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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

To link to this article: http://dx.doi.org/10.1080/15427528.2015.1113222

Published online: 09 Feb 2016.

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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

JOURNAL OF CROP IMPROVEMENT 67


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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))


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)


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



7/20

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



8/20

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.



9/20

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.



10/20

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.



11/20

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.



12/20

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 .



13/20

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.



14/20

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



15/20

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



16/20

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.

References

Allen, R. T. 1979. The occurrence and importance of ground beetles in agricultural andsurrounding habitats. In Carabid beetles, 485–505. Netherlands: Springer.

Altieri, M. A. 1999. The ecological role of biodiversity in agroecosystems. Agr. Ecosyst.Environ. 74(1):19–31.

Baguette, M., and T. Hance. 1997. Carabid beetles and agricultural practices: Influence of soil

ploughing. Biol. Agric. Hortic. 15(1–4):185–190.Barber, H. S. 1931. Traps for cave-inhabiting insects. J. Elisha Mitchell Sci. Soc. 46(3):259–266.Barker, G. M. Ed. 2002. Molluscs as crop pests Wallingford, UK: CABI Publishing.

Bohan, D. A., A. C. Bohan, D. M. Glen, W. O. Symondson, C. W. Wiltshire, and L. Hughes.

2000. Spatial dynamics of predation by carabid beetles on slugs. J. Anim. Ecol . 69(3):367–379.

Bohan, D. A., A. Boursault, D. R. Brooks, and S. Petit. 2011. National-scale regulation of the

weed seedbank by carabid predators. J. Appl. Ecol . 48(4):888–898.



17/20

Brussaard, L., P. C. De Ruiter, and G. G. Brown. 2007. Soil biodiversity for agricultural

sustainability. Agr. Ecosyst. Environ. 121(3):233–244.Byers, R. A., and W. C Templeton. 1988. Effects of sowing date, placement of seed, vegetation

suppression, slugs, and insects upon establishment of no-till alfalfa in orchardgrass sod.

Grass Forage Sci. 43(3):279–289.Celano, V., and H. Hansen. 1999. La Carabidofauna e l ’Aracnofauna di una bonifica della

Laguna di Venezia. Bollettino del Museo Civico di Storia Naturale di Venezia 49(1998):55–97.

Chailleux, A., E. K. Mohl, M. Teixeira-Alves, G. J. Messelink, and N. Desneux. 2014. Natural

enemy-mediated indirect interactions among prey species: Potential for enhancing biocon-

trol services in agro-ecosystems. Pest Manag. Sci. 70(12):1769–1779.Chancellor, R. J., and R. J. Froud-Williams. 1986. Weed problems of the next decade in

Britain Crop Prot . 5:66–72.Chauhan, B. S., G. S. Gill, and C. Preston. 2006. Tillage system effects on weed ecology,

herbicide activity and persistence: A review. Anim. Prod. Sci. 46(12):1557–1570.Creamer, N. G., M. A. Bennett, B. R. Stinner, J. Cardina, and E. E. Regnier. 1996. Mechanisms

of weed suppression in cover crop-based production systems. Hort. Science 31(3):410–

413.Cross, J. V., M. A. Easterbrook, A. M. Crook, D. Crook, J. D. Fitzgerald, P. J. Innocenzi, C. N.

Jay, and M. G. Solomon. 2001. Review: Natural enemies and biocontrol of pests of

strawberry in northern and central Europe. Biocontrol. Sci. Techn 11(2):165–216.Douglas, M. R., and J. F. Tooker. 2012. Slug (Mollusca: Agriolimacidae, Arionidae) ecology

and management in no-till field crops, with an emphasis on the mid-Atlantic region. J.Integr. Pest Manag . 3(1):C1–C9.

Driessen, P., J. Deckers, O. Spaargaren, and F. Nachtergaele. 2001. Lecture notes on the major

soils of the world. World Soil Resources Reports 2000, 94, Food and Agriculture

Organization (FAO).

Dubie, T. R., C. M. Greenwood, C. Godsey, and M. E. Payton. 2011. Effects of tillage on soil

microarthropods in winter wheat. Southwest Entomol . 36(1):11–20.Duyck, P. F., A. Lavigne, F. Vinatier, R. Achard, J. N. Okolle, and P. Tixier. 2011. Addition of

a new resource in agroecosystems: Do cover crops alter the trophic positions of generalist

predators? Basic Appl. Ecol . 12(1):47–55.Eitzinger B., and M. Traugott. 2011. Which prey sustains cold-adapted invertebrate generalist

predators in arable land? Examining prey choices by molecular gut-content analysis. J. Appl. Ecol . 48:591–599.

Errouissi, F., S. Ben Moussa-Machraoui, M. Ben-Hammouda, and S. Nouira. 2011. Soil

invertebrates in durum wheat (Triticum durum L.) cropping system underMediterranean semi arid conditions: A comparison between conventional and no-tillage

management. Soil Till Res. 112(2):122–

132.Fujii, Y. 2001. Screening and future exploitation of allelopathic plants as alternative herbicides

with special reference to hairy vetch. J. Crop Prod . 4(2):257–275.Gebauer, J. 2002. Survival and food choice of the grey field slug (Deroceras reticulatum) on

three different seed types under laboratory conditions. Anz. Schadl. 75(1):1–5.

Glen, D. 2002. Integrated control of slug damage. Pesticide Outlook 13(4):137–141.Glen, D. M., and W. O. C. Symondson. 2003. Influence of soil tillage on slugs and their

natural enemies. In Soil tillage in agroecosystems, ed. A. El Titi, 207–227. Florida: CRCPress.

Godan, D. 1983. Pest slugs and snails. Biology and control . Berlin, Germany: Springer Verlag.Grime, J. P. 1998. Benefits of plant diversity to ecosystems: Immediate, filter and founder

effects. J. Ecol . 86(6):902–910.



18/20

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

2004. Biodiversity effects on soil processes explained by interspecific functional dissim-

ilarity. Science 306(5698):1019–1020. van Helsdingen, P. J. 2009. Araneae. In: Fauna Europaea Database European spiders and their

distribution — Distribution —Version 2009. 1. http://www.european-arachnology.org;

Accessed 11 October 2012.

Hinkle, M. K. 1983. Problems with conservation tillage. J. Soil Water Conserv . 38(3):201–206.Holland, J. M. 2002. The agroecology of Carabid beetles. Andover, Hampshire UK: 1–356.Holland, J. M. 2004. The environmental consequences of adopting conservation tillage in

Europe: reviewing the evidence. Agr. Ecosyst. Environ. 103(1):1–25.Holland, J. M., and Reynolds, C. J. 2003. The impact of soil cultivation on arthropod

(Coleoptera and Araneae) emergence on arable land. Pedobiologia 47(2):181–191.Hommay, G., Kienlen, J. C., Jacky, F., and Gertz, C. 2003. Daily variation in the number of

slugs under refuge traps. Ann. Appl. Biol . 142(3):333–339.Honek, A., Martinkova, Z., and Jarosik, V. 2003. Ground beetles (Carabidae) as seed

predators. Eur. J. Entomol . 100(4):531–544.

Jeannel, R. 1941. Faune de France, n°39, Coléoptères Carabiques, première partie. Librairie dela Faculté des Sciences, Paris.

Jeannel R. 1942. Faune de France, n°40, Coléoptères Carabiques, deuxième partie. Librairie de

la Faculté des Sciences, Paris.

Jones, M. N., and N. D. Bryan. 1998. Colloidal properties of humic substances. Adv. Colloid.Interfac. 78(1):1–48.

Kato-Noguchi, H. 2003. Isolation and identification of an allelopathic substance in Pisumsativum. Phytochemistry 62(7):1141–1144.

Koskinen, W. C., and C. G. McWhorter. 1986. Weed control in conservation tillage. J. Soil Water Conserv . 41(6), 365–370.

Kromp, B. 1999. Carabid beetles in sustainable agriculture: a review on pest control efficacy,

cultivation impacts and enhancement. Agr. Ecosyst. Environ. 74(1):187–228.Kulagowski, R., and A. Chailleux. 2015. Short-term effects of conversion from reduced tillage

to direct-seeding mulch-based cropping systems. J. Crop Improvement . 29(5):650–668.Lacoste A. and R. Salanon. 2005. Éléments de géographie et d ’ écologie. 2e éd. Armand Colin,

Coll. Fac. Géographie, France.

Lahmar, R. 2010. Adoption of conservation agriculture in Europe: Lessons of the KASSA

project. Land Use Policy 27(1):4–10.Landis, D. A., S. D. Wratten, and G. M. Gurr. 2000. Habitat management to conserve natural

enemies of arthropod pests in agriculture. Ann. Rev. Entomol . 45(1):175–201.Langdale, G. W., R. L. Blevins, D. L. Karlen, D. K. McCool, M. A. Nearing, E. L. Skidmore, A.

W. Thomas, D. D. Tylerv, and, J. R. Williams. 1991. Cover crop effects on soil erosion by wind and water. Cover crops for clean water. Soil and Water Conserv. Soc., Ankeny, IA,

15–22.

Lawley, Y. E., J. R. Teasdale, and R. R. Weil. 2012. The mechanism for weed suppression by a

forage radish cover crop. Agron. J . 104(2):205–214.Locke, M. A., and C. T. Bryson. 1997. Herbicide-soil interactions in reduced tillage and plant

residue management systems. Weed Sci. 45:307–320.Lundgren, J. G., and K. A. Rosentrater. 2007. The strength of seeds and their destruction by

granivorous insects. Arthropod.-Plant Inte. 1(2):93–99.Lundgren, J. G., J. T. Shaw, E. R. Zaborski, and C. E. Eastman. 2006. The influence of organic

transition systems on beneficial ground-dwelling arthropods and predation of insects and

weed seeds. Renew Agr. Food Syst . 21(04):227–237.


http://www.european-arachnology.org/http://www.european-arachnology.org/


19/20

Malézieux, E. 2012. Designing cropping systems from nature. Agron. Sustain Dev . 32(1):15–29.

Martinkova, Z., P. Saska, and A. Honek. 2006. Consumption of fresh and buried seed by

ground beetles (Coleoptera: Carabidae). Eur. J. Entomol . 103(2):361–364.Moonen, A. C., and P. Barberi. 2008. Functional biodiversity: an agroecosystem approach.

Agr. Ecosyst. Environ. 127(1):7–21.

Nentwig, W., A. Hänggi, C. Kropf, and T. Blick. 2003. Central European spiders. An internetidentification key. http://www.araneae.unibe.ch (Version 04.2013); Accessed 11 October 2012.

Northfield, T. D., D. W. Crowder, R. Jabbour, and W. E. Snyder. 2012. Natural enemy

functional identity, trait-mediated interactions and biological control. In Trait-mediated indirect interactions: Ecological and evolutionary perspectives, 450–465. New York:Cambridge University Press.

Oraze, M. J., and A. A. Grigarick. 1989. Biological control of aster leafhopper (Homoptera:

Cicadellidae) and midges (Diptera: Chironomidae) by Pardosa ramulosa (Araneae:Lycosidae) in California rice fields. J. Econ. Entomol . 82(3):745–749.

Peigné, J., B. C. Ball, J. Roger-Estrade, and C. David. 2007. Is conservation tillage suitable for

organic farming? A review. Soil Use Manage. 23(2):129–

144.Phillips, R. E., R. L. Blevins, G. W. and Thomas. 1980. No-tillage agriculture. Science 208:6.Pianezzola, E., S. Roth, and B. A. Hatteland. 2013. Predation by carabid beetles on the

invasive slug Arion vulgaris in an agricultural semi-field experiment. B Entomol. Res. 103(2):225–232.

R Development Core Team. 2009. R: A language and environment for statistical computing.

R Foundation for Statistical Computing, Vienna, Austria.

Ratnadass, A., P. Fernandes, J. Avelino, and R Habib. 2012. Plant species diversity for

sustainable management of crop pests and diseases in agroecosystems: A review. Agron.Sustain. Dev . 32(1):273–303.

Renkema, J. M., G. C. Cutler, D. Blanchard, and A. Hammermeister. 2014. Using ground

beetles (Coleoptera: Carabidae) to control slugs (Gastropoda: Pulmonata) in salad greensin the laboratory and greenhouse. The Canadian Entomologist 1–12. DOI: http://dx.doi.org/10.4039/tce.2014.8.

Ribera, I., G. N. Foster, I. S. Downie, D. I. McCracken, and V. J. Abernethy. 1999. A

comparative study of the morphology and life traits of Scottish ground beetles

(Coleoptera, Carabidae). Ann. Zool. Fenn. 36(1):21–37.Rice, E. L. 1974. Allelopathy . New York: Academic Press.Roberts, M. 1985. The spiders of Great Britain and Ireland . Volume 1: Atypidae to

Theridiosomatidae. Colchester: Harley Books.Saska, P., and A. Honek. 2004. Development of the beetle parasitoids, Brachinus explodens

and B. crepitans (Coleoptera: Carabidae). J. Zool . 262(1):29–

36.Schrim, M., and R. A. Byers. 1980. A method for sampling three slug species attacking sod-

seeded legumes [Derocerus reticulatum, Derocerus laeve]. Melsheimer EntomologicalSeries.

Scopel, E., B. Triomphe, F. Affholder, F. A. M. Da Silva, M. Corbeels, J. H. V. Xavier, R.

Lahmar, S. Recous, M. Bernoux, E. Blanchart, I. De Carvalho Mendes, S. De Tourdonnet.

2013. Conservation agriculture cropping systems in temperate and tropical conditions,

performances and impacts. A review. Agron. Sustain. Dev . 33(1):113–130.Séguy, L., S. Bouzinac, and O. Husson. 2006. In Biological approaches to sustainable soil

systems, eds. N. T. Uphoff, A. S. Ball, E. C. M. Fernandes, H. R. Herren, O. Husson, M. V.Laing, C. Palm, J. Pretty, P. Sanchez, N. Sanginga, J. Thies, 323–342. Boca Raton: CRC

Press.


http://www.araneae.unibe.ch/http://www.araneae.unibe.ch/


20/20

Settle, W. H., H. Ariawan, E. T. Astuti, W. Cahyana, A. L. Hakim, D. Hindayana, and A. S.

Lestari. 1996. Managing tropical rice pests through conservation of generalist natural

enemies and alternative prey. Ecology 77(7):1975–1988.Shearin, A. F., S. C. Reberg-Horton, and E. R. Gallandt. 2007. Direct effects of tillage on the

activity density of ground beetle (Coleoptera: Carabidae) weed seed predators. Environ.Entomol . 36(5):1140–1146.

Shenk, M., and J. L. Saunders. 1984. Vegetation management systems and insect responses inthe humid tropics of Costa Rica. Int. J. Pest Manage. 30(2):186–193.

Sigsgaard, L. 2000. Early season natural biological control of insect pests in rice by spiders-

and some factors in the management of the cropping system that may affect this control. In

European Arachnology 2000 Proceedings of the 19th Colloquium of Arachnology, Aarhus,

57–64.

South, A. 1992. Terrestrial slugs. Biology, ecology and control . London, UK: Chapman andHall, Ltd.

Stinner, B. R., and G. J. House. 1990. Arthropods and other invertebrates in conservation-

tillage agriculture. Ann. Rev. Entomol . 35(1):299–318.

Stubbs, T. L., A. C. Kennedy, and W. F. Schillinger. 2004. Soil ecosystem changes during thetransition to no-till cropping. J. Crop. Improvement 11(1):105–135.Suenaga, H., and T. Hamamura. 1998. Laboratory evaluation of carabid beetles (Coleoptera:

Carabidae) as predators of diamondback moth (Lepidoptera: Plutellidae) larvae. Environ.Entomol . 27(3):767–772.

Suenaga, H., and T. Hamamura. 2001. Occurrence of carabid beetles (Coleoptera: Carabidae)

in cabbage fields and their possible impact on lepidopteran pests. Appl. Entomol. Zool . 36(1):151–160.

Symondson, W. O. C., D. M. Glen, C. W. Wiltshire, C. J. Langdon, and J. E. Liddell. 1996.

Effects of cultivation techniques and methods of straw disposal on predation by

Pterostichus melanarius (Coleoptera: Carabidae) upon slugs (Gastropoda: Pulmonata) in

an arable field. J. Appl. Ecol . 33:741–753.Teasdale, J. R. 1996. Contribution of cover crops to weed management in sustainable

agricultural systems. J. Prod. Agric. 9(4):475–479.Teasdale, J. R., and C. L. Mohler. 2000. The quantitative relationship between weed emer-

gence and the physical properties of mulches. Weed Sci. 48(3):385–392.Teasdale, J. R., L. O. Brandsaeter, A. Calegari, and F. S. Neto. 2007. Cover crops and weed

management. In Non-chemical weed management: Principles, concepts and technology , eds.M. K. Upadhyaya, and R. E. Blackshaw, 49–64. Wallingford, UK: CABI.

Trautner, J., and K. Geigenmuller 1987. Tiger beetles, ground beetles. Illustrated key to the

Cicindelidae and Carabidae of Europe. Germany: Margraf, Aichtal.

Vernavá, M. N., P. M. Phillips-Aalten, L. A. Hughes, H. Rowcliffe, C. W. Wiltshire, and D. M.Glen. 2004. Influences of preceding cover crops on slug damage and biological control

using Phasmarhabditis hermaphrodita. Ann. Appl. Biol . 145(3):279–284.Vos, V. C.A., J. van Ruijven, M. P. Berg, E. T. Peeters, and F. Berendse. 2011. Macro-

detritivore identity drives leaf litter diversity effects. Oikos 120(7):1092–1098.Welch, K. D., and J. D. Harwood. 2014. Temporal dynamics of natural enemy-pest interac-

tions in a changing environment. Biol. Control 75:18–27.Weston, L. A. 1996. Utilization of allelopathy for weed management in agroecosystems.

Agron. J . 88(6):860–866.Yenish, J. P., J. D. Doll, and D. D. Buhler. 1992. Effects of tillage on vertical distribution and

viability of weed seed in soil. Weed Sci. 429–433.


short-term effects of conversion to direct seeding mulch-based cropping systems on macro-fauna and...

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