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	14	0.179	1.27
Cerastium fontanum	9	0.140	1.66		Agrostis gigantea	9	0.134	3.17		Myosotis arvensis	8	0.169	2.10
Cirsium arvense	4	0.097	3.25		Arenaria serpyllifolia	8	0.107	2.13		Poa annua	17	0.161	1.31
Vulpia bromoides	9	0.087	2.05		Medicago lupulina	3	0.100	3.19		Cerastium fontanum	3	0.121	3.55
Crepis capillaris	2	0.077	4.74		Anchusa arvensis	2	0.086	4.24		Anagallis arvensis	6	0.116	3.82
Bromus hordeaceus	1	0.076	4.90		Polygonum aviculare	5	0.085	2.35		Viola arvensis	7	0.114	2.21
Anagallis arvensis	5	0.065	2.39		Myosotis arvensis	7	0.085	2.60		Vulpia bromoides	3	0.109	2.82
Myosotis discolor	11	0.037	1.44		Crepis capillaris	5	0.081	3.27		Erodium cicutarium	4	0.083	4.40
Achillea millefolium	11	0.036	1.54		Cerastium glomeratum	6	0.058	3.17		Taraxacum officinale	4	0.081	2.67
Fallopia convolvulus	4	0.036	2.96		Chenopodium album	18	0.043	1.32		Chenopodium album	13	0.068	1.73
Veronica serpyllifolia	13	0.029	1.68		Myosotis discolor	4	0.038	2.59		Holcus mollis	2	0.067	4.30
Agrostis gigantea	5	0.028	2.29		Veronica serpyllifolia	7	0.031	1.79		Rumex acetosella	2	0.035	3.87
Sonchus asper	6	0.019	3.10		Cerastium fontanum	3	0.030	2.71		Geranium molle	14	0.035	1.77
Poa annua	4	0.018	3.81		Coronopus didymus	1	0.023	4.90		Holcus lanatus	1	0.028	4.90
Anchusa arvensis	1	0.015	4.90		Sagina erecta	2	0.016	4.77		Atriplex patula	1	0.024	4.90
Acer pseudoplatanus	2	0.010	3.47		Acer pseudoplatanus	1	0.014	4.90		Cirsium arvense	3	0.020	3.15
Holcus lanatus	1	0.009	4.90		Rumex acetosella	2	0.014	3.69		Coronopus didymus	5	0.019	2.95
Atriplex patula	1	0.006	4.90		Vulpia bromoides	1	0.013	4.90		Quercus robur	1	0.018	4.90
Chenopodium album	5	0.005	3.08		Cirsium arvense	2	0.012	4.55		Acer pseudoplatanus	2	0.017	3.40
Conyza canadensis	1	0.004	4.90		Atriplex patula	2	0.009	4.25		Veronica serpyllifolia	8	0.017	1.98
Juncus bufonius	2	0.003	4.23		Senecio sylvaticus	1	0.003	4.90		Fallopia convolvulus	2	0.009	4.25
Coronopus didymus	1	0.001	4.90		Conyza canadensis	1	0.003	4.90		Galinsoga parviflora	3	0.008	2.90
Rumex acetosella	1	0.001	4.90		Senecio jacobaea	1	0.003	4.90		Plantago major	2	0.006	4.27
Solanum nigrum	1	0.001	4.90		Geranium molle	2	0.002	3.39		Crataegus monogyna	1	0.006	4.90
Stellaria media	1	trace	4.90		Poa annua	2	0.001	3.59		Juncus bufonius	2	0.005	3.44
										Sisymbrium officinale	1	0.005	4.90
										Bromus hordeaceus	1	0.003	4.90
										Pinus sylvestris	1	0.002	4.90
										Polygonum aviculare	1	0.002	4.90
										Solanum nigrum	3	0.002	2.89
										Lamium purpureum	1	0.001	4.90
										Cardamine hirsuta	1	0.001	4.90
										Persicaria maculosa	1	0.001	4.90
										Aira praecox	1	trace	4.90
										Sagina erecta	1	trace	4.90
Total biomass		47.180	0.23		Total biomass		40.500	0.30		Total biomass		46.062	0.40
Tripleurospermum density		11.1	1.29		Tripleurospermum density		27.5	3.30		Tripleurospermum density		11.2	1.91
Species		17.417	0.17		Species		14.042	0.22		Species		16.917	0.17
Total species		43			Total species		43			Total species		53
B) Cultivated spring 2003
October					March					May
Species	Frequency	Mean	CV		Species	Frequency	Mean	CV		Species	Frequency	Mean	CV
Anisantha sterilis	18	7.121	1.180		Chenopodium album	24	16.832	0.599		Chenopodium album	24	16.347	0.615
Tripleurospermum inodorum	23	5.956	1.201		Anisantha sterilis	10	7.774	1.906		Capsella bursa-pastoris	24	7.626	0.580
Chenopodium album	19	5.579	1.081		Anchusa arvensis	3	4.524	2.823		Anisantha sterilis	7	4.503	2.002
Holcus mollis	9	4.511	2.184		Spergula arvensis	13	3.737	2.703		Spergula arvensis	18	3.877	2.349
Bromus hordeaceus	15	3.695	1.042		Capsella bursa-pastoris	24	2.882	0.502		Erodium cicutarium	6	1.272	2.737
Capsella bursa-pastoris	21	2.854	0.937		Bromus hordeaceus	7	2.760	2.123		Lamium amplexicaule	11	0.932	2.481
Myosotis arvensis	17	2.388	1.952		Veronica persica	23	2.237	0.614		Galinsoga parviflora	21	0.865	1.181
Arenaria serpyllifolia	19	1.857	1.197		Sonchus asper	18	1.865	3.981		Veronica persica	19	0.823	0.986
Spergula arvensis	5	1.404	2.920		Tripleurospermum inodorum	23	1.835	1.389		Agrostis gigantea	10	0.725	2.703
Sonchus asper	16	1.243	2.051		Fallopia convolvulus	11	1.158	1.676		Geranium molle	20	0.631	1.318
Achillea millefolium	14	0.980	2.809		Anagallis arvensis	13	0.663	2.150		Sonchus asper	14	0.388	1.522
Vicia hirsute	23	0.768	1.385		Elytrigia repens	1	0.344	4.899		Bromus hordeaceus	3	0.377	3.210
Cirsium arvense	7	0.743	4.612		Senecio vulgaris	3	0.255	4.272		Artemisia vulgaris	4	0.270	2.630
Fallopia convolvulus	4	0.604	2.690		Vulpia bromoides	3	0.241	2.820		Poa annua	14	0.255	1.946
Papaver dubium	16	0.503	1.992		Galinsoga parviflora	12	0.238	1.513		Holcus mollis	6	0.254	3.541
Anchusa arvensis	1	0.438	4.899		Matricaria recutita	5	0.218	2.928		Anchusa arvensis	2	0.159	4.322
Polygonum aviculare	4	0.415	3.995		Polygonum aviculare	11	0.205	1.956		Thlaspi arvense	1	0.142	4.899
Veronica persica	14	0.398	1.989		Viola arvensis	16	0.168	1.857		Fallopia convolvulus	5	0.110	2.349
Lamium amplexicaule	5	0.353	2.554		Achillea millefolium	12	0.138	2.912		Tripleurospermum inodorum	5	0.101	2.756
Rumex acetosella	4	0.295	4.543		Lamium amplexicaule	9	0.118	1.873		Coronopus didymus	2	0.090	4.533
Elytrigia repens	2	0.202	4.193		Sisymbrium officinale	5	0.114	2.493		Persicaria maculosa	6	0.063	2.343
Viola arvensis	11	0.163	1.921		Myosotis arvensis	7	0.110	2.262		Vicia hirsuta	3	0.061	4.230
Matricaria recutita	3	0.144	3.065		Vicia hirsuta	10	0.086	2.090		Achillea millefolium	10	0.048	1.602
Coronopus didymus	4	0.112	3.474		Senecio sylvaticus	4	0.080	2.974		Senecio sylvaticus	3	0.033	2.915
Veronica arvensis	11	0.105	1.496		Medicago lupulina	4	0.067	3.020		Myosotis discolor	2	0.029	4.109
Anagallis arvensis	10	0.096	2.055		Cirsium arvense	4	0.063	3.138		Holcus lanatus	1	0.026	4.899
Agrostis gigantea	8	0.088	2.397		Agrostis gigantea	7	0.055	2.070		Myosotis arvensis	3	0.024	2.903
Vulpia bromoides	3	0.083	4.325		Artemisia vulgaris	3	0.048	3.083		Senecio vulgaris	1	0.024	4.899
Solanum nigrum	9	0.070	1.787		Papaver dubium	5	0.022	2.637		Papaver dubium	3	0.023	3.645
Cerastium fontanum	3	0.068	4.472		Solanum nigrum	6	0.022	2.038		Viola arvensis	5	0.022	2.399
Medicago lupulina	5	0.057	3.921		Geranium molle	2	0.019	4.376		Cirsium arvense	5	0.020	2.912
Galinsoga parviflora	7	0.054	2.001		Rumex acetosella	5	0.019	3.348		Trifolium dubium	2	0.020	3.893
Arabidopsis thaliana	16	0.048	1.625		Coronopus didymus	4	0.013	2.915		Solanum nigrum	6	0.018	2.534
Atriplex patula	1	0.044	4.899		Crepis capillaris	3	0.013	3.991		Rumex acetosella	1	0.018	4.899
Myosotis discolor	4	0.034	3.718		Veronica arvensis	7	0.013	2.348		Medicago lupulina	2	0.007	3.417
Senecio sylvaticus	1	0.028	4.899		Arabidopsis thaliana	1	0.009	4.899		Arabidopsis thaliana	5	0.006	2.397
Lamium purpureum	2	0.028	4.532		Persicaria maculosa	1	0.009	4.899		Veronica arvensis	2	0.006	4.547
Poa annua	7	0.028	2.038		Poa annua	2	0.004	3.587		Anagallis arvensis	3	0.005	3.279
Crepis capillaris	1	0.027	4.899		Holcus lanatus	1	0.003	4.899		Crepis capillaris	1	0.003	4.899
Galium aparine	1	0.016	4.899		Arenaria serpyllifolia	2	0.003	3.388		Vulpia bromoides	1	0.003	4.899
Senecio vulgaris	1	0.010	4.899		Myosotis discolor	2	0.003	3.587		Lamium purpureum	1	0.002	4.899
Stellaria graminea	2	0.009	3.624							Arenaria serpyllifolia	2	0.002	3.822
Trifolium dubium	2	0.009	4.068							Cerastium glomeratum	1	0.002	4.899
Gnaphalium uliginosum	1	0.003	4.899							Matricaria recutita	1	0.000	4.899
Geranium molle	2	0.001	3.587
Rumex obtusifolius	1	0.000	4.899
Total mass		43.629	0.394		Total mass		48.977	0.308		Total mass		40.209	0.335
Chenopodium density		24.0	9.76		Chenopodium density		59.5	7.51		Chenopodium density		46.3	3.37
Species		15.500	0.135		Species		13.583	0.278		Species		11.917	0.198
Total species		46			Total species		41			Total species		44

   Notes: Frequency out of n = 24 quadrats per treatment (6 pseudoreplicates in each of 4 blocks), mean = mean dry mass per species (n = 24), CV = coefficient of variation in species dry mass (standard deviation in dry mass/mean dry mass) is a measure of within-plot heterogeneity (patchiness) in species abundance, total biomass is mean biomass per quadrat; species is the mean number of species within each 50 × 25 cm quadrat, and total species is the number of species found in all 24 quadrats receiving a particular timing of disturbance.