Exp Appl Acarol (2006) 40:175–188 DOI 10.1007/s10493-006-9044-z
EVects of agroforestry on phytoseiid mite communities (Acari: Phytoseiidae) in vineyards in the South of France Ziad Barbar · Marie-Stéphane Tixier · Brigitte Cheval · Serge Kreiter
Received: 19 April 2006 / Accepted: 24 November 2006 / Published online: 20 January 2007 © Springer Science+Business Media B.V. 2007
Abstract The abundance and diversity of phytoseiid mites were surveyed from April to September 2003 to 2005 in vineyards (Grenache and Syrah cultivars) coplanted with rows of Sorbus domestica or Pinus pinea and in monoculture plots of grapes in the South of France. Densities of phytoseiid mites were diVerent on the two tree species, with P. pinea a more suitable host than S. domestica. Typhlodromus (Typhlodromus) exhilaratus was the dominant species occurring on grapes and on co-planted rows of S. domestica and P. pinea, whereas T. (T.) phialatus was the most abundant species in monoculture plots of both S. domestica and P. pinea. Factors determining the dominance of T. (T.) phialatus over T. (T.) exhilaratus in monoculture trees are discussed. In this study, agroforestry management did not aVect phytoseiid diversity in vineyards, but did aVect phytoseiid density, especially in 2005. The results obtained in 2003 and 2004 are not easy to discuss in this regard because of the low densities of mites observed during these 2 years (very dry climatic conditions and pesticide applications). Keywords Phytoseiidae · Typhlodromus exhilaratus · Typhlodromus phialatus · Vineyards · Pinus pinea · Sorbus domestica
Introduction Management of vegetational diversity in uncultivated areas surrounding crops or inside Welds may increase natural enemy occurrence (density and diversity) and ultimately limit pesticide applications (Altieri and Letourneau 1982; Lozzia and Rigamonti 1998; Escudero and Ferragut 1999; Zacarias and Moraes 2002).
Z. Barbar (&) · M.-S. Tixier · B. Cheval · S. Kreiter Laboratoire d’Acarologie, ENSAM – INRA, Unité d’Ecologie animale et de Zoologie agricole, 2 Place Pierre Viala, 34060 Montpellier cedex 1, France e-mail:
[email protected]
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Predatory mites belonging to the family Phytoseiidae are well known because of the eYcacy of some species in controlling pest mites on many crops, including grape (McMurtry and Croft 1997). High densities of these predators have often been reported on native or weedy vegetation neighbouring vineyards (Boller et al. 1988; Tsolakis et al. 1997; Kreiter et al. 2000, 2006; Tixier et al. 2000a, b; Duso et al. 2004; Barbar et al. 2005) and also on herbaceous and Xowering plants (wild or cultivated) in vineyards and apple orchards (Flaherty 1969; Coli et al. 1994; Nicholls et al. 2001). Findings such as these have been discussed as an application of conservation biological control. Another possibility of enhancing natural enemy occurrence in cropping systems is agroforestry, the combination of trees and/or shrubs with crops. This type of cropping system has been found to modify microclimatic conditions, and enhance alternative habitats and food resources for natural enemies (Stamps and Linit 1998; Altieri and Nicholls 2002). However, to date no published studies exist to show the impact of agroforestry management on predatory mite communities in vineyards. This paper presents the results of a study on phytoseiid mite communities in vineyards co-planted with tree species chosen for their high commercial wood value: Sorbus domestica L. and Pinus pinea L. The hypothesis tested is that vineyards in an agroforestry system would have higher mite abundance and diversity than vineyards in monoculture.
Materials and methods The study site The experiments were carried out in grape vineyards located in Restinclières (15 km North of Montpellier, Hérault, France, the area has a sub-humid Mediterranean climate). The vineyards contained two cultivars: Syrah and Grenache, which were planted in 1997 on a reclaimed fallow (30-years-old) (Table 1). Syrah has a hirsute leaf undersurface, whereas Grenache has a glabrous leaf. In the same year, rows of S. domestica and P. pinea were planted within the vineyards. The cropland was surrounded by uncultivated areas mainly composed of Pinus halepensis Viller and Quercus coccifera L. Eight to ten phytosanitary treatments a year (fungicides against powdery and downy mildews, plus insecticides against Scaphoideus titanus Ball) were applied to all grape crops (Table 2). The pesticides used were selected to minimise their side
Table 1 Characteristics of the sampled plots in the study site of Restinclières (Hérault–France)
Surface Plantation density Rows of Syrah cv Rows of Grenache cv Rows of S. domestica Rows of P. pinea
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Experimental Experimental Monoculture Monoculture grape crop I grape crop II grape crop P. pinea
Monoculture S. domestica
4600 m² 2.5 £ 1 m 11 9 4 –
2500 m² 4£3m – – 5 –
3900 m² 2.5 £ 1 m 10 12 – 5
1000 m² 2.5 £ 1 m 9 11 – –
3400 m² 3£2m – – – 12
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Table 2 Active ingredients of pesticides used in grape crops in Restinclières (Hérault–France) from 2003 to 2005 Diseases and pests sprayed
Active ingredients used
Date of spraying
Number of Applications/year
Dose/ha
Downy mildew
Fosetyl aluminium 50% and Folpel 50%
2003 2004
May–June May
2 2
2.9–3.5 kg
Powdery mildew
Scaphoideus titanus
2005
May–June
2
Azoxystrobine
2003 2004
June June
1 1
2.4 l
Dimethomorph 9% + Mancozeb 60%
2004 2005
July July
1 1
2.5 kg
Difenoconazol
2003 2004
June June
2 2
0.14 l
Quinoxyfen
2003 2005
June–July May
2 1
0.21 l
Sulfur Lambda–cyhalothrin Chloropyrifos–ethyl
2005 2003 2004 2005
July June–July June–July June–July
1 3 3 3
6 kg 0.14 l 1.5 l
eVects on phytoseiid mites whenever possible. No acaricide was used either prior to or during the 3-year study. Because of the close proximity of the co-planted rows of S. domestica and P. pinea, these too received the same pesticide applications. In contrast, no pesticides were applied to the monoculture tree crops (S. domestica and P. pinea). Two kinds of plots were included in the studies: (1) vineyard plots comprising coplanted rows of S. domestica (grape crop I) or of P. pinea (grape crop II), (2) control plots comprising monoculture grape, monoculture S. domestica or monoculture P. pinea. All of them were located within an area of less than 5 km2 (Table 1; Fig. 1). Sampling Sampling was conducted Wve, six and three times a year (from April to September) in 2003, 2004, and 2005, respectively. The sampling unit for grape and S. domestica was a leaf, and for P. pinea a 10 cm shoot, all of them randomly collected on the plants. On each date, the following samples were randomly collected: (1) In grape crops I and II, 30 samples of each crop per row (Table 1), keeping the two vine cultivars separate (30 leaves of each vine cultivar). (2) In the monoculture tree plots, sixty leaves per plot of grape S. domestica, and P. pinea were also randomly sampled. The leaves were put in plastic bags, and brought back to the laboratory on ice. Phytoseiids were counted and removed from each grape and S. domestica leaf using a binocular microscope at 40£ magniWcation. As it was diYcult to directly remove mites from the shoots of P. pinea, mite extraction was undertaken using the “soaking-checking-washing-Wltering” method (Boller 1984). Only mobile stages of mites were taken into account. In all cases, each collected leaf or shoot of grape, S. domestica or P. pinea was considered as a replicate for statistical analysis. To compare densities of phytoseiid mites found on trees and on vines, leaves of S. domestica, P. pinea and vine were considered equivalent.
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Fig. 1 Experimental and monoculture grape crops and monoculture plots of P. pinea and S. domestica in Restinclières (Hérault–France)
Mite identiWcations All mite samples were mounted in Hoyer’s medium on slides and identiWed using a phase and interferential contrast microscope, according to the taxonomic keys of the generic revisions of Chant and McMurtry (1994) for Typhlodrominae and Phytoseiinae, and those of Chant and McMurtry (2003a, b, 2004) for Ambyseiinae Amblyseiini, Neoseiulini and Kampimodromini. The slides were kept in the mite collection of the Laboratoire d’Acarologie of Montpellier and data computerised in the corresponding database. Data analysis Variance analysis (Kruskal–Wallis test), followed by a Newman–Keuls mean comparison test ( = 5%) (Statistica® version 7.1, 2005) were carried out to compare (1)
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mean mite density (all species) found on co-planted trees and in monoculture plots of S. domestica and P. pinea; (2) mean densities of Typhlodromus (Typhlodromus) exhilaratus Ragusa and Typhlodromus (Typhlodromus) phialatus Athias-Henriot found on S. domestica and P. pinea co-planted with grape or in monoculture; (3) mean mite density on the two grape cultivars; (4) mean mite density on the vines in grape crops I and II (agroforestry) and in the grape monoculture; (5) mean mite density on S. domestica, P. pinea and grape.
Results Phytoseiid mite diversity Seven phytoseiid mite species were found, two being dominant. In all the grape crops (agroforestry or monoculture), T. exhilaratus was the prevailing species (more than 98%) (Fig. 2). The other species (T. phialatus, Paraseiulus triporus [Chant & Yoshida-Shaul] and Typhlodromus [Typhlodromus] pyri Scheuten) were only observed on grapes in 2003. Five species were found on co-planted P. pinea: T. exhilaratus was the prevalent one (87.0%) followed by T. phialatus (12.1%). Other species (Typhlodromus [Anthoseius] recki [Wainstein], Kampimodromus aberrans [Oudemans] and Neoseiulus bicaudus [Wainstein]) were observed sporadically. In the monoculture plot of P. pinea, the main species was T. phialatus (96.9%). Typhlodromus exhilaratus was the only species observed on co-planted S. domestica, whereas T. phialatus was the prevalent species in the monoculture plot (66.7%).
Typhlodromus exhilaratus
Typhlodromus phialatus
Others species
100%
80%
60%
40%
20%
0% Vine crop I
Co-planted Vine crop II Co-planted S. domestica P. pinea
Monoculture Monoculture Monoculture vine crop S. domestica P. pinea
Fig. 2 Percentage of phytoseiid mite species in the sampled plots in the study site of Restinclières (Hérault–France)
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Phytoseiid mite densities on S. domestica and P. pinea Sorbus domestica Overall mite density was very low during the three study years, both for co-planted or monoculture trees. Abundance was, however, slightly higher in monoculture [H(1,2520) = 4.04; P = 0.0443], but this was due to the data of May 2003 [H(4,900) = 19.68; P = 0.026] and July 2004 [H(5,1080) = 64.97; P = 0.003] (Fig. 3). No signiWcant diVerence [H(1,540) = 0.468; P = 0.493] was observed between the two cropping systems in 2005 (Fig. 3). Of the two dominant species, T. exhilaratus densities were not signiWcantly diVerent between the two cropping systems [3-year data set: H(1,2520) = 2.47; P = 0.116; 2003: [H(1,900) = 0.41; P = 0.522]; 2004: [H(1,1080) = 1.43; P = 0.23]; 2005: [H(1,540) = 2.37; P = 0.120]) (Fig. 4), wheras T. phialatus was never observed on co-planted S. domestica. Pinus pinea The highest densities of mites were observed in the monoculture in regard to co-planted trees [3-year data set: H(1,2940) = 86.87; P < 0.001; 2003: H(1,1050) = 15.87; P = 0.001]; 2004: [H(1,1260) = 112.06; P < 0.001], and 2005: [H(1,630) = 12.73; P < 0.0001] (Fig. 5). However, this only concerns T. phialatus [3-year data set: H(1,2940) = 296.59;
Fig. 3 Mean number of phytoseiid mites per leaf and § SE in the two S. domestica modalities in Restinclières (Hérault, France) from 2003 to 2005 (— co-planted S. domestica; ---- monoculture S. domestica plot)
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Fig. 4 Mean numbers of Typhlodromus exhilaratus per leaf and § SE. in P. pinea and S. domestica (co-planted or not with vine) in Restinclières (Hérault, France) from 2003 to 2005
Fig. 5 Mean numbers of phytoseiid mites per leaf and § SE. in the two P. pinea modalities in Restinclières (Hérault, France) from 2003 to 2005 (— co-planted P. pinea; ---- monoculture P. pinea plot)
P < 0.001. Separate years-2003: H(1,1050) = 104.83; P < 0.001; 2004: H(1,1260) = 151.71; P < 0.001; 2005: H(1,630) = 67.97; P < 0.0001] (Fig. 6). In contrast, densities of T. exhilaratus were signiWcantly higher in co-planted P. pinea than in the monoculture plot [3-year data set: H(1,2940) = 67.01; P < 0.0001. Separate years-2003: H(1,1050) = 51.21; P < 0.0001; 2004: H(1,1260) = 18.82; P < 0.0001; 2005: H(1,630) = 3.88; P = 0.048] (Fig. 4). The same variations in time were observed for the co-planted trees and trees in the monoculture plot, and diVered for the three years. In 2003 and 2005, the highest
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Fig. 6 Mean numbers of Typhlodromus phialatus per leaf and § SE in P. pinea and S. domestica (co-planted or not with vine) in Restinclières (Hérault, France) from 2003 to 2005
densities were observed early in the season, whereas in 2004, they were observed in July. Phytoseiid mite abundance on grape As T. exhilaratus was the dominant species (more than 98%) in all grape plots during three years, comparisons will only concern the densities of this species. In all the grape crops (agroforestry or monoculture), mite densities were signiWcantly higher on Syrah than on Grenache [H(1,18480) = 98.78; P < 0.001]. The highest densities were observed in 2005 (mean per leaf: 0.22 § 0.01) and the lowest in 2004 (0.06 § 0.004) (Table 3). No signiWcant diVerence was observed between mite densities found in agroforestry grape and monoculture grape (Table 3) [H(2,18480) = 6.05; P > 0.05] when data from the three years were pooled. However, for each year diVerences existed. In 2003, mite abundance did not signiWcantly diVer between grape crop I and monoculture grape, but a signiWcant diVerence [H(2,6900) = 141.97; P < 0.001] (Fig. 7) was observed between these two latter plots and grape crop II where the lowest densities were observed. In 2004, phytoseiid densities were signiWcantly higher in grape crop II than in the others [H(2,7920) = 47.78; P < 0.0001] (Fig. 7). In 2005, densities were signiWcantly diVerent in all three grape crops [H(2,3960) = 15.28; P = 0.0005], higher densities being observed in agroforestry grape (I and II) (Fig. 7). Density variation in time was diVerent for each year, but equivalent for the three plots considered. In 2003, the highest densities were observed in April and May [H(4,6600) = 854.24; P < 0.001] (Fig. 7), whereas in 2004, densities were very low early in the season [H(5,7920) = 228.289; P < 0.001]. In 2005, the highest densities were found in September [H(2,3960) = 273.28; P < 0.001]. Phytoseiid mite densities on P. pinea, S. domestica and grape The highest mite densities were found in the monoculture plot of P. pinea during 3 years [H(5,23940) = 223.41; P < 0.001] (Fig. 8) followed by these observed on coplanted P. pinea, Syrah and Grenache. The lowest densities of phytoseiid mites were observed on monoculture and co-planted trees of S. domestica.
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Table 3 Results of variance analysis and mean comparison test ( = 5%) on phytoseiid mites densities in sampled grape crops in Restinclières (Hérault–France) (means sharing a common letter are not signiWcantly diVerent) Comparison
Kruskal–Wallis test
Newman–Keuls-Mean
Mite density
Sampling date
P < 0.001
2003 2004
0.10 § 0.004 b 0.06 § 0.004 c
2005
0.22 § 0.010 a
Grape cultivars
P < 0.001
Grenache Syrah
0.07 § 0.003 b 0.14 § 0.005 a
Grape crops
P > 0.05
Experimental grape crop I Experimental grape crop II
0.11 § 0.005 a 0.10 § 0.004 a
Monoculture grape crop
0.10 § 0.004 a
Fig. 7 Mean number of phytoseiid mites per leaf and § SE in the diVerent grape crop modalities in Restinclières (Hérault, France) from 2003 to 2005 (— grape crop I; ---- grape crop II; .... monoculture grape crop)
Comparison of phytoseiid mite densities on Sorbus domestica and Pinus pinea For all the sample dates, the densities of T. exhilaratus were higher on co-planted P. pinea than on co-planted S. domestica and on monoculture plots of P. pinea and S. domestica [H(3,5460) = 162.43; P < 0.001]. However, the densities of T. phialatus in the monoculture plot of P. pinea were higher than on co-planted P. pinea and in monoculture plots of S. domestica [H(3,5460) = 567.93; P < 0.001].
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Fig. 8 Mean numbers of phytoseiid mites per leaf and §SE in the diVerent modalities of S. domestica, P. pinea and grape cultivars in Restinclières (Hérault, France) during 3 years (CS and MS: co-planted and monoculture S. domestica consecutively; CP and MP: coplanted and monoculture P. pinea, G and S: Grenache and Syrah cvs.)
Discussion Phytoseiid mite diversity in grape crops and on S. domestica and P. pinea Sorbus domestica and P. pinea in monoculture or co-planted with grape shelter phytoseiid mites. However, diVerent species dominated among the cropping systems: T. exhilaratus on co-planted and T. phialatus in monoculture. Typhlodromus exhilaratus was also the prevailing species on grape, whereas T. phialatus was dominant in uncultivated areas surrounding the experimental plots (Barbar et al. 2005). Furthermore, previous studies at the same site showed a dispersal of these two species in the vine plot (Tixier et al. 2006). InterspeciWc predation between T. exhilaratus and T. phialatus could aVect the establishment of these species in the grape crops. Meszaros et al. (2006) showed in lab experiments that adult females of T. exhilaratus could feed on larvae and protonymphs of T. phialatus and had a greater fecundity than T. phialatus. However, these characteristics are not suYcient to explain the spatial segregation between T. phialatus and T. exhilaratus, and especially the dominance of T. phialatus in monoculture tree plots or in uncultivated areas surrounding the grape crops. Thus, other factors are certainly involved. Pesticides could aVect the development of these two species (Castagnoli and Liguori 1987; Grande and Ingrassia 1988; Rodrigues et al. 2002). DiVerences in suceptibility between the two species to the pesticides applied would perhaps explain the spatial segregation observed. In Spain and Portugal, authors have reported high densities of T. phialatus in vineyards poorly sprayed or unsprayed with insecticides (Garcia-Mari et al. 1987; Pereira et al. 2003). However, no comparative study on pesticide eVects on these two species exists. Therefore, in a separate study, we studied the side eVects of two pesticides (a fungicide and an insecticide) on populations of T. phialatus and T. exhilaratus which suggests a negative eVect of these pesticides on phytoseiids and especially on T. phialatus (Barbar et al. submitted). Phytoseiid mite densities in grape crops Mite densities were diVerent between the two vine cultivars, the highest densities being observed on the most hirsute leaf cultivar, which agrees with previous results
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(Karban et al. 1995; Tixier et al. 1998, 2006; Duso and Vettorazzo 1999). Many studies have shown the close relationships between phytoseiid mite species and their leaf substrate. A hirsute leaf with domatia aVects leaf hygrometry and also food occurrence (especially pollen) (O’Dowd and Willson 1989; Karban et al. 1995; Walter 1992, 1996; Kreiter et al. 2002). However, more studies are clearly needed to better characterise the links between vine cultivar characteristics and phytoseiid density and diversity. Phytoseiid abundance in the grape agroforestry crops was sometimes higher compared to the monoculture grape crop. However, this observation cannot be generalised. During 2003 and 2004, mite abundance in grape crops did not seem to be aVected by the presence of the two tree species, but the densities of phytoseiids were very low. In 2003, particularly dry weather conditions and use of an insecticide (“lambda-cyhalothrin”) (applied in the context of the obligatory control of S. titanus in French vineyards; Auger, personal communication) may have aVected densities of phytoseiids in grape crops and on co-planted trees, and may have had a consecutive eVect in 2004. For these reasons, the 2003 and 2004 data are very diYcult to discuss in regard to the agroforestry eVect, and if an eVect does exist, it may be hidden by high mortality due to the dry conditions and side eVects of the pesticides. In 2005, phytoseiid population density increased and a positive eVect of agroforestry on mite densities on grape was observed, no matter what the species of co-planted tree. In this case, trees may have been acting as reservoirs for these predators, providing alternative food and refuge, as several studies have shown for plants surrounding vineyards (Boller et al. 1988; Tsolakis et al. 1997; Kreiter et al. 2000; Tixier et al. 2000a). However, further studies are needed to conWrm these trends and to determine the mechanisms leading to a greater abundance due to the presence of the trees. Molecular typing of populations of T. exhilaratus on co-planted trees versus vines are planned in order to determine the degree of mite migration between the two habitats. The management of S. domestica did not show a strong eVect on phytoseiid densities, while monoculture P. pinea was colonised by higher phytoseiid densities. In this latter case, pesticide application could have aVected mite densities. However, if we compare the densities of T. exhilaratus, the prevalent species in grape crops, S. domestica (monoculture or co-planted) sheltered the same densities of T. exhilaratus, and densities were higher on co-planted P. pinea than in monoculture. This seems to show the low eVect of pesticide application of T. exhilaratus densities on coplanted trees. The lowest densities on monoculture P. pinea could be due to competition with T. phialatus. Phytoseiid mite densities on S. domestica and P. pinea Higher densities of mites and a higher frequency in time were observed on P. pinea than on S. domestica (monoculture or co-planted). Pinus pinea seems to be a more suitable host plant than S. domestica between the two types of plant management and for the two diVerent phytoseiid species encountered. Factors such as food availability could aVect mite densities (Zacarias and Moraes 2002). Phytophagous mites (Tetranychidae, Tenuipalpidae, and Eriophyidae) have been observed on these two host plants (Barbar, unpublished data). It can be hypothesised that diVerent density and diversity of prey (especially high densities of Tenuipalpidae on P. pinea) could have a diVerent impact on phytoseiid mite development on S. domestica and P. pinea
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(Kreiter et al. 1993; McMurtry and Croft 1997; Duso and Vettorazzo 1999). More speciWc studies are required to test this hypothesis and to draw conclusions about the impact of food on phytoseiid densities on S. domestica and P. pinea. Even though phytoseiids were found on the co-planted trees, one wonders if these trees are really a better source of phytoseiids than the grapes themselves. Densities of phytoseiids observed on co-planted P. pinea were similar to those observed on Syrah, whereas densities observed on co-planted S. domestica were much lower. All factors being equal, a row of P. pinea would act much the same as a row of Syrah, providing equivalent mite densities. However, volumes of canopies (numbers of leaves, height of trees, plantation densities) of grape and tree rows are diVerent. To make a true comparison of mite abundance between vine and tree rows, some sort of biomass index between the diVerent crop types would be required. In conclusion, plantation of trees within grape vineyards seems to aVect mite density, noted especially in 2005, when mite densities were high and when the negative impacts of external factors were low (e.g., temperature, relative humidity and pesticide applications). However, these trends have to be conWrmed by additional experiments. Furthermore, agroforestry management does not aVect phytoseiid mite diversity in grape vineyards. Nevertheless, co-planted trees could constitute a reservoir for these predators. Exchanges between vine and trees of populations of the same dominant species could occur in a way that would enhance biological control of pest mites of adjacent crops. These hypotheses, however, have to be conWrmed by further experiments, including molecular typing of populations of T. exhilaratus on vine and on co-planted trees, in order to better quantify the rate of exchange between crop types. Acknowledgements We thank the conseil général de l’hérault for Wnancial contribution. We thank also Mr. Thierry Vacher, the vineyard manager. This study is included within the granted project: “Programme de Recherche Intégrée en AgroforesTerie (PIRAT)”, 1999–2007. We also kindly thanks Michael Costello for english improvement of the paper.
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