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Lichens

Chapter 1: LICHENS
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The work surveys the development of lichen study, then analyzes the thallus and its composite nature, detailing algal gonidia, fungal hyphae, and their physiological interactions. It classifies and describes thallus morphologies—crustaceous, squamulose, foliose, fruticose, radiate, and stratose-radiate—examining cortical, gonidial, and medullary tissues and attachment structures. Special lichen features such as soredia, isidia, cephalodia, cyphellae, and breathing pores are defined and their formation and dispersal discussed. Reproductive structures and life-cycles are treated, including apothecia, perithecia, ascospore development, and asexual propagation. A glossary, bibliography, and numerous figures support identification, cultivation experiments, and taxonomic considerations.

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Title: Lichens

Author: Annie Lorrain Smith

Release date: March 14, 2020 [eBook #61616]
Most recently updated: October 17, 2024

Language: English

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*** START OF THE PROJECT GUTENBERG EBOOK LICHENS ***

Cambridge Botanical Handbooks

Edited by A. C. Seward and A. G. Tansley

LICHENS


CAMBRIDGE UNIVERSITY PRESS
C. F. CLAY, Manager
LONDON: FETTER LANE, E.C.4

LONDON: H. K. LEWIS AND CO., Ltd.,
136, Gower Street, W.C.1

LONDON: WHELDON & WESLEY, Ltd.,
28, Essex Street, Strand, W.C.2

NEW YORK: THE MACMILLAN CO.

BOMBAY } MACMILLAN AND CO., Ltd.
CALCUTTA }
MADRAS }

TORONTO: THE MACMILLAN CO. OF CANADA, Ltd.

TOKYO: MARUZEN-KABUSHIKI-KAISHA

ALL RIGHTS RESERVED


LICHENS

BY
ANNIE LORRAIN SMITH, F.L.S.
ACTING ASSISTANT, BOTANICAL DEPARTMENT, BRITISH MUSEUM

CAMBRIDGE:
AT THE UNIVERSITY PRESS
1921

PRINTED IN GREAT BRITAIN


PREFACE

The publication of this volume has been delayed owing to war conditions, but the delay is the less to be regretted in that it has allowed the inclusion of recent work on the subject. Much of the subject-matter is of common knowledge to lichenologists, but in the co-ordination and arrangement of the facts the original papers are cited throughout. The method has somewhat burdened the pages with citations, but it is hoped that, as a book of reference, its value has been enhanced thereby. The Glossary includes terms used in lichenology, or those with a special lichenological meaning. The Bibliography refers only to works consulted in the preparation of this volume. To save space, etc., the titles of books and papers quoted in the text are generally translated and curtailed: full citations will be found in the Bibliography. Subject-matter has been omitted from the index: references of importance will be found in the Table of Contents or in the Glossary.

I would record my thanks to those who have generously helped me during the preparation of the volume: to Lady Muriel Percy for taking notes of spore production, and to Dr Cavers for the loan of reprints. Prof. Potter and Dr Somerville Hastings placed at my disposal their photographs of the living plants. Free use has been made of published text-figures which are duly acknowledged.

I have throughout had the inestimable advantage of being able to consult freely the library and herbarium of the British Museum, and have thus been able to verify references to plants as well as to literature. A special debt of gratitude is due to my colleagues Mr Gepp and Mr Ramsbottom for their unfailing assistance and advice.

A. L. S.

London, February, 1920


CONTENTS

PAGE
Glossary xix
Errata xxii
Introduction xxiii
CHAPTER I
HISTORY OF LICHENOLOGY
A. Introductory 1
B. Period I. Previous to 1694 2
C. Period II. 1694-1729 5
D. Period III. 1729-1780 6
E. Period IV. 1780-1803 9
F. Period V. 1803-1846 10
G. Period VI. 1846-1867 15
H. Period VII. 1867 and after 18
CHAPTER II
CONSTITUENTS OF THE LICHEN THALLUS
I. LICHEN GONIDIA
1. GONIDIA IN RELATION TO THE THALLUS
A. Historical account of Lichen Gonidia 21
B. Gonidia contrasted with Algae 22
C. Culture Experiments with the Lichen Thallus 24
D. Theories as to the Origin of Gonidia 25
E. Microgonidia 26
F. Composite Nature of Thallus 27
G. Synthetic Cultures 27
H. Hymenial Gonidia 30
I. Nature of Association between Alga and Fungus 31
a. Consortium and symbiosis
b. Different forms of association
J. Recent views on Symbiosis and Parasitism 36
2. PHYSIOLOGY OF THE SYMBIONTS
A. Nutrition of Lichen Algae 39
a. Character of algal cells
b. Supply of nitrogen
c. Effect on the alga
d. Supply of carbon
e. Nutrition within the symbiotic plant
f. Affinities of lichen gonidia
B. Nutrition of Lichen Fungi 44
C. Symbiosis of other Plants 45
II. LICHEN HYPHAE
A. Origin of Hyphae 46
B. Development of Lichenoid Hyphae 47
C. Culture of Hyphae Without Gonidia 49
D. Continuity of Protoplasm in Hyphal Cells 51
III. LICHEN ALGAE
A. Types of Algae 51
a. Myxophyceae associated with Phycolichens
b. Chlorophyceae associated with Archilichens
B. Changes induced in the Alga 60
a. Myxophyceae
b. Chlorophyceae
C. Constancy of Algal Constituents 63
D. Displacement of Algae Within the Thallus 64
a. Normal displacement
b. Local displacement
E. Non-gonidial Organisms associated with Lichen Hyphae 65
F. Parasitism of Algae on Lichens 65
CHAPTER III
MORPHOLOGY
I. GENERAL ACCOUNT OF LICHEN STRUCTURE
ORIGIN OF LICHEN STRUCTURES
A. Forms of Cell-Structure 67
B. Types of Thallus 68
a. Endogenous thallus
b. Exogenous thallus
II. STRATOSE THALLUS
1. CRUSTACEOUS LICHENS
A. General Structure 70
B. Saxicolous Lichens 70
a. Epilithic lichens
aa. Hypothallus or protothallus
bb. Formation of crustaceous tissues
cc. Formation of areolae
b. Endolithic lichens
c. Chemical nature of the substratum
C. Corticolous Lichens 77
a. Epiphloeodal lichens
b. Hypophloeodal lichens
2. SQUAMULOSE LICHENS
A. Development of the Squamule 79
B. Tissues of Squamulose Thallus 81
3. FOLIOSE LICHENS
A. Development of foliose Thallus 82
B. Cortical Tissues 82
a. Types of cortical structure
b. Origin of variation in cortical structure
c. Loss and renewal of cortex
d. Cortical hairs
C. Gonidial Tissues 87
D. Medulla and Lower Cortex 88
a. Medulla
b. Lower cortex
c. Hypothallic structures
E. Structures for Protection and Attachment 91
a. Cilia
b. Rhizinae
c. Haptera
F. Strengthening Tissues of Stratose Lichens 95
a. Produced by development of cortex
b. Produced by development of veins or nerves
III. RADIATE THALLUS
1. CHARACTERS OF RADIATE THALLUS
2. INTERMEDIATE TYPES OF THALLUS
3. FRUTICOSE AND FILAMENTOUS THALLUS
A. General Structure of Thallus 101
Cortical Structures
a. The fastigiate cortex
b. The fibrous cortex
B. Special strengthening Structures 103
a. Sclerotic strands
b. Chondroid axis
C. Survey of Mechanical Tissues 105
D. Reticulate Fronds 106
E. Rooting Base in Fruticose Lichens 108
IV. STRATOSE-RADIATE THALLUS
1. STRATOSE OR PRIMARY THALLUS
A. General Characteristics 111
B. Tissues of Primary Thallus 112
a. Cortical tissue
b. Gonidial tissue
c. Medullary tissue
d. Soredia
2. RADIATE OR SECONDARY THALLUS
A. Origin of the Podetium 114
B. Structure of the Podetium 114
a. General structure
b. Gonidial tissue
c. Cortical tissue
d. Soredia
C. Development of the Scyphus 117
a. From abortive apothecia
b. From polytomous branching
c. From arrested growth
d. Gonidia of the scyphus
e. Species without scyphi
D. Branching of the Podetium 119
E. Perforations and Reticulation of the Podetium 120
F. Rooting Structures of Cladoniae 121
G. Haptera 122
H. Morphology of the Podetium 122
I. Pilophorus and Stereocaulon 125
V. STRUCTURES PECULIAR TO LICHENS
1. AERATION STRUCTURES
A. Cyphellae and Pseudocyphellae 126
a. Historical
b. Development of cyphellae
c. Pseudocyphellae
d. Occurrence and distribution
B. Breathing-Pores 129
a. Definite breathing-pores
b. Other openings in the thallus
C. General Aeration of the Thallus 132
2. CEPHALODIA
A. Historical and Descriptive 133
B. Classification 135
I. CEPHALODIA VERA
II. PSEUDOCEPHALODIA
C. Algae that form Cephalodia 136
D. Development of Cephalodia 137
a. Ectotrophic
b. Endotrophic
c. Pseudocephalodia
E. Autosymbiotic Cephalodia 140
3. SOREDIA
A. Structure and Origin of Soredia 141
a. Scattered soredia
b. Isidial soredia
c. Soredia as buds
B. Soralia 144
a. Form and occurrence of soralia
b. Position of soraliferous lobes
c. Deep-seated soralia
C. Dispersal and Germination of Soredia 147
D. Evolution of Soredia 148
4. ISIDIA
A. Form and Structure of Isidia 149
B. Origin and Function of Isidia 151
VI. HYMENOLICHENS
A. Affinity with other Plants 152
B. Structure of Thallus 153
C. Sporiferous Tissues 154
CHAPTER IV
REPRODUCTION
I. REPRODUCTION BY ASCOSPORES
A. Historical Survey 155
B. Forms of Reproductive Organs 156
a. Apothecia
b. Perithecia
C. Development of Reproductive Organs 159
1. DISCOLICHENS
a. Carpogonia of gelatinous lichens
b. Carpogonia of non-gelatinous lichens
c. General summary
d. Hypothecium and paraphyses
e. Variations in apothecial development
aa. Parmeliae
bb. Pertusariae
cc. Graphideae
dd. Cladoniae
2. PYRENOLICHENS
a. Development of the perithecium
b. Formation of carpogonia
D. Apogamous Reproduction 174
E. Discussion of Lichen Reproduction 177
a. The Trichogyne
b. The Ascogonium
F. Final Stages of Apothecial Development 181
a. Open or closed apothecia
b. Emergence of ascocarp
G. Lichen Asci and Spores 184
a. Historical
b. Development of the ascus
c. Development of the spores
d. Spore germination
e. Multinucleate spores
f. Polaribilocular spores
II. SECONDARY SPORES
A. Reproduction by Oidia 189
B. Reproduction by Conidia 190
a. Rare instances of conidial formation
b. Comparison with Hyphomycetes
C. Campylidium and Orthidium 191
III. SPERMOGONIA OR PYCNIDIA
A. Historical Account of Spermogonia 192
B. Spermogonia as Male Organs 193
C. Occurrence and Distribution 193
a. Relation to thallus and apothecia
b. Form and size
c. Colour
D. Structure 196
a. Origin and growth
b. Form and types of spermatiophores
c. Periphyses and sterile filaments
E. Spermatia or Pycnidiospores 201
a. Origin and form
b. Size and structure
c. Germination
d. Variation in pycnidia
F. Pycnidia with Macrospores 204
G. General Survey 205
a. Sexual or asexual
b. Comparison with fungi
c. Influence of symbiosis
d. Value in diagnosis
CHAPTER V
PHYSIOLOGY
I. CELLS AND CELL PRODUCTS
A. Cell-membranes 209
a. Chitin
b. Lichenin and allied carbohydrates
c. Cellulose
B. Contents and Products of the Fungal Cells 213
a. Cell-substances
b. Calcium Oxalate
c. Importance of calcium oxalate
C. Oil-cells 215
a. Oil-cells of endolithic lichens
b. Oil-cells of epilithic lichens
c. Significance of oil-formation
D. Lichen-acids 221
a. Historical
b. Occurrence and examination of acids
c. Character of acids
d. Causes of variation in quantity and quality
e. Distribution of acids
E. Chemical grouping of acids 225
I. ACIDS OF THE FAT SERIES
II. ACIDS OF THE BENZOLE SERIES
Subseries I. Orcine derivatives
Subseries II. Anthracene derivatives
F. Chemical Reagents as Tests for Lichens 228
G. Chemical Reactions in Nature 229
II. GENERAL NUTRITION
A. Absorption of Water 229
a. Gelatinous lichens
b. Crustaceous lichens
c. Foliose lichens
d. Fruticose lichens
B. Storage of Water 232
C. Supply of Inorganic Food 232
a. In foliose and fruticose lichens
b. In crustaceous lichens
D. Supply of Organic Food 235
a. From the substratum
b. From other lichens
c. From other vegetation
III. ASSIMILATION AND RESPIRATION
A. Influence of Temperature 238
a. High temperature
b. Low temperature
B. Influence of Moisture 239
a. On vital functions
b. On general development
IV. ILLUMINATION OF LICHENS
A. Effect of Light on the Thallus 240
a. Sun lichens
b. Colour-changes due to light
c. Shade lichens
d. Varying shade conditions
B. Effect of Light on Reproductive Organs 244
a. Position and orientation of fruits with regard to light
b. Influence of light on colour of fruits
V. COLOUR OF LICHENS
A. Origin of Lichen-Colouring 245
a. Colour given by the algal constituent
b. Colour due to lichen-acids
c. Colour due to amorphous substances
d. Enumeration of amorphous pigments
e. Colour due to infiltration
CHAPTER VI
BIONOMICS
A. Growth and Duration of Lichens 252
B. Season of Fruit Formation 255
C. Dispersal and Increase 256
a. Dispersal of crustaceous lichens
b. Dispersal of foliose lichens
c. Dispersal of fruticose lichens
D. Erratic Lichens 258
E. Parasitism 260
a. General statement
b. Antagonistic symbiosis
c. Parasymbiosis
d. Parasymbiosis of fungi
e. Fungi parasitic on lichens
f. Mycetozoa parasitic on lichens
F. Diseases of Lichens 268
a. Caused by parasitism
b. Caused by crowding
c. Caused by adverse conditions
G. Harmful Effect of Lichens 269
H. Gall-Formation 270
CHAPTER VII
PHYLOGENY
I. GENERAL STATEMENT
A. Origin of Lichens 272
B. Algal Ancestors 273
C. Fungal Ancestors 273
a. Basidiolichens
b. Ascolichens
II. THE REPRODUCTIVE ORGANS
A. Theories of Descent in Ascolichens 273
B. Relation of Lichens to Fungi 275
a. Pyrenocarpineae
b. Coniocarpineae
c. Graphidineae
d. Cyclocarpineae
III. THE THALLUS
A. General Outline of Development of Thallus 281
a. Preliminary considerations
b. Course of evolution in Hymenolichens
c. Course of evolution in Ascolichens
B. Comparative Antiquity of Algal Symbionts 282
C. Evolution of Phycolichens 283
a. Gloeolichens
b. Ephebaceae and Collemaceae
c. Pyrenidiaceae
d. Heppiaceae and Pannariaceae
e. Peltigeraceae and Stictaceae
D. Evolution of Archilichens 287
a. Thallus of Pyrenocarpineae
b. Thallus of Coniocarpineae
c. Thallus of Graphidineae
d. Thallus of Cyclocarpineae
AA. LECIDEALES
aa. Coenogoniaceae
bb. Lecideaceae and Gyrophoraceae
cc. Cladoniaceae
1. Origin of Cladonia
2. Evolution of the primary thallus
3. Evolution of the secondary thallus
4. Course of podetial development
5. Variation in Cladonia
6. Causes of variation
7. Podetial development and spore-dissemination
8. Pilophorus, Stereocaulon and Argopsis
BB. LECANORALES
aa. Course of Development
bb. Lecanoraceae
cc. Parmeliaceae
dd. Usneaceae
ee. Physciaceae
CHAPTER VIII
SYSTEMATIC
I. CLASSIFICATION
A. Work of Successive Systematists 304
a. Dillenius and Linnaeus
b. Acharius
c. Schaerer
d. Massalongo and Koerber
e. Nylander
f. Müller-Argau
g. Reinke
h. Zahlbruckner
B. Families and Genera of Ascolichens 311
C. Hymenolichens 342
II. NUMBER AND DISTRIBUTION
1. ESTIMATES OF NUMBER
2. GEOGRAPHICAL DISTRIBUTION
A. General Survey 343
B. Lichens of Polar Regions 345
C. Lichens of the Temperate Zones 348
D. Lichens of Tropical Regions 352
III. FOSSIL LICHENS
CHAPTER IX
ECOLOGY
A. General Introduction 356
B. External Influences 357
a. Temperature
b. Humidity
c. Wind
d. Human Agency
C. Lichen Communities 362
1. ARBOREAL 363
a. Epiphyllous
b. Corticolous
c. Lignicolous
2. TERRICOLOUS 367
a. On calcareous soil
b. On siliceous soil
c. On bricks
d. On humus
e. On peaty soil
f. On mosses
g. On fungi
3. SAXICOLOUS 371
a. Characters of mineral substrata
b. Colonization on rocks
c. Calcicolous
d. Silicicolous
4. OMNICOLOUS LICHENS 376
5. LOCALIZED COMMUNITIES 378
a. Maritime lichens
b. Sand-dune lichens
c. Mountain lichens
d. Tundra lichens
e. Desert lichens
f. Aquatic lichens
D. Lichens as Pioneers 392
a. Soil-formers
b. Outposts of vegetation
CHAPTER X
ECONOMIC AND TECHNICAL
A. Lichens as Food 395
a. Food for insects
b. Insect mimicry of lichens
c. Food for the higher animals
d. Food for man
B. Lichens as Medicine 405
a. Ancient remedies
b. Doctrine of “signatures”
c. Cure for hydrophobia
d. Popular remedies
C. Lichens as Poisons 410
D. Lichens used in Tanning, Brewing and Distilling 411
E. Dyeing Properties of Lichens 411
a. Lichens as dye-plants
b. The orchil lichen, Roccella
c. Purple dyes: orchil, cudbear and litmus
d. Other orchil lichens
e. Preparation of orchil
f. Brown and yellow dyes
g. Collecting of dye-lichens
h. Lichen colours and spectrum characters
F. Lichens in Perfumery 418
a. Lichens as perfumes
b. Lichens as hair-powder
G. Some minor Uses of Lichens 420
Appendix 421
Addendum 422
Bibliography 423
Index 448

GLOSSARY