A modelling approach using seedbank and soil properties to predict the relative weed density in organic elds of an Italian

pre-alpine valley
` & G ZANIN* S OTTO*, M C ZUIN*, G CHISTE
*National Research Council (CNR), Institute of Agro-environmental and Forest Biology, Legnaro (PD), Italy, and Istituto Agrario San Michele allAdige (ISMAA), San Michele allAdige (TN), Italy Received 18 September 2006 Revised version accepted 27 March 2007

Summary
In 1996, a study was conducted on the seedbanks of a pre-alpine valley in northern Italy which had been organically farmed since 1986. The seedbanks were evaluated using soil cores taken from 16 organic elds located at various altitudes and seed numbers were determined using the seedling emergence method. Thirteen soil properties were also evaluated. In 2003, the germinable seedbank was assessed in ve other elds chosen at random. Soil properties were evaluated by the same method as in 1996. Using the data of the rst 16 elds as the analysis data set and those of the latter ve as an independent validation data set, a quadratic weed seedbank-soil properties model was built with partial least square regression analysis. The model estimates the relative abundance of the various species as the sum of

the contribution of individual soil properties and has a high predictive capacity. With a novel graphic approach, it is possible to describe the nonlinear relationship between each soil property and weed species relative abundance, giving a rational, quantitative, explanation as to why some species are widespread (e.g. Chenopodium album, Galinsoga parviora and Chenopodium polyspermum), whereas others tend to concentrate in specic elds (e.g. Spergula arvensis). The approach can, when some hypotheses hold, give a rational basis for the explanation of the relative abundance of species in a weed community and constitutes a useful methodology for study and research. Keywords: seedbank, germinable seed, soil properties, quadratic model, partial least square regression analysis.

` G & ZANIN G (2007). A modelling approach using seedbank and soil properties to predict OTTO S, ZUIN MC, CHISTE the relative weed density in organic elds of an Italian pre-alpine valley. Weed Research 47, 311326.

Introduction
The interest in linking soil quality and weed management derives from the belief that greater understanding of key soilweed relationships will lead to the design of agro-ecosystems with greater capacity and opportunity to suppress weeds (Gallandt et al., 1999). There is great interest in predicting the presence and spatial distribution of species from studies of the soil properties (Streibig et al., 1984; Andreasen et al., 1991; Milberg & Hallgren, 2000). Soil properties and weed abundance are known to vary spatially in agricultural elds and landscapes. However, the mechanisms giving rise to

spatial heterogeneity of weeds are poorly understood. Only recently have the weed management implications of precision agriculture increased interest in this topic, with the purpose of evaluating whether the abundance of weeds are consistently associated with a variety of site properties (Walter et al., 2002). These authors stated that the conclusions of other studies must be noted with caution because of the dierent methods used, for example Hausler and Nordmeyer (1995) reported that the distribution of Polygonum amphibium L. was similar to the distribution of high soil phosphorus concentration and clay content and low sand content. However, the distribution of Veronica hederifolia L. was similar to that

` , 16 - 35020 Correspondence: S Otto, National Research Council (CNR), Institute of Agro-environmental and Forest Biology, Via dellUniversita Legnaro (PD), Italy. Tel: (+39) 498272884; Fax: (+39) 498272818; E-mail: stefan.otto@ibaf.cnr.it 2007 The Authors Journal Compilation 2007 European Weed Research Society Weed Research 47, 311326

312 S Otto et al.

of sand content. Nordmeyer and Dunker (1999) found signicant correlations between Viola arvensis Murr. and pH, phosphorus and magnesium contents. They also found correlations between Stellaria media L. (Vill.) and organic matter, phosphorus and magnesium contents, amongst others. Recent multivariate analysis techniques (Kenkel et al., 2002) allow the qualitative relationships between abundance of species and soil properties to be identied, provided that the number of variables does not exceed the number of observations. A quantitative study of such relationships, for a data set with many variables but few objects, is possible using partial least square (PLS) regression analysis (de Jong, 1993; Rannar et al., 1994). The aim of this study was to determine whether weed seedbank composition could be predicted from soil properties. For this, a model was constructed that links weed seedbank composition and soil properties (analysis data set) using a specic regression analysis. An external validation was performed with an independent data set of germinable seedbank composition (validation data set). A method to represent graphically the quantitative relationships between soil properties and the relative abundance of weed species is then suggested.

cover crops in combination with one or two stale seedbed preparations. Rotation is the primary means for maintaining soil fertility and achieving weed, insect pest and disease control. The type of tillage used in the previous year, preceding crop, altitude and coding of the 16 elds are reported in Table 1.
The model

We developed a model of weed seedbank composition (Yi, dependent variables or response) and soil properties (Xj, independent variables or predictors) using a PLS polynomial regression, a technique that can explain one or more Y given one or more X, even with a small number of objects (observations, that are the rows of the matrices): Y1 ; Y2 ; . . . Yn f X1 ; X2 . . . Xm Error 1

Materials and methods

Site information

The Val di Gresta is situated in north-eastern Italy. It is a small valley (approx. 3000 ha), 4001300 m a.s.l., between Lake Garda and the River Adige. Average annual rainfall (over a period of 20 years) is 1192 mm. Average maximum and minimum temperatures are 13.6C and 4.5C, respectively, with high day night temperature uctuations. The soils lack or have very little gravel, are fairly deep and easily tilled, with variable organic matter content (2565 g kg)1) and cation exchange capacity. The pH also varies (4.68.3), even on a local scale, and the most uniform values are found at low altitudes, while the lowest values are found at high altitudes, probably due to accentuated leaching phenomena. Since the mid-1980s, horticultural produce has been grown on an ever-increasing area using organic farming techniques. The typical crops are autumn-harvested long-storage vegetables. Organically farmed crops currently cover more than half the arable land in the valley, amounting to approximately 150 ha. Weed management and cultivation practices (harrowing and inter-row hoeing) currently form the basis of weed control in the organically farmed elds. Thermal weeders are less widely used. Important weed suppression methods are

where Ys are weed species relative abundance in a certain eld (percentage of total weed number), and together constitute the Y-matrix. The Xs are the soil properties in the elds and together constitute the X-matrix. The Ys and Xs constitute the analysis set. The PLS algorithm, originally proposed by Wold et al. (1984), can analyse data with strongly collinear, noisy and numerous Xs (Wei et al., 2007) and works by breaking down the X-matrix as the product of two smaller matrices, much like principal component analysis (PCA): (i) the loading matrix, which contains a few vectors (the so-called latent variables, LVs) obtained as linear combinations of the original Xs; (ii) the score matrix, which contains information on the objects, described in terms of LVs, instead of the original variables. The main dierence is that PCA obtains the principal components that represent at best the structure of the X-matrix, whereas PLS obtains the LVs under two constraints: (i) they must represent the structure of the X-matrix and Y-matrix; (ii) they must maximise the tting between the Xs and Ys. More precisely, with the LVs, the original Xs are transformed to a set of x-scores (as in PCA). Similarly, the Ys are used to dene another set of components known as the y-scores. The x-scores are then used to predict the y-scores, which in turn are used to predict the response variables with a multistage process. An alternative approach could be univariate multiple regression, but in this case the focus is for the best t of each Yi independently, or the multiple linear regression. However, this technique works well when the number of objects is much larger than the number of variables and when the Xs are independent and uncorrelated. In this study a quadratic function f is used, which is exible and can approximate linear, exponential and asymptotic relationships. Therefore, given m soil

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A weed seedbanksoil properties model 313

Table 1 Sampling year, preceding crop and tillage, altitude, eld code in the 16 elds of the analysis set (A) and ve elds of the validation set (V). Fields sorted by ascending altitude Sampling year 1996 1996 1996 1996 1996 1996 1996 1996 1996 1996 1996 1996 1996 1996 1996 1996 2003 2003 2003 2003 2003 Altitude (m a.s.l.) 530 580 610 660 800 830 840 850 1000 1010 1030 1060 1180 1200 1220 1260 750 840 900 940 1160 Field code A0530 A0580 A0610 A0660 A0800 A0830 A0840 A0850 A1000 A1010 A1030 A1060 A1180 A1200 A1220 A1260 V0750 V0840 V0900 V0940 V1160

Preceding crop Carrot Cabbage Carrot Leek Savoy cabbage leek Potato Field pumpkin Carrot Chicory Carrot Potato Carrot Celery Potato Cauliower potato leek Cabbage Field pumpkin Field pumpkin Carrot Field pumpkin Cabbage

Preceding tillage Ploughing + roto-tillage Ploughing Ploughing No tillage No tillage Ploughing + roto-tillage Roto-tillage No tillage No tillage Ploughing + roto-tillage No till + manure Ploughing No tillage No tillage No tillage Roto-tillage Roto-tillage Roto-tillage Roto-tillage Roto-tillage Roto-tillage

Set A A A A A A A A A A A A A A A A V V V V V

properties, the relative abundance (Yi) of the i-th species is calculated as:
2 2 Yi I a11 X1 a12 X1 a21 X2 a22 X2 2 Error . . . am1 Xm am2 Xm

2 where I is the intercept and aij are regression coefcients; in our model we set I = 0. The algorithm used in our study assumes that the Xs and Ys have been normalised, separately for the analysis and the validation set, and was not constrained so that the sum of all Y variables must be less than or equal to 100%. For each species of the n species in the analysis set an R2 value is calculable: 2P 3 n ^i 2 Yi Y 6 i 1 7 6 7 R2 3 i 1 4P n 5 2  Yi Y
i 1

taking different numbers of LVs into account. For this the Predicted Sum of Squares Statistic (PRESS) is used. The value of PRESS for each species i in the validation set is: s v X ^ iN 2 =v 1 PRESSi YiN Y 5
i 1

^ iN are the observed and calculated relative where YiN and Y abundance in normalised units, and v the number of objects in the validation set. Then the overall PRESS of a set of n species is: PRESS
n X i1

PRESSi =n

^i is the calculated abunwhere Yi =observed abundance, Y dance and Y is the observed mean abundance. The average value of R2 is then: Average R2 Y
13 X i1

R2 i =n

When too many LVs are included in the model the average R2 (Y) will increase, but a serious overt will result and the model will have little or no predictive capacity for an independent data set of observations (validation set). It is then necessary to test the predictive capacity of the model

The PLS regression with a second-degree polynomial model was performed with STATISTICA (StatSoft Inc., 2005). For each n weed species, the PLS regression calculates two regression coefcients (of rst order, C1, and quadratic, C2) for each of m 13 soil properties, for a total of (n*m*2) coefcients. Because of the quadratic model between predictors (Xs) and responses (Ys) (Eqn 8), immediate interpretation of the effect of the regression coefcients could be difcult. The relationships between each soil property and the relative density of each weed species included in the model become clear when proper double normalised graphs are used. Obviously the validity of relationships is restricted to the range of the data set used. In such a graph, the normalised relative density of a species (y axis) is plotted against the normalised values of the various soil properties (x axis) of the analysis set.

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314 S Otto et al.

The normalised values (XN) are calculated from the original values (X) as: XN X Mso =Sso 7

where Mso is the mean and Sso the standard deviation of the soil properties in the analysis set. Furthermore, the calculated relative density of each species can be obtained as a sum of the partial relative density of this species as established by the contribution of the single soil properties. Let X be a value of a soil properties and XN the normalised value, let C1 and C2 be the regression coefcients of rst and second order for a species: then the abundance of the species (in normalised units, YN) as a consequence of the soil properties is given by the following quadratic equation:
2 YN C1XN C2 XN 1 1=u

where u is the number of objects in the analysis set, and the term 1 u is included in the PLS algorithm to take into account the size of the analysis set (Wold et al., 1989). This relative density can be interpreted as the contribution of the soil properties to the total relative density, that is then given by the sum of all partial abundance referable to all soil properties. The total relative density in original units is then calculated as: Y YN Ssp Msp 9

where Msp is the mean and Ssp the standard deviation of the relative density of the species in the analysis set.
Soil sampling and analysis

At the end of winter 1996, 25 soil samples were taken from each of the selected elds with a core sampler 7 cm in diameter by 25 cm depth, this number being considered sucient for estimating the semi-quantitative composition of the seedbank (Dessaint et al., 1996). Fields consisted of uniform plots 300-900 m2. In 1996 and 2003, another 10 soil samples were taken from each eld, bulked and analysed using the Italian standard methods for soil properties (Anonymous, 1999).
Evaluation of seedbank and germinable seedbank

The methods used to evaluate the ora diered in 1996 and 2003: the seedbank was evaluated in 1996, the germinable seedbank in 2003. For the 1996 soil samples, seedbank evaluation was done according to the seedling emergence method (Zanin et al., 1989). The individual cores were arranged singly on plastic trays. Seedlings were identied weekly and counted by species. The experiment, conducted in a temperature-controlled greenhouse, lasted for 18 months.

The seedlings from each core were summed and the seedbank was expressed as number of seeds m)2. The 1996 data set, e.g. soil seedbank in 16 elds estimated with the seedling emergence method, has been used as analysis set in the PLS regression model. In 2003, the germinable seedbank was monitored in ve sites within three permanently marked small plots of 1.0 m2 each, where, once or twice a week from March to October, weed seedlings were counted and removed. The method is analogous to that used by Zhang et al. (1998) in a similar study. Mickelson and Stougaard (2003) showed that in demographic research the use of permanently marked small plots is advantageous, because increasing the proportion of total area sampled improves precision. The sampled surface in 2003 was 0.331.00% of the total eld area. These same percentages could be obtained taking 195 soil cores with a 7 cm diameter core sampler. Under the assumption that germinable seedbank composition is a direct consequence of the soil seedbank, the 2003 data set, e.g. number of seedlings counted in the permanently marked small plots in ve elds, was used as validation set in the PLS regression model. On the basis of information and direct observations on crop management, we assume that the time lag between the two data sets (7 years) did not aect the link, if it exists, between seedbank and germinable seedbank, because (i) herbicides are not used in the selected elds; (ii) there is no irrigation; (iii) crop rotations are very varied; and (iv) general crop practices are unchanged. The selection pressure of crop management in those elds had been generally low and similar for all weed species. Finally, we hypothesised that in elds with this type of crop rotation and management, the seedbank is the result of various seed rains after dierent crop species (Beuret, 1984; Trresen & Skuterud, 2002), so there is no strong eect due to the last crop preceding the soil sampling. All the conclusions we draw are valid, if these hypotheses are true. The importance of each weed species within the seedbank community can be expressed by its relative abundance index (RAI), which is computed as follows: let C the counts of elds where a species has been found, N the total number of elds (16), then C N is the absolute frequency (AF). Given that 95 species are found, then 95 AF are calculable, and their sum is the total absolute frequency (TAF). The relative frequency (RF) is given by AF TAF. The relative density (RD) is the number of seedlings of a species as a fraction of total seedlings number, then RAI: (RD+RF) 2. The RAI accounts for both species density and pattern, thus limiting problems arising from weed patchiness (Derksen et al., 1993). The weed species nomenclature presented in the results follows Flora Europea (Tutin et al., 19641983).

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A weed seedbanksoil properties model 315

Results
Soil characteristics

The main physico-chemical characteristics of the 21 sampled elds were very variable (Table 2). 73.8 percent of values of the validation set fall within the range of the analysis set.
Flora characteristics Seedbank (Analysis set)

The total number of seeds ranged from 3473 to 19760 seeds m)2, with an average of 9370 4878. There was wide variability, both within and between altitudes: for example, 3473 seeds m)2 were found in eld A0850 and almost three times that (9921 seeds m)2) in A0840. The highest seedbank was 19760 seeds m)2 (A1060), equal to six times the poorest eld (A0850). Neither altitude nor tillage effects were remarkable.
Germinable seedbank (Validation set)

A total of 95 weed species were counted in the 16 sampled elds of the analysis set. The number of species per eld ranged from 25 to 41, with an average of 32.9 5.11 (standard deviation). Among these species, eight were found in at least 14 elds and three [Chenopodium album, S. media, Capsella bursa-pastoris (L.) Medik.] were common to all elds, whereas 14 were restricted to only one eld (rare species).

A total of 49 weed species were counted in the ve elds of the validation set. The number of species per eld ranged from 20 to 30 averaging 26.6 4.22. Among these species, 27 were found in at least three elds and 10 (C. album, Chenopodium polyspermum L., Galinsoga parviora Cav., Amaranthus spp. and others) were common to all elds, whereas 17 were restricted to only one eld [rare species, e.g. Digitaria sanguinalis (L.) Scop.]. The total number of weeds ranged from 476 to 1759 seeds m)2, with an average of 1210 612. Half of the

Table 2 Soil properties (X-matrix) in the 16 elds of the analysis set (A) and 5 elds of the validation set (V) Set A A A A A A A A A A A A A A A A A A A A V V V V V V V V V Field A0530 A0580 A0610 A0660 A0800 A0830 A0840 A0850 A1000 A1010 A1030 A1060 A1180 A1200 A1220 A1260 Mean SD Max Min V0750 V0840 V0900 V0940 V1160 Mean SD Max Min SA* 500 520 440 480 320 440 460 500 220 420 460 400 400 500 380 440 430 77 520 220 470 600(+) 270 520 430 458 123 600 270 SI 260 260 280 280 300 300 270 260 420 340 340 340 340 290 380 330 312 47 420 260 370 320 490(+) 400 490(+) 414 75 490 320 CL 240 220 280 240 380 260 270 240 360 240 200 260 260 210 240 230 258 49 380 200 160()) 80()) 240 80()) 80()) 128 72 240 80 OM 34.7 48.3 56.7 46.2 54.8 44.9 55.3 43.6 46.5 45.7 51.2 53.1 48.0 59.7 50.8 53.2 49.5 6.1 59.7 34.7 31.0()) 40.0 49.0 55.0 45.0 44.0 9.1 55.0 31.0 pH 8.00 7.85 7.80 7.50 7.55 7.35 7.70 7.95 7.90 7.67 7.80 7.65 5.70 7.15 4.85 6.00 7.28 0.93 8.00 4.85 7.89 7.14 7.44 7.73 7.68 7.58 0.29 7.89 7.14 CaCO3 139 1008 304 64 105 40 368 544 480 136 1444 240 60 232 31 60 328 392 1444 31 384 34 116 200 46 156 144 384 34 P2O5 72 37 101 89 119 129 204 49 116 117 142 104 34 116 32 180 102 50 204 32 192 24()) 7()) 30()) 124 75 80 192 7 K2O 468 117 420 245 396 240 480 162 195 207 178 207 390 152 65 162 255 131 480 65 208 540(+) 532(+) 150 438 374 183 540 150 CaO 12.32 13.44 12.04 11.48 15.68 9.86 16.80 11.48 12.94 12.88 12.94 12.99 4.40 14.84 2.52 3.58 11.26 4.21 16.80 2.52 7.06 11.12 7.74 6.72 2.86 7.10 2.95 11.12 2.86 MgO 760 480 500 540 1320 450 340 230 260 290 220 280 270 260 150 230 411 289 1320 150 225 972 381 237 544 472 308 972 225 B 0.385 0.265 0.395 0.400 0.620 0.320 0.445 0.310 0.350 0.375 0.485 0.810 0.300 0.250 0.340 0.300 0.397 0.143 0.810 0.250 0.300 0.450 0.600 0.420 0.370 0.428 0.112 0.600 0.300 CEC 17.0 19.5 20.7 19.7 26.8 20.1 23.0 19.1 25.0 19.6 22.4 24.9 20.9 21.9 20.0 21.0 21.4 2.5 26.8 17.0 20.3 51.7(+) 31.9(+) 26.3 20.0 30.0 13.1 51.7 20.0 CaCO3act 79 81 78 54 80 35 94 75 74 72 66 71 42 79 30 37 65 20 94 30 40 30 32 25()) 7()) 27 12 40 7

*Code = soil properties (units); SA = sand (g kg)1); SI = silt (g kg)1); CL = clay (g kg)1); OM = organic matter (g kg)1); pH = pH (H2O); CaCO3 = total calcium carbonate content (g kg)1); P2O5 = available phosphorus (mg kg)1); K2O = available potassium (mg kg)1); CaO = available calcium (g kg)1); MgO = available magnesium (mg kg)1); B = water-soluble boron (mg kg)1); CEC = cation exchange capacity (cmol+ kg)1); CaCO3act = active calcium carbonate content (g kg)1). Values of soil properties of the validation set which fall outside the maximum (+) or minimum ()) of the analysis set range are indicated.
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316 S Otto et al.

seedbank species were found in the germinable seedbank, because of the lack of rare species. Referring to the RAI ranking, taking into account the rst ten species of the analysis set (75% of total seedbank), eight were in the rst 10 species of the validation set (again, 75% of the total germinable seedbank). It means that a given correlation between seedbank and the germinable fraction exists only in terms of rank for the species with the highest relative density and relative frequency. Otherwise, relative densities of the species in the two data sets were almost uncorrelated. Signicant correlations were found in only 6 of 156 cases, i.e. between C. album and Sonchus oleraceus L. in the analysis set, G. parviora and Galinsoga ciliata (Raf.) Blake in the analysis and validation sets, D. sanguinalis and Amaranthus spp., Spergula arvensis L. and G. parviora and S. arvensis and G. ciliata in the validation set. In this study, the correlation between seedbank and germinable seedbank cannot be calculated, but is most probably very low or absent, given the additional eect of the low rate of emergence of viable seed content. For example, Zhang et al. (1998) found positive correlations, even if only 37% of the germinable seedbank was capable of producing seedlings in the eld. Barralis et al. (1996) found an average rate of emergence of 5.54%, with high variability between species. Therefore, simple correlation analysis may not be the right approach for constructing a weed ora composition using a set of available surveys.
Model variables

Fitting of the model and regression coefcients

The lowest value of PRESS was obtained with eight LVs (Table 4). The general predictive capacity of the model was good, with an average R2 (Y) of 0.68: for some species it was very good (e.g. G. parviora, C. polyspermum, S. media, S. oleraceus), for others less so (e.g. C. album, Amaranthus spp., V. persica, D. sanguinalis) (Table 5).
The double normalised graph and example of calculation of relative density

All 13 soil properties were taken as predictor variables, each entered twice in the regression model (at rst and second degree). So the X-matrix had 13 original variables (Table 2) but the model had 26 predictors. Even if PLS regression analysis can run with a number of response variables greater than that of predictor variables, for a model with a reasonable predictive capacity a choice has to be made between the 102 theoretical response variables, e.g. the complete weed species list,. In this study, the selected response variables were the main nine species in the 21 elds of the entire data set: C. album, G. parviora, C. polyspermum, Portulaca oleracea L., Amaranthus spp., S. media, S. arvensis, Chaenorrhinum minus (L.) Lange, D. sanguinalis, and four species not dominant in any eld but with overall importance, i.e. C. bursa-pastoris (RAI rank = 6), S. oleraceus (RAI rank = 7), Veronica persica Poir. (RAI rank = 9) and G. ciliata (RAI rank = 10). As a result, the Y-matrix had 13 variables (Table 3). These species are annuals, with no dispersal adaptations and a persistence of >1 year, except for C. minus (persistence <1 year) (Thompson et al., 1997).

It is possible to graphically represent the quantitative relationships between soil properties and the relative density of weed species. The algorithm calculated scaled regression coecients for the calculations of normalised predicted Y-values, and soil properties, except soil texture, that varied in very dierent ranges. Therefore, to display all the results (13 species 13 soil properties) in a simple graphical format, normalised units were used. In Fig. 1 the normalised relative density of C. album (y-axis) was plotted against the normalised values of the sand, clay, silt and organic matter content of the 16 elds of the analysis set (x-axis). In Val di Gresta elds, the sand content ranged from 220 to 520 g kg)1, with mean = 430 77 (Table 2). After normalisation, the range was from )2.742 to +1.175, i.e. the range of the solid line Sand. Figures 24 show the double normalised graph of the selected species for the 16 elds of the analysis set. The calculated relative abundance of each species included in the PLS regression model can be obtained as a sum of the partial abundance of this species as established by the contribution of the single soil properties. Consider, for example, the abundance of C. album in eld A0610. This eld has 440 and 280 g kg)1 of sand and silt, respectively, that are 0.131 and )0.685 in normalised units. The PLS regression model with eight LVs establishes that the abundance of C. album, in normalised units, in eld A0610 as a consequence of the sand content, is given by the following quadratic equation: C : album norm: units 0:075 0:131 0:004 0:1312 1 1=16 0:013 10

where )0.075 and 0.004 are the regression coefcients. The result )0.013 means that the sand content in this eld slightly reduces the abundance of C. album below the mean calculated on the whole analysis set (i.e. in normalised units, zero). Figure 1 shows clearly that the estimated C. album abundance declines monotonically with increasing sand content. This does not necessarily imply that C. album will

2007 The Authors Journal Compilation 2007 European Weed Research Society Weed Research 47, 311326

Table 3 Relative density (% of total density) of the 13 selected weeds species (Y-matrix) in the 16 elds of the analysis set (A) and ve elds of the validation set (V); relative density of other species and total seed number

2007 The Authors Journal Compilation 2007 European Weed Research Society Weed Research 47, 311326

RAI rank Field 5.9 7.7 8.9 40.5 0.5 1.1 0.1 1.2 0.0 23.8 20.0 1.1 0.0 0.0 0.1 0.0 6.9 11.6 40.5 0.0 9.1 3.2 2.4 5.6 3.5 4.7 2.7 9.1 2.4 0.9 20.1 7.9 0.9 11.3 6.6 62.7 12.3 3.3 2.0 0.5 0.6 3.5 4.1 0.2 0.2 8.6 15.5 62.7 0.2 2.2 1.0 2.8 1.3 0.6 1.6 0.9 2.8 0.6 1.1 0.0 0.5 0.8 9.6 11.1 9.2 6.3 4.8 0.6 2.5 0.3 12.2 0.5 1.1 0.0 3.8 4.4 12.2 0.0 0.2 8.2 3.7 3.2 3.7 3.8 2.9 8.2 0.2 0.6 2.6 0.1 15.4 10.0 5.8 5.8 4.2 7.2 1.9 4.7 4.0 11.2 2.3 15.7 5.2 6.0 4.8 15.7 0.1 0.6 3.5 2.0 2.1 4.4 2.5 1.5 4.4 0.6 0.3 0.0 0.1 0.5 0.9 0.3 1.0 0.6 0.5 0.4 0.5 25.3 5.9 0.3 0.4 0.0 2.3 6.3 25.3 0.0 2.3 4.1 3.4 3.5 1.4 2.9 1.1 4.1 1.4 7.6 0.3 0.0 0.1 0.9 0.5 0.0 0.0 8.0 1.9 15.5 0.1 0.5 0.8 0.0 0.0 2.3 4.4 15.5 0.0 0.0 0.6 1.5 0.0 4.0 1.2 1.7 4.0 0.0 1.4 30.7 1.6 7.7 0.0 0.0 0.0 42.8 0.0 1.6 0.0 0.1 2.1 0.1 0.0 0.0 5.5 12.5 42.8 0.0 0.0 0.4 0.0 0.0 0.0 0.1 0.2 0.4 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 20.7 1.1 27.1 3.1 8.2 27.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 0.0 0.2 0.1 20.8 0.3 0.0 0.0 0.0 0.1 0.0 0.2 1.3 0.0 0.0 0.0 1.4 5.2 20.8 0.0 0.0 0.2 0.0 0.2 0.0 0.1 0.1 0.2 0.0 5.2 7.7 0.0 0.3 1.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 5.8 1.3 2.5 7.7 0.0 27.1(+) 0.0 0.0 0.0 0.0 5.4 12.1 27.1 0.0

1 Chenop. album

2 Galinso. parvio.

3 Chenop. polysper.

4 Amarant. spp.

5 Stellaria media

9 Veronica persica

6 Capsella bursa-p.

7 Sonchus olerace.

10 Galinso. ciliata

11 Portula. olerac.

17 Spergula arvensis

19 Chaenor. minus

28 Digitaria sanguin.

Other species 16.0 13.2 16.1 13.8 32.8 56.5 13.6 25.2 25.3 15.7 4.3 2.4 26.1 40.9 28.1 61.2 24.5 16.7 61.2 2.4 44.7 50.7 28.9 26.8 2.5 30.7 18.8 50.7 2.5

Tot seed (n) 6739 3931 8528 9516 7790 3942 9921 3473 8226 16401 16557 19760 6490 9693 13593 5366 9370 4878 19760 3473 628 476 1485 1704 1759 1210 612 1759 476

A0530 A0580 A0610 A0660 A0800 A0830 A0840 A0850 A1000 A1010 A1030 A1060 A1180 A1200 A1220 A1260 Mean SD Max Min V0750* V0840 V0900 V0940 V1160 Mean SD Max Min

8.0 5.0 56.0 11.9 5.5 12.4 7.3 2.1 15.6 14.8 3.5 14.9 23.9 14.2 45.7 0.4 15.1 15.3 56.0 0.4 8.6 25.8 34.0 54.3 21.3 28.8 17.0 54.3 8.6

52.6 12.7 8.5 8.0 0.7 3.2 0.1 5.4 35.4 28.7 46.5 0.4 0.0 1.4 0.1 0.0 12.7 17.8 52.6 0.0 5.2 0.5 13.5 1.0 57.4(+) 15.5 24.0 57.4 0.5

0.2 0.0 0.0 0.0 5.1 2.4 0.1 0.0 0.0 8.6 2.0 50.7 13.3 14.8 7.7 0.0 6.5 12.8 50.7 0.0 0.2 2.0 7.8 2.0 1.2 2.6 3.0 7.8 0.2

A weed seedbanksoil properties model 317

RAI, relative abundant index. *Values of relative density of the validation set which fall outside the maximum (+) or minimum ()) of the analysis set range.

318 S Otto et al.

Table 4 Fitting model parameters with increasing number of latent variables (LVs) Average R2 (Y) 0.139 0.211 0.350 0.437 0.494 0.552 0.637 0.682 0.776 0.817 0.856 0.900 0.941 0.972 1.000

LVs (n) 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

PRESS 0.975 0.965 0.951 0.937 0.915 0.907 0.831 0.822 0.920 0.929 0.971 1.180 1.793 1.945 2.226

C. album relative density (norm. val.)

0.5

0.3

0.1

0.1

0.3 3

Soil properties (norm. val.)

Fig. 1 The double normalised graph of the relationships between relative density of Chenopodium album and soil texture ( sand; silt; clay).

PRESS, predicted sum of squares statistic.

Table 5 Fitting parameters of the selected species in the model with eight latent variables Species Chenopodium album Galinsoga parviora Chenopodium polyspermum Amaranthus spp. Stellaria media Veronica persica Capsella bursa-pastoris Sonchus oleraceus Galinsoga ciliata Spergula arvensis Portulaca oleracea Chaenorrhinum minus Digitaria sanguinalis Average (Y) R2 0.383 0.943 0.910 0.311 0.951 0.250 0.519 0.954 0.980 0.539 0.847 0.983 0.298 0.682 PRESS 0.841 1.025 0.381 0.801 0.811 0.833 0.709 0.864 1.129 0.579 0.882 1.024 0.808 0.822

The same calculation approach applies to all other soil properties. The nal relative abundance of C. album calculated for eld A0610 is therefore the result of the single contribution of all soil properties, some of them decreasing, others increasing the abundance with respect to the mean calculated for the whole analysis set. This same calculation applies to all species for all elds. Taking into account that a soil cannot have high sand, high silt and high clay contents all at the same time, it is therefore possible to rationalise why some species are widespread, while others tend to concentrate in specic elds.
Chenopodium album

PRESS, predicted sum of squares statistic.

dominate in elds with low sand content, because the effect of other soil properties can mask that of sand, indeed C. album abundance is also favoured by a high silt content and average clay content. The contribution of the silt content is given by: C : album norm: units 0:110 0:685 0:055 0:6852 1 1=16 0:101 11

where 0.110 and )0.055 are the regression coefcients. Also the silt content pushes the abundance of C. album below the overall mean, and Fig. 1 shows that in this eld silt is not enough to strongly increase C. album abundance. With a high silt content, the steep slope of the line Silt indeed suggests a high abundance of C. album.

Chenopodium album was the species with the highest relative density in elds A0610, A0830, A1180, A1220 of the analysis set and also characterised elds V0840, V0900 and V0940 of the validation set. In this group of elds, tillage diered widely, in agreement with the results of Trresen and Skuterud (2002) concerning the indierence of this species to changing tillage. The species seemed to be favoured by high silt and average clay contents; its abundance was always inversely proportional to the sand content. It was also favoured by low pH, in agreement with the result of Basset and Crompton (1978). The relationship with organic matter content suggests that it was favoured by quite high or quite low (extreme) organic matter contents, even if not so penalised when this content is average. It appears to be penalised by high P2O5 and MgO contents and favoured by high K2O contents.

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A weed seedbanksoil properties model 319

0.5 0.3 0.1 0.1

C. album

1.2

G. parviflora

1.0 0.6

G. ciliata

0.8

0.4

0.2 0.2 0.6 3 0.2

0.0

0.3 3

0.4 3 0.3 0.1

0.1 0.1

C. polyspermum

S. oleraceus

Amaranthus spp.

0.0 0.1

0.3 0.5 0.7 3

0.3 0.2 0.5 0.7 3 0.6 0.4 0.2 0.4 3

0.4 S. media 0.2

V. persica
0.4

C. bursa-pastoris

0.2

0.0 0.0 0.2 0.0 0.2 0.4 3 0.2 3

0.4 3

P. oleracea
0.4

0.2 S. arvensis 0.1

1.0 0.6

C. minus

0.0

0.1 0.2

0.2 0.2 0.6 1.0 3

0.4 0.3 0.8 3 0.4 3

D. sanguinalis
0.1

0.1

0.3

0.5 3

Fig. 2 Relationships between soil texture (x-axis, normalised values) and the relative density (y-axis, normalised values) of species included in the model ( sand; silt; clay).

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320 S Otto et al.

0.6

C. album

2.0 1.6

G. parviflora

1.6 G. ciliata 1.2 0.8

0.4 1.2 0.2 0.8

0.4 0.4 0.0 0.0 0.2 3 0.4 3 0.5 0.0 0.4 3

C. polyspermum
0.3

S. oleraceus
0.3

Amaranthus spp.

0.3 0.1 0.1 0.1 0.1 0.1

0.1

0.3 3 0.6

0.3 3

0.3 3 0.8

S. media

0.3 0.1

V. persica

C. bursa-pastoris

0.2

0.4

0.2

0.1 0.3 0.5 3

0.0

0.6

0.4

1.0 3 0.4

0.8 3

P. oleracea

0.8 S. arvensis 0.6

C. minus

0.0 0.4 0.2 0.4 0.0 0.8 0.2

1.2 3

0.4 3

0.6 3

D. sanguinalis
0.2

0.0

0.2

0.4 3

Fig. 3 Relationships between soil chemical properties (x-axis, normalised values) and the relative density (y-axis, normalised values) of species included in the model ( organic matter; pH; cation exchange capacity; CaCO3act).

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A weed seedbanksoil properties model 321

0.4

C. album
0.6

G. parviflora

0.6

G. ciliata

0.0 0.2

0.2

0.4

0.2

0.2

0.8 3 2 1 0 1 2 3 4

0.6 3.5 2.5

0.6 3 0.3

2.5

C. polyspermum

S. oleraceus

Amaranthus spp.

0.1

1.5 1.5 0.5 0.5 0.3 0.5 0.5 0.5 1.5 3 1.4 1.0 0.4 0.6 0.2 0.0 0.2 0.6 3 0.4 0.4 3 0.4 2 1 0 1 2 3 4 1.5 3 2 1 0 1 2 3 4 3 0.8 2 1 0 1 2 3 4 0.1

S. media
0.8

V. persica

C. bursa-pastoris

0.0

P. oleracea
0.8 0.4

S. arvensis
2.0 1.6

C. minus

0.4

0.0

1.2 0.8 0.4

0.0

0.4

0.4

0.8 0.0 0.4 3

0.8 0.8

1.2

D. sanguinalis

0.4

0.0

0.4

0.8

Fig. 4 Relationships between soil chemical properties (x-axis, normalised values) and the relative density (y-axis, normalised values) of species included in the model ( P2O5; K2O; CaO; MgO; B).

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322 S Otto et al.

Galinsoga parviora and Galinsoga ciliata

Galinsoga parviora was the species with the highest relative density in elds A0530, A1000, A1010, A1030 of the analysis set and eld V1160 of the validation set. The abundancetexture relationships found were very interesting: G. parviora abundance was favoured by a variety of conditions, e.g. with low sand or clay content or high silt content, provided that clay is scarce. Galinsoga parviora did not appear to be sensitive to pH, CaCO3act or CEC, while it seemed to be favoured when organic matter content was low, and in this it resembles C. album. Galinsoga parviora was favoured by average contents of P2O5, CaO, MgO, B and high K2O content. The wide range of distribution of G. parviora with respect to texture and pH was also reported by Columa (1983). The preference for such dierent textures can be explained by an interaction eect between texture and humidity as reported by Rai and Tripathi (1983). Looking at the relationships found in Val di Gresta it seems that G. parviora and G. ciliata were favoured by the same critical soil parameters, and that therefore the dierence in abundance of G. ciliata in comparison with G. parviora could be explained by the dierent fertility of the species under the pressure of crop management.
Chenopodium polyspermum and Sonchus oleraceus

an expression of the same aptitude to colonise similar environments. The similarity of the relationships found for these two species was notable for all the soil properties.
Amaranthus spp.

This species characterised eld A0660 of the analysis set, while it was scarce in those of the validation set. Amaranthus spp. was favoured by sand, silt, CaCO3, CaO contents that were not extreme, and low clay content. Texture conditions favourable to Amaranthus spp. were in agreement with that reported by Holm et al. (1997). A relationship with pH existed such that a very low pH was unfavourable. However, Montegut (1984) demonstrated that Amaranthus spp. was indierent to pH. Abundance of Amaranthus spp. seemed higher when organic matter or CEC were low. Amaranthus spp. was also favoured where P2O5 and K2O were average and was penalised by high B content.
Stellaria media

Chenopodium polyspermum was the species with the highest relative density in eld A1060 and characterises elds A1180 and A1200 of the analysis set, but was not the main species in any eld of the validation set. Chenopodium polyspermum was clearly favoured by average conditions of texture and its abundance decreased as conditions became extreme. Therefore, C. polyspermum seemed to have complementary behaviour to that observed for C. album. The relationships with sand and clay appeared to follow one another, and the abundance was higher when the sum of this was average, almost as if a kind of substitution eect exists between them. Complementary to C. album, the abundance of C. polyspermum was proportional to organic matter and CEC. It was more sensitive than C. album to CaCO3act content and like C. album, it was penalised by too high pH. The favourable conditions of texture, pH and CaCO3act were in agreement with those reported by Columa (1983). Chenopodium polyspermum was also favoured by average contents of P2O5, K2O, MgO and CaO; instead, there was a strong direct proportionality with B content. In Val di Gresta, a positive correlation was noticed in the elds of the analysis set between the abundance of C. polyspermum and that of S. oleraceus. It was clearly

This species was characteristic of eld A0840 of the analysis set, while it was scarce in those of the validation set. Stellaria media abundance was favoured by soil with high sand and clay contents, and low silt content, but was penalised by low organic matter, CEC and pH. Stellaria media was favoured by soils rich in P2O5, K2O and CaO, but was indifferent to the B content. The relationship with K content was dissimilar to that with Mg, indeed it was opposite. These two elements are often paired, e.g. in the expression Mg+K content, but this might not always be the case as suggested by S. media in this study. The preference of S. media for sandy-clay, not acid soils, is in agreement with Hallgren (1990), while that for high organic matter and P2O5 contents agrees with that reported by Walter et al. (2002).
Veronica persica, Veronica hederifolia, Veronica verna

The species V. persica was found in 14 elds of the analysis set, even if it was the main species in none of them; it was also found in all elds of the validation set, with a maximum relative density of 8.17%. Therefore, V. persica was a widely-distributed species, and the graph of the relation with soil properties explains why: it was indierent to sand content, the relationship with silt was the opposite to that of clay, and the three parabolas of soil texture relationships intersect in the point (0.0). This can be interpreted as meaning the species has average abundances in soils with average texture. It was generally indierent to pH and did not appear to be

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A weed seedbanksoil properties model 323

penalised by low pH values. Veronica persica was favoured by intermediate organic matter content, was relatively abundant with low P, K and CaO contents, and is considered a typical species of organic farming or ` rberi et al., 1998). of soils with a low chemical input (Ba Veronica hederifolia and Veronica verna L., two Veronica species of minor importance, were also found in Val di Gresta. Veronica hederifolia was found in seven elds, V. verna only in two. Both species were always found together with V. persica, i.e. the elds with V. verna were a subset of the elds with V. hederifolia, which were a subset of those with V. persica. Notably, the two elds where V. verna was present (A0800 and A0840) were characterised by average or low CaCO3 content. The characteristic calcifuge behaviour of V. verna is in agreement with Jauzein (1995), however a clear preference for specic soil texture was not evident. It is interesting to analyse which soil properties are changed or lost passing from the set of V. persica soils to the V. hederifolia subset. Veronica persica was found in 14 of 16 elds, so the V. persica set includes a wide range of soil properties. The narrowing of the V. hederifolia set was due to the fact that this species had a small set of preferences, e.g. V. hederifolia was not found where pH was very low. Similarly, the reduced presence of V. verna could be linked to a further narrowing of the set of preferences, and V. verna was found only where the clay, organic matter, K2O, CaO, B and CEC contents were high or very high.
Capsella bursa-pastoris

mediate and pH and CaCO3 contents must not be low. It is worth noting that the relationship with organic matter (i.e. the parabolas shape) was opposite to that found for C. album. The pH range (between 7.85 and 7.95) and the high CaCO3 content in the elds where P. oleracea was very abundant, are partially in agreement with Miyanishi and Cavers (1980).
Spergula arvensis

This species was typical of eld A1260 and was also very abundant in eld A1200 of the analysis set. However, it was virtually absent from elds of the validation set. Spergula arvensis was very abundant where the clay content was low and there was equilibrium between sand and silt. It was favoured by low but not extreme values of pH, as reported by Jauzein (1995), and average conditions of CEC. The favourable conditions of soil texture agreed with that reported by Columa (1983), and, in part, by Hallgren (1990) who suggested that the abundance of S. arvensis was proportional to the sand content. The inverse relationship with CaCO3act and B contents was of note, the latter being very marked. In Val di Gresta, S. arvensis was only found at about 1200 m a.s.l., because of the compatibility of its biological cycle with lower temperatures (Radics et al., 2000).
Chaenorrhinum minus

This species was found in all elds of the analysis set, although it was never the dominant species. It was also found in all elds of the validation set, with a maximum relative density of 4.35%. Therefore, C bursa-pastoris was a widely-distributed species, like V. persica, and indeed their relationships with soil texture and the main chemical properties were very similar, with C. bursapastoris less sensitive to organic matter content and CEC, but more sensitive to pH.
Portulaca oleracea

This species was the main species in elds A0580 and A0850 of the analysis set, whereas it was very scarce or absent in those of the validation set. Relationships between P. oleracea abundance and soil texture were very clear: this species was favoured by soil rich in sand and poor in silt and the clay content was of minor importance. It was very abundant in elds A0850 and A0580, which were very rich in sand (A0580 had the overall highest sand content) but scarce in silt (A0580 had the overall lowest silt content). The organic matter content must be inter-

This species was characteristic of eld A0800 of the analysis set, and was virtually absent from those of the validation set. Chaenorrhinum minus abundance was favoured by soil with high clay content and poor in silt, and was insensitive to the sand content. The abundance of C. minus was average when sand, silt and clay were average, and these conditions are in agreement with Columa (1983). The correspondence average abundance in average texture was also in agreement with Jauzein (1995). Soil pH appeared to be of little importance, whereas abundance was favoured by a high CEC (eld A0800 has the highest CEC). A clear direct proportionality exists with MgO content, the other elements being of little importance. In this study, C. minus did not appear to be a fertility indicator, in contrast to the results of Caputa (1984).
Digitaria sanguinalis

This species is thinly spread throughout the valley and was ranked 20 in the ora list of the analysis set in terms of relative density. However, it characterised eld V0750 of the validation set. Digitaria sanguinalis was favoured by average sand and silt contents and in this respect it

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324 S Otto et al.

was very similar to Amaranthus spp., while clay content seemed to have little importance. The abundance of this species appears to be independent of pH, but proportional to organic matter content and CEC value. It was also more abundant where MgO content was high and B content low.

Discussion
The seedbank in the 16 elds was fairly consistent, with a range of between 34 and 200 million seeds ha)1, values not unlike those found in other research on arable soils. Annual weeds predominated, in some elds reaching almost 100%. The most distinctive result was observed in the seed population structure. The seedbank was characterised by a large number of species (32.7, range 2541), in agreement with many other authors who have found that organic farming or relaxed management usually amplies populations of rare species (Hebden et al., 1998). The results indicated that the variability of the seedbank was also very high in elds with the same management and at the same altitude. This indicates that relationships between weed species density and soil properties are eld-specic, or even site-specic within a eld, because of the dierent preferences of the weed species for the various soil types (Dessaint, 2000; Walter et al., 2002). This is the case even if some weed species are sometimes ambiguous indicators, i.e. they are not only found in their preferred habitat (Jauzein, 1995). The germinable seedbank was characterised by fewer species, and in some cases by a high density of one or two, suggesting that weed ora composition can also be dramatically simplied in organic farming. A strong presence at high density and frequency levels of continuously fruiting annuals (e.g. G. ciliata, G. parviora, P. oleracea and S. media) and true monocarpic annual weeds (e.g. C. album, C. polyspermum and Amaranthus spp.) was also noted in the valley. The relative density of weeds in the germinable seedbank can be predicted using seedbank composition and soil properties. Hypothesising a quadratic relationship between weed abundance and soil properties in a PLS regression analysis, the overall predictive capacity of the resulting model was quite good and for many species very high. Part of the remaining unexplained variability was probably caused by tillage eect and sampling error, e.g. due to a validation set that does not span the range for the analysis set. Furthermore, the elementary relationships between soil properties and weed species abundance can be plotted in a simple and informative way in a two-dimensional graph, which can help in understanding the behaviour and spread of some species. The use of quadratic relationships increases the

analysis complexity, whereas a model with only rstorder terms would be simple. When the nal abundance of a species is seen as a result of the single soil property contribution, it is possible to rationalise why some species are widespread, whereas others tend to concentrate in specic elds. For example, when a species is slightly favoured by some soil properties and slightly penalised by others, it can reach high relative abundance in elds with quite dierent characteristics (i.e. widespread in various types of soils) (e.g. C. album, G. parviora and C. polyspermum). When, instead, the abundance of a species has a strong proportionality with certain soil properties, and a eld with extreme values (e.g. very high or very low) of those properties exists, then the species will have very high or very low relative abundance in that eld (e.g. S. arvensis for organic matter and boron, P. oleracea for silt). In contrast, in some other cases relative abundance could be higher when a certain soil condition value is average (e.g. P. oleracea for organic matter). In Val di Gresta, S. arvensis was found in abundance in two elds. Therefore, it appears that some arable weeds, now considered uncommon, could return to having detectable populations with appropriate management systems (Beveridge & Naylor, 1999). This in particular in soil with low clay content and equilibrium between sand and silt, sub-acid pH, and low B content. Portulaca oleracea abundance seemed to be favoured by soil rich in sand and poor in silt and with average organic matter content. The two Chenopodium species were not always abundant in the same eld; the texture preferences highlighted by the model indicate it would be unlikely to nd them both with high relative density. Soil pH was not a discriminating property, whereas C. polyspermum seems to reach higher relative abundance with increasing levels of organic matter and cation exchange capacity. Amaranthus spp. was also abundant when organic matter and cation exchange capacity were low, provided that pH was not. The soil properties can explain the presence and relative abundance of weed species in Val di Gresta. Although climatic factors generally inuence the distribution of a species on a large scale, soil factors are a major cause of local distribution patterns, even if the eect of seed dispersal pattern is taken into account. Crop management inevitably adapts environmental conditions and the ora variability on a landscape scale. However, on a smaller scale, and when the assumptions on crop management described in the Materials and methods section hold, it is still possible to identify the links between soil properties and species abundance.

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A weed seedbanksoil properties model 325

Acknowledgements
The research was nanced by the Italian CNR and in part by the INRM in the program Miglioramento delle tecniche di controllo delle malerbe in un sistema di economia corta: il caso della Val di Gresta in Trentino. The authors are particularly grateful to Michela Luise, Massimo Bonetti and Alberto Cappelletti, the Consorzio Ortofrutticolo Val di Gresta and to all the farmers they met in the valley.

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