ): 26. 1971. Lectotype

(cf. Grolle, 197 la): Chile, Prov. Magallanes, I.

Desolation, Pto. Angosto, Dusen (G, non vidi).

ECOLOGY— In the Falklands, this species seems

to be restricted to sheltered high altitude cliffs,

where it may be found in crevices, under ledges,

etc. Like Cryptochila grandiflora, it is part of a

combination of hepatics frequently found associ-

ated in this habitat, i.e., Austrolophoziafuegiensis,

Balantiopsis bisbifida, C. grandiflora, Herzogo-

bryum teres, Pachyglossa spp., and Triandrophyl-

lum subtrifidum, and often occurs mixed with these

Hepaticae.

PHYTOGEOGRAPHY— Valdivian + Magellanian

+ Falklands -Falkland Is. (610-685 m); Tierra

del Fuego; Patagonian Channels; Valdivian region

north to 39°52'S; Tristan da Cunha (see fig. 44).

FALKLAND SPECIMENS SEEN -EAST FALKLANDS.

Mt. Usborne Region: ridge between Mts. Usborne 1 &

2, 685 m (2554F). WEST FALKLANDS. Port Howard:

passSW of Mt. Maria summit, ca. 610 m (3097C, 3120,

3129B, see Grolle, 1 97 1 a). Mt. Adam: ridge S of northern

lake, 610 m (3030B, 3037); summit of southernmost

peak, 685 m (2986Q.

Jamesoniella

Jamesoniella (Spruce) ("airing, in Lees, London

Cat. Brit. Moss. Hep. 25. 1881. Jungerman-

nia subg. Jamesoniella Spruce, J. Bot. 1 4: 202,

230. 1876.

Jamesoniella colorata (Lehm.) Schiffn. Figure 45.

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

99

100

FIELDIANA: BOTANY

44

45

FIG. 44. Distribution of Cryp-

tochila paludosa (Steph.) Grolle,

plus Tristan de Cunha. Open cir-

cle indicates precise locality un-

known.

FIG. 45. Distribution of Ja-

mesoniella colorata (Lehm.)

Schiffn. (after Grolle, 1971a).

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

101

Jungermannia colorata Lehm., Linnaea 4: 366. 1829.

Jamesoniella colorata (Lehm.) Schiffn. in Engler

& Prantl, Natiirl. Pflanzenfam. 1(3): 83. 1893.

Lectotype(/Kfe Grolle, 197 la): South Africa, Cape

Prov., Table Mt., Ecklon (s, non vidi).

Jungermannia malouina Gott., Ann. Sci. Nat. Bot.

IV, 8: 337. 1857, syn. fide Grolle (197 la). Ja-

mesoniella malouina (Gott.) Steph., Bull. Herb.

Boissier 1(10): 1027. 1901 (= Spec. Hep. 2: 90)

("maluina"). Lectotype (cf. Grolle, 197 la): Falk-

land Is., Lesson16 (PC, non vidi).

For complete synonymy see Engel (1978); only

Falkland Islands species are included here.

ECOLOGY— Quite common in dwarf shrub and

Cortaderia heaths, where it grows on soil or more

often in crevices, etc., of rock outcrops. It was not

found below 60 m.

PH YTOGEOGRAPHY — Pa n-so u t li-U> m p e r at i- in

Distribution— The species has not penetrated north

beyond temperate regions in any of the sectors. It

has, however, penetrated into the subantarctic re-

gion, as it occurs on Marion and Prince Edward

Islands, Crozet Is., Kerguelen Is., and Macquarie

I. Grolle (197 la) stated the records from Neo-

tropical regions are erroneous and are mostly spec-

imens of J. rubricaulis. Grolle also states the report

from South Georgia by Stephani (1911) is based

upon a specimen which is actually Cryptochila

grandiflora (see fig. 45).

LITERATURE RECORDS (FALKLANDS)— Anony-

mous (Bonner, 1 966, as /. maluina; Kiihnemann,

1937, 1949, as J. colorata & J. maluina; Spruce,

1 876, as Jungermannia); Hooker (G. L. & N. 1 847,

as Jungermannia; Hooker & Taylor, 1844, and

Taylor & Hooker, 1847, as Jungermannia; Spruce,

1876, as Jungermannia); Lesson (Stephani, 1901,

as J. maluina); Skottsberg, Port Stanley (Grolle,

197 la; Skottsberg, 1913; Stephani, 1905a, as /.

colorata & J. maluina), Port Louis (Stephani,

1905a), Port Louis, Mt. Adam, Port William

(Skottsberg, 1913), Hornby Mts. and between Ar-

row Harbour House and Victoria Creek-Lafonia

(Skottsberg, 1913, as J. maluina); Skottsberg &

Halle, Port Stanley, Mt. Adam, Roy Cove, Wed-

dell Is. (Stephani, 1911), Hornby Mts. (Stephani,

1 9 1 1 , as J. maluina).

FALKLAND SPECIMENS SEEN -EAST FALKLANDS.

Port William Region: Charles Point, 1954, Pettingill 3

(CINC, F); Sparrow Cove, 1954, Pettingill 45 (cmc, F).

"Gottsche (1857) states the original collection was

made in 1825. However, Moore (1968) stated Lesson,

who was physician and assistant to Dumont d'Urville,

was in the Falklands from 20 November- 18 December

1822.

Stanley Region: Sapper Hill, 2 km W of Port Stanley,

ca. 150 m, Ochyra 2872/80 (F); Sapper Hill, 2.5 km W

of Port Stanley, ca. 100 m, Ochyra 2907/80 (F); Sapper

Hill, 135 m (2422); Mt. William, rock pinnacles, 185 m,

Taylor 224 (AAS, F); between Mt. William and Tumble-

down Mt., 170 m, Longton & Smith. Longton 978 (AAS,

F); Tumbledown Mt., 155-230 m (2374, 23828-c. per.);

Wireless Ridge, near Stanley, 60 m, Longton & Smith,

Longton 1016 (AAS, F); N peak of Two Sisters, 275 m,

Dollman s.n. (Greene 3558a) (AAS); summit of Mt. Kent,

455m (2760/1). Mt. Usborne Region: ridge between Mts.

Usborne 1 & 2, 685 m (2553C-C. per.); The Gap, 275-

290 m (2464); below The Gap, ca. 90 m (2555, 2555).

WEST FALKLANDS. Fox Bay Region: valley E of Su-

livan House, ca. 75 m (3530, 3533). Hill Cove Region:

summit of West French Peak, 290 m (2959); valley near

Hill Cove House, Valient in s.n. (MANCH). Roy Cove Re-

gion: Roy Cove, Vallentin s.n. (MANCH). Weddell Island:

summit of Mt. Weddell, 380 m (3341).

Jungermannia

Jungermannia L. Spec. PI. ed. 1. 1131. 1753.

Jungermannia crassula Nees & Mont.

Jungermannia crassula Nees & Mont., Ann. Sci. Nat.

Bot. II, 5: 54. 1836. Solenostoma crassula (Nees

& Mont.) Steph., Bull. Herb. Boissier 1(5): 497.

1 90 1 (= Spec. Hep. 2: 59). Original material: Juan

Fernandez Is., Bertero (G, PC, s, w), cited in Vana

( 1974) (non vidi).

Jungermannia domeikoana Mont., Ann. Sci. Nat. Bot.

Ill, 4: 349. 1845, syn. fide Stephani (1901). Orig-

inal material: southern Chile, without specific lo-

cality, Gay (BM, FH, G, PC, s, w), cited in Vana

(1974) (non vidi).

Solenostoma obtusiflorum Steph., Kongl. Svenska Ve-

tenskapsakad. Handl. 46(9): I6.f.5a. 1911, syn.

fide Vana (1974). Original material: Juan Fer-

nandez Is., Mas Afuera, Skottsberg (G, L, s), cited

in Vana (1974) (non vidi).

Solenostoma rostratum Steph., Kongl. Svenska Ve-

tenskapsakad. Handl. 46(9): 16./ 5b. 1911, syn.

fide Vana (1974). Original material: Juan Fer-

nandez Is., Mas Afuera, Skottsberg (G, L, s), cited

in Vana (1974) (non vidi).

Jungermannia simplicissima Steph., Kongl. Svenska

Vetenskapsakad. Handl. 46(9): 17./ 5c,d. 1911,

syn. fide Vana (1974). Original material: Chile,

Prov. Chiloe, I. Chiloe, Pto. Quellon, Halle &

Skottsberg (G, s), cited in Vana (1974) (non vidi).

Jamesoniella nana Steph., Spec. Hep. 6: 101. 1917,

syn. fide Grolle (197 la). Original material: Falk-

land Is., Halfway Cove, 1907, Halle (G, hb. Grolle),

cited in Vana ( 1 974), (UPS), cited in Grolle (1971)

and Vana (1974) (non vidi).

REMARKS— Stephani (1911) used the name S.

obtusiflorum, but altered the spelling to "obtusi-

102

F1ELDIANA: BOTANY

flos" in Species Hepaticarum (Stephani, 1917, p.

82).

ECOLOGY— Collected at a single locality in the

Falklands, on a stream bank of a dwarf shrub heath.

PHYTOGEOGRAPHY — Valdivian-Falkland —

Tristan da Cunha, Inaccessible I., Gough I.; Falk-

land Is.; Valdivian region (West Patagonia north

to 36°50'S); Juan Fernandez Is. (Mas Afuera, Mas

a Tierra). (Within the Falklands it was found only

in the Port Stanley region, and the cooler temper-

atures and higher rainfall may account for its pres-

ence there.)

LITERATURE RECORDS (FALKLANDS)— Dusen11,

Halfway Cove (Bonner, 1966, as Jamesoniella

nana\ Vana, 1974).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Stanley Region: headwaters of Mullet Creek Stream, ca.

60 m (3161A, 3172).

Notes on Jungermannia Species

Jungermannia stereocauli Bory in Gaudichaud,

Botanique, Pt. 4, p. 1 30. In Freycinet, Voyage

autour du monde . . . sur les corvettes de S.

M. rUranieel la Physicienne. 1827 (1826).

This species was described in Gaudichaud (1827)

for a Falkland Islands specimen Gaudichaud col-

lected. I have not been successful in numerous

attempts, over a period of many years, to locate

this specimen. It could not be found at PC, or at

BM, BR, DS, E, FI, G, GL, H, L, Or W. It is quite

impossible to ascertain the identity of the species

from the original one line diagnosis: "Foliis ovato-

concavis, integerrimis, imbricatis." The species

name refers to the lichen Stereocaulon, and the

only member of that genus mentioned in the Gau-

dichaud Falkland collection is S. turfosum Bory.

It was thought that perhaps the lichen might still

have the J. stereocauli on the thallus, and that a

separate collection of the hepatic had not been

made. However, Lamb (1977, p. 326) stated that

neither Bory's type, nor any other authentic Ste-

reocaulon specimen from the Falklands, is known.

The following comments by Stafleu and Cowan

17 Stephani cited Dusen as the collector of the speci-

men; this, however, is incorrect, as Dusen did not visit

the Falklands. Bonner ( 1 966) stated for the type of this

species, "West Falkland, Halfway Cove, s.d., Dus£n s.n.,

in hb. G." Halfway Cove was visited by Halle and Skotts-

berg, and Stephani determined their hepatic collections.

In this case Halle made the collection.

(1976, p. 921) are relevant: "Gaudichaud gave all

his collection to P [and PC]. The collections made

on the voyage in 1' Vranie and la Physicienne (1817-

1820) went directly to P. Part of the collections

were lost during the voyage when the Vranie

stranded in the Falkland Islands (lies Malouines)."

Therefore, the possibility exists that a Gaudichaud

description of J. stereocauli was saved but the

specimen was lost. See also Sayre (1975, p. 326).

Family GYMNOMITRIACEAE

Gymnomitriaceae Klinggr., Die hoheren Crypto-

gamen Preussens 16. 1858.

There are nine genera in the Gymnomitriaceae;

all are concentrated in arctic or arctic-alpine re-

gions. Herzogobryum, however, is strictly austral,

and members of that genus are typical of the alpine

flora. Acrolophozia is also strictly austral: A. fue-

giana occurs in the American sector, A. sulcata

Hassel is endemic to South Georgia, and A. pec-

tinata Schust. is found in New Zealand.

Acrolophozia

Acrolophozia Schust., Rev. Bryol. Lichenol. 34:

259. 1966; J. Hattori Bot. Lab. 26: 216, 279.

1963, nom. inval. sin. descr. lat.

Acrolophozia fuegiana Schust.

Acrolophozia fuegiana Schust., Nova Hedwigia 15:

494, pi 63. 1968. Holotype: Argentina, Terr.

Tierra del Fuego, C. Garibaldi, ca. 900 m, Schus-

ter 58317 (non vidi).

ECOLOGY- PHYTOGEOGRAPHY— This subantarc-

tic rare species was found at a single station in the

sheltered high altitude cliff vegetation growing with

Clasmatocoleafulvella and Herzogobryum teres on

rock under a rock overhang. The Falkland ecology

of A. fuegiana appears to be quite similar to that

of the type, which was gathered on Cerro Gari-

baldi, Tierra del Fuego, "ca. 900 m, on damp ledges

in alpine tundra . . . associated with Pachyglossa

dissitifolia, Clasmatocolea sp. (a species with poly-

stratose leaves) [= Evansianthus georgiensis], and

Herzogobryum teres" (Schuster 1 968b). All the as-

sociates listed above are found in the sheltered

high altitude cliff vegetation in the Falklands. The

species, which is also known from South Georgia

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

103

(Hassel de Menendez, 1 980a), is very closely allied

to A. pectinata Schust. of South Island, New Zea-

land. It is significant to note that the New Zealand

species is known from ca. 1370-1460 m, on soil

over shaded ledges in the snow tussock (alpine)

zone.

FALKLAND SPECIMEN SEEN- EAST FALKLANDS. Mt.

Usborne Region: ridge between Mts. Usborne 1 & 2, 685

m (2521C-C. per.).

Herzogobryum

Herzogobryum Grolle, Rev. Bryol. Lichenol. 32:

160. 1964 (1963).

Chondrophyllum Herz., Rev. Bryol. Lichenol. 21: 46.

1952, non Chondrophyllum Kylin, Acta Univ.

Lund. n. s. 20(6): 442. 1924 (Rhodophyta).

Key to Falkland Islands Species of Herzogobryum

1 . Cells of leaf margins with considerably thicker walls than median lamina cells; plants monoecious;

antheridia paired; perianth not exserted beyond bracts H. erosum

1 . Cells of leaf margins with thinner walls than median lamina cells; plants dioecious; antheridia single;

perianth exserted far beyond bracts 2

2. Leaves cucullate; leaf margins plane to weakly crenulate; leaf apex not fissured

H. vermiculare

2. Leaves concave; leaf margins strongly crenulate; leaf apex often fissured H. teres

Herzogobryum erosum (Caning. & Pears.) Grolle.

Fig. 46.

Cesia erosa Carring. & Pears., Pap. & Proc. Roy. Soc.

Tasmania 1887: 8. pi. 6, f. 1-19. 1888. Gymno-

mitrium erosum (Carring. & Pears.) Bast., Pap. &

Proc. Roy. Soc. Tasmania 1887: 244. 1888 ("ero-

sa"). Herzogobryum erosum (Carring. & Pears.)

Grolle, Oesterr. Bot. Z. 1 13: 231. 1966. Original

material: Tasmania, Bastow s.n. (BM), cited in

Grolle (1966a)(m?H vidi).

Cesia stygia var. denticulata Berggr., New Zealand

Hep. 4. 1898, syn. fide Grolle (1966a). Acolea

denticulata (Berggr.) Steph., Bull. Herb. Boissier

1(2): 143. 1901 (= Spec. Hep. 2: 4). Gymnomit-

rium denticulatum (Berggr.) K. Mull. (Freib.) Rev.

Bryol. Lichenol. 20: 176-177. 1951. Lectotype

(/KteGrolle, 1 966a): New Zealand, South I., West-

land, Kelly's Hill, 1200 m, 1874, Berggren 2870

(LD, non vidi).

ECOLOGY— Collected only once in the Falk-

lands, on sandy soil under a rock ledge of sheltered

high altitude cliffs.

PHYTOGEOGRAPHY— Subantarctic in Distribu-

tion—South Georgia; Falkland Is.; Tierra del Fue-

go, Isla de los Estados; southern Patagonian Chan-

nels, Brunswick Peninsula; Tristan da Cunha; 500-

2000 m in the New Zealand sector, occurring on

Campbell I., Stewart I., and north to Tasmania

(see fig. 46).

FALKLAND SPECIMEN SEEN -WEST FALKLANDS.

Mt. Adam: summit of southernmost peak, 685 m

(2986A—C. per. + $).

Herzogobryum teres (Carring. & Pears.) Grolle.

Figure 47.

Jungermannia teres Carring. & Pears., Pap. & Proc.

Roy. Soc. Tasmania 1887: 9. pi. 7,f. 1-16. 1888

("Jungermania"). Jamesoniella teres (Carring. &

Pears.) Steph., Bull. Herb. Boissier 1(10): 1037.

1901 (= Spec. Hep. 2: 100). Herzogobryum teres

(Carring. & Pears.) Grolle, Oesterr. Bot. Z. 113:

223. 1966. Lectotype (cf. Grolle, 1966a): Tas-

mania, Mt. Wellington, Bastow s.n. (MANCH, non

vidi).

Acolea magellanica Mass. & Steph. in Massalongo,

Atti Reale Inst. Veneto Sci. 87: 237. pi. 3, f. 5-

7. 1927, syn. fide Grolle (1966a). Original ma-

terial: Argentina, Terr. Tierra del Fuego, I. de los

Estados, Penguin Rookery, Spegazzini s.n. (non

vidi).

Jamesoniella inflexo-limbata Herz., Trans. & Proc.

104

FIELDIANA: BOTANY

46

FIG. 46. Distribution of Herzogobryum erosum (Carring. & Pears.) Grolle.

FIG. 47. Distribution of Herzogobryum teres (Carring. & Pears.) Grolle.

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

105

Roy. Soc. New Zealand 68: 41. 1938, syn. fide

Grolle (1966a). Lectotype (cf. Grolle, 1966a): New

Zealand, North I., Ml. Tongariro, 12 Jan. 1933,

Moore 130 (JE, non vidi).

ECOLOGY— In the Falklands, this taxon seems

restricted to sheltered high altitude cliffs, where it

occurs very commonly with Austrolophozia fue-

giensis, Balantiopsis bisbifida, Cryptochila spp.,

Pachyglossa spp., and Triandrophyllum subtrifi-

dum. It grows in moist, protected rock crevices,

frequently under rock overhangs. Schuster ( 1 968b,

p. 499) included this species as an associate of

Acrolophoziafuegiana, which was collected at Cer-

ro Garibaldi, Tierra del Fuego at "ca. 900 m, on

damp ledges in alpine tundra" (p. 499), an eco-

logical situation which apparently is quite similar

to the sheltered high altitude cliffs in the Falklands.

PHYTOGEOGRAPHY— Antarctic in Distribu-

tion—South Shetland Is. (Ochyra and Vana, 1989);

South Orkney Is. (Smith, 1972); Falkland Is.; Ti-

erra del Fuego; Tristan da Cunha; Gough I.;

Campbell I.; 1200-2300 m in New Zealand and

Tasmania; Marion I. (montane) (fig. 47).

FALKLAND SPECIMENS SEEN- EAST FALKLANDS.

Mt. Usborne Region: ridge between Mts. Usborne 1 &

2, 685 m (2575-c. $, 2517 A, 2521A-C. 6, 2527, 2528-

c. per. + 3, 2530, 2554G). WEST FALKLANDS. Port

Howard: pass SW of Mt. Maria summit, ca. 610 m

(3087-c. per., 3098-c. $, 3112B, 3315B-C. <5, 3127 'A-

c. $, 3129C). Mt. Adam: ridge S of northern lake, 610

m (30 30 A -c. per. + <3, 3036B).

Herzogobryum vermiculare (Schiffn.) Grolle. Fig-

ure 48.

Gymnomitrium vermiculare Schiffn. in Naumann,

Forschungsr. Gazelle 4(4): 2. pi. l,f. 9,10. 1890.

Herzogobryum vermiculare (Schiffn.) Grolle, J.

Hattori Bot. Lab. 28: 103. 1965. Original mate-

rial: Kerguelen Is., Successful Harbor, 1 874, Nau-

mann s.n. (w), cited in Grolle (1965d) (non vidi).

Chondrophyllum cucullatum Herz., Rev. Bryol. Li-

chenol. 21: 46. / 1-2. 1952, syn. fide Grolle

( 1 965d). Herzogobryum cucullatum (Herz.) Grolle,

Rev. Bryol. Lichenol. 32: 160. 1963. Original ma-

terial: Chile, Prov. Aisen, C. Tesoro, ca. 1000 m,

1 3 Feb. 1 940, Schwabe 39cp.p. (JE), cited in Grolle

(I965d)(non vidi).

ECOLOGY— Found in a moist feldmark (700 m),

on Azorella cushions, soil over rock, and in rock

crevices of sheltered high altitude cliffs. It is very

frequently mixed with other hepatics, particularly

with Cryptochila spp., Pachyglossa spp., and Her-

zogobryum teres.

PHYTOGEOGRAPHY— Subantarctic in Distribu-

tion—South Georgia; Falkland Is.; southern South

America only at ca. 1 000 m in the Valdi vian region

(44°19'S); Kerguelen Is.; Marion I. (see fig. 48).

LITERATURE RECORD (FALKLANDS)— Anony-

mous (Grolle, 1 969).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Mt. Usborne Region: ridge between Mts. Usborne 1 &

2, 685 m (25555); summit of Mt. Usborne 1, ca. 700 m

(2505D). WEST FALKLANDS. Port Howard: pass SW

of Mt. Maria summit, ca. 6 10 m (3097 B-c. per., 3110A,

3111.3129A). Mt. Adam: summit of southernmost peak,

685 m (2986B).

Family SCAPANIACEAE

Scapaniaceae Migula, Krypt.-Fl. Deutschl. 1: 479.

1904.

Two genera of this basically Laurasian family

occur regionally— Diplophyllum and Scapania.

Scapania is the largest genus, with ca. 58 species.

Scapania gamundiae Schust., of southern South

America, is the only austral member of the genus

and also stands alone in having subequally bilobed

leaves.

Diplophyllum

Diplophyllum (Dum.) Dum., Recueil Observ. Jun-

germ. 15.1835, nom. cons. Jungermannia sect.

Diplophyllum Dum., Syll. Jungerm. Eur. In-

dig. 44. 1831.

Diplophyllum acutilobum Steph.

Diplophyllum acutilobum Steph., Kongl. Svenska Ve-

tenskapsakad. Handl. 46(9): 83. / 32f. 1911.

Original material: Chile, Prov. Maga llanos. W end

of L. Fagnano, Halle & Skottsberg s.n. (s), cited

in Grolle (1965b) (non vidi).

REMARKS— See comments in Schuster (1974, p.

193).

ECOLOGY— Rare; collected at a single locality,

on soil under a rock overhang of sheltered high

altitude cliffs.

PHYTOGEOGRAPHY— Valdi vian + Magellanian

+ Falklands— Falkland Is.; Tierra del Fuego, Lago

Fagnano; Patagonian Channels; the Valdivian re-

gion north to 40°42'S.

106

FIELDIANA: BOTANY

48

FIG. 48. Distribution of Herzogobryum vermiculare (Schiffn.) Grolle.

FIG. 49. Distribution Blepharidophyllum clandestinum (Mont.) Lac.

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

107

FALKLAND SPECIMEN SEEN -WEST FALKLANDS.

Mt. Adam: ridge S of northern lake, 610 m (3027— c.

per. + i).

Family BLEPHARIDOPHYLLACEAE

Blepharidophyllaceae (Schust.) Schust. ex Engel,

comb. nov. Scapaniaceae subf. Blepharido-

phylloideae Schust., Hep. Anthoc. N. Amer.

3: 175. 1974. Blepharidophyllaceae (Schust.)

Schust. in Clarke & Duckett, Syst. Assoc. Spe-

cial Vol. 14: 74. 1980, nom. inval.

This family of two genera is strictly austral in

distribution. Blepharidophyllum has five species,

three centered in the American sector and two

Australasian. The southern South American Kru-

nodiplophyllum is monotypic.

Blepharidophyllum

Blepharidophyllum Angstr., Ofvers. Forh. Kongl.

Svenska Vetenskapsakad. 30: 151. 1873.

Key to Falkland Islands Species of Klepharidophyllum''

1 . Leaves not keeled, not conduplicately bilobed, bifid to the middle, the insertion curved; leaf cells

with large knotlike trigones; perianth mouth with long cilia which consist of long cylindrical cells

B. densifolium

1. Leaves keeled, conduplicately bilobed, divided 0.75-0.9 into 2 lobes, the insertion nearly transverse;

leaf cells ± equally thick walled, or with thin walls and small trigones; perianth mouth with teeth

and short spines, the latter of 1-2 massive, extended cells 2

2. Leaf margins irregular and with numerous small teeth, at least on antical margin of dorsal lobe;

leaf segments apiculate; cell walls thick, trigones absent B. clandestinum

2. Leaf margins entire; leaf segments triangular, not apiculate; cell walls thin, with small trigones

B. gottscheanum

Blepharidophyllum clandestinum (Mont.) Lac.

Figure 49.

Plagiochila (Scapania) dandestina Mont., Ann. Sci.

Nat. Bot. II, 19: 247. 1843. ? Scapania dandes-

tina (Mont.) G. L. & N., Syn. Hep. 73. 1844.

Jungermannia dandestina (Mont.) Hook. f. &

Tayl. i/i Hooker f., Bot. Antarc. Voyage 1(2): 434.

1 847. Martinellia dandestina (Mont.) Trev., Mem.

Reale 1st. Lomb. Sci. Lett. Ill, 4: 41 1. 1877. Bal-

antiopsis dandestina (Mont.) Schiffn. in Engler &

Prantl, Naturl. Pflanzenfam. 1(3, 1): 112. 1893.

Diplophyllum dandestinum (Mont.) Steph., Bih.

Kongl. Svenska Vetenskapsakad. Handl. 26 (III,

6): 61. 1900. Blepharidophyllum dandestinum

(Mont.) Lac., Genera Hep. 27. 1910. Clandarium

clandestinum (Mont.) Schust., New Manual

Bryology 1: 541. 1983. Original material: (fide

Grolle, 1965b, exd. B. gottscheanum): Chile, Prov.

Magallanes, Pto. Gallant, "Inter caespites Hypni

fluitantis," Hombron (PC, non vidi).

Balantiopsis incrassata Mitt., J. Linn. Soc., Bot. 15:

197. 1876, syn.fide Mitten (1879). Original ma-

terial: Kerguelen Is., Hill NW of Mt. Crozier, Ea-

ton (NY!).

18 Key adapted and modified from Grolle (1965b).

ECOLOGY— Rather local in occurrence in the

Falklands, where I found it on soil in a Cortaderia

heath, on the bases ofBlechnum magellanicum in

a dwarf shrub heath, and on a sheltered high al-

titude cliff.

PHYTOGEOGRAPHY — Amphiatlantic Temper-

ate—Falkland Is.; Tierra del Fuego; Patagonian

Channels; Valdivian region (to 42°30'S in West

Patagonia); also on Kerguelen (see fig. 49).

FALKLAND SPECIMENS SEEN -EAST FALKLANDS.

Mt. Usborne Region: ridge between Mts. Usborne 1 &

2, 685 m (2525A)- below The Gap, ca. 90 m (2566).

WEST FALKLANDS. Weddell Island: Waterfall Val-

ley, west of settlement, ca. 125 m (3313A—C. $, 3318).

Blepharidophyllum densifolium (Hook.) Angstr.

ex Mass. Figure 50.

Jungermannia densifolia Hook., Musci Exot. 1 : pi. 36,

f. 1-4. 1818. ? Scapania densifolia (Hook.) Nees

in G. L. & N., Syn. Hep 72. 1844. Diplophyllum

densifolium (Hook.) Mitt., J. Linn. Soc., Bot. 1 5:

69. 1876. Martinellia densifolia (Hook.) Trev.,

108

FIELDIANA: BOTANY

Mem. Reale 1st. Lomb. Sci. Lett. Ill, 4: 4 1 1 . 1 877.

Blepharidophyllum densifolium (Hook.) Angstr.

ex Mass., Nuovo Giorn. Bot. Ital. 17: 208. 1885.

Original material: Argentina, Terr. Tierra del

Fuego, I. de los Estados, Menzies (NY!, s, w), cited

inGrolle(1965b).

Jungermannia chloroleuca Hook. f. & Tayl., London

J. Bot. 3: 467. 1844, syn.fide Bescherelle & Mas-

salongo ( 1 889). Scapania chloroleuca (Hook. f. &

Tayl.) G. L. & N., Syn. Hep. 662. 1847. Schis-

tocalyx chloroleuca (Hook. f. & Tayl.) Lindb., J.

Linn. Soc., Bot. 13: 185. 1873, nom. illeg. Mar-

tinellia chloroleuca (Hook. f. & Tayl.) Trev., Mem.

Reale 1st. Lomb. Sci. Lett. Ill, 4: 41 1. 1877. Ble-

pharidophyllum vertebrale var. /3 chloroleucum

(Hook. f. & Tayl.) Mass., Nuovo Giorn. Bot. Ital.

1 7: 208. 1885. Blepharidophyllum densifolium var.

7 chloroleucum (Hook. f. & Tayl.) Schiffn. in Nau-

mann, Forschungsr. Gazelle 4(4): 9. 1 890. Diplo-

phyllum vertebrale var. /3 chloroleucum (Hook. f.

& Tayl.) Mass., Atti Reale 1st. Veneto Sci. 87:

221. 1927. Original material: Chile, Prov. Ma-

gallanes, I. Hermite, Hooker (NY!).

Scapania pycnophylla De Not., Mem. Reale Accad.

Sci. Torino II, 16: 215. pi. 3, f. 1-6. 1855, syn.

fide Bescherelle & Massalongo (1889). Martinellia

pycnophylla (De Not.) Trev., Mem. Reale 1st.

Lomb. Sci. Lett. Ill, 4: 411. 1877. Wlepharido-

phyllum pycnophyllum (De Not.) Angstr. ex Mass.,

Nuovo Giorn. Bot. Ital. 17: 208. 1885. Blephar-

idophyllum densifolium var. < pycnophyllum (De

Not.) Schiffn. in Naumann, Forschungsr. Gazelle

4(4): 9. 1890. Diplophyllum pycnophyllum (De

Not.) Steph., Result. Voyage S. Y. Belgica 6(5):

6. 1901. Original material: Chile, "Prov. Valpa-

raiso, Valparaiso," Puccio (NY!).

Blepharidophyllum fuscum Besch. in Bescherelle &

Massalongo, Mission Sci. Cap Horn 5: 209. 1889,

nom. nud. pro syn. Blepharidophyllum densifol-

ium var. 6 fuscum (Besch.) Schiffn. in Naumann,

Forschungsr. Gazelle 4(4): 9. 1 890.

ECOLOGY— In southern South America, the

species is quite common and exhibits a wide eco-

logical amplitude. It is common in mounds of

bryophytes in the very wet Magellanian moorland

region, as well as in the mesic, deciduous forested

zones. The species is surprisingly local in the Falk-

lands, where it was found in pools, rock crevices,

and between Bolax as well as Azorella cushions

on sheltered high altitude cliffs. It rarely occurs on

soil of dwarf shrub heaths. On Marion and Prince

Edward Islands B. densifolium is the dominant

plant species in a component of the "Swamp Com-

plex" termed a "Blepharidophyllum densifolium

bog" by Huntley (1971). This "nodum" is char-

acterized by nearly stagnant anaerobic conditions

and by a deep, primarily bryophyte carpet with

usually less than 20% vascular plant cover. (See

also Gremmen, 1982, who has a "Juncus

scheuchzerioides-Blepharidophyllum densifolium

complex" for the same islands.)

PHYTOGEOGRAPHY— Amphiatlantic Temper-

ate—South Georgia; Falkland Is.; Tierra del Fue-

go; Patagonian Channels and north in the Valdi-

vian region (West Patagonia) to 36°50'S. Quite

well represented in the subantarctic, on Kerguelen

Is., Crozet Is., Marion and Prince Edward Is., and

Gough I. (see fig. 50).

FALKLAND SPECIMENS SEEN -EAST FALKLANDS.

Stanley Region: Sapper Hill, 2 km W of Port Stanley,

100 m, Ochyra 2876/80 (F); S slopes of Mt. William,

1 85 m, Longton & Smith, Longton 989 (AAS, F); summit

ridge of N peak of Two Sisters, 245-290 m (2727). Mt.

Usborne Region: ridge between Mts. Usborne 1 & 2, 685

m (2526); summit of Mt. Usborne 1, ca. 700 m (2490,

2493A, 2494, 2504A). WEST FALKLANDS. Mt. Adam:

ridge S of northern lake, 610 m (3032).

Blepharidophyllum gottscheanum (nolle. Figure

51.

Blepharidophyllum gottscheanum Grolle, J. Hattori

Bot. Lab. 28: 69. / 4. 1965. Clandarium gotts-

cheanum (Mont.) Schust., New Manual Bryology

1: 541. 1983. Holotype: Chile, Prov. Magallanes,

I. Desolation, Dusen (JE, non vidi) cited in Grolle

(1965b).

ECOLOGY— I found this species at but a single

locality in the Falklands, where it grew in a Cor-

taderia heath at 90 m.

PHYTOGEOGRAPHY— Valdivian + Magellanian

+ Falklands— Falkland Is.; western Tierra del

Fuego; Patagonian Channels and north to 40°45'S

in the Valdivian region (West Patagonia) (see fig.

51).

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Mt. Usborne Region: below The Gap, ca. 90 m (2562B,

2578B).

Family GEOCALYCACEAE

Geocalycaceae Klinggr., Die hoheren Cryptoga-

men Preussens 34. 1858 ("Geocalyceae").

Geocalycaceae, composed of 20 genera, are

worldwide in distribution but much better repre-

sented in the Southern Hemisphere. Several small

genera are restricted to south temperate-subant-

arctic regions (e.g., Evansianthus and Pigafettoa

[each 1 sp.], Leptophyllopsis [2 spp.], Pachyglossa

[4 spp.], and the larger genera are best represented

there [e.g., Clasmatocolea, Heteroscyphus]). The

largest genus is Chiloscyphus (s. lat. to include

Lophocoled).

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

109

50

,

FIG. 50. Distribution of

Blepharidophyllum densifolium

(Hook.) Angstr. ex Mass.

FIG. 51. Distribution of

Blepharidophyllum gottscheanum

Grolle. Open circle indicates pre-

cise locality unknown.

110

FTELDIANA: BOTANY

Chiloscyphus

Chiloscyphus Corda in Opiz (ed.), Beitrage zur

Naturg. ( 1 2): 65 1 . 1 829 ("Cheilocyphos"); coir.

Dumortier, Sylloge Jung. Europ. Indig. 67.

1831, nom. et orth. cons. Lophocolea (Dum.)

Dum., Recueil Observ. Jungerm. 17. 1835.

Jungermannia sect. Lophocolea Dum., Syll.

Jungerm. Eur. Indig. 59. 1831.

Key to Falkland Islands Species of Chiloscyphus

1. Leaves consistently or at least predominantly bifid 2

2. Leaf lobes and marginal teeth caducous, often giving leaf apices a ragged appearance; leaves often

with accessory teeth and laciniae; leaf margins entire to highly dentate. Underleaves deeply bifid,

with margins 1-laciniate, and lobes spreading; perianths twice as long as broad; either margin of

innermost bract to 8-dentate-laciniate; plants dioecious [Leptophyllopsis irregularis]

2. Leaf lobes and marginal teeth, if present, persistent, leaf apices never with a ragged appearance;

leaves not with accessory teeth and laciniae; leaf margins entire to 1-dentate 3

3. Leaves bifid to (0.3)0.4-0.5; leaf segments frequently ventrally sulcate 4

4. Leaf segments widely divergent; leaves basically weakly to strongly cuneate, the lamina

widest in distal third; underleaf margins mostly entire, sporadically with a small tooth . .

C. divaricatus

4. Leaf segments ± parallel with lamina margins at most only sporadically divergent; leaves

basically ovate to subrectangular, the lamina widest in basal third; underleaf margins usually

with 1 lacinium or tooth C. bispinosus

3. Leaves bidentate or bifid to at most 0.25; leaf segments plane, never ventrally sulcate .. 5

5. Underleaves connate with the leaves on both sides. Leaves symmetrically ovate or nearly

so to subrectangular in shape, the sinus truncate or lunate C. sylvaticus

5. Underleaves connate on only one side or completely free 6

6. Median leaf cells (46)52-78 /*m x 33-65 jmi; perianth terete or obscurely triangular

below. Dorsal leaf margin straight or nearly so, entire; ventral margin rounded, especially

towards the apex, entire to 1-dentate; underleaves elongate-ovate in shape, as wide as

or slightly wider than stem, connate on one side C. textilis

6. Median leaf cells 17-48(52) x 20-43 Aim; perianth triangular below 7

7. Leaf lobes variable; 1 or both commonly rather broadly rounded (often with 1 lobe

rounded and the other triangular), or both narrowly triangular. Lobes asymmetric,

with one lobe often larger than the other; trigones distinct C. humifusus

1. Leaf lobes consistent in shape, narrow to broadly triangular or piliferous. Lobes ±

symmetric 8

8. Leaf segments ending in a uniseriate row of 1-6 cells, the segments narrow to

broadly triangular. Underleaf margins entire to 1-dentate or occasionally ciliate-

small laciniate; plants monoecious C. lentus

8. Leaf segments ending in a uniseriate row of 4-15 cells, the segments piliform.

Underleaves deeply bifid (nearly always almost to base), the lamina margins

usually with a single long lacinium-lobe; perianths (3.1)3.6-5.3 x longer than

broad, the lobes narrowly triangular, dentate-laciniate; plants monoecious ....

C. leptanthus

1 . Leaves (well developed) undivided, emarginate or 1-dentate 9

9. Living plant whitish to light to yellow-green or tawny, without wall pigments. Gynoecia isophyllous

or subisophyllous: bracteole 0.75-0.95 the area of the bracts 10

10. Underleaf apices plane to retuse to bidentate; underleaves/re<7w^rt//y convex to ± canaliculate,

the margins entire. Underleaves connate on one side; leaf apices entire .... C. austrigenus

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA 1 1 1

10. Underleaves bifid to ca. 0.5 or more; underleaves plane and not convex, the margins entire

to 4-dentate-laciniate 11

1 1 . Leaves expanded near ventral base and here often reflexed; underleaves 0.5 x stem width

to as wide as stem. Leaf apices broadly rounded, never retuse; underleaf segments often

differing in size and configuration; stems thick, fleshy C. elatus

\ 1 . Leaves not expanded near ventral base; underleaves never less than stem width ... 12

1 2. Leaf apices variable, broadly rounded to truncate to retuse to emarginate to 1 - or

bidentate; underleaves as wide as to 1.5x stem width; underleaf margins entire to

2-dentate-small laciniate C. sabuletorum

12. Leaf apices broadly rounded to truncate, never retuse; underleaves (l-)3x stem

width; underleaf margins to 4-dentate-large laciniate C. semiteres

9. Living plant warm to castaneous brown, with wall pigmentation. Gynoecia strongly anisophyllous:

bracteole ca. 0.25-0.3 as large as bracts, 0.5-0.75 bifid, underleaf-like and often with 1 lateral

lacinium on 1 or both sides. Perianths most often on abbreviated lateral-intercalary branches

without vegetative leaves C. hookeri

Chiloscyphus austrigenus (Hook. f. & Tayl.) Engel

& Schust. Figure 52.

Jungermannia austrigena Hook. f. & Tayl., London

J. Bot. 3: 466. 1 844. Lophocolea austrigena (Hook,

f. & Tayl.) G. L. & N., Syn. Hep. 702. 1847.

Chiloscyphus austrigenus (Hook. f. & Tayl.) Engel

& Schust., Nova Hedwigia 39: 411. 1984. Orig-

inal material: Chile, Prov. Magallanes, I. Hermite

and/or Cabo de Hornos, Hooker s.n. (BM!— c. per.;

NY!— c. per.; s!; w, cited in Grolle, 1962).

Jungermannia cavispina Hook. f. & Tayl., London J.

Bot. 3: 463. 1844, syn. fide Bescherelle & Mas-

salongo (1 889). Lophocolea cavispina (Hook. f. &

Tayl.) Fulf., Mem. New York Bot. Gard. 1 1 : 483.

1976, nom. illeg. (ICBN, Art. 57). Lectotype (fide

Engel, 1978): Falkland Is., 1843, Hooker s.n. (FH!).

Lophocolea triseriata Steph., Bih. Kongl. Svenska Ve-

tenskapsakad. Handl. 26 (III, 6): 45. 1900, syn.

fide Fulford (1976). Original material: Chile, Prov.

Aisen, R. Aisen Valley, 13 Jan. 1897, Dusen s.n.

(NY!, s!— c. per., UPS!).

Leioscyphus grandistipus Steph., Kongl. Svenska Ve-

tenskapsakad. Handl. 46(9): 37. / 13c,d. 1911,

syn. fide Grolle ( 1 962). Leptoscyphus grandistipus

(Steph.) Kiihnem., Revista Centre Estud. Doct.

Ci. Nat. Buenos Aires 1: 176. 1937. Mylia gran-

distipa (Steph.) Kuhnem., Lilloa 19: 341. 1949.

Lectotype (fide Engel, 1 978): Falkland Is., Hornby

Mts., 19 Dec. 1907, Skottsberg 346 (s!).

Lophocolea falklandica Steph., Kongl. Svenska Ve-

tenskapsakad. Handl. 46(9): 44./ 16e. 191 1, syn.

fide Grolle (1962). Lectotype (fide Engel, 1978):

Falkland Is., Westpoint I., 8 Dec. 1907, Skotts-

berg 351 (s!— c. S).

REMARKS— See Engel (1978).

Chiloscyphus austrigenus is the correct name for

this species, and not Lophocolea cavispina, as Ful-

ford (1976) would have it; see Engel (1979) and

Grolle (1979). Further, Fulford erroneously in-

cluded several synonyms of this species. Chiloscy-

phus otiphyllus is a well-differentiated species, with

Chiloscyphus notophylloides Mass, and Leioscy-

phus oppositifolius Steph. as synonyms (see Engel,

1978). Lophocolea magellanica Schiffn. is a syn-

onym of Clasmatocolea humilis var. humilis (fide

Engel, 1980).

Several syntype specimens of Lophocolea falk-

landica have been misdetermined. These speci-

mens are referable to Balantiopsis erinacea and

Chiloscyphus elatus. The description (especially of

the underleaf) and figures, however, agree with the

syntypes referable to Chiloscyphus austrigenus. I

have designated a Stockholm (S) specimen from

Westpoint Island, Falkland Islands, as the lecto-

type. Chiloscyphus elatus is mixed with C. austri-

genus in the lectotype packet.

ECOLOGY— This taxon is rather common in and

typical of the bryoflora of streams in Cortaderia

and dwarf shrub heaths. It is especially luxuriant

when submerged, and then frequently forms thick,

pure, extensive masses. It occasionally may be

found on stream banks.

PHYTOGEOGRAPHY— Valdivian + Magellanian

Falklands— Falkland Is.; Tierra del Fuego; Pata-

gonian Channels; Valdivian region (north to

39°52'S); Tristan da Cunha; and Inaccessible I.

(see fig. 52). There have been no authentic reports

from Andean Patagonia; the report in Stephani

(1911) from Estrecho Miguens is based upon a

specimen of Clasmatocolea vermicularis. The re-

ports from the New Zealand sector require con-

firmation.

LITERATURE RECORDS (FALKLANDS)— Anony-

mous (Kiihnemann, 1937, as Leptoscyphus gran-

distipus, Lophocolea austrigena & L. falklandica;

112

FIELDIANA: BOTANY

52

FIG. 52. Distribution of Chi-

loscyphus auslrigenus (Hook. f. &

Tayl.) Engel & Schust., plus Tris-

tan da Cunha. Open circle indi-

cates precise locality unknown.

FIG. 53. Distribution of Chi-

loscyphus elatus (Gott.) Engel &

Schust.

53

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

113

Kiihnemann, 1949 as L. austrigena, L. falklandica

& Mylia grandistipus; Rodway, 1916); Abbott

(Fulford, 1976, as Lophocolea cavispina); Hooker

(G. L. & N. 1 847 as J. cavispina; Taylor & Hooker,

1847, as J. cavispina; Stephani, 1906); Lechler,

Mt. William, Port Stanley (Fulford, 1976, as L.

cavispina); Skottsberg (Stephani, 1 922, as Leioscy-

phus grandistipus & Lophocolea falklandica), be-

tween Arrow Harbour House and Victoria Creek

(Skottsberg, 1913, as Leioscyphus grandistipus);

Hornby Mts. (Grolle, 1 962); Port Stanley (Fulford,

1976, as Lophocolea cavispina); Port Stanley, Port

Louis (Skottsberg, 1913; Stephani, 1905a); West-

point Is. (Stephani, 1913, as L. falklandica);

Skottsberg & Halle, Murrell River (Stephani, 1911;

Fulford, 1976, as Lophocolea cavispina).

FALKLAND SPECIMENS SEEN— without specific locality,

Capt. Abbott (BM, NY-C. per.). EAST FALKLANDS.

1 843, Hooker, syntypes of Jungermannia cavispina (BM,

s). Stanley Region: Port Stanley, 28 Mar. 1902, Skotts-

berg s.n. (s); Port Stanley, Lechler 99 (PC); headwaters of

Mullet Creek Stream, ca. 60 m (3160, 3164, 3168-3169);

Murrell River, 6 Nov. 1907, Halle (s, UPS); Mt. William,

Lechler 99 (NY); Moody Brook, near Stanley, 8 m, Long-

ton & Smith, Longton 961 (AAS, F). Mt. Usborne Region:

valley SW of Mt. Usborne, ca. 60 m (2647, 2651, 2652B,

2656). WEST FALKLANDS. Mt. Adam: basin E of

summit, 580-595 m (2999). Westpoint Island: 8 Oct.

1907, Skottsberg 351, syntype of Lophocolea falklandica

(UPS); near The Waterfall, 30-90 m (2864, 2870-2873,

2877). Weddell Island: Waterfall Valley, W of settle-

ment, ca. 125 m (3311, 3314, 3317, 3331-3332).

Chiloscyphus bispinosus (Hook. f. & Tayl.) Engel

& Schust.

Jungermannia bispinosa Hook. f. & Tayl., London J.

Hot. 3: 378. 1844. Lophocolea bispinosa (Hook,

f. & Tayl.) G. L. & N., Syn. Hep. 162. 1845.

Chiloscyphus bispinosus (Hook. f. & Tayl.) Engel

& Schust., Nova Hedwigia 39: 41 1. 1984. Orig-

inal material: Campbell I., Hooker (non vidi).

ECOLOGY -Collected only once in the Falk-

lands, on soil over rock in a dwarf shrub heath.

PH YTOGEOGRAPH Y — Amphi pacific Temper-

ate -Macquarie I.; Campbell I.; New Zealand;

Tasmania; Australia (Victoria); Falkland Is.; Isles

of Scilly (Paton, 1974); Isle of Colonsay, Scotland

(Wallace, 1979). Reported from southern South

America by Massalongo (1885, 1927); these, how-

ever, are erroneous, for they are misdetermina-

tions of Chiloscyphus leptanthus (fide NY!, VER!).

FALKLAND SPECIMEN SEEN— WEST FALKLANDS.

Weddell Island: summit of Mt. Weddell, 385 m (3347).

Chiloscyphus divaricatus (Hook. f. & Tayl.) Engel

& Schust.

Lophocolea divaricata Hook. f. & Tayl., London J.

Bot. 5: 367. 1846, non L. divaricata Herz., Arch.

Esc. Farm. Fac. Ci. Med. Cordoba 7: 19. 1938.

Jungermannia divaricata (Hook. f. & Tayl.) Hook,

f. & Tayl. in Hooker f., Bot. Antarc. Voyage 1(2):

437. 1847, non J. divaricata Sm. in Sowerby, En-

glish Bot. 10: 719. 1800 (= Cephaloziella), necJ.

divaricata Nees, Enum. Plant Crypt. Javae 60.

1830 (cf. Acromastigum). Chiloscyphus divari-

catus (Hook. f. & Tayl.) Engel & Schust., Nova

Hedwigia 39: 414. 1984. Original material: Chile,

Prov. Magallanes, I. Hermite, Hooker (NY!).

ECOLOGY— This species is rare not only in the

Falklands, but also in southern South America.

Falkland plants are restricted to tussock associa-

tions, where the species occurs on moist litter on

the sides or bases of tussock (Poa flabellata) ped-

estals, or between the pedestals. It usually occurs

in admixtures with other hepatics, such as Chi-

loscyphus lentus or Chiloscyphus sabuletorum.

PHYTOGEOGRAPHY— Valdivian + Magellanian

+ Falklands— Falkland Is.; Tierra del Fuego; Pat-

agonian Channels (Brunswick Peninsula); and

Valdivian region (45°17'S, 73°43'W). Within the

Falklands, it was found only in the Port Stanley

region, and Beauchene Is. (52°54'S, 59°09'W).

LITERATURE RECORD (FALKLANDS) — Smith,

Beauchene Is. (Smith & Prince, 1985).

FALKLAND SPECIMENS SEEN— Beauchene Is.: "The

Hole," 25 m, Smith 3018, 3019d(AAS, F); above "Blinn's

Cove," SE side of island, 10-20 m, Smith 3016 (AAS, F).

EAST FALKLANDS. Kidney Island: north of shanty

(2788B).

Chiloscyphus elatus (Gott.) Engel & Schust. Figure

53.

Jungermannia elata Gott., Ergebn. Deutsch. Polar-

Exped. 2(16): 450.pl. 7,f. 3-6. 1890. Lophocolea

elata (Gott.) Steph., Wiss. Ergebn. Schwed. Siid-

polar-Exped. 4(1): 7. 1905. Chiloscyphus elatus

(Gott.) Engel & Schust., Nova Hedwigia 39: 414.

1 984. Original material: South Georgia, 1 882, Will

(M), cited in Grolle (1962) (non vidi).

Leioscyphus set istipus Steph., Bih. Kongl. Svenska Ve-

tenskapsakad. Handl. 26 (III, 17): 16. 1901, syn.

fide Grolle (1962). Leptoscyphus setistipus (Steph.)

Kiihnem., Revista Centra Estud. Doct. Ci. Nat.,

Buenos Aires 1: 177. 1937. Mylia setistipa (Steph.)

Kuhnem., Lilloa 19: 341. 1949. Original mate-

rial: Chile, Prov. Magallanes, R. Azopardo, 1 896,

Dusen 56 (G, s), cited in Grolle (1962) (non vidi).

ECOLOGY— This species seems to be restricted

to stream habitats in dwarf shrub heath and Cor-

114

FIELDIANA: BOTANY

taderia associations, where it was, without excep-

tion, submerged or nearly so. This species, with

C. austrigenus, is characteristic of the submerged

stream bryoflora.

PHYTOGEOGRAPHY— Magellanian-Falkland —

South Georgia; Falkland Is.; Fuegia (Isla Grande

de Tierra del Fuego); southern Patagonian Chan-

nels (Brunswick Peninsula). The report from Gru-

po Evangelistas in Stephani (1911) requires in-

vestigation (see fig. 53).

LITERATURE RECORD (FALKLANDS)— Skottsberg,

(?)Eastpoint Is. (Grolle, 1962).

FALKLAND SPECIMENS SEEN -EAST FALKLANDS.

Lyall, as Jungermannia vasculosa (FH). Mt. Usborne Re-

gion: valley SW of Mt. Usborne, ca. 60 m (2667). WEST

FALKLANDS. Fox Bay Region: near mouth of Cheek's

Creek, ca. 12 m (3496A); near peat cuttings in Ram

Paddock, 45-60 m (3518); valley E of Sulivan House,

ca. 75 m (3524, 3528). Westpoint Island: Skottsberg as

L. falklandica (o, UPS); near The Waterfall, 39-90 m

(2559).

Chiloscyphus hookeri Engel

PHYTOGEOGRAPHY— Magellanian-Falkland —

Falkland Is., Tierra del Fuego, and Patagonian

Channels north to 49°02'S.

REMARKS— Fulford (1976) placed Chiloscyphus

hookeri in the synonymy of Leptoscyphus palli-

dovirens, regarding the two taxa as "two extremes

of another variable species of Leptoscyphus" where

she transferred the species. Over the last decade I

have studied many collections of both species, and

regard the two taxa as quite distinct entities which

do not even approach one another. See Engel

( 1 972b, 1978) for key characters which I have used

to differentiate the species. I have outlined reasons

for classifying C. hookeri in Chiloscyphus (see En-

gel, 1972b,p. 154; see also Engel & Schuster, 1984,

p. 409).

This species has two varieties (see Engel, 1 972b);

only the typical variety occurs in the Falklands. It

differs from var. constant ifolius Engel in the vari-

able leaf apices— broadly rounded to truncate to

retuse to single lobed to bifid, even on a single

axis.

Chiloscyphus hookeri Engel var. hookeri

Chiloscyphus hookeri Engel var. hookeri. J. Hattori

Bot. Lab. 36: I50.pl. 12. 1973.

Holotype— Chile, Prov. Magallanes, I. Hermite,

Hooker 12, mixed with syntype plants of Junger-

mannia pallido-virens (NY!).

ECOLOGY— Rather common in dwarf shrub

heaths, especially over 155 m, where it grows un-

der rock overhang or Blechnum magellanicum

cover in deep rock crevices, etc., more rarely on

stream banks in this association, and less common

under grass cover in Cortaderia heaths.

FALKLAND SPECIMENS SEEN -EAST FALKLANDS.

Port William Region: summit ridge of Mt. Low, ca. 245

m (3227-3228); ridge on Engineer Point Peninsula, 1 8-

25 m (2820— c. per.). Stanley Region: S slopes of Mt.

William, 185 m, Longton & Smith, Longton 984 (AAS,

F); Tumbledown Mt., 155-230 m (2372B-C. 9, 2386);

Goat Ridge, ca. 180 m (3197— c. per., 3209-c. per.);

summit ridge of N peak of Two Sisters, 245-290 m

(2704. 2710C-C. per.). Mt. Usborne Region: The Gap,

275-290 m (2462A-C. S); S side of The Gap, 260-275

m (2454 -c. per.). WEST FALKLANDS. Port Howard:

Freezer Rocks, on E slope of Mt. Maria, 320 m (3137).

Hornby Mts: 19 Dec. 1907, Halle & Skottsberg 223 as

Lophocolea pallido-virens (UPS). Fox Bay Region, sum-

mit of East Head, 180 m (3437); near peat cutting in

Ram Poddock, 45-60 m (3511-c. S). Mt. Adam, 13

Dec. 1907, Halle & Skottsberg 223 as Lophocolea pal-

lido-virens (UPS). Weddell Island: summit of peak NE of

Mt. Weddell, 335 m (3369E); summit of Mt. Weddell,

380 m (3334. 3337-c. a, 3343A).

Chiloscyphus humifusus (Hook. f. & Tayl.) Engel

& Schust.

Jungermannia humifusa Hook. f. & Tayl., London J.

Bot. 3: 472. 1 844. Lophocolea humifusa (Hook,

f. & Tayl.) G. L. & N., Syn. Hep. 695. 1847.

Chiloscyphus humifusus (Hook. f. & Tayl.) Engel

& Schust., Nova Hedwigia 39: 416. 1984. Orig-

inal material: Kerguelen Is., Hooker (NY!; s!, ex

"Herbarium Lehmannianum").

REMARKS— This is a distinctive species which

may be readily identified by the following features:

1) variable leaf lobes: one or both are commonly

rather broadly rounded (often with one lobe

rounded and the other triangular), or both are nar-

rowly triangular; 2) leaf lobe asymmetry: the ven-

tral lobe is usually larger and the dorsal is often

reduced to a low, rounded projection or is absent

altogether (see Taylor & Hooker, 1847, pi. 159, f.

5); in some populations, however, the ventral lobe

is often smaller than the dorsal; 3) underleaves

bifid nearly to the base, with the lamina between

sinus base and underleaf base only a few cells high;

4) underleaf lobes setaceous, sometimes quite

markedly so; 5) trigones distinct, medium and

straight-sided to large and slightly bulging; and 6)

underleaf lamina with a pronounced lacinium or

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

115

cilium. The last is a curious appendage that varies

from erect to spreading. At times its attachment

to the underleaf is quite subtle, and it superficially

appears to insert on the stem. I have not seen

"stipules sometimes with a setaceous process at

each side running back from their insertion along

the stems" as described by Hooker and Taylor

(1844) in the protologue. However, I have ob-

served (in the type) long-setaceous underleaf seg-

ments that extend distally to the basal-lateral por-

tion of the next underleaf above, and superficially

appear to originate there. It might possibly be this

condition that was the basis for Taylor and Hook-

er's figures and mention of underleaf processes. It

is doubtful that they are rhizoids, for Taylor and

Hooker ( 1 847) illustrate them as multicellular.

Intercalary branches in Engel 3382 are plastic

in origin and may be ventral, ventral-lateral, or

lateral. Lateral branches may issue from above or

below the leaf insertion and near the median por-

tion of the leaf or near its ventral base. They may

often be found between the underleaf base and the

ventral base of the leaf immediately basal to it.

Most intercalary branches remain dormant and

"budlike"; some elongate slightly, and are very

narrow compared to the main axis. I know of no

other southern South American Chiloscyphus with

intercalary branches of this kind. Frullania-type

branches are also common.

Fulford (1976) placed the species as a synonym

of Lophocolea leptantha.

ECOLOGY— Rather rare in Falklands; my collec-

tions are from rocky outcrops above 335m where

over rock or on soil under partial rock cover. The

Longton and Smith plant was collected at a much

lower elevation (30 m) on moist sandy sheltered

vertical north-facing rocks.

PHYTOGEOGRAPHY — Subantarctic — Kerguelen

Is.; Falkland Is. This is the first authentic record

of this species from the Falkland Islands. Stephani

(1911) reported Chiloscyphus humifusus from the

Falklands, and later Kuhnemann (1937, 1949) in-

cluded this reference in his catalogs. The reports

are in error, as the specimen is a misdetermination

of Leptoscyphus patagonicus. The first of the two

reports of the taxon in Stephani (191 1) is a Valle

Frias collection, which is a misdetermination of

Clasmatocolea humilis. Further, in the Uppsala

Herbarium, the following specimens are misde-

termined by Stephani as L. humifusa: 1896, Dusen

31 from Punta Arenas (a specimen of Clasmato-

colea rigens mixed with Chiloscyphus leptanthus)

and 1896, Dusen 304 from Ushuaia (a specimen

of Chiloscyphus textilis). All remaining records of

the species from southern South America require

confirmation.

FALKLAND SPECIMENS SEEN— EAST FALKLANDS.

Stanley Region: S. shore of Murrell River, opposite Islet

Point, 30 m, Longton & Smith, Longton 1026 (AAS, F).

WEST FALKLANDS. Port Howard: pass SW of Mt.

Maria summit, ca. 610 m (3081, 3105, 3125). Weddell

Island: summit of peak NE of Mt. Weddell, 335m (3382,

3398).

Chiloscyphus lentus (Hook. f. & Tayl.) Engel &

Schust.

Jungermannia lenta Hook. f. & Tayl., London J. Bot.

3: 379. 1 844. Lophocolea lenta (Hook. f. & Tayl.)

G. L. & N., Syn. Hep. 162. 1845. Chiloscyphus

lentus (Hook. f. & Tayl.) Engel & Schust., Nova

Hedwigia 39: 418. 1984. Original material: Auck-

land Is., Hooker (non vidi).

Jungermannia secundifolia Hook. f. & Tayl., London

J. Bot. 3:471. lS44,syn.fideMillen(\&55). Loph-

ocolea secundifolia (Hook. f. & Tayl.) G. L. & N.,

Syn. Hep. 693. 1 847. Leclolype (fide Engel, 1978):

Falkland Is., Hooker (FH!).

REMARKS— See Engel (1978).

ECOLOGY— This species is common on bases of

tussock in the tussock association. It is also present

on outcrops in dwarf shrub heaths where it may

be found on soil under rock overhangs and in crev-

ices. It was also found on bark of Hebe, Notho-

fagus, and fir (the latter two in the grove of planted

trees at Hill Cove settlement).

PHYTOGEOGRAPHY — Amphipacific — Reported

from South Sandwich Is. and South Georgia

(Grolle, 1972b; Longton & Holdgate, 1979, both

as L. secundifolia); Falkland Is.; Tierra del Fuego;

southern Patagonian Channels (Brunswick Pen-

insula); the Valdivian region in the American sec-

tor; Macquarie, Auckland, Campbell, and Snares

Is., New Zealand, Tasmania, and Australia in the

New Zealand sector.

LITERATURE RECORDS (FALKLANDS)— Anony-

mous (Kuhnemann, 1937, 1949, as L. secundifo-

lia); Hooker (G. L. & N., 1847, as L. secundifolia;

Taylor & Hooker, 1847, as /. secundifolia; Ste-

phani, 1906, as L. secundifolia); Skottsberg, Port

Stanley, Port Louis (Stephani, 1905a, as L. secun-

difolia), Port Stanley (Skottsberg, 1913, as L. se-

cundifolia); Skottsberg & Halle, Port Louis, Half-

way Cove (Stephani, 1911, as L. secundifolia);

Smith, Beauchene Is. (Smith & Prince, 1985).

FALKLAND SPECIMENS SEEN— Beauchene Is.: Above

Blinn's Cove, SE side of island, 10-20 m., Smith 3015

(AAS, F). EAST FALKLANDS. 1843, Hooker, syntypes

116

FIELDIANA: BOTANY

of Jungermannia secundifolia(BM, NY— c. per. + <5); Lyall

as L. secundifolia (FH). KIDNEY ISLAND, 1954, Pet-

tingill 16 (CMC, F), 77 (CINC); N of shanty (27 88 A, 2792.

2795. 2796. 2797, 2802-c. per. + a). Port William

Region: S shore of Cape Pembroke Peninsula, near Surf

Bay, sea level (2432. 2434— c. per. + S); ridge on En-

gineer Point Peninsula, 18-25 m (2837— c. 3 + young

9). Darwin Settlement: Boca House on Brenton Lock,

sea level (2689-c. sporo. + S). WEST FALKLANDS.

Fox Bay Region: summit of East Head, 1 80 m (3438A —

c. sporo., 3442— c. per. + <j). Hill Cove Region: settle-

ment, 15 m (3049— c. S + 2, 3050- c. 3 + 2, 3056— c.

S + 9). Westpoint Island: steep slope & cliffs facing The

Woolly Gut (292 3). Weddell Island: summit of peak NE

of Mt. Weddell, 335 m (3380-c. 5 + 2, 3390-c. per.

+ <5). New Island: ridge between Bold Hill and Bold

Point, 105 m (^265. 3269).

Chiloscyphus leptanthus (Hook. f. & Tayl.) Engel

& Schust. Figure 54.

Jungermannia leptantha Hook. f. & Tayl., London J.

Bot. 3: 471. 1844. Lophocolea leptantha (Hook,

f. & Tayl.) G. L. & N., Syn. Hep. 694. 1847.

Chiloscyphus leptanthus (Hook. f. & Tayl.) Engel

& Schust., Nova Hedwigia 39: 418. 1984. Lec-

totype(/Kfe Engel, 1978): Chile, Prov. Magallanes,

I. Hermite, Hooker s.n. (NY!— c. sporo.).

Jungermannia alternifolia Hook. f. & Tayl., London

J. Bot. 4: 83. 1 845, s>77../Kfc Fulford ( 1976). Loph-

ocolea alternifolia (Hook. f. & Tayl.) G. L. & N.,

Syn. Hep. 695. 1847. Lectotype (/kfc Engel, 1978):

Falkland Is. (non New Zealand), Lyall s.n. (FH! —

c. sporo.).

Lophocolea gibbosa Mont., Ann. Sci. Nat. Bot. Ill, 4:

351. 1845, syn.fide Fulford (1976). Original ma-

terial: Southern Chile, without specific locality,

Gay (PC!— c. per.).

Lophocolea cunninghamiiSleph., Bull. Herb. Boissier

6(8): 652. 1906 (= Spec. Hep. 3: 68), syn. fide

Fulford (1976). Original material: Chile, Prov.

Magallanes, Pto. Eden, ex hb. Kew, Apr. 1868,

Cunningham 56 (G!).

Lophocolea monoica Steph., Kongl. Svenska Vetensk-

apsakad. Handl. 46(9): 48. / 19k-p. 1911. syn.

fide Fulford (1976). Lectotype (fide Engel, 1978):

Falkland Is., Port Stanley, Sapper Hill, 29 Oct.

1907, Skottsberg s.n. (UPS!— c. per.).

REMARKS— See remarks in Engel (1978).

The type locality of Lophocolea alternifolia has

been confused. The original description cites New

Zealand as the type locality. A single sheet in the

Taylor herbarium (FH) has the following three sep-

arate specimens: a) a packet labeled "Falkland Is-

lands" but crossed out with "New Zealand" writ-

ten below, collected by Lyall; b) a small packet

along with a glued specimen labeled "number 1";

and c) a portion of b which has both been placed

in a small packet and glued to the sheet and labeled

"number 2." Taylor has noted below the speci-

mens that number 1 is from the Falkland Islands

and collected by Lyall and was communicated by

J. D. Hooker, 1844 (the following note is also in-

cluded here "with Chilosc. retusatus Tayl."), and

number 2 is from New Zealand, also collected by

W. Lyall and communicated by W. Wilson.

Taylor and Hooker ( 1 847, p. 436) stated of Jun-

germannia alternifolia: "Hab. Falkland Islands"

and "This species was erroneously described (Lon-

don J. Bot. I.e.) as a native of New Zealand; from

whence we have never seen specimens." I believe

that specimens labeled numbers 1 and 2 are merely

segregates of a single Falkland Islands-Lyall col-

lection, as both contain the following identical

complex of plants: Chiloscyphus leptanthus, a rath-

er flaccid form of Leptoscyphus expansus with

truncate leaf apices which are frequently emargin-

ate, a moss, and bits of grass. In addition, I have

studied a syntype collection from the Hooker

Herbarium of the British Museum, and this Lyall-

collected specimen bears the Falkland Islands

locality only. This specimen also contains, in ad-

dition to Chiloscyphus leptanthus, the Leptoscy-

phus expansus, a moss, and a grass. Specimen a

above, which is only a small segregate, has the

moss but not Leptoscyphus expansus or the grass.

The confusion surrounding this species likely orig-

inated during curating of the specimen after it left

Hooker and before it was studied by Taylor.

I consider the three Farlow plants to be one

collection that has been subdivided, and since two

of the specimens have perianths, one with a cap-

sule, I have designated these specimens as the lec-

totype. The British Museum specimen (syntype)

appears sterile.

The description and figures of Lophocolea mo-

noica in Stephani (1911) are somewhat confusing.

Part of this confusion arises out of the heteroge-

neous assembly of syntype material. The syntypes

from Territory Chubut (Arroyo Carbon), and Azo-

pardo-Tal are misdeterminations of Clasmatoco-

lea rigens. The syntypes from Isla Dawson, Falk-

land Islands (Rabbit Island) and Falkland Islands

(Sapper Hill) are plants of Lophocolea monoica.

The figures and descriptions of vegetative material

are of L. monoica, while those of the perianth

clearly do not correspond to the long, narrow peri-

anths normally produced by this taxon. The peri-

anth drawn and described may be that of a very

immature one, or one of C. rigens.

Stephani (Spec. Hep., Vol. 6, p. 283) stated for

L. monoica that plants are ""dilute brunnea" and

revised the description of the perianth for the

species, stating ""Perianthia anguste oblonga (7 mm

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

117

54

FIG. 54. Distribution of Chiloscyphus leptanthus

(Hook. f. & Tayl.) Engel & Schust. Open circle indicates

precise locality unknown.

longa, 2.5 mm laid) apice ad V3 triloba, lobis late

ligulatis, irregulariter multispinosis" The descrip-

tion of the perianth here fits that of L. monoica,

i.e., it corresponds to the vegetative material de-

scribed and drawn in 1911. Further, in 1922 Ste-

phani stated the species was collected in the Falk-

land Islands by Skottsberg. Thus, Stephani in the

latter reference has excluded South American col-

lections. Of the syntypes from the Falklands, that

from Sapper Hill has several plants with leaves

concave and decurved, and thus best agrees with

the description. The UPS collection by Skottsberg

s.n. (29 Oct. 1927) from the Falkland Islands (Sap-

per Hill) was designated the lectotype of Lopho-

colea monoica in Engel (1978).

The material of L. monoica agrees well with L.

leptantha and is here treated as a synonym.

Fulford (1976) considerably extended the dis-

tribution of this species by including, as synonyms,

C. humifusus from Kerguelen and three taxa from

Brazil. I accept C. humifusus as a distinct species,

and question the Brazil names as synonyms. For

the present I am restricting C. leptanthus to those

regions indicated below.

ECOLOGY— This taxon is rather rare in the Falk-

lands, occurring in wet depressions, on wet stones

partially submerged in streams, and on soil of dwarf

shrub heaths. The species was also found on the

bases of and in between tussock plants in a tussock

association. On Beauchene Is. the species occurs

on moist tussock litter between Poajlabellata ped-

estals, and label data accompanying the Smith col-

lection notes that it is "widespread in small

amounts— the only 'abundant' bryophyte on the

island". It also occurs in the grove of planted trees

(mainly hardwood) at Hill Cove settlement and

on the floor of the coniferous plantation at Gov-

ernment House in Stanley. See comments in Engel

(1978) for the ecology of the species in the Bruns-

wick Peninsula, where it is rather common.

PHYTOGEOGRAPHY— Valdivian + Magellanian

+ Falklands— Falkland Is.; Tierra del Fuego; Pat-

agonian Channels; the Valdivian region (West Pat-

agonia to 43°54'S). I have not seen the specimen

on which the Juan Fernandez report is based, and

its presence there is to be regarded as questionable.

There are no authentic reports of this species from

Andean Patagonia; the report of L. monoica from

Arroyo Carbon in Stephani (191 1) is a misdeter-

mination of Clasmatocolea rigens (see fig. 54).

LITERATURE RECORDS (FALKLANDS)— Anony-

mous (Kuhnemann, 1937, 1949 as L. cunning-

hamii & L. monoica); Hooker (Taylor & Hooker,

1847, as Jungermannia alternifolid); Pettingill—

Charles Point (Fulford, 1976); Skottsberg (Ste-

phani, 1 922 as L. monoica), Port Stanley, Rabbit

Is. (Skottsberg, 1913, as L. monoica), Port Louis

(Skottsberg, 1913; Stephani, 1905a, both as L.

cunninghamii); Smith, Beauchene Is. (Smith &

Prince, 1985).

FALKLAND SPECIMENS SEEN— Beauchene Is.: "The

Hole," 25 m, Smith J079c(AAS); ca. 25 m to N of fresh-

water pool at SW end of island, Smith 3014 (AAS, F).

EAST FALKLANDS. Lyall, syntype of Jungermannia

alternifolia (BM). Kidney Island: N of shanty (2797, 2801 —

c. per. -I- 3). Port William Region: Charles Point, 1954,

Pettingill 4 (CINC, F); N side of Gypsy Cove, sea level

(3253); ridge on Engineer Point Peninsula, 1 8-25 m (2814,

2816, 2824— c. per. + <3, 2835 -c. per. + $). Stanley

Region: Port Stanley, Government House, 1 5 m, Long-

ton & Smith, Longton 1032 (AAS); Port Stanley, 8 Nov.

1902, Skottsberg s.n. as L. cunninghamii (UPS— c. per.);

Moody Brook, 8 m, Longton & Smith, Longton 963

(AAS). Darwin Settlement: S side of Carcass Bay, Darwin

Harbour (2682-c. per. + $). WEST FALKLANDS. Hill

Cove Region: settlement, 15 m (3052— c. per. + <J, 3053—

c. per. + S); Hill Cove, Vallentin s.n. (MANCH). Roy Cove

Region: Vallentin 58 (MANCH). King George Bay Region:

Rabbitt Island, 21 Nov. 1907, Halle & Skottsberg 219,

syntype of Lophocolea monoica (UPS— c. per.).

Chiloscyphus sahuletorum (Hook. f. & Tayl.) En-

gel & Schust.

118

FIELDIANA: BOTANY

Jungermannia sabuletorum Hook. f. & Tayl., London

J. Bot. 3: 469. 1844. Lophocolea sabuletorum

(Hook. f. & Tayl.) G. L. & N., Syn. Hep. 697.

1 847. Chiloscyphus sabuletorum (Hook. f. & Tayl.)

Engel & Schust., Nova Hedwigia 39: 422. 1984.

Lectotype (/ate Engel, 1978): Falkland Is., Hooker

(FH!).

Jungermannia rivalis Hook. f. & Tayl., London J. Bot.

3: 469. 1844, syn. fide Engel (1978). Lophocolea

rivalis (Hook. f. & Tayl.) G. L. & N., Syn. Hep.

701. 1847. Lectotype (fide Engel, 1978): Falkland

Is., Port Louis, Hooker (NY!— c. $).

REMARKS— See Engel (1978).

ECOLOGY— This species is quite common in the

Falklands where it is present in a variety of hab-

itats. It occurs on stream banks and in stone runs

ofCortaderia heaths and on stream banks of dwarf

shrub heaths. Is is quite common in the Poafla-

bellata association where it grows on soil and grass

bases of nearly pure tussock stands, as well as in

rock crevices of coastal rocks at the margin of the

tussock community.

PHYTOGEOGRAPHY— Valdivian + Magellanian

-I- Falklands— Falkland Is.; Fuegia (Isla Grande

de Tierra del Fuego); southern Patagonian Chan-

nels (Brunswick Peninsula); Valdivian region;

Tristan da Cunha.

LITERATURE RECORDS (FALKLANDS)— Anony-

mous (Kiihnemann, 1937, 1949, as L. rivalis); Bir-

ger. Port Stanley (Stephani, 1905a, as L. rivalis);

Hooker (G. L. & N., 1847).

FALKLAND SPECIMENS SEEN -EAST FALKLANDS.

Hooker, syntype of Jungermannia sabuletorum (BM);

Lyall, as L. sabuletorum (FH); Hooker, syntype of Jun-

germannia rivalis (FH— c. 6); Lyall, as L. rivalis (FH— c.

6); Hooker, as L. grisea (BM— c. <J). Kidney Island: shore-

line, 16 Feb. 1954, Pettingill 11. 18, 20-22, 24, 47-48

(CINC, F); N of shanty (2781-2782, 2787, 2789); SE shore

between landing bay and Southeast Point (2804-2806,

2807 B, 2809, 2810-c. young per., 2811-2812). Port

William Region: Port William, Sept. 1850, Lechler as

Jungermannia rivalis (NY); N side of Gypsy Cove, sea

level (3250— c. young 9); ridge on Engineer Point Penin-

sula, 1 8-25 m (2575). Stanley Region: Port Stanley, Sept.

1 850, Lechler 100, as L. vasculosa (S); Port Stanley, 1 953,

Pettingill 35 (CINC, F); S shore of Port Stanley Harbour,

10 m, Taylor 203 (AAS, F); foot of Mt. William, 25 m,

Taylor 227b (AAS, F); Moody Brook, near Stanley, 8 m,

Longton & Smith, Longton 959, 962 (AAS, F); headwaters

of Mullet Creek Stream, ca. 60 m (3169, 3174, 3177).

Mt. Usborne Region: below The Gap, ca. 90 m (2589-

2590); valley SW of Mt. Usborne, ca. 60 m (2547 -c.

6, 2654, 2675-c. S). WEST FALKLANDS. Westpoint

Island: steep slope and cliffs facing The Woolly Gut

(2899-c. 3, 2903, 2904-c. <5, 2905-c. S, 2915-2916.

2936); near The Waterfall, 30-90 m (2557, 2557, 2567);

west base of Black Hog Hill, adjacent to Devils Nose,

ca. 18 m (2881-2884, 2886-2887, 2890); NE shore ad-

jacent to Cape Terrible (2879-2880). New Island: shore

of Ship Harbour, N of Ship Island (3291, 3294 -c. S).

Chiloscyphus semiteres (Lehm.) Mitt. Figure 55.

Jungermannia semiteres Lehm., Linnaea 4: 363. 1829.

Chiloscyphus semiteres (Lehm.) Lehm. & Lin-

denb. in G. L. & N., Syn. Hep. 190. 1845. Lopho-

colea semiteres (Lehm.) Mitt., J. Linn. Soc., Bot.

16: 188. 1877. Original material: South Africa,

Cape Prov., ". . . ostlichen Seite des Teufelberges

. . . ," Ecklon (W), cited in Grolle (1959a).

Lophocolea aequifolia Nees & Mont., Ann. Sci. Nat.

Bot. II, 5: 55. 1 836, syn. fide Grolle (1959a). Orig-

inal material: Juan Fernandez Is., Apr. 1 830, Ber-

tero (non vidi).

Chiloscyphus canaliculatusG. L. & N., Syn. Hep. 710.

1847, syn. cf. Mitten (1854). Jungermannia ca-

naliculata Hook. f. & Tayl., London J. Bot. 3:

563. 1 844, non Jungermannia canaliculata Nees,

Hep. Javanicae 10. 1830. Lophocolea canalicu-

lata (G. L. & N.) Steph., Bull. Herb. Boissier II,

6(9): 786. 1906 (= Spec. Hep. 3: 86). Original

material: New Zealand, Hooker (non vidi).

Lophocolea heterophylloides Nees in G. L. & N., Syn.

Hep. 157. 1 845, syn.fide Grolle (1959a). Original

material (fide G. L. & N., 1 845): "Habitat in Nova

Hollandia (Sieber in Hb. N. sub n. 54. fructifera

et sterilis sub n. 47.) ad flu men Cygnorum legit

Febr. 1 843 James Drummond (Hb. Taylor, Lg.

et G.) sterilem; ft in Chili in Hb. L. sub nomine

Lophocol. tumidae Ldbg. asservatur" (non vidi).

Chiloscyphus lindenbergianus Nees in G. L. & N., Syn.

Hep. 187. 1 845, syn.fide Grolle (1959a). Original

material: South Africa, Table Mt., Drege (w), cit-

ed in Grolle (1959a).

Lophocolea undulata Mont. Ann., Sci. Nat. Bot. Ill,

4: 351. 1845, syn fide Grolle (1959a). Original

material: "Chile australiori reperit cl. Gay. Herb.

Mus. Par." Gay (non vidi).

Lophocolea krauseana Steph., Bull. Herb. Boissier II,

6(8): 658. 1906 (= Spec. Hep. 3: 74), syn. fide

Grolle (1959a). Original material: Chile, Prov.

Valdivia, Corral, "typus in herb. Stephani," Krause

(non vidi).

Lophocolea dalliana Steph., Bull. Herb. Boissier II,

6(9): 785. 1906 (= Spec. Hep. 3: 85), syn. fide

Grolle (1959a). Original material: New Zealand,

without specific locality, "typus in herb. Ste-

phani," Dall (non vidi).

Lophocolea fernandeziensis Steph., Kongl. Svenska

Vetenskapsakad. Handl. 46(9): 44./ 16 f, g. 1 9 1 1 ,

syn.fide Grolle (1959a). Original material: Juan

Fernandez Is., Mas a Tierra, Valle Colonial,

Skottsberg (non vidi).

Lophocolea calbucensis Herz., Beih. Bot. Centralbl.

60(B): 6./ 3a-f. 1939, syn. fide Grolle (1959a).

Original material: Chile, Prov. Llanquihue, Cal-

buco, Schwabe (non vidi).

Lophocolea (?) magnistipula S. Amell, Rev. Bryol.

Lichenol. 23(1-2): 175. / 2a,b. 1954, syn.fide

Grolle (1959a). Original material: South Africa,

Cape Prov., Kasteel Poort, 1951, S. Arnell 1101

(UPS), cited in Grolle (1959a) (non vidi).

REMARKS— Chiloscyphus semiteres appears al-

lied to C. sabuletorum, and may be distinguished

from that species by underleaf features. Chiloscy-

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

119

120

FIELDIANA: BOTANY

phus semiteres has underleaves to 3 times the stem

width, with more ornamented margins and a ten-

dency to have larger laciniae, while C. sabuletorum

has underleaves 1-1.5 times the stem width, with

less ornamented margins and a tendency to pro-

duce laciniae or merely teeth.

ECOLOGY— This apparently is a very rare species

in the Falklands, where it was collected on soil

under rock overhang in a Hebe stand in a dwarf

shrub heath.

PH YTOGEOGRAPH Y — Pan-south-temperate in

Distribution— Falkland Is., Valdivian region; Juan

Fernandez Is.; "Frai Jorge"; Tristan da Cunha;

Inaccessible I.; Campbell and Stewart Islands; New

Zealand; Tasmania; Australia (widespread); Lord

Howe I. and South Africa. Also present in the cool,

oceanic Isles of Scilly (see Paton, 1965); Adelan-

thus lindenbergianus and Chiloscyphus bispinosus

are the only other Falkland taxa with an outpost

in the British Isles. The taxon seems to be restrict-

ed to rather warm temperate regions, especially in

the American sector (it is absent from the Magel-

lanian region entirely). In the Falklands, the species

is restricted to Westpoint Island, which is known

to have some of the highest temperatures in the

archipelago (see fig. 55). Fulford (1976) cites three

Pettingill collections as L. semiteres from Kidney

Is. (Falklands), which has an ecology quite differ-

ent from that of Westpoint Is. Pettingill 20 and 47

are actually C. sabuletorum; I have not seen no.

52.

LITERATURE RECORDS (FALKLANDS)— Skotts-

berg, Westpoint I. (Fulford, 1976).

FALKLAND SPECIMEN SEEN— WEST FALKLANDS.

Westpoint Island: steep slope and cliffs facing The Wool-

ly Gut (2930).

Chiloscyphus sylvaticus (Mitt.) Engel & Schust.

Figure 56.

Lophocolea sylvalica Mitt, in Thomson & Murray,

Rep. Sci. Results Challenger (Bot.) 1(3): 84. 1884.

Chiloscyphus sylvaticus (Mitt.) Engel & Schust.,

Nova Hedwigia 39: 424. 1984. Original material:

Juan Fernandez Is., Sounders (non vidi).

Lophocolea papulosa Steph., Kongl. Svenska Vetensk-

apsakad. Handl. 46(9): 50, / 18a,b. 1911, syn.

fide Fulford (1976). Original material: Juan Fer-

nandez Is., Mas a Tierra, Valle Colonial, Skotts-

berg (o, UPS), cited in Fulford (1976) (non vidi).

REMARKS— See comments in Engel (1978).

ECOLOGY— This species is occasionally encoun-

tered on stream banks in Cortaderia and dwarf

shrub heaths. It is rare on the bases of Blechnum

56

57

FIG. 56. Distribution of Chiloscyphus sylvaticus Mitt.

FIG. 57. Distribution of Chiloscyphus texlilis (Hook,

f. & Tayl.) Engel & Schust.

ENGEL: FALKLAND ISLANDS HEPATICAE AND ANTHOCEROTOPHYTA

121

magellanicum in the latter association and on cliffs

in Hebe stands of dwarf shrub heaths.

PHYTOGEOGRAPHY— Valdivian + Magellanian

+ Falklands — Falkland Is., southern Patagonian

Channels (Brunswick Peninsula); the Valdivian re-

gion (West Patagonian zone); and Juan Fernandez

Is. (see fig. 56).

FALKLAND SPECIMENS SEEN— without specific locality,

Capt. Abbott as L. humifusa (BM). Stanley Region: head-

waters of Mullet Creek Stream, ca. 60 m (3173— c. 6,

3 18 IB). Mt. Usborne Region, valley SW of Mt. Usborne,

ca. 60 m (2650, 26598-c. very young 2, 2660). West

Falklands. Westpoint Island: near The Waterfall, 30-90

m (2845). Weddell Island: Waterfall Valley, W of set-

tlement, ca. 125 m (3308— c. $, 3309— c. $, 3310, 3325).

Chiloscyphus textilis (Hook. f. & Tayl.) Engel &

Schust. Figure 57.

Jungermannia textilis Hook. f. & Tayl., London J.

Bot. 3: 468. 1844. Lophocolea textilis (Hook. f.

& Tayl.) G. L. & N., Syn. Hep. 696. 1847. Chi-

loscyphus textilis (Hook. f. & Tayl.) Engel &

Schust., Nova Hedwigia 39: 424. 1984. Lectotype

(fide Engel, 1978): Falkland Is., Hooker (FH!-C.

«).

Lophocolea campanulata Steph., Bih. Kongl. Svenska

Vetenskapsakad. Handl. 26 (III, 6): 34. 1900, syn.

fide Fulford (1976). Original material: Chile, Prov.

Aisen, R. Aisen Valley, ca. 200 m, 19 Jan. 1897,

Dusen 260 (G!— c. per., without specific locality,

NY!— c. per., UPS!— c. per.).

REMARKS— Lophocolea campanulata Steph. and

Chiloscyphus campanulatus Steph. are not based

upon the same type as stated by Fulford (1976, p.

469). Stephani (Spec. Hep. 3: 208) erroneously

listed "Chile (Dusen)" as the sole provenance and

collector of C. campanulatus. The species is ac-

tually based on a Himalayan plant (cf. Bonner,

1962-63 and Stephani, Icones Hep. Chiloscyphus

no. 63).

PHYTOGEOGRAPHY— Valdivian + Magellanian

+ Falklands— Falkland Is., Tierra del Fuego; Pat-

agonian Channels; Valdivian Region (West Pata-

gonia reportedly north to 43°57'S, Andean Pata-

gonia in P. N. Nahuel Huapi); and Juan Fernandez

Is. There have been no reports of the species in

West Patagonia north of Isla Guaitecas. I regard