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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 SeedingMulch-Based Cropping Systems on Macro-Faunaand Weed Dynamics
Rémy Kulagowski, Laura Riggi & Anaïs Chailleux
To cite this article: Rémy Kulagowski, Laura Riggi & Anaïs Chailleux (2016) Short-Term Effects
of Conversion to Direct Seeding Mulch-Based Cropping Systems on Macro-Fauna and WeedDynamics, Journal of Crop Improvement, 30:1, 65-83
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Short-Term Effects of Conversion to Direct Seeding
Mulch-Based Cropping Systems on Macro-Fauna and WeedDynamics
Rémy Kulagowskia, Laura Riggib, and Anaïs Chailleuxc
aChamber of Agriculture of Alpes de Haute Provence, Oraison, France; bDepartment of Ecology, SwedishUniversity of Agricultural Sciences, Uppsala, Sweden; cCIRAD, UPR HortSys, Montpellier, France
ABSTRACT
Agroecosystem biodiversity could provide essential servicessuch as pest control. One approach currently used to promote
ecosystem services in agricultural systems is to reduce tillageand increase plant diversity. In this study, we assessed theshort-term effects of conversion from reduced tillage (RT) todirect seeding mulch-based cropping systems (DMC) on thedynamics of arthropods (detritivores and predators), and majorpests (slugs and weeds). The study was conducted in twocommercial fields: one cropped with sorghum (Sorghum bicolor L.) and one with maize ( Zea mays L.). We found that bothbeneficial and detrimental groups monitored were more abun-dant in DMC than in RT treatment and that the dominantspecies differed between treatments. Because of their majorrole in agroecosystems by contributing to the control of weedseeds, insects, and slugs, carabid beetles (Carabidae) were
investigated in greater detail, and the results showed theirdiversity was also higher in DMC than in RT. The dominantspecies found were Poecillus cupreus and Pseudofonus rufipes inthe maize and sorghum fields, respectively. The increase inbiological control agents shortly after conversion suggestedthat cover crops should be considered as a pest managementtool, even on a short-term scale.
ARTICLE HISTORY
Received 1 September 2015Accepted 23 October 2015
KEYWORDS
Adoption; biological control;conservation agriculture;ecosystem services; maize;sorghum
Introduction
Biodiversity underpins many ecosystem processes; hence increased biodiver-sity in agroecosystems could provide important ecosystem services for farm-
ers (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 Rémy Kulagowski [email protected] Chamber of Agriculture of Alpes de HauteProvence, Avenue Charles Richaud, 04700 Oraison, France.
JOURNAL OF CROP IMPROVEMENT
2016, VOL. 30, NO. 1, 65–83
http://dx.doi.org/10.1080/15427528.2015.1113222
© 2016 Taylor & Francis
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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 methodsthat aim at maintaining biodiversity and enhancing the sustainability and
autonomy of agroecosystems (Malézieux 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. Inmany 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 bio-diversity 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 agroeco-
systems (Lundgren et al. 2006; Northfield et al. 2012). However, their effec-
tiveness 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 prac-tices on macro-fauna and potential ecosystem services are lacking in Europe
66 R. KULAGOWSKI ET AL.
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(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 con-
version 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 inarable 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 43°N and longitude 5°E, 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 12.9°C. 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 opera-tions 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 Rotation
M Rape or winter pea Durum wheat MaizeS Rape or winter pea Durum wheat Sorghum or sunflower
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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 para-
meters was carried out during spring and summer 2012.
Table 2. Relevant crop management practices for the study carried out: soil preparation, maincropping operations, and pest management practices.
Maize (field M) Sorghum (field S)
Month Date Operations Date Operations
August 2011 30/08/2011 Only for the DMC†
treatment: Cover crop
direct sowing;irrigation: two times
15 mm
25/08/2011 Only for the DMC
treatment: Cover crop
direct sowing;irrigation: 25 mm
September 2011 27/10/2011 Only for the RT‡
treatment: 12 cm
plowing
10/11/2011 Only for the RT
treatment: 13 cm
plowing
February 2012 28/02/2012 Only for the RT
treatment: 15 cm soil
loosening
27/02/2012 Only for the RT
treatment: 8 cm
depth tine harrowing
March 2012 14/03/2012 Herbicide treatment:
Glyfoflash® 3 L ha−1
(glyphosate 360 g L−1)
28/03/2012 Herbicide treatment:
Glyfoflash® 3 L ha−1
(glyphosate 360 g L−1)
29/03/2012 Maize sowing (cv. Maggi
CS®): 81 000 seeds ha−1
(seed treatment:
Cruiser® (thiametoxam
350 g L−1))
30/03/2012 Herbicide treatment:
Trophée® 5 L ha−1
(acetochlore 400 g L−1)
+ Lagon® 0.5 L ha−1
(isoxaflutole 75 g L−1
and aclonifen 500 g
L−1)
April 2012 21/04/2012 Molluscicide treatment:
Sluxx® 6 kg ha−1
(ferricphosphate 29.7 g kg−1)
May 2012 11/05/2012 Herbicide treatment:
Elumis® 0.4 L ha−1
(mesotrione 75 g L−1
and nicosulfuron 30 g
L−1)
11/05/2012 Sorghum sowing (cv.
Solarius®): 350 000
seeds ha−1; Row
treatment: Super 45
(0-45-0) 90 kg ha−1 +
Belem® 12 kg ha−1
(cypermethrine 8 g
kg−1)
17/05/2012 Herbicide treatment:
Elumis® 0.4 L ha−1
(mesotrione 75 g L−1
and nicosulfuron 30 gL−1)
05/06/2012 Irrigation beginning 20/06/2012 Irrigation beginning
August 2012 31/08/2012 Irrigation end
(410 mm)
21/08/2012 Irrigation end
(260 mm)
October 2012 17/10/2012 Harvest 04/10/2012 Harvest
† DMC = direct seeding mulch-based cropping system, ‡ RT = reduced tillage.
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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 ha−1 in RT and 18.84 (±0.94) t ha−1 in DMC,
and 5.45 (±0.20) t ha−1 for RT and 7.60 (±0.32) t ha−1 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 ha−1 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 thecollaboration 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 (2011–2012) and characteristics on 15December 2011.
Composition
Dry matter
(DM) (t ha−1)
Nitrogen content
(% of DM) C/N
Maize (field M) Field pea (10 kg ha−1), grasspea
(10 kg ha−1), lentil (5 kg ha−1), fenugreek
(3 kg ha−1), common vetch (5 kg ha−1),
faba bean (10 kg ha−1).
2.8 4.1247 10.18
Sorghum
(field S)
Field pea (28 kg ha−1), grasspea
(28 kg ha−1), faba bean (28 kg ha−1),
lentil (9.5 kg ha−1), soybean (16 kg ha−1),oat (14 kg ha−1), radish (6 kg ha−1).
4.0 3.6471 11.5
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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 measuresacross 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
Shannon–Wiener index and the Simpson index. The Shannon–Wiener (H ’ )index was calculated as follows (Lacoste and Salanon 2005):
H 0 ¼ XS
i¼1
pilog2 pi
where pi ¼niN is the proportional abundance of each species, and S is the total
number of species. The Shannon–Wiener index is commonly used to char-
acterize 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) wascalculated as follows:
J 0 ¼ H 0=H max
where H max is the log2 of the total number of species (Lacoste and Salanon2005). 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 func-
tional 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: χ 2 = 4.96, df = 1, P = 0.026; field S: soiltreatment: χ 2 = 8.00, df = 1, P = 0.0047) irrespective of the date (field M:
soil treatment*date: χ 2 = 0.96, df = 1, P = 0.326; field S: soil treatment*-date: χ 2 = 0.66, df = 1, P = 0.4161), but remained low in the sorghum
70 R. KULAGOWSKI ET AL.
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field (i.e., under 20 weeds m−2). The weed density varied significantly
across the weeks (field M: date: χ 2 = 21.36, df = 1, P < 0.001; field S:date: χ 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 abun-dant weed species in both fields, peaking at 32 plants m−2 (maize field)
and 25 plants m−2 (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), Fumariaofficinalis L. (Fumariaceae), Veronica spp. (Scrophulariaceae), andChenopodium album L. (Chenopodiaceae) were occasionally recorded. Inthe 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) wereoccasionally recorded.
Figure 1. Weed population dynamics over time for the reduced tillage (RT) and direct seedingmulch-based cropping system (DMC) treatments for each field. Mean numbers of weeds per m2
(±SEM) are shown.
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Slugs were only found in the spring, with higher densities in the DMC
treatment (field M: soil treatment: χ 2 = 14.00, df = 1, P = 0.00019; field S: soil
treatment: χ 2
= 4.00, df = 1, P = 0.0463) (Figure 2). The interaction betweenthe soil treatment and the date was significant in both fields (field M: soil
treatment*date: χ 2 = 40.2, df = 1, P < 0.001; field S: soil treatment*date: χ 2 =6.9, df = 1, P = 0.0084). Two slug species recorded were Deroceras reticula-tum (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 arachnidspecies found belonged to the following families: Gnaphosidae,
Figure 2. Slug population dynamics over time for the reduced tillage (RT) and direct seedingmulch-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 mollusci-cide treatment.
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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 Lycosidaefamily, was the most affected by the treatment, with densities in the DMC
treatment reaching nearly two-fold that of the RT treatment (in June andJuly).
Table 4. Results of aboveground arthropod fauna statistical analyses (GLM with adapted disper-sion laws) for the two fields. P -values for the soil treatment and date factors and their interactionfor the three arthropod groups studied are shown.
Detritivores Carabids Spiders
Maize (field M) Soil treatment < 0.001 0.035 0.001
Date < 0.001 < 0.001 < 0.001
Soil treatment*Date 0.596 0.501 0.281Sorghum (field S) Soil treatment < 0.001 < 0.001 < 0.001
Date < 0.001 < 0.001 < 0.001
Soil treatment*Date 0.397 0.014 0.011
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. Meannumbers of individuals per trap (±SEM) are shown.
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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 wereinvestigated 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 Pseudofonusrufipes in the maize and sorghum fields, respectively. They were both presentthroughout 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, withmore 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 beforebeing trapped again in August and September, reaching more than 30
individuals/trap. In sorghum, all species other than P. rufipes were found atlow levels, while in RT a slight increase in Calathus fucipes was observed atthe 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 individualsranging from 0 to 5 individuals/trap. The biodiversity indexes were relatively
low for the two treatments, with the appearance of new species at low levelsnoted 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 recordedin 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 monitored—predators, detritivores, and pests—weremore abundant in the DMC treatment than in the RT treatment.
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T a b l e
5 . C a r a b i d b e e t l e s p e c i e s f o u n d i n t h e e x p e r i m e n t , t h e i r r e l a t i v e a b
u n d a n c e a n d s t a t i s t i c a l r e s u l t s o f t h e i m p a c t o f t h e s o i l t r e a t m e n t ( e i t h e r i n i n t e r a c t i o n
w i t h t h e d a t e f a c t o r o r n o t ) : ( + ) h i g h e r
d e n s i t y ,
( - ) l o w e r d e n s i t y ,
( = ) n o d
i f f e r e n c e s ,
( n o n e ) a b s e n c e o f t h e
s p e c i e s .
F o o d p r e f e r e n c e s b a s e d o
n p u b l i s h e d r e s u l t s
a r e p r e s e n t e d .
M a i z e ( f i e l d M )
S o r g h u m
( f i e l d S )
R e l a t i v e
a b u n d a n c e
P - v a l u
e
R e l a t i v e
a b u n d a n c e
P - v a l u e
S p e c i e s
F o o d p r e f e r e n c e s
R T
‡
D M C
†
S o i l
t r e a t m e n t
f a c t o r
I n t e r a c t i o n S o i l
t r e a
t m e n t * D a t e
R T
D M C
S o i l
t r e a t m e n
t
f a c t o r
I n t e r a c t i o n S o i l
t r e a t m e n t * D a t e
A m a r
a a e n e a ( D e
G e
e r 1 7 7 4 )
M a i n l y p h y t o p h a g o
u s ( s e e d d i e t ) o c c a s i o n a l l y p r e d a t o r
( R i b e r a e t a l . 1 9 9 9 )
.
+
-
0 . 3
4
<
0 . 0
0 1
+
-
0 . 1
6
<
0 . 0
0 1
A n c h o m e n u s d o r s a l i s
( P o
n t o p p i d a n
1 7 6 3 )
G e n e r a l i s t p r e d a t o r
( R i b e r a e t a l . 1 9 9 9 ) .
-
+
0 . 0
0 1
0 . 4
7 9
-
+
0 . 0
0 5
0 . 6
4 4
B a d i s
t e r u n i p u s t u l a t u s
( B o
n e l l i 1 8 1 3 )
N o n s e e d d i e t ( L u n d g r e n e t a l . 2 0 0 6 ) .
=
0 . 6
3
0 . 8
1
n o n e
+
0 . 0
0 5
0 . 6
4 4
B r a c h
i n u s c r e p i t a n s
( L .
1 7 5 8 )
E c t o p a r a s i t o i d ( p a r t i c u l a r l y A m a r a s p . )
( S a s k a a n d H o n e k
2 0 0 4 ) .
+
n o n e
<
0 . 0
0 1
1
n o n e
+
0 . 0
0 5
0 . 6
4 4
B r a c h
i n u s s c l o p e t a
( F a
b r i c i u s 1 7 9 2 )
E c t o p a r a s i t o i d ( C e l a n o a n d H a n s e n 1 9 9 9 ) .
-
+
0 . 0
0 1
0 . 9
7 3
n o n e
n o n e
0 . 0
0 5
0 . 6
4 4
C a l a t h u s f u s c i p e s
( G o e z e 1 7 7 7 )
P r e d a t o r , o c c a s i o n a
l l y p h y t o p h a g o u s ( R i b e r a e t a l . 1 9 9 9 ) ( s l u g
c o n s u m p t i o n [ C r o s s e t a l . 2 0 0 1 ] ) .
-
+
0 . 0
5 6
0 . 0
3 8
=
0 . 8
1
0 . 4
7
D o l i c h u s h a l e n s i s
( S c
h a l l e 1 7 8 3 )
P r e d a t o r ( S u e n g a a
n d H a m a m u r a 1 9 9 8 ,
2 0 0 1 ) .
-
+
0 . 2
9 9
0 . 0
0 2
-
+
0 . 0
0 1
0 . 0
9 7
H a r p a l u s a f f i n i s
( S c
h r a n k 1 7 8 1 )
M a i n l y p h y t o p h a g o
u s ( R i b e r a e t a l . 1 9 9 9 ) ( S e e d c o n s u m p
t i o n
[ H o l l a n d 2 0 0 2 ] )
n o n e
+
0 . 6
7
<
0 . 0
0 1
n o n e
+
<
0 . 0
0 1
1
P o e c i l u s c u p r e u s
( L .
1 7 5 8 )
P o l y p h a g o u s ,
( H o l l a n d 2 0 0 2 )
-
+
0 . 1
1
0 . 6
1
-
+
0 . 0
0 1
0 . 2
8 5
P s e u d
o o p h o n u s
r u f
i p e s
( D e g e e r 1 9 7 4 )
P o l y p h a g o u s ,
( H o l l a n d 2 0 0 2 ) ( s e e d a n d s l u g c o n s u m p t i o
n
[ M a r t i n k o v a e t a l . 2
0 0 6 ] ) .
-
+
0 . 0
0 8
0 . 0
0 9
-
+
0 . 2
0 . 2
3
P t e r o s t i c h u s
m e
l a n a r i u s
( l l l i g e r 1 7 9 8 )
G e n e r a l i s t p r e d a t o r
, o c c a s i o n a l l y p h y t o p h a g o u s ( R i b e r a e
t a l .
1 9 9 9 ) ( s l u g c o n s u m
p t i o n [ P i a n e z z o l a e t a l . 2 0 1 3 ] ) .
+
-
0 . 0
4 3
0 . 9
2
-
+
<
0 . 0
0 1
0 . 0
0 4
†
D M C
=
d i r e c t s e e d i n g m u l c h - b a s e d c r o p p
i n g s y s t e m ,
‡
R T =
r e d u c e d t i l l a g e .
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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 (Peigné 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 nutrientresources, 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 phenom-
enon 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 thefarmers 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 alter-
native under DMC management, but this should be assessed in furtherexperiments. Another way to improve weed control, using an agroecological
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.
76 R. KULAGOWSKI ET AL.
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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 whenmixed 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 inthe 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 treatmentwas 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, Vernavá 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 treat-
ment 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 protec-tion and food resources for this group. The lower detritivore abundance
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during summer could have been caused by (i) unfavorable climatic condi-
tions 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 impor-
tant role as alternative prey for generalist predators, such as carabid beetlesand 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 treatmentcould 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), wastrapped, 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 poly-phagous species that are not of major interest for biological control.
Although Carabus species are well-known slug predators (e.g., Holland2004; 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 couldrecolonize undisturbed fields, which could require additional time.
Interestingly, the slug predator Pterostichus melanarius exhibited a key func-
tional slug control trait (Northfield et al. 2012; Welch and Harwood 2014): itwas more abundant at the beginning of the season, when slugs are the most
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detrimental to crops. This species should thus be promoted using conserva-
tion 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, evenunder 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 practi-
tioners 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 Peigné (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 Rémy
Kulagowski.
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