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Adviser m Agricultural Botany

_ University COLLEGE OF NortH WALES

BANGOR P

Bangor
JARVIS & FOSTER, PRINTERS

MCMXIX

Cornell University

Library

The original of this book is in
the Cornell University Library.

There are no known copyright restrictions in
the United States on the use of the text.

http://www.archive.org/details/cu31924003370685

PASTURE STUDIES:

SOME RESULTS

CONTENTS

Introduction

Methods of Examination
Tuer History OF SPECIES IN PASTURES.

Areas Examined -
Table I. Introduction of Species

Table II. Some Particulars of Areas

Seeds- Mixtures

Aggregate Results

Table III. Aggregate Results

Individual Species

Table IV. Seeds- Mixtures and Percentage Frequency
Results for Gramineae and Leguminosae

Table V. Percentage Frequency and Specific
Frequency Results for Natural Orders other
than Gramineae

Summary aod Conclusions

References

16a

166
52
57

INTRODUCTION.

The main object of this publication is to place on record
certain results obtained by the writer from the study of pasture
herbage. A few of the present results and others have already
been embodied in a paper written jointly by Mr. R. G. Stapledon,
M.A., and the present writer, and published in the Journal of

Agricultural Science (18), but the greater part of the results are
now published for the first time.

METHODS OF EXAMINATION.

(a) Percentage Frequency.—This method was originally used
by Armstrong (1), and was adopted by Stapledon (15) in a
modified form. It has been re-defined as “the number of plants
of each species per unit area” expressed in percentages of the
total herbage (18).

This method has been further modified to some extent by
the present writer, as follows :—

(t) The plant unit has been re-defined :

i. For most plants, the plant unit shall consist of any
portion of a plant possessing an independent shoot, which
can be separated from others with at least three rootlets still
attached to it. Ifa portion possesses six rootlets and it can
be separated into two in such a way that each portion
possesses three rootlets, then it counts as two plant units.

ii. The plant unit of Z+ifolium repens shall be, “ any
portion of runner up to three inches in length and bearing
functioning rootlets.”

iii. The plant unit for Vardus stricta shall be as nearly
as possible comparable in size to that of Festuca ovina.?

iv. In a few cases the plant unit used is, “ any individual
tiller or shoot irrespective of roots.” This definition applies to

1 Figures in heavy type and in brackets refer to list of references at the end,

2 This definition was adopted owing to the plant’s peculiar habit. It is
obviously unsatisfactory, and in future work it is intended to bring this plant,
in spite of its peculiar habit, under the general definition (i.)

2 PASTURE STUDIES: SOME RESULTS.

Juncus spp., Carex spp., Luzula spp., Schoenus nigricans, and

plants of a similar habit.

2. A new turf-sampling mesh was introduced.

In 1916, the 6in. x 6in. mesh previously used for taking
specimen turves was partly replaced with a new mesh, r2in. x rin.
This new mesh has since been used exclusively on pastures which
are fairly well grazed, and on such it is quite satisfactory. Where
the herbage is rank, the 6in. x 6in. mesh is still used.

(b) Percentage Area.—This method was also originally used
by Armstrong (1), but the present writer has found it effective
and reliable only in the estimation of the area covered by herbs
which are easily identified, and, in a pasture, grow quite near the
ground. Only results obtained for Z+ifolium repens by this
method will be referred to.

(¢) Specific Frequency.—This is a qualitative method intro-
duced by Stapledon (15) (17). It has been re-defined as “ the
number of times a species (without regard to the number of plants
of that species), occurs per 100 readings within the 6in. x 6in.
mesh (18). This method is exceedingly useful for ascertaining
the distribution of plants which are easily identified in pastures,
but much less satisfactory in the case of grasses. The writer has
used it mostly on young pastures for ascertaining the distribution
of leguminous and other plants which are easily identified 27 stfu.

AREAS EXAMINED.

Up till quite recently, relatively little work had been done to
ascertain to what extent and for how long, species whose seeds
are commonly included in seeds-mixtures used for laying land
down to pasture persist in such pastures.

The present writer commenced his investigations along these
lines in 1913 on land the history of which was well known to him.

All the areas for which results are here given are situated on
the farm of Budloy, near Maenclochog, Pembrokeshire.? This

1 In some respects, these definitions resemble that adopted by Yapp,
Johns and Jones (22).

? A few results obtained elsewhere are given to illustrate certain points,

é

AREAS EXAMINED, 5

farm may be described as a mixed hill farm with no natural first,
nor even second class land. It is about 160 acres in extent, and
svaries in altitude from 5ooft. to g4oft. Less than half the total
area is cultivable.

The soil is of three main types :—

Type i.: Thin, rather light loam, sedentary on shale. Very
,small areas of this type are found between sooft. and 6coft.,
and also between 6ooft. and 7ooft. while the greater part of the
area between 7ooft. and gooft. is of this type.

Type ii: Thin, slightly heavier loam on local drift. This
Idcal drift consists chiefly of shale with some felsitd?/boulders.
This soil is of a silty character, and is very liable to cake on the
surface when rain follows immediately after thorough cultivation
to a fine tilth. Most of the land regularly cultivated and the best
permanent pasture fields are on this type.

Type iii.: Thin, acid peat on felsite rock. Most of the land
above 8ooft. is of this type. All this land is uncultivable, owing
to the thin nature of the peat and the presence of felsite blocks.

The character of the herbage on soil-types i. and ii. will be
most clearly seen from the tables. ‘The most important arable
land weeds on the farm are :—

Agrostts spp.

Chrysanthemum segetum (local),

Galeopsis Tetvahit (local).

Arrhenatherum elatius var. tuberosum (local).

Polygonum Persicaria,

Stellaria media,

Cirsium arvense.

Ranunculus repens.

These chiefly occur on soil-type ii, The land on soil-type i.
has been little cultivated, but on this Rumex Acetosella was a
prominent weed in places.

Soil-type iii, carries a distinct type of herbage which may be
described as a Cal/una heath. Calluna vulgaris is dominant, but
it is accompanied by Arica Tetvalix, Vaccinium Myrtillus, Festuca

4 PASTURE STUDIES: SOME RESULTS.

ovina, Deschampsia flexuosa, eat
Scirpus caespitosus, etc.t No quantitative examination of this
heath herbage has been made.

The cultivation of the land on the farm varies somewhat
according to circumstances. As this seems to be important, full
details will be given for each area examined. The main features
are, that the length of the rotation is usually 10 to 15 years, and
that the land is well cultivated and manured while under cultiva-
tion, but that it very rarely happens that the entire area which is
seeded down to pasture in any one year has been entirely under
a cleaning crop? in the course of that rotation.

The land is always seeded down to grass with a corn cover-
ing crop, usually dredge-corn.3 Since the rotation is a somewhat
irregular one, no land is laid down to grass in some years.

The seeds-mixtures used on the farm have varied considerably
in recent years, but formerly they were of a simple character.
Fortunately, the species included in all seeds-mixtures used from
1883 to 1917, with the exception of those used in 1901, 1902 and
1903,+ are known. During this period 23 seeds-mixtures were
used, and the various species and commercial varieties included
are given in Table I.

TaBLE I.: Showing the Years in which various “seeds ”5 were
included in Seeds-mixtures on Budloy.

1. Lolium perenne: All seeds-mixtures 1883-1917.

2. “Broad Red Clover”®: All seeds-mixtures except in
igi and 1912.

3. Plantago lanceolata: 1884, 1887-91, 1898, 1902 (?),
1905, ’06, ’o9, ‘12.

4. Lolium italicum : 1891, ’97, 98, 1906-77.

5. Trifolium hybridum : 1898 (2), 1905-17.

' Calluna vulgaris is everywhere dominant in the older herbage, but the
abundance of the other species varies in different places according to the slope,
etc, After buming, Cad/na does not become dominant far a few years,

? Potatoes, Mangels, Turnips, etc.

3 A mixture of barley and oats.

4 Some details are known for these years alsa

5 The word ‘‘ seeds” is used throughout in the commercial or common sense,
§ The Commercial names of the different varieties are given,

AREAS EXAMINED. 5

6. Phleum pratense: 1901, ’02, ’08, '11, 713-17.
4. Dactylis glomerata: 1903-1917.
8. “ Cowgrass” (Zrifolium pratense): 1905-08, 1911-13.
9. Medicago lupulina: 1905, ’o6.

10. Festuca elatior: 1908, ’13.

11. Ditto var. pratensis: 1908, 13, ‘17.

12. F. duriuscula: 1908-13.

13. Poa pratensis: 1908, ’16.

14. “White or Dutch Clover”: 1908-13.

15. Anthylis Vulneraria: 1908-17. ,

16. “ Hybrid Cow Clover”: 1909.

17. Festuca ovina: 191l.

18. Cynosurus cristatus : 1911, 16, 717.

19. Anthoxanthum odoratum: 1911.

20. Medicago sativa: 1911, ’12.

21. Cichorium Intybus: 191t, 12, °16, 717.

22. Potertum Sangutsorba: 19tt, ‘12.

23. Carum Petroselinum: 1911, ’12.

24. Arrhenatherum elatius: 1912, '13, ‘17.

25. “Late-flowering Red Clover”: 1912, ’16, ‘17.

26. Alopecurus pratensis: 1913.

27. Poa trivals: 1916, 717.

28. “Wild White Clover”: 1916, ’17.

29. Trifolium dubium: 1916.

30. Achillea Millefolum : 1916, ’17.

The evolution of the seeds-mixture on this particular farm
as shown by these data is seen to have been relatively slow, but
even in the absence of expert advice, it had been proceeding
prior to 1908. After this, the seeds-mixtures were more or less of
an experimental character, although used on the most extensive
scale possible on the farm. The details of the seeds-mixtures
used on Areas 1-8 are given in Table IV., p/éa, except that a few
miscellaneous plants are not included.

1 For these, see under ‘‘ Individual Species,” p. 15 ef seg.
? P Pp. 15 ef seg

6 PASTURE STUDIES: SOME RESULTS.

AREA 1,

Altitude : 650 feet.

Soil: Type ii. (p. 3).

History: In grass 1898-1910; oats 1910; bastard fallow ;
dredge corn 1911 ; oats 1912; bastard fallow ; dredge corn 1913,
and seeded down to permanent pasture with the seeds-mixture
shown in Table IV. The area was well manured with farmyard
manure and phosphates during this period of cultivation, but
was only partly under a cleaning crop.

Hay was taken in 1914, farmyard manure was applied winter
1914-15, and hay again taken in 1915. It has been continuously
grazed since, without further manuring.

AREA 2. Adjoining Area 1.

Altitude: 700 feet.

Soil: Mostly Type ii. (p. 3), but partly of Type i.

History: In grass 1895-1905 without manuring ; hay 1905 ;
oats 1906; bastard fallow; dredge-corn 1907; bastard fallow ;
partly dredge-corn and partly oats 1908, and then seeded down to
grass ; well manured with farmyard manure and phosphates while
under cultivation, but no part of it under a cleaning crop ; mown
for hay in 1909 and 1910; subsequently grazed; 5 cwt. Basic
Slag per acre applied in winter 1913-14. Cynosurus cristatus was
very prominent in the hay crop of 1905.

The area is bounded to the north by uncultivable land in
which Ulex europaeus (used for horse fodder) is dominant in most
places. In other places, Rubus spp., and Preris aguilina are
prominent and Agrostis tenuis, Anthoxanthum odoratum, Dactylis
glomerata, Festuca rubra, Holcus lanatus, and Cynosurus cristatus
are more or less abundant.

AREA 3. Part of the same field as Area 2.

Altitude: 650 feet.

Soil; Type ii. (p. 3).

History: As for Area 2 up till 1905 ; oats 1905, and under
cultivation for five successive seasons, during which time the

AREAS EXAMINED, 7

whole area was under a cleaning crop; well manured while under
cultivation ; seeded down to pasture 1909 ; mown for hay 1910»
and subsequently grazed without further manuring.

Arga 4. Adjoining Areas x and 2.

Altitude: 675 feet.

Soil: Type ii. (p. 3).

History : In grass probably for about 10 years prior to 1898 ;
ploughed in the spring of 1898 and put through a course of
cultivation similar to that of Area 1; laid down to grass in 1903;

mown for hay in 1904; subsequently grazed without further
manuring.

The details of the seeds-mixture used are not known, but it
is known that Lolium perenne, Dactylis glomerata, ‘Broad Red
Clover,” and probably Plantago lanceolata were included.

Ara 5. Adjoining Area 1 diagonally.

Altitude: 600 feet.

Soil: Type ii. (p. 3); slightly deeper and lighter than Area 1.

History: Grazed continuously with very little manuring for at
least 50 years prior to 1903; oats 1903; bastard fallow; dredge-
corm 19043 oats 1905; bastard fallow; dredge-corn 1906, when
it was seeded down ; well manured while under cultivation, and
partly under a cleaning crop in 1904 and 1905; mown for hay
1907; dressed with lime compost winter 1907-8 ; mown for hay
1908 ; subsequently grazed without manure.

Holcus lanatus was very prominent in the 1g08 hay crop.

AREA 6,

Altitude: 850 feet.

Soil: Type i. (p. 3)

History ; It is known to have been under cultivation about
1855, and that during that period of cultivation it was partly under
a cleaning crop. It was also manured at that time with peat

8 PASTURE STUDIES: SOME RESULTS.

ashes and guano,? and to some extent with farmyard manure, but
there is no evidence to show that it was limed.

The conditions under which the area was then laid down to
grass are not known, nor its treatment in the years immediately
following. It is known, however, that in a few years it was again
merged in the sheep walk of which it previously formed a part,
and was allowed to revert to its natural herbage without any inter-
ference. About 1898, however, it was given a light dressing of
Basic Slag, which had very little apparent effect. By this time
the herbage was of a grass-heath type, dotted with Ulex europaeus
and a few Ulex Gallii bushes closely trimmed by sheep. Patches
of Callina vulgaris also occurred on the thinnest soil. After 1898
Lotus corniculatus colonies were fairly common, and some weak
Trifolium repens plants were to be found.

The area was ploughed in the winter of 1906-7 ; oats 1907;
oats 1908; bastard fallow ; partly dredge-corn and partly rape
1909 ; oats 1910; bastard fallow ; dredge-corn 1911, and seeded
down to pasture; well manured with farmyard manure and
phosphates, and to a slight extent with kainit and nitrate of soda
while under cultivation; mown for hay 1912; subsequently
grazed, chiefly by cattle up till 1918, when it was again thrown
open to sheep and very heavily grazed. In 1918 it was also badly
scorched in the drought of the early summer, as the soil is thin,
on a strong slope with a southern aspect. A dressing of Basic
Slag was applied in the spring of 1916.

AREA 7. Part of the same field as Area 6,

Altitude: 750 feet.

Soil: Type i. (p. 3)

History: As for Area 6 up till 1906 ; ploughed, winter 1907-8
and put through exactly the same course of cultivation as Area
6, with similar manuring ; laid down to grass 1912, but, as will be
seen from Table IV., the seeds-mixture used differed widely from

1 This fact is of considerable interest, as the first cargo of this manure only
reached England in 1840 (5), and the farm is in a very inaccessible locality.

AREAS EXAMINED. 9

that used on Area 6. The after treatment was also different in
that Area 7 was not mown for hay.

Area 8. Adjoining Area 6.

Altitude: 850 feet.
Soil: Type i. (p. 3).

ffistory ; This area has formed part of the sheep walk already
referred to for an unknown period (probably over a hundred
years), but there is evidence that it was under cultivation at some
remote period. The presence of imperfectly burnt limestone
cores shows that it has been limed, but otherwise it is not known
to have been manured. At the time of examination, a consider-
able proportion of the field was over-run by Ulex europaeus, but
on the part examined only scattered bushes were found, and in
between, the herbage was typical grass heath.

It will be seen that these eight areas fall into two clearly defined
groups. Areas 1-5 are mainly on soil-type ii, and have generally
been regularly cultivated on the long rotation system. The slope
of these is also much more gentle than that of Areas 6-8, and
they are much more easily accessible.

Areas 6-8 are on soil-type 1. The character of the herbage
natural to these situations at a late stage of stabilisation when
untended is shown by the analyses of the herbage of Area 8
(Tables IV. and V., pp. 16a, 164).

The results obtained for these eight areas are especially
valuable on account of the facts that some of them have been
repeatedly examined, and that the soil conditions for each group
are almost perfectly comparable. The examinations in each year
were also made for all areas in August and September.

Further particulars are given in Table II. (p. 10).

ft seems probable that the plant had been sown on part of the field for
fodder purposes.

to PASTURE STUDIES: SOME RESULTS.

TasB._e II.

Showing Altitude, Soil-type, and Extent of each Area, the Age of
the Pasture, and Method of Examination employed.*

Area I 2 3 4 5 6 7 8
Altitude | 650ft. | 7ooft. | 650ft. | 675ft. | Gooft. | Ssoft. | 750ft. | S5oft.
Soil-type ii. ii. ii. ii. ii. i. i. i,
Extent Sac. | 5ac. | Zac. | 3ac. | 6ac. | 6ac. gac. | 2ac,
Age
34 Sp. fr. _ = eS = = = a
44 Both _ — — — | P.c. fr. | Sp. fr. _
53 P.c. fr. | Both _ _ — Both | P.c. fr. _
64 — | P.c. fr. _ _ — |{P.c. fr. 35 =e
73 = ” Sp. fr. = == ” cae =
8h — Both | P.c. fr. _ _ _— = iss
93 — |P.c. fr. | P.c. fr, _ _ _ aners ies
104 — | Both _ — | Sp. fr. _— ~ ne
133 —_ _ — | Both _ _ = —
100 (?) _ _ _ _ _ _- _ Both

*The altitudes are given in feet and the age in years. Sp. fr. = Specific
Frequency method; P.c. fr.= Percentage Frequency method. For con-
venience, these abbreviations are largely used in the text.

SEEDS-MIXTURES.

The chief constituents of the seeds-mixtures used for all
these areas except Areas 4 and 8 are shown in Table IV. (p. 16a),
The full details of that used on Area 4 are not known. It was,
however, fairly simple, and included no grasses except Lolium Spps
and Dactylis glomerata, It also included Trfohum pratense
(“Broad Red Clover”), and possibly, but not ptobably, Z

SEEDS-MIXTURES. II

hybridum, but no other clover seed. It also probably included
the seed of Plantago lanceolata.

Nothing is known as to the character of the seeds-mixture
used on Area 8,

AGGREGATE RESULTS.
Tas_e III.
Aggregate Results obtained by the Percentage Frequency Method.

Grami Legumi: Other Orders,

ore “eed. wea. Not Not Not
Sown. | sown. | S°¥™ | sown. | S°¥™ | Sown.

I 650 43 19°7 | 36°7 | 27°5 o'2 ml E559
I ” 54 13'2 | 36°2 | 31°8 ol _ 18°8
2 700 53 1o'5 | 56°4 | 12°4 0"4 — | 20'5
2 + 63 80 58°3 17°4 o'7 _ 15°5
2 ” 7k 75 | 52:0 | 187 9°9 — | 20°9
2 ” 84 64 55°0 18°2 o'7 _ 19°4
2 ” 94 4°8 | 644 | 15°7 08 — | 141
2 55 104 41 63°3 15°9 03 _ 16°5

3 650 8h 8:5 49°4 18'0 03 o'”7 23'1
3 ” 94 50 | 484 | 23°71 13 = 22°2
4" | 675 133 18 64°0 o'4 159 2'2 16'2

6 850 4k 73°3 8:8 8-7 32 ol 6'0
6 Pr 54 51°5 12'2 | 27°6 o'9 _ 77
6 ” 64 47'8 1570 | 24'7 3'°0 — 93
6 | os 7h | 489 | 324 | 89 | mt — | 87
7 | 750 sk | rrr | 6472 | 109 | 38 | O9 | 93
7 | » 6 | s7 | 747} 99 | 04 | o2 | 88
8 850 | 100'(?) 78's 4 81

*Details of seeds-mixture used not known.

12 PASTURE STUDIES: SOME RESULTS.

It is obviously impossible directly to compare the results for
Areas 1-5 with those for Areas 6-8 as the natural factors are
different for the two groups. For each of the groups, however,
the soil and altitude factors may be regarded as relatively constant,
and for Areas 6 and 7 the treatment of the land was so nearly
similar that this also may be considered almost constant. The
treatment of Areas 1-5, although mainly similar, was different in
respects which may be of considerable importance, while the
composition of the seeds-mixture varied considerably for each
area.

The aggregate results obtained by the P.c. fr. method are
given in terms of sown species and those not sown, but where the
seeds-mixture is widely different, the significance of these aggregate
results becomes less. This is due to the fact that in no case
probably would the soil be quite free from living plants or at least
of seeds which might produce plants in the young pasture, when
laid down to grass (3). In addition, no provision could be made
against the recolonisation of the ground by indigenous plants
whose seeds might be introduced by natural means.

The term “ sown species” used here must therefore include
all the plants of those species the seeds of which had been
included in the seeds-mixture, whether their presence be the result
of sowing the seed or not.

The difficulty which occurs under this consideration is how-
ever very much reduced by the fact that most of the areas have
been examined in successive years.

GRAMINEAE.
(2) Sown.

As will be seen from Table III., the tendency on all these
areas is towards a decrease in the proportion of sown grasses from
year to year. This is very clearly shown by the results for Area 2
where there was a steady decrease from 10-5 per cent. at 54 years
to 4-1 per cent. at rof} years. Assuming that this decrease con-
tinued at the average rate of the last three years for the further
period of three years (up till 13} years), the sown grasses would

AGGREGATE RESULTS. 13

then have been reduced to less than 1 per cent. This would
apparently not occur, as on Area 4 there was still a P.c. fr. of 1-8
at 134 years.

The large decrease on Area 1 from 19-7 per cent. at 44 years
to 13:2 per cent. at 54 years, strongly suggests that the decrease
on these areas was at first rapid, but became slower lateron. The
results for Areas 1 and 3 do not agree with those for Area 2 at
corresponding periods however, although there was relatively little
difference between Areas 2 and 3 at g$ years. These differences
may be due either to the different treatment given to the areas, or
to the composition of the seeds-mixtures used.

The results for Area 6 are in marked contrast with those for
the other areas, even with those for Area 7, which was in most
respects comparable with it. A P.c. fr. of 73-3 at 44 years is
exceptionally high, as also are the other results in succeeding years
on the same area.*

Areas 6 and 7 had similar soil conditions, and their treatment
was mainly similar throughout. The only difference in treatment
was, that Area 6 was mown for hay at 1} years while Area 7 was
grazed. It may be that this placed Area 7 at a disadvantage—
that by being grazed in the first season a large proportion of the
sown grasses was uprooted. This is probably not the case, as in
this, and even in the following year, it was grazed with cattle only.

The difference in altitude would be expected to give the
advantage to Area 7,? so that apparently the chief cause of the
difference was the difference in the composition of the seeds-
mixtures.

(4) Not sown.

Areas 2 and 4 appear to agree fairly closely, but again there
was a considerable difference between Areas 2 and 3 and between
Areas 1 and 2. These differences are probably partially explained
when the individual species are considered,3 but they may also
have some significance in connection with the treatment of the

* Cf. (18).

2 This would perhaps depend however on the nature of the seeds sown,
8 See Anthoxanthum odoratum, p. 27, especially,

14 PASTURE STUDIES: SOME RESULTS.

land. No part of Area 2, in which the unsown grasses were most
prominent, was under a cleaning crop while under cultivation ;
Area 1 was partly under a cleaning crop, while Area 3 was entirely
under sucha crop. Areas 1 and 3 were not examined at corres-
ponding ages, and it is possible that the unsown grasses had
increased rapidly on Area 3. It may be borne in mind, however,
that Area r was dressed with farmyard manure after it was laid
down to grass.

In spite of the fact that Area 6 had not been under a clean-
ing crop, the percentage of unsown grasses was exceedingly low
up to 64 years. In the following year they increased rapidly, but
this was probably due to (a) change in the method of grazing, (4)
a period of drought (see p. 8). At 74 years, however, these
grasses were very much less developed on Area 6 than on Area 7
even at 54 years. The increase on the latter area at 64 years was
probably due to the same cause as the increase on Area 6 at 7}
years.

LEGUMINOSAE.

The results for Areas 1 and 2 in respect of sown leguminous
plants differ very markedly. Other peculiar features of the results
are, the sudden changes both on Areas 2 and 6, and the difference
between Area 4 and all the others, but as this is a question
mainly of Zrifotium repens, these features are better pursued
when the individual species are discussed.

OTHER ORDERS.

The seeds of some plants belonging to other Natural Orders
were included in the seeds-mixtures for some of the areas. The
P.c, fr. results obtained for these plants are almost negligible, so
they will be considered under the various species.

The aggregate results for the unsown species do not show
any regular sequence. The most marked feature is the difference
between the area groups, 1-4 and 6-8. The results for Area 8
show that in the old grass-heath the miscellaneous plants formed
as high a proportion of the herbage as on Areas 1-4, but on Areas

1 See under ‘‘ Individual Species,” p. 15 ef seg.

AGGREGATE RESULTS. 15

6 and 7 they formed a much smaller proportion. As will be seen
from the detailed results (Table IV.), although Area 8 agreed with
Areas 1-4 in the proportion of these plants, it differed markedly
from them in other respects. It would appear therefore that the
cause of the low figures for Areas 6 and 7 is, that the heath-plants
proper had been very largely eradicated, and that for some reason
they had not been fully replaced by others.

The relatively high results for Area 3 may be connected with
the fact that the area was under a cleaning crop while under
cultivation, and that therefore these plants had not to compete
to the same extent with other indigenous plants. On the
whole, these plants seem to show some tendency to decrease on
Area 2, but on Area 4 at 134 years their total P.c. fr. was higher
than on Area 2 at 94 years and at 104 years.

INDIVIDUAL SPECIES.
GRAMINEAE.

Results for Gramineae were obtained by the Percentage
Frequency method only. These results and also the weight of
seed sown per acre are given in Table IV., p. 16a.

Lolium italicum.

This grass is usually regarded as either an annual or a
biennial, so that, although the seed was included in the seeds-
mixtures for most of the present areas, since none of them was
examined within the first three years it is not surprising that the
grass should not have been recorded.

Lolium perenne.

Results obtained by Armstrong in the Market Harborough
district of Leicestershire and Northamptonshire (1) show that
this grass may contribute largely to the herbage of some pastures.
On different types of pastures, his results were as follows :—

On an extreme type of poor pasture, 1 per cent.

On other inferior old pastures, 14-1 to 44-6 per cent.
On excellent recent pastures, 28 to 47:2 per cent.
On first rate old pastures, 33-4 to 39-3 per cent.

16 PASTURE STUDIES: SOME RESULTS.

The highest percentage frequency result obtained by the
present writer was 20-2 on a good old pasture at an altitude of
yo feet,* and the P.c. fr. of the grass on the best pasture examined
by him was 4-6.?

The present areas are not comparable with any of these.
Armstrong’s results were obtained by a different method, while
the conditions of soil and altitude were much more favourable on
the other two areas mentioned than on those for which the results
are here given.

The grass was sown on all the present areas, with the possible
exception of Area 8, in quantities ranging from 6 to 18 lb. per
acre.3 Since the grass has been sown on that part of the farm
which is under regular rotation for many years (Table I.) it is not
known whether it is truly indigenous to Areas 1-5 or not, but, as
shown by the results for Area 8, it is not indigenous to Areas 6,
7 and 8.

The highest P.c. fr. for Arcas 1-4 was 10-3 on Area 1 at 44
years. On this area it decreased to 6-1 per cent. at 54 years, and
at that time it was slightly less abundant than on Area 2 at the
corresponding period, although a heavier seeding had been given
to the former area.

On Area 2 its decrease was slower from 54 years onwards than
on Area 1 at an earlier date, but it was continuous and regular, so
that at ro} years it had a P.c. fr. of only 3-0. It was rather more
abundant on Area 3 at 8} years than on Area 2 at the correspond-
ing period, but the results for the two areas at g years are almost
identical in spite of the difference in quantity of seed sown, while
on Area 4 at 134 years there was still a P.c. fr. of 1-1.

These results indicate quite clearly that under the conditions
of these areas the grass decreases from year to year, at least up to
about 134 years, but that even in that length of time it does not
completely disappear, and that, in fact, up to about 83 years it
may be quite as abundant as in some excellent Welsh pastures

(see above).
1College Farm, Aber, Bangor.
218 years old pasture in Anglesey at 200 feet,
3 Probably more cn Area 4.

TABLE IV.—Gramineae and Leguminosae. Seeds sown, in Ib. per acre and Percentage Frequency Results. 16a

AREA. I 2 3 4 5 6 7 .
2 o 5 o ra D o

G2] a | a 32) sb | op | ] om | ot [os [32] oy | ot 143] ob [S232 { a | a | oe | oe PSE] sb | ob | r00:

Beep Me I ag 3 yrs. | yrs. | yrs. | yrs. | yrs.) yrs. Ee | yrs. | yrs. ae yrs. | 3 = 2 a yrs. | yrs. | yrs. | yrs. | 8 Z yrs. | yrs. § yrs
Lolium italicum .. wel 5 —| —-] 5 —; —| —| —}] —); —f] 3 Sex ai BP as] 4 5 Saul ser eS Sa a ex each =
L. perenne oi if BR | RORZ GSD] ae 6:6) 62) Sar 47) 3:6) Ztof 8 5.81 3°57 ? ears |r arg 63 | ai2a re] 6 o8; —] —
Dactylis glomerata wel 5 5°51 4°34 3 2:1 | £4.) @°7 | OO) Os) OF}: 22 27) go, oe o7{ TET 34] ao] 45] so] 27% 44] 776) 3°57 —
Festuca elatior et var. «| 3 13) err 4% | 203i) O84), "ORS! ora) Coe, at eae ee ee el Se ee) el ee ee ee ee ee) ee es
F. duriuscula .., weal 3 —| —f 22; —}| —| —} —}] —: —7] 2 —; —|} —| —]} —] 2 I'l] 06} O73] 2 227 EO SS
F. ovina sie wt — 1} —]| —} —}| —] —] orf —} —}] —F — fs Ps ee ah i ee jer Geis | SEM a a Ae 1°3} 163
F. rubra oon wl —] 2°09] 4°47 —] 79] Or] 7°7] 5°9| 71] 79] | 2S] 23] —] 867 —] —] ro} or] T —} —| T Pe =
Phleum pratense wa| 1:2] O75} 3 EG. G'2)|| 1083) oy | Ors) Ca 2 POMP a) ea al @ 19| 06} o4f —} —}| — -—]F —
Arrhenatherum elatius he O'S: Ose |e eee ee ee ef ee ee oe eee eee) ee ee cee ee] ee eee est eet eS
Alopecurus pratensis aeel, OE EE ee ee ee ee me mee ee ee | ee ee eet ee ee eS he
Poa pratensis”... wf) —f —] 2) | = ob) oa) oo oa Soe ee | ea ff a] — Pe ea ae ae] Ce) eh
P. trivialis pas wal — | 3°38] 2°37 — | 1°93] 2°09] 20] go] 2°37 4:7] — S76) ss | vorg hf || | GRR) GH oS ee) eee ee
P. annua a | — | 84] o6f —| —| —]}| TY —~—] —]} -] —- fr : a0 ces call ii sees Pcs ee eh ee |) EE | ote
Cynosurus cristatus wf — |] T ] 38> — | S31) g2]10°9] 94] 12:2}14:7) —} 88] 667 — ]137] —] 1 | 43°4)33°7] 304] 32.0] -—| 13) —] —
Anthoxanthum odoratum ...J) — | 2°5| 2°37 — | 189) 21'2)11°3} 12:1] 12°2] Zo} — | 61] Got —° gS) — 4} go) 62] z-2/121]f —| 08] rofl os
Agrostis spp... wf — | 143} 13°5] — | 47°5; 171 | 149] 15°22) 2r'9] 205} — | 16:9] 23°04 — | 222] —} —] 38, 91] 86! 27-2] — | 58-7! 69:3] 42°9
Tlolcus lanatus ... | —| 48! o3f —] 2°95) 26} 50) 82) 87) 68) —] 71( a3f — | S4l —] —|] 36) o9] 33) 48] —| —] os} —
Arrhenatherumelatius var. ...) — — Tr —] o1}] or; o1) o'2} —| oF] — = — —.o3f — aes = oa oe oes —_ <=, _ a
Aira caryophyllea wd — {| —tl —>P —f or! —}| —)| —} —}| —f —]} —| —t -|) —7F —FT —| —|] —| —| —F —}) —7 —] —
Sieglingia decumbens op SS SS Sa SS SS a ee ee ee ee eh | Oe O89). re! orf a Sag crea a 8e8
‘* Broad Red Clover” ad 3 } 2 } I | ? | og] 4 ar eee ee (eee (end (eee | ees ene ares

=.) ves see TE | SAP) seed | cell ee

“‘Cowgrass” ses] oR 2 T —{ —-| -—J 1 2 or | 2 —| —| 2 _
“¢ Hybrid Cow Clover”... —/| —] —— —j} —| —]| —}] —} —] YI O15 | re ae, | Umer (Pee meee | eee P| PR Re Se | |
“ Late-flowering Red Clover”) — | --| —] —/ —]| —] —! —] —] —} ~—-/| —}] -—# —/] —]7 —} —]}] —| —) —| —f 2) _
Trifolium hybridum eit || Sh ag Se pS ey See ea SS a —~| —T| ? —{ 141 2 | o2] 03] T —i 2 ee een) ee
‘* White or Dutch Clover” ... 41275) 3r8) 2 | 124/174) 187/182) 157/159] 14 | 180] 231], — lis1, —} 2 84|27°3|247| So} 2 |ro9! gg) —
Anthyllis Vulneraria sea} —| — 4/ —| —-?| —} —] —-] - 4} —{ —] —| —] —] 2 coe | eee || eel es I ae ee eee
Medicago sativa... wel —f| —} —F—} —} —-!) -—] — tt omy os a mH nH er! OT KI 2 el gecesi eee ees I bad) Ca =
M. lupulina ts vel _ = — — — = a == =] = —_ — —_ —_ iL —_ _ = oe a ss = _ eo

*T = Traces,

TABLE V.—Percentage Frequency and Specific Frequency Results for the most important plants other than Gramineae.

Percentage Frequencies.

Specific Frequencies.

AREAS.

Age of Pasture (years).

Trifolium pratense?
T. hybridum? aay
T. repens!

Lotus spp.

Plantago lanceolata
Achillea Millefolium
Ranunculus repens...
Bellis perennis
Prunella vulgaris ..,
Cerastium vulgatum
Sagina procumbens
Luzula campestris ...
Potentilla erecta
Leontodon autumnalis
Senecio Jacobaea ...
Rumex Acetosa
Centaurea nigra
Plantago major
Rumex Acetosella ...
Potentilla anserina...

Taraxacum vulgare

Chrysanthemum Leucanthemum

Polygala vulgaris ...
Veronica officinalis
Viola canina ae
Hieracium Pilosella
Euphrasia officinalis
Thymus Serpyllum
Galium hercynicum

Calluna vulgaris...

eee

Ow

lone @

rod 84 | 9d 13!

09} 0'6 | 02] 2°0

o'2}0'4 | 06] or

T J[o3) T Jor

44

8),

LSS)

n a

,

24
16

15

13

166
7 | 8
4k f100?
10 | —
60} —
22 | 40
26 | Io
—]10
38 | —
14] —
224 7
12] —
16] —
? ?
€4 | 87
2 =
$4.) =
—J|i1o
44 22
— | 22
ame ay 4
2155
18175
4 | 42

1 See Table IV. for quantities of seed sown.

*T=Traces.

* Heavy type indicates that the species was sown on the area,

INDIVIDUAL SPECIES. 7

They also suggest that the decreass is rapid up to the fifth
year, and that after, there is no definite relationship between the
amount of seed sown, and the P.c. fr. of the grass, at least where
not less than 8 lb. per acre is sown.

Since the grass is exotic to Areas 6 to 8, it is interesting to
note that after a lighter seeding it had a higher P.c. fr. on Area 6
than on Area 1 at 44 years. Here, however, there was a greater
decrease in the following year, and a further decrease in the next
year. At 74 years its P.c. fr. on Area 6 had fallen to 1-8, but the
system of grazing had then been changed, and the sudden decrease
from 64 years to 74 years is probably connected with this, and
possibly also with the fact that the area suffered rather severely
from drought during this period.

With a light seeding (6 Ib. per acre) the grass was but slightly
represented on Area 7 at 54 years, and it was not recorded on this
area at 64 years.

The difference in the amount of seed sown on Areas 6 and 7
was considerable, but it does not seem alone sufficient to account
for the difference in the P.c. fr. of the grass on the two areas at
corresponding periods.

From the results for Area 6 it is seen, however, that even
where not indigenous the grass may persist in fair quantity up to
64 years, and in some quantity up to at least 74 years in spite of
very heavy grazing with sheep at the end of this period.

Since the presence of the grass in the pasture was desired,
these results, (with the possible exception of those for Area 7),
fully justify the inclusion of the seed in the seeds-mixture, but at
the same time they show that under these conditions the grass
does not form a very high proportion of the total herbage after a
few years, as on Area 1 it was the second most plentiful grass at
44 years, while on Area 2 at 10} years it had fallen to the seventh
place, and on Area 6 it had fallen from second place at 4} years
to sixth place at 7} years.

1Stapledon, however, found the grass ‘‘wonderfully resistant and
persistent ” in drought (15).

18 PASTURE STUDIES: SOME RESULTS.
Dactyhis glomerata.

The highest percentage obtained by Armstrong for this grass
was 9:5 on one of the inferior old pastures examined by him (1).
On another inferior old pasture he found only o-9 per cent., while
on first-rate old pasture it ranged from o-2 per cent. to 4-5 per
cent.

On the best pasture examined by the present writer its P.c.
fr. was 1-1,% while it was not recorded on some of the best old
pastures examined. The highest P.c. fr. obtained was 9-5 on a
four-year-old pasture after a seeding of 8 lb. per acre.?

The seed of the grass was not included in the seeds-mixtures
used on the farm on which the present areas are situated until
1903, but the grass was more or less abundant in a permanent
hay meadow and along hedgerows up to 700 to 750 feet even
before this, so that it may be regarded as more or less strongly
indigenous to situations up to 700 feet.

The seed was included in the seeds-mixtures for all the
present areas except Area 8,

On Areas 1-4 the highest P.c. fr. obtained was 5-5 on Area 1
at 44 years. It decreased somewhat on this area in the following
year, but even then it was considerably higher than on Area 2
(where a lighter seeding had been given) at the corresponding
date. On Area 2 it decreased fairly rapidly up to 74 years, but it
still had a P.c. fr. of 0-5 at 104 years.

On Area 4, it had persisted rather better apparently, but it is
not known what quantity of seed had been sown on this area. It
had also persisted better on Area 3 than on Area 2. This may be
due to the fact that Area 3 had been more thoroughly cleaned
while under cultivation.

Although not indigenous to Areas 6 and 7, the grass was
established on these areas even better than on Areas 1-4. On
Area 7 especially it had a very high P.c. fr. at 54 years, while on
Area 6 it appeared to be holding its ground from 4} years to 64
years. Asinthe case of Loum perenne, however, it decreased

1 Anglesey, 200 feet, 18 years.
2 College Farm, Aberystwyth, 400 feet.

INDIVIDUAL SPECIES. 19

considerably both on Area 6 and on Area 7 under the influence
of heavy sheep grazing and (or) drought in 1918.

The results for all these areas taken together suggest that the
grass has held its own better on the lighter and drier soil (Areas
6 and 7), and there is some indication also that there may bea
distinct relationship between the amount of seed sown and the
P.c. fr. of the grass for some years where the other conditions are
fairly equal. There is a suggestion, however, that in some cases
at least, thorough cleaning of the land is advantageous to the grass

(Area 3), although on Area 7 it was by far the most successful
sown grass.

Festuca elatior et var. pratensis.

These grasses were only recorded on one of the grazed
pastures examined by Armstrong, and.on this they formed 1 per
cent. of the herbage (1).

On the best pasture examined by the writer, they had a
P.c. fr. of 6, but they have not been recorded on any of the other
old pastures examined. Their presence to the extent of 6 per
cent. on the pasture referred to may not mean that they were
indigenous. The pasture was but 18 years old, and they may
have persisted from sowing. It is more probable, however, that
their presence was due to the fact that cattle are regularly winter-
fed on the field, and the seed might be repeatedly introduced in
the hay. That this is at least possible is shown by the fact that
although the grasses are completely exotic to Areas 1-8 and even
to the district, they appeared in small quantity on a permanent
hay meadow on the farm in a few years after the seed had first
been sown on Area 2.

Only a very small quantity of seed was sown as an experi-
ment on Area 2, but the grasses were recorded at each examination
up to 9# years.

On Area 3 a heavier seeding was given, and although the
P.c. fr. was low at 44 years, it was slightly higher than that given
by a heavier seeding of Pileum pratense (Table IV., p. 16a), and
in the following year the difference was still greater as the Festuca
spp. had increased while Phlewm pratense had decreased,

20 PASTURE STUDIES: SOME RESULTS.

Thus it is shown that although exotic, one or the other or
both these grasses may persist to some extent up to 9% years
under the conditions prevailing on these areas.

Festuca duriuscula*

Festuca duriuscula is the commercial name under which the
seed of a variety of F. ovzna seme was sold before the war. It is
said by Hunter (7) to be the seed of F. ovina (L.), var. duriuscula
(Koch), but in any case it is different in many respects from any
fine-leaved fescue found by the present writer to be indigenous to
any of the present areas, and even to any areas in any part of
Wales investigated by him.

As will be seen from Table IV., its seed was included in the
seeds-mixtures for several of the present areas, but even though
sown in quantities from 2 to 3 lb. per acre on Areas 1-3 the grass
was not recorded on any of these areas.

On Areas 6 and 7 it was somewhat more successful. On
Area 6 it was not separated from other varieties of / ovina
which may have been present at 44 years, but after a seeding of
2 lb. each of this variety and another commercial variety of
£”, ovina, the total P.c. fr. was only 2-7. Itis probable, however,
that this was mainly #. duriuscula, as in other years other varieties
were only present in very small quantities, and / duriuscula was
decreasing at such arate that at 74 years it was represented by
only 0-3 per cent.

On Area 7 the grass had a P.c. fr. of 2-7 at 54 years, but
here also it decreased to 1-2 per cent. by the following year.

These results show that the grass is useless for such situations
as Areas 1-4, at least after 44 years, and that even on situations
such as 6 and 7, it is of relatively little value. They certainly do
not justify the claims generally made for the grass (2), (7), (19).

1The results for the various forms of fine-leaved fescues are not given
separately by Armstrong (I).

2Much of the seed sold in recent years under this name is that of quite
a different grass—probably even a different species. These grasses have been
the subject of a special investigation by the writer for some time.

INDIVIDUAL SPECIES. 21
Festuca ovina,

Seeds sown under this name apparently may be those of any
species or variety of fine-leaved fescue which are available in
quantity. Those sold as F. ovina tenuifolia usually conform to a
distinct type.

It is not known which of these types was sown on Area 6, but
in view of the fact that at 54 years only o-1 per cent. of / ovina
was recorded, this is of little importance. In the two years which
followed, the grass completely disappeared, although, as shown by
the fact that it hada P.c. fr. of 16-3 on the old grass-heath of
Area 8 adjoining, the grass is strongly indigenous.

This shows, not only that the sown variety was valueless for
this situation, but that even the indigenous variety had not
re-colonised the ground to an appreciable extent at 74 years.

On Area 7, where no seed of this name was sown, traces
were found at 54 years, and a percentage frequency of 1-3 at 64
years, so that evidently for some reason the grass was re-colonising
this area at a faster rate than Area 6. This may be chiefly due to
the difference in the competing plants, especially the great develop-
ment of Cynosurus cristatus on Area 6.

Festuca rubra.

A creeping variety of this grass is indigenous to Areas 1-4,
and it may also be found in very small quantities in the shade of
gorse bushes on Area 8. It is probable therefore that it was also
to be found to a slight extent on Areas 6 and 7 in similar situa-
tions in the original grass-heath. The results for Area 8, however,
show that it is not indigenous to the open grass-heath.

The seed of this creeping variety is not obtainable com-
mercially, and the seed of no variety of F. rubra was sown on any
of these areas. Vet, on Area 2 especially, it formed an important
part of the herbage (7-9 per cent.) even at 54 years, and with
some fluctuation in the intervening years, it had exactly the same
P.c. fr. at 104 years.

On Area 1, on the other hand, it had only a P.c. fr. of 4-4 at
54 years, while the highest for Area 3 was only 2-8 at 8} years.

22 PASTURE STUDIES: SOME RESULTS.

Observation has shown that where this creeping variety is
strongly indigenous it may prove a rather troublesome arable land
weed, and therefore, unless the land is rigorously cleaned while
under cultivation, the grass may persist bodily. Even where the
land is put under a cleaning crop, small portions of the rhizomes
may survive. It would be expected therefore that under condi-
tions otherwise equal the grass would be most prevalent in a young
pasture where the land had been least perfectly cleaned.

The results for Areas 1-3 perfectly agree with this. Area 2
was not even partly under a cleaning crop; Area r was partly
under a cleaning crop while Area 3 was entirely under such a
crop.

A percentage frequency of 8-6 at 134 years on Area 4, which
had been partly under a cleaning crop, does not contradict the
explanation, as here the grass had had a longer period to re-
colonise the ground.

The fact that Areas 2 and 4 agree so closely, suggests that on
this land, under the conditions prevailing, the grass reaches a
certain stability at about 7 to 9 per cent. of the total herbage.

On Areas 6 and 7 the grass, as might have been expected
from the fact that it is only weakly indigenous, was but poorly
represented. On Area 6it had a P.c. fr. of 1-0 at 44 years, but traces
only were left at 64 years, and it was not recorded at 74 years. This
suggests that it was gradually decreasing from a more prominent
position occupied prior to 4} years. Since the area was not
under a cleaning crop, the shade provided by the growing corn
and by the hay crop would probably favour its development, but
when open pasture conditions followed, it rapidly decreased.

The results for Area 7 do not confirm this theory, as here
traces only were found at 54 years, and a P.c. fr. of 1-3 at 64
years. This area was, however, much less well grazed than Area
6, and it is possible that the shade provided by the rough
Agrostis herbage was sufficient for its development.

In no case was there any indication that the grass is rejected
by stock, so that the only apparent objection to it is its presence
as an arable land weed,

INDIVIDUAL SPECIES. 23

Phieum pratense.

This grass is a complete exotic to all the present areas, and
its seed was only introduced to the farm in 1901.' Its seed was
sown on Areas 1, 2, 3 and 6 in quantities from 2 lb. to 4 lb. per
acre.

It was prominent in the hay crop taken from Area 2 at 2}
years, but at 54 years it had only a P.c. fr. of 1-5, and in the next
year it decreased to o-2, Even at 104 years, however, it had not
completely disappeared.

With a somewhat heavier seeding its initial history was some-
what similar on Area 1, but here and on Area 3 its decrease was
rather more rapid.

On Area 6, in spite of the drier situation, it had higher per-
centage frequencies than on Area 1 at corresponding periods,
although the seeding was lighter. It was not recorded on this
area at 74 years.

Apparently therefore, this was one of the least satisfactory
grasses sown, but it evidently does persist for some years even on
these situations. Here it seems to be more valuable for hay than
in a pasture.

Arrhenatherum elatius.

The typical species is an exotic to these areas, but its seed
was included in the seeds-mixtures for Areas 1 and 7. It was not
recorded on Area 7, and its highest P.c. fr.on Area 1 was 0-3
after a seeding of 2 lb. per acre. This grass is therefore
apparently of very little value for these situations.

Its variety, 4. elatius tuberosum, is indigenous to Areas 1-4,
and on these areas is a more or less troublesome arable land
weed. In these pastures, its highest P.c. fr. was 0-7 on Area 2
at 104 years, and its next highest was 0-3 on Area 4 at I 3% years
Pasturing conditions, therefore, do not exterminate it, but even if
it were a desirable pasture plant, it was not sufficiently abundant

1 The grass must be regarded as a complete exotic in spite of the fact that
for several years a small colony, about a foot in diameter, existed in a very old
pasture field on the farm, which was at that time continuously mown for hay,
This field was remote from the present areas,

24 PASTURE STUDIES: SOME RESULTS.

on these areas to counterbalance its objectionable character as
an arable land weed.

Poa pratensis.

On most of the pastures examined by Armstrong (1), this
grass was absent, and only in one case was it found by him to
reach = per cent. of the total herbage.

Sutton (19) expresses the opinion that “it should be service-
able in Wales,” but the present results give no confirmation to
this view.

Only on Area 2, and then only in very small quantity, was it
sown, and its highest P.c. fr. on this area was 0-6, while it was not
recorded on the area in three out of six examinations. It was not
recorded on Areas 1 and 3, but it had a P.c. fr. of o-1 on Area 4
at 134 years, and on Area 6 at 54 years.

On all pastures examined by the writer its highest P.c. fr.
was 3:2."

Poa trivials.

Armstrong’s results (I) show that the prevalence of this grass
varies in some cases from month to month. He found its average
range to be from o-8 to 5-5 per cent. of the total herbage on first-
rate old pastures.

Its seed was not sown on any of the present areas, but the
grass was recorded in percentage frejuencies ranging from o-g to
5:6 on Areas 1-4.2, It was most strongly represented on Area 3,
but it was more strongly represented on Area 1 than on Area 2 at
corresponding periods. On Area 1 it decreased from 4$ to 54
years, but on Area 2 it increased almost regularly from 5% years
to 104 years. The difference between Area 2 at ro4 years and
Area 4 at 13$ years is very marked, and suggests that a rapid
decrease would follow on Area 2.

On these areas, however, the grass is fairly strongly indigen-
ous, but on Areas 6-8, as shown by its absence from areas 7 and
8, it is not indigenous under natural conditions. Yet it was

1 Anglesey, an old pasture of good quality at 50 feet,
2 This grass was not separated from P. annia on Area 3 at 84 years,

INDIVIDUAL SPECIES. 25

recorded on Area 6 at 54, 64 and 74 years in quantities up to 1-6
per cent., but it seems to have been adversely affected by the
heavy grazing with sheep and (or) drought in 1918.*

Poa annua.

This was generally of little or no importance, but it had the
very high P.c. fr. of 8-4 on Area 1 at 4} years. Probably this
was due to the open condition of the young pasture.

Cynosurus cristatus.

Armstrong (I) found this grass to be more abundant on
inferior old pastures than on better pastures, but even in first-rate
old pastures he foundits range to be from 4-4 to 11-3 per cent.
of the total herbage. On the best pasture examined by the
present writer it had a P.c. fr. of 12-9 at 18 years. It is not
known whether it had been sown in this field or not.

The grass is strongly indigenous to Areas 1-4, but exotic to
Areas 6-8 under natural conditions. Its seed was not intentionally
sown on the farm until 1911, but it was very prominent on
Areas 1, 2 and 3 in the pastures which preceded that of the
rotation under investigation. It was especially prominent in the
hay crop taken from Area 2 in 1905.

Its seed was not included in the seeds-mixtures for Areas 1-4,
and since it very rarely occurs as an impurity in other seed samples
(8), (x), (14), its occurrence on these areas must be considered
as more or less natural.

On Area 1 its development was very weak at 44 years, but it
had increased considerably at 54 years. Even then, however,
its P.c. fr. was much lower than on Area 2 at the corresponding
period. On the latter area it had a P.c. fr. of 8-1 at 54 years,
and with a small set back at 84, it increased regularly from year
to year to 14-7 at ro$ years. By this time it had exceeded the
P.c. fr. on Area 4 at 134 years, so that probably on that area there
had been a slight decrease. On Area 3 it was considerably less
well developed than on Area 2 at corresponding periods, and even
showed a decrease from 84 years to 94 years.

tcf Stapledon (15).

26 PASTURE STUDIES: SOME RESULTS.

The soil conditions on all four areas are very similar, but the
treatment of the land and the seeds-mixtures were different. It is
difficult to estimate the influence of the seeds-mixture on the
development of the grass. In habit, the grass most closely
resembles Lohum perenne, but the results for the two grasses are
not complementary, although on Area 2 itself, they were very
nearly so. The total for the two grasses on the other areas differ
considerably from those for Area 2.

The difference in the treatment of Areas 1, 2 and 3 has
already been referred to, but how this could effect the difference
is not at all clear. The grass is not included by Stapledon in his
list of arable land weeds, and therefore, presumably, it does not
persist bodily through a period of cultivation. This would also
be expected from its caespitose habit.

It has been mentioned that the grass was very plentiful in the
hay crop taken from Area 2 in 1905. As the hay was mown late
in the season it is probable that a large quantity of seed was shed
in harvesting, and would be ploughed in in the following winter.
It is not known how prevalent the grass may be on land investi-
gated for dormant seeds by Brenchley (3), but it is not mentioned
by her as having been produced from dormant seeds, so that
there is no direct evidence to show that the seeds of the grass are
capable of lying dormant in the soil for any appreciable length of
time. At the same time, this is strongly suggested by the present
results.

Another factor which may have had considerable effect is the
presence of uncultivable land adjoining Areas 2 and 4. The grass
is fairly abundant in the more open spaces of this land, which
would therefore afford a convenient centre of seed distribution.
Area 2 lies between this uncultivable land and Areas 1 and 3, but
the grass is also fairly abundant on hedge banks surrounding these
ateas. The seeds are apparently quite unsuited to wind distribu-

1(17) Table I.
In a few experiments of a similar kind conducted by the writer, no plants

of this grass were obtained, although the soil was taken from land where the
grass is abundant, ‘

INDIVIDUAL SPECIES. 27

tion, but the areas are open to sheep in winter and early spring,
so that they may be the distributing agents.

As shown by Table IV., p. 16a, the grass was not recorded on
Area 8, and it is absent from other land adjoining Areas 6 and 7, so
that it may be assumed to be naturally exotic to these. Seed was
sown on Area 6 at the rate of 1 lb per acre, and the results, as
shown by Table IV., are remarkable. The grass decreased
considerably from 44 years to 54 years, and again decreased some-
what in the following year, but with the heavy grazing with sheep
it again increased, so that even at 74 years it still had a P.c. fr. of
32-0, and was the most prevalent grass.

The effectiveness of the seeding on this area is strikingly
shown by a comparison of the results with those for Area 7, where
the grass was not sown. On the latter area the grass had a P.c. fr.
of 1-3 at 54 years, but was not recorded at 64 years. How far
the difference in the herbage of the two areas generally may be
due to this factor alone, it is not possible to say, but since the soil
and general treatment of the two areas were practically identical,
the suggestion that this was the controlling factor is very strong.
At least, it appears to be a very important factor, and the
difference in the herbage of the two areas well illustrates the
importance of a suitable seeds-mixture. It also shows how well a
plant which may be exotic or at most only very weakly indigenous
to land in its neglected state may flourish on such land when
properly cultivated and manured.

Anthoxanthum odoratum.

This grass was not recorded by Armstrong on first-rate old
pastures, on excellent recent pastures, nor on an exreme type of
poor pasture, and only up to 3 per cent. of the total herbage on
other pastures (I).

On the best old pasture examined by the present writer it
had a P.c. fr. of 4:5, and on the best pasture of all (at 18 years),
37:

It is undoubtedly indigenous to all the present areas, as even
on Area 8 it formed o-5 per cent. of the herbage,

28 PASTURE STUDIES: SOME RESULTS.

The difference in the development of this grass on Areas 1-4,
on none of which had it been sown, is even more striking than
that of Cynosurus cristatus. At 54 years its percentage frequency
was 2-3 on Area rt and 18-9 on Area 2, and on the latter area it
increased to 21-2 at 64 years. There was a great decrease in the
following year, but then again it remained practically constant
until another sudden decrease occurred at ro} years, and by this
time there was little difference between Area 2 and Area 3 at 9}
years, but the grass had now a considerably lower P.c. fr. than on
Area 4 at 134 years.

The grass probably does not survive a period of cultivation
to any great extent either bodily or as dormant seeds (17), (3),
but its “seed” is apparently fairly well suited to rapid dispersal
both by wind and animals. Since it occurs in considerable
quantity on the uncultivable ground adjoining Areas 2 and 4, this
ground probably formed a very effective seed-distributing centre,
and this may be the chief cause of its great abundance on Area 2
io the early part of its history. Areas 1 and 3 were much more
remote from this uncultivable land.

A gradual decrease from the high percentage frequencies of
Area 2 at 54 and 64 years might be expected in subsequent years,
but the sudden decrease from 21-2 at 65 years to 11-3 at 74 years is
difficult to explain. A dresssing of Basic Slag was appiied to the
area at 5 years, but this was followed by a slight increase at 54
years. The manure may have a slight encouraging effect on the
grass itself, but this may be so much greater on other plants as
to put this grass at a disadvantage in competition. This theory
is not borne out by other results obtained.?

A small amount of seed was included in the seeds-mixture
for Area 6, but not in that for Area 7. The seeding seems
to have been very effective, as the grass formed a very much
larger proportion of the herbage of the pasture,on the former

10Qn Area 6, the grass decreased somewhat in the year immediately
following the application of Slag, and increased slightly in the following year.
The ultimate effect of Slag on the grass is in some cases at least relatively very
small, See (23).

INDIVIDUAL SPECIES. 89

than on the latter area. Even with this small seeding, how-
ever, the grass was much less represented on Area 6 than on
Area 2 up to 64 years, but at 74 years it was more developed on
Area 6. The increase on Area 6 at 74 years seems to indicate
that this grass is encouraged by heavy grazing with sheep, but
during the examinations abundant evidence was found that this
was not due to their neglect of the plant itself.

The behaviour of the grass on these areas suggests that under
these conditions it is not a very strenuous competitor, although it
may recolonise the land rapidly and become very prominent in a
young pasture where a suitable seed-distributing centre is near
at hand. In the absence of such a centre it colonises but slowly,
and in such cases, if the presence of the grass is desired in the
pasture, colonisation on suitable soil may be greatly accelerated
by the inclusion of even a very small quantity of the seed ina
seeds-mixture.

Foleus lanatus.

Armstrong’s results (I) show that the presence or absence of
this grass in a pasture is no reliable indication of the quality of
the pasture, but ‘results obtained by Hall and Russell (6)' seem
to show that on the whole it is more plentiful on the better
feeding pastures.

The seeds of the grass are never intentionally included in
seeds-mixtures, but low grade seed samples of Lolium spp.
especially may contain considerable quantities of them (8),
(II), (14).

It was not recorded on Area 8 (grass-heath) nor on Area 7
at 54 years, and only in very smail quantity on the latter area at
64 years, but on Area 6 it was fairly well represented except at 54
years. The cause of its abnormally low proportion in this year as
compared with other years is not known, but it may be a
temporary effect of the Basic Slag which was applied at 4 years,
The manure had no such effect on the grass on Area 2 however,
as on that area there was a continuous increase from 2-5 per cent.

1 These results were obtained by quite a different method of analysis, so
they are rarely referred to in this paper.

30 PASTURE STUDIES: SOME RESULTS.

at 54 years to 8-7 per cent. at 9$ years, followed by a decrease to
68 per cent. at 104 years.

The grass was much more abundant on Area 1 than on
Area 2 at corresponding periods. On the former area there had
been a sudden increase from 4-8 per cent. at 44 years to 9-3 per
cent. at 54 years. This large and sudden increase may be
temporary only, but such a change was not met with on Area 2.
The grass seemed to behave differently also on Areas 2 and 3, as
on the latter area there was a marked decrease from 84 to 94
years. On Area 4, however, its P.c. fr. at 134 years was very near
that for Area 2 at g} years.

On the whole, therefore, this grass seems to increase fairly
gradually up to about ro years on this soil, but then acquiresa
degree of stability. On soil-type i. (Areas 6-8) its developmemt
seems to depend very largely on the species it has to compete
with. On Area 7, where its chief competitor was Agrostis tenuis,
it made very little progress up to 63 years, but on Area 6, where
A. tenuis was apparently kept well in ckeck by other species, it
was more successful.

Agrostis spp.

Only traces of these grasses were found on an extreme type
of poor pasture investigated by Armstrong, but on first-rate old
pastures he found them in quantities up to ro per cent. of the
total herbage (1). On the best pasture investigated by the
present writer they had a percentage frequency of 27:1. The
highest P.c. fr. found was on a neglected 12 year old pasture,
where it reached g2-9 of the total herbage.?

Both A. alba and A. tenuis occur on Areas 1-4, but they
were not separated in the examinations. On these areas, they are
probably the most troublesome of all arable land weeds, and they
are very difficult to eradicate even with the most thorough, clean-
ing. Brenchley (3) obtained 43 “seedlings” from the soil from
two holes Gin. x 6in. x gin. in arable land “under ordinary
farm management,” and they were recorded by Stapledon (£7) ia

2 Great Orme, Llandudno, 500 feet,

INDIVIDUAL SPECIES. 3r

land under Roots, Cereals and Seeds. It is not surprising
therefore, to find the genus strongly represented even in a young
pasture, and although a thorough cleaning may have a consider-
able effect for some years in some cases, the fact that the grasses
had a higher P.c. fr. on Area 3 at 94 years than on Area 2 at the
corresponding period suggests that after some lapse of time this
has little effect.

The genus was less strongly represented on Area 1 than on
Area 2 at corresponding periods, but the difference was not very
great. On Area 2, however, there was a considerable decrease
from 64 to 74 years. This, although there was no apparent
change in the season immediately following its application (64
years), may be connected with the application of Basic Slag.
Between 84 and 9} years there was again a large increase, and
the P.c. fr. of these grasses was very nearly the same for Area 2 at
g} years, Area 2 at o$ years and ro} years, and Area 4 at 134
years, but even then the. genus was less represented than on the
best pasture examined by the writer.

On Areas 6-8 only Agrostis tenuis was present. It formed
42-9 per cent. of the total herbage of the old grass-heath (Area 8),
but it formed 58-7 per cent. of the total herbage of Area 7 even
at 54 years, and this increased to 69-3 per cent. under heavy
grazing at 64 years. The same influence of very heavy grazing
was found on Area 6, where the grass increased from 8-6 per cent.
at 64 years to 27-2 per cent. at 74 years. On this area the grass
had an exceedingly low P.c. fr. at 44, 54 and 64 years in spite of
the fact that the area had not been under a cleaning crop.

Since the soil conditions and the general treatment were
practically identical for the two areas, the difference in the
development of Agrostis must apparently be attributed to either
of the following two causes :—

(a) As already stated, a crop of hay was taken from
Area 6 at 1} years from seeding, while Area 7 was grazed
at the corresponding period. Area 6 was, however, put at a
disadvantage to the extent that it was not compensated for
this extra crop taken by extra manuring, but in spite of this,

32 PASTURE STUDIES: SOME RESULTS.

Agrostis was incemparably more prevalent on Area 7 in later
years. As Area 7 was grazed with cattle only, at this period,
it is not probable that a serious amount of the sown seed-
lings would be uprooted. On the whole, therefore, it does
not seem probable that the difference was due to this cause.

(6) The seeds-mixtures for the two areas were quite
different in somerespects, and the success of Cynosurus
eristatus especially, and to a smaller extent of Anthoxanthum
odoratum and Lolium perenne on Area 6 seems to indicate
that this was the main cause. The development of Dactylis
glomerata evidently had no appreciable influence against
Agrostis, as this was even better developed on Area 7 than
on Area 6.

Atra caryophyllea.

This grass is not indigenous to any of the present areas, and
it was only recorded once. It is known to have been present in
considerable abundance in the very young pasture of Area 3, to
which its seed was probably introduced as an impurity in other
grass seeds.*

Steglingia decumbens.
This grass was not recorded on Areas 1-4, but it formed 18-8
per cent. of the herbage of the old grass-heath of Area 8. It was

also found in all examinations of Areas 6 and 7, but on the former
area in relatively very small amounts.

LEGUMINOSAE,

In addition to the Percentage Frequency examinations, the
areas were examined for these plants by the Specific Frequency
method.

Trifolium pratense.

The seeds of a number of so-called varieties of this plant are
obtainable commercially, and it has been shown that not only
may these varieties differ considerably from each other in some

1Its seed is a common impurity of Festuca duriuscula and Cynosurus
cristatus seed (8), (II),

INDIVIDUAL SPECIES. 33

respects, but that even plants obtained from seed sold under a
similar name, if they have been produced in different countries,
may differ in behaviour when grown in this country (12).

At least four of these “varieties” have been used on the
farm on which the present areas are situated, and it is very
probable that these bought merely as “ Broad Red” or “ Red”
Clover seeds may have had different countries of origin.

Only on Areas 4 and 6 was a single variety sown, so that on
the other areas it can not be determined which variety may have
left persistent plants.’ This does not seem to be of very great
importance here however, as the results obtained are all very low.

Traces were recorded by the percentage frequency method
on Area 2 at 64 and 74 years, on Area 3 at $4 years, and on
Area 6 at 54 years, while on Area 6 also a P.c. fr. of o-1 was
recorded at 44 years. The only other Pyc. fr. result was 0-4 per
cent. on Area 4 at 134 years.

The Specific Frequency results are more definite, although
relatively few were obtained. After a seeding of ‘Broad Red”
and “Cowgrass” a Sp. fr. of 19 was obtained on Area 1 at 34
years, and a Sp. fr. of 1 at 44 years. With a heavier seeding, but
of a similar kind, the plant was not recorded on Area 2 at <$
years, but it had sp. frequencies of 2 and 3 at 84 years and ro}
years repectively, while a relatively light seeding of ‘ Broad
Red ” and “ Hybrid Cow Clover” gave a Sp. fr. of 2 on Area 3 at
7% years, and the same Sp. fr. was obtained on Area 5 at 104
years with a heavy seeding of “Broad Red” and ‘ Cowgrass.”
A Sp. fr. of 6 was found on Area 4 at 134 years.

From these results it appears that under these conditions the
development of the plant has two distinct periods. It decreases
rapidly from sowing, so that at 44 or 5$ years it has practically if
not quite disappeared. After this, itseems to increase slowly up
to 134 years at least.

It seems very probable that the second period of develop-

1These ‘‘varieties” can not be separated by botanical characters fram
each other, nor from the indigenous plants, in the vegetative state,

34 PASTURE STUDIES: SOME RESULTS.

ment is due to recolonisation by the indigenous variety, but this
has not been proved.’

The plant is probably completely exotic to Areas 6-8,
“ Cowgrass” was sown on Area 6 and on Area 7 at the rate of
alb, per acre, and on the latter area 2lb. per acre “ Late-flowering
Red Clover ” seed was also sown.

On Area 6, the plant was recorded in the P.c. fr. examina-
tions at 44 and at 51 years, but its development was very small,
as at 54 years it had only a Sp. fr. of 2. On Area 7, however, it
hada Sp. fr. of ro at 4$ years, so that in this situation, either
owing to the conditions or treatment or owing to the difference in
the varieties of seeds sown, it has persisted to a greater extent than
on Area 1,

Trifolium hybridum.

This plant is not indigenous to any of the areas, but its seed
was included in the seeds-mixtures for all except Area 8 and
possibly Area 4.

The results obtained show that its persisting powers are
generally small under some conditions. It was not recorded on
Areas 1-4 by the P.c. fr. method, and only to the extent of a
Sp. fr. of 2 on Area 1 at 34 years by the other method.

On Area 7 it was not recorded by either method, but on
Area 6 it was much more successful, as on the latter area it was
recorded by the P.c. fr. method at 44, 5 and 64 years, but not
at 74 years, and it also had a Sp. fr. of 15 at 54 years.

On Area 6, the plant did remarkably well throughout. It
produced a relatively large amount of herbage up to the fourth
year, but no determinations were made before 44 years. In the
third summer it produced ripe seeds quite freely in tufts surround-
ing cattle droppings, and it is possible that its relatively long
persistence on this area is largely due to this. In whatever way
it did persist, it did so on this area to a much greater extent than
Trofolium pratense, but on Area 7 it was much less successful than
the latter plant.

1On anold pasture field on the farm, a remarkable development of the
indigenous variety followed the application of Basic Slag.

INDIVIDUAL SPECIES. 35

Here again probably we see the effect of the other consti-
tuents of the seeds-mixtures, as the seeds of this plant were
included in equal quantities for the two areas,

Trifolium repens.

On an oat stubble following ley, the present writer has found
a Sp. fr. of 18 for this plant,? but it is not included in a list of
arable land weeds given by Stapledon (17). Brenchley (3)
obtained 131 seedlingsof this plant from the soil from four holes
(6in. x 6in. x gin) in a field which had been under grass for some
years.

The present writer collected ripe seed in 1916, and in April,
1917, a portion of these were treated with fine emery paper.
These, and others which had not been treated, were then placed
under similar germinating conditions, and the following germina-
tion results were obtained :—

1o days. 25 days. 2 mths, 4 mths. 6 mths. 8 mths,
Treated: i... (66? was GP ase OF ace BG. 86 sce 863
Untreated.e; J asx Fo as FF «ee TO ws WO ace 72

At the end of eight months the test on the treated seeds was
discontinued, and the untreated seeds were allowed to dry
gradually, and to remain in the dry condition for four months.
At the end of this time there were 81 per cent. hard+ seeds left,
while 7 per cent. had become mouldy.

The hard seeds were then collected and treated with emery
paper, but in this process a further 7 per cent. of the original
number were lost, leaving 74 per cent. for further testing,

In three days, 48 per cent. of the original number had
germinated normally, and 5 per cent. abnormally.s The test was

lAnglesey, poor soil at 150 feet.
2 Numbers in percentages of total seeds tested,
8 Some seeds had probably escaped treatment.

*7.e, seeds which appear normal and healthy although they have not
germinated under conditions which produce rapid germination of other
apparently similar seeds.

5This was probably due to the emery paper treatment,

36 PASTURE STUDIES: SOME RESULTS.

allowed to run for five months, and in the meantime a further 25
per cent. germinated normally, leaving only 1 per cent. which was
still hard.”

Thus seeds which were capable of immediate germination,
provided their coats were made permeable to water, remained
dormant for eight months under conditions which in other seeds
produced rapid germination, and after being allowed to dry for
four months were still capable of normal germination after treat-
ment with emery paper.

All these results leave little room for doubt that dead ripe
seeds of this plant may remain dormant in the soil for a very long
period, and still be capable of producing normal seedlings when
by some means their coats are rendered permeable.’ It is to be
expected therefore that where a pasture field in which there is a
fair development of the plant is brought under cultivation, the
plant may appear in the subsequent pasture in some quantity
although no seed be sown.

The seed of at least two varieties of this plant are obtainable
commercially. These are known as “ White or Dutch Clover”
( Trifolium repens) and “Wild White Clover” (Z. repens, var.
silvestre), (7), (20).2. The value of these two varieties of seeds in
the formation of a pasture has been examined by the writer, and
the results published elsewhere (9). These results showed that
on an average, with the same weight of seed, at 24 years from
sowing, 19-7 per cent. of the ground on which “Wild White
Clover” seed had been sown was covered by the plant, as com-
pared with 2-13 per cent. of the ground on which “ White or
Dutch” seed had been sown.

1 The conditions of the experiment are in some ways different from those
which occur in the seil. It is possible that soil water has a ditlereut effect
from tap water. Soil organisms (except animals) are not likely to have any
effect on them as the hard seeds of the experiment were quite immune from the
attacks of fungi.

“It is difficult to understand why the wild form shonld be regarded by

seedsmen as the variety, as it is very much more probable that the ‘‘ Dutch ”

is the variety, produeed by continued cultivation and unconscious selection of
the wild form,

~ INDIVIDUAL SPECIES. 37

Tn an experiment on the College Farm, Bangor, the follow-
ing results were obtained :—

Variety of seed: Dutch Dutch Wild None
Ib. of seed per acre : 2 I I —
Sp. fr. at 14 years: gI 98 99 85
Sp. fr. at 2 years: 97 100 100 86

These results are inconclusive as between the two varieties,
but they show clearly that even without seeding the plant may be
well distributed in a young pasture.

The Specific Frequency method is generally unsatisfactory as
a means of comparing the development of this plant, as even
where not strongly represented it may give the maximum Sp. fr.
This is clearly shown by the results for Area 2, where the plant
had a Sp. fr. of roo and a P.c. fr. of 12-4 at 54 years, anda Sp. fr.
of 98 anda P.c. fr. of 18-2 at 84 years. For this reason, the
P.c. fr. results chiefly will be made use of.

‘“ White or Dutch” seed was sown on Areas 1, 2, 3, 6 and 7,
but not on the other areas. That the prevalence of the plant in
the ensuing pasture at 54 years does not depend on the amount
of this seed sown is clearly shown by the results for Areas 1 and 2.
At this date the plant had a P.c. fr. of 31-8 on Area 1, where only
$b. per acre had been sown, while it had only a Pvc. fr. of 12-4
on Area 2 from 2lb. seed per acre. Since the soil, altitude, and
grazing-are similar on the two areas the cause of this difference
must be soughtelsewhere. It may partly be due to the difference
in the other components of the seeds-mixtures, but this does not
seem probable. It may also be due to the high development of
Cynosurus cristatus and Anthoxanthum odoratum on Area 2.
The most probable cause, however, is the difference in treatment
—partly the treatment while under cultivation, and partly the
subsequent treatment, and especially that Area 2 was given a
dressing of farmyard manure after one hay crop had been taken,
while Area 2 was not manured until the winter following this

examination.

1Jtis believed also that the phosphatic manure applied to Area 1 when
sown down to grass was Basic Slag, while Superphosphate was used for Area 2.

38 PASTURE STUDIES: SOME RESULTS.

The sudden increase of the plant on Area 2 at 64 years was
undoubtedly due to the application of Basic Slag (16), (13), but
the effect of the Slag was not very lasting here, as at 84 years the
plant was almost equally well developed on Area 3, which had not
been slagged. On this latter area, however, the relatively high
development of the plant may be due to the thorough cleaning of
the land, but at 9} years it was only slightly more abundant on
Area 2 than on Area 4 at 134 years, which had not been slagged,
and on which the plant had not been sown, so that apparently the
effect of the Slag on this plant was exhausted in three years.

The plant was not recorded by either method of examination
on Area 8, but it is known that weak plants were to be found in
the former grass heath of Area 6. ‘‘ White or Dutch” seed was
sown on Areas 6 and 7, but the plant had a P.c. fr. of only 8-4 on
the former area at 44 years. Following a dressing of Basic Slag,
however, it increased to 27-3 in the following year, and at 64
years it still had a P.c., fr. 24-7. Its decrease was then as rapid as
its previous increase, probably to a great extent owing to the
exhaustion of the effect of the Slag, but partly also probably to the
effect of the drought! The same quantity of seed was sown on
Area 7, but this area was not examied by the P.c. fr. at 44 years.
The Sp. fr. of the plant at that date was relatively low (sp. fr. = 60).
and its P.c. fr. at 54 years, following a dressing of Basic Slag as on
Area 6, at the corresponding date, was very much lower than on
the latter area, so that here again apparently the effect of the
seeds-mixture as a whole is seen.

From these results it is seen that the amount of ‘“ White or
Dutch” seed sown may bear no relationship to the development
of the plant at 54 years, and that at 134 years at most, its inclusion
has no apparent effect on some soils. Its development in the
pasture appears to be very largely a matter of cultivation, manur-
ing and treatment, but other results (9) show that “ Wild White
Clover” seed may hasten the development of the plant very
considerably in a pasture.

1Jt has been found elsewhere that heavy grazing with sheep has no such
effect on the plant. Cf. Stapledon on the effect of drought (15).

INDIVIDUAL SPECIES, 39

Anthyllis Vulneraria.

This plant is not indigenous to this farm, but, as shown by
Table I., the seed has often been included in seeds-mixtures in
recent years. It was first introduced on Area 2 in 1908, but at
the rate of only fb. per acre. On this area rare plants were to be
found up to 24 years, but on Areas 1 and 2, where it was also
sown, it was not found after 14 years.

On Area 6, it gave remarkable results in the hay crop at
1} years from a seeding of 2lb. per acre, but it had almost
completely disappeared at 24 years except on the thinnest soil,
and even here it was not recorded in P.c. fr. examinations at 44
years and subsequently, nor in the Sp. fr. examination at 54 years.
It also developed fairly well up to 14 years on Area 7, but it then
disappeared, and was not recorded in the detailed examinations of
this area,

Medicago sativa (Exotic).

This was sown on Areas 6 and 7, but it made a very poor
development, and was not recorded in the examinations.
Medicago lupulina (Exotic).

This was sown on Area 5, but was not recorded in the Sp. fr.
examination at 10} years.

Trifolium dubium (Not sown).

The seed of this plant has never been intentionally sown on
the farm, but it often occurs as an impurity in seed sarnples of
Trifolium repens and TJ. hybridum (8), (II). The plant is some-
times plentiful in a well-manvred permanent hay field on the
farm, but otherwise it is not very noticeable.

On the present areas its highest P.c. fr. was 2-6 on Area 6 at
6} years, but by the following year its P.c. fr. on this area was
only o-g. Its highest Sp. fr. was 4 on Area 1 at 4 years, and on
Area 6 at 54 years.

Lathyrus sp, (Not sown).

Traces of this genus were found on Area 2 at 6}, 74, 84 and
gd years, but it was not otherwise recorded,

40 PASTURE STUDIES : SOME RESULTS.
Lotus spp. (Not sown).

L. corniculatus was the only species recorded on Areas 6-8,
but traces of Z. udéginosus were also found on Areas 1-5. The two
species were not separated in these examinations.

The Percentage Frequency method of examination is net
very satisfactory on young pastures for these plants owing to their
habit of frequently occurring in colonies. The Sp. fr. method is
probably more reliable, as the plant seldom reaches a high Sp. fr,,
and therefore its relative distribution is a good guide to com-
parison. The results obtained by the P.c. fr. method are however
given in Table V., p. 16d.

The highest P.c. fr. for Areas 1-4 was o-8 on Area 2 at of
years, and on Area 4 at 134 years.

The genus was not recorded on Area 1 by the Sp. fr. method,
but the other records for these areas ranged from a Sp. fr. of 2 on
Area 3 at 7} years, to 37 on Area 4 at 13} years. The results
for Area 2 are irregular, as the Sp. fr. was lower at 84 years than
at 54 years, but it was higher at 104 years than at 54 years. On
this area, therefore, there appeared on the whole to be a tendency
to increase, but not to the same extent as on Area 4. The
difference between Areas 1, 2, 3 and 5 are capable of explanation
by reference to the treatment of the land while under cultivation,
together with the assumption that the tendency to increase is real
on these areas, as Area 2 was not undera cleaning crop. But
the high Sp. fr. on Area q is difficult to explain as this area was
also partly under a cleaning crop. This area had not been
manured, however, since it was laid down to grass, but Basic Slag
does not usually depress these plants (23). It may be that the
difference between Areas 2 and 4 is merely an indication that in
the later history of a young pasture these plants develop very
rapidly.

The highest P.c. fr. and the highest Sp. fr. were obtained on
the old grass-heath (Area 8). On both Area 6 and Area 7 the
P.c. fr. results indicate a decrease with an increase in the age of
the pasture. It is very probable that this plant is capable of

INDIVIDUAL SPECIES. 4t

surviving cultivation bodily,t and that on Area 6 especially it was
less suitable for the new conditions created, and therefore suffered
in competition with other plants. The Sp. fr. for the plant on
this area at 54 years was very low considering that the area had
not been under a cleaning crop, and the difference between this
and the Sp. fr. for Area 7 at 44 years is very marked. In respect
of this plant, Area 7 was again reverting much more rapidly than
Area 6 towards the grass-heath condition of Area 8.

OTHER ORDERS.*

The plant unit in most of the species included under this
heading is relatively large,? and as they rarely form a very large
proportion of the herbage the P.c. fr. results are generally low.
In most cases, therefore, the Sp. fr. method is more satlsfactory,
and the discussion of these species will be mainly based on the
Specific Frequency results.

Results were obtained for a few species which are not
included in the tables owing to the fact that their development
was very small. Some of these were sown species.

Cichorium Intybus (Exotic).

The seeds of this plant were included in the seeds-mixtures
for Areas 6 and 7 at the rate of 2lb. per acre.

It had a P.c. fr. of o-r on Area 6 at 44 years, but otherwise
was not recorded by this method. It bad Sp. fr. of 3 and 2 on
Areas 6 and 7 at 5} and 44 years respectively, so that although
not indigenous, it had persisted to some extent for over five years,

Poterium Sanguisorba (Exotic).

This was sown on Areas 6 and 7 at the rate of 3lb. per acre.
It was not recorded in the P.c. fr. examinations, but it had Sp.
frequencies of 3 and 2 respectively on these areas, the former
area examined at 54 years and the latter at 44 years.

1 This is suggested by a comparison of Areas 2 and 3.
*See Table V., p. 16%
% Definition i,, pr Is

42 PASTURE STUDIES: SOME RESULTS.

Carum Petroselinum (Exotic).
This was also sown on Areas 6 and 7 at the rate of 2lb. per
acre, but the plant was not recorded in the examinations.

Plantago lanceolata.

The seed of this plant has often been included in seeds-
mixtures on this farm (Table I., p. 4), and it was included in those
for Areas 3, 5 and 7, and also probably in that for Area 4. Its
seed is also often found in samples of Z7ifolium pratense secd (8),
(II), (14), so that probably a small amount would be introduced
in this way.

Its Sp. fr. was quite low on Area x both at 34 and at 4}
years, and in spite of the fact that it had been sown on Area 3 its
Sp. fr. was considerably lower on this area than on Area 2, where
it had not been sown. On Area 5, where also it had been sown,
its Sp. fr. was only 23 at rod years, as compared with 27 on Area 2
at the corresponding period. Its extreme development, however,
was on Area 4, where it had a Sp. fr. of 81 at 134 years.

Stapledon found this plant to occur in some quantity as an
arable land weed, and in considerable abundance in pastures of
less than a year old (‘‘seeds”) (17), but Brenchley found its seeds
quite unimportant amongst buried weed seeds (3). It is quite
possible, therefore, that the relative abundance of the plant on
Area 2 is due to the imperfect cleaning given to the land. It is
surprising, however, to find that the seed sown had been relatively
so ineffective on Areas 3 and 5.

On Area 7 a seeding of 3lb. per acre was considerably more
effective, as here there was a Sp. fr. of 26 at 44 years as compared
with a Sp. fr. of 6 on Area } at 54 years, where the seed had not
been sown. As shown by a Sp. fr. of ro for Area 8 (old grass-
heath), the plant was more or less indigenous to these areas.?

Achillea Millefolium.

The seed of this plant was not sown on any of the present
areas, and it had not been sown on the farm until some time after
these areas had been laid down to grass. It is not included by

'Qn Area 8 it was chiefly found along sheep tracks.

INDIVIDUAL SPECIES. 43

Stapledon in his list of arable land weeds (17), and Brenchley
shows that its seeds rarely lie dormant except in the soils of very
old grass land (3). It was not recorded by the present writer on
an oat stubble, and the following results obtained on well cleaned
land’ at 50 feet also show that the plant may not occur where not
SOWN :—

Seed sown in lb. per acre: o ¥% Yy %
Sp. fr. at 1} years: ° 9 23 27
eo 2 wt © 145 335 35

On the other hand, the following results obtained on
imperfectly cleaned land,? show that it may occur in some quantity
even in young pastures, and that in some such cases the inclusion
of seed in the seeds-mixture may have no appreciable effect :—

Seed sown in lb. per acre: nil yy

Sp. fr. at rd years: 6 4
i Sp ES ay 2 8
we cg 2 4s 8

On the whole, these results tend to show that the plant does
not usually persist through a period of cultivation which includes a
cleaning crop, but that where the land is imperfectly cleaned,
especially at relatively high elevations, it may persist by some
means. Its obvious success, at least in some cases, when sown
on well-cleaned land, shows that although it may not appear
spontaneously in the young pasture, the conditions in those cases
are not such as to inhibit its growth.

In view of this, the results for Apeas 1-8 are especially inter-
esting. The plant had a Sp. fr. of ro on the old grass-heath
(Area 8), but it was not recorded on Area 7 at 44 years, nor on Area
6 at 54 years. In this case, therefore, it had been completely

1 These are average results for four plots each over three acresin extent,
and six plots each a fourth of an acre in extent, all in the same field, at the
College Farm, Aber, Bangor.

2Penmachno, 775 feet, The two areas are the two halves of the same
field,

44 PASTURE STUDIES: SOME RESULTS.

eradicated, although no cleaning crop had been taken. On
Area 1, which had been partly under a cleaning crop, it had a

Sp. fr. of only 2 both at 3} years and at 4% years. It is rather
surprising, therefore, to find that on Area 3, which had been

entirely under a cleaning crop, it had a Sp. fr. of 41 at 7k years.
In fact, it was probably only slightly less prevalent on this area
than on Area 2, which had not been even partly under a cleaning
crop. The development of the plant on Area 2 was, however,
very rapid. There isa very marked difference between this area
at 54 years and Area 1 at 44 years, but this would naturally be
assumed to be due to the difference in cultivation were it not for
the results for Area 3, and the results for Areas 4 and 5 would
not contradict this conclusion as each of these had been partly
under a cleaning crop.

If we assumed that for some unknown reason? the develop-
ment of the plant on Area 3 was abnormal, the other results would
be straight-forward. They would show that under the conditions
prevailing for Areas 1-5 the plant is strongly indigenous, and that
although the land may be put partly under a cleaning crop the
plant may be very prevalent on relatively young pastures, but that
where the land is not even partly put under a cleaning crop its
development is very rapid, its Sp. fr. rising from 39 at 5$ years to
76 at 104 years.

On Area 3, however, the effect of a thorough cleaning was
exceedingly small.

Under the conditions prevailing on Areas 6-8 on the other
hand, its appearance may be delayed very considerably even
though the land is not put under a cleaning crop.

Other results mentioned show that in some cases at least the
inclusion of the seed in the seeds-mixture may be very effective
where the land is thoroughly cleaned, and that with low quantities
of seed, the distribution of the plant in the young pasture bearsa
definite relationship to the amount of seed sown. In some cases,
however, the inclusion of the seed may be ineffective.

} The seed of the plant never occurs in any appreciable quantity in other
seed samples (8), (II), (14).

INDIVIDUAL SPECIES. 45
Ranunculus repens.

Stapledon found this plant to be one of the most important
arable land weeds (17), and Brenchley classifies it amongst
“arable or grass land plants ” (3).

It was not recorded on Area 8 (old grass-heath), but it was
recorded on all the other areas by both methods of examination.
On Areas 6 and 7 it was, however, much less important than on
most of the other areas.

The P.c. fr. results for Areas 1-4 on the whole indicate that
itis most prevalent in the young pasture under the prevailing
conditions, and that it decreases with the increasing age of the
pasture. The results for Area 3, however, are very high as com-
pared with those for Area 2 at corresponding periods.

This greater development of the plant on Area 3 is also
apparently indicated by the Sp. fr. results, as on this area the
plant had a Sp. fr. of 78 at 74 years, as compared with a Sp. fr. of
62 on Area 2 at 84 years. The latter result may be slightly
abnormal however, as on the same area there was a Sp. fr. of 70 at
rod years.

The Sp. fr. results for Areas 1-5 as a whole seem to indicate
that the plant increases up to 44 or 54 years, and then decreases
fairly rapidly. The very low Sp. fr. on Area 4 may be due to
competition with other plants, especially vantago lanceolata,
which had a very high Sp. fr. on this area. If anything, these
results tend to show that a thorough cleaning of the land is
favourable to the development of the plant, but there is little to
show to what extent it is affected by the composition of the seeds-
mixture used.

Bellis perennis.

This plant may also occur in considerable quantity as an
arable land weed (17).

It was not recorded on Area 8, and its development on

\The seeds of the remaining plants are never intentionally included in
seeds-mixtures, but those of some of them are frequently found in other seed
sdtaples (8), (2), (44).

46 PASTURE STUDIES: SOME RESULTS.

Areas 6 and 7 was relatively very small, as on these areas its

highest Sp. fr. was 18 on Area 6 at 54 years.
It was, however, very widely distributed on the other areas,

and gave considerably higher P.c. fr. results than Ranunculus
repens.

The results for Areas 1, 2, 4 and 5 are, on the whole, fairly
regular, and from these it would appear that the plant increases
up to 54 years or perhaps even somewhat later, but then decreases
fairly rapidly. The results for Area 3 are different, and they show
that the plant may have a Sp. fr. very little short of the maximum
even at 74 years, and a very high P.c. fr. at 9} years. It is
difficult to find any cause for this difference except that Arca 3
had been more thoroughly cleaned than the other areas while
under cultivation.

Prunella vulgaris.

This is regarded by Brenchley as a grass land plant (3), but
Stapledon found it to occur frequently as an arable land weed,
and especially in pastures less than a year old (17). Its seed is
fairly frequently found in other seed samples, especially in those
of Zrifolkum hybridum aud T. repens (8), (11), (14).

Tt had a Sp. fr. of 7 even on Area 8 (grass-heath), but on
Area 6, at 54 years, it had a Sp. fr. of 54, while on Area 7 (which
has been found to be approaching the grass-heath condition at a
more rapid rate than Area 6 in other respects) it had only a Sp. fr.
of 22 at 44 years.

On Areas 1, 2, 4 and 5 the results are again fairly regular,
showing a maximum development at about 54 years. The Sp. fr.
results for Area 3 agree fairly well with the others, but the P.c, fr.
results for this area and for Area 4 are altnormally high.?

Cerastium vulgatum.

This plant was recorded in all examinations except on
Area 8, but its development was small on other areas. On Areas

1 These areas were all examined, August - September in each year, and at
this time there are usually a fairly large number of seedlings of this plant. The
Pre, fr. results, therefore, depend very largely on the number of seedlings
produced,

INDIVIDUAL SPECIES. 47

1-4 it was much more widely distributed, and had a much higher
P.c. fr. up to 54 years than later. Different from species already
mentioned, this plant decreased very rapidly in most cases, so that
on Area 2 its Sp. fr. had decreased from 67 at 5} years to 4 at 84
years. Its Sp. fr. on Area 5 at 104 years was relatively very high
as compared with the other areas. This may be due to the
slightly lighter soil.

Sagina procumbens.

Owing to the habit of this plant, it is not easy to detect it
where the turf is fairly thick and the plants small, so that the
P.c, fr. results may be more reliable than those obtained by the
Sp. fr. method.

The highest P.c. fr. was obtained on Area 2 at 7} years, and
apparently, on Areas 1-5 it reached its maximum at about this
time, and then decreased fairly rapidly.

It was not recorded on Area 8, and on Areas 6 and 7 it was
but poorly represented except on Area 6 at 64 years. In respect
of this plant Area 6 appeared to be reverting to the heath con-
dition more rapidly than Area 7.

Luzula campestris.

Few records were obtained for this plant by the Sp. fr.
method. Its highest P.c. fr. was obtained on Area 8 (grass-
heath), but as compared with other areas, especially Area 6, at the
corresponding periods, it was well represented on Area 7 both at
54 and 64 years. Only traces were recorded on Area 6, so that
in respect of this plant Area 7 was reverting to heath condition
most rapidly.

The plant was not recorded on Area 1 at 44 years nor on
Area 3 at 84 years, andit -was but poorly represented on these
areas a year later in each case.

The results for the other areas seem to show that on
this land it tends to increase with the increasing age of the
pasture, although its P.c. fr. was lower at 104 years than at of}

1Qn old pastures examined by the writer, its highest P.c. fr. was 1°3 on
a soil inclined to be gravelly or sandy (College Farm, Bangor).

48 PASTURE STUDIES: SOME RESULTS.

years on Area 2, as on Area 4 at 134 years it had a very high
percentage frequency.

Potentilla erecta,

As shewn by a Sp. fr. of 87 on Area 8, this is a typical grass-
heath plant.t It is recorded as an abundant arable land weed in
exceptional cases by Stapledon (17), and results obtained by the
present writer on land which had been brought under cultivation
from a rough grass heath at an elevation of 150 feet in Anglesey,?
shows that where the land is not thoroughly cleaned it may occur
in considerable quantity both in the stubble and in the young
pastures. These results were as follows :—

(t) Sp. fr. on pasture 1% yearsold: 66
(2) “ % 2% ‘3 46
(3) ‘i - 2 i 60
(4) stubble following (3) 46

In these cases, the plants have in most cases undoubtedly
survived the period of cultivation bodily, as shown by their strong
root-stocks. It is net surprising, therefore, to find the plant fairly
widely distributed in most of the present areas.

On Area r it was not recorded except in very small quantity
at 34 years, and on Area 3, which had been thoroughly cleaned,
it was net recorded in the Sp. fr. examination at 7} years, and
only to a very slight extent by the P.c. fr. method at 9} years.
Even on Area 2, which had not been partly under a cleaning
crop, it only had a Sp. fr. of 3 at 54 years, but this had increased
to 17 at 84 years, and the same Sp. fr. was again found at 10}
years, while on Area 4 adjoining, its Sp. fr. was also 17 at 134
years. On Area 5 at rod years, however, it was much less widely
distributed.

Thus, on this type, the plant probably survives a period of
imperfect cultivation to some extent, and then increases fairly
slowly, but its rate of increase becomes much less when it has
attained a Sp. fr. of about 17, and therefore in respect of this

tof. (21).
? Ceidio ‘Isaf, Llanerchymedd,

INDIVIDUAL SPECIES. 49

plant these conditions do not immediately produce the typical
grass-heath.

On Areas 6 and 7 the increase of the plant was much more
rapid, but in this case again Area 7 was reverting to the grass-
heath type much more rapidly than Area 6,

Leontodon autumnats.

This plant is obviously not a grass-heath plantas it was not
recorded on Areas 7 and 8, and only in minute quantities on
Area 6.

It was of frequent occurrence on Areas 1-5, and on Area 4
it reached a Sp. fr. of 65 at 134 years. On Areas 3 and 5 its
increase was less rapid than on Area 2.t Qn the whole it would
seem that the plant is capable of surviving a period of imperfect
cultivation, and that under the conditions prevailing, its distribu-
tion in the pasture extends with the age of the pasture.

Senecio Jacobaea.

The development of this plant in a pasture is usually inter-
fered with by cutting, or sometimes by grazing with sheep. It
never flourishes where sheep graze during the summer.

It was of frequent occurrence on most of the present areas,
but its development had in most cases been interfered with. On
Area 5 for instance, it was not recorded, although it is known to
" have been present in some quantity before the area was grazed
with both cattle and sheep during the summer. Up till 1918
Areas 6 and 7 were only grazed by sheep in the spring, so that
the development of the plant had been little interfered with. The
plant was recorded at each examination—reaching a Sp. fr. of 18
at 54 years—on Area 6, but it was not recorded on Area 7, so that
in respect of this plant again Area 7 was nearer the heath type
than Area 6.

Areas 2 and 3, since they were parts of the same field, were
also subject to the same grazing conditions, but here the plant
was obviously much more prevalent on Area 3 than on Area 2.

1In the Sp. fr. of this area at 54 years the plant was confused with
Taraxacum vulgare, so that the results are not given in Table V., p. 164,

50 PASTURE STUDIES: SOME RESULTS.

This may be connected with the different treatment given to the
two areas while under cultivation.

Rumex Acetosa,

This was not recorded on Areas 6, 7 and 8, and only in small
quantities on the other areas. It seems to be most prevalent on
the young pastures, but the highest Sp. fr. occurred on Area 5 at
rod years. This is interesting in view of the fact that this was
the only area recently limed, as it is generally understood that
“lime applications tend to check its growth” (5).

Centaurea nigra.

This plant is recorded by Stapledon as an arable land weed
at high elevations in Mid-Wales (17), and it is likely that where
it is fairly strongly indigenous it may survive a period of imperfect
cultivation, as the writer has found it to have specific frequencies
of 4, 8 and 8 at 14, 2 and 24 years respectively under such con-
ditions.*

It was not recorded on Area 8, and only in minute quantities
on Areas 6 and 7, but on the other areas it was more plentiful.
The effect of thorough cleaning of the land is probably seen in
the relatively low Sp. fr. of 7 on Area 3 at 74 years as compared
with specific frequencies of 14 and zo at 84 and 104 years respec-
tively on Area 2, It had but a low Sp. fr. on Area 5 at 10} years,
but it is generally believed that it is kept in check by grazing with
sheep. Is is obviously a more important constituent of the
herbage of the older than the younger pastures under the condi-
tions prevailing for Areas 1-5.

Plantago major.

This was but poorly represented, but it is shown to be at its
highest development on the younger pastures.
Rumex Acetosella.

Although not recorded on Area 8, this plant had high specific
frequencies on Areas 6 and 7. On these areas, it was a trouble-
some weed while the land was under cultivation, but the P.c. fr,

1 Penmachno, 775 feet.

INDIVIDUAL SPECIES, 51

results show that it was decreasing in the pasture. This decrease
had not proceeded very faron Area 7 at 44 years, but the plant
was less widely distributed on Area 6 at 54 years.

Except on Area 1, where it had specific frequencies of 7 and
10 at 34 and at 4} years respectively, the plant was rarely
recorded on the other areas.

Potentilla anserina.

This was chiefly met with on Area 2, where its maximum
Sp. fr. was 24 at 84 years. This seems to be abnormally high, as
on the same area its Sp. fr. was 8 and 9 at 54 and ro years
respectively,

It was not recorded on the slightly lighter soils of Argas 5, 6,
7 and 8.

Taraxacum vulgare.

Traces of this plant were found on Area 6 at 74 years, but
otherwise it was not recorded on Areas 6,7 and 8. Its chief
development was on Areas 2 and 4, and on Areas 1-4 its Sp. fr.
seemed to be increasing with the age of the pasture.

Chrysanthemum Leucanthemum.—This was chiefly recorded
on Area 2, and it appeared to be decreasing with the increased
age of the pasture.

Polygala vulgaris—This plant was recorded in minute
quantities on Area 2 at 74 years and later, and on Area 4 at 134
years, but otherwise it was not recorded except to a small extent
on Area 6 at 54 years, and on Area 8, where it hada Sp. fr. of
to. It is obviously a plant of the older pastures and the grass-
heath.

Veronica officinalis—This was also recorded on Area 2 in
minute quantities at 84 years and later, and also on Area 4, It
had a Sp. fr. of 22 on Area 8, and it was represented on Areas
6 and 7. On these areas, it seems to come in sooner than
Polygala vulgaris, but it is primarily a grass-heath plant.

Viola ¢anina,—This is also a grass-heath plant, but it was

52 PASTURE STUDIES: SOME RESULTS.

recorded on Area 2 at $4 years (Sp. fr.=1). It had a high Sp.
‘fr. on Area 8, but it was not recorded on Areas 6 and 7.

Hieracium Pilosella was recorded on Areas 3, 6, 7 and 8,
but on the former three areas in very small quantities.

Euphrasia officinalis was only recorded on Areas 4 and 8,
and it had a considerably higher Sp. fr. on the former than on the
latter area,

Thymus Serpyllum.—This grass-heath plant had a Sp. fr.
of 55 and a P.c. fr, of 4-2 on Area 8, and minute quantities were
found on Areas 6 and 7.

Galium hercynicum is another typical grass-heath plant, and
it was recolonising Area 7 at a more rapid rate than most of these
plants, but it was recolonising Area 6 much more slowly.

Calluna vulgaris had a fairly wide distribution on Area 8,
although, as shown by the P.c. fr. results, it formed but a very
small proportion of the total herbage. It was also recorded on
Areas 6 and 7, but its Sp. fr. was small.

The following species were also recorded, but in very small
quantities :—

Hypochoeris radicata (Areas 1, 2, 6, 7), Veronica serpyltifolia
(Areas 2, 3, 5), V. Chamaedrys (Areas 1, 3, 5, 7), Centaurium
umbellatum (Areas 1, 2), Ulex europaeus (Area 8), Vaccinium
Myrtillus (Areas 6, 8), Afentha sp. (Areas 1, 2, 3), Pterts aguilina,
Anagallis arvensis (Area 2), Hypericum humifusum (Areas 7, 8),7
Cirsium arvense (Areas 1, 3,5, 7)

SUMMARY AND CONCLUSIONS.

The botanical composition of the herbage of pastures on two
types of soil has been studied in considerable detail with respect
to the history of the individual species by the Percentage
Frequency and the Specific Frequency methods. Five areas

1On Area 8, on the sites of burnt gorse bushes,

SUMMARY AND CONCLUSIONS. 53

have been investigated, three of them several times in consecutive
years, on one type, and three areas on the other type. The
history of each area has been given, and also in the case of six of
the areas the chief details of the seeds-mixtures used in laying
them down to grass (Table IV., p. 16a),

Some variation has been found in the behaviour of the same
species on different areas on the same soil type, and considerable
differences on the different soil types.

In drawing conclusions for the two area groups it is necessary
to bear in mind not only that they are not comparable in soil, but
that their history is also quite different. Areas 1-4 have been in
regular cultivation on the long rotation system, so that relatively
short-lived plants would tend to be perpetuated, while more
permanent plants, unless capable of surviving cultivation, would
be eradicated or would have no chance of recolonising the ground
unless especially adapted. These areas are also in constant
communication with other parts of the farm, and particularly with
the farm buildings, as they are largely grazed by cattle, so that
although perhaps not actually manured they are still affected by
their position. Even Area 5, which had been under grass for a
long period prior to the rotatioa during which it was examined,
would have been affected by its proximity to the farm buildings,
and all these areas are practically surrounded by land treated in
the same way.

Areas 6 and 7, on the other hand, had only been through
one period of cultivation since they had been for a long period in
the grass-heath condition. They were also remote from the farm
buildings, and formed part ofa sheep-walk. There was practically
no communication between them and the land under regular
cultivation, and they were almost entirely* surrounded by grass-
heath or by uncultivated land. In this case, therefore, plants
which are short-lived under the grass-heath conditions would have
disappeared, and even when the land was brought under cultiva-
tion they would not easily be re-introduced, This is probably why

1 Entirely except for a few yards where Area 7 joins the cultivable part ef
the field in which Area 4 is situated,

64 PASTURE STUDIES: SOME RESULTS.

such plants as Ranunculus repens, Bellis perennis and Cynosurus

cristatus were so poorly represented on Area 7, and in the case of
the two former plants, on Area 6.

THE EFFECT OF ALTITUDE, SLOPE, ASPECT, ETC.

Since, as shown above, the two groups of areas differed very
greatly in what appear to be more important respects, the
influence of these factors. can not accurately be estimated. The
most marked differences that may be traceable to these are, that
Festuca rubra is very prevalent on Areas 1-4, but negligible on
Areas 6-8, and that both Agrostis alba and A. tenuis occurred on
the former areas and only 4. ¢enwzs on the latter.

THE EFFECT OF CULTIVATION,

Areas 6 and 7 were similarly treated throughout their period
of cultivation, so that the effect of this can not be seen in their
case. At the same time, it is very difficult to believe that the
great difference between the herbage of the two areas was entirely
due to the difference in the composition of the seeds-mixtures,
but there is no evidence te the contrary.

From the results for these areas, one point in particular
becomes prominent, viz., that the inclusion of a cleaning crop is
not necessarily the only means of eradicating undesirable plants.
As will be seen from a comparison of the results for Areas 6 and
8, the disappearance of the grass-heath plants from Area 6 was
remarkable. In particular, the suppression of Agrostis tenuis may
be noted, as this grass is generally considered to be perhaps the
most difficult of all toget rid of. Thus, although the writer has
several times used the words “imperfectly cleaned” to indicate
land which had not been under a cleaning crop, the distinction
does not seem to be absolute.

Areas 1, 2 and 3 were not cultivated in exactly the same
way. Even Area 2, however, although it had not been under a
cleaning crop, had been carefully cleaned as far as growing cereal
crops in succession allowed. On these areas, some results, such
as the greater promineace of Festuca rubra, and the occurrence of

SUMMARY AND CONCLUSIONS. 58

particular plants on Area 2 in greater abundance than on other
areas, and the smaller proportion of Agrostis spp. in the early
history of Areas 1 and 3, and also possibly the greater prevalence
of some miscellaneous plants such as Ranunculus repens, Bellis
perennis, etc., on Area 3, have been considered as showing the
effect of different forms of cultivation.

THE EFFECT OF AFTER-TREATMENT.

Ithas been suggested that this might have been to some
extent the cause of the difference between Areas 6 and 7, but on
consideration, the suggestion has not been found tenable. A
difference in manuring has left some distinct results, but on these
areas it did not affect any species very materially except Z7zfolium
repens, and even in the case of this plant the effect was only
temporary.

THE EFFECT OF SEEDS-MIXTURES.,

On Areas 1-4 few marked differences traceable to this cause
were met with. A difference in the quantity of the seed of some
species, such as Dactylis glomerata, seems to have had some
lasting effect, but in the case of other plants, such as Lolium
perenne, the effect was generally at most temporary. The
inclusion of the seeds of some plants which are for some reason
exotic to the areas on which they were sown had some marked
results. Thus, although it did not change the character of the
herbage to any appreciable extent, Festuca e/atior persisted for at
least 9$ years on Area 2. The most marked results were
obtained, however, by the inclusion of the seeds of Cywosurus
cristatus and Anthoxanthum odoratum in the seeds-mixture for
Area 6.

On the other hand, the seeds of some exotic plants, such as
Festuca duriuscula and Arrhenatherum elatius, were practically
useless,

It is quite obvious from these results that the seeds-mixtures
used on Areas 1-4 and on Area 7 at least were not successful
after five or six years, It is true that some desirable plants madg

56 PASTURE STUDIES: SOME RESULTS.

their appearance in some quantity at this time, but a large pro-
portion of undesirable plants also appeared, and taking Area 2
as an example, it would seem that the seeds,Cynosurus crsistatus
and Poa trivialis at least should also have been included in the
seeds-mixture, so that they might not start at a disadvantage in
competition with other plants. It is probable also that the
inclusion of a greater quantity Festuca elatior and (or) its variety
£. elatior var. pratensis would give good results. On the other
hand, Féstuca duriuscula might have been excluded, and also
possibly Poa pratensis, Trifolium repens and Anthyllis Vulneraria.
In this case, it would seem to be quite unnecessary to add
Achillea Millefolium, but it might apparently have been included
with advantage in the seeds-mixtures for Areas 6 and 7, while
Carum Petroselinum and Arrhenatherum elatius might have been
omitted.

REFERENCES.

(t) ARMsTRoNG, S. F.: The Botanical and Chemical Composi-
of the Herbage of Pasture and Meadows. Jour. Agr. Sci.,
vol, ii, 1907.
(2) AnmstRonG, S. F.: British Grasses and their Employment
in Agriculture. Camb. Univ. Press, 1917.
(3) BRencHLEY, W. E,: Buried Weed Seeds. Journ. Agr. Sci.,
vol. ix., 1918.
(4) Crowruer, C., and Ruston, A. G.: The Influence on Crop
and Soil of Manures applied to Permanent Meadow.
Lb. vol, vii. 1915-16.
(5) Cyclopedia of Modern Agriculture, The Standard: Gresham
Publ. Co., London, 1911.
(6) Hatt, A. D., and Russet, E. J.: On the Causes of High
Nutritive Value and Fertility of the Fatting Pastures of
Romney Marsh. /ourn. Agr. Sci., vol. iv., 1911-12.
(7) Hunter, James: 35¢h Price List of Agricultural Seeds
Chester, 1917.
(8) Jenkin, T. J.: Seed-Testing and Report on Seeds Tested in
1913-15. Univ. Coll. of N. Wales, Bangor, 1916.
(9) Jenkin, T. J.: Ordinary White Clover Seed v. Wild White
Clover Seed. Journ. Bd. Agr., vol. xxili., March, 1917.
‘,, (10) Jonss, A. E., and StapLepon, R. G.: The Improvement of
Upland Pastures. Univ. Coll., Aberystwyth, 1916.

(11) Official Seed-Testing Station: First Annual Report. Journ,
Bd. Agr., vol. xxv., Sept., 1918.

(12) Report on Nationalities and Varieties of Red Clover. Univ.
of Leeds and Yorkshire Council for Agr. Educ., Bulletin
57, 1906.

(13) SOMERVILLE, WiLL1aM: Influence on the Production of
Mutton of Manures applied to Pastures. Journ, Bd, Agr.
Supplement, vol, xvii., Jan., 1911,

58 PASTURE STUDIES: SOME RESULTS.

(14) STAPLEDON, R. G.: Report on the Condition of the Seed
Trade in the Aberystwyth College Area. Univ. Coll.,
Aberystwyth, 1914.

(15) StaPLepon, R. G.: Pasture Problems: Drought Resistance.
Journ. Agr. Sct., vol. v., 1913.

(16) StapLepon, R. G.: Pasture Problems: The Response of
Individual Species under Manures. JZ, vol. vi., 1914.

(17) StapLepon, R. G.: On the Plant Communities of Farm
Land. Anais Bot., vol. xxx., 1916.

(18) Staprepon, R. G., and Jenxin, T. J.: Pasture Problems:
Indigenous Species in Relation to Habitat and Sown
Species. Journ. Agr. Sci., vol. vili., 1916.

(19) Sutron, Martin J.: Permanent and Temporary Pastures.
London: Simpkin, Marshall. Popular Edition, 1902.

(20) Sutton’s Farmers’ Year Book. Reading: Sutton & Sons,

1919.
(21) TansLey, A. G.: Zypes of British Vegetation. Camb. Univ.
Press, 1911.

(22) Yapr, R. H., Jouns, D., and Jones, O. T.: The Salt
Marshes of the Dovey Estuary. Journ. L£cology, vol. iv.,
1916, and vol. v., 1917.

(23) Jenkin, T. J. : Unpublished Results.