Short-Term Effects of Conversion To Direct Seeding Mulch-Based Cropping Systems On Macro-Fauna and Weed Dynamics

Journal of Crop Improvement
ISSN: 1542-7528 (Print) 1542-7536 (Online) Journal homepage: http://www.tandfonline.com/loi/wcim20
Short-Term Effects of Conversion to Direct Seeding

Mulch-Based Cropping Systems on Macro-Fauna
and Weed Dynamics
Rmy Kulagowski, Laura Riggi & Anas Chailleux
To cite this article: Rmy Kulagowski, Laura Riggi & Anas Chailleux (2016) Short-Term Effects
of Conversion to Direct Seeding Mulch-Based Cropping Systems on Macro-Fauna and Weed
Dynamics, Journal of Crop Improvement, 30:1, 65-83
To link to this article: http://dx.doi.org/10.1080/15427528.2015.1113222
Published online: 09 Feb 2016.
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Date: 10 February 2016, At: 02:42
JOURNAL OF CROP IMPROVEMENT

2016, VOL. 30, NO. 1, 6583
http://dx.doi.org/10.1080/15427528.2015.1113222
Short-Term Effects of Conversion to Direct Seeding

Mulch-Based Cropping Systems on Macro-Fauna and Weed
Dynamics
Rmy Kulagowskia, Laura Riggib, and Anas Chailleuxc
Downloaded by [Rmy Kulagowski] at 02:42 10 February 2016
a
Chamber of Agriculture of Alpes de Haute Provence, Oraison, France; bDepartment of Ecology, Swedish
University of Agricultural Sciences, Uppsala, Sweden; cCIRAD, UPR HortSys, Montpellier, France
ABSTRACT
ARTICLE HISTORY
Agroecosystem biodiversity could provide essential services

such as pest control. One approach currently used to promote
ecosystem services in agricultural systems is to reduce tillage
and increase plant diversity. In this study, we assessed the
short-term effects of conversion from reduced tillage (RT) to
direct seeding mulch-based cropping systems (DMC) on the
dynamics of arthropods (detritivores and predators), and major
pests (slugs and weeds). The study was conducted in two
commercial fields: one cropped with sorghum (Sorghum bicolor
L.) and one with maize (Zea mays L.). We found that both
beneficial and detrimental groups monitored were more abundant in DMC than in RT treatment and that the dominant
species differed between treatments. Because of their major
role in agroecosystems by contributing to the control of weed
seeds, insects, and slugs, carabid beetles (Carabidae) were
investigated in greater detail, and the results showed their
diversity was also higher in DMC than in RT. The dominant
species found were Poecillus cupreus and Pseudofonus rufipes in
the maize and sorghum fields, respectively. The increase in
biological control agents shortly after conversion suggested
that cover crops should be considered as a pest management
tool, even on a short-term scale.
Received 1 September 2015

Accepted 23 October 2015
KEYWORDS
Adoption; biological control;

conservation agriculture;
ecosystem services; maize;
sorghum
Introduction
Biodiversity underpins many ecosystem processes; hence increased biodiversity in agroecosystems could provide important ecosystem services for farmers (Altieri 1999; Moonen and Barberi 2008). While many studies have dealt
extensively with the relationship between diversity and ecosystem services in
natural ecosystems, few have focused on this relationship in agricultural
ecosystems. Increasingly, research suggests that the level of natural regulation
in agroecosystems is largely dependent on the level of plant and animal
biodiversity present (Altieri 1999; Ratnadass et al. 2012). However, changes
in food demand, conversion to modern, high-input agriculture, land-use
CONTACT Rmy Kulagowski
kulagowski.remy@gmail.com
Provence, Avenue Charles Richaud, 04700 Oraison, France.
2016 Taylor & Francis
Chamber of Agriculture of Alpes de Haute
66
R. KULAGOWSKI ET AL.
changes, and the globalization of agricultural markets have caused rapid

agricultural biodiversity loss. In particular, crop-management practices have
been shown to directly affect the stability and functioning of agroecosystems
through their impacts on functional biodiversity, potentially disrupting
trophic webs (Grime 1998; Duyck et al. 2011). Intercropping, agroforestry,
shifting cultivation, and conservation agriculture are examples of methods
that aim at maintaining biodiversity and enhancing the sustainability and
autonomy of agroecosystems (Malzieux 2012; Chailleux et al. 2014).
Conservation tillage is currently promoted to sustainably improve soil
quality. It involves soil-management practices that minimize disruption of
the soil structure, composition, and natural biodiversity, thereby reducing
erosion and degradation. Soil tillage adversely affects soil macro-fauna
because of direct mortality or as a result of indirect losses via dispersal
caused by habitat deterioration (Shearin et al. 2007). The two main concerns
regarding conservation tillage are increases in slug and weed populations. In
many parts of the world, slugs are serious pests of cereals, oilseeds, protein,
and vegetable crops (Godan 1983; South 1992; Barker 2002), but were
unknown as major pests until conservation-tillage practices were adopted
along with changes in cropping patterns (Stinner and House 1990; Glen
2002). Conservation tillage could also increase weed infestation (Phillips
et al. 1980; Hinkle 1983; Koskinen and McWhorter 1986) and alter the
species composition, favoring perennials and annuals (mostly grasses) that
do not require seed burial (Chancellor and Froud-Williams 1986).
As conservative soil management plays a major role in maintaining biodiversity in agricultural fields (Brussaard et al. 2007), it should be regarded as
a tool to improve ecosystem services in pest management. Many different
soil-conservation practices exist, from reduced tillage (RT) to direct seeding
mulch-based cropping systems (DMC), with variable effects on soil quality
and biodiversity. In conservation-tillage practices, cover crops can also be
used to avoid soil erosion (Langdale et al. 1991) and affect soil quality and
humidity. Hence, conservation practices impact biodiversity through (i)
reduced soil disruption and (ii) cover-crop introduction (Landis et al. 2000;
Ratnadass et al. 2012).
Generalist predators, such as carabids, staphylinids, and spiders, have been
shown to provide important natural biological control services in agroecosystems (Lundgren et al. 2006; Northfield et al. 2012). However, their effectiveness in controlling pests is negatively affected by intensive crop
management, such as tillage (Kromp 1999). In Europe, conservation tillage
is a recent practice as compared with North and South America (Holland
2004). Most studies assessing the impact of this practice on natural enemies
and pests have therefore been carried out in North America (Allen 1979),
and field studies concerning the impacts of different soil conservation practices on macro-fauna and potential ecosystem services are lacking in Europe
67
(Kromp 1999). Therefore, we carried out on-farm experiments in southern

France on maize and sorghum fields to assess the short-term effects of DMC
adoption in fields managed through RT. We focused on RT to DMC conversion because farmers more easily adopt RT that does not require a drastic
change in habits, contrary to DMC that is a new soil-management strategy
(Lahmar 2010; Scopel et al. 2013). Thus, conversion generally occurs step by
step, with a first conversion from plowing to RT and a second from RT to
DMC, which requires accurate knowledge of the ecological processes
(Kulagowski and Chailleux 2015). This study (i) assessed the seasonal
dynamics of the aboveground arthropod community and of the major pests
(i.e., slugs and weeds) on maize and sorghum crops, and (ii) involved a
detailed analysis of carabid beetle diversity and abundance. Our aim was to
evaluate the impact of soil practices when fields are managed by farmers
using their regular practices. The first objective was to obtain data for further
improvement of aboveground arthropod-mediated ecosystem services in
arable fields, and the second was to assess any benefits of DMC adoption
on a short-term scale.
Materials and methods

Study site and crop management description
This study was conducted on two commercial fields of two farms located in
the same catchment basin (latitude 43N and longitude 5E, altitude: 376 m)
at Oraison, France. The area experiences an inland Mediterranean type
climate (i.e., sunny with low humidity). It rains less than 90 days per year,
with an irregular pattern during the summer. The mean annual rainfall is
695 mm, with a mean annual temperature of 12.9C. The two fields had a
clayey loam soil, which is classified under the Food and Agriculture
Organization (FAO) system (Driessen et al. 2001) as a typical Fluvisol.
The trial was set up in autumn 2011 in fields that were previously
managed under reduced-tillage practices. Conditions were similar between
the treatments in each plot from the non-crop period. All cultivation operations were conducted by farmers; thus, the two fields differed slightly in their
crop rotation and management practices. The experiment was carried out on
one field cropped with maize (field M) and on one field cropped with
sorghum (field S). Table 1 contains information on the crop rotations used.
Soil-management practices applied to each field during the experiment are
Table 1. Crop rotations per field.
Field
M
S
Rotation
Rape or winter pea
Rape or winter pea
Durum wheat
Durum wheat
Maize
Sorghum or sunflower
68
Table 2. Relevant crop management practices for the study carried out: soil preparation, main
cropping operations, and pest management practices.
Maize (field M)
Month
August 2011
Date
30/08/2011
September 2011 27/10/2011
February 2012
28/02/2012
March 2012
14/03/2012
29/03/2012
30/03/2012
April 2012
21/04/2012
May 2012
11/05/2012
17/05/2012
August 2012
05/06/2012
31/08/2012
October 2012
17/10/2012
Operations
Only for the DMC
treatment: Cover crop
direct sowing;
irrigation: two times
15 mm
Only for the RT
treatment: 12 cm
plowing
Only for the RT
treatment: 15 cm soil
loosening
Herbicide treatment:
Glyfoflash 3 L ha1
(glyphosate 360 g L1)
Maize sowing (cv. Maggi
CS): 81 000 seeds ha1
(seed treatment:
Cruiser (thiametoxam
350 g L1))
Trophe 5 L ha1
(acetochlore 400 g L1)
+ Lagon 0.5 L ha1
(isoxaflutole 75 g L1
and aclonifen 500 g
L1)
Molluscicide treatment:
Sluxx 6 kg ha1 (ferric
phosphate 29.7 g kg1)
Elumis 0.4 L ha1
(mesotrione 75 g L1
and nicosulfuron 30 g
L1)
Elumis 0.4 L ha1
(mesotrione 75 g L1
and nicosulfuron 30 g
L1)
Irrigation beginning
Irrigation end
(410 mm)
Harvest
Sorghum (field S)
Date
25/08/2011
Operations
Only for the DMC
treatment: Cover crop
direct sowing;
irrigation: 25 mm
10/11/2011
Only for the RT

treatment: 13 cm
plowing
Only for the RT
treatment: 8 cm
depth tine harrowing
Glyfoflash 3 L ha1
(glyphosate 360 g L1)
27/02/2012
28/03/2012
11/05/2012
Sorghum sowing (cv.

Solarius): 350 000
seeds ha1; Row
treatment: Super 45
(0-45-0) 90 kg ha1 +
Belem 12 kg ha1
(cypermethrine 8 g
kg1)
20/06/2012
21/08/2012
Irrigation beginning
Irrigation end
(260 mm)
Harvest
04/10/2012
DMC = direct seeding mulch-based cropping system, RT = reduced tillage.
shown in Table 2. Field M was irrigated using a pivot irrigation system and
field S using a hose reel irrigation system. Care was exercised to ensure that
irrigation was similar between treatments in each field. Monitoring of parameters was carried out during spring and summer 2012.
69
Experimental design
Two soil treatments were set up in each field: (1) DMC and (2) RT (tillage to
15 cm deep and without a cover crop). Three replicates (i.e., plots) were
performed for each treatment in a homogeneous area in the center of each
field to avoid edge effects. The experimental area was 150 x 28 m. The
treatments were randomized; each plot was 50 x 14 m. Grain yields for the
maize field were 15.80 (0.56) t ha1 in RT and 18.84 (0.94) t ha1 in DMC,
and 5.45 (0.20) t ha1 for RT and 7.60 (0.32) t ha1 for DMC in the
sorghum field (Kulagowski and Chailleux 2015).
The cover crops in the DMC treatment were consistent across fields,
consisting of a mixture of species, mainly legumes, with low C/N ratio and
biomass of around 3 t ha1 at the time of the first frost (Table 3).
Weed abundance and diversity were monitored using a quadrat (0.25 m2),
from seeding to harvest for maize and until full recovery of the inter-row for
sorghum. Two random samples were monitored for each plot once a week.
Traps creating 0.25 m2-wet artificial refuges (Schrim and Byers 1980;
Hommay et al. 2003) were used to monitor slug density and diversity. One
trap was placed in each plot. Aboveground arthropods were collected weekly
from crop seeding to harvest using one Barber pitfall trap (Barber 1931;
Kromp 1999) per plot. Slugs were counted before seeding until the end of the
crop sensitive stages (i.e., with a two-month interval). Collected arthropod
specimens were identified down to the family level when species could not be
identified using a binocular microscope and determination keys (with the
collaboration of the Luberon Regional Nature Park, Apt, and PSH Unit,
INRA, Avignon) (Jeannel 1941, 1942; Roberts 1985; Trautner and
Geigenmuller 1987; Nentwig et al. 2003; Helsdingen 2009).
Statistical analyses
All statistical analyses were performed using R software (R Development

Core Team, 2009) with the geepack package. For statistical analyses,
Table 3. Cover crop composition in the direct seeding mulch-based cropping system (DMC)
treatment for each field during the previous winter (20112012) and characteristics on 15
December 2011.
Composition
Maize (field M) Field pea (10 kg ha1), grasspea
(10 kg ha1), lentil (5 kg ha1), fenugreek
(3 kg ha1), common vetch (5 kg ha1),
faba bean (10 kg ha1).
Sorghum
Field pea (28 kg ha1), grasspea
(28 kg ha1), faba bean (28 kg ha1),
(field S)
lentil (9.5 kg ha1), soybean (16 kg ha1),
oat (14 kg ha1), radish (6 kg ha1).
Dry matter
(DM) (t ha1)
2.8
Nitrogen content
(% of DM)
4.1247
C/N
10.18
4.0
3.6471
11.5
70
aboveground arthropods were separated depending on their functionality

(Northfield et al. 2012): (i) predators, which mainly consisted of carabid
beetles and arachnids, each evaluated separately, and (ii) detritivores.
Density differences among treatments in predators (carabid beetles and
arachnids), detritivores, and pests (weeds and slugs) were analyzed separately
using generalized estimating equations (GEE) adapted to repeated measures
across time based on the Poisson distribution. The soil treatment and the
date were included as factors in the model.
For carabid beetles, two biodiversity indexes were calculated, the
ShannonWiener index and the Simpson index. The ShannonWiener (H)
index was calculated as follows (Lacoste and Salanon 2005):
H0
S
X
pi log2 pi
i1
where pi nNi is the proportional abundance of each species, and S is the total
number of species. The ShannonWiener index is commonly used to characterize species diversity in a community. It accounts for both the abundance
and evenness of a species and can range from 0.5 (low diversity) to 5 (high
diversity) (Lacoste and Salanon 2005).
An equitability index, also called evenness, the Simpson index (J 0 ) was
calculated as follows:
J 0 H 0 =Hmax
where Hmax is the log2 of the total number of species (Lacoste and Salanon
2005). This index can range from 0 to 1, and is at minimum when a large
proportion of the total community is represented by a small number of
species.
Results
The trapped aboveground arthropods generally belonged to beneficial functional groups. Therefore, we focused our results on these main groups:
predators (mainly consisting of carabid beetles and arachnids) and
detritivores.
Pests
The weed density was significantly higher in the DMC treatments for both
crops (field M: soil treatment: 2 = 4.96, df = 1, P = 0.026; field S: soil
treatment: 2 = 8.00, df = 1, P = 0.0047) irrespective of the date (field M:
soil treatment*date: 2 = 0.96, df = 1, P = 0.326; field S: soil treatment*date: 2 = 0.66, df = 1, P = 0.4161), but remained low in the sorghum
71
Figure 1. Weed population dynamics over time for the reduced tillage (RT) and direct seeding
mulch-based cropping system (DMC) treatments for each field. Mean numbers of weeds per m2
(SEM) are shown.
field (i.e., under 20 weeds m2). The weed density varied significantly
across the weeks (field M: date: 2 = 21.36, df = 1, P < 0.001; field S:
date: 2 = 8.87, df = 1, P = 0.0029), with the highest levels obtained in July
and August (Figure 1). Lolium perenne L. (Poaceae) was the most abundant weed species in both fields, peaking at 32 plants m2 (maize field)
and 25 plants m2 (sorghum field), and reaching higher levels in the DMC
treatments. In the maize field, the main weeds found were L. perenne,
Solanum nigrum L. (Solanaceae), Amaranthus retroflexus L.
(Amaranthaceae). Representatives of Sonchus spp. (Asteraceae), Fumaria
officinalis L. (Fumariaceae), Veronica spp. (Scrophulariaceae), and
Chenopodium album L. (Chenopodiaceae) were occasionally recorded. In
the sorghum field, L. perenne and Sonchus spp. (Asteraceae) dominated;
however, Sonchus spp. (Asteraceae) died before reaching full development,
possibly because of competition with sorghum (personal observation). In
addition, Papaver rhoeas L. (Papaveraceae), Amaranthus retroflexus L.
(Amaranthaceae), and Chenopodium album L. (Chenopodiaceae) were
occasionally recorded.
72
Figure 2. Slug population dynamics over time for the reduced tillage (RT) and direct seeding
mulch-based cropping system (DMC) treatments for each field. Mean numbers of slugs per m2
(SEM) are shown. S indicates the date of sowing, and T indicates the date of the molluscicide treatment.
Slugs were only found in the spring, with higher densities in the DMC
treatment (field M: soil treatment: 2 = 14.00, df = 1, P = 0.00019; field S: soil
treatment: 2 = 4.00, df = 1, P = 0.0463) (Figure 2). The interaction between
the soil treatment and the date was significant in both fields (field M: soil
treatment*date: 2 = 40.2, df = 1, P < 0.001; field S: soil treatment*date: 2 =
6.9, df = 1, P = 0.0084). Two slug species recorded were Deroceras reticulatum (Gastropoda: Pulmonata) and Arion hortensis (Gastropoda: Pulmonata);
however, A. hortensis was only trapped in the maize field at low densities.
Predators
The statistical results are presented in Table 4. Most of the arachnid

species found belonged to the following families: Gnaphosidae,
73
Table 4. Results of aboveground arthropod fauna statistical analyses (GLM with adapted dispersion laws) for the two fields. P-values for the soil treatment and date factors and their interaction
for the three arthropod groups studied are shown.
Maize (field M)
Detritivores
< 0.001
< 0.001
0.596
< 0.001
< 0.001
0.397
Carabids
0.035
< 0.001
0.501
< 0.001
< 0.001
0.014
Spiders
0.001
< 0.001
0.281
< 0.001
< 0.001
0.011
Sorghum (field S)
Soil treatment
Date
Soil treatment*Date
Soil treatment
Date
Soil treatment*Date
Figure 3. Aboveground arthropod fauna population dynamics over time for the reduced tillage
(RT) and direct seeding mulch-based cropping system (DMC) treatments for each field. Mean
numbers of individuals per trap (SEM) are shown.
Lycosidae, Philodromidae, Pisauridae, Salticidae, Sparassidae, and

Thomisidae. Their densities reached the highest levels in June and July
in both fields (Figure 3). The Pardosa genus, belonging to the Lycosidae
family, was the most affected by the treatment, with densities in the DMC
treatment reaching nearly two-fold that of the RT treatment (in June and
July).
74
Potentially predatory ground beetles found belonged mainly to the

Carabidae (Coleoptera) family and Staphylinidae (Coleoptera) were only
occasionally recorded. Carabid densities peaked in July in the maize field
and in August in the sorghum field, and were constantly higher in maize
than in sorghum.
Because of their relevance for biological control, carabid beetles were
investigated in further detail (Table 5). Eleven species of carabid beetles
were recorded; species varied significantly in their abundance and period of
activity. The dominant species found were Poecillus cupreus and Pseudofonus
rufipes in the maize and sorghum fields, respectively. They were both present
throughout the cropping season, but their population dynamics differed, with
P. cupreus population peaking in June and July at more than 170 individuals/
trap, and P. rufipes population peaking later in the season, in August, with
more than 30 individuals/trap. In the maize field, Anchomenus dorsalis was
the second most abundant species, followed by Pterosticus melanarius.
Pterosticus melanarius peaked in April, and then almost disappeared before
being trapped again in August and September, reaching more than 30
individuals/trap. In sorghum, all species other than P. rufipes were found at
low levels, while in RT a slight increase in Calathus fucipes was observed at
the end of August and the other species remained at very low levels, with
single individuals occasionally trapped. In the DMC treatment, species other
than P. rufipes were at higher levels than in RT, with numbers of individuals
ranging from 0 to 5 individuals/trap. The biodiversity indexes were relatively
low for the two treatments, with the appearance of new species at low levels
noted in August and September in the maize field (Figure 4). Differences
were more clear-cut with the Shannon index, which was generally higher in
DMC (Figure 4).
Detritivores
Detritivores were represented by the Anthicidae (Coleoptera), Julidae

(Julida),
Scarabidae
(Coleoptera),
Sylphidae
(Coleoptera),
and
Armadillidiidae (Isopoda) families, and the highest abundance was recorded
in the maize field, with more than 50 individuals/trap in June in the DMC
treatment (i.e., two-fold that of the RT treatment) (Figure 3 and Table 4). In
the sorghum field, populations remained low but were also significantly more
abundant in the DMC treatment (Table 4).
Discussion
Result trends were consistent between the two fields irrespective of the crop
and farm. Every group monitoredpredators, detritivores, and pestswere
more abundant in the DMC treatment than in the RT treatment.
Polyphagous, (Holland 2002) (seed and slug consumption

[Martinkova et al. 2006]).
Generalist predator, occasionally phytophagous (Ribera et al.

1999) (slug consumption [Pianezzola et al. 2013]).
Mainly phytophagous (Ribera et al. 1999)(Seed consumption none

[Holland 2002])
Polyphagous, (Holland 2002)
-
0.043
0.008
0.11
+
+
0.67
0.299
0.056
0.001
< 0.001
0.63
0.001
Soil
treatment
factor
0.34
Ectoparasitoid (particularly Amara sp.) (Saska and Honek

2004).
Ectoparasitoid (Celano and Hansen 1999).
+
none
Nonseed diet (Lundgren et al. 2006).
Predator, occasionally phytophagous (Ribera et al. 1999) (slug

consumption [Cross et al. 2001]).
Predator (Suenga and Hamamura 1998, 2001).
DMC
-
RT
+
Food preferences
Mainly phytophagous (seed diet) occasionally predator
(Ribera et al. 1999).
Generalist predator (Ribera et al. 1999).
DMC = direct seeding mulch-based cropping system, RT = reduced tillage.
Species
Amara aenea (De
Geer 1774)
Anchomenus dorsalis
(Pontoppidan
1763)
Badister unipustulatus
(Bonelli 1813)
Brachinus crepitans
(L. 1758)
Brachinus sclopeta
(Fabricius 1792)
Calathus fuscipes
(Goeze 1777)
Dolichus halensis
(Schalle 1783)
Harpalus affinis
(Schrank 1781)
Poecilus cupreus
(L. 1758)
Pseudoophonus
rufipes
(Degeer 1974)
Pterostichus
melanarius
(llliger 1798)
Relative
abundance
0.92
0.009
0.61
< 0.001
0.002
0.038
0.973
0.81
0.479
Interaction Soil
treatment*Date
< 0.001
P-value
Maize (field M)
DMC
-
none
none none
none
none
RT
+
Relative
abundance
< 0.001
0.2
0.001
< 0.001
0.001
0.81
0.005
0.005
0.005
0.005
Soil
treatment
factor
0.16
0.004
0.23
0.285
0.097
0.47
0.644
0.644
0.644
0.644
Interaction Soil
treatment*Date
< 0.001
P-value
Sorghum (field S)
Table 5. Carabid beetle species found in the experiment, their relative abundance and statistical results of the impact of the soil treatment (either in interaction
with the date factor or not): (+) higher density, (-) lower density, (=) no differences, (none) absence of the species. Food preferences based on published results
are presented.

75
76
Figure 4. Biodiversity index dynamics over time for the reduced tillage (RT) and direct seeding
mulch-based cropping system (DMC) treatments in each field. Mean index (SEM) are shown.
Weed densities were up to three-fold higher under DMC than under RT

management. The results were in accordance with those of previous studies
where DMC increased the weed population because of a lack of physical
destruction and of seed burial (Peign et al. 2007). Cover crops have been
considered a means to overcome this negative effect (Creamer et al. 1996;
Teasdale 1996). The establishment of a winter cover crop in the DMC fields
was expected to outcompete weeds, during the intercrop period, for nutrient
resources, light, and space, thus reducing weed infestation (Teasdale et al.
2007; Lawley et al. 2012). Cover crop mulch is also expected to limit weed
germination and development (Teasdale and Mohler 2000). This phenomenon may have occurred here, but may not have been sufficient to achieve a
similar weed level as in the RT treatment. This may be attributed to the facts
that (i) the cover crop residues were not persistent enough (because of low C/
N) to provide an effective light shield, and (ii) the herbicide strategy was
more adapted to RT than DMC, thus explaining the better results obtained in
the RT treatment. The herbicide strategy was that usually applied by the
farmers who owned the field. This strategy mainly relied on a root absorption
mode of action, but the presence of residues on the surface in the DMC
treatment may have created a physical barrier between the sprayed herbicides
and roots. Moreover, when the soil surface horizon has a high level of
organic matter, as is generally the case under DMC, herbicide molecules
may be adsorbed by colloids and degraded by microorganisms (see, e.g.,
Locke and Bryson 1997; Jones and Bryan 1998; Chauhan et al. 2006). Hence,
an herbicide with a foliar absorption mode of action may be a better alternative under DMC management, but this should be assessed in further
experiments. Another way to improve weed control, using an agroecological
77
strategy, would be to favor cover crops with allelopathic effects (Weston

1996) (i.e., the harmful effect produced in one plant through toxic chemicals
released into the environment by another, Rice 1974, p.1). Among the cover
crop species used in the experiment, field pea and faba bean (Fields M and S),
and oat (Field S) are known to have some allelopathic effects (Fujii 2001;
Kato-Noguchi 2003), but allelopathic species are not always efficient when
mixed with other species (Creamer et al. 1996).
The other major pests at the experimental site were slugs. The lower
presence of slugs in the RT treatment could be explained by the tillage,
which killed them directly and destroyed their shelters (Glen and
Symondson 2003). Conversely, in the DMC treatment, mulch provided
shelters and food sources for slugs, and maintained favorable conditions
(humidity) for their dispersal (Glen and Symondson 2003). Despite the
higher slug densities in the DMC treatment, the final yield (see the
Materials and Methods section) was higher in the DMC treatment than in
the RT treatment. Other studies have shown an increase in the number of
slugs in no-till conditions as compared with tillage, but rarely has there been
any evidence of economic consequences (Stinner and House 1990), possibly
because the plants compensate the lower population density by a greater
development and yield. In addition, the experiment revealed differences in
slug densities between the two fields. This may be related to the time elapsed
between the cover crop destruction and the crop sowing, and the crop sowing
date itself. Sorghum was sown on May 11th, six weeks after destruction of the
cover crop and in the absence of slugs, and thus no molluscicide treatment
was necessary. Conversely, maize was sown on March 29th, two weeks after
destruction of the cover crop and large slug populations were present. Other
studies have highlighted the importance of the sowing timing to limit slug
damage (Byers and Templeton 1988; Douglas and Tooker 2012). The choice
of cover crop composition is also a key factor to reduce slug infestation. For
example, Vernav et al. (2004) observed more slugs in a crop after a clover
(Trifolium pratense) or vetch (Vicia villosa) cover crop than after ryegrass
(Lolium perenne). In our experiment, the cover crops included faba species,
with high nitrogen content, which is generally very palatable for slugs
(Gebauer 2002). When compared with the findings of other studies, these
results highlighted the many different impacts of cover crops, which could
thus be used to improve slug and weed control.
The overall abundance of arthropods was also higher in the DMC treatment than in the RT treatment. Other studies have shown the same trend
between conventional tillage and no-till (i.e., the arthropod diversity was
higher when the soil was not disturbed) (Shenk and Saunders 1994; Stubbs
et al. 2004; Dubie et al. 2011; Errouissi et al. 2011). Highest detritivore
abundance was found in DMC, as expected, as mulch provides both protection and food resources for this group. The lower detritivore abundance
78
during summer could have been caused by (i) unfavorable climatic conditions in the summer (drought), (ii) mulch degradation across time, leading to
fewer food resources and less shelter, and (iii) an increase in the abundance
of their predators. Detritivores, which are widely known to improve the soil
quality (e.g., Heemsbergen et al. 2004; Vos et al. 2011), also play an important role as alternative prey for generalist predators, such as carabid beetles
and spiders, when target prey are scarce. This leads to complex indirect
interactions that can indirectly enhance biological control (Settle et al.
1996; Sigsgaard 2000; Eitzinger and Traugott 2011; Chailleux et al. 2014).
Aboveground predators were also more abundant in the DMC than in the
RT treatment. Similar findings were reported by Holland and Reynolds
(2003) when comparing plowed and non-plowed plots. In our study, the
higher abundance of Pardosa sp. (i.e., hunter species) in the DMC treatment
could be interesting for biological control, as some species of this genus have
been reported to be biocontrol agents of midges and plant- and leaf-hoppers
(Oraze and Grigarick 1989; Sigsgaard 2000). Carabid beetles are known to
feed, depending on species, on eggs and juvenile slugs (Symondson et al.
1996; Bohan et al. 2000) or weed seeds (Honek et al. 2003; Lundgren and
Rosentrater 2007; Bohan et al. 2011). Our experimental design did not allow
us to determine whether such predation occurred, but we observed that the
increase in the carabid population alone was not sufficient to offset the
increase in slug and weed numbers on a short-term scale. Indeed, direct
destruction of slugs and the absence of shelters in RT appeared to keep their
populations at a low level (Yenish et al. 1992; Glen and Symondson 2003).
Pterosticus melanarius, a known slug predator (Symondson et al. 1996), was
trapped, but its preference for one of the two treatments was not clear-cut in
our experiment, with opposite trends observed between the sorghum and the
maize fields. Indeed, unlike most carabids, this species does not seem to be
disrupted by soil tillage (Baguette and Hance 1997).
Carabids were further investigated because of their important role in the
biological control of weeds and slugs. The diversity indexes of this group
were relatively low in the two treatments and the dominant species (i.e., P.
cupreus in maize and P. rufipes in sorghum) are both opportunistic polyphagous species that are not of major interest for biological control.
Although Carabus species are well-known slug predators (e.g., Holland
2004; Pianezzola et al. 2013; Renkema et al. 2014), none were collected in
this study. Indeed, this genus is very sensitive to the regular disruption of
arable habitats by cultivation practices (Kromp 1999). This may explain why
slug control was low in the monitored fields, but the Carabus genus could
recolonize undisturbed fields, which could require additional time.
Interestingly, the slug predator Pterostichus melanarius exhibited a key functional slug control trait (Northfield et al. 2012; Welch and Harwood 2014): it
was more abundant at the beginning of the season, when slugs are the most
79
detrimental to crops. This species should thus be promoted using conservation biological control strategies. However, as noted earlier, soil practices
promoting this species seem unclear as trends were opposite between the two
fields, maybe because of the different crop species grown.
Our results showed that soil practices and cover crops had a marked
impact on fields regarding species abundance in the short-term, even
under different field conditions. Our findings indicate that cover crops
should be regarded as a tool to improve ecosystem services, not only on a
long-term scale, but also when converting to DMC by (i) favoring natural
enemies and (ii) disfavoring pests. The functional traits of cover crops
(e.g., low attractiveness for slugs, allelopathy, and biomass production for
weed competition) should be identified to facilitate choices for practitioners and DMC adoption.
Acknowledgments
We express our thanks to the following farmers for providing access to the study sites and for
crop management: Guy Giraud and Robert Ristorto. We thank Caroline Bertrand (Chamber
of Agriculture of Alpes de Haute Provence) for technical assistance; Yvan Capowiez (PSH
Unit, INRA, Avignon), Christophe Mazzia (Avignon University, Avignon), and Pierre Frapa
(Luberon Regional Nature Park, Apt) for their help in arthropod identification. We are
grateful to Josephine Peign (ISARA, Lyon) for helpful comments on the experimental design
and Alain Ratnadass (UPR HortSys, CIRAD, Montpellier) for useful comments on an earlier
version of the manuscript.
Funding
We thank the Chamber of Agriculture of Alpes de Haute Provence for funding Rmy
Kulagowski.
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Short-Term Effects of Conversion to Direct Seeding

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Short-Term Effects of Conversion to Direct Seeding

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Agroecosystem biodiversity could provide essential services

Received 1 September 2015

Adoption; biological control;

Chamber of Agriculture of Alpes de Haute

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changes, and the globalization of agricultural markets have caused rapid

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(Kromp 1999). Therefore, we carried out on-farm experiments in southern

Materials and methods

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September 2011 27/10/2011

Only for the RT

Sorghum sowing (cv.

DMC = direct seeding mulch-based cropping system, RT = reduced tillage.

JOURNAL OF CROP IMPROVEMENT

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All statistical analyses were performed using R software (R Development

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aboveground arthropods were separated depending on their functionality

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The statistical results are presented in Table 4. Most of the arachnid

JOURNAL OF CROP IMPROVEMENT

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Lycosidae, Philodromidae, Pisauridae, Salticidae, Sparassidae, and

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Potentially predatory ground beetles found belonged mainly to the

Detritivores were represented by the Anthicidae (Coleoptera), Julidae

Polyphagous, (Holland 2002) (seed and slug consumption

Generalist predator, occasionally phytophagous (Ribera et al.

Mainly phytophagous (Ribera et al. 1999)(Seed consumption none

Ectoparasitoid (particularly Amara sp.) (Saska and Honek

Nonseed diet (Lundgren et al. 2006).

Predator, occasionally phytophagous (Ribera et al. 1999) (slug

DMC = direct seeding mulch-based cropping system, RT = reduced tillage.

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Weed densities were up to three-fold higher under DMC than under RT

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JOURNAL OF CROP IMPROVEMENT

strategy, would be to favor cover crops with allelopathic effects (Weston

Downloaded by [Rmy Kulagowski] at 02:42 10 February 2016

JOURNAL OF CROP IMPROVEMENT

Downloaded by [Rmy Kulagowski] at 02:42 10 February 2016

Downloaded by [Rmy Kulagowski] at 02:42 10 February 2016

Downloaded by [Rmy Kulagowski] at 02:42 10 February 2016

JOURNAL OF CROP IMPROVEMENT

Heemsbergen, D. A., M. P. Berg, M. Loreau, J. R. Van Hal, J. H. Faber, and H. A. Verhoef.