BIOLOGICAL

CONSERVATION

Biological Conservation 121 (2005) 157–166

www.elsevier.com/locate/biocon

Reed cutting affects arthropod communities, potentiallyreducing food for passerine birds

Martin H. Schmidt a,*, Ga€etan Lefebvre b, Brigitte Poulin b, Teja Tscharntke a

a Fachgebiet Agrar€okologie, Universit€at G€ottingen, Waldweg 26, 37073 G€ottingen, Germanyb Station Biologique de la Tour du Valat, Le Sambuc, 13200 Arles, France

Received 16 July 2003; received in revised form 12 December 2003; accepted 25 March 2004

Abstract

Winter cutting of Common Reed Phragmites australis (Poaceae) is increasing in Camargue, southern France, potentially affecting

wetlands of high conservation value. We studied its impact on arthropods by comparing four cut and uncut sites with special respect

to avian prey availability in the breeding season. The two most important prey groups for breeding passerines, spiders (Araneida)

and beetles (Coleoptera), were identified to species in trap catches and diet samples. The arthropod assemblages differed significantly

between cut and uncut sites. Phytophagous and saprophagous species showed contrasting responses. Numbers of homopterans

increased in cut reed beds, where green Phragmites stem density was higher. Saprophagous woodlice decreased, presumably due to

the reduced amount of litter. Densities of some vegetation-dwelling spider and beetle species were lower at cut sites, including two of

the most preferred prey species for passerine birds. Consequently, large-scale mechanically harvested reed beds host altered ar-

thropod communities, missing major food components used by vulnerable passerines. However, reed cutting on a small scale may

increase habitat heterogeneity and species richness on a landscape level. To contribute to reed bed conservation, EU schemes should

reward management that leaves uncut reed patches in the proximity of cut areas to permit their recolonization by arthropods.

� 2004 Elsevier Ltd. All rights reserved.

Keywords: Araneae; Coleoptera; Phragmites australis; Disturbance; Biodiversity; Conservation; Habitat management

1. Introduction

Monospecific stands of Common Reed Phragmites

australis are widely distributed in European wetlands.

They host a particular fauna with an outstanding pro-

portion of habitat specialists, including insects, spiders

and birds (Flade, 1994; Ostendorp, 1999; Tscharntke,

1999). Several of the bird species are considered vul-

nerable in Europe (e.g., Great Bittern, Purple Heron,Moustached Warbler), and many reed beds are part of

nature reserves. They are often cut in winter either for

commercial or conservation purposes. Cut reed provides

thatching and other building material, and harvesting is

considered to counteract the silting up of reed beds and

to enhance plant species diversity in the undergrowth

(Cowie et al., 1992; Decleer, 1990; Hawke and Jos�e,

* Corresponding author. Tel.: +49-551-392358; fax: +49-551-398806.

E-mail address: m.schmidt@uaoe.gwdg.de (M.H. Schmidt).

0006-3207/$ - see front matter � 2004 Elsevier Ltd. All rights reserved.

doi:10.1016/j.biocon.2004.03.032

1996). Cut reed beds are drained in winter to facilitatemechanical harvest, and irrigated in spring to enhance

growth (Hawke and Jos�e, 1996; Poulin and Lefebvre,

2002). In natural reed beds, green shoots emerge from a

perennial rhizome in spring, grow over the summer, and

die in autumn to persist as standing dry matter for

several years. In harvested reed beds, the dry stems are

cut about 20–30 cm above the ground in winter. With

the removal of dry reed, nest building of early breedingpasserines is prevented (Baldi and Moskat, 1995; Poulin

and Lefebvre, 2002), overwintering stages of arthropods

are removed or destroyed (Ditlhogo et al., 1992;

P€uhringer, 1975), litter input is reduced (Cowie et al.,

1992; Hawke and Jos�e, 1996), and temperature fluctu-

ations and radiation at the ground level increase (Cowie

et al., 1992; Decleer, 1990). The reed grows thinner,

shorter and more densely, resulting in an increasedprimary production (Cowie et al., 1992; Ostendorp,

1999).

158 M.H. Schmidt et al. / Biological Conservation 121 (2005) 157–166

Old reed stems serve as a source for arthropods to

recolonize young reed in spring (Gessner, 1950;

P€uhringer, 1975). Hence, winter cutting might reduce

the abundances of arthropod species, and this may affect

the passerines that prey upon them. A number of studiesaddressed the ecological impact of reed cutting in Aus-

tria (Kampichler et al., 1994), Belgium (Decleer, 1990;

Gryseels, 1989), England (Cowie et al., 1992; Ditlhogo

et al., 1992), France (Poulin and Lefebvre, 2002), Ger-

many (Ostendorp, 1995, 1999; Kube and Probst, 1999),

Hungary (Baldi and Moskat, 1995) and The Nether-

lands (Mook and van der Toorn, 1982; Graveland,

1999). Five of them focused on the vegetation, four onbirds and only three (Decleer, 1990; Ditlhogo et al.,

1992; Kampichler et al., 1994) on the arthropod fauna.

Kampichler et al. (1994) compared reed infestation by

endophagous insects in two cut and three uncut sites. In

the study of Ditlhogo et al. (1992), arthropods in 30� 40

m cut patches within an old reed bed showed mainly

short-term changes, presumably due to quick recolon-

ization. Decleer (1990) found edge effects in reed bedsless than 20 m wide where neighbouring habitats altered

the spider assemblage. Ostendorp (1999) observed that

stem density kept rising, and stem size falling, during the

first 3 years of consecutive cutting. Hence, the most

significant changes in the arthropod community are

expected to occur in large reed beds after several years of

cutting. We studied the effect of cutting on species

composition, diversity and relative abundance of ter-restrial arthropods, and analysed their importance as

food for insectivorous breeding passerines. The direct

impacts of reed cutting and water management on the

passerine assemblage are dealt with elsewhere (Poulin

and Lefebvre, 2002; Poulin et al., 2002). Here, we focus

on the following hypotheses:

1. Reed cutting affects arthropod species and functional

groups differentially, affecting the composition of thearthropod assemblage.

2. Reed cutting reduces arthropod prey for breeding

passerines.

2. Materials and methods

The study was carried out in Camargue, an alluviallowland of 1400 km2 in southern France comprising

8000 ha of reed beds of which 25% is cut annually

(Mathevet, 2001). Eight study sites were selected based

on their size (>15 ha), homogeneity of reed cover and

accessibility. Four of the studied reed beds had been cut

annually over the last 5 years or more, and four had not

been cut over the last 5 years. Vegetation-dwelling (hy-

pergeic) arthropods were sampled between 20 May and14 June 1999 at four plots per site, 100 m distant from

each other and at least 50 m from the habitat edge. Two

trapping methods were used on each of the four plots.

One yellow bowl of 27.8 cm diameter was placed on a

wooden support 1 m above ground (‘‘water traps’’).

Additionally, six transparent plastic mugs of 7.3 cm

diameter were bound to 12 reed stems each with wire, 60

cm above ground (‘‘stem traps’’). Stem traps effectivelycaptured vegetation-dwelling arthropods, while attract-

ing less flower-visiting insects than the water traps.

Captures from water traps and stem traps were com-

bined to increase sample size, whereby 67% of the

spiders and beetles came from the stem traps. Ground-

dwelling (epigeic) arthropods were captured with six

pitfall traps each at two plots per site, consisting of the

same mug as the stem trap buried down to the soilsurface. Epigeic arthropods were sampled between 18

and 27 July, when the ground was dry at all sites. All

traps were filled with 3 cm of a mixture of monoethylene

glycol and water (1:3) with some drops of detergent and

exposed for 6 days. For each site, the capture data

represent 24 water-trap days plus 144 stem-trap days for

hypergeic arthropods and 72 pitfall-trap days for epigeic

arthropods. Arthropods were preserved in 80% ethanol.Diet samples were taken from birds captured with mist

nets simultaneously to the water- and stem-trapping of

arthropods at each site. Birds were forced to regurgitate

and released 15–30 min later (see Poulin and Lefebvre,

2002 for details). Regurgitated items were preserved in

70% ethanol.

Water level and salinity were measured in May (only

water level) and December of the study year. Charac-teristics of the vegetation layer were measured in 24

quadrats of 50� 50 cm per site at the end of July. The

parameters measured were: number of green and dry

reed stems, total length (from ground level to the highest

leaf tip) of one randomly chosen green stem, thickness

of the litter layer, number of plant species and percent

coverage of all vascular plants apart from Phragmites

australis (referred to as ‘‘plant cover’’). Arthropods fromstem traps and diet samples were counted and identified

to order. Spiders and beetles accounted for 55% of the

prey items taken by birds. They were further identified

to species, or to morphospecies in the case of some

beetles (see Appendix A). 87% of the spiders and beetles

taken by birds could be identified to species. For the

comparison among prey species, dry mass was estimated

from the body length according to Rogers et al. (1976)and Lang et al. (1997).

Capture data were log transformed (Jongman et al.,

1995) to reduce skewness, and percent values (plant

cover) were arcsine-transformed. Arithmetic mean-

s� SE are given in figures, tables and text. Direct gra-

dient analysis (RDA, Jongman et al., 1995) was used to

visualize the correlations between habitat factors and

arthropod abundance patterns. Differences in arthropodorders, spider and beetle species between cut and uncut

sites were tested with multivariate Monte-Carlo-

Permutation tests (MCP, ter Braak and Smilauer, 1998).

M.H. Schmidt et al. / Biological Conservation 121 (2005) 157–166 159

We used species-wise error rates according to t tests

with separate variance estimates (StatSoft Inc., 2000).

Bonferroni corrections would be overly conservative

(Legendre and Legendre, 1998), and the multivariate

tests as well as the fact that 21 out of 115 comparisons inappendix fell below an a-level of 0.05 (p < 0:0001 ac-

cording to Bernoulli-equations) confirm that the overall

pattern is significant (Moran, 2003).

3. Results

The eight reed bed sites showed a marked variation inreed growth, with mean stem length ranging from 87 to

234 cm at uncut sites, and 91–160 cm at cut sites. There

were fewer dry stems (4.8� 4.8 m�2 vs. 224� 45 m�2;

t ¼ 4:82; p ¼ 0:016), and a thinner litter layer (8.4� 0.8

cm vs. 15.9� 1.5 cm; t ¼ �4:32; p ¼ 0:01) at cut than

uncut sites. Additionally, the litter layer in cut reed beds

was looser, consisting mostly of leaves and almost no

decomposing stems, except for the upright stubbles.Mean density of green stems was 85% higher at the cut

sites (198� 31 m�2 vs. 107� 5 m�2; t ¼ 2:89; p ¼ 0:06).There was no significant difference between cut and

uncut sites in stem length (129� 17 cm vs. 165� 31 cm;

t ¼ �1:02; p ¼ 0:36), salinity (3.0� 1.0 gL�1 vs. 3.1�1.2 gL�1; t ¼ �0:064; p ¼ 0:95), water level in May

(7.9� 4.5 cm vs. 2.2� 1.8 cm; t ¼ 1:17; p ¼ 0:31) or

December (4.1� 2.8 cm vs. 15.5� 5.3 cm; t ¼ �1:90;p ¼ 0:12) and plant cover (12.3� 3.4% vs. 10.0� 2.4%;

t ¼ 0:557; p ¼ 0:60). In contrast to our expectations,

plant species richness was similar at cut and uncut sites

(5.0� 1.3 vs. 5.0� 1.0; t ¼ 0:00; p ¼ 1:0). It was not

related to salinity (r ¼ �0:106; p ¼ 0:80), but negativelyto water level in May (r ¼ �0:735; p ¼ 0:038).

Stem traps captured 14,242 arthropods, including

1933 beetles and 520 spiders. Another 901 beetles and285 spiders were captured in water traps, and 7370

beetles plus 5287 spiders in ground pitfall traps. Ar-

thropod distributions clearly segregated between cut

and uncut sites according to orders (Fig. 1(a)), spider

species (Fig. 1(b)) and beetle species (Fig. 1(c)). At the

order level, cutting had a significant impact on the

assemblage (MCP: p ¼ 0:03) and is the environmental

variable that explains the largest proportion ofvariance in the capture data (k ¼ 42%). Aphids

(Aphidina) occurred in 18, and leafhoppers (Auc-

henorrhyncha) in 4.5 times higher numbers in cut than

in uncut reeds (Fig. 1(a); for test statistics see Ap-

pendix A). Spiders, and both larval and adult moths

(Lepidoptera) were captured in smaller numbers at cut

than uncut reed beds. The strongest difference was

shown by saprophagous woodlice (Isopoda), whichwere captured at none of the cut, but all of the uncut

sites in stem traps. Woodlice were significantly less

abundant in cut (5.5� 3.8) than uncut reed beds

(1004� 311; t ¼ �6:9; p < 0:01) in the pitfall traps as

well. In the analyses of spider and beetle species

(Fig. 1(b) and (c)), cutting and the resulting thinner

litter layer and lower density of dry stems were sig-

nificantly correlated with differences in communitycomposition among sites (MCP: p ¼ 0:03). Cutting

had the highest coefficient of determination of the

environmental variables, explaining 41% of the vari-

ance in the spider assemblage, and 25% of the vari-

ance in the beetle assemblage. A species list is given in

Appendix A. The jumping spider Marpissa canestrinii

(Salticidae), immatures of the nocturnal hunting

Clubionidae, Clubiona juvenis (Clubionidae), the am-busher Tibellus oblongus (Philodromidae) and the orb-

web weaver Hypsosinga (Araneidae; sums of H. heri

and a second, yet un-described species) occurred in

lower numbers in cut than in uncut reed beds

(Fig. 1(b)). On the ground, the money spider Gnath-

onarium dentatum (Erigoninae), the wolf spider Pirata

piraticus (Lycosidae), Pachygnatha clercki (Tetragna-

thidae) and the sheet-web weaver Microlinyphia impi-

gra (Linyphiinae) were more numerous in cut reed

beds (Fig. 1(b)). Immature Gnaphosidae had higher

densities in the uncut sites. The dominant, semiaquatic

Cyphon laevipennis (Scirtidae), tended to be more nu-

merous on the vegetation of cut reed beds, together

with the coccinellids Anisosticta novemdecimpunctata

and Hippodamia tredecimpunctata (Fig. 1(c)). Dromius

longiceps (Carabidae), Cerapheles terminatus (Mal-achiidae), Philonthus salinus (Staphylinidae) and Cor-

ticaria sp. (Lathridiidae) were less abundant on cut

reeds. On the ground, Acupalpus parvulus (Carabidae),

Dermestes undulatus (Dermestidae), Philontus punctus

(Staphylinidae) and Paracymus aeneus (Hydrophilidae)

were more abundant in cut reed beds, while Bembidion

fumigatum (Carabidae), Silpha tristis (Silphidae) and

Stenus sp. (Staphylinidae) were more abundant inuncut reed beds (Fig. 1(c)).

Diet samples were taken from 110 Reed Warblers

(Acrocephalus scirpaceus Hermann, 1804), 66 Bearded

Tits (Panurus biarmicus Linnaeus, 1758), 19 Mous-

tached Warblers (Acrocephalus melanopogon Tem-

minck, 1823), 19 Reed Buntings (Emberizia schoeniclus

Linnaeus, 1758) and 14 Great Reed Warblers (Acro-

cephalus arundinaceus Linnaeus, 1758). Of 1814 preyitems, 27% were beetles (21 species) and 28% were

spiders (27 species). All but two spider species had also

been trapped. Other prey groups included Diptera

(10%), Hymenoptera (9%), Rhynchota (8%; mostly

Homoptera) and Gastropoda (5%). The spiders Clubi-

ona juvenis (Cluionidae), C. phragmitis and Marpissa

canestrinii (Salticidae) contributed most to the bird

diet (Table 1), C. juvenis and M. canestrinii beingnegatively affected by cutting. Other species were

eaten in similar numbers, but had distinctly lower

biomasses.

(a)

(b)

(c)

Fig. 1. Ordination of the arthropod assemblages at cut and uncut reed beds (RDA), based on all environmental variables and the relative abundances

of: (a) hypergeic arthropod orders; (b) spider species; (c) beetle species. The diagrams on the left show orders respective species (dots), those on the

right show sites (boxes) and environmental variables (arrows) that significantly affected community structure (MCP: p < 0:05). Filled squares

represent cut sites, open squares uncut sites. Species are labelled, if they were affected by cutting with at least marginal significance (t test: p < 0:10).

Acp, Acupalpus parvulus; Ann, Anisosticta novemdecimpunctata; Aph, Aphidina; Ara, Araneida; Auc, Auchenorrhyncha; Bef, Bembidion fumigatum;

Cet, Cerapheles terminatus; Cli, immature Clubionidae; Clj, Clubiona juvenis; Cos, Corticaria sp.; Cyl, Cyphon laevipennis; Deu, Dermestes undulatus;

Drl, Dromius longiceps; Gnd, Gnathonarium dentatum; Gni, immature Gnaphosidae; Hit, Hippodamia tredecimpunctata; Hyh, Hypsosinga heri and

sp.; Iso, Isopoda; Lea, Lepidoptera adults; Lel, Lepidoptera larvae; Mac, Marpissa canestrinii; Mii, Microlinyphia impigra; Paa, Paracymus aeneus;

Pac, Pachygnatha clercki; Php, Philonthus punctus; Phs, Philonthus salinus; Pip, Pirata piraticus; Sit, Silpha tristis; Sts, Stenus sp.; Tio, Tibellus

oblongus.

160 M.H. Schmidt et al. / Biological Conservation 121 (2005) 157–166

Table 1

Spider and beetle species accounting for a minimum of five items or 1% of the dry mass content in the diet of reed bed passerines

Family Species N m (mg)

Carabidae Agonum viduum 3 29

Chlaenius tristis 1 25

Chrysomelidae Epitrix pubescens 6 1

Coccinellidae Coccidula scutellata 32 14

Heteroceridae Heterocerus obsoletus 9 27

Phalacridae Stilbus oblongus 11 2

Scirtidae Cyphon laevipennis 393 213

Silphidae Silpha tristis 1 44

Other Coleoptera (>13 species) 37 >51a

Araneidae Larinioides folium 6 65

Singa lucina 31 97

Clubionidae Clubiona juvenis 86 270

Clubiona phragmitis 43 541

Dictynidae Argenna patula 94 60

Erigoninae Gnathonarium dentatum 31 9

Lycosidae Arctosa fulvolineata 6 93

Pirata piraticus 10 44

Philodromidae Philodromus glaucinus 21 26

Tibellus oblongus 2 27

Salticidae Marpissa canestrinii 26 252

Myrmarachne formicaria 17 68

Other Araneida (>15 species) 139 >75a

N , number of individuals; m, calculated dry mass of all individuals.aDry mass of the 19 (Coleoptera) resp. 23 (Araneida) individuals, that could be identified to species.

M.H. Schmidt et al. / Biological Conservation 121 (2005) 157–166 161

4. Discussion

Reed cutting significantly altered habitat structure,

the overall arthropod community, as well as spiders and

beetles at the species level, reducing two of the most

important prey species for breeding passerines. The ef-

fects of reed cutting on habitat structure are similar to

those reported from higher latitudes (Cowie et al., 1992;

Hawke and Jos�e, 1996; Ostendorp, 1999), except that wedid not observe higher plant species richness in cut reed

beds. This might be related to the particular water

management of Mediterranean cut reed beds, which

prevents the colonisation of terrestrial plant species

(Poulin and Lefebvre, 2002). The differences in water

level were not significant, but corresponded qualitatively

to the expectations of higher levels in cut reed in spring

due to irrigation, and lower levels in winter due to theartificial drawdown to facilitate harvest. The uncut sites

varied widely in abiotic conditions. Some sites were

relatively dry, with short-growing reeds that hardly

flowered, while others provided good growing condi-

tions with mean stem lengths almost reaching 2.5 m. In

spite of this environmental variability, cutting had a

stronger impact on arthropods both on the vegetation

and on the ground than any other environmentalvariable.

The higher abundance of herbivores (Homoptera)

in cut reeds was probably related to several factors:

high density of green stems, good visibility of green

stems following the removal of dry stalks, more humid

microclimate, low abundance of some predators (spi-ders and predacious beetles), and chemical changes

within the host plant. Thin reed stems are known to

be less resistant to herbivores due to reduced silicate

content (Tscharntke, 1990). On the other hand, low

numbers of woodlice in cut reeds are probably asso-

ciated with the lower availability of litter, which serves

as both food and microhabitat. On the vegetation, a

number of spider and beetle species were less abun-dant at the cut sites. Depending on the biology of

each species, this may be due to the removal or de-

struction of hibernating individuals during harvest

(Ditlhogo et al., 1992; Fr€omel and H€olzinger, 1987),

or to the lack of dry stems and panicles as a foraging

and resting substrate. Spiders such as Clubiona and

Marpissa were observed to deposit their egg sacs

within these structures. Furthermore, the recoloniza-tion of cut reeds is expected to be faster by herbivores

than predators, as higher trophic levels are more af-

fected by disturbance (Holt et al., 1999; Kruess and

Tscharntke, 2000). The hypergeic beetles that were

positively affected by cutting were either aphid ene-

mies (Coccinellidae) or semiaquatic (Cyphon laevipen-

nis). This is probably a consequence of the increased

availability of food resources (aphids) and larvalhabitat (water through spring irrigation), respectively.

Ground-dwelling spiders and beetles preferred cut or

uncut reed beds depending upon species. Species more

common in the uncut sites are probably influenced by

factors similar to those affecting hypergeic species.

162 M.H. Schmidt et al. / Biological Conservation 121 (2005) 157–166

With the autumn rainfalls, many epigeic arthropods

leave the ground to hibernate in dry stems (Obrtel,

1972; P€uhringer, 1975; Neumann and Kr€uger, 1991)

and are removed during reed harvest. Gnaphosidae

hide in reed litter during the day, and suitable brokenstems are scarce on the ground of cut reed beds.

Other epigeic arthropods may, however, be favoured

by the increased humidity in cut reeds, which lasts

longer into the arid Mediterranean summers due to

the artificial spring irrigation. Denis (1954) mentions

drought as a major restriction for spiders in the Ca-

margue, leading to a depression in the abundance and

activity of many species during the summer months.Overall, cutting modifies the spider community to-

wards higher abundances of Linyphiidae (both Erig-

oninae and Linyphiinae), Lycosidae and, partly,

Tetragnathidae (Pachygnatha) at the expense of fami-

lies such as Clubionidae, Gnaphosidae, and Salticidae.

This resembles the difference observed between pe-

rennial open habitats (e.g., fallows) and regularly cut

hay meadows in the European agricultural landscape,although different species are involved (Ekschmitt

et al., 1997; Marc et al., 1999). Cattin et al. (2003)

observed a similar mowing effect in wet meadows of

high conservation value. The shift can be attributed to

the removal of the vegetation layer, which prevents

the accumulation of litter. Linyphiidae are good col-

onizers, and they can probably reach higher densities

in disturbed habitats because of reduced competitionwith other spiders (Łuczak, 1979). The marked reac-

tion of the spider community to cutting emphasizes

their suitability as bioindicators (Marc et al., 1999), a

quality shared by beetles at the species but not the

family level in our study.

Cutting reduced the availability of major prey

species of birds. Thereby, species determination has

turned out to be essential for two reasons. First, in-dividual biomass could be considered, which varied by

nearly two orders of magnitude within prey orders

consumed by single bird species, leading to high dif-

ferences between the number of individuals consumed

and their biomass contribution (e.g., Coleoptera eaten

by the Great Reed Warbler). Second, birds were se-

lective within prey order, and some preferred species

were differently distributed across habitats than theoverall taxon. An obstacle to species-level analysis of

bird prey is the difficulty of covering the whole prey

spectrum. However, given the small body sizes of

groups not determined to species in the bird diet (e.g.,

Homoptera, Diptera), the scarcity of very rewarding

large spider species in cut reed may indicate an overall

degradation of foraging opportunities during the

breeding season. Studies on nestling food of ReedWarblers (Cramp, 1992) have also revealed a higher

contribution of spiders to the diet when estimated by

weight rather than by number.

On a regional scale, patches of cut reeds may increase

arthropod diversity by providing habitat to groups as-

sociated with this type of management. However, some

spiders and beetles, especially within the vegetation-

dwelling species, avoid cut reed beds at least duringspring. It is therefore advisable for species conservation

to preserve uncut reed beds of suitable size that cover

the whole range of abiotic conditions typical of an area.

An approach to maintain higher densities of cutting-

sensitive arthropods and birds in cut reed beds would be

to leave patches or strips uncut at regular intervals.

When size and spacing of the uncut patches are appro-

priate, arthropods could hibernate in them and recolo-nize the neighbouring cut areas in spring. The uncut

patches would also provide nesting opportunities to

passerines (Baldi and Moskat, 1995). EU-subsidies for

reed harvesters in France have resulted in an opposite

trend of increasing large-scale mechanical harvest of

reed beds and embankment to improve water control,

which conflicts with the conservation goals (Mathevet,

2001). Further research is needed to determine the po-tential of spatially explicit reed management schemes,

which could be integrated in the agri-environment policy

to better conserve the fauna without necessarily re-

stricting reed harvest.

Acknowledgements

We are grateful to Michael D. Eyre, Steve J. Ormerod

and two anonymous referees for commenting on earlier

drafts of this manuscript. Jean-Philippe Paul assisted in

the field, and the following persons identified arthropod

and plant species: Louis Bigot (Dytiscidae and Hydro-

philidae), Gerhard Brunne (Carabidae), Manfred D€oberl(Chrysomelidae: Alticinae), Helmut F€ursch (Coccinelli-

dae), Nicolas Gompel (div. Coleoptera), Patrick Grillas(plants), Horst Kippenberg (Chrysomelidae), Bernhard

Klausnitzer (Scirtidae), Stefan Kroo� (Staphylinidae),

Andr�e Mauchamp (plants), Sylvain Piry (Apionidae and

Curculionidae), Jean-Philippe Tamisier (Carabidae) and

Olivier Villepoux (Araneida). We are indebted to all re-

servemanagers who allowed us to work in their reed beds:

St�ephan Arnassant (Syndicat mixte pour la Protection

et la Gestion de la Camargue Gardoise), Eric Coulet(SNPN-R�eserve Nationale de Camargue), Jean-Laurent

Lucchesi (Marais du Vigueirat) and Robert Ducasse

(Environment – Sollac M�editerran�ee), and to St�ephanArnassant (Bouva�u), Philippe Morane (Je m’en Repens),

Jean-Louis Perret (Charnier, Gallician) and R�egis Vianet(Marais Pont de Rousty) for allowing us to sample in

harvested reed beds. This study was supported by the

Fondation Sansouire and carried out under a researchpermit issued by the Centre de Recherche sur la Biologie

des Populations d’Oiseax (CRBPO) at the Mus�eumd’Histoire Naturelle de Paris.

M.H. Schmidt et al. / Biological Conservation 121 (2005) 157–166 163

Appendix A

Arthropods captured at three or more reed beds

Cut Uncut t a

Spiders

Araneidae Hypsosinga heri (Hahn, 1831) & nov. sp.2;3;5 0.3� 0.3 1.8� 0.6 )2.1 0.10

Singa lucina (Audouin, 1836)1–4 0.3� 0.3 0.8� 0.5 )0.9 0.43

Clubionidae Clubiona juvenis (Simon, 1878)1–5 7.9� 0.7 19.1� 6.5 )2.3 0.09

Clubiona phragmitis (C.L. Koch, 1843)1–5 3.5� 1.2 5.3� 1.7 )0.6 0.54

sp. (immature) 11.2� 2.2 81.4� 23.7 )3.2 0.04

Dictynidae Argenna patula (Simon, 1874)1–5 9.0� 3.1 4.8� 1.4 1.2 0.29Erigoninae Erigone vagans (Audouin, 1827)5 3.5� 2.2 0.8� 0.5 1.3 0.27

Gnathonarium dentatum (Wider, 1834)2–5 101� 29.6 4.1� 1.8 5.8 <0.01

Gnaphosidae Drassylus lutetianus (L. Koch, 1866)3 1.5� 0.9 11.3� 4.6 )1.7 0.17

sp. (immature) 0.8� 0.3 15.3� 5.8 )3.7 0.02

Trachyzelotes fuscipes (L. Koch, 1867) 6.5� 5.5 )1.8 0.17

Zelotes serotinus (L. Koch, 1866) 3.0� 1.2 5.5� 4.5 0.0 0.98

Linyphiinae Bathyphantes gracilis (Blackwall, 1844)3 3.0� 0.4 2.6� 1.4 0.8 0.47

Meioneta rurestris (C.L. Koch, 1836) 0.8� 0.5 0.5� 0.3 0.3 0.77Microlinyphia impigra (O.P.-Cambridge, 1871) 9.5� 3.0 0.3� 0.3 5.6 <0.01

Liocranidae Phrurolithus festivus (C.L. Koch, 1835) 0.3� 0.3 0.5� 0.3 )0.7 0.54

Lycosidae Arctosa fulvolineata (Lucas, 1846)3;4 21.0� 8.3 10.8� 5.3 1.2 0.29

Arctosa leopardus (Sundevall, 1833) 1.4� 0.8 0.3� 0.3 1.2 0.31

sp. (immature) 484� 145 325� 89 1.0 0.37

Pardosa prativaga (L. Koch, 1870) 3.1� 1.3 4.0� 3.0 0.2 0.82

Pardosa proxima (C.L. Koch, 1848) 1.6� 1.3 0.5� 0.5 0.7 0.51

Pirata latitans (Blackwall, 1841) 0.5� 0.3 0.8� 0.5 )0.3 0.77Pirata piraticus (Clerck, 1757)1;2;5 58.4� 10.7 8.5� 3.3 3.2 0.04

Trochosa ruricola (De Geer, 1778) 0.9� 0.6 2.0� 1.1 )0.9 0.42

Philodromidae sp. (immature) 0.3� 0.3 2.0� 1.4 )1.2 0.31

Philodromus glaucinus (Simon, 1871)2;3;4 3.9� 1.7 1.3� 1.3 1.3 0.23

Tibellus oblongus (Walckenaer, 1802)4 1.1� 0.5 )2.7 0.08

Salticidae sp. (immature) 1.3� 0.9 2.3� 0.9 )0.9 0.42

Marpissa canestrinii (Ninni, 1868)1–5 0.3� 0.3 2.1� 0.4 )4.0 0.01

Tetragnathidae Pachygnatha clercki (Sundevall, 1823)2;3 25.3� 4.6 0.8� 0.8 7.4 <0.01Thomisidae Ozyptila bicuspis (Simon, 1932) 0.3� 0.3 0.8� 0.3 )1.4 0.21

Beetles

Anthicidae Anthicus laeviceps (Baudi, 1877)3 1.1� 0.7 1.8� 1.8 0.0 0.97Cyclodinus bremei (La Fert�e, 1848) 6.1� 4.4 0.8� 0.8 1.5 0.20

Pseudotomoderus compressicollis (Motschul-

sky, 1839)

15.1� 12.6 9.4� 5.9 0.2 0.83

sp. 1.0� 0.4 1.5� 1.0 )0.1 0.91

Byrrhidae Bothriophorus sp. 16.6� 9.1 9.5� 8.8 1.2 0.28

Cantharidae Cantharis sp. 0.3� 0.3 0.5� 0.3 )0.7 0.54

Carabidae Acupalpus parvulus (Sturm, 1825) 7.9� 4.3 0.5� 0.3 2.9 0.04

Agonum thoreyi Dejean, 1828 0.5� 0.3 43.5� 40.9 )1.2 0.31Agonum viduum (Panzer, 1797)1 81.2� 31.4 622� 369 )1.7 0.14

Badister collaris (Motschulsky, 1844) 16.6� 11.2 2.0� 0.8 0.7 0.51

Badister meridionalis (Puel, 1925) 1.1� 0.6 1.3� 0.8 )0.1 0.94

Bembidion fumigatum (Duftschmid, 1812) 2.0� 0.4 )7.5 <0.01

Bembidion rivulare (Dejean 1831) 41.0� 30.7 31.0� 18.0 0.1 0.93

Brachinus exhalans (Rossi, 1792) 3.1� 2.0 61.3� 60.6 )0.5 0.65

Brachinus nigricornis (Gebler, 1829) 15.4� 5.2 168� 97 )0.3 0.76

Carabus clathratus (von Linn�e, 1761) 1.0� 0.7 4.3� 4.3 )0.3 0.81

Appendix A (continued )

Cut Uncut t a

Chlaenius spoliatus (Rossi, 1790) 1.8� 1.2 1.5� 1.5 0.4 0.73

Chlaenius tristis (Schaller, 1783)1 123� 56 9.5� 8.2 1.9 0.11Dromius longiceps (Dejean, 1826)1;3 0.3� 0.3 6.8� 0.5 )10.3 <0.01

Dyschirius laeviusculus (Putzeys, 1848) 1.2� 0.9 2.5� 2.2 )0.3 0.78

Oodes gracilis (Villa, 1833) 7.9� 3.1 21.3� 6.5 )1.3 0.24

Paratachys bistriatus (Duftschmid, 1812) 0.3� 0.3 1.0� 0.7 )0.9 0.40

Poecilus cupreus (Linnaeus, 1758) 0.3� 0.3 11.3� 10.6 )1.2 0.33

Pterostichus elongatus (Duftschmid, 1812) 20.6� 16.9 22.8� 13.7 )0.3 0.77

Pterostichus cursor (Dejean, 1828) and gracilis

(Dejean, 1828)

5.4� 1.0 76.0� 44.2 )1.9 0.14

Stenolophus mixtus (Herbst, 1784) 2.8� 1.5 0.3� 0.3 1.8 0.15

Trechus quadristriatus (Schrank, 1781) 3.5� 2.8 1.8 0.17

Coccinellidae Anisosticta novemdecimpunctata (Linnaeus,

1758)

1.3� 0.5 2.8 0.07

Coccidula scutellata (Herbst, 1783)1;2;3;5 2.4� 1.6 0.3� 0.3 1.3 0.28

Coccinella septempunctata (Linnaeus, 1758) 1.3� 0.8 0.3� 0.3 1.1 0.32

Hippodamia tredecimpunctata (Linnaeus, 1758) 5.3� 3.3 2.4 0.09

Corylophidae sp. 7.3� 1.7 13.7� 5.9 )0.6 0.55Cryptophagidae Atomaria sp. 2.0� 0.9 7.3� 4.1 )1.1 0.32

Curculionidae Bagous argillaceus (Gyllenhal, 1836)3;5 0.3� 0.3 0.5� 0.3 )0.7 0.54

Mononychus punctumalbum (Herbst, 1784) 2.3� 1.7 5.8� 5.1 )0.3 0.79

Dermestidae Dermestes undulatus (Brahm, 1790) 2.8� 1.0 3.0 0.06

Heteroceridae Heterocerus obsoletus (Curtis, 1828)3;5 3.6� 2.4 0.8� 0.8 0.9 0.39

Histeridae sp. 7.0� 4.6 2.1 0.12

Hydraenidae sp. (1) 1.6� 0.7 0.8� 0.5 0.9 0.41

sp. (2) 0.5� 0.3 1.0� 0.7 )0.4 0.67Hydrophilidae Cercyon sp.5 5.1� 3.4 16.8� 13.9 )0.2 0.82

Enochrus and Cymbiodyta sp.1;3 5.3� 0.8 5.3� 2.5 0.6 0.61

Paracymus aeneus (Germar, 1824) 6.5� 2.4 5.2 0.01

Latridiidae Corticaria sp. 0.5� 0.3 31.5� 23.5 )2.4 0.08

Malachiidae Attalus sp. 0.6� 0.4 27.0� 25.7 )1.2 0.30

Cerapheles terminatus (M�en�etri�es, 1832) 0.4� 0.4 43.8� 14.3 )5.3 0.01

Clanoptilus strangulatus (Abeille de Perrin,

1885)

0.3� 0.3 30.0� 29.7 )1.0 0.38

Nitidulidae sp. 1.3� 0.6 15.2� 13.7 )0.7 0.50

Oedemeridae sp. 0.3� 0.3 1.3� 0.9 )1.0 0.39

Phalacridae Stilbus oblongus (Erichson, 1845)2–5 6.9� 3.6 2.8� 0.9 1.0 0.35

Pselaphidae sp. 29.1� 20.0 6.0� 2.7 1.6 0.16

Ptiliidae sp. 13.6� 5.7 7.6� 3.6 0.9 0.41

Scirtidae Cyphon laevipennis (Tournier, 1868)1–5 222� 42 105� 38 2.2 0.09

Cyphon padi (Linnaeus, 1758)3 6.3� 2.4 4.0� 3.1 1.2 0.30

Silphidae Silpha tristis (Illiger, 1798)1 0.5� 0.3 27.8� 17.9 )3.7 0.02Staphylinidae Astrapaeus ulmi (Rossi, 1790) 3.5� 2.5 )2.0 0.13

Paederus sp. 1.1� 0.8 9.8� 4.0 )1.8 0.13

Philonthus sp. 0.5� 0.5 6.3� 4.0 )1.8 0.15

Philonthus punctus (Gravenhorst, 1802) 10.0� 3.8 1.3� 0.5 3.0 0.03

Philonthus salinus (Kiesenwetter, 1844) 6.7� 2.6 57.0� 27.5 )2.3 0.08

sp. (1) 27.9� 4.8 28.4� 11.0 0.5 0.62

sp. (2) 5.7� 2.0 8.3� 2.2 )1.0 0.37

sp. (3) 10.9� 4.0 10.8� 6.2 0.3 0.75sp. (4) 1.3� 0.6 2.8� 1.8 )0.5 0.63

Stenus sp. 3.0� 1.4 )4.2 0.02

Throscidae Trixagus sp. 2.0� 1.7 15.8� 8.9 )1.2 0.30

164 M.H. Schmidt et al. / Biological Conservation 121 (2005) 157–166

Appendix A (continued )

Cut Uncut t a

Orders

Acarina 109� 102 34.2� 14.4 )0.5 0.66Aphidina 252� 159 14.2� 4.0 3.9 0.02

Araneida 39.9� 6.2 84.4� 12.3 )3.3 0.02

Auchenorrhyncha 194� 32 42.9� 13.9 4.0 0.02

Brachycera 52.3� 2.6 51.0� 8.7 0.4 0.74

Coleoptera (adults) 214� 36 242� 84 0.0 0.97

Coleoptera (larvae) 3.0� 1.2 4.9� 3.9 0.1 0.91

Collembola 24.9� 14.2 383� 212 )1.7 0.15

Heteroptera 2.0� 1.4 1.0� 0.7 0.4 0.69Hymenoptera 34.8� 11.7 63.7� 15.1 )1.5 0.21

Isopoda 78.0� 37.2 )8.8 <0.01

Lepidoptera (adults) 0.3� 0.3 4.6� 1.7 )4.1 0.01

Lepidoptera (larvae) 1.0� 0.4 6.3� 2.3 )2.5 0.05

Nematocera 949� 338 418� 159 1.5 0.18

Psocoptera 1.0� 0.7 0.8� 0.5 0.2 0.87

Thysanoptera 11.0� 3.4 6.3� 3.3 1.4 0.22

Species occurring in less than three reed beds, with the number of indiduals in cut versus uncut reed in brackets. (i) Spiders: immature Agelenidae

(0/2); Agroeca cf. cuprea (Menge, 1866) (1/1); Argyroneta aquatica (Clerck, 1757) (3/0); immature A. aquatica (5/0); Ceratinella brevipes (Westring,

1851) (1/0); Dictyna brevidens (Kulczynski in Chryzer & K., 1897)3;5 (2/0); Diplostyla concolor (Wider, 1834) (8/1); immature Dolomedes sp. (4/0);

Donacochara speciosa (Thorell, 1875)3 (3/0); Enoplognatha mordax (Thorell, 1875) (2/0); Entelecara omissa (O.P.-Cambridge, 1902) (0/1); Erigone atra

(Blackwall, 1841) & E. dentipalpis (Wider, 1830) (10/0); Gongylidiellum murcidum (Simon, 1881) (1/1); immature Hahnidae (0/3); Haplodrassus

macellinus (Thorell, 1871) (0/8); Heliophanus cf. flavipes (0/1); Tenuiphantes tenuis (Blackwall, 1852) (1/0); Micaria pulicaria (Sundevall, 1832) (1/3);

immature Mimetidae (0/1); Minyriolus pusillus (Wider, 1834) (0/3); Myrmarachne formicaria (De Geer, 1778)3;5 (0/2); Mysmena leucoplagiata (Simon,

1879) (0/3); Ozyptila simplex (O.P.-Cambridge, 1862) (0/1); Pardosa cribrata (Simon, 1876) (2/0); P. vittata (Keyserling, 1863) (1/0); Pirata tenuitarsis

(Simon, 1876) (0/2); P. piscatorius (Clerck, 1757)5 (0/3); Porrhomma microphthalmum (O.P.-Cambridge, 1871) (1/1); Runcinia lateralis (C.L. Koch,

1838) (1/1); Silometopus cf. ambiguus (O.P.-Cambridge, 1905) (0/1); Tegenaria nemorosa (Simon, 1916) (0/2); Tetragnatha sp. (1/0); T. extensa (Linn�e,

1758) (1/0); immature Tetragnathidae (3/0); Theridion sp. (1/0); T. hemerobius (Simon, 1914) (1/0); Tibellus maritimus (Menge, 1874) (4/0); Titanoeca

albomaculata (Lucas, 1846) (1/2); immature Titanoecidae (1/0); Trachyzelotes cf. barbatus (L. Koch, 1866) (0/3); T. pedestris (C.L. Koch, 1837) (0/2);

Xysticus sp. (1/0); X. kochi (Thorell, 1845) (2/0); Zelotes tenuis (L. Koch, 1866) (2/2). (ii) Beetles: Achenium cf. depressum (Grav., 1802) (1/0);

Acupalpus exiguus (Dej., 1829) (0/5); Agapanthia villosoviredescens (Deg., 1775) (0/1); Agriotes sordidus (Ill., 1807)4 (0/2); Amblystomus metallescens

(Dej., 1829) (1/0); Anisodactylus virens (Dej., 1829) (1/0); Anotylus cf. rugifrons (Hochh., 1849) (1/0); Aphtona lutescens (Gyll., 1808) (0/3); Bagous

petro (Hbst., 1795) (1/0); Bembidion azurescens (D.T., 1877) (1/0); B. varium (Ol. 1795) (8/0); B. iricolor (Bedel, 1879) (0/1); Bledius cf. furcatus (Ol.,

1811) (15/0); Bruchidae sp. (0/1); Calosoma maderae (F., 1775) (1/0); Cantharis cf. livida (L., 1758) (0/4); Carabus alysidotus (Illiger 1805) (0/1);

Cartodere delamarei (Dajoz, 1962) (2/0); Cassida vittata (Vill., 1789) (0/1); Chaetocnema conducta (Motsch.) (0/1); C. tibialis (Ill., 1807) (0/5);

Coccinella undecimpunctata (L.,1758) (13/0); Coelostoma orbiculare (F.) (0/13); Colotes cf. maculatus (Cast.) (1/0); Copelatus haemorrhoidalis (F.,

1787) (0/1); Cordicomus gracilis (Panzer, 1797) (1/3); Corimalia pallidula (Grav.) (0/1); C. tamarisci (Gyllenthal, 1838) (0/1); Cryptocephalus rugicollis

(Olivier, 1791) (0/3); Cryptopleurum minutum (F., 1775) (2/1); Cyclodinus humilis (Germar, 1824) (0/4); Cylindera germanica (L., 1758) (0/4); Dapsa

trimaculata Motsch. (0/1); Dyschirius chalybaeus biskrensis (Putzeys 1846) (1/0); Epitrix pubescens (Koch, 1803)3 (0/2); Euconnus sp. (3/0); Formi-

comus pedestris (Rossi, 1790) (0/28); Graphoderus cinereus (L., 1758) (4/1); Harpalus distinguendus (Duft., 1812) (0/2); Hippodamia variegata (Goeze,

1777) (1/0); Hydaticus seminiger (Degeer, 1774) (0/3); Hydraenidae sp.3 (1/7); Hydrochara caraboides (L., 1758) (1/0); Ilybius fuliginosus (F., 1792) (1/

0); Lampyridae sp. (0/1); Latridiidae sp. (7/0); Limnoxenus niger (Zschach.) (0/1); Longitarsus lycopi (Foudr., 1860) (0/1); Malthinus sp. (1/1);

Microlestes corticalis (Duft., 1812) (8/2); M. minutulus (Goeze, 1777) (1/4); Migneauxia sp. (0/1); Monolepta erythrocephala (Olivier, 1790) (0/1);

Mordellidae sp. (0/3); Neocrepidodera impressa (Fabricius, 1801) (0/1); Notaris scirpi (F., 1792) (1/3); Noterus crassicornis (M€ull., 1776) (1/0);

Odacantha melanura (L., 1767) (0/28); Oedemeridae sp. (2/0); Oxythyrea funesta (Poda., 1761)1 (0/4); Pachnephorus bistriatus (Muls.) (0/1); Philonthus

sp. (0/3); Poecilus puncticollis (Dej., 1829) (0/1); Pogonus chalceus (Marsh., 1802) (1/1); Polystichus connexus (Fourcr., 1785) (1/0); Protapion nigritarse

(Kirby, 1808) (1/0); Pseudoophonus rufipes (Degeer, 1774) (3/1); Psilothrix sp. (1/0); Scaphisoma sp. (0/1); Scarabaeidae sp; (1/0), Scraptiidae sp. (0/2);

Scymnus gr. apetzi (Mulsant and Rey, 1846) (0/1); S. haemorrhoidalis (Hbst., 1797) (0/1); Sitona discoideus Gyllenthal (0/1); Sphenophorus piceus

(Pall., 1776) (0/5); Staphylinus dimidiaticornis (Gemm., 1851) (0/2); Stenolophus skrimshiranus (Steph., 1828) (1/0); S. teutonus (Schrk., 1781) (0/1).

Mean�SE for cut versus uncut sites. t Tests with separate variance estimates on log-transformed data. Superscript indicates in which bird species’

diet the arthropod species was found: 1Acrocephalus arundinaceus; 2Acrocephalus melanopogon; 3Acrocephalus scirpaceus; 4Emberizia schoeniclus;5Panurus biarmicus.

M.H. Schmidt et al. / Biological Conservation 121 (2005) 157–166 165

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