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to the General Library
University of California, Berkeley

Libris

BEATRIX
JONES

LANDSCAPE
ARCHITECTURE

REEF POINT GARDENS
LIBRARY

PLANT LIFE

ALPINE SWITZERLAND

A Typical Damp Pasture Association. The Globe Flower ( Trollius europceus,
Linn.), the Aconite-leaved Buttercup (Ranunculus aconitifolius, Linn.),
and the White Veratrum ( Veratrum album. Linn.).

[Frontispiece.

PLANT LIFE IN
ALPINE SWITZERLAND

BEING AN ACCOUNT IN SIMPLE LANGUAGE

OF THE NATURAL HISTORY OF ALPINE

PLANTS

BY E. A. NEWELL AEBER

M.A., F.L.S., F.G.S.

TRINITY COLLEGE, CAMBRIDGE

UNIVERSITY DEMONSTRATOR IN PALJJOBOTANY

ILLUSTRATED BY 48 PLATES OF PHOTOGRAPHS
FROM NATURE, AND 80 FIGURES IN THE TEXT

LONDON

JOHN MURRAY, ALBEMARLE STREET, W.

1910

E. A. NEWELL ABBEB

ldd'1

tANDSCAPE

ARCHITECTURE

A. A.

AND

D. V. S.

a<f> olv TT OLVT a

517

PREFACE

A FEW words may be necessary to explain the
scope and intention of the present volume.

It has often been remarked, and with some
measure of truth, that botanists, in common with
other scientists, have in the past done little to bring
the results of their researches within the reach of
the layman, in a language in which they can be
commonly understood and appreciated. The present
volume attempts, no doubt imperfectly, the difficult
task of trying to draw attention to some points of
botanical interest among the better-known members
of the Alpine flora of Switzerland.

As the result of repeated visits to the Alps,
now extending over a period of more than twenty
years, I have been struck on many occasions by
the inadequate information, at present available, for
those who are interested in the habits of Alpine
plants, but have not had the good fortune to have
been able to devote much time previously to a
serious study of Botany. Again and again I have

vii

viii PREFACE

watched the natural sequence of events. The
tourist, arriving among the High Alps perhaps for
the first time, is at once struck by the extreme beauty
and richness of the flora. I have noticed that, in
nine cases out of ten, he will devote more attention to
the Alpine flowers than he is at all likely to bestow on
our British wild plants. He is anxious, and rightly
so, to ascertain the name of this or that plant, which
he may happen to have come across, usually quite unex-
pectedly, in his wanderings and excursions. Yet, to
determine the genus and species is often an extremely
difficult task for the layman, and in certain cases
even for a trained botanist. If, rather by good luck
than otherwise, perhaps by resort to a book contain-
ing crudely coloured illustrations of some of the
commoner Swiss plants, the name of a flower is
obtained, as a rule all further interest in the plant
ceases. It is either thrown away or perhaps
"pressed," and, as often as not, forgotten.

Yet, to the trained botanist, the name of a plant
is frequently the least interesting matter in connection
with it. He too, like the layman, may have to take
some pains to find out the genus and species, but,
once these have been ascertained, a whole host of
fascinating, even absorbing, interests may be pre-
sented by almost any Alpine plant. Originally a
knowledge of each of these was won from the
domain of ignorance by the researches of some
botanical student of Alpine vegetation. The results
of his scientific explorations are buried in a host of

PREFACE ix

scattered, sometimes obscure, and inaccessible
scientific periodicals, or in the transactions of
learned societies, and may have been published in
almost any one of the European languages. This
" literature," as it is called, is naturally beyond the
ken of the layman, whereas the trained botanist is
familiar with, and is constantly brought into contact
with, these memoirs, and is an expert in the use of
such means as exist, whereby one can reach all that
is known, or has been written, on a particular subject.
Thus the botanist is at a great advantage, an
advantage which has always appeared to me to be
somewhat unfair.

It should here be stated clearly that this work is
not intended to give any aid towards ascertaining the
names of Alpine plants. Even the photographs
which illustrate it are not published for that purpose.
To determine a species or genus, the reader must
make use of some one or other of the systematic
floras of Switzerland, the more useful of which are
indicated in Appendix III. In this volume I propose,
rather, to attempt to explain, in simple language,
some of the features presented by Alpine plants
which appear to me to be of special interest. So
far as possible, technical terms will be avoided.
It will be assumed, however, that the reader is
familiar with the ordinary features of a plant,
especially of the flower. A summary of this sub-
ject will be found in Appendix II. ; whereas the
first Appendix contains a list of the commoner

x PREFACE

technical terms applied to the Higher Plants, including
those used or mentioned in this volume.

In the preparation of this volume I have, in
addition to new and original observations, drawn
freely on many sources of information, which are too
numerous to permit of specific mention. I would,
however, record my indebtedness to the works
of Kerner, Christ, and Schroeter, mentioned in
Appendix III. ; to various scientific memoirs by
Professor Bonnier and other members of the French
School of Experimental Alpine Cultivation ; and to
those of the Swiss School of Ecology, of which
Professor Schroeter of Zurich is the head.

I am also indebted to many friends for information,
advice, and criticism. To my wife I owe many
thanks for the drawings of the text-figures, which are
original, except where the contrary is stated in each
case, and also for much help in many other directions,
including the reading of the proof-sheets. To my
friend Mr John Parkin, M.A., of Trinity College, I
would express my thanks for much information, and
for the value of his co-operation in field work in
Switzerland.

The photographs, which illustrate the letterpress,
were all taken in Switzerland during the past four
years, expressly for this work. In a majority of
cases the plants were photographed in situ, and the
prints and negatives are "untouched." For the
larger number I must plead responsibility, but I am
greatly indebted to my friend Major George Dixon

PREFACE xi

(of the 5th Border Regiment), M.A., Trinity College,
and to Miss Gertrude Bacon, for many interesting
negatives, and for the enthusiasm with which they
have assisted me in this direction. The source
of each photograph is indicated in the List of
Illustrations.

E. A. NEWELL ARBER.

TRINITY COLLEGE, CAMBRIDGE,
3rd June 1910.

CONTENTS

CHAPTER I
INTRODUCTORY— Swiss NATIONAL FLOWBKS : THE EDELWEISS

AND THE ALPENROSES . * . . , » . l

CHAPTER II

TYPICAL FLOWERS OF THE ALPINE PASTURES— THE ANEMONES

AND GENTIANS . . . . * . 28

CHAPTER III
TYPICAL FLOWERS OF THE ALPINE PASTURES (continued}— THE

SOLDANELLAS, THE PRIMULAS, THE ANDROSACES, THE

SAXIFRAGES, THE CAMPANULAS, AND THE RAMPIONS . 61

CHAPTER IV
ROCK PLANTS OF THE PASTURES . , * . . 93

CHAPTER V
INTERESTING PLANTS OF THE ALPINE PASTURES . . .116

CHAPTER VI

PLANTS OF THE ALPINE MEADOWS . -.-;. * . . 142

CHAPTER VII
THE HIGH ALPINE PLANTS . . * . . .169

xiii

xiv CONTENTS

CHAPTER VIII PAGE

THE HIGH ALPINE PLANTS (continued)— MAESH PLANTS . 199

CHAPTER IX
THE ALPINE THICKETS AND FORESTS . . . . 220

CHAPTER X
THE SHADE PLANTS OF THE ALPINE FORESTS *-.?-, £ >«•. ;> 246

CHAPTER XI
ADAPTATIONS AMONG ALPINE PLANTS 8 -J . . .268

CHAPTER XII

THE GEOGRAPHICAL DISTRIBUTION, AFFINITIES, AND ORIGIN OF

THE Swiss ALPINE FLORA . '>* . . . 294

APPENDICES

I. GLOSSARY OF BOTANICAL TERMS . . * . 307

II. THE STRUCTURE OF THE FLOWER . . '. . 322

III. BOOKS ON THE Swiss ALPINE FLORA . 333

INDEX ........ 337

LIST OF ILLUSTRATIONS

PLATES

A Typical Damp Pasture Association. The Globe
Flower (Trollius europceus, Linn.), the Aconite-leaved
Buttercup (Ranunculus aconitifolius, Linn.), and
the White Veratrum ( Veratrum album, Linn.) . Frontispiece

[E.A.N.A., photo.]
PLATE

'Fig. 1. The Edelweiss (Leontopodium alpinum,\
Cass.)

I. -{ (Major Dixon, photo.] V Face

Fig. 2. The Alpine Rose (Rosa alpina, Linn.)

[Major Dixon, photo.]}

PAGE

II. The Common Alpenrose (Rhododendron ferru-
gineum. Linn.) .....

[Major Dixon, photo.]

III. A Typical Alp or Mountain Pasture, with the Stone

Pine (Pinus cembra, Linn.) in the Foreground
(6,300 feet) . . . .

[Major Dixon, photo.]

IV. The Spring Anemone (Anemone vernalis, Linn.)

[E.A.N.A., photo.]

'Fig. 1. Mature Fruits of the Spring Anemone'
(Anemone vernalis, Linn.) . .

[E.A.N.A., photo.]

Fig. 2. Mature Fruits of the Alpine Anemone
(Anemone alpina, Linn.) . *

[Major Dixon, photo.]j

VI. A Group of Yellow Alpine Anemones (Anemone
alpina. Linn., var. sulphurea, Linn.)

[E.A.N.A., photo.]

VIII.

xvi LIST OF ILLUSTRATIONS

PLATE
VII. The Alpine Anemone (Anemone alpina. Linn.)

Fig. 1. Young Flowers of the White Variety
growing out of the Involucre. [E.A.N.A., photo.]
Fig. 2. Male Flowers of the Yellow Variety.

[Major Dixon, photo.]

Fig. 1. Young Fruits of the Alpine Anemone>
(Anemone alpina, Linn.)

[E.A.N.A.,photo.]

Fig. 2. The Narcissus-flowered Anemone (Ane-
mone narcissi/lora, Linn.)

[Major Dixon, photo.] J

IX. A Plant of the Common Bell Gentian (Gentiana
acaulis, Linn.), the Corolla cut open to show the
Stamens and the Ovary . . V

[E.A.N.A., photo.]

Fig. 1. The Spring Gentian (Gentiana verna,
Linn.) f . . . ,

[Miss Bacon, photo.]

Fig. 2. The Flowers of the Field Gentian
(Gentiana campestris, Linn.)

[Major Dixon, photo.]-)

XI. The Flowers of the Spotted Gentian (Gentiana
punctata, Linn.) ,

[E.A.N.A., photo.]

XII. The Alpine Soldanella (Sddanella alpina, Linn.)
Flowering in the Snow ,

[Mayor Dixon, photo.]

XIII. The Bird's-eye Primrose (Primula farinosa, Linn.) .

[E.A.N.A., photo.]

Fig. 1. The Auricula (Primula auricula, Linn.) .

[E.A.N.A., photo.]

XIV. •[ Fig. 2. The Dwarf Androsace (Androsace chamce-

Willd.) . . .

[Miss Bacon, photo.] j

XV. A Bank of the Rough Saxifrage (Saxi/raga aspera,
Linn.), with the Rock Catchfly (Silene rupestris,
Linn.), and a Rampion (Phyteuma, sp.).

[B.A.N.A., photo.]

PAGE

Face 40

PLATE

XVI. -

XVII. •

XVIII. •

LIST OF ILLUSTRATIONS

Fig. 1 . Leaf Rosettes of the Evergreen Saxif rage>
(Saxifraga aizoon, Jacq.)

[E.A.N.A., photo.]

Fig. 2. The Rock Catchfly (Silene rupestris,
Linn.) ......

[Major Dixon, photo.]'

rFig. 1. The Harebell (Campanula rotundifolia^
Linn.) . . \

[E.A.N.A.,photo.]

Fig. 2. The Mont Cenis Campanula (Cam-
panula cenisia, Linn.) . *

[Major Dixon, photo.]-

Fig. 1. The Spider's Web House-leek (Semper--
vivum arachnoideum. Linn.) .

[E.A.N.A.,photo.]

Fig. 2. The Bearded Campanula (Campanula
barbata. Linn.) .

• [E.A.N.A., photo.]

XIX. The Tufted Campanula (Campanula thyrsvidea,
Linn.) » * ^ ••*

[E.A.N.A., photo.]

XX.

Fig. 1. Leaves of the White Dryas (Dryas octo-
petalay Linn.) . » . . fc

[E.A.N.A., photo.]

Fig. 2. The Round-headed Rampion (Phyteuma
orbicular et Linn.) V . . .

[E.A.N.A.tphoto.]

XXI. The Rosettes of the Spider's Web House-leek
(Sempervivum arachn&ideum, Linn.)

[E.A.N.A., photo.]

XXII. The Migration of the Mountain House-leek
(Sempervivum montanum, Linn.)

[E.A.N.A., photo.]

XXni. The Migration of the Mountain House-leek
(Sempervivum montanum, Linn.)

[E.A.N.A., photo.]

xvii

PAOE

Face 76

100

102

XV111

PLATE

XXIV. <

LIST OF ILLUSTRATIONS

Fig. 1. Rosettes of the Mountain House-leek
(Sempervivum montanum, Linn.) ,

[Major Dixon, photo.]

Fig. 2. Rosette of a Sempervivum attacked by
a Parasitic Fungus ....

[E.A.N.A., photo.]''

XXV. A Plant of the White Dryas (Dryas octopetala,
Linn.) .

[Miss Bacon, photo.]

XXVI. A Typical Alpine Carpet Plant (Globularia
nudicaulis, Linn.) * , .

[E.A.N.A.,photo.]

XXVII. The White VQwimm (Veratrtim album, Linn.) .
Fig. 1. Leafy Shoots. IE.A.N.A., photo.]

Fig. 2. Inflorescence. [B.A.N.A.,photo.]

XXVIII. The White Veratrum (Veratrum album, Linn.) .
Fig. 1. Young Shoots in Spring beginning to

expand. [Major Dixon, photo.]

Fig. 2. Root-stock and Contractile Roots.

[E.A.N.A., photo.]

Fig. 1. The Leaves of a Lady's-mantle (Alche-
milla\ with Drops of Water exuding from
the Pores on the Margins . . ,

[E. A. N. A., photo.]

Fig. 2. The Carline Thistle (Carlina acaulis,
Linn.) in Bad Weather . .

[E.A.N.A.,photo.]

XXIX. -i

XXX. H

Fig. 1, The Alpine Buttercup (Ranunculus^
alpestrisy Linn.) ....

[E.A.N.A., photo.]

Fig. 2. Fruits of the Mountain Avens (Geum
montanum, Linn.) ....

[Major Dixon, photo.]

PAGE

Face 104

106

110

120

122

126

128

PLATE

XXXI.

XXXII. -i

LIST OF ILLUSTRATIONS

•Fig. 1. The Black Nigritella (Nigritella angusti-
folidy Kick.) .....

[Major Dixon, photo.]

hFig. 2. The Lesser Butterfly Orchid (Hdbenaria
Ufolia, R. Br.) .
{Major Dixon , photo.]'

Fig. 1. The Spring Crocus (Crocus vernus, All.)'
Flowering in the Snow

[Major Dixon, photo.]

Fig. 2. Section of an Alpine Meadow, showing
the Peat Soil.

[Major Dixon, photo.]-*

XIX

PAGE

Face 134

XXXIII.

XXXIV.

XXXV.

XXXVI,

Typical High Alpine Cushion Plants
Fig. 1. The Sessile-flowered Moss Campion
(Silene exscapa, All.) [Major Dixon, photo.]
Fig. 2. The Swiss Androsace (Androsace hel-
vetica. Gaud.) [Major Dixon, photo.]

High Alpine Cushion Plants . i 4

Fig. 1. Cushion of the Glacial Androsace
(Androsace glacialis, Hopp.)

[Miss Bacon, photo.]

Fig. 2. Cushion of the Dwarf Alsine (Alsine

Sedoidesy'FlC&l.) [Miss Bacon, photo.]

Fig. 1. The Eritrichium (Eritrichium nanum,
Schrad.)

[Miss Bacon, photo.]

Fig. 2. The Flowers and Bulbils of the Vivi-
parous Polygonum (Polygonum viviparum,
Linn.), » . •

[E.A.N.A., photo.])

Fig. 1. The Purple Saxifrage (Saxifraga^
oppositifolia, Linn.) ....

[Major Dixon, photo.]

Fig. 2. The Androsace-like Saxifrage (Saxi-
fraga androsacea, Linn.)

[Major Dixon, photo.]

154

180

182

184

186

LIST OF ILLUSTRATIONS

PLATE

PAGE

XXXVII. A High Alpine Carpet Plant, the Reticulate

Willow (Salix reticulata, Linn.) . . Face 188

[E.A.N.A.,photo.]

XXXVIII. 1

fFig. 1. Vital's Androsace (Androsace vitaliana,1
Lap.) ' .

[Miss Bacon, photo.]

Fig. 2. A Colony of the Common Butterwort
(Pinguicula vulgaris, Linn.) ^'

[Major Dixon, photo.]J

XXXIX. An Alpine Marsh of the Marsh Marigold (Galtha
palustrist Linn.) . 4 « '

[E.A.N.A.,photo.]

XL. A Damp Pasture covered with the Aconite-
leaved Buttercup (Ranunculw aconitifolius,
Linn.) . . . '••/'

[E.A.N.A.,photo.]

'Fig. 1. A Colony of the Alpine Butterwort'
(Pinguicula alpina, Linn.) .

[E.A.N.A.,photo.]

XLI.

Fig. 2. The Kosette of Leaves of the Common
Butterwort (Pinguicvla vulgaris. Linn.),
showing Flies and a Moth caught and
digested by the Leaves

[E.A.N.A., photo.]

'Fig. 1. The Cowberry (Vaccinium vitis-idcea,'
Linn.) . ....

[Major Dixon, photo.]

Figs. 2 and 3. The Flowers of the Alpine
Soldanella (Soldanella alpina, Linn.)

[Major Dixon, photo.]

Fig. 4. Two Flowers of the Blue Honeysuckle
(Lonicera ccerulea, Linn.) .

[E.A.N.A., photo.]}

XLIII. The Atragene (Clematis alpina, Miller) .

Fig. 1. The Flowers. Fig. 2. The Fruits.

[E.A.N.A.,photo.]

XLII.

198

210

212

222

230

PLATE

XLIV.

LIST OF ILLUSTRATIONS

Fig. 1. Rock Colonisation : A Bare Slab of"
Rock in a Forest showing a Primitive Soil
composed of Pine-needles .

[Major Dixon, photo.]

Fig. 2. The Lichen (Usnea) growing on the
branches of a Coniferous Tree

[E.A.N.A.,photo.]j

XLV. The Linnsea (Linncea borealis, Gronov.) .

XLVI. J

XLVII. 4

Fig. 1. The Box-leaved Polygala (Pdygala?
chamcebuxw, Linn.) . . . .

[E.A.N.A., photo.]

Fig. 2. The Two-flowered Violet (Viola biftora,
Linn.). . . . , *

[Major Dixon, photo.]J

Fig. 1. The Flowers of the Martagon Lily
(Lilium martagon^ Linn.) .

[E.A.N.A., photo.]

Fig. 2. The May Lily (Maianthemum convert-
laria, Weber) '*

[Major Dixon, photo.]-1

XXI
PAGE

Face 238

248

252

258

FIGURES IN THE TEXT

FIG. PAGE

I. The Leaf and Scales of the Common Alpenrose (Rhodo-
dendron ferrugineum, Linn.) . . . .26

1. Transverse section of a leaf, showing the scales on the

lower surface. Magnified 14 times.

2. Transverse section through a scale, on the right side of

which a stoma is seen sheltered by the scale.
Magnified 190 times.

3. Surface view of a scale. Magnified 190 times.

II. Fruits of the Alpine Anemone (Anemone alpina, Linn.), in
various stages of development. Magnified twice

1. In fully-opened flower. 2. Intermediate stage. 3. The
mature fruit.

xxii LIST OF ILLUSTRATIONS

FIG. PAGE

III. The Flower of the Field Gentian (Gentiana campestris,

Linn.), showing the scale-like outgrowths from the throat

of the corolla. Magnified 3 times . . . . 55

IV. 1 and 2. The Alpine Soldanella (Soldanella alpina, Linn.)

1. Longitudinal section of the flower.

2. Section across the flower on the level of the scales.

„ 3 and 4. The Small Soldanella (S. pusilla, Baumg.) . 65

3. Longitudinal section through the flower.

4. Section across the flower in the plane of the anthers.

(All after Schroeter.)

V. Diagrammatic Views of the Heterostyled Flowers of a

Primula § ,* , * * > , 68

1. Flower with short style and high stamens.

2. Flower with long style and low stamens.

VI. Section through a Chalk-gland on the edge of a Leaf of a

Saxifrage. Highly magnified . . . .76

VII. A Plant of the Hound-leaved Campanula (G. rotundifolia,
Linn.), grown in diffuse light. All the leaves are cordatej
like the seed-leaves. (After Goebel) *' » . 84

VIIL The Stamens and Carpels of Scheuchzer's Bell-flower
(Campanula Scheuchzeri, Vill.), illustrating the mech-
anism of the stylar brush. Enlarged . . .86

1. United anthers surrounding the style in the unopened

flower.

2. The anthers separating and curling backwards.

3. The empty anthers coiled in tight spirals, the style; with

the stylar brush, bearing pollen, and the three stigmas
beginning to unfold.

IX. A Fruit (Capsule) of Campanula. Enlarged . . 88

X. The Flowers of the Hound-headed Rampion (Phyteuma

orbiculare, Linn.), in various stages. Magnified . . 92

1. The united petals of a young flower, with the calyx below.

2. The petals separating in an older flower.

3. The petals further separated, and the style growing

through the tip of the tube formed by the petals.

4. Mature flower with free petals, showing the style with

stylar brush, and the three expanded stigmas.

LIST OF ILLUSTRATIONS xxiii

FIG. PAGE

XI. 1. The Flower of the Lesser Butterfly Orchid (ffabenaria

bifolia, Rothb.). Enlarged . . . .137

„ 2. A Pollinium. Much enlarged.

XII. The Pollinium of an Orchid as removed from the flower,
adhering by the disc to the end of a needle. Much
magnified ..,,... 139

1. The position immediately after removal from the

flower.

2. The position assumed shortly afterwards.

XIII. The Underground Stem or Corm of the Spring Crocus

(Crocus vernus, All.), in Spring. (The scale-leaves have
been removed.) . . . , . ..153

XIV. Stages in the development of the Fruit of the Water

Avens (Geum rivale, Linn.) . . . .158

XV. Section of a Flower of the Field Pansy (Viola tricolor,

Linn.) . . « , * ' * * 16°

XVI. Ripe Fruit of the Field Pansy (Viola tricolor, Linn.),
shooting out its seeds. The top valve has not yet
lost any seeds ; the left valve has lost one row ; the
right valve, two rows. Enlarged v . . 163

XVII. The Corm or Underground Stem of Colchicum, the

Meadow Saffron . • f , . . . 167

XVIII. Plants of the Harebell (Campanula rotundifolia, Linn.).
That on the left grown in the Lowlands ; that on the
right in an Alpine garden. (After Bonnier) . , 201

XIX. The Common Butterwort (Pinguicula vulgaris, Linn.) . 213

1. Transverse section of a leaf, showing the glands on

the upper surface. Somewhat enlarged.

2. A side view of a gland. \,r .

3. Asurfaceviewof a gland. /Much enlarged.

XX. A Staminode from a Flower of the Grass-of-Parnassus

(Parnassia palustris, Linn.). Much magnified . . 217

XXI. Stamen of a Bilberry (Vaccinium) . . . . 223

xxiv LIST OF ILLUSTRATIONS

FIG. PAGE

XXII. Two Flowers of the Mountain Honeysuckle (Lonicera
alpigena, Linn.), with Ovaries partially united.
(After Hermann Miiller) . . . * 226

XXIII. The Fruit of the Mountain Honeysuckle (Lonicera

alpigena, Linn.), formed by the complete union of the
two berries. The scars of the two calyx rings can
still be seen ...... 227

XXIV. A Flowering Branch of the Knot-foot (Streptopus

amplexifolius, D.C.), showing the flower-stalks
united to the axis for the length of the internode
above their insertion, and the flowers thus hanging
below the leaf next above that in the axil of which
they arise " » , » » 256

XXV. The Leaves of the Wood Sorrel (Oxcdis acetosella, Linn.) 264

1. In the day position.

2. In the sleep position.

XXVI. 1. The Flower of a Buttercup (natural order Ranun-
culaceae); 2. Petal of a Buttercup, showing the
Honey-gland at the base .... 328

XXVII. The Flower of a Pea (natural order Leguminosae) . 329

XXVIH. The Flower of a Saxifrage (natural order Saxi-

fragacese) . . . . . . , 329

XXIX. The Head of Flowers of the Groundsel, Senecio (natural

order Compositse). Enlarged . . . 330

XXX. The Flower of a Crocus. A typical Monocotyledonous

Flower (natural order Iridaceas). Reduced . # 331

PLANT LIFE IN
ALPINE SWITZERLAND

CHAPTER I

INTRODUCTORY — SWISS NATIONAL FLOWERS: THE
EDELWEISS AND THE ALPENROSES

NOT so very long ago — in fact, within the recollection
of many — the object of a botanical study of a region
such as Alpine Switzerland was merely to compile a
list of the genera and species l of plants found within
its limits. Such a catalogue, which the botanist terms
a flora, though a highly necessary preliminary stage
in the progress of our knowledge, is now no longer
regarded as "the be-all and the end-all" of such
enquiries. It is true that by its aid we learn the names
of the plants we meet with in our wanderings among
the Alps, and how to distinguish them from one
another. This being accomplished, we may compare
the Alpine flora of one country, such as Switzerland,
with that of another, Britain ; thus an extremely

1 gee footnote, p. 15.

2 INTRODUCTORY

interesting comparison may result. But we may seek
to know more of these Alpines than a "flora" can
tell us. We have come to recognise that all plants
are living beings like ourselves, face to face with
certain difficulties of existence, such as adverse climatic
conditions, and competition with their neighbours.
How are they adapted to meet these conditions,
and so to survive in the struggle for existence ?
How are we to explain in these terms the various
characteristics of Alpine plants, their similarities
and dissimilarities to one another?

It is this newer and wider view of the vegetation
of a district such as the Alps which has opened
up such an interesting field for study and reflection.
We pass beyond the limits of a flora, or catalogue
of species, to enquire how these plants can live under
conditions so dissimilar to those which prevail in
Britain, and with which we are more familiar. We
may well adopt the attitude of one of the earliest
of the great German botanists, Christian Konrad
Sprengel, who, in 1793, sought to unravel the origin,
meaning, and uses of the various features exhibited
by plants, starting with the hypothesis that the wise
Author of Nature had not created even a single hair
without a definite purpose,1

We shall adopt a somewhat similar standpoint

1 "Ueberzeugt, dass der weise Urheber der Natur auch nicht ein
einziges Harchen ohne eine gewisse Absicht hervorgebracht hat," C. K.
Sprengel, Das Entdeckte Geheimniss der Natur, in Bau und in der
Befruchtung der Blumen ; Berlin, 1793, p. 1.

THE ALPINE ZONE 3

here. We have still much to learn as regards the
life of Swiss Alpine plants. Many of their striking
characteristics cannot be satisfactorily explained at
present. However, on others we are now beginning
to get some light, as the result of scientific enquiries,
conducted for the most part within recent years.

In the present chapter we will endeavour to
illustrate these principles by reference to some well-
known Swiss Alpines. For the moment, however,
we may first pause to enquire : What is an Alpine
plant ? This is a question which it is not possible
to answer accurately. The term is an arbitrary one,
and, so far as Switzerland is concerned, an Alpine
plant is best defined as one flourishing within the Alpine
zone, another term which itself implies a still more
arbitrary distinction.

Switzerland, although a small country, varies
greatly as regards altitude. It is well known that as
we ascend towards the mountains from the lowlands
of northern Switzerland, the flora changes gradually.
We can recognise, as we pass upwards, at least three
zones, which we may call the Lowland, the Subalpine,
and the Alpine. The general character of the flora of
each zone is distinct, though the zones merge
gradually one into the other, and many plants may
occur in two, or even all three, zones. As a rule,
however, with several noteworthy exceptions, each
plant flourishes abundantly in one particular zone
alone, and is only feebly represented in, or even
entirely absent from, the others.

4 INTRODUCTORY

The Lowland zone includes the plains and the
small hills in the north and west of Switzerland.
Its flora is identical, for the most part, with that
of temperate Northern and Western Europe, includ-
ing Britain and much of France and Germany. In
some places, however, there is also a very distinct
intermixture of southern plants, derived from the
subtropical Mediterranean flora lying to the south
of the great mountain chain of Central Europe, of
which the Swiss Alps form only one link. Such an
intermixture even exists in Britain, for a few plants,
especially certain Heaths, also belong to the
Mediterranean flora, and there has been much
speculation as to how they managed to reach Britain.

In Switzerland, the Lowland zone is essentially
the zone of the Vine, and it extends upwards to an
altitude of 3,000 to 3,500 feet, above which grapes will
not, as a rule, ripen.

The next zone, the Subalpine, reaches to about
5,000 feet, the exact height varying locally according
to the physical conditions, such as exposure, situation,
etc. Beech forests are the great natural feature of
this zone. Many Lowland plants can still flourish
at this altitude, and members of the Alpine flora
creep downwards, so that the vegetation of the
Subalpine zone is largely a mixture of species found
also in the zone above, and in that below. In no
case is there any sudden change in the flora as we pass
from one zone to another — merely a gradual transi-
tion. A large number of Lowland plants become less

THE ALPINE ZONE 5

and less frequent as we pass upwards, and the
Alpine species, little by little, become more numerous
and important as characteristic features of the
flora.

A few species are confined to the Subalpine region,
just as some Lowland plants do not extend beyond
their particular zone; but these are not sufficiently
numerous to characterise the region in which they
occur.

The Alpine zone will here be regarded as beginning
roughly at a height of 5,000 feet. At this elevation
the Beech, as a rule, ceases to flourish, and coni-
ferous forests of Spruce, Pine, and Larch replace
it and become important landmarks as regards
scenery and vegetation. It is the plants growing
above this altitude which we shall term Alpine
plants, and with these alone we are concerned in this
volume.

As we have already indicated, all zonal limits are
purely arbitrary, owing to the exceedingly gradual
nature of the change in the vegetation as we ascend the
Alps. This is well illustrated by the fact that almost
all the botanists who have attempted to define the
Alpine zone within rigid limits, have arrived at totally
different conclusions. Thus a great Swiss authority,
Dr Christ, distinguishes the region above the limit of
the Pine and Larch forests as the Alpine zone — a
region which begins at more than 1,000 feet above
that here adopted. The late Mr John Ball, another
authority, divides the whole of Switzerland into two

6 INTRODUCTORY

zones : a lower, extending to the limit of deciduous1
trees ; and an upper, including all above this limit,
except a glacial region, where the soil is only free from
snow for two or three months in summer. This Glacial
or High Alpine region may well be distinguished as a
separate zone.

Prof. Schroeter has recently published a com-
parative table showing how no less than twenty-five
botanists, between 1808 and 1904, have attempted to
subdivide Swiss vegetation in regard to altitude. No
two schemes agree even remotely ; and this diversity of
opinion well illustrates the impossibility of attempting
to define zones of altitude at all rigidly in regard to
vegetation.

The altitude of 5,000 feet, which we will, then, take
as the mean lower limit of the Alpine zone for
Switzerland generally, is as natural a dividing line as
can be found. Not only does it indicate the average
lower limit of the Coniferous forests, but at this
height all the physical conditions which we term
Alpine — such as shortness of the flowering season and
intensity of the illumination — are typically in force.
Further, the majority of the plants flourishing at or
above this elevation, possess all those peculiarities
of architecture, or, as the botanist terms it, " habit,"
which we associate especially with Alpine plants.
Biologically, then, these plants are Alpine, as well as by

1 Trees which shed their leaves in autumn. For this and other
technical terms, see Appendix I., p. 307. The Conifers, except the
Larch, are all evergreen.

VEGETATION AND ALTITUDE 7

elevation, even if we restrict the High Alpine or so-
called Glacial plants to a separate category. Many of
the latter, as we shall see in a later chapter, exhibit
the same peculiarities, though in a more marked
degree.

The gradual nature of the change in the vegetation
as we pass upwards from the Alpine to the High
Alpine region is usually better realised nowadays,
than the changes from the Lowland to the Subalpine
and the Subalpine to the Alpine zones. In these days
of mountain railways, when most travellers reach the
Alpine zone by train, the transitions between the
lower zones are apt to be overlooked. The slow
journey on foot, still happily necessary to reach the
High Alpine region, offers a far better chance of study
and reflection on this point. It is thus worth while
to those botanically inclined, for once at any rate, to
pass leisurely, on foot, from the lowlands to the Alpine
zone, as, for instance, along the Rhone valley and up
either the Zermatt or Saas Thai. In this way we
shall have an excellent opportunity of studying the
gradual nature of the change as we ascend.

It may be here mentioned, perhaps, that care
should be taken in regard to altitude in selecting a
Swiss centre for botanical explorations. The great
majority of Swiss health resorts1 do not lie within
the Alpine zone. Grindelwald, Caux, and Chamonix
(Savoy), for instance, are 1,500 feet too low. The

1 A list of Swiss resorts, classified according to altitude, is given in
the introduction to Baedeker's Switzerland.

8 INTRODUCTORY

higher the centre chosen, the more interesting the
flora.

With regard to the number of species of Flowering
Plants (Angiosperms) found within the Alpine zone,
estimates naturally vary. Ball calculated that in the
Alpine region of the whole chain of the Alps, 1,117
species are found, belonging to 279 genera and 60
natural orders. In the corresponding zone of the
Swiss Alps, I estimate the number of species as
about 900 out of a total of 2,350 Swiss Angiosperms.
Of these, some 300 are confined entirely to the Alpine
zone, while roughly 250 are also lowland plants, and
in many cases members of the British flora. The
rest occur both in the Alpine and Subalpine regions.

We now pass to some consideration of the
difficulties which a plant has to overcome in order
to survive the rigours of an Alpine climate. If we
are to understand and appreciate these difficulties,
we must arrive at some clear conception of the
internal economy of the plant itself. The plant has
two duties to perform. A duty to itself, to maintain
its own existence, and a duty to the next generation
— reproduction. We will now briefly discuss the first
of these.

The plant is a living being which may be likened
to a complicated steam engine or other piece of
machinery in motion. It is fashioned for one end
alone — the existence of the species. All plants are
built up of an enormous number of very small units,
termed cells. The cells differ very much in form,

VITAL PROCESSES IN PLANTS 9

substance, etc., and may be either living or dead.
All living cells contain that mysterious and complex
substance known as protoplasm, which is the seat of
vitality and of all the vital processes, such as growth.
The protoplasm of one cell is in communication with
that of the cells which surround it. Thus the whole
plant is really a mass of protoplasm, divided up into
minute compartments by cell walls, through which,
however, it is continuous from cell to cell.

Protoplasm is a very unstable substance. It is
constantly in a state of flux, some parts being built
up into other substances and others broken down. It
is on this fact that vitality depends.

We have therefore in each cell a machine capable
of building up and breaking down, not only substances
derived from without, but its own substance. In
order that this machinery may work, it requires food
— that is, raw material from which to manufacture a
finished product.

Both in animals and plants, the food necessity is
ever present, but plants obtain their food in quite a
different way to animals. Animals can make use of
substances of an extremely complex chemical nature,
built up chiefly of the elements carbon, hydrogen,
oxygen, and nitrogen.

On the other hand, the plant makes use of relatively
simple substances. The food of the green plants is
obtained partly from the atmosphere by means of the
leaves, and partly from the soil by the roots. Whereas
the Higher Animals have the power of locomotion,

10 INTRODUCTORY

and live in a single medium, the atmosphere, or in
water, most of the Higher Plants are fixed, and live
partly in the soil (or in water) and partly in the
atmosphere. The roots buried in the soil absorb
water and certain essential mineral salts, especially
those of nitrogen, but not of carbon. These raw
materials are passed on to the protoplasmic machinery
from cell to cell. This function is known as absorp-
tion.

On the other hand, the leaves perform the function
known as assimilation. They can withdraw carbon
dioxide from the atmosphere. This gas passes
through into the leaf by means of numerous minute
pores, which the botanist terms stomata. This simple
raw food material, with the addition of water derived
from the soil by means of the roots, is first of all
converted into a complex substance, such as sugar,
by the aid of the special portions of the protoplasm
of the leaf, which contain the green colouring matter
known as chlorophyll. It is the chlorophyll which
gives the green coloration to the majority of plants.
Chlorophyll is able, in the presence of light and other
favourable physical conditions, to absorb energy,
which gives it the power to convert the simple gases
of the atmosphere into complex sugars, which are
passed on in turn to the protoplasm of the growing
cells.

The protoplasm thus receives, and incorporates
into its own substance, raw food material from two
sources : the atmosphere, which supplies the carbon

VITAL PROCESSES IN PLANTS 11

compounds ; and the soil or water, which furnishes the
other elements. The raw food material is converted
into complex substances by the protoplasm itself, and
these are made use of for growth, reproduction, and
other vital processes. It is, in fact, the coal which
keeps the machinery in motion.

Now, if the roots of an Alpine plant are buried
in a frozen soil, one source of food- supply is, for a
time at least, cut off, and the plant must depend on
such reserves of water and mineral salts as it has in
hand. On the other hand, the leaves, if covered with
snow, are in darkness, and cannot assimilate, and
here another source of raw material fails for a time.
Plants can, however, by storing up reserves, survive
for a long period, during which both sources of food-
supply are cut off; but if this period is unduly
prolonged for any reason, the plant, having used up
all its reserves, may die, for the machinery can no
longer continue in motion.

In plants, then, as in animals, food is essential to
the life of the protoplasm, and is built up into its
substance to form new protoplasm. On the other
hand, protoplasm being unstable, is constantly break-
ing down, and the simpler substances which result are
returned to the atmosphere. In man and all the
higher animals, and also in plants, there is a twofold
exchange with the atmosphere. Man breathes by
means of his lungs — that is to say, he is constantly
exhaling carbon dioxide and water vapour, and inhaling
oxygen — a process known as respiration. If animals

12 INTRODUCTORY

alone existed on the earth, all the oxygen in the
atmosphere1 would have been long ago exhausted
and replaced by carbon dioxide. But the whole
economy of nature depends on the fact that in plants
the process, in the main, is just the reverse. It is
true that plants respire exactly like animals, absorb-
ing oxygen and giving off carbon dioxide ; but in the
case of a green plant, we can only detect this process
in the dark. This is because, in the presence of
sunlight, it is masked by the opposite, and far more
vigorous, process of assimilation, in which oxygen
is given out and carbon dioxide absorbed.

One further essential function of plants remains to
be discussed. All plants immersed in the atmosphere
are constantly losing water by evaporation — a loss
which has to be made good by absorption from the
soil by means of the roots. This process is known
as transpiration. It is not simply due to evaporation,
but is controlled by, and intimately connected with,
the vital processes of the plant.

With Alpine plants the tendency to excessive
transpiration in summer is very marked, owing to
the high day temperature and the dryness of the
atmosphere. The water vapour passes out through
the pores or stomata which occur on the leaf (Plate
XXIX., Fig. 1) and sometimes on the stem and
flowers also. Consequently we shall find in many

1 The atmosphere consists approximately of 23 parts by weight of
oxygen and 77 parts of nitrogen and a very small quantity of carbon
dioxide, about 3 to 6 parts in 10,000 measures.

REPRODUCTION OF OFFSPRING 13

Alpine plants special protections in connection with
these organs, in order to guard against excessive
transpiration.

Such briefly are the vital functions which the
plant has to perform to fulfil its duty to itself — the
maintenance of its own existence. We shall find that
many of the peculiarities of Alpine plants are to be
explained as special adaptations, in order to ensure
that these functions shall not be interfered with by
the influence of the severe climatic conditions, which
the plant has to face. Unless we have some com-
prehension of the vital processes of Alpine plants,
we shall naturally not possess the key to an
understanding of many of the peculiarities of habit
which they present, and thus it has seemed well to
enter here, at the outset, into these matters somewhat
at length.

The plant has also a second duty : the production
of offspring. The means by which this is effected
are more generally understood, and need only a passing
reference here. In the plants which stand highest of
all in a botanical sense, the Flowering Plants (Angio-
sperms), with which we are here alone concerned, the
essential features of the flowers, are as follows.

The male organs — stamens — produce the pollen
grains, which in turn furnish sperms, which fertilise
the eggs contained in the ovules. The ovules are
borne by and enclosed in modified leaves termed
carpels. Both the male and female organs may be
found in the same flower, or in separate unisexual

14 INTRODUCTORY

flowers, borne either on one plant or on two different
plants. In a very large number of cases, the male
and female organs, especially where they occur in the
same flower (the so-called hermaphrodite flowers), are
enclosed in a floral envelope or perianth, often
differentiated into an outer series, the calyx, composed
of sepals, and an inner series, the corolla, composed of
petals. The floral envelope may, however, be very
poorly developed in some flowers. It plays many
different parts in various plants, as we shall see. It
serves partly to protect the young developing sexual
organs in the unopened flower-bud, and in many cases
it also furnishes a conspicuous advertisement for the
allurement of insects, which carry the pollen of one
plant to the female organs of another, and so effect
cross-pollination.1

It will be our object in the present volume to
bear constantly in mind the influence of the severe
climatic conditions of the Alpine world on the vital
processes of the plant, both those relating to existence
and to reproduction. We shall find that many of the
peculiarities of Alpines are alone intelligible from this,
the biological standpoint, which is entirely different
from that of the systematic or descriptive Botanist.

We may illustrate these principles by reference to
the two national flowers of Switzerland : the Edelweiss
and the Alpenroses. Both will be found to show
interesting adaptations to their special surroundings.

1 Further information on the structure of the flower will be found
in Appendix II.

THE EDELWEISS 15

THE EDELWEISS.

The Edelweiss, Leontopodium alpinum, Cass.1
(natural order Composite, the Daisy family), Plate I.,
Fig. 1, about which so much romance has been woven,
and which is commonly believed to grow only in
situations almost unapproachable even to the most
hardy mountaineer, must be pronounced a complete
fraud in this respect. Scarcely a year passes but one
hears of some fatal accident as an unnecessary
corollary to the desire to gather this plant. Yet every
season huge masses of Edelweiss, which must
approach to tons in the aggregate, are gathered by
the Swiss peasants, or even grown in the lowlands
of northern Switzerland, for sale to the tourist. This
sale, I suspect, is a ready one, not so much because
of the interesting and unusual appearance of the
plant itself, as because of its reputed associations.

Yet the Edelweiss is not a rare plant. It might
almost be called common. Those who have really

1 Every plant has a Latin Christian name and Surname. The
Surname, or generic name, indicates the genus to which the plant belongs,
and the Christian name is the specific name, or name of the species.
The latter is regarded as an adjective, and is placed after the noun, the
generic name, the customary order in the Latin tongue. Thus JBellis
perervnis, the Common Daisy, is a member of the genus jBellis, and the
particular species, perennis. The name should always be written : Bellis
perennis, Linn. Linn, is a contraction for Linneeus, the name of the
botanist who founded the species, and who is called the " authority "
for the name. The necessity for quoting the authority arises from the
fact that in many cases the same plant has been described by two
botanists quite independently under different names, the oldest or first
description being regarded as alone valid.

16 INTRODUCTORY

interested themselves in Alpine plants are aware that
it can often be gathered near many of the Alpine
centres without the trouble of forsaking a well-
made path. This is true, for instance, of some of
the hills above Zermatt and Saas. Yet if we were
to set forth in order to collect this plant, our chances
of coming across it would usually be quite small, unless
we were guided by the experience of others.1 The
explanation is that the Edelweiss, while not a rare
plant, is exceedingly local in its distribution. It
does frequently occur among the most inaccessible
of crags, but even there it is often not to be seen.
On the other hand, it will sometimes cover a stony,
dry, almost level alp by the acre. Why its distribution
should be so local is a question which cannot be fully
answered at present. The fact remains that it is
usually restricted to the driest of situations in which
plants can flourish. Many other Alpines, such as
the Saxifrages, cling to the crevices of a cliff, but
these nearly all require some degree of moisture in
the scanty soil, or some situation well exposed to the
weather. Where the conditions are such that a
minimum degree of moisture is alone available, there
the Edelweiss will outstrip its competitors, and succeed
in the struggle for existence. Where it occurs in
surroundings in which other plants can flourish, there
it must compete against them for a bare livelihood.
Thus, as a rule, Edelweiss is restricted to the

1 Edelweiss is usually to be found in one or more localities near all
the great Alpine centres of Switzerland in the month of August,

PLATE I.

FIG. 1. — The Edelweiss (Leontopodium alpinum, Cass.).

FIG. 2.— The Alpine Rose (Rosa alpina, Linn.).

[To face v. 16.

THE EDELWEISS 17

driest and barest rocks, barren of other plants ; and
since such localities are relatively infrequent, the
Edelweiss is a local plant, though often exceedingly
abundant where it does occur. On the other hand,
in places where the circumstances that prevail appear
to be in every way adapted to its needs, the Edelweiss
is often conspicuous by its absence.

Thus the sentimental value of the Edelweiss does
not really depend so much on its rarity or difficulty of
collection, as on the fact that the localities in which
it grows are comparatively few and far between. It
is one of the most local of Alpine flowers, a fact in
itself of great botanical interest.

The Edelweiss is not a British plant, and the name,
though by now almost completely anglicised, is a
combination of two German words : edel = precious
and weiss = white.1 So we see that romance is bound
up in the very name itself.

Let us now examine the features of this plant
which give rise to the idea of whiteness. If we study
a specimen with a hand-lens, we shall find it is covered,
completely and thickly, with long, woolly hairs.
These hairs consist of empty cells. It is one of the
properties of light, that when it falls on innumer-
able, minute, transparent particles, we receive the
impression which we term white. A good and well-
known example is the foam of waves breaking on
the sea-shore. Water itself is colourless, but when
it is broken up into small particles or bubbles, as

1 Cf. the German Edelstein= precious stone.

18 INTRODUCTORY

in the case of sea foam, our eyes experience the
sensation which we call whiteness. The hair cells of
the Edelweiss produce a similar effect.

The Higher Plants have, as a rule, green leaves and
often green stems, and this is true also of the Edel-
weiss, but the green colour is here masked by the
coat of hairs.

The leaves of the Edelweiss, like those of many
other Alpines, are arranged in a small rosette just
above the soil. A single stalk springs from the
leaves bearing what, at first sight, appears to be a
solitary flower, but which in reality is a very com-
plicated structure consisting of several flower-heads,
each with a large number of individual flowers. It
is one of the peculiarities of the order Composite
to which the Edelweiss belongs, that the flowers
should be all massed together into one or more heads.
The single heads of a Daisy or a Sunflower, for
instance, are not flowers, but collections of a large
number of flowers, seated on a broad receptacle. If
we cut one of these heads through with a pocket-
knife lengthwise, we can see the receptacle, and also
separate the individual flowers from one another.1

At the same time, the head performs all the
functions of a single flower, and is in itself an adapta-
tion designed for that very purpose. In the Edelweiss,
however, the heads are very small and yellowish in
colour, and, further, they are grouped together into

1 For a full account of the head of a Composite, see Appendix II.,
p. 330.

THE EDELWEISS 19

what appears at first sight to be a single head.
Thus, what is apparently a single flower is really a
very complicated structure. There is a large central
head, composed of many flowers and equivalent to
the inflorescence of a Daisy or a Sunflower, sur-
rounded by a varying number, usually five, of other
smaller heads, the whole being wrapped round by
woolly leaves which are called bracts. These
bracts are the conspicuous part of the so-called
Edelweiss "flower."

But to return to the woolly coat : What is the
purpose of this adaptation ? Why is it present in this
plant and not in others ?

It must be remembered that the Edelweiss usually
flourishes in very dry situations, where there is com-
paratively little moisture in the soil. In this respect
it is like a plant growing in a desert. Consequently
it has to husband such water as it can absorb from
the soil by means of its roots with the greatest care.
Were it to transpire rapidly (see p. 12) — i.e., give
off water-vapour — the supply would run short, and
the stream of water passing from the soil to the
atmosphere, by means of the root, stem, and
leaves, would soon cease. Thus some contrivance
must be arrived at, which will prevent an undue loss
of water from the surface of the plant by evapora-
tion. The hairy coat is this contrivance, and therein
lies the great point of botanical interest presented by
this plant. The fact that, not only the leaves, but
the head- stalk, and even' the heads of flowers, all of

THE ALPENROSES 21

two more of its near relatives — the Everlastings,
belonging to the same order — which are much more
frequent in the Alps. One of these, Antennaria dioica,
Gartn., with white or rose-coloured heads, is fairly
common with us in Britain. The other, Antennaria
carpathica, Bl. and Fing., is a High Alpine, and has
brownish heads. Both have a similar cottony coat to
that of the Edelweiss, though shorter, but the woolly
bracts surrounding the flower-heads, which are so
prominent in the case of the latter, are very much
smaller and less conspicuous.

They grow in dry, stony places, and, as in the
case already discussed, their cottony coats prevent
excessive loss of moisture.

THE ALPENROSES.

The Alpenroses, of which there are two — the
Common Alpenrose, Rhododendron ferrugineum, Linn.,
and the Hairy Alpenrose, R. hirsutum, Linn, (natural
order Ericaceae, the Heath family) — are, equally with
the Edelweiss, national flowers in Switzerland.

They are, however, very much more abundant and
more easily obtained, and so are more in evidence
as personal decorations, on the not-infrequent
occasions on which the Swiss peasants make sport or
holiday. In the little hamlet of Sils, in the Engadine,
an excellent botanical centre, there are two rival inns :
the Edelweiss and the Alpenrose. Both names are
frequently used to designate hotels all over Switzer-

22 INTRODUCTORY

land, and possibly in other villages they may be
found in as close proximity as at Sils.

The Alpenrose, it may be said at once, is merely a
small-flowered Rhododendron, of blood relationship
to the Rhododendrons and Azaleas of our English
gardens, many of which were derived originally from
the mountainous regions of India, especially the
Himalayas. The genus is a large one, but does not
occur in Britain, unless we include in it Loiseleuria
procumbens, Linn., the Trailing Azalea.

The word Rhododendron is derived from two
Greek words, signifying respectively a rose and a tree.
For the Swiss species, however, the name "alpen-
rose " has become almost completely anglicised. At
the same time, some confusion appears to exist on this
point. It is often spelt or pronounced as if it were
two separate words : Alpine Rose. This is a common
error, and regrettable because there is in Switzerland
a true Alpine Rose (Rosa alpina, Linn.), Plate I.,
Fig. 2, which flourishes commonly in shrubby places
in the pastures, in much the same situations as some
of our wild roses in Britain.

The true Alpine Rose is, moreover, interesting
botanically as being "a rose without a thorn," since
prickles are often absent from the upper part of the
stems and from the leaves, though they may frequently
occur near the base of the plant. Occasionally the
whole shrub is well armed with thorns or prickles.

On the other hand, the single German word Alpen-
rose, means the "rose of the pastures." The term

THE ALPEN, OR PASTURES 23

alp J is used by the Swiss in a very definite sense, and
the alpen, or pastures, are not only of great economic
importance to this essentially pastoral people, but are
actually owned by them in common. The alpen high
above the valley (Plate III.) are the mountain pastures,
which provide food for the cows during the height
of summer. The word forms part of the names of
such well-known Swiss resorts as the Bel Alp, the
Riffelalp, both of which were originally, and are still
largely, places to which the cattle migrate in search of
grazing-ground in summer-time.

In Switzerland the commune is a highly developed
and ancient institution, and each commune usually
owns one or more alps, in the sense that its burghers
have the freeborn right of pasturing their cows on
certain alpen. Whereas in the villages the meadows
are for the most part private property, the pastures, or
alpen, are held in common by the inhabitants. In the
spring and summer, the hay growing in the meadows
is far too valuable to become the immediate fodder
for the cattle, for it has to be carefully cultivated
and stored up to feed them during the long winter
months. Thus in summer-time the cattle belonging
to the commune are sent to the pastures high up in
the hills, and these are only occasionally cut with the
scythe, and then late in autumn. As the summer
passes, the cattle climb higher and higher from alp to
alp in search of a fresh food-supply. The milk is
also made into cheese in the far-away chalets perched
1 The term "The Alps," is also derived from this word.

24 INTRODUCTORY

up on the mountain-side, where those in charge of
the cows are often completely isolated for weeks from
the world : so much so, that in many valleys the
church bells of the principal villages are rung at twelve
noon each day, to pass the time to those of the distant
alpen. This is the case, for instance, at Davos
Platz, where in addition the bells of St Johann
Kirche are rung on Saturday evening, usually about
six o'clock, to warn those on the mountains of the
approach of the Sabbath.

We have dwelt on the alpen, or pastures, at some
length here, for it is essential that they should be
clearly distinguished from the valley meadows, cut
regularly with the scythe, usually three times during
the summer months. The pastures are in many
respects as distinct botanically from the meadows as
they are often far away from them. In the following
chapters we shall discuss some of the typical Alpines
of the pastures and the meadows.

With this digression we must now return to our
Alpenroses. They are evergreen shrubs, two, three,
or more feet in height, often occurring in great
abundance in the higher pastures, among the Alpine
thickets, in or on the borders of the forests, especially
where they fringe the margins of the Alpine lakes.
Of the two species, the Common Alpenrose, Rhodo-
dendron ferrugineum, Linn. (Plate II.), is by far the
most abundant in Switzerland. It is easily recog-
nised by the fact that the under-sides of the older
leaves exhibit a rusty-brown appearance, though the

PLATE III.

«8 *

^ ^
CO ^

THE ALPENROSES 25

new leaves of the current year at the tips of the
branches are green beneath. There are also no hairs
in this species on the edges of the leaf.

The Hairy Alpenrose, Rhododendron hirsutum,
Linn., is more common on limestone soils. At one
time it was believed to be entirely confined to
calcareous rocks, but it is now known that this is not
the case (p. 117). This shrub is very similar in appear-
ance to the Common Alpenrose, but all the leaves
are green beneath, and are fringed with hairs along
their margins. In both, the numerous stem branches
are bare of leaves except near the tips.

There is nothing of much interest to add as
regards the flowers of the Alpenroses. But the
leaves of both species are specially protected, as in the
case of the Edelweiss, against excessive evaporation
(transpiration; see p. 12), though here in quite a
different way and in a less marked degree. The
leaves of all members of the Heath family (Ericaceae)
are evergreen — that is to say, they are not all shed at
once in autumn, but new leaves gradually replace the
old ones, which last in the case of the Alpenroses
from three to four years, so that the shrubs are
never leafless even in winter. The great majority
also show some adaptation against excessive transpira-
tion, though form of the adaptation varies greatly in
different cases.

In the Common Alpenrose the rusty-brown colour
of the under-side of the older leaves is the outward
and visible sign of the means whereby excessive

INTRODUCTORY

evaporation is prevented. In the Hairy Alpenrose a
precisely similar arrangement is met with : only, the
older leaves do not turn brown below.

If we were to examine such a leaf carefully under
a microscope, by means of thin sections cut with a
razor, we should find that the lower surface is covered
with a number of green or rusty brown coloured

FIG. I.— The Leaf and Scales of the Common Alpenrose
(Rhododendron ferrugimum, Linn.).

1. Transverse section of a leaf, showing the scales on the lower surface.

Magnified 14 times.

2. Transverse section through a scale, on the right side of which a stoma is

seen sheltered by the scale. Magnified 190 times.

3. Surface view of a scale. Magnified 190 times.

scales. Text-fig. I., 1, shows a transverse section of
a leaf and the scales below. In Text-fig. I., 2, we
see a scale cut in transverse section, on the right-
hand side of which a pore, or stoma, is seen sheltered
by the scale. In Text-fig. I., 3, the surface view of the
scale is depicted. The scales are broad, flat structures

THE ALPENROSES 27

attached to the leaf by a very short stalk. In the
Common Alpenrose the lower surface of the leaf is
thickly studded with these scales, while in the Hairy
Alpenrose the scales are relatively fewer, and are
always green in colour.

The function of the scales is to protect the pores,
or stomata, which are confined to the lower surface of
the leaf. In this way, the danger of excessive loss of
water by evaporation from the leaves through the
pores is decreased. The adaptation, though entirely
different in form, is thus fashioned to the same end
as that which we have just discussed in the case of
the Edelweiss. It is also much less marked than in
the latter, the different conditions under which the
two plants grow being sufficient to account for the
dissimilarities observed. Thus the Edelweiss and
Alpenroses furnish excellent illustrations of how
Nature attains her ends by different means.

CHAPTER II

TYPICAL FLOWERS OF THE ALPINE PASTURES — THE
ANEMONES AND GENTIANS

" WHAT is the best time of year to visit the Alps in
order to see the Alpine flora at its best?" is a
question frequently asked. The answer is : The period
of late spring and early summer, i.e., from the middle
of June to the middle of July, preferably the last two
weeks of June. The early spring flowers begin to
appear at the end of April and during May, the
precise date varying somewhat with the particular
season, and depending on the depth of winter snow
and the period at which the spring thaw begins in
earnest. These are among the most beautiful of
Alpine plants, and well repay an early visit. But in
June many of them may still be found in flower in
the higher pastures at a height of from 6,000 to 8,000
feet and upwards. At this time of year also the fruits
of the early spring species are often mature, and in
many cases are interesting objects of study, too
frequently neglected.

Again, at the end of June, the Alpine meadows,

JUNE IN THE ALPS 29

with their beautiful flora, quite distinct as a whole
from that of the pastures, are ripe for the scythe,
though as yet uncut. Once the meadows have
yielded their first crop of hay, which is usually the
case early in July, or even in June, one of the great
floral beauties of the Alpine valleys has for a time
departed. By the end of July, and in early August,
though a few summer plants such as the Pinks are
now in flower, the height of the flowering season is
over.

A good illustration is furnished by the EiSelberg,
above Zermatt, which is almost entirely bare of
blossom by the end of July, when the annual stream
of summer visitors commences. Yet in June it is
famous as possessing one of the richest and most
varied floras in the Alps, and especially for certain
plants which are rare elsewhere. On the other hand,
the High Alpine region of the Gorner Grat (3,000 feet
above the Riffelalp) is often by the end of July nearly
free from snow, and furnishes an excellent collecting
ground rich in the dwarf plants of the High Alpine
zone.

It is perhaps unnecessary to add that everything
is in the favour of a visitor in early summer. Greater
comfort in travelling and in hotels, as well as less
intense midday temperature, contribute to the enjoy-
ment of "the season" for Alpine plants.

One of the great peculiarities of the Alpine flora
is the tendency to "rush into flower" at the earliest
possible moment in spring-time. With us in Britain,

30 TYPICAL FLOWERS OF ALPINE PASTURES

the flowering period is more generally distributed
throughout the spring and summer, and even extends
into the autumn.

In the Alps, the great majority of plants are in
flower, or have passed the flowering stage, by the
beginning of July, though some continue to flower
during the early summer. Moreover, a large number
of Alpine plants are specially constructed with a
view to flowering at the earliest possible moment, as
we shall see in subsequent chapters. There are only
two other floras which exhibit this peculiarity in the
same degree — the vegetation of the Arctic regions,
and of the Cape (S. Africa).

The chief reason which has led to this peculiarity
is to be found in the shortness of the summer season
in the High Alps. The flowering period is but one
stage, almost the preliminary step, in the process of
reproduction. At its close much remains to be done
before the seeds are ripened. Time must also be
allowed for their distribution, and for them to obtain
a firm hold in their new surroundings, before the
winter's snow rings down the curtain.

It may be of interest to note the succession of
flowering among some of the earlier spring flowers.
The dates naturally vary somewhat from year to year,
and according to the locality.

Dr Christ states that a Swiss botanist, named
Brugger, observed at St Moritz, in the Upper
Engadine, the following order of flowering among
the heralds of spring. In the particular year in

SPRING FLOWERS 31

question, the snow on the pastures did not disappear
completely until the 3rd May. Yet on 22nd March,
forty-two days previously, the first Spring Gentians
(Gentiana verna, Linn.) were in flower, and with
them, or very slightly later, the Spring Potentil
(Potentilla verna, Linn.) and the Spring Anemone
(Anemone vernalis, Linn.) also appeared. On 2nd
April, Crocus vernus, All., sprang into flower in
the meadows, while, on the day following, the
Coltsfoot (Tussilago farfara, Linn.) showed on the
dry, stony banks. On 18th April, the Bird's-eye
Primrose (Primula farinosa, Linn.) and the Hairy
Primrose (P. hirsuta, All.) also appeared. By 24th
April, such characteristic species in the pastures as
the Bell Gentian (Gentiana acaulis, Linn.), the Alpine
Anemone (Anemone alpinav&r. sulphur ea, Linn.), and
Poly gala chamcebuxus, Linn., were in flower. By
18th May, the Long-spurred Violet (Viola calcarata,
Linn.) and the Oxlip (Primula elatiort Jacq.) had
flowered, and these were quickly followed by a host
of other plants. The Alpenroses did not, however,
bloom until 20th June, and even this date was quite
early for these plants at an elevation of 6,000 feet.

If we make a preliminary survey of the plants to
be found near some Swiss resort in an Alpine valley,
about 5,000 feet in altitude, we shall find that we can
classify them roughly according to the kind of locality
or habitat in which they flourish. Some are rock
plants, others are only found in the shade of the
Alpine forests. We shall also observe that the plants

32 TYPICAL FLOWERS OF ALPINE PASTURES

of the Alpine meadows are, for the most part, distinct
from those which flourish in the pastures. We can
thus distinguish certain groups or associations of
Alpine plants, each fitted or adapted to flourish under
special circumstances. On the other hand, certain
Alpine species will be found to belong to more than
one association, and to be widely distributed through-
out the valley.

The Alpine pastures and the valley meadows are
typical associations on a large scale. The Alpine
forests afford another instance. The conditions of
life which prevail in these three types of habitat are
very different, and since certain plants flourish
in one and not in the others, it may be assumed that
such species are specially suited to the particular
conditions under which they live, and are not adapted
to the circumstances which prevail elsewhere. Con-
versely, those species which are generally distributed
and frequent in more than one association, have
remained less specialised, and are therefore to be
found more widely. They, however, are comparatively
few in number.

Within the great associations, such as the pasture
plants or the forest plants, are a number of smaller
communities, the units which build up the great
associations. Thus in the pastures we find typical sub-
associations in the rock plants and in the inhabitants
of the Alpine marshes. In the forests, we have the
plants forming the forests themselves, and other species
which thrive in their shade.

PASTURE PLANTS 33

In the present volume we propose to group
Alpine plants according to their associations, large
or small, and to discuss the more typical and
interesting members of each. The first few chapters
will be devoted to the plants of the pastures,
beginning with some of the more characteristic and
abundant genera. We shall also devote chapters to
some of the smaller associations, such as the rock
plants, the marsh plants, and the inhabitants of the
artificial modifications of the pastures — the meadows.

We will then discuss the High Alpines — i.e., the
inhabitants of the highest pastures — and finally the
Alpine thickets and forests and their smaller associa-
tions, the shade plants. The last two chapters will
be devoted to a general summary of some of the
peculiarities and other interesting features of Alpine
plants, and to the theories advanced as to the origin
of the Alpine flora of Switzerland.

The pasture plants form a very large association
belonging to many different families. Of these, how-
ever, four are especially well represented by a large
number of abundant species in the Alps, and
their members contribute the most characteristic
feature of this association. The Buttercup family
(natural order Kanunculacese), represented by the
Anemones, the Alpine Buttercups, and several other
genera, the Gentian family (natural order Gentian-
acese) by the Gentians (genus Gentiana), the Primrose
family (natural order Primulacese) by the Soldanellas,
Primulas, and Audrosaces, and the Bell-flower family

34 TYPICAL FLOWERS OF ALPINE PASTURES

(natural order Campanulacese) by the Campanulas or
Bell-flowers, and the Rampions, are the most typical
of all Alpine orders. We shall in this and the
following chapters pay special attention to these
plants, and thus gain a general idea of the chief and
most characteristic members of the Alpine pastures.

THE ANEMONES.

The Anemones or Wind-flowers (natural order
Kanunculaceae, the Buttercup family) are very
abundant in spring in the Alps, though the number
of species is not very large. The Spring Anemone
(Anemone vernalis, Linn.) is one of the earliest plants
to flower in the pastures, when the snow begins to
melt. Our two British Anemones, the Wood
Anemone (Anemone nemorosa, Linn.), so common in
plantations in early spring, and the Pasque-flower
(Anemone pulsatilla, Linn.), a much rarer plant, grow-
ing chiefly on chalk-downs and other limestone soils,
both occur in Lowland Switzerland, but not in the
Alpine zone.

The Alpine Anemones fall into two natural
groups, of which we may take the Spring Anemone,
the Alpine Anemone, and the Narcissus-flowered
Anemone as typical members.

THE SPRING ANEMONE.

We will begin with the Spring Anemone (A.
vernalis, Linn.) (Plate IV.). By the time the annual

PLATE IV.

The Spring Anemone (Anemone vernalis, Linn.).

[To face p. 34.

THE SPRING ANEMONE 35

influx of tourists has reached the Alps, this plant is
long past flowering in the lower pastures, and we
must ascend to 7,000 to 8,000 feet, if we wish to see
it in its prime.

In many respects the Spring Anemone recalls our
English Pasque-flower (A. pulsatilla, Linn.), to which
it is closely related. Just above the ground there is
a small rosette of leaves, which are much cut and
divided. The rather long leaf-stalks end below in a
broad sheathing base. One or more solitary flowers,
each mounted on its own flower-stalk, springs from
among the leaves. At first, when the flowers are
young, the stalks are short, and at this stage the
flower itself projects but little beyond the rosette of
leaves. As, however, the flower-buds mature, the
stalk lengthens and carries up the flower. Just
below the flower itself, we find several narrow, much-
divided structures, which are really three leaves,
much dissected, arranged in a circle on the flower-
stalk. These form what the botanist calls an
involucre, the presence of an involucre being char-
acteristic of the Anemones. This structure serves to
some extent to protect the young flowers.

The perianth members of the flowers, usually five
or six in number, are of a beautiful, violet or pinkish-
violet tint externally, though white inside. Within
the perianth we find numerous stamens and ovaries,
all arranged in a spiral fashion on the receptacle.

The chief interest of this plant lies in the long,
silky, yellowish-brown hairs, which clothe both the

36 TYPICAL FLOWERS OF ALPINE PASTURES

flower-stalks, the involucre segments, and the outer
surface of the perianth members. These hairs add
greatly to the beauty of the plant (Plate IV.). They
are commonly regarded as serving to keep the plant
warm during the season of melting snows. It is very
unlikely, however, that this is their real use. It is
more probable that they tend to lessen the risk of
excessive transpiration (see p. 12) from the upper
portion of the plant. It must be remembered that
when this Anemone flowers, the soil in which the
roots are buried is at a very different temperature
from the atmosphere. The sun during the snow-
melting season is hot, but the soil is still either
frozen or very cold. Hence if the above-ground
portion of the plant were to lose water too rapidly,
the loss could not be made good by the roots. The
hairs probably serve to lessen this risk in much the
same way as those of the Edelweiss, discussed in the
last chapter.

The fruit of the Spring Anemone (Plate V., Fig. 1)
is a very common " find " in the pasturages in summer.
It is not only beautiful, but botanically interesting.
It is closely similar to, though smaller than, that of
the Alpine Anemone, which we will shortly describe
in detail (p. 39).

Before leaving the Spring Anemone, we may
mention that in the Zermatt and a few other valleys
of Canton Valais, another species, Haller's Anemone
(Anemone halleri, All.), is found, which very closely
resembles the Spring Anemone in many points. This

PLATE V.

Q. c

'S.I

*S 1

I a

THE ALPINE ANEMONE 37

plant, however, loses its leaves in autumn, whereas
those of the Spring Anemone persist throughout the
winter.

THE ALPINE ANEMONE.

The Alpine Anemone (Anemone alpina, Linn., and
its variety sulphur ea, Linn.) (Plates VI. and VII.)
is the most striking of all the Swiss Anemones, and
one of the most handsome of Alpine plants. It is
the large white, or more often sulphur-yellow, "Wind-
flower " of the pastures. It flowers in June and July,
according to the altitude, and in many districts it is
exceedingly common. It varies in height from six
inches to a foot or more. It has rather large, much-
divided leaves mounted on long stalks, which spring
from the stem just above the surface of the soil. The
flowers, which are also borne on fairly long flower-
stalks, are solitary. As in the Spring Anemone, there
is an involucre (see p. 35) of three leaves on the
flower-stalk below the flowers, but here the leaves
are large and highly divided, and altogether much
more like foliage leaves. In the photograph on
Plate VII., Fig. 1, two young flowers just expand-
ing can be seen still partly sheltered or enclosed by
the involucre, exactly as we noticed in the case of the
Spring Anemone. As the flowers mature, the stalks
between the involucres and the flowers grow rapidly,
and thus the flowers are carried up out of the
involucre.

In the typical Alpine Anemone, the perianth

38 TYPICAL FLOWERS OF ALPINE PASTURES

members are either white, or white slightly tinged
with blue on the outside. In the yellow-flowered
variety, sulphured, which is sometimes regarded as a
distinct species, the flowers are a beautiful, uniform,
pale sulphur-yellow colour. Curiously enough, this
variety is in Switzerland generally much more
abundant than the true " alpina " with white flowers.
Often, however, they may be found growing side by
side. At one time it was thought that the sulphur
variety flourished only on granite soils, but this is
really not the case, and the difference in colour
between the species and the variety bears no relation
to the soils on which they grow.

The masses of these flowers (Plate VI.), forming,
as it were, miniature forests of Anemones, which are
often to be seen on the steeper slopes of the pastures,
are among the most wonderful sights in the Alpine
world. When the flowers are mature, the perianth
members open out and catch the sunlight on their inner
sides, thus greatly adding to their conspicuousness.

The plants are only moderately hairy, and in this
respect contrast with the Spring Anemone or the
Edelweiss, where the conspicuousness of the whole
plant, as we have seen, is largely increased by its
hairy coat.

The flowers of the Anemones are constructed much
like those of a Buttercup (see Appendix II., p. 328) :
except that the perianth is not differentiated into
calyx and corolla. If we look closely at the flowers
growing in profusion on some bank in the pastures,

PLATE VI.

A Group of Yellow Alpine Anemones (Anemone alpina, Linn.,
var. sulphured. Linn.).

[To face p. 38.

THE ALPINE ANEMONE 39

we shall find that many of them are entirely male,
the ovaries or female organs having been completely
suppressed, and stamens only being present. This
remarkable phenomenon — the occurrence of male
unisexual flowers, in addition to flowers with both
sexes (hermaphrodite) — is probably far from infrequent
among Alpines. It occurs in the case of the White
Veratrum (Veratrum album, Linn.) (p. 123), another
typical pasture plant, and also in the White Dryas
(p. 128) and the Mountain Avens (p. 128). Its
significance is not yet clearly understood, but may
be connected with the fact that the flowers of the
Alpine Anemone are quite devoid of honey, pollen
forming the sole attraction to insects.

In the photograph on Plate VI. the lowest flower
has stamens only, and two at least of the six other
flowers are also male, the highest flower being
typically bisexual. Two male flowers are also seen
on Plate VII., Fig. 2.

The fruit-head of the Alpine Anemone (Plate V.,
Fig. 2) is a very beautiful structure. The individual
fruits consist of a small sac below, enclosing a single
seed, prolonged above into a long, feathery structure
called an awn. A very large number of these awned
fruits are borne in a head. It is interesting to examine
different stages in the formation of this fruit (Plate
VIII., Fig. 1, and Text-fig. II.). The awn grows in
length exceedingly rapidly. At the beginning of the
flowering stage, the ovary, which later forms the fruit,
will be found to be rather small in comparison (Text-

40 TYPICAL FLOWERS OF ALPINE PASTURES

fig. II., 1). When the flowering stage has passed
(Plate VIII., Fig. 1), the perianth members fall off
much earlier than in the case of the Spring Anemone

FIG. II. — Fruits of the Alpine Anemone (Anemone alpina, Linn.) in
various stages of development. Magnified twice.

1. In fully-opened flower. 2. Intermediate stage. 3. The mature fruit.

(cf. Plate IV.), and the stamens wither. The fruits,
or rather the awns, have begun to increase in length
(Text-fig. II., 2). They continue to grow rapidly,
the awns developing short hairs and a spiral twist

PLATE VII.

o c

£ 13

I <

THE NARCISSUS-FLOWERED ANEMONE 41

below, until they reach a length of 1 to 2 inches in
the mature stage (Text-fig. II., 3). The fruits are
now ready for distribution (Plate V., Fig. 2).

If we choose a ripe fruit-head on a windy day
and detach the awns with their seeds, and cast them
to the wind, we shall find that they are admirably
adapted for travelling in the air, and will often cover
considerable distances from the parent plant, in much
the same manner as the parachute-like fruits of the
Dandelion, with which everyone is familiar. Thus
the awn is an air-flying device, and extremely effective
as a means of distribution.

Awned fruits are not common among Swiss
Alpines, though they are conspicuous in the case of
some other Anemones, such as the Spring Anemone,
the White Dryas, and the Creeping and Mountain
Avens. The fruits of our common Traveller's Joy
(Clematis vitalda, Linn.) are familiar examples of the
same nature, though this plant does not occur in
Alpine Switzerland.

THE NARCISSUS-FLOWERED ANEMONE.

The beautiful Narcissus - flowered Anemone
(Anemone narcissiftora, Linn.) (Plate VIII., Fig. 2)
is in many respects a marked contrast to the Alpine
and Spring Anemones. It is not so abundant, and
is apt to be rather local in its distribution. It grows
chiefly in the pastures where the flowers are thick and
the grass long, and especially where the soil is fairly

42 TYPICAL FLOWERS OF ALPINE PASTURES

moist but not wet. It differs from all the other Swiss
Anemones in the grouping of the flowers into stalked
heads, which the botanist terms umbels, the flower
arrangement so characteristic of the great family of
plants known as the Umbelliferse, which includes the
Parsley and Hemlock. Usually there are from five
to eight flowers in the umbel, but the number varies
somewhat.

The flowers are white in colour, often streaked or
"blushed" with delicate rose-pink on the outside.
Just below the point at which the stalks of the
individual flowers unite, an involucre of three leaves
is seen, which are much less divided than in the case
of the other Anemones already discussed.

The Narcissus-flowered Anemone is also distin-
guished by the fact that the fruits are not prolonged
into a hairy awn. They are, on the contrary, quite
simple, resembling those of our British Wind-flower
(Anemone nemorosa, Linn.), and are not specially
adapted to travel in the air. The flowers, like those
of the Alpine Anemone, are entirely devoid of honey,
in which they are rather exceptional among Alpine
flowers.

THE GENTIANS.

The Gentians (natural order Gentianaceae, the
Gentian family), like the Anemones, are highly
characteristic of Alpine regions, though by no means
confined to them. In Britain, in addition to the
Yellow wort (Chlora perfoliata, Linn.), the Centaury

PLATE VIII.

THE GENTIANS 43

(Erythrcea centaurium, Pers.), the Buckbean (Meny-
anthes trifoliata, Linn.), and other genera not found
in Alpine Switzerland, we have five Gentians, all of
which occur in the Alps, except the Marsh Gentian
(Gentiana pneumonanthe, Linn.), which does not
extend beyond the Lowlands. One other British
species, the Autumn Gentian (G. amarella, Linn.), is
believed to be very rare in Switzerland, occurring only
in the Lower Engadine.

As opposed to our five British Gentians, there are
no fewer than eighteen species to be found within the
Alpine zone in Switzerland, many of them being widely
distributed and often abundant. Three of these are
especially in evidence in the High Alpine region.

The best-known Alpine Gentians are those which
possess blue flowers. But it must not be imagined
that all Swiss Gentians have blue flowers, though this
is true of the majority. Some species, as we shall see,
have yellow or red corollas. The blue-flowered plants
are, however, very much in evidence in the Alps, and
this is somewhat remarkable, for blue, as a colour, is
not so strikingly conspicuous as red or yellow.

The colours of Alpine flowers have been the
subject of repeated investigations at the hands of
botanists, especially in recent years. At one time it
was thought that there was actually a larger percent-
age of blue-flowered plants within the Alpine zone
than in the plains. This would appear probable,
when we remember that, in addition to many Gentians,
other blue- or purple-flowered plants are abundant,

44 TYPICAL FLOWERS OF ALPINE PASTURES

such as the Bell-flowers, the Rampions, the Forget-
me-nots, and the Eritrichium, many Geraniums, the
Globularias, the Alpine Toadflax, certain Monkshoods,
and the Opposite-leaved Saxifrage. But when we come
to statistics, we find, as is so often the case, that not
only are our first impressions not confirmed, but they
are shown to be erroneous. Dr Fisch has pointed out
that the colour proportions among Alpines are about
30 per cent, white-flowered species to 27 per cent,
yellow-flowered, and 19 per cent, red-flowered and 24
per cent, violet- or blue-flowered species. In the
flora of Davos, he finds that only 36*8 per cent, of the
total species have red, blue, or violet flowers, which
appears to be about the general average, and is quite
comparable to the proportion found among species
growing in the plains.

The blue-flowered Alpines are, however, specially
noticeable, owing to the intense depth of the colora-
tion and the large number of the individual flowers.
It has been shown repeatedly that the pigment which
is contained in the petals, and to which the colour is
due, increases in intensity as we pass from the plains
to the Alpine zone. While this is the general rule for
all colours as well as blue, it does not hold good in
every case. The flowers of some species, such as the
Wood Geranium (Geranium sylvaticum, Linn.), are
stated to be less intensely coloured in the Alps than
in the plains, while in other species there appears to
be no appreciable difference in the depth of the
colour in the two cases.

THE COLOURS OF FLOWERS 45

As to the cause of the greater intensity of colour
found in many Alpine flowers, it is not possible to
conclude finally at present. There are two alter-
natives. Either it is due to the greater intensity of the
illumination, or it may be a special adaptation among
Alpines to serve as an insect advertisement. There is
much to be said for both theories. Prof. Bonnier has
made experiments subjecting various plants, under
suitable conditions and with necessary precautions, to
the influence of a strong continuous light. He finds
that the chlorophyll, or green colouring matter of
the leaves, is thereby rendered more intense, and
the chlorophyll grains more numerous and more
evenly distributed. If this is the case with chloro-
phyll, it may also be the same with the pigment
granules in the petals. In many flowers, especially
those with yellow, orange-yellow, or orange-red
corollas, the pigment is solid and in the form of
granules, or, to state it more accurately, is contained
in minute specialised portions of the protoplasm
(p. 9) called chromoplastids, just as chlorophyll
(p. 10) is also held in small specialised protoplasmic
bodies. In a large number of red, blue, and purple
flowers, however, and also in some yellow flowers, the
pigment is held in solution in the cell sap.

On the other hand, it is known that there is a
larger percentage of flowers which are cross-pollinated
by means of insects, especially by butterflies, in the
Alps than in the plains. Some of these flowers are
specialised to certain groups of insects — that is, they

46 TYPICAL FLOWERS OF ALPINE PASTURES

are so constructed that only certain kinds of insects
can reach the honey legitimately. It has also been
proved experimentally that certain insects favour a
flower of one colour, and will avoid or overlook a
flower of another. Blue flowers, for instance, are,
as a rule, "bee flowers," while many white flowers
are visited by small flies. It is therefore possible
that in many cases not only the colour, but the
increased density of the pigment, met with in Alpine
flowers, may be primarily due, not to the special
physical conditions of the Alpine world, but to a
specialisation in favour of a particular class of insect
visitor.

In connection with the colours of Alpine flowers,
it may be remarked that certain species which
normally bear coloured flowers are occasionally found
to produce white flowers. We are all familiar with
the white Heather; and white forms of Alpine flowers,
such as of the Common Bell Gentian or a Bearded
Campanula, are equally prized on account of their
rarity in the Alps. The tendency to produce occasional
white flowers is greatest in those plants with blue,
pink, or red flowers, and least among the yellow-
flowered species. White-flowered plants will also
sometimes assume a yellowish hue. In others, again,
such as the Spring Crocus or the Field Pansy ( Viola
tricolor, Linn.), the colours, or rather the combinations
of colours, of the flowers are always fluctuating. The
causes which lie at the root of these colour-changes
are complex, and are not yet fully understood. It is

THE ALPINE GENTIANS 47

thus impossible to enter into the matter here, though
it may be stated that in some cases the white flowers
occasionally found are instances of fresh variations ;
whereas in others, the lack of a colour pigment in the
petals, or the development of an exceptional pigment,
may represent a reversion to an ancestral type. At
any rate, these variations in colour have no connection
with the chemical nature of the soil, as was formerly
thought possible, but they arise from deep- seated
tendencies, which find their expression in the exist-
ence of the individual, and the evolution of the race.

But to return to the Alpine Gentians : we will
commence with the blue-flowered species, which are
universally regarded as among the most interesting of
Alpine plants. These fall naturally into three groups.
First we have the Gentians with star-like flowers.
The corolla formed by the united petals consists of a
narrow tube below, the free portions of the petals
expanding above into radiating lobes, which, when
the flower is open, are spread out nearly at right
angles to the tube. Between the free portions of the
petals, small lobes occur, each divided into two.
When the flower is closed, the free portions of the
petals point upwards and are twisted together. The
Spring, Bavarian, and Snow Gentians belong to this
group. In these the leafy stem is short, though it is
quite obvious.

Next we have the Bell Gentians with very short
stems, and corollas in the form of a large bell,
the mouth pointing upwards to the sky when the

48 TYPICAL FLOWERS OF ALPINE PASTURES

flower is open. The Common Bell Gentian and the
Broad-leaved Gentian belong here. The former is
often called the Stemless Gentian.

The third group, the Fringed Gentians, or Gentian-
ellas, are usually much-branched plants with corollas
shaped like those of the first group. But the
entrance to the corolla tube, or the throat of the
corolla, as it is termed, is closed by a fringe of scales,
which are outgrowths from the inner surface of the
petals.

THE SPRING AND BAVARIAN GENTIANS.
The Spring Gentian (Gentiana verna, Linn.) (Plate
X., Fig. 1), is, as we have seen, one of the earliest
flowers to deck the Alpine pastures on the retreat of
the winter's snow. The Bavarian Gentian (Gentiana
bavarica, Linn.) does not flower until later, usually in
July, and is perhaps often more in evidence in the
High Alpine region than in the Alpine zone. The
two plants are both perennials, very similar in appear-
ance and liable to be mistaken for one another.
The underground stems produce numerous, very leafy
shoots, and flowering shoots ending in a single flower
of an intense azure-blue, especially in the case of the
Bavarian Gentian. In the Spring Gentian, the leaves
are more or less elliptical in shape and pointed. The
leaves on the flowering shoot are arranged in one or
two pairs, which are smaller than those forming the
rosettes close to the ground. The Bavarian Gentian
has blunt, spoon-shaped or egg-shaped leaves, and

THE SNOW GENTIAN 49

three or four pairs of leaves on the flowering shoots,
nearly similar in size to those below.

The chief feature of interest presented by these
two Gentians is the intensity of the blue coloration
of the flowers, a feature in which they are perhaps
only rivalled in the Alps by Eritrichium nanum
(p. 183).

The flowers are scentless. Their method of
fertilisation is essentially similar to that of the Bell
Gentian, which we shall describe in detail shortly.
Both the Spring and Bavarian Gentians are adapted
to cross-pollination by the agency of butterflies and
moths, the Hawk -Moth (Macroglossa stellatarum)
being their most frequent and important visitor. The
insects are attracted by the brilliant coloration, and
by the honey secreted at the base of the ovary. The
small double-toothed lobes or appendages between
the free portions of the petals serve to protect the
entrance to the corolla tube and to keep out "unbidden
guests," which can perform no service to the plant by
effecting cross-pollination.

THE SNOW GENTIAN.

The tiny little Snow Gentian (Gentiana nivalis,
Linn.), fully -grown specimens of which almost
resemble in size the seedlings of many other plants,
is interesting as being an annual, and not a perennial,
like the majority of Alpine Gentians. The whole
plant is very slender, and does not exceed 4 to 6

50 TYPICAL FLOWERS OF ALPINE PASTURES

inches in height. The stem is simple or branched,
and bears a few small leaves, the upper ones arranged
in pairs. It ends in a small solitary flower, like that
of the Spring Gentian, but smaller.

The Snow Gentian, like the Spring Gentian, is a
British plant, though it is rare with us, and is only
found on a few of the higher Scotch mountains. In
the Alps it is a very common plant in the pastures,
and is not, as perhaps the name Snow Gentian implies,
by any means necessarily confined to high elevations.
In fact, its upward range ceases at about 9,900 feet,
while it is much commoner at elevations of 5,000 to
6,000 feet. It is thus not a High Alpine plant at all.

The flowers of this species are extraordinarily
sensitive to sunlight. They are nearly always closed
unless the sun is shining very brightly, and, the
moment the sun disappears behind a cloud, the
flowers may shut with considerable rapidity. They
thus sometimes open and shut many times in the
course of an hour, changes in temperature acting as a
signal to the plant to open or close its flowers. It
should also be observed that when the flowers shut,
not only are the free portions of the petals held erect,
but they twist together in a spiral.

THE COMMON BELL GENTIAN.

We now pass to consider the Bell Gentians, which
have the largest corollas of the blue-flowered Alpine
Gentians. We will take the Common Bell Gentian,

PLATE IX.

A Plant of the Common Bell Gentian (Gentiana acaulis, Linn.), the Corolla
cut open to show the Stamens and the Ovary.

THE COMMON BELL GENTIAN 51

often called the Stemless Gentian (Gentiana acaulis,
Linn.) (Plate IX.), as a typical example. The plant is
quite unmistakable, and is especially frequent on
limestone soils. The stem is very short, though not
absent altogether, as the somewhat unfortunate
specific name implies. The leaves are arranged in a
beautiful little rosette on the stem, just above the
surface of the ground. From the rosette springs a
single flower, of large size, mounted on a stalk
which varies from 1 to 4 inches in length. The flower
is of a deep blue colour, and is shaped like a church-
bell, the mouth of the bell being turned upwards,
facing the sky.

There are no other Swiss Gentians at all similar
to the Common Bell Gentian, with the exception of
Gentiana excisa, Presl. ( = G. latifolia, Gren. and
Godr.), and G. alpina, Vill., both of which are
probably only varieties of the same plant, though by
some they are regarded as distinct species.

The large size of the flowers of the Common Bell
Gentian prompts us to peep within the bell, and to
study the form of the male and female organs, which
we shall find are full of interest. First of all, we
will slit open the corolla from base to summit
with a needle, or a pin, in the manner shown in
Plate IX. The photograph also shows the rosette
of rather leathery leaves, and the flower-stalk with
its two pairs of smaller leaves. Outside the bell and
attached at its base, we find a tubular calyx composed
of five small, green leaves united together. The five

52 TYPICAL FLOWERS OF ALPINE PASTURES

petals forming the bell are almost completely fused,
the recurved portions at the edge of the open bell
being alone free.

Internal to the corolla we find five stamens, which
spring from the base of the bell. Their stalks are
quite free from one another, but the pollen-producing
organs, the anthers, are united together in a ring
closely embracing the style or upper portion of the
ovary. In the photograph the ovary is seen between
the stalks of the stamens, and the style, with its two
stigmas, above the united anthers.

The union of the anthers, and their close proximity
to the style, is part of a simple and interesting
mechanism for ensuring cross-fertilisation, by the
agency of some insect visitor, which will carry the
pollen of one flower to the stigmas of another. Cross-
fertilisation is essential to most of the Flowering Plants,
and is brought about either by animal, especially
insect, visits, or by wind transference. The result of
cross-fertilisation is renewed vitality to the stock.
While self-fertilisation may be the rule in a minority
of plants, yet in the majority, as Charles Darwin
showed, continuous self-fertilisation is harmful, for
the stock weakens and the seeds tend to become
sterile, and the plant may even become totally extinct.
At the same time, many plants which are usually
cross-fertilised are capable of occasional self-fertilisa-
tion when by some accident the chance of cross-
fertilisation has been missed.

In a large number of flowers adapted to cross-

THE COMMON BELL GENTIAN 53

fertilisation, special contrivances exist which tend
to make self-fertilisation impossible. A very simple
method is the arrangement whereby the male organs
ripen and shed their pollen before the stigmas
are mature. This happens in the case of the Bell
Gentian. A close examination of a flower such as
that seen on Plate IX., or, better still, a comparative
study of several flowers in different stages of develop-
ment, some quite young and hardly open, others fully
mature, will enable us to follow the details clearly.

The Bell Gentian is fertilised by humble-bees.
A bee, visiting a young flower in search of the honey
secreted at the base of the ovary, has to push its way
through one of the spaces between the stalks of the
stamens. The anthers at this stage are quite ripe and
dehisce each by means of two long slits. They open
outwards — i.e., towards the corolla and away from the
ovary. In the young flower, the stigmas are not
mature, but the pollen is ripe ; and when a bee forces
its way between the anther stalks, it shakes a cloud
of pollen dust out of the anthers on to its own back.
Later, when the bee seeks another flower in a more
advanced condition, in which the pollen has all been
shed, but where the stigmas are mature, it deposits
some of the pollen on them, when it enters the
flower. If by any chance the pollen of a flower
should reach the stigma of the same flower, it is
usually ineffective, because the stigmatic surfaces are
not ripe.

Kerner states, however, that self-fertilisation may

54 TYPICAL FLOWERS OF ALPINE PASTURES

take place in the following manner and circumstances.
The ordinary method, described above, depends to
some extent on the occurrence of fine sunny weather,
when the flowers are mature. If the summer is wet,
bees are not on the wing and the flower remains
closed. In this case the pollen, when mature, falls to
the bottom of the bell and there accumulates. Later,
when the stigmas are ripe, the stalk of the flower
lengthens, and the closed bell, instead of pointing
directly upwards, is inverted. The pollen falls down
along the grooves inside the folded bell, and thus,
when the bell is shaken by the wind, reaches the
stigmas, and the plant is self-fertilised. This process
may also occur at night, when the flowers are always
closed.

Thus the difference in the position of the bell of
this Gentian, whether held vertically upright or
pointed earthwards, has an important biological
significance. The two positions may be observed
in almost any large patch of this plant in the Alps.

THE FRINGED GENTIANS, OR GENTIANELLAS.

We now reach our third group of blue-flowered
Gentians, the Fringed Gentians, or Gentianellas, which
possess a fringe of much-divided scales just inside
the throat of the corolla. The Field Gentian (Gentiana
campestris, Linn.), a fairly common British plant, and
the Delicate Gentian (Gentiana tenella, Rottb.), a
somewhat rare High Alpine, are the two chief repre-
sentatives of this group in Alpine Switzerland.

PLATE X.

THE FIELD GENTIAN

Both are much-branched plants bearing many
flowers, as a rule, and the sepals and petals are only
four and not five in number in each case, a character
which easily distinguishes them
from the other Alpine Gentians.

THE FIELD GENTIAN.

The Field Gentian (Gentiana
campestris, Linn.) (Plate X., Fig.
2) is a many - flowered annual
plant, with a rather peculiar
calyx. Two of the sepals are
much broader than the other
pair.

The blue fringe of scales at
the mouth of the corolla is very
conspicuous, and is deeply cut
into long, narrow segments

(Text-fig. III.). ItS probable FlG.m. -The Flower of the

object is to keep out "unbidden

guests" — that is, certain small

Showin the scale-like out-

in-

g
th

growths from the throat
of the corolla. Magnified
3 times.

insects, especially creeping
sects, which might visit the flower
in search of honey or pollen,
but are of no service to the plant as cross-pollinators.
The flower is specially adapted to bees and butterflies,
which alone are powerful enough to force aside the
fringe closing the throat of the corolla, and possess a
sufficiently long proboscis or tongue to reach the honey
secreted by the nectaries at the base of the corolla

56 TYPICAL FLOWERS OF ALPINE PASTURES

tube. The arrangement for cross-fertilisation here is
similar to that described in the case of the Common
Bell Gentian : except that both the anthers and
stigmas are ripe at the same time, or the stigmas
may mature slightly before the anthers.

The flowers of the Field Gentian, like those of the
Snow Gentian, are extremely sensitive to light. They
are usually closed in the absence of bright sunlight.

THE DELICATE GENTIAN.

The Delicate Gentian (Gentiana tenella, Eothb.)
resembles the Snow Gentian somewhat in habit. It
is a dwarf plant, with a small rosette of leaves on the
surface of the soil, from which numerous erect
branches arise, often comparatively long and leafless.
Each branch ends, as a rule, in a single flower.

Scales occur at the mouth of the corolla tube,
which are similar to those found in the Field Gentian,
but shorter and less finely divided. The fertilisation
of the flower is effected by the same agency, and
the scales appear to perform the same function as in
those found in the Field Gentian. In the Delicate
Gentian, both the male and female organs mature
simultaneously.

THE YELLOW- AND KED-FLOWERED GENTIANS.

The Swiss Alpine Gentians with yellow and red
flowers differ in many remarkable respects from the
blue-flowered species, which we have just been con-

THE YELLOW GENTIAN 57

sidering. The habit is quite dissimilar. The plants
are all perennials, and very much larger. The long,
leafy stems are rarely less than a foot high, and often
reach several feet in length. The opposite pairs of
leaves are large, and the flowers are arranged in
whorls in the axils of the higher, alternating pairs of
leaves. These plants possess a stout root-stock below
ground. Two species have yellow, and two red,
flowers. They frequently form large colonies in the
stony pastures.

THE YELLOW GENTIAN.

The Yellow Gentian (Gentiana lutea, Linn.) is
quite dissimilar in several respects from all the other
Swiss Gentians. The yellow petals are almost
entirely free from one another, and not united into a
tube. The honey is secreted from an annular swelling
at the base of the ovary, above the points of attach-
ment of the stalks of the stamens.

The Yellow Gentians are tall plants arising from
a stout root-stock below ground, which is often 2 to
3 feet in length, so that the total length of the plant
is sometimes nearly 6 feet. The upper portion of the
root-stock is the stem, which is covered with numerous
ring-like scars, marking the points of attachment of
the leaves of former summers. These root-stocks
often attain to a considerable age, forty-three years
being recorded in one case. It is also stated that the
plant is several years old before it produces any
flowers.

58 TYPICAL FLOWERS OF ALPINE PASTURES

The roots of Gentiana lutea contain a bitter
substance, which is used in medicine as a tonic.
Similar substances are also obtained from G. punctata
and G. purpurea, the roots being often collected in
Switzerland for this purpose. The Swiss also make a
liqueur, Gentian brandy, from some species.

The leaves are large and rather like those of
Veratrum album. They are about 10 inches long,
and 2 inches at their broadest. The yellow flowers
are borne from three to nine in a whorl. The petals
are free and spreading. The anthers and stigmas
mature about the same time, and so, no doubt, some
self-fertilisation takes place.

The flowers of the Yellow Gentian are regarded as
approximating closely to the primitive type of Gentian
flower, from which the blue-flowered and other
species have been evolved, largely by specialisation
to particular groups of insects. Here the petals are
free, whereas in all the other species they are united
to form bells or long tubular corollas, adapted to
fertilisation by long-tongued insects, such as humble-
bees and butterflies. The yellow colour is also a
primitive feature, whereas blue is a derived colour,
and is especially attractive to certain groups of
insects, such as bees.

The nectary is also more primitive in Gentiana
lutea than in any other species. In other Gentians it
is found at the base of the corolla — that is, farther
from the entrance to the flower and out of reach
except to those insects possessing long tongues.

PLATE XI.

The Flowers of the Spotted Gentian (Gentiana punctata, Linn.).

THE SPOTTED AND PURPLE GENTIANS 59

The various contrivances which tend to close the
mouth of a tubular corolla, such as scales at the
throat, which we have noticed in discussing the blue-
flowered species, are all less primitive devices to keep
out unbidden guests, and adaptations in favour of
particular kinds of insect visitors. Whereas the
flowers of the Yellow Gentian are open to almost all
comers and the honey is not protected. A very
different state of affairs is met with in the Field
Gentian, as we have seen, where a long, narrow tube,
closed with scales at the mouth, must be penetrated
before the honey is reached.

Again, the fact that in the Yellow Gentian the
anthers and stigmas ripen simultaneously, and not at
different periods, points to a greater degree of
primitiveness than that met with in the other species.

THE SPOTTED, PURPLE, AND HUNGARIAN GENTIANS.

The remaining Swiss Gentians with yellow and
red flowers possess many points of similarity in
habit to the Yellow Gentian. The Spotted Gentian
(Gentiana punctata, Linn.) (Plate XL) has yellow
flowers, ornamented with reddish-brown or purple
spots. The petals are all united to form a bell, and
the calyx is also bell-shaped, with five very dis-
similar teeth. The Purple Gentian (Gentiana purpurea,
Linn.) has also a bell-shaped corolla, red outside and
yellowish within, but the calyx is incompletely
united on one side. The Hungarian Gentian (Gentiana
pannonica, Scop.), which is less abundant in Switzer-

60 TYPICAL FLOWERS OF ALPINE PASTURES

land than the preceding species, has a dark red bell
corolla, spotted with black, and a bell-shaped calyx,
the teeth of which are bent backwards.

In the union of the petals into a bell-shaped
corolla, in the position for the nectaries at the base
of the corolla, and in the colour of the flowers — the
evolution of red or purple from yellow — these flowers
show a marked advance on the primitive features met
with in Gentiana lutea. The male and female organs
also ripen at different periods, and not simultaneously,
as in the Yellow Gentian.

CHAPTER III

TYPICAL FLOWERS OF THE ALPINE PASTURES

(continued]

THE SOLDANELLAS, THE PRIMULAS, THE ANDROSACES,
THE SAXIFRAGES, THE CAMPANULAS, AND THE
RAMPIONS.

IN the present chapter we will consider some of the
other genera characteristic of the Swiss pastures,
beginning with the members of the Primrose family —
the Soldanellas, Primulas, and Androsaces, and then
passing to the Saxifrages, the Campanulas, and the
Eampions.

The Primrose family (natural order Primulacese) is
very well represented in the Alpine zone, one of the
most interesting genera being the Soldanellas.

THE SOLDANELLAS.

The Soldanellas, or, as the Germans call them,
" Alpenglockchen " (the little bells of the pastures), are
among the earliest flowers to bloom in the pastures
when the snow begins to melt ; they are often extremely

62 TYPICAL FLOWERS OF ALPINE PASTURES

abundant. They also occur on the margins of woods,
flowering from the first sign of spring until June or
July.

The genus is not found in Britain. In the Swiss
Alpine zone there are two common species : the
Alpine Soldanella (S. alpina, Linn.) and Soldanella
pusilla, Baumgarten, which we may call the Small
Soldanella. The former bears two to three flowers
on each flowering shoot (Plate XLIL, Figs. 2 and 3),
the large, pale lilac flowers being bell- shaped and
drooping. The petals are deeply fringed, the fringe
extending for half the entire length of the corolla. In
the Small Soldanella, the fringe is shorter, and does
not exceed one-third of the length of the petals, and
only one flower is, as a rule, borne on each flower-
stalk. The bell-like corollas are here usually violet
in colour. The leaves of both species are thick, dark
green, heart-shaped structures, mounted on long stalks,
which arise from the stem just below the surface of
the ground. The leaves persist throughout the winter,
and are recumbent on the soil. Below ground there
is a fairly stout stem giving off numerous roots.

Like the Crocus, which we shall discuss in a later
chapter devoted to meadow flowers, the Soldanellas
often bloom before the winter's snow has melted.
A photograph of Soldanellas flowering in the snow is
shown on Plate XII. The flower matures gradually
during the winter months. At first it is quite a
small object, down among the bases of the leaves.
Long before the snow has begun to melt, the plant

PLATE XII.

The Alpine Soldanella (Soldanella alpina, Linn.) Flowering in the Snow.

[To face p. 62.

THE SOLDANELLAS 63

is at work. The energy for growth is supplied by
the reserve food material stored up during the previous
autumn, not only in the underground stem, but in
the leathery, evergreen leaves themselves. As the
flower matures, the flower-stalk is pushed up through
the snow. It was formerly thought, on the authority
of the great Austrian naturalist, Kerner, that the
plant had the power of melting the snow and forming
a little dome-shaped cavity above the flowers. It
was believed that the heat necessary to melt the
dome was derived from the respiration (p. 11)
accompanying growth, that is, the conversion of the
raw food material into the substance of new tissues.
We are not, however, quite sure now whether this
plant really has the power of melting the snow above
it. No doubt dome-shaped cavities often occur
above the plant. One of these is seen cut across, in
the middle of the photograph on Plate XII., and
rather to the left-hand side, on the margin of the
snow. It is also a matter of common observation
and experiment, that young developing flower-buds
do set free considerable heat, their temperature rising
sometimes as much as 2° to 3° Centigrade above that
of the atmosphere. There was, therefore, much
inherent probability that Kerner's account was correct.
However, it is now doubted whether the heat set free
by the developing plant is sufficient to melt the snow
above it, and cases have been observed where there
is no dome-shaped structure to be found. What
does appear to be clear is, that once a dome-shaped

64 TYPICAL FLOWERS OF ALPINE PASTURES

structure is formed above the plant, to whatever
cause we may attribute its origin, the flower-stalk
grows rapidly and pierces it before the sun has melted
it away from above. The flower-stalks have thus,
under certain conditions at any rate, the power of
piercing the snow. It will be observed that the stalk
of the individual flower is arched, and that the bell-
shaped corolla droops from it. In the passage
upwards to the light, the bell is thus saved from
injury, for it is the arched flower-stalk which actually
bores through the snow.

We have here a good example of a common
feature among Alpine plants — the tendency to flower
at the earliest possible moment in the spring. The
summer season is very brief, and the period before
the plant is again buried in the snow all too short
for the work which lies in front of it.

The leaves, in which, as we have seen, are stored
the reserves for spring growth, are exhausted and
perish after the plant has reached the light. New
leaves are formed later in the season, and in them are
laid by the fresh reserves for the following winter.

The flowers of the Alpine Soldanella are worth
examining closely (Text-fig. IV.). The five stamens
spring from the corolla, as in all members of the
Primrose family, and when mature are firmly pressed
against the style or upper portion of the pistil. The
pollen, even when mature, is thus closely held between
the inner surface of the anthers and the style. When,
however, an insect, visiting a flower in search of the

THE SOLDANELLAS

honey secreted by the nectary at the base of the
corolla, inserts its proboscis between the style and
the obliquely placed stamens, the moment the former
is separated from the latter a shower of pollen falls
on to the insect's head. This, as likely as not, is

FIG. IV.

1 and 2. The Alpine Soldanella (Soldanella alpina, Linn.).

1. Longitudinal section of the flower. 2. Section across the flower on the level
of the scales ; *, scales ; n, nectary ; k, calyx. Fig. 2 shows how the
ring of five scales closes the entrance to the lower portion of the flower.

3 and 4. The Small Soldanella (8. pusilla, Baumg.).

8. Longitudinal section through the flower. 4. Section across the flower in
the plane of the anthers. (All after Schroeter.)

carried to another flower of the same species, and
placed on the stigma at the tip of the style as the
insect enters the flower.

The interior of the flower of an Alpine Soldanella
can only be approached by flying insects. The
drooping position of the bell, with mouth directed

66 TYPICAL FLOWERS OF ALPINE PASTURES

downwards, and possibly also the fringed edge, bars
access to insects crawling upwards from below by
means of the flower-stalk. But, in this plant, as it
were, to make doubly sure that only insects with a
long and stout proboscis shall reach the honey, and
incidentally cross-fertilise the plant in the manner
already indicated, the nectary is also protected, or, as
it is called, " concealed." If we examine a flower of this
species more closely (Text-fig. IV., 1 and 2), we shall
find that just below the level of the insertion of the
stamens on the corolla, and alternating with them,
there are five membranous scales projecting across
the base of the bell (Text-fig. IV., 2). These scales
roof in a little chamber above the ovary, and act like
trap doors, having to be raised or pressed down by
the insect before it can get at the honey. Thus this
plant is specially adapted to cross-fertilisation only
by a particular class of insects which possess a long
and strong proboscis.

In the Small Soldanella (Text-fig. IV, 3 and 4)
these scales are absent, and the tips of the stamens
are not produced into horns, as in the Alpine
Soldanella.

THE PRIMULAS.

We are familiar in Britain with several Primulas,
especially the Primrose, Cowslip, and Oxlip. All of
these also occur in Switzerland, but the Oxlip is the
only one which is commonly met with, within the
Alpine zone, at comparatively low elevations. In

THE PRIMULAS 67

addition to these three species, there is another less
commonly known British plant, the Bird's-eye
Primrose, which occurs in the north of England,
especially on the hills. Of all the Swiss Alpine
Primulas, this is by far the most abundant.

In Alpine Switzerland there are also several
Primulas of great interest and beauty which do not
occur wild with us. Of these, the Auricula, the original
parent of our cultivated Auriculas, stands first. It has
yellow flowers and characteristic leaves, which are
thick, fleshy, and dusted with a white waxy powder.
It thus stands in strong contrast to the Primrose and
Cowslip, so common in this country, which have green
wrinkled leaves.

Next we have the rare Alpine, the Long-flowered
Primula, with leaves which are green and wrinkled
above, but covered below with a waxy powder like
that found on both sides of the leaf of the Auricula.

Lastly, we have five other Alpine species with
violet or rose-coloured flowers, and leaves which are
green and not powdery.

We will commence with the Oxlip (Primula
elatior, Jacq.). The individual flower-stalks are all
mounted on a long common flower-stalk, the whole
forming an umbel, in the manner which we have
already noticed in the case of the Narcissus-flowered
Anemone. This arrangement of the flowers is
characteristic of the Primulas, and occurs even in the
Primrose (P. vulgaris, Huds.), where, however, the
common flower-stalk is short, and sunk beneath the

68 TYPICAL FLOWERS OF ALPINE PASTURES

level of the soil. For this reason it is often over-
looked.

The Oxlip is a good plant on which to observe
the very interesting adaptation to cross-fertilisation
known as heterostylism. If we slit open with a
needle or pin the corollas of a number of flowers, we
shall be able to distinguish two types. One set will

FIG. V. — Diagrammatic Views of the Heterostyled Flowers
of a Primula.

1. Flower with short style and high stamens.

2. Flower with long style and low stamens.

be found to possess a long style (the prolongation of
the ovary) extending almost to the throat of the
corolla tube (Text-fig. V., 2), and five short stamens,
seated on the corolla itself, near the base of the
corolla tube.

Other flowers will be found to be the exact
opposite (Text-fig. V., 1). The style here is very short
and the stamens are very long, and occupy the same

PLATE XIII.

The Bird's-eye Primrose (Primula far inos a, Linn.).

[To face p. 68.

CROSS-POLLINATION IN PRIMULAS 69

relative position as the head of the style in the first
set of flowers. The great naturalist, Charles Darwin,
showed that these differences constitute a special
mechanism or contrivance to ensure cross-fertilisation.
For example, a bee, visiting a long-stamened flower,
would get dusted with pollen around the base of its
proboscis or tongue, and this pollen could not fail to
be deposited on the stigma of the next long- sty led
flower it visited. Darwin found by experiment that
a full yield of seed is only obtained when the pollen
from a flower with long stamens is transferred to the
stigma at the top of a long-styled flower, or when the
pollen from short stamens is transferred to a flower
with a short style. This is " legitimate pollination."
If by any chance illegitimate pollination takes place —
that is, from a short stamen to a long style, or vice
versa — the seeds that result are few, and more or less
sterile. A similar adaptation is met with in other
Alpine Primulas, including P. farinosa, and also in
one of the Androsaces. In some Lowland plants, for
example in Lythrum, three kinds of flowers occur with
different lengths of stamens and styles.

The Bird's-eye Primrose, Primula farinosa, Linn.
(Plate XIII), is so called because the pale lilac flowers
have a yellow " eye " or ring round the throat of the
funnel-shaped corolla. This plant is one of the
earliest spring blossoms in the damper pasturages,
where it flowers in countless millions. The leaves
are green and smooth above, but are covered below by
a white, mealy wax or bloom.

70 TYPICAL FLOWERS OF ALPINE PASTURES

Kerner states that the wax on the lower side of
the leaf tends to protect the plant by hindering the
access of water to the minute pores or stomata which
exist on the lower surface of the leaves of the rosette,
placed close to the ground. Were the leaves to
become saturated with water, the gases of the atmo-
sphere would no longer have free access to the leaf,
and thus the whole internal economy of the plant
would come to a standstill. The presence of a layer
of wax prevents the lower surface from becoming
easily wetted. This can be demonstrated by immers-
ing a leaf in water for a few minutes. It will then be
found that whereas the upper surface, where there is
no wax, is easily wetted, the lower remains quite dry.

The Auricula (Primula auricula. Linn.) (Plate
XIV., Fig. 1) is, as we have said, one of the original
parents of our cultivated Auriculas. The Primulas,
as a whole, are very apt to form hybrids — that is,
crosses between, not two individuals of the same
species, but of two different species. Our garden
Auriculas are all derived from a cross between P.
auricula, Linn., and P. hirsuta, All., which gives
a hybrid (P. pubescens), and this is the stock from
which yet other hybrids can be obtained.

The hybrid, as we should expect, combines the
characters of both parents. The corolla limb is partly
yellow and partly red or violet, the yellow colour
being derived from the Auricula, the red or violet from
the other parent. The question of the inheritance of
characters in hybrids is a very interesting one, and

PLATE XIV.

THE AURICULA "71

much research has recently been done on this point.
It would, however, involve a too lengthy botanical
preface to permit us to discuss it here.

The Auricula flourishes chiefly on limestone soils,
in fairly dry situations. It is extremely abundant in
June, for instance, on the terraces of the natural rock
garden of the Engstlen Alp (Canton Berne), where
the limestone rocks are weathered into fantastic
shapes, owing to their solubility and lack of resistance
to decay under the influence of atmospheric agencies.
This pasturage resembles an artificial rock garden
covering thousands of acres in extent, and in the
crevices of the decayed limestone crags many an
Alpine plant of interest flourishes in addition to the
Auricula.

The rosette of leaves of the Auricula placed close
to the ground is well protected against the danger of
undue evaporation of moisture, under the hot suns of
early summer, by the waxy covering or mealy bloom,
with which both the upper and lower surfaces are
thickly dusted. The leaves themselves are really
green, though the colour is masked by the mealy
powder. In addition, there are special tissues for
water storage within the substance of the leaf, as its
thick semi-succulent nature would lead one to imagine.
It will be noticed that the waxy bloom is not confined
to the leaves, but also occurs both on the common
and the individual flower-stalks, as well as at the
throat of the corolla itself.

The Long-flowered Primula (Primula longiflora,

72 TYPICAL FLOWERS OF ALPINE PASTURES

All.) is a rare plant, occurring chiefly in the Enga-
dine and in the Zermatt and Saas valleys. It is
remarkable for the length of the corolla tube as
compared with the calyx. It is believed to be the
only European Primula which does not possess long-
styled and short-styled flowers (see p. 68). The
leaves are similar to those of the Bird's-eye Primrose,
the lower surface being covered with a waxy powder.
Of the other species of Swiss Primulas, there is
little of interest to relate. Their leaves do not possess
a mealy powdering on either surface, and their rose-
or violet-coloured flowers have relatively short, common,
and individual stalks. The Hairy Primula (P. hirsuta,
All.) and the Entire-leaved Primula (P. integrifolia,
Linn.) are the commoner species.1 In the former the
leaves are thickly covered with sticky glandular hairs,
while in the latter they have only a few such hairs on
the margins. The Hairy Primula has strong-scented
flowers, and in the autumn the glandular hairs on
the leaves turn bright red in colour. It flourishes
on bare rocks, especially in granitic and gneissic
regions. As we have already remarked, it is one of
the parents of the cultivated Auricula.

THE ANDROSACES.

Closely allied to the Primulas are the Androsaces,
also members of the order Primulacese, and charac-

1 Three other species, P. viscosa, All., P. oenensis, Thorn., and P.
glutinosa, Wulf, are found in Canton Graubunden. They are dis-
tinguished with difficulty from the above and from one another.

THE ANDROSACES 73

teristic plants of Alpine habitats. The genus does
not occur in Britain. These plants are well known
to horticulturists for their aversion to the plains, it
being very difficult, if not impossible, to cultivate some
of them in England. In the Swiss Alps there are
some eleven species, all much alike in external form.
Several of them are confined to the High Alpine
region, for the genus as a whole is strictly Alpine.

The habit (Plate XIV., Fig. 2) is quite like that of
a Primula ; only, the plant as a whole is much smaller.
There is, just above the ground, the same little rosette
of leaves, from which springs a common flower-stalk
ending in an umbel of flowers. The corollas resemble
those of the Primula in shape. The flowers are,
however, distinguished by the fact that between
each of the five lobes of the petals, we find a little
scale, not unlike that which we have already noticed
in the case of some Gentians. These five scales tend
to narrow the entrance to the mouth of the corolla.
The Androsaces, except Vital's Androsace, do not
possess long-styled and short-styled flowers (p. 68).

In the High Alpine species, the umbel is frequently
reduced to a single flower. This is a marked feature
in plants, which in the lower Alpine regions possess
inflorescences of several flowers. As we ascend higher
and higher, the number of flowers in the inflorescence
decreases, and finally only one remains. This is the
case in the Saxifrages, the Harebells, and many other
genera besides the Androsaces.

The flowers of most of the Swiss species are white,

74 TYPICAL FLOWERS OF ALPINE PASTURES

often with a yellow or red "eye." In one species,
however (A. carnea, Linn.), they are rose coloured,
and in another (A. vitaliana, Lap., Plate XXXVII.,
Fig. 1) they are yellow. It often happens, however,
that the flowers of species which are usually white
may be rose-coloured.

The commonest and most widely distributed
species is Androsace chamcejasme, Willd., the Dwarf
Androsace (Plate XIV., Fig. 2), in which the whole
plant except the flowers is covered with long hairs,
especially on the edges of the leaves. The Obtuse-
leaved Androsace, A. oUusifolia, AIL, and the Red-
flowered Androsace, A. carnea, Linn., are also com-
mon, especially in western Switzerland. They have
short hairs on the leaves and flower-stalks, but are
otherwise very similar in habit.

THE SAXIFKAGES.

The Saxifrages (natural order Saxifragacese) are
among the most characteristic of Alpine plants.
Though the individual flowers are often rather small
and not very showy, they are rendered conspicuous
by the mass of bloom borne by each little colony of
these plants on some rocky shelf (Plate XV.).

Britain is rich in Saxifrages, possessing no
fewer than thirteen species, some confined to our
highest mountains, others common in the meadows
and woods of the Lowlands. Two of these occur
in Alpine Switzerland, where there are also to be
found about eighteen other species. Of these eighteen,

PLATE XV.

THE SAXIFRAGES 75

some seven are confined to the High Alpine region,
and these we will reserve for a later chapter.

The habit of many of these Saxifrages is quite
typical of that of the majority of Alpine plants. The
plant is, so far as possible, buried in the scanty soil,
especially the roots and the very short stem. Above
the soil, one or more compact rosettes of leaves are
pressed close to the surface of the ground. These char-
acteristic rosettes often form a beautiful leaf-mosaic
(Plate XVI., Fig. 1), each leaf being arranged in
regard to its neighbours so as to cut off as little light
from it as possible. The really conspicuous part of
the plant is the flowering shoot, which often bears
several leaves and numerous flowers, and may vary
from a few inches to a foot or more in height.

The individual species are not, as a whole, very
dissimilar, and in some cases may only be distinguished
with difficulty from one another. These plants are
thus perhaps less interesting than those of many
other Alpines.

The Alpine Saxifrages may be divided into two
groups, the first of which, containing some six species,
has undivided leaves, bearing a row of conspicuous,
white chalk-glands along the margins. These glands
are clearly seen on Plate XVI., Fig. L

In Saxifraga aizoon, Jacq. (the Evergreen Saxi-
frage) (Plate XVI., Fig. 1), the leaves are strap-shaped,
and each chalk-gland lies in a little notch on the
edge of the leaf. The glands themselves are really
invisible to the naked eye. All that we see here are

76 TYPICAL FLOWERS OF ALPINE PASTURES

the little heaps or accumulations of calcium carbonate
or chalk, which mark the position of the glands.
We have already shown (p. 10) that the leaves of
all plants have numerous but very minute pores
or openings, by which the atmosphere has free
entrance into the substance of the leaf itself,
and by which the gases evolved by the internal
mechanism of the leaf pass back to the atmo-

FIG. VI.— Section through a Chalk-gland on the edge of a Leaf
of a Saxifrage.

to, the water stomata ; g, the chalk-gland ; ».&., the vascular bundle of
the leaf. Highly magnified.

sphere. In most plants these pores or stomata
can be opened and closed. The chalk-glands or
water stomata of the Saxifrages, on the contrary, are
differently constructed, and remain always open. In
order to prevent an undue amount of water- vapour
escaping from the leaves — a matter of great
importance to plants which, like the Saxifrages, grow
in dry situations with but a limited supply of moisture
in the soil — a small quantity of calcium carbonate in

PLATE XVI.

I!
•s-i

THE SAXIFRAGES 77

solution is secreted by the leaf, and this exudes at the
pores. As the water evaporates, the chalk crystallises
out, and blocks the mouth of the pore. In warm
weather and in direct sunlight, the opening of the
pore is almost completely closed in this manner. At
night, however, when the temperature is lower, more
water is secreted, which dissolves some of the calcium
carbonate, and thus a freer passage for the gaseous
exchange is afforded. In this way the chalk-glands
control the rate at which the leaves lose water to
the atmosphere.

It is a curious fact that Saxifrages with chalk-glands
may often be found growing in abundance on rocks
composed of granite or on schists, which contain very
little or no lime. Yet by means of their roots these
plants can obtain from the soil sufficient lime or
chalk to render the incrustation-mechanism of the
leaves quite efficient. The roots of plants have a
peculiar property, known to botanists as "selective
capacity," which enables them to gather in or absorb a
sufficient quantity of a substance in the soil, even
when it exists only in extremely minute quantity.
Thus plants can appropriate, if they need it, a
considerable quantity of one particular substance, to
the exclusion of others. For instance, sea-weeds
absorb from sea- water sufficient phosphorus — one of
the essential elements to the life of all plants — though
the amount of phosphorus in sea-water is so exces-
sively small that it is quite impossible to estimate it.
Closely allied to the Evergreen Saxifrage is the Thick-

78 TYPICAL FLOWERS OF ALPINE PASTURES

leaved Saxifrage, S. cotyledon, Linn., which is the
largest and most magnificent species occurring in
Switzerland. It is not, however, common, except
on granite rocks in Transalpine Switzerland and the
St Gotthard region. It occurs also on the slopes
of the Brevent, above Chamonix, where the tall-
branched flowering shoots, 2 feet high, are con-
spicuous objects. Its leaves possess chalk-glands
like those of S. aizoon.

The Purple Saxifrage (S. oppositifolia, Linn.) is a
British plant of frequent occurrence in our moun-
tains. In Switzerland it is common hi stony and
rocky places, especially in the High Alpine zone,
and not infrequently is much in evidence in the
Alpine region also. It is easily distinguished from
all other Swiss Alpine Saxifrages by the solitary
purple flowers, and the very small evergreen leaves
placed in crowded pairs opposite one another. Each
leaf has a single chalk-gland, situated at the blunt,
somewhat thickened tip. The small creeping stems
are much branched, and form a cushion composed
of little tufts of leafy shoots (see p. 186) which seldom
rise more than an inch above the surface of the
ground. Only one flower is borne at the end of
each branch, and these flowers, huge in size in com-
parison with the leaves (Plate XXXVI., Fig. 1),
form a conspicuous advertisement to attract the insect
world, especially butterflies. Thus cross-fertilisation
is ensured.

There is also another but much less frequent

THE SAXIFRAGES 79

Alpine Saxifrage with purple flowers and opposite
leaves. This is the Two-flowered Saxifrage (S. biflora,
All.). Here, however, the flowers are not solitary,
but borne two to five together, and the leaves are more
distant from one another. Otherwise the resemblance
to S. oppositi/olia is marked.

We now turn to another series of Saxifrages, in
which, instead of a single chalk-gland existing at the
tip of the leaf, as in S. oppositi/olia and S. biflora, or
a line of glands occurring all round the edge, as in S.
aizoon and S. cotyledon, we find only a limited number
of chalk-glands, usually situated near the tip. The
Glaucous Saxifrage, S. ccesia, Linn., is the commonest
of these species, S. diapensioides, Bellard, being rarer
and confined to the Canton Valais. The leaves of S.
ccesia are borne in rosettes of a bluish - green tinge,
and are bent or arched backwards almost from the
base. Otherwise this plant is in no degree remark-
able as compared with other Saxifrages.

The species of Saxifrage in which chalk-glands are
absent from the leaves are more numerous, but are
not, for the most part, deserving of special notice.
The greatest contrast which they present is chiefly in
the leaves. The Yellow-flowered Saxifrage (Saxifraga
aizoides, Linn.), a frequent British Alpine, is also
common, especially in damp places, in Switzerland.
The leaves are narrow, rather thick, quite smooth and
shining, and are not arranged in rosettes. The flowers
are worth noticing on account of their red pollen and
yellow honey nectaries. The sepals and petals are

80 TYPICAL FLOWERS OF ALPINE PASTURES

both yellow and about the same length, so that at
first sight the flower appears to have ten petals.

The Star-leaved Saxifrage (S. stellaris, Linn.) has
thin egg-shaped or oblong leaves, toothed at the top,
and borne in tufts near the ground, while the flower-
ing shoots are entirely destitute of leaves. A small
flower-leaf or bract is found at the base of each flower-
stalk. The petals are white, two yellow spots
occurring on each. It is a British plant, not
uncommon in our mountains.

The Saxifrages last mentioned, and also the Eound-
leaved Saxifrage (S. rotundifolia, Linn.), are members
of this genus which love damp, shady spots, and thus
differ in their choice of habitat from many of the
other Alpine species, which flourish on dry stony
ground, or on exposed rocky ledges. This difference
is indicated by their leaves, which are much larger
and thinner.

The Eound-leaved Saxifrage (S. rotundifolia, Linn.)
has large, thin, heart- or kidney-shaped leaves, lobed
and toothed, and white petals spotted with yellow
and red. It is often a foot or more in height
and much branched. The numerous flowering shoots
bear leaves similar to those below, but smaller. The
flowers of the Kound-leaved Saxifrage are interesting
from the manner in which the stamens shed their
pollen, one by one — a peculiarity which is, however,
shared by many other plants, including the Grass-of-
Parnassus (p. 216). When the flower first opens, the
stamens are as yet unripe, and they bend backwards

THE SAXIFRAGES 81

with the petals. Then in a day or so, one of the ten
stamens becomes erect and moves in towards the
centre of the flower, where it remains until it has
shed its pollen on to the back of any insect that
may visit the flower in search of honey. At the
end of twenty-four hours or more, it bends back to
its original position, and not till then does the next
stamen begin to go through the same performance.
After all the ten stamens have shed their pollen
one by one in this way, and not until then, the two
stigmas of the ovary mature. Thus the female organs
of the flower cannot be fertilised by the pollen of the
same flower.

As a typical example of the Saxifrages inhabiting
dry stony places, we may instance the Rough
Saxifrage (Saxifraga aspera, Linn.) (Plate XV.), which
is widely distributed in the Alps. It has a very near
relative, the Moss-like Saxifrage (S. bryoides, Linn.),
which is perhaps only a High Alpine variety of the
former. The flowering shoots of the Eough Saxifrage
are leafy, and the thick leaves are rough with long
hairs.

Further, many of the leaves bear large leaf-buds
in their axils, which in the High Alpine (S. bryoides)
are as long as the leaves themselves. The flowers of
this species are also interesting botanically, from the
fact that the ovary is superior or free from the calyx
tube, whereas in the majority of Saxifrages the ovary
is more or less united with the calyx tube.

The Eough Saxifrage, like all the other Swiss

82 TYPICAL FLOWERS OF ALPINE PASTURES

Alpine species, except Saxifraga controversy Stern-
berg, is a perennial plant. The latter species is quite
exceptional in being an annual.

The compact form of the colony is especially
noticeable here, and is due to the fact that in the
Alps a great struggle for existence is everywhere in
evidence. Other plants tend to intrude into a colony
of Saxifrages or other Alpines, and to rob them of
their possession of the soil. In fact, what may be
not unfairly termed plant slums, comparable as
regards crowding to the worst slums of our great
cities, though infinitely more beautiful, are to
be seen on every hand in the Alps. Wherever
the ground is unoccupied, there is strong compe-
tition among the neighbouring plants to seize upon
it and to establish themselves, to the exclusion
of others. The competition, however, is not only for
the possession of the soil, but also for light and
air — matters of equal importance to the plant.
The photograph on Plate XV. shows a bank of the
Rough Saxifrage, and is a typical example of an Alpine
plant slum. The colony of the Saxifrage is here
holding its own very successfully, although other plants
have intruded into it. At the right-hand corner
flowers of the Eock Catchfly (Silene rupestris, Linn.)
(see Plate XVI., Fig. 2) are evident, and in this region
a struggle between these two plants is in progress.
Towards the left-hand side, two plants of a Rampion
are seen with heads of flowers borne on long stalks.
These are being overwhelmed by the advance of the

THE CAMPANULAS, OR BELL-FLOWERS 83

Saxifrage colony. Their leaves are arranged in the
form of rosettes, pressed close to the ground, and it is
absolutely essential for the welfare of the plant that
the leaves should remain fully exposed to the light
and air. In this case, however, they are being fast
engulfed by the advancing colony of the Saxifrage
with its tall growth of flowering shoots, which form a
miniature forest.

THE CAMPANULAS, OR BELL-FLOWERS.

The Campanulas, or Bell-flowers (natural order
Campanulacese), are very much in evidence in the
Alps, and add appreciably to the strikingly large
number of blue-flowered plants in that region.

The common British Harebell (Campanula rotundi-
folia, Linn.) (Plate XVII., Fig. 1) merits the dis-
tinction of being one of the most abundant of all
Alpines in every sort of locality within this zone.
In the Davos Valley, for instance, the intense blue
of its flowers is noticeable, both in the meadows,
on the higher and lower pasturages, and on the
margin of the Pine forests ; in fact, almost every-
where.

The specific name rotundifolia, or round-leaved, is
often regarded as a misnomer, for the leaves visible at
the time of flowering are all long and narrow. The
first-formed leaves of the young seedling, or cotyledons,
as the botanist terms them, of which there are two,
and a few of the leaves which succeed them are,
however, rounded or heart-shaped. The cotyledons

84 TYPICAL FLOWERS OF ALPINE PASTURES

of the Harebell thus differ entirely in shape and form
from the mature leaves of the adult plant, and this is
also the case in many other plants. It has been found
that in the Harebell these leaves are really shade-
leaves (see p. 247) adapted
to the conditions which
prevail when the young
seedling is forcing its way
up to the light between its
tall neighbours in whose
shade it starts its existence.
If a Harebell is grown from
the seedling stage through-
out the whole summer in a
really shady place, all or
many of its leaves may be
heart shaped (Text - fig.
VII.). The long, narrow
leaves, on the other hand,
with which we are more
familiar, are sun - leaves,
. VII.-A Plant of the Round- adapted for full exposure to

leaved Campanula (C. rotundi- the Summer Sunlight.
folia, Linn.), grown in diffuse T . . P „

light. All the leaves are cordate, In many Alpine Valleys,

c£Ud!) see(Weaves* (After as at Saas, another species,

Campanula Scheuchzeri,

Vill., named after the celebrated Swiss geologist of
Zurich, occurs in place of our British Harebell.
Scheuchzer's Bell-flower is not a British plant,
although it is so like our Harebell. The flowers

PLATE XVII.

FIG. 1. — The Harebell ( Campanula rotundifolia, Linn.).

FIG. 2. — The Mont Cenis Campanula (Campanula cenisia, Linn.).

[To face p. 84.

THE CAMPANULAS 85

are larger, and the flower-stalks less branched, and
bearing only one to five flowers on each plant. The
flower-buds droop in this species, whereas in the
Harebell they are almost erect.

The flowers of either species are worthy of
examination. The drooping position of the corolla,
the mouth being directed downwards, serves both to
protect the pollen and to prevent the entrance of
unbidden guests (see p. 275) in the shape of crawling
insects, which serve no purpose useful to the plant.
For, to most Alpines, though not all, cross-fertilisa-
tion by means of insects, which carry the pollen or
male generative dust from one flower to fertilise the
female organs of another, is quite essential. In the
case of the Bell-flowers and the Rampions, we shall
find, if we examine the flowers, that a special
mechanism, termed the "mechanism of the stylar
brush," exists, which is adapted to ensure cross-
fertilisation, and also guards against the 'possibility
of self-fertilisation. Thus we see that an intimate
relationship and interdependence exists between
these plants and members of the insect world. We
shall later discuss other examples (p. 268).

If we examine an unopened head of a very young
flower (Text-fig. VIII., 1), dissecting away the corolla,
we shall find the five anthers of the stamens com-
pletely covering the style. The anthers at this stage
are united at their margins, though their stalks at the
base remain free. The nectar or honey, which attracts
insects, is secreted by the ovary, which is completely

86 TYPICAL FLOWERS OF ALPINE PASTURES

covered in by the ring of united anthers. The
anthers ripen before the ovary, and shed their pollen
on the inner side, on to the style, which in this region
is studded with little hairs, called collectively the
stylar brush, to which the pollen adheres.

FIG. VIII.— The Stamens and Carpels of Scheuchzer's Bell-flower (Campa-
nula Scheuchzeri, VilL), illustrating the mechanism of the stylar brush.
Enlarged.

a = anther ; /= filament ; * = style ; 6 = stigma.

1. United anthers surrounding the style in the unopened flower.

2. The anthers separating and curling backwards.

3. The empty anthers coiled in tight spirals, the style, with the stylar brush,

bearing pollen, and the three stigmas beginning to unfold.

Let us now examine a flower which is just opening
(Text-fig. VIII., 2). We find the style has begun to
lengthen, and the anthers, having shed their pollen
on to the brush, are bending away from one another.
The separation and downward curvature of the
anthers occur suddenly, if the slightest touch is
given to the stamens when the flower is just opening.
Thus, if an insect visits the flower at this stage, it is

THE STYLAR BRUSH MECHANISM 87

pretty sure, when seeking the nectar, to come in
contact with the stylar brush, and to dust its head
with pollen, which it carries to another flower. If
the anthers have not all begun to curve, the touch of
the insect proboscis will cause them to bend
backwards.

In a still older flower, the style, with the brush, has
greatly elongated and further has opened at the tip
into three little flaps, which curve slightly backwards
(Text-fig. VIIL, 3). It is on the upper or newly
exposed faces of these flaps that the stigmatic surface
lies, on which the pollen from another flower is
deposited by an insect visitor. The pollen fertilises
the ovules in the ovary below. The five stamens at
this stage are coiled into tight spirals.

We see that by this arrangement self-fertilisation
is almost impossible. The pollen on the stylar brush
cannot reach the stigmatic surfaces above it, except
by insect agency, and an insect visiting a flower is
likely to touch the stigmas with pollen brought from
another plant, as it enters the flower; while, as it
leaves, the projecting flaps prevent the pollen of the
same flower being deposited on the stigmatic surfaces.

The mechanism of the stylar brush is found not
only throughout the order Campanulacese, but also in
the very large family of Composites. It is, however,
most favourably studied in the Bell-flowers on account
of the comparatively large size of the organs.

If we should find ourselves among the Alps in late
summer or in autumn, the fruits of the Campanulas or

88 TYPICAL FLOWERS OF ALPINE PASTURES

Bell-flowers will, on examination, prove instructive
objects. The fruit has the form termed a capsule,
and contains many seeds. Just as the flower is
directed mouth downwards, so the fruit is inverted.
The base of the capsule becomes the top and its
apex the bottom of the fruit. The capsule remains
attached to the plant, and instead of
the walls of the ovary or capsule
splitting apart and thus exposing the
seeds, as is the case in many other
plants, for instance, the Violet (p. 162),
three to five small triangular flaps or
valves will be found near the base —
that is, at the top of the inverted
fruit. These flaps are very sensitive
to moisture, closing in wet weather,
and curling backwards when the air
js <jry. When the seeds are ripe
and the air * the valves curl

through which the backwards, and the seeds are shaken

seeds escape. ' .

out or the capsule by the wind,
through the triangular clefts, often with considerable
force, and they are thus spread to a considerable
distance from the parent.

We have so far only discussed Campanula rotundi-
folia and C. Scheuchzeri. There are, however, some
seven other species to be found in the Alpine zone,
and one occurring rarely in the High Alps. None of
the British species, with the exception of the Harebell,
are found above 5,000 feet in Switzerland. Many of

PLATE XVIII.

THE CAMPANULAS 89

them, however, occur in Lowland and Subalpine
Switzerland, where other fine Bell-flowers are also
conspicuous.

Another Alpine species, which is very common, is
the Bearded Campanula (C. barbata, Linn.) (Plate
XVIII., Fig. 2). It is frequent in the pastures and
in shady spots in July. In contrast to the intense
deep blue colour of Campanula rotundifolia, and
especially C. Scheuckzeri, the bell of the Bearded
Campanula is of the most delicate shade of pale
Cambridge blue. Between each of the calyx-lobes
there is a little triangular appendage, turned back-
wards against the base of the bell. The edge of the
lobes of the corolla is also fringed with a beard of
hairs, hence'the specific name " barbata." These hairs
project over the mouth of the inverted bell (Plate
XVIII., Fig. 2), and it seems probable that their
function, like that of the five little triangular, back-
ward^ directed appendages of the corolla, is to
prevent the entrance into the bell of small creeping
insects in search of nectar, which might otherwise
crawl along the flower- stalk and rob the honey.
Such unbidden guests (p. 275) would be of no ser-
vice to the plant, for they would probably not visit
another flower of the same species, and thus no cross-
fertilisation would result.

The three Campanulas above described are the
commoner Alpine species. C. cenisia, Linn.1 (Plate
XVII., Fig. 2), is a somewhat rare High Alpine form,

1 The Mont Cenis Campanula.

90 TYPICAL FLOWERS OF ALPINE PASTURES

occurring on rocky places at Mattmark and elsewhere
at an elevation of about 7,000 to 10,000 feet. Each
stem bears only a single erect flower, the corolla of
which is deeply divided. It is interesting to com-
pare the dwarf habit and solitary flowers of
this species with the larger, much-branched flower-
ing shoots of the Campanulas growing at lower
elevations.

The rarest of all the Swiss Bell-flowers is Cam-
panula excisa, Schleicher, confined to a few valleys in
the cantons of the Valais and Tessin. It may be
collected in the Saas Thai, on the Simplon, in the
Binnen Thai, and in a few other localities. The
Incised Bell-flower is so called from the fact that
the base of each of the lobes of the corolla is cut
away in a beautiful curve, and its flowers are thus
easily distinguished from those of the other Alpine
species. The precise object or advantage of this
peculiarity does not appear to be known at present.

We will notice one further Alpine Campanula —
a very remarkable one. It must not be imagined that
all Bell-flowers have blue flowers like the Harebell.
There is one Swiss species, the Tufted Campanula
(Campanula thyrsoidea, Linn.) (Plate XIX.), in which
the flowers are pale yellow, and the whole habit is
quite unlike that of the other Alpine Bell-flowers.
The plant has a rosette of hairy leaves close to the
ground, from which springs a stout stem 6 inches to
a foot in height, bearing numerous closely-set leaves
and ending in a dense spike of yellow flowers.

PLATE XIX.

The Tufted Campanula (Campanula thyrsoidea, Linn.).

[To face p. 90.

THE RAMPIONS 91

The whole plant is rough with hairs, cotton-like
hairs being even found on the pale yellow corolla.
Unlike the other Alpine Campanulas, this plant is a
biennial — that is to say, it takes two years to mature.
In the first year, when the seed germinates, only the
root and a rosette of leaves close to the ground are
formed. From this, in the second year, the stout,
leafy flower-stem shoots up, producing in July a dense
spike of flowers. Having set its seed, the plant dies.
It is not at all an uncommon plant, though much less
abundant than the three species of Bell-flower first
mentioned. It grows in the pastures, with often a
preference for somewhat shady places, near Rosa
alpina, Linn., or other shrubs.

THE EAMPIONS.

The blue flowers of the Eampions, genus Phyteuma
(natural order Campanulacese, the Bell-flower family),
are conspicuous in the Alpine meadows and pastur-
ages in the early summer. They are borne either in
long cylindrical spikes or short rounded heads. There
are several species, three of which are confined to
High Alpine habitats. Of the two British species,
the Eound-headed Rampion (Phyteuma orUculare,
Linn.) (Plate XX., Fig. 2) is very frequent in the
Alpine zone, but the other (P. spicatum. Linn.) does
not occur except at lower elevations.

The flowers of the Eampion are in some respects
very like, in others very unlike, those of their near
allies, the Bell-flowers.

92 TYPICAL FLOWERS OF ALPINE PASTURES

If we examine the young flower-buds, we shall
find that the petals are all united into a closed tube
(Text-fig. X., 1). At a later stage the petals begin to
split apart at the base and the style pushes through
the tip of the still united upper portions (Text-fig. X.,
2 and 3). Finally, the petals separate altogether and

-k.

FIG. X. — The Flowers of the Round-headed Rampion (Phyteuma
orbiculare, Linn.), in various stages. Magnified.

k — calyx; c— corolla; a = anther; s= style; 6 = stigma.

1. The united petals of a young flower, with the calyx below.

2. The petals separating in an older flower.

3. The petals further separated, and the style growing through the tip of the

tube formed by the petals.

4. Mature flower with free petals, showing the style with stylar brush, and

the three expanded stigmas.

curl apart, leaving the style fully exposed (Text-fig.
X., 4). The same mechanism of the stylar brush
exists here as in Campanula (p. 85). The Eampions,
however, differ from the Bell-flowers in the mature
petals being quite free from one another, though
they are slightly united when young.

PLATE XX.

FIG. 1.— Leaves of the White Dryas (Dry as octopetala, Linn.).

FIG. 2. — The Round-headed Rampion (Phyteuma orbiculare, Linn.).

[To face p. 92.

CHAPTER IV

ROCK PLANTS OF THE PASTURES

IN the preceding pages we have noticed some of the
most characteristic of Alpine genera represented in
the upland pastures, though some of their species
occur in the meadows, forests, or in other habitats.
The flowers of the pastures are the crowning glory
of the Alps, and we shall therefore devote the present
and the following chapters, to the consideration of
other members of this most interesting assemblage
of plants.

The pastures naturally vary in their physical
features. Some are dry and stony, while others more
closely resemble fertile meadows. Even in a typical
fertile pasture, rocky boulders, or rock masses, each
with its own little flora, are frequently conspicuous.
The conditions under which plant life flourishes on
the rocks and on the dry stony slopes, with their
poor soils and small water supply, are naturally in
marked contrast to those which pertain in the normal
or typical pasture with its rich soil, often well watered
by some neighbouring stream.

94 ROCK PLANTS OF THE PASTURES

As a rule, the plants which grow on the rocky
portions of an upland pasture are not those which
abound in the normal pasture. We will devote
the present chapter to the rock plants of the pas-
tures, including with them those which grow under
similar physical conditions on the dry^ bare, stony
slopes.

The study of the colonisation of bare ground or
virgin soil by plant life, whether at home or in the
Alps, is a most interesting occupation. Many will
have noticed how in England some artificially
made new ground suitable for plant life, such as a
railway embankment, becomes gradually populated,
the Coltsfoot being, as a rule, the first to seize upon
the opportunity to establish itself. In the Alps,
fresh areas of rock are constantly being exposed,
either by the washing away of the soil and its vegeta-
tion by streams, especially in time of flood, or by
avalanches in winter, or again by soil-slip, the ever-
present tendency of the soil of the sides of the valley
to slide downwards. In other cases, landslips on a
large scale not infrequently lay bare the rock of a
mountain side, which formerly was densely clothed
with vegetation. How do plants establish them-
selves on such new ground ? What are the first
species to take advantage of the fresh opportunity ?

We have already called attention to the struggle
for existence, the competition for room, light, and air,
among Alpine plants. It follows that any opportuni-
ties will naturally be quickly seized upon, where fresh

ROCK COLONISATION 95

space is found to be available. But all plants are not
capable of taking advantage of the fact that a new
area of bare rock has recently become exposed in
their neighbourhood. We know that if we transfer
a plant of a damp meadow species to, or sow its seeds
on, some bare and dry rocky ledge, the chances are
very greatly against the survival of the species in its
strange habitat. The plants which are most likely to
survive on the new ground are those which grew
formerly under conditions as nearly as possible similar
to those which prevail in the area in which colonisa-
tion is being begun afresh. These are the rock plants
of the pastures. They are the advance-guard of
vegetation in its march from the normal pasture to
the bare untenanted rocks, exposed from time to
time by geological agencies.

The colonisation of new ground is effected in the
great majority of cases by seeds which, in a very
large proportion of Alpine plants, are distributed by
the agency of the wind. Where outposts on some
bare ledge have become established, the advance of
vegetation may be furthered by some asexual method
such as the formation of runners and offsets, which
tend to distribute the species still further. But
initially it is the wind-blown seed which is the
coloniser in nine cases out of ten.

It is doubtful if a seed falling on absolutely bare
rock will survive in any instance. Certainly it often
happens that seeds perish in this way, for without
some kind of soil, however primitive, the chances of

96 ROCK PLANTS OF THE PASTURES

their survival are very small indeed. It need hardly
be pointed out that in any case there is always a
"high mortality" among seeds. A much larger
number are always produced than can ever possibly
survive, and the whole rationale of seed production is
that, while many are certainly doomed to perish, there
is a distinct chance that a few, perhaps only one, may
survive, and so the species will continue in being.

We must, therefore, study the formation of a
primitive soil, if we wish to understand all the stages
in colonisation. Let us consider a large slab of rock
recently laid bare. The surface of the slab will
probably not be quite smooth. Smaller or larger
irregularities in the surface will exist, and further,
the rock, under the influence of what the geologist
terms "weathering," will soon begin to crack in
various directions, and at the same time the small
irregularities of the surface will be accentuated.

It is around these irregularities and in the cracks
that the primitive soil accumulates. It may be
formed initially by those lowly plants known as
Lichens (p. 291), especially the Crustaceous Lichens,
forming the yellowish- or greenish-white crust on the
surface of the rocks, so commonly seen in the Alps.
These Lichens are really composed of two plants — one
an Alga or pond-weed, and the other a Fungus, living
together. After a time the body of the Lichen dies,
but it continues to remain attached to the rock.
The dead Lichens tend to hold any rain-water which
falls on them for a time, and small wind-blown vege-

PRIMITIVE SOILS 97

table fragments and dust collect round them, and so
a primitive soil is built up. If the seed of a rock
plant should have the good fortune to come to rest
on such a spot, its chances are good. Some sort of
soil at least exists, and that not entirely free from
moisture.

Sometimes Mosses play the chief part in the for-
mation of primitive soils, but these plants are rarely
the first colonisers, though often the second, where
some primitive soil already exists.

A very common soil of this nature in the Alps
consists of little matted masses of pine-needles — that
is, of the leaves of Pines, Spruces, or Larches. These
are blown for considerable distances by the wind, and
may come to rest in the cracks on the bare face of a
rock, or cling to the little knobs or irregularities of the
surface (Plate XLIV., Fig. 1). They form small
masses interlocked together, which are wonderfully
permanent, though not fixed to the rock face in any
way. They retain a considerable portion of the rain-
water that falls on them and collect humus and dust.
Thus in the end a primitive soil results.

One of the greatest of the rock colonisers in the
Alps is the genus Sempervivum, the House-leek,
which plays a part somewhat similar to the Coltsfoot
with us in Britain. The House-leeks are extremely
interesting plants in many directions, as we hope
to show.

98 ROCK PLANTS OF THE PASTURES

THE HOUSE-LEEKS.

The House-leeks, genus Sempervivum (natural
order Crassulaceae, the Stonecrop family), are among
the most striking Alpine plants in dry rocky situations.
The leaves are thick, fleshy, or succulent, and arranged
in rosettes (Plate XXIV., Fig. 1) close to the ground.
The flowering stems also bear similar leaves, but these
are smaller and more scattered in their arrangement.
The leaves of the rosette are held erect — that is to say,
the apex of the leaf points directly upwards — and thus
the sunlight only falls obliquely on their surfaces.
This is another adaptation which tends to reduce the
loss of water given off by the leaves. The leaves have
also water-storage reservoirs in their tissues, and so
are well adapted to the dry barren soils on which they
live fully exposed to the summer sun.

In Britain, one species of House-leek (Sempervivum
lector 'urn, Linn.) is common on cottage roofs and on
old walls. This plant is also not infrequent in
Switzerland. In the Alps there are also several
species with large, handsome, rose-coloured or yellow
flowers, which do not occur in Britain.

The Spider's - web House - leek (Sempervivum
arachnoideum, Linn.) (Plate XVIII., Fig. 1) has
peculiar rosettes in which the tips and edges of the
leaves are all bound together by a white network of
long hairs, the whole meshwork resembling a spider's
web, hence the specific name (Plate XXL). Where
the plant grows in very shady places, the network

PLATE XXI.

The Rosettes of the Spider's Web House-leek (Sempervivum
arachnoideum, Linn.).

[To face p. 98.

THE HOUSE-LEEKS 99

may be less pronounced, and the rosette appear
to be green and not white. On the other hand,
where fully exposed to the sun, the spider's web is
usually very much in evidence.

The object of these matted hairs would appear to
be to bind together the leaves of the rosette as com-
pactly as possible, and thus ensure that each leaf is
held erect, and that the risk of excessive loss of water
by transpiration (p. 12) is reduced to a minimum.
The fact above mentioned, that, in shady places, where
there is less danger from evaporation, the web is much
less in evidence or is only feebly developed, supports
this view.

The hairs of the spider's web, as Fraulein Dintel
of Vienna has shown, are modified glandular or
secretory hairs. Glandular hairs are abundant on the
leaves and flowering shoots of many Alpine species of
this genus. Fraulein Dintel finds that it is the secre-
tion of such hairs which forms the means whereby
they are bound together into the web.

Another common House-leek to be found on
Alpine rocks is the Mountain House-leek (Semper-
vivum montanum, Linn.) (Plate XXIV., Fig. 1,
Plate XXII., and Plate XXIIL), which like the
preceding species has rose-coloured flowers but no
spider's web on the leaves, though they possess
ordinary, small glandular hairs. This plant is per-
haps the most abundant representative of the genus in
the Alps.

We may take these two plants — the Spider's -web

100 ROCK PLANTS OF THE PASTURES

and Mountain House-leeks — as typical of the genus in
the Alps, and study them in regard to their adapta-
tions to the particular conditions under which they
there live. When the seed germinates on the primi-
tive soil of some freshly exposed rock, a little rosette
of leaves is first formed. The next step is the
formation of a colony of such rosettes. This is done
by means of what are termed runners and offsets,
quite like those of our ordinary garden Strawberry
Plants. From the parent plant, in the axil of one of
the leaves of the rosette, a thin, prostrate stem is put
out, which grows for some little distance along the
surface of the rock. At or near its end, a second
rosette of leaves is formed, which in turn produces
other runners and offsets.

If we remove from the soil a colony of Semper-
vivum such as that of the Mountain House -leek
figured on Plate XXIV., Fig. 1, we shall have no
difficulty in making out the runners and their off-
sets. In most cases where these plants grow on
flat-topped rocks with plenty of room all round, the
runners are very short and new rosettes or buds are
formed close to the parent, and so a very compact
colony is produced. It is thus scarcely possible,
unless the plant is removed from the soil, to make
out the relationships of the colony. The runners
connecting the rosettes will be found to persist for
a long time, and tend to bind the individuals of the
colony together as a whole.

Compactness of growth, which is here well

PLATE XXII.

•2

.1
S

43
H

THE MIGRATION OF HOUSE-LEEKS 101

illustrated, is characteristic of many Alpines. The
cushion plants, p. 179, and the carpet plants, are equally
compact, though entirely different in habit. The
compactness of the colony, cushion or carpet, tends to
reduce to a minimum the risk of intrusion of other
plants into the colony.

In the photographs of Sempervivum arachnoideum
and S. montanum, on Plates XVIII., Fig. 1, and
XXIV., Fig. 1, it is obvious that room for the
extension of the colony can be found on its margins.
The colony can advance and overwhelm the present
occupants of the soil. Let us, however, examine
cases where space is restricted, and see how the
plant meets the difficulty.

The photographs on Plates XXII. and XXIII.
show two colonies of the Mountain Sempervivum
growing in the crevices of an old wall bounding a
meadow near Saas Fee. For a time the plants have
been quite at home, but now the necessity for further
space to accommodate the growing colony has become
pressing. We notice that the colony is no longer
compact. We can now see the runners, which are
very much longer than they are under normal circum-
stances. Each bears a few small leaves, and ends in
a rosette-bud or offset. The runner arises in the axil
of a leaf of the parent rosette.

We notice in the photograph on Plate XXII.
the crowded nature of the rosettes, and, further,
that the whole colony is tilted upwards on its side
to face the light. For this reason, some of the

102 ROCK PLANTS OF THE PASTURES

runners appear to be shooting straight up into the
air. As a matter of fact, this is merely due to
the circumstance that the rosettes are tilted through
a high angle and the runners are always produced
at right angles to the rosettes. Other runners are
growing over the sides of the lichen-covered rocks,
and on the right-hand side of the picture two runners
are seen going round the corner, as it were, to another
crevice to seek "fresh Woods, and Pastures new."

In the photograph on Plate XXIII., a colony is
seen boldly letting itself down over the face of the
rocks from ledge to ledge. The runners seen on the
left-hand side are obviously creeping or marching
down hill. On the right, the plant, by means of its
enormously elongated runners, has, as it were, made
a ladder of itself and is descending over the miniature
precipice. The relation of the runner to the rosette
can be clearly seen in this photograph.

In the two Alpine species described here, the
runners are wonderfully persistent. Kerner has,
however, described another species (S. globiferum,
Linn. = S. soboliferum, Sims), which does not occur
in Switzerland, in which the young rosettes soon
become detached from the thread-like runners, and
are blown by the wind from one rocky ledge to
another, and eventually find refuge in some crevice,
where a new colony is founded. This vegetative
means of distribution has not, however, been observed
in the case of either the Spider's-web or the Mountain
House-leek.

PLATE XXIII.

The Migration of the Mountain House-leek (Sempermvum montanum. Linn.).

[To face p. 102.

HOUSE-LEEKS AND STONECROPS 103

The name Sempervivum, meaning "ever-living," is
not inappropriate for the House-leeks. It is true that
the individual rosettes do not live beyond a few
years at most, but their place is constantly being
taken by new rosettes. The result is, the colony
presents much the same appearance from year to
year at each season. The old, dead rosettes persist
for a long time beneath the new rosettes, and
go to increase the humus and thus enrich the poor
soil on which these plants manage to flourish.
Even the withered flower- stalks of the previous
year often remain attached to the rosettes. Many
small wind-blown particles of vegetable matter
and dust also collect round the colony, and thus
the soil constantly receives fresh additions from
without.

Other rock plants, which play an important part
as colonisers of fresh ground, are the Stonecrops or
Sedums, belonging to the same natural order as the
House-leeks, and, like them, fleshy, succulent-leaved
plants adapted to dry habitats, and some of the
Saxifrages which we have already considered in
Chapter III. In the Edelweiss, discussed in Chapter
I., we have a typical rock-plant of a different habit,
but equally adapted to similar dry situations, though
not a frequent coloniser.

We may now turn to Alpine species which not
only occur on rocky ledges, but are also frequent on
dry stony and semi-bare patches in the pastures, on
the bare moraines of glaciers, or on the debris of

104 ROCK PLANTS OF THE PASTURES

torrents, habitats which all present similar difficulties
to the plant as regards existence.

Among the first of these to be noticed as
extremely abundant in such situations throughout the
Alps, are the Wild Thymes (Thymus serpyllum, Linn.,
and T. chamcedrys, Fries, natural order Labiateae,
the Mint family). These plants produce flowers of
two sizes, the larger being hermaphrodite, and
the smaller possessing only female organs. The
difference between the large- and the small-flowered
plants is noticeable even at some little distance.

There are two other British plants belonging to
the Pea family (natural order Leguminosae) which
are often very much at home in the drier, stony
portions of the pastures and on the moraines of
glaciers.

THE BIRD'S-FOOT TREFOIL.

The Bird's-foot Trefoil (Lotus corniculatns, Linn.)
is remarkable for its indifference to the nature of
the soil, the degree of moisture, and situation. It
will flourish under almost any conditions in the Alps,
though it is most conspicuous on dry ground, where
it meets with less competition from its fellow- Alpines.
Its wide distribution over Europe and Central Asia,
and even in Australia, is, no doubt, due to its adapta-
bility to varied physical conditions.

It is extraordinary what a large area a single
plant of this Trefoil manages to cover in the Alps. It
is a perennial plant, held fast between the stones by a

PLATE XXIV.

— Rosettes of the Mountain House-leek (Sempercicum montanum, Linn.).

FIG. 2.— Rosette of a Sempercivum attacked by a Parasitic Fungus.

[To face p. 104.

THE LADY'S-FINGERS 105

long, stout root-stock. Above ground the leaves
radiate out in all directions from a very short stem.
The flowers, clustered in little umbrella-like heads
(umbels), are mounted on long stalks which extend
even further than the leaves. The whole spreading
habit adds greatly to the conspicuousness of the plant,
and its advertisement to the insect world is thereby
increased. As we shall see, when we come to
describe the carpet plants of the Alps, such as Dryas
octopetala, this spreading habit is characteristic of
many Alpines, and the fact that many flowers are
borne on the same plant is explained by the neces-
sity for a large seed production, since the chances
of the survival of an individual seed are smaller than
in the plains, owing to the severer physical conditions
which it has to combat.

The Bird's-foot Trefoil, as we should perhaps
expect, when we consider its indifference to habitat, is
a plant with many varieties. The flowers may vary in
colour, even on the same plant ; while the corolla is
usually yellow, in some cases it may be reddish or
even wholly red.

THE LADY'S-FINGERS.

The other British member of the Leguminosse,
the Lady's-fingers, or Kidney Vetch (Anthyllis vulner-
aria, Linn.), is almost as abundant as the Bird's-foot
Trefoil, with which it is often associated. The flowers
are also borne in umbels, though much larger than in
the previous plant. They are easily recognised by

106 ROCK PLANTS OF THE PASTURES

the hairy, inflated calyx formed by the united sepals.
The leaves also end in large terminal leaflets, an inch
or more in length. It is a biennial plant, existing
for two years only.

In this species also, the flowers are very variable
in colour. Usually yellow, they may be almost white,
or again more or less red, or entirely so. They are
frequently cross-fertilised by butterflies in the Alps,
though in the plains, humble-bees are the chief and
most useful visitors. Like the Bird's-foot Trefoil, it
is one of the most important pioneers of vegetation
on the bare stony patches of the pastures, on the
moraines of glaciers, and the debris brought down by
streams.

THE WHITE DRYAS.

The White Dryas (Dryas octopetala, Linn., natural
order Rosacese, the Eose family) is one of the most
beautiful of Alpine plants, flourishing in similar
habitats to the preceding. Its large white or yellowish-
white flowers form a welcome landmark on many a
bare patch in the pastures.

The habit of the White Dryas (Plate XXV.) is
very characteristic of that of many Alpines. It is
what is called a carpet plant. Other examples of
carpet plants will be found in the Trailing Azalea,
the Alpine Juniper, and Globularia cordifolia.

A carpet plant is really a very dwarf, recumbent
shrub : one might almost say a miniature tree. The
plant is woody and not herbaceous. The stem is very

PLATE XXV.

THE WHITE DRYAS 107

short and buried in the soil. Just above the ground
a very large number of long prostrate branches spread
over a considerable area, packed closely together.
The branches bear numerous little tufts of leaves, and
thus a green carpet of close texture, often occupying
many square feet in extent, is woven over the soil.

The stems of many of these carpet plants reach a
great age. In the case of Dryas, as many as a
hundred years have been recorded on the evidence of
the rings of growth of the woody tissues of the stem,
as seen in transverse section. Thus these lowly
plants are as permanent as many of the trees of a
forest.

This type of habit has many advantages. It
ensures space for the production of a very large
number of flowers and consequently seeds. This we
have seen to be a prime necessity for many Alpine
plants. The close, compact nature of the carpet is
very successful in preventing the intrusion of other
plants on the same ground. Nothing can live beneath
it. In the Alps the struggle for room leads to a
daily war among plants. Not only does a carpet
plant hold its ground successfully, but by the increase
in the length of the branches it can increase its
holding and oust out other plants which happen to be
situated near the margin of the carpet. In winter-
time also, when all the world is wrapped in snow,
this particular habit is no doubt extremely well
adapted to withstand the weight of the overlying
snow, and thus to ensure the plant against injury.

108 ROCK PLANTS OF THE PASTURES

The leaves, which, as we have seen, are borne in
little tufts on the branches, are oblong in shape, deeply
toothed, and mounted on long stalks. The upper
surface is of a shining, deep green colour, and quite
free from hairs. Below, the leaves are covered with
a thick, felt-like coat of white, downy hairs, as is seen
in the photograph on Plate XX., Fig. 1.

The leaves are evergreen and may persist for
four or five years. In winter-time, when covered
with snow, they are rolled on themselves, with the
hairy, lower surface innermost. The young leaves
are also covered over and protected by the downy
lower surfaces of the older.

The hairs on the lower side of the leaf serve to
protect the pores or stomata (see p. 10), which are
confined to this surface. They both prevent too great
evaporation of moisture from the leaf itself in dry
weather, and, in wet, ensure that the lower surface,
pressed close to the ground, does not become
thoroughly soaked with rain-water, and thus that the
gaseous interchange between the leaf and the atmo-
sphere (p. 10) be not hindered.

The large and beautiful flowers, borne on stalks
2 to 3 inches in length, are remarkable for the fact that
the sepals and petals vary from eight to ten, though
the former is the more usual number. The fruits,
which have long, feathery awns, enclosing a single
seed at the base, resemble those of the Spring and
Alpine Anemones (pp. 36 and 39) and the Mountain
Avens (p. 128), the latter being a near relative of the

THE TRAILING AZALEA 109

Dryas. They are adapted to travel long distances on
a windy day.

There are only three living species of Dryas. Like
many other Alpine plants, however, they are widely
distributed, occurring not only in the mountains of
Europe and Asia, but in the Arctic regions and in
North America.

THE TRAILING AZALEA.

The Trailing Azalea (Loiseleuria procumbens,
Desvaux, also known as Azalea procumbens^ Linn.,
natural order Ericaceae, the Heath family) is another
typical carpet plant (p. 106) of the Alps. The
numerous reddish-brown branches are imperfectly
clothed with leaves. The leaves are small and oval
in shape, set nearly at right angles to the branch and
arranged usually in four rows. The flowers are
usually borne in little groups near the ends of the
branches, each arising in the axil of a leaf. They are
small and rose coloured.

In comparison with many other Azaleas and
Rhododendrons (two genera now usually regarded
as identical), especially those characteristic of the
Himalayas, which are now so common in cultivation,
our Alpine Trailing Azalea, with its lowly habit and
small leaves and flowers, presents a marked contrast.
By some botanists it is still regarded as a typical
Azalea, while others include it in a separate genus,
Loiseleuria, on the ground that the flowers are
perfectly regular in their construction. There is

110 ROCK PLANTS OF THE PASTURES

only one species, which has, however, a wide
distribution.

In Alpine Switzerland the Trailing Azalea is fairly
common in all sorts of habitats. It may occur in
woods which are not very dense (p. 248), on flat-
topped hills resembling moorlands, and elsewhere,
though it generally inhabits some slab of rock or
some patch of dry stony ground. It occurs in the
Highlands of Scotland, where it also forms a carpet
on the flat dry hilltops. When grown in the Low-
lands, it may abandon its prostrate, trailing habit and
become erect.

The small leaves of this plant are worth examining.
It will be found that they are rolled inwards at the
edges. The stomata or pores are situated on the
lower surface in two grooves near the edges, which
are filled with hairs and further protected by the
incurving of the leaf at the margins. Similar adapta-
tions to guard against excessive loss of moisture from
the leaf are found in the leaves of the Alpine Heath
(Erica carnea), the Ling (Calluna vulgar is), and the
Black Empetrum (Empetrum nigrmi).

THE ALPINE GLOBULARIAS.

In the Alpine Globularias, of which there are two
species — the Eound-leaved Globularia (G. cordifolia,
Linn.) and the Bare-stemmed Globularia (G. nudi-
caulis, Linn., natural order Selagineae, the Selago
family) — we have a family of plants which does not

PLATE XXVI.

THE GLOBULARIAS AND BUTTERCUPS 111

occur in Britain. In Alpine Switzerland they are an
invasion from the Mediterranean subtropical flora.

The Alpine Globularias are typical carpet plants, in
habit quite like those which we have just discussed.
They are common on flat-topped rocks and in stony,
dry places, and are easily recognised by their little
dense heads of blue flowers borne erect on long stalks,
which are leafless except for one or two very small
bracts. The arrangement of the flowers in compact
heads closely resembles that characteristic of the
order Composite, the Daisy family, though here
there is no involucre of bracts below the flower-
head or capitulum. The flowers are two-lipped, and
are for the most part fertilised by butterflies. In
both species the leaves may occur in little rosettes
on some portions of the trailing stems, while else-
where they are scattered in their arrangement. In
the Round-leaved Globularia (the leaves of which, by
the way, are more spoon- shaped than round, despite
the specific name) numerous runners are produced.
These are not found in the case of the Bare-stemmed
Globularia ; cf. also Geum reptans and Geum alpinum,
p. 127.

THE ALPINE BUTTERCUP.

The Buttercups, or Ranunculi (natural order
Ranunculacese), are particularly numerous -in the
Alps. Some have yellow flowers, others white.
They flourish under a great variety of circumstances,
and are remarkable for being very little modified

112 ROCK PLANTS OF THE PASTURES

externally in response to the physical conditions under
which they grow.

The Alpine Buttercup (Ranunculus alpestris, Linn.)
is often an abundant representative of the genus on
rocks or on rocky ground, especially on calcareous
soils at an elevation of about 6,000 feet. Its
chief peculiarity is the dwarf habit, which, however,
it shares with other Buttercups at high elevations.
The whole plant is only from 2 to 4 inches in height.
The leaves, which spring from a very short stem, are
stalked, heart-shaped, and lobed. The nerves are
very conspicuous on the upper surfaces, as in some
other Alpines ; cf. Salix reticulata, p. 188. The white
flowers are solitary, and each is borne erect on a
long stalk (Plate XXX., Fig. 1).

The whole plant is remarkable for being entirely
free from hairs, unlike the majority of Alpines.

THE ALPINE PINKS.

In the month of July in the Alps, some of the
spring flowers, especially on the drier stony slopes in
the mountain pastures, are replaced by newcomers,
among which are the Pinks. Three species are
fairly common in the Alpine zone, while others
occur at lower elevations, and one rare species is
confined to the High Alps.

The Large - flowered Pink (Dianthus superbus,
Linn., natural order Caryophyllacese, the Pink
family), flourishing in meadows and on the edges
of woods, is the handsomest of the Alpine species.

THE ALPINE PINKS 113

It is easily recognised by its pale pink corolla, deeply
cut into a fringe of delicate segments.

The flowers have a scent resembling oil of cloves.
The Wood Pink (Dianthus sylvestris, Wulf., also
known as D. inodorus, Steud.), on the other hand, is
scentless, and has solitary flowers. It is perhaps the
commonest species all over Switzerland, especially in
rocky places. In the Carthusian Pink (Dianthus
Cartktmanorum, Linn.), the dark red flowers are
borne in clusters, and the leaves are blue-green. In
our cultivated Sweet - William, which is a Pink, we
find similar flower-clusters. In addition to the erect
flowering shoots, we sometimes find at the base of
this plant small, more or less prostrate shoots, each
bearing only a single flower, which is female and does
not possess stamens.

There is nothing very striking, biologically, to
relate of the Alpine pinks. Their tufted habit and
erect, grass-like leaves, peculiarities also shared by
other Alpine plants, such as the Anthericums (natural
order Liliaceae), are adaptations fitting them to exist
in situations fully exposed to the sun, and on soils
which are comparatively dry.

THE ALPINE TOADFLAX.

The Alpine Toadflax (Linaria alpina, Mill,
natural order Scrophulariacese, the Foxglove family)
is a characteristic plant on the dry stony debris
bordering on Alpine streams and in other localities
physically similar. We are familiar in Britain with

114 ROCK PLANTS OF THE PASTURES

two Toadflaxes, the handsome Yellow Toadflax
(Linaria vulgaris, Mill) and the Ivy-leaved Toadflax
(L. cymbalaria, Mill), the latter common on old
walls. These do not occur within the limits of the
Alpine zone. They are there replaced by the Alpine
Toadflax, with its violet or deep ultramarine blue
flowers, touched with yellow at the throat of the
corolla. This plant is thus a further example of an
Alpine replacement (see p. 266), and adds another to
the large number of the blue-flowered plants of the
Alps (p. 43).

In the Alpine zone this species is usually a
perennial. Prof. Bonnier of Paris has experimented
with this and other Alpine plants, in respect to its
duration of life at different altitudes. The vast
majority of Alpines are perennials, but certain annuals,
such as the Snow Gentian (Gentiana nivalis, Linn.),
p. 49, and Saxifraga controversy Sternberg, p. 82, or
biennials such as Campanula thyrsoidea, Linn., p. 90,
also manage to flourish very well. Prof. Bonnier
finds that many Lowland plants, which are annuals or
biennials in the plains, may become perennials if
transplanted to the Alpine zone. The Alpine Toad-
flax, on the other hand, if removed to the Lowlands,
is found to become an annual or biennial. The
duration of the life of the individual plant is thus,
at least to some extent, adapted to the physical
conditions under which it lives. Owing to the
shortness of the summer in the high Alps, there is
not sufficient time to carry out the life's work in

THE ALPINE TOADFLAX 115

one season, as in the plains, and consequently some
different plan of operation has to be adopted, and
this finds its expression in the perennial habit.

The flowers of the Alpine Toadflax, which, like
many other members of the same order, only possess
four perfect stamens, the fifth being suppressed, are
remarkable for the spur formed by one of the petals,
a rare occurrence in this family. The petals are also
so shaped that the corolla is closed at the throat.
The same feature is also seen in the Snap-Dragons
(Antirrhinum). The pollen and the honey are thus
hidden. This is a special adaptation, to ensure that
only large and strong insects, such as certain bees, can
force open the throat of the corolla. Further, such
insects must possess a long tongue to reach the
nectar secreted in the spur, and incidentally cross-
fertilise the flower. This flower is thus specialised for
certain insects alone.

There are many other Alpine plants flourishing in
rocky situations, such as some of the Potentillas
(natural order Eosaceae) or the Eock Catchfly (Silene
rupestris, Linn., natural order Caryophyllacese) (Plate
XVI., Fig. 2), with its widely spreading branches,
but most of them are not known to present any very
striking points of biological interest.

CHAPTER V

INTERESTING PLANTS OF THE ALPINE PASTURES

IN Chapters II. and III. we have reviewed some of
the more characteristic plants of the Alpine pastures.
There remain others, which, unlike those considered
in the last chapter, are rare in dry rocky habitats, but
inhabit the typical, grassy alpen or pastures. We
will consider some of the more interesting of these
in the present chapter.

The distribution of Alpine species in the Swiss
Alps is very uneven. Some districts are rich in
species more or less confined to them, so far as
Switzerland itself is concerned. For instance, the
valleys of Canton Valais, especially the Zermatt and
Visp Thalen, or the high Alpine valleys of the Upper
Engadine and Davos, with their tributaries, are remark-
able in this respect, and form the finest collecting
grounds for Alpine plants in Switzerland. Other
districts, such as the Bernese Oberland, are relatively
poor. In a later chapter we will discuss the theories
put forward to account for this inequality of distribu-
tion. For the present, we will simply bear in mind
that it exists.

116

THE EFFECT OF SOILS 117

At first sight it might appear that the nature of
the soil may be a controlling element in determining
the distribution of a species. Some plants appear to
occur in Switzerland only where granite or schist
forms the rock from which the soil is derived. Others
seem to frequent only limestone soils. A third set of
plants appear to be quite indifferent as to soil. We can
thus distinguish three groups : the calcicoles, confined
to limestone soils ; the calcifuges, which occur only
where lime is absent from the soil ; and a third, those
which are indifferent as to soil.

It is frequently asserted that the white-flowered
Anemone alpina occurs on all sorts of soil, while the
still commoner variety with yellow flowers, often called
Anemone sulphur 'ea, is only to be found on non-
calcareous soils. Again, the rarer Hairy Alpenrose
(Rhododendron hirsutum) is believed to be calcicole,
whereas the commoner species, Rhododendron ferru-
gineum, is indifferent in its tastes. The Auricula
(Primula auricula) is also stated to be calcicole.
It is true that in many localities this is the case.
The Auricula, for instance, flourishes exceedingly
on the limestone rocks of the Engstlen Alp (Canton
Berne). But from a botanical standpoint, in order
to estimate the absolute effect of the constituents
of a soil, as factors controlling the distribution of
plants in the Alps, it is of importance to know
whether each species is always restricted to one
particular soil.

Professor Bonnier, whose work we have already

118 INTERESTING PLANTS OF ALPINE PASTURES

referred to more than once, has specially enquired into
this problem. For this purpose he studied the flora of
three mountain regions, widely separated : the French
Alps of Dauphine, the Austrian Alps, and the
Carpathians. The distribution of the same plant was
observed in each of these three districts, especially in
relation to its soil. He found that some species, such
as the Glacial Buttercup (Ranunculus glacialis),
p. 193, the Stemless Catchfly (Silene acaulis), and
the Mountain House-leek (Sempervivum montanum),
p. 99, were calcifuge in two regions, though not in
the third.

Other plants, such as the Alpine Anemone
(Anemone alpina), p. 37, the Alpine Buttercup
(Ranunculus alpestris), p. 112, and the "White Dryas
(Dryas octopetala], p. 106, may be indifferent as to
soil in one or even two districts, yet in a third they
are distinctly calcicole. The Edelweiss (Leonto-
podium alpinum), p. 15, proved to be calcifuge in
Dauphine, indifferent as to soil in Austria, and
calcicole in the Carpathians. Hardly any Alpine
is confined to limestone soils in all three regions,
and only three species are absolutely calcifuge.

It is thus obvious that while in one country such
as Switzerland a plant may be almost entirely
calcicole, it is quite likely that, in one of the other
mountain ranges of Southern Europe, it will be found
to be indifferent as to soil, and thus the influence of
the soil alone on distribution is, at the most, local
and not absolute.

CHALET PLANTS 119

At the same time, the nature of the soil has
naturally a very profound influence on vegetation
locally. A good example is seen in the case of
" chalet plants," so called because they flourish in
abundance close to the picturesque wooden cow-
chalets of the pastures, and are but rarely found
elsewhere. This is probably due to the fact that
round the chalets the soil is relatively rich in humus
in comparison with many other localities, the humus
being derived from the manure which accumulates
close to the chalets, and which is to some extent
spread around them when exposed to rain.

Many of these chalet plants are common weeds.
Our well-known Dandelion (Taraxacum offitinale?
Weber, natural order Composite, the Daisy family),
with its beautiful little parachute fruits, adapted
for travelling long distances in the air, is very
common in this position, though also widely distributed
in other localities. Our two British Stinging Nettles,
Urtica urens, Linn., the Small Nettle, and Urtica
dioica, Linn., the Common Nettle, are also often
abundant near chalets, though rare or absent else-
where. The latter species flourishes exceedingly at
6,000 feet or more in the Alps, producing very tall,
vigorous plants, 3 feet or more in height.

The Alpine Gagea (Gagea Liotardi, Schult.,
natural order Liliaceae, the Lily family), allied to
the Star-of-Bethlehem and our British Yellow Gagea,

1 So called because parts of the plant were formerly used as
medicine; cf. Officinal.

120 INTERESTING PLANTS OF ALPINE PASTURES

is another plant which is rare except in damp places
near cow-chalets, as on the Engstlen Alp (Canton
Berne).

The Alpine Dock (Rumex alpinus, Linn., natural
order Polygonaceae, the Dock family), with very
large leaves borne in massive clumps, is frequently
to be seen near chalets. It is largely cultivated by
careful manuring, and is made use of as a fodder. The
leaves are cut two or three times during the course of
the summer and boiled down, the concoction being
laid by for the use of the cattle in winter. The
German-speaking Swiss call this dock "Blacken,"
and the patches of dock "Blackengarten." The
same name is also sometimes found applied to certain
pastures, such as the Blacken Alp on the Surgnen
Pass, where this plant is abundant.

THE WHITE VERATRUM.

We will now discuss some further pasture plants
of interest, beginning with the White Veratrum
(Veratrum album, Linn., natural order Liliaceae, the
Lily family) (Frontispiece, and Plates XXVII. and
XXVIII.), which is a stout herb very common in
the pastures. The stem is tall and the leaves light
green in colour, large and broad. The flowers are
greenish -white, the perianth leaves being widely open,
and the anthers globular in form. The specific name,
" album," = white, if it refers to the flowers, is certainly
a misnomer, for their colour is, as a rule, much nearer
green than white.

PLATE XXVII.

THE WHITE VERATRUM 121

The leaves are interesting structures (Plate
XXVII., Fig. 1). The upper surface is concave
and the leaf is folded lengthways, so that a series
of ridges separated by grooves are formed. It is
believed that these ridges and grooves help to
direct rain, falling on the leaf, to the soil imme-
diately above the large underground stem. The
water collects in the concave leaves and trickles down
in the grooves to the base of each, and thus falls on
the soil immediately below the insertion of the spirally
arranged leaves on the stem. In this way a larger
quantity of water is probably brought within the
reach of the underground stem, than would be the
case if the rain-water dripped from the tips or edges
of the leaves. The fact is easily verified by the
experiment of pouring some water from a neighbour-
ing stream over the plant, and watching its course to
the soil.

It is interesting to compare the young shoot of
a Veratrum (Plate XXVIII., Fig. 1) as it appears
forcing its way up above ground when the snow
is melting, with the fully expanded, mature plant
and its spreading leaves (Frontispiece, and Plate
XXVII., Fig. 1). The young shoot somewhat
resembles a compact form of cabbage. The leaves
are all held erect and closely wrapped round one
another, their tips being directed upwards, the whole
forming a compact, conical bud of large size, with a
few scale leaves at the base. If we cut one of
these shoots in two lengthways, we shall find the

122 INTERESTING PLANTS OF ALPINE PASTURES

whole plant, as we recognise it later when fully
expanded, present in miniature or almost tabloid
form ! The internodes of the stem and its branches —
that is to say, the portions of the stems between the
nodes where the leaves are borne — are very short
and compressed. As the stem grows, the internodes
lengthen rapidly.

The underground stem of Veratrum is a stout
stock, which is worth digging up and examining. It
contains a highly poisonous substance, the alkaloid
known as veratrin. The underground stem is, or was,
called by herbalists, the "white hellebore root," a
misnomer, for the stock is botanically a stem or
rhizome, and not a root.

If we examine the thick roots borne by this
underground stem, we shall find they are wrinkled
transversely (Plate XXVIII., Fig. 2). We have
here a good case of what are called "contractile
roots." Similar contractile roots occur in the case
of many other Alpines, such as some of the large
Gentians with yellow or red flowers, and they are
also very common among bulbous and tuberous
plants. By means of these roots, the rhizome,
tuber, or bulb, as the case may be, is being pulled
downwards continuously, and lowered, so to speak,
into the soil, especially when the plant is young.
The "Californian Lily," for instance, produces one
great, thick, contractile root annually, which draws
down the tuber into the soil, from two -fifths to
three-fourths of an inch each year. In another case,

PLATE XXVIII.

5 H

£0

THE WHITE VERATRUM 123

the seedlings of the British Cuckoo Pint (Arum
maculatum, Linn.) have been observed to be dragged
downwards a distance of 2 inches between May and
October.

It is impossible to attempt to explain here the
mechanism whereby the contraction is effected. To
do so would involve a detailed account of the internal
structure and functions of the root. It may, however,
be said, that certain internal cells have the power
of contracting, and when this tissue shrinks it
shortens the external tissues, which no longer fit it,
but are drawn into wrinkles, like a glove too long for
the finger. It is naturally important that the under-
ground stem, as it increases in size, should be buried
deeper and deeper in the soil, to escape the frosts of
winter, and it is by the contraction of these roots
that this is effected.

The flowers of Veratrum are interesting from the
fact that while those of the main flowering shoot
usually contain both sexes (hermaphrodite), the flowers
of the lateral branches are generally male only. In
the plant figured on Plate XXVII., Fig. 2, it was
found that thirty-two hermaphrodite flowers occurred
on the main axis, against forty-nine male flowers on
the lateral branches. We have already noticed in
the case of the Alpine Anemone, p. 39, that male
flowers are often more abundant than hermaphrodite.
Occasionally in Veratrum, some of the upper flowers
are entirely female, without any trace of stamens.

The flowers of Veratrum are often wholly green,

124 INTERESTING PLANTS OF ALPINE PASTURES

yet they are conspicuous objects, quite apart from the
large, shining, green leaves. Conspicuousness to the
insect world is a matter of the highest importance to
the majority of Alpine plants, which depend on insect
visits to ensure cross-fertilisation (p. 271). In the
greater number of cases, as in Veratrum, it is the
corolla or perianth which forms the conspicuous
advertisement to the insect world on the part of the
plant. The corolla of a Pansy or Violet (p. 161) is a
good example. In some plants, however, the stamens
play this part. The Meadow Eues (genus Thalictrum,
natural order Kanunculaceae), of which there are
several species in the Alps, including the beautiful
Thalictrum aquilegifolium, with its lilac-coloured
stamens, and the Alpine Willows, p. 189 (genus Salix,
natural order Salicaceae), are examples of this class.
In other flowers, it is the calyx, and not the corolla,
which serves as the attractive organ; as, for instance,
in the Globe-flower (genus Trollius), p. 207, and the
Monkshoods (genus Aconitum), p. 130, both belonging
to the Eanunculacese. In other plants, it is the bracts
or leaves of the inflorescence, as in Bupleurum and
Astrantia (natural order Umbelliferse), the Edelweiss,
p. 19, and the Spurges (genus Euphorbia, natural order
Euphorbiaceae), which perform this function. These
facts illustrate the variety of means by which nature
attains a single end.

THE CARLINE THISTLE 125

THE CARLINE THISTLE.

The Carline Thistle (Car Una acaulis, Linn.,
natural order Composite, the Daisy family) is quite
unmistakable from its habit of flowering flat on the
surface of the ground, in many an Alpine pasture
and mountain-side (Plate XXIX., Fig. 2). A real
stem is present, despite the specific name : only, it
is very short, and buried as much as possible in the
soil. This type of habit, known to botanists as the
geophilous habit, is characteristic of Alpine plants
as a whole. Usually the underground development
of an Alpine plant greatly exceeds that above
ground. Not only is the root system in many Alpines
extremely well developed, but the underground stem,
comparatively safe from the attacks of the winter's
frost, is often a large structure modified to serve as
a storehouse for reserve food laid by to enable the
plant to begin work again in the spring at the earliest
possible moment.

From the short underground stem of the stemless
thistle a large number of spiny leaves radiate, all
of them being closely pressed to the soil. In their
centre a single, large, pale yellow thistle-head, 2 to 3
inches in diameter, occurs, which in its turn is closely
pressed to the leaves. The flower-heads are ripe in
August, and usually persist throughout the following
winter. They are very interesting, from the fact
that they act like weather-glasses. The flower-
leaves (bracts) of the head, arch upwards and form

126 INTERESTING PLANTS OF ALPINE PASTURES

a penthouse over the flowers on the approach of wet
weather, so as to protect the pollen (Plate XXIX.,
Fig. 2). On sunny days the bracts bend backwards
on to the leaves, and leave the flowers fully exposed
to the heavens.

When the flower-head is young and its diameter
relatively small, the length of the bracts is also
short. But as the flower grows and increases in
size, so the bracts also lengthen. Thus the roof or
penthouse grows in proportion with the dimensions
of the whole flower-head, and protection is at all
times efficient.

It has been found experimentally that it is not a
change in the intensity of the light or of temperature,
which stimulates this Thistle to erect its penthouse,
but a variation in the amount of humidity or moisture
in the air. For this reason these flower-heads are
often used as weather-glasses or hygrometers.

The heads of the Carline Thistle are wonderfully
conspicuous, especially when expanded, and this is
largely due to the silvery white, glistening, inner
surface of the strap-shaped bracts. The flowers of
the head are much visited by humble-bees, which carry
the pollen from one flower to the stigma of another.
The stylar-brush mechanism, already described on
p. 85, exists in this, as in all other members of the
Composite.

PLATE XXIX.

o c

THE CREEPING AND MOUNTAIN AVENS 127

THE CREEPING AND MOUNTAIN AVENS.

The genus Geum (natural order Rosaceae, the
Rose family) is represented by four species in the
Alps. Two of these are British plants chiefly confined
to the Alpine meadows (p. 157). The other two,
sometimes placed in a distinct genus, Sieversia, are
the Creeping Avens (Geum reptans, Linn.) and
the Mountain Avens (Geum montanum, Linn.), both
common plants on the pastures, with a preference
for dry soils.

The Avens are easily mistaken for Buttercups or
even Anemones. In some respects the family to
which they belong closely resembles the Buttercup
family. It is distinguished by the fact that the sepals
are united below into a cup, on which the petals and
stamens are borne. If a flower is cut in half length-
ways with a pocket knife, this can easily be seen. In
the Buttercup family all the organs of the flower are
quite free from one another, and attached separately
to the receptacle.

The Creeping Avens produce little runners like
those of the Strawberry or of the House-leeks (p.
100). These take root a few inches away from the
parent stem, and at this point a bud is formed,
which grows into a new plant. The Mountain Avens,
on the other hand, does not form runners.

The leaves are compound, and all spring from a
short stem just above the ground. Each has a
large terminal leaflet, and the other leaflets become

128 INTERESTING PLANTS OF ALPINE PASTURES

smaller and smaller towards the base. In the case
of the Creeping Avens the leaflets are pointed
and have sharp teeth, while in the Mountain Avens
both the leaflets and teeth are rounded.

The yellow flowers, which are large and handsome,
are borne singly on long stalks, pinkish-brown in
colour. The calyx is double, a peculiarity also shared
by several other genera of Eosacese, including the
Potentillas, a very abundant Alpine race, and the
Strawberry. Six outer members of the calyx alternate
with six longer inner sepals. The petals are usually
six in number, while the stamens are very numerous.
The pinkish-brown fruits have feathery plumes, like
those of the White Dryas already mentioned (Plate
XXX., Fig. 2).

The flowers of the Avens are often unisexual, and
the same holds good also in the case of the White
Dryas. Some will be found to be perfect — that is to
say, they possess both stamens and carpels — while
others contain stamens only. This tendency to derive
unisexual male flowers, from perfect or hermaphrodite
organs, is not infrequent among Alpine species of
Rosaceae and Ranunculacese. We have already noticed
another instance, in the case of the Alpine Anemone
(p. 39).

THE LONG-SPURRED VIOLET.

The Long-spurred Violet (Viola calcarata, Linn.,
natural order Violaceae, the Violet family) is frequent
in June and July on the slopes of the higher pastures

PLATE XXX.

FIG. 1. — The Alpine Buttercup (Ranunculus alpestris, Linn.).

FIG. 2.— Fruits of the Mountain Avens (Geum montanum, Linn.).

[To face p. 128.

THE LONG-SPURRED VIOLET 129

all over Switzerland, between 6,000 and 9,000 feet.
It is often so abundant that it forms a regular carpet
of flowers of a pale violet-blue colour on the mountain-
side. The leaves are toothed. The flowers are
exceptionally large for Violets, and possess a very
long, slender spur, in which the honey is stored.
Like many other Alpine plants, they are fertilised by
butterflies, being specially adapted to this class of
insects. The long spur is a device for placing the
nectar out of the reach of any other insects except
butterflies, which alone possess a sufficiently long and
slender tongue or proboscis to penetrate into the spur.
This spur is more than f of an inch in length, and is
very narrow, being only about ^ of an inch across.
Even if the proboscis of some other insect were
long enough, the extreme narrowness of the spur
would prevent it reaching the nectar.

The great naturalist, Hermann Mueller, to whom
we owe our present knowledge of the fertilisation
of Alpine flowers, observed that in the Alps, nine
different butterflies visited no less than 194 flowers of
this Violet in the space of 6f minutes.

The mechanism to ensure cross-fertilisation is
otherwise practically identical with that to be
described later in the case of the Field Pansy
(p. 160).

In the High Alpine region, another but similar
species, the Mont Cenis Violet (Viola cenisia, Linn.),
occurs infrequently. The leaves differ in being quite
uncut at their margins.

130 INTERESTING PLANTS OF ALPINE PASTURES

THE ALPINE MONKSHOODS.

The Alpine Monkshoods, genus Aconitum (natural
order Kanunculacese, the Buttercup family), are several
in number. We will notice two : a blue-flowered
species, Aconitum napellus, Linn., and the yellow
flowered, Aconitum lycoctonum, Linn. The flowers
are interesting from their peculiar construction.
There are five coloured sepals, one being much larger
than the rest, and fashioned in the form of a helmet.
The petals may be eight in number, but they are all
very small, except a pair which are covered in by the
helmet-like sepal. These are converted into long-
stalked honey-glands, of a peculiar form, with a large
nectary at the apex. The flowers are specialised for
cross-fertilisation through the agency of humble-bees,
other insects being unable to reach the nectaries
protected by the helmet-like sepal. Aconitum is not
known to occur in any region of the world from
which humble-bees are absent. Curiously enough,
these insects often rob the flowers of the honey by
biting through the helmet, instead of seeking the
nectar by the natural entrance to the flower. In
such cases, of course, the plant loses the chance of
being cross-pollinated. Sometimes every flower in
the inflorescence is thus mutilated.

The tuberous roots, and also the leaves of the
Monkshoods, contain a very poisonous alkaloid,
known as aconitine. One-fiftieth part of a grain is
said to be a fatal dose. The alkaloid obtained from

THE ALPINE BUGLE 131

A. tycoctonum is less powerful than that from A.
napellus.

THE ALPINE BUGLE.

The Alpine Bugle (Ajuga pyramidalis, Linn.,
natural order Labiatese, the Mint family), is a strik-
ing plant, on account of its pagoda-like habit. It is
often to be found growing in the shade of rocks.
The build of this species is wonderfully regular and
symmetrical, almost formal. Like all Labiates, the
stem is square and the leaves are placed in opposite
pairs. The pairs of leaves alternate very regularly,
so that there are four longitudinal rows of leaves
in all, one on each side of the square stem.

In the Alpine Bugle the leaves are larger and
longer than in our British species, and decrease in size
much more gradually toward the top of the plant,
thus constructing the characteristic pagoda-like habit.
They are often tinged with a bright-red colour.

The flowers are borne in complicated and com-
pressed inflorescences in the axils of the leaves. They
occur in abundance from the base of the plant to
the very apex. Unlike the Swiss Lowland species
and some of the British Bugles, the Alpine plant
does not form runners. This fact is the more curious,
for runners are characteristic of many Alpine plants
such as the House-leeks, the Creeping Avens, and
others.

132 INTERESTING PLANTS OF ALPINE PASTURES

THE ALPINE CLOVER.

In the Alpine pastures and meadows various
species of Clover are conspicuous, some of which
have beautiful yellowish - brown flower - heads. We
will notice here one species, the Alpine Clover
(Trifolium alpinum, Linn., natural order Leguminosae,
the Pea family), in which the rose-coloured flowers
are very large and handsome. As compared with
Clovers in general, the heads contain only compara-
tively few flowers, which are much larger than those
of the other Swiss species. Here we see another
good example of increase in the size of the individual
flowers so characteristic of many Alpines. A few
large flowers in a head compete successfully with
heads containing many smaller flowers. The flowers
have a very strong, sweet scent.

The fruits are also interesting, from the fact that
the calyx remains attached to the one- or two-seeded
pod after the corolla has fallen, and aids as a flying
apparatus for the distribution of the fruit by the
wind.

The Alpine Clover in Switzerland is a calcifuge —
that is, it avoids calcareous soils. It has a great
underground development of root and stem, as we
shall find if we attempt to unearth a plant. It is
a perennial, and may attain to a considerable age.

THE ALPINE ORCHIDS 133

THE ARNICA.

Among the many Alpine Composites (natural
order Composite), the flowers of the Arnica (Arnica
montana, Linn.), of large size, and deep orange-yellow
colour, are quite common and unmistakable. The
roots and leaves of this plant contain a bitter
resinous substance, arnica, together with a volatile
oil which, in the form of a tincture, is made use of
medicinally, chiefly in connection with sprains and
bruises.

The leaves are borne close to the ground in a
cross-shaped rosette, and from this springs a long
flower-stalk, bearing one to three flowers and one or
two pairs of opposite leaves. The flowers of the
disc are good objects for the study, by means of a
hand-lens, of the mechanism of the stylar brush
(p. 85). When the flowering stage is over, the long
ray florets all droop and hang vertically, and thus
the heads assume a very characteristic appearance.

THE ALPINE ORCHIDS.

The Orchids of the Swiss Alpine regions are very
numerous, and though the individual flowers are
generally rather small, they are arranged in an
extremely graceful manner, which entitles them to be
reckoned among the beauty plants of the Alps. The
most handsome of all the Swiss Orchids, with very
large flowers in comparison to the rest, the Lady's
Slipper (Cypripedium calceolus, Linn.), is not un-

134 INTERESTING PLANTS OF ALPINE PASTURES

common in shady places in the Subalpine zone, though
rarely found in the higher region.

The Orchids are abundant in the Alps in three
types of habitat : (1) very wet, marshy, or boggy
places, especially where some rivulet is running
sluggishly; (2) steep banks in the pastures, which,
on the other hand, appear at least superficially to be
rather dry habitats ; (3) in the forests. Many of the
Orchids flourishing in these situations are British
plants, such as the Spotted Orchis (Orchis maculata,
Linn.), the Frog Orchid (Habenaria viridis, R. Br.),
and the Fragrant Habenaria (H. conopsea, Benth.).
We will here notice two Swiss Orchids, the Black
Nigritella (Nigritella angustifolia, Rich. = N. nigra,
Eeichb.) (Plate XXXI., Fig. 1), and its near relative
the Lesser Butterfly Orchid (Habenaria bifolia, R.
Br.) (Plate XXXI., Fig. 2). The latter is a frequent
British plant, while the former does not occur
with us.

THE BLACK NIGRITELLA.

The Black Nigritella is one of the best known of
Swiss Alpines and is a universal favourite. The little
conical heads (really dense spikes) of blackish-red
flowers are quite unmistakable. They are often very
abundant in the pastures in all sorts of situation,
especially on steep banks.

It is worth while to dig up a plant of Nigritella
and to examine it closely. If we unearth it carefully,
we shall find, buried deep in the soil, a short stem,

PLATE XXXI.

THE BLACK NIGRITELLA 135

and then below, in addition to ordinary roots, a
pair of tuberous roots, white and fleshy, which
are of different ages (Plate XXXL, Fig. 1). Both
are storehouses for reserve food material. The
larger tuber contains the nourishment for the flower-
ing stem, and was in existence last year. The
smaller tuber dates from the present year, and in
it will be stored reserves during the summer to
further the growth of next year's shoot. The same
state of affairs will be found in the case of the Lesser
Butterfly Orchid. The leaves are rather thick and
narrow. They wrap round one another at the base,
and are enclosed in one or more scale leaves, of a
brownish colour and thin papery texture. At the
base the leaves are white and fleshy, no chlorophyll
(see p. 10) being developed below ground because
light is absent. The flower-stalk springs from among
the leaves which invest it at the base.

We thus see that the Black Nigritella, like most
Swiss Orchids, is a pronounced geophyte (see p. 125).
Its storehouses of reserve food lie buried deep in the
soil, away from the dangers of winter frosts.

The flowers, like those of many other Orchids, have
a strong scent, in this case resembling that of vanilla.
Vanilla itself is obtained from the fruits of a tropical
climbing Orchid, known botanically as Vanilla plani-
folia, Andr. The structure of the flowers of the
Orchids, as is well known, is specially adapted to
insect visitors, and to cross-fertilisation by their
agency. Those of the Black Nigritella are perhaps

136 INTERESTING PLANTS OF ALPINE PASTURES

rather small for examination, and we may therefore
postpone a description of them until we come to
discuss the similar structure of the Lesser Butterfly
Orchid. We may merely note that in the Black
Nigritella a very short spur is found.

THE LESSER BUTTERFLY ORCHID.

The Lesser Butterfly Orchid (Habenaria bifolia,
R Br.), sometimes called Platanthera Mfolia, Kich.
(Plate XXXI., Fig. 2), is easily recognised by the two
very large and broad leaves, and by the white flowers
with long spurs, twice the length of the ovary. The
general habit is otherwise not dissimilar to Nigritella,
except that the leaves, other than the two broad ones
arising from the base, are very few and reduced to
scales, and the spike is cylindrical in form and much
less dense.

In shape the flowers are supposed to possess some
resemblance to a butterfly, just as other Orchid flowers
are likened to bees, spiders, and flies. Curiously
enough, the Lesser Butterfly Orchid is visited by
some nocturnal insects, and especially by Hawk-
moths (Sphinx). The clove-like scent of the flowers
is strongest towards evening, and attracts night-flying
insects. Only those with a sufficiently long and
slender tongue or proboscis can reach the honey
stored in the long spur.

Let us now examine one of the flowers to see
how they are specialised for their insect visitors. The
floral envelope in the Orchids is not differentiated into

THE LESSER BUTTERFLY ORCHID

137

calyx and corolla, bub consists of a perianth of three
outer, and three inner, floral leaves. Of the former,
one is directed upwards, and two of the inner series
lie just inside it. The other perianth members are

Fio. XI.

1. The Flower of the Lesser Butterfly Orchid (Habenaria bifolia, R. Br.).

Enlarged.

6r, The bract, in the axil of which the flower arises ; or, the twisted ovary ;
sp, the long spur of the labellum ; I, the labellum ; e, entrance to the
spur ; stt the stigma ; a, the anther.

2. A Pollinium. Much enlarged.

p, The pollinium proper ; *, the stalk ; d, the disc.

spreading. The lip (labellum) — that is, the third
member of the inner series — is directed forwards, and
forms a platform upon which the insect alights. In
many Orchids this lip is very different in form and
colour from the other perianth segments, and becomes
the most conspicuous portion of the whole flower.

138 INTERESTING PLANTS OF ALPINE PASTURES

In the present instance, however, the structure of the
flower is relatively simple. The labellum only differs
in being rather longer than the other segments, and
is often green at the tip. The base is produced
backwards in the form of a long spur, which contains
honey.

There is only one stamen in all Swiss Orchids
except the Lady's Slipper. This plant possesses a
flower with two stamens, somewhat different in
construction from those of the other Swiss genera.

The structure and relationships of the male and
female organs in the Orchids are highly peculiar.
The single stamen is united with the style of the
ovary to form a short column, of which the anther
forms the apex, and is placed immediately above the
stigma and over the entrance to the spur. Further,
the anther produces, not the usual dust of pollen
grains, but two club-shaped bodies mounted on short
stalks, and each attached below by a viscid adhesive
disc. The upper portion of the club, orpollinium as it
is called, consists of small compact masses of pollen
grains, united together by elastic threads. When ripe,
these club-shaped pollinia can easily be detached by
loosening the adhesive discs with a needle or a pin, or
even a sharp pencil point, and studied under a lens.
If they are carefully watched, it will be found that
they move. At first they are erect, but shortly they
bend on themselves by the stalk, in a forward
direction.

These features are all adaptations to cross-fertili-

THE LESSER BUTTERFLY ORCHID 139

sation by means of insects. An insect seeking the
entrance to the spur is quite likely to detach one or
both of the club-shaped pollinia, which adhere to its
head by the viscid discs. Before the insect seeks
another flower, these structures, at first erect, bend
forwards, and so when the head of the insect is

FIG. XII.— The Pollinium of an Orchid as removed from the flower, adhering
by the disc to the end of a needle. Much magnified.

1. The position immediately after removal from the flower.

2. The position assumed shortly afterwards.

inserted into the spur of the next flower, the pollinia
come into contact with the stigma, to which they
adhere. Thus cross-fertilisation is effected.

In the Lesser Butterfly Orchid the structure of the
anther and stylar column is relatively simple. The
club-shaped pollinia are naked, and not covered in as
in many other Orchids. They are therefore easily
seen.

Another feature common to most Orchids is that
the ovary is twisted, and thus the whole flower is
turned through a semicircle (180°), so that the
labellum, which is really the upper lip, comes to be

140 INTERESTING PLANTS OF ALPINE PASTURES

the lower. In the Black Nigritella, however, the
ovary is exceptional in that it is not twisted, so
that the labellum is in its proper position above the
entrance to the flower.

THE MEDITERRANEAN HEATH AND THE LING.

In the woods and thickets, and on moor-like
expanses in the Alps, we find little tufts of the
Mediterranean Heath or of the Ling growing in much
the same situation as with us in Britain. The Ling
is our common British (Calluna vulgaris, Salisb.,
natural order Ericaceae, the Heath family), the flowers
of which are interesting because the conspicuous
portion is the pink calyx, as long as, or longer than,
the corolla, which it almost entirely conceals. On
the other hand, only one of our five British heaths
is found either in Lowland or Alpine Switzerland.
This, the Mediterranean Heath (Erica carnea, Linn.),
is, in fact, the only Swiss species of the genus. It
does not occur in Great Britain, but only in Ireland.
It is really an immigrant from the Mediterranean
flora, and as such its presence in Alpine Switzerland
can be readily understood. How it got into southern
Ireland and yet not into England, is a more puzzling
problem.

In this species the leaves are arranged in whorls
of four, and the vase-shaped corolla is bright pink or
crimson in colour. In the Alps the flowers are
believed to be chiefly fertilised by butterflies, though
some observers state that bees perform this office.

HEATH AND LING 141

Both the Ling and the Mediterranean Heath
usually grow in places more or less fully exposed to
the sun. Their leaves, instead of being flat, are rolled
at the edges, so that the lower surface lines a
groove. By partially closing one's hand, and imagin-
ing that the palm and the lower surface of the
fingers correspond to the lower surface of the leaf,
one can roughly imitate the groove formed by the
inrolling of the edges of the leaf. The groove is
filled with minute hairs, springing from the under-
side of the leaf, which interlock together and choke
the groove. The stomata or pores, through which
water- vapour passes, are found only on the underside
of the leaf, and the object of the adaptation of the
hair-filled groove is to prevent an excessive loss of
water by evaporation from the leaf through the
pores — a matter of importance to a plant growing
fully exposed to the sun, and with often a limited
water-supply available in the soil for its roots.

CHAPTER VI

PLANTS OF THE ALPINE MEADOWS

THE meadows, which clothe the floors of many of the
Alpine valleys over 5,000 feet in altitude, and extend
for some little distance up their sides, are rich in
Alpine flowers. The wonderful massing and play of
colour in the meadows in spring, before they are
reaped for the first time, constitutes one of the chief
glories of the Alps. It is a sight which is missed
by the great majority of those who visit Alpine
Switzerland. By the beginning of July the meadows
have, as a rule, been cut, and a second crop is
growing vigorously, which will again be reaped in
August. In many Alpine valleys, yet a third crop
of hay is gathered in at the end of September or
in early October. But after the first yield, the
meadows never exhibit the same wealth of blossom
as in spring. To see them in their full glory, a pil-
grimage in the latter part of June is necessary.

The Alpine meadows, unlike the alpen or pastures,
which we have seen to be owned by the village com-
munes, are, as a rule, the private property of the

142

CULTIVATION OF THE MEADOWS 143

peasants. They are of immense importance, for the
successive crops of hay which they yield during the
summer furnish the greater portion of the fodder for
the cattle during the long winter months, and it is on
the cattle, or rather the cows, that the Swiss peasant
relies for subsistence almost exclusively. For this
reason the meadows are always very carefully tended.
A liberal coating of manure is applied to them usually
twice a year, and, where necessary, irrigation channels
are made to supply them with the maximum of
moisture. All these processes, including the cutting
with the scythe, have their bearing on the botany of
the meadow, as we shall see.

From a botanical point of view, these Alpine
meadows are interesting in many ways. But it must
always be borne in mind, especially when comparing
the flowers of the pastures with those of the meadows,
that the former are natural gardens, whereas the
meadows are highly artificial. The periodic cutting
with the scythe, and coating of manure regularly
applied, help to induce a strong, dense growth of
vegetation, consisting not so much of members of the
Grass family as of various other Alpine plants. Con-
sequently, some species which cannot grow without
plenty of space, light, and air, are crowded out, and
exist only in the pastures. The constant manuring
of the soil, which adds to the amount of humus or
decayed vegetable material naturally present in the
soil, while favourable to the growth of some species,
is unsuited to others, for plants differ much in

144 PLANTS OF THE ALPINE MEADOWS

their requirements in this respect. Then, again,
periodic mutilation by the scythe has a profound
influence on the habit of the plants, and this, again, is
a factor which eliminates certain plants from the
meadows. Again, as we shall see, the typical Alpine
meadows are damp associations, requiring a soil and
surroundings which are the reverse of dry. Hence
many plants, which have adapted themselves to the
drier pastures, are absent from the meadows, for
many typical Alpine meadows are little removed
from marshes, so far as the water contents of the soil
are concerned.

The exact time at which the meadows are first cut
varies from year to year. Sometimes when the winter's
snow has melted early, and the spring season has
been forward, the middle of June will see the
harvesters at work. In other years, a late spring
implies that the meadows will not be cut until the
first or even the second week in July. The nature of
the crop also varies somewhat from year to year. At
one time it will be comparatively short, and well under
3 feet in height, but of a thick, very compact growth.
In another year, the growth is thinner and the height
considerably greater.

To understand the conditions under which meadow
plants flourish, it may be well to consider the whole
year's cycle of a meadow.

In winter-time, for several months, on an average
from the middle of December to the middle of April,
the meadows are continuously covered with several

THE MEADOWS IN SPRING 145

feet of snow. Both before and after this period,
intermittent snowfalls occur, which by a lowering of
the temperature act as temporary checks to growth
during the most vigorous period of this phase of
plant life.

By the end of March or the beginning of April,
the snow begins to melt on the higher pastures, and
the first spring flowers appear. The meadows, how-
ever, are not uncovered for some weeks later. When
the snow at last disappears, scarcely a green blade is
to be seen. This is a peculiarly disagreeable season,
when but few have been tempted to study Alpine
vegetation, and the meadows, like the pastures, have
an ugly, deep yellowish-brown hue, rather reminding
one of a patch of common or grass land which has
recently been on fire.

The contrast between such a meadow in the
middle of April, and the same meadow in the middle
of June, with its wealth of colour and its harvest 3
feet high, is one of the most remarkable to be found in
nature. Few comparisons, to my mind, give one a
better idea of the huge outlay of energy which is being
expended on growth, or of the rapidity of that process
in the Alps.

It cannot be too clearly borne in mind that when
the covering of winter snow disappears, the flowering
period begins, and a great race against time is in
progress. Rather, in the case of many of the plants
concerned, the "full speed ahead" signal, as we shall
see, has been given before the snow has entirely melted.

146 PLANTS OF THE ALPINE MEADOWS

Everywhere there is a rush to get into flower. For
this object everything has been long prepared. The
plant seems to be aware that the season is short and
will last but a few months at the best, and that during
this period temporary checks to growth will frequently
occur in the shape of sudden lowerings of the tempera-
ture and transient snowfalls.

During the short summer period, the plant has to
perform its duty to the next generation — the reproduc-
tion of offspring. The actual flowering period is but
one stage in this process. Time is required to set
and ripen the seed, to distribute it, and to allow it a
fair opportunity of taking a firm hold in its new
surroundings before the mantle of snow cuts it off
from the outside world. All these processes are often
comparatively lengthy. Hence the race against time.

But other work equally important has to be
performed during this brief season. Most Alpine
plants are perennials, and during the short summer
months such a plant has varied duties to itself to
perform. When it reaches the light, it has not only
to manufacture its food -supply by the agency of
its leaves, and thus maintain its own existence and
make good the costly outlay of energy on repro-
duction, but it has, as often as not, to store up during
the summer those reserves which are to carry it
through the long, dark winter months. Thus, the
brief summer season is, indeed, a busy time. In a
temperate climate such as that of Britain, the period
available for these processes is quite long in com-

THE MEADOWS IN SPRING 147

parison with that of an Alpine or Arctic region : hence
the necessity for haste in the Alps. It is due to this
fact that the majority of Alpine plants "rush into
flower " at the earliest moment.

But there is one class of plants for which the race
is even more severe. In the High Alpine region the
snow does not disappear until much later — perhaps
the end of July. The conditions of May and June in
the valleys resemble those of July and August in the
High Alpine regions above 7,000 feet, and, by the
end of August, snow may ring down the curtain
once again. Thus for such plants the period of
energy is much shorter, and the struggle against
time still keener, while the work to be accomplished
remains much the same.

But to return to the Alpine meadows, just laid
bare by the melting snow, and of an ugly brownish
hue : the peasant now seizes the earliest opportunity
for a rich dressing of manure, often applied in semi-
liquid form from a primitive tank-like wooden box on
wheels. Within a few weeks the effect is magical.
A dense growth springs into existence, and increases
rapidly until, often before the end of June, the scythe
is at work.

The hay is made in the usual way as with us in
England, except that in many districts it is heaped up
to dry in the form of miniature stacks on cross-shaped
wooden frames placed, at this season, in the meadows.
At other times these frames may be often seen ranged
in rows outside, or on the walls of the chalets. When

148 PLANTS OF THE ALPINE MEADOWS

dry, the hay is carried up the slopes in truly
enormous bundles by a staggering Switzer, or by one
of his female relations, bo the hay chalets, which are
frequently perched up in the air on the points of
pillars of stone. Similar foundations are often used
for our hayricks in England. These pillars raise the
hay chalets above the level of the winter's snows, and
so secure the crop against damp.

Now we reach the second stage. The moment
the plant is mutilated by the scythe, it starts growth
again, helped by the influence of a fresh dressing of
manure. The new growth, however, is not so
vigorous as the first. The earlier crop represents a
growth prepared for during the late autumn, and to
some extent during the winter, but especially in the
months of early spring. Now, however, both the
period is shorter, and damage by the scythe has to be
made good. The strong young winter buds have
gone, and the new ones, less matured, are further
weakened by the fact that the leaves of the plant
which supply the energy for growth have also fallen
before the reaper.

The scythe, however, does not cut down the plant
quite close to the ground. The stem which remains
has first to heal its wounds, which it does by the
growth of a pad of corky tissues over the injured
part. Next, new leaves are developed afresh, and
lateral buds, already existing in a dormant state in the
axil of leaves right at the base of the stem, which
have escaped the scythe, now grow out into branches.

PECULIARITIES OF THE MEADOWS 149

The chief result of mutilation by the scythe is the
development of a branched system from the stem,
which originally may have been quite unbranched.
The whole character of the plant is more straggling
and less compact, owing to the prevalence of the
branched habit, and the flowers are smaller and less
conspicuous. Thus the second crop of hay differs
remarkably from the first, and the Alpine meadows,
when ready for the scythe for the second time, never
present the same appearance as they did in spring.
When a third crop is harvested, the contrast is even
greater. Such is briefly a year's history of an Alpine
meadow.

The meadow plants form an interesting association
(see p. 32), well worthy of study, and of comparison
with the flora of the neighbouring alpen or pastures.
An English meadow in June is fair to look upon,
but an Alpine meadow is still finer. This fact is due
to several causes. Although our British fields are
rich with Buttercups and Daisies, as well as other
flowers, yet the Grasses, with their comparatively
inconspicuous flowers, form by far the greater pro-
portion of the plant inhabitants of the meadow. In
the Alps, on the other hand, Flowering Plants of the
Dicotyledonous class with conspicuous flowers are
quite as numerous individually as the Grasses, and
sometimes even outnumber them.

The greater beauty of an Alpine meadow is also
due to the more intense coloration of the flowers,
and to the extremely robust or " well-grown " habit

150 PLANTS OF THE ALPINE MEADOWS

of the plants themselves. These plants frequently
grow in clumps, which produce a wealth of bloom,
forming conspicuous masses of colour when the whole
meadow is viewed in the aggregate. We frequently
notice large colonies of Lychnis, or Silene, of Bell-
flowers and Rampions, of Polygonum, of Geranium, or
Polemonium, each adding its note of colour, and con-
tributing to the harmony of the meadow.

Another peculiarity of the Alpine meadows is
that, in the majority of cases, they are comparatively
damp or even wet places. Many of the plants of the
Alps, with the most conspicuous flowers, thrive best
in damp soils. The valley meadows of the Engadine
or the Zermatt region are, for the most part, typical
damp meadows. Those near Saas Grund, on the
other hand, on drier soils are carefully irrigated, in
order that they may receive the maximum amount of
moisture available. On the Rieder Alp, above Morel,
in the Rhone valley, we find typical water meadows
in which a large number of marsh plants, such as
Trollim europceus, the Globe Flower, flourish.

One reason why the soil of an Alpine meadow is
much damper than that of a typical English meadow,
is the fact that it is well watered by the swiftly flowing
streams and their tributaries, which are invariably
to be seen traversing the fertile valleys in the Alps.
Another and equally important factor is the nature
of the soil, which frequently consists entirely of
peat, one of the most powerful water - retainers
among soils. The peat itself is formed by the slow

MEADOW FLOWERS 151

but constant accumulation of vegetable debris through
a period extending over many centuries.

In the photograph on Plate XXXII., Fig. 2, a
section or cutting of a typical valley-meadow is seen.
The soil is composed entirely of peat, which is here
more than 4 feet thick. In the lower portion, the
white bleached masses, which are seen projecting,
are the roots of trees, probably of Pines, which once
covered the area now forming the meadow, like those
seen at some little distance to the right of the
photograph. It will be noticed that the actual soil
on which the meadow plants are seen growing is very
thin. It should be also pointed out that the hay of
this particular meadow had been cut some weeks
before the photograph was taken, so in this case the
crop is small.

If we were to make an analysis of an Alpine meadow,
we should perhaps be surprised to find how many of
its inhabitants are British plants. Several of our
British Buttercups, such as the Bulbous Buttercup
(Ranunculus bulbosus, Linn.), the Field Buttercup (R.
acris, Linn.), are abundant. The Bistort (Polygonum
Ustorta, Linn.), the Ox-eye Daisy (Chrysanthemum
leucanthemum, Linn.), are characteristic. The Hare-
bell (Campanula rotundifolia, Linn.), and the common
Field Pansy (Viola tricolor, Linn.), are frequent. The
Water Avens (Geum rivale, Linn.), and Jacob's Ladder
(Polemonium cceruleum, Linn.), are often common.
Some of our British Geraniums, Campions, especially
Silene cucubalus, Wibel, and Catchflies, such as Lychnis

152 PLANTS OF THE ALPINE MEADOWS

dioica, Linn., add greatly to the colour-mass of many
an Alpine meadow. In late autumn the Meadow
Saffron (Colchicum autwnnale, Linn.) reigns supreme.
On the other hand, many of the conspicuous
elements of the meadow flora are not found
in Britain. Of the characteristic Swiss plants
discussed in Chapters II. and III., only the
Campanulas and Rampions are frequent in the
meadows. Scheuchzer's Campanula (Campanula
Scheuchzeri, Vill.), and C. rhomboidalis, Linn., with
various species of Eampion, increase the percentage of
blue-flowered meadow plants. To them are added
many other non-British plants, too numerous to
mention here, for it is not our purpose to construct a
full list of the meadow plants of Alpine Switzerland.
The above-mentioned species may serve to indicate
some of the commoner inhabitants to be met with in
most Alpine meadows.

At the same time, if we compare the floras of two
Alpine meadows some little distance apart, we shall
often find that they are dissimilar, or rather that
the plants which are specially abundant in one
meadow are less frequent in another. The meadows
thus vary noticeably among themselves as regards
their most abundant constituents, this, in fact, being
one of their chief characteristics.

We will now study some of the commoner
meadow plants, beginning with the Spring Crocus.

THE SPRING CROCUS

153

THE SPRING CROCUS.

The Spring Crocus (Crocus vernus, All., natural
order Iridacese, the Iris family) is nearly always the
first flower to appear in the meadows on the melting
of the winter's snow (Plate XXXI., Fig. 1).

It is what botanists call a pronounced geophyte —

Fio. XIII.— The Underground Stem or Conn of the Spring Crocus
(Crocus vernus, All.), in Spring.

The scale-leaves have been removed.

a, Corm developed from the base of last year's flowering shoot ; 6, corm
of previous year ; c, scars of scale-leaves ; d, remains of last year's
flowering shoot ; *, buds which will flower this year ; /, smaller buds
which may flower next year.

that is to say, the plant spends a large proportion
of its life underground. The stem has a peculiar
structure, which is often spoken of by the horti-
cultural fraternity as a bulb, but which is more
accurately described as a corm (Fig. XIII.). It is a

154 PLANTS OF THE ALPINE MEADOWS

thick, solid body which serves mainly as a store-
house for reserve food materials. Externally it is
sheathed in a few light brown scale-leaves of thin
papery texture. Numerous roots spring from the
base of the corm. From this underground stem, buds
grow out, one or more of which are flower-buds.
The central portion of each bud, the flower, is
pushed out first, and appears above the soil some
considerable time before the leaves, which in the
young bud surround it, reach the light After the
flower has died down, the base of the bud which bore
it swells out into a new corm for next year.

We shall understand how the Crocus manages to
come into flower at the earliest possible moment, if
the flower-bud, borne by the corm, is cut open length-
ways. It will then be found that the flower is
already developed in miniature. All the parts, such
as the six perianth leaves, the three stamens, and the
ovary, are easily recognised.

Thus we see that long before the snow of the
meadow begins to melt, the flower is ready. On
the coming of spring, all the plant has to do is to
lengthen the flower-axis below the flower, and thus
push up the flower to the light through the leaves,
which for the time being remain dormant.

The Crocus, like some other Alpine spring flowers,
frequently does not wait for the snow to entirely
disappear. We shall often notice, where some patch
of winter's snow still lingers in a meadow, the state
of affairs seen in Plate XXXIL, Fig. 1. Here the

PLATE XXXII.

FIG. 1.— The Spring Crocus (Crocus vernus, All.) Flowering in the Snow.

FIG. 2.— Section of an Alpine Meadow, showing the Peat Soil.

[To face p. 154.

THE SPRING CROCUS 155

flowers have forced their way up through the snow,
and flowering is in full swing. We have already
discussed (p. 62) another instance of this curious
haste to arrive at the flowering stage, when dealing
with the Soldanellas of the pastures.

Once the process of flowering is over and done
with, much work has still to be performed if the
results of fertilisation are to be carried through to a
successful ending. The next thing the plant does is
to send up the leaves. If we enquire why the leaves
appear after the flowers and not before, the answer is
quite simple. The work of flowering demands a
continuous supply of energy, just as an engine
requires energy obtained from coal to enable it to
perform work. In the case of the Crocus, sufficient
energy for the flowering stage is stored up in the
corm or underground stem, in the shape of reserve
food materials. Hence new food materials, furnished
by the green leaves, are not for the moment required.
But once the flowers are over and withered, fresh
food-supplies are required, both to ripen the seed and
to store up new reserves in the new corms of the
coming year. These supplies are manufactured by
the leaves when once they reach the light.

Another peculiarity of the Crocus, which is of
interest, is the varied coloration of the flowers.
Some are nearly white, others are almost wholly
purple, others, again, yellowish or pinkish. In
some flowers, again, all three colours are combined.
The significance of these fluctuations in colour is not

156 PLANTS OF THE ALPINE MEADOWS

yet fully understood. No doubt the plants are
thereby rendered extremely conspicuous to butterflies,
by which they are often fertilised, though it is
possible that self-fertilisation is the rule. In the case
of the Autumn Saffron, which we shall shortly discuss,
the colour has become " fixed " or constant, and this
would appear at first sight just as effective as an
insect advertisement. However, it does not appear
that the manner of fertilisation of the Saffron flowers
is known with certainty at present.

THE VIVIPAROUS POLYGONUM.

The Viviparous Polygonum (Polygonum mviparum,
Linn., natural order Polygonacese, the Dock family), is
a very frequent plant in the Alpine meadows.
Everyone is familiar with the beautiful pink spikes
of flowers of its near relative the Bistort, Polygonum
Ustorta, Linn., which is a much more conspicuous
plant, often contributing largely to the colour scheme
of the meadows. The Viviparous Polygonum (Plate
XXXV., Fig. 2) is easily recognised by the slender
flower-spike, partly composed of small white or flesh-
coloured flowers in the higher portion, and numerous
little red "bulbils" below.

These "bulbils" are not flowers, though they
occupy the position of flowers. They are really little
buds — minute leafy shoots— which become detached
from the spike and fall to the ground. They root
themselves to the soil, and grow into new plants.
Thus, should the Viviparous Bistort fail to produce

THE WATER AVENS 157

any seed owing to the flowers of the spike having
missed fertilisation, the perpetuity of the plant is at
any rate secured by means of these asexually produced
"bulbils."

The ancient term "viviparous," sometimes still
applied to this means of propagation, and still sur-
viving in the specific name, is extremely misleading.
When it was first used, the whole nature of these
bulbils was misunderstood. The bulbils are in no
way connected with any sexual organs, though in
position they replace them. They are purely vegeta-
tive, and the term viviparous should be confined to
certain animals, where it has a definite meaning and
significance.

Several other Alpine plants, such as the Grass,
Poa alpina, Linn., and a Saxifrage, Saxifraga cernua,
Linn., also produce bulbils. This means of repro-
duction is sometimes met with in Lowland plants,
though not so frequently as in Alpine and Arctic
species, which is probably due to the fact that it
is a saving of time for a plant to reproduce itself by
bulbils rather than by seeds. The Viviparous Poly-
gonum is itself an Alpine plant in Britain.

THE WATER AVENS.

The frequent occurrence of the Water Avens
(Geum rivale, Linn., natural order Rosaceae, the Rose
family), in the meadows of the Davos and other Alpine
valleys, affords a sure indication of the dampness of
the soil of the typical Alpine meadow. With us in

158

PLANTS OF THE ALPINE MEADOWS

Britain, this plant flourishes chiefly in marshes and
wet ditches.

The drooping flowers, with dusky red petals, are
interesting from several points of view. Like the
Potentillas, this plant possesses a double calyx. The
outer five sepals, known botanically as the epicalyx,

FIG. XIV.— Stages in the development of the Fruit of the Water Avens
(Qeum rivale, Linn.).

are small, and alternate in position with the five larger
inner sepals.

The fruit of the Water Avens (Text-fig. XIV.)
furnishes material for an interesting study. It is
especially adapted for distribution by animal agency.
Each of the carpels grows out into a long awn as
the fruits mature. A well-marked hook is developed

THE FIELD PANSY 159

at the end of the awn, and this catches in the fur or
coat of any animal passing, and thus the seed at the
base of the awn is carried to a distance from the
parent plant. It eventually becomes detached, and
starts life on its own account.

By comparing different stages in the formation of
the mature fruit, it will be found that the hook is not
formed at the actual tip of the awn. As the awn
matures, it develops a twist or kink at a little distance
from the end. At a later stage, the end of the awn
above the kink is thrown off altogether, and a sharp
point now terminates the fruit above the kink. The
apparatus is now mature. The pointed end, in con-
junction with the kink below it, acts quite like a fish
hook, and tends to stick into any rough substance
with which it is brought in contact. The stages in
the formation of the mature awn are shown in Text-
fig. XIV.

THE FIELD PANSY.

One of the first British plants which will be recog-
nised in an Alpine meadow is the Field Pansy or
Heart's-ease (Viola tricolor, Linn., natural order
Violacese, the Violet family). Though a humble plant,
it is interesting in several respects. The variation in
the colours of different flowers, and variety of colour
often observed in a single flower, ranging from purple
and yellow to white, is a feature in which this species
contrasts very markedly with the two other Violets
common in the Alpine zone, which we shall describe

160 PLANTS OF THE ALPINE MEADOWS

in other chapters. The flowers of the Long-spurred
Violet (Viola calcarata, Linn.) (p. 128), are a uniform
pale blue; those of the Two-flowered Violet (V. biftora,
Linn.) (p. 253) are yellow.

The flowers of all the Violets are specially modified
or adapted to their insect visitors, and we may

FIG. XV.— Section of a Flower of the Field Pansy (Viola tricolor, Linn.).

o, The flower-stalk; 6, sepal; c, petal; d, stamen; e, nectary;
/, spur of petal ; g, ovary ; h, stigma.

perhaps choose this opportunity of describing the
flower of this common species in detail, in order
to compare those of the Long-spurred and Two-
flowered Violets. A section through a flower is shown
in Text-fig. XV. All the organs there seen can easily
be observed by dissecting a flower with a needle under
a hand-lens.

THE FLOWER OF THE PANSY 161

Of the five petals, one is larger than the rest, and
serves as a landing-stage for the bees and butterflies
visiting the flower in search of the nectar, which
is hidden away in the spur or backward tubular
prolongation of the large petal. The bright colours
of the petals, as a whole, serve to attract these insects,
and the numerous streaks or lines of colour, all
leading to the centre of the flower, serve as "honey
guides," directing the insect to the narrow opening at
the mouth of the flower, through which the knob-
like stigma projects. If we dissect away the petals,
we shall expose the five anthers, which are mounted
on very short stalks, and closely applied to the
ovary. The anthers open inwards towards the pistil,
and above each anther a membranous appendage is
found. From the bases of two of the stamens, two
glistening, white, spur-like nectaries, secreting honey,
hang freely in the spur of the large petal. They will
be easily seen if the spur is carefully slit open. In
the centre of the flower, the ovary is produced
upwards into a club-shaped hairy stigma, on one side
of which is a small pit or groove, below which a little
triangular flap or valve can be seen. The stigmatic
surface is situated in this groove.

Now let us imagine that a bee or butterfly has
just alighted on the large petal. As it thrusts its
tongue or proboscis into the opening of the flower to
get at the honey in the spur, any pollen, derived from
a previously visited flower, adhering to its head comes
into contact with the stigmatic surface, which lies in

162 PLANTS OF THE ALPINE MEADOWS

the groove immediately above the entrance to the
flower. Thus the plant visited is cross-fertilised.
As the insect pushes its proboscis between the ring
of stamens and the style, its head becomes dusted
with pollen, exuded from the inner side of the anthers,
and this pollen is carried away to another plant. All
risk of transferring the pollen of one flower to the
stigma of the same flower, as the insect withdraws its
head, is prevented by the little triangular flap above
mentioned, which is automatically forced upwards by
the insect's head and thus shields the stigma.

In addition to the larger and more showy flowers
of the Field Pansy, adapted to cross-pollination, other
much smaller and less conspicuous flowers may be
found to occur, which are self-pollinated. In other
species of Violet, such as the Sweet Violet (Viola
odorata, Linn.), which is not Alpine, some of the
flowers are not only self-pollinated, but never open,
and after fertilisation, bury themselves (i.e., the whole
flower) in the soil.

The fruits of the Field Pansy are also very interest-
ing. The ovary bears many small seeds in a single
chamber. When ripe, the fruit becomes erect, though
formerly pendulent, and in dry weather the ovary
splits into three valves, each of which has usually
three rows of seeds. The valves dry up and contract,
and thus the seeds are pressed firmly against one
another, and are shot out one by one, often to a con-
siderable distance. A space of 3 feet has been
recorded. We can imitate this mechanism for seed

THE CAMPION

163

distribution by holding in the hand a group of several
balls. If pressure is put on the outer members of
the group, one of the inner balls is squeezed out.

Text-fig. XVI. illustrates the way in which the
seeds are ejected. The seeds
of each row are shot out in
regular order. The upper
valve has still its three rows
of seeds; the valve below
to the left has lost one row,
while that on the right has
lost two rows. When all the
seeds are distributed, the valves
of the capsule close on them-
selves.

THE CAMPION.

A very common plant in
the Alpine meadows is the FIG. xvi.— Ripe Fruit of the
Campion (Stone cucubalus,
Wibel = Silene inflata, Sm.,
natural order Caryophyllace*,
the Pink family). This plant
forms massive clumps, which
are quite unmistakable. The smooth, glossy, hairless
stems bear numerous white flowers, with a very
inflated or swollen calyx, the veins of which are very
prominent. Some of the flowers contain both male
and female organs, others only stamens, and others,
again, only an ovary, the last being distinctly smaller
flowers,

seeds- Enlarge

lost one row? the right valve,

164 PLANTS OF THE ALPINE MEADOWS

The inflated calyx certainly adds greatly to the
conspicuousness of the flowers. The object of this
adaptation is, however, still a debated question. The
theory is that the swollen calyx serves to prevent
the flower being "robbed" of its honey. The plant
is visited by humble-bees and butterflies. These
insects will often try to rob a flower of its honey
by biting through the base of the calyx and corolla,
instead of entering the flower in the legitimate
manner. It is obvious that in such cases the
insect can be of no service to the plant in the way
of cross-pollination. It is imagined that the inflated
calyx in some way protects the plant by making
robbery of the honey impossible, or at least more
difficult. If this is so, it can only be by deceiving
the insect, for the tongue or proboscis of a humble-
bee or butterfly is sufficiently long to penetrate both
calyx and corolla and the empty space between these
organs. It may be that the insect, having bitten
through the calyx, is deceived by imagining that the
space between the calyx and corolla, where there is
no honey, is really the space surrounding the ovary
where honey should be found. Finding no honey
there, it assumes that some other insect visitor has
been beforehand, and thus desists from further efforts.
In view of the considerable intelligence possessed
by such insects, it is doubtful, however, if so trans-
parent a device would prove successful; hence this
theory is not above suspicion.

THE MEADOW SAFFRONS 165

THE EED LYCHNIS.

The Red Lychnis (Lychnis dioica, Linn.), belong-
ing to the same family as the Campion, and like-
wise a common British plant, is another meadow
species of some interest. It is a very near relative
of the White Lychnis (L. vespertina, Siboth.), of
which it is very probably only a variety with red
flowers adapted to pollination in the day-time,
whereas in the White Lychnis the flowers are closed
by day and open only at night, when they are visited
by moths. White is a more conspicuous colour than
red at night-time, though not by daylight. The
Red Lychnis is fertilised by bees. Both these plants
have unisexual flowers, the male and female flowers
being borne on different plants. Bisexual flowers
are also sometimes found.

THE MEADOW SAFFRONS.

If we visit the Alps in September, we shall find
that the meadows in autumn present a very similar
appearance to that noticed in early spring. They
are thick with a Crocus-like plant, often called the
Meadow Crocus, with pink or lilac flowers. This is
really the Meadow Saffron, and in most cases it is the
common British Saffron (Colchicum autumnale, Linn.,
natural order Liliaceae, the Lily family). There is
another species sometimes found in cantons Tessin
and the Valais, the Alpine Saffron (CokMcum alpinum,

166 PLANTS OF THE ALPINE MEADOWS

D. C.), a rare plant, but very similar to the commoner
species.

The Meadow Saffron, though somewhat similar in
habit to the Spring Crocus, is no relative of that plant,
but belongs to a distinct family. It is easily dis-
tinguished by the six stamens, whereas the Crocus has
only three. But, like the Crocus, the Meadow Saffron
is a pronounced geophyte (p. 125), the whole plant
being buried deep in the soil, perhaps a foot or more
below the surface, at which depths frosts fail to pene-
trate. Only at certain seasons of the year does any
portion of the plant appear above ground.

If we take the trouble to unearth a plant by
digging out a large sod of turf a foot or more deep —
no easy task, and a delicate operation needing some
care, if it is to be performed without injury to the
plant — we shall find a little underground stem. The
structure of this stem or corm is different from that of
the Crocus, and is illustrated in Text-fig. XVII.

In autumn, when the flowers dot the meadows,
there are no leaves to be seen above ground. The
leaves, which are narrow, though broader than those
of a Crocus, do not appear until the following spring,
when, in the absence of flowers, they are easily over-
looked. All that we see above ground in the autumn,
are the upper parts of extraordinarily long flowers.
They rise perhaps 4 inches above the soil, and extend
below ground for another 9 to 12 inches, and are thus
of a total length of a foot or more.

Two flowers nearly always arise from each under-

THE MEADOW SAFFRON

167

ground stem. A few brownish scale-leaves, below
the flowers, serve to protect them when they are
pushed up through the soil. The perianth of six
united floral-leaves has the form of a very long,
funnel-like tube, reaching down to the corm. The six
stamens spring from the tube at various levels, but

FIG. XVII.— The Corm or Underground Stem of Colchicum, the
Meadow Saffron.

1. Corm seen from the front. 2. Longitudinal section through corm.

a, corm ; 6, sheathing leaf; c, flowering axis of last year ; d, foliage leaf;
0, leaf on flower-stalk ; /, next year's corm.

the ovary is right at the base of the tube, a foot deep
in the soil. Three very slender, thread-like styles
extend from the ovary, nearly the whole length of the
tube of the perianth.

In describing the flowers of Primula (p. 68), we
drew attention to the occurrence of two forms of
flowers, one with a long style and low stamens, the

168 PLANTS OF THE ALPINE MEADOWS

other with a short style and high stamens. It is
asserted that the Meadow Saffron possesses three
forms of flowers, in which the styles are long, short,
and intermediate in length respectively, and is thus
similar to the well-known case of Lythrum described
by Darwin.

The flowers are probably fertilised by bees. The
seeds remain enclosed in the ovaries, deep in the soil,
throughout the winter, and it is not until the following
spring that they are pushed up above ground with
the leaves, by the growth of the region of the stem
just below the ovaries. Often, however, only one fruit
comes to maturity. It is probable that the whole
structure of the flower is so designed that the fruits
shall remain buried in the soil throughout the winter,
and the seeds thus escape injury from frost. It is
obvious that, to a plant flowering so late in the year
as the Saffron, some contrivance, which will allow
time for the seeds to mature, and preserve them
unharmed from winter frosts, must exist.

The seeds of the Meadow Saffron contain a power-
ful, poisonous alkaloid, known as colchicin, which is
sometimes made use of medicinally.

CHAPTER VII

THE HIGH ALPINE PLANTS

WE now pass to a discussion of the peculiarities of
the highest plant assemblage in the Alps, the species
of the High Alpine region.

It is often thought or assumed that, above a
certain height vaguely imagined to be somewhere
between 10,000 and 13,000 feet, vegetation, or at any
rate Flowering Plants, cannot exist in the Alps. This
idea is entirely fallacious. It is true that in the
higher mountain regions, vegetation is much more
scanty than at elevations of 5,000 or 6,000 feet; but
wherever the physical conditions are in the least degree
favourable, there plants will be found, whatever the
altitude may be.

The physical conditions that an Alpine growing
at a height above 10,000 feet in the Alps has to combat
are no doubt extremely severe. The first necessity for
its existence is absence of snow or ice for a sufficiently
long period from some sheltered spot. Given a flower-
ing season of adequate length, many of the High Alpine
species will be able to overcome most of the other

169

170 THE HIGH ALPINE PLANTS

difficulties. On the other hand, the extreme short-
ness of the flowering season at high altitudes is
probably the factor which, more than any other, limits
the upward distribution of Alpines. Time for
flowering alone is not sufficient. A certain length
of time afterwards must also be available to set
and distribute the seed, to enable the offspring to
take a firm hold in its new home, and to allow the
parent to manufacture reserves against the coming
winter.

The so-called snow-line, or imaginary line above
which the snow continues to lie all through the
summer, varies in elevation according to the aspect,
situation, and other physical conditions of any parti-
cular locality. In general, it lies between 8,500 and
10,500 feet, but is sometimes lower or even higher.
Though above this elevation the coating of snow is
permanent, it is by no means continuous. Kocks
fully exposed to the sun, steep slopes, and precipitous
crags quickly lose their snowy covering, for a time
at least, in summer, though the periods during
which they are free from snow may be only short and
intermittent.

But bare rock alone will rarely furnish a liveli-
hood sufficient to permit the seed of a High Alpine
to establish itself. Some sort of primitive soil, such
as those which we have already discussed (p. 96),
must in most cases be present. In the High Alpine
region the thalli of Crustaceous Lichens appear to
contribute in a large degree to the building up of

DIFFICULTIES OF EXISTENCE 171

primitive soils, and around them wind-blown dust
and debris of all sorts accumulate.

Once a seed becomes established on a primitive
soil on some rocky ledge, free from snow perhaps for
less than two months in the height of summer, it
has to face certain other difficulties. The gap
between the extremes of temperature to which it will
be exposed is enormous. The temperature in the
sun at midday may rise to nearly 20° C. ( = 68° F.), and
sink several degrees below zero C. at night-time.
In winter-time it may fall to - 25° C. or more. Fur-
ther, the difference between sun and shade tempera-
ture in the daytime is much greater than at lower
elevations, so that whenever the sun is hidden by
a passing cloud, a sudden and considerable drop in
temperature takes place.

Another difficulty often to be faced is the lack of
an adequate water-supply for the roots. This is not
due, as is sometimes supposed, to the coldness of the
soil, for it has been shown that, in summer at
any rate, the soil is several degrees warmer than the
atmosphere in the shade, and that the soil is generally
at a higher relative temperature in the Alps than in
the lowlands. But at high altitudes rain rarely falls,
and such water as penetrates to the roots is derived
from melting snow. Since the supply is precarious,
many High Alpine plants possess hairy coats or
other adaptations, designed, as much as possible, to
reduce transpiration or loss of water by evaporation.

Other drawbacks to life in high altitudes are the

172 THE HIGH ALPINE PLANTS

excessive intensity of the light, the relatively stronger
ultra-violet rays, the increased rarity of the atmo-
sphere, the greater force of the wind, and the scarcity
of insect visitors. Yet, strange to say, plants will
overcome all these difficulties successfully, if only a
favourable chance offers.

If the highest of the Swiss peaks were free from
snow and ice for only six weeks or two months in
summer, there is every reason to believe that they
would accommodate a large and varied population of
Alpine plants. The rich flora of the Gorner Grat
(10,290 feet), near Zermatt, an exceptionally favour-
able situation for plant life in the High Alps, enables
us to form some idea of the vegetation we should
meet with at even greater elevations, if the conditions
were similar.

The highest mountain in the Alpine Chain, Mont
Blanc, is 15,782 feet high. It is partly in France
(Savoy), and partly in Italy. Monte Eosa is partly
in Switzerland and partly in Italy, and reaches
15,217 feet. The highest mountain entirely in
Switzerland is the Dom, between the valleys of
Zermatt and Saas ; it is 14,942 feet high.

The greatest height at which a flowering plant has
been found in Switzerland is about 14,107 feet
(4,275 metres), which proves the assertion pre-
viously discussed : that there is no real upward limit
to vegetation in the Alps.

In other parts of the world, vegetation far exceeds
this altitude. On the north side of the Himalayas,

PLANTS AT GREAT ELEVATIONS 173

Flowering Plants occur as high as 18,000 feet, while
in the Bolivian Andes, species have been found
growing at 18,700 feet, if not higher.

The late John Ball, a great authority on Alpine
plants, relates how, when botanising on the Aletsch
Glacier (Bernese Oberland), the largest snowfield in
Europe, he found no less than forty plants in
flower, including the Common Thyme and the still
commoner Dandelion, on a slope of fine debris, clear
of snow, at an elevation of about 10,700 feet. This
is by no means an exceptional instance.

De Saussure, one of the earliest naturalists to
devote serious attention to nature in the Alps, and
the famous leader of the first party to reach the
summit of Mont Blanc, related how, in 1796, he
found Silene acaulis growing on that mountain
at an elevation of 11,450 feet, and Androsace
glacialiSy near the Col de Geant, at about the same
height. The celebrated Swiss botanist, Dr Christ,
states that at least thirteen Flowering Plants have
been found on the Theodule Pass, 10,900 feet,
between Zermatt and Breuil. Here the mean tem-
perature for the year is known to be - 5*59° C.,
the minimum - 21 4° C., and the maximum -f 151° C.

The well-known " Jardin " of the Mer de Glace,
above Chamonix, is perhaps the most reputed of the
higher localities for Alpine flowers in the Alps. The
flora of this favoured spot, on a moraine of the glacier,
having an area of about 7 acres, and forming, as it
were, an island in a sea of ice, has been repeatedly

174 THE HIGH ALPINE PLANTS

recorded. Nearly a hundred Flowering Plants, as
well as a large number of Mosses and Lichens, have
been found. Yet the height of the " Jar din " is only
9,140 feet, which is low in comparison with the other
localities mentioned above. It is nowadays possible
to reach by train to even greater heights, where the
high Alpine flora may be studied !

"We may now enquire what is the highest recorded
species for Switzerland. According to Prof. Schroeter,
Ranunculus glacialis, the Glacial Buttercup, has this
honour. This plant has been found at 14,107 feet
(4,275 metres) on the Finsteraarhorn, the giant of
the Bernese Oberland. Seven other species are
known to occur at or above 13,200 feet (4,000
metres) — namely, Achillea alrata, Saxifraga aspera,
var. bryoides, S. moschata, on the Finsteraarhorn;
Androsace glacialis, on the Lauteraarhorngipfel ; and
Saxifraga muscoides, S. biflora, and Gentiana brachy-
phylla, at 13,860 feet on the "shoulder" of the
Matterhorn.

If we study the distribution of plants within the
Alpine zone — that is, at elevations above 5,000 feet in
altitude — we shall find that between eighty and ninety
species of Flowering Plants are only found in the
higher regions, and do not occur in the lower portion
of this zone. At elevations of above 8,000 or 8,500
feet, we shall notice that the flora is composed of
species such as the Bavarian Gentian, which are
also abundant in the lower Alpine region, and in
addition a number of other plants occur, many of

PLANTS AT GREAT ELEVATIONS 175

which are of rare or local occurrence, and are only
exceptionally met with at lower elevations.

Heer found that between 8,580 and 9,135 feet, 336
species occur in Switzerland as a whole, 294 in
Canton Grisons, 206 in Canton Valais, and 150 in
Canton Berne. As we ascend, the numbers gradually
decrease; at an elevation of about 10,200 feet, the
figures were 120, 32, 118, and 17 respectively; at
11,260 feet the number of species sank to 13, 4, 18,
and 6, while above 12,870 feet only 6 species occur in
the whole of Switzerland, none in the Grisons, 2
in the Valais, and 5 in Canton Berne. Thus the
Valais is much richer in High Alpine species, and
the Bernese Oberland much poorer, than any other
district in Switzerland.

These figures give only an approximate result, for
estimates of the number of species in any district
differ according as the author regards certain plants
as distinct species or only as varieties — a question
which is always one of great difficulty, and
quite impossible to settle, for variation is character-
istic of plants as of animals. But whether we regard
Heer's figures as rather too high or too low is
immaterial. The important point is, that of 336
species occurring at about 8,600 feet, a large number
consist of plants common in the lower Alpine region,
while others are what we will term High Alpines, and
are almost unknown in the lower zone.

Schroeter has made an analysis of the flora
known to occur at, and above, 10,725 feet ( = 3,250

176 THE HIGH ALPINE PLANTS

metres). Of the 73 species or varieties instanced,
40 are Alpine, 30 are High Alpine (Nivial), one,
Thymus serpyllum, is a Lowland plant, and two,
Gentiana verna and Phyteuma corniculatumy are
Subalpine and widely distributed. These figures give
a good idea of the proportion of Alpine to High
Alpine species at this great elevation.

We can thus subdivide the Alpine zone into a
lower and a higher region, the latter characterised by
the presence of certain species, the High Alpines. At
the same time, the transition from the lower Alpine to
the higher Alpine region is perfectly gradual, just as
is the transition from the Subalpine to the Alpine
zone.

The High Alpine species are perhaps the most
interesting of all Swiss plants. They are often
spoken of as constituting the Nivial or Glacial flora
of Switzerland. They present certain peculiarities of
habit which at first sight may appear to be unique.
As a matter of fact, it will be found, if the lower
and higher Alpine floras are closely compared,
that most of these peculiarities are shared by plants
growing in the lower Alpine region, described in the
preceding chapters, though in a less pronounced
degree.

It is customary to speak of the "Alpine habit" as
characteristic of plants growing at great elevations in
the Alps. By this is usually implied nothing more
than their dwarf stature. Perhaps the most striking
feature of Alpine vegetation as a whole is, that the

THE DWARF HABIT 177

plants are distinctly shorter, and much more compact
in build than in the Lowlands. There is a marked
absence in Alpine plants of those features which
gardeners term "leggy" or "weedy." The stems are
relatively shorter, and in many cases are buried below
the surface of the soil. The axes of the flowering shoots
are also less extended. Yet the average Alpine plant
can hardly be called a dwarf, if by that term we imply
a plant which only rises from 1 to 4 inches (2 to 10
cm.) above the level of the soil.

Among the lower Alpine species, such plants
as the Alpenroses, the Yellow- and Eed-flowered
Gentians, the White Veratrum, the Monkshoods, and
the Martagon Lily are very far from being dwarfs. On
the other hand, the majority of the lower Alpines
are short in stature, though not true dwarfs. The
rosette and carpet plants may be regarded as the best
examples of the dwarf plants, but these types of habit
are far from being universal.

When we turn to the High Alpine flora, we find
a decidedly larger number of dwarf plants. The
stature of these species is on the average distinctly
shorter than in the lower region. Yet all High
Alpine plants are not dwarfs, for Adenostyles leuco-
phylla, Reich., is often 3 feet high, Doronicum
(Aronicum) scorpioides, Lam., is from 6 inches to 2
feet in height, and Empetrum nigrum. Linn., is a
shrub varying from 6 to 18 inches in height. On
the other hand, the majority of High Alpine species,
as contrasted with their near relatives flourishing in

178 THE HIGH ALPINE PLANTS

the lower Alpine region, are distinctly smaller plants
and shorter in stature, even if they cannot all be
termed true dwarfs.

The term "Alpine habit," if applied in any
other sense than size, has no botanical significance.
There are several types of Alpine habit or build,
which, with one exception, are found both in the
lower and the higher Alpine regions. Thus, apart
from the cushion plants, which are only represented by
one or two species in the lower Alpine zone, the "archi-
tectural" peculiarities of High Alpines only differ from
those of the lower Alpines in degree, not in kind.
H We will now discuss the different types of habit
met with in the High Alpine region : the cushion plants,
the carpet plants, the rosette plants, and the normal
but dwarf habits.

We have seen that among the Alpines of Switzer-
land many of the species are British plants. When
we turn to the High Alpine plants, it is interesting to
find that several, though a much smaller proportion,
are also British. The Alpine Cerast (Cerastium
alpinum, Linn.), the Crowberry (Empetrum nigrum,
Linn.), the Mountain Lloydia (Lloydiaserotina, Sweet),
the Scotch Asphodel (Tqfieldia palustris, Huds.), and
the Reticulate and Dwarf Willows, are, for example,
almost confined in Switzerland to the High Alpine
region. Several other plants, common both in the
lower and higher Alpine regions, such as the Moss
Campion (Silene acaulis, Linn.) and the Purple
Saxifrage (S. oppositifolia, Linn.), are also British.

HIGH ALPINE CUSHION PLANTS 179

THE CUSHION PLANTS.

The cushion species present a type of plant
architecture or habit essentially characteristic of the
High Alpine region. In the flora of the lower
Alpine region, only two cushion plants, the Moss
Campion and the Purple Saxifrage, are met with, and
these, as we have seen, are also abundant in the higher
Alps.

The cushion plants (Plates XXXIII. and
XXXIV.) are constructed on a definite plan.
There is a simple, unbranched stem buried deep
in the soil. Just above the surface of the soil, the
stem gives off a very large number of leafy shoots or
branches, radiating out, as it were, from the centre
of an hemisphere. The main branches give rise to
secondary branches, also clothed with leaves, which all
grow out to about the same length, and the whole of
the shoots are crowded together into the smallest
possible space (Plate XXXIV.), thus giving rise to a
compact, cushion-shaped structure. The exact form
or shape of the cushion varies in different species, as
illustrated on Plates XXXIV. and XXXV. The
cushions often grow to a very large size. The largest
cushion of the Moss Campion we have measured
was near the Hotel Weissmies (9,180 feet) above
Saas Grund. It was nearly circular, and the diameter
was 3 feet 4 inches, and the height about 4 inches.

Cushion plants, it may be of interest to add, are
also found in many other parts of the world besides

180 THE HIGH ALPINE PLANTS

the high Alps, where one or more of the physical
conditions are similar. They are found in the Arctic
and Antarctic regions, in deserts such as the Sahara,
and in the Steppe region of Russia.

THE COMMON Moss CAMPION.

The moss-like cushions of the Common Moss
Campion (Silene acaulis, Linn., natural order Caryo-
phyllacese, the Pink family) are one of the most
familiar sights in Alpine Switzerland. They are
dark green in colour and flat topped. The leaves,
which arise in opposite pairs, are short, narrow,
and awl-shaped. Each cushion produces an enormous
number of rose-coloured or pink flowers, each borne
singly on a very short flower-stalk, arising from the
axil of a leaf.

There are three kinds of cushions : those bearing
flowers with both stamens and carpels, which are
rare ; those with male only, and those with female
flowers, in which stamens are absent. The last are
smaller than the others.

This British plant is not only common at 5,000 feet
in the Alps, but extends upwards to nearly 12,000 feet.

THE SESSILE-FLOWERED Moss CAMPION.

The Sessile-flowered Moss Campion (Silene
eacscapa. All.) (Plate XXXIII., Fig. 1) is a very near
relative of the species just discussed, and may be
only a variety of it. It differs in having only a very
short flower-stalk, which is winged, and in certain

PLATE XXXIII.

FIG. 1.— The Sessile-flowered Moss Campion (Silene exscapa, All.).

FIG. 2.— The Swiss Androsace (Androsace helvetica, Gaud.).
Typical High Alpine Cushion Plants.

[To face p. 180.

THE HIGH ALPINE ANDROSACES 181

peculiarities of the calyx and fruits. Its cushions
are commonly mistaken for those of Silene acaulis,
but they are more densely tufted, more spherical in
shape, and the leaves shorter. The flowers are also
smaller and paler. It is generally believed to be
confined in Switzerland to the High Alpine region.
It is not a British plant.

THE HIGH ALPINE ANDROSACES.

The Androsaces of the lower Alpine region
described in Chapter III. are typical rosette plants.
Yet all the High Alpine species possess the cushion
habit. The Swiss Androsace (Androsace helvetica,
Gaud., natural order Primulacese, the Primrose family)
(Plate XXXIV., Fig. 1) is a typical example. This
plant builds cushions, resembling an hemisphere in
shape, and sometimes 6 inches high. These are formed
by a large number of crowded branches, each branch
clothed with very small, blunt, over-lapping leaves,
which form compact, cylindrical, bud-like growths at
the ends of the branches. The leaves are covered
with simple hairs. The flowers are borne on very
short stalks, the corollas being white with a yellow
centre. This plant is not uncommon in the fissures
of calcareous rocks at great elevations. The cushions
are said to reach a considerable age, 50 to 60 years
being reported in one case.

The much rarer Imbricated Androsace (Androsace
imbricata, Lam.) builds cushions similar to those of
the Swiss Androsace. They are, however, easily

182 THE HIGH ALPINE PLANTS

distinguished by the dense, grey felt of star-shaped
hairs which covers the blunt lance-shaped leaves.
The flowers are rose coloured or white, with a red
" eye " at the throat of the corolla.

In the next three species, the cushions are
fashioned on a somewhat different plan. They are
hemispherical in shape, but much less compact. The
leaves at the end of the branches, instead of forming
dense bud-like growths, are arranged in fairly open
rosettes. The older leaves below the rosettes tend to
die off, so that the cushion is only leafy towards the
surface of the sphere. Thus we have here a combina-
tion of the cushion and rosette habits.

The Glacial Androsace (Androsace glacialis,
Hopp.) (Plate XXXIV., Fig. 1) is a typical
example, and possesses rose-coloured corollas. A
rare species, found only in Canton Tessin, Char-
pentier's Androsace (Androsace charpentieri, Heer.),
is very similar, but the flowers are mounted on
longer stalks, and differ also in a number of
other details. The Downy Androsace (Androsace
pubescens, D. C.) has cushions quite like those of
the last two species, but white flowers with a yellow
" eye." It occurs chiefly in the Valais.

Vital's Androsace (Androsace vitaliana, Lap.,
= Aretia vitaliana, Murr.1) (Plate XXXVIII., Fig. 1)

1 Also known as Gregoria vitaliana, Duby. According to Index
Kewensis, this plant should be called Douglasia vitaliana, B. and H.f .
I have, however, here included it in the genus Aiidrosace, to which, even
if it should be referred to a separate genus, it is very nearly related.

PLATE XXXIV.

THE DRABA AND ERITRICHIUM 183

forms loose, spreading cushions, the ends of the leafy
branches being resetted. Numerous large, yellow
flowers spring from the cushion, and these differ
from other Androsaces in the greater length of the
corolla tube, which is twice the length of the calyx.
The free portions of the corolla spread to form a
saucer-shaped structure. It is a rare plant, confined
to Canton Valais.

THE PYRENEAN DRABA.

The Pyrenean Draba (Draba pyrenaica, Linn. =
Petrocallis pyrenaica, Linn., natural order Cruciferse,
the Crucifer family), is another example of a High
Alpine cushion plant, whose near relatives in the
Alpine region proper, such as Draba aizoides, Linn.,
and D. tomentosa, Wahl., are rosette plants. At first
sight the cushion of the Pyrenean Draba, with its
pink or lilac flowers, might be easily mistaken for
that of Androsace glacialis. It is, however, at once
distinguished by the four petals, and the Crucifer
type of the flower as a whole, and by the fact that
the leaves at their tips are cleft into three or more
lobes. It is not a very abundant plant, but is not
infrequent on calcareous rocks and debris in the High
Alpine region.

THE ERITRICHIUM.

The Eritrichium (Eritrichium nanum, Schrad.,
natural order Boraginese, the Borage family) (Plate
XXXV., Fig. 1) is in some respects the "belle"

184 THE HIGH ALPINE PLANTS

of the High Alpines, on account of the wonderful
bright, azure blue of the flowers, for which it is
famous. It is closely allied to the Forget-me-nots,
one species of which, Myosotis alpestris, Schmidt, is
frequent in the Alpine and High Alpine regions.
They so closely resemble one another, that the
Forget-me-not is sometimes mistaken for the Eritri-
chium. The two are not uncommonly associated.

The plant forms rather loose cushions of highly
branched, leafy shoots crowded together. The leaves
are covered with shining, silky hairs, which are clearly
seen in the photograph on Plate XXXV., Fig. 1).
The cushions apparently attain to a considerable age,
for, like nearly all the High Alpines, the Eritrichium
is a perennial. A cushion thirty years old has been
recorded. The flower-stems are leafy and a few
inches in length. They bear, as a rule, from three to
six flowers, arranged in a complicated inflorescence.

THE HIGH ALPINE ALSINES.

There are two High Alpine species of the genus
Alsine (natural order Caryophyllaceae, the Pink
family) which are cushion plants. The Dwarf Alsine
(Alsine sedoides, Fradl^Cherleria sedoides, Linn. =
Arenaria Cherleria, Hook.f.) (Plate XXXV., Fig. 2)
builds very compact, hemispherical cushions composed
of an enormous number of small branches, clothed
with awl-shaped leaves placed in opposite pairs. The
flowers are interesting from the fact that they are
small and comparatively inconspicuous, and thus the

PLATE XXXV.

THE HIGH ALPINE SAXIFRAGES 185

plant contrasts rather markedly with those already
described. The flowers are borne singly, and are
shortly stalked. As a rule, the corolla is entirely
absent, or, if present, the petals are very reduced and
minute. The sepals forming the conspicuous portion
of the flower have membranous margins.

The other species, Alsine aretioides, M. K. ( = A.
octandra, Schur.), is confined to calcareous soils in
Canton Valais. It builds similar cushions. The
flowers in this case possess petals which, like the
sepals, are four in number, while there are eight
stamens, an unusual occurrence in this order of
plants.

THE HIGH ALPINE SAXIFRAGES.

Many of the Saxifrages, as we have seen (Chapter
III.), are typical rosette plants. A few form cushions
by a close aggregation of bud-like shoots. Such
species are not, however, confined to the High
Alpine region, but also occur at lower elevations.

Sometimes in the High Alpine region Saxifraga
moschata, Wulf ( = S. varians, Sieb.), builds cushions,
though the plant is only tufted at lower altitudes.
Saxifraga bryoides, Linn., which probably is simply
a High Alpine variety of the Rough Saxifrage
(Saxifraga aspera, Linn.) (see p. 81), forms cushions
by an aggregation of many bud-like shoots, arising in
the axils of the leaves of older branches. The
yellowish-white flowers are borne on erect shoots.

We have already discussed the Purple Saxifrage

186 THE HIGH ALPINE PLANTS

(Saxifraga oppositifolia, Linn.) (Plate XXXVI.,
Fig. 1) in Chapter III. (p. 78). We may, however,
notice here that the tufted leafy branches of this
plant form a loose cushion, less compact than those
of many of the High Alpines. Though the Purple
Saxifrage is found at great elevations in Switzerland,
it is not a true High Alpine, for it is also common
in the Alpine and Subalpine zones.

The large size of the flowers in comparison with
the leaves (Plate XXXVI., Fig. 1) is a striking feature
of the plant. Often when this Saxifrage is in full
bloom, the crowded flowers almost completely hide the
cushion. The same feature is characteristic of many
other cushion plants, though it is not universal.

THE HIGH ALPINE CARPET PLANTS.

We have already discussed the peculiarities of a
typical carpet plant in the case of the White Dryas
(p. 106), the Trailing Azalea (p. 109), and other Alpines
described in Chapter IV.

The Trailing Azalea is essentially a High Alpine
plant. Another High Alpine which will be found
described in Chapter IX., the Crowberry (Empetrum
nigrum, Linn.), is a low, spreading shrub which, while
not a true carpet plant, has some characteristics in
common with typical instances of that kind of habit.

In the High Alpine region, the Willows form
perhaps the most perfect examples of carpet plants
to be found in the whole Alps.

PLATE XXXVI.

* c

A a

"? s

<u §

II
Is

THE RETICULATE WILLOW 187

THE HIGH ALPINE WILLOWS.

We are accustomed to think of the Willows
(natural order Salicacese) as being good-sized trees,
as with us in England. Even in the Alps the Willows
manage to hold their own, owing to their marvellous
power of adapting themselves to a set of physical con-
ditions, entirely different from those met with in the
plains. As they ascend the Alps, they discard the
tree habit. They gradually become reduced to dwarf
shrubs, and then finally in the High Alpine regions
to carpet plants, which are only slightly woody. The
gradual dwarfing of a tree-form, as the altitude
increases, is well seen in other plants, such as the
Juniper and the Mountain Pine, which will be dis-
cussed in a later chapter, but nowhere is it found
perfected to the degree met with in the High Alpine
Willows.

The carpet-forming Willows do not belong to the
same species as those found in the plains, nor are
they regarded as varieties of Lowland species, as is
the case with Juniperus communis, var. nana, Willd.
There are two Carpet Willows confined to the High
Alpine region, which we will now compare.

THE RETICULATE WILLOW.

The Reticulate Willow (Salix reticulata, Linn.)
(Plate XXXVII.) is the best known and the
handsomest of the Alpine Willows. It is usually
very abundant on old or new moraines of glaciers,

188 THE HIGH ALPINE PLANTS

which are excellent places on which to study these
Willows, as well as other plants.

The habit of this species, as seen on Plate
XXXVII., is that of a typical carpet plant. There
is a stem, for the most part buried in the scanty
soil, which gives off numerous branches, radiating
in all directions and placed close to the ground.
The plant may attain to a good age. Prof. Schroeter
records forty-one years in one case. In other Alpine
Willows the period may be even greater.

The large, elliptical leaves are very characteristic.
The upper surface is smooth, shining, and dark green
in colour; the veins are extremely prominent, and
form a well-marked mesh- work or reticulation, clearly
seen on Plate XXXVII. The prominence of the net
veins is a special feature of this Willow ; hence the
specific name "reticulata." On the lower surface the
leaves are covered with a fairly thick felt of bluish-
white cotton-hairs. If we examine young leaves, we
shall find that they are hairy all over. As the leaf
matures, the hairs disappear completely from the
upper surface. The leaves are quite entire — i.e., not
toothed at the margin. They are mounted on long,
pinkish leaf-stalks. Sometimes the leaves are some-
what rolled at the edges, an adaptation which, like
the felt of hairs on the lower surface, serves to protect
them against undue loss of water by evaporation, in
the manner already explained.

The flowers of the Willows are very different to
those of all the other Flowering Plants discussed in

PLATE XXXVII.

I e

!§>

THE RETICULATE WILLOW 189

this volume. They are very reduced structures
without either sepals or petals, and the male and
female organs are borne on separate plants. Each
male flower consists solely of two or more stamens,
in the axil of a modified leaf or bract. Each female
flower is composed of a single carpel, also in the axil
of a bract. Both types of flower also contain a honey
gland. The flowers, whether male or female, and the
bracts, are arranged in dense spikes called catkins.

The photograph on Plate XXXVII. is of a male
plant of the Eeticulate Willow. The erect male
catkins are borne on rather long, leafless stalks,
springing from the ends of branches. The bracts of
the catkins are brown in colour. The withered
stamens, of which there are two to each flower in this
species, can just be seen with the use of a hand-lens,
the anthers projecting beyond the bracts. The
catkins were past their prime when the photograph
was taken.

The female flowers produce a large number of
very small seeds. Each seed has a little tuft of hairs
at the base, which helps it to fly on a windy day and
so distributes it far from the parent. It will be often
noticed that the Willow carpets growing on moraines
are covered with a whitish fluff, somewhat resembling
cotton-wool. This fluff consists of enormous numbers
of seeds tangled together by their hairs. The Alpine
Willow-herbs (Epilobium, natural order Onagraceee)
produce similar seeds, with hairs at the base, which
often become entangled in much the same way.

190 THE HIGH ALPINE PLANTS

The Eeticulate Willow ascends to 10,450 feet on
Monte Eosa. It is fond of calcareous rocks. It is,
of course, a British plant, though confined to some
of our loftiest hills in Scotland.

THE DWARF WILLOW.

The Dwarf Willow (Salix herbacea, Linn.) was
described by Linnaeus as " the smallest of all trees "
(minima inter omnes arbores). It is essentially
similar in habit to the Eeticulate Willow, from which
it differs only in detail. It is also a British plant.

The leaves are smaller, possessing only a short
stalk, and are quite destitute of hairs underneath.
They are green and shining on both sides, and have
a net nervation. They are finely but bluntly toothed
at the margin. The catkins are very small and few
flowered, and are borne on short stalks.

There are several other Willows — such as Salix
glauca, Linn., which is not infrequent in the High
Alps, and has lance-like leaves with long, straight,
silky hairs on both sides, also Salix retusa, Linn.,
S. ccesia, Vill., and others which are found in the
Alpine zone — but only the Eeticulate and Dwarf
Willows are exclusively High Alpine.

HIGH ALPINE EOSETTE PLANTS.

In the typical rosette plant, the above-ground
portion of the stem is very short, and the bases of
the leaves are all crowded together in the form of a

ROSETTE PLANTS 191

rosette. This plan of architecture is by no means
confined to Alpine species. Our British Whitlow-
grass (Draba verna, Linn.), a Lowland species, is a
typical rosette plant. This habit is, however, more
common in the Alpine zone than in the Lowlands.
We have already noticed some typical Alpine rosette
plants in Chapters II. and III.

Within the Alpine zone, as has been already
pointed out, a large number of species tend to become
marked geophytes — that is to say, the stem is buried
as deeply as possible in the soil, and the upper
portion is reduced. The consequence is that the
spaces (internodes) between the points of attachment
of the leaves to the stem (nodes) are shortened. Thus
the great feature of a rosette plant is, in botanical
jargon, the suppression of the internodes of the stem.
We can actually see this condensation of the inter-
nodes if we compare specimens of the Bavarian
Gentian (Gentiana bavarica) from the lower and higher
Alpine regions. At the lower level, this plant is not
a rosette plant. Quite long spaces or internodes
occur between the lower leaves. From higher
habitats, however, the internodes will be found to
become shorter and shorter, and finally an imbricated
or overlapping rosette results.

It may be doubted whether the flora of the Higher
Alpine region is so rich in rosette plants as the lower.
Probably there are actually fewer above 8,000 feet
than below. Some of the typical Alpine rosette
species, as we have seen, take to cushion building, so

192 THE HIGH ALPINE PLANTS

to speak, in the High Alps, as, for instance, the
Androsaces. On the other hand, the Gentians are
represented by two High Alpine species, Gentiana
brachyphylla, Vill., and G. tenella, Rottb. (see
Chapter II.), which are rosette plants, but the High
Alpine Saxifrages, Saxifraga androsacea, Linn. (Plate
XXXVI., Fig. 2), S. Seguieri, Spr., S. muscoides, All.
( = S. planifolia, Lap.,) S. exarata, Vill., and S. aphylla,
Sternb. ( = S. stenopetala, Gaud.), are all tufted or
cushion-building species, and not true rosette plants.
Of the three High Alpine Rampions, Phyteuma
pamiflorum, Linn. (P. pedemontanum, Schulz), alone
has true rosettes, for P. humile, Schleich, and P.
hemisphwricum, Linn., are tufted plants. The blue-
flowered Arabis c&rulea, Haenke, however, with
Draba Wahlenbergii, Hartm., and D. carinthiaca,
Hoppe ( = D. johannis, Host.), all three Cruciferous
species, go to swell the number of rosette plants in
the High Alpine zone.

HIGH ALPINE DWARF PLANTS.

For want of a better term, we may include under
the heading of High Alpine dwarf plants, those species
growing at great elevations in the Swiss Alps, which,
as regards their habit, are not markedly dissimilar
except in stature, from their near relatives in the
Lowlands. Many of them rarely exceed 4 inches in
height, and they are often less. The leaves, for the
most part, are borne just above the surface of the
ground, and though they are not arranged in true

THE HIGH ALPINE BUTTERCUPS 193

rosettes, they are in many instances tufted. We will
now notice a few of the more interesting examples.

THE HIGH ALPINE BUTTERCUPS.

There are four Buttercups, if we include the
Kue-leaved Callianthemum, which are confined to
the upper portion of the Alpine zone, the High Alpine
region. Of these the Glacial Buttercup (Ranunculus
glacialis, Linn., natural order Eanunculacese, the
Buttercup family) is especially interesting because, as
we have seen, it is the highest plant found in
Switzerland. It flourishes on very damp, gravelly
or rocky places, and often on fully exposed, sunny
slopes. The whole plant varies from 2 to 7 inches in
height. There is a very short stem rooted in the
gravelly soil, bearing leaves mounted on fairly long
stalks. Each leaf is divided into three portions,
which, again, are either lobed, cut, or divided. It is
characteristic of the Glacial Buttercup that the leaves
of different plants vary enormously in the degree to
which the three segments are lobed or divided.

We should rather expect to find in a plant living
at such great elevations — for the Glacial Buttercup
rarely occurs lower than 7,600 feet — that the leaves
would be covered with a thick coat of hairs, affording
them some protection against the severe climatic
conditions of these high places. Yet, except for the
calyx, the whole plant of the Glacial Buttercup is^ as a
rule, quite smooth or nearly hairless, and, externally
at least, shows no special adaptation to its particular

194 THE HIGH ALPINE PLANTS

environment. It is only in the most elevated habitats
that the very dwarf specimens become covered with
hairs.

The flowers of this plant are very beautiful and
are easily distinguished from all other Buttercups by
the fact that the outer surface of the calyx is thickly
covered with reddish-brown hairs. Usually only one
flower, but sometimes as many as three, are borne on
each flowering shoot, which also carries a few leaves,
smaller in size than those found below. The colour
of the corolla varies from pure white to rose-pink, or
very dark pink-red. It is very inconstant, great
differences in colour being often remarked among the
flowers borne on the same plant, and even in the
different petals of the same flower. The petals have
a small honey nectary at the base on the inner side,
and these organs, again, vary greatly in size and com-
plexity, in different examples of the plant. Otherwise
Ranunculus glacialis is quite a typical Buttercup,
though of dwarf stature.

The three other Buttercups have white flowers, and
also offer an interesting contrast when the shape of
the leaves is compared.

The Pyrenean Buttercup (Ranunculus pyrenceus,
Linn.), which is common at elevations from 6,000 to
8,000 feet or more, has one to three grass-like, lance-
shaped leaves, blue-green in colour. The sepals are
hairless. It is a dwarf plant, from 3 to 12 inches in
height.

The rarer Parnassus-leaved Buttercup (Ranunculus

THE ALPINE POPPY 195

parnassifolius, Linn.), 2 to 6 inches high, shares with
the Pyrenean Buttercup the distinction of being the
only other Alpine species, which has undivided and
non-lobed leaves. The thick leaves are here large
and heart-shaped, and borne on leaf-stalks. The
nerves on the upper surface are very well marked.
The leaves on the flowering shoots sheathe the axis
at their base. The whole of the lower portion of
the plant is apt to be very hairy.

The Rue-leaved Callianthemum (Callianthemum
rutcefolium, Reichb.), sometimes included in the genus
Ranunculus, and its variety C. coriandrifolium, Reichb.,
by some considered as a distinct species, are among
the rarer and more local High Alpines. They have
highly compound leaves with long stalks, the segments
of the leaves resembling those of the Rue (Ruta).
The petals are white, with a yellow claw.

THE ALPINE POPPY.

The Alpine Poppy (Papaver nudicaule, Linn.,
natural order Papaveraceae, the Poppy family) is a
beautiful High Alpine of local occurrence, chiefly on
calcareous soils. It varies from about 2 to 6 inches
in height ; otherwise it closely resembles the ordinary
Lowland Poppies in habit, except that the flowering
shoots are leafless. It is, however, a perennial plant,
whereas many of the Poppies of the plains are annuals.
This plant is remarkable for the large number of
variations to be found in some of its principal

196 THE HIGH ALPINE PLANTS

characters. The varieties are regarded by some
authors as constituting distinct species.

The plants may have white flowers, and the leaves
be destitute of hairs (Papaver alpinum, Linn.), or the
flowers may be white with a yellow base, or again, the
flowers may be yellow and the leaves hairy (P. pyren-
aicum, Willd. =P. aurantiacum, Lois.). The shape
of the lobes of the much-divided leaves also varies in
great degree.

THE ALPINE BITTERCRESS.

The Alpine Bitterer ess (Cardamine alpina, Willd.,
natural order Cruciferse, the Crucifer family) is the
representative of the Ladies' Smock or Cuckoo-flower
(Cardamine pratensis, Linn.) in the High Alpine
regions. It is a dwarf plant, from 1 to 5 inches in
height, with many long-stalked, undivided leaves
below, and numerous leafy shoots above. The
simple undivided leaves contrast with the compound
leaves of the Ladies' Smock.

THE SHORT-STEMMED HUTCHINSIA.
The Short-stemmed Hutchinsia (Hutchinsia brevi-

iSy Hoppe) is possibly only a High Alpine variety
of the Alpine species, H. alpina, E. Br. Both
differ from our British Kock Hutchinsia (H. petrcea,
K. Br.) in the flowering stems being simple, un-
branched, and leafless. The white petals are also
much longer than the calyx, and therefore more

THE HIGH ALPINE POTENTILLAS 197

conspicuous, and the whole flower, though small, is
larger than that of Hutchinsia petrcea. H. brevicaulis
differs chiefly from H. alpina in the shorter flowering
shoot, and the more compact and dwarf habit. In these
three species, all of which occur in Switzerland, one
can trace each stage in the production of a dwarf
High Alpine.

THE FRIGID AND SMALLEST POTENTILLAS.

The genus Potentilla (natural order Kosaceae, the
Rose family) is well represented in the Alps by
numerous species. Several British plants, including
the Tormentil (Potentilla tormentilla, Neck.) and the
Spring Potentilla (P. verna, Linn.), are frequent in the
Alpine zone, where also many non-British species,
such as the beautiful Large-flowered Potentilla (Poten-
tilla grandiftora, Linn.), are as abundant. In the High
Alpine region we find two very dwarf species, both
with stems only 1 to 2 inches high, and usually only
one or two very small flowers on each shoot. The
leaves of the Frigid Potentilla (Potentilla frigida,
Vill.) are dull green, and very hairy on both sides.
Those of the Smallest Potentilla (Potentilla minima,
Hall.) are smooth on the upper surface, and bright green
in colour. The Scotch Sibbaldia (Potentilla Sibbaldi,
Haller, = Sibbaldia procumbens, Linn.) also occurs in
the High Alpine region in Switzerland.

198 THE HIGH ALPINE PLANTS

THE ALPINE OX-EYE DAISY.

The Ox-eye Daisy (Chrysanthemum leucanthemum,
Linn. = Leucanthemum vulgare, Lam.) is very abundant
nearly everywhere in the Alpine zone. In the high
Alpine region we meet with the Alpine Ox-eye Daisy
(Chrysanthemum alpinum, Linn.), a much smaller plant.
Whereas the flowering stems of the former species are
1 to 2 feet tall, those of the latter are only 1J to 4
inches in height. The Alpine Ox-eye Daisy is, on the
whole, closely similar to the Common Ox-eye Daisy,
though it differs in some characters as regards the
leaves.

PLATE XXXVIII.

FIG. 1. — Vital's Androsace (Androsace vitaliana, Lap.).

FIG. 2. — A Colony of the Common Butterwort (Pinguicula vulgarix. Linn.).

[To face p. 198.

CHAPTER VIII

THE HIGH ALPINE PLANTS (continued) — MARSH PLANTS

THE methods and processes, by which some of the
results achieved by Science are attained, are apt to be
beyond the imagination of the lay mind. Most people
are perhaps aware that the earth has on several
occasions not only been measured but weighed. It is
less widely known that Alpine plants have actually
been made. How these matters are accomplished
is too often regarded as the secret of the scientist,
and certainly it would be difficult to demonstrate the
method of ascertaining the volume and weight of the
earth, without assuming a considerable knowledge of
the groundwork of physical science. But the making
of an Alpine plant is quite simple of comprehension.

We have discussed in the previous chapters the
various types of habit, which are characteristic of
Alpine plants. When, therefore, we say that Alpine
plants have been made, we mean that Lowland plants
transplanted to the High Alps have been found to
assume one or other of those peculiarities of form

199

200 THE HIGH ALPINE PLANTS

and structure, which we have seen to be distinctive of
Alpine habitats.

Several botanists in the past have paid special
attention to this matter. The experimental researches
of Prof. Bonnier of Paris on the adaptation of plants
to Alpine climates are of particular importance in
this connection. We will now glance briefly at his
methods and results. Prof. Bonnier studied a number
of species, among others the Harebell (Campanula
rotundifolia, Linn.), the Kidney Vetch (Anthyllis
vulneraria, Linn.), the Bird's-foot Trefoil (Lotus
corniculatus, Linn.), the Ling (Calluna vulgaris,
Salisb.) and the Rock Silene (Silene rupestris, Linn.).
Strong, well-grown examples of these plants from the
Lowlands were divided into two halves, as nearly
similar as possible. One half of each plant was
transported to one or other of the experimental gardens
on the Mont Blanc range, situated at 3,460 feet and
7,590 feet respectively, or in the Pyrenees to one of
three small gardens at 2,470 feet, 4,950 feet, and
7,920 feet respectively. The other half was cultivated
at Paris (105 feet above sea-level). The soil used in
each case was identical, and everything was precisely
similar, except the physical conditions of the climate
of each experimental station, which varied with the
altitude above sea-level.

The experiment was a particularly fair one, for
the plant grown as a "control" in the plains at Paris
was derived originally from the same individual as
the plant in one of the Alpine gardens. In such

EXPERIMENTAL CULTIVATIONS

201

experiments "controls," or plants grown under
normal conditions, are always used as the basis
of comparison with the plants which are being
investigated,

The results of Prof. Bonnier's experiments were
remarkable. It will not be necessary to discuss here
the different effects pro-
duced on each of the
very large number of
species with which he
worked. The con-
clusions were similar
in each case, though
differing in detail. In
several instances he
found that the plants
grown in the Alpine
regions became dwarf
plants, possessing many

Of the Characteristics FIG. XVIIL— Plants of the Harebell

P A , . . . (Campanula rotundifolia, Linn.).

Of Alpine Species; in That on the left grown in the Lowlands ;
Short, he made Alpine that on the right in an Alpine garden.

(After Bonnier.)

plants.

The case of the Harebell (Campanula rotundifolia,
Linn.) — of which two examples are seen in Text-fig.
XVIII., taken from Prof. Bonnier's memoir — will
afford a good illustration of the effect of the High
Alpine climate. The plant on the left is the " control "
grown at Paris. The one on the right is the other
part of what was originally the same plant, grown in

202 THE HIGH ALPINE PLANTS

one of the Alpine gardens. The latter is a dwarf
plant. The stem is shorter, and the lower leaves
closer together. The flowering shoot is much shorter
and more hairy. It bears only a single flower,
which is, however, larger than any one of the flowers
borne in the branched raceme of the Lowland plant.
The coloration of the Alpine specimen is also a
very much deeper blue.

The following are, in general, the conclusions to
which Prof. Bonnier was led, though in some cases
comparatively little modification was observed. The
whole habit of the species cultivated in the mountains
was much shorter and more dwarf than in the plains.
Sometimes they only reached one-tenth of the height
of the Lowland examples. The stem was shorter, and
much buried in the earth. The underground portions
of the stem and roots were better developed, the
above-ground stems and shoots were more hairy and
spreading, and tended to cling closer to the soil.
The internal structure of both stem and leaf was
profoundly modified. The leaves were, in general,
nearer together, more hairy, relatively thicker and
smaller, and of a much deeper green colour, more
chlorophyll (see p. 10) being developed. The flowers
were relatively larger and more strongly coloured.

The three principal factors of the Alpine climate
which call forth these changes are the more intense
illumination, the drier atmosphere, and the lower
average temperature. The first has a particularly
powerful influence.

EXPERIMENTAL CULTIVATIONS 203

Some earlier experiments by the Viennese botanist,
Kerner, made between 1875 and 1880, are worthy of
consideration in conjunction with Prof. Bonnier's
more recent work.

Kerner established an experimental garden near
the summit of the Blaser (7,243 feet), a mountain in
Tyrol, and the controls were grown in the botanical
garden at Vienna. Of a large number of annuals
raised from seed, many on the Blaser perished from
the severe frosts in spring. Those which survived and
flowered possessed extremely short internodes or
lengths of stem between the leaves. Also, the number
of internodes developed was often little more than half
those found in the controls. The number of flowers
was less, and they were smaller than in the controls.

A biennial plant, an Umbellifer, experimented
with, produced only five umbels as against twenty
found in the control. The internodes were again half
as numerous, and the whole plant was less than a
quarter the height of the Lowland example. So also
with the perennials. In the case of the Grass-of-
Parnassus, the stem in the Alpine cultivation was
only one-third to one-quarter the height of the control,
and the size of the leaves and of the flowers was
smaller. In many species the flowers were more
intensely coloured. Thus, Kernel's researches agree
in several points with those of Professor Bonnier,
and he, too, was inclined to lay special stress on the
influence of the intense illumination of the Alps, as
being the chief factor concerned in these modifications.

204 MARSH PLANTS

MARSH PLANTS.

The concluding part of this chapter will be devoted
to the Alpine marsh plants — not a very large class.

To avoid excessive competition, and to mitigate
the struggle for existence, plants have specialised in
different directions. Some have taken to the water
entirely, others to marshes or perpetually damp
localities, while a third group favours soils with high
water-contents, such as peat. On the other hand,
certain plants, such as the Edelweiss, already dis-
cussed (p. 15), have specialised in quite the opposite
direction, and flourish on soils and in situations
where the water-supply is of the scantiest nature.
There remain a third set of plants which exist under
conditions midway between these two extremes.
There is no doubt that all the Higher Plants
( Angiosperms) which have become aquatic or adapted
to marshy places have been derived from the last
class, for although in the first place all land plants
were evolved from aquatic ancestors, these ancestors
existed at a very distant geological period, long before
the Higher Plants came into existence. In the case
of present-day aquatics, we have merely an interesting
case of a return to the far-away and remotely primitive
habitat.

In the present chapter we may consider the plants
of the Alpine marshes, and more especially the species
which are restricted to open situations, in which the
soil is, as a rule, constantly moist.

THE MARSH MARIGOLD 205

THE MARSH MARIGOLD,

True aquatics are almost unknown in Alpine
Switzerland. One or two species of Pondweed
(Potamogeton, natural order Naiadacese), occurring in
Alpine lakes, are the sole representatives of what in
the Lowlands is an important plant association. Nor
are marsh plants so numerous as at lower elevations.
The British Marsh Marigold (Caltha palustris, Linn.,
natural order Banunculaeeee, the Buttercup family)
(Plate XXXIX.) is probably the most abundant
representative of this class of plants in the Alpine
region. Its flowers are interesting from the fact that,
as compared with the Buttercup, its near relative, it
has no petals. There are five large, yellow sepals,
however, which are brightly coloured, and resemble
the petals of a Buttercup rather closely, and also serve
as an attractive insect advertisement. They furnish
a good illustration of how Nature attains to the same
end by a variety of means.

The Alps are an excellent hunting-ground in
which to pursue a study of plants, which are also
members of the British flora. Paradoxical as it may
seem, our British flora can, in some respects, be better
studied in the Alps than at home. Many of our
more interesting British plants are rare or of very
local occurrence, and unless we happen to be in a
certain district at the right time of year, and further,
to possess a more or less exact knowledge of the
places in which they flourish, our chances of coming

206 MARSH PLANTS

across them are small. On the other hand, many of
the plants, which are local with us, are in the Alps
often extraordinarily abundant. Nearly all British
Alpines are extremely common in Switzerland. Thus
those who are to some degree familiar with our
British plants can extend their knowledge by further
studies within the Swiss Alpine zone.

It is also a matter of common remark that some
of the most frequent of Alpine plants are also
abundant with us in England. Here is our British
Marsh Marigold, there our British Harebell, flourish-
ing, if anything, more vigorously than with us ! Thus
those who are familiar with our wild plants will find
many old friends within the Alpine zone in Switzerland.
There are at least 250 Lowland species which ascend
to heights of 5,000 feet or more in the Alps. Further,
a very large number, certainly a majority, of Swiss
Alpine plants which do not occur in Britain are very
closely related to species or genera found wild with
us. We will, however, reserve for the last chapter
some discussion on the relationship of the Swiss
Alpine flora to that of Britain and Northern Europe,
and the theories as to its origin.

The photograph of a Caltha Marsh on the Engstlen
Alp (Canton Berne), shown on Plate XXXIX., gives
a good idea of how vigorously the Marsh Marigold
thrives at an elevation of over 6,000 feet in the Alps.

PLATE XXXIX.

THE GLOBE-FLOWER 207

THE GLOBE-FLOWER.

The Globe-flower (Trollius europceus, Linn.)
(Frontispiece), belonging to the same family as Caltha,
is an abundant plant in more or less wet habitats in
the Alps. With us in Britain it is much less frequent
than the Marsh Marigold. In Switzerland it does
not flourish, as a rule, in decidedly marshy places where
Caltha may be found, for while apparently requiring
a very damp soil, it is a much less pronounced marsh
plant than the latter. At the same time, its distribu-
tion is rigidly restricted in accordance with the water
contents of the soil, though its requirements in this
respect are less exacting.

The flowers of Trollius are very interesting and
somewhat exceptional. They never open. Here,
again, it is the sepals, and not the petals, which form
the conspicuous floral envelope. They may be ten to
fifteen or more in number, and are yellow in colour.
They are all bent on themselves, so that they converge
towards the summit of the flower, overlapping one
another, and forming, as it were, a dome-shaped roof
over all the other parts or organs of the flower ; hence
the name Globe-flower. The sepals here perform the
same function as in Caltha. If we dissect them away,
we shall find they enclose an equal, or nearly equal
number, usually about thirteen, small, flat honey-
glands, which by some are regarded as the real
petals, considerably modified to serve as nectaries.
Others hold that they are more probably derived by

208 MARSH PLANTS

modification of some of the outer stamens. The
real stamens, which are numerous, lie more internally
still, and then, in the centre of the flower, we find
several carpels quite free from one another. Honey-
glands, even more highly modified, occur in the flowers
of several other members of the same family, Kanun-
culacese, as, for instance, in the Hellebores (Christmas
Eose, etc.), where they are fairly large and horn-like
in shape, and in Eranthis, the Winter Aconite, where
they are tubular.

The presence of honey-glands and a copious
supply of honey implies that this flower is fertilised
by insects ; yet the dome-shaped roof of the converging
sepals never opens. How, then, do the insects manage
to get inside? The mystery can be explained by
anyone who will take the trouble to watch these
flowers for a few minutes on a bright sunny day.
Before very long, one or more small flies will be seen
to alight on the sepals, push their way between them,
and disappear bodily into the interior. Later on
they will be seen to creep out again. It is these flies,
attracted by the honey, which serve as pollen carriers
from flower to flower. No bee can get at the honey
because of the roof of sepals. Thus we have here an
interesting contrivance for cross-pollination.

Although the nectar, copiously secreted in the
honey-glands, is strictly reserved for certain small flies,
other insects, which are unbidden guests, make
determined attempts to reach the honey and to rob
the flower. Their chief difficulty is that their bodies

THE ACONITE-LEAVED BUTTERCUP 209

are too big to allow them to creep between the
sepals.

The photograph shown in the Frontispiece is
fortunate in that it exhibits examples of both the
bidden and unbidden guests of this flower. On the
top of the highest flower seen in the photograph, one
of the small flies, a legitimate guest, is just visible
(though not very distinctly) returning to the world.
On the left side of the same flower, a much larger
insect, a robber, probably a beetle, is at work. Unable
to get within by legitimate means, it tries to bite
through, and to tear off the sepals. Several of these
flowers show sepals, the margins of which have been
bitten away, and which consequently have turned
slightly brown at the edges.

THE ACONITE-LEAVED BUTTERCUP.

A conspicuous plant in the Alps, growing under
much the same conditions as the Globe-flower and
belonging to the same family, is the white Aconite-
leaved Buttercup (Ranunculus aconitifolius, Linn.)
(Frontispiece and Plate XL.). With our neigh-
bours, the double-flowered variety of this plant is
known as the "Fair Maid of France."

In Britain we have a group of white-flowered
Buttercups of aquatic habit known as the Water
Ranunculi, which flourish in ponds, wet ditches, and
gently flowing streams. These plants are specially
adapted to their habitat, the submerged leaves
being highly divided and quite different in form from

210 MARSH PLANTS

those which float or project above the surface of the
water. These Water Ranunculi are entirely absent
from the Swiss Alpine region. Their place is taken by
such species as the Aconite-leaved Buttercup, which is
extremely partial to damp, but not too wet, soils. In
Plate XL. a photograph of an Alp covered with this
plant is seen. But it will be noticed that it does not
grow everywhere. It is distributed in bands, and
these bands represent the portions of the pasture
where the soil is dampest. The higher and drier
areas are unoccupied. Thus we have here a veritable
hygrometric chart of an Alp, the lines of highest
water contents being indicated by this plant. Similar
zones may also be frequently observed in Alpine
meadows, traversed by one or more damp ditches or
depressions, which are, as a rule, marked out as bands
of pure white by this Buttercup.

Ranunculus aconitifolius is a large, highly
branched, spreading plant, the leaves resembling
those of the Winter Aconite (Eranthis). A variety
of this plant, very similar in many respects, but with a
leaf like that of a Plane (Platanus), is by some
regarded as a distinct species (R. platantfolius, Linn.).
It, on the contrary, occurs chiefly in fairly dry
situations in the Alps.

THE BUTTERWORTS.

The Butterworts, members of the genus
Pinguicula, belong to an order of insectivorous
plants, the Lentibulariacese, which stands near to the

PLATE XL.

THE BUTTERWORTS 211

large order Scrophulariacese. There are three species
of British Butterworts, and in Britain we have also
other insectivorous representatives of the family in
the genus Utricularia, or Bladderworts. The latter
also occur in Lowland Switzerland, but are rare within
the Alpine zone, though one species has been observed
in boggy pools on the Julier road, above Silvaplana,
at 6,400 feet.

The Swiss Alpine Butterworts are easily distin-
guished. The Alpine Butterwort (Pinguicula alpina,
Linn.) has large white flowers, while the Common
Butterwort (Pinguicula vulgaris, Linn.) and the Large-
flowered Butterwort (Pinguicula grandiflora, Lam.)
have blue flowers. All the species are essentially
marsh plants, though they will flourish in many other
situations in which the soil is usually very damp.

We will commence with the Alpine Butterwort.
This is a British plant, though with us it is very rare,
and confined to a few districts on the west coast of
Scotland. In Switzerland, however, it is common
and widely distributed.

The build of the plant is characteristic. There is
a little rosette of light green leaves, close to the
ground, of which we shall have more to say presently.
From the rosette springs a single, long flower-stalk,
terminating in a solitary white, occasionally yellowish,
or even slightly purple flower (Plate XLI., Fig. 1).
The flower is built much on the same plan as that of
the members of the Scrophulariacese. Both the sepals
and the petals are united so that each forms a two-

212 MARSH PLANTS

lipped structure. The corolla has also a short spur,
and often two yellow spots on the lower lip. But
there are only two stamens, and the ovary has only a
single compartment, in which the seeds are arranged
in much the same manner as in the Primroses and
Androsaces.

But we are not concerned here so much with the
flower as with the leaves of this very interesting plant.
If we examine the pale, yellowish-green leaves of the
rosette, we shall find that they are sticky on the upper
surface. This stickiness results from the presence of
numerous glands, or secretory organs, arranged on the
upper surface of the leaf. These glands secrete a
viscid fluid, which plays an important part in the
economy of the plant, as we shall see. By their
means it obtains part of its food-supply. Many of
the leaves will be found to be concave, for the edges
curve somewhat inwards and upwards towards the
centre of the leaf.

A cross-section of a leaf is shown in Text-fig.
XIX., 1. On the upper surface, two kinds of glands
will be seen, some which are stalked and others which
are stalkless, the latter being the more numerous. In
Text-fig. XIX., 3, a surface view of one of the stalk-
less glands is seen, while Text-fig. XIX., 2, shows a
side view of the same. Both of these are highly
magnified under the microscope. The glands them-
selves can, however, be made out by examining the
upper surface of a leaf with a powerful hand-lens.

Kerner states that there may be as many as 25,000

PLATE XLI.

THE BUTTERWORTS

213

of these glands on a square centimetre of the leaf,
and consequently a plant with a rosette of six to nine
leaves is estimated to possess about half a million.

In the photograph on Plate XLL, Fig. 2, two rosettes
of Pinguicula leaves are seen, thriving on a cushion of
damp moss, which is itself attached to the root of a
Spruce Fir. It will be noticed that numerous remains
of dead insects, chiefly flies, occur on the leaves of the

FIG. XIX. — The Common Butterwort (Pinguicula vulgaris, Linn.).

1. Transverse section of a leaf, showing the glands on the upper surface.
Somewhat enlarged. 2. A side view of a gland. Much enlarged.

3. A surface view of a gland. Much enlarged.

larger rosettes, and in one case, on the uppermost
leaf of the smaller rosette, growing below and slightly
to the right of the larger rosette, the remains of a
moth are clearly seen.

This plant has the power, not only of catching
small insects on its leaves, but of digesting and absorb-
ing them. What happens is briefly as follows. The
glandular hairs of the upper surface secrete a sticky
fluid, which attracts insects, probably under the

214 MARSH PLANTS

impression that honey is to be obtained there gratis.
If the insect is a small one, it becomes firmly glued to
the surface of the leaf by mucilage secreted by the
glands. Somewhat later, another substance, a
digestive fluid, known scientifically as a ferment, is
also secreted by the part of the leaf with which the
insect is in contact. The ferment has the power of
reducing to a liquid state and digesting the insect, all
except the indestructible chitinous investment of the
body. Finally, the products are absorbed by the leaf
itself, and go towards its food-supply. The chitinous
investment of the insect remains attached until the
mucilage disappears, and it is then blown away by
the wind, or washed away by rain, and the trap for
fresh insects is set again.

Such is, briefly, one of the most interesting of
phenomena among the Higher Plants. The insecti-
vorous habit, though not rare, is infrequent in the
vegetable kingdom. Examples occur in the case of
our British Sundews (Drosera) and Bladderworts
(Utricufaria), some of which are also found in Lowland
Switzerland. Extreme or highly evolved adaptations
for a similar purpose are to be seen in the curious
tropical Pitcher plants (Nepenthes), commonly
cultivated in our greenhouses. Those who may be
interested to pursue this subject further should read
Charles Darwin's "Insectivorous Plants," which
contains a whole chapter devoted to the Butterworts.

In this connection, it may be worth while to call
attention to some further points, discussed by Darwin,

THE BUTTERWORTS 215

in the case of English specimens of the Common
Butterwort, Pinguicula vulgaris, Linn. This species
also occurs in Alpine Switzerland (Plate XXXVIII.,
Fig. 2). In one case Darwin records that 142 insects
were caught by 32 leaves, giving an average of 4*4
insects per leaf. In addition, the small leaves of a
Heath and other plant fragments were often blown
by the wind on to the leaf accidentally, to which they
adhered. He was also able to show that when
objects which contain little or no soluble matter come
to rest on the leaves, there is no secretion by the
glands. But where the substance is nitrogenous, the
secretion is copious, and the material absorbed by
the leaf contributes to the food-supply of the plant,
and helps to compensate for the limited extent of the
root system by which nutriment is derived from the
soil.

The fact that the leaves of the Butterworts contain
a special substance was known for some hundred
years before Science discovered its true nature.
The ancient herbalist, John Gerarde, writing in 1597,
says, " The husbandmen's wives of Yorkshire, do use
to annoint the dugs of their kine with the fat and
oilous juice of the herbe Butterwoort, when they
are bitten with any venemous worm, or chapped,
risted, and hurt by any other meanes." Kerner also
states that a similar use of this plant is made in
Switzerland. Linnaeus, more than 150 years ago,
related that the Common Pinguicula, which is
frequent in the Arctic regions as well as in the

216 MARSH PLANTS

Alps, is used by the Laplanders to curdle milk.
This can be easily confirmed by the experiment of
placing a few pieces of the leaves in a little new milk,
and letting it stand for some hours.

The other Alpine species of Butterwort, the
Common Butterwort (Pinguicula vulgaris, Linn.), and
the rarer Large-flowered Butterwort (Pinguicula
grandiflora, Lam.), the latter not occurring in Britain,
are biologically similar to the Alpine Butterwort, with
which they are often associated in marshy places and
on wet soils, such as a crevice in a rock filled with
damp moss. They both have blue flowers and longer
spurs than the Alpine Butterwort.

THE GRASS-OF-PARNASSUS.

TheGrass-of-Parnassus (Parnassiapalustris, Linn.,
natural order Saxifragacese, the Saxifrage family) is
another Lowland marsh plant, common in Britain,
which ascends to the Alpine zone in Switzerland,
where it is abundant. Like Caltha and many marsh-
loving species, the whole plant is smooth and devoid
of hairs. The leaves, which spring from a very
short perennial stem, have fairly long stalks, and are
heart shaped. Each year the stem sends up an erect
flowering stem, which bears a single leaf, and ends in
a large white flower, the structure of which is very
interesting.

There are five small, green sepals, and five white
petals, the latter with conspicuous translucent veins,
which add considerably to the attractiveness of the

THE GRASS-OF-PARNASSUS

217

flower. More internally we find five normal fertile
stamens, and then five barren stamens, greatly
modified in form. The latter are branched structures,
reminding one of a group of small pins arranged in a
cushion in a fan-like manner (Text-fig. XX.). The
ovary is seen in the centre of the flower, and divides
above into four stigmas. The fertile stamens, when
young, are bent inwards over the ovary. They shed
their pollen one at a time, and
then bend backwards and out-
wards, one by one, the filament
or stalk also elongating con-
siderably.

There has been much
speculation as to the use of
the modified stamens to the
plant. The flower is cross-
fertilised by flies. The anthers
shed their pollen before the
stigmas are ripe, and thus
self- fertilisation is impossible.

Honey is secreted at the base, and on the inner
side of the modified stamens. But at first sight
the little yellow knobs, glistening in the sun at the
tops of the pin-like lobes of the branches of these
organs, appear quite like drops of honey. These are
really false nectaries, and not true honey-glands.
Flies are constantly being deceived by them, and
have been watched licking the false glands under the
impression that honey would be forthcoming.

FIG. XX. — A Staminode from
a Flower of the Grass-of-
Parnassus (Parnassia palus-
trisy Linn.). Much magni-
fied.

218 MARSH PLANTS

The precise function of these false nectaries cannot,
however, be said to be fully understood at present.
They may be additional allurements to flies — i.e., insect
advertisements — or they may constitute a sort of fence,
which forces the insect to enter the flower, and reach
the concealed honey in a particular way, which is
favourable to cross-pollination.

THE ALPINE LOUSEWORTS.

The Louseworts, genus Pedicularis (natural order
Scrophulariacese, the Foxglove family), are quite
characteristic plants in marshy places in the Alpine
zone. More than ten species occur, as opposed to two
in Britain, so the genus is a very successful one in
Alpine Switzerland. They are quite similar in build
to our British Louseworts, except Pedicularis verticil-
lata, Linn., in which the leaves are whorled. The
flowers, as a rule, are, however, rather larger than
with us, and are either yellowish-red or reddish -black,
according to the species.

The interesting point to notice about the Louse-
worts is that they are semiparasites. This peculiarity
they share with several other genera, nearly related
and belonging to the same section of the family, such
as Euphrasia, Eyebright ; Melampyrum, Cow-wheat ;
Rhinanthus, Battle; and Bartsia — all common in
Alpine Switzerland.

In temperate floras parasitic Flowering Plants are
rare, though they are common in the tropics. They
are plants which attach themselves to other living

PARASITES AND SEMIPARASITES 219

plants — the hosts, as they are called — and draw from
them the whole or part of their nourishment. The
Dodder is a familiar instance of a plant wholly
parasitic on Thyme and a variety of other hosts.
The Mistletoe is a semiparasite. It attaches itself
to the branches of the Apple and some other
trees, and while it draws part of its nutriment
from the host, yet, by means of its leaves, it manu-
factures a portion, at least, of its food-supply itself.
In the Alps parasites of either type are rare, with
the exception of the members of the order Scrophu-
lariacese mentioned above.

Euphrasia and the other related genera are semi-
parasitic on the roots of Grasses, which they rob of
part of their food-supply. If we dig up a Lousewort,
and separate the plant from the turf very carefully,
we shall find that its roots are often attached to those
of certain Grasses. At the same time, these plants, by
means of their leaves, manufacture a portion of their
nutriment.

CHAPTER IX

THE ALPINE THICKETS AND FOEESTS

THE shrubby plants, forming the Alpine thickets and
the fringe of the Pine forests, are very interesting,
and well repay attention.

The Alpenroses and the Alpine Rose, which are
commonly found growing in such positions, have
been already discussed in the first chapter. The
former are perhaps the most characteristic plants of
the Alpine thickets, and frequently form regular
terraces on the hillsides. With them are often
associated many other plants, especially the Bilberries
and Honeysuckles, which we will discuss in the
present chapter.

It is rather remarkable that the fruits of nearly all
the shrubby plants of the Alpine thickets, with the
exception of the Alpenroses and a few others, are
berries or other succulent fruits, for bird-life is
remarkably scarce in the Alpine world. In the
Lowlands such fruits are greedily devoured by birds,
which distribute the seeds to some distance from the
parent plant, the fleshy pulp of the berry being an

THE BILBERRIES 221

adaptation to attract birds. In the Alps, however,
while a certain number of berries are no doubt eaten
by Black-Cock, Ptarmigan, Grouse, and other birds,
such as the Snow-Finch, the Bing-Ouzle, and the
Alpine Chough, the number of berries produced
annually seems to be greater than the demands of
bird-life require.

THE BILBEKRIES.

Under this name we may group together the three
Alpine species of Vaccinium, all of which are common
British plants. This genus is a member of the Heath
family (natural order Ericacese), though by some it is
placed in a separate order, Vacciniacese. Vaccinium
myrtillus, Linn., is the True Bilberry ; V. uliginosum,
Linn., is the Bog Vaccinium ; and V. vitis-idcea, Linn.,
the Cowberry or Ked Whortleberry (Plate XLII.,
Fig. 1). All three are often associated in the Alps.

Vaccinium myrtillus, Linn.

The True Bilberry (V. myrtillus) is distinguished
from the other two species by the fact that the leaves
are toothed, and the stem is triangular in section.
The leaves are thin, and are shed each autumn. The
berry is blue-black in colour.

The flowers are by no means conspicuous, and
quite scentless. The corollas are pale, greenish-white
in colour, globular in form, and nearly as broad as
long.

The plant is a low, thick shrub, with long under-

222 THE ALPINE THICKETS AND FORESTS

ground runners, by whose aid it manages to cover
large areas.

V actinium uliginosum, Linn.

The Bog Vaccinium has a round stem. The leaves
are not toothed, but, like those of the True Bilberry,
they are thin, and shed in the autumn. When young
they have a bluish-green tinge, and are much veined.
The berry is also blue-black in colour.

The bush of this species is taller and more upright
than that of the other Bilberries. Otherwise the
plants are very similar.

Vaccinium vitis-idcea, Linn.

The Cowberry (Plate XLIL, Fig. 1) is easily
distinguished from the other species by the red
berry and the thick evergreen leaves, which are
not toothed at the edges. The shrub has also
numerous runners below the surface of the soil,
from which fresh shoots spring.

The leaves are rolled at their edges, the under side
being studded with glands, which appear to the naked
eye as brown dots. The corolla is also shaped
quite differently from that of either of the two other
Bilberries.

Like the other Bilberries, Vaccinium vitis-idcea
has its enemies. It is largely attacked by parasitic
Fungi, which grow both in the leaves and in the berry,
greatly to the detriment of the plant.

PLATE XLII.

FIG. 1. — The Cowberry (V actinium vitis-i<l&a, Linn.).

FIGS. 2, 3. — The Flowers of the Alpine Soldanella (Soldanella alpina, Linn.).

FIG. 4. — Two Flowers of the Blue Honeysuckle (Lonict,ra ccvridea, Linn.).

[To Jace p. 222.

STAMENS OF THE BILBERRIES

223

The stamens of the Bilberries, especially the True
Bilberry and the Bog Vaccinium, are interesting, and
should be examined with a pocket-lens (Text-fig.
XXI.). Each stamen consists of a flattened stalk,
bearing above and on its inner side two flagon-shaped
structures, placed side by side. Each flagon is really
a half-anther, and it opens at the top by a pore,
through which the pollen escapes. From the stalk,
continued up the back of the
flagon, a little horn-like process
projects from each half-anther.
When a bee visits a flower in
search of the nectar at the base of
the stamens, it touches these horns
with its proboscis, and thus shakes
the anther, and is dusted with pollen
thrown out through the pores at the
top of the stamen. This pollen may
be carried to another flower, in
which the stigmas are ripe, and
thus cross-fertilisation is effected.

In the Ked Whortleberry the half-anthers are not
horned, but the tips of the flagons are produced into
long spout-like structures, with a pore opening at the
top of each. In this species the mouth of the corolla
is not contracted — an adaptation to ersure that the
insect strikes the horns of the stamens — but is widely
open, and the spout-like tips of the stamens no doubt
play a similar part to the horns, in conjunction with
the contracted throat of the corolla found in

FIG. XXI.— Stamen of
a Bilberry ( Vaccinium).

p, pores at the apex of
the anthers, through
which the pollen is
shed; a, anther; /,
filament.

224 THE ALPINE THICKETS AND FORESTS

Vacdnium myrtillus and V. uliginosum. Horned
anthers, though somewhat different in shape, are also
found in Arctostaphylos alpina, another shrub, which
we shall mention presently, belonging to the same
family, Ericaceae.

THE ALPINE HONEYSUCKLES.

The Alpine Honeysuckles are exceedingly in-
teresting plants. Unlike our English Honeysuckle,
or Woodbine (Lonicera periclymenum, Linn.) (natural
order Caprifoliaceae, the Honeysuckle family) — the
only species which is regarded as a true native
of Britain, though two others commonly occur
naturalised — the Honeysuckles of Alpine Switzerland
are not climbing plants, but erect bushes — often, in
fact, large shrubs. They are frequent in the dwarf
thickets and on the margins of forests, and often
extend considerably higher than the tree limit. They
are commonly associated with the Alpenroses and
Bilberries.

There are three species in the Alpine zone, which
differ greatly as regards their flowers. These are
not nearly so conspicuous as in our British Woodbine,
where they are bunched together in heads, and thus
gain in conspicuousness by massing. In the Alpine
species the flowers are borne in pairs, arising in
the axils of the leaves, by which they are partly
hidden.

THE BLACK HONEYSUCKLE 225

Lonicera nigra, Linn.

The Black Honeysuckle (Lonicera nigra, Linn.),
so called from its black berry, bears two flowers, placed
side by side on the end of a very long common
flower-stalk, usually more than three or four times
the length of the flowers. As in the other Alpine
Honeysuckles, the flower arrangement is really in
threes, one flower being central, and the other two
lateral, on either side of the central flower. But in
these plants only the lateral flowers are present, the
central one being entirely suppressed. This type of
flower arrangement, which may be well studied in
the Eagged Robin or the Eock Catchfly (Plate XVI.,
Fig. 2), is termed by the botanist a simple cyme or
dichasium.

In the typical cyme, each flower is subtended by
a leaf or bract. In the Alpine Honeysuckles this
bract may or may not be present, but two small leaf-
lets occur, which are known as bracteoles. Thus the
inflorescence here consists of the two lateral flowers
of a cyme, each with a pair of bracteoles at the base.

The interesting point in regard to the flowers and
fruits of the Black Honeysuckle, in comparison with
the other species described below, is that the two
ovaries of the flowers, and later the two berries, are
slightly united for less than half their length at the
base. The four bracteoles of the flowers are also
united in pairs, and in the flowering stage sheathe the
lower portions of the ovaries. The two berries, which

226 THE ALPINE THICKETS AND FORESTS

are quite distinct, although partly united in one plane,
are longer than the bracteoles.

A similar state of affairs is found in the Lowland
plant (Lonicera xylosteum, Linn.), which sometimes
occurs in Britain.

Lonicera alpigena.
Let us now compare the flowers of the Mountain

FIG. XXII. — Two Flowers of the Mountain Honeysuckle (Lonicera alpigena,
Linn.), with Ovaries partially united. (After Hermann Mtiller.)

Honeysuckle (Lonicera alpigena, Linn.) (Text-
fig. XXII.) The two flowers are here also mounted
on long flower-stalks, but only a single berry results,
which is red. If the ovaries of the flowers are
examined, it will be found that they are completely
united in the median plane. The result is a double

THE MOUNTAIN HONEYSUCKLE

227

berry, formed by the fusion of the two ovaries ; and
as the fruit ripens and becomes globose, the distinction
between the two ovaries, from which it originates, is
gradually lost. The double berry, however, bears
two scars near the apex, marking the position of the
calyx of each flower (Text-fig. XXIII). The
bracteoles in this species are
small and unimportant.

Curiously enough, while the
formation of a double berry is
constant wherever this plant is
found in Switzerland, in India,
where it also occurs, the berries
are always free and not united.

The fruit of the Mountain
Honeysuckle forms an ex-
cellent illustration of the
botanical axiom, that the ex-
planation of many features
presented by flowers is to
be sought for in the fruit. The flower is merely a
stage towards the fruit, and the fruit is only a con-
trivance for the distribution of one or more seeds at a
distance from the parent plant. In the case of the
Mountain Honeysuckle, the idea appears to be, that
if the seeds of two flowers are contained within a
single berry, they will have a greater chance of being
all distributed, should some bird devour the berry,
than if two berries containing the same number of
seeds were produced.

FIG. XXIII.— The Fruit of the
Mountain Honeysuckle
(Lonicera al<pigenay Linn.),
formed by the complete union
of the two berries.

The scars of the two calyx rings
can still be seen.

228 THE ALPINE THICKETS AND FORESTS

Lonicera ccerulea, Linn.

In the flowers of the third Alpine species, the Blue
Honeysuckle (Lonicera ccerulea, Linn.), a very curious
state of affairs is found, quite unlike that in L. nigra
and L. alpigena. The leaves of this shrub are also
very different, being delicate in texture, to some
extent transparent, and of a bluish-green colour,
especially beneath. The berry is black, with a bluish
bloom. The flower-stalks are short.

If we examine the flowers, we shall find that the
two ovaries appear to be entirely united (Plate XLII.,
Fig. 4). As a matter of fact, they are quite free from
one another, and what we see externally is a sheath
formed by the union of the bracteoles (p. 225), which,
when the ovaries are mature, entirely enclose them,
and are partly united to them.

As the fruit ripens, the growth of the bracteolar
sheath keeps pace with the growth of the ovaries
containing the seeds, and forms the fleshy substance
and the skin of the single " false berry " which is not
very dissimilar in appearance to that of the Mountain
Honeysuckle. Here, however, it is the bracteolar
sheath and not the ovary walls, which form the outer
substance of the berry, and this assumes eventually a
bluish-black hue. In the peculiar origin of the " false
berry," the Blue Honeysuckle stands quite alone
among the members of the genus.

A very striking feature of the Alpine flora is its
extreme poverty in climbing plants, whereas our

THE ALPINE CLEMATIS 229

British flora contains quite a fair percentage of species
which climb by some means or other, while in the
forests of the tropics the number is very much greater.
We have already noticed that the climbing Honey-
suckles do not occur in the Alps, In the Alpine zone
there are also no Convolvuli, no Ivies, and no Traveller's
Joy. Neither are any Vetches nor Peas, with
modified climbing leaves or tendrils, indigenous to
this zone. Of the hook-climbers — plants which
scramble up over other plants by means of recurved
hooks — certain species of Bramble (Rubus) and
Bedstraw (Galium) are frequent in Alpine Switzerland,
as in Britain. There remains only one other climbing
plant, the Atragene or Alpine Clematis.

THE ATRAGENE OR ALPINE CLEMATIS.

The Alpine Clematis or Atragene (Clematis alpina,
Miller, also known as Atragene alpina, Linn., natural
order Kanunculaceae, the Buttercup family) (Plate
XLIII.) is a near relative of our British Traveller's
Joy (Clematis vitalba, Linn.), which occurs also in
Lowland Switzerland and ascends to nearly 3,000
feet. It is a woody plant, not infrequent in the
Alpine thickets, climbing up over other plants,
though not, perhaps, very common in the lower
portion of the Alpine zone.

This genus is remarkable among the Buttercup
family as being the only one which has opposite leaves.
The Alpine Clematis climbs, not by means of tendrils,
but with the aid of its long, sensitive leaf- stalks^ which

230 THE ALPINE THICKETS AND FORESTS

twine round any support within their reach. The leaf-
stalks are so sensitive, that mere contact with the
support is sufficient to stimulate them to twine
around it.

The flowers (Plate XLIIL, Fig. 1) are large and
handsome, and mounted singly on long stalks.
Externally there are four long, lance-shaped sepals,
deep blue in colour. These enclose four spade-
shaped petals, much smaller in size, and whitish in
colour. There are many stamens and carpels, as in
most of the other members of the family.

The fruits (Plate XLIIL, Fig. 2), enclosing each
a single seed, have long feathery awns, quite com-
parable to those we have already described in the
case of certain Anemones, the White Dryas, and the
Avens. They are distributed by wind.

THE BEARBERRIES.

The Red Bearberry (Arctostaphylos uva-ursi,
Sprengel) and the Alpine Bearberry (A. alpina,
Sprengel (natural order Ericaceae, the Heath family),
both British plants, are common in Switzerland, on
the hillsides, and in the neighbourhood of the Alpine
thickets. They are low carpet plants, highly branched,
each branch covered with leaves, which in the Eed
Bearberry are thick, leathery, entire, and evergreen,
while in the Alpine Bearberry they are thin, toothed,
and shed at the end of summer. A hillside covered
with the Alpine Bearberry in autumn furnishes one
of the most wonderful sights to be seen in the Alps,

PLATE XLIII.

FIG. 1.— The Flowers.

FIG. 2 — The Fruits.
The Atragene (Clematis alpina, Miller).

[To face p. 230.

THE CROWBERRY 231

on account of the intense, ruby-red colour, which
the leaves assume just before they are shed.

The flowers are fashioned quite like those of the
Alpine Heath (Erica earned). The stamens are horned,
and the same mechanism to ensure cross-fertilisation
exists as in the Vacciniums described on p. 223. The
fruits are berry-like, but botanically of the type
termed drupes, with one to five stones, as in the
Crowberry, next to be described. Those of the Eed
Bearberry am red, while the Alpine Bearberry has
black fruits.

THE CROWBERRY.

The Crowberry (Empetrum nigrum, Linn., natural
order Empetracese, the Crowberry family) is a low,
spreading, heath-like shrub, rarely more than a foot
in height, common in the dwarf thickets of the High
Alpine regions, and often associated with the Trailing
Azalea. There is only one species of this genus,
which, however, is very widely distributed, occurring
in Britain and Northern Europe, and even in the
Andes of South America. The plants often reach
a considerable age in the Alps, although the stem
and the annual rings of growth are very small indeed.

The branches are closely set with leaves, which
are needle-like in shape, and evergreen. The general
form of the leaf resembles that of a Heath. The leaf
is rolled, so that the edges meet below. The margins
are interlocked with hairs, and thus enclose an oval
cavity, lined by the lower surface of the leaf on which

232 THE ALPINE THICKETS AND FORESTS

the stomata and also glandular hairs are situated.
This is another adaptation, similar to those to which
attention has already been drawn, for reducing the
risk of excessive loss of moisture from the leaf by
means of the stomata, which are here protected by
their position in the half-closed chamber formed by
the rolling in of the leaf.

The flowers, which are very small and stalk-
less, are often produced in great numbers near the
tips of the twigs. As a rule, each plant is either
male or female — that is, all the flowers on one plant
belong to one sex. The pollen of the male flowers
is carried to the stigmas of the female by the agency
of the wind. In some cases, though rather rarely,
the flowers are hermaphrodite — i.e., they contain both
male and female organs in the same flower.

The fruit is like a berry externally, but, in its
structure, it more closely resembles that of a Cherry
or a Peach, in that the inner portion is hard or
stony. This type of fruit is distinguished botanically
as a drupe. In the drupe of the Crowberry, there are
from six to nine stones. The fruits are not eaten
in Switzerland, but, in Scandinavia, the Laplanders
and Finns make use of them. In Northern Europe
these fruits grow naturally much larger and are
more juicy than in the Alps, as is also the case with
the fruits of several other Alpines which occur in
Scandinavia.

THE JUNIPER AND ALDER 233

THE DWARF JUNIPER.

The Dwarf Juniper (Juniperus communis, Linn.,
var. nana, Willd., class Coniferse, natural order
Cupressaceae, the Cypress family), which is very
frequent in the Alpine and High Alpine zones, is
regarded as simply a variety of the Common Juniper.
In habit it is a large carpet plant, the branches,
densely clothed with leaves, being pressed close to
the ground, so that the height of the plant is quite
small. This form of habit is no doubt well calculated
to withstand the great weight of the winter snow,
which lies over the shrub for several months each
year.

It is often abundant in or near Alpine thickets,
and also grows commonly on rocks. In some respects
it is an important coloniser of bare rocky places, like
the plants discussed in Chapter IV. It is probably
the highest woody plant occurring in Switzerland,
having been recorded on Monte Eosa at an altitude
of 11,700 feet. The fruit is a berry, and here again
we have another example of an Alpine shrub with
a succulent fruit.

THE GREEN ALDER.

The Green Alder (Alnus viridis, D. C., natural
order Betulaceae, the Birch family) is a common
shrub in the thickets, bordering the mountain
streams. It is interesting as being one of the few
representatives in the Alpine zone of the tree

234 THE ALPINE THICKETS AND FORESTS

families, with deciduous leaves and catkin-like
inflorescences, so characteristic of the Lowlands of
Europe. Dwarf Willows, p. 187, also occur, especially
in the High Alps.

The Green Alder has no very striking peculiarities.
It resembles the Alders of the plains, except that it is
rather dwarfed in stature.

THE CONIFEROUS FORESTS.

The Coniferous forests are a highly characteristic
feature of the lower portion of the Alpine zone.
They consist essentially of two trees : the Spruce
and the Larch, the former, as a rule, prevailing.
Thus, although it is customary to speak of the Pine
forests of the Alpine region, this term is not strictly
accurate in a botanical sense, since neither the Spruce
nor the Larch are true Pines. They are indeed very
closely related to them — so closely, that whether
we call the forests Coniferous or Pine is a small
matter, if we bear in mind that the only true Pines
found in Alpine Switzerland, the Stone or Arolla
Pine, and the Mountain Pine, are not nowadays
forest formers in the Swiss Alps.

The race of plants, the Coniferse, to which the
Larch, Spruce, Pines, Firs, and Junipers belong,
is quite distinct from that to which all the other
Alpines discussed in this volume are assigned —
namely, the Flowering Plants, or Angiosperms.
These two races differ greatly, not only in their
general structure, and in their life histories; but it

THE CONIFEROUS FORESTS 235

is impossible, without entering on a lengthy and
technical botanical discussion, to indicate precisely
in what these differences consist. It may suffice to
say that the Conifers do not bear flowers. The
organs which produce the pollen and the ovules,
which when fertilised become seeds, are borne on
special, complicated fertile shoots known as cones.
These produce either pollen or ovules, but never
both, and are thus spoken of as male or female
cones. Both types may occur on the same or on
different trees, according to the genus. All the
Coniferse are wind fertilised.

At a height of about 5,000 feet (which we here
regard as the lower limit of the Alpine zone), the
Beech, so characteristic of the Subalpine region, is, as a
rule, entirely replaced by the Spruce and Larch.
Forests in which these trees figure largely are of
course common also in the Lowland and Subalpine
zones, but they are there associated with other trees,
not found in Alpine habitats.

The Coniferous forests extend upwards to a height
which depends on a great variety of circumstances.
Often the upward limit is as low as 6,000 feet,
but in the Zermatt region it is as high as 7,600 feet.
No general rule can be laid down, for the upper
frontier depends on various local conditions such as
the height of the floors of the neighbouring Alpine
valleys, the nearness or remoteness of the snow-line,
and the situation, or aspect of the particular district.
The best way to view the vertical distribution of

236 THE ALPINE THICKETS AND FORESTS

the Pine forests is in relation to the altitude of the
floors of the Alpine valleys on the one hand, and the
snow-line on the other. The upper limit of the
forests, which is as a rule very sharply defined,
bears a definite relation to both. In the case of
a wooded Alpine valley, the following sequence can
generally be determined : —

Immediately above the valley
floor and the Alpinemeadows
Snow. the forests rise, clothing the

valley -sides for 1,000 or
Pastures. 1,500 feet. The gap between

the upper frontiers of the

Forests. forest and the snow-line is

filled by the treeless

Alpine Meadows.
Valley floor. pastures, extending upwards

for some 2,000 to 2,600 feet.

It has already been explained (p. 170) that the height
of the snow-line varies in different parts of the Alps.
The upper limit of the forests, for the most part, varies
in height according to the position of the snow-line
in that particular district, and keeps at a respectful
distance of 2,000 feet or more from it.

Further, just as the mean level of the floors of the
Alpine valleys varies in elevation, so the vertical dis-
tribution of the Larches and Spruces which clothe their
sides changes. Probably exposure to wind is one of
the great factors that determines the upward limit of
the forests. So long as they are in some measure pro-
tected by the "brow of the hill," the trees can flourish.

THE CONIFEROUS FORESTS 237

Of course, many desolate, treeless valleys are to
be found in the Alps. Here the forests have either
been entirely destroyed by human agency, or there
may be some other special reason, such as a
particularly sunless aspect or configuration, or the
fact that in winter the valley is the track of frequent
avalanches, or that the soil is barren or absent.

Forests were at one time very much more
extensive in Switzerland, as a whole, than they are
to-day. How greatly they have been destroyed can
be appreciated by a study of some of the older Swiss
maps. Many localities, which are there indicated as
thickly forested, are almost treeless to-day, or perhaps
only a group of Spruces or Larches survives. The
same is also the case in England, which in the Middle
Ages was a highly wooded country.

In 1862, according to official estimates, quoted by
Dr Christ, the forest area comprised only 15*4 per
cent, of the whole of Switzerland. It was estimated
that the country consisted of: —

Barren land . '».:• + . 31*6 per cent.

(including snowflelds,
glaciers, etc.)

Pastures .... 33'0 „
Arable land . . . "' . ' 20*0 „

Forest . 15*4

lOO'O

The percentage of forested land in Switzerland was
then less than half that met with in the Jura. No
doubt at the present day the percentage is still
smaller.

238 THE ALPINE THICKETS AND FORESTS

The great enemy of the forest has been man.
Before the introduction of State legislation, and the
institution of forestry police in 1876, the Switzer had
for centuries cut down the trees ruthlessly, sometimes
even wholesale, without any thought for the morrow.
The wood was chiefly required for fuel, though it was
alsip much made use of in building. Sometimes a
commune sold a whole forest for a tithe of its value
to speculators, who promptly cleared the ground.
In other cases, wholesale destruction was undertaken
in order to increase the acreage of the pastures or
meadows.

Nowadays, happily, a better condition of affairs
prevails. The forests are protected by stringent laws
and regulations with regard to cutting, thinning, and
replanting. In many situations, exposed to avalanches
in winter, all interference with the forests is forbidden.
But the damage has been done.

Great as has been the destruction wrought by man,
he is not responsible for the whole. Every year,
especially in winter-time, hundreds, perhaps thousands,
of noble trees perish beneath avalanches, Alpine
storms, or lightning. In addition, the goats and
cattle are a constant menace to the seedling plants,
the former being particularly partial to their young
shoots. Again, these trees have other enemies. They
are liable to be attacked by certain specific diseases,
either of insect origin, or caused by parasitic fungi,
such as the Larch disease, which in recent years has
destroyed many a fine giant in Switzerland.

PLATE XLIV.

FIG. 1. — Rock Colonisation : A Bare Slab of Rock in a Forest showing a
Primitive Soil composed of Pine-needles.

FIG. 2. — The Lichen ( Usnea) growing on the Branches of a Coniferous Tree.

[To face p. 238.

THE CONIFEROUS FORESTS 239

On the other hand, the Lichen (Usnea barbata)
(Plate XLIV., Fig. 2), known as the Old Man's
Beard Lichen, so frequently seen depending from the
boughs, especially the dead branches of the Spruce
or Larch, to which it is attached, is not, despite
statements to the contrary, known to be directly
harmful to the tree.

It has been well said that "the Pine forests play a
most important part in the natural economy of the
Alps, and their preservation is a matter of vital
consequence to the future inhabitants." What follows
when a country has been deprived of its forests is well
seen in certain districts in India and Ceylon. The
nature of the climate has been changed, and has
become drier and hotter than formerly. A forested
country tends to produce a more humid atmosphere
than a treeless district, and, in Switzerland, the
atmosphere is probably drier to-day than it was a
hundred years ago. Again, to a greater degree than
is usually realised, the forests afford the only efficient
safeguard which prevents the valleys and lower
pastures being overwhelmed with avalanches in
winter and floods in summer. Often the former fail
to melt completely for several summers after their
fall, and the ruin of the Alpine meadows and pastures
is rendered still more complete by the confusion of
rocky debris which they bring down in their train.

We will now discuss the trees forming the Alpine
forests.

240 THE ALPINE THICKETS AND FORESTS

THE SPRUCE.

The Spruce, Spruce Fir, or Norway Spruce (Picea
excelsa, Link.) ( = Pinus picea, Dur. = Abies excelsa,
Poir.), like the Larch, is a familiar tree in Britain,
where it has been much planted and become natural-
ised, though not originally a native. Young Spruces
with us, as in Germany, are universally used as
Christmas trees. The Spruce is easily distinguished
from the Pines by the fact that the leaves are borne
singly on the branches and spirally arranged, and not
in groups of two or more on very short shoots. The
female cone terminates the shoot formed during
the previous year. It is first erect, but later becomes
pendulent, when the seeds, which may number over
300, drop out and are distributed by the wind, aided
by the wing-like expansion of the seed.

As the specific name "excelsa" implies, it is one
of the loftiest trees found in Europe. Some specimens
are said to reach 150 to 180 feet in height. In the
Alps the average height is considerably less, and the
Spruce is usually overtopped by the Larch. Immense
forests of Spruce occur also in Scandinavia and
Northern Russia.

The Spruce is an extremely useful tree. It yields
a resin from which turpentine is extracted, and it is
also largely used as timber in the construction of the
chalets so characteristic of the pastures. The deep,
reddish-brown colour of the old Alpine chalets is
caused by changes in the resinous substances of the

THE LARCH 241

wood, brought about by long exposure to the intense
light and heat of the Alpine summers. It may be of
interest to note in this connection that amber is simply
the fossil resin of a species of Pine, Pinus succinifera,
long since extinct. Specimens of the wood of this
tree are known, containing pieces of amber, or fossil
resin, in the resin-forming tissues, which are quite
similar to those of living Conifers.

THE LARCH.

The Larch, Larix europcea, D. C. ( = Abies larix,
Poir.), is easily distinguished from the Pines and the
Spruce and from all European evergreen Coniferse,
by the fact that the leaves are shed each autumn.
These, like those of the Pines, and unlike the Spruce,
are borne on short branches. They occur in clusters
of fifteen to thirty. They are long, fine, soft, and
needle-shaped. The cones formed in early spring are
small, erect, and brightly coloured. They ripen during
a single year, whereas those of the Pines require two
or more years before the seeds mature. The male
cones terminate short, leafless shoots, and the female,
short, leafy branches.

The Larch rarely forms pure forests in the Alps.
As a rule, it is mixed with the Spruce. The dis-
tribution of these trees is best estimated in winter-
time, when the light brown of the leafless Larches is
sharply contrasted with the dark green of the evergreen
Spruces. Pure woods of Larch do, however, occur

242 THE ALPINE THICKETS AND FORESTS

exceptionally in certain localities, as in the Zermatt
and Saas Valleys. At one time it was thought that
the Larch altogether avoided limestone soils, but it
is now known that this is not the case. The Larch,
like the Spruce and Pines, yields resin and turpentine,
and the bark is sometimes used in tanning.

THE STONE OR AROLLA PINE.

The Stone Pine, or Arolla Pine as it is sometimes
called in Switzerland, Pinus cembra, Linn. (Plate III.),
is a very handsome tree, easily distinguished from all
the other Alpine Conifers, by the fact that the leaves
are arranged in little bundles of five, on very short
shoots, and by the absence of any wing to the seed.
It is a tall tree, sometimes reaching a height of
70 feet.

There is reason to believe that extensive forests of
Stone Pines once existed in Switzerland, but most of
these have long ago perished beneath the woodman's
axe. At the present day the Stone Pine is distinctly
uncommon, and, as a rule, only isolated examples or
small groups are met with at infrequent intervals.
It is perhaps the rarest as well as the most handsome
of Alpine Conifers. Dr Christ states that forests of
these Pines, much gnarled and twisted, occur on the
slopes of the Kieine Scheidegg (on the Grindelwald
side) and elsewhere in the Bernese Oberland. Some
of the finest examples are found in the Valaisian
valleys on the south side of the Rhone valley, especi-

THE MOUNTAIN PINE 243

ally at Arolla and on the Kiffelalp. Some grand
trees of this species also occur on the Engstlen Alp
(Canton Berne). A fine forest of Arolla Pine mixed
with Larch is traversed by the path to Fuorcla Surlej
from Silvaplana in the Engadine.

The Stone Pine produces large cones. The seeds
are nearly as big as a hazel-nut and are edible.

THE MOUNTAIN PINE.

The Mountain Pine (Pinus montana, Mill), is
nearly related to the Scotch Fir (P. sylvestris) from
which it is usually to be distinguished by the dwarf
habit alone. The leaves are borne in pairs on short
shoots, and the seeds are small and winged, but the
cones are stalkless, or nearly so, whereas those of the
Scotch Fir are stalked.

In the Pyrenees, the Mountain Pine forms
extensive forests, but this is rarely, if ever, the case in
Switzerland, though considerable masses of this tree
may occur exceptionally.

In one respect the Mountain Pine is one of the
most extraordinary plants to be found in the Alps.
The habit varies from that of an erect tree, 30 feet
high, to that of a low straggling shrub, 7 feet or less
in height. The largest trees, with stout, erect, and
straight trunks and smaller lateral branches, are apt
to be mistaken for those of other species. Some of
the best examples of this type occur in Canton
Grisons, especially at Wolfgang (the pass traversed

244 THE ALPINE THICKETS AND FORESTS

by the railway between Klosters and Davos), near
Lenzer Heide, and at the Maloja Pass around the
Chateau Belvedere. In the Valais also, between
Almagell and Saas Fee, good examples of the erect
tree form can be seen.

As a rule, however, the specimens of this tree
met with are dwarfs or semi-dwarfs. Either the
lateral branches, at some little distance from the base
of the trunk, grow more vigorously than the main
stem, or these branches arise just above the level of
the soil, and are quite prostrate on the ground, only
the smaller leafy shoots being erect. Thus a bush
habit, in certain respects not unlike that of the
Alpenroses or the Bilberries, is attained. In addition,
all sorts of minor variations may be observed. Some-
times the bush is fairly symmetrical, at others it is
extremely unsymmetrical, the plant being one-sided
and growing more or less in a straight line along the
ground, or down a steeply sloping bank. Curiously
enough, the cones are almost as variable in shape
and symmetry as the stems.

The Mountain Pine is of quite common occurrence,
especially above the limit of the Spruce and Larch
forests. Usually the dwarf bushes grow singly or
in small clumps, though here and there they may
form little miniature forests, inviting comparison with
the Coniferous woods at lower elevations.

THE SILVER FIR 245

THE SCOTCH PINE.

The Scotch Pine or Fir (Pinus sylvestris, Linn.), so
familiar to us in Britain, is rarely found above 5,000
feet in the Alps, though it is common enough in the
Lowlands and the Subalpine zone. The same is true
of the Silver Fir (Abies pectinata, D. C.) ( = Pinus
picea, Linn.), which is a characteristic Subalpine tree,
only rarely occurring in the Alpine zone.

CHAPTER X

THE SHADE PLANTS OF THE ALPINE FORESTS

IN the present chapter we will discuss some of the
commoner and more interesting plants, chiefly
herbaceous, which thrive for the most part only in
the shade of the Alpine forests and thickets. The
physical conditions which prevail in such habitats are
markedly different from those of the open pastures.
In the first place, the intensity of the illumination is
much less. Here and there direct sunlight may
penetrate in fine weather, but as a rule only very
diffuse light prevails. Again, the soil is generally
rich in humus or vegetable debris, derived from the
thick carpet of discarded leaves of the Spruce or
Larch which covers the forest floor. Many plants
avoid soils rich in humus, while others are very
partial to them. Another difference is found in the
fact that the shade plants are, to some extent,
protected from the weather, especially the wind.
Their roots also probably receive less moisture than if
they grew on the open Alp.

Of all the characteristic conditions under which

246

SHADE PLANTS 247

such plants live, the diffuse nature of the illumination
has probably the most powerful influence, not only on
the external form, but on the internal structure of the
plant (see p. 202). The dependence of the green
plant on light, for the maintenance of assimilation
through the agency of the chlorophyll, or green
pigment of the leaves, etc., has been already
explained (p. 10).

This being so, we should naturally look to the
leaves of shade plants for some indication of the
conditions under which they grow. In many cases,
but not by any means all, it will be found that the
leaves of shade plants are larger, broader, and thinner,
and at the same time more intensely green than those
of plants thriving fully exposed to the sun. They are
often hairless or only slightly hairy, but no rule can
be laid down as to the presence, or absence, of hairs
on the leaves of shade plants. Some, while hairless
on the upper surface, are covered with a thick hairy
felt on the lower. The leaves of the Lily of the
Valley, the May Lily (Plate XLVIL, Fig. 2), and
the Two-flowered Violet (Plate XLVL, Fig. 2) are
quite typical of a shady habitat. We also find in
a number of cases, that where the shoots bear many
scattered leaves, the successive leaves are separated
by fairly long internodes, as in Solomon's Seal.

If we examine closely any large Spruce forest, the
influence of sunlight can be readily seen. Where the
trees are dense and their branches interlace over-
head, the carpet of " needle " leaves will be found to

248 SHADE PLANTS OF THE ALPINE FORESTS

be almost entirely bare of herbaceous undergrowth,
the absence of which seems only to intensify the
prevailing gloom, through which no direct sunlight
ever penetrates. But here and there the trees are more
scattered, or some monarch Larch or Spruce has fallen.
A fair amount of sunlight reaches such spots on sunny
days, and a more vigorous undergrowth will be
observed, though the vegetation is still distinctly
sparse. In the less dense portions of the forest,
where several trees have fallen, either from natural
causes or beneath the axe, the glade will be found
thronged with plants all vigorously competing for
the available sunlight.

A flowery glade in a Larch forest presents a
pleasing contrast to the condition of affairs met with
in the denser Spruce woods. However, the under-
growth of a typical Swiss forest is never really
luxuriant, nor can it compare with the dense vegeta-
tion beneath the giants of a tropical forest, where the
intensity of the illumination is relatively greater, and
the air more highly charged with moisture.

We will now discuss some characteristic species
of the forest shade.

THE LINN^A.

Probably there is no more dainty or delicate plant
to be found in the whole of the Alps than the slender
Linnsea of the Pine forests (Plate XLV.). Linncea
borealis, Gronov. (natural order Caprifoliacese, the
Honeysuckle family), is the name plant of the great

PLATE XLV.

THE LINNJ3A 249

Swedish naturalist, Linnaeus (1707-1778), the Father
of Botany. Linnaeus invented the system, now
universally adopted, of giving two Latin names to
each animal and plant, the one generic and the other
specific, in order to distinguish them from one
another. He also laid the foundations of the present
system of classification of both animals and plants,
and thus practically called the sciences of Botany and
Zoology into being.

The Linnaea, which is a common plant in
Scandinavia, was well known to Linnaeus, with
whom it was a particular favourite. He specially
selected this plant to bear his name, and although
the term Linncea borealis was first recorded by
Gronovius by the wish and consent of Linnaeus, it
was the latter who first pointed out its true affinities.
A spray of Linncea will be found engraved on nearly
all portraits of the great northern naturalist, and he
himself adopted it as his crest.

There is only a single species of this plant in
existence, but it is very widely distributed. It is
abundant in Scandinavia and the Arctic regions. It
is found throughout the whole Alpine chain of
Central Europe, and also in the mountains of Asia
and North America. It even occurs in a few localities
in England and Scotland, as one of our rarer British
plants.

The Linnaea is a little creeping or trailing shrubby
plant, a foot or more in length (Plate XLV.). It
frequently grows on flat-topped rocks, and is fond of

250 SHADE PLANTS OF THE ALPINE FORESTS

soils rich in humus. The leaves are rather small,
and broadly egg-shaped. They are arranged in
opposite pairs, and are evergreen. Here and there,
the stems send up erect flowering branches, each of
which has two or three pairs of leaves, and terminates
in a long flower-stalk branching into two at the
top. From each fork, a single, graceful, white or
pale pink flower droops. The sepals and petals of
the calyx and corolla are five in number, but
there are only four stamens. The flowers have a
faint vanilla-like scent, which is said to be more
powerful by night than by day. The Laplanders
make a decoction of the flowers, which they use as a
remedy for certain complaints, such as rheumatism.

In some cases the lower portion of the plant is
almost hairless, in others it is distinctly hairy. The
calyx and the flower-stalks, however, are nearly always
covered with sticky bristles, which can be readily
seen under a hand-lens. They probably serve as a
protection against unbidden guests in the shape of
crawling insects, which otherwise might creep up
from below, and rob the flower of its pollen. Such
insects would only very rarely visit another flower,
and thus could be of no use to the plant as cross-
pollinators. The drooping position of the flower is
an additional protection against the visits of all but
flying insects, which alone can be of service to the
plant.

The sticky hairs on the lower portion of the
flowers also help in the distribution of the fruits.

THE BOX-LEAVED POLYGALA 251

The ovary in the fruit becomes a nut, and remains
enclosed in the two little leaves on the flower-stalk,
just below the flower, which at this stage increase
greatly in size. They are covered with glandular
hairs, which catch in the furry coat of any passing
animal, and thus the fruits are distributed to a
distance from the parent plant.

THE BOX-LEAVED POLYGALA.

We are all familiar with our little Common Milk-
wort (Poly gala vulgaris, Linn., natural order Poly-
galaceae, the Milkwort family), which, with other
Alpine species, is frequent also in Switzerland. But
in the Alpine woods and forests we often find another
species, the Box-leaved Polygala (Polygala chamce-
buxus, Linn.) (Plate XL VI., Fig. 1), forming little
carpet-like patches on the rocks. The whole habit
or build of this plant is quite unlike that of the
Common Milkwort. It is a low creeping shrub, with
leathery evergreen leaves, like those of the Box
(Buxus), and quite unlike the typical shade-leaves of
most Alpine forest plants. There is a good reason
for this difference. The fact is that the Box-leaved
Polygala is an immigrant from the Mediterranean
flora, like the Alpine Heath (Erica carnea, Linn.) and
Biscutella Icevigata, Linn., among other southern
plants which have successfully invaded the Alps. It
has retained several of the characteristic features of
the subtropical flora of Southern Europe, such as, for
instance, the evergreen leaves and the shrubby habit.

252 SHADE PLANTS OF THE ALPINE FORESTS

The flowers are produced quite early in spring, in
April or May. They have even been recorded in
bloom on Christmas Eve at altitudes of over 5,000
feet.

The flowers bear a curious resemblance to those
of a member of the Pea family (Leguminosse), but as
a matter of fact, they are constructed quite differently.
This will be seen by anyone who takes the trouble to
compare a flower of the Kidney Vetch (Anthyllis
vulneraria, Linn.), or of the Bird's-foot Trefoil (Lotus
corniculatus, Linn.), usually to be obtained not far
away, with that of this Polygala. The structure of
the typical Leguminous flower will be found described
in Appendix II. Here the conspicuous organs are
formed entirely by the petals, the upper petal being
the "standard," the two side petals the "wings" and
the two lower petals uniting to form the "keel" The
calyx remains small and green.

In the Box-leaved Polygala, on the other hand,
two of the sepals at the sides of the flower are much
enlarged, and are bright yellow in colour, spreading
or bending outwards to form the wings. Hence in
this plant the showy portion of the yellow or rose-
coloured flowers, borne singly or in pairs in the axils
of the leaves, is formed partly by the calyx, and
partly by the corolla.

Now we turn to the petals of the Polygala. As
a rule, there are only three, two being absent
altogether or rudimentary. The lower petal, how-
ever, is very large, and is shaped quite like the boat-

PLATE XLVI.

FIG. 1. — The Box-leaved Polygala (Polygala chamwbuxusi Linn.).

FIG. 2.— The Two-flowered Violet (Viola biflora, Linn.).

[ To face p. 252.

THE TWO-FLOWERED VIOLET 253

shaped keel of the Leguminous flower, while the two
upper petals form a sort of standard. There are only
eight stamens, all united into a bent tube, split on
one side. In this, again, we find a curious resemblance
to the Pea family, where, however, there are ten
stamens. The anthers open by pores. The style
lies inside the tube of stamens, and is likewise bent
in conformity with the shape of the keel.

The explanation of the curious parallelism between
the shape of the flower of this Polygala and that of
a Leguminous plant is to be sought for in the mode of
pollination. The whole structure of the flower is
adapted to that end, and though it differs in some
details, the manner of fertilisation is quite similar
to that of a typical member of the Pea family (see
p. 328).

The Box-leaved Polygala has the distinction of
being the sole member of the genus, in which the
stem is woody, all the other species being herbaceous.

THE TWO-FLOWERED VIOLET.
One of the most delicate and graceful plants of
the Alpine forests is the little Yellow-flowered Violet
(Viola biflora, Linn,, natural order Violacese, the
Violet family) (Plate XLVL, Fig. 2). This plant
loves the damp, shady dells among the Pines, where
it forms characteristic clumps. The slender shoots
bear usually two large, kidney-shaped leaves, quite
typical examples of shade-leaves (p. 247). They are
thin and delicate structures, light-green below, but

254 SHADE PLANTS OF THE ALPINE FORESTS

much darker in colour above. The veins are very
prominent.

Despite the specific name, "bifiora," each shoot
usually bears only a single flower, and not two, in
the Alps. The flowers are the smallest of any Swiss
Violet. They are bright yellow in colour, streaked
with brown. The plant attracts flies, which abound
in the forests, and is cross-pollinated by their agency.
The mechanism for ensuring cross-fertilisation is
much the same as in the case of the Field Pansy,
described on p. 161.

The flowers of this Violet probably correspond
more closely to the ancestral form, than those of any
other Swiss member of the genus. In the Field
Pansy (Viola tricolor, Linn.), the colour has become
wholly or partly changed from yellow to blue, and
the size of the flower has been greatly increased in
order to make them acceptable to humble-bees and
butterflies. In the case of the Long-spurred Violet
(Viola calcarata, Linn.) (p. 128), the flowers have
specialised for butterflies exclusively. To this end,
the colour has become "fixed" to a pale blue, the
size of the flower still further enlarged, and the spur
enormously lengthened.

THE MAY LILY.

One particular section (Convallariese) of the Lily
family, Liliacese, consists almost entirely of shade-
plants, which differ in several important respects from
other members of the order. The underground stems

THE MAY LILY 255

are of the type known botanically as rhizomes, and
not bulbs. The foliage is of the large, thin, shade-
leaf type, and the fruits are berries.

The best-known British example of this group,
the Lily of the Valley (Convallaria majalis, Linn.), is
rarely, if ever, found within the Alpine zone, though
it is sometimes frequent at lower elevations. Above
5,000 feet, its place in the forests is taken by the May
Lily (Maianthemum convallaria, Weber = Smilacina
bi/olia, Schult.) (Plate XL VII., Fig. 2), one of our
rarest British plants.

From the creeping root-stock, this plant sends
up each year a flowering shoot, bearing two large,
shortly-stalked, heart-shaped leaves, placed at different
levels on the shoot. Some other Alpine plants produce
only two leaves each year, such as the Lesser Butterfly
Orchis (Habenaria bifolia) (Plate XXXI., Fig. 2), and
the Twayblades (Listera ovata, E. Br., and L.
cordat^ E. Br.), though this is a comparatively rare
feature among either British or Swiss plants. The
flowering shoot of the May Lily ends in an inflores-
cence (raceme) of white flowers, which are probably
smaller than those of any other plant belonging to
the same group. They are interesting from the fact
that there are only four perianth members and four
stamens, whereas the usual number found in this
order is six.

The three British species of Solomon's Seal (Poly-
gonatwri), though rare or infrequent with us, are to be
found in the Alpine woods. Another plant, which,

256 SHADE PLANTS OF THE ALPINE FORESTS

however, is not British, the Knot-foot, Streptopus
amplexifolius, D. C. (Text-fig. XXIV.) is also some-

Fio. XXIV.— A Flowering Branch of the Knot-foot (Streptopus

amplexifolius, D. C.).

Showing the flower-stalks united to the axis for the length of the internode
above their insertion, and the flowers thus hanging below the leaf next
above that in the axil of which they arise.

times met with, though it is rather local in its
distribution. It is an interesting species in many
respects. The flowering shoot is zigzag in form and

THE HERB PARIS 257

knotty. It bears several large leaves, not very unlike
those of the May Lily in shape, but stalkless and
attached directly to the axis, which they clasp by
their bases. The flowers arise singly or in pairs, in
the axil of each leaf, and are mounted on long stalks.
The most curious point about this plant is that
the flower-stalk is united with the portion of the
axis between the leaf in whose axil it arises, and
the next leaf above it. The free portion of the
flower-stalk lies below the upper leaf, and is bent at
right angles, so that the flower hangs downwards.

THE HERB PARIS.

Another forest shade plant, belonging to the same
section of the Lily family as the May Lily, is the
Herb Paris (Paris quadrifolia, Linn.). This species
is one of the most curious and unmistakable plants
in the Alps, as regards its build or habit. The
white, underground rhizome sends up each year a
flowering shoot, about a foot high, which ends in a
single flower. A few inches below the flower are
four, very large, egg-shaped, net-veined leaves, arranged
in the form of a cross, and attached directly to the
axis. The parts of the flower are also arranged in
fours, and not in whorls of three, as in typical
Liliaceous flowers. Further, the outer and inner
whorls of the perianth are dissimilar. The four outer
perianth members are much broader than the four
narrow, inner members, and both are yellowish-green
in colour. There are also eight stamens.

258 SHADE PLANTS OF THE ALPINE FORESTS

Though two whorls of four are usually met with,
other numbers, such as 3, 5, 6, or 7, may occur both
in the perianth and stamens, and then, as a rule, a
corresponding number is found in the whorl of leaves
below the flower. The flowers do not produce honey,
and have an unpleasant smell.

THE MARTAGON LILY.

The Martagon Lily, or Turk's Cap (Lilium
martagon, Linn.) (Plate XLVIL, Fig. 1) is a tall
leafy plant about 3 feet high, with turban-shaped
flowers. It is not common, though it is sometimes to
be found in the forests and thickets, and may occur
even in the Alpine meadows, as at Saas Fee. The
flowers are borne in a large inflorescence, containing
twenty-five or more drooping flowers. The tips of the
perianth members are curled upwards and backwards,
hence the name Turk's Cap Lily. On their inner sides
they are rose-coloured and spotted with dark brown.
They are thick in substance, and have a waxy surface.
Outside, as seen in unopened buds (Plate XLVIL,
Fig. 1), the perianth segments are clothed with
matted white hairs. Each, near its base on the inner
surface, has a deep groove, arched in and protected
by a little flap on either side. This groove is full
of honey, which can be made to ooze out by pinching
the segment at its sides.

The stamens produce great quantities of rust-
coloured pollen, the anthers swinging freely on their
stalks.

PLATE XL VI I

THE MEZEREON 259

The flowers are not very conspicuous, and by day-
time have hardly any scent. But towards evening
they emit a sweet odour, which attracts night-flying
insects such as moths, which are of service as cross-
pollinators. The flower is so constructed that it can
only be entered by insects on the wing. The droop-
ing position, and the recurved perianth segments are
designed for this very purpose, and the latter, with
their smooth, waxy, inner surface furnish no " alight-
ing place " for an insect. To get at the honey, the
moth or other visitor has to hover on the wing below
the flower, and thus its proboscis or tongue comes in
contact with both the stamens and the stigma.
These organs both mature at the same time, so it
sometimes happens that the pollen is deposited by
the insect on the stigma of the same plant, and thus
self-fertilisation, as well as cross-fertilisation, may
take place.

Though the flower, both in the bud and when
fully opened, droops as above mentioned, when the
fruit is mature, the stalk straightens out and the
capsule is borne erect.

THE MEZEREON.

The Mezereon (Daphne mezereum, Linn., natural
order Thymeleacese, the Daphne family) is one of the
three Daphnes, and the only British species, found
in Alpine Switzerland, though the Spurge Laurel
(Daphne laureola, Linn.) occurs at lower elevations.
It is built on much the same lines as the Alpenroses,

260 SHADE PLANTS OF THE ALPINE FORESTS

to which it has an extraordinary resemblance in
habit. It is an erect shrub, 1 to 3 feet high, much
branched at the base, the lower portions of the shoots
being bare and leafless, though showing numerous
leaf scars, marking the position of former leaves.
Little tufts of leaves are borne near the ends of the
branches, and are shed each autumn. In this respect
the Mezereon differs from the Alpenroses, which are
evergreen.

In the spring the flowers appear before the leaves,
as in our British Coltsfoot and other plants. They
are borne in clusters, each of several flowers, in the
axils of the leaves of the previous year. The leaves
themselves disappeared last autumn, but their position
is indicated by the scars on the branches. The
flowers are destitute of a corolla. The four sepals,
however, are united into a rose-coloured tube, and
perform the attractive function. They are sweet
scented, and much visited by flies and other insects.
The fruit is a red berry.

Although the Mezereon is frequent in the forests,
it is not confined to that particular habitat, but is often
to be found growing in the rocky pastures. The two
other species, Daphne alpina, Linn., and D. striata,
Tratt, are easily distinguished by the fact that the
flowers are arranged in umbrella- shaped clusters, and
appear with or after the leaves. The former species
has white flowers, the latter white or rose-coloured
sepals. The calyx tube of Daphne striata is longer
and narrower at the mouth than that of the Mezereon,

THE WINTERGREENS 261

and is specially adapted to fertilisation by insects
with long tongues, such as hawk-moths. The strong,
sweet scent is emitted chiefly in the evening.

THE WINTERGREENS.

All the five British species of the genus Pyrola
(natural order Pyrolacese, the Wintergreen family,
by some botanists included in the Ericaceae, to which
they are closely related) occur in Switzerland, and
most of them may be met with in the Alpine forests.
A non-British species (Pyrola chlorantha, Sw.) is also
found in Switzerland, but it is a rare plant.

The Wintergreens are herbaceous perennials with,
as the name implies, evergreen leaves. The leaves
are stalked, and spring from the creeping stem, close
to the surface of the ground. From among them
erect flowering shoots arise, bearing either a single
flower or a number of flowers arranged in a raceme.
The petals are quite free from one another, and fall
off one by one, when the flower begins to fade. The
anthers open by pores at the tips.

The Single-flowered Wintergreen (Pyrola uniftora,
Linn. ( = Moneses grandiflora, Gray) has a single
solitary pendulous flower, much larger than those of
the other species. The flowers, though conspicuous,
are quite honey less. In the other species the
inflorescence is a raceme and the flowers are smaller.
The Larger Wintergreen (Pyrola rotundifolia, Linn.)
differs from the Common Wintergreen (P. minor,
Linn.) and the Intermediate Wintergreen (P. media,

262 SHADE PLANTS OF THE ALPINE FORESTS

Swartz.), chiefly in certain peculiarities of the styles.
These three species are very nearly related, and are
often difficult to distinguish. The Serrated Winter-
green (Pyrola secunda, Linn.) is easily recognised by
the fact that all the flowers of the raceme or
inflorescence are turned to one side. The leaves are
also more strongly toothed than in the other species.

The pollen-shedding mechanism of the Winter-
greens offers a very pretty study, and is worth investi-
gation. If we examine the pendulous flowers of
Pyrola secunda or P. rotundifolia, we shall find that
when they have just opened, the stalks of the stamens
are bent in an S-shaped manner, and are held in
position by the pressure of the petals. The stalks
are in a high state of tension, like a bent spring. At
this stage the tips of the anthers, where the pores are
placed, are directed towards the base of the flower.
When an insect visits the flower, and pushes back the
petals, the pressure on the spring-like stalk is released,
and it straightens out. The result is that the anther
performs a sort of somersault, turning through 180°,
the tips now pointing to the mouth of the flower.
At the same time, the jerk caused by the release of
the spring shakes out a shower of pollen dust through
the pores on to the back of the insect. This process
can be imitated by simply pressing back the petals of
a flower in which the stamens are ripe, having first
carefully noted the position of the anthers in the
undisturbed condition.

In the Single-flowered Wintergreen the arrange-

THE WOOD SORREL 263

ment is somewhat different. The petals here are
spreading, and do not hold the stamens in position.
The latter, however, have spring-like stalks, which
are so arranged that, when an insect visits the flowers,
they become released, and straighten out in much the
same way as in the other species above described.

The fruit or capsule of the Wintergreens, contain-
ing many very small seeds, opens by little slits. The
fruits, like the flowers, are pendulous, so the slits arise
at the top of the fruit, which is really the base. No
valves are, however, formed, such as are found in the
fruits of the Bell-flowers (p. 88, Text-fig. IX.). In wet
weather, when there is a serious danger of the
seeds, still enclosed in the capsule, becoming damp,
the slits close.

THE WOOD SORREL.

Our graceful little Wood Sorrel (Oxalis acetosella,
Linn., natural order Oxalidacese, the Oxalis family), is
often to be found in the forests of the lower region of
the Alpine zone. The creeping, slender root-stock
sends up a number of long- stalked, characteristic
leaves, each composed of three egg-shaped, delicately
green leaflets. These leaves are believed to be those to
which the name " Irish Shamrock " was first applied,
though the leaves of a species of Clover are now
more generally used as that emblem.

The flowers are borne singly on long stalks, which
arise among the leaves. They are large and white or
pinkish-white, the petals being extremely delicate.

264 SHADE PLANTS OF THE ALPINE FORESTS

The leaves of the Wood Sorrel are of great
interest botanically, for they possess the power of
movement — a rare occurrence among the Higher
Plants. At night, or in very strong sunlight, each of
the three leaflets falls, so as to hang with the apex
pointing directly downwards, and thus to lie parallel
with the leaf-stalk (Text-fig. XXV., 2). Under
ordinary circumstances, in daylight the leaflets are

1. 2.

FIG. XXV.— The Leaves of the Wood Sorrel (Oxalis acetosella, Linn.).
1. In the day position. 2. In the sleep position.

held at right angles to the stalk (Text-fig. XXV., 1).
This means that they receive the maximum of
illumination, whereas in very strong sunlight, by
assuming the second, or sleep position as it is called,
the light only falls very obliquely on the delicate
leaflets. At night, the sleep position protects the
leaflets from excessive loss of heat by radiation.
The mechanism of movement is seated in the very
short stalks of the leaflets, and the changes in

THE WOOD SORREL 265

position are effected by certain tissues becoming more
turgid or more flaccid, by absorption or loss of water.
According as these tissues are swollen or limp, so
the leaflet is held horizontal or sinks to the vertical
sleep position.

It may also be mentioned here that the minute
chlorophyll grains, which contain the chlorophyll
(p. 10) or green pigment of the leaf, have the power
of changing their position in the cells, according
to the intensity of the light. In dull or diffuse
light, they spread themselves so as to obtain the
maximum illumination. In stronger sunlight, they
collect around the walls of the cells, and place them-
selves parallel to the rays of light, so as to receive less
illumination, while, after long-continued sunlight, they
all gather together in groups at the ends of the cells.

The fruits of the Wood Sorrel are interesting
from the fact that the seeds are shot out one by
one, often to considerable distances. The pod, or
capsule as it is termed botanically, is five-chambered,
each chamber possessing two seeds, placed one above
the other, and attached to the central column. Each
seed possesses two coats. Some of the cells of the
inner layers of the external coat, when mature, are
in a high state of strain. They become very tense or
turgid. When each chamber of the fruit opens, the
inner strained layers of the external seed-coat swell
up and burst the outer, non-strained layers, and thus
the external seed-coat is turned inside out. This
violent jerk shoots out the seed itself, enclosed in its

266 SHADE PLANTS OF THE ALPINE FORESTS

shining black internal coat. Thus the Wood Sorrel
presents the curious case of a seed shooting itself off
from the parent fruit by sacrificing one of its seed-
coats. We have already seen how in the Field
Pansy (p. 162) a similar effect is attained in quite a
different way.

THE ALPINE LETTUCE.

Everyone is familiar with our British Coltsfoot
(Tussilago farfara, Linn.), so common on dry bare
slopes, such as newly-made railway embankments,
where it is often the first plant to obtain a footing.
The pale yellow flower-heads of this weed appear
in spring, some little time before the large, broad,
heart-shaped leaves.

In Alpine Switzerland the Coltsfoot is not very
abundant, though, as with us, it is common enough in
the plains. It is then replaced by another near rela-
tive, the Alpine Lettuce (Homogyne alpina, Cass.,
natural order Composite, the Composite family).
Such replacements will be frequently noticed within
the Alpine zone. Our Common Daisy (Bellisperennis,
Linn.) is replaced by the Alpine Daisy (Bellidastrum
michelii, Cass.), while in the High Alpine region the
Alpine Ox-eye Daisy (Chrysanthemum alpinum, Linn.)
takes the place of the Common Ox-eye Daisy (C. leu-
canthemum, Linn.) of the Alpine, Subalpine, and Low-
land zones.

The Alpine Lettuce is, however, a very different
plant to the Coltsfoot. It is essentially a shade plant,

THE ALPINE LETTUCE 267

and, though not confined to the forests, it favours, as a
rule, only the shadier spots in the pastures. The
Coltsfoot, on the other hand, is a marked sun plant,
living in situations not only fully illuminated, but of
the driest description. On the other hand, the Alpine
Lettuce loves a damp situation, a shady retreat, and a
large percentage of humus in the soil. Its leaves are
much smaller, heart-shaped or kidney-shaped, and
very dark bluish-green on the upper surface. The
nerves are very prominent, and, on the lower surface,
are covered with hairs. The leaves appear at the
same time as the flower-heads.

Only a single flower-head is produced, and this is
borne on a long, brown stalk. The flowers are of a
dull, brownish-red tinge, which is quite unmistakable.
The corollas of the outer ray (female) flowers are not
strap-shaped, as in the Coltsfoot, or Arnica, or Ox-
eye Daisy, but divided into thread-like segments — a
very rare feature among the Compositae.

CHAPTER XI

ADAPTATIONS AMONG ALPINE PLANTS

THE present chapter will be devoted to a discussion
of certain adaptations found among Alpine plants.
In the previous sections we have repeatedly drawn
attention to the mutual dependence or alliance which
exists between a large number of Alpine flowers and
members of the insect world. We may now enter
on a more general consideration of this subject.

FLOWERS AND INSECTS.

It is a matter of common observation that insect
life abounds in Alpine Switzerland ; so do con-
spicuous and beautiful flowers, and" for the good
reason that a large percentage of Alpine plants are
specialised for cross-pollination through insect agency.
They have found that, to ensure and maintain the
fertility and robustness of the stock, the ovules of one
flower must be fertilised by pollen brought from
another plant, and not by the pollen of the same
flower. The two chief agencies which act as pollen
carriers are insects and the wind. Such is a plain

FLOWERS AND INSECTS 269

statement of the case from the plant's standpoint.
From the insects' point of view, an abundance of
flowers is essential, for many of them live entirely on
a diet of honey or pollen, or both combined. We
have no evidence that insects ever visit flowers for
philanthropic or disinterested motives. They go bent
on the serious business of marketing, just as many
human beings go daily to the markets to obtain the
necessaries of life.

These facts must always be carefully borne in
mind. They form the key to the origin of many of
the varied peculiarities in the form and structure of
the flower, to be met with in the Alps. The botanist
who first clearly noted these facts was Joseph Gottlieb
Kolreuter (1733-1806) of Karlsruhe. He was shortly
afterwards followed by Christian Konrad Sprengel,
to whom reference has already been made (p. 2).
Thus Kolreuter and Sprengel are the honoured Fathers
of the study of the manner of pollination of Flowering
Plants.

There is no doubt whatever, from the evidence of
palaeontology, that insects existed long prior to the first
appearance of the Flowering Plants (Angiosperms).
It is conjectured that the earliest primitive flowers
were wind pollinated, but owing to the probability
that they were soon visited by insects, who robbed
them of their pollen, the plants appear to have
determined that, if they must suffer robbery, they
might as well make use of the insects in some way,
and consequently they hit on the happy idea of

270 ADAPTATIONS AMONG ALPINE PLANTS

making them the pollen distributors. So successful
was the move in this direction, that it quickly became
" fashionable " to adapt the flowers especially to
insect visitors. Since then many plants have become
more and more specialised, as regards the flowers,
in relation to insects, and all sorts of floral mechanisms
have been devised, some of which have been discussed
in the preceding chapters.

We will now notice some further points in this
connection. The late Hermann Mtiller, our greatest
authority on the fertilisation of Alpine flowers, calcu-
lated that the proportion of insect- to wind-fertilised
plants in the Alps was as follows : —

Insect-fertilised species . 590 = 84 per cent.
Wind-fertilised species > 109 = 16 „

Total species 699 100 „

Another prominent worker in the same field, Loew,
found that the proportions in the plains and lowlands
of Central Europe is : —

Insect-fertilised species . 981 = 78 per cent.
Wind-fertilised species ^ 271 = 22 „

Total species . 1252 J100 „

Thus there is a relatively larger number of insect-
fertilised plants in the Alps than in the Lowlands, and
a smaller number of wind-fertilised plants.

The chief insect-fertilisers in the Alps are butter-
flies, moths, humble-bees, and honey-bees. Certain
flies also cross-pollinate some flowers, such as the
Globe-flower (p. 207) and the Two-flowered Violet

FLOWER ADVERTISEMENTS 271

(p. 253). Miiller showed that in the Alps, butterflies
play a more important part in this connection than
in the plains.

Flower Advertisements.

Flowers may be said to be the most ancient of
advertisers. Those which are pollinated by insect
agency specially lay themselves out to attract insects.
Their advertisements are of two main types : colour
and scent. As a rule, in the Alps both exist together ;
in other cases colour alone is relied on, and the
flowers are scentless. But many plants go even a
stage further : they provide " free samples " of honey
or pollen for some or all of the insects whose com-
pound eyes they have managed to attract by their
advertisements.

Conspicuousness may be more or less confined to
the flowers themselves, or other parts of the plant
may share in it. We have already seen (p. 124)
how diflerent parts of the flower may be specialised
as the conspicuous organs. We have also discussed,
in the case of the Gentians (p. 58), the evolution
of the coloured pigments of flowers. Frequently one
or more organs or sets of organs are greatly enlarged,
and highly coloured, and are thus rendered more
conspicuous. A good example of this may be found
in a comparison of the corollas of the Field Pansy
(p. 161) or Long-spurred Violet (p. 128) with that of
the Two-flowered Violet (p. 253). Or, again, con-
spicuousness may be increased by the massing of a

272 ADAPTATIONS AMONG ALPINE PLANTS

large number of flowers in an inflorescence, to form
a head, as in the Rampions or Composites. Or,
again, if the inflorescences become one-sided, all the
flowers turning in one direction, as in the Bearded
Campanula, and many Boragineae, conspicuousness
is markedly increased.

As regards scent, the odours of many flowers are
well known to be characteristic, such as those of the
Mignonette, or Jasmine. Yet other quite unrelated
plants often possess exactly the same kind of scent.
A good example is the vanilla scent, typical of the
fruits of a tropical climbing Vanilla Orchid, which is
possessed by many other flowers, among them the
Black Nigritella (p. 135) and the Linnaea (p. 250).

Again, many plants possess what to us are
unpleasant or nauseous odours, which apparently are
quite acceptable to certain insects. Such flowers are
rare in the Alpine zone, the most unpleasant being
Thalictrum aquilegifolium, Linn., which has the Elder-
odour.

We now turn to the "free samples" offered to
insect visitors. Pollen alone is rare, though in the
Alps the flowers of the Alpine Anemone (p. 37)
and the Narcissus-flowered Anemone (p. 41), the
Alpine Poppy (p. 195) and Thalictrum aquilegifolium,
Linn., possess no nectar. As a rule, however, nectar
or honey together with pollen are the inducements
which entice insects to enter the flower.

Honey is secreted by special glands known as
nectaries, the position and shape of which varies in

CONCEALMENT OF HONEY 273

different flowers. In the case of the Yellow Gentian
(p. 58), we saw that the nectary was a ring-shaped
swelling at the base of the ovary. In other flowers
certain of the stamens are modified to form honey-
glands, and have quite lost their original function.
Such cases are well seen in the Globe-flower, the
Hellebores, and the Monkshoods. In the Violets
(p. 161) and Orchids, as well as many other
plants, the nectary or nectaries are situated in a
spur.

In some plants, such as the Yellow Gentian and
the Marsh Marigold, the nectar and pollen are freely
open to all insect visitors, which arrive on the wing,
though not to crawling insects. In other cases the
flowers are specialised for certain insects alone, such
as humble-bees and butterflies, which possess very
long probosces or tongues. This implies concealment
of the honey or some adaptation by which such
visitors can alone enter the flower. In the previous
chapters we have discussed several instances. In
some Gentians, the throat or entrance to the corolla
is closed, by means of scales, to all except the more
powerful insects, and the same adaptation is met with
in other plants. Again, as in members of the Pea
family, the flower may be so shaped that only certain
insects can force their way within. The honey may
be concealed at the end of a long corolla tube of fine
bore, as in Daphne striata (p. 260), or in slender spurs,
as in Viola cakarata (p. 129), so that it can only be
reached by long-tongued insects. In the case of the

274 ADAPTATIONS AMONG ALPINE PLANTS

Globe-flower (p. 207), only very small flies can reach
the honey.

The different contrivances to be met with in
flowers adapted to particular insect visitors are
extremely numerous, but it will not be possible to
enter into this subject fully here. These remarks,
supplementary to the observations scattered through
the preceding chapters, may, however, serve to indicate
the chief points of interest of this nature to be met
with among Swiss Alpine plants.

Before we leave the subject, we may explain how
insects recognise the flowers they visit. This subject
has been much discussed, and there is still a great
deal to be learnt in the matter, but certain facts seem
now clearly established. The eyes of insects are com-
pound, and further, they are immovable. The number
of "facets " in a single eye may vary from 4 to 25,000.
Despite this fact, insects are extremely short-sighted.
At one time it was thought that each facet reflected
the image of a flower, so that if 10,000 facets occur in
a single eye, the insect would receive 10,000 different
images of the flower from one eye alone. This
view, multiple vision as it was called, is now
abandoned, and it is believed that each facet only
receives the image of a small part of an object, the
whole object being seen by the total sum of the
images of each facet. This may perhaps explain the
short-sightedness of insects. At any rate, it appears
clear that though insects are attracted to a particular
flower from a considerable distance, it is by the sense

UNBIDDEN GUESTS 275

of smell, and not by sight. When the insect begins
to get near the flower, then the colours attract its
eyes, and often " honey guides" (p. 161) are furnished
by the plant to help it to find the entrance.

Protections against Unbidden Guests.

We have seen how certain flowers are specialised
for certain types of insects, and we are now in a
position to appreciate the safeguards possessed by
many Alpines, whereby insects or unbidden guests,
which can perform no service to the flower by acting
as cross-pollinators, are warned off* or excluded.
Some plants, not so well adapted to meet this
difficulty, are frequently robbed of their honey or
pollen, by insects which are not likely to visit another
flower of the same species.

Unbidden guests are of two classes : those
which approach the flower on the wing, and those
which crawl up from below by means of the flower-
stalk. The former are excluded, as we have seen, by
the concealment of the honey, or by the special shape
of the flowers, which precludes their entrance, or by
the closure of the throat of the flower by special
means such as scales, or by the presence of tufts of
hairs or bristles inside the flower, which prevent
some insects penetrating as far as the honey. In
certain species, such as the Martagon Lily (p. 259)
and Lychnis vespertina (p. 165), which are adapted
for fertilisation by night-flying Lepidoptera, the plants

276 ADAFPATIONS AMONG ALPINE PLANTS

do not emit scent in the daytime, and thus do not
attract many visitors.

The inflated calyx of Silene cucubalus (p. 164)
probably serves to protect the honey from robber
bees, who try to steal it by biting through the base of
the corolla.

Some subtropical and tropical plants have remark-
able methods of protecting themselves from unbidden
guests whether visitors to the flowers or leaf-
destroyers. They actually hire other insects to
protect them, and produce nectaries outside the
flowers, as an inducement to their protectors to stay
and fight their battles for them. Kerner states that
the flower-heads of Centaurea alpina, and some other
Composites in Southern Europe, maintain a colony of
ants by means of special nectaries, on the involucral
bracts of the flower-heads, on which they feed. These
ants act as a body-guard against certain beetles or
other unbidden guests, which eat the flower-buds.
The ants are able to drive away the beetles, ejecting
formic acid if necessary, and they thus perform a
good service to the plant. Several other tropical
plants go much further. They form a group called
myrmecophilous plants. They not only feed, but
actually house standing armies of fighting ants, which
keep off leaf-cutting insects and other unwelcome or
injurious visitors.

We now turn to the other class of unbidden
guests, the crawling insects. A very simple device,
which is sometimes adopted, to keep such insects

WIND- AND SELF-POLLINATION 277

away from the flower, is the production of nectaries
for their benefit on other parts of the plant. Or, again,
the stems, leaves, and the stalks of the flowers may
be armed with stiff hairs, prickles, or thorns, which
prevent soft-skinned creeping animals, or very small
insects from climbing up to the flowers. Hairs
frequently occur on the calyx and on the petals,
which serve the same function, especially where the
flower droops, the bell of the corolla pointing directly
downwards to the soil. Examples of these types
of protection occur in the Bearded Campanula
(p. 89), the Tufted Campanula (p. 91), and the
Linnaea (p. 250),

In other plants, the flower-stalks are rendered
slippery by a coating of wax, or the very smooth,
polished surface of the petals affords no hold for the
insect climber, or the secretion of some sticky material
by glandular hairs, on the stems or leaf-stalks, or
calyces, serves the same function. In other cases
different contrivances are met with, all fashioned to
the same end, but the above will perhaps suffice as
some indication of the commoner types of protection
met with in the Alps.

Wind-pollination and Self-pollination.
Although, as we have seen, insect pollination is the
rule among Alpine plants, there are many exceptions
which are either wind-pollinated or self-fertilised. The
Coniferous trees and shrubs, for instance, are all wind-
pollinated. The same is true of the Green Alder, the

278 ADAPTATIONS AMONG ALPINE PLANTS

whole of the Alpine Grasses, Eushes, and Sedges,
and some others. In such plants the individual
flowers are nearly always small and inconspicuous,
and either devoid of calyx and corolla, or possess
a very reduced, inconspicuous perianth. Another
striking feature of wind-pollinated plants is that the
flowers are often aggregated into dense clusters,
or inflorescences.

Self-fertilisation — the fertilisation of the ovules of
a flower by pollen derived from the anthers of the
same flower — is also common, and the seeds that
result may be either quite fertile or abortive.
Many flowers, which are normally cross-pollinated
by insect agency, may, if no insect visits the flower
or no foreign pollen is brought to their stigmas,
fertilise themselves at the end of the flowering stage.
Some flowers possess special adaptations to ensure
self-fertilisation, if cross-pollination fails.

Some plants produce flowers which never open
— cleistogamous, as they are called botanically — in
addition to ordinary flowers. This phenomenon is
not very frequent in the Alps, though it is sometimes
to be found in the case of certain Gentians, Gentiana
tenella, Rottb., and G. campestris, Linn., and in the
Wood Sorrel. As a rule, such cleistogamous flowers
in the Alps are of quite normal structure, though,
either through lack of sufficient light or warmth, they
never open. In the Lowlands some plants produce
flowers which never open under any circumstances,
such as certain Violets and Dead Nettles (e,g.,

PARTHENOGENESIS 279

Lamium amplexicaule, Linn.), which are differently
constructed from the normal flowers borne by the same
plants.

The extraordinary phenomenon called partheno-
genesis, which sometimes occurs both in plants and
animals, in which the ovule is able to develop
normally without having been fertilised at all by the
male sperm produced by the pollen grain, has so far
only been observed in the Alps in the case 'of certain
species of Lady's Mantle, genus Akhemilla (natural
order Eosaceae, the Rose family). Another case
is known in the Arctic regions — namely, Antennaria
alpina, Gsertner ( = Gnaphalium alpinum, Linn.), a
plant nearly related to the Swiss Everlastings (see
p. 21). Not only are no male flowers of this species
known in the Arctic regions, but the female flowers,
which in this genus are borne on different plants
to the male, regularly set their seed without having
been pollinated at all.

Before leaving this subject, we may note that in
the tropics certain flowers are fertilised by animals
other than insects, such as bats, humming, and
honey-sucking birds, snails, and slugs.

The Protection of Pollen.

Everyone who has travelled in the Alps has been
impressed, often in a disagreeable manner, by one fact
— namely, the variableness of the weather. In the
mountains the changes in the condition of the weather
are often extremely rapid. For instance, the early

280 ADAPTATIONS AMONG ALPINE PLANTS

morning may show a bright sun, shining in a
cloudless sky : yet before the afternoon, everything
is changed. The peaks are wrapped in clouds,
mists fill the valleys, and rain descends in torrents,
or snow may take its place. The traveller caught
afield under such changed conditions compared
with those under which he set forth, seeks some
form of protection from the weather. But how
do the flowers meet this difficulty? Here is a
meadow, or there an Alp in full bloom. That is
to say, the stamens of the flowers are shedding the
precious pollen dust, on which so much depends. The
pollen, in nearly every case, is ruined at once if
thoroughly soaked. It is drowned, as it were, and all
the labour of the plant and flower will, in that case,
be in vain. This is a possibility against which an
Alpine plant has to be specially on its guard.

The contrivances to be found among Alpine
flowers are as varied as they are interesting. In
many cases the permanent drooping position of the
flower at the end of a bent stalk meets the case
admirably. We have already discussed examples of
pendulous flowers, as, for example, the Alpine Bell-
flowers, the Soldanellas, and the Water Avens. In
other plants the flower is so placed that the tube of
the corolla is held horizontally parallel to the soil,
and thus the petals shield the stamens. This method
of protection is especially effective when the throat
or entrance to the corolla tube is contracted or closed
in some way. Examples of such plants are to be

POLLEN PROTECTION 281

found in the Alpenroses, the Primulas and Androsaces,
the Butterworts, Violets, and Monkshoods.

More rarely the stamens are never exposed at all
to the weather. In the Globe-flower (p. 207), the
sepals form an arched roof over the stamens, while
in the flowers of the Pea family (Leguminosae) and
in the Box-leaved Polygala (p. 253), the stamens
remain enclosed in the petals.

But in other plants where the flowers are held
with their stamens freely exposed to the weather,
special contrivances exist to protect the pollen. We
have seen how, in the case of the Carline Thistle
(p. 125), the bracts of the flower-head curve up over
it in bad weather. The flowers of the Anemones,
Gentians, Crocus, Colchicum, and others close entirely.
In other plants, such as the Common Daisy and the
Jacob's Ladder (p. 151), the flower-stalks curve at
night and in bad weather, so that the flowers, instead
of pointing upwards, nod towards the soil. Other
plants in which the stamens are freely exposed, such
as the Thalictrums, the Plantains, the Globularias,
and the Lady's Mantles (Alchemilla), meet the
difficulty by simply closing the anthers and ceasing
tp emit pollen until a more favourable season. The
anthers of Thesium alpinum, Linn., are said to close
within thirty seconds of their being moistened.

It is obvious that the heavy night dews of the Alps
must have an effect similar to that of a slight shower of
rain, as regards danger to the pollen. Hence, most of
the types of movement mentioned above are normally

282 ADAFIATIONS AMONG ALPINE PLANTS

to be observed towards evening. It is probable that
rapid changes in the illumination give the signal to
the plant to close its flowers or anthers, or to droop
the flower or flower-head. Atmospheric changes of
a marked nature are usually foreshadowed in the
Alps by some variation in the intensity of the
illumination.

These special contrivances to prevent the flower
or the stamens becoming water-logged are extremely
interesting botanically, for they necessitate the move-
ment of one or more sets of organs. Yet, speaking
generally, the power of movement is as comparatively
rare among plants as it is common among animals.
Hence the mechanisms which effect the opening and
closing of flowers and the like are of great importance,
though unfortunately it is not possible to enter into
the matter here in detail.

SEEDS AND FKUITS.

In the preceding chapters we have called attention
to some of the chief difficulties to be faced, as regards
climate, by plants living at high altitudes in Switzer-
land. It must not, however, be imagined that the
Alpine region, even with its manifold disadvantages,
is unsuited to support plant life. This is far from
being the case. The High Alps are extremely favour-
able to life, provided only that plants can adapt
themselves to the special conditions which there rule.
Early flowering and perennial existence are two of
the necessary conditions. Another is an abundance

SEEDS AND FRUITS 283

of seed-production, implying in many cases a wealth
of blossom.

A large number of the seeds set and ripened each
year are doomed to failure. The seeds or fruits are
the new colonisers, which, transported to a distance
from the parent plant, by one agency or another,
seize upon such new ground as is available. They
have to fight not only against climatic conditions,
sufficiently severe to eliminate the majority of the
seeds annually distributed, but to compete for space,
light, air, and moisture with the seeds or seedlings
of other plants, or to contend with an established
adult population, for the most part of a perennial
nature. The existence and welfare of the species
is dependent on the prosperity of the new generation,
which begins with the seed. The first step is the
successful establishment of the seedling, if possible at
some distance from the parent. It may now be well
to consider how this object is furthered among
Alpine plants.

Early flowering and an abundance of flowers are
simply the means whereby a few, out of a large
number of seeds matured, may gain a hold in their new
surroundings, before the short summer and autumn
merges into winter. The contrivances for the dis-
tribution of the seed or a group of seeds, either free
or still enclosed in the fruit, are as varied in the Alps
as in the plains of Britain, but they are not, for the
most part, dissimilar to those met with in this
country.

284 ADAPTATIONS AMONG ALPINE PLANTS

In the majority of cases the seeds are shed
individually from the fruits, and distributed chiefly by
the agency of wind, which plays an especially
important part as a disseminator within the Alpine
zone. Dr Vogler has shown that the number of wind-
distributed species in the Alps is nearly 60 per cent,
of the whole flora — that is, more than half as much
again as the number in the Lowlands (38 per cent.).
The percentage of animal-distributed plants is only
3 per cent., whereas in the plains it is 15 per cent.

The great majority of Alpine species have very
minute seeds, as in the case of the Saxifrages,
Campanulas, and Primulas, which are shed individu-
ally from the capsule or fruit. As a rule, these seeds
possess no special adaptation to wind-distribution,
beyond their small size and lightness.

Some seeds, however, possess tufts of hairs which
render them admirably adapted for wind-distribution,
as in the case of the Willowherbs (EpiloMum) and the
Alpine Willows (Salix). More frequently the seed
remains enclosed in the fruit, which may develop
hairs adapted to dispersal by the wind. Such fruits
are common to many members of the order Com-
positae. The parachute-like hairs of the fruit of
the Dandelion (Taraxacum) afford an excellent
example.

In the case ofAtragene (p. 230), Dryas (p. 108), and
some species, but not all, of Anemone (pp. 36 and 39),
the style of the ovary develops into a long, feathery
awn, which enables the fruit, which here encloses

DISTRIBUTION OF SEEDS 285

only a single seed, to travel considerable distances
with the wind (Plate XLIII, Fig. 2, and Plate V.).

In a few plants the seeds are thrown out by the
agency of the fruit itself, as has been already explained
in the case of the Violets (p. 162). In other cases,
such as the Geranium, the ovary divides into five
fruitlets, which are shot off from the plant to a
considerable distance by the sudden splitting of the
fruit.

The fruits, distributed by the agency of animals,
are of two kinds. The first are succulent and edible
and attract animals, especially birds. The second
possess some form of clinging apparatus, such as hooks,
by which they become attached to passing animals,
and are thus carried to a distance from the parent
plant.

We have in an earlier chapter (p. 220) com-
mented on the number of plants producing berries in
the Alps. The other types of succulent fruits are
few in number. In the Wild Strawberry (Fragaria)
the receptacle of the flower becomes fleshy, and the
fruits, each of which only contains a single seed, are
the little black bodies scattered over the red flesh of
the ripe fruit. The " hips " of the Eoses (Rosa) are
complicated structures, in which the persistent and
flesh-coloured calyx tube of the flower grows
round and encloses the true fruits, each of which
also contains a single seed.

Hooked fruits are comparatively few in the Alps.
We have already noticed (p. 158) the hooked style of

286 ADAPTATIONS AMONG ALPINE PLANTS

the Geums. The fruits of species of the Cleavers
(Galium aparine, Linn.), also a common British
plant, are covered with hooked bristles, the clinging
power of which is too well known to need description.
It is believed that some seeds in the Alps are
distributed by ants. There is a delicate little wood-
land plant, the Cow-wheat, Melampyrum sylvaticum,
Linn, (natural order Scrophulariacese, the Foxglove
family), common in the Pine woods, and, like the
Louseworts (p. 218), a semiparasite on the roots of
other plants. The fruits contain two (sometimes
only one) white, shining seeds, which are of quite
unusual appearance, and very conspicuous. If the
ripe fruit be squeezed slightly, the seeds will pop out
suddenly, and are shot away for some little distance.
If we place some of these glistening, white seeds
among the roots of a Pine, about which there are
sure to be a number of ants busily engaged, we
shall find that the ants instantly turn their attention
to them and eagerly carry them off to the nest, as if
they had at length found a great prize. Whether
the seeds of the plant are distributed only by the
agency of ants is as yet "not proven."

THE DISEASES OF ALPINE PLANTS.

It is too little recognised that plants, like animals,
are subject to various diseases, often infectious and
frequently fatal. Alpine plants form no exception to
the rule. These diseases arise from two sources :
parasitic plants and parasitic animals. The former

THE DISEASES OF PLANTS 287

are, as a rule, the more numerous, as well as the
more disastrous. The subject of plant disease is too
difficult and complicated to be discussed here, except
in the barest outline. We may, however, learn how
to recognise the conspicuous signs of disease so
commonly to be met with in the Alpine zone.

We will first notice the effects of certain diseases
due to the presence of parasitic plants, living on or
within the host, as the plant attacked by them is
termed. These parasites nearly all belong to the
group of Fungi, a race of plants consisting of threads
of cells, destitute of green colouring matter or chloro-
phyll. The Mushroom is one of the higher and more
advanced of the Fungi, as regards organisation. The
Moulds, Mildews, tod Rusts are other examples,
lower in the scale. Some of them are saprophytes,
living on dead and decayed vegetable or animal
matter. Others are parasitic on or in living tissues.

Some fungal parasites may only infect certain
portions of the host plants, such as the leaves, at any
rate during some stages of the disease. They are
thus local in their effect. They injure certain portions
of the leaves or the stems of the host plants, but
they do not, as a rule, imperil their existence. But,
more commonly, the injuries are more widely spread
throughout the plant. The very small threads of
colourless cells — often quite invisible to the naked
eye unless many thousands of them are matted
together into a felt, as in the Moulds and Mildews —
not only rob the host of its own nourishment, but

288 ADAPTATIONS AMONG ALPINE PLANTS

actually eat up the substance of the host plant itself.
If the leaves are attacked, the plant loses its power of
assimilating (p. 10). If the flowers or fruits are
attacked, as in the case of the Smuts on wheat, few
or no seeds result. The loss on crops of wheat, rye,
oats, and other cereals may amount to many millions
sterling in one country alone in a single year, when a
particularly bad epidemic of disease has prevailed.

The commonest signs of disease are the presence
of spots, patches, or swellings on the leaves or shoots
of the host, often dark in colour, or again, brightly
pigmented. Little pustules of various colours also
break out on the surface of the stem or leaf. Often
the plant loses some of its green colour, and becomes
whitish, or shows some other obvious signs of ill
health. The number of plant diseases known is
almost endless. Each disease has its own symptoms,
and these depend on the nature of the life history of
the particular parasite in question.

One special feature connected with many plant
diseases, is that the host has the power of accom-
modating itself to the disease in various ways. The
presence of a parasite, whether plant or animal, may
stimulate the host plant to adapt itself against the
invader by producing certain modifications of its
normal form. The growth of the branch or other
organ attacked may be either arrested or greatly
increased, or some other abnormality may result. In
some cases the host becomes reconciled, as it were,
to the parasite, and goes out of its way to provide for

THE DISEASES OF PLANTS 289

its needs. Thus the gravity of the attack is often
minimised, and the disease restricted to local areas.

The well-known "Witches' Brooms," frequently
seen on Cherry, Spruce, and many other trees, afford
an excellent example of abnormal growths in response
to the attack of a Fungus. These shoots are always
quite erect, however drooping the ordinary branches
may be. They arise from diseased buds which grow
abnormally. Such shoots never produce flowers or
fruits. They are branches specialised for the fungus
in which the parasite finds a home suited to its needs,
and thus the disease is localised, and does not spread
throughout the whole plant.

Diseased organs are thus frequently dissimilar in
appearance to healthy structures. This is often well
seen in the case of the House-leeks (Sempervivum)
(see p. 98). If we examine the rocks in summer-time,
in a locality where these plants are abundant, we
shall often find that some rosettes occur in which the
leaves are either very pale or red in colour. They
are also distinctly larger, as well as three or more
times as long as those of the healthy rosettes (cf.
Plate XXIV., Fig. 2, with Fig. 1 of same Plate. These
rosettes are attacked by an internal parasitic Fungus,
known as Endophyllum sempervim, and in the large
size of the diseased leaves we see an adaptation to
the invader.

Another excellent example of dissimilarity between
a healthy and a diseased plant may be frequently
observed in the case of a Spurge (Euphorbia cyparis-

290 ADAPTATIONS AMONG ALPINE PLANTS

sias, Linn., which is a very abundant plant in the
Alps, especially in dry places. The healthy shoots are
much branched, and bear numerous large umbels of
complicated inflorescences which resemble flowers,
the golden-yellow bracts of the inflorescence simulat-
ing a corolla. Frequently among the clumps of this
plant we shall find small unbranched shoots, bearing
pale leaves, sometimes slightly larger than the leaves
of the healthy plant, and often rust coloured on the
lower surface. These shoots never bear flowers. A
dense, little rosette of leaves is often found at the
apex. These are shoots specialised to a parasitic
Fungus (Uromyces pisi), allied to the Eusts of
wheat, which lives within them, and thus the parasite
is for the most part localised to such shoots, and
does not spread indiscriminately through the whole
plant.

A familiar instance of response to disease is the
production of galls. These may be caused either by
plant invaders or by parasitic animals, chiefly insects.
The tissues of the part of the host plant attacked by
the parasite, be it a leaf or be it a branch, are pro-
foundly modified by the presence of the invader, and
stimulated to renewed, and often excessive, growth,
and the formation of new tissues or organs of unusual
shape and size, which are called galls. The well-
known "B/obin's Pin-cushions" on Rose leaves, and
the "Oak Apples" on Oak twigs, are two common
examples out of many thousands of varieties of
galls, due to insect attacks. Galls of various forms

THE LICHENS 291

frequently occur on the leaves of Alpenroses, the
Alpine Currant, and many other plants.

In other cases galls are produced by the stimulus
of parasitic fungi. Galls of this nature may often be
observed on the leaves of the Alpenroses. A large
spongy body, spherical or oval or somewhat irregular in
shape, arises from a portion of the leaf, and may grow
almost to the size of an apple, though it is frequently
much smaller. This gall is of a pale yellow colour,
often tinted with rose on the side turned towards the
sunlight. These galls have been called Alpenrose
Apples. The Fungus which stimulates these new
formations is known as Exobasidium rhododendri.

THE LICHENS.

It is often imagined that the large Lichens, especi-
ally the greyish-green Beard Lichen ( Usnea barbata)
(Plate XLIV., Fig. 2), which festoon the branches of
the Spruce and other Conifers in Alpine Switzerland,
are parasitic upon these trees. This is not, however,
correct. The Lichens simply attach themselves to the
trees, to which they do little or no harm. Many plants,
especially in the tropics, though much more rarely in
temperate climates such as that of Western Europe,
are what is called epiphytic. That is to say, while
they attach themselves to other plants, they do not live
parasitically at their expense. They neither penetrate
nor destroy their tissues, nor rob them of their
food- supply. The attached plant remains entirely
self-supporting.

292 ADAPTATIONS AMONG ALPINE PLANTS

The Lichens are exceedingly abundant plants in
the Alps, and very interesting organisms from many
points of view. Though in this volume we are only
concerned with the higher members (in the botanical
sense) of the vegetable kingdom, occurring within the
Alpine zone, we may, however, here make an excep-
tion and add a few remarks on the subject of the
Lichens.

In addition to the Bearded Lichens, another, a
very different type, may be observed on almost any
large boulder. It will be frequently noticed that
yellow, green, or brown patches are to be seen firmly
encrusted on bare rocks, forming what might almost
be described as films, adhering closely to the surface,
and without any definite outline. These patches are
the Crustaceous Lichens.

The Lichens are composite organisms ; that is to
say, the body of the Lichen is built up of two plants
living together, closely intertwined. One of these
plants is a member of the Algae, the group to which
the Seaweeds, etc., belong. The other is a Fungus.
The Alga, which is green, and the Fungus, which is
colourless, are so closely interwoven together that the
resulting structure, or Lichen, is quite dissimilar from
any single Alga or Fungus. The constituents of the
Lichen are not individually visible to the naked eye,
and it requires careful microscopic examination to
detect them.

The life history of a Lichen is too complicated to
be entered into here. It may, however, be added

SYMBIOSIS 293

that the two constituents of the Lichen have come, as
it were, to an agreement as to the part which each
is to play. The Alga, by means of its green chlorophyll,
undertakes the manufacture of the food-supply, while
the Fungus looks after the reproduction of the species.
This extraordinary mutual benefit society of two quite
different plants — a condition which the botanist calls
symbiosis — is not confined to the Lichens, though they
form by far the largest group of symbiotic plants.

CHAPTER XII

THE GEOGRAPHICAL DISTRIBUTION, AFFINITIES, AND
ORIGIN OF THE SWISS ALPINE FLORA

IN this, the concluding chapter, we will briefly review
some points of interest in regard to the geographical
distribution of Swiss Alpines, and the theories
advanced to account for the origin of the Alpine flora
of Europe.

INEQUALITIES OF DISTRIBUTION.

It has been already pointed out that some districts
of Alpine Switzerland are much richer in species, and
especially in the rarer plants, than others. Also the
converse — that certain Swiss Alpines are confined to
particular districts. It was stated nearly forty years
ago, by a great authority, that whereas the Bernese
Oberland only offers a single species not found
elsewhere in Switzerland, the chain of Alps of the
Valais, lying to the south of the Ehone valley, contains
sixty-three species peculiar to it. Whether these
figures would be accepted as accurate or not to-day is
really immaterial to the argument. The Alps of the

294

RARE ALPINE PLANTS

295

Cantons Valais and Grisons (Graubunden) are
certainly much richer in Alpine species than any
other district on the north side of the main Alpine
chain.

Among the rarities confined to the Valais are : —

Anemone halleri, All.
Silene vallesia, Linn.
Oxytropis fcetida, D. C.
Potentilla multifida, Linn.
Trifolium saxatile, All.

Saxifraga diapensioides,

Bell.

Valeriana celtica, Linn.
Senetio uniflorus, All.

In the Engadine, and elsewhere in the Grisons, the
following species are found : —

Papaver nudicaule, var.

rhceticum, Leresche.
Dianthus glacialis, Hanke.
Sempervivum wulfeni,

Hopp.

Senecio carniolicus, Will.
Valeriana supina, Linn.
Pedicu laris jacquini,
Koch.

which are either unknown elsewhere in Switzerland
or only occur in the southern Canton of Tessin.

Rare species, common to both the Valais and
Grisons, but practically confined to these Cantons, are,
among others : —

Arabis halleri, Linn.
Sempervivum funckii,

Braun.
Pedicularis incarnata,

Jacq.

Potentilla nivea, Linn.
Phyteuma humile,
Schleich.

296 DISTRIBUTION OF SWISS ALPINES

Primula longiflora, All., and Phyteuma pauci-
florum, Linn., are rare plants found in the Valais,
Orisons, and Tessin.

How can these inequalities of distribution be
explained ? Some of the older observers attributed
the facts to the effect of soil or climate alone. There
is, however, little doubt that this explanation is
insufficient. A celebrated Swiss botanist, the late
Alphonse de Candolle of Geneva, has left us a more
plausible theory.

It can scarcely be doubted, even by the most
sceptical, that the Alps were at one time more heavily
glaciated than they are at present. The retreat of
the glaciers, with certain exceptions, has been marked
even within the last few decades. The amount of
glaciation appears, however, to have been as unequal
in the past as it is at present, and it is to this fact
that De Candolle attributes the observed inequalities
of distribution among Alpine plants. The southern
chain of the Valais and the eastern Ehaetian Alps
appear to have been less glaciated than the central
Oberland region and northern Switzerland generally ;
and further, the glaciers there began to retreat earlier
than in the Oberland. The consequence has been, on
this theory, that the Valais and Grisons were being
colonised by Alpines, while the Bernese Alps remained
a vast waste of ice or snow.1

1 According to Prof. Jaccard, the number of species to be found in
any locality is directly proportional to the diversity of its ecological
formations.

AFFINITIES OF THE FLORA 297

The question, however, is naturally bound up with
the greater problem of the origin of the Alpine flora
of Switzerland, and its relation to the vegetation on
the north and south of the Alps. To this question
we will now turn. Before we attempt to summarise
the theories which have been advanced, we may first
set out the facts which such theories should explain.

THE AFFINITIES OF THE Swiss ALPINE FLORA.

The mountain ranges of Central Europe form an
incomplete chain, stretching roughly west and east,
beginning with the Pyrenees, then the Swiss and
Austrian Alps, and finally the Carpathians. The
widest gaps in the chain are between the Pyrenees
and the Swiss Alps, and between the Austrian Alps
and the Carpathians. The Alpine floras of these
mountain regions are essentially similar, many species
being common to them all. Although, in the Swiss
Alpine region, there is probably no species which
is not found elsewhere, there are, on the other hand,
many plants in the Austrian Alps and the other
ranges, which do not occur in Switzerland. At the
same time, there is a distinct unity in type and marked
relationship between the floras of all these Alpine
chains.

Further, many Swiss Alpine species occur so far
afield as the mountains of Central Asia, such as the
Himalayas and the Altai. The Edelweiss (p. 15),
for instance, is found in the Pyrenees, the Swiss
Alps, the Tyrol, the Carpathians, and the Himalayas.

298 AFFINITIES OF SWISS ALPINES

Schroeter states that 275 Swiss Alpines occur in the
Pyrenees, 399 in the Austrian Alps, 231 in the
Carpathians, 129 in the Altai, and 72 in the Hima-
layas. We see, therefore, that many Alpines are
very widely distributed, though confined to Alpine
regions, and for the most part unknown in the plains
between the various links in the chain of mountains
in Southern Europe. We may picture these plants
as occupying islands surrounded on all sides by a sea
of Lowland vegetation.

We now pass on to compare the Swiss Alpine
flora with those of the plains to the north and south
of the Alps.

One of the most striking features of the vegetation
of Alpine Switzerland is its close affinity with that of
the Lowlands of temperate Western and Central
Europe, north of the Alps, especially with the floras
of Britain, France, and Germany.

We have seen (p. 205) that many British plants
are abundant in the Swiss Alpine region. Further,
the great majority of the species which there occur,
and which are not found in the plains to the north
of the Alps, belong to genera or families highly
characteristic of the latter region. This affinity is, if
anything, somewhat closer than that of the floras of
the Pyrenees and Tyrol to the same Western
European flora.

When, however, we compare the Swiss Alpine
flora with the subtropical Mediterranean flora to the
south of the Alps, the difference is exceedingly

ARCTIC SPECIES 299

striking. Comparatively few representatives of the
southern flora which is in its general aspect quite
distinct, are found in the Swiss Alpine region.
Among the few Mediterranean types in the Alps are
the Crocus (p. 153), the Globularias (p. 110), Biscutella
Icevigata, Linn, (a very common Cruciferous plant,
especially abundant on dry banks), the Mediterranean
Heath (Erica cornea, Linn.), and species of Hutchinsia.
Others occur in Tessin (Ticino), which is the only one
of the Swiss Cantons which lies wholly on the south
side of the Alps. Here Mediterranean types are
naturally more abundant, the climate, race, language,
and flora being, as we should expect, thoroughly
Italian.

The Swiss Alpine flora must next be compared
with that of the Arctic regions of Northern Europe.
It is a very remarkable fact that at least 30 per cent.,
or nearly one-third, of the Swiss Alpine species
flourish in the far northern frigid zone. Such
plants as Dryas octopetala, Linn. (p. 106), Ranunculus
glacialis, Linn. (p. 193), and Saxifraga oppositifolia,
Linn. (p. 186), are nearly as abundant in the Arctic
regions as in Switzerland, and no fewer than 130 other
species are common to both the floras. On the other
hand, several important Swiss genera, such as the
Gentians, are either absent or very poorly represented
in Arctic Europe.

The flora of temperate North America (United
States) is quite unlike that of Western Europe, yet
in the Kockies and other ranges we find a number of

300 THE ORIGIN OF THE FLORA

plants common in Alpine Switzerland, such as Caltha
palustris, Linn. (p. 205), Dryas octopetala, Linn,
(p. 106), Androsace chamcejasme, Jacq. (p. 74),
Pyrola uniflora, Linn. (p. 261), Primula farinosa, Linn,
(p. 69), Saocifraga oppositifolia, Linn. (p. 186), Juni-
perus communis, var. nana, Willd. (p. 233), and
Empetrum nigrum, Linn. (p. 231). These are
associated with a large number of plants unknown
to Europe either generically or specifically, such as
Claytonia and Gaillardia.

Thus we see that the distribution of the Swiss
Alpines beyond the Alps affords an exceedingly
interesting study.

THE ORIGIN OF THE Swiss ALPINE FLORA.

Any theory, which is to account adequately for
the origin of the Swiss Alpine flora, must explain the
facts of the present-day distribution and affinities of
these plants, which we have just passed under review.

The study of this problem has thus naturally two
sides : the historical and the geographical. We now
pass to consider the historical or geological facts.

The Alps, like the other mountain regions of
Europe, Asia, and North America, began to come
into existence as a mountain range during the
Oligocene period, and the uplifting of these and other
"massifs" continued during Miocene times. They
are thus quite modern structures in a geological
sense. The elevation of these vast areas was
probably comparatively rapid at some epochs, while

THE GLACIAL PERIOD 301

slow and gradual at others, and has possibly not yet
entirely ceased. During the next geological period,
the Pliocene, the mean temperature of Europe and
North America became gradually lowered by the
setting in of new climatic conditions, which eventually
culminated in the Great Ice Age.

The epoch during which this great event took
place is known as the Pleistocene or Glacial Period,
and according to one very probable view, we are at
present living in the latter part of this very period.

The Glacial Period no doubt did not set in
suddenly. The foci, so to speak, of the wave of
long-continued cold appear to have been located at
the Poles, both at the beginning, during the period of
maximum intensity, and during the last lingering
phase. The cold increased in intensity slowly and
gradually. As the temperature fell at the North
Pole, a wave of cold began to creep further and
further southward over the greater part of Central
Europe and North America. When the Ice Age
reached its maximum, a very large portion of the
British Isles, Central and Southern Europe, Canada
and the United States, became highly glaciated — that
is to say, these areas resembled Greenland, as it is
to-day. The Alps, and all the other mountain
regions of the Northern hemisphere, were almost
entirely clothed in a mantle of snow and ice extend-
ing to the foothills or almost to the plains.

In all probability the advance of the wave of low
temperature was exceedingly gradual, and several

302 THE ORIGIN OF THE FLORA

thousands of years may have elapsed between the
period when the temperature first began to fall at the
Pole and the glaciation of the British Isles.

There was also probably more than one period of
maximum low temperature. Cycles of severe cold,
each of considerable length, seem to have alternated
with more genial, Interglacial Periods, when the ice
retreated for a time from Britain and the plains of
Central Europe. Finally, the climate became per-
manently temperate, the ice retreated to the polar
circle, and to the higher peaks of the European
and American mountains, and thus was ushered in
the era of to-day.

The Glacial Period then was a long-continued
epoch, the changes of temperature being extremely
slow and gradual. Periods of cold alternated with
more genial epochs. As the tides ebb and flow,
so the wide- spread glaciation appears to have now
advanced, now retreated.

We will not attempt to discuss here the arguments
which have been put forward to account for the
occurrence of such glacial periods. These remain
the province of the mathematician, physicist, and
astronomer. We may, however, add that the late
Tertiary Glacial Period is not the only ice age of
which geological evidence exists. Nor will we
concern ourselves with the proofs of the general
glaciation of Europe and North America. As
Darwin, in the " Origin of Species" (p. 310), remarks :
" The ruins of a house burnt by fire do not tell their

THE ANCIENT ARCTIC FLORA 303

tale more plainly than do the mountains of Scotland
and Wales, with their scored flanks, polished surfaces,
and perched boulders, of the icy streams with which
their valleys were lately filled."

There is little doubt that some members of the
floras of Europe and North America, both of the
plains and the mountains, date back to a period prior
to the Ice Age. The history of both these floras
begins in the Miocene period.

We now turn to the explanation of the origin of
the Alpine flora of Europe, which is most generally
accepted, and examine how far it will explain the
present geographical distribution of Alpine plants,
and the historical facts as above outlined. This
theory was elaborated many years ago by Forbes,
Charles Darwin, Sir Joseph Hooker, Asa Gray, and
others. Briefly, it is as follows : — It is believed that
in Miocene and Pliocene times a flora existed in
Europe and North America, which was essentially
similar in both these regions. The Alps no doubt
also then possessed their Alpine flora. In the Far
North, however, there was another flora, which we
may speak of as the ancient Arctic flora. This,
then, was the state of affairs prior to the Ice Age.

With the setting in of glacial conditions at the
Pole, these floras were driven, each in their turn,
further and further southward. The ancient Arctic
flora retreated before the growing ice cap of the Arctic
regions, and migrated into North America (Canada,
and the United States) and into Europe. The

304 THE ORIGIN OF THE FLORA

temperate Miocene flora of North America was
driven, in its turn, towards the south, and there being
no mountain ranges stretched across the continent
to bar its way, it found a haven in what are now the
tropics, and in some cases, it is believed, even migrated
into the Southern hemisphere. In Europe, however,
the retreat of the Miocene flora was barred by
the Alps and other links in the incomplete chain of
mountains, which stretches from the west to the east
across the continent. The flora, brought up sharp
against the highly-glaciated Alps, became almost
entirely extinct. A few survivals, however, preserved
by some good fortune, still lingered.

With the advancing wave of cold, the Alps and
the Rockies became gradually more and more glaciated,
and the ancient Alpines were driven downwards to
the Lowlands, and there mingled with the ancient
Arctic plants from the Far North.

At the periods of maximum intensity of cold, the
plains of Europe and North America were populated
by Arctic and Alpine species.

When, however, the ice again retreated northward,
many of the ancient Arctic plants returned to the
polar region and the old temperate, Miocene types
to North America, and there mingled with some
Arctic species which finally made their homes in the
Lowlands.

In Europe, as we have seen, the Miocene flora
had been practically exterminated, and the present
Western European flora would seem to have originated

DIFFICULTIES AS TO THE THEORY 305

from some of the ancient Arctic plants, which became
established in the plains. Both in Europe and North
America, as the glaciation decreased, some of the
ancient Arctic, as well as the ancient Alpine species,
ascended the mountains, and gave rise eventually to
the modern Alpine floras of these continents.

Such is the bare outline of the theory which has
been advanced to account for the origin of the Alpine
flora. It is quite impossible to discuss the evidence
for, or against, it here. At any rate, it will explain
the facts of the present-day distribution of Swiss
Alpine plants, which we have already enumerated. It
will explain why some of the Alpines of Europe and
North America are identical, while the Lowland floras
of these regions are very dissimilar. It will explain
the isolated seclusion of Alpines on the various
detached mountain chains of Europe and elsewhere
without any appeal to the theory of multiple centres
of origin of species. It furnishes a reasonable historic
connection between the facts of the past and the
present.

At the same time, it must be pointed out that
difficulties exist, either as to the theory as a whole,
or as to some of its essential components, which have
led such authorities as Dr Christ and the late John
Ball to dissent from it. The relationship between
the present Alpine and Arctic floras is the chief
stumbling block. A large number, in fact the majority,
of Swiss Alpines do not occur in the Arctic. It has
been urged that Central Asia was the original home

306 THE ORIGIN OF THE FLORA

of the Alpine flora. It has been pointed out that,
even if the Alpine flora has been derived from an
Arctic flora, which, according to the theory, existed
in the Far North before the incoming of the Glacial
Period, we have still to explain the origin of this
ancient flora, and above all, why at that time it
possessed an Arctic character !

The recent experimental work of Prof. Bonnier,
to which we have more than once alluded, has a
profound significance for those who are inclined, in
part at any rate, to distrust the theory. Bonnier has
shown that the conditions which prevail in the Arctic
regions are by no means identical with those in the
High Alps, and, what is even more important, that a
plant grown in the Arctic region differs from one
grown in the Alps. It is thus clear that the effect of
physical conditions alone is greater than was formerly
supposed.

Until, however, we know not only how one species
is derived from another, but which particular genera
and species have given rise to other genera and
species, it is not probable that we shall be able to
settle the matter finally. Was Eritrichium nanum
derived from a species of Myosotis? Did Anemone
alpina originate from Anemone pulsatilla, or perhaps
A. vernalis? These are questions to which at present
there is no answer ; and until we can decide such
matters, we shall not see clearly on the subject of
the origin of the Alpine flora of Switzerland.

APPENDIX I
GLOSSARY OF BOTANICAL TERMS

Absorption, the act of imbibing a liquid or a gas (see p. 10).
Achene, a dry, one-seeded, indehiscent fruit — e.g., the individual

fruits, commonly miscalled "seeds," of a Strawberry or

Buttercup (p. 285).
Actinomorphic, the term used to describe a flower which can

be divided into two symmetrical halves by two or more

planes of symmetry, as opposed to Zygomorphic (q.v.).
Acyclic, a term used to describe a flower whose parts are

arranged spirally instead of in whorls ; cf. Cyclic.
Adhesion, the union of dissimilar parts of the flower, such as

the petals and sepals ; cf. Cohesion.
Adventitious Roots, those which arise as outgrowths from the

stem or leaves, and are not branches of the main root.
Alkaloid, a general term applied to organic bases, which occur

in many plants ; cf. Aconitine (p. 130).
Alternate, applied to leaves which are arranged spirally on

the stem, and are not opposite to one another. Also

known as scattered.
Analogous, having the same function, but not necessarily the

same structure ; cf. Homologous.
Androecium, the male organs or stamens of a flower, considered

as a whole.

807

308 APPENDIX I

Anemophilous, a term applied to flowers which are cross-
pollinated by the agency of wind ; cf. Entomophilous.

Angiosperm, a Flowering Plant whose ovules are enclosed in
ovaries. The term is used in opposition to Gymnosperm

(?.».).

Annual, a term used to describe plants which pass through
their whole life-history in one year, and then die.

Anther, the upper portion of the stamen consisting of the
pollen-sacs and the tissue between them.

Apetalous, without petals, or with very small rudimentary
petals — e.g., the flowers of the Willow.

Apocarpous, applied to the gynaeceum of a flower in which all
the carpels are free from one another — e.g., the Marsh
Marigold (Caltha).

Aquatic, living in water.

Asexual, a term applied to reproduction by means of organs
other than the stamens and carpels — e.g., Bulbils of
Polygonum viviparum (p. 156).

Assimilation, the process by which the plant converts raw food
material into its own substance. The term is used especi-
ally in reference to Carbon Assimilation — that is to say,
the formation of organic substance from carbon dioxide
and water by green plants in sunlight (p. 10).

Association, a community of plants living together (p. 32).

Awn, a prolongation of an organ, usually thread-like ; cf. the
awned fruits of Anemone alpina (p. 39).

Axil, the angle formed, on the upper side, at the point of
attachment of a lateral organ to the main organ — e.g., the
angle above the attachment of a leaf to the stem.

Axis, a term generally used to imply the stem. The root may
also be termed an axis. Such organs as leaves are
appendages of the axis.

Berry, an indehiscent fruit, containing several seeds surrounded
by a juicy or fleshy pulp — e.g.9 the Bilberry and Gooseberry.

GLOSSARY OF BOTANICAL TERMS 309

Biennial, a plant which requires two years to complete its life-
history. In the second year it produces flowers and fruit,

and then dies.

Bisexual = Hermaphrodite (q.v.).
Bract, the leaf or modified leaf on the axis of an inflorescence,

in the axil of which the flower arises. (The bracts are

often entirely suppressed, as in the Crucifer family.)
Bracteole, the leaf, or one of the leaves, borne on the axis of

the flower itself, above the bract and below the flower.

(Bracteoles are often entirely absent) (p. 225).
Bud, a short stem axis, crowded with overlapping, young,

undeveloped leaves, arising close to one another.
Bulb, a modified stem, often subterranean and disc-like in form,

bearing a number of succulent leaves, containing reserve

materials — e.g., an Onion or a Lily bulb.
Bulbil, a deciduous bud, capable of reproducing the species, and

often containing reserve materials (p. 156).
Calcicole, applied to plants which flourish best on calcareous

(limestone) soils (p. 117).
Calcifuge, applied to plants which will not grow on calcareous

(limestone) soils (p. 117).
Calyx, the sepals of a flower considered as a whole ; the outer

series of a differentiated floral envelope (generally green).
Capitulum, or Head, a type of inflorescence in which all the

flowers are stalk less, and arranged on a terminal expansion

of the axis — e.g.-) the inflorescence of the Composites.
Capsule, a many-seeded dry fruit, composed of two or more

carpels, which open in various ways to allow the seeds to

escape (p. 88).
Carpel, the modified leaf, bearing and enclosing the ovules.

The pistil consists of one or several carpels, which may

be free or united together.
Catkin, a close spike of unisexual apetalous flowers, which may

be shed as a whole — e.g., Willow Catkins (p. 189).

310 APPENDIX I

Cells, the ultimate units of which plant tissues are built up

(p. 8).

Cell Sap, the watery contents of the cell.
Chalk-gland, a glandular tissue secreting water and calcium

carbonate (p. 76).
Chlorophyll, the green colouring matter of plants. It is

generally localised in special granules within the cells,

known as chlorophyll corpuscles or Chloroplasts (p. 10).
Chromoplasts, special portions of the protoplasm containing

colouring matter (p. 45).
Claw, the narrowed base of the petal.

Cleistogamous, flowers which never open, and which are self-
fertilised (p. 278).
Cohesion, the union of similar parts of the flower, such as the

union of all the petals ; cf. Adhesion.
Cone, as applied to the Coniferae, the fertile shoots, bearing

spirally arranged scales subtending carpels or bearing the

pollen sacs (p. 235).
Coniferae, the group of plants, including the Pines and Firs,

which bear fertile shoots or cones (q.v.) of a special type.
Connective, the part of the stalk or filament of the stamen

connecting the two pairs of pollen sacs.
Contractile Boots, special roots which pull the stem down into

the soil (p. 122).
Corm, or solid bulb, is a modified swollen underground stem

which serves as a storehouse for reserve food material. It

is often covered externally with scale-leaves (pp. 153, 166).
Corolla, the petals of a flower considered as a whole ; the inner

series of a differentiated floral envelope, usually white or

coloured.
Corona, an outgrowth from the corolla or perianth — e.g., the

" trumpet " of a Daffodil.
Cotyledon, the first or one of the first pair of seed-leaves

produced by a young seedling plant (p. 83).

GLOSSARY OF BOTANICAL TERMS 311

Cross-fertilisation, the act of impregnation of the egg or ovum
of one flower by the male gamete (q.v.) of another
flower (pp. 52, 268).

Cross-pollination, the dusting of the stigma of one flower by
the pollen brought from another (p. 268).

Crustaceous, forming a crust, closely adherent to the sub-
stratum— e.g., Crustaceous Lichens (p. 292).

Cyclic, a term applied to leaves or parts of a flower arranged in
whorls; cf. Acyclic.

Cyme, a form of inflorescence in which the main axis ends in a
flower, and the succeeding flowers are produced on succes-
sive, lateral axes. A cyme is also called a centrifugal
inflorescence, because the oldest flower is placed centrally
— e.g.9 Lychnis (p. 225).

Deciduous, applied to those plants whose leaves do not persist
from year to year (cf. Evergreen), but are shed each
autumn.

Decurrent, applied to leaves which are prolonged downwards
on the axis — e.g., leaves of Thistles.

Dehiscent, opening at one or more points so as to allow the
contents to escape. Applied to fruits and pollen sacs.

Dichasium, a form of cyme, in which two lateral branches of
nearly equal strength arise below the flower which
terminates the main axis. These lateral branches may
each again give rise to two branches, and so on.

Dichogamous, a term applied to flowers in which the stamens
and carpels ripen at different times, and thus self-pollina-
tion is prevented.

Dicotyledon, a plant having two seed-leaves. The Dicoty-
ledons form the majority of the flowering plants; cf.
Monocotyledon.

Dioecious, unisexual male and female flowers borne on different
plants (p. 189).

Dimorphism = Heterostylism, flowers of two or more forms

312 APPENDIX I

differing in the relative lengths of the stamens and styles

— e.g.) Primula^ Ly thrum (p. 68).
Disc-florets, the flowers occurring in the central portion of the

capitulum of the Composite ; cf. Ray Florets.
Drupe, a fleshy, indehiscent fruit, containing one or more seeds.

The outer fruit-coat is fleshy or pulpy, the inner is hard

and stony — e.g., a Cherry and a Peach (pp. 231-2).
Bbracteate, without bracts (q.v.).
Ecology, or (Ecology, the study of plant life in relation to its

environment.
Emergence, an outgrowth from the surface tissues of a plant

organ, such as a prickle on a rose stem.
Endosperm, the store of food material laid up in certain seeds

outside the embryo. The embryo absorbs it during its

germination — e.g., the horny part of a grain of maize.
Entire, applied to leaves the margins of which are not

toothed.

Entomophilous, a term applied to flowers which are cross-
pollinated by the agency of insects ; cf. Anemophilous.
Epicalyx, the outer series of a double calyx — e.g., in Potentilla,

Dryas, Fragaria (pp. 128, 158).
Epigynous, applied to flowers in which the calyx tube

completely encloses the ovary, and the corolla and stamens

appear to be placed on the top of the ovary.
Epipetalous, applied to stamens borne upon petals (p. 64).
Epiphyte, a plant which grows upon another plant, but is not

a parasite (p. 291).
Etiolation, the condition of a plant which has been grown in

absence of sunlight. The stems are long and weak, the

leaves small, and the colour is yellowish-white instead of

green.
Evergreen, a term applied to leaves which last for more than

one season, and to plants bearing green foliage all the

year (p. 25).

GLOSSARY OF BOTANICAL TERMS 313

Exalbuminous, applied to seeds which have no endosperm, and

in which the embryo occupies the whole cavity of the

seed.

Exstipulate, possessing no stipules.
Extra-floral, outside the flower,; a term applied to some

nectaries which are situated on leaf-stalks, etc., instead of

in their usual position within the flower (p. 276).
Extrorse, applied to anthers which are so turned that they

open outwards, away from the centre of the flower.
Family = Natural Order (q.v .).
Ferment, a substance in the plant which produces chemical

changes, without itself contributing to the resulting

products (p. 214).
Filament, the stalk of a stamen.
Flaccid, limp, flabby, as opposed to turgid (q.v.).
Floral envelope, the modified leaves surrounding the stamens

or carpels, or both, in a flower, and placed below them.

It may be undifferentiated (a perianth) or differentiated

into calyx and corolla.
Floral leaf, a leaf modified to form one of the parts of a flower

— e.g., a sepal, a petal, a stamen, or a carpel.
Flower, a shoot bearing modified leaves devoted to sexual

reproduction. The flower may consist of stamens or

carpels alone, or both, with or without a floral envelope.

In a hermaphrodite flower the stamens are always placed

below the carpels.

Follicle, a dehiscent fruit, composed of a single carpel contain-
ing several seeds.
Fruit, the ripened carpel or carpels of a single flower, enclosing

one or more seeds.
Function, the part or role performed by any organ — e.g.,

reproduction is the function of the flower.
Gall, an abnormal growth, caused by an insect or a fungus

(p. 290).

314 APPENDIX I

Gamete, the sexual unit, male or female, consisting of a

naked mass of protoplasm, motile or non-motile.
Gamopetalous, applied to flowers in which the petals are all

united together ; cf. Polypetalous.
Genus (plural Genera), a group or collection of nearly related

species, possessing certain characters in common by which

they are distinguished from other groups or genera

(pp. 15, 249).

Geophilous ; see Geophyte.
Geophyte, a plant which develops its aerial organs more or

less completely beneath the surface of the soil (see p. 125).
Germination, the first act of growth of a seed.
Glabrous, without hairs.

Gland, a definite secreting organ, usually superficial (p. 212).
Glandular Hair, a hair with an enlarged apex, containing a

secretion (pp. 72, 99).
Globose, spherical.
Gymnosperm, a flowering plant whose ovules are not enclosed

in carpels ; cf. Angiosperm. The Gymnosperms include

the Coniferae (q.v.).
Gynaeceum, the carpels or female organs of a flower, considered

as a whole.

Habit, the external form of the plant, its shape or build.
Habitat, the particular kind of locality in which a plant

flourishes — e.g., a marsh, a forest.
Head of Flowers = Oapitulum (q.v.).
Herbaceous, not woody.
Hermaphrodite, applied to flowers which possess both male

and female organs.
Heterostylism ; see Dimorphism.
Holophyte, a plant which obtains all its own nourishment

itself, and does not live parasitically or saprophytically.
Homologous, having the same type of structure, but not

necessarily the same function ; cf. Analogous.

GLOSSARY OF BOTANICAL TERMS 315

Honey, or Nectar, the substance secreted by many flowers to

attract insects.

Honey-glands, the organs which secrete honey.
Humus, decomposing organic matter in the soil (pp. 119, 143).
Hybrid, a cross between two species or races (p. 70).
Hypogynous, applied to flowers in which the calyx and corolla arise

directly from the receptacle and in which the ovary is superior.
Imbricated, overlapping, like the tiles on a roof.
Indehiscent, applied to fruits which do not open to allow the

seeds to escape ; cf. Dehiscent.
Inferior, applied to the ovaries of those flowers in which the

calyx tube encloses the ovary, as in epigynous and some

perigynous flowers ; cf. Superior.
Inflorescence, the mode of branching of the floral axis; the

manner in which the flowers are arranged on the primary

and lateral shoots.
Insectivorous, applied to plants which capture insects and

absorb nutriment from them.
Internode, the portion of the axis between the insertion of two

successive leaves (p. 191).
Introrse, applied to stamens in which the anthers open inwards,

towards the centre of the flower.
Involucre, the whorl or rosette of bracts below an inflorescence

or a single flower (see pp. 35, 111).
Irregular = Zygomorphic (q.v.).
Keel, a term applied to the two lower, united petals of the

flowers of members of the Leguminosse.
Labellum, or Lower Lip, applied to the enlarged and

irregularly shaped member of the inner whorl of perianth

members in the Orchids (p. 137), and to a similarly placed

petal in other flowers.
Lamina, the blade of a leaf.
Lichens, a group of thalloid plants, consisting of Algae

associated with Fungi (see p. 291).

816 APPENDIX I

Loculus, the cavity of an ovary or anther.

Monocotyledon, a plant having only one seed-leaf. Lilies,

Crocuses, and many other bulbous and tuberous plants,

with the Grasses and Rushes, come under this heading;

cf. Dicotyledon.
Monoecious, applied to plants which bear unisexual male and

female flowers on the same plant.
Natural Order, or Family, a collection or group of nearly

related genera, possessing certain characters in common,

whereby they can be distinguished from other groups.

Sometimes the order contains only one genus.
Nectar = Honey (q.v.).

Nectary, an organ secreting honey or nectar (pp. 272, 273).
Nitrogenous, containing nitrogen.

Node, the point of insertion of a leaf on an axis (p. 191).
Nut, a dry, indehiscent fruit with a woody pericarp — e.g.,

Hazelnut.

Offset, the budjformed at the end of a runner or stolon (p. 100).
Opposite, applied to leaves which are arranged in pairs at the

same level on the stem ; cf. Alternate.
Organ, a part of a plant which serves a definite function — e.g.,

a leaf, a stamen.
Ovary, the female portion of the flower, consisting of one or

more closed carpels, enclosing one or more ovules.
Ovule, the female organ enclosed in the ovary, which when

fertilised becomes a seed.
Panicle, a branched raceme (q.v .).
Pappus, the hairs or scales developed at the summit of the

fruits of many Compositae. Derived originally from the

calyx — e.g., Dandelion fruits.
Parasite, a plant living in or on another plant (the host), from

which it derives part or all of its food-supply (p. 287).
Parthenogenesis, the development of an embryo from an ovule

without fertilisation (p. 279).

GLOSSARY OF BOTANICAL TERMS 317

Pedicel, the stalk of a single flower of an inflorescence.
Peduncle, the stalk of an inflorescence or of a solitary flower.
Perennial, a plant which lives for more than two years and

does not perish after producing flowers and fruit, but

flowers again the succeeding season ; cf. Biennial and

Annual.
Perianth, a floral envelope, not differentiated into two distinct

series, calyx and corolla.

Pericarp, the wall of the ovary as developed in the fruit.
Perigynous, applied to flowers where the calyx tube surrounds,

but does not enclose, the ovary.
Petal, a unit of the corolla, or inner series of a differentiated

floral envelope.
Petiole, a leaf stalk.
Phanerogam, a Flowering Plant ; a wide term, including the

Gymnosperms and Angiosperms.
Phyllotaxis, the plan of arrangement of the leaves on the

stem.

Pistil = Gynaeceum (q.v.).
Pollen, the yellow dust formed in the anthers of the stamens ;

the male spores which produce the male gametes.
Pollen-grains = Pollen.
Pollen sac, the sporangium or sac in which the pollen-grains

are formed in the anther.
Pollination, the act of dusting the stigmatic surface of the

pistil with pollen.

Pollinium, the pollen mass of an Orchid (p. 138).
Polygamous, bearing hermaphrodite and unisexual flowers on

the same or different individuals of the same species.
Polypetalous, applied to flowers in which the petals are free

from one another ; cf. Gamopetalous.
Prickle, a pointed Emergence (q.v.).
Protandry, the form of dichogamy, in which the male organs

(andrcecium) mature before the female (gynaeceum).

318 APPENDIX I

Protogyny, the form of dichogamy in which the female organs

(gynaeceum) mature before the male (andrcecium).
Protoplasm, the viscous or jelly-like substance, which forms

the essential part of the "cells'" or individual units of

which all living creatures, both plants and animals, are

built up (p. 9).
Raceme, a simple inflorescence in which stalked flowers are

borne laterally on a central axis, the oldest flowers

being at the base.

Rachis, the axis of a compound leaf.
Ray Florets, the outer flowers of the capitulum of the

Compositae, which often differ from the Disc Florets

(?•»•)•

Receptacle, the portion of the axis of the flower which bears
the floral envelope, and the male and female organs ; the
axis bearing the florets in the Compositae (p. 18).

Regular = Actinomorphic (q.v.).

Reserves, food material stored in the plant for future use

Respiration, the gaseous exchange between a plant and the

atmosphere, corresponding to the breathing of animals.

Oxygen is absorbed and carbon dioxide given out (p. 11).
Rhizome, a creeping or prostrate, subterranean stem, bearing

erect, leafy shoots.
Root-stock = Rhizome (q.v.).
Rosette, applied to a group of leaves, arranged in a very close

spiral, the internodes between them being very short

(p. 75).
Runner, a slender, prostrate stem-branch, usually rooting

(p. 100).
Saprophyte, a plant deriving its nourishment from decaying

organic matter (p. 287).
Scale, a disc-like outgrowth, usually of superficial origin,

especially on leaves (p. 26).

GLOSSARY OF BOTANICAL TERMS 319

Scale-leaves, reduced leaves which have generally only a

protective (p. 167) or storage function.
Scape, a floral axis arising from the ground, and terminating in

a single flower — e.g., Daffodil.
Schizocarp, a splitting fruit, in which the pericarp divides into

two or more one-seeded portions.
Seed, a fertilised ovule.
Seed-coat, a covering of the seed, derived from one or more

integuments of the ovule (pp. 265, 266).
Selective Capacity, the power possessed by roots of selecting

certain special substances from among those in solution in

the soil (p. 77).
Self-pollination, the act of placing the pollen of one flower on

the stigma of the same flower. This results in self-
fertilisation (pp. 52, 278).
Semiparasite, a plant deriving part of its nutriment from the

tissues of another plant ; cf. Parasite (p. 218).
Sepal, a unit of the calyx, or outer series of a differentiated

floral envelope.
Sessile, stalkless.
Shade-leaves, leaves growing under conditions of poor

illumination (pp. 84, 247, 253).

Shrub, a woody perennial, without an erect or main trunk.
Spadix, a fleshy spike,

Spathe, a sheath-like leaf, enveloping an inflorescence.
Species (plural Species), a classificatory unit ; a unit of a

genus ; one sort or kind of plant (pp. 15, 249).
Sperms = male gametes (q.v .).

Spike, a simple inflorescence, in which the flowers are stalk-
less and attached directly to a central axis ; a sessile

raceme.

Spiral = Acyclic (q.v.).
Sporangium, a sac containing spores.
Spur, a prolongation of a perianth member, usually tubular.

320 APPENDIX I

Stamen, an individual member of the androecium ; the male

leaf of a flower.
Staminode, a modified stamen, no longer pollen producing ; a

reduced stamen (p. 217).
Standard, a term applied to the large, upper petal of a

Leguminous flower.
Stigma, the receptive portion of the pistil, on which the pollen

is deposited.

Stipulate, possessing stipules.
Stipules, the paired appendages (often leaf-like) occurring at

the base of some leaves.
Stolon = Runner (q.v .). .-
Stoma (plural Stomata), a pore or minute opening in the

outer layer (epidermis) of a leaf (p. 10).
Style, the upper portion or portions of the pistil, bearing the

stigma or stigmas.
Sun-leaves, leaves growing under conditions of strong

illumination (p. 84).
Superior, a term applied to an ovary which is free from, or not

enclosed by, the floral envelope ; cf. Inferior.
Symbiosis, the living together of two dissimilar organisms:

either two plants or animals, or a plant and an animal

(p. 293).

Sympetalous = Gamopetalous (q.v.).
Syncarpous, applied to the gynseceum of a flower, in which the

carpels are united together to form a single ovary.
Teeth, minute, pointed lobes on the margins of leaves.
Tendril, a special organ, usually thread-like, adapted for

climbing.
Thallus, a vegetative body without differentiation into stem

and leaf.
Transpiration, the act of giving off water from the leaves of a

plant. The water passes off in the form of vapour, through

the pores or stomata (p. 12).

GLOSSARY OF BOTANICAL TERMS 321

Tuber, a short, thick, underground stem, stored with food

material.

Turgid, distended with sap.
Umbel, an inflorescence in which the flowers are stalked, the

stalks all radiating from a common point on the axis

(p. 42).
Unisexual, applied to flowers which contain either male or

female organs, but not both (p. 39).
Variation, a departure from type (p. 47).
Versatile, applied to anthers which are balanced on the

filament, and so swing freely.
Viviparous, germinating while attached to the parent plant

(p. 157).
Water-stomata, special pores on the leaves of some plants

which exude water (p. 76).
Whorled, applied to foliar and floral leaves, which are arranged

in a circle, at one level on the axis.
Wing, a prolongation of a seed or fruit ; the lateral petal of a

Leguminous flower.
Zygomorpbic, applied to flowers, which are not divisible into

two similar halves in more than one plane of symmetry.

APPENDIX II
THE STRUCTURE OF THE FLOWER

THE botanist's idea of what constitutes a flower is not exactly
the same as that implied in our common speech. Even small,
green, inconspicuous objects, such as the little structures which
go to make up a Willow catkin, come under the botanical
category of flowers, though they do not possess the brightly-
coloured, expanded petals which are ordinarily associated
with the word. The definition of a flower, from a botanical
standpoint, is that it is a shoot, bearing leaves specialised for
purposes of reproduction. The shoot is, as it were, telescoped,
so that the floral leaves, instead of being separated from one
another, like the leaves on an ordinary branch, are closely
crowded together. This is not peculiar to flowers alone.
The same thing happens in the crowded leaf rosettes of
Sempervivum (Plate XXIV., Fig. 1) and other plants. The
floral leaves are, however, not merely crowded on the
shortened axis or receptacle, but they are also usually arranged
in successive circles, known as whorls, instead of being in a
continuous, spiral series.

Let us now examine the floral leaves, and notice how they
are modified for the ultimate purpose of setting seed, which
will reproduce the plant. Starting from the outside of the
flower — in other words, from the base of the floral shoot — we
first meet with a group of leaves, the perianth, which have

822

THE STRUCTURE OF THE FLOWER 323

nothing directly to do with reproduction, but at the same time
are of great indirect value in connection with it. The perianth
leaves may be all alike (cf. Crocus, Text-fig. XXX.), or they
may be differentiated into two series : an outer, known as the
calyx, and an inner, the corolla (cf. Buttercup, Text-fig. XXVI.).

The individual members of the calyx are called sepals.
They are commonly green, and more nearly resemble ordinary
foliage leaves than do any other of the parts of the flower.
In some cases, as in the Rose, they may be remarkably leaf-like.
The purpose which they serve in the economy of the flower is
that of protection. In the bud, they commonly enclose all
the other floral parts. The corolla consists of petals, which
usually differ from the sepals in being larger and more delicate
in texture, and either white or gaily coloured, instead of green.
Like the sepals, they enclose and protect the rest of the flower,
but in a very large number of cases they have another and more
special function — namely, that of attracting insects to visit the
flower, for a purpose to which we will return later.

Immediately within the petals, we meet with the first of the
" essential organs " of the flower — namely, the group of stamens
constituting the androecium. Each stamen consists of a
slender stalk or filament, terminating in a head or anther
of four little sacs, two on each side of the connective, as
the top of the filament is called. The stamen, like the other
parts of the flower, may be regarded as a metamorphosed leaf.
In some flowers, such as that of the White Waterlily, the
leafy nature of the stamens is particularly obvious. The
object of the stamen's existence is the production of the yellow
dust, known as pollen, with which the anther sacs are filled.
When the stamens are ripe, the anthers open in various ways,
and the pollen escapes. The stamens are the male organs,
and the pollen gives rise to the male gametes or sperms, without
whose aid no seeds can be set. We will return to the subject
of the fate of the pollen after considering the pistil.

324 APPENDIX II

The central structure of the flower is the female organ,
which is known as the pistil or gynocceum. As the androecium
is made up of stamens, so the gynseceum is made up of one or
more carpels, either free or more often fused together. Each
carpel may be regarded as essentially a leaf, bearing immature
seeds, or ovules as they are called, along its margin. Its nature
is, however, obscured by the fact that the edges of the leaf
are fused together, so as to enclose the ovules. The structure
of a pistil, consisting of a single carpel, can be best understood
by splitting open a pea-pod, down the edge along which the
peas are attached. The pod, when opened out, is seen to be
only slightly changed from a leaf folded along its midrib.

The part of the carpel which contains the ovules is called
the ovary. It is prolonged upwards, as a columnar or thread-
like outgrowth, called the style, which terminates in a special
portion intended for the reception of the pollen grains, and
known as the stigma. The stigma is frequently hairy or sticky,
so as more easily to catch and retain the pollen grains.

A better crop of seeds is usually produced when pollen
from another flower is deposited on the stigma — or in other
words, when cross-fertilisation, and not self-fertilisation, takes
place. Cross-pollination is brought about either by the
arrival of pollen blown by the wind, or brought by insect
visitors which, attracted by the coloured perianth or by a sweet
scent and a prospect of honey, fly from flower to flower, and
thus unintentionally convey the pollen of one flower to the
stigma of another.

When the pollen-grain reaches the receptive surface of the
stigma, it grows out into a long slender tube (only visible
under the microscope), which travels down the tissues of the
style into the cavity of the ovary, and, advancing towards one
of the ovules, enters it by a tiny aperture in its outer coat.
The male fertilising element, or sperm, passes down the tube,
enters the ovule, and fuses with a cell within it known as the

THE STRUCTURE OF THE FLOWER 325

egg-cell. The next generation may be said to begin with the
fertilised egg-cell, which develops into a plant embryo.

The form of the embryo, at the stage when it is ready to
begin independent life, may be seen by examining a ripe pea.
Within the seed-coat we find two thick, whitish bodies, which
occupy nearly the whole of the interior and are known as the
cotyledons. Between them we can see a tiny root {the radicle),
and a bud which will form the leafy shoot (the plumule). The
cotyledons are really the two first leaves of the young plant,
which have given up the usual form and appearance of leaves
in order to act as storehouses of food material for the young
plant to draw upon before it has expanded its green leaves and
can nourish itself independently.

In other cases, such as the wheat and the coffee-bean, the
embryo does not occupy the whole interior of the seed, but lies
more or less embedded in a store of food material which is
known as the endosperm. The cotyledons do not in these cases
store the food themselves, but they have the power of sucking
it out of the endosperm.

The Flowering Plants are divided into two main groups, the
Dicotyledons and Monocotyledons, according as their seedlings
have two seed leaves, or only one. These two groups also differ
in floral characters. For instance, the dicotyledonous flower
has, as a rule, four or five parts in each whorl, and the perianth
is often differentiated into calyx and corolla, while the
monocotyledonous flower has its parts in threes, and the
perianth is undifferentiated.

We must now return to our description of flower structure,
and consider some of the chief modifications which the
general type may undergo. A typical flower contains
both gynseceum and androecium, and is called hermaphrodite.
But it is possible to have unisexual flowers, which may be
either wholly male, or wholly female. This is brought about
by the suppression either of the gynseceum or androecium

326 APPENDIX II

(cf. Veratrum, p. 123, and Dianthus, p. 113). Sometimes both
male and female flowers occur on the same plant, which is then
termed monoecious. In other cases, specialisation is carried
still further, one plant bearing only male, and another only
female, flowers. Such plants are described as dioecious.

Apart from the actual loss of one or other of the " essential
organs," there are several other directions in which flowers have
become modified. One of these is the union of similar members.
For instance, when the sepals or the petals are free from one
another, the flowers are called polysepalous or polypetalous,
but these members may unite among themselves, and then the
flowers are termed gamosepalous or gamopetalous. The stamens,
again, may unite by their filaments into a single group, when
they are called monadelphous, or into several groups, di-, tri-,
or polyadelphous. If they unite by their anthers, they are
spoken of as syngenesious.

In describing the gynseceum, we took a pea-pod as a typical
illustration of a carpel. Many gynaecea, however, consist of
more than one carpel. These carpels may either be free, as in
the Buttercup (Text-fig. XXVL), or fused together more or less
completely. In the Saxifrage (Text-fig. XXVIIL), the two
carpels are united in the ovary region, but the styles and
stigmas are free ; in the Crocus (Text-fig. XXX.), the fusion has
gone further, and only the three stigmas are free ; while in the
Primula (Text-fig. V.), in which there are five carpels, they
are completely fused throughout. A gynaeceum consisting of
fused carpels is spoken of as syncarpous.

Another way in which the flower may become modified is
through the union of dissimilar members. For instance, the
stamens, instead of growing freely from the receptacle, are
sometimes united for the greater part of their length with
the corolla (cf. Primula, Text-fig. V.). A more profound
modification is brought about by the hollowing out of the top
of the receptacle, and its union with the lower part of the

THE STRUCTURE OF THE FLOWER 327

calyx. This produces the calyx cup, which is very well seen in
the Rose. The gynaeceum is enclosed in the calyx cup, or, as
it is sometimes called, the receptacular cup, and the stamens,
petals, and free parts of the sepals arise from the edge of it.
This kind of flower is called perigynous, while the simpler type,
in which the receptacle is more or less conical, is called
hypogynous (cf. Buttercup, Text-fig. XXVI.).

As a further development, the gynaeceum may fuse
completely with the calyx tube, so that all the floral parts
appear to grow on the top of the ovary (cf. Crocus, Text-
fig. XXX., and Groundsel, Text-fig. XXIX). The flower is
then said to be epigynous^ and the ovary inferior.

Besides the floral parts which we have mentioned, special
structures called nectaries occur in some flowers. These take
various forms, but agree in one essential feature — namely, that
they secrete honey. In many flowers the nectaries are modified
stamens, as in the Globe-flower (p. 207), where they are horn-
shaped structures. In this case the modified stamen both secretes
the honey, and forms a receptacle to hold it. In the Pansy, on the
other hand (Text-fig. XV.), processes from two of the stamens
secrete the honey, which drops into a spur-like receptacle,
hollowed out of the base of the front petal. In the Lesser
Butterfly Orchid (Text-fig. XI., 1), one of the perianth members
is spurred like the Pansy ; but here it is the tissue of the spur
itself which secretes the honey. In the Buttercup (Text -fig.
XXVI., 2), honey oozes out from a patch of glandular tissue
concealed behind a little scale at the base of the petal.

Starting from such a flower as the Buttercup, there seem to
be several lines along which evolution may proceed. One of
the most important steps involves the union of the petals — that
is to say, polypetaly is replaced by gamopetaly. On the other
hand, the plant, instead of becoming more highly evolved in
regard to the corolla, which aims at the attraction of insects,
may decide to dispense with it altogether, or to reduce it to

328 APPENDIX II

a merely protective structure. In so doing, it must give up the
idea of insect-pollination in favour of self-pollination or wind-
pollination.

These distinctions are made use of in classification. The
Dicotyledons fall into three classes, known as the Polypetalce,
Gamopetdlce and Apetalce, according to whether their petals are
free, united, or absent. Within each of these groups we find
another tendency operating — namely, to proceed from a
hypogynous form of flower to a perigynous, and eventually to
an epigynous type.

We may illustrate some of the chief varieties of floral

FIG. XXVI.

1. The Flower of a Buttercup (natural order Ranunculaceae).

2. Petal of a Buttercup, showing the Honey-gland at the base.

structure by reference to five flowers, of which text-figures are
given here. Each diagram represents the flower cut in half
longitudinaUy. The Buttercup (Text-fig. XXVI.) is a
hypogynous flower, polysepcdous and polypetdlous. Both the
stamens and carpels are numerous, and free from one another.
Each carpel contains a single seed. The Buttercup is called a
regular flower, because all the parts of each whorl are alike in
shape and size.

The next text-figure (XXVII.) represents a Pea flower, in
which the five petals differ amongst themselves in size and

THE STRUCTURE OF THE FLOWER

329

shape, giving an irregular flower, which is symmetrical about
one plane only — namely, that seen in the figure. There are ten
stamens, nine of which are united, while one remains free, and

FIG. XXVII.— The Flower of a Pea (natural order Leguminosse).
thus the androecium is di-adelphous. The result of one stamen
being free is to leave a slit in the tube formed by the filaments,
through which the insect visitor can pass its proboscis when

FIG. XXVIII.— The Flower of a Saxifrage (natural order Saxifragaceee).
seeking for honey. The gynocceum, as we have already mentioned,
is monocarpellary (see also p. 252).

330 APPENDIX II

In the Saxifrage (Text-fig. XXVIII.) we have an example
of an incompletely epigynous flower. The petals and stamens
grow from the edge of the calyx tube, which is fused with the
lower part of the gynceceum. The gynccceum consists of two
carpels, not unlike two short pea-pods fused together.

The Groundsel (Text-fig. XXIX.) belongs to a more

FIG. XXIX. —The Head of Flowers of the Groundsel, Senecio (natural
order Composite). Enlarged.

advanced type, in which a number of tiny flowers orjlorets are
all crowded together into a head or capitulum, surrounded by
an involucre of bracts. Thus we have an inflorescence or
collection of flowers simulating a single flower. The Groundsel
is an example of the natural order Composites, to which the
Edelweiss (Plate L, Fig. 1) also belongs. The flowers are all
epigynous. The calyx is represented by hairs, which, later on,

THE STRUCTURE OF THE FLOWER

331

make it easy for the wind to waft the fruits along and
so distribute them in new situations. There are two types of
floret, the outer or ray florets, which have strap-shaped corollas
arid no stamens, and the hermaphrodite disc florets, which have
a symmetrical corolla of five united petals and five epipetalous
stamens. The latter are syngenesious,
and form a hollow cylinder round the
style. The ovary, which contains a
single ovule, is prolonged into a slender
style, divided at the top into a bifid
stigma. One disc floret is represented
cut in half to show the stamens.

The Buttercup, Pea, Saxifrage, and
Groundsel are all Dicotyledons. Text-
fig. XXX. shows the flower of the
Crocus, a Monocotyledon. Here the
perianth, which is epigynous, is not
differentiated into calyx and corolla,
but the segments are all alike and
coloured. The stamens are attached to
the perianth tube. The three carpels are
united and form the ovary and style, but
the curious funnel-shaped and toothed
stigmas are free from one another.

Among the Monocotyledons we find
a great variety of flowers, showing
polypetaly, gamopetaly, and apetaly, and
also hypogyny and epigyny, precisely
as among the Dicotyledons.

The arrangement of the flowers on the plant is by no
means haphazard. Sometimes the flowers arise singly, just as
leafy branches may occur, in the axils of the leaves. This
happens, for instance, in the Violets, but the fact is not obvious
at first sight, because the stem and the leaf bases are hidden away

FIG. XXX.— The Flower of
a Crocus. A typical Mo-
nocotyledonous Flower,
(natural order Iridacese).
Reduced.

332 APPENDIX II

underground. In the Violets also, a pair of small, scale-like
leaves occur part of the way up the flower-stalk or peduncle.
Such simplified leaves are known as bracteoles.

In a very large number of plants the flowers occur in special
groups, and then, instead of arising in the axils of ordinary
foliage leaves, they commonly arise in the axils of simplified
leaves, called bracts. Such aggregations or groups of flowers
are known as inflorescences. Inflorescences fall under two heads :
indefinite or racemose, and cymose. In the racemose inflorescence,
the main axis does not end in a flower, and thus its growth can
continue indefinitely. The older flowers are at the base, and
the younger near the apex. A simple inflorescence of this type
is called a raceme if each flower has a special stalk or pedicel
(cf. Plate XLVIL, fig. 1), and a spike, if each flower is stalkless
or sessile, as in the Lesser Butterfly Orchid (Plate XXXI., fig. 2).
If the raceme is complicated by branching, we have the form
of inflorescence called a panicle. If the main axis of the
inflorescence is suppressed altogether, so that all the pedicels
start from one point, we get an umbel (cf. Plate XIII.) ;
and if the pedicels also are suppressed, and the flowers are
crowded together on a flattened expansion of the apex of the
peduncle, we have a head or capitulum (Text-fig. XXIX.).

In the other type of inflorescence, the main axis terminates
in a flower, and the younger flowers appear below it as lateral
outgrowths. Such an inflorescence is known as a cyme. If
two lateral flowering branches grow out below the first flower,
we have a dichasium; if there is only one, a monochasium.
When all the branches of a monochasium are developed on the
same side, a scorpioid cyme is produced. Of this type of
inflorescence the Forget-me-not is a good example.

APPENDIX III

BOOKS ON THE SWISS ALPINE FLORA

FLORAS

The best flora in J&nglish is

A. GREMLI'S The Flora of Switzerland: For the Use of Tourists and Field-
Botanists (Translated from the Fifth German Edition by Leonard
W. Paitson. David Nutt, 1889),

which is now, unfortunately, out of print. A useful book,
however, is

K. W. v. DALLA-TORRE'S The Tourist's Guide to the Flora of the Alps
(Translated and Edited by Alfred W. Bennett. Swan Sonnenschein,
Le Bas & Lowrey, 1886. 5s.).

Both these works are handy for the pocket and for use in
the field, the latter especially. Neither of them are, however,
really suited to the lay reader, nor are they illustrated.
Gremli's Flora is a standard work, written in technical botanical
language, often severely contracted. The artificial system of
Linnaeus is made use of as a key to the genera, and this, while
now an archaic contrivance, has its advantages. Gremli's
flora includes not only the Alpine plants, but the whole
vegetation of Lowland Switzerland.

Dalla-Torre's Tourist Guide is written, so far as possible, in
non-technical language, which is not contracted. It is not
confined to the Swiss Alpine flora, but treats of the Alpine

833

334 APPENDIX III

floras of all the Western and Central European Mountains.
The Lowland plants are not included.

Another useful work which may be recommended is

J. HOFFMANN'S Alpine Flora: For Tourists and Amateur Botanists (Trans-
lated by E. S. Barton (Mrs A. Gepp). Longmans, Green & Co.,
1903. 7s. 6d. net).

This book deals only with Alpine plants, but is not confined to
the Swiss Alps. The language is made as simple as possible. The
book is illustrated by 40 coloured plates, containing 250 figures
of typical Alpines, in execution of more than average merit. It
is, however, rather large for the pocket, though not unwieldy.

The following works, in the absence of any satisfactory
descriptive flora in English, couched in language intelligible to
the layman, will be found very useful in Switzerland : —

BENTHAM & HOOKER'S Handbook of the British Flora (Reeve & Co.
7th Edition, 1908. 9s. net),

and the corresponding volume of plates by

FITCH & SMITH— Illustrations of the British Flora (Reeve & Co. 7th
Edition, 1908. 9s. net).

It contains descriptions of many Swiss Alpines which are
also British plants, but of course a large number of Swiss
species are not discussed. It is especially useful, however, for
determining the families and genera.

The following work, consisting of coloured plates of Swiss
Alpines,

C. & L. SCHROETER'S Taschenflora des Alpen-Wandereres (A. Raustein,
Zurich. 7 fr. 50 c.),

enjoys a wide popularity in Switzerland, and is a very useful
possession.

The photographs in

SOMERVILLE HASTINGS' Alpine Plants at Home (First Series. Go wans'
Nature Books, No. 20. 6d. net),

afford excellent illustrations of many typical Swiss Alpines.

BOOKS ON THE SWISS ALPINE FLORA 335

LARGER WORKS ON THE ALPINE FLORA

A large work in two volumes, with many coloured plates,
A. W. BENNETT'S The Flora of the Alps (J. C. Nimmo, 1897. 15s. net),

may be consulted. It is written on much the same lines as
Dalla-Torre's Tourist Guide, above mentioned, and also includes
the Alpine floras of the various mountain ranges in Central
and Western Europe.

Of the many recent works in German or French, the
following may be mentioned : —

The latest Swiss flora (in German) is by

H. SCHINZ and R. KELLER : Flora der Schweiz, zum Oebrauche auf
Exkursionen, in Schulen, und beim Selbstunterricht (A. Raustein,
Zurich. 2nd Edition, 1905 ; 3rd Edition, 1909. 6 fr. 80 c.).

The work is in the highest degree technical, and the
language contracted. It is, however, a very complete account.

G. BONNIER and G. DE LAYENS' Flore complete de la France et de la Suisse
(Paris, Librairie generate de 1'Enseignement, 1908. 11 frs.),

is written with as few technical words as possible, and is well
illustrated by over 5,000 figures.

WORKS ON THE NATURAL HISTORY, ECOLOGY,
AND DISTRIBUTION OF SWISS ALPINE PLANTS

KERNER'S The Natural History of Plants (Translated from the German by
Prof. F. W. Oliver, in 2 or 4 large 4to volumes. Cassell & Co.
1st Edition, 1894 ; 2nd Edition, 1905).

This great work, a most fascinating book, especially useful
to the layman, contains in simple language a full account
of plant life in the Alps, among much other information on
plants generally.

C. SCHROETER'S Das Pflanzenleben der Alpen. Eine Schilderung der Hoch-
gebirgsflora (A. Raustein, Zurich, 1908),

336 APPENDIX III

a thick volume, containing many illustrations, is of the highest
value, especially from the ecological standpoint. It is to be
hoped that it will one day be translated into French, if not
into English. It is hardly suited, however, to the layman.

H, CHRIST'S Der Flora der Schweitz (Translated into French by Tieche under
the title La Flore de la Suisse et ses Origines. H. Georg: Bale,
Geneva, Lyons. 2nd Edition, with Supplement, 1907),

is a standard work, especially on the distribution and origin of
the Swiss Alpines.

For a summary of recent literature, especially by the
Zurich School of Botany, on the ecology of the Swiss flora,
see T. W. Woodhead, in the Naturalist for May and June 1908.

INDEX

(For references to the technical terms mentioned in the text,
see APPENDIX I., p. 307.)

Abies excelsa, Poir., see the Spruce
Abies larix, Poir., see the Larch
Abies pectinata, D. C., see the Silver

Abnormal growths, 288-291
Accommodation to parasitic plants,

288-291

Achillea atrata, Linn., 174
Aconite, the Winter, 208, 210
Aconitine, 130

Aconitum, see the Monkshoods
Aconitum lycoctonum, Linn., 130-131
Aconitum napellus, Linn., 130-131
Adenostyles leucophylla, Reich., 177
Advertisements, flower, 78, 271-275
Affinities of the Alpine Flora, 297-

300
Ajuga pyramidalis, Linn., see the

Alpine Bugle

Alchemilla, see the Lady's Mantle
Alder, the Green, 233-234, 277
Aletsch Glacier (Bernese Oberland),

173

Algae, the, 96, 292, 293
Alkaloids, 122, 130, 168
Almagell (near Saas), 244
Alnus viridis, D. C., see the Green

Alder

Alpenglockchen, 61
Alpenrose Apples, 291
Alpenrose, the Common, 21-27, 117
affinities of, 22
and soils, 177
837

Alpenrose, the Common-— continued

habitat of, 24

scales on leaves of, 26-27

transpiration of, 25
Alpenrose, the Hairy, 21-27, 117

and soils, 25, 117

scales on leaves of, 27

specific characters of, 25
Alpenroses, the, 14, 20, 21-27, 177,
220, 224, 244, 259, 260, 281, 291

derivation of name, 22

galls on, 291

time of flowering of, 31
Alp (Alpen), 23, 24
Alpine plant defined, 3, 5
Alpine Rose, see Rose, the Alpine
Alpine species, number of, 8
Alpine zone defined, 3, 5
Alpines in the Arctic regions, 299,

305-306

Alps, the, as barriers, 304
Alps, the Austrian, 297, 298
Alsine aretioides, M. K., 185
Alsine octandra, Schur., see A.

aretioides, M. K.
Alsine sedoides, Froel, see the Dwarf

Alsine

Alsine, the Dwarf, 184-185
Alsines, the High Alpine, 184-185
Altai, the, 297, 298
Amber, 241
America, North, 109, 249, 303, 304,

305

America, South, 231
Andes, the, 231

338

INDEX

Andes, the Bolivian, 173
Androsace carnea, Linn., see the

Red-flowered Androsace
Androsace chamwjasme, Willd., see

the Dwarf Androsace
Androsace charpentieri, Heer, see

Charpentier s Androsace
Androsace, Charpentier 's, 182
Androsace glacialis, Hopp., see the

Glacial Androsace
Androsace helvetica, Gaud., see the

Swiss Androsace
Androsace imbricata, Lam., see the

Imbricated Androsace
Androsace obtusifolia, All., see the

Obtuse-leaved Androsace
Androsace pubescens, D. C. , see the

Downy Androsace
Androsace, the Downy, 182
Androsace, the Dwarf, 74, 300
Androsace, the Glacial, 173, 174, 182,

183

Androsace, the Imbricated, 181-182
Androsace, the Obtuse-leaved, 74
Androsace, the Red-flowered, 74
Androsace, the Swiss, 181
Androsace vitaliana, Lap., see Vital's

Androsace

Androsace, Vital's, 73-74, 182-188
Androsaces, the, 33, 61, 69, 72-74,

181-183, 192, 212, 281
Androsaces, the High Alpine, 181-

188, 192
Anemone alpina, Linn., see the

Alpine Anemone
Anemone alpina, var. sulphurea, see

the Yellow Alpine Anemone
Anemone halleri, All., see Haller's

Anemone

Anemone, Haller's, 86-37, 295
Anemone narcissiftora, Linn., seethe

Narcissus-flowered Anemone
Anemone nemorosa, Linn., see the

Wood Anemone
Anemone pulsatilla, Linn., see the

Pasque Flower
Anemone, the Alpine, 36, 87-41, 108,

117, 118, 123, 128, 272, 306
and soils, 38
fruits of, 39-41
involucre of, 37
male flowers of, 39

Anemone, the Narcissus-flowered,

34, 41-42, 67, 272
fruits of, 42
local occurrence of, 41
umbels of, 42
Anemone, the Spring, 31, 34-37, 38,

41, 108, 306
fruits of, 36
hairs on, 35-36
involucre of, 35
rosette of leaves of, 35
time of flowering of, 31, 34
Anemone, the Wood, 34, 42
Anemone, the Yellow Alpine, 37-41,

117

and soils, 38
yellow flowers of, 38
time of flowering of, 31
Anemone vernalis, Linn., see the

Spring Anemone
Anemones, the, 33, 84-42, 127, 230,

281, 284
Animal-distributed fruits and seeds,

158-159, 251, 284, 285-286
Annuals in the Alps, 114
Antarctic regions, the, 180
Antennariat see the Everlastings
Antennaria alpina, Gaertn. , 279
Antennaria carpathica, Bl. and

Fing., 21

Antennaria dioica, Gaertn. , 21
Anthericums, the, 113
Anthyllis vulneraria, Linn., see the

Lady's-fingers

Antirrhinum, see the Snap-Dragons.
Anthers, closing of, 281
Ants as bodyguards to plants, 276
Ants as seed distributors, 286
Apple, the, 219
Arabis coerulea, Haenke, 192
Arabis halleri, Linn. , 295
Arctic flora, the ancient, 303-306
Arctic regions, percentage of Alpines

in, 299

Arctic regions, the flora of, 30, 109,
157, 180, 215, 249, 279, 299, 303,
305, 306

Arched flower-stalks, 64
Arctostaphylos alpina, Sprengel, see

the Alpine Bearberry
Arctostaphylos uva-ursi, Sprengel,
see the Red Bearberry

INDEX

339

Arenaria cherleria, Hook.f., see

the Dwarf Alsine
Aretia vital iana, Murr. , see Vital's

Androsace

Arnica (medicinal), 133
Arnica montana, Linn., see the

Arnica

Arnica, the, 183, 267
Arolla, 243

Arolla Pine, see the Stone Pine
Arum maculatum, Linn., see the

Cuckoo-Pint
Asia, 109, 249
Asia, Central, 104, 297, 305
Asia, Northern, 20
Asphodel, the Scotch, 178
Astrantia, 124
Atmosphere, the, 10, 11, 202
composition of, 12
dryness of, 12, 202
Atragene alpina. Linn., see the

Alpine Clematis
Atragene, the, see the Alpine

Clematis

Auricula, the, 67, 70-71, 117
and soils, 71, 117
hybrids of, 70-71

protections against transpiration, 71
Auriculas, the cultivated, 67, 70

origin of, 70
Australia, 104

Austrian Alps, the, 118, 297-298
Avalanches, effects of, 94, 237, 238, 239
Avens, the, 230, 286
Avens, the Creeping, 41, 111, 127-

128, 131
Avens, the Mountain, 39, 41, 108,

111, 127-128

Avens, the Water, 151, 167-159, 280
Awned fruits, 36, 39-41, 108, 128, 158-

159, 230, 284
Azalea procumbens, Linn., see the

Trailing Azalea
Azalea, the Trailing, 22, 106, 109-110,

186, 231
Azaleas, the, 22, 109

Ball, John, 5, 8, 173, 305
Bartsia, 218

Bats as flower pollinators, 279

Bearberry, the Alpine, 224, 230-231

Bearberry, the Red, 230-231

Bearberries, the, 230-231

Bedstraw, the, 229

Beech, the, 5, 235

Bees as cross-pollinators, 46, 55, 69,

115, 161, 165, 168, 223, 270
Beetles as unbidden guests, 209,

276

Bel Alp, the, 23
Bell-flower, Scheuchzer's, 84-87, 88,

89, 152

mechanism of stylar brush, 85-87
unbidden guests, protections

against, 85
Bell-flower, the Bearded, 46, 89, 272,

277

appendages on corolla, 89
the beard of hairs, 89, 277
Bell-flower, the Incised, 90
Bell-flower, the Mont Cenis, 89-90
Bell-flower, the Tufted, 90-91, 114,

277

biennial, 91, 114
hairs on, 91, 277
yellow flowers of, 90
Bell-flowers, the, 34, 44, 61, 88-91,

92, 150, 152, 263, 280, 284
distribution of the seeds, 88
fruit, 88, 263

stylar brush mechanism, 85-87
Bell Gentians, see Gentians, the Bell
Bellidastrum michelii, Cass. , see the

Alpine Daisy
Bellis perennis, Linn., see the

Common Daisy
Berne, Canton, 175
Bernese Oberland, the, 116, 174,

175, 242, 294, 296
Berries, 220-222, 225, 227, 233, 255,

260, 285

Best time of year for flowers, 28-29
Betulacece, see the Birch family
Biennials, 106, 114
Bilberries, the, 220, 221-224, 231, 244

pollination of, 223-224
Bilberry family, the, 221
Bilberry, the True, 221-222, 223, 224
Binnen Thai, the, 90
Bird life in the Alps, 220-221, 285
Birch family, the, 233

340

INDEX

Bird's-foot Trefoil, the, 104-105, 200,

252

Biscutella Icevigata, Linn., 251, 299
Bistort, the, 151, 156
Bittercress, the Alpine, 196
Black-Cock, the, 221
Blacken, 120
Blacken Alp, 120
Blackengarten, 120
Bladderworts, the, 211, 214
Blaser, the (Tyrol), 203
Blue-flowered Alpines, 43-46, 47, 49,

114

Bolivia, 20
Bonnier, Prof., 45, 114, 117, 118,

200-202, 203, 306
Borage family, the, 183, 272
Boragineae, see the Borage family
Box, the, 251
Brambles, the, 229
Breuil, 173
Brevent, the, 78
British plants in the Alps, 151-152,

178, 205-206
Brugger, C. G., 30
Buckbean, the, 43
Bugle, the Alpine, 131
Bupleurum, 124
Buttercup family, the, 33, 34, 111,

124, 127, 130, 205, 207, 209,

229
Buttercup, the Aconite-leaved, 209-

210

Buttercup, the Alpine, 111-112, 118
Buttercup, the Bulbous, 151
Buttercup, the Field, 151
Buttercup, the Glacial, 118, 174, 193-

194, 299
Buttercup, the Parnassus - leaved,

194-195

Buttercup, the Pyrenean, 194-195
Buttercups, the, 33, 38, 127, 149, 205
Buttercups, the High Alpine, 193-

195
Butterflies as cross-pollinators, 45,

49, 55, 58, 78, 106, 111, 129, 156,

161, 254, 270, 271, 273
Butter wort family, the, 210
Butterwort, the Alpine, 211-214
glands on the leaves, 212-215
insectivorous habit of, 213-215
Butterwort, the Common, 211-216

Butterwort, the Large-flowered, 211,

216
Butterworts, the, 210-216, 281

number of insects caught by, 215

uses of leaves of, 215-216
Buxus, see the Box

Calcareous soils (or rocks), 112, 117-

118, 132, 181, 183, 190, 195
Californian Lily, the, 122
Ga llianthemum co riandrifo Hum ,

Reichb., 195
Callianthemum rutcefolium, Reichb.,

see the Rue-leaved Callian-
themum
Callianthemum, the Rue-leaved, 193,

195
Calluna vulgaris, Salisb., see the

Ling
Caltha palustris Linn., see the

Marsh Marigold
Calyx, inflated, 106, 163-164, 276
Campanula barbata, see the Bearded

Bell-flower
Campanula cenisia, Linn., see the

Mont Cenis Bell-flower
Campanula excisa, Schleicher, see

the Incised Bell-flower
Campanula rotundifolia. Linn., see

the Harebell

Campanula rhomboidalis, Linn., 152
Campanula scheuchzeri, Vill. ,

Scheuchzer's Bell-flower
Campanula thyrsoidea, Linn.,

the Tufted Bell-flower
Campanulaceae, the, 34, 83-92
Campanulas, the, see the Bell-flowers
Campions, the, 150, 151, 163-164, 276
Campion, the Common Moss, 173,

180
Campion, the Sessile-flowered Moss,

180-181

Campions, the Moss, 178, 179
Canada, 301, 303
Cape, the (South Africa), 30
Caprifoliaceae, see the Honeysuckle

family
Cardamine alpina, Willd., see the

Alpine Bittercress

see

INDEX

341

Cardamine pratensis, Linn., see the

Ladies' Smock
Carlina acaulis, Linn., see the

Carline Thistle

Carpathians, the, 20, 118, 297, 298
Carpet plants, the, 101, 105, 106-107,

109, 111, 177, 178, 186-188, 230,

233
Carpet plants, the High Alpine, 186-

188
Caryophyllaceae, the, see the Pink

family

Catchflies, the, 151
Catchfly, the Rock, 82, 115, 200,

225
Catchfly, the Stemless, see the Moss

Campion
Caux, 7

Centaurea alpina. Linn., 276
Centaury, the, 42
Cerast, the Alpine, 178
Cerastium alpinum, Linn., see the

Alpine Cerast
Ceylon, 239
Chalet plants, 119
Chamonix, 7, 78, 173
Chateau Belvedere (Maloja), 244
Cherleria sedoides, Linn., see the

Dwarf Alsine
Cherry, the, 232, 289
Chlora perfoliata, Linn., see the

Yellow-Wort
Chough, the Alpine, 221
Christ, Dr, 5, 30, 173, 237, 242, 305
Christmas Roses, the, 208
Chrysanthemum alpinum, Linn., see

the Alpine Ox-eye Daisy
Chrysanthemum leucanthemum, Linn. ,

see the Ox-eye Daisy
Claytonia, 300
Cleavers, the, 286
Cleistogamous flowers, 278-279
Clematis alpina, Miller, see the

Alpine Clematis
Clematis, the Alpine, 229-230, 284

climbing organs of, 229
Clematis vitalba, Linn., see the

Traveller's Joy
Climbing plants, 228-229
Closing of flowers, 281-282
Clover, the Alpine, 132
Clovers, the, 263

Col de Giant, 173

Colchicin, 168

Colchicum alpinumt D. C., see the

Alpine Saffron
Colchicum autumnale, Linn., see the

Meadow Saffron
Colonies of Saxifrages, 82-83
Colonies of Semper vivums, 100
Colonisation of new ground, 94-97,

283

Colours of flowers, 43-47, 254, 271
Colours of flowers, intensity of, 45,

149, 184, 202
Colour variations in Alpine Flowers,

38, 46-47, 155-156, 159
Coltsfoot, the, 31, 94, 97, 260, 266,

267

time of flowering of, 31
Commune, the, 23, 142, 238
Compositse, the, see the Daisy

family

Composite organisms, 292
Composites, the, see the Daisy

family

Concealed honey, 66, 273-274, 275
Coniferse, the, 233, 234-235, 241, 277,

291
Coniferous forests, the, 5, 6, 220, 284-

245, 248, 286
former distribution, 237
destruction of, 238
legislation respecting, 238
upward limits of, 235-236
vertical distribution of, 235-236
Conspicuousness of flowers, 124, 164,

271-272

Contractile roots, 122-123
Contrivances to protect the pollen,

281-282
Convallaria majalis, Linn., see the

Lily of the 'Valley
Convallarieae, the, 254
Convolvuli, the, 229
Cowberry, the, 221-223
Cowslip, the, 66, 67
Cow-wheat, the, 218, 286
Crassulaceae, see the Stonecrop

family
Crawling insects as unbidden guests,

275
Crocus, the Meadow, see the Meadow

Saffron

342

INDEX

Crocus, the Spring, 31, 46, 62, 152,

153-156, 166, 281, 299
colours of flowers of, 155, 156
time of flowering of, 31, 153-155
the corm of, 153-155
Crocus vernus, ALL, see the Spring

Crocus
Cross-fertilisation, advantages of, 52,

268
Cross - pollination through insect

agency, 268-275
Crowberry family, the, 231
Crowberry, the, 110, 177, 178, 186,

231-282, 300
Crucifer family, the, 183, 192, 196,

299
Crustaceous Lichens, see Lichens,

Crustaceous
Cuckoo-flower, the, see the Ladies'

Smock

Cuckoo-Pint, the, 123
Cudweeds, the, 20
Cultivations, experimental Alpine,

199-203

Cupressaceae, see the Cypress family
Currant, the Alpine, 291
Curving of flower stalks, 281
Cushion plants, 101, 178, 179-186
Cypress family, the, 233
Cypripedium calceolus, Linn. , see the

Lady's Slipper

Daisy family, the, 15, 18, 87, 111, 119,

125, 133, 266, 267, 272, 276, 284
Daisy, the Alpine Ox-eye, 198, 266
Daisy, the Common, 15 (footnote),

18, 19, 149, 266, 281
Daisy, the Common Ox-eye, 151,

198, 266, 267

Dandelion, the, 41, 119, 173, 284
Daphne alpina, Linn. , 260
Daphne family, the, 259
Daphne laureola, Linn., see the

Spurge Laurel
Daphne mezereum, Linn., see the

Mezereon

Daphne striata, Tratt, 260-261, 273
Darwin, C., 52, 69, 168, 214-215, 302-

303

Dauphine Alps, the, 118
Davos, 24, 44, 83, 116, 157, 244
Davos Platz, 24

St Johann Kirche at, 24
deCandolle, A., 296
Deception of insects by flowers, 217
de Saussure, 173

Destruction of the forests, 237-239
by man, 238
by goats and cattle, 238
by avalanches, 238
by disease, 238
effect of, 239
Dew and pollen, 281
Dianthus, see the Pinks
Dianthus carthusianorum, Linn., see

the Carthusian Pink
Dianthus glacialis, Hanke, 295
Dianthus inodorus, Steud., see the

Wood Pink
Dianthus superbus, Linn., see the

Large-flowered Pink
Dianthus sylvestris, Wulf., see the

Wood Pink
Dintel, Fraul., 99
Diseases of Alpine plants, 286-291
Diseases of forest trees, 238
Distribution (geographical), in-
equalities of, 116, 294-297
Distribution (geographical), wide,

298
Distribution of fruits by animals,

159, 285-286
Distribution of seeds and fruits, 283-

286

Districts rich in rare species, 294-296
Dock family, the, 120, 156
Dock, the Alpine, 120
Dodder, the, 219
Dom, the, 172
Doronicum (Aronicum) scorpioides,

Lam., 177
Double Berry, 227
Douglasia vitaliana, B. and H.f.,

see Vital's Androsace
Draba aizoides, Linn., 183
Draba carinthiaca, Hoppe, 192
Draba johannis, Host., see D.

carinthiaca, Hoppe
Draba pyrenaica, Linn., see the

Pyrenean Draba
Draba, the Pyrenean, 183

INDEX

343

Draba tomentosa, Wahl., 183
Draba vema, Linn. , see the Whitlow-
grass

Draba wahlenbergii, Hartm., 192
Drooping of flowers, significance of,

277, 280

Drosera, see the Sundews
Dryas octapetala, Linn., see the

White Dryas

Dryas, the White, 39, 41, 105, 106-
109, 118, 128, 186, 230, 284, 299,
300

age of stem, 107
carpet plant, 106-107, 186
distribution of, 299-300
hairs on leaves, 108
soils, 118

Dwarf plants, 176-177, 201-202, 244
Dwarf plants, the High Alpine, 192-

198
Dwarf Willows, the, 187-190. 234

Early flowering in the Alps, 29, 30,

64, 147, 154, 282, 283
Edelweiss, the, 14, 15-20, 36, 38, 103,

118, 124, 204, 297
and soils, 118
derivation of name, 17
hairy coat of, 17-20, 38
local distribution of, 16-17
the heads of, 18-19
Edible seeds and fruits, 243, 285
Elder-odour, the, 272
Empetraceae, see the Crowberry

family
Empetrum nigrum, Linn., see the

Crowberry

Endophyllum sempervivi, 289
Engadine, the, 21, 30, 72, 116, 150,

295

Engadine, the Lower, 43
Engstlen Alp (Canton Berne), 71,

117, 120, 206, 243
Epilobium, see the Willow-herbs
Eranthis, see the Winter Aconite
Erica carnea, Linn. , see the Mediter-
ranean Heath
Ericaceae, see the Heath family

Eritrichium nanum, Schrad. , see the

Eritrichium

Eritrichium, the, 44, 49, 188-184, 306
Erythrcsa centaurium, Pers. , see the

Centaury

Euphorbia, see the Spurges
Euphorbia cyparissias, Linn., 290
Euphorbiaceae, see the Spurge family
Euphrasia, see the Eyebright
Europe, Central, the flora of, 298,

301, 302
Europe, Northern, the flora of, 231,

232, 299
Europe, Southern, the flora of, 118,

276, 298, 301
Europe, Western, the flora of, 291,

298, 299, 304

Everlastings, the, 21, 279
Evolution of the flowers of the

Gentians, 58-59
Exobasidium rhododendri, 291
Experimental cultivation of Alpines,

199-203

Explosive fruits, 162, 265, 285
Extinction of the Miocene European

flora, 304
Eyebright, the, 218, 219

« Fair Maid of France," 209

False Berry, 228

False nectaries, 217-218

Finns the, 232

Finsteraarhorn, the, 174

Fir, the Silver, 245

Fir, the Scotch, see the Scotch Pine

Firs, the, 234

Field Pansy, the, 129, 159-168

Fisch, Dr, 44

Flies as cross-pollinators, 46, 208-209,
217-218, 254, 260, 274

Flower advertisements, 45, 124, 218,
270, 271-275

Flower, structure of, 13, and Appen-
dix II.

Flowering in the snow, 62-64, 154-155

Flowers and insects, 268-277

Flowers that do not open, 278-279

Flying insects as unbidden guests,
275

344

INDEX

Forbes, E., 303

Forested land, percentage of, in

Switzerland, 237
Forests, Alpine, 32, 33, 234-245
Forests, the, and dry ness of the

atmosphere, 239

Forget-me-nots, the, 44, 184, 306
Formic acid, 276
Fossil resin, 241
Foxglove family, the, 113, 211, 218,

219, 286

Fragaria, see the Strawberry
France, flora of, 4, 172, 298
Fringed Gentians, the, 48, 54-56
Fruits, distribution of, 282-286
Fungi, parasitic, 222, 287, 289
Fungi, the, 96, 222, 287, 289, 292,

293
Fuorcla Surlej (Engadine), 243

Gagea liotardi, Schult, see the

Alpine Gagea
Gagea, the Alpine, 119
Gagea, the Yellow, 119
Gaillardia, 300
Galium, see the Bedstraws
Galium aparine, Linn., see the

Cleavers
Galls, 290-291
Gentian Brandy, 58
Gentian family, the, 33, 42
Gentian, the Autumn, 43
Gentian, the Bavarian, 47-49, 174,
191

flowers scentless, 49

pollination of, 47

time of flowering, 48
Gentian, the Broad-leaved, 48, 51
Gentian, the Common Bell, 31, 46,
47, 48, 50-54

cross-pollination of, 52-53

self-pollination of, 54

structure of flower, 51-53

time of flowering, 31
Gentian, the Delicate, 54, 56, 192,
278

habit, 55

scales on corolla, 56

Gentian, the Field, 54, 66-56, 59,

278

annual, 55
habit, 55

flowers sensitive to light, 56
scales on corolla, 55
unbidden guests, 55
Gentian, the Hungarian, 59-60
Gentian, the Marsh, 43
Gentian, the Purple, 57, 58, 59
habit, 57
roots, 58
Gentian, the Snow, 47, 49-50, 56,

114

annual, 49, 114
closing of the flowers, 56
sensitiveness to sunlight, 56
Gentian, the Spotted, 58, 59
Gentian, the Spring, 31, 47, 48-49,

50, 176

flowers scentless, 49
pollination of, 49
time of flowering, 31
Gentian, the Yellow, 67-59, 60, 177,

273

age of plants, 57
flowers primitive, 58
habit, 57
petals free, 57
pollen exposed, 273
roots of, 58
Gentiana acaulis, Linn., see the

Common Bell Gentian
Gentiana alpina, Vill., 51
Gentiana amarella, Linn., see the

Autumn Gentian
Gentiana bavarica, Linn., see the

Bavarian Gentian
Gentiana brachyphylla, Vill., 174,

192
Gentiana campestrfe, Linn., see the

Field Gentian
Gentiana excisa, Presl., see the

Broad-leaved Gentian
Gentiana latifolia, Gren. and Godr. ,

see the Broad-leaved Gentian
Gentiana lutea, Linn. , see the Yellow

Gentian
Gentiana nivalis, Linn., see the Snow

Gentian

Gentiana pannonica, Scop., see the
Hungarian Gentian

INDEX

345

Gentiana pneumonanthe, Linn., see

the Marsh Gentian
Gentiana punctata, Linn., see the

Spotted Gentian
Oentiana purpurea, Linn., see the

Purple Gentian
Oentiana, see the Gentians
Oentiana tenella, Rottb., see the

Delicate Gentian
Oentiana verna, Linn. , see the Spring

Gentian

Gentianaceae, see the Gentian family
Gentianellas, the, 48, 54
Gentians, the, 33, 42-60, 73, 122, 192,

271, 273,278,281,299
Gentians, the Fringed, 48, 54
Geographical distribution of Alpines,

the, 294-306
Geological history of the Alpine

flora, the, 300-306
Geranium gylvaticum, Linn. , see the

Wood Geranium
Geranium, the Wood, 44
Geraniums, the, 44, 150, 151, 285

the fruits of, 285
Gerarde, John, 215
Germany, flora of, 4, 298
Oeum montanum, Linn., see the

Mountain Avens

Geum reptans, Linn., see the Creep-
ing Avens
Oeum rivale, Linn., see the Water

Avens

Geum, see the Avens
Glacial flora of the Alps, the, 7, 176
Glacial Period, the, 301-306
Glacial region, the, 6, 7
Glaciation of the British Isles, the,

302
Glaciers, the retreat of the, 296,

304-305
Globe-flower, the, 124, 150, 207-209,

270, 273, 274, 281
Olobularia cordifolia, Linn. , see the

Round-leaved Globularia
Olobularia nudicaulis, Linn., see the

Bare-stemmed Globularia
Globularia, the Bare-stemmed, 110-

111
Globularia, the Round-leaved, 106,

110-111
Globularias,the, 44, 110-111, 281, 299

Globularias, the Alpine, 110-111
Onaphalium alpinum, Linn., see

Antennaria alpina, Gsertn.
Gnaphalium, see the Cudweeds
Gorner Grat, the, 29, 172
Grasses, the, 149, 157, 219, 278
Grass-of-Parnassus, the, 80, 203, 216-

218

staminodes of, 217-218
Graubunden, Canton, see Grisons
Gray, Asa, 303
Great Ice Age, the, 301-305
Greenland, 301
Orefforia vitaliana, Duby, see Vital's

Androsace
Grindelwald, 7, 242
Grisons, Canton (Graubunden), 72,

175, 295, 296
Gronovius, 249
Grouse, the, 221

Habenaria bifolia, R. Br., see the

Lesser Butterfly Orchid
Habenaria conopsea, Benth., see the

Fragrant Habenaria
Habenaria viridis, R. Br., see the

Frog Orchid

Habenaria, the Fragrant, 134
Habit, change of, 114
Habit, the Alpine, 176-178, 199-203
Hairs aiding distribution of seeds

and fruits, 189, 284
Hairs as protections to plants, 89,

99, 250, 277
Harebell, the, 73, 88-89, 151, 200,

201, 206

the cotyledons, 83-84
mechanism of stylar brush, 85-87
origin of specific name, 83
shade-leaves of, 84
sun-leaves of, 84
Hawk-moths as cross-pollinators, 49,

136, 261
Hay of the Alpine meadows, the,

142-149

Heart's-ease, see the Field Pansy
Heath family, the, 21, 25, 109, 134,

221, 224, 230, 261

346

INDEX

Heath, the Alpine, see the Mediter-
ranean Heath
Heath, the Mediterranean, 110, 140-

141, 231, 251, 299
Heather, White, 46
Heaths, the, 4, 215
Heer, O., 175
Height, greatest at which plants

occur, 172-174
Hellebores, the, 208, 273
Herb Paris, the, 257-258
High Alpine region, the, 6-7, 169-178
Highlands of Scotland, the, 110
Himalayas, the, 20, 22, 109, 172, 297,

298

Hips of Roses, the, 285
Homogyne alpina, Cass., see the

Alpine Lettuce
Honey and insects, 269-275
Honey-bees, see Bees
Honey guides, 161, 275
Honeyless flowers, 39, 261, 272
Honey - sucking birds as flower

pollinators, 279

Honeysuckle family, the, 224, 248
Honeysuckle, the Black, 225-226,

228

berries of, 225-226
union of ovaries of, 225
Honeysuckle, the Blue, 228-229

false-berry of, 228
Honeysuckle, the Mountain, 226-227,

228

double berry of, 227
Honeysuckles, the, 220, 224-229
Hook-climbers, 229
Hooked fruits, 285
Hooker, Sir J., 303
Horned stamens, 223, 231
House-leek, the Mountain, 99-102,

118

migration of, 101-102
runners of, 101-102
House-leek, the Spider's-web, 98-99

hairs on leaves of, 98-99
House-leeks, the, 97, 98-108, 127,

131, 289
habit, 98

origin of generic name, 103
Humble-bees as cross-pollinators,

53, 58, 106, 126, 130, 254, 270,

273

Humming-birds as pollinators, 279
Hutchinsia alpina, R. Br., see the

Alpine Hutchinsia
Hutchinsia brevicaulis, Hoppe, see

the Short-stemmed Hutchinsia
Hutchinsia petrcea, R. Br., see the

Rock Hutchinsia
Hutchinsia, sp., 299
Hutchinsia, the Alpine, 196, 197
Hutchinsia, the Rock, 196-197
Hutchinsia, the Short-stemmed, 196,

197
Hygrometer plants, 125-126

Ice Age, see the Great Ice Age
Illumination, intensity of, 45, 172,

202, 203, 246, 247, 264, 265, 282
India, 20, 22, 227, 239
Inequalities of distribution of alpines,

294-297

Insect-capturing plants, 211-215
Insect-fertilised flowers, number of,

270

Insect parasites, 238, 290-291
Insectivorous plants, 211-215
"Insectivorous Plants," by C.

Darwin, 214

Insects and flowers, 268-277
Insects : how they recognise flowers,

274

Insects, the eyes of, 274
Insects, the food of, 269
Insects, the number visiting Viola

calcarata, Linn., 129
Intensity of colour, see Colours of

flowers

Interglacial periods, 302
Ireland, 140

Iridaceae, see the Iris family
Iris family, the, 153
Irish Shamrock, the, 263
Italy, 172, 299
Ivies, the, 229

Jaccard, Prof., 296

Jacob's Ladder, the, 150, 151

INDEX

347

Japan, 20

Jardin of the Mer de Glace, 173-174

Jasmine scent, the, 272

Julier Road, the, 211

Juniper, the Alpine, see the Dwarf

Juniper

Juniper, the Common, 233
Juniper, the Dwarf, 106, 187, 233,

300

Junipers, the, 234
Juniperus communis, Linn., var.

nana, Willd., see the Dwarf

Juniper
Jura, the, 237

Kerner, Prof., 53, 63, 70, 102, 203,

212, 215, 276
Kidney Vetch, the, 252
Kleine Scheidegg, the, 242
Klosters, 244
Knot-foot, the, 256-257
Kblreuter, J. G., 269

Labiateae, see the Mint family
Ladies' Smock, the, 196
Lady's-fingers, the, 105-106
Lady's Mantles, the, 279, 281
Lady's Slipper Orchid, the, 133-134,

138

Lamium amplexicaule, Linn., 279
Laplanders, the, 216, 232, 250
Larch disease, the, 238
Larch, the, 5, 6, 97, 234, 235, 236,

237, 239, 240, 241-242, 244, 246,

248

Larix europcea, D. C. , see the Larch
Laurel, the Spurge, 259
Lauteraarhorngipfel, the, 174
Leaf-cutting insects, 276
Leguminosae, see the Pea family
Lentibulariaceae, see the Butterwort

family

Lenzer Heide, 244
Leontopodium alpinum, Cass. , see the

Edelweiss
Lepidoptera, night-flying, 275

Lesser Butterfly Orchid, see Orchid,

the Lesser Butterfly
Lettuce, the Alpine, 266-267
Leucanthemum vulgare, Lam., see

the Ox-eye Daisy
Lichen, the Old Man's Beard, 239,

291

Lichens as rock colonisers, 96
Lichens, the, 174, 239, 291-293
Lichens, the Crustaceous, 96, 170,

292

Light, intensity of, see Illumination
Liliacese, see the Lily family
Lilium martagon, Linn., see the

Martagon Lily
Lily family, the, 113, 119, 120, 254,

257, 258

Lily of the Valley, the, 247, 255
Lily, the Martagon, 177, 268-259, 275
Lily, the May, 247, 255, 257
Limestone soils, 25, 71, 77, 117-118,

242
Linaria alpina, Mill., see the Alpine

Toadflax
Linaria cymbalaria, Mill., see the

Ivy-leaved Toadflax
Linaria vulgaris, Mill. , see the Yellow

Toadflax

Ling, the, 110, 140-141, 200
Linncea, borealis, Gronov., see the

Linncea

Linncea, the, 248-251, 272, 277
distribution, 249
distribution of the fruits, 251
origin of name, 249
scent of, 272
Linnaeus, Carl, 15 footnote, 190,

215, 249

Listera cordata, R. Br., 255
Listera ovata, R. Br., 255
Lloydia serotina, Sweet, see the

Mountain Lloydia
Lloydia, the Mountain, 178
Localisation of disease in plants,

288-291

Loew, Dr, 270
Loiseleuria procumbens, Desv., see

the Trailing Azalea
Lonicera alpigena, Linn., see the

Mountain Honeysuckle
Lonicera ccerulea, Linn., the Blue

Honeysuckle

348

INDEX

Lonicera nigra. Linn., see the Black

Honeysuckle
Lonicera periclymenum, Linn., see

the Woodbine

Lonicera xylosteum, Linn. , 226
Lotus corniculatus, Linn., see the

Bird's-foot Trefoil
Louseworts, the, 218-219
Lowland plants in the Alps, 206
Lowland zone defined, 4
Lychnis dioica, Linn., see the Red

Lychnis

Lychnis, the Red, 151, 165
Lychnis, the White, 165, 275
Lychnis vespertina, Siboth., see the

White Lychnis
Lythwim,sp., 69, 168

Macroglossa stellatarum^ 49
Maianthemum convallaria, Weber,

see the May Lily
Maloja Pass, the, 244
Marigold, the Marsh, 205-206, 207,

216, 273, 300
Matterhorn, the shoulder of the,

174

Mattmark, 90

May Lily, the, see Lily, the May
Meadows, the Alpine, 23, 24, 28, 29,

32, 33, 142-152
Mechanism of stylar brush, 85-87,

92, 133
Mediterranean flora, the, 4, 111, 140,

251, 298-299
Mediterranean species in the Alps,

299

Melampyrum, see the Cow-wheat
Melampyrum sylvaticum, Linn., see

the Cow-wheat
Menyanthes trifoliata, Linn., see the

Buckbean

Mezereon, the, 259-261
Migration of the ancient Arctic flora,

303-304

Mildews, the, 287
Milkwort family, the, 251
Milkwort, the Common, 251
Mint family, the, 104, 131

Miocene flora of Europe, the, 304
Miocene flora of North America, the,

304

Miocene Period, the, 300, 303
Mistletoe, the, 219
Moneses grandiflora. Gray, see the

Single-flowered Wintergreen
Monkshoods, the, 44, 124, 180-131,

177, 273, 281

Mont Blanc, 172, 173, 200
Monte Rosa, 172, 190, 233
Morel, 150
Mosses, 174

Mosses as rock colonisers, 97
Moths as cross-pollinators, 49, 165,

259, 270

Moulds, the, 287
Movement, power of, 264-265, 281-

282

Mueller, H., 129, 270-271
Multiple vision of insects, the, 274
Mushroom, the, 287
Mutilation of plants by scythe, 144,

148

Myosotis alpestris, Schmidt, 184
Myrmecophilous plants, 276

Naiadacese, see the Pondweed

family
Narcissus - flowered Anemone, see

Anemone, the Narcissus-
flowered

Nepenthes, see the Pitcher Plants
Nettle, the Common, 119
Nettle, the Small, 119
Nettles, the Dead. 278
Nigritella angustifolia. Rich. , see the

Black Nigritella
Niqritella nigra, Reichb., see the

Black Nigritella
Nigritella, the Black, 184-136, 140,

272

ovary of, 140
scent of, 135, 272
tuberous roots of, 134-135
Nivial flora, the, 176
North America, 299, 300, 301, 302,

303, 304, 305

INDEX

349

Oak apples, 290

Oak, the, 290

Oats, 288

Oligocene Period, the, 300

Onagraceae, see the Willow-herb

family

Orchid, the Frog, 134
Orchid, the Vanilla, 135, 272
Orchid, the Lesser Butterfly, 134,
135, 136-140, 255

flower structure, 136-138

method of pollination, 138-139

scent of, 136

twisted ovary, 139-140
Orchids, the Alpine, 133-140, 273

types of habitat, 134
Orchis maculata, Linn., see the

Spotted Orchis
Orchis, the Spotted, 134
"Origin of Species," the (by

C. Darwin), 302-303
Origin of the Alpine Flora, 300-306
Oxalidaceae, see the Oxalis family
Oxalis acetosella, Linn. , see the Wood

Sorrel

Oxalis family, the, 263
Oxlip, the, 31, 66, 67-69

heterostylism of, 68

time of flowering, 31
Oxytropis fcetida, D. C., 295

Pansy, the Field, 46, 124, 151, 159-

163, 254, 266, 571
cross-pollination of, 161-162
fruits of, 162-163, 266
Papaver alpinum, Linn., see the

Alpine Poppy
Papaver aurantiacum, Lois., see the

Alpine Poppy
Papaver nudicaule, Linn., see the

Alpine Poppy
Papaver pyrenaicum, Wild., see the

Alpine Poppy

Papaveraceae, see the Poppy family
Parasitic animals, 286, 290, 291
Parasitic plants, 238, 286-291
Paris, 200-201

Paris quadrifolia, Linn., see the

Herb Paris
Parnassia palustris, Linn., see the

Grass-of-Parnassus
Pasque-flower, the, 34, 306
Pastures, the Alpine, 28, 24, 32, 33,

34, 93, 94
Pea family, the, 104, 105, 132, 252,

253, 273, 281
Peach, the, 232

Peat of the Meadows, the, 150-151
Pedicularis incarnata, Jacq., £95
Pedicularis jacquini, Koch., 295
Pedicularis, see the Louseworts
Pedicularis verticillata, Linn., 218
Petrocallis pyrenaica, Linn., see the

Pyrenean Draba

Phyteuma corniculatum, Clairv., 176
Phyteuma hemisphcericum, Linn. ,

192

Phyteuma humile, Schleich., 192, 295
Phyteuma orbiculare, Linn., see the

Round-headed Rampion
Phyteuma pauciflorumt Linn., 192,

296
Phyteuma pedemontanum, Schulz, see

P. pauciflorum, Linn.
Phyteuma spicatum, Linn., 91
Phyteuma, see the Rampions
Picea excelsa, Link., see the Spruce

Fir

Pigments of Flowers, 45, 47, 271
Pine-forests, see the Coniferous

Forests

Pine-needles and primitive soils, 97
Pine, the Mountain, 187, 234, 243-

244

Pine, the Scotch, 243, 245
Pines, the, 97, 151, 234, 240, 241,

242
Pinguicula alpina. Linn., see the

Alpine Butterwort
Pinguicula grandiflora, Lam., see

the Large-flowered Butterwort
Pinguicula, see the Butter worts
Pinguicula vulgaris, Linn., see the

Common Butterwort
Pink family, the, 112, 115, 163, 180,

184

Pink, the Carthusian, 113
Pink, the Large-flowered, 112-113
Pink, the Wood, 113

350

INDEX

Pinks, the Alpine, 29, 112-113
Pinus cembra, Linn., see the Stone

Pine
Pinus montana, Mill., see the

Mountain Pine

Pinus picea, Dur., see the Spruce
Pinus picea, Linn., see the Silver Fir
Pinus succinifera, Goepp., 241
Pinus sylvestris, Linn., see the Scotch

Fir

Pitcher Plants, the, 214
Plane, the, 210
" Plant slums, "82
Plantains, the, 281
Platanthera bifolia, Rich., see the

Lesser Butterfly Orchid
Platanus, see the Plane
Pleistocene Period, the, 301
Pliocene Period, the, 301, 303
Poa alpina, Linn., 157
Polemonium coeruleum, Linn., see

the Jacob's Ladder
Pollen and insects, 269-274
Pollen, protection of, 279-282
Polygala chamcebuxus, Linn., see the

Box-leaved Polygala
Polygala, the Box-leaved, 31, 261-

253, 281

cross-pollination of, 252-253, 281
Mediterranean origin of, 251
time of flowering of, 31, 252
Polygala vulgaris, Linn., see the

Common Milkwort
Polygalaceae, see the Milkwort

family
Polygonaceae, the, see the Dock

family

Polygonatum, see the Solomon's Seal
Polygonum bistorta, Linn., see the

Bistort
Polygonum, the Viviparous, 156-157

bulbils of, 156-157
Polygonum viviparum, Linn. , see the

Viviparous Polygonum
Polygonums, the, 150
Pondweed family, the, 205
Pondweeds, the, 205
Poppy family, the, 195
Poppy, the Alpine, 105-196, 272, 295
Potamogeton, see the Pondweeds
Potentilla frigida, Vill., see the

Frigid Potentilla

Potentilla grandiftora, Linn., see the

Large-flowered Potentilla
Potentilla minima, Hall, see the

Smallest Potentilla
Potentilla multifida, Linn., 295
Potentilla nivea, Linn., 295
Potentilla sibbaldi, Haller, see the

Sibbaldia

Potentilla, the Frigid, 197
Potentilla, the Large-flowered, 197
Potentilla, the Smallest, 197
Potentilla, the Spring, 31, 197

time of flowering, 31
Potentillas, the, 115, 128, 158, 187
Potentilla tormentilla, Neck., see the

Tormentil
Potentilla verna, Linn., see the

Spring Potentilla
Primitive soils, 95-97, 170-171
Primrose Family, 33, 61, 64, 72, 181
Primrose, the, 66, 67
Primrose, the Bird's-eye, 31, 67, 69-

70, 300

function of wax on leaf of, 69-70
time of flowering, 31
Primula auricula, Linn., see the

Auricula

Primula elatior, Jacq. , see the Oxlip
Primula farinosa, Linn., see the

Bird's-eye Primrose
Primula glutinosa, Wulf. , 72
Primula hirsuta, All. , see the Hairy

Primula

Primula integrifolia, see the Entire-
leaved Primula
Primula longiflora, AIL, see the

Long-flowered Primula
Primula cenensis, Thorn. , 72
Primula pubescens, 70
Primula, the Entire-leaved, 72
Primula, the Hairy, 31, 70, 72

time of flowering, 31
Primula, the Long-flowered, 67, 71-

72, 296

Primula viscosa, All. , 72
Primula vulgaris, Huds., see the

Primrose

Primulacese, see Primrose Family
Primulas, the, 33, 61, 167, 212, 281,

284
Protections against unbidden guests,

275-277

INDEX

351

Ptarmigan, the, 221

Pyrenees, the, 20, 200, 243, 297, 298

Pyrola chlorantha, Sw., 261

Pyrola media, Swartz, see the Inter-
mediate Wintergreen

Pyrola minor, Linn. , see the Common
Wintergreen

Pyrola rotundifolia, Linn., see the
Larger Wintergreen

Pyrola secunda, Linn., see the
Serrated Wintergreen

Pyrola, see the Wintergreens

Pyrola uniflora, Linn., see the
Single-flowered Wintergreen

Pyrolacese, see the Wintergreen
family

Ragged Robin, the, 225
Rampion, the Round-headed, 91
Rampions, the, 34, 44, 61, 82, 85,

91-92, 150, 152, 192, 272
stylar brush mechanism, 92
the petals, 92
Ranunculacese, see the Buttercup

family

Ranunculi, the Water, 209-210
Ranunculus alpestris, Linn., see the

Alpine Buttercup
Ranunculus aconitifolius, Linn., see

the Aconite-leaved Buttercup
Ranunculus acris, Linn., see the

Field Buttercup
Ranunculus bulbosus, Linn., see the

Bulbous Buttercup
Ranunculus glacialis, Linn. , see the

Glacial Buttercup
Ranunculus parnassifolius, Linn.,

see the Parnassus-leaved Butter-
cup

Ranunculus platanifolius, Linn. , 210
Ranunculus pyrenceus, Linn., see the

Pyrenean Buttercup
Rare Alpine species, 294-296
Rattle, the, 218

Replacements, Alpine, 114, 266
Reserve food materials, 11, 63, 64,

125, 135, 146, 154, 155, 170
Resin, 242
Retreat of the Glaciers, the, 296

Rhsetian Alps, the, 296
Rhinanthus, see the Rattle
Rhododendron ferrugineum, Linn.,

see the Common Alpenrose
Rhododendron hirsutum, Linn., see

the Hairy Alpenrose
Rhododendrons, see the Alpenroses
Rhone Valley, the, 7, 150, 242, 294
Rieder Alp, the, 150
Riffelalp, the, 23, 29, 243
Riffelberg, the, 29
Ring- Ouzel, 221
Robbers, insect, 130, 164, 208-209,

269, 275, 276

Robin's Pin-cushions, 290
Rock Catchfly, see Catchfly, the

Rock

Rocky Mountains, the, 299, 304
Rolled leaves, 110, 141, 188, 222, 231
Rosa alpina, Linn., see the Alpine

Rose

Rosaceae, see the Rose family
Rose family, the, 106, 115, 127, 157,

197, 279
Rose, the Alpine, 22, 91, 220

absence of thorns, 22
Rosette plants, 73, 75, 79, 98-103,

177, 178, 181, 182, 185, 190-192
Rosette plants, the High Alpine,

190-192

Rubus, see the Brambles
Rue, the, 195

Rues, the Meadow, 124, 281
Rumex alpinus, Linn., see the Alpine

Dock

Rushes, the, 278
Russia, 180, 240
Rusts of Wheat, the, 287, 290
Ruta, see the Rue
Rye, 288

Saas Fee, 101, 244, 258

Saas Grund, 150, 179

Saas Thai, 7, 16, 72, 84, 90, 172, 242

Saffron, the Alpine, 165-166

Saffron, the Autumn, see the

Meadow Saffron
Saffron, the Meadow, 152, 156, 165-

168, 281

352

INDEX

Saffron, the Meadow — continued
corm of, 166-167
dimorphism of, 168
fruits of, 168
habit of, 166-167
Sahara, the, 180

Salicaceae, see the Willow family
Salix casia, Vill., 190
Salix glauca, Linn., 190
Salix herbacea, Linn., see the Dwarf

Willow
Salix reticulata, Linn., see the

Reticulate Willow
Salix retusa, Linn., 190
Savoy, 172
Saxifraga aizoides, Linn., see the

Yellow-flowered Saxifrage
Saxifraga aizoon, Jacq., see the

Evergreen Saxifrage
Saxifraga androsacea, Linn., 192
Saxifraga aphylla, Sternb., 192
Saxifraga aspera, Linn., see the

Rough Saxifrage
Saxifraga aspera, var. bryoides, 174,

185

Saxifraga biftora, All., see the Two-
flowered Saxifrage
Saxifraga bryoides, Linn., see the

Moss-like Saxifrage
Saxifraga cassia, Linn., see the

'Glaucous Saxifrage
Saxifraga cernua, Linn., 157
Saxifraga controversa, Sternb., 82,

114
Saxifraga cotyUdon, Linn., see the

Thick-leaved Saxifrage
Saxifraga diapensioides, BelL, 79,

295

Saxifraga exarata, Vill., 192
Saxifraga moschata, Wulf., 174, 185
Saxifraga muscoides, All., 174, 192
Saxifraga oppositifolia, Linn., see

the Purple Saxifrage
Saxifraga planifolia, Lap., see S.

muscoides, All.
Saxifraga rotundifolia, Linn., see the

Round-leaved Saxifrage
Saxifraga seguieri, Spr., 192
Saxifraga stellaris, Linn., see the

Star-leaved Saxifrage
Saxifraga stenopetala, Gaud., see S.

apnylla, Sternb.

Saxifraga varians, Sieb., see S.

moschata, Wulf.
Saxifragaceae, see the Saxifrage

family

Saxifrage family, the, 216
Saxifrage, the Evergreen, 75-77, 79
Saxifrage, the Glaucous, 79
Saxifrage, the Moss-like, 81
Saxifrage, the Opposite-leaved, see

the Purple Saxifrage
Saxifrage, the Purple, 44, 78-79, 178,

179, 185-186, 299, 300
Saxifrage, the Rough, 81-82, 185
Saxifrage, the Round-leaved, 80
Saxifrage, the Star-leaved, 80
Saxifrage, the Thick-leaved, 78, 79
Saxifrage, the Two-flowered, 79, 174
Saxifrage, the Yellow-flowered, 79-

80
Saxifrages, the, 16, 61, 73, 74-83,

185-186, 284

chalk-glands on leaves of, 75-79
Saxifrages, the High Alpine, 185-

186

Scandinavia, 232, 240, 249
Schroeter, Prof., 6, 174, 175, 188,

298

Scotland, 50, 110, 190, 211, 249, 303
Scrophulariaceae, see the Foxglove

family

Seaweeds, the, 77, 292
Sedges, the, 278
Sedum, see Stonecrops
Seeds, 282-286
Seeds, distribution of, 88, 132,

162-163, 189, 220-221, 250-251,

282-286

Selago family, the, 110
Selagineae, see the Selago family
Semiparasites, 218-219
Sempervivum, see the House-leeks
Sempervivum arachnoideum, Linn.,

see the Spider's-web House-leek
Sempervivum funckii, Braun, 295
Sempervivum globiferum, Linn., 102
Sempervivum montanum, Linn., see

the Mountain House-leek
Sempervivum soboliferum, Sims, 102
Sempervivum tectorum, Linn., 98
Sempervivum wulfeni, Hopp. , 295
Senecio carnioliceus, Will., 295
Senecio uniflorus, AIL, 295

INDEX

353

Shade plants of forests, 31, 32, 33,

246-267

Shamrock, the, 263
Shoots specialised to parasites, 288-

291
Shortness of the flowering season, 6,

30, 114, 146, 170
Sibbaldia procumbens, Linn., see the

Sibbaldia
Sibbaldia, the, 197
Sieversia, see the Avens
Silene, see the Campion
Silene acaulw, Linn., see the Common

Moss Campion
Silene cucubalus, WibeL, see the

Campion

Silene exscapa, All., see the Sessile-
flowered Moss Campion
Silene inflata, Sm., see tne Campion
Silene rupestris, Linn., see the Rock

Catchfly
Silene, the Rock, see the Rock

Catchfly

Silene vallesia, Linn., 295
Sils (Engadine), 21-22
Silvaplana (Engadine), 211, 243
Simplon, the, 90
Slugs as flower pollinators, 279
" Slums, plant, * p. 82
Smilacina bifolia, Schult., see the

May Lily
Smuts, the, 288

Snails as flower pollinators, 279
Snap-Dragons, the, 115
Snow, Crocus flowering in, 62, 154-

155

Soldanella flowering in, 62-64
Snow-Finch, the, 221
Snow-line, the, 170, 236
Soil, temperature of, 36, 171
Soils, 34, 38, 47, 72, 77, 117-119, 151-

152, 296

Soils, primitive, 95-07, 170
Soldanella alpina, Linn., see the

Alpine Soldanella
Soldanella pusilla, Baumg,, see the

Small Soldanella
Soldanella, the Alpine, 62-66
cross-pollination of, 64-66
flowering in the snow, 62-64
Soldanella, the Small, 62-66
flowering in the snow, 62-64

Soldanellas, the, 33, 61-66, 155, 280
Solomon's Seal, the, 247, 255
Sphinx, see Hawkmoths
Sprengel, C. K.,2, 269
Spring flowers, early, 28, 81
Spruce Fir, see the Spruce
Spruce, the, 5, 97, 213, 234, 235,

236, 237, 239, 240-241, 242, 244,

246, 247, 248, 289, 291
Spruce, the Norway, see the Spruce
Spurge family, the, 124
Spurges, the, 124, 289, 290
Star-of-Bethlehem, the, 119
State legislation in regard to forests,

238

Steppe region, the, 180
St Gotthard region, the, 78
Stinging nettles, 119
St Moritz, 30
Stonecrop family, the, 98
Stonecrops, the, 103
Stone Pine, the, 234, 242-243
Strawberry, the, 100, 127, 128
Strawberry, the Wild, 285
Streptopus amplexifolius, D. C., see

the Knot-foot
Stylar brush mechanism, 85-87, 92,

126

Subalpine zone defined, 4
Succulent fruits, 220-221, 285
Sundews, the, 214
Sunflower, the, 18, 19
Surenen Pass, the, 120
Sweet-William, the, 113
Symptoms of disease, 287-291

Taraxacum officinale, Weber, see the

Dandelion
Tasmania, 20
Temperature, lowering of, during

the Ice Age, 301-304
Temperature, variations in, 146,

171, 173

Tertiary period, the, 302
Tessin (Ticino) Canton, 90, 165, 182,

295, 296, 299

Thalictrum aquilegifolium, Linn. , 272
Thalictrum, see the Meadow Rues
The"odule Pass, the, 173

354

INDEX

Thesium alpinum, Linn., 281
Thistle, the Carline, 125-126, 281
Thymeleacae, see the Daphne family
Thymes, the, 104, 173, 219
Thymus chamcedrys, Fries., 104
Thymus serpyllum, Linn., 104, 176
Ticino, see Tessin
Toadflax, the Alpine, 44, 118-115
Toadflax, the Ivy-leaved, 114
Toadflax, the Yellow, 114
Tofieldia palustris, Huds., see the

Scotch Asphodel
Tormentil, the, 197
Transalpine Switzerland, 78
Transpiration, protections against

excessive, 13, 19, 25-26, 36, 76-

77, 98-99, 108, 110, 113, 141,

171, 188, 231, 232
Traveller's Joy, the, 41, 229
Treeless higher pastures, 236-237
Trifolium alpinum, Linn., see the

Alpine Clover
Trifolium saxatile, All. , 295
Trollius europceus, Linn., see the

Globe-flower
Tuberous roots, 130, 135
Turk's Cap, the, see the Martagon

Lily

Turpentine, 240, 242
Tussilago farfara, Linn., see the

Coltsfoot

Twayblades, the, 255
Tyrol, the, 20, 203, 297, 298

Ultra-violet rays, 172
Umbelliferse, the, 124
Unbidden guests, 49, 55, 85, 89, 208-

209, 250, 275-277
Unisexual flowers, 13, 104, 113, 123,

128, 163, 165, 180, 189, 232,

267
United States, the flora of the, 299,

301, 303

Uromyces pisi, 290
Urtica dioica, Linn., see the Common

Nettle
Urtica urens, Linn., see the Small

Nettle

Usnea barbata, see the Old Man's

Beard Lichen
Utricularia, see the Bladderworts

Vacciniaceae, see the Bilberry family
Vaccinium myrtillus, Linn., see the

True Bilberry

Vaccinium, see the Bilberries
Vaccinium, the Bog, 221-224
Vaccinium uliginosum, Linn., see

the Bog Vaccinium
Vaccinium vitis-idcea, Linn., see the

Cowberry
Valais, Canton, 36, 79, 90, 116, 165,

175, 182, 183, 242, 244, 294-296
Valeriana celtica, Linn., 295
Valeriana supina, Linn., 295
Vanilla, 135
Vanilla planifolia, Andr., see the

Vanilla Orchid
Veratrin, 122
Veratrum album, Linn., see the

White Veratrum
Veratrum, the White, 39, 58, 120-

124, 177

contractile roots of, 122-123
habit, 120

leaves directing rain, 121
young shoots of, 121
Vetch, the Kidney, 105-106, 200
Vienna, 203
Vine, the zone of, 4
Viola biflora, Linn., see the Two-
flowered Violet
Viola cakarata, Linn., see the

Long-spurred Violet
Viola cenisia, Linn., see the Mont

Cenis Violet
Viola odorata, Linn. , see the Sweet

Violet
Viola tricolor, Linn., see the Field

Pansy

Violaceae, see the Violet family
Violet family, the, 128, 159, 253
Violet, the Long-spurred, 31, 128-

129, 160, 254, 271, 273
Violet, the Mont Cenis, 129
Violet, the Sweet, 162

INDEX

355

Violet, the Two-flowered, 160, 247,

258-254, 270, 271
Violets, the, 88, 124, 273, 278, 281,

285

Visp Thalen, 116
Vogler, Dr, 284

Wales, 303

Water-storage tissues, 71, 98

Weather-glass plants, 125-126

Weissmies Hotel, 179

Wheat, the, 288

White Hellebore root, the, 122

White varieties of flowers, 46

Whitlow-grass, the, 191

Whortleberry, the Red, see the

Cowberry

Willow family, the, 187
Willow, the Dwarf, 178, 190
Willow, the Reticulate, 112, 178, 187-
190

age, 188

flowers, 188-189

habit, 188

nerves of leaves, 188

seeds and their distribution, 189
Willow-herb family, the, 189
Willow-herbs, the Alpine, 189, 284
Willows, the High Alpine, 124, 186,

187-190, 284

Wind distribution of seeds, 41, 88,
108-109, 128, 132, 189, 230, 240,
284-285

Wind-flowers, see Anemones
Wind-pollination, 235, 268, 269, 277-

278

Wintergreen family, the, 261
Wintergreen, the Common, 261
Wintergreen, the Intermediate, 261
Wintergreen, the Larger, 261-262
Wintergreen, the Serrated, 262
Wintergreen, the Single-flowered,
261-268, 300

pollination of, 262
Wintergreens, the, 261-263

fruit of, 263

stamens of, 262-263
Witches' Brooms, 289
Wolfgang (Davos), 243
Woodbine, the, 224
Wood Sorrel, the, 268-266, 278

fruits of, 265

leaves of, 263-264

movements of leaves, 264

movements of chlorophyll grains,
265

Yellowwort, the, 42

Zermatt, 7, 16, 29, 36, 72, 116, 150,

172, 173, 235, 242
Zonal limits, arbitrary nature of,

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is practically a new book. A good deal of fresh material has been added, the latest
statistics have been included and the whole text has been thoroughly overhauled and
re-set, bringing everything completely up to date.

" An excellent and up-to-date text-book. . . . The complete knowledge of the soil and
the part it plays in the nutrition of the plants requires investigation along three Hues,
which maybe roughly classed as— chemical, physical or mechanical, and biological. It
is exactly these witu which the author deals, and although it is in no sense an exhaustive
treatise, a general outline has been given of all the recent investigations which have
opened up so many soil problems, and thrown new light on difficulties that are experienced
in practice." — Gardeners' Chronicle.

A HANDY
BOOK OF HORTICULTURE

AN INTRODUCTION TO THE THEORY AND PRACTICE
OF GARDENING.

BY F. C. HAYES, M.A.,

Rector of Raheny ; Lecturer in Practical Horticulture in Alexandra College, Dublin.

With Illustrations. Crown 8vo. as. 6d. net.

" Not so big that It need frighten the ardent amateur, nor so much of a primer that it
may be disdained by the fairly accomplished gardener, it has a good scheme. The first
part, consisting of eight chapters of general principles, in simple, non-technical language.
is a model of useful information in a small space ; the second part deals with departments
of gardening ; the third, with types of flowers, and the fourth is a calendar to work by."

—Daily Chronicle.

ELEMENTS OF AGRICULTURE

A TEXT-BOOK PREPARED UNDER THE AUTHORITY OF
THE ROYAL AGRICULTURAL SOCIETY OF ENGLAND.

BY W. FREAM, LL.D.

yth Edition. With numerous Illustrations. Crown 8vo. 33. 6d.

THE BOOK OF THE

ROTHAMSTED EXPERIMENTS

BY A. D. HALL, M.A. (Oxon.),

President of the Rothamsted Experimental Station ; First President of the
South-Eastern Agricultural College.

ISSUED WITH THE AUTHORITY OF THE LAWES AGRICULTURAL TRUST COMMITTEE.

With Illustrations. Medium 8vo. IOS. 6d. net.

CONTENTS.

BIOGRAPHICAL INTRODUCTION — THE SOURCES OF THE NITROGEN OF VEGETATION —
METEOROLOGICAL OBSERVATIONS— THE COMPOSITION OF THE ROTHAMSTED SOIL
— EXPERIMENTS UPON WHEAT — EXPERIMENTS UPON BARLEY — EXPERIMENTS UPON
OATS— EXPERIMENTS UPON ROOT CROPS GROWN CONTINUOUSLY ON THE SAME LAND
— EXPERIMENTS UPON THE CONTINUOUS GROWTH OF LEGUMINOUS CROPS — EXPERI-
MENTS UPON GRASS LAND MOWN FOR HAY EVERY YEAR— EXPERIMENTS UPON CROPS
GROWN IN ROTATION, AGDELL FIELD — NITRIFICATION AND THE COMPOSITION OF
DRAINAGE WATERS — THE FEEDING EXPERIMENTS — MISCELLANEOUS ENQUIRIES —
Append ix— INDEX.

JOHN MURRAY, ALBEMARLE STREET, W.

RETURN ENVIRONMENTAL DESIGN LIBRARY
TO—^ 210 Wurster Hall 642-4818

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UNIVERSITY OF CALIFORNIA, BERKELEY
FORM NO. DD1 3, 74m, 3/78 BERKELEY, CA 94720

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Full text of "Plant life in Alpine Switzerland; being an account in simple language of the natural history of Alpine plants"

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