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






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


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. 


Chap. Page. 


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 .... 










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 


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 

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 

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 

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- 

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 

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. 



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 

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 

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- 


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. 



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 : — 



in Britain. 

in Europe. 

Targe seeds 



Small seeds 

... 17-6 


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 

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 

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 

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 

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. 


Origin of the British Flora, 

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


Abundance of minute seeds 
(Heaths, Rushes, Saxi- 
frages, Caryophyllacese, 

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

Edible fruits with hard 
stones (Blackberry, Haw- 
thorn, Holly, Arbutus, 

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- 

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

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). 



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- 


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 

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 

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 

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 

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 

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. 



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 

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 


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- 

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 

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. 


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

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. 


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. 


Hippuris vulgaris. 

Carex dioica. 

Apium nodiflorum. 




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 

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. 

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. 


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


(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. 

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

Deposits containing Fossil Plants 


Thalictrum flavum. 


Nuphar luteum. 

Hydrocotyle vulgaris. 


Solanum Dulcamara. 

Atriplex patula. 

Alnus glutinosa. 

Ceratophyllum demersum. 

Stratiotes aloides. 
Sparganium ramosum. 
Potamogeton lucens. 


Zannichellia palustris. 
Scirpus pauciflorus. 


Eriophorum angustifolium. 

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 

Ranunculus aquatilis. 


Papaver somniferum. 

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. 



Scirpus lacustris. 


Carex rostrata. 
Isoetes lacustris. 

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. 


(l. — Lower Peat. u. — Upper Peat.) 

Ranunculus sceleratus 

repens . 

Oxalis Acetosella 
Ilex Aquifolium 
Acer campestre 
Prunus communis 


Avium , 

Rubus fruticosus 

*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. 



Polygonum viviparum. 
Salix herbacea. 



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. 


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. 


Juncus (i*) 


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 

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. 


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

Ranunculus aquatilis. 


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. 
Carex, several species, 
Phragmites (?). 

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. 


Stellaria media. 


Potentilla Tormentilla. 

Hippuris vulgaris. 

Myriophyllum spicatum. 

Polygonum aviculare. 



Zannichellia palustris. 
Eleocharis palustris. 
Carex dioica. 

2 sp. 


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 


(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. 


(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 


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 (?). 


Polygonum Persicaria. 

Viola palustris. 

Salix polaris. 

Prunus Avium. 


Rubus Idaeus. 


Crataegus Oxyacantha. 

Potamogeton crispus. 





Sonchus arvensis. 


Menyanthes trifoliata. 


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. 

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. 


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

Thalictrum minus (?). 
Ranunculus aquatilis. 


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. 



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. 


Lingua (?). 


Caltha palustris. 
Viola palustris. 
Lychnis diurna. 


Stellaria media. 


Montia fontana. 
Linum. i 

Oxalis Acetosella. 
Prunus spinosa. 


Spiraea Ulmaria. 
Rubus Idaius. 


Potentilla Tormentilla (?). 


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. 

Happisburgh, Norfolk. 

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

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. 


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

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

2 sp. 

Najas marina. 
Scirpus lacustris. 



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. 


(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 

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



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. 


Origin of the British Flora. 

Corylus Avellana. 

Quercus sessiliflora. 

Fagus sylvatica. 


Populus tremula. 

Platanus (?). 

Juglans regia (?). 

Empetrum nigrum. 

Ceratophyllum submersum. 



Sparganium minimum. 


Potamogeton natans. 

cf. polygon i- 

Potamogeton rufescens. 

cf. colorata. 


cf. praelonga. 



Potamogeton compressa. 




cf. trichoides. 

— manna. 

Najas major. 


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


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. 

Pinus sylvestris. 



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. 


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. 


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- 

Eupatorium cannabinum. 

Bidens tripartita. 

Taraxacum officinale. 

Menyanthes trifoliata. 

Lycopus europaeus. 
Ajuga reptans. 
Rumex maritimus. 

crispus Q). 

Urtica dioica ( } ) [one 


Betula nana. 

Alnus glutinosa [perhaps 

Carpinus Betuius [deriv- 

Salix myrsinites. 



Ceratophyllum demersum. 
Taxus baccata [ deriv- 

Sparganium ramosum. 
Alisma Plantago. 
Potamogeton rufescens. 

78 Origin of the British Flora. 

Potamogeton pusillus. Scirpus lacustris. 

trichoides. Blysmus rufus. 

pectinatus. Carex incurva (?). 

Scirpus pauciflorus. Chara. 

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 

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 

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

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- 

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. 


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. 


Nymphaea alba. 


Brasenia ovulum. 

Caprea (?). 

Tilia platyphyllos (J). 

Populus tremula. 

Ilex aquifolium. 

Ceratophyllum submersum. 

Acer campestre. 


Comarum palustre. 

Taxus baccata. 

Hippuris vulgaris. 

Pinus sylvestris. 


Picea excelsa. 

Galium palustre (?). 

Stratiotes aloides. 

Vaccinium Oxycoccos. 

Echinodorus ranunculoides(?) 

Menyanthes trifoliata. 

Potamogeton natans. 

Betula verucosa. 

Najas marina. 


Scirpus lacustris. 


pauciflorus (?). 

Carpinus Betulus. 

Cladium Mariscus. 

Corylus Avellana. 

Carex, several sp. 


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, 


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. 


(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. 


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. 


(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. 

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


(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. 


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

Ranunculus aquatilis. 


Potentilla Tormentilla. 


Hippuris vulgaris. 

Heracleum Sphondylium. 
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. 


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

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


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. 


Oxalis Acetosella. 

Spiraea Ulmaria. 
Rubus Idseus. 


Potentilla Tormentilla. 


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. 


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. 


Rumex obtusifolius. 


Euphorbia Helioscopia. 
Alnus glutinosa. 
Quercus Robur. 

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


Eleocharis palustris. 
Scirpus pauciflorus. 


Carex dioica. 





* Holcus lanatus. 

* 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. 

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. 

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

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. 


(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 

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 (?). 


(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.) 


Betula. (Bark and wood.) 

Alnus glutinosa. (Leaves and catkins.) 




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. 


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

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, 

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. 

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. 

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, 


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. 








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

Flos-cuculi. • 

Stellaria aquatica. 



Montia fontana. 
Hypericum perforatum. 


Ilex aquifolium. 
Rhamnus Frangula. 

Prunus spinosa. 


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

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


Chaerophyllum temulum. 
Anthriscus sylvestris. 
CEnanthe fistulosa. 


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


Valerianella olitoria. 
Scabiosa succisa. 


Origin of the British Flora. 

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


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. 

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


lucens (?). 



Ruppia maritima. 
Zannichellia pedunculata. 
Najas minor. 



Mercurialis perennis. 
Betula alba. 

Eleocharis acicularis. 


Scirpus lacustris. 
Carex muricata. 





(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- 

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

Potamogeton natans. 



Scirpus maritimus (?). 


Phragmites communis. 

Osmunda regalis. 



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, 


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 

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 

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, 

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. 

Neolithic and Postglacial : — 

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

Interglacial : — 

West Wittering, Sussex ; Mundesley, Norfolk (old river 

Preglacial (Cromer Forest-bed) : — 

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

io6 Origin of the British Flora. 

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. 


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 


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 

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. 

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, 

Late Glacial: — 

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

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. 


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, 

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). 


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 

Former Distribution of British Plants. 1 1 3 

Acer campestre, L. 

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 

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. 


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, 

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. 

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). 

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. 


Unknown fossil in Britain. 

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

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. 


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. 

Neolithic :— 

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

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

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. 

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. 

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). 

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- 

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 

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. 

Airdrie, Lanark ; West Wittering, Sussex. 


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. 


Unknown fossil in Britain. 

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

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. 

Roman Period : — 

Silchester, Hampshire. 

Neolithic: — 

Hailes, near Edinburgh; Elie, Fife. 

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

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). 

Unknown fossil in Britain. 

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

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 

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. 

Interglacial: — 

West Wittering, Sussex. 

Neolithic: — 

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


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 

Aster Tripolium, L. 
Interglacial: — 

West Wittering, Sussex. 

Neolithic: — 

Redhall, near Edinburgh. 

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

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


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. 


130 Origin of the British Flora. 

Cnicus palustris, L. 
Neolithic: — 

Hailes, near Edinburgh ; Redhall, near Edinburgh. 

Interglacial: — 

Southelmham, Suffolk (fruits small) ; West Wittering, 


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. 


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. 

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 

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 

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. 

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. 

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. 


Neolithic : — 

Redhall, near Edinburgh ; Megaceros-marls of Central 

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, 

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, 

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. 


OXYRIA digyna, Hill. 
Late Glacial : — 

Corstorphine, near Edinburgh. 

Roman Period : — 

Silchester, Hampshire. 

140 Origin of the British Flora, 

Interglacial : — 

West Wittering, Sussex. 

Late Glacial : — 

Hoxne, Suffolk (bed C). 

Interglacial : — 

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

Early Glacial : — 

Beeston, Norfolk (base of Arctic Freshwater bed). 

Preglacial (Cromer Forest-bed) : — 

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

Neolithic : — 

Redhall, near Edinburgh. 

Late Glacial : — 

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

Interglacial : — 

West Wittering, Sussex ; Endsleigh Street, London. 

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). 

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. 

Unknown fossil in Britain. 

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


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. 


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. 

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 

Interglacial: — 

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

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. 

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. 

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, 

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 

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 

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, 

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 

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, 


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. 

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. 


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). 

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). 


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. 

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 

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. 

Interglacial : — 

West Wittering, Sussex. 

Recorded from the Oak Zone in Gotland (Gunnar 


Unknown fossil in Britain. 

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

Former Distribution of British Plants. 155 

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). 


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). 

Neolithic : — 

Drope, Glamorgan ; Hailes, near Edinburgh. 

Interglacial : — 

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

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

156 Origin of the British Flora. 

Neolithic : — 

Cowden Glen, Renfrewshire. 

Interglacial : — 

West Wittering, Sussex. 

Preglacial (Cromer Forest-bed) : — 

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

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). 

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). 

Neolithic : — 

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

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. 

Interglacial : — 

West Wittering, Sussex. 

Interglacial : — 

Endsleigh Street, London; Southelmham, Suffolk. 

Neolithic : — 

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

Late Glacial : — 

Hoxne, Suffolk (bed C). 

Interglacial : — 

Hoxne, Suffolk (bed D). 

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. 

Neolithic : — 

Cowden Glen, Renfrewshire. 

Late Glacial : — 

Hoxne, Suffolk (bed C). 

Preglacial (Cromer Forest-bed) : — 

Common at most localities — drupes often very large. 

Unknown fossil in Britain. 

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

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 


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. 

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. 


1 62 Origin of the British Flora. 

Preglacial (Cromer Forest-bed): — 

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

Interglacial : — 

Southelmham, Suffolk. 

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

Late Glacial : — 

Dronachy, Fife (?). 

Interglacial : — 

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

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


Neolithic : — 

Hailes, Redhall, and Broughton, near Edinburgh. 

Late Glacial : — 

Hoxne, Suffolk (bed C). 

Interglacial : — 

Hoxne, Suffolk (bed D). 

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 

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. 

Neolithic : — 

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


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 


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 

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 , 

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 


Found in calcareous tufa of doubtful age at Dursley, in 

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). 


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 

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). 



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. 











S M 






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 S 




Time of the British Flora. 


Rubus Idaeus, L. ... 

fruticosus, Z. 

caesius, L. ... 

saxatilis, L. 


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 


















S M 






S I 

S M 


Table showing the Range 


(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 W 

E W 





E S 




Time of the British Flora. 


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 















W S 



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 S 



S I 













Time of the British Flor^a. 


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 S 









E S 











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 M 

E S 








S I 




S I 




Time of the B^'itish Flora. 


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? W 







E W 





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, 

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. 

Regions, plants of the, 39, 99, 


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. 



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, 

segetum, 32, 128. 

Cicuta virosa, 122. 
Cladium Mariscus, 81, 




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, 

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, 

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, 

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, 

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, 

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 


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, 

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, 

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- 

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. 



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, 

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, 

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. 



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, 

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, 

Mammoth, 54, 67, 71, 76, 80, 81. 
Man, first appearance of, 37, 38. 
flora modified by, 11, 12, 14-17, 

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- 

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. 



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, 

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, 

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, 


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, 

Pedicu la ris pa lustris, 135. 
Pengelly, W., 59. 
Peniland Firth, acorns carried across, 

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, 

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, 

Poplar, 61, 76, 81, 150. 
Poppy, 24, 32, 63, 98. 
Populus canescens, 71, 150. 



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, 

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- 

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. 



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, 

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, 

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. 



Stone, Hants, 52, 88, 91, 92, 94, 95, 

Storms, effects of, 12, 23- 
Storrie, J., 57. 
Strahan, A., 57, 60. 
Strait of Dover, 30, 36, 39, 46, 47, 97. 

Stratigraphical evidence of age, 3, 48, 

Stratiotes aloides, 81, 89, 153. 
Struggle for existence, 21 . 
Submerged forests, 30, 46, 57, 58, 

69, 76, 93, 100, 145, 147, 151, 

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, 

Sunshine, relation to plant-life of, 10-13, 

18, 24, SI- 
Sussex, 29, 32, 41, 43-45, 88, 94-96, 98- 

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, 

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, 

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. 



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, 

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, 

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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9^ The origin of the British 

R4 flora