# Mediterranean vineyard soil seed bank characterization along a slope/disturbance gradient: Opportunities for land sharing

Martin Faucher, Séraphine Grellier, Clémence Chaudron, Jean-Louis Janeau, Gabrielle Rudi, Fabrice Vinatier

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## ► To cite this version:

Martin Faucher, Séraphine Grellier, Clémence Chaudron, Jean-Louis Janeau, Gabrielle Rudi, et al.. Mediterranean vineyard soil seed bank characterization along a slope/disturbance gradient: Opportunities for land sharing. Agriculture, Ecosystems & Environment, 2024, 361, pp.108821. 10.1016/j.agee.2023.108821. hal-04300713

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1 Mediterranean vineyard soil seed bank characterization along a slope/disturbance

2 gradient: Opportunities for land sharing

3

4 Martin Faucher¹*, Séraphine Grellier², Clémence Chaudron³, Jean-Louis Janeau⁴,

5 Gabrielle Rudi¹,⁵, Fabrice Vinatier¹

6

7 ¹ LISAH, Univ. Montpellier, AgroParisTech, INRAE, Institut Agro, IRD, Montpellier,

8 France

9 ² Department of Spatial planning and Environment Engineering of Polytech Tours,

10 CITERES UMR7324 CNRS, University of Tours, Tours, France

11 ³ UMR 1434 SILVA, Université de Lorraine, AgroParisTech, INRAE, Faculté des

12 Sciences et Technologies, F-54506 Vandœuvre-lès-Nancy, France

13 ⁴ iEES-Paris, Institut d'Ecologie et des Sciences de l'Environnement de Paris, (IRD,

14 Sorbonne Université, CNRS, INRAE, UPEC, Université Paris Diderot), Paris, France

15 ⁵ G-EAU, Univ. Montpellier, AgroParisTech, BRGM, CIRAD, INRAE, Institut Agro,

16 IRD, Montpellier, France

17

18 *Corresponding author: Martin Faucher, martin.faucher@inrae.fr

19 Abstract

20 The Mediterranean region is predicted to experience more intense rainfall events

21 separated by severe droughts due to climate change. In agrosystems, the

22 intensification of rainfall on dry bare soils will lead to an increase in runoff and

23 erosion rates, especially on slopes. A common method to reduce erosion is to cover

24 the soil with vegetation. To save costs, this vegetation could be provided by the soil

25 seed bank. Vineyards situated on slopes are part of the typical Mediterranean

26 landscape; thus, developing vegetation cover inside vineyards could be a solution to

27 fight growing erosion risks in addition to providing other valuable services. However,

28 the relationship between slope, erosion and seed loss can vary. Another key

29 ingredient in the development of spontaneous vegetation cover is the spatial

30 dispersal of seeds from one place to another, but the amount of seeds displaced

31 depends on their respective dispersal modes. In this study, we sampled the soil seed

32 bank at two sites characterized by either a low or high frequency of vegetation

33 removal. The soil seed bank was sampled along a transect starting upslope of the

34 vineyards to the bottom of neighboring agricultural ditches. Germination trials were

35 carried out to define the soil seed bank in terms of species and seed density. Each

36 identified species was linked with a main dispersal mode. Our main objective was to

37 assess the effect of vegetation cover on seed loss inside a vineyard, as well as the

38 potential differences in the soil seed bank along our transect and the influence of

39 dispersal mode on those differences. Despite previous contradictory literature,

40 vegetation did not seem to protect interrows from seed loss caused by runoff.

41 Overall, our results support the use of the soil seed bank to provide spontaneous

42 vegetation cover to limit present and future erosion rates and advocate for relaxing
43 vegetation removal operations.

44

45 Keywords: soil seed bank, germination method, dispersal, agrosystem

46

# 1. Introduction

47 In the Mediterranean area, rainfall events are predicted to become more intense and 48 droughts more severe due to climate change (Cozannet et al., 2022). In 49 agrosystems, rainfall events will increase soil erosion caused by runoff, a 50 phenomenon already considered problematic for more than three decades (Busico et 51 al., 2023). Soil erosion causes multiple problems leading to a decrease in agricultural 52 productivity (Pimentel & Burgess, 2013). A common practice to reduce erosion is to 53 protect the soil with vegetation cover (Wu et al., 2020; Baraibar et al., 2021).

54 The presence of vegetation inside agrosystems would have additional 55 benefits alongside erosion reduction. This idea is at the core of the concept of land 56 sharing, which advocates for maintaining and/or increasing the surface of 57 noncultivated zones found within agrosystems (i.e., hedgerows, field edges, ditches, 58 etc.) (Green et al., 2005; Fischer et al., 2014). To be effective, land sharing requires 59 a sufficient pool of plant biodiversity. This pool can originate from the biodiversity in 60 adjacent habitats or directly from the soil seed bank (Erfanzadeh et al., 2010); thus, 61 seeds can disperse through space or time (Plue and Cousins, 2018).

62 Using the soil seed bank to produce vegetation cover could be considered, 63 but erosion has been associated with seed loss inside the field (Han et al., 2011; 64 Janeau et al., 2022); thus, the potential of spontaneous vegetation cover could be 65 threatened by the process it aims to reduce. Moreover, while the link between 66 erosion and seed loss has been observed to be direct (García-Fayos, Bochet and 67 Cerdà, 2010; Bochet, 2015), seed loss can also occur without erosion when light 68 seeds are removed by runoff water whose force is otherwise too weak to displace 69 soil particles (de Rouw et al., 2018); thus, vegetation cover may reduce erosion but 70 not seed loss. To summarize, runoff can potentially reduce the potential of the soil 71 seed bank to produce sufficient vegetation cover regardless of the surface state.

72 Most studies of the soil seed bank in agrosystems revolve around avoiding 73 seeds from the soil seed bank from germinating to limit weeds in agrosystems (Mahé 74 et al., 2021; Zhang et al., 2021; Adeux et al., 2023). There have been comparisons 75 of the plant diversity between the field and its adjacent zones but those studies 76 mostly investigated the aboveground vegetation (Fried et al., 2012; Minarro, 2012; 77 Barrio et al., 2013; Holland et al., 2017; Cursach et al., 2020), and those that 78 investigated the soil seed bank did not sample outside the cultivated surface 79 (Roschewitz et al., 2005; Fracchiolla et al., 2016; Carpio et al., 2020). Given the lack 80 of linkability between the seed bank and the aboveground vegetation (Andreasen et 81 al 2018), we need to confirm that the difference in diversity between the plot and its 82 surroundings is also found in the seed bank.

83 Together with the field edges, the soil seed bank of agricultural ditches is also 84 not well known. While plant communities inside ditches have been studied
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(Blomqvist et al., 2006), recent observations of the soil seed bank are lacking.

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However, ditches are interesting from a restoration perspective because they act as

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ecological corridors (Herzon and Helenius, 2008; Dollinger et al., 2015); thus, they

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can accelerate spatial dispersal and improve soil seed bank diversity.

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Most spatial dispersal modes can be grouped into four main vectors: wind

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(anemochory), water (hydrochory), animals (zoochory), and gravity (barochory).

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Usually, spatially dispersed seeds become a part of the soil seed bank (Dekker,

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1999), but the relative contribution of the different dispersal syndromes to soil seed

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bank replenishment is still unclear (Benvenuti, 2007). Therefore, studying the soil

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seed bank could also provide information on the potential plant community and the

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impact of different dispersal modes on soil seed bank replenishment.

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To evaluate the potential of vegetation to reduce seed loss and the effect of

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practices and dispersal modes on the soil seed bank, we set up an experiment to

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determine the soil seed bank distribution within two Mediterranean vineyards and

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their adjacent environments, namely, their respective field edges and bordering

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ditches. The soil seed bank was then described in terms of seed density and species

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diversity at each sampled point. We formulated three main hypotheses: (1) The

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effect of the slope on the soil seed bank will be different between tilled and vegetated

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interrows. We also expect (2) operations within the field to deplete the soil seed

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bank; thus, the soil seed bank will be denser and more diverse in the field edges and

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the ditches than within the field. Grouping species into anemochory, hydrochory,

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barochory, and zoochory, we expect that (3) the effect of slope or position in the

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study site on the soil seed bank will be different depending on the dispersal mode.

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## 2. Materials and methods

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- 2.1. *Study sites*

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Two study sites in the Hérault Department (France) were selected for the experiment

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(Fig. 1).

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<figure><img src="image_3.png" /></figure>

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<figure><img src="image_4.png" /><figcaption>Figure 1. Location of study sites and visualization of interrows in the Roujan study site (A) and the tilled (B) and vegetated (C) interrows of the Combaillaux study site</figcaption></figure>

The first study site, herein designated as the P1, is in the Roujan catchment (43°30′N, 3°19′E), situated west of Montpellier, France. The Roujan catchment has been monitored since 1992 as a part of the long-term observatory ORE-OMERE (Molénat et al., 2018). The Roujan catchment area is a small catchment area (<1 km²) mainly occupied by vineyards framed by a network of drainage ditches dug by farmers to drain some of the water from the soil, collect runoff and limit erosion. The catchment is characterized by a sub-Mediterranean subhumid climate with a long dry period, and the majority of rainfall occurs in spring and autumn. The average rainfall is 600 mm.year⁻¹ (Molénat et al., 2018). The vineyards that constitute most of the Roujan catchment area belong to different producers. The maintenance of the drainage ditches is the responsibility of the owners of the vineyards they cross and typically consists of removing the vegetation by mowing or burning the vegetation to preserve the water flow capacity. More rarely, dredging/chemical weeding operations are carried out. Within the Roujan catchment, P1 is a vineyard of approximately 0.3 ha of *Pinot Noir* grape variety planted on Calcaric Regosols with a mean slope of approximately 15%. Rows are oriented from east to west. The vineyard is bordered by other vineyards and a drainage ditch located downstream of the most sloping part of the vineyard. All interrows of P1 are chemically weeded, and the calendar of practices has not changed in the last 5 years before the sampling took place. When sampling took place, P1 was dominated by *Poaceae* and *Brassicaceae* in the field and the field edge, and by *Rubus fruticosus* inside the ditch.

The second study site, referred to as the P2, is in a vineyard situated north of Montpellier in Combaillaux (UTM 43.659903, 3.767451, first sampled interrow). The climate there is also classified as Mediterranean and subhumid, with an average rainfall of approximately 760 mm.year⁻¹. The vineyard is HVE-certified (Haute Valeur
Environnementale - High Environmental Value), which requires a low use of synthetic chemical inputs, a significant share of vegetated interrows, and a share of diversified noncultivated zones situated above a threshold value. P2 has a surface area of approximately 3 ha of Merlot grape variety, clay-limestone soil, and a mean slope of approximately 20%. Similar to the P1, rows are oriented from east to west. An agricultural ditch downstream of the vineyard collects run-off water. This ditch separates the vineyard from another field left fallow for part of the year and sowed the remaining time. In this adjacent field, faba beans were sown during soil seed bank sampling. Every second row is mown and plowed after the harvest and then sown, and the remaining rows are simply mown. When sampling occured, P2 was dominated by a mix of Fabaceae and Poaceae, and the ditch was mostly composed of Rubus fruticosus and Clematis vitalba. After sampling in April 2022, a Trifolium mixture was sown, and the faba bean seedlings produced by the adjoining land were sown in September 2022.

As the soil seed bank could be influenced by its surrounding areas, we estimated landscape diversity inside a 500-m radius buffer around the study site following the methodology from Carpio et al. (2020). We classified orthoimages from 2021 (source: IGN) in vineyards, other agricultural land and seminatural habitats to obtain the surface proportion of each land cover. We digitized surrounding fields mainly composed of agricultural ditches to obtain a cumulative linear length inside the radius. Consequently, P1 is primarily composed of other vineyards (approximately 70%) and other agricultural lands (approximately 20%), with a small amount of seminatural habitats (approximately 10%). The buffer encompasses more than 8 km of agricultural ditches. P2 is composed of vineyards (approximately 55%) and seminatural habitats (approximately 40%). The buffer encompasses approximately 3 km of agricultural ditches.

Soil analyses were conducted at both study sites on soil samples taken alongside the soil seed bank samples. Samples were analyzed for pH, total carbon and nitrogen, organic matter content, cation exchange capacity, and concentrations of P, Mg and Na. Data were ultimately removed from the analysis due to a lack of significant results (Appendix A).

## 2.2. Soil seed bank sampling protocol

Soil seed bank sampling was carried out along a 50-meter transect consisting of 8 points starting upslope within the field until the ditch at the bottom of the study site. The first four sampling points are situated within the field, with T1 at the top of the slope and T4 at the bottom of the slope. Sampling was carried out using a corer that allowed the sampling of 10 cm diameter cylinders at a depth of 10 cm. The number of interrows sampled was determined so we had at least 3 interrows with the same vegetation state. For each point, 4 samples were collected. In the P1, 3 interrows were sampled with a spacing of one interrow between each of them (Fig. 2). The first row of the vineyard was not sampled to avoid edge effects. In total, the
189 sampling area covered approximately 350 m²

<figure><img src="image_5.png" /><figcaption>Figure 2. Representation of the Roujan vineyard (P1) and sample distribution. Each sample point represents 4 samples taken on a straight line approximately 20 cm from each other.</figcaption></figure>

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In the P2, 6 interrows were sampled, one with permanent plant cover and the other plowed after harvest (around mid-September) and then sown in April 2022. The sampling area covered approximately 2000 m² (Fig. 3).

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In total, 286 soil samples were collected for the soil seed bank of the two study sites, representing 9 interrows, 8 sampling points and 4 replicates at each sampling point, except one in the P1, where we were only able to collect 2 soil samples because the soil layer was too thin to collect the other two samples.

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<figure><img src="image_6.png" /><figcaption>Figure 3. Representation of the Combaillaux vineyard (P2) sampling distribution and detailed sampling pattern. Each sample point represents 4 samples taken on a straight line approximately 20 cm from each other.</figcaption></figure>

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## 2.3. Seedling emergence method

Three days after the collection of the soil samples in the field (in April 2022), the 286 soil samples were placed in perforated plastic boxes (200x170x75 mm) on a layer of draining sand and covered with potting soil to facilitate germination. Before use, the potting soil was sterilized to ensure that it did not contain any organisms that could influence the development of the seeds (granivores, fungi, insects, etc.). The sterilization process included successive oven drying and then freezing the soil. The boxes were then placed randomly in an air-conditioned greenhouse (average T° = 22 +/- 3°C) on 3 separate benches. To ensure that the samples were not contaminated by exogenic seed input or seeds from other samples, one control box containing only sand and potting soil was placed on each bench. The samples were manually irrigated 5 days a week with tap water to maintain a constant moisture content throughout the experiment.

To trigger the emergence of the seeds of certain species from dormancy, a cold stratification period at 1°C was carried out for one month. The boxes were then placed back in the greenhouse in a randomized arrangement different from that prior to vernalization. The samples were also shuffled to stimulate germination (Price et al., 2010; Erfanzadeh et al., 2016).

Samples were monitored regularly to count and remove individuals sufficiently developed to be identifiable at the species level if possible and at the genus level otherwise using the Latin nomenclature as available in the WFO Plant List classification (https://wfoplantlist.org/plant-list/). Poaceae were regularly replanted in individual pots to achieve a more precise identification without impacting the remaining seeds that may still germinate within the samples.

The spontaneous germination experiment took place over nine months. It was divided into two cycles: from April 2022 to July 2022 and from September 2022 to December 2022. The end of the experiment occurred after 2 weeks with the absence of seed emergence. Residual soil was not combed because the added information has been considered negligible in previous studies (Niknam et al., 2018).

## 2.4. Soil seed bank sampling representativity

To gain insight into how many species we might have missed, we used calculations developed for plant species composition surveys. This method is based on computing the theoretical species accumulation curve (SAC) by calculating the sum of generalized Simpson entropies based on subsampling using jackknife methods (Zhang and Zhou, 2010, summarized in Marcon, 2015). This method forecasts the effort needed to reach the “true” number of species in the sampled population. According to this method, we could have missed approximately 10 species in our sampled soil seed bank, which means that our results might have a bias of approximately 10%.

## 2.5. Dispersal modes
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## 2.6. Statistical analyses

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Species observed throughout the emergence period have been linked to their main dispersal mode using the TRY database (Kattge et al., 2020). Most species had several dispersal modes; thus, the most observed mode was selected. Among the 96 different species identified, 16 were primarily dispersed by anemochory, 11 by barochory, 15 by hydrochory and 54 by zoochory.

Eight different equations were used for generalized linear model (GLM) construction (Tab. 1), with separate GLMs constructed for each study site. All data manipulation and statistical analyses were performed using R 4.2.2 (R Core Team, 2023) software. The Shannon index was computed for every sample using the entropart package (Marcon and Hérault, 2015) with a Chao, Wang & Jost correction (Chao, Wang and Jost, 2013) to take undetected species into account. The number of germinated seeds observed per box (=E) was related to the area sampled (=S, equal to 0.008 m²). Thus, the number of germinated seeds per surface unit obtained for each box, referred to as the "E/S ratio" (in number of plants per m²), was used for statistical analysis (Gaudichet et al., 2022). Vegetation cover was visually inspected at multiple points in every interrow sampled. In the Combaillaux vineyard, vegetation cover was on average between 90 and 100% inside the vegetated interrows and under 5% inside the unvegetated interrows. In the Roujan vineyard, vegetation cover was on average under 5% in all the studied interrows. Consequently, the analyses that included the effect of vegetation cover were only conducted on data collected within the field in the P2, and vegetation cover was included as presence/absence data in the models. Concerning the effect of dispersal mode on soil seed bank distribution, the number of germinated seeds observed for each main dispersal mode dispersal mode defined in the previous section was weighted by the number of species with this dispersal mode to avoid having an inflated E/S ratio caused by an overrepresentation of one particular dispersal mode.

The effect of the slope on the soil seed bank was tested using samples taken from the first 4 sampling points, which were used as a proxy for the slope gradient within the field. The sampled transect points were grouped into three zones: points T1 to T4 in the field (field zone), points T5 to T7 in the field edge (field edge zone), and points taken in T8 (ditch zone).

The effect of landscape was not analyzed since the information would have been identical for every sample in each study site.

Because the number of germinated seeds was not normally distributed, our hypotheses were tested by constructing GLMs with a Poisson distribution, which eliminates the requirement for linearity and homoscedasticity of the data. Overdispersion of the residuals was detected; thus, a correction was applied using a quasi-GLM (Zuur et al., 2009). Based on a visual inspection, normality of the residuals and homogeneity of variances were met. Models were then tested using
296 ANOVA tests.

297 To confirm that greenhouse position did not impact emergence, a GLM linking
298 the E/S ratio to the sample location within the greenhouse was also constructed and
299 tested with ANOVA. Greenhouse position did not have an impact on the number of
300 seeds that germinated (F = 0.44, p = 0.66).

301

302 Table 1. Variables used for generalized linear model (GLM) construction. Separate GLMs were constructed for

303 the two study sites, except for Equations 5 and 6, including the effect of the vegetation cover, which were only

304 constructed using data from inside the field of the Combaillaux study site. Vegetation cover was included as

305 presence/absence data in the models

<table><thead><tr><th>Eq</th><th>Y</th><th>Explanatory variable(s)</th></tr></thead><tbody><tr><td>1</td><td>E/S ratio</td><td>slope</td></tr><tr><td>2</td><td>Shannon index</td><td>slope</td></tr><tr><td>3</td><td>E/S ratio</td><td>zone</td></tr><tr><td>4</td><td>Shannon index</td><td>zone</td></tr><tr><td>5</td><td>E/S ratio</td><td>Interaction between zone and vegetation cover</td></tr><tr><td>6</td><td>Shannon index</td><td>Interaction between zone and vegetation cover</td></tr><tr><td>7</td><td>E/S ratio</td><td>Interaction between zone and dispersal mode</td></tr><tr><td>8</td><td>Number of species over the total number of species for the dispersal mode</td><td>zone</td></tr></tbody></table>

306

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308 Differences between the coefficients computed for each GLM were tested by

309 performing post hoc Tukey tests with the emmeans package (Lenth et al., 2023).

310

## 3. Results

311 A total of 96 species were identified during our experiment, 52 of which were found

312 in both study sites (Tab. 2).

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314 In the P2, the E/S ratio in T1 (upslope) was significantly higher (6160 +/- 6203

315 seedlings/m²) than the E/S ratio in T2 (steepest part of the slope) (2900 +/- 1969

316 seedlings/m²). However, we could not show such an effect of slope in P1 (Fig. 4).
<figure><img src="image_7.png" /><figcaption>Figure 4. Distribution of the E/S ratio (in log scale) by sampling point within the field for P1 (Roujan vineyard) and P2 (Combaillaux vineyard). The slope effect was tested by ANOVAs on two separate GLMs constructed with data collected in P1 and the P2. T1 is situated before the start of the slope, T2 is at the start of the slope, T3 is at the end of the slope and T4 is at the bottom of the slope. In the plot displaying data from the P2, different letters indicate a significant difference between GLM coefficients.</figcaption></figure>

<table><caption>Table 2. Number of species, first five most observed species and their respective contribution in samples found in each study site, in both study sites, and overall</caption><thead><tr><th></th><th>P1<br>Roujan vineyard</th><th>P2<br>Combaillaux vineyard</th><th>Shared</th><th>Total</th></tr></thead><tbody><tr><td>Number of species observed</td><td>68</td><td>81</td><td>52</td><td>96</td></tr><tr><td>Most observed species</td><td><i>Diplotaxis erucoides</i><br>(10.6%)</td><td><i>Setaria pumila</i><br>(13.3%)</td><td><i>Setaria pumila</i> (12.1%)</td><td><i>Setaria pumila</i><br>(11.1%)</td></tr><tr><td>2nd most observed species</td><td><i>Euphorbia prostrata</i><br>(9.3%)</td><td><i>Portulaca oleracea</i><br>(12%)</td><td><i>Euphorbia prostrata</i><br>(11.6%)</td><td><i>Euphorbia prostrata</i><br>(10.7%)</td></tr><tr><td>3rd most observed species</td><td><i>Poa annua</i> (9%)</td><td><i>Chenopodium album</i><br>(11.5%)</td><td><i>Portulaca oleracea</i><br>(10.3%)</td><td><i>Portulaca oleracea</i><br>(9.5%)</td></tr><tr><td>4th most observed species</td><td><i>Erigeron sumatrensis</i><br>(7%)</td><td><i>Euphorbia prostrata</i><br>(11.2%)</td><td><i>Chenopodium album</i><br>(10.1%)</td><td><i>Chenopodium album</i><br>(9.3%)</td></tr></tbody></table>
<table><tr><td>5th most observed species</td><td>Lysimachia foemina (6%)</td><td>Polygonum aviculare (8.5%)</td><td>Polygonum aviculare (7.2%)</td><td>Polygonum aviculare (6.6%)</td></tr></table>

## 3.1. In-field effect of vegetation cover

The mean E/S ratio was significantly higher in samples originating from vegetated interrows (5329 +/- 4419 seedlings/m²) than in samples originating from tilled interrows (3773 +/- 3109 seedlings/m²; t = 1.99, p<0.05).

The mean Shannon index was also significantly higher in samples originating from vegetated interrows (2.42 +/- 0.48) than in samples originating from tilled interrows (1.99 +/- 0.41; t = -4.61, p<0.01).

We could not show a significant interaction of slope and vegetation cover on the E/S ratio (F = 1.9, p = 0.12).

However, the interaction of slope and vegetation cover had a significant positive effect on the Shannon index (Fig. 5). The mean Shannon index from T3 (slope end) was significantly lower (1.95 +/- 0.43) in tilled interrows than in vegetated interrows (2.67 +/- 0.46; p < 0.01).

<figure><img src="image_8.png" /></figure>

<caption>Figure 5. Shannon index computed for each sample grouped by slope point sampled for P2 (Combaillaux vineyard) only in tilled and vegetated interrows. The combination of the slope point and the presence of vegetation on the Shannon index was tested with ANOVA. T1 is situated before the start of the slope, T2 is at the start of the slope, T3 is at the end of the slope and T4 is at the bottom of the slope. The difference between all coefficients was tested with Tukey tests between all slope points, and different letters are used when differences are significant.</caption>
350 3.2. Zone effect

351 - 3.2.1. Effect of zone on the E/S ratio

352 Zone (i.e., the sample origin between the field, field edge and ditch) had a significant 353 effect on the E/S ratio at both study sites. In the P1, the mean E/S ratio was lower 354 within (1426 +/- 803 seedlings/m²) than outside the field (field edge: 3860 +/- 3953 355 seedlings/m²; ditch: 3144 +/- 1524 seedlings/m²). In the P2, the mean E/S ratio was 356 lower at the field edge (3839 +/- 2760 seedlings/m²) than in the ditch (5759 +/- 2755 357 seedlings/m²; Fig. 6).

<figure><img src="image_9.png" /></figure>

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Figure 6. Distribution of the E/S ratio (in log scale for readability purposes) in each sample, grouped by location in the study site, for P1 (Roujan vineyard) and P2 (Combaillaux vineyard) separately. The "field" location represents samples taken within vineyard rows, the "field edge" location represents samples taken between the rows and the ditch, and the "ditch" location represents samples taken within agricultural ditches. The location effect was tested with ANOVAs, the difference between coefficients was tested with Tukey tests, and different letters are used when differences are significant.

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366 - 3.2.2. Effect of zone on the Shannon index

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368 Zone had a significant effect on the Shannon diversity index at both sites (Fig. 7).

369 In the P1, the mean Shannon index was only significantly higher in the ditch (2.74 +/- 0.65) than in the field (2.13 +/- 0.64, p < 0.05), whereas in the P2, both the field edge (2.48 +/- 0.49, p < 0.01) and the ditch (3.49 +/- 0.49, p < 0.01) had a higher mean 370 Shannon index than the field (2.20 +/- 0.49). The mean Shannon index was also 371 significantly higher in the ditch than in the field edge (Fig. 7).

372
<figure><img src="image_10.png" /><figcaption>Figure 7. Shannon index computed for each sample grouped by location sampled for P1 (Roujan vineyard) and P2 (Combaillaux vineyard). The "field" location represents samples taken within vineyard rows, the "field edge" location represents samples taken between the rows and the agricultural ditch, and the "ditch" location represents samples taken within the agricultural ditches. The location effect was tested with ANOVAs. Differences between coefficients were tested with Tukey tests, and different letters are used when differences are significant.</figcaption></figure>

## 3.3. Effect of dispersal mode on the soil seed bank

### 3.4.1. Effect of dispersal modes on the E/S ratio

The dispersal mode affected the overall E/S ratio observed at both study sites (Fig. 8). At both sites, barochory was more prevalent than other dispersal modes (Fig. 8). Additionally, in the P2, hydrochory was more prevalent than anemochory and zoochory (Fig. 8; Appendix B).

The dispersal mode had a significant effect on the E/S ratio observed in each zone for both study sites (Fig. 9; Appendix B).

In the P1, barochory was more prevalent than other dispersal modes at the field edge and in the ditch. In the P2, anemochory and zoochory were rarer than barochory and hydrochory in the field. Zoochory was also rarer than barochory at the field edge, and barochory was more frequent than hydrochory (Fig. 9; Appendix B).
<figure><img src="image_11.png" /><figcaption>Figure 8. Share of emergences for each dispersal mode for P1 (Roujan vineyard) and P2 (Combaillaux vineyard) separately. Significant differences between dispersal modes using Tukey tests are represented using stars, one for each dispersal mode significantly different from the others.</figcaption></figure>

<figure><img src="image_12.png" /><figcaption>Figure 9. Mean E/S ratio by species for each dispersal mode, grouped by zone sampled, for P1 (Roujan vineyard) and P2 (Combaillaux vineyard). The “field” location represents samples taken within vineyard rows, the “field edge” location represents samples taken between the rows and the agricultural ditch, and the “ditch” location represents samples taken within the agricultural ditches. Significant differences between dispersal modes using Tukey tests are represented using stars, one for each dispersal mode significantly different from the others.</figcaption></figure>
405

### 3.4.2. Influence of dispersal modes on the number of species

406

407

408

409

410

411

The dispersal mode had a significant effect on the number of species found in each zone at both study sites. More anemochorous species were observed outside the field than within both study sites. Additionally, the maximum number of anemochorous species was found within the ditch in both study sites. Last, more barochorous species were found in the field edge than in the field for both study sites (Fig. 10; Appendix B).

412

413

In the P1, more barochorous species were found in the ditch than in the field.

414

415

416

417

In the P2, the ditch hosted more hydrochorous species than the field and the field edge. Similarly, fewer zoochorous species were observed in the field than in the field edge and the ditch, and there were also fewer zoochorous species in the field edge than in the ditch (Fig. 10; Appendix B).

<figure><img src="image_13.png" /></figure>

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Figure 10. Contribution of each zone to the total number of species, grouped by dispersal syndrome, for P1 (Roujan vineyard) and P2 (Combaillaux vineyard) separately. Significant differences between zones using Tukey tests are represented using stars, one for each zone significantly different from another.

422

# 4. Discussion

423

424

## 4.1. Effect of vegetation cover in interrows on seed loss reduction

425

426

427

428

Our first hypothesis that the effect of the slope on soil seed bank density would be different between tilled and vegetated interrows was not supported. This result coincides with other works observing an absence of a link between seed loss and
erosion (de Rouw et al., 2018). In their study, de Rouw et al. (2018) explained the
possibility for seeds to be dispersed without the occurrence of soil erosion by the
lower discharge necessary to move small and light seeds compared to the threshold
value over which erosion starts, which might be the explanation for our results.

Nevertheless, the overall higher number of seeds that germinated in
vegetated interrows and the higher species diversity gives credit to the expected
benefits of vegetation cover on sloping conditions, which are more diverse than
erosion control. While the benefits of vegetation cover on the soil seed bank of
woody crops have already been observed in different orchards (Carpio et al., 2020),
the spatial configuration of a vineyard and the slope are differences important
enough to consider these results noteworthy.

## 4.2. Soil seed bank diversity along the transect

441

The soil seed bank was found to be denser within the ditch than in the field in Roujan
but not in Combaillaux. Our second hypothesis that the soil seed bank would be
denser and more diverse inside the field margins and ditches than within the field
was thus partially validated. Since chemical weeding inside the field is prevalent in
Roujan and nonexistent in Combaillaux, it could be the factor that explains this
difference between sites. Indeed, chemical weeding has been linked with soil seed
bank depletion in previous studies (Andreasen et al., 2018).

The higher species diversity in the ditch found in both study sites is in
accordance with the literature obtained on aboveground vegetation studies (Fried et
al., 2012; Minarro, 2012; Holland et al., 2017; Cursach et al., 2020), which is
interesting because the soil seed bank and the aboveground vegetation are not
always similar in agrosystems (Andreasen et al., 2018). The results in the
Combaillaux vineyard also indicate that interrows benefit from the field edge and the
ditch as potential biodiversity sources, while the Roujan vineyard benefits only from
the ditch. Thus, reducing chemical inputs benefits both the field and the field edges,
which is in accordance with the expectations of land sharing (Fischer et al., 2014).

Like landscape in general, other factors, such as altitudinal gradient
(Erfanzadeh et al., 2013), need to be accounted for when considering recolonization
processes that stem from the soil seed bank of neighboring habitats.

Our results also need to be nuanced due to differences between our two
study sites in both local conditions: annual precipitation and soil conditions, and the
surrounding environment in terms of land uses and cumulative length of ditches. The
higher length of ditches in Roujan than in Combaillaux could explain the higher
diversity in the ditches, whereas the higher amount of seminatural habitats in
Combaillaux could counterbalance the difference between sites. These doubts could
be dispelled by implementing a new protocol based on more sites covering the
region's soil and climate diversity to study the interparcel variability of the seed bank.
470

## 4.3. Dispersal mode and soil seed bank distribution

471

472

473

474

475

476

477

478

479

480

481

482

483

At both sites, dispersal mode played a role in soil seed bank distribution. Our results show that barochory is well represented within vineyards and their surrounding habitats, as was already reviewed by Benvenuti (2007) for agricultural environments in general. Barochory was prevalent outside the field both in terms of seed density and species in Roujan. However, the results were mixed in Combaillaux, where barochory was mostly observed within the field in terms of seed density but outside the field in terms of the number of species. Because the mean slope is 15% in Roujan and 20% in Combaillaux, the beneficial effect of slope on dispersal by barochory is not straightforward in our results. In Combaillaux, it is possible that practices favored a few opportunistic species within the field, whereas conditions outside the field allowed more species with different dispersal modes to develop.

484

485

486

487

488

489

490

491

492

Concerning the distribution of hydrochorous species, we only observed more species in the ditch for the Combaillaux site and found no differences in terms of the number of seed counts. The absence of a higher number of hydrochorous species within the ditch in Roujan is likely due to the site position on the upper side of the Roujan catchment, which makes it a likelier site for seed detachment than retention due to its hydrological regime and its slope (Calçada et al., 2015; Fraaje et al., 2017; West et al., 2020; Su et al., 2022). An additional explanation could be the length of the ditch network, which is almost 3 times longer in Roujan than in Combaillaux; thus, seeds have more space to disperse.

493

494

495

496

497

498

499

500

501

502

As agricultural ditches can act as biodiversity corridors and refuges (Herzon and Helenius, 2008; Dollinger et al., 2015), it is not surprising to observe more zoochorous species within the ditch than within the field, although this result was observed only in Combaillaux. In Combaillaux, presence of carabids, small mammals and ants was observed, whereas in Roujan only ants were observed. This difference could be explained by the landscape, especially by the share of seminatural habitats, which is higher in Combaillaux (55%) than in Roujan (10%), because seminatural habitats are important biodiversity refuges (Garibaldi et al., 2021; Tscharntke et al., 2021).

503

# Conclusion

504

505

506

507

508

509

510

511

To our knowledge, this study was one of the first to study the effect of management practices on the soil seed bank from inside a field to the bottom of an agricultural ditch. Another noteworthy result of this study is the observation that vegetation inside the interrows did not significantly reduce the depleting effect of the slope on the soil seed bank distribution, despite the well-known role of erosion reduction on vegetation and seed loss. These preliminary results, should they be confirmed by further studies, could influence weeding practices in vineyards in order
to maximize potential benefits from a rich soil seed bank inside agroecosystems.. To confirm the related hypothesis we made to explain some of our results, further work investigating the influence of landscape on the soil seed bank is needed.

515

516

517

518

Acknowledgments

520

The authors wish to thank the two anonymous reviewers for their helpful comments, which helped to highlight the strengths of our study. The authors warmly thank Pauline MARTIN and Léo BOULAIS for their high involvement in the seedling emergence experiment. The authors are also very grateful to Léa GENTY, David FAGES, Manon LAGACHERIE, Sandrine NEGRO, and Sébastien TROIANO for their technical support in the experiment and collection of the data. The authors also wish to thank the Environmental Research Observatory OMERE (http://wwwobs-omereorg), whose policy drives the availability of part of the data used in the current study. Last, the authors wish to thank the vinegrowers for giving them access to their vineyards.

530

The PhD of M.F. was funded through LabEX AGRO 2011-LABX-002 (under the I-Site Muse framework) coordinated by Agropolis Fondation. The experimental part of the study was funded by both LabEX AGRO 2011-LABX-002 and INRAE Metaprogram Biosefair (AMI-2021). For the purpose of Open Access, a CC-BY public copyright license has been applied by the authors to the present document and will be applied to all subsequent versions up to the Author Accepted Manuscript arising from this submission.

537

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708
<table><thead><tr><th>Field</th><th>Interrow</th><th>Transect</th><th>PH</th><th>CARBONE (g/kg)</th><th>SOM (g/kg)</th><th>N (g/kg)</th><th>C/N</th><th>PHOSPHATE</th><th>MAGNESIUM</th><th>MgO (mg/kg)</th><th>POTASSIUM</th></tr></thead><tbody><tr><td>P1</td><td>1</td><td>T1</td><td>8.3</td><td>11.6</td><td>19.95</td><td>1.3</td><td>8.92</td><td>155</td><td>7.7</td><td>155.16</td><td>3.72</td></tr><tr><td>P1</td><td>1</td><td>T2</td><td>8.5</td><td>9</td><td>15.48</td><td>0.69</td><td>13.04</td><td>93</td><td>6.53</td><td>131.58</td><td>2.77</td></tr><tr><td>P1</td><td>1</td><td>T3</td><td>8.2</td><td>17.4</td><td>29.93</td><td>1.31</td><td>13.28</td><td>163</td><td>11.64</td><td>234.55</td><td>9.73</td></tr><tr><td>P1</td><td>1</td><td>T4</td><td>8.6</td><td>11.3</td><td>19.44</td><td>0.92</td><td>12.28</td><td>131</td><td>9.15</td><td>184.37</td><td>5.79</td></tr><tr><td>P1</td><td>1</td><td>T5</td><td>8.3</td><td>15.3</td><td>26.32</td><td>1.29</td><td>11.86</td><td>114</td><td>10.84</td><td>218.43</td><td>6.29</td></tr><tr><td>P1</td><td>1</td><td>T6</td><td>8.3</td><td>17.7</td><td>30.44</td><td>1.54</td><td>11.49</td><td>136</td><td>11</td><td>221.65</td><td>6.21</td></tr><tr><td>P1</td><td>3</td><td>T1</td><td>8.5</td><td>10.2</td><td>17.54</td><td>0.82</td><td>12.44</td><td>89</td><td>09.09</td><td>183.16</td><td>3.16</td></tr><tr><td>P1</td><td>3</td><td>T2</td><td>8.5</td><td>10.2</td><td>17.54</td><td>0.8</td><td>12.75</td><td>85</td><td>7.58</td><td>152.74</td><td>4.23</td></tr><tr><td>P1</td><td>3</td><td>T3</td><td>8.6</td><td>9.3</td><td>16</td><td>0.9</td><td>10.33</td><td>84</td><td>7.25</td><td>146.09</td><td>3</td></tr><tr><td>P1</td><td>3</td><td>T4</td><td>8.5</td><td>15.5</td><td>26.66</td><td>1.4</td><td>11.07</td><td>276</td><td>10.09</td><td>203.31</td><td>7.15</td></tr><tr><td>P1</td><td>3</td><td>T5</td><td>8.5</td><td>16</td><td>27.52</td><td>1.38</td><td>11.59</td><td>104</td><td>12.19</td><td>245.63</td><td>5.9</td></tr><tr><td>P1</td><td>3</td><td>T6</td><td>8.3</td><td>17.3</td><td>29.76</td><td>1.64</td><td>10.55</td><td>111</td><td>10.91</td><td>219.84</td><td>5.3</td></tr><tr><td>P1</td><td>5</td><td>T1</td><td>8.1</td><td>25.3</td><td>43.52</td><td>2.26</td><td>11.19</td><td>221</td><td>10.7</td><td>215.61</td><td>5.88</td></tr><tr><td>P1</td><td>5</td><td>T2</td><td>8.5</td><td>12.1</td><td>20.81</td><td>01.01</td><td>11.98</td><td>95</td><td>10.23</td><td>206.13</td><td>3.33</td></tr><tr><td>P1</td><td>5</td><td>T3</td><td>8.5</td><td>11</td><td>18.92</td><td>0.81</td><td>13.58</td><td>91</td><td>7.31</td><td>147.3</td><td>03.03</td></tr><tr><td>P1</td><td>5</td><td>T4</td><td>8.3</td><td>9.6</td><td>16.51</td><td>0.91</td><td>10.55</td><td>92</td><td>6.74</td><td>135.81</td><td>04.01</td></tr><tr><td>P1</td><td>5</td><td>T5</td><td>8.1</td><td>17.8</td><td>30.62</td><td>1.37</td><td>12.99</td><td>100</td><td>9.15</td><td>184.37</td><td>11.35</td></tr><tr><td>P1</td><td>5</td><td>T6</td><td>8.5</td><td>8.8</td><td>15.14</td><td>01.03</td><td>8.54</td><td>98</td><td>7.19</td><td>144.88</td><td>4.15</td></tr><tr><td>P2</td><td>1</td><td>T1</td><td>8.3</td><td>22.9</td><td>39.39</td><td>1.89</td><td>12.12</td><td>127</td><td>11.19</td><td>225.48</td><td>4.51</td></tr><tr><td>P2</td><td>1</td><td>T2</td><td>8.6</td><td>17.3</td><td>29.76</td><td>1.2</td><td>14.42</td><td>192</td><td>10.43</td><td>210.16</td><td>5.54</td></tr><tr><td>P2</td><td>1</td><td>T3</td><td>8.6</td><td>11.7</td><td>20.12</td><td>0.73</td><td>16.03</td><td>59</td><td>14.7</td><td>296.21</td><td>4.21</td></tr><tr><td>P2</td><td>1</td><td>T4</td><td>8.5</td><td>12.1</td><td>20.81</td><td>1</td><td>12.1</td><td>123</td><td>9.77</td><td>196.87</td><td>05.06</td></tr><tr><td>P2</td><td>1</td><td>T5</td><td>8.3</td><td>19.3</td><td>33.2</td><td>1.5</td><td>12.87</td><td>247</td><td>12.64</td><td>254.7</td><td>9.58</td></tr><tr><td>P2</td><td>1</td><td>T6</td><td>8.1</td><td>27.5</td><td>47.3</td><td>2.35</td><td>11.7</td><td>311</td><td>13.8</td><td>278.07</td><td>12.59</td></tr><tr><td>P2</td><td>1</td><td>T7</td><td>8.3</td><td>22.7</td><td>39.04</td><td>1.58</td><td>14.37</td><td>208</td><td>11.2</td><td>225.68</td><td>7.61</td></tr><tr><td>P2</td><td>3</td><td>T1</td><td>8.2</td><td>29.2</td><td>50.22</td><td>1.97</td><td>14.82</td><td>196</td><td>11.4</td><td>229.71</td><td>8.58</td></tr><tr><td>P2</td><td>3</td><td>T2</td><td>8.5</td><td>17.6</td><td>30.27</td><td>1.38</td><td>12.75</td><td>206</td><td>12.16</td><td>245.02</td><td>7.67</td></tr><tr><td>P2</td><td>3</td><td>T3</td><td>8.4</td><td>19.3</td><td>33.2</td><td>1.45</td><td>13.31</td><td>188</td><td>13.15</td><td>264.97</td><td>10.02</td></tr><tr><td>P2</td><td>3</td><td>T4</td><td>8.2</td><td>19.1</td><td>32.85</td><td>1.42</td><td>13.45</td><td>280</td><td>12.1</td><td>243.82</td><td>7.87</td></tr><tr><td>P2</td><td>3</td><td>T5</td><td>8</td><td>24.1</td><td>41.45</td><td>1.98</td><td>12.17</td><td>264</td><td>13.67</td><td>275.45</td><td>11.18</td></tr><tr><td>P2</td><td>3</td><td>T6</td><td>8.3</td><td>26.9</td><td>46.27</td><td>1.61</td><td>16.71</td><td>235</td><td>14.38</td><td>289.76</td><td>12.58</td></tr><tr><td>P2</td><td>5</td><td>T1</td><td>8.5</td><td>14.9</td><td>25.63</td><td>0.99</td><td>15.05</td><td>147</td><td>10.48</td><td>211.17</td><td>6.77</td></tr><tr><td>P2</td><td>5</td><td>T2</td><td>8.5</td><td>14.1</td><td>24.25</td><td>0.92</td><td>15.33</td><td>77</td><td>11.2</td><td>225.68</td><td>6.96</td></tr><tr><td>P2</td><td>5</td><td>T3</td><td>8.5</td><td>9.8</td><td>16.86</td><td>0.73</td><td>13.42</td><td>62</td><td>10.05</td><td>202.51</td><td>5.37</td></tr><tr><td>P2</td><td>5</td><td>T4</td><td>8.4</td><td>15</td><td>25.8</td><td>0.98</td><td>15.31</td><td>163</td><td>11.05</td><td>222.66</td><td>7.6</td></tr><tr><td>P2</td><td>5</td><td>T5</td><td>8.2</td><td>20.4</td><td>35.09</td><td>1.59</td><td>12.83</td><td>253</td><td>11.93</td><td>240.39</td><td>10.76</td></tr><tr><td>P2</td><td>5</td><td>T6</td><td>8.4</td><td>15.2</td><td>26.14</td><td>1.14</td><td>13.33</td><td>94</td><td>8.46</td><td>170.47</td><td>4.96</td></tr><tr><td>P2</td><td>5</td><td>T7</td><td>8.2</td><td>26.8</td><td>46.1</td><td>2.23</td><td>12.02</td><td>180</td><td>11.49</td><td>231.52</td><td>7.41</td></tr><tr><td>P2</td><td>2</td><td>T1</td><td>8.6</td><td>9.9</td><td>17.03</td><td>0.76</td><td>13.03</td><td>63</td><td>8.54</td><td>172.08</td><td>3.27</td></tr><tr><td>P2</td><td>2</td><td>T2</td><td>8.5</td><td>10.3</td><td>17.72</td><td>0.86</td><td>11.98</td><td>60</td><td>10.74</td><td>216.41</td><td>4.85</td></tr><tr><td>P2</td><td>2</td><td>T3</td><td>8.5</td><td>13.6</td><td>23.39</td><td>1.1</td><td>12.36</td><td>110</td><td>9.83</td><td>198.07</td><td>6.73</td></tr><tr><td>P2</td><td>2</td><td>T4</td><td>8.3</td><td>18.9</td><td>32.51</td><td>1.4</td><td>13.5</td><td>233</td><td>9.67</td><td>194.85</td><td>9.68</td></tr><tr><td>P2</td><td>2</td><td>T5</td><td>8</td><td>24.6</td><td>42.31</td><td>1.91</td><td>12.88</td><td>187</td><td>10.87</td><td>219.03</td><td>08.04</td></tr></tbody></table>
<table><tr><td>P2</td><td>2</td><td>T6</td><td>8.4</td><td>14.3</td><td>24.6</td><td>01.02</td><td>14.02</td><td>124</td><td>8.59</td><td>173.09</td><td>5.66</td></tr><tr><td>P2</td><td>2</td><td>T7</td><td>8.2</td><td>19.4</td><td>33.37</td><td>1.62</td><td>11.98</td><td>179</td><td>10.35</td><td>208.55</td><td>07.02</td></tr><tr><td>P2</td><td>4</td><td>T1</td><td>8.5</td><td>14</td><td>24.08</td><td>01.02</td><td>13.73</td><td>127</td><td>9.29</td><td>187.19</td><td>3.47</td></tr><tr><td>P2</td><td>4</td><td>T2</td><td>8.5</td><td>11.2</td><td>19.26</td><td>0.85</td><td>13.18</td><td>32</td><td>6.82</td><td>137.42</td><td>3.13</td></tr><tr><td>P2</td><td>4</td><td>T3</td><td>8.5</td><td>12</td><td>20.64</td><td>01.08</td><td>11.11</td><td>111</td><td>9.26</td><td>186.59</td><td>7</td></tr><tr><td>P2</td><td>4</td><td>T4</td><td>8.3</td><td>16</td><td>27.52</td><td>1.44</td><td>11.11</td><td>148</td><td>10.52</td><td>211.98</td><td>12.08</td></tr><tr><td>P2</td><td>4</td><td>T5</td><td>8.1</td><td>20</td><td>34.4</td><td>1.67</td><td>11.98</td><td>183</td><td>11.86</td><td>238.98</td><td>9.65</td></tr><tr><td>P2</td><td>4</td><td>T6</td><td>8.3</td><td>20.4</td><td>35.09</td><td>1.44</td><td>14.17</td><td>131</td><td>10.22</td><td>205.93</td><td>6.43</td></tr><tr><td>P2</td><td>4</td><td>T7</td><td>8.4</td><td>18.7</td><td>32.16</td><td>1.46</td><td>12.81</td><td>128</td><td>9.29</td><td>187.19</td><td>6.61</td></tr><tr><td>P2</td><td>6</td><td>T1</td><td>8.5</td><td>14.9</td><td>25.63</td><td>1.23</td><td>12.11</td><td>130</td><td>10.24</td><td>206.34</td><td>5.89</td></tr><tr><td>P2</td><td>6</td><td>T2</td><td>8.5</td><td>12</td><td>20.64</td><td>0.95</td><td>12.63</td><td>61</td><td>10.75</td><td>216.61</td><td>8.48</td></tr><tr><td>P2</td><td>6</td><td>T3</td><td>8.6</td><td>8.6</td><td>14.79</td><td>0.87</td><td>9.89</td><td>70</td><td>8.62</td><td>173.69</td><td>6.55</td></tr><tr><td>P2</td><td>6</td><td>T4</td><td>8.4</td><td>14.4</td><td>24.77</td><td>01.03</td><td>13.98</td><td>111</td><td>11.36</td><td>228.9</td><td>11</td></tr><tr><td>P2</td><td>6</td><td>T5</td><td>8.3</td><td>17.4</td><td>29.93</td><td>1.35</td><td>12.89</td><td>178</td><td>11.71</td><td>235.96</td><td>10.97</td></tr><tr><td>P2</td><td>6</td><td>T6</td><td>8.1</td><td>25.6</td><td>44.03</td><td>2.25</td><td>11.38</td><td>163</td><td>12.16</td><td>245.02</td><td>7.71</td></tr><tr><td>P2</td><td>6</td><td>T7</td><td>8.5</td><td>16</td><td>27.52</td><td>01.06</td><td>15.09</td><td>144</td><td>9.58</td><td>193.04</td><td>4.65</td></tr></table>
<table><thead><tr><th>K2O (mg/kg)</th><th>SODIUM</th><th>Na2O (mg/kg)</th><th>CEC</th></tr></thead><tbody><tr><td>175.21</td><td>0.66</td><td>20.46</td><td>81</td></tr><tr><td>130.47</td><td>0.37</td><td>11.47</td><td>73</td></tr><tr><td>458.28</td><td>1.31</td><td>40.61</td><td>125</td></tr><tr><td>272.71</td><td>0.45</td><td>13.95</td><td>91</td></tr><tr><td>296.26</td><td>0.89</td><td>27.59</td><td>107</td></tr><tr><td>292.49</td><td>0.53</td><td>16.43</td><td>109</td></tr><tr><td>148.84</td><td>0.5</td><td>15.5</td><td>99</td></tr><tr><td>199.23</td><td>0.6</td><td>18.6</td><td>98</td></tr><tr><td>141.3</td><td>0.5</td><td>15.5</td><td>86</td></tr><tr><td>336.77</td><td>0.94</td><td>29.14</td><td>99</td></tr><tr><td>277.89</td><td>0.72</td><td>22.32</td><td>109</td></tr><tr><td>249.63</td><td>0.6</td><td>18.6</td><td>111</td></tr><tr><td>276.95</td><td>0.86</td><td>26.66</td><td>145</td></tr><tr><td>156.84</td><td>0.78</td><td>24.18</td><td>112</td></tr><tr><td>142.71</td><td>0.54</td><td>16.74</td><td>84</td></tr><tr><td>188.87</td><td>0.39</td><td>12.09</td><td>77</td></tr><tr><td>534.59</td><td>0.7</td><td>21.7</td><td>108</td></tr><tr><td>195.47</td><td>01.05</td><td>32.55</td><td>88</td></tr><tr><td>212.42</td><td>0.62</td><td>19.22</td><td>129</td></tr><tr><td>260.93</td><td>0.73</td><td>22.63</td><td>107</td></tr><tr><td>198.29</td><td>1</td><td>31</td><td>209</td></tr><tr><td>238.33</td><td>0.67</td><td>20.77</td><td>126</td></tr><tr><td>451.22</td><td>0.48</td><td>14.88</td><td>136</td></tr><tr><td>592.99</td><td>0.92</td><td>28.52</td><td>148</td></tr><tr><td>358.43</td><td>0.63</td><td>19.53</td><td>132</td></tr><tr><td>404.12</td><td>0.82</td><td>25.42</td><td>151</td></tr><tr><td>361.26</td><td>0.6</td><td>18.6</td><td>120</td></tr><tr><td>471.94</td><td>0.69</td><td>21.39</td><td>113</td></tr><tr><td>370.68</td><td>0.63</td><td>19.53</td><td>129</td></tr><tr><td>526.58</td><td>0.66</td><td>20.46</td><td>126</td></tr><tr><td>592.52</td><td>01.09</td><td>33.79</td><td>131</td></tr><tr><td>318.87</td><td>0.51</td><td>15.81</td><td>106</td></tr><tr><td>327.82</td><td>0.75</td><td>23.25</td><td>138</td></tr><tr><td>252.93</td><td>0.7</td><td>21.7</td><td>142</td></tr><tr><td>357.96</td><td>0.67</td><td>20.77</td><td>128</td></tr><tr><td>506.8</td><td>0.7</td><td>21.7</td><td>129</td></tr><tr><td>233.62</td><td>0.63</td><td>19.53</td><td>112</td></tr><tr><td>349.01</td><td>0.83</td><td>25.73</td><td>164</td></tr><tr><td>154.02</td><td>0.67</td><td>20.77</td><td>83</td></tr><tr><td>228.44</td><td>0.77</td><td>23.87</td><td>159</td></tr><tr><td>316.98</td><td>0.55</td><td>17.05</td><td>126</td></tr><tr><td>455.93</td><td>0.56</td><td>17.36</td><td>121</td></tr><tr><td>378.68</td><td>0.82</td><td>25.42</td><td>128</td></tr></tbody></table>
<table><tr><td>266.59</td><td>0.41</td><td>12.71</td><td>100</td></tr><tr><td>330.64</td><td>0.57</td><td>17.67</td><td>125</td></tr><tr><td>163.44</td><td>1</td><td>31</td><td>87</td></tr><tr><td>147.42</td><td>0.95</td><td>29.45</td><td>138</td></tr><tr><td>329.7</td><td>0.89</td><td>27.59</td><td>105</td></tr><tr><td>568.97</td><td>0.65</td><td>20.15</td><td>94</td></tr><tr><td>454.52</td><td>0.77</td><td>23.87</td><td>131</td></tr><tr><td>302.85</td><td>1</td><td>31</td><td>117</td></tr><tr><td>311.33</td><td>0.46</td><td>14.26</td><td>130</td></tr><tr><td>277.42</td><td>0.79</td><td>24.49</td><td>111</td></tr><tr><td>399.41</td><td>0.68</td><td>21.08</td><td>131</td></tr><tr><td>308.51</td><td>0.6</td><td>18.6</td><td>133</td></tr><tr><td>518.1</td><td>0.54</td><td>16.74</td><td>125</td></tr><tr><td>516.69</td><td>0.59</td><td>18.29</td><td>140</td></tr><tr><td>363.14</td><td>1.27</td><td>39.37</td><td>149</td></tr><tr><td>219.02</td><td>0.53</td><td>16.43</td><td>96</td></tr></table>
<table><thead><tr><th>Name</th><th>Description</th></tr></thead><tbody><tr><td>Field</td><td>Study site where the sample was taken, P1 is the Roujan vineyard and P2 is the Combaillaux vii</td></tr><tr><td>Interrow</td><td>Index of the interrow sampled</td></tr><tr><td>Transect</td><td>Position along the transect sampled, T1 is upslope of the field</td></tr></tbody></table>
neyard
<figure><img src="image_14.png" /></figure>

Appendix B. Mean E/S ratio (number of seedlings counted divided by the surface of one sample) values and standard deviation for each study site (P1 : Roujan study site, P2 : Combaillaux study site) and each dispersal mode considered, separated by the zones accounted for in the statistical analyses. Combined effect of dispersal mode and zone sampled was tested by performing an ANOVA on a generalized linear model for each site separately, and differences between zones and between dispersal modes were tested using Tukey tests. Significant differences between dispersal modes are represented by different lowercase letters, and differences between zones are represented by different uppercase letters.

<table><thead><tr><th rowspan="2"></th><th colspan="2">Anemochory</th><th colspan="2">Barochory</th><th colspan="2">Hydrochory</th><th colspan="2">Zoochory</th></tr><tr><th>P1</th><th>P2</th><th>P1</th><th>P2</th><th>P1</th><th>P2</th><th>P1</th><th>P2</th></tr></thead><tbody><tr><td>Study site</td><td>34.6 +/- 23.1<sup>a</sup></td><td>32.5 +/- 32<sup>a</sup></td><td>128 +/- 258<sup>b</sup></td><td>133 +/- 218<sup>c</sup></td><td>34 +/- 42.5<sup>a</sup></td><td>87.3 +/- 102<sup>b</sup></td><td>28.9 +/- 28.4<sup>a</sup></td><td>37.2 +/- 34.5<sup>a</sup></td></tr><tr><td>Field</td><td>22.0 +/- 12.84<sup>A</sup></td><td>20.96 +/- 17.24<sup>Ab</sup></td><td>43.95 +/- 33.64<sup>A</sup></td><td>160.04 +/- 282.66<sup>Aa</sup></td><td>24.53 +/- 14.04</td><td>115.79 +/- 123.27<sup>Aa</sup></td><td>23.6 +/- 18.2</td><td>26.56 +/- 29.22<sup>Ab</sup></td></tr><tr><td>Field edge</td><td>38.95 +/- 28.14<sup>Bac</sup></td><td>28.93 +/- 24.44<sup>Ba</sup></td><td>248.64 +/- 445.41<sup>Bb</sup></td><td>122.72 +/- 135.7<sup>Bb</sup></td><td>51.78 +/- 78.51<sup>a</sup></td><td>49.47 +/- 60.88<sup>Aa</sup></td><td>39.76 +/- 44.81<sup>a</sup></td><td>38.02 +/- 29.8<sup>Ba</sup></td></tr><tr><td>Ditch</td><td>47.71 +/- 20.93<sup>Ca</sup></td><td>77.5 +/- 43.88<sup>C</sup></td><td>150.69 +/- 166.67<sup>Bb</sup></td><td>61.53 +/- 44.28<sup>AB</sup></td><td>33.23 +/- 13.85<sup>a</sup></td><td>75.36 +/- 43.86<sup>B</sup></td><td>28.77 +/- 20.97<sup>a</sup></td><td>74.47 +/- 40.12<sup>C</sup></td></tr></tbody></table>