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393

RELATIVE GROWTH-RATE: ITS RANGE AND ADAPTIVE
SIGNIFICANCE IN A LOCAL FLORA

BY J. P. GRIME AND RODERICK HUNT

Unit of ComparativePlant Ecology (N.E.R.C.), Departmentof Botany,
The University,Sheffield S10 2TN

INTRODUCTION
The needfor comparativeexperiments
Laboratory experimentation in plant ecology has evolved very largely as an attempt to
pursue investigations which began with fieldwork. With the widespread development of
plant growth-room facilities an alternative approach is possible. This is to measure the
characteristics of plants under a variety of controlled conditions, and to use the results
to predict their field ecology (Grime & Hodgson 1969). One advantage of this approach
is that predictions can be tested against descriptions of the field ecology obtained by
independent field investigation.
In the long term, however, the most important advantage of the predictive approach
is that many growth-room investigations can be reproduced or extended wherever there
are adequate facilities: hence data collected on different species or genotypes and in
various laboratories can be compared directly. When comparable data are available for
a large number of species drawn from a wide range of habitats it may be possible to esti-
mate the limits of variation of a particularplant attribute, to place an individual measure-
ment in context and to attempt to judge its ecological significance.
The investigation describedin this paper is an attempt to examine the range and pattern
of variation in a local flora of one particularplant attribute-the maximum potential rate
of dry matter production. Although most data have been obtained from only one field
population per species the number of species is large and includes representatives from
all the major dry terrestrial habitats of the area. Uncertainty concerning the extent to
which each sample is representative of the species does not, therefore, invalidate the
exercise either as an estimate of the range of variation or as an attempt to recognize
differences between groups of species of contrasted ecology.

Screeningfor relative growth-rate
In a comparison of the physiology of growth among flowering plants of the Britishflora
it may be expected that genetically-determineddifferences will be exhibited, both with
respect to the circumstances optimal for dry matter production and in the maximum
rates of dry matter production actually achieved at the optimum. A number of
comparative studies has revealed such differences in what may be called 'maximum
potential relative growth-rate'* (Blackman & Wilson 1951a, b; Bradshaw et al.
1958; Blackman & Black 1959; Bradshaw, Chadwick et al. 1960a; Bradshaw, Lodge et
* Henceforwardnotated, after the style of Evans (1972, p. 633), as Rmax.

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394 RGR in a localflora
al. 1960b; Rorison 1960; Bradshaw et al. 1964; Jarvis & Jarvis 1964; Jefferies & Willis
1964; Clarkson 1965, 1967; Grime 1965a; Grime & Jeffrey 1965; Myerscough & White-
head 1966, 1967; Hutchinson 1967, 1970a, b; Pollard & Wareing 1968; Rorison 1968;
Higgs & James 1969; Foulds 1970). Despite the facts that (i) the number of species in-
cluded in each investigation was relatively small, and (ii) many were undoubtedly grown
under sub-optimal conditions, there were nevertheless strong indications that species
drawn from contrasted habitats often differedconsistently with respect to Rmx. Hence this
attribute of the flowering plant appears to be a promising candidate for extensive study.
In order to examine further any possible ecological significance in Rmax it would be
necessary, in the first place, to determine it over a wide range of species differing in
ecology. However, it is now well known that relative growth-rate in plants is simultan-
eously subject to genetic, ontogenetic and environmental control. Any attempt to make
meaningful comparisons between species with respect to what is, in effect, a function
close to their 'economy in working' (Blackman 1919) would face the considerable
difficulty of deciding upon what basis and over what time period to study their growth.
Such a study would certainly have to be conducted under controlled conditions in which,
as far as possible, no environmental factors were limiting, in order that the resulting
rates of growth might be internally rather than externally determined (Briggs 1928).
Once these comparisons had been made on a large scale another, more specific,
objective could be achieved; this would be an examination of the relationship between
Rmx and site productivity. The work of Njoku (1959), Jarvis & Jarvis (1964) and Grime
(1965b) supports the statement that high Rmax is associated with success in potentially
productive situations. Conversely, it is possible to deduce from work such as that of
Hackett (1967) and Rorison (1968) that low Rax is associated with a poor performance
in such circumstances. In unproductive situations (and in situations subject to environ-
mental stress) a low Rmx appears to be an advantage to certain species (Bradshaw et al.
1964; van den Driessche & Wareing 1966; Clarkson 1967; Hutchinson 1967; Parsons
1968a; Rorison 1968; Higgs & James 1969; Foulds 1970; Hunt 1970) whereas under
similar conditions a high Rmx may place certain species at a selective disadvantage
(Ashton 1958; Grime & Jeffrey 1965; Loach 1967; Parsons 1968b; Rorison 1968; Hunt
1970). Such hypotheses would be confirmed if it could be demonstrated that Rmaxwas,
per se, of positive or negative adaptive significance. In addition (or alternatively) the
importance of Rma might be seen to derive from its linkage to other variable characteris-
tics which themselves influence the fitness of the plant in the field.
This paper describes an initial attempt to make such comparsions and tests of hypo-
thesis. A variety of native species has been grown in monoculture in a standardized,
potentially-productive environment and measurement has been made of the rates of
growth attained in the phase of more or less exponential growth which follows shortly
after germination. Since growth in this phase is normally the most rapid in the life-
history of the plant (and since the external environmental factors were designed, as far as
possible, to be present in non-limiting quantities) these measurements may be regarded
as provisional estimates which, for the majority of the species, approximate to Rmx.
These comparisons minimize variation resulting from morphological and develop-
mental differences between species. These estimates of Rm, are examined in the light
of the species' known field distribution and an attempt is made to assess their eco-
logical significance. Only data relating to dry matter production are presented and dis-
cussed. Subsequent publications will deal with the influence of seed weight, leaf area
phenomena and the distribution of dry matter between root and shoot.

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Table 1. Thefrequency of occurrenceof each
The habitats tabulated are:
1, Lakes, ponds, canals and ditches; 2, Rivers and streams; 3, Unshaded mire; 4, Shaded mire; 5, Rock outcrops; 6, Limestone scree; 7, Cliffs; 8, Walls; 9
stone quarry heaps; 21, Quarry heaps on acidic strata; 22, Scrub; 23, Hedgerows; 24, Limestone woodlands; 25, Woodlands on acidic strata; 26, Broad-lea
In each case column a refers to the percentage occurrence of the species in forty or more 1-m2 samples and column b gives the ranking of the species accor
Habitat 1 2 3 4 5 6 7 8 9 10 11
Species ab ab ab ab ab ab ab ab a b a b a b
Acer pseudoplatanus (seedlings) - - 3 2 1 - - - - 2 -
Achillea millefolium - - - 6 65 3 1 2 2 5 21 21
- 5 60 - 1 - - 4 555 30 23 16 26
Agropyron repens - - - -
Agrostis canina 3 - 19 7 928 1
A. stolonifera 14 5 42 1 40 1 16 13 16 21 - 4 8 14 65 3 37 20 26 18
A. tenuis 1 - 10 25 544 30 8 - 10 14 6 20 646 42 18 70 2
Aira praecox - - - - 580 - - - - -
Alopecurus geniculatus 3 2 9 30 -2 - 9 35
A. pratensis - - - - 2725 1924
Anisantha sterilis - - - - 2 - 4 1 2 2 -

Anthoxanthumodoratum - - 9 30 4 8 41 22 15 4 3 2 45 16 35 12
Anthriscus sylvestris - - - - - 1 5 27 2 22 29 6
Arabis hirsuta - - - 10 34 14 21 4
Arenaria serpyllifolia - - - 26 10 3 2 1 1
Arrhenatherumelatius - - 1 5 44 21 15 97 1 12 10 19 5 6 46 12 35 5 54
Betula pubescens (seedlings) - - 1 4 6 65 - 4 2 1 2
Bidens tripartita - - 1 - - - - 1 - -
- - - 7 52 - 1 - -
Brachypodium pinnatum - - - -
B. sylvaticum - - 1 832 3
Briza media - - 6 65 8 32 - - 5 554
Calluna vulgaris -4 - 21 15 - 11 13 1 - - 2
- - - 18 18 41 6 10 14 1 - 554
Campanula rotundifolia - - -
Cardamine flexuosa - 6 14 6 47 27 3 1 -
C. pratensis 2 - 20 5 928 - - - 1 - 2 16 26
- - 3 - 2 387 - -
Carexflacca
C. panicea - - - 2 - - - - -
Catapodium rigidum - - - - 3 - I - - -
Centaurea nigra - 3 10 34 - 527 1 12 35 5 54
Cerastiumholosteoides - - 7 43 - 25 11 3 1 1 18 21 80 3 49 8
Chamaenerion angustifolium - - 1 4 8 41 - 14 8 16 8 -
- - - - - - 1 21 16 - -
Chenopodium album -
Cirsium vulgare - - 1 6 65 2 1 3 2 12 32
- - - - 1 - 1 - - - -
Cllnopodium vulgare - - - - - - -
Convolvulus arvensis 3 12 32
- - 4 - 3 - - 1 1 47 14 35 12
Cynosurus cristatus
Dactylis glomerata- - 4 - 36 3 14 21 23 2 19 5 11 35 80 3 51 6
Deschampsiacespitosa 1 2 8 3 10 25 1 3 - 4 2 2 14 29
D. flexuosa - - 11 7 544 32 6 261 2 - - 554
- - - 2 1 2 -
Digitalis purpurea - - - - -
Draba muralis - - -
7 12 4 15 10 3 1 - 1 8 14 -
Epilobium hirsutum - - - -
3 - 13 13 2
Eriophorum angustifolium -
Festuca giganteaI - 1 811 - 4 -
F. ovina - - 4 - 38 2 54 4 19 4 4 935
F. rubra - 2 10 5 - 47 1 65 2 21 3 28 2 551 55 9 53 5
2 - 3 1811 - - - - -
Filipendula ulmaria - - - - - -
Galeobdolon luteum- - 732 1
Galium aparine - - 1 1314 - - 3 2 16 24 7
G. palustre 1 - 16 9 1217 - - -
G. saxatile - - 8 33 - 5 80 - 1 1 -9 35
G. verum - - - 6 65 - - - - -
Geraniumrobertianum - - 1 544 7 52 65 2 10 14 8 14 1
Geum urbanum - - - - - - 2 1 - -
Glyceria fluitans 10 8 12 9 10 25 7 32 - - - - - - 5 54
G. maxima 6 14 - 8 33 -
Helianthemum chamaecistus - - - 6 65 22 15 -
Helictotrichon pratense - - 7 52 24 14 3
Heracleumsphondylium- - 3 637 10 34 3 4 1 551 35 21 12 32
Holcus lanatus 2 6 14 23 5 4 14 24 5 41 527 14 9 19 20 70 6 70 2
H. mollis 3 4 12 15 24 4 665 - 4 4 2 541 7 43
Hordeum murinum - - - -- 1 - - -
Juncuseffusus 5 22 2 31 2 10 25 1 -1232
J. squarrosus - - 2 - - -
Koeleria cristata - - - 16 21 19 18 4 -
- - - - - - - - 2
Lathyrus montanus
L. pratensis - 1 - 3 - - 1 27 25 7 43
Lemna minor 53 1 4 13 15 189 1 3 - -
Leontodon hispidus - - - 16 21 2712 1210 - - - 2
Loliumperenne - 2 1 - 10 34 - 2 5 27 16 24 82 2 51 6
Lotus corniculatus - - - 13 26 8 32 3 - 1 541 7 43
Luzula campestris - 3 - 665 8 32 2 1 - 17 33 35 12
- - - - - - - - - - -
Lycopersicon esculentum cv. Ailsa Craig - - -
Matricaria matricarioides - 1 1 - 49 8
- - - 18 18 - 4 1 6 46 2
Medicago lupulina - - - -
Melica nutans 3
Mentha aquatica 4 - 11 17 20 - - - 1
Milium effusum - - - -
Minuartia verna - - 1 -
- - - -
Myosotis sylvatica -
Nardus stricta - 647 1 1
- - -7 52 3 2 1 -
Origanum vulgare - - -
Oxalis acetosella 1 2 1 22 15 2
Phalaris arundinacea 3 8 11 7 43 928 - - - 2
Plantago lanceolata- - 2 - 36 3 5 41 11 13 4 551 50 12 26 18
P. major - - 3 1 665 - - 5 27 2017 10 37 19 24
Poaannua - 8 33 4 12 28 3 11 13 71 2 15 34 21 21
P. pratensis 2 7 43 - 28 9 11 24 719 22 3 6 46 2031 479
P. trivialis 1 15 7 29 3 451 3 5 41 622 40 1 28 12 871 604
Polygonum aviculare- - 1 1 3 - - -1 72 143
P. convolvulus- - - - - - - - 40 8 - -

P.persicaria-
- 2 - - - - 1 506 2
Potentilia erecta- - 833 4 - - - - - - 7 43
Poterium sanguisorba - - 752 11 24 1 - - - -
Prunlln. ,lunriel 7 43 1 9 39 3 1 5 51 27 25 1429

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. 2 17 20 . . Cliffs. 15 23 5 49 . . . 15. . . . . . - 19 5 11 35 80 3 51 6 35 16 . . . . . 50 3 1 . . . - 1 3 2 12 32 12 53 . 11 33 3 5 51 27 25 14 29 1643 2 8. . . 2 21 32 .. . 2 . . 7. . . - 4 . . .. 1 . 1 . . 26. . 4 . 2 .. 1 - 22 3 6 46 20 31 47 9 33 18 7 19 35 9 6 26 37 35 7 30 14 15 19 48 5 . 31. . . 3 15 18 S 2 2 5 21 21 16 43 1 7 52 . 15 18 - 3 2 45 16 35 12 58 4 14 9 3 . . 2. 219 533 36 2 16 24 7 . 11 39 4 . . . . . . 5 . . . . . . 8 40 35 12 35 6 8 51 . . 17 28 - 8 25 4 . . 7 64 1 . . . -3 2 12 32 . 30 8 . 3 . - 1 . 2 . . 11 11 5 56 . . 1 . . . I . . - 5 27 16 24 82 2 51 6 7 64 2 21 19 4 . 15 27 12 32 5 49 5 30 6 61 .. 19 21 . 49 3 14 15 . 17 28 - 1 18 21 80 3 49 8 31 21 2 15 29 . S . . . 2 . 20 17 37 9 10 22 5 65 - 11 13 71 2 15 34 21 21 2 3 4-7 14 15 4 3 6 71 1 48 2 9 43 . 14 3 18 11 - . . . . 2 . . . . 35 2 . . . . 2 10 38 20 13 .. -1235 . . 9 46 8 21 . 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 a b a b a b a b a b a b a b ab ab a b a b a b a b a b a b ab c 2 . 8. Wasteland and heath on limestone. - . . 10 19 . . . 2 . . 9. . 12 32 12 32 10 22 6 61 . 4 25 12 1 40 1 28 12 87 1 60 4 9 58 1 63 1 2 2 26 13 50 4 53 1 9 43 .167. 30. . . 1 . . 2 2 25 16 44 2 2 . 26 5 5 33 4 1 21 16 . 642 2 5 49 . . 2 . - I . . - 1 5 51 35 21 12 32 4 . . . .. 46 9 . 5 42 . . . 1 . Broad-leaved plantations. 12 32 . . 5 33 - 1 2 2 . 12 16 50 32 . . . - I . - 19 5 6 46 12 35 5 54 27 25 . . . . . . . 19 21 5 56 2 . 4 2 . . 3121 . . .194 on Mon. . . 1 . . 4 .4 1 .. 29 10 16 13 34 10 30 11 15 25 10 22 54 4 .. .. . - . . . Pastures on acidic strata. 455 33 9 15 25 8 28 28 20 . . 764 463 . 5 65 . 2 . 7 42 3 . 455 3 . 7 40 12 32 38 5 3 . 15 29 . 2 5 33 3 . 2 . 21 2 6 36 - 1 . 2 15 18 20 1 . . 2 . . . . 13. 5 33 . . 11 39 . 7 43 29 24 1111 . . . . . 9 43 ?. 25 22 - 2 . 2 . 513 --. . . 2 2 .. . 22 32 5 22 5 56 . . . . . . I . - 4 555 30 23 16 26 . 2. . . -. 52 3 18 16 38 11 . . . 11 38 2 . . .. . . 958 . . . 9 43 . . 5 42 - 5 27 2 22 29 6 2 . 1 27 25 7 43 5 72 . Limestone pastures. . . . 28. . . 8 - 1 . . 522 -. . 9 35 25 27 46 3 . 32 14 .. . 1 . 2 10 21 4 6 20 6 46 42 18 70 2 62 3 41 5 13 35 364 43 7 21 16 15 25 3 . . . 12. . - 1 1 . 642 . . . . 14. 3 10 38 - 1 . . 1 12 35 554 7 64 . 3 . . 5 54 35 16 97 1 3 10 20 15 27 9 37 2 . 45 9 . Enclosed pastures. - . - 1 . . -3 8 14 -. 39 8 22 9 17 25 39 5 32 13 28 11 55 2 . 19 10 . 3 2 7 6 17 13 30 9 1 1 . 19 21 3 . . . 25 16 14 15 15 23 30 14 20 13 2 . . 8 25 60 2 10 1 -i . . . 26 14 14 29 2 3 32 14 . . 49 8 .. . 2 . 2 . 7 4 . . 2 . 2 - 8 14 1 . 4 50 3 12 14 9 19 20 70 6 70 2 36 14 4 48 4 62 1 21 19 45 2 40 8 20 13 48 5 . No ranking is given for species with a percentage occurrence of less than five. . Meadows. . .65. 12 17 6 42 3 10 38 . 55 2 30 6 2 45 2 57 2 30 8 31 18 . . I . 9 35 . 3 . . 27. 3 . Verges. 2 . . . 2 5 36 . . . . . . 1 50 6 . 16 43 . . 10 21 1 1 47 14 35 12. . . . 15 23 45 5 18 16 - 1 6 46 2 . 12 35 . 2 7 64 . 9 22 50 3 13 2 1 2 .. 11. . 4 . .. 1 - -4 . . 2 1 1 8 21 4 6 48 5 49 . 29. . . . 6 61 12 16 . 2 . . . I 1 . . . .. . Coniferous plantations. . . 1 2 4 5 56 . 26 14 6 48 5 49 . . . 9 35 75 1 582 6 26 552 3 2 . 7 64 . . . . Coal Woodlands on acidic strata. I . 9 4 11 18 . 17 33 35 12 55 6 12 10 1 2 17 25 2 . . 7 52 4 5 56 5 49 5 36 5 65 - . . 5 54 -. . . 72 1 . 549 . . . . . Walls. . . 11 39 4 . . . . - . 1 1 .. 9 22 5 33 3 4 2 5 41 7 43 4 9 14 8 50 20 11 4 2 . . 33 18 . . 3 . 11 33 7 40 . .. 554 53 8 1 . 3 . Limestone scree. . . 4 .iuency of occurrenceof each of the species includedin the investigation in vegetationsamplesfrom thirty-onehabitats in th 6. - 1 . . 20 23 . . . 4 . 29 10 2 . . 2 . 12 37 . 1 5 41 7 43 44 10 2 3 . 5 49 . 4 . .. 2 16 8 . .. . . Wasteland and i b gives the ranking of the species according to percentage occurrence. 4 . 3 45 7 9 . 2 13 20 -. 3 . 3 27 25 19 24 . . . . . -4 . 2 . . . 5 . 9 43 . 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . 37 - . . 3 . . 14 32 . . . . 58 4 . . 18 26 5 8 5 42 - 28 2 5 51 55 9 53 5 64 2 3 21 19 12 17 51 3 41 27 17 21 5 36 66 1 . . -. 2 . 12 16 10 21 36 5 5 54 27 25 . 12 . . 3 . . 642 2 . . . . . . - 5 27 20 17 10 37 19 24 . ... 10 19 7 4 5 51 50 12 26 18 40 12 1 20 23 12 17 3212 17 20 12 32 5 36 38 11 . 3 . 37 1 34 1 - 1 2 . 1 . . Soil heaps. . 14 32 . 8 51 6 7 3 . . .. . 7 43 . 51 - This content downloaded from 69. 7 33 25 12 5 4 2 2 14 29 24 30 4 . . 10. . . . 2 16 26 4 . 2 2 - 8 14 65 3 37 20 26 18 4 . 2 . 3 7 42 . . 2 20 35 24 7 .. . . . . . 25 27 . . 4 . 40 8 .. . 2 2 3 . . Paths.. 642 . . Arable.

. . 1 II 1 1151 . . 3 . 9 22 5 33 3 4 8 20 7 11 29 12 12 16 34 11 19 7 Lower Bradway. . . 2 2 68 1 22 12 62 1 21 4 Lathkilldale. . 1 . . Wentworth Woodhouse. . . . I Ashover. Derbys.. .. . . W.. W. . - - - - - - -~~8 1036 6" 27 1 Via Gellia. . W. Pebley Quarry. 5 30 6 22 1 . . 8 38 . . 7 24 2 5 14 1 8 59 16 10 Deepcar. 1 4 Broomhill. . . 12 30 18 13 12 47 5 32 Coombsdale.- . . .. Sheffield -. Derbys. Derbys. 43 . . 6 51 2 28 13 630 Bramley. 3 2 32 12 2 Markland.. Derbys. . Riding . 3 . W. W. . . Coombsdale. . . . . Riding 32 . . Derbys. . -~~7 43 2410 . W. . ... 20 21 22 23 24 25 26 27 28 29 30 31 Source b a b a b a b a b a b a b a b a b ab a b a b 3 15 18 9 28 17 6 18 5 14 8 4 . . . . . Commercial source 33 6 I Roche Abbey. 2 Via Gellia. . 7 41 6 27 26 14 195 Two sources* 1 . . W. I Cromford Canal. Coal mine heaps. . . .. . 1 . . .1 .. Riding 9 4 11 18 . 18 . . 2 . Derbys. . .. . . 38 . . I 2 . . Riding 1 .. 5 . 6 27 18 24 7 27 Coombsdale. 13 42 . . Lead mine heaps. . 59 - 51 . . .- . W. . . . Via ellia. . . 31. 2 .. 14. 1 . . . . 14 36 . . 1 1 2 1 1 17 19 78 I 4 Orgrev. -6 71 . W. 6 51 .. . 2 . . . . . . . Derbys. . W. Derbys. . Cadeby. . Riding 28 . 2 ... . 51 6 7 3 . 5 32 Calver Slough. . Riding . . 12 30 6 27 2 1 Maltby. . .. Lindrick. . 15. .. . . . Lime- heath on limestone. 2 . 12 16 50 3 32 8 16 7 13 10 5 14 1425 39 4 9 55 1512 Cadeby Common. Derbys. Cinders. . Via Gellia. Riding --. . . Maltby. . . -. W. . . 11 51 . .. . Sheffield . 1 . . 21 2 6 36 . . 7 31 8 19 4 1 . I-. . Riding 5 42 . 19 10 . . Riding 10 19 . 1 . . 5 39 2 2 1 ..194 on Mon. . . Derbys. . 17. . 4 2 2 Coombsdale. 3 Clumber. Orgreave.. . Derbys. . . 65 . 1 2 . . Winnats Pass. . . . . . Notts. .. . . 1 2 Wharncliffe Chase. .. . .. . . . . . . . . . . . . . . .. 9 22 50 3 13 16 5 30 3 4 39 9 2 37 10 13 14 Via Gellia. 2 7 33 25 12 5 38 1 8 20 4 57 4 25 9 56 2 20 5 Orgreave. 30 3 31 2 27 3 4 5 27 6 71 54 1 Temple Normanton.65. I Walkley. 8 59 . . Lindrick. 2 . 18.167. . Riding . 50 3 . 11 . . ' 18 18 16 13 . .. . Northumberland 37 1 34 1 . . . . . . Derbys. 4 50 3 12 20 6 28 3 1 37 10 627 39 9 726 Via Gellia. Bradwell. . 8 25 60 2 10 24 8 19 6 22 2 25 15 2 14 32 630 Via Gellia. . W. Whitley Wood.. 51 . Lathkilldale.. . 4 . 5 33 . 2 -Smithy Wood. . .. W. Derbys. Riding 5 . Derbys. . -. 2 15 18 20 10 4 . 36 11 41 3 1 4 Markland Grips. .erbrhv. . 22 16 24 10 17 27 920 Waleswood. . . . 1 -1 Orgreave. . . . 1 2 2 . 3 . . . Millers Dale. 1 Litton. Riding . 35 . . . . . W. 2 Orgreave. W. 32 . . W. Derbys. . Derbys. . Sheffield 61 .. . . . . . . 4 .. . 14 25 2 20 19 11 17 Waleswood. Coombsdale. Roche Abbey. W. . W. Riding . W. . W.. Riding . . Derbys.. W. 2 3 Maltby Common. Sheffield . I I . Derbys. . . . Riding 4 2 . . . . 42 6 3 Millers Dale. Riding . 1 1 1 Black Carr Plantn. Riding 14 . W. 14 36 2 Millers Dale. . . 2 . . Derbys. . Soil heaps. I . ice of less than five.. Derbys. . 4 25 12 1 1 4 64 2 31 7 50 4 1610 Grangemill. Derbys. W. .. . . . Riding 61 . 43 . . . . Derbys. . . Wasteland and heath on acidic strata. -. W. . Riding 22 . . . 1 . . . Derbys. Manure and sewage waste: 20. . W. . Riding 65 . 1 Maltby. . 4 . . W. . . Cressbrookdale. . . 43 . .I. Riding . . Maltby. W. Derbys. . This content downloaded from 69. . . Cressbrookdale. . 14 . Riding 10 19 7 32 16 7 2 612 Anston Stones Wood. . 5 38 7 24 4 2 1 . Riding 20 .. Derbys.from thirty-onehabitats in the Sheffieldregion acidic strata. . 764 1 Orgreave. 1 1 Mansfield.. . 19. . . . 455 . . - . . . Riding 61 . .. 1 . Riding 28 . . Derbys. . 2 . . . 2 5 33 3 . . .. Riding . .. . 35 5 10 15 2 1 6 51 4 9 55 2 Ashington. I . Bricks and mortar. . Riding 35 . . Derbys. W. 1 Anston Stones Wood. . . .. . Riding . - . . 6 51 4 408 1 Maltby. . W. . Derbys. 11 . . Riding 43 . Litton. Riding . W. 3 . . . 1 . 5 42 . 8 38 2 1824 3 Maltby. . . Riding . . 43 . .. .. W. . . . . Derbys. . 12 20 5 30 2 . W. . . Mansfield. . . 2 1 Via Gellia. 1 1 Cressbrookdale. . Riding . 579 . Derbys. Riding . 1 6 51 4 4 3 Taddington Wood. Derbys. Lathkilldale. 14 25 2 16 28 2 Monksdale. .- 9 . 4 . . . 4 . 4 8 19 .. . Riding 5 33 . Sheffield 12 30 . . .. W. . . Notts. 1 . . . 21 9 5 33 36 3 12 12 1 . 8 25 . W. W. . W.. . Riding 10 21 . . . . 26 5 5 33 4 13 11 17 6 26 4 12 30 6 27 15 31 629 Walkley. Derbys. . 12 16 . . . 3 . W. . 1 1 Calver Slough. . Riding 14 3 18 11 . . 42 8 10 18 7 64 10 19 Worksop. 1 . Notts. . W. Derbys. . 1 . . 6 27 955 1 LathkiUdale. 1 4 . 29 12 627 2 4 Dore. . . 1 4 4 923 Wharncliffe Chase. 15 18 . . 1 . 15 22 67 2 . . 1 . 14 36 ... Notts. 4 Orgreave. 1 . 5 8 . 2 10 21 4 1 1 4 46 6 37 5 6 71 11 17 Wortley. Derbys. .-. Orgreave..- - . 2 Orgreave. Riding 65 . 1 Conisborough. . . . . 2 Broomhead Moor. 1 . . Riding 26 5 8 5 42 . . 12 47 .. . Riding . . Derbys. 12 30 . 4 . Derbys. W. . . .. .. Riding . Via Gellia. Derbys. . Riding . 532 Wharncliffe.. . . . W. . . . . . W. 1 3 6 27 2 18 9 Balderstone. 16. 5 30 14 9 11 9 21 17 31 6 41 7 38 2 Broomhill. Orgreave. . Riding 7 6 17 13 30 9 1 35 1 25 3 24 6 7 14 1 5 79 26 3 Lathkilldale. 3 . Riding 3 45 7 9 28 3 .. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . . Riding I . . . . . 12 16 10 21 36 3 2 5 28 4 . W. . . 1151 1 Monksdale. W. W.

1 19 17 - . . . . . . mnjo. 1 6 65 2 1 3 2 12 32 . . . - Filipendula ulmaria . 1 Centaurea nigra . . 3 12 22 7 39 2 Salix cinerea ssp. - . . . 13 13 2 . . . . . - Rorippa nasturtium-aquaticum . 6 47 4 . - G. 1 8 14 - Epilobium hirsutum . 3 . . . . . 47 1 65 2 21 3 28 2 5 51 55 9 53 5 2 . . 2 Trifolium medium T. 7 43 . 2 . . 72 1 7 43 Polygonum aviculare . panicea 2 . . . 3 . 4 2 2 14 29 D. 13 26 8 32 3 . 1 21 16 . . 11 17 20 . . I . . verum . - Eriophorum angustifolium . . . 3 2 16 24 7 G. . 7. . 1 3 . 36 3 5 41 11 13 4 5 51 50 12 26 18 P. . 12 28 . 16 26 S. . . - Chenopodium album - Cirsium vulgare . . 2 . 8 33 4 12 28 . flexuosa . . . . - Stellaria media . Ailsa Craig . . 2 Plantago lanceolata. . . . 1 . 2 1 . . . - Cllnopodium vulgare . - 7 12 4 15 10 3 1 . . 6 47 1 1 . - Veronica arvensis . . 14 8 16 8 . 16 21 19 18 4 . . . . . . . . 9 35 G. . - Silene dioica 1 18 11 2 4 1 . - Thymus drucei . - Matricaria matricarioides . 1 . squalidus . . . 2 Lathyrus montanus L. . 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . . - Festuca gigantea . . 17 33 35 12 . . 7 43 Poterium sanguisorba . . . 1 5 41 7 43 Luzula campestris . 1 3 . . 1 . . 8 33 4 . - . 554 Glyceria fluitans G. rubra . repens . 2 8 33 . . 1 Galium aparine . 4 . . . 1 1 47 14 35 12 Cynosurus cristatus Dactylis glomerata. 3 1811 . 1 5 44 7 52 65 2 10 14 8 14 1 Geum urbanum . atrocinerea (seedlings) . 9 39 11 24 5 27 - Sedum acre . 5 80 . . 8 33 . . . . - Minuartia verna . . 2 2 . . . . . 38 2 54 4 194 4 . 12 15 35 2 8 32 3 13 10 15 26 2 2 Vaccinium myrtillus. . 554 Sieglingia decumbens . . 2 10 5 . 7 52 . convolvulus 408 - P. . 4 . - Oxalis acetosella. 1 . 7 52 11 24 1 . - P. . 394) This content downloaded from 69. .Carex flacca . . . . . - Convolvulus arvensis 3 1 12 32 . . . 7 19 5 27 1 - S. . --- Myosotis sylvatica - Nardus stricta . .194 on Mon. . . . 3 20 17 52 10 44 11 8 9 31 2 6 4 12 17 . . 7 52 3 2 1 - Origanum vulgare . . persicaria . . 1 1 2 Zerna erecta. - Juncuseffusus 5 22 2 31 2 10 25 . . . 3 1 65 3 10 37 - Succisa pratensis 1 1 . . . . - . . . 1 . . . . 1 . 7 43 1 9 39 3 1 . 8 14 . 3 . 12 28 5 41 2 3 Senecio jacobaea. - C. 2 5 27 16 24 82 2 51 6 Lotus corniculatus . 2 7 43 . . . . . 665 8 32 2 1 . 1 27 25 7 43 Lemna minor 53 1 4 13 15 18 -1 3 - Leontodon hispidus . . - 3 . 1- Milium effusum . . .167. . 8 33 . . 1 4 8 41 . . . maxima 6 14 . 25 11 3 1 1 18 21 80 3 49 8 Chamaenerion angustifolium . acetosella . . . 1 1314 . . obtusifolius 1 2 4 544 . 1 50 6 . . . - Galeobdolon luteum. -2 1 . 1 1 . 1 6 46 2 2 Viola riviniana. . 1 1 2 43 5 . - . . . 5 27 20 17 10 37 19 24 Poaannua . . 2 . 12 32 J. .32 . 1 25 15 72 15 77 1 . . . - Lycopersicon esculentum cv. 28 9 11 24 7 19 22 3 6 46 20 31 47 9 P. . 3 . 16 9 12 17 . . - Rumex acetosa I 13 13 7 52 5 41 1 1 4 65 7 47 9 R. 2 Taraxacumofficinale 4 2 31 7 11 24 18 6 22 3 11 35 50 12 28 16 Teucrium scorodonia . - Catapodium rigidum . . 6 65 . . mollis 3 4 12 15 24 4 665 . . . . . - Helianthemum chamaecistus . . . . . 1 2 2 2 14 29 R. . 2 1 1 . 5 54 . 6 65 5 41 . 1 . ovina . . . 2 Vaccinium vitis-idaea . . . . . . 5 80 32 10 527 1 . . vulgaris . 7 52 24 14 3 - Heracleumsphondylium. - Sanicula europaea . pratensis. 2 Loliumperenne . 3 11 13 71 2 15 34 21 21 P. 36 3 14 21 23 2 19 5 11 35 80 3 51 6 Deschampsiacespitosa 1 2 8 3 10 25 1 3 . . 11 7 544 32 6 26 1 2 . 1 1 . . 13 26 27 13 2 . . . . 2 . . . 16 21 27 12 12 10 . .65. 21 15 35 9 10 14 7 18 3 . . 3 6 37 10 34 3 4 1 5 51 35 21 12 32 Holcus lanatus 2 6 14 23 5 4 14 24 5 41 5 27 14 9 19 20 70 6 70 2 H. 7 52 . - Geraniumrobertianum . . . . 2 21 - Digitalis purpurea 2 Draba muralis . . . 18 18 . pratensis . - Prunella vulgaris . . trivialis 1 15 7 29 3 45 1 3 5 41 6 22 40 1 28 12 87 1 60 4 . . . . 6 47 1 3 8 32 1 4 5 51 - Tussilagofarfara . 2 Potentilla erecta . . . saxatile . - Sarothamnus scoparius Scabiosa columbaria . . . . . - Ulex europaeus . . . . In addition to the local sour * Festuca ovina was collec (Facing p. . . . . 1 . 3 1 6 65 . 3 10 34 527 1 12 35 5 54 Cerastiumholosteoides . - Helictotrichon pratense . . squarrosus . 6 65 22 15 . 49 8 . . 5 51 27 25 14 29 Ranunculusrepens 6 14 6 14 244 244 1 . . - Urtica dioica . - Melica nutans 3 Mentha aquatica 4 . - 10 8 12 9 10 25 7 32 . . . 8 41 . . . palustre 1 . . . . . - Koeleria cristata . . . 1 2 1 22 15 2 Phalaris arundinacea 3 8 11 7 43 9 28 . 12 28 . 4 1 6 46 2 Medicago lupulina . . 4 4 2 5 41 7 43 Hordeum murinum . . - . . 1 1811 4 - F. 4 . . . 2 The data presented in this paper were obtained using seed collected from the sources listed on the right-hand side of the table. 10 34 . . . 9 35 F. . . . .

. 2 . . . . 1 10 21 . 3 . . 15 29 2 19 21 8 40 7 42 5 36 46 9 . . . . -. 18 26 5 8 5 42 - 28 2 5 51 55 9 53 5 64 2 3 21 19 12 17 51 3 41 27 17 21 5 36 66 1 . . . I . .. 4 50 3 12 14 9 19 20 70 6 70 2 36 14 4 484 62 1 21 19 45 2 40 8 20 13 48 5 . . 9 37 12 32 8 28 11 38 . . 8 51 . J 0. 2 . - 8 14 .. 15 23 45 5 18 16 - 1 6 46 2 . . - . . . . 3 10 38 - 1 . 1 . 12 ... -19 21 5 56 2 . . . . 6 36 10 21 7 1 . 10 19 7 4 551 50 12 26 18 4012 1 20 23 12 17 32 12 17 20 12 32 5 36 38 11 . . 2 . . . 2 . I 3I - 958 1 . 37 1 34 1 - 1 2 .. 1 5 41 7 43 44 10 2 3 . . 2 5 36 . . 2 . - 5 27 20 17 10 37 19 24 . . 2 . 20 23 . 5 56 17 21 33 7 . 1S 27 5 56 . 18 41 . 584 . . W. 7 52 4 . . . . 32 14 . 1 6 . . . W. 4 . . . 48 5 .. 51 3 . I 1 . 2 . . 11 33 3 . - 1 5 51 35 21 12 32 4 . 10 19 . . . 11 33 2 . 4 .. . . . . . .. . Riding and Wye and Crundale N. 7 43 29 24 11 11 . 6 61 . . . . . 64 1 11 36 5 49 3 943 . 17 33 35 12 55 6 12 10 1 2 17 25 2 . . - .. . 1 . 2 15 18 20 1 . .. 7 52 22 9 . 25 22 - 2 . 1 1 4 65 7 47 9 31 21 3 8 50 . 39 8 22 9 17 25 39 5 32 13 28 11 55 2 . . . 28 13 2 11 33 35 7 37 9 10 22 38 11 . . 2 764 . - 1 27 25 7 43 572 . . . 2 . .. 1 - . In addition to the local sources. 3 13 ~. . 2 2 3 . . . R.. . . . . . . 5 54 35 16 97 1 3 10 20 15 27 9 37 2 . . . 12 37 . .. 3 . Riding (acidic substratum) and Via Gellia. collections of Brachypodium pinnatum from Crich. . 642 . . 1 .. 11 11 56 26 26 . . . -3 2 12 32 . - 1 . . 29 10 2 . 219 5 33 36 2 16 24 7 . . . . 2 13 20 - . 446 . . - 2 2 2 1 25 15 72 15 77 1 36 14 3 24 18 . 28 13 . . - 1 1 . . . 1921 . 52 3 18 16 38 11 . 2 21 32 . - . . . 4 25 12 1 40 1 28 12 87 1 60 4 958 1 631 2 2 26 13 50 4 53 1 9 43 .. 522 . 2 2 - . . . . 2 12 53 415 . -. 2 40 12 1 . 2 . 10 55 . 661 12 16 .. . . . . . . 4 . - . . . . . . 459 -. . . .. . . -.. 6 36 - 3 12 22 7 39 2 . . . 19 . . . . 16 26 33 18 . - 1 . . 3 . . . 2 4 . . - 13 10 15 26 2 2 2 . . 2 16 8 . . . . 9 46 8 21 . . . - 19 5 11 35 80 3 51 6 35 16 . 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . . . 33 la I . 2 4 . . 11 36 32 13 10 32 .. 35 2 . - 7 18 3 . 57_2 . . . 51 27 25 14 29 16 43 . 4 2 ... . - 1 . 2 . 937 - . 1 - . . .65. . . 2 10 38 20 13 . . 15 23 30 14 20 13 2 . . . . 5 42 . 25 16 14 15 . . - 5 27 16 24 82 2 51 6 764 2 21 19 4 . - 1 50 6 . 28 13 . . 26 14 14 29 2 3 32 14 . . . 1 22 3 6 46 20 31 47 9 33 18 7 19 359 6 26 9 37 35 7 30 14 15 19 48 5 . 4 . - of the table... . 9 35 75 1 58 2 .. 12 32 12 32 10 22 6 61 . 2 -8 51 - 3 20 17 52 10 44 11 .. . . . and of Zerna erecta from Cric * Festuca ovina was collected from two sources: Fox House. . . . 8 40 35 12 35 6 8 51 . . . . 26 5 5 33 4 1 21 16 .. . . 8 25 4 . . N. . . . 5 65 .. 943 1 12 35 5 54 7 64 . . . 2 9 37 7 42 5 36 8 51 . 2 20 35 . 23 18 18 19 22 17 8 28 12 35 . . 15 23 5 49 . 2 15 23 42 6 40 4 3 .. . . . . . 1 . 2 . 25 27 . . 25 27 -. (calcareous substratum). 8 25 60 2 10 1 . This content downloaded from 69.. -3 . 1 ~ ~ ~ .. .. 1 - . 45 5 33 9 15 25 8 28 28 20 . 3 . . . 17 28 - 1 18 21 80 3 49 8 31 21 2 15 29 . 5 - . . . . - 1 3 2 12 32 12 53 . . 1 . - . 2 . .. 1 4 5 51 . . 9 4 11 18 - . .. . . 764 463 . . . . 12 32 . -. . 9 22 5 33 3 4 2 5 41 7 43 4 9 14 8 50 20 11 4 2 . . . 1 . . 7 33 25 12 5 4 2 2 14 29 24 30 4 . 26 14 6 48 5 49 . 40 8 . . 5 33 5 54 16 43 . 4 --43 . 12 35 . 23 15 45 5 3 8 51 . . . 10 21 1 1 47 14 35 12. . 3 2 7 6 17 13 30 9 1 1 . . 2 . . 2 2 25 16 44 2 2 . . . 11 11 5 56 . - 11 13 71 2 15 34 21 21 2 3 47 14 15 4 3 6 1 48 2 9 43 . . - 1 . 6 48 2 . . . . . 5 33 . . 407 8 21 . . . . . . . 17 28 - .. 8 51 . . .. 2 22 3 11 35 50 12 28 16 22 32 . . 9 35 25 27 46 3 . . 2 5 27 1 . I*.. - 1 19 17 . . - 4 . . Q L 1 65 3 10 37 . . .. . - 3 . . 556 5 49 5 36 5 65 - -4 . 4 . 29 10 . 2 . - . 5 554 2 . - 8 14 . . 3 . . . .194 on Mon. . . . 5 56 . 58 3 1 6 46 2 2 7 64 . 2 . 2 . 16 43 . . . . 3 . 12 17 6 42 3 10 38 . 6 42 . 10 19 . 8 51 6 7 3 49 8 . . . .167. 2 17 20 .. . . 46 9 . . . . . 11 39 4 . . 31 21 . 6 42 2 . . 8 5 7 33 - 1111 . . 2 5 33 3 ~. . . 7 42 3 . Derbys. . 7 4 . 58 . . . 7 43 . . 15 29 . 20 17 37 9 10 22 5 65 . . . .. . . . . . . . . . 12 16 503 32 . - . . . 11 39 . 626 55 2 3 2 . . . 7 64 1 . 4 . . 5 65 . 1 2 2 2 14 29 . . 6 42 2 5 49 . 72 1 . . 28 13 . 2 .. -. . . 21 19 38 3 6 42 27 12 42 6 . 6 48 15 25 43 3 3 15 14 651 17 1 1 . 6 36 . 2 8 14 1 .. .. ..

I . . 2 . W. . . . . Riding 9 . . . Derbys.. . . . 2 . . . . . Derbys. . 1 . . Notts. . . 30 3 31 2 27 3 4 5 27 6 71 54 1 Temple Normanton. . Via Gellia. .-. ... . I Black Carr Plantn. . . W. 651 4 408 1 Maltby. 12 16 50 3 32 8 16 7 13 10 5 14 14 25 39 4 9 55 15 12 Cadeby Common. . 6 27 955 1 Lathkilldale. .- . Sheffield 51 . W. . Riding . . Derbys. I1 1 I-. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions .. Riding 3 13 16 1 2 Anston Stones Wood. .... .. Derbys. Riding . 14 25 2 20 19 11 17 Waleswood. . . . W. 17 19 10 18 13 42 4 Deepcar. Northumberland 37 1 34 1 . Riding 7 6 17 13 30 9 1 35 1 25 3 24 6 714 1 5 79 26 3 Lathkilldale. . . . . . . . 12 47 . . Riding . 4 3 Coombsdale. .. 12 20 5 30 2 . . .. Coombsdale. 1 . . . Derbys. Markland. . 2 Orgreave. 15 22 67 2 . . . . . Derbys. W. Derbys. 764 1 Orgreave. Riding 51 . . Winnats Pass. W. Sheffield .. . . . Riding .. . . Pleasley Vale.. . -2 . 38 . Notts. Derbys. W. . Derbys. Derbys. Derbys. Commercial source 33 6 . W. . .. 4 Orgreave. . Riding 28 . -. . . Derbys. . W. . 3 6 27 2 2 Broomhill. 5 33 . Lindrick. W. Riding . 1 Orgreave. . Roche Abbey. . . . 4 6 27 1 3 Central Sheffield . . 51 . . . Derbys. . . 7 31 8 19 4 1 . 35 5 10 15 2 1 6 51 4 9 55 2 Ashington. . . . .. ... . .. 26 5 5 33 4 13 11 17 6 26 4 12 30 6 27 15 31 6 29 Walkley. . 5 . 1 . . . . 4 25 12 1 1 4 .. .. . . . W. W. . 10 36 6 27 1 1 Via Gellia. . Riding 28 . W. . 4 8 19 . -. . 3 .65. . W. . Cressbrookdale.. . . Riding 5 . . 1 . . Derbys... Riding 35 . . I1 . Riding ... . . 2 . 4 . Derbys. Sheffield 5 33 . Derbys. . . 6 36 . . . Pebley Quarry. . 2 .. Sheffield 12 30 . . . W. 1 2 2 . . . . . 5 42 . .. 11 . . W. . . Lathkilldale. 43 . . 2 . . 1 . . . . . Mansfield. . .. . . 1 2 . .. .. . 3 9 17 1 8 38 10 18 1 3 Orgreave. Derbys. 43 1 . . . 7 24 2 5 14 1 8 59 16 10 Deepcar. 7 41 6 27 26 14 19 5 Two sources* 1 . ..167. 764 1 Lathkilldale. W. . W. . 11 .. Riding 51 . I Roche Abbey. .. . . . . Wentworth Woodhouse.. 1 . . . . . were used. Derbys. Derbys. Lindrick. . W. Derbys. 14 36 . 1 4 4 9 23 Wharncliffe Chase. . .. Riding 10 19 . . -18 18 16 13 .. . . 46 6 31 7 2 5 32 Norton. 2 2 1 5 14 6 .. . Riding . 5 30 6 22 1 . W.. 42 6 3 Millers Dale. W. . . 6 51 6 27 4 Wortley.39 .. Riding 9 4 11 18 . Notts. Riding 65 . Sheffield -. . Via Gellia.. Bradwell. . . and of Zerna erecta from Crich. . Derbys. . . 12 47 .. . Cressbrookdale. Sheffield 15 14 65 1 17 13 10 15 10 14 9 10 29 12 8 22 14 32 9 20 Coombsdale.194 on Mon. Maltby. 3 . Riding 51 . . . . Derbys. . I Ashover.. . W. . . .. W. Juniper Top. 14 36 . 8 59 ... 11 51 . W. Riding . 4 . . . . 79 .. Riding . Derbys. 36 11 41 3 1 4 Markland Grips. . . . Bradwell. . . . 1 1 1151 .. 2 . W. W. 2 3 Maltby Common. . b1f . . . 21 9 5 33 36 3 12 12 1 . 65 . . . N. 1 . . . 3 14 13 Wigtwizzle... . This content downloaded from 69. 18 18 2 26 14 11 16 Loxley. Derbys. . .- 9 . W. Riding 10 19 . . W.. . 1 2 1 1 17 19 78 1 . . Riding 61 .. . . . Lathkilldale. 12 30 18 13 12 47 5 32 Coombsdale. .. Surrey. . . -1 . . Sheffield .. . Riding 14 . . Derbys. 2 . 11 . 64 2 31 7 50 4 16 10 Grangemill. . .. 3 . . Derbys. Orgreave.. Derbys. 2 . . W. Riding 22 . . . W. . Winnats Pass. W. 5 32 Wharncliffe.. Bawtry. . . Riding . . Riding . 65 . Riding . 1 . ... W. 7 64 . 2 5 33 3 . . W. W. 16 28 . . . ... . 2 . . .. . Riding 26 5 8 5 42 . 22 16 24 10 17 27 9 20 Waleswood. 10 19 . Derbys. . . . . 2 20 19 . Derbys. . 1 522 14 14 2 9 20 Norton. 10 15 1 4 . . . . .. 2 1 Via Gellia... 1 . I . 2 Via Gellia. 1 Anston Stones Wood. Uerbys. 3 2 Norton. . . Derbys. 7 43 24 10 . . Fircester. . .. .. . . (calcareous substratum). .. Maltby. . Riding 6 36 1 4 2 1 6 27 2 19 7 Yarncliffe.. . 2 7 33 25 12 5 38 1 8 20 4 57 4 25 9 56 2 20 5 Orgreave. . 14 36 2 Millers Dale. .- - -. Wilts. j . ^5 IQ&2 161l 12 12 9 17 4 . -. 12 16 . 8 5 7 33 . Riding - . . W. 9 12 3 2 1 3 4 21 17 1 Ranby. .. 4 . . . .. . - Thorpe Common. . 6 36 10 21 7 32 2 2 . . . . 61 . 5 39 2 2 1 . W. 1 .. 1 . W. . . W. . . .. -6 71 . . Sheffield . 65 . W. .. 47 5 24 10 14 32 3 Broomhill. Whitley Wood. Riding 1 . . 3 Froggatt Edge. . Derbys. . 2 . . .. . 14 . Millers Dale... . 2 Broomhead Moor. . 1 Litton. Derbys. . 4 . Orgreave. I Cromford Canal.. Derbys. 51 . 51 6 7 3 . . --. 35 . N. 1 . .. . . . . I Deepcar. 4 50 3 12 20 6 28 3 1 37 10 6 27 39 9 7 26 Via Gellia. 3 . 5 8 .. 8 25 60 2 10 24 8 19 6 22 2 25 15 2 14 32 6 30 Via Gellia. 1 .. 12 30 . 1 6 51 4 4 3 Taddington Wood. . a . 838 2 18 24 3 Maltby. Derbys.. Litton. 9 22 5 33 3 4 8 20 7 11 29 12 12 16 34 11 19 7 Lower Bradway. . . . 12 30 6 27 2 1 Maltby. I Walkley. Derbys. . 3 .. . . . Riding. Riding 61 . Riding 20 . Derbys. . .. . . 2 . 43 . . . 11 31 1 Monksdale.. Derbys. . 2 -Smithy Wood. .. Derbys. . Riding .. 1 . . . 1 1 Calver Slough. . 1 10 21 . Riding . .. . W. W. 732 167 2 612 Anston Stones Wood. . 2 . . . R. 1 Broomhill. . W. Derbys. . .. Riding . Riding .. Derbys. . 2 2 68 1 22 12 62 1 21 4 Lathkilldale. . 1 . W. . Riding . 1 . . 2 15 18 20 10 4 . . . .. Riding 43 . . . Riding 5 8 3 . 2 . . Maltby. and Monksdale. . . . 4 Orgreav. Cressbrookdale. . Sheffield 3 4 859 1 Peak Forest. . . . Notts. . 2 Orgreave. Riding . . . . . .. . .. 8 59 . . . W. 8 25 . . . . W. . Riding 10 21 . . 1 . . . Mansfield. .

* Nomenclaturefollows that of Clapham. The list (Table 1) includes species characteristicof the more ancient landscape and also plants associated with habitats which are either disturbed or of recent origin. could not be represented properly because seeds or seedlings were not readily available and because the experimental conditions appeared to be unfavourable climatically for certain of these species. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . P. Data from an unpublished vegetation survey (J. Even on a smaller scale than at present such a task would be totally impracticable. estimations should be made under the growth conditions optimal for that species. each species was represented by a collection of seed or seedlings from one local field site. are now excluded from further consideration. to ensure measurement of the true Rmax of which each species is capable. additional comparisons were possible. both aquatic. In the case of tomato (an important naturalized species on sewage waste) and in several coniferous tree species commercial seed sources were used. in which separate analyses were carried out using seed from different sources. The majority of the species selected were herbaceous plants of widespread occurrence in Britain. The essential features of the survey area have been described elsewhere (Lloyd. In Table 1 this ranking is indicated in columns headed b.167. GRIME ANDRODERICK HUNT 395 MATERIALS AND METHODS Species The species were chosen primarily in order to allow comparisons in rates of seedling growth to be made between groups of flowering plants characteristicof local examples of the major inland habitats of the British Isles.194 on Mon. The extent to which this sample of species represents the complete list of plants to be found in each habitat is shown in Table 2. However. Grime & Rorison 1971. because the number of species involved was large (132). The environment selected for the measurement of Rmx in this large number of species therefore had to some extent to be a compromise between what was theoretically desirable and what was experimentally practicable.65. obtained from such a series of experiments. The data from this survey also allow the species recorded in each habitat to be ranked in order of percentage occurrence in the samples. became large enough to permit separate consideration. B This content downloaded from 69. notably legumes and woody species. columns headed a). that of vernal woodland species. J.*Festuca ovina and Zerna erecta. Yorkshire (Table 1.Tutin & Warburg(1962). In order to facilitate comparisons between groups defined by these secon- dary criteria. P. Hodgson. Grime & J. in pre- paration) have made it possible to estimate the frequency of occurrence of each of these species in the major habitats in an area of approximately 900 square miles (2400 km2) around Sheffield. A natural consequence of selection on ecological grounds was the inclusion of species differing widely in other plant characteristics such as life-history. With the exception of Brachypodiumpinnatum. These two. An important ecological group. R. Satisfactory representation was obtained in all but the first two habitats. Experimentalconditions General considerations Ideally. G. additional species were selected in order that certain under-represented groups. quantity of seed reserve and morphology of the root or shoot. Grime & Lloyd 1973). In order to find these optimal conditions it would be necessary to subject each species to a wide range of factorially-combined levels of the necessary growth-factors (sensu Lockhart 1965) and then to adopt as Rmax the highest value of relative growth-rate.

Lakes.194 on Mon. canals and ditches 32 4 2. Comparisons of Rmaxmade among many species in a single set of environmental conditions therefore may result in an under-estimation of Rmaxin species adapted to survive in environments far removed from that used in the investigation. Parsons (1968b).396 RGR in a local flora The level of supply of water and mineral nutrients was considerably higher than that commonly experienced by many of the species in the field. Wastelandand heath on limestone 57 10 31. Broad-leavedplantations 54 6 27. despite inaccuracies. Arable 37 8 10. Meadows 60 9 11. (1964) recorded reduced yields in greenhouse-grown Festuca ovina and Nardus stricta at nitrogen levels above 81 parts/106. Woodlandson acidic strata 62 5 26. Rorison (1968) and others suggests that normally unproductive species are often able to grow faster in monoculture in a productive environment than in conditions more nearly approaching those of their natural habitat. However. Quarryheaps on acidic strata 65 10 22. The representationof species examined among those of common occurrence in habitats 1-31 (Table 1. the work of Hackett (1967). Cliffs 53 5 8. Walls 67 9 9. Coal mine heaps 52 7 16. Rivers and streams 44 4 3. Enclosedpastures 64 10 12. such comparisons would differentiate between species of low and high This content downloaded from 69. Wastelandand heath on acidic strata 81 9 It seems likely that some species achieve their true Rmax only at rates of supply of growth-factors which would be harmful to other less demanding species. Hedgerows 56 7 24.65. Limestonewoodlands 47 8 25. markedly sub-optimal growth is restricted to species in which the true Rmaxis low. Scrub 61 5 23. Bricksand mortar 68 10 19.167. a concentration which is suboptimal for some arable species in the field. Limestonescree 60 9 7. there is evidence in the litera- ture and from this investigation to suggest that. Lead mine heaps 73 7 17. Manureand sewage waste 58 9 20. facing p. Paths 81 9 30. Cinders 60 8 18. Coniferousplantations 58 6 28. Soil heaps 64 10 15. However. ponds. although Bradshaw et al. Hence. under productive conditions. Pastureson acidic strata 78 8 14. Limestonepastures 66 9 13. Unshadedmire 58 8 4. Shadedmire 46 6 5. Rock outcrops 60 8 6. Table 2. 394) Percentagerepresentation Number representedfrom of specieswith an the ten most frequently occurrence of 5% or more occurring species Habitat in the habitat 1. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . Limestonequarryheaps 55 8 21. Verges 69 9 29.

99 0'62 0. Here. Species 15 20 25 30 (P< 0-05) Agrostistenuis 1-31 1-58 1-52 1-36 0'29 Arrhenatherum elatius 0. The controlled-environment rooms described by Rorison (1964) were used for this purpose. or nearly so. Illumination of the growing plants in these rooms was provided by thirty 5-ft (152-cm) 80 watt 'warm white' aged fluorescent tubes which supplied a maximum of 38 0 W m-2 visible radiation (0'0545 cal cm-2 min -) at plant height. It seems that in each case a day temperature of 20? C is at.99 1.22 0.65. 20 25 or 30? C. and using methods closely similar to those of the main series of experiments except that for each species there was a treatment in which temperaturewas held continuously at 15. 18 h. Arrhenatherumelatius. five grasses Agrostis tenuis.11 Festucaovina 1. Table 3. it is nevertheless the case that the daily light energy input to plants in the growth-room was less than that which they would receive in the majority of open field situations at temperate latitudes. P. Although arrangements were made to ensure that each plant received this level of radiation with no shading by adjacent plants. This is above the normal mean summer air temperature of most of the field situations near Sheffield but is not outside the experience of any of the species used.23 Zernaerecta 0'72 1 03 0'99 0-82 0 11 Temperatureregime A uniform day temperature of 20? C was chosen.167.06 1. J.194 on Mon.42 1-16 0. This figure is comparable to mean daily outdoor values for only two or three winter months in the year and is only in the region of 24% of the mean daily value for similar latitudes to Sheffield in June (de Vries 1955). Festuca ovina and Zerna erecta were grown under conditions. was chosen to encourage a high productivity (Hughes & Evans 1963). Light regime The scale of the operation required a cumulative series of standardized experiments. These results are summarized in Table 3. The daily total visible radiation supplied was 2-46 MJ m-2 (equivalent to 58-95 cal cm-2 day-').31 1.S. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . Des- champsiaflexuosa. The relationshipbetween temperatureand mean relativegrowth-rate (week-1) over the period 2-5 weeks after germination in five grass species (data of Mahmoud 1973) Temperature (? C) L.37 Deschampsiaflexuosa 0'96 1. R (mean relative growth-rate) was calculated from Fisher's (1920) formula: (loge 5W-loge 2W)/T where W and 2 W This content downloaded from 69. Since the present experiments were concluded the work of Mahmoud (1973) has confirmed the suitability of the chosen temperature regime for near-optimal growth in several native species.12 0-85 0. At night the temperaturewas lowered to 15? C to decrease dry weight losses due to respiration and to give a crude approximation to a 'normal' diurnal temperature variation which may be of physiological importance to many species. GRIME HUNT ANDRODERICK 397 Rmaxproviding that the environment selected allows Rmaxto be attained. or near to.05 1. Experiments carried out by Went (1957) con- firm that high yields may be obtained in a variety of different species grown at this tem- perature. the optimum temperature for the realization of Rmax given the background of the other conditions used.D. A long day-length. in species of high potential growth-rate.

ovina and Scabiosa columbaria were grown in the controlled-environment room in sand/solution culture using both the standard solution and a solution in which the major mineral nutrients nitrogen. the volume of sand to be used in the experiments was fixed at 500 cm3. The choice therefore rested with sand/solution culture.398 RGR in a localflora are whole-plant dry weights at 5 weeks and 2 weeks respectively and T is the time interval. No differences with respect to final yield or R were detectable at P<0. F. From inspection of the curve of yield v. phos- phorus. Solution culture. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . therefore. although not suffering in this respect. There were clear and significant increases in final dry weight yield with each increase in pot size. A special container (Fig. assumed the more efficient hexa- gonal section. Four pot sizes over the range 115-2280 cm3 were used and the sand was supplied daily with an excess of full nutrient solution. transparent polyethylene tubing. 3 weeks. To provide a shoot enclosure the polyethylene walls of the container were continued upwards for 10 cm above the This content downloaded from 69. only 128 could be accommodated at once. Soil was rejected as a possi- bility because of its variable and uncharacterizednutrient and water supplying capacity. Tables 40. Helianthus annuus L. 41) was selected because of its reported suitability for the growth of a wide variety of crop species. Other important design considerations for this container were that it should provide a standardized rooting and aerial environment in which the roots and shoot of each seed- ling were completely isolated from those of its neighbour. Rooting-medium The choice of rooting-medium presented few problems. on this basis.167. Festuca gigantea.05 between treatments after four weeks' growth. was also rejected because of the difficulty of devising a single plant-support system suitable for a wide range of plant sizes and growth forms. It was thought that few. It was suspected that the levels of mineral nutrients supplied might be super-optimal for the less productive species. potassium..65. con- structed from 4-in (10-2-cm) flat. With this design it was possible to fit up to 360 into a growth room at one time and still leave space for a thermohygrograph and dishes of germinating seeds. Rooting-volume A test was made to determine the minimum volumes of sand and solution necessary to support the highest yields expected in the projected experiments. 1) was. when closely packed. were grown in the controlled environment room for four weeks under a temperatureregime of 20?/15? C with an 18-h day. Deschampsiaflexuosa. Design of container Conventional flowerpots use growth-room space inefficiently: at 500 cm3 each. if any. Nutrient solution The choice of a suitable mineral nutrient solution was governed by the need to provide the more productive species with more than adequate supplies of mineral nutrients. The 'Long Ashton' solution (Hewitt 1966. In order to examine this possibility four species drawn from various types of unproductive vegetation. sand volume it was clear that yields of sunflower of around 4 g dry matter could be expected under these conditions from a volume of 500 cm3 of sand.Seedlings of sunflower. This provided con- tainers with flexible walls which. calcium and magnesium were supplied at only one-fifth of the stand- ard rate.194 on Mon. native species would approach this yield after only four weeks' growth and.

1.194 on Mon. J. 1). This content downloaded from 69. foil lining lOAcm / Sj Polyethylene 10/cm /tubing / 500 cm3 sand __ . 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . GRIME AND RODERICK HUNT 399 surface of the sand. Vertical section through a seedling container. P.65. parts of adjacent seedlings with a drop in light intensity of only 18% between the mouth of the container and the surface of the sand.167. / / /Heat-sealed base / / /Supporting grille _ CD 1CD C Cf _ )O c_ = |[\~~~~ ~ \ Inverted plastic pot I \// Waterproof tray To waste FIG. General arrangement The containers were supported in rows on a PVC-coated heavy-gauge steel grille. / Glass-fibre cloth over ' ' / /drainage holes * */ / ': . At the top of this extension the tubing was turned back on itself for a further 10 cm and a lining of aluminium foil was inserted into the space between the two layers of plastic (see Fig. /. This device ensured a complete isolation of the aerial Lights 93 cm from sand surface I I /Seedling I // / /Aluminium /J/q .

167. Seeds were set out in Petri dishes on moist- ened Whatman No. extra to the main series. Bedfordshire. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . When containers were watered at this rate solution still emerged from the bottoms and tests indicated that any accumulated nutrients were flushed out by this process. An initial harvest. was carried out on these seedlings as they came in from the field.). It was subjected to further purification in poly- ethylene bins by an acid-washing procedure similar to that described by Hewitt (1966). the 'sheltered' position of the sand surface and the relatively high humidity in the growth room (maintained additively at a level continuously above 60% R.65.400 RGR in a localflora This system allowed completely free drainage from each container and ensured that the excess solution emerging from the bottom of the containers was not allowed to collect outside or to contact any other container. After each experiment the sand was re-used following a repeat of the whole acid-washing process. On the basis of this information seeds could be imbibed for different lengths of time so that simultaneous peaks of germination were attained by up to eighteen species prior to the start of each experiment. The adequacy of this rather low rate of watering was owing to the small exposed-surface/ volume ratio of the sand. Medium: Sand+ Hewitt nutrient solution Visible radiation at plant height: 38-0 W m-2 (0-0545 cal cm-2 min-') Day temperature: 20+ 0 5? C Night temperature: 15+0 5? C Day-length: 18 h to (06.00) Relative humidity: always above 60% Dates of experiments: 24 November 1967-24 May 1973 Design of experiments: 12-18 species x 4 harvests x 5 replicates.00 00. Randomization The containers were placed in two randomized blocks. or because the seeds had some special or lengthy pre-germination requirements which could not easily be provided. Procedure Sand treatment The sand used was a pure. 1 filter paper. in the environment described above and daily counts of percentage germination were made. Several woodland species came into this category. the non-porous sides of the containers. It was found that the sand in each tube could be kept uniformly moist by the addition of 20 ml of solution on alternate days. Leighton Buzzard. one on each side of the growth This content downloaded from 69. In these cases material for the experiments was provided in the form of newly-germinated seedlings transferred directly from the field to the experiment. washed silica sand obtained from Messrs Arnolds Quarries Ltd. Lemna minorwas grown from single fronds using the same techniques except that solution culture replaced sand. Germination In order that seedlings of comparable age could be obtained simultaneously for different species a simple germination test was previously performed on seeds of all species considered for inclusion in the series. At the time of collection these seedlings all possessed cotyledons in good condition and all had either no true leaves or the first one (or pair) visible.194 on Mon. Seedlings of some species which it was desired to include in the experiments could not be obtained in the normal way either because no seed was available. A summary of the experimental design and environmental conditions is set out below. H.

One hundred seeds were weighed together in an air-dried condition and a mean weight was obtained. J. for more than one species. Measurements Measurements of mean 'seed' weight were made prior to each experiment. This measurement should perhaps more correctly be called a 'disseminule weight' since in some species (e. on some occasions. The root measure- ment was made with roots in a straightened but not stretched condition. GRIMEAND RODERICKHUNT 401 cabinet. Leaf areas. Dry weights.194 on Mon.g. were included initially in the root weights and then. This precaution was taken only to minimize the possible effects of differences between species with respect to maintenance procedure. Although these are all described here only data on dry weights are dealt with in this paper. including grasses.Plants were removed from the sand and the distances from the sand surface to the furthest points of the root and shoot were measured. Spatial arrangementof containers Towards the end of each experiment a majority of the more productive species pro- This content downloaded from 69. This design allowed plants to 'settle-in' to the environment before any measurements were taken and was a sufficiently short period to ensure that even the most productive species were still near their exponential phase of growth at the end of the measurements. if any. The remains of the seed. Harvesting The duration of the experiments was fixed at five weeks with harvests at two. At the time of harvesting a number of measurements was carried out.167. On removal from the oven the roots and shoots were parted and weighed separately. A visual record of the sizes of the various species at different stages of growth was obtained by making silhouettes on 'UNAX' paper of whole repre- sentative seedlings for each species at each harvest. a previous calibration giving their equivalent areas. Poterium sanguisorba)the pericarp was included. Leaves were removed from each seedling before drying and silhouettes were made on 'UNAX' 3M3 semi-dry diazo blackline paper. Images of the leaves were then cut out and weighed. four and five weeks. For a few species it was not possible to harvest exactly at these times and harvests were taken within three days of the designated dates. three. In these cases the actual time intervals were recorded exactly and used in computation. Linear measurements. that in growth rooms of this design. The shoot measurement was made between the sand level and the growing point of the shoot in species of erect habit and between the sand level and tip of the largest leaf in other species. Some data have been analysed both with and without the seed dry weight component. P. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . Seedlings were planted one per container and selected for harvesting at random. Washed plants were placed in paper envelopes and dried at 100? C for at least 48 h.65. Whole-plantsilhouettes. spatial variation in environ- ment causes only minor differences in the performance of plants growing in different positions. Rorison (1968) has shown. detached and weighed separately.

No significant differencesin Rma could be detected at P < 005 between experi- ments. These new plants then grew for the remaining two weeks of the experiment and were harvested at the same time as the five-week harvest. therefore. the removal of early harvests had created sufficient space in the growth- room to allow spacing out of the remaining containers to ensure that no shading of adjacent smaller plants occurred. It seems likely that among the plants which were allowed to remain in the experiment and which therefore contributed to the total variability of the data there were some individuals which suffered less obvious checks on their growth.65.and five-week harvests. By this time. In order to increase the effective capacity of the cabinet in some of the experiments a 'staggered' planting schedule was adopted. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . Special problems During the course of the experiment. It was. Seedlings showing severe stunting and/or death of leaves were removed from the experiment and discarded. fungal attack or disturbance during the application of nutrient solution. Repeatability If data from a series of experiments are to be combined it is advisable. The initial planting was of material for the three-. This was apparently related to drying of the surface layers of the sand and was rectified by repeating the experiments using containers of sand which were allowed to stand in 10 cm of deionized water during the first week.402 RGR in a local flora duced sufficient shoot material to emerge from the mouths of their containers. Such a situation is unavoidable in this type of experiment where one cannot entirely separate the variability inherent in the seed population from that en- gendered by experimental conditions or techniques. experienced a high rate of seedling mortality during the first week after planting. four. The environment itself may be monitored by instruments which ensure an exact repetition of environmental conditions from experi- ment to experimentbut the technical and procedural repeatabilityis not so easily checked. to establish that the experiments have been carried out in a comparable manner. When some containers had been in the growth room for two to three weeks it was found that a crust of algae up to 2 mm thick had built up over the exposed upper surface of the sand. In one way this was an advantage because it prevented the displacement of the sand which tended to occur when nutrient solution was applied.g. Silene dioica was chosen for this purpose. This content downloaded from 69. decided that at intervals throughout the period of the investigation experiments would be repeated on one species as a check on the constancy of the experi- mental procedure and environmental conditions. even where there has been strict control of the environment.194 on Mon. a small proportion of the seedlings suffered setbacks in the form of desiccation. e. When the three-week harvest was removed from the growth-room the plants for the two-week harvest were introduced. Where it was sus- pected that this crust caused asymmetrical penetration of the solution into the surface layers of the sand it was broken into small sections with a mounted needle and good drainage was restored. Thymus drucei. however. A number of small-seeded species. The final value of Rmx presented for this species is a combined value obtained from a pooling of all such data (105 plants in all).167.

Although any log. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . as a result of various internal and/or external influences. In general terms. Other species had a high initial R which. no doubt. In some cases this decline was smooth.65. this particularform of analysis often imposed its own pattern on the values of R obtained. Hughes & Freeman (1967). Many showed true exponential growth over the whole of the period studied. with no statistically- significant departuresfrom log.-linear dry weight increases with time. Working along broadly the same lines as those of Hughes & Freeman (1967). although other applications were borne in mind.d Y/dX. was held at a steady level for some time and then finally decreased (a log. growth apparently possessed no clear-cut pattern and often appeared to be following an intermediate path between various of these trends. this analysis was not successful. Hunt & Parsons (1974) constructed a program principally for the analysis of the present data. the cubic's approximations to the simpler trends resulted in an artificially erratic behaviour of R (and other functions) with time.194 on Mon. over the period studied. the period two to five weeks after germination encompassed some part of an exponential phase of growth. GRIME AND RODERICK HUNT 403 QUANTITATIVE ANALYSIS Generalrequirements The analysis of the large body of data obtained from these experiments presented problems of its own. either uniformly or at a decreasing rate. P. A few species exhibited combinations of these trends: R began with a low value which then increased. even from reasonably uniform raw data. Much of this problem. plant growth-analysis functions of the form 1/ Y.-sigmoidal trends in the data were described adequately. Although in the great majority of the present species. J. Because of the scale of the operation a single. 338) have described the advantages which this approach enjoys (see Hunt & Parsons (1974) for a full bibliography). In the remaining species. With only four or five harvest occasions. Among others. Unacceptably large standard errors also resulted. however. stemmed from the application of relatively high-order regressions to data covering only a small number of harvest occasions: the program was originally constructed to serve a system of frequent small harvests (in this context it can produce generally-acceptableresults as shown by Hunt & Burnett (1973)). In this particular case the method was desirable because of its ability to smooth out any small deviations from general trends with time and to draw on all of the relevant data for each species in calculating individual values of R. Therefore development of a further computer program.167.-sigmoidal growth pattern). the precise patterns of growth obtained varied considerably from species to species. which suffered less in this respect. was undertaken. p. In general. declined towards the end of the period of observations. in others it was of increasing steepness. but the data themselves required a variety of analytical approaches. Approachesthroughregression analysis Rather than adopt the more conventional harvest-interval approach to the growth- analysis it was decided to calculate growth-functions from regressions fitted to the raw data. Z/ Y and 1/Z. Still other species began growth with low values of R which increased. This program derives R as the differential (slope) of a third-order (cubic) polynomial fitted to the natural logarithms of successive estimates of whole-plant dry weight. computerized pro- cedure was desirable. As a first attempt at an analysis of the data the computer program described by Hughes & Freeman (1967) was used.d Y/dX were derived from lines or curves This content downloaded from 69. Radford (1967) and Evans (1972.

their Fig. two degrees apart in twenty- eight cases and three degrees apart ("no-fit"/cubic) in one case. values have been calculated by a number of methods as a specific example of the ways in which different methods of treating the data can yield different results. be viewed with some suspicion. 210 were linear.-cubic regression. however. This value for 11. A simple approach which has been * Significancedecidedat P < 005 by a test of the ratio regressionmean square/residualmean square. apply to phases of growth in which R was limited by external factors. following Evans (1972)) and that given by the log. Derivation of Rmax When an analysis had been established which provided reasonably accurate descrip- tions of the experimental data a problem remained in deciding on which of the fitted values of R best represented the concept of Rmaxand provided the most legitimate and informative basis for the comparisons between species discussed in the introduction. sixty-eight were cubic and two were "no-fit". were one degree apart in forty-nine cases. 104 were quadratic. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . l(a) and (c) for the log.35R (prefixes indicate the period in days. The possibility of making estimates across the board at a common time or at a common dry weight was rejected because this would often involve comparisons between plants at very different stages of development and might.194 on Mon. cf. This content downloaded from 69.404 RGR in a local flora fitted to the natural logarithms of variates Y and Z v. a total of 384 fitted regressions (some subsequently rejected).-quadratic regression is this value significantly higher than that of the overall mean (P <0 05). In these 192 sets of data the orders of polynomial fitted to loge W and to logeLAwere the same in 114 cases. Comparisonswith conventionalanalyses Table 4 contains relative growth-rates calculated for Holcus lanatus. in some species. The two higher-order regressions indicate substantially higher initial values of R but only in the case of the log. time. progressions both in W and in leaf area. Analysis via this program removed much of the artificial pattern and variability imposed on most of the present results by the Hughes & Freeman program (see Hunt & Parsons (1974). The application of Fisher's (1920) formula (p. In this case the quadratic regression gives a significantly better fit to the data than does the linear. In preparation for a further publication on leaf area phenomena. but similarcalculations made between individual harvest occasions indicated values that initially were higher and finally were lower than this.167.65. A further facility giving only a mean and standard error was included for cases where no significant trends with time existed in the (logged) raw data. have been analysed by the Hunt & Parsons program in each of 192 sets of data. To quote Hunt & Parsons. 2(a) and (c) for the log. 'Of this total. in the present body of data the simpler trends with time are the most frequent and the logarithms of W and LA follow trends that are often similar but are by no means exclusively so. gives no significant improvement over the quadratic. therefore.1. X.* The cubic.- linear regression must. all decided at P <0-05.-quadratic regression and their Fig. 397) gave the overall value of R as 1-61week. their Figs 1 and 2). LA. whilst adding substantially to the size of the confidence limits on the fitted values of R. Here. For plots of logeW and R in this case see Hunt & Parsons (1974). A 'stepwise' regression procedure was incorporated in which the most apposite of a series of polynomials from the first to third order (inclusive) could be fitted independently to loge Y and to logeZ according to statistical tests built into the program.' Thus.

in species exhibiting exponential growth Rmaxis equal in value to R. P. since the capacity. for rapid growth (over however brief a period) is itself a phenomenon of physiological and ecological interest.194 on Mon. and these are reviewed in the discussion. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . of the quadratic curves and for some of the cubic curves the maximum slope was. It is merely the fitted slope at a point close to this maximum (although for all Table 4. The figure in bracketsis the 95% con- fidencelimit. In particular. Relative growth-rate calculatedfor Holcus lanatus by various methods Relative growth-rate(week-1) Fisher's(1920) formula: 11-35R 161 11-20R 1-75 20-28R 1 59 28-35R 1. quadraticregression: 11R 2-01 (0-28) 2oR 1-70 (0-12) 28R 1-40 (0-12) 35R 111 (0-28) LogeW v. linear regression: 11-35R 1-56 (0-10) LogeW v. time. cubic regression: 1R 2-08 (0-69) 20R 1-68 (0-22) 28R 1-38 (0-21) 35R 1-18 (0-69) The notation follows that of Evans (1972) in which R and R respectively representinstantaneousrelative growth-rateand mean relative growth-rate overa statedperiodof time. Rmaxis an instan- taneous rate and is not necessarily sustained for any substantial period of time. time. Furthermore. This procedure is not without its limitations. Notwithstanding the foregoing difficulties these approximations to Rmaxhave been used in a first attempt to explore the possible ecological significance of this quantity.-curvilinear growth is exhibited. Cases in which the analy- ses have produced what might be considered as anomalous results will be discussed individually.46 LogeW v. at the first harvest occasion).The prefixesindicatethe timesor periodsinvolved (measuredin days from planting).65. Neither is this estimate of Rmaxinvariably the value of the true maximum slope of the log W curve. or lack of capacity. time. This content downloaded from 69. GRIMEAND RODERICKHUNT 405 explored here is to take the highest value of R obtained for each species during the period of observations. J.167. in fact. where log. however.

01 1-32 1-16 0-07 1.11 Glyceriafluitans 1-33 0. sylvaticum 1-35 0-13 1-06 1-63 1-02 0-04 0-94 1-12 Briza media 1.09 0.406 RGR in a localflora Table 5.01 0-67 0-12 0-39 0-94 Festucagigantea 1-44 0-20 1-02 1-85 1-04 0-06 0.10 1-30 1-72 1-17 0-04 1-10 1-25 G. pratensis 1-15 0-08 0-98 1-33 0-76 0-12 0-50 1-03 Carexflacca 1-38 0-05 1-28 1-47 1-26 0-04 1-17 1-35 C. confidence limits (CL) are given at P<0-05 Rmax R Value Standard Lower Upper Value Standard Lower Upper error CL CL error CL CL Acerpseudoplatanus 0-34 0'09 0.19 C.11 1-33 Agrostiscanina 1-41 0-08 1-23 1-59 1-41 0'08 1-23 1-59 A.96 0.08 1-27 1-61 Chenopodium album 2-12 0-34 1-39 2-85 1-25 0.19 1.10 0-97 1 40 Dryas octopetala 1-12 0-22 0-65 1-59 0-58 0-07 0-43 0-74 Epilobiumhirsutum 1-83 0-06 1-70 1-96 1-83 0-06 1-70 1-96 Eriophorumangustifolium 0-71 0-14 0-41 1.09 083 1.99 0-81 0-08 0-63 0'99 Digitalispurpurea 1-21 0-15 0-87 1-54 1-21 0-15 0-87 1-54 Draba muralis 1-24 0-07 1 09 1-39 1-18 0. Values obtainedfor Rma.90 0-21 0-47 1-33 0-48 0-07 -0-34 0-62 Galeobdolonluteum 0-75 0-06 0-63 0-88 0-75 0-06 0-63 0-88 Galiumaparine 1-51 0.09 114 1-52 1-33 0.01 0.11 1-61 1-36 0-12 1.91 116 F.10 1-07 1-51 Anisanthasterilis 2-28 0-12 2-03 2-53 1 43 0-07 1 29 1-57 Anthoxanthum odoratum 0-94 0-08 0-77 1.11 0-06 0-97 1 24 Callunavulgaris 0-35 0-05 0-25 0-45 0-35 0-05 0-25 0-45 Campanularotundifolia 0-81 0-17 0-45 1-17 0-81 0-17 0-45 1-17 Cardamineflexuosa 1.11 1-33 1-21 0-05 1.01 0. maxima 1-08 0-12 0-82 1-33 1 08 0212 0-82 1-33 This content downloaded from 69.34 0.10 1-38 A.09 085 1-20 B.01 1-32 G.01 1 27 Geumurbanum 0-73 0-15 0-35 1'11 0-73 0-15 0-35 1.11 1-61 Airapraecox 0-87 0-07 0-73 1 00 0-87 0-07 0-73 1 00 Alopecurusgeniculatus 1-24 0-07 1.90 Agropyronrepens 1-21 0-05 1.09 1 49 1. stolonifera 1-48 0.11 0-42 0-93 0-68 0.09 171 2-06 1-30 0-12 1-06 1-55 Brachypodium pinnatum* 1-03 009 0-85 1-20 1-03 0. saxatile 1-52 0-25 0-97 2-06 0-79 0'10 0-59 1 00 G.10 1-26 1-70 A. panicea 0-68 0.11 1-71 2-21 1-70 0.11 0-42 0-93 Catapodiumrigidum 1 60 0-18 1 23 1-98 1 09 0-06 0-95 1-22 Centaureanigra 1-13 0-07 0-98 1-29 1-13 0-07 0-98 1-29 Cerastiumholosteoides 1 46 0.16 0-52 Achilleamillefolium 1. verum 1-12 0-14 0-82 1-41 1-12 0-14 0-82 1-41 Geraniumrobertianum 1-14 0-06 1 01 1 27 1-14 0-06 1. pratensis 1-29 0.11 0-06 0-97 1 24 1. rubra 1-18 0-07 1 03 1-34 1-18 0-07 1 03 1-34 Filipendulaulmaria 0-95 0-12 0-64 1-21 0-95 0-12 0-64 1-21 Fraxinusexcelsior 0.10 1-26 1-70 1-48 0.09 126 1-65 Chamaenerion angustifolium 1-44 0-08 1-27 1-61 1-44 0.09 114 1-52 G.16 0-52 0.194 on Mon. ovina 1 00 0-07 0-86 1-14 1 00 0-07 0-86 1-14 F. tenuis 1-36 0-12 1.10 1-07 1-51 1-29 0.09 083 1.palustre 1-16 0-07 1. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions .10 1-03 1-47 Cirsiumvulgare 1-59 0-25 1-03 2-14 0-89 0-14 0-58 1-21 Clinopodiumvulgare 0-71 0-04 0-62 0-81 0-71 0-04 0-62 0-81 Convolvulusarvensis 2-44 0-26 1-84 3 03 1-36 0-13 1 07 1-64 Cynosuruscristatus 1-54 0-06 1-41 1-67 1-54 0-06 1-41 1-67 Dactylisglomerata 1-31 0-07 1-16 1-45 1-31 0-07 1-16 1-45 Deschampsiacespitosa 1-45 0-06 1-33 1-57 1-45 0-06 1-33 1-57 D.167.11 0-94 0-08 0-77 1.10 1-38 1 24 0-07 1. flexuosa 0-81 0-08 0-63 0. and R (week-l).09 126 1-65 1 46 0.65.11 Anthriscussylvestris 0-52 0-07 0-38 0-66 0-52 0-07 0-38 0-66 Arabishirsuta 1-32 0-12 1 05 1-60 1-32 0-12 1 05 1-60 Arenariaserpyllifolia 1-22 0-14 0-92 1-52 1-22 0-14 0-92 1-52 Arrhenatherum elatius 1-30 0-07 1-15 1 46 1-30 007 1-15 1 46 Betulapubescens 0-80 0-06 0-68 0-93 0-80 0-06 0-68 0-93 Bidenstripartita 1-89 0.

squalidus 228 008 2-12 2-45 1-40 009 1-21 1-59 S.09 0-35 P.11 0-46 0-93 0-70 0.167. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . atrocinerea 1-06 009 0-88 1-25 1-06 009 088 1-25 Saniculaeuropaea 0-77 0-13 0-50 1-05 0-47 0-05 0-38 0-57 Sarothamnusscoparius 0-63 005 0-52 073 063 005 0-52 073 Scabiosacolumbaria 1-26 0-07 1.65.10 0-95 1-39 1-17 0.01 006 0-88 1-14 Lycopersiconesculentum 1-64 0-15 1-33 1-95 1-02 007 0-87 1-16 Matricariamatricarioides 1. convolvulus 1-92 0-26 1-37 248 1-35 008 1-18 1-52 P. squarrosus 1-62 0-08 1-46 1-77 1-42 007 1-27 1-57 Koeleria cristata 094 005 0-84 1 04 094 005 0-84 1 04 Lathyrusmontanus 0-49 0-06 0-37 0-61 0-46 0-06 0-34 0-58 L.11 1 42 1-13 0-06 1 00 1 26 Sedumacre 0-71 0-08 0-54 0-88 0-71 0-08 0-54 0-88 Seneciojacobaea 1-24 0-06 1-12 1-38 1-24 0-06 1-12 1-38 S.09 0-23 0-60 0-33 0-08 0-16 0-50 P. sylvestris 0-36 0-04 0-28 0-44 0-36 0-04 0-28 0-44 Plantagolanceolata 1-70 0.91 1'19 1-05 006 091 119 Rumexacetosa 1-71 0-15 1-39 204 1-36 005 1 26 1 47 R.05 006 0-92 1.10 1-50 1. laricio 0-33 0-05 0-21 0-44 0-29 0-05 0-18 0-40 P. GRIMEAND RODERICKHUNT 407 Table 5 (contd) Rmax R Value Standard Lower Upper Value Standard Lower Upper error CL CL error CL CL Helianthemumchamaecistus 0-70 0.11 0-46 0-93 Helictotrichonpratense 0-75 004 0-66 0-84 0-75 004 0-66 0-84 Heracleum sphondylium 0-58 0-06 0-47 0-70 0-58 0-06 0-47 0-70 Holcus lanatus 2-01 0-13 1 73 2-28 1-56 005 1 46 1-66 H.10 Menthaaquatica 1.11 007 0-96 1-27 Myosottssylvatica 1-15 0-06 1-03 1-28 1-15 0-06 1-03 1-28 Nardusstricta 0-71 0-07 0-55 0-86 0-71 0-07 0-55 0-86 Origanumvulgare 1-46 0-10 1-25 1-68 1-46 0 10 1-25 1-68 Oxalis acetosella 0-51 0-04 0-42 0-60 0-51 0-04 0-42 0-60 Phalarisarundinacea 1-24 0-12 098 1-50 1 24 0-12 0-98 1-50 Picea abies 0-42 0.11 0-59 1-06 0-83 0'11 0-59 1-06 Poteriumsanguisorba 1-35 0-18 0-97 1-72 0-92 0-06 0-80 1-04 Prunellavulgaris 0-86 0-07 0-72 1. mollis 1-44 007 1-28 1-60 1 44 007 1-28 1-60 Hordeummurinum 1-76 0-16 1-41 2-10 1-18 0-08 1 01 1-35 Juncuseffusus 1-03 0-08 0-85 1-21 1-03 008 0-85 1-21 J. sitchensis 0-22 0-05 0. pratensis 1-26 007 1-12 1-40 1-26 007 1-12 1-40 P.30 008 1-13 1-46 Miliumeffusum 1 11 007 0-96 1-27 1. obtusifolius 1-49 0-14 1-20 1-78 1-49 0-14 1-20 1-78 Salix cinereassp.pratensis 073 010 051 095 073 010 051 095 Lemnaminor 2-45 008 229 2-61 206 044 1-62 2-50 Leontodonhispidus 0-89 007 0-75 1-03 0-89 007 0-75 1-03 Loliumperenne 1. trivialis 1 40 0-12 1-15 1-66 1-40 0-12 1-15 1-66 Polygonumaviculare 1-43 006 1-32 1-55 1-43 006 1-32 1-55 P.00 0-86 0-07 0-72 1'00 Ranunculusrepens 1-39 0-16 1-05 1-74 0-93 0-06 0-81 1-04 Rorippanasturtium- aquaticum 1-05 006 0.10 0-95 1-39 Medicagolupulina 1'14 0-06 1.17 0.30 006 1-17 1-43 1-30 006 1-17 1-43 Lotus corniculatus 1.19 Luzulacampestris 1.01 1-27 1-14 0-06 1.10 099 005 0-88 1.01 006 0-88 1-14 1.90 1-40 008 1-24 1-56 P. acetosella 1-55 008 1-39 1-72 1-55 008 1-39 1-72 R.09 1-56 1-92 P.194 on Mon. major 1-61 009 1-42 1-80 1-61 009 1-42 1-80 Poa annua 2-70 0-20 2-28 3-11 1 74 0. J. vulgaris 1-63 0-17 1-27 1-98 0-84 006 0-72 0-97 Sesleriaalbicans 0-75 0-05 0-65 0-86 0-75 0-05 0-65 0-86 Sieglingiadecumbens 0-60 006 046 073 060 006 046 073 This content downloaded from 69.09 0-35 0-22 0-05 0.30 008 1-13 1-46 1. P.01 1-27 Melica nutans 099 005 0-88 1.19 1 05 006 0-92 1. persicaria 1-29 006 1-17 1-42 1-29 006 1-17 1-42 Potentillaerecta 0-83 0. nigra ssp.

This result agrees well with that of Jarvis & Jarvis (1964) who collected infor- mation on R for eleven woody species and reported values equivalent to 0-053-0-822 week. Both groups showed a distribution which was strongly single-humped (Fig. The distribution is strongly biased towards the right. 2 as histo- grams with a class-interval of 0-25 week-'. The overall distribution (Fig.v.65. Dryas octopetala. vitis-idaea 0-23 0-07 0. 2(f)).11 2-20 2-66 2-09 0'10 1-88 2-31 Succisapratensis 0-76 0-07 0-61 0. shrubs and undershrubs. so there is no real differ- ence between the lower limits of the distributions shown in Fig.19 0-06 1-07 1-32 1.g.).09 0-53 0-92 Trifoliummedium 0-82 0-04 0-74 0'90 0-82 0-04 0-74 0'90 T. 2(b) and (c)) and which extended over a wide range. Ulex europaeus). 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions .10 0-37 0-23 0-07 0'10 0-37 Veronicaarvensis 1-18 0-13 0'90 1-45 1-18 0-13 0'90 1 45 Violariviniana 0-65 0-07 0-50 0-79 0-65 0-07 0-50 0-79 Zernaerecta* 1 04 0-04 0-96 1. 2(b) are occupied by woody species (q. RESULTS Values for Rmaxand R (calculated from the linear regression of loge W on time) are given in Table 5. Standard errors are included to provide a simple index of the variability of these estimates for each species and 95%/confidence limits allow individual tests of significance to be made. came into this category. but the right-hand tail is slightly more extensive than that on the left. All fourteen of the woody species had low values of Rax (Fig. 2(a)) is single-humped. e.grassland This content downloaded from 69. Distributions of Rmaxwithin the present sample of species are given in Fig. Little difference emerged between Dicotyledones and Monocotyledones with respect to the distribution of Rax. All of the legumes studied had o1w to medium values of Rax (Fig. 2(e) shows the frequency-distribution for species which are normally annual in habit. The range of values. completely absent from the bottom three class-intervals and virtually absent from the fourth (Aira praecox only).19 0-06 1-07 1-32 Teucriumscorodonia 1 08 0'09 0-88 1-29 1 03 0-08 0-85 1-21 Thymusdrucei 0-72 0'09 0-53 0-92 0-72 0. Errors in the estimation of Rmaxwere not taken into account in preparingthese histograms and some of the higher values had wide confidence limits. Deciduous and coniferous tree species.91 Taraxacumofficinale 1. extending from Picea sitchensis to Poa annua. repens 1-26 0-08 1 10 1-42 1 26 0-08 1 10 1-42 Tussilagofarfara 1-32 0-08 1-15 1-49 1-32 0-08 1-15 1-49 Ulex europaeus 0-74 0-06 0-61 0-87 0-74 0-06 0'61 0-87 Urticadioica 2-35 0-06 2-22 2-48 2-20 0-05 2-10 2-31 Vacciniummyrtillus 0-52 0-08 0-35 0-70 0-52 0-08 0-35 0-70 V.91 0-76 0-07 0-61 0.167. This group included woody shrubs of low Rax (Sarothamnusscoparius.for growth in the seedling phase.11 * Includingdata suppliedby R. Law.11 1-04 0-04 0-96 1.408 RGR in a localflora Table 5 (contd) Rmax R Value Standard Lower Upper Value Standard Lower Upper error CL CL error CL CL Silene dioica 2-27 0-15 1-98 2-56 1-32 0-07 1-18 1 46 Stellariamedia 2-43 0.194 on Mon. 2(d)). The very lowest classes in Fig. 2(b) and (c). represents more than a twelve-fold difference in values of Rmax. Hence this right-hand tail may not be genuine and it would be unwise to speculate on the slight skewness of this distribution. Fig.

(e) annual species.194 on Mon.(c) Monocotyledones.(d) woody species.(b) Dicotyledones.167. This content downloaded from 69.65. P. The frequencydistributionof Rmaxin (a) the whole sampleand (hatchedarea).Class-intervalsare 0-25 week-1. J. GRIME AND RODERICK HUNT 409 30 ( (a) (b) 20 I0- U 30 (d) (e) Lf) 20 10- 0 I 2 3 0 I 2 3 0 I 2 3 Rma (week-') FIG. (f) Papiliona- ceae (legumes). 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . 2.

many of these differences are due largely to the present methods This content downloaded from 69.-linear model..410 RGR in a local flora species of low Rmax(Lathyrus spp. This is an inevitable consequence of the quantitative analysis adopted.65. This value was considered by Evans to be among the highest values of R ever recorded and yet. DISCUSSION Accuracy of the data Before considering the physiological and ecological significance of the data it is neces- sary to comment on the reliability of the estimates of Rmaxobtained. in the present investigation. Digitalis purpurea. The first is inaccurate curve-fitting owing either to variability in the data or to methodological limitations and the second is restriction of growth by the experimental conditions. Many of the values of Rmax in Table 5 are high in comparison with other values reported in the literature.Medicago lupulina. In view of the small size of this sample it would be unwise to attach a general significance to the distribution of Rmax within legumes. corresponding to a mean RGR over the period of 2-07 [week-l]' (work of Shamsi 1970). Festuca ovina.) and grassland species of low to moderate Rmax (Lotus corniculatus. Variations in environment apart. their Figs 1(c) and 2(c)).). since the highest values of R are more likely to be obtained from the early parts of log. cf. For example. Trifolium spp. Evans (1972) reported '[the dry weight of] seedlings of Epilobium hirsutum increased. Certain species also had relatively variable final yields where there had been seedling mortality during the experiment. The variability of the estimates of Rmaxgiven in Table 5 increases with the absolute value of this function. 85-fold in 15 days. Zerna erecta.-curvilinear regressions than from the 'overall' log. Two possible sources of error can be identified. viz. the transformation of all values of W to natural logarithms before analysis had removed the systematic increases in variability which occurred with increases in W. No significant differences in Rmax were revealed within the three species for which more than one seed source was used (Brachypodiumpinnatum. depending as they do on a mixture of analytical methods. In other words.167. face new problems of heterogeneity of variance in the fitted growth-functions. Geum urbanum.Origanumvulgare. comparisons made across a large body of data such as this. leaving only the contributions of random and experimental errors. The estimation of slopes from curvilinear regressions becomes progressively less certain as the order of polynomial increases (see Hunt & Parsons (1974). For these species the values listed in Table 5 are derived from pooled data of all populations used. It is in the elimination of non-significant and unnecessary high-order terms in the regressions of loge W and logiLA on time that Hunt & Parsons' analysis represents an improvement on that of Hughes & Freeman (1967)..194 on Mon. this value is exceeded by Rmx in nine different species. see Table 1 footnote). Scabiosa colum- baria and Veronicaarvensis. Curve-fitting Some insight into the variability of the data was obtained by examination of the fitted estimates of final whole-plant dry weight (W): the variability of values obtained showed no systematic trend with absolute values when plotted on a logarithmic scale. It is considered that this is a small price to pay for the more realistic description of the trends in each of the sets of data that this analytical procedure provides. Nevertheless. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions .

This illustrates one of the natural disadvantages of the present 'broad-front' style of investigation. although many of these values are high.167. although our value of R for Epilobium hirsutumitself is lower (1-83 week. this phenomenon was most probably due to external constraints. The values given by Elkington (1971) for these two latter species were based on preliminary analyses only and should be replaced by the values listed in the present Table 5. The size of the values obtained thus suggests that the experimental conditions were suitable for near-maximal growth-rates in the more productive species even if only for a relatively brief period. Despite these sources of error and for the reasons discussed on pp.-curvilinear progressions of W on time and values of Rmx were taken from the early parts of the curves. However. Comparatively slow growth in tree c This content downloaded from 69. Even so. To have increased the number of harvests and the space available to each seedling would have reduced substantially the number of species which it was possible to include in the investigation. are likely to fall marginally below the true maxima. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . Restriction of growth by externalfactors In certain species growth curves flattened considerably towards the end of the experi- ment. In other species. Thephysiological basis for differencesin Rmax Rma and 'woodiness' In Fig. Where this coincided with the onset of flowering. Senecio vulgaris.the effect appearedto be a normal feature of development. J. however.). quoted results obtained for the growth of this species in the present investigation. Elkington (1971). e. These were apparent in species such as Poteriumsanguisorba. they give way to more moderate values when calculated over a more extended period. Hence. It was compared with values for Poa annua and Urtica dioica respectively equivalent to 1-90 and 2-27 week-l. Shamsi's result for E. fast-growing species. A decline in relative growth-rate occurred in many of the very highest-yielding species. A more likely explanation is that the roots became 'pot-bound'. Hordeum murinumand Lycopersicon esculentum. All of these cases exhibited significant log. again calculated from the present data over the whole period of growth.g. however. e. to bear in mind that the values of Rmaxpresentedhere are first estimates and particularly in the case of large. therefore. 2(d) it was clearly established that seedlings of the tree species included in the investigation yielded consistently low values of Rma. GRIMEAND RODERICKHUNT 411 of estimating R. each of which ultimately formed a rosette of large leaves which tended to experience mutual shading within the container. P.g. The over- all spread of final yields obtained in these experiments has. in presenting physiological data for a Biological Flora account on Dryas octopetala. Plantago major and Centaureanigra.In these species the growth form was such that mutual shading could probably be discounted.194 on Mon.65. in Bidens tripartita. more recent experiments indicate that some of the most rapidly-growing species achieve Rmaxprior to the 2-5 week period covered by this investigation. before flattening had taken place.Polygonumconvolvulusand Chenopodiumalbum. A value of R covering the whole period of observations (14-35 days after planting) and equivalent to 0-58 week-l was reported. suffered a check or truncation at the higher end of the scale. It is neces- sary. The values of Rmaxreported here are instantaneous values and are not necessarily sustained for any substantial period of time. 395-7 it seems reasonable to con- clude that broad comparisons of R are possible from the present body of data. hirsutumis still exceeded by R in two species here (Stellaria media and Urtica dioica).

simple leaves (e.are those which as estab- lished plants in the field are known to develop very long. Leaf form varies from large. However. In these plants.g.or higher is hetero- geneous with respect to seedling morphology. Deschampsiaflexuosa.g.g. leaf shape and the form of the mature plant. low Rmaxappears to be due to the translocation of photo- synthate into a swollen rootstock. Epilobiumhirsutumand Urtica dioica) and the shortest (e. With the exception of Festuca rubra. to expect that the values measured are a reliable guide to the rates of dry matter production commonly achieved in nature. appears to commence with the appearance of the first leaf. The fast-growing Dicotyledones include some of the tallest species (e. It would be naive.g. Rmaxandplant morphology Several plant attributes are known to exercise a controlling effect on relative growth-rate and. The grasses of low Rmaxinclude a high proportion of small tussock species. The grasses are invariably broad-leaved but both erect forms (e. The group of species which achieves values of Rmaxof 1-5 week.g. vitis-idaea). e.A further observation is that many of the slower growing species. as a preliminary to formal analysis some general points can be made with regardto the relationships between Rax and plant morphology.194 on Mon. those illustrated by Salisbury (1952)). e. Vaccinium myrtillus and V.65. all of the narrow-leaved grasses examined (Nardus stricta. e. Holcus lanatus and Bromus sterilis) and low-growing forms (e.167. The ecological significance of this pheno- menon will be considered later.g.g. techniques of growth-analysis will be used to assess the relative importance of leafiness and leaf efficiency in determining Rmx in each species. often swollen. Aira praecox and Koeleria cristata) had low values of Rmx. Dryas octopetala.g. However. Moreover. therefore.e. The ecological significance of differencesin Rmax Extrapolations to field conditions The estimations of Rmaxin this investigation were carried out on seedlings growing under conditions far removed from those obtaining in the natural habitats of the species concerned. Plantago major and Cerastium holosteoides). devoted to woody tissue during early seedling develop- ment. Even where seedlings are growing under productive conditions and in the absence of competition This content downloaded from 69. in a subsequent paper. Loach 1970) and has been attributed to the expenditure of photosynthate on woody tissue. in fact. Helian- themum chamaecistus. Rumex obtusifolius)to tripinnate structures (e. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . often in low rosettes. These are the Umbellifers Heracleum sphondyliumand Anthriscus sylvestris. Cynosurus cristatus and Agrostis stolonifera) are represented. Achillea millefolium). < 1 0 week. since all these species may be broadly described as microphyllous other explanations for the low rates of dry matter production may be involved. Thymus drucei. perhaps surprisingly.g. Among the slow-growing dicotyledons the majority of the species have small leaves. tap-root systems (see. Lotus corniculatusand Prunella vulgaris.412 RGR in a localflora seedlings has been recorded previously (Jarvis & Jarvis 1964. It is tempting to apply the same explanation also to the slow growth-rates of the six undershrubs examined (Calluna vulgaris. for example. a process which. With few exceptions the herbaceousspecies of low Rmax(i. it remains to be confirmed that a significantproportion of the dry matter of these species is. Viola rivinianaand Campanularotundifolia. Festuca ovina.or on creeping shoots. The slow-growing species include two which are exceptional in that they attain a large stature at maturity. a process concomitant with a slow rate of expansion of leaf area.1) are small in stature both as seedlings and as mature plants. Lotus corniculatus.

GRIME AND RODERICK HUNT 413 Table 6. Arable 2-43 11-00 18-55 19-69 8-44 15-54 22-83 23-54 +2-99 +8-61 +7-02 ?9-80 10. Lead mine heaps 11-65 23-00 11-89 2029 17-71 27-59 17-74 24-10 + 545 7-91 + 546 + 14-78 17.167. Themeanfrequency of occurrenceof species belongingtofour classes of Rma.65. in habitats 3-31 (Table 1) For each habitatand classof Rmaxthe upperfigurerefersto the meanpercentageoccurrenceof thesespeciesin quadratsof the habitat. Rock outcrops 6-45 12-11 7-92 6-73 13-28 18-53 14-60 13-59 + 3-19 + 488 + 3-28 3-98 6. Limestonescree 18-86 26-55 16-50 5-20 23-96 28-54 22-06 13-02 +6-47 + 1149 + 10-50 2-82 7. Coal mine heaps 6-38 6-50 14-30 7-80 12-82 13-78 20-35 13-35 + 6-77 + 3-66 + 499 + 7-71 16. Unshadedmire 4-77 7-87 8-85 700 11-92 14-88 15-33 13-79 + 263 3-92 + 3-66 + 474 4. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . Cinders 5-22 9-88 18-16 13-06 12-25 16-29 2350 19-16 + 424 + 499 + 5-25 + 5-69 This content downloaded from 69. Habitat Rmaxclass (week-1) < 1-0 1-0-1-24 1-25-1-44 > 144 3. P.The lowerfigureis the sameafterangulartransformation and with 95% confidencelimits added. Limestonepastures 21-88 22-94 18-83 16-75 25-79 25-99 23-88 21-34 + 5-83 + 8-46 + 5-06 11-48 13.194 on Mon. Shadedmire 4-33 8-44 12-24 10-63 10-79 15-27 18-56 16-66 +3-64 +6-33 +5-11 +?904 5. Meadows 44-40 34-33 43-69 59-00 41-78 34-22 40-61 50-42 +11-67 +20-14 +10-81 +32-73 11. Soil heaps 6-17 9-88 21-13 17-41 13-23 15-27 25-35 22-02 +6-80 6-24 + 554 6-83 15. Walls 1-83 5-18 6-80 4-11 7-42 10-91 12-78 10-31 +1-54 +4-14 +3-69 +2-91 9. Enclosedpastures 7-27 15-36 28-86 29-43 14-12 21-31 30-31 31-33 +4-23 +6-31 + 758 + 14-10 12. J. Pastureson acidic strata 22-64 17-20 10-31 2-83 25-25 21-48 15-75 9-30 +15-10 20-65 +7-31 314 14. Cliffs 5-13 825 560 407 11-65 14-99 12-14 10-70 + 359 +523 +3-21 +2-83 8.

Bricks and mortar 4-00 14-33 20-38 23-00 11-30 19-63 24-57 26-81 +6-88 +9-83 +?558 +8-13 19. Quarryheaps on acidic strata 15-20 550 1000 - 21-40 13-55 18-27 ?13-36 +8-26 8-48 22.414 RGR in a localflora Table 6 (contd) Habitat Rmaxclass (week-1) <1-0 1 0-1 24 1-25-1-44 > 144 18. Manureand sewage waste . Limestonewoodlands 12-22 5-10 11-00 700 18-73 11-97 16-68 13-36 +7-41 +?415 + 656 + 1090 25. Broad-leavedplantations 5-20 2-11 7-50 5-71 10-96 7-98 14-46 12-76 + 600 + 200 + 420 5-49 27. Limestonequarryheaps 10-54 18-88 20-65 12-54 16-76 23-81 24-76 18-77 +6-40 + 683 ? 644 + 655 21. Woodlandson acidic strata 8-60 2-73 6-82 6-20 15-21 9-03 13-02 13-66 +6-29 +2-10 +4-41 +6-67 26.167. 1343 17-78 22-85 20-16 23-65 26-67 + 854 + 509 ? 820 20. Wastelandand heath on limestone 10-60 17-50 12-56 11-36 17-18 21-79 18-20 17-55 +3-77 +7-10 ?4-71 +6-90 31. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions .65. Scrub 8-07 2-62 8-21 5-25 14-73 8-78 15-08 11-71 + 429 + 197 + 3-70 + 5-73 23. Hedgerows 33-33 17-00 27-00 17-14 33-33 22-40 29-66 22-50 +4411 + 16-51 +?981 + 1312 24. Paths 3-77 985 17-14 17-36 10-87 16-46 22-61 20-80 + 170 + 550 + 5-01 + 10-54 30.19 460 12-40 10-56 15-86 11-17 +5-31 +3-20 +3-47 +3-20 This content downloaded from 69. Coniferousplantations 6-00 2-14 685 440 11-74 7-90 13-12 11-14 +854 +2-86 +5-13 +6-72 28. Verges 7-59 15-93 21-14 14-27 13-83 20-48 25-28 20-47 +4-69 +822 + 596 ? 657 29. Wastelandand heath on acidic strata 7-00 455 9.194 on Mon.

but it is already clear that high yields may coincide either with a combination of high seed weight and high Rmax(e.g. A significant reduction (P<0-05) in the frequency of slow-growing species (i. It is apparent also that species of low Rmaxattain only low frequency on paths. Festuca ovina. Fig.g. 15. Poa annua and Polygonum spp. Urtica dioica and Stellaria media). J. < 10 week-1) was detected in a number of productive habitats. 10.seed weight and yield after five weeks' growth in the present experiments will be examined in a later publication. Grime 1966). 3. In Table 6 species have been classified according to Rax into four equally-populated class-intervals and the mean frequency of occurrence of the species in each class has been calculated for twenty-nine of the thirty-one habitats represented in Table 1.e. However. From Table 6. it is clear that there is a statistically-significantreduction in the frequency of species of high Rma (i. GRIMEAND RODERICKHUNT 415 such as in the early stages of recolonization of disturbed ground it is necessary to recog- nize that plant characteristics apart from Rmaxwill exert a major influence upon dry matter production. Such a relationship is suggested by the fact that species associated with fertile habitats such as arable fields (e. Nardus stricta. Stellaria media and Urtica dioica) have higher values of Rmax than species characteristic of unproductive habitats such as unfertilized pastures (e. Hordeummurinumand Polygonum convolvulus)or with that of small seed weight and exceptionally high Rmax (e. 3 the contribution of annuals to each class of Rax in selected habitats has been distinguished by hatching. These include various types of spoil. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . 29 and 31) the in- crease in the contribution of annuals. 3 allows a closer inspection of the nineteen habitats in which there is evidence of a skewed distribution in Rmax. limestone scree and unenclosed pastures on acidic strata. 19.e. is generally of secondary importance to the increase in abundance of perennial plants of high Rmax. the possibility remains that values of Rm. therefore.Helictotrichonpratense and Sesleria albicans). 1964.g.167.g. there is confirmation that productive and un- productive vegetation types differ with respect to the potential maximum growth-rates This content downloaded from 69. Chenopodiumalbum. These figures show clearlythat in each of the habitats in which there is a bias among the commoner constituent species towards high Rmax(Habitats 9.g. 14. 17. it has already been noted (Fig.194 on Mon. P. The data collected in the present investigation provide an opportunity to test the strength of this correlation.65.) or manure heaps (e.. The indicatorvalue of Rax It has been acknowledged already that the intrinsic and extrinsic constraints on R in plants growing under natural conditions are such that it is pointless to extrapolate to the field in absolute terms from laboratory determinations of Rax. In the histograms of Fig. Sieglingia decumbens. 23. From Table 6 and Fig. 2). 18. This arises from the fact that in many of the fertile habitats the preponderance of fast-growing species also coincides with physical disturbance and the presence of a high proportion of annual plants which. Bradshaw et al. In exploring the correlation between site fertility and Rmaxa complication must be recognized. include many species of high Rmx. Angular transformations have been added to permit statistical tests. enclosed pasture and arable land. The relationships between Rmax. > 15 week. where it occurs.) in the flora of two unproductive habitats. are a clue to the relative productivity of species growing under field conditions.g. In the early seedling phase the most important of these is the initial capital for growth provided by the seed reserves. 11. Correlations between potential growth-rate and site fertility have been com- mented upon or are evident in a number of publications (e.

. 13. Paths. 16.The system of numberingof the habitatsfollows that used in Table 1: 6. 17. Quarryheaps on acidic strata. Verges. Manure and sewage waste. Limestonequarry heaps.194 on Mon..ColumnR shows specieswhichweresimilarlyabundantin the habitat but which were not screenedfor Rma. 31.. Enclosed pastures. C.65..167. Coal mine heaps. 15. I DI i w A B C D R A B C D R (31) FIG. 1-0-124. This content downloaded from 69. 23..D. Limestonescree. Limestonepas- tures. Hedgerows.12. Arable. 1 25-1 -44.10. 20. 29. 30.28. Cinders.the hatchedareasreferto annual species. Wasteland and heath on acidic strata.. All specieswhich occurredin 10% or more of the quadrats are included. 11. 21. B. Soil heaps. 3. 18. Meadows. Pastures on acidic strata. 10. 14. The distribution of Rmax between four equally-populated class-intervals plotted for nineteenof the thirty-onehabitatsgiven in Table 1. (6) (9) (10) I0 20 (I )2) (13) 10- (14) (15) j (16) t (17) (18) (19) (20) (21) (23) 10 II^ 7 . Bricks and mortar. The class-intervals used are A. Wasteland and heath on limestone. > 1-44week-'1. 19. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions .Throughout.9. Lead mine heaps. < 1-0.

In order to explain the marked differences between common vegetation types with respect to the potential growth-rates of the component species it is necessary to consider the significance of Rmaxin relation to the very different processes which may determine the success or failure of species in different types of habitat.. In the ruderal. vegetation apparently excluding the majority of other plants by means of their superior competitive ability above and below ground (but see Pigott (1971) for an alternative interpretation of the performance of U. This combination of attributes is conducive to the efficient capture of light. Hence. P. high Rmaxfacilitates rapid completion of the life-cycle rather than exclusive occupation of the habitat. at the other extreme. It is possible that genetic characteristicsconducive to rapid growth in productive conditions become disadvantageous when the same plants are subjected This content downloaded from 69. however. and in the expansion of leaf area.65. by annuals and short-lived perennials. dioica under productive conditions). Earlier in this discussion it was pointed out that high Rmaxcoincides with a wide range in morphology. At one extreme high Rmaxis associated with tall stature. GRIME HUNT 417 of the major constituent species. the significance of high Rma appears to be rather different from that which it has in com- petitive species. The need to consider the concomitants of high Rmaxis even more obvious when an attempt is made to explain the low incidence of potentially fast-growing species in unproductive habitats. Here. It is likely. mineral nutrients and space and is particularly well developed in species such as Epilobium hirsutum. It may be helpful to recognize Rma as one of the more easily quantified expressions of a group of physio- logical attributes which are under common genetic control or show parallel responses to natural selection. It is apparent that at the two extremes of this morphological range the capacity to achieve a high rate of dry matter production under favourable circumstances is incorporated into distinct strategies. Moreover. More precise comparisons on the basis of individual vegetation samples and in which the frequency of each species within the sample is used to calculate weighted means for Rmax(Grime 1974) suggest that in natural vegetation there may be a sensitive adjustment of general levels of Rmaxprevailing in a community in response to variation in site fertility. In the ruderal. that in ruderals of productive. great plasticity in deployment of photosynthate between root and shoot and a rapid rate of response to environmental variation especially in the extension growth of stems. These may include. it may be suggested that species of high Rmax possess other attributes which place them at an advantage in productive or disturbed environments.194 on Mon. With respect to both the competitive and the ruderal strategy the significance of high Rma may extend beyond the capacity for rapid dry matter production. A marked contrast to this 'competitive' strategy is provided. high Rm. account must be taken of the fact that the habitat types recognized for the purposes of these comparisons are extremely broad and are likely to be rather heterogeneous both in vegetation and in productivity. Harper 1961). the capacity for extensive lateral spread above and below ground and the tendency to deposit a dense layer of litter on the ground surface. for example. ANDRODERICK J. occurs in association with a relatively short life-history and a growth pattern in which a large proportion of the photosynthate is directed into seeds (Salisbury 1942. arable land high Rmx will confer a considerable advantage in situations where fertility allows rapid growth and the onset of competition occurs at an early stage of colonization. water. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . Chamaenerionangustifolium and Urtica dioica which frequently occupy ex- tensive areas of productive. petioles and roots. relatively undisturbed.167.

and perhaps most important. Clarkson1967. The resultsof the present investigationare consistentwith those of previousworkerswho have drawnattentionto the fact that habitatssubjectto environmentalstress.194 on Mon. Loach 1967. Therelativescarcityof speciesof low Rmax in productivevegetationsuggeststhat these plants have a lower competitiveability. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions .particularlyin the absenceof more extensiveevidenceon the lack of intraspecificvariationin Rm.With respectto failureundercertainother forms of stress.418 RGR in a local flora to environmentalextremes.Investiga- tions into the physiology of shade-tolerantplants (Grime 1966. Scabiosa columbaria and Poterium sanguisorba.Moreover.The explanationfor this findingappearsto be that species of low Rmax are ill-adaptedeitherfor the replacementof foliageby regrowthof establishedplantsor for rapidseedlingestablishmentin areasof baregroundcreatedby disturbance.stillunidentified. Beadle 1954. Parsons 1968b. there is a strong probabilitythat the pathologydiffersbetweenstressesand possibly also betweenspeciesexperiencingthe samestress(Loach1970).The first is a low respiratoryrate and the second is the extendedlife-spanof individualleavesandroots.slow-growingplantsmakemodestdemandsand arethereforelesslikelyto exhausttheresourcesof theirimmediateenvironment.However. Hutchinson 1967.Higgs & James 1969)and severeshading(Grime 1965a.167.65.thus allowingthe build-upof reserveswithinthe plant. the investigations whichhavebeencarriedout so far havenot identifiedthe featureswithinthe physiology of fast-growingspecieswhichaccountfor theirgreatersusceptibility. 1964.in manycases.These species are all plants of relativelyunproductivevegetation and are to be found in the field in associationwith speciesof ratherlow Rmax. Mahmoud& Grime 1974)suggest that species of low Rmaxpossess two characteristicslikely to preventrapiddeteriorationof plants in whichgrowthhas been arrestedby some form of environmentalstress.speciesof low Rmax may be betterfittedto surviveperiodsin whichlittle or no growthis possible.Loach 1970) tendto be colonizedby speciesof low potentialgrowth-rate..For this reasonno attempthas generallybeen made to commentupon the significanceof differencesin Rmaxbetween speciesof similardistribution. Among the documentedinvestigationsof the susceptibilityof fast-growingspeciesto stressare the resultsobtainedby Grime(1965b. In this case it appearsthat the This content downloaded from 69.underconditionsof protractedand extremestress.each of which has a moderatevalue of Rmax (in the range 1-3-1-6). As a corollaryto thesecommentson the ecologyof fast-growingspecies. Bradshawet al. Special cases The mainconcernin this paperhas been to examinethe generalrelationshipbetween Rmax and plant distributionand it wouldbe unwise. such as mineralnutrientdeficiencyand heavy metal toxicity. Thirdly. Thereareat leastthreewaysin whichinherentlylow ratesof growthcouldbe adaptive to conditionsof stress.in particularnutrientdeficiencies (Kruckeburg1954.In the firstplace. 1966)and Loach (1967)whichsuggest that inflexiblyhigh respirationratesare implicatedin the rapiddeteriorationand mor- tality of deeply-shadedseedlings.The plant characteristics which cause speciesof high Rmax to be vulnerableto environmental stressesare. theremay be lower rates of incorporationof photosynthateand mineralnutrientsinto structure. The first concerns the species Galium saxatile.Secondly. Slow-growingspecies are also infrequentin habitatssuchas pathsin whichthe vegetationexperiencesa high intensityof damageor disturbance.it is necessary to referto the adaptivesignificanceof low valuesfor Rmax. to regardthe single determinationgiven here for each speciesas definitive.two particularcasesmeritfurtherinvestigation. Hackett 1967.

T. The reverse was true of several stable. Neal and Carmen Rathey for technical assistance. in part. W. A second anomaly is provided by the two biennial Umbellifers. road verges).194 on Mon. The work was supported. unproductive habitats. SUMMARY Estimations have been made of the maximum potential relative growth-rate (Rma) attained in the exponential phase by 132 species of flowering plants including representa- tives from each of the dry terrestrialhabitats of the Sheffieldregion. J. Grasses and forbs included a wide range of growth-rates and in both. Curtis. Earlier in this discussion it was pointed out that the low relative growth-rate of the seedlings of these two species may be due to the translocation of photosynthate into the rootstock.) in the Department of Botany at the University of Bristol during part of the preparation of this paper. Despite their extremely low relative growth-rates both of these species achieve high frequencies of occurrence in three of the habitats dominated by species of high Rma (meadows. productive environment and fitted growth-curves were used to derive various growth-analysis parameters. all the species of low Rmaxexamined were species which as seedlings and mature plants tend to be small in stature. The adaptive significance of Rmaxand its contribution to the determination of her- This content downloaded from 69. Allen. Species of moderate Rmaxwere ubiquitous. With the exception of the woody plants and the biennial herbs. Parsons of the University of Bristol Computer Centre for computing assistance and to Professor E. J. P. Woody species exhibited a bias towards low values of Rmaxand a similar trend was evident among fine-leaved grasses.65. The period of growth between two and five weeks after germination was studied in a standardized. may in part be related to the relatively large seeds which confer an impetus to seedling development which is unusual among native British herbaceous plants. The possibility that Rmaxis of adaptive significancein the field was tested by examining the frequency of species of low or high Rmaxin vegetation samples from a range of habitat types. V. doubtless at the expense of leaf development. Hargreaves.H. M. G. The remarkableability of the seedlings of these two species to establish in vegetation composed of potentially fast-growing perennial species and simultaneously to build up a root storage organ. by the Natural Environment Research Council. a process which begins at a very early stage of seedling development and which provides the capital for production of the flowering shoot in the second year. A. Annual plants were most frequent in the high Rmax category. ACKNOWLEGMENTS The authors are grateful to Dr J. A. Anthriscussylvestris and Heracleum sphondylium. high values of Rmaxwere associated with a variety of growth forms. In several disturbed and/or productive habitats fast-growing species were pre- dominant and species of low Rmaxwere virtually or completely absent. Heracleumsphondylium and Anthriscussylvestris. A. 2 Dec 2013 13:17:26 PM All use subject to JSTOR Terms and Conditions . GRIME AND RODERICK HUNT 419 relatively high values of the three species are due to the occurrence soon after germina- tion of a very brief period of rapid growth which then gives way to rates more typical of species from unproductive habitats.167. Yemm for providing facilities for one of us (R. hedgerows. Thanks are due to I. Janet Buckhorn. Law for permission to quote their unpublished data and to R. Hodgson and R.

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