Edaphoclimatic drivers of the effect of extensive vegetation management on ecosystem services and biodiversity in vineyards

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Agriculture, Ecosystems and Environment 339 (2022) 108115

Available online 12 August 2022

Edaphoclimatic drivers of the effect of extensive vegetation management on ecosystem services and biodiversity in vineyards

Carmen Chapela-Oliva

^a

, Silvia Winter

^b

, Raúl Ochoa-Hueso

^a^,^c^,^*

aDepartment of Biology, IVAGRO, University of C´adiz, Campus de Excelencia Internacional Agroalimentario (CeiA3), Campus del Rio San Pedro, 11510 Puerto Real, C´adiz, Spain

bInstitute of Plant Protection and Integrative Nature Conservation Research, University of Natural Resources and Life Sciences, Gregor-Mendel-Str. 33, A-1180 Vienna, Austria

cDepartment of Terrestrial Ecology, Netherlands Institute of Ecology (NIOO-KNAW), P.O. Box 50, 6700 AB Wageningen, the Netherlands

A R T I C L E I N F O Keywords:

Biodiversity Climate Cover crops Ecosystem services Meta-analysis Modulating factors Soil

Vineyards Vegetation cover

A B S T R A C T

Vineyards are a very important perennial woody crop globally, but they are also among the most intensively managed agroecosystems. This has resulted in biologically impoverished and highly eroded vineyards.

Environmentally-friendly inter-row vegetation management, particularly the use of cover crops, could contribute to avoid erosion and regenerate soil biodiversity in vineyards. In this study, we updated a global meta-analysis on the effects of inter-row extensive vegetation cover management, particularly through the use of cover crops, on ecosystem services, including supporting, regulating and provisioning services, and biodiversity in vineyards. We also analyzed the role of environmental variables (climate, including precipitation- and temperature-related bioclimatic variables and soil properties, including pH and texture) and irrigation in modulating these effects.

The presence of extensive vegetation cover consistently increased biodiversity, as well as supporting ecosystem services in irrigated vineyards and regulating services in rainfed vineyards. Provisioning services, which were evaluated as grape yield, were slightly negatively affected in rainfed vineyards, but not in irrigated ones. The effects of vegetation cover on ecosystem services varied depending on the climate and edaphic characteristics of vineyards. For example, supporting ecosystem services were favored in acidic soils and were also positively related to the precipitation of the wettest quarter, whereas regulating services were particularly enhanced in alkaline soils and in locations with lower temperatures of the wettest quarter. Biodiversity was especially favored in locations with lower precipitation seasonality. Taken together, our study indicates the importance of developing strategies for the adaptive management of extensive vegetation covers tailored to the climatic and edaphic conditions of each vineyard. This adaptive management, combined with irrigation and potentially other locally- tailored adaptive strategies, could also contribute to further mitigate potential negative effects of vegetation cover on grape production while maximizing other ecosystem services such as provisioning and supporting services and biodiversity.

1. Introduction

Vineyards are very important perennial woody crop globally (OIV, 2019). The world surface of this type of cultivation is 7.4 million hect- ares, regardless of the purpose for which they are grown (wine or table grapes), and including those areas that do not yet have productive capacity (OIV, 2019). The leading countries in terms of vineyard area worldwide are Spain with 969,000 ha, followed by China and France (OIV, 2019). The planting of vineyards, whose origin dates back more

than 7000 years in the Middle East (Armenia and Georgia) (Riera, 2014), is one of the oldest and most profitable agricultural practices in the world (OIV, 2019). Traditionally, vineyards have hosted high levels of biodiversity at different taxonomic levels, including plants, birds, and insects (Paiola et al., 2020), as they were managed as multifunctional systems including fruit trees, annual crops, and other forms of land use such as pastures (Nicholls et al., 2008). However, vineyards are currently among the most intensively managed agroecosystems due to the intensive use of pesticides such as fungicides, herbicides and

* Corresponding author at: Department of Biology, IVAGRO, University of C´adiz, Campus de Excelencia Internacional Agroalimentario (CeiA3), Campus del Rio San Pedro, 11510 Puerto Real, C´adiz, Spain.

E-mail address: [email protected] (R. Ochoa-Hueso).

Contents lists available at ScienceDirect

Agriculture, Ecosystems and Environment

journal homepage: www.elsevier.com/locate/agee

https://doi.org/10.1016/j.agee.2022.108115

Received 19 October 2021; Received in revised form 20 July 2022; Accepted 22 July 2022

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insecticides, as well as frequent soil tillage, which decrease their biodiversity at multiple trophic levels (Nicholls et al., 2008). These intensive practices have, thus, resulted in biologically impoverished and highly eroded vineyards (G´omez et al., 2011; Winter et al., 2018). For this reason, it is important to develop sustainable land management strategies and practices that contribute to preserve or even regenerate the biodiversity and ecosystem services in agroecosystems such as vineyards, including soil fertility and nutrient cycling (supporting services), carbon sequestration, hydrological regulation, and pest and disease control (regulating services) (Daily, 2003; Reid et al., 2005;

Tscharntke et al., 2012). This need also raises the general challenge of how to promote biodiversity and ecosystem services in vineyards and other crops while maintaining production and the quality of crops (i.e., provisioning services) to feed a constantly growing population (Paiola et al., 2020).

One of the ways that have been proposed to avoid soil erosion and regenerate biodiversity in vineyards is through inter-row vegetation management, particularly through the use of cover crops (Lu et al., 2000). The use of cover crops in the inter-row spaces of vineyards is a traditional technique implemented since the times of Romans that has been widely replaced by tillage to control weeds, particularly in Medi- terranean areas (Blackshaw et al., 2001). Cover crops are a living vegetation cover that protects the soil and can be both temporary and permanent, seeded or spontaneous, as well as in association with other plants in an intercrop, relay, or in rotation (Garcia et al., 2018). Most cover crops are legumes and grasses, although they can belong to any plant family or even form complex communities made up of a multitude of species belonging to different functional groups (Garcia et al., 2018).

Cover crops can be used to recover degraded soils because they offer a wide variety of functions, such as increasing soil carbon sequestration and fertility, restoring soil structure and aggregation, reducing agro- chemical residues, improving water infiltration, competing with un- wanted weeds, providing beneficial mycorrhizal inoculum sources, as well as increasing the biodiversity of agroecosystems at multiple trophic levels, among others (Abad et al., 2021a; Garcia et al., 2018). For example, it has been shown that when winegrowers used cover crops instead of tilling the soil, erosion rates were drastically reduced (Ruiz-Colmenero et al., 2011; Winter et al., 2018). The positive effects of the use of these crops for pest control in vineyards have also been demonstrated (Berndt et al., 2006; Sanguankeo and Le´on, 2011), although some plant species can also cause an increase in pest species by acting as host plants and providing favorable habitats for pests (Begum et al., 2006). These disparate results suggest a context-dependent response in the ability of cover crops to provide certain ecosystem services in vineyards. However, the role that environmental factors such as climatic conditions (e.g., temperature and rainfall) and soil properties (e.g., pH and texture) may have in controlling the effect of inter-row vegetation cover management on the biodiversity and ecosystem services in vineyards around the world remains largely unknown.

Macro-climatic conditions, including precipitation and temperature, as well as variations in soil properties such as pH and texture due to different soil parent material have been long known as critical environmental factors controlling the metabolism of grapevines and their associated microbial communities, thus determining wine yield and quality, finally contributing to the expression of wine terroir (Bokulich et al., 2014). This suggests that these environmental factors could also be important drivers of the response of vineyards to inter-row vegetation management (Schultz and Stoll, 2010; Xu et al., 2017). For example, in dry climates, cover crops could be advantageous because they help to retain soil water (Ruiz-Colmenero et al., 2011). However, their use in dry climates could also imply that at critical times for vine physiology, such as during the characteristic summer drought period in Mediterra- nean climates, there is competition for soil resources such as nutrients like nitrogen and water between the cover crops and the vines (Ward et al., 2012). This has led to poor adoption among winegrowers in certain areas, particularly where irrigation is not feasible because it is

not economically viable, or not allowed by Protected Designations of Origin (PODs). Thus some authors have suggested the importance of granting subsidies for the establishment of cover crops depending on the macro-climatic location of the vineyards as part of the agri-environmental policies of each country or region (Winter et al., 2018). In addition, it is possible that different ecosystem services are influenced in different ways by various environmental variables, both edaphic and climatic, and even that there is a compromise on what type of services (provisioning, regulating, and supporting) are maximized or reduced depending on these environmental conditions. This knowledge would allow us to make intelligent adaptive management decisions that could help us to maximize biodiversity and the provision of ecosystem services in vineyards.

In this study, we carried out an update of a global meta-analysis on the effects of vegetation management intensity on ecosystem services in vineyards around the world (Winter et al., 2018), focusing here on the role of inter-row vegetation cover management, particularly cover crops. We also sought to analyze the role that macro-climatic variables, soil properties linked to parent material characteristics, and irrigation play in modulating such effects, for which we currently lack information. We only considered the role of those climatic variables that are considered decisive for the functioning of vineyards and vine physiology, including: (i) mean annual rainfall and temperature, as indicative of long-term climatic conditions; (ii) mean temperature of the wettest quarter and precipitation of the warmest quarter, broadly indicative of climatic conditions during the plant growing season; and (iii) precipitation of the wettest quarter and mean temperature of the coldest quarter, broadly indicative of potential climatic constraints linked to water and temperature stress. We predict that, globally, the effects of the presence of cover crops on ecosystem services and biodiversity in vineyards will be positive (Winter et al., 2018). Furthermore, we predict that these effects will be more positive in arid climates, where a greater accumulation of organic matter, a higher rate of nutrient cycling, and greater diversity under vegetation have been demonstrated compared to adjacent bare soil areas (Ochoa-Hueso et al., 2018). This is because plants can act as a shelter, buffering extreme temperatures and increasing soil moisture values, resulting in greater fertility (Ochoa-- Hueso et al., 2018). We also predict that the responses of different types of ecosystem services to inter-row management will depend on the type of ecosystem service considered. We predict that climatic variables that are representative of conditions during the growing season will be particularly important in the response of regulating (e.g., carbon sequestration and pest regulation) and provisioning (e.g., grape yield) services to extensive vegetation cover management. In contrast, we predict that supporting ecosystem services will be more affected by soil variables such as pH and texture, due to the known role of these variables in controlling the abundance of organisms and their activity in soils, and irrigation due to the mitigation of water competition.

2. Materials and methods 2.1. Bibliographic search

To carry out this study, we collected data published in the literature on the effects of extensive vegetation management (cover crops) on ecosystem services and biodiversity in vineyards on a global scale.

Ecosystem services, defined as the benefits that people obtain from ecosystems, include regulating services (e.g., regulation of the carbon and water cycles, and pest control), supporting services (e.g., soil fertility and nutrient cycling), and provisioning services (in this study we only considered crop yield, while indicators of crop quality were not considered) (Daily, 2003; Reid et al., 2005; Tscharntke et al., 2012).

As a starting point, we used the meta-analysis of Winter et al. (2018), whose database was updated for this work with studies published between 2016 and 2020 in SCOPUS and Web of Science. In addition, we added information for geographic (latitude and longitude) and edaphic

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variables (pH, and texture [percentage of clay and sand]) for each study site. Novel aspects of our study include the explicit consideration of edaphoclimatic drivers and irrigation for controlling the response of the different ecosystem services in relation to inter-row management , which is a critical knowledge gap for the successful implementation of adaptive management strategies in sustainable viticulture (Darnhofer et al., 2010; Garcia et al., 2018; Ripoche et al., 2011). The keywords used for the search were those previously used by Winter et al. (2018).

The selection criteria applied were: (1) field studies had to include plots treated with cover crops that were managed extensively and controls without cover crops (i.e., bare soil established with tillage and/or herbicides); (2) the studies should also contain information regarding means and any measure of dispersion of the dependent variable, either in numerical values or represented in graphs; and (3) should preferably provide information regarding the geographical location of the study site and soil characteristics. This resulted in a subset of the original meta-analysis published by Winter et al. (2018).

2.2. Data extraction

The following information was obtained for each article: country, ecosystem service category (provisioning, regulating, supporting, or biodiversity), type of ecosystem service (e.g., carbon sequestration, pest control, etc.), irrigation, type of control used (e.g., bare soil or conventional management), sample size, latitude, longitude, soil pH, percentage of sand and clay, as well as the mean and standard deviation of the variable of interest under control and treatment conditions. Based on the geographical coordinates, we also obtained data on the previously mentioned bioclimatic variables (i.e., mean annual rainfall, mean annual temperature, mean temperature of the wettest quarter, precipitation of the warmest quarter, precipitation of the wettest quarter, and mean temperature of the coldest quarter, and precipitation seasonality) from WorldClim 2 (Fick and Hijmans, 2017). When the numerical values of the control and treatment means were not directly provided and were only represented in figures, we used the Digitize package (Poisot, 2011) in R (version 4.0.2), which provides a reliable estimate of the numerical data from its graphical representation.

2.3. Numerical calculations and statistical analyses

Once the numerical values of the control and treatment means were obtained, the natural logarithm of the response ratio (lnRR) was calculated as the estimate of the size of the effect associated with the presence of cover crops, according to Eq. (1):

lnRR = ln (Xt/Xc) (1)

where Xt andXc are the means of the treatment and the control, respectively. Obtaining a value of zero indicates that there is no change in the response of the variable due to cover crops, while a positive or negative value represents an increase or decrease in the response as compared with the control (bare soil inter-rows). In addition, the vari- ance of the natural logarithm of the response was calculated according to Eq. (2):

VlnRR = ( St²

nt Xt² )

+ ( Sc²

nc Xc² )

(2) where St and Sc are the standard deviation of the treatment and the control respectively and nt and nc are the sample size of the treatment and control.

Prior to analyses, we summarized the dataset so we would only have one observation per article (overall analyses), or one observation per article and service type (service-specific analyses). These data were used to evaluate the effect of cover crops (overall effects, and effects with and without irrigation separately) on ecosystem services, for which we used the rma.mv function from the metafor package in R (Viechtbauer, 2010).

To explore the relationship between the response of ecosystem services to the presence of cover crops and climatic and edaphic variables, we used Pearson correlations using the “rcorr” function of the Hmisc package (Harrell, 2020). We then used structural equation models (Eisenhauer et al., 2015) in order to evaluate the causality of the modulating role of a series of climatic and soil variables on the size of the response of ecosystem services to the presence of cover crops in vineyards. This SEM approach offers the opportunity to evaluate the direct and indirect effects that various variables may have on the response variable based on an a priori conceptual model based on previous knowledge (Eisenhauer et al., 2015). To do this, we used the piece- wiseSEM package (version 2.1.0) (Lefcheck, 2015) in R, where a set of linear equations are evaluated individually. To run the linear models, we used the "lm" function of the Stats package. A good fit was assumed when the p-value of Fisher’s C was greater than 0.05.

3. Results

The literature search between 2016 and 2020 resulted in 17 articles, of which only five were relevant to our study. After updating the literature, our meta-analysis included results from a total of 71 articles (Table 1). Most of the articles conducted their study in those countries with the highest wine production such as Italy, France, Spain, United States, Australia, Chile, South Africa and Germany (Fig. 1). Approxi- mately 64% of the dataset comprised studies in rainfed vineyards (n = 46), and 36% in irrigated vineyards (n = 26; one study had both rainfed and irrigated vineyards). The most predominant type of ecosystem service in our dataset was regulating services (n = 47 observations, of which a majority of observations corresponded to soil carbon sequestration and pest regulation), while biodiversity was the least represented response variable (n = 11 observations).

Extensive vegetation cover management increased the supply of biodiversity and ecosystem services in vineyards (Fig. 2), especially supporting services, and regulating services. However, provisioning ecosystem services (grape yield) were not significantly affected (marginally significant effect; P = 0.085). Extensive vegetation cover management was particularly important for regulating services and biodiversity in rainfed vineyards. However, extensive vegetation cover management favored supporting ecosystem services particularly in irrigated vineyards. Provisioning services were not significantly affected when vineyards were irrigated. However, without irrigation, we found a negative effect of Extensive vegetation cover management on provisioning services (i.e., grape yield). The effect of cover crops on irrigated vineyards was comparable to the effect on non-irrigated ones when all ecosystem services were analyzed together.

Multiple environmental variables, particularly pH, precipitation of the wettest quarter, and precipitation of the warmest quarter, modulated the effect of extensive vegetation cover management on the supply of different ecosystem services and biodiversity. The response of supporting ecosystem services was more positive when the mean temperature of coldest quarter was higher and the pH was lower (Fig. 3; Fig. 4-A), whereas regulating services were more favored in high-pH soils where precipitation of the wettest quarter was also higher (Fig. 3; Fig. 4-B). The significant effect of pH on supporting services was only found when other drivers were jointly considered in our SEM. Further exploration by category of service type within regulating services showed that soil carbon sequestration improved regardless of the environmental context (Fig. S1), while pest control was particularly favored at places with higher precipitation of the warmest quarter (Fig. S1). Biodiversity benefited more in locations with lower precipitation seasonality (Fig. 3).

Regarding provisioning services, only irrigation had a marginally significant effect in our SEM (Fig. 4-D), while we did not find any corre- lation between soil and climatic variables and the effect of extensive vegetation cover management on grape yield, either considered as a whole or separately for irrigated and rainfed vineyards (Fig. S1).

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Table 1

Summary of the 71 articles used for this study.

Publication Country Climate Latitude Longitude Provisioning Regulating Supporting Biodiversity

Adams, 2011 USA Mediterranean 35,674 -120,677 X X

Amaral et al., 2012 Brazil Oceanic -23,417 -51,950 X

Barrio et al., 2011 Spain Mediterranean 37,550 -4617 X X

Barroso et al., 2016 Portugal Mediterranean 38,567 -7900 X

Bartoli & Dousset, 2011 France Oceanic 46,416 4815 X

Baumgartner et al., 2005 USA Mediterranean 36,317 -121,233 X

Belmonte et al., 2016 Italy Mediterranean 44,468 8627 X

Blanco-P´erez et al., 2020 Spain Mediterranean 42,433 -2500 X

Blavet et al., 2009 France Mediterranean 43,860 3939 X

Botton et al., 2010 Brazil Oceanic -29,404 -51,719 X

Bouffaud et al., 2016 France Oceanic 47,000 4500 X X

Bruggisser et al., 2010 Swiss Oceanic 47,083 7133 X

Burns et al., 2015 USA Mediterranean 38,330 -122,300 X

Cap´o - Bauç `a et al., 2019 Spain Mediterranean 39,521 2521 X X X

Celette et al., 2005 France Mediterranean 43,721 4174 X

Cluzeau et al., 2013 France Oceanic 45,833 -0.917 X X

Coll et al., 2011 France Mediterranean 43,191 2817 X X

Coll et al., 2012 France Mediterranean 43,191 2817 X

Costello & Danne, 1998 USA Mediterranean 36,583 -119,450 X

Costello & Danne, 2003 USA Conti-steppe 36,600 -119,517 X X

Costello, 2010a USA Conti-steppe 36,600 -119,517 X

Costello, 2010b USA Conti-steppe 36,600 -119,517 X

Daane & Costello, 1998 USA Mediterranean 36,600 -120,183 X

Danne et al. (2010) Australia Conti-steppe -34,467 139,000 X

English-Loeb et al., 2003 USA Conti-steppe 42,760 -79,945 X

Favretto et al., 1992 Italy Mediterranean 42,416 11,478 X X

Garcia-Diaz et al., 2017 Spain Conti-steppe 40,417 -3703 X

Giese et al., 2014 USA Oceanic 36,350 -80,767 X

Gomez et al., 2011 France Mediterranean X

Guzm´an et al. (2019) Spain Mediterranean 37,633 -4783 X X X

Hall et al. (2020) Europe Mediterranean X

Hanna et al., 2003 USA Conti-steppe 36,831 120,024 X

Ingels et al., 2005 USA Mediterranean 38,450 -121,350 X X

Irvin et al., 2016 USA Mediterranean 33,557 -117,014 X X

Isaia et al., 2006 Italy Oceanic 44,900 8207 X

James et al., 2015 USA Mediterranean 38,879 -77,007 X

Kazakou et al., 2016 France Mediterranean 43,611 3877 X X X X

Kehinde & Samways, 2012 South Africa Mediterranean -33,000 18,000 X

Kehinde & Samways, 2014a South Africa Mediterranean -33,000 18,000 X X

Kehinde & Samways, 2014b South Africa Mediterranean -33,925 18,425 X

King & Berry, 2005 USA Mediterranean 38,700 -121,583 X

Klymenko, 2014 Ukraine Oceanic 44,600 33,533 X

Lee & Steenwerth, 2013 USA Mediterranean 43,450 -79,683 X

Marques et al., 2010 Spain Mediterranean 40,350 -3367 X X

Mercenaro et al., 2014 Italy Mediterranean 40,558 8322 X

Morlat & Jacquet. 2003 France Oceanic 47,350 -0.467 X

Muscas et al., 2017 Italy Mediterranean 40,558 8322 X X

Novara et al., 2011 Italy Mediterranean 37,655 13,015 X

Okur et al., 2015 Turkey Mediterranean 38,612 27,426 X X X

Ovalle et al., 2010 chili Mediterranean -35,967 -72,283 X

Paoletti et al., 1998 Italy Oceanic 44,413 12,201 X

Peregrina et al., 2012 Spain Oceanic 42,443 -2515 X

P´er`es et al., 2008 France Oceanic 46,083 4667 X X

P´erez -´Alvarez et al., 2013 Spain Oceanic 42,443 -2726 X X

P´erez -´Alvarez et al., 2015 Spain Oceanic 42,443 -2726 X X

Pou et al. (2011) Spain Mediterranean 39,650 -2800 X X

Probst et al., 2008 France Oceanic 47,950 7283 X X

Reeve et al., 2016 USA Mediterranean 45,245 -123,070 X X

Reinecke et al., 2008 South Africa Mediterranean -33,724 18,956 X

Rodriguez Lovelle et al., 2000 France Oceanic 44,838 -0.579 X

Ruiz-Colmenero et al. (2011) Spain Mediterranean 40,342 -3608 X X

Ruiz-Colmenero et al. (2013) Spain Mediterranean 40,342 -3608 X

Rusch et al., 2017 France Oceanic 44,840 -0.581 X

Salom´e et al., 2016 France Mediterranean 43,828 4563 X X

Sharley et al., 2008 Australia Conti-steppe -34,217 142,200 X

Smith et al., 2008 USA Mediterranean 36,240 -121,310 X

Steenwerth & Belina. 2008 USA Mediterranean 36,317 -121,233 X X

Steenwerth et al., 2013 USA Mediterranean 38,195 -121,150 X

Sweet & Schreiner. 2010 USA Mediterranean 43,936 -120,575 X X

Trigo-C´ordoba et al., 2015 Spain Mediterranean 42,360 -8117 X X

Virto et al., 2012 Spain Oceanic 42,823 -1.651 X X

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4. Discussion

In this study, we have shown how extensive vegetation management, particularly the presence of cover crops, favors biodiversity and the

supply of two ecosystem services that are important for the functioning of sustainable vineyards (i.e., supporting and regulating services). In addition, considered as a whole, extensive vegetation cover management had a minor, but non-significant, negative impact on provisioning Fig. 1. Representation of the geographical location of the study sites considered in this meta-analysis.

Fig. 2. Effects of vegetation cover (lnRR) on biodiversity and each of the ecosystem services (ES), with and without taking into account the irrigation variable. The values in brackets belong to number of observations. The points with error bars show the mean effect size and the 95% confi- dence interval. The error bars that do not cross the zero line indicate significant differences associated with vegetation cover compared to bare soil (asterisks are indicating the associated significance levels: + p < 0.1, * p < 0.05; **

p < 0.01; *** p < 0.001). NS = non-significant effects.

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Fig. 3. Relationships between the effects of vegetation cover (lnRR) on biodiversity and different ecosystem services (supporting and regulating) and edaphoclimatic variables. Rainfed vineyards = red dots. Irrigated vineyards = blue dots.

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services (i.e., grape yield), which is the ecosystem service that many winegrowers typically seek to maximize the most, particularly in the case of large operators and members of cooperatives (Pennerstorfer and Weiss, 2013). However, this negative effect was completely offset by irrigation. Our study, thus, indicates that the positive effects of extensive vegetation cover management on biodiversity and the rest of the ecosystem services outweigh the possible negative effects associated with the loss of grape production, particularly where irrigation is feasible. It has also been suggested that, in humid climates, cover crops can reduce excessive growth and vines and consequently reduce the time needed for expensive and time-consuming canopy thinning operations in the vineyards (Pou et al., 2011). Furthermore, there might addition- ally exist positive effects of extensive vegetation cover management for regulating the characteristics of grape juice and wines (Abad et al., 2021b), which may contribute to compensate potential yield losses.

Moreover, the effects of extensive vegetation cover management on biodiversity and ecosystem services also varied depending on the climatic conditions and edaphic characteristics of each location, suggest- ing the importance of developing strategies for the adaptive management of vegetation covers that are appropriate to the environmental conditions of each vineyard.

The use of cover crops has been widely described as promoting ecosystem services in a wide variety of agroecosystems, including vineyards (Garcia et al., 2018) (Winter et al., 2018). Such positive effects may be particularly enhanced when coupled with the well-established benefits obtained from the reduction of soil tillage, particularly in perennial woody crops; these benefits include less mixture of soil

horizons, greater amount of soil organic matter, greater cation exchange capacity, greater retention of nutrients, less soil compaction, and greater water infiltration and rooting depth (Garcia et al., 2018; Triplett and Dick, 2008). Cover crops may be composed of exotic or native species.

The composition of the seed mixture and the soil seed bank can thus also influence the provision of services; for example, native cover crops increased pest control in Australian vineyards (Danne et al., 2010) and arthropod biodiversity in South African vineyards (Eckert et al., 2020) as compared to exotic species. Grass-dominated cover crops have also been shown to reduce plant species richness across European vineyards in comparison to spontaneous vegetation or mixed cover crops (Hall et al., 2020), whereas aboveground biomass of inter-row vegetation was the main factor improving soil related ecosystem services in Spanish vineyards (Guzm´an et al., 2019). Cover crops consisting of communities of diverse herbaceous plants that produce flowers tend to have more positive effects on pollinators (Campos et al., 2017; Kratschmer et al., 2019, 2018). Consequently, the type, composition, and management of the cover crops in the inter-rows of vineyards can play an important role for the provision of ecosystem services and biodiversity, but could not be investigated here due to the lack of precise information. This aspect should thus be investigated in more detail, particularly under contrast- ing edaphoclimatic conditions.

Cover crops and extensive vegetation management can increase the supply of supporting and regulating ecosystem services through various mechanisms. For example, cover crops photosynthesize and transfer part of that carbon to the soil in the form of litter or as rhizodeposits, thus improving belowground trophic networks (Cheng and Baumgartner, Fig. 4. Structural equation models related to the response of biodiversity and ecosystem services in vineyards to extensive vegetation management. Solid black lines indicate positive associations, while dashed black lines indicate negative associations. +p < 0.1; * p < 0.05; * * p < 0.01; * ** p < 0.001. Gray lines indicate non- significant relationships. TWetQ = temperature of wettest quarter; PWetQ = precipitation of wettest quarter; PWarmQ = precipitation of warmest quarter; PSea

=precipitation seasonality; n = sample size.

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2004). This contributes to soil formation and water retention, improving nutrient cycling rates and, thus, enhancing the supply of both supporting and regulating ecosystem services. In addition, the interaction that takes place between the roots of grapevines and the roots of cover crops can help crops to combat abiotic stress and disease, increasing the exchange of essential nutrients such as nitrogen and phosphorus and, consequently, improving soil fertility (Celette and Gary, 2013; Cheng and Baumgartner, 2006). Such interactions can also result in the exchange of beneficial microbes between the roots of cover crops and vines (Vuki- cevich et al., 2016). Cover crops can also increase the structural stability of the soil by mitigating the direct impact of rain, improving infiltration and acting as a barrier against runoff and soil erosion (Burlotto, 2008;

Ruiz-Colmenero et al., 2013).

Despite the benefits mentioned above, it has also been shown that cover crops can compete with vines for soil resources such as water and nutrients, especially in climates with water limitations, such as for example Mediterranean climates in the summer season (Ward et al., 2012). This has led to a comparably low level of implementation of this type of soil management in areas with water deficit, which could addi- tionally be influenced by business structures of the wineries where grape producers are keener to maximize production and are thus less likely to maintain cover crops (Schütte and Bergmann, 2019). In our study, provisioning service (grape yield) was the only service that was compromised by the presence of cover crops, but this negative effect was completely cancelled out when irrigation was implemented. This un- derlines the importance of adequate water management for the successful implementation of cover crops in vineyards. However, irrigation may not always be possible, or even desirable. Hence, applying other sustainable practices that may help vines cope with the extra competition imposed by the presence of cover crops will be particularly important for a widespread adoption. These may include careful timing of cover crop management, as well as the inoculation with microbial biostimulants, including mycorrhizal strains, that may be able to supply extra water and nutrients, thus limiting the negative effects of competition (Abad et al., 2021a; Trouvelot et al., 2015).

Despite the consistent results found (this study and Winter et al., 2018), soil properties and the climate of each locality also had a modulating role of the effects of cover crops on ecosystem services in vineyards. Furthermore, the modulating role of environmental variables varied for each type of ecosystem service. In our study, soil pH was one of the main variables that acted as a modulator of this response. This coincides with the key role of soil pH as a regulator of nutrient availability to plants and of the composition and activity of soil microbial communities (Fierer and Jackson, 2006; Ramos Vázquez and Zúñiga Dávila, 2008). Also consistent with the key role of pH in agriculture, management practices aimed at correcting the soil pH level by adding calcium and magnesium salts have been long used in order to create an ideal environment for the development of plants and soil organisms (Quiroga et al., 2017). Specifically, regulating ecosystem services were more favored in alkaline soils. We speculate that this could be attributed to the fact that, under alkaline conditions, soil activity and biota abundance are often constrained, and the presence of cover crops may mitigate this possibly by acidifying the soil in the inter-rows and by creating micro-environmental conditions that are more conducive for soil organisms to thrive. In contrast, supporting ecosystem services were reduced at high pH levels, but this effect was only evident in the SEM.

Moreover, soil carbon sequestration and pest control did not show evidence of a link with pH, indicating that the negative effects of pH on supporting services is the result of considering several supporting services together. Our results, thus, suggest that soils with neutral pH may show a more balanced response in their supply of supporting and regulating services in the presence of cover crops.

In addition to pH, climate played a very important role as a regulator of ecosystem services in the presence of cover crops. The two most important climatic variables were, on one hand, temperature of the wettest and precipitation of the warmest quarter, broadly representing

the specific conditions during the growing season, and precipitation of the wettest quarter, most likely linked to water scarcity. In particular, lower temperatures of the wettest quarter favored regulating services.

This may be due to the lower development of pests, including fungal diseases, which are favored in conditions of higher temperature and humidity (Bois et al., 2017; Delp, 1954). On the other hand, supporting services were favored by a higher precipitation of the wettest quarter, which is probably associated with more favorable conditions for soil biota due to higher soil moisture. Actually, the abundance and activity of soil organisms is known to be tightly linked to water availability.

Similarly, cover crops may also particularly benefit from greater precipitation during the wet season, which may indirectly also affect soil biota abundance and activity and, hence, the provision of supporting services. Biodiversity was, in turn, favored in vineyards with lower precipitation seasonality, which occurred in locations closer to the equator. These conditions may imply more favorable conditions for a wider range of species across trophic levels, thus magnifying the positive effects of cover crops on biodiversity. Understanding how the climate regulates the effects of cover crops on ecosystem services in vineyards is key in a context of climate change in which historically important wine growing areas are becoming increasingly arid (Van Leeuwen et al., 2019), whereas areas that were previously not suitable for quality wine production could gain importance; for example, southern parts of En- gland and Sweden (Nicholas, 2015). It is thus likely that the importance of a flexible adaptive management of cover crops that are well-adapted to various local edaphoclimatic conditions will become increasingly more important in the face of climate change. Our study suggests that considering climatic variables that capture conditions during the growing season or climatic harshness, particularly in terms of rainfall, may be particularly relevant.

The magnitude by which cover crops increased regulating and supporting services as well as biodiversity was much greater than the negative effect that they had on provisioning services. However, it is important to take into account the usual farmer’s vision of obtaining the maximum economic performance from the crop, in this case the vineyard. Therefore, any management practice that may cause a minimal negative economic impact, or an uncertain positive impact, is not likely to be accepted, and thus implemented, by growers, especially if they sell their grapes to cooperatives or larger producers (Schütte and Bergmann, 2019). For this reason, it is extremely important to establish locally-adapted economic incentives that favor the use of cover crops to offset potential yield losses or increased costs and at the same time contribute to the overall supply of ecosystem services and biodiversity.

Some of these incentives may be targeted at allowing the implementation of potentially costly irrigation systems that would only need to be activated during specific periods of the year, when competition among vines and cover crops is highest. These incentives could also be targeted at the implementation of other management measures that minimize the impact of competition by cover crops on vines, including the addition of microbial biostimulants formulated to increase the tolerance of vines to drought or to increase its ability to capture water and nutrients. Among the most promising biostimulants that could aid in this, mycorrhizal inoculants and Trichoderma are notorious.

5. Conclusions

Taken together, we have shown that implementing cover crops in the inter-rows of vineyards provides a positive net effect on ecosystem services. The most favored variables were those linked to biodiversity, supporting services in irrigated vineyards and regulating services in rainfed vineyards, while provisioning services were slightly negatively affected. For this reason, we believe that it is of utmost importance to establish locally adapted agricultural policies that promote the use of cover crops. On the other hand, we demonstrated the importance of climate, and soil variables in modifying the effects of cover crops in vineyards, particularly pH and climatic variables linked to conditions

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during the growing season or climatic harshness. Irrigation was also key to mitigate the negative effects of cover crops on grape yield. Of course, we acknowledge that our study is based on modelled climatic data that may not be able to capture the climatic conditions during the study period of the specific vineyards, and thus should be interpreted with care. Nevertheless, we believe that this article provides some of the first evidence of the importance of considering among-site differences in terms of soil and climatic conditions when understanding the benefits, and also potential drawbacks, of the implementation of cover crops in vineyards. Taken together, our results, which are extremely important in the current context of climate change, widespread environmental degradation, and greener environmental legislation, emphasize the importance of a flexible, locally adapted management of the vine-soil- cover crop interaction for the future sustainability of vineyards.

Declaration of Competing Interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

Data availability

Data will be made available on request.

Acknowledgements

R.O.-H. is financially supported by the Ramón y Cajal program from the Ministry of Science and Innovation (RYC-2017 22032) and projects (Plan Andaluz de Investigación, Desarrollo e Innovación (2020 [Ref. 20_00323], Spanish State Research Agency [PID2019-106004RA- I00/AEI/10.13039/501100011033], and Junta de Andalucía [GOPC- CA-20-0001]). S.W. was financed through the research project SECBI- VIT, which was funded through the 2017–2018 Belmont Forum and BiodivERsA joint call for research proposals, under the BiodivScen ERA- Net COFUND programme, with the funding organisations: Spanish State Research Agency/Spain, BMBF/Germany, ANR/France, NWO/

Netherlands, UEFISCDI/Romania, FWF/Austria (Grant #I 4025-B32) and the NSF/USA (Grant #1850943).

Appendix A. Supporting information

Supplementary data associated with this article can be found in the online version at doi:10.1016/j.agee.2022.108115.

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