THE

ORIGIN OF THE BRITISH FLORA.

THE ORIGIN

OF THE

BRITISH FLORA

BY

CLEMENT REID,

F.R.S., P^.L.S., F.G.S.,

OF THE GEOLOGICAL SURVEY OF ENGLAND AND WALES.

LONDON :

DULAU & CO., 37 SOHO SQUARE, W.
1899.

HE

Ua

LONDON :
Prin'ed by Straxgkwavs and Sons,
Tower St., Cambridgt Circu$. W.C.

PREFACE

While embodying in this book the results which
I have accumulated during the past twenty years,
I should like to take the opportunity of thanking
the many friends who have assisted me. The
first to do so were Mr. Carruthers and Professor
A. G. Nathorst, whose work, in fact, led me to
undertake these studies. In the troublesome work
of determining the plants I have been greatly aided
by the constant courtesy and assistance of the
officers of the Botanical Department of the British
Museum, especially of my friends Mr. E. G. Baker
and Mr. A. B. Rendle. At Kew also I have
received every facility for the work, and to Mr.
J. G. Baker, the late keeper of the Herbarium, I
owe much. Messrs, G. and H. Groves have also
assisted me at various times with specimens of
recent plants which I was unable to obtain for
myself, and others have been received from
friends whose names are too numerous to mention.
With regard to the geological material that I
have obtained from others, specimens have been

vl Preface,

received from so many sources that I must leave
the reference at the head of each locality to speak
for itself, only acknowledging the special aid that
has been given by Mr. James Bennie, in collecting
the plants of the ancient silted-up lakes of the
Scottish Lowlands. For the constant encourage-
ment of Sir Archibald Geikie, Director-General of
the Geological Survey, I am also very grateful.

CONTENTS.

Chap. Page.

I

I. — Introduction

II. — The Present Flora of Britain
III.— Means of Dispersal ....
IV.— Changes in Geography and Climate

v.— Deposits Containing Fossil Plants.

VI.— Former Distribution of British Plants

Appendix.— Table Showing the Range in Time
OF THE British Flora ....

lO

20

33

48

97
171

THE

ORIGIN OF THE BRITISH FLORA.

CHAPTER I.

Introduction.

In the year 1876, happening to be engaged on the
Geological Survey of East Norfolk, I was led to
commence observation on the plants of the Preglacial
* Cromer Forest-bed.' At first I confined my efforts to
collecting the animals and plants, some of the latter being
afterwards determined by Mr. Carruthers. But it soon
became obvious that, in order to obtain any satisfactory
knowledge of the subject, it was necessary to collect and
study the ripe seeds and fruits of our British plants, and to
devote much of my leisure to the work of comparison ;
fossil seeds had seldom been collected in this country, and
recent plants with perfectly ripe seeds were seldom to be
found in our herbaria.

From a study of the plants of the Cromer Forest-bed,
the work gradually expanded into an examination of any
Newer Tertiary plants that could be found in Britain, and
as during the past twenty years my employment on the
Geological Survey of England has necessitated a close
scrutiny of our Newer Tertiary deposits, especially in the
south and east of England, I have been brought continu-
ally face to face with the problems of the origin of our

B

2 Origin of the British Flora.

fauna and flora, and the relations these bear to the climatic
changes through which this country has passed.

Moreover, this life spent principally in field, and moor,
and forest has forced me to observe how each changing
season is marked by corresponding adaptations in the
animals and plants, such as enable the species to preserve
themselves, to multiply, and to spread ; or, if adaptation
fails at any point, through some climatic irregularity, how
sweeping and rapid may be the extermination of all except
some few accidentally favoured individuals. While col-
lecting seeds and fruits for comparison with the fossils
I was compelled particularly to observe their many
adaptations for dispersal, and also their times of ripening,
and the abundance or scarcity of ripe seeds.

It was impossible under such circumstances to avoid
seeing the close connexion which must exist between the
present geographical distribution of plants and animals
and bygone changes in climate and in physical geography.
Edward Forbes' * essay was read and read again ; but it
soon became apparent that his brilliant generalisations,
though far in advance of the date when they were written,
were only partially true. Much of his reasoning was
fallacious.

To explain the presence of Arctic and of Iberian plants
in Britain, he showed that outliers of the Arctic flora stranded
on our mountain peaks could be accounted for by an
appeal to the climatic conditions of former days, when a
similar flora covered the whole of our Islands, and was not
confined to isolated mountains. He did not see, apparently,
that the use of this reasoning precluded the use of the

* 'On the Connexion between the Distribution of the existing
Fauna and Flora of the British Isles, and the Geological Changes
which have affected their area, especially during the epoch of the
Northern Y)x\iV—Mem. Geol. Survey^ Vol. I., pp. 336-432 (1846).

Introduction. 3

converse hypothesis of a warm climate continuous from
Preglacial times to account for the Iberian plants in the
west of Ireland and in Cornwall. Either might be true,
but scarcely both ; for the Irish and Cornish plants are not
such as could survive a colder climate like that postulated
by Forbes to explain the migration of the Arctic species.
We have obtained direct evidence, since Forbes wrote,
that all Ireland was at one time strongly glaciated, and
also that Arctic plants once occupied the lowlands of
Devonshire.

This problem of the origin of our flora is one which can
be solved, I think, by the historical method, and that
seems to be the proper mode of attacking it. No doubt
the imperfection of the geological record is so great as to
make the task an exceedingly difficult one ; for nowhere
have we yet discovered a continuous sequence of deposits,
all fossiliferous, such as would give a connected history of
our recent animals and plants from their first appearance in
Britain to the present day. The exact order of succession
of the deposits, of the physical changes, of the climatic
alternations, and of the waves of migration, is still uncer-
tain ; though a definite historical record is gradually being
built up by the comparison and correlation of numerous
overlapping chronicles, each recording at most some three
or four of the subordinate stages or periods. This work of
correlation, as already mentioned, has been greatly
facilitated by a detailed examination of extensive areas,
and a close study of the geology of the more recent deposits.
In this way I have been enabled to trace the connexion
between the strata, and often to speak with confidence as to
the date of groups of fossils which otherwise would have had
to remain as isolated finds. My own researches have been
largely aided and supplemented by the examination of
material obtained from friends working in districts which I

4 Origin of the British Flora.

have had but slight opportunity of studying. This has
especially been the case with regard to the lacustrine
deposits of the Scottish Lowlands, so minutely examined
by Mr. James Bennie. The results of these investigations
will be found summarised in Chapters IV. and V. of this
work.

In the examination of our recent flora I have looked
at the plants mainly from the point of view of the field-
naturalist. Their climatic and geographical distribution ;
the periods of ripening, and the means of dispersal of their
seeds ; their competition with other plants ; and their
dependence on, or destruction by animals, were the circum-
stances especially noted — more so than critical distinctions
of varieties and sub-species. Not that these distinctions
are considered unimportant, but mainly because of the
difficulty of studying them without a complete herbarium,
too heavy to transport during constant changes of station.
Moreover, botanists have almost ignored the essential
distinction between a varietal form due to local conditions,
and a true sub-species or race ; for many of our named
sub-species have evidently no more claim to such rank than
have luxuriant garden specimens. Forms, for instance, of
the water-crowfoot {Ranunculus aquatilis) or of the lesser
spearwort {R, Flammuld) growing in a well-manured horse-
pond or ditch have no claim to rank as sub-species, unless
they can be found also under more natural conditions, and
come true from seed. Again, the prostrate maritime form
of broom found in Cornwall {Cytisus scoparius, var. pro-
stratus) has similarly no claim to varietal rank, for Mr.
Mitten tells me that seeds gathered by him grew in his
garden into the common erect form of broom. A botanical
visit to the Dingle Promontory, in Kerry, in company with
Mr. Edmund Baker, produced several instances of this
sort. We examined Saxifraga umbrosa and its allied

Introduction, 5

forms, of which we found several, each occupying well-
defined small areas, and apparently possessing definite
characters. But, as more and more of the patches were
examined, these distinctions were found to melt away ; for
each fresh patch yielded a slightly different form, so that
finally we were able to obtain a nearly complete series of
intermediates seeming to connect the extreme kS. umbrosa
with the extreme 5. Geum, all of them living within a
small area under similar conditions. Pinguicida vulgaris
and P . grandiflora, on the other hand, we found growing
together in abundance, and quite distinct except at one
spot where, below a rock on which both grew, we found a
number of hybrids. In this case the allied forms, some-
times only ranked as sub-species, are both good species, and
have different geographical distributions, though they over-
lap at more than one point. Botanical books are full of
similar anomalies, often due to a natural desire to announce
the discovery of a form new to Britain ; but for the student
of geographical distribution varietal names founded on
such material are worse than useless. For they tend to
confound sub-species, which, if found in isolated areas show,
in all probability, a transportation of the seeds from one
to another, with varieties or forms, which will reappear
wherever the parent species is subject to particular con-
ditions.

A flora like that of the British Islands may be studied
in so many different ways, that it will be well to define at
once the standpoints from which it is viewed in the following
pages. I do not propose, nor do I feel competent, to
touch on the questions of the evolution of the species, or
of their relationship to each other ; what will be attempted
in Chapter 11. is, to give a sketch of the existing flora as
a whole, to note its composition, and the distribution of
the species. Chapter III. will deal with the means of

6 Origin of the British Flora.

dispersal of the various species which constitute our flora,
with special reference to the present and past distribution
of the plants. Finally, I propose to give an historical
account of each species as far back as geological evidence
will yet allow it to be traced.

It may be considered presumptuous to attempt such a
task ; but, though the following Chapters are most imper-
fect, yet they may do good by directing attention to lines
and methods of research which are as yet little appreciated.
The section on the geological history of our flora, being a
record of the actual distribution in space and time of our
plants from direct observation, will perhaps be the one to
which botanists will most readily turn. It may be sug-
gested, however, that the section on means of dispersal is
equally important, and that the connexion between the
difi*erent Chapters is so close that it is impossible properly
to appreciate the relationship of the living plants to their
fossil representatives without a study of the subject from
various points.

Though the present volume is professedly occupied with
a discussion of the origin of the British flora, it should not
be forgotten that in questions of geographical distribution
it is impossible to separate animals from plants, for many
plants are directly dependent on certain animals for means
of dispersal. Moreover, certain animals are dispersed by
the same means as flowering plants, have the same difficul-
ties to contend with, are no less dependent on climatic
conditions, and are almost equally tied to a single spot
during the lifetime of the individual. The land mollusca
in particular are in these respects so like the more sedentary
species of flowering plants that I have not hesitated to
speak of them where they help to illustrate the subject
under consideration. Beetles, I believe, would also be of
use ; but of this order I have unfortunately no knowledge,

Introduction. 7

and at present few of the numerous fossil species occurring
in our Pleistocene deposits have been determined. Fresh-
water mollusca, freshwater fish, and amphibia seem to obey
the same laws of geographical distribution as aquatic
plants : the species are usually of wide range, provided the
barriers are not excessively broad or high, and the
climatic conditions are suitable.

The geological sketch has been greatly condensed ; for
it is obviously impossible to deal with so complicated a
subject in a limited space, and all that can be done is to
give some indication of the climatic conditions, local
peculiarities, and character of the flora at each spot where
plant-bearing deposits are found. The thorny subject of
bygone alternations of climate is perforce discussed, for
it lies at the root of our inquiry, I have also been obliged
to deal with another equally vexed question, the submer-
gence or elevation of the land in Pleistocene times; for
this obviously has a most important bearing on the possible
survival of plants within our Islands. In discussing the
past climatic changes, while giving the preference to the
evidence derived from remains of plants belonging to
existing species, I have not hesitated to supplement this by
an appeal to other groups of organisms, or to inorganic
geology ; for an assemblage of Arctic mammals, a group
of Arctic or desert mollusca, a morainic deposit, or erratics
brought by floating ice in an Arctic sea, are as good
evidence of climate as a group of plants, and are often
discoverable in strata in which no plants are preserved.

Perhaps it will be asked why, if the British flora is to
be treated from standpoints which involve a consideration
of climatic and geographic changes such as cannot be
merely local, a still wider view is not taken, and this flora
dealt with as a mere outlier of the Palaiarctic one } To
this I may reply, firstly, that the fossil plants of the periods

8 Origin of the British Flora.

dealt with are at present almost unknown outside Britain,
Sweden, and North Germany, and speculation would have
to take the place of an appeal to direct evidence. Secondly,
that Britain is not by any means simply an outlier of the
continent of Europe. Its flora is an insular one of peculiar
character, unlike that of any part of Europe, and unlike
that of an oceanic Island. Few, if any, of the species are
confined to Britain ; but the Islands contain a selection of
the continental species best adapted for dispersal, and best
able to hold their own in a changing climate. Britain,
within the lifetime of existing species, has been subjected
to many fluctuations of climate, which have left their mark
on the flora. On these fluctuations was superimposed a
series of orographic changes, such as must have tended
greatly to modify local conditions, and must sometimes
have aided, sometimes have hindered, the dispersal of the
seeds.

The following pages deal, therefore, with an insular
flora of exceptional type ; in the building up of which
selection and sweeping extermination have played so
vigorous a part, that the flora now consists largely of an
assemblage of the more readily dispersed of the Palaearctic
species. Time has not permitted any large amount of
variation or formation of sub-species in these Islands ; and
in this our flora is totally different from the more ancient
floras of oceanic islands, which were beyond the reach of
such violent climatic fluctuations as have afiected Britain.

There is one point which needs explanation before we
proceed further. I have been obliged in the following
pages to go back to the popular and original use of the
term ' seed.' . Of the two senses the popular one seems to
be by far the most useful scientifically, for it refers to the
thing that is sown, not to an embryo with or without

Introduction. 9

certain appendages and coverings, which in function may
be quite indistinguishable from others belonging to the
fruit. A seed, therefore, for our present purposes is the
one-seeded unit of dispersal. All our British fruits, with
the single exception of that of the Cornel, divide into such
one-seeded portions, which tend to be dispersed separately,
so that the young plants do not interfere with each other.
These units may be seeds in the strict botanical sense, or
they may be complete one-seeded fruits ; sometimes they
are stones or carpels, one-seeded, or at any rate with only
one of the seeds properly developed ; in other cases they
include the dried calyx, or other parts of the flower or
receptacle. Constant explanation would be needed if an
attempt were made to define botanically what part of the
fruit is referred to in each case — it is more convenient to
accept the perfectly understood popular usage.

lO

CHAPTER II.

The Present Flora of Britain.

When the British Flora is carefully studied, it is found
to be composed of numerous elements, and can be divided
into several well-marked groups. The grouping of the
species, however, varies according to the point from which
they are viewed. Disregarding purely botanical affinities,,
which are not under consideration in this volume, the
assemblages necessarily differ according as the flora is
looked at from the standpoint of relationship of the plants
to climatic conditions ; or from the standpoint of habitat^
including variations in soil, and shelter ; or again, from
that of local distribution. No one of these methods will
enable the plants to be grouped into 'provinces' satisfactory
for all purposes. Each set of conditions overlies and
modifies the distribution which either of the others alone
would tend to bring about.

If we begin with the broadest classification, that based
on climatic conditions, we find at once that this is not
merely a question of average, or of extreme temperature.
It is temperature plus amount of moisture, modified in
various ways by the season at which the rain falls, the
amount of sunshine, and the season at which the sun is
felt. A flowering plant has varying needs at different
seasons ; and the satisfying of these is so essential to the
existence of the species — not necessarily, I would remark,
the same thing as essential to the existence of the in-
dividual— that, if the conditions are unfavourable for any

The Present Flora of Britain. ii

one of them, the plant cannot maintain itself. The seed
must have the right temperature, soil, and amount of
moisture to enable it to germinate and grow. The young
plant must have sufficient vigour to defend itself against
parasites or aggressors — not like the wheat which cannot
grow among our ordinary weeds, and depends on human
protection. The climatic conditions at the time of
flowering must be favourable, or the ovule may not be
fertilised. For the ripening of the seed a certain critical
temperature must be reached, and maintained for a
sufficient time. The cold or wet in the winter must not
be such as to destroy the seed before it has germinated.
All these conditions must be favourable or the plant can-
not establish itself An annual plant must seed every
year, and go through the whole round safely, or it will be
destroyed. A perennial plant need seed and grow from
seedlings only once in a generation.

As instances of what is meant by these remarks I will
take a few common plants. The horse-chestnut grows
well even as far north as Bergen in Norway, and in Britain
it produces abundance of ripe seeds every year ; but even
in the south of England, as far as I am aware, it never
succeeds in establishing itself from self-sown seeds. The
common elm {Ulmus campestris), on the other hand, in
England only produces perfect seed about once in forty
years. Forty years is far less than the lifetime of an elm,
and if the tree seeds once in a lifetime, and the seed
germinates, the species may establish itself. Perfect seeds
have not come under my observation, and I cannot there-
fore say whether this elm does grow from seedlings. It
is generally said only to occur where planted. The
butcher's broom {Ruscus aculeatus) is an instance of a
plant which just manages to hold its own. After watching
its fruiting for twelve years in succession, I find that as a

12 Origin of the British Flora.

rule only about one plant in fifty produces any fruit, and
these are not only few in number, but, as they ripen in
November, an early winter may prevent them ripening
at all. The plant being perennial and hardy can survive,
but it has evidently reached its northern limit in Britain.*
The sycamore, maritime pine, and common rhododendron
^R. ponticuni) are instances of plants undoubtedly intro-
duced, which seed and grow freely from seedings in the
South of England. That they were not till lately members
of our flora is evidently due to geographic, not to climatic
conditions.

We cannot point to any British annuals which do not
seed freely in some part of the Islands, for the sufficient
reason that an annual which cannot seed well may be
entirely exterminated by a single exceptional season.
This points to a probable explanation of the curious
tendency noticed in the floras of small oceanic islands,
for genera ordinarily annual and herbaceous to be repre-
sented by perennial species. This may be explained in
the following way. In many annual plants a few in-
dividuals become biennial ; these in an island devastated
by an exceptional gale at flowering time, by a swarm of
locusts, or other adverse conditions, would be the only ones
to survive, and natural selection would thus tend to
perpetuate the biennial or perennial forms which so
characterise these islands. This change of annual into
perennial forms, however, in all probability has had little
effect on the British plants ; for the Islands, besides being
too large, are sufficiently close to the Continent to receive
occasional seeds or pollen of the same species, which by
intercrossing would tend to keep the species true..

* The exceptionally warm and dry summer and autumn of 1898,
however, caused Ruscus to fruit so freely in Hampshire that I counted
upwards of forty ripe berries on each of several plants.

The Present Flora of Britain, 13

Climatic conditions cause two very distinct floras to be
represented in Britain. The lowland flora is in the main
the temperate flora of the neighbouring lowlands of
Belgium and France. The upland flora, on the other hand,
consists of numerous more or less isolated outliers of the
flora which overspreads the lowlands of the Arctic Regions
and occupies the mountains of Scandinavia. This latter
assemblage is found at higher and higher elevations as it is
traced southvvard, and is confined to hills sufficiently high
to have an average temperature approaching that met with
at the sea-level within the Arctic Circle. As the fall of
temperature is about i^ Fahr. for every 300 feet of elevation,
a sub-arctic climate is found over a considerable area in
Scotland, and on a certain number of isolated hills in
England, Wales, and Ireland. The seeds of the British
Alpine plants are invariably small and usually very minute,
a peculiarity that will be again alluded to.

Local conditions govern the distribution of large groups
of species. First, there are the sea-coast plants, which are
all confined to a narrow belt near the sea. This flora is
very uniform throughout Britain, though some of the
species are found only on the south coast and a few only
on the east.

The seeds of maritime plants are of various descriptions,
and often of large size. Many of them are scattered far
and wide by the sea, though the plants only establish
themselves where a suitable habitat occurs. Thus the
sea-coast flora includes a good many plants like the sea-
kale {Crambe maritima), which tend to appear sporadically
wherever the habitat is suitable and to disappear again after
a few years — as though dispersal were easy, and the range
of the species was limited by climatic rather than by other
considerations. Many of the sand-dune or shingle-beach

14 Origin of the British Flora.

species are more properly desert plants, and are only-
confined to the coast because in Britain we have no other
suitable regions. -^

The aquatic flora consists largely of species of wide
range, which have a remarkable power of reaching isolated
rivers, lakes, or ponds. Though some of these species are
confined to limited areas, most of them tend to re-appear
wherever the local conditions are favourable. They are
apparently more limited in their northerly range by un-
favourable climate than by diflliculty of crossing barriers.
Several of the aquatic plants of limited range are almost
confined to the East Anglian broads and rivers ; but this
limitation is evidently due to the more extensive and
connected waterways of that district, rather than to other
conditions. Not one of our aquatic plants is a member
of the Alpine flora, or belongs to the Lusitanian group
found in Cornwall and in the West of Ireland.

Among the marsh and peat-moss plants are many of
which the local distribution is evidently governed by
climate and geographical position, and is not dependent
on soil or amount of rainfall. A large group of these
plants consists of upland forms, such as the Arctic willows
and sedges. Another set is confined to the Eastern Counties;
though these are few in number, notwithstanding the large
area of swampy ground there found. A third group is
confined to the South-west of England, or to the West of
Ireland.

The anomalies in the distribution of our peat-moss
and marsh plants are very striking, especially as this flora
probably has been less aflected by human agency than
any other, except the Alpine. Man may have drained
a certain number of swamps, and thus exterminated some
species, principally in the Fenland ; but it is not probable
that he has had much to do with the introduction of new

The Present Flora of Britain. 15

species, or the transfer to other widely separated localities
of species already in Britain. Marsh plants, of all the
groups, are the least likely to be introduced accidentally
or on purpose by man.

Many of the heath or barren-land plants might be
classed equally well as marsh species, for gravelly or
sandy areas tend to become peaty and waterlogged in
our climate. The most marked characteristic of this flora
is the occurrence in it of certain gregarious plants, which
occupy definite areas in enormous profusion, though
entirely absent from others equally suitable. Several of
our heaths, for instance, are very local, though all of them
occur abundantly where found at all. The British plants
which have a marked western geographical distribution
within the Islands nearly all belong to the marsh and
heath groups.

Of the other open-land groups, that belonging to good
soil and clayey meadows is surprisingly restricted, and
many of the species are probably late introductions. It
is not difficult to see the reason why we have so few
species characteristic of our wide areas of clayey pasture.
These, till recent times, were woodland, not open prairie,
and since the destruction of the woods they have been
under cultivation or closely grazed. We have therefore
nothing equivalent to the prairie vegetation of North
America or other drier climates. Several plants confined
to the eastern counties belong, however, to this group ; for
there the dry cutting winds of winter probably always
prevented the forest growth from extending to the sea,
even where the soil was richest. The other meadow
species have generally a wide range throughout Britain,
wherever the climate is suitable.

Our woodland plants are extremely difficult to deal
with, partly on account of the wholesale destruction of the

1 6 Origin of the British Flora.

ancient forests, partly because of the extensive planting,
which has introduced trees belonging to other districts
and has profoundly modified our woodland flora. To take
one or two instances, the Hornbeam is one of the principal
ancient trees of Essex and other south-eastern counties;
but in the New Forest it only occurs sporadically, near
houses and villages, and such would seem to be its ordinary
mode of occurrence in most parts of Britain. We cannot,
however, say positively that it can only be reckoned as
indigenous over a certain limited area, though the evidence
points in that direction. The Scotch Pine is equally
doubtful, for it was abundant throughout Britain when
our existing peat-mosses began to form ; it afterwards
disappeared throughout the south of England ; but now
that it has been re-introduced it seeds freely and is fast
spreading, especially in Hampshire and Dorset. It is
probable that as far back as Roman times trees were
planted round the villas for shade and beauty, and Roman
officers would probably have given preference to southern
forms which reminded them of their native lands. Thus
such trees as the Horse Chestnut, Spanish Chestnut,
Sycamore, Lime, and probably the Vine and Fig-tree,
would be introduced. Some of the trees died out, others
established themselves from seedlings and still remain;
but except through the negative evidence of the geological
record there seems to be no satisfactory way of telling
which of our rarer trees were thus introduced.

Besides the forest-trees, we have a large number of
plants which are confined to woods ; we have also several
species of land-snails, which are similarly restricted to
ancient forest and are not found in modern plantations.
The moisture and shelter of our woods make the general
character of the undergrowth fairly uniform throughout
Britain ; though we possess a large number of woodland

The Present Flora of Britain. ly

plants which are confined to a few widely separated
localities. Some of the Liliacece and Boraginece^ for
instance, though abundant where they occur, are curiously
local, most of them being absent from extensive areas
apparently as well suited for their growth as those in
which they are found. In the altered state of our woods
these anomalies are particularly difficult to understand,
for the plants usually do not appear to group themselves
into assemblages confined to special districts, and the
distribution of each species has to be studied separately.
Not one of our woodland mollusca or plants, unless the
Arbutus be reckoned as a forest species, falls into the
special groups confined to the eastern counties, to Cornwall,
or to the West of Ireland. It is a question whether the
absence of Lusitanian woodland species may not be due
merely to the destruction of forests in Cornwall and in
the West of Ireland; but this cannot be determined till the
sub-fossil plants of the forests buried under the recent peat
in these districts have been collected and examined. It is
possible that some of the difficulties may be cleared up
when we have studied each patch of ancient woodland,
however small ; for by searching small isolated patches of
old forest we can often find outliers of the sedentary wood-
land mollusca and plants, such as probably once extended
over wide areas now bare or under cultivation.

A certain number of our plants are confined to lime-
stone rocks or to calcareous soils ; but it will be sufficient
here to remark that none of them is characteristically
eastern or western, and that scarcely anything is yet
known of any of them in the fossil state.

In addition to these classifications according to climate
or habitat, there is yet another, certain species being
eastern and others western. Though we have a con-

C

1 8 Origin of the British Flora.

siderable number of plants which are confined to the
Eastern Counties, they, or at any rate the majority of
them, have not a correspondingly eastern distribution on
the Continent, and so many of them occur throughout the
greater part of Europe, that the present local distribution
in Britain may be, after all, climatic rather than geo-
graphical. The Eastern Counties are considerably drier
and more sunny than the others, in this agreeing more
nearly with the mainland of the Continent.

Our western plants, on the other hand, are very
peculiar, for we find in Cornwall and Devon, and also in
the West of Ireland, groups of plants characteristic of the
Pyrenean region. These plants occur usually not as
rarities but in profusion, so that in parts of the West
of Ireland the common species which carpet the hill-sides
are Iberian forms unknown elsewhere in Britain. There
is also another peculiarity which must be taken into
account when we discuss the origin of these outliers —
though Pyrenean plants occur both in the south-west of
England and in the West of Ireland, the species found in
the two districts are not the same. Thus Cornwall pos-
sesses two of the Pyrenean heath-plants. Erica ciliaris
(another outlier of which occurs in Dorset) and Erica
vagans; while the four found in the West of Ireland,
Erica Mackayi, Erica mediterranean Dabeocia polifoliay
and Arbutus Unedo, are all different from the Cornish
ones. The only western plants common to the two
regions are three spurges, two of which are sea-coast
species. Nearly all the Pyrenean plants found in the
British Islands, including the only tree belonging to this
group, have minute seeds, the numerous large-seeded trees
and plants which are associated with them in Spain not
extending into Britain.

Three American plants also occur in Ireland, but the

The Present Flora of Britain. 19

distribution of these is too peculiar to permit of any
attempt at explanation in the present state of our know-
ledge as to the former range of these species. Spiranthes
Romanzoviana occurs in Cork, and in North America
and Kamtschatka ; Sisyrinchium angustifolium is found
in bogs in Galway and Kerry, and also in Arctic and
Temperate North America ; Eriocaulon septangulare is an
aquatic plant occurring in Skye and the West of Ireland,
and also in North America.

From the above notes it will be seen that Britain shows
signs of a geographical distribution of plants largely in-
dependent of that due to climate; or, perhaps we should say,
not governed by existing climatic conditions. The cause of
these peculiarities will be best discussed when we have
examined into the means of dispersal possessed by dif-
ferent plants ; but it will be as well at once to say that
the subject is beset with difficulties, and at every turn we
meet with instances of anomalous distribution, such as
make a botanist inclined to suggest 'accidental introduc-
tion by man ' were it not that many of the species are
marsh or woodland forms, long established and most un-
likely to be brought by human agency in any form. Per-
haps future research may show that many of the outliers
were once less isolated, and that the present distribution
is not so unaccountable as it seems. Such has already
been shown to be the case with many mammals and
mollusca, which geology proves had once a much wider
distribution ; but the flora of our Later Tertiary deposits
has not yet been collected and studied so thoroughly as
has the fauna.

20

CHAPTER III.

Means of Dispersal.

When the adaptation of plants for dispersal is spoken
of, one thinks of winged seeds, or of clinging burrs, of
floating nuts, of succulent fruits which tempt birds, or of
other obvious adaptations. These, however, form only a
few of the contrivances made use of by nature to aid
plants to hold their own and to extend their range. On
considering what is necessary to the existence of a species,
it soon becomes evident that modes of dispersal that seem
to be merely accidental really depend on some modifi-
cation of the seed or plant. They are often alternative
methods without which the very life of the species would
be in danger.

No plant of the Temperate Regions — I do not speak
of Tropical species — would be likely to hold its own for
long periods if it were confined to a single station. The
sweeping climatic waves which time and again have
passed over our latitudes within the life-time of the exist-
ing species must have compelled every one now found in
Britain to move. When deep snow and ice smothered our
uplands, the Alpine flora had to descend to the lowlands ;
when a warmer climate returned, the Arctic plants had to
leave the low ground and again climb the heights The
lowland plants, on the other hand, with few exceptions,
had to leave the country when the Reindeer, Arctic Fox,

Means of Dispersal. 2 1

and Lemming inhabited Salisbury Plain, and the Arctic
Birch and Bearberry grew in the lowlands of South Devon.
The Temperate flora has returned again ; but the fact that
the whole, or nearly the whole, of our plants have been
compelled at least twice, probably many more times, to
migrate long distances, shows that the British flora as it
now exists must be a flora highly specialised for dispersal.
In this respect it is probably more specialised than any
tropical flora, which has been developed in an unvarying
climate, but under a struggle for existence more violent to
the individual.

We should expect to find, therefore, that the British
flora consists of a selection of the more mobile plants of
Europe, without the accompanying sedentary forms. As
the best illustration of what is meant, we may take the
proportions of plants with minute seeds and of plants with
large seeds to the total number, in orders represented both
in the flora of Britain and in that of Europe ; the numbers
not including plants that have seeds, either large or small,
modified in special ways for dispersal over long distances.
The approximate percentages are as follows : —

Percentage

Percentage

in Britain.

in Europe.

Targe seeds

24-5

31-3

Small seeds

... 17-6

12-4

The composites, which at first sight appear to form an
order particularly adapted for dispersal, constitute, how-
ever, a much smaller proportion of the British than of the
continental plants. This, I believe, is due to the general
deficiency in our flora of prairie vegetation — the majority
of the composites are prairie species, and until the last
thousand years Britain, while possessing a temper-
ate climate, was mainly woodland, so that there

22 Origin of the British Flora.

was only comparatively small area suited to their
needs.

Before studying more minutely the means of dispersal
available, it may be well to ask, in this connexion, what
are the requirements that are usually essential to the life of
the species. In the first place, it is necessary that the
seed should be sown beyond the limit of the patch of soil
exhausted by the parent plant. For this a very slight
mobility is requisite. Secondly, in the case of British
plants, some method is ordinarily needed by which
they are enabled to cross barriers, such as rivers or
straits, or tracts of desert in which the plant cannot
flourish.

I use the term ' desert ' as implying areas unfavourable
to any particular species. A desert from the human
standpoint is a sandy waste without water, which is
unsuitable for the plants and animals useful to man.
Such an area may be ga}" with flowers, and is no desert to
the Gorse or Horned Poppy — the desert to them is the
luxuriant meadow or forest, which they cannot overpass
unless their seeds are carried by some rapid messenger.
To a water-plant the dry land is a desert ; to a mountain
plant the lowlands are desert ; to the lowland plants the
mountain is a desert ; and to go further, to certain plants
everything but limestone rock is a desert. Consequently
the British Isles consist not only of an Archipelago with
numerous islands, but from the points of view of different
plants the area forms quite different Archipelagos, of low-
lands with scattered mountain tops, of non -calcareous
country with isolated limestone, or of dry land with scat-
tered lakes.

In gregarious plants, such as heaths and rushes, the
necessity for scattering the seeds beyond the shadow of,
and beyond the soil exhausted by the parent species, may

Means of Dispersal, 23

mean that only the outer individuals of each cluster,
presumably on the average those that have already been
selected by the dispersing agency, have much chance of
propagating themselves. In the case of small-seeded gre-
garious plants like the heaths, without highly specialised
means of dispersal, this difficulty probably tends to keep
the seeds small and chaffy, so as easily to be scattered by
the wind. The berry-bearing heath-plants on the other
hand, though equally gregarious, have seeds fewer, larger,
heavier, and with thicker walls. These latter have been
modified for dispersal by birds. The small-seeded heaths
without special adaptation for dispersal are often singularly
local ; though occurring in profusion, they tend to
occupy widely separated areas, and are absent from
other districts equally favourable. The berry-bear-
ing species are of more general occurrence in suitable
localities, though individually they may not be so
abundant.

Other species have special methods of throwing the
seeds beyond the shadow of the parent plant. The Gorse,
Wood-sorrel, Geranium, and Spurge forcibly eject their
seeds from the ripe pod or capsule. The acorn is attached
lightly for some time after it is ripe, and grows at the end
of a thin branch which, lashed by the October gales, flings
the acorn as boys throw clay-pellets from the end of a
switch. Many umbelliferous plants have a similar mode of
scattering their seed ; for when ripe the carpophore splits
and the seeds hang loosely by their upper ends to the two
whip-like filaments. At the same period the withered
plant hardens and becomes very elastic, so that any
passing animal causes it to spring back and throw off the
seeds, which unless thus scattered, tend to hang on till
they decay. This process one can study in a patch of
these withered umbellifers, part of which is accessible to

24 Origin of the British Flora.

animals, and part of which is cut off by a fence so that it
has remained undisturbed. Umbelliferous plants which
possess burrs, however, behave quite differently. They are
less tall and springy, and, like other plants with burrs, are
so arranged as to scrape the burrs against any passing
animal, but usually not to fling them.

Many plants have capsules so arranged as to scatter the
seeds when forcibly disturbed, but not otherwise to drop
them. The Poppies, Wild Hyacinth, Henbane, and various
caryophyllaceous plants, have capsules erect in fruit and
opening above, and the stems become stiff and elastic when
the seeds are ripe. In some plants such as Erodium, the
seed can actually crawl away from the parent. Certain
trees, such as the Ash, Maple, Hornbeam, and Pine possess
winged fruits which when detached by a breeze tend to be
carried short distances, clear of the shadow of the parent,
though the seed itself is of considerable weight. They com-
bine in this way the advantages of a large embryo, which
gives the young plant a copious store of nutriment to draw
from while it is competing with the short herbage, with a
seed sufficiently mobile to reach places where it can obtain
sunshine and new soil.

The majority of our plants, as already remarked, have
other means of dispersal, which will enable the species
occasionally to overleap barriers — a faculty very different
and probably far more important than the slow spreading
over short stages that has just been spoken of. Here it
may be pointed out that this conquest of the land foot by
foot or yard by yard is insufficient to account for the
present distribution of our flora. It cannot surmount
barriers, and will not account for the mode of occurrence
of such a plant as Erica ciliaris^ which occupies in profusion
two compact areas, one in Cornwall and one in Dorset,
and has every appearance of spreading in each case from a

Means of Dispersal. 25

single seed accidentally transported from some distant
region. The British flora is full of anomalies of this sort.
I may also point out as a geologist that sufficient time
cannot be allowed for this method of spreading, even on
the unwarrantable supposition that our plants could find a
continuous belt of suitable country all the way from
Central Europe, or whatever country they were obliged to
take refuge in during the Glacial Epoch, to the furthest
point they have now reached. Though the Postglacial
period counts its thousands of years, it was not indefinitely
long, and few plants that merely scatter their seed could
advance more than a yard in a year ; for, though the seed
might be thrown further, it would be several seasons before
an oak, for instance, would be sufficiently grown to form a
fresh starting point. The oak, to gain its present most
northerly position in North Britain after being driven out
by the cold, probably had to travel fully six hundred miles,
and this without external aid would take something like a
million years. I doubt whether anything like this time
has elapsed since the Arctic flora occupied the lowlands of
the south of England and the reindeer inhabited Central
France.

Most of our plants have special adaptations for dispersal
over long distances, and, as the different modes of trans-
portation must necessarily lead to different geographical
distributions in different orders, a classification of plants
and animals founded solely on method of migration ought
to throw much light on some obscure problem in geo-
graphical distribution. I am afraid, however, that at
present we have not sufficient direct evidence and can only
speak in a general way of these facilities; though new
observations are made from day to day, and Darwin
collected a large body of evidence on this subject.* The
* Orison of Species^ 6th edition, pp. 323-330.

26

Origin of the British Flora,

main directions in which British plants are specially adapted
for dispersal are the following : —

Modification.

Abundance of minute seeds
(Heaths, Rushes, Saxi-
frages, Caryophyllacese,
&c.).

Abundance of large edible
seeds (Oak, Pine, Horn-
beam, Ivy, &c.).

Edible fruits with hard
stones (Blackberry, Haw-
thorn, Holly, Arbutus,
&c.).

Winged seeds (many Com-
posites, Willows, &c.).

Winged seeds with lax
hairs ( Willow - herbs,
Willows, Bulrush, &c.).

Burrs and hooked seeds.

Floating seeds.

Cut- leaved submerged water-
plants ( Water - crowfoot,
.Water-milfoil, &c.).

Mode of Dispersal.
Readily moved by accidents
of all sorts.

Eaten or dropped by birds ;
most are destroyed, but
some are transported un-
injured.

Eaten by birds and mam-
mals ; seeds passed unin-
jured.

Transported by wind.

Cling to feathers or fur.

Transported by water.

Collapse and cling when re-
moved from the water ;
stems fragile, and broken
pieces grow. Carried on
legs of mammals or of
wading birds.

The first group, the minute-seeded plants, is a very
large one, and it will readily be understood that the plants
belonging to it include nearly all the British species which
show strikingly anomalous distribution. Nearly all of our
Alpine plants, of the Lusitanian species found in Ireland

Means of Dispersal. 27

and Cornwall, and of the peculiar eastern-county plants
belong to this group, the larger seeded species found
associated with them on the Continent being absent.
These plants seem therefore to possess in a pre-eminent
degree the power of crossing seas like that which separates
Ireland from the Pyrenees. They are probably trans-
ported freely by migrating birds, either on their feet or in
their feathers ; but the moist-soil species must also have
been carried in profusion in the cakes of mud which adhere
to the flanks of oxen that have rested in a moist meadow
till the earth has dried on them. Before fences were made,
the migrating horses, oxen, and bisons, in this way must
have carried such seeds for long distances, and any adhering
to the head of an animal would be carried across an arm
of the sea uninjured. It must be remembered, however,
that the autumn migration of mammals, which is the
migration when nearly all the seeds are ripe, would have
been southward in Britain, and consequently could only carry
plants in that direction. The northward migration taking
place in spring, few seeds would be carried, except such as
had become entangled in the fur and were shed with it
next summer. Wading and swimming birds, on the other
hand, commonly corne to Britain from the north and east
in autumn, leaving the colder districts at a time when the
seeds are ripe, thus bringing the smaller ones to this
country. This is probably the reason why so large a pro-
portion of the minute-seeded Arctic plants are found in
Britain, though many of the species only occur in small
numbers and at various scattered localities.

The next group, that containing the plants with large
edible unprotected seeds, is a small one in this country ;
but it is of especial importance on account of the difficulty
the species present when we try to account for their pre-
sence in these Islands, except on the hypothesis of a former

28 Origin of the British Flora.

greater continuity of the land. The difficulty is so real
that I have devoted particular attention to the attempt to
discover in what manner large soft seeds, which cannot be
carried in fur or feathers, and are killed by digestion, can
be transported across deserts. It will be shown in Chapter
IV. that since suitable climatic conditions came into
existence there has been no sufficient change of land or
sea to give a continuous land passage from the Continent
for these plants — yet, here they are and their presence
must be explained.

The British plants to which these remarks particularly
apply are the following : — the Oak, Beech, Ash, Maple,
Privet, Spindle Tree, Ivy, Flags, Convolvulus, various
Mallows, White and Yellow Waterlilies, and Apple. In
each of these, except sometimes in the Waterlilies and
Apple, the fruit is eaten for the nutriment contained in the
seed itself, which is therefore generally destroyed. No
doubt in many of these plants the seeds are occasionally
dispersed by rivers ; but this will only scatter them along
the lower part of the same river-basin or at most some
distance along shore ; it will not carry Waterlilies to
isolated lakes or to other river basins, nor can dry-soil
plants be carried thus to scattered islands.

The largest edible seed we have is the acorn ; if it can
be transported freely for considerable distances uninjured,
the difficulty in the other cases must be more apparent
than real. In peat-mosses, on open chalk downs, and in
ploughed fields, often a mile or more from the nearest
mature tree, one constantly finds seedling Oaks, which last
a few months or, perhaps, a couple of years, and then die,
the conditions being unfavourable. I have for several
years noted the position of these seedling oaks, finding
them in places where no mammal would take the acorns.
For instance, they are common in any of the New Forest

Means of Dispersal. 29

peat-bogs that are within a mile of an Oak-tree. They
are common also in some places on the top of the escarp-
ment of the South Downs, half a mile from Oaks, and 300
or 400 feet above them. They are always associated with
empty acorn-husks, stabbed and torn in a peculiar way.
In October and November rooks feed in the Oak-trees, and
I have long felt convinced that they were mainly responsible
for the dispersal of acorns. On October 29th of 1895, in
the middle of an extensive field, bordered by an oak-copse
and scattered trees, I saw a flock of rooks feeding and
passing singly backwards and forwards to the Oaks. On
driving the birds away, and walking to the middle of the
field, I found hundreds of empty acorn-husks, and a
number of half-eaten pecked acorns. It was noticeable
that many of them were not shed acorns, but were accom-
panied by acorn-cups, the stalks of which had been bitten
to tear them off the tree. The reason for the selection of
acorns in cups is probably that they are easier to carry —a
shed acorn must be an awkwardly large and slippery thing
for a rook's beak, one with a stalk will be more convenient.
Several uninjured acorns were found, one, almost uninjured,
had been driven by a single peck deep into the soft soil of
a mole-hill.

In this way oak-woods must spread rapidly ; but we
still want observations as to the extreme distance to which
acorns are thus carried. I have seen seedling Oaks at a
distance of a mile from the nearest tree (not necessarily the
tree from which the acorn came) and have found the
characteristically torn husks somewhat further away.*
Mr. J. J. Armistead, moreover, recordsf that he once
found a young Oak in a sheltered ravine among sea-cliffs
on the northern coast of Hoy, Orkney. The tree was

* Nature^ No. 1358, vol. liii., p. 6 (1895).
t Zoologist^ p. 19 ( 1 891).

30 Origin of the British Flora.

over six feet high. A few Rock Doves bred near the
place, and he concluded that an acorn had been brought by
one of these birds, but where from ? Unless it had been
picked up on the sea-shore, it must have been carried a
long way indeed. It could hardly have been brought by
man, as the place was very remote, as well as difficult of
access. Rooks occasionally cross the Pentland Firth.
The distance of the north of Hoy from the nearest point
where Oaks grow is fully as great as is the distance across
the Strait of Dover ; it is probably more than twice as
great as was the gap between England and France at the
period when the Oak was re-introduced after the Glacial
Epoch. Not only have the cliffs of Dover and of Calais
steadily receded through the inroads of the sea, but when
the ' submerged forests ' flourished both the English and
the French Coasts seem to have been bordered by a wide
belt of flat land covered with Oaks, the stumps of which are
now found rooted in the ancient soil as much as forty feet
below the present sea-level.

The transportation of large edible seeds for such long
distances uninjured is probably of exceptional occurrence,
and is more probably due to rare accidents than to special
adaptation. Some years ago I found, for instance, in an
old chalk-pit the remains of a wood-pigeon which had met
with some accident. Its crop was full of broad-beans, all
of which were growing well, though under ordinary circum-
stances they would have been digested and destroyed.
As fully half at least of the birds that are hatched must
die by various accidents before the following season, it is
evident that this dispersal of the contents of their crops
must be of daily occurrence. A pigeon would easily cross
the Strait of Dover in half-an-hour, and in the days when
raptorial birds and wild cats were plentiful, many must
have been struck down with their last meal undigested.

Means of Dispersal. 31

Accidents of this sort, however, are not absolutely-
necessary for the dispersal of the seeds ; for a considerable
number, even of such soft seeds as that of the Ivy, are
passed with their vitality unimpaired. This is often the
case when the bird or other animal has been feeding
greedily ; and at such times the bird may throw up great
part of its food undigested, especially if it is startled.'

Birds, especially young birds, as Professor Lloyd
Morgan has shown, learn by experience, and try various
unsuitable foods. This must often lead to their eating
indigestible, poisonous, or aperient fruits, which are not
commonly taken. So many fruits have medicinal qualities
that these in many cases may be special adaptations to aid
the dispersal of the seeds. The migrating bird in its first
year is constantly coming across plants new to it, and this
at times when it is too tired and hungry to discriminate.

Mammals also must have greatly aided the dispersal of
seeds in former times, for an ox, a deer, or a horse falling
over the cliffs of France would tend to drift with the
prevalent south-west wind till it was thrown upon the
English Coast, where wolves and foxes would pull it to
pieces, dragging the remains beyond the reach of the sea,
and perhaps burying parts, with the undigested vegetable
food still contained in the stomach.

It is needless to multiply instances, enough has been
said to show that the special modes of transportation
studied by Lyell and Darwin, added to the accumulated
accidents of some thousands of years, are sufficient to
account for the introduction of the whole of our native
plants, without the necessity for any continuous land con-
nexion between the different islands, or with the Continent.
Indeed the constant rain of seeds over our Islands
is probably on such a scale that were it not for the
* E. M. Langley, iV<3:/«r^, December 15th, 1898.

32 Origin of the British Flora.

circumstance that most of them must fall on ground that is
already occupied, we should continually have to record the
introduction of new species. New plants are rarely intro-
duced at the present day, merely because all the species
occurring within a reasonable distance have already had
their chance, and those that were suited to our climate
established themselves long ago. The modern introduc-
tions are mainly weeds of cultivation that cannot compete
with the native plants on uncultivated ground, or are
species from distant lands.

As instances of how readily our native plants will
occupy any tract newly made fit for them, I will mention
two or three cases that have particularly struck my
attention. When the new railway to Cromer was made,
the turf and top soil were pared off for a long distance, but
nothing more was done for several months. Next summer
the route of the new line was marked by a scarlet ribbon,
which could be seen stretching across the country, the
newly bared sub-soil having been taken possession of by a
profusion of poppies. A new embankment on the Bourne-
mouth line near Brockenhurst, again, for several years was
gay with corn-marigolds, which have since died down and
mostly disappeared. A still more remarkable case is seen
in the rapidity with which aquatic plants and animals
spread to a newly dug pond. In fact, so continuous is
this migration that we can get a fair idea how long a pond
has been made, and has contained water, by the number of
species of aquatic plants and mollusca that it yields. A
mediaeval fish-pond or moat contains a much more varied
fauna and flora than is found in a newly dug dew-pond
on the Chalk Downs, though it is surprising how many
species find their way to these ponds.*

* See Reid, ' The Natural History of Isolated Ponds,' Trans.
Norfolk Nat. Soc, Vol. V., pp. 272-286 (1892).

33

CHAPTER IV.

Changes in Geography and Climate.

When we discuss the origin of the British flora or
fauna it is impossible to assume, as we can in the case
of certain oceanic islands, that the process has been no
more than the gradual introduction of the plants, under
unchanging climatic conditions, into an area of limited
and almost unvarying extent, holding unchanging relations
with the nearest land, and till that time unoccupied by
any other flora. Both geographical and climatic changes
have played an essential part in shaping our flora as we
now see it. Moreover, except in part of our country
immediately after the retreat of the ice, each plant intro-
duced seems to have been brought into an area already
clothed with vegetation, though, under a changing climate,
the native plants may have become less adapted for the
station than were the intruders. It will be necessary,
therefore, to trace out the changes of land and sea which
have affected our islands since the existing plants and
animals first made their appearance here; though, as was
suggested in the last chapter, I greatly doubt whether in
islands so near a continent the actual junction or isolation
is of such great importance as has been imagined. Plants
can certainly overleap barriers more easily than is usually
thought. In various indirect ways, however, former geo-
graphical changes must greatly have facilitated the dis-

D

34 Origin of the B7dtish Flora.

persal of the species, and a short discussion of the principal
changes that can be shown to have taken place may assist
in explaining some of the anomalies in geographical
distribution.

It is useless for our present purpose to go back to any
distant geological period, for in Britain there exists so vast
a break in the series of Tertiary strata that we are unable
to bridge it. Our Middle Tertiary flora, which can be
studied in the Oligocene strata of Hampshire, is a sub-
tropical one, not allied to that now occupying the country.
The history of the succeeding Miocene Period in these
islands is a complete blank, for we have no fossiliferous
deposits of that age, and all we can say is, that the
Miocene appears to have been a period of great earth-
movement and folding, under which the surface con-
figuration of Britain was completely changed. Whether
Britain was then under water or was mainly dry land we
do not know. Certain of the Miocene plants found on
the Continent are living European species — probably none
of them now British — and the flora as a whole begins to
show a distinct affinity with that now occupying the
southern parts of the Continent.

Throughout the Pliocene Period there is evidence of
the slow refrigeration which culminated in the Glacial
Epoch ; but unfortunately, as far as the botany is con-
cerned, this climatic change cannot be followed, for plants
only occur in the newest stage of the period. The whole
of the strata of Older Pliocene age yet discovered in Britain
are of marine origin, and were laid down at some distance
from land in a warm sea. The Coralline Crag of Suffolk
yields, however, a few drifted land-shells, and at its base
contains bones of land animals, washed out of some older
deposit ; but there are in it no determinable plant-remains.
A few pieces of much decayed worm-eaten drift-wood are

Changes in Geography and Climate. 35

all that I have seen, and, as these might well have drifted
across the Atlantic with the Gulf Stream, they are of no
value for our present inquiry; the rolled fragments of
phosphatised or silicified palm-wood in museums do not
really belong to the period of the Crag, they are washed
out of the underlying London Clay.

During the earlier stages of the Newer Pliocene Period
the climate was still somewhat warmer than at the present
day, as is indicated by both the marine and the land
mollusca. Britain then seems to have taken somewhat its
present shape, for we find in our eastern counties traces
of a shore-line, parallel to the existing one, and of an
adjoining area of dry land, on which flourished various
mammals and mollusca. Of the associated plants we as
yet know nothing, mainly, I believe, because collectors
who examine the Red Crag desire to obtain mollusca or
mammals, and do not look for the fruits and seeds, which
moreover in a marine deposit, even of littoral origin, are
usually rare and badly preserved. The land and fresh-
water mollusca of the lower part of the Red Crag are
mainly south- European ; those of the Upper Red Crag
and of later Crag Deposits are more northern — there is
still a slight admixture of extinct forms, even in the newest.

Only in the latest deposits belonging to the Pliocene
Period can we find a copious land fauna and flora, and, as
far as the plants now inhabiting Britain are concerned,
history begins with the Cromer F'orest-bed; all before is
prehistoric and speculative. The so-called Forest-bed
consists of a series of estuarine and lacustrine strata, laid
down apparently by the ancient Rhine, which at that
period seems to have crossed a low area now occupied by
the shallow southern half of the North Sea.*

* 'Geology of the Country around Cromer' (1882); 'Pliocene
Deposits of Britain' (1890), Memoirs Geological Survey.

;^6 Origin of the British Flora.

We cannot speak confidently on the point, but the
evidence suggests that the general outline of the British
Isles did not greatly differ from that which now holds,
the principal difference probably being, that the Strait of
Dover had not then been cut, and that England was
connected with Belgium and Holland by a wide alluvial
plain. The legible records of the period here referred to
are confined to the eastern part of the counties of Norfolk
and Suffolk, though deposits probably of the same age,
but containing no fossils, occur in several other of the
eastern and southern counties. At one spot only, outside
East Anglia, are fossils apparently of this age to be found.
Dewlish, in Dorset, has yielded a few bones of the
characteristic elephant, Elephas meridionalis ; but no other
fossils could be discovered. If the deposit is of the same
age of the Forest-bed, it certainly suggests that the main
contours of the land were already shaped ; though most of
the valleys, in that region at any rate, are of later date.
The climate indicated by the plants and animals of the
Cromer Forest-bed is very like that which we now enjoy;
the warmth of the Miocene and early Pliocene Periods had
passed away, but the cold of the Glacial Epoch had not
yet swept off the numerous large mammals, nor trans-
formed the character of the vegetation.

The Pliocene Period, with its temperate and gradually
cooling climate, was separated from the present era by a
period of which the exact history is still obscure. We
know that this Pleistocene Period was characterised by
more than one wave of intense cold, which, for a time
must profoundly have modified the fauna and flora of
Britain. It was also marked by milder intervals, suffi-
ciently long for the temperate plants to re-appear ; and
also by a period of drought, which brought the fauna of
Central Asia into continental Europe, and in a minor

Changes in Geography and Climate. 37

degree affected the climate of Britain. Man first appeared
in Britain during the latter half of the Pleistocene Period ;
or rather we should perhaps say, that we have as yet no
satisfactory evidence of his earlier incoming.

The physical geography of Britain during the period
which saw the formation of the Glacial and Palaeolithic
deposits is still uncertain in many points. I think, how-
ever, that the evidence warrants us in saying that no very
great changes affected the boundaries of land and sea.
Submergence of part of the land took place; but in the
South and East of England at any rate, only to a limited
extent, perhaps 1 50 feet. There was also a time when the
land stood at a greater elevation; though in England this
elevation above the present level does not appear to have
exceeded 70 feet. Coast-lines have also been cut back in
the course of time by the incessant action of the waves,
and in other places shingle-beaches or sand-dunes have
slightly encroached on the sea. But all these changes can
scarcely have been sufficient greatly to modify the outline
of Britain ; though in indirect ways their influence on the
flora must have been considerable. The changes which
modified the Pleistocene fauna and flora were of an
exceptional character; for, besides the enormous fall and
the great oscillations in the temperature, the accumulation
of vast uninhabitable deserts of ice and snow must have
blotted out all plant life over great part of Britain. These
deserts must also have affected the migration of the
Arctic plants in ways that even yet have been scarcely
recognised.

An attempt will be made to give an outline of the
succession of events as far as the history can be traced ;
but it may be necessary to warn readers that I have been
led to interpret the records somewhat differently from other
geologists. Approaching the subject from the point of

38 Origin of the British Flora.

view of a naturalist, the comparative importance of the
different stages and of the different agencies, and even the
reading of the physical geography, will assume an aspect
very unlike that ordinarily laid before the student. To
the extreme glacialist the ' Pleistocene * is equivalent to the

* Glacial ' Period, and the scattered relics of Interglacial
mild epochs are judged to be of small importance. It may
be thought that the following notes go to the opposite
extreme. I believe, however, that the accumulation of ice
and snow merely marked two or more culminating epochs
in a period when the climate was at least as commonly
temperate as Arctic. The geological evidence for this I
have already published (see also below ' Hoxne,' p. JJ ;

* Selsey,' p. 88 ; ' West Wittering,' p. 94.)

The appearance of man in this country is sometimes
thought to mark a new era ; but, as far as our present
information goes, it was long before he had much influence
on the character of the fauna and flora. Palaeolithic man
was only one more carnivorous animal added to a fauna
which already possessed several quite as dangerous, and
apparently occurring in greater numbers. He did not
cultivate the ground, and therefore would not introduce
weeds of cultivation. We do not know whether he often
crossed the narrow seas ; though it is doubtful whether an
occasional canoe, not freighted with vegetable produce,
would greatly aid in the dispersal of plants which could be
carried by so many other messengers. It was not till
Neolithic man appeared, with domesticated animals, culti-
vated plants, and probably with more seaworthy canoes,
that the human race took a leading part in the dispersal
of seeds. It still remains to be seen how large a proportion
of our plants were unrepresented in Britain before his
days.

We have now to trace in a few words the succession of

Changes in Geography and Climate, 39

events during this somewhat obscure period. The unmis-
takably Preglacial records cease, as already observed, with
the temperate Cromer Forest-bed. Then succeeds a marine
stratum showing a submergence of perhaps fifty feet,
which cannot greatly have altered the outline of the
country, though at present little is known about this epoch.
Next follows a colder period, with Arctic plants ; and as
these occur just above the present sea-level, and lie evenly
on the strata below without deeply channelling them, the
height of the land at the commencement of the Glacial
Epoch, in Norfolk at any rate must have been almost the
same as it is now.

The freezing of the shallow land-locked North Sea, and
the steady accumulation of snow, which could neither
escape nor melt sufficiently fast, seems next to have
resulted in the formation of an ice-sheet continuous with
that pouring down from Norway and the Baltic, and this
ice-sheet overspread the east of Britain as far south as the
Thames. Whether the Arctic flora had sufficient time
thoroughly to occupy Britain before this mantle over-
whelmed the lowlands seems somewhat doubtful, for the
only routes the plants could follow were across the North
Sea, or the more southerly land-passage by the isthmus
through which the Strait of Dover has now cut. The
absence of any comparatively large-seeded northern plants,
such as the Larch, Scandinavian Alder, or Arctic Poppy,
either in a recent or in a fossil state, suggests that the
small-seeded species that we do find were brought by
birds, either across the sea or across the desert of ice, and
did not come by land. To this epoch, when the drainage
of a large part of Europe was poured into the North Sea,
but could not escape northward on account of the ice,
belongs probably the severance of England from the
Continent, for the water was forced to cut itself a new

40 Origin of the British Flora.

channel across the low neck of land just beyond the
southern limit of the ice-sheet. Other parts of Britain
were hidden under ice-sheets whose gathering grounds had
other centres, and the result seems to have been the total
blotting out of the flora over the area north of the Thames
and Severn, with the possible exception of certain high
hills which rose above the ice. Even these were probably
so smothered with snow that only the steeper crags were
bare in summer.

The condition of the greater part of Britain during the
climax of the Glacial Epoch will not, therefore, greatly
interest the botanist. The flora was so nearly extermin-
ated that the interest is transferred to the non-glaciated
strip between the Thames and Severn and the English
Channel, and to a very small non-glaciated area in South
Wales. In these parts only could the Arctic plants and
mammals live, and the whole of Britain was so cold that
the temperate species must have entirely disappeared.

Many naturalists will disagree with the statement that
has just been made ; for it has become almost an article of
faith that there were certain warm corners in these Islands
where the Temperate animals and plants could survive, and
where the peculiar Lusitanian flora of Cornwall and of the
West of Ireland lingered on till the renewed warmth
enabled the plants again to spread. It will be necessary
therefore briefly to summarise the evidence on which the
opinion above expressed has been founded.*

The temperature of the sea and of the air do not neces-
sarily correspond in the same regions ; we will, therefore,
first discuss the evidence as to the lowest temperature of
the seas round Britain. For this purpose the former
southern limit of the formation of shore ice, or ' ice-foot,'

* See ' The Climate of Europe during the Glacial Epoch,' Natural
Science^ Vol. I., No. 6, pp. 427-433 (1892).

Changes in Geography and Climate. 41

ought to give a fairly accurate idea as to the temperature
of the water. No doubt a large iceberg may travel a
long distance through comparatively warm water before it
entirely melts away ; but shore-ice, such as forms every
winter in the Arctic Regions, once fringed our south coast,
and beset the shores of Brittany and of the Channel
Islands. When, in the spring, the ice became detached,
it transported its burden of included rocks hither and
thither, even across the Channel. We thus find on Selsea
Bill erratics weighing several tons, but undoubtedly derived
from Bognor or from the Isle of Wight, Others, equally
large, have come from the Channel Islands and the coast of
Brittany ; one block of granite is like that of Cornwall.
The transportation of large erratic blocks for distances of
at least a hundred miles, shows that the temperature of the
water in the spring, though sufficiently high to dislodge
the ice, was yet too low to melt it rapidly. Even with a
strong wind a flat mass of shore-ice would take several
days to cross the Channel. In order to compare this ice-
laden English Channel with existing seas, it is necessary
to travel northward, till we cross the isotherm of 32° P.,
and are near the Arctic Circle.

Thus far we ha^'e dealt solely with the temperature of
the sea. We will now turn to the evidence as to the
temperature of the air during the same period in the South
of England ; and for this we can employ both physical and
biological data. The country north of the Thames and
Severn, buried under ice, must have been bordered by a
wide strip of barren land, with dwarf birch and willow, but
without trees. In this belt flourished also a mammalian
fauna like that now inhabiting similar belts in the Arctic
Regions, for in the area lying between the ice-sheet and
the ice-cold English Channel it would be impossible to
have a mean temperature much above the freezing point.

42 Origin of the British Flora,

Remains of this boreal fauna and flora have now been
found at several places in the south of England. A large
assemblage of Arctic mammals has been discovered near
Salisbury, and it includes such thoroughly boreal forms as
the Musk Ox, Arctic Fox, and Lemmings. Even in what
is now one of the warmest parts of our Islands, Arctic
plants occur in the fossil state ; for Bovey Tracey, in
Devon, yields the Dwarf Birch, and the Bearberry. This
leaves no place of retreat within these islands for the
Temperate animals and plants. All Ireland was glaciated,
so nothing could live there, except perhaps a few Arctic
plants on the mountain-tops. All England was under ice,
except the extreme south ; and there the climate was too
cold for temperate plants to live. It may be suggested
that the Scilly Islands were warmer, and perhaps they
were somewhat better than Devon and Cornwall. But this
will not account for the preservation of the Lusitanian
Species, for most of them are not found on the Scilly
Islands, and plants like the Arbutus would be killed by a
climate only slightly more severe than that now found in
Ireland.

After the passing away of the ice there was a return to
genial conditions, which lasted so long that during this
' Inter-glacial ' period a series of physical changes took
place, and there was time for the Arctic species to die out
and for a large Temperate fauna and flora to occupy the
country. We do not yet know the history of some of the
stages, as there are several gaps in the record ; the changes,
however, were slow and gradual, allowing time for valleys
to be deepened and again silted up, for sea-cliffs to be cut
back, and for plants to spread far and wide over new
districts. During the greatest intensity of the cold, as we
have shown, there seems to have been a submergence of a
few feet. Then comes a break for which the records have

Changes in Geography and Climate. 43

not yet been discovered. The next stage known shows a
submergence of about 140 feet, with a sea slightly warmer
than that now washing the coast of Sussex. The marine
mollusca are species living in the English Channel at the
present day, mixed with a few that do not now range north
of the Bay of Biscay. We know nothing of the plants of
this stage, and it is probable that the warmth of the sea
was mainly the result of its greater depth, which allowed
ocean currents more freely to enter.

After this submergence the land rose gradually, the
climate apparently remaining unaltered, till we again find
freshwater and estuarine deposits, laid down when sea-level
must have been the same as at the present day, or slightly
lower. These deposits contain a prolific fauna and flora,
which includes several southern animals and plants, but no
northern ones. Then succeeds another transition stage,
about which we at present know very little, followed by a
second glaciation, less severe than the former one, or
perhaps characterised rather by a dry cold, which did not
permit of so great an accumulation of snow and ice, though
the northern parts of Britain were again glaciated.

I may be permitted at this point to say a few words on
the subject of the recurrence of Glacial Epochs, for it is a
matter that closely concerns the student of the geographical
distribution of animals and of plants. It will be unneces-
sary to enter into theoretical questions as to the cause of
these climatic oscillations, for they are evidently due to
something entirely unconnected with changes in the
physical geography of Britain or of Western Europe.
These notes are merely a chronicle of the climatic and
geographical changes for which we have direct evidence; a
true connected history of Britain since it became a recog-
nisable unit cannot yet be written. It will be observed
that neither of the doctrines commonly taught seems to be

44 Origin of the British Flora.

borne out by the evidence above mentioned. We have no
indication in our Tertiary or later deposits of a number of
alternating Glacial and Interglacial Epochs, such as are
required on the theory of Croll * adopted by Professor
James Geikie.-|- On the other hand, the evidence is
perfectly clear that this country saw two cold Epochs, and
certain indications make one suspect that there may have
been a third, less rigorous. The exact succession of events
is at present very difficult to follow ; for it is unsafe to
compare isolated records, which belong to different regions,
and may not belong to the same period. We need more
excavation and close examination of localities such as
Hoxne and the Selsey Peninsula, where several stages can
be studied in chronological order, with no possibility of
mistakes in the succession.

The wind-borne 'loess' of Central Europe, with its
desert or sand-dune mollusca and mammals, belongs
apparently to the second cold period just alluded to.
Only slight indications of this dry climate have been
discovered in Britain, and, though it may have marked an
important stage in the building up of our flora, we know
little about its plants in the south, while nearer the glaciated
area those found are common Arctic forms. It is always
difficult to obtain botanical evidence of a bygone period
of drought, for desert-plants seldom find their way into
lacustrine deposits, and porous sub-aerial deposits like
drift-sand or loess are the worst possible for the preserva-
tion of plant-remains, though they may be full of calcareous
fossils.

The South of England during the second period of

glaciation seems to have suffered from dry cold winters,

which froze the ground unprotected by snow, and allowed

the summer rains to fall on soils rendered impervious by

* Climate and Time. t Great Ice Age.

Changes in Geography and Climate. 45

deep freezing. This led to enormous and rapid denudation,
over areas where the rain now sinks in and is slowly given
out as springs. Masses of loose flint and chalk debris were
swept off the South Downs and spread out in a wide sheet
extending several miles over the lowlands, and over the
Interglacial deposits already described. Even in Cornwall
the rubbly drift known as ' head ' seems to have marked a
similar stage. It is difficult to believe that anything but a
poor Arctic vegetation could have withstood these condi-
tions, and the Arctic plants of Devon may belong to this
cold epoch, rather than to the older one represented by the
erratics of Sussex and the Boulder Clay near London.
The Arctic mammals found near Salisbury may belong to
the same stage, they are migratory or else Steppe species.

The stage that follows — the transition from the
Palaeolithic to the Neolithic — is unfortunately one of the
most obscure, and I can only suggest that the break is
more apparent than real, and that one follows the other in
close succession. No doubt there is generally a marked
difference between deposits of Palaeolithic and those of
Neolithic age, the older series occupying terraces far above
the reach of any flood, while the more recent series lie in
the bottoms, or below the bottoms, of existing valleys. It
may prove, however, that the climatic change and the
difference in the position of the deposits are related as cause
and effect, little change having really occurred in the
contours of the country. As soon as the climate amelio-
rated, frozen soil would no longer cause erosion and
deposition to act in the peculiar way above described.
The older deposits would be left stranded at all elevations,
and denudation and deposition would at once change to
the ordinary types caused by river action in a Temperate
climate. With the climate, the fauna and flora would also
change; and at the same time the race of hunters would

46 Origin of the British Flora.

give place to a higher race that tilled the ground and had
domesticated animals. These, however, are merely sug-
gestions ; for a systematic study of the deposits of this
stage also, at some point where they give a continuous
record, will probably solve the riddle.

The Neolithic and later periods do not call for any
lengthy description. At first the land stood at an elevation
some 60 or 70 feet above its present level, so that many of
the river-valleys were cut to that depth below the sea, and
much of the English Coast was fringed with a broad strip
of alluvium, which probably almost connected our island
with Belgium and France. The climate during this epoch
was Temperate, for in the lowest ' submerged forests ' the
Oak is the most abundant tree. Then gradual and inter-
mittent submergence flooded the lower parts of the valleys,
and caused them to be silted up by the deposits of rivers
that no longer had sufficient fall to scour their beds. In
some of the peaty deposits or old vegetable soils that mark
stages of rest in their process of submergence, we find
polished stone weapons, and relics of cultivated plants and
of domesticated animals. The flora of these deposits,
however, is still very imperfectly known ; but all the plants
are species still found in Britain, though the occurrence in
South Wales in a * submerged forest ' of Najas marina, a
plant now confined to Norfolk, shows that the local
distribution may have been slightly different.

Since the close of the Neolithic Period, changes in
physical geography have been slight, and have consisted
mainly in the continuous silting up of the flooded valleys,
and in the cutting back of the coast-line by the waves.
This latter process, it should be remembered, has been
sufficiently marked to increase the width of the Strait of
Dover, which in places is also being deepened by the
scour of the tides. When our present flora entered the

Changes in Geography and Climate. 47

»

country, at the close of the Glacial Epoch, it was far
easier for animals and plants to cross from the Continent
than it is now.

The reader will probably rise from the perusal of this
chapter with a confused idea of many small changes in
the limits of sea and land; which, however, were of no
very great importance as bearing on the past history of
our flora. This impression is, I believe, the correct one;
for, after twenty years* work at deposits belonging to the
periods here dealt with, I am greatly impressed with the
smallness and multitude of the changes, and with the
gradual way in which they occurred, as is demonstrated
wherever we can discover continuous records. The
climatic changes, on the other hand, though perhaps
equally gradual, were most thorough and sweeping ;
inevitably they must have been accompanied by corre-
sponding changes in the flora.

48

CHAPTER V.

Deposits containing Fossil Plants.

If we desire to trace out the past history of our native
plants, we must study such relics as are preserved in
various stratified deposits, especially those of which the
geological position can be proved by other evidence.
The questions are often asked : — Where are these relics
to be found, and what is the method of research adopted ?
These are questions the answers to which will not be
found in any text-book, nor, apparently, are they known
to most geologists. It will be useful, therefore, to give
a short description of the sort of deposits which have
proved most prolific, and of the methods that have usually
been adopted to obtain the plant-remains. This will be
followed by an account of the fossiliferous strata already
examined, with the leading characteristics of each, such
as date, nature, and origin of the deposits, general
character of the included fauna and flora, notes of any
local circumstances which must have affected the plants,
and finally, a list of the plants. This will occupy a good
deal of space ; but it is all information needed by the
local geologist or botanist, and will, I hope, aid in the
study of past history of the floras of our different counties
and districts. References have been added to published
authorities, from which a fuller account of the geology and
zoology can be obtained; but in every case, unless other-

Deposits containing Fossil Plants. 49

wise stated, I have myself determined the plants included
in the lists.

Various points have to be taken into consideration if
we desire to avoid failure or useless labour in our search
for seeds or leaves in a determinable state. The deposits
most likely to yield satisfactory results are not such as
one would at first sight select as best for the purpose.
On consideration, it will readily be understood that a
wide-spread peat-moss will yield little but remains of bog-
plants; an extensive lacustrine deposit will contain few
but aquatic species; a broad alluvial flat may only pre-
serve plants of the marsh and wet meadow. The work of
collecting at best is very laborious, and, in order to obtain
with the least amount of trouble an insight into the fossil
botany of any particular period or district, it is best, where
practicable, to select for examination the deposits of a
small stream which flowed through a varied country.
These will yield not only seeds of .the aquatic and marsh
plants that lived on the spot, but also of a variety of dry?
soil plants and trees which grew on sandy or rocky banks
overhanging the channel. They will also yield seeds of
numerous species which grew somewhat further away, and
were brought by birds and dropped from the overhanging
boughs ; and will contain winged seeds transported by the
wind.

It may be thought that plants of all these descrip-
tions will be found in a lake or peat-bog, and no doubt it
is so ; but they will be so rare, and mixed with so large a
proportion of seeds belonging to some few aquatic plants,
that the time spent in searching for them will be largely
increased. I speak of this from, personal experience ; for,
through an imperfect appreciation of this difficulty, much
time was lost in my earlier work, and samples of clay,
collected and washed with great labour, often yielded

B

50 Origin of the British Flora.

nothing but thousands of seeds belonging to half a dozen
aquatic plants, which were already quite well known. As
an example, we may take the flora of the Cromer Forest-
bed, which is still a small one, for the deposits belonging
to it are parts of a wide>spread alluvial plain, with shallow
pools and broads. Yet the* collection of the plants has
given me ten times the trouble that was needed to obtain
a much larger number of species at West Wittering. This
latter deposit is of estuarine origin — it therefore contains
mingled fresh-water, estuarine, and sea-coast plants ; it is
the deposit of a very small stream — the proportion of dry-
soil species is therefore exceptionally large, and their seeds
are unusually well preserved ; moreover, the stream within
a mile crossed the edges of a most varied series of strata,
including chalk, stiff clay, loose sand, marl, loam, and
gravel — the flora is therefore as good an epitome of that
of the surrounding district as could be obtained by the
examination of several- deposits, each of which only fulfils
some of these conditions. It may be added that, while the
best fossiliferous deposits in the Forest-bed are commonly
stiff clays or peaty-beds, difficult to take to pieces without
injury to the fossils, the strata at West Wittering are
sandy loams, which, when dried and placed in a sieve in
water, quickly fall to pieces and leave the seeds un-
injured.

Deposits like that just mentioned, though giving the
best general view of the flora of a district, are unsatisfac-
tory in certain respects; for they seldom yield well-
preserved leaves, and many species having soft seeds can
only be recognised by the leaves. In order to discover
leaves of any plants, other than the small tough-leaved
Arctic ones, it is commonly necessary to split up laminated
lacustrine clays, or masses of bog-iron ore ; but, unfortu-
nately, Pleistocene clays are seldom sufficiently firm to

Deposits containing Fossil Plants, 51

allow of handling in this way. Or else we must search
the masses of tufa deposited by calcareous springs; for
these yield beautifully preserved casts of deciduous leaves,
and may also contain impressions of the succulent leaves
of non-deciduous plants ; they are almost useless, however,
for the study of seeds, which are generally too small to be
recognisable in hollow impressions in a somewhat coarse-
grained matrix.

Want of time has prevented me from undertaking so
thorough an examination of the plants of the newest
deposits as they deserve, or as has been made in Sweden
by Professor A. G. Nathorst and Dr. Gunnar Andersson.
We happen, however, to possess a large series of deposits
of somewhat earlier date than any of those found in
Scandinavia ; it seems best, therefore, to devote attention
more particularly to the plants contained in them. Pre-
glacial plants are extremely rare in Europe, and the Inter-
glacial flora has only been studied at a few localities in
North Germany, principally by Dr. Carl Weber and
Professor A. Nehring.

It only remains to add a few words as to the position
in time of the various deposits to be described. They are
here divided into Preglacial, Early Glacial, InterglaciaU
Late Glacial, and Neolithic. The whole of the historic
period, from the invasion of the Romans downwards, has
purposely been omitted, not because it is of little import-
ance in the history of the flora as we now see it, but
because collecting has not yet been done with sufficient
accuracy to fix the century to which the deposits belong.
Without this, the identification of the included plants
would be of little value. One exception only has been
made. A certain number of plants from Silchester are
mentioned, as these were found in carefully selected
material obtained by Mr. A. H. Lyell during excavations

52 Origin of the British Flora.

at the Roman town, and were certainly contemporaneous
with the Roman occupation.

It is quite possible that more than one Interglacial
Period is represented in the deposits and plants that I
have examined; but the classification has been kept pur-
posely as simple as possible. It so happens, also, that the
most prolific of the Interglacial deposits in the South and
East of England all seem to fall into a single period —
that immediately succeeding the greatest intensity of the
cold. Those that remain have at present yielded so poor
a flora, which consists so exclusively of species of wide
range, that from a botanical point of view they are of little
importance. The botanical characteristics of the different
periods may be summarised thus : —

Preglacial (latest Pliocene).

Found on the coasts of Norfolk and Suffolk.

Plants are aquatic and wet-meadow species and forest
trees.

All yet known are now natives of Britain except Trapa
natanSf Najas minor, and Picea excelsa.

Associated with many large mammals, the majority of
which are now extinct.

Early Glacial.
Found at a few localities on the Norfolk coast.
Northern plants, including Salix polaris and Betula
nana; no forest trees except Birch and Alder.

Interglacial.

Southern and Eastern Counties (Hoxne beds D and E,
Hitchin, Grays, Selsey, Stone, West Wittering ; also
Deuben, Griinenthal, Klinge, Fahrenkrug, Lauenburg,
Honerdingen).

Deposits containing Fossil Plants, 53

Plants still living in the same district, mixed with a
few southern forms, those already determined being Acer
monspessulanum, Najas graminea, and Najas minor, and in
Germany Brasenia purpurea.

No northern species.

The associated mammals and mollusca suggest a climate
somewhat drier and sunnier than that now possessed by
the South of England.

Late Glacial.

Throughout Scotland and England as far south as
London and Devonshire. (Crianlarich, Hailes, Corstorphine»
Bridlington, Hoxne bed C, London, Bovey Tracey, &c.)

Numerous Arctic plants, all of which, except Salix
polaris, are still to be found on the mountains of Scotland.

Neolithic.

Including * submerged forests' and early peat mosses.
(Hailes upper bed, Redhall, Woolwich, Blashenwell, Barry
Docks, &c.)

Flora Temperate. Cultivated plants and weeds of
cultivation occasionally appear. Extensive Oak forests.
Pine common in the South of England. This flora is
better known in Scandinavia than in Britain ; it has been
divided by Swedish and Danish botanists into several
stages characterised by different trees (see p. 92).

Space will not allow me to give in full the evidence on
which the deposits are referred to different periods. Where
possible, the stratigraphical position has been studied; but
in certain cases where direct evidence of superposition is
not available I have dated the deposits according to the
affinities of the included fauna and flora. The animals
are, for this purpose, of more value than the plants, for
they change more rapidly ; plants, however, yield the best

54 Origin of the British Flora.

evidence of former climate. It will be seen that the date
of certain of the deposits is unmistakable; and particular
attention having been devoted to these and to their in-
cluded plants, any doubt as to the age of the remainder
is of comparatively little consequence botanically — nearly
all their plants can be authenticated from specimens found
in deposits of known age. The localities are placed in
alphabetical order, partly as being most convenient, partly
to avoid any appearance of forcing the correlation, as
might be suggested if they were here grouped into periods.
The principal foreign localities are added for purposes of
comparison ; but I have seen few of the plants from these.
It will be noticed that in the German Interglacial deposits
Brasenia purpurea^ a water-lily not now living in Europe,
is a common fossil, though it has not yet been discovered
in Britain.

Admiralty Offices, London.

(Abbott, ' The Section exposed in the foundations of
the New Admiralty Offices,' Proc. Geol. Assoc, Vol. XII.,
pp. 346-356. 1892.)

Associated with or below remains of Mammoth, Hippo-
potamus, and Rhinoceros are found a few plants, the only
determinable species being: —

Betula nana. Ceratophyllum demersum.

AiRDRiE, Lanark.

(Dunlop, ' Note on a Section of Boulder-clay, containing
a Bed of Peat,' Trans. Geol. Soc. Glasgow, Vol. VI 1 1.,
pp. 312-314. 1888.)

The peat is classed as Interglacial on account of its
occurrence between two beds of Boulder-clay. It contains
beetle-remains and the following species of plants : —

Deposits containing Fossil Plants. 55

Ranunculus aquatilis.

Betula nana.

Prunus Padus.

Empetrum nigrum

Potentilla Comarum.

Potamogeton.

Hippuris vulgaris.

Carex dioica.

Apium nodiflorum.

panicea.

Carduus.

rostrata.

Menyanthes trifoliata.

Isoetes lacustris.

Albert Dock, North Woolwich, Essex.

(Spurrell, 'On the Estuary of the Thames and its
Alluvium,' Proc, Geol. Assoc, Vol. XL, pp. 210-230. 1889.)

A ' Submerged Forest ' and peat bed beneath the
Roman layer yields the following plants : —

Cornus sanguinea. Quercus Robur sessiliflora.

Betula alba. Taxus baccata.

Alnus glutinosa. Phragmites communis.
Corylus Avellana.

Allenton, near Derby.

(Arnold-Bemrose, ' Discovery of Mammalian Remains
in the Old River-gravels of the Derwent near Derby,
Quart. Journ. Geol. Soc, Vol. LI I., pp. 497-500.' 1896.)

The plants were found associated with Hippopotamus
(apparently a whole skeleton), Elephant and Rhinoceros.
This fauna is probably Interglacial.

Ranunculus aquatilis. Valeriana officinalis.^

sceleratus. Eupatorium cannabinum.

Flammula. Leontodon autumnalis.

repens. Taraxacum officinale.

bulbosus (?). Ajuga reptans.

Sardous. Atriplex.

Viola palustris. Eleocharis palustris.

Montia fontana. Scirpus pauciflorus.

Rubus Idaeus. Carex.

Potentilla. Isoetes lacustris.
Hydrocotyle vulgaris

56 Origin of the British Flora,

Bacton, Norfolk.

(Reid, ' Pliocene Deposits of Britain,' Memoirs Geol.
Survey, 1890; Reid, 'Geology of the Country around
Cromer,' Memoirs Geol. Survey, 1882.)

The Cromer Forest-bed at Bacton yields cones of
Pinus sylvestris and Picea excelsa, and rhizomes of
Osmunda; the principal fossiliferous localities are, how-
ever, close to Ostend Gap, a short distance to the south-
east, and are described under that heading.

Ballaugh, Isle oy Man.

('Report of Committee on Irish Elk Remains in the
Isle of Man,' Rep. Brit. Assoc, for 1897, p. 346. 1898.)

The deposits examined were as follows, Bed A being
the most recent : —

A. Peat, with caddis cases and eggs of insects.

Ranunculus Flammula. Hydrocotyle vulgaris.
Potentilla Tormentilla. Potamogeton.

B. Sand without fossils.

C. Sandy silt with Lepidurus {Apus) glacialis.

Ranunculus aquatilis. Carex.
Poterium officinale. Schoenus (?).

Salix herbacea.

D. Loamy Peat.

Ranunculus aquatilis. Littorella lacustris.

Flammula. Potamogeton crispus.

repens. Carex.

E. Gravel without fossils.

F. Chara Marl with Megaceros hibemicus.

Ranunculus aquatilis. Potamogeton natans.

Flammula. sp.

repens. Carex.

Littorella lacustris. Chara, 2 sp.
Empetrum nigrum.

Deposits containing Fossil Plants, 57

A is Recent or Neolithic. C, from the occurrence of
Salix herbacea and Apus glacialis, is classed as Late
Glacial. D and F are provisionally classed with C, but
may belong to a milder, Interglacial, period. These
deposits, and those found at Close y Garey, occupy
silted-up hollows in the glacial gravel. It is not yet clear
whether the poverty of the flora, and the entire absence
thus far of remains of dry-soil plants, is due to the
barren water-logged character of the gravel-flat, or is
characteristic of the flora of the Isle of Man at these
periods.

Barry Docks, Glamorgan.

(Strahan, ' On submerged Land-surfaces at Barry, Gla-
morganshire. With Notes on the Fauna and Flora by
Clement Reid,' Quart. Journ. Geol. Soc, Vol. LIT.,
pp. 474-489. 1896.)

The newest of these, about 4 feet below mean tide,
yielded a polished Neolithic implement and also, according
to Mr. Storrie, logs of Willow, Pine, and Oak. An
associated shell-marl was full of freshwater shells and
seeds of: —

Rumex crispus. Potamogeton.

Atriplex. Najas marina.

Salix (leaves). Chara, 2 species.

Najas marina is now confined to east Norfolk.

The second peat, or land-surface, is composed mainly
of sedges {Scirpus maritimus) and lies about 9 feet below
mean-tide level.

The third peat is composed of large timber and matted
Sallow and Reed, with seeds of Valeriana officinalis and
Carex. It lies 20 feet below Ordnance Datum, but shows
no sign of the influence of salt water.

The fourth peat is a true submerged land-surface, full

58 Origin of the British Flora.

of Oak-roots in place, indicating a soil above the reach of
the sea. It lies 35 feet below mean tide level, and points
to a subsidence of fully 55 feet. The plants are: —

Crataegus Oxyacantha. Quercus Robur.

Cornus sanguinea. Salix Caprea.

Corylus Avellana. Sparganium.

The whole of the deposits belong in all probability to
the Neolithic Period.

Beeston, Norfolk.

(Reid, ' Piocene Deposits of Britain,' Mem. GeoL Survey.
1890; Reid, * Geology of the Country around Cromer/
Mem. GeoL Survey. 1882.)

Two distinct plant-bearing deposits are represented at
Beeston. The lower is a peaty loam full of seeds of
Temperate plants, and belongs to the Preglacial Cromer
Forest-bed. The upper, and newer, is an Early Glacial
stratified loam with leaves of Arctic plants, at the base of
the Boulder Clays. At one spot an intermediate deposit
is perhaps represented ; this is here classed as the base
of the Arctic bed.

Plants from the Cromer Forest-bed : —

Thalictrum flavum. Alnus glutinosa.

Ranunculus aquatilis. Ceratophyllum demersum.

repens. Stratiotes aloides.

Nuphar luteum. Alisma Plantago.

Stellaria aquatica. Potamogeton pectinatus.

Poterium officinale. trichoides.

Hippuris vulgaris. heterophyllus.

CEnanthe Phellandrium. Najas marina.

Carduus. Scirpus pauciflorus (?).

Stachys palustris. caespitosus.

Atriplex patula. fluitans (?).

Rumex Acetosella. Carex (several sp.j.

maritimus. Isoetes lacustris.

Betula alba.

Deposits containing Fossil Plants, 59

Plants from the base of the Arctic bed : —

Thalictrum minus (?). Rumex maritimus.

Ranunculus aquatilis. Betula nana.

Rubus Idseus. Alnus glutinosa.

Poterium officinale. Ceratophyllum demersum.

Hippuris vulgaris. Zannichellia palustris.

Myriophyllum spicatum. Eleocharis palustris.

Galium boreale (.?). Scirpus lacustris.

Tanacetum vulgare. Isoetes lacustris.
Menyanthes trifoliata.

Plants of the Arctic bed : —

Salix polaris.

Black Burn, East Tarbet.

From the Clyde Beds at this place Mr. David Robertson
obtained seeds of Rubus Idceus. The sub-Arctic character
of the associated marine moUusca causes these deposits to
be here classed as Late Glacial.

Blashenwell, Dorset.

(Reid, ' An Early Neolithic Kitchen-midden and Tufa-
ceous Deposit at Blashenwell, near Corfe Castle,' Proc.
Dorset Field Club, Vol. XVII., pp. 67-75. 1897.)

The calcareous tufa contains only recent species of
mammals and mollusca, with charcoal and unpolished flint
implements of early Neolithic type. The plant-remains
are impressions of leaves and twigs, with decayed wood and
hazel-nuts. The only determinable plants found were : —

Ulmus montana (.?). Quercus Robur.

Corylus Avellana.

BovEY Tracey, Devon.

(Pengelly and Heer, ' On the Lignite Formation of
Bovey Tracey,' Phil Trans., Part II. 1862 ; Nathorst, ' On
the Distribution of Arctic Plants during the Post-glacial
Epoch,' /^^^^. Pot., n. s.. Vol. II., p. 225. 1873.)

6o Origin of the British Flora.

Some clays overlying the Eocene lignite deposits have
yielded a few Arctic plants. It is not clear whether these
should be classed as Early or Late Glacial. They yield : —

Arctostaphylos Uva-ursi. Salix cinerea.
Betula alba. Pinus.
nana.

Bridlington, Yorkshire.

(Nathorst, * Ueber neue Funde von fossilen Glacial-
pflanzen,' Engler's Bot. Jahrb., 1881, p. 431.)

A hollow in the boulder-clay, filled with peaty marl,
is here classed as Late Glacial from its stratigraphical
position and the occurrence in it of Betula nana.

Broughton, Edinburgh.

From a peaty deposit at this spot Mr. James Bennie
has recently obtained a few plants, probably of the same
date as those from the Neolithic deposits at Hailes and
Redhall. There is nothing characteristic in the list: —

Ranunculus aquatilis. Polygonum aviculare.

Flammula. — -. Persicaria.

Lingua. Rumex.

repens. Potamogeton.

Stellaria media. Scirpus setaceus.

Montia fontana. Eriophorum.

Carduus. Carex.
Atriplex (?).

Caerwys, Flintshire.

(Maw, ' On the occurrence of extensive Deposits of
Tufa in Flintshire,' Geol. Mag., Vol. III., p. 253. 1866;
Strahan, ' Geology of Flint, Mold, and Ruthin,' p. 1 50,
Mem. Geol. Survey. 1890.)

An extensive deposit of calcareous tufa at this place is
full of leaves ; but the date cannot be fixed, as the tufa

Deposits containing Fossil Plants, 6i

is still being formed. The part from which the leaves
were collected is probably of Neolithic age.

Pyrus Aucuparia. Salix cinera.

Hedera Helix. Caprea.

Betula alba. Populus tremula.

Casewick, Lincolnshire.

(Morris, ' On some Sections in the Oolitic District of
Lincolnshire,' Quart. Journ. Geol. Soc.^ Vol. IX., p. 3 1 7. 1853;
Reid, ' Pleistocene Plants from Casewick . . . .' ibid. Vol.
LI IL, pp. 463, 464. 1897.),

An ancient alluvial deposit fills a channel in Oolite.
The age is uncertain, as there is nothing peculiar among
the fossils. Though here classed as Neolithic it may be
of older date.

Nuphar luteum. Rumex crispus.

Galium Aparine. Ceratophyllum demersum.

Atriplex patula. Scirpus lacustris.

Close y Garey, Isle of Man.

(' Report of Committee on Irish Elk Remains in the
Isle of Man,' Rep. Brit. Assoc, for 1898, p. 550. 1899.)

The deposits occupy a silted-up hollow, like that at
Ballaugh, in glacial gravel. The plants are : —

B. Peat:—

Ranunculus Flammula. Carduus crispus.
Viola palustris. Menyanthes trifoliata.
Rubus fruticosus. Empetrum nigrum.
Potentilla Tormentilla. Potamogeton.
Comarum. Carex, 4 sp.

C. Megaceros-marl : —

Ranunculus repens. Empetrum nigrum.

Viola palustris. Potamogeton.

Potentilla Comarum. Carex, 4 sp.

Myriophyllum spicatum. Chara.
Rumex obtusifolius.

62 Origin of the British Flora.

E. Loam at the base of the marl : —

Betula alba. Carex.

Potamogeton.

B is Recent or Neolithic. C and E correspond with
the marl at Ballaugh (which see), and are classed pro-
visionally as Late Glacial.

CORSTORPHINE, NEAR EDINBURGH.

(Bennie, 'Arctic Plants in the old Lake Deposits of
Scotland,' Ann. Scot. Nat. Hist., 1894, pp. 46-52.)

In the lower part of the lacustrine deposits filling a
silted-up lake are numerous seeds and leaves of Arctic
plants. The deposit is probably Late Glacial, and con-
temporaneous with those of Hailes and Dronachy.

Ranunculus aquatilis. Menyanthes trifoliata.

repens. Oxyria digyna.

Viola palustris. Betula nana.

Stellaria media. Salix repens.

Rubus. herbacea.

Dryas octopetala. polaris.

Potentilla. reticulata.

Poterium officinale. Empetrum nigrum.

Hippuris vulgaris. Potamogeton.

Myriophyllum spicatum. Eleocharis palustris.

Taraxacum officinale. Scirpus pauciflorus.

Andromeda Polifolia. lacustris.

Loiseleuria procumbens. Carex, 2 sp.

Gorton, Suffolk.

(Raid, * Notes on the Sections at Gorton, seen during
the recent visit of the members of the Geological Con-
gress,' Trans. Norfolk Nat. Soc, Vol. IV., pp. 606-609.
1889.)

A bed of lignite and clay, belonging to the Preglacial
Cromer Forest-bed, here yields abundant seeds.

Deposits containing Fossil Plants

63

Thalictrum flavum.

Ranunculus.

Nuphar luteum.

Hydrocotyle vulgaris.

CEnanthe.

Solanum Dulcamara.

Atriplex patula.

Alnus glutinosa.

Ceratophyllum demersum.

Stratiotes aloides.
Sparganium ramosum.
Potamogeton lucens.

trichoides.

Zannichellia palustris.
Scirpus pauciflorus.

lacustris.

Eriophorum angustifolium.
Carex.

CowDEN Glen, Renfrewshire.

(Geikie, 'Great Ice Age,' 3rd edit., pp. 102-104. 1894;
Bennie, ' On Things New and Old from the Ancient Lake
of Cowdenglen, Renfrewshire,' Trans. Geol. Soc. Glasgow,
1891, pp. 213-225.)

This deposit has given rise to much discussion, Pro-
fessor James Geikie, Mr. Bennie, and others maintaining
that the peat is Interglacial and interbedded between two
masses of boulder-clay. Some writers, however, consider
the upper boulder-clay to be merely a landslip. The plants
have a recent appearance, and include the Opium Poppy,
a species cultivated in Neolithic times. I think it safer,
therefore, not to consider them older than the Neolithic
period.

Ranunculus aquatilis.

Flammula.

Papaver somniferum.
Viola.

Montia fontana.
Rubus Idaeus.
Poterium officinale.
Hippuris vulgaris.
Myriophyllum spicatum.
Galium palustre.
Menyanthes trifoliata.
Pedicularis palustris.
Galeopsis Tetrahit.
Ajuga reptans.
Polygonum lapthifolium.

Rumex crispus.
Betula alba.
Corylus Avellana.
Salix repens.
Pinus sylvestris.
Alisma Plantago.
Potamogeton perfoliatus.

pusillus.

pectinatus.

Scirpus lacustris.

pauciflorus.

Carex rostrata.
Isoetes lacustris.
Chara.

64 Origin of the British Flora.

Crianlarich, Perthshire.

Peaty loam with leaves of Arctic plants was found in a
railway cutting, and a sample given to me by Mr. J. R.
Dakyns yielded the subjoined species. The exact relation
of the deposit to the old moraines is not perfectly clear,
though the plant-bed would seem to be the newer of the
two, and therefore Late Glacial.

Dryas octopetala. Salix herbacea.

Betula alba. reticulata.

nana. Empetrum nigrum.

Salix repens.

Cromer, Norfolk.

(Reid, ' Pliocene Deposits of Britain,' Mem. Geol.
Survey. 1 890 ; Reid, ' Geology of the Country around
Cromer,' Mem, Geol. Survey. 1882.)

The Preglacial Cromer Forest-bed at Cromer itself is
mainly of estuarine origin, and yields therefore only drift-
wood and cones of Scotch and Spruce Firs. About
three-quarters of a mile north-west of Cromer black mud
belonging to the lower part of the Forest-bed is full of
aquatic plants. The species are : —

Ranunculus aquatilis. Potamogeton praelongus.

Myriophyllum spicatum. Eriophorum angustifolium.
Potamogeton lucens.

A full list of plants from the Cromer Forest-bed of all
localities will be found in the Table, p. 171.

Crossness, Essex.

(Spurrell, ' On the Estuary of the Thames and its
Alluvium,' Proc. Geol Assoc, Vol. XL, pp. 210-230. 1889.)

Two beds of peat or ' submerged forest ' are here met
with beneath the estuarine deposits of the Thames and
underlying the Roman layer. The deposits are synchronous
with those at Tilbury and at the Albert Dock.

Deposits containing Fossil Plants.

65

(l. — Lower Peat. u. — Upper Peat.)

Ranunculus sceleratus

repens .

Oxalis Acetosella
Ilex Aquifolium
Acer campestre
Prunus communis

domestica

Avium ,

Rubus fruticosus
Rosa

*Pyrus communis
Crataegus Oxyacantha
Hippuris Vulgaris
CEnanthe Phellandrium
Hedera Helix .
Cornus sanfjuinea

L. Sambucus nigra . L. U.

L. U. Viburnum Opulus . L. U.

U. Fraxinus excelsior . L. U.

L. U. Polygonum Hydropiper L.

L. U. Mercurialis perennis . L. U.

L. U. Betula alba . . L. U.

U. Alnus glutinosa . . L. U.

L. U. Corylus Avellana . L. U.

L. U. Quercus Robur. . L. U.

L. U. Salix . . . U.

L. U. Taxus baccata . . L. U.

L. u. Iris Pseudacorus . U.

TJ. Sparganium ramosum L. U.

U. Scirpus lacustris . L.

L. U. Phragmites communis L. U.
L. U.

Deuben, Saxony.

(Nathorst, * Die Entdeckung einer fossilen Glacialflora
in Sachsen, am aussersten Rande des nordischen Dilu-
viums/ Kongl. Vetenskaps-Akad. Forh., 1894, pp. 519-544.)

Arctic plants are here found nearly as far south as
Lat 50°. The species recorded are : —

Batrachium confervoides (?)
Stellaria graminea (?).
Saxifraga oppostifolia.

Hirculus.

aizoides.

Polygonum viviparum.
Salix herbacea.

retusa.

myrtilloides.

arbuscula (?).

Dronachy, Fife.

(Bennie, ' Arctic Plant-beds in Scotland,' Ann. Scottish
Nat. Hist., 1896, pp. 53-56.)

Lacustrine deposits, like those of Hailes and Corstor-
phine, were laid open during the construction of a new

* Determined by Prof Marshall Ward from wood.

F

66 Origin of the British Flora.

railway about half a mile from Auchtertool. The plants
sent to me by Mr. Bennie were : —

Thalictrum flavum (?J. Salix polaris.

Ranunculus aquatilis. reticulata.

Viola palustris (?). Empetrum nigrum.

Hippuris vulgaris. Potamogeton, 2 sp.

QEnanthe. Eleocharis palustris.

Menyanthes trifoliata. Scirpus pauciflorus.

Betula nana. fluitans (.^).

Salix herbacea. Carex, 2 sp.

Drope, Glamorgan.

My colleague Mr. Cantrill has obtained some seeds
and freshwater shells from beds of peat and marl in a
railway cutting near Cardiff. There is nothing character-
istic among the fossils, though the assemblage and the
relations of the deposits both suggest the Neolithic period.

Viola palustris (?).

Potamogeton hetero-

Hippuris vulgaris.

phyllus (.?).

Menyanthes trifoliata.

Potamogeton natans.

Betula alba.

Carex.

Juncus (i*)

Chara.

DuRSLEY, Gloucester.

The calcareous tufa used for building is full of leaves ;
but, as the tufa is still forming, it^ is difficult at present to
date the different parts of the sheet. Leaves of Hazel,
Elm, and Hartstongue were found by Miss M. A. Reid and
myself.

Elie, Fife.

(Bennie and Scott, * The Ancient Lake of Elie,' Proc,
R. Phys. Soc , Edinburgh, Vol. XH., pp. 148-170. 1893.)

The occurrence of bones of Sheep and Rabbit, with
capsules of Flax and seeds of Fool's Parsley, suggests that

Deposits containing Fossil Plants.

67

the lacustrine deposit of Elie is not older than the Neolithic
period. The flowering plants are : —

Ranunculus aquatilis.
Flammula.

repens.

Viola (?).
Lychnis alba.

diurna (?).

Stellaria media.
Li num.
Rubus Idaius.
Potentilla Tormentilla.
Hydrocotyle vulgaris.
iEthusa Cynapium.

Valeriana ofificinalis.
Cnicus lanceolatus.
Menyanthes trifoliata.
Ajuga reptans.
Atriplex patula.
Polygonum Persicaria.
Corylus Avellana.
Iris Pseudacorus.
Potamogeton.
Carex, several species,
Phragmites (?).
Chara.

Endsleigh Street, London.

(Hicks, ' On the Discovery of Mammoth and other
Remains in Endsleigh Street . . . .' Quart. Joum. Geol.
Soc, Vol. XLVIIL, pp. 453-468. 1892.)

A clayey loam containing bones of Mammoth and
numerous seeds, fills the lower part of a hollow eroded
in the London Clay. From its position at the base of
the Drift, and its resemblance to other deposits in the
neighbourhood of London, it is most probably Interglacial,
though newer than the boulder-clay of Middlesex.

Rumex obtusifolius.
Luzula (?) maxima (?).
Potamogeton obtusifolius.

Ranunculus aquatilis.
sceleratus.

repens.

Stellaria media.

Geranium.

Potentilla Tormentilla.

Hippuris vulgaris.

Myriophyllum spicatum.

Polygonum aviculare.

Persicaria.

crispus.

Zannichellia palustris.
Eleocharis palustris.
Carex dioica.

2 sp.

Chara.

Fahrenkrug in Holstein.

(Weber, ' Ueber die diluviale Flora von Fahrenkrug in
Holstein,' Engler's Bot. Jahrb., Beiblatt 43. 1893.)

68 Origin of the British Flora.

The deposits described are apparently of Interglacial
date. Like those of Lauenburg and KHnge, they yield
seeds of Brasenia purpurea^ a plant which has not yet been
found fossil in Britain. Its recent range is very wide,
though it does not include any part of Europe.

Nuphar luteum. Ceratophyllum submersum.

Nymphaea alba. demersum.

Brasenia purpurea. Taxus baccata.

Tilia platyphyllos. Pinus sylvestris.

Acer campestre. Picea excelsa.

Vaccinium uliginosum. Stratiotes aloides.

Oxycoccus. Typha.

Fraxinus. Potamogeton natans.

Menyanthes trifoliata. Najas major.

Myrica Gale. Scirpus lacustris.

Betula. sp.

Alnus. Eriophorum vaginatum.

Quercus sessiliflora. angustifolium (?).

Corylus Avellana. Carex echinata.

Fagus sylvatica. Goodenoughii (i*).

Salix aurita. Phragmites communis.
cinerea. Aira caespitosa (?).

Faskine, Lanark.

(Bennie, * On the occurrence of Peat with Arctic Plants
in Boulder Clay at Faskine, near Airdrie, Lanarkshire,'
Trans, Geol. Soc. Glasgow, Vol. X., pp. 148-152. 1895.)

The Boulder Clay here contains masses of transported
peat full of moss and leaves of Arctic willows. Though
here provisionally classed as Interglacial, they may per-
haps be of the same date as the Late Glacial deposits of
Hailes and Corstorphine.

Viola palustris. Salix herbacea.

Stellaria. Sedges — 3 or 4 species.

Potentilla (.''). Isoetes lacustris.
Hippuris vulgaris.

Deposits containhig Fossil Plants. 69

Fenland.

(Skertchly, 'Geology of the Fenland,' p. 320, Me^n.
GeoL Survey. 1877; Miller and Skertchly, 'The Fenland
past and present,' p. 341. 1878.)

The peat and ' submerged forests ' of the Fenland yield
numerous remains of trees. The following list of the
plants was compiled by Mr. A. Bell, but I have not seen
the specimens, and cannot trace the authorities for some
of the species. Betula nana is unrecorded elsewhere in
deposits of so recent a date.

Fraxinus. Salix Caprea.

Ulmus. repens.

Betula alba. Taxus baccata.

nana. Pinus sylvestris.

Quercus Robur. Juncus aquaticus.

Fagus sylvatica. Lastraea.

FiLLYSIDE, NEAR EDINBURGH.

(Bennie, ' The Raised Sea-Bottom of Fillyside.' Proc.
R. Phys. Soc. Edinburgh, Vol. XL, pp. 215-237. 1892.)

Some drifted seeds occur associated with the marine
shells. The deposit is here classed provisionally as Neo-
lithic, for the fauna and flora consists entirely of recent
British forms, without the Arctic species found in the
Clyde Beds.

Ranunculus Flammula. Taraxacum officinale.

repens. Stachys palustris.

Viola. Ajuga reptans.

Lychnis diurna (i*). Atriplex patula.

Stellaria media. Rumex.

Montia fontana. Mercurialis perennis.

Rubus Idseus. Alnus glutinosa.

Sambucus nigra. Carex.

70 Origin of the British Flora.

Garvel Park, Greenock.

(Robertson, ' On the Post-tertiary Beds of Garvel
Park, Greenock.' Trans. Geol. Soc. Glasgow , Vol VII.,
pp. 1-37. 1881.)

Marine clays belonging to the Clyde Beds contain a

sub- Arctic fauna, and are therefore classed as Late Glacial ;

they ought, perhaps, to be included in the Neolithic series,

for dug-out canoes are stated to have been found at some

places in these clays. The plants from Garvel Park sent

to me by Mr. Thomas Scott do not suggest an Arctic

climate, such as is apparently indicated by the marine

fauna.

Ranunculus repens. Bartsia Odontites.

Lychnis Flos-cuculi. Atriplex patula.

Rubus Idaeus. Rumex crispus.

CEnanthe Lachenalii. Sparganium ramosum (?).

Taraxacum officinale. Isoetes lacustris.

Gayfield, Edinburgh.

From a peaty deposit Mr. James Bennie has recently
obtained a number of leaves and seeds. At this locality,
as at Hailes, two different plant-beds are apparently repre-
sented. The three Arctic Willows suggest a climate like
that of the North Cape ; the Hawthorn and Wild-Cherry
point to a climate as mild as that now possessed by the
Scottish Lowlands.

Ranunculus aquatilis.

Atriplex (?).

repens.

Polygonum Persicaria.

Viola palustris.

Salix polaris.

Prunus Avium.

herbacea.

Rubus Idaeus.

reticulata.

Crataegus Oxyacantha.

Potamogeton crispus.

Myriophyllum.

sp.

Carduus.

Carex.

Sonchus arvensis.

Phragmites.

Menyanthes trifoliata.

Isoetes.

Deposits containing Fossil Pla^its. 71

Grays, Essex.
(Tylor, ' On Quarternary Gravels/ Quart. Journ. Geol.
Soc, Vol. XXV., p. d>2>. 1869 ; Reid, ' Pleistocene Plants
from Casewick, Shacklewell, and Grays,' ibid. Vol. LIII.,
p. 464. 189;.)

The plants collected by Prestwich occur associated with
or below the remains of Mammoth and Corbicula fluminalis.
They point distinctly to a temperate climate and mild
winters, for the Ivy is extremely sensitive to winter cold.
Both the character of the flora and the position of the
deposit suggest correlation with the temperate plant-beds
of Hoxne, which lie between the Boulder Clay and the
deposit with Arctic species.

Ranunculus repens. Populus canescens (?).

Rubus fruticosus. Salix sp.

Rosa. Potamogeton.

Hedera Helix. Cyperus (?).

Ulmus (?). Phragmites (.?).

Alnus glutinosa. Grass nodes.
Quercus Robur, var. sessiliflora. Equisetum.
Corylus Avellana.

Greenock (Roxburgh St.).

From the Clyde Beds (Late Glacial or Neolithic)

Mr. Thomas Scott obtained the following species : —

Potentilla Tormentilla (i*). Carex.

Taraxacum officinale. *Anthoxanthum odoratum.

Thymus Serpyllum. *Poa trivialis.

Atriplex patula.

Grunenthal, Holstein.

Weber, Neues Jahrb. Mineralogie, Geologie . . . . 189 1,
Vol. II., pp. 62-85, 228-230; and 1893, Vol. I., pp. 94-96.)

* These I think are recent specimens ; they are not in the same
state of preservation as the others, and are therefore omitted in the
summary. It is extremely difficult to prevent the adherence of light
grass-seeds when removing lumps of clay.

72 Origin of the British Flora.

The cuttings for the North Sea Canal showed silted-up
channels with Pleistocene plants. One at Beldorf exposed
a trough cut in the boulder-clay, filled with deposits yielding
temperate plants, at the top of which occurred a layer with
Betula nana, the whole being levelled up and hidden by
recent peat. This intercalation of a temperate flora be-
tween the boulder-clay and an Arctic plant-bed agrees
with the succession found at Hoxne in Suffolk. Another
channel at Grossen-Bornholt is apparently of the same
date. The plants occur in several different beds, full
details being given in Dr. Weber's papers.

Ranunculus. Alnus glutinosa.

Nuphar luteum. Carpinus Betulus.

Nymphsea alba. Corylus Aveilana.

Brasenia purpurea. Ouercus Robur.

Tilia platyphyllos. Salix pentandra (?).

Ilex aquifolium. Caprea.

Acer. Ceratophyllum demersum.

Prunus Avium. Juniperus communis.

Hippurus vulgaris. Picea excelsa.
Myriophyllum spicatum. Pinus sylvestris.

Trapa natans. Stratiotes aloides.

Galium uliginosum. Typha.

Vaccinium Vitis-Idaea. Potamogeton natans.

Myrtillus. Najas flexilis.

Andromeda Polifolia. Eriophorum.
Fraxinus excelsior {}). Carex panicea.
Menyanthes trifoliata. Holcus.
Betula alba. Phragmites.
nana.

Hailes, near Edinburgh.

(J. Geikie, 'Great Ice Age,' 3rd edit., p. 99. 1894;
Bennie, ' Arctic Plants in the old Lake Deposits of Scot-
land,' Ann. Scottish Nat. Hist., 1894, pp. 46-52.)

Two plant-bearing deposits are found at this spot.
The lower one rests immediately on the Boulder Clay and

Deposits containing Fossil Plants.

73

yields Arctic plants in the hollows between the boulders,
the species sent to me by Mr. Bennie being : —

Thalictrum minus (?).
Ranunculus aquatilis.

repens.

Viola palustris.
Lychnis diurna.
Stellaria media.
Oxalis Acetosella.
Hippuris vulgaris.
Taraxacum officinale.
Andromeda Polifolia.
Menyanthes trifoliata.

Stachys palustris.
Ajuga reptans.
Polygonum aviculare.
Alnus glutinosa.
Salix herbacea.

polaris,

reticulata,

Empetrum nigrum.
Eleocharis palustris.
Scirpus pauciflorus.
Isoetes lacustris.

The newer deposit, resting immediately upon this
Arctic plant-bed, contains Temperate species, among which
will be observed various plants usually considered to be
only present in this country as weeds of cultivation. The
species collected by Mr. Bennie are as follows : —

Ranunculus aquatilis.

Flammula.

Lingua (?).

repens.

Caltha palustris.
Viola palustris.
Lychnis diurna.

Flos-cuculi.

Stellaria media.

uliginosa.

Montia fontana.
Linum. i

Oxalis Acetosella.
Prunus spinosa.

Padus.

Spiraea Ulmaria.
Rubus Idaius.

fruticosus.

Potentilla Tormentilla (?).

Comarum.

Crataegus Oxyacantha.

Hippuris vulgaris.
yEthusa Cynapium.
Sambucus nigra.
Valeriana officinalis.
Chrysanthemum segetum.
Matricaria inodora.
Cnicus palustris.
Lapsana communis.
Menyanthes trifoliata.
Pedicularis palustris.
Lycopus europaeus.
Stachys palustris.
Galeopsis Tetrahit.
Ajuga reptans.
Atriplex patula.
Polygonum Persicaria.
Rumex crispus.
Mercurialis perennis.
Betula alba.
Alnus glutinosa.
Corylus Avellana.

74 Origin of the British Flora.

Quercus Robur. Scirpus lacustris.

Pinus sylvestris. Carex dioica.

Potamogeton heterophyllus. echinata.

perfoliatus. canescens.

pusillus. flava.

Eleocharis palustris. Isoetes lacustris.
Scirpus pauciflorus. Chara.
setaceus.

Happisburgh, Norfolk.

(Reid, 'Geology of Cromer/ Mem. Geol. Survey, 1882;
Reid, * Pliocene Deposits of Britain,' Mem. Geol. Survey,
1890.)

Slabs of clay-ironstone full of leaves and twigs are
thrown up by storms at this spot. They belong to the
lower part of the Preglacial Cromer Forest-bed. This
locality is the only one where determinable Preglacial
leaves are found in any quantity.

Cornus sanguinea. Fagus sylvatica.

Ulmus. Salix, 2 sp.

Betula alba. Pinus sylvestris.

Alnus glutinosa. Picea excelsa (cone).
Quercus Robur.

HiTCHiN, Hertfordshire.

(Reid, *The Palaeolithic Deposits at Hitchin and their
Relation to the Glacial Epoch,' Proc. Royal Soc, Vol. LXI.,
pp. 40-49. 1897.)

The plant-bearing deposits rest in a hollow eroded
in the Glacial beds, underlie brick-earth with Palaeolithic
implements, and apparently correspond with the Inter-
glacial deposits at Hoxne, though the overlying stratum
with Arctic plants has not been discovered at Hitchin.

Ranunculus aquatilis. Montia fontana.

sceleratus. Prunus spinosa.

: repens. Poterium officinale.

Deposits containing Fossil Plants.

75

Pyrus torminalis (?).
Hippuris vulgaris.
Myriophyllum.
Cornus sanguinea.
Sambucus nigra.
Eupatorium cannabinum.
Fraxinus excelsior.
Menyanthes trifoliata.
Lycopus europaeus.
Ajuga reptans.

Alniis glutinosa.
Quercus Robur.
Ceratophyllum demersum.
Sparganium.
Potamogeton crispus.

2 sp.

Najas marina.
Scirpus lacustris.

sp.

Carex.

HoLMPTON, Yorkshire.

(Reid, ' Geology of Holderness,' p. 85, Mem. Geol.
Survey. 1885.)

A hollow in boulder-clay, filled with peaty loam,
contains freshwater shells, and leaves of Betula nana, it is
classed therefore as Late Glacial.

HONERDINGEN, HaNOVER.

(Weber, ^ Ueber die fossile Flora von Honerdingen und
das nordwestdeutsche Diluvium, Abh. Naturw. Ver.
Bremen. Vol. XIII., pp. 413-468. 1896.)

The deposits are considered by Dr. Carl Weber to be
of Interglacial date. He compares them with those of
Fahrenkrug, Griinenthal, Klinge, and Lauenburg. The
occurrence of Platanus, Juglans, and Najas Jiexilis is
noticeable.

Thalictrum flavum.
Ranunculus Lingua.
Nuphar luteum.
Nymphaea alba.
Tilia platyphyllos.

parvifolia.

intermedia.

Ilex Aquifolium.
Rhamnus Frangula.
Acer platanoides.

Rubus Idaius.
Hippuris vulgaris.
Cornus sanguinea.
Fraxinus excelsior.
Menyanthes trifoliata.
Myrica Gale.
Betula pubescens.

alba (.?).

Alnus glutinosa.
Carpinus Betulus.

76

Origin of the British Flora.

Corylus Avellana.

Quercus sessiliflora.

Fagus sylvatica.

Salix.

Populus tremula.

Platanus (?).

Juglans regia (?).

Empetrum nigrum.

Ceratophyllum submersum.

demersum.

Typha.

Sparganium minimum.

simplex.

Potamogeton natans.

cf. polygon i-

folia.
Potamogeton rufescens.

cf. colorata.

graminea.

cf. praelonga.

perfoliata.

crispa.

Potamogeton compressa.

obtusifolia.

pusilla.

rutila.

cf. trichoides.

— manna.

Najas major.

flexilis.

Scirpus lacustris.
Eriophorum vaginatum (?).
Carex acuta ij).
cf. acutiformis.

rostrata.

Phragmites communis.
Taxus baccata.
Juniperus communis.
Pinus sylvestris.
Abies pectinata.
Picea excelsa.
Equisetum palustre.
Polystichum cf. Thelypteris.

Hornsea, Yorkshire.

(Phillips, 'Geology of Yorkshire,' 3rd edit. Part I.,
pp. 75-79. 1875 ; Reid, 'Geology of Holderness,' Mem.
Geol. Survey, pp. 79-83. 1885.)

Peaty mud fills a valley cut through the Glacial
deposits. It contains Mammoth (.-*), Irish Elk, Lion, and
Bos primigenius {?). The mollusca and plants are all
recent British forms.

Prunus Padus.

Alnus glutinosa.
Quercus Robur.
Salix.

Pinus sylvestris.

Potamogeton.

Chara.

The stratigraphical position of this deposit and its resem-
blance to other ' Submerged forests ' suggest a Neolithic or
Late Glacial Age. The occurrence of the Mammoth, Lion,

Deposits containing Fossil Plants.

77

and Bos primigenius — if these were really obtained from
it — point, on the other hand, to an earHer period.

HoxNE, Suffolk.

(' Report of the Committee on the Relation of Palaeo-
lithic Man to the Glacial Epoch,' Rep. Brit. Assoc, for
1896, pp. 400-415, 1897.)

Ancient Alluvial deposits fill a channel newer than, and
eroded through, the chalky boulder-clay, but independent
of the existing valley system. Several plant-bearing zones
are seen in direct superposition, A being the most recent : —
A, B. — Brick-earth and gravel with Palaeolithic implements
freshwater shells and bones of Elephant.

Alnus (?). Chara.

Potamogeton.

C. — Black earth, with freshwater shells, leaves and seeds of
Arctic plants.

Ranunculus aquatilis.

— sceleratus.

repens [deriv-
ative (?)].

Caltha palustris.

Viola palustris.

Stellaria media.

Montia fontana.

Rhamnus Frangula [worn
and derivative].

Rubus Idaeus.

Poterium officinale.

Hippuris vulgaris.

Myriophyllum spicatum.

CEnanthe Phellandrium.

Sambucus nigra [deriv-
ative].

Eupatorium cannabinum.

Bidens tripartita.

Taraxacum officinale.

Menyanthes trifoliata.

Lycopus europaeus.
Ajuga reptans.
Rumex maritimus.

crispus Q).

Urtica dioica ( } ) [one

seed].

Betula nana.

Alnus glutinosa [perhaps
derivative].

Carpinus Betuius [deriv-
ative].

Salix myrsinites.

herbacea.

polaris.

Ceratophyllum demersum.
Taxus baccata [ deriv-
ative].

Sparganium ramosum.
Alisma Plantago.
Potamogeton rufescens.
crispus.

78 Origin of the British Flora.

Potamogeton pusillus. Scirpus lacustris.

trichoides. Blysmus rufus.

pectinatus. Carex incurva (?).

Scirpus pauciflorus. Chara.
setaceus.

D. — Lignite with Temperate plants.

Ranunculus aquatilis. Rumex Acetosella (?).

sceleratus. Urtica dioica (?).

Lingua. Alnus glutinosa.

cf. repens. Carpinus Betulus.

Montia fontana. Corylus Avellana.

Rhamnus Frangula. Ceratophyllum demersum.

Rubus Idaius. Taxus baccata.

Rosa canina. Sparganium ramosum.

Pyrus torminalis if). Alisma Plantago.

CEnanthe Phellandrium. Potamogeton pusillus.

Sambucus nigra. trichoides.

Eupatorium cannabinum. Eleocharis acicularis.

Bidens tripartita. Scirpus pauciflorus.

var. with setaceus

four equal awns. lacustris

Mentha aquatica. Blysmus rufus.

Lycopus europaeus. Eriophorumangustifolium.

Stachys (?). Carex distans (?).

Rumex maritimus. ampullacea (?).

: crispus.

E. — Clay with freshwater shells, fish-bones, and Temperate
plants.

Ranunculus Lingua. Ceratophyllum demersum.

repens. Sparganium ramosum.

Rubus Idaeus. Potamogeton trichoides.

Hippuris vulgaris. Zannichellia palustris.

Rumex maritimus. Scirpus lacustris.

Alnus glutinosa. Carex.

As beds D and E, containing a temperate flora, lie

above the Glacial deposits and below loam with Arctic
plants, they are classed as Interglacial. Bed C, containing

Arctic plants, is newer than the boulder-clays of Suffolk,
and is therefore called Late Glacial.

Deposits containing Fossil Plants. jq

Ireland (120 miles west of Dublin).
(Reid, 'The Origin of Megaceros-marl,' Irish Natu-
ralist, May, 1895.)

A sample of the marl which yields the skeletons of
the Irish elk was sent to me by Mr. W. Williams of
Dublin. The exact locality was not stated. The deposit
is a Chara-marl full of seeds of Pond-weeds, with a few other
plants.

Ranunculus aquatilis. Eleocharis palustris.
Myriophyllum spicatum. Carex (}).
Littorella lacustris. Scirpus (.?).
Potamogeton crispus. Chara (several species).
praelongus.

The exact date of these marls is still uncertain, for the
associated deposits have not yet been properly examined.
Above similar Megaceros-marls at Ballaugh in the Isle of
Man is found peat with Salix herbacea. No fossil Arctic
plants have yet been found in Ireland, and the deposit is
therefore provisionally classed with the Neolithic peat-
mosses.

Kelsey Hill, Yorkshire.

(Reid, 'Geology of Holderness,' pp. 74, 75. Mem.
Geol. Survey. 1885.)

Peaty clay caps an isolated sand- hill rising about 40 feet
above the Humber marsh. The plants are : —

Ranunculus aquatilis. Phragmites.

Potamogeton.

The exact age of the deposit is doubtful ; for, though
provisionally classed as Neolithic, it may be Late Glacial.

KiLMAURS, Ayrshire.

(Bennie, ' Note on the Contents of Two Bits of Clay
from the Elephant Bed at Kilmaurs in 1817,' Proc. R.

8o Origin of the British Flora.

Phys. Soc, Edinburgh, 1885, pp. 451-459; Craig, 'On the
Post-Pliocene Beds of the Irvine Valley, Kilmaurs, and
Dreghorn Districts,' Trans. Geol. Soc. Glasgow, 1887,
pp. 213-226.)

The deposits occur beneath Boulder Clay, they yield
remains of Mammoth and of the following species of
plants : —

Ranunculus aquatilis. *Potamogeton Zizii or
Potentilla (?). heterophyllus.

Hippuris vulgaris. Zannichellia palustris.

Myriophyllum spicatum. Chara.

*Potamogeton rufescens (?). Isoetes lacustris.

Kirk Michael, Isle of Man.

(Lamplugh, Annual Rep. Geol. Survey for 1895, p. 13.)

The plants occur in a peaty layer at a depth of 1 5 feet.
They probably belong to the same period as the upper beds
at Ballaugh, and the Arctic plant-beds near Edinburgh.

Ranunculus aquatilis. Salix herbacea.

Viola palustris. Potamogeton.

Potentilla Comarum. Eleocharis palustris.

Hippuris vulgaris. fCarex alpina.

Menyanthes trifoliata. sp.

KiRMiNGTON, North Lincolnshire.

(Reid, 'Geology of Holderness,' pp. 58, 59,69, 70, Mem.
Geol. Surrey. 1885.)

Estuarine warp, peat, and shingle occur at a height of
about 80 feet above the sea. The peat is a mass of the
common Reed, among which I could find no other plants.
The warp contained : —

Scirpus fluitans. Phragmites communis.

* Determined by Mr. A. Bennett,
t Determined by Mr. C. B. Clarke.

Deposits containing Fossil Plants. 8i

Apparently a littoral deposit of the same date as the
Interglacial marine gravels which occupy the lower grounds,
and lie between two boulder-clays.

Klinge bei Cottbus, Prussia.

CNehring, * Ueber Wirbelthier-Reste von Klinge,' Neues
Jahrb. filr Mineralogie, 1895, pp. 183-208 ; Nehring, ' Ueber
Elephas-yio\2.x^x\., aus dem diluvialen Torflager von Klinge
bei Cottbus,' Gesellsch. naturf. Freunde^ 20th Oct., 1896.)

The deposit is probably of Interglacial date. The plants
are associated with remains of Mammoth, Rhinoceros, Horse,
Reindeer, and a species of Megaceros closely allied to the
Irish Elk. The determination of the plants is mainly due
to Dr. C. Weber.

Thalictrum flavum.

Salix aurita.

Nuphar luteum.

cinerea.

Nymphaea alba.

repens.

Brasenia ovulum.

Caprea (?).

Tilia platyphyllos (J).

Populus tremula.

Ilex aquifolium.

Ceratophyllum submersum.

Acer campestre.

demersum.

Comarum palustre.

Taxus baccata.

Hippuris vulgaris.

Pinus sylvestris.

Myriophyllum.

Picea excelsa.

Galium palustre (?).

Stratiotes aloides.

Vaccinium Oxycoccos.

Echinodorus ranunculoides(?)

Menyanthes trifoliata.

Potamogeton natans.

Betula verucosa.

Najas marina.

odorata.

Scirpus lacustris.

Alnus.

pauciflorus (?).

Carpinus Betulus.

Cladium Mariscus.

Corylus Avellana.

Carex, several sp.

Quercus.

Polysticum Thelypteris.

Lauenburg an der Elbe.

(Keilhack, * Ueber ein interglaciales Torflager im
Diluvium von Lauenburg an der YXh^l Jahrb. der k.preuss.
geolog. Landesanstalt fiir 1884. Berlin, 1885 ; Nathorst,

G

82 Origin of the British Flora,

' Eine Probe aus dem Torflager bei Lauenburg an der
Elbe,' Natiirwissensch. Wochenschrift, ^.th Nov., 1894.)

The exact relation of this peat to the Glacial deposits
is not clear, and Professor Nathorst suggests that more
than one plant-bed is represented. The occurrence of
Brasenia suggests an Interglacial date.

Nymphasa alba. Lycopus europaeus.

Brasenia purpurea. Ulmus.

Corydalis fabacea. Alnus glutinosa.

Viola. Carpinus Betulus.

Arenaria trinervia. Corylus Avellana.

Tilia platyphyllos. Quercus Robur.

Geranium columbinum. Salix aurita (.?).

Rhamnus Frangula. Salix repens.

Acer platanoides. Pinus sylvestris,

Trapa natans. Picea vulgaris.

Cornus sanguinea. Larix europaea.

Viburnum Opulus {}). Iris Pseudacorus.
Lysimachia Nummularia (J). Sparganium.

Vaccinium Oxycoccos. Potamogeton.

Fraxinus excelsior. Carex Pseudo-cyperus.
Menanythes trifoliata.

MUNDESLEY, NORFOLK.

(Reid, ' Geology of the Country around Cromer,' pp. 36,
37, 83, 84, 118, 119, and folding plate, Mem. Geol. Survey^
1882; also 'Pliocene Deposits of Britain,' pp. 166-169.
ibid. 1890.)

At this locality three different plant-bearing deposits
are represented. The oldest is Preglacial, and belongs to
the Cromer Forest-bed, which is here divisible into an
upper and a k>wer freshwater deposit, between which is a
mass of estuarine gravel. The lower freshwater bed
consists of laminated peat, full of fruits of Trapa natans^
but containing little else. The middle or Estuarine divi-
sion, contains bones of extinct mammals, much drift-wood,
artd cones of Pinus sylvestris and Picea excelsa. The

Deposits containing Fossil Plants. Zt^

upper freshwater bed is a thin seam of lacustrine clay, full
of seeds of aquatic and marsh plants. The flora of this
deposit is so uniform at most of the localities, that it is
needless to repeat the list.

Above the Forest-bed lies an Early Glacial flood-loam
or loess- like deposit containing bones of a Spermophilus
and leaves and seeds of Arctic plants, the species being : —

Hippuris vulgaris. Salix polaris.

Still higher, and cutting through the boulder clay, is
seen an old river channel, subsequently silted-up with
Alluvial mud containing remains of the water-tortoise
{Emys lutaria), shells of Hydrobia marginata^ and plants,
the species observed being Nuphar luteum^ Ceratophyllum
demersum^ and Salix. This deposit is probably equivalent
to beds D and E at Hoxne, and is here provisionally
classed as Interglacial.

Northampton.

An Alluvial deposit of uncertain age yields the
following plants, sent to me by Mr. H. N. Dixon: —

Nuphar luteum. Polygonum.

Stellaria media. Mercurialis perennis.

Prunus spinosa. Alnus.

Padus. Corylus Avellana.

Sambucus nigra. Quercus Robur.

OsTEND, Norfolk.

(Reid, * Geology of the Country around Cromer,' pp.
41-43, 62-65, Mem. Geol. Survey. 1882; * Pliocene Deposits
of Britain,' pp. 171, 195, ibid. 1890.)

Two distinct plant-bearing deposits are here repre-
sented. The older belongs to the Preglacial Cromer
Forest-bed, and contains cones of Picea excelsa and fruits
of Trapa 7iatans. The newer is Early Glacial, contains

84 Origin of the British Flo7^a.

Arctic plants, and corresponds to the similar deposit at
Mundesley, Salix polaris being abundant. The older deposit
yields : —

Thalictrum flavum. Alnus glutinosa.

Ranunculus aquatilis. Corylus Avellana.

Hippuris vulgaris. Quercus Robur.

Trapa natans. Taxus baccata.

Cornus sanguinea. Pinus sylvestris.

Menyanthes trifoliata. Picea excelsa.
Rumex maritimus.

The Early Glacial bed contains : —

Hippuris vulgaris. Potamogeton.

Betula nana. Carex.

Salix polaris.

OVERSTRAND, NORFOLK.

(Reid, * Geology of the Country around Cromer,' and
* Pliocene Deposits of Britain,' Mem. Geol. Survey, 1882
and 1890.)

At this locality the Preglacial Cromer Forest-bed is
full of drift-wood and fir-cones, and its upper, freshwater
division contains seeds of Cratcegus Oxyacantha — a plant
unknown elsewhere in Preglacial deposits. The other
plants are all common to several localities and the list
need not be repeated.

OVERTOUN, NEAR BEITH, AYRSHIRE.
(Craig, Trans. Geol. Soc. Glasgow, Vol. IV., p. 145.)
Hazel nuts are here said to occur between two masses
of till.

Oxford.

(A. M. Bell, ' On the Pleistocene Gravel at Wolvercote^
near Oxford,' Rep. Brit, Assoc, for 1894, p. 663.)

A Pleistocene alluvial deposit at Wolvercote, near

Deposits containing Fossil Plants.

85

Oxford, shown to me by the late Professor A. H. Green,
yields the following species of plants : —

Ranunculus aquatilis.
sceleratus.

repens.

Potentilla Tormentilla.

Viola.

Hippuris vulgaris.

Heracleum Sphondylium.
Potamogeton.
Eleocharis palustris.
Scirpus lacustris.
Carex rostrata.

This plant-bed lies above a deposit with bones of Bison
and Palaeolithic implements, and is of uncertain age.

Pakefield, Suffolk.

( Reid, ' Pliocene Deposits of Britain,' pp. 1 77- 1 79. Mem,
GeoL Survey. 1890.)

A silted-up channel belonging to the Preglacial Cromer
Forest-bed occurs at about two hundred yards south of the
Lighthouse Gap. It has yielded about fifty species of
flowering plants, among which are a few dry-soil species
unknown elsewhere in deposits of this date. The plants
are : —

Thalictrum flavum.
Ranunculus aquatilis.
sceleratus.

repens.

Nuphar luteum.
Viola palustris.
Hypericum quadrangulum.
Acer campestre.
Prunus spinosa.
Spiraea Ulmaria.
Rubus fruticosus.
Potentilla.
Pyrus Aria.
Trapa natans.
Myriophyllum spicatum.
Hydrocotyle vulgaris.
CEnanthe Lachenalii.
Phellandrium.

Heracleum Sphondylium.
Cornus sanguinea.
Bidens tripartita.
Lapsana communis.
Picris hieracioides.
Mentha aquatica.
Lycopus europaeus.
Atriplex patula.
Polygonum Persicaria.
Rumex maritimus.

sp.

Euphorbia amygdaloides.
Betula alba.
Alnus glutinosa.
Carpinus Betulus.
Corylus Avellana.
Quercus Robur.
Ceratophyllum demersum.

86 Origm of the British Flora.

Taxus baccata. Najas minor.

Sparganium ramosum. Eleocharis palustris (?).

Alisma Plantago. Scirpus pauciflorus.

Potamogeton lucens. lacustris.

trichoides. Carex remota.

Zannichellia palustris. paludosa (?).

pedunculata. riparia.

Najas marina. Phragmites communis.

Parkstone, Dorset.

(Reid, ' On Charred Pine-wood from Dorset Peat
Mosses/ Proc. Dorset Nat. Hist. Field Club, Vol. XVI.,
p. 14. 1895-)

The bottom layers of a peat-moss here contain trunks
and cones of Pinus sylvestris.

Redhall, near Edinburgh.

(Henderson, * On some Sections of Boulder Clay, Peat,
and Stratified Beds exposed in a Quarry recently opened
at Redhall, Slateford, near Edinburgh.' Trans. Geol. Soc.^
Edinburgh, Vol. II., p. 391. 1874 ; Geikie, ' Great Ice Age,'
3rd Edit., pp. icx), lOi. 1894; Bennie and Scott, ' The
Ancient Lakes of Edinburgh/ Proc. R. Phys. Soc, Edin-
burgh, Vol. X., pp. 126-154. 1889.)

The Deposit at Redhall, like that of Cowden Glen,
has been generally accepted as of Interglacial age. The
many weeds of cultivation, capsules of Flax, and pieces of
charcoal, that it contains can scarcely belong to any earlier
period than the Neolithic. The flora closely corresponds
with that of the upper bed at Hailes.

Ranunculus aquatilis. Lychnis Flos-cuculi.

Flammula. Stellaria media.

Lingua. uliginosa.

repens. Spergula arvensis.

Caltha palustris. Montia fontana.

Fumaria ofificinalis. Hypericum quadrangulum.

Viola palustris. elodes.

Lychnis diurna. Linum.

Deposits containing Fossil Plants.

87

Oxalis Acetosella.
Prunus.

Spiraea Ulmaria.
Rubus Idseus.

fruticosus.

Potentilla Tormentilla.

Comarum.

Alchemilla arvensis.
Poterium officinale.
Rosa canina 7).
Crataegus Oxyacantha.
Sambucus nigra.
Valeriana officinalis.
Eupatorium canabinum.
Bidens cernua.
Crysanthemum segetum.
Matricaria inodora.
Tussilago Farfara.
Senecio sylvaticus.
Cnicus lanceolatus.

palustris.

Centaurea Cyanus.
Lapsana communis.
Crepis virens.
Leontodon autumnalis.
Taraxacum officinale.
Sonchus arvensis.
Menyanthes trifoliata.
Pedicularis palustris.
Lycopus europaeus.
Prunella vulgaris.

Stachys palustris.
Galeopsis Tetrahit.
Ajuga reptans.
Littorella lacustris.
Atriplex patula.
Polygonum aviculare.

Persicaria.

Rumex obtusifolius.

crispus.

Euphorbia Helioscopia.
Alnus glutinosa.
Quercus Robur.
Pinus.

Juncus glaucus (?).
Sparganium ramosum.
Alisma Plantago.
Potamogeton perfoliatus,

pusillus.

Eleocharis palustris.
Scirpus pauciflorus.

setaceus.

Carex dioica.

echinata.

canescens.

panicea.

flava.

* Holcus lanatus.
Agrostis.

* Poa trivialis.

* Dactylus glomerata.

Sand le Meer, Yorkshire.

(Reid, 'Geology of Holderness,' p. 84, Mem. Geol.
Survey. 1885.)

In a ' submerged forest ' of the ordinary type, opposite
the mouth of a small valley, the plants observed were : —

Prunus Padus. Salix.

Alnus glutinosa. Juncus.

Corylus Avellana. Potamogeton.

Quercus Robur. Carex.

* Probably recent specimens.

88 Origin of the British Flora.

Selsey, Sussex.

(Reid, ' The Pleistocene Deposits of the Sussex Coast,
and their Equivalents in Other Districts,' Quart. Journ.
Geol. Soc, Vol. XLVIIL, pp. 344-366. 1892.)

Carbonaceous river-mud here overlies Glacial erratics

and underlies the Palaeolithic deposits. The river-mud is

apparently of Interglacial date, and corresponds closely in

position and fossil contents with the strata found at West

Wittering and Stone. The plant-remains consist of drifted

seeds, Acer monspessulanum giving a southern aspect to

the flora : —

Acer monspessulanum. Atriplex patula.

Prunus Avium. Polygonum aviculare.

Padus. Quercus Robur.

Rubus fruticosus. Zannichellia palustris.

Rosa. Scirpus pauciflorus (.?).

Ajuga reptans. Carex distans (i*).

Shacklewell, Middlesex.

(Prestwich, ' On a Fossiliferous Deposit in the Gravel
at West Hackney,' Quart. Journ. Geol. Soc.^ Vol. XL,
p. 107. 1885; Reid, * Pleistocene Plants from Casewick,
Shacklewell and Grays.' Ibid, Vol LI 1 1., pp. 463, 464.
1897.)

Peaty clay is found beneath 8 or 10 feet of gravel.
Though none but British species of Mollusca or plants
have yet been discovered, the geology suggests a con-
siderable antiquity. The plants are Temperate species : —

Ranunculus repens. Lycopus europaeus.

Rubus Idaeus. Alnus glutinosa.

Rosa. Quercus Robur.
Eupatorium cannabinum.

SiDESTRAND, NORFOLK.
(Reid, ' Geology of the Country around Cromer.' 1882 ;
and * Pliocene Deposits of Britain,' Mem. Geol. Survey.
1890.)

Deposits containing Fossil Plants. 89

The upper part of Preglacial Cromer Forest-bed is here
a bed of blue lacustrine clay, full of freshwater shells and
seeds of aquatic and marsh plants. Dry-soil species and
forest trees are absent. Among the plants are Trapa
natans and Stratiotes aloides.

SiLCHESTER ROMAN STATION, HAMPSHIRE.

(Hope and Fox, ' Excavations on the site of the Roman
City at Silchester, Hants, in 1890-1897,' ArchcsologiayW o\s,
L.-LVn. 1 891-1898.)

Some material sent to me in April of the present year
(1899) by Mr. A. H. Lyell, from one of the excavations
made to explore this station, contained a number of seeds
which belong to the date of the Roman occupation. They
do not include cultivated plants of any sort, with the
doubtful exception of a single capsule of flax ; but among
them are several weeds of cultivation, the seeds of which
are small and starved, as though growing on exhausted
land that for some years had been out of cultivation or
occupation.

Thalictrum flavum. Conopodium denudatum.

Ranunculus Flammula. ^thus aCynapium.

Lingua. Sambucus nigra.

repens. Galium.

Sardous. Chrysanthemum Leucan-

Caltha palustris. themum.

Papaver Argemone. Mentha aquatica.

Thlaspi arvense. Prunella vulgaris.

Lychnis Flos-cuculi. Stachys arvensis

Stellaria media. Galeopsis Tetrahit.

graminea. Atriplex.

Hypericum perforatum. Polygonum Aviculare.

Linum usitatissimum (?). Rumex conglomeratus.

Spiraea Ulmaria. Eleocharis acicularis.

Rubus fruticosus. , palustris.

Potentilla Tormentilla. Carex.
Alchemilla arvensis.

90 Origin of the British Flora.

\

Southampton Docks,

(Shore and Elwes, ' The New Dock Excavation at
Southampton,' Proc. Hants Field Club for 1889, pp. 43-56.)

A bed of peat and shell-marl beneath the sea-level
yields Neolithic implements, and is said to contain Oak,
Beech, Hazel, Birch, and Pine, besides decomposed remains
of Scirpus laaistris, Carex, Myrica Gale, heaths, Pteris
aquiiina. A small sample given me by Mr. Whitaker
contained seeds of: —

Rubus Idaeus. Corylus Avellana.

Sambucus nigra. Scirpus maritimus (?).

SOUTHELMHAM, SUFFOLK.

(Candler, ' Observations on some Undescribed Lacus-
trine Deposits at Saint Cross, South Elmham, in Suffolk,'
Quart. Journ. Geol Soc, Vol. XLV., pp. 504-510. 1889.)

The plant- bearing stratum yields bones of Elephant,
and probably agrees with the Interglacial beds D and E at
Hoxne. It overlies the Boulder Clay, but is not overlain
by any newer deposit.

Thalictrum flavum. . Ceratophyllum demersum.

Ranunculus aquatilis. Stratiotes aloides.

sceleratus. Alisma Plantago.

- Flammula. Potamogeton heterophyllus.

Crataegus Oxyacantha. perfoliatus.

Hippuris vulgaris. crispus.

Myriophyllum spicatum. obtusifolius.

Hydrocotyle vulgaris. trichoides.

QEnanthe Phellandrium. Zannichellia palustris.

Cnicus palustris (.?). Scirpus pauciflorus.

Taraxacum officinale. — caespitosus.

Menyanthes trifoliata. fluitans.

Lycopus europaeus. lacustris.

Rumex maritimus. Carex riparia.

Alnus glutinosa. rostrata.

Deposits containing Fossil Plants. 91

Stoke Newington, London.

(W. G. Smith, 'Man the Primeval Savage,' pp. 288, 289.
8vo, London, 1894.)

The Palaeolithic deposits here have yielded various
plants ; but the specimens deposited in the British Museum
by Mr. Worthington Smith are not at present available.
He mentions: —

Clematis Vitalba. (Leaves.)

Vitis vinifera. (Wood ; perhaps a recent specimen.)

Rubus (?), (Fragments of stems, with thorns.)

Ulmus.

Betula. (Bark and wood.)

Alnus glutinosa. (Leaves and catkins.)

Corylus.

Taxus.

Pinus.

Rushes. \

Sedges. > Impressions.

Grass. )

Aspidium Filex-mas. (Impressions of pinnae.)

Osmunda regalis. (Fronds and rhizomes.)

Stone, Hampshire.

(Reid, ' A fossiliferous Pleistocene Deposit at Stone, on
the Hampshire Coast,' Quart. Journ. GeoL Soc, Vol. XLIX.,
pp. 325-328. 1893.)

Carbonaceous river-mud here underlies the Palaeolithic
gravels. It is evidently equivalent to the Interglacial
deposits of West Wittering and Selsey, and contains
remains of Elephant and of the following plants : —

Ranunculus sceleratus. Myriophyllum.

repens. Caucalis nodosa.

Arenaria peploides. Valeriana officinale.

Acer monspessulanum. Mentha aquatica.

Rubus fruticosus. Lycopus europaeus.

Potentilla. Atriplex patula.

Rosa. Polygonum aviculare.

92 Origin of the British Flora,

Rumex. Zannichellia palustris.

Urtica. Eliocharis acicularis.

Quercus Robur. Scirpus lacustris.

Sparganium. Carex riparia (?).

Alisma Plantago. rostrata.

Potamogeton heterophyllus. muricata.

trichoides. Phragmites.

Ruppia maritima.

Sweden.

(Gunnar Andersson, ' Svenska Vaxtvarldens Historia,'
8vo, Stockholm, 1 896 ; and ' Geschichte der Vegetation
Schwedens/ Englers Bot. Jahrb., Bd. XXII., pp. 434-550.
1896.)

The extensive literature relating to Swedish Quaternary
fossil plants has been brought together by Dr. Gunnar
Andersson, who refers the deposits to the following five
zones, all corresponding, apparently, with our late Glacial
and Neolithic, no plant-bearing strata of Interglacial or
of Preglacial date being yet known in Sweden.

5 Spruce Zone.

4 Oak Zone.

3 Pine Zone.

2 Birch Zone.

I Dryas Zone.

No. I corresponds in all probability with the Arctic
plant-beds of Hailes, Corstorphine, and Gayfield.

As Dr. Gunnar Andersson records no fewer than 133
species of flowering plants, the table is too long here to be
reproduced ; but the range in Sweden, where it supple-
ments the British records, is mentioned in the next
Chapter under the heading of each species.

Tilbury, Essex.

(Spurrell, * On the Estuary of the Thames and its
Alluvium,' Proc. Geol. Assoc, Vol. XL, pp. 210-230. 1889.)

Deposits containing Fossil Plants, 93

The ' Submerged Forests ' met with during the excava-
tion of Tilbury Docks apparently belong to the same
period as those seen at Crossness and at the Albert Dock;
they underlie a layer with Roman remains, but the small
list of plants includes nothing characteristic of any par-
ticular date : —

Sambucus nigra. Quercus Robur sessiliflora.

Betula alba. Sparganium ramosum.

Alnus glutinosa. Carex.

Corylus Avellana. Phragmites communis.

Trimingham, Norfolk.

(Reid, 'Geology of the Country around Cromer/ 1882 ;
and ' Pliocene Deposits of Britain,' Mem. Geol, Survey,
1890.)

Good sections of the Preglacial Cromer Forest-bed can
be seen at the foot of the cliffs and on the foreshore at
Trimingham. The Early Glacial freshwater deposits may
also be represented there ; but I have not yet been able to
find any of the characteristic Arctic plants in them.

Twickenham, Middlesex.

(Leeson & Laffan, * On the Geology of the Pleistocene
Deposits in the Valley of the Thames at Twickenham.
, . . .' Quart. Joum. Geol. Soc.^ Vol L., pp. 453-462.
1894.)

A small silted-up channel is here found beneath the
Thames gravel. It is of interest as yielding mammals
which perhaps point to a transition between the Palaeo-
lithic and Neolithic periods. The species are the Bison,
Reindeer, Horse, and Bos longifrons. The plants include
Galeopsis Tetrahit, usually a weed of cultivation ; but there
are no definite signs of cultivated plants or domesticated

94 Origin of the British Flora.

animals. From the presence of Reindeer the deposit is
classed as Late Glacial.

Stellaria media. Potamogeton rufescens.

Montia fontana. Zannichellia palustris.
Heracleum Sphondylium. Eleocharis palustris.

Galeopsis Tetrahit. Scirpus lacustris.

Atriplex. Carex panicea.

Polygonum Persicaria. Phragmites.
Rumex crispus.

West Runton, Norfolk.

(Reid, ' Geology of the Country around Cromer,' 1882 ;
and 'Pliocene Deposits of Britain,' Mem. Geol. Survey.
1890.)

The upper part of the Preglacial Cromer Poorest-bed
is here represented by a mass of peat filling a shallow
channel. It is full of remains of animals and plants, but
the latter are not usually well preserved, and have not yet
been properly collected. They seem to include a some-
what larger proportion of dry-soil species than is usually
found in deposits of this age.

West Wittering, Sussex.

(Reid, ' The Pleistocene Deposits of the Sussex Coast.
. . . .' Quart. Journ, Geol. Soc, Vol. XLVIIL, pp. 344-
361. 1892.)

The plant-bearing strata yield a Temperate flora, but
contain at their base far-travelled erratic blocks, derived
from an earlier glacial deposit, and are overlaid by brick-
earth of Late Glacial date. The plant-bed corresponds
with those at Selsey and Stone, and contains remains of
Elephant, Rhinoceros, with some freshwater shells no
longer living in Britain. Local conditions being excep-
tionally favourable, the flora is unusually varied, fresh-
water, estuarine, sea-coast, marsh, dry-soil, woodland, and

Deposits containing Fossil Plants,

95

limestone species all being represented by their seeds.
Leaves, except badly preserved fragments, are not found.
The 94 species in the subjoined list were obtained by
Mrs. Reid and myself by washing about two hundred-
weight of the loamy sand. Only two of those determined
are now extinct in Britain, though among the undetermined
seeds are several well-marked forms, which do not belong
to any living British plants, but cannot yet be identified.
Najas minor and N. graminea are both southern forms.
Acer monspessulanum, so common at Selsey on the east,
and at Stone on the west, has not yet been found at West
Wittering. The exotic species probably number nearly
10 per cent. ; but in the absence of good collections of ripe
seeds their determination is very difficult.

Thalictrum flavum.
Ranunculus aquatilis.

hederaceus.

sceleratus.

Lingua.

repens.

bulbosus.

Sardous.

parviflorus.

Caltha palustris.
Nuphar luteum.
Nymphaea alba.
Chelidonium majus.
Silene maritima.
Lychnis diurna.

Flos-cuculi. •

Stellaria aquatica.

media.

Holostea.

Montia fontana.
Hypericum perforatum.

quadrangulum.

Ilex aquifolium.
Rhamnus Frangula.

Prunus spinosa.

Avium.

Spiraea Ulmaria.
Rubus fruticosus.
Potentilla Tormentilla.
Alchemilla arvensis.
Poterium officinale.
Rosa.

Hippuris vulgaris.
Myriophyllum spicatum.
Hydrocotyle vulgaris.
Apium graveolens.

nodiflorum.

Chaerophyllum temulum.
Anthriscus sylvestris.
CEnanthe fistulosa.

Phellandrium.

Angelica sylvestris.
Cornus Sanguinea.
Sambucus nigra.
Viburnum Opulus.

Lantana.

Valerianella olitoria.
Scabiosa succisa.

96

Origin of the British Flora.

Eupatorium cannabinium.
Aster Tripolium.
Senecio aquaticus.
Cnicus lanceolatus.

palustris.

Lapsana communis.
Hieracium Pilosella.
Taraxacum (?).
Glaux maritima.
Menyanthes trifoliata.
Solanum Dulcamara.
Verbascum Thaspus.
Mentha aquatica.
Lycopus europaeus.
Stachys palustris.
Ajuga reptans.
Atriplex patula.
Polygonum Persicaria.
Rumex conglomeratus.
obtusifolius.

Corylus Avellana.
Quercus Robur.
Salix cinerea.
Ceratophyllum demersum.
Sparganium ramosum.
Alisma Plantago.
Sagittaria sagittifolia.
Potamogeton natans.

heterophyllus.

lucens (?).

densus.

trichoides.

Ruppia maritima.
Zannichellia pedunculata.
Najas minor.

grammea.

crispus.

Mercurialis perennis.
Betula alba.

Eleocharis acicularis.

palustris.

Scirpus lacustris.
Carex muricata.

distans.

rostrata.

riparia.

Weymouth.

(Gepp, ' Fossil Plant-remains in Peat,' Journ. Botany^
Vol. XXXIII., pp. 180-182. 1895.)

Cakes of compressed peat thrown up near Weymouth
have yielded a few plants, among which is the white water-
lily. The peat is apparently derived from the seaward edge
of Lodmoor, which lies to the north-east of Weymouth.
Its age is not earlier than Neolithic, and at present it is
impossible to say whether it may not be entirely Post-
Roman.

Nymphaea alba. .
Prunus Padus.
Myriophyllum spicatum.
Sambucus nigra.
Alnus glutinosa.
Ceratophyllum demersum.
Sparganium ramosum (.?).

Potamogeton natans.

lucens.

perfoliatus.

Scirpus maritimus (?).

Carex.

Phragmites communis.

Osmunda regalis.

97

CHAPTER VI.

Former Distribution of British Plants.

I HAVE set down in this chapter what is known of the
past history of our British Plants ; but the species about
which we have as yet been able to learn anything amount
only to about one-sixth of the flora, though constant
additions are being made to the number. Under these
circumstances, and in face of the imperfection of the record
in Pliocene times, I doubt whether it would be of much use
to attempt any minute analysis of the list ; all that can be
done with advantage, is to draw attention to the leading
changes in geographical distribution that have already
been proved.

Variations caused by climatic changes were spoken of
in Chapter IV. In the course of time, however, there have
been other changes in distribution ; for it is obvious that a
flora driven south by a cold wave, on its return when the
climate has again become genial is not likely to consist
of exactly the same species. The chances of dispersal
cannot be twice alike. When the mammals and birds
change, the relative power of spreading possessed by the
different plants must change also; when England is con-
nected with the Continent, and the Rhine flows to Norfolk,
heavy seeds must have easier travelling than when Britain
becomes an island. Other differences in geographical dis-
tribution seem to be the result of accident — one plant has
accidentally been introduced and has had time to spread,

H

98 Origin of the British Flora.

a later comer needing the same station finds the ground
pre-occupied. All the proved cases will now be brought
together; but, as this chapter is merely a record of facts,
it does not seem advisable at present to deal with the
converse side of the question, that is to say with the
noticeable absence of many of our most common living
species. This deficiency, also, may be apparent only, not
real, and till we have a fuller knowledge of the fossil
plants it is undesirable to throw out suggestions which
to-morrow's work may show to be founded on nothing
more than the incompleteness of our search.

The exotic plants which have as yet been recorded as
British fossils are only six ; but I may repeat, in a more
general sense, the remarks already made, with regard to
the plants of one locality, and say that in reality the pro-
portion of exotic species must be considerably greater.
These are the plants for the determination of which it is
most difficult to obtain the necessary material. Botanists
seldom collect plants in fruit, and, if they do, the ordinary
method of preserving specimens is not suitable, as most of
the seeds that are ripe, or nearly ripe, fall out and are lost
in drying. My own collection of recent seeds and fruits
includes only a small proportion of exotic forms ; but I
have examined various fossil seeds which certainly do not
belong to any living British plant, and are quite determin-
able, if only sufficiently complete continental collections
were available.

Papaver somniferum has only been found at Cowden
Glen, and in face of the great uncertainty as to the age
of the peaty deposit at that place I do not feel prepared
to accept it as a true fossil, though the opium-poppy was
apparently grown in Switzerland in Neolithic times.

Acer monspessulanum occurs in Interglacial deposits at
Selsey, in Sussex, and at Stone, in Hampshire. It now

Former Distribution of British Plants. 99

flourishes throughout the Mediterranean region, and extends
into Central Europe. This maple grows well in gardens
in the South of England and seeds freely; though I have
not heard of any case in which it has spread from self-
sown seedlings. Mr. A. R. Wallace has undertaken some
experiments in Dorset with a view to ascertain whether
this plant can establish itself under natural conditions.

Trapa nutans is found in the Cromer Forest-bed, but
does not seem to have re-entered this country after it had
been driven out by the cold. It is an aquatic plant still
living as near as the South of Sweden, and has large
edible fruits known as water-chestnuts. Its absence in
Britain seems to be unconnected with changed climatic
conditions.

Salix polaris occurs abundantly in Glacial deposits,
both Early and Late ; but it has now completely dis-
appeared from Britain. It grows within the Arctic
Regions, and on the highest mountains of Scandinavia.

Picea excelsa was common in the East of England in
Preglacial times. It is apparently another large-seeded
plant that has been unable to re-establish itself here, now
that Britain is separated from the Continent. There is
nothing in the modern distribution of the spruce-fir to
suggest that it is unsuited for our present climate, though
this tree does not tend to spread from seedlings as do
Pinus sylvestris and Pinus maritima.

Najas graminea has only been found in the Interglacial
deposit at West Wittering in Sussex. Its recent distri-
bution is throughout the Tropics of the Old World, and
also in the Mediterranean Region. In Britain it has been
introduced at one spot, where it grows in a canal which
receives waste hot water from a mill.

Najas minor occurs in Preglacial deposits, and at
West Wittering. It also belongs to warmer climates.

lOO Origin of the British Flora.

ranging throughout Europe except in the north and in
Britain. It is living in the Rhine.

A certain number of our fossil-plants, though still living
in Britain, formerly had a range markedly different. The
majority of these species are northern forms, which formerly
occupied our lowlands, but on the passing away of the
cold of the Glacial Epoch could only live on our mountain
tops. They are Dry as octopetala, Arctostaphylos Uva-ursi,
Androineda Polifolia^ Loiseleiiria procumbens, Oxyria digyna,
Betula nana, Salix Myrsinites, Salix herbacea, Salix reti-
culata, Carex alpina. The Temperate species of which the
ancient distribution within Britain was markedly different
from that now existing were only three or four.

Quercus Robur appears at one time to have grown at
higher elevation ; for remains of well-grown oaks occur
occasionally in peat mosses above the limit of any but
stunted trees.

Pinus sylvestris seems to have been abundant through-
out Britain during part of the Neolithic Period, for its
cones are abundant at the base of peat-mosses and in
' submerged forests.' It afterwards disappeared from the
South of England and only recently has been re-introduced.

Potamogeton trichoides occurred in Sussex and Hamp-
shire in Interglacial times ; it is now confined in Britain to
Norfolk, Suffolk, and the West of Ireland.

Najas marina, now confined to a single locality in
Norfolk, was formerly widely distributed. It has now
been found fossil in Norfolk, Suffolk, Hertfordshire, and
Glamorgan.

Clematis Vitalba, L.
Interglacial (?) : —

Stoke Newington, London.

Mr. Worthington Smith has recorded leaves of this
plant from a Palaeolithic deposit at Stoke Newington.

Former Distribution of British Plants. loi

Thalictrum minus, L.
Late Glacial: —

Hailes, near Edinburgh, associated with Salix polaris

Early Glacial: —

Beeston, Norfolk (base of Arctic Freshwater bed).

Two small sharp-ribbed fruits have been found at
Hailes and are doubtfully referred to this species. Two
small fruits from Beeston, the one oval the other elongated,
sharp-ribbed and obscurely stalked, may also be referred to
T. mimis. In each case the fruits are considerably smaller
than my recent specimens.

Thalictrum flavum, L.
Roman Period : —

Silchester, Hampshire.
Interglacial: —

West Wittering, Sussex ; Southelmham, Suffolk.
Preglacial (Cromer Forest-bed): —

Beeston, Sidestrand, Mundesley, Ostend ( Norfolk ) ;
Gorton, Pakefield (Suffolk).

Also at Honerdingen in Hanover (Carl Weber) ; Klinge
bei Cottbus, Prussia, where it is associated with Brasenia^
Najas marina, &c. ; and in the Pine Zone (Neolithic ?) in
Sweden (Gunnar Andersson).

Ranunculus aquatilis, L.
Neolithic: —

Kelsey Hill, Yorkshire ; lacustrine deposits of the
Scottish Lowlands at Redhall, Hailes, Broughton, Elie,
and Cowden Glen ; Megaceros-marl of Ireland.

I02 Origin of the British Flora.

Late Glacial: —

Hoxne, Suffolk (Bed C); Kirk Michael, Isle of Man;
Ballaugh, Isle of Man; Hailes, near Edinburgh (lower bed);
Corstorphine, near Edinburgh; Dronachy, Fife; Gayfield,
near Edinburgh.

Interglacial : —

West Wittering, Sussex ; Endsleigh Street, London ;
Hitchin, Hertfordshire; Hoxne, Suffolk (bed D); South-
elmham, Suffolk ; Allenton, Derby ; Airdrie, Lanark ;
Kilmaurs, Ayrshire.

Early Glacial : —

Beeston, Norfolk (at base of the Glacial deposits).
Preglacial (Cromer Forest-bed): —

Beeston, Cromer, Sidestrand, Trimingham, Mundesley,
Ostend (Norfolk) ; Pakefield, Suffolk.

Several varieties occur fossil ; but the characters of the
fruit do not seem to be sufficiently constant in the recent
state to allow of any determination of sub-species from
fruit alone.

Ranunculus hederaceus, L.
Interglacial : —

West Wittering, Sussex.

Probably common elsewhere, but included among the
forms of R. aquatilis.

Ranunculus sceleratus, L.

Neolithic : —

Crossness, Essex (lower peat).

Late Glacial: —

Hoxne, Suffolk (bed C).

Former Distribution of British Plants, 103

Interglacial : —

Stone, Hampshire ; West Wittering, Sussex ; Endsleigh
Street, London ; Hitchin, Hertfordshire ; Hoxne, Suffolk
(Bed D); Southelmham, Suffolk ; AUenton, near Derby.

Preglacial (Cromer Forest bed) : —
Pakefield, Suffolk.

Ranunculus Flammula, L.
Roman Period : —

Silchester, Hampshire.
Neolithic : —

Ballaugh, Isle of Man (bed A) ; lacustrine deposits of
the Scottish Lowlands, Redhall, Hailes, Broughton, Cowden
Glen, Elie ; Fillyside, near Edinburgh (in raised beach).

Late Glacial: —

Ballaugh, Isle of Man (bed D).
Interglacial : —

Southelmham, Suffolk ; Allenton, near Derby.

Ranunculus Lingua, L.
Roman Period : —

Silchester, Hampshire.
Neolithic : —

Broughton, near Edinburgh ; Redhall, near Edinburgh ;
Hailes, near Edinburgh (one doubtful specimen).

Interglacial : —

West Wittering, Sussex; Hoxne, Suffolk (beds D and E).

Ranunculus acris, L.

I can find no trace of this species in deposits as old
as the Roman occupation.

I04 Origin of the British Flora,

Ranunculus repens, L.
Roman Period : —

Silchester, Hampshire.

Neolithic : —

Crossness, Essex (upper and lower peats); Fillyside,
near Edinburgh (raised beach) ; Lacustrine deposits of the
Scottish Lowlands, Redhall, Hailes, Broughton, Elie.

Late Glacial : —

Hoxne, Suffolk (bed C) ; Ballaugh, Isle of Man ; Close
y Garey, Isle of Man ; Hailes, near Edinburgh (lower bed);
Gayfield, near Edinburgh; Corstorphine, near Edinburgh;
Garvel Park (Clyde Beds).

Interglacial : —

Stone, Hampshire ; West Wittering, Sussex ; Grays,
Essex ; Endsleigh Street, London ; Shacklewell, London ;
Hitchin, Hertfordshire ; Hoxne (beds D and E) ; Allenton,
Derby.

Preglacial (Cromer Forest-bed) —

Beeston, West Runton, Sidestrand, Mundesley (Norfolk);
Pakefield (Suffolk).

Ranunculus bulbosus, L.
Interglacial : —

West Wittering, Sussex ; Allenton, near Derby (a
doubtful carpel).

Ranunculus Sardous, Crantz.
Roman Period : —

Silchester, Hampshire.

Interglacial: —

West Wittering, Sussex ; Allenton, near Derby.

Former Distribution of British Plants. 105

Ranunculus parviflorus, L.
Interglacial : —

West Wittering, Sussex.

Ranunculus Ficaria, L.
This has not yet been found in a fossil state; the
carpels, however, are softer than in most of the other species
of Ranunculus^ and are less likely to be preserved.

Caltha palustris, L. •
Roman Period : —

Silchester, Hampshire.
Neolithic : —

Redhall, near Edinburgh ; Hailes, near Edinburgh.

Late Glacial : —

Hoxne, Suffolk (bed C).

Interglacial : —

West Wittering, Sussex.

Preglacial (Cromer Forest-bed) : —
Mundesley, Norfolk.

NUPHAR LUTEUM, L.
Neolithic and Postglacial : —

Hampton Waterworks, Middlesex ; Casewick, Lincoln-
shire; Northampton (old river bed).

Interglacial : —

West Wittering, Sussex ; Mundesley, Norfolk (old river
deposit).

Preglacial (Cromer Forest-bed) : —

Beeston, West Runton, Cromer, Overstrand, Sidestrand,
Mundesley (Norfolk); Corton, Pakefield (Suffolk).

io6 Origin of the British Flora.

NYMPHiEA ALBA, L.
Recent Alluvium (?) : —

Happisburgh, Norfolk ; Weymouth.
Interglacial : —

West Wittering. Also at Griinenthal and Fahrenkrug,.
in Holstein, and Klinge bei Cottbus, Prussia, associated
with Brasenia^ &c. (C.Weber) ; at Honerdingen in Hanover
(C. Weber) ; and in Sweden in the Birch, Pine, Oak, and
Spruce Zones (Neolithic) (Gunnar Andersson).

The exact date of the Alluvium at Happisburgh and
at Lodmoor, near Weymouth, cannot be fixed.

Brasenia purpurea, Mich.

This species, though found in Africa, Asia, Australia,
and America, is unknown living in Europe.

It occurs in the fossil state in Russia, Denmark,
Germany, and Switzerland (Gunnar Andersson); but has
not yet been found in Britain.

PAPAVER SOMNIFERUM, L.

Neolithic (.?)—

Cowden Glen, Renfrewshire.

The Opium Poppy was cultivated in Neolithic times,,
and its seeds have been found in the Swiss Lake-dwellings.
The deposit at Cowden Glen is considered by Professor
James Geikie to be of Interglacial age ; but the occurrence
in it of Papaver somniferum suggests a more modern date.

Papaver Argemone, L.
Roman Period : —

Silchester, Hampshire.

Former Distribution of British Plants. 107

Chelidonium matus, L.
Interglacial: —

West Wittering, Sussex.

Five well-preserved and characteristic seeds have been
found.

FUMARIA OFFICINALIS, L.

Neolithic : —

Redhall, near Edinburgh.

This plant has only been found associated with flax and
weeds of cultivation.

Thlaspi arvense, L.
Roman Period : —

Silchester, Hampshire.

Cakile maritima, Scop.

Unknown fossil in Britain, but occurs in the Oak Zone
(Neolithic) in Gotland (Gunnar Andersson).

Viola palustris, L.
Neolithic : —

Drope, Glamorgan (?) ; Fillyside, near Edinburgh (raised
beach.)

Lacustrine deposits of the Scottish Lowlands, Hailes,
Redhall, near Edinburgh ; Elie, Fife (?).

Late Glacial: —

Hoxne, Suffolk (bed C) ; Kirk Michael, and Close y
Garey, Isle of Man ; Hailes (lower bed) ; Corstorphine,
Gayfield, near Edinburgh ; Dronachy, Fife.

Interglacial : —

AUenton, near Derby ; Faskine, Lanark.
Preglacial (Cromer Forest-bed) :—

Mundesley, Norfolk; Pakefield, Suffolk.

io8 Ofigin of the British Flora.

SiLENE MARITIMA, With.
Interglacial : —

West Wittering, Sussex.

Lychnis alba, Mill.
Neolithic : —

Elie, Fife.

Lychnis diurna, Sibth.
Neolithic : —

Lacustrine deposits of the Scottish Lowlands, Hailes,
Redhall, Elie(?); Fillyside, near Edinburgh (raised beach) (?).
Late Glacial: —

Hailes, near Edinburgh (lower bed).
Interglacial : —

West Wittering, Sussex.

Lychnis Flos-cuculi, L.
Roman Period : —

Silchester, Hampshire.

Neolithic: —

Redhall, near Edinburgh (capsules and seeds); Hailes,
near Edinburgh.

Late Glacial: —

Garvel Park (Clyde Beds).
Interglacial : —

West Wittering, Sussex.

Stellaria aquatica, Scopoli.
Interglacial : —

West Wittering, Sussex.

Preglacial (Cromer Forest-bed): —

Beeston, Norfolk.

Former Distribution of British Plants. 109

Stellaria media, Cyr.
Roman Period : —

Silchester, Hampshire.
Neolithic : —

Fillyside, near Edinburgh (raised beach) ; lacustrine
deposits of the Scottish Lowlands, Hailes, Redhall, Elie,
Broughton.

Late Glacial: —

Twickenham, Middlesex ; Hoxne, Suffolk (bed C) ;
Hailes, near Edinburgh (lower bed); Corstorphine, near
Edinburgh.

Interglacial : —

West Wittering, Sussex; Endsleigh Street, London.

Preglacial (Cromer Forest-bed): —
Overstrand, Norfolk.

Stellaria Holostea, L.
Interglacial : —

West Wittering, Sussex.

Stellaria graminea, L.
Roman Period : —

Silchester, Hampshire.

Stellaria uliginosa, L.
Neolithic : —

Redhall, near Edinburgh ; Hailes, near Edinburgh.

Arenaria trinervia, L.

Not yet found fossil in Britain ; but recorded by
Keilhack and Nathorst from Lauenburg an der Elbe, where
it is associated with Brasenia purpurea.

no Origin of the British Flora,

Arenaria peploides, L.
Interglacial : —

Stone, Hampshire.

Spergula arvensis, L.
Neolithic : —

Redhall, near Edinburgh.

Montia fontana, L.

Neolithic : —

Fillyside, near Edinburgh (raised beach) ; lacustrine
deposits of the Scottish Lowlands, Redhall, Hailes
Broughton, near Edinburgh; Cowden Glen, Renfrewshire.

Late Glacial : —

Twickenham, Middlesex; Hoxne, Suffolk (bed C).

Interglacial : —

Hitchin, Hertfordshire; Hoxne, Suffolk (bed D); Allen-
ton, near Derby; West Wittering, Sussex.

Hypericum perforatum, L.
Roman Period : —

Silchester, Hampshire.

Interglacial : —

West Wittering, Sussex.

Hypericum quadrangulum, L.
Neolithic (?) —

Redhall, near Edinburgh.

Interglacial: —

West Wittering, Sussex.

Preglacial (Cromer Forest-bed) : —
Pakefield. Suffolk.

Former Distribution of British Plants. 1 1 1

Hypericum elodes, L.

Neolithic (?) —

Redhall, near Edinburgh.

TiLIA PLATYPHYLLOS, Scop.

Unknown fossil in Britain.

Recorded from Griinenthal and Fahrenkrug, in Holstein
(associated with Brasenia, 8ic.) (C. Weber) ; Lauenburg an
der Elbe (with Brasema, &c.) (Keilhack and Nathorst);
Klinge bei Cottbus, Prussia (with Brasema, Najas marina,
&c.) (C.Weber); Honerdingen, in Hanover, associated with
Tilia parvifolia and T. intermedia (C. Weber).

Tilia europea is recorded from the Pine and Oak Zones
in South Sweden (Gunnar Andersson).

LiNUM, sp.

Roman Period : —

Silchester, Hampshire.

Neolithic: —

Lacustrine deposits of the Scottish Lowlands, Redhall,
Hailes, Elie.

Capsules and seeds of flax are so common at Redhall
as to suggest that bundles of the plant were steeped there.
Flax is known to have been cultivated in Neolithic times.

Geranium columbinum, L.

No species of Geraniicm has yet been found fossil in
Britain, with the doubtful exception of a seed from Ends-
leigh St., London.

G. columbinium is recorded by Keilhack from Lauen-
burg an der Elbe (associated with Brasenid).

112 Origin of the British Flora,

OXALIS ACETOSELLA, L.
Neolithic : —

Crossness, Essex (upper bed) ; Redhall, near Edinburgh ;
Hailes, near Edinburgh.

Late Glacial : —

Hailes, near Edinburgh (lower bed).

Ilex Aquifolium, L.
Neolithic: —

Crossness, Essex (upper and lower peats).

Interglacial : —

West Wittering, Sussex.

Also recorded from Griinenthal, in Holstein ; Klinge
bei Cottbus, in Prussia (associated with Brasenia^ Najas
marina, 8ic,) ; Honerdingen, in Hanover (C. Weber).

Rhamnus Frangula, L.
Interglacial : —

Hoxne (bed D) ; West Wittering, Sussex.

Recorded from Honerdingen, in Hanover (C. Weber) ;
Lauenburg an der Elbe (A. G. Nathorst) ; and from the
Pine and Oak Zones in Sweden (Gunnar Andersson).

VlTIS VINIFERA, L.

A portion of a stem discovered by Mr. Worthington
Smith in the Palaeolithic deposits of Stoke Newington was
so determined by Mr. Carruthers. The specimen has
unfortunately been mislaid, but Mr. Smith thinks that it is
probably a recent stem accidentally introduced ; he has
found several such in ground disturbed during the Roman
occupation.

Former Distribution of British Plants. 1 1 3

Acer campestre, L.
Neolithic:—

Crossness, Essex (upper and lower peats).

Preglacial (Cromer Forest-bed) — :

Pakefield, Suffolk.

Also recorded from the Oak Zone in Sweden (Gunnar
Andersson) ; and from Fahrenkrug, in Holstein, and
Klinge bei Cottbus, in Prussia (associated with Brasenia)
(C. Weber).

Acer monspessulanum, L.
Interglacial : —

Stone, Hampshire ; Selsey, Sussex.

This maple lives throughout the Mediterranean region
and extends into central Europe ; it grows well in gardens
in the South of England and seeds freely.

Acer platanoides, L.

Recorded by Gunnar Andersson from the Oak Zone in
South Sweden ; and by Keilhack from Lauenburg an der
Elbe, associated with Brasenia. It has not been found in
Britain.

PRUNUS SPINOSA, L.
Neolithic: —

Crossness, Essex (upper and lower peats) ; North-
ampton ; Hailes, near Edinburgh.

Interglacial : —

Hitchin, Hertfordshire ; West Wittering, Sussex.

Preglacial (Cromer Forest-bed): —

West Runton, Overstrand, Happisburgh (Norfolk),
Pakefield, Suffolk.

I

114 Origin of the British Flora,

Prunus domestica, L.
Neolithic : —

Crossness, Essex (upper peat).

In • Etudes d'Ethnographie Prehistorique. — II. Les
Plantes cultiv^es de la Periode de Transition au Mas-d'Azil,'
{Anthropologic, Vol. VII., No. I., pp. 1-24), Monsieur Ed.
Piette has given a good account and figures of the early
cultivated species of Prunus. We have not yet obtained
in Britain sufficient material for a similar analysis.

Prunus Avium, L.
Neolithic : —

Crossness, Essex (upper and lower peats) ; Gayfield,
Edinburgh.

Interglacial: —

West Wittering, Sussex ; Selsey, Sussex.

Prunus Padus, L. •
Neolithic : —

Northampton ; Hornsea, Yorkshire ; Sand le Meer,
Yorkshire ; Hailes, near Edinburgh.

Interglacial : —

Selsey, Sussex ; Airdrie, Lanarkshire.
Recorded from the Pine and Oak Zones (Neolithic) in
Sweden (Gunnar Andersson).

Spiraea Ulmaria, L.
Roman Period : —

Silchester, Hampshire.

Neolithic : —

Hailes, near Edinburgh ; Redhall, near Edinburgh.

Former Distribution of British Plants. 115

Interglacial : —

West Wittering, Sussex.

Recorded also from the Pine and Spruce Zones
(Neolithic) in Sweden (Gunnar Andersson).

Preglacial (Cromer Forest-bed) : —
Pakefield, Suffolk.

RUBUS Id^US, L.
Neolithic : —

Southampton Docks (?) ; Lacustrine deposits of the
Scottish Lowlands at Hailes, Redhall, Elie, Cowden Glen ;
Fillyside, near Edinburgh, (raised beach).

Late Glacial : —

Hoxne, Suffolk (bed C) ; Gayfield, near Edinburgh ;
Garvel Park (Clyde Beds) ; Black Burn, East Tarbet
(Clyde Beds).

Interglacial : —

Shacklewell, London ; Hoxne, Suffolk (beds D and E) ;
Allenton, near Derby.

Early Glacial: —

Beeston, Norfolk (at the base of the Arctic plant-bed).

RUBUS FRUTICOSUS, L.
Roman Period : —

Silchester, Hampshire.

Neolithic : —

Crossness, Essex (upper and lower peats).

Hailes, near Edinburgh ; Redhall, near Edinburgh.

ii6 Origin of the British Flora,

Interglacial : —

Stone, Hants ; West Wittering, Sussex ; Selsey,
Sussex ; Grays, Essex.

Preglacial (Cromer Forest-bed) : —

Overstrand ; Mundesley ; Pakefield.

RUBUS C^SIUS, L.

Unknown fossil in Britain.

Recorded with doubt by Gunnar Andersson from the
Oak Zone in South Sweden.

RUBUS SAXATILIS, L.
Unknown fossil in Britain.

Recorded by Gunnar Andersson from the Pine, Oak,
and Spruce Zones in Sweden.

Dryas octopetala, L.
Late Glacial : —

Corstorphine, near Edinburgh ; Crianlarich, Perthshire.
This species, though rare fossil in Britain, is very
abundant in similar deposits in Scandinavia.

POTENTILLA TORMENTILLA, Neck.

Roman Period : —

Silchester, Hampshire.

Neolithic : —

Ballaugh, Isle of Man (bed A) ; Close y Garey, Isle of
Man (bed B); Hailes, near Edinburgh; Redhall, near
Edinburgh; Elie, Fife.

Former Distribution of British Plants. 1 1 7

Late Glacial: —

Roxburgh Street, Greenock (?).

Interglacial : —

Endsleigh Street, London; West Wittering, Sussex.

POTENTILLA COMARUM, Nestl.
Neolithic :—

Redhill, near Edinburgh; Hailes, near Edinburgh.
Late Glacial : —

Kirk Michael, Isle of Man; Close y Garey, Isle of
Man.

Interglacial : —

Airdrie, Lanarkshire.

Alchemilla arvensis, Lam.
Roman Period : —

Silchester, Hampshire.

Neolithic: —

Redhall, near Edinburgh.

Interglacial : —

West Wittering, Sussex.

As only a single achene has been sent to me from
Redhall, this may possibly be a recent specimen acci-
dentally introduced. At West Wittering achenes are
fairly common.

POTERIUM OFFICINALE, Hook.
Neolithic : —

Redhall, near Edinburgh; Cowden Glen, Renfrewshire.

1 1 8 Origin of the British Flora.

Late Glacial: —

Hoxne, Suffolk (bed Q; Ballaugh, Isle of Man (bed C);
Corstorphine, near Edinburgh.

Interglacial : —

West Wittering, Sussex; Hitchin, Hertfordshire; South-
elmham, Suffolk.

Early Glacial: —

Beeston, Norfolk (at base of the Arctic Freshwater bed).
Preglacial (Cromer Forest-bed) : —

Beeston, Sidestrand, Mundesley, in Norfolk.

Rosa.
Neolithic: —

Crossness, Essex (upper and lower peats) ; Redhall,
near Edinburgh (?).

Interglacial : —

West Wittering, Sussex ; Selsey, Sussex; Stone, Hamp-
shire; Grays, Essex; Shacklewell, London; Hoxne, Suffolk
(bed D).

Prickles and achenes are not uncommon at several
localities. The achenes are always short in proportion to
their breadth, and very small; the prickles are generally
curved and small. I have seen nothing approaching to
the common living forms of R, canina^ L. ; and the fossils
more suggest a species with small globose fruits.

Pyrus torminalis, Ehrh.
Interglacial : —

Hitchin, Hertfordshire; Hoxne, Suffolk (bed D).

Former Distribution of British Plants. 1 1 9
Pyrus Aria, Sm.

Preglacial (Cromer Forest-bed) : —
Pakefield, Suffolk (leaves).

Pyrus Aucuparia, Gaert.
Neolithic : —

Caerwys, Flintshire.

Recorded from the Pine, Oak, and Spruce Zones

(Neolithic) in Sweden (Gunnar Andersson).

Pyrus communis, L.
Neolithic : —

Crossness, Essex (upper and lower peats), (wood de-
termined by Professor Marshall Ward).

Crataegus Oxyacantha, L.

Neolithic : —

Barry Docks, Glamorgan; Crossness, Essex (upper
and lower peats); Redhall, near Edinburgh; Hailes, near
Edinburgh ; Gayfield near Edinburgh.

Interglacial: —

Southelmham, Suffolk.

Preglacial (Cromer Forest-bed) : —

Overstrand, Norfolk.

In the Swedish peat- mosses only the form Cratcegus
monogyna occurs (Gunnar Andersson).

Saxifraga oppositifolia, L.

Unknown fossil in Britain.

Recorded from Deuben, in Saxony (Nathorst), asso-
ciated with Salix kerbacea, &c.

I20 Origin of the British Flora.

Saxifraga Hirculus, L.
Unknown fossil in Britain.
Recorded from Deuben, in Saxony (Nathorst).

Saxifraga aizoides, L.

Unknown fossil in Britain,

Recorded from Deuben, in Saxony (Nathorst).

HiPPURIS VULGARIS, L.
Neolithic : —

Drope, Glamorgan ; Crossness, Essex (upper peat) ;
Hailes, near Edinburgh ; Cowden Glen, Renfrewshire.

Late Glacial : —

Hoxne, Suffolk (bed C) ; Kirk Michael, Isle of Man ;
Corstorphine, near Edinburgh ; Hailes, near Edinburgh
(lower bed); Dronachy, Fife.

Interglacial ; —

West Wittering, Sussex ; Endsleigh Street, London ;
Hitchin, Hertfordshire ; Hoxne, Suffolk (bed E) ; South-
elmham, Suffolk ; Faskine, Lanark ; Airdrie, Lanark ;
Kilmaurs, Ayrshire.

Early Glacial : —

Beeston, Norfolk (at base of Arctic Freshwater bed) ;
Ostend, Norfolk.

Preglacial (Cromer Forest-bed) : —

Beeston, Cromer, Sidestrand, Mundesley, Ostend (Nor-
folk).

Former Distribution of British Plants. 1 2 1

Myriophyllum spicatum, L. .
Neolithic : —

Lacustrine deposits of the Scottish Lowlands, Gayfield,
Cowden Glen ; Megaceros-marls of Ireland.

Late Glacial: —

Hoxne, Suffolk (bed C); Corstorphine ; Dronachy, Fife;
Close y Garey, Isle of Man (bed C).

Interglacial : —

West Wittering, Sussex ; Endsleigh Street, London ;
Hitchin, Hertfordshire (?) ; Southelmham, Suffolk ; Kil-
maurs, Ayrshire.

Early Glacial : —

Beeston, Norfolk (3 doubtful fruits).

Preglacial (Cromer Forest-bed) : —

Cromer, Overstrand, Sidestrand, Mundesley, in Norfolk ;
Pakefield, in Suffolk.

Myriophyllum alternifolium, L.

Unknown fossil in Britain.

Recorded from the Pine Zone (Neolithic) in Sweden
{Gunnar Andersson).

Trapa natans, L.

Preglacial (Cromer Forest-bed) : —

Sidestrand, Mundesley, Ostend (Norfolk) ; Pakefield, in
Suffolk.

This plant does not occur in Britain in deposits later
than the Cromer Forest-bed. In Scandinavia, however, it
is common in Postglacial peaty deposits, extending also

122 Origin of the British Flora.

into Finland, At Lauenburg an der Elbe it occurs asso-
ciated with Brasenia (Keilhack) ; and at Griinenthal, in
Holstein, under similar conditions (Weber) ; both these
deposits being apparently of Interglacial date. As a living
species it still lingers in Southern Sweden, and is more
common in Southern Europe, where the fruits are eaten.

Hydrocotyle vulgaris, L.
Neolithic (?)—

Ballaugh, Isle of Man (bed A) ; Elie, Fife.
Interglacial : —

West Wittering, Sussex ; Southelmham, Suffolk ;.
Allenton, near Derby.

Preglacial (Cromer Forest-bed) : —
Gorton, Suffolk ; Pakefield, Suffolk.

Apium graveolens, L.
Interglacial : —

West Wittering, Sussex.

Apium nodiflorum, Reich.
Interglacial:—

Airdrie, Lanark ; West Wittering, Sussex.

CiCUTA VIROSA, L.

Unknown fossil in Britain.

Recorded from the Pine, Oak, and Spruce Zones
(Neolithic) in Scandinavia (Gunnar Andersson).

Carum Carui, L.

This plant is only represented by a single well-preserved
fruit from Redhall. In the absence of corroborative

Former Distribution of British Plants. 123

evidence, and considering how largely carraway-seeds are
used for cakes, it does not seem advisable to Include this
species in the list of British fossil plants.

SlUM LATIFOLIUM, L.

Unknown fossil in Britain.

Recorded from the Pine and Oak Zones (Neolithic) in
Sweden (Gunnar Andersson).

CONOPODIUM DENUDATUM, Koch.
Roman Period : —

Silchester, Hampshire.

Ch^erophyllum temulum, L.
Interglacial : —

West Wittering, Sussex.

Anthkiscus sylvestris, Hoffm.
Interglacial : —

West Wittering, Sussex.

OEnanthe fistulosa, L.
Interglacial: —

West Wittering, Sussex.

GEnanthe Lachenalii, Gmel.
Late Glacial: —

Garvel Park (Clyde Beds).

Preglacial (Cromer Forest-bed) : —

Mundesley, Norfolk ; Pakefield, Suffolk.

124 Origin of the British Flora.

GEnanthe Phellandrium, Lam.
Neolithic: —

Crossness, Essex (upper peat).

Late Glacial: —

Hoxne, Suffolk (bed C).

Interglacial : —

West Wittering, Sussex; Hoxne, Suffolk (bed D);
Southelmham, Suffolk.

Preglacial (Cromer Forest-bed): —

Sidestrand, Mundesley, Beeston, in Norfolk ; Pakefield,
in Suffolk.

^THUSA CYNAPIUM, L.
Roman Period : —

Silchester, Hampshire.

Neolithic: —

Hailes, near Edinburgh; Elie, Fife.

In each case there is some doubt as to the age of the
deposit.

Angelica sylvestris, L.
Interglacial: —

West Wittering, Sussex.

Also recorded from the Oak Zone (Neolithic) Gotland
(Gunnar Andersson).

Peucedanum palustre, Mcench.
Unknown Fossil in Britain.

Recorded from the Oak Zone (Neolithic) in Gotland
(Gunnar Andersson).

Former Distribution of British Plants. 125

Heracleum Sphondylium, L.
Late Glacial : —

Twickenham, Middlesex.

Preglacial (Cromer Forest-bed): —

Pakefield, Suffolk.

Caucalis nodosa, Scop.
Interglacial: —

Stone, Hampshire.

Hedera Helix, L.
Neolithic: —

Crossness, Essex (upper and lower peats) (Spurrell);
Caerwys, Flintshire.

Interglacial: —

Grays, Essex.

Recorded also from the Oak Zone (Neolithic) in
Gotland (Gunnar Andersson).

CORNUS SUECICA, L.
Unknown fossil in Britain.

Recorded from Gotland, in South Sweden, associated
with Arctic plants.

CORNUS SANGUINEA, L.
Neolithic : —

Barry Docks, South Wales; Crossness, Essex (upper

and lower peats); Albert Dock, Essex.

Interglacial : —

West Wittering, Sussex; Hitchin, Hertfordshire; Hoxne,
Suffolk (a single fruit, not found in place and perhaps
recent).

126 Origin of the British Flora.

Preglacial (Cromer Forest-bed) : —

Happisburgh, Norfolk ; Pakefield, Suffolk.

Sambucus nigra, L
Roman Period : —

Silchester, Hampshire.

Neolithic : —

Southampton Docks (peat below sea-level) ; Crossness,
Essex (upper and lower peats); Northampton (old river-
bed) ; Hailes, near Edinburgh ; Redhall, near Edinburgh ;
Fillyside, near Edinburgh (in raised beach).

Late Glacial : —

Hoxne, Suffolk (bed C — perhaps derivative from bed D).

Interglacial : —

West Wittering, Sussex ; Hitchin, Hertfordshire ;
Hoxne, Suffolk (bed D).

Viburnum Opulus, L.
Neolithic : —

Crossness, Essex (upper and lower peats).

Interglacial : —

West Wittering, Sussex.

Viburnum Lantana, Linn.
Interglacial : —

West Wittering, Sussex.

Galium boreale, L.
Early Glacial: —

Beeston, Norfolk (one badly preserved fruit of Galium
perhaps belongs to this species).

Former DistribiUion of British Plants, 127

Galium palustre, L.
Neolithic: —

Cowden Glen, Renfrewshire.

Recorded by C. Weber with doubt from Klinge bei

Cottbus, Prussia, associated with Brasenia.

Galium uliginosum, L.

Unknown fossil in Britain.

Recorded from an Interglacial deposit at Griinenthal,
in Holstein (Weber).

Galium Aparine, L.

Neolithic (.?)—

Casewick, Lincolnshire.

Valeriana officinalis, L.
Neolithic: —

Barry Docks, South Wales; lacustrine deposits of the
Scottish Lowlands, Hailes, Redhall, Elie.

Interglacial: —

Allenton, near Derby ; Stone, Hampshire.

Valerianella olitoria, Moench.
Interglacial: —

West Wittering, Sussex.

SCABIOSA SUCCISA, L.
Interglacial: —

West Wittering, Sussex.

EUPATORIUM CANNABINUM, L.
Neolithic: —

Tilbury Docks (peat below sea-level) ; Redhall, near

Edinburgh.

128 Origin of the British Flora.

Late Glacial: —

Hoxne, Suffolk (bed C).
Interglacial: —

West Wittering, Sussex ; Shacklewell, London ; Hitchin,
Hertfordshire ; Hoxne, Suffolk (bed D) ; AUenton, near
Derby.

Aster Tripolium, L.
Interglacial: —

West Wittering, Sussex.

BiDENS CERNUA, L.
Neolithic: —

Redhall, near Edinburgh.

The fruit from Hoxne, referred in 1888 to this species,

is a variety of B. tripartita with four equal awns.

BiDENS TRIPARTITA, L.

Late Glacial: —

Hoxne, Suffolk (bed C — a starved fruit).

Interglacial : —

Hoxne, Suffolk (bed D), associated with a variety
having four equal awns.

Preglacial (Cromer Forest-bed): —

Mundesley, Norfolk; Pakefield, Suffolk.

Chrysanthemum segetum, L.
Neolithic: —

Hailes, near Edinburgh ; Redhall, near Edinburgh.

Chrysanthemum Leucanthemum, L.
Roman Period : —

Silchester, Hampshire.

A single fruit of this species, sent to me as from the

Former Distribution of British Plants. 129

Arctic bed at Hailes, near Edinburgh, is probably recent;
it is not carbonised, and the ribs are light-brown.

Matricaria inodora, L.
Neolithic: —

Redhall, near Edinburgh ; Hailes, near Edinburgh.

Only found associated with weeds of cultivation.

Tanacetum Vulgare, L.
Early Glacial: —

Beeston, Norfolk (base of the Arctic Freshwater bed).

TUSSILAGO Farfara, L.
Neolithic: —

Redhall, near Edinburgh (a single fruit).

Senecio sylvaticus, L.
Neolithic: —

Redhall, near Edinburgh.

Senecio aquaticus, Huds.
Interglacial: —

West Wittering, Sussex.

Carduus crispus, L.
Neolithic: —

Close y Garey, Isle of Man (bed B).

Cnicus lanceolatus, L.
Neolithic: —

Redhall, near Edinburgh (fruits rather small) ; Elie, Fife.
Interglacial: —

West Wittering, Sussex.

K

130 Origin of the British Flora.

Cnicus palustris, L.
Neolithic: —

Hailes, near Edinburgh ; Redhall, near Edinburgh.

Interglacial: —

Southelmham, Suffolk (fruits small) ; West Wittering,

Sussex.

Centaurea Cyanus, L.
Neolithic: —

Redhall, near Edinburgh.

With weeds of cultivation and flax-seeds.

Lapsana communis, L.
Neolithic: —

Redhall, near Edinburgh; Hailes, near Edinburgh (fruit
very small).

Interglacial : —

West Wittering, Sussex.

Preglacial Cromer Forest-bed): —
Pakefield, Suffolk.

PiCRIS HIERACIOIDES, L.

Preglacial (Cromer Forest-bed) : —
Pakefield, Suffolk.

Crepis virens, L.
Neolithic : —

Redhall, near Edinburgh.

Hieracium Pilosella, L.
Interglacial : —

West Wittering, Sussex.

No trace of any other species of Hieracium has yet
been found fossil in Britain.

Former Distribution of British Plants, 131

Leontodon autumnalis, L.
Neolithic :—

Redhall, near Edinburgh.

Interglacial : —

Allenton, near Derby.

Taraxacum officinale, Web.
Neolithic : —

Redhall, near Edinburgh ; Fillyside, near Edinburgh
(raised beach).

Late Glacial : —

Hoxne, Suffolk (bed C); Hailes, near Edinburgh
(lower bed) ; Corstorphine, near Edinburgh ; Garvel Park
(Clyde Beds) ; Roxburgh Street, Greenock (in glacial clay).

Interglacial : —

Southelmham, Suffolk ; Allenton, near Derby ; West
Wittering, Sussex.

SONCHUS ARVENSIS, L.
Neolithic : —

Lacustrine deposits of the Scottish Lowlands, Redhall,

and Gayfield, near Edinburgh.

Vaccinium Oxycoccos, L.

Unknown fossil in Britain.

Recorded from Fahrenkrug in Holstein (C. Weber) ;
Lauenburg an der Elbe (Keilhack); Klinge bel Cottbus,
in Prussia (C. Weber) ; at all three localities being asso-
ciated with Brasenia purpurea. Also from the Pine, Oak,
and Spruce Zones in Sweden (Gunnar Andersson).

132 Origin of the British Flora.

Vaccinium ViTis-lDiEA, Linn.

Unknown fossil in Britain.

Recorded from the Pine, Oak, and Spruce Zones in
Sweden (Gunnar Andersson) ; also in deposits apparently
of Late Glacial date at Griinenthal, in Holstein (Weber).

Vaccinium uliginosum, L.

Unknown fossil in Britain.

Recorded from Fahrenkrug in Holstein (associated with
Brasenia) (C. Weber) ; and in the Dryas, Birch, Pine, Oak,
and Spruce Zones in Sweden (Gunnar Andersson).

Vaccinium Myrtillus, L.

Twigs are recorded from Cowden Glen by Dr. Craig.
I can find no recognisable seeds or leaves of this species
in the material sent me by Mr. Bennie, and doubt whether
twigs alone would be sufficient for determination.

Recorded from a deposit apparently of Late Glacial
date at Griinenthal, in Holstein (C. Weber).

Arctostaphylos alpina, Spreng.

Unknown fossil in Britain.

Recorded from the Dryas Zone in Gotland (Gunnar
Andersson).

Arctostaphylos Uva-ursi, Spreng.

Late Glacial (.?)—

^ . Bovey Tracey, Devon (A. G. Nathorst).

•' Also recorded from the Dryas, Birch, Pine, and Oak
Zones in Gotland, Sweden (Gunnar Andersson).

Former Distribution of British Plants. 133

Andromeda Polifolia, L.

Late Glacial: —

Hailes, near Edinburgh (lower bed); Corstorphine, near
Edinburgh.

LOISELEURIA PROCUMBENS, Desv.
Late Glacial : —

Corstorphine, near Edinburgh.

Glaux maritima, L.
Interglacial : —

West Wittering, Sussex.

Fraxinus excelsior, L.
Neolithic : —

Crossness, Essex (upper and lower peats), wood de-
termined by Prof. Marshall Ward.

Interglacial : —

Hitchin, Hertfordshire.

In Sweden confined to the Oak Zone (Gunnar Anders-
son) ; recorded from Honerdingen in Hanover, associated
with Platanus, Juglans, and Najas flexilis (C. Weber).

Menyanthes trifoliata, L. .
Neolithic : —

Drope, Glamorgan ; Redhall, near Edinburgh ; Gay-
field, near Edinburgh ; Hailes, near Edinburgh; Elie, Fife;
Cowden Glen, Renfrewshire ; Montrose (in peat below 20
feet of estuarine deposits).

134 Origin of the British Flora.

Late Glacial : —

Hoxne, Suffolk (bed C) ; Kirk Michael, Isle of Man ;
Hailes, near Edinburgh (lower bed); Gayfield and Cor-
storphine, near Edinburgh ; Dronachy, Fife.

Interglacial : —

West Wittering, Sussex; Hitchin, Hertfordshire; South-
elmham, Suffolk ; Airdrie, Lanark.

Early Glacial : —

Beeston, Norfolk (base of Arctic Freshwater bed).

Preglacial (Cromer Forest-bed) : —

Cromer, Mundesley, Happisburgh (in Norfolk).

Myosotis sylvatica, Hoffm.

Recorded from Gotland, Sweden, by Dr. Gunnar
Andersson. No species of Myosotis has yet been found
fossil in Britain, M, lingulata being a wrong determination.

SOLANUM Dulcamara, L
Interglacial : —

West Wittering, Sussex.

Recorded from the Pine and Oak Zones in South
Sweden (Gunnar Andersson).

Preglacial (Cromer Forest-bed) : —
Corton, Suffolk.

Verbascum Thaspus, L.
Interglacial : —

West Wittering, Sussex.

Former Distribution of British Plants. 135

Bartsia Odontites, Huds.
Late Glacial : —

Garvel Park (Clyde Beds), four seeds received from Mr.
Thos. Scott.

Pedicularis palustris, L,
Neolithic : —

Hailes, near Edinburgh ; Redhall, near Edinburgh ;
Cowden Glen, Renfrewshire.

Mentha aquatica, L.
Roman Period : —

Silchester, Hampshire.

Interglacial : —

Hoxne, Suffolk (bed D) ; West Wittering, Sussex ;
Stone, Hampshire.

Preglacial (Cromer Forest-bed) : —
Pakefield, Suffolk.

This plant may be common at other localities, tlie
small size of its nutlets having caused it to be overlooked
till specially searched for.

LYCOPUS EUROPyEUS, L.
Neolithic : —

Redhall, near Edinburgh ; Hailes, near Edinburgh.

Late Glacial : —

Hoxne, Suffolk (bed C).

Interglacial : —

West Wittering, Sussex ; Stone, Hampshire ; Shackle-
well, London ; Hitchin, Hertfordshire ; Hoxne, Suffolk
(bed D) ; Southelmham, Suffolk.

1 36 Origin of the British Flora.

Preglacial (Cromer Forest-bed) : —

Mundesley, Norfolk ; Pakefield, Suffolk.

Thymus Serpyllum, L.
Late Glacial : —

Greenock (Roxburgh Street), in Clyde Beds.

Prunella vulgaris, L.
Roman Period : —

Silchester, Hampshire.
Neolithic : —

Redhall, near Edinburgh.

Stachys palustris, L.
Neolithic : —

Redhall, near Edinburgh ; Hailes, near Edinburgh ;
Fillyside, near Edinburgh (raised beach).

Late Glacial : —

Hailes, near Edinburgh (lower bed).
Interglacial : —

Hoxne Suffolk (bed D), one badly preserved nutlet
perhaps belongs to this species ; West Wittering, Sussex.
Preglacial (Cromer Forest-bed) : —

Beeston, Norfolk.

Stachys sylvatica, L.

Unknown fossil in Britain.

Recorded from the Dryas and Pine Zones (Late
Glacial and Neolithic) in Sweden (Gunnar Andersson).

Stachys arvensis, L.
Roman Period : —

Silchester, Hampshire.

Former Distribution of British Plants, 137

Galeopsis Tetrahit, L.
Roman Period : —

Silchester, Hampshire.
Neolithic : —

Hailes, near Edinburgh ; Redhall, near Edinburgh ;
Cowden Glen, Renfrewshire. —

Late Glacial : —

Twickenham, Middlesex.

The occurrence of Galeopsis Tetrahit^ often considered
to be a weed of cultivation, at Twickenham associated with
Reindeer, Bison, and Bos longifroiis, but not with extinct
mammals, suggests a transition period between Palaeolithic
and Neolithic.

AjUGA REPTANS, L.
Neolithic : —

Redhall, near Edinburgh ; Hailes, near Edinburgh ;
Fillyside, near Edinburgh (raised beach) ; Elie, Fife ;
Cowden Glen, Renfrewshire.

Late Glacial : —

Hoxne, Suffolk (bed C) ; Hailes, near Edinburgh
(lower bed).

Interglacial : —

Selsey, Sussex ; West Wittering, Sussex ; Hitchin,
Hertfordshire ; Allen ton, near Derby.

LiTTORELLA LACUSTRIS, L.

Neolithic : —

Redhall, near Edinburgh ; Megaceros-marls of Central
Ireland.

Late Glacial : —

Ballaugh, Isle of Man.

13^ Origin of the British Flora.

Atriplex patula, L.
Neolithic : —

Casewick, Lincolnshire ; Hailes, near Edinburgh ;
Redhall, near Edinburgh ; Fillyside, near Edinburgh
(raised beach) ; Elie, Fife.

Late Glacial :•»

Garvel Park (Clyde Beds) ; Roxburgh Street, Greenock
(Clyde Beds).

Interglacial : —

Stone, Hampshire ; West Wittering, Sussex ; Selsey,
Sussex.

Preglacial (Cromer Forest-bed) : —

Beeston, West Runton, Sidestrand (in Norfolk) ; Corton,
Pakefield (in Suffolk).

Polygonum aviculare, L.
Roman Period : —

Silchester, Hampshire.

Neolithic : —

Redhall, near Edinburgh ; Broughton, near Edinburgh.

Late Glacial : —

Hailes, near Edinburgh (lower bed).

Interglacial : —

Endsleigh Street, London ; Stone, Hampshire ; Selsey,
Sussex.

Polygonum Hydropiper, L.
Neolithic : —

Crossness, Essex (lower peat).

Former Distribution of British Plants. 139

Polygonum Persicaria, L.
Neolithic : —

Lacustrine deposits of the Scottish Lowlands, Hailes,
Redhall, Gayfield, Broughton, EHe.

Late Glacial : —

Twickenham, Middlesex.

Interglacial : —

West Wittering, Sussex ; Endsleigh Street, London.

Preglacial (Cromer Forest-bed): —
Pakefield, Suffolk.

Polygonum lapathifolium, L.
Neolithic (?) :—

Cowden Glen, Renfrewshire.

Polygonum amphibium, L.
Unknown fossil in Britain.
Recorded by Dr. Gunnar Andersson from South Sweden.

Polygonum viviparum, L.
Unknown fossil in Britain. 1

Recorded from the Dryas Zone of Gotland (Gunnar

Andersson) ; and from Deuben, in Saxony (A. G.

Nathorst).

OXYRIA digyna, Hill.
Late Glacial : —

Corstorphine, near Edinburgh.

RUMEX CONGLOMERATUS, Murr.
Roman Period : —

Silchester, Hampshire.

140 Origin of the British Flora,

Interglacial : —

West Wittering, Sussex.

RUMEX MARITIMUS, L.
Late Glacial : —

Hoxne, Suffolk (bed C).

Interglacial : —

Hoxne, Suffolk (beds D and E) ; Southelmham,
Suffolk.

Early Glacial : —

Beeston, Norfolk (base of Arctic Freshwater bed).

Preglacial (Cromer Forest-bed) : —

Beeston, West Runton, Overstrand, Sidestrand, Ostend
(in Norfolk) ; Pakefield (in Suffolk).

RUMEX OBTUSIFOLIUS, L.
Neolithic : —

Redhall, near Edinburgh.

Late Glacial : —

Close y Garey, Isle of Man (in Megaceros-marl).

Interglacial : —

West Wittering, Sussex ; Endsleigh Street, London.

RUMEX CRISPUS, L.
Neolithic : —

Barry Docks, Glamorgan ; Casewick, Lincolnshire ;

Redhall, near Edinburgh ; Hailes, near Edinburgh ;

Cowden Glen, Renfrewshire.

Late Glacial : —

Twickenham, Middlesex ; Hoxne, Suffolk (bed C) (i*) ;
Garvel Park (Clyde Beds).

Former Distribution of British Plants. 141

Interglacial : —

Hoxne, Suffolk (bed D) ; West Wittering, Sussex.
Preglacial (Cromer Forest-bed) : —

Sidestrand, Norfolk (?).

RUMEX Hydrolapathum, Huds.

Unknown fossil in Britain.

Recorded from the Pine and Oak Zones (Neolithic) in
Gotland (Gunnar Andersson).

RUMEX ACETOSELLA, L.
Interglacial : —

Hoxne, Suffolk (bed D) (?).

Preglacial (Cromer Forest-bed) : —

Beeston, Norfolk.

It is noteworthy that our two aquatic docks, R.
aquaticus and R. Hydrolapathum^ are still missing in the
fossil state in Britain.

HiPPOPHAE RHAMNOIDES, L.
Unknown fossil in Britain.

Recorded from Norrland in Sweden in calcareous
tufa (Nathprst), and from Gotland (Gunnar Andersson).

VTSCUM ALBUM, L.

Unknown fossil in Britain.

Recorded from the Oak Zone (Neolithic) in Gotland
(Gunnar Andersson).

Euphorbia Helioscopia, L.
Neolithic: —

Redhall, near Edinburgh, with weeds of cultivation.

142 Origin of the British Flora.

Euphorbia amygdaloides, L.

Preglacial (Cromer Forest-bed) : —

Mundesley, Norfolk ; Pakefield, Suffolk.

Mercurialis perennis, L.
Neolithic: —

Crossness, Essex (upper and lower peats) ; Hailes, near
Edinburgh; Fillyside, near Edinburgh (raised beach).

Interglacial: —

West Wittering, Sussex.

UlMUS MONTANA, Sm.

Blashenwell, Dorset ; Dursley, Gloucester (in calcareous
tufa of doubtful age); Digby Fen (elm-wood recorded by
Skertchly from a depth of lO feet).

Interglacial: —

Grays, Essex (badly preserved leaves).

Preglacial (Cromer Forest-bed) : —
Happisburgh, Norfolk.

In each case the leaves are small, and more like
U. montana than like U. campestris. The difference in
the leaves is very slight, and I have not yet been able
to obtain the more characteristic fruit.

Urtica dioica, L.
Late Glacial : —

Hoxne, Suffolk (bed C) (one seed, perhaps derived from
the bed below).

Interglacial : —

Hoxne, Suffolk (bed D;.

Former Distribution of British Plants. 143

Myrica Gale, L.
Unknown fossil in Britain.

Recorded from the Oak Zone in Gotland, Sweden
(Gunnar Andersson); from Fahrenkrug in Holstein asso-
ciated with Brasenia (Carl Weber) ; and from Honerdingen,
in Hanover, associated with Platanus, Juglans, and Najas
fiexilis (C. Weber).

Betula alba, L.
Neolithic: —

Drope, Glamorgan ; Southampton Docks (recorded
by Messrs. Shore & Elwes); Albert Dock, N. Woolwich;
Crossness, Essex (in lower and upper peats), (Spurrell);
Whittlesey Mere, Fenland (Skertchly) ; Caerwys, Flint-
shire ; Hailes, near Edinburgh ; Cowden Glen, Renfrew-
shire. Also common in peat-mosses nearly everywhere.

Late Glacial : —

Bovey Tracey, Devon (A. G. Nathorst) ; Close y
Garey, Isle of Man ; Crianlarich, Perthshire.

Interglacial : —

West Wittering, Sussex.

Preglacial (Cromer Forest-bed) : —
Common nearly everywhere.

Betula nana, L.
Late Glacial: —

Bovey Tracey, Devon (Heer); Hoxne, Suffolk (bed C);
Holmpton, East Yorkshire ; Bridlington, Yorkshire (Na-
thorst) ; Corstorphine, near Edinburgh ; Dronachy, Fife ;
Crianlarich, Perthshire.

144 Origin of the British Flora.

Interglacial: —

Admiralty Offices, London; Airdrie, Lanark.

Early Glacial: —

Beeston, Norfolk; Ostend, Norfolk.

AlNUS GLUTINOSA, Linn.
Neolithic: —

Submerged peats of the Thames Valley at Tilbury
Docks, Albert Docks, and Crossness ; Kings Lynn ; Horn-
sea, E. Yorkshire; Sand le Meer, E. Yorkshire; Redhall,
near Edinburgh; Hailes, near Edinburgh; Fillyside, near
Edinburgh (raised beach).

Late Glacial : —

Hoxne, Suffolk (bed C;; Hailes, near Edinburgh (lower
bed).

Interglacial: —

Grays, Essex ; Shacklewell, London ; Hitchin, Hert-
fordshire; Hoxne, Suffolk (beds D and E) ; Southelmham,
Suffolk.

Early Glacial : —

Beeston, Norfolk (base of Arctic Freshwater Bed).

Preglacial (Cromer Forest-bed) : —

At most localities in Norfolk and Suffolk.

Carpinus Betulus, L.
Interglacial : —

Hoxne, Suffolk (bed D).

Preglacial (Cromer Forest-bed) : —
Pakefield, Suffolk.

Former Distribution of British Plants. 145

Also recorded from Lauenburg an der Elbe (Keilhack) ;
from Klinge bei Cottbus, in Prussia (Carl Weber); and
from Grlinenthal in Holstein (Carl Weber), in each case
associated with Brasenia, 8ic.

CORYLUS AVELLANA, L.
Neolithic : —

Southampton Dock (submerged peat) ; Blashenwell,
Dorset (in tufa) ; Barry Docks, Glamorgan ; Albert Dock,
Essex ; Whittlesey Mere, Fenland (peat at 20 feet) ;
Northampton ; Sand le Meer, East Yorkshire ; Hull ;
Hailes, near Edinburgh ; Cowden Glen, Renfrewshire ; and
common in the * submerged forests ' nearly everywhere.

Interglacial : —

West Wittering, Sussex ; Grays, Essex (a doubtful
fragment of a nut); Hoxne, Suffolk (bed D); Overtown,
near Beith, Ayrshire (between two tills) (Mr. C. Craig).

Preglacial (Cromer Forest-bed) : —

Ostend, Norfolk; Pakefield, Suffolk.

The hazel, though so abundant in Postglacial deposits,
is rare in the Interglacial and Preglacial strata.

QUERCUS ROBUR, L.
Neolithic : —

Common in the * submerged forests ' everywhere ;
Blashenwell, Dorset (in tufa) ; Northampton (old river
bed) ; at base of peat mosses in Yorkshire up to a height of
1000 feet; Hailes and Redhall, near Edinburgh.

Interglacial : —

Stone, Hampshire ; West Wittering, Sussex ; Selsey,
Sussex ; Grays, Essex ; Shacklewell, London ; Hitchin,
Hertfordshire; Hoxne, Suffolk (Prestwich).

146 Of'igm of the British Flora.

Preglacial (Cromer Forest-bed): —

Ostend, Norfolk; Happisburgh, Norfolk; Pakefield,
Suffolk.

Castanea sativa, Mill.

(Ridley, /^2/r«. BoL, 1885, P- 253.)

Charcoal of Chestnut was discovered by Mr.H. N. Ridley
associated with Palaeolithic implements between Crayford
and Erith in Kent.

The Chestnut is not usually considered to be a native
of Britain ; but Mr. Ridley suggests that owing to the
value of the fruit any trees found would be enclosed and
become private property at an early date. I have not yet
discovered any corroborative evidence ; but as the tree is a
dry-soil species it can only be expected to occur rarely
in the fossil state. Large beams of Chestnut are not
uncommon in old castles and abbeys ; these may be of
foreign origin, for they are associated with building-stone
which has undoubtedly come by water.

Fagus sylvatica, L.
Neolithic : —

Southampton Docks (Shore and Elwes) ; Crossness,
Essex (wood determined by Marshall Ward) ; Fenland
(A. Bell).

Preglacial (Cromer Forest-bed) : —

Happisburgh, Norfolk.

Also recorded from Fahrenkrug, in Holstein, associated
with Brasenia (Carl Weber); and from Honerdingen, in
Hanover, associated with Juglans and Platamis (Carl
Weber).

Former Distribution of British Plants. 147

Salix pentandra, L.

Unknown fossil in Britain.

Recorded from the Pine Zone (Neolithic) in Gotland
(Gunnar Andersson) ; and doubtfully from Griinenthal, in
Holstein (Carl Weber).

Salix cinerea, L.

Neolithic (?) :—

Caerwys, Flintshire (in calcareous tufa).

Late Glacial : —

Bovey Tracey (Heer and Nathorst).

Interglacial : —

West Wittering, Sussex.

Preglacial (Cromer Forest-bed) : —

Mundesley (Nathorst).

Also recorded from Fahrenkrug in Holstein (C.Weber);
from Klinge bei Cottbus, Prussia (C.Weber); in the Birch,
Pine and Oak Zones (Neolithic) in Sweden (Gunnar
Andersson).

Salix aurita, L.

Unknown fossil in Britain.

Recorded from Fahrenkrug and Klinge, associated with
Brasenia (C. Weber); from the Birch, Oak, and Spruce
Zones in Sweden (Gunnar Andersson).

Salix Caprea, L.
Neolithic : —

Barry Docks, Glamorgan ; Fenland (A. Bell) ; Caerwys,
Flintshire.

148 Origin of the British Flora.

Also recorded from Sweden in the Birch, Pine, and
Oak Zones (Gunnar Andersson) ; and from Griinenthal, in
Holstein, associated with Brasenia (Carl Weber).

Salix phylicifolia, L.
Unknown fossil in Britain.

Recorded from Sweden in the Dryas (Late Glacial), and
Birch and Spruce (Neolithic) Zones (Gunnar Andersson).

Salix nigricans, Sm.

Unknown fossil in Britain.

Recorded from Sweden in the Pine, Oak, and Spruce
Zones (Gunnar Andersson).

Salix repens, L.
Neolithic : —

Fenland (A. Bell); Barnwell, Cambridge; Cowden
Glen, Renfrewshire.

Late Glacial : —

Corstorphine, near Edinburgh ; Crianlarich, Perthshire.

Also recorded from Klinge bei Cottbus in Prussia
(Carl Weber) ; and from Lauenburg an der Elbe (Keilhack).
In each case it is associated with Brasenia.

Salix lanata, L.
Unknown fossil in Britain.

Recorded from the Pine Zone in Norrland (Gunnar
Andersson).

Salix Arbuscula, L.
Unknown fossil in Britain.
Recorded from Sweden in the Birch and Spruce Zones

Former Distribution of British Plants. 149

(Gunnar Andersson) ; and with doubt from Deuben, in
Saxony (A. G. Nathorst).

Salix Myrsinites, L.
Late Glacial : —

Hoxne, Suffolk (bed C).

Salix herbacea, L.
Late Glacial : —

Hoxne, Suffolk (bed C); Ballaugh, Isle of Man; Kirk
Michael, Isle of Man ; Hailes, near Edinburgh (lower
bed) ; Corstorphine, near Edinburgh ; Gayfield, Edinburgh ;
Dronachy, Fife; Crianlarich, Perthshire.

Interglacial : —

Faskine, Lanark.

Also recorded from Deuben in Saxony (A . G. N athorst)
and from various localities further north.

Salix polaris, Wahlb.
Late Glacial : —

Hoxne, Suffolk (bed C); Hailes, near Edinburgh (lower
bed) ; Gayfield, Edinburgh ; Corstorphine, Edinburgh ;
Dronachy, Fife.

Early Glacial : —

Beeston, Mundesley, Ostend (in Norfolk).

Salix reticulata, L.
Late Glacial : —

Hailes, near Edinburgh (lower bed); Gayfield, Edin-
burgh; Corstorphine, Edinburgh; Dronachy, Fife; Crian-
larich, Perthshire.

T50 Origin of the British Flora,

POPULUS CANESCENS, Sm.

Interglacial : —

Grays, Essex ; some leaves collected by Prestwich
suggest this species, though they may belong to P. tremula.
The specimens have suffered from long keeping.

POPULUS TREMULA, L.
Neolithic : —

Caerwys, Flintshire, in calcareous tufa.

Recorded from Klinge bei Cottbus, in Prussia (Carl
Weber); from Honerdingen, in Hanover (Carl Weber);
also from the Birch, Pine, Oak, and Spruce Zones in
Sweden (Gunnar Andersson).

Empetrum nigrum, L.
Neolithic : —

Close y Garey, Isle of Man (beds B and C).
Late Glacial : —

Hailes, near Edinburgh (lower bed) ; Dronachy, Fife ;
Corstorphine, near Edinburgh (lower bed); Crianlarich
Perthshire; Ballaugh, Isle of Man.

Interglacial : —

Airdrie, Lanark.

Also recorded from Honerdingen, in Hanover, asso-
ciated with Platanus^ Juglans, Najas, &c. (Carl Weber).

Ceratophyllum demersum, L.

Neolithic (?) :—

Casewick, Lincolnshire.

Late Glacial : —

Hoxne, Suffolk (bed C).

Former Distribution of British Plants. 1 5 1

Interglacial : —

West Wittering, Sussex; Admiralty Offices, London
(Abbott); Hitchin, Hertfordshire; Southelmham, Suffolk;
Hoxne, Suffolk (beds D and E) ; Mundesley, Norfolk (old
valley deposit).

Early Glacial : —

Beeston, Norfolk (base of Arctic Freshwater bed).
Preglacial (Cromer Forest-bed) : —

Common at nearly all localities.

JUNIPERUS COMMUNIS, L.

Unknown fossil in Britain.

Recorded from the Dryas, Birch, Pine, Oak, and Spruce
Zones in Sweden (Gunnar Andersson) ; also from Honer-
dingen, in Hanover, associated with Platamis, Juglans, Sic.
(Carl Weber) ; and from Griinenthal, in Holstein, asso-
ciated with Brasenia (Carl Weber).

Taxus baccata, L. _
Neolithic : —

Common in peat below the sea-level in the Thames
Valley and Fenland ; Portobello, near Edinburgh.

Interglacial : —

Hoxne, Suffolk (bed D).

Preglacial (Cromer Forest-bed) : —

Mundesley, Bacton, Happisburgh (in Norfolk); Pake-
field (in Suffolk).

PiCEA EXCELSA, Link.
Preglacial (Cromer Forest-bed) : —

Cromer, Mundesley, Bacton, Happisburgh, in Norfolk.

152 Origin of the British Flora.

Unknown in Britain in later deposits. Recorded from
the Spruce Zone (Neolithic) in Sweden (Gunnar Andersson);
and from Fahrenkrug, and GrUnenthal in Holstein (Carl
Weber) ; from Klinge bei Cottbus, Prussia (Carl Weber) ;
and from Honerdingen, in Hanover, associated with
Platanus,Juglans, &c. (Carl Weber).

PiNUS MONTANA, Mill.

Preglacial (Cromer Forest-bed) : —

Determined by Heer and figured by Saporta from the
Cromer Forest-bed, but I can find no specimens belonging
to this species. Small cones of P. sylvestris may have
been mistaken for P. montana.

PiNUS SYLVESTRIS, L.
Neolithic : —

In * submerged forests ' and at the base of peat-mosses
nearly throughout Britain and in Ireland.

Late Glacial : —

BoveyTracey, Devon (Heer); Hoxne, Suffolk (bed C) (?).

Preglacial (Cromer Forest-bed) : —

Common at various localities in Norfolk, especially
Cromer and Happisburgh.

The distribution in space and time of the Scotch Pine
is very peculiar. Abundant in the Preglacial Strata of
Norfolk, it has not been found in any of the Interglacial
Deposits in Britain, though occurring at Fahrenkrug and
Griinenthal in Holstein. In Late Glacial times it reap-
pears at Bovey Tracey, in Devon, and perhaps at Hoxne,
in Suffolk. During the Neolithic period it seems to have
been one of our commonest trees ; but afterwards disap-

Former Distribution of B^'itisk Plants. 153

p eared from the southern half of England ; though, when
re-introduced, it flourishes and spreads rapidly from
seedlings.

Stratiotes aloides, L.
Interglacial : —

Southelmham, Suffolk.
Preglacial (Cromer Forest-bed) : —

Beeston, Sidestrand (in Norfolk); Gorton (in Suffolk).

The fruits described by Nehring, Potonie, and myself
as Paradoxocarpus (or Folliculites) carinatus have since
been shown to belong to Stratiotes aloides. They occur
abundantly at Klinge bei Cottbus,- in Prussia, and at
Fahrenkrug in Holstein.

Iris Pseudacorus, L.
Neolithic : —

Crossness, Essex (upper peat) ; Elie, Fife.

Sparganium ramosum, Curtis.
Neolithic : —

Barry Docks, Glamorgan ; Crossness, Essex (upper and
lower peats); Tilbury, Essex; Redhall, near Edinburgh.

Late Glacial : —

Hoxne, Suffolk ibed C) ; Garvel Park (Clyde Beds)
(two very small and doubtful carpels).

Interglacial : —

West Wittering, Sussex ; Hitchin, Hertfordshire ;
Hoxne, Suffolk (beds D and E).

Preglacial (Cromer Forest-bed) : —

Beeston, Norfolk ; Pakefield, Suffolk ; Corton, Suffolk.

154 Origin of the British Flora.

Sparganium simplex, Huds.

Unknown fossil in Britain.

Recorded from Honerdingen, in Hanover (Carl Weber).

Sparganium minimum, Fr.

Unknown fossil in Britain.

Recorded from Honerdingen, in Hanover (Carl Weber).

Alisma Plantago, L.
Neolithic : —

Redhall, near Edinburgh; Cowden Glen, Renfrewshire.

Late Glacial : —

Hoxne, Suffolk (bed C).

Interglacial : —

Hoxne, Suffolk (bed D) ; Southelmham, Suffolk ; West
Wittering, Sussex ; Stone, Hampshire.

Preglacial (Cromer Forest-bed) : —

Beeston, Overstrand, Sidestrand, Mundesley,in Norfolk;
Pakefield, in Suffolk.

SAGITTARIA SAGITTiEFOLIA, L.
Interglacial : —

West Wittering, Sussex.

Recorded from the Oak Zone in Gotland (Gunnar
Andersson).

• SCHEUCHZERIA PALUSTRIS, L.

Unknown fossil in Britain.

Recorded from the Dry as and Oak Zones in Sweden
(Gunnar Andersson).

Former Distribution of British Plants. 155

POTAMOGETON NATANS, L.
Neolithic : —

Drope, Glamorgan.

Late Glacial : —

Ballaugh, Isle of Man (bed F).
Interglacial : —

West Wittering, Sussex.

Also recorded from Klinge bei Cottbus, in Prussia ;
Fahrenkrug in Holstein \ Honerdingen, in Hanover (Carl
Weber) ; and from the Birch, Pine, and Oak Zones in
Sweden (Gunnar Andersson).

POTAMOGETON RUFESCENS, Schrad.

Late Glacial : —

Twickenham, Middlesex ; Hoxne, Suffolk (bed C).

Interglacial : —

Kilmaurs, Ayrshire (?) (Bennett).

Also recorded from Honerdingen, in Hanover, asso-
ciated with Platanus, Juglans, &c. (Carl Weber).

POTAMOGETON HETEROPHYLLUS, Schreb.
Neolithic : —

Drope, Glamorgan ; Hailes, near Edinburgh.

Interglacial : —

West Wittering, Sussex ; Stone, Hampshire ; South-
elmham, Suffolk; Kilmaurs, Ayrshire (or P. Zizii)
(Bennett).

Preglacial (Cromer Forest-bed) :—
Abundant at most localities.

156 Origin of the British Flora.

POTAMOGETON LUCENS, L.
Neolithic : —

Cowden Glen, Renfrewshire.

Interglacial : —

West Wittering, Sussex.

Preglacial (Cromer Forest-bed) : —

Between Cromer and Runton, in Norfolk ; Corton,.
Pakefield, in Suffolk.

POTAMOGETON PR.ELONGUS, Wulf
Neolithic : —

Megaceros-marls of Central Ireland.

Preglacial (Cromer Forest-bed) : —

Between Cromer and Runton.

Recorded from the Dryas, Birch, and Pine Zones (Late
Glacial and Neolithic) in Sweden (Gunnar Andersson);
and with doubt from Honerdingen, in Hanover, with
fuglans, Platanus, &c. (Carl Weber).

POTAMOGETON PERFOLIATUS, L.
Neolithic : —

Redhall, near Edinburgh ; Hailes, near Edinburgh ;
Cowden Glen, Renfrewshire.

Interglacial : —

Southelmham, Suffolk.

Also recorded from Honerdingen, in Hanover, asso-
ciated with Juglans, Platanus , &c. (Carl Weber).

POTAMOGETON CRISPUS, L.
Neolithic : —

Gayfield, near Edinburgh; Mes^aceros-marls of Central
Ireland.

Former Distribution of British Plants. 157

Late Glacial : —

Hoxne, Suffolk fbed C) ; Ballaugh, Isle of Man (bed D).

Interglacial : —

Endsleigh Street, London ; Hitchin, Hertfordshire ;
Southelmham, Suffolk.

Preglacial (Cromer Forest-bed) : —

Beeston, Sidestrand,Trimingham,Mundesley, in Norfolk.

POTOMAGETON DENSUS, L.
Interglacial : —

West Wittering, Sussex.

POTAMOGETON OBTUSIFOLIUS, Mert. & Koch.
Interglacial : —

Endsleigh Street, London; Southelmham, Suffolk.

POTAMOGETON PUSILLUS, L.
Neolithic : —

Hailes, near Edinburgh ; Redhall, near Edinburgh ;
Cowden Glen, Renfrewshire.

Late Glacial : —

Hoxne, Suffolk (bed C).

Interglacial : —

Hoxne, Suffolk (bed D).

POTAMOGETON TRICHOIDES, Cham.
Late Glacial: —

Hoxne, Suffolk (bed C).

Interglacial : —

West Wittering, Sussex ; Stone, Hampshire; Southelm^
ham. Suffolk ; Hoxne, Suffolk (beds D and E).

158 Origin of the British Flora.

Preglacial (Cromer Forest-bed) : —

Beeston, Sidestrand, Mundesley, in Norfolk ; Pakefield,
Gorton, in Suffolk.

POTAMOGETON PECTINATUS, L.
Neolithic : —

Cowden Glen, Renfrewshire.

Late Glacial : —

Hoxne, Suffolk (bed C).

Preglacial (Cromer Forest-bed) : —

Common at most localities — drupes often very large.

POTAMOGETON FILIFORMIS, Nolte.
Unknown fossil in Britain.

Recorded from Gotland in the Dryas, Birch, and Pine
Zones (Gunnar Andersson).

RUPPIA MARITIMA, L.
Interglacial : —

West Wittering, Sussex ; Stone, Hampshire.
Recorded from the Pine and Oak Zones in Sweden
(Gunnar Andersson).

Zannichellia palustris, L.
Late Glacial : —

Twickenham, Middlesex.
Interglacial : —

Stone, Hampshire; Selsey, Sussex; Endsleigh Street,
London; Southelmham, Suffolk; Hoxne, Suffolk (bed E);
Kilmaurs, Ayrshire.

Former Distribution of British Plants. 159

Early Glacial : —

Beeston, Norfolk (base of Arctic Freshwater Bed).
Preglacial (Cromer Forest-bed) : —

Abundant nearly everywhere.

Zannichellia pedunculata, Reichb.
Interglacial : —

West Wittering, Sussex.

Preglacial (Cromer Forest-bed) : —

Sidestrand, Norfolk ; Pakefield, Suffolk.

At Pakefield a remarkable spinose form of drupelet
occurs.

ZOSTERA MARINA, L.

Not certainly known fossil in Britain, though Zostera-
like foliage occurs in estuarine deposits.

Recorded with doubt by Gunnar Andersson from South
Sweden and Gotland.

Najas flexilis, Rostkov.

Unknown fossil in Britain.

Recorded from Gotland, and from the^^^^/z/.y-formation
(Pine Zone) in Sweden (Gunnar Andersson) ; also from
Honerdingen, Hanover, and Griinenthal, Holstein, in Inter-
glacial deposits (C. Weber).

This plant in Europe is only known living in the West
of Ireland, Scotland, South Sweden, and Gotland.

Najas marina, L.
Neolithic : —

Barry Docks, Glamorgan.

Interglacial : —

Hitchin. Hertfordshire.

1 60 Origin of the B^Htish Flora.

Preglacial (Cromer Forest-bed) : —

Beeston, in Norfolk ; Pakefield, in Suffolk.

Also recorded from the Pine and Oak Zones of South

Sweden and Gotland (Gunnar Andersson) ; from Klinge

bei Cottbus in Prussia, Fahrenkrug in Holstein, and Honer-

dingen in Hanover (Carl Weber).

Najas graminea, Delile.
Interglacial : —

West Wittering, Sussex.

A widely dispersed Tropical species, which extends into
the Mediterranean region, and occurs as an accidental
introduction into Britain in a canal which receives waste
hot water from a mill. As a fossil it has only been
recorded at West Wittering.

Najas minor, Allione.
Interglacial : —

West Wittering, Sussex.
Preglacial (Cromer Forest-bed) : —

Pakefield, Suffolk.

A plant of the Mediterranean region, and of central
Europe as far north as the Rhine ; it is unknown living in
the north or in Britain.

Eleocharis acicularis, Sm.
Roman Period : —

Silchester, Hampshire.
Interglacial : —

Hoxne, Suffolk (bed D); West Wittering, Sussex;
Stone, Hampshire.

Former Distribution of British Plants. i6i

Eleocharis palustris, Br.
Roman Period : —

Silchester, Hampshire.

Neolithic : —

Hailes, near Edinburgh (upper bed) ; Redhall, near
Edinburgh; Megaceros-marls of Central Ireland.

Late Glacial :—

Twickenham, Middlesex; Kirk Michael, Isle of Man;
Corstorphine, near Edinburgh ; Hailes, near Edinburgh
(lower bed) ; Dronachy, Fife.

Interglacial : —

West Wittering, Sussex ; Endsleigh Street, London ;
Allenton, near Derby.

Early Glacial : —

Beeston, Norfolk (base of the Arctic Freshwater Bed).

Preglacial (Cromer Forest-bed) : —
Pakefield, Suffolk.

SCIRPUS PAUCIFLORUS, Lightf.
Neolithic : —

Hailes, near Edinburgh ; Redhall, near Edinburgh ;
Cowden Glen, Renfrewshire.

Late Glacial : —

Hoxne, Suffolk (bed C); Stair, Ayrshire; Hailes, near
Edinburgh (lower bed) ; Corstorphine, near Edinburgh ;
Dronachy, Fife.

Interglacial : —

Southelmham, Suffolk; Hoxne, Suffolk (bed D); Allen-
ton, near Derby.

M

1 62 Origin of the British Flora.

Preglacial (Cromer Forest-bed): —

Beeston and Mundesley, in Norfolk ; Gorton and Pake-
field, in Suffolk.

SCIRPUS C^SPITOSUS, L.
Interglacial : —

Southelmham, Suffolk.

Preglacial (Cromer Forest-bed) : —
Beeston and Mundesley, in Norfolk.

SCIRPUS FLUITANS, L.
Late Glacial : —

Dronachy, Fife (?).

Interglacial : —

Southelmham, Suffolk; Kirmington, Lincolnshire (?);
Stone, Hampshire (?).

Preglacial (Cromer Forest-bed) : —
Beeston, Norfolk.

SCIRPUS SETACEUS, L.

Neolithic : —

Hailes, Redhall, and Broughton, near Edinburgh.

Late Glacial : —

Hoxne, Suffolk (bed C).

Interglacial : —

Hoxne, Suffolk (bed D).

SCIRPUS LACUSTRIS, L.
Neolithic : —

Crossness, Essex (lower peat); Casewick, Lincolnshire;
Hailes, near Edinburgh ; Cowden Glen, Renfrewshire.

Former Distribution of British Plants. 163

Late Glacial : —

Twickenham, Middlesex ; Hoxne, Suffolk (bed C) ;
Hailes, near Edinburgh (lower bed) ; Corstorphine, near
Edinburgh.

Interglacial : —

Stone Hampshire; West Wittering, Sussex; Hitchin,
Hertfordshire; Hoxne, Suffolk (beds D and E); Southelm-
ham, Suffolk.

Early Glacial : —

Beeston, Norfolk (base of Arctic Freshwater bed).

Preglacial (Cromer Forest-bed) : —

Beeston and Mundesley, in Norfolk ; Gorton and Pake-
field, in Suffolk.

SCIRPUS MARITIMUS, L.
Neolithic : —

SoutJiampton Docks (?) (a single damaged nut); Barry
Docks, Glamorgan.

SCIRPUS SYLVATICUS, L.

Unknown fossil in Britain.

Recorded from Vernitsa, St. Petersburg (Gunnar
Andersson and Berghell) ; also from the Pine and Spruce
Zones in Sweden (Gunnar Andersson).

"^
Blysmus rufus, Wahlb.
Late Glacial : —

Hoxne, Suffolk (bed C).

Interglacial : —

Hoxne, Suffolk (bed Dj.

164 Origin of the British Flora.

Eriophorum vaginatum, L.

Unknown fossil in Britain.

Recorded from Fahrenkrug, in Holstein (Carl Weber);
and from the Pine (?) Oak (?) and Spruce Zones in Sweden
(Gunnar Andersson).

Eriophorum angustifolium, Roth.

Interglacial : —

Hoxne, Suffolk (bed D).

Preglacial (Cromer Forest-bed) : —

Between Cromer and Runton, Norfolk ; Corton, Suffolk.

Recorded also from the Pine and Oak (?) Zones
(Neolithic) in Sweden (Gunnar Andersson).

Cladium Mariscus, Br.

Unknown fossil in Britain, the fruits recorded from the
Cromer Forest-bed not belonging to this plant.

Recorded from Klinge bei Cottbus, in Prussia (Carl
Weber); and from the Birch (?) Pine, and Oak Zones in
Gotland (Gunnar Andersson).

Carex dioica, L.
Neolithic : —

Redhall, near Edinburgh; Hailes, near Edinburgh.

Late Glacial : —

Roxburgh Street, Greenock (Clyde Beds).

Interglacial : —

Endsleigh Street, London ; Airdrie, Lanark.

Former Distribution of British Plants. 165

Carex muricata, L.
Interglacial : —

West Wittering, Sussex; Stone, Hampshire.

Carex echinata, Murr.
Neolithic : —

Hailes, near Edinburgh; Redhall, near Edinburgh.

Also recorded from Fahrenkrug, in Holstein (Carl

Weber).

Carex remota, L.

Preglacial (Cromer Forest-bed) : —
Pakefield, Suffolk.

Carex alpina, Sw.
Late Glacial : —

Kirk Michael, Isle of Man (determined by C. B. Clarke).

Carex canescens, L.
Neolithic : —

Hailes, near Edinburgh ; Redhall, near Edinburgh.

Carex panicea, L.
Neolithic : —

Redhall, near Edinburgh.

Late Glacial : —

Twickenham, Middlesex.

Interglacial : —
Airdrie, Lanark.

Carex distans, L.
Interglacial : —

West Wittering, Sussex; Hoxne, Suffolk (bed D) (?).

1 66 Origin of the British Flora.

' Carex flava, L.
Neolithic : —

Hailes, near Edinburgh; Redhall, near Edinburgh.

Carex filiformis, L.

Unknown fossil in Britain.

Recorded from Vernitsa, St. Petersburg ( Gunnar
Andersson and Berghell); also from the Birch, Pine, Oak,
and Spruce Zones in Sweden (Gunnar Andersson).

Carex Pseudo-cyperus, L.

Unknown fossil in Britain.

Recorded from Lauenberg an der Elbe (Carl Weber) ;
and from the Pine, Oak, and Spruce Zones in Sweden
Gunnar Andersson).

Carex paludosa. Good.

Preglacial (Cromer Forest-bed) : —
Pakefield, Suffolk.

Carex riparia, Curtis.
Interglacial : —

Stone, Hampshire; West Wittering, Sussex; Southelm-
ham, Suffolk.

Preglacial (Cromer Forest- bed) : —

Overstrand, Mundesley (?) in Norfolk ; Pakefield, in
Suffolk.

Also recorded from the Pine (?) and Oak Zones in
South Sweden (Gunnar Andersson).

Former Distribution of British Plants. 167

Carex rostrata, Stokes.
Neolithic : —

Cowden Glen, Renfrewshire.

Interglacial : —

West Wittering, Sussex; Stone, Hampshire; Southelm-
ham, Suffolk ; Hoxne, Suffolk (bed D) (?) ; Airdrie, Lanark.

Carex vesicaria, L.

Unknown fossil in Britain.

Recorded from the Pine and Oak Zones in Sweden
(Gunnar Andersson).

Phragmites communis, Trin.
Neolithic : —

Thames Valley (common) ; Barry Docks, Glamorgan ;
Kelsey Hill, Yorkshire.

Interglacial : —

Stone, Hants ; West Wittering, Sussex ; Kirmington ,
Lincolnshire.

Preglacial (Cromer Forest-bed) : —
Common nearly everywhere.

Stems, leaves, or nodes of grass are common at most
localities ; but the only species in a determinable state
appears to be Phragmites communis. Anthroxanthum'
odoratum, Holus lanatus^ Poa trivialis, and Hordeum
distichum have all been recorded ; but in each case I think
that the specimens are recent and do not belong to the
deposit in which they are said to occur. Extreme care
is needed to prevent the introduction of grass-seeds, which
are dispersed by the wind and adhere to the surface of
the clays containing the fossil plants.

1 68 Origin of the British Flora.

Pteris aquilina, L.

Though often stated to occur in a fossil state I have
seen no undoubted specimens in Britain.

Recorded from South Sweden (Gunnar Andersson).

Athyrium Filix-fcemina, Roth.

Unknown fossil in Britain.

Recorded from the Oak Zone in Gotland (Gunnar
Andersson).

SCOLOPENDRIUM VULGARE, Sm.

Found in calcareous tufa of doubtful age at Dursley, in
Gloucestershire.

Lastr^a Thelypteris, Presl.

Unknown fossil in Britain.

Recorded from the Oak and Spruce Zones in South
Sweden (Gunnar Andersson) ; and from Klinge bei Cottbus,
in Prussia (Carl Weber).

OSMUNDA REGALIS, L.

Neolithic : —

' Submerged forests ' near Liverpool.

Preglacial (Cromer Forest-bed) :—

Common at Mundesley, Norfolk ; rare elsewhere.

Equisetum palustre, L.
Unknown fossil in Britain.
Recorded from Honerdingen, in Hanover (Carl Weber).

Former Distribution of British Plants. 169

Equisetum limosum, Sm.

Though fragments possibly belonging to this species
are not uncommon in peaty deposits in Britain, I have
seen no determinable specimens.

Recorded from Lauenburg an der Elbe (Keilhack).

Equisetum hyemale, L.

Unknown fossil in Britain.

Recorded from the Pine Zone in Norrland, in Sweden,
in calcareous tufa (Nathorst) ; and from Gotland (Gunnar
Andersson).

Isoetes lacustris, L.
Neolithic : — •

Hailes, near Edinburgh; Cowden Glen, Renfrewshire.

Late Glacial : —

Garvel Park (Clyde Beds), (J. B. Balfour) ; Hailes, near
Edinburgh (lower beds) ; Gayfield, Edinburgh.

Interglacial : —

Allenton, near Derby; Kilmaurs, Ayrshire; Faskine,
Lanark ; Airdrie, Lanark.

Early Glacial : —

Beeston, Norfolk (base of Arctic Fresh-water-bed).

Preglacial (Cromer Forest-bed) : —

Beeston, Norfolk (a single specimen, perhaps from the
Early Glacial bed above, which here rests immediately on the
Cromer Forest-bed).

171

APPEiNDIX.

Table showing the Range in Time of the British Flora.

E - England.
W = Wales.
S = Scotland.
M = Isle of Man.
I = Ireland.

D — Denmark.

F — Finland.

G — North Germany.

jV = Norway.

S = Sweden.

Clematis Vitalba, L.

flavum, L.

Ranunculus aquatilis, L

hederaceus, L.

sceleratus, L. .

Flammula, L. .

Lingua, Z. ....

repens, L

bulbosus, Z. .

Sardous, Crantz,

parviflorus, Z. .

Caltha palustris, Z

Nuphar luteum, Z

Nymphaea alba, Z

Papaver somniferum, Z.

Argemone, Z.

Chelidonium majus, Z.
Fumaria officinalis, Z.

Thlaspi arvense, Z

Cakile maritima, Scop.
Viola palustris, Z. ,,,...

The foreign range is
only recorded where
the species is unknown
in the equivalent de-
posits in Britain.

E?

E

ES
E
E
E
E
E
E
E
E
E
E
E

ESM

E
M

ESM

ES

ESM

6-
ES I

E

S M

S
ES

S
E
E?

S.?

s?
ws

172

Table showing the Range in

Silene maritima, With.

Lychnis alba, Mill.

diurna, Sibth....

Flos-cuculi, L.

Stellaria aquatica, Scop.

• media, Cyr. ...

Holostea, L. ...

graminea, Z. ...

uliginosa, L. ...

Arenaria trinervia, L

— — ^ peploides, L

Spergula arvensis, L

Moniia fontana, Z

Hypericum perforatum, L

quadrangulum, L.

elodes, L

Tilia platyphyllos, Scop

europaea, Z

Linum, sp

Geranium columbinum, Z

Oxalis Acetosella, L

Ilex Aquifolium, Z

Rhamnus Frangula, Z

Vitis vinifera, Z

Acer campestre, Z

monspessulanum, Z

Prunus spinoga, Z

domestica, Z

— — ' — Avium, Z

— Padus, Z

Spiraea Ulmaria, Z

E
E
E?
G
E
E

E

ES
E

S
S

ES

S?
S

s

s?

s?
s

s?
s?

s?

ES
E

5

E S

E

ES
ES

S

Time of the British Flora.

173

Rubus Idaeus, L. ...

fruticosus, Z.

caesius, L. ...

saxatilis, L.

Rosa

Pyrus torminalis, Ekrk

Aria, Sm ,

Aucuparia, Gaert.

communis, L

Crataegus Oxyacantha, L.
Saxifraga oppositifolia, L.

Hirculus, L.

aizoides, L ,

Dryas octopetala, L

Potentilla Tormentilla, Neck

Comarum, Nestl.

Alchemilla arvensis, Lam

Poterium officinale, Hook

Hippuris vulgaris, L

Myriophyllum spicatum, Z

alternifolium, Z. ...

Trapa natans, Z

Hydrocotyle vulgaris, Z

Apium graveolens, Z

nodiflorum, Reich

Cicuta virosa, Z

Sium latifolium, Z

Conopodium denudatum, Koch.

Chaerophyllum temulum, Z

Anthriscus sylvestris, Hoffm

CEnanthe fistulosa, Z

E

E.?

ES
ES

G

E

E

ES

ES

S

s.?

M
ESM

G

G

G
ESM
ESM

ES
ES

S

s

S M

S
S?

S
ES

W

E

EWS

EWS
S I

S
FS
S M

174

Table showing the Range

m

(Enanthe Lachenalii, Gmel.

Phellandrium, Lam. ...

iEthusa Cynapium, L

Angelica sylvestris, L...

Peucedanum palustre, yI/^«^^. ...

Heracleum Sphondylium, L

Caucalis nodosa, Scop

Hedera Helix, L

Cornus suecica, L

sanguinea, L

Sambucus nigra, L

Viburnum Opulus, L

Galium boreale, L. ...

palustre, L. ...

uliginosum, L.

Aparine, L. ...

Valeriana officinalis, L

Valerianella olitoria, Moench.

Scabiosa succisa, L

Eupatorium cannabinum, L

Aster Tripolium, L

Bidens cemua, Z

tripartita, L

Chrysanthemum segetum, L. ...

Leucanthemum, L,

Matricaria inodora, L

Tanacetum vulgare, L

Tussilago Farfara, Z.

Senecio sylvaticus, L

aquaticus, //i?/^j

E
E

E
E
E
E

Gf
G

E
E
E
E
E

E

S?

S

E W

E W

ES

E

S.?

E?
WS

E S

S

S.?
S.?
S?

s.?

Time of the British Flora.

175

Carduus crispus, L

Cnicus lanceolatus, Willd.

palustris, Willd. ....

Centaurea Cyanus, L

Lapsana communis, L

Picris hieraciodes, L

Crepis virens, L

Hieracium Pilosella, L

Leontodon autumnalis, L. .
Taraxacum officinale, Wed.
Sonchus arvensis, L.

Vaccinium Oxycoccos, L ,

Vitis-Idaea, L

uliginosum, L

MyrtilluS; L

Arctostaphylos alpina, Spreng. ...

Uva-ursi, Spreng.

Andromeda Polifolia, L

Loiseleuria procumbens, Z^^jz/. ...

Glaux maritima, L

Fraxinus excelsior, L.

Menyanthes trifoliata, L

Myosotis sylvatica, Hoffm

Solanum Dulcamara, L

Verbascum Thaspus, L

Bartsia Odontites, Huds

Pedicularis palustris, L

Mentha aquatica, L

Lycopus europaeus, L

Thymus Serpyllum, L

Prunella vulgaris, L

E.?

E

E

ES

E
E

ES

ESM

M
S
S

S?

s

s.?
s
s?

s

s

s

E
W S

s
s

176

Table showing the Range in

Stachys palustris, L.

\ — sylvatica, L.

arvensis, L...,

Galeopsis Tetrahit, L.

Ajuga reptans, L

Littorella lacustris, L

Atriplex patula, L

Polygonum Aviculare, L

Hydropiper, L

Persicaria, L

— ^— lapathifolium, L

— — — ^— amphibium, L

viviparum, L

Oxyria digyna, Hill.

Rumex conglomeratus, Murr.

maritimus, L

obtusifolius, Z

' crispus, L.

< — Hydrolapathum, Huds

Acetosella, L

Hippophae rhamnoides, L

Viscum album, L

Euphorbia Helioscopia, L

amygdaloides, Z

Mercurialis perennis, L

Ulmus montana (?) Sm

Urtica dioica, L

Myrica Gale, L

Betula alba, L

nana, Z

Alnus glutinosa, Z

E
E
E
E

E?

E
E
E
G
E

ES
E

S

E

ES

M

S

s

GS
S

E
M

ES

E?

E S
ES
ES

S
S

s

S I
ES

S

E

S

s

5

s

EWS

5

S
S
S?

ES
E

S
EWSI

ES

Time of the British Flor^a.

^77

Carpinus Betulus, L

Corylus Avellana, L

Quercus Robur, L

Castanea sativa, Mill.

Fagus sylvatica, L

Salix pentandra, L

cinerea, L

aurita, L

Caprea, L

phylicifolia, L

nigricans, Sm

repens, L

lanata, L

Arbuscula, L

Myrsinites, L

herbacea, L

polaris, Wahlb

reticulata, L

Populus canescens, Stn. ...
tremula, Z

Empetrum nigrum, L

Ceratophyllum demersum, L

Juniperus communis, L ,...

Taxus baccata, L

Picea excelsa, Lznk

Pinus sylvestris, L

Stratiotes aloides, L

Iris Pseudacorus, L

Sparganium ramosum, Curtis ...

simplex, Huds

minimum, Fr.

E

ES

E
E?

G
Gf

E

G

G

E?
G
S
E
G
E
G

G?

E

ESM

E S

S

ES

E WS
E WS

E

W

s

EW

S

S
E S

s
s

w

M

E

S

ES

ES I

ES
E WS

X

178

Table showing the Range in

Alisma Plantago, L

Sagittaria sagittifolia, L

Scheuchzeria palustris, L

Potamogeton natans, L

rufescens, Schrad.

heterophyllus, Schreb.

— lucens, Z

praslongus, Wulf.

perfoliatus, Z

crispus, L

densus, L

obtusifolius, M. ^ K,

pusillus, L

trichoides, Chatn...

pectinatus, L

filiformis, Nolte. ..

Ruppia maritima, L

Zannichellia palustris, L

pedunculata, Reichb.

Zostera marina, L

Najas flexilis, Rostkov

marina, Z

graminea, Delile

minor, Allione

Eleocharis acicularis, Sm

palustris, Br.

Scirpus pauciflorus, Zz^A^

ca^spitosus, Z

fluitans, Z

setaceus, Z

lacustris, Z

E

E
E

E
S?
ES
E
G
E
E
E
E
E
E

E

ES
E

G
E
E
E
E
E
E
E
E
E
E

S
M
E

E M

ESM
E S

S?

E

ES

S

W
WS

s

I
s

S I

s?

s

w

S I

s

s

ES

Time of the B^'itish Flora.

179

Scirpus maritimus, L

sylvaticus, Z.

Blysmus rufus, Schrad.

Eriophorum vaginatum, L

angustifolium, Roth.

Cladium Mariscus, Br

Carex dioica, L

muricata, L

echinata, yJ/z/r^

remota, L

alpina, Sw

canescens, L

panicea, L

distans, Z

flava, L

filiformis, L

Pseudo-cypems, Z

paludosa, Good.

riparia, Curtis

rostrata, Stokes

vesicaria, Z

Phragmites communis, Trin

Pteris aquilina, Z

Athyrium Filix-foemina, Roth. ...

Scolopendrium vulgare, Sin

Lastrsea Thely pteris, Presl.

Osmunda regalis, Z

Equisetum palustre, Z

limosum, Z

hyemale, Z

Isoetes lacustris, Z

E?

E
G
E
G

ES
E
G

E
ES

E ES

M

E? W

S

s
s
s
s

s
s

s

s
s

s
s
s

E W

S
E?

S
E

i8i

INDEX.

Abbott, W. J. L., 54, 151.

Accidental dispersal of plants, 20, 21,

26, 30, 31, 97.
Acer campestre, 24, 28, 68, 85, 113.

numspessulanum, 53, 88, 95, 98,

99, "3-
— — platanoides, 75, 82, 113.

Pseudo-platanus, 12, 16, 24, 28,

Acorns, dispersal of, 23, 29, 30.
Adaptation for dispersal, 2, 4, 20-32.

to environment, 2, 4.

Admiralty Offices, London, 54.
yEtkusa Cynapium, 66, 89, 124.
Airdrie, Lanark, 54, 68.

Ajuga reptans, 137.

Albert Dock, N. Woolwich, 55, 93.

Alchemilla arvensis, 117.

Alder, 39, 52, 144.

Alisma Plant ago, 154.

ranunculoides, 81.

AUenton, near Derby, 55.
Alnus glutinosa, 39, 52, 144.
incana, 39.

Alpine flora of Britain, 2, 3, 13, 14, 20,

21, 25-27, 40, 53.
Alternations of climate, 2, 7, 8, 20, 21,

33-47. 97-
Amphibia, dispersal of, 7.
American plants in Britain, 18, 19.
America, prairie vegetation of, 15.
Andersson, Dr. Gunnar, 51, 92, loi,

106, 107, 111-116, 119, 121-125,

131-134, 136, 139, 141, 143, 147-

152, 154-156, 158-160, 163, 164,

166-169.
Andromeda Polifolia, 100, 133.
Angelica sylvestris, 124.
Animals and plants, interdependence of, i

4, 6, 23-32, 97. \
geographical distribution of,

2, 3. 6, 7.

Animals, means of dispersal of, 6, 7.

use of fossil, in classification,

S3. 54-
Annual plants, conditions needed by,

II, 12.
Anthriscus sylvestris, 123.
Apium graveolens, 122.

nodiflorutn, 122.

Apple, 28.

Apus glacialis, 56, 57.

Aquatic plants, distribution of, 14.

means of dispersal of, 26.

Arbutus, 17, 18, 26, 42.
Arctic mammals in Britain, 7, 20, 21,
40-42, 45, 81, 83, 93, 94.

plants in Britain, 2, 13, 14, 20, 21,

25-27, 37. 39. 40. 50. 52, S3. 79. 99.
loo.

Regions, plants of the, 39, 99,

loo.

Willows, 14, 57-59, 62, 64, 65,

68, 70. 73. lly 78, 83, 84, 99, 149.
Arctostaphylos alpina, 132.

Uva-ursi, 21, 23, 42,

100, 132.
A renaria peploides, 1 10.

trinervia, 82, 109.

Armistead, J. J., on dispersal of acorns,

29, 30.
Arnold- Bemrose, H. H., 55.
Ash, 24, 28, 69, 82, 133.
Aster Tripoliuni, 128.
Atriplex patula, 138.
Athyrium Filix-foeniina, 168.
Auchtertool, 66.
Ayrshire, 66, 79, 80, 84.

Bacton, 56.

Baker, E. G. , on Kerry Plants, 4. 5.

l82

Index.

Balfour, Prof. I. B., 169.

Ballaugh, 56, 57, 61, 62, 79, 80.

Baltic ice-sheet, 39.

Barren-lands, plants of, 15.

Barriers, influence of, 7, 14, 19, 22, 24-

33, 39, 46, 47.
Barry Docks, 46, 53, 57, 58.
Bartsia Odontites, 135.
Beach-plants, 13, 14.
Beans carried by wood-pigeon, 30.
Bearberry, 21, 23, 42, 100.
Beech, 28, 69, 146.
Beeston, 58, 59.
Beetles, fossil, 6, 7, 54.
Beith, Overtoun near, 84.
Beldorf, Holstein, 72.
Belgium, lowland flora of, 13.
Bell, Alfred, 69, 146-148.

A. M. , 84.

Bennett, Arthur, 80, 155.

Bennie, J., 4, 60, 62, 63, 65, 66, 68, 69,

70. 72, 73. 79. 86, 132.
Bergen, Horse-chestnut at, 11.
Berghell, Hugo, 163, 166.
Berry-bearing plants, dispersal of, 23.
Betula alba, 52, 143.
Betula nanUy 21, 41, 42, 52, 60, 69, 72,

75, 100, 143, 144.
Bidens cernua, 128.

tripartita, 128.

Biennial plants, origin of, 12.
Birch, Arctic, 21, 41, 42, 52, 100.
Birds carrying seeds, 23, 26-32, 97.

great mortality among, 30.

eating poisonous seeds, 31.

Blackberry, 26, 115. 116.
Black Burn, East Tarbet, 59.
Blashenwell, 53, 59.

Blysmus rufus, 163.

Bog iron-ore, plants of, 50, 74.

Boraginece, distribution of the, 17.

Bos longifrons at Twickenham, 93,

Bournemouth Pine, 12, 99.

Bovey Tracey, 42, 53, 59, 60.

Bracken, 168,

Brasenia purpurea, 53, 54, 68, 72, 81,

82, 106.
Bridlington, 53, 60.
Broads, flora of the, 14.
Broom, prostrate variety of, 4.
Broughton, near Edinburgh, 60.
Bulrush, 26, 76.

Burrs, dispersal of, 20, 24, 26.
Butcher's Broom, 11, 12.

Cae;wys, Flintshire, 60, 61.

Cakile maritinia , 107.

Calcareous soils, plants of, 17.

Calcareous tufa, plants in, 51, 59-61, 66.

Caltha paluslrls, 105.

Candler, C. , 90,

Canoes in Clyde Beds, 70.

Cantrill, T. C. , 66.

Capsules, use of, 23, 24.

Cardiff, Drope near, 66.

Carduus crispus, 129.

Carex alpina, 100, 165,

canescens, 165.

dioica, 164.

distans, 165.

echinata, 165

filiformis, 166.

flava, 166.

muricata, 165.

paludosa, 166.

panicea, 165.

Pseudo-cyperus, 166,

remota, 165.

riparia, 166.

rest rata, 167.

vesicaria, 167.

Carpinus Betulus, 16, 24, 26, 75, 144.

Carruthers, W. , i, 112.

Carum Carui, 122, 123.

Caryophyllacece, dispersal of, 24, 26.

Casewick, 61.

Castanea saiiva, 16, 146.

Caucalis nodosa, 125.

Centaurea Cyanus, 87, 130.

Central Europe, plants of, 18. 25, 99.

Ceratophylluin demersum, 150, 151.

Chcerophyllum temuhim, 123.

Chara-marl, 56, 79.

Charcoal in plant-beds, 59, 86.

Chelidonium majus, 107,

Cherry, 70, 114.

Chestnut, 16, 146.

Chrysanthemum Leucanthemum, 128,

129.
segetum, 32, 128.

Cicuta virosa, 122.
Cladium Mariscus, 81,

164.

Index.

183

Clarke, C. B., 80, 16 c;.

Clay ironstone, plants in, 50, 74.

Clematis V it alba, 91, 100.

Climate and Elevation. 13.

Climatic changes, 2, 3, 7, 8, 20, 21, 33-

47. 97-
Climatic conditions, relations of plants

to, 10-13, 18, 70, 71.
Close y Garey, Isle of iVIan, 57, 61, 62.
Clyde Beds, 59, 69, 70, 71.
Cnicus lanceolatus, 129.

palustris, 130.

Coast-lines, cutting back of, 37.

Cold and warm climatic waves, 2, 3, 7,

8, 20, 21, 33-47, 97.
Competition among plants, 4, 11, 21,

98.
Composites, distribution and dispersal

of, 21, 26.
Conopodium denudatum, 123.
Continent, connection of Britain with

the, 97.
Continental climate, 18,

flora, 8, 18, 21, 25, 99.

Convolvulus, difficulty of dispersal of,

28.
Coralline Crag, 34, 35.
Corbicula fluminalis, jx.
Corfe Castle, Blashenwell near, 53, 59.
Cork, Spiranthes at, 19.
Corn-flower, 87, 130.
Corn Marigold, 32, 87.
Cornus sanguinea, 9, 125, 126.

suecica, 125.

Cornwall, 3, 4, 14, 15, 17, 18, 24, 27,

40-42, 45.
Correlation of plant-bearing deposits,

52-54.
Corstorphine, Edinburgh, 53, 62, 65,

68, 92.
Corton, Suffolk, 62, 63.
Corydalis fabacea, 82.
Corylus Avellana, 66, 84, 145.
Cowden Glen, 63, 86, 98, 106, 132.
Craig, R., 80, 84, 132, 145.
Crambe maritima, 13.
CratcBgus Oxyacantha, 26, 70, 84,

119.
Crepis virens, 130.
CrianlaTich, Perthshire, 53, 64.
CroU, Dr., 44.
Cromer, 32, 64.

Cromer Forest-bed. i, 35, 36, 39,50-52,
56, 58, 59, 62, 63, 64, 74, 82-86, 88,
89. 93. 99. 121, 171-179.

Cross-fertilisation by wind-borne pollen
from the Continent, 12.

Crossness, Essex, 64, 65, 93.

Cultivated plants, n, 46, 53, 89, 137.

Cytisus scoparius, var. prostratus, 4,

Daheocea polifolia, 18.

Dakyns, J. R., 64.

Darwin, C, on dispersal of seeds, 25, 31.

Denmark, fossil plants of, 53.

Derbyshire, 55.

Derwent, gravels of the, 55.

Desert plants, 14, 22, 44.

Deserts, barriers caused by, 22, 28.

Deuben, Saxony, 52, 65.

Devon, 3, 14, 18, 21, 42, 45, 53. 59, 60.

Dewlish, 36.

Dew-ponds, plants found in, 32.

Digestion of soft seeds, 27, 28, 30, 31.

Dingle Promontory, plants of the, 4, 5.

Dispersal, means of, 2, 4, 6, 13, 14, 20-

32, 97.
Distribution of British plants, 2-19.
Dixon, H. N., 83.
Dorset, 16, 18, 24, 36, 53, 59, 86, 96,

99.
Doves, acorns carried by, 30.
Drainage of swamps, 14.
Dronachy, 62, 65, 66.
Drope, Glamorgan, 66.
Dry as octopetala, loo, 116.
Dry-soil plants, fossil, 50, 85, 94.
Dunlop, R., 54.
Dursley, 66.

Early Glacial flora, 52, 171-179.

East Tarbet, 59.

Edinburgh, 52, 53, 60, 62, 65, 68, 69,

70, 72-74, 80, 86, 87, 92, III.
Eleocharis acicularis, 160.

palustris, 161.

Elevation of the sea-bed, 7, 37, 46.

, relation to temperature of, 13.

Elie, Fife, 66, 67.

Elm, II, 59, 66, 69, 74, 142.

1 84

Index.

Elwes, J. W. , 90, 146.

Evtpetrum nigrutn, 150.

Emys lutaria, 83.

Endsleigh Street, London, 67.

English Channel, former temperature of

the, 41, 46.
Environment, adaptation to, 2, 4.
Epilobium, 26
Equisetum hyemale, 169.

limo\um, 169.

palustre, 76, 168.

Erica, 15, 18, 22-27.
Eriocaulon septangulare, 19.
Eriophorum angustifolium, 164.

vaginatum, 164.

Erodium, 24.

Erratics on the Sussex coast, 7, 41, 45,

52, 88, 94.
Essex, 16, 52, 55, 64, 65, 71, 92,

93-
Euonymus, 28.

Enpatorium cannabinum, 127, 128.
Euphorbia amygdaloides, 85, 142.

Helioscopia, 141.

, sea-coast, 18.

Evolution, 5, 12.

Exotic plants in Britain, 95, 98-100.
Extermination by climatic changes, 2,
8, 40.

by drainage, 14.

of non-seeding plants.

Fagus sylvatica, 28, 69, 146.

Fahrenkrug in Holstein, 52, 67, 75.

Faskine, Lanark, 68.

Fenland, 14, 69.

Fertilisation by wind-borne pollen from

the Continent, 12.
Fife, 65, 66, 67.
Fig-tree, 16.

Fillyside, near Edinburgh, 69.
Finland, Trapa nutans in, 122.
Fish, dispersal of freshwater, 7.
Flag, 28, 153.
Flax, 66, 86, 89, iii.
Flintshire, 60.
Floating seeds, 20, 26, 28.
Folliculites, see Stratiotes.
Fool's Parsley, 66, 89, 124.

Forbes, Edward, on distribution of the
existing fauna and flora of Britain,

2, 3-

Forest plants, 15-17, 19, 94.

France, lowland flora of, 13.

Fraxinus excelsior, 24, 28, 69, 82, 133.

Fruits, deficient collections of recent, i,
98.

how dispersed, 2, 4, 6, 13, 14,

20-32, 97.

poisonous, 31

succulent, 20, 26.

tendency to divide into one-
seeded portions, 9.

Fruits, winged, 24, 26.

Fumaria officinalis, 86, 107.

Galeopsis Tetrahit, 93, 137.
Gales, effects of, 12, 23.
Galium Aparine, 127.

boreale, 126.

palustre, t.2j.

uliginosum, 127.

Galway, 19.

Garvel Park, Greenock, 70.
Gayfield, Edinburgh, 70, 92.
Geikie, Prof. J., 44, 63, 72, 86, 106.
Geographical distribution of British

plants, 2-8, 10, 13-19, 25-30, 97-

100
Geological record, imperfection of the,

3. 97. 98.
Geological Survey, work of the, 1-4.
Gepp. A., 96.
Geranium, 23.

columbinum, 82, iii.

Germany, Pleistocene plants of, 8,

51. 52. 54, 65, 67, 71. 72, 75.

81, 82.
Glacial Epochs, recurring, 43, 44.
Glaciation of England, 3, 33, 36-45.

Hampshire, 41.

Ireland, 3, 40, 42.

Sussex, 7, 41, 45, 52, 88,

91. 94-

Wales, 40.

Glamorgan. 46, 57, 66, 100.
Glaucium flavum, 22.
Glaux maritima, 133.
Gloucestershire, 66.

Index.

185

Gorse, 22, 23.

Grass-seeds, dispersal of, 71, 87, 167.
Grays, Essex, 52, 71.
Green, Prof. A. H., 85.
Greenock, 70, 71.
Gregarious plants, 15, 22, 23.
Grossen-Bornholt, Holstein, 72.
Griinenthal, Holstein, 52, 71, 75.

Habitat of plants, 4, 10, 13.

Hackney, 88.

Hailes, near Edinburgh, 52, 53, 60, 62,

65, 68, 70, 72-74, 80, 86, 92.
Hampshire, 16, 32, 34, 41, 52, 88-92,

94. 95. 98, 100.
Hanover, 52, 75.
Happisburgh, 74, 106.
Hartstongue, 66, 168.
Hawthorn, 26, 58, 70, 84, 119.
Hazel, 66, 84, 145.
Heaths, 15. 18, 22-27.
Hedera Helix, 26, 28, 31, 61, 65, 71,
125.

Heer, Dr. O., 59, 147, 152.

Henbane, 24.

Henderson, J., 86.
Heracleuvi Sphondylium, 125.

Hertfordshire, 52, 74, 75.

Hicks, Dr. H., 67.

Hieracium Pilosella, 130.

Hippophae rhamnoides, 141.

Hippopotamus, 54, 55.

Hippuris vulgaris, 120.

Historical method in botany, 3, 6.

Hitchin, 52. 74, 75.

Holderness, 53, 60, 75-77. 79-8i. 87.

Holly, 26, 72, 75, 112.

Holmpton, 75.

Holstein, 52, 67, 71, 72, 75.

Honerdingen, Hanover, 52, 75.

Hope and Fox, Messrs., 89.

Hornbeam, 16, 24, 26, 75, 144.

Horned poppy, 22.

Hornsea, Yorks, 76, 77.

Horse chestnut, ii, 16.

Hoxne, 44, 52, 53, 71, 72, 74, 'jt, 78, 90.

Hoy, Orkney, 29, 30.

Human agency, flora modified by, 11,
12, 14-17. 19-

Hydrobia marginata, 83.

Hydrocotyle vulgaris, 122.
Hyoscyamus, 24.
Hypericum elodes, 11 1.

perforatum, no.

quadrangulum, no.

Iberian plants in Britain, 2-5, 14, 15,

17, 18, 24, 27, 40, 42.
Ice, floating, in the English Channel, 7,

41. 43. 45. 52, 88, 91.
Ice-sheets, extermination by, 3, 40.
lies. Aquifolium, 26, 72, 75, 112.
Imperfections of the geological record,

3. 97. 98.
Insular floras, 8, 12, 33, 97.
Intercrossing of British with continental

plants, 12.
Interglacial periods, 36, 38, 42-44, 52,

53. 63, 68, 72, 75, ^^, 78, 81, 83,

84, 86, 88, 91, 92, 94, 95, 98, 106,

171-179.
Introduced plants, 11, 12, 14-16, 19, 31,

32, 38, 46, S3, 73, 86, 87, 89, 93,

98, 137, 141.
Ireland, Alpine plants of, 13.

American plants in, 19.

fossil plants of, 79.

Lusitanian flora in, 2-5, 14, 15,

17, 18, 26, 27, 40, 42.
Iris, 28, 82, 153.
Isle of Man, 56, 57, 61, 62, 79, 80.

Wight, 41.

Isoetes lacustris, 169.

Isolated ponds, plants of, 14, 28, 32.

Ivy, 26, 28, 31, 61, 65, 71, 125.

Juglans, 75, 76.

/uncus, 22, 26, 66, 69, 87.

Juniperus communis, 76, 151.

Kamtschatka, Spiranthes Romanzo-

viana in, 19.
Keilhack, Dr. K., 81, 109, in, 113,

122, 131, 145, 148, 169.
Kelsey Hill, Yorks, 79.
Kerry, Iberian plants of, 4, 5, 19.

i86

Index.

Kessingland, 85, 86.

Kilmaurs, 79.

Kirk Michael, 80,

Kirmington, 80, 81.

Klinge bei Cottbus, 52, 68, 75, 81.

Lacustrine deposits, plants of, 49.

Laffan, G. B., 93.

Lakes, plants of, 14, 28, 32,

Lamplugh, G. W. , 80.

Land-connection not necessary, 31.

Lanark, 54, 68.

Langley, E. M. , on birds and seeds, 31.

Lapsana communis, 130.

Larch, 39, 82,

Large-seeded plants, how dispersed, 18,

26-31, 99.
Larix, 39, 82.
Lastrcea Thelypteris, 168.
Late Glacial, 53, 92, 94, 171-179.
Lauenburg an der Elbe, 52, 68, 75, 81,

82.
Leeson, Dr. J. R., 93.
Lemmings at Salisbury, 21, 42.
Leontodon azitumnalis, 131.
Lepidurus glacialis, 56, 57.
Lesser Spearwort, 4, 103.
Ligustrum, 28.

Liliacece, distribution of the, 17,
Lime-tree, introduction of the, 16.
Limestone plants, 17, 22, 95.
Lincolnshire, 61, 80, 81.
Linum, 66, 86, 89, iii.
Littorella lacustris, 137.
Local conditions, 4, 7, 10, 13-17.
Locusts destroying plants, 12.
Lodmoor, near Weymouth, 96, io6.
Loess, 44.

Loiseleurla procumbens, 100, 133.
London, 53, 54, 67, 88, 91.
Lowland flora, 13.
Lowlands, fossil Arctic plants of the,

20, 21, 100.
Lusitanian plants in Britain, 2-5, 14, 15,

17, 18, 24-27, 40, 42.
Lychnis alba, 108.

diuma, 108.

Flos-cuculi, 108.

Lycopus europcsus, 135, 136.
Lysimachia Numtmdaria, 82.

Lyell, A. H., 51, 89.

Lyell, Sir C, on dispersal of seeds, 31.

Malva, 28.

Mammals carrying seeds, 26-28, 31,

97-
Mammoth, 54, 67, 71, 76, 80, 81.
Man, first appearance of, 37, 38.
flora modified by, 11, 12, 14-17,

19.
Maple, Lesser, 24, 28, 68, 85, 113.

South European, 98, 99, 113.

Sycamore, 12, 16, 24, 28.

Maritime Pine, 12, 99.

plants, 13, 14, 18, 50, 94.

Marsh plants, 14, 15, 19, 22, 94,
Matricaria inodora, 129.
Maw, G., 60.
Meadow plants, 15, 21.
Mediterranean region, plants of the, 99,

113, 160.
Megaceros-marl, 56, 61, 76, 79.
Mentha aquatica, 135.
Menyanthes trifoliata, 133, 134.
Mercurialis perennis, 142.
Middlesex, 53, 54, 67, 88, 91, 93.
Migration of mammals, 27.
Miller, S. H., 69.
Miocene flora, 34, 36.
Mitten, W., on Cytisus scoparius, 4.
Moisture, relation to plant-life of, 10-

13-
MoUusca, distribution of land and

freshwater, 6, 7, 16, 17, 19.
Montia fentana, no.
Montrose, 133.
Morainic deposits as evidence of

climate, 7.
Morgan, Prof. C. Lloyd, on instincts of

young birds, 31.
Morris, Prof. J., 61.
Mountain Ash, 6r, 119.
Mountain plants in Britain, 2, 3 13,

14, 25-27, 40, 53.
Mundesley, 82, 83, 84.
Musk Ox, 42.
Myosotis sylvatica, 134.
Myrica Gale, 143,
Myriophyllum alternifoliutn, 121.
spicattim, 121.

Index.

187

Najasjlexilis, 75, 76, 159.

graminea, 53, 95, 99, 160.

marina, 46, 57, 76, 81, 86, 100,

159, 160.
minor, 52, 53, 86, 95, 99, 100,

160.
Nathorst, Prof, A. G., 51, 59, 60, 65,

81, 82, 109, III, 112, 119, 120, 132,

139, 141, T43. 147. 149. 169.
Natural seJejtion in insular floras, 12, 21.
Nehring, Prof. A., 51, 81, 153.
Neolithic Period, 38, 45, 46, 53, 58-

71, 76, 79, 86, 90, 92, 93, 96, 98,

106, III, 171-179.
New Forest, 16, 32.
Norfolk, I, 14, 15, 18, 27, 32, 35, 36, 39,

50-52, 56-59, 64, 74, 82-85. 88,

89, 93, 94. 97, TOO.

Northampton, 83,

North Sea Canal, 72.

North Sea, freezing of the, 39.

Norway, horse-chestnut in, 11.

Nuphar luteum, 28, 68, 72, 75, 81, 83,

105.
Nymphcea alba, 28, 68, 72, 75, 81, 82,

96, 106.

Oak, 23, 25, 26, 28-30, 46, 53, 57, 58,

100, 145, 146.
Oceanic islands, plants of, 8, 12, 33.
CEnantheJistulosa, 123.

Lachenalii, 123.

Phellandrium, 124.

Opium Poppy, 63, 98, 1 06.
Orkney, seedlmg oak in, 29, 30.
Osmunda rigalis, 91, 96, 168.
Ostend, Norfolk, 56, 83, 84.
Outliers in botany, 19, 22, 23, 40.
Overstrand, 84,
Overtoun, Ayrshire, 84.
Oxalis Acetosella, 23, 112.
Oxford, 84, 85.
Oxyria digyna, 100, 139.

Pakefield, 85, 86.
Palaearctic flora, 7.

Palaeolithic man, 38, 45, 74, ^jj, 78, 85,
88, 91, 93, 100.

Pjpaver Argemene, io5.
sovmiferum, 63,

106.

Paradoxocarpus^ see Stratiotes.
Parkstone, Dorset, 86,
Pear, 65, 119,
Peat-mosses, Oaks in, loo.

Pine in, 16, 86, 100.

plants of, 14, 15, 22, 49,

79-
Pedicu la ris pa lustris, 135.
Pengelly, W., 59.
Peniland Firth, acorns carried across,

30-
Perennial plants, conditions needed by,

II, 12.

in oceanic i-slands, 12.

Perthshire, 53, 64.

Peucedanum. palustre, 124,

Phillips, Prof. J., 76.

Phraginiies communis, 57, 80, 167.

Physical geography, past changes in, 2,

7, 8, 27, 28, 33-47.
Picea excelsa, 52, 64, 82, 8j, 99, 151,

152.
Picris hieraciodes, 85, 130.
Piette, Ed,, on cultivated forms of

Prumis, 114.
Pinguecula grandifiora, 5.

vulgaris, 5.

Pinus maritima, 12, 99.

montana, 152.

sylvestris, 16, 24, 26, 53, 57, 64,

82, 86, 99, 100, 152, 153.
Plantations, influence of, 16.
Pliocene flora, 34-36, 97.
Platafius, 75, 76.
Pods, ejection of seeds from, 23.
Poisonous fruits, 31.
Pollen, effects of wind-borne, on island

plants, 12.
Polygonum amphibium, 139.

■ Aviculare, 138.

Hydropiper, 138.

la pathi folium, 139.

Persicaria, 139.

viviparum, 139.

Polystichum, 76, 81.

Ponds, plaats of isolated, 14, 28,

32-
Poplar, 61, 76, 81, 150.
Poppy, 24, 32, 63, 98.
Populus canescens, 71, 150.

i88

Index.

Populus tremula, 6i, 76, 81, 150.
Postglacial period, length of the, 25.
Potamogeton crispus, 156, 157.

densus, 157.

filiformis, 158.

heterophyllus , 155.

lucens, 156.

nutans, 155.

obtusifolius, 157.

pectinatus, 158.

— perfoliatus, 156.

prcelongus, 156.

— pusillus, 157.

rufescens, 155.

trichoides, 100, 157, 158.

Zzszz. 155.

Potentilla Comarum, 117

Tormentilla, 116, 117.

Poterium officinale, 117, 118.
Potoni^, H., 153.

Prairies, plants of, 15, 21, 22.
Preglacial flora, i, 2, 35, 36, 39, 50-52,
58, 62, 63, 64, 74, 82, 83, 84-86, 88,

89. 93. 99. 171-179-
Prestwich, Sir J., 71, 88, 145.
Privet, 28.

Provinces, botanical, 10.
Prunella vulgaris, 136.
Prunus Avium, 114.

domestica, 114.

Padus, 114.

spinosa, 113.

Prussia, 52, 68, 75, 81.

Pteris aquilina, 168.

Pyrenean plants in Britain, 2-5, 14, 15,

17, 18, 24-27, 40, <2.
Pyrus Aria, 85, 119.

Aucuparia, 61, 119.

communis, 65, 119.

Mains, 28.

torminalis, 75, 78, 118.

Quercus Robur, 23, 26, 28-30, 46, loo,
145, 146.

Rabbit, bones of, 66.
Races and subspecies, 4, 5,
Rainfall, 10, 11, 14.

Ranunculus acris, 103.

aquatilis, 4, 26, lOl.

bnlbosus, 104.

Ficaria, 105.

Flammula, 4, 103.

hederaceus, 102.

Lingua, 103.

parviflorus, 105.

re pens, 104.

Sardous, 104.

sceleratus, 102.

Red Crag, 35.

Redhall, near Edinburgh, 53, 60, 86, 87,

III.
Reeds, 57, 80, 167.
Reid, Miss M. A., 66.
Reindeer, 20, 81, 93, 94.
Renfrewshire, 63, 70, 86, 98, 106, 132.
Rhamnus Frangula, 112,
Rhine, old course of the, 35, 97.
Rhinoceros, 54, 55, 81, 94.
Rhododendron, 12.
Ridley, H. N.. 146.
Ripening of seeds, period of, 2, 4.
Robertson, D. , 59, 70.
Rock Doves, acorns carried by, 30.
Romans, plants introduced by the, 16,

51, 89, 103, 112.
Roman Period, 51, 52, 89, 93, 96, 171-

179.
Rooks carrying acorns, 29, 30.
Rosa, 118.
Rubus ccBsius, 116.

fruticosus, 26, 115, 116.

IdcBus, 59, 115.

saxatilis, 116.

Rumex Ace to sella, 141.

conglotneratus, 139, 140.

crispus, 140, 141.

Hydro lapathtim, 141.

maritivius, 140.

obtusifolius, 140.

Ruppia maritima, 158.
Ruscus aculeatus, 11, 12.
Rushes, 22, 26, 66, 69, 87.
Russia, 163, 166.

Sagittaria sagittifolia, 154.

Saint Cross, Suffolk, 90.

Salisbury, Arctic mammals of, 21, 42, 45.

Index,

189

Salix Arbuscula, i48> 149.

— \ aurita, 147.

Caprea, 147, 148.

cinerea, 147.

furbacea, 100, 149.

lanata, 148.

Myrsinites, 100, 149.

nigricans, 148.

pentandra, 147.

phylicifolia, 148.

Polaris, 52, 53, 99, 149.

repens, 148.

reticulata, 100, 149.

Sambucus nigra, 126.
Sand-dune plants, 13, 14.
Sand le Meer, 87.
Sandy heaths, plants of, 15.
Saporta, G. de, 152.
Saxifraga aizoides, 120.

Geum, 5.

Hirculus, 120.

oppositifolia, 119.

umbrosa, 4, 5.

Saxony, 52, 65.

Scabiosa succisa, 127.

Scandinavia, alpine plants of, 13, 99.

Scheuchzeria palustris, 154.

Scilla nutans, 24.

Scilly Islands, 42.

Scirpus ccBspitosus, 162.

Jluitans, 162.

lacustris, 162, 163.

maritimus, 57, 163.

faucfjlorus, 161, 162.

setaceus, 162.

sylvaticus, 163.

Scolopendrium vulgare, 168.
Scotland, alpine plants of, 13.
Scott, Thos., 66, 70, 71, 86, 135.
Sea-coast plants, 13, 14, 18, 50, 94.
Sea, dispersal of plants by the, 13.
Sea-kale, 13.
Sedentary plants, 21.
Seeds, deficient collections of, i, 98.

definition of the term, 8, 9.

how preserved in a fossil state,

49-51-
how transported, 2, 4, 6, 13, 14,

20-32, 97.

period of ripening, 2, 4,

ripe, essential to the plant, 11,12.

size of, 13, 18, 21, 24, 26, 27, 99.

Seeds, winged, 24, 26.
Selsey, 52, 88, 91, 94, 95, 98.

erratics, 7, 41, 45, 52, 88, 91.

Senecio aquaticus, 129.

sylvaticus, 129.

Sheep, bones of, 66,
Shacklewell, 88.

Shingle-beaches, plants of, 13, 14,
Shore, T. W., 90, 146.

Shore-ice in the English Channel, 7, 41,

45, 52, 88, 91.
Sidestrand, 88, 89.
Silchester, 51, 89.
Silene maritima, 108.
Sisyrinchium angustifolium, 19.
Sium latifolium, 123.
Skertchly, S. B. ]., 69.
Skye, 19.

Slow spreading of plants, 24, 25.
Small-seeded plants, 13, 18, 21, 26,

27.
Smith, W. G , 91, loo, 112.
Snails, distribution of, 6, 7, 16, 17, ig.
Solanum Dulcamara, 134.
Sonchus arvetisis, 131.
Southampton, 90.
South Downs, 29, 32, 45.
Southelmham, 90.
South Wales, 40, 46, 53, 57, 58, 66.
Spanish Chestnut, 16, 146.
Sparganium minitnum, 76, 154.

ramosum, 153.

simplex, 76, 154.

Specialisation of the British flora, 20,

21, 25, 26.
Spergula arvense, no.
Spermophilus with Arctic plants, 83.
Spindle-tree, 28.
Spircea Ulmaria, 114, 115.
Spiranthes Romanzoviana, 19.
Sporadic appearance of plants, 13, 16.
Spurge, 18, 23, 85, 141, 142.
Spurrell, F. C. J., 55, 64, 92.
Stachys atvensis, 136.

palustris, 136.

sylvaticus, 136.

Stellaria aquatica, 108.

graminea, 109.

Holostea, 109.

media, 109.

' — uliginosa, 109.

Stoke Newington, 91.

190

Index.

Stone, Hants, 52, 88, 91, 92, 94, 95,

98.
Storms, effects of, 12, 23-
Storrie, J., 57.
Strahan, A., 57, 60.
Strait of Dover, 30, 36, 39, 46, 47, 97.

99.
Stratigraphical evidence of age, 3, 48,

53.
Stratiotes aloides, 81, 89, 153.
Struggle for existence, 21 .
Submerged forests, 30, 46, 57, 58,

69, 76, 93, 100, 145, 147, 151,

152.
Submergence of the land, 7, 34, 37, 39,

43, 46.
Subspecies and vaiieties, 4, 5, 8, 102.
Succulent fruits, dispersal of, 20, 26.
Suffolk, 14, 15, 18, 27, 36, 44, 52,

62, 72, 74, 'jj, 78, 85, 86, 90,

100.
Sunshine, relation to plant-life of, 10-13,

18, 24, SI-
Sussex, 29, 32, 41, 43-45, 88, 94-96, 98-

100
Sweden, fossil plants of, 8, 51, 53, 92,

99, T2I.

Switzerland, Opium Poppy in, 98, 106.
Sycamore, 12, 16, 24, 28.

Tanacetum vulgare, 129.

Taraxacum officinale, 131.

Taxus baccata, 151.

Temperature, relation to plant-life of,
10-T3.

Tertiary floras, 34-36.

Thalirtrum fiaviun, loi.

minus, loi.

Thames, alluvial deposits of the, 53, 55,
92. 93

Thlaspi arvensc, 107.

Thymus Serpylluvi, 136.

Tilbury, 92, 93.

Tilia enr>p(Ea, 16, 75, iii.

intermedia, 75, iii.

platyphyllos, 75, 81, 82, in.

Trapa natans, 52, 82, 83, 85, 89, 99,
121, 122.

Tropical floras contrasted with Tem-
perate floras, 20, 21.

Trimingham, 93.

Tufa, plants in, 51, 59-61, 66.

Tussilago Farfara, 129.

Twickenham, 93, 137.

Tylor, Dr. A., 71,

Typha, 26, 76.

Ulex, 22, 23.

Ulmus campestris, 11, 142.

montana, 66, 69, 74, 142.

Umbelliferous plants, dispersal of, 23,

24.
Upland flora, 13, 14.
Urtica dioica, 142.

Vaccinium Myrtillus, 132.

Oxycoccos, 82, 131.

uliginosum, 132.

Vitis-Idcea, 132.

Valeriana officinalis, 57, 127.
Valerianella olitoria, 127.
Varietal forms, 4, 5, 8, 102.
Verbascutn Thaspus, 134.
Vernitsa, 163, 166.
Viburnum Lantana, xib.

Opulus, 82, 126.

Vine, 16, 91, 112.
Viola palustris, 107.
Viscum album, 141.
Vitis vinifera, 16, 91, 112.

Wales, Alpine plants of, 13.

foss'l plants of, 46, 53, 57, 58, 60,

61, 66.
Wallace, A. R., 99.
Walnut, 75, 76.

Ward, Prof. M., 65, 119, 133, 146.
Warm and cold waves, 2, 3, 7, 8, 20,

21. 33-47. 97-
Water-chestnut, 52, 82, 83, 85, 89, 99,

121, 122.

crow.f^oot, 4, 26.

lily, 28.

milfoil, 26,

tortoise, 83.

Waterlogged soils, 15.

Index.

191

Weber, Dr. Carl, 51, 67, 71, 72, 75, 81,
loi, 106, III, 112, 113, 122, 127,
131, 132, 133, 143, 145-148, 150-
152, 154-156, 159, 160, 164-166.

Weeds of cultivation, it, 31, 32, 38, 53,
73, 86, 87, 89, 93, 98, 137, 141.

West Runton, 94

West Wittering, 50, 52, 88, 91, 94-96,
99.

Weymouth, 96, 106.

Wheat, II.

Whi taker, W., 90.

Wild hyacinth, 24.

Williams, W., 79.

Willow-herb, 26.

Willows, Arctic, 14, 57; 58, 59, 62, 64,
65, 68, 70, 73, ^^, 78, 84, 99, 100,
149.

dispersal of seeds of, 26.

Wiltshire, 21, 42, 45.

Wind, dispersal of seeds by, 24, 26.

Winged seeds, 24, 26.

Wolvercote, 84, 85.

Woodland plants, 15-17, 19, 94.

Wood-pigeons, dispersal of acorns and
beans by, 30.

Woods, importance of examining an-
cient, 17,

Wood-sorrel, 23.

Woolwich, 53, 93.

Yew, 151.

Yorkshire, 53, 60, 75, 76, 'j'j, 79, 87.

Zannichellia palusiris, 158, 159.

pedunculata, 159.

7.0 st era marina, 159.

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