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Diss. ETI! No. 13698
Insect
diversity
The effects of
in
agricultural grasslands:
management and landscape
structure
A dissertation submitted to the
Swiss Federal Institute
for the
Doctor
of
of
Technology Zurich
degree
of
Natural Sciences
presented by
Manuela Di Giulio Müller
Dipl.
zool.
University
bora L7
July
of Zurich
1970
from Bettlach SO
accepted
on
the recommendation of
Prof. Peter J. Edwards, examiner
Dr. Erhard
Meister, coexaminer
Dr. Bernhard Merz, coexaminer
2000
Table of contents
1
Summary
3
Zusammenfassung
5
General introduction
Chapter
i.
Enhancing
insect
management and
Chapter
2, Insect and
Chapter
3. The
bug
diversity in agricultural grasslands:
landscape structure
plant diversity patterns
fauna
(Heteroptera)
Schaffhauser Randen
updated
checklists of
(SH)
of
in
The roles of
agricultural grasslands
and Rottal
Heteroptera
in the
23
36
(LU). Switzerland, with
of the Cantons Luzern and
Schaffhausen
Chapter
plant diversity and food quality on
of plant feeding heteropteran bugs (Heteroptera:
4. The influence of host
survival
larval
62
Stenodemini)
Conclusions
Curriculum vitae
79
1
Summary
study investigates the influence of management and landscape structure on
insect diversity of hay meadows. The aim of the thesis is to examine what are the most
important environmental factors affecting the insect fauna of meadows and how insect
diversity can be enhanced m agricultural landscapes. Meadows have been chosen as the
study system because m Switzerland they cover a large part of the agriculturally used
The four
area, and they therefore represent an important structure of cultural landscapes.
chapters of the thesis deal with the following questions: l) How does management
This
affect the insect
of
diversity
grasslands?
Do different
grassland types
have characteristic
2) How important are the various grassland types for species
landscape level? How does the surrounding landscape affect the insect
insect communities?
diversity
single meadows? 3) How valuable arc the meadows of the research area
bug fauna of Switzerland? 4) How does plant diversity influence the insect
community
for the
a
on
of
diversity of grasslands'?
oligophagous insects?
Does
plant di\crsity
host
bugs (Heteroptera) were chosen as
they are ecologically very diverse and
The true
because
habitat for this insect group. The research
Northern Switzerland and
with
with
a
number of
a
belongs
nested block
area
agricultural
habitats
grassland
were
represented by the four enclaves (area),
grassland management types (officially referred
study.
(landscape type)
investigated
we
to as medium intensive and
replicate
selected three
area
land. Four such enclaves
selected for the
and within each block
we
diversity,
extensive forested
with two levels. The block level
and for each management type
of
because meadows represent a typical
is located in the Schaffhauser Randen in
less isolated enclaves of
design
development
indicator group for insect
an
to the Jura formation. It is an
mixtures of arable and
varying
chose
more or
affect the larval
We
was
two
extensive),
sites.
presented m chapter 1 indicate that management greatly affects the
heteroptcran bug diversity of grasslands. Extensive meadows are richer in species than
medium intensive ones. Further, species differ m their response to management, and
thus species composition is also affected. Two species appear to benefit from intensive
The
data
management and
species develop
are
two
most
abundant in medium intensive meadows.
generations per year and
these characteristics enable them to
affected
arc
intensive
relatively mobile; it is argued that
highly disturbed habitats. Six species,
are
survive in
adversely
by
extensively managed meadows. Most of
development takes place in spring. Frequent
however,
Both of these
management and
these
and
overwinter
arc
most
eggs
as
early cutting, therefore,
abundant
and
may
m
larval
damage
reproducing. Ground
living species which need a warm and dry microclimate are also adversely affected by
intensive management. Medium intensive meadows are characterised by a dense and
closed vegetation and therefore do not represent suitable habitats for these species.
the larval
The
stages and prevent the
bug
from
insects
fauna of extensne meadows differs
The medium intensive sites, however, show
species
richness and composition.
appear to
have
densities but
are
2
Chapter
explained by
be
Most
often restricted to
examines
m
order
to
greatly
of the
one or a
maintain
areas
investigated.
only in individual areas and
species can reach locally high
occur
rare
lew sites. It
species
whether the differences between
the structure of the
between the
rather small variation in respect to their
Many bug species
restricted ranges.
many different sites
a
developing
and
is
therefore crucial to preserve
dnersity
areas in
surrounding landscape.
landscape level.
species composition can
at
the
The results indicate that the
2
surrounding might affect the bug fauna of a meadow
and that some species apparently benefit from the presence of this habitat type.
However, it is concluded that the spatial scale used in this study was too coarse to
understand in detail how insects are affected by their surrounding habitats.
proportion
of arable land in the
Chapter
bug
3
investigates
the
importance
of the meadows in the research
findings
updated species list
and
of
the
fauna of Switzerland. Based
on
my
own
of
on
data from
a
area
for the
similar
for the
study
Canton
Heteroptera
sites investigated from a
given.
conservation point of view, the species list is compared to data from other regions of
Switzerland (based on literature and on a comparable study which has been conducted
in the Canton Luzern). The results indicate that the extensively managed meadows of
the Schaffhauser Randen are relatively very species rich habitats for the heteropteran
conducted in the Randen
Schaffhausen
an
To
is
evaluate
the
value
fauna of Switzerland.
Chapter 4 explores how host plant diversity and food quality affect the larval
development of two bug species (Stenodemmi) which arc strictly phytophagous and
capable of feeding on a wide range of different grass species, but arc normally found on
only a few species. They change their host plants during ontogenesis and it has been
suggested that this represents a response to the changing protein content of the hosts. In
a laboratory experiment the development of insects reared on grass monocultures was
compared with that when they were reared on mixtures of four species. In addition the
host grasses were grown under two nitrogen regimes to test if protein content is the key
factor determining host plant switching. The results indicate that host plant diversity
significantly increases larval survival and that crude protein content is not the main
factor responsible for the effect. Other important factors might be the form and
availability of nitrogen, the balance of amino acids, the water content of the host and the
content of mineral and trace elements. Furthermore, secondary compounds and
mechanical or chemical plant defences may also play an important role. A highly
diverse meadow might therefore enhance insect diversity by increasing larval survival
of herbivores and, indeed, the field data indicates that host plant diversity is positively
correlated with the
species
richness of Stcnodemini.
3
Zusammenfassung
Diese
Arbeit untersucht den Einfluss
struktur auf die Insektenvielfalt
von
von
Nutzungsintensität
und Landschafts¬
Mähwiesen. Das Ziel der Dissertation ist aufzuzei¬
gen, welche Umweltfaktorcn die Tnsektenfauna
von
Mähwiesen bestimmen und wie die
Agrarlandschaften gefördert werden kann. Als Untersuchungsobjekt
gewählt worden, weil sie in der Schweiz einen grossen Teil der landwirt¬
schaftlich genutzten Fläche ausmachen und ein wichtiges Element der Kulturlandschaft
darstellen. Die vier Kapitel der Arbeit behandeln folgende Fragestellungen: 1) Welchen
Einfluss hat die Bewirtschaftungsintensität auf die Tnscktenvielfalt? Können für die ver¬
schiedenen Wiesentypen spezifische Lebensgemeinschaften definiert und charakterisiert
werden? 2) Wie viel tragen die verschiedenen Wiesentypen zur Artenvielfalt auf Land¬
schaftsebene bei? Wie beeinflusst die umgebende Landschaft die Tnsektenvielfalt einer
Wiese? 3) Wie wertvoll sind die Wiesen des Untersuchungsgebietes für die Wanzen¬
Tnscktenvielfalt in
sind Wiesen
fauna der Schweiz?
4) Welchen Einfluss hat die Pflanzcnvielfalt auf die Insektenvielfalt
einer Wiese? Hat die
Wirtspflanzenvielfalt
einen Einfluss auf die
Entwicklung oligo-
pbagerInsekten?
Indikatorgruppe für die lokale Wanzenvielfalt wurden Wanzen (Heteroptera)
gewählt, weil sie ökologisch sehr vielfältig sind und weil Wiesen typische Habitate die¬
ser Tnsektenordnung darstellen. Die Untersuchungen wurden im Schaffhauser Randen
durchgeführt, dem östlichsten Ausläufer des Tafeljuras. Die landwirtschaftlich genutz¬
ten Gebiete auf den Hochflächen sind durch ausgedehnte Wälder voneinander getrennt.
Vier solche Gebiete wurden ausgewählt, die sich in ihren Anteilen an Gras- und Acker¬
land unterscheiden. Als Versuchsanordnung wurde ein Blockdesign gewählt, mit zwei
Nutzungsintensitäten (extensiv und mittel intensiv) pro Gebiet und drei Wiederholungen
pro Gebiet und Nutzungsintensität. Insgesamt wurden 24 Flächen erfasst.
Als
Im
Kapitel
1 wird
gezeigt,
dass die
Bewirtschaftung
die Wanzen vi elf alt einer Wiese
stark beeinflusst. Extensive Wiesen sind diverser als mittel intensive und weil die Wan¬
Nutzung reagieren, wird auch die Zusammensetzung
der Lebensgemeinschaften
Nutzung verändert. Die Reaktionen der häufigsten
Arten werden mit Hilfe von Varianzanalysen geprüft. Zwei Alten reagieren positiv auf
die intensivere Bewirtschaftung und sind in mittel intensiven Wiesen häufiger als in
zenarten
unterschiedlich auf die
durch die
extensiven. Beide Arten entwickeln mehrere Generationen pro Jahr und sind relativ
bil,
was
ihnen das
Wanzenarten
Überleben
in stark
ermöglichen scheint. Sechs
beeinträchtigt und sind
Bewirtschaftung
gestörten Habitaten
werden durch die
mo¬
zu
klar
hingegen
häufiger als in mittel intensiven. Viele dieser Arten überwintern
und beginnen ihre Entwicklung erst im Spätfrühling. In früh und häufig
in extensiven Wiesen
im Eistadium
geschnittenen Wiesen können sie ihre Entwicklung deshalb nicht abschhessen und ver¬
schwinden längerfristig aus diesen ITabitaten. Wanzenarten die am Boden leben und für
ihre Entwicklung auf ein warmes und trockenes Mikroklima angewiesen sind, werden
ebenfalls durch eine intensive Nutzung beeinträchtigt. Mittel intensive Wiesen zeichnen
sich durch eine dichte und geschlossene Vegetation aus und sind deshalb für diese Tiere
ungeeignete Habitate.
Die Wanzenfauna der extensiven Wiesen unterscheidet sich stark zwischen den vier
Gebieten. Die mittel intensiven Flächen hingegen gleichen sich stärker in ihrer Artenzahl und -Zusammensetzung. Viele Wanzenarten kommen
und scheinen eine
können
zwar
nur
in einzelnen Gebieten
vor
kleinräumige Verbreitung zu haben. Die meisten der seltenen Arten
häufig auftreten, sind aber auf einzelne Flächen beschränkt. Es
lokal
4
braucht deshalb viele verschiedene Wiesen
Region
zu
erhalten. Im
unterschiedliche
sen
Kapitel
Strukturierung
daraufhin, class der Anteil
zenfauna
um
das ganze
Spektrum
an
Arten in einer
2 wird untersucht, ob die Gebietsunterschiede durch die
der Landschaft erklärt werden kann. Die Resultate wei¬
an
Ackerland in der
umgebenden
Landschaft die Wan¬
Wiese beeinflussen kann und dass gewisse Arten durch diesen Habitat¬
einer
typ gefördert werden. Es hat sich aber auch gezeigt, dass die gewählte räumliche Skala
zu grob ist, um verstehen zu können, wie Insekten durch die umgebende Landschaft
beeinflusst werden.
aufgrund dieser Arbeit eine aktualisierte Artenliste für den Kanton
Schaffhausen vorgestellt. Dafür werden Daten einer Studie einbezogen, die ähnliche
Fragestellungen bearbeitet und im gleichen Untersuchungsgebiet durchgeführt wurde.
In
Kapitel
3 wird
Um den Wert der untersuchten Wiesen für den Artenschutz abschätzen
die Artenlistc
mit
Daten
aus
Literaturdaten und Resultate
Luzern untersucht hat. Der
ser
anderen
einer
Regionen
Arbeit
der Schweiz
verwendet,
Vergleich zeigt,
zu
verglichen.
können, wird
Dafür werden
die extensive Wiesen im Kanton
dass die extensiven Wiesen des Schaffhau-
Randens wertvolle Flabitate für die Wanzenfauna der Schweiz darstellen.
Kapitel 4 wird untersucht, ob die Vielfalt von Wirtspflanzen die larvale Ent¬
wicklung phytophager Wanzen (Heteroptera: Stcnodemini) beeinflusst und ob der Roh¬
proteingehalt der Nahrung dabei eine entscheidende Rolle spielt. Die Larvcncntwicklung von Tieren, die mit Mischungen aus vier verschiedenen Pflanzcnarten gefüttert
In
worden sind, wird mit
derjenigen
von
Insekten
verglichen,
denen reine Monokulturen
an Wirtspflanzen die lar¬
zeigen,
vale Mortalität senken und dass der Proteingchalt der Pflanzen nicht alleinc dafür ver¬
antwortlich sind. Die vorteilhafte Wirkung eines vielfältigen Pflanzenangebots für her¬
gefüttert
worden sind. Die Resultate
dass eine Vielfalt
bivore Insekten scheint demnach durch verschiedene Faktoren verursacht
wie zum
Beispiel
oder die unterschiedliche
fältiger
Zusammensetzung
und
Verfügbarkeit
von
werden,
Stickstoff. Ein viel¬
Pflanzcnbestand kann daher die Tnscktenvielfalt fördern, indem
Entwicklung
zu
sekundäre Pflanzenstoffc, chemische oder mechanische Abwehrstoffe
er
die larvale
der Insekten positiv beeinflusst und die Mortalität senkt. Felddaten unter¬
stützen diese Resultate und /eigen, dass die Artenzahl
Artenzahl ihrer
Wirtspflanzen
zunimmt.
oligophagcr
Herbivoren mit der
5
General Introduction
species rich landscapes of Central Europe were generated
during the Middle Ages by traditional small scale and extensive management
(Mühleberg & Slowik 1997). They are characterised by both temporal and spatial
heterogeneity (Macdonald & Smith 1990). In the last fifty years, however, agricultural
practices have changed and strongly altered agricultural landscapes. The mechanisation
of cultivation and the use of artificial fertiliser and pesticides have greatly increased
agricultural production but at the same time led to an impoverishment of the landscape
The
heterogenous
and
heterogeneity and the use of
pesticides resulted in a major decrease m species diversity (lsler-Htibscher 1980;
Mühlcbcrg & Slowik 1997). These negative effects have led to growing public concern
about the environmental problems caused by modern agricultural practices, and to an
increasing demand for food produced in an ecologically sensitive way. Further, the
and to environmental
Convention of Rio in the
biodiversity
and the
The decrease
pollution.
1990's resulted
m
m
habitat
greater
awareness
about the issue of
negative consequences of species loss. In parallel major changes
have occurred in world
which have
economics
greatly
altered the rural
economies
of
Europe. In many countries the time had come for major changes m
agriculture policy. For example, the Swiss government introduced subsidies for
extensive management and ecological compensation areas; these were intended partly to
enhance species diversity m agricultural landscapes and partly to support Swiss
agriculture without causing damaging effects. The new policy led to a great interest in
management techniques to enhance biodiversit> m agricultural landscapes and they
of
most countries
were
therefore
National
investigated
Science
in the
Foundation.
Priority Programme
This
thesis
is
Environment
(SPPE) of the Swiss
of that programme
part
and has
the
objectives, firstly, to examine how management and landscape structure affect insect
diversity of agricultural grasslands and, secondly, to evaluate management techniques to
enhance species diversity m agricultural landscapes.
investigated the influence of different grassland management
types on the species diversity of insects (for an overview see Curry (1994) and
Gerstmeier & Lang (1996)). Although landscape structure may influence the effect of
management on species diversity, most studies have focused on individual sites, without
taking into account the surrounding landscape (Saunders, Hobbs & Margules 1991).
However, landscape structure can be important: m a mosaic of different grassland types
Several studies have
insects have
Furthermore,
better chance to find suitable habitats, shelter
a
the
grassland types
are
destructive
present,
effect
since
of
mowing
insects may
may
escape
to
which investigates the influence of management and
or
food
over
the
when
season.
different
mitigated
adjacent meadows. This thesis,
landscape structure on insect
be
diversity of grasslands, is concerned with both the local system and with the influence
of the surrounding landscape. A survey was made of insect diversity in grasslands under
extensive
and medium
intensive
indicator group for insect
management
the North
of Switzerland.
As
an
bugs (Heteroptera) were chosen because
ecologically
including phytophagous, saprophagous and
they
predatory species (Dollmg 1991). Furthermore, some species arc generahsts while other
are specialists. Both larval stages and adult individuals live m the same habitat and
respond sensitively to environmental changes (Morns 1969, 1979; Achtziger 1995; Otto
1996). Previous studies have shown that the richness of the bug fauna correlates
strongly with total insect diversity (Duelh & Obnst 1998). Finally, it is a manageable
are an
diversity
the true
m
very diverse group,
group in terms of numbers of species.
6
Only few studies have investigated the dispersal and dispersion of arthropods in
agricultural landscapes, though dispersal ability is a key factor determining how species
are affected by management and the surrounding landscape (Samways 1994; Forman
1995; Mortimer, Hoi her & Brown 1998). The sink-source model (Pulliam 1988; Howe
& Davis 1991; Watkinson & Sutherland 1995) and the mctapopulation concept
(Harrison 1993;
Reich & Grimm 1996; Han ski
1997)
are
theoretical concepts which
recognise the essential role of dispersal for long term persistence of organisms in patchy
habitats. Apart from Southwood (I960), few authors have examined the dispersal ability
of heteropteran bugs. A few pest species have been thoroughly investigated, but
otherwise only little is known about the ecology of species in this insect order (Brown
1965; Dingle 1968; Dmglc & Arora 1973). We designed a field experiment to
investigate how rapidly new habitat patches are colonised by bugs and which species
disperse
most
readily.
Although, the relationship between species richness of insects and plants has been
investigated in several studies, the results are contradictory and the underlying
mechanisms are not yet entirely understood (Strong, Lawton & Southwood 1984). The
relationship is dependent on various factors, including the age of a habitat and the stage
of succession. Early in a succession there may be a positive correlation between insect
and plant diversity, but later the structure of the vegetation may become a more
important factor than plant species numbers influencing the species richness of insects
(Southwood, Brown & Reader 1979). Tn this thesis the relationship between species
richness of plants and heteropteran bugs was investigated using the insect data from the
field study and in close collaboration with Sibylle Studer (2000) who studied the
vegetation of the same sites. In the laboratory the influence of plant diversity on a guild
of plant feeding bugs (Heteroptera: Stenodemini) was investigated experimentally. The
effect of host plant diversity and food quality in terms of crude protein content on larval
development and survival of two bug species {Leptopterna dolobrata L. and Notostira
crratica L.)
Outline
was
studied.
of this thesis
grassland management on the heteropteran
2 explores the species patterns on the landscape level and compares them
with those of plants and butterflies, which were investigated by Studer (2000) and
Widmer (2000). In addition, the influences of the surrounding landscape on
heteropteran bug diversity of grasslands are discussed. Chapter 3 analyses the faunistic
data gained in the field study in terms of the abundance and ecological affinity of the
various species and discusses the biogeography of rare species in respect to data from
Germany. Chapter 4 presents the results of the laboratory experiment on the effect of
host plant diversity on larval survival of two phytophagous bug species.
Chapter 1
fauna. Chapter
examines the influence of
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Chapter 1. Enhancing insect diversity
management and landscape structure
in
The roles of
Manuela Di Giulio, Peter J. Edwards & Erhard Meister
Accepted by Journal of Applied Ecology
Summary
1.
During
to
preserve and
of this
variety of methods have been applied in Switzerland
enhance biological diversity in agricultural systems. The purpose
to evaluate grassland management techniques in respect to their
the last few years
study
is
a
managed area itself and
species diversity at a landscape scale.
effectiveness within the
tribute to
to examine
how these
areas con¬
heterogenous landscape in part of the Swiss Jura we examined insect diver¬
sity in grasslands subject to different management. Four study areas with varying
landscape structure were selected and in each area, meadows of two grassland
management types were investigated.
2.
In
3.
The true
4.
The variance of the
a
bugs (Heteroptera) have been chosen as an indicator group for insect di¬
versity on the basis of previous work which had shown that the richness of the
bug fauna correlates strongly with total insect diversity.
heteropteran species
data is
management components. Area accounts for
the interaction management
diversity
is
greater
m
x area
more
35.4%,
for 7.2% of the
extensively managed
ones; extensive sites have
partitioned
into
spatial (area)
and
management for 29.7% and
species
variance. The
species
meadows than in medium intensive
individuals and show
a more even
rank abun¬
dance distribution.
5.
differ in their responses to management. Two species benefited
from intensification whereas six species were affected negatively by intensive
Individual
species
management. The influence of management
on
individual
species
is discussed in
species do not appear to respond to management;
these are mostly widespread species which occur in various types of habitats, and
polyphagous species which live in a wide range of grasslands but which show a
certain affinity to managed meadows.
detail. Two main groups of
6.
grasslands can enhance both
local and regional insect diversity in agricultural landscapes. Extensively man¬
aged meadows are species rich habitats which support some rare and specialised
species. In contrast the bug community of medium intensive meadows is domi¬
nated by more widespread and less specialist species.
Our
study
indicates that extensive management of
Key-words: grassland management, Heteroptera, landscape structure, species diversity,
Switzerland
INTRODUCTION
Agricultural landscapes are characterised by a very high spatial and temporal het¬
erogeneity which is determined largely by human activities. Little is known about how
species are affected by such heterogeneity and how they persist in agricultural systems
(Macdonald & Smith 1990). For example, few studies have investigated the dispersal
9
agricultural landscapes (Samways 1994; Forman 1995;
Mortimer, Hollier & Brown 1998), though there is information on some groups, notably
butterflies (Fry 1994; Sutcliffe & Thomas 1996), spiders (Bishop & Riechert 1990;
Thomas, Hoi & Everts 1990; Thomas 1996), carabid beetles (Thomas, Wratten &
Sotherton 1991; Kinnunen & Tiainen 1994; Forman 1995) and wild bees (Steffanof
dispersion
and
Dewentcr
in
arthropods
1998).
grassland
In contrast, there have been several studies of the influence of
manage¬
and Gerstmeicr &
Lang
Curry (1994)
example mowing, have a direct effect on in¬
sects by damaging or killing individuals or removing them from the site. Immobile and
relatively immobile species or stages are specially affected, and in the long term the
whole population may be threatened (Volkl et al. 1993). Indirect effects may also occur
mainly through changing the habitat; for example, if management alters the plant spe¬
cies composition and the structure of the vegetation, it is also likely to affect the micro¬
climate and other aspects of the microhabitat. Since many arthropods respond very sen¬
sitively to the microclimatic conditions such indirect effects can strongly influence
community composition (Franz 1931; Fewkes 1961: Curry 1994; Gerstmeicr & Lang
insect
ment on
(1996).
diversity;
for
an
overview
sec
Some management operations, for
1996). Lcafhoppers (Auchenorrhyncha) arc strongly affected by indirect effects of
grassland management. They are closely associated with particular plant species and
reach their greatest diversity in species-rich, highly structured meadows. Many species
the sward during the
occupy definite layers in a grassland and move vertically within
season. Changes in the species composition and in the structure of the plant community
due to management therefore affect the community of lcafhoppers greatly (Andrzcjewska 1965;
Curry 1994).
Although it is recognised that landscape structure also affects the species composi¬
tion at a particular site, most studies of management have focused on the site alone,
without taking into account the surrounding landscape (Saunders, Flobbs & Margulcs
1991). However,
over
the
course
insects have
of
a season
a
in
a
better chance to find suitable habitats, shelter
can
escape to
1995)
grassland types. Furthermore, the
grassland types arc pres¬
where they find new habitats. Indeed
when different
mitigated
adjacent meadows
1991; Watkinson & Sutherland
Reich & Grimm 1996; Hanski
1993;
metapopulation concept (Harrison
the sink-source model
and the
food
mosaic of different
destructive effect of mowing may be
ent, since insects
or
(Pulliam 1988;
Howe & Davis
dispersal for long
Moreover,
species differ in their
patchy
organisms
persistence
responses to management, it is important to support a high diversity of management
types in order to maintain and enhance species diversity at a regional level (Morris &
Rispin 1993; Dietl 1995).
1997)
are
theoretical concepts which
of
term
m
recognise the
habitats.
essential role of
since
study on the effects of both management and landscape
species diversity of agricultural grasslands in Switzerland. We chose the
true bugs (Heteroptera) as an indicator group for insect diversity in general for various
reasons. Firstly, the bugs are an ecologically very diverse group, including phytopha¬
gous, saprophagous and predatory species (Dollmg 1991). Furthermore, some species
are generalists while other are specialists. Secondly, both larval stages and adults live in
the same habitat and respond sensitively to environmental changes (Morris 1969, 1979;
Achtziger 1995; Otto 1996). Thirdly, previous studies have shown that the richness of
the bug fauna correlates strongly with total insect diversity (Duelli & Obrist 1998). FiThis paper is part of
structure
on
a
10
ecological diversity, the Heteroptera
of species occurring in grasslands.
their
nally,
and
terms
of the numbers
despite
is
a
manageable
group in
developed to enhance biological diver¬
sity in agricultural systems. The aim of our study is to evaluate how contrasting grass¬
land management techniques influence insect diversity in agricultural landscapes and to
assess to what extent insect diversity of grasslands is affected by landscape structure. In
this paper we focus on the effect of management on the species composition of the het¬
In Switzerland different schemes have been
eropteran fauna. The effects of landscape
position
Giulio,
of
in
in
a
separate paper (Manuela Di
and methods
area
and
The research
ble
presented
communities will be
preparation).
Material
Research
heteropteran
structure on the relative abundance and com¬
1) and
study
area
sites
is located in the Schaffhauser Randen
of the Swiss
forms part of the hills
(northern Switzerland,
Jura. The soils
are
Ta¬
nutrient poor
yearly precipitation in Schaffhausen (437 m a.s.l.) is 866 mm,
and the highest mean monthly are m summer and lowest in late winter. The mean tem¬
perature is 7.8 °C, with a maximum mean monthly temperature in July (23.2 °C) and a
limestones. The average
January (-3.9 °C) (SMA 1998).
minimum in
gion with several
more or
The
study
site is
agricultural
by offering
policy
follow
specified
Table l.The four
given
in
areas are
meters
re¬
agricultural
they
subsidies if
grassland management.
situated
above
extensive forested
land. Local
encourages farmers to maintain extensive meadows
forms of
an
sea
in
different
level. The
municipalities
area
of the Canton Schaffhausen. The altitude is
of the mdn idual enclaves has been estimated based
on
1:5000 maps.
Community
Area
ALTITUDE
Area op
COORDINATES
Enclave
Zeigli/Mösli
Hemmental
800
30 ha
8° 34'E/47° 45'N
Hinterranden
Hemmental/
830
40 ha
8° 34'E/47° 44'N
670
100 ha
8° 36'E/47° 44'N
780-830
240 ha
8° 36'E/47° 46'N
Siblinçen
Klosterfcld
Hemmental
Merishauscr Randen
Merishausen
study we chose a nested block design with two levels. At the block level
(landscape type) we selected four enclaves (= areas) ranging in size from 30 to 240 ha
(area) with varying proportions of arable and grassland habitat. Within each block (area)
we investigated two grassland management types (officially referred to as medium in¬
tensive and extensive), and for each management type wc selected three replicate sites.
The management types are defined by two main factors: the number of cuts per year,
and the type and amount of fertiliser used, Medium intensive meadows are regularly
fertilised with slurry and are cut 2-3 times per year. The time of the first cut is approxi¬
mately the end of May. The extensive meadows are not fertilised and arc cut once or
twice per year. The time of the first cut is July. We selected the grassland types on the
basis of the plant species composition and later we interviewed the farmers about the
amount and type of fertiliser used, the number of cuts, the time of cutting and the previFor
our
11
management. In these interviews
ous
lected medium intensive
grasslands
we
had
learned that in the
been
only
the other
in
managed
the last few years. The management contrast in this
of Hinterrandcn the
area
area was
se¬
less intensive way for
a
therefore smaller than in
areas.
plant species composition was assessed in summer 1997 using six randomly lo¬
cated Ixlm2 quadrats per site. A total of 110 plant species were recorded in the 24 sites,
with the number of species sampled per site ranging from 21 to 48. The extensive and
medium intensive meadows showed clear differences in plant species richness and in
species composition. The community of the extensive meadows was dominated by
The
Bromus
erectus
and there
The medium intensive sites
and the
vescens
heteropteran bugs
diameter of 40
cm
net over
sulting
a
were
was
were
in five
subsamples
also restricted to the
was
from
September.
May
to
The adult insects
a
(Studer 2000).
m.
The net
per site at each
were
period
good,
were
two
was
summer
ie
a
by making
weeks
using
of 1997
cloth suitable for
a
standard¬
The sweep-net had
sampling
insects in
100 sweeps with the
after every twentieth sweep,
emptied
date.
sampling
Sampling
was
only
carried out
between 10.00
Every
a.m.
site
and 17.30 p.m.; the
Sampling
sampling order
was
determined with the aid of
sampled
between 7 and 9 times
entomological
handbooks
1952; Wagner & Weber 1964; Wagner 1966. 1967, 1970-1971, 1975). The
R. Heckmann
Statistical
All
and Ch.
(1990).
species
Rieger (Germany).
determinations
were
(Wag¬
nomen¬
checked
Analysis
For the
sample
(Germany)
re¬
minimum of 17°C and sunshine.
clature followed Günther & Schuster
by
the
heavy
collected every
varied between weeks.
of the fields
ner
30
sampled during
fitted with
distance of about 100
when the weather conditions
was
was
(Remane 1958; Bornholdt 1991; Otto 1996).
and
vegetation. Samples
the
number
species
ised sweep-net method
a
were
dominated
Methods
Sampling
The
mean
species number of 42 in the sample of 6 quadrats.
by Arrh en ath en un elatius and Trisetum fla-
was a mean
analysis
of the data, all
samples
were
per meadow. Rank abundance distributions
pooled
are a
over
time, resulting in
one
useful method to describe and
account both
species
bug
communities of the two management types. Canonical Correspondence Analysis
(CANOCO, version 3.1 by ter Braak 1987-1992) was used to study the influence on bug
communities of management factors and spatial effects. The variance of the bug species
data was partitioned into spatial (area) and management components (Borcard, Legendre
& Drapeau 1992). Number of cuts, time of the first cut, and number of plant species
sampled per meadow were included in the model as management variables. The four ar¬
eas investigated (enclaves) were included as a spatial factor. The apparent species com¬
position of communities is strongly influenced by sample size and rare species tend to
be recorded only in large samples. To minimise sampling effects we omitted both rare
species (< 3 individuals) and sites that were sampled fewer than eight times. The latter
compare communities
(Magurran 1988) because they take into
number and their abundance. We used rank abundance distributions to compare the
concerns
24 most
three sites in the
common
area
Merishausen and
species (> 14 individuals)
amine how individual
species respond
to
one in
analyses
the
area
of variance
management.
Hinterranden. For the
were
conducted to
ex¬
12
Results
Influences of grassland management
on
insect
diversity
bugs comprising 93 species characteristic of meadows were
recorded in the 24 sites (occasional vagrants from wooded habitats were excluded). A
full species list will be presented elsewhere (Di Giulio, Beckmann & Schwab 2000).
A total of 5608 adult
In three of the four
strate
clearly
areas
(Figs
la to
lc),
a
very
communities of the extensive meadows demon¬
while in the fourth
The overall pattern
(Fig. Id).
high abundance
structures are similar
cies to have
bug
rank abundance distribution than those of the medium inten¬
a more even
sive meadows
the
while the
area
the
community
the intensive sites is for
in
majority
sive meadows. This contrast is clearest within areas,
from
(Hintcrranden)
arc
as
a
few spe¬
less abundant than in exten¬
the communities vary
strongly
area to area.
1000
1000
b)
r
c
100
Xs
10
extensive
<
~U—
0
10
20
30
40
50
10
60
20
1000
1000
n
O
100
S
100
cd
O
Ö
3
10
extensive
extensive
10
<
<
intensive
20
10
30
40
managed
Hinterranden. Three
m
different
sites
30
Rank
I. Rank abundance distributions of
meadows
20
10
Rank
Fig.
d)
c)
O
d
40
Rank
Rank
Ö
30
hctciopteran
areas:
extensively and medium intensively
Zelgli/Mösli, c) Mcnshauser Randen, d)
have been pooled for this analysis.
communities of
a) Klosteifeid, b)
per management type and
area
explains 72.3% of the total variance (total inertia: 1.578, sum
eigenvalues: 1.141, Monte Carlo permutation test: P 0.01; Fig. 2). Of
this, area accounts for 35.4% (eigenvalue 0.559, P
0.01) of the species variance, man¬
and
P
for
29.7%
the interaction management x area
0.01),
(eigenvalue 0.468,
agement
for 7.2%. The plant species number was included into the variable 'management' and
we therefore cannot say how much variation if explains.
The ordination model
of all canonical
=
=
=
13
1UU 10
27 7
O
2
50%
35 4!
iummmttl
u
cd
>
D
unexplained
il
area
D management
7.2
\
area
H management
A(V„
Fig
2.
L
Paititioning
^^NW,\\NNy.W'
by en\ispecifications)
of the species
vanance
lonmcntal vatiables (see tevt foi
respectively) of the ordina¬
tion analysis (Table 2, Fig. 3) account for 47.2% of the species variance, while the third
and fourth axes (eigenvalues 0.158 and 0.077 respectively) for 14.9%. The ordination
diagrams show that the axes one and two separate the area Klosterfeld from Hinterranden, and also the extensively managed meadows (July cut and one cut) from the more
intensively managed ones (Fig. 3a). The extensive meadows are grouped in the upper
part of the diagram of axes one and two, while the medium intensive ones are found m
the lower part. The medium intensne sites are closely gioupecl, indicating that their het¬
The first and second
eropteran communities
separated
are
(eigenvalues
very similar.
from those of the other
rated from the other
edly
axes
areas
different species
by
areas
axes one
0.554 and 0.191
Further, the extensive sites of Klosterfeld
by
the first
and three,
axis.
Hmterranden
indicating
is
clearly
that these sites have
a
are
sepa¬
mark¬
compositions (Figs 3a and b). The plant species number is the
only non nommai variable m the model and is correlated
though the correlation is not very strong (Table 2).
with the first and second axes,
14
N
b)
rpi
M
x
O
4-
Z*
z. «z
number of
plant species
K1
Km
2 cuts
#H
*,'Klosterfeld
Hjfi,K
ff
axis
.Zelgji
Hinterranden#
+0 7
Hinterr.randeTr
,
u
M*
-0.7
1
.
1
#
3 cuts
\<-
cnvhonmcnlal vaiiablcs
-,.»Z
Z*
given for
j^Zelgli
axis3i
»M
40 7
Z
Z»
»June
»
Z
K*
k
»K
June
»Z
ï3 cuts
9 -«K
Fig. 3. Ordination diagrams based on
represented as points (centroids),
are,
•
Merishausen
•
K«w 3 cuts
1 cut
,tv'
C
•H
Mayj,i
2 cuts
i)t
Klosterfeld-,1
h»»H
a
^
»M
-0 7
2 cuts
»
c
Juy
a
Canonical
non
Correspondence Analysis (CCA).
nominal variables
the number of cuts per year. The
capital
as
vectors in the
Nominal variables
diagram.
letters indicate the
areas:
Different
Z:
arc
symbols
Zelgli/Mösli,
K:
species compositions and
community structures are represented by points close to each other in the diagrams. Total inertia:
1.578, eigenvalue first axis: 0.554, eigenvalue second axis: 0.191, eigenvalue third axis: 0.158, eigen¬
Klosterfeld. M: Merishauser Randen. H: ITinterranden. Sites with similar
0.01. For
was tested using a Monte Carlo permutation lest: P
analysis four sites (three in Merishauser Randen and one in ITinterranden) were omitted because
they were sampled less frequently than the other ones (see text for specifications).
value fourth axis: 0.077. The model
=
this
Table 2. The correlation matrix is based
the first four environmental
on a
Canonical
Correspondence Analysis
and shows the values of
axes.
Env. ax. 4
Env. ax. 2
Klosterfeld
0.8807
0.2660
-0.2635
-0.2660
Area 2: Zelgli/Mösli
-0.1488
0.0289
0.7723
0.0133
Area 3: Hjnerranden
-0.6090
-0.3265
-0.7059
0.1146
Area 4: Merishauser Randen
-0.0983
0.0814
0.3996
0.1649
1
-0.2656
0.6369
0.2167
-0.0080
2 CUTS
-0.0469
-0.3609
-0.2097
0.3909
3 CUTS
0.5121
-0.4222
0.0022
-0.6430
May cut
0.3777
-0.4323
0.2351
0.1534
June
0.4391
-0.4653
0.1505
-0.2255
July cut
-0.6070
0.666
-0.2832
0.0646
Number
-0.4240
0.4058
-0.001
0.0170
Area 1
:
CUT
cut
of plant species
Env.
3
Env. AX.l
Variables
ax.
15
There is
no
significant
opteran bugs (t2i
38
=
0.96, P
correlation between the
=
0.35, R2
species
0.04; Fig. 4).
=
number of
and heter¬
plants
r
34
(1)
O
en
30
•
?fi
-
Pw
on
ci
X>
4-1
22
o
*-<
(1>
1«
x>
H
s
!z;
14
-
10 L
18
'
22
34
30
26
Number of
Fig.
for
a
vegetation
tot the
Species
is
46
50
plant species
plants and heteropteran bugs are not correlated.
grassland site and is based on 6 Ixl m2 quadrats
recorded data for the bugs.
number of
species
4. The
Each point
42
38
different
and the
response to management
species level, eight out of 24 species are significantly influenced by manage¬
ment. Because the timing of the first cut and the number of cuts arc not independent
variables we cannot be certain which factor is more important for the bug community.
However, one or the other factor usually correlated more strongly with the occurrence of
At the
species.
Only
are
Nabis
species appear
pseudoferus (Fijo
two
0.05) which
are
evidence that N.
Six
ceraea
species
=
3.55, P
most abundant m
pseudoferus
arc
recticornis
=
<
5.39,
by cutting
affected
P
=
intensive, frequently
is reduced
adversely
(F2,io
management: these
4.57, P <
0.05) and Notoslira erratica (F?.,2o
to benefit more from medium intensive
<
by
0.05),
in
cut meadows
July (7*2,20
=
(Fig. 5).
9.2, P
intensive management. Of
Peritrechus
<
There is
0.01; Fig. 6).
these, Mégalo-
gracilicornis, (F?„20
=
13.36, P
<
3.96, P < 0.05) are apparently reduced by fre¬
biguttatus (F220
the
of
the
by
timing
cutting. These species occur most abundantly
in extensive meadows (Fig. 5). The abundances of Adelphocoris seticornis (Fo,2o
5.0,
0.0
P < 0.05; F220
P
P
Hadrodemus
<
<
0.001; F2,2o
8.19,
L),
13.29,
m~jlavum (F2ao
13.95, P <
4.6, P < 0.05) and Leptopterna dolobrata (F7 2o
9.26, P < 0.01; F220
0.001) appear to be reduced by both frequency of cutting and by early cuts (Figs 5 and
0.001)
and Adomerus
=
quency of cut but not
=
=
=
6).
=
=
=
16
N.
pseudoferus
t
rf-i
m
u
2
3
1
a
fi
ab
S
i
_
—-
—
2
1
3
M. recticornis
ii
ri
0)
2
1
3
F. dolobrata
m-flavum
/F
a
b
b
-
ab
A. seticornis
Af. erratica
r
a
O
b
fi
cd
b
Ä
It?
-u-
fi
3
2
1
P.
2
1
A.
gracilicornis
i
3
biguttatus
5
b
21
b
b
5.1
-1
Number of cuts
Fig. 5. Responses
of individual species
to
the number of cuts
(mean
± sd). The
pairwise comparison
was
Tukey test. Significant differences between the number of cuts are marked with dif¬
3.55. P < 0.05) and V. erratica (F2,20
ferent letters (a/b). N. pseudoferus (F220
4.57, P < 0.05) are
5.0, P <
most abundant in frequently cut sites. In contrast the abundances of A. seticornis (F2.2o
9.26, P < 0.01), M. recticornis
0.05), H. m-flavum (7A20 =13.29, P < 0.001), L dolobrata (F2,20
13.36. P < 0.001) and A. biguttatus (F2,20
3.96, P <
5.39, P < 0.05). P. gracilicornis (F220
(F2io
conducted with
a
=
=
=
=
are
=
=
=
0.05)
by frequent cutting.
reduced
A.
6
A. seticornis
pseudoferus
a
4
a
a
2
b
0
LXl
!
A
4-
ii
DJ
May
June
X
-2
July
June
May
July
o
G
H.
ö
L, dolobrata
m-flavum
<
^t
x
LI
May
June
July
May
June
July
Time of the first cut
6.
Responses of
species to the time of the first cut (mean ± sd). The pairwise comparison
Tuke) test. Significant differences between the, time of cut are marked with dif¬
ferent letters (a/b). July cut reduces the abundance of N. pseudoferus (F220
9.20, P < 0.01). May and
June cut reduces the abundances of A. seticornis (F12Q
P
H.
<
0.01).
8.11,
m-flavum (F2.20 4.6, P <
Fig.
was
individual
conducted with
a
=
=
0.05)
and L. dolobrata (F
,n
=
13.95, P
<
0.001 ).
~
17
Sixteen
show
species
no
response to management
These
intensity.
are:
Lygus
pra¬
Carpocoris fuscispinus, Berytinus minor, Polymerus unifasciatus, Sirongylocoris steganoides, Piagiognathus chrvsemthemi, Stenodema laevigatum, Trigonotylus caelestialum, Adelphotensis, Lygus ntgulipennis, Dolycoris baccarum, Eurygasler
coris
maura,
lineolatus, Catoplatus fabricii, Nabis brevis, Chi amy dolus pulicarius and Haïtiens
apterus.
Discussion
Responses of the bug community
presented here show that management effects are clearly important (Figs 2
and 3) for the species composition of the heteropteran bug community. They account for
30% of species variance and the ordination diagrams indicate that the extensive and me¬
dium intensive meadows have differing species compositions (Figs 3a and b). The ex¬
tensively managed meadows show a high variation in species composition, especially
The data
between areas,
light
indicating
conditions
or
that apart from
management several other factors like aspect,
species composition of these sites. The medium
similar communities, suggesting that management is
isolation affect the
intensive sites, however, have very
important factor influencing the species composition of these meadows. Fur¬
ther, species diversity tends to be greater in extensively managed meadows than in me¬
the most
dium intensive
more even
Tn three
ones.
areas
the extensive sites have
rank abundance distribution
(Figs
la to
ever, is different. The rank abundance distributions
grams
(Figs
3a and
b)
One
1c).
for Hinterranden show that the
area
a
(Hinterranden), how¬
and the ordination dia¬
(Fig. Id)
bug
individuals and show
more
communities
are
very similar
types. As already mentioned, the medium intensive grasslands have
in both management
vegetation does
species composition re¬
clearly
changes
mains typical of medium intensive and extensive sites elsewhere (Sibylle Studer, un¬
published data); this suggests that the bug communities react more quickly than plants
been
managed
less
intensively
to
in the last few years.
in management
not reflect the recent
Interestingly,
the
and the
so
management changes (Mortimer, Hollier & Brown 1998).
explain most of the species
variance (72.3%). The interaction term is very small (7.2%) indicating that the spatial
and management variables are clearly separable in their effects (Borcard, Legendre &
Drapeau 1992). Spatial effects account for an important part of the total variation (35.4
The environmental variables included in the CCA model
%)
and have
a
strong effect
on
the structure and
assemblage
of
bug communities (Figs 2
(eg differences in for¬
and 3). It is not yet clear whether this is due to historical factors
mer
land use)
or to
present day differences in landscape
structure. However the obser¬
geographic distributions of ter¬
patchier,
species having restricted ranges than
is the case for terrestrial vertebrates and plants (Colwell & Cocldington 1994; Otto
1996). The possibility that the structure of the landscape surrounding a site interacts
with management effects on the insect community will be addressed in a more detailed
spatial analysis of the data using G1S,
vation is consistent with
various
restrial invertebrates tend to be
studies which show that
with
more
plants and het¬
eropteran bugs (Fig. 4) and only a weak relationship between plant species number and
the species composition of the bug fauna as represented m the CCA (Table 2, Figs 3a
and b). Our findings are supported by the study of Otto (1996) who showed that the het¬
eropteran fauna is significantly correlated with the structure of the plant community but
Interestingly,
there
is no
correlation between the
species
number of
18
strongly by the
structure and the microclimatic conditions in the grasslands than by plant species rich¬
ness per se. As many bug species are zoophagous or polyphagous herbivores the pant
species richness might play a less important role than the structural diversity of the
not with the
richness.
species
Apparently
the
bug
fauna is affected
more
habitat.
Responses of individual species
Although
the trends associated with management
are
clear, species do differ in their
N. erratica is
responses to management, and not all arc adversely affected. For example,
most abundant in medium intensive meadows (Fig. 5). This mirid bug has at least two
freshly cut
is not yet clear how this species survives the frequent disturbances by cut¬
be that it rccolomses meadows soon after the cut (Remane 1958; Morris
even
in
1969, 1979; Gibson 1980; Otto 1996). Bockwinkel (1990) has shown that
at
generations
meadows. It
ting.
It may
per year. It feeds
on
of the adults take shelter
portion
surprising
since the females
males, in contrast,
good
are
are
grass leaves and
m
adjacent
can
find its food
least
a
pro¬
meadows. If this is the case, then it is rather
flight muscles. The
(Kullenberg 1944; Doll¬
very weak fliers due to their reduced
fliers and
to be
seem
quite
mobile
ing 1991).
generations per year which
lives in various habitats eg grasslands, cereals fields and field margins (Remane 1958;
Southwood & Leston 1959; Pen cart 1987). It is a predatory species that feeds mostly on
N.
pseudoferus
smaller insects
is
a
macropterous nabid with
(Péricart 1987).
grassland
habitats and
(Figs
5 and 6).
seem to
cies with several
Macropterous
per year have
turbed habitats than those with
to
be the
case
more
nabids
be fast and strong fliers
generations
three
Little is known about its
benefit from intensive management and is
site
two to
only
one
a
it appears to
abundant in the medium intensive
are
ephemeral
general, insect spe¬
persisting in highly dis¬
often associated with
(Southwood I960).
better chance of
In
(Southwood 1960; Morris 1979). This
for both N. erratica and N.
pseudoferus.
and is
reduced
biology, though
In contrast to these
appears
species,
M.
abundant in extensive meadows
by cutting
high proportion of grasses (Morris 1979; Otto 1996). Lar¬
val development takes place in spring when the intensively managed grasslands are cut
for the first time. This phase is the most sensitive of the whole life-cycle and therefore
susceptible to disturbances. The life-cycle may therefore explain the sensitivity of the
recticornis is
(Fig. 5)
species
or
severely
fallow land with
to intensive
be slow to colonise
more
a
management. Moreover this species is
new
habitats (Southwood
a
poor flier and therefore may
I960).
very poor soils
(Kullenberg 1944)
development (McNeill 1973; McNeill
We might expect that intensively managed meadows with a high
&
nitrogen availability would be a favourable habitat for this species. However, frequent
and early cutting reduces this species severely (Figs 5 and 6) and the life-cycle may be
the critical factor. Since the females lay their eggs on the bottom part of grass-stems
(Kullenberg 1944), damage to the eggs is probably not the reason that it is restricted to
extensive sites. Larval development, however, may be the critical phase since this takes
place in June when the medium intensive meadows are cut (Kullenberg 1944; Wagner
It is known that L. dolobrata does not
nitrogen is a
Southwood 1978).
and that
occur on
factor which limits its larval
1952; Southwood & Leston 1959).
gracilicomis is
lygaeid bug that occurs
P.
extensively managed meadows (Fig. 5).
dry places (Wagner 1966), though little more is
restricted to
in
warm,
It is
a
known
19
The
biology.
about its
heterogeneous
the
dry microclimate on
thermophilic species (Otto 1996).
warm
and
and open
ground
vegetation
of extensive sites
provides
and therefore ideal conditions for
a
xero-
biguliatus is also a xero-tiiermophilic species and therefore favoured by extensive
management (Fig. 5). Moreover it lives on Mclampyrum spp. where it feeds mostly on
the roots (Southwood & Leston 1959; Wagner 1966). It is therefore restricted to the
habitats of its hostplant which is found in extensive but not in intensive meadows (Lauber & Wagner f 996).
A.
H.
m-flavum is
a
monophagous species
Southwood & Leston 1959). Like its host
it
plant,
on
is
(Figs
(Southwood
& Leston
1959),
therefore
pratensis (Wagner 1952;
extensively managed
egg and develops in early
Salvia
restricted to
5 and 6). This species, too, overwinters
meadows
summer
that Ihes
as an
early cutting
may
damage
the larval
stages.
by frequent and early cutting
(Figs 5 and 6). The females lay their eggs in the upper part of the hostplants stem and
larval development begins in spring (Kullenberg 1944; Wagner 1952; Southwood &
Leston 1959). Cutting in May or June therefore may destroy its eggs or damage the lar¬
A. seticornis lives
on
Vicia
and
species
is
reduced
val stages.
For the
species
that do not appear to
respond
to
management,
two
main groups
can
be identified:
Widespread species which occur in various types of habitats: F. pratensis, L. rugulipennis (Southwood & Leston 1959; Morris 1979), C. fuscispinus (Wagner 1966), D.
baccanun (Wagner 1966), T. caelestialum (Remane 1958; Otto 1996), A. lineolatus
(Kullenberg 1944; Southwood & Leston 1959), S. laevigatum (Wagner 1952; Remane
1958; Otto 1996), P. imifasciatus (Kullenberg 1944; Wagner 1952), E. maura (South1)
wood I960) andN. brevis (Southwood 1960; Péricart 1987).
2) Polyphagous species, which live
certain
affinity
to
managed
berg 1944; Otto 1996),
fabricii (Péricart
in
a
wide range of
grasslands
and which show
a
(Kullen¬
pulicarius (Otto f996),
C.
Otto
Morris
1979;
1944;
1996),
(Kullenberg
5. minor (Péricart 1984; Otto 1996).
meadows: Ch.
H. aplerus
P. chrvsanthemi
1983; Otto 1996) and
be
species
rich habitats and valuable for specialised insect species. In contrast, the bug community
of medium intensive meadows is dominated by more widespread and less specialist spe¬
cies. Frequent and early cutting reduces the abundance of many species, and especially
certain specialised or rare species like the xero-thei-mophilic clement of the Swiss bug
fauna. The more widespread species, however, do not show a consistent response to
In
conclusion, this study shows that extensively managed meadows
management and
some
may
even
benefit from medium intensive management. Our
schemes in Switzerland, is effective
are
used in
niques
our
study
practice,
we
have
and
we can
which would support
re¬
supported by agricultural
grasslands,
maintaining insect diversity in agricultural land¬
mainly investigated only management techniques which
sults indicate that the extensne management of
scapes. Tn
can
as
m
therefore not say whether there
even
more
species. Moreover,
our
are
management tech¬
study
reveals that the
extensively managed meadows can vary greatly in respect to their species composition,
suggesting that to maintain species richness on a landscape level it is crucial to preserve
a
range of sites in
various
locations of
a region.
Acknowledgements
We
to Bernhard
greatful
are
Merz, Andrea Schwab, Sibylle Studer and Karin Ull¬
manuscript, to Philippe Jeanneret for statistical assistence, to Robyn Siin/i for her help in the field and in the laboratory, to Bruno Koch
for his help in finding an excellent research area and suitable study sites, to Sibylle
rich for their critical
Studer for
reading
permission
to
of the
unpublished data and to the farmers of Hemmental, Siblinpermission to work on their sites. Further we thank two
use
gen and Merishausen for
anonymous reviewers for their useful comments
This work
earlier version of the
manuscript.
by the Swiss National Science Foundation (5001-044634/1)
Schaffhausen (Kantonales Plantings- und Naturschutzamt).
was
and the Canton
on an
funded
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23
Chapter
2. Insect and
Manuela Di Giulio,
plant diversity patterns
Sibylle Studer,
in
Luzia Widmer & Peter I. Edwards
Summary
J.
The purpose of this study is to investigate the effects of management and landscape
structure on plant and insect diversity of agricultural grasslands. In this paper we
focus
of different meadow types and their contribution to the
of the research area. Further, we investigate whether variation in
the local
on
regional diversity
diversity
species number and abundance of individual
surrounding land use.
2.
species
between sites is related to the
(Schaffhauser Randen) is located in the North of Switzerland and
belongs to the Swiss Jura. Four study areas with varying landscape structure were
selected and meadows of two grassland management types were investigated in each
lire research
area
area.
bugs (Heteroptera) and the butterflies (Lepidoptcra: Rhopaloccra,
Hesperiidae, Zygaenidae) were chosen as indicator groups for local insect diversity
because they differ ecologically m various ways and can be expected 1o respond to
their environment differently.
3.
The
4.
Extensive meadows
true
are more
species
rich than medium intensive sites and contain
specialised insect and plant species. They therefore contribute more
regional species diversity. In contrast to the flowering plants and the
strongly
butterfly fauna, the heteropteran communities differ greatly between the areas
investigated. It is argued that the heteropteran bugs appear to be more limited by
dispersal than the plant and butterfly species. These findings are supported by a field
experiment which investigated how rapidly new habitat patches arc colonised and
and
more rare
to the
by
5.
which
bug species.
indicate that
heteropteran bug communities of meadows in different areas
explained by the surrounding land use. Multiple regression analyses
several bug species do respond to the structure of the landscape
surrounding
site.
The differences
can
partly
be
a
m
the
Key words: grassland management, Heteroptera. landscape structure, Lepidoptcra, local
diversity, regional diversity, Vegetation
INTRODUCTION
From
structures
aeroplane, agricultural landscapes appear as a
and colours. The heterogeneity of the substrate,
an
mosaic of various forms,
and the various types of
development of this
mosaic structure. A landscape is composed of local ecosystems, which are relatively
homogenous and more or less clearly limited (Forman 1986. 1995). Our study is
concerned with both the ecosystem and the landscape levels. If investigates the effects
of management and landscape structure on plant and insect diversity of agricultural
grasslands. In this paper we describe the local diversity of different meadow types and
their contribution to the regional diversity, or as suggested by Whittaker (1972), we
investigate the alpha diversity of differently managed meadows and discuss their
contribution to the gamma diversity of the research area. Our study was carried out in an
natural disturbance and human activities all contribute to the
24
extensive forested
region
in
Randen,
which has several
enclaves
arc
situated
on a
the north
more
or
of Switzerland, known
of 650 to 850
plateau
m
as
the Schaffhauser
land. These
agricultural
a.s.l. and have different mixtures of
varying proportions of shrubs and woods. Four such
enclaves were selected to investigate the influence of landscape structure on plant and
insect diversity. In each enclave (area), differently managed meadows were chosen to
assess the influence of management on species diversity.
arable and
grassland
habitats with
bugs (Heteroptera) and the butterflies (Lepidoptcra: Rhopalocera,
Hcsperiidae, Zygaenidae) as indicator groups for local insect diversity because they
differ ecologically in various ways and therefore perceive and respond to their
environment differently. A first difference is that the bugs are an ecologically very
diverse group, including phytophagous, saprophagous and predatory species (Dolling
1991), while butterflies are strictly phytophagous and therefore closely bound to the
vegetation. Secondly, the larval stages and adult individuals of bugs live in the same
habitat, while the different stages of butterflies can live in contrasting habitats. Thirdly,
butterflies are a highly mobile group and most species are strong fliers, while bugs have
a wide range of dispersal abilities, with flightless species but also very mobile ones.
Finally, from a practical point of view, both groups arc manageable in terms of numbers
of species.
We chose the true
agricultural landscapes it is important to
know how they move and how they perceive their environment (Macdonald & Smith
1990; Samways 1994). Rather little information is available for bugs and we therefore
designed an experiment to assess dispersal ability of bugs. The main goal of this
cxpcritnent was to investigate how rapidly new habilat patches are colonised and by
To understand how
which
species
survive in
species.
species diversity has been influenced by the history and the
landscape. We therefore describe briefly how the Schaffhauser
development
Randen developed and what were the most important factors determining the spatial
structure of this landscape. During the Middle Ages the area was cleared of forest to
gain charcoal for the smelting of iron ore. Only later were the clear-cut areas used
agriculturally. The fields on the plateau have been used for several hundred years to
cultivate cereals and hay for livestock. For most of this period the agriculture was very
extensive and was organised in a special system called 'Dreizelgenwirtschaft'. The land
belonging to a community was divided into three sectors which were cultivated
differently. A communal plan defined which crop was to be cultivated annually in each
sector respectively on which piece of land, and the farmers had to keep strictly to this
plan. There was a system of crop rotation on every field which included a fallow period
when the land was used to produce hay for the livestock or as grazing (Bronhofer 1956;
Russenberger 1984).
The present pattern of
of the
The forest
was
practised today
and
reached the
never
regularly
represented
as
occurs
land
mtenshely
a
a
less
height they
totally
sharply
different habitat
common
The trees
do
kind of wood pasture
in Switzerland
was
used but the management had little in
in the forests of Central
were regularly
Europe.
nowadays. Moreover, the forest
so
that
it
remained open.
cut
was
Thus,
type from the dense production forest which
defined and the transitional
areas
provided
habitats
(Schiess-Buhlcr & Schiess 1997). In the mid 191'1 century,
political
and economic
the
grazed
these forests
today. Moreover, the transition between forest and
upheavals,
with that
for wood
population
very
as
a
agricultural
species rich
result of both
of the Randen communities decreased
25
and most of the arable land
the
plateau was abandoned. The higher areas of the
remaining agricultural area was converted to
grassland which was managed extensively. Thus most of the Randen was covered with
either forest or extensively managed meadows for a period of about fOO to 150 years
until the mechanisation and intensification of the agricultural production commenced in
the 196()'s. Subsequently a large part of the grassland was turned into arable land and
most of the remaining grasslands were intensified through increased use of fertiliser and
machinery (Schiess-Btihler & Schiess 1997; Zehnder 1998). These developments led to
a decline in the species richness of the region, which has been documented for both the
vegetation (Isler-Hiibscher 1980) and the butterfly fauna. A comparison of the butterfly
fauna recorded in the 1920's with new data from the 1990's showed that during this
period about half of the species had declined in their abundance while 20% had
disappeared altogether (Schiess-Btihler & Schiess 1997).
Randen
were
rcafforested
on
and
the
Material and Methods
Research
area
and
The research
is part of
a
sampling design
area
is located in the Schaffhauser Randen
chain of limestone hills which forms the Swiss Jura. The soils
from nutrient poor limestones. The average
is 866
a.s.l.)
mean
(northern Switzerland)
mm.
The average
monthly temperature
in
mean
yearly precipitation
annual temperature
July (23.2 °C)
and
7.8
is
a minimum
in
are
in Schaffhausen
°C, with
a
and
derived
(437
m
maximum
January (-3.9 °C) (SMA
1998).
For
study we chose a design with two levels, the landscape and the site level. At
the landscape level we selected four enclaves (areas) with varying proportions of arable
and grassland habitats (Fig. 1). Within each area we investigated two grassland
management types (officially referred to as 'medium intensive' and 'extensive'), and for
each management type we selected three replicate sites. The butterfly community was
investigated in only three areas (without Hinterranden). The management types are
defined by two main factors: the number of cuts per year, and the type and amount of
fertiliser used. Medium intensive meadows are regularly fertilised with slurry and cut 23 times per year. The time of the first cut is around the end of May. The extensive
our
meadows
July.
are
not
fertilised and
arc cut once or
twice per year. The time of the first cut is
26
Fig.
6
3
0
Kilometer
agricultural used ai cas resemble enclaves in a sea of forest and have different mixtures of
arable and grassland habitats. Zelgli/Mosli has onh grasslands; more than half are extensively
managed meadows, the test compuse both medium and little intensive meadows. Hmterrandcn has
1. The
only
little arable land, the rest
The
areas
grassland
of Chlosterfcld
aie
giasslands
with
a
high propoition of extensively used
and Merishauser Randen
consist
of
a
meadows.
mixture of arable land
and
habitats.
sampled using a standardised sweep-net method
1996). The sweep-net had a diameter of 40 cm
(Remane
and samples were collected ever\ two weeks by making one hundred sweeps with the
net over a distance of about 100 m, Sampling was only carried out when the weather
conditions were good, ie a minimum of 17°C and sunshine. Sampling was also
restricted to the period between 10.00 a.m. and 17.30 p.m.; the sampling order of the
fields was varied between weeks. Every site was sampled between 7 and 9 times from
May to September. The nomenclature of the bugs follows Günther & Schuster (1990).
The butterflies were counted using a transect method. Sampling was carried out
between 10 a.m. and 4 p.m., on days when temperatures were over 17°C, wind speed
The
heteropteran bugs
were
1958; Bornholdt 1991; Otto
maximum of 3 Beaufort and there
was
below
was
sampled eight
a
times from
site and the butterflies
one
transect was
were
about
May
to
August.
recorded within
15 minutes per site
a
was
at
least 60% sunshine.
Every
site
m was walked at every
corridor. The average time spent for
A transect of 200
5
m
(Widmer 2000).
To
assess
plant species
27
quadrats were randomly located witbm each site. Percentage cover
species (top cover; summing up to 100%) was estimated in each quadrat just
before the first cut. For statistical analysis the six subsamples were pooled.
composition,
six
Im
of all
For both management types
a
hierarchical
was
level
related to the
use
types
at the level
the
number
surrounding
land
use.
were
correlated with all others. The
models.
Finally
calculated.
zones
were
remaining
developed
variables
recalculated the models with
we
proportions
of different land
of 10m, 30m, 50m and 100m
surrounding
analysed
strongly
partial regression
several steps. We first
intcrcorrelated and excluded those which were most
each site. The regression models
which variables
and the entire
For this purpose the
measured in the GIS for
were
area
was
conducted to estimate
region (Lande 1996). For
Two-way ANOVAs were
average species
conducted to assess how management and area affect the species numbers of insects and
plants. The landscape elements of the four areas investigated were mapped in summer
97 and entered into a GIS. Partial regression models were used to analyse whether
variation in species number and abundance of individual species between sites was
species diversity
each
of the site, the
analyses
m
were
only
used to define
the
significant
variables.
Dispersal experiment
intensively managed sites by planting 50
individuals of each of nine different flowering plant species in plots of 5m x 5m.
Subsequently these plots are referred to as 'flower patches'. As control plots we selected
five comparable sites (in terms of topography and management intensity) and plots
within the treated meadow (outside of the flower-patch plot). The following plant
We enhanced
plant diversity
in five
species, none of which occur in medium intensive meadows, were introduced: Lathyrus
pratensis, Sctnguisorba minor, Salvia pratensis, Onobrychis viciifolia, Thymus
pulegioides, Leontodon hispidus, Daucus carota, Cenlaurea jacea and Knautia
arvensis. The experiment was set up at the beginning of June 1998. Sampling started
one month later and was continued at three weeks intervals until the end of September.
On each occasion 20 subsamples of 0.02 m" (total area 0.4 m2) were taken with an
insect suction sampler of the type 'Vortis' (Arnold 1994). The results showed, that the
suction sampler collected mostly species which live on the ground and not those living
in (he vegetation layer. We therefore changed the sampling method in the second year
and used a standardised sweep net method with 50 sweeps per plot (Remane 1958;
Bornholdt 1991; Otto 1996). In the second year
June,
out
when the control meadows
were
when the weather conditions
was
sampling
started in
first lime.
April and finished in
Sampling was only carried
good, ie a minimum of 17°C and sunshine.
period between 10.00 a.m. and 17.30 p.m. We
were
also restricted to the
Sampling
varied the
sampling
cut for the
order of the fields between weeks.
One-way ANOVAs were conducted to test the influence of the flower patches on
bug species number. We compared the flower patch with the control plots within the
treated site and outside. We also analysed the data qualitatively by investigating
whether the bug species found exclusively in the flower patches were specialised on one
of the introduced plant species.
Results
Spatial effects
There
on
were
species numbers
fewer
extensive, though the
butterfly species in
partitioning of species
the medium
intensive sites
within and between
areas
than
was
in
the
similar for
28
the two management
per
area
was
increased
sites, the
For
by
35% from the
mean
plants
increased
in
by
types (Fig. 2). For the extensive sites, the
28 while the total number in all three
number of
area
level to the
species per
extensive sites the
was
44 per
area
number of extensive sites varied little
areas.
bug species
intensive sites the
number increased
higher
by
was
species
mean
The
particularly
a higher
number
was
r
Level
and
species
plants per
number of butterflies,
site,
area
mean
heteropteran bugs
and in total (me
+
area
41% up to 34.
62.75
was
plants
up to 74. Plant
variation. The
by
min./max.).
mean
and
in the
species
species
number of
50% up to 72. In medium
bug species
1;
(Table
Tukey post-hoc test).
mean
Medium intensive sites
2. Mean
by
number of
28.75, while the
100r
Fig.
per
by 41%
36.25 and increased
Extensive sites
100
Species
between areas, while the
54%. Moreover, the
in the Chlosterfeld
plants
and increased
number of the medium intensive sites showed
in the extensive sites
species
number
20.33 and increased
number of
33% up to 93 for the three
medium intensive sites
was
number of
43.
level. For the medium intensive
regional
area was
mean
areas
mean
total
was
number is
Species
significantly
62.
29
Table t shows that
management but
management and
Table 1.
Two-way
area
on
by
not
plant
and
butterfly species number is influenced significantly by
bug species number is affected significantly by both
area, while
area.
ANOVAs of the effect of management and
species numbers of plants, bugs and
the
butterflies. The species numbers of the insects
species
numbers
MS
F-ratio
p-value
0 0001
plant
the
log-transformed,
wcie
wcic
square-root transformed.
source
of variation
df
Plants
Management
I
6.011
30.045
Area
3
0 121
0 607
0 621
Interaction
3
0.886
4.426
0.02
15
0.200
1
0.055
7 25
0.017
Area
3
0.056
7.12
0.003
Interaction
3
0.016
2 01
0 156
Residual
15
0.008
Total
22
Residual
Total
Bugs
Management
Butterflies
1
0.J29
8.04
0 015
Area
2
0.052
3.23
0.076
Interaction
2
0.001
0 07
0 933
Residual
12
0.016
Total
17
Management
Species
response
to
the
surrounding landscape
structure
surrounding a site has no effect on the number of
bugs, butterflies or plant species (multiple regression analyses). Furthermore, the
number of plant and butterfly species do not differ significantly between areas and we
therefore did not carry out any further analyses (Table J). The bug species numbers,
however, differ strongly between the four areas and we therefore investigated how the
present landscape structure affects the bug fauna. We computed multiple regressions to
analyse how single species respond to the surrounding landscape considering zones of
The land
varying
use
in
a zone
up to 10m
widths around the
site.
In total
we
analysed
13
species
with
more
than 25
individuals.
respond to the surrounding landscape structure
(Adelphocoris seticornis, Carpocons fuscispinus, Leptopterna dolobrata, Lygus
pratensis, Nabis pseudoferus and Plagiognathus chrvsanthemi). Six species, however,
are significantly affected by the structure of the surrounding landscape (Table 2). The
abundance of Adelphocoris lineolatus, Notostira erratica, Lygus rugulipennis and
Trigonotylus caelestialium increases with the proportion of arable land. For N. erratica
Seven
species
this is the
addition,
case
T.
seem
only
to
independent of the distance. In
proportion of roads in the
positively
abundance of Polymerus unifascialus increases with the
up to 50
m,
but all others
caelestialium correlates
surrounding up to 30 m.
proportion of grassland
erratica
not
The
arc
with the
surrounding (Table 2). Chlamydatus pulicarius, N.
and Stenodema laevigatum are affected differently by site area (Tables 2 and
in
the
30
3), C puhtanus
erratica aie moie
Table 2
is
abundant
most
stiongly
Regiession analysis shows the
distance
10
30
m
B
R2
Adelphotons lineolatus
ai
able land
R2
able land
ai
Tiiqonotdns
cackstiahum
R"
loads
R2
R7
0 0005
-
0 68
-.0 0001
=
0 57
<0 000l
4 68
R2
0 39
-
p value
4 71
R2
0 42
=
4 13
R2
0 4
^.0 0001
m
B
R2
=-0 6
4 47
R2
0 "U
0 002
-
p value
=
0 46
2 "s
0 004
2 89
0 004
3 62
0 0003
0 009
5^6
0 0^
4 7
Ol
118
0 71
R'
=
R;
0 4^
0 4t
=
R2
=
0 37
.nable land
124
0 008
101
0 03 3
0 95
0 05
0 78
0 13
site ai ea
0 74
0 001
0 6S
0 004
0 64
0 007
0 63
0 012
Pohmeiusunifasctatus
R2
gtassland
fable 3
-
The p
0 04
0 5
=
«.0 0001
3 7
0 36
=
4 69
Notostaaeiiatiea
R°
0 03
stmctuie
î 00
m
B
R"
0 54
4 2
=-0 17
196
=
landscape
species lo
50
ni
p value
R1
0 001
and N
laevigatum
conelation
B
0 37
=
2 8^
able land
ai
p value
3 63
Lygus lugidipen/iis
thepaitial
while S
ones.
single bug
icsponses of
value îcpiesent the coetficients of
Specie s I Butter
large meadows,
m
associated with small
J
wo
bug
=
species
Species
aie
0 05
onl\ aliected
site
Stenodema
R
laeMgatum
R
the site
0 01
=
0 25
2 55
0 01
R2
-=
0 26
3 33
0 008
aiea
0 03
=0 18
0 95
siteaica
by
R2
0 24
=0 17
105
ai ea
=
2 19
P value
B
Chlamvdatus pulicaiiiis
R°
0 13
153
0 03
The first
v\e set up the di speis al expeiiment was
veiy dry and hot m
and
Randen,
many of the mtioduced plants died in the first month of
Few plants achieved any new giowth oi floweied However, despite
summet
the Schaffhausei
the expenment
these difficulties, tout
aftei
plant
species not otheiwise piesenl
in
the
did become established and two of them floweied, at least
m
the
weie
thetefoie able to achieve
sunoundmg vegetation
peiiod of the study We
local
mciease in plant
species richness In the llowei
patches a total of 27 bug species and 114 individuals weie recoided There
significant difleience (one-way ANOVAs) between the mean numbeis of bug
a
and contiol
is
no
species
in
the flowei
patches (6
and outside the tieated sites
(4 2
± 2
65) and
± 2
the iccoided species is specialised on
fauna iccoided is typical foi medium
piep)
17)
m
The
the control
patches
withm
(2 6
± 2
07)
qualitative analysis showed that none of
the mtioduced plants species In general, the bug
intensively managed meadows (M Di Giuho, m
31
DISCUSSION
Spatial effects
highly fragmented habitats
and represent a only small fraction of the former grassland. Until the mid 20th century
the plateau was covered with extensive, continuous grassland and various studies
suggest that the species diversity was considerably higher than at present (Zoller 1954;
Isler-Hübscher 1980; Schiess-Bühlcr & Schiess 1997). The extensively managed
meadows are about 100 to 150 years old and arc remnants of formerly much larger areas
The meadows of the Schaffhauser Randen
of continuous
created
intensification
by
has been
grasslands (Bronhofer 1956).
a
longer period
over
richness
are
In contrast the medium intensive sites
the last few decades and
are
of
to reach
were
varying age. Since there
an equilibrium with the
grasslands
suiprising that, in contrast to the medium
grasslands only show little variation in plant species
for the extensive
habitat conditions and management, it is
intensive
plateau
not
grasslands, the extensive
(Fig. 2) and plant species composition. Moreover,
medium intensive meadows is
more
fertiliser used than the extensive
ones,
the management of the
variable in respect to the amount and type of
which arc not fertilised at all. Insects, however,
different pattern of variation. The extensive sites investigated vary greatly in
both their bug species number (Fig. 2 ) and the composition of the bug fauna (M. Di
show
a
Giulio. in
prep.), indicating
The
species
probably
sites
number of butterflies varies much less between sites within
because adult insects
are not
much of
Given the
were
bug fauna is determined by various environmental
example aspect, isolation and previous management.
that the
factors besides management, for
a
history
at one time
are
an area
mobile and unsuitable habitats between the
(Fig. 2),
grassland
barrier for them.
of land
use
change
arable land, most of the
there must have recolonised these
areas
and the fact that most, if not all, of these sites
plant
at one
species which presently occur
present species composition of
and animal
stage. The
particular the differences between sites, will reflect at least in part the
differing dispersal abilities of different taxa. For plants, no regional differences in
species composition and species richness were detected between the four areas. Rare
for regional patterns,
species with a scattered distribution, which often are responsible
I9lh
are missing. After the period of arable land use until the mid
century, when these
grasslands developed, there were probably only few grasslands that could have served
sites, and in
vegetation cover can establish within a relatively short
time (Zoller 1954). slowly dispersing species probably were clearly at a disadvantage.
Therefore the plant species which occur arc mainly those which arc common and
apparently readily dispersed. This hypothesis is supported by a separate study
conducted in the same study area (Studer 2000) where exactly the same set of species
was found in 50 years old grasslands that were ploughed during the Second World War
and in older grasslands that were not ploughed for about 150 years. However, species
as
seed
sources.
Since
a
dense
easily are now rare in the Randen and are confined to
sites which have not been ploughed. Additionally the intensification of the land use in
the region led to a fragmentation of the grasslands, especially of the extensively used
ones and, as a consequence, to a reduction m rare species. The butterflies are obviously
that do not
disperse
or
establish
strong fliers and therefore the distances between suitable habitats within and between
might not be a barrier for them. This is supported by the our findings that we
only small differences in species number (Table 1) and species composition
(Widmer 2000) between the areas suggesting that most species which occur are not
limited by dispersal. The bugs show the strongest differences between areas, both in
areas
found
32
species number (Table 1) and species composition (M. Di Giulio, in prep.). Of the
regional diversity only 50% of the species occur in individual areas (Fig. 2), indicating
that many bug species occur only locally and have very restricted ranges. Many bug
species seem not to be very mobile and therefore habitat fragmentation represents a
problem for their populations.
species than the other areas, though
general,
history has been similar in all
four areas. In fact, Chlosterfeld, due to its vicinity to the village of Hemmental, has been
used mainly as arable land and may have been managed more intensively in the past
few decades (Bronhofer 1956; Keel 1993). We think it likely that climatic conditions
are chiefly responsible for the greater species richness. Most bug species that occur
exclusively in the Chlosterfeld are associated with very warm and dry habitats and are
typical for chalk grasslands (R. Heckmann, pers. comm.). The Chlosterfeld is located
about 100 m lower, is more open and less shady than the other areas and the snow melts
sooner (pers. obs.). The climatic conditions therefore differ from those of the other areas
and may favour species which prefer warm and dry habitats. Nevertheless, we also have
an indication that the spatial structure of the surrounding landscape influences the bug
community. We could show that at least some species, like the quite common species L.
rugulipennis, A. lineolatus, T. caelestialium and N. erratic prefer a mixture of arable
and grassland (Table 3). Plowever, we also found several less frequent species, like
Eurydema oleraceum. Aelia acuminata, Eurygaster maura or Aptus mirniicoides
(Wagner L966; Péricart 1987) which occur exclusively in mixed enclaves. Areas with a
mixture of arable land and grassland habitats provide more niches and habitats for
insects and are therefore more heterogenous.
The
the
fauna of the Chlostcrfeld is richer in
bug
reasons are not
obvious. In
and
the management
bug species
response
to
landscape
structure
dispersal experiment were negative. Although we achieved a
modest increase of the local plant species diversity, there was no apparent effect on bug
species directly, and no 'new' species appeared in the community. In view of the rather
poor growth of the introduced plants, it would be a mistake to read too much into these
findings. However, they do support the general conclusion that many grassland bugs are
relatively immobile and their distribution is limited by habitat fragmentation. There is
an interesting contrast in this respect to species found on flower strips planted in arable
The results of the
land, which appear
to
be
more
mobile. Ullrich & Edwards
occurring in the flower
bug species
plants within
a
barrier to these
strips
few weeks. Distances of
can
over
colonise
100
m
do
(1999)
newly
found that
created
not seem to
some
patches
present
a
of the
of host
significant
species.
The management of the
adjacent
fields appears to have
significant effect on the
species diversity of plants and insects. However, several bug species do respond to the
structure of the landscape surrounding a site. A. lineolatus, L. rugulipennis, T.
caelestialium and N. erratica are most abundant m grasslands that arc surrounded by
arable land (Table 2). They are widespread species which feed also on crops and occur
frequently in arable land (Kullenberg 1944; Wagner 1952, 1970-1971, 1975;
Afscharpour 1960; Vans 1972; Hattwig 1997; Fasterbrook 1997). For these species
arable land might represent an additional habitat or one they can switch to under
unfavourable
erratica
are
conditions
and
therefore
very mobile and have
a
enhance
their
no
local
abundance.
AU
but N.
high flight activity (Southwood I960; Morris 1969,
only over short
1979; Gibson 1980). Previous studies have shown thai /Y. erratica flies
distances and that the females
are
very weak fliers because of their reduced
flight
33
(Kullenberg 1944; Dolling 1991; Bockwinkel 1990). The restricted dispersal
ability may explain why this species is influenced by crop fields only up to 50 m. T.
caelestialium is positively influenced by the roads to a distance of up to 30 m. In the
research area, the roads arc mostly un metalled tracks with wide grassy margins. We
therefore suggest that the abundance of T. caelestialium is positively correlated with the
road margins rather than the roads themselves. This miricl bug is specialised on various
with a high amount of
grasses (Southwood & Leston 1959). Undisturbed road margins
muscles
grasses may
represent
an
additional habitat and therefore enhance their local abundance.
field margins in the
Only
(Southwood 1960).
vicinity
seem to
be
important
even
though
it is
a
strong flier
Only one species, P. unifasciatus. is significantly influenced by the proportion of
grassland in the surroundings (Table 2). This ohgophagous species feeds on Galium sp
(Kullenberg 1944; Wagner 1952). Galium spp. occur in a wide range of meadow types
and are very common in the research area. The proportion of grassland probably
correlates strongly with the abundance of this plant species, ie the bugs host plant. P.
unifasciatus is therefore positively affected b) the abundance of its host plant.
Interestingly no other species were influenced by the proportion of grassland in the
surrounding. Many common species which are frequently found in meadows, like A.
seticornis, L. dolobrata, L. pratensis, N. pseudoferus and P. chrysanthemi (Kullenberg
1944; Southwood and Leston 1959; Wagner 1970-1971. 1975)
seem not to
be affected
grassland in the surroundings. The bug fauna responds to the
structure, the species composition and the management of meadows and therefore from
a bug's perspective not all meadows are suitable habitats (Remane 1958; Curry 1994;
Otto 1996). Our scale might therefore be too coarse to detect differences from a bugs
view and we would have to work on a much finer scale to analyse specific spatial
by
the presence of
effects
on
Site
bugs (Wiens 1989;
area
affects three
increases with the habitat
species (Tables 2 and 3), The abundance of C. pulicarius
size, a relationship that has already been found in previous
1976). The opposite is the case for S. laevigatum and N.
(Simbcrloff 1974.
erratica, both species that are
studies
Kotliar & Wiens 1990).
most
abundant in small meadows.
quantitative approach to investigate how landscape structure affects the
plants and insects and how individual species respond to the spatial
structure of their surrounding. One problem with this approach is that only very
abundant species can be investigated. Species which occur only locally or less
frequently cannot be analysed properly for want of data, although these species would
be particularly interesting from a conservation point of view. Moreover, the scale turned
out to be too coarse and therefore only very rough pattern could be detected.
Nevertheless, the study has yielded valuable insights into what are the most important
factors affecting the regional plant and insect diversity in the Schaffhauser Randen.
We used
a
communities of
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40
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Beiträge
zur
36
3.
Chapter
The
Schaffhauser Randen (SH) and Rottal
of
agricultural grasslands in the
(LU), Switzerland, with updated checklists
(Heteroptera)
fauna
bug
of
of the Cantons Luzern and Schaffhausen
Heteroptera
Manuela Di Giuho, Ralf Heckmann & Andrea Schwab
Submitted
the
to
der Schweizerischen
Mitteilungen
Entomologischen Gesellschaft
Abstract
45
investigated
We
Schaffhauser Randen
in
Rottal
(Canton
findings
give
in
Wc
The
different
recorded
140
management
types
the
in
biogeography
on
bugs
in
the Canton
the literature and
our
own
for the Cantons of Schaffhausen and Luzern: From
specics-hsts
of
of
species
the Canton Luzern. Based
the Canton Schaffhausen 207
Luzern
three
of
(Canton Schaffhausen) and 31 extensively managed meadows
Luzern).
Schaffhausen and 78
we
meadows
bug
some
species
special
Three species from the Schaffhauser Randen
harwoodi CHINA, 1936,
Physalocheila
and Strongylocoris steganoides (J.
Strongylocoris leucocephalus should
Keywords: grassland, Heteroptera.
known and 201
are
species ol
are
interest is
new
from the Canton
discussed in detail.
records for Switzerland:
Polymerus microphlhalmus (WAGNER, 1951)
1875). The previous records of
SAHl BERG,
be revised.
Canton Luzern, Canton
Schaffhausen, Switzerland
Introduction
bugs (Heteroptera) arc an ecologically very diverse group, including
phytophagous, saprophagous and predatory species (Dolling, 1991). Furthermore,
some species are generahsts while others are specialists. The larval stages and adult
individuals live m the same habitat and respond sensitively to environmental changes
(Morris, 1969, 1979; Achtziger, 1995; Otto, 1996). Previous studies have shown
that the richness of the bug fauna correlates strongly with total mscct diversity (Duelli
& Obrist, 1998). These properties and the fact that despite their diversity, they are a
manageable group in terms of numbers of species, make the heteropteran bugs an
attractive insect group for ecological studies m agricultural systems. Unfortunately, little
is known about the distribution and biology of these insects. In Switzerland, the number
of publications concerning Heteroptera is Jess than half the corresponding number for
Baden-Württemberg m Germany (Otto, 1994; Reeger, 1996). According to the
literature, 758 species of Heteropteia are known m Switzerland (Otto, pers. comm.,
1997) and 724 m Baden-Württemberg, Germany (REEGER, 1996).
True
The
aim
of this paper
Switzerland, with
focus
is
to
contribute to the
the
knowledge
of the true
of Schaffhausen and Rottal
bugs
of
The
(Luzern).
regions
presented here come from various studies at the Federal Research
Station for Agriculture and Agroecology Zürich-Reckenholz about the influence of
management techniques on biodiversity in rural landscapes (financial support by the
Swiss National Science Foundation is acknowledged). The ecological results will be
presented m separate papers (DiGtutto, m prep.; Schwab, m prep.).
a
on
faumstic records
Material
and
Mimons
One research
Tab. 1 and
Fig. 1)
area
is
and
is
located
m
the Schaffhauser Randen
pint of the Swiss Jura. The soils
arc
(Canton Schaffhausen,
nutrient-poor limestone.
37
Matertal and Methods
One research
Fig. 1) and
Tab. 1 and
is
rainfall occurring
is
m
7.8
summer
°C, with
Schaffliauscn
m
(Canton Schaiihausen,
the Schaffhauser Randen
in
part of the Swiss Jura. The soils
yeaily precipitation
The average
temperature
is located
area
and the lowest
maximum in
m
(437
m
are
nutrient-poor limestone.
a.s.l.)
866 mm, the
is
late winter. The average
July (23
2 °C) and minimum
in
mean
highest
annual
January (-3.9
we imesltgated
grassland
in four
and
intensive
extensive)
intensive,
moderately
(medium
management types
locations
four
all
In
and
I
different locations of the Schaffhauser Randen (Tab.
Fig. 1).
°C) (SMA, 1998).
45 meadows of three
From 1997 to 1999,
grassland types were investigated. The medium intensive meadows are
regulaily fertilised with slurry and cut 2-3 times per year. The vegetation is dominated
by Arrhenatherum elatuis and Triselum flavescens. The moderately intensive meadows
are fertilised occasionally with manure and cut twice per year. No single plant species is
dominant but Trifolium pratensis and Tnsetum flavescens reach the highest mean cover
all
three
values. The extensive meadows
community
is
dominated
sampled during
at
least
by
not fertilised and
are
Bromus credits
one summer
are
cut once or twice
per year. The
(Stlder, unpublished data). All
by sweep-netting,
but
frequency
sites were
and number of
sampled with a suction sampler of
sweeps per
the type Vorüs' (ARNOin, 1994) and 18 with Barber pitfall traps (CURTIS, 1980).
site
varied. Besides, 10 meadows
weie
,
*
Fig
The leseaich
1
located
aieas aie
m
i
the Noith of Sw itzeiland
(Canton Schaffhausen) and the Cenüal
patt of Switzei land (Canton Luzein)
Tab
I
The five investigated
Klosteifeld
aieas
situated
the Canton Schaffhausen and Canton Luzein.
Hemmental
(SH)
Menshausen
m
(Sil)
Menshausen
780-830
683T)00/291'500-
8°36'E/46°44-N
6S'7'500/29F000
Rottal
The other
1 and
750-830
heights (LU)
Fig. 1).
study
The
RuswilBultisholz
sites are located m the eastern
area is
composed
of
hilly
650 000/220'000
hills of Rotlal
Molasse and
a
few
8° 08'F / 47° 07'N
(Canton Luzern,
moraines.
Tab.
The soils tend
38
monthly temperature in July (17.4 °C) and the minimum in January (0.7 °C) (SMA, 1998). The whole area is used very intensively for agriculture. During
1999. 31 extensively managed meadows were investigated in an area of approximately
8 km2 in the communities of Ruswil and Buttisholz. The vegetation is very
maximum
mean
heterogeneous, in
sites beinç dominated b\
some
crass
and in others by forbs. Overall,
plants, all
of intensively used meadow s w Inch are heavily
early
Also common arc Lolium multiflorum and Dactylis glomerata, both of
them characteristic species of intensive management and wet soils. All sites were
sampled monthly from May to August with a standardised sweep net method and with
Barber pitfall traps during 10 weeks.
repens, Poa trivialis and Ranunculus repens
Trifolium
of them typical
and frequently.
Tf not
the
most
abundant
fertilised and cut
specified
in the text, the
and Péricart
1975)
arc
of Wagner
keys
(1972, 1990)
(1952, 1966, 1967, 1971, 1973,
used for the determination of
were
species.
R.
HECKMANN discussed the faunistic records.
Results
In
the
total, 167 species have been recorded in both regions, which amounts to 22 % of
(Otto, pers. comm.) known species of Switzerland. Most species were
758
caught only in pitfall traps or
of endangered species for
samplers (Tab.
Switzerland, those of Baden-Württemberg (RlEGTR, 1986), Bayern (ACHTZIGER et al.,
1992) and Germany (Günther et al., 1998) have been taken as a basis for assessing the
conservation values of the investigated sites. In the Schaffhauser Randen, we recorded
140 species (12'080 specimens). 29 species (21 %) arc listed in the Red Lists for
Germany. In Rottal, 78 bug species (1' 129 specimens) were recorded. Ten species
account for more than 80 r/c of all specimens recorded and only 10 species (13 %) arc
listed in at least one of the Red Lists for Germany (Tab. 2).
collected with
a
sweep net
Altogether
in Tab.
and
and
As
only
there
11 species
is
no
were
Red
3 I species have been found that do not
2). Fourteen of
them
species probably
the forest edge. That
these
(156)
2).
suction
drifted
were
by
found
in
List
belong
species characteristic of the herbal
by
a
habitats
(see
"*"
Schaffhausen and 22 in Luzern. Most of
wind from trees and shrubs in the
also live in moist meadows is confirmed
than in the Schaffhauser meadows
to grassy
much
(Judex, 1999).
higher
layer
rate
nearby
woodlands
of the forest
edge
can
of vagrants in the Luzern
39
Tab. 2.
(Canton
Luzern). Columns: M:
Merishausen. IT: Hinterranden. K: Klosterfeld, Z: Zelgli. SIT and LU.: Total
number of individuals recorded in the two Cantons. Met.: Sampling method
used; n: Sweep net, s: Suction sampler, p: Pit fall traps. R.L.: f: BadenWürttemberg (D), 2: Bayern (D). 3: Germany. "*": Species specialised on
plants not normally occurring in meadows (trees, shrubs, ferns) and which
probably have been drifted by the wind. The nomenclature of the species list
from Ceralocombidae to Miridae (excluding the arrangement in the subfamily
Mirinae) follows the Catalogue of the Heteroptera of the Palaearctic Region
(Aukcma & Ricger, 1995, 1996. 1999). The subfamily Mirinae (changed
names adapted from the above catalogue) of Miridae and the Pentatomorpha
follow the Verzeichnis der Wanzen Mitteleuropas (Günther & Schuster, 1990).
Species
list of the
Schaffhausen)
investigated
and
the
meadows in the Schaffhauser Randen
Rottal
M
Species
(Canton
heights
II
K
Z
SH
LU
Met.
R.L.
s
2
P
1
Ceratocombidae
+
6
Saldiila orthochila (Fieber.
+
1
Saldula saltatoria
+
Ceratocombus
coleoptralus (Zctlcrsledt, 1819)
Salmdae
3
Saldula c-allntm (Fieber,
1859)
1859)
(Linnaeus, 1758)
2
+
n
11
n/p
TlNGIDVE
Acalypia carinaia ('Panzer. 1806)
Aealvpla marginale! (Wolff, 1S04)
Campylosteira venia (Fallen. 1826)
Catoplatus jabricii (Stâl, 1868)
+
Kaiama iricomis (Schrank, 1801)
+
+
4
1
P
90
3
n/p
n/s/p
n/p
n/s/p
18
+
+
+
+
+
+
+
+
%Physaiochcüa harwoodi China, 1936
Tingis reticulata Hcrrich-Schaffcr, 1835
71
+
22
+
1
9
n
5
1,2
2
2.3
2,3
2
2,3
n/p
Nabidae
Himacerus
major i.A. Costa, 1842)
(O. Costa, 1834)
"Himacerus apterus (Fabricuis, 1798)
Nabis flavomarginalus (Scholtz, 1847)
+
Nabis brevis brevis Scholl/.. 1847
+
Iliinacerus ininnicoides
Nabis jenes
Nabis
(Lmnacus, 1758)
pseudoferus pseudoferus
Romano, 1949
Nabis punetatus punelatiis A. Costa, 1847
Nabis nigosii.s
1
P
20
n/p
3
n
34
4
7
+
+
3
n
+
+
+
+
5
18
+
4-
+
+
131
2
n/p
n/p
n/s/p
+
+
+
11
n/s/p
(Linnaeus, 1~5S1
+
+
2
18
n
+
10
1
1
n
+
2
40
n
1
1
1.2
Antiiocoridak
"'Acoinpocoris alpinus
Anlhocoris
neinonun
'''Temnostclluis
i
Reuter. 1875
Linnaeus. 1761)
+
+
pusilliis (Herrich-Schàffer,
P
1835)
Onus
majuseulus
(Reuter, 1879)
Orias minimis (Linnaeus. 1"dX)
+
+
+
3
9
n
6
n
2
40
M
Species
II
K
z
SIT
LU
Met.
Miridae
pleridis (Fallen, 1807)
Campyloneiira virgiila (Herrich-Schäffer. 1835)
Dicyphus anntdalus (Wolff, 1804)
Dicvphus globulifer (Fallen, 1829)
Dicyphus errans (Wolff, 1 S04)
*Dicyphus pal/icltis (Hcinch-Schäffcr. 1836)
^Dicyphus slaehydis slachxdis J. Sahlberg, 18"8
+
Deraeocoris inorio (Boheman, 1852)
-r
Deraeocoris ruber (Linnaeus, 1758)
+
^Rrvocoris
4
f
+
to
dolobrata (Linnaeus, 1758)
i
Stenodema
4
+
+
+
+
laevigata (Linnaeus. 1758)
2
t-
+
+
+
n
1
n
6
n/p
1
n
n
to
n
15
n/p
n/p
40
n
n/p
n/pi
+
7
78
+
164
8
+
222
3
Megaloceraea recticornis (Geoffroy,
Nolo.stira clongata (Geoffroy, 1785)
1785)
Noloslira erratica (Linnaeus. 1758)
caelestialium (Kirkaldy, 1902)
+
;
+
+
i
|
+
1
1
n
+
+
1065
177
n/s/p
+
+
+
157
1
n/s
1
+
+
!
+
+
Adelphocoris
seticornis (Fabricius. 1775)
+
1
+
+
Brachvcoleus
pilieomis pilicornis (Panzer.
,
1805)
n
n/p
+
i
+
Phytocoris longipcnnis Flor. 1860
Adelphocoris lineolatus (Goeze, 1778)
'
n
4152
Stenodema vi reus (Linnaeus, 1767)
Trigonotxlus
n
9
+
i
Stenodema ealcarata (Fallen, 1807)
Stenodema holsala (Fabricius, 1787)
n
1
2
Pithanus maerkelii (Herrich-Schäffer, 1838)
Leplopterna
1
473
n
4
n
+
696
n/p
+
1
n
;
Calocoris
a)finis (Herrich-Schäffer, 1835)
alpeslris (Meyer-Dùr, 1843)
1
+
2
""Calocoris
Closlerolinus bielavatus (Herrich-Schäffer, 1835)
Closlerolflinus
norvégiens
+
*Giypocoris sexgiillalus (Fabncius, 1777)
+
n
8
n
322
n/p
1
+
(Gmelin, 1790)
3
12
+
n
1
n
1
'*Rhabdomiris slrialcllus strialellus (Fabricius,
n
1794)
Hadrodemus
in-fhivuin (Goc/e,
1778)
+
Slenolus binolatus (Fabricius, 1794)
+
""Dichrooscxliis intermedins Reiner, 1885
~f
+
+
4-
239
I
n
1
n
2
n
4
1
n
4-
2
+
Apolvgus liieorum (Meyer-Dui. 1843)
Apolygus spinolae (Me_\ci-Diir. 1841)
L.ygocoris pabutinus (Linnaeus, 1761)
'-l.vgocorn viridis (Fallen, 18Ü7)
Lygus pratensis (Linnaeus. P5S!
+
+
+
+
+
677
1
n/s
Lygus rugulipennis Poppius.
+
+
4-
+
371
161
n/s/p
+
4
+
+
l^ll
Lygus wagneri Remane. 1955
Orlhops basalts (A. Costa, 1853)
Orlhops campestris (Linnaeus, P58)
+
Orlhops
+
kalmii (Linnaeus. 1758)
4
+
+
Polxmerus nigrila (Fallen. 1S29)
+
'
n
n
f
2
n
1
n
to
+
+
n
4
"''Pinalilus visicola (Puten. 1888)
IS071
n
3
6
"Pinalitus alomarius (Me)er-Dür, 1843)
Charagochilus gxllenltalii (Fallen
n
35
+
+
19
+
+
13
n
1
n
i
n/s/p
n
41
II
K
Z
SU
4-
f
4-
+
745
+
2
n
-r
+
+
+
11
n
+
+
44
1
n/s
2
3
n
Species
M
Polyments lnicwphtulinus i Wagner. 1951)
Polvmenis unifasciatus (Fabricius. 1794)
Capsodes gotlucus gothicus (Linnaeus, 1758)
Capsus aler (Linnaeus. 1758)
+
Haïtiens aplerus apterus (Linnaeus. 1758)
+
Orl/iocephalus
Orthocephalus
4-
coriaceus (Fabiicius, 1777)
+
Globiceps fulvicollis Jakovlev. 1877
^Heterocordylus tiunidieornis (Herrich-Schäffer.
Met.
2
n/s/p
1
saltator (Hahn, 1835)
Strongylocoris steganoidcs (J. Sahlberg, 1875)
^Blepharidoptcrus angiilalus (Fallen. 1 807)
Globiceps flavomaculaius (Fabricius, 1794)
LU
+
+
+
n
39
n/p
t
1
2
|
'
4-
i
n
3
|
4-
R.L.
2
n
n
+
23
n/p
+
1
n
1835)
Heterotoma
1
planicornis (Pallas, 1772)
'^Malacocoris chlonzans (Panzer, 1794)
*Cremnocephalus alpestris Wagner, 1941
Hallodapus ntfescens (Burmcislcr. 1835)
Ainbylvlus
nasutus
(Kirschbaum.
1856)
1
+
0
4-
n
n
i
2
2
n
1
8
n
1,2,3
i
i
*Alractoiomits
inagnicornis (Fallen, 1807)
+
'*Alraclotomtis
parvuhis
+
3
4-
l
Renier, 1878
Campylonuna verbasei tMe>er-Diir. 1843)
Chlamydalus pulicarius (Fallen. 1807)
Chlamydalus pulltis (Reuici, 1870)
+
4,
l
+
4"
*Harpoeera thoracica (Fallen. 1807)
Macrotyhis herrichi (Reutei. 1873)
Megalocolcus inolliculiis (Fallen, 1807)
+
^Phoenicocoris obscurellus (Fallen, 1829)
+
arbuslonun arbiistorum
+
215
n
n
1
2
n
1
11
+
11
+
2
2
+
+
;
4
+
11
n
1
n
10
n/p
1
+
n/s/p
+
+
+
Criocoris crassieornis (Hahn, 1834)
Plagiognalhiis
n
1
+
n
2
n
(Fabricius. 1794)
Plagiognalhiis chrysantheini (Wolff, 1804)
pinclella (Zetlcisledt, 1828)
perrisi (Mulsanut Rev, 1852)
piceae Reutei. 1878
'"Plesiodema
4-
+
+
+
1274
n/p
1
4-
n
3
n
2
n
Psallus haemalodes (Gmelin, 1790)
3
n
^Psallus varians varians (Herrich-Schäffer. 1841)
4
n
"Psallus
"Psallus
ARADIDAE
1
1
"'Aradiis cinnainoineus (Panzer, 1794)
4
î
+
2
P
Piesmatidae
Piesma maculaluin
(Lapone.
1832)
4-
+
2
4-
43
n
Berytidae
Berylinus clavipes (Fabricius. 1775)
+
Bervlinus minor
+
+
4-
1
n
2
+
.i
+
16
n/p
1,2
+
1
n
2
+
l
n
(Hcrrich-Sehaffcr, 1835)
Berylinus crassipes
Bervlinus
iHernch-Schhft'cr.
monlivaspts
to
+
1835)
(\le\ei-Diir. 1841)
Berylinus signoreti (Fieber. I 859")
Campsocoris pimetipes iGcrmar,lS22)
!
+
9
n/p
n/s/p
42
M
Species
II
k
Z
se
LU
Met.
i
2
n
1
n
1
n
R.L.
Lygaeidaf
Nysius
encae
(Schilling 1829)
+
>Kleidoeens it\edae (Pan/ci, 179/)
Cymus
1861
inelanoLcplialus Fiebei,
Isclinodemiis sabulcii (Fallen
Plalxplaxsahnu (Schilling
18291
Plmlhistis biei ipenms (Latiulle
Piopislelhus
o
1829)
1S07)
4
holosaiLCiis (Scholl/, 1845)
r
19
n
+
14
n/p
t
3
n
latus Douglas iV. Scolt. 1871
+
2
P
1865
+
2
n/s
-r
8
4
1
+
1
Di
vmus
Di
vmus i veil
Dou.il
is
& Scolt
Dnimts s\hattcus (Fibuuus 1775)
1
11
b
Gasliodes abtelwn Beigioth. 1914
Scoloposlethus aftinis (Schilling, 1829)
Seolopostethus thomsoni Reutei 1874
Stygnocons milieus (Fallen, 1807)
Slxgnocoiis sabulosus (Schilling 1829)
Giaplopellus lynccus (Fabnuus 1775)
+
f
+
2
1
Rlnpaiochiomus pun (Linnaeus 1758)
Peiitiechiis qiaahcoinis Pulon 1877
Meçalonotits cluia^ni ihiiagia (Fabnuus,
+
+
+
1
1
1,2 3
n/p
2
n
P
1
n
s/p
2
s
53
n/p
46
P
57
n/p
n/s/p
1,2
l
n
1
1
n
2
n
1
n
11
n
6
n
97
1
2
2
1794)
Fmbletlus \eibasn (Fibiuius
1803)
1
Pyrriiocoridve
Pyiihoeons aplerus (Ï
CoRrio
innaetis
1758)
+
VL
Svminastus ihoinbciis (Linnaeus
1767)
4
Coieus inaiguialus (Linnaeus. 1738)
Conciliais dtnliculalus (Scopoli
4
1763)
i
+
4
i
Alydidae
Alydus calcul
alio, (Linnaeus
1758)
+
1
--
_-
Riiop
vi in vi
Coiizus Inoscvami (Linnaeus, 1758)
+
+
3
n
Rhopalus panimpunclaUc Schilhn» 1829
Rhopahis subnipc (Gniclin 1790)
Sliclopleiiius abutiloii abutiloii (Rossi, 1790)
Sliclopleiiiuspiinclaloneivosus (Goczc, 1778)
1
-t-
2
n
Plaiaspidvi
3
n
1
3
n
4-
1
n
16
n
17
n/p
i
Coptosoma saiiellatum (Gioihoj 1785)
Cydnid
f
\e
Sehinis lueliiosus MulsanuV. Rt\
1866
1
'
Caullwphoius uupii s>u\ llomtli 1 b>S1
Xdomeius bcpitkwis (1 innaius '"^5)
Legnolu s pu ipi M Fallen ISO?
I
+
+
F
<-
e
4
|
r
b
2
n
1
+
90
n'p
1
n
2
3
_
Thyreocoridvi
ThvHOcons Haiabaeoidcs (Linnia,s
1758)
4
1
n
43
M
Species:
IT
K
+
+
Z
SH
LU
Met.
22
1
n
R.L.
Scutelleridae
Eurygaster maura (Linnaeus. 1758)
Eurygaster testiidinaria (Geoffroy, 1758)
2
n
Pent.vtomidae
Sciocoris
lnacrocephahis
Sciocoris
niicropluhalrnus
Fieber, 185 1
4-
Flor, 1860
Aelia acuminata (Linnaeus. 1758)
+
Neolliglossa ptisilla (Gmelin. 1789)
3
n
1.3
50
n
2
8
n
>
+
n
I
Rubicoiiia intermedium (Wolff, 1811)
n
+
?
6
n
Paloinena viridissima (Poda, 1761)
+
3
4
n
Drvocoris vernal is (Wolff, 1804)
+
1
97
8
n/s/p
3
Paloinena
prasina (Linnaeus, 1761)
+
Carpocoris fiiscispimis (Boheman, 1849)
Carpocoris piirpiireipennis (Dc Gccr, 1773)
Dolycoris
4-
+
+
+
4-
4-
+
4-
23
+
+
+
135
4-
+
baccanun (Linnaeus. 1758)
n
n
n/s/p
n/p
Eurvdema oleraceum (Linnaeus. 1758)
+
*PenlaIoma
4-
2
n
+
1
n/p
rufipes (Linnaeus.
1758)
Troilus luridiis (Fabricius, 17"75)
8
1
"''Anna cuslos (Fabneius. P94)
Zicrona caerulea (Linnaeus, 1758)
Total number of
species
167
4-
+
4-
+
IS
112
41
70
46
140
2,3
n
n/p
78
2
44
Discussion
General discussion
publications concerning the heteropteran fauna
the cntomologically more interesting southern part
Recent faunistic
mostly deal with
especially the Canton
Ticmo where till
now
of Switzerland
of the country,
395 species have been found
(PÉRICART,
Rezbanyai-Rksfr, 1992; Otto, L992, 1994, 1995,
L984;
1996; Otto & Burki, 1996; Otto & Rezbanyai-Rlsfr, 1996; Rampazzi & Detiiier,
GöLLNER-SaiHiDiNG &
1997; Rezbanyai-Reser, 1993, 1997). The other Cantons
the fauna is much less well known than that of
various
regions
investigated and
Baden-Württemberg.
well
are not
of
knowledge in the various Cantons of Switzerland
Middle-Europe (Günther & Schuster, 1990),
Switzerland (Otto, unpublished list), Baden-Württemberg (RiEGER, 1996) and the
Landkreis Konstanz m Baden-Württemberg (Hlckmann, 1989, L990, 1993, 1996,
1999; Heckmann & Rteger, m prep.).
Tab. 3 gives
m
relation
a
survey of the faunistic
data
to
One way to
available
assess
the
from
ecological
differences between
regions
of
a
country is
at differences in species number of families known to be indicators for
areas
to
indicator
look
varying
Tingidae, Reduviidac, Lygaeidae,
Pentatomidae increase from boreal and alpine
types. The numbers of
of families like the
species
Bcrytidae and the
mediterranean and tropical habitats.
families of xero-thermophily. On the
Rhopalidae,
to
climate
Coreidae
and Anthocoridae decrease
These taxa
other hand
can
therefore be used
species
as
numbers of Miridae
indicators for
direction,
more temperate
being
climates (Gollner-Scheiding, 1989a). Although the data are not complete it is
interesting to note that there is a good fit of the data from SH, KN and even TG which
are located close together (Tab. 3). Moreover, there are big differences between these
areas and the Canton Luzern which has the highest proportions of Miridae and the
lowest proportions of xero-thermophihe Tingidae. Reduviidae, Berytidae, Lygaeidae,
Coreidae and Rhopalidae.
We found
more
than
m
one
the
same
fifth of all known species of Switzerland in the meadows
oi" the Schaffhauser Randen and the Rottal. The sites in the
however,
arc
much
more
and
much
diverse than those in Luzern and
oï
region
of Schaffhausen,
found
nearly twice as
endangered species. Most of the
restricted to dry and extensively
we
higher proportion
species
are
xero-thermophilic species, which are
managed meadows (Di GiULiO. m prep.). The climatic conditions in the Schaffhauser
Randen not only favour these species directly, but in combination with the soil
conditions, have limited the intensification of agricultural production. It is known for
other taxa (eg butterflies and flowering plants) that many species could persist here
which have disappeared from the other regions of Switzerland (Isler-Hubsciier, 1980;
many
a
latter
Schtess-Buiiler & Schiess. 1997).
rare or
Tab.
1928).
KN
45
TG
EU
7,0
LU
10
Sil
4
TI
Absolute Numbers
BW
10
401
IS
271
14
284
13
164
7
156
7
CH
KN
Ti
Percentage Numbers
BW
4,8
; si i
!
LU
TG
2.4
| 12.1
j
6,8
1,1
0,6
10 7
8,7
__3.0_
1.9
11,4
5.2
j
8,7
2.4
5,4
I
6 2
7,3
2.6
6.5
2.4
1.8
6.3
2.7
1.9
1,9
3,8
Miridae
5,9
2.1
2,3
20
;
!
:
j
3. List of absolute and relative numbers of ecologically most important families caugbt in different regions.Columns:
EU: Middle-Europe (Günther & Schuster, 1990). CH: Switzerland (Otto, unpublished list & pers. comm.. 1997:
uncontrolled literature data). BW: Baden-Württemberg (Rieger. 1996: literature data) KN- Landh-eis Konstanz
(Heckmann. 1989. 1990, 1993. 1996. 1999; Heckmann & Rieger. in prep., unpublished verified records). TI: Canton
Ticino (Péricart, 1984: Rezbanyai-Reser. 1993: Olio. 1994. 1995, 1996: Otto & Bürki. 1996; Rezbanyai-Reser. 1997:
the 12 unpublished records of Dethier excluded). SIT: Canton Schaffhausen (Tab. 5). LU: Canton Luzern (Tab. 4).
Thurgau (Blöchlinger. verified records of unpublished list & pers. coram., 1999 partly based on
Other families: Hydrocorisae. Amphibiocorisae and the "rest"' of Geocorisae.
TG: Canton
Hofmânner.
EU
j
:
22
; CII
4^
74
49
i
1,7
2,7
7
4,5
8
!
8
3,9
7
4,6
!
2,3
11
5,1
14
!
0,6
16
3,4
IS
2,4
1.5
36,1
Nabidac
Tingidae
1,8
0,6
56.5
!
2.0
5.6
40.8
3,4
36,7
7
6,9
24,0
4.6
1.7
0.5
12
1,8
1.7
39.9
2.6
15,2
9
1,4
39,0
1.7
15.1
7
J,3
35,8
23
15.2
18
1.3
15.3
23
14 6
[joy
33
4
2.5
!
36
30
1.6
52
1
5
6,6
Anthocoiiddc
19
8
j
7
1
1~k
21 9
4,7
25
2
1
24,8
4,7
2
5
Î2
4,9
58
5
15,0
18.9
jj.
10
18
!
!
1.8
62
10
17.5
19,5
1,7
10
7
28
17.5
!
6
110
10
19,5
36,0
95
10
17
26
20.1
38,7
1
116
14
I
*
68,2
100
1
23
43
23.7
100
63
263
44
201
31
207
67
383
72
411
141
724
157
758
71,4
76
14
15
Rcduvndae
Beiytidae
155
i
52
70
251
t««t
37,8
Lygaeidae
27
;
!
i
Coreidae
17
\
!
Rhopalidae
Other families
|
Pentatomidae
Species
j
number
46
Canton Luzern
Faumstic data
near
ïlasle
are
available from the Canton Luzern for the
(Rezbanyai,
upland
1980; Gollner-Schtiding, 1981) and for the
moor
Baimoos
surroundings
of
Vogelwarte Sempach (Gollner-Scheiding. 1982). Later the Vogelmoos near
(Gollner-Schftding, 1990) was investigated and occasional records for
different places in the Canton of Luzern were published (Gollner-Schetding, 1989b).
The record of Panaoriis adspersus Muls. & Rey was added from the Museum of
Natural History in Pans (Péricart, 1998) and 35 species by an investigation near
Rottal (Judex, 1999). Tlnrty species of water and seniiaquatic bugs (Nepomorpha and
Gerromorpha) were found m the Wauwiler Ebene and 18 in the Mösliwciher near
Schötz (Wipraciitiger, 1999 a, b). We found 78 species for Rottal, 26 of them being
m Tab. 4). Altogether 201
new records for this Canton (see
species have been
the
Neudorf
'"""
recorded in the Canton Luzern.
publications of GöLLNER-ScnETDiNG (1981, 1982) Orlhops basalis A.C. was
published as Ortliops kahnii L. The misidentification of these two species was
apparently due to a mistake in the key of Wagnfr (1967, 1971) which later was revised
In the
(RlEGER, 1985).
47
Tab. 4. Check list of
Heteroptera
from
for the Canton Luzern. The nomenclature of the
(excluding the subfamily
Catalogue
Heteroptera of the Palaearcric Region
(Aukcma & Riegcr, 1995, 1996, 1999). The subfamily Mirinae (the names
were adapted from the above catalogue) of Miridae and the Pentatomorpha
follow the Verzeichnis der Wanzen Mitteleuropas (Günther & Schuster,
species
list
Ceratocombidae
Mirinae) follows the
1990). For complete
different families
are
Miridae
to
of the
names
sec
Tab.
2.
The number of
noted. The asterisks indicate the 26
species in
species
new
the
for
the Canton.
Check
Heteroptera for
list ok
Nepidae
Nepa
2
cinerea L.
the
201
Canton Luzern
Gerridae
6
Aquarius pallidum
F.
Gerris argenlatus Sebum.
Ranalra linearis L.
CORIXIDAE
12
Gerris
gibbifer
Schum.
Micronecla scliollzi Ficb.
Gerris lacustris L.
Callicorixa praeusla Ficb.
Gerris
odontogasler Zeit.
Con.\a punctata 111.
Gerris thoracicus Schum.
Hesperocoma linnaei Ficb.
Hesperocorixa sahlbergi Ficb.
Saldidae
Paraeorixa
Ficb.
conanna
Macrosaldida variabilis 14.-S.
Sigara nigrolineata Ficb.
Sigara semisiriata Ficb.
Sigara striata L.
Sigara
Sigara
'''Saldula c-album Fieb,
Saldula sallatoria L.
TlNGlDAF.
distincte Fieb.
Sigara falleni
''Kalama iricornis Schrank
.
Natjcoridae
1
Ilvocoris cimicoides L.
glauca
"Tingis
reticulata II.-S.
Nabidae
NOTONECTIDAE
Noloiiecta
4
"Acalypla carinata Pz.
*Acalvpla marginale WIT.
Fieb,
lateralis Lch
4
Charloscirla cincla II.-S.
1
8
"Hiniacerus
major
A. C.
Himacents minnicoides 0. C.
L.
Noloiiecta maculate F.
'"Hiniacerus apterus F.
Noloiiecta viridis Dele.
Nabis limbalus Dhlbm.
Pleidae
1
Plea minulissima Lch.
Nabis
Mesoveliidae
Mesovelia furcata Muls. &
"'Nabis brevis Sz.
Nabis
1
Rey
Hydrometridae
2
Hydromctra gracilenta Horv.
Rem.
Nabis rugosus L.
Anthocoridae
9
Acompocoris alpinus
Anlhocoris
Hydromel in slagnorum L.
Veliidak
ferns 1,.
pseudoferus
confiisus
4
Anlhocoris
neinonim
Teimiostellius
Micmvelia reticulata Burin.
Orius lalicollis Reut.
Veiia
caprai Tarn.
Reut.
Anlhocoris limbalus Ficb.
Microvelia pygmaea Dut",
Velia sciulii Tarn.
Reut.
Oriiis
L
pusillus
majusculus
GVws miiiulus L.
GV/hs victims Rib.
U.S.
Rent.
48
Lygus punctalus Zell.
Za ç»i pratensis L.
Lyells rugulipennis Popp.
1
CïMICTOAE
Cimex lectularius L.
Reduvhdae
Reduvius
1
Lygus wagneri Rem.
Orlhops basalis A. C.
"'Orthops catnpestris L.
Orthops monlanus Schill.
"Pinalitus alomarius Mey.-D.
P£_rsonahj.sL.
100
Miridae
pleridis Fall.
Monalocoris jilicis L.
"'Campyloneiira virgule 1-1.-S.
Dicyphus epilobii Rcut.
Dicyphus errans Wff.
Dicyphus hyalinipennis Burin.
*Dicxphus pallidas H.-S.
Dicyphus slachydis I. Shlbg.
Alloeotomiis germaniciis E. Wgn.
Alloeotomus golhicus Fall.
''''Bryocoris
Pinalitus cervinus U.S.
Pinalitus rubricalus Fall.
''Pinalitus visicola Put.
Agnocoris
Deraeocoris ruber L.
Deraeocoris lulescens Schill.
Pilhanus inaerkelii IL-S.
Leplopterria dolobrata
Teralocoris
pallidum
J,
"Haïtiens apte rus L.
L.
"Orlhocephalus coriaceus F.
B/epharidoptenis angulatiis Fall.
Blep/iaridoplerus diaphanus Kb.
Shlbg.
Stenodema calearata Fall.
Stenodema holsata F.
Stenodema
laevigata
Drxophilocoris /lavoqiiadriiiiaciilaliis
L,
""Globiceps flavomaciilatus
""Helerotoma planicornis Pali.
recticornis Gcolïr.
Mecomma ambulans Fall.
Noloslira el on gala Gcolïr.
Orlhotyliis marginalis Rcut.
Orlholyliis nassalus F.
Noloslira erratica L.
Trigonolvlus
caelestialium Kirk.
Phvlocoris dimidiatus Kb.
Phylocoris
intricaïus Flor
Phvlocoris
longipennis
Flor
Kb.
Pant il ins lunicatus F.
^Adelphocoris lineolatus Gz,
Adelphocoris seticornis F.
Calocoris ajjinis U.S.
Calocoris alpestris Mey.-D.
Calocoris roseomaculalus De G.
Closlerolinus biclavanis U.S.
Closlerotonnis norvégiens
Grypocoris sexguitaius F.
Gm.
Rhabdomiris striate!Iw. F,
Slenolus binolaliis F.
Apolvgus
lucoruin
I Orlhotyliis prasinus Fall.
[
Phylocoris liliac nliac F.
Phylocoris pini
De G.
F.
Stenodema virais L.
Megaloceraea
rubieundus Fall.
iripusiiilaiiis F.
Camplozyguni aeqnale Vill.
Charagochilus gyllenhalii Fall.
Polymerus holosericeus Hahn
''Polymeries unifasciatus F.
Capsus aler L.
Liocoris
Mey.-D.
tcygocoris pabulums L,
Lygocoris nigicollis Fall.
Lvgocoris contaminai us Fall.
Lygocoris viridis Fall,
coccineus Mey.-D.
Crcinnoccphalus alpestris E. Wgn.
Pilophorus clavatus L.
Pilophorus confusiis Kb.
Pilophorus perplexus Doug. & Sc.
Pseiidoloxops
"Ambvlvlus
nasillas
Kb.
Alraclolomus kolenalii Flor
magiiicornis Fall.
Compsidolon salicellum U.S.
'"Clilamydalus pulicariiis Fall.
Harpoccra thoracica Fall.
"'Megalocoleus inollicuhis Fall.
Orthonotus nififrons Fall.
Atraclotomus
Phoenicocoris obscurellus Fall.
Phyllis coryli L.
Phyllis inelanocephalus
L.
Placochiliis seladonicus Fall.
Plagiognalhiis
arbustoriiiii F.
Psallus belitleli Fall.
Psallus
ambiguus Fall.
perrisi Muls. & Rey
; "'Psallus
49
Miridae
Coreidae
CONT.
Coreus
''Psallus piccae Rcut,
Psallus
fallen/
Rcut.
Rhopalidae
_
Psallus haemalodcs Gm.
Psallus
lepidus
varians
i
Fieb
Psallus mollis Muk &
Psallus
Rc>
varians II -S
Eurygaster
Eurygaster
vAradus cinnamoineus Pz
Eysarcoris
Berytinus minor II -S.
|
19
Lygaeidae
scnatilis
auicsccns
Cyinus inelanoicphalus Fieb
1
Penlatoma
'
Picromenis bidens L.
Z(c;
Bgtth
Gaslrodes grossipe.s De G.
Scolopostelhiis iifjmis SCHILL.
Scoloposlelhus thomsoniRruT.
Stygnocons rusticiis F\ll,
Pachvbrachius jiacticollis Schill.
Panaoriis adspersus Ml'LS, & REY
Rhvparochromus pini L
penicillatus
PI
\i I
Meçalonolus chiraçra
F.
Paloinena
1,1»»«
Ischnodemus sabuleti Fall
Pentrechus nubilus F
12
\H\
fabricii Kiik.
prasina L.
Carpocoris piirpiireipennis De
Dolycons baccanun L.
Eurvdema dominuhiin Scop.
Rhaplugasler nebulosa Pd.
Hahn
Dist.
Gaslrodes abiclum
lesludinaria Geo ffr.
Carpocoris fiiscispimis Boh.
Kleidocerys icsedae Pz.
glandicolor
2
L.
1
'•Nysius ericae Schill.
Cymes
maura
I Paloinena viridissima Pd.
Scop.
Nysuis cvirioidcs Spin.
Cvmus
L.
Pextatomidae
Berytidae
Peiitrechus
Rhopalus siibrufus Gill.
S7z< lopleurus crassicornis
Scltelleridae
Aradidae
Spilostethus
L.
marginatus
,
1
1
,
rufipes
oust os
o»a
G.
L.
F.
caerulea L.
ACANTIIOSOMATIDAE
Acaiilhosoma haemonhoidale L.
Elasinosletlius minor FTorv.
Elastnostetliiis interstinetiis L.
Elasinucha
grisea
L.
4
50
Schaffhausen
Canlon
species of Miridae
species was published by
The earliest records for the Canton Schaffhausen include 23
collected
S. Sfttfr
by
(Mfyer-Dur, 1843).
(1864-1866),
Frey-Gessner
19th century. In
Dur (1843). Much
A list of 76
most of them also collected
by
S. Setler in the midst of
by MEYERlater the occurrence of two Saldid species was reported (Dethtfr &
Péricart, 1990). Together with our results, wc present here a list of 207 species of
Heteroptera for the Canton Schaffhausen, 1 18 of them being new records for the Canton
the
(see
ai-"
We
his list, he included
a
revised list of the Mirids recorded
in Tab. 5).
collection,
able to check the determinations of" Frey-Gessner because his
not
were
which
Schaffhaiisen,
probably deposited
destroyed by bombing of
was
was
History Museum of
Schaffhausen during world-war II (A.
only updated the names (AuKEMA &
the
m
Natural
MÜLLER, pers. comm.). Therefore, we have
RffiGER, 1995, 1996, 1999; HEISS & PÉRICART. 1983; PÉRICART, 1984, J998; STICHEL,
1955-1962). However,
1.
Velia
currens
(1864-1866).
that species
the
F.
following
was
four records
are
in the lists
published
problematic:
for the Canton of FREY-GESSNER
later, Velia caprai Tam. and Velia saulii Tam. were separated from
distinct species (Tamanini, 1947), It is therefore not clear which of the
Much
as
Probably it was Velia caprai TAM., which
common in the southern part of Baden-Württemberg, while Velia currens F.
in Ticino (DETHIER & MATTHEY. 1977).
three species
2.
was
actually
Aconipocoris
Aconipocoris
found.
alpinus
pygmaeus Fall,
REUT,
so
was
that
we
described
in
1875
and
is also very
occurs
only
separated from
species was
do not know which of the two
found.
Taphropeltus hamulatus Thoms. was described
as a synonym of Taphropeltus contractus H.-S. it was
know which of the two species was really collected.
3.
4.
Sehirus morio L.
(1864-1866).
a
was
published
being considered
separated again; we do not
in 1870. After
later
in the lists for the Canton of FREY-GESSNER
At that time Sehirus luctuosus MULSANT & Rey had not been described
separate species
front S. morio. We
luctuosus Muls. & Rey which is found
Middle-Europe.
assume
that the record
frequently,
probably
as
deals with S.
while S. morio L, is seldom found in
51
lab
Check list ol
5
species
follows
fiom
the
Catalogue
199s
Riegei
above
Hcteiopteia
list
1996
Wanzen
the Cmton Schaffhausen
of the
to
Mnidae
Ileteioptu
Miiidae
and
Alittdeuiopas (Gunthei
k i is i or
Hi
<k.
i lrop ii ra i or i m
ClRVlOCOMHIDVL
Ceiatoeombus
Nr pro
vi
Ranali
a
names weie adapted fiom the
Puititomoipha follow the Veizeichms det
Schustei
1990) Foi full names see Tab 2 '"?"
C
\abis
1
colcoptiatus
Zelt
ai is
bithocons
f\<
Lch
capiat copiai 1
am
P\
tienlatus Schum
qolampis
RIiMiot
De G
I
ons
bidentala Gz
iiacundus Pd
76
Miridvi
album Fieb
Lampxloneuia MiqulaU S
Du \phus iiniuilatiis Wii
'saldula oithodula Ficb
Saldula sallatona L
IING1D
5
culicifoimis
vagabundus
Pin mata aassipcs 1
u
e
S
Reut
campeslnsT
aeons
/ mpiLOiis
'
S\iDm\r
Saldula
Du
10
\1
Fall
\phus qlobiilifa
Di
i cu oloi is moi w
Di
nu
Boh
Aeahpta cannula Pz
\calypta marqinata WI1
\( ahptapaisula 1 all
Dciacocoiis lulescens Schill
[z-iamina hutiim Fill
Steno lema holsala I
"-
/1 plop le
i
ons
nut
''Campx/ostcua \eina Fill
*("aloplalus/abncii Stil
Stenodema
Diet]la celui Schi ink
\otostua
Dicnla
lupu/iU
kalama
Pin
Vi/i
Still ink
lu da han\ oodi Chin
Pioslemma aillitla I
'~Ilunac<ins miimicohit
"•Nabis
flax
Nabis bu
Nobis je
i
"-
Nvbidw
omaiuiuilus
i is
ins
dolobiala I
lae\i%ata
1
leclicoims
Gcolli
elongate Gcolh
si n a eiiatica
L
Tnjonolylus caelestialium Kuk
Plniocoits longipenms Floi
[delphocons lineolatus Gz
\di Iphoeons seticoims F
Biachycoleus pilicoims Pz
kCalocons ajfuns U
S
Caleçons loseomaculalus De G
Closleiolomus biclcnalus H
I
latus
O C
Sz
Sz
%Nabispsaidofaus
"~Nabis piuu
s
nibci I
Meçaloceiciea
S
tnioinis
satoi
'
I
nanoium
majuscules
/ mpicons
Micio\clia icliculala Buim
Ganyai
Reut
RlDLMIDU
iid vr
Gerrid
7
alpnuis
Onus minimis L
Strata fallait Ficb
f Vcha
L
7annoslclhuspusillusll
Onus
Siçaia slilata L
Vn
207
lai« pocons pyginacus Tall
'
L
VI
um
nnosus
\( ompocons
Hi spaocoiixa inoesta Ficb
Sigai a Jossen
species foi the Canton
Vmiiocoridal
"•
CORIXID
new
Sciiatthausen
\nt on
1
luit
subfamily Mumae)
Region (Aukema &
the
doubtfull histoi leal iccoids The astcnsks indicate the 118
On c
(he
(excluding
the Palaeatctic
oi
i
The nomcnclatuic of the
1999) The subtamih Mumae (the
of
catalogue)
foi
Ceiatocombidac
\
Rem
C
( loste lolomits
fuhomaeulatiis
Closte lotonius nonegicus Gm
S
De G
52
Miridae
Grypocoris
cont.
sexyuitaltis F.
Rhabdomins striatedliis F,
i
1
'
*Hcidrodemus m-llavum Gz
'
Slenotus binotatus F.
*Dichrooscvtus intermedins Reut.
Apolygus Iiicorum Mey.-D.
*Apolygus spinoleie Mey.-D.
"'Lygocoris pa bid inns L.
""Lygus pratensis L.
*Lygus rugulipennis Popp.
^Lygtis wagneri Rem.
"'Orlhops basales A. C.
"'Orlhops campesti
"Orlhops kabnii L.
Psallus variabilis Falk
3
Aradidae
"'Aradits ciimamoineus Pz.
| Aradits
I Aradits
conspiciuis U.S.
depressus F.
! Piesmatidae
Piesma
1
capilatum
2
WIT
"Piesma maciilatuiu
Lap.
Berytidae
is L.
7
Berylinus clavipes F.
Berylinus minor U.S.
"'Berylinus crassipes U.S.
j "'Berylinus monlivagus Mey.-D.
J "'Berxlinus signoreli Fieb.
1
Pinalitus rubricalus Fall.
Charagochilus gylleiihalii Fall.
Polymerus holosericeus Hahn
"'Pedvmerus nigrila Fall.
''"Polymerus unifasciatus V.
""Polymerus mierophlalmiis F. Wgn.
*Capsodes golhicus gotlucits L.
"'Capsus aler I
,
!
"'Gampsocoris punclipes
tipiilarius L.
Germ.
Neides
Lygaeidae
"'Nysius
„
"'Haïtiens aptcnis
""Plagiognalhiis arbitsionini F.
"Plagiognalhiis chrvsanlheiniWfii.
"•Ples'wdcina pinetella Zelt.
Psallus ambiguiis Fall.
25
ericcte Schill.
""Isclmodemus sahuleti Fall.
aplerus L,
Globiceps sphaegiformis Rossi
^Globiceps jlavomaculatus V.
"'Globiceps fuivieollis Jak.
Macropiaxpreysslcri Fieb.
Hclerogasler urticae F,
'"Plalyplax salviae Schill.
Plinilusiis brevipennis Latr.
Propistethus holosericeus Sz.
'Drvmus leans Doug. & Sc.
"Dryinus rycii Doug. & Sc.
*
Dry m us sylvalicus F.
Hclerocordxlus lumidicornis U.S.
"fGastrodes abielmn
'Orlhocephalus coriaccus F.
"Orlhocephaltts solictor Halm
Strongylocoris luridus Fall,
*Slrongylocoris slcganoides I. Shlbg.
Cvllccons histrionius L.
''Malacocoris clilonzans Pz.
Orlhotyliis nassatus F.
Orlhotyliis viridinervis Kb,
'-'"Hallodapus rufeseens Bui'ill.
'"Ambylyitis nasiitus Kb.
-'"Alraclolomus magnicornis Fall.
'''Alraclotomus pervitins Rent.
Campylomma vcrbasci Mey.-D.
*Chlamydaius pulicarius Fall,
"'Chlainydatus pullus Rcut.
Criocoris crassicornis Ilahn
Hoploinaclius thiinbergii
Fall.
'
Bgrlll.
'"Scoloposlelhus afjinis Schill.
''"Scolopostethus thoiusoni Rent.
IPaphropellus contractus U.S. ?
"'Stygnocoris nislicus Fall.
i Stygnocoris sabiilosus Schill.
1
Piiclnbrachius fracticollis Schill.
Aellopus atrcUus Gz.
"Graptopeltus lynccus F.
""Rhxparochromuspini L.
Xanlhochiliis qiiadralus F.
'*Peritrechiis gracdiconiis Put.
""Megalonolus chiragra F.
"'Macrolxlus herrichi Reut,
Emblelhis verbasci F
*Megalocoleiis mol Had us Fall.
Monosynainma bohananni Fall.
Oiicolylus viridiflavus Gz.
Plerolmetiis
""Phoenicocoris obseurelhi.s Fall.
staphilinijormis
Pyrriiocoridae
I "-Pyrrhocoris aplerus
Schill.
1
L.
53
Coreidae
Acantiiosomatidae
eicuteeingiikitus
Goiiocerus
Elasinuclia ferrugeita F.
rlwmbeus L.
Syromaslus
Enoplops scapha
''•'Coreus
Gz,
[
F.
inarginattis
L.
'!'Coriotneris denticulatiis
Scop^
Alydidae
1
Alxdus ceilcareitus L.
Rhopalidae
5
^Corizus hvoscvami L.
"•Rhopaliis pariirnpunctalus Schill.
"''Rhopaliis siibnifus Gm.
""Sticlopleurus abutiloii Rossi
""Slicioplcnnis piinctatonervosns Gz._
Plataspidae
_
1
seutellatum Geo ffr.
Coptosoma
Cydnidae
Sehirus luctuosus Mills. &
Rey
Trilomegas bicolor L.
'•'Canlhophorus iinpressus Ilorv.
""Adomcnis
bigullatus
*Legncjiui2icipc.s
L.
Fall.
Thyreocoridae
"'1'hvreocoris scarabaeoidcs L.
Scutelleridae
Odonloscelis
"•'Eurygaster
^Eurygaster
fuliginosa
L.
L.
maura
testiidinaria Geo ffr.
Pentatomidae
"'Sciocoris
''Sciocoris
18
macrocephaliis Fieb.
microphthalmia, Flor
Sciocoris umbrmus WIT.
*
Aelia acuminata L.
'Neolliglossa pusilla
dm.
*Rubiconia intermedium Wff.
"'Paloinena prasina L.
^Paloinena viridissiina Pd.
Dryocoris
venudis Wff.
Chlorochroa juniperina I..
""Carpocoris fuseispinus Boh.
'"Carpocorispiirpiireipennis De
"'Dolycoris
baccanun L,
'•'Eurydema oleraceum L.
"Penlaloma
rujipes
L.
*Troilus luridus F.
Anna
cuslos
F.
'"Zicrona caerulea L.
G.
Cvphostethus
trisirialus F,
54
Biogeographv of
following,
In the
species
lists of
species
rare
discuss
we
the Red list and those
on
Germany
are
our
rarely
very
found
and
common
findings
own
m
in
detail with
Switzerland. Some
widespread
in both
critical focus
a
species
on
on
the Red
Baden-Württemberg
and
Switzerland and will not be discussed here. These
orthochila FlFB.,
arc Saldula c-album FlEB., Saldula
Wff.,
marqinata
Acalvpta
Catoplatus fabricii S TAL, Kalama tricornis
alpinus Retjt., Calocoris alpestris Mey.-D., Grypocoris
hernchi REUT., Bentinus signoreli FlEB., Ischnodemus
Macrotydus
F.,
sexguttatus
sabuleti
FALL., Adomerus biguttatus L.,
Legnolus picipes FALL., Rubiconia
SCHR.,
Aconipocoris
intermedium Wff. and Zicrona caerulea L.
compared our results for Schaffliauscn (Tab. 5) with
those ol Baden-Württemberg and the dry habitats of the Hegau (Landkreis Konstanz).
For the Landkreis Konstanz 411 bug species have been recorded (Heckmann, 1989,
1990, 1993, 1996, 1999; Heckmann & Reeger, m prep.; Heckmann, unpublished
data). In total 21 species of the Schaffhausen bug fauna have not been found in the
Canton Schaffhaiisen
Landkreis Konstanz
Light
of
our
so
-
We
far.
recently
recorded species ha\c not been found in the Landkreis
Konstanz, though
venia
it is a well investigated region. All species except Campylosteira
Phvsatocheila
harwoodi CHINA and Sciocoris tnicrophlhalmus FLOR, are
FALL.,
\ero-thermophihc species and most are on the Red List of Germany: Saldula orthochila
FlEB., Campylosteira venia F Ml.. Phvsatocheila harwoodi China, Brachycoleus
pilicornis Pz., Hallodapus rufescens Burm., Sxromastus rhombeus L., Sciocoris
niacrocephalus FlFB. and Sciocoris microphthalmus FLOR.
Thirteen species
early records but were neither found by us nor
Llesperocorixa moesta FlEB., Sigara fossarum LCH., Dictyla
lupuli II.-S., Calocoris roseomaculatus De G.. Strongylocoris luridus FALL., Globiceps
sphaegifornns Rossi, Neides tipularhis L., Aellopits a trahis F., Xanthochilus quadratus
F., Sxromastus rhombeus L., Sciocoris timbrants Wff., Chlorochroa juniperina L. and
Elasmucha fcmiqata F Also Neides tipitlanus L Aellopits atratus F., Xanthochilus
quadratus F. Sxromastus rhombeus L. and Chlorochroa juniperina L. are xerothermophilic and are mostly Red List species in Germany. Strongylocoris luridus Fall,
are
mentioned
m
the
in the Landkreis Konstanz.
,
is
found in the Black Forest
Elasmucha
Forest
ferrugata
(Kifss, 1961)
eleven species in
Rhine
Valley
1.
is
and lives
& STRAUSS.
known from the
1992) and lives
nearby
special
on
Jasionc
mont ana.
Wutachschlucht in the Black
Vaccinium mxrtillus. The distribution of the other
on
Baden-Württemberg corresponds
north of Basel
The species of
(Rieger
F. also
with hot and
dry
habitats
along
the
(Heckmann, 1996).
interest
found
by
us are-
Ceratocomhus
coleoptratus (ZhTTFRSTEDi, 1819); In Switzerland this species has
only recentl) from the Canton Ttcmo (OiTO, 1994). It was found several
times in Baden-Württemberg and is common m the Landkreis Konstanz (Heckmann &
RiEGER, m prep.). It is rarely collected because of its small size and the fact that it
occurs hidden m moss. Mostly it is recorded by pitfall
traps. In Bayern it has been put
been found
on
2.
the Red list.
Acalypta
carmaia
Ceratocomhus
arc
(Panzfr,
coleoptratus
Zftt
1806)
and
can
is
be
known fot Vaud, Bern, Zurich and Wallis
often
found
m
caught easily
VRT. 1983).
the
with
(Pfric
It
same
habitats
pitfall-traps.
is on
as
Records
the Red list for
55
Baden-Württemberg and Bayern but seems to be common m the south of
Württemberg (Rieger, 1981) and the Landkreis Konstanz (Heckmann, 1990).
3.
Campy] osteva
Switzerland H
is
venta
(Fallen, 1826)
lives hidden between
known from Bern and Graubunden
mosses
Baden-
and lichens. In
(PÉRICART, 1983) but
few record
Baden-Württemberg (HÜEBER, 1891; Meess, 1907; FISCHER, 1961;
is on the Red list of Bayern and Germany and should be also for
Reeger, 1981).
aie
for
known
It
Switzerland.
4. ThysatocheUa harwoodi China.
1936
(Fig. 2);
New record for Switzerland! The
known from
species lives on Acer pseudoplatanus and is recorded very seldom. It is
seveial places (Heiss, 1978) m fyrol (Austria), from Baden-Württemberg (RTEGER.
1981; Voigt, 1983) and from Bayern (Singer, 1952). This
with the known distribution
Fig
2
Physatocheila hatwoodi The
wider
5.
m
anow
shows thenanow black
the othci species of the genus. The
Deraeocoris morio
body length
in
record
corresponds
is
aica
of the
about 3 5
clytna which
(Photo: M.
mm.
is much
Di
Giulio)
xero-therinophilic species lives on
the Ticino (Gollner-Schfiding & RezbanyaiBaden-Württemberg and in the Landkreis Konstanz
(BotiEMAN. 1852):
Thymus and is known only from
RESER, 1992) and several places m
(ITECKMANN& RlFGER,
new
area.
This
pi'Cp.).
Brachycoleus pilicornis pilicornis (PANZER, 1805): This thermophilic species lives on
Euphorbia verrucosa. It is knov\n from the Cantons of Basel, Zürich. Aargau (FreyGi ssner,
1864-1866) and Thurgau (Blociilinger, pers. comm.). In BadenWürttemberg it is known from se\eral places and also from warm places in the nearby
6.
Wutachschlucht
m
the Black Forest
(Kt
fss.
1961).
Polymerus
nticrophtlialmus (Wagnfr. 1951): New record for Switzerland! This
species lives on Galium, often together with P. unifasciatus F.. and is common in the
Llegau and sparsely found m other parts of Baden-Württemberg. Probably there is no
Swiss record because it has been confused with P. unifasciatus F.
7.
Strongylocoris steganoidcs (J. Sahlberg, 1875) was synonymised with S.
leucocephalus by Reuter (1888). Kritshenko (1951) revised the genus and separated
these two species. Wagner (1973) treats it as a subspecies, and it might therefore have
been misidentified (RiEGFR, 1997). In Baden-Württemberg S. steganoidcs is widespread
and very common whereas S. leucocephalus is a rare species. In the Landkreis Konstanz
8.
56
S.
only
the
steganoidcs
has been found
(HECKMANN
same
prep.).We assume that
leucocephalus should be checked.
& RffiGER, in
is true for Switzerland and the records of S.
Hallodapus rufescens (BURMEISTER, 1835) is known in Switzerland only from the
Vaud, Bern (FREY-GESSNER, 1864-1866), Wallis (CERUTTI, 1937a) and
Graubünden (Valbella; Dl GlULlO, unpublished data). In Baden-Württemberg it is only
9.
Cantons
(Rieger, 1976) and from the Kniebis in the northern
known from the Suebian Alb
comm.).
Black Forest (RiFGER, pers.
Berylinus crassipes (Herrich-Schäffer, 1835) lives on Cerastium. Records arc
known from Aargau and Vaud in Switzerland (PÉRICART, 1984), the Wollmatinger Ried
in the Landkreis Konstanz (LIeckmann, 1990) and a few other places m BadenWürttemberg (Kless, L961: Rieger, 1976; Heckmann, 1996).
10.
1841): This xero-thermophilic species is
Berylinus montivagus (Meyer-Dur,
11.
known
from
only
Switzerland
m
Graubünden
(Frey-Gessner,
(Péricart, 1984) and Neuchâtel (Barbalaf, 1991) but is probably
is known from the Hohentwiel in the Landkreis Konstanz and from
Baden-Württemberg (Sciimtd,
12.
The
1967; Heckmann & Rifger, in
prep.).
latus DOUGLAS & SCOTT, 1871: This is the second record for Switzerland.
Drymus
species
only
has
(Péricart, 1998).
been recorded in the Canton Basel
and from few other
places
in
There
are
Heckmann & Rteger, in
(Heckmann, 1990;
Baden-Württemberg (Rieger
few records in the Landkreis Konstanz
prep.)
Aargau
widespread. It
other places in
1871)
more
&
Strauss, 1992).
gracilicornis PUTON, 1877: This xerophilic species is known in
Switzerland from Wallis and Ticino (Péricart, 1998). In Baden-Württemberg it is
mostly found along the valleys of the Rhine and the Neckar and also in the Llegau
13.
Peritrechus
(Heckmann, 1996).
14.
Canthophorus impressus HORVATH,
species
for
Switzerland.
It
has
(Gollner-Schtiding, 1989b)
1992).
The
species
in
(he
Cantons
Obwalden
(Göllner-Sceteiding & Rezbanyat-Reser,
and Ticino
Thesium and is
on
previously
widespread
m
Baden-Württemberg (RiEGER
(Heckmann, 1999). Before Reeger (1997) the determination of the females
was unsure
were
found
1992) and has been found in the Wollmatinger Ricd in the Landkreis
& STRAUSS,
Konstanz
lives
been
1881: This is the third reliable record of this
probably
15. Sciocoris
(Rampazzt
(1864-1866)
Baden-Württemberg.
and therefore the records of C. dubius Scop, in Frey-Gessner
impressus
as
this
is
microphthalmia FLOR,
&
Württemberg
C.
Dfthier,
it is common
1997)
the
more common
species
in
1860
and
(Fig. 3): This species is known
Tyrol (Heiss, 1977); however,
from Ticino
in
Baden-
(RfLGER, pers. comm)
macrocephalus Fieber, 1851 (Fig. 3): Reliable records exist only from
Graubünden (Voellmy & Sauter, 1983), Neuchâtel (Barbalat, 1991), Ticino
(Göllner-Scheiding & Rezbanyai-Reser. 1992) and Thurgau (Coll. Blociilinger,
Miillheim). There are no checked records from Baden-Württemberg (RiEGER, 1996) and
the Fürstentum Liechtenstein (Bernhardt, 1992), but the species is found frequently in
Tyrol (Heiss, 1977). Reports also exist from Vaud, Wallis, Aargau, Zürich and
16. Sciocoris
Graubünden (Frey-Gessner,
1864-1866). However,
if
remains uncertain whether these
reports differentiate between S. macrocephalus and S. microphlhahnus.
57
Fig
3
Fieb
(uppet pait)
flowei pait)
bv the smuated latcnil
The
body length
is
Canton Luzei
f
maciocephalus
Sciocoiis
rioi
Acalypla
n
-1
cannata
fiom
species,
the
mm
(P
\NZER,
1806)
see
Ptrtcart, 1988)
is
w ei e
found
(auows) and by
m
the
miciophthalmus
longei eye stalks
Giuho)
the Canton Luzern-
discussion of Canton Schaff hausen
and Basel
Wallis
fiom Sciocons
distinguished
In Suitzciland theie
(A COSTA, 1842).
Cantons
be
gins ol the head
R ETcekinann and M Di
(Photos
inteiestmg species
out
2 Hunaceius mcifoi
Deihier &
about 6
can
mai
attnbuted to
a
A
imfhei
only
aie
two
îecoid
mistake of the
lecoids of this
1987,
(Ptrtcart,
name
The
Kaiseistuhl
actually found m Baden-Wuittcmbcig (Kaiseistuhl in Baden) and not m the
Aaigau Several îecoids come fiom Baden-Wuittcmbeig along the Rhine Vallev
(Heckmann 1996) and fiom the Landkreis Konstanz This is the fust tecoid foi
was
species
Canton
Ccntial Switzeiland
s
Pinalitus
which lives
1866)
(Meyer-Dur, 1843)
atomanus
on
Picea
and
tcccntly
Badcn-Wuittembeig
fiom Ticino
is
Heckmann, 1996) and
4
to the
the Black Foicst
Suebian Alb (Rieger peis
Pinalitus visicola (Puton, 1888)
This species lives
species
Aatgau (FrfY-Gessner,
(RAMPA7ZI & DriHTFR,
îestiicted to
alpme-boieal
In Switzeiland this
known fiom Bern, Basel and
is
1864
The distnbution
1997)
(Rieger
)
&
in
1992,
Strauss,
comm
on
Viscum album and has been
lecoided
In the Landkreis
m Switzeiland onl> m the Canton Wallis (Ceruttt, 1937b)
Konstanz, the distnbution conesponds with that of Viscum Its distnbution foi Baden-
Win ttembei g has been desciibed
icflcct the
canopy,
is
m
Heckmann & RiEGFR
of the species
dependence
îaicly sampled
on
(in pi ep )
The spai
Viscum and the fact that its habitat
se
iecoids
mostly
in
the
Conci usions
The Red list of
Badcn-Wuittembeig is veiy helpful m înteipieting the îaie species
of Switzeiland Some \cio-theimophihc species iccoided moie oi less fiequently m the
south of Baden-Wuittembcig especially m the valley ot the Rhine and Hegau also occui
m
the Canton Schaffhausen
Switzeiland and
Mittelland
m
I he same species aie iccoided in the southern pait of
the Cantons Basel and Aaigau, but not in the Inneischweiz and
good candidates foi a Red list of Switzeiland
Hallodapus infest em Burm Bervtmus cuissipes
II S Berytinus monlnaçus Me\ -D Bentinus signoreti I ifb
Dtymus latus Doug &
Pentrechus
Pui
}eibasci
Embletlus
Sc,
F, Sciocons macrocephalus
giacihconus
FrhB and Sciocons miciophthalmus FlOR Fiom oui findings m luzein Hunacerus
include
,
majoi A
These species
Deuicocous
which
mono
ate
BOEl
,
,
C and Pinalitus \isicola PUT should be added
,
to
the list
58
Many species found in the ancient records of Meess (1900, 1907) and FreyGessner ( 1864-1866) appear to have decreased in abundance during the last century, eg
tipitlarius L., Aellopus atralus F.. Xanthochilus quadratus F., Sciocoris
umb ri nus WFF., Chlorochroa juniperina L. and Elasmucha ferrugata F. However, the
sparse data available on the distribution of heteropteran bugs do not allow a thorough
analysis of the situation in Switzerland, The data of Baden-Württemberg, however,
support the conclusion that these species decreased in the last century (LIeckmann,
Neides
1996). We
that this declines
assume
in both
agriculture production
were a
Germany
consequence of
intensification of
increasing
and Switzerland.
Acknoledgments
general assistance, H. BLÖCEfLlNGER, Cil. RiEGER and
Studer for permission to use unpublished data, to P.J. Edwards, Ch. Rieger and
Reser for their critical reading of the manuscript, to A. Müller for the permission
the material of the entomological collection of the Swiss Federal Institute
use
Technology (ETIIZ).
We thank B. Merz for
This work
was
funded
and
5001-44636)
Naturschutzamt).
and
by
the Swiss National Science Foundation
Canton
the
Schaffhausen
(Kantonales
S.
L.
to
of
(5001-044634/1
Plantings- und
Zusammenfassung
Im Schaffhausei
(Kanton Schaffhausen)
Randen
Wiesentypen
Kescherfang,
untersucht.
Wanzenarten
gefunden.
Dabei
und
Bodenfallen
wurden
ein
Auf dem
wurden zwischen
verschiedene
drei
Sauggerät.
Insgesamt
Gemeindegebict
wurden
Literatur und
neue
aufgrund
aus
1936,
(Kanton
festgestellt.
In
nämlich
angewendet,
untersucht
Luzern)
unseren
eigener Funde stellten wir fur beide Kantone Artenlisten
und
wurden
140
31
Untersuchungen
zusammen:
Aus dem
dem Kanton Luzern 20t Arten. Die
aus
Arten wird diskutiert.
Biogeographie ausgewählter
Drei Arten
Wiesen
Arten fur den Kanton Schaffliauscn und 26 für den Kanton Luzern. Anhand der
Kanton Schaffhaiisen sind bisher 207 Wanzenarten bekannt,
CHINA,
45
Ruswil/Rutlisholz
extensiv bewirtschaftete Wiesen untersucht und dabei 78 Arten
fanden wir 118
1999 drei verschiedene
und
1997
Sammelmethoden
Physatocheila harwoodi
tniciophtlialmus (WAGNER, 1951) und Strongylocoris steganoides (.1.
bisherigen Funde von Strongylocoris leucocephalus sollten überarbeitet werden.
dem Schaffhauser Randen sind Neunachweise für die Schweiz:
Polymeius
Sahlberg, 1875). Die
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Schötz, Kt.
62
Chapter 4. The influence of host plant diversity and food quality
of plant feeding heteropteran bugs (Heteroptera: Stenodemini)
on
larval survival
Manuela Di Giuho & Peter J. Edwards
Abstract
laboratory experiment we investigated the influence of host plant diversity and
food quality, m terms of nitrogen content, on larval survival of two plant feeding
bug species (Heteroptera, Minclac: Leptopterna dolobrata L., Notostira erratica L.).
Both species are strictly phytophagous and capable of feeding on a wide range of
different grass species, but are normally found on only a few ones. They typically
In
a
plants during ontogenesis and it has been suggested that this
represents a response to the changing protein content of the hosts. We compared the
development of animals reared on grass monocultures with that when they were
their host
change
four species. In addition the host grasses were grown under
two different nitrogen regimes to test whether nitrogen content is the key factor
reared
on
mixtures
host
determmtng
plant switching.
species had
plants, but only
Both
a
L.
significantly higher
dolobrata
Furthermore, there
content.
host
o\
plants
survival
showed
was no
a
when
rate
significant
feeding
response to
several host
on
food
and the survival rate of N. erratica larvae. Moreover, there
evidence that
our
mixtures have a
nitrogen
correlation between the nitrogen content of the
results cannot be
explained by
a
sampling effect,
better chance than the monocultures of
Our results suggest that the level of host
containing
in
strong
good
a
the
is
which the
food
factor
plant nitrogen
plant.
responsible for the diversity effect and that at least some Stenodemini need a variety
of host plants during larval development. In a field survey we found a positive
is
not
main
correlation between the number of Stenodemini species and the number of grasses,
indicating that diverse meadows can support more grass bug species. Our study
plant diversity can
phytophagous insects.
indicates that
survival of
food
Key-words:
enhance insect
host
quality, herbtvory. Heteroptera,
diversity by increasing
plant diversity,
insect
the larval
diversity
Introduction
partly because the chemical
composition of their tissues are very different from those of plants. In particular, the
protein content of insects is much higher than that of plants and the proportions of the
acids differ between animal and plant proteins (Southwood 1973;
ammo
various
Mattson 1980; White 1993). Various studies show that nitrogen content can be a crucial
factor for the development and repioduction ol heibivores (Strong, Lawton and
Herbivory presents
Southwood
major
difficulties
to
1984). In addition, the allocation of
changes considerably
insects,
nutrients to structures
like stems, fruits
during the course of plant
development.
plants is not only generally low but also
varies
during the season. Ohgophagous and polyphagous insects may respond to
changes m nutrient availability by switching from a plant with a low protein level to one
with a higher protein concentration (McNeill and Southwood 1978).
and flowers
both
seasonally
and
Thus, the protein content of
The Stenodemini
feeding
on a
strictly phytophagous and capable of
species, but are normally found on a restricted
wide range ol different grass
are
63
species. They change their host plants during ontogenesis and it has been
suggested that this represents a response to the changing protein content of their hosts
(Braune 1971; McNeill 1971,1973; Gibson 1976, 1980). However, there are other
possible advantages of changing host plant. Gibson (1976) has observed that
Leplopterna dolobrata L. and Stenodema laevigatum L. switch to unusual host plants in
set
a
of
year of extreme
by
the
drought.
and in
drought,
This
suggests
particular
that
to
host
grass
species that have
plant change
may
also
unfavourable climatic conditions. Indeed, the species richness of
may be
switch
factor
important
their host plants
an
affecting
the
performance
of
a
plant feeding
not
been affected
compensate
for
pfant community
insects which
can
in this way.
Although, the relationship between species richness of insects and plants has been
investigated in several studies, the results are contradictory and the underlying
mechanisms are not yet entirely understood (Strong, Lawton and Southwood 1984). In
recent years several diversity experiments have been designed to investigate how
species diversity influences ecosystem function, but the conclusion remain controversial
and insects have rarely been included in these experiments (Huston 1997; Hector and al.
1999). Our study investigates the effect of plant diversity on the development and larval
survival of phytophagous bug species and contributes to the debate about the role of
species diversity in ecosystems. In a laboratory experiment we examine how host plant
species diversity affects the growth and survival rate of Stenodemini. We compare the
larval survival of insects reared on grass monocultures with that of insects reared on
mixtures of four species. To test whether total nitrogen content is the key factor
responsible
for host
plant switching,
the
plants
have been cultivated under two different
nitrogen regimes. It is known that total nitrogen
with
nitrogen fertilisation, though
(Meister
species
and Lehmann
content of grasses tends to increase
the reaction of individual grass
1990). The experiment
was
L. dolobrata and Notostira erratica L. which
species
is variable
conducted with the Stenodemini
are common
in Swiss meadows.
The two species have different feeding habits; N. erratica feeds only on leaves, whereas
L. dolobrata is both leaf and seed feeding. They can easily be found in a sufficiently
laboratory experiments and they both have been reared successfully in
the laboratory (Braune 1971, 1983; Dolling 1973). If grass diversity increases the
survival rate of Stenodemini then wc might expect diverse meadows with several grass
species to support a higher number of Stenodemini than species-poor meadows. We test
this hypothesis with data from a field survey carried out in the north of Switzerland
(Canton Schaffhausen, Switzerland) in which the bug fauna and the plant species of 24
high
number for
meadows of two different management types
Material
and
were
recorded.
Methods
Field survey
representing two administratively
(regular use of slurry; 2-3
recognised grassland management types
cuts per year) and extensive (no fertiliser; one or two cuts per year). Twelve replicate
sites of each management type were selected. A description of the research area and a
detailed analysis of the effect of management on species diversity are presented in a
separate paper (Di Giulio, m preparation). The plant species composition was assessed
using six quadrats of one square meter per site. The extensive meadows were dominated
by Bromus erectus, while the medium intensive sites were dominated by
Arrhenatherum elatius and Trisetum flavescens (Studer 2000).
In
summer
1997
we
investigated
24 meadows
-
medium intensive
64
sampled using a standardised sweep-net method
(Remane 1958; Born hol dt 1991; Otto 1996). The sweep-net had a diameter of 40 cm
and samples were collected every two weeks by making one hundred sweeps with the
net over a distance of about 100 m. Sampling was only carried out when the weather
conditions were good, îe a minimum of J7°C and sunshine. Sampling was also
restricted to the period between 10.00 a.m. and 17.30 p.m.; the sampling order of the
fields was varied between weeks. Every site was sampled between 7 and 9 times from
May to September.
The
heteropteran bugs
were
Experimental design
experiment was to investigate the effect of plant species diversity
number of species per se
upon larval survival. The important variable was therefore the
rather than which species were present in the mixture; replicates were therefore
represented by several different combinations of the same number of species (Tilman et
al. 1996). Total nitrogen content was included in the experimental design as a second
factor m order to separate the nitrogen effect from the diversity effect (Huston 1997).
We chose a fully factorial design with the two factors: plant diversity and nitrogen
content. Both factors had two levels; thus insects were offered either one or four plant
species which had been raised on either high or low nitrogen. For each treatment there
were five replicates. The low nitrogen level was selected to correspond about to the
typical nutrient input in an extensive meadow (with no fertiliser input), and the high
level to one of a medium intensive meadow (40-60 mJ slurry per ha and year). The
species in both the monocultures and the mixtures were chosen randomly out of a
The aim of the
species pool comprising eight grass species which were cited in the literature as host
plants and were very common in the study area (Table 1). To compare the single
monocultures with the mixtures and to exclude the sampling effect (Wardle 1999) we
also examined the three monocultures which were not m the sampling design (P.
trivialis, F. ovina and F. pratensis). These treatments could only be carried out with N.
erratica as we had not enough larvae of L. dolobrata. The insects were reared on high
nitrogen
treatment
plants.
65
Table L The table shows the full list of the grass
(column: Mono)
chosen monocultures
host
plants
species
experiment and the randomly
quoted in the literature as
selected for the
and mixtures. All
species
are
of N. erratica and L. dolobrata.
References
Mixtures
Mono
Species
I
2
3
4
x
X
X
Arrhcnateritm clatius
X
X
Dactylis glotnerata
X
X
5
Gibson i976
Kullenberg 1944;
Osborn 1918;
x
Southwood &
Kullenberg 1944;
Leston 1959; Braune 1971; Gibson 1976
Festuca ovin a
Festuca
x
pratensis
X
X
Holen s lanaliis
X
Loliiini perenne
X
Phleuin prat en
X
sc
X
X
X
Poa trivialis
Kullenberg 1944; Osborn 1918;
Gibson 1976
1971; Gibson 1976
Braune
1944
x
Kullenberg
x
Kullenberg 1944; Southwood
x
X
Cultivation and chemical
Gibson 1976
Kullenberg 1944;
X
X
X
Kullenberg 1944;
Osborn 1918;
X
& Leston 1959
Kullenberg L944
'
analysis of'the food plants
The grass seeds were of local provenance and had been provided by the Swiss
Federal Research Station of Agroccology and Agriculture. The cultivation method is a
procedure used
inputs on the growth
standardised
nutrient
study
the research station to
at
and
development
(U. Walthcr, pers. comm.). The plants
the effects of different
leguminous species
of various grass and
grown out-of-doors under
were
a
mobile roof.
raining so that
be controlled, while temperature, humidity and light
rainfall and nutrient
regime corresponded closely to natural conditions. Each species was cultivated
separately in double-walled pots of 23.5 cm diameter which were filled with 9 kg of
loamy soil. The soil was fertilised with 6 g KMFPOa before sowing. The nitrogen
They
were
fertiliser
drained
dry
input could
prepared
was
added to
a
weather but remained covered when it
uncovered in
pot
at
through
one
any
a
solution
time. The
the pots and
refilled into the pots
level
as
so
that
was
no
fertilised
(2,5
pots
g
F1
were
of CaNcGô
regularly
x
was
4LI20),
and 100 ml
watered and the
surplus
water
collected in separate containers, from where it
nutrients
only
were
washed out. The grasses grown at
was
a
was
low
The pots of the
weeks after
germination.
monthly and thus received five times as much
the whole experimental period. The food grasses were cultivated from the
end of March to the end of August. Food plants for both bug species came from the
same batches of material over the whole experimental period.
nitrogen
high nitrogen
nitrogen over
were
treatment
were
once, ten
fertilised
plant
material
the
et al.
1983; Houba
these
two to three weeks from the
The total
nitrogen
content of the
Kjeldahl method (Novozarasky
analyses was collected every
Mid August.
Rearing of the
The insect
a
circular
was
determined
et
al.
by
a
modification of
1989). Plant material for
beginning
of
May
to
insects
rearing
opening
cages,
made from
in the front with
a
acrylic plexiglass, measured 25x25
gauze tube for
handling
cm
and had
the insects. Each cage
was
equipped with a small pot for the food plants and a water dispenser which was filled
daily. Freshly cut grass leaves and stems served as food and were provided every
second day. We fed only stems and leaves to both bug species, although under natural
conditions L. dolobrata feeds on leaves, flowers and seeds. To provide food in equal
66
quantities
all animals
fed
were
a
combined two stems of each grass
eight stems and
species. The specimens
total of
leaves. For the mixtures
were
reared under
a
we
constant
photoperiod of 16 h light/8 h dark at a constant temperature of 20°C. The humidity in
the climate chamber varied between 60 and 90 %, but it was lower in the cages. Low
humidity in climate chambers can be responsible for a high mortality at moult when
laboratory (McNeill 1973). We therefore sprayed
water into the cages daily m
humidity, though this did not
completely prevent larval death. Every day dead individuals were removed and the
number and larval stage of the larvae determined. The adult insects were killed the day
rearing nymphs
of L. dolobrata
m
the
order to increase the
after the final moult.
The
experimental
collected
(Canton
material for L. dolobrata
the field at the
m
beginning
comprised
total duration
Larvae of N.
recorded
daily.
(20-22 d) corresponded closely
of the larvae
erratica
was
were
populations in
put eight larvae.
of June from three
Graubiinden, Switzerland). In each cage
development
first instar larvae which
we
were
the Safienial
The state of
The duration from moult to moult and the
with the values
given by
reared from adult females which
were
McNeill
(1971).
collected at the
July in the Schaffhauser Randen m the North of Switzerland. These
females were kept in the climate chamber and pro\ided with freshly cut grass material.
In the first few days we observed several females laying eggs on the upper part of the
stems. The stems were replaced every second day and those with eggs were placed on
humid tissues and kept in transparent plastic boxes in the climate chamber. After about
12 to 14 days the first larvae hatched and they were immediately placed in the cages (20
per cage). The first adult males hatched after about 22 days and the first females after 25
days; after 35 days all insects had hatched.
beginning
Statislical
of
analysis
Least squares
regression
was
conducted to
analyse
the influence of food
species
the species number of Stenodemini. We used one-way ANOVA to test how
diversity on
nitrogen fertilisation affected the total nitrogen content of the grasses during the season.
Two-way ANOVA was conducted to analyse the influence of crude prolem content and
host plant diversity on larval survival of A. erratica and L. dolobrata. As a measure for
larval survival we calculated the proportion of the larvae which emerged as adults.
Larvae killed during handling (in total 12 out of 620 individuals) were subtracted from
the total number of larvae put info the cages at the beginning. We tested two models for
the N. erratica data. The first model comprised all monocultures tested, while the
second one included just the five monocultures actually used in the experimental design.
We also tested two models for the data of L dolobrata. The first model included only
fully hatched adults. In the second one we also included insects which had died during
the final moult, because the total number used was rather small and the mortality high at
the final moult.
Results
Influence of grass diversity
on
the
The number of grass species
species
number
the field
of Stenodemini
samples ranged from 6 to 13, while the
number of Stenodemini species sampled per site ranged from 2 to 4. Despite the small
numbers of bug species the species number of Stenodemini increased significantly (P <
0.05) with the number of grass species (Fig. 1, Table 2).
in
67
Table 2.
Regression model of the influence of grass species
on the species number of Stenodemini. The
number
values
were
In-transformed before the anahsis
REGRESSION MODE1
R2
=
p(0
SE
Variable
0.162
Constant
-0 088
0 535
-1.64
0 87
Grass species number
0 554
0.242
2.291
0 03
-a
o
c
4
£
Î5
3
<D
.Q
ZJ
^
en
CD
°
CD
1
'
Q.
00
6
8
7
Species
11
10
9
12
13
14
number of grasses
the
species number of Stenodemini correlates positively with
< 0 05). Each point is lor a different
of
number
(P
glasses
species
The
Fig.l
grassland
site and
is
based
on
6
Ixf
m? quadrats
for the grass species
and the recorded data for the buss.
Food
quality
experiment the use of nitrogen fertiliser increased the nitrogen content of the
24.89, P < 0.00001) and there were also
grasses significantly (ANOVA: Fi,7s
differences in the temporal variation in nitrogen content between the two nitrogen
treatments (Fig. 2). There was much more variation in nitrogen content between species
in the low nitrogen plants than m the high nitrogen treatment. However, in the high
nitrogen plants nitrogen content dropped sharply between June and July in all species.
The two bug species used in the experiment do not develop at the same time of year and
therefore they experienced differing nitrogen levels. L. dolobrata was reared in May and
June, while N. erratica developed m Jul)' and August. During May and June the
nitrogen content was on average higher and varied more than m July and August; food
quality for L. dolobrata was therefore better but varied more than for N. erratica (Fig.
In the
=
2).
68
50
jC
CD
"(1)
T3
CD
40
30
high nitrogen input
c
8,20
o
c
10
.05.
20.5.
14.6.
12.8.
27.7.
50
CD
*
40
"CD
X3
CO
30
ow
nitrogen input
c
©
O)
o
CD
20
10
4.05.
20.5.
14.6,
12.3.
27.7.
Date
•
Fig.
A. elatius
D.
H. lanatus
L. perenne
glomerata
F. ovina
F.
P. trivialis
Ph.
pratensis
pratense
development of the nitrogen content is shown ovei the expeinnental period. The grasses
high nitrogen input (uppei part) were teitthscd legularly, while the ones with low nitrogen
input (lower part) weie feitihsed once in Ma\ (see text loi specification). The grey boxes show
appioxtmately the period during which the two bug species developed. The dark grey box
indicates the development time of L. dololvata. the light grey box the one of N. erratica. The
piotcm level was high, but \anecl strongly in late spimg when L. dolobrata developed
(May/June), while it was low and \aned less in summer, when N. erratica developed
(July/August).
2. The
with
Influence of host plant
diversity and food
quality
on
larval survival
ofL. dolobrata
22.86. P < 0.001) and food qualify (Model
plant diversity (Pi l6
increased
larval survival of L. dolobrata (Fig. 3,
0.002) drastically
Moreover, there was a significant interaction between the two factors (Model
13.91, P < 0.002). Both the effect of nitrogen level and the interaction term
Host
13.91, P
=
<
I:
PL16
Table
=
3).
I: Fus
=
tended to
69
the insects which died at the final moult
excluding
0.066).
increase the larval survival when
(Model II: P1J6
=
0,059, P
=
Two-way ANOVA of the influence of nitrogen level and diversity
Table 3.
on
the survival of I.
were included which reached the adult stage, while in Model II
insects which died at the final moult were excluded (specifications see text). The proportion of
dolobrata. In Model I all adults
hatched adults
was
transformed for the
taken
as
a
measure
for larval survival. The values
F-ratio
p-valuc
0.059
3.91
0.066
0.117
7.8
0.013
3.91
0.066
d.f.
MS
F-ratio
p-value
d.f.
MS
N-Level
1
0.125
13.91
0,002
1
I
0.206
22.86
0.0002
1
13.91
0.002
t
0.059
Interaction term
I
0.125
Residual
16
0.009
16
0.015
Total
19
0.025
19
0.013
w
(square)-
Model 11
Source of variation
0.5
arcsine
analysis.
Modt;1I
Diversity
were
r
0.4
TJ
CO
~a
CD
0.3
o
05
0.2
O
.2
o
Q.
O
0.1
0.0
v.,
Q_
monoculture
-0.1
mixture
high
low
N-level
Fig. 3. Interaction plot of the two-way ANOVA of the effect of food quality and
plant diversity on larval sun ival of L. dolobrata. For this analysis the insects
which died at the final moult were included (Model I). The nitrogen level
represents the
two
nitrogen
values of the two treatments
high
regimes
are
and
low fertiliser
input.
Mean
shown.
either the low
high
ranged from 0 to 75% (Fig. 4). Larval
survival on mixtures of high nitrogen plants ranged from 12.5 to 75%, while that on
mixtures of low nitrogen plants ranged from 0 to 25%. There was a significant
interaction between nitrogen level and plant species number.
No larvae of L. dolobrata survived
nitrogen plants,
whereas survival
on
on
monocultures
mixtures
on
or
70
1.0
0.8
T3
CO
"O
CD
SI
O
0.6
03
Ci
2
"c
o
Q.
O
0.4
0.2
1
1.
Q.
0.0
4
Number of
Larval survival of insects reared
that of insects reared
on
cases
plant species
4. The larval survival of L. dolobrata increased with the
plants.
cases
10
4
1
Fig.
case
id CclSGS
on
species
monocultures
number of host
was
compared
to
mixtures of four species. The dots indicate the
of different species combinations at each level of
The size of the dots represents the number of cases.
replicates
species
richness.
mortality of insects reared on monocultures differed from that of insects
on
(Fig. 5). The majority of animals reared on monocultures died before
they had reached the fifth larval stage, while insects reared on mixtures showed not only
lower mortality overall but a more equally distributed mortality between the larval
Larval
reared
mixtures
stages.
3
2
Larval
4
on
Table 4 and
erratica very
Monocultures
during
larval
5
L. dolobrata
monocultures died earlier than those reared
Influence of host plant diversity
Mixtures
—
stage
Fig. 5. 'The frequency distribution shows the mortality of
Insects reared
\zzz3
and food
quality
on
on
mixtures
larval survival
development.
species.
of four
qf'N.
erratica
Fig. 6 indicate that host plant diversity increased the survival rate of N.
22.66, P < 0.001), while food quality had no effect (PU6
greatly (Fp16
=
71
0.65). There was no interaction between food quality and diversity (PU6
0.27, P
0.61). The same effects were found when including all monocultures into the
0.47;
0.55, P
model (diversity: FU6
16.28, P < 0.001; nitrogen level: FU6
0.63, P
interaction term: P].i6
0.44).
0.21,
P
=
=
=
=
=
=
=
Table 4.
level and
Two-way ANOVA of the influence of nitrogen
diversity on the survival of N. erratica. The design
factorial, with five monocultures and five
as
a
measure
for larval survival. The values
(square)-transformed
for the
was
mixtures
The proportion of hatched adults
nitrogen levels.
=
fully
for both
was
were
taken
arcsme
analysis.
p-value
F-ratio
Source of variation
d.f.
MS
N-EEVEL
1
0.007
0,21
Diversity
I
0.748
22.66
interaction term
1
0.009
0.27
Residual
16
0.033
Total
19
0.042
0.65
0 0002
0 61
I.U
0.9
tn
.+_*
0 8
U
"O
CO
0.7
T5
CD
SZ
o
OR
CO
sz
0.5
o
c
o
0.4
r
O.M
o
CI.
o
U.2
CL
monoculture
0.1
low
mixture
high
N-level
Fig.
6. Interaction
plot
eriatica.
monocultures and five mixtures for
The nitrogen level represents the two nitrogen
nitrogen levels,
high
and low fertiliser input. Mean values of the two treatments
100%
to
were
One
to
88%, while
(Fig. 7). On four monocultures survival
the monocultures of P. ovina
71%)).
For this
both
On monocultures survival ranged from 0
50
quality and
analysis the fully
of the two-way ANOVA of the effect of food
plant diversity on larval sur\i\al of As
factorial design was chosen with li\e
or
more
of these grass
lowest survival rates, which
survival rate
(P.
ovina:
survival rates
(50
and 67%).
ranged
88%)
was
(88%,), F.
species
are
on
regimes
shown.
mixtures it ranged from
higher than on mixtures; these
pratensis (82%) and L. perenne (67 and
were
was
also included in the mixtures with the
from 50 to 82%. The monoculture with the
highest
also included in the mixtures with the two lowest
72
1 case
4
1
Number of
9
2
•
5 cases
cases
plant species
Fig. 7 .The larval survival of N. erratica increased with the species number of the host
plants. Larval survival of insects reared on monocultures was compared to that
of insects reared
on
mixtures of four species. All monocultures tested in the
analysis, that are five monocultures of low
included in this
experiment
nitrogen plants and eight of high nitrogen plants. The dots indicate the
replicates of different species combinations ai each level of species richness.
are
The size of the dots represents the number of
Larval
higher
mortality
occurred
for insects reared
90
on
mainly
cases.
in the first and second larval
monocultures than for insects reared
on
stage and
mixtures
was
much
(Fig. 8).
r
80
70
>,
60
o
I
5°
°"
40
^
30
-
20
i~n
Mixtures
«a
Monocultures
10
0
_
2
3
4
5
Larval stage
Fig. 8. The frequency distribution shows the mortality of A', erratica during larval development. The
larval mortality of insects reared on monocultures was very high m the first and second larval
stage, but stayed low in the later stages. The mortality of larvae reared on mixtures was
generally
low. For this
and five mixtures.
analysis the fully factorial design
was
chosen with five monocultures
73
The
mean
nitrogen
nitrogen content of plant
experiment (27.7.) and after two weeks
calculated
content was
beginning of the
significant correlation
material collected at the
(12.8.).
There
and the
mean
was no
nitrogen
monocultures of the
content of the
high nitrogen
food
as
the average
between the larval survival of TV. erratica
plants (Fig. 9;
treatment were
t <;
=
included in this
-0.88, P
=
0 41). All
analysis.
175
§155-
of hatched adults
Proportion
Fig.
9
Lai val suivival of \
content
of the food
cuatiea was not con
plants
1.00
0.75
0.50
0.25
0.00
To calculate the
elated with the
mean
mean
nitiogen content,
nitrogen
we
used
That
and 12
was at
8)
plant matenal collected at two diffeient dates (27 7
the beginning of the evpeitmenf and aftei two weeks Foi this analysis we
included all eight monocultuics of the high nitiogen tiealmcnt
Discussion
Both Stenodemini species have
a
significantly higher
plants, but only m the case of L.
by nitrogen content (Figs 3, 4, 6,
survival rate when
several host
dolobrata
affected
7 and 9, Tables 3 and
is
feeding
on
there evidence that survival
is
4). These results
plant nitrogen is the main factor
responsible for the observed host plant changes as has been hypothesised (Gibson 1976.
1980; McNeill 1971, 1973), We suggest that other factors may also be responsible for
the positive effect of host plant diversity on the survival of the two bug species.
Previous studies indicate that insects respond to several components of the diet and not
suggest that neither
only
to total
nitrogen
variation
level
nor
content. Other
of nitrogen, the balance of
ammo
oi
host
important 1 actors
can
be the form and
availability
acids, the water content of the host and the content of
minerals and trace elements. Furthermore,
secondary compounds and mechanical or
important role (McNeill and Southwood
plant defences may also pla\ an
1978; Strong, Lawton and Southwood 1984). N. erratica larvae reared on monocultures
showed the highest mortality in the first larval stage, while it was much lower later m
chemical
development (Fig. 8). Mortality of insects reared 011 monocultures of A. elatius (100%),
IL lanatus (100%) and D glome rat a (77.5%) was extremely high, although these plants
are
usual hosts of older larvae and adults. The fust instar larvae died very
they
were
put
into the cages,
indicating
starved. We suggest that chemical
young larvae from
feeding
on
avoid the negative effects ol
or
these
the
that
soon
after
they could not feed on these grasses and
plant defences may have prevented the
mechanical
plants. How ever, larvae reared on mixtures could
plants by choosing between various species and
74
therefore had
that
observed
who
Stenodemini larvae.
The
majority
larval
are
important
A
of larvae reared
stage while,
distributed
instar
supported by Gibson (1976)
certain grass species are avoided particularly by very young
Larval mortality of L. dolobrata showed a different pattern (Fig. 5).
better chance of survival. Our
a
when
on
reared
findings
are
monocultures died between the second and fourth
on
mixtures,
mortality
larval
was
more
evenly
development time. This indicates that the second to the fourth larval
sensitive stages, during food quality and a variety of host plants play an
over
role.
problem
inherent in all
this experiment
a
positive
diversity experiments
effect of
mixtures
is the so-called
could result
contain
that mixtures of several
sampling effect;
because there is
simply
good quality plant,
species
higher probability
of a high nitrogen content, than monocultures (Huston 1997). However,
indicate that the diversity effect cannot entirely be explained in this way. No
dolobrata survived
on
a
monocultures, while most of the larvae reared
in
a
in terms
the results
larvae of L.
mixtures of
on
high nitrogen plants did reach the adult stage (Fig. 4). The effect is less clear for N.
erratica, though in most cases survival on mixtures was greater than on any of the
monocultures (Fig. 7). However, the values of the four mixtures lie below those of the
best monocultures, indicating that some monocultures are better than some mixtures. It
is interesting that the grass species of the best four monocultures are also included in the
mixtures with the lowest survival rates, indicating that larval survival on mixtures is not
determined by the best plant species in the mixture. The benefit of the mixtures
therefore results from the various plant species in the mixture and not exclusively from
the best
The
species.
content of the grasses
nitrogen
until autumn
(Fig. 2),
trends winch
1980). Bug species which develop
in
those
emerging
species used m the experiment.
summer or
drops
occur
m
autumn
towards mid-summer and remains low
in natural
spring
therefore
populations of
have generally
(Gibson 1976), which is also the
grasses
(Gibson
better food than
case
for the two
plants of N. erratica had much lower nitrogen
(Fig. 2). In fact, N. erratica can produce two
generations per year. The first generation develops during spring while the second one
emerges in late summer, indicating that this species can cope with a wide range of
nitrogen levels (Gibson 1976). Further, Gibson (1980) showed that TV. erratica does not
fully exploit the nitrogen available in a sward and apparently does not select within a
sward for plants with an above average nitrogen content. Our results support these
findings. Larval survival of TV. erratica seems not to be affected by the nitrogen
treatment and the nitrogen content of the host plants (Table 4, Figs 6 and 9). L.
dolobrata. however, does seem to need both host plant diversity and good food quality
(Table 3, Figs 3 and 4), Our observation that this species only occurs in diverse but not
excessively nutrient-poor meadows supports these findings. Several studies show that L.
dolobrata has a high nitrogen demand during its development. McNeill (1971, 1973)
demonstrates that it changes from leaf to seed feeding at a time when protein content of
the leaves drops drastically and plant nutrients are increasingly allocated to the
development of stems and seeds. Furthermore, it fully exploits the available nitrogen in
a sward by feeding on hosts with an above-average nitrogen content (Gibson 1980).
Some noticeable features of its hfe-history may be responsible for this high nitrogen
demand. Under natural conditions this species develops and reproduces within a period
contents than those of P.
of
about
two
and
a
The food
dolobrata
half
months
(Braune
1971;
McNeill
197L,
1973).
Two
morphological forms of females occur in this species: the brachypterous or short winged
morphs produce many more eggs than the long winged macropterous morphs which
75
dispersal function. Because of their high egg production the brachypterous
female morphs need a higher amount of nitrogen and, indeed, nitrogen seems to be the
crucial factor for the trade-off between dispersal and reproduction. A shortage of
nutrient rich seeds and flowers induces the development of macropterous females.
These disperse but postpone the development of eggs and produce on average fewer
have
a
1918; Braune 1983).
offspring (Osborn
To
summarise, the experiments show that
at least some Stenodemini
species
arc
plants for their larval development and survival. The
plants appears not to be the crucial factor for host plant
nitrogen
switching during ontogenesis. The survey study supports these results (Fig. I, Table 2),
indicating that there is a positive correlation between the numbers of grass and
Stenodemini species. Species rich meadows therefore may support more Stenodemini
species by offering better conditions for larval development.
dependent
variety
a
on
of host
content of the host
Acknowledgements
Lehmann, Andreas Rtiegger and Hansueli Briner for the grass
seeds, Josef Lehmann, Ueli Walther, Ernst Brack and Robert Richli for their assistance
We thank Josef
tips on cultivating the food grasses and Stefan Bosshard for his help at rearing the
bugs in the climate chamber. We are grateful to Sibylle Studer for permission to use
unpublished data, to Franz Schubiger and the technical staff of the Swiss Federal
Research Station for Agroeeology and Agriculture for the chemical analysis of the plant
and
material,
to Josef Lehmann
and Walter Dietl for their comments
on
the reaction of
for
grasses to fertilisation and to Bernhard Mer/, Sibylle Studer and Erhard Meister
their critical reading of the manuscript. Finally, we thank Bernhard Schmid for his
assistance with the
This work
and the Canton
experimental design.
by the Swiss National Science Foundation (5001-044634/L)
Schaffhausen (Kantonales Plantings- und Naturschutzamt).
was
funded
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77
Conclusions
Grassland management affects both species richness and species
insect
communities.
Extensive
meadows
arc
more
diverse
than
composition of
medium intensive
specialised insect species while the
medium intensive sites are characterised by common and widespread bug species. Our
study supports the findings of previous studies and further indicates that even a slight
intensification of management reduces bug species diversity in grasslands (Chapter I).
The extensive meadows of the Schaffhauser Randen are particularly valuable habitats
for some rare xcro-thermophilic bug species The climatic and soil conditions m this
region apparently favour these species and have also limited the intensification of
agricultural production. This is also the reason why many species of other taxa (eg
butterflies and flowering plants) persist m this area but have disappeared from other
regions of Switzerland. As few data are available on the distribution of individual bug
species m Switzerland, we cannot say how the heteropteran fauna changed m the last
decades, but we think that the situation is similar to that for other taxa (Chapter I & 3).
The bug fauna vanes both within and between grassland management types. The
extensive sites arc specially variable m species nchness and composition. These
lmdmgs suggest that apart fiom management, lactois like structure of the surrounding
landscape, aspect and location strongly influence the communities. The medium
intensive meadows, however, show only little variation, indicating that management is
the most impoitant factor determining the heteropteran fauna of these sites (Chapter I).
meadows; they
We found
(Chapter
contain
a
number of
rare
and
correlation between the species numbers of bugs and plants at a site
1). However, the number of ohgophagous bug species (eg Stenodemini)
no
significantly with the species number of their host plants, suggesting one
reason why flonstically diverse meadows can support more herbivorous insect
species.
Previous studies have shown that Stenodemini change their host plants during
ontogenesis and it has been suggested that this repiesents a response to the changing
piotcm content of the hosts. In a laboratoiv expenment we examined how host plant
diversity and food quality affect larval survival. The results suggest that host plant
diversity significantly increases the larval survival but that the crude protein content is
not the mam factor responsible for this effect. At least some herbivorous bugs seem to
depend on a variety of host plants for their development, and there is evidence that plant
diversity can increase insect diversity by enhancing the larval performance (Chapter 4).
increases
In
the
flowering plants and the butterfly fauna, the heteropteran
greatly between the four areas investigated (Chapter 1 & 2). Many
bug species occui only m one or two areas and seem to have restricted ranges,
suggesting that their distribution is limited bv then dispersal abilities. Fragmentation of
the habitat therefore represents a major problem for their populations. In general plant
and butterfly species associated with grassland seem to have colonised the areas more
equally, perhaps because they aie less limited by dispersal. We had expected insect
diversity to be greater in the area containing only grasslands (Zelgli/Mösli) than in areas
with a mixture of grassland and arable land habitats. Interestingly this was not the case,
contrast
communities
and there
is
perhaps
to
differ
even
an
indication that species richness
because these
is
greater
in
areas
of mixed land
heterogenous and present a wider range of
occur only m areas of mixed land use and
bug
tor some common species we could show that then abundance increases
significantly
with the proportion of arable land in the surrounding landscape (Chapter 2). Another
reason for the difference may be that the grassland area is dominated
by extensively
use,
niches
We found several
areas
are more
species which
78
managed
not to
meadows.
be cut before the first of
both the
escape
July.
regulations
alternative
the
and
habitats.
recommended to enhance insect
for subsidies, extensive meadows
Most farmers therefore cut the meadows
weather conditions and regulations
simultaneously. Moreover,
insect, particularly species
to
to the
According
allow,
and most sites
as soon as
cut
almost
cutting very fast and many
stages with very limited mobility have no chance to
A rotational management of grasslands is therefore
diversity at a landscape level.
machinery
used makes the
are
are
79
Curriculum vitae
Name
Di Giuho Müller
Vorname
Manuela
Geburtsdatum
17. Juli 1970
S t aats an geh öri gkei t
Italien, Schweiz
Bürgerort
Bettlach
Matura
Wirtschaftsgymnasium in Solothurn
Mafuritätsabschluss; September 1989
Studium
Studium der
sität
(SO)
Zoologie
und der Umweltwissenschaften
logischen
am
Institut für Umweltwissenschaften und
Dissertation
am
Zoo¬
Agrarö ko logic und Landbau
Forschungsanstalt
für
Insect
diversity
landscape
in
des Teufelsab-
Geobotani sehen Institut der ETHZ und der
structure
Leitung: Prof. P.J. Edwards
Aktuelle Anstellunt
am
Museum der Universität Zürich
Diplomarbeit: Die phytophage Insektengemeinschaft
biss (Succisa pratensis)
and
der Univer¬
Zürich, 1990-1996
Abschluss 1996
Dissertation
an
Wissenschaftliche Mitarbeiterin ETHZ
Eidg.
(FAL), 1997-2000
The effects of management

Fulltext - ETH E-Collection

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