Ecological ArchivesE085-113-A1

Michael J. Crawley. 2004. Timing of disturbance and coexistence in a species-rich ruderal plant community. Ecology 85:3277–3288.

Appendix A. Tables showing (TableA1) the effects of autumn 2002 and spring 2003 cultivation on botanical composition at Pound Hill in July 2003, (Table A2) nested ANOVA for total biomass, and (Table A3) the effects of long-term timing of cultivation (October, March, or May) over the period 1992–2001 on species biomass in July 2003.

Table A1. The effects of autumn 2002 and spring 2003 cultivation on botanical composition at Pound Hill in July 2003. See Notes at end of table for details.

Autumn 2002 cultivation

Spring 2003 cultivation

Species

Biomass
mean

 

SE

Frequency
(%)

Species

Biomass
mean

 

SE

Frequency
(%)

               

Tripleurospermum inodorum

16.36

1.741

90.3

Chenopodium album

12.92

1.207

93.1

Anisantha sterilis

4.58

0.873

56.9

Anisantha sterilis

6.47

1.303

48.6

Papaver dubium

4.26

0.573

88.9

Capsella bursa-pastoris

4.45

0.448

95.8

Vicia hirsuta

3.51

0.602

63.9

Spergula arvensis

3.01

0.962

50.0

Veronica persica

2.53

0.409

79.2

Tripleurospermum inodorum

2.63

0.587

70.8

Capsella bursa-pastoris

2.11

0.381

83.3

Bromus hordeaceus

2.28

0.505

34.7

Veronica arvensis

1.90

0.177

100.0

Anchusa arvensis

1.71

0.901

8.3

Aphanes inexpectata

1.20

0.200

80.6

Holcus mollis

1.59

0.707

20.8

Arabidopsis thaliana

1.13

0.138

91.7

Sonchus asper

1.17

0.533

66.7

Holcus mollis

1.11

0.421

26.4

Veronica persica

1.15

0.152

77.8

Anchusa arvensis

0.83

0.471

12.5

Myosotis arvensis

0.84

0.339

37.5

Myosotis arvensis

0.75

0.203

50.0

Fallopia convolvulus

0.62

0.178

27.8

Arenaria serpyllifolia

0.72

0.141

56.9

Arenaria serpyllifolia

0.62

0.182

31.9

Sonchus asper

0.47

0.094

62.5

Lamium amplexicaule

0.47

0.172

34.7

Matricaria recutita

0.31

0.097

23.6

Erodium cicutarium

0.42

0.244

8.3

Agrostis gigantea

0.28

0.164

38.9

Achillea millefolium

0.39

0.193

50.0

Viola arvensis

0.27

0.065

43.1

Galinsoga parviflora

0.39

0.084

55.6

Anagallis arvensis

0.26

0.100

31.9

Vicia hirsuta

0.31

0.084

50.0

Myosotis discolor

0.19

0.052

48.6

Agrostis gigantea

0.29

0.137

34.7

Cerastium glomeratum

0.17

0.050

50.0

Cirsium arvense

0.28

0.234

22.2

Achillea millefolium

0.16

0.061

37.5

Anagallis arvensis

0.25

0.102

36.1

Polygonum aviculare

0.16

0.061

25.0

Geranium molle

0.22

0.066

33.3

Lamium amplexicaule

0.15

0.046

25.0

Polygonum aviculare

0.21

0.116

20.8

Fallopia convolvulus

0.15

0.086

16.7

Papaver dubium

0.18

0.073

33.3

Crepis capillaris

0.13

0.072

13.9

Elytrigia repens

0.18

0.128

4.2

Cerastium fontanum

0.10

0.034

20.8

Viola arvensis

0.12

0.031

44.4

Medicago lupulina

0.10

0.036

12.5

Matricaria recutita

0.12

0.053

12.5

Bromus hordeaceus

0.08

0.056

4.2

Rumex acetosella

0.11

0.091

13.9

Vulpia bromoides

0.07

0.025

18.1

Vulpia bromoides

0.11

0.053

9.7

Sagina erecta

0.06

0.040

22.2

Artemisia vulgaris

0.11

0.051

9.7

Poa annua

0.06

0.017

33.3

Senecio vulgaris

0.10

0.075

6.9

Sisymbrium officinale

0.06

0.054

2.8

Poa annua

0.10

0.036

31.9

Galium aparine

0.05

0.025

11.1

Coronopus didymus

0.07

0.038

13.9

Cirsium arvense

0.04

0.022

13.9

Thlaspi arvense

0.05

0.047

1.4

Chenopodium album

0.04

0.009

48.6

Senecio sylvaticus

0.05

0.020

11.1

Erodium cicutarium

0.03

0.025

5.6

Medicago lupulina

0.04

0.020

15.3

Taraxacum officinale

0.03

0.015

5.6

Veronica arvensis

0.04

0.012

27.8

Veronica serpyllifolia

0.03

0.005

40.3

Sisymbrium officinale

0.04

0.020

6.9

Rumex acetosella

0.02

0.010

6.9

Solanum nigrum

0.04

0.010

29.2

Coronopus didymus

0.01

0.008

9.7

Persicaria maculosa

0.02

0.011

9.7

Acer pseudoplatanus

0.01

0.006

6.9

Cerastium fontanum

0.02

0.021

4.2

Atriplex patula

0.01

0.009

5.6

Myosotis discolor

0.02

0.012

11.1

Holcus lanatus

0.01

0.010

2.8

Arabidopsis thaliana

0.02

0.007

30.6

Geranium molle

0.01

0.005

22.2

Atriplex patula

0.01

0.015

1.4

Quercus robur

0.01

0.006

1.4

Crepis capillaris

0.01

0.010

6.9

Juncus bufonius

tr

 

5.6

Lamium purpureum

0.01

0.009

4.2

Galinsoga parviflora

tr

 

4.2

Holcus lanatus

0.01

0.009

2.8

Conyza canadensis

tr

 

2.8

Trifolium dubium

0.01

0.006

5.6

Plantago major

tr

 

2.8

Galium aparine

0.01

0.005

1.4

Crataegus monogyna

tr

 

1.4

Stellaria graminea

tr

 

2.8

Senecio sylvaticus

tr

 

1.4

Gnaphalium uliginosum

tr

 

1.4

Senecio jacobaea

tr

 

1.4

Cerastium glomeratum

tr

 

1.4

Solanum nigrum

tr

 

5.6

Rumex obtusifolius

tr

 

1.4

Pinus sylvestris

tr

 

1.4

 
 
 
 

Lamium purpureum

tr

 

1.4

 
 
 
 

Cardamine hirsuta

tr

 

1.4

 
 
 
 

Persicaria maculosa

tr

 

1.4

 
 
 
 

Aira praecox

tr

 

1.4

 
 
 
 

Stellaria media

tr

 

1.4

 
 
 
 
 
 
 
 
 
 
 
 

Total

44.53

1.698

 

Total

44.27

1.831

 

Mean species

16.25

0.389

 

Mean species

13.67

0.373

 

Total species

59

 
 

Total species

53

 
 
 
 
 
 
 
 
 
 

Dominance p1

0.37

 
 

Dominance p1

0.29

 
 

   Notes: Mean = mean dry mass per species in n = 72 quadrats 50 × 25 cm (i.e., averaged over the 3 long-term timings of cultivation), SE = standard error of mean species dry mass, frequency (%) is frequency out of a maximum of 72 quadrats expressed as a percentage. Species are ranked according to their mean dry mass. tr = "trace" means an average dry mass less than 0.001 g per plot. Total is mean biomass per 50 × 25 cm quadrat (the average of the sum of all species' dry masses); species is the mean number of species per 50 × 25 cm quadrat, total species is the species richness across all 72 quadrats, dominance, p1, is the proportion of total biomass made up by the most abundant species. Nomenclature follows Stace (1997). Species shown in bold are not represented in the 20 most abundant species in the opposing cultivation timing.

 

Table A2. Nested ANOVA for total biomass. See Notes at end of table for details.


 
df
Sum of squares
Mean square
F value
Pr (F)

           
Error: Block          
   Residuals
3
0.124317
0.041439
   
           
Error: Month %in% Block          
   Month
2
0.087777
0.0438883
0.1577355
0.8575047
   Residuals
6
1.669439
0.2782398
 
 
           
Error: Cultivation %in% (Block/Month)
1
0.023272
0.0232722
0.098471
0.7608241
   Cultivation
2
0.899069
0.4495345
1.902091
0.2046483
   Cultivation:Month
9
2.127033
0.2363370
 
 
   Residuals          
           
Error: Within          
   Residuals
120
11.96894
0.0997412
   

   Notes: This shows that there was very little variation in total biomass from block to block (variance = 0.041), no significant effect of Month of cultivation 1992–2002 on total biomass (P = 0.86), and no significant interaction between Month of cultivation and timing of cultivation in 2002-2003 (P = 0.20). The main effect of autumn 2002 or spring 2003 cultivation on total biomass was also not significant (P = 0.76). The variance attributable to differences in total biomass between the pseudoreplicated quadrats within half-plots is measured by the residual variance of ln(total + 0.01) = 0.0997 (5 df in each of 24 half plots = 120 df) and is much greater than the variation from block to block (0.041).

 

Table A3. The effects of long-term timing of cultivation (October, March, or May) over the period 1992–2001, on species biomass in July 2003; (a) following cultivation in autumn 2002; (b) following cultivation in spring 2003. See Notes at end of table for details.

A) Cultivated autumn 2002

October cultivation

 

March cultivation

 

May cultivation

 

Frequency

Mean

CV

   

Frequency

Mean

CV

   

Frequency

Mean

CV

Tripleurospermum inodorum

24

14.300

0.84

 

Tripleurospermum inodorum

24

23.762

0.49

 

Tripleurospermum inodorum

16

11.450

1.58

Vicia hirsuta

24

8.706

0.66

 

Veronica persica

24

5.412

0.78

 

Anisantha sterilis

16

8.319

1.25

Anisantha sterilis

19

5.102

1.00

 

Papaver dubium

20

3.385

1.29

 

Capsella bursa-pastoris

24

5.324

0.74

Papaver dubium

24

4.802

0.71

 

Veronica arvensis

24

1.574

0.73

 

Papaver dubium

19

4.379

1.48

Holcus mollis

15

3.262

1.73

 

Vicia hirsuta

14

0.994

1.68

 

Aphanes inexpectata

24

2.476

0.85

Myosotis arvensis

21

1.988

1.29

 

Capsella bursa-pastoris

23

0.748

0.83

 

Anchusa arvensis

6

2.398

2.81

Arenaria serpyllifolia

22

1.802

0.84

 

Anagallis arvensis

12

0.601

2.23

 

Veronica arvensis

24

2.205

0.85

Veronica arvensis

24

1.802

0.75

 

Viola arvensis

11

0.444

1.87

 

Veronica persica

21

1.945

1.09

Arabidopsis thaliana

24

1.317

0.65

 

Fallopia convolvulus

6

0.390

3.17

 

Arabidopsis thaliana

23

1.766

0.83

Aphanes inexpectata

23

0.948

1.29

 

Sonchus asper

20

0.386

1.28

 

Sonchus asper

18

0.913

1.05

Matricaria recutita

7

0.428

2.15

 

Anisantha sterilis

6

0.309

2.51

 

Vicia hirsuta

9

0.825

1.81

Polygonum aviculare

12

0.405

2.04

 

Arabidopsis thaliana

18

0.288

1.34

 

Agrostis gigantea

13

0.666

3.53

Veronica persica

12

0.280

2.00

 

Aphanes inexpectata

12

0.247

1.97

 

Myosotis discolor

20

0.497

1.35

Capsella bursa-pastoris

13

0.267

1.67

 

Achillea millefolium

9

0.193

2.56

 

Matricaria recutita

5

0.323

3.10

Cerastium glomeratum

15

0.263

2.52

 

Matricaria recutita

5

0.173

2.71

 

Arenaria serpyllifolia

11

0.284

1.71

Viola arvensis

12

0.202

1.58

 

Sisymbrium officinale

1

0.162

4.90

 

Lamium amplexicaule

8

0.275

1.96

Medicago lupulina

6

0.188

2.23

 

Lamium amplexicaule

10

0.153

2.31

 

Achillea millefolium

6

0.257

2.88

Sagina erecta

12

0.169

3.45

 

Bromus hordeaceus

1

0.150

4.90

 

Crepis capillaris

3

0.234

4.15

Galium aparine

8

0.150

2.31

 

Holcus mollis

2

0.138

4.85

 

Cerastium glomeratum