INTRODUCTION

In agroecosystems, polyphagous benecial ar-

thropod predators such as carabid and staphylinid
beetles and spiders can efciently reduce the popu-
lations of some pest insects and slugs (Best and
Beegle, 1977; Edwards et al., 1979; Chiverton,
1986; Clarke et al., 1994; Kromp, 1999). Thus,
emphasis has recently been placed on studies to
create and manage farmland habitats for augmenta-
tion of such benecial predators (Thomas et al.,
1991; Lys and Nentwig, 1992; Lys et al., 1994;
Kromp, 1999). These studies indicate that eld
margins and semi-natural habitats with dense vege-
tation harbor high abundance and species richness
of benecial predators. These habitats not only
support feeding and reproductive activity but also
function as overwintering refuges for the predators
(Desender, 1982; Sotherton, 1984, 1985; Thomas
and Marshall, 1999; Pffner and Luka, 2000).
Some studies have shown that predatory epigaeic
insects which had overwintered in eld margins
emigrated into adjacent farmland in spring
(Coombes and Sotherton, 1986; Thomas et al.,
1991, 2002). However, these studies were restricted
to cereal and forage crop elds in Europe and
North America (Kromp, 1999).
In Japan and other temperate Asian regions, rice,
vegetables and fruits are intensively cultivated on
narrow plains and foothill areas of mountains.
Farmland management practices in Japan are dif-
ferent from those in Europe and North America.
Assemblages of carabids and other predators in
Japan thus may be distinct from those in other
world areas, although Luff (2002) compared the
carabid assemblages of various world farmlands in-
cluding Japan mainly at the genus level and dis-
cussed how dominant genera were common among
Appl. Entomol. Zool. 38 (4): 449459 (2003)
449
Ground beetles (Coleoptera: Carabidae) and other insect predators
overwintering in arable and fallow elds in central Japan
Kazuo YAMAZAKI,
1,
*
Shinji SUGIURA
2
and Koji KAWAMURA
2
1
Osaka City Institute of Public Health and Environmental Sciences; Tennoji, Osaka 5430026, Japan
2
Laboratory of Forest Ecology, Graduate School of Agriculture, Kyoto University; Kyoto 6068502, Japan
(Received 5 February 2003; Accepted 26 May 2003)
Abstract
To clarify assemblage patterns of overwintering ground beetles (Coleoptera: Carabidae) and other insect predators in
farmland habitats for the purpose of proper land management to enhance benecial predators, we collected carabid
and other insect predators at eight plots including arable and fallow rice and vegetable elds and a bank of an adjacent
irrigation pond in central Japan. In total, 159 adults and 268 larvae of 33 carabid species, and 178 individuals of at
least 17 species of other insect predators were collected by the quadrat sampling method. In rice elds, both the num-
ber of species and no. of individuals of overwintering carabid beetles increased as the soil became dry and vegeta-
tional succession proceeded, whereas in fallow vegetable elds carabids decreased according to succession. Similar
trends were conrmed in other insect predators. Variations of carabid species richness and abundances among the
plots might be attributed to soil water content, vegetation and prey availability. In early-successional fallow vegetable
elds, the larvae of the carabid genus Harpalus overwintered with high density; this appeared to be because the n-
gergrass Digitaria ciliaris (Poaceae), whose seeds were a potential food for Harpalus, were densely vegetated there.
In a dry fallow rice eld and early-successional vegetable elds, benecial predators such as Dolichus halensis
(Coleoptera: Carabidae), Agrypnus binodulus (Coleoptera: Elateridae), and soldier beetle (Coleoptera: Cantharidae)
larvae hibernated with high densities. For proper farmland management to augment insect predators, it is desirable to
maintain fallow rice and vegetable elds as relatively dry habitats and at early successional stages. Ploughing fallow
elds in winter may reduce overwintering predacious insect larvae.
Key words: Carabidae; hibernation; benecial predators; pest control; habitat preference
* To whom correspondence should be addressed at: E-mail: K.Yamazaki@iphes.city.osaka.jp
temperate world farmlands. Therefore, in order to
use carabid beetles and other benecial predators
as biological control agents in Japan, the assem-
blage patterns in each habitat and habitat prefer-
ences of component species should be claried.
In Japan, Habu and Sadanaga (1961, 1963,
1965, 1969, 1970a, b, 1971) described larval mor-
phology of carabid beetles and noted their biology
in cultivated elds and paddy elds based on labo-
ratory rearings. Carabid assemblages and faunae
have been described from paddy elds, forage crop
eld, cabbage eld, g orchards and vineyards dur-
ing the warm season using pitfall-trapping (Torii,
1974; Yano et al., 1989, 1995; Yahiro et al., 1992;
Ishitani et al., 1994; Ishitani and Yano, 1994; Ishi-
tani, 1996; Suenaga and Hamamura, 2001; Yano,
2002). Yahiro and Yano (1997) reported long-term
data on a carabid assemblage caught by a light trap
set at a farmland area which comprised rice, cereal
and vegetable elds, orchards and irrigation ponds.
However, in Asia including Japan, no quantitative
study on overwintering carabid assemblages in
farmland and adjacent semi-natural habitats has
been reported. Quantitative assemblage data of hi-
bernating carabids and other insect predators in
each farmland habitat is useful to create and man-
age overwintering refuges for the predators.
Carabid life-cycle patterns (breeding season and
hibernating developmental stage) vary across cli-
matic regions and habitats (Murdoch, 1967; Paar-
mann, 1979; Andersen, 1984). However, the life
cycles of most carabid species excluding the tribe
Carabina are not well known in Japan (for Cara-
bina see Sota, 1985). Collecting ground beetles in
winter helps to elucidate their life cycles, espe-
cially their hibernating stages. Information on life
cycles of benecial carabid species may be essen-
tial for proper land-management practices to aug-
ment those already in use; In Europe, deep plough-
ing during autumn or winter is known to reduce the
numbers of overwintering carabid beetles and other
predators (Kromp, 1999; Pffner and Luka, 2000).
This negative effect to carabids may be severe for
hibernating larvae compared with overwintering
adults, since the larvae appear to be more vulnera-
ble to physical disturbance than the adults.
In the present study, we sampled overwintering
carabid beetles and other insect predators in arable
and fallow elds (rice and vegetables) with differ-
ent successional stages and a bank of adjacent irri-
gation ponds in central Japan. Firstly, assemblages
of carabids and other predators in each habitat are
described, and the habitats with high abundance
and species richness of the predators are explored.
Similarities between assemblages are also ana-
lyzed. Overwintering developmental stages and
breeding seasons of the sampled carabid beetles
are compared among habitats. Then, habitat prefer-
ences for benecial species are examined, and in
turn land management practices to enhance bene-
cial insect predators against pest insects in Japa-
nese agroecosystems are discussed.
MATERIALS AND METHODS
Study site. The study site was located in a rural
area of Son-enji (3448N, 13543E, ca. 100 m
above sea level), Hirakata City, Osaka Prefecture,
central Japan (Fig. 1). The landscape of this area is
well preserved, comprising mosaics of coppice
woodlands, rice and vegetable elds, irrigation
ponds and streams, and therefore relatively rare
plants and insects that prefer grassland and marsh-
land habitats were still extant (Kanky-kagaku Co.,
2001). There were many arable and fallow rice and
vegetable elds at the study site. Fallow elds were
at various successional stages. These farmlands
were surrounded by secondary forests composed
mainly of the Japanese red pine Pinus densiora
and the oak Quercus serrata.
Overwintering ground beetles and other insect
predators were sampled at the following eight plots
which were within a 1 km
2
area. Table 1 shows the
environmental conditions of sampling plots. Age
after cultivation in each plot was estimated by con-
sulting with farmers and based on vegetational suc-
cession. Vegetational succession advanced from A
to C in rice elds, and from D to G in vegetable
elds.
Methods. We arbitrarily set three quadrats
(1.6 m1.6 m) per plot on the ground except the
two plots (D: 2 quadrats and F: 1 quadrat), that
were unfortunately ploughed while sampling. All
the quadrats except the bank of an irrigation pond
were set on relatively at and horizontal ground.
Ground inclination of three quadrats on the pond
bank was ca. 50. We then dug up each quadrat to a
depth of 40 cm using hoes, and inspected the soil
thus obtained for ground beetles and other insect
predators (see Yamazaki et al., 1999, 2002). The
450 K. YAMAZAKI et al.
sampling was conducted between December 1999
and March 2000 and in March 2001. As to ants,
only queens were caught because of the difculty
and the labor intensity required for sampling work-
ers. In the present study, we focused on epigaeic
and soil macro-predators, but did not examine
predators on plant surfaces such as lacewings and
syrphids and other micro-predators. The collected
carabid larvae were identied according to the de-
scriptions of Habu and Sadanaga (1961, 1963,
1965, 1969, 1970a, b, 1971) and our collections
(Yamazaki et al., 1999, 2002).
The abundance and species richness of carabid
beetles and other predators per quadrat were com-
pared among the sampling plots. A one-way
ANOVA model was used to compare the number of
species and abundance among the plots in carabid
beetles and other predators, and Scheffs range
test was also used to clarify the difference. Plots D
and F were excluded from these analyses due to the
lack of an adequate data set. The similarities
among assemblages were compared using Wards
minimum-variance clustering method. Prior to the
one-way ANOVA and cluster analysis, the no. of
individuals was logarithmically transformed. Over-
wintering developmental stages and breeding sea-
sons in carabids were compared among the plots.
Breeding seasons were determined on the basis of
the descriptions by Kurosa (1959), Habu and
Sadanaga (1961, 1963, 1965, 1969, 1970a, b,
1971), Sota (1985) and Ishitani (1996). Chi-square
test was used to ascertain whether the percentage
(no. of individuals) of larvae and autumn-breeders
differed among the plots.
Overwintering Ground Beetles in Rice and Vegetable Fields 451
Fig. 1. Study site of the ground beetles and other insect predators overwintering in farmland habitats. A to H denote sampling
plots.
Table 1. The environmental conditions of sampling plots
Plots Land use
Years after
Dominant plants
abandonment
A Arable rice eld Astragalus sinicus (Leguminosae), grasses
B Wet fallow rice eld 1 Oenanthe javanica (Umbelliferae), Stellata sp. (Caryophyllaceae)
C Dry fallow rice eld 1 Erigeron canadensis (Asteraceae), A. sinicus
D Fallow vegetable eld 1 0.5 Digitaria ciliaris (Poaceae), E. canadensis
E Fallow potato eld 1 D. ciliaris
F Fallow vegetable eld 2 1 Solidago altissima, Artemisia indica var. maximowiczii (Ateraceae)
G Old fallow eld 3 Miscanthus sinensis, Pleioblastus fortunei (Poaceae)
H Bank of irrigation pond Mostly bare ground
RESULTS
Overview of predator assemblages
In a total of eight plots, 605 individuals of cara-
bid beetles and other insect predators belonging to
at least 50 species were collected by the quadrat
sampling method. Among them, carabid beetles
predominated in the assemblage, with 427 individ-
uals (70.6% of all the predators) belonging to 33
species (66.0% of all the predator species) that
were collected. The collected carabid beetles con-
sisted of 159 adults (37.2%) of 27 species and 268
larvae (62.8%) of nine species (Table 2). In addi-
tion, one adult and 63 larvae (36.0% of all other
predators) of four click beetle species (Elateridae)
were sampled, and 47 larvae (26.4%) of several
soldier beetle species (Cantharidae) and 26 adults
and four larvae (16.9%) of ve rove beetle species
452 K. YAMAZAKI et al.
Table 2. Collected carabid beetles overwintering at eight plots of arable and fallow elds. Numbers without and in
parentheses are adults and larvae, respectively. denotes none found.
Plot
Carabid species
Breeding
Rice elds Vegetable elds Pond Total
season
a
A B C D E F G H
Carabus yaconinus yaconinus Bates S 1 1
Leptocarabus kumagaii Komiya et Kimura A (5) (5)
Scarites terricola pacicus Bates S 3 3
Lesticus magnus (Motschulsky) S 1 1
Pterostichus sulcitarsis Morawitz S 2 2
P. haptoderoides japanensis Lutshnik A 10 3 13
P. longinquus Bates S 1 23 1 25
P. microcephalus (Motschulsky) S 2 2
Agonum chalcomus (Bates) ? 2 3 3 8
Colpodes japonicus (Motschulsky) ? 1 1
Dolichus halensis (Schaller) A (1) (6) (8) (1) (16)
Amara congrua Morawitz S 15 3 18
Am. chalcites Dejean S 12 1 13
Anisodactylus signatus (Panzer) S 1 1 2 13 17
An. puctatipennis Morawitz S 1 1
?Harpalus vicarius Harold A (18) (18)
H. jureceki (Jedlicka) A (3) (2) 2 1 3(5)
H. griseus (Panzer) A (5) (2) (7)
H. tridens Morawitz A (13) (13) (26)
H. sinicus Hope A (75) (14) 3 3(89)
H. niigatanus Schauberger A 1(2) (7) (92) 1(101)
H. chalcentus Bates S 5 8 13
Oxycentrus argutoroides (Bates) ? 3 3
Trichotichnus congruus (Motschulsky) ? 1 1
T. noctuabundas Habu ? 1 1
Bradycellus subditus (Lewis) ? 13 13
B. grandiceps (Bates) ? 6 6
Acupalpus inornatus Bates S 1 1
Stenolophus iridicolor Redtenbacher S 5 5
S. fulvicornis Bates S 1 1
Anoplogenius cyanescens (Hope) S 2 2
Chlaenius naeviger Morawitz S 1 1
Carabidae Gen. sp. ? (1) (1)
Total 4 33(1) 63(8) 6(104) 14(134) 27(21) 5 7 159(268)
a
S: spring, A: autumn. Breeding seasons of each species were determined based on Kurosa (1959), Habu and Sadanaga
(19611971), Sota (1985) and Ishitani (1996).
(Staphylinidae) were also collected (Table 3).
Assemblage patterns according to vegetational
succession
The number of species and the no. of individuals
of carabids per quadrat were signicantly different
among the sampling plots (one-way ANOVA,
species number, df5, F6.24, p0.01; no. of in-
dividuals, df5, F17.73, p0.0001) (Fig. 2). In
rice elds (plots AC), both the no. of species and
individuals of carabids increased, from arable A to
C (one year old elds), as the soil became dry and
succession proceeded. On the other hand, in fallow
vegetable elds (plots DG), there was a tendency
for overwintering carabid beetles to decrease as
succession proceeded: The no. of species was simi-
lar among plots DF, but plot G harbored a small
number of carabid species (Fig. 2a). The no. of in-
dividuals decreased according to vegetational suc-
cession (Fig. 2b).
The number of species and the no. of individuals
of predators other than carabids per quadrat were
also signicantly different among the sampling
plots (one-way ANOVA, species number, df5,
F4.96, p0.05; No. of individuals, df5,
F7.20, p0.01) (Fig. 3). In rice elds, both
Overwintering Ground Beetles in Rice and Vegetable Fields 453
Table 3. Collected insect predators except Carabidae at eight plots of arable and fallow elds. Numbers in and without
parentheses are adults and larvae, respectively. denotes none found.
Plot
Predator species Rice elds Vegetable elds Pond Total
A B C D E F G H
Histeridae
Atholus pirithous (Marseul) 1 1
Staphylinidae
Olophrum arrowi Scheerpeltz 1 1
Paederus fuscipes (Curtis) 17 17
Lathrobium unicolor Kraatz 1 1
Tachyporus celatus Sharp 1 1 1 3 6
Aleochara sp. 1 1
Staphylinidae Gen. sp. (2) (1) (1) (4)
Elateridae
Agrypnus binodulus (Motschulsky) (23) (35) (58)
Ampedus sp. (2) (2)
Melanotus cete Candeze 1 1
Melanotus sp. (1) (2) (3)
Cantharidae
Cantharidae Gen. spp. (1) (7) (6) (28) (4) (1) (47)
Coccinellidae
Coccinella septempunctata L. 3 3
Formicidae
Crematogaster osakensis Forel
a
10 5 15
Vespidae
Vespa tropica pulchra Buysson
a
2 2
Reduviidae
Peirates turpis Walker 1 1
Gryllotalpidae
Gryllotalpa orientalis Burmeister 1 (2) (5) 1 2(7)
Anisolabididae
Euborellia plebeja (Dohrn) 6 6
Total 1(1) 17(9) 22(36) 3 6(64) 5(5) 1(6) 2 57(121)
No. of quadrats 3 3 3 1 3 2 3 3 21
a
Queens.
species richness and abundance of the other preda-
tors increased with succession, similar to the case
of carabid beetles. In vegetable elds, although the
abundance of the other predators was high at plot E
(fallow potato eld) (Fig. 3b), the no. of species
and abundance were similar among the other plots.
Among the eight plots, the average number of
species of the other predators per quadrat was sig-
nicantly correlated to that of carabids (Spear-
mans correlation coefcient by rank test, N8,
rs0.79, p0.05) (Figs. 2a, 3a), although the aver-
age no. of individuals of other predators per
quadrat was marginally correlated to that of cara-
bids (Spearmans correlation coefcient by rank
test, N8, rs0.64, p0.08) (Figs. 2b, 3b). This
nding suggested that habitat preferences in cara-
bids and other predators were similar.
Predator assemblages at eight plots
Predator assemblage patterns at each plot are
briey described as follows (Tables 2, 3): (A) The
arable rice eld harbored very small numbers
of insect predators (Figs. 1, 2); (B) Wet fallow
rice eld: A hygrophilous carabid, Pterostichus
longiquus was the dominant species (23 individu-
als, 67.6% of carabid individuals), and Acupalpus
inornatus, Stenolophus iridicolor and Paederus
fuscipes were found only from this plot; (C) Dry
fallow rice eld: Carabid (71 individuals of 12
species), staphylinid (four individuals of three
species) and elaterid beetles (23 Agrypnus binodu-
lus larvae) were relatively dominant, and Lepto-
carabus kumagaii, Bradycellus subditus and the
earwig Euborellia plebeja were specic to the plot.
Two carabid species of the genus Amara were
454 K. YAMAZAKI et al.
Fig. 2. Carabid species number and abundance at eight plots of arable and fallow elds. (a) Species number per quadrat, (b)
No. of individuals per quadrat. Means1SE. Different alphabetical letters above the bars indicate statistical difference at p0.05
level of signicance by Scheffs range test. Before analysis, no. of species and individuals were logarithmically transformed.
Fig. 3. Species number and abundance of insect predators other than carabid beetles at eight plots of arable and fallow elds.
(a) Species number per quadrat, (b) No. of individuals per quadrat. Means1SE. Different alphabetical letters above the bars indi-
cate statistical difference at p0.05 level of signicance by Scheffs range test. Before analysis, no. of species and individuals
were logarithmically transformed.
abundantly collected (27 individuals); (D) Fallow
vegetable eld 1: This plot was characterized by
the high abundance of Harpalus larvae (98 individ-
uals of four species). (E) Fallow potato eld: Cara-
bid (148 individuals of eight species), elaterid (36
individuals of two species) and cantharid beetles
(28 individuals) were abundant. Larvae of ve
Harpalus species occurred with high density simi-
lar to plot D; (F) Fallow vegetable eld 2: The
carabids Scarites terricola pacicus, P. sulcitarsis
and B. grandiceps were found only in this plot; (G)
Old fallow eld: The abundance of predators was
very low (Figs. 2, 3), but two species of the carabid
genus Stenolophus were only in this plot; (H) Bank
of irrigation pond: Both species richness and abun-
dance of predators were very low (Figs. 2, 3). Sil-
vicolous species, Carabus yaconinus yaconinus,
Chlaenius naeviger and the hornet Vespa tropica
pulchra were sampled.
Similarities among assemblages
Dendrograms of Fig. 4 show the similarities
among the assemblages of the plots in carabids and
other predators, respectively. For carabid assem-
blages, the fallow eld plots D and E were similar,
but three rice eld plots (AC), especially the dry
fallow rice eld (C), were distinct from each other
(Fig. 4a). For other predator assemblages, the
arable rice eld (A) and the bank of the pond (H)
were similar, possibly due to the small number of
predators sampled (Figs. 2, 3). The dry fallow rice
eld (C) bore a distinct assemblage of insect preda-
tors from other plots (Fig. 4a).
Overwintering developmental stages and breed-
ing seasons in sampled carabids
Table 4 shows the overwintering developmental
stages and breeding seasons in the collected cara-
bid beetles at each plot. In rice elds (AC), there
was a tendency for the percentage of the no. of
species and individuals of larvae that overwintered
to increase somewhat from arable (A) to fallow (B)
and (C). In vegetable elds (DG), as to both no.
of species and individuals however, the percentage
of larvae that hibernated decreased markedly and
the percentage of adults increased with vegeta-
tional succession. Plots D and E had many over-
wintering Harpalus larvae (cf. Table 2). From the
pond bank (H), a small number of adults was
found.
With regard to breeding seasons, in the dry fal-
low rice eld (C), autumn-breeders were found,
while in both the arable rice eld (A) and wet fal-
low rice eld (B), no autumn-breeders were col-
lected. In fallow vegetable elds (DG), with re-
gard to both the no. of species and individuals, the
percentage of autumn-breeders decreased and the
percentage of spring-breeders tended to increase as
succession proceeded. A small number of adult
carabids collected at the pond bank (H) comprised
both spring- and autumn-breeders.
DISCUSSION
Assemblage patterns according to vegetational
succession
In this study, we could not examine arable veg-
etable elds and old rice elds, and our sampling
was restricted to only eight plots. Therefore, the
ndings presented in this paper are snapshots of
overwintering predators in farmland. The present
results may, however, reect some important as-
semblage patterns during vegetational succession
in farmlands. In rice elds (from A to C), overwin-
Overwintering Ground Beetles in Rice and Vegetable Fields 455
Fig. 4. Dendrograms showing the assemblage similarities of (a) carabid and (b) other insect predators hibernated at eight plots
in arable and fallow elds. Wards minimum-variance clustering method was applied. Prior to clustering, no. of individuals was
logarithmically transformed. AC: rice elds, DG: vegetable elds, H: pond bank.
tering carabid beetles increased as the soil became
dry and vegetational succession proceeded, while
in fallow vegetable elds (from D to G) carabids
decreased according to succession (Fig. 2). These
trends also applied more or less to other insect
predators (Fig. 3). The increase in carabids in the
dry fallow rice eld may be attributed to the loss of
water content in the soil and vegetational develop-
ment. In the dry fallow rice eld, the carabids P.
haptoderoides japanensis, two Amara species and
B. subditus were abundantly found (Table 2). Since
these carabid species are known to inhabit rela-
tively dry elds and orchards (Habu and Sadanaga,
1961, 1963; Ishitani and Yano, 1994; Ishitani,
1996), these xerophilous species appeared to over-
winter where they were active during the warm
season. Moreover, the low water content in the soil
may be advantageous for carabids to hibernate.
Frmbs (1994) reported that even hygrophilous
species living in peat bogs migrated to drier hum-
mocks for overwintering. In Europe, high densities
of overwintering carabids and other predators have
been found in dense vegetation (Sotherton, 1984;
Thomas et al., 1991; Pffner and Luka, 2000).
Dense vegetation such as tussock-forming grasses
provides less variable temperature environments
(Bossenbroek et al., 1977). Such temperature
buffering properties of dense vegetation may favor
overwintering carabids. In addition, vegetation fos-
ters many prey species for carabid predators.
Thomas et al. (1992) found that a carabid, Deme-
trias atricapillus fed on prey even in winter, and
that D. atricapillus abundantly overwintered at
dense vegetation areas, but that no signicant rela-
tionships were found between the no. of D. atri-
capillus and their potential prey among habitats.
In fallow vegetable elds, carabid abundance of
larval stage and autumn-breeders decreased from D
to G, according to vegetational succession (Table
4). This was because many carabid larvae of the
456 K. YAMAZAKI et al.
Table 4. Overwintering developmental stages and breeding seasons of carabid beetles collected at
eight plots of arable and fallow elds
Plot
Rice elds Vegetable elds Pond Total
A B C D E F G H
Overwintering stages
No. of species Adults 3 5 8 3 2 7 4 5 24
Larvae 0 1 2 5 6 3 0 0 6
Adults/larvae 0 0 1 0 0 0 0 0 3
Total 3 6 11 8 8 10 4 5 33
Larvae (%) 0.0 16.7 18.2 62.5 75.0 30.0 0.0 0.0 18.2
No. of individuals Adults 4 33 63 6 14 27 5 7 159
Larvae 0 1 8 104 134 21 0 0 268
Total 4 34 71 110 148 48 5 7 427
Larvae (%)
a
0.0 ab 2.9 a 11.3 a 94.5 c 90.1 c 43.8 b 0.0 ab 0 ab 62.8
Breeding seasons
No. of species Spring 1 4 6 2 1 4 4 2 16
Autumn 0 0 4 6 1 5 0 1 9
Unknown 2 2 2 0 6 1 0 2 8
Autumn (%) 0.0 0.0 33.3 75.0 12.5 50.0 0.0 20.0 27.3
No. of individuals Spring 1 30 36 3 13 16 5 2 106
Autumn 0 0 19 107 134 26 0 1 287
Unknown 3 4 16 0 1 6 0 4 34
Total 4 34 71 110 148 48 5 7 427
Autumn (%)
b
0.0 abc 0.0 a 26.8 b 97.3 d 90.5 d 54.2 c 0.0 abc 14.3 abc 67.2
a
The percentages of larval stage differed among plots (chi-square test, df7, c
2
263.4, p0.0001).
b
The percentages of autumn breeders differed among plots (chi-square test, df7, c
2
23.51, p0.0001).
a, b
The values with different alphabetical letters are signicantly different (sequential Bonferroni test (Rice, 1989), a0.05).
genus Harpalus were found from the one-year-old
fallow elds D and E but no Harpalus was found
from the more than three-year-old fallow eld G
(Table 2). All the sampled Harpalus species except
H. chalcentus are known as autumn-breeders and
overwinter as adults and larvae (Habu and
Sadanaga, 1961, 1965, 1970a). Many Harpalus
species larvae feed on the seeds of Poaceae, Polyg-
onaceae, Asteraceae and so on (Kirk, 1972; Jor-
gensen and Toft, 1997; Kromp, 1999; Tooley and
Brust, 2002). Since at plots D and E, D. ciliaris
densely covered the ground, the Harpalus larvae
sampled appeared to feed on the seeds of D. cil-
iaris and hibernated at the sites. Yamazaki et al.
(1999) reported that Harpalus larvae (especially H.
capito and H. sinicus) overwintered with high den-
sities in marshy ground along a river where grasses
vegetated. As vegetational succession proceeded,
D. ciliaris and other grasses declined, possibly re-
sulting in the loss of Harpalus larvae in plot G.
Luff (2002) compared dominant carabid genera
among world farmlands and stated that Harpalus
species were particularly predominant in the Japa-
nese agricultural fauna.
Habitat preferences in benecial species and
farmland management
Although these Harpalus larvae are not preda-
tors but seed-feeders of weeds, some of those
adults have omnivorous feeding habits, sometimes
feeding on insects (Ishitani, 1996). Furthermore,
certain Harpalus species can be used for biological
control of injurious weeds (Brust, 1994; Kromp,
1999; Tooley and Brust, 2002). Therefore, aug-
mentation of Harpalus species in fallow elds ap-
pears to be benecial for farmland management.
Ploughing fallow elds in winter may reduce the
populations of these Harpalus larvae, since larval
stage appears to be more vulnerable than adult
stage due to the fragility of larval integuments.
Dolichus halensis is a benecial predacious
carabid attacking larvae of the diamondback moth
Plutella xylostella and other lepidopteran and
planthopper pests (Habu and Sadanaga, 1963;
Wakisaka et al., 1991), and in the present study the
plots D and E harbored this species with relatively
high densities (Table 2). This carabid is an autumn-
breeder and overwinters as larvae, so ploughing
fallow elds in winter may negatively affect this
species.
Among other insect predators, Agrypnus binodu-
lus (Elateridae) larvae and cantharid larvae were
abundantly found, especially in the dry fallow rice
eld C and fallow potato eld E. A. binodulus larva
is known to be predacious (Kurosa, 1959; hira,
1968), but its feeding preferences are not known.
In Europe, cantharid larvae are voracious predators
feeding on earthworms, aphids and other inverte-
brates (Langenstck et al., 1998) and have high
dispersal ability in farmlands (Traugott, 2002). In
Japan, however, although the taxonomy of adult
cantharid species has been progressively devel-
oped, its larval biology remains little known
(Imasaka, 1998). Bionomics of these elaterid and
cantharid species should be claried to use these
potentially benecial predators for pest control.
In conclusion, the ndings of the present study
suggest that, in farmland management, in order to
increase the numbers of many carabids and other
predators, the rice elds and vegetable elds
should be maintained in a considerably dry state
(but, to sustain regional biodiversity, some rice
elds should maintain permanent waters; e.g. Hi-
daka, 1998). Furthermore, because the old (more
than three-year-old) fallow elds harbored only a
small number of carabid and other predators, it is
recommended to maintain fallow elds at an early
successional stage. In addition, ploughing fallow
elds in winter may reduce the predators, espe-
cially carabid, elaterid and cantharid larvae, since
larvae appear to be more vulnerable to physical
disturbances than adult stages. To validate these
farmland management practices in Japan, further
eld data on predatory insects overwintering in
farmlands of various districts and experimental
studies that manipulate management practices are
required.
ACKNOWLEDGEMENTS
We thank the farmers for permitting us to conduct the pres-
ent study and two anonymous reviewers for improving the
manuscript. Thanks are also due to Dr. Y. Hayashi (Kawanishi
City, Hyogo Prefecture) for identifying staphilinid beetles.
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